CONTENT - International Society of Zoological Sciences

XXth International Congress of Zoology, Paris, 26-29th August, 2008 

Foreword 

The XXth International Congress of Zoology was promoted by the International 

Society of Zoology (ISZ) and the French Zoological Society (SZF). It was jointly 

organized by University Pierre and Marie Curie (Paris 6), the National Museum of 

Natural History of Paris (MNHN) and the Faculty of Sciences of Orsay (University 

Paris-South). It was strongly supported by the Ile-de France District, the Linnean 

Society of London, the International Union of Biological Science (IUBS), research 

organisations (CNRS, INRA), the French Foreign office, the University Paris 7, the 

Jacques Monod Institute and various other sponsors. It was held in Paris from the 

26th to 29th August, with 460 participants from 40 different countries. 

The congress was for mainly held at the University Paris 6, in the Great 

Lecture Hall of the MNHN and, for the Symposium “Evolution and Development”, in 

the Faculty of Sciences of Orsay. 

The programme comprised 4 plenary lectures (including the opening lecture 

given by Prof. Jules Hoffmann, President of the French Academy of Sciences) and 

26 symposia, thus giving most participants the possibility to present their research, 

with a total of 223 oral presentations (including 97 invited lectures) and 138 posters. 

The different symposia covered most research areas of animal biology: systematics, 

biodiversity, evolution, ecology, reproduction, development, comparative immunology, 

ecotoxicology, marine biology, behaviour, acoustic communication, animal societies. 

One symposium was specially devoted to the teaching of Zoology, including several 

lectures followed by a round-table discussion. 

Owing to the generosity of our sponsors, it was possible to support a large 

number of participants (invited speakers and symposia organizers). Particular 

support was provided to young participants through poster prizes based on the 

quality of contributions (and age of contributors). 

The present volume contains the abstracts of all oral and poster presentations. 

It represents an updated version of that provided to Congress participants. 

Jean-Marc JALLON, René LAFONT and Jean-Loup d’HONDT

CONTENT 

Plenary Lectures 1 

S1 -Contemporary approaches in Systematic Zoology 3 

S2-Mechanisms of speciation 11 

S3-A/B-Animal genitalia in evolution-in honour of William G.Eberhard 14 

SYSTEMA NATURAE 250. The International Commission on Zoological Nomenclature 

S4-PLENARY SESSION. The Linnean Ark-250 years of animal names 18 

S5 A/B-Current issues in animal nomenclature 20 

S6-Paleozoology and comparative anatomy:exceptional conservations 24 

S7-Paleontology and Evolution 26 

S8-Biodiversity and ecology of protists and S9-Genomics and cell biology of protists 28 

S10-Sex differentiation in vertebrates 31 

S11-Evolution and development 38 

S12-Transitions from clonal to sexual reproduction: key variations of a key process 44 

S13-Comparative Immunology 45 

S14-Venomous animals and their venoms 47 

S15-Integrative biology of acoustic communication 49 

S16-Genes, individuals, societies: current trends in social insects 52 

S17-Phenotypical plasticity and behavior 57 

S18-Interdisciplinary approaches in ecology: from individuals to populations and communities 64 

S19-Animals ecotoxicology 73 

S20-Marine organisms and symbiotic systems in extreme environments 

S21-The ecological and evolutionary consequences of global climate evolution on population, 

77 

species and ecosystem 87 

S22-Studies of invasive animal species 90 

S23-Comparative organogenesis in Animals 94 

S24-Lamarck’s Philosophie Zoologique: 200 years 97 

S25-A tribute to Darwin 99 

S26-Diversity in teaching Zoology 101 

Author index 103

ICZ2008 - Abstracts 

The rise and fall of hox genes clusters 

Denis Duboule 

National Research Centre ‘Frontiers in Genetics’. Department of 

Zoology and Animal Biology, University of Geneva, Sciences III and 

School of Life Sciences, Ecole Polytechnique Fédérale, Lausanne, 

Switzerland 

During development, the body plan of all bilateria is organized by a 

set of homeobox containing genes members of the Hox gene family. 

While all bilaterian animals have a related set of Hox genes, the 

genomic organisation of this gene complement comes in different 

flavours. In some unrelated species, Hox genes are clustered in the 

genome, yet, in others, they are not. This indicates that the bilaterian 

ancestor had a clustered Hox gene family and that, subsequently, 

this genomic organisation was either maintained or lost. Remarkably 

enough, the tightest genomic organisation, in terms of overall size 

and resistance to foreign repeats, is found in vertebrates, raising the 

embarrassingly finalistic possibility that vertebrates have maintained 

best this ancestral configuration. 

Alternatively, could such an improved genomic organisation have coevolved 

along with the emergence of vertebrates, possibly linked to 

the whole genome amplification events which occurred at this time, a 

possibility that is at odds with our current perception of evolutionary 

mechanisms? When discussing the why’s and how’s of Hox gene 

clustering, we need to account for three points: the mechanisms of 

Hox gene cluster evolution; the underlying biological constraints and, 

finally, the developmental modes of the animals under consideration. 

By integrating these parameters, some general conclusions emerge 

that can both help solve the aforementioned dilemma and impact 

upon our understanding of the evolution of genome structurefunction 

relationships in vertebrates, in particular concerning largescale 

gene regulation. 

The Antimicrobial Defense of Drosophila : a paradigm for Innate 

Immunity 

Jules Hoffmann 

IBMC, CNRS UPR 9022, 15 rue René Descartes, 67084 Strasbourg, 

France 

Insects have long been known to be particularly resistant to microbial 

infections. A long list of scholars have established over many 

decades that this powerful host defense is a multifaceted process 

with both cellular and humoral arms. In the early 90ies, our group 

has chosen the fly Drosophila melanogaster to address the 

molecular mechanisms of the inducible antimicrobial defense. As is 

the case for other insects, Drosophila responds to an immune 

challenge by synthesizing potent antimicrobial polypeptides. We 

have asked three sets of questions : (1) what is the identity of these 

inducible molecules in Drosophila ? (2) how is the expression of the 

genes encoding these peptides controlled during the immune 

response to infection ? (3) how does the fly recognize the state of 

infection, and does it discriminate between various infecting agents? 

The presentation will review our current understanding regarding 

these three sets of questions. It will highlight the fact that, namely 

through its highly tractable genetics, Drosophila is now the best 

known biological model for innate immunity in metazoans. Parallels 

will be drawn with recognition receptors, intracellular signalling 

cascades and effector molecules in other groups, as well in 

evolutionary ancient phyla as in more recently evolved groups, 

namely in mammals. The presentation will propose the conclusion 

that the molecular mechanisms of innate immune defenses evolved 

very early in evolution (Sponges, Cnidaria) and that their general 

characteristics have persisted without major changes in all 

metazoans, including mammals. 

Ferrandon D, Imler JL, Hetru C, Hoffmann JA. (2007) The Drosophila 

systemic immune response: sensing and signalling during bacterial 

and fungal infections. Nat Rev Immunol. 7:862-74. 

Lemaitre B, Hoffmann J. (2007) The host defense of Drosophila 

melanogaster. Annu Rev Immunol 25:697-743. 

Plenary lectures 

- 1 - 

Zoological systematics/phylogeny at the beginning of the 21st 

century 

Claus Nielsen 

Zoological Museum, The Natural History Museum of Denmark, 

University of Copenhagen, Universitetsparken 15, DK-2100 

Copenhagen, Denmark 

Darwin founded modern biology by proposing that the biological 

diversity is the result of evolution through ”descent with modification”, 

but he did not use his revolutionary ideas in his systematic studies. 

Haeckel introduced the terms ontogeny and phylogeny and gave a 

completely modern definition of a monophyletic group. Hennig 

sharpened phylogenetic reasoning by distinguishing between 

plesiomorphies and apomorphies and introduced the useful sistergroup 

concept. Thus, the theoretical background for a classification 

which reflects evolution, viz. a phylogenetic or cladistic classification, 

has been available for half a century. However, there is no generally 

agreed phylogeny for the main groups of animals, which is what I will 

concentrate on in this lecture, and this seems to be the result of 

various problems with the different methods of analysis. 

The “classic”, speculative method of arranging the animal groups 

according to certain ideas about the evolution and based on 

available information of morphology and embryology has been used 

for more than a century. Hennig analyzed his trees in an attempt to 

distinguish plesiomorphies, apomorphies and convergence, but the 

number of characters and taxa soon made his “manual” method 

inadequate. The introduction of computer aided cladistic programs 

has greatly enhanced the possibilities for analyzing large numbers of 

taxa with many characters. Much effort has been put on the 

elaboration of the statistical programs, whereas there has been 

comparatively little emphasis on the data matrices. This is 

unfortunate. Every entry in a data matrix reflects the author’s idea 

about homology, because only homologous characters should be 

considered. The early optimism saying that one should just enter 

everything and then let the programs sort things out has turned out 

not to be realistic. Characters like coeloms, segmentation and 

blastopore fate have innate problems of definition and homology. 

Inherent is of course the problem of weight of characters and 

polarization of changes. Most matrices contain errors, and several 

analyses have shown that changes in few characters may change 

the topology drastically. The cladistic analyses of morphological 

characters must be characterized as objective analyses of subjective 

character matrices. 

The analytical methods based on sequence data have given us 

excellent information about phylogeny of many groups and forced 

new interpretations of several of the accepted morphological 

characters, but the interrelationships especially of the so-called 

lophophorate phyla and of the basal metazoan radiation still appear 

unresolved. My limited knowledge of the methods and problems of 

molecular phylogeny does not suffice for a thorough discussion, but I 

will present a number of examples to demonstrate the problem. 

This could sound pessimistic, but at the end I will try to demonstrate 

that a combination of all the different types of information with due 

reference to continuous evolution of fundamental biological functions, 

such as feeding and locomotion, can indeed produce a probable 

phylogeny of the basal metazoans, which is then open to subsequent 

testing by new studies.

Aristotle founder of Zoology: when art imitates nature 

Rosa-Maria Polymeni 

National and Kapodistrian University of Athens, Faculty of Biology, 

Department of Botany, Panepistimioupolis, Athens 157 84, Greece 

Aristotle, one of the biggest philosophers and scientists of the Greek 

Classical era is the founder of Zoology. Furthermore his versatile 

genius together with his voluminous works on natural history allows 

considering him as co-founder, with Theophrastus, of biology. 

Aristotle had a penetrating and in the same time broad mind 

combined with an empirical manner of study. It is likely that about 

400 works are attributed to him. Greeks in classical years, never 

dispute the primacy of nature compared to the human creations: […] 

if natural products could also be produced by art, they would move 

along the same line that the natural process actually takes […]. 

Indeed, as a general proposition, the arts either on the basis of 

nature, carry things further than nature can, or they imitate nature 

(Aristotle, On Nature 199a). Descriptions of numerous creatures 

given by Aristotle in detail include information on the colours. In the 

work “On colors” that is thought to be written by his pupil and 

colleague Theophrastus, Aristotle rejects the active projection of 

visual rays by the eye. The real object of vision is color, which is a 

property of the surface of things. Such old but advanced ideas could 

be proved and be usable only in our days using new technological 

tools. In the frame of this approach, we studied the microsculpture of 

the skin surface of salamanders of Greece; it seems likely that colour 

reflects the state of surface and it depends on a relationship between 

light and the corporeal quality of the matter. 

- 2 - 

ICZ2008 - Abstracts 

Aristotle and Theophrastus founders of the Lyceum: when art 

imitates nature 

Sophia Rhizopoulou 

National and Kapodistrian University of Athens, Faculty of Biology, 

Department of Botany, Panepistimioupolis, Athens 157 84, Greece 

Aristotle's successor as head of the Lyceum at Athens was 

Theophrastus (371–286 BC), a Greek Peripatetic philosopher and 

pupil of Aristotle; he studied at Athens under Aristotle, and when 

Aristotle was forced to retire in 323 BC he became the head, in the 

academy in Athens founded by Aristotle. Under Theophrastus the 

enrolment of pupils and auditors rose to its highest point; also, he 

was responsible for the first botanical garden. Evidently, among 

other issues, colour has attracted their interest and attention. Ideas 

of early thinkers were not -and could not have been- grasped on a 

scientific level without knowledge of a kind that lay far in the future. 

Colour on the surfaces of living tissues could hardly have been 

visualized without a corresponding reference to the microscale 

parallel. Millions of years before man made manipulated synthetic 

structures, biological systems were using forms at the nanoscale 

level to produce striking optical effects; according to Theophrastus 

art imitates nature (the Causes of Plants II.18.2). We studied the 

microsculpture of floral surfaces and pollen grains –from plants 

grown in the Mediterranean Basin and referred by Theophrastus in 

his classical texts (Enquiry of the plants and the Causes of plants)– 

by using advanced microscopic techniques; to the best of our 

knowledge imaging of petals has not been reported hitherto. Our 

findings on microsculpture may be linked with aspects on colour 

revealed from ancient literature; likewise, visual perception of living 

tissues reminds us of classical fragments.

ICZ2008 – Abstracts S1 

S1 - Contemporary approaches in Systematic Zoology 

Molecular phylogeny of the genus Gerbillus (Rodentia 

Gerbillinae) in Tunisia 

Awatef Abiadh 1 , Paolo Colangelo 2 , Tahar Lamine Cheniti 1 and 

M’barek Chetoui 1 

1 Laboratoire d’Ecologie Animale. Faculté des Sciences de Tunis. 

Université de Tunis El Manar II, 1092 Tunis, Tunisia 

2 Departamento di Biologia Animale dell’Uomo, Universita di Roma 

« La Sapienza ».Via Barelli 50, 00161 Roma, Italy 

Although it forms an important component of the mammalian fauna of 

Northern Africa, the taxonomic and phylogenetic relationship within 

the genus gerbillus are still ambigous. The present study introduce 

finding based on the hole sequence of the cytochrome b (1140bp) 

mitochondrial gene of six species Gerbillus campestris, G gerbillus, G 

tarabuli, G latastei, G nanus and G simoni. Our result shows that 

species are monophyletic and they are divided into two groups. There 

is an ancestral group with stable karyotype and the other is 

characterized by a considerable number of chromosomal 

rearrangements. 

DNA barcoding in Birds: comparing the performance of three 

mitochondrial genes (cox1, cob, 16S) in species recognition 

Mansour Aliabadian 1,2 , Mohammad Kaboli 3 , Vincent Nijman 1 and 

Miguel Vences 4 

1 

Institute for Biodiversity and Ecosystem Dynamics and Zoological 

Museum, University of Amsterdam, Mauritskade 61, 1092 AD 

Amsterdam, The Netherlands 

2 

Department of Biology, Faculty of Sciences, Ferdowsi University of 

Mashhad, Mashhad, Iran 

3 

Department of Fishery and Environment, Faculty of Natural 

Resources, Tehran University, Iran 

4 

Division of Evolutionary Biology, Zoological Institute, Technical 

University of Braunschweig, Spielmannstr. 8, 38106 Braunschweig, 

Germany 

For comparative zoology, proper recognition of basal taxa is 

indispensable, and in this the choice of a suitable gene with high 

phylogenetic resolution is crucial. For the goals of species 

identification in animals, the cox1 has been introduced as standard 

marker. Making use of the difference in intra- and interspecific genetic 

variation–the DNA barcoding gap- has been successful in species 

identification across a wide array of taxa but in some cases failed to 

delimit the species boundaries of closely allied allopatric taxa or of 

hybridising sister taxa. Here, we extend the sample size of prior 

studies in birds for cox1 (756 species) and target especially species 

that are known to be closely related and/or are known to hybridise. 

We compare the effectiveness of cox1 barcodes for the identification 

of bird species on a global scale. To obtain a larger data set, we 

include also DNA sequences of two other mitochondrial genes 

cytochrome b (2087 species) and 16S (498 species), which allows 

inclusion of many additional taxa. The three genes taken together, the 

analysed sequences comprise 2719 species of birds. Our results 

showed that a wide gap exists between intra- and interspecies 

divergences for both cox1 and cob whereas no such gap is seen for 

16S. Most of the closely related species that are known to hybridise, 

however, showed average divergences that were intermediate 

between intraspecific and interspecific distances for both cox1 and 

cob. This study confirms, DNA barcoding in closely related and 

potentially hybridising species should not rely on mtDNA alone. 

- 3 - 

Molecular phylogeny of sea spiders (Arthropoda, Pycnogonida) 

Juliette Arabi and Alexandre Hassanin, 

MNHN, Département Systématique et Evolution, UMR 5202 – 57, rue 

Cuvier, CP 51, 75231 Paris Cedex 05, France. 

Sea spiders or pycnogonids constitute a class of marine arthropods, 

which are found from shallow waters to abyssal depths in all oceans 

and seas. They present a large range of sizes and forms, but 

members of this group are clearly united by several unique 

morphological characters, such as the prominent external proboscis, 

the ovigers and multiple gonopores. They contain 1334 described 

species currently ranged in ten or eleven families. The phylogeny of 

sea spiders remains controversial, and recent molecular studies 

revealed conflicting results for inter-familial relationships, the position 

of the genus Rhynchothorax, the monophyly of Ascorhynchidae and 

its basal divergence with respect to all other families. Several nonexcluding 

hypotheses can be proposed to explain these conflicts: 

misidentification of the taxa, taxa sampling, missing data, divergent 

signal between molecular markers, tree misrooting, over-interpretation 

of the results, or the use of inappropriate methods or models for tree 

reconstruction. To test these hypotheses, the phylogeny of 

Pycnogonida was reassessed in a first approach by the analysis of all 

18S rRNA sequences available in the nucleotide databases, and then 

by the analysis of six genes for 34 taxa. The six markers were 

analysed separately and also combined to benefit the signals of each 

of them. 

DNA taxonomy and barcoding of monogenean parasites - 

lessons from Gyrodactylus 

Lutz Bachmann, Haakon Hansen and Tor A. Bakke 

PO Box 1172 Blindern, 0312, Oslo, Norway 

DNA taxonomy and barcoding use nucleotide sequence data to 

achieve comprehensive species descriptions that facilitate reliable 

species diagnostics and rapid assessment of biodiversity, both of 

which are of great importance for parasitologists. Such molecular 

approaches have been applied to the monogenean genus 

Gyrodactylus, in particular to G. salaris, the cause of serious 

gyrodactylosis on Atlantic salmon. Here, we discuss, using the 

example of G. salaris and related species, why DNA barcodes, 

although powerful for biodiversity assessment, are insufficient to 

appropriately characterize parasite species – from a parasitological 

point of view – in the absence of additional data on and infection 

biology and morphology. 

Cave-dwelling springtails (Collembola) from Georgia 

Shalva Barjadze and R. Djanashvili 

Laboratory of Entomology, Georgian Institute of Zoology, 

Chavchavadze av. 31, 0179, Tbilisi, Georgia 

The investigation of the Georgian cave dwelling springtails began at 

the beginning of 1960s, but unfortunately collapsed in 1980s. 28 

species of the springtails are recorded from the caves of Western 

Georgia, united in the 3 orders, 8 families and 21 genera. A restudy of 

R. Djanashvili’s collection permits us to add 13 species, namely 

Arrhopalites caecus (Tullberg, 1871), A. pygmaeus (Wankel, 1860), 

Ceratophysella cavicola (Börner, 1901), Desoria trispinata 

(MacGillivray, 1896), Deuteraphorura variabilis (Stach, 1954), 

Folsomia spinosa Kseneman, 1936, Neelus murinus Folsom, 1896, 

Oligaphorura schoetti (Lie-Pettersen, 1896), Onychiuroides 

granulosus (Stach, 1930), Proisotoma minuta (Tullberg, 1871), 

Pseudachorutes dubius Krausbauer, 1898, Willowsia buski (Lubbock, 

1871), W. nigromaculata (Lubbock, 1873) to the springtail list of 

Georgia.

S1 ICZ2008 - Abstracts 

Contribution to the aphid fauna (Hemiptera: Aphidoidea) of 

Imereti region (Western Georgia) 

Shalva Barjadze 1 , N. Bakhtadze 2 , G. Bakhtadze 2 , N. Kintsurashvili 2 , N. 

Chakvetadze 1 and N. Zhukovskaya 2 

1, Laboratory of Entomology, Institute of Zoology of Georgia, 

Chavchavadze av. 31, 0179, Tbilisi, Georgia 

2 Institute of Zoology of Georgia, Laboratory of Karyosystematics and 

Biochemistry, Chavchavadze av. 31, 0179, Tbilisi, Georgia 

Aphids were investigated in Imereti region in 2004-2007. Of the 

aphids collected 17 species were first record for the abovementioned 

territory. Also, apterous and alate viviparous female, oviparous female 

and alate male of Macrosiphum symphyti sp. nov. living on 

Symphytum asperum (Boraginaceae) were described and illustrated. 

Aphis salsolae was recorded for the first time from Transcaucasia. 

After this investigation, aphid species number increased up to 48 

species in this region, which are united in the 5 subfamilies and 27 

genera. It was studied peculiarities of the aphids’ distribution in the 

landscapes of investigated territory. 

This abstract is a result of the designated project, which has been 

fulfilled by financial support of Georgian National Science Foundation 

(Grant № GNSF/ST 06/6-086). 

Effect of parasitism by Varroa destructor on morphometric 

characters of northern Algeria honeybees (Apis mellifera 

intermissa) 

Messaouda Belaid 1 , Salaheddine Doumandji 2 , Djamila Benaziza 3 and 

Assia Habbi 4 

1 

Department of Biology. Faculty of Sciences. University M’Hamed 

Bougara, Boumerdes, Algeria 

2 

Department of Zoology. National Institute of Agricultural El Harrach 

Alger, Algeria 

3 

Ecole Normale supérieure Kouba, Algeria 

4 

Apiary Oued Aissi Tiziouzou, Algeria 

Varroa destructor represents one of the major enemies of honeybees. 

It was found in Algeria in 1981 in apiary Oum Theboul el kala (East 

Algeria). Recently, several studies have been performed on honeybee 

morphometry in various localities from East Algeria. The results 

shows the heterogeneity in the population of Apis mellifera intermissa. 

The parasite had no significant effects on both the length of antenna 

and width of the posterior wing but affected the fresh body weight, 

length of tongue, length and width of the anterior wing, and length of 

tibia. In order to investigate the effects of Varroa on honeybees in 

Northern Algeria, 21 morphometric characters were used; 10 to 30 

worker bees per colony and two or five colonies were sampled at 

each location. 

Inventory and identification of some thrip species in coastal and 

sub-coastal regions of Algeria 

Hassina Benmessaoud-Boukhalfa, F Mouhouche and F Belmazouzi 

Département de zoologie agricole et forestière, Institut national 

agronomique, 16200 Alger, Algeria 

During the last decade, the presence of thrips on crops has 

manifested itself in the increase in viral diseases transmitted by 

certain species. This study proposes a first approach to these 

Thysanoptera, which have been very little studied in Algeria. An 

exploration and identification of thrips found on different host plants in 

coastal and sub- coastal Algeria were made for 2 consecutive years. 

The taxonomic study enabled the determination of 5 species. There 

were two Tubulifera-Phlaeothripidae, being Gynaikothrips ficorum 

(Marchal, 1908), a species strictly limited to Ficus retusa, and 

Haplothrips tritici (Kurdjumov, 1912), harvested from Anacylus 

clavatus and Avena sterilis, plus two Terebrantia-Thripidae, i.e. 

Odontothrips loti (Haliday, 1852), found on jasmine and Pittosporum 

tobira, and Frankliniella occidentalis (Pergande, 1895) collected from 

roses, Cucurbita pepo and Cucumis sativus. The fifth species was 

Limothrips cerealium (Haliday, 1836), which was harvested from 

Triticum durum, Triticum aestivum and Avena sterilis. 

- 4 - 

The Glypheoidea: a molecular study performed on the newly 

discovered species in the Coral sea 

Marie-Catherine Boisselier 1 , Nicolas Vidal 1 , Céline Bonillo 2 and 

Bertrand Richer de Forges 3 

1 

UMR 7138 – MNHN, 43 rue Cuvier, 75005 Paris, France. 

2 

IFR 101 – MNHN, SSM, 43 rue Cuvier, 75005 Paris, France. 

3 

UMR 7138 – IRD, B.P. A5, 98848 Nouméa Cedex, Nouvelle- 

Calédonie. 

The Glypheoidea is a group a lobster-like decapod crustaceans. Like 

monoplacophorans and the crossopterygians, they were long known 

from fossil specimens only. A male specimen of a living species, 

caught off the Philippines in 1908 at a depth of 185 meters, was kept 

in the collections of the Smithsonian Institution in Washington DC, but 

its identity was not recognized until 1975, when it was described as 

Neoglyphea inopinata by Forest & de Saint Laurent. Subsequent 

expeditions in 1976, 1980, and 1985, captured additional specimens 

at exactly the site of the Albatross 1908 station, and another 

specimen was trawled in the Timor Sea. In October 2005, a single 

female specimen of another glyphaeid species was discovered on a 

seamount in the Coral Sea, near New Caledonia, at a depth of 400 

meters. Nicknamed “Jurassic shrimp”, it received some attention from 

the media. This specimen represents a second genus and species of 

Recent Glyphaeoidea, and has been named Laurentaeglyphea 

neocaledonica (Richer de Forges, 2006; see also Forest, 2006). A 

molecular study based on 5 genes (two mitochondrial genes: COI and 

16S and three nuclear genes: H3, 18S and 28S) was performed in 

order to test the phylogenetic relationships of these living fossils 

among all the major decapod infraorders within the suborder 

Pleocyemata and to confirm or infirm Glypheoidea as the sister group 

of Astacidae. 

What the Internal Transcribed Spacer 2 of the Nuclear Ribosomal 

DNA Have to Say About the Systematics of Triatoma dimidiata 

Latreille, 1811 (Hemiptera: Reduviidae) from Jutiapa, Guatemala 

and Other Central American Populations? 

Andrea A. Cabrera 

Laboratorio de Entomología Aplicada y Parasicología, Escuela de 

Biología, Universidad de San Carlos de Guatemala. 12 calle 2-05 

zona 2, El Zapote, Guatemala 

Chagas disease is caused by the protozoan parasite Trypanosoma 

cruzy, and is transmitted by Triatominae bugs, mainly Triatoma 

dimidiata in Central America. Given that evidence suggests that the 

ribosomal DNA internal transcribed spacer 2 (ITS-2 rDNA) may 

become the DNA marker by excellence for species distinction and 

may as well be very useful for population differentiation within 

Triatominae. The sequences ITS-2 of three populations of T. dimidiata 

from Jutiapa, Guatemala, were analyzed and compared with other 

populations from Guatemala, El Salvador, Honduras, Nicaragua, 

Panama and Mexico. The sequence comparison, and phylogenetic 

analysis indicate that ITS-2 is a useful marker to establish some 

relationships in a population level for T. dimidiata. It was not possible 

to differentiate the populations from Jutiapa, El Salvador and 

Honduras, with no nucleotide difference detected among the 

sequences. Little differences were detected with Nicaraguan 

population. Two different clades were also formed, one with the 

population of Panama and the other with the populations of Mexico 

and Quiche (Guatemala). These results are consistent with the 

denoted forms of T. dimidiata dimidiata, T. d. capitata and T. d. 

maculipennis. The great differences in nucleotide composition and the 

early separation of the clade of Peten support the specific status of 

this population as T. sp. aff. dimidiata. This separation suggests that T. 

dimidiata populations followed different evolutionary paths and that 

problems may appear in the establishment of an effective control of 

the bug.


Towards a new paradigm for oceanic island biogeography: the 

contribution of molecular studies on landbirds 

Alice Cibois 1 , Jean-Claude Thibault 2 and Eric Pasquet 2 

1 Natural History Museum, Geneva, Switzerland 

2 Muséum National d’Histoire Naturelle, Paris, France. 

Two landmark theories dominated the field of island biogeography for 

the last decades: Mac Arthur and Wilson's dynamic equilibrium and 

Nelson and Platnick's vicariance biogeography. Recently, Heaney 

formulated a series of hypotheses that summarize aspects of an 

emerging paradigm, based on a number of studies using molecular 

approaches, and especially temporally calibrated phylogenies. These 

hypotheses include the frequency of long distance dispersal over 

large oceanic barriers, the occurrence of colonization from islands to 

continents, the intra-archipelago radiations, and the progression rule 

(sequential speciation inside an archipelago of volcanic origin). After 

reviewing these main points, we present here additional examples on 

landbirds, based on molecular phylogenies, which can be added to 

Heaney's proposal: for instance the existence of multiple colonization 

of a single island or archipelago, and the occurrence of "old" taxa on 

"young" islands. Moreover, human related extinctions on islands, for 

vertebrates, and particularly for birds, have been particularly 

numerous and we highlight the influence of these extinctions in the 

reconstruction of biogeographical scenarios. 

Is the biodiversity of moths in Africa still a black hole or is it 

becoming a little bit greyish? 

Jurate De Prins 

Royal Museum for Central Africa, Leuvensesteenweg 13, 3080 

Tervuren, Belgium 

While celebrating the 250 th anniversary of Linnaeus’ Systema Naturae, 

the lepidopterist community has a nice opportunity to evaluate the 

progress which was achieved in inventorying the Lepidoptera species 

over these years. However, we must sadly conclude that we still 

cannot provide even rough numbers of Lepidoptera species one can 

expect in tropical regions. Kristensen et al. (2007), in their overview 

on inventorying the moth and butterfly diversity, cited Heppner (1991) 

and provided the number 20,491 which could represent the 

Lepidoptera species diversity in the Afrotropical region. This 

commonly used estimation of species richness in the Afrotropical 

region is significantly underestimated. Herewith, we are presenting a 

taxonomic database of Afrotropical moths, which currently contains 

24,461 species-group names (18,670 valid species and subspecies). 

The database is still far from being complete. However, we already 

have recorded all Afrotropical species of the families Tineidae, 

Gracillariidae, Tortricidae, Crambidae, Saturniidae, and Sphingidae, 

while for all the other families at least the list of South Africa (Vári et al. 

2002) is included. Currently, we are working on the completion of the 

inventory of the Afrotropical moth species. Our data are based on the 

original fieldwork in different African areas and on published 

references. According to our estimation, the generally accepted figure 

of ca. 21,000 of the expected Lepidoptera species in the Afrotropical 

region, which was announced last year in the Linnaeus Tercentenary 

memorable publication (Kristensen et al. 2007) should probably be 

doubled or even tripled. 

- 5 - 

Can we resolve basal hexapods relationships? 

Cyrille A. D'Haese 

UMR 5202 CNRS "Origine, Structure et Evolution de la Biodiversité", 

Département Systématique et Evolution, Muséum National d'Histoire 

Naturelle, CP50 – Entomologie, 45 rue Buffon, 7500 Paris, France 

Despite many attempts to elucidate basal Hexapoda phylogeny, 

relationships of the primitively wingless hexapods in relation to 

Pterygota and the rest of Arthropoda remain equivocal and highly 

controversial. Hexapoda include three entognathous orders 

(Collembola, Protura and Diplura) and three ectognathous groups 

(Archeognatha, Zygentoma and Pterygota). Once it was understood 

that Apterygota (Collembola, Diplura, Protura, Archaeognatha and 

Zygentoma) were based on a symplesiomorphy (i.e. absence of 

wings), it was considered that Hexapoda should be divided in two 

clades: Entognatha and Insecta s.str. (= Ectognatha). The resolution 

of relationships among the entognathan orders Protura, Collembola 

and Diplura is controversial from the morphological and molecular 

points of view. Entognatha were traditionally considered monophyletic 

comprising Elliplura (Collembola + Protura) plus Diplura. However, 

some authors viewed Diplura as the sister group of Insecta. The 

monophyly of Diplura itself has been questioned. Among the Insecta 

s.str., the relationships between Archaeognatha, Zygentoma and 

Pterygota is better understood, but some questions remain such as 

potential paraphyly of the silverfish Lepidotricidae and Zygentoma 

s.str. with respect to other dicondylous insects. The position of 

Hexapoda within Arthropoda is also contentious and the very 

Hexapoda monophyly itself has been questioned recently. 

Background on traditional views on Apterygota phylogeny and recent 

results are commented. Estimate of basal hexapods relationships 

(addressing the placement of Protura, Diplura, Collembola, 

Archaeognatha, and Zygentoma with respect to Pterygota, 

Chelicerata, Myriapoda, and Crustacea) based on new data 

comprising a broad taxonomic and character sampling is proposed. 

Females do count: Documenting Chironomidae (Diptera) species 

diversity using DNA barcoding 

Torbjørn Ekrem 1 , Elisabeth Stur 1 , Kaare Aagaard 1 & Paul Hebert 2 

1 

Norwegian University of Science and Technology, Museum of 

Natural History and Archaeology, NO-7491 Trondheim Norway 

2 

Biodiversity Institute of Ontario, 579 Gordon Street, University of 

Guelph, Guelph, Ontario 

Canada N1G 2W1 

Because they are difficult to identify, female Chironomidae are usually 

neglected in biodiversity studies. We employed DNA barcoding to 

investigate if their inclusion would substantially increase the species 

count in an inventory of cold water springs. All specimens were 

sampled with emergence traps to ensure that both males and females 

were collected from the same surface area. When possible, 3-5 

specimens of each male morphotype were sequenced using standard 

DNA barcoding protocols. Most of the 216 males that were sequenced 

could be identified to species using existing literature, and all 66 

species were discriminated by barcodes. Barcode results further 

indicated the presence of 58 species among 96 females which were 

selected for analysis based on their morphological diversity. In total, 

93 species were recognised by morphology and barcode clusters. Of 

these, 31 species were represented by both sexes, 35 species by 

males and 27 species by females. Thus, despite the much lower 

sample size, almost 30% of the species were only represented by 

females. This result might reflect a high proportion of parthenogenetic 

species, biased sex ratios or general species rareness. We conclude 

that the inclusion of female chironomids, particularly at sites with 

many rare species or when sample sizes are small, will aid 

comprehensive species coverage.


Characteristics of habitats and reproduction of taiga birds in 

Yakutia 

Nikolai I. Germogenov 

Institute for Biological Problems of Cryolithozone SB RAS, 41 Lenin 

avenue, 677980 Yakutsk, Russia 

Yakutia is a large high-latitude region in the north-east of the 

Palearctic with extreme rigid conditions for bird habitats and their 

breeding comparable with adjacent areas by richness and abundance 

of avian fauna (308 species of 19 orders). Taiga as a natural zonal 

system plays a leading role for 87.0% of the regional avian fauna 

including 89.0% of nesting species. It is distinguished with 

supercontinental climate, landscape diversity, broad zonality and 

vertical belts, mountain dominance, specific heat-and water condition 

of perennially frozen grounds, low bioproductivity. Because of high 

watering and afforestation of the area its nest complex is rich in wood- 

and waterbirds. 

Reduced vegetative period is determined with deficit of the nesting 

time on the background of unpredictable conditions for breeding 

according to years and their seasonal instability, which become more 

expressed northwards and especially by mountains. This is leveled 

with local ecological-climatic conditions i.e. warming effect of water 

bodies, thermoarid effect of depressions, cryoarid influence of plains, 

well-developed floodplains, azonal landscapes. 

To maintain their proper existence and reproduction many birds 

change their stereotypes, which were previously acquired in the area 

optimum. The season of nesting is significantly shorter, proceeds in 

much earlier phenological conditions and its initial stages are shifted 

for a calendar subsequent time. Having abundant monocyclic species 

with small clutches, of them over 80% lay similar number of eggs or 

sometimes even greater than in the other parts of the area. 

Birds are mainly carnivorous, plastic when choosing food depending 

on its abundance and availability. Impoverishment of food resource 

composition under an increasing mosaic pattern while spreading and 

simultaneous abundance of some food items causes narrowing of 

nutrition spectra and limitation of the number of main food items for 

birds and their nestlings. 

Rare and protected birds in South Yakutia 

Nikolai I.Germogenov, Victor G. Degtyarev, Arkady P. Isayev, Nikolai 

N. Egorov and Vasily V. Okoneshnikov 


avenue, 677980 Yakutsk, Russia 

South Yakutia is the largest quickly developing industrial region of 

Siberia and Far East due to its natural resources development (gold, 

gas, oil, coal, wood, water power) and their supply through project oilgas 

pipelines, network of car and railway lines, transmission lines 

purposed at inner and outer markets. Besides, the region is 

distinguished with rich biodiversity; here live the Evenks – indigenous 

not numerous North people leading traditional nature management 

(hunting, reindeer breeding). 

A weak ornithological component of biodiversity as the basis of 

regional ecological balance is rare and endangered species. 69 rare 

bird species including 51 protected are currently recorded in Yakutia, 

of them 16 joined the Russia Red Data Book (additionally to casual 

Mandarin duck and Asian Marble murrelet), 9 are in IUCN Red List of 

Threatened Animals (additionally to Falcated teal, Yellow-breasted 

bunting and Black-tailed godwit), even more species are in 

Threatened Birds of Asia, CITES Supplements, Bonn Convention on 

conservation of migratory animal species, migratory birds and their 

habitats signed by Russia and a number of countries. Yakutia has key 

ornithological territories of national and international importance. In 

some parts of the Aldan basin during migrations over 10% of the 

world Siberian crane population, nearly 1% of Baikal teal are recorded; 

large breeding groups of Hooded crane and preserved local Siberian 

jay population is also reported here. 

The coming transformation of South Yakutia ecosystems demands 

optimization of species and territorial forms of biodiversity 

conservation. 

- 6 - 

Eastern population of Siberian crane: structure, habitats, nesting 

and migration 

Nikolai I. Germogenov, Nickita G. Solomonov, Anatoli E. Pshennikov, 

Andrey G. Degtyarev, Sleptsov M. Sleptsov, Nickolai N. Egorov, Inga 

P. Bysykatova, Maria V. Vladimirtseva and Basili V. Okoneshnikov 


avenue, 677980 Yakutsk, Russia. 

Siberian crane Grus leucogeranus (Pallas, 1773) is an endangered 

species of the world fauna. It is represented by a western population 

comprising a few pairs and nesting in West Siberia with their winter 

grounds in Iran and an eastern one breeding in northeastern tundras 

of Yakutia. Owing to the efforts of scientific nature conservation 

agencies of the countries concerned including communities the latter 

is in relatively safety condition – up to 4000 birds are reported on their 

winter grounds in China. This presentation summarizes research 

materials conducted with participation of foreign partners during bird 

banding and use of Platform Transmitter Terminals (PTT) within the 

range of the population in 1990-2007. 

There are three megapopulations of nesting Siberian crane over 

Yakutia. The study of the Indigirka reproductive habitat shows that 

Siberian cranes inhabit wetlands (near lake and boggy sites) located 

as high as 14-19 m a.s.l. neighboring upon large lakes in a mosaic 

and irregular pattern. These wetlands comprise 21-26% of the territory, 

their state is defined by permafrost condition. Because of abrasia and 

soil ice thaw the habitat areas are reducing. In the optimal grounds 

nesting density is 5 pairs/100 km 2 . The pairs keep the same sites, 

which have nests used many times or once irrespective of the 

breeding participation. 

Brood occurrence is 4.3-83.0% but availability of very successful 

breeding cycles is small (12.5%). The population stock are pairs 

(89%), of them 50% are aged 20 years old and over. 

In Yakutia fall migration proceeding in a narrow corridor is very busy 

and lasts 20-25 days. In the north-east of China in Jilin and Laonin 

Provinces transit concentrations of 500-1500 birds are observed for a 

month. 

On the diversity and phylogeography of freshwater snails of the 

genus Pachychilus in Guatemala: How many species are there? 

Maria Gomez 1 , Frank Köhler 2 ; Thomas von Rintelen 1 and Matthias 

Glaubrecht 1 

1 

Museum für Naturkunde, Humboldt-Universität, Invalidenstrasse 43, 

10115 Berlin, Germany 

2 

Malacology Section, Division of Invertebrate Zoology, Australian 

Museum, 6 College Street, Sydney NSW 2010, Australia 

Situated right in the heart of Central America, one of earth’s 

biodiversity hotspots, Guatemala is characterized by two rugged 

mountain chains that were uplifted during a dynamic geological history. 

The dramatic contrasts between the lowlands and highlands promote 

a huge diversity of ecosystems that harbour a mega-diverse and due 

to their geographical isolation largely endemic biota. 

The present study focuses on one of the more conspicuous elements 

of the macro-invertebrate fauna of the rivers of Guatemala, snails of 

the genus Pachychilus I. & H. Lea, 1850. These molluscs are 

considered as an essential element of the riverine food-web. However, 

there is little understanding of their systematics, diversity, and ecology. 

A phylogeographic study based on samplings from a large number of 

localities across the whole country and on two mitochondrial genes 

(COI, 16S), provides evidence for the existence of various well 

differentiated and likely endemic river clades. 

In contrast to this significant genetic differentiation, we found that 

morphological variation between most species is often subtle. We 

conclude that due to the high degree of superficial similarity, one 

might underestimate the true diversity of the group. In fact, we are 

convinced that most of the 27 described taxa from Guatemala should 

be considered as valid species. These species are usually restricted 

to rather small areas, such as single rivers or river systems. We 

anticipate that future revisionary work will probably lead to the 

identification of further, yet unrecognized species.


Absence of the articular disk in the Tasmanian Devil 

temporomandibular joint 

Katsuhiko Hayashi 1 , Masashi Sugisaki 1 , Koji Kino 2 and Takayuki 

Ishikawa 2 

1 

Department of Dentistry, Jikei University School of Medicine, Tokyo 

Japan, 

2 

Temporomandibular Joint and Occlusion, Comprehensive Oral 

Health Care, Comprehensive Patient Care, Graduate School, Tokyo 

Medical and Dental University, Tokyo Japan 

The articular disk of the temporomandibular joint is a constant 

structure in the Mammalia. According to Parson’s report in 1900, 

however, it was absent in four animals: the Armadillo, two kinds of 

Monotremes (Echidna and Ornithorhynchus anatinus), and the 

Tasmanian Devil. Thereafter, no research has been done to confirm 

this observation. The aim of this study was to determine by 

anatomical and histological examination whether the Tasmanian devil 

has an articular disk in its temporomandibular joint. 

Fresh corpses of eight Tasmanian Devils were obtained from the 

School of Zoology, University of Tasmania. They were dissected and 

the structure of the temporomandibular joint was carefully observed 

anatomically. Then, the temporomandibular joint was removed, 

immersed in 10% buffered formaldehyde solution, decalcified in 10% 

ethylenediaminetetraacetic acid solution and embedded in paraffin. 

Serial sagittal sections were cut and stained with hematoxylin and 

eosin for histological examination. In all cases, gross observation and 

dissection revealed the absence of an articular disk. Histologically, the 

surface layer of both the condyle and the glenoid fossa consisted of a 

quite thick fibrous tissue as compared with those of other 

mammalians. A synovial membrane-like structure was observed in the 

anterior and posterior parts of the fibrous structure of the condyle. We 

confirmed the absence of an articular disk in the Tasmanian Devil’s 

temporomandibular joint. Furthermore, a thick fibrous layer on the 

surface of both the condyle and the glenoid fossa, might play a role as 

a buffer against hard jaw movement, instead of articular disk. 

Distribution pattern of Antedon in the Mediterranean: a story of 

vicariance following the Messinian salinity crisis 

Lenaig Hemery 1 , Marc Eleaume 1 , Pierre Chevaldonne 2 , Agnès Dettai 3 

and Nadia Ameziane 1 

1 

Museum National d’Histoire Naturelle, UMR 5178-BOME, 

Département Milieux et Peuplements Aquatiques ; 

2 

Centre 

d’Océanologie de Marseille, CNRS-UMR 6540 DIMAR, Station Marine 

d’Endoume ; 3 Museum National d’Histoire Naturelle, UMR 7138, 

Département Systematique et Evolution, Paris, France 

Crinoids constitute one of the five extant classes of the phylum 

Echinodermata. Crinoid classifications generally separate the stalked 

sea lilies from the stakless comatulids. However, phylogenetic 

reconstruction based on morphologic and molecular data seem to tell 

a more complex story. The so-called “stalked” crinoids appear para- 

or polyphyletic and the loss of the stalk may have happened several 

times. The main goal of this work was to come to a better 

understanding of the inter-relationships within the crinoids and among 

the species of the genus Antedon, because Antedon may appear to 

be a model organism to study the recolonization in the Mediterranean 

after the messinian salinity crisis, ~ 5 My ago. The phylogenetic 

reconstruction of the crinoids was based on one nuclear (18S) and 

one mitochondrial gene (COI) amplified for 65 specimens 

representing 23 out of 28 of the recognized crinoid families. The 

Antedon phylogeny was done using the mitochondrial cytb gene for 

27 specimens representing seven nominal species. Phylogenetic 

trees were generated using the maximum parsimony and maximum 

likelihood criterion. Results show that the genus Antedon is not 

monophyletic. Atlantic and Mediterranean species form a well 

supported clade. Antedon bifida is distributed in the Atlantic Ocean 

and the Alboran Sea while A. mediterranea is distributed in the rest of 

the Mediterranean. These results suggest that the divergence 

between A. bifida and A. mediterranea may have occurred by 

vicariance after the setting of an ecological barrier to larval dispersion 

in the Alboran Sea following to the messinian salinity crisis. 

- 7 - 

Diversification in Mesoamerica: the case of hummingbirds 

(Trochilidae) 

Blanca E. Hernández-Baños 1 , Nandadevi Cortés-Rodríguez 1 , 

Gabriela M. García-Deras 1 and Jaime García-Moreno 2 

1 Departamento de Biología Evolutiva, Facultad de Ciencias, 

Universidad Nacional Autónoma de México, Mexico City. 2 Centre for 

Biodiversity Conservation Mexico and Central America, Conservation 

Internacional, San Pedro, Costa Rica 

The Trochilidae (hummingbirds) is one of the most species-rich 

families of birds (approximately 331 species distributed in 104 genera) 

and, therefore, it is considered a model system of evolutionary 

diversification. The phylogeny of major clades is well supported by 

analyses of DNA sequences, but the relationships at the genus and 

species levels are not clear. In particular, in Mesoamerica (North of 

Mexico to Panama) there are 89 hummingbird species pertaining to 

41 genera, which have been considered to have originated and 

diversified within Mesoamerica. However, this hypothesis has been 

proposed considering only groups distributed in the highlands and it is 

not clear if it applies when groups inhabiting the lowlands are included 

in the analyses. We present the results of molecular phylogenetic 

studies, employing mitochondrial and nuclear genes, of two groups 

inhabiting the highlands (the genus Lampornis and the Lampornis 

amethystinus complex) and of two groups distributed in the lowlands 

of Mesoamerica (Amazilia rutila and Cynanthus latirostris complexes). 

Our phylogenetic analyses indicated that both the highland and the 

lowland taxa had their origin and diversified within Mesoamerica. Our 

analyses also showed that for the highlands taxa the Isthmus of 

Tehuantepec (Mexico) has been an important barrier promoting 

differentiation between populations, whereas Neovolcanic Belt 

(Mexico) has not. In contrast, in the lowland taxa we found 

differentiated evolutionary units without clear physical barriers, with 

the exception of populations inhabiting the Tres Marías islands 

(Mexico). 

A hidden diversity of snakes in Australia 

Marie Jacquot 1 , Steve Donnellan 2 , S. Blair Hedges 3 , William R. 

Branch 4 , Richard Thomas 5 and Nicolas Vidal 1,3 

1 

UMR 7138, Systématique, Evolution, Adaptation, Département 

Systématique et Evolution, 

C. P. 26, Muséum National d’Histoire Naturelle, 43 Rue Cuvier, Paris 

75005, France. 

2 

Evolutionary Biology Unit, South Australian Museum, North Terrace, 

Adelaide, SA 5000, Australia. 

3 

Department of Biology, 208 Mueller Laboratory, Pennsylvania State 

University, University Park, PA 16802-5301 USA. 

4 

Bayworld, P.O. Box 13147, Humewood 6013, South Africa. 

5 

Department of biology, PO box 23360, University of Puerto Rico, 

San Juan, Puero Rico 00931-3360, USA. 

Snakes are considered to be well-known in terms of diversity (~3070 

sp.), and have a slower rate of discovery of new species than frogs, 

lizards, or mammals. However, the Scolecophidia, or blindsnakes 

(~370 sp.), which burrow and occur on most continents, are unusual 

in having a very conservative morphology linked to their very 

specialized fossorial mode of life. This causes difficulties in 

distinguishing species. Here, we show that mitochondrial DNA 

sequences of blindsnakes from Australia (genus Ramphotyphlops) 

reveal a hidden diversity, corresponding to a rate of an undescribed 

species for each described species. This suggests that global 

blindsnake diversity may be underestimated by more than 50%. A 

greater diversity of these mostly tropical vertebrates with small ranges 

would have implications for protected areas, biodiversity hotspots, and 

conservation practices.


Patterns of skeleto-muscular connectivity: brackets across 

development and evolution 

Georgy Koentges 1 , B. Ryll 1 , K. Vance 1 , Sacha Ott 1 , D. Woodcock 1 , D. 

Rand 1 , Toshiyuki Matsuoka 4 , Paul Tafforeau 2 and Per E. Ahlberg 3 

1 Warwick Systems Biology Centre, University of Warwick, CV4 7AL, 

UK ; 2 European Synchrotron Radiation Facility, Grenoble, France ; 3 

Evolutionary Biology Centre, Uppsala University, Sweden ; 4 Kyoto 

University, Japan 

Our work concerns questions of how transcriptional decision making 

in cells of vertebrates is controlled in such a way that it affects 

patterning in embryos, resulting adult morphologies and their deeper 

macroevolutionary transformations (Koentges, Nature Feb 2008). This 

has three aspects. Based on previous discoveries of cryptic 

boundaries of cell populations carrying distinct molecular and cellular 

identities (Matsuoka et al. Nature 2005) and that are precisely 

reflected in muscle attachment systems we have now gone into the 

fine-histology of fossils, using latest synchrotron-based imaging, in 

order to trace muscle attachment patterns and thereby cellular 

lineages in fossils many hundred million years old. I will report latest 

findings in this area. Secondly, I will try to outline the basics of genetic 

fate mapping that allow us to trace the anatomical impact of early 

embryonic decision making processes with single cell resolution. We 

will look at other cryptic boundaries in the vertebrate head. Thirdly, I 

will investigate the mechanics of the transcriptional process by 

showing latest results from our efforts in Warwick to discover cisregulatory 

regions, to study their function in massively parallel singlecell 

assays using a novel imaging platform we have established and 

describe their action and combinatorial logic in mathematical terms (of 

stochastical differential equations). This might provide a substantive 

experimental and theoretical foundation for a future functional 

comparative analysis of gene-regulatory regions affecting patterns 

and speeds of gene expression across evolutionary history, and can 

help us explain fundamental aspects of gene-regulatory change in a 

macro-evolutionary perspective. 

Towards cyber exchange facilities for systematic studies 

Elise Kuntzelmann, Visotheary Rivière-Ung and Régine Vignes-Lebbe 

MNHN, 43 rue Buffon, 75005, Paris, France 

Taxon names are the main access keys to biological information and 

the link between the real biological world and the conceptual world 

proposed by systematists through the description and characterization 

of taxa. The generalization and the integration into current taxonomic 

work of knowledge base management systems may be the next 

revolution in taxonomy. International projects are already opening the 

way of this new, delocalized, taxonomy. As an example, the european 

union project EDIT1 aims to contribute to change current taxonomic 

practices. EDIT is a 5 years european network started in 2006 

including 21 major natural history institutions and several 

organisations. An important goal of this program is to reduce 

fragmentation, encourage durable integration of institutions and to 

promote collaborative research using biodiversity informatics. A main 

part of EDIT is dedicated to biodiversity informatics and to the creation 

of an internet platform for cybertaxonomy based on a Common Data 

Model (CDM). The aim of this platform is to support the taxonomic 

work process through applications and services. In this context 

software tools for descriptive data were inventoried and a number of 

them selected for further testing. In order to formalize data exchange 

between all the tools, an XML based standard has been adopted by 

the Taxonomic Data Working Group (TDWG2): the Structured 

Descriptive Data (SDD). We focus here on the program Xper2 3. It is 

a state-of the- art taxonomic management system for the storage, 

edition and on line distribution of taxonomic knowlege. To do this, 

Xper©˜ has is own data format but it can also be connected to the 

cyberplatform thanks to the SDD format. There is a complete 

export/import procedure from Xper©˜ to SDD format. Taxonomy has 

become a planetary-scale science and deserves a planetary-scale 

tool (Wheeler, 2004). All this contributes to the common objective of 

main inter-connected projects like GBIF, BIOTA, Catalogue of Life etc.: 

to provide a free web access to biodiversity information. 

- 8 - 

Barcoding gene COI fails to distinguish between two fiddler 

crabs (Brachyura: Ocypodidae: Uca) across their entire range of 

geographic overlap 

Richard B. Landstorfer 1 , Christoph D. Schubart 1 and Darryl L. Felder 2 

1 Fakultät für Biologie I, Universität Regensburg, 93040 Regensburg, 

Germany ; 2 Dept. of Biology, University of Louisiana at Lafayette, 

Lafayette 70504, USA 

Uca minax (Le Conte, 1855) and Uca longisignalis Salmon & Atsaides, 

1968 are two closely related fiddler crab species from the 

northwestern Atlantic. Uca longisignalis is endemic to the northern 

Gulf of Mexico including northwestern Florida and northern Texas. 

The geographic range of Uca minax is wider and includes most of the 

American East Coast up to Massachusetts with a disjunct distribution 

across the Peninsula of Florida. Following the description of Uca 

longisignalis by Salmon and Atsaides, there was a long-standing 

debate on the validity of this species. However, following a redescription 

in 1982, it has been accepted by most systematists. 

Several diagnostic morphological traits, like differences in pubescence 

and in color, allow us to distinguish the two sister species. 

Intraspecific allozyme divergences in trans-Floridian populations in 

Uca minax have also raised the question of whether the Gulf of 

Mexico hosts an endemic lineage of this species. Our studies include 

populations in the region of sympatry for the two species as well as 

regions where Uca minax (along the Carolinian Province) and Uca 

longisignalis (south-central Texas) occur alone. Samples of at least 

ten specimens each from separate populations were examined for 

morphological characters and color; thereafter, six to ten specimens 

from eight populations were used for genetic examination with the 

barcoding gene cytochrome oxidase I (COI). Results are presented as 

phylogenetic networks. Both species are characterized by high 

haplotype diversities, but limited geographic structuring. The amount 

of gene flow within and between species was calculated with AMOVA. 

As opposed to the morphology, our COI analysis does not allow 

distinction between these two species, suggesting a very recent 

separation, possibly during Pleistocene isolation. This is one more 

example, where COI barcoding methods fail to recognize actual 

species diversity. 

Phylogeography of the Siberian roe deer (Capreolus pygargus 

pallas, 1887) 

YunSun Lee, Nickolay Markov, Inna V. Voloshina, Alexander I. 

Myslenkov and Irina Sheremetyeva 

Seoul National University College of veterinary Medicine 85-803, San 

56-1, Sillim-dong, Gwanak-Gu, Seoul 151-742, South Korea., 151-742, 

Seoul, KOREA 

The Siberian roe deer (Capreolus pygargus) is widely distributed in 

Eurasia, including Russian Siberia, Yakutia, Transbaikalia, Russian 

Far East, Northern Mongolia, Northern China and Korea. In a number 

of these areas, the population fluctuate significantly due to overhunting 

and habit! at degradation. This has resulted in the formation of 

fragmented and isolated populations within their range. Information on 

the genetic diversity and phylogeographic structure of the Siberian roe 

deer populations would be very helpful for the proper management of 

the populations. In the present study, mitochondrial cytochrome b and 

control region sequences of roe deer samples from Russian Siberia, 

Trans-Baikal, Amurskyi region, Primorskyi Krai, Korean peninsula and 

Jeju Island were utilized to evaluated genetic diversity and 

phylogeographic structure of the species. The phylogenetic trees and 

network analysis indicated that there were three main lineages in 

Siberian roe deer. Two of them, present in mainland, do not show any 

geographic affinities suggesting historical gene flow among the 

populations. The third group is composed entirely of individuals from 

Jeju Island. Nucleotide and haplotype diversities of C. pygargus in 

Jeju Island were much lower than those in Russia a! nd mainland 

Korea probably due to founder effect during the spatial isolation of the 

population from the mainland. In contrast, the levels of genetic 

diversity in mainland groups are comparable to other populations of C. 

pygargus and C. capreolus. To understand their detailed 

phylogeographic structure, analysis with a faster evolving genetic 

markers such as microsatellites would be needed.


Surprizing findings in the teleostean large-scale 

interrelationships 

Blaise Li, Agnès Dettai and Guillaume Lecointre 

MNHN, cp 26, 43 rue Cuvier, 75005, Paris, France 

Molecular phylogenetics comes to an age where different teams work 

on various genes sequences for large comparative samples of 

biodiversity. As a result a number of orders (e.g. Lophiiformes, 

Tetraodontiformes, Gadiformes, Zeiformes, etc.) are now present in a 

same data matrix, a phenomenon that was rare ten years ago and 

almost non existent from morphological matrices; and prone to new 

phylogenetic discover! ies. Our strategy is to sequence nuclear 

phylogenetic markers appropriate for acanthomorph (spiny teleosts: 

16,000 species) large-scale interrelationships, analyze them 

separately and simultaneously, and extract from these informations an 

reliability index for clades. A number of reliable clades are rather 

surprizing for classical zoology: labrids ans scarids are not close to 

cichlids; flatfishes are related to jacks, remoras and barracudas; 

mackerels are more closely related to stromateioids than to billfishes; 

gadiforms are closely related to zeioids; lophiiforms go with 

tetraodontiforms and acanthuroids, caproids and chaetodontids; 

blennioids and gobiesociforms are closely related to atherinomorphs... 

Among new results, howellids, epigonids and lateolabracids group 

together; moronids, elassomatids and centrarchids group together; 

plesiopids are related to mugiloids; indostomids go with anabantoids 

and swamp eels while sticklebacks are related to cottoids and 

zoarcoids. 

A completely new picture of higher teleost interrelationships emerge 

from several genes and teams where a number of "classical" groups 

are polyphyletic (like paracanthopterygians, acanthopterygians, 

percomorphs, perciforms, percoids, trachinoids, labroids, scombroids, 

serranids, gasterosteiforms, zeiforms) while others are maintained 

because they appear monophyletic (gadiforms, lampridiforms, 

tetraodontiforms, lophiiforms, pleuronectiforms, notothenioids, 

cottoids...). 

The Varan-ID project : Online determination for monitor lizards. 

Öna Maiocco 1 , Aurélien Miralles 2 and Régine Vignes-Lebbe 3 

1 2 nd year Master student at MNHN; 2 Laboratoire évolution, genomes 

et spéciation, CNRS, 91198 Gif-sur-Yvette CEDEX, France; 

3 Laboratoire Informatique et Systématique UPMC, UMR 5143 

Paléobiodiversité, MNHN CP 48, 57 rue Cuvier, 75231 Paris cedex 05, 

France 

Monitor lizards are very popular and subjected to an intense trade 

over the world, for their skins as well as for herp-breeding. A lot of 

information on monitor lizards is available for the general public but 

identification keys remain the prerogative of scientific publications. As 

many people involved with monitor lizards are not herpetologists 

(keepers, customs officers, etc.) and do not have such literature at 

hand, we have created an online general key dealing with the 60 

monitor lizards species. Varan-ID is available at the url: 

http://lis.snv.jussieu.fr/apps/xper/data/varanID. It uses a free software 

(Xper2) developed by the LIS (Laboratory of Computer Science for 

Systematics, Pierre and Marie Curie University). It offers multiple 

access to six knowledge bases describing all Varanus species and 

online determination keys. Such free-access keys are useful to make 

identification successful in a lot of contexts, for example even if some 

parts of the animal are missing. The knowledge bases include 

structured taxonomic descriptions and illustrations. They can be linked 

to specimens in collection databases, and to any additional electronic 

resources (molecular databases, bibliographic references etc.) without 

duplicating information. The update of such a system is greatly 

facilitated and it allows bringing closer taxonomic research and users 

of taxonomy. Varan-ID displays data on Varanus and several results 

computed automatically on each knowledge base: species diagnoses, 

printable keys, readable taxonomic forms, similarities etc. For further 

information on the new generation of computer-aided tools for, see 

the EDIT program for cybertaxonomy: http://www.etaxonomy.eu/wp.php?menuwhat=1&wp=5&action=1 

and the website 

dedicated to the DELTA system : http://delta-intkey.com. 

- 9 - 

Animal taxonomy in the light of recent progress in 

developmental biology 

Alessandro Minelli 

University of Padova, Department of Biology, Via Ugo Bassi 58 B, I 

35131 Padova, Italy 

Evolutionary developmental biology (evo-devo) has its twin roots in 

two disciplines - evolutionary biology and developmental biology - 

which until recently have been progressing along independent routes 

and have independently contributed to progress in animal taxonomy. 

Today, even if still in its infancy, evolutionary developmental biology 

can effectively contribute in original way to the main steps of 

phylogenetic analysis: (a) evaluation of homology, especially in 

helping the identification of developmentally independent units; (b) 

selection of characters; and (c) assessment of character polarity. 

We can expect that important progress in animal taxonomy will derive 

from a careful evaluation of the evolvability of traits. The relationship 

between taxonomy and evo-devo can be also considered from the 

reciprocal perspective, of how taxonomy can contribute to progress in 

evo-devo. This contribution is being obtained (a) by providing 

informed insights towards the selection of new model species for 

experimental studies, (b) by offering the tools of cladistic methodology 

for an employ in the study of heterochrony, and especially (c) by 

providing phylogenetic scenarios onto which to map comparative data 

concerning developmental processes or their outcome. 

Phylogenetic relationships of the Asian palm civets (Hemigalinae 

& Paradoxurinae, Viverridae, Carnivora) 

Marie-Lilith Patou 1 , Régis Debruyne 2 , Andrew P. Jennings 1 , Akbar 

Zubaid 3 , Jeffrine Japning Rovie-Ryan 4 , F. Satoshi 5 and Géraldine 

Véron 1 

1 Unité Origine, Structure et Evolution de la Biodiversité (CNRS UMR 

5202), Département Systématique et Evolution, MNHN, CP 51, 57 rue 

Cuvier, 75231 Paris Cedex 05, France ; 2 McMaster Ancient DNA 

Centre Department of Anthropology McMaster University Chester 

New Hall Rm. 524 1280 Main Street West Hamilton, Ontario, Canada 

- L8S 4L9; 3 School of Environmental & Natural Resource Sciences, 

Faculty of Science & Technology, University Kebangsaan Malaysia, 

43600 UKM, Bangi, Malaysia; 4 Ex-Situ Conservation Division, Dpt of 

Wildlife & National Parks (DWNP), Peninsular Malaysia, Ministry of 

Natural Resources and Environment (NRE), KM 10 Jalan Cheras, 

Kuala Lumpur, Malaysia; 5 Institute for Biodiversity, DWNP, NRE, 

28500 Bukit Rengit Lanchang, Pahang, Malaysia. 

The Viverridae (Mammalia, Carnivora), one of the least studied 

groups of carnivorans, include two subfamilies of Asian palm civets: 

Hemigalinae and Paradoxurinae. The relationships between and 

within these two subfamilies have never been thoroughly tested using 

an extensive molecular sample set. In this study, we gathered 

sequences of four genes (two mitochondrial: Cytochrome b and ND2 

and two nuclear: β-fibrinogen intron 7 and IRBP exon 1) for eight of 

the eleven extant species representing these two subfamilies in order 

to shed some light on their evolutionary history. The results showed 

that: (1) the Asian palm civets (Hemigalinae and Paradoxurinae) have 

a single origin and form the sister group of the (Genettinae + 

Viverrinae) clade, (2) the Hemigalinae (including the otter civet 

Cynogale bennettii) are monophyletic, (3) the Paradoxurinae are 

monophyletic and (4) the small-toothed palm civet (Arctogalidia 

trivirgata) is an early offshoot within the Paradoxurinae, exhibiting 

peculiar morphological characters. Using a relaxed molecular clock 

analysis, the differentiation of the (Hemigalinae + Paradoxurinae) was 

inferred to occur in the Late Oligocene / Early Miocene.


Integrative taxonomy in a hyperdiverse group of marine molluscs: 

the Turrinae (Gastropoda, Conoidea). 

Nicolas Puillandre 1,2 , Sarah Samadi 1 , Marie-Catherine Boisselier 1 and 

Philippe Bouchet 2 

1 

MNHN, Service de Systématique Moléculaire, 43 rue Cuvier, 75005 

Paris, France. 

2 

MNHN, 55 rue Buffon, 75005 Paris, France. 

Conoidea, which includes subfamily Turrinae, is a hyperdiverse group 

of marine gastropods, with about 10000 living species (Bouchet 1990) 

defined mostly by shell characters. The great and sometimes 

continuous variability of these morphological characters renders the 

delimitation of species complicated. Recently, molecular taxonomy 

has received increasing attention (Vogler 2006, Arhens 2007) to 

better define species boundaries in poorly known groups, where 

traditionnal characters are difficult to assess. Recent cruises in West- 

Pacific have permitted the achievement of a sampling comprising 

almost 1000 specimens of Turrinae. Species were not a priori defined 

in order to accurately estimate intra and interspecific variabilities. 

Sequencing was performed for two genes: the COI mitochondrial 

gene used for the barcode project and a portion of the 28S nuclear 

gene. Species boundaries were defined using the “barcode gap” as a 

threshold but also phylogenetic reconstructions associated to the 

method described in Pons et al. 2006. More than 80 clusters were 

thus delimited, mostly including several specimens. Geographic and 

morphologic informations were then used to discuss alpha-taxonomy 

of the group. If some clusters appeared easily recognisable on the 

basis of morphological characters, some gave the impression to 

include specimens with identical shells, while several shell 

morphologies were observed in others. Indeed, molecular taxonomy 

appears to be a fast and accurate way to define clusters of individuals 

within Turrinae. 

New perspectives in the systematics of Limnadiid clam shrimps 

(Crustacea, Branchiopoda, Spinicaudata): cyst morphology 

provides solid elements to the revision of this group in the world 

Nicolas Rabet 

UPMC, UMR 7138 CNRS UPMC MNHN IRD, Case 05, 7 quai St 

Bernard, F-75005 Paris, France. 

The shape and surface ornamentation of cysts, resting embryos, are 

very useful for systematic studies of Branchiopoda. These characters 

were already frequently used in Anostraca at the genus and 

sometimes species level (Thiéry 1996) but some important specific 

variations were also recently described (Thiéry et al. 2007). Within the 

Spinicaudata the cyst shells are always spherical, except in 

Limnadiidae where they can also be cylindrical or spiralled with 

numerous intermediate shapes. Contrary to Anostraca, the systematic 

based on adult morphology is very difficult in limnadiid clam shrimps 

because morphology is highly variable and classical taxonomy is 

generally very confused. Some studies suggested that the variation of 

shape and ornamentation of cysts are the best morphological 

characters for the taxonomy of the genus Eulimnadia in America (Belk, 

1989, Martin, 1989). Here I present the examination of numerous 

specimens from different parts of the world (including newly collected 

samples and samples from museum collections). In each population 

the cyst morphology is relatively stable, confirming its interest 

throughout the world to identify species of Eulimnadia and also of 

other genera (Metalimnadia and a new genus). In addition, cyst 

morphology allows species identification in the absence of adult 

specimens by sampling soils from temporary pools, which could 

increase rapidly our knowledge on this group. 

Thiéry A. 1996. In: Grassé PP, ed., Traité de Zoologie, Anatomie, 

Systématique, Biologie 7: 287–351. Paris: Masson. 

Thiéry A, Rabet N, Neve G. 2007. Biol J Linnean Soc 90(1): 55-60. 

Belk D. 1989. J Crust Biol 9(1): 115-125. 

Martin JW. 1989. J Crust Biol 9(1): 104-114. 

- 10 - 

EDIT: creating a new environment in taxonomic institutions 

Simon Tillier 

EDIT, CP43, Muséum national d'Histoire naturelle (UMR 7138), 57 

rue Cuvier, 75231 Paris cedex 05, France 

Methodological, scientific and technical progress is rapidly 

transforming taxonomy and its environment, making these times more 

exciting than ever for taxonomists. Pivotal to the development of 

taxonomy are the rapidly expanding fields of high throughput DNA 

sequencing, automated digital data-gathering and biodiversity 

informatics. Incorporating these technologies will be critical to the 

future of the science of taxonomy. 

To employ these new tools, scientific collaborators and users of 

taxonomy will require new ways of working and interacting with 

taxonomists. Taxonomic scientists integrated into interdisciplinary 

teams will be an essential way of working. There will have to be 

substantial, even radical, changes in how taxonomy is done and its 

supporting infrastructure operated, to exploit these opportunities to the 

full. 

In support of these radical changes in the way taxonomy is done and 

made available to its users, EDIT is a consortium of the largest 

taxonomic institutions in the world which unite their strengths to 

develop and accompany the necessary change in infrastructure 

planning, functioning and management. Beside progress toward 

integration of research infrastructures, EDIT is elaborating the 

computer tools which will be instrumental to further developments of 

taxonomic research, and has set up projects for All Taxa Biodiversity 

Inventories in Europe experimenting new standards and methods. 

Metazoan phylogenetics: “surprising new results” and the 

deceptive phylogenetic signal 

Johann-Wolfgang Wägele 

Zoological Research Museum Alexander Koenig (Bonn, Germany): 

During the last years a large number of new, often mutually 

incompatible hypotheses on the phylogeny of animals have been 

published, most of them based on molecular data. The Ecdysozoa 

hypothesis is not compatible with the Coelomata, the clade 

Myriochelata disrupts the Mandibulata and Tracheata, the Serialia is 

not compatible with the Conchifera, and at the base of the metazoan 

tree we find alternatively sponges, placozoans, or ctenophorans. Each 

author or team of authors claim to have found convincing evidence. 

Obviously, some profound misunderstandings are misleading us. A 

major issue is the missing analysis of data quality. Statistical support 

values on trees are not adequate signs for the quality of data. 

Different types of long branch artefacts require different measures. 

Improvement of substitution models helps, but does not compensate 

for the absence of phylogenetic information in raw data. New tools for 

the evaluation of the suitability of alignments still have to be 

developed.


Connectivity and speciation processes in an oceanic 

seamounts system: comparative phylogeography of 

Gastropods with contrasting reproductive strategies 

Magalie Castelin 1 , Philippe Bouchet², Marie-Catherine Boisselier 1 

and Sarah Samadi 1 

1 Department of Systematic and Evolution, The Natural History 

Museum, Paris 43 rue Cuvier 75005, France; 

2 Department of Systematic and Evolution, The Natural History 

Museum, Paris 55 rue Cuvier 75005, France; 

Species distribution and speciation processes in the deep-sea 

remain largely unknown. It has been suggested that seamounts, 

which vary greatly in their faunal assemblage and display many 

original species, may be center of speciation. Their spatial 

distribution represents a fragmented habitat, which may locally 

increase speciation rate by breaking up species in small isolated 

populations. Another viewpoint suggests that seamounts, which are 

highly productive oases, receive large trophic input that allow 

abundance of species and multiplication of ecological niches. As 

such, seamounts could serve as suitable place for parapatric 

speciation by accommodating species to colonize new ecological 

niche. The phylogeographic survey of seven Gastropods species 

with contrasted reproductive strategies from seamounts near New 

Caledonia reveals patterns supporting either allopatric or parapatric 

speciation models. Allopatric diversification pattern is found for 

organisms with low dispersal abilities, whereas parapatric pattern is 

observed for a couple of species with high dispersal abilities. In this 

last case the two species have distinct bathymetric distribution. 

The geography of speciation 

Jerry Coyne 

CNRS, Avenue de la Terrasse, 91198, Gif-sur-Yvette, France 

One of the most contentious areas of speciation is its biogeography: 

do populations usually need to be geographically isolated to become 

new species, or can those species form in the presence of some 

gene flow. In this talk I review the theory and data bearing on the 

likelihood of parapatric and sympatric speciation in nature, 

concentrating on work that has been done in the last five years. 

Ecological adaptation retraced by molecular changes in 

Odorant Binding Proteins (OBP) in the Drosophila simulans 

complex 

Jean-Luc Da Lage 1 , Delphine Legrand 1 , Takashi Matsuo 2 and Marie- 

Louise Cariou 1 

1 

LEGS, CNRS, avenue de la Terrasse, bâtiment 13, 91198, Gif-sur- 

Yvette, France 

2 

Department of biological sciences, Tokyo Metropolitan University, 

Tokyo, Japan 

Adaptation to an exclusive food resource provides an evolutionary 

advantage and may lead to reproductive isolation. Among the closely 

related species of the simulans complex, Drosophila simulans and D. 

mauritiana are generalist while D. sechellia evolved as a strictly 

specialized species on the toxic ripe fruit of Morinda citrifolia 

(Rubiaceae). The OBP family plays a major role in odor recognition, 

and the Obp57d/e region is clearly involved in the attraction of D. 

sechellia to its host plant. We thus performed a population genetics 

analysis to detect selection signature in the three species. Selection 

was found in D. sechellia and, unexpectedly, in D. mauritiana, but 

not in D. simulans. In addition, tissue-specific expression patterns of 

the two genes Obp57d and Obp57e were investigated and compared 

among species in the presence or absence of the toxic resource. 

Linking fine scale molecular analysis with biological traits, this study 

provides an interesting insight in the comprehension of the 

mechanism of adaptation. 

S2 - Mechanisms of speciation 

- 11 - 

The relative importance of habitat choice and assortative mating 

during rapid ecological divergence 

Fabrice Eroukhmanoff 1 , Sébastien Guéchot 2 , Anders Hargeby 3 and 

Erik I. Svensson 1 

1 

Section for Animal Ecology, Ecology Building, Lund University, SE- 

223 62 Lund, Sweden 

2 

Sup Agro, 2 Place Pierre Viala, 34060 Montpellier, France 

3 

Division of Biology, Linköping University, 581 83 Linköping, 

Sweden 

The question of when and how diverging populations start restraining 

gene flow is critical to the understanding of the origin of species 

(Schluter 2000, Rundle et al. 2000). Assortative mating might 

emerge early on during divergence, but under intense selection, 

migration modifications are favored and will lead to rapid speciation 

(Yukilevich & True 2006). If habitat choice is strong enough between 

populations inhabiting different environments, this will lead to 

allopatric speciation. Reproductive isolation might then emerge 

secondarily through reinforcement (Yukilevich & True 2006). These 

two types of barriers to gene flow can interfere with each other if they 

evolve jointly and it is quite uncertain how well gene flow is limited 

under this scenario. We studied two ecotypes of a freshwater isopod 

(Asellus aquaticus), which started to diverge in parallel in two lakes 

of southern Sweden twenty years ago (Hargeby et al. 2004). We 

report that populations from different ecotypes are sexually isolated 

but not populations from similar ecotypes, regardless of whether they 

originate from the same or different lakes. In addition, we provide 

evidence that habitat isolation between ecotypes has also evolved in 

at least one lake, and now constitutes the main barrier to gene flow 

between ecotypes. In conclusion, the early and simultaneous 

emergence of sexual and habitat isolation has efficiently restrained 

gene flow during this rapid divergence event. However, our study 

reveals that rather than assortative mating (Nosil et al. 2002), habitat 

choice could play an acute role in the early stages of ecological 

speciation. 

Feeding habit causes divergence of skull shape in bats 

Allowen Evin 1,2 1, 2 

and Michel Baylac 

1 

MNHN – CP 50, Origine, Structure et Evolution de la Biodiversité, 

45 rue Buffon, 75005 Paris, France 

2 

Plate-Forme de morphométrie, IFR 101, MNHN – CP 50, 45 rue 

Buffon, 75005 Paris, France 

Within the bat species complex of Myotis myotis, M. blythii and M. 

punicus, studies using 3D geometric morphometrics have shown that 

skull evolution does not match molecular phylogeny (Evin et al. 2007; 

2008). Since some of the observed patterns of skull evolution implied 

different regions of the skull that are potentially linked with feeding, 

we hypothesized that this divergence, between phylogeny and 

phenotypic similitude, could result from functional convergences. 

To test this hypothesis we investigate the patterns of skull variation 

within and between taxa. Most cases reveal that the patterns of 

species evolution closely follow within-species patterns of variability, 

i.e. the least-resistance lines of evolution as defined by Schluter 

(1996). The single exception is that of M. blythii whose patterns of 

differenciation diverged, implying skull parts that are involved in 

mastication process. A closer examination of the within and between 

patterns of skull differenciation using Marroig & Cheverud's test 

(2004) for selection and drift indicates that some components of skull 

shape variation are compatible with selection. These components 

correspond to the separation of M. blythii from the rest of the 

complex and they deal with skull parts specifically involved in 

mastication. Finally, we used a partial-least squares approach to 

analyze the covariation between skull variation and feeding habits. 

Once the effect of diet removed, the residual shapes match closely 

the phylogenetic relationships. 

All investigations therefore are congruent with a divergence of M. 

blythii by diversifying selective pressures related to feeding habits.


Closing the ring: biogeography of the salamander ring species 

Ensatina eschscholtzii 

Shawn R. Kuchta 1 , Duncan Parks 2 , Rachel Lockridge Mueller 3 and 

David B. Wake 4 

1 

Lund University, Department of Animal Ecology, Ecology Building, 

Sölvegatan 37, SE-223 62, Lund, Sweden 

2 

Mt. Angel Seminary, St. Benedict, OR 97373, USA 

3 

Department of Biology, Colorado State University, Fort Collins, CO 

80523-1878, USA 

4 

Museum of Vertebrate Zoology, Department of Integrative Biology, 

University of California Berkeley, Berkeley, CA 94720-3160, USA 

Ring species exhibit a circular arrangement of populations around a 

central barrier, with reproductively isolated parts overlapping at one 

point in the ring, yet morphological and genetic intergradation 

elsewhere. They evolve when two or more lineages descend from a 

common ancestor and become reproductively isolated while 

maintaining their connectivity through a chain of interbreeding 

populations. The salamander Ensatina eschscholtzii is a classic 

example of a ring species. In the original biogeographical scenario, 

the point of ring closure was situation in southern California, USA. 

Here we develop an alternative biogeographic scenario that is 

informed by the geomorphological development of California, and 

situates the point of ring closure in central coastal California. To 

distinguish between these two biogeographical alternatives, mtDNA 

sequence data was collected from 385 individuals from 224 

populations, and a Bayesian phylogeny was inferred. The two 

biogeographical scenarios were tested against our Bayesian 

topology, including the associated Bayesian 95% credible set of 

trees. Our Bayesian topology contradicts the new biogeographic 

hypothesis. 

Evolution of the chromosomal organization in the Sophophora 

subgenus of Drosophilidae : the nucleolus organizer region 

Laurence Monti 1,2 , Nicole Chaminade 2 , Jean-Luc Da Lage 2 , Marie-Louise 

Cariou 2 , Françoise Lemeunier 2 and Sylvie Aulard 2 

1 present address: Laboratoire Stress, Défenses et Reproduction des 

Plantes, Université de Reims, Champagne, Ardenne. Campus Moulin de 

la Housse Bat 8 – BP 1039 51687 Reims cedex 2, France 

2 Laboratoire Evolution, Génomes et Spéciation, CNRS, Avenue de la 

Terrasse, 91198 Gif-sur-Yvette cedex, France 

In Drosophila, the heterochromatic sequences are found into specific 

chromosomal regions and submitted to rapid evolution. Due to its 

heterochromatic localisation, and its highly conserved structure in Insects, 

the nucleolus organizer region (NOR) is a useful tool to study the 

rearrangements of the chromosomal architecture. 

Using fluorescent in situ hybridization, we have analysed 54 species 

belonging to different groups of the Sophophora subgenus. 

The presence of one NOR on each sexual chromosome (X and Y), already 

known in Drosophila melanogaster, has been detected for the 16 other 

species studied in the melanogaster group. However, NORs on the Y 

chromosome of D. simulans and D. sechellia have lost the 28s rDNA 

genes. 

In the 26 species of the montium group, the NORs are also localised on the 

sex chromosomes but their number can vary between 1 and 3, which 

implies the loss or the gain of a NOR in some species. In conclusion, the 

presence of NORs on sexual chromosomes is an ancestral feature for 

these two groups. 

On the contrary, the 12 species of the ananassae group show quite 

different pattern, most of them showing only one NOR, on chromosome 4. 

This could be related to data on chromosomal evolution that point 

out the hypothesis of a translocation between the X or the Y 

chromosome (depending on authors) and chromosome 4 in D. 

ananassae. Such an event could explain the presence of the NOR 

on this autosome without rejecting that NORs on sexual 

chromosomes are ancestral for the Sophophora subgenus. 

- 12 - 

The Crassostrea oyster species in China 

Haiyan Wang 1,2 , Guofan Zhang 1 , Xiao Liu 1 and Ximing Guo 2 

1Institute 

of Oceanology, Chinese Academy of Sciences, Qingdao, 

Shandong, PRC 

2 

Haskin Shellfish Research Laboratory, Institute of Marine and 

Coastal Sciences, Rutgers University, Port Norris, New Jersey, USA 

China is the home to many oyster species. Crassostrea species are 

most common and commercially important in China. There is 

considerable confusion about the classification of Crassostrea 

species in China. In the past five years, we collected and analyzed 

thousands of oysters at typical sites along the coast of China. By 

comparing the morphology and sequences of 16S, COI and 28S 

gene, we found seven Crassostrea species occurring along the coast 

of China. One of our studies is on the taxonomic status of the red 

and white forms of C. rivularis (Gould, 1861), which shows that the 

red oyster is the same species as C. ariakensis; the white oyster is 

the same species as a newly described species, C. hongkongensis 

(Wang et al., 2004). Another study is on the classification of small 

cupped oysters along the coast of China, which includes three 

species, C. gigas, C. angulata and C. sikamea. Recently we found 

that C. nippona and C. iredalei were also presented in China. Our 

studies clarified the classification of Crassostrea species in China, 

providing basic information for oyster classification, aquaculture and 

the protection of oyster resources in China. 

Guo X., S. Ford and F. Zhang. 1999. J. Shellfish Res., 18:19-31. 

Wang H. and X Guo, 2008. J. Shellfish Res., 27(3):481-487. 

Wang H., X. Guo, G. Zhang and F. Zhang. 2004. Aquaculture (242): 

137-155. 

Wang H., G. Zhang, X. Liu and X. Guo. 2008. J. Shellfish Res., 

27(3):495-503. 

The unexpected ecology of allopatric speciation 

John Wiens 

New York, 11794-5245, Stony Brook, U.S.A. 

Allopatric speciation is widely considered to be the most common of 

the thre geographic modes. Yet, there has been suprisingly little 

focus on the evolutionary and ecological processes that cause 

species to become allopatric. In this talk, I will discuss recent 

research from my lab looking at this neglected aspect of speciation. 

Consideration of how a species becomes split into allopatric populat! 

ions leads to an unexpected view: that the key to geographic 

isolation is not necessarily the adaptive divergence of different 

populations, but rather the failure of an ancestral species to adapt to 

environmental change in part of its geographic range. Thus, the 

ecological similarity of species over time (niche conservatism) may 

often be essential to their geographic isolation; if all species could 

quickly adapt to any environmental conditions, there would be few 

geographic barriers to gene flow and little allopatric speciation. I will 

describe how we have tested and supported this model of allopatric 

speciation through niche conservatism in North American 

salamanders. However, the situation is more complicated in tropical 

salamanders, where closely related species tend to occur in 

dissimilar environments. Explanations for these divergent patterns 

will be addressed, along with their implications for the causes of the 

latitudinal diversity gradient. Research on the evol! utionary ecology 

of allopatry will be placed within the larger conceptual framework of 

speciation research. Finally, the relationships between speciation, 

niche conservatism, and other topics in ecology and evolution will be 

discussed (e.g., responses to climate change, spread of invasive 

species, community assembly, historical biogeography).


Role of expression differentiation in adaptation and speciation: 

comparative analysis of Drosophila simulans and 

Drosophila sechellia 

François Wurmser 1 , David Ogereau 1 , Tristan Mary-Huard 2 , 

Dominique Joly 1 and Catherine Montchamp-Moreau 1 

1Laboratory Evolution Genomes and Speciation, CNRS UPR9034, 

Av de la terrasse 91198 Gif sur Yvette CEDEX, and University Paris 

Sud, France ; 2 UMR AgroParisTech/INRA 518, 16 rue Claude 

Bernard 75231 Paris cedex 05, France 

Variation in gene expression can cause important phenotypic 

variation between and within species, and is likely to play a central 

role in processes driving adaptation and speciation. Our goal is to 

detect genes involved in these processes by comparing populations, 

species and interspecific hybrids. 

We have conducted a comparative study of the transcriptome of 

different Drosophila populations and species: 3 subsaharian African 

populations and one European population of the cosmopolitan 

generalist Drosophila simulans, and a population of the insular 

endemic specialist D. sechellia. We also examined 

simulans/sechellia sterile hybrid males to detect misregulation 

following Dobzhansky-Muller's model of genetic incompatibility. We 

used microarrays carrying complete cDNAs of about 5000 genes of 

D. melanogaster to compare whole-body RNAs of 7-days-old males 

between each condition. 

The statistical analysis revealed structuration between African 

populations of the ancestral area and the European population, 

which is consistent with published data on microsatellite variation. 

The comparisons between D. simulans and D. sechellia have 

revealed 304 consistently differentially expressed genes, while only a 

few between the hybrids and their parents. Within the former gene 

list, it is worth noting the overrepresentation of genes involved in 

hormone functionalisation and thus hormone regulation, and/or in 

detoxification processes. Genes involved in Hormone catabolism 

were also overrepresented. These pleiotropic genes being involved 

in reproduction will be worth further exploration for their potential role 

in the simulans/sechellia speciation. 

- 13 -


S3 - Animal genitalia in evolution-in honour of William G.Eberhard 

Dragonflies (Aeshnidae, Corduliidae, Gomphidae and 

Libellulidae), of the Llanos Orientales, Colombia: species, inter 

and intraspecific variation of the genitalia 

Catalina Amaya-Perilla 1 and Gonzalo Fajardo 2 

1 Universidad Jorge Tadeo Lozano, Facultad de Ciencias Naturales, 

Programa de Biología Ambiental, Bogotá, Colombia 

2 Universidad Jorge Tadeo Lozano, Facultad de Ciencias Naturales, 

Programa de Biología Marina, Bogotá, Colombia 

The knowledge about the genitalia of Anisoptera (Insecta:Odonata) 

is very little since these individuals are very difficult to capture and 

mark. In Colombia there is no information about this subject, 

therefore, the purpose of this study is to add information about the 

variation of the genitalia for the suborder Anisoptera in the Llanos 

Orientales Meta department of Colombia. We sampled the 

specimens of the suborder Anisoptera (Odonata) on 11 sampling 

areas (farms): 8 sampling areas of gallery forest and savannah, 2 

sampling areas of Andean foothills and 1 sampling area of lower 

mountain forest. We used entomological nets in two different 

seasons from 2003 to 2007 and one season of 2008. All the 

collected material was preserved through immersion in acetone for 

12 hours, and was later determined to genera. We explored the 

variation between the genitalia of the specimens collected with 

Elliptic Fourier analysis and semilandmarks. We found in total 575 

specimens and 24 genera distributed in 4 families: Aeshnidae, 

Corduliidae, Gomphidae and Libellulidae. The genitalia studied for 

the males showed inter and intra specific variation, showing a 

significant variation between the genera studied. These differences 

could be used as tools for the evolutionary relationships among the 

genitalia of the group. 

The secret sexual life of Pea Crabs – Reproductive Morphology 

of European Pinnotheridae (Crustacea, Decapoda, Brachyura) 

Carola Becker 1 , Dirk Brandis 2 , Volker Storch 3 and Michael Tuerkay 1 

1 

Research Institute Senckenberg, Senckenberganlage 25, 60325 

Frankfurt/Main, Germany 

2 

Zoological Museum, University of Kiel, Hegewischstr. 3, 24105 Kiel, 

Germany 

3 

Zoological Institute, University of Heidelberg, Im Neuenheimer Feld 

230, 69120 Heidelberg, Germany 

Pea Crabs are not larger than a thumbnail and live commensally 

inside a variety of invertebrates. While juveniles of both sexes still 

look very similar - being agile swimmers, partly free-living and just 

occasionally found inside the host - a metamorphosis takes place in 

the female after mating which results in a conspicuous sexual 

dimorphism. From now on, the female focuses her life on feeding 

and breeding hidden inside the host, while the male remains freeliving, 

only occasionally found together with the female in the host. 

Pea Crabs have a quite complicated life cycle and due to their cryptic 

way of life it is hard to observe their sexual behaviour. Instead of that, 

I examined the functional reproductive morphology in order to 

understand basic adaptions and possible processes in copulation, 

sperm storage and fertilisation. I used histological methods, Electron 

Microscopy (SEM & TEM) and Confocal Laser Scanning Microscopy 

(CLSM). 

The pinnotherids’ female spermatheca shows a high degree of 

complexity: While a holocrine glandular tissue is located in the 

ventral “insemination area” where oviduct and vagina are connected 

to the spermatheca, an apocrine glandular epithelium lines out the 

dorsal “sperm-storage area” of the spermatheca, that is described 

here for the first time for true crabs. 

The significance of the morphology and function of the male and 

female reproductive systems for understanding the evolutionary 

pathways of the sexual biology is discussed in the context of other 

true crab families. 

- 14 - 

Sexual coevolution and the evolution of genital traits in 

Lepidoptera 

Carlos Cordero 1 , Víctor Sánchez 2 and Blanca Hernández 3 

1 Departamento de Ecología Evolutiva, Instituto de Ecología, 

Universidad Nacional Autónoma de México (UNAM), Mexico City. 

2 Posgrado en Ciencias Biológicas, Instituto de Ecología, UNAM. 

3 Departamento de Biología Evolutiva, Facultad de Ciencias, UNAM. 

The convergence and divergence of male and female interests 

during sexual reproduction generates reciprocal selection pressures 

between the sexes. The continuous development and elaboration of 

male and female traits resulting from these selection pressures is 

known as sexual coevolution. We present sexual coevolution 

hypotheses for the evolution of two genital traits widely distributed in 

the Lepidoptera. (1) Signa are sclerotized structures located on the 

inner wall of the corpus bursa of females, whose main function is 

tearing open spermatophores. We propose that polyandry would 

favor the evolution of thick spermatophore envelopes that take 

longer to be broken, thus decreasing female remating rate, and that 

this would select for signa that allow females to recover control of 

their remating rate. We tested this hypothesis by a comparative 

study and found: (a) the predicted association between mating 

pattern and the presence of signa in a sample of 37 taxa, and (b) the 

predicted association between thickness of the spermatophore 

envelopes and the presence and characteristics of signa in a sample 

of Heliconius and Eueides (Nymphalidae) species with different 

mating system. (2) The cornuti are sclerotized structures of the 

endophallus, which in some species break off and remain within the 

corpus bursae of females. We describe the diversity of cornuti, 

propose hypotheses on their function, and discuss ways of testing 

these hypotheses. Our analyses suggest that sexual coevolution has 

played a major role in the evolution of genitalia in the Lepidoptera. 

Sexually antagonistic coevolution and rapid divergent evolution 

of animal genitalia 

William G. Eberhard 

Smithsonian Tropical Research Institute, and Universidad de Costa 

Rica 

A recent summary by Hosken and Stockley concluded that current 

evidence strongly favors sexual selection as the primary force driving 

rapid divergent evolution of genitalia, but that it is not clear whether 

sexually antagonistic coevolution (SAC) or cryptic female choice 

(CFC) is responsible. I will explain in this talk why I believe the 

current balance is more strongly tilted against SAC than they imply. 

SAC has the virtue of generating relatively clear predictions. The 

evidence against it includes large surveys of insects and spiders 

(involving many thousands of species) that found: a lack of the 

predicted correlation between rapid divergent evolution of male 

genitalia and the behavioral contexts in which male-female interests 

are more likely to be in conflict in different species; a general lack of 

the predicted female defensive coevolution in groups with speciesspecific 

male genitalia; only weak genital diversification in groups 

with especially intense male-female conflicts; and a strong trend 

toward allometric scaling patterns that are opposite of those 

predicted by SAC. This evidence does not rule out SAC for particular 

cases at particular moments in evolution; but the combination of very 

large sample sizes and the lack of traces of trends predicted by SAC 

implies that the effect of SAC on rapid divergent genital evolution 

must be small. If SAC has acted, it has apparently been brief, weak, 

or inconsistent; most of the modern diversity of genitalia is 

apparently due to some other factor.


Investigations on giant sperm in Drosophila bifurca 

Magali Evanno 1,2 , Christophe Bressac 1 and Dominique Joly 2 

1 Institut de Recherche sur la Biologie de l’Insecte, UMR CNRS 6035, 

Faculté des Sciences, 37200 Tours, France 

2 Laboratoire Evolution, Génomes et Spéciation, CNRS UPR 9034, 

91198 Gif sur Yvette Cedex, France and Université Paris-Sud 11, 

91405 Orsay Cedex, France 

The theory of anisogamy predicts that, for the same investment in 

energy, males will produce smaller and more spermatozoa, while 

females will produce fewer but larger oocytes with many reserves. 

Accordingly, some organisms appear to be evolutionary paradoxes, 

as Drosophila bifurca which produces spermatozoa measuring 6 cm, 

representing 20 times the body length. The reproductive tract of 

those males contains a coiled structure between the testes and the 

seminal vesicles, namely the roller. At the testicular proximal end, 

brother spermatozoa of the same cysts are coiled up together before 

transiting through the roller, then individualizing and rolled as pellets 

in the seminal vesicles, i.e. the male storage organs. In D. bifurca, 

sperm present a unique shape with extraordinarily long flagellum 

rolled up around large quantities of cellular material. In order to 

understand the physiological mechanisms that allow such giant 

sperm to coil and roll them individually, we observed and analyzed 

the changes of their shape in different saline buffers and under 

different osmotic pressure conditions. We also studied their motility 

and sprawling parameters according to time in a reference solution 

(Hyes’ringer). Finally we focused on the identification of the cellular 

material using fluorescent dyes, and its apparition or disappearance 

kinetics respectively in the male and female reproductive tracts. 

Thanks to our results and electron microscopic study, we will discuss 

the origin (testis or roller) of the cellular material and the evolutionary 

significance of such male donation in the context of sexual selection 

hypotheses (i.e. nuptial gift or mating plug). 

Sexual selection and asymmetric genitalia 

Bernhard A. Huber 

Alexander Koenig Research Museum of Zoology, Adenauerallee 160, 

53113 Bonn, Germany 

There is a wide consensus that sexual selection has been 

responsible for the immense diversity of genital morphology, but one 

particular aspect has largely been neglected: adaptive genital 

asymmetries. Such asymmetries are extremely common in insects, 

but almost entirely absent in spiders. In the latter, only five 

independent origins are known; in contrast, genital asymmetry is in 

the groundplan in some insect orders and has evolved multiple times 

convergently in others. 

Insect genital asymmetry is overwhelmingly limited to the male. I thus 

propose that sexual selection is involved, and I argue that changes in 

mating position are the centerpiece in the insect route to asymmetry. 

Available evidence strongly suggests that the plesiomorphic 

neopteran mating position is a female-above position. Changes to 

male-dominated positions have occurred frequently, and some of the 

resulting positions require abdominal twisting, flexing, and 

asymmetric contact between male and female genitalia. Insects with 

their median unpaired sperm transfer organ may adopt a one-sided 

asymmetric position and still transfer the whole amount of sperm. 

Spiders with their paired sperm transfer organs can only mate in 

symmetrical or alternating two-sided positions without foregoing 

transfer of half of their sperm. 

Based on this premise, I propose two major hypotheses for the 

evolution of insect genital asymmetry. One explains morphological 

asymmetry as a mechanical compensation to evolutionary and 

behavioural changes of mating position. Not the asymmetry per se is 

advantageous, but the newly adopted mating position. The second 

hypothesis predicts a split of functions between right and left sides. 

In this hypothesis, asymmetry per se is advantageous. 

- 15 - 

Differences in structure and function of genitalia between the 

subfamilies of longhorn beetles (Coleoptera: Cerambycidae) 

Lasse Hubweber and Michael Schmitt 

Zoologisches Forschungsmuseum Alexander Koenig, Adenauerallee 

160, 53113 Bonn, Germany 

Genitalia of beetles are usually described only for taxonomic reasons 

without any idea on the functionality of the various structures. 

Exceptions to this can be found only in very detailed studies on 

single species. This study relates to the genital structures of the 

various longhorn beetle subfamilies, ideas concerning the function of 

the structures and the phylogeny of Cerambycidae. 

Specimens of almost 150 species of cerambycids have been 

collected, dissected and studied with SEM. In addition to that, 

copulating pairs of about 20 species have been frozen with ice spray 

and dissected or studied histologically. 

Especially male genitalia show various characters typical for higher 

taxonomic categories. For example the parameres show a large 

variability in Cerambycinae, while the parameres in most species of 

Lamiinae look relatively uniform and different to Cerambycinae. 

Internal sac structures look very different in the various subfamilies. 

Small spines pointing backwards are the most common armature of 

the internal sac. There are less differences between the groups in 

female genitalia. At least the ovipositor is built in variable ways in the 

different subfamilies. In most species, the connection between the 

mates during copulation is given through the long internal sac and 

ovipositor only. Median lobe and parameres are in contact with the 

female only at the beginning of copulation. 

The comparison of genital structures of various subfamilies provides 

new phylogenetic characters. The functional reason for the length of 

the internal sac is its function as element of connection and fixation 

between mates. 

Male-female coevolution of reproductive organs in Drosophila 

Dominique Joly 

CNRS-UPR 9034, Laboratoire Evolution, Génomes et Spéciation, 

91198 Gif-sur-Yvette Cedex, and Université Paris-Sud 11, 91405 

Orsay Cedex, France, 

The male genitalia of Drosophila species exhibit great morphological 

diversity even between closely related species. This morphological 

diversity is most likely due to sexual selection processes. In most 

species, the external (epandrium) and internal (hypandrium) 

chitinized structures are commonly used as diagnostic criteria for 

species identification. Contrasting with these structures, the soft 

internal reproductive organs are generally poorly considered mainly 

due to increased morphological bias that artificially overestimates 

within species variability. However, alongside with the external 

structures, the internal ones have to be considered in postcopulatory 

sexual selection as they may drive differential interactions between 

sperm and oocytes. This is even more crucial in internally fertilizing 

organisms where females have the ability to store sperm, because it 

is then possible for the ejaculates of several males to compete 

during the course of the female's reproductive life. Such competition 

is likely a major driving force of sexual selection. Sperm length is one 

of the traits of the internal male apparatus that tightly correlate with 

the internal female morphology. Variation in sperm length is 

enormous within taxonomical groups and leads recurrently to sperm 

gigantism, particularly in non-vertebrate species. However, the 

anatomical, cytological, and physiological requirements of both 

males and females associated with giant sperm are poorly 

understood. In Drosophila, we analysed the extant of the malefemale 

coevolution of internal reproductive organs and identified 

specific features of the male tract that are associated with giant 

sperm.


Male terminalia variation in the rainforest dwelling Drosophila 

teissieri contrasts with the sperm pattern and species stability 

Dominique Joly 1 , Marie-Louise Cariou 1 , Tendai Mhlanga- 

Mutangadura 2,3 and Daniel Lachaise 1,# 

1 CNRS UPR 9034, Laboratoire Evolution, Génomes et Spéciation, 

91198 Gif-sur-Yvette Cedex, France, and Université Paris-Sud 11, 

91405 Orsay Cedex, France 

2 Department of Biological Sciences, University of Zimbabwe, P.O. 

Box MP 167, Mount Pleasant, Harare, Zimbabwe ; 3 Present adress: 

Department of Veterinary Pathobiology, University of Missouri, 

Columbia, MO 65211 

# Our late colleague and friend 

It is commonly recognized that speciation does not necessarily result 

from extensive variation between populations, and what the 

speciation process per se consists of still remained an outstanding 

question. We advocate here that the variation of male terminalia 

does not necessarily result in noticeable reproductive isolation. Here 

we report on the strictly Afrotropical forest-dwelling continental 

species Drosophila teissieri whether there is invariance or variance 

of traits central to sexual selection processes (i.e. male terminalia 

and sperm length) compared to traits which are generally assumed 

to vary more neutrally (i .e. allozymes). Three geographic categories 

can be recognized: the central populations of the Guineo-Congolese 

forest block (Cameroon-Congo), the outlying large populations of 

West Africa to the west of the Dahomey gap (Guinea and Ivory 

Coast), and isolated marginal populations scattered in East Africa 

(e.g. Silinda in Zimbabwe). Although we concur with the argument 

that the species integrity requires some degree of stability, our data 

suggest that its components do not obey the variance/invariance 

alternative consistently. Male terminalia and allozymes show 

extensive variation while sperm length distribution is strikingly similar 

between the categories. It is therefore inferred that this trait might be 

one of the major targets of stabilizing selection. The relative 

invariance of sperm distribution feature between populations is only 

made possible due to considerable and consistent variance within. 

Finally, it is suggested that the striking fit between the extent of 

sperm heteromorphism (within male) and sperm polymorphism 

(between males) is instrumental in maintaining the species integrity. 

Evolutionary causes and consequences of mating plugs in 

spiders 

Stefan H. Nessler 1 , Jutta M. Schneider 1 and Gabriele Uhl 2 

1 

University of Hamburg, Biocenter, Department of Ethology, Bonn, 

Germany 

2 

University of Bonn, Institute of Zoology, Department of 

Neuroethology, Bonn, Germany 

In animal mating systems there is a high selective advantage for any 

mechanism that helps to avoid sperm competition. Relatively obvious 

means by which males can secure their fertilization success is to 

prevent or impede the female from re-mating with rival males by 

plugging her copulatory opening. Mating plugs are a common 

phenomenon for spider taxonomists: plugs made of secretion are 

often found on the female copulatory opening and in some taxa 

males are known to mutilate their genitalia in order to leave the 

broken off parts inside the female genitalia. In this talk we will review 

the information available to date on the effectiveness of the two 

types of plugs. As to secretory plugs there is preliminary evidence 

that in some species the mating plug is only effective when male and 

female substances interact. These observations strongly suggest 

that cryptic female mate choice can shape the effectiveness of male 

secretions. As to male genital mutilation there is a recent increase in 

experimental and comparative data. Since male genital mutilation 

predominantly occurs in sexually cannibalistic species we will ask if 

genital mutilation in males is the cause or the consequence of sexual 

cannibalism. 

- 16 - 

Insect genitalia, sexual selection and sperm ageing 

Klaus Reinhardt 

Western Bank, S10 2TN, Sheffield, UK 

Bill Eberhard has both united and revolutionised taxonomy and 

evolutionary biology by proposing that animal genitalia have evolved 

by sexual selection context, particularly so through male genitalic 

courtship enabling cryptic female choice. Using several insect 

species, I will here test some of Bill Eberhard's explicit predictions 

about male genitalia allometry and about corresponding female 

genitalia diversification. Cryptic female choice also incorporates the 

specific case of sperm choice by females. I will explore the possibility 

that genitalia have functions other than courtship and will propose 

that intra-ejaculate variation in sperm quality may also be related to 

genitalia complexity. 

Functional Morphology of Copulatory Organs in Coleoptera 

Phytophaga (Insecta) 

Michael Schmitt and Susanne Düngelhoef 

Zoologisches Forschungsmuseum Alexander Koenig, Adenauerallee 

160, 53113 Bonn, Germany 

Originally, the male copulatory organ (aedeagus) of the 

Curculionoidea and the Chrysomeloidea is composed of a median 

lobe and a tegmen with basal struts and distal parameres. Within the 

taxon Phytophaga (= Pseudotetramera), the parameres have been 

reduced several times. Comparison of different types of parameres 

and median lobes and aedeagi lacking parameres and investigation 

of dissected pairs in copula revealed that (1) parameres do not 

provide mechanical coupling, (2) mechanical footing is provided by 

the endophallus, (3) the flagellum serves as a means for sperm 

transfer, and (4) the male possibly perceives tactile stimuli from the 

bursa copulatrix (inside the female) during copulation by means of 

sensilla in the membrane of the endophallus. 

In several evolutionary lines the sclerotization of the aedeagus has 

been reduced. Mechanical and behavioural interaction between male 

and female copulatory organs have been studied morphologically 

and by observation of live copulating pairs of beetles.


Sexual selection and genital evolution in mammals 

Paula Stockley 

Mammalian Behaviour & Evolution Group, Faculty of Veterinary 

Science, University of Liverpool, Leahurst Campus, CH64 7TE, UK. 

Animal genitalia evolve rapidly and divergently, with the result that 

closely related species often have conspicuously different genital 

anatomy. Following the pioneering work of William G. Eberhard, 

there is now growing evidence that sexual selection may be largely 

responsible for this diversity. In the current presentation I will review 

evidence that the genital anatomy of male mammals is subject to 

sexual selection, with emphasis on understanding diversity in the 

relative size of particularly variable traits such as penile spines and 

the baculum or os penis. After providing a broad overview of 

comparative trends in genital diversity among primates and other 

taxa, I will focus on a case study of post-copulatory sexual selection 

in wild house mice (Mus musculus domesticus) that provides new 

evidence for an association between male genital anatomy and 

reproductive success in a mammal. Overall, the evidence presented 

indicates that sexual selection is likely to be a significant factor 

explaining certain aspects of genital evolution in mammals. However, 

as for other taxa, distinguishing between alternative models of 

genital evolution by sexual selection may prove a significant 

challenge. 

- 17 - 

Morphological and phylogenetic implications of female genitalia 

in micronetine spiders (Araneae, Linyphiidae) 

Lihong Tu 1,2 and Gustavo Hormiga 2 

1 

College of Life Sciences, Capital Normal University, Beijing100037, 

P. R. China 

2 

Department of Biological Sciences, The George Washington 

University, Washington, DC 20052, USA 

The genital morphology of spiders provides a rich source of 

phylogenetic data and can help our understanding of many issues in 

evolutionary biology and phylogenic reconstruction. Male palpal 

morphology has played a critical role in phylogenetic studies, often 

dominating character data matrices. In contrast, very limited 

information is available on the morphological details of the female 

genitalia (the epigynum). The present paper focuses on the female 

genitalia of micronetine spiders, a group of linyphiids with a very 

complex epigyna. We have studied the external and internal 

morphology of the epigynum of representatives of more than 30 

micronetine genera. A growing body of evidence suggests that the 

so called ducts used to transmit sperm (copulatory and fertilization 

ducts) are furrows derived from integument folds. We try to use this 

model to explain variations in different linyphiid groups. A set of 

comparative morphological characters is presented here. We discuss 

the phylogenetic implications of these new findings and whether 

there is coevolution between male and female genitalia.


S4 - SYSTEMA NATURAE 250 - The Linnean Ark-250 years of animal names 

Daniel Rolander’s Diarium Surinamicum and its Insects 

James Dobreff 

Lund University, Research Fellow Manager, Daniel Rolander Project, 

Götgatan 81, 5tr, SE- 116 62 Stockholm, Sweden 

This talk will explain why Linnaeus apostle Daniel Rolander’s 

Diarium Surinamicum has remained, until very recently, unknown to 

nearly all of the scientific community and why he has suffered a 

horrendous reputation in his Swedish homeland. Finally, Rolander’s 

treatment of insects will be presented along with several key 

passages from the historical critical edition of the Latin text which the 

speaker is currently editing. 

Rolander (c.1723-1793) had been one of Linnaeus’s most promising 

students. He resided with Linnaeus for four years, while tutoring Carl 

Jr. In 1754 he was selected to document the Dutch colony of 

Suriname according to Linnaean methods. After seven months in 

Suriname and arduous journeys to and from Suriname, he returned 

to Sweden in 1756. His health had suffered from two serious fevers. 

He refused to show his specimens or journal notes to Linnaeus, 

which eventually caused their complete estrangement. He completed 

the last draft of the Diarium in Denmark in 1765. Besides a few short 

passages, it was never published. 

The speaker is currently preparing a critical edition of the Latin text 

and was the coordinating translator of the forthcoming English 

translation of the Diarium. 

Linnaeus - A passion for order 

David Quammen 

Bozeman, Montana, USA 

Carl Linnaeus stated in his Philosophia Botanica (1751) “If you do 

not know the names of things, the knowledge of them is lost too". 

Seven years later he extended to animals the Latin binomial system 

already established by him for plants. Although Linnaeus built on the 

earlier work of Aristotle, Fuchs, Ray and Pitton de Tournefort, he is 

justifiably regarded as the "father of taxonomy". Despite his 

classifying Homo sapiens with monkeys and apes, he was no 

evolutionist but a committed creationist. However, his religious 

beliefs did not prevent him from striving to embrace the entire 

diversity of nature for its own sake as well as for its practical uses. 

Linnaeus's colossal achievements constitute an act of human 

heroism, and a 250-year long legacy that we have a moral duty both 

to preserve and to build upon. 

- 18 - 

The naming of threatened animal and plant species – a matter of 

life and death 

Gordon McGregor Reid 

North of England Zoological Society, Zoological Gardens Chester, 

Cheshire, UK 

Following the Systema Naturae of Carolus Linnaeus (1707-1778) the 

naming of species and other taxa remains the fundamental activity of 

taxonomists. There are now >1.8 million scientifically named, living 

organisms: 13 new 

amphibian species were recognized each month. Counteracting 

such trends is an increased number added to the IUCN Red List of 

species threatened with extinction, including 43% of freshwater 

fishes, 32% of amphibians, 23% of mammals and 12% of birds in 

2007. Costs to implement the IUCN Amphibian Conservation Action 

Plan are estimated at >€257 million. The price of saving individual 

species can be high, up to €32 million for Californian condor; and an 

individual elephant in a zoo breeding programme may cost >€45, 

000 per annum to maintain. Conservation strategies organized in 

situ or ex situ often depend on the biological validity and 

nomenclatural stability of taxonomic names and the scientific cum 

philosophical integrity of species concepts; and are subject to costly 

legal challenges by land developers. The contemporary recognition 

(or de-recognition) of new species or subspecies among longestablished 

animal groups such as Amazon parrots, elephants, 

jaguars and orang-utans has substantial wildlife management 

implications. Today, justification and conservation prioritisation of 

names can be a matter of life or death in terms of species survival. 

Fossils and Linnaean classification 

Hans-Dieter Sues 

National Museum of Natural History, Washington, DC, USA 

As the remains of long extinct organisms, fossils are inherently 

incomplete. Even the most exquisitely preserved specimens contain 

but a fraction of the total original biological information about the 

organisms documented by them. Consequently, classifying fossils 

has proven to be challenging ever since the introduction of the 

Linnean system of classification. Fossils represent critical evidence 

for understanding of the evolution and diversity of Life, and thus must 

be included into classifications based on extant organisms. This was 

already recognized by paleontologists working in a pre-Darwinian 

framework. Over the years researchers have proposed various 

paleontological species concepts (e.g., chronospecies), but none of 

these has proven useful and been widely adopted. Species of extinct 

organisms can be based on the presence of unique morphological 

character-states or unique combinations of features, as is commonly 

done for extant species. However, it is important to keep the nature 

of such extinct taxa in mind when using them to address biological 

questions such as detailed changes in biodiversity over geological 

time.


Linnaean Classifications: from Ark to Battlestar 

Quentin D. Wheeler 

International Institute for Species Exploration, Arizona State 

University, Tempe, Arizona, USA 

With the recognition of the biodiversity crisis, the missions of 

taxonomists to explore, describe and classify species have taken on 

unprecedented importance and urgency. Although species 

discovery rates are higher than the historic averages, they are 

insufficient to meet the needs of the world for reliable taxonomic 

information or to comprehensively document species diversity for 

posterity. A fusion of the traditional goals of taxonomy with grid 

computing and digital technologies is defining a fundamentally new 

approach to species exploration: cybertaxonomy. Collaborations 

with historians, philosophers and sociologists can help identify and 

avoid repeating the mistakes of the past that have undermined 

support for taxonomy in recent decades. Essential to success are 

recognizing the unique needs of taxonomy as an independent 

science and opening access to the resources needed by taxonomists 

to do their work. The “Taxon Knowledge-Community” model 

provides for both individual scholarship and highly coordinated teams 

making rapid progress in taxonomy. All the necessary theory, 

methods, and technologies for transforming taxonomy into an 

effective international science are in place or in sight. We need now 

the vision, courage and individual and institutional leadership to 

realize taxonomy’s potential as a modern leading collaborative 

science. Rather than a mentality of gathering species onto an ark of 

knowledge when and however we can, we must now adopt an 

aggressive strategy to attack and conquer taxonomic ignorance on a 

planetary scale. 

- 19 - 

The Linnaean Ark 

Edward O. Wilson 

Harvard University, Cambridge, MA, USA 

The outstanding fact of 2008 concerning the Great Linnaean 

Enterprise is that now, 250 years after the publication of the tenth 

edition of the Systema Naturae, biologists probably have discovered 

and classified fewer than 10 percent of the species of organisms on 

Earth. That estimate manifestly applies to fungi, as well as the most 

hyperdiverse of small invertebrates and-dramatically-the bacteria, 

archeans, and viruses. About 1.8 million species of organisms have 

been diagnosed and given formal names. But the real number? It 

should be thought an astonishing failure of science that biologists do 

not know within the nearest order of magnitude the total number of 

species composing the biosphere. 

This quarter millennial, 2008, will be remembered as occurring 

during a turning point in the history of systematics. Should society 

wish to make such a project big science, it is now possible to 

complete the discovery of most of the remaining, unknown 90 

percent of species within a generation-say 25 years, or one-tenth the 

time it took to reach the point where we are now. A powerful 

motivation exists in the publicly recognized dangerous change 

occurring in the global environment. The technologies are newly 

available to accomplish the job. They include genomics, highresolution 

digital photography-and the internet-based Encyclopedia 

of Life, designed to make everything known about every species to 

anyone, anywhere, any time, on command. 

The ICZN has a major challenge and responsibility to guide the 

classification system within the swiftly growing database, and to 

provide juridical decisions for nomenclatural conflicts arising within it.


S5 - SYSTEMA NATURAE 250 - Current issues in animal nomenclature 

1758: Binomen; 2008 Open, Enhanced Descriptions 

Donat Agosti 1 , Terry Catapano 2 , Norman F. Johnson 3 , Richard Pyle 4 

and Zhi-Qiang Zhang 

1 Berne, Switzerland ; 2 Columbia University, New York, USA ; 3 Ohio 

State University, Columbus, Ohio, USA ; 4 Bishop Museum,Honolulu, 

Hawaii, USA ; 5 Landcare, Auckland, Nez Zealand 

Linnaeus's goal for Systema Naturae was to have a synthesis and 

catalogue of all species known at that moment together in one place. 

It remained from its invention until today the ultimate reference. 

Since all the knowledge of the species included was printed in a 

single publication, it could be shipped easily, and used as the basis 

of all the later work - and it had such a tremendous effect that we 

cannot master its legacy: we do not have a Systema Naturae edito 

2008, not even a list of all the described taxa. 

Now, 250 years later, with the Internet we have the medium to 

provide everybody access at once to all the taxonomic knowledge 

about species (and of course anything linked to it), we can produce 

e-descriptions shared at once by everybody, and assure that all edescriptions 

are linked to a registry of at least all the recently 

published names. 

In this contribution we describe how such an e-description could look, 

using examples recently published in Zootaxa and PLoS-ONE. 

Issues are discussed regarding technical aspects (TaxonX XML to 

produce semantically enhanced publications), Life Science Identifiers 

(LSIDs) to link to external databases like ZooBank and GenBank, 

specimen records or bibliographic citations, hosting of descriptions 

on dedicated servers such as Plazi, building a descriptions archive 

and distributing them; legal aspects regarding copyright, and how we 

envision the future. A future that will include registration of taxonomic 

names and descriptions, tight integration of descriptions with all the 

underlying databases, and journal production workflow producing 

semantically enhanced publications that can be harvested by robots, 

and thus will be part of the electronic taxonomic knowledge sphere. 

Finally, the development of the Code will be discussed in the light of 

such tremendous opportunities, which will allow linking hundreds of 

millions of pages and databases - and if the nomenclatural domain 

is part of Web2.0, increasingly accelerated descriptions of the world's 

taxa. 

Freshwater animal diversity assessment (FADA) – a project 

documenting biodiversity in continental aquatic ecosystems 

Estelle V. Balian 1 , Hendrik Segers 1 , Christian Lévêque 2 and Koen 

Martens 1 

1 

Freshwater Laboratory, Royal Belgian Institute of Natural Sciences, 

Vautierstraat 29 B-1000, Brussels, Belgium 

2 

Antenne IRD, MNHN-DMPA, 43 rue Cuvier, Case Postale 26, Paris 

cedex 05 75231, France 

A consortium of more than 100 taxonomists has compiled data on 

specific and generic biodiversity from all animal groups in inland 

waters worldwide (*) . The diversity and distribution of vertebrates, 

insects, crustaceans, molluscs and a suite of minor phyla is 

compared. Whereas the available data on vertebrates and some 

emblematic invertebrate groups such as Odonata (dragonflies and 

damselflies) allow for a credible assessment, data are mostly 

deficient for several other groups. This is owing to knowledge gaps, 

both in geographical coverage of available data and/or lack of 

taxonomic information. These gaps must urgently be addressed, 

either by liberating date from inaccessible repositories or by fostering 

further taxonomic research. A similar effort is required to compile 

environmental and ecological information in order to enable crosslinking 

and analysis of these complementary data sets. Only in this 

way will it be possible to analyze information on freshwater 

biodiversity for sustainable management and conservation of the 

world’s freshwater resources. 

Balian, E., C. Lévêque, H. Segers & K. Martens, 2008. Freshwater 

Animal Diversity Assessment. Hydrobiologia 595: 637 pp. Reprinted 

as Developments in Hydrobiology 198. 

- 20 - 

The Catalogue of Life – an e-Science Systema Naturae for the 

future 

Frank Bisby 

Species 2000 Secretariat, School of Biological Sciences, University 

of Reading, Reading, RG6 6AS, UK 

A significant success in the Linnaean enterprise was to synthesise a 

single comprehensive taxonomic backbone that could be distributed 

widely for practical use, and that collated a professional quality 

catalogue of the entire span of organisms. In the modern world the 

practical need for such a system becomes ever more pressing, as in 

indexing biodiversity knowledge on the internet, documenting global 

biodiversity as a whole, and for globalising biodiversity science in 

relation to food, ecosystem biology and modelling climate change. 

The Catalogue of Life is a global scale community-wide programme 

intended to collate such a backbone using a distributed e-science 

system, gathering expertise from across the taxonomic profession. It 

is focused on establishing a sustainable and functionally complete 

system that includes a species checklist and taxonomic hierarchy for 

all known extant Plants, Animals, Fungi and Micro-organisms. 

Bisby FA, Roskov YR, Orrell TM, Nicolson D, Paglinawan LE, Bailly 

N, Kirk PM, Bourgoin T, van Hertum J, eds (2008). Species 2000 & 

ITIS Catalogue of Life: 2008 Annual Checklist. CD-ROM; Species 

2000: Reading, UK. 

Documenting marine megabiodiversity 

Philippe Bouchet 

Muséum National d'Histoire Naturelle, 57 rue Cuvier, 75005 Paris, 

France. 

With one-fourth of all described marine biota, molluscs are the 

beetles of the sea. It is estimated that we currently know 80,000 valid 

described mollusc species (53,000 marine), with a yearly increment 

of about 580 species (350 marine). The shells of molluscs are both a 

blessing and a curse. They are a blessing because the fossil record 

of molluscs is exceptionally good, the global sampling of molluscs is 

the best for marine invertebrates, and post mortem remains at 

intensively studied coral reef sites show that 28% of the species are 

simply never collected alive. Shells are also a curse. Shell collecting 

has fueled taxonomic inflation, and there is a vast purgatory of 

nominal species that have not been recently critically re-evaluated: 

this is undoubtedly the main reason why there is not yet a world 

register of mollusc species. Additionally, most mollusc species have 

historically been named based on their shells alone - sometimes 

even atrociously "beach worn". There is concern for a broadening 

gap between documenting the molluscan diversity of the world, 

which involves assigning names, and backing this exercise with 

sound nomenclature. Admittedly, molluscan systematics is not the 

only branch of zoology that suffers from the instability of names in 

taxonomic limbo, but the magnitude of the problem is certainly 

unique to malacology: there are probably 100-150,000 molluscan 

species still to be named, and there is a graveyard of perhaps 

100,000 nominal species based on name-bearing types of (very) 

questionable taxonomic value.


The Codes: is reconciliation possible? 

Benoit Dayrat 

School of Natural Sciences, University of California, P.O. Box 2039, 

Merced, CA 95344, USA 

There have been several historical attempts to change zoological 

nomenclature, including some of its most fundamental rules, such as 

the Linnaean binominal form of species names. For instance, today’s 

systematists would probably be amazed at some daring proposals of 

new forms of species names made in the 1960s, when numerical 

taxonomy was the cause of intense debates in systematics. 

Developed in the recent years, the International Code of 

Phylogenetic Nomenclature, or PhyloCode, proposes new rules to 

defining clade names. Since May 2007, the PhyloCode is ‘complete,’ 

for it also includes an article on species names. The least we can 

say is that the PhyloCode and some reactions to it have been quite 

divisive within the systematic community. Although this may be 

sociologically regrettable, challenging scientific debates should not 

be viewed as a bad thing. There are undeniable points of 

incompatibility between the current versions of the Code (ICZN) and 

the PhyloCode. However, both codes also share several important 

points and goals, and one should not exclude that they could 

potentially be reconciled or, at least, co-exist and influence each 

other. Whether or not this is possible will be discussed: special 

attention will be given to the treatment of species names in the 

PhyloCode and its potential consequence on Linnaean binomials. 

Flying after Linnaeus: Dipteran names since Systema Naturae 

(1758) 

Neal L. Evenhuis 1 , Thomas Pape 2 , Adrian C. Pont 3 and Christian F. 

Thompson 4 

1 

Department of Natural Sciences, B.P. Bishop Museum, Honolulu, 

Hawaii, USA 

2 

Natural History Museum of Denmark, Zoological Museum, 


3 

Oxford University Museum of Natural History, Oxford, United 

Kingdom 

4 

Systematic Entomology Laboratory, Washington, DC, USA 

Since the start of zoological nomenclature, new names for flies 

(Diptera) have been proposed at an average of more than eight 

hundred names per year. 

Linnaeus (1758: Systema Naturae) and Fabricius (1805: Systema 

Antliatorum) provided the first and the last comprehensive 

summaries of contemporary knowledge of extant flies respectively. 

Today, version 10 of the BioSystematic Database of World Diptera 

(BDWD) contains nomenclatural data for 156,599 extant and extinct 

species in 154 families and 11,671 genera, which is about 10% of 

the described diversity of the world biota. The BDWD is a 

nomenclator, which allows for the retrieval of the single correct name 

for each dipteran, and by providing a framework for organizing and 

integrating current and future knowledge it is a comprehensive portal 

to knowledge about all flies. 

Various statistics are presented to assess the growth in knowledge, 

from species accumulation curves and regional patterns of 

taxonomic output, to the number of workers, their productivity and 

error rates. With an estimated 98% of all Diptera names entered in 

the BDWD, the challenge now is to assure the highest quality 

through appropriate community participation, particularly by 

specialists and peers. At present, 15% of the entries have been 

verified and peer-reviewed. 

- 21 - 

Web taxonomy - the future, or a distraction? 

Charles J. Godfray 

University of Oxford, UK 

There is little argument that the World Wide Web and related 

technologies have a huge amount to offer taxonomy. More 

controversial is whether taxonomy should become a wholly webbased 

enterprise, dispensing with the paper outputs that have been 

its mainstay since the days of Linnaeus. Indeed, critics of web 

taxonomies have argued that it is a dangerous distraction, 

undermining the scientific basis of taxonomy and redirecting 

resources away from more important areas of the field. I will discuss 

the pros and cons of web-based taxonomies and explore whether 

methods of working on the web can be devised to address the major 

worries of its critics. Taxonomy is currently faced with the challenges 

of addressing the major policy needs of the biodiversity community, 

as well as the scientific challenges of incorporating the flood of 

sequence and related molecular data that will snowball over the 

coming years. I will argue that moving to the web will be critical for 

the subject to survive and flourish in the face of these pressures. 

The Encyclopedia of Life 

James Hanken 

Harvard University, Museum of Comparative Zoology, Cambridge, 

MA, USA 

The Encyclopedia of Life is a global biodiversity initiative that was 

launched officially in May 2007. Its fundamental goal is deceptively 

simple: provide a website for every species of living organism. In 

reality, EOL is an ambitious, if not audacious, project that seeks to 

readily make available online and at no cost as much biological 

information regarding a given species as can be obtained legally and 

in digital form. In addition to its core activities involving Biodiversity 

Informatics and construction of Species Sites, key components 

include Biodiversity Synthesis, Education and Outreach, and 

Scanning and Digitization of the primary literature of comparative 

biology and taxonomy since Linnaeus. The Encyclopedia of Life is 

not a taxonomic activity per se, but it has the potential to 

tremendously enhance the work of practicing taxonomists and 

systematists, especially those in developing countries, and thereby 

facilitate the discovery, naming and classification of Earth’s biological 

diversity. At the same time, EOL requires—and seeks—the active 

participation and involvement by the professional taxonomic 

community in order to succeed and achieve its full potential as a 

reliable and up-to-date source of information for scientists, educators, 

conservationists, environmental planners, government policymakers, 

students, laypeople, and other “consumers” of biological data 

worldwide. The Encyclopedia of Life is one of several current 

projects that together may soon realize the grand vision of a 

seamless global biodiversity informatics infrastructure for use by 

science and society. 

Future taxonomy today: new tools applied to accelerate the 

taxonomic process. 

Norman Johnson 

Ohio State Univ., 1315 Kinnear Road, 43212, Columbus, Ohio, USA 

In the 250 years since the publication of the tenth edition of Systema 

Naturae only a small fraction of the total number of species of 

animals have been formally described and named. In the face of the 

many threats to biodiversity, substantial progress in the disco! very 

and description of species is needed. The Platygastroidea Planetary 

Biodiversity Inventory project seeks to treat a substantial number of 

species in the course of five years. The community of experts on this 

taxon is small and scattered around the world. New methods and 

tools are required in order to maximize the efficiency of the time 

devoted to research on these parasitic wasps as well as to 

accelerate and enhance the formal taxonomic publication process. 

The taxonomic workflow is modelled, dividing the tasks into two 

discrete domains: research and publication. The tools and methods 

developed and used in each domain in the project are described and 

illustrated.


e-Publish or Perish 

Sandra Knapp 1 and Debbie Wright 2 

1 

Department of Botany, The Natural History Museum, Cromwell 

Road, London SW7 5BD, UK 

2 

Wiley-Blackwell, 9600 Garsington Road, Oxford, OX4 2DQ, UK 

Access to the published literature of science is increasingly 

becoming digital. Journals provide on-line early access, and as the 

Internet reaches more corners of the world, scientists are turning to 

electronic publications as a first port of call in doing research. As the 

world becomes more and more digitally organised and accessible, it 

is critical that taxonomic work is not left behind to languish in printonly 

journals. Taxonomic publication is perhaps unique in the 

scientific literature in remaining relevant for nomenclatural purposes 

“forever”, thus creating a conservative and cautious attitude amongst 

taxonomists for changes in publication methods. The two central 

pillars of all four organismal codes (zoology, botany, bacteria and 

cultivated plants) are validity of publication and the principle of 

priority, but the rules governing publication are arguably the most 

important, as it is through reference to publications that hypotheses 

as to taxon identity are supported or rejected. We will discuss the 

main issues besetting acceptance of electronic publication for 

nomenclatural acts – essentially new names, synonymies and 

typifications. These are 1) archiving, 2) accessibility, 3) date of 

publication and 4) type of electronic medium. We will also touch on 

how electronic publication might interface with registration of names, 

another subject of considerable debate in some taxonomic 

communities. Issues surrounding access to new names and 

nomenclatural acts transcend their medium of publication, but as our 

world becomes digital, taxonomists must not be left behind. The 

products of taxonomy are just too useful to suffer that fate. 

Linnaeus – Sherborn - Zoobank 

Ellinor Michel 1 and Andrew Polaszek 2 

1 International Commission on Zoological Nomenclature, c/o Natural 

History Museum, Cromwell Road, SW7 5BD, London, UK 

2 Dept. of Entomology, Natural History Museum, London, UK 

The monumental task of cataloguing the scientific names of animals, 

begun 250 years ago by Linnaeus, is entering a new golden age. 

Within the 823 pages of the 10 th edition of Systema Naturae 

Linnaeus catalogued 313 genera and 4,397 species. The journal 

Zoological Record, currently the most exhaustive annual catalogue 

of newly-published animal names and nomenclatural acts in zoology, 

includes approximately 1.7 million names in the database on which it 

is based - the Index of Organism Names. Zoological Record started 

in 1864 and has continued uninterrupted to the present day, 

cataloguing about 16,000 new names every year. Between 1902 and 

1933 Charles Davies Sherborn published the formidable Index 

Animalium - 11 volumes (including the indexes) and more than 9,000 

closely-typed pages covering animal names from 1758-1850. 

Sherborn personally researched and checked information for Index 

Animalium almost exclusively from the original sources. It is therefore 

astonishing that 20 years after Sir Robert May lamented the absence 

of a "centralized computer index of these recorded species" in the 

journal Science (referring to all organisms), such a catalogue is still 

incomplete. The encouraging news is that efforts over the last four 

years are bringing together several initiatives, including those 

mentioned above but also several that we will hear about during this 

meeting. These initiatives, together with the establishment of 

ZooBank as a registry for animal names and nomenclatural acts, 

suggest that the combined vision of Linnaeus, Sherborn, May, 

Wilson and others will soon become a reality. 

- 22 - 

Future taxonomy - bigger, better, faster. 

David Patterson 

Informatics lead, 7 MBL Street, MA 02543, Woods Hole, USA 

Taxonomy is usually seen as the source of our catalog of biodiversity, 

but taxonomists also have been managers of information about 

organisms. The informatics role has been made possible through two 

features that were embedded in taxonomic practices established by 

Linnaeus, the use of names and h! ierarchies. With the maturation of 

the internet into a near universal network interconnecting distributed 

information, taxonomy has new devices available to it to assist in its 

role of managing information. The start of 21st century is a time of 

special opportunity to reinvent taxonomy with much greater 

relevance and visibility. This requires taxonomists to acknowledge 

and develop their role as informaticians. Taxonomy can extend 

beyond the development of the catalog of diversity, but to do so it 

must commit to communal processes rather than individual products, 

to eschew parochiality, to set an aggressive agenda of getting all 

useful information into publicly accessible internet-accessible forms, 

and to reassess familiar issues such as nomenclature to develop the 

use of names as metadata in on-line information management. The 

incorporation of taxonomic practices into the design of data bases, 

software, and services is referred to as taxonomic intelligence. A 

taxonomi! cally intelligent names-based infrastructure underpins the 

Encyclopedia of Life project. EOL has confirmed the capacity of a 

names-based infrastructure to interconnect any data object for any 

taxon, to represent alternative views of taxa and systematics, to 

create new and accessible identification tools, to accelerate 

discovery and the quality of taxonomic judgments, and the evolution 

of taxonomic concepts. 

Zoobank : reviewing the first two years, and preparing for the 

next 250 

Richard Pyle, Miguel Alonso-Zarazaga, Nina G. Bogutskaya, 

Philippe Bouchet, Denis J. Brothers, Daphne G. Fautin, Mark J. 

Grygier, Bruce R. Halliday, Maurice Kottelat, Frank-Thorsten Krell, 

Sven O. Kullander, Gerardo Lamas, Susan Lim, Shunsuke F. 

Mawatari, Alessandro Minelli, Peter K.L. Ng, Thomas Pape, Laszlo 

Papp, David J. Patterson, Gary Rosenberg, Pavel Stys, Jan van Tol, 

Zhi-Qiang Zhang 

International Commission on Zoological Nomenclature, 

http://www.iczn.org/ 

In 2005, the ICZN Secretariat and Commissioners proposed 

“ZooBank” as a web-based registry of zoological names and 

nomenclatural acts. The ZooBank web site was launched as a 

prototype on January 1 st , 2008, coinciding with the 250 th anniversary 

of the official start of Zoological Nomenclature. At its launch, the 

ZooBank registry included all 4,819 names established in the 10 th 

edition of Linnaeus’ Systema Naturae, as well as five new fish 

species names established in an article published concurrently with 

the launch of ZooBank. 

ZooBank is not intended to replace existing nomenclatural catalog 

databases, and it makes no assessment or judgment of the 

taxonomic content of any published work. ZooBank assigns unique 

registration identifiers in four information domains of relevance to the 

ICZN Code: nomenclatural acts (including new names and other acts 

that affect existing names), publications, authors, and type 

specimens. These identifiers are envisioned as pointers to 

authoritative information concerning zoological nomenclature, and 

are expected to become integral to current and future efforts to index 

taxonomic content. 

The complete implementation details of the ZooBank registry are 

currently being discussed, developed, and tested, with involvement 

from ICZN Commissioners, nomenclatural data managers, and the 

taxonomic community at large. Many questions concerning technical 

implementation details, content sourcing and prioritization, 

information quality standards, and scenarios for mandatory 

registration are open to discussion. As much as there is an urgent 

need to answer these questions soon, there is also the need to “get it 

right”, ensuring a solid foundation for the next 250 years of zoological 

taxonomy.


The all genera index 

David Remsen 

GBIF, Universitetsparken 15, 2100, Copenhagen, Denmark 

The All Genera Index (AGI) represents an effort to identify all current 

and historic code-compliant genera names. It is an index that is 

dynamically derived from a range of partnering nomenclatural 

databases accessible through a common point of discovery. The 

indexed nomenclatural records are accessed through a common 

taxonomic names architec! ture being developed by GBIF and 

partners. The AGI application includes a comprehensive names 

management taxonomy that places all the genera within a 

provisional taxonomic framework. A comprehensive index of all 

genera provides a number of useful outcomes. First, as a 

nomenclatural reference, it will provide the means to avoid the 

creation of new generic homonyms – not only with names of animals, 

but also of plants and other organisms. Second, it will serve to 

identify all existing homonymous genera. In so doing, it will also 

identify the genera that are lexically distinct. As genera are a 

component of all species and infra-species names, a provisional 

taxonomic assignment for all genera can be extended to include all 

species combinations. This can provide a framework for biodiversity 

data mobilization initiatives such as GBIF and the Biodiversity 

Heritage Library to better organize the names associated with the 

data objects they index and serve, particularly those names not 

accounted for within taxonomic contexts available to these initiatives. 

This will serve to demonstrate the need for these and other data 

mobilization initiatives to embed ta! xonomic methodologies into 

generalized biodiversity data management. 

250 years of Swedish Taxonomy 

Fredrik Ronquist 

Swedish Museum of Natural History, Box 50007, SE-10405 

Stockholm, Sweden 

Linnaeus and his contemporaries aimed at nothing less than a 

complete inventory of the flora and fauna of the planet. At least in 

Sweden, the goal seemed easy to reach. However, biologists later 

realized how daunting the task actually was even in Sweden. The 

entomologists, in particular, were discovering a staggering number of 

species. Linnaeus listed some 1,600 insect species from the country. 

By 1920, almost 14,000 more had been added. While decreasing 

ranks of stubborn taxonomists kept at it, mainstream biologists lost 

patience in the effort and turned their attention to the fundamental 

principles of biology instead, focusing on easy-to-study model 

organisms. It was not until the Convention on Biological Diversity put 

the inventory of the planet's diversity on the top of political agendas 

that the Swedish taxonomy effort was revitalized through the 

Swedish Taxonomy Initiative. Launched in 2002, the project aims to 

complete the inventory of the Swedish flora and fauna in 20 years. It 

is now clear that the task was more difficult than initially thought. The 

biggest challenges appear to be among the insects, comprising half 

of Swedish macroscopic diversity. In 2003, the insect fauna was 

estimated at 25,000 species. Since then, some 2,000 species have 

been added, about one third of which are new to science. Although 

taxonomists are now working actively on many of the critical groups, 

we estimate that 4,000 species still remain to be discovered. The 

missing species are predominantly parasitic or saprophagous, 

demonstrating significant bias in earlier biodiversity maps. 

Provisional nomenclature: the on-ramp to taxonomic names 

David Schindel and Scott Miller 

Consortium for the Barcode of Life, Smithsonian Institution 

Published names are the basis of information exchange and retrieval 

in taxonomy, but the preparation of formal names for publication is 

proving too slow. Identification of specimens to species normally 

involves three steps: (1) sorting into groups, often based on a single 

character system; (2) refining those groups through detailed analysis 

of accepted, multiple character systems; and (3) associating the 

groups with published species names based on review of literature 

and type specimens. DNA barcoding, a standardized molecular 

technique, can be used by non-experts for first-stage sorting into 

DNA sequence clusters. 

Specimens that have passed through the second step sometimes 

cannot be associated with known species. 

- 23 - 

The third step, validating the species’ novelty, requires extensive 

comparison with types. Ironically, formal species names cannot be 

used publicly during the validation process, to protect the name’s 

priority. Pre-published names have become trade secrets, protected 

against piracy so authors may get their due credit. As a result, 

validity of proposed taxonomic concepts can only be judged after 

publication. 

During the long validation phase, researchers often use provisional 

species labels, not names, associated with voucher specimens and 

GenBank records. They support information management for 

"interim taxonomy" (Erwin 1991) and can be treated as provisional 

species concepts. 

A system of provisional labels would enable researchers to publish 

interim results and conduct critical comparisons. Provisional labels 

would have to be stable and unique, and would be cited in proposing 

the formal name. Provisional taxonomy and informatics can 

accelerate a more collaborative, consensus-based team approach to 

taxonomy. 

Grasping the taxonomic diversity of Life: barcoding and 

integration of infrastructures 

Simon Tillier 

EDIT, CP43, Museum national d’Histoire naturelle (UMR 7138), 57 

rue Cuvier, 75231 Paris cedex 05, France 

Since the arrival of phylogenetics, taxonomy has undergone more 

changes in its principles, methods and techniques in the last 

decades than ever before since Linnaeus’ time: computer-aided 

data processing, web access to information and genetic characters 

constitute a revolution with heavy consequences on the usage of 

names, specimens and taxonomic concepts. 

By allowing access to names and related taxonomic concepts 

illustrated by a specimen through genetic characters using the web 

and phylogeny algorithms, DNA barcoding reunites classical 

identification with the full array of these new methods. 

Simultaneously the expert is not involved in the identification process 

itself, as the operational species concept is defined by the algorithm, 

and de facto the barcode voucher replaces the holotype as the link 

between the name and the concept. This results in increased 

responsibility for taxonomists in the application of names through 

initial identification of the voucher. For users, it opens a new world by 

providing access to names and concepts at an unprecedented speed, 

providing a chance to apprehend the actual taxonomic diversity and 

to better connect local identifications to a single global classification. 

To implement these new developments in daily taxonomic work, for 

the benefit of users (who will have access to identification) and of 

taxonomists (who will be able to concentrate on scientifically creative 

tasks), infrastructure is crucial and must be unobtrusive. However, 

taxonomic infrastructures are not only underdeveloped with respect 

to new developments, but also heavily fragmented as the result of 

their history. It is the role of networks, and particularly of EDIT at EU 

level, to set up the institutional organisation which will allow transition 

from the infrastructures of Linnean times to the new global 

distributed taxonomic system. 

250 years: enough animal nomenclature 

Alfried P. Vogler 

Imperial College London and Natural History Museum 

Although deceptively simple, the binomial system lacks practical 

utility in animal nomenclature, in particular in highly diverse groups. 

Open access and encyclopaedia-style resources facilitate the access 

to taxonomic information but they do not fundamentally deal with the 

problem of recognising and identifying the named groups. DNA 

sequences can overcome these issues. As studies in recent years 

have shown, sequence variation in animals is highly clustered and 

therefore provides an ideal system to delimit groups which can be 

easily re-identified with existing search algorithms. This obviates the 

need for a binomial system in animal nomenclature, as clusters of 

sequence variation can be coded with any identifier. The validity of 

such DNA-based system requires that the recognised entities have 

biological (reproductive coherence) and historical (common ancestry) 

relevance, and that they discriminate between groups. Unless the 

work of the past 250 years is ignored, the DNA-based taxonomy also 

needs to be linked to the existing system. None of this is a problem 

in principle. The talk will show the application of the approach to the 

insects.


S6 - Paleozoology and comparative anatomy:exceptional conservations 

Non-bilaterians in the early fossil record 

Stefan Bengtson 

Swedish Museum of Natural History, Stockholm, Sweden 

Given their basal positions in the metazoan phylogenetic bush, 

sponges, ctenophores and cnidarians should be expected to have 

played a major role in the first grand diversification of animals, during 

the Neoproterozoic–Cambrian transition, some 570–530 million 

years ago. The fossil record is far from clear with regard to these 

events, however, partly because preserved characters in nonbilaterian 

fossils are often too few and ambiguous for meaningful 

phylogenetic analysis. The Neoproterozoic–Cambrian record of 

exquisitely preserved embryos, body fossils and skeletal 

components of non-bilaterians is growing in extent and quality, 

shedding light on the early diversification of metazoans. The fossils 

may share primitive characters that have been lost in living 

representatives of the major groups, which helps us identify 

relationships within the early branches of the metazoan bush, 

including extinct lineages. A very early ctenophore-like fossil, 

sponge-like fossils with cnidarian features, and embryos of strong 

cnidarian aspect will be used to illustrate these points. 

Remarkably preserved marine invertebrates from the Silurian of 

Herefordshire, England 

Derek E.G. Briggs 1 , Derek J. Siveter 2, 3 , David J. Siveter 4 and Mark D. 

Sutton 5 

1 Department of Geology & Geophysics, Yale University, PO Box 

208109, New Haven, CT 06520-8109, USA 

2 Geological Collections, University Museum of Natural History, 

Oxford OX1 3PW, UK 

3 Department of Earth Sciences, University of Oxford, Parks Road, 

Oxford OX1 3PR, UK 

4 Department of Geology, University of Leicester, Leicester LE1 7RH, 

UK 

5 Department of Earth Sciences and Engineering, Imperial College, 

London SW7 2BP, UK 

A remarkable assemblage of marine animals is currently emerging 

from study of a 425 million-year-old (Silurian) volcanic ash in 

Herefordshire, England. Tiny animals, most just a few mm in 

dimension, are preserved in carbonate concretions. The fossils 

cannot be extracted by mechanical or chemical means – nor are they 

visible to x-radiography or scanning methods. The specimens must 

be ‘prepared’ by grinding them away microns at a time to produce 

high resolution 3-D reconstructions that can be manipulated on a 

computer. The fidelity of preservation of even soft-part anatomy 

allows the morphology of these animals to be investigated at a 

comparable level of detail to their modern counterparts. The rare and 

spectacular fossils include many animals that are very poorly 

represented in the fossil record, as they lack any biomineralized 

skeleton. The Herefordshire deposit represents one of very few 

exceptionally preserved fossil biotas (Lagerstätten) known from the 

~70 Ma period between the Cambrian and Devonian; it provides a 

unique and important window on Palaeozoic marine life. The biota 

includes a diverse suite of arthropods: a stem-group chelicerate and 

crustacean, a phyllocarid, ostracods, a larval barnacle, a pycnogonid 

and a marrellomorph. It also includes radiolarians, a diversity of 

sponges, a polychaete worm, an aplacophoran-like mollusc, a 

platyceratid gastropod, a brachiopod with pedicle, several 

echinoderms including an asteroid with preserved tube-feet, and 

many other organisms of uncertain affinity. These fossils are yielding 

critical information on the evolutionary history and relationships of 

living invertebrate taxa. 

- 24 - 

Bone growth marks suggest protracted growth in Apteryx (Aves, 

Neornithes, Ratitae) 

Jacques Castanet, Estelle Bourdon, Jorge Cubo and Armand de 

Ricqlès 

Equipe « Squelette des Vertébrés », UMR CNRS 7179. UPMC, 2, pl. 

Jussieu, case 19,75005 Paris, France 

Skeletochronology is a broadly used method that utilizes bone 

growth marks (BGMs) to infer life history traits in tetrapods. In 

modern birds, however, the presence of BGMs and their use for 

individual aging remains controversial. Because most living birds 

achieve their complete skeletal development in less than one year, 

BGMs are either absent or scarce and restricted to the outer part of 

bone cortices. A BGM pattern similar to that of non avian reptiles is 

known in some non-neornithine birds and in extinct New Zealand 

moas (Dinornithiformes). Till recently, BGMs were regarded as 

unknown in living ratites (Turvey et al. Nature, 2005: 435). This is 

now contradicted by the discovery of BGMs in the long bones of 

Apteryx australis (Apterygidae). The occurrence of 7 or 8 wellmarked 

lines of arrested growth (LAGs) in hindlimb bone cortices of 

an adult suggests that this individual was at least 7 or 8 years old. In 

the femur, the first 3 or 4 LAGs are separated by layers of parallelfibered 

bone, weakly vascularized by primary vascular canals. In the 

tibiotarsus and tarsometatarsus, these inner LAGS are locally erased 

by bone remodelling. The 4 outer LAGs are closer to each other and 

located in bone periphery. This LAG pattern and the decrease of 

vascular density towards the periosteum suggests that Apteryx, in 

contrast to other living birds, does not reach its adult body size until 

up to 4 years of age and subsequently shows a prolonged periostic 

osteogenesis during at least 4 more years. 

Cambrian representatives of the tongue worms: ontogeny and 

taxonomy 

Christopher Castellani, Dieter Waloszek and Andreas Maas 

1Workgroup Biosystematic Documentation, Helmholtzstr. 20, 

University of Ulm, 89081 Ulm, Germany 

During the last twenty years, several authors have tried to solve the 

problem related to the systematic position of an enigmatic group of 

parasites, the Pentastomida (tongue worms), but without convincing 

success. Two major assumptions result from this controversy. The 

first, supported mainly by investigations of sperm morphology and 

nucleic acids, assumes close alliance to the eucrustacean in-group 

Branchiura (fish lice). The second, founded on morphological, 

embryological, anatomical and fossil data, points to a derivation from 

even before the Arthropoda s. str. level, i.e. before achievement of 

arthropod characteristics such as sclerotised body tergites, 

compound eyes, or segmented limbs with exopods. In 2004, new 

material of 3D-preserved (‘Orsten’-type-preserved) Cambrian 

representatives of the tongue worms was obtained from a piece of 

limestone rock collected in Västergötland, Sweden. In the framework 

of the EU programme Molmorph we study more than 60 specimens 

of different sizes, using two major techniques, SEM with 

standardised views and biometry combined with statistical tools. 

Preliminary measurements suggest that the new material might 

represent not only different instars, at least three, but also contains 

different species, at least two. The large material permits to 

investigate all major external features of the fossils in detail. These 

will be used in a re-evaluation of the taxonomy status of all Cambrian 

fossil taxa described so far. In the light of the new data, we expect to 

contribute further to a solution of the unclear systematic position of 

Pentastomida.


Study of scale morphology of blackspot threadfin Polydactylus 

sextarius (Bloch & Schneider, 1801) (Actinopterygii: 

Polynemidae) using scanning electron microscope (SEM) 

Hamid Reza Esmaeili, Fatemeh Laghaei and Azad Teimory 

Department of Biology, College of Sciences, Shiraz University, Iran 

Shape, size, and number of scales are suitable tools in fish 

taxonomy and using it dates back to first half of 19 th century when 

Agassiz (1833-1843) used it in fish taxonomy for the first time. He 

classified the fishes into 4 group based on the scale morphology. 

Besides its use in classification, scale morphology is an important 

tool in determining the diet of piscivorous predators or in the 

paleontological analysis. During the late 19 th century and the first half 

of the 20 th century, and with the great advancements in the field of 

light microscopy, the importance of scale morphology in systematics 

increased significantly. The importance of scale morphology used in 

classification was strengthened with the introduction and 

development of scanning electron microscopy (SEM), so that scales 

of many different fish species have been studied using SEM (Jawad 

et al., 2007). With regards to the importance of scale morphology, in 

this research work, ultrastructure of scale of blackspot threadfin fish 

Polydaactylus sextarius from Persian Gulf, were studied. To study 

scale morphology, the scales from below the dorsal fin and lateral 

line were removed, cleaned in suitable solution and were prepared 

for the scanning electron microscopy. Results showed that the focus 

in this ctenoid scale located in the posterior part of scale and is clear 

and has granules. Few radii were found only on the anterior part of 

scale. All the radii are in primary type (originating from the focus and 

reach to the scale margin). These radii divided the anterior part of 

scale into several regions. Hence the scale is sectioned. Tongue or 

lobes of the anterior part are very distinct. Presence of the lepidonts 

on the crest of circuli is another character of this fish scale. Inter 

circular space is without granules. The circuli ends at the posterior 

part of scale and the small granules named tubercles are formed. On 

the most posterior part of scale, tubercles are replaced with the 

sharp conical structures called ctenii. It seems that focus shape, 

number, shape and size of lepidonts, ornamentation of tubercles and 

ctenii are important in taxonomy of this fish. 

- 25 -


Evolution of main groups of Bryozoa during Phanerozoic : 

present data 

Françoise P. Bigey 

9 rue de Douai F -75009 Paris France 

Among lophophorates, bryozoans appear as peculiar organisms 

especially on account of significant extent in present marine 

environments and in many past marine environments as well. It 

concerns stenolaemates and gymnolaemates. Fossil 

phylactolaemates, fresh water colonies without mineralised skeleton, 

are revealed by statoblasts only, in some cases on account of 

difficult preservation. Unlike brachiopods, the other main lineage of 

lophophorates, the oldest bryozoans were described from ordovician 

strata. 

If main prevailing paleozoic orders (cystoporates, trepostomates and 

fenestrates) are extinct, they were replaced by other ones as 

cheilostomes, since Upper Jurassic and, predominating now. 

Mention must be done of ctenostomates, free of calcareous skeleton, 

but recognised by immuration process since Ordovician. 

Cyclostomates, widely developed in Jurassic and Cretaceous, are 

known from Ordovician too. 

Bryozoan evolution is characterised by links with main biological 

crisis, particularly the permo-triassic one, quite drastic for these 

organisms, even though few of them survived in Lower Triassic. 

Recovery in marine environment occurred only during Jurassic. From 

Middle Cretaceous cyclostomates/cheilostomates ratio in 

assemblages begins to reverse for the benefit of last ones. This ratio 

did not widely evolved since Eocene. 

Bryozoan systematics as a whole (fossil and recent forms) is 

established from skeletal characters. Phylogenenetic relations of 

cyclostomates, for instance is based on wall types. Cladistic analysis 

is used to propose models at different levels of organisation. For 

recent bryozoans data from DNA are promising. 

Why are there more genera of Mutillidae (Hymenoptera) with 

wingless males in Southern Africa than elsewhere? 

Denis J. Brothers 

School of Biological and Conservation Sciences, University of 

KwaZulu-Natal (Pietermaritzburg), Private Bag X01, Scottsville, 3209 

South Africa 

The velvet ants of the family Mutillidae (Hymenoptera: Vespoidea) 

are larval parasitoids of the enclosed immatures of other insects, 

specially other aculeate Hymenoptera (bees and wasps). Mutillid 

females are all completely apterous, with the mesosomal (thoracic) 

sutures entirely fused (or almost so) so that the mesosoma is a 

simple box-like structure. Males are generally fully winged, with the 

mesosoma essentially unmodified, and with almost all sutures 

articulating. This dimorphism has been related to biology: wings 

would be an encumbrance and susceptible to damage for females 

searching underground or in burrows for hosts, and good motility 

(flight) would be required for the discovery of mates by males, 

avoidance of inbreeding and possibly dispersal. Associated with the 

dimorphism in wing development is differentiation in other features 

(such as colour pattern), so that the two sexes of a single species 

can generally not be associated on the basis of morphology. 

Nevertheless, there are several species (and genera) in which males 

show wing reduction to varying degrees, with loss of flight ability, 

from brachypterous with the mesosoma scarcely modified through 

several intermediate steps to completely apterous and with the 

mesosomal sutures entirely fused as in the female. Apterous males 

generally look very similar to their females. 

This poster provides an illustrated survey of the range of wing 

reduction in male Mutillidae, its distribution across higher taxa, 

genera and geographically, and attempts to draw some conclusions 

and identify explanatory patterns. 

S7- Paleontology and Evolution 

- 26 - 

Palaeontology: a load of old stones? 

Simon Conway Morris 

Downing Street, CB2 3EQ, Cambridge, England 

Some years ago the distinguished evolutionary biologist John 

Maynard Smith welcomed palaeontology back to the High Table; a 

very English type of invitation with the implication that one would be 

subject to a series of terrifying conversations whilst drinking 

inordinate quantities of wine. In fact the last few years may have 

been quite benign for palaeontology, with wide-spread interest 

amongst evolutionary biologists, as well as being marked by a 

steady stream of hi! gh profile papers in Nature and Science (and, of 

course, elsewhere). But as we all know to rest on one's laurels can, 

sooner or later, lead to a strong and distressing smell of 

decomposition. As ever we need to look to the future. In this 

presentation I will outline a few areas where we should be able to 

contribute to the wider conversations in evolution and the earth 

sciences. How well do we know life in very deep time, especially in 

the nether regions of the Precambrian? Are we any closer to 

explaining the Cambrian explosion? Is the influence of mass 

extinctions over-rated? Can we identify directionality and increasing 

complexity, even progress, from the fossil record? Does anybody still 

seriously subscribe to the metaphor of Stephen Jay Gould that were 

we to re-run the tape of life we would end up with a completely 

different biosphere? So too may we finally declare punctuated 

equilibrium dead and buried? What real links exist between the 

fashionable areas of evo-devo and the transformations we see in the 

fossil record? Where are the real puzzles in terms of identifying 

transitional groups and why apparently are they so difficult to 

decipher? Do we radically under-estimate the importance of nonuniformitarian 

worlds: What is ecology really like in deep time? Will 

we be in any position to advance our understanding of the evolution 

of behaviour, as! well as sensory systems, communication and 

brains, even the emergence of consciousness? And to finish on an 

absurd note: What future for astrobiology? 

The Lilliput Effect in Cretaceous-Tertiary (K-T) Planktonic 

Foraminifera 

Norman MacLeod 

Palaeontology Department, The Natural History Museum, Cromwell 

Road, London, UK, SW7 5BD 

The stratigraphic record of K-T planktonic foraminifera represents an 

outstanding target for probing the dynamics of the Lilliput Effect. 

Preliminary analysis indicates this event began prior to the 

emplacement of bolide impact debris. Event initiation is marked by a 

rapid decrease in test size among K-T survivor taxa and defined by 

the maintenance of these small-size populations, along with the 

appearance of small, fully Danian species, for the first 40,000– 

200,000 years of the Danian. The atypically small size of the latter is 

a direct result of speciation from the former. Correction for 

phylogenetic covariation reveals the presence of a strong 

phylogenetic signal in generic size data. In particular, it appears the 

extinction of larger-sized Cretaceous species had little effect on the 

evolutionary size dynamics of the Lilliput faunas. Finally, morphotype 

analysis shows that the K-T Lilliput interval represents a transition 

between faunas exhibiting a diversity of trochospiral and flaring tests 

to those composed almost exclusively of rounded trochospiral forms. 

This, in turn leads to a subdivision of the K-T planktonic foraminiferal 

Lilliput event into a two-stage structure: Stage 1 (Zone P0) 

representing a small-sized, flared test-dominated fauna and Stage 2 

(Zone P1a) representing a slightly larger sized rounded trochospiredominated 

fauna. The entire event exhibits a duration of c. 300,000 - 

500,000 years. Application of this data analysis strategy to the 

succeeding Cenozoic interval illustrates the important role phylogeny 

should play in understanding the evolutionary history of organismal 

size.


Assessing confidence intervals for stratigraphic ranges of 

higher taxa: the case of Lissamphibia 

David Marjanović and Michel Laurin 

CNRS, UMR 7179, Équipe Squelette des vertébrés : aspects 

fonctionnels et évolutifs, U. Paris 6, Case 19, 4 place Jussieu, 75252 

Paris cedex 05, France 

To evaluate stratigraphic evidence for the time of origin of the clade 

of extant amphibians (Lissamphibia), we attempt to establish a 

confidence interval on the lower bound of the stratigraphic range of 

this clade. This is based on the stratigraphic distribution of 1207 

fossiliferous localities that have yielded lissamphibians, the relative 

area of sedimentary rocks from various periods (upper Paleozoic to 

present) exposed on the continents, and ten exponential-growth 

models of lissamphibian diversity that differ according to the 

assumed effects of three major biological crises and the assumed 

starting times of lissamphibian diversification. This method does not 

rely on a phylogeny of Lissamphibia, but only assumes that fossils 

which belong to Lissamphibia have been identified as such. The 

results suggest a more recent (Permian) origin of Lissamphibia than 

advocated in most recent molecular studies, which proposed an 

origin in the Late Devonian or Early Carboniferous. Our results are 

also more compatible with the monophyly than the polyphyly of the 

extant amphibians, but depend heavily on poorly constrained 

assumptions about lissamphibian extinction rates during biological 

crises. Counts of lissamphibian diversity through time that consider 

ghost lineages and stage durations show moderate declines across 

the Cretaceous-Paleogene and Oligocene-Miocene boundaries. 

Sharks as indicators of trophic structure within ‘mid’ 

Cretaceous watermasses 

Emma-Louise Nicholls 

Department of Earth Sciences, University College London, Gower 

Street, London, WC1E 6BT , UK 

The Elasmobranchii were apical predators during the Cretaceous 

period and their presence crucial for regulating the balance of 

oceanic food-webs. In the ‘mid’ Cretaceous a period of transgressive 

episodes caused a eustatic rise, meaning the destruction of shallow 

water environments as well as the deepening and of large expanses 

of ocean. The subsequent effects of the ‘mid’ Cretaceous 

transgressive episodes upon the radiation, diversity and 

palaeobiogeography of the Elasmobranchii are being investigated in 

order to establish the effects of the transgressions upon the 

Cretaceous ecosystems. This research is the first study to use 

organisms of high trophic level as palaeoenvironmental indicators, as 

sharks are virtually unique amongst vertebrates in yielding 

statistically large numbers of specifically identifiable remains. In 

order to establish the diversity and spatial variations of the 

elasmobranch faunas, sediment from a number of well-defined timeplanes 

during critical intervals of the sea-level rise are being sampled. 

Localities within the UK include the Anglo-Paris Basin, the North Sea 

Basin and the intervening shallows of the East Midlands Shelf. 

The Case of the Lamnid-Orectolobe. Where does 

Palaeocarcharias belong? 

Emma-Louise Nicholls and David J. Ward 

Department of Earth Sciences, University College London, Gower 

Street, London, WC1E 6BT, UK 

Palaeocarcharias stromeri is an Upper Jurassic selachian from the 

Solnhofen area of southern Germany and Cerin, France. Three 

specimens have been recovered including the near-complete 

holotype at Jura Museum in Eichstätt. Its fusiform body plan is 

reminiscent of the Orectolobid morphology though the teeth, that 

show linear gradient monognathic heterodonty, are high-cusped 

which is a morphology considered unique to the Lamniformes. 

Subsequently, Palaeocarcharias was originally placed within the 

Lamniformes though more recent analyses have suggested the 

Genus be placed within the Orectolobiformes, Carcharhiniformes or a 

Genus Palaeocarchariformes. 

- 27 - 

This study uses character state analysis of Lamniformes, 

Orectolobiformes and Carcharhiniformes in order to re-assess the 

systematic affinities of the Genus. 

A new species of Confuciusornis from the Early Cretaceous of 

northwestern China 

Zihui Zhang 1,2 , Chunling Gao 3 , Qingjin Meng 3 , Jinyuan Liu 3 , Lianhai 

Hou 2,4 and Guangmei Zheng 1 

1 

College of Life Sciences, Beijing Normal University, Beijing 100875, 

China 

2 

College of Life Sciences, Capital Normal University, Beijing 100037, 

China 

3 

Dalian Natural Museum, Dalian 116023, China 

4 

Institute of Vertebrate Paleontology and Paleoanthropology, 

Chinese Academy of Sciences, PO Box 643, Beijing 100044, China 

A new species of Confuciusornis —the oldest known beaked bird is 

erected based on a nearly complete fossil from the Early Cretaceous 

Yixian Formation of western Liaoning, northeast China. 

Confuciusornis feducciai is the largest and shows the highest ratio of 

the forelimb to the hindlimb among all known species of 

Confuciusornis, the skeletal qualitative autapomorphies including a 

V-shaped furcula, a rectangular deltopectoral crest, the absence of 

an oval foramen at the proximal end of the humerus, the very slender 

alular digit, a relatively much longer ischium which is 2/3 rds the length 

of the pubis. Comparison with other described species strongly 

suggests the new specimen a valid distinctive taxon, and provides 

further evidence for diversification in an Early Cretaceous avian 

genus. Anatomical features suggest an arboreal habit of the new bird. 

Some notes on the life style of confuciusornithids (Aves, 

Confuciusornithiformes, Confuciusornithidae) 

Andrei Zinoviev 

Department of Biology, Tver State University, Chaikovskogo pr., 70a, 

Tver, 170002, Russia 

Confuciusornithids is a group of Early Cretaceous birds from 

Liaoning Province of China. They possess a mosaic of primitive and 

derived characters, making them extremely important for 

understanding the evolution and early radiation of birds. By analyzing 

peculiarities of confuciusornithids’ morphology (skeleton, plumage, 

horny sheaths of beak and claws) and taphonomy, we propose the 

most plausible reconstruction of their life style. Being strikingly similar 

in body outlines to modern tropic birds (Phaethontidae) these 

creatures fed on fish, catching it on the wing from the surface layer of 

freshwater reservoirs. They rested (and, probably, nested) in 

canopies of adjacent abundant tropical vegetation. 

Confuciusornithids were not swimming birds like gulls and ducks; 

neither had they perched like modern passerines. Wing digit II, 

devoid of alula and equipped with the large sharp claw, helped 

Confuciusornis to climb in canopies in the way similar to that of 

hoatzin’s chick. Unable to fly from the ground, Confuciusornis must 

have climbed tree trunks to get airborne. In doing so it helped itself 

with digit IV, free of primaries and furnished with large hooked claw. 

A couple of remarkable elongated rectrices, developed in some 

specimens of Confuciusornis, most likely served for sexual display.


S8 - Biodiversity and ecology of Protists S9 – Genomics and cell biology of Protists 

A global perspective of the biodiversity of heterotrophic 

nanoflagellates as the most abundant heterotrophic eukaryotes 

Harmut Arndt 1* , Frank Nitsche 1 , Frank Scheckenbach 1 and Claudia 

Wylezich 1,2 

1 Department of General Ecology, Zoological Institute, University of 

Cologne, D-50968 Koeln (Cologne), Germany 

2 Institute for Baltic Sea Research, D-18119 Rostock, Germany 

Protist diversity is still controversially discussed. One group of 

protistologists assumes a relatively low number of protistan species 

compared to their low size due to assumed high rates of dispersal 

and low speciation rates. Another group of protistologists assumes 

high protistan diversity similar to other groups of eukaryotes claiming 

a high number of cryptic species behind each nominal species. We 

will summarise investigations of heterotrophic nanoflagellates 

regarding their diversity and distribution patterns. We will include 

samples from the largest, though seldom investigated parts of the 

biosphere, deep sea, groundwater and polar regions. There seems 

to exist biogeographies in nanoflagellates. The endemic and 

cosmopolitan distribution of certain taxa will be discussed. Regarding 

the knowledge of nanoflagellate biodiversity, we are still at the 

beginning and we will point to several problems associated with a 

current estimate of biodiversity patterns. According to our recent 

estimates, protists as the most abundant eukaryotic key players in 

most ecosystems and as the evolutionary oldest and 

phylogenetically most diverse eukaryotes should account for a large 

proportion of Earth’s eukaryotic diversity. The knowledge of 

biogeographic characteristics of nanoprotists is also important from 

the viewpoint of overall biological theory. Are small organisms 

different from large organisms? At least regarding their importance 

for the matter transfer in both aquatic and terrestrial ecosystems 

protists are much more important than larger eukaryotes. The 

knowledge of diversity patterns of nanoprotists seems to be 

fundamental for understanding the Earth’s biodiversity patterns and 

ecosystem functioning and may give us also a hint for a better 

understanding of evolutionary processes. 

The genome of Paramecium 

Jean Cohen 

Centre de Génétique Moléculaire, CNRS, 91198 Gif-sur-Yvette 

cedex. 

Ciliates such as Paramecium are the only unicellular eukaryotes 

known to separate germinal and somatic functions. Diploid but silent 

micronuclei transmit the genetic information to the next sexual 

generation. Polyploid macronuclei express the genetic information 

from a streamlined version of the genome but are replaced at each 

sexual generation through reproducible rearrangements of the 

zygotic genome involving elimination of repeated sequences, precise 

excision of unique-copy internal eliminated sequences (IES), and 

amplification of the cellular genes to high copy number. The 

macronuclear genome of Paramecium tetraurelia was recently 

sequenced by a shotgun approach, providing access to the gene 

repertoire. The 72-Mb assembly represents a consensus sequence 

for the somatic DNA which revealed the presence of nearly 40,000 

genes, most of them arising through at least three successive wholegenome 

duplications, which is produced after sexual. An overview of 

genomic and postgenomic data obtained with the Paramecium 

genome will be presented. 

Aury et al. 2006. Global trends of whole-genome duplications 

revealed by the ciliate Paramecium tetraurelia. Nature 444, 171-178. 

Duret et al. 2008. Analysis of sequence variability in the 

macronuclear DNA of Paramecium tetraurelia: a somatic view of the 

germ line. Genome Research, 18, 585-596. 

- 28 - 

Bacterial Endocytobionts Variety in Ciliophora 

Sergei I. Fokin 

Department of Invertebrate Zoology, St. Petersburg State University, 

199034, Russia 

Different bacteria, which could be considered facultative or, 

sometimes, permanent endocytobionts (Eb) can occupy ciliate’s cell. 

Till now over 230 ciliate species were recorded as hosts of different 

intracellular bacteria. However, approximate number of species of 

free-living ciliated protists is about 9000-20000. Thus the number of 

recorded Eb is, apparently, just a small part of the real biodiversity of 

this mostly undiscovered world. In the presentation is mentioned 

some new material concerning Eb variety, its categories and 

interaction with a host cell. Special attention is paid to the 

endocytobiosis between some ciliates and highly infectious bacteria 

Holospora or other alpha-proteobacteria, as well as to life cycles and 

strategies of Eb of different ciliates. From ecological point of view we 

can assume that association with Eb sometimes is useful for the host 

cell, especially in unstable environment like littoral marine zone or 

river’s estuary. However, quite low percentage of bacteria infection in 

most ciliate’ populations, revealed so far, appears to indicate that 

majority of Eb are either parasites or commensals, but not true 

symbionts. Dynamic of some bacterial infections, first of all infectious 

ones, in natural ciliate’ populations looks epidemic. In some years 

and places the number of infected populations increases and then 

infected cells become rare for a long time. There is a great need for a 

more extensive field-based research in this key sub-discipline of 

symbiosis. 

Rare ciliate species (Ciliophora, Protista) revealed from 

brackish water habitats with oxygen deficiency. 

Sergei I. Fokin 1,2 , Letizia Modeo 1 , Ilaria Andreoli 1 , Filippo Ferrantini 1 , 

Franco Verni 1 and Giulio Petroni 1 

1 

Department of Biology, University of Pisa, 56126, Italy 

2 Department of Invertebrate Zoology, St. Petersburg State 

University, 199034, Russia 

Sonderia pharyngea. Cell size 80-130 μm; all ventral ciliary rows 

extending from posterior end; cytopharynx is almost tubular, 

sometimes as long as a half of the body; there are few trichocysts. 

Found in salinity 5-12‰. 

Parablepharisma bacteriophora. Cell size 70-110 μm; has about 50 

membranelles on the left side of the large vestibulum; undulated 

membrane is very conspicuous – up to 30 μm long; 2 micronuclei; 

contractile vacuole closed to posterior end; According to SSrRNA 

sequence it clustered with Spirotrichea, very basal to the group. 

Found in salinity 22‰. All body surfaces of both mentioned species 

are covered with sulfur bacteria. 

Copemetopus sp. (subsalsus?). Cell size 150-300 μm; has about 50 

membranelles on the left side of large vestibulum (up to half of the 

body) and additional 8-12 units of very long cirri (up to 80 μm) 

decorate external apical-left part of the mouth area. Cilia tuff in 

anterial part of dorsal side; macronucleus has dumbbell shape; 

micronuclei are numerous (9-13). According to SSrRNA sequence it 

is clustered with Spirotrichea, very basal to the group. Found in 

salinity 5-8‰. 

All mentioned rare species were isolated in Italy. Probably previous 

single finds of the cilates connected with insufficient investigation of 

the habitats.

ICZ2008 – Abstracts S8-9 

From cows, cockroaches, and tiny protozoa: symbiosis and the 

evolution of life 

Johannes H.P. Hackstein 

IWWR, Fac. Science, Radboud University Nijmegen, NL-6525 ED 

Nijmegen, The Netherlands 

Evolution has been characterized by Darwinists as a 4 milliard’s 

years war - ignoring that nearly all life on earth is somehow 

integrated into an extremely complex web of symbiotic associations. 

Man, for example, hosts much more bacteria in his gastrointestinal 

tract than he has neuronal cells in his brain. Molecular approaches 

are beginning to unravel the incredible complexity and the functions 

of these symbionts that make man a “super-organism”. Studying a 

cow, we learn that the rumen of a single cow hosts more than 10 16 

bacteria, 10 – 100 milliards of ciliated protozoa and 100 millions of 

anaerobic fungi. 

The evolution of ruminants started some 40-60 million years ago, 

and the obvious success of these animals would be impossible 

without their symbionts. Other examples are found among the 

arthropods: cockroaches and termites host protozoa, bacteria and 

methanogenic archaea in their guts. The association between 

anaerobic protozoa and methanogenic (endo)symbionts potentially 

exemplifies the origin of the eukaryotic cell as a fusion product of a 

hydrogen-producing and a hydrogen-consuming cell. Certain 

anaerobic protozoa host intracellular methanogens because they 

possess “hydrogenosomes”, organelles that provide the 

endosymbionts with hydrogen for methane formation. While 

genomics have proven the endosymbiotic origin of the mitochondria 

(and plastids), the origins of “hydrogenosomes” and “mitosomes” 

remained an enigma until recently. It was not until our discovery of a 

“missing link” between hydrogenosomes and mitochondria that 

hydrogenosomes (and mitosomes) could be identified as a kind of 

mitochondria that – by evolutionary tinkering (bricolage) – adapted to 

life under anaerobic conditions. 

Paramecium actin cytoskeleton as a key player in Holospora 

infection 

Elena V. Sabaneyeva 1 , Konstantin A. Benken 1 , Maria E. Derkacheva 2 , 

Sergei I. Fokin 3 and Ilya N. Skovorodkin 2,4 

1 

Department of Cytology and Histology, St. Petersburg State 

University, St. Petersburg, Russia. 

2 

Institute of Cytology, Russian Academy of Sciences, St. Petersburg, 

Russia. 

3 

Department of Invertebrate Zoology, St. Petersburg State University, 

St. Petersburg, Russia 

4 

Department of Medical Biochemistry and Molecular Biology, 

Biocenter Oulu, Laboratory of Developmental Biology, University of 

Oulu, Oulu, Finland 

Holospora obtusa (Alphaproteobacteria) is an obligate 

endonucleobiont inhabiting the macronucleus of the ciliate 

Paramecium caudatum. The bacterium is ingested by the host in the 

course of phagocytosis but it quickly escapes from the digestive 

vacuole and reaches the target nucleus. The mechanism of bacterial 

transportation in the host cytoplasm has been enigmatic so far. An 

actin-based mechanism of transportation has been proposed for H. 

obtusa on the basis of electron microscopy data (Görtz, Wiemann, 

1989). The present study was undertaken to verify this hypothesis. 

Immunocytochemical investigation of the early stages of infection 

using antibodies against paramecium actin 1-1 and experimental 

infection of GFP-actin transfected paramecia showed that host actin 

contributes to the bacterial escape from the phagosome. The 

involvement of actin microfilaments in Holospora invasion was further 

supported by cytochalasin D treatment which resulted in a significant 

decrease in the rate of nuclear infection. However, antibody labeling 

demonstrated that the trails left by H. obtusa in the paramecium 

cytoplasm contained an actin-related protein (Arp3), but not actin 1-1 

itself. This observation suggests a rapid disassembly of F-actin and 

its quick loss from the trail, which might be due to unconventional 

character of actin cytoskeleton in Paramecium. 

Involvement of actin depolymerization factor cofilin in this process is 

under investigation. The data obtained in this study evidence for 

participation of actin in trafficking H. obtusa in the paramecium 

cytoplasm. 

Görtz, H.-D., Wiemann M. 1989 Eur J Protistol 24: 101-109. 

- 29 - 

Ancient soil protozoa isolated from eastern arctic permafrost 

Anastassia Shatilovich, Lubov Shmakova, Alexander Mylnikov and 

David Gilichinsky 

Institutskaya, 2, 142 290, Pushchino, Moscow region, Russia 

Permafrost is the unique environment which is capable to protect 

microorganisms for long-term preservation. The purpose of this study 

is to detect viable forms of protozoa in permanently frozen sediments 

of late Pleistocene and Holocene age, isolate and characterize these 

organisms. A total of 200 samples of Eastern Arctic permafrost and 

buried soils (including burrows of fossil rodent) were screened for 

presence of viable protozoa, while within 28 probes the 

representatives of all main groups, namely, naked amoebae, 

heterotrophic flagellates and ciliates were detected to the 20 m depth. 

Forty-three species and forms of ancient protozoa from fifteen supergeneric 

taxa were found in the investigated sediments. The longevity 

of protozoans cryoconservation corresponds to the permafrost age. 

The organisms found at the upper permafrost boundary are not older 

than a few hundred years, and single, the most ancient finding is 

dated to the middle Pleistocene (200,000-300,000 years). The 

general tendency of increasing the number and diversity of protozoa 

within buried soils and burrows of fossil rodent could be explained by 

more favorable conditions of cryoconservation in rich plant debris, 

and a relatively rich initial fauna. All observed organisms are 

common representatives of soil protistofauna and most of selected 

species are able to produce cysts. 

New approaches to analysis of energetics in plankton 

populations 

Télesphore Sime-Ngando 

LMGE, Laboratoire ‘Microorganismes: Génome et Environnement’, 

UMR CNRS 6023, Université Blaise Pascal, Clermont-Ferrand II, 

63177 Aubière Cedex, France. 

New approaches have been designed to unravel the functions of 

small heterotrophic eukaryotic microorganisms in pelagic 

ecosystems. In these systems, small heterotrophic eukaryotes, often 

known as heterotrophic flagellates (HF), are currently considered to 

include keystone groups of microorganisms in microbial loops and 

food webs which play significant roles in the transfer of prokaryotic 

carbon to higher trophic levels. Two decades ago small HF were 

regarded as a ‘homogenous’ functional group in the plankton 

including eukaryotic, non-photosynthetic and motile microorganisms, 

with similar morphological, physiological and ecological 

characteristics and almost exclusively composed of bacterivorous 

organisms. Recent investigations providing extensive data on 

sequences of DNA from ribosomal genes has changed the 

classification of these microorganisms and increased our knowledge 

of the taxonomic diversity within microbial communities. However, 

we are not much farther ahead today than we were two decades ago 

in our overall understanding of the ecological functions of these 

microorganisms. In reality, HF is a heterogeneous polyphyletic 

grouping of microorganisms, including a large number of genetically 

unrelated organisms from different phyla with different ecological 

functions. A new understanding of the diverse functions from HF in 

aquatic ecosystems is of paramount importance in aquatic 

environmental microbiology.

S8-9 ICZ2008 - Abstracts 

Biodiversity and ecology of freshwater ciliates 

Thomas Weisse 

Institute for Limnology of the Austrian Academy of Sciences, 

Mondseestr. 9, Mondsee, A-5310, Austria 

Ciliates are abundant, diverse and quantitatively important players in 

freshwater food webs. Their quantitative significance as consumers 

of protist and bacterial production, and food for larger metazoans 

such as Daphnia and copepods was recognized mainly during the 

1980s and early 1990s. Biogeography and biodiversity of ciliates has 

been controversially discussed during the past decade. The 

speculation that ciliates and other free-living protists are 

cosmopolitans, found everywhere where a suitable habitat exists, 

without any geographic isolation provoked intensive field and 

laboratory studies to demonstrate that ciliates do have a 

biogeography. Detailed taxonomic research by a few specialists 

revealed that, while many species are globally dispersed, endemics 

are also common among soil and freshwater ciliates. With the advent 

of molecular techniques to identify and classify ciliates based upon 

their genes it became obvious that ciliate diversity is considerably 

larger than derived from morphology-based alpha-taxonomy. 

Similarly, ecophysiological laboratory experiments revealed large 

inter- and even intraspecific differences. The response to food, 

temperature, pH and predators has been identified as the major 

environmental factors controlling the occurrence and distribution of 

freshwater ciliates. It now appears that ecotypes, adapted to a 

particular habitat, are common among widespread ciliates. 

Measurements of the actual rates of dispersal and the frequency of 

sexual reproduction in the field are currently the major challenges for 

a better understanding of the biodiversity of freshwater ciliates. 

- 30 -


S10 - Sex differentiation in vertebrates 

Sex-specific differences of the skeleton in North African gazelles 

(Mammalia, Artiodactyla, Bovidae) 

Maria Andrés 2 , Gema Maria Alcalde 1 , Beatriz Azanza 2,1 , Jorge 

Morales 1 and Maria Teresa Alberdi 1 

1 Departamento de Paleobiología, Museo Nacional de Ciencias 

Naturales, Consejo Superior de Investigaciones Científicas, José 

Gutiérrez Abascal 2, 28006 Madrid, Spain 

2 Departamento de Ciencias de la Tierra, Facultad de Ciencias, 

50009, Universidad de Zaragoza, Zaragoza, Spain 

Ruminants display frequently sexual dimorphism in body size and 

also in horns and upper canines if they are present. Sex-specific 

differences in size and morphology of bones is explored in three 

captive-bred North African gazelle species (Nanger dama, Gazella 

dorcas and Gazella cuvieri) from the “Estación Experimental de 

Zonas Áridas” (Almería, Spain). Morphologically female skulls are 

characterized by a more asymmetric basioccipital with less prominent 

anterior tuberosities, thinner jugular processes, and less depressed 

supraorbital fosses that are also more separate from each other. 

Apart from the pelvis, sexual differences are detected in the 

morphology of atlas and axis. In females, the atlas presents less 

developed wings with parallel lateral edges and the axis has smaller 

and less divergent transverse apophysis and a flat superior edge of 

the spinous process. PCAs run on each of these element databases 

found that between 93-100 % of the total variance is explained by the 

three first principal components. PC1 reflects general size, whereas 

PC2 and PC3 reveal variations of shape. Males and females are 

arranged by the three first components confirming important sexual 

differences in size and morphology of skull, pelvis, atlas and axis. But 

the biggest species, N. dama, shows dimorphism in all skeletal 

elements, even in the palatal region of skull and in the teeth. Sexual 

arrangements were checked by means of Discriminant Analysis with 

a classification rate between 69-76% in teeth and between 71-100 % 

in skull, 99% in either atlas or axis, and 95% in pelvis. 

Goat as model for studying R-Spondin involvement in ovarian 

differentiation in Mammals 

Aurélie Auguste, Fatemeh Montazer-Torbati, Ayhan Kocer, Maëlle 

Pannetier, Lauriane Renault, Béatrice Mandon-Pépin, Corinne 

Cotinot and Eric Pailhoux 

INRA – UMR Biologie du Développement et Reproduction – 78350 

Jouy en Josas. 

In mammals, once the chromosomal sex XY or XX is set up at 

fertilization, the presence of SRY gene (Sex-determining Region of Y) 

will determine the fate of the gonad by initiating testis differentiation. 

In the female counterpart, key genes for ovarian differentiation have 

been isolated from genetics studies of female-to-male XX sexreversal: 

the PIS locus (Polled Intersex Syndrome) in goat, and 

recently RSPO1 in human. 

In goat, the pleiotropic PIS mutation is a 11.7 kb deletion 

encompassing any genes, but acting on the transcriptional regulation 

of at least 3 genes, PISRT1, PFOXic and FOXL2, distal from the 

deletion. Compare to the normal situation (PIS +/- and PIS +/+ ), the 

expression of these three genes is abolished in the ovaries of XX 

foetuses homozygous for the deletion (PIS -/- ). So, it seems that one 

of the PIS-regulated genes have an “anti-testis” effect. Among the 

three genes, only FOXL2 encodes a conserved protein belonging to 

the forkhead family making it a strong anti-testis candidate gene. 

As shown in human a R-Spondin gene (RSPO1) is involved in a 

female-to-male XX sex reversal. Consequently, R-Spondin 

expression patterns were studied in XX PIS -/- male. RSPO2 was 

detected with a female sex-dimorphique expression, and was absent 

in XX mutants. This leads us to assume that RSPO2 could be a 

direct or indirect target of FOXL2 in ovarian differentiation. 

To further confirm our presumption, different studies are under 

progress: (i) cellular localization of RSPO2 in ovary; (ii) if co-localized 

with FOXL2, studies of RSPO2 promoter. 

- 31 - 

Identification of new transcripts involved in ovine ovary 

development 

Adrienne Baillet 1,2,3 , Béatrice Mandon-Pépin 1,2,3 , Cédric Cabau 4 , 

Elodie Poumerol 1,2,3 , Eric Pailhoux 1,2,3 and Corinne Cotinot 1,2,3 

1 

INRA, UMR 1198 Biologie du développement et reproduction, F- 

78350 Jouy en Josas, France 

2 

ENVA, UMR 1198 Biologie du développement et reproduction, F- 

78350 Jouy en Josas, France 

3 

CNRS, FRE 2857, F-78350 Jouy en Josas, France 

4 

INRA, SIGENAE, UR83 Unité de Recherches Avicoles, 37380 

Nouzilly, France 

The aim of this study was to isolate by suppressive subtractive 

hybridization genes differentially expressed between the two main 

steps of ovarian development in sheep: onset of prophase I meiosis 

(55dpc) and follicle formation (82dpc). 

Two subtractive libraries (55/82; 82/55) were constructed and 7296 

clones were obtained. Among them, 6080 clones were sequenced 

and grouped into 2101 unique contigs using SIGENAE bioinformatics 

facilities. Then these contigs were compared with databases from 

different mammalian species (human, bovine and ovine) and 

annoted. In both libraries, 99% of contigs possessed an Unigene 

annotation and 1% were unknown (21/2101). 

The expression profile of all unknowns contigs were determined by 

RT PCR in fetal ovary, testis and in a pool of somatic tissue. Three of 

them showed an ovary specific expression that was confirmed by 

real time RT-PCR. Furthermore, we have found one gene never 

described in female ovary. This gene showed a high expression level 

during female meiosis I while its expression remained low in fetal 

testis and during the other stages of ovary development. 

Further studies such as 5’ and 3’RACE analysis to obtained fulllength 

transcripts and ISH to determine cellular localisation are 

currently in progress for these genes. Moreover, investigations of the 

folliculogenesis library are in under progress. 

In parallel, these 2101 contigs will be used to develop a custommade 

macroarray, dedicated to ovine ovary differentiation, in order to 

evaluate ovarian transcriptomes in different physiological and 

physiopathological conditions. 

Sex determination and sex chromosome evolution in the 

platyfish Xiphophorus maculatus 

Astrid Böhne 1 , Christina Schultheis 1 , Delphine Galiana-Arnoux 1 , 

Comelia Schmidt 2 , Qingchun Zhou 2 , Alexander Froschauer 2 , Yvonne 

Selz 2 , Catherine Ozouf-Costaz 3 , Béatrice Ségurens 4 , Arnaud 

Couloux 4 , Sylvie Bernard-Samain 4 , Stefan Chilmonczyk 5 , Frédéric 

Brunet 1 , Jean-François Baroiller (6) , Helena D’Cotta 6 , Julien Bobe 7 , 

Yann Guigen 7 , Manfred Schartl 2 and Jean-Nicolas Volff 1 

1 Ecole Normale Supérieure de Lyon, Institut de Génomique 

Fonctionnelle, Université de Lyon, France; 2 University of Würzburg, 

Physiologische Chemie I, Würzburg, Germany; 3 Muséum National 

d'Histoire Naturelle, Département Systématique et Evolution, Paris, 

France; 4 Genoscope, Evry, France; 5 INRA, Jouy en Josas, France; 

6 CIRAD, Montpellier, France; 7 INRA-SCRIBE, Rennes, France 

In contrast to the situation observed in mammals and birds, fish 

display an amazing diversity of sex determination systems. Almost 

nothing is known about the molecular and evolutionary basis of this 

phenomenon. Bacterial artificial chromosome (BAC) contigs 

including the sex-determining region of the platyfish Xiphophorus 

maculatus and covering 2.5-3.5 megabases of the X and Y 

chromosomes have been constructed and sequenced. Thirty-nine 

BAC clones have been sequenced so far to completion, leading to 

the identification of 63 sex chromosomal genes. Interestingly, eleven 

of these genes are also located on human sex chromosomes too, 

but no synteny was found with other fish gonosomes. Through 

comparison between X and Y sequences, the sex-determining region 

has been delimited. Gene candidates have been identified for all loci 

involved in pigmentation, cancer formation and sexual development 

that are linked to the master sex-determining gene of the platyfish. 

Numerous genes, including the melanocortin hormone receptor gene 

mc4r, are amplified in the sex-determining region of X. maculatus.


Gonad-specific genes were identified, including two genes with so far 

unknown function organized in tandem and exclusively expressed in 

oocytes. One gene, called swimy, is present on the Y but not on the 

X chromosome and represents an excellent candidate for the master 

sex-determining gene. This gene encodes a ubiquitously expressed 

protein, with different predicted domains involved in nucleic acid 

binding and protein-protein interactions. Interestingly, through the 

use of an alternative exon in spermatogonia, a testis-specific longer 

isoform is produced, with different domains involved in protein 

modification. 

The battle of sexes: patterning the gonad 

Blanche Capel, Danielle Maatouk, Yuna Kim, Leo DiNapoli and 

Lindsey Barske 

Department of Cell Biology, Duke University Medical Center, Durham, 

NC 27710 

Although genetic sex is established at fertilization, Alfred Jost 

demonstrated that the decision determining whether an embryo 

develops as male or female occurs later in gestation, in the gonad. 

The gonad arises as a uniquely bipotential primordium, 

indistinguishable in XX and XY embryos, and harboring the potential 

to develop as either a testis or an ovary. Studies suggest that cells in 

the early gonad are influenced by opposing signals that hold them in 

a bipotential state. In mammals, transient expression of the Y-linked 

gene Sry in a subset of cells triggers a global shift in this balance 

toward the testis fate. Although Sry normally acts as the genetic 

switch to initiate male development of the gonad, the sex 

determination pathway can be manipulated by alterations in the 

underlying antagonistic signaling pathways. For example, disruption 

of the Fgf signaling pathway leads to male to female sex reversal, 

while disruption of the Wnt/Rspo signaling pathway leads to partial 

female to male sex reversal. Recent evidence suggests that these 

opposing pathways may act by influencing the intracellular 

antagonism between SOX9 and ß-catenin. This work leads to the 

hypothesis that Sry is a mammalian invention that is superimposed 

on an underlying signaling network that may regulate sex 

determination across vertebrates in species where Sry is absent and 

sex is determined by other genetic elements or by environmental 

cues. 

Rspo1, an essential gene for ovarian differentiation in mammals 

Anne-Amandine Chassot 1 , Elodie Grégoire 1 , Giovanna Camerino 2 , 

Dirk de Rooij 3 , Andreas Schedl 1 and Marie-Christine Chaboissier 1 

1 INSERM, U636, F-06108 Nice, France; Université de Nice-Sophia 

Antipolis, Laboratoire de Génétique du Développement Normal et 

Pathologique, F-06108 Nice, France 

2 Dipartimento di Patologia Umana ed Ereditaria, Sezione di Biologia 

Generale e Genetica Medica, Universita di Pavia, Via Forlanini 14, 

27100 Pavia, Italy 

3 Center for Reproductive Medicine, Academic Medical Centre, 1105 

AZ Amsterdam, The Netherlands 

The sex of an individual is determined during development through 

the fate of the gonad. Gonad sex determination is controlled by a 

balance between two different pathways. While the expression of Sry 

and Sox9 is sufficient to induce the male developmental program, we 

have shown that the transcriptional function of β-catenin is activated 

in XX embryonic gonads, and antagonizes the male determining 

pathway. Rspo1 is an activator of the Wnt/β-catenin signalling 

pathway required for ovarian differentiation and ablation of Rspo1 

induces female-to-male sex reversal, similar to human XX patients 

with mutations in RSPO1. Rspo1 is not only involved in the 

differentiation of somatic cells, but is also required for normal germ 

cell differentiation and commitment to meiosis. These results 

demonstrate that Rspo1 is a crucial gene required for sex 

determination in mammals. 

- 32 - 

Sex differentiation of the urodele amphibian Pleurodeles waltl 

Dominique Chardard, Amand Chesnel, Hélène Dumond, Mathieu 

Gelhaye, Chia-I Ko, Sandra Kuntz, Angelina Wallacides and 

Stéphane Flament 

EA 3442 Aspects cellulaires et moléculaires de la reproduction et du 

développement, Faculté des Sciences. Université de Nancy 1. 

Boulevard des Aiguillettes. BP 239., 54506 Vandoeuvre-lès-Nancy 

Cedex. France 

In the sexually dimorphic urodele amphibian Pleurodeles waltl, sex 

determination obeys female heterogamety. Female (ZW) and male 

(ZZ) genotypes can be deduced from tail biopsy samples submitted 

to a biochemical analysis of the sex linked marker peptidase-1. The 

first histological changes in the differentiating gonad are observed at 

the end of larval life (stage 53): germ cells are localized in the cortex 

of the ovary but in the medulla of the testis. Meiosis entry occurs 

before metamorphosis in females (stage 54 + 2 months) whereas it 

occurs only two months post-metamorphosis in males. Experiments 

using organotypic cultures suggest that retinoic acid is involved in 

this event. The differentiation of the testis is very astonishing: the 

caudal extremity initiates spermatogenesis whereas the cephalic part 

remains in an undifferentiated status and contains primordial germ 

cells whose meiosis entry is delayed. Besides, the differentiation of 

the testis takes place lifelong and leads to several lobes named 

“multiple testis”. This model displays a few original features that can 

be helpful for the understanding of sex differentiation. First, when 

genetically female larvae (ZW) are reared at 32°C (instead of 20°C) 

during a thermo-sensitive period (stage 42 to stage 54), they develop 

as phenotypic fertile males. Several studies have demonstrated that 

estrogen synthesis is inhibited in case of temperature-induced sex 

reversal and have pointed out steroids as key factors in sex 

differentiation pathway. Second, parabiosis experiments can be 

performed. Ovarian development is impaired but testis differentiation 

is normal suggesting a role of other hormones, maybe AMH. 

Temperature-induced male differentiation in the Nile tilapia: 

gonad gene expression using female monosex populations & 

divergent thermo-sensitive lines 

Helena D’Cotta 1 , Elodie Pepey 1 , Stephane Wessels 2 , Srisupaph 

Poonlaphdecha 1 , Birgit Reinelt 2 , Gabriele Hörstgen-Schwark 2 and 

Jean-François Baroiller 1 

1 

CIRAD, Upr20, Dept. Persyst, Campus International de Baillarguet, 

F-34398 Montpellier, France 

2 

Institute of Animal Husbandry and Genetics, Albercht Thaer-Weg 3, 

G-37075 Göttingen, Germany 

Sex in the Nile tilapia, Oreochromis niloticus is governed by the 

interactions between a complex genetic sex determination system 

and the influence of temperature. High temperatures applied during a 

critical period of sex differentiation can induce masculinisation in 

some progenies originating from both domestic and wild stocks. 

Differences in thermo-sensitivity were observed between the wild 

populations and both paternal and maternal effects have been 

demonstrated, suggesting that it is a heritable trait. This has been 

confirmed by the development of divergent lines for thermosensitivity. 

Two thermo-sensitive divergent lines were selected with 

regard to their sex ratios showing high (>90% males) and low 

responses (only 54%) when treated to 36°C temperatures. At least 

one of the mechanisms by which temperature exerts its effects on 

sex differentiation is through a down-regulation of aromatase Cyp19. 

Expression profiles were analysed in monosex XX populations 

treated to 27°C and 36°C temperatures showing that temperature 

up-regulates Sox9s, Amh, IGFs, Dax1 and down-regulates other 

genes such as aromatase Cyp19-1a. Variation in these gene 

expression levels was associated with the percentage of males 

obtained by temperature treatments. The divergent thermosensitive 

lines have been used to complement our study done on the monosex 

populations. Expression of Cyp19-1a was also analysed in these two 

divergent lines.


foxl2 and sox9 show in situ sex reversal in patients with various 

forms of disorders of sex development 

Marc Fellous 1 , Remko Hersmus 2 , Reiner Veitia 1 , Francis Jaubert 1 

and Leendert H.J. Looijenga 2 

1 

University Paris 7, Institut Cochin, Inserm 567, 24 Rue du Fg St 

Jacques, 75014 Paris, France 

2 

Department of Pathology, Erasmus MC - University Medical Center 

Rotterdam, Josephine Nefkens Institute, Daniel den Hoed Cancer 

Center, Rotterdam, The Netherlands 

Under normal physiological conditions, the gonadal sex formation of 

testicular or ovarian tissue, is highly separated in mammals, and 

determined by chromosomal sex constitution. 

In normal gonads SOX9 protein expression was found to be 

restricted to the development and maintenance of Sertoli cells. In 

contrast, FOXL2 was only found in granulosa cells, as well as 

stromal cells in early development. 

However, there are a number of pathological exceptions to this rule: 

both testicular and ovarian tissue can be formed, either in a single 

gonad, or in two different gonads. This pathological condition is 

classified as ovo-testicular Disorder of Sex Development (DSD) 

This study for the first time investigates the presence of both SOX9 

and FOXL2 in gonads of patients with various forms of DSD. This 

study demonstrates the novel finding that in DSD patients, the 

formation of either ovarian or testicular development can only be 

visualized using immunohistochemistry for FOXL2 and SOX9, 

respectively. 

The results demonstrate that SOX9 is a highly informative marker for 

testicular development and FOXL2 for ovarian development, either 

present in isolated or mixed constitution. Although SOX9 and FOXL2 

could be present within a single histological context, it was never 

found to be expressed at high levels within the same cell. These 

observations demonstrate for the first time the additional value of 

immunohistochemistry for SOX9 and FOXL2, compared to 

morphology alone, to diagnose the presence of either ovarian or 

testicular differentiation or both, especially in patients with DSD. 

Gender-bending chemicals and the mammalian fetal gonad 

Paul Fowler 

IMS, Foresterhill, AB25 2ZD, Aberdeen, UK 

Most mammals adhere to a general pattern of gonadal development 

and then subsequent sexual differentiation along the lines outline by 

Alfred Jost over 40 years ago. For development of a reproductively 

competent male, establishment of testes leads to androgen action, 

in-utero and post-natally, which is critical for the development of a 

urogenital system based on Wolffian ducts. In contr! ast, females 

develop ovaries as a result of a different cascade of genes and 

establish a urogenital system based on Mullerian ducts. It is well 

recognised from experimental studies that excess androgen 

masculinises the female fetus while excess oestrogen interferes with 

male fetal masculinisation, in both instances impairing fertility. While 

there are mammals, including the spotted hyena and several mole 

species, which deviate from this basic pathway, in the majority of 

mammals disturbance of endocrine signalling adversely affects 

reproductive development. However, over the last 20 years there has 

been increasing concern about the potential detrimental effects of 

environmental chemicals, including heavy metals and endocrinedisrupting 

compounds, on reproductive development and health in 

humans, domestic species and wildlife. These chemicals have 

complex mechanisms of action and additive effects in complex 

mixtures and at low doses. The levels of exposure that developing 

mam! mals receive and the effects of the exposures remain poorly 

understood. In addition the data concerning the effects of exposure 

are sometimes contradictory. However, there is sufficient evidence 

for so-called “gender-bending” actions of fetal exposure to these 

environmental chemicals. In this presentation the basic processes in 

male and female reproductive development and the evidence for 

their disturbance by exposure to environmental chemicals will be 

discussed. 

Collaborators: Sharpe, R.M., Evans, N.P., Rhind, S.M., Cotinot, C., 

Fischer, B., Pocar, P., Sinclair, K., Lea, R.G., O’Shaughnessy, P.J. 

Funding: Wellcome Trust, European Commission (FW7), Grampian 

Endowments, Chief Scientist Office, Scotland. 

- 33 - 

The cres/testatin subgroup, a reproductive tract specific 

subgroup of genes within the cystatin family 2 protease 

inhibitors in sexual development 

Jessica Frygelius, Anna Wedell and Virpi Töhönen 

CMM L8:02, SE-171 76, Stockholm, Sweden 

Testis differentiation is initiated with the expression of the Y-linked 

factor Sry in XY gonads. Factors, such as Sox9, Fgf9 and Dhh, have 

been identified downstream of Sry and proven to cause gonad 

development failures or sex reversals in humans or in mice models 

when disrupted. Characterization of novel genes will help to clarify 

basic mechanisms behind gonad development as well as facilitate 

diagnostics of gonad dysgenes! is. Testatin was previously isolated 

by our group in a screen searching for novel genes expressed in 

early mouse sex differentiation. Testatin is specifically up regulated 

in the developing testis just after expression of Sry. Testatin belongs 

to the Cres/testatin subgroup of cystatin family 2 protease inhibitors 

that show a reproductive tract restricted expression (testis, 

epididymis, ovary, pituitary) in contrast to the broad expression 

profile of family 2 cystatins, which implies a specialized function 

within the reproductive tract. We generated a testatin knockout mice 

to evaluate the role of testatin in male sexual development. We 

observed normal fetal testis development and fertility in male testatin 

knockout animals (1). An explanation could be redundancy between 

the subgroup members. We evaluated the expression profile of the 

Cres/testatin subgroup in fetal testis with real-time PCR and in situ 

hybridization and we show that three of the subgroup members 

together with testatin (Cres, cystatin SC, Cystatin TE-1) are 

expressed in mouse fetal testis (2). In conclusion, future generation 

of conditional triple knockout mice would further evaluate the role of 

the Cres/testatin subgroup in male sexual development. 

Gene expression profiling during gonadal differentiation in 

chicken 

Marina Govoroun, Gwenn-Aël Carré, Isabelle Couty and Jean-Pierre 

Brillard 

SRA INRA-Centre de Tours, 37380 Nouzilly, France 

In birds the sex is determined by ZZ/ZW chromosome system in 

which the female is heterogametic. Several genes involved in 

mammalian gonadal differentiation have been identified in the 

chicken, and their expression patterns investigated but their 

function(s) as well as the molecular pathways in which they are 

involved, are still poorly understood. In the light of current knowledge, 

this study aimed at identifying new actors of chicken gonadal 

differentiation in order to improve our understanding of the female 

and male molecular pathways in birds. Based on real time RT-PCR, 

we studied the expression profiles of over 100 candidate genes 

during gonadal differentiation. Our set of candidates included ligands, 

receptors, signaling molecules, enzymes and transcription factors, 

known for theirs roles in sex differentiation, reproduction and 

embryogenesis in other species. The hierarchical clustering of genes 

based on the similarity of theirs temporal expression patterns 

allowed the statistical identification of gene clusters with sex or/and 

stage, or/and side specific signatures. For instance, the analysis of 

the distribution of genes in clusters suggested a differential 

involvement of the members of the TGFβ system in chicken testicular 

and ovarian differentiation along with, for some of them, their 

implication in the mechanisms underlying gonadal left-right 

asymmetry which features the ovarian development in birds. In 

addition, we also identified new Z-linked actors of chicken testicular 

differentiation. To conclude, the findings of this study gave new 

insight on the molecular mechanisms of chicken gonadal 

differentiation and provided good candidate genes for further 

functional analysis.


New mechanisms involved in meiosis prevention by fetal mouse 

testes 

Marie J. Guerquin, Clotilde Duquenne, Jean-Baptiste Lahaye, René 

Habert and Gabriel Livera 

Laboratory of Differentiation and Radiobiology of the Gonads, 

INSERM-U566, CEA/DSV/DRR/SEGG/LDRG, Univ. Paris 7 – Denis 

Diderot, F-92265 Fontenay aux Roses, France 

Fetal testes and ovaries differentiate from morphologically identical, 

bipotential gonads. In mice, the proliferative germ cells (GC) remain 

very similar in male and female gonads until about 13.5 days postconception 

(dpc). At this age, female GC initiate meiosis whereas in 

the testis, GC undergo mitotic arrest with all GC having entered the 

quiescence phase by 15.5 dpc. Recent findings indicate that retinoic 

acid (RA) is the key factor in committing GC toward the female 

pathway as it induces meiosis in mouse fetal germ cells. GC in the 

fetal testis are protected against the effects of RA by CYP26b1, a 

male-specific enzyme that degrades RA. In this study, we evidenced 

distinct testicular pathway involved in the prevention of the fetal 

meiosis. Using a co-culture model in which an undifferentiated XX 

gonad is cultured next to a fetal or neonatal testis, we demonstrated 

that the testis prevented the meiosis initiation in the XX gonad. This 

testicular effect was function of the stage of the testis and was not 

correlated with the expression of Cyp26b1. Moreover, addition in the 

medium of RA or ketoconazole, an inhibitor of Cyp26b1, did not 

prevent the testicular meiotic inhibitory effect on the GC of the cocultured 

ovary. We evidenced that this testicular effect was due to 

secreted factor(s) as conditioned medium of fetal testes also inhibit 

meiosis in the XX gonad. Lastly, molecular weight cut-off 

experiments indicated that the factor’s weight is higher than 10 kDa. 

In conclusion, our results demonstrate that diffusible testicular factor 

specifically produced during fetal and neonatal lifes have the 

potentiality to prevent meiosis independently of the activity of 

Cyp26b1. 

Correlation between ovarian steroidogenesis and β-endorphin 

in the Lizard Uromastyx acanthinura: Immunohistochemical 

approach. 

Sadlia Hammouche 1 , Thérèse Gernigon 1 and Jean-Marie Exbrayat 2 

1 Laboratoire de Recherche en Zones Arides, Faculté des Sciences 

Biologiques, Université des Sciences et de la Technologie de 

Houari Boumediene, PB 39 El Alia, Bab Ezzouar, Alger, Algeria 

2 Laboratoire de Biologie Générale, Ecole Pratique des Hautes 

Etudes, Université Catholique, 25 rue du Plat, 69288 Lyon cedex 02, 

France. 

In Mammals, opioid peptides are involved in various physiological 

process including reproductive function. The major site of 

biosynthesis are hypothalamus, solitary bundle nucleus and 

hypophysis intermediary lobe. β-endorphin, one of opioid peptides 

was also synthesized in the ovary. In Uromastyx acanthinura, the 

localization of this peptide and sex steroid was investigated by the 

immunohistochemical approach. The β-endorphin is strongly 

distributed in the granulosa cells and oocyte cytoplasm of the 

previtellogenic follicles of sexually quiescent lizards (winter) when 

steroidogenesis is interrupted. In spring, the signal becomes low, or 

event absent, in the vitellogenic and previtellogenic follicules. The 

granulosa cells of the previtellogenic ones show an important 

synthesis of 17β estradiol. Females that do not undergo 

vitellogenesis in spring show the same profile as winter quiescent 

females. These findings represent the first evidence of the presence 

of β-endorphin in the ovary of Uromastyx acanthinura. The seasonal 

variations observed in the reproductive cycle suggest that this opioid 

peptide is involved in the modulation of seasonal steroidogenesis. 

- 34 - 

Adrenal cortex contents of androstenedione in rabbit 

(Orynctolagus cuniculis) 

Faroudja Kandsi and Fatima Hadj-Bekkouche 

FSB/USTHB, 16111, Alger, Algeria 

The exploration of the adrenal androgen activity with the aim of 

knowing the endocrine physiology of the domestic rabbit in local 

populations of Algeria had been carried. An histology and a 

morphometric measurements of the glands and an evaluation of its 

contents of androstene-dione hormone are realized. Male rabbits of 

37 days (n=14), 60 days (n=5) and 6 months old (n=2), from the 

experimental farm (ITELV) are sacrificed. Adrenal glands are quickly 

taken, streamlined, the left (awkward) gland is fixed in Bouin 

Hollande for the histological study and the right adrenal gland are 

crushed in the phosphate buffer pH 7.4 for the dosage of the 

androstenedione by RIA kit. The histo- morphometric study shows 

that the zonation of the adrenal cerebral cortex is net in the animals 

of 37 days, the thicknesses of glomerulosa, fasciculata and 

reticularis zones expressed in percents are respectively 9.36 %, 

62.99 %, 27.63 % of the total thickness of the cerebral cortex, and 

these values are not different from those observed in rabbits of 60 

days and 6 months. The adrenal content of androstene-dione in 100 

mg of adrenal weight are 2.90 ng at 37 days, 4.54 ng at 60 days and 

1.34 ng in 6 month-old rabbits. Even though, in the rabbit, the 

adrenal cerebral cortex, according to its production of androstenedione 

(which is the metabolite of the DHEA and a potential direct 

precursor of the testosterone) might participate in the installation of 

the puberty. 

Gonadal development of freshwater turtle (Malayemys 

macrocephala) embryos exposed to environmentally relevant 

doses of Cadmium 

Noppadon Kitana 1 , Sarun Keithmaleesatti 2,3 and Kumthorn 

Thirakhupt 1 

1 Department of Biology, Faculty of Science, Chulalongkorn 

University, Bangkok 10330, Thailand 

2 Inter-department of Environmental Science, Graduate School, 

Chulalongkorn University, Bangkok 10330, Thailand 

3 Department of Environmental Sciences, Faculty of Science, Khon 

Kaen University, Khon Kaen 40002, Thailand 

Cadmium contaminated sediment plumes have formed on the Mae 

Tao River Basin, Tak Province, Thailand, possibly as a result of zinc 

mining activities. We are using the freshwater turtle (Malayemys 

macrocephala) as a sentinel to monitor the reproductive effects of 

exposure and, by inference, the potential for human health impacts. 

Since contaminant may affect turtle by exposing through eggshell as 

well as maternal transfer via yolk, we thus examined the effect of 

cadmium on turtle embryos. Freshly laid eggs of M. macrocephala 

were collected from a reference site with no history of cadmium 

contamination. Representative eggs were analyzed for cadmium 

contamination by an ICP/ES to verify that the cadmium content in 

egg yolk is non-detectable. The effect of cadmium was determined 

using eggs subjected to in ovo exposure to cadmium chloride. The 

doses used in this study (0, 3, 30 and 300 μg of total cadmium/g egg 

weight) were based on concentration in soil at Mae Tao area. Eggs 

were kept in an incubator at a constant temperature that yields 1:1 

sex ratio until hatch. It was found that low dose of cadmium 

significantly prolonged the hatching time compared to that of the 

control. However, the hatching success and hatching weight are not 

significantly different among groups. Since cadmium may interfere 

with processes of gonadal development, results on sex ratio and 

gonadal development of these hatchlings will be presented. The 

potential of the environmentally relevant dose of cadmium on turtle 

development and its implication for sentinel system will be discussed.


Modulation and immunohistochemical localization of P450 

aromatase on Algerian Psammomys 

Rachid Menad and Thérèse Gernigon-Spychalowicz 

Laboratoire de Recherche sur les Zones Arides (LRZA), Equipe 

Reproduction des petits Vertébrés, Faculté des Sciences 

Biologiques (FSB), Université des Sciences et de la Technologie 

Houari Boumediene (USTHB), BP 32, El Alia 16111 Alger, Algérie 

Epididymal function is androgen dependent, but recent evidence 

indicates that estrogens are also important. They have a major 

importance in fertility. Estrogens receptors invalidation in the mouse 

induces an abnormal epididymal phenotype and infertility. The aim of 

this study is research of aromatase on sand rat epididymis and to 

highlight its modulation during castration, castration then treatment 

and efferent ducts ligation. Three groups were constituted: castrated 

groups for one month, castrated groups during 36 days then treated 

during 15 days by testosterone, and animals having undergone 

efferent ducts ligation. The organs were taken in Hollande Bouin for 

immunohistochemistry. In order to locate P450 aromatase 

responsible for the estrogen production, indirect method was applied 

by using anti-mouse aromatase antibodies against human 

aromatase. The research of P450 aromatase on proximal epididymis 

does not show positive reactions on lumen, epithelium and 

conjunctive tissue in the control. Castration, castration then 

testosterone treatment and efferent ducts ligation does not show 

positive reaction on all compartments of proximal epididymis. In 

distal epididymis, P450 aromatase shows proximal cauda epididymis 

localization. This staining increases and becomes more intense 

towards the distal cauda principal cells cytoplasm and not the clear 

cells. Castration experiments show enzyme disappearance and 

testosterone treatment induces its reappearance. In the animals 

having undergone efferent ducts ligation, the positive reaction 

persists only in distal part of cauda epididymis; it always remains in 

the cytoplasm of the principal cells. The absence of P450 aromatase 

in proximal epididymis excludes its expression in this segment or its 

testicular origin. 

Its immunolocalization on principal cells cytoplasm of the distal 

epididymis and its androgen-dependence lets him to allot a 

modulating role of T/E balance other than T/DHT balance and does 

not exclude estrogens impact on epididymal function. 

Development of a goat-specific anti-SRY antibody: preliminary 

results on goat SRY protein expression 

Fatemeh Montazer-Torbati, Aurélie Auguste, Maëlle Pannetier, 

Ayhan Kocer, Lauriane Renault, Béatrice Mandon-Pépin, Corinne 

Cotinot and Eric Pailhoux 


Jouy en Josas, France 

In mammals, the SRY gene (Sex-determining Region of Y 

chromosome) is the master regulator of male sex determination. 

Despite 15 years of studies, the molecular and cellular mechanisms 

operating downstream SRY remain undefined. The SRY gene is not 

well-conserved among mammals both in terms of protein structure 

and gene expression. In mouse, Sry is first expressed around 10.5 

days post-coïtum (dpc), reaches a peak of expression at 11.5 dpc, 

and is extinguished shortly after 12.5 dpc. In sheep, SRY expression 

is first detected in male gonads at 23 dpc, remains highly expressed 

during 18 days between 27 and 44 dpc and decreases but is not 

totally absent from 49 dpc until a few days after birth. In the postpartum 

sheep testis, SRY is expressed again from 12 dpp until 

adulthood. By contrast in goat, SRY mRNAs are detectable since the 

earliest studied stage (25 dpc) until adulthood. In order to determine 

the cellular and sub-cellular localization of SRY protein at different 

developmental stages in goat, we have developed polyclonal 

antibodies against the C-terminal part of goat SRY protein, excluding 

the HMG-box. Our objective is to confirm the specificity and 

effectiveness of our antibodies against goat SRY protein and to 

compare the developmental expression profiles of SRY proteins 

between goat and sheep. 

- 35 - 

Impact of phtalates on human fetal testis in vitro 

Vincent Muczynski 1 , Romain Lambrot 1 , Charlotte Lécureuil 1 , 

Delphine Moison 1 , Hervé Coffigny 1 , Catherine Pairault 1 , Gaelle 

Angenard 1 , René Frydman 2 , René Habert 1 and Virginie Rouiller- 

Fabre 1 

1 

LDRG, Unit of Gametogenesis and Genotoxicity, INSERM U566, 

University Paris 7; CEA, 18 route du panorama, 92265 Fontenay aux 

Roses, France 

2 

Service of Gynecology and Obstetrics, Antoine Béclère Hospital, 

Clamart, France 

During the last decades, a decrease in sperm production and an 

increase in genitalia abnormalities have been described in human. 

They are suspected to belong to the same Testicular Dysgenesy 

Syndrome. This syndrome may result from a defect in testis 

development during fetal life, due to an increasing exposure to 

endocrine disruptors (Sharpe et Skakkebaek, 1993), strongly 

widespread in the environment, among which we found the 

phthalates. One of the most important phthalates is the di(2ethylhexyl)-phthalate 

(DEHP) which is metabolized in its major active 

compound: MEHP. Exposure of rodents to phthalates impairs testis 

functions. Moreover, recent clinical investigations have reinforced 

this hypothesis (Swan et al. 2005; Main et al. 2006). 

We have studied the effects of MEHP, with doses from 10 -6 M to 10 - 

4 M, on the development of human fetal testis. The testes have been 

obtained form legal pregnancy abortion (7 to 12 weeks). This study 

has been realized in vitro in an organotypic culture system, 

previously developed in our laboratory (Lambrot et al. 2006). 

With the higher dose, we show for the first time that MEHP have a 

negative effect on the number of human gonocytes by increasing 

their apoptosis without any changes in their proliferation rate. But 

contrary to the rat and whatever the dose used, MEHP does not 

affect testosterone production of the human fetal testis. 

Sharpe, R. M., et al. (1993). Lancet 341(8857): 1392-5. 

Swan, S. H., et al. (2005). Environ Health Perspect 113(8): 1056-61. 

Main, K. M., et al. (2006). Environ Health Perspect 114(2): 270-6. 

Lambrot R, et al. (2006) J Clin Endocrinol Metabolism, 91: 2696-703. 

Sex differentiation in mammals: what about polled goats? 

Ayhan Kocer, Maëlle Pannetier, Lauriane Renault and Eric Pailhoux 


Jouy en Josas, France 

In goats, the PIS (Polled Intersex Syndrome) mutation is responsible 

for both the absence of horns in males and females and sex-reversal 

affecting exclusively XX individuals. The mode of inheritance is 

dominant for polled trait and recessive for sex-reversal. In XX PIS -/- 

mutants, expression of testis-specific genes is observed very 

precociously during gonad development. Nevertheless, a delay of 4- 

5 days is observed in comparison with normal testis differentiation in 

XY males. By positional cloning, we have demonstrated that the PIS 

mutation is a 11.7-kb regulatory-deletion affecting the transcriptional 

expression of 3 genes, PISRT1, PFOXic and FOXL2 which should 

act synergistically to promote ovarian differentiation. The 

transcriptional extinction of these 3 genes leads, very early, to testisformation 

in XX homozygous PIS -/- mutants. 

Since the discovery of this mutation in 2001, progresses have been 

made in order to understand the molecular functioning of this 

complex locus and the role of the different PIS-regulated genes. Our 

current understanding of this locus will be presented in regards with 

our main results obtained these last years, demonstrating that for 

some aspects goat gonad differentiation seems more closely related 

to non-mammalian vertebrates than to mouse.


A diversified role for SOX9 gene in amphibian gonads 

May Penrad-Mobayed 1 , Anwar El Jamil 1 , Rasha Kanhoush 1 , Solange 

Magre 2 , Brigitte Boizet-Bonhoure 3 and Caroline Perrin 1 

1 

Institut Jacques Monod, UMR 7592, CNRS/Universités Paris VI & 

Paris VII, Paris, France 

2 

Physiologie de l’axe gonadotrope, UMR 7079, CNRS/Université 

Paris VI, Paris, France 

3 

Institut de Génétique Humaine, UPR1142, Montpellier, France 

Our purpose is to identify common mechanisms implied in the 

acquisition of gonad identity in vertebrates. SOX9 is one of the highly 

conserved genes, which seems to be a key factor in gonad 

differentiation. Its role in the male determining pathway was clearly 

confirmed in mammals. Accumulating evidences showed that this 

gene is also expressed in male specific manner in reptiles and birds. 

However, its function in gonad differentiation in fishes and 

amphibians remains to be elucidated. Indeed, RT-PCR studies 

showed that SOX9 mRNA is expressed during gonad development 

of both sexes. 

To approach the role of SOX9 gene in amphibians, we carried out in 

situ hybridization and immunostaining studies in one urodelan and 

two anurans species: Pleurodeles waltl, Xenopus laevis and 

Xenopus tropicalis. Results show that SOX9 mRNA is expressed at 

the later stage of gonad differentiation in both sexes. However, 

SOX9 protein expression greatly differs between the male and the 

female gonads. In the testis, SOX9 protein expression is restricted to 

the nuclei of Sertoli-like cells whereas in the ovary, SOX9 protein is 

initially detected in the cytoplasm of previtellogenic oocytes and then 

translocated into nucleus at the vitellogenic stage. These data 

suggest that SOX9 have a diversified role in gonad differentiation 

throughout vertebrate evolution. 

Microdissection and DOP-PCR as a way to study the evolution 

of an old sex chromosome pair using 2 sister-species of tilapia 

with different sex determination systems, O. aureus and O. 

niloticus 

Srisupaph Poonlaphdecha 1 , Elodie Pepey 1 , Olivier Coriton 3 , Jean- 

Pierre Coutanceau 2 , Angélique D’Hont 5 , Thomas D. Kocher 4 , 

Catherine Ozouf 3 , Jean-François Baroiller 1 and Helena D’Cotta 1 

1 CIRAD, UPR20, Dept. Persyst, Campus International de Baillarguet, 

F-34398 Montpellier, France 

2 MNHN, CNRS, IFR 101 43, rue Cuvier, F-75231 Paris Cedex 05, 

France 

3 INRA, UMR 118 Amélioration des plantes et Biotechnologies 

Végétales, 35650 Le Rheu, France 

4 University OF Maryland, Dept. of Biology, College Park 20742, 

2421 Maryland, USA 

5 CIRAD, UMR-DAP, Dept. Bios, Ave Agropolis, F-34398 Montpellier, 

France 

Within the tilapia group, the sex determining locus has been located 

on linkage group 1 (LG1) in Oreochromis niloticus, whereas in O. 

aureus both LG1 and LG3 are sex-linked. Using specific BAC clones 

as probes for FISH, LG3 and LG1 have been respectively located on 

the largest and on a smaller chromosome pairs. The largest pair 

presents various traits of a relatively old sex chromosome, whereas 

LG1 seems to be at an early stage of sex chromosome evolution. 

Evidence for interactions between the 2 sex linked loci have been 

demonstrated at least in O. aureus, with minor factors (genetic and 

environmental factors) also modulating sex ratios. We have taken 

advantage >3-fold larger size of one of the chromosome pairs to 

microdissect it in order to search for genes specific to this large pair. 

This chromosome was microdissected from metaphase preparations 

of homogametic genotypes (XX and YY for O. niloticus, and ZZ for O. 

aureus). Microdissected chromosomes were amplified by DOP-PCR 

and the DOP-products were then used to screen a gonadal cDNA 

library. Positive clones were hybridized by FISH on metaphase 

spreads of different genotypes from both species. We evidenced 

positive signals located on the large chromosomes of both strands 

indicating specificity of the probes, despite the presence of large 

amounts of repetitive sequences on these large sex chromosomes. 

The conservation of the structure of the large pair between the 2 

species is discussed. 

- 36 - 

Sex determination and gonadal development in the common 

toad (Bufo bufo) 

Alvaro Roco 1 , Rafael Díaz de la Guardia Quiles 1 , Rosanna Falconi 2 , 

Francesco Zacanti 2 , Jean David Durusel 2 , Juan Antonio Marchal 1 , 

Antonio Sánchez 1 and Monica Bullejos 1 

1 

Dpto. de Biología Experimental, Facultad de Ciencias 

Experimentales, Universidad de Jaén, 23071 Jaén 

(bullejos@ujaen.es); 

2 

Departamento de Biología Evolutiva 

Experimental, Facultad de Ciencias Matematicas Fisicas y Naturales, 

Universidad de Bolonia, Italia. 

Amphibians have been traditionally used as animal models in 

embryology. Nevertheless, the knowledge about the mechanisms 

involved in sex determination and differentiation are very scarce in 

this group. 

In the species Bufo bufo there are no sex chromosomes 

morphologically distinguishable. However, by analysing the sex ratio 

of the offspring of sex reversed animals it is possible to conclude that 

the females are the heterogametic sex in this species. That is, the 

species Bufo bufo has a ZZ/ZW chromosome system for sex 

determination. 

Regarding to gonadal development, the species of the genus Bufo, 

have the particularity of developing undifferentiated ovaries (Bidder’s 

organs) in the cranial portion of male and female gonads. 

To study gonadal development in B. bufo, we are cloning in this 

species orthologous genes to those involved in gonadal development 

in other vertebrate groups. The study of the expression pattern of 

these genes during gonadal differentiation, and its relation with the 

morphological changes that take place, will enable us to establish 

their role in this process. SOX9 gene, involved in Sertoli cell 

differentiation, is among the genes we are studying. This gene codes 

for a transcription factor closely related to SRY gene, is male-specific 

and necessary for testis differentiation. In B. bufo this gene is 

expressed only in developing male gonads, starting at the stage 

where morphological differences start to be observed between 

developing testes and ovaries. 

Identification of novel sex determining genes by copy number 

analysis of patients with disorders of sexual development 

Andrew Sinclair 1 , Stefan White 1 , Hinda Daggag 1 , Lavina Gordon 1 , 

Henrik Bengtsson 2 , Terence P. Speed 2 and Eric Vilain 3 

1 

Murdoch Children’s Research Institute, Melbourne, Australia 

2 

Walter and Eliza Hall Institute for Medical Research, Melbourne, 

Australia 

3 

Division of Medical Genetics, UCLA School of Medicine, Los 

Angeles, USA 

Disorders of sexual development, ranging in severity from genital 

abnormalities to complete sex reversal, are surprisingly common. 

The cause of these problems is most often the failure of the complex 

network of genes that regulate development of testes or ovaries. 

Mutations in the critical testis-determining genes SRY and SOX9 

account for approximately 20% of XY females with complete gonadal 

dysgenesis. We have little idea about what other genes to account 

for the remaining 80% of patients. In contrast, 90% of XX males with 

gonadal dysgenesis are due to Y translocations that include SRY. 

We have collected DNA from 34 patients with gonadal dysgenesis 

(XX males lacking SRY and XY females without mutations in SRY), 

which have been hybridized to Affymetrix Genome-Wide Human 

SNP Array 6.0. Copy number analysis has been performed with a 

custom designed algorithm. So far two causative rearrangements 

have been identified in known genes (a duplication of DAX1 and a 

deletion of ~ 1 Mb in the upstream regulatory region of SOX9). Other 

rearrangements have been found encompassing candidate sex 

determination genes identified in mouse models, and there have 

been several genes deleted or duplicated that are not listed in the 

database of genomic variants. These are currently being confirmed 

and de novo status is being checked in parental DNA. A set of the 

most interesting candidate genes is being sequenced to identify 

small mutations. The combination of these powerful approaches is 

helping us to identify new genes and their regulatory regions 

involved in sex determination.


Histological and immunohistochemical analysis of the vagina 

during estrous cycle of the algerian wild Libyan jird (Meriones 

libycus) 

Souaâd Smaï-Hamdidouche 1 , Thérèse Gernigon-Spychalowicz 1 , 

Jean Marie Exbrayat 2 

1 Laboratoire de Recherche des Zones Arides (LRZA), Equipe 

Reproduction des Petits Vertébrés, Faculté des Sciences 

Biologiques (FSB), Université des Sciences et de la Technologie 

Houari Boumediene (USTHB), BP 32, DZ-16111 El Alia, Algeria. 

2 Université de Lyon ; Laboratoire de Biologie Générale de 

l’Université Catholique de Lyon, Laboratoire Reproduction et 

Développement Comparé, EPHE, 25 rue du Plat, F669288 Lyon 

Cedex 2, France. 

The female reproductive cycle of Meriones libycus, a nocturnal 

gerbillidae rodent, living in Algerian Sahara (Beni-Abbes, 30°7 N, 

2°10 W) is characterized by a short breeding period at spring and 

beginning of summer, and a long quiescent sexual period from 

summer until the end of winter. In order to understand the 

physiological variations and hormonal control affecting the genital 

tract, some histological and immunohistological methods were used 

to understand the cyclic variations in vaginal tissues. 

During the breeding cycle, vaginal epithelial cells showed cyclical 

variations. More particularly, keratinisation and exfoliation of 

epithelial cells were observed during the estrus phase; a proliferative 

activity of basal cells was also observed at this moment. 

Immunohistochemical methods revealed that the distribution of 

estrogen and progesterone receptors performed cyclic pattern. The 

highest labelling of all the hormone receptors was observed on the 

basal cells layer, showing that these ones can be implicated in 

vaginal regeneration. 

A conserved role for R-SPONDIN1 in vertebrate ovary 

development 

Craig A. Smith #1 , Christina M. Shoemaker #2 , Kelly N. Roeszler 1 , 

Joanna Queen 2 , David Crews 2 and Andrew H. Sinclair 1 

1 

Department of Paediatrics, The University of Melbourne, and 

Murdoch Childrens Research Institute, Royal Children’s Hospital, 

Melbourne, Victoria, 3052, Australia 

2 

Section of Integrative Biology, The University of Texas, Austin, 

Texas, USA 

# 

These authors contributed equally 

R-SPONDIN1 (RSPO1) is a novel regulator of the Wnt/β-catenin 

signalling pathway. Loss-of-function mutations in human RSPO1 

cause testicular differentiation in 46, XX females, pointing to a role in 

ovarian development. Here we report the cloning and comparative 

expression analysis of R-SPONDIN1 orthologues in the mouse, 

chicken and red-eared slider turtle, three species with different sexdetermining 

mechanisms. Gonadal RSPO1 gene expression is 

female up-regulated in the embryonic gonads in each species at the 

onset of sexual differentiation. In the embryonic mouse gonad, 

Rspo1 mRNA is expressed in the somatic cell lineage of females, 

with little or no expression in germ cells. In the chicken embryo, 

RSPO1 expression becomes elevated in females at the time of 

ovarian differentiation, coinciding with female-specific activation of 

the FOXL2 gene and estrogen synthesis. RSPO1 protein in chicken 

is localised in the outer cortical zone of the developing ovary, the site 

of folliculogenesis and oocyte differentiation. Chicken R-SPO1 

expression is down-regulated in embryos treated with an aromatase 

enzyme inhibitor, indicating that it is influenced by oestrogen. In the 

red-eared slider turtle, which exhibits temperature-dependent sex 

determination, female up-regulation of RSPO1 occurs during the 

temperature-sensitive period, when gonadal development is 

responsive to temperature. Accordingly, RSPO1 expression is 

temperature-responsive, and is down-regulated in embryos shifted 

from female- to male-producing incubation temperatures. These 

results indicate that RSPO1 is up-regulated in the embryonic gonads 

of female vertebrates with different sex-determining mechanisms. 

Taken together, the findings indicate that R-SPONDIN1 is an ancient, 

conserved part of the vertebrate ovary-determining pathway. 

- 37 - 

Effects of laboratory exposure to cadmium-contaminated field 

environments on gonadal development of guppy (Poecilia 

reticulata) 

Jirarach Srijunngam, Kingkaew Wattanasirmkit and Noppadon 

Kitana 

Department of Biology, Faculty of Science, Chulalongkorn University, 

Bangkok 10330 Thailand 

Exposure to low levels of cadmium, a known endocrine disruptor, 

may induce changes in gonadal development and differentiation 

resulting in adverse effects on animal reproduction. In Mae Sot 

District of Tak Province, Thailand, concerns have been raised over 

the contaminated sediment plume resulted from runoff and natural 

irrigation system that leach cadmium from zinc mining area into 

agricultural area. Although there is no report of acute toxicity on 

aquatic animals in this area, the impact of long-term environmental 

exposure to cadmium on their reproduction are of attention. In this 

study, Poecilia reticulata was used to investigate effects of laboratory 

exposure to cadmium-contaminated field environments on the 

neonate fish. Experimental aquaria were set up using water and 

sediment samples collected from contaminated and reference sites 

in Tak Province. Guppies at the age of 1 day post partum were 

raised in each aquarium for 8 weeks. After exposure, all guppies 

were sampled for histological study of the gonad. Growth of the fish 

in term of means standard length is not significantly different 

between guppies raised in the reference site or contaminated site 

conditions. The difference in sex ratio and gonadal development and 

differentiation will be compared. The implications on the effects of 

exposure to environmentally relevant dose of cadmium on sex 

differentiation and gonadal development of the neonate guppies will 

be discussed. 

Retinoic acid triggers meiosis entry in the urodele amphibian 

Pleurodeles waltl 

Angelina Wallacides, Armand Chesnel, Dominique Chardard, 

Stéphane Flament and Hélène Dumond 

EA 3442 Aspects cellulaires et moléculaires de la reproduction et du 

développement, Nancy-Université, Boulevard des Aiguillettes, BP 

239, 54506 Vandoeuvre-lès-Nancy cedex, France 

Pleurodeles waltl is a urodele amphibian that displays a genetic 

mode of sex determination. Sex differentiation can later be 

modulated either by temperature or hormonal treatment. Like in 

mammals, germ cell lineage is specified during embryonic 

development through cell-cell interactions. Recently, retinoic acid 

(RA) was demonstrated to promote meiosis entry in female mice 

foetuses while this event is delayed until puberty due to the 

degradation of RA in male gonad. To further investigate the 

timecourse of germ cell specification and differentiation in 

Pleurodeles waltl, we analysed the expression of the germ cell 

marker VASA and the meiosis marker DMC1 during normal 

development. We show that germ cell specifically express VASA 

protein at hatching, enter meiosis in late larval life in females while 

this event occurs two months after metamorphosis in males. We set 

up a protocol for organotypic cultures of gonad-mesonephros (GM) 

complexes and demonstrate that RA is necessary and sufficient to 

induce meiosis entry in P. waltl. Furthermore, we analysed Raldh2 

and Cyp26b1, the enzymes required for RA synthesis and 

degradation respectively, in steroid-induced sex reversal conditions. 

The data indicate that meiosis entry depends on Cyp26b1 repression 

in the gonad. Taken together, these results confirm with molecular 

data the late specification of germline in urodeles, indicate for the 

first time that RA- dependent meiosis entry could be a conserved 

mechanism of germ cells differentiation between urodeles and 

mammals and provide evidence for complex crosstalks between 

steroid production and RA biosynthesis in the course of sex 

differentiation.


Myogenesis in the articulate brachiopods Argyrotheca cordata 

(Risso, 1826), Argyrotheca cistellula (searles-wood, 1841) and 

Terebratalia transversa (Sowerby, 1846) 

Andreas Altenburger and Andreas Wanninger 

University of Copenhagen, Institute of Biology, Research Group for 

Comparative Zoology, Universitetsparken 15, Building 3, 2100 


We investigated development and muscle formation in larvae of the 

articulate brachiopods Argyrotheca cordata, A. cistellula and 

Terebratalia transversa using immunocytochemistry combined with 

confocal laser scanning microscopy. Full grown larvae are threelobed 

and express two pairs of bristles. Muscle development in the 

three species investigated shows a number of similarities. As such, 

the first anlagen of the musculature develop in the bristle pouches 

and the pedicle lobe. Late-stage larvae show a network of 

longitudinal muscles running from the apical to the pedicle lobe as 

well as transversal muscles situated in the apical lobe. Strong 

muscles attach to both the bristles pouches and the pedicle lobe. We 

found only few similarities between the larval myoanatomy of 

brachiopods and the hitherto investigated representatives of the two 

other lophophorate phyla, Phoronida and Ectoprocta. This may be 

due to an early evolutionary split of the ontogenetic pathways of 

Brachiopoda, Phoronida and Ectoprocta. 

The evolutionary origin of the bilaterian CNS 

Detlev Arendt 

EMBL Heidelberg, Germany 

Animal nervous systems are composed of neuron types specialized 

for functions as diverse as light perception or hormone secretion. 

While some animals such as cnidarians have few neuron types, 

human neuron types count in hundreds. Understanding the evolution 

of neuron types is key to understanding the evolution of animal 

nervous systems 

The recent advent of molecular fingerprints for defining cell types 

sets the stage the study of neuron type evolution. Each neuron type 

displays a unique profile of expressed genes, which encode 

transcription factors, microRNAs and differentiation gene batteries. 

Since many of these genes are deeply conserved in animal evolution, 

this allows the identification of homologous cell types over large 

evolutionary distances. Molecular fingerprint comparisons also allow 

identifying, within a given species, sister cell types that are related by 

evolutionary diversification. 

In my lab, we are currently generating an expression profiling atlas 

for all cell types of the developing Platynereis nervous system by a 

novel technique, Wholemount In Silico Expression Profiling. This 

technique provides a close-to-complete inventory of neuron type 

molecular fingerprints in the Platynereis brain. We use this to identify 

homologous cell types between annelids and other bilaterians such 

as insects and vertebrates. This approach will be exemplified for the 

mushroom bodies, the associative centre of the annelid brain, and 

for the various types of photoreceptor cells that harbour the 

Platynereis brain. I will discuss the implications of these data for our 

understaning of the origin of the bilaterian brain. 

Evolution and development of the ascidian neural gland 

complex 

Hélène Auger, Shungo Kano, Laurent Legendre and Jean-Stéphane 

Joly 

INRA MSNC Group, DEPSN, Institut A. Fessard, CNRS, 1 Avenue 

de la Terrasse, 91198 Gif-sur-Yvette, France. 

The cerebral ganglion of adult ascidians is apposed to the neural 

gland complex. This later has three parts: a ciliated funnel, a ciliated 

duct and one neural gland. I will first report our progress on the study 

of the ciliated funnel epithelium. 

S11 - Evolution and Development 

- 38 - 

We have looked for genes expressed in this cell type, proposed to be 

related to some of the vertebrate pituitary endocrine cells. These 

experiments are based on in situ hybridization experiments, and on 

micro-array experiments on fluorescent epithelia micro-dissected 

from GFP expressing transgenic lines. The neural gland is another 

organ whose homology with vertebrate organs is still debated. It is a 

spongious sac with a folded epithelium. It has been suggested that it 

is involved in osmoregulation, with the ciliated duct generating an 

unidirectional influx of seawater into the neural gland. In Deyts et al. 

(2006), we reported that the ascidian neuropeptide G-proteincoupled 

receptors, the vertebrate homologues of which are involved 

in homeostasis, are expressed in the neural gland complex. More 

recently, ISH on thick sections revealed that vertebrate markers 

specifically found in the choroid plexus, such as Transthyretin or 

Gelsolin, are also expressed in the ascidian neural gland. These 

results highlighted similarities between the ascidian neural gland and 

the vertebrate circumventricular organs. Circumventricular organs 

are located along the midlines of vertebrate brain ventricles. They 

appear to be associated with multiple functions, at first as 

transducers of information between the blood, neurons and the 

cerebrospinal fluid. Among them, the vertebrate choroid plexus, a 

neural circumventricular organ, produces large quantities of 

cerebrospinal fluid. 

These findings, together with electrophysiological experiments and 

cell lineage analysis through metamorphosis should allow us to 

determine more precisely the ontological origin and morphogenesis 

of the neural gland and its function in ascidians. This leads us to 

raise hypotheses on the origin in chordates of neuroendocrine and 

endocrine cell types associated with nervous system. 

The segmented ancestor of protostomes and the origin of 

parasegmental patterning 

Guillaume Balavoine 

Centre de Génétique Moléculaire du CNRS, avenue de la Terrasse, 

91198, Gif-sur-Yvette, France 

The comparison of the genetic processes of segment formation in 

three distant groups of bilaterian metazoans (arthropods, insects and 

annelids) has opened a heated debate that considerably impacts on 

our conceptions of metazoan evolution: was the last common 

ancestor of protostome and deuterostome animals a metameric 

organism? Processes of segment formation can be divided in three 

steps: the proliferation of axial tissues, the production of a periodic 

segmentation signal along this axis and the patterning of the subparts 

of individual segments. While the ?cascade? of metamere 

patterning genes of the fruitfly and the known mechanisms of somite 

formation in vertebrates have little in common, intriguing similarities 

have been uncovered in other model animals. Gene expression and 

interference data suggest that the Notch pathway may be involved in 

synchronising a periodic segmental signal in some arthropods 

(spiders, centipedes) in a similar way as in vertebrates. Also, a 

number of genes have been shown to present similar ?segment 

polarity? patterns in the marine annelid Platynereis dumerilii and 

arthropods. Pharmacological treatments suggest that the hedgehog 

pathway is involved in regulating the individual segmental patterns in 

Platynereis in the same way as in arthropods. The spatial 

relationships between these annelid ?segment polarity? patterns are 

remarkably similar to their arthropod homologues and leave few 

doubts on the segmented architecture of the protostome ancestor 

body. Additionally, they suggest that annelid segments are 

homologous with arthropod parasegments. A scenario for the origin 

of arthropod parasegmental patterning is proposed.


Development under stress. Fluctuating asymmetry of Rana 

perezi from Monegros arid zone (NE Spain) 

Carmen Burghelea, Dragos Zaharescu and Antonio Palanca 

Animal Anatomy Laboratory, Faculty of Sciences, Vigo University, 

campus Lagoas Marcosende 36310, Vigo, Spain 

Fluctuating asymmetry (FA) has been promoted as a measure of 

developmental instability, reflecting disorders of developmental origin 

in stressed environments. 

We investigated skeletal FA of R. perezi from rice paddies of 

Monegros, an arid region in NE Spain that undergone intensive 

agriculture during the last decade. FA was assessed on a sample of 

318 individuals using image analysis of X-ray radiographies. 

We compared the degree of FA in multiple traits for juveniles, males 

and females. All size corrected asymmetries were significantly 

different from 0. 

The femur showed the highest FA in all developmental stages with a 

decrease towards the adults, probably for being a higher functional 

trait. However for other characters such as metatarsus, humerus, 

coracoids and the maxillary arch, FA has increased to adult stage, 

suggesting that these latter ones may be less capable to cope with 

the developmental disorders during the ontogeny. 

Glycosphingolipids expression in neural tissues of three rat 

strains 

Vedrana Čikeč Čulić 1 , Jasminka Rešić 1 , Maja Tomasović 2 , Tatijana 

Zemunik 3 and Anita Markotić 1 

1 

Department of Biochemistry, Split University School of Medicine, 

Šoltanska 2, 21000 Split, Croatia 

2 

Department of Pediatrics, Clinical Hospital Split, 21000 Split, 

Croatia 

3 

Department of Biology, Split University School of Medicine, 

Šoltanska 2, 21000 Split, Croatia 

Many neurological disorders and injuries such as Parkinson's 

disease, multiple sclerosis, stroke, traumatic injury to CNS etc. have 

movement abnormalities as major symptom. Many animal models of 

neurological disease and injury exist, especially in rats, and most 

common used strains are Lewis, Wistar and Sprague-Dawley. 

Glycosphingolipids, both neutral and gangliosides are, according to 

their role, differently distributed in neural tissues and synaptic 

mechanisms. Therefore, comparison of different types of 

glycosphingolipid expression in neural tissues of three rat strains 

was analyzed. The aim was to see the possible connection of 

glycosphingolipid type in neural tissue depending of animal type. 

Three different male, age-matched rat strains were used: Lewis, 

Wistar and Sprague-Dawley. Glycosphingolipids were isolated from 

animal tissues by standard procedure. Distribution of 

glycosphingolipids in brain, cerebellum and pons of each group of 

animals were analyzed by orcinol-staining. Cerebellum of Lewis rats 

showed slightly lower expression of glycosphingolipids GD1b, GD1b, 

GT1b and GQ1b in comparison to cerebellum of Wistar rats, and 

extremely lower expression of same glycosphingolipids in Sprague- 

Dawley cerebellum. Moreover, Wistar brain and pons showed 

enhanced expression of those glycosphingolipids comparing to 

Sprague-Dawley rats. Finally, the highest expression of complex 

glycosphingolipids was found in brain of Lewis rats. We can 

conclude that differences in glycosphingolipid expression between 

neural tissues of three laboratory rat strains exist, which suggests 

that special glycosphingolipids play important role in special neural 

synapses, but to final conclusion, advanced immuno-staining of 

individual antibodies should be performed. 

Axis specification mechanisms in the dogfish S. canicula : 

implications on the origin and evolution of extraembryonic 

tissues in jawed vertebrates 

Marion Coolen 1 , A. Gombault 1 , Corinne Da Silva 2 , Patrick Wincker 2 , 

Arnaud Menuet 1 and Sylvie Mazan 1 

1 CNRS-UMR 6218, Université d'Orléans, 45071 Orléans, France 

2 GENOSCOPE (CEA) and UMR CNRS 8030, Université d’Evry, 2 

rue Gaston Crémieux, 91057 Evry, France 

- 39 - 

The genetic mechanisms that control the establishment of early 

polarities and their link with embryonic axis specification and 

patterning seem to substantially diverge across vertebrates. A 

particularly intriguing issue is that in amniotes, these processes rely 

on signals secreted by extraembryonic territories, which have no 

clear equivalent in amphibians and teleosts. In order to gain insight 

into the underlying unity of axis specification mechanisms, we have 

used an evo-devo approach, aimed at reconstructing the ancestral 

state of jawed vertebrates. To this end, we have achieved an 

extensive molecular characterization of the blastula and early 

gastrula in a chondrichthyan, the dogfish Scyliorhinus canicula, 

chosen both for its phylogenetic position as an outgroup to 

osteichthyans and its developmental characteristics. A molecular 

characterization relying on analysis of a wide range of genetic 

markers leads to the identification of presumptive embryonic 

territories, including the organizer. More unexpectedly, it also points 

to the presence, with comparable relative locations, of candidate 

homologues of extraembryonic territories of amniotes known to play 

key parts in axis specification in the mouse such as the distal 

visceral endoderm, extraembryonic visceral endoderm or 

extraembryonic ectoderm. Finally, in ovo pharmacological treatments 

at blastula stages support the conservation of essential roles of 

Nodal signaling in their specification and in axis formation. These 

data suggest that the subdivisions of the ectoderm and endoderm 

into territories homologous to the major embryonic and 

extraembryonic components of amniotes have an ancient origin in 

jawed vertebrates. They lead to a unifying model of axis specification 

mechanisms, thus providing a comparative framework to reassess 

conservation and divergence of the major vertebrate model 

organisms. 

Sauka-Spengler, T., Baratte, B., Lepage, M. and Mazan., S. 2003. 

Dev. Biol. 263, 296-307. 

Coolen, M., Sauka-Spengler, T., Nicolle, D., Le-Mentec, C., 

Lallemand, Y., Da Silva, C., Plouhinec, J.L., Robert, B., Wincker, P., 

Shi, D.L. and Mazan, S. 2007. PLoS ONE, 2, e374. 

Variation of a complex structure: the joint effects of mutations 

and of developmental temperature on the Drosophila wing 

Allan Debelle and Vincent Debat 

UMR 5202 Origine, Structure et Evolution de la Biodiversité, 

Département Systématique et Evolution, Muséum National d'Histoire 

Naturelle, 45 rue Buffon, 75005, Paris, France 

While environmental effects have traditionally been considered a 

nuisance in developmental genetics, phenotypic plasticity – a change 

in phenotype due to the environment – is becoming a central theme 

in evolutionary developmental biology. It is now universally accepted 

that environment does alter gene expression and morphogenesis. In 

spite of this, a surprisingly small number of studies have directly 

investigated the joint effects of genetic and environmental 

manipulations onto a complex structure. Here we used heterozygous 

insertional mutations altering 17 genes known for their role in the 

formation of the drosophila wing. The flies were raised at two 

developmental temperatures (18 and 28°C). Landmark based 

geometric morphometrics were used to analyse the variation of the 

wing size and shape. Our results show that (i) the temperature alters 

the effect of most of the mutations; (ii) reciprocally most mutations 

change the effect of temperature on wing size and shape (in other 

words mutations affecting wing formation also change wing plasticity) 

(iii) both temperature and mutations alter the patterns of wing shape 

variation. Interestingly, we found that temperature systematically 

changes the main direction of wing shape variation among 

individuals, but not the stochastic component of variation as 

measured by fluctuating asymmetry. Considering that phenotypic 

variation is the raw material on which selection acts, deciphering the 

processes by which environment impacts the directions of available 

variation is of critical importance. Our results suggest that systematic 

environmental manipulations would be very beneficial to genetic 

studies to gain a proper understanding of organism’s variational – 

and thus evolutionary – properties.


New insights on the nematogenesis process give new elements 

to understand the acquisition of the nematocyte 

Elsa Denker, Nicolas Rabet, Eric Bapteste and Michael Manuel 

UPMC Univ Paris 06, UMR 7138 CNRS UPMC MNHN IRD, Case 05, 

7 quai St Bernard, F-75005 Paris, France. 

Nematocytes are a synapomorphy of Cnidaria probably including 

Myxozoa (Jimenez-Guri et al., 2007). The fundamental importance of 

this cell in living Cnidaria or Myxozoa suggests that its acquisition 

probably conditioned the evolutionary success of the whole group. In 

order to understand the evolution of this particular cell, we 

investigated nematogenesis in a new model, the tentacle bulb of 

Clytia (Hydrozoa), a promising experimental system (Denker et al., 

2008). Firstly, we investigated the expression of homologues of 

bilaterian neural genes. Together with data from the literature, 

coming from development or neurophysiology, we confirmed that the 

cnidocyte is a very specialised neural cell. Then, we investigated the 

nematocyst, the typical extrusive organelle of the nematocyte. We 

made a comparison of extrusive organelles occurring in various 

lineages using data from the literature. In all cases except the 

Myxozoa, we were led to exclude primary homology with the 

cnidarian nematocyst on the basis of deep ultrastructural differences, 

thus excluding endosymbiotic origin of these organelles contrary to 

previous proposals (Okamura & Canning, 2003). In order to 

understand particularities of nematocysts, we also investigated the 

capacity of PGA (poly gamma-glutamate) synthesis, a polymer 

accumulating in the nematocyst and known only in Cnidaria among 

Eukaryotes. 

We found that cnidarian genomes harbour one homologue of a 

bacterial gene known to be implicated in PGA synthesis and 

accumulated indications that this gene was horizontally transferred 

before the diversification of Cnidaria. 

We propose a scenario in which the ancestral nematocyte was a 

neuro-sensory cell, which acquired an extrusome with particular 

efficiency. 

Jimenez-Guri E, et al. 2007. Science 317(5834): 116-118. 

Denker E, et al. 2008. Develop Biol 315(1): 99-113. 

Okamura B, Canning EU. 2003. TREE18(12): 633-639. 

Evolutionary conservation of the glycoprotein hormone alpha 

and ß subunit gene precursors 

Sandra Dos Santos, Claire Bardet, Damien Habert and Bruno Quérat 

UMR5166 USM501 CNRS-MNHN RDDM BP32, 57 rue Cuvier, 

75231 Paris cedex 05, France 

The pituitary gonadotropins and thyrotropin are heterodimers 

composed of a common alpha subunit (SU) and a specific βSU. Two 

genes were recently identified in vertebrates and in some protostome 

genomes that present structural similarities with the alpha and β SUs 

and were thus named glycoprotein alpha2 (GPA2) and β5 (GPB5). 

Urochordates, the sister group of vertebrates and more basal 

deuterostomes have no pituitary gland and no acknowledgeable 

pituitary hormone genes. But they do have GPA2 and GPB5 related 

genes that might thus represent the molecular precursors of the 

alpha and β glycoprotein hormone SUs. In this study, we show that 

GPA2 and GPB5 are organized in inverse tandem in basal 

deuterostomes, an organization that is not conserved in vertebrate 

genomes. However, some of the genes found in the vicinity of the 

tandem in basal deuterostomes are also present close to vertebrate 

glycoprotein hormone β SU genes. GPA2 and GPB5 are expressed 

in the gonads of the cephalochordate amphioxus, but also in 

intestine and other non-endocrine tissues. We also have identified a 

glycoprotein hormone receptor-like gene in the amphioxus genome. 

The localization of its expression is in progress. Our results bring 

new arguments for parental relationships between GPA2 and GPB5 

and the vertebrate glycoprotein hormone SUs but further studies are 

necessary in order to assess whether a possible GPA2/GPB5 

heterodimer could represent the functional glycoprotein hormone 

ancestral form. 

- 40 - 

Evolution of the Neural Crest: New Insights From Urochordates 

William R. Jeffery 

Department of Biology, University of Maryland, College Park, MD. 

20742 USA 

New insights from ascidians on the chordate ancestry of the neural 

crest (NC) will be discussed. Ascidians have neural crest-like cells 

(NCLC), defined by migration from the dorsal midline, expression of 

some vertebrate NC markers, and development into body pigment 

cells. These characters suggest that primordial NC cells were 

already present in the common ancestor of the vertebrates and 

urochordates, which have been recently inferred as sister groups. 

The primitive role of NCLC may have been pigment cell dispersal 

and development. Later, additional functions may have appeared in 

the vertebrate lineage, resulting in the evolution of definitive NC cells. 

I will also discuss new evidence showing that the ascidian 

homologues of vertebrate NC specifier genes, including foxD3, cMyc, 

and twist-like, and NC effector genes, such as those encoding 

rhoABC and cadherins, are expressed in Ciona intestinalis NCLC. In 

contrast, the ascidian homologues of vertebrate neural plate border 

specifier genes, such as msxb, pax3/7, and dlx3/5, are not 

expressed in Ciona NCLC. These results suggest that the definitive 

vertebrate NC was elaborated to include many additional functions 

by co-option of neural plate border genes into a pre-existing genetic 

cascade responsible for NCLC development. 

Evolution of segmentation: Regulation of spider segmentation 

by Wnt8 

Alistair P. McGregor, Matthias Pechmann, Evelyn E. Schwager, 

Natália M. Feitosa, Sarah Kruck, Manuel Aranda and Wim G. M. 

Damen 

University of Cologne, Institute for Genetics, Evolutionary Genetics, 

Zülpicher Straße 47, D-50674 Köln, Germany 

One outstanding question in animal evolution is whether the last 

common ancestor of the three segmented phyla, chordates, annelids 

and arthropods, was itself segmented. In vertebrates, somitogenesis 

is regulated by FGF, Wnt and Delta/Notch signaling. Our previous 

analysis of Delta/Notch signaling in spiders suggested that minimally 

this aspect of the regulation of vertebrate somitogenesis and 

arthropod segmentation is similar. Our latest data show that another 

important regulator of vertebrate somitogenesis, Wnt8, is also 

required for spider segmentation. Knockdown of Wnt8 expression in 

Achaearanea tepidariorum resulted in failure to properly establish a 

posterior growth zone and caused a breakdown of posterior 

segmentation. Our results suggest Wnt8 has a similar role in 

vertebrates and spiders in establishing and maintaining a posterior 

segmentation zone and may be part of an ancient regulatory network 

for segmentation that could have been present in the common 

ancestor of segmented animals.


Loss of an enzymatic activity involved in sterol metabolism 

during evolution: the interesting case of a cactophilic 

Drosophila species 

Virginie Orgogozo, Sophie Murat and Chantal Dauphin-Villemant 

Équipe Biogenèse des Stéroïdes, Laboratoire Protéines Biochimie 

structurale et fonctionnelle, FRE2852 CNRS, Université Pierre et 

Marie Curie, 7 Quai St Bernard, F-75005 Paris, France 

Host specialization in insects that feed on plants provides an 

excellent opportunity to study the genetic basis of ecological 

adaptation. The specialization of Drosophila pachea towards its 

single host plant, a cactus species named Lophocereus schottii, is 

remarkable in that it involves both ecotoxicological and hormonal 

interactions between plant and insect. While D. pachea is one of the 

few fly species that is resistant to toxic alkaloid compounds of 

Lophocereus schottii, it requires 7-dehydrogenated sterols produced 

by this cactus to survive, because it has lost the capacity to 7,8dehydrogenate 

cholesterol (first enzymatic step of the ecdysone 

biosynthesis pathway). 

In insects, this enzymatic reaction appears to be catalyzed by a 

Rieske-domain enzyme encoded by the neverland gene (Yoshiyama 

et al, 2006). To determine the genetic basis of loss of 7,8dehydrogenation 

in D. pachea during evolution, we analyzed the 

evolution of neverland in D. pachea and other Drosophila species. 

We found that several amino acids that are otherwise conserved 

across insects have changed in D. pachea Neverland protein. Our 

study suggests that neverland may have acquired a new function in 

D. pachea. 

Evolutionary trends of the pharyngeal dentition in 

Cypriniformes (Actinopterygii, Osteichthyes) 

Emmanuel Pasco 1 , Paul Tafforeau 2 , Jérôme Adrien 3 , Laurent Viriot 4 

and Vincent Laudet 1 

1 Institut de Génomique Fonctionnelle, UMR 5242, Ecole Normale 

Supérieure de Lyon, 46 allée d’Italie, 69364 Lyon cedex 07 France 

2 European Synchrotron Radiation Facility, BP220, 6 rue Jules 

Horowitz, 38043 Grenoble Cedex, France 

3 Mateis, UMR 5510, INSA Lyon, Blaise Pascal, 7 avenue Jean 

Capelle, 69621 Villeurbanne Cedex, France 

4 International Institute of Paleoprimatology, iPHEP, CNRS UMR 

6046 Faculté SFA, Université de Poitiers, 40 avenue du Recteur 

Pineau, 86022 Poitiers Cedex, France 

The freshwater fish order Cypriniformes is characterized by the 

absence of oral teeth and the presence of pharyngeal teeth located 

on the fifth ceratobranchial. Among this order, members of the 

superfamily Cobitoidea always display one row of pharyngeal teeth 

with a conical shape whereas members of the superfamily 

Cyprinoidea, and more especially the family Cyprinidae, display one, 

two or three tooth row(s) with different shapes. It is thus interesting to 

understand the evolutionary mechanisms responsible for this 

diversity. We performed a preliminary survey to explore the diversity 

of cypriniforme dentitions and infer evolutionary trends. A total of 

forty cypriniforme species have been scanned using either 3D 

conventional microtomography or synchrotron microtomography. 

Dental formulas and tooth shape were determined for each of them. 

By comparing these morphological results with updated molecular 

phylogenies of the Cypriniformes, we propose that the ancestral 

condition of Cyprinidae was a 1-row dentition with conical shape. 

Moreover, we observed that the appearance of a new tooth row 

happened several times during the evolution of Cyprinidae. Our 

model suggests that two contrasting evolutionary pathways were 

taken by the two superfamilies: a single row with numerous teeth for 

Cobitoidea, multiple rows with few teeth for Cyprinoidea. Finally, 

factors such as feeding habits and constraints due to the size don't 

seem to explain entirely the evolution of cypriniforme pharyngeal 

dentition. 

- 41 - 

How can sponges bring new insights on the origin of neurosensory 

cells? 

Emmanuelle Renard (Deniel), Eve Gazave, Alexander Ereskovsky, 

Jean Vacelet, Pascal Lapébie and Carole Borchiellini 

DIMAR (diversité et évolution des écosystèmes marins), Centre 

d'océanologie de Marseille- Université de la Méditerrannée, rue de la 

batterie aux lions, 13007 Marseille, France 

The capacity of all cells to respond to stimuli implies the conduction 

of information at least on short distances. In multicellular organisms 

more complex systems of integration and coordination of activities 

are necessary. In most animals the processing of information is 

realized by a "true" nervous system. Among the most basal taxa, 

sponges, unlike all other metazoans, are nerveless so that it is 

traditionally assumed that neuro-sensory cells originated only once, 

in Eumetazoa. 

Nevertheless, new data may lead to reconsider this hypothesis: 

i) Sponges are interestingly able to react upon external stimuli, to 

contract, and display diurnal rhythms. Nervous-system like 

coordination mechanisms was evidenced more recently when action 

potentials and GABAergic coordination mechanism were reported. 

ii) Genomic analyses show that, despite their apparent 

morphological simplicity, sponges harbour an unexpected genetic 

complexity. Notably genes orthologous to those implicated in the 

nervous system patterning or required for sensory and neural 

functions in Eumetazoa were found. 

Two competing hypothesis have been proposed: on one hand, 

recent phylogenomic trees placing Ctenophora at the basis of 

Metazoa suggest a more complex history involving either 

convergent acquisition of neuro-sensory cells in ctenophores and 

(cnidarians + bilaterians), or their secondary loss in sponges. On the 

other hand, ultrastructural data suggest that some sponge cells may 

be sensory-cells and may thus be regarded as universal common 

ancestral sensors. 

We will discuss how the study of non bilaterian models and in 

particular expression data of sponge genes may help testing these 

three conflicting views. 

The origin of biramous appendages - a new view on arthropod 

limb evolution 

Gerhard Scholtz 

Philippstr. 13, 10115, Berlin, Germany 

The impressive functional and structural diversity among arthropod 

limbs is commonly interpreted as a variation of two or three basic 

limb types that are identified by the number of branches and axes. 

Despite some recent gene expression studies, however, it is still not 

clear how the various limb branches are formed in terms of 

hierarchies of axis differentiation and how arthropod limbs evolved. 

Accordingly, the interpretation of fossil and recent limb types is con! 

tentious. Moreover, there is no consensus how and when a proper 

biramous limb evolved. In my talk I discuss the various hypotheses 

on arthropod limb interpretation and limb evolution. Based on recent 

cell-lineage and gene expression studies of our group the hypothesis 

is developed that biramous arthropod limbs evolved by a split of the 

primary proximodistal axis of the limb bud. Furthermore, it is 

suggested that proper biramous limbs occurred later in arthropod 

evolution than generally thought. These ideas are supported by a 

variety of data and are testable by future investigations.


The nervous system in orthonectid Intoshia variabili 

George Slyusarev 

Saint Petersburg State University, Russia 

The nervous system in the female orthonectid Intoshia variabili was 

revealed by anti-serotonin labeling. The nervous system consists of 

two pairs of symmetrically arranged nerve cells. The neuron bodies 

lie between the ciliated cells and the muscle cells in the anterior part 

of the female body. The anterior nerve cells are multipolar, they are 

brought closer together than the posterior pair of neurons and are 

located dorsally. A few processes run anteriorly and “ventrally” from 

the anterior neuron pair, forming a well developed plexus. Some part 

of these processes appears to get in contact with a provisional 

sensory organ. Two posterior bipolars are located laterally and each 

has a long process running further posteriorly and lying between the 

epithelial and the muscle cells. The processes do not reach the very 

end of the body, The nervous system in the orthonectid I. variabili is 

distinctly bilateral. 

The finding of the nervous system in the orthonectids should put an 

end to the dispute concerning phylogenetic relationships between 

Orthonectida and Diciemyda. The phylums Orthonectida and 

Diciemyda cannot be joined in a single group Mesozoa. 

Cloning of cyclic AMP responsive element binding protein 1 

(CREB1) cDNA in the earthworm Eisenia fetida 

Kosuke Watanabe 1 , Sumitaka Hase 2 , Toshinobu Shimoi 1 , Hiroto 

Ogawa 3 , Kohji Hotta 1 and Kotaro Oka 1 

1 Center for Biosciences and Informatics, School of Fundamental 

Science and Technology, Keio University, Hiyoshi, Kohoku-ku, 

Yokohama, Kanagawa, Japan ; 2 Faculty of Science and Technology 

Experiment Education Support Center, Keio University, Hiyoshi, 

Kohoku-ku, Yokohama, Kanagawa, Japan ; 3 Department of Biology, 

Faculty of Medicine, Saitama Medical University, Iruma-gun, Saitama, 

Japan 

The earthworm can be classically conditioned by weak vibration as 

conditioned stimulus and by light as unconditioned stimulus. Our 

previous study showed that both de novo mRNA and protein 

synthesis are required for long-term memory (LTM) formation in the 

earthworm, Eisenia fetida. In other words, both transcription and 

translation are needed for LTM formation. However, the neurons 

involved in LTM formation remain unknown. Cyclic AMP responsive 

element binding protein 1 (CREB1) plays an essential role in LTM 

formation as a transcription factor in various animals including 

Lymnaea stagnalis. The aim of our study is to observe localization of 

CREB1 gene in the central nervous system of the earthworm, and 

identify the neurons involved in LTM formation. 

As a first step, we obtained a partial sequence of CREB1 homolog in 

the earthworm, using sets of degenerate primers designed to target 

a bZIP domain which is well conserved over hundreds of species of 

CREB1. Next, we determined a complete nucleotide sequence of 

CREB1 homolog by RACE-PCR, and it was found that obtained 

CREB1 has also two domains, bZIP and P-box, like other animals’ 

CREB1. Our study is the first report of existence of CREB1 in 

annelids, which will shed light on the evolution of molecular 

mechanism in LTM. 

- 42 - 

Inheritance of Color Phenotype in the Sea Urchin Lytechinus 

variegatus 

Maria L. Wise and Daniel Rittschof 

Duke University Marine Laboratory, Beaufort NC 28516, USA 

Lytechinus variegatus is a variably colored sea urchin common 

throughout the western Atlantic and Caribbean, from Beaufort, North 

Carolina to Brazil. Field sampling indicates that coloration in L. 

variegatus varies with geographic location and is more variable in 

individuals in some locations than in others. The color phenotype of 

the spines can be white, green, purple, lavender and pink and is 

often a combination of two or more colors. Pawson and Miller’s 1982 

experimental crosses of L. variegatus from Florida and Bermuda 

demonstrated a genetic link to color inheritance and in this study we 

expand on these experiments by creating a series of crosses 

involving several color morphs. Early stage juveniles (< 4mm in 

horizontal diameter) of all color crosses are very similar in 

appearance - translucent white with a central lavender band on the 

spines. Color change occurs when juveniles reach approximately 5 

mm in horizontal diameter and continues until the adult color 

phenotype is evident (approximately 15 mm in diameter). Color in the 

F1 generation is complex suggesting it is a multigene trait for both the 

spines and test. Experiments to create an F2 generation are ongoing 

and may help shed light on the mode of color inheritance. 

Brain development in cephalopod molluscs studied by means of 

immunohistochemistry and gene expression analysis 

Tim Wollesen and Andreas Wanninger 

Department of Biology, Research Group For Comparative Zoology, 

University of Copenhagen, Copenhagen, Denmark 

Among invertebrates, cephalopod molluscs such as squids, cuttlefish, 

octopuses and nautiluses are known for their sophisticated behavior 

and cognitive abilities resembling those of vertebrates. Their central 

nervous system (CNS) represents an integrative part of their 

evolutionary success and has been subject of a wealth of studies 

employing classic neuroanatomical methods. However, to date there 

are virtually no comparative investigations on the distribution of 

neurotransmitters within the developing CNS of cephalopods. In this 

study we compare the gross anatomy of the CNS of adults and 

developmental stages of selected coleoid cephalopod species by 

means of immunohistochemistry combined with confocal laser 

scanning microscopy. We investigated the distribution of the neural 

markers FMRFamide and acetylated α-tubulin in combination with 

phalloidin staining in the brain of the pygmy squid Idiosepius 

notoides, the cuttlefish Sepia officinalis, the loliginid squid Loligo 

vulgaris, the sepiolid squid Euprymna scolopes and the octopod 

Argonauta argo. Moreover, FMRFamidergic gene expression during 

CNS development of Idiosepius are provided. This approach aims at 

characterizing the expression of certain neural markers in the various 

brain lobes during ontogeny in order to generate a highly resolved 

atlas of cephalopod neurogenesis in time and space. 

The neuropil in the CNS of all above mentioned cephalopod species 

is intensely labeled by phalloidin and exhibits strong α-tubulinergic 

immunoreactivity. FMRFamidergic structures show a restricted 

distribution and comprise cell somata within the cerebral ganglia and 

neurites of the circumoesophageal nerve ring. Although certain traits 

are shared among the species studied, others are highly species 

specific.


In search of the evolutionary meaning of natriuretic peptides for 

cardiac function in terrestrial animals 

Shigeru Yoshida, Naoto Fuwa, Yoshie Ishijima, Ruriko Mukai, J. 

Yamaguchi, T. Tsuchikura and Teruki Hagiwara 

Department of Life Science, School of Science & Engineering, Kinki 

University, Higashi-Osaka City 577-8502, Japan 

Life first emerged in sea water, and some animals left the water as 

the concentration of sea-water Na + increased gradually over billions 

of years. Natriuretic peptides (NPs), which are released from the 

heart and keep the concentration of body fluid Na + ([Na + ]o) at a low 

level by increasing Na + excretion, first appeared in fish during animal 

evolution and are active in terrestrial vertebrates. The aim of the 

present work is to elucidate the relationship between NPs and the 

cardiac function of mammals, the most highly developed terrestrial 

life, since the heart possesses receptors for NPs. 

Electrophysiological properties of the beating heart were examined 

by the suction-electrode method in adult mice. The amplitude of the 

action potential and its differentiated record increased when [Na + ]o 

was raised from a normal concentration (140 mM) to 190 mM, 

indicating a larger Na + influx in the atrium. In contrast, contractile 

force of the atrium, measured with a strain gauge, decreased when 

[Na + ]o was elevated. The suppressed contractility of the atrium, 

however, recovered when atrial NP (ANP; 1-10 nM) was added to 

the high-Na + solution. This reinforcing effect was greater in high-Na + 

media than in normal solution. From the perspective of evolution, the 

reason why some aquatic animals escaped from the water seems to 

be to supply mitochondria with a sufficient amount of oxygen for the 

manufacture of ATP. In addition, such mitochondria are so 

vulnerable to high Na + that terrestrial animals had to lower body fluid 

Na + with ANP. 

- 43 -


S12 - Transitions from clonal to sexual reproduction: key variations of a key process 

The function of sperm donating species to the evolution and 

persistence of unisexual salamanders (caudata: ambystomatidae) 

Jim Bogart 

Department of Integrative Biology, University of Guelph, N1G 2W1, 

Guelph, CANADA 

Unisexual salamanders in the genus Ambystoma are common 

around the Great Lakes region in eastern North America. Based on 

mitochondrial DNA sequences, they originated from a single 

hybridization event that involved a female that shared a common 

ancestor with Kentucky A. barbouri 2.4 to 3.9 million years ago. 

Individual unisexual salamanders reproduce by stealing sperm from 

donors of normally bisexual species so their reproductive mode is 

described as kleptogenesis. As many as 22 different diploid and 

polyploid unisexual biotypes are known and they all possess at least 

one A. laterale genome. One or more other nuclear genomes have 

been derived from sperm donors that may involve males of five 

distinct species. Sperm may serve only to stimulate egg 

development (gynogenesis) but can be incorporated to replace a 

nuclear genome or to elevate ploidy. Genome replacement is 

considered to be an essential evolutionary strategy to rectify meiotic 

problems that arise from recombinations and translocations among 

gynogenetic offspring and to introduce novel genetic combinations 

that have a selective advantage. Molecular tools such as 

microsatellite DNA loci, expressed sequence tags (EST), and 

genomic in situ hybridization (GISH) can be used to identify the 

genomic constitution of individuals and to test hypotheses pertaining 

to the role of sperm donors in the co-evolution of the disparate 

cytoplasmic and nuclear components in unisexual salamanders. 

Bogart, J. 2003. Genetics and systematics of hybrid species. Pp. 

109-134, In D. M. Sever (edt.), Reproductive Biology and Phylogeny 

of Urodela. M/s Science Inc., Enfield NH. 

Bogart, J. P., K. Bi, J. Fu, D.W.A. Noble, and J. Niedzwieki. 2007. 

Unisexual salamanders (genus Ambystoma) present a new 

reproductive mode for eukaryotes. Genome 50: 119-136. 

Bi, K., J. P. Bogart, and J. Fu. 2008. Genealogical relationships of 

unisexual salamanders of the genus Ambystoma inferred from 

intergenomic exchanges and 45S rDNA cytotypes. Chromosome 

Research 16: 275-289. 

Sex in asexuals: how occasional sex determines the fate of 

parthenogenetic populations 

Thomas D'Souza 

Auf der Morgenstelle 26, 72076, Tuebingen, Germany 

Sex is expensive compared to asexual reproduction. Nevertheless 

sex is more widespread among multi-cellular organisms. Many 

theories have been developed to solve this so-called paradox of sex 

and to clarify why sex is advantageous. Particularly when combining 

these theories, the benefits of sex seem to outweigh the costs. 

Consequently, asexuals are usually considered as evolutionary 

dead-ends. This ra! ises the question whether asexual systems are 

always truly clonal or whether they have cryptic forms of sexuality 

that enhances their viability. This is important as, at least 

theoretically, a limited amount of sexuality is sufficient to compensate 

for the long-term costs of clonality. Here, the advantages of 

occasional sex will be demonstrated using the planarian flatworm 

Schmidtea polychroa as an example. Parthenogenetic S. polychroa 

are simultaneous hermaphrodites, which require sperm from a 

partner in order to trigger embryo development (pseudogamy). As 

parthenogens are hermaphroditic, they produce fertile sperm and do 

not rely on sexual sperm donors. Parthenogenesis in S. polychroa is 

not always strictly clonal, but sometimes contains occasional, sexual 

process. This mixed reproduction mode may combine the benefits of 

sex and asex and increases the survival of parthenogenetic 

populations in two different ways. Firstly, occasional sex promotes 

the investment in male reproductive organs of parthenogens, which 

is essential for rare sex among parthenogens. 

- 44 - 

Secondly, occasional sex in parthenogenetic S. polychroa leads to 

an increase in fitness variance at the local population level, which in 

turn increases mean fitness, which can be interpreted as a sign! 

ature of a more effective natural selection. 

Sexual reproduction in triploid forms of planarian Dugesia 

ryukyuensis 

Kazuya Kobayashi 1 , Hirotsugu Ishizu 1 , Motonori Hoshi 2 and Midori 

Matsumoto 1 

1 

Biosciences and Informatics, Keio University, Hiyoshi 3-14-1, 

Kouhoku-ku, Yokohama, 223-8522, Japan. 

2 

University of the Air, Wakaba 2-11, Mihama-ku, Chiba, 261-8586, 

Japan. 

Triploidy has generally been considered to be an evolutionary dead 

end due to problems of chromosomal pairing and segregation during 

meiosis. Thus, the formation of tetraploids and diploids from triploid 

types is a rare phenomenon. In the planarian Dugesia ryukyuensis, 

both fissiparous and oviparous triploids occur in nature. It is believed 

that oviparous triploids reproduce via pseudogamy (spermdependent 

parthenogenesis). We have successfully induced 

experimental sexualization of fissiparous triploids in D. ryukyuensis. 

Following sexualization, the triploids develop hermaphroditic gonads 

and other reproductive organs and begin reproducing by copulation 

instead of fission. In the present study, we demonstrated that 

inbreeding of the triploid planarian D. ryukyuensis resulted in both 

diploid and triploid offspring in nature. In the triploids of D. 

ryukyuensis, chiasmata between homologous chromosomes were 

observed in both female and male germ lines. This result suggests 

that both diploid and triploid offspring of this species are produced 

bisexually by zygotic fusion between sperm and eggs. Hence, this 

phenomenon may be a novel mechanism in planarian for escaping 

the triploid state. 

Males in a thelytokous strain of tardigrade 

Atsushi C. Suzuki 

Department of Biology, Keio University, Hiyoshi, Yokohama 223- 

8521, Japan 

Tardigrades are generally gonochoristic and propagate by sexual 

reproduction. A number of tardigrade species exhibit secondary 

sexual characters, with males in some Eutardigrada being 

distinguished by modified claws. Many moss-dwelling tardigrades, 

however, have thelytokous mode of parthenogenesis. One of such 

tardigrade is Milnesium tardigradum. Males in this species with 

modified claws are usually rare and many populations appear to 

have only parthenogenetic reproduction. The presence of males in 

small numbers, being greatly different from the sex ratio of 1:1, had 

not been considered as heterogony so far. Instead, it was interpreted 

to be the result of mixed populations that have parthenogenetic and 

amphimictic reproduction. Here the author reports that males have 

emerged at a very low frequency in a thelytokous strain of Milnesium 

cf. tardigradum, which has been maintained since 2000. Some 

individuals of this strain had modified claws characteristic of males 

on the first pair of legs. A small testis filling with many spermatozoa 

was observed in one of such specimen. Although copulation has not 

been observed yet, an interesting activity of a male against a female 

appeared to be a sexual behaviour. The frequency of the emergence 

of males in the cultured strain is so low that any environmental factor 

that may generate males has not been determined yet. It is unknown 

if these males could actually function as males in reproduction. 

However, they might show some possibility of genetic exchange 

among the clonal populations.


Insights into evolutionary ecology of insect immunity 

Boran Altincicek and Andreas Vilcinskas 

Interdisciplinary Research Center, Institute of Phytopathology and 

Applied Zoology, Justus-Liebig-University of Giessen, Heinrich-Buff- 

Ring 26-32, D-35392 Giessen, Germany 

The evolutionary success of insects regarding diversity in species 

and ecological niches reflects their prominent ability to control a wide 

array of pathogens. Here, we have identified immune-inducible 

genes in phylogenetically distant insects using subtractive 

suppression hybridization. Results from the apterygote Thermobia 

domestica, the aphid Acyrthosiphon pisum, the drone fly Eristalis 

tenax, and the genetically tractable model beetle Tribolium 

castaneum revealed the immune induced expression of genes 

encoding proteins involved in signaling, defense mechanisms, stress 

response, and cellular homeostasis. However, we also noted 

differences, especially on antimicrobial peptides (AMPs). We 

identified thaumatins in Tribolium (ancient antifungal plant peptides) 

that are absent from most other insects and numerous putative E. 

tenax specific AMPs. In the aphid we have found lysozyme activity 

but no detectable inhibitory activities against live bacteria. We 

identified a lysozyme gene but no homologues of known AMPs in our 

cDNAs or in 90.000 public sequences. Instead, we discovered that 

viviparous offspring generation was accelerated upon wounding. We 

argue that aphids increase terminal reproductive investment and limit 

antibacterial defense in response to a survival threat. Obtained 

results suggest an ancestral complexity of insect immunity and that 

ecology and pathogens are likely to have had a particularly important 

role in the diversification of the insect immune system. 

Autoimmunity : mechanisms of breakdown and restoration of 

self 

Jean-François Bach 

Académie des Sciences, 23 quai de Conti, 75006, Paris, France 

Autoreactive B and T cells with high affinity receptors for auto 

antigens are negatively selected during ontogenesis. However, 

normal individuals harbour low affinity autoreactive B and T cells. A 

central question of autoimmunity is to determine the mechanisms by 

which autoreactive T cells may become pathogenic after breakdow! 

n of the operational self tolerance which prevails in healthy subjects. 

Initial studies indicated an important role for anergy, an antigenspecific, 

silencing of auto-reactive T cells. Recent data suggest a 

more important role for regulatory T cells. It is not yet determined, 

however, whether the onset of autoimmune diseases is due to the 

decline of regulatory T cell function or number has documented in 

some experimental models such as day 3 thymectomy in mice or to 

progressively acquired resistance of effecter cells to regulatory T 

cells. The problem is complicated by the multiplicity of regulatory T 

cell subsets. These considerations pave the way to a number of 

therapeutic approaches aiming at restoring self tolerance in subjects 

with autoimmune diseases, as recently achieved using anti-CD3 

monoclonal antibodies. 

S13 - Comparative Immunology 

- 45 - 

Comparative Immunology 

Edwin L. Cooper 

Laboratory of Comparative Immunology Department of Neurobiology 

David Geffen School of Medicine at UCLA University of California at 

Los Angeles Los Angeles California 90095-1763 

Phagocytosis in unicellular animals represents the most ancient and 

ubiquitous form of defense against foreign material. Unicellular 

invertebrates can phagocytose for food and defense. Multicellular 

invertebrates and vertebrates possess phagocytic cells and have 

evolved more complex functions attributed to immuno-defense cells 

that specialized into cellular and humoral immune responses. Thus 

invertebrates and vertebrates possess: innate, natural, non-specific 

(no memory) non-anticipatory, non-clonal, germ line (hard wired); 

vertebrates possess: adaptive, induced, specific (memory), 

anticipatory, clonal, somatic (flexible). The early expression of 

proteins involved in cell interaction in metazoa demonstrates that 

these proteins evolved before the origin of animals and were later coopted 

for development. A similar situation exists with respect to 

components of the signaling system, immunity and development. 

With multicellularity, clearly numerous immune response 

characteristics are not possible in unicellular forms or even those 

that straddle the divide between unicellularity and multicellularity, 

beginning with colonial/social protozoans. Still it is instructive to 

elucidate a hierarchy of animals based upon immunologic 

characteristics and how they parallel other physiological traits. 

Evidence is presented that the most primitive of invertebrates prior to 

the evolution of multicellullar organisms possess varying degrees of 

complexity at the molecular level of those hallmarks that now 

characterize the immune system. It is at the level of the annelids, 

notably the earthworm where phagocytosis and the precursors of 

natural killer cell activity are first dissociated. This negates the 

commonly held view that all invertebrates are only capable of 

phagocytic responses –their single innate system. 

Evolution of Adaptive Immune Systems in Vertebrates 

Max D. Cooper 

Emory University, Department of Pathology and Laboratory Medicine, 

1462 Clifton Road NE, DSB, Room 403C, Atlanta, Georgia 30322, 

USA 

Two surprisingly different types of adaptive immune systems have 

evolved in vertebrates. The surviving jawless vertebrates, lamprey 

and hagfish, have a lymphocyte antigen receptor repertoire that is 

potentially as large as that of the antibody repertoire (>10 14 ) in mice 

and humans. However, unlike our Ig-based T cell receptors and B 

cell receptors for antigens, the agnathan variable lymphocyte 

receptors (VLR) are composed of multiple leucine rich repeats (LRR) 

and an invariant stalk region tethered via glycosyl-phosphatidylinositol 

anchorage to the lymphocyte surface. The diverse VLR 

genes are generated by a gene conversion mechanism for a 

multistep, piecewise assembly process in which flanking LRR 

sequences are stitched into an incomplete germline VLR gene. The 

VLR diversity is based upon differences in sequences and numbers 

of constituent LRR modules. Monoallelic assembly and expression of 

unique VLR genes by individual cells results in the generation of a 

clonally diverse lymphocyte population, members of which can be 

selected for specific immune responses. Immunized lamprey 

undergo antigen induced lymphocyte activation, proliferation and 

differentiation to produce multivalent VLR antibodies with precise 

specificity for protein and carbohydrate antigenic determinants on 

bacteria, viruses, and mammalian blood cells. The secreted lamprey 

antibodies are composed of eight or ten identical subunits held 

together at their base by disulfide bonds. Remarkable antigen 

specificity, avidity, stability and ease of molecular engineering 

suggest many potential biomedical uses for monoclonal VLR 

antibodies.


Diversification of the immunoglobulin superfamily receptors in 

the immune systems of Metazoa 

Louis Du Pasquier 

Institute of Zoology and Evolutionary Biology. University of 

Basel ,Vesalgasse 1, CH-4051 Basel, Switzerland 

Restricted to the animal kingdom, the immunoglobulin domains build 

up the most diversified family of leukocyte receptors involved in 

immunity (immunoglobulin superfamily: IgSF). Some of them 

regulate lymphocyte functions, other recognize specifically the 

antigens. The conservation of regulatory receptors controlling the 

proliferation, migration, activation and inhibition of leukocytes, is 

visible at the structural, signalling and chromosomal localization level. 

The case of leukocyte receptor complex–like molecules will be 

presented in arthropods, urochordates, birds and mammals. Some Ig 

domains were recruited as part of different types of variable antigen 

receptors, selection acting on different segments of the domain 

susceptible to bind a ligand. Such receptor genes were duplicated, 

building multigene families .In some species they were made 

susceptible to be diversified by somatic processes using different 

convergent ways. Whether at the DNA or the RNA level these 

mechanisms can randomly diversify the variable domain sequence, 

the isotype composition and the signalling capacities of the receptors 

and this during the life of an individual or during the course of an 

immune response, Example will be given in arthropods, molluscs 

and vertebrates. Relatives of the enzymes RAG and AID that control 

rearrangement, gene conversion, heavy chain class switch and 

somatic hypermutation in gnathostomes exist in some protostomes 

(RAG) or agnathans (AID). The questions arise whether all of this 

potentially very large and randomly acquired diversity is expressed, 

useful and how autoimmunity is avoided. It is a matter of selection, 

the evolutionary aspects of which are not well understood. 

Sexual chemical cues enhance non-specific immunity in male 

mice 

Ekaterina Litvinova 1 , Elena Goncharova 2 , Andrew Garms 1 and 

Michail Moshkin 1,3 

1 Institute of Systematics and Ecology of Animals, Siberian Division 

of the Russian Academy of Sciences, Novosibirsk, Frunze st., 11, 

Russia, 

2 Institute of Chemical Biology and Fundamental Medicine, Siberian 

Division of the Russian Academy of Sciences, Novosibirsk, Russia 

3 Institute of Cytology and Genetics, Siberian Division of the Russian 

Academy of Sciences, Novosibirsk, Russia 

Scent of females as signal to reproduction stimulates male mice to 

search of a potential breeding partner that is coupled with infection 

risk due to: the skin wound as a result of intermale aggressiveness, 

sexual contacts with infected partners, or an inspection of their 

marks, which are harbored a variety of pathogens. We hypothesized 

scent induced immunoredistribution in favour of non-specific immune 

defense against infection risks related with breeding behavior. 

According to the hypothesis female scent triggers migration of 

leucocytes from blood to peripheral areas, in particularly to skin for 

resistance to wounding, and to airways for resistance to viral and 

bacterial agents. Evidences of this idea we have got in study on 

mature male mice that were kept in two different conditions – with or 

without female scent. Histological study of lungs showed intervention 

of leukocytes to perivenous area that was higher in male mice, which 

have got daily new portion of soiled bedding from female cages or 

one intranasal application of fresh female’s urine, in comparison with 

males, which were kept without female scent. Scent induced 

reallocation of leucocytes to upper airways enhanced stress-reaction 

to intranasal application of LPS and increased resistance to 

respiratory infection. We have found 80% surviving of scent treated 

male mice after intranasal infection with murine flu virus (A/WSN/33). 

It was significantly higher (א 2 =6.7; P


Central role for venom in predation by the komodo dragon 

and the giant extinct megalania 

Bryan Fry 

Bio21 Institute, University of Melbourne, 3010, Parkville, Victoria, 

Australia 

It has previously been argued that the Komodo dragon, Varanus 

komodoensis, evolved to prey upon pygmy elephants (Stegodon 

sondaari) and other large preys, but it remains unclear how this 

might have been achieved. Here we investigate three possible 

mechanisms and evaluate their likely role in the predatory ecology 

of V. komodensis; (a) a powerful bite-force, (b) hypertoxic bacteria 

and (c) envenomation. Using three-dimensional (3D) computer 

modelling we show that bite force in V. komodoensis is particularly 

weak, p! recluding the use of a bone-crushing action typical of 

mammalian carnivores. We also find no convincing evidence for 

the role of pathogenic bacteria in prey capture and reject the 

notion that V. komodoensis utilises toxic, bacteria-loaded saliva as 

part of its predatory ecology. We demonstrate here that V. 

komodoensis possesses a highly complex venom system, with the 

capacity to induce coagulopathic and shock-inducing hypotensive 

bioactivity, and venom volumes sufficient to immobilise large prey 

such as pygmy elephants. We further propose that the much larger 

and related Australian fossil varanid, Varanus (Megalania) prisca, 

was also a venomous predator capable of dispatching all but the 

largest Australian Pleistocene megafauna. Such extraordinary 

animals may well have had a role in shaping Aboriginal cultures as 

well as that of Homo floresiensis. 

Does the venomous function originally derive from the innate 

immunity? 

Max Goyffon 1 and Grazyna Faure 2 

1 Muséum National d'Histoire naturelle, Paris 

2 Institut Pasteur, Paris 

Most of the venom toxins are peptides or proteins. In the buthid 

scorpions, the structure of the venom neurotoxins is closely related 

to the structure of circulating defensins and other arthropod 

defensins, with the existence of a consensus sequence containing 

three disulfide bridges (Cociancich et al., 1993). Defensins, as 

other antibacterial peptides, are considered as ancestral molecules 

possessing conservative structures which are present in all living 

animals. On the other hand, scorpions are among the most ancient 

terrestrial arthropods and the composition of the buthid venoms is 

relatively simple and devoid of enzymes with toxic activity. A 

recent work (Whittington et al., 2008) describes the presence of 

three genes coding for venom defensin-like peptides in platypus 

(Ornithorhynchus anatinus) which are the main components of its 

venom. It was also observed that snake venoms contain similar 

toxins (crotamine and crotamine-like peptides). Because platypus 

are considered as primitive mammals in vertebrates, the authors 

concluded that a convergent evolution in venoms has repeatedly 

selected genes coding for proteins containing specific structural 

motifs which can be used as templates for venom toxins. In a work 

on the molecular origin and evolution of the snake venom 

proteome, Fry (2005) has shown the presence of defensins and 

lectins, and noted the importance of the cystein cross-linked 

ancestral proteins in the composition of snake venoms. Besides, 

the natural circulating inhibitors which protect the snakes against 

their own venoms are now considered as proteins of the innate 

immunity (Faure, 2000 ; Perales & Domont, 2002), due to their 

structural homologies with other proteins of the innate immune 

system. Thus, the close relationship of immune proteins and 

venom toxins as well as of immune proteins and natural inhibitors 

which neutralise neurotoxins is clearly evident (Kaplan, 2007; 

Faure & Goyffon, 2008). We consider in consequence that the 

conjunction of these results strongly support the hypothesis that 

the venomous function has originally derived from the innate 

immune function. 

S14 - Venomous animals and their venoms 

- 47 - 

The diversity of venoms and toxins. Marine animals as drug 

providers 

Dietrich Mebs 

Zentrum der Rechtsmedizin, University of Frankfurt, Kennedyallee 

104, D-60596 Frankfurt, Germany, 

In animals the production of biologically active compounds is 

performed either by protein synthesis or through complex 

metabolic pathways leading to secondary metabolites. Peptide and 

protein toxins are secreted by exocrine glands and are applied by 

various injecting devices for prey acquisition or in defense. Other 

compounds are synthesized in various organs and are used to 

prevent infection, overgrowth or as defensive agents. They may 

also be obtained from the diet, stored in the body and sequestered 

via the food chain. 

Animals are most efficient drug developers. Exploring their toxin 

arsenal provides new structures and compounds for curing human 

ailment and diseases. Sponges and other marine invertebrates 

contain a huge array of new entities with mostly unknown activities. 

Marine snails feed on sponges and incorporate the toxic 

metabolites using them in their own defense. Conus snails contain 

in their venom a huge diversity of peptides. Mutations and 

posttranslational modifications make these toxin mixtures highly 

diverse and complex. Selection pressure and adaptation to special 

needs shape the structure and specificity of the peptides for 

certain targets such as receptors and ion-channels. In a process of 

accelerated evolution these compounds are perfectly designed 

and surpass any synthetic product. Exploration and exploitation of 

biodiversity for health is an exciting adventure and a most 

promising enterprise. 

Venomics, the venomous system genome project 

André Ménez 1 †, Dietrich Mebs 2 , Reto Stöcklin 3 

1 

Muséum National d´Histoire Naturelle, 57 Rue Cuvier, 75005 

Paris, France 

2 

Zentrum der Rechtsmedizin, University of Frankfurt, 

Kennedyallee 104, D-60596 Frankfurt, Germany 

3 

Atheris Laboratories, case postale 314, CH-1233 Bernex-Geneva, 

Switzerland. 

Sponsored by the European Union, the new Venomics project 

(CONCO) aims to study the genetics, transcriptomics and 

proteomics of the evolutionary tripartite combination: animal, 

venomous system and toxins. Using marine snails of the genus 

Conus as typical representatives of venomous animals, the project 

is anticipated to clarify these fundamental aspects and their 

association with evolutionary processes. Practical consequences 

of this research include the discovery of novel biopharmaceuticals 

in the venom of cone snails. The genome and transcriptome of one 

cone snail species will be exhaustively studied, venom will be 

fractionated and submitted to proteomic analysis to generate a 

“natural library” of compounds, and peptides will be synthesized 

forming the basis of a “synthetic library”. The biological activity of 

peptides from these two libraries are being tested on a variety of 

physiological targets, selected peptides with potential therapeutic 

value are further characterized in vivo. Moreover, the biodiversity, 

ecology and molecular evolution of a wide range of Conus species 

are studied. This first Venomics project involves 18 European 

laboratories, the J.Craig Venter Institute (USA) and the non-forprofit 

Toxinomics Foundation, which has been created to 

synchronize the activities of the groups.


Evolutionary and ecological relationships of the venoms of 

the related coral snakes Micrurus dissoleucus and Micrurus 

mipartitus 

Camila Renjifo 1 , Alain Riveros 2 , Armando Sanchez 2 , Juan Manuel 

Renjifo 3 , Henry Aceros 2 , Darío Riascos 2 , Jairo Maldonado and 

Gabriel Pascual 

1 Pontificia Universidad Javeriana, Bogotá-Colombia Facultad de 

Ciencias Básicas, Departamento de Biología, Colombia 

2 Pontificia Universidad Javeriana, Bogotá-Colombia, Facultad de 

Medicina, Departamento de ciencias fisiológicas, Colombia 

3 Universidad del Magdalena, Santa Marta-Colombia, Facultad de 

Ciencias, Departamento de Biología, Colombia 

The venoms of coral snakes (genus Micrurus) clinically produce 

flaccid paralysis; the high-rate of mortality results from respiratory 

failure. While studies have investigated potential human effects, 

the neurotoxic effects have been poorly investigated in terms of 

venom evolutionary relationships. The purpose of this study was to 

investigate the physiological effects on neuromuscular junction of 

the venom of the closely related species Micrurus dissoleucus and 

Micrurus mipartitus to be able to compare their venoms in terms of 

ecology. While both species are likely to be specialist-feeders on 

reptiles, they occupy very different habitats. M. dissoleucus lives 

mainly in xeric to semiarid or seasonal dry regions while M. 

mipartitus is found in a wide range of habitats including lower 

montane wet forest and cloud forest. We examined the 

neurotoxicity of M. dissoleucus and M. mipartitus venoms in chick 

biventer cervicis muscle preparations and the venoms were also 

compared by mass spectrometry. M. dissoleucus and M. mipartitus 

venom produced a progressive decrease in the amplitude of 

miniature end-plate potentials, with the indirect stimuli, until these 

were abolished, both venoms significantly inhibited contractile 

responses to the exogenous nicotinic agonists (i.e. ACh and CCh) 

but not KCl, showing mainly a post synaptic effect with different 

doses and times of blocking. These results allow for a relation of 

toxicity to ecology and shed additional light on the forces driving 

venom evolution. 

Evolution of venomous reptiles 

Nicolas Vidal 

UMR 7138, Systématique, Evolution, Adaptation, Département 

Systématique et Evolution, C.P. 26, Muséum National d’Histoire 

Naturelle, 43 Rue Cuvier, Paris 75005, France 

The evolution of the venomous function is considered to be a key 

innovation driving ecological diversification in advanced snakes. 

Recent phylogenetic, toxicological, and histological results 

demonstrate a single early origin of venom in squamate reptiles in 

the Jurassic that may also have been a key factor in the adaptive 

radiation and subsequent ecological success of several lizard 

lineages. The well-supported anguimorph/iguanian/snake clade, 

named Toxicofera, represent ~4600 out of ~7900 extant squamate 

species, or 58 % of the total squamate species diversity. These 

results provide new insights into the evolution of the venom system 

in squamate reptiles and open additional new avenues for 

biomedical research and drug design using hitherto unexplored 

venom proteins. Among snakes, the caenophidian venom 

apparatus has experienced extensive evolutionary tinkering 

throughout its history. All traits, ranging from biochemical 

(specialization of the venoms) to dentition and glandular 

morphology, have changed independently, resulting in many kinds 

of toxins and diverse delivery systems. Rear-fanged—or more 

correctly defined, non front-fanged—caenophidians possess 

complex venoms containing multiple toxin types, while the frontfanged 

venom system appeared three times independently: once 

early in caenophidian evolution with viperids, once within 

atractaspidines, and once with elapids. Further a reduction of the 

venom system is observed in species in which constriction has 

been secondarily evolved as the preferred method of prey capture 

or dietary preference has switched from live prey to eggs or to 

slugs and snails. 

- 48 - 

Variation in snake venom composition; evidence for natural 

selection and adaptation to diet 

Wolfgang Wüster, Axel Barlow, Catharine E. Pook and Nicholas R. 

Casewell 

Bangor University, School of Biological Sciences, Bangor LL57 

2UW, UK 

Variation in venom composition in snakes is a ubiquitous 

phenomenon at all taxonomic levels. The causes of this variation 

have been subject to considerable debates, especially at low 

taxonomic levels (within species and between closely related 

species). Some authors have argued for the importance of natural 

selection for different prey items, whereas others suggest that the 

high lethal potential of most snakes makes natural selection for 

different prey types unlikely. 

This lecture will summarise the available evidence on causes of 

venom variation in snakes. There is ample evidence for the likely 

role of natural selection from correlation between variation in 

venom composition and diet, examples of high levels of resistance 

to venom in some prey animals, the reduction in venomous 

function in snakes feeding on undefended prey, and apparent 

specific action of venom against the main prey of the snakes. 

This information is complemented with novel data on variation in 

venom composition, gene expression and diet in saw-scaled vipers 

(Echis): the four main species groups in the genus differ 

profoundly in the proportion of arthropods in the diet. Venom 

toxicity to scorpions reflects the importance of arthropods in the 

diet of the species groups, and gene expression patterns reflect 

adaptation of venom composition to physiologically highly 

divergent prey types.


MEMRI of the olfactory pathway into the frog brain Xenopus 

laevis 

Renaud Boistel 1,4 , Brigitte Gillet 2 , Christelle PO 2 , Anthony 

Sébillot 3 , Peter Cloetens 4 , André Mazabraud 5 , Nicolas Pollet 5 

S15 - Integrative biology of acoustic communication 

1 Muséum National d’Histoire Naturelle, UMR 7179 Mécanismes 

Adaptatifs : des Organismes aux Communautés, Département 

Ecologie et Gestion de la Biodiversité, 57, rue Cuvier CP55, F- 


2 Laboratoire de RMN Biologique, ICSN, CNRS, Gif-sur-Yvette, 

France 

3 WatchFrog S.A.S. RCS Evry 484 872 551, Code APE 731 Z 

4 European Synchrotron Radiation Facility,, B.P. 220, F-38043 

Grenoble, France 

5 CNRS UMR 8080, Bât. 445, Université Paris XI, 91405 Orsay, 

France 

In terrestrial frogs, metamorphosis involves a shift in olfaction from 

sensing chemicals in water to detecting airborne odours. Some 

frogs such as Xenopus have an aquatic life and the structure of 

their olfactory organ is therefore the subject of considerable 

interest as genetic models. In adult Xenopus, the principal cavity is 

always filled with air, even when the frog is submerged. On the 

other hand, the vomeronasal organ is filled with fluid throughout 

life. Xenopus is reported to be able to find its way to new ponds 

over distances of several kilometres. The only available cue seems 

to be olfaction. This is supported by Du Plessis (1966) who 

suggested that one function of smelling airborne odours may be in 

locating pond habitats. Knowledge on the detection of odours from 

the air is lacking. We aim to investigate airborne olfaction in 

Xenopus laevis in vivo by a novel method, Manganese-enhanced 

Magnetic Resonance Imaging (MEMRI). We obtained a good T1 

contrast in the afferent projection and the olfactory bulb during time. 

MEMRI proves to be useful and offers new perspectives as a 

molecular imaging method to visualize functional neural circuits in 

the brain in vivo. 

Vocal communication in groups of neighbouring skylarks 

(Alauda arvensis): syntax always matters? 

Elodie Briefer, Fanny Rybak and Thierry Aubin 

Université Paris 11, NAMC, CNRS-UMR8620, Bioacoustics Team, 

Bat. 446, 91405 Orsay cedex, France 

The skylark is a territorial species of open landscape in which pairs 

settle in stable territories during the breeding season. Due to the 

heterogeneity of the habitat, territories are gathered in patches 

spaced by few kilometres. Males produce complex songs as a part 

of the territorial behaviour. We showed that, in a given patch, 

males (neighbours) share several sequences of syllables in their 

songs, whereas males settled in different patches (strangers) have 

no sequences in common. To test the hypothesis that these 

shared sequences support a group signature, we made playback 

experiments with ‘chimeric’ signals: songs of strangers where the 

sequences shared by neighbours were artificially inserted. 

Behavioural responses to playbacks indicated a reduced 

aggression toward neighbour songs compared to stranger ones. 

Furthermore, the same level of responses, observed when a 

‘chimeric’ and a neighbour song were broadcast, indicated that 

shared sequences are recognised as markers of the 

neighbourhood identity. To test neighbour recognition, further 

playback experiments were performed. Neighbour and stranger 

songs were broadcast from the territory boundary shared by the 

subject and its adjacent neighbour whose song was played back, 

and from the opposite boundary. Subjects exhibit a lower 

aggression towards neighbour songs at the shared boundary but 

display the same level of aggression to the two categories of 

songs at the opposite one. Thus, males showed a spatial 

categorisation of their proximate neighbours. Acoustic analyses 

revealed that individual information is potentially carried by the 

syllable organisation. Thus, the syntax may code for distinct 

messages and different identities. 

- 49 - 

Effects of fish size on agonistic sounds in the clownfish 

Amphiprion akallopisos: implications for the role of the 

swimbladder in the sound production mechanism. 

Orphal Colleye 1 , Pierre Vandewalle 1 , Bruno Frédérich 1 , Margarida 

Casadevall 2 and Eric Parmentier 1 

1 Laboratoire de Morphologie Fonctionnelle et Evolutive, 

Département des Sciences et Gestion de l’Environnement, 

Université de Liège, Institut de Chimie, Bât B6c, 4000 Liège, 

Belgium 

2 Unitat de Biologia Animal, Facultat de Ciències, Universitat de 

Girona, Campus de Montilivi s/n, 17071 Girona, Spain 

Clownfishes (i.e. Amphiprion akallopisos) are territorial fishes that 

use sound production to defend their anemone territory. In this 

context, agonistic interactions are frequent and are often 

associated with sounds that are called “pops”. In sonic fishes, the 

swimbladder is often involved in sound production, mainly acting 

as resonance chamber. Currently, the potential role of the 

swimbladder in sound production has not been demonstrated yet 

in clownfishes. 

In the present study, we thus compared the sonic features 

between individuals of different sizes and in different sexual status 

(juvenile, male and female). The role of the swimbladder in sound 

production was also tested using an experimental filling with 

physiological liquid. 

In A. akallopisos, fish size is highly correlated with swimbladder 

size. Sound analyses showed that dominant frequency and pulse 

duration are highly related to fish size (r² = 0.95). No variations in 

sounds are related to sex. The experimental filling of the 

swimbladder with physiological liquid significantly modified the 

acoustic features. 

In A. akallopisos, frequency and pulse duration are directly related 

to fish size, and thus to swimbladder size. This study highlights the 

role of the swimbladder in the sound production mechanism in 

clownfishes, acting as a resonance chamber. 

Vocal competition between two hybridizing seabird species 

Charlotte Curé 1,2 , Nicolas Mathevon 1,2 and Thierry Aubin 1 

1 Equipe ‘Communications acoustiques’, Laboratoire Neurobiologie, 

de l’Apprentissage, de la Mémoire et de la Communication UMR 

CNRS 8620, Université Paris-Sud, F-91405 Orsay cedex, France. 

2 Laboratoire d’Ecologie et Neuro-Ethologie Sensorielles EA3988, 

Université Jean Monnet, F-42023 Saint-Etienne cedex 2, France. 

Whereas the impact of sexual selection on the evolution of vocal 

signals has been extensively studied in song-learning birds, it has 

been so far largely neglected in non song-birds, especially the 

intra-sex component of sexual selection. Using the opportunity 

offered by shearwaters where mates take turns to incubate and 

specifically interact with same-sex individuals during nest defence, 

we investigated the relationship between intra-sex vocal 

interactions and hybridization in two allopatric closely related 

species, the Yelkouan and the Balearic shearwaters (Puffinus 

yelkouan and P. mauretanicus). Acoustic analysis of the territorial 

call recorded in the allopatric populations of each species revealed 

that this call carries both reliable sex- and species-specific 

signatures. Calls of hybrid birds showed an intermediate structure 

to those of parental species, underlining the genetic basis of 

vocalization structure. While birds from the hybridization zone and 

from the Yelkouan population responded equally to playback of 

both parental vocalizations, the Balearic reacted less strongly to 

heterospecific calls than to conspecific calls. This asymmetry 

between both parental species in intra-sex competition may have 

important fitness consequences, in that Yelkouan individuals are 

more likely to introgress into Balearic populations than the reverse. 

Moreover, as Balearic males less readily respond during intra-sex 

interaction with Yelkouan individuals than females do, Yelkouan 

introgression into the Balearic population may be primarily driven 

by males.


Temporal and amplitude parameters affecting mate choice 

selectivity in Nezara viridula (L.) (Heteroptera: Pentatomidae). 

Maarten de Groot, Andrej Čokl and Meta Virant-Doberlet 

Department of Entomology, National Institute of Biology, Večna pot 

111, Ljubljana, Slovenia 

While on the same plant, males and females of the southern green 

stink bug Nezara viridula use species and sex specific vibrational 

signals transmitted through the substrate for recognition and 

localization of conspecifics. We investigated the selectivity of the 

behavioural responses of males to natural and altered female 

calling song (FCS) in one- and two-side play-back experiments on 

bean plants. We tested the influence of two temporal parameters 

(signal and interval duration) and signal amplitude on triggering of 

recognition (male calling) and searching. For all altered FCSs, 

significantly more males were calling than searching. The results 

show that the signal and interval duration were positively 

correlated with the male calling and searching. In general, in twoside 

playback experiments with natural and altered FCSs of equal 

amplitude, more males responded with calling and searching 

behaviour than in one-side stimulation experiments. In this 

situation only signals with shorter than normal interval induced less 

response, while duration of the signal had no effect on male 

responsiveness. In two-side playback experiments with signals of 

different amplitudes, more males were searching when stimulated 

bilaterally by natural FCS. Fewer males were calling and searching 

when altered FCSs were played back simultaneously. Results 

indicate that males’ behavioural responses decreased in line with 

the decrease of natural FCS amplitude. In conclusion, males are 

able to discriminate between natural and altered FCS when played 

simultaneously. Furthermore, the structural and intensity 

differences of the FCS are important factors which underlie the 

complex mate searching behaviour of N. viridula males. 

Phenotypic plasticity and flexibility in the crowing behaviour 

of a vocal non-learner species: the domestic Japanese quail 

(Coturnix c. japonica) 

Sébastien Derégnaucourt and Manfred Gahr 

Max Planck Institute for Ornithology, Postfach 1564, D82305 

Starnberg (Seewiesen), Germany 

Previous studies have shown that species-specific behaviours 

gradually emerge, via incomplete patterns, to the final complete 

adult form. A classical example in the acoustic domain is birdsong, 

a learned behaviour. In contrast to birdsong development, little is 

known about ontogenetic changes of the vocalizations of vocal 

non-learners species. This was the purpose of our study, using the 

crowing behaviour of the domestic Japanese quail (Coturnix c. 

japonica) as an example. Young males were maintained in social 

isolation from the age of 3 weeks to 4 months, and their complete 

crowing activity was continuously recorded. We observed 

developmental changes in crow structure, both at the temporal and 

at the spectral levels. Speed and trajectories of these 

developmental changes did exhibit an unexpected high interindividual 

variability. These ontogenetic changes present some 

similarity to that of song in songbirds, both in general 

developmental changes in vocalization structure and in plasticity of 

form development. We also observed some daily changes in the 

temporal pattern of the crow: crows emitted at night were longer 

than crows emitted during the day. Such vocal changes were also 

observed when quails were transferred from a regular light-dark 

cycle to constant light: crows emitted in constant light were shorter 

than crows emitted in a normal light-dark cycle. These results 

suggest that, like in songbirds, melatonin, which is produced in 

darkness, might affect the temporal pattern of birds’ vocalizations. 

Studies on vocal non-learners could shed light on the specificity 

and evolution of vocal learning. 

- 50 - 

Mechanisms of avian songs and calls 

Ole Næsbye Larsen 

Institute of Biology, University of Southern Denmark, DK-5230 

Odense M, Denmark 

The avian vocal organ, the syrinx, is a specialized structure 

located rather inaccessibly in an air sac close to the heart where 

the trachea bifurcates into the two primary bronchi. The syrinx of 

different avian taxa varies so much in position and morphology that 

it has been used for taxonomy. It consists of a skeletal framework, 

flexible membranes or soft tissue masses, labia, stretched 

between elements of this framework, and the syringeal muscles. 

Until a decade ago most of our knowledge about syringeal 

mechanics was based on such indirect evidence as 

electromyography, emitted sound, and anatomy. The use of thin, 

flexible endoscopes has made direct observation of the syrinx 

possible in situ. The effects of direct muscle stimulation on the 

syringeal aperture have identified adductor and abductor muscles, 

confirming results from electromyographic studies. Endoscopic 

observations have revealed the dynamics of syringeal 

reconfiguration during phonation, which in most bird species 

investigated results in simultaneous movement of soft tissue 

masses (the medial and lateral labia in songbirds and lateral 

tympaniform membranes in non-songbirds) into the bronchial 

lumen where they collide. High-speed video-filming during sound 

production has revealed that sound pulses coincide with short 

duration formation of slots between the soft tissue masses forming 

a pneumatic valve, which suggests that the avian sound 

generating mechanism is a similar to that in the human larynx. 

Lately studies have revealed surprising properties of the syringeal 

muscles and physical models of the syrinx have given us new 

insight into the workings of this fascinating organ. 

Testosterone early treatment affects production of song 

learning in male zebra finches 

Albertine Leitão and Manfred Gahr 

Max Planck Institute for Ornithology, Eberhard-Gwinner-Str. 82319 

Seewiesen, Germany 

Songbirds as well as humans exhibit vocal learning. In male zebra 

finches the sensory and motor phases overlap. Juveniles start to 

sing and memorize the father’s song from 25 post-hatching days 

(PHD) and they have to be exposed at least ten days with the 

tutor’s song to be able to produce an accurate copy of the tutor, 

when adult ca. 90PHD. Song learning and production are 

controlled by a discrete neural circuit, which undergoes profound 

developmental changes during the time when song is learned. 

First singing starts when synaptic connections between two song 

control areas, HVC and RA, are established, sometime after 

PHD25. To understand how testosterone influences different 

stages of behavioural and neural development, we implanted 

juvenile males at PHD16 with testosterone (T) or placebo (P) 

pellets. We removed the tutor at PHD25 and monitored 

continuously their vocalizations until adulthood. We compared 

song development in T and P groups with birds implanted with 

placebo but that were kept exposed to the tutor’s song until PHD35 

(C, control) and with juveniles implanted with testosterone that 

were raised in acoustic isolation (no tutor). We found that 

administration of T induces a shift in the onset of song production: 

birds implanted with T started to sing at 19PHD, instead of 25 for C 

and P groups. Moreover, while not exposed to the tutor’s song at 

the ‘critical’ period T birds were nonetheless able to learn the 

tutor’s song as C birds did. P birds produced a song similar to 

isolate birds. We discuss these behavioural changes with our 

findings in the organisation of the neural vocal system.


Spectral energy distribution as a tool for determining the 

presence of pups in wolf chorus howls 

Vicente Palacios 1 , Luis Llaneza 1 , Benjamin Dugnol 2 , Carlos 

Fernández 2 , Gonzalo Galiano 2 and Julián Velasco 2 

1 

A.RE.NA. Asesores en Recursos Naturales, S. L., C/ Perpetuo 

Socorro, 12 - entlo B, 27003, Lugo, Spain 

2 

Departamento de Matemáticas, Universidad de Oviedo, C/ Calvo 

Sotelo, 33007, Oviedo, Spain 

Researchers need reliable methods that provide accurate data on 

the distribution, abundance, and population trend of a species in 

order to advise wildlife managers. The number of packs is often 

used as an indicator for determining the status of the wolf, Canis 

lupus, over large areas. As a general rule, only the dominant pair 

in a pack breeds and pup presence is taken as an evidence of the 

existence of a pack. Response to simulated howls is commonly 

used by wolf researchers to locate wolf litters during the summer. 

Nevertheless, acoustic structure of chorus howls is complex and 

discriminating in the field the presence of pups in a chorus is 

sometimes a difficult task due to the highly modulated 

vocalizations emitted, especially as pups grow up. In this study we 

test the potentiality of analysing the spectral energy distribution of 

choruses for distinguishing vocalizations emitted by pups from 

those emitted by adult wolves. We analyzed wolf chorus howls 

recordings with known pack composition. Choruses were divided 

into segments of five seconds of duration. For each segment we 

measured some variables related to the spectral energy 

distribution of the signal and identified the types of vocalizations 

present. Our results show that the frequency at which the 

maximum energy peak occurs could be an important variable for 

developing an easy way to discriminate between adults and pups 

vocalizations. 

- 51 -


S16 - Genes, individuals, societies: current trends in social insects 

Colony relocation in the ant Aphaenogaster senilis: role of 

foragers and reminiscence of acquired experience 

Aurore Avargues-Weber 1,2 and Thibaud Monnin 1 

1 Laboratoire Écologie & Évolution CNRS UMR 7625, Université 

Pierre et Marie Curie, 7 quai Saint Bernard, 75005 Paris, France 

2 New address: Centre de Recherches sur la Cognition Animale, 

CNRS UMR 5169, Université Paul Sabatier, 118 Route de 

Narbonne, 31000 Toulouse, France 

We studied the pattern of colony emigration of the ant 

Aphaenogaster senilis, which reproduces by colony fission, and its 

ability to improve at emigrating through acquired experience. 

Emigrations were triggered in the laboratory by opening the nest 

while providing a new nest 1.6 m away (n=10 colonies). We video 

recorded foragers and nurses entering or exiting this arrival nest. 

Nest relocation was characterized by a synchronised emigration: 

the number of workers present in the arrival nest increased 

following a sigmoid curve. The duration of emigration was 

relatively constant between colonies, while the delay before 

emigrating varied. The queen relocated in the middle of the 

emigration, when many workers were on the move. Six to eleven 

weeks after the emigration, foragers were removed from the 

colonies and a second emigration was triggered. We expected this 

emigration to take longer to be completed, yet we found the 

opposite. This may stem from experience acquired during the first 

emigration, as has been shown in the ant Temnothorax albipennis. 

In A. senilis the experience acquired lasted much longer than in T. 

albipennis, and this may be because T. albipennis nurses are 

passively transported by foragers, while A. senilis nurses actively 

walk to the destination nest, so that only the latter gain experience 

by emigrating. In T. albipennis experienced individuals are old 

foragers, with a short life expectancy, while in A. senilis young 

nurses also are experienced. 

Choosing an appropriate index to construct dominance 

hierarchies in animal societies 

Alok Bang 1 , Sujata Deshpande 1 , Annagiri Sumana 1,2 and 

Raghavendra Gadagkar 1,3 

1 

Centre for Ecological Sciences, Indian Institute of Science, 

Bangalore, India 

2 

Indian Institute of Science Education and Research, IIT 

Kharagpur Extension Centre, Salt Lake City, Kolkata, India 

3 

Evolutionary and Organismal Biology Unit, Jawaharlal Nehru 

Centre for Advanced Scientific Research, Jakkur, Bangalore, India 

There is a vast diversity in methods of computation of dominance 

ranks in behavioural literature, but rarely the rationale behind using 

a particular index has been explained. In this study, we analyzed 

three dominance indices, viz. Frequency-based Index of 

Dominance (FDI), Clutton-Brock et al.’s Index (CBI) and David’s 

Score (DS). Apart from FDI that is proposed by our group to 

calculate dominance ranks in Ropalidia marginata and Ropalidia 

cyathiformis, primitively eusocial paper wasps, the other two 

indices were chosen based on popularity and recommendations by 

other groups. Dominance ranks obtained from the behavioural 

data were subjected to correlation analysis and later, the indices 

were evaluated based on the number of unique ranks (ranks 

without ties) they attributed. We found that the three indices 

attributed similar ranks; however, FDI gave significantly more 

number of unique ranks. Using artificial data sets, each index was 

later tested on two parameters, viz. non-interacting pairs and 

reversals. The three indices attributed similar ranks for artificial 

data sets with varying percentages of non-interacting pairs or pairs 

showing reversals. The numbers of unique ranks given by the 

three indices were not significantly different. At high percentage of 

non-interacting pairs, FDI emerged as the best index, whereas at 

higher reversal rates, DS seemed to be a better index. Our results, 

thus, show that the choice of index to be used should be decided 

depending on the properties of the index as well as the social 

structure and interaction network of the species to which it is 

applied. 

- 52 - 

A multi-scale study of ant distribution in the Pyrenees 

Moutains 

Abel Bernadou 1,2 , Xavier Espadaler 2 , Régis Céréghino 3 and 

Vincent Fourcassié 1 

1 Centre de Recherches sur la Cognition Animale, CNRS UMR 

5169, Université de Toulouse, 118 route de Narbonne, F-31062 

Toulouse Cedex 4, France; 2 Departament de Biologia Animal, de 

Biologia Vegetal i d'Ecologia, Facultat de Ciències, Universitat 

Autònoma de Barcelona, E-08193 Bellaterra, Spain; 3 EcoLab, 

CNRS UMR 5245, Université de Toulouse, 118 Route de 

Narbonne, F-31062 Toulouse Cedex 4, France 

As other mountain ranges in the world, the insect fauna of the 

Pyrenees (SW France) present interesting cases of endemism 

making it an area of great interest for entomologists. Surprisingly 

however, relatively few studies have investigated the ant fauna of 

these mountains. Here we present a survey of the distribution of 

ant species along an altitudinal gradient in two Pyrenean valleys 

located in the central part of the ridge and on the opposite sides of 

the mountain range. The influence of environmental factors on the 

distribution of ant species has traditionally been investigated at two 

widely different scales, either the microhabitat or the regional scale. 

However, this does not tell much about the key parameters 

underlying the distribution of a given species because the factors 

intervening at large scale, e.g. altitude, meteorological conditions, 

can impinge on the factors intervening at small scale, e.g. 

vegetation type. We simultaneously examined the distribution of 

ants at three different scales: large (transect - 2000 m 2 ), 

intermediate (100m 2 centered on the sample point) and small 

(sample point - 1m 2 ). We used a neural network method of 

classification to investigate the relationship between the spatial 

distribution of ants and the factors considered at the three spatial 

scales. The results show that both the factors considered at large 

scale (altitude and sun exposure) and those considered at 

intermediate and small scales (land cover) contribute to explaining 

the differences observed in the distribution of ant species. 

The mechanism of queen selection during queen replacement 

in a fission-performing ant 

Blandine Chéron, Claudie Doums and Thibaud Monnin 

Laboratoire d’Ecologie CNRS UMR 7625, Université Pierre et 

Marie Curie, 7 Quai St-Bernard, 75005 Paris 

Fission is a mode of colony foundation where young queens 

(gynes) mate near the nest, and then leave it with a group of 

workers to found a new colony. This mode of colony founding 

allows for the replacement of the queen by one of her mated 

daughters, so that colonies reproducing by fission are potentially 

immortal. 

The Mediterranean ant Aphaenogaster senilis obligatorily 

reproduces by colony fission and is monogynous (a single queen 

per colony). When a colony loses its queen, several gynes are 

produced, which gives rise to a process of queen selection to 

restore the monogyny. This selection could result from competition 

between queens, queen choice by workers or a combination of 

both. Some characteristics of young queens (e.g. age or weight) 

may correlate with the probability that they will inherit the colony. 

Additionally, in a polyandrous species workers could favour gynes 

which are more related to them (nepotism). 

We developed microsatellite markers for A. senilis and showed 

that queens mate only once. We investigated queen selection 

using orphaned colonies. The mean number of gynes produced 

per colony was two (range 1-4) and the mean age difference 

between two successive gynes was 12 days (range 1-29). We 

observed aggressions from both gynes and workers. The oldest 

gyne was more likely to survive and a hierarchy seemed to be 

established, with the oldest gyne dominating the others. Thus, we 

suggest that during queen replacement in A. senilis, 

supernumerary gynes are produced as an insurance, since they 

are generally produced with some delay and rarely become the 

new queen.


How many nestmates ? A key issue in ants’ social 

organization 

Claire Detrain and Jean-Louis Deneubourg 

Unit of Social Ecology, Université Libre de Bruxelles, 50 Avenur F. 

Roosevelt, B-1050 Brussels, Belgium 

Social insects have evolved means of sharing information about 

many aspects of their everyday life. Each ant behave or make 

decisions based on its own experience but also integrate grouplevel 

information provided directly or indirectly by nestmates. Cues 

are the keystones of this socially acquired information: they are 

namely used by ants as “statistical” tools to assess the occupancy 

level of a location and hence to determine the potential for 

amplification processes and emergence of cooperative behaviour 

(1,2). Cues about nestmate density can be acquired by a direct 

sampling of the others, through antennal contacts and/or food 

exchanges. These direct cues are most useful for work 

organization by providing ants with a precise- albeit local and 

punctual- information about the origin, status or recent activities of 

encountered ants (3). Another efficient way for ants to assess their 

social environment is through indirect cues – through the 

perception of chemical cues passively laid by conspecifics (4). 

Unlike direct encounters, such area marking provides to the 

receiver ant an estimate of average ant density without necessarily 

requiring a time-consuming sampling effort. It is an integrative cue 

resulting from the summation through time of tracks left by 

nesmtates and hence reflecting their activity level in the nest 

surroundings. Furthermore, it appears as a unique way for ants to 

“smell the past” since compounds ratios and concentrations vary 

depending on how intensively and for how long one area has been 

foraged by nestmates. 

(1) Detrain, C. & Deneubourg, J.L (in press). Social cues and 

adaptive foraging strategies in ants. In: Food Exploitation by Social 

Insects: An Ecological, Behavioral, and Theoretical Approach 

(Jarau S. & Hrncir M. eds) 

(2) Detrain C. & Deneubourg J.L. 2006. Self-organization in a 

superorganism: do ants behave like molecules? Physics of life 

Reviews 3:162-187. 

(3) Gordon D.M. & Mehdiabadi N. 1999. Encounter rate and task 

allocation in harvester ants. Behav. Ecol. Sociobiol. 45: 370-377. 

(4) Devigne C., Renon A. & Detrain C. 2004. Out of sight but not 

out of mind: modulation of recruitment according to home range 

marking in ants. Anim. Behav. 67: 1023-1029. 

Using social insects for seed dispersal: the case of 

myrmecochory 

Claire Detrain and Pablo Servigne 

Unit of Social Ecology, Université Libre de Bruxelles, 50 Avenue F. 

Roosevelt, B-1050 Brussels, Belgium 

Ants are one of the rare invertebrate groups that participate into 

seed dispersal. In the first phase of this myrmecochory process, 

ants removed seeds from the parent plant, behave as centralplace 

foragers and bring them back to the nest. We show that this 

centripetal movement of seeds towards the ant nest follows a 

species-specific dynamics. For the two tested plant species 

(Chelidonium majus, Viola odorata, the insectivorous Myrmica 

rubra ants remove seed items in larger number and at higher 

speed than the aphid-tending Lasius niger workers what supports 

the hypothesis of a convergence between odours of elaiosomes 

and insect preys. Within the nest, seed nutritive bodies (i.e. 

elaiosomes) are discarded and eaten by ants or larvae. Then 

begins the second phase of myrmecochory, during which the still 

viable seeds do not interest ants anymore: they are removed from 

the nest and dispersed to outside final locations. We demonstrate 

that this centrifugal movement is also highly specific with Myrmica 

ants rejecting seeds from the nest at quicker rates than Lasius 

workers. Such a difference may be due to the higher propensity of 

insectivorous species to remove waste items due to higher 

sanitary constraints on their nest-confined social life. This raises 

questions on how plant seeds may have evolved ways to be 

attractive to ant species that are the most likely to promote their 

dispersal. 

- 53 - 

How individual foraging behaviour is affected by social 

environment in ants 

Renée Fénéron 1 , Marie-Claire Malherbe 1 , Vincent Fourcassié 2 and 

Stéphane Chameron 1 

1 Laboratoire d’Ethologie Expérimentale et Comparée (CNRS UMR 

7153), Université Paris-Nord, Villetaneuse, France 

2 Centre de Recherche en Cognition Animale (CNRS UMR 5169), 

Université Paul Sabatier, Toulouse, France 

Division of labour amongst eusocial animals implies that most 

individuals contribute energy, time or work capacity rather than 

direct reproduction to their group. In insect societies, workers 

benefit the colony fitness through the altruistic behaviours they 

perform. Among the worker's behavioural tasks, foraging is of 

utmost importance because adult survival and larvae growth 

entirely depend on food provisioning and colony energetic 

reserves. As foragers support a high mortality risk, theories predict 

that colony-level selection should regulate foraging behaviour most 

strictly in social contexts where the loss of foragers is costly. It is 

especially the case when colonies are small and food demands 

high. We then investigated how individual foraging behaviour is 

affected by social environment, namely the numbers of workers 

and larvae, in the ant Ectatomma tuberculatum (Formicidae, 

Ectatomminae). We analysed the behaviour and foraging paths of 

individual foragers in experimental groups varying in worker and 

larvae composition. Rate of foraging was more important in large 

groups but individual foraging effort higher in small ones. Trip 

duration decreased with the amount of larvae, independently of the 

colony size; both the decrease in walking distance, stop numbers 

and feeding duration were responsible for shorter trips. Our results 

complement the numerous studies on optimal foraging, 

demonstrating that foragers through their behaviour and 

movement patterns continue to be influenced by the social 

environment they have temporarily left. Discussion focuses on the 

ants' ability to assess colony needs and the regulation 

mechanisms of task allocation on the basis of threshold models. 

Path efficiency of ants foraging in tunnel networks with 

different branching geometries 

Vincent Fourcassié, Simon Garnier, Aurélie Guérécheau, Christian 

Jost, Grégory Gerbier, Maud Combe and Guy Theraulaz 

Centre de Recherches sur la Cognition Animale, CNRS UMR 5169, 

Université de Toulouse, 118 route de Narbonne, F-31062 

Toulouse Cedex 4, France 

Some species of ants forming large colonies use a system of mass 

chemical recruitment to explore collectively novel areas. Workers 

lay a trail more or less permanently and this quickly leads through 

a process of self-organization to the emergence of a network of 

interconnected trails 1 . When a food source is discovered a 

recruitment trail is established over existing exploratory trails and 

the question arises as to whether ants are able to establish this 

trail along the shortest possible path going from their nest to the 

food source. To answer this question we investigated in the 

Argentine ant Linepithema humile the collective performance of 

workers moving in artificial networks of tunnels in which several 

interconnected paths can be used to reach a single food source. 

We used two networks of same length but differing in the geometry 

of their branching (symmetrical, i.e. in which the angles between 

tunnels is the same, or asymmetrical). For both networks most 

experiments ended with the establishment of the trail along one of 

the shortest path 2 , which shows that ants did not orient randomly in 

the network. Moreover, the traffic was more concentrated along 

the shortest paths in asymmetrical than in symmetrical networks. 

Experiments conducted at the individual level show that ants 

reaching an asymmetrical bifurcation prefer to orient on the branch 

that deviates less from their initial direction. This bias could partly 

explain the results observed at the collective level in the path 

chosen by ants for the establishment of a recruitment trail. 

1 Edelstein-Keshet, L., Watmough, J. & Ermentrout, G. B. 1995. 

Behav Ecol Sociobiol, 36, 119-133. 

2 Vittori, K., Talbot, G., Gautrais, J., Fourcassie, V., Araujo, A. F. R. 

& Theraulaz, G. 2006. J Theoret Biol, 239, 507-515.


What makes a termite queen – from ecology to genes 

Judith Korb 

Behavioral Biology, University of Osnabrück, D-49069 Osnabrück, 

Germany 

Termites are the oldest social insects. Despite their apparent 

similarity to ants, they evolved social life independently, probably 

from a wood-nesting cockroach ancestor. We studied the ultimate 

and proximate factors influencing cooperation in a drywood termite, 

Cryptotermes secundus, with an ancestral life type of dwelling in 

dead wood. Associated with this life type is a flexible development 

in which workers can become sterile soldiers, winged sexuals that 

disperse and found a new colony, or replacement reproductives 

that inherit the natal breeding position when the same sex 

individual of the colony dies. We showed that the ‘decision’ of 

individuals to stay in the colony or leave it as winged sexual is 

influenced by ecological factors, such as food availability or 

parasite pressure, and social factors such as colony size. However, 

it is not affected by the number of young present in the nest. These 

results together with the finding that the workers do not care for 

young, suggest that the workers do not stay to raise siblings. 

Rather they seem to stay because they can inherit the colony as 

replacement reproductive. Proposing that the opportunity to 

become a replacement reproductive builds a major backbone for 

the evolution of termite sociality, we investigated genes involved in 

reproductive development. In a cross-species comparison we 

isolated three genes which were generally highly overexpressed in 

female replacement reproductives. These genes are good 

candidates to play a crucial role in caste determination and 

reproductive division of labor. 

Molecular basis of foraging and defense behaviors in the ant 

Pheidole pallidula 

Christophe Lucas 1,2 and Maria B. Sokolowski 1 

1 Department of Biology, University of Toronto, 3359 Mississauga 

Rd. Mississauga, Ont. L5L 1C6, Canada 

2 present address: Department of Ecology and Evolution, 

University of Lausanne, Biophore, CH-1015 Lausanne, 

Switzerland 

Social insects are spectacular examples of behavioral adaptations 

with sterile sisters performing specialized work in the colony. 

However, little is known about the molecular basis of their 

behavioral specialization and flexibility. 

The ant Pheidole pallidula has two sub-castes of workers, majors 

and minors, which are specialized in defense and foraging 

respectively. However, majors are able to help minors in foraging 

activities depending on the needs of the colony. Thus, there is 

plasticity in the subcaste behavioral repertoires. 

Here we investigate the molecular underpinnings of subcaste 

specialization and plasticity in foraging and defense by studying 

the foraging (for) gene. The behavioral effects of for which 

encodes a cGMP dependent protein kinase (PKG) were first 

described in Drosophila melanogaster, where for allelic variation 

affects foraging behaviors (Osborne et al. 1997). In Apis melifera, 

for expression is different between foragers and nurses (Ben- 

Shahar et al. 2002). Here we show that for is involved in 

behavioral flexibility and specialization of P. pallidula's sub-caste. 

Specifically, majors have higher PKG enzyme activities than 

minors, their PKG activities are lower in the presence of a foraging 

stimulus and higher in the presence of an alien intruder. 

Furthermore, pharmacological activation of PKG decreases 

foraging and increases defense behaviors. Finally, both the 

number and localization of neuron clusters in which FOR-PKG is 

expressed differ between the brains of majors and minors. 

Together these results suggest that the foraging gene modulates 

responses to task-related stimuli in the ant colony. 

Ben-Shahar Y, Robichon A, Sokolowski MB, Robinson GE (2002) 

Influence of gene action across different time scales on behavior. 

Science 296:741-744. 

Osborne K, Robichon A, Burgess E, Butland S, Shaw RA, 

Coulthard A, Pereira HS, Greenspan RJ, Sokolowski MB (1997) 

Natural behavior polymorphism due to a cGMP-dependant protein 

kinase of Drosophila. Science 277:834-836. 

- 54 - 

Evolution of queen size polymorphism in the neotropical ants 

Ectatomma Smith 1958 

Alejandro Nettel 1 , Chantal Poteaux-Leonard 1 , Dominique 

Fresneau 1 and Jean-Paul Lachaud 2 

1 Laboratoire d‘Ethologie Expérimentale et Comparée (LEEC) 

CNRS UMR 7153 - Université Paris 13. 99, avenue J-B. Clément, 

93430, Villetaneuse, France 

2 Centre de Recherches sur la Cognition Animale (CRCA) UMR 

5169 - Université Paul-Sabatier, 118 route de Narbonne, Bât. IVR3, 

31062 Toulouse, France 

The evolutionary implications of ant queen-size polymorphism, a 

phenomenon sparsely present throughout the Formicidae, is a 

central question in the study of the reproductive strategies of social 

insects. This polymorphism entails the presence of normal sized 

queens, macrogynes, and size-reduced queens, microgynes. 

Microgynes have been related to different evolutionary trajectories 

including parasite queens (inquilinism), alternative dispersal 

agents at a lower energetic cost, and dependent colony-founding 

specialists. Parasitic forms are thought to originate from queenpolymorphic 

populations that undergo sympatric speciation. Two 

different instances of microgyny have been described within two 

widely distributed species of the neotropical genus Ectatomma. 

Microgynes in E. tuberculatum are parasites and have recently 

been described as a new species, E. parasiticum. On the other 

hand, evidence suggests that E. ruidum microgynes are a 

dispersal-related morph. These two cases of queen-size 

dimorphism have been described only for colonies from southern 

Mexico. The aim of our study is to understand the evolution of 

microgyny in Ectatomma by answering the questions: Is there 

evidence of a common origin of microgyny in E. ruidum and E. 

tuberculatum? If not, is microgyny a recent, derived characteristic 

within each species? Our results based on the molecular 

phylogeny of the whole Ectatomma genus using mitochondrial and 

nuclear sequences show that microgyny has evolved twice 

independently and that dissimilar processes were involved in the 

origin of mycrogyne-bearing populations in E. ruidum and E. 

tuberculatum. We present our results and their implications to our 

understanding of the evolution of alternative reproductive 

strategies in ants. 

Genomic response to sex: A microarray-based study of 

queens in the ant Cardiocondyla obscurior 

Jan Oettler 1 and John Wang 2 

1 

Universität Regensburg, Biologie I, Germany 

2 

University of Lausanne, Department of Ecology and Evolution, 

Suisse 

Previous research documented a dramatic effect of mating on 

aging in the ant C. obscurior: Queens that have been mated with 

sterilized males show a similar lifespan compared to queens 

mated with fertile males. By contrast, virgin queens have a much 

lower life expectancy (50% lower). We were interested in why 

those queens that were mated with sterilized males still show 

physiological traits of fully functional queens although fecundity is 

as low as in virgin queens, i.e. why does aging appear to be 

uncoupled from fecundity. To begin investigating the molecular 

basis for this uncoupling, we used a cDNA microarray developed 

for the closely related Solenopsis invicta to profiled gene 

expression differences between virgin queens, queens mated to 

normal males and queens mated to sterilized males. We found 

many differentially regulated genes among the three different 

queen classes, and of particular interest in our preliminary analysis, 

some of these genes are involved in metabolism and juvenile 

hormone production.


Mating biology of Leptothorax gredleri (Hymenoptera, 

Formicidae) 

Angelika Oppelt 1 , Klaus Hartfelder 2 and Juergen Heinze 1 

1 

Biologie I, Universität Regensburg,Universitätsstr.31, D-93040 

Regensburg, Germany 

2 

Departamento de Biologia Celular e Molecular e Bioagentes 

Patogênicos, Faculdade de Medicina de Ribeirão Preto, 

Universidade de São Paulo, Brazil 

Leptothorax gredleri is a European ant species, in which female 

sexuals attract males by “female calling" and mating occurs on the 

ground. This allows observing sexual behaviour in flight cages and 

thus investigating the mating biology of ants, which previously has 

often been neglected. We studied sperm transfer, mate choice, 

and cuticular hydrocarbon profiles of sexuals in this species and in 

addition investigated gene expression patterns to learn more about 

the reproductive investment of males. 

A topological approach to the characterisation of termite 

nests 

Andrea Perna 1 , Guy Theraulaz 1 , Pascale Kuntz 2 , Stéphane 

Douady 3 and Christian Jost 1 

1 Centre de Recherches sur la Cognition Animale, CNRS UMR 

5169, Université Paul Sabatier, 118 route de Narbonne, 31062 

Toulouse Cedex 4, France 

2 Ecole Polytechnique de l’Université de Nantes, rue Christian 

Pauc, La Chantrerie, 44306 Nantes, France 

3 Laboratoire Matière et Systèmes Complexes, Bâtiment 

Condorcet, Université Paris Diderot - CC7056, 75205 Paris cedex 

13 France 

Termites build some of the most complex nests observed in the 

animal world. The nests can have elaborate and distinctive forms 

typical of the species that produced them; sometimes, however, 

strong similarities are found between nests built by 

phylogenetically unrelated taxa, suggesting that genetically coded 

behavioural rules strongly interplay with physical constraints 

imposed by the building material and environment. Inside the nests 

of most species are large mazes of interconnected chambers and 

galleries : complex transportation networks spanning the three 

dimensions. 

How can the insects orient themselves inside these networks? Are 

there central points where all the paths converge? How long is the 

effective distance a termite has to cover from one particular point 

to another? 

The answers to these questions all depend on the particular 

topology of the network of galleries. Starting from a topological 

analysis of the gallery system of termites Cubitermes [Perna et al. 

Naturwissenschaften DOI : 10.1007/s00114-008-0388-6] we will 

proceed to discuss the potentialities of a “topological” approach to 

better understand the internal organization of social insects nests. 

In particular, by characterizing gallery networks with tools 

developped in the field of graph theory, we can analyze the 

functions of transport and exchange of information or material 

associated to different nests. We can also obtain compact yet 

informative descriptors of the complex arrangement of internal nest 

structures. The level of description is su±ciently abstract to allow 

direct comparisons between nests built by different species and 

evaluate directly architectural similarities and differences. 

- 55 - 

Evolutionary relationships within the Pachycondyla apicalis 

species complex 

Chantal Poteaux-Léonard, Ronara S. Ferreira and Dominique 

Fresneau 

Laboratoire d‘Ethologie Expérimentale et Comparée (LEEC), 

CNRS UMR 7153 - Université Paris 13, 99, avenue J-B. Clément, 

93430 Villetaneuse, France 

Ponerines are considered as one of the oldest 

phylogenetic ant assemblages. They are 

characterised by high species diversity, a vast 

geographic distribution area, and considerable 

variations in their anatomical features. However, 

they remain primitive in their general social 

organisation. Because of misinterpretations of 

the biological traits examined and biased 

phylogenetic reconstructions based on 

morphological data, the phylogeny of 

“ primitive ” ants has been subject to reassessment 

during the past decade. In the 

Pachycondyla apicalis complex, ongoing speciation 

has resulted in several closely-related, 

sympatric taxa. Even though a recent taxonomic 

revision has been published, species limits are 

difficult to assess in P. apicalis due to their 

conservative morphology. There is still 

disagreement about the actual number of species 

and their relationships. To clarify these 

questions, we performed separate and combined 

molecular phylogenetic analyses based on nuclear 

and mitochondrial DNA sequences. Our aims are to 

reconstruct a robust phylogenetic hypothesis 

using a dense taxon sampling, and to investigate 

the evolution of behavioural traits (e.g. social 

structure, nest location, mating strategy, among 

others) among the examined taxa. 

Rescue behavior in Cataglyphis cursor ants: an altruistic 

behavior or a simple reaction toward individuals in distress? 

Alexandra Scohier and Elise Nowbahari 

Laboratoire d’Ethologie Expérimentale et Comparée UMR CNRS 

7153, Université Paris Nord, 93430 Villetaneuse, France. 

Cooperation is an interaction between, at least, two individuals to 

accomplish a task. Cooperation frequently involves altruistic 

behavior. A particular pattern, the so-called nestmate rescue 

behavior, is expressed by Cataglyphis ants when they are 

collected in the field. In this research, we created an experimental 

procedure, not only to study this behavior in the laboratory, but 

also to determine whether it should be considered as real rescue 

behavior toward nestmates or if it is only a reaction to an individual 

in distress. Our results clearly show that C. cursor ants help only 

their nestmates by employing specific behavioral strategies. When 

nestmates are in distress, ants display a particular helping 

behavior that is excavation, dragging the ant and moving sand. 

However, when a stranger ant or a prey is encountered buried in 

sand, ants exhibited agonistic behavior (i.e threatening, biting and 

gaster flexion). Thus, C. cursor ants discriminate between different 

distress situations and adapt their behavior accordingly. Future 

exploration of this behavior, coupled with biochemical analysis (the 

study of the alarm signal and of the cuticular hydrocarbons by gas 

chromatography) is planned to determine the nature of the signal, 

itself, as well as to determine whether all colony members can act 

as helpers and whether all ants in the colony are helped equally.


From individual to social behaviour: ant colonies as complex 

systems 

Ana Sendova-Franks 

School of Mathematical Sciences, CEMS, University of the West of 

England, Bristol, Frenchay Campus, Coldharbour Lane, Bristol 

BS16 1QY, UK 

Ant colonies have appealed to our imaginations and challenged 

our comprehension for hundreds of years. Today they feature in 

the World Wide Web encyclopedia, Wikipedia, as one of the 

classic examples of a complex system. However, we are only 

beginning to understand how ant colonies are organised. It is still 

not widely appreciated that social insects in general and ant 

colonies in particular represent a prime model experimental 

system for studying complexity for three main reasons. First, there 

is an evolutionary link between individual and society where the 

individual has a relatively high degree of independence. Second, 

both levels of organization are relatively tractable experimentally. 

Third, we can explore whether individual complexity plays a crucial 

role. This is a key topical example of how zoology transcends 

solely the study of animals and links with general theory and 

application, in this case our understanding of how the multiple 

interactions between entities at one organizational level (microlevel) 

influence the behaviour at a higher or macro-level. My 

presentation will be based on phenomena such as the division of 

labour, worker Spatial Fidelity Zones inside the nest, brood sorting, 

building, the improvement of collective performance with 

experience and the distribution of food. All have been studied 

through experimental manipulation and the meticulous study of the 

behaviour of individuals within their societies in colonies of the ant 

Temnothorax albipennis - a species that is particularly amenable to 

studies attempting to link phenomena at the colony (macro-) level 

to behaviour at the individual (micro-) level. 

- 56 - 

Altruist in insect societies and beyond: voluntary or enforced? 

Tom Wenseleers 

Laboratory of Entomology, Zoological Institute, Katholieke 

Universiteit Leuven, 

The extreme altruism of the social insect worker caste has puzzled 

researchers for decades. Inclusive fitness theory suggests that 

close relatedness has been key in promoting such altruism. 

Recent theory, however, shows that the intermediate levels of 

relatedness found within insect societies are too low to directly 

cause the extreme altruism observed in many species. Instead, 

recent results show that workers are frequently “coerced” into 

acting altruistically. For example, workers are deterred from laying 

eggs by egg killing, and female larvae are prevented from 

developing into queens by food control. This shows that the 

altruism seen in many modern-day insect societies may not be 

voluntary but enforced. In this talk I will argue that enforced 

cooperation in fact is a widespread phenomenon, that occurs not 

just in insect societies, but also in social vertebrates, humans and 

interspecific mutualisms. 

F.L.W. Ratnieks & T. Wenseleers (2008) Altruism in insect 

societies and beyond: voluntary or enforced? Trends in Ecology 

and Evolution 23: 45-52. 

T. Wenseleers & F.L.W. Ratnieks (2006) Enforced altruism in 

insect societies. Nature 444: 50.


Does early sibling aggression in Lynx fit facultative siblicide 

theories? 

Anastasia Antonevich 1 and Sergey Naidenko 2 

A.N. Severtsov Institute of Ecology and Evolution, Leninsky pr.33, 

Moscow, 119071, Russia 

One spontaneous and sometimes lethal sibling fight occurs in 

more than 50% of European lynx twins and triplets during 6-7 

week of cubs life (Sokolov et al., 1994). The same siblicidal fights 

were described in zoos and found in Iberian lynx litters (Vargas et 

al., 2005). Facultative siblicide can evolve when the fitness benefit 

gained by a dominant offspring, exceeds the cost of reduced 

inclusive fitness. The main environmental component expected to 

control sibling aggression is food, fights can lead to individual 

advantages in feeding and growth as well as deaths of broodmates 

(Drummond, 2001). According to the Challenge Hypothesis the 

androgens level is up-regulated during social challenges (Wingfield 

et al., 1990) including sibling aggression period (Ferree et al., 

2004). We tested these hypotheses for facultative in Eurasian lynx. 

Fights were recorded both in twins and triplets. Contrary to 

expectation aggression in lynx litters was not directly dependent on 

food supply and intralitter competition for food. None the less, 

fights occurred mostly in the litters with lower growth rate and led 

to weight gain advantages for winners. Winners also got priority in 

food access. Although predictions of the resource-tracking 

hypothesis were not supported, growth rate differences reveal the 

resource competition role in sibling aggression. No enhancement 

of androgens or cortisole levels were found during the fighting 

period but plasma androstendione was elevated 3 weeks before 

fighting period in aggressive comparing to non-aggressive cubs. 

Spontaneous fights in lynx appear to be unique phenomenon that 

nevertheless fits main siblicide patterns. 

Diversity of the baltic clam Macoma balthica L. at local and 

geographical scales (Barents Sea) 

Larisa Basova, Mikhail Gantsevich, Eugene Genelt-Yanovsky and 

Petr Strelkov 

Leninskie Gory 1, build.12, 119991, Moscow, Russia 

The bivalve mollusc Macoma balthica is common and often 

dominant member of littoral soft-bottom communities along the 

North European coasts. Macoma demonstrates considerably high 

inter- and intra-population variability in size, growth and 

morphological characters such as shell shape, colour and hinge 

structure. Spatial variation of some of these parameters was early 

studied at local and (or) geographical scales! . However there is no 

common opinion about the level of variation and factors causing 

the variability. The goal of our study was to analyze spatial 

variation of M. balthica growth and morphology at the scale of Kola 

Bay, the biggest fjord in the Russian sector of the West Barents 

Sea. Results of our study show that the southern segment of the 

Kola Bay and open shore localities represents two the most 

different marine environments. Top of the Bay is a typical estuary 

while oceanic conditions predominate in open shore localities. 

Central and northern segments of the Bay can be described as 

transitory area. Populations of M. balthica are in fact the most 

different between the top of the Bay and open coast in respect of 

shell shape and color, hinge plate abnormalities, and longevity. At 

the same time growth rate is similarly high in two regions, and is 

depressed in the transitory area. Latitudinal clines in growth rate, 

longevity, size and shell shape of M. balthica were early revealed 

in Europe. The result of our investigation shows that variation of all 

listed parameters in the Kola Bay is comparable to that in the 

whole Europe. 

S17 - Phenotypical plasticity and behavior 

- 57 - 

A theoretical study of alternative female mating strategies 

Josefa Bleu, Carmen Bessa-Gomes and David Laloi 

Laboratoire Écologie & Évolution (UMR 7625), Université Pierre et 

Marie Curie, 7 quai St Bernard, 75 252 Paris Cedex 05, France 

Alternative mating strategies are increasingly recognized as being 

common in many species. Although alternative female mating 

strategies have been less studied than male ones, they may also 

be frequent. For example, both monandrous and polyandrous 

females of Lacerta vivipara are observed simultaneously. The aim 

of this study is to unravel the behavioural mechanisms that can 

lead to the coexistence of singly and multiply-mated females in the 

same population. We focused on the role of female mate choice 

(either by trade-up or by selection given a threshold of mating 

acceptance). We explored the role of external factors (density, 

sex-ratio and cost of multiple mating). This analysis was conducted 

with an evolutionary individual based model. Trade-up alone 

allows the coexistence of monandrous and polyandrous females at 

the beginning of a reproductive season, but if the season is long 

enough, all females become polyandrous. When adding a 

threshold to first mate acceptance, the evolutionary simulations 

show that populations converge towards coexisting strategies with 

a high percentage of monandrous females, while the acceptance 

threshold evolves towards similar values among females. The 

selected threshold is very sensitive to the demographic factors 

whereas the monandrous / polyandrous females' ratio is relatively 

stable. An increase of the cost of multiple mating selects for higher 

threshold and consequently for more monandrous females despite 

the risk of remaining unmated at the end of the season. 

Sex-specific flexibility of parental care allows for the sexual 

conflict to be resolved in King penguins, Aptenodytes 

patagonicus 

Hélène Corbel, Sylvie Geiger and René Groscolas 

Institut Pluridisciplinaire Hubert Curien, UMR 7178 CNRS, 

Département Ecologie, Physiologie et Ethologie, 23, rue Becquerel, 

67087 Strasbourg Cedex 2, France 

In biparental species, sex differences in reproductive costs and 

mate availability entail a conflict over parental investment between 

sexual partners. In King penguins, breeding cycles last more than 

one year and are just separated by moult, resulting in high 

availability of mates for a extended period. On the other hand, 

breeding success depends on the hatching date so that 

reproduction is highly time-constrained. Because of size 

dimorphism and male-biased sex-ratio, sexual conflict would be 

resolved through female-initiated divorce resulting in 50% of brood 

being deserted by mother during rearing. Aiming at determining 

the mechanisms underlying the resolution of sexual conflict, we 

investigated sex-specific correlates of the transition from 

reproduction to moult. We focused on prolactin (Prl) and T4 levels, 

involved in care and the initiation of moult, respectively, together 

with offspring signals. In females, care was inflexible and required 

elevated Prl to be maintained. The termination of care was 

characterized by a concerted decrease of Prl and body condition 

and by antagonism between Prl and T4, possibly to prevent from 

the initiation of energy-demanding moult while feeding the chick. In 

males, the intensity of care was linked to Prl and chick solicitation. 

Accordingly, males partially compensated for the desertion of 

females. Both body condition and T4 tended to increase at the end 

of rearing, indicating that males were able to initiate moult and 

associated energy storage while feeding the chick. Therefore, 

flexibility of paternal care together with overlapping of male 

reproduction and moult allow for energy-restrained females to 

desert.


Nest biology of Monoeca haemorrhoidalis (Smith, 1854) 

(Apidae) at the Atlantic Rainforest of Southern Brazil 

Léo Correia da Rocha-Filho 1 and Gabriel A. R. Melo 2 

1 Departamento de Biologia, Universidade de São Paulo, Av. 

Bandeirantes 3900, Monte Alegre, 14040-901, Ribeirão Preto, SP, 

Brazil ; 2 Departamento de Zoologia, Setor de Ciências Biológicas, 

Centro Politécnico, Universidade Federal do Paraná, 81531-990, 

Curitiba, PR, Brazil 

The genus Monoeca is distributed over almost the entire 

Neotropical region, with ten described species. The nesting biology 

of a few species, Monoeca sp., M. schrottkyi, M. lanei and M. 

xanthopyga, has been studied by previous authors. The aim of this 

study was to investigate the nesting ecology of M. haemorrhoidalis. 

This study was conducted between Setember/2005 and 

January/2007, in an area of Atlantic Rainforest. Four nest 

aggregations were studied, as well as other smaller aggregations 

of M. haemorrhoidalis also found along the reserve trail. M. 

haemorrhoidalis females construct their nests in clay soil, in dense 

aggregations, with density values varying from 4 to 27 nests/m 2 . 

The period of nest construction and cell provisioning started at the 

end of October/2005 and stopped by the end of February/2006. 

During this period, plant species of the families Orchidaceae, 

Styracaceae and, mainly, Malpighiaceae, were the most important 

pollen and floral oils resources that were utilized in brood cell 

provisioning. Males and females gathered nectar in a great variety 

of plant species. M. haemorrhoidalis is a univoltine and seasonal 

species and its nest habits are similar to those already observed 

for other Monoeca species. 

Natural enemies and other species associated with Monoeca 

haemorrhoidalis (Smith, 1854) (Apidae) 

Léo Correia da Rocha-Filho 1 and Gabriel A. R. Melo 2 

1 Departamento de Biologia, Universidade de São Paulo, Av. 

Bandeirantes 3900, Monte Alegre, 14040-901, Ribeirão Preto, SP, 

Brazil 

2 Departamento de Zoologia, Setor de Ciências Biológicas, Centro 

Politécnico, Universidade Federal do Paraná, 81531-990, Curitiba, 

PR, Brazil 

The bee species Monoeca haemorrhoidalis, the largest one in the 

genus, occurs in the Atlantic rainforest of southeastern and 

southern Brazil. This study focused in the interactions between M. 

haemorrhoidalis and its natural enemies and associates. Nest 

aggregations were studied in an area at the transition between 

Dense and Mixed subtropical Rainforest, in southern Brazil. During 

the nest activities, between October/2005 and February/2006, 

thirty-two animal species were observed at the nesting sites. 

Nonetheless, association with M. haemorrhoidalis was confirmed 

only for the cleptoparasitic species Protosiris gigas (Apidae), 

Heterostylum maculipennis (Bombyliidae) and Megaselia sp. 

(Phoridae), Tetraolytta gerardi and Tetraonyx distincticollis 

(Meloidae), the predators Pyrogaster moestus (Lampyridae) and 

Pachycondyla harpax (Formicidae), the cleptobiont species 

Acromyrmex niger (Formicidae) and the parasitoids Physocephala 

bipunctata (Conopidae), Pseudomethoca melanocephala e 

Hoplocrates specularis (Mutillidae). Moreover, pathogenic fungi 

developed in stored food of brood cells and in M. haemorrhoidalis 

cocoons and Meloidae larvae, causing the death of these species. 

The cleptoparasitic bee P. gigas was one of the main causes of M. 

haemorrhoidalis mortality. During the peak of activity of P. gigas, M. 

haemorrhoidalis had a substantial decrease in its nest construction. 

This can be an important adaptation of the host species against its 

cleptoparasite. 

- 58 - 

Phenotypic plasticity, genotype x environment interaction, 

and the (un)reliability of animal mating signals 

Michael D. Greenfield 

IRBI (Institut de Recherche sur la Biologie de l’Insecte), CNRS 

UMR 6035, Université François Rabelais, Parc de Grandmont, 

Tours 37200, France 

The potential for animals to ‘deceive’ one another via ‘dishonest’ 

signals remains a major question in behavioral ecology. The 

current view, that on average an individual’s signals are reliable 

indications of potential ability, is based on the argument that 

receivers would be selected to ignore signals that are habitually 

unreliable, which, in turn, would select against the production of 

such signals. Formal analyses as well as observations and 

experimental findings are largely consistent with this generalization. 

A very different, and more insidious, problem for reliable 

communication, particularly in mating, is posed by the plasticity of 

signal traits across environments: Because genotypes may 

respond in different ways to environmental changes across space 

or time, a given genotype may exhibit the ‘superior’ (mating) signal 

in one environment but the ‘inferior’ one in another. Thus, (mating) 

signals may not be reliable indications if the environment changes 

across generations or offspring disperse to different environments. 

This conundrum does not necessarily challenge the primacy of 

signal reliability, but it points out fundamental weaknesses in our 

understanding of signal evolution that result from ignoring 

unpredictable environmental variation. Here, I present findings on 

genotype x environment interaction (gei) in the male courtship 

song of the ultrasonic pyralid moth Achroia grisella. I show how 

signal unreliability can arise and discuss the restricted 

circumstances under which reliability might yet persist. I then 

present recent findings on genetic variance, phenotypic plasticity, 

and gei in female response and preference in A. grisella, and I 

show how reliability in animal communication can increase or 

decrease when both male signal and female response traits are 

subject to gei. 

Prenatal stress influences behavioural features in young birds 

Floriane Guibert 1 , Cécilia Houdelier 1 , Sophie Lumineau 1 , Kurt 

Kotrschal 2 , Erich Möstl 3 and Marie-Annick Richard-Yris 1 

1 

UMR CNRS 6552 Ethologie animale et humaine, Université de 

Rennes 1, 35042 Rennes, France 

2 Konrad-Lorenz-Forschungsstelle, University of Vienna, A-4645 

Grünau, Austria 

3 

University of Veterinary Medicine, Department of Natural 

Sciences, Biochemistry, Veterinärplatz 1, A-1210 Vienna, Austria 

The individual’s behavioural development is notably influenced by 

his social environment and particularly by his mother. This 

maternal effect occurs after youngster’s birth but also before. In 

mammals, prenatal maternal stress influences the setting up of 

youngster’s behaviour; this influence results from the modulation of 

the mother’s plasmatic levels of steroid hormones. In birds, a 

similar maternal influence also exists, implicating a modulation of 

steroids levels in the egg. Indeed, egg’s hormonal levels are 

influenced by laying females’ environment and an artificial steroids’ 

enrichment of eggs modulates the behavioural phenotype of 

resulting chicks. However, no study has investigated yet the whole 

mechanism of maternal prenatal influence in birds (i.e. from the 

mother to the offspring). Thus, our aim was to analyse the impact 

of stress on the laying female on her eggs’ hormonal composition 

and on the behavioural features of her offspring. Therefore, we 

applied stressors on laying females in a precocious bird, the 

Japanese quail. Our results showed that the prenatally stressed 

chicks appeared to be more emotive with a higher social 

motivation and also a slight increase of steroids in the eggs of 

stressed females. Thus, in this study, we show for the first time 

that, in birds, stress on the laying female has a significant impact 

on the behaviour of her offspring via a hormonal change in her 

eggs.


Sex differences in cognition 

Susan D Healy and Anjanette Harris 

Institute of Evolutionary Biology, Kings Buildings, EH3 9JT, 

Edinburgh, UK 

Tests of spatial cognition produce the best substantiated of the 

purported differences in cognitive abilities between males and 

females (mammals). Although there are at least seven extant 

hypotheses as to why this sex difference might have evolved, 

there are few data to differentiate amongst them. Additionally, 

although the difference is always in the male’s favour, even spatial 

cognition tests do n! ot always result in a sex difference. There are 

at least two hormonal explanations for this, due to the effects of 

sex steroids and stress hormones on cognitive performance. For 

example, fluctuations in oestrogen across an oestrous or 

menstrual cycle result in lower spatial performance by females, but 

only at certain stages of that cycle. Female performance will, then, 

may be as good as males at some stages of the cycle and poorer 

at others. Additionally, or alternatively (it is not yet clear), stress 

has a differential impact on males and females, such that cognitive 

performance in females is typically poorer under acute stress 

conditions. We argue that stress, as imposed via the testing 

situation (often a Morris water maze task) may explain the 

occurrence of sex differences in spatial condition in the laboratory 

rat, at least. If so, we suggest that the laboratory rat (and, perhaps 

laboratory testing) is not an appropriate model system for 

addressing questions on th! e evolution of sex differences in 

spatial cognition in mammals. 

Ecological factors involved within adjustment of European 

hedgehog (Erinaceus europaeus) to urbanization 

Pauline Hubert 1,2 , Romain Julliard 3 , Sylvie Biagianti 1 and Marie- 

2, 4 

Lazarine Poulle 

1 Laboratoire d’éco-toxicologie, EA 2099, URVVC, Université de 

Reims Champagne-Ardenne, 51687 Cedex 2 Reims, France 

2 Centre de Recherche et de Formation en Eco-éthologie (2C2A- 

CERFE), 5 rue de la Héronnière, 08240 Boult-aux-Bois, France 

3 Centre de Recherche sur la Biologie des Populations d’Oiseaux 

(CRBPO), Muséum national d’Histoire naturelle, 55 rue Buffon, 

75005 Paris, France 

4 Laboratoire de Parasitologie - Mycologie, EA3800, IFR 53, 

Université de Reims Champagne Ardenne, 51100 Reims, France. 

European hedgehog is a wild mammal considered as an "urban 

adapter” since it is able to maintain and even increase its 

population despite urbanization. Adjustment of its populations to 

specific conditions of the urban environment, especially through an 

increase of their density, is an important indicator of the ecological 

plasticity of the species. The objective of the present study was to 

identify the ecological factors responsible for the density of 

hedgehog population being higher in cities than in the countryside. 

The study was conducted on a 4213 ha area located in the 

Ardennes region, North-eastern France. It includes the Sedan city 

(21000 inhabitants) and the neighbouring rural area. Forty three 

transects of 500m each were spread out on the study area in order 

to perform visual catch of hedgehog at night, with infrared 

binoculars. Transects were walked twelve times from June 2006 to 

October 2007, leading to 127 encounters. General Linear Models 

(GLM) were built to identify the ecological factors explaining the 

number of individuals observed per transect. Tested variables 

were i) earthworm and arthropod biomass per transect (main food 

resource) estimated with sampling methods applied to the 5 types 

of habitat present in the study area, ii) presence of pet food and 

anti-slug pellet or pesticides obtained by means of a verbal 

questionnaire and iii) proximity of a badger, Meles meles sett 

(hedgehog’s predator). Hedgehog density, estimated from distance 

of hedgehog localisations to transects, and importance of factors 

explaining hedgehog presence were then compared between 

urban and rural areas. 

- 59 - 

Some aspects of the Great Warbler' behavior in wet land 

(Romania) 

Constantin Ion 

Bd. Carol I, no. 20A, Faculty of Biology, 700505, IASI, Romania 

The Great Reed Warbler is one of the most common warbler form 

the wet areas of Romania. It can be found especially around the 

reeds beds plain and it is visiting our country form spring until 

autumn. The spring migration takes place in the second part of 

April. At the end of the April –the beginning of May, the great reed 

warbler males start to delimitate the nesting territories. 

These territories cover generally a surface of 100-250 m ². The 

territoriality behavior can be seen as a form of aggression and also 

competition, when individuals of the same species compete for a 

mate and for environmental resources (food, shelter, from 

predators, wind and heavy rain). 

While delimit the territory these birds manifest the supremacy of 

one warbler to another through emitting complex songs and 

showing intimidating positions. 

The establishment of the warbler territory is closely connected with 

the meteorological conditions of the environment in which they live. 

Rain or low temperatures are the main factors detrimental to the 

establishment of territories. These territories can be contiguous 

from one warbler to another or buffer spaces may exists between 

them. We observed that territories have an opening to grassland 

were they catch insects, and another opening towards the open 

water were they take shelter, favor the success of the reproduction. 

The building of the nest last 5-6 days, the deposition of the eggs 

( 4-5 ) takes place 3-6 days (one egg - one day). The female is 

hatching 10-14 days. After 30 days the chicks became good flying 

and able to have an independent life. 

The great reed warbler´s behavior is correlated with the breeding 

stage. If the warblers have eggs, they leave the nest when a male 

comes around. If the warblers have broods in their nest, they will 

become very agitate, aggressive, and will try to sting the male in 

order to defend the chicks. 

Individual fate within an ant clone: the nature vs. nurture 

debate under the microscope 

Emmanuel Lecoutey, Fabien Ravary, Nicolas Châline and Pierre 

Jaisson 

Université Paris 13, Laboratoire d’Ethologie Expérimentale et 

Comparée UMR CNRS 7153, 99 av. JB Clément, 93430- 

Villetaneuse, France 

The life cycle of some insular populations of the Asian tropical ant 

species Cerapachys biroi Forel is likely to be unique worldwide. In 

effect, these populations lost males and queens and maintain 

themselves through the workers’ parthenogenesis. Moreover, all 

the individual workers oviposit, participate to reproduction, making 

copies of themselves, at least during the first weeks of their adult 

life (Ravary & Jaisson, 2004). Eggs are laid altogether in a short 

round, which results in a cyclic production of generations of 

workers (Ravary & Jaisson, 2002). All workers of a same 

generation are homogenous concerning age as they hatch 

together within in a few hours. The C. biroi callow workers usually 

stay within the nest displaying nursing behaviour and then become 

foragers while aging. This species feed exclusively on ant brood of 

alien ant species which makes easy to control strictly their diet at 

the laboratory. Finally, hierarchy as well as worker polymorphism 

(potential causes of division of labour) are absent within colonies. 

These life history characteristics together allowed us to set up an 

experimental design in which young workers of the same clone 

and of the same age were reared in the same conditions, except 

for the foraging experience: 50% were let to catch preys whereas 

50% were not permitted to get a foraging success. Then, the fate 

of these individuals was dramatically oriented accordingly: those 

who experienced foraging successes became foragers and those 

who experienced foraging failures turned towards nursing tasks 

(Ravary et al. 2007). This direct demonstration of the role of 

experience in the determinism of social behaviour reminds recent 

knowledge about cell differentiation within organisms (Chang et al. 

2007) and may contribute to revive the superorganismic 

conception of insect societies (Wilson & Sober 1989).


Reproductive behavioral plasticity in Père David’s deer 

Zhigang Jiang 1,2 , Yan Zeng 1,2 , Zhenyu Zhong 3 , Chunwang Li 1 and 

Linyuan Zhang 3 

1 

Key Laboratory of Animal Ecology and Conservation Biology, 

Institute of Zoology, Chinese Academy of Sciences, Beijing, 

100101, China. 

2 

Graduated School of Chinese Academy of Sciences, Beijing, 

China 100049, China. 

3 

Beijing Père David’s Deer Park 

Père David’s deer is a polygamous species; one to several stags 

rut at the same time, a “Harem master” defends the receptive 

female herd and tries to monopolize the opportunity of mating. 

However, reproductive behavior in Père David’s deer is highly 

plastic. First, the males have different rut tactics: “Harem master 

tactic”, “Challenger tactic” and “Bachelor tactic”, rut season is 

extended for a long period, e.g. rut lasts as long as two months in 

the Beijing Père David deer population. Thus, the time to 

commence rut is plastic in stags. Female “estrus window”, which is 

composed of the individual females in estrus, drifts among years. 

That means unknown factor controls the estrus in the females, or 

females are able to adjust their estrus. We conducted a study on 

the reproductive behavior of Père David’s deer from 1994 to 2005. 

We observed the behaviors of males and females and used five 

polymorphic microstatellite loci which were screened out from 84 

pairs of species-transferred primers to study the mating system in 

the reintroduced Père David’s deer populations in China. The 

study identified multi stags sired offspring in the population. 

“Harem masters” could not monopolize breeding opportunities, 

Challengers also had chance to breed. While the plasticity in 

female estrus window presumably is related to female cryptic sex 

choice, on the other hand, to be a “Harem master” is not the “The 

winner takes all” tactics. The multi-paternity in Père David’s deer 

indicates the rut tactics like “Challenger tactic” has its evolutionary 

fitness as well. 

Relationships between morphological traits, behavioural 

characteristics, and habitat selection in Iranian Wheatears 

using Fourth-Corner Problem analysis 

Mohammad Kaboli 1 , Mansour Aliabadian 2 , Saina Habibi 1 , Parisa 

Mehrandish 1 and Roger Prodon 3 

1 

Department of Fishery and Environment, Faculty of Natural 

Resources, University of Tehran, Tehran, Iran 

2 

Department of Biology, Faculty of Science, Ferdowsi University of 

Mashhad, Mashhad, Iran 

3 

Laboratoire Ecologie et Biogéographie des Vertébrés (EPHE), 

Centre d’Ecologie Fonctionnelle et Evolutive, UMR 5175, 1919 

route de Mende, 34293 Montpellier cedex 5, France 

Ecological segregation in the Wheatear genus Oenanthe is a 

complex and intriguing question. Here we studied habitat variables, 

behavioural characteristics and morphological traits of 11 

Wheatear species that are distributed sympartrically in Iran. We 

recorded 19 behavioural characteristics and measured 36 habitat 

variables within a 100-m radius around observed birds. We also 

measured 21 biometrical variables in each species. We compared 

variable-to-variable correlation between these three data matrices 

using Legendre et al.’s Fourth-Corner Problem method. We first 

related, through the matrix A1 (presence/absence of species in 

sampling sites), the morphological traits of the species (mean 

values of the variables for each species; matrix Cmorphology) to the 

habitat features measured on each sampling site (matrix Bhabitat). 

We then related, through A1, the behavioural traits of the species 

(mean values of the variables for each species; matrix Cbehaviour) to 

the habitat features measured on each sampling site (matrix 

Bhabitat). We lastly related, through A2, the morphological traits of 

the species (matrix Cmorphology) to the behavioural variables 

measured, on each sampling site, on the "principal" species of 

Wheatears of the site (matrix Bbehaviour). Our results showed that 

morphological traits, particularly flight and foot-leg apparatuses, 

represent a good synthesis of overall behaviour traits. Since 

Morphological traits result from compromise between different 

selection pressures like foraging mode and migration pattern, 

seems plays an important role in determining the ecological range 

of a species. Likewise, habitat features are also correlated with 

behavioural traits, but less tightly. 

- 60 - 

This correlation between behaviour and habitat could be explained 

by this fact that many behavioural variables are defined on the 

habitat-basis (e.g. types of perch used). But the proportion of 

significant relationships between habitat and behaviour remains 

the same when we consider only the "purely" behavioural variables 

(type, speed, and frequency of movements). Finally, The match 

between morphology and habitat is relatively loose that may partly 

be due to the fact that habitat variables are more likely depend on 

proximal factors, especially the sampling conditions (e.g., unequal 

frequency of different landscape features in different sampling 

areas, random component of the detection of the species). 

Control of copula duration: how males and females control 

sperm transfer 

Joshka Kaufmann, Jean-François Le Galliard and David Laloi 

Laboratoire Ecologie & Evolution - UMR 7625, Université Pierre et 

Marie Curie / ENS / AgroParisTech / CNRS, Paris, France 

In polygynandrous mating systems, sexual conflict can arise on the 

number of partners and on the reproductive investment of each 

sex. Particularly, traits that control sperm transfer can have quite 

different optima with regard to female and male fitness. Indeed, 

female behaviour may have evolved to ensure fertilization, favour 

preferred male through longer copulation or promote postcopulatory 

mate choice. Males may strategically allocate their 

sperm according to female reproductive quality, level of sperm 

competition and further mating opportunities. Behavioural 

mechanisms have been poorly studied in taxa, such as reptiles, 

where sperm transfer is driven by copula duration rather than by 

the number of copulations. In the present study, we investigate in 

the common lizard whether males and females control copula 

duration. In a sequential mating context, we study how mate 

choice and copulatory behaviours vary with individual quality (e.g. 

coloration, social dominance, fitness-related physical 

performances) and mating history. We principally disentangle the 

respective role of both sexes in this conflict for sperm transfer. 

Both sexes become more choosy in second mating opportunities, 

female preference being based on male ventral coloration while 

males prefer virgin females. Our results show that females control 

copula duration and hence sperm transfer and may therefore bias 

paternity. Nevertheless, no evidence was found that females 

favour high-quality males through longer copulation. Here, we 

suggest that female lizards may use both pre-copulatory mate 

choice and control of copula duration to maximize the genetic 

benefits of multiple mating. 

Is discrimination of song quality by shortened-wings female 

Canaries (Serinus canaria) reduced ? 

Alexandre Lerch and Laurent Nagle 

Laboratory of Ethology and Comparative Cognition, Université de 

Paris X-Nanterre ; Bat BSL, 1er étage; 200, avenue de la 

République, 92 000, Nanterre cedex 01, France 

Because of their greater investment in reproduction, females are 

supposed to be choosier than males when mating (Trivers, 1972). 

While numerous studies investigated this aspect of mate choice, 

most of them have focused on male traits that maintain female 

preferences (Jennion & Petrie, 1997). 

Nevertheless, consensus between females in the selection of male 

phenotypic features is not the rule, and more subtle mechanisms 

take place. Indeed, variations in female preferences do exist, 

especially in some contexts of predation (Simcox et al., 2005) or 

costly competition (Fawcett and Johnstone, 2003). In birds, a case 

of reduced discrimination for male coloration has been reported in 

female zebra Finches with shortened wings (Burley & Foster, 

2006). 

In canaries (Serinus canaria), song is known to be the main 

secondary sexual feature, and a special phrase (« A » phrase) is 

reported to elicit a high level of sexual responses by females 

(Vallet et al., 1998). In our study, we investigate the power of 

discrimination toward songs of different quality, in a group of 

female canaries tested in two conditions: 1) with intact wings 

(control group) and 2) with cut wings. Our preliminary results 

suggest that birds with shortened wings show a weaker preference 

toward high quality songs.


Plasticity of the gastrointestinal tract of the burmese python 

Jean-Hervé Lignot, Cécile Helmstetter, Robert Pope, Alain 

Ackermann and Stephen Secor 




Morphological changes observed in the gastric and intestinal lining 

of fed and fasting Burmese pythons were studied using 

immunohistochemistry, Western blotting, scanning and 

transmission electron microscopy techniques. In the stomach, 

oxyntopeptic cells of fasting animals are filled with zymogen 

granules and possess a thick tubulovesicular system that is 

transformed into apical digitations projecting into the lumen of the 

gastric crypts soon after feeding. This allows the proton pump 

sequestered into the tubulovesiular system of fasting animals to 

actively excrete proton ions immediately after ingestion of the prey. 

In the intestine, fasting animals possess hypotrophied enterocytes 

with small apical microvilli and numerous lysosomes and related 

organelles such as numerous apical multivesicular bodies, 

embodied particles and large lamellar bodies filled with concentric 

rings of lipid membranes. 

Fed individuals, however, show enlarged absorbing cells that can 

be filled with lipids as well as elongated microvilli. A new cell type 

within the mucosal epithelium is also described that has an apical 

crypt. This cell type is only present in the proximal part of the 

intestine, is connected to the b! asal membrane, is devoid of large 

lipid droplets, possesses a large nucleus, and is less stained than 

its neighbouring absorbing enterocytes. In fed animals the crypt is 

usually filled with a multi-layered spheroid particle made of calcium 

and phosphorus. Gut plasticity is therefore of crucial importance in 

Burmese pythons and relies on rapid and massive morphofunctional 

changes as well as cell components recycling and 

trafficking. 

Thermogenesis of digestion in juvenile burmese pythons 

Jean-Hervé Lignot and Thibault Poolny 




Burmese pythons experience dramatic increases in metabolism 

during meal digestion that can directly modify thermogenesis. 

Temperature data loggers were surgically implanted adjacent to 

the stomach and small intestine or inserted inside the prey. 

Infrared images of fasting pythons and up to 120 hours after 

feeding were also collected from th! e snakes’ skin. The effects of 

environmental temperature on postprandial thermogenesis were 

evaluated by recording skin and body temperatures during the 

digestion of a rodent meal equalling 20% of snake body mass at 

room temperatures of 20, 25, 30, and 35°C. We also identified the 

effects of meal size on thermogenesis by recording temperatures 

following the ingestion of meals equalling 10, 20, or 30% of snake 

body mass. At 20°C, skin and body temperatures of digesting 

pythons were barely elevated above room temperature. Between 

25 and 35°C, skin and body temperatures rapidly increase after 

feeding and peaked 20-24 hours after feeding at 1–2.5°C above 

room temperature. Thereafter, body temperatures remained 

elevated before returning to room temperature 50-80 hours after 

feeding. With an increase in meal size, pythons experience a 

larger increase in skin and body temperatures and a longer 

duration of elevated temperatures. For 10% meal, body 

temperature rose by 1.2°C and remained elevated for 60 hours, 

whereas for the 30% meals, body temperature increased by 2.5 °C 

and remained elevated for 80 hours. A by-product of the work of 

digestion, this endogenous production of heat is beneficial to the 

python in increasing the rate of meal digestion and assimilation. 

- 61 - 

Learning of a heterocolonial, heterospecific template in adult 

workers of the wood ant, Formica rufa (Hymenoptera: 

Formicidae): a neuropharmacological approach 

Sylvie Marques and Stéphane Chameron 

Laboratoire d’Ethologie Expérimentale et Comparée, UMR CNRS 

7153, Université Paris 13, 99 Avenue JB Clément, 93430 

Villetaneuse, France 

Social insects are characterized by colonial closure that relies on 

their ability to discriminate between nestmates and non-nestmates. 

In ants, recognition mechanisms involve the matching of perceived 

chemical cues to an internal template of colony odour, which has 

been acquired during early adult life in many species. Intercolonial 

relationships in ants can be ruled by the “dear-enemy effect”, 

which consists in a differential treatment of familiar neighbours and 

strangers. Such discrimination can arise either from differential 

olfactory distance between close and distant strangers, or from 

true learning process of neighbours’ colonial visa. 

We present an experimental paradigm where heterospecific 

colonies have been settled as neighbours in the lab. We monitored 

agonistic behaviours in dyadic encounters before and after the 

familiarisation period. Our results show that workers adjusted their 

social behaviour to the stranger’s familiarity level, leading to the 

conclusion that adult workers of the Formica rufa-group can form a 

heterospecific, colony-specific template, in addition to the 

homocolonial visa early learned. 

From a cognitive point of view, heterocolonial visa acquisition is 

probably based on an aversive conditioning process. 

We investigated the role of dopamine, which is known to mediate 

negative reinforcement in invertebrates, by providing experimental 

ants with either dopamine or a dopaminergic antagonist 

(fluphenazine). We report here first evidence that social odour 

learning could indeed be correlated to dopaminergic activity. 

We conclude that our paradigm of social odour learning in ants 

provide a promising model for the investigation of the cognitive and 

neuropharmacological bases of recognition mechanisms in social 

insects. 

Is the drop in the breeding success of white storks related to 

a poorer reproductive ability of released captive birds ? 

Sylvie Massemin-Challet 1 , Delphine Michard-Picamelot 2 , Joël M. 

Durant 3 and Yvon Le Maho 1 

1 

CNRS, CEPE, UPR 9010, associated with Louis Pasteur 

University, 23 rue Becquerel, F-67087 Strasbourg, France 

2 

Mission of Scientific and Technical Culture, ULP, Strasbourg, 

France 

3 

Centre for Ecological and Evolutionary Synthesis (CEES), 

Department of Biology, University of Oslo, P.O. Box 1050 Blindern, 

NO-0316 Oslo, Norway 

A decrease in populations of white storks has been observed in 

Western Europe until 1950. Conservation steps were conducted by 

releasing young storks born in captivity. In the Alsace region 

(France), the reproductive success (number of fledglings) of 

breeding pairs was, however, higher before than after 

reintroduction with a population mainly constituted of birds 

released from captivity. The aim of this three year study was to 

determine if the breeding success in the field depends to the 

realising of captive birds. Reproductive success was lower in 

captive pairs than in free-living pairs both in the Alsace region and 

other parts of France, where no reintroduction has been managed. 

In captivity, food competition within brood had a negative effect on 

the survival of nestlings and on the maximum gain of the two lasthatched 

nestlings in broods. If no influence of hatching order was 

observed on asymptotic body mass, this growth flexibility can have 

an impact on the breeding efficacy of birds, as suggested by other 

studies. In conclusion, one of the factors contributing to the drop of 

reproductive success in areas with reintroduction management 

may be the quality of future breeders released in the field. To 

determine the consequences of overall reproductive success of 

captive birds released in the field on the reproductive success of 

the white stork population, we will compare the reproductive 

success between released and wild birds.


Mammalian (Human) oral tactile imprinting 

Elsie Mobbs 

Department of Perinatal & Infant Mental Health Service, Liverpool 

Hospital and The University of Sidney, The Children’s Hospital at 

Westmead Clinical School, PO Box 36 Westmead, 2145 NSW, 

Australia 

In the situation of maternal deprivation in the newborn critical or 

sensitive period, across the mammalian spectrum body part 

sucking as displacement of the oral imprint on the maternal nipple 

or teat can be observed. Without human intervention in this 

pathological state the newborn dies. 

Calves are recorded as orally taking to the genitalia of other 

prematurely weaned calves and ingesting urine during sucking 

which upsets the digestive system. Kangaroo joeys generally take 

to digits, tail or the cloth of their human replacement pouch or 

swaddling. Monkeys, being more flexible than human primates, 

also take to their tails or penis as well as digits. The imprinted 

sucking object, a reaction to a decoy stimulus feature chosen by 

the infant to replace the maternal nipple/teat, often becomes 

excoriated and in the case of the penis especially, gangrenous. 

Sucking is used by infant mammals to form an emotional 

relationship with the mother rather than it being a need to suck. In 

human infants this relationship is directed towards a stimulus 

feature, either the mother's nipple, an inanimate object such as a 

provided dummy/pacifier, or on to the self as in body-part/thumbsucking 

which often becomes intractable. 

1. Mammalian oral imprinting is on a stimulus feature of the 

mother. 

2. Attachment is when the infant can visually differentiate, 

recognize and follow the mother (lambs approximately 12 

hours and zebras about a week). 

3. Bonding is what adults do. 

4. All three states are linked by the seeking of the object of 

affection and emotional distress by its absence. 

Can pill bugs (Armadillidium vulgare, Isopoda, Crustacea) 

estimate cliff depth with reference to length of antennae? 

Tohru Moriyama 

3-15-1 Tokida, 386-8567, Ueda, Japan 

The length of the antennae of pill bugs in the test group was 

extended by attaching Teflon tubes. Each individual was placed at 

the top of a set of stairs consisting of five steps. The distance 

between the steps was 5, 10, 15 and 17 mm, in order from the first 

to the fifth step. The distance between the third and fourth steps 

(15mm) was the ! critical distance that the extended antennae 

could just reach. Two other groups were also tested: A free-walk 

group, in which the antennae were also extended, but they were 

allowed to move in an arena before placing them on the stairs, and 

a control group, in which the antennae were not extended. Most of 

individuals in the test group descended to the fourth step and the 

median maximum reachable step was 4.0. However, most 

individuals in both the free-walk and control groups failed to 

descend to the fourth step, and the median maximum reachable 

steps were 2.0 and 2.5, respectively. Statistical tests showed that 

the value for the test group was significantly larger than both the 

others. These results suggest that individuals in the test group 

reached the fourth step because they depended on mechanical 

stimulation from the extended tubes, while those in the free-walk 

group didn’t reach the fourth step because they were aware of the 

length of the extensions to their antennae du! ring free-walking in 

the arena and could estimate that the distance was too great to go 

down without falling. 

- 62 - 

Selective forces driving latitudinal clines in Iberian Calopteryx 

damselflies 

David Outomuro and Francisco J. Ocharan 

Departamento de Biología de Organismos y Sistemas, University 

of Oviedo, Oviedo, E-33071, Spain 

Latitudinal morphological clines in ectotherms are generally 

caused by natural selection. However, in the species in which 

sexual selection plays a major role in specific divergence, these 

clines might be deeply modified, especially regarding to secondary 

sexual traits (SST). Calopteryx damselflies are an excellent model 

to study these processes, since their SST are strongly influenced 

by intra- and interspecific sexual selection, and due to its wide 

distribution, they are exposed to a variety of environmental 

conditions. 

Three metapopulations of sympatric Calopteryx virgo meridionalis 

and Calopteryx xanthostoma were studied in the Iberian Peninsula, 

located at three different latitudes. Using principally discriminant 

analyses, latitudinal differences in size variables and SST were 

found. Size showed an increase southwards, although a slightly 

decrease was observed in the intermediate population. Regarding 

to SST, males showed an increase of wing spot southwards, C. 

xanthostoma females showed a decrease of pseudopterostigma 

corrected length northwards and meridionalis females showed an 

increase of wing pigmentation intensity southwards. Moreover, 

males and meridionalis females showed broader wings 

southwards. 

An environmental hypothesis better explains size clines, based on 

a converse Bergmann’s rule, as an adaptative response to 

latitudinal change of temperature. Clines in SST for both sexes are 

better explained by a balanced interaction hypothesis, which 

involves a balance between intra- and interspecific sexual 

selection forces. This hypothesis is explained by latitudinal 

differences in relative abundances, in such a way that the most 

abundant species would displace the other one. An ecological trait 

displacement would be produced in order to reduce reproductive 

effort costs. 

Predation of Honeybees by hornets 

Agnès Rortais 1 , Alexandros Papachristoforou 2 and Gérard Arnold 1 

1 

Laboratoire Populations, Génétique, Evolution CNRS, UPR 9034, 

Avenue de la Terrasse, 91198 Gif-sur-Yvette, France 

2 

Laboratory of Apiculture, Aristotle University of Thessaloniki, 

Greece 

Hornets are predators of honeybees worldwide. They attack 

colonies for proteins (bees) and carbohydrates (honey). However, 

as a result of co-evolution, honeybees which live in sympatry with 

such predators have developed efficient strategies to defend their 

nest. Conversely, species that have not evolved with these 

predators may be at risk when exposed to them as illustrated by 

the case of imported Italian honeybees (A. m. ligustica) to Asia 

(Ken et al. 2007). 

This study describes a case-study of co-evolution between the 

Cypriot honeybee Apis mellifera cypria and the oriental hornet 

Vespa orientalis. While thermo-balling has been described as a 

strategy developed by Asian honeybees (A. cerana) to kill their 

predator, V. mandarinia and V. simillima, another strategy namely 

asphyxia-balling has been found in Cyprian honeybees and is 

further presented (Papachristoforou et al. 2007). 

In the light of this study, discussion is made on the case of V. 

velutina which recently invaded France (Villemant et al. 2006), 

exposes honeybee colonies to great levels of predation, and may 

have an impact on colonies because local and naïve honeybees (A. 

m. mellifera) did not evolve with this predator. A better 

understanding of the way such prey-predator couples (A. mellifera 

spp-V. velutina) operate is necessary to prevent further honeybee 

colony losses in Europe.


The Dynamic epigenome, physiology, behavior and pathology 

Moshe Szyf 

Department of Pharmacology and Therapeutics, McGill University 

Montreal Canada 

Epigenetic patterns are sculpted during development and shape 

the diversity of gene expression programs in the different cell types 

of the organism. The epigenome of the developing foetus is 

especially sensitive to maternal nutrition, exposure to 

environmental toxins as well as psychological stress. Epigenetic 

alterations have the same phenotypic consequences as genetic 

differences. However, in difference from the genetic sequence, 

which is fixed, the epigenetic profile is somewhat dynamic. Thus, 

the responsivity of the epigenome to the environment continues 

throughout life. Exposure of the young rodent pup to different 

intensities of maternal care, differentially affects the epigenome 

and the behavior of the pup into adulthood. We will propose here 

a mechanism linking behavioral exposures such as maternal 

behavior and epigenetic programming. This mechanism illustrates 

a possible conduit between the external environment and the 

epigenome, which could explain epigenetic programming early in 

life as well as its dynamic nature throughout life. We will discuss 

the prospect that similar epigenetic variations laid down during 

early life play a role in generating inter individual differences in 

human behavior and we will present data from different human 

cohorts. We will illustrate how early childhood experience is 

marked in humans the brain and in blood cells. In summary, we 

propose that the epigenome mediates between the dynamic 

environment and our static genomes and provides a molecular link 

between nurture and nature between the social and chemical 

environment and phenotype. 

Control of female pheromones in Drosophila melanogaster 

Claude Wicker-Thomas 

Legs, UPR 9034, 91198 Gif sur Yvette, France 

Drosophila melanogaster produces sexually dimorphic 

pheromones, with C23 and C25 monoenes produced in males and 

C27 and C29 dienes produced in females. We have molecularly 

and functionally characterized the genes involved in their 

biosynthesis and have begun the study of their regulation. Female 

pheromones are controlled by hormones (ecdysone) and 

neurotransmitters (dopamine), which act on the second 

desaturation step leading to dienes. No such control has been 

evidenced in males. Here we review how this control can be 

exerted and present data on pheromones and courtship behaviour. 

Conditional response of feeding on zebra fish (Danio rerio) 

Zhongneng Xu 

Institute of Hydrobiology, Jinan University, 510632, Guangzhou, P 

R China 

Conditional response of feeding on zebra fish (Danio rerio) was 

investigated under laboratory conditions. Different light colors, 

different training time, and different levels of progesterone and MS- 

222 were set in this experiment. Conditional response of feeding 

on zebra fish - the fish swimming to the light side - induced by light 

can be established in two weeks. Red light affected more 

significantl! y the fish conditional response than yellow light and 

blue light. The male fish swam faster than females in the groups 

exposed to progesterone. In the treatments of adding MS-222, 

males’ speeds were also faster than females’ without photic 

stimulation and similar to females’ if photic stimulation was 

performed, but the speeds of both females and males were slower 

than those in treatments without MS-222. Additionally, another 

conditional response of feeding on zebra fish - the fish swimming 

to the side opposite to the light when it was turned on - induced by 

light could also be established. 

- 63 - 

Wing loading adjustment in mallards (Anas platyrhynchos): a 

generalization of the mass starvation predation risk theory in 

large birds? 

Cédric Zimmer, Mathieu Boos, Odile Petit and Jean-Patrice Robin 

Institut Pluridisciplinaire Hubert Curien, Département Ecologie 

Physiologie Ethologie, UMR 7178 CNRS/ULP, associé à 

l’Université Henri Poincaré - Nancy 1, 23 rue Becquerel, 67087 

Strasbourg Cedex 2, France 

In passerine birds, body reserves are optimized to minimize 

starvation and predation risks. To check this theory in larger birds 

with different body fuel storage strategies and higher starvation 

capacities, two groups of mallards (G1, G2) maintained in outdoor 

aviaries were disturbed (respectively 2x15 and 4x20 min daily) at 

one-month intervals during one-week sessions with a radiocontrolled 

car. Birds’ take-off flights, body mass, food intake and 

wing loading data were recorded and compared to an undisturbed 

control group. The number of take-offs was similar between 

successive sessions (P>0.5) and sexes (P>0.45), but was two-fold 

higher in G2 than in G1 (P


S18 - Interdisciplinary approaches in ecology: from individuals to populations and communities 

Red List project in France: an example with freshwater 

crustaceans 

Yoann Allanic 1 , Patrick Haffner 1 , Florient Kirchner 2 , Danielle 

Defaye 3 and Pierre Noël 3 

1 Muséum national d'Histoire naturelle, Service du Patrimoine 

naturel, DEGB, USM 0308, CP 53, 57 rue Cuvier, F-75231 Paris 

cedex 05 France 

2 Comité français de l’UICN (Union Internationale pour la 

Conservation de la Nature), 26 rue Geoffroy Saint-Hilaire, F- 

75005 Paris France 

3 Muséum national d'Histoire naturelle, DMPA / BOME -UMR n° 

5178, CP 53, 57 rue Cuvier, F-75231 Paris cedex 05 France 

The IUCN Red List is a tool providing information on the global 

conservation status of plant and animal species in the world. 

Based on a set of criteria and quantitative thresholds, the 

methodology used to evaluate species extinction risk has been 

developed through an extensive consultation process with the 

scientific community. Since 1963, the Red List helps Governments, 

NGOs and multilateral agencies to establish conservation priorities 

and make biodiversity-related decisions. In 2003, guidelines have 

been published to apply the methodology to countries or any subglobal 

entity. To fulfil its engagement as part of the Convention on 

Biological Diversity and to reach its target to halt biodiversity loss 

by 2010, France adopted in 2004 a national strategy for 

biodiversity and launched in 2007, the Red List project. It aims to 

measure species extinction risk and progress towards the 2010 

target. It is coordinated by the IUCN French Committee and the 

Muséum national d'Histoire naturelle, and based on a partnership 

with all key organisations for species conservation. After the first 

chapters on vertebrates to be published in 2008, the conservation 

status of invertebrate species has to be evaluated. We choose to 

illustrate the project with freshwater crustaceans, whose evaluation 

according to IUCN criteria will provide the first overview in France 

of the level of threat on these species. More than 4,200 crustacean 

species are known in France, of which about 1,270 live in 

freshwaters. The Red List project will allow to document and 

evaluate the situation of about 700 of these species. 

Interplay between trail recruitment and allee effects: the case 

of Tetranychus urticae 

Aina Astudillo Fernandez and Jean Louis Deneubourg 

CP231, Campus la plaine, Boulevard du triomphe, 1050, Brussels, 

Belgium 

Trail laying behaviour leading to recruitment is frequent among 

animals. This behaviour can result in collective migrations to 

unknown destinations. The interest of migrating collectively to an 

unsure destination is not obvious for subsocial species. We 

hypothesize that natural! selection of this type of migration is 

closely related to the existence of Allee effects on population 

growth. Through mathematical modelling (differential equations 

and stochastic simulations) we study how collective migration can 

influence the dynamics of a network of populations, when Allee 

effects are at stake. We focus our work on the two-spotted spider 

mite, a phytophagous pest of recognised agricultural importance. 

This sub-social mite systematically spins silk trails as it moves, 

which attracts its followers. Moreover, evidence suggests that its 

population dynamics are subject to Allee effects. Its aggregative 

behaviour under collectively spun webs is responsible for an 

enhanced survival in groups, which can result in a positive 

dependence between population size and population growth. 

Experimental data on migration and population dynamics of 

Tetranychus urticae is thus used to parametrise the models. Our 

results suggest that passive recruitment is only advantageous for 

species presenting some kind of Allee effect. Thereby we evidence 

a possible correlation between the selection of trail laying 

behaviour and the presence of Allee effects. 

- 64 - 

Movement Ecology of fire salamanders: Integration of 

ecological and genetic data and implications for conservation 

Shirli Bar-David 1 , Nir Peleg 2 , Ori Segev 2 , Naomi Hill 2 , Alan R. 

Templeton 3 and Leon Blaustein 3 

1 

Mitrani Department of Desert Ecology, Jacob Blaustein Institutes 

for Desert Research, Ben Gurion University, Sde Boqer Campus, 

84990, Israel 

2 

Institute of Evolution, Faculty of Sciences, University of Haifa, 

31905, Israel 

3 

Department of Biology, Washington University, St. Louis, MO 

63130-4899 USA 

Dispersal may be particularly important for the regional persistence 

of amphibian species that are structured as metapopulations and 

experience local extinctions. The conventional wisdom is that the 

fire salamander (Salamandra infraimmaculata), an endangered 

species in Israel, has strong pond fidelity – i.e. returns each year to 

the same site to breed – and low dispersal ability. This would 

result in a number of isolated populations rather than a linked 

metapopulation. We explored movements of fire salamander 

among breeding sites on Mt. Carmel, northern Israel, and the 

implications for population structure and persistence. During five 

breeding seasons (November-March) capture-recapture surveys 

were conducted around several breeding sites, and along unpaved 

roads connecting them. DNA samples taken from captured 

individuals were used for a genetic survey with microsatellite loci. 

Most of the recaptures were at the same site as the initial capture. 

However, several salamanders were recaptured at least one 

kilometer away from the first site. These movement distances are 

considerably larger than documented in the literature and indicated 

potential connectivity between breeding sites. These findings are 

also supported by the genetic analysis: assignment tests indicated 

potential migrants between breeding sites. We examined the 

potential implications of habitat fragmentation, i.e. isolation of a 

breeding site, on population persistence. The high probability of 

local extinction found by the analysis highlights the severe 

consequences of fragmentation. We conclude that in order to 

maintain a viable population of fire salamander in Mt. Carmel, local 

conservation units should include several breeding sites and 

landscape connectivity between them. 

Brooding and reproductive success in the hybrid sparrow 

Passer domesticus x P. hispaniolensis Linné, 1758 (Aves, 

Ploceidae), in Boudouaou 

Nassima Behidj-Benyounes 1 and Salaheddine Doumandji 2 

1 

Département de Biologie, Faculté des Sciences, Université de 

Boumerdès, 35000 Algeria 

2 

Laboratoire d’Ornithologie, Département de Zoologie Agricole et 

Forestière, Institut National Agronomique, El-Harrach, 16200, 

Algeria 

The hybrid sparrow is considered a pest bird species in Algeria. 

Several studies have been performed on this avian species and 

more particularly on its behaviour. 

In the present study, a study of the nesting of this bird has been 

carried out. We have followed the number of clutches and the fate 

of the eggs of the hybrid sparrow at Boudouaou (Algeria) during 

the course of 2 consecutive years, 2005 and 2006. 

The results show that the hybrid sparrow is a species which 

produces 4 egg clutches per year. The average percentage 

success per clutch varies from clutch to clutch and from year to 

year for the same brood period.


Important stopovers and limit of both breeding populations of 

Black storks in Europe 

Damien Chevallier, Yvon Le Maho and Sylvie Massemin-Challet 

IPHC, Département d’Ecologie, Physiologie et Ethologie, CNRS, 

ULP, 23 rue Becquerel, F-67087 Strasbourg Cedex 02, France 

Sixteen black storks (Ciconia nigra) were tracked by satellite 

during autumnal and spring migration in order to identify their 

important stopover sites. They were followed between breeding 

site in Europe and wintering site in West Africa from 1998 through 

2006. The data obtained during the migration have permitted to 

describe movement patterns, timing of migration, routes followed, 

speed, stopovers and their connections between them, in Europe 

and Africa. Among 29 important stopover sites identified, 5 were 

considered as important ones. 

It is known that the west and east breeding populations of black 

storks in Europe pass by two ways to go the African continent, 

Gibraltar for the first one and Bosphore Detroit for the second one. 

Our results show that birds start the spring migration from Africa, 

travel a distance equal to the “accessibility distance” (maximum 

distance covered without stopover) until Spain, stay on one 

stopover and go directly on a breeding ground. This “accessibility 

distance” used one or two times by birds corresponds exactly at 

the separation line between these two populations. 

This study highlights the importance of stopover location on the 

migration way for black storks. The protection of the most 

important stopovers seems to be essential in black storks and 

other bird species to assure good nutritional conditions during the 

migration. 

How physical oceanography helps in understanding the 

foraging ecology of the southern elephant seal 

Anne-Cécile Dragon 1 , Frédéric Bailleul 1 , Young-Hyang Park 2 and 

Christophe Guinet 1 

1 

Centre d’Etudes Biologiques de Chizé, CNRS, 79360 Villiers-en- 

Bois, France 

2 

Laboratoire d’Océanographie Physique, Muséum National 

d’Histoire Naturelle, 43 rue Cuvier, 75005 Paris, France 

Studies of predator movements contribute in understanding 

individual foraging processes. In the ocean, those spatial 

processes result from individual decisions taken in response to 

physiological but also environmental constraints. In the Southern 

Ocean polar frontal zone, where deep diving female southern 

elephant seal concentrate their foraging activity, mesoscale eddies 

structure and enhance the primary production with a likely effect 

on the spatial structure of top predator’s prey fields. The optimal 

foraging theory predicts that predators should adjust their 

movement’s pattern in relation to prey density. The aim of this 

study was to investigate the change in the movement and diving 

patterns of 4 adult female southern elephant seals equipped with 

temperature and salinity satellite-relayed data loggers. Argos 

tracks were analysed using a hidden Markov model: each step and 

turn of the animals’ trajectories was thus assigned to a behavioural 

state between two statistically distinct states: travelling and 

foraging. The first passage time technique applied on the duration 

of the bottom part of the dive allowed to identify favourable 

foraging zone at a finer scale. These favourable foraging zones 

were all found to be part of portion of the trajectory classified as in 

a foraging state. We finally correlated the detected foraging zones 

with several hydrographical variables such as sea level anomalies 

and the loggers’ temperature and salinity profiles. The most 

favourable foraging zones were related to the presence of both 

cyclonic and anticyclonic eddies. Elephant seals used the centre 

and edges of cyclonic eddy but only the edges of the anti-cyclonic 

ones and adjusted their diving behaviour accordingly 

- 65 - 

Mimetic interactions shape the ecological structure of 

butterfly communities 

Marianne Elias 1,2 , Zach Gompert 3 , Keith Willmott 4 , Julia Robinson- 

Willmott 4 and Chris Jiggins 2 

1 Institute of Evolutionary Biology, University of Edinburgh, West 

Mains Road, Edinburgh EH9 3JT; 2 Department of Zoology, 

University of Cambridge, Downing Street, Cambridge CB2 3EJ; 3 

Department of Botany, University of Wyoming, 1000 E. University 

Avenue, Laramie, WY 82072, USA; 4 McGuire Center for 

Lepidoptera Research, Florida Museum of Natural History, 

University of Florida, Gainesville FL, USA 

Niche phylogenetic conservatism is common at broad scales. 

Conversely, fine scale niche at community level is much more 

labile, due to complex interplay between common ancestry and 

interspecific interactions such as competition, predation, parasitism 

or mutualism. Yet surprisingly few studies have examined fine 

niche structure in communities of related, interacting species in a 

phylogenetic context. Here we use a multifaceted approach putting 

ecological data from a highly diverse community of mimetic 

butterflies in the upper Amazon into a phylogenetic perspective. 

Müllerian mimetic butterflies, where several unpalatable species 

have converged in wing patterns that advertise their toxicity to 

predators, are one of the most spectacular examples of mutualistic 

ecological adaptation. While the adaptive value of mimicry has 

been clearly demonstrated in the field and in the lab, an 

outstanding puzzle remains: up to ten distinct mimicry complexes 

may occur in the same community. If selection is so strong, why do 

communities not converge on a single colour pattern? Using 

microhabitat measurements for butterflies and their avian 

predators and a phylogeny of local butterfly species we show that 

1) mimicry complexes and their avian predators are partitioned in 

the ecological space in a similar way, and 2) community niche 

structure is likely the result of predation-driven selection, which 

overcomes common ancestry. Niche partitioning mediated by 

predation and possibly competition thus maintain species and 

mimetic diversity at the community level. We discuss the role of 

mimicry in diversification of groups of mimetic butterflies in light of 

these and recently published results. 

The Space-Lifetime Hypothesis: viewing organisms in four 

dimensions, literally. 

Lev Ginzburg and John Damuth 

100 North Country Road, 11733, Setauket, NY, USA 

Much of the debate about alternative scaling exponents may result 

from unawareness of the dimensionality appropriate for different 

data and questions; in some cases, analysis has to include a 

fourth temporal dimension, and in others, it does not. Proportional 

scaling simultaneously applied to an organism and its generation 

time, treating the latter as a natural fourth dimension, produces a 

simple explanation for the 3/4 power in large-scale interspecies 

comparisons. Analysis of data sets of reduced dimensionality (e.g., 

data sets constructed such that one or more of the four dimensions 

are fixed), results in predictably lower metabolic exponents of 2/3 

and 1/2 under one and two constraints, respectively. Our spacelifetime 

view offers a predictive framework that may be useful in 

developing a more complete mechanistic theory of metabolic 

scaling.


Status of terrestrial bird’s communities in cuban ecosystems 

during the winter residence 

Hiram González 1 , Alejandro Llanes 1 , Daysi Rodríguez 1 , Eneider 

Pérez 1 , Bárbara Sánchez 1 , Patricia Rodríguez 1 , Ramona Oviedo 1 y 

Alina Pérez 2 

1Instituto de Ecología y Sistemática, 2 ECOVIDA, Cuba 

In our study it was determined the influence of the different types 

of forests and regions in the distribution, composition, abundance 

and status of residents and migratory bird’s communities during 

the winter residence between the years 1993 and 2003. We 

worked in 33 localities of 10 regions of Cuba to evaluate the 

terrestrial bird’s communities. The methods of circular plots and it 

captures with ornithological mist nets were applied. It was 

evaluated the structure and composition of the vegetation types by 

means of the method of vegetation plots. We banded 13846 birds 

of 104 residents and migratory birds’ species and it was 

demonstrated by the counts and captures that there are significant 

differences in the richness and abundance of bird’s communities 

between sampling areas and the same ones they are given 

fundamentally by the migratory birds. The variables of the 

vegetation with more influence in the composition and abundance 

of bird´s communities are the height of the forest, the canopy 

covering, the density of trees and of bushes. The migratory birds 

presented a high fidelity to the winter territories, because 93.7% 

used the same microhabitat every year. The most important 

regions for the birds during the winter residence were Zapata 

Swamp, the Peninsula of Guanahacabibes, Coco Key, Pinares of 

Mayarí, Sabinal Key and the Alejandro of Humboldt Park, which 

are proposed as Importance Birds Areas. 

Status of birds of important birds area of Alejandro de 

Humboldt park, Cuba 

Hiram González Alonso 1 , Eneider Pérez Mena 1 , Daysi Rodríguez 

Batista 1 , Patricia Rodríguez Casariego 1 , Alejandro llanes Sosa 1 , 

Gerardo Begué Quiala 2 y Arturo Hernández Marrero 1 

1 

Instituto de Ecología y Sistemática 

2 

Unidad de Servicios Ambientales, Parque Alejandro de Humboldt, 

Cuba 

We have carried out evaluations of birds communities in five types 

of vegetation in the localities of Cupeyal del Norte, Piedra la Vela 

and Baitiquirí inside the National Park Alejandro of Humboldt, 

using the methods of surveys, circular plots and geo-referencing. 

The samplings were carried out in the months of December, 

January and March during 2005 to 2007. 

Based on species richness and relative abundance of their 

populations, we demonstrate considerable similarity among avian 

communities in the different forested ecosystems, relative 

abundance, diversity (H´) and the indexes of Simpson and 

Equitativity (J´) were always obtained in the evergreen forest and 

the pine forests. Also, the largest numbers of endemic and 

threatened species were detected in the evergreen forest and pine 

forests. We determined the distribution of the threatened species 

inside the Park, and in particular the abundance of pairs of Bee 

Hummingbirds (Mellisuga helenae). 

- 66 - 

Wild Felids densities in mountain ecosystems of Costa Rica: 

differential abundances and methodological issues 

Jose F. Gonzalez-Maya 1 , Diego Zarrate 2 , Catalina Amaya-Perilla 3 

and Jan Schipper 4 

1 

Proyecto de Conservación de Aguas y Tierras, Las Alturas, Coto 

Brus, Costa Rica 

2 

ProCAT Colombia, Calle 127 b # 45-76, Bogotá, Colombia. 

3 

ProCAT Colombia, Calle 127 b # 45-76, Bogotá, Colombia. 

4 

IUCN - Global Mammal Assessment. Virginia, USA. 

Mountain ecosystems are important areas for felid species 

conservation in Mesoamerica because they represent an important 

habitat for these animals and for general biodiversity. However, 

this habitats still mostly unknown about population, ecology and 

conservation status as a result of numerous causes - including 

logistics and isolation. The present study was carried out in the 

Talamanca region Costa Rica, an important mountain ecosystem 

of high endemicity and both biological and cultural importance for 

Mesoamerica. During 2006 and 2007, camera-traps were 

deployed in the study area in three sampling arrays on an 

elevation gradient (between 1000 and 2000 m). The minimum 

convex polygon covered by the cameras was 3.92, 19.08 and 

75.66 km 2 respectively. Different buffer distances were calculated 

using the data from each sampling and the integrated data of the 

maximum distance moved (MDM). Jaguar estimates vary from 

5.42 ± 2.30 to 11.54 ± 3.40, puma density vary from 4.30 to 44.01 

individuals/100 km 2 and ocelots density vary from 6,35 to 

15.90/100 km 2 . Differential densities and abundances were 

determined due to elevation and other environmental and 

ecological variables; including prey exploitation and habitat quality 

for each species. Current results indicate a good jaguar, puma and 

ocelot densities in the region however, the methodology still needs 

to be corrected for neo-tropical cat samplings and this type of 

ecosystems. A continued research is needed in the area in order 

to understand density patterns across the area and the influence of 

human variables on the distribution of felids and prey species. 

Effects of Experimentally Increased Stress Hormones levels 

on Survival in a long-lived bird: the Black-legged Kittiwake 

Aurélie Goutte 1 , Frédéric Angelier 2 , Borge Moe 3 , Claus Bech 3 and 

Olivier Chastel 1 

1 Centre d’Etudes Biologiques de Chizé-CNRS, 79360 Beauvoir 

sur Niort, France ; 2 Smithsonian Migratory Bird Center, National 

Zoological Park, Washington, DC 20008, USA; 3 Norwegian 

University of Science and Technology, NO-7491 Trondheim, 

Norway 

The life-history theory predicts the existence of a trade-off between 

the current reproductive episode and survival (i.e. future 

reproduction) but the underlying mechanisms are poorly 

understood. Often considered as the ‘stress hormone’, 

corticosterone (CORT) is thought to allow energy storing and 

adjust reproductive behaviour and physiology appropriately to the 

conditions encountered. Plasma levels of CORT rapidly rise in 

response to stressful conditions, which shifts energy investment 

away from reproduction and is believed to favour long-term 

survival. However, bearing elevated CORT levels over a prolonged 

period of time may compromise survival by impairing 

immunocompetence, cognitive abilities and increasing proteolysis. 

In the present study, we investigate the relationship between 

annual survival probability and experimentally increased 

corticosterone levels in Black-legged Kittiwake Rissa tridactyla. 

Corticosterone levels were higher during the two first days after 

implantation in CORT-implanted birds than in control ones, and 

then return to the initial level. Furthermore, CORT-implanted birds 

had a higher annual mortality rate than control ones. These results 

show that transitory high levels of corticosterone have strong 

consequences on survival. We suggest that the exogenous 

implantation of CORT might inhibit the endogenous secretion of 

CORT. This negative feedback would reduce the abilities of 

individuals to adopt an adequate behaviour in order to cope with a 

stressful perturbation, and thus would compromise survival.


Wildlife Depredation on Livestock in Medog County, Tibet, 

China 

Guangpu Guo 1 , Endi Zhang 2 , Pengju Wu 3 and Min Chen 2 

1 

School of Life Science and Technology, Tongji University, 

Shanghai, 200092 China 

2 

School of Life Science, East China Normal University, Shanghai, 

200062 China 

3 

Dongguan University of Technology, Dongguan Guangdong, 

523808 China 

The importance of the eastern Himalaya as an important center of 

biological diversity has long been recognized. Yarlung Tsangpo 

River flows eastward through southern Tibet before looping north 

then toward the plains of India. And it flows all through Medog 

County. The style of life and work affected the environment while 

the wildlife brings them damages. 

The main depredated livestock were cattle, horses, mules and pigs 

and the main predators was tigers, and also bears and wild dogs. 

Most depredations (91.37%) occurred in 1990s and in Gedang 

Xiang. There were 97.37% families who suffered from losing 

livestock. Even though the depredation occurrences were not 

significantly different between months and seasons, they showed 

an increasing trend from spring to winter. 

Horses and mules were depredated most by tiger during 

1990~1999 with an average of 10.60 ± 10.15 heads per year, and 

it showed a significant difference among months with the most 

occurrences in November (18.82%) and the least from August to 

September (1.18%, 2.35% and 1.18% individually). Cattle also 

killed mostly in November but less in May (0.77 ± 1.91) and 

September (1.14 ± 1.05). Livestock were preyed by tiger mainly in 

grassland and forest-grass with cattle also in forest-shrub-grass 

environments. There were 29.47% depredated livestock dragged 

by tiger for a certain distance. 

Wild dogs were often in groups of 2~7 individuals and mainly 

preyed on cattle and pigs; black bear preyed mainly on cattle. It is 

supposed that there would be 5 tigers in Gedang Xiang in those 

years. 

Mediterranean molluscs: an inventory in Port-Cros national 

park, France 

Elsa Marangoni and Pierre Noël 

Muséum national d'Histoire naturelle, DMPA / BOME -UMR n° 

5178, CP 53, 57 rue Cuvier, F-75231 Paris cedex 05, France 

Molluscs appeared some 600 MY ago and started diversification. 

Eight classes are recognized presently viz. Solenogastra, 

Caudofoveata, Polyplacophora, Monoplacophora, Gastropoda, 

Cephalopoda, Bivalvia and Scaphopoda. most of them 

represented within the 2045 species or so recognized to be 

present in the Mediterranean. This work presents preliminary 

results about the inventory of molluscs Port-Cros national park and 

its surroundings. 

A collection gathered by Priest Olivier all along the second half of 

XIXth century in Porquerolles Island was studied first; it includes 

some 274 species. It is one of the few shell collections from XIXth 

century for Mediterranean. 

Quite a number of shells were also obtained from the "Miladou" 

wreck, a roman ship, lying 42 m deep since 100 yrs BC in the Port- 

Cros national park. When she sank, some of the shells were on 

board (obvious for species from Red Sea), and some others could 

also have settle there as larvae later on. Subsequently, octopuses 

living in amphora brought also their own contribution... A total of 36 

species were present. It is well known that Humans were 

interested long ago by sea-shells for different reasons: food, 

decoration, jewel, money, religion... The question of the origin of 

these shells will be discussed. 

Our own findings brought other species not reported yet from the 

studied area. This work is a contribution to the knowledge of 

marine biodiversity in southern France 

- 67 - 

Vertebrates of Buçaco’s National Forest, Centre of Portugal 

Milena Matos and Carlos Fonseca 1 

Department of Biology/CESAM, University of Aveiro, 3810-193 

Aveiro, Portugal 

Buçaco’s National Forest (wall fenced area with approximately 105 

ha) constitutes an exclusive natural heritage in Portugal and in the 

World, due to its history, architecture and nature values. 

Biologically, it represents one of the best dendrological collections 

in Europe. However an unknown fauna is associated to the 

amazing and well-described flora diversity. The only faunistic study 

of this forest took place more than 100 years ago, comprising a 

survey/inventory of the animals present at that time. 

The vertebrate species (except Chiroptera) identification as well as 

the assessment of each vertebrate local distribution was 

conducted in this forest during the last three years. To reach these 

goals, several methodologies were applied for each group of 

vertebrates. 

117 species (fishes: 5, amphibians: 10, reptiles: 14, birds: 66, 

mammals: 22) were confirmed in this forest ecosystem and their 

distribution mapped. 

The occurrence of several protected and endemic species from all 

presented vertebrates groups (e.g. Chondrostoma oligolepis, the 

gold-striped salamander Chioglossa lusitanica, the Schreiber’s 

green lizard Lacerta schreiberi, the booted eagle Hieraatus 

pennatus and the Lusitanian pine vole Microtus lusitanicus) 

reinforces the importance of this natural area in terms of 

Portuguese fauna’s conservation. 

This study will provide the basis for further investigations focused 

on regional biodiversity and habitat fragmentation, among other 

subjects. 

The knowledge and science education of Buçaco’s National 

Forest’s biodiversity can contribute to the environmental 

sensitization of the visitors (more than 100 thousand/year) in what 

concerns to the maintenance of natural balances and the 

protection of ecosystems. 

Does Thermo-period influence growth and activity of the asp 

viper? 

Catherine Michel and Xavier Bonnet 

CEBC CNRS, UPR 1934, Villiers en Bois 79360 Beauvoir sur Niort, 

France 

Daily and seasonal variations of ambient temperatures determine 

activity and metabolism of animals. The vast majority of species 

are ectotherms and cannot maintain constant their body 

temperature through physiological means; the effects of ambient 

temperatures apply with force in these organisms. We 

experimentally examined the influence of thermo-period 

(temperature fluctuations) on the behaviour and physiology of a 

reptile, the aspic viper (Vipera aspic). This snake faces strong 

natural daily and annual temperature variations. During three 

months, we placed four groups of snakes under 4 contrasted 

thermal regimes: 1) a 2400h natural regime, 2) a 1200h 

accelerated regime, 3) a slow 4800h regime, and 4) a stable 

regime. For the first 3 groups, the ambient temperature fluctuated 

from 19°C to 29°C (mean=23°C) to mimic daily variations recorded 

during the peak of activity in the field (e.g. May). For the fourth 

group, the temperature was maintained at 23°C. Overall, the mean 

temperature over the whole experiment (3 months) was of 23°C for 

each of the four groups. For that, we used four climatic chambers 

where both temperature and hygrometry were controlled. We 

recorded locomotor activity, feeding success, digestion efficiency 

and growth rates. There was no significant effect of thermo-period 

on activity (p=0.163), digestion efficiency (p=0.850), ecdysis 

frequency (p=0.290), body mass changes (p=0.982) and growth 

rate (p=0.999). This study suggests that mean temperatures might 

be determinant, but not fluctuations. This notion challenges the 

simplistic paradigm that suggests that maximal body temperatures 

chosen by reptiles correspond necessarily to thermal optimum.


Reproductive potential of an entomopathogenic nematode, 

Steinernema feltiae, on different hosts (Galleria mellonella, 

Hyphantria cunea and Leptinotarsa decemlineata) 

Nona Mikaia 

Dept. of Biology, Sukhumi University, 9, Jikia, Tbilisi 0186, Georgia 

Research into the comparative efficiency of reproduction of an 

entomopathogenic nematode, Steinernema feltiae, on several 

insect species (the wax moth Galleria mellonella, the fall webworm 

Hyphantria cunea and the Colorado potato beetle Leptinotarsa 

decemlineata) has been carried out. Experiments were performed 

under laboratory conditions at 25 0 С and 75% relative humidity. For 

nematode (n) infection of these insect by S. feltiae, 150 insects 

were used, with 10 individuals of each species in a total of 15 Petri 

dishes. Consequently, insects of each species were present in 5 

Petri dishes, using nematode densities of 10 n/ml, 20 n/ml, 50 n/ml, 

100 n/ml and 120 n/ml in the 5 dishes. At such concentrations the 

nematode suspension has been applied to each insect species. 

Death of the insects occurred within 48 hours. After 48 hours the 

dead insects were placed on a white trap and the nematodes 

started replication. The output increased with time, with invasive 

nematode larvae emerging from the host insect into distilled water 

during 12 days from G. mellonella, 11 days from H. cunea, and 14 

days from adults of L. decemlineata. Continuous records of 

nematode emergence from the dead insect’s bodies showed that 

the greatest numbers (91.8%) of nematodes were obtained on 

days 4-7 from larvae G. mellonella, 94.5% on days 2-7 from larvae 

of H. cunea, and 98.4% on days 1, 2, 4 and 7 from L. 

decemlineata adults. 

On the basis of the data for the reproduction of S. feltiae on its 

generally accepted host G. mellonella, it is concluded that 

laboratory populations of H. cunea and L. decemlineata may 

alternatively be used. 

Raven’s (Corbus corax) body temperature change during 

annual life cycle 

Nadejda I. Mordosova and Andrey I. Anufriev 

Institute of Biological Problems of Criolitozone SD RAS, Lenina pr., 

41, 677980, Yakutsk, Russia 

Yakutia is situated in the north-east of Siberia and it is one of the 

coldest regions in Russia. Annual range of temperature changes is 

over 100ºC according to absolute value, in winter temperature falls 

to - 60 0 C. The data presented in the report show raven’s body 

temperature changes during annual life cycle in conditions of 

natural temperature background. The thermograph DS 1922 L-F5 

was inserted into its front part of body between wings. The body 

temperature was measured once in 60 minutes. 8062 

measurements of temperature were analyzed during 11 months. 

During the annual cycle raven’s body temperature was stable and 

it remained on a relatively high level. The average day body 

temperature changed in the following way: the highest temperature 

was registered in summer months: May (39.63 ± 0.044°C), June 

(39.89 ± 0.040°C), July (39.62 ± 0.096°C), after the average day 

temperature was lowering, reaching the minimum temperature in 

January (38.51 ± 0.057 0 C). The absolute low body temperatures of 

raven (35,5-36°C) were registered at night time from December to 

February, the highest temperatures (42.0-42.5°C) were registered 

in daytime during the whole year. Annual dynamics of the average 

daily body temperature had the same direction with the 

temperature of environment. The daily dynamics of body 

temperature during the whole year remained the same: high 

temperature in daytime and low temperature at night. In summer 

months the body temperature was higher than in winter according 

to absolute value. The daily dynamics in winter and summer were 

5-6°C at absolute value. Thus, the raven’s body temperature 

remains on a relatively high level during the whole year. This 

species demonstrates high degree of adaptation to the wide range 

of environmental temperatures in cold climatic conditions of 

Yakutia. 

- 68 - 

Quantifying the dynamics of marine invertebrate 

metacommunities: what processes can maintain high 

diversity with low densities in the Mediterranean Sea? 

Charlotte Moritz 1,2 , Nicolas Loeuille 3,4 , Jean-Marc Guarini 1,2 and 

Katell Guizien 1,2 

1 UPMC Univ Paris 06, UMR 7621, LOBB, Observatoire 

océanographique, F-66651, Banyuls/mer, France ; 2 CNRS, UMR 

7621, LOBB, Observatoire océanographique, F-66651, Banyuls/ 

mer, France ; 3 UPMC Univ Paris 06, UMR 7625, Ecologie et 

Evolution, F-75005, Paris, France ; 4 CNRS, UMR 7625, Ecologie 

et Evolution, F-75005, Paris, France 

The Mediterranean Sea hosts 5.6% of the world benthic 

invertebrate species on 0.82% of the ocean surface. In addition, 

Mediterranean ecosystems are characterized by low densities (and 

biomasses) compared to other oceans, a feature often attributed to 

their oligotrophic environment. Oligotrophic conditions can induce 

reduced growth rates and increased mortality rates through 

increased competition for food between individuals. To study the 

diversity vs. density patterns in benthic invertebrate populations, a 

theoretical model was developed. This model simulates benthic 

coastal ecosystems, composed of a mosaic of habitats, in which 

populations of different species of invertebrates are structured in a 

metacommunity. The connectivity between communities is 

ensured by passive larval dispersal favoured by exchanges 

between benthic and pelagic compartments due to hydrodynamics. 

In the Mediterranean Sea, the microtidal regime induces a lower 

and more variable coastal connectivity compared to macrotidal 

basins. The model describes minimal population dynamics 

including basic processes (growth, mortality, reproduction and 

effects of interactions between individuals) and incorporating 

fluxes of larvae (finally recruited as juveniles) between the patches 

of the metacommunity. Mathematical analysis of model properties 

revealed that negative interactions (intra- and interspecific 

competitions) have a stabilizing effect on interacting organisms 

when gains by recruitment are higher than losses by mortality. 

In addition, low mortality rates and low connectivity which 

decreases negative local interactions maintain high regional 

species diversity with low local densities. This property suggested 

that oligotrophy cannot be the only factor leading to the high 

diversity-low density pattern observed in the Mediterranean Sea. 

Use of intensive sampling methods for evaluating wolf 

abundance and population trends in small areas: possible 

applications in management at broader scales 

Vicente Palacios 1 , Victor Sazatornil 2 , Ana Guerra 3 , Mónica 

Almeida 4 and Emilio J. García 1 

1 

A.RE.NA. Asesores en Recursos Naturales, S. L., C/ Perpetuo 

Socorro, 12 - entlo B, 27003, Lugo, Spain 

2 

C/9, nº 69, Turó de Sant Pau, 08193 Cerdanyola del Vallès, 

Barcelona. Spain 

3 

R. Manuel Marques, 15-1-frente. 1750-169 Lisboa. Portugal 

4 

R. António José de Almeida. Lote 1137-A, Quinta do Conde 2, 

2975-316 Quinta do Conde. Portugal 

Animal abundance and population trends are two of the most 

important parameters for the conservation and management of a 

species. Estimating animal abundance requires standardized 

application of a technique allowing researchers to detect changes 

with an appropriate accuracy. Sierra de la Culebra is a hunting 

reserve in north-western Iberian Peninsula. We conducted an 

intensive survey consisting in sampling transects along roadways 

in part of the reserve to detect wolf scats in the winter of 2008. In 

this small area, five wolf litters were born in the summer of 2007. 

The study area was divided into 51 square cells (4 km 2 ) and the 

sampling transects covered 3.5 km/cell. The transects were 

sampled in February and four weeks later. We obtained an index 

reflecting the rate at which scats are deposited along established 

roadways (0.019 scats/km/day). Representing the scat index by 

cell and analyzing spatial distribution of scats, we could define 4 to 

5 areas with higher scat concentrations, corresponding to the 

places where the wolves were more frequently seen during the 

winter. These results suggest that these intensive surveys in small 

areas could be useful in estimating the number of packs living in a 

determinate area.


We are investigating if the scat abundance index obtained could 

be used in estimating relative abundance, allowing 

us to obtain an accurate estimation of wolf population trends. 

Moreover, selecting several small areas of interest for the 

conservation and management of wolves, this method could be 

useful in monitoring population trends over large areas. 

Range development of a reintroduced deer (Cervus elaphus) 

population in Southeast of Portugal 

Rosana Peixoto 1 , António Mira 1 and Pedro Beja 2 

1 Biology Department - Conservation Biology Unit, University of 

Évora, Pólo da Mitra, 7002-554 Évora, Portugal 

2 ERENA – Natural Resources Management, Rua Robalo Gouveia 

1-1A 1900-392 Lisboa - Portugal 

As the number of endangered species grow, the use of 

reintroduction as a conservation tool against species extinction 

increases. Translocations of common deer species, as the red 

deer (Cervus elaphus), may also help improving our understanding 

of reintroduction requirements and planning the reintroduction for 

endangered ungulates. A reintroduction program has been 

developed in the Herdade da Coitadinha –Noudar Nature Park, 

Portugal and includes an intensive post release monitoring for 

three years. This present work focus on the home range sizes and 

spatial preferences on the 11 radio collared females red deer after 

the first 13-months restocked monitoring program. The animals 

were tracked twice a day two or three times per week between 

December 2006 and December 2007. Displacement activity was 

monitored by radio telemetry (locations calculated by L.O.A.S. 

software) and personal observation. MCP (Minimum Convex 

Polygon) was used to calculate the total area and Kernel to 

describe home ranges. Spatial preferences were determined by 

QGis – GRASS software. The monitored deer territory had shown 

average values of 9.56 km 2 . The total territory covered by the 

eleven females was 19.99 km 2 . The maximum territory was 11.36 

km 2 and the minimum 6.97 km 2 . Main occupation area remains 

inside the farm, southeast/northwest direction. This area is 

characterized by the presence of denser stands of Holm oak, 

grasslands and scrublands and also Holm oak open woods. During 

reproduction period most of the mature females demonstrated 

preference to a not disturbance eucalypt stand area located 

northeast from the release point. 

Impact Assessment of Wind Turbines over local winged 

vertebrate communities 

Nuno Pinto 1 , Milena Matos 2 and Eduardo Ferreira 3 

University of Aveiro, 3810-193 Aveiro. Portugal, 3810-193, Aveiro, 

Portugal 

The study here presented refers to the update of bats and birds 

checklist and monitoring campaigns carried out in the 

implementation area of a power plant at the Natura 2000 site from 

Serra da Lousã. The power plant includes 25 wind turbines placed 

through an NE-SW axis, at two different locations, around 

1200m.a.s.l. Presented results refer to the period preceding the 

construction of the power plant as well as the first year of activity. 

A checklist of species using the area before and during 

construction period was compiled for the build up of a reference 

state. This should allow the assessment of anthropogenic impact. 

The bird and bats checklists include four and five threatened 

species or higher status by IUCN, respectively. For chiropters, a 

supplementary area of 10Km radius was searched for shelters. 39 

potential shelters were found and, out of these, 12 were monitored 

for six months. After construction of the power plant, use of the 

space by monitored species as well as mortality rates and other 

impacts were monthly assessed. Until now, no considerable 

adverse impacts were detected for both groups. During the first 

year of activity, estimated mortality was zero individuals. 

Monitoring will continue for two more years. However, data 

collected until the moment suggests that this power plant does not 

present a risk for winged vertebrates using the area. If these 

results stand for the following years, we could consider this case 

study as a successful implementation of “effectively clean” energy 

production. 

- 69 - 

Fauna of golf courses: the example of the National Golf 

(France) 

Rose-Line Preud'Homme 1 , Jean-Philippe Siblet 1 , Nathalie 

Machon 1 , Jérôme Paris 2 and Pierre Noël 3 

1 Muséum national d'Histoire naturelle, Service du Patrimoine 

naturel, DEGB, USM 0308, CP 53, 57 rue Cuvier, F-75231 Paris 

cedex 05, France ; 2 Fédération Française de Golf, 68 rue Anatole 

France, F-92309 Levallois-Perret cedex, France ; 3 Muséum 

national d'Histoire naturelle, DMPA/BOME - UMR n°5178, CP 53, 

57 rue Cuvier, F-75231 Paris cedex 05, France 

Playgrounds are highly frequented areas, often strongly managed. 

There biodiversity is thus considered very poor. In order to assess 

biodiversity of Golf Courses, the French Golf Federation and the 

Muséum national d'Histoire naturelle decided to carry out 

inventories of several taxonomic groups in one of the most famous 

golf courses in France: the National Golf at Saint-Quentin near 

Paris. This golf was established 20 years ago in an agricultural 

landscape, a habitat usually hosting a low animal biodiversity. 

Since then, artificial ponds and lakes have been created; trees, 

bushes and flower beds have been planted. 

The present investigation established the presence of a richer 

fauna than expected. We found mammals (wild boars, fox, bats, 

moles, rabbits, hares, shrews...), birds (at least 80 species among 

which kingfishers, cormorants, warblers, tits,...), amphibia (frogs, 

toads, newts), fishes (white amur carps, pumkinseeds), insects 

(many species including beetles, butterflies, moths, grasshoppers, 

dragonflies, dipterians, ants...), molluscs (snails and slugs), 

spiders, crustaceans (woodlices), centipedes, and plancton 

(different zoological groups). 

In conclusion, it appears that the lesser anthropized grounds of the 

National Golf Course could be at least as rich as the surrounding 

habitats. Furthermore, since hunting and fishing is not permitted, 

wildlife is assumed to be protected on these areas. However, the 

intensive management of other parts of the golf could have a 

negative influence on the dynamics of animal populations. The 

inventories we performed will help to define the best management 

actions for biodiversity preservation. The question of the Golf 

Course as ecological corridor among natural habitats will also be 

discussed. 

The study of the gene H2 and microsatellites located within tcomplex 

region from wild populations of house mice (Mus 

musculus) 

Larisa D. Safronova, Sergey G. Potapov, Anna I. Chekunova and 

Varos G. Petrosyan 

A. N. Severtsov Institute of Ecology and Evolution RAS, 117071, 

Moscow, Leninski pr., 33, Russia 

The proximal region of the chromosome 17 in house mouse 

includes the complex system of genes which charactereterized by 

four non-overlapping paracentric inversions span a 20cM region. 

The gene H2 and microsatellite analysis of mice from wild 

populations of Russia (Volgograd, Rostov, Saratov areas and 

Kalmykia), from another states (Armenia, Bulgaria, Iran, Mongolia) 

and also of laboratory stocks of house mice carrying t-haplotypes 

(t 12 , t w5 , t w12 , t w73 ) has been carried out using PCR method. Specific 

for t-carrying mice allele variant of the exon 2 of the MHC class II 

A beta gene has been found out in laboratory stock tw 12 , 2 wild 

mice from Kalmykia and 1 mice from the Saratov area. The 

microsatellite amplificated are DNA sequences that located at the 

chromosome 17 in the region from 7,6 till 18,8 cM. Each pair of 

primers of D17Mit (16, 21, 23, 28, 32, 57, 63, 78) recognized from 

3 to 6 nucleotide sequence variants of different size. The patterns 

of microsatellite DNA fragments were specific for each specimen, 

and in the most of cases two variants of nucleotide sequences 

have been detected. The variants of patterns of microsatellite 

nucleotide sequences from the laboratory stocks were the most 

similar to Volgograd and Rostov regions while those from Bulgaria 

and Saratov region were similar to each other. Interestingly, 

microsatellite specimens from Armenia and Iran significantly differ 

from all other patterns. Thus variants of nucleotide sequences 

located in t-complex region can be used as markers for 

identification of Mus representatives from wild populations. This 

study was supported by RFBR 06-04-48866.


The Lusitanian pine vole and the Mediterranean pine vole in a 

sympatry area: using presence signs to discriminate among 

similar species 

Sara M. Santos 1 , António P. Mira 2 and Maria Luz Mathias 1 

1 Centre of Environmental Biology, Department of Animal Biology, 

Faculty of Sciences, University of Lisbon, Campo Grande, Lisbon 

1749-016, Portugal 

2 Unit for Conservation Biology, Department of Biology, University 

of Évora, Pólo da Mitra, 7002-554 Évora, Portugal 

The Lusitanian pine vole (Microtus lusitanicus) and the 

Mediterranean pine vole (Microtus duodecimcostatus) are sister 

species with an allopatric occurrence pattern and a narrow band of 

potential sympatry in central Portugal. Our aim was to determine if 

presence signs can be used to discriminate the presence of the 

two species in an area of sympatry (Northern Alentejo) and, if so, 

which characteristics achieve best classification accuracy. 

A total of 175 trapping plots were sampled across the study area. 

Previous to traps placement, ten presence signs were randomly 

selected for measurements of three variables: mean diameter of 

soil mounds, proportion of entrance holes (number of presence 

signs with entrance hole / number of presence signs) and mean 

diameter of entrance holes. 

Based on a classification tree analysis, our results show that 

presence signs can be used to discriminate between the two 

species of pine voles in the studied sympatry area. The 

characteristic that most discriminates the groups is the proportion 

of entrance holes: plots with > 0.8 correspond to M. lusitanicus 

presence (i.e. mostly entrance holes with few or none soil mounds 

present), while plots with < 0.8 correspond to M. duodecimcostatus 

(i.e. mostly soil mounds with few or none entrance holes). 

Competition, historical geography, and community assembly 

rules – birds of the Solomon Islands 

Daniel Simberloff 1 and Michael Collins 2 

1 

University of Tennessee, Knoxville, Tennessee 37996 USA 

2 

Hampden-Sydney College, Hampden-Sydney, Virginia 23943 

USA 

The birds of the Solomon Islands were crucial in the development 

of community assembly rules, but their distribution patterns have 

not been rigorously examined to consider alternative hypotheses 

for which species are found on which islands. A detailed study of 

checkerboard distributions – pairs of species that never occupy the 

same island – suggests that Pleistocene and current geography 

combined with dispersal limitation may well explain the great 

majority of them. Interspecific competition is rarely a cogent 

explanation for such non-overlapping distributions in this system. 

In fact, the checkerboard metaphor does not represent Solomon 

Islands bird ranges; rather, mutually exclusive, ecologically similar 

species pairs are almost always regionally allopatric, and the 

boundaries between their ranges almost always correspond to 

Pleistocene and present dispersal barriers. 

Current status of black-capped marmot in North East Siberia 

Nickita G. Solomonov 

Institute for Biological Problems of Cryolithozone SB RAS, 41 

Lenin avenue, 677891 Yakutsk, Russia 

Black-capped marmot Marmota camtchatica Pall from Yakutia is 

represented with two populations of subspecies forms. The 

Yakutia subspecies M. c. bungei Kastch inhabits North East 

Yakutia occurring in mountain ecosystems of the Moma, Chersky, 

Verkhoyansk mountain-ridges reaching the Yana and Lena 

downstreams along the Kharaulakh Range. Baikal marmot 

(Barguzin) M. c. doppelmayeri Birula occurs in the upstream of the 

Aldan in the Zverev mountain-ridge, in the basins of the Olekma, 

Tokko and Chara. In the 50-60s of the last century distribution and 

biology of Yakutia marmot were carefully studied by Kapitonov, as 

for Baikal subspecies we have only scanty information. We 

gathered detailed evidence on number and state of Yakutia 

subspecies during All-union census of marmot number in 1984. 

- 70 - 

According to these data endangered Kondek population of Yakutia 

marmot and south-Yakutia population of Baikal subspecies 

entered on the YASSR Red Book in 1987. Further wildlife 

biologists of our Institute – Yu. Revin, Yu. Lukovtsev, N. 

Solomonov, V. Vasiliev, I. Oklopkov, F. Yakovlev including the 

researchers from other institutions – N. Zheleznov, V. Lyamkin, G. 

Boyeskorov studied different aspects of black-capped marmot 

biology and distribution. Vasiliev and Semenov defended Ph.D. 

theses on marmot hibernation and characteristics of its biology. 

Because of decline in numbers and fragment spreading of both 

Yakutia black-capped marmot subspecies they joined in the Red 

Book of Sakha in 2003. Black-capped marmot is the most 

important intermediate between marmots of the “marmota” and 

“lobec” groups. This gives rise to the interest for researchers 

concerning to its origin and settling during a historical span. By 

level of morphological differentiation black-capped marmot 

subspecies M. c. kamtchatica and M. c. doppelmayeri especially 

differ at the level of independent species (Kapitonov 1978; 

Baryshnikov, et al. 1981; Gromov, Erbayeva 1995). Important 

immune-genetic differences are found in all three marmot 

subspecies (Tolnerovskaya, et al. 1990). We assume that G. 

Boyeskorov’s proposal (1999) to regard Marmota camtchatica a 

superspecies is reasonable 

Ecophysiological adaptations of small mammals to the 

conditions of the cold climate (9) 

Nickita G. Solomonov 1 , Andrey I. Anufriev 1 , Alexander K. 

Akhremenko 1 , Tatyana N. Solomonova 1 , Vladimir N. Vasiliev 2 , 

Innokenti M. Okhlopkov 1 and Victor T. Sedalischev 1 

1 Institute for Biological Problems of Cryolithozone SB RAS, 41 

Lenin avenue, 677891 Yakutsk; 2 Department of Biological 

Resources MNP RS(Ya), 18, Sverdlov str., 677005, Yakutsk 

Already A.F. Middendorf (1869) indicated the most important 

adaptations of Siberian animals to cold: well-developed fur cover, 

ability to store fat reserves, adaptive behavior to the conditions of 

cold. In the 20th century schools of thought headed by physiologist 

A.D. Slonim and zoologists N.I. Kalabukhov and S.S. Schwartz 

contributed to the development of this problem. 

Our research on adaptation of animals was set in the Yakut State 

University and Institute of Biology Ya.B. SB USSR Academy of 

Sciences in the 60s-early 70s of the last century. The main objects 

of investigation are representatives of Sciuridae: Marmota 

camtschatica, Spermophilus undulatus, Spermophilus parryi and 

Tamias sibiricus; voles: Clethrionomus rutilus, Microtus gregalis, 

Microtus hyperboreus, Microtus oeconomus, synanthropic rodents: 

Ondatra zibethica, Mus musculus, Rattus norvegicus. 

Some interesting features, metabolism level and content of 

biologically active substances in tissues such as vitamin A and C 

were studied. A particular attention was given to the study of 

spatial and biological structure of populations, pattern of 

colonization over the territory, food and protective conditions, 

features of dynamics number. It has been determined that 

abundant species adapted to the conditions of the cold climate due 

to active metabolism, well developed chemical and behavioral 

thermoregulation, variability of morphophysiological parameters. 

The aboriginal forms adapt to cold at the expense of deeper 

physiological adaptations because of highly developed physical 

thermoregulation and strong seasonal metabolism changes. A 

disorder in the functioning of eco-geographical regulations 

according to Bergman and Allen causing a hypobiotic state has 

been observed in the forms most adapted to the cold. Very similar 

are the trends of physiological-biochemical variability during winter 

hibernation of small mammals and hypobiotic states in large 

mammals (Yakut horse, moose, reindeer).


Siberian chipmunk Tamias sibiricus (Laxmann, 1769) in 

Yakutia 

Tatyana N. Solomonova 


Lenin avenue, 677891 Yakutsk, 

Chipmunk is a common forest species. In Yakutia the northern 

boundary of its range goes along the latitude 67-69N reaching 71N 

in the downstream of the Lena, 69-70°N in the Olenek River basin, 

69 along the Yana and the Indigirka and 67 30' in the Kolyma area. 

The animal does not occur in the northern thin forests. In the 

mountain of the north-east of Yakutia the species inhabit the sites 

at the altitude 1300 m a.s.l. Most modern researchers consider 

that morphological parameters of the Yakutia chipmunk are within 

the size limits of the nominal subspecies T. sibiricus Laxmann. The 

Yakutia subspecies s. jacutensis Ognev, described in 1936 has 

slight differences, mainly in skin coloring. Chipmunk occupies a 

peculiar niche in the forest ecosystems: unlike a tree squirrel 

species the animal is a wood-shrub form, inhabits sparse forests 

with shrub growth occurring often in the outskirts, makes hollows 

and dwelling places on the ground under tree roots, under fallen 

trees, in holes not high from the ground. It feeds on seeds of 

woody-shrub plants, dog-rose, many herb species, berries, 

Vaccinium vitis-iedae, in particular. Larch and pine seeds the 

chipmunk gets from the fallen last year cones. 

In spring until mid-summer it consumes much greenery. Population 

is maintained through reproduction of one but large-sized litter. 

Regular hibernation is characteristic of the Yakutia chipmunk 

population though it is less expressed unlike larger ground 

squirrels and marmots. On the other hand Siberian chipmunk 

hibernation is noted for a longer period than in north-American 

chipmunks where the climate conditions are more favorable. It 

lasts 5 months with repeated arousals in 7-8 days. The 

temperature dependence of metabolism rate has been studied in 

hibernating chipmunks within the temperature range -5 to 9 °C. 

The least metabolism rate is fixed at the temperature 3-5 °C; 

amounting 0, 049±0.0017 ml O2/g.hr. Decrease of the ambient 

temperature causes a sharp rise of oxygen uptake. Minimal mass 

losses are recorded at 3-5 °C, the longest periods of hypothermia 

proceed in the same intervals. Duration of a torpid state was much 

shorter in the artificial hole with colder temperature conditions. 

Bats (Chiroptera, Vespertilionidae) in Yakutia (north-eastern 

Siberia) 

Tatiana N. Solomonova and Andrey I. Anufriev 



There are 5 Chiroptera species of Vespertilionidae family recorded 

in Yakutia. Eptesicus nilssoni is spread over the whole southern 

half of the territory. In the Lena River basin Northern bats are 

found in the Buotama (tributary) Mouth and in the vicinity of 

Olekminsk. The Vilyui basin is rich in this species all along this 

river. East of the Lena the Northern bat is sighted in the upper 

course of the Tamma; the Ungra River (left confluence of the 

Aldan); easternmost of the Zapadny Yangi Mountains; lower 

course of the Maya. The northernmost point of this bat finding in 

Yakutia is the Vilyui River basin in its midstream in the place of the 

Markha River confluence (63 0 30’ N, 119 0 E). Information related to 

the Northern bat habitat in the Kolyma River area has not been 

confirmed. Plecotus auritus within the limits of Yakutia has been 

reported in the Tokko River valley near the Tyanya outfall; the 

Tokko valley; on the riverside of the Amga; this species animals 

were regularly found among hibernating bats in the gypsum pit. 

The northernmost finding of the Common long-eared bat in eastern 

Siberia is the neighborhood of Churapcha settlement (62 0 N; 

132 0 30’ E). In Yakutia, Myotis brandti was first recorded during 

wintering in the gallery of the gypsum pit near Olekminsk (60 0 30’ N; 

120 0 30’E) in November, 2002. 

In 2006 one specimen of Brandt’s bat was found in summer 100 

km south of its wintering range. All places of Мyotis daubentoni 

finding are in the southern sector of Yakutia. The northern 

boundary line of the area for this species runs along 61-62 0 N. The 

animal was recorded in Nyurba situated in the Vilyui basin 

(63 0 30’N; 119 0 E), it is the northernmost point of East Siberia. 

- 71 - 

5 records of Мyotis ikonnikovi are reported in 3 geographical 

points of Yakutia: one specimen was caught as far as 25 km from 

Tommot (town) in the upstream of the Aldan (59 0 N; 126 0 E); one 

– in the area of the mid-Ungra, the upper part of the Aldan basin 

(57 0 30’ N; 124 0 30’ E), three – during wintering in the galleries of 

the gypsum pit, respectively. 

All Chiroptera species in Yakutia are found during their wintering in 

the galleries of the gypsum pit. Experimental research of the 

hibernation course in Eptesicus nilssoni, Plecotus auritus and 

Мyotis daubentoni showed that Hiroptera survival in the frosty 

period without food is maintained due to long-time periods of 

hypothermia (97-98% of budget time) and a short-time (1-2%) stay 

in normothermia. 

Information on Mallophaga fauna of Passeriformes in Yakutia 

Olga N. Stepanova and Nickolay I. Germogenov 



499 individuals from 36 passerine species (28 migratory-breeding 

and 8 resident) were examined, of them we found 3383 

Mallophaga specimens (34 species) on 323 birds (23). Invasion 

extensiveness is 64.7%, abundance index - 6.8 specimens. 

Parasites were absent on Brown shrike (n=3), Northern skylark 

(n=3), Yellow wagtail (n=5) и Water pipit (n=5), Olive-backed pipit 

(n=1), Red-winged thrush (n=1), Siberian rubythroat (n=1), Redflanked 

bluetail (n=11), Arctic warbler (n=2), Pallas`s reed bunting 

(n=8), Gray bullfinch (n=1), Gray-crowned rosy (n=4), House 

sparrow (n=8). 

The greatest Mallophaga diversity is recorded on large resident 

wintering species - Siberian jay, Northern raven, Carrion crow, 

Spotted nutcracker and small migratory – Dusky Thrush, Common 

redpoll. Mallophaga fauna complex comprises 6 genera, 3 families 

- Menoponidae and Ricinidae from Amblycera suborder, 

Philopteridae family from Ischnocera suborder. 12 Mallophaga 

species are new for science, not described earlier. Members of 

Amblycera are dominant by family number while Ischnocera by the 

species number. 

8 species of 2 genera represent Mallophaga of Menoponidae 

family parasitizing on different hosts (Genus Menacanthus is 

mostly rich in species), 1 genus and 4 species of Ricinidae family 

are characteristic parasites of buntings and thrushes. 

Philopteridae family represented by 3 genera is distinguished with 

great species diversity (22). Parasites of this family occur on many 

hosts. Philopterus genus is abundant in species number both in 

Philopteridae family and in other families. 

The Breeding Biology and the Ecology of Grebes 

(Podicepiformes) on the Campenesti ponds (Romania) 

Alexandru Nicolae Stermin, Alin David and Ioan Coroiu 

Babes-Bolyai University, Faculty of Biology and Geology, 

Department of Biology and Ecology, Cluj-Napoca, Romania 

The focus of this study were three protected species of 

Podicepiformes, Podiceps cristatus, Podiceps grisegena and 

Tachybaptus ruficollis, species that have never been studied 

before in Romania. Observations were made between 2005 - 

2007 on the Campenesti Ponds, located in the Feiurdenilor valley, 

an area under intense human pressure. Among the studied 

aspects of the species were the annual passage, the 

distribution, habitat use and the breeding biology, taking into 

account the breeding success and the mortality, with the factors 

influencing it. 

The results show a direct dependence on the total surface and 

density of vegetation, and less so on the depth of the water. 

Correlations were found on the nest-building strategies, the density 

of vegetation and the water depth. The connection between the 

temperature and the effective of the population during the annual 

passage was also revealed, in the spring and autumn. 

On the Campenesti ponds, the breeding success decreases with 

the increased vegetation exploitation, with the increase in the 

density of submerse vegetation, and with the decrease of water 

depth in the ponds.


Integrating ethological and morphological features, ecological 

requirements and biodiversity across taxonomical groups to 

found water quality bioindicators 

Antonio Torralba-Burrial and Francisco J. Ocharan 

Dpto. Biología de Organismos y Sistemas, Universidad de Oviedo, 

E-33071 Oviedo, Spain 

Rivers play a fundamental role in the global hydrological cycle, and 

provide important services to humans and biodiversity. The search 

of simple, easy to use and cheap rivers ecological status 

bioindicators can contribute to their biodiversity conservation. Here, 

dragonflies which presence was related with a high ecological 

status rives were found. In the fluvial network of Aragon (NE Spain) 

108 reaches were sampled. Reach ecological status was assigned 

using IBMWP score (summatory of benthic macroinvertebrate 

communities’ taxa richness weight to taxa pollution sensibility). 

Using multivariate methods (IndVal, SIMPER) dragonfly species 

related with high ecological status reaches were identified: 

Cordulegaster boltonii (Donovan, 1807), Boyeria irene 

(Fonscolombe, 1838) and Onychogomphus uncatus (Charpentier, 

1840). Definitive choice of C. boltonii and B. irene was based on 

their morphological features (size and colour patterns), behaviour 

(adults patrolling river reaches persistently), distribution (wide in 

Aragon) and low taxonomical diversity of genera in the area (only 

one species of Boyeria in Continental Europe; two of 

Cordulegaster in Iberian Peninsula, but Cordulegaster bidentata 

with a very narrow distribution and habitat). These species can be 

identified easily by trained voluntaries in almost all of their life cycle 

phases (larvae, exuviae and imagines). Moreover, both species 

shown indicator values to class I and II of ecological status 

(allowed by European Water Framework Directive), reinforced their 

bioindicator characteristic. 

Coexistence of Aphidius species: Does interspecific 

competition structure the parasitoid community? 

Joan van Baaren 1 , Cécile Le Lann 1 , P. Le Guigo 2 , L. Krespi 2 , 

Yannick Outreman 3 , Jean-Sébastien Pierre 1 and Jacques van 

Alphen 1 

1 UMR CNRS ECOBIO, University of Rennes 1, France; 

2 Ecobiologie des Insectes Parasitoïdes, University of Rennes 1, 

France ; 3 UMR BIO3P, Agrocampus Rennes, France 

Three species of Aphidius parasitoids exploit the wheat aphid 

Sitobion avenae in western Europe. Aphidius rhopalosiphi is a 

habitat specialist and the only species present in wheat fields in 

winter. It is the commonest species throughout spring. Two 

generalists, A. avenae and A. ervi, invade wheat fields by the end 

of spring. Wheat aphids defence themselves actively when 

attacked by parasitoids and emit alarm pheromone when attacked. 

Behavioural analyses of interspecific host discrimination, 

competition, patch exploitation were realized. A. rhopalosiphi is 

sensitive to host defensive behaviour, and exploits host patches 

only partially. Aphidius avenae displays a particular behaviour, with 

wing fluttering before attacking a host, which results in suppression 

of defensive behaviour. This allows them to exploit patches with 

alerted aphids abandoned by A. rhopalosiphi. Aphidius ervi has a 

fecundity twice that of A. rhopalosiphi, is less sensitive to the host 

defence behaviour and is a better competitor than A. rhopalosiphi 

in multi-parasitized hosts. Interactions between A. avenae and A. 

ervi are limited due to the low densities of these species. When 

females of the three species encounter each other in a host patch 

they display no aggressive behaviour to each other. 

We showed that the three species can coexist with little overlap in 

host use, because they have different phenologies and densities, 

different host exploitation strategies and different life histories. 

- 72 - 

Behavioral and endocrine responses to environmental 

challenges in water vole males (Arvicola terrestris) 

Evgenii Zavjalov and Ludila Gerlinskaya 

Institute of Systematics and Ecology of Animals, SB RAS, Frunze 

street 11, Novosibirsk 630091, Russia 

Nobody doubts that stress reaction to both non-social and social 

stimuli plays a key role in the population responses to 

environmental challenges. We studied the multi-annual changes of 

food conditions, demographic and territorial structures, endocrine 

status and immunocompetence of water vole males in local 

population during 6 reproductive seasons. Multi-annual variations 

of food conditions and demography let us to separate these 6 

years on 3 time periods. Two years with minimum of available food 

were combined in food shortage (FS) period. Shortage of 

reproductive resources (SRR) embraced 2 years, when percent of 

females in studied local populations dropped below than 20%. 

Resources abundant (RA) period combined the 2 years when food 

and reproductive resources were higher than multi-annual 

averages. 

At both periods of resource shortage the non-equal space 

distributions of animals was observed in the local population. At 

this time overlapping of individual home ranges increased in 

comparison with RA. But the average home ranges size did not 

change between selected periods. Remarkably, maximum of skin 

wounds as index of aggression was found only in male trapped at 

SRR. The fecal testosterone also was highest in this time. Highest 

aggression in SRR period was coincided with minimal marking of 

animals that was estimated by numbers of latrine. Fecal 

corticosterone increased in both FS and SRR compare with RA. 

Antibody titer (IgG) was higher in males of RA in comparison with 

SRR and FS. Thus, ethological and physiological reactions of male 

in local population are determined by environment conditions.


Evaluation of the biological activity of alkaloids of Pergularia 

tomentosa (Asclepiadaceae) on Vth instar nymphs of Locusta 

migratoria: an histopathological approach 

Fatma Acheuk 1 , Bahia Doumandji - Mitiche 2 , Karima Ait kaci 1 and 

Fethia Fazouane 1 

1 

Department of Biology, Faculty of Sciences, University of 

Boumerdes, Algeria. 

2 

Department of agricultural and forest zoology, National 

Agronomic Institute, El-Harrach, Algeria 

The desert locust Schistocerca gregaria and the migratory locust 

Locusta migratoria occupy a particular place among insect pests. 

They constitute a quasi-permanent threat for cultures and pastures, 

and the current methods of control use liquid insecticides whose 

active components belong to different chemical families. These 

preparations are at the same time effective on locusts but also 

harmful for other animal species of the biotope, in connection with 

their accumulation in the treated ecosystems. 

The use of extracts of plants found in the biotope of these locusts 

and displaying an insecticidal effect may offer an alternative in the 

field of anti-locust fight. In the present study, we studied the 

toxicological impact of a crude mixture of alkaloids extracted from 

the sheets and stems of Pergularia tomentosa on the histology of 

the digestive tract of fifth instar nymphs of the migratory locust. 

Our results showed that the insecticidal effect of this treatment 

resulted in a disruption of the epithelium of mesenteron and gastric 

cæca, together with a slight disorganization of the musculature. 

These histopathological effects are accompanied by diarrhoea and 

a reduction of food intake. 

Hepatic endogenous defense potential of propolis after 

mercury intoxication 

Monika Bhadauria 1 , Sangeeta Shukla 1 , Ramesh Mathur 1 , Om 

Prakash Agrawal 2 , Sadhana Shrivastava 1 , Sonia Johri 3 , Deepmala 

Joshi 1 , Varsha Singh 1 , Deepak Mittal 1 and Satendra Kumar Nirala 1 

1 

Reproductive Biology and Toxicology Laboratory, 

2 

Insect 

Physiology and Biochemistry, School of Studies in Zoology, Jiwaji 

University, Gwalior-474011, India ; 3 Boston College for 

Professional Studies, Gwalior, India 

Exposure to mercuric chloride (HgCl2; 5 mg/kg body weight; i.p.) 

induced oxidative stress in mice and substantially increased the 

LPO and GSSG level along with corresponding decrease in the 

GSH and various antioxidant enzymes in liver and also increased 

the activities of liver marker enzymes in serum. Therapy with 

propolis extract; a resinous wax-like beehive product (200 mg/kg 

orally, after mercury administration) for 3 days inhibited LPO and 

formation of GSSG level along with increase in liver GSH level. 

Release of serum transaminases, alkaline phosphatase, lactate 

dehydrogenase and γ-glutamyl transpeptidase were significantly 

restored after propolis treatment. The activities of antioxidants 

enzymes i.e., superoxide dismutase, catalase, glutathione-Stransferase 

and glucose-6-phosphate dehydrogenase were also 

concomitantly restored towards normal level after propolis 

administrations. The observations clearly demonstrated that 

propolis treatment augments the antioxidants defense against 

mercury induced toxicity and provides evidence that it has 

therapeutic potential as hepatoprotective agent. 

S19 - Animals ecotoxicology 

- 73 - 

Solving the problem of predation: further insights as to how 

barnacles survive as sessile organisms 

John St J. S. Buckeridge and Jessica Reeves 

Earth & Oceanic Systems Research Centre, RMIT University, 

Melbourne, Australia 

During a routine chemical analysis of the stalked ibliform barnacle 

Chaetolepas calcitergum Buckeridge and Newman 2006, a peak 

conforming to more than 15% bromine was detected. Although the 

bromine ions occur in seawater (up to 65ppm), this level of 

accumulation, in the soft tissue of the barnacle, is extraordinary. 

Organic concentration of bromine compounds is known in a 

number of invertebrates, such as algae and sponges, but this is 

the first record of elevated bromine in barnacles. It is proposed that 

the high accumulation of bromine compound(s) is most likely a 

defense mechanism. The paper concludes with a review of the 

mechanisms deployed by barnacles to repel predators. 

Combining ecophysiology and ecotoxicology: field studies 

and experimental approach using the blue mussel Mytilus 

edulis 

François Leboulenger, Julie Letendre, Fabrice Durand, Jean- 

Michel Danger and Frank Le Foll 

Laboratoire d’Ecotoxicologie – Milieux Aquatiques, EA 3222, 

IFRMP 23,Université du Havre, France 

The coastal zone is characterized by a high variability of abiotic 

parameters linked to the tidal cycle and by the presence of diverse 

chemical compounds in the water. These particular conditions 

represent a multiple stress that resident organisms have to deal 

with. Interactive effects of physiological and toxicological stresses 

were investigated in the blue mussel (M. edulis) by studying a 

series of biomarkers such as the activities of antioxidant enzymes 

and MXR protein, and DNA strand breaks, in gills and digestive 

gland of mussels sampled at high shore (HS) and low shore (LS) 

in a reference site (Yport, France) and a contaminated site (Le 

Havre harbour, France) during the tidal cycle. The results indicate 

that patterns of activities of antioxidant enzymes are affected by 

environmental contamination as well as by ecophysiological 

stresses imposed by tidal conditions, and that the combination of 

the two parameters can induce a specific response of these 

activities. 

Biomarker measurements and differential analysis of 

transcriptomes and proteomes were performed in gills of control or 

exposed mussels which were maintained for 2 weeks in an original 

experimental design where the animals could be either 

permanently immersed or submitted to a regular immersionemersion 

alternation. For each exposure or maintenance 

conditions, a specific pattern of messenger and protein expression 

level was recorded, showing that the expression of many 

functional categories of genes are affected by stress. 

New data on the typology and activities of hemocytes in M. edulis 

will be presented.


In vitro methods for the detection of algal neurotoxins to 

secure shellfish and fish consumption 

Aurélie Ledreux 1,2 , Sophie Krys 2 and Cécile Bernard 1 

1 Muséum National d’Histoire Naturelle, USM 505/EA 4105 

Ecosystèmes et interactions toxiques, 57 rue Cuvier, CP 39, 


2 Agence Française de Sécurité Sanitaire des Aliments, Unité 

Toxines, Polluants Organiques et Pesticides, 23 avenue du 

Général de Gaulle, 94706 Maisons-Alfort cedex, France 

The global occurrence of toxic algal blooms has increased in 

frequency and intensity over the past 20 years, resulting in an 

increase in the number of shellfish production area affected by 

phycotoxins (Hallegraeff, 2003). 

To keep shellfish and fish consumption secure, the risk 

management associated to phycotoxin contamination in shellfish 

and fish is based on monitoring of toxins in seafood destined to 

human consumption (Yasumoto, 1995). The available chemical 

methods, while being efficient for the detection of identified toxins 

are inadequate for the detection of emerging toxins or unknown 

analogues. As a consequence, detection of algal toxins is still 

based on mouse bioassays in spite of their poor sensibility, their 

lack of specificity and the ethical issue. 

Alternative assays are required which are reliable, sensitive and 

able to detect toxic agents depending on the molecular mechanism 

by which they affect biological systems. In vitro methods are 

attractive since they are inexpensive and allow a rapid screening 

of a large number of samples. 

The aim of this study was to determine if the specific detection of 

three neurotoxins families (saxitoxins, brevetoxins and palytoxin 

and analogues), acting on two different cellular targets can be 

performed by using a single neuroblastoma cell-line (Neuro-2a). 

To that end, pure toxins have been tested in order to evaluate the 

sensibility of the Neuro-2a cell based assay. The specificity of the 

assay for each type of toxins was achieved by the use of two 

antagonists: ouabain and veratridine. The matrix effect was 

assessed by testing algal extracts and mussel extracts. 

Bioaccumulation of heavy metals in the tissues of two 

flatfishes (Platichtys flesus and Solea vulgaris) from the outer 

coast and Ría of Vigo, NE Atlantic (Spain) 

Lazhar M’Hadhbi 1,2 , Dragos Zaharescu 2 , Tahar Gharred 3 , Antonio 

Palanca 2 and Moncef Boumaïza 1 

1 Hydrobiology Unit, Enironmental Biomonitoring Laboratory, 

Faculty of Sciences, Bizerte, Zarzouna 7021, Tunisia 

2 Animal Anatomy Laboratory, Vigo University, 36310 Vigo, 

Spain 

3 Marine Biotechnological Institute of Monastir,Tunisia 

Concentrations of ten heavy metals (Cd, Pb, Hg, Cu, Zn, As, Se, 

Ag, Mn, Sn) in liver, muscle and kidney of European flounder 

(Platichtys flesus) and the sole (Solea vulgaris) from Ria of Vigo 

and its coastal zone (Spain) were measured from April 2006 to 

February 2007 and the relationships between fish size (length and 

weight), age, gender, season and metal concentrations in the 

tissues was investigated. 

The average metal concentrations in different organs varied in the 

following ranges: Cd (0.01- 2.08), Pb (0.02- 2.91), Hg (0.015- 2.13), 

Cu (2.5- 46.71), Zn (12.7- 996.7), As (1.2- 151.9), Se (1.56- 34.02), 

Ag (0.01- 5.81), Mn (0.27- 57.26), Sn (0.01 to 5.07) µg g -1 dry 

weight. Fish kidney and liver registered highes levels of Cd, Cu, Pb 

and Zn, whereas the muscles registred highest Hg concentrations. 

Platichtys flesus tended to accumulate more quantities than Solea 

vulgaris. A comparison of metals concentration in fishes from both 

sites shown higher bioaccumulation in those exemplars from the 

Ria of Vigo. This means a tendency of higher pollution levels in the 

estuarine environment. Seasonal variability generally indicated 

increased levels of metals bioaccumulation during winter, the 

period of heavy rains and high flows from the surrounding Vigo 

cityscape. 

Although, generally, metals levels in the tissues of fish in the area 

are not harmful for humans, measures shall be taken to lower 

heavy metals inputs into Ria of Vigo. 

- 74 - 

Global quantitative analysis of protein phosphorylation status 

in liver of fish exposed to the Cyanotoxin Microcystin 

Mélodie Malécot 1 , Karim Mezhoud 1 , Danièle Praseuth 2,3 , Arul 

Marie 4 , Simone Puiseux-Dao 1 and Marc Edery 1 

1 USM 505/EA 4105 Écosystèmes et interactions toxiques, 

Département de « Régulations, développement et diversité 

moléculaire », Muséum national d’Histoire naturelle, 12 rue Buffon, 

F-75231 Paris Cedex 05, France ; 2 INSERM, U 565 and 3 MNHN, 

USM 503 Laboratoire des Régulations et dynamique des génomes, 


moléculaire », CNRS, UMR 5153, Acides nucléiques: dynamique, 

ciblage et fonctions biologiques, Muséum national d’Histoire 

naturelle, 57 rue Cuvier, CP26, F-75231 Paris Cedex 05, France ; 

4 Plateforme de spectrométrie de masse et de protéomique, 


moléculaire », Muséum National d’Histoire Naturelle, 63 rue Buffon, 

F-75231 Paris Cedex 05, France 

Microcystins (MCs) are hepatotoxins inhibiting protein 

phosphatases PP1 and PP2A. These peptides produced by some 

freshwater cyanobacteria are getting attention due to their acute 

toxicity and tumor-promoting activity. To date, the detailed 

mechanisms underlying their toxicity are unknown. MC-leucinearginine 

(MC-LR) is the most toxic and the most commonly 

encountered MC variant in aquatic environment. Toxicological 

investigations on the liver of the aquatic model, the medaka fish, 

have been performed with differential proteome analyses of MC- 

LR-treated (by 2 exposure routes; water or food contamination) 

and untreated medaka fish in order to investigate the 

mechanisms of establishment of early responses to the toxin. 

Cytosolic, membrane and organelle proteins from livers were 

resolved by 2D electrophoresis and detected using stains specific 

for phosphoproteins and for whole protein content. Overall, more 

than 100 spots were found to vary significantly on the proteomic 

2D maps or on the phosphoproteomic 2D maps. Of these, 32 

proteins could be identified by mass spectrometry. Among them, 

phenylalanine hydroxylase, keratin 18 (type I) and grp78 showed 

variations in phoshoryl content that could be directly associated 

with inhibition of PP activity. The other identified proteins 

exhibited variations in their expression level. The identified 

proteins appear to be involved in cytoskeleton assembly, cell 

signalling, oxidative stress and apoptosis. The reported 

methodology should be widely used to a number of tissues and 

organisms, thus helping in the search for biomarkers of MC-LR 

contamination. 

Study of the insecticidal activity of alkaloids of the 

hairybroom (Cytisus triflorus L’Hér.) and of the bacterium 

Bacillus thuringiensis against the desert locust Schistocerca 

gregaria 

Hakima Mohand Kaci 1 , Karima Ait-Kaci 1 , Bahia Doumandji- 

Mitiche 2 and Fethia Fazouane 1 

1 Faculty of Sciences, Department of Biology, University of 

Boumerdes, Algeria; 2 El Harrach National Agronomy Institute, 

Alger, Algeria 

In many regions of Africa and Asia (including Algeria), in addition 

to abiotic factors such as climate, food security relies mainly on 

crop protection. Crops are under attack from locusts, e.g. the 

desert locust S. gregaria. Among existing forms of control, are 

those involving entomopathogenous microorganisms, e.g. the 

bacterium B. thuringiensis, or acridicide natural products, e.g. 

those extracted from the shrub C. triflorus. In the present work, we 

compare the effects of B. thuringiensis and alkaloids of C. triflorus 

on digestive tract histology of the of adult locusts and on three 

physiological parameters: heart rate, respiratory rate and 

haemocyte number. The digestive tract anatomy of control and 

treated animals appeared identical. Histological examination of the 

three parts of the digestive tract (stomodeum, mesenteron and 

proctodeum) of locusts treated with alkaloids and by the bacterium 

showed that histological changes occurred in the three parts.


In animals treated with alkaloids, we noted a tear in internal 

longitudinal muscles and external circular muscles in the 

stomodeum and proctodeum and destruction of some epithelial 

cells in the medial intestine. We also observe a tear in external 

circular muscles and destruction of some epithelial cells in the 

stomodeum of bacterial-treated individuals. The two agents show a 

clear decrease in the opening number of metathoracic stigma and 

heartbeat; the effect of C. triflorus alkaloids on respiratory rate was 

greater than that of B. thuringiensis, although the heartbeat rate 

was almost the same for both test agents. However, the bacterium 

has a greater effect on haemolymph, since the treatment causes a 

considerable decrease in haemocyte number. In conclusion, it 

appears that the two control agents show significant effects on the 

physiology of S. gregaria. It is important, therefore, to extend 

studies on these two control agents for use in integrated control. 

Effects of water-borne copper on the isozymes and on the 

ultrastructure of gills and hepatopancreas in Macrobrachium 

rosenbergii 

Na Li 1,3 , Qiang Ma 1 , Jian Yang 2 , Yunlong Zhao 1 and Jean-Claude 

Brochon 3 

1 School of Life Science, East China Normal University, Shanghai 

200062, China; 2 Key Laboratory of Ecological Environment and 

Resources of Inland Fisheries, Freshwater Fisheries Research 

Center, Chinese Academy of Fishery Sciences, Wuxi 214081, 

China ; 3 CNRS, UMR8113, Laboratoire de Biotechnologies et 

Pharmacologie Génétique Appliquée, Ecole Normale Supérieure 

Cachan, 61 avenue du Président Wilson, 94235 Cachan cedex, 

France 

Present study is focused on elucidating toxic effects of water-born 

copper on giant freshwater prawns Macrobrachium rosenbergii. 

After 7 days exposure to copper (Cu 2+ ) concentrations ranging 

from 0.01 mg/l to 0.5 mg/l, three isozymes, including malate 

dehydrogenase, alkaline phosphatase, and esterase, were 

analyzed and compared by use of polyacrylamide electrophoresis 

(PAGE) and biochemical staining. Results indicated that 

electrophoretic patterns of isozymes showed a copper 

concentration-related difference. Low doses of copper stimulated a 

strong expression for three isozymes. Electrophoretic patterns of 

malate dehydrogenase and alkaline phosphatase gradually 

became weaker or even lost as the copper level increase; on the 

contrary, esterase patterns exhibited an increased molecular 

heterogeneity in higher copper concentrations. Meanwhile, 

transmission electron microscope (TEM) was adopted to study the 

ultrastructure differences of gills and hepatopancreas in M. 

rosenbergii, results showed a significant structural damage as the 

increase of copper level: compared with the control group, 

basement membranes and mitochondira in gills were seriously 

damaged. Cuticle electron density distributed inhomogeneously; 

infolded basement membrane, circularized nucleus, disintegrated 

nuclear membrane; decreased mitochondria number and size 

were observed in gills. Similarly, flowing out of karyoplasms, partly 

falling microvilli, decreased mitochondrion, partly disappeared 

mitochondrial cristae, thinned matrix were observed in 

hepatopancreas. Present studies indicated that, exposure to 

elevated copper levels might damage the ultrastructure of gills and 

hepatopancreas in M. rosenbergii and further weaken their normal 

physical activities; isozymes were quite sensitive to the 

environment stress and changes of isozymes electrophoretic 

patterns could be effective biomarkers to environment 

contamination. 

Combined effects of gallic acid and propolis on berylliuminduced 

hepatorenal toxicity 

Satendra Kumar Nirala 1,4 , Pei Qiang Li 2 , Monika Bhadauria 3 and 

Guang Qin Guo 1 

1 School of Studies in Zoology, Jiwaji University, Gwalior-474011, 

India ; 2 Institute of Genetics, School of Basic Medical Sciences, 

Lanzhou University, Lanzhou, 730000, PR China ; 3 National 

Laboratory of Applied Organic Chemistry, Lanzhou University, 

Lanzhou, 730000, PR China ; 4 Institute of Cell Biology, School of 

Life Sciences, Lanzhou University, Lanzhou, 730000, PR China 

- 75 - 

The combined effect of gallic acid (3,4,5-trihydroxy benzoic acid; 

GA; 50 mg/kg, i.p.) and propolis (200 mg/kg, p.o.) was evaluated 

against beryllium induced biochemical and morphological 

alterations of liver and kidney. Female albino rats were exposed to 

beryllium nitrate (1mg/kg, ip) daily for 28 days followed by 

treatment of above mentioned therapeutic agents either 

individually or in combination for 5 consecutive days. Exposure to 

beryllium increased its concentration in serum, liver and kidney 

and caused significant alterations in cytochrome P450 enzymes, 

microsomal lipid peroxidation and protein contents. Beryllium 

administration significantly altered the aspartate 

aminotransaminase, alanine aminotransaminase, lactate 

dehydrogenase, γ-glutamyl transpeptidase, bilirubin, creatinine 

and urea in serum; activity of acid phosphatase, alkaline 

phosphatase, adenosine triphosphatase, glucose-6-phophatase 

and succinic dehydrogenase, triglycerides, cholesterol, protein 

contents, glycogen contents, lipid peroxidation and glutathione 

level in liver and kidney. Beryllium exposure induced severe 

alterations in hepatorenal morphology proving its toxic 

consequences at cellular level. Individual administration of GA and 

propolis recovered some of the studied parameters moderately 

towards control. Interestingly, GA in conjunction with propolis 

reversed the alterations of all the variables more towards control 

concluding beneficial effects of combined therapy over mono 

therapy in alleviation of beryllium induced systemic toxicity. 

Biomarkers in Crassostrea gigas larvae in a transplant 

experiment in Archachon harbour (France) 

Françoise Quiniou 1 , Gautier Damiens 2 , Xavier Caisey 1 , Catherine 

Mouneyrac 3 , Hélène Budzinski 4 and Michèle Roméo 2 

1 Département Biogéochimie et Ecotoxicologie IFREMER - Centre 

de Brest, B.P. 70, 29280 Plouzané, France ; 2 EA ECOMERS 

Ecosystèmes Côtiers Marins et Réponses aux Stress, Faculté des 

Sciences, BP 71, 06108 Nice Cedex 2, France ; 3 Centre d’Etude 

et de Recherche sur les écosystèmes aquatiques, IBEA/UCO, BP 

10808, 49008 Angers Cedex 01, France and Service 

d’Ecotoxicologie, SMAB-ISOMER, Faculté des Sciences, BP 

92208, 44322 Nantes Cedex 3, France ; 4 LPTC Université de 

Bordeaux I, 3405 Talence, France 

Early embryo-larval stages of bivalve molluscs are frequently used 

in marine exotoxicology; however laboratory studies cannot 

accurately simulate natural conditions. Field experiments can 

eliminate artefacts resulting from artificial laboratory conditions 

affecting the real toxicity and bioavailability of contaminants. Two 

field experiments were conducted (July 2002 and June 2004) in 

which fertilized gametes from Crassostrea gigas were placed into 

a special container (device patented by Ifremer). Embryos kept in 

the containers were immersed into two sites of Arcachon harbour 

(A: entrance of the harbour and P) for 48 h (field temperature 22 ± 

1°C, salinity 32.0 ± 0.5). The containers were ballasted so as to 

keep them at a depth of 1 meter under the surface. Larvae were 

collected and biomarkers (acetylcholinesterase AChE, catalase 

CAT, glutathione transferase GST activities, Thiobarbituric acid 

reactive substances TBARS, metallothionein concentrations MT) 

determined in the samples as well as metal and PAH (polycyclic 

aromatic hydrocarbons) concentrations in larvae and sediments 

collected under the containers. The results show that GST activity 

and TBARS levels were significantly higher at P than at A. MT and 

copper concentrations in larvae tended to increase from A to P 

whereas AChE activity showed a trend to be lower at P. Organics 

in larvae showed a tendency to be higher at A than P. Pollutant 

concentrations in sediments showed the same trends as larvae. At 

P, larvae seemed to be submitted to an oxidative stress inducing 

lipid peroxidation of the membranes (shown by TBARS and GST 

activity), this stress might be caused by copper present in the 

surrounding environment. Larvae, caged in A (entrance of the 

harbour) where water masses flow, seemed to be affected by low 

molecular weight PAHs such as naphtalene which are soluble 

compounds (contrary to high weight compounds). Moreover, the 

percentage of abnormal larvae was higher at A (40 %) than P (60 

%) suggesting that A was more exposed to embryotoxic pollutants 

than P. Nevertheless both sites may be considered as submitted 

to anthropogenic influence. As conclusion, transplantation of 

larvae together with chemical and biomarkers measurements 

constitute an “active biomonitoring” providing early warning signal 

of pollution.


Honeybees as bioindicators in urban areas (Seine-Saint-Denis, 

France) 

Agnès Rortais, Yves Loublier and Gérard Arnold 

Laboratoire Populations, Génétique, Evolution CNRS, UPR 9034, 

Avenue de la Terrasse, 91198 Gif-sur-Yvette – France 

Foraging activity in honeybees is an important component of the 

colony’s life. When visiting plants, honeybees bring back products 

(nectar, pollen) to the colonies that mirror the diversity and quality 

of the environment. In addition, honeybees being very sensitive to 

environmental pollution, any behavioural changes among foragers 

are an indicator of an environmental stress. Based on these 

features, the present study describes tested methods for the use of 

honeybees as bioindicators, in particular in urban areas. 

In the region of Ile-de-France, three sites showing different types 

of anthropogenic disturbances (urban, agricultural and seminatural) 

were selected. In these sites, five hives were monitored in 

2007 and 2008 during the flowering period (April to September). In 

particular, pollen traps were set at two hives and bee traps, bee 

counters, cameras and scales on the three other hives. Detail 

analyses were conducted on pollen identification, forager’s activity 

and behaviour, and colony’s development 

Preliminary data are described for the urban site based in the 

Seine-Saint-Denis department which is recognized for its activities 

in the field of urban biodiversity conservation (ODBU: Observatoire 

Départemental de la Biodiversité Urbaine – Departmental 

Observatory of urban biodiversity). 

One of the main striking results of this study is the great taxonomic 

richness found at this site and the presence of exotic species. 

These results highlight the importance of conserving urban areas 

as reservoirs of nectariferous and melliferous resources for 

pollinators. Further studies need to identify the relative importance 

of generalist (like honeybees) versus specialist pollinators in such 

a floral environment. 

Uncertainty and ignorance in assessing environmental effects 

of toxic chemicals: the example of endocrine disruption 

Helmut Segner 

Centre for Fish and Wildlife Health, University of Bern, Switzerland 

Ecotoxicological risk assessment aims to estimate at what 

concentrations chemicals may occur in the environment and at 

what concentrations they are likely to cause adverse effects in 

exposed biota and ecosystems. The toxicological data used as 

basis in the risk assessment process are derived mainly from 

laboratory tests relying on few “model” species and on apical, nonmechanistic 

endpoints. Further, the focus of these tests is on acute 

rather than on chronic effects. 

The extrapolation from such reductionistic approaches to 

environmental scenarios necessarily involves uncertainty. 

However, the extrapolation additionally suffers from ignorance on 

the biological functions and processes that are at risk by the 

exposure to toxic chemicals. As an example to illustrate the 

inherent problems in assessing the impact of chemicals on biota, 

the presentation will discuss the case of endocrine disruption, i.e. 

the disturbance of hormone systems by environmental compounds. 

- 76 - 

Cellular damage induced by organic mercury and its 

prevention by combination therapy 

Varsha Singh 1 , Sadhana Shrivastava 1 , Mohammed Abdullah 2 and 

Sangeeta Shukla 1 

1 Reproductive Biology and Toxicology Laboratory, School of 

Studies in Zoology, Jiwaji University, Gwalior-474011, India 

2 Trace Element - Institute for UNESCO, 1 place de l'Ecole, BP 

7021, 69342 Lyon Cedex 07, France 

Heavy metals such as cadmium, mercury (Hg), lead etc. are major 

environmental and occupational hazards. These non-essential 

elements are toxic even at very low doses and non-biodegradable 

with a very long biological half-life. Mercury is one of these heavy 

metals, which leads to several health hazards in human and 

animals. In this study, we addressed the putative uptake pathways 

involved along the nephrons, the mechanisms of intracellular 

sequestration and detoxification of mercury. We also tackled the 

question of the possible therapeutic means to decrease its toxic 

effects. For this purpose, albino rats were intoxicated with mercury 

(Dimethyl mercury; 10 mg/kg, p.o., once only). The degree of 

kidney damage was evaluated biochemically and 

histopathologically. Quantity of SALP, GGT, urea, uric acid, BUN 

and creatinine were increased in serum after Hg administration. 

Renal lipid peroxidation was raised significantly, whereas GSH 

contents and the enzymatic activities of GSH cycle were found to 

be declined. Significant accumulation of Hg was also noticed in 

kidney. Three days curative treatment of glutathione (GSH; 0.30 

mM/kg, i.p.) and its combination with lipoic acid (LA; 100 mg/kg, 

p.o) and magnesium (Mg; 10 mg/kg, p.o.) recovered in all the 

biochemical estimations. Protective activity of GSH + Mg was 

statically compared with GSH + LA and is better in preventing 

oxidative stress as confirmed by recouped endoplasmic reticulum 

and mitochondrial assembly in ultra structural study. Distribution 

study of mercury demonstrated that GSH + Mg exert more 

pronounced effects (P < 0.001) rather than GSH + LA and per se 

groups.


S20 - Marine organisms and symbiotic systems in extreme environments 

Investigation of the intertidal Macrobenthic communities in 

the Yangtze estuary, China 

Chuanguang An and Yunlong Zhao 

School of life science, East China Normal University, Shanghai, 

China 

The Yangtze River, is a river of medium sediment concentration. 

Its length, runoff volume and sediment transporting capacity are 

ranked respectively as third, fourth and fifth in the world. The 

highly productive tidal flat of the Yangtze estuary serves as habitat 

and breeding ground for many commercially valuable fishes and 

invertebrates, and migration station for shorebirds. In order to 

obtain integrative inshore data, the distribution patterns and faunal 

composition of intertidal macrobenthic assemblages were studied 

in the Yangtze estuary (30°19′-31°53′N, 121°18′-122°00′E) from 

June 2006 to October 2007 as a part of a national project. 

According to longitudinal and vertical gradients of the estuary, 116 

sites were sampled in the spring and autumn respectively for 

qualitative and quantitative investigation. The macrobenthic 

species distribution and community structure are related to salinity, 

depth, sediment characteristics and artificial activities. The 

analyses clearly reveal distinct gradients in diversity, abundance, 

and biomass along the vertical and longitudinal gradients. 

Compared with the previous data, the species composition and 

distribution of macrobenthos of Yangtze estuary have been 

remodeled, due to effects of artificial activities and other factors. 

Macrobenthos assemble mainly in a less than 100 m wide narrow 

strip between the sea dam and seawall, while the sedentary 

benthos, which seldom or never existed before in this area, 

increased in number and became a boom in seawall. In conclusion, 

despite the effects of a boom for some species, the resources of 

benthic macrofauna is for the most part reduced. 

Vestimentiferan tubeworms from vents and seeps: how 

different are they? 

Ann C. Andersen 1,2 

1 Université Pierre et Marie Curie - Paris 6, Laboratoire Adaptation 

et Diversité en Milieu Marin, 2 CNRS, UMR 7144, Equipe 

Ecophysiologie: Adaptation et Evolution Moléculaires, Station 

Biologique, Place Georges Teissier, BP 74, 29682 Roscoff Cedex, 

France 

Vestimentiferan tubeworms are polychaete annelids belonging to 

the Siboglinidae that have neither mouth nor gut, but live in 

symbiosis with chemosynthetic bacteria stored intracellularly in 

their trophosome. This symbiotic relationship implies that the worm 

gets oxygen, but also carbon dioxide and sulphides from its 

surroundings, and brings them through the blood circulation to the 

bacteria that in turn use these inorganic elements to build organic 

compounds for the benefit of their host. The role of the gill in 

assimilating these inorganic molecules depends on the particular 

species habitat: vents or seeps. Internal transport of these 

molecules relies heavily on the extracellular hemoglobins 

dissolved in their blood and coelomic fluid. Till now about 16 

vestimentiferan species have been described worlwide in either 

hydrothermal vents, cold seeps or even both. Riftia pachyptila, 

Jones 1981, lives in the hydrothermal vents from the East Pacific 

Ridge, as also Ridgeia piscesae, (Jones 1985) Southward 1995, 

the latter having two different morphotypes: one close to 

hydrothermal chimneys, and the other in cracks of basaltic fields. 

Escarpia southwardae, Andersen 2004, lives in the East Atlantic 

cold seeps and Lamellibrachia n. sp. in Mediterranean carbonate 

crusts. We compare these four species regarding their morphology, 

molecular phylogeny, gill functional anatomy and hemoglobins 

structure and function. This synthesis aims to enlight, whether 

there could be possible links between the 

morphological/physiological characteristics and the habitats 

conditions of these tubeworms. 

- 77 - 

Multiple new nuclear markers for the exploration of the 

adaptative speciations in arctic Nototheniids 

Mohamed Berkani 1 , Agnès Dettai 1 , Arnaud Couloux 2 , Corinne 

Cruaud 2 , Anne-Claire Lautredou 1 , Sophie Sanchez 1 and Guillaume 

Lecointre 1 

1 MNHN, UMR7138, Dpt Systématique et Evolution, Paris, France 

2 Génoscope, Centre National de Séquençage, Evry, France 

It has been proposed that some groups of organisms of the 

Antarctic Ocean have diversification patterns characteristic of a 

species flock type of adaptive speciation. Nototheniidae (Teleostei) 

are endemic to the region, and among the most promising 

organisms for this type of study. While recent studies on 

nototheniid fishes have resolved most of the relationships within 

the group, some nodes of the tree are repeatedly unresolved or 

displaying contradictory support depending on the markers used in 

the study. We have explored more than ten markers looking for 

information to solve the basal nototheniid polytomy and the 

interrelationships within the genus Trematomus. Some of these 

markers had been previously used for teleost phylogeny (TMO4c4, 

rhodopsin retrogene, IRBP gene 1), but several new markers have 

been developed for this study and present more variability than the 

ones previously available. These show promise for the resolution 

of the interrelationships of closely related species and 

phylogeographic studies in some other acanthomorph groups, 

where the problem of insufficient variability arises repeatedly in 

nuclear markers. The comparison of the trees resulting from the 

separate analyses of each marker highlights some areas of conflict. 

Overall, the new data clarifies the interrelationships of nototheniid 

fishes, partially solving the basal part of the nototheniid tree and 

allowing to draw some hypotheses on the relationships within the 

Trematomus “bush at the top”. The position of several species 

never included before in a molecular study could also be clarified. 

Comparative study of the lipidic profile of three Atherina 

populations of Tunisia: Atherina boyeri (Bizerta sea), Atherina 

lagunae (Bizerta lagoon) and Atherina sp. (Kerkannah’s 

Islands) 

Nawzet Bouriga 1 , Salah Selmi 2 , Eric Faure 3 and Monia Trabelsi 1 

1 

Unité de Biologie marine. Faculté des Sciences de Tunis, 2092 

Campus universitaire, Tunisia 

2 

Institut National des Sciences et Technologies de la Mer. 

Laboratoire de Biotechnologie Marine, La Goulette 2060, Tunisia 

3 

Laboratoire Systématique Evolutive, case 5, EA 2202 

« Biodiversité », place Victor Hugo, Université de Provence, 13331 

Marseille cedex 3, France 

Atherina are teleostean fish characterised by a capacity to tolerate 

enormous variations in temperature and salinity, which allowed 

them to occupy seas, lagoons and estuaries. In Tunisia, various 

studies based on morphological, morphometric, and genetic 

parameters allowed several local populations to be distinguished. 

In the present work, we examined possible differences or 

variations in the lipidic profiles of Atherina sp. (insular population), 

Atherina lagunae (lagoon population) and Atherina boyeri (marine 

population) as a means of identifying each population and 

investigating the possible existence of new specie that we aim to 

characterize. The total lipid content of all examined specimens was 

nearly the same, i.e. about 6% of the wet weight. Saturated fatty 

acids constitute the majority of the fatty acids pool, reaching 

43.54%, 36.96% and 33.64% in marine, lagoon and insular 

Atherina respectively. The total polyenes content was 27% in 

Atherina boyeri and Atherina sp. In these two populations, 

eicosapentaenoic acid, docosahexaenoic acid and linoleic acid are 

the predominant fatty acids, while n-6 polyunsaturated fatty acids 

of Atherina sp. were significantly higher (14.44%) than in Atherina 

boyeri and Atherina lagunae (5.97% and 6.78% respectively). The 

index PUFAn-3/PUFAn-6 shows a significant level, indicating a 

tendency to accumulate n-3 fatty acids in Atherina boyeri and 

Atherina lagunae, and n-6 fatty acids in Atherina sp.


How to quantify endosymbionts in the gills of the 

hydrothermal vent mussel Bathymodiolus.azoricus ? 3D-FISH 

versus qPCR 

Isabelle Boutet 1 , Arnaud Tanguy 1 , François H. Lallier 1 , Sébastien 

Halary 2 , Sébastien Duperron 2 and Françoise Gaill 2 

1 UPMC-Paris 6 & CNRS, UMR 7144 Adaptation et Diversité en 

Milieu Marin, Station Biologique de Roscoff, 29680 Roscoff, 

France 

2 UPMC-Paris 6 & CNRS, UMR 7138 Systématique, Adaptation et 

Evolution, 7 quai St Bernard, 75005 Paris, France 

The mussel Bathymodiolus azoricus forms dense colonies around 

the deep sea hydrothermal vents of the mid-Atlantic ridge. They 

derive most if not all of their metabolic needs from the two types of 

endosymbiotic chemoautotrophic bacteria they harbour in their gills. 

The two types of gamma-proteobacteria, a methanotroph and a 

thiotroph, are found in variable amounts in the bacteriocytes, 

apparently as a function of methane and sulfide availability in the 

mussel environment. Here we compare two methods aiming at 

quantifying each type of bacteria. The first one (3D-FISH) 

measures volumes occupied by each type of symbiont in 

bacteriocyte sections from a vent mussel gill filament using 

fluorescence in situ hybridization with 16S rRNA-based specific 

probes coupled to three dimentional microscopy and image 

analysis carried out by a dedicated software. The other method 

(qPCR) uses the same probes and some others targeting specific 

metabolic genes to measure the relative expression of these 

genes in gill extracts. Qualitatively, the two methods give 

congruent results and confirm the impact of local environmental 

parameters on symbiont abundances. 

A novel view on relationships between Lucinidae (Mollusca: 

Bivalvia) and intracellular sulfur-oxidizing bacteria 

Terry Brissac 1 , Olivier Gros 2 , Audrey Caro 3 and Hervé Merçot 1 

1 UMR 7138 CNRS UPMC MNHN IRD “Systématique, Adaptation, 

Evolution”, Equipe: Génétique & Evolution, UPMC, 7 quai St 

Bernard, 75252 Paris cedex 05, France ; 2 UMR 7138 CNRS 

UPMC MNHN IRD “Systématique, Adaptation, Evolution”, Equipe: 

Symbiose, UAG, UFR SEN, Dpt de Biologie, BP592, 97159 

Pointe-à-Pitre cedex, Guadeloupe, France ; 3 UMR-CNRS 5119 

Laboratoire Ecosystèmes Lagunaires, CC 93, Université de 

Montpellier II, 34095 Montpellier cedex 5, France 

Associations between marine invertebrates and chemoautotroph 

bacteria constitute a large field for the study of symbiotic 

associations. In these interactions, transmission of the symbiont 

must represent the cornerstone to permit the persistence of the 

association through generations. Within Bivalvia, in some cases 

like Solemyidae or Vesicomyidae, the vertical transmission mode 

of the symbiont occurs without any ambiguity. However, within 

Lucinidae, the transmission mode is described in the literature as 

environmental, symbionts being acquired by the new host 

generations from the environment. Nevertheless, the precise origin 

of the symbiont which infects lucinids is still unknown, and we can 

pose two hypotheses. Symbionts’ origin could be: (i) bacteria 

released after multiplication in a bivalve host of the parental 

generation, (i.e. environmental transmission), (ii) free-living 

bacteria which multiply only in the environment; in which case we 

would not have transmission, but merely recruitment of nearby 

bacteria by the Lucinidae. Our observations show that after 

internalization of symbiosis competent bacteria from the 

environment, it appears that symbionts can not divide inside the 

bacteriocytes even if genome replication stays always active (Caro 

et al., 2007). Moreover, it seems that symbionts are not released 

by their host and at last are destroyed via lysosomal degradation 

process (Liberge et al., 2001). According to these observations the 

environmental transmission hypothesis can be rejected, and this 

relationship seems more correspond to a predation than a 

mutualistic relationship. Moreover the association seems to be 

advantageous only for the bivalve and constitutes a dead-end for 

the bacteria. 

- 78 - 

Lucinidae/Sulfur-oxidizing bacteria: Ancestral heritage or 

environment-dependant association? 

Terry Brissac 1 , Olivier Gros 2 and Hervé Merçot 1 

1 

UMR 7138 CNRS UPMC MNHN IRD “Systématique, Adaptation, 

Evolution”, Equipe: Génétique & Evolution, UPMC, 7 quai St 

2 

Bernard, 75252 Paris cedex 05, France ; UMR 7138 

“Systématique, Adaptation, Evolution”, Equipe: Symbiose, UAG, 

UFR SEN, Dpt de Biologie, BP592, 97159 Pointe-à-Pitre cedex, 

Guadeloupe, France 

In symbiosis, a question which arises concerns the origin and the 

evolutionary story of symbiotic couples: the two partners could 

have co-evolved or not from an ancestral couple (Distel et al., 

1994). Analysis of diversity and host/symbionts phylogenies 

comparison could give us some evidences to discriminate between 

these two hypotheses. The genetic diversity of gill-endosymbionts 

associated to Lucinidae was studied via sequencing of 16S rRNA. 

In Distel et al. (1988; 1994) a bacterial species is specifically 

associated to each host species. In a second study, 6 host species 

harbor the same bacterial species (Durand & Gros, 1996), results 

which permit us to pose the hypothesis that thes associations are 

not strict and are constituted according to the bacterial species 

present in the environment. The comparison of hosts and 

symbionts phylogenies of other lucinids from Philippines, confirms 

that there is no specificity of association between Lucinidae and 

their symbionts. Considering that 16S rRNA being maybe too less 

discriminative to point a bacterial diversity at an intra-specific level, 

we develop at the laboratory a MLST (Multi Locus Sequence 

Typing) analysis with different markers (dnaE, gyrB, rpoB, ITS, 

aprA) and analyzed species used by Durand & Gros (1996). 

Different bacterial haplotypes were retrieved, which one specific to 

two host species sampled in superficial strata of sediments. This 

specificity is independent from the geographic localization and 

does not constitute a phylotype. So, we sought to know if there 

exists an ecotype i.e. a bacterial strain only localized in superficial 

strata of the sediments. 

Effects of host starvation (Codakia, Bivalve, Lucinidae) on its 

symbiotic population 

Audrey Caro 1 , Marc Bouvy 1 and Olivier Gros 2 

1 UMR-CNRS 5119 Laboratoire Ecosystèmes Lagunaires, CC 93, 

Université. Montpellier II, 34095 Montpellier cedex 5, France 

2 UMR 7138 CNRS UPMC MNHN IRD “Systématique, Adaptation, 

Evolution” Equipe: Symbiose, UAG, UFR SEN, Dpt de Biologie, 

BP592, 97159 Pointe-à-Pitre cedex Guadeloupe 

The association between bivalves and chemoautotrophic 

symbionts allows the host to fix carbon either by autotrophic or 

heterotrophic pathways, namely a mixotrophic diet. Codakia 

orbicularis, a tropical lucinid bivalve, lives in association with 

sulfur-oxidizing bacteria, supporting the autotrophic pathway, 

despite the presence of a reduced digestive tract which sustains 

filter feeding nutrition. To evaluate the effect of host starvation on 

the symbiotic population, long term experiment was designed, 

maintaining freshly collected bivalves in artificial seawater during 

several months. The modification in the symbiotic population 

housed in the gills of the bivalve, namely in bacteriocytes, was 

monitored by transmission electron microscopy (TEM) and 

fluorescence in situ hybridization (FISH). A major consequence of 

host starvation consists in a gradual decrease in the symbiotic 

population. Quantification of the fluorescence FISH signal through 

gill section showed that one third of the initial symbiotic population 

disappears each month of starvation. TEM observations reveal, 

according to the presence of lysosomes along with the 

disappearance of bacteriocytes, that symbionts could be digested 

by the host. The capability of Codakia to survive in starvation 

conditions for long period seems to rely mainly on symbiont 

digestion. These results support evidence that symbiont digestion 

could represent an important part in host nutrition in natural 

conditions.


Integument, moult cycle and bacterial ectosymbionts 

relationships in the deep-sea hydrothermal vent shrimp, 

Rimicaris exoculata : a review 

Philippe Compère 1 , Magali Zbinden 2 , Laure Corbari 1 , Marie-Anne 

Cambon-Bonavita 3 , Gilles Lepoint 1 , Bruce Shillito 2 and Françoise 

Gaill 2 

1 Université de Liège, Dept. Sciences et Gestion de 

l’Environnement, Institut de Chime (B6c), allée de la chimie, 3, B- 

4000 Liège, Belgique 

2 UMR 7138 ‘Systématique, Adaptation et Evolution’, Université 

Pierre et Marie Curie, 7 Quai St Bernard, F-75252 Paris Cedex 05, 

France ; 3 IFREMER, Laboratoire de Microbiologie des Environnements 

Extrêmes, Centre de Brest, BP 70, F-29280 Plouzané, 

France 

The Rimicaris exoculata shrimps are considered as the primary 

consumers dominating the fauna of Mid-Atlantic Ridge (MAR) 

hydrothermal vent ecosystems. These shrimps harbour in their 

dilated gill chambers an important ectosymbiotic community of 

chemoautotrophic bacteria associated with iron oxide deposits. 

Observations in scanning and transmission electron microscopy 

realised over the past 5 years give new insights in the 

establishment and functioning of the supposed ectosymbiosis in 

close relation with the shrimp integument. The distribution of the 

bacteria give evidence of three functional compartments in the gill 

chambers while their internal ultrastructure suggests the presence 

of various bacterial strains and metabolisms as also supported by 

genetic analyses. The survey of the moulting stages and the 

determination of their ratio in the shrimp population reveal a high 

moulting rate and the periodic bacterial re-colonisation of the gill 

chamber by the bacterial symbionts after each exuviation. TEM 

observations also showed the deposition of cuticle and the 

ultrastructure of the epidermis at each moulting stage. 

Characteristics of the gills as well as of inner walls of the gill 

chambers (inner branchiostegites) colonised by the bacteria are 

those of permeable/transporting tissues (thin cuticle, numerous 

mitochondria, membranes infoldings, sulphide-oxidising bodies) 

suggesting the possibility of symbiont-host transtegumental 

transfers. The bacterial metabolism and the hypothesis of direct 

nutritional transfer through the shrimp integument are investigated 

using cold and radioactive tracers. The authors thank the belgian Fund 

for Joint Basic Research (FRFC-Belgium, conv. n° 2.4594.07.F) and the 

IFREMER (France) for the financial support. 

Do the hydrothermal vent fluid characteristics influence on 

the ectosymbiotic bacteria and associated minerals in the 

shrimp Rimicaris exoculata ? 

Laure Corbari 1 , Marie-Anne Cambon-Bonavita 2 , Magali Zbinden 3 , 

Françoise Gaill 3 and Philippe Compère 1 

1 Univ. Liège, Lab. de Morphologie fonctionnelle et évolutive, Unité 

de Morphologie ultrastructurale, allée de la chimie, 3, 4000 Liège, 

Belgium; 2 Laboratoire de Microbiologie et Biotechnologie des 

Extrêmophiles, Ifremer, centre de Brest, BP 70, F-29280 Plouzané, 

France; 3 UMR CNRS 7138 ‘Systématique, Adaptation et Evolution’, 

UPMC, 7 Quai St Bernard, 75252 Paris Cedex 05, France 

The shrimps Rimicaris exoculata are the primary consumers 

dominating the fauna of many hydrothermal vent sites Mid-Atlantic 

Ridge (MAR). They harbour in their gill chambers a rich 

ectosymbiosis of chemosynthetic bacteria that are supposed to be 

fuelled by vent fluid reduced compounds. Up to now, most of the 

recent studies considered shrimps from the Rainbow vent field 

(36°14.0’N) while it has an atypical fluid composition, rich in Fe 2+ 

and relatively poor in HS - . They considered the distribution of 

bacteria in the shrimp gill chambers as well as their morphotypes, 

phylotypes and metabolisms. They also showed the rapid 

development of the bacterial colonisation in phase with the shrimp 

moult cycle and the deposition of heavy bacteria-associated Fe 3+ - 

oxides. Interestingly, the TAG vent field (26°08.0’N), characterised 

by a sulphide-rich and iron-poor, fluid, also harbours an important 

shrimp population. For comparison, the bacterial population and 

the associated-minerals were characterised by electron 

microscopy and EDAX in shrimps from the TAG vent fields and 

followed throughout the shrimp moult cycle. 

- 79 - 

Differences were neither obtained in the shrimp moult cycle nor in 

the bacterial colonisation of the new cuticle after exuviation. The 

bacteria morphotypes also appear identical in both sites. Genetic 

analyses have indicated that the bacteria are of the same genera 

(mainly epsilon and gamma Proteobacteria) but the species seems 

to be different between TAG and rainbow. In contrast, obvious 

differences are revealed in the deposition of mineral. In TAG 

shrimps, a diffuse sulphide precipitate, giving grey-coloured 

shrimps, precedes iron oxide deposition that occurs less rapidly 

than in Rainbow shrimps. Moreover, the importance of minor 

elements (Si, Ca, P) is increased in accordance with the TAG vent 

fluid composition. The authors thank the Belgian fund for Joint 

Basic Research (FRFC-Belgium; conv. n° 2.4594.07.F) and 

Ifremer (France) for the financial support. 

New digestive symbioses in the hydrothermal vent 

Amphipoda Ventiella sulfuris. 

Laure Corbari 1 , Françoise Gaill 2 and Philippe Compère 1 


l’Environnement, Institut de Chime (B6c), allée de la chimie, 3, B- 

4000 Liège, Belgium ; 2 UPMC, UMR CNRS 7138 ‘Systématique, 

Adaptation et Evolution’, 7 Quai St Bernard, 75252 Paris Cedex 05, 

France 

Ventiella sulfuris is the most abundant amphipod species 

inhabiting the Eastern Pacific Rise (EPR 9°N) vent fields. This 

endemic species is commonly encountered near colonies of 

Pompeii worms Alvinella pompejana. That these species could 

live in a close trophic association never was further investigated. V. 

sulfuris specimens were collected during the oceanographic cruise 

LADDER II to the Bio9 (9°50.3’ N, 2508m depth) hydrothermal 

vent site. Looking for associated microorganisms, the integument 

and the digestive tract of the amphipod were observed in light 

microscopy (LM) and electron microscopy (SEM, TEM). The 

cuticle surface of the outer body and appendages appeared free of 

microorganisms. In contrast, the observation of the digestive tract 

revealed two important bacterial colonisations located in the 

mesenteron and in the proctodeum respectively. Both exhibit 

typical characteristics of symbioses. In the mesenteron, long 

bacteria are inserted between microvilli of endodermic epithelial 

cells. In the proctodeum, large densities of rods cover the cuticular 

walls. Examination of the gut content showed abundant fragments 

of annelid cuticle, most probably from A. pompejana, as well as 

mineral particles and diatom frustules. These results reveal new 

potential bacterial symbioses in vent crustaceans and point out the 

importance of amphipods in the trophic relationships of 

hydrothermal vent ecosystems. 

The authors thank the belgian Fund for Joint Basic Research 

(FRFC-Belgium, conv. n° 2.4594.07.F) for the financial support. 

The LADDER project was funded by NSF Ocean Sciences grant 

OCE-0424953. 

Temperature resistance studies on deep-sea vent shrimp 

Delphine Cottin, Juliette Ravaux, Nelly Léger and Bruce Shillito 

UPMC, UMR 7138, "Systématique, Adaptation et Evolution", 

75005 Paris, France 

The deep-sea hydrothermal vent ecosystem is an extreme 

environment characterized by great spatial and temporal 

instabilities and especially in terms of temperature. In this 

environment, the temperature can vary from 2°C to more than 

50°C in a few centimeters. As a consequence, in addition to a high 

ambient hydrostatic pressure, the vent fauna have to deal with 

harsh and highly unstable thermal conditions with frequent 

temperature burst. The caridean shrimp Rimicaris exoculata and 

Mirocaris fortunata dominate the vent megafauna at many of the 

hydrothermal vent field at the Mid-Atlantic Ridge. The first one, 

Rimicaris exoculata which forms dense swarm around the black 

smoker chimneys, is found in the hottest end part of the edifice, 

where it maintains close proximity to the superheated fluid. The 

second one, Mirocaris fortunata, is more broadly distributed across 

the vent-fluid influence gradient and thus is supposed be a less 

thermotolerant species. We performed in vivo experiments on R. 

exoculata and M. fortunata in pressurized aquaria to determine 

their upper thermal limit and we found that both species does not 

tolerate sustained exposure to temperature above 37°C. These


results show firstly that tolerance to heat is not a pre-requisite for 

life on a smoker wall and secondly that temperature resistance 

does not appear to be a crucial factor for explaining differences in 

distribution of shrimp species in a given vent sites. Moreover, the 

thermal resistance of vent shrimp is comparable to those of other 

non-vent tropical caridean species and thus does not place them 

amongst the most thermophilic metazoans. In this study on deepsea 

vent shrimp, we also investigate the influence of pressure on 

temperature resistance properties since pressure-dependant 

thermal characteristics have been reported for various biological 

systems. First, we determined the upper thermal limit of M. 

fortunata specimens originating from different sites and depths 

(850 to 1700 m) and secondly we evaluated the upper thermal limit 

of R. exoculata specimens originating from 2300 m, at different 

pressures in pressurized aquaria. In the case of M. fortunata 

originating from different depths, it appears that temperature 

resistance properties are not influence by depth of occurrence. 

However, for R. exoculata originating from 2300 m depth and 

pressurized at different pressures, the temperature at which the 

first signs of thermal discomfort occur seems to vary with pressure. 

Further studies are required to better understand the role of 

pressure in colonization processes. 

Symbioses between deep-sea mussels (Mytilidae: 

Bathymodiolinae) and chemosynthetic bacteria: diversity, 

function and evolution 

Sébastien Duperron 

UPMC-Paris 6 & CNRS, UMR 7138 Systématique, Adaptation et 

Evolution, 7 quai St Bernard, 75005 Paris, France 

Mussels of the subfamily Bathymodiolinae thrive around chimneys 

emitting hot fluids at deep sea hydrothermal vents, as well as at 

cold seeps and on sunken organic debris such as sunken wood 

and whale falls. Despite the absence of light-driven primary 

production in these deep-sea ecosystems, mussels succeed 

reaching high biomasses thanks to chemosynthetic, carbon-fixing 

bacterial symbionts located in their gill tissue (1). Since the 

discovery of mussel symbioses about three decades ago our 

knowledge has increased, mainly regarding large vent and! seep 

species, and new findings are published regularly. Recent studies 

have pointed out new interesting facts such as the presence of 

multiple symbionts in the gills of two cold seep mussels (2), the 

distinct intra- or extracellular localization of bacteria in the gills of 

mussel species from the same habitat (3), and the direct influence 

of immediate environmental parameters on symbiont densities (4). 

The study of smaller, less spectacular, species from cold seeps 

and organic falls has also shed new light on the diversity and 

evolution of mussel symbioses. In this talk, we’ll summarize 

current knowledge about symbiosis in Bathymodiolinae, with an 

emphasis on bacterial diversity and evolution. Future prospects will 

be discussed. 

Riftia pachyptila: a hydrothermal vent tubeworm as poster 

child for thioautotrophic symbioses 

Horst Felbeck 

Scripps Institution of Oceanography, University of California San 

Diego 0202, La Jolla, CA 92093, USA 

Riftia pachyptila has no digestive tract as an adult and is 

dependend on nutritional supply through intracellular 

chemoautotrophic symbionts harbored in a special tissue, the 

trophosome. The symbionts are acquired new in each generation 

of the host. Their metabolic interactions with the host and the 

origin of the free-living form have remained unclear for many years. 

Recent information gained from proteomics and the sequencing of 

the majority of the symbiont’s genome provides new clues about 

its identity, metabolic capabilities and the presence of nonexpressed 

genes. Apparently, only one species of symbiont, or 

possibly several strains of one species, populates a specimen of R. 

pachyptila. The name Endoriftia persephone has been proposed 

for the symbiont. The symbiont may have two ways to fix CO2 as 

autotrophic sources of carbon for itself and the host, the Calvin- 

Benson cycle and the reductive tricarboxylic acid cycle. The 

activity of the reductive tricarboxylic acid cycle could explain the 

unusual carbon isotope ratio characteristic for the tubeworm. The 

- 80 - 

relative significance of the two ways of carbon fixation may be 

dependend on the environmental conditions. Non expressed 

genes hint to possible capabilities of the symbiont as a heterotroph 

while free-living between generations of the host. 

Cell interactions induced by bacterial infection processes 

within the lateral zone of gill filaments of the lucinid Codakia 

orbiculata. 

Sylvie Gustave, Nathalie Elisabeth and Olivier Gros 

UMR-CNRS 7138, Systématique-Adaptation-Evolution, Equipe 

«Symbiose», Université des Antilles et de la Guyane, UFR des 

Sciences Exactes et Naturelles, Département de Biologie. B.P. 

592. 97159 Pointe-à-Pitre Cedex, Guadeloupe, France. 

Codakia orbiculata (Montagu, 1802) is a shallow-water lucinid 

which inhabits low sulfide sediments in seagrasses of Thalassia 

testudinum, and harbors sulfuroxidizing endosymbiotic bacteria 

within bacteriocytes of its gill filaments. Here, we attempted to 

evidence the mechanisms that might underlie the adaptative 

plasticity of the cells in the lateral zone of gill filaments. Two sets of 

starved individuals (3-month and 5-month starvation periods) were 

put back in their natural environment then collected at daily interval 

before treatment with thymidine analogue bromodeoxyuridine 

(BrdU) which is a cell division marker, or with cytosine analogue 

cytosine arabinoside (Ara-C) which is an antimitotic agent. 

Technical approaches were made by CARD-FISH, 

immunoshistochemistry, and histology in order to monitor the 

bacterial infection processes in correlation with the variation of cell 

organization within gill filaments, and to detect the new synthetized 

cells. In starved individuals, non symbiotic granule cells become 

the majority cells while bacteriocytes disappeared, whereas during 

bacterial colonization, these cells had been partially replaced by 

bacteriocytes which became predominant. One explanation of 

such cell variability might be cell proliferations regulated by the 

number of endosymbionts within the bacteriocytes. 

3D FISH for the quantification of methane- and sulphoxidising 

endosymbionts in bacteriocytes of the hydrothermal vent 

mussel Bathymodiolus azoricus 

Sébastien Halary 1 , Virginie Riou 3,4 , Françoise Gaill 1 , Thomas 

Boudier 2 and Sébastien Duperron 1 

1 UMR 7138 Systématique Adaptation Evolution, équipe 

Adaptation aux Milieux Extrêmes, Université Pierre et Marie Curie, 

7 quai St Bernard, 75005 Paris, France; 2 UMR 7101 Neurobiologie 

des Signaux Intercellulaires, Université Pierre et Marie Curie, 9 

Quai St Bernard, 75005 Paris, Franc; 3 IMAR Centre of the 

University of Azores, Department of Oceanography and Fisheries, 

Rua Cais de Santa Cruz, 9900 Horta, Portugal; 4 Department of 

Analytical and Environmental Chemistry, Vrije Universiteit Brussels, 

Pleinlaan 2, 1040 Brussel, Belgium 

Dual endosymbioses involving methane- and sulphur-oxidising 

bacteria occur in the gills of several species of mussels from deepsea 

hydrothermal vents and cold seeps. Variations of total and 

relative abundances of symbionts depending on local 

environmental parameters are not yet understood, due to a lack of 

reliable quantification of bacteria in the host tissue. Here we report 

the first attempt to quantify volumes occupied by each type of 

symbiont in bacteriocyte sections from a vent mussel, 

Bathymodiolus azoricus, using fluorescence in situ hybridization 

(FISH) coupled to 3D microscopy and image analysis carried out 

by a dedicated software which we developped. Bacteriocytes from 

mussels recovered at different vent sites displayed significantly 

different abundances of bacteria. Specimens kept in aquaria at 

atmospheric pressure and exposed to an artificial pulse of sulphur 

displayed an increase in absolute and relative abundance of 

sulphur-oxidisers within their bacteriocytes. Distributions of all 

measured parameters fitted normal distributions, indicating that 

bacteriocytes from a specimen tend to display similar behaviours. 

This study shows that symbiont volume quantification is tractable 

using 3D FISH and confirms the impact of local environmental 

parameters on symbiont abundances.


Biodiversity and colonization of Antarctic cold waters: the 

importance of symbioses associated to echinoids 

Clotilde Hardy 1 , Thomas Saucède 1 , Bruno David 1 , Chantal De 

Ridder 2 and Thierry Rigaud 1 

1 

Biogéosciences, CNRS/Université de Bourgogne, 6 bvd Gabriel, 

21000 Dijon, France 

2 

Laboratoire de Biologie Marine, Université Libre de Bruxelles, 

1050 Bruxelles, Belgium 

Among the main factors that control the structure of Antarctic 

benthic communities, the nature of the bottom and interspecific 

relations play an important part. Hard bottoms are not a common 

feature of Antarctic marine environments, and hard substrates, 

either biotic or abiotic, are generally colonized by many epibiotic 

invertebrates. Since primary spines of ctenocidarids - one 

subfamily of highly endemic Antarctic echinoids - are devoid of 

epithelium and of any anti-fouling system, they offer a hard 

substrate to many invertebrates. Even as one of the most speciose 

group of invertebrates in the Antarctic, echinoids (and their 

symbionts) may be at last of some importance for the diversity of 

Antarctic benthic ecosystems. To test the hypothesis, abundance, 

distribution, richness, diversity and similarity of symbionts present 

on two ctenocidarids (Notocidaris mortenseni and Ctenocidaris 

nutrix) have been compared with those of epibionts present on 

abiotic substrates (drop stones). Echinoids and stones were 

collected during the expedition ANTXXIII/8 (winter 2006-2007) in 

contrasted stations and areas of the Weddell Sea (Atka Bay), 

South Shetlands (Elephant Island) and Antarctic Peninsula (Larsen 

embayments in which iceshelves very recently collapsed). Our 

results demonstrate the importance of cidarids for local diversity 

and show contrasted patterns, likely due to contrasted conditions 

(latitude, depth and currents) prevailing in sampling areas. In the 

Larsen area, symbiotic communities distinguish by a low diversity 

and a strong similarity with epibionts present on stones, what is 

congruent with the conditions prevailing in Larsen: relatively poor 

and newly colonized habitats, both for benthic and symbiotic 

communities. 

Feeding and gut symbionts of decapod crustaceans 

associated to deep-sea wood falls: ultrastructural, trophic and 

molecular approaches 

Caroline Hoyoux 1 , Magali Zbinden 2 , Sarah Samadi 3 , Gilles 

Lepoint 1 , Pierre Becker 4 , Chantal De Ridder 4 , Renaud Berlémont 5 , 

Moreno Galleni 5 , Françoise Gaill 2 and Philippe Compère 1 


l’Environnement, B-4000 Liège, Belgium; 2 Université Pierre et 

Marie Curie, UMR 7138 CNRS, F-72252 Paris cedex 05, France; 3 

Muséum National d’Histoire Naturelle, UMS 2700 CNRS, F-72252 

Paris cedex 05, France; 4 Université Libre de Bruxelles, Biologie 

Marine, B-1000 Bruxelles, Belgium; 5 Université de Liège, Dept. 

des Science de la Vie, B-4000 Liège, Belgium 

Almost unknown few years ago, wood falls in the deep sea are 

now the subject of recent studies showing their importance as 

nutriments on the deep-sea floor. They support original 

ecosystems, with a macrofauna dominated by molluscs and 

crustaceans that belong to taxa also found in other reducing 

chemosynthetic environments as hydrothermal vents and whale 

falls. Because previous studies pointed out digestive and 

chemosynthetic bacterial symbioses in mollusc species from 

sunken woods, many questions arise about crustaceans, their role 

in the degradation of wood, their feeding biology and their 

microbial associations. 

An ultrastructural survey of the gut content and gut lining in 12 

crustacean species collected during 4 oceanographic cruises on 

sunken wood accumulations, near Pacific islands (Solomon, 

Vanuatu), pointed out 2 xylophagous and 2 detritivorous species 

(Munidopsis spp. and Callianassa spp.) harbouring resident, 

potentially symbiotic, gut microorganisms. These microorganisms 

mainly consist of bacteria and fungi attached to the hindgut cuticle 

lining. Determinations of stable isotope ratio ( 13 C/ 12 C, 15 N/ 14 N) are 

carried out to specify or confirm the diet and the trophic level of the 

species, as well as the possible implication of the gut 

microorganisms. Currently, the identification of the gut 

microorganisms is in progress by sequence analyses of the 16S 

- 81 - 

rRNA gene. Finally, finding genes for cellulose hydrolysis is in 

process by metagenomic analysis of bacterial gut communities. 

The authors thank the National fund for Scientific Research 

(FNRS-Belgium conv. FRFC n° 2.4.594.07.F) for the financial 

support. 

Extraordinary high diversity of siliceous sponges play an 

important ecological role in the deep Southern Ocean 

Dorte Janussen 

Forschungsinstitut und Naturmuseum Senckenberg, Senckenberganlage 

25, D-60325 Frankfurt am Main, Germany 

Sponges are important in marine benthic ecology by providing 

habitates for a variety of other organisms, ranging from microbial 

to megafaunal communities. Furthermore, large siliceous sponges 

(mainly of the class Hexactinellida) commonly possess prominent 

siliceous “needles”, which remain as spicule mats after death of 

the sponges and thus structure the sea floor and provide attractive 

substrates for the settlement of larvae. Rich associations of 

Porifera are thus linked with high benthic diversity and abundance 

of other animal phyla. This is obvious in the Southern Ocean, 

particularly on the Antartic shelf, where extraordinarily high sponge 

abundance and biomass are correllated with diverse associations 

of especially crustaceans, polychaetes and echinoderms. From 

bathyal to abyssal depths, most of the largely endemic Antarctic 

shelf sponge fauna is successively replaced by cosmopolitan taxa, 

and simultanously, the distribution of sponges and other sessile 

animals is becoming increasingly patchy. However, the diversity of 

especially the Hexactinellida (glass sponges) at above-species 

taxonomic levels is significantly higher in the Antarctic deep-sea 

than on the shelf. Especially bathyal depths between 1000m- 

2000m are characterized by rich associations of siliceous sponges, 

including both shallow and deep water taxa. This is due to the fact 

that on the Southern Ocean slope, eurybath species of endemic 

shelf species (e. g. Rossella spp.) co-occur with cosmopolitan 

deep-sea taxa (such as Bathydorus and Caulophacus spp.). 

With their structuring properties and large body size of many 

Antarctic siliceous sponges, these hghly diverse taxa play a crucial 

role for the colonization of deep-sea bottoms. 

Long-term monitoring of sunken wood colonization in the 

shallow water mangrove swamp of Guadeloupe (F.W.I.) 

Mélina Laurent 1 , Nadine Le Bris 2 , Françoise Gaill 3 and Olivier 

Gros 1 

1 UMR-CNRS 7138, Systématique-Adaptation-Evolution, Equipe 

« Symbiose ». Université des Antilles et de la Guyane. UFR des 

Sciences Exactes et Naturelles, Département de Biologie. B.P. 

592. 97159 Pointe-à-Pitre Cedex, Guadeloupe. France. 

2 IFREMER Département Etude des Ecosystèmes Profonds. Zone 

de la Pointe du Diable, BP70. 29280 Plouzané. France. 3 UMR- 

CNRS 7138, UPMC, 7 Quai Saint Bernard. 75005 Paris, France 

We realized experimental immersion of Cocos nucifera wood 

pieces in a mangrove swamp of Guadeloupe (16’N 61.5W) West 

Indies. Regular sampling allowed to study the kinetic of wood 

colonization from the very first days to several months of 

immersion. The organisms observed during the first week of 

immersion, are the ciliates Heterotrichea and Oligohymenophora 

and few nematodes. Bryozoans appear after 10 days and annelids 

after 20 days. After the first month, the abundance of ciliates and 

nematodes decrease, while several invertebrates including new 

ciliates, platyhelminthes, ascidians, annelids, bryozoans, 

crustaceans, gastropods, and few vertebrates as Gobiidae and 

Bleniidae colonize the wood. 

After the second month, the fauna abundance and diversity 

increase, with the apparition of cnidarians and echinoderms. 

Chemical monitoring (pH and hydrogen sulfide) realized with 

autonomous probes revealed sulfide contents reaching almost 

millimolar levels at the wood surface within 2.5 days through the 

first month, before a rapid decrease to sub-micromolar levels. A 

significant relationship between the presence of organisms and 

sulfide levels was suggested. This hypothesis was sustained by 

the detection of bacterial ectosymbionts using hybridization 

experiments, SEM and TEM observations in some of the 

invertebrates encountered.


Late colonizers, which do not possess bacterial coat, seemed to 

take advantage of the decrease of sulfide levels, while the 

symbiotic organisms more adapted to the high sulfide levels 

disappear. This study combining chemical and biological 

approaches allowed a better characterization of sunken wood 

colonization process. 

Molecular taxonomy and speciation patterns in mussels 

associated to organic falls 

Julien Lorion and Sarah Samadi 

MNHN, Service de Systématique Moléculaire 57 rue Cuvier F- 


The biodiversity in deep sea marine environments is still poorly 

known, what slows down the study of the evolutionary processes 

such as speciation and dispersion. In this context, hydrothermal 

vents and cold seeps have stirred up the attention of the biologists. 

Indeed in this extreme environmental conditions inhabit original 

communities of endemic o! rganisms. Among the most studied 

organisms living on those ecosystems are the Bathymodiolinae 

mussels. Bathymodiolinae mussels (Bivalvia: Mytilidae) were 

thoroughly studied in hydrothermal vents and cold seeps, where it 

was shown that they have symbiotic relationships with 

chemosynthetic bacteria. Other species inhabiting less known 

ecosystems such as sunken vegetable material and decaying 

bones of marine vertebrates form a clade with the Bathymodiolinea. 

Both plant material and bones, produce dihydrogene sulphides, 

from lignin or lipids respectively, when decaying on the sea floor 

allowing chimiosynthesis. Some species inhabiting sunken woods 

were yet shown to have chemosynthetic symbionts, among which 

the thiotrophic bacteria are the most common ones. Using an 

integrative taxonomy approach, we give new insights to the 

diversity of mussel species associated to organic falls and discuss 

which factors (such as geographic barriers or symbiotic 

relationships) drives the speciation in these reducing environments. 

Age, growth and mortality of largescaled scorpionfish 

(Scorpaena scrofa, Linnaeus, 1758) in the eastern Adriatic Sea 

Sanja Matić-Skoko, Miro Kraljević, Armin Pallaoro and Jakov 

Dulčić 

Institute of oceanography and fisheries, Spli, Croatia, Meštrovićevo 

šetalište 63, P.O.box. 500, 21 000 Split, Croatia 

The age, growth and mortality parameters of Scorpaena scrofa 

from the eastern middle Adriatic Sea were studied. Total lengths 

(TL) of 350 specimens ranging from 13.5 to 58.2 cm were obtained 

by trammel net fishing (2000-2005). Total weight was ranging from 

60.0 to 3650.0g. The mean lengths, as well as the age frequency 

distributions of males and females were not significantly different. 

Scales showed clearly the ring pattern. The opaque ring was 

deposited during the summer months. The length-weight 

relationship showed a positive allometric growth (b = 3.165; R 2 = 

0.989). The parameters of the von Bertalanffy growth equation 

were: L∞ = 68.20 cm; K = 0.084 per year; t0 = -1.378 year; R 2 = 

0.981. This study revealed that S. scrofa is a relatively slow 

growing and long-lived species with a life span in excess of 25 

years. The mortality parameters were: Z=0,27, M=0,22 and F=0,05. 

Description of two new Desmodora species: marine 

ectosymbiotic Nematode from Thalassia testudinum 

environment in Guadeloupe (F.W.I.) 

Leslie C. Maurin 1 , Janura Rzeznik-Orignac 2 and Olivier Gros 1 


« Symbiose ». Université des Antilles et de la Guyane, 

Guadeloupe. France. 

2 UMR-CNRS 5178 Biologie des Organismes Marins et 

Ecosystèmes, Département Milieux et Peuplements Aquatiques, 

MNHN, CP 53, 61 rue Buffon, 75231 Paris cedex 05, France 

- 82 - 

Two new morphotypes of marine nematodes belonging to the 

genus Desmodora (Desmodoridae) are found from calcareous 

sand of Thalassia testudinum sediment in Guadeloupe (F.W.I.). 

These two nematodes are characterized by an unstriated head 

region of thickened cuticle which forms a conspicuous cephalic 

capsule. Data obtained by molecular in situ hybridisation and 

electron microscopy (SEM and TEM) prove that these nematodes 

are associated with extracellular prokaryotes belonging to the 

gamma-proteobacteria. These two species present a thick 

bacterial coat constituted by a single morphotype of bacterial 

ectosymbionts. The average size of these bacteria is 3µm which 

are positioned all along the nematode’s body except in the 

cephalic region. The ultrastructural analyses of these bacteria 

show the presence of white inclusions located in the periplasmic 

space which are considered as elemental sulphur granules due to 

Raman microspectrometry data analysis. Such spectra permitted 

to detect this compound only in the bacterial coat of these 

Desmodora individuals. Nevertheless, within the Desmodoridae 

family only the Stilbonematidae subfamily is characterized by an 

obligate ectosymbiosis with sulphur-oxidizing bacteria covering 

their cuticle in a manner that is characteristic for the genus, and 

even species (Polz et al., 1992). In the actual classification the 

genus Desmodora belongs to the Desmodoridae family but is not 

included in the Stilbonematidae subfamily. 

Due to the observations realized in this study, these two 

ectosymbiotic Desmodora collected in the tropical sulfidic 

environment of Thalassia testudinum could have an emended 

classification in order to be transferred with the other ectosymbiotic 

nematodes of the Stilbonematidae subfamily. 

The impact of wood fall on marine Nematode abundance and 

diversity in the mangrove swamp sediment (Guadeloupe, 

F.W.I.) 

Leslie C. Maurin 1 , Nadine Le Bris 2 , and Olivier Gros 1 


« Symbiose ». Université des Antilles et de la Guyane, 

Guadeloupe. France ; 2 IFREMER Département études des 

écosystèmes profonds, Plouzané, France 

Abundance and diversity of nematode assemblage in the sediment 

surrounding an experimentally implanted sunken wood box in the 

mangrove swamp of Guadeloupe (F.W.I) were investigated at 15 

days during 3 months after placement. Samples of nematodes 

were taken at 0, 1, 2 and 5m distance away from the sunken wood 

box. The chemical analyses of: [H2S] measured by an autonomous 

probe (NKE, Inc.) directly to the bottom of the mangrove swamp 

while [O2] was obtained using microsensors from push cores 

realized at the same distance with an analysis at 0, 1, 2, 5, and 

10cm of depth. 

Preliminary results are shown indicating that the sediment is 

anoxic in the first millimeters of the top muddy layer for all sites 

analysed (close and far to the wood box). Concerning the H2S 

measurement, in the first 15 days, an enrichment of the sediment 

was observed in the vicinity of the wood box with a high 

concentration (millimolar content) from 2cm depth. The same 

concentrations were found at only 5cm depth for the distant sites. 

At 1 month, sulphide emission by the degradation of sunken 

woods is low and the sediment looses its high [H2S] concentration 

in surface. First observations concerning the nematofauna along a 

transect from experimentally sunken woods until 5m distance 

enable to show the presence of selective and non-selective 

deposit feeders (1A and 1B) nematodes close to the wood and a 

majority of predators/omnivores (2B) in the sediment far away the 

wood box. Preliminary data concerning the quantification and the 

distribution of nematodes through a long period (1.5 months) will 

be presented in order to connect chemical measurements with 

abundance and diversity (family and trophic groups) of nematodes 

along this transect. These results will permit to show the impact of 

the sulphide emission by the wood fall degradation in mangrove 

swamp on the nematofauna.


Chromosomal evolution in the Notothenia clade 

(Acanthomorpha, Nototheniidae) 

Catherine Ozouf-Costaz, Mélyne Hautecoeur 2 , Jean-Pierre 

Coutanceau 1 , Céline Bonillo 1 , Laura Ghigliotti 3 , Agnès Dettai 1 , G. 

Duhamel 2 and 

Eva Pisano 3 

1 

CNRS-MNHN, UMR7138, Dpt Systématique et Evolution, Paris, 

France 

2 

MNHN, UMR 5178 BOME, Dpt Milieux et Peuplements 

Aquatiques, Paris, France 

3 

Universita di Genova, Dpt di Biologia, Genova, Italy 

Antarctic fish suborder Notothenioidei (Acanthomorpha) exhibit a 

high rate of endemicity and represent a well known model of 

adaptive radiation in marine extreme environment. Within the 

family Nototheniidae, the clade Notothenia includes Antarctic and 

sub-Antarctic species, which display habitat ontogenic shifts 

between larvae and juveniles (pelagic) and adults (that live in 

shallow waters, and partly feed with algae). This phylum is clearly 

separated from other nototheniid clades on the base of 

morphological and molecular characters. It is characterized by 

common traits in chromosome diploid numbers that are particularly 

low (Notothenia coriiceps: 2n = 22; Notothenia rossii: 2n = 24; 

Notothenia angustata: 2n = 26, Paranotothenia magellanica: 2n = 

26, Paranotothenia microlepidota: 2n = 26), compared with the 

most frequent diploid number occurring in notothenioid fishes (2 = 

48). Their karyotypes are mainly composed of large metacentric 

chromosomes arising from Robertsonian rearrangements. Only 

one species, Notothenia (Indonotothenia) cyanobrancha, has 2n = 

48 chromosomes and juveniles that become benthic at very early 

stages. Its phylogenetic position is currently under revision with 

several molecular markers. We present here a detailed analysis of 

these species karyotypes, including location of 5S and 28S 

ribosomal genes by FISH. Main chromosome characters have 

been mapped onto the Notothenia cladogram, including extragroups 

chosen among other nototheniid clades so that 

chromosome changes and inter-relationships within this clade 

could be re-discussed. 

Natural and experimental sunken wood: analysis of 

substrates and associated fauna 

Marie Pailleret 1,2,3 , Nima Saedlou 3 , Catherine Privé-Gill 3 , Françoise 

Gaill 1,2 and Magali Zbinden 1,2 

1 Laboratoire « Systématique, évolution, adaptation », UMR 7138, 

université Pierre et Marie Curie, bâtiment A, 4e étage, 7, quai 

Saint-Bernard, 75252 Paris cedex 05, France ; 2 AMEX, UMR 

7138, CNRS, Université Pierre et Marie Curie, IRD, MNHN, 4, 

place Jussieu, 75252 Paris cedex 05, France ; 3 Laboratoire de 

paléobotanique et paléoécologie, UMR 5143, MNHN, case postale 

48, 57, rue Cuvier, 75231 Paris cedex 05, France 

Samples of natural sunken wood collected near Vanuatu (>500 mdepth) 

are identified based on histological observations in order to 

know their diversity, to infer their geographical origin and to 

appreciate their degradation state. Diversity of the associated 

fauna is also studied. Investigation of eventual specific 

associations between wood species and organisms constitute the 

second step of the study. Preliminary results concern two wood 

samples: they belong to two different families (Asteraceae or 

Onagraceae for the first sample; Fabaceae for the second one); 

they may have a local geographical origin as sample 1 may come 

from Polynesia and sample 2 may be endemic from Vanuatu; no 

degradation in their histological structure was observed. The two 

selected samples showed completely different colonisation 

patterns, which could be due to differences in chemical 

composition, to a selection of wood by fauna or to time elapsed 

since sinking. To bring comparative data, an in-situ experiment 

was established off the Nouméa coast. Four species of wood and 

three monocots were immersed at a depth of 900-1000 m during 

20 months. The seven samples did not equally respond to the 

immersion. One of them housed the most abundant and diversified 

fauna, the other samples were mostly colonised by two taxa. Major 

cell-wall degradation was also noticed on the densely colonised 

sample. 

- 83 - 

Bacterial symbioses in Nautiloids excretory organs: some 

evolutionary and functional aspects 

Mathieu Pernice 1 , Silke Wetzel 2 , Olivier Gros 3 , Gaute Lavick 2 , 

Renata Boucher-Rodoni 1 and Nicole Dubilier 2 

1 

UMR 5178 Biologie des Organismes Marins et Ecosystèmes, 

Département Peuplements et Milieux Aquatiques, Muséum 

National d’Histoire Naturelle, 55 rue Buffon, 75005 Paris, France 

2 

Max Planck Institute for Marine Microbiology, 28359 Bremen, 

Germany 

3 

UMR 7138 Systématique, Adaptation, Evolution, Département de 

Biologie, université des Antilles et de la Guyane B. P. 592, 97159 

Pointe à Pitre Cedex, Guadeloupe, France 

Symbiosis is an important driving force of metazoan evolution and 

the study of symbiotic associations in ancient lineages might 

provide further insight into the origin of several major adaptations. 

In this respect, symbiotic associations concerning the excretory 

organs of Nautilus (Cambrian origin: ca 500 mya), are of particular 

interest. Indeed, conversely to what is known in others 

cephalopods, in Nautilus most of the excretory processes (filtration, 

reabsorption, secretion) are assumed by the highly specialized 

pericardial appendages. In this study, we report that nautiluses 

from various geographical areas (Nautilus macromphalus from 

New Caledonia, and Nautilus pompilius from Philippines and from 

Vanuatu) harbour a high density of betaproteobacteria and 

spirochete phylotypes in their pericardial appendages. They were 

characterized by using various molecular approaches (16S rRNA 

phylogeny, CARD-FISH) and electron microscopy (TEM). This 

dual symbiosis concerns the genus Nautilus as it is not related to 

geographical origin of the specimens. 

CARD-FISH analyses relate bacteria distribution to the functional 

ultrastructure of the host organ, suggesting a symbiotic 

contribution to the excretory metabolism. First analyses by using 

ex-vivo incubations of the symbiotic complex in controlled medium 

suggest a bacterial implication in nitrogen metabolism of the host. 

Such symbiosis being rare among marine invertebrates, Nautilus 

bacterial symbiosis provides a great opportunity to investigate the 

influence of host-microbes interactions on a major physiological 

adaptation during the course of marine invertebrates’ evolution. 

Comparative phylogeography of deep-sea hydrothermal vent 

species along the east pacific rise 

Sophie Plouviez, Claire Daguin, Frédérique Viard, François Lallier 

and Didier Jollivet 

Equipe Ecophysiologie : Evolution et Adaptation Moléculaires, 

Station Biologique, BP 74, 29680 Roscoff, France 

Deep-sea hydrothermal vents are widely distributed along oceanic 

ridges throughout the globe. Venting is caused by deep-sea water 

infiltration into the oceanic basaltic crust where it is overheated 

and charged of metallic elements. The hot uplifted fluid (around 

350°C) is mixed to the cold and well-oxygenated bottom sea-water 

and precipitated to form large sulfide edifices. Hydro! thermal vent 

species are strictly associated with these sulfidic emissions as 

their nutritional needs only rely on chemolithoautotrophic bacteria. 

Deep-sea hydrothermal vent sites are highly fragmented and often 

separated by hundreds of kilometres with a rapid and highly 

variable turn-over both in time and space, which implies good 

dispersal capacities in order to (re)colonize new habitats. Dispersal 

is mainly restricted to the sea bottom layer along the ridge axis. 

Consequently, breaks that offset ridge axis (like transform faults) 

could disrupt gene flow and thus may promote genetic breaks, 

geographic structure of populations and, ultimately speciation. 

Here, we compared several phylogeographic patterns from 

different gastropod and polychaete species in order to test the 

possibility of vicariant processes and propose a global history of 

colonisation of vent species along the East Pacific Rise (EPR). 

First coalescence results using sequences of the Cytochrome 

Oxydase I mit! ochondrial gene indicated low geographic structure 

in nearly all studied species. Most networks displayed a star-like 

structure of same age, suggesting a recent demographic 

expansion. Breaks to gene flow are detected at different latitudes 

and often shared between groups of species and could be 

attributed to putative vicariant events.


Dispersal and colonization strategy of the whale bone eating 

worm Osedax 

Florence Pradillon, Masaru Kawato, Katsunori Fujikura, Chikayo 

Noda and Yoshihiro Fujiwara 

Extremobiosphere Research Center, Japan Agency for Marine- 

Earth Science and Technology (JAMSTEC), 2-15 Natsushima-cho, 

Yokosuka, Kanagawa, Japan 

In the deep-sea, whale falls bring massive organic enrichment in 

single pulses that significantly impact on local deep-sea 

communities. Whale falls provide island habitat for a specialized 

fauna adapted to live on whale remains 1 . The Siboglinid 

polychaetes Osedax exhibit a unique relationship to whale bones. 

Females grow a root organ that harbor symbiotic heterotrophic 

bacteria and that infiltrate bones and extract lipid and collagen for 

nutrition 2 . Since the discovery of this genus, 10 species have been 

reported, all associated to decaying bones, but in a wide range of 

environmental condition 3-6 . The success of Osedax in colonizing 

such a patchy habitat over broad geographic and bathymetric 

ranges implies efficient dispersal and colonization strategies. 

We analyzed colonization patterns of Osedax species on a sperm 

whale carcass implanted at 925 m depth in Sagami Bay (Japan). 

Seven different species successively colonizing the bones were 

identified using molecular techniques. O. japonicus that colonizes 

a close but shallow (225 m) sperm whale fall site 7 was not found, 

whereas 2 of the early colonists in Sagami Bay are also known 

from distant sites in the western Pacific (off California). This 

suggests very efficient dispersal possibly limited by depth related 

environmental parameters. We investigated embryonic and larval 

development in order to understand mechanisms underlying 

dispersal and colonization efficiency. Embryonic development 

exhibits remarkably rapid rates even at low temperature. This 

feature is usually associated with short dispersal range, suggesting 

that spatial and temporal frequency of suitable bones may be 

essential for maintaining Osedax population. 

Disclosing chemosynthesis in bivalve species from mud 

volcanoes in the gulf of Cadiz 

Clara F. Rodrigues, Gordon Webster, Andrew Weightman and 

Marina R. Cunha 

Campus Universitario de Santiago, 3810-093, Aveiro, Portugal 

Symbiotic associations with thiotrophic and methanotrophic 

bacteria occur in a wide array of animal species that live in 

reducing environments with high sulfide and methane 

concentrations, such as hydrothermal vents, deep-sea whale fall, 

sunken wood, and cold seeps. Since the discovery in 1999 of the 

first mud volcano in the Gulf of Cadiz, about 30 other sites at 

depths ranging from 200 to 3900 m, with varying degrees of 

hydrocarbon-rich gas seepage activity have been located and 

sampled. Chemosynthetic bivalve species include the widespread 

species Acharax sp. and Solemya sp but also other species like 

Lucinoma sp., Thyasira vulcolutre and Bathymodiolus mauritanicus. 

Stable isotopes analyses 13 C, 15 N, 34 S) show the major role of 

chemoautotrophic derived C in the food web of mud volcanoes. 13 C 

values measured in solemyids, lucinid and thyasirids specimens 

support the hypothesis of a thiotrophic nutrition. On the other hand, 

isotopic signatures of Bathymodiolus specimens collected were in 

line for methanotrophic nutrition. Molecular identification of 

chemosynthetic prokaryotic endosymbionts associated with these 

species was carried out using PCR-DGGE analysis of bacterial 

16S rRNA genes and analysis of bacterial clone libraries. 

Phylogenetic analysis of 16S rRNA genes from the gill tissues 

indicate that the bacterial sequences found were related to sulfuroxidizing 

endosymbionts from other deep-sea chemosynthetic 

environments. The divergence observed for Bathymodiolus 

mauritanicus (stable isotope values extremely depleted in 13 C 

suggested methano-trophic nutrition but molecular studies 

revealed the presence of sulfur-oxidizing bacteria) may be 

indicative of the occurrence of dual symbiosis but only further 

studies can corroborate that. 

- 84 - 

Transcriptomics study of the annelid host Riftia pachyptila 

symbiotic with chemosynthetic bacteria 

Sophie Sanchez 1 , Stéphane Hourdez 2 and François H. Lallier 2 

1Equipe 

Horloge circadienne, Laboratoire Arago, BP 74, 66650 

Banyuls sur mer, France 

2, 

Equipe Ecophysiologie : Evolution et Adaptation Moléculaires, 

Station Biologique, BP 74, 29680 Roscoff, France 

The giant hydrothermal vent tubeworm Riftia pachyptila (Siboglinid 

Polychaete Annelid) is probably one of the most extensively 

studied organisms that are symbiotic with sulfide-oxidizing bacteria. 

Located in a specialized internal organ, the trophosome (TR), 

these symbionts depend on the host for their supply in O2, CO2 

and H2S that are acquired from the environment by the branchial 

plume (BR) of the worm. We used a global molecular approach by 

Subtractive Suppression Hybridization (SSH) to try to identify 

proteins involved in the metabolites exchanges in these tissues. 

Four subtractive cDNA libraries were constituted, with the body 

wall (BW) as a reference tissue (BR-BW, BW-BR, TR-BW, BW- 

TR). More than 700 sequences were obtained with an average of 

45 different cDNAs per library, of which half could be identified. 

The differential expression of the most interesting transcripts was 

confirmed by quantitative PCR. A new carbonic anhydrase 

transcript (RpCAbr) specifically expressed in the gills was obtained 

in addition to the one already known (RpCAtr) that is mostly 

expressed in the trophosome (checking by fluorescent in situ 

hybridization). A myohemerythrin transcript, obtained in the TR- 

BW library, appears clearly specific of the trophosome tissue. 

Given that it is a particularly rare pigment in Annelids, this 

transcript was overexpressed in a heterologous system to explore 

a more functional aspect. In the branchial plume tissue, the Major 

Vault Protein could constitute an interesting clue for further studies 

about the transport of metabolites. 

Changes in lipid quality and fatty acids profile of sardine oil 

(Sardina pilchardus) during storage: The effects of 

temperature and α-tocopherol 

Salah Selmi 1 , Nawzet Bouriga 2 , S. Sadok 1 , Irineu Batista 3 , Narcisa 

Bandarra 3 and Maria Leonor Nunes 3 

1 

Institut National des Sciences et Technologies de la Mer, Port La 

Goulette 2060, Tunisia 

2 

Unité de Biologie marine, Faculté des Sciences de Tunis, 2092 

Campus universitaire, Tunisia 

3 

Department of Technological Innovation and Upgrading of 

Fishery Products, INRB/IPIMAR, Av. de Brasília, 1449-006 Lisboa, 

Portugal 

Sardine oil obtained through industrial mince preparation was 

stored at two different temperatures (+4C° and +35°C) during 28 

days with or without the addition of α-tocopherol. The peroxide 

values and fatty acid composition were determined to monitor 

oxidative lipid changes during storage. Peroxide values increased 

significantly (p < 0.05) during storage to reach 4.36 mEq O2/kg oil 

and 24.9 mEq O2/kg oil at +4°C and +35°C, respectively. Oil 

treated with α-tocopherol showed lower hydroperoxyde levels than 

did the control lot (3.3 mEq O2/kg oil and 21.47 mEq O2/kg oil at 

+4°C and +35°C, respectively). 

At +4°C, saturated and monounsaturated fatty acids remained 

constant after storage. Polyunsaturated fatty acids (34.05%) 

decreased significantly at the end of storage (31.3%). At +35°C, a 

50% decrease of polyunsaturated fatty acids was observed after 

28 days of storage, reaching 18.2%. Saturated and 

monounsaturated fatty acids instead showed a significant increase. 

It is concluded that α-tocopherol has a positive affect on the 

oxidation of sardine oil stored at +35°C.


Fat reserves and moisture content in relation to sexual cycle 

of sardine, Sardina pilchardus (Walbaum., 1792), in the 

eastern middle adriatic fishery grounds 

Gorenka Sinovčić and Bosiljka Mustać 

Institute of oceanography and Fisheries, Split, Croatia 

The fat reserves, moisture and sexual cycle of sardine, Sardina 

pilchardus (Walbaum, 1792), were studied from monthly random 

samples of purse seine catches from March 2004 to February 

2005. Catches were realised in offshore (Dugi Otok) and inshore 

waters (Virsko more) of the Mid Adriatic Sea. A total of 1,219 

sardine specimens were collected, out of which 668 were males 

and 541 females. Fish were measured, weighed and sexed. The 

sexual cycle analysis was based on the temporal evolution of 

gonadosomatic index and gonad mass. Gonadosomatic index 

(GSI) was calculated by expressing the monthly gonad weight as a 

proportion of the total body weight. The fat content was examined 

on the basis of monthly analyses of mesenteric fat in the visceral 

cavity and by determining the amount of lipid content in sardine 

tissues using Soxhlet’s s method. The total length of sardine 

ranged from 13.0 to 19.0 cm and the mass ranged from 16.72 to 

51.45 g. The reproductive period was from October to May, 

coinciding with the highest gonad weights and gonadosomatic 

indices. The mean percentage of mesenteric fat grades in visceral 

cavity points to the greatest fat quantities in August, when grade 4 

(fattest fish) presence amounted to 72%. The value is the result of 

an increase of mesenteric fat started in June and proceeding in the 

successive months until October. During autumn, a decrease trend 

is evident and it becomes pronounced in winter and spring when 

the lowest mesenteric fat quantities are recorded. Thereafter an 

increase in mesenteric fat for sardine was recorded, indicated by 

the records of grades 3 and 4. Due to such variations in 

mesenteric fat and tissue lipid during year, i.e. its greatest levels in 

summer (out of spawning) and lowest levels in the colder part of 

the year (during sardine spawning), it was assumed that quantity 

of sardine fat was affected by its sexual cycle. Lipid content in 

terms of sardine tissue dry weight showed the same trends as 

mesenteric fat - highest amounts during resting phase of 

reproduction (41.1%) and lowest during the peak of spawning (1.0- 

2.2%). Amounts of total lipids and moisture in sardine tissues 

showed that females had more fat content and less moisture than 

males. An inverse correlation between fat content and sexual 

cycle on one side and lipid content and moisture on the other has 

been noted. 

Nutritional strategies of Pectinodonta sp., a gastropod 

associated with sunken woods 

Magali Zbinden 1,2 , Marie Pailleret 1,2,3 , Juliette Ravaux 1,2 , Sébastien 

Halary 1,2 , Françoise Gaill 1,2 and Sébastien Duperron 1,2 

1 UPMC, Laboratoire Systématique Adaptation et Evolution, 7 Quai 

St Bernard, 75252 Paris, France; 

2 CNRS, UMR7138, Systématique, Adaptations, Evolution, AMEX, 

7 Quai St Bernard, 75252 Paris, France; 

3 CNRS UMR 5143 Paléodiversité et Paléoenvironnements, 

Laboratoire de Paléobotanique, UPMC, 12 rue Cuvier, 75005 Paris, 

France 

The occurrence of sunken wood and vegetal debris in deep 

oceanic environments has been known for a long time (Murray 

1895; Wolff 1979). 

Sunken woods are abundant in the deep sea, at all depths, with 

higher densities near estuaries, and in sedimentary accumulation 

basins. Wood reaches abyssal depths in sufficient quantity to 

support locally the development of long-lasting ecosystems, 

playing a key role in the deep sea (Cayré & Richer de Forges 

2002). 

The faunal diversity that colonize sunken woods is widespread, 

comprising gastropod, polyplacophoran and bivalve molluscs, 

decapod and peracarid crustaceans, polychaetes, and 

echinoderms (Turner 1973, 1977, Cayré & Richer de Forges 2002). 

Until now, the only well known organisms able to degrade wood 

(thanks to symbiotic association with cellulolytic bacteria) and to 

use it as a food source are wood boring mollusca (Teredinidae and 

Xylophagainae) and isopodes crustacea (Limnoridae) (Gareth 

Jones et al. 2001). 

- 85 - 

During several cruises dedicated to sample sunken woods around 

Vanuatu, some species were observed stuck in hollows that they 

seem to dig at the surface of dregded woods. The most numerous 

species observed is a true limpet (Patellogastropoda) of the genus 

Pectinodonta, the species being still undescribed. 

The present work combine various approaches (microscopic 

observations of the radula, gills, and gut content; molecular 

methods to determine occurrence, localisation and phylogenetic 

position of associated microorganisms, and enzymatic assays to 

look for cellulolytic activity) to determine wether wood is a food 

source for this species and what are the terms and conditions of its 

degradation. 

New insigths on the metabolic diversity among the epibiotic 

microbial communitiy of the hydrothermal shrimp Rimicaris 

exoculata 

Magali Zbinden 1 , Bruce Shillito 1 , Nadine Le Bris 2 , C. de Villardi de 

Montlaur 1 , E. Roussel 3 , F. Guyot 4 , Françoise Gaill 1 and Marie- 

Anne Cambon-Bonavita 3 

1 

UMR CNRS 7138, Systématique, Adaptations et Evolution, 

UPMC, 7 Quai Saint Bernard, 75252 Paris cedex 05, France: 

2 

Département Environnement Profond, Ifremer DRO, BP 70, 

29280 Plouzané, France; 

3 Laboratory of Microbiology of Extreme Environments, Ifremer 

Brest, LM2E, UMR 6197, BP 70, 29280 Plouzané, France; 

4 

Institut de Physique du Globe de Paris, Laboratoire de 

Minéralogie-Cristallographie, Université Paris-Jussieu, Tour 16, 

Case 115, 4, place Jussieu, 75 252 Paris Cedex 05, France 

The shrimp Rimicaris exoculata (Williams and Rona, 1986) 

dominates the megafauna of some of the Mid Atlantic ridge 

hydrothermal vent sites. This species harbors a rich community of 

bacterial epibionts inside its gill chamber. Literature data indicate 

that a single 16S rRNA phylotype dominates this epibiotic 

community, and is assumed to be a sulfide-oxidizing bacteria. 

However attempts of cultivation were not successful and did not 

allow to confirm it. The aim of our study was to test the hypothesis 

of sulfide oxidation in the gill chamber, by a multidisciplinary 

approach, using in vivo experiments at in situ pressure in the 

presence of sulfide, microscopic observations and a molecular 

survey. Morphology of microorganisms, before and after treatment, 

was analyzed to test the effect of sulfide depletion and re-exposure. 

Our observations, as well as molecular data indicate a wider 

diversity than previously described for this shrimp’s epibiotic 

community. We observed occurrence of bacterial intracellular 

sulfur- and iron-enriched granules and some methanotrophic-like 

bacteria cells for the first time. Genes that are characteristic of 

methane-oxidizing (pmoA) and sulfide-oxidizing (APS) bacteria 

were identified. These results suggest that three metabolic types 

(iron, sulfide and methane oxidation) may co-occur within the 

epibiont community associated with Rimicaris exoculata. As this 

shrimp colonizes chemically contrasted environments, the relative 

abundance of each metabolic type could vary according to the 

local availability of reduced compounds.


New records of host-parasite interactions between garfish 

Belone belone (Linnaeus, 1761) and Copepod Peniculus 

fistula von Nordmann, 1832 (Siphonostomatoidea, Penelidae) 

in the Adriatic sea 

Barbara Zorica, Gorenka Sinovčić and Olja Vidjak 

Institute of Oceanography and Fisheries, Šetalište I. Meštrovića 63, 

P.O. Box. 500, 21000 Split, Croatia 

Garfish, Belone belone (L., 1761), is a migratory pelagic species 

that occurs in the north eastern Atlantic, Mediterranean and Black 

Sea. In the Adriatic Sea this species is widely distributed in costal 

and open sea waters. Four samples of garfish individuals were 

caught by purse seine and beach seine in the eastern mid-Adriatic 

Sea between February and March 2008. Out of 224 analysed 

specimens of garfish, 22.3% were infested by crustacean 

ectoparasite identified as Peniculus fistula von Nordmann, 1832. 

The ectoparasites ranged from 1 to 7 per host with a mean 

intensity of 1.64. The parasites were attached to the fins - 62.2% to 

the ventral fins, 20.7% to the pectoral fins, 11.0% to the anal fin, 

4.9% to the dorsal fin and 1.2% to the caudal fin. There was no 

correlation between the total length of the host and the number of 

ectoparasites (R=0.0715, p=0.622). The present study has been 

undertaken not only to register parasitic copepod P. fistula as a 

parasite on B. belone (L., 1761) for the first time, but also to 

contribute to the knowledge of pennellid fauna in this region. 

Besides, this parasite species might potentially be used as one of 

the biological tags for garfish stock discrimination in the 

Mediterranean and Adriatic Sea, inclusively. 

- 86 -


S21 - The ecological and evolutionary consequences of global climate evolution on population, 

species and ecosystem 

The Global Biodiversity Information Facility (GBIF): free and 

open access to biodiversity data 

Anne-Sophie Archambeau, Eric Chenin, Régine Vignes-Lebbe, 

Adil El Filali and Justine Cheval 

GBIF France, MNHN Géologie, CP 48, 43 Rue Buffon, 75005 

Paris, France 

The Global Biodiversity Information Facility (GBIF, 

http://www.gbif.org) is an international organization that is working 

to make the world's biodiversity data accessible anywhere in the 

world. Currently, 42 countries and 37 international organizations 

share their data and catalyzed agreements on many of the 

standards and protocols required to make disparate datasets 

compatible. 

Over 145 million of scientific data records from over 2000 datasets 

from 242 institutions worldwide are now accessible through the 

GBIF data portal (http://data.gbif.org). The two types of data 

currently being shared are: 

• Species occurrence records (based on specimens and 

observations) - information about the occurrence of species at 

particular times and places. 

• Names and classifications of organisms - information on the 

names (both scientific and common) used for species and on the 

classification of those organisms into taxonomic hierarchies. 

Most of the data shared by the GBIF network can be mapped 

geospatially, which allows a vast array of analysis such as 

ecological niche modelling. Different web services are also 

available and can be reused by others applications. A global 

biodiversity commons is thus now a reality, allowing access to 

previously inaccessible records, and analyses which were 

previously impossible. 

The national GBIF nodes develop tools to enlarge the GBIF 

activities and help dataproviders to connect their data to the GBIF 

portal. The French node (http://www.gbif.fr) can be contacted at 

the following address: gbif@gbif.fr and the following number: +33 

(0)1 40798065. 

Diversity, endemism and conservation priorities of desert fish 

in Algeria 

Rachid Bouhadad 

FSB/USTHB, Algiers, Algeria 

Two hydrographic networks exists in Algeria: the first is an actual 

basin, gathering the rivers, the natural lakes and other point of 

water in the North of the country, as for the other it’s a fossilized 

basin covering several points of water in the Sahara. The desert 

freshwater could be considered poorly studied - consequently, the 

systematic and the biogeography data are changed from an author 

to another one for several species, notably morph metric data’s. 

This part of view is perfectly verified by the recent work, which 

used molecular markers and other tools on the genus Barbus 

(detailed in text). The list of Algerian freshwater is established 

according to our investigations (which cover all the hydrographic 

network in the north and the Sahara), the species cited in literature 

which are often conserved in Museums 

According to the list of fish established, it provides that the 

Cyprinidae, the Cobitidae and the Cichlidae families are 

respectively more represented. In an other fact, endemic species 

(Barbus deserti and the catfish genus Clarias have been gathered 

(but threatened for the first species). 

The genus Barbus largely abundant as wild populations, is 

represented by diploid, tetraploid and hexaploid species in the 

North of Africa. According to the ploïdy diversity (three levels), and 

osteological characters, the genus Barbus has been considered as 

a phylogenetic assemblage. Five different groups have been 

identified in the sahara/: the diploid species (endangered) are 

restricted to the Ahaggar mountains (protected area actually), this 

situation seems to coincide with the morphological and taxonomic 

distribution proposition (Pellegrin, 1939) with some exceptions 

according to a study realised with biochemical markers. 

- 87 - 

The effect of climate change on bird communities 

Denis Couvet, Frédéric Jiguet and Romain Julliard 

UMR 5173 MNHN-CNRS-UPMC, 55 rue Buffon, 75005 Paris, 

France 

Birds offer the advantage that data are numerous, due to a longterm 

effort to organize a network of observers who collect data on 

numerous sites. 

Analyses of the results in France for the last twenty years shows 

very clearly changes of distribution towards northern latitudes. 

We will discuss which species are moving, how it is related to their 

life-histories, and how much these moves are as important as 

expected, or if one can consider there is a lag relative to climate 

changes. 

We will finally discuss functional consequences that can be 

derived from these different results. 

The impact of global warming on polar seas: expected 

changes on Antarctic Echinoid fauna and forecasts for the 

future? 

Bruno David 1 , María Eugenia Manjón-Cabeza 2 , Francina Moya 2 , 

Thérèse Choné 1 , Thomas Saucède 1 and Chantal De Ridder 3 

1 

Biogéosciences, CNRS/université de Bourgogne, 6 bvd Gabriel, 

21000 Dijon, France 

2 

Dpto. Biología Animal, Universidad de Málaga, Campus de 

Teatinos s/n, 29071 Málaga, Spain 

3 

Laboratoire de Biologie Marine, Université Libre de Bruxelles, 

1050 Bruxelles, Belgium 

Global models propose climate evolution scenarios that forecast a 

global warming strongly impacting Polar seas. In Antarctica, 

several studies have shown that fauna is already affected by 

global warming and that some species display new geographical 

distributions. In this context, it is important to set precisely the 

present day biogeographic distributions in order to have access to 

the dynamic of changes, and to the potential vulnerability of 

species. To address this question, biogeographic comparisons of 

echinoid fauna have been performed to settle links between South 

American and West Antarctica. Indeed, southward migrations of 

cold temperate fauna can be expected as the Magellanic area is in 

connection with the Antarctic Peninsula via South Georgia and 

South Sandwich islands and via ridges surrounding the Scotia Sea. 

Depending on their feeding habits (carnivorous, algivorous or 

omnivorous), trophic plasticity, and life strategies (brooders or 

indirect developers), we can expect the echinoids to cope in 

different ways with environmental changes. The data analyzed 

cover more than 150 years of exploration, up to the most recent 

cruises. Comparisons have been done considering several subregions 

and involve analyses of raw distribution data of the whole 

echinoid fauna, but also take in consideration more specific 

aspects such as reproductive strategies, feeding habits, or 

phylogeny. All these aspects allow to set hypotheses for the 

evolution of the echinoid fauna in the forthcoming decades, 

particularly regarding what group(s) would be more prone to 

migration/extinction processes as enhanced by global warming.


Changes in phenology of the ground beetle Pterostichus 

madidus (Coleoptera: Carabidae) as evidenced from a longterm 

dataset 

Gabor Pozsgai 

Craigiebuckler, AB15 8QH, Aberdeen, UK 

Long-term data from the UK Environment Change Network (ECN) 

were analysed to investigate whether there is a phenological 

change in life history of Pterostichus madidus. Pitfall trap data 

were available from 12 ECN sites across Britain, mo! st of which 

have been established for over 11 years. All ECN sites follow the 

same sampling protocol making data were suitable for trend 

analysis. Weather and vegetation datasets were also available for 

these sites. Pitfall trap lines – normally three at each site – were 

categorized to broadleaf woodland, arable field, calcareous 

grassland, coniferous woodland, permanent pasture, dry heather 

and acid grassland vegetation types. Trend analysis was carried 

out first using all the available capture data, then the datasets 

grouped by vegetation type and ECN site. Few Pterostichus 

madidus were caught on four sites and in three vegetation types, 

so these were eliminated from the analysis. Pterostichus madidus 

appeared significantly earlier, if tested the pooled data, at three 

sites and in two vegetation types. There were also trends, although 

not significant, of earlier appearance, in three vegetation types and 

at two sites. In many cases adult beetles also disappeared 

significantly earlier, possibly due to disadvantageous 

environmental conditions in late summer. These results suggest 

that in the last 11 years the Pterostichus madidus phenology has 

changed with, the adults tending to appear earlier. Global warming 

may be an important factor driving these changes. 

Comparing modelling procedures for forecasting variations in 

species distribution due to climate change 

Raimundo Real, Ana Luz Márquez, Alba Estrada and Jesús 

Olivero 

Dpto. Biología Animal, Universidad de Málaga, 29071 Málaga, 

Spain 

Given that mountain species are presumed to be more at risk than 

other species as a consequence of climate warming, we selected 

one amphibian (Baetic midwife toad), one reptile (Lataste’s viper), 

one bird (Bonelli’s eagle), and one mammal (Iberian wild goat) 

species present in Andalusian mountains (S Spain) to model their 

distributional response to climate change during this century. The 

climatic forecasts for the whole century were provided by the 

National Meteorological Institute of Spain, which adapted the 

global circulation models CGM2 and ECHAM4 to Spain and 

produced expected temperature and precipitation values according 

to the A2 and B2 emission scenarios for each circulation model. 

We tested several approaches to forecast future distribution. We 

modelled the response of the species to spatial, topographic, 

human, and climate variables separately. We compared each of 

these explanatory models using Akaike Information Criterion, and 

produced a combined model weighting those of each factor 

(spatial, topographic, human, and climatic) according to Akaike 

weights. This procedure overestimated the best model, and the 

other factors were neglected in the final model. We also produced 

a combined model using stepwise selection of the variables 

previously selected according to each factor. In this way every 

factor was represented in the final explanatory model of the 

distributional response of the species to environmental conditions. 

In the latest part of our approach, we forecasted the distribution of 

the species by replacing current climatic values with those 

expected according to each climate change scenario, while 

preserving spatial, topographic and human variables. 

- 88 - 

Global climate change and temperature effects on pest 

insects 

Mark Schlueter 

Georgia Gwinnett College, School of Science and Technology, 

1000 University Center Lane, Lawrenceville, Georgia 30043 USA 

Global climate change or elevated annual temperatures affects 

animal activity and reproduction. In insects, increased 

temperatures may decrease development time from egg to adult, 

increase metabolic rates, increase food consumption rates, and 

affect many other factors. 

In our laboratory, we have investigated the effects of temperature 

on reproductive rates and development time in Tribolium 

castaneum and T. confusum. Temperature has an interesting 

effect on the reproduction and development of Tribolium beetles. 

Increased temperature results in a much higher number of eggs 

laid by females; however, higher temperatures also decrease the 

chance an egg will hatch successfully. Similarly higher 

temperatures speed up pupa development time; however, 

increased temperature also decreases the chance of a successful 

pupation. Extreme temperatures may even cause infertility. Overall, 

Tribolium beetles exposed to higher temperatures exhibit faster 

colonization rates, increased activity, and increased food 

consumption rates. 

Understanding how global warming or temperature affects pest 

insects is very important to humans. Just consider the flour beetles, 

Tribolium. Each gram of flour consumed by Tribolium beetles is 

one less gram for humans. Stored food products have been rapidly 

declining in the past few years due to increased demands by 

human consumption and use of food products to create ethanol. 

Any additional loss from sources like pest insects may result in 

even greater food shortages. 

The effects of climate change on birth seasonality in freeranging 

populations of hanuman langurs: a review of 

hypotheses and theoretical predications 

Arun Srivastava 

2000 85th Street, 07047, North Bergen, USA 

This paper discusses the ultimate and proximate causes of birth 

seasonality in 23 populations of Hanuman langur (Semnopithecus 

entellus) and seasonal pattern shown by each population within 

populations. Review of literature! and data collected through 

survey questioner and direct field observations are consistent with 

the hypothesis that food availability is the most important ultimate 

cause of birth seasonality. Stepwise multiple regression equation 

was use to estimate how the degree of birth seasonality is affected 

by ecological and life history variables. This model shows that two 

factors rainfall and latitude affect the degree of birth seasonality. 

The degree of seasonality increases with latitude. To understand 

which of the three alternative birth strategies are followed (reduce 

energy stress during peak lactation; wean infants during peak food 

availability; or store reserve during peak energy availability), we 

compared the location of birth peak in relation to peak in food 

availability for those populations from which the data were 

available. Most populations can form a typical pattern of births 

concentration before the peak in food availability allowing peak 

lactation or weaning to take! place before the start of the lean 

season. The current understanding of climate change due to 

contraction of the continental ice sheet and the amount of solar 

radiation it is predicted that the Asian monsoon will boost the 

occurrence of severe floods in Indian subcontinent and drought in 

western Indonesia through an Indian Ocean Dipole (IOD). The 

implications of such climatic conditions on the survival and 

reproduction of other endangered primate species inhabiting south 

Asian region could bring significant challenges for conservation.


Ecological effects of climate change: an overview 

Nils Chr. Stenseth 

Centre for Ecological and Evolutionary Synthesis (CEES), 

University of Oslo, Norway 

This presentation will review a broad spectrum of studies 

demonstrating the ecological effects of climate change – covering 

terrestrial, marine and freshwater systems. Most of the reviewed 

studies are based on the statistical analysis of long-term 

monitoring data. The trust of the presentation is that we can 

prepare us for what might happen (and hence ought to be done) 

by looking backwards in time trying to understand how climate 

variation has effected ecological processes in the past. 

Phylogeographic pattern of the northern edge freshwater crab, 

Sinopotamon yangtsekiense: genetic divergence and 

evolutionary history inferred from mtDNA sequences 

Hongying Sun 1 , Xiuling Lu 1 , Naifa Liu 2* , Qi Li, Kaiya Zhou 1 , 

Daxiang Song 1 

1 Jiangsu Key Lab for Biodiversity and Biotechnology, College of 

Life Sciences, Nanjing Normal University, Nanjing 210046, China 

2 Lanzhou University, Lanzhou 730000, China 

Sinopotamon is one of the endemic genera to Mainland China, and 

a total of 75 species in this genus were recoded in the past nearly 

50 years. Of these, only a few distributed north to the Qinling 

Montains and the Huai River, and even distributed to both sides of 

the Yellow River. The Qinling Montains and the Huai River serve 

as natural boundary between southern subtropical and northern 

warm temperate in Chinese zoogeographic fauna. S. yantsekiensis 

is one of the representatives distributed in the drainages south 

from the Yangtze River and north to the Yellow River located in 

north of the Qinling Mountains, with rang across the natural 

transition between Palaearctic and Oriental regions. We used 

phylogeographic approach to elucidate the evolutionary history of 

S. yantsekiensis lineages restricted in the northern range of the 

species’ distribution; the three sub-clade statues of the species in 

the current taxonomy were also assessed with the use of partial 

sequences of mitochondrial cox1 and nad5 genes, based on 83 

individuals from 13 populations throughout the species’ range. Two 

major evolutionary lineages were recovered in our phylogenetic 

analyses. The northern lineages comprised of populations 

spanning from upper reaches of the Huai River to the Yellow River 

drainages, and was basal group in the phylogeny. The southern 

lineages included populations located in the lower reaches of the 

Yangtze River and Huai River. The divergence for the splits 

between the northern edge and elsewhere lineages occurred in the 

middle of Pliocene (2.72 Myr). Their evolutionary history was 

discussed. Grant from the National Natural Science Foundation of 

China Key Project No. 30600010 to DXS. 

- 89 - 

Insights inside population genetic data on effects of global 

climate changes 

De-Xing Zhang 

Institute of Zoology, Chinese Academy of Sciences, Beijing 

100101, China 

Scientists urge a deeper understanding of the ecological effects of 

global climate change, aiming to make reliable predictions of its 

consequences. Conventional experimental studies are faced with 

some ineluctable challenges, since both the spatial and temporal 

scales required for mimicking the process of climate changes are 

difficult, if not impossible, to implement in research practice, 

leaving aside the complexities of such processes. Here lie the 

issues that genetic studies of populations can complement. Global 

climate change, and other environmental stresses with strength 

and duration, will leave genetic imprints in populations of many 

species. This provides a unique window for gaining insights into 

ecological effects of past climate changes, such as the Pleistocene 

glaciations. Therefore, genetic data of natural populations 

preserved rich information about historical global changes the 

consequences of which can be examined by comparative studies 

of multiple species and across geography. A fairly wealthy amount 

of studies already attempted in this area and produced 

encouraging results. Currently, what is restricting in such 

retrospective investigations is not the data but the analyses that 

follow. 

Ecological Consequence of Global Climate Change on 

Population Abundances of Animals in China 

Zhibin Zhang 

State Key Laboratory of Integrated Pest Management of Insects 

and Rodents in Agriculture, Institute of Zoology, Chinese Academy 

of Sciences, Beijing 100101, P.R. China 

Recent IPPC report has reconfirmed the accelerated global 

warming trend of our earth in the new centaury. During past 

millennia, China has experienced obvious warm and cold phases 

or periods. Understanding how temperature oscillations influence 

animal spatial distribution and temporal abundances is extremely 

valuable in predicting potential impact of the on-going global 

warming on biological, ecological and social systems. China has 

unique records of many important animals such as elephant, 

panda, locust, rodent-borne plague ect. And recently, there has 

been great progress in climate reconstruction of temperature, 

precipitation, ect using ice core, pollen, tree ring or historical 

records. All of these make it possible in analyzing relationship 

between population dynamics and climate change. Besides, the 

irregular El Nino and Southern Oscillation (ENSO) has been also 

recognized to be important factor in affection climate anomalies in 

China. ENSO significantly alter precipitation and temperature 

pattern in China, thus it is likely to affect population dynamic of 

animals. This paper will give a brief review on recent progress in 

studying relationship between population dynamics and global 

climate change in China, with emphasis on discussing population 

change of many important endangered species and agricultural 

pests or diseases in China during past millennia or centuries.


Assessing the invasion potential of the Colorado potato 

beetle, Leptinotarsa decemlineata, to Northern Europe 

Sanna Boman 1 , Alessandro Grapputo 2 , Leena Lindström 1 , Anne 

Lyytinen 1 , Johanna Mappes 1 and S. Piiroinen 1 

1 

Department of Biological and Environmental Science, P.O. Box 

35, FI-40014 University of Jyväskylä, Finland 

2 

Department of Biology, University of Padua, Via U. Bassi, 58/B, 

35121 Padova, Italy 

To prevent or manage invasions of alien species, it is vital to study 

their invasion potential before they colonize new areas. We 

assessed the potential of a well-known invader, the Colorado 

potato beetle (Leptinotarsa decemlineata), to expand north of its 

current range in Europe by studying genetic variability in life history 

traits from four geographically distinct populations (Russia, Estonia, 

Poland and Italy). We further assessed the cold tolerance during 

development and whether mild stress (pesticide) confers cost in 

terms of metabolic rate and fat content. Our results show that 

European L. decemlineata still exhibit adaptive genetic variation in 

life-history traits, indicating that there is potential for evolution in 

these traits. In particular the northern population (St. Petersburg, 

Russia) of L..decemlineata developed significantly faster than the 

central one (Poland), indicating adaptation to a short growing 

season. Beetles were very cold tolerant as exposing larvae to 

subzero temperature (simulating night frosts) did not incur mortality. 

We found also that pesticide selection (i.e. parents exposed to 

pesticides) can cause significant costs in their offspring (smaller 

size, higher metabolic rate), which impairs overwintering success 

and thereby may retard their invasion potential and colonization 

success. This study demonstrates the importance of considering 

both ecological and evolutionary aspects when assessing invasion 

risk. 

A preliminary account of the green leafhopper Jacobiasca 

lybica in the northen vineyard of Algeria 

Farid Bounaceur 

16, street Hassan Badi el Harrach Algiers, 16200, El harrach 

Algiers, Algeria 

The dynamics of populations and damage on three industrial vines 

newly introduced in Algeria has been studied in a vineyard to the 

west of the plain of the Mitidja by numbering of larvae on whole 

plants and adults captured by yellow chromoactive traps during 

two consecutive years 2005 to 2006. The follow-up of populations 

showed the succession of four generations and those from the 

month of April until the month of August for the two years of the 

follow-up. The exam of infestations showed that the Merlot vine 

accuses big preferences in relation to cabernet sauvignon and the 

Syrah, indeed the number of green leafhopper observed on these 

last vines in 2nd and 3rd generations passes the recommended 

percentage extensively. The leaf damage was observed in all 

variety, so Merlot variety accused the most important damage, 

80% of their leaf were attacked, 45% about Cabernet-Sauvignon 

and 25% for Syrah variety. 

About the mealybug Planococcus ficus in northern 

vineayards of Algeria 

Farid Bounaceur, Farid Boustila, Atika Guendouz-Benrima and 

Bahia Doumandji-Mitiche 

16, street Hassan Badi el Harrach Algiers, 16200, El harrach 

Algiers, Algeria 

The scales on vines were seen from nearly a century as secondary 

pests. Recent experiments have shown their involvement in the 

transfer and the spread of the disease of winding vines, which may 

affect somewhat taking into account their control. Under a 

monitoring pest of vineyard in Algeria, mealybug proved as an 

important major economic pest in Northen Vineyard of Algeria. In 

S22 - Studies of invasive animal species 

- 90 - 

this study we describe and evaluate, infestations, estimate the 

damage and understand the sensitivity of type of wines in Mitidja 

areas Northern of Algeria. The monitoring was conducted during 

2007 season, during spring and summer, the results have shown 

differences between grapes and appellations, rates vary 

infestations, damage estimated seem to be important mainly for 

the some variety Mitidja. 

Reintroduction of Chinese water deer(Hydropotes inermis) 

to Shanghai, China 

Min Chen and Endi Zhang 

School of Life Science, East China Normal University, Shanghai 

200062, China 

Chinese water deer ( Hydropotes inermis ) is classified as the 

second class of protected animals in China, and ranked as LR/nt in 

IUCN red data book. The distribution and population of the water 

deer in China were declined rapidly, the species suffered habited 

lost. Based on partly survey, the totally number of the Chinese 

water deer was estimated not more than 10,000 in the field. It is 

necessary to making strategy on conservation and management to 

recovery the species. Assessing the feasibility for reintroduction of 

the Chinese water deer based on textual research of the deer 

populations in the history, survival condition, investigation of public 

attitude and local condition. The water deer only disappeared in 

Shanghai only about 100 years, and most surveyed local people 

would like to invest to reintroduction of the water deer. The project 

selected a plot in Shanghai to the reintroduction and aims to 

recovery Chinese water deer in Shanghai. A group of Chinese 

water deer have been released into a Park on the round-the-city 

greenbelt area between rural and urban of Shanghai in 2007. If the 

water deer can inhabit Shanghai and no incline to invasive local 

ecology by monitoring of this population, we will plan to release the 

deer to open areas around the city. 

A profile of bird invasions – how are they related to European 

politics? 

François Chiron and Salit Kark 

The Biodiversity Research Group, Dept. of Evolution, Systematics 

and Ecology, The Institute of Life Sciences, The Hebrew University 

of Jerusalem, Jerusalem 91904, Israel. 

We questioned whether European politics in the 20 th century were 

responsible for less introductions of birds in Eastern Europe than 

in Western Europe. We focused on the consequences of the socalled 

Cold War (1949-1991) that split Europe into two political and 

economic blocs. During this period, every aspect of trade and 

abilities of humans to travel have been affected, isolating the 

Eastern Europe from the rest of the world. To test embargo effects 

on bird introductions in Eastern Europe, we assessed variations of 

introduction efforts between the Eastern and the Western blocs, 

before, during and after the Cold War. We expected that the 

embargo has drastically disrupted introductions of birds imported 

from other continents. 

We used a very recent dataset on bird invasions we generated as 

part of DAISIE 1 consortium. Introduction histories are presented in 

the context of trade and people movement regulation policies and 

impacts of the Cold War were tested using an experimental design 

of analysis. Our results pointed out the effect of the embargo on 

the number of birds transported and released in Eastern Europe 

during the Cold War. Further, we showed the importance of 

multilateral agreements involving countries of a same bloc on 

pattern of introductions. 

This study suggests that politics can affect human activities and 

subsequent introductions of birds at the scale of Europe. Any large 

scale study should consider economic and political history as a key 

factor to understand bird invasion patterns. 

1 Delivering Alien Invasive Species Inventories for Europe


Exotic and introduced fish species of Iran and their impacts 

on native fishes 

Hamid Reza Esmaeili and Azad Teimory 

Department of Biology, College of Sciences, Shiraz University, 

Shiraz 71454, Iran 

In an effort to enhance, restore or re-establish fishery resources, 

species have been moved across the globe and introduced in 

totally new environments. 

As in many countries of the word, many fish species have been 

introduced to Iran and some of them have helped boost production 

in composite fish culture and also control of malaria. Introduction of 

fish species to Iran, dates back to the 1920s when mosquito fish 

Gambusia was introduced as an anti malaria agent, and to the late 

1930s when Cyprinus carpio was introduced for aquaculture 

purposes. Since then, about 27 more fish species have been 

introduced to inland waters of Iran and many more have also been 

translocated. The exotic fish species of Iran belong to 8 orders and 

8 families (Cyprinidae, Percidae, Salmonidae, Acipenseridae, 

Anguilidae, Esocidae, Mugilidae and Poecilidae). The species are 

dominated by the Cypriniformes (10 species). Some of exotic 

fishes (ex. C. carpio, and Gambusia holbrooki) have already 

established breeding populations, some others are regularly 

stocked by fisheries department of Iran (ex. Hypophthalmichthys 

molitrix, Ctenopharyngodon idella) and few others are occasionally 

recorded from natural waters. Aquaculture, sport fishing, control of 

malaria, ornamental purpose, research activities, demonstration in 

national fairs and accidental introduction have been the main 

reasons for this introduction. Pseudarasbora parva has been 

introduced accidentally along with the exotic carps. Due to lack of 

previous data, it is difficult to estimate the impact of these fish 

introduction on the native aquatic organism. However, predation, 

competition with native fishes, genetic changes through the 

hybridization, habitats changes, introduction of parasites and 

diseases are the main impacts of introduced fish species. Beside 

the invasion of exotic fishes, other threats including habitat 

degradation, environmental pollution, population growth, intensive 

aquaculture, unusual methods of fishing (toxin, dynamite) and 

drought have affected the ichthyodiversity of Iran. We briefly 

discuss the issues related to the introduction of exotics and their 

possible impacts on native fish fauna with certain possible 

measures to protect ichthyodiversity. 

Raccoon (Procyon lotor) as an invasive species for north of 

Iran 

Azita Farashi 1 , Bagher Nezami 2 , Ehsan Hadipour 2,3 , Hossein 

Alinezhad 4 , Mohammad Dehdar Dargahi 5 , Najmeh Mahdianzadeh 

1 , Mansour Aliabadian 6 and Mohammad Kaboli 1 

1 Department of Fishery and Environment, Faculty of Natural 

Resources, University of Tehran, Tehran, Iran; 2 Islamic Azad 

University, Science and Technology Branch, Tehran, Iran; 

3 Department of Environment of Lahijan, Gilan province, Iran ; 

4 Department of Environment of Rasht, Gilan province, Iran; 

5 Environment Science, Islamic Azad University, Lahijan Branch, 

Iran; 6 Department of Biology, Faculty of Sciences, Ferdowsi 

University of Mashhad, Mashhad, Iran 

Raccoon (Procyon lotor), as an invasive mammal, has been 

reordered from several European and Asian countries. For first 

time in 1991, Raccoon has been reordered from Iran, at Lavandvil 

Wildlife Refuge (48° 52' 20" E, 38° 20' 45" N) near Iran-Azerbaijan 

border. Since then incident of this species has not taken into 

account seriously. Protection of this wildlife refuge by Iranian 

Department of Environment has created a safe and suitable region 

for breeding and dispersal of Raccoon into neighboring areas. In 

this study, we recorded and observed the exponential dispersal of 

Raccoon in Iran using direct observation, camera trapping, and 

filling questionnaires from local people. Results confirmed that 

Raccoon is already reached to about 250 Km far from its original 

introduction place, either by replacing other native species or by 

local people as pet. The first evidences of effect of this invasive 

species on local wildlife have been recorded on breeding birds 

including, 

- 91 - 

Common Pheasant (Phasianus colchicus), Pygmy Cormorant 

(Phalacrocorax pygmeus. 

It seems there is a significant relationship between Raccoon’s 

density and recreational zone in Lavandvil Wildlife Refuge; 

because garbages, which have left over from visitors, of this 

protected area serve as good sources of food. Another important 

factor, which has increased the population density as well as the 

distribution of this invasive species into new habitats, is lacking of 

any natural predators. Apart of beautiful appearance which makes 

a pet animal of this species among local people, informing local 

people of consequences of distributing this species needs a 

serious programming. 

Early phases of a successful invasion: mitochondrial 

phylogeography of the common genet (Genetta genetta) 

within the Mediterranean Basin 

Philippe Gaubert, José-Antonio Godoy, Irene Del Cerro and 

Francisco Palomares 

UR IRD 131, DMPA – MNHN 43 rue Cuvier 75005 Paris France 

The Mediterranean Basin, connected by cultural exchanges since 

prehistoric times, provides an outstanding framework to study 

species translocations. We address the early phases of the 

invasion of the common genet (Genetta genetta), a small 

carnivoran introduced from Africa to Europe during historical times, 

by assessing mitochondrial nucleotide variability in 134 individuals 

from its native and invasive ranges. We propose that the cooccurrence 

in Maghreb of two divergent mitochondrial lineages 

(autochthonous and western African) is du! e to secondary contact 

through intermittent permeability of the Saharan belt during the 

Plio-Pleistocene. Estimates of coalescence time and genetic 

diversity, in concert with other available evidences in the literature, 

indicate that the origin of European populations of common genets 

is in Maghreb, possibly restricted to northern Algeria. The 

autochthonous mitochondrial lineage of Maghreb was the only 

contributor to the European pool, suggesting that translocations 

were associated to a cultural constraint such as a local use of the 

species, which might have artificially excluded the western African 

lineage. Haplotype network and Nested Clade Analysis provide 

evidence for independent events of introductions throughout Spain 

(Andalucia, Cataluña, and the Balearic Isl.), acting as a 

‘translocation hotspot’. Our demographic analyses support a 

scenario of very recent introduction of a reduced number of 

individuals in Europe followed by rapid population expansion. We 

sugg! est that an exceptional combination of factors including 

multiple translocations, human-driven propagation across natural 

barriers, and natural processes of colonization allowed by a wide 

ecological tolerance, promoted the successful spread of the 

common genet into Europe.


Mammal invasions in Europe: overview of patterns, trends, 

impacts and possible responses 

Piero Genovesi 

Istituto Nazionale per la Fauna Selvatica, Ozzano Emilia (BO), 

Italy 

The recently completed DAISIE program, co-funded by the 

European Commission within the 6 th Research Programme, has 

implemented the most updated and comprehensive inventory on 

alien species, storing data on over 10,000 alien species and 

45,000 introduction events recorded in Europe. 

As far as mammals are regarded, we recorded the presence of 

around 88 alien species introduced in the different areas of Europe 

after 1500, and of 397 events of introductions. 

Alien mammals account for about 15% of the total mammal fauna 

of Europe, and this proportion arises to over 20% if we only 

consider terrestrial species. Mammal invasions are increasing at 

very high speed, and the rate of new arrivals per year has passed 

from 0.03 estimated before 1800, to 1 new specie per year 

recorded in recent years. 

An analysis of the pathways and vectors of mammal introduction 

shows the increasing role of intentional import when compared to 

accidental transport of species. In recent years several 

introductions regarded pet species, confirming an increasing 

importance of pet trade as a vector of invasions. 

Among alien species, mammals are probably the most harmful 

taxonomic group; introduced mammals cause in fact severe 

impacts to biological diversity, are responsible for many extinctions, 

but also significantly affect agriculture, forestry and other economic 

activities. It is therefore urgent to improve our ability to prevent and 

respond to mammal invasions. On the basis of the reviewed 

information I provide some priorities to address the threats posed 

by alien mammals in Europe. 

Parasitism and biotic interactions: the cost of infection goes 

beyond the individual level 

Vincent Médoc, Jean-Claude Moreteau anf Jean-Nicolas Beisel 

Laboratoire des Interactions Ecotoxicologie, Biodiversité, 

Ecosystèmes (LIEBE), UMR CNRS 7146, Université Paul 

Verlaine-Metz, 57070 Metz-Borny, France 

Complex-life cycle parasites rely on the trophic pathway to reach 

the successive hosts. These so-called “manipulator” parasites are 

assumed to acquire the ability to modify various phenotypic traits 

in their intermediate hosts to make them vulnerable to the 

predation by definitive ones. Infection also affects how 

manipulated preys perceive the wide range of predators out of the 

parasite’s life cycle, expanding the cost of infection over the single 

host/parasite association. 

The freshwater amphipod Gammarus roeseli serves as an 

intermediate host for the bird parasite Polymorphus minutus. We 

investigated the cost of infection by P. minutus on the biotic 

interactions that occur between G. roeseli and two non-hosts, the 

invasive amphipod Dikerogammarus villosus, and the native 

stickleback Gasterosteus aculeatus. Infected amphipods 

interacting with D. villosus showed a prompt escape and then 

sheltered close to the water surface whereas uninfected 

individuals remained benthic. In water scented by sticklebacks, 

both prey types showed an increased refuge usage, but the 

response was exacerbated in infected preys. Consequently, in 

microcosm experiments, infected G. roeseli became out of reach, 

or at least less attractive, to the two predatory species, and thus 

were less preyed upon than uninfected specimens suffering from 

an increased predation pressure. From the parasite’s perspective, 

the induced alterations could be of selective importance in 

preventing the dead-end inside non-hosts of P. minutus. Beyond 

host/parasite association, these results suggest that infection can 

affect community dynamic through a shift in energy allocation, 

modifying the availability of certain prey types according to their 

infection status. 

- 92 - 

Invertebrate invasions in Europe: patterns and trends 

Alain Roques 

INRA UR0633, Zoologie Forestière, Orléans, France 

Within the framework of the European project DAISIE (Delivering 

Alien Invasive Species Inventory in Europe), a first continental 

inventory of the terrestrial invertebrate species alien to Europe has 

been realized. This compilation revealed that 1501 exotic species 

at least have right now established on the continent, and for the 

majority they arrived accidentally. A significant acceleration of the 

establishment process was observed since the 1990s, probably 

due to globalization, During this period, the contribution of Asia 

became gradually prevalent compared to North America. About a 

half of the alien species have phytophagous habits and the trade 

of ornamental plants, in all its forms (cut flowers, plants for planting, 

seeds, bonsais,…) appeared to be the dominant pathway of arrival 

in Europe. However, more than 60% of these species remained for 

the moment confined to man-made habitats such as gardens and 

parks, urban areas and cultivated fields whilst natural and seminatural 

habitats such as forests remain little colonized. Whether 

this situation reflects a lower resistance of man-made habitats to 

invasion or the alien populations arriving in these habitats need an 

accumulation phase before spreading in natural habitats remains 

still unclear. Insects largely dominate the invertebrate species alien 

to Europe, with a majority of Coleoptera and Homoptera. A 

comparison of the relative representation of the orders and families 

of insects among the entomofauna native and alien to Europe 

indicated that Homoptera, especially aphids and scales, are 

significantly more represented in the alien entomofauna; By 

contrats, Hymenoptera and Diptera are uner-represented. More 

generally, small-sized alien insects with sucking habits or 

developing in concealed habitats (seeds) established much more 

in Europe than large-sized external defoliators. These results 

suggest to define at taxa level and/or guild level groups at risk for 

future invasion. Other patterns possibly facilitating the introduction 

and establishment such as prolonged diapause, polyphagy, 

fecundity, … will be discussed during the conference. 

Widespread distribution of the invasive Artemia franciscana 

(Crustacea, Anostraca) in France: morphological, genetic and 

morphometric data 

Romain Scalone 1 , Mounia Amarouayache 2 and Nicolas Rabet 3 

1 Institut fur Spezielle Botanik und Botanischer Garten Johannes 

Gutenberg, Universitat Mainz Bentzelweg 9A 55099 Mainz. 

Germany. ; 2 Laboratoire Bioressources Marines, Université Badji 

Mokhtar, Annaba, Algeria; 3 UPMC, UMR 7138 CNRS UPMC 

MNHN IRD, Case 05, 7 quai St Bernard, F-75005 Paris, France 

New parthenogenetic and sexual populations of Artemia were 

found along the French Atlantic and Mediterranean coasts. In 

order to determine the specific status of these populations, we 

used referential populations of A. franciscana (from USA) and A. 

salina (from Algeria, Tunisia and Egypt). We present a genetic 

study using the variability of the developmental caudal gene in 

these populations (see Copf et al., 2003). In addition, a 

morphological investigations of the frontal knob and penis of the 

male specimens using the scanning electronic microscopy (SEM) 

was conducted to support the molecular data. The usefulness of a 

third morphological trait (the ratio of spine clusters on the frontal 

knob) for the Artemia systematics is discussed in the light of a 

statistical analysis using morphometric traits. 

This study reveals that all French sexual populations of Artemia 

belong to the species A. franciscana (Kellog, 1906) and not to A. 

salina (Linnaeus, 1758) accepted as the sexual native species in 

Mediterranean area. Recent publications on south of Europe and 

North of Africa (Amat et al., 2005; Mura et al., 2006) are congruent 

with our conclusions. The progressive replacement of the 

parthenogenetic and endemic Artemia species by a sexual and 

introduced species coming from North America, is fast and 

recently seems to be accelerated. The new French Artemia 

distribution is an additional example of an American invasive 

species probably introduced by human and decreasing the 

European biodiversity.


Spreading in France of the asiatic hornet, Vespa velutina 

nigrithorax : 2004-2007 distribution and invasion potentialities 

Claire Villemant 1 , Olivier Gargominy 2 , Quentin Rome 1 , Jean 

Haxaire 3 and Agnès Rortais 4 

1 MNHN Département Systématique et Evolution, UMR 5202 

CNRS, CP 50, 45 rue Buffon, F-75005 Paris, France ; 2 MNHN 

Département d'écologie et gestion de la biodiversité, Service du 

patrimoine naturel ; 3 « Le Roc », F-47310 Laplume ; 4 Université 

de Tours, Institut de Recherche sur la Biologie de l'Insecte (IRBI), 

UMR 6035 CNRS, Parc de Grandmont, F-37200 Tours, France ; 5 

Laboratoire Evolution, Génomes et Spéciation – LEGS, CNRS, 

UPR 9034, F-91198 Gif-sur-Yvette, France 

The accidental introduction of the Asian hornet Vespa velutina in 

south-west of France represents a new threat to biodiversity. This 

hornet, naturally distributed in mountainous regions from north 

India and China to Indonesia, is notably known as a fierce predator 

of pollinators, mainly honeybees (Abrol, 1994, Tan et al., 2007). In 

2004, the variety V. velutina nigrithorax was recorded, for the first 

time, in the Lot-et-Garonne department, and since then, it spread 

out very quickly across 17 departments throughout the Aquitaine 

region and its surroundings (Haxaire et al., 2006, Villemant et al., 

2006). As demonstrated by the rapid growth of colonies, the 

species adapted perfectly well to its new environment so that 

eradication is no longer possible. In its natural range of distribution, 

V. velutina occurs under various climatic conditions (continental, 

tropical and mountainous) and it feeds on a wide range of 

resources (insects and fruits). Therefore, further expansion is 

expected to occur, in the near future, to other European countries 

where the hornet can find suitable environmental conditions to 

develop. Thus, mapping the invasive hornet distribution proved to 

be essential to follow the wasp spreading and try to infer the 

modalities of expansion, in order to better perform control 

measures aiming at slowing down the invasion. We presented 

here the first V. velutina invasion maps, drawn from the 2004-2007 

nest records registered in the INPN database (Inventaire national 

du Patrimoine naturel, http://inpn.mnhn.fr) and inferred the 

spreading riks in Europe by using the Maxent model developed for 

modelling species geographic distributions (Phillips et al., 2006). 

A new enemy of honeybees in Europe: the invasive Asian 

hornet Vespa velutina 

Claire Villemant 1 , A. Perrard 1 , Quentin Rome 1 , Olivier Gargominy 2 , 

Jean Haxaire 3 , Eric Darrouzet 4 and Agnès Rortais 5 

1 MNHN Département Systématique et Evolution, UMR 5202 

CNRS, CP 50, 45 rue Buffon, F-75005 Paris, France ; 2 MNHN 

Département d'écologie et gestion de la biodiversité, Service du 

patrimoine naturel ; 3 « Le Roc », F-47310 Laplume ; 4 Université 

de Tours, Institut de Recherche sur la Biologie de l'Insecte (IRBI), 

UMR 6035 CNRS, Parc de Grandmont, F-37200 Tours, France ; 

5 Laboratoire Evolution, Génomes et Spéciation – LEGS, CNRS, 

UPR 9034, F-91198 Gif-sur-Yvette, France 

In 2004, the Asian hornet V. velutina nigrithorax was recorded, for 

the first time, in France in the Lot-et-Garonne department. Since 

then, the wasp spread out very quickly across 17 departments 

throughout the Aquitaine region and its surroundings. As 

demonstrated by the rapid growth of colonies, the species adapted 

perfectly well to its new environment so that its eradication is no 

longer possible. Further expansion is definitely expected to occur, 

in the near future, to other French departments as well as to other 

European countries where the hornet can find suitable 

environmental conditions (climate and resources) to develop. In its 

natural range of distribution, V. velutina occurs under various 

climatic conditions (continental, tropical, and mountainous) and 

feeds on a wide range of resources (various insects and fruits). 

Pollinators, mainly honeybees, being its main preys, the Asian 

hornet thus represents a new threat to European beekeeping 

activities. We presented here a short description of the hornet, its 

main biological features and the first invasion map in France, 

drawn from the 2004-2007 nest records registered in the INPN 

database (Inventaire national du Patrimoine naturel, 

http://inpn.mnhn.fr). 

- 93 - 

Mapping V. velutina distribution proved to be essential to follow the 

wasp invasion and to try to infer its modalities of expansion in 

order to perform control measures aiming at slowing down the 

invasion. 

Chemotaxis of the Pinewood Nematode, Bursaphelenchus 

xylophilus, to volatiles associated with Host Pine, Pinus 

massoniana, and its vector, Monochamus alternatus 

Lili Zhao and Jianghua Sun 

State Key Laboratory of Integrated Management of Pest Insects 

and Rodents, Institute of Zoology, Chinese Academy of Sciences, 

Beijing 100101, P. R. China 

The pinewood nematode (PWN), Bursaphelenchus xylophilus, the 

most important invasive species in pine forests of Asia, is 

transported to new pine hosts by vector beetles of the genus 

Monochamus. Third-stage dispersal juveniles (JIII) aggregate in 

pupal chambers around the vector as it matures. We demonstrate 

that the ratio of three terpenes (α-pinene and β-pinene and 

longifolene at 1:2.7:1.1) released by larval Monochamus alternatus 

strongly attract JIII while the ratio of these three terpenes (1:0.1:0.01) 

found in healthy xylem of Pinus massoniana attracts only the 

propagative stage (Jn) of the nematode. The results show that the 

nematode can distinguish specific ratios of terpenes and adjust 

behaviors for adaption to environmental change. The ratio of these 

semiochemicals may play an important role in nematode 

communication as it does in insects. We suggest that the volatiles 

produced by the host plants could be the basis of chemoecological 

relationship between plant parasitic nematodes and their vector 

insects. In addition, capture of JIII with terpene baited trap tubes 

deployed for 2hr in the field is demonstrated. Subsequently a new 

rapid sampling method using a baited trapping tube for attracting 

third-stage dispersal pinewood nematode juveniles (JIII) within two 

to six hours without injuring the pine tree or altering the wood 

product. This simple, effective and rapid sampling method is 

superior to the traditional laborious sampling by the Baermann 

funnel technique and has important implication at either ports-ofentry 

or field detection of this worldwide quarantine pest.


Organogenesis in vitro using undifferentiated cells and their 

molecular approaches in Vertebrate Development 

Makoto Asashima 

Graduate School of Arts and Sciences The University of Tokyo, 

ICORP Organ Regeneration Project, Japan Science and 

Technology Agency (JST), Japan 

In early development of animals, the prototype of formation is 

completed during the period from fertilization to the larval stage. 

Regular gene expressions and visceral formation, as well as other 

important phenomena involved in the features of embryonic 

uniformity, occur as an embryonic development program 

progresses. The degree of understanding of these phenomena in 

the terms of molecular language is a major problem in both 

developmental and cell biology. On this occasion, mechanisms 

underlying visceral formation, morphological formation, the 

structural development, and structural function become issues. 

Under these circumstances, visceral formation of 22 types of 

organs, such as the pancreas, kidney, eyeball, etc., using 

undifferentiated cells from frogs and the types of genes used for 

this formation via controlled expression, are outlined in the former 

half of this article. 

In the latter half of this article the possibility of visceral formation 

being achieved by using murine ES cells is described. Visceral 

formation from human stem cells is also described with reference 

to knowledge obtained by other investigators. The possibility of invitro 

visceral formation from murine ES cells has also become an 

issue. We have achieved various types of visceral formation, 

including those of nerves, smooth muscle, myocardium, the 

intestinal tract and fat cells, by using signal molecules such as 

activin, RAR and RXR. In connection with this, concentrations of 

activin and RA and changes in differentiation morphology and 

function are also discussed. The structure and function of these 

patterns of in-vitro visceral formation have become important 

issues. 

Embryonic and larval development of garpike Belone belone 

(Linnaeus, 1761) 

Jakov Dulčić, Branimir Baždarić, Leon Grubišić and Pero Tutman 

Institute of Oceanography and Fisheries, Split, POB 500, 21000 

Split, Croatia 

Eggs of garpike Belone belone (L.) were fertilized artificially and 

incubated under natural temperature and salinity conditions in the 

laboratory. Peculiarities in egg morphometry and in the course of 

embryogenesis of the Adriatic garpike were studied. The eggs, 

measuring 3.07±0.75 mm in diameter are equipped in viscous 

filamentous processes. Yolk-sac is homogeneous and does not 

contain oil globules. The embryonic development takes 330 D 0 (at 

temperatures varied between 19.4 0 C and 22.3 0 C). The Adriatic 

garpike embryonic mortality was high and exceeds 90%. The 

larvae carry a residual yolk sac and are fully prepared to live 

independently in water and to feed efficiently. The newly hatched, 

fully pigmented larvae measured 10.50±0.35 mm and were 

equipped with a residual 1.96 mm 3 yolk sac. Yolk sac was 

absorbed 17-48 h after hatching and immediately thereafter the 

larvae catches air bubble in purpose of making swimming bladder. 

A peculiar detail of the garpike embryogenesis is a certain 

extension of the embryonic development at the expense of the 

larval stage duration. 

S23 - Comparative organogenesis in Animals 

- 94 - 

Muscular developmental asynchrony during the early growth 

phase in the king penguin chick 

Aude Erbrech, Nathalie Guérin, Delphine Verrier, Catheline 

Gaudin, René Groscolas and Jean-Patrice Robin 

Institut Pluridisciplinaire Hubert Curien, Département Ecologie 

Physiologie Ethologie, UMR 7178 CNRS/ULP, associé à 

l’Université Henri Poincaré, Nancy 1, 23 rue Becquerel, 67087 

Strasbourg Cedex 2, France 

Continuous growth and steady parental food supply is a general 

trend in the rearing of young. However, in some species, food 

supply may be reduced seasonally for prolonged periods. Little is 

known about the impact of this energy constraint on the obligatory 

maturation of tissues. This was investigated in king penguin chicks 

which are reared by parents for one year. We compare the 

development of the pelvic and pectoral muscles in early growth, 

before the transitory growth arrest induced by winter food 

restriction. Data, obtained from freshly predated birds with 

developmental indices similar to those of free-living ones, were 

compared with adults. Muscle mass increased more slowly than 

total body mass in the pectoral girdle (P


Comparative studies on egg capsules ultrastructure in two 

species of Tardigrada: Dactylobiotus dispar Murray, 1907 and 

Isohypsibius granulifer Thulin, 1928. 

Izabela Poprawa and Magdalena Rost-Roszkowska 

Department of Animal Histology and Embryology, University of 

Silesia, Bankowa 9, 40-007 Katowice, Poland 

The eggs of Dactylobiotus dispar and Isohypsibius granulifer, 

similar to other Tardigrada eggs (Węglarska 1982), are covered 

with two shells: the vitelline envelope and the chorion. In both 

cases the vitelline envelope is very thin but the chorion has a 

multilayered structure. In both analyzed species the chorion is 

composed of 3 layers: (1) the inner layer of medium electron 

density; (2) middle, labyrinthine layer; (3) the outer layer of 

medium electron density. Differences between those two species 

are connected with all layers thickness and the structure of 

chorion’s surface. 

The labyrinthine layer of D. dispar is thinner than the same layer in 

the chorion of I. granulifer whereas the outer layer is thicker than 

the outer one of I. granulifer. The chorion surface of D. dispar is 

covered with small, conical processes with hooked tips. Such 

processes are not observed on the surface of L. granulifer eggs. 

Both egg capsules do not possess micropylar opening. 

Węglarska B. (1982) Ultrastructural study of the formation of egg 

envelops in Macrobiotus richtersi (Eutardigrada). In. Nelson D.R. 

(ed.) Proceedings of the third international symposium on the 

Tardigrada. East Tennessee State University Press, pp. 115-128. 

Extracellular vacuoles in developing midgut epithelium of 

Melasoma saliceti (Chrysomelidae, Coleoptera) embryos 

Magdalena Rost-Roszkowska and Jerzy Klag 

Department of Animal Histology and Embryology, University of 


Embryonic development of Melasoma saliceti lasts 7 days in 22 0 C. 

Stomodaeum and proctodaeum invaginate at the anterior and 

posterior ends of the germ band. Together with the ectodermal 

tissue the endoderm cells also enter into the embryo. The 

endoderm cells start to migrate over the yolk mass as two lateral 

bands of cells. On the fifth day of development the endoderm cells 

spread over the ventral and dorsal sides of the yolk and form the 

single layered primordium of the midgut epithelium whose cells 

become connected by specialised intercellular junctions of zonula 

adherens type. At the same time, between the cells, the 

intercellular spaces widen in some places. Initially the wide 

intercellular spaces have irregular outlines, but later they widen, 

acquire spherical shapes and transform into peculiar extracellular 

vacuoles. Each vacuole is enclosed within a band of junctional 

complex. The complexes are formed by zonula adherens, septate 

and gap junctions. On the sixth day of development the 

endodermal epithelium cells change shape from flat to cuboidal 

and eventually into columnar. At the same time, the vacuoles grow 

and become filled with electron dense content. The cell 

membranes enclosing the vacuoles form long, densely spaced 

microvilli eventually filling the vacuole completely. At the beginning 

of seventh day of development the apical surfaces of the 

embryonic midgut cells are completely smooth but the extracellular 

vacuoles come close to the midgut surface. At the end of the 

seventh day, shortly before hatching the extracellular vacuoles 

open to the midgut lumen and the microvilli spread over the apical 

surfaces of the midgut epithelium. Thus in very short time larval 

midgut, completely covered with fully formed brush-border is 

formed. The epithelial cells remain connected by zonulae 

adherentes continuous junctions and gap junctions. The septate 

junctions are seen only sporadically. The above-presented way of 

embryonic “preformation” of larval midgut microvilli in a 

holometobolous insect has not been described. Similar 

phenomenon of microvilli preformation was observed during 

differentiation of regenerative cells in some insect species. 

- 95 - 

Fine structure of the midgut epithelium of two species 

belonging to Archaeognatha Lepismachilis notata and 

Machilis hrabei (Insecta) 

Magdalena Rost-Roszkowska 1 and Jitka Vilimova 2 

1 Department of Animal Histology and Embryology, University of 


2 Charles University, Faculty of Science, Department of Zoology, 

128 44 Praha 2, Vinicna 7, Czech Republic 

Two primitive wingless insect taxa Archaeognatha and Zygentoma 

have originally formed one order Thysanura. Later, they were 

distinguished as two distinct orders Archaeognatha and 

Zygentoma, the basal taxa of Insecta. Recently some authors have 

suggested a recovery of the order Thysanura, thus the 

classification and relationship of Archaeognatha and Zygentoma 

are not still completely resolved. We analyzed the midgut 

epithelium and processes of its degeneration and regeneration in 

Thermobia domestica and Lepisma saccharina, which belong to 

Zygentoma (Rost 2006). In our present studies we compare the 

ultrastructure of the midgut epithelium in Lepismachilis notata and 

Machilis hrabei (Archeognatha) with those of Zygentoma. 

Midgut epithelium of both archaeognathans Lepismachilis notata 

and Machilis hrabei is composed of columnar epithelial cells and 

regenerative cells, which form regenerative cell groups. In both 

analyzed species the midgut epithelium resembles that of T. 

domestica and L. saccharina (Zygentoma). The cytoplasm of the 

epithelial cells shows distinct regionalization in arrangement of 

organelles, so basal, perinuclear and apical regions are 

distinguished. The cytoplasm of the regenerative cells is poor in 

organelles. They are capable of intensive proliferation and 

differentiation in epithelial ones. The first sign of their differentiation 

is appearance of cisterns of RER and SER, and numerous 

mitochondria in their cytoplasm. 

Rost M.M. 2006. Comparative studies on regeneration of the 

midgut epithelium in Lepisma saccharina L. and Thermobia 

domestica Packard (Insecta; Zygentoma). Annals of Entomological 

Society of America 99: 910-916. 

Molecular construction of tissues and organs of the Ciona 

intestinalis larva 

Nori Satoh 

Department of Zoology, Graduate School of Science, Kyoto 

University, Kyoto 606-8502, Japan 

Ascidians, or sea squirts, are sessile marine invertebrate 

chordates. Although the adult is a filter feeder, the fertilized egg 

develops quickly into so-called tadpole larva. The larva is 

composed of only ~2600 cells, which constitute a small number of 

organs including epidermis, central nervous system (CNS), 

endoderm and mesenchyme in the trunk, and notochord and 

muscle in the tail. This represents the most simplified chordate 

body plan. Specification of ascidian embryonic cells takes place 

very early at the cleavage stage; developmental fates of epidermis, 

CNS, endoderm, notochord and muscle are restricted by the 110cell 

stage. Together with well-characterized lineage of embryonic 

cells, the ascidian embryo may provide an appropriate 

experimental system to explore genome-widely the genetic 

cascades underlying embryonic cell specification. The draft 

genome of Ciona intestinalis has been decoded in 2002, and its 

~120 Mbp euchromatic genome contains ~16,000 protein-coding 

genes. Among these is a fundamental set of conserved chordate 

genes encoding transcriptional factors and cell signaling molecules. 

A thorough examination of Ciona gene expression (the 

transcriptome) is ongoing, including large-scale EST analyses 

(~700,000 ESTs), cDNA sequencing, in situ hybridization, and 

microarray analyses. All of the information is now available with a 

new Web site of our lab. Here we wish to discuss our current 

approach using transgenic lines and microarray to disclose genes 

involved in the formation of the notochord, central nervous system, 

and muscle of the Ciona larva.


Organogenesis of digestive organs in vertebrates 

Sadao Yasugi 

Faculty of Pharmaceutical Science, Teikyo Heisei University, 2289, 

Uruido, Ichihara, Chiba 290-0193, Japan 

The embryonic digestive organs in the vertebrates consist of 

endodermal epithelium and mesodermal mesenchyme. The former 

later fulfils functions such as production of digestive enzymes and 

absorption of nutrients, whereas the latter differentiates into 

connective tissue layers and smooth muscle layers. It has been 

well established that interactions between epithelium and 

mesenchyme are prerequisite for the normal development of the 

digestive organs. In general, the differentiation of the epithelium 

requires specific influence of the mesenchyme, and the 

differentiation of the mesenchyme and its nervous system is 

regulated by the epithelium. 

Recent studies carried out in my laboratory have revealed 

importance of some factors in the mesenchymal and epithelial 

tissues in the development of the chicken digestive organs, 

especially the stomach. For example, the determination of regional 

specificities of the epithelium in the primitive gut is often under the 

influence of the mesenchymal growth factors such as BMP, FGF 

and Wnt. On the other hand, the differentiation of the mesenchyme 

is controlled by sonic hedgehog expressed in the epithelium. 

The digestive organs possess essentially the same structures 

among the vertebrates. For example the stomach has glands cells 

of which produce and secrete digestive enzyme pepsin. The small 

intestine forms villus structures and epithelial cells produce 

dissacharidase and absorb nutrients. This means that factors 

acting in the development of the chicken digestive organs are also 

active in other vertebrates. In fact, above-mentioned growth actors 

and sonic hedgehog are known to be expressed in the developing 

stomach epithelium and mesenchyme of the mammals. Thus roles 

of these factors elucidated with he chicken embryo must also be 

important in other vertebrates. 

- 96 -


Against Weismann: transformism in French marine stations 

(1872-1914) 

Josquin Debaz 

S24 - Lamarck’s Philosophie Zoologique: 200 years 

EHESS – GSPR, 131 boulevard Saint-Michel, 75005 Paris, France 

At the end of the 19th Century, a whole generation of French 

biologists discovered the transformists thesis with Haeckel's work. 

It should be noticed however that the great figures of this 

generation were concurrently the actors of the development of 

marine stations. For scientists as Perrier, Giard, Delage, Dantec, 

Cuénot, the writings of Darwin and Lamarck were going hand in 

hand with a new vision of their discipline. All these young scientists 

were trained under Henri de Lacaze-Duthiers figure and its 

methodological school, which, if they were not opposed to Darwin 

as a scientist, did not intend to recognise his theories, outside any 

experimental process. Though, It was less the skepticism of their 

master than a conception of the discipline which was inherited, and 

therefore the goal to anchor transformism in an experimental 

practice. 

With the arrival of neo-darwinist thesis, the separation of two 

designs of heredity reactivate the older debate between epigenesis 

and preformation. Those french biologist, as epigenetists, felt 

strongly reluctant to Weismann's work. This opposition was often 

studied like a fight about inheritance of acquired characters, 

without pointing the opposition between two strong reductionisms, 

one based on embryo structures, the other on the indissociable 

unity of zygote. Marine invertebrates embryology studies, lead in 

these marine stations, were central in the debate, even if the two 

camps gave opposite readings of them. 

The general denomination of this group as 'neo-lamarckian' hide 

the great heterogeneity in their positions. Hence, the re-discovery 

of Mendel's laws had raised these differences. Crystallising the 

group against an interpretation of this event which drive biology 

towards the direction they reject. The reaction that can be noticed 

in the periodicals, in particular in the scientific Bulletin, was to 

widen the field, to publish every different voices, sometimes in 

order to give them a higher critic. 

Creationist conceptions of teachers (Primary and Secondary 

schools teaching biology or language) across nineteen 

countries 

Pierre Clément 1 and Marie Pierre Quessada 2 

1 LEPS-LIRDHIST, Université Lyon 1, France 

2 IUFM & Université Montpellier 2, France 

Because anti-evolutionist organizations are militating, teaching 

evolution at school is currently under contest in several countries. 

However no study across many countries was currently available 

to analyse the eventual creationist ideas among teachers dealing 

with Life and Evolution. Investigating the conceptions about this 

topic of 7,050 in-service and pre-service teachers from 19 

countries (Europe, Africa and Middle East), we have found 

significant proportions of creationist conceptions, and a wide 

variation of their amount across countries. The frequency of radical 

creationist conceptions is more related to the national economical 

level, to the personal degree of believing in God and practising 

religion, to the teacher's educational level and to the importance of 

teaching Evolution at school, than belonging to a particular 

religious group (e.g. Muslim or Christian; Catholic. Protestant or 

Orthodox). 

The data exposed here are coming from 19 countries, in the 

context of the European research project BIOHEAD-Citizen 

(Biology, Health and Environmental Education for better 

Citizenship: STREPS FP6, Priority 7, n° 506015). We thank all the 

colleagues who worked in each country, with the following team 

leaders: Farida Khammar (Algeria), Ivette Béré Yoda (Burkina 

Faso), Nicos Valanides (Cyprus), Tago Sarapuu (Estonia), Anna- 

Lisa Rauma (Finland), Franz Bogner (Germany). Attila Varga 

(Hungary), Adriana Valente (Italy), Iman Khalil (Lebanon), Jurga 

Turcinaviciene (Lithuania), Paul Pace (Malta), Sabah Selmaoui 

- 97 - 

(Morocco), Elwira Samonek-Miciuk (Poland), Graça Carvalho 

(Portugazl), Adrienne Kozan (Romania), Mame Seyni Thiaw 

(Senegal), Mondher Abrougui (Tunisia), Stephen Tomkins (UK). 

We also thank François Munoz and Charline Laurent (team of P. 

Clément) for the statistical analyses. 

Phenotypic Plasticity: a modern Avatar of Lamarckian thought 

Jean R. David 

Lab. Evolution, Génomes, Spéciation, CNRS1198 Gif sur Yvette, 

France, and Muséum National d’Histoire Naturelle, Département 

de Systématique et Evolution. Paris, France 

In his ‘Philosophie Zoologique’ (1809), Lamarck proposed that the 

diversity of living beings was not the result of a divine creation but 

the consequence of a natural process of progressive changes over 

time : this is now called biological evolution. Lamarck idea was 

obviously right, but changing over generations with modifications 

required an explanation of hereditary transmission. Observing that 

phenotypes could be modified by the environment, often in an 

adaptive direction, he proposed that acquired characters should be 

heritable, at least under some conditions. A century later, the 

rediscovery of Mendelian heredity proved that acquired characters 

were not heritable and, as a consequence, the influence of 

Lamarck ideas was more or less forgotten. However, ,the 

phenotypic variability in some groups may be spectacular, for 

instance in insects like Aphids, while the genome is not modified. 

Some researchers remained, however, interested in such 

variations, and their study now corresponds to a recognized field of 

modern biology, that is phenotypic plasticity. The challenge is to 

understand how this plasticity, which is basically non-heritable, can 

nonetheless be seen by natural selection, so that phenotypic 

plasticity itself may exhibit genetic changes overtime between 

populations or species. A classical approach is the Reaction Norm, 

that is the response curve of a phenotype along an environmental 

gradient. Examples of the diversity of reaction norms will be given, 

mostly using the Drosophila model, and showing that in many 

cases plasticity has an adaptive significance. Plasticity is an 

interaction between the environment and the genome, which by 

itself may be selected. For the moment, the genetic mechanisms 

of such interactions remain however to be worked out. 

Lamarck and Bergson on Progressive Evolution. No need for 

an "intelligent designer". 

Francis Dov Por 

The Hebrew University Jerusalem, Israel 

The central progressivist evolutionary theme of Lamarck, is of 

equal importance for the zoological philosophers of today. For 

Lamarck, progress is the result of the forces of nature, seen as an 

inexorable physico-chemical agent. It is not a linear "Scala 

Naturae", but a branching evolutionary tree which strives upward 

along many ramifications and the primate-human line is only the 

most advanced of the different shoots. 

Bergson ,in "Evolution Creatrice" which completes its centenary, 

saw progressive evolution as irreversible, neither mechanistic, nor 

teleological. His "élan vital" is seen by him as a natural force ,yet to 

be discovered ,like gravity in its time. 

Co-founders of the "modern synthesis", like Dobzhansky and 

Huxley, accepted zoological progress without however proposing 

any specific mechanism for it. Suspicions of finalism , theology and 

of anthropocentrism, led to the presently dominant ultradarwinistic 

.dogma .The field has been left open to a pervasive 

theological progressivism. 

Building of ever higher energy hungry animals and of more energy 

efficient ecosystems was emphasized by Lotka (1922) and 

recently by Vermeij (2004). Chaisson (2001) sees an evolution of 

more complex and more energy dissipating organisms in the open 

system of the globe. MEP, the law of maximum entropy production 

(Kleidon and Lorenz,2005) is the drive behind animal progress 

throughout geological history and the physical expression of 

Lamarck's "force de la nature" and Bergson's "élan vital".


It can be attributed to a demiurgic Ềtre Supreme, like Lamarck, son 

of the Revolution did, but this is not necessary for explaining 

animal progress . 

1) Chaisson, E. 2001, Closmic Evolution. The Rise of Complexity 

in Nature. Harvard University Press 

2) Keidon, A. and Lorenz, R.D. 2005. Non-equilibrium 

Thermodynamics and the Production of Rntropy. Life,Earth and 

Beyond. Springer Verlag 

3) Lotka, A.J,1922.Contribution to the Energetics of 

Evolution.PNAS 8 p.147 

4) Verm eij, G. 2004. Nature: An Economic History. Princeton 

University Press 

D’Omalius d’Halloy, Lamarck’s nice student 

Marie-Claire Groessens-Van Dyck 

Morren Foundation, Catholic University of Louvain-la-Neuve, 

Belgium 

The well-known Belgian geologist J.J. d’Omalius d’Halloy has been 

an assiduous Lamarck’s student before to become an assiduous 

defender of the Lamarck’s idea of the transformation of species. In 

1848, he stood up particularly for this point of view in a lecture to 

the Belgian Academy of Sciences. In this lecture, he pointed up a 

phenomenon just observed by his famous colleague from 

university of Louvain, the zoologist P.J. Van Beneeden. He saw in 

it what will be call later a case of neoteny and immediately pointed 

out this phenomenon as a mechanism able to transform species. 

On his side, P.J. Van Beneden claimed with reason the 

misinterpretation of his observation. This paper shows how the 

geologist has very well interpreted the consequences of a 

phenomena he has understood in an absolutely wrong way ! 

Baldwin Effect and Phenotypic Plasticity. 

Pierre Jolivet 

67 Boulevard Soult, F-75012-Paris, France 

Introduction : Genes and environment are essentially linked in the 

production of the phenotype. Baldwin effect is supposed to be 

produced when a biological trait becomes innate as a result of first 

being learned. If a mutation increases fitness, it will tend to 

proliferate in the population. Baldwinian evolution should be able 

to reinforce or weaken a genetic trait. 

Methods: Lamarckian evolution can arise from purely Darwinian 

evolution. Our concept arose mostly from the observation of ant 

domatia in plants and of controversial phenomena as heredity of 

callosities and coaptations among animals. 

Discussion: In 1896, James Mark Baldwin proposed a theory, 

entitled « a new factor of evolution », which has been called later 

on the Baldwin Effect. It is, roughly speaking, the genetic 

assimilation of an acquired character, physical or mental ; it is the 

result of an interaction of evolution with learning by individuals over 

their lifetime. Baldwin is remembered today exclusively for this 

paper. Recently, Erika Crispo (2007) and Mary Jeanne Eberhard 

(2003) pointed out that the two related evolutionary theories 

pertaining to phenotypic plasticity, those of James Mark Baldwin 

and of Conrad Hal Waddington, differ. Both theories have been 

often confused, and the purpose here is to stick to Baldwin Effect, 

which despite the ideas expressed by Baldwin himself, is a sort of 

an effort to reconcile the, otherwise irreconciliable, lamarckism and 

darwinism. In effect, phenotypic plasticity could allow an individual 

to genetically absorb a partially successful somatic change, 

through mutations, which might otherwise be useless to the 

individual. The Baldwin Effect says that a biological trait becomes 

innate as a result of first being learned. In fact, it is a sequential 

process in which acquired characters, somations, become genetic 

characters. The learned response to an environment change 

evolves on a genetic basis. The effect has been always 

controversial, but there are instances, which cannot be explained 

except by turning to the Baldwin Effect. Besides several 

simulations on software seem to have successfully confirmed the 

correctness of the theories. 

- 98 - 

Conclusions : 112 years of interest in the Baldwin effect produced 

thousand of papers. Roughly speaking, it means a selection of 

genes which reinforces the genetic basis of a variant of a 

phenotype. Modern authors concentrate on the phenotype, not the 

genotype, as the central driving force of the evolution. Lamarck 

was not fully wrong after all. 

Plant/animal frontier (1780-1830) 

Denis Lamy 

Muséum national d’histoire naturelle, Département Systématique & 

Evolution, UMS CNRS 2700, Taxonomie et Collections, CP 39, 57 

rue Cuvier, F-75231 Paris Cedex 05, France 

The distribution of the beings following a scale or a line from the 

minerals to the plants then to the animals was currently accepted 

by the naturalists of the 18 th century. Lithophytes and zoophytes 

were considered as the intermediate between minerals and plants, 

plants and animals respectively. By the way the frontiers between 

the kingdoms were not well defined. By the end of the 18 th century, 

the development of the study of the more simple animals and 

plants led to the distinction between animals and plants. Rejecting 

the concept of zoophytes, J.B. Lamarck proposed two independent 

series; but he compared the organization and the development of 

the more imperfect organisms in the animal and plant kingdoms. 

J.J. Virey accepted a common origin, more or less aquatic 

organisms, to the two kingdoms, developing in two diametrically 

opposite lines. A few years after, in front of the difficulty to place 

microscopic organisms, Bory de Saint Vincent proposed a new 

regnum, named ‘Psychodiaires’, including the ‘ébauches’ of plants, 

animals and minerals. These propositions will be discussed, 

namely in terms of their impact on the elaboration of animal and 

plant classifications. 

Lamarck and the beginning of life 

Stéphane Tirard 

Centre François Viète, Université de Nantes, France 

In 1802, in his book, Recherches sur l’organisation des corps 

vivans… Lamarck presented, for the first time, his evolutionary 

theory, in which animal series begin with spontaneous generation. 

We want to study, in Lamarck’s works, the distinction between 

spontaneous generations and the notion of primordial beginning. 

Firstly, we will show that spontaneous generations are completely 

included in the general process of evolution and constitute a 

simple, but complete, model of the lamarckian mechanisms of the 

transformations of organisms. 

Secondly, it seems very important to study the analogy established 

by Lamarck between spontaneous generation and fecundation. 

Thirdly, we will examine how, in Lamarck’s works, the notion of 

perpetual beginning dominates primordial beginning.


The neo-selectionist theory of genome evolution 

Giorgio Bernardi 

Laboratory of Molecular Evolution, Stazione Zoologica Anton 

Dohrn, Villa Comunale, 80121 Naples, Italy 

The vertebrate genome is a mosaic of GC-poor and GC-rich 

isochores, megabase-sized DNA regions of fairly homogeneous 

base composition that differ in relative amounts, gene density, 

gene expression, replication timing and recombination frequency. 

At the emergence of warm-blooded vertebrates, the gene-rich, 

moderately GC-rich isochores of the cold-blooded ancestors 

underwent a GC increase, which was similar and conserved in 

mammals and birds. Neither the GC increase nor its conservation 

can be accounted for by the random fixation of neutral or nearly 

neutral single nucleotide changes (i.e., the vast majority of 

nucleotide substitutions), or by a biased gene conversion process 

occurring at random genome locations. Both phenomena can be 

explained, however, by the neo-selectionist theory of genome 

evolution which fully accepts Ohta’s nearly neutral view of point 

mutations, but proposes in addition (i) that the AT-biased 

mutational input present in vertebrates pushes some DNA regions 

below a certain GC threshold; (ii) that these lower GC levels cause 

regional changes in chromatin structure which lead to deleterious 

effects on replication and transcription; and (iii) that the carriers of 

these changes undergo negative (purifying) selection, the final 

result being a compositional conservation of the original isochore 

pattern in the surviving population. 

How molecules changed the vertebrate tree. 

Wilfried W. de Jong 

Dept. Biomolecular Chemistry, Radboud University, Nijmegen, The 

Netherlands 

Since Darwin’s times ideas about the evolutionary relationships 

between the major groups of vertebrates have continuously been 

revised. While the division of vertebrates into higher taxonomic 

units like classes, subclasses and orders was often quite obvious, 

resolving their successive radiations from a common ancestor 

remained a matter of much controversy. Proposed relationships 

greatly depended on the subjective weight given by individual 

investigators to specific morphological characters or 

paleontological data. As a result consensus was often lacking, and 

prevailing opinions were largely dictated by the most dominant and 

outspoken schools. In that sense the vertebrate tree itself evolved 

according to Darwinian principles. 

Initially, the advent of molecular approaches further increased 

controversies. Early protein and DNA sequences generally 

confirmed firmly established morphological groupings, but often 

produced conflicting or ambiguous results with regard to higher 

order relationships. Especially confusing have been the 

discrepancies between sequence analyses based on mitochondrial 

and nuclear genes. Informative examples will be presented of 

phylogenetic conflicts that at the time have seriously damaged the 

confidence in molecular data. However, the current availability of 

long DNA sequences from many different nuclear genes, 

combined with a better taxon sampling and more sophisticated 

methods of analysis, have gradually stabilized the vertebrate tree. 

Especially useful for distinguishing between alternative topologies 

have been the so-called « rare genomic changes » such as 

insertions/deletions and retroposons. It will be demonstrated how 

these characters can provide unambiguous and objective 

resolution, taking examples mainly from mammalian interordinal 

studies. 

During the past few years consensus has been reached about the 

major branching patterns of the vertebrate tree, which deviates in 

some important aspects from previous morphological opinions. 

The presentation will summarize the current robust achievements, 

with special attention to mammals, birds and reptiles. It will also 

emphasize the remaining uncertainties. These mostly concern 

persistent trichotomies, which probably reflect periods of rapid 

speciation, and may well turn out to be intrinsically unresolvable. 

S25 - A tribute to Darwin 

- 99 - 

Darwin and the Barnacles: Insights and dreadful blunders 

Jean S. Deutsch 

Université P et M Curie (Paris 6), UMR 7622 Biologie du 

Développement, Paris, France. 

Barnacles (Cirripedes) are without any doubt Charles Darwin’s 

favourite animals. He spent no more than eight years studying 

them. He wrote monographs on living and on fossil cirripedes in 

1851 and 1854. Darwin himself was ambiguous as in his 

autobiography he writes both that his work on cirripedes was “of 

considerable value” and that he made “dreadful blunders”. The 

master word of both, to my view, is ‘homology’. I will review his 

contributions to cirripedes’ biology on both sides, using our present 

knowledge. I will propose an interpretation of his “blunders” in the 

context of the structural concept of homology derived from 

Geoffroy Saint-Hilaire and Owen. 

A 2008 look at The origin of species 

Michel Morange 

Centre Cavaillès and IHPST, ENS, 45 rue d’Ulm, 75005 Paris, 

France 

The publication of The origin of species played a decisive role in 

the general acceptance of the fact of evolution, through the 

proposal by Darwin of a reasonable mechanism to account for it. 

Paradoxically, this mechanism – variation and selection – was not 

accepted by most of his contemporaries. I will underline the limits 

of Darwin’s work. Darwin accepted the inheritance of acquired 

characteristics, had a vague idea of the nature of competition 

between organisms, and no mechanism at hand to justify the 

reliable transmission of variations through generations. In his 

subsequent works, Darwin more or less successfully 

complemented his initial model. 

The present theory of evolution is only distantly related to the 

contributions of Darwin. But the meticulous way in which Darwin 

collected informations coming from very different disciplines, and 

how he cautiously assembled these data remain lessons of good 

scientific practice.


Evolution of eyes and visual tasks 

Dan-Eric Nilsson 

The Lund Vision Group, Lund University, Dept. of Cell and 

Organism Biology, Zoology Building, Helgonavagen 3, 22362 Lund, 

Sweden 

Vertebrates, cephalopods, and arthropods have prominent eyes 

that are involved in most of the behaviours these animals display. 

But it is not the eyes and brains themselves that provide the 

selective advantage driving evolution of their visual systems. 

Rather, it is the ability to perform visual tasks that drive the 

evolution of vision. Naturally, early stages in the evolution of these 

complex visual organs must have been simpler, and served fewer 

visual tasks. Hence, eye evolution is driven by a consecutive 

accumulation of visual tasks. Each task ads to the requirements on 

eye structure, making it gradually more complex. For these 

reasons, reconstructions of eye evolution should ideally be based 

on an understanding of the sequential addition of visual tasks. In 

particular, it is interesting to ask what the first visual tasks might 

have been, and what requirements these would have placed on 

the structure and function of early eyes. With this objective, we 

have investigated vision in a group of simple and phylogenetically 

basal animal, the box jellyfish. Behavioural experiments indicate 

that these animals use vision primarily for positioning in the habitat, 

and for negotiating obstacles. To serve these tasks, the eyes are 

tuned for low spatial frequencies and are colour blind. The findings 

indicate that low resolution is not just sufficient, but in fact 

desirable in early stages of eye evolution. The lecture will analyse 

early phases of eye evolution and identify visual tasks and neural 

mechanisms that were involved in the first imaging visual systems. 

- 100 -


Teaching zoology in Paris: from the accomplishments - and 

mistakes - of the past towards a realistic future 

Armand de Ricqlès 

UPMC P6, Osteohistologie UMR 7179, case courrier 19, 75252 

Paris cedex 05, France 

French Zoology had its most glorious decades from the mid 

Eignteenth century (Buffon's time) to the end of the nineteenth 

century (opening of the "Galerie de Zoologie" at the Paris Museum 

was in 1889).Since then, as a "mother science",the aims, practices 

and teaching of zoology have tended to be split and spread among 

several "daughter sciences", often causing critical "struggle for 

survival" in a restricted academic ecosystem. By the middle of the 

last century, fields such as embryology, cells and tissues biology, 

general biology, comparative anatomy, physiology, paleontology 

and, lately, genetics were well delineated disciplins in the french 

academic world, often anxious to be recognized as such and apart 

from zoology, and with their own teaching. 

Accordingly, to zoology sensu stricto were left the description and 

classification of animals, with the relevant specialized subdisciplines 

(entomology, herpetology, ichthyology...) and fields 

which still had not conquered autonomy and recognition of their 

own, such as marin biology, ecology, ethology, ecophysiology and, 

to some extent, evolutionary biology. This trend had massive 

negative consequences on the teaching of zoology by the nineteen 

sixties and, accordingly, on how zoology was valued by the 

students. Because the heavy official program in zoology at the 

University was a mandatory part of the curriculum required by the 

students preparing "les concours" (agregation) to become 

appointed teachers in the State national education system, the 

trend was to offer a more and more practical, standardized but 

dogmatic teaching. This teaching was very facts rich and well 

organized but not questions-inspiring, and far away from most "hot 

spots" of the current scientific endeavors. Above all, it almost 

completely lacked of historical backgrounds and of epistemological 

and critical dimensions (there were happy exceptions!). The 1968 

riots brought severe blows to this system, the structure of 

traditional Zoology as a well-established and respected field within 

Academia being one of the casualties. 

Since then, zoology has managed to survive in practice and in 

spirit, here and there, often within its various "daughter sciences", 

to the point that it is very difficult now to offer any meaningful and 

operational definition of what zoology is, or should be, and of what 

should not be recognized as zoology within the full realm of 

functional or comparative-evolutionary sciences dealing with 

animal life. 

If it is possible to learn from the past, what little will survive of 

zoology in the French Academic ecosystem will be rooted in what 

raises current questions and interest in the mind of students at the 

level of research. For instance, systematics was dogmatically 

taught sixty years ago as a definite corpus of frozen knowledge 

fixed forever (and boring to most). My generation has pushed it 

back on the bench of a living research endeavor and it is why 

many students love it now. Every part of zoology may experience 

similar rejuvenation. 

The Beagle in the 21st Century 

Motonori Hoshi 

The Open University of Japan, 2-11 Wakaba, Mihama-ku, Chiba 

261-8586, Japan 

Darwin and Wallace’s papers on natural selection were read in 

their absence at the Linnean Society of London meeting of 1st July 

1858 when the remarkable and revolutionary theories were given 

their first public exposure. It is believed that Darwin conceived the 

idea of natural selection during his five-year journey around the 

world as chronicled in his book, The Voyage of the H.M.S. Beagle 

(1839). 

S26 - Diversity in teaching Zoology 

- 101 - 

Through my personal experiences of teaching zoology in Japan for 

decades, I have become convincing that contemporary students of 

biology need the Beagle in the 21st Century to practice the never 

fading words, Study Nature Not Books. In this paper, I will discuss 

more details of the Beagle in the 21st Century and propose an 

international project to implement this dream. 

The future of Zoology undergraduate curriculum: a canadian 

perspective. 

Anthony Russell 

2500 University Drive NW, T2N 1N4, Calgary, Alberta, Canada 

In this presentation, I will explore the structure of the Zoology 

curriculum in the North American arena, using Canada as my 

primary focus of a curriculum built upon a course-credit, liberal arts 

background. I will compare this with other approaches to 

curriculum design in other areas of the world and contemplate the 

concept of what might reasonably be expected of Zoology 

graduates in the North American system. This will include 

considerations of curriculum constraint imposed by entry into 

Medicine, Veterinary Medicine, Dentistry and the like requiring at 

least two years of undergraduate training in a liberal arts program. 

I will summarize what progress has been made in curriculum 

design. I will conclude by exploring curriculum design changes in 

courses in which I teach, injecting inquiry-based and blended 

learning approaches, and the degree of acceptance of these 

changes by students. 

Show and tell: college style – a method to improve student 

engagement 

Mark Schlueter 

1000 University Center Lane, Lawrenceville, Georgia, 30043, 

Lawrenceville, USA 

Two major challenges that exist in college teaching today are: (1) 

getting students actively involved inside the classroom and (2) 

getting students to learn outside the classroom. One teaching 

method that has been very successful in overcoming both of these 

challenges is the “Show and Tell” method. In my zoology class, 

students are assigned a particular animal group from which they 

must choose a single species. Each student begins by searching 

the Internet for information and videos. Then, the student returns 

to the classroom and gives a 7-minute presentation. Each lecture 

class begins with 2 student presentations that serve to introduce 

the day’s topic. Each presentation begins with several short videos 

(60-90 seconds) that immediately engage the audience, followed 

by several slides of interesting facts. 

The presentation concludes with a question slide in which the 

audience must answer 1-2 focus questions about the animal. 

Survey data indicates a significant amount of zoology learning 

outside the classroom while researching the Internet for the 

presentation. Also, students indicate a high degree of satisfaction 

using the Internet to track down entertaining videos and 

information. Most students reported rewatching the videos and 

viewing related topics after class. This teaching method has 

additional benefits. Students develop presentation skills and use 

technology to research information. Students go beyond the 

textbook and get a better understanding of the biodiversity of the 

planet. Video clips provide a safe way to view dangerous and 

exotic animals. Finally, this method can upgrade passive lecture 

classes into active learning environments.


Kinorhynchs murdered by bachelors! should we care? 

Gerhard Scholtz 

Philippstr. 13, 10115, Berlin, Germany 

Kinorhyncha is a small group of minute bilaterians showing a 

hidden lifestyle. An ecological impact on humans is virtually absent. 

Hardly any layperson and even many biologists ever heard of 

these animals. Nevertheless, based on their anatomy they 

suddenly play an important role for our understanding of metazoan 

evolution, in particular the origin of segmentation. I use this animal 

group a! s an example to ask the question of whether it is 

important to have zoological knowledge. Germany has a strong 

tradition in morphology and comparative embryology of 

invertebrates. This field of research had always tight connections 

to phylogeny and evolution of the animal kingdom and produced 

numerous hypotheses and theories which are relevant for general 

biological questions. Teaching and thus the knowledge of these 

issues decreased dramatically during the last years. This is 

reflected in the low number of filled and advertised university 

positions dealing with morphology and comparative embryology. 

The introduction of the new Bologna curricula and degrees even 

enforces this development and might lead to an irreversible 

interruption of the morphological tradition. 

- 102 - 

Jeanne Villepreux-Power, the mother of aquariology 

Claude Vast 

Aquarium du Limousin, 87000 Limoges, France 

Jeanne Villepreux-Power was born in a French village (Juillac) in 

1794 and died in this city in 1871. 

She lived in the town of Messsina for 20 years after her marriage. 

She made the list of natural riches and colected plants, animals 

and fossils of the Sicilia Island. 

She devoted one’s self to experimental research about marine 

animals. 

She was first (1832) to create and use the aquarium in glass and. 

imerged cages for observation and experimentation in aquatic 

environments. She was famous all through Europe for her works 

on the Argonauta argo shell. She was recognized by Prof. Richard 

Owen (1804-1892) of British Museum as the "mother" of 

aquariology. 

She was, the first woman in the Catania Accademia and a 

correspondent member of the London Zoological Society (plus 16 

other academies). 

She authored « Observations et experiences physiques sur 

plusieurs animaux marins et terrestres » as well as several other 

papers. The « Guida per la Sicilia » is a brillant and erudite 

inventory of the Island environment.

ICZ2008 – Abstracts Addendum 

Author index 

- 103 -


Abdullah Mohammed 82 

Abiadh Awatef 3 

Aceros Henry 52 

Acheuk Fatma 79 

Ackermann Alain 65 

Adrien Jérôme 44 

Agosti Donat 23 

Agrawal Om Prakash 79 

Ahlberg P. 8 

Ait-Kaci Karima 79, 80 

Akhremenko Alexander K. 76 

Alberdi Maria Teresa 33 

Alcalde Gema Maria 33 

Aldana Elis 78 

Aliabadian Mansour 3, 64, 99, 100 

Alinezhad Hossein 100 

Alizadeh-Eghtedar Hanieh 12 

Allanic Yoann 68 

Almeida Mónica 74 

Alonso-Zarazaga Miguel 25 

Altenburger Andreas 41 

Altincicek Boran 48, 50 

Amarouayache Mounia 102 

Amaya-Perilla Catalina 71 

Ameziane Nadia 13, 67 

An Chuanguang 84 

Andersen Ann C. 84 

Andreoli Ilaria 31 

Andrés Maria 33 

Angelier Frédéric 71 

Angenard Gaelle 37 

Antonevich Anastasia 61 

Anufriev Andrey I. 73, 76, 77 

Arabi Juliette 3 

Aranda Manuel 44 

Archambeau Anne-Sophie 95 

Arendt Detlev 41 

Arndt Harmut 31 

Arnold Gérard 66, 82 

Asashima Makoto 103 

Asem Alireza 3, 4, 99 

Astudillo Fernandez Aina 68 

Aubin Thierry 53 

Auger Hélène 41 

Auguste Aurélie 33, 37 

Aulard Sylvie 15 

Avargues-Weber Aurore 56 

Azanza Beatriz 33 

Bach Jean-François 48 

Bachmann Lutz 4 

Baillet Adrienne 33 

Bailleul Frédéric 70 

Bakhtadze G. 4 

Bakhtadze N. 4 

Bakke Tor A. 4 

Balavoine Guillaume 41 

Balian Estelle V. 23 

Bandarra Narcisa 92 

Bang Alok 56 

Bapteste Eric 43 

Barberousse Anouk 12 

Bar-David Shirli 68 

Bardet Claire 43 

Barjadze, Shalva 4 

Barlow Axel 52 

Baroiller Jean-François 33, 35, 38 

Barske Julia 68 

Barske Lindsey 34 

Basova Larisa 61 

Batista Irineu 92 

Baylac Michel 14 

Baždarić Branimir 103 

Bech Claus 71 

Becker Carola 17 

Becker Pierre 89 

Behidj-Benyounes Nassima 69 

Beisel Jean-Nicolas 101 

Beja Pedro 74 

Belaid Messaouda 4 

Benaziza Djamila 4 

Bendifallah L. 4 

Bengtson Stefan 27 

Bengtsson Henrik 39 

Benken Konstantin A. 32 

Berkani Mohamed 84 

Berlémont Renaud 89 

Bernadou Abel 56 

Bernal Ximena 55 

Bernard Cécile 80 

Bernard-Samain Sylvie 33 

Bessa-Gomes Carmen 61 

Bhadauria Monika 79, 81 

Biagianti Sylvie 63 

Bigey Françoise P 29 

Birmani Nadir Ali 6 

Bisby Frank 23 

Blair Hedges S. 8 

Blaustein Leon 68 

Bleu Josefa 61 

Bobe Julien 33 

Bogart Jim 47 

Bogutskaya Nina G. 25 

Böhne Astrid 33 

Boisselier Marie-Catherine 5, 11, 14 

Boizet-Bonhoure Brigitte 38 

Boman Sanna 99 

Bonami Jean-Robert 94 

Bonfante-Cabarcas Rafael 78 

Bonillo Céline 5 

Bonnet Xavier 73 

Bonyadi Naeini Alieh 5 

Boos Mathieu 67 

Borchiellini Carole 44 

Borisov Zakhar Z. 76 

Boucher-Rodoni Renata 91 

Bouchet Philippe 11, 14, 23, 25 

Boudier Thomas 88 

Bouhadad Rachid 95 

Boumaïza Moncef 80 

Bounaceur Farid 99 

Bourdon Estelle 27 

Bouriga Nawzet 84, 92 

Boustila Farid 99 

Boutet Isabelle 85 

Bouvy Marc 85 

Branch William R. 8 

Brandis Dirk 17 

Bressac Christophe 18, 85 

Briefer Elodie 53 

Briggs Derek E.G. 27 

Brillard Jean-Pierre 36 

Brissac Terry 85 

Brochon Jean-Claude 81 

Brothers Denis J. 25 

Brunet Frédéric 33 

Buckeridge John St J. S. 79 

Budzinski Hélène 82 

Bullejos Monica 39 

Burghelea Carmen 42 

Bysykatova Inga P. 7 

Cabau Cédric 33 

Cabrera Arreola Andrea Alejandra 5 

Caisey Xavier 82 

Cambon-Bonavita Marie-Anne 86, 93 

Camerino Giovanna 34 

Capel Blanche 34 

Cariou Marie-Louise 14, 15, 19 

Caro Audrey 85 

Carré Gwenn-Aël 36 

Casadevall Margarida 53 

Casewell Nicholas R. 52 

Castanet Jacques 27 

Castelin Magalie 14 

Castellani Christopher 27 

Catapano Terry 23 

Causse René 81 

Céréghino Régis 56 

Chaboissier Marie-Christine 34 

Chakali Gahdab 69, 72 

Chakvetadze N. 4 

Chalghaf Mohamed 93 

Châline Nicolas 63 

- 104 - 

Chameron Stéphane 57, 65 

Chaminade Nicole 15 

Chang Muhammad Saleem 69 

Chardard Dominique 34, 40 

Charry Maria 86 

Chassot Anne-Amandine 34 

Chastel Olivier 71 

Chebbi Nadia 99 

Chebouti Yahia 69 

Chebouti-Meziou Nadjiba 69 

Chekunova Anna I. 75 

Chen Min 71, 100 

Chenin Eric 95 

Cheniti Tahar Lamine 3 

Chéron Blandine 56 

Chesnel Armand 34, 40 

Chetoui M’barek 3 

Cheval Justine 95 

Chevaldonne Pierre 13 

Chevallier Damien 69 

Chilmonczyk Stefan 33 

Chiron François 100 

Choné Thérèse 95 

Cibois Alice 5 

Čikeč Čulić Vedrana 42 

Čikeš Keč Vanja 34 

Ciofi Claudio 68 

Clément Pierre 109 

Clobert Jean 70 

Coffigny Hervé 37 

Cohen Jean 31 

Čokl Andrej 53 

Colangelo Paolo 3 

Colleye Orphal 53 

Collins Michael 75 

Combe Maud 57 

Compère Philippe 86, 89 

Conway Morris Simon 29 

Coolen Marion 42 

Cooper Edwin L. 48 

Cooper Max D. 48 

Corbari Laure 86 

Corbel Hélène 61 

Cordero Carlos 17 

Coriton Olivier 38 

Cornette Raphaël 67 

Coroiu Ioan 77 

Correia da Rocha-Filho Léo 62 

Cortés-Rodríguez Nandadevi 7 

Cotinot Corinne 33, 37 

Cottin Delphine 87 

Couloux Arnaud 33, 84 

Coutanceau Jean-Pierre 38 

Couty Isabelle 36 

Couvet Denis 95 

Coyne Jerry 14 

Crews David 40 

Cruaud Corinne 84 

Cubo Jorge 27 

Cunha Marina R. 92 

Curé Charlotte 53 

Curtis Nicholas E. 91 

D’Cotta Helena 33, 35 

D’Haese Cyrille 6 

D’Hont Angélique 38 

Da Lage Jean-Luc 14, 15 

Da Silva Corinne 42 

Daggag Hinda 39 

Daguin Claire 91 

Dahlgren Thomas G. 93 

Damen Wim G. M. 44 

Damiens Gautier 82 

Damuth John 71 

Danger Jean-Michel 79 

Dargahi Mohammad Dehdar 100 

Darrouzet Eric 102 

David Alin 77 

David Bruno 88, 95 

David Jean R. 106 

Dayrat Benoit 24 

de Groot Maarten 53


de Jong Wilfried W. 108 

de Ricqlès Armand 27, 109 

De Ridder Chantal 88, 89, 95 

de Rooij Dirk 34 

de Villardi de Montlaur C. 93 

Debat Vincent 42 

Debaz Josquin 106 

Debelle Allan 42 

Debruyne Régis 10 

Defaye Danielle 67, 68 

Degtyarev Victor G. 6, 7 

Del Cerro Irene 101 

Deneubourg Jean-Louis 57, 68 

Denis Françoise 72 

Denker Elsa 43 

Derégnaucourt Sébastien 54 

Derkacheva Maria E. 32 

Deshpande Sujata 56 

Detrain Claire 57 

Dettai Agnès 9, 13, 84 

Deutsch Jean S. 108 

Dharejo Ali Murtaza 6 

Díaz de la Guardia Quiles Rafael 39 

DiNapoli Leo 34 

Djanashvili R. 4 

Dobreff James 21 

Donnellan Steve 8 

Dos Santos Sandra 43 

Douady Stéphane 59 

Doumandji Salah Eddine 4, 69 

Doumandji-Mitiche Bahia 79, 80, 99 

Doums Claudie 56 

Dov Por Francis 106 

Dragon Anne-Cécile 70 

D'Souza Thomas 47 

Du Pasquier Louis 49 

Dubilier Nicole 87, 93 

Duboule Denis 1 

Dugnol Benjamin 54 

Dulčić Jakov 90, 103 

Dumond Hélène 34, 40 

Düngelhoef Susanne 19 

Duperron Sébastien 85, 87, 88, 93 

Duquenne Clotilde 36 

Durand Fabrice 79 

Durant Joël M. 65 

Durusel Jean David 39 

Eberhard William G. 17 

Edery Marc 80 

Egorov Nickolai N. 6, 7 

El Filali Adil 95 

El Jamil Anwar 38 

Eleaume Marc 13 

Elias Marianne 70 

Elisabeth Nathalie 88 

Erbrech Aude 103 

Ereskovsky Alexander 44 

Eroukhmanoff Fabrice 14 

Errouissi Faiek 96 

Esmaeili Hamid Reza 28, 100 

Espadaler Xavier 56 

Estrada Alba 97 

Evanno Magali 18 

Evenhuis Neal L. 24 

Evin Allowen 14 

Exbrayat Jean-Marie 36, 40 

Fajardo Gonzalo 17 

Falconi Rosanna 39 

Fang Hongxia 15 

Farashi Azita 100 

Fassatoui Chiheb 70 

Faure Eric 84 

Faure Grazyna 51 

Fautin Daphne G. 25 

Fazouane Fethia 79, 80 

Feitosa Natália M. 44 

Felbeck Horst 87 

Felder Darryl L. 8 

Fellous Marc 35 

Fénéron Renée 57 

Fernández Carlos 54 

Ferrantini Filippo 31 

Ferreira Eduardo 74 

Ferreira Ronara S. 59 

Flament Stéphane 40 

Fokin Sergei I. 31, 32 

Fonseca Carlos 72 

Fourcassié Vincent 56, 57 

Fowler Paul 35 

Frédérich Bruno 53 

Fresneau Dominique 58, 59 

Froschauer Alexander 33 

Fry Bryan 51 

Frydman René 37 

Frygelius Jessica 35 

Fu Daliang 74 

Fujikura Katsunori 91 

Fujiwara Yoshihiro 91 

Fusani Leonida 68 

Fuwa Naoto 46 

Gachal Ghulam Sarwar 11, 69 

Gadagkar Raghavendra 56 

Gahr Manfred 54 

Gaill Françoise 85, 86, 88, 89, 90, 93 

Galiana-Arnoux Delphine 33 

Galiano Gonzalo 54 

Galleni Moreno 89 

Gantsevich Mikhail 61 

Gao Chunling 30 

Garcia Manuel Ruiz 86 

García Emilio J. 74 

García-Deras Gabriela M. 7 

García-Moreno Jaime 7 

Gargominy Olivier 102 

Garms Andrew 49 

Garnier Simon 57 

Gaubert Philippe 101 

Gaudin Catheline 103 

Gaudron Sylvie 88 

Gazave Eve 44 

Geiger Sylvie 61 

Gelhaye Mathieu 34 

Genelt-Yanovsky Eugene 61 

Genovesi Piero 101 

Gerbier Grégory 57 

Gerlinskaya Ludila 78 

Germogenov Nickolai I. 6, 7, 77 

Gernigon-Spychalowicz Thérèse 36, 37, 40 

Gharred Tahar 80 

Ghelem Mohamed 69, 72 

Gilichinsky David 32 

Ginzburg Lev 71 

Glaubrecht Matthias 7 

Godfray Charles J. 24 

Godoy José-Antonio 101 

Gombault A. 42 

Gomez Maria 7 

Gompert Zach 70 

Goncharova Elena 49 

Gonzalez-Maya Jose F. 71 

Gordon Lavina 39 

Goutte Aurélie 71 

Govoroun Marina 36 

Goyffon Max 51 

Grant Jonathan 78 

Grapputo Alessandro 99 

Gratiashvili Nana 58 

Greenfield Michael D. 62 

Grégoire Elodie 34 

Groessens-Van Dyck Marie-Claire 106 

Gros Olivier 88, 89, 90, 91 

Groscolas René 61, 103 

Grubišić Leon 103 

Grygier Mark J. 25 

Guarini Jean-Marc 73 

Guéchot Sébastien 14 

Guendouz-Benrima Atika 99 

Guérécheau Aurélie 57 

Guérin Nathalie 103 

Guerquin Marie J. 36 

Guerra Ana 74 

Guibert Floriane 62 

- 105 - 

Guigen Yann 33 

Guinet Christophe 70 

Guizien Katell 73 

Guo Guang Qin 81 

Guo Guangpu 71 

Guo Ximing 15 

Gustave Dit Duflo Sylvie 88 

Guyot F. 93 

Habbi Assia 4 

Habert Damien 43 

Habert René 36, 37 

Habibi Saina 64 

Hackstein Johannes H.P. 31 

Hadipour Ehsan 100 

Hadj-Bekkouche Fatima 36, 39 

Haffner Patrick 68 

Hagiwara Teruki 46 

Halary Sébastien 85, 88, 93 

Halliday Bruce R. 25 

Hammouche Sadlia 36 

Hanken James 24 

Hansen Haakon 4 

Hardy Clotilde 88 

Hargeby Anders 14 

Harris Anjanette 63 

Hartfelder Klaus 59 

Hasan Hassan 72 

Hase Sumitaka 45 

Hassanin Alexandre 3 

Haxaire Jean 102 

Hayashi Katsuhiko 7 

Healy Susan D. 63 

Heinze Juergen 59 

Helmstetter Cécile 65 

Hemery Lenaig 13 

Hernández-Baños Blanca E. 7, 17 

Hersmus Remko 35 

Hill Naomi 68 

Hmida Leila 70 

Hoffmann Jules 1 

Hoke Kim 55 

Hormiga Gustavo 20 

Hörstgen-Schwark Gabriele 35 

Hoshi Motonori 47, 109 

Hotta Kohji 45 

Hou Lianhai 30 

Houdelier Cécilia 62 

Hourdez Stéphane 92 

Hoyoux Caroline 89 

Huber Bernhard A. 18 

Hubert Pauline 63 

Hubweber Lasse 18 

Ion Constantin 63 

Isayev Arkady P. 6 

Ishijima Yoshie 46 

Ishikawa Takayuki 7 

Ishizu Hirotsugu 47 

Jacquot Marie 8 

Jaisson Pierre 63 

Janussen Dorte 89 

Jaubert Francis 35 

Jeffery William R. 43 

Jennings Andrew P. 10 

Jiang Zhigang 15, 64 

Jiggins Chris 70 

Jiguet Frédéric 95 

Joger Ulrich 30, 43 

Johnson Norman F. 23, 24 

Johri Sonia 79 

Jolivet Pierre 107 

Jollivet Didier 91 

Joly Dominique 16, 18, 19 

Joly Jean-Stéphane 41 

Joshi Deepmala 79 

Jost Christian 57, 59 

Julliard Romain 63, 95 

Kaboli Mohammad 3, 64, 100 

Kandsi Faroudja 36 

Kanhoush Rasha 38 

Kano Shungo 41 

Kark Salit 100


Kaufmann Joshka 64 

Kawato Masaru 91 

Kazemi Noureini Sakine 43 

Keithmaleesatti Sarun 37 

Khan Muhammad Munif 6 

Khondkaryan Lusine 96 

Khosravani Azar 39 

Kim Yuna 34 

Kino Koji 7 

Kintsurashvili N. 4 

Kirchner Florient 68 

Kitana Noppadon 37, 40 

Klag Jerzy 104 

Knapp Sandra 25 

Ko Chia-I 34 

Kobayashi Kazuya 47 

Kocer Ayhan 33, 37, 38 

Kocher Thomas D. 38 

Koentges Georgy 8 

Köhler Frank 7 

Korb Judith 58 

Kotrschal Kurt 62 

Kottelat Maurice 25 

Kraljević Miro 90 

Krasnov Boris 96 

Krell Frank-Thorsten 25 

Krespi L. 78 

Kruck Sarah 44 

Krys Sophie 80 

Kuchta Shawn R. 15 

Kullander Sven 0. 25 

Kuntz Pascale 59 

Kuntz Sandra 34 

Kuntzelmann Elise 8 

Labidi Imen 96 

Labutin Yuri V. 76 

Lachaise Daniel 19 

Lachaud Jean-Paul 58 

Laghaei Fatemeh 28 

Lahaye Jean-Baptiste 36 

Lallier François H. 85, 91, 92 

Laloi David 61, 64 

Lamas Gerardo 25 

Lambrot Romain 37 

Lamy Denis 107 

Landstorfer Richard B. 8 

Lapébie Pascal 44 

Larsen Ole Næsbye 54 

Laudet Vincent 44 

Laurent Goulven 107 

Laurent Mélina 89 

Laurin Michel 29 

Lautredou Anne-Claire 84 

Lavick Gaute 91 

Le Bris Nadine 89, 90, 93 

Le Foll Frank 79 

Le Galliard Jean-François 64 

Le Guigo P. 78 

Le Lann Cécile 78 

Le Maho Yvon 65, 69 

Leboulenger François 79 

Lecointre Guillaume 9, 84 

Lecoutey Emmanuel 63 

Lécureuil Charlotte 37 

Ledreux Aurélie 80 

Lee YunSun 9 

Legendre Laurent 41 

Léger Nelly 87 

Legrand Delphine 14 

Leitão Albertine 54 

Lemeunier Françoise 15 

Lepoint Gilles 86, 89 

Lerch Alexandre 64 

Letendre Julie 79 

Lévêque Christian 23 

Li Blaise 9 

Li Chunwang 15, 64 

Li Na 81 

Li Pei Qiang 81 

Li Zhongqiu 15 

Lignot Jean-Hervé 65 

Lim Susan 25 

Lindström Leena 99 

Litvinova Ekaterina 49 

Liu Jinyuan 30 

Liu Xiao 15 

Liu Yuanmin 110 

Livera Gabriel 36 

Lizano Eliezer 78 

Llaneza Luis 54 

Lockridge Mueller Rachel 15 

Loeuille Nicolas 73 

Looijenga Leendert H.J. 35 

Lopes Peterson L. 44 

Lorion Julien 89 

Louadi Kamel 4 

Loublier Yves 82 

Lu Lin 78 

Lucas Christophe 58 

Lumineau Sophie 62 

Lyytinen Anne 99 

M’Hadhbi Lazhar 80 

Ma Qiang 81 

Maas Andreas 27 

Maatouk Danielle 34 

Machon Nathalie 75 

MacLeod Norman 29 

Magre Solange 38 

Mahar Mukhtiar 11 

Mahdian-zadeh Najmeh 100 

Maiocco Öna 9 

Maldonado Jairo 52 

Malécot Mélodie 80 

Malherbe Marie-Claire 57 

Mandon-Pépin Béatrice 33, 37 

Manjón-Cabeza María Eugenia 95 

Manuel Michael 43 

Mappes Johanna 99 

Marangoni Elsa 72 

Marchal Juan Antonio 39 

Marie Arul 80 

Marjanović David 29 

Markotić Anita 42 

Markov Nickolay 9 

Marques Sylvie 65 

Márquez Ana Luz 97 

Martens Koen 23 

Mary-Huard Tristan 16 

Massemin-Challet Sylvie 65, 69 

Mastrototaro Francesco 99 

Mathevon Nicolas 53 

Mathias Maria Luz 75 

Mathur Ramesh 79 

Matić-Skoko Sanja 90 

Matos Milena 72, 74 

Matsumoto Midori 47 

Matsuo Takashi 14 

Matsuoka T. 8 

Maurin Leslie C. 90 

Mawatari Shunsuke F. 25 

Mazan Sylvie 42 

McGredor Reid Gordon 21 

McGregor Alistair P. 44 

Mdelgi Esma 70 

Mebs Dietrich 51 

Mecelem Dalila 72 

Médoc Vincent 101 

Mehrandish Parisa 64 

Melo Gabriel A. R. 62 

Memon Nasreen 12 

Menad Rachid 37 

Ménez André 51 

Meng Qingjin 30 

Menuet Arnaud 42 

Merçot Hervé 85 

Mezhoud Karim 80 

Mhlanga-Mutangadura Tendai 19 

Michard-Picamelot Delphine 65 

Michel Catherine 73 

Michel Ellinor 25 

Mikaia Nona 73 

Miller Scott 26 

- 106 - 

Minelli Alessandro 9, 25 

Mira António P. 74, 75 

Miralles Aurélien 9 

Mirzanejad-asl Hafez 10 

Missaoui Hechmi 93, 99 

Mittal Deepak 79 

Modeo Letizia 31 

Moe Borge 71 

Mohand Kaci Hakima 80 

Moison Delphine 37 

Monnin Thibaud 56 

Montazer-Torbati Fatemeh 33, 37 

Montchamp-Moreau Catherine 16 

Monti Laurence 15 

Morales Jorge 33 

Morange Michel 108 

Mordosov Innokenti I. 76 

Mordosova Nadejda I. 73 

Moreteau Jean-Claude 101 

Moritz Charlotte 73 

Moriyama Tohru 66 

Moshkin Michail 49 

Möstl Erich 62 

Mouneyrac Catherine 82 

Moya Francina 95 

Muczynski Vincent 37 

Mukai Ruriko 46 

Muricy Guilherme 11 

Mustać Bosiljka 38 

Mylnikov Alexander 32 

Myslenkov Alexander I. 9 

Nagahama Yoshitaka 103 

Nagle Lucien 64 

Naidenko Sergey 61 

Nejlaoui Zohra 66 

Nessler Stefan H. 19 

Nettel Alejandro 58 

Nezami Bagher 100 

Ng Peter K. L. 25 

Nicholls Emma-Louise 30 

Nie Haiyan 74 

Niedbała Wojciech 10 

Nielsen Claus 1 

Nijman Vincent 3 

Nilsson Dan-Eric 108 

Nirala Satendra Kumar 79, 81 

Nitsche Frank 31 

Noda Chikayo 91 

Noël Pierre 68, 72, 75 

Nouira Said 66, 96 

Nowbahari Elise 59 

Nunes Maria Leonor 92 

Ocharan Francisco J. 66, 77 

Oettler Jan 59 

Ogawa Hiroto 45 

Ogereau David 16 

Oka Kotaro 45 

Okhlopkov Innokenti M. 76 

Okoneshnikov Vasily V. 6, 7 

Olivero Jesús 97 

Oppelt Angelika 59 

Oraie Hamzeh 39 

Orrell Tom 12 

Orus Ilyas 10 

Ott S. 8 

Oukil Salah 81 

Outomuro David 66 

Outreman Yannick 78 

Ozouf-Costaz Catherine 33, 38 

Pailhoux Eric 33, 37, 38 

Pailleret Marie 88, 90, 93 

Pairault Catherine 37 

Palacios Vicente 54, 74 

Palanca Antonio 42, 80 

Pallaoro Armin 90 

Palomares Francisco 101 

Pannetier Maëlle 33, 37, 38 

Papachristoforou Alexandros 66 

Pape Thomas 24, 25 

Papp Laszlo 25 

Paris Jérôme 75


Park Young-Hyang 70 

Parks Duncan 15 

Parmentier Eric 53 

Pasco Emmanuel 44 

Pascual Gabriel 52 

Pasquet Eric 5 

Patou Marie-Lilith 10 

Patterson David 25 

Pechmann Matthias 44 

Peixoto Rosana 74 

Peleg Nir 68 

Penrad-Mobayed May 38 

Pepey Elodie 35, 38 

Perna Andrea 59 

Pernice Mathieu 91 

Perrard A. 102 

Perrin Caroline 38 

Petit Odile 67 

Petroni Giulio 31 

Petrosyan Varos G. 75 

Pierce Sidney K. 91 

Pierre Jean-Sébastien 78 

Piiroinen S. 99 

Pinto Nuno 74 

Plouviez Sophie 91 

Polaszek Andrew 25 

Polymeni Rosa-Maria 2 

Pont Adrian C. 24 

Pook Catharine E. 52 

Poolny Thibault 65 

Poonlaphdecha Srisupaph 35, 38 

Pope Robert 65 

Poprawa Izabela 104 

Potapov Sergey G. 75 

Poteaux-Léonard Chantal 58, 59 

Poulle Marie-Lazarine 63 

Poumerol Elodie 33 

Pozsgai Gabor 96 

Pradillon Florence 88, 91 

Praseuth Danièle 80 

Preud'Homme Rose-Line 75 

Privé-Gill Catherine 90 

Prodon Roger 64 

Pshennikov Anatoli E. 7, 76 

Puillandre Nicolas 11, 12 

Puiseux-Dao Simone 80 

Pyle Richard L 23, 25 

Qadri Ayaz Hussain 11 

Quammen David 21 

Queen Joanna 40 

Quérat Bruno 43 

Quessada Marie-Pierre 109 

Quiniou Françoise 82 

Rabet Nicolas 11, 43, 102 

Rahimian Hassan 5 

Rajabizadeh Khosro 11 

Ramalho Lais V. 11 

Rand D. 8 

Rastegar Pouyani Eskandar 30, 43 

Rastegar-Pouyani Nasrullah 3, 4, 11, 

30, 39, 96 

Ravary Fabien 63 

Ravaux Juliette 87, 93 

Real Raimundo 97 

Reeves Jessica 79 

Reinelt Birgit 35 

Reinhardt Klaus 19 

Remsen David 12, 26 

Ren Qian 49 

Renard (Deniel) Emmanuelle 44, 110 

Renault Lauriane 33, 37, 38 

Renjifo Camila 52 

Renjifo Juan Manuel 52 

Rešić Jasminka 42 

Rhizopoulos Sophia 2 

Riascos Darío 52 

Richard-Yris Marie-Annick 62 

Richer de Forges Bertrand 5 

Rigaud Thierry 88 

Riou Virginie 88 

Rittschof Daniel 45 

Riveros Alain 52 

Rivière-Ung Visotheary 8 

Robin Jean-Patrice 67, 103 

Robinson-Willmott Julia 70 

Roco Alvaro 39 

Rodrigues Clara F. 92 

Rodríguez Claudina 78 

Roeszler Kelly N. 40 

Romdhane Mohamed Salah 70 

Rome Quentin 102 

Roméo Michèle 82 

Ronquist Fredrik 26 

Roques Alain 101 

Rortais Agnès 66, 82, 102 

Rosenberg Gary 25 

Rost-Roszkowska Magdalena 104 

Rouiller-Fabre Virginie 37 

Roussel E. 93 

Rovie-Ryan Jeffrine Japning 10 

Russell Anthony 110 

Ryan Michael 55 

Rybak Fanny 53 

Ryll B. 8 

Rzeznik-Orignac Janura 90 

Sabaneyeva Elena V. 32 

Sadiguettaf Hassiba 39 

Sadok S. 92 

Saedlou Nima 90 

Safronova Larisa D. 75 

Samadi Sarah 11, 12, 14, 89 

Sanchez Armando 52 

Sanchez Sophie 84, 92 

Sánchez Antonio 39 

Sánchez Víctor 17 

Santos Sara M. 75 

Sari Alireza 12, 92 

Satoh Nori 104 

Satoshi F. 10 

Saucède Thomas 88, 95 

Sazatornil Victor 74 

Scalone Romain 102 

Scharti Manfred 33 

Scheckenbach Frank 31 

Schedl Andreas 34 

Schindel David 26 

Schipper Jan 71 

Schlueter Mark 97, 110 

Schmidt Cornelia 33 

Schmitt Michael 18, 19 

Schneider Jutta M. 19 

Scholtz Gerhard 45, 111 

Schubart Christoph D. 8 

Schulteis Christina 33 

Schwager Evelyn E. 44 

Schwartz Julie A. 91 

Scohier Alexandra 59 

Secor Stephen 65 

Sedalischev Victor T. 76 

Segers Hendrik 23 

Segev Ori 68 

Segner Helmut 82 

Ségurens Béatrice 33 

Selmi Salah 84, 92 

Selmi Slaheddine 66 

Selz Yvonne 33 

Sendova-Franks Ana 60 

Servigne Pablo 57 

Shaikh Abdul Manan 12 

Shaikh Muhammad Yousuf 69 

Shams-Shoaei, Seyedeh Zahra 12 

Shatilovich Anastassia 32 

Sheibani-Tezerji Raheleh 92 

Sheikh. Muhammad Yousuf 11 

Shenbrot Georgy 96 

Sheremetyeva Irina 9 

Shi Dazhao 13, 16 

Shi Xiu-Zhen 49 

Shillito Bruce 86, 87, 93 

Shimoi Toshinobu 45 

Shmakova Lubov 32 

Shoemaker Christina M. 40 

- 107 - 

Shrivastava Sadhana 79, 82 

Shukla Sangeeta 79, 82 

Siblet Jean-Philippe 75 

Simberloff Daniel 75 

Sime-Ngando Télesphore 32 

Sinclair Andrew H. 39, 40 

Singh Varsha 79, 82 

Sinovčić Gorenka 34, 38, 94 

Siveter David J. 27 

Siveter Derek J. 27 

Skovorodkin Ilya N. 32 

Sleptsov Sleptsov M. 7 

Slyusarev George 45 

Smaï-Hamdidouche Souaâd 40 

Smith Craig A. 40 

Sokolowski Maria B. 58 

Solomonov Nickita G. 7, 75, 76 

Solomonova Tatyana N. 76, 77 

Song Yanling 74 

Speed Terence P 39 

Srijunngam Jirarach 40 

Srivastava Arun 97 

Stenseth Nils Chr. 97 

Stepanova Olga N. 77 

Stermin Alexandru Nicolae 77 

Stockley Paula 20 

Stöcklin Reto 51 

Storch Volker 17 

Strelkov Petr 61 

Stys Pavel 25 

Sues Hans-Dieter 21 

Sugisaki Masashi 7 

Sumana Annagiri 56 

Sun Jianghua 102 

Sutton Mark D. 27 

Suzuki Atsushi 47 

Svensson Erik I. 14 

Szyf Moshe 67 

Tafforeau Paul 8, 44 

Tanguy Arnaud 85 

Taylor Paul D. 11 

Teimory Azad 28, 100 

Templeton Alan R. 68 

Theraulaz Guy 57, 59 

Thibault Jean-Claude 5 

Thirakhupt Kumthorn 37 

Thomas Richard 8 

Thompson Christian F. 24 

Tillier Simon 13, 26 

Tirard Stéphane 107 

Töhönen Virpi 35 

Tomasović Maja 42 

Torralba-Burrial Antonio 77 

Torres Keila 78 

Trabelsi Monia 84 

Troudi Dhaker 93 

Tu Lihong 20 

Tuerkay Michael 17 

Tutman Pero 103 

Uhl Gabriele 19 

Vacelet Jean 44 

Valade Pierre 67 

van Alphen Jacques 78 

van Baaren Joan 78 

van Tol Jan 25 

Vance K. 8 

Vandewalle Pierre 53 

Vasiliev Vladimir N. 76 

Vast Claude 111 

Veitia Reiner 35 

Velasco Julián 54 

Vences Miguel 3 

Verna Caroline 93 

Verni Franco 31 

Véron Géraldine 10 

Verrier Delphine 103 

Viard Frédérique 91 

Vidal Nicolas 5, 8, 52 

Vidjak Olja 94 

Vignes-Lebbe Régine 8, 9, 95 

Vilain Eric 39


Vilcinskas Andreas 48, 50 

Vilimova Jitka 104 

Villemant Claire 102 

Vinokurov Vladimir N. 76 

Virant-Doberlet Meta 53 

Viriot Laurent 44 

Vladimirtseva Maria V. 7 

Vogler Alfried P. 26 

Volff Jean-Nicolas 33 

Voloshina Inna V. 9 

von Rintelen Thomas 7 

Wägele Johann-Wolfgang 13 

Wake David B. 15 

Wallacides Angelina 34, 40 

Waloszek Dieter 27 

Wang Deng 13 

Wang Haiyan 15 

Wang Jin-Xing 49, 50 

Wang John 59 

Wanniger Andreas 41, 46 

Ward David J. 30 

Watanabe Kosuke 45 

Wattanasirmkit Kingkaew 40 

Webster Gordon 92 

Wedell Anna 35 

Weightman Andrew 92 

Weisse Thomas 32 

Wenseleers Tom 60 

Wessels Stephane 35 

Wetzel Silke 91 

Wheeler Quentin D. 22 

White Stefan 39 

Wicker-Thomas Claude 67 

Wiens John 16 

Willmott Keith 70 

Wilson Edward O. 22 

Winckler Patrick 42 

Wink Michael 30, 43 

Wise Maria 45 

Wollesen Tim 46 

Woodcock D. 8 

Wright Debbie 25 

Wu Pengju 71 

Wurmser François 16 

Wüster Wolfgang 52 

Wylezich Claudia 31 

Xu Jianping 83 

Xu Zheng-Long 49 

Xu Zhongneng 67 

Yamaguchi J. 46 

Yang Jian 81 

Yasugi Sadao 105 

Yin Haiping 83 

Yoshida Shigeru 46 

You Zhangqian 15 

Zacanti Francesco 39 

Zaharescu Dragos 42, 80 

Zarrate Diego 71 

Zavjalov Evgenii 78 

Zbinden Magali 86, 89, 90, 93 

- 108 - 

Zemunik Tatijana 42 

Zeng Yan 64 

Zhang De-Xing 98 

Zhang Guofan 15 

Zhang Jianjun 13, 16 

Zhang Linyuan 64 

Zhang Qiong 74 

Zhang Shuyong 93 

Zhang Zhibin 98 

Zhang Zhi-qiang 23, 25 

Zhang Zihui 30 

Zhang Endi 71, 100 

Zhao Lili 102 

Zhao Xiao-Fan 49 

Zhao Yunlong 81, 84 

Zheng Guangmei 30 

Zheng Youfeng 74 

Zheng Zhigao 74 

Zhong Zhenyu 64 

Zhou Qingchun 33 

Zhou Xiangking 83 

Zhou Yi 78 

Zhukovskaya N. 4 

Zimmer Cédric 67 

Zimmermann Gabrielle 67 

Zinoviev Andrei 30 

Zorica Barbara 34, 94 

Zubaid Akbar 10

CONTENT - International Society of Zoological Sciences

You also want an ePaper? Increase the reach of your titles

Delete template?

Save as template?