PhD Arthur Decae 2010 - Ghent Ecology - Universiteit Gent

Diversity and distribution of
mygalomorph spiders in the
Mediterranean region.
Arthur Emile Decae

Cover drawing:
Nemesia randa Decae 2005 Female, dorsal view. Drawing by the author.

Voor
Jean-Pierre MAELFAIT
Ik heb je veel te kort gekend.
“Almost the whole summer was lost with this agonizing labor. At last on a quite trifling
occasion I came nearer to the truth.”
Johannes Kepler in Mysterium Cosmographicum 1596.
English translation from D.J. Boorstin 1983
The Discoverers Vol. 1

UNIVERSITEIT GENT
Faculteit Wetenschappen
Onderzoeksgroep Terrestrische Ecologie (TEREC)
Diversity and distribution of mygalomorph spiders in the Mediterranean
region.
Diversiteit en verspreiding van mygalomorfe spinnen in het Middellandse Zeegebied.
Arthur Emile Decae
Thesis submitted to the University of Ghent in order to acquire the PhD-degree in Sciences.
Proefschrift ter verwerving van het doctoraat in de wetenschappen aan de Universiteit van
Gent
Promotor: Prof. Dr. D.Bonte
examencommissie: Dr. R. Jocqué & Prof. Dr. F. Hendrickx
leescommissie: Prof. Dr. J. Mertens, Dr. R. Bosmans, Dr. J. Bosselaers

Contents
(page numbers and running heads present in the hard cover are absent in this
PDF File)
Acknowledgements. 1
Chapter 1 General introduction. 2 - 15
Chapter 2 Trapdoor spiders of the genus Nemesia Audouin, 1826
on Majorca and Ibiza: taxonomy, distribution and behaviour
(Araneae, Mygalomorphae, Nemesiidae). 16 -46
Chapter 3
Chapter 4
Chapter 5
Chapter 6
Chapter 7
Taxonomic Review of the Portuguese Nemesiidae
(Araneae, Mygalomorphae). 47 -68
Iberesia, a new genus of trapdoor spiders (Araneae, Nemesiidae)
from Portugal & Spain. 69 -79
Sub-generic diversity and distribution in the Mediterranean
trapdoor spider genus Nemesia Audouin 1826
(Araneae, Mygalomorphae, Nemesiidae). 80 - 103
The genus Ummidia Thorell, 1875 in the western Mediterranean,
a review (Araneae, Mygalomorphae, Ctenizidae). 104 -119
Systematics of the trapdoor spider genus Cyrtocarenum
Ausserer, 1871 (Araneae, Ctenizidae). 120 - 134
Chapter 8 General conclusions. 135 - 149
Summary/Samenvatting. 150 - 153
Literature. 154 -162
Epilogue. 163

Acknowledgements
Because this thesis consists of a series of papers that have already been published, or that are
submitted for publication, acknowledgements are provided separately with each paper or in
each chapter. The thesis could not have been written however without the help of others not
directly involved in the published material.
My special thanks go to Dr. Rudy Jocqué and the late Prof. Dr. Jean–Pierre Maelfait for their
warm enthusiasm, active help and provision of the conditions in which this project could
succeed. I furthermore thank Jean-Pierre for taking-up the position as my promotor and for
the valuable advice he gave me in the earlier stages of the work. I am also very grateful to
Prof. Dr. Dries Bonte for taking over the difficult task of being my promotor after the tragic
and much too early death of Jean-Pierre and to Prof. Dr. Luc Lens for his friendly help and
willingness to give me a place to work at the TEREC laboratories. I thank the members of the
promotion committees Prof. Dr. D. Bonte, Dr. R. Jocqué, Prof. Dr. F. Hendrickx, Prof. Dr. J.
Mertens, Dr. R. Bosmans and Dr. Jan Bosselaers for their enthusiasm to contribute to my
studies and I thank all my colleague PhD students for the friendly reception they provided me
at the TEREC laboratories. I thank the University of Gent for accepting me as a student and
for financing the expenses made in order to successfully carry out this thesis work. I am also
greatly indebted to many personal friends and family members for their mental support and
the constant interest they showed in the progression of my spider work. Finally I thank Nollie
Hallensleben, my wife and life’s companion for over forty years, for her help in preparing this
thesis.
A thesis usually marks the beginning of a scientific career and is therefore accompanied by a
curriculum vitae and a list of publications aimed to help the young doctor at a prospective
professional start. That is not the case here. This thesis is written at the end of a professional
career in which science did not play a central role, but was constantly present as a matter of
special interest and useful background to achieve all sorts of practical goals. The motivation
to nevertheless produce this thesis is that I hope to encourage interest in a branch of science
that, after a providential start in the last quarter of the nineteenth century, got bogged down in
its own confusion and the changing interests in the biological sciences. The study of
mygalomorph spiders started off in the Mediterranean where early savants of Natural History
such as J.T. Moggridge, E. Simon, A. Ausserer, O. Pickard-Cambridge, T. Thorell and others
carried out their pioneering research. Their work, building a solid taxonomic basis for
understanding mygalomorph spider diversity in the Mediterranean Region, unfortunately has
never been satisfactorily finished. When in the course of the twentieth century the focus of
biological science shifted from classical Systematics and Taxonomy to more predictive and
process orientated studies, attention for biological description faded and no further progress
was made in the knowledge of the Mediterranean mygalomorph spider fauna. In the
awakening of modern biodiversity studies however, there is a renewed interest in questions of
species identity and identification. Knowledge about how many species there are and which
species occurs where is important again. Due to the unfinished taxonomical work these
questions, when asked in relation to Mediterranean mygalomorph spiders, are very difficult to
answer. Nevertheless, these questions are now asked by a number of young researchers that
live and work in countries bordering the Mediterranean or that are located in the
Mediterranean climate zone. As an older arachnologist I hope to be able to act as a mediator
between the generations providing helpful background information for new studies in
Phylogenetics and Phylogeography. This thesis is therefore dedicated to all these young
researchers whose names feature in the acknowledgements of the here collected papers.
1

Chapter 1
General introduction.
Arthur Decae
August 2010
Front-page illustration. Cladograms of higher classifications of spiders (Order Araneae) Top:
conventional following Platnick & Gertsch 1976. Bottom: here proposed alternative cladogram
based on functional apomorphies. Lateral views of spider bodies, appendages omitted.

Motivation for this thesis
This thesis is written as an effort to contribute basic knowledge to the study of Arachnology,
hoping this will help future research in a branch of biology that I regard as a potentially rich
source of important scientific insight. All papers in this thesis contain original information on
Mediterranean mygalomorph spiders provided in order to facilitate and stimulate more
detailed future research and understanding of these fascinating, yet little known, animals.
Aim
The central goal of this thesis is to gain insight in the historical build-up of regional
mygalomorph diversity in the expectation that this will aid a more general understanding of
growing faunal complexity as it has occurred on earth.
Personal Note
When in 1983 I took my exams for a Master Degree in Biology at the University of
Groningen one of the members in the jury asked me if I did not think that my studies had been
too narrowly focused at just one animal group. I had worked on subjects in taxonomy,
biogeography, ecology, ethology and comparative morphology of sensory organs all in one
genus of mygalomorph spiders; the genus Cyrtocarenum Ausserer 1871. To my relief, and
before I could answer, another member on the committee countered that there would have
been no critical questions on this point if the one animal group had been fruit flies
(Drosophila melanogaster), rats (Rattus norvegicus) or zebra fish (Brachydanio rerio). The
point of course is that fruit flies, rats, zebra fish and a hand full of other animals are widely
considered ideal model organisms for a range of interests in zoological research and that it
remains to be seen if we could learn anything important from mygalomorph spiders. In fact, I
believe we can learn something new and important from indiscriminately which animal we
choose to study, but that the study of mygalomorph spiders offers more.
The study of spiders in general offers more than the study of other animal groups because
spiders are unique in producing a wide range of intricate and seemingly intelligent and/or
geometric constructions made of a combination of material found in nature and self produced
excretions: such as saliva and silk. These constructions, nests, webs, drag lines, cocoons,
trapdoors, wrappings, ties and a number of other functional devices, may be seen as material
expressions of complex behavior that no other animal, save the human species, leaves
behind 1 . To produce their complex constructs spiders of course possess a range of
morphological, physiological and neurological qualities not found in any other animal group
and it is only because spiders are not of any obvious economical or medical interest that
araneology (the study of spiders) is a relatively remote branch of the biological sciences 2 .
This is to be regretted, because spiders not only possess all these unique qualities of direct
biological interest, they also preserve, within their diversity, the reflection of an evolutionary
history that dates back to the beginning of terrestrial animal life (Decae 1984). Although the
fossil record of spiders is relatively poor, spiders do not fossilize easily (Selden & Penney
2010), much of the history of spiders can be read from extant species. Apparently spiders
have been a marked biological success throughout evolutionary time, surviving in great
1 Beavers, birds, termites, bees and other animals produce amazing constructions in size and
complexity, but rarely as versatile, intricate, geometrical and multifunctional as spiders and
humans do (see e.g. Von Frisch 1974, Hansell 2007).
2 Although properties of spider silk and venom in particular might offer prospects of medical,
agricultural and other applications (e.g. Bailey & Chada 1968, Sterling et. al 1992, Patrick &
Canard 1997, Novak 2001, Scheller et. al. 2001, Escoubas & Rash 2004) it seems to be very
difficult to economically exploit spider products.

diversity (Penney et. al. 2003) all cataclysmic mass extinctions that mark the divisions of the
geological timescale and that have punctuated the histories of many animal groups. There is
no clear evidence of major evolutionary bottle necks 3 in the contemporary spider fauna. Three
major lineages all dating far back in geological time have living representatives (see frontpage
illustration). The primordial Mesothelae have 5 known genera and 87 species in eastern
Asia, the primitive Mygalomorphae have 320 genera and 2643 known species distributed on
all continents, being absent only from circum polar regions, and the Araneomorphae finally
with 3777 genera and 38523 species 4 have an even wider cosmopolitan distribution.
Moreover, in their degree of evolutionary development the Araneomorphae contain both
rather generalized forms such as the lampshade web spiders 5 (Hypochilidae), the tube web
spiders (Segestriidae) and the daddy-long-legs spiders (Pholcidae) and superbly specialized
lineages such as the orb weavers (Araneidae), the wolf spiders (Lycosidae) and the jumping
spiders (Salticidae). Paleontology has shown that many extant spider families can be traced
back in time deep into the Mesozoic era and some Mesothelae even into Paleozoic times
(Selden & Penney 2010). This means that a well developed phylogenetic tree of spiders would
provide a rare telescopic view into the evolutionary history of a major animal group. Over the
passed decades arachnologists have greatly advanced in the reconstruction of this tree but
advances have been biased towards the more advanced Araneomorphae whereas knowledge
of the primitive Orthognatha (Mesothelae + Mygalomorphae) is relatively underdeveloped
(Coddington 2005). This is to be regretted again because precisely the living existence of
these primitive spiders in great abundance and diversity alongside advanced forms give
spiders their unmatched evolutionary scope. Moreover, the very effective mechanism for
aerial dispersal (ballooning), characteristic for most araneomorph spiders, is lacking in most
orthognathe lineages (Main 1976, Coyle 1985b). This means that whereas araneomorph
distributions (at lower taxonomic levels) are generally to be understood in terms of ecological
opportunity and capacity for dispersal, orthognathe spiders tend to be ‘locked-up’ in their
centers of origin as distinct local endemics. This special character of orthognathe spiders
means that our telescopic view of spider evolution can be focused in space as well as in time
and that primitive spiders have outstanding qualities for studies in historical biogeography and
other aspects of evolutionary biology.
To conclude this motivation for working on mygalomorph spiders I’ll turn back to my earlier
studies at the University of Groningen, where after graduation, I successfully applied for a
three year grant to study the ecology and behavior of Neotropical mygalomorph spiders at the
Smithsonian Tropical Research Institute (STRI) in Panama. My supervisor at STRI, the late
Dr. Mike Robinson, remarked on my arrival that he was sure mygalomorphs were present in
the area but that, except for the large bird-spiders (Theraphosidae), virtually nothing was
known about their whereabouts and their identities. A study of ecology and behavior
necessarily had to be preceded by a study in mygalomorph taxonomy if future results were to
be comprehensively communicated to the scientific community. In a nut shell this is the
situation that students of mygalomorphs still encounter today. In most parts of the world little
can be studied on mygalomorphs until their taxonomy is reasonably resolved. The problem is
that studies of mygalomorph taxonomy have a reputation of being difficult and frustrating
3 Although the Permian-Triassic extinction event might have seriously affected spider
diversity and modern cladograms (e.g. Penney et. al 2003 Fig. 1) suggest that most spider
radiation occurred after the P-T event.
4 As a result of new discoveries of species and genera and changing insights following
taxonomical revisions these numbers constantly change. The numbers given here follow the
World Spider Catalog version 10.5 (Platnick 2010).
5 Vernacular names follow Jocqué & Dippenaar-Schoeman 2006

(Coddington 2005) and therefore lags behind other developments in biological knowledge.
The rewards of studying mygalomorphs however will be in keeping with the efforts and a
much improved vista on the evolutionary history of a major animal group can thus be
obtained.
Outline and objectives
The Mediterranean mygalomorph fauna, as it is presently known, contains representatives of
seven different spider families, eleven genera and approximately 120 species (see Chapter 8
general conclusions). Current knowledge of Mediterranean Mygalomorphae largely rests on
old and often incoherent information that has accumulated over the past 200 years. However
all families and genera need thorough taxonomical revision before deeper biological insight
can be developed. This elementary though necessary work is beyond the scope of this thesis.
However two genera − Ummidia Thorell 1875 and Cyrtocarenum Ausserer 1871 − have been
fully revised on all material available (Chapters 6 & 7) and the complex and diverse genus
Nemesia Audouin 1826 has been revised for local geographic regions (the Balearic Islands in
Chapter 2 and Portugal in Chapter 3). To structure future research on the Mediterranean
Nemesiidae a genus level revision was carried out (Chapter 4) resulting in the splitting-off of
a new genus (Iberesia Decae & Cardoso 2005) from traditional Nemesia stock, and the
establishment of two subgenera (Pronemesia and Holonemesia) within Nemesia (Chapter 5).
Within both Pronemesia and Holonemesia several distinct species-groups are recognized and
discussed (Chapter 5). Work on the remaining mygalomorph genera occurring in the
Mediterranean Region −Atypus, Brachythele, Chaetopelma, Cteniza, Cyrtauchenius,
Ischnocolus, Macrothele and Idops− is restricted to descriptions of their species diversity and
Mediterranean distributions. The probable regions of origin for all these genera and their
affinities to related genera outside the Mediterranean are discussed in Chapter 8.
The central object of study is the Mediterranean mygalomorph spider fauna. To provide an
informative backdrop for understanding the arachnological information contained in this work
the history and biological importance of the Mediterranean Region as an area of high
biodiversity will be described below separate from an exposé on mygalomorph spiders. The
general background information on Mygalomorphae and their presumed sister-group the
Liphistiomorphae is intended to shed a new and original light on spider phylogeny and spider
evolution that departs from conventional arachnological views.
The Mediterranean Region
Why study the Mediterranean
Obviously there are many very good answers to this question depending on whatever your
line of interest is. One very good reason is that, for people, the Mediterranean is a very
pleasant place to be. This is not only evidenced by the huge success of modern mass tourism
in the region, it is also evidenced by early Homo erectus visitors coming straight from Africa.
H. erectus settled in the Mediterranean at least half a million years ago (Dennel & Roebroeks
1994, Turner 1999) and since that early period mankind has never left the beautiful places that
surround the bluest of all seas. Within the Mediterranean Region humans developed their
incredible success as a biological species. They started off as small tribes equipped with
stones and sticks looking for whatever nature would provide, but in the benign environment of
the Mediterranean they became farmers, citizens, soldiers, empire builders, discoverers of the
world and of the universe. For the rise of human civilization the Mediterranean certainly is a
‘hotspot’.

Recently the Mediterranean has acquired another qualification for being a hotspot. Within the
field of biological conservation the Mediterranean is proclaimed one of the world’s hotspots
of biodiversity (Meyers et.al. 2000). This qualification is used by the International Union for
the Conservation of Nature (IUCN) in order to focus human attention on the impoverishing
effects on nature of unbound exploitation of the environment. The Mediterranean, as a source
of natural wealth and beauty, is of special significance because of its unique evolutionary
history that makes it one of the richest biological regions outside the tropics (Naveh 2007).
Moreover the Mediterranean flora and fauna contains a very high proportion of endemic
species, species that only occur in the Mediterranean, which gives the region the status of
being of critical importance for nature conservation 6 . Present knowledge of Mediterranean
biodiversity however is based on only a very thin slice of the local flora and fauna. It is
mainly focused on vascular plants, vertebrates and very few invertebrate groups 7 . Very little
information is available for the bulk of smaller animals and plants that actually build the
world’s biodiversity. This means that, from a biodiversity point of view, there is much to
discover in and around the Mediterranean, providing numerous outstandingly good reasons to
study the region.
A first question coming-up from the biodiversity perspective would be; where does it come
from Why is the Mediterranean so rich in endemic and other species of plants and animals
To answer that question we will have to go far back in time, long before the first human
settlements in the region, to the birth of the Mediterranean itself and its subsequent history. To
do that, we will have to turn to geology and geography and we may start with a map of the
world. The map of the world, as we all know it (Fig. 1), is only a reflection of the current
coastlines and the distribution of land and water over the surface of the planet. Since a number
of scientific discoveries and realizations, settling in human conscience over the past 250
years 8 , we now know that the map of the world is changing continuously and has changed
dramatically over millions of years of geological time. The main features of geography, such
as oceans, seas, continents and mountain ranges have not always been where we see them
today, but they have come and gone in different configurations. This shifting of continents
and oceans has had major effects on the evolution of plant- and animal species. Evolutionary
processes have broadly followed the dynamics in the earth’s crust to produce the regional
florae and faunae of the different continents, including that of the Mediterranean.
In connection to the here presented studies of Mediterranean mygalomorph spiders this idea
of related shifts in geography and faunal composition led to asking three fundamental
questions: (1) what is the origin of the Mediterranean, (2) why does it have such a high
species diversity, and (3) how did it get its mygalomorph spider fauna
Fig. 1 Familiar map of the World
showing existing coastlines and the
distribution of land and water over
the earth’s surface. Source: Google
maps of the world.
6 For actual numbers see www.biodiversity/hotspots.org/...
7 See IUCN, Red List on: www.iucn.org/redlist/
8 Discoveries of (1) ‘deep time’ (Hutton 1788, Lyell 1830-1833), (2) evolution by natural
selection (Darwin 1859), (3) continental drift (Wegener 1916), (4) seafloor spreading (Hess
1954, 1960).

Origin of the Mediterranean
Questions of origin are always difficult to approach because we have to find a starting point.
In the case of the Mediterranean this is particularly difficult, but we might start from the idea
that the forces that broke-up the ancient super continent Pangaea, some 170 million years ago,
led to the formation of the separate ‘continental plates’ that much later in time, were to form
the Mediterranean. This formation of the Mediterranean finally took place in a gigantic
‘tombola’ of collisions and segregations of fractions of the earths crust (see footnote 7 for
discoveries and discoverers of continental drift and seafloor spreading). The main processes to
be considered here are: (1) the opening of the North Atlantic Ocean and the vanishing of the
ancient Tethys Ocean, (2) the movements of the African and Arabian continental plates, (3)
birth of the Mediterranean Sea. For information on these phenomena I lean heavily on the
following sources: D. Ager’s (1980) celebrated book The Geology of Europe; Rögl &
Steiniger’s (1984) Chapter 10 in Fossils & Climate (Wiley Books) and Rögl’s (1999) short
discussion of the Mediterranean Palaeogeography in Geologica Carpatica.
1. The Atlantic and Tethys Oceans
The formation of the Atlantic Ocean started
near the equator with cracking open the
solid continental block of Pangaea, roughly
at a place where we now find the Caribbean.
A subsequent rapture moved from there to
the southwest to separate North and South
America. A second giant rapture moved
northeast to separate North America from
Africa and Europe. In the process southern
Europe was broken-up in a series of microplates
that formed an archipelago within the
northwestern Tethys Ocean (Fig. 2). It all
must have happened early in Jurassic times,
approximately 170 million years ago
(Pannekoek 1973), when giant dinosaurs
roamed the earth. The rapture of Pangaea
separated the northern continents from one
large southern continent that is here denoted
as Neogondwana 9 . To the east the young
Atlantic Ocean had a seaway connection with
the western parts of the ancient Tethys Ocean.
This inter-oceanic seaway is the location where
the Mediterranean was going to be formed. In
time the Tethys Ocean would be replaced by the
Indian Ocean (late Mesozoic to mid Cenozoic)
Fig. 2 Showing the young Atlantic Ocean (approx.
150myBP) separating the northern from the still
united southern continents and the seaway
connection with the western Tethys. Source:
R.Blakey.ucc.nau.edu/
through the northward movement of the Indo-Australian plate and the rotation of the Arabian
plate away from Africa to close the Atlantic-Tethys connection so shaping the Mediterranean.
9 An earlier separation between northern and southern continental blocks broadly denoted as
Laurasia and Gondwana had existed in the Palaeozoicum (400myBP-350myBP) before the
formation of Pangaea, hence the term Neogondwana for the southern continental block that
originated from the breakdown of Pangaea.

2. The African-Arabic Continent
Around 120myBP the African-Arabian
plate broke away from the great southern
continent of Neogondwana and drifted
towards the northeast ever closer to the
European archipelago. These movements
of the African-Arabian Plate in the late
Eocene and early Oligocene made the
old Tethys Ocean finally disappear
between 35myBP and 25myBP. In the
western parts of the largely shallow
seaway that remained, the Mediterranean
was gradually taking shape. The
Neoeuropean 10 archipelago was pushedup
against the Mesoeuropean mainland
and the formation of the modern
European continent slowly took shape.
To the south, the continuous stable and
passive continental front of Africa
contrasts sharply with the fragmented
archipelago of Neoeurope along the northern
shores of the Proto-Mediterranean (Fig.3). A
following anti-clockwise rotation of Arabia away
from Africa was going to terminate the last
remnants of the Tethys Ocean. To the north of the
Proto-Mediterranean the Paratethys became a
more and more isolated epicontinental sea in
Central and eastern Europe and in western Asia.
Fig. 3 Proto-Mediterranean and Paratethys
developing during late Oligocene- early Miocene
times (25myBP-15myBP). Source:
R.Blakey.ucc.nau.edu/
3. Birth of the Mediterranean Sea
The final formation of the Mediterranean Sea took shape in the Miocene between
approximately 20myBP and 10myBP. The rotation of the Arabian plate more and more closed
off the connection of the eastern Mediterranean with the former Tethys Ocean, and Africa
pushing-up in the West formed a separate western Mediterranean Basin surrounded by rising
Alpine mountain ranges. Within this western zone the former islands of the Maghreb, the
Betic micro-plate, Iberia and the Tyrrhenian micro-plate (including most of what is now the
Italian mainland) formed a separate geographical unit. In the Northeast continental microplates
that had existed as islands in the eastern Neoeuropean Archipellago (Taurus, Pindos,
Adriatica, Pannonica) were pushed against southwestern Asia reducing the Paratethys to a few
isolated basins that were to form the Black Sea, Caspian Sea and other smaller basins (Fig. 4).
Connections between the Paratethys and the Mediterranean and between the Mediterranean
and the Atlantic Ocean were closed. In the course of the Miocene this would change. The
disappearing shallow seaway to the East would finally shut off the Mediterranean from the
10 Terminology from Ager 1980, the Geology of Europe, who distinguishes between
Eoeurope in the far north, Palaeoeurope in Scandinavia and the British Isles, Mesoeuropes for
the Varischian effected parts of Central Europe and Neoeurope for the largely southern Alpine
regions.

Indian Ocean and the Strait of Gibraltar would open (probably periodically) to connect the
western Mediterranean with the Atlantic again. The western Mediterranean in this period was
a distinct sea from the eastern Mediterranean. Opening of the Strait of Sicily in time would
renew the connection between the eastern and western parts of the Mediterranean Basin. The
familiar geography of the region is now becoming visible although the major peninsulas and
archipelagos as well as the connection between the Black Sea (Bosporus – Dardanelle
connection) still had to develop (Fig. 4).
Fig. 4 Mediterranean in the early Miocene (approx. 20myBP). The western Mediterranean is separated
from the eastern Mediterranean, The Paratethyis is reduced to isolated basins and the European
Continent is taking its final shape. Source: R.Blakey.ucc.nau.edu/
Mediterranean Biodiversity
What is the connection between the above exposé on plate tectonics and the high biodiversity
in the Mediterranean The geological history sketched-out here shows that the Mediterranean
was formed at the crossroads of the African, Asian and European continents (Fig. 3). So, parts
of the native florae and faunae of these three continents are expected to have met and mixed
within the Mediterranean leading to an increase of species richness. Moreover one of the three
contributing continents, Europe, had existed as an extensive archipelago of larger and smaller
islands for many millions of years. In this archipelago the evolution of island endemics must
have been high and the contribution from Europe to the later Mediterranean flora and fauna
must have been particularly rich. This reasoning however only relates to terrestrial
biodiversity. The repeated opening and closing of seaways and the constant formation of
separate aquatic basins both in the Mediterranean and the Paratethys must have had further
effects of the enrichment on the marine flora and fauna of the Mediterranean. These basic
conditions for the development of a particularly rich biodiversity in the Mediterranean have
prevailed throughout the geological formation of the region to our present time. And there is
even more than the historical high diversity of terrestrial and marine biomes in the

Mediterranean. Recently a very diverse deep-sea fauna was discovered that is believed to
have originated from special conditions of fragmentation and isolation of local deep water
holes that pertained oceanic conditions during the Messinian salinity crisis. The Messinian
salinity crisis has been a major event (6myBP) in the history of the Mediterranean in which
the Atlantic connection was blocked and much of the sea bottom fell dry (Danovaro et.al
2010). Another instant of severe fragmentation and isolation of local elements of flora and
fauna has occurred repeatedly in the course of numerous Pleistocene glaciations (Médial &
Diadema 2009). During the height of these glaciations, when the permafrost covered most of
Europe and the permafrost front reached the Alps, the particularly heterogenic habitat of the
Mediterranean produced local refuge zones in which species could survive. Again the
repeated fragmentation and isolation of populations is expected to have produced increasing
biodiversity. Finally the present flora and fauna of the Mediterranean still survives in, and is
adapted to the small scale heterogeneity that is typical for the Mediterranean environment,
where soil depth, micro-relief, availability of moisture, nutrients, rocky outcrops, tree cover,
litter cover and shade are all patchily distributed (Rundel 1998). All this has attributed to the
very high species diversity in the Mediterranean that needs to be studied before too severe
destruction overcomes the natural resilience of the local flora and fauna. In conclusion, the
rich species diversity of the Mediterranean can not be fully understood in terms of prevalent
ecological conditions, it must be understood in the context of complex historical and
environmental factors that go back deep into geological time. The history of Mediterranean
biodiversity is expected to be reflected in the diversity and distributions of smaller species
with strong qualities for survival and a limited capacity for dispersal. Such species are
particularly common among mygalomorph spiders.
Arachnology
Arachnology is the scientific study of spiders and spider-like animals. Spiders are descendents
of an ancient animal group, the Chelicerata, traces of which are found as fossils in rocks
dating back to the Cambrian Period (>500myBP). This is the period when recognizable
animals appear for the first time in nature (Meglitsch 1972). At that time all animals were
aquatic in habit and no complex forms of life occurred on the land at all. Still fully aquatic,
during the Silurian period (430-413myBP), the Chelicerata had diversified in several animal
groups, one of which was the Order Arachnida. These early arachnids might well have been
among the very first animals that invaded the land (Decae 1984). Such an invasion of animal
groups into a new and pristine environment promotes adaptive radiation and the arachnids
showed strong diversification probably leading to the origin of all fifteen taxa we presently
recognize (Table 1). Although we have no Paleozoic records for the small and fragile bodied
Schizomida and Palpigradi the major radiation of arachnids is likely to have occurred in the
Devonian and Carboniferous periods (Dunlop & Selden 2009). In the course of evolutionary
history only four arachnid Orders became extinct and eleven Orders are still alive today
(Table 1).
Because the Arachnida were early colonizers of the terrestrial environment it is only to be
expected that initially they were adapted to sheltered habitat situations where risks of
desiccation were minimized. In fact, nearly all arachnids that have survived the eons of
geological time are still adapted to very sheltered habitat situations. They survive in cracks
and crevices only to venture out at under conditions of high humidity or darkness. Many
occur in caves, under litter, moss, stones, logs or rubbish, and only very few have adapted to
exposed living conditions. True spiders, such as the araneomorph orb-web weavers
(Araneidae), jumping spiders (Salticidae), wolf spiders (Lycosidae) and a few other groups
are aberrant arachnids in a sense that they have adapted to living in exposed conditions, often
in bright sun shine and hot climates. Also among spiders, these exposed living groups are the

most seen and studied groups, but actually they represent the exception rather than the rule. It
is insufficiently realized by students of spiders (arachnologists) that the most conspicuous
spiders naturally draw most of the attention and that much hypothesizing and theorizing in
Arachnology is biased to explaining the ways of these ‘aberrant’ spiders. Most spider species,
as virtually all their arachnid relatives, are nocturnal and secretive creatures that live in hidden
and sheltered positions and that, because of this, have attracted little attention from zoologists.
Following conventional taxonomical wisdom the Order Araneae (spiders) contains two
Suborders: Mesothelae and Opisthothelae. The Mesothelae contain only one family of living
spiders: Liphistiidae. The Opisthothelae are further subdivided into two Infraorders (see frontpage
illustration): Mygalomorphae and Araneomorphae (Platnick & Gertsch 1976). Only the
Mygalomorphae are of further concern here.
Mygalomorphae
Mygalomorph spiders are a prime example of a group of understudied secretive living
arachnids. Nevertheless, the mygalomorph spiders are a major group of animals in themselves
(320 genera, 2643 species known). They occur in most terrestrial and some semi-aquatic
environments on all continents except Antarctica. In many habitats they are among the
dominant predators of ground living arthropods and some species at least incidentally prey on
other animal groups including small vertebrates (Raven 2010). Mygalomorphae are primitive
spiders that have retained many ancestral traits including their way of living. Just as in the
true spiders of the Infraorder Araneomorphae however, some successful diversification has
taken place within the Mygalomorphae. The large bird-spiders of the family Theraphosidae
are a fine example. Some species in this family that have adopted a more actively hunting and
wandering way of life, have been very successful. Their biological success is reflected in the
highest known species diversity among mygalomorph spiders and an almost worldwide
distribution in the warmer climate zones.
Table1. List of Arachnida taxa, with their first appearance in the fossil record and their status of being,
living or extinct (after Dunlop & Selden 2009). Note the Paleozoic origin of all taxa except
Schizomida and Palpigradi. Fossil traces of these two taxa of small and fragile bodied animals have
not as yet turned up in Paleozoic rocks.
TAXON OLDEST RECORD GEOLOGICAL PERIOD STATUS
myBP
1 Scorpiones 428 Silurian Living
2 Trichonotarbida 419 Silurian Extinct
3 Phalangiotarbida 411 Devonian Extinct
4 Opiliones 410 Devonian Living
5 Acari 410 Devonian Living
6 Pseudoscorpiones 392 Devonian Living
7 Uraraneida 392 Devonian Extinct
8 Ricinulei 319 Carboniferous Living
9 Thelyphonida 319 Carboniferous Living
10 Haptopoda 312 Carboniferous Extinct
11 Araneae 312 Carboniferous Living
12 Amblypygi 312 Carboniferous Living
13 Solifigae 308 Carboniferous Living
14 Schizomida 34 Oligocene Living
15 Palpigradi 5 Pliocene Living

The funnel-web spiders of the families Dipluridae and Hexathelidae may also be regarded as
advanced mygalomorph spiders in their adopted life strategies 11 . They build elaborated webs
for capturing prey and in this respect closely resemble some of the ‘true spiders’ (e.g.
members of the family Agelenidae).
Most mygalomorph spiders however have retained an ancestral lifestyle that they share with
the truly primitive Liphistiomorphae (the segmented trapdoor spiders of SE Asia). The central
aspect of this ancestral lifestyle is the construction of tunnels in which the spider spends most,
if not all of its life. Because individual mygalomorph spiders may live for many years their
burrows need to be solid and stable structures. Usually the burrow tunnels are dug in the
ground (occasionally in rotting logs), in few species they exist as tunnel shaped extensions,
socks or cells on rock faces or tree trunks. Because we cannot travel back in geological time
we can never be sure that tunneling actually is the ancestral lifestyle of all spiders and the best
evidence to prove it is when we can collect sufficient circumstantial evidence. A good piece
of circumstantial evidence would be if tunneling was found to be the habit among the majority
of primitive Orthognatha (Mygalomorphae + Mesothelae). And actually it is.
Currently 16 families of orthognate spiders are recognized; one family in the Mesothelae and
fifteen families in the Mygalomorphae (Table 2). At least thirteen of these families (>80%)
have species that dig holes in the ground. Three of these families (Table 2) are not specially
noted for the burrows they dig. In eleven families of orthognate spiders however, burrow
construction is the dominant lifestyle. Actually the great majority of the species included in
these families are obligate burrowers. In other words, excavating burrows in the ground is by
far the majority lifestyle of primitive spiders living today and this alone should serve as strong
circumstantial evidence to defend the proposition that burrowing is the ancestral lifestyle for
all spiders. The fact that this burrowing lifestyle is ancestral for all spiders is insufficiently
realized in Arachnology. This is the basis of much misconception in attempts to understand
spider evolution. The prime misconception originates from interpretations of the function of
the orthognate chelicerae, the abdominal position of the spinnerets, the compact build with
short strong legs of Orthognatha (particularly the rear two pairs of legs) and the function of
the remarkable narrow and flexible pedicel that is present in all spiders. Interpretations of
these structures in arachnological literature are usually strict morphological and not
functional. Nevertheless understanding the adaptive function of this typical orthognate spider
morphology has major consequences for our view on the phylogeny of spiders in general. The
questions of why do primitive spiders have orthognate chelicerae, why are spinnerets located
on the abdomen, why do Orthognatha have stubby legs and why do all spiders have this
vulnerable narrow stalk, the pedicel, to connect their two bulky body parts, are rarely asked.
The reason is I think psychological; it is because our conception of nature is shaped by our
experience, perception and interest, and since we are predominantly visual animals our
experience, perception and interests are primarily fed by the things we readily see. Orthognate
spiders do not belong to those objects we readily see alive, we only find them dead in
museum collections and therefore very few arachnologists have ever looked at primitive
spiders beyond the things visible in dead specimens; static morphology.
If a civil engineer is asked to develop a machine for tunneling in the earth he/she will comeup
with a machine as shown in Fig. 1. The machine shown is called a TBM (tunnel boring
machine). The essence of its construction is that it has a strong cutting and scraping device up
front, a compact body, a mechanism for grip on the tunnel wall that allows it to be driven
11 Mecicobothriidae and/or Microstigmatidae may also fall in this category, although little is
known of their ways of life.

forward with great force, and a facility for stabilizing the tunnel walls at its rear end (if it is
tunneling in soft rocks). Now if we look at a mygalomorph spider through the eyes of a civil
engineer we would recognize the perfect TBM (Fig. 2). The only difference between a
mechanical TBM and an orthognate spider is the absence of a conveyor belt to remove the
dug-out material. The problem of soil removal from the tunnel under construction is solved
differently in spiders. The key solution to this problem is that spiders have a particularly
narrow and extremely flexible waist, the pedicel that allows it to pivot in the narrow shaft of
its tunnel and to carry-out particles of dug-out soil.
Observation of a trapdoor spider
excavating its burrow immediately
shows the prime functions of the
orthognate chelicerae (scraping,
digging and carrying soil), the stubby
legs (strong grip on the burrow wall),
abdominal spinnerets (stabilizing the
tunnel behind the spider) and the
narrow pedicel (allowing the spider to
pivot for soil removal).
If we look at spiders in this way it is
clear that they differ very much from
their putative sister group the
Amblypygi. The forward orientation
of the chelicerae may superficially be
the same in orthognate spiders and
the Pedipalpi (Amblypygi + Uropygi
+ Schizomida. Harvey 2003) their
function is fundamentally different.
None of the Pedipalpi use their
chelicerae for digging, in orthognate
spiders however, digging is done solely
with the aid of the fangs and chelicerae.
Comparison of the morphology of the
two hind pairs of legs in orthognate
spiders and the Pedipalpi reveals that in
spiders these legs are short and very
strong, against being quite slender to
very slender in the Pedipalpi. The
pedicel is narrow as well in Pedipalpi
as in orthognate spiders, presumably
because all these animals have to move
in the confines of cracks and crevices,
but although there are no comparative
Fig. 1 Tunnel Boring Machine (TBM). One of many models,
all of basically similar design, offered by construction
companies on the internet for infrastructural tunnel drilling
projects.
Fig. 2 Atypus affinis depicted as a TBM.
data that I know of, I would predict that the pedicel of spiders is much more flexible than that
of the Pedipalpi. Finally, spiders are the only creatures on earth that have developed an
abdominal spinning apparatus, working with extreme precision and ideally suited and placed
for stabilizing loose soil during construction work. Coyle (1981) has studied and described all
the above mentioned aspects of burrow construction behavior for Ummidia in detail. My
personal observations confirm that Nemesia, Iberesia, Cteniza, Cyrtauchenius and
Cyrtocarenum all construct their burrows in an identical manner as is reported for Ummidia.

Coyle (1981) also negates the common misconception that the front legs of the Ctenizidae are
equipped with ‘digging spines’. He shows that the front legs of Ummidia play no role in
digging and the same is true for Cteniza and Cyrtocarenum. The strong short spines on the
distal parts of the front legs and palps function in prey capture, rather like the sharp conical
teeth of sharks and crocodiles that prevent prey from escaping.
If we look at spiders as originally tunneling animals the current phylogeny in which
Mygalomorphae is placed as the sister group of Araneomorphae (Platnick & Gertsch 1976)
appears unlikely. An alternative cladogram, as earlier proposed by Bristowe (1933) seems to
be more in accordance to reality (see front-page illustration bottom). The specially digging
adapted chelicerae of the Liphistiidae and Mygalomorphae appear to be a perfect functional
synapomorphy for a Suborder Orthognatha as well as the strong and often specially for lock
and grip adapted legs III and IV. This view on the basic phylogeny of the Araneae would
bring Goloboff’s (1993) remarkable decision for using Mesothelae (instead of the cladistically
established sister-group Araneomorphae 12 ) for rooting his Reanalysis of Mygalomorph Spider
Families in an enlightening perspective. The here described ideas on spider phylogeny would
also provide a more comprehensible view on spider evolution in general in which it is seen as
the result of two highly successful waves of adaptive radiation; one in two dimensional space
(Orthognatha) and one in three dimensional space (Labidognatha).
As described above, mygalomorph spiders are secretively living animals. Although some
species build elaborate webs in very visible locations (e.g. Macrothele calpeiana in southern
Spain), others hide either in crevices or under stones. The majority of Mediterranean
Mygalomorphae however are trapdoor spiders that inhabit self dug burrows up to 30cm deep
in the ground. The entrances of the burrows are usually closed by a hinged door that is often
perfectly camouflaged. The study of trapdoor spiders therefore needs field experience. The
best guide here is Moggridge’s (1873, 1874) book Harvesting Ants and Trapdoor Spiders.
Although the book is somewhat antiquated it perfectly explains the way in which trapdoor
spiders can be best studied in the field. Also it shows the critical importance of looking further
than plain morphology for understanding Arachnology and illuminates, in superb plates and
figures the amazing works of tunneling and underground construction works of trapdoor
spiders. As for the distinct morphology of mygalomorph spiders I can refer to numerous
general text books in Arachnology (reference to some of which are to be found in the section
Literature) as well as various internet sites. Here only the main points of identification are
given.
Distinguishing Mygalomorphae from Araneomorphae
Mygalomorph spiders constitute a primitive branch of the spider family tree that is
distinguished from the true spiders (araneomorph spiders) by a number of conspicuous
morphological characters (Fig. 3). The main external differences between mygalomorph
spiders and araneomorph spiders are found in the anatomy of the biting organs, in the
organization of the respiratory organs and in the development of the spinning organs (Fig. 3).
The biting organs (chelicerae) in mygalomorph spider are orientated forward and act in a
parallel fashion whereas the chelicerae of araneomorph spiders project downward and act in
opposition as in pincers (Fig. 3).
12 Araneomorphae are cladistically indicated to be the sister group of Mygalomorphae
(Platnick & Gertsch 1976), nevertheless, and somewhat against the ‘rules of cladistics’
Goloboff (1993) chooses Mesothelae as outgroup for his cladistic revision of the
Mygalomorphae apparently just because this is more convenient.

The external respiratory organs of mygalomorph spiders are located on the anterior half of the
ventral abdomen as two pairs of book lungs (visible as roughly circular light colored patches).
Most araneomorph spiders have only the anterior pair of book lungs and a more posterior
located tracheal opening that is missing in mygalomorph spiders.
The external spinning organs (spinnerets) of mygalomorph spiders differ from those of other
spiders in the absence of anterior median spinnerets or their homolog, the reduction or
absence of anterior lateral spinnerets and the sub-segmentation of the basal segment of the
posterior lateral spinnerets (Raven 1985).
Table 2. All sixteen currently recognized orthognate spider families with notes on their principal
lifestyles after Jocqué and Dippenaar-Schoeman 2006. Note that in 11 out of 16 families the
production of silk-lined burrows is the dominant lifestyle. Regarding the 6 families that are not
primarily burrowing; three families (Theraphosidae, Dipluridae and Hexathelidae) are known to
contain at least some species that excavate burrows (personal observations) and only three are not
known to excavate burrows at all. These last three families ( Mecicobothriidae, Microstigmatidae and
Paratropididae) are also the families of which very little is known about their habits.
FAMILY LIFESTYLE
1 Actinopodidae Silk-lined burrow with trapdoor
2 Antrodiaetidae Silk-lined burrow with trapdoor or silk collar
3 Atypidae Silk-lined burrow with sock-like extension
4 Barychelidae Silk-lined burrow (retreat) with trapdoor
5 Ctenizidae Silk-lined burrow (retreat) with trapdoor
6 Cyrtaucheniidae Silk-lined burrow with trapdoor
7 Dipluridae Silk-lined retreat in crevices
8 Hexathelidae Silk-lined retreat in crevices
9 Idiopidae Silk-lined burrow with trapdoor
10 Liphistiidae Tubular burrow with trapdoor
11 Mecicobothriidae Silk-line retreat in crevices
12 Microstigmatidae Free-living encrusted with earth
13 Migidae Silk-lined burrow (retreat) with trapdoor
14 Nemesiidae Silk-lined burrow (retreat) with trapdoor
15 Paratropidae Cursorial in leaf litter
16 Theraphosidae Silk-lined burrow, arboreal retreat or cursorial
Fig. 3 Distinguishing Mygalomorphae
from Araneomorphae. A-C ventral views,
B-D frontal views. Differences in
morphology of chelicerae, book lungs and
spinnerets.

Chapter 2
Trapdoor spiders of the genus Nemesia Audouin 1826 on Majorca and Ibiza:
taxonomy, distribution and behaviour (Araneae, Mygalomorphae, Nemesiidae).
Arthur Decae
Research 1995-2005
Bulletin of the British arachnological Society (2005) 13(5): 145-168.

Abstract
Seven species of the trapdoor spider genus Nemesia (family Nemesiidae) have recently been
found in the Balearics, five on Majorca and two on Ibiza. Only one of these species N. brauni
(L. Koch 1882) had previously been named and described. Another species, N. bristowei sp.
n., had been reported from Majorca by Bristowe (1941, 1952), but it was never formally
described. The remaining five species are new. Here, supplementary notes and new figures
are given for N. brauni, and six new species are described and figured for the first time.
Additional information on the natural history, behavior and distribution of all seven species is
provided. The six new species are: N. bristowei, N. seldeni sp. n., N. randa sp. n. and N.
santeugenia sp. n., all from Majorca; N. ibiza sp. n. and N. santeulalia sp. n., from Ibiza. All
species are regarded as endemic to the Balearics.
The information on the individual species is preceded by a review of the morphology of
Nemesia at the generic level in order to discuss diagnostic characters for distinguishing
species.
Introduction
Among the Mygalomorphae, the genus
Nemesia is comparatively rich in species. A
survey of Platnick's (2003) catalogue shows
that Nemesia currently ranks fifth in a list of
303 mygalomorph genera (Table1). Large
genera usually have large areas of
distribution. The largest mygalomorph
genus, Idiops, occurs in Africa, Asia and
South America, and the second largest
genus, Aphonopelma, although concentrated
in North America, also has South American
representatives. According to Platnick's list,
Nemesia also has an almost cosmopolitan
distribution. The reality of this wide
occurrence, however, is biogeographically
difficult to explain, because of a curious
discontinuous distribution, with single
species recorded from China, Afghanistan, Mozambique and Cuba and approximately fifty
species reported from one relatively small geographical zone. In fact the distribution of
Nemesia is concentrated around the western, central, and parts of the eastern Mediterranean. It
is bordered by Alpine mountain ranges in the north, the Sahara desert in the south and the
Atlantic Ocean in the west (see Chapter 2, Fig. 5). In North Africa the genus is reported from
as Far East as Egypt, and in southern Europe as far east as Cyprus 13 . Nemesia is not known
from Anatolia, where it seems to be replaced by the related genus Brachythele Ausserer 1871.
The high species diversity of Nemesia in the restricted geographical zone around the western
and central Mediterranean might tentatively be explained by the combined effects of poor
powers of dispersal and strong allopatric speciation in an area that has been fragmented for
millions of years by tectonic activity and/or that has seen numerous relict populations formed
in an area that, in the not too distant past, has been affected by Pleistocene glaciations. The
reality of such speculation, however, remains to be investigated, but it could possibly explain
why so many, if not all, Nemesia species seem to be local endemics.
13 Not shown in Fig. 1.

The questions of whether these species really are local endemics and whether Nemesia
contains as many species as the catalogues suggest (Roewer 1942: 40 species, 6 subspecies;
Bonnet 1958: 37 species, 4 subspecies; Platnick 2003: 49 species, 4 subspecies) are of present
concern. Currently there are good arguments to believe that the lists of species are overestimations
of the real numbers, because species reported from Asia, southern Africa and the
Caribbean probably belong to different genera and revisions of Mediterranean species will
surely reveal synonymies. On the other hand the list of Nemesia species may yet increase
considerably in length because particularly the North African, Balkan, Greek, Spanish and
Italian faunas are still very incompletely known and, as this study shows numerous new
species can still be discovered.
Material and methods
In order to find the most useful morphological
characters for diagnosis at the species level, a
survey of about forty Nemesia species
(described and undescribed) present in the
author's collection and in the collection of
the MNHN in Paris was conducted.
This survey resulted in the present review of
the genus and in special attention being paid to
Genus name No. species No. subspecies
Idiops 90 1
Aphonopelma 90 0
Misgolas 61 0
Avicularia 54 2
Nemesia 49 4
Table 1 Ranking of the most diverse mygalomorph
genera according to Platnick 2003.
character variation in the morphology of leg IV, spinnerets, fangs, eye-formation and sexual
organs of the different species discussed.
The specimens reported on are housed, or will be deposited, in the following collections:
British Museum of Natural History (BMNH), London (type material of N. brauni); Museum
National d' Histoire Naturelle (MNHN), Paris (numbers starting with ARl4); Natural History
Museum (NMR), Rotterdam (numbers starting with 9972.40). The collection numbers are
given with the references to the material studied under the heading of each species discussed.
All spiders described here (with the exception of the male of N. brauni) were collected from
their natural burrows. Field data and burrow characteristics were noted in situ and parts of the
burrows (burrow entrance tubes with trapdoors intact and some burrow bottoms) or whole
burrows were collected for study in the laboratory. The samples so taken contained only adult
female and juvenile spiders. Some juvenile spiders from Majorca (often in their natural
burrows) were reared in captivity in an attempt to produce adult males (successful in only one
species, N. bristowei). Other Majorcan spiders, both adult and juvenile, were kept alive to
study their behavior. The spiders from Ibiza were not studied alive. Spiders used for
taxonomic research were killed in the deep-freeze compartment of a refrigerator at -21 °C and
preserved in 70% ethanol. These spiders were studied and drawn with the aid of a CETI-
MEDO 2 stereomicroscope equipped with an ocular micrometer, a drawing mirror and a cold
light source. All specimens were studied fully submerged in 70% ethanol, and fixed in
position by supporting them with insect-pins stuck in the polystyrene bottom of a small dish.
Measurements of body parts were taken by positioning that part horizontally with respect to
the microscope's objective and having both points of measurement simultaneously in sharp
focus (Figs. 2-5).
Three descriptive formulae are used:
1. Leg IV; summarizing the relative lengths of the metatarsus, tibia and femur (e.g. T4>
F4=M4, means that tibia IV is longer than both femur IV and metatarsus IV which are of
equal length). The leg segments were measured along the prolateral side of the right leg (see
Fig. 5).

2. PSP; summarizing the prolateral spine pattern on all patellae of a single specimen such as a
holotype: e.g. p=0-0, I=l-l, II=I-l, III=3-3, IV=l=0, means that there are no prolateral spines on
the palp patellae, one prolateral spine on patellae I and II (one left and one right), three spines
left and right on patellae III and one spine left and none right on patellae IV.
3. PSPvar; is used to describe the variation in the patellar prolateral spine pattern in a sample
of several spiders (e.g. paratypes): p=1(0-2), I=1, II=1(2), III= 1(0-2), IV=0, means that
usually there is one prolateral spine op the palp patella, but occasionally none or two spines,
that on patella I, invariably a single prolateral spine was observed, that patella II usually has
one prolateral spine, but occasionally two, etc.
Abbreviations are as follows: BL = total length body, CL = length carapace, CW = width
carapace, Ca = length cephalic part, Ch = height cephalic part, Th = thorax height, AR =
width anterior eye row, PR = width posterior eye-row, El = length eye-formation, Clyp =
height clypeus, ALE diameter anterior lateral eye, PLE = diameter posterior lateral eye, POP
= periocular pigmentation, M4 = length metatarsus IV, T4 = length tibia IV, F4 = length
femur IV, PSP = number of prolateral spines on patellae, PMS = posterior median spinnerets,
PLS = posterior lateral spinnerets.
Methods of measurement used are as follows: CL/CW = length carapace/ width carapace,
CL/Ca = length carapace/ caput length, Ch/Th = caput height/ thorax height, l/w =
length/width, AR/PR = width anterior eye row/ width posterior eye-row, AR/El = width
anterior eye row/ length eye-formation, ALE/PLE = diameter anterior lateral eye/ diameter
posterior lateral eye. All methods of measurement and description are shown in Figs. 2-8.
Measurements of body parts are in mm, and measurements of burrow parts are in cm.
Figs. 2-8 Measurements and abbreviations. 2 Body, dorsal: BL=body length, CL=carapace length, CW=carapace
width, Ca=caput length; 3 Carapace, lateral: Ch=caput height, Th=thorax height; 4 Eye-formation:
AR=width anterior row, PR=width posterior row, EI=eye-formation length, ALE=diameter anterior lateral
eye, PLE=diameter posterior lateral eye, Clyp=clypeus height; 5 Leg IV, prolateral: F4=femur IV length,
T4=tibia IV length, M4=metatarsus IV length; 6 Fang with smooth keel; 7 Fang with serrated keel; 8 Fang
with irregular keel.

Genus Nemesia Audouin, 1826
Nemesia is a genus of small to large Mediterranean trapdoor spiders (body length of adults 9-
31 mm). The generally dull brownish color, relatively long legs and distinctly recurved fovea
distinguish Nemesia species in the field readily from sympatric Ctenizidae and
Cyrtauchenidae that usually have a procurved fovea, are blackish in color, and are more
compactly built. Morphologically Nemesia species are not easily distinguished from each
other. Particularly the females of different species can be very similar in appearance, and
males also vary little in their anatomy. This inconspicuous anatomical variation has
undoubtedly contributed to the taxonomy of Nemesia having been confused virtually from the
start (Thorell 1870; Pickard-Cambridge in Moggridge 1874: 270-274). Another factor adding
to the confusion probably resulted from the different and personal styles of description that
authors have used (e.g. Ausserer 1871, 1875; Thorell 1875; Simon 1914; Franganillo 1920;
Frade & Bacelar 1931; Bacelar 1933). Modern descriptions of spider taxa usually follow a
more or less standard format that discusses the various body parts in a given order (carapace,
eye group, chelicerae, etc.) defining both qualitative (e.g. color, shape, pattern) and
quantitative (e.g. counts, measurements, ratios) characters. This modern method produces
improved prospects of finding character states of diagnostic value at all taxonomic levels. The
morphological character description of the genus Nemesia below focuses on finding such
characters of diagnostic value at the species level.
Species level diagnostic characters within Nemesia (Figs. 2-8):
Size: Larger and smaller bodied Nemesia species exist (in the field sometimes in close
proximity). Three different size-classes of Nemesia species are recognized here:
(a) Small sized species: adult BL ♂ 9-10, ♀ 11-17.
(b) Medium sized species: adult BL ♂ 13-14, ♀15-23.
(c) Large sized species: adult BL ♂ 17-18, ♀20-31.
Color: All Nemesia species are brownish in general appearance, but pigmentation of the
carapace, basal segment of the chelicerae, legs, palps, abdomen and spinnerets may show
species specific patterns. Particularly color differences between leg segments and the degree
of color contrast between the chelicerae and carapace may be of diagnostic value.
Carapace: The shape of the carapace and the degree of elevation of the head region (caput)
above the fovea may vary between species and is here reflected in the ratios CL/CW, CL/Ca
and Ch/Th (Figs. 2-3). Pubescence: In some Nemesia species the carapace, chelicerae, femora
and other leg-segments are clothed with a dense cover of fine pubescent hairs, in other species
these body parts may be devoid of pubescence. Nemesia males may differ in the possession of
"fringe setae", curved bristles on the edge of the carapace directed outwards. These fringe
setae may be found along the full length of both lateral sides of the carapace, only locally, or
they may be absent. Eye-formation: Variation in the shape of the eye-formation is reflected in
the ratios AR/PR and AR/El (Fig. 4). Variation in the position of the eye-formation relative to
the anterior edge of the carapace is expressed as Clyp (Fig. 4), a measure of the clypeus in
mm. Fovea: The shape of the fovea may vary from species to species and is here described as
curved, angular, with or without a central longitudinal groove, etc., with relevant illustrations
in the dorsal habitus drawings of the different species.
Fang: The inner surface of the fang carries a sharp longitudinal keel that may be either
smooth, irregularly notched or neatly serrated (Figs. 6-8). Cuspules: Usually, but not always,
present. When present they may form short single rows, two more or less parallel rows, or
irregular groups on the proximal margin of the maxillae. Sigilla: Three pairs of round or oval
sigilla may, or may not, be visible on the sternum; the anterior and median pairs usually touch
the sternum's margin opposite coxae I & II; the position and shape of the posterior pair may
be of diagnostic value. Scopulae: Always present in females on the palp tarsi and tarsi and

metatarsi I & II, but they may or may not extend onto the tibiae. In some species the typical
scopula-hairs are replaced by dense pubescence on the anterior tibiae, thus forming a distinct
pseudoscopula (Decae 1995). Spines: Descriptions of spine patterns are a major source of
confusion in Nemesia taxonomy. Owing to their conspicuous presence, spines and spine
patterns were used extensively as discriminative characters at the species level in early
literature. Usually these patterns were reported in the form of descriptions where figures
would have been less ambiguous. In more recent literature (Blasco 1986a; Cardoso 2000)
spine patterns in Nemesia have been regarded as taxonomically virtually useless, owing to
their extreme variability down to the individual level where different spine patterns are
frequently found on the left and right sides of a single spider. However, on certain faces of
some leg and palp segments, spine patterns may contain useful taxonomic information. Here
the prolateral spine formulae for all patellae are given (PSP, PSPvar), and those on patella III
and tibia III are figured (Figs. 17, 30, 37, 44, 51, 58, 65).
Leg IV: The relative lengths, measured
along the prolateral margin of the
metatarsus, tibia and femur, are of
diagnostic value at the species level and
are given in the ratios M4:T4:F4 (Fig. 5).
Metatarsal combs: The metatarsal
preening combs on legs III and IV that
were reported to be of diagnostic value in
Nemesia by Raven (1985: 96) are
unambiguously found only distally on
metatarsus IV in three Balearic species (N.
ibiza, N. randa and N. bristowei).
Spinnerets: The spinneret morphology of
Nemesia shows important, but so far
underrated, taxonomic characters for
species level taxonomy. The PMS may be
absent as in N. brauni (Fig.18), reduced,
having none or few apical spigots as in N.
bristowei (Fig. 31), or fully functional,
having spigots distributed over the distal
and ventral surface as in N. seldeni (Fig.
52). The PLS also show variation,
particularly in the spigot development on
the ventral surface of the basal segment.
Spigots may be absent, restricted to the
distal half of the segment (most species),
or distributed widely over the ventral
surface of the segment as in N. seldeni
(Fig. 52). Finally, the apical spigots on the
distal segment of the PLS may also show
important differences between species. All
species have a dense field of spigots at the apex of the PLS that are roughly arranged in
circular concentric rings of smaller spigots on the periphery around larger spigots more
centrally, with a few distinct "macro-spigots" in the centre. The number of macro-spigots can
be of diagnostic value. Spermathecae: The receptacles of the spermathecae provide a key
character for species identification in female Nemesia (see also Blasco 1986a). In general
three broad types of shape can be distinguished: unipartite with no distinct divisions between
Figs. 9-12 Nemesia brauni L. Koch, male. 9 body, dorsal; 10
distal end of right palp; retrolateral; 11 ditto, prolateral; 12 leg I
clasper, prolateral. Scale lines = 2mm (9), 1mm (10-12).

parts as in N. santeugenia (Fig. 46), bipartite with distinct proximal and distal parts as in N.
brauni (Fig. 19), and tripartite with a median part clearly separating the proximal and distal
parts as in N. ibiza (Fig. 67). Furthermore, the shape of the receptacles may differ from
species to species: the parts of the receptacles may lie in line (straight) as in N. bristowei (Fig.
32), the median part may be bent, or doubly bent (not seen in any of the Balearic species but
shown by Blasco 1986a for N. simoni O.P.-Cambridge 1874 (Fig. lb) and N. manderstjernae
Ausserer 1871 (Fig. 1g)), or the receptacles may be twisted in the median part as seen in N.
seldeni (Fig. 53). Finally, a third useful diagnostic character may be found in the density and
degree of coverage of the receptacles with glandular tissue. The whole receptacle may be
densely covered as in N. santeugenia (Fig. 46), coverage may be thin as in N. ibiza (Fig. 67),
or it may be locally dense and locally thin as in N. seldeni (Fig. 53) where the glandular tissue
is concentrated proximally on both receptacles. Clasper: The structure of the clasper on the
tibia and metatarsus I of Nemesia males may vary somewhat from species to species.
Particu1arly the ventral "clasper field" of short stiff hairs and cuspules on the metatarsus
(Figs. 12, 25) can be of diagnostic value.
Bulb: The palpal bulb of Nemesia males shows little variation in the proximal parts; the shape
of the embolus and its ornamentation (or lack of it) with denticles, combs or ridges on its tip,
however, is of important diagnostic value at the species level.
Other characters: Nemesia species are remarkably similar in most other features of their
morphology, and the clearest diagnostic characters are typically found in burrow structure,
trapdoor construction and in other aspects of behavior and distribution (Moggridge, 1873,
1874). Available information on these characters is given following the morphological species
descriptions.
Nemesia brauni L. Koch 1882 (Figs. 9-19, 68, 75, 79, 85)
Nemesia braunii L. Koch 1882: 642, p1. 20, fig. 21 (D ♂♀).
Nemesia brauni: Simon 1892: 113; Reimoser 1919: 6; Frade & Bacelar 1931: 226, Figs. 7-8 (♂).
Type: Material in the BMNH examined by P. Hillyard (pers. comm.).
Other material examined: Majorca: 1♂ in E. Simon's collection at the MNHN (labeled N.
brauni L.K. Palma AR 4491). 8 ♀♀, leg. A. E. Decae: 1♀, Porto Sóller, 39.783°N, 2.663°E,
13 September 1997 (AR 14191); 1♀, Arta, 39.725°N, 3.325°E, 16 September 1997
(9972.4004); 1 ♀, Santa Maria, 39.691°N, 2.713°E, 10 April1997 (AR 14193); 1 ♀, Santa
Eugenia, 39.640°N, 2.830°E, 29 March 2000 (9972.4002); 2 ♀♀, roadside between Inca and
Puebla, 39.766°N, 2.992°E,13 October 2000 (9972.4003; AR 14192); 1 ♀, roadside near
Campos, 39.415°N, 2.998°E, 16 April 2002 (9972.4005); 1 ♀, Massis de Randa, 39.534°N,
2.925°E, 18 April 2003 (AR 14190).
Diagnosis: Nemesia brauni differs from all other described Nemesia species by the absence of
PMS and the presence of a row of three tiny denticles just proximal to the embolus tip (Fig.
10). The absence of PMS in N. brauni has been confirmed by a study of the type material,
BMNH collection, by P. Hillyard (pers. comm.).
Notes: Machado 1944 reported the absence of PMS for N. hispanica L. Koch 1871, but a
check of the type material of this species in the BMNH by P. Hillyard (pers. comm.) revealed
that the PMS, although reduced, are present in N. hispanica. At least two other Nemesia
species lacking PMS are known from the Iberian Peninsula, but these have not yet been
formally described. These two species differ from N. brauni, and from each other, in the
morphology of their sexual organs and it might be argued that the absence of PMS is
sufficient to group them with N. brauni in a separate genus. Further study of probable
phylogenetic relationships between these species is necessary however, before revision of the
genus Nemesia can usefully be attempted.

Description: L. Koch 1882 described both sexes and figured the male palp. Koch's
descriptions are elaborate and accurate, but lack detailed information on the morphology of
the embolus, clasper, fang structure, spermathecae and spinnerets that is of diagnostic
importance. Supplementary information on these aspects is provided here, as well as
measurements of some body parts and figures that are considered to be of diagnostic value.
Figs. 13-19 Nemesia brauni L. Koch, female. 13 habitus dorsal; 14 habitus ventral; 15 carapace, lateral; 16
eye-formation, dorsal; 17 patella and tibia III, prolateral; 18 spinnerets, ventral (note absence of PMS); 19
spermathecae, dorsal. Scale lines = 2mm (13-15) 1mm (16-18).
Male (n= 1): BL = 15.6, CL = 6.6, CW = 5.4. Leg IV: M4>T4>F4. PSP: p = 1-1; I = 2-2; II =
1-2; III = 2-2; IV = 1-1. Body dorsally as in Fig. 9. Carapace longer than wide, CL/CW = 1.2.
Similar to female in distinct morphological characters such as large size, absence of PMS,
relatively wide clypeus (Clyp =0.39), shape of eye-formation (AR/PR = 0.96; AR/El = 2.00),
and presence of lighter longitudinal zones on basal segment of chelicerae clothed with silvery
white pubescence. The male studied differs from the females in having a low caput, an
irregular serrated fang ridge (Fig. 8), and presence of dorsal spines on metatarsi and tibiae I
and II. It differs also in relative lengths of segments in leg IV and prolateral spine pattern on
patellae. Embolus (Figs. 10-11) neither shortened nor elongated; tip sigmoid, curved, and
furnished with three tiny denticles (Fig. 10a), evident when embolus examined under high
magnification in retrolateral view. Clasper as in Fig.12.
Female (n=8): BL = 20-26, CL= 7.3-9.2, CW =6.2-7.9. Leg IV: T4>F4>M4. PSPvar: p = 0; I
= 0; II = 0; III = 0; IV = 0. Dorsal aspect (Fig. 13). Carapace slightly longer than wide,
CL/CW = 1.1-I.2, caput strongly elevated (Fig. 15), Ch/Th = 2.2-2.7. Clypeus wide, Clyp =
0.43-0.65. Eye-formation (Fig. 16): posterior row slightly longer than anterior row, AR/PR =

1.01-1.08, more than twice as wide as long, AR/EI = 2.08-2.39, grouped on and around a
sloping rather than steep ocular tubercle; anterior laterals larger than posterior laterals,
ALE/PLE = 1.05-1.31. POP broken between median eyes and between median and lateral
eyes, PME clearly smallest. Fang ridge smooth (Fig. 6). Leg III: one very strong retrolateral
spine on tibia III; no prolateral spines on tibia III and patella III (Fig. 17). Ventral aspect (Fig.
14). Spinnerets (Fig. 18): PMS absent, PLS with spigots on ventral surface of basal segment
restricted to distal half. Spermathecae (Fig. 19): bipartite, short "mushroom" shaped, evenly
covered with dense glandular tissue.
Burrow (Fig. 68): Nemesia brauni digs a burrow that is closed at the entrance by a thick,
plug-like trapdoor (typical cork-door, as illustrated in Moggridge (1873: pl. 8) and here in Fig.
79). The depth of the burrow seems to vary with soil conditions and underground obstacles
such as stones and roots. Some adult spiders were found in shallow burrows, 10-13cm deep,
in stony and very compact soils, but generally burrows of N. brauni are between 17cm and 25
cm deep. The burrow shaft is a simple, unbranched tube with a diameter of 1.2-2.0cm in adult
females. The burrow-walls are compacted and apparently impregnated with a liquid excretion
from the oral opening (observed in captive spiders) to form a hard plaster wall. This hard
plaster can be used to construct burrow tubes in hollow spaces underground. That the spiders
can construct their nests in this way is shown when they build hard clay cells from loose soil
material packed with the spiders in containers for transport. Silk is used only sparsely in
burrow construction by N. brauni. Only the uppermost parts of the burrow-wall are covered
with a thin sheet of silk that extends upwards to form a narrow, ribbon-like "hinge" that
connects the burrow lining with the much thicker silken cover of the under surface and
bevelled edges of the trapdoor.
Behavior: The adult N. brauni female is an aggressive spider that will vigorously strike at any
object (or organism) entering the burrow. During daylight hours, however, the spiders usually
react to disturbances by running up the burrow and pulling the trapdoor firmly closed by
anchoring the claws of the first two pairs of legs in the silken sheet that covers the underside
of the trapdoor. Around dusk N. brauni spiders come to the entrance of their burrow to lie in
wait for passing prey. At first they just open their trapdoors by a small crack like most other
trapdoor spiders, but with the growing darkness they come out more and more, until finally
the trapdoor is wide open and the spiders sit fully exposed in the burrow entrance (Fig. 75).
From this position the spider strikes at, and may even pursue, prey within a range of up to at
least 10 cm around the burrow entrance. After a prey is captured away from the burrow, the
spider will pause for a few seconds before picking up the prey with the fangs and supporting it
with the palps. Apparently it has a "memory" for the location of the burrow, because usually it
will turn around to carry the captured prey straight back to the burrow. If the trapdoor has
remained open during the spider's actions it descends, head first, into the burrow tube and
then turns around and closes the door with its front legs. Often, however, the vigorous action
of the spider rushing from the burrow in pursuit of prey causes the door to fall back, closing
the burrow entrance behind the hunting spider. On its return to the closed burrow the spider
sometimes has some difficulty in opening the trapdoor from the outside. If the spider does not
succeed in opening the door with its front legs it will turn around to lift the door with the
claws of the fourth leg before reversing back into its burrow. This action shows that the
behavior of a trapdoor spider is not necessarily as stereotyped as sometimes supposed.
When it captures a large prey, the spider will remain in the closed burrow for many hours,
presumably feeding. When the capture is a small prey the spider may be seen in the ambush
position again ready to capture a second prey while still eating the first.
Distribution (Fig. 85): Nemesia brauni appears to be the most common trapdoor spider on
Majorca. It has a wide distribution on the island, but it appears to be rare in the far southeast
of the island, where it seems to be largely replaced by N. randa (Fig. 84). Koch 1882 reported

N. brauni also from Minorca. Its presence there has not been checked in the course of this
study.
Nemesia brauni may be found in various habitats and situations and occurs locally in close
association with all other Majorcan Nemesia species. It tends to be present in more exposed
locations and occurs more frequently in horizontal surfaces than other Nemesia species on
Majorca, although numerous burrows of N. brauni were also found in steep and vertical
surfaces, both on natural slopes and in clay fillings between rocks of man-made stone walls.
The species was found inland up to an altitude of over 800 m, but also at sea level no more
than 2 m from the flood line.
Nemesia bristowei sp. n. (Figs. 20-32, 69, 76, 80, 84)
Types: Holotype ♂ (AR 14208), 17 September 1997, Majorca, Porto Só11er, 39.793°N,
2.673°E, burrow in steep clay-bank along a road between farmland. Paratypes: Majorca: 1♂
(9972.4010), same data; l ♂ (9972.4009), same locality, 11 September 1997; 3 ♀♀
(ARI4205; 9972.4007; 9972.4006), same locality, 11,17, 20 September 1997 respectively;
3♀♀ (9972.4008; AR14206-7), 2 April 1996, Esporles/LaGranja, 39.656°N, 2.576°E,
burrows in steep roadside bank between pine forest and farm fields. All leg. A. E. Decae.
Males were collected as juveniles and reared in captivity.
Etymology: The species is named in honor of W. S. Bristowe who discovered the species first
on Majorca near Genova, 39.56°N, 2.60°E, in January 1930 and published about it in 1941
and 1952.
Diagnosis: Nemesia bristowei seems to be closely related to N. randa, with which it shares
the characteristic forwardly directed ALE (Figs. 29, 36) that seems to be unique to these two
Majorcan endemics. It differs from N. randa, however, in its smaller adult size, the glabrous
carapace and the presence of prolateral spines on patella III. It differs from all other Nemesia
species by the construction of a remarkable, cog-wheel shaped trapdoor and the regular
molded counter shapes in the burrow margin that neatly receive the teeth of the cog-wheel
when the door is closed (see Bristowe 1941: 257, fig. 26 and the photographs presented in
Figs. 76, 80).
Description: Male (holotype): BL = 10.3, CL = 3.9, CW = 3.3. Leg IV: F4=M4>T4. PSP: p =
l-l; I = 2-2; II = 2-1; III = 2-2; IV = 0-0. Dorsal aspect (Fig. 20). Carapace slightly longer than
wide, CL/CW = 1.2, glabrous, light yellowish brown, caput elevated (Fig. 22) and darker in
color than thorax, Ch/Th = 1.4, few setae on crest of caput and on clypeus, fringe setae
restricted to posterior margins of carapace. Clypeus narrow, Clyp = 0.10. Eye-group less than
twice as wide as long, AR/El = 1.88, PR slightly longer than AR, AR/PR = 0.96, ALE much
larger than PLE, ALE/PLE = 1.43, and characteristically forwardly orientated. Ocular tubercle
steep in front sloping behind. Fovea deep, smoothly recurved. Chelicerae brown, darker than,
and contrasting with carapace, stiff spiny setae in longitudinal group on dorsal crest and finer
setae in narrow lateral line, pubescence absent. Promargin of cheliceral furrow with six teeth,
all somewhat spaced apart, second and fifth teeth largest. Rastellum with two strong teeth
apically, standing out in field of smaller rastellar teeth. Fangs with neatly serrated keel (Fig.
7) along promargin. Ventral aspect (Fig. 21). Maxillae slightly darker than sternum and
ventral surface of coxae, longer than wide (l/w = 1.8), with few cuspules near prolateral
proximal margin. Sternum light yellow, slightly longer than wide (1/w = 1.3), evenly covered
with black setae. Anterior and median pairs of sigilla round, touching sternum margin,
posterior pair placed slightly away from margin. Labium greyish, dome shaped, wider than
long, evenly covered with fine setae, proximal setae strongest. Labial furrow wide and
glabrous. Legs lighter yellow than carapace, with numerous spiky spines on metatarsi, tibiae,

and femora; all metatarsi and tibiae with dorsal, ventral pro- and retrolateral spines, prolateral
spines on patellae I and II, pro- and retrolateral spines on patella III, no spines on patella IV,
numerous short dorsal and dorso-lateral spines on all femora, spine patterns differ on left and
right legs. Dense pubescence on all femora. Palps (Figs. 23-24): color and pubescence as legs,
dorsal apical spine-group on tibia with two spines left and three right, cymbial spines
Figs. 20-25: Nemesia bristowei sp. n. male. 20 habitus dorsal; 21 habitus ventral; 22 carapace lateral; 23
distal end of right palp, prolateral; 24 distal end of left palp, retrolateral; 25 left leg I clasper, prolateral.
Scale lines = 2mm (20-22), 1mm (23-25).
restricted to distal half of segment. Abdomen light greyish brown with vague dorsal pattern of
darker blotches, fully covered with fine hairs. Spinnerets (as in female, Fig. 31) creamy
yellow. PMS small, spiky, with one apical spigot. PLS basal segment without spigots, as long
as median and distal segments together. Apical spigots grouped around one large central
spigot. Bulb (Figs. 23-24): embolus long, slender and slightly bent. Tibial spur (Fig. 25)
inwardly curved. Ventral clasper field on metatarsus I with short sharp spikes not closely
grouped.
Variation (n=3): Males vary considerably in size, BL = 9-10.5, CL = 3.2-3.9, CW = 2.8-3.3.
Shape of carapace rather constant, CL/CW = 1.2. ALE/PLE = 1.36- 1.55, AR/PR = 0.94-0.96.
Fovea varies from smoothly recurved to almost straight. Slight variation was found in relative
lengths of femur IV and metatarsus IV, F4/M4 =1.00-1.02. Most conspicuous variation in
spine patterns, which differ considerably on left and right appendages of all individuals and
between all spiders studied. PSPvar: p = 1(0); I = 2(1); II = 2(1); III = 2(1); IV = 0(l). Number
of macro-spigots on distal segment of PLS may vary from 1-3.
Female (n=6): BL = 12-17, CL = 4.8-5.7, CW = 3.8- 4.6. Leg IV: F4>T4> M4. PSPvar: p =
1(0-2); I = 1; II = 1(2); III = 1(0-2); IV = 0. Dorsal aspect (Fig. 26). Larger and more
compactly built than males. Carapace color pattern as in male, CL/CW = 1.2. Caput (Fig. 28)
elevated, Ch/Th = 1.7-1.9. Clypeus variable in width, Clyp = 0.12-0.24. Ocular tubercle as in
male. Eye- formation (Fig. 29), AR/PR = 0.92-1.00, POP unbroken, ALE directed forwards
and larger than PLE, ALE/ PLE = 1.25-1.62. Fovea recurved, sometimes somewhat extended

laterally. Chelicerae strong, color and setae as in male. Rastellum more strongly developed
than in male. Fang keel serrated (Fig. 7) or rarely smooth (in one specimen). Ventral aspect
(Fig. 27).
Maxillae with small anterior apical process, cuspules well developed, in irregular rows.
Sternum (1/w =1.4-1.6), posterior sigilla oval, about their largest diameter from sternum edge,
anterior and median sigilla hardly visible. Labium with anterior edge slightly convex. Legs
with very strong spines on anterior metatarsi and tibiae, scopula restricted to tarsi and
metatarsi (no pseudoscopula on tibiae). One or two prolateral spines on patella III (Fig. 30a,
Fig. 26-32. Nemesia bristowei sp. n., female. 26 habitus dorsal; 27 habitus ventral; 28 carapace, lateral; 29
eye-formation, dorsal; 30 patella and tibia III, prolateral (note variation in spine pattern right (a) and left (b);
31 spinnerets, ventral (note vestigial PMS); 32 spermathecae, dorsal. Scale lines = 2mm (26-28), 1mm (30),
0.5mm (29, 31-32).
b), rarely none. Retrolateral apical comb on metatarsus IV not evident in all specimens. Palp
with numerous strong spines and spine pairs on both lateral sides of tibia; scopula restricted to
tarsus. Abdomen light greyish brown with vague, irregular dorsal pattern of darker patches.
Spinnerets (Fig. 31) creamy yellow, PMS vestigial with only one or two fine apical spigots,
conical, PLS apical field spigots grouped around 1-3 macro-spigots. Spermathecae (Fig. 32)
bipartite, slender "mushroom" shape (proximal part of receptacle tubular, distal part globular),
evenly covered with thin glandular tissue.
Burrow (Fig. 69): The burrow of N. bristowei is immediately recognizable by the peculiar,
intricate shape of the trapdoor and burrow rim (Fig. 80). Bristowe (1941: 257) first found and
illustrated this "cogwheel- door". Bristowe 1952 reported that he had not been able to describe
the species, because his collection was destroyed during World War II. It took nearly half a
century before Selden 1997 rediscovered this remarkable trapdoor spider in 1989. The tube of
the burrow has no special features, but is a narrow cylindrical hole (diameter 0.6-0.9cm) about
8-15cm deep, that often widens somewhat near the bottom. The largest diameter of the
trapdoor is about 1.15cm. The walls and bottom of the tube are lined with a dense sheet of
silk. In this respect it is similar to the burrow of N. randa (Fig. 70), but different from that of
N. brauni (Fig. 68), in which only the upper parts of the burrow are lined with silk.
Behavior: Adult N. bristowei females are small, rather shy spiders that have a much more
defensive attitude than N. brauni. The peculiar trapdoor, the construction of which was
described by Bristowe (1941: 257), seems to function to improve the closing of the burrow.

The "teeth" on the edge of the door fit closely between the notches on the burrow rim (Fig.
80), providing what seems to be a "locking" system. This system of closure appears to
function in keeping the door shut on the vertical or even slightly overhanging surfaces in
which these spiders frequently build their burrows. When disturbed, N. bristowei, as all
trapdoor spiders so far observed, runs up to the trapdoor to pull it shut. The spider reacts to
excavation of the burrow by retreating to the bottom of the burrow, where it sits still with its
legs drawn close to the body (Fig. 69). Only when taken from the burrow will it try to escape
by suddenly dashing away with surprising speed. In captivity the spiders will, given the
opportunity, readily construct a new burrow both in sloping and horizontal soils. Shortly after
a spider has constructed a new trapdoor it may be found hunting by lying in ambush behind an
only slightly cracked open door (Fig. 76). The tips of tarsi I and II rest against the inside of
the notches on the burrow rim, but no part of the spider extends from under the trapdoor. The
spider will react aggressively to small prey passing the burrow within a distance of less than
1cm. Small isopods, with body lengths up to about half the body length of the spider, are
readily captured in the usual trapdoor spider fashion, by a quick and aimed dash forwards.
Small beetles, flies and ants will also be captured and eaten by the spiders in the laboratory.
The natural prey of N. bristowei is difficult to establish because this species, as all other
Majorcan Nemesia species, removes indigestible remains of eaten prey from the burrow by
"throwing" them out of the burrow entrance. In the laboratory these remains, compressed into
compact little balls, are invariably found in the container in which the spider is kept, some
hours after prey is captured. That the spiders are inclined to keep their burrows clean is also
evident from the traces of defecation usually found on the cover of the container some
centimeters above the trapdoor. Apparently these spiders eject feces with some force from the
burrow opening.
Annual cycle: The males of N. bristowei described here were collected as juveniles in
September 1997. They left their burrows as adults (in captivity) one year later in September
1998. Because this period of the year coincides with the peak in male wandering of Nemesia
in the field (121 records between 1 September and 30 November in pitfall trap catches of
males in Portugal, Spain, France and Italy) it is supposed here that males of N. bristowei
wander predominantly in autumn. They would do so after a period of aestivation in summer
when the burrows of most Mediterranean trapdoor spiders are found to be "locked" with silk
spun around the inner edges of the trapdoor to seal it to the burrow rim and/or by a thick clay
plug positioned directly under the trapdoor. The last feature has not been observed in N.
bristowei. In spring most females show development of eggs in the ventral part of their
abdomen. Egg-sacs and spiderlings may be found in the burrows of females between May and
September. It seems that N. bristowei does not practice extended brood care as do some
Nemesia species in southern France.
Distribution (Fig. 84): The distribution of N. bristowei seems to be restricted to the western
half of Majorca, where the species is very common in the mountain range of the Serra de
Tramuntana. It seems to be rare anywhere east of the line Palma-Pollença. Within its area of
distribution N. bristowei seems to be a specialist of steep surfaces. Although burrows are also
found on gradually sloping banks, the densest aggregations of burrows are found on nearly
vertical or even overhanging surfaces, often of eroding clay banks. The spiders may also be
found within human settlements in garden walls, parks and even in the walls of houses.
Nemesia randa sp. n. (Figs. 33-39, 70, 78, 82, 84)
Types: Holotype ♀ (AR14198), 2 Apri12002, Majorca, Castelix de la Pau, 39.550°N,
2.898°E, burrow in steep bank of white clay within patch of pine forest. Para-types: Majorca:
3 ♀♀ (AR14199; AR14200; 9972.4011), 1 April 1998, between Randa and Monturi,

39.539°N, 2.974°E, burrows in steep clay-bank along road between mixed forest and
cultivated fields; 1 ♀ (AR14201), Randa, 39.531°N, 2.904°E, 8 April 1995, burrow in steep
clay bank in garigue; 2 ♀♀ (9972.4012; 9972.4013), between Llucmajor and Porreres,
39.503°N, 2.972°E, 5 April 1995 and 18 April 2002, burrows in steep roadside bank in mixed
forest. All specimen leg. A. E. Decae.
Etymology: The specific name is a noun in apposition taken from the geographical area, the
Massis de Randa, where it was first discovered.
Diagnosis: Nemesia randa differs from all other known Nemesia species, with the exception
of N. bristowei, in the relatively large size and forward orientation of the ALE and the narrow
clypeus (Fig. 36). It differs from N. bristowei by the larger size of adult female spiders and
other characters mentioned in the diagnosis of N. bristowei. The male is unknown.
Description: Female (holotype): BL = 20.6, CL = 7.6, CW = 6.0. Leg IV: T4> F4> M4. PSP:
p = 2-2; I = 1-1; II = 1-1; III = 0-0; IV = 0-0. Dorsal aspect (Fig. 33). Carapace longer than
wide, CL/CW=1.3, different shades of brown and yellow in suffused, indistinct pattern of
darker central zones and lighter lateral zones, cervical grooves distinctly dark brown, setae in
three longitudinal, somewhat irregular parallel rows, with strongest setae in central row and in
small groups on ocular tubercle and along anterior edge of clypeus. Carapace indistinctly, but
fully, covered with fine whitish pubescent hairs. Caput elevated, Ch/Th = 2.0. Clypeus
narrow, Clyp = 0.12. Eye-group (Fig. 36) on steep ocular tubercle, less than twice as wide as
long, AR/El = 1.69, anterior row slightly longer than posterior row, AR/ PR = 1.04, ALE
orientated forward and much larger than other eyes, ALE/PLE = 1.57. POP unbroken. Fovea
not smoothly recurved, but somewhat angular, dropping away from base of caput. Chelicerae
dark chocolate brown, contrasting with yellowish carapace, broad glabrous zones between
dorsal setae-field and lateral zones with fine hairs. Dorsal setae merging distally with field of
stronger spines surrounding rastellum teeth. Cuticle of cheliceral furrow distinctly lighter in
color, pro-margin of furrow with seven strong teeth, second and third (counting from fang
base) largest, two proximals (6th and 7th) somewhat separated. Rastellum on small process
distal and prolateral of fang base. Fangs rather short and blunt, with serrated keel (Fig. 7).
Legs yellowish brown, slightly lighter than carapace; ventral and lateral surfaces of femora III
and IV, ventral surfaces of patella I and tibiae I and II conspicuously lighter creamy white, all
femora with dark central longitudinal zone dorsally. Scopulae strongly developed on tarsi and
metatarsi I and II, extending slightly onto distal ends of tibiae I and II. Strong prolateral
ventral spines in longitudinal rows on metatarsi and tibiae I and II, also more dorsally on
metatarsus III, tibia III with two short prolateral spines (Fig. 37), spines on metatarsus and
tibia IV thinner and fewer than on those segments of other legs. Slender spiny setae in
longitudinal rows and distal groups dorsally on all femora. Metatarsus IV with retrolateral
apical comb. Palp similar color to anterior legs, scopula on tarsus not extending onto tibia,
one pro- and one retrolateral proximal spine on tarsus (left pro-lateral spine absent), fine
ventral spines in tarsal scopula, rows of strong pro- and retrolateral spines on tibia, two
prolateral spines on patella, a larger spine more ventrally and a smaller spine more dorsally
placed, femur as legs. Ventral aspect (Fig. 34). Maxillae longer than wide (l/w = 1.8),
extending into indistinct prolateral distal process, slightly darker than coxae and sternum,
except for creamy white anterior zone of maxillary scopula. Ventrally evenly covered with
setae; well developed row of cuspules along proximal margin. Sternum longer than wide (l/w
= 1.2), widest between coxae II and III, yellow, evenly covered with fine setae of varying
size, strongest setae along edges, three pairs of sigilla, anterior and median pairs touching
margin, posterior pair more centrally placed (twice their diameter from sternum edge).
Labium slightly darker than sternum, dome shaped, wider than long, evenly covered with
setae. Labial furrow wide, glabrous, distinctly bicolored, yellowish along sternum, more
greyish along labium. Abdomen greyish, evenly covered with fine setae, faint dorsal pattern
of darker lines and chevrons. Spinnerets (Fig. 38) similar color to ventral abdomen; PMS

Fig. 33-39: Nemesia randa sp. n., female. 33 habitus dorsal; 34 habitus ventral; 35 carapace lateral; 36 eyeformation,
dorsal; 37 patella and tibia III, prolateral; 38 spinnerets, ventral; 39 spermathecae dorsal. Scale lines
= 2mm (33-35), 1mm (36-38), 0.5mm (39).
reduced, spiky, with two apical spigots, PLS short, basal segment longer than median and
distal segment together, ventral spigots only on distal half of basal segment, apical spigots
grouped around three macro-spigots. Spermathecae (Fig. 39) bipartite, proximal part conical,
distal part globular, both parts evenly covered with not very dense glandular tissue.
Variation (n = 7): Females are medium to large trapdoor spiders, BL= 18-24, CL=6.8-8.4.
Carapace, CL/CW=1.2-1.3, CL/Ca=1.7. Caput Ch/Th=1.7-2.2. Clyp=0.07-0.20. ALE/PLE=
1.26-1.55. AR/PR= 1.03- 1.08. Fang ridge may be either neatly serrated (Fig. 7) or more
irregular (Fig. 8). PSPvar: p = 1 (2), I.= I, II = I, III=0(l), IV=0. Leg IV: F4>T4≥M4. PMS
reduced, 2-3 apical spigots. PLS apical spigots grouped around 2-3 macro spigots. Male:
unknown.
Burrow (Fig. 70): The first impression of the burrow of N. randa in the field is that of a
classical cork-burrow as first described in detail by Moggridge 1873 for N. carminans
(Latreil1e 1818) in southern France, and as reported here for N. brauni from Majorca and N.
ibiza from Ibiza. On closer inspection, however, there are some interesting differences. The
silken lining of the burrow tube is not restricted to the upper parts of the burrow, as it is in all
three species mentioned above, but it extends all the way down the walls of the tube and also
covers the bottom of the burrow (Fig. 70, dashed line). The trapdoor of N. randa also differs
from that of the classical "cork-type" in that it not only fits snugly into the entrance opening
of the burrow, but it also extends over the edges of the burrow opening, resting on some
irregular extensions (tags) constructed on the upper side of the door (Fig. 82). In this respect
the trapdoor of N. randa is somewhat like the "cog-wheel" door of N. bristowei, although not
nearly so regularly sculptured. Another similarity between the trapdoors of N. randa and N.
bristowei, in which both differ from a typical cork-door, is the width of the hinge. In a typical

cork- door, as built by the three above-mentioned species, the hinge is a narrow, ribbon-like
silken strap that allows the door, when open, to rotate slightly. The trapdoors of both N. randa
and N. bristowei, however, have a much broader silken hinge that limits the door to simple
straightforward opening and closing movements. Like the burrow of all other species
mentioned in this paragraph, the burrow of N. randa is a simple tube without side
excavations.
Behavior: In the field the burrows of N. randa may be found on rather horizontal and
gradually sloping soil surfaces but, like N. bristowei, it is most frequently found on very
steep or vertical clay walls. Particularly where N. randa is found syntopically with N. brauni
there seems to be some habitat partitioning in which N. brauni predominantly occupies the
more horizontal ground and N. randa the steeper surfaces. A curious observation, that so far
has no clear explanation, is that whereas most trapdoor spider species orient their trapdoors
in the same position with respect to the slope of the terrain, N. randa orients its door
differently. In most trapdoor spiders the hinge connects the trapdoor to the highest (most
uphill) point of the burrow entrance and the door opens widely facing downhill, but in N.
randa the trapdoor almost invariably opens sideways with respect to the slope, or even uphill
(also observed though less frequently in N. bristowei). In temperament N. randa is a rather
un-aggressive spider that will bite only as a last defense after capture. It is also "shy" in its
hunting posture, which is essentially similar to that described for N. bristowei (Fig. 78). The
prey capture range of N. randa, owing to the spider's larger size, extends to a somewhat
greater distance of about 2 cm from the burrow rim. Prey handling and the ejection of preyremains
and feces from the burrow are as described for N. bristowei. In captivity N. randa
was found to be more reluctant to construct a new burrow than the other Majorcan Nemesia
species. Some spiders wandered around their clay-filled containers for weeks before they
finally started digging, and some never dug. When the spiders finally constructed a burrow, it
was indistinguishable from a natural burrow, and usually the spiders were found hunting
normally the night after the construction work was finished. In captivity N. randa would take
a wide range of arthropod prey, but none larger than about half the size of the spider. As N.
bristowei, and in contrast to N. brauni and N. seldeni, N. randa was never seen to leave the
burrow in pursuit of prey.
Distribution (Fig. 84): Nemesia randa was found to have a restricted distribution on Majorca,
where it is common around the central massif of the Massis de Randa and in the south-eastern
parts of the island. It inhabits similar habitats to N. brauni, with which species it is sometimes
found in close association.
Nemesia santeugenia sp. n. (Figs. 40-46, 71, 86)
Types: Holotype ♀ (AR 14197), 1 April 1997, Majorca, Santa Eugenia, 39.611°N, 2.876°E,
burrow in horizontal ground on edge of pine growth. Paratype: 1 ♀ (9972.4014), Majorca,
Bon Ani, 39.590°N, 3.082°E, 16 October 2000, burrow in sloping ground on hillside pine
forest. Both leg. A. E. Decae.
Etymology: The specific name is a noun in apposition taken from the village Santa Eugenia,
in the vicinity of which it was first found on Majorca.
Diagnosis: Nemesia santeugenia is a medium to large sized Nemesia species from Majorca
that shares a characteristic type of spermathecae with N. santeulalia sp. n. from Ibiza (cf.
Figs. 46 and 60). The spermathecae are compact (without a clear division between proximal
and distal parts), short, more or less "potato- shaped" and densely covered with glandular
tissue over their total surface. In their external morphology N. santeugenia and N. santeulalia
are also very similar, although some differences were observed in the relative sizes of ALE
and PLE (ALE/PLE= 1.2-1.4 (n=2) for N. santeugenia and ALE/PLE=0.9-1.2 (n=6) for N.

santeulalia), the broken POP in the first species that is not seen in the second, and the
difference in reduction of the PMS (cf. Figs. 45 and 59). The limited material of N.
santeugenia available for study and the absence of males of both species, however, make the
morphological distinction between these species problematic. The main argument to regard N.
santeugenia distinct from N. santeulalia rests on conspicuous differences in their behavior
and burrow construction as observed in the field. While N. santeugenia was aggressive and
ready to escape as a reaction to disturbance, N. santeulalia remained passive and tried to hide
in a dead-ended side tube of the main burrow (Fig. 73). In N. santeugenia the side tube
reaches the surface (probably closed off only by a thin silk sheet, Fig. 71), while in N.
santeulalia it does not reach the surface. Another remarkable difference in behavior is that N.
santeulalia retains indigestible remains of prey packed in the bottom and lower walls of the
burrow (Fig. 73), but such storage of debris was not found in the burrows of N. santeugenia
(Pig. 71). The male is unknown.
Note: Blasco (1986a: 347, fig. 2F) shows comparable spermathecae for N. ariasi Simon 1914
that were reproduced "après Buchli". A check of an unpublished manuscript by Buchli (1968:
79, fig. 44E) that shows these spermathecae suggests that Blasco was mistaken, because
Buchli attributes this type of spermathecae not to N. ariasi, but to N. hispanica. Buchli must
have been confused about the identity of N. hispanica because, in the same manuscript, he
shows N. hispanica (p. 79, fig. 43C) as having no PMS. A check of the type material of N.
hispanica in the BMNH by P. Hillyard (pers. comm.) has shown that this species has PMS. In
the course of the present study (of around forty different species), no other species except N.
santeugenia and N. santeulalia have been observed to have the described type of
spermathecae; the spermathecae of N. ariasi have not been studied because the female of this
species is unknown (Roewer 1942).
Description: Female (holotype): BL = 23.5, CL = 8.2, CW = 6.3. Leg IV: T4>P4≥M4. PSP: p
= 3-3; I = 2-2; II = 2-2; III = 2-2; IV = 1-0. Dorsal aspect (Fig. 40). Carapace longer than
wide, CL/CW = 1.3, brown, caput elevated (Ch/Th = 8.8) and slightly lighter brown than
thorax, no clear pattern, stronger setae in line on caput crest, fine setae on anterior clypeus
edge, dense white pubescence on most of carapace. Clypeus wide, Clyp = 0.27. Eye-group
(Fig. 43) on and around sloping, not steep, ocular process, rectangular, AR/PR = 0.99, twice
as wide as long, AR/El = 2.00, POP faint and broken-up, ALE largest, ALE/PLE = 1.41.
Fovea recurved at base of caput, curving out laterally. Chelicerae strong, bicolored brown
(dorsally lighter than laterally), fine black setae along dorsal crest, merging distally with
rastellum, fine whitish pubescence in longitudinal zones on lateral faces. Cheliceral furrow
promargin with five teeth, distal tooth strongest. Rastellum consists of five strong spines
apical to fang base. Fangs very strong with smooth prolateral keel (Fig. 6). Legs yellowish
brown, lighter yellow laterally and ventrally on femur IV and on proximal parts of coxae II-
IV; scopulae over full length of tarsi and metatarsi, and on tibiae I & II extending as thin
pseudoscopula on patellae I & II; spines in rows on lateral or ventral sides of all metatarsi,
tibiae and patellae, spiny setae dorsally on femora; prolateral spines on patella and tibia III
(Fig. 44). Palp: color as legs, scopula on tarsus and ventrally and laterally on distal half of
tibia, three prolateral spines (2 in line and 1 more ventrally) on patella. Ventral aspect (Fig.
41). Maxillae almost twice as long as wide, l/w = 1.96, with small anterior-apical process,
cuspules in irregular rows along

Figs. 40-46: Nemesia santeugenia sp. n., female. 40 habitus dorsal; 41 habitus ventral; 42 carapace lateral;
43 eye-formation, dorsal; 44 patella and tibia III, prolateral; 45 spinnerets, ventral; 46 spermathecae, dorsal.
Scale line = 2mm (40-42), 1mm (43-46).
prolateral proximal edge. Sternum centrally lighter brown than along edges, similar shading
as ventrally on coxae, evenly covered with setae of varying size, anterior and median sigilla
round and marginal, posterior sigilla oval and sub-marginal (about their largest diameter from
sternum edge). Labium more greyish than sternum, dome-shaped, wider than long, even1y
covered with fine setae proxirmally and stronger setae distally. Labial furrow wide, glabrous,
co1our as sternum edge. Abdomen brownish grey with complex dorsal-latera1 pattern of
brownish patches and chevrons. Spinnerets (Fig. 45) same color as ventral abdomen, PMS
reduced, spiky, with few apica1 spigots, PLS basa1 segment with few fine spigots near
ventral-dista1 edge, apica1 spigots evenly distributed around four macro-spigots.
Spermathecae (Fig. 46) unipartite, compact, a1most as broad as long, "potato-shaped", with
dense cover of glandular tissue.
Variation (n=2): BL = 15-23.5, CL = 5.9-8.2. Carapace, CL/CW = 1.3-1.5, CL/Ca = 1.6.
Caput Ch/Th = 1.9-2.2. Clyp = 0.19-0.27. Eye-formation: AR/PR = 0.99- 1.02, AR/El = 2.00-
2.06, ALE/PLE = 1.21-1.41. Male: Unknown.
Burrow (Fig. 71): Nemesia santeugenia inhabits a typica1 "branched wafer-door burrow".
The burrows were found in relatively soft sandy soils on gently sloping surfaces covered with
debris of conifer needles, fallen leaves, bits of grasses and moss. The thin, flexible trapdoor
(wafer door; Moggridge, 1873, 1874) is camouflaged by spun-in debris from the surrounding
surface and 1oosely closes the burrow entrance by falling over the entrance opening. The

urrow runs almost vertically into the ground and may reach a depth of 30 cm. Four to five
centimeters behind the trapdoor a side tube branches off from the main burrow. This side tube
reaches the soil surface a few centimeters from the trapdoor, where its surface opening is
covered with a thin sheet of silk that is well camouflaged among the plant debris. On the
lower side of the burrow entrance some "linear litter", as described by Main 1957, is spun into
the burrow rim.
Behavior: The only spider of this species kept in captivity never settled down properly. After
days of hiding inactively in a shallow depression made in the clay bottom of its container, it
started digging down to produce a burrow. The burrow was never completed however, and no
trapdoor was built. Nevertheless, in the six months that the spider was kept in this way it
captured four isopods that wandered near the open burrow. It ate these preys and removed the
compacted chitin remains by working them out of the burrow entrance.
Distribution (Fig. 86): Nemesia santeugenia seems to be remarkably rare. Among 120
Nemesia spiders collected on Majorca, only 2 specimens appeared to be N. santeugenia.
These two specimens came from 1ocations about 25 km apart in the central regions of the
island.
Nemesia seldeni sp. n. (Figs. 47-53, 72, 77, 81, 83)
Types: Holotype ♀ (ARI4202), 4 Apri11996, Majorca, Randa, 39.503°N, 2.972°E, burrow in
sloping clay-bank along forested roadside. Paratypes: Majorca: 2 ♀♀ (AR14203;
9972.4015), 8 Apri11995, Randa, 39.503°N, 2.972°E, same habitat; 1♀ (AR14204), 9 April
1995, Randa, 39.503°N, 2.972°E, same habitat; 1♀ (9972.4017), 4 April 1996, Llucmajor,
39.500°N, 2.937°E, same habitat: 1 ♀ (9972.4016).13 October 2000, Inca-Puebla, 39.767°N,
2.986°E, horizontal ground, edge of cultivated field. All leg. A. E. Decae.
Etymology: The species is named after Dr P. A. Selden who rediscovered Bristowe's N.
bristowei in 1989 on Majorca near Estellencs (39.655°N, 2.4800E), and provided the author
with the first specimens for study.
Diagnosis: Females of N. seldeni are morphologically very similar to those of N. maculatipes
Doleschall, in Ausserer 1871 and N. sanzoi Fage 1917. They differ from both those species by
the relatively elongated carapace, CL/CW ≥ 1.4 (n=6) in N. seldeni, CL/CW = 1.3 (n = 5) in
the other two species, the lower caput (Ch/Th = 1.2 vs. 1.5-1.7), the prolateral scopula on
tibiae I and II restricted to the distal half of the segment (vs. extending over the full length of
the tibiae), and the prolateral spine pattern on patella III (usually 2 spines, rarely 3, in N.
seldeni, vs. usually 3 spines, rarely 2, in N. maculatipes and N. sanzoi). The male is unknown.
Description: Female (holotype): BL = 15.4, CL = 5.5, CW = 3.7. Leg IV: T4>F4>M4. PSP: p
= 0-0; I = l-l; II = 2-2; III = 2-2; IV = 0-0. Dorsal aspect (Fig. 47). Carapace distinctly longer
than wide, CL/CW = 1.5, with distinct narrow dark purplish lateral edges, and clear pattern of
darker brown zones radiating from fovea over lighter, yellowish background. Brown zones
cover lateral slopes of caput and follow cervical grooves and all shallow radial furrows.
Clypeus narrow, Clyp = 0.17, light yellowish brown, as is cuticle at sides of eye-group and on
central caput. Caput (Fig. 49) slightly elevated, Ch/Th = 1.2. Fine setae in one longitudinal
row on central caput and in irregular group on clypeus and ocular tubercle. Fine hairs in
irregular patches predominantly on darker colored zones and along carapace edge. Eyes (Fig.
50) compactly grouped on and around steep ocular tubercle, eye-formation less than twice as
wide as long, AR/El = 1.83, AR slightly shorter than PR, AR/PR = 0.98, ALE larger than
PLE, ALE/PLE = 1.21, PLE pearly, other eyes greyish. POP unbroken. Fovea narrow, deep,
recurved, with smal1 central longitudinal groove. Chelicerae dorsally appearing somewhat
darker than carapace, distinctly bicoloured with lighter and darker brown zones, lighter zones

Figs. 47-53: Nemesia seldeni sp. n., female. 47 habitus dorsal; 48 habitus ventral; 49 carapace lateral; 50 eyeformation
dorsal; 51 patella and tibia III, prolateral; 52 spinnerets, ventral; 53 spermathecae, dorsal (m =
macula). Scale lines =2mm (47-49), 1mm (51), 0.5mm (50, 52-53).
covered with fine hairs and stronger setae that merge distally with rastellum. Cheliceral
furrow with six promarginal teeth, increasing in size from proximal to distal. Rastellum of
five strong teeth. Fangs with smooth promarginal keel (Fig. 6). Legs yellow with darker
longitudinal zones dorsally on femora and inconspicuous maculae on outer surfaces; dense
scopulae on tarsi and metatarsi I and II extending onto prolateral distal half of tibiae I and II.
Fine pro- and retrolateral spines in ventral rows on metatarsi and tibiae I and II and more
laterally and dorsally on III and IV, lateral spines on patellae I-III (see PSP), prolateral spines
on tibia and patella III (Fig. 51), dorsal spines on all femora. Palp: color as legs, scopula on
tarsus and tibia, one pro- and one retrolateral proximal spine on tarsus; double row of short
ventral spines dividing tarsal scopula, no patellar spines, group of long dorsal spiny setae
distally on femur. Ventral aspect (Fig. 48). Maxillae almost twice as long as wide, extending
into small anterior distal process, yellow, except for creamy white anterior zone of maxillary
scopula, evenly covered with setae, only one cuspule in proximal prolateral corner. Sternum
longer than wide (l/w=l.44), yellow, evenly covered with fine setae, three pairs of oval sigilla.
Labium twice as wide as long, darker than sternum, evenly covered with setae, labial furrow
wide and glabrous. Abdomen ovoid, densely covered with fine greyish-white hairs, pale grey
cuticle with yellowish sheen ventrally and irregular dorsal pattern of dark greyish brown
blotches around similarly colored central longitudinal zone, fine setae dorsally. Spinnerets
similar in color to ventral abdomen, PMS well developed with fine spigots ventrally and
distally, PLS short, lateral macula (Fig. 52), basal segment longer than median and distal
segments together, spigots evenly spread over ventral surface of basal and median segments
and apically grouped around one macro-spigot. Spermathecae (Fig. 53) tripartite, proximal
part short with dense glandular tissue, middle part twisted with dense glandular tissue
proximally and thin glandular tissue distally, distal part digitiform with thin glandular tissue.

Variation (n= 6): Adult females small Nemesia species, BL = 13.7-16.3, CL = 4.2-5.8.
Carapace, CL/CW = 1.4-1.5, CL/Ca = 1.5-1.7. Clyp = 0.14-0.19. Eye-formation: AR/PR =
0.96-1.02, AR/El = 1.86-2.04, ALE/PLE = 1.07-1.25. Chelicerae in some specimens not
contrasting with color of carapace. Maxillary anterior-apical process relatively well developed
in some specimens, less so in others, one or more cuspules (in irregular rows) may be present.
PSPvar: p = 0(2); I = 1(2); II = 2; III = 2(1); IV = 0. Maculae on leg segments and basal
segment of PLS more or less distinct. Male: unknown.
Burrow (Fig. 72): The structure of the burrow of N. seldeni is roughly similar to that of N.
santeugenia. Both have a thin flexible wafer-door (Moggridge, 1873, 1874), which closes off
the main burrow tube at the soil surface, both have a side tube that branches off from the main
burrow a few centimeters behind the trapdoor and that opens onto the soil surface, both have
only the upper half of the burrow (including the inside of the side tube) lined with silk, and
both have linear plant material spun into the burrow rim (Fig. 81). Two differences between
the burrows of N. seldeni and N. santeugenia have been noticed: first, the diameter and depth
are smaller in N. seldeni (probably because N. seldeni is a smaller spider), and secondly, N.
seldeni constructs a second, smaller trapdoor that closes off the side tube (Fig. 72, d). A
second trapdoor has not been observed in N. santeugenia.
Behavior: Nemesia seldeni spiders, captured in the field, readily constructed new burrows in
the laboratory. Usually the spiders could be induced to dig their burrow in a desired position,
by providing them with an artificial impression about 1 cm deep in the clay soil in the centre
of the container. The main burrow tube with trapdoor was built first, and the side tube,
including a smaller trapdoor, appeared later. In the field the larger, first trapdoor, usually
faces down the slope, while the second smaller trapdoor usually opens uphill (Fig. 72).
Observations on captive spiders showed that the spiders invariably take up their ambush
positions at dusk under the larger, first trapdoor. The second trapdoor was found to serve
probably several functions. It may be used for escape when the spider is threatened inside its
burrow, as has been seen in several other Nemesia species that build similar burrows,
although this escape function was not definitely established in the course of this study on N.
seldeni. When capturing N. seldeni in their natural burrows these spiders were inclined to
retreat to the bottom of their burrow and, when the top half of the burrow was removed, the
spiders would dash out from the remnants of the main burrow in an effort to escape collection.
The smaller, second trapdoor definitely serves purposes other than solely as a possible escape
route. One function is an enlargement of the hunting area by serving as an "early-warning"
system. In captivity N. seldeni, while waiting in ambush at the larger trapdoor, was seen to
react to small isopods crawling over the smaller trapdoor by swiftly retreating from its
ambush position and launching an immediate attack from the second trapdoor. Generally,
however, prey was captured from the larger trapdoor within a range of up to several
centimeters away from the burrow entrance. Like N. brauni, N. seldeni was found to leave the
burrow completely in pursuit of prey that may be captured at distances of several centimeters
away from the burrow. The second trapdoor also functions in waste disposal. Indigestible
chitin remains of prey and exuviae are pushed out of this smaller door. Nemesia seldeni seems
less strictly nocturnal in its hunting activity than most other Nemesia species and was
sometimes found, both in the field and in captivity, to lie in ambush during daylight hours.
Distribution (Fig. 83): Nemesia seldeni has a wide distribution in the western and central
parts of Majorca, where it prefers moist and shady positions. It is particularly common in
humus-rich soils under vegetation and in creek banks. In preferring these moist and shady
habitats it is often separated from other Nemesia species that seem to prefer more exposed and
often less stable soils, although N. seldeni has also been found in close proximity ( < 1m) to
all other Nemesia species found on Majorca.

Nemesia santeulalia sp. n. (Figs. 54-60, 73, 86)
Types: Holotype ♀ (AR14194), 8 May 1986, Ibiza, Santa Eulalia del Rio, 38.987°N, 1.530°E,
burrow in clay-bank along road between pine forest and village edge. Paratypes: Ibiza: 4 ♀♀
(AR14195; AR14196; 9972.4022; 9972.4023), 9-10 May 1986, Figueritas, 38.911°N,
1.416°E, horizontal and slightly sloping ground in wasteland on building site; 1 ♀
(9972.4021), 4 May 1986, San Antonio, 38.977°N, 1.316°E, in sandy slope on village edge.
All specimen leg. A. E. Decae.
Etymology: The specific name is a noun in apposition taken from the village Santa Eulalia
del Rio, the type locality, on Ibiza.
Diagnosis: Differs from all other known Nemesia species, except N. santeugenia sp. n., in the
morphology of the spermathecae (Fig. 60). For observed differences in the morphology,
burrow structure and behavior between N. santeulalia and N. santeugenia, see diagnosis of
the latter species. The male is unknown.
Description: Female (holotype): BL = 22.7, CL = 7.7, CW = 5.7. Leg IV: T4>F4>M4. PSP: p
= 3-3; I = 2-2; II = 2-1; III = 2-; IV = 0-0. Dorsal aspect (Fig. 55). Carapace longer than
wide, CL/CW = 1.4, central dark brown zones with irregular fine black line patterns radiating
from fovea over yellowish brown background, posterior and lateral edges marked with narrow
purplish black line, darker zones covered with black and silvery white pubescence. Caput
strongly elevated, Ch/Th = 1.9, fine setae on caput crest, ocular tubercle and posterior end of
thorax. Clypeus wide, Clyp = 0.27. Eye-group (Fig. 57) rectangular, AR/PR=0.99, slightly
more than twice as wide as long, AR/El = 2.03, ALE as large as PLE, ALE/PLE = 1.00,
posterior eyes pearly, anterior eyes greyish. POP dense and unbroken. Fovea smoothly
recurved, centrally narrow; laterally distinctly widened. Chelicerae dark brown, contrasting
with carapace, darker brown in areas with setae or hair cover, lighter in glabrous areas,
narrow band of silvery white pubescence laterally, cheliceral furrow with six promarginal
teeth, distal five equally strong, most proximal tooth smaller and placed somewhat apart.
Fangs strong, blunt with smooth keel (Fig. 6). Legs yellowish brown, similar color to
sternum, with darker longitudinal zone dorsally on femora and dark pigmented maculae
distally on all outer surfaces of femora. Spines on metatarsi, tibiae and femora thin. Scopulae
on tarsi and metatarsi I and II extending over full length of tibiae onto patellae. Prolateral
spines on tibia and patella III (Fig. 58). Palp: color as legs, but without lateral maculae on
femora, tarsal scopula extending halfway along tibia, three prolateral spines on patella (2 in
line and 1 more ventrally). Ventral aspect (Fig. 54). Maxillae dark brown, with creamy white
anterior scopula zone, longer than wide (l/w = 1.7), well developed row of cuspules along
proximal margin. Sternum warm yellowish brown, darker around edges, evenly covered with
setae, three pairs of well-developed sigilla, posteriors oval and placed slightly away from
margin. Labium almost as long as wide, dome-shaped, strongly elevated, separated from
sternum by narrow dark brown furrow. Abdomen evenly covered with fine hairs, dorsally
with dense irregular pattern of dark lines and patches on light background, ventrally light grey
with yellow book lung covers. Spinnerets (Fig. 59): PMS lighter than PLS, PMS small with
several apical spigots, PLS proximal segment longer than median and distal segments
together, spigots on apical spigot field in concentric circles around four macro-spigots.
Spermathecae (Fig. 60) unipartite, compact, almost as broad as long, "potato-shape", with
dense cover of glandular tissue.
Variation (n=6): Adult females, medium to large spiders, BL = 18-23, CL= 6.7-7.7.
Carapace, CL/ CW =1.3-1.4, CL/Ca = 1.5-1.6, Ch/Th= 1.9-2.0. Eye- formation: AR/PR =
0.95-0.99, AR/El = 1.97-2.06, ALE/PLE = 0.9-1.2. PSPvar: p = 2(3); I = 2(1); II = 2(0-1);
III = 2(1); IV = 0. Male: unknown.

Figs. 54-60 Nemesia santeulalia sp. n. female. 54 habitus, ventral; 55 habitus dorsal; 56 carapace lateral; 57 eyeformation,
dorsal; 58 patella and tibia III, prolateral; 59 spinnerets, ventral; 60 spermathecae, dorsal. Scale lines
= 2mm (54-56), 1mm (57-60).
Burrow (Fig. 73): Adult N. santeulalia females dig a 15-20 cm deep burrow, closed at the
entrance by a thin flexible trapdoor (wafer-door). The maximum diameter of the trapdoor
measured in the field was 1.5 cm, and the largest entrance diameter of the burrow measured
was 1.4 cm. Two to three centimeters behind the trapdoor a side tube branches off from the
main burrow. The side tube slopes upwards to end in a cul-de-sac (ten out of ten
observations) just below the soil surface. The whole inside of the burrow, including the side
tube and the underside of the trapdoor, is lined with one continuous, densely woven sheet of
silk. Near the bottom of the main tube the silk is thinner than in the higher parts of the
burrow. Indigestible remains of prey are packed into the burrow bottom or into the lower
wall of the main tube. In building a dead-ended side tube, the full lining of the burrow with
silk and the storage of chitinous remains in the walls and bottom of the burrow, N.
santeulalia of Ibiza differs markedly from N. santeugenia from Majorca, from which species
it is morphologically difficult to distinguish.
Behavior: Females of N. santeulalia seem to spend the day rather passively. Insects placed on
the trapdoor in daylight hours, or lightly teasing the burrow entrance with a blade of grass,
failed to trigger any observable reaction from the spider. Stronger teasing, however, resulted
in the spider coming to the burrow entrance to pull the door firmly closed. After dark the
spider reacts differently to both stimuli. An insect placed on or near the trapdoor, or teasing of
the burrow rim with a grass blade, elicit an instant aggressive response as the spider darts
forwards with great accuracy in the direction of the stimulus. Nemesia santeulalia hunts from
behind an almost closed trapdoor. Most wafer-door building species extend the distal parts of
the first, second and third pairs of legs from under the door when lying in wait for prey (see N.
seldeni, Fig. 77), but no parts were observed to be extended in N. santeulalia. On disturbance

of the burrow (by the spider collector), N. santeulalia will initially hold on to the trapdoor to
keep it closed. When the trapdoor is forced open the spider will retreat to the bottom of the
burrow and wait there head-up. When further disturbed, it will finally run up the burrow again
to hide in the side tube. The spiders were not aggressive and were hardly ever observed to
bite. Analysis of the indigestible remains of prey found in the burrow bottoms and walls
revealed head -capsules, legs and mandibles of ants, legs and elytra of beetles and possibly
remains of woodlice. Probably such ground-dwelling arthropods constitute the main prey of
N. santeulalia, although other types of prey cannot be ruled out on the grounds of this
observation alone 14 .
Distribution (Fig. 86): Nemesia santeulalia was found to be common in quite widely
separated places on Ibiza. It has not so far been found on the nearby island of Formentera.
Nemesia ibiza sp. n. (Figs. 61-67, 74, 85)
Types: Holotype ♀(AR 14067), 3 May 1986, Ibiza, San Juan, 39.081°N, 1.510°E, burrow in
clay-fillings between limestone rocks. Paratypes: IBIZA: 1 ♀ (ARI4065), 3 May 1986, same
data as holotype; 2 ♀♀ (AR 14066; 9972.4018), 10 May 1986, Figueritas, 38.911°N,
1.416°E, horizontal and slightly sloping ground in wasteland on building site; 2♀♀
(9972.4019, 9972.4020), 8 May 1986, Santa Eulalia del Rio, 38.987°N, 1.530°E, in clay-bank
along road between pine forest and village edge. All specimen leg. A. E. Decae. All locations
mentioned here are on Ibiza, but the species was also found near Punta de la Anguila on the
nearby island of Formentera.
Etymology: The specific name is a noun in apposition taken from the type locality, the island
of Ibiza, where it was found to be common and widely distributed.
Diagnosis: Females of N. ibiza can be distinguished from an other known Nemesia species by
the "hour- glass" shape of the spermathecae (Fig. 67), particularly the proximal narrowing of
the receptacula seems to be characteristic. The male is unknown.
Note: Nemesia ibiza seems to be closely related to N. hispanica, because some differences
observed in the morphology of the two species (eye-formation, spine pattern, morphology of
legs III and IV) may fall within the range of geographical variation of N. hispanica. However,
on the grounds of the observed differences, the limited material of N. hispanica, and the
absence of males of both species, N. ibiza is here regarded as a distinct species.
Description: Female (holotype): BL = 20.4, CL = 7.4, CW = 6.7. Leg IV: F4>T4>M4. PSP: p
= 2-2; I = 1-1; II = 1-1; III = 2-2; IV = 0-0. Dorsal aspect (Fig. 62). Carapace slightly longer
than wide, CL/CW = 1.1, yellowish brown with "leaf pattern" of darker zones radiating from
fovea and along cervical grooves and radial furrows, setae along anterior edge of clypeus and
on ocular tubercle. Caput elevated, Ch/Th = 1.9, groups of fine setae on each side of central
longitudinal row of stronger bristles on crest of caput, pubescence absent, but widely
dispersed fine hairs present. Clypeus yellowish, relatively wide, Clyp = 0.36. Eye-group (Fig.
64) almost rectangular, AR/PR = 0.96, less than twice as wide as long, AR/El = 1.87, ALE
largest, ALE/PLE = 1.17, anterior eyes greyish, posterior eyes pearly. POP broken between
AME and lateral eyes. Fovea not smoothly recurved, but somewhat angular. Chelicerae warm
reddish brown, darker than and contrasting with overall color of carapace, with narrow dark
brown glabrous zone dorso-laterally, cheliceral furrow with seven promarginal teeth, distal
14 Ants, beetles and woodlice were observed to be captured and eaten by other Nemesia species in captivity. Other types of
prey such as earthworms, maggots, flies and earwigs, were captured and eaten and even food items that cannot possibly be
part of their natural diet, such as pieces of prawn and tiny meatballs placed on the burrow rim were accepted as food. These
observations indicate that Nemesia trapdoor spiders are food generalists.

Figs. 61-67: Nemesia ibiza sp. n., female. 61 habitus, ventral; 62 habitus, dorsal; 63 carapace, lateral; 64 eyeformation,
dorsal; 65 patella and tibia III, prolateral; 66 spinnerets, ventral; 67 spermathecae, dorsal. Scale lines
= 2mm (61-63), 1mm (64-67).
tooth largest. Rastellum on small process. Fangs short, blunt, with irregularly serrated keel
(Fig. 8). Legs mostly uniform yellowish brown, femora III and IV laterally and ventrally
lighter whitish yellow, scopulae on tarsi and metatarsi I and II not extending onto tibia I,
numerous spines dorsally and dorso-laterally on femora, ventrally and ventro-laterally on
tibiae and metatarsi I-III, two dorsal spines on metatarsus III, prolateral spines on tibia and
patella III (Fig. 65), tibia IV with three long slender prolateral spines along ventro-lateral
edge, metatarsus IV with retrolateral apical comb. Palp: colored as anterior legs, scopula
extending onto tibia, two parallel rows of strong pro-lateral spines and three parallel rows of
strong retrolateral spines on tibia. Ventral aspect (Fig. 61). Maxillae yellowish brown, almost
twice as long as wide (1/w= 1.9), evenly covered with curved setae, well defined double rows
of cuspules along proximal edge, anterior scopula zone white. Sternum yellow, slightly longer
than wide (1/w = 1.1 ), evenly covered with setae, three pairs of oval sigilla, anterior and
median pairs touching edge, posterior pair their longest diameter from sternum edge. Labium
wider than long, almost rectangular, dome-shaped, strongly elevated, separated from sternum
by wide glabrous furrow. Abdomen grey dorsally with dark lateral chevrons, ventrally lighter
and more uniformly grey, whole abdominal surface evenly covered with fine hairs. Spinnerets
(Fig. 66) yellow, contrasting with grey of ventral abdomen, PMS reduced with few apical
spigots, basal segment of PLS as long as median and distal segments together, spigots in
apical field tightly grouped, with smaller spigots in circles around three central macro-spigots.

Spermathecae (Fig. 67) tripartite, "hourglass-shape", evenly covered with very thin glandular
tissue.
Variation (n=6): Females medium sized spiders, B L= 19-21, CL= 6.5-8.0. Carapace,
CL/CW = 1.1-1.2, CL/Ca = 1.6, Ch/Th = 1.9-2.5. Clyp = 0.3l-0.36. Eye- formation: AR/PR =
0.96-l.00, AR/El = 1.87-2.14, ALE/ PLE = 1.18-1.38. POP entirely or indistinctly broken
between AME. PSPvar: p = 1(2-3); I = 1(2); II = 1(2); III = 2(0-1); IV = 0. Leg IV:
F4>T4≥M4. Spinnerets: PMS with two or three fine spigots on apex. Fang-keel: serrated
(rarely smooth). Abdominal pattern more or less distinct. Male: unknown.
Burrow (Fig. 74): Nemesia ibiza builds a typica1 cork- burrow, with a thick trapdoor that fits
within the entrance of the burrow-tube. The depth of the burrow is usually around 12 cm, but
may vary between 7-16 cm. The maximum diameter of the trapdoor measured in the field was
1.8 cm, and the diameter of the burrow entrance was 1.75 cm. There are no internal sidediggings,
but the burrow may bend in various ways, probably around underground obstacles.
Most of the burrows are dug in steep or vertical clay-fillings between natural rock-layers or in
constructed stone walls. The silk lining of the burrow is thickest near the burrow-entrance and
vanishes completely a few centimeters down the shaft. Indigestible chitin-chips and remains
of prey are stored in a compacted package pushed into the lower burrow-wall, or in the
burrow-bottom. Identified prey- remains were similar to those found for N. santeulalia.
Figs. 68-74: Longitudinal sections of Nemesia burrows showing different tube shapes and types of trapdoors,
extent of silk burrow lining (dashed lines), presence or absence of prey remains in pockets associated with the
burrows (pr), presence of a second trapdoor (d), and positions in which the spiders were usually encountered
during collection. 68 N. brauni (aggressive, ready to attack at burrow bottom); 69 N. bristowei (defensive
posture, inactive at burrow bottom); 70 N. randa (defensive posture, inactive at burrow bottom); 71 N.
santeugenia (halfway up burrow ready to escape); 72 N. seldeni (halfway up burrow ready to escape); 73 N.
santeulalia (hiding in side branch); 74 N. ibiza (aggressive, ready to attack at burrow bottom). Arrows indicate
opening direction of trapdoors. Scale line = 1cm.

Behavior: Some spiders were found to be active during daylight hours, lying in wait for prey
behind a cracked-open trapdoor. Generally however, N. ibiza only becomes active at dusk,
when the spiders come to the entrance of the burrow to lie in wait for prey passing near to the
burrow, as do virtually all other trapdoor spiders. Nemesia ibiza seems to defend its burrow
somewhat more actively, by pulling the door closed when it is disturbed, than is the case in N.
santeulalia. Although both species show this behavior as a reaction to attempts to open the
trapdoor from the outside, this "door-holding" was rarely observed in the latter species and
virtually always in N. ibiza.
Distribution (Fig. 85): Nemesia ibiza was widely and commonly found on Ibiza. It appeared
to be rarer on Formentera where it was found only in association with clay-fillings in stone
walls.
Figs 75-78 Different ambush positions of Majorcan Nemesia species. 75 N. brauni; 76 N. bristowei,
77 N. seldeni; 78 N. randa.
Discussion
Excluding the four species (see introduction) from China, Mozambique, Afghanistan and
Cuba listed in Platnick 2003, which on biogeographical grounds are unlikely to be members
of the genus, Nemesia is distributed in a rather restricted geographical area in southern Europe
and in Africa north of the Sahara (Chapter 2 Fig. 5). In this relatively small distribution zone,
about fifty different species are currently recognized. Many, if not all, of these species appear
to be local endemics. Several observations may support this view of Nemesia being a
regionally distributed genus composed of numerous locally endemic species both on islands
and in continental regions.

Island endemics: Of the 49 "Mediterranean" species and subspecies listed in Platnick 2003
twelve species (24%) are known exclusively from islands, and only two (N. cellicola and N.
maculatipes) species (4%) are recorded from both mainland and island locations 15 .
Widespread island endemism is also supported by records, partly as yet unpublished, of
endemic Nemesia species from Crete, Corfu, Malta, Sicily, Sardinia, Corsica, Elba,
Montecristo, Ile d' Alboran, Majorca and Ibiza.
Mainland endemics: Local endemism is also found in continental areas. In southern France,
where Nemesia was first studied in some detail, Moggridge 1873, 1874 discovered that the
Nemesia faunas east and west of the river Rhone differed remarkably in their species
composition. East of the Rhone he found N. carminans, N. congener O.P-Cambridge 1874, N.
e/eanora O.P-Cambridge 1873 and N. manderstjernae 16 , while west of the Rhone he found N.
caementaria (Latreille 1799), N. dubia O.P-Cambridge 1874 and N. simoni. Simon 1914
added N. raripila from the Pyrenees to the Nemesia fauna of southwest France. In a current
study of the Nemesia fauna of Portugal (Decae & Cardoso in prep.) endemism again seems to
be a remarkable feature, because none of the six species collected from various sites
throughout Portugal has so far been found in Spain. Since some of the Portuguese collections
were made close to the Spanish border, it is unlikely that all these species are absent from
Spain, but their presence there still has to be established. This study of the Portuguese
Nemesia fauna also shows that the presence of species that had previously been reported from
Portugal, but that have their type localities in distant lands (N. dubia from Montpellier, N.
meridionalis Costa, 1835 from Naples), could not be reconfirmed, which suggests that the
distribution of these species is more restricted than previously supposed. The information
presented here on the species of Majorca and Ibiza also underlines the view that local
endemism is strong in Nemesia, because the Nemesia fauna of Majorca appears to differ from
that of nearby Ibiza, and because it shows that even on an island the size of Majorca, species
distributions can be geographically separate (see Fig. 84 for the distributions of N. bristowei
and N. randa).
Whether Nemesia really is a regional genus composed of numerous locally endemic species,
however, requires further investigation. Too little is currently known about the identity of
individual species in large parts of the distribution range (Spain, Italy, Balkans, Greece, North
Africa) and virtually nothing is known about possible interspecific, let alone phylogenetic,
relationships between species. Early suggestions of such relationships (Simon 1914; Frade &
Bacelar 1931) were unavoidably speculative owing to the very limited amount of information
then available.
New collections and the discovery of better diagnostic characters (particularly spermathecae
and spinneret morphology) have provided the opportunity for the following fresh speculations
on possible interspecific relationships within Nemesia.
Based on the morphology of the spermathecae, a broad division between eastern and western
Nemesia species may exist. The eastern species characteristically have narrow, elongated
spermathecae that are medially "twisted", as illustrated here for N. seldeni (Fig. 53). Western
species have simpler built and much broader spermathecae as shown here in Figs. 19, 32, 39,
15 There is some doubt about the correctness of distribution data for the two species reported from both mainland and island
locations, first, because the reported distributions are not easily explained and show unlikely disjunctions, and secondly,
because these two species, N cellicola (Audouin, 1826) and N. maculatipes Ausserer, 1871, are species that (for different
reasons) are easily misidentified. N. cellicola is the type species of the genus, and in early collections, before information on
the diversity of Nemesia was available, was reported mistakenly from various wide1y separated locations. Nemesia
maculatipes seems to carry its diagnostic character in its name (the maculae on legs and PLS), but it is now clear that the
maculate pattern is present in different, only distantly related, Nemesia species and that such species are easily confused.
16 The distinction between the last two species is currently unclear.

Figs 79-82 Trapdoors of Majorcan Nemesia species. 79 N. brauni; 80 N. bristowei, 81 N. seldeni;
82 N. randa.
46, 60 and 67. In the central areas of the distribution range, species with these two different
types of spermathecae overlap. Nemesia carminans, which occurs in an area between the river
Rhone in south-central France and the city of Genoa in northern Italy, is currently the species
with "western-type" spermathecae that extend its distribution range furthest to the east.
Nemesia dubia, with a distribution range between Montpellier (southern France) and eastern
Spain (Blasco 1986b), is currently the most westward extending species with "eastern- type"
spermathecae 17 . It is currently unknown if a similar phenomenon occurs on the north African
side of the Mediterranean, but on Mediterranean islands it is evident from what is reported
here, that of the seven Balearic species recognized six have "western-type" spermathecae and
only N. seldeni has "eastern-type" spermathecae.
Interspecific relationships of N. seldeni: Nemesia seldeni appears to belong to a tribe of small
(♀ BL = 13-17 mm), wafer-door building Nemesia species that is here designated the
maculatipes-group. Typical, but not unique for the species in this group, is the presence of
dark pigmented blotches (maculae) on the outer faces of leg-segments (mainly on the femora,
patellae and tibiae) and on the outer face of the basal segment of the PLS. Females of the
maculatipes-group differ from those of other Nemesia species by a combination of characters
that include "eastern-type", twisted, receptacles (not yet established in all species), a relatively
elongated carapace, dense pubescence on carapace and legs, distinct color pattern with a
17 Although Blasco 1985 shows more ‘western-type’ spermathecae for N. dubia from Catalonia.

wedge-shaped yellow zone on the crest of the caput, smooth fang, a retrolateral spine on
patella III, and well developed PMS with spigots on the ventral surface. The maculatipesgroup
is distributed on islands in the western basin of the Mediterranean (Majorca, Corsica,
Sardinia, Sicily and smaller islands). The group contains several species, some of which may
be synonymous. Further study is necessary to confirm the taxonomy of this group. Species
supposedly included in the maculatipes-group are: N. maculatipes, N. sanzoi, N. pavani
Dresco 1978, N. fertoni Simon 1914, N. arenicola Simon 1892, N. albicomis Simon 1914, N.
kahmanni Kraus 1955 and maybe other species.
Interspecific relationships of N. santeugenia and N. santeulalia: Nemesia santeugenia from
Majorca and N. santeulalia from Ibiza seem to be close relatives and both endemic to the
Balearics. Morphologically the two species are hard to distinguish. The most prominent
difference is found in the PMS, which are more strongly developed and carry more spigots in
N. santeulalia than in N. santeugenia. This feature may, or may not, be related to a behavioral
difference shown in the structure of the burrow. The burrow of N. santeulalia is fully lined
with silk while that of N santeugenia is only partly lined. Other behavioral differences
reflected in the structure of the burrow are the storage of indigestible prey-remains in the
burrow walls and floor by N. santeulalia, and the dead-ended side tube that this species
builds, in contrast to the open-ended side tube and the removal of prey-remains from the

urrow by N. santeugenia. Based on their spermathecae structure, both species probably have
their closest relatives to the west, possibly in Morocco (N. ariasi figured in Blasco 1986a) or
on the Iberian Peninsula (N. hispanica figured in Buchli 1968). More probably, however, the
closest relative is an as yet unidentified species collected by Buchli in Spain.
Interspecific relationships of N. brauni: Nemesia brauni differs strongly in behavior and
morphology from all other Majorcan Nemesia species. Particularly the absence of PMS places
this species apart, as does its aggressive behavior and its extrovert way of hunting. The closest
relatives of N. brauni are probably several species found on the Iberian peninsula (both in
Portugal and Spain) with which it shares its large size, aggressive attitude, thick cork-door,
burrow type and absence of PMS. A Portuguese species that has long been regarded as N.
hispanica (Frade & Bacelar 1931; Machado 1944; Cardoso 2000) was recently compared with
the type of N. hispanica and found to be a different, as yet undescribed, species.
Interspecific relationships of N. bristowei and N. randa: Both species appear to be endemic to
Majorca, where they occur in separate geographical areas (Fig. 84). Nemesia bristowei and N.
randa show a number of interesting similarities in morphology (carapace shape, eyeformation,
"western-type" spermathecae), behavior (reclusive lifestyle, simple fully silk-lined
burrows, modified trapdoor) and ecology (predominantly found on steep surfaces) that might
indicate a mutual relationship. According to their type of spermathecae N. bristowei and N.
randa seem to be "western" species, of which the closest relatives should be expected on
nearby islands or on the Iberian Peninsula. However, N. bristowei, with its remarkable cogwheel
shaped trapdoor, might have a relative towards the east, on Sardinia, where an as yet
undescribed species seems to build a somewhat similar trapdoor (though more "star-shaped")
that has been photographed by and published in Kullmann & Stern 1975 (see also Bellmann
1997 p. 37).
Interspecific relationships of N. ibiza: This species seems to be a member of a larger tribe of
cork-door building Nemesia species that has a wide distribution on the Iberian mainland and
in southern France. The group probably contains N. hispanica, N caementaria and N.
carminans. In particular N. ibiza and N. hispanica seem to be closely related. If these two
species could be proved to be synonymous this would provide the first definite record of a
species with a distribution that includes both mainland and island locations.
Finally, it is unfortunate that the present study is rather incomplete. Particularly the absence of
males of most species described here is an important omission. To complete the picture of the
Balearic Nemesia fauna it will be necessary to obtain the missing males and also to study the
Nemesia species that occur on Minorca and the smaller islands in the Balearic archipelago. It
is hoped that samples from these locations and males of all species will become available for
study shortly in order to complete this study and to improve our understanding of Nemesia
diversity and distribution in the region.
Acknowledgements
I wish to thank Peter Merrett for his critical comments and thorough editorial work on the
manuscript, Paul Selden for sharing his discovery of N. bristowei with me and for providing
me with the first specimens of this species for study, Paul Hillyard for his study of the
spinneret morphology of the type specimens of N. brauni and N. hispanica in the BMNH in
London, Christine Rollard for her friendly reception and provision of facilities for study of the
collection in the MNHN in Paris, Gilbert Caranhac for helping in the survey of the MNHN
collection and providing information on the male of N. brauni, and Nollie Hallensleben for
comments and corrections of the manuscript.

Chapter 3
Taxonomic Review of the Portuguese Nemesiidae (Araneae, Mygalomorphae).
Arthur Decae, Pedro Cardoso & Paul Selden
Research 2006
Revista Ibérica de Aracnología (2007) Vol. 14: 1-18.
Iberesia machadoi ♀

Abstract
The occurrence of the trapdoor spider family Nemesiidae in Portugal is reviewed on the basis
of recently collected material. The new sample of well over a hundred specimens (of which
97 were used for description) collected from locations throughout the country, contains six
different species, five of which we classify in the genus Nemesia, and one species that we
place in the recently erected genus Iberesia. Three species, N. athiasi, N. fagei and N.
uncinata, could be recognised from descriptions in the literature although no type material
was found. New information and figures of these species are provided. Two species, N.
bacelarae sp. n. and N. ungoliant sp. n. are newly described and illustrated. A description of
the female of N. uncinata and extended information on Iberesia machadoi are presented for
the first time. The Portuguese species list is discussed and updated.
Introduction
The trapdoor spiders of the family Nemesiidae collected in the course of a wildlife
conservation study in Portugal Cardoso 2004 proved difficult to identify at the species level.
The problem arose from a lack of up to date taxonomic information in the literature and the
unavailability of type material. Knowledge of Portuguese trapdoor spiders largely rests on the
publications of Frade & Bacelar 1931 and Bacelar (1932, 1933a, 1933b), who worked at a
time when modern key characters to distinguish species of European Nemesiidae (particularly
the morphology of sexual organs and spinnerets) were not practically used (Decae & Cardoso
2005). Our enquiries and searches for the type material of N. athiasi, N. gravieri, N. berlandi,
and N. fagei were unsuccessful. Notwithstanding this drawback we could recognise most
species from their original descriptions in the literature. Here, we present an update of the
species list of Portuguese Nemesiidae (see Cardoso 2000a for an earlier version) with
descriptions of two new species, Nemesia bacelarae sp. n. and N. ungoliant sp. n., and the
first description of the female of N. uncinata Bacelar 1933a. We recognise the following six
nemesiid species to occur in Portugal: N. athiasi Franganillo 1920, Nemesia bacelarae sp. n.,
N. fagei Frade & Bacelar 1931, N. uncinata Bacelar 1933a, N. ungoliant sp. n. and Iberesia
machadoi Decae & Cardoso 2005. New information on all these species is provided.
Material
A total of 97 specimens were used for description in this study. The bulk of the material (66
specimens) was collected in a program using pitfall traps carried out by P. Cardoso and coworkers
between 1997 and 2003 (see Decae & Cardoso 2005 for more detailed information).
This sample contained primarily male spiders of four different species (I. machadoi, N.
bacelarae, N. ungoliant, N. uncinata). Also included in this study were handcollected
specimens of N. fagei, from H. Buchli’s 18 1961/1962 collections1 (n = 15), N. uncinata
females from P. Selden’s 1996 collection (n = 4), N. athiasi, N. ungoliant and I. machadoi
females from A. Decae’s 1999 collection (n = 12).
Methods and Terminology
Methods used in collecting are described in Decae & Cardoso 2005. The methods of study
follow Decae 2005 and Decae & Cardoso 2005. SEM photography was carried out by
Cardoso at the Zoological Museum of the University of Copenhagen. Specimens chosen for
SEM work were gradually transferred from 70% to 100% ethanol in 10% steps. They were
then critical-point dried, coated in platinum and placed in appropriate stoppers for SEM.
18 Buchli’s collection contains two specimen of special interest. One is a female collected in 1937 by H. Mendez
at Foz do Rio Mira. It is present with the remarkable ‘burrow plug’ which N. fagei constructs (see Frade &
Bacelar 1931 and Bacelar 1933b). The other is a male collected by F. Frade in August 1935.

Abbreviations. Most terms and abbreviations used are standard in arachnological literature
and/or are previously explained in Decae 2005 and Decae & Cardoso 2005. For convenience
the following abbreviations are here given: BL = total body length measured from the apex of
the chelicerae to the most caudal tip of the abdomen; CL = length carapace, CW = width
carapace; SL = length sternum; SW = width sternum; LP = length palp; LL1 = length leg I;
LL2 = length leg II; LL3 = length leg III; LL4 = length leg IV; AR = length anterior eye row;
PR = length posterior eye row; EL = length eye-formation; Clyp = clypeus length measured in
a straight line from the anterior edge of the left ALE (anterior lateral eye) to the anterior edge
of the carapace; POP = deep black periocular pigmentation; MTF4 ratio = relative lengths of
metatarsus, tibia and femur of leg IV; PMS = posterior median spinnerets; PLS = posterior
lateral spinnerets. MNHN = Museum National d’ Histoire Naturelle, Paris; NMR =
Natuurhistorisch Museum Rotterdam; ZMUC = Zoological Museum University Copenhagen.
Qualitative characters. Traditional descriptions of Nemesia species have focused on colour
and spine patterns, and descriptions of the eye formation. Such information is of descriptive
value but is rarely sufficient for species diagnosis. Descriptions of the male palpal organ
found in the literature are of diagnostic value although older information and figures are
generally too superficial to make conclusive determinations possible. Here, we give more
detailed diagnostic information on the morphology of the sexual organs of both males and
females and descriptions of the following qualitative characters that were found to be of
diagnostic value: Crest zone, the colour pattern of the integument and the settings of setae on
the crest of the caput were found to vary consistently between most species. Pubescence, the
presence or absence and the colour of pubescent hair on the carapace, chelicerae and leg
segments may be of important diagnostic value in Nemesiidae at the species level. POP, the
pattern of the deep black pigmentation between and around the eyes has diagnostic value.
Fovea, the shape of the fovea and in particular the presence or absence of a short central
groove that runs perpendicular to the main recurved transversal groove of the fovea contains
diagnostic information. Fang ridge, the ventral prolateral ridge on the fangs may be smooth,
neatly serrated or irregularly broken (see Decae 2005 Figs. 2.6−2.8) in different species (and
sometimes in different sexes of one species). Furrow teeth, the number of teeth and the
position of the largest tooth on the prolateral edge of the cheliceral furrow is rather constant
within species, and variable between species. In our descriptions we number the furrow teeth
from distal to proximal (e.g. ‘tooth 5 largest’, in a row of six teeth means that the tooth most
proximal but one is the largest in the row). Cuspules, the number, pattern and shape of the
maxillary cuspules may vary between species. Spinneret morphology, (Figs. 38, 40, 42, 44,
46, 48) is an important diagnostic character in Nemesia species both to distinguish females
and to relate conspecific males and females. PMS, are always reduced in Portuguese Nemesia
and always absent in Iberesia (see Decae & Cardoso 2005 Figs. 1-9 & 1-10), the shape of the
PMS and the pattern of spigots however, are of important diagnostic value at the species level
in Nemesia. PLS, the relative length of the basal segment (seen in ventral view) and the
pattern of spigots on this segment are of diagnostic value. Maculae, the presence of dark
pigmented blotches (maculae) on the external leg segments and/or on the external basal
segment of the PLS (Fig. 38) is a conspicuous character that has had much attention in
literature and has led to much confusion in Nemesia taxonomy. The presence or absence of
maculae is of diagnostic value when used in combination with other characters. MTF4 ratio,
the relative lengths of metatarsus, tibia and femur of leg IV are of diagnostic value and given
here e.g. as Me4>Ti4; Ti4=Fe4 (metatarsus IV is longer than tibia IV, and tibia IV is as long
as femur IV).

Distribution of species. Although our knowledge on species distribution in the Iberian
Peninsula is rapidly growing, it is still far from complete. This is especially true for taxa, such
as the Nemesiidae, that contain a large number of endemics that usually are very restricted
in space (Melic 2001). A way of trying to overcome this impediment is to use ecological
niche modelling. Using confirmed and georeferenced data on the one hand, and data on
climatic variables in space on the other, it is possible to infer the potential distribution ranges
of species. Most of the existing techniques try to determine the range of environmental
variables in which the species is known to exist and extrapolate the known distribution to sites
where the species has not actually been found, but where environmental conditions are similar
to the ones found in the known area of distribution (e.g. Segurado & Araújo 2004, Elith et al.
2006). This bioclimatic envelope, when transferred to a spatial analysis tool, is reflected in a
map with the potential distribution of the species. The potential distribution is a measure of
adequacy of the area suitable to the species, taking the climatic variables into account. From
all the options available, some require both presence and absence data and others rely on
presence data alone. Given that absence data are usually very hard to obtain for spiders
(and most invertebrates) our options are somewhat restricted. We have chosen to use the
DOMAIN method (Carpenter et al. 1993) for a number of reasons: (1) it uses presence-only
data; (2) it is one of the best methods overall, usually as efficient as more complex methods
(e.g. Elith et al. 2006; Hernandez et al. 2006); (3) it is simple to use and moreover
implemented in free GIS software (http://www.diva-gis.org); (4) its results are easily
amenable to expert knowledge scrutiny. Expert knowledge is critical with automated methods,
because these usually do not consider historical and biological factors, which may restrict the
true distribution of the species to an area smaller than its potential distribution. Because we
realise that our ‘expert knowledge’ of the species we discuss is necessarily incomplete we
have chosen to present tentative modelling of the distributions of Portuguese nemesiids with
likely extensions into Spain. To do so we have primarily relied on the DOMAIN results. The
maps produced by this method are scaled, with probabilities of presence attributed to 1 km 2
squares covering the entire Iberian Peninsula. We used a conservative approach, with the cutoff
value being the maximum value for probability of occurrence that would connect all areas
with known records (between 60% and 80% in all cases). We then proceeded to eliminate
areas that were part of the predicted potential distribution but that contained no known records
and that remained isolated, irrespective of their size. In this way we have limited the potential
distribution to an area that should be closer to the realised niche. To make our results
amenable to future research the following detailed information is provided: we used 19
bioclimatic variables as provided by the Worldclim database (see Hijmans et al. 2005) derived
from temperature and rainfall values, with a spatial resolution of about one square kilometre:
annual mean temperature, mean monthly temperature range, isothermality, temperature
seasonality, maximum temperature of warmest month, minimum temperature of coldest
month, temperature annual range, mean temperature of wettest quarter, mean temperature
of driest quarter, mean temperature of warmest quarter, mean temperature of coldest quarter,
annual precipitation, precipitation of wettest month, precipitation of driest month,
precipitation seasonality, precipitation of wettest quarter, precipitation of driest quarter,
precipitation of warmest quarter, precipitation of coldest quarter. In the resultant modelling
we claim only to visualise potential (not actual) distributions. Of course, any model is only as
good as the data that it relies on, but we believe to have created distribution maps that
approach reality better than maps showing only the few known localities for each species
(Figs. 49−54).

Taxonomy
Nemesia athiasi Franganillo, 1920 Figs. 1, 2, 13, 14, 25, 26, 37, 38, 49
Types. Nemesia athiasi Franganillo 1920: p. 140. Female, Portugal. Frade & Bacelar 1931: p.
237. Bacelar 1932: p. 18, 21. Roewer 1942: p. 179. Bonnet 1958: p. 3036. Melic 2001: p. 85.
Cardoso 2000a: p. 20. New synonymy: N. gravieri Frade & Bacelar 1931: 228, f. 12–13
(Dm). Bacelar 1932: p. 20. Schenkel 1938 p. 1-14: Perez de San Román and de Zárate 1947.
Cardoso 2000a: p 20. Cardoso 2000b: p. 35. Melic 2001: p. 77. New synonymy: N.
meridionalis Frade & Bacelar 1931. (Misidentification in part, as reported from Portugal): p.
228–230. Bacelar 1932: p. 20. Bacelar 1933b: 292-293. Buchli 1969: p. 178, 191. Blasco
1985: p. 9. Blasco 1986: p. 345-346.Cardoso 1998. Cardoso 2000a: p. 20. Cardoso 2000b: p.
35.
Diagnosis. Differs from most Nemesia species by the occurrence, absent in some males, of
maculae (dark pigmented blotches) on the external surfaces of leg segments and the basal
segment of the PLS (Fig. 38). This character is shared with the species of the N. maculatipesgroup
(Decae 2005 p. 166-167) that is distributed in Italy and on islands in the western
Mediterranean. N. athiasi, however, differs from the N. maculatipes- group by having serrated
fang ridges (smooth ridges in the N. maculatipes-group), short, straight spermathecae (Fig.
37), (long and twisted in the N. maculatipesgroup), and a sub-apical serrated ridge on the
embolus tip (Fig. 26), (absent in the N. maculatipes-group).
Comments. We have not been able to find Franganillo’s type material, nor the specimen
studied by Frade & Bacelar that led them to conclude that N. athiasi Franganillo, is
insufficiently described and probably synonymous with N. meridionalis and/or N. sanzoi
(Frade & Bacelar 1931 p. 237). Also, we did not find the type of N. gravieri Frade & Bacelar
1931 that we regard synonymous with N. athiasi. We have indications (but no certainty) that
all this material is lost. Our results and conclusions are based on recently collected spiders
from various locations in Portugal in which we recognise the species from the texts of the
original descriptions of N. athiasi Franganillo 1920 and N. gravieri Frade & Bacelar 1931.
Description. Males (N = 14)
Measurements (mm). BL=11.2–14.9, CL=4.5–5.9, CW=3.7–4.9, SL=2.4–2.8 SW=1.9–2.3
LP=6.1–7.2, LL1=16.1–17.8, LL2=15.9–17.5, LL3=15.7–17.6, LL4= 20.6–22.8.
Patellar spine formulae PSPvar (n=14) [p=0 (1–2); I=0 (1–2); II=1(2–3); III=1(0–2–3);
IV=2(1–3)] RSPvar (n=14) [p=0–1; I=0–1–2; II=1–2–3; III=2 (1–3); IV=3 (1–2–3–4–6)]
Qualitative characteres. General appearence medium sized Nemesia species with
a distinct colour pattern on the carapace (Fig. 1), maculae might be vague or absent.
Pubescence black on slopes of caput, white in narrow longitudinal zones on the basal
segments of the chelicerae. Crest zone wide yellowish brown tapering towards the fovea, crest
setae in one well developed though somewhat irregular row, flanked on either side by rows of
smaller setae. Caput low (Fig. 13). POP dark black connecting all eyes. Fovea usually (7 out
of 9 specimens) without a central groove. Fang ridge serrated (rarely smooth: 1 out of 14
specimens). Furrow teeth six (no. 5 is largest, no. 1 smallest). Cuspules usually 4 to 6 in
mostly irregular rows, total range observed 0–7. Spinnerets as in female. Maculae clearly
present on legs and basal segment of PLS in 8 out of 19 observed specimens, less clear or
even absent in the other 11 specimens. MTF4 ratio Me4≥Ti4>Fe4.
Description. Females (n = 6)
Measurements (mm) BL=13.3–20.0, CL=4.6–7.1, CW=3.7–5.9, SL=3.1–4.3, SW=2.8–3.5,
LP=6.7–11.7, LL1=10.6–18.6, LL2=9.4–17.0, LL3=9.2–16.2, LL4=14.6–23.8.

Patellar spine formulae. PSPvar (n=5) [p=0; I=0; II=0 (1); III=0 (1); IV=2(1–3)] RSPvar
(n=5) [p=0 (1); I=0 (1–2); II=0 (1–2); III=2; IV=2 (1)]
Qualitative characters. General appearence (Fig. 2) as given for males, but maculae always
present (Fig. 38). Pubescence locally dense on carapace, black radiating from the fovea. Crest
zone wide, distinct in colour, tapering towards the fovea, single row of crest-setae. Caput
elevated (Fig. 14). POP connecting all eyes. Fovea without central longitudinal
groove. Fang ridge serrated. Furrow teeth usually six (7 and 8 in two specimens). Cuspules
club-shaped 4 tot 14 in irregular row, or double rows. Spermathecae (Fig. 37). Spinnerets
(Fig. 38). PMS reduced, somewhat coneshaped. PLS spigots restricted to the ventral distal
half, length proximal segment ≥ median + distal segment. Maculae present on the legs, not
always distinct on the spinnerets. MTF4-ratio Ti4>Fe4>Me.
Distribution (Fig. 49). Table 1. Summarises all currently recorded collection sites for N.
athiasi. The species is distributed in north-eastern, western and southern Portugal. Dots in Fig.
49 indicate locations where N. athiasi was collected. On grounds of the method we used for
range estimation (Carpenter et al. 1993) the species is expected to be distributed towards the
east into Castilla y León and Andalusia, Spain.
Natural history and nest type. A population of N. athiasi was found near Porto de Mós
(N39º36.140, W08º49.046) at the entrance of the Parque Natural das Serras de Aire
Candeeiros by Decae on 17 July 1999. The burrows are dug in pockets in limestone outcrops
on a garigue covered slope at the edge of a forest. Locally burrows occur close together, with
trapdoors of adult spiders less than 10 cm apart: in other places burrows occur more isolated.
All burrows had branching tubes (e.g. see Moggridge 1874p. 214, Plate XVII) with two
surface openings. Each entrance is closed off by a dorsally (highest part of the burrow rim)
hinged trapdoor, both of approximately similar size and shape. The typical
‘wafer-type’ trapdoors that Moggridge described for some Nemesia species in southern
France are flexible silken constructs. The wafer-door of N. athiasi is stiffer. Adult females
have trapdoors with diameters ranging from 13 mm to 17 mm, the corresponding entrance
openings are between 7mm and 8.5mm. In July the spiders are probably in aestivation
because the trapdoors are sealed with silk from the inside. No egg-sacs or juvenile spiders
were found in the burrows of adults. The branching burrow shafts meet at a depth of approx.
5 cm behind the trapdoors. Here they converge into a single shaft that extends to 15–25 cm
into the ground. The upper 2/3 of the burrow shaft, including both entrance tunnels, is lined
with thin silk that extends upward into the burrow rim, the hinge and the under covering of
the trapdoors. The deepest parts of the burrows are apparently not lined with silk. In some
burrows remains of prey were found. Analysis of remains shows ants, beetles and woodlice
being regular prey of N. athiasi. Wandering males were collected between August 13th and
November 28th.
Nemesia bacelarae sp. n. Figs. 3, 4, 15, 16, 27, 28, 39, 40, 50
Types. Male holotype: leg. P. Cardoso, nr. PMC0522A coll. 05 September 2001, deposited at
the MNHN Paris no. AR14324. Female paratype: leg. P. Cardoso, nr. BAP06 coll. 23 April
2002, deposited at the MNHN Paris no. AR14325. Other paratypes (5 males and 1 female)
included in the type collection are deposited in the NMR under the numbers: 9972.4042 -
9972.4047, and six males and one female are deposited in the ZMUC (under collection
numbers yet to be appointed).
Type locality. Bruçó, Tras-os-Montes, Portugal (N41º15.253, W06º43.542). A reforestation
area planted with Pseudotsuga menziesi.

Etymology. Named in honour of Amélia Bacelar who laid the basis for our knowledge of the
trapdoor spider fauna of Portugal.
Diagnosis. Differs from all known Nemesia species by the ‘fish-hook’ tooth on the embolus
(Fig. 28). Females differ from all Nemesia species by the spermathecae with a wide proximal
part, narrow median part that bends toward ventral, to connect with the globular distal part
(Fig. 39).
Description. Male holotype.
Measurements (mm) BL=11.9, CL=5.0, CW=3.9, SL=2.6, SW=2.1, LP=6.3, LL1=14.8,
LL2=13.5, LL3=12.6, LL4=17.6.
Patellar spine formulae. PSP [p=1–1; I=1–2; II= 2–2; III= 2–2; IV= 2–2]
RSP [p=0–0; I=0–0; II= 0–0; III= 1–1; IV= 1–1]
Qualitative characters. Dorsal habitus (Fig. 3).Carapace light brown, anterior and lateral caput
greyish brown, lateral caput speckled and blotched with irregular light brown dots,
pubescence greyish covering the whole carapace, crest setae only in anterior half of the crest
zone, fringe-, and thorax setae present but not very strongly. Clypeus darkest part of caput,
length equals approx. diameter ALE, 9 marginal setae in two sets of different strength. Caput
low, narrow in front (Figs.3 and 15). Eye-group rectangular almost twice as wide as long,
AR/EL = 1.8, AR/PR = 1.0, ALE largest ALE/PLE =1.1, AME less than one diameter apart,
POP dark shiny black connecting all eyes. Fovea: smoothly recurved, central groove absent.
Chelicerae light brown, not contrasting with the colour of the carapace, lateral longitudinal
zones with short setae and whitish pubescence, seven furrow-teeth, rastellum not very strong,
fang-keel serrated. Maxillae colour as sternum, distal process small and rounded, six
maxillary cuspules in irregular rows on either side. Sternum light yellowish brown, setae
cover even, but absent from three small central areas, edge setae longer and stronger than
those more centrally placed, anterior sigilla difficult to observe, centrals round touching
sternum edge, posteriors egg-shaped and less than their longest diameter from the sternum
edge. Labium light colour not contrasting with sternum, labial furrow shallow. Legs and palps
dorsal colour as carapace, ventral as sternum, no pattern, dorsal femora with dense black
pubescence (most prominent in legs III and IV), spine patterns no external spines on first
three appendages, trichobothria patterns as typical for the genus 19 , scopulae very light and not
extending on the tibia, paired claws with well developed double rows of denticles. Abdomen
evenly covered with fine hairs and bristles, dorsal light greyish yellow with slightly darker
anterior setae field, vague cardiac line and two indistinct chevrons just above the spinnerets,
ventral uniform light yellow. Spinnerets similar colour as ventral abdomen, PMS reduced
club-shaped, PLS basal segment and median segments approx. equal length, distal segment
shorter with well developed apical spigot field. Clasper hook somewhat flattened, smoothly
bent towards prolateral. Clasper field on slightly bulging integument just proximal of ventral
middle metatarsus, and placed towards prolateral. Bulb with clear fishhook embolus. MTF4
ratio Ti4≥Me4, Me4=Fe4.
Description. Female paratype
Measurements (mm) BL=17.2, CL=6.1, CW=4.6, SL=3.2, SW=2.5, LP=9.0, LL1=13.7,
LL2=12.5, LL3=11.9, LL4=18.7.
19 In approx. forty species of Nemesia and Iberesia examined the patterns of trichobothria were found to be
constant. On the tarsi a central dorsal zigzag line, on the metatarsi a compact distal group followed by an almost
straight line, on the tibia two, towards proximal, diverging lines. Distal bothria in all groups and lines longest,
proximals shortest.

Patellar spine formulae. PSP [p=0–0; I=0–0; II= 0–0; III= 2–2; IV= 0–0] RSP [p=0–0; I=0–0;
II= 0–0; III= 0–0; IV= 0–0]
Qualitative characters. Dorsal habitus (Fig. 4). Carapace different shades of brown without a
clear pattern, darkest on the lower and posterior lateral caput, around the fovea and in two
narrow lines along the implant on the chelicerae, crest zone lighter brown, crest setae in one
row with parallel groups of finer setae on either side, fringe setae indistinct,
back pubescence in and along the cervical furrows, white pubescence in the lower crest zone.
Clypeus length approx. equal to diameter of ALE, short row of 5 strong forward curved
marginal setae. Ocular process anterior rounded with small group of backward curved setae.
Caput slightly elevated (Fig. 16), abruptly narrowing towards fovea (Fig. 4). Eye group
rectangular, AR/PR= 1.0, more than twice as wide as long, PR/EL= 2.2, ALE largest,
ALE/PLE = 1.3, AME one their diameter apart. POP broken between AME and between
AME and the lateral eyes. Fovea lightly recurved without central groove. Chelicerae darker
brown, contrasting with the colour of the carapace, crest of the basal segment lighter brown, a
narrow lighter zone with white pubescence longitudinally on flanks, eight furrow teeth all
strongly developed and closely set, rastellum triangular fields of strong rigid spikes directly
bordering the fang implant. Fang ridge irregularly serrated. Maxillae with small anterior distal
process, cuspules club shaped, placed in rows. Sternum yellowish brown, darker along the
edges, dense cover of black setae along the edges and in the lateral zones, less dense in the
central parts. Sigilla well defined anterior and median pair round, posterior pair oval. Labium
twice as wide as long, labial furrow glabrous, wide, gently sloping. Legs and palps ventral
coxae shading in colour from warm orangeyellow on the maxillae to pale creamy yellow on
coxae IV. Legs generally coloured as carapace, ventral and lateral femora lighter creamy
yellow, spines most numerous and with strongest development on metatarsi III and IV and
lateral spines on Tibia IV, trichobothria patterns as typical for the genus, scopula dense on
palp tarsus, tarsi and metatarsi I and II, more lightly extending on distal palp tibia and
prolateral tibiae I and II. Abdomen evenly covered with fine hairs and bristles, uniform grey
dorsally, brownish ventrally. Spermathecae (Fig. 39) as described in diagnosis. Spinnerets
(Fig. 40). PMS cone shaped with a narrow top, PLS basal segment longer than median +
distal segment together, maculae absent. MTF4 ratio Ti4>Fe4>Me4.
Variation.
Males (n = 12)
Measurements (mm). BL=11.0–13.0, CL=3.9–5.4, CW=3.0–4.2, SL=2.1–3.2, SW=1.7–2.2,
LP=5.7–6.7, LL1=13.0–16.1, LL2=12.0–14.8, LL3=11.3–14.2, LL4=17.0–20.2.
Patellar spine formulae. PSPvar [p=2–1; I=2 (0–1); II=2(1); III=2(1–3); IV=2(0–
3)] RSPvar [p=0; I=0; II=0 (1); III=1 (2); IV=1 (0–3)]
Females (n = 3)
Measurements (mm). BL=16.5-17.9, CL=5.7-6.3, CW=4.5-4.8, SL=3.2-3.5, SW=2.5-2.6,
LP=9.0-9.4, LL1=13.7-13.9, LL2=12.5-13.0, LL3=11.9-12.5, LL4=18.7-19.1.
Patellar spine formulae. PSPvar [p=0; I=0; II=0 (1); III=2; IV=0] RSPvar [p=0; I=0; II=0;
III=0; IV=0.
Distribution. (Fig. 50), Table 1. Summarises all currently recorded collection sites for N.
bacelarae. The species is distributed in north-eastern, central western Portugal. Dots in Fig.
50 indicate sites where N. bacelarae was collected. The method we used for range estimation
(Carpenter et al. 1993) expects the species also to be found in southern Portugal and in
Castilla y León, Spain.

Natural history and nest type. All specimens known were collected in pitfall traps; hence no
information on the burrow of N. bacelarae is presently available. The species was collected in
various habitat types ranging from natural oak forests to cultured plantations
of Eucalyptus spec., and from maquis type of bush land to reforestation areas. Wandering
males were collected between September 5th and November 5th.
Nemesia fagei Frade & Bacelar 1931 Figs. 5, 6, 17, 18, 29, 30, 41, 42, 51
Types. Nemesia fagei Frade & Bacelar 1931 (Dmf); Bacelar 1933b: 291–292; 1937: 1571-
1576, f. 3–4 (Dm); Wiehle 1960: 459, f. 2 (m); Blasco 1986: 346, f. 1A (f). Cardoso 2000a:
p20.
Diagnosis. Males differ from all known Nemesia species by the shape of the embolus being
regularly curved, and pointed under low magnification Fig. (29) and slightly scooped and
twisted under high magnification Fig. (30). Females are distinguished by the concentration of
glandular tissue in the proximal part of the spermathecae extending into a collar that connects
the proximal part of the spermathecae with the distal part (Fig. 41). It is the only species
known to construct a curious ‘bullet-shaped’ burrow plug (Frade & Bacelar
1931; Bacelar 1933b).
Comments. All specimens studied are from Buchli’s collection that is housed in the MNHN in
Paris. Most of the specimens were apparently hand collected as juveniles by Buchli in 1961
and 1962 and raised to adulthood in his laboratory.
Description. Males (n = 4)
Measurements (mm). BL=7.0–11.0, CL=2.8–4.3, CW=2.1–3.1, SL=1.6–2.2, SW=1.2–1.7,
LP=4.1–6.1, LL1=8.1–12.8, LL2=6.9–11.2, LL3=6.6–10.5, LL4=9.9–15.1
Patellar spine formulae. PSPvar [p=0; I=0; II=1(2); III=1(0); IV=3(0–2)] RSPvar [p=0;
I=1(0); II=1(0); III=0 20 ; IV=1].
Qualitative characters. General appearance small, light coloured 21 Nemesia with
dark line around the carapace and dark dorsal pattern on the abdomen (Fig. 5). Caput low
(Fig. 17). Pubescence absent (very thin in 1 specimen in 4). Crest zone single row of strong
central setae flanked by rows of fine bristles. POP connecting all eyes. Fovea central groove
generally absent (present in 1 specimen in 4). Fang ridge serrated (smooth in 1 specimen in
4). Furrow teeth six. Cuspules absent. PMS and PLS as described for female. Maculae absent.
MTF4 ratio Ti4>Me4, Me4=Fe4.
Description. Females (n = 13)
Measurements (mm). BL=11.5–17.2, CL=3.7–5.2, CW=2.8–4.1, SL=2.1–3.0, SW=1.5–2.3,
LP=5.1–8.0, LL1=7.9–11.9, LL2=6.8–10.7, LL3=6.0–10.1, LL4=11.2–14.9]
Patellar spine formulae. PSPvar [p=0; I=0; II=0; III=0; IV=0(1–3)] RSPvar [p=0(1); I=0;
II=0(1); III=0; IV=0]
Qualitative characters. General appearance as described for males (Fig. 6).
Pubescence absent. Crest zone slightly darker in colour than surrounding integument. Caput
slightly elevated (Fig. 18). POP usually broken between the AME and the other eyes. Fovea
central groove generally absent (present in 1 specimen out of 13). Fang ridge serrated. Furrow
teeth six. Cuspules 3 to 5 in rows (rarely 1 or 2). Spermathecae (Fig. 41). Spinnerets (Fig. 42).
20 Groups of short strong spines on this segment, no singular spines.
21 The fact that the specimens used in this description have been preserved for over 40 years in alcohol might
have had an effect on their colour.

PMS reduced digitiform. PLS spigots on the ventral basal segment restricted to the distal half.
Maculae absent. MTF4 ratio Ti4>Fe4>Me4.
Distribution (Fig. 51). Table 1. Summarises all currently recorded collection sites for N. fagei.
To date, the species has only been found in coastal Alentejo and the Algarve. On the basis of
the method we used for range estimation it is expected to be found in coastal southwestern
Spain unless the Guadiana River proves to be a barrier to its dispersal.
Natural history and nest type. Details of the remarkable habits and nest-type of this species, as
well as other information on its natural history, are given in Frade & Bacelar 1931 and
Bacelar (1933b, 1937). We have no reliable information on the season that the males of N.
fagei emerge and wander in search of females. The males in Buchli’s collection were reared
in captivity and emerged in March and in May. One field record of a male of N. fagei was
taken in August 1935.
Nemesia uncinata Bacelar 1933 Figs. 7, 8, 19, 20, 31, 32, 43, 44, 52
Types. N. uncinata Bacelar 1933a: 285–287, f. 1-3 (Dm). Bonnet 1958: p. 3043. Jerardino et.
al. 1988: p. 358. Jerardino et. al. 1991: p. 145. Cardoso 2000a: p. 20. Cardoso 2000b: p. 35.
Melic 2001: p. 77.
Diagnosis. differs from all other Nemesia species by the grossly enlarged bulb with short
ornamented embolus (Figs. 31–32) and the simple dome shaped spermathecae (Fig. 43).
Comments. N. uncinata was described by Bacelar (1933 p. 287) as the possible male of N.
hispanica. Although the male of N. hispanica remains unknown it is clear that N. uncinata is a
separate species.
Description. Males (n = 2)
Measurements (mm) BL= 14.8, CL=6.6, CW=1.3.
Patellar spine formulae. PSPvar [p=1; I=2; II=2; III=3; IV=3] RSPvar [p=0; I=1; II=1;
III=1(0); IV=1]
Qualitative characters. General appearance medium to large sized Nemesia of an overall
slender and brownish appearance, with bands of silvery pubescence on the chelicerae and the
anterior carapace, and black pubescence on the central carapace (Fig. 7). Carapace with
narrow purplish fringe line. Caput low (Fig. 19). Chelicerae contrast in colour with carapace.
Dorsal femora with dense black pubescence and numerous spines on tibiae and metatarsi of
all legs. POP connecting all eyes. Fovea crescent shape, central groove absent. Fang ridge
smooth. Cuspules absent. Spinnerets as in female. Maculae absent. MTF4 ratio
Ti4>Me4, Me4=Fe4.
Description. Female reference specimen
nr. 22/08/96-2, Praia da Oura, Albufeira, Algarve (N37º05.273, W08º13.477) leg. P. Selden.
Measurements (mm) BL=20.5, CL=7.7, CW=5.9, SL=4.1, SW=3.1, LP=11.7, LL1=18.0,
LL2=17.0, LL3=16.2, LL4=23.5.
Patellar spine formulae. PSP [p=2–2; I=2–2; II=2–2; III=2–2; IV=0–0] RSP[p=0–0; I=0–0;
II=0–0; III=0–0; IV=0–0].
Qualitative characters. General appearance the adult female is a medium-sized Nemesia of an
overall slender shape and brownish colour with conspicuous silvery pubescence on the
chelicerae and the anterior carapace, the warm brown chelicerae contrast in colour with the
yellowish carapace (Fig. 8). Carapace yellowish brown with narrow purplish
fringe line, anterior carapace darkest with two lighter patches on either side lateral of the eye
group, dense silvery pubescence, caput-thorax junctions rich reddish brown. Caput elevated

(Fig. 20). Ocular process steep in front. Eye group posterior row longer than anterior row
(AR/PR = 0.93), width > length (PR/HE = 2.19), AME one diameter apart, diameter ALE =
PLE. POP broken between ALE-PLE and between ALE-AME. Chelicerae dark reddish
brown with longitudinal zones of brilliant white pubescence, colour contrasts with that of
carapace. Fangs: long, slender, sharp with a smooth keel. Rastellum compact group of strong
teeth on a small rastellar process. Abdomen dorsal, central zone, creamy white with a
complex pattern of dark pigmentation, lateral and ventral zones greyish. Fine setae with
undercover of fine hair. Ventral prosoma yellow, brownish around the edges. Sternum with
evenly distributed
setae, posterior sigilla ‘egg-shaped’. Labium slightly darker than sternum, labial furrow
comparatively narrow. Maxillae five strong somewhat spiky cuspules in irregular rows. Legs
and palps uniform brown except for the dark dorsal zone and the white pubescence on all
femora. Spermathecae (Fig. 43). Spinnerets (Fig. 44), PMS vestigial digitiform. PLS basal
segment longer than medium and distal segment together, spigots on proximal segment in a
triangular field, colour slightly lighter than ventral abdomen.
Variation.
Females (n = 4).
Measurements (mm). BL=20.5–23.2, CL=7.7–8.6, CW=5.9–6.6, SL=4.1–4.5, SW=3.1–3.5,
LP=11.7–12.5, LL1=18.0–19.4, LL2=17.0–18.4, LL3=16.2–17.5, LL4=23.5–26.6.
Patellar spine formulae. PSPvar [p=2; I=2; II=2; III=2; IV=0] RSPvar [p=0; I=0; II=0; III=0;
IV=0]
Distribution (Fig. 52). All currently recorded collection sites for N. uncinata are given in
Table 1 and are indicated as dots in Fig. 52. The estimated distribution range (dark zones in
Fig. 52) is restricted to the Algarve.
Natural history. We have no information about the structure of burrow of this species. The
species is apparently not uncommon in the far south of Portugal, though very limited in its
distribution. The males wander in autumn. Two males were collected in pitfall traps in a
Eucalyptus plantation in the first week of October. A third male (not included in our
description) was collected by S. Huber during heavy rain in the last week of October 2006
near Alte in the Algarve.
Nemesia ungoliant sp. n. Figs. 9, 10, 21, 22, 33, 34, 45, 46, 53
Types. Male holotype: leg. P. Cardoso, nr 5004-1 collected 24 Sept. 2002, deposited at the
MNHN Paris Nr. AR14327; Female paratype: leg. P. Cardoso, nr. 5003-2 collected 24 Sept
2002, deposited at the MNHN Paris Nr. AR14328. Other paratypes (11 males and 2 females)
included in the type collection are deposited in the NMR under the numbers: 9972.4029–
9972.4034, and seven males are deposited in the ZMUC (under collection numbers yet to be
appointed).
Type locality. Barrenta (N39º34.316, W08º45.686)
Eucalyptus plantation.
Etymology. Named after Tolkien’s mythical spider acting in “Silmarillion” and other
writings.
Diagnosis. The dark, shiny appearance, the lack of pubescent hair on the carapace, the double
row of crest setae, and the high elevation of the caput in females, N. ungoliant resembles the
species grouped in the ‘subgenus’ Haplonemesia (Simon 1914). It differs from the
Haplonemesia in lacking the diagnostic palpal spine pattern of this group, the much smaller
size of the adult spiders, the morphology of the spermathecae which are straight (compare

Fig. 45 with Blasco 1986 p.346 Fig.1c) instead of bent and the embolus being slightly hooked
instead of smoothly curved (compare Fig. 33 with Frade & Bacelar 1931 p.223 Figs. 1-2).
Description. Male holotype.
Measurements (mm) BL=10.4, CL=4.2, CW=3.4, SL=2.2, SW=1.7, LP=5.5, LL1=12.6,
LL2=12.4, LL3=12.2, LL4=16.4.
Patellar spine formulae. PSP [p=1–1; I=1–1; II= 2–2; III= 2–2; IV= 1–1] RSP [p=0–0; I=0–0;
II= 0–0; III= 1-1; IV= 1–1].
Qualitative characters. Dorsal habitus (Fig. 9). Carapace colour almost uniform
brown, with slighter darker areas around the eyes and the fovea, pubescence absent, crest
setae in irregular central longitudinal group setae of different strength flanked on either side
by a sub-row of thinner setae, fringe setae. Clypeus shorter than one diameter of ALE with
five equal-sized edge setae. Fringe setae and thorax setae strongly developed. Caput slightly
elevated (Fig. 21). Eye group trapezium with AR as shortest parallel side (AR/PR=0.9), width
> 2x height (AR/EL=2.3), ALE largest (ALE/PLE=1.4), AME’s less than their diameter apart.
POP dark black, connecting all eyes. Ocular process low and sloping both in front and behind.
Fovea sickle shaped recurved, central groove absent. Chelicerae basal segment uniform
brown, not contrasting with colour of carapace, lateral lines indistinct, seven furrow teeth,
fang-keel serrated. Maxillae distal process very small, cuspules, three on either side, in short
rows and pointed in shape. Sternum colour uniform yellowish brown, same colour as ventral
coxae and maxillae, uniform setae cover, anterior and median sigilla round, posterior sigilla
oval, distanced about their longest diameter from the sternum edge. Labium colour slightly
darker than sternum, labial furrow shallow, wide, glabrous. Legs and palps brown with
greyish longitudinal dorsal zones and black pubescence dorsally and laterally on the femora,
spines on all segments including the ventral tarsi, trichobothria patterns as typical for the
genus, scopula thin, all paired claws with double combs, maculae absent. Abdomen dorsally
dark colour, ventrally lighter, dense group recurved setae in anterior dorsal group, no clear
colour pattern. Spinnerets similar colour as ventral abdomen. PMS club-shaped, reduced.
PLS basal segment longer than median + distal segment, maculae absent. Clasper hook strong
and inwardly curved, clasper field restricted to central ventral metatarsus. MTF4 ratio
Me4

zones indistinct, seven furrow teeth. Fang ridge serrated. Maxillae with small anterior distal
process, five club shaped cuspules in irregular rows. Sternum uniform yellowish brown,
evenly covered with black setae, anterior and median sigilla round, posterior sigilla oval,
distanced about their longest diameter from the sternum edge. Labium slightly darker in
colour than sternum and separated by a wide shallow furrow. Legs and palps: dorsal anterior
appendages slightly darker brown than legs III and IV, all appendages uniform colour with the
exception of the lateral and ventral femora and the ventral coxae IV that are lighter coloured,
no lateral spines on external tibiae and metatarsi I & II, trichobothria as typical for the genus,
scopula on palp-tarsi and tarsi and metatarsi I and II, only slightly extended on distal
prolateral tibiae, combs on paired claws III and IV reduced, maculae absent. Abdomen
greyish yellow with vague dorsal pattern of darker chevrons and irregular blotches.
Spermathecae (Fig. 45) as described in diagnosis. Spinnerets (Fig. 46). PMS reduced, coneshaped
with one or two spigots at the apex. PLS basal segment with few small spigots in
distal half, more than twice as long as median + distal segments, no clear maculae, but some
separate irregular darker pigmented blotches present. MTF4 ratio Me4

the northern-most parts of the country. The method that was used for range estimation
indicates that the distribution of I. machadoi may extend into Andalusia, southern Spain. Dots
in Fig. 54 indicate actual collections sites of I. machadoi.
Natural history and nest type. I. machadoi constructs a classical cork-type burrow (Moggridge
1873, Plates VII and VIII) of approx. 20 cm deep which is often dug in exposed patches of
soil (road and creek banks, bare ground along railway tracks and on hill
slopes). The species occurs in a variety of habitat types that range from shady woodlands to
dry garigue and from public parks to olive yards. Egg-sacs are produced in summer when the
spiders aestivate closing their burrow with a thick clay plug just under the trapdoor.
Wandering males were collected in autumn, between October 17th and November 19th.
Key to Portuguese Nemesiidae species based on the morphology of genital structures
Males
1. Proximal bulb enlarged, very wide and bulbous, distal bulb very strong with short embolus,
tip of embolus sharply bent and ‘gouge shaped’ with a strong triangular side tooth (Figs. 31–
32)………………………………............................................................N. uncinata
2. Embolus broad, abruptly tapering near the tip and sharply bent convex side with a small but
conspicuous ‘serrated ridge’ (Figs. 25–26)…………………………. ...N. athiasi
3. Distal embolus elongated and slightly sigmoid with a distinct ‘fish-hook’ proximal on the
embolus tip (Figs. 27–28) ……………………………..........................N. bacelarae
4. Embolus slightly hooked with a smooth, ‘curled-up’ ridge near the embolus tip (Figs. 33–
34) .........................................................................................................N. ungoliant
5. Embolus regularly curved, ‘claw-shaped’ with a slightly twisted tip (Figs. 29–30)
...............................................................................................................N. fagei
6. Embolus long, somewhat irregularly curved, ending in a short counter curve at the tip.
Embolus tip slightly enlarged, ‘torpedo-shaped’ and preceded by a sharp ridge with two
prominent teeth (Figs. 35–36)……….. ................................................I. machadoi
Females
1. Spermathecae dome shaped evenly covered with dense glandular tissue (Fig. 43)
..............................................................................................................N. uncinata
2. Spermathecae short (approx. as long as wide), proximal tubular part approx. as short as
distal globular part……………….......................................................4.
3. Spermathecae distinctly longer than wide…………………...........6.
4. Proximal part conical, distal part of spermathecae spherical thinly covered with glandular
tissue. (Fig. 37) ...................................................................................N. athiasi
5. Proximal part short and tubular, distal part of spermathecae flattened on top, glandular
tissue concentrated in a thick, dense collar around the connection of the proximal and distal
parts (Fig. 41) ................................................................................... N. fagei
6. Proximal part longest, distally gradually tapering and connecting directly with distal part
(Fig. 45) ……………………............................................................N. ungoliant
7. Proximal part wide at the basis, abruptly tapering before connecting under an angle with the
distal part (Fig. 39) ..........................................................................N. bacelarae
8. Proximal part wide and irregularly shaped, a distinct narrow tubular middle part connecting
the proximal to the distal part (Fig. 47) …………………………...I. machadoi
Discussion
In her Inventário das Aranhas migalomorfas da Península Ibérica Amélia Bacelar 1932
reported the occurrence of six different Nemesia species in Portugal. Three species, N.
berlandi, N. gravieri and N. fagei, were newly described one year earlier from specimens of

Portuguese origin found in the collection of the Museum Bocage, in Lisbon (Frade & Bacelar
1931). The other three species, N. dubia, N. hispanica and N. meridionalis, with type
localities in France, Spain and Italy respectively, were identified in specimens collected in
Portugal (Frade & Bacelar 1931). A year later Bacelar 1933a added a seventh species to the
Portuguese list by describing N. uncinata from a male spider collected in southern Portugal.
The species list of Nemesiidae occurring in Portugal we present here is reduced to six species
again, although our removal of N. berlandi is only tentative. Furthermore there are indications
of the presence of still other species present within Portuguese borders, but we have no
conclusive information so far. Only one species of Bacelar’s 1932 original list, N. fagei, still
features on our list. All changes we made in the list and our motivation to do so are discussed
below.
N. berlandi Frade & Bacelar 1931 was described from female specimens (the male has not
been described) collected near Fagilde in north-central Portugal. The species has, to our
knowledge, not been reported from any other location and a search by Cardoso at the type
locality has not been successful. Also we have not been able to trace any specimen in museum
collections attributed to this species by earlier workers in the field, although Blasco (1986
p.346 Fig. 2E) figures the spermathecae of N. berlandi with a note: d’après Buchli.
Though we have examined Buchli’s unpublished manuscript that gives the original illustration
of the figure Blasco reproduced, we failed to find the specimen that was used to produce it.
Buchli’s figure of the spermathecae of N. berlandi does not fit any of the species in our
sample and we are therefore inclined to regard N. berlandi as a possible seventh species,
occurring in Portugal that escaped our collection efforts.
N. gravieri was described from a single male specimen collected at Alcacerdo Sal, southcentral
Portugal. This species, as N. berlandi, has been reported from its type locality only.
Contrary to N. berlandi however, N. gravieri is abundantly present in our sample. It is
immediately recognised from the conspicuous maculae on the legs and the basal segment of
the PLS. This obvious character is mentioned in the species description of N. gravieri (Frade
& Bacelar 1931 p. 230), and also in the earlier description of N. athiasi Franganillo, 1920 22
probably collected in the Lisbon area. The maculate character, in Bacelar’s days, was
considered diagnostic for yet another Nemesia species (N. meridionalis Costa 1835) from
Italy that is known from the female only (Frade & Bacelar 1931 p. 228). Consequently Frade
& Bacelar diagnosed all maculate females in the collection of the Museum Bocage Lisbon as
N. meridionalis and included this species on the Portuguese species-list (Bacelar 1932). They
went on to regard Franganillo’s 1920 description of N. athiasi as insufficient and claimed that
his species is probably synonymous with N. meridionalis (Frade & Bacelar 1931 p. 237). It is
a bit of a puzzle why Frade & Bacelar did not record the first maculate male they found in
Portugal also as N. meridionalis, but instead described it as a new species, N. gravieri.
Currently, several maculate Nemesia species are known (see Decae 2005 for a detailed
discussion) and it is clear that the species occurring in Portugal differs from those occurring
elsewhere (see our diagnosis of N. athiasi). Since Franganillo clearly described maculae in N.
athiasi, we regard N. gravieri as a junior synonym of N. athiasi. We also remove N.
meridionalis from the Portuguese list on grounds here explained and we regard the specimens
diagnosed by Frade & Bacelar as N. meridionalis from Portugal to be N. athiasi.
N. dubia Pickard-Cambridge 1874 is another Nemesia species with a confused taxonomic
history. It is clear however, that N. dubia is common in southwestern France and probably
22 “Mamillarum superiorum primus articulus magna macula nigra ornatus.” Franganillo 1920. p. 140.

also in Catalonia (type locality eastern Pyrenees). We have studied several specimens of both
sexes collected at the type locality and nearby areas that make us conclude that the
morphology of the sexual organs of this species clearly differs from all species we have found
in Portugal. Therefore we remove N. dubia from the Portuguese species list.
N. hispanica Koch 1871 has long been considered the most commonly occurring trapdoor
spider in Portugal. However, we have argued before (Decae & Cardoso 2005) that this idea
rested on misidentification. N. hispanica is not present in our sample of Portuguese species,
but because it is common in Spain, it might well be discovered on the Portuguese side of the
border in the future. Until then however, we remove N. hispanica from the Portuguese species
list.
N. uncinata was described from a male specimen, and although Bacelar expressed some doubt
about the separate identity of N. uncinata, stating that it might be the unknown male of N.
hispanica (Bacelar 1933 p. 287). It is now clear that, on grounds of the unusual morphology
of the sexual organs in both sexes, N. uncinata is a distinct species and properly features on
the Portuguese species list.
Acknowledgements
We thank the Portuguese Institute for Nature Conservation, in particular all the people that
made our work possible in the National Park of Peneda-Gerês, the Nature Parks of Douro
Internacional, Serras de Aire e Candeeiros, Arrabida, Sudoeste Alentejano e Costa Vicentina
and Vale do Guadiana, and the Nature Reserve of Paúl do Boquilobo. We also thank Christine
Rollard and Gilbert Caranhac of the MNHN in Paris for making Buchli’s collection of N.
fagei available for study and Siegfried Huber for sending specimens and sharing his new field
data on N. fagei, N. uncinata and I. machadoi with us. We thank Barbara York Main, Brent E.
Hendrixson and Nollie Hallensleben for their valuable comments and suggestions that helped
improve the text. Special thanks go to Israel Silva, Sérgio Henriques, Luis Crespo and Ana
Cerveira who did much of the fieldwork.

LOCATION Latitude Longitude I. machadoi N. athiasi N. bacelarae N. fagei N. uncinata N. ungoliant
Albergaria 41º47.718 08º08.180 X
Albufeira 37º05.274 08º15.115 X
Alcácer do Sal 38º22.347 08º30.773 X
Algodôr 37º44.956 07º48.027 X X
Aljustrel 37º52.830 08º09.898 X
Alte 37º14.000 08º11.000 X X X
Bairro 39º34.087 08º37.022 X X
Barrenta 39º34.316 08º45.686 X X
Braciais 37º37.948 07º34.199 X
Bruçó 41º15.253 06º43.542 X
Cabo Espichel N 38º24.936 09º12.988 X
Castelo de Vide 39º24.958 07º27.339 X
Castro Marim 37º13.110 07º26.516 X X
Coimbra 40º12.428 08º25.770 X
Corredoura 37º44.774 07º38.530 X X
Corte da Velha 37º41.239 07º43.716 X
Escusa 39º23.544 07º24.580 X
Faro 37º00.909 07º56.056 X
Fátima 39º37.072 08º39.161 X
Fonte d'Aldeia 41º25.372 06º24.251 X
Foz do Mira 37º43.281 08º47.305 X
Guadalupe 38º34.234 08º01.272 X
Guarda 40º32.314 07º16.036 X
Herdade do Pinheiro 38º27.689 08º43.199 X
Lagos 37º05.965 08º40.264 X X
Mata do Solitário 38º27.711 09º00.117 X
Mata do Vidal 38º29.198 08º59.552 X X
Mazouco 41º10.131 06º47.855 X
Mértola 37º38.576 07º39.673 X X
Palão 41º07.655 06º49.225 X X X
Palmela 38º33.946 08º54.132 X
Paúl do Boquilobo 39º23.385 08º32.478 X X
Penina 37º09.712 08º33.918 X
Picote 41º22.900 06º20.700 X
Picotino 41º12.410 06º45.646 X
Planalto de St António 39º30.469 08º42.239 X
Pomarão 37º34.500 07º32.100 X
Porto de Espada 39º21.308 07º21.024 X
Porto de Mós 39º36.140 08º49.046 X X
Praia da Falésia 37º05.328 08º10.130 X
Praia da Oura 37º05.273 08º13.477 X
Praia da Rocha 37º07.031 08º32.134 X
Ramalhais 39º53.949 08º31.490 X
Relva do Lobo 37º09.118 08º53.081 X X
Ribeira de Limas 37º49.233 07º37.065 X
Sagres 37º00.193 08º56.678 X X
São Domingos 37º39.676 07º29.289 X
São Martinho do Porto 39º30.684 09º08.114 X
Serra do Caldeirão 37º14.341 07º56.232 X
Serro Ventoso 39º33.372 08º50.293 X X
Setúbal 38º31.435 08º53.535 X
Sintra 38º48.062 09º23.008 X
Tavira 37º07.585 07º39.000 X X
Terras do Risco 38º27.765 09º02.038 X
Tó 41º18.950 06º34.442 X
Vale da Rasca 38º30.680 08º58.837 X X X
Vale Garcia 39º32.989 08º35.272 X X X X
Vila Chã da Braciosa 41º24.940 06º21.395 X
Vila Real de St António 37º11.671 07º24.926 X
LOCATION Latitude Longitude I. machadoi N. athiasi N. bacelarae N. fagei N. uncinata N. ungoliant
Table 1 Summary of locations (X) where nemesiids have been found in Portugal. Latitude in degrees North, longitude in
degrees West.

1 2 3 4
5 6 7 8
9 10 11 12
Figs. 1-12 Nemesiidae from Portugal (impression of comparative sizes and dorsal habitus). Males left of
conspecific females. 1-2 Nemesia athiasi; 3-4 N. bacelarae; 5-6 N. fagei; 7-8 N. uncinata; 9-10 N.
ungoliant; 11-12 Iberesia machadoi. Scale line = 10mm.

13
15
14
16
17
19
18
20
21
23
22
24
Figs. 13-24 Nemesiidae from Portugal, lateral prosoma comparative size and shape. Males above.
13-14 Nemesia athiasi; 15-16 N. bacelarae; 17-18 N. fagei; 19-20 N. uncinata; 21-22 N. ungoliant;
23-24 Iberesia machadoi. Scale line 10mm.

25 26 27 28
29
30 31 32
33
34 35 36
Figs 25-36 Bulb structure, general bulb shapes, and close-ups of diagnostic structures on embolus tip. 25-26
Nemesia athiasi; 27-28 N. bacelarae (arrow indicates fish hook); 29-30 N. fagei; 31-32 N. uncinata; 33-34 N.
ungoliant; 35-36 Iberesia machadoi.

37
43
39
45
41
47
38
40 42
44 46 48
Figs. 37-48 Diagnostic characters in females (not to scale). Spermathecae (above), spinnerets (below). 37-38
Nemesia athiasi (arrow indicates maculae); 39-40 N. bacelarae; 42-42 N. fagei; 43-44 N. uncinata; 45-46 N.
ungoliant; 47-48 Iberesia machadoi.

49
50
51
52
53
54
Figs 49-54 Estimated areas of distribution (black zones) of Portuguese Nemesiids. 49 Nemesia athiasi; 50 N.
bacelarae; 51 N. fagei; 52 N. uncinata; 53 N. ungoliant; 54 Iberesia machadoi.

Chapter 4
Iberesia, a new genus of trapdoor spiders (Araneae, Nemesiidae) from Portugal
& Spain.
Arthur Decae & Pedro Cardoso
Research autumn 2005
Revista Ibérica de Aracnología (2005) Vol. 12: 3-11.

Abstract
A new genus, Iberesia, is formed to receive species endemic to the Iberian Peninsula
traditionally included in Nemesia Audouin 1826, but to be distinguished by the absence of
posterior median spinnerets.
Iberesia includes I. machadoi sp. n., the type species, which is widely distributed in Portugal
as well as two species transferred from Nemesia: I. brauni (Koch 1882) comb. n. from the
Balearics and I. castillana (Frade & Bacelar 1931) comb. n. from Avila, central Spain. We
group Iberesia with Nemesia and Brachythele in the tribe Nemesiini Raven 1985. The
characters on which this grouping is based are discussed. Also discussed is the taxonomic
confusion about the identity of Nemesia hispanica (L. Koch 1871) in relation to that of I.
machadoi sp. n. The type of Nemesia hispanica is considered to be an adult female.
Introduction
At least three authors to date failed to notice the absence of the posterior median spinnerets
(PMS), present in most mygalomorphs (Raven 1985), in some species previously placed in
the genus Nemesia. L. Koch 1882 described Nemesia brauni from a male and a female but,
despite the detailed description, the absence of the PMS was not noted. Much later, in newly
describing Nemesia castillana, Frade & Bacelar 1931 also failed to note the absence of the
PMS. António de Barros Machado 1944 was the first to report the reduction of the posterior
median spinnerets (PMS) in several Nemesia species and the complete absence of these
spinnerets in a species he took to be Nemesia hispanica L. Koch 1871. Based on these
observations, Machado (1944 p. 26) concluded that N. hispanica was not only the first
ctenizid known to possess only one pair of spinnerets (the posterior laterals, PLS), but also
that Nemesia is the only spider genus in which the number of spinnerets was variable (either
one or two pairs are present). From our point of view on mygalomorph systematics
Machado’s conclusions have to be modified. First, Nemesia is no longer placed in the family
Ctenizidae since Raven 1985 transferred it to the family Nemesiidae; second, we find that the
species that Machado studied was not N. hispanica because a re-examination of the type
specimen in the BMNH on our request (Hillyard pers.comm.) revealed that N. hispanica
actually possesses PMS. Our studies on newly collected Nemesiidae indicate that the PMS
(although present in N. hispanica) are missing in several described and undescribed Iberian
species, one of which is common and widespread in Portugal. From this we conclude that
Machado’s observation on the missing PMS in Portuguese Nemesiidae refers to a hitherto unnamed
species that we here describe as Iberesia machadoi sp. n., and that this species (on
grounds of the shared absence of the PMS) should be grouped with I. brauni (comb. n.) from
the Balearics, and I. castillana (comb. n.) from central Spain in the new genus that we have
named Iberesia gen. n.
Methods and Terminology
Methods: Males of Iberesia machadoi sp. n. studied were collected in a pitfall trapping
program running between 1997 and 2003 at various locations in Portugal. Females were hand
collected between 1999 and 2003. Information provided on I. brauni (L. Koch 1882) comb.n.
is based on a recent study by Decae 2005, for I. castillana (Frade & Bacelar 1931) comb.n.
the male type specimen in the MNHN was re-examined. The pitfall trapping program was
conducted by Cardoso and co-workers in a series of protected areas in Portugal. These were
the Nature Parks of Arrábida (PNA) during 1997 and 1998, Douro Internacional (PNDI)
during 2001, Serras de Aire e Candeeiros (PNSAC) during 2002 and Vale do Guadiana
(PNVG) during 2000 and 2003, the Nature Reserve of Paúl do Boquilobo (RNPB) during
2002. The traps were filled with ethylene glycol 50% diluted with water, placed in each of the
main habitat types present at each area, and checked every 2 weeks during about one year at

each site. All specimens were transferred to 70% ethanol, studied and photographed fully
submerged in this medium under a CETI-MEDO-2 binocular, as described by Decae 2005.
Terminology: most terms used are standard in arachnological literature (e.g. Decae 2005 p.
146). The following terms are of descriptive or diagnostic value in the Nemesiini
(Nemesiidae, Nemesiinae). Crest-zone: a usually contrasting lighter coloured narrow
longitudinal zone running centrally over the highest part of the caput in most Nemesiini. In
Iberesia, the crest-zone is generally indistinct in males and females (Figs. 1-2). Crest-setae:
recurved setae placed in the crest-zone, usually in one straight longitudinal row, but
occasionally more irregularly placed in both sexes. Fringe-setae: outwardly directed setae
along lateral and caudal edges of the carapace, variable in conformation between species and
distinct only in males. Furrow-teeth: teeth arranged in prolateral longitudinal rows along the
cheliceral furrow can be diagnostic in their total number and relative sizes. The teeth are
numbered from distal to proximal (Fig. 3). Super-spine: distinctly stronger built, mostly
curved and obliquely upward directed spine set in a particularly large socket on retrolateral
tibia III (Fig. 4). Clasperfield: zone with specialised short setae, often in species specific
setting, on the ventral metatarsus I of males (Fig. 6).
Descriptive formulae and ratios
Spine patterns are variable and difficult to interpret unambiguously in Nemesiini, and have
therefore been of minor use in systematic descriptions of this group (Blasco 1986; Cardoso
2000; Decae 2005). However, the patellar spine patterns can be interpreted relatively
unambiguously and therefore are of taxonomical value, particularly if the observed variation
can be summarised in a short formula. The following descriptive formulae are used to
summarise the patellar spine patterns (and variation therein) for I. machadoi. (See also Decae
2005): prolateral (PSP) and retrolateral (RSP) patellar spine formulae for the patterns in one
specimen such as the male holotype and the individually described female paratype. An
example: PSP [p=0-0; I=1-1; II=1-1; III=2-1; IV=0-0] means that there are no prolateral
spines on either of the palp patellae, one prolateral spine on patellae I & II (one left and one
right), two spines right and one spine left on patellae III, and no prolateral spines on patellae
IV. PSPvar and RSPvar are used to summarise the variation in patellar patterns in the type
sample (7 males and 10 female paratypes). An example: RSPvar [p=1(0-2); I=1; II=1(2);
III=1(0-2); IV=0] means that usually there is one retrolateral spine on a palp patella, but there
might be none or two, there is invariably one spine on retrolateral patella I; patella II usually
has one retrolateral spine, but occasionally two and so on. Quantitative characters of the eyegroup
are summarised in three ratios: anterior row width/eye-group length (AR/EL), anterior
row width/ posterior row width (AR/PR), maximal diameter anterior lateral eye/maximal
diameter posterior lateral eye (ALE/PLE). The eyes are measured as in Decae 2005.
Length/width ratios are abbreviated as (l/w) under the description of the segment concerned.
Abbreviations of institution names: MNHN (Museum National d’Histoire Naturelle, Paris),
NMR (Natuurhistorisch Museum Rotterdam), BMNH (Natural History Museum, London).
Iberesia new genus
Type species: Iberesia machadoi sp. n. Other species included: Iberesia brauni (L. Koch
1882) comb. n., Iberesia castillana (Frade & Bacelar 1931) comb. n.
Etymology: the generic name is a noun derived from the name of the Iberian Peninsula where
the genus was discovered and appears to have a wide distribution.
Genus diagnosis: member of the tribe Nemesiini sensu Raven (1985 p.47). Iberesia differs
from other Nemesiini (Nemesia & Brachythele Ausserer 1871) by the absence of the posterior

median spinnerets (PMS) and consequently the possession of only one pair of spinnerets (Fig.
9), the posterior laterals (PLS). Other diagnostic characters are: the short, single rows of spiky
cuspules (n= 3 – 6) along the proximal edge of the maxillae (Fig. 7), the presence of a superspine
(Fig. 4) retrolaterally on tibia III of the females.
Genus description: Robustly built Nemesiini (total lengths males (mm) 12.5-18.5; females
17.5- 31.5) of a general brownish appearance. Strong sexual dimorphism. Females with
elevated caput, smooth fang keel, no patellar spines, reduced combs on inner paired claws of
legs III, IV. Males with low caput, serrated or smooth fang keel, spines on leg patellae, well
developed combs on inner paired claws of legs III, IV. Males may have distinct colour
patterns and dense pubescence on the carapace (Fig. 1), whereas conspecific females have
thin pubescence and indistinct colour patterns (Fig. 2).
Key to the species of Iberesia (males):
1. Embolus tip smooth and only slightly bent …....……………………..I. castillana (Fig. 14)
.- Embolus tip clearly bent and furnished with 2 or 3 tiny denticles …………………….. 2
2. Embolus tip sigmoid, denticles on convex side……………………….I. machadoi (Fig. 12)
.- Embolus tip bent, three minute denticles on concave side ……………….I. brauni (Fig. 13)
Iberesia machadoi new species (Figs 1-4, 6-7, 9, 12, 15)
Nemesia hispanica (misidentification): Frade & Bacelar 1931 p. 233-234. Bacelar 1932 p.20-21.
Machado 1944 p. 25-26. Buchli 1968 p. 70, Fig. 43c. Blasco 1986 p. 345,347 fig. 2D. Cardoso 2000 p.
31-36.
Etymology: the species is named in honour of António de Barros Machado who first
recognised the taxonomic importance of spinneret morphology in the Nemesiidae.
Distribution: the species occurs in many locations throughout Portugal, but it appears to be
more common in central and southern Portugal than in the north (see map 1).
Type:male holotype (leg. P. Cardoso PMC0018D, collected 23 Oct. 2000) deposited at the
MNHN, Paris Nr. AR14312.
Type locality: Algodôr, South Portugal, UTM 29SPB07, habitat Quercus ilex woodland.
Paratypes: collected at several locations in central and southern Portugal; five males from
Ribeira de Limas UTM 29SPB28; three females from Mértola UTM 29SPB16 (leg. P.
Cardoso); three females from Terras do Risco UTM 29SMC95 (leg. A. Decae). All deposited
at MNHN (AR14313 – AR14323). One male from Braciais UTM 29SPB26 (leg. P. Cardoso);
four females, one from each of the following locations: Guadalupe (Évora) UTM 29SNC86;
Castelo de Vide 29SPD36; Escusa (Marvão) 29SPD36; Porto de Espada 29SPD45 (leg. A.
Decae). All deposited at NMR (9972.4024 – 4028).
Diagnosis: differs from other Iberesia species by the morphology of the embolus tip that is
distinctly sigmoid (Fig. 12) and bears two tiny denticles on the convex side. Females differ
from those of I. brauni (the female of I. castillana is not known) by the tripartite
spermathecae (Fig. 15).

Description of male holotype 23
Measurements (mm): total length 17.6, carapace 7.0 long 6.5 wide, sternum 3.5 long 3.5 wide,
lengths legs (measured from bases of paired claws to femur – trochanter junction) I = 22.2, II
= 22.4, III = 21.7, IV = 28.4, length palp 10.0.
Patellar spine formulae: PSP [p=1-1, I=2-2, II=2-2, III=2-2, IV=2-2] RSP [p=0-0, I=1-1, II=1-
0, III=1-1, IV=1-1].
Carapace: (colour in alcohol) distinct pattern with dark head and warm yellow thorax, dark
colour extends from caput over central carapace to caudal edge (Fig. 1), caput moderately
elevated, dense silvery pubescence concentrated on the caudal flanks of caput and irregularly
located elsewhere on carapace, crest-setae in a somewhat irregular row, crest-zone indistinct,
fringe-setae fine procurved on posterior carapace, recurved from coxae III forward, dense
silvery pubescence between fringe-setae on carapace edge. Clypeus: slightly higher than
diameter ALE. Eye-group: eight eyes in two rows, compactly grouped on distinct tubercle,
posterior row longest, periocular pigmentation lighter around AME, shining black connecting
all other eyes. (AR/EL 2.1), (AR/PR 0.9), (ALE/PLE 1.1). Fovea: wide, recurved, central
longitudinal groove indistinct. Chelicerae: dark brown, not contrasting with colour of caput,
strong setae along dorsal crest distally merging with rastellum, zones of dense silvery
pubescence dorsally and laterally, ventrally row of six prolateral furrow teeth, numbers 1, 2
and 5 larger than others (Fig. 3), rastellum well developed, fang-keel serrated. Maxillae:
slightly darker than sternum, (l/w = 1.75), anterior distal process distinct, cuspules spiky
grouped in short rows of four. Sternum: yellow, setae evenly distributed, lateral setae slightly
stronger built, three distinct pairs of sigilla, all placed more than their diameter from edge.
Labium: darker than sternum, shape shorter than wide (l/w 0.5), labial furrow wide and
glabrous. Legs: ventrally yellow grading to darker brown dorsally, spines prominent on all
metatarsi, tibiae, patellae and femora, absent from other segments, trichobothria in irregular
longitudinal row on dorsal tarsi, distinct distal group and proximal row on dorsal metatarsi,
two parallel but alternating rows on dorsal tibiae, scopulae only distinct on ventral tarsi I & II:
all paired claws with two distinct combs of teeth: metatarsi and tibiae of leg I slightly shorter
than of leg II, tibia IV shorter than either metatarsus or femur IV, metatarsus IV slightly
longer than femur IV. Palp: colour, spine and trichobothria patterns as legs. Abdomen: dense
dorsal pattern of irregular dark brown and light yellow patches, ventrally uniformly yellow.
Spinnerets: PMS absent, PLS similar colour as ventral abdomen, basal segment longer than
median and distal segment together, distal segment very short and rounded (domed). Clasper
(Fig. 6): clasper-field of short distally curved spiky hairs on ventral metatarsus I, tibial spur
robust, on prominent tubercle and only slightly curved. Bulb (Fig. 12): long, slender embolus
with sigmoid, denticulate tip.
Description of female paratype:
Specimen female paratype (leg. A. Decae 02/07/99-3) deposited, MNHN, AR14314.
Measurements (mm): total length 28.7, carapace 9.4 long 8.3 wide, sternum 5.2 long 4.8 wide,
lengths legs (measured as in holotype) I = 20.8, II = 18.3, III = 18.0, IV = 26.6, length palp
15.0.
Patellar spine formulae: PSP [p=0-0; I=0-0; II=0-0; III=0-0; IV=0-0] RSP [p=0-0; I=0-0;
II=0-0; III=0-0; IV=0-0].
23
The male of this species has recently been described by Cardoso 2000 as the presumed male of N. hispanica.
To avoid confusion we have selected another specimen (than those described by Cardoso) to function as
holotype. The description given here largely overlaps with that of Cardoso 2000 and slight differences noted are
regarded to result from intraspecific morphological variation.

Carapace: pattern indistinct (Fig. 2), flanks of caput slightly darker than other parts of
carapace, crest-zone indistinct, crest-setae few and fine, fringe-setae absent, pubescence fine
silvery white and concentrated on posterior caput. Clypeus: as wide as height of eyeformation.
Caput: strongly elevated. Eyes: eight in two rows of equal length, compactly set on distinct
tubercle, periocular pigmentation distinctly broken between AME and ALE and between ALE
and posterior eyes, (AR/EL 1.9), (AR/PR 1.0), (ALE/PLE 1.2). Fovea: recurved,
somewhat hooked, deep central depression. Chelicerae: massive, dark brown contrasting with
colour of carapace (Fig. 2), dorsal setae and silvery pubescence present but less developed
than in males, furrow teeth as for male: rastellum well developed with several strong rigid
spines on protracted apical process. Fang with smooth, not serrated, keel. Maxillae: slightly
darker than sternum, (l/w 1.65) distal process indistinct, cuspules spiky, in short rows.
Sternum and Labium as in male. Legs: colour as in male, short spines only on ventral sides of
tarsi, metatarsi and tibiae I & II and on lateral tibiae and metatarsi III, super-spine on
retrolateral tibia III distinct (Fig. 4), trichobothria as in male, scopulae on tarsi and metatarsi I
& II extending to distal tibia I, combs on paired claws tarsi III & IV somewhat irregular and
reduced. Leg formula 4-1-2-3, tibia IV longer than either metatarsus IV or femur IV, femur
IV longer than metatarsus IV. Palp: colour like legs: scopula dense on ventral tarsus with
interspersed short spines: one lateral spine proximally on either side of tarsus, tibial spines
located ventrally and lightly built: claw smooth. Abdomen: uniform greyish with faint pattern
of irregular darker patches dorsally. Spinnerets (Fig. 9): PMS absent, PLS slightly darker in
colour than abdomen, otherwise as for male. Spermathecae (Fig. 15): tripartite, basal part
wide, connected by narrow central part to domed distal part, glandular tissue evenly dense on
all parts except proximally on distal part where this tissue is absent.
Variation
Males (N=7): total length 16.7-18.2; carapace 6.7-8.2 long, 5.9-6.5 wide; sternum 3.1-4.1
long, 3.3-3.5 wide; leg lengths: I = 19.0-22.2, II = 19.1-22.4, III = 17.9- 21.7, IV = 23.9-28.4;
palp length 9.2-10.0; eye-group: AR/EL 1.8-2.1, AR/PR 0.9-1.0, ALE/PLE 0.9-1.1. Patellar
spine formulae: PSPvar [p=0(1); I=2(0-1); II= 2(0-1); III=0(1); IV=1(0- 3)] RSPvar [p=0;
I=0(1-2); II=0(1-2); III=1; IV=1(0-3)]
Females (N=10): total length 19.9-29.1; carapace 6.5- 10.0 long, 5.6-8.4 wide; sternum 3.7-
5.2 long, 3.7-4.0 wide; leg lengths I = 14.8-20.8, II = 13.5-18.9, III = 13.7-18.0, IV = 20.4-
26.6; palp length 11.0-15.5; eyegroup:
AR/EL 1.8-2.3, AR/PR 1.0-1.0, ALE/PLE 1.0-1.5 Patellar spine formulae: There are no
spines, prolateral or retrolateral, in any of the 10 specimens studied.
Iberesia brauni (L. Koch 1882) comb. n. (Figs. 13,16)
Nemesia braunii L. Koch 1882: 642, pl. 20, fig. 21(mf).
Nemesia brauni Simon 1892: 113. Reimoser 1919: 6. Frade & Bacelar 1931: 226, figs. 7-8 (m). Decae
2005: 148-151, figs. 9-19 (mf).
Type material: BMNH (Hillyard, pers. com), see Decae 2005 p. 148-151.
Type locality: Palma, Majorca, Spain.
Diagnosis: differs from all other Iberesia species by three minute denticles on the concave
embolus tip.
Remarks: Koch’s 1882 original description of this species is detailed but it does not mention
the absence of the PMS. In fact, this character was never mentioned in any paper on I. brauni
until Decae 2005 who in a recent study on newly collected specimens from Majorca,
redescribed the species and gave new information on the morphology of the sexual organs and
spinnerets, and furthermore on its distribution and behaviour.

Figs. 1-11. Iberesia machadoi, dorsal habitus 1. Male 2. Female (arrow indicates crest-zone) 3. Chelicera prolateral
view showing furrow-teeth, scopula and fang serrations 4. Super-spine on retrolateral tibia III in I. machadoi 5.
Ordinary spines on retrolateral tibia III in N. hispanica 6. Clasper in male I. machadoi spur (S) and field (F) 7. single
row of spiky maxillary cuspules, I. machadoi 8. Double rows of knob-shaped cuspules, N. hispanica 9-11. spinnerets
ventral view 9. I. machadoi 10. N. hispanica 11. Brachythele spec.

Iberesia castillana (Frade & Bacelar 1931) n.comb. (Fig. 14)
Nemesia castillana Frade & Bacelar 1931:222-226, figs. 5-6 [holotype MNHN AR4341, pot 58,
Simon nr. 6045, examined].
Type locality: Avila, Spain.
Diagnosis: differs from all other Iberesia species by the absence of denticules near the tip of a
curved long slender embolus
Remarks: Frade & Bacelar 1931 described this species from an adult male specimen that they
found, unnamed, in the collection of E. Simon (MNHN Paris 6045/AR4341, Pot 58). We
recovered the type specimen and found the original description correct although somewhat
incomplete. Particularly, the observations on spinneret morphology and embolus structure, in
retrospect, were important omissions. We found that the PMS are absent in the type specimen
and therefore we place the species in Iberesia. The female of I. castillana remains unknown.
Distribution: I. castillana has only been reported from its type locality.
Figs. 12-14. Iberesia species, comparative bulb morphology (similar orientations) 12. I. machadoi 13. I.
brauni 14. I. castillana Figs. 15-16. Iberesia comparative morphology of spermathecae in dorsal view 15. I.
machadoi (tripartite) 16. I. brauni (bipartite).

Burrows: Only the burrows of I. machadoi and I. brauni have
so far been observed. Both species construct a typical ‘cork
nest’ as described and beautifully illustrated for Nemesia
caementaria by Moggridge (1873 p. 94 Plate VIII, see also
Decae 2005 p. 150, figs. 68, 75, 79).
Discussion of Iberesia relationships
In his cladistic and systematic revision of the
Mygalomorphae, Raven 1985 delimited the tribe Nemesiini
to include only the two south-western Palaearctic genera
Nemesia and Brachythele. The recurved shape of the fovea
and the conformation of the spur on tibia I of
males were proposed to be the synapomorphies that unite the
Nemesiini, although the morphology of the tibial spur differs
markedly between Nemesia (single spur) and
Brachythele(dual spurs). Here we propose an extension of the
tribe Nemesiini to include a third genus, Iberesia gen. n., that
shares the foveal shape with both Brachythele and Nemesia
and the conformation of the single tibial spur with Nemesia
alone. We base the genus level taxonomy within the
Nemesiini on the variation in spinneret morphology. In
Brachythele, the posterior median spinnerets (PMS) are well
developed, widely separated and the distal segment of the
posterior lateral spinnerets (PLS) is digitiform (Fig. 11); in
Nemesia the PMS are reduced, closely set and the
distal segment of the PLS is domed (Fig. 10); in Iberesia,
the PMS are absent and the PLS are as in Nemesia (Fig. 9).
Further, non-diagnostic macro-morphological characters
shared by all Nemesiini are: the presence of eight eyes in a
rectangular group on a distinct ocular tubercle; scopulae on
tarsi and metatarsi I & II; biserially dentate paired claws
(in both sexes but particularly well developed in males);
Map 1. Currently known distribution of
Iberesia
machadoi. Black dots are collection sites
of material
used in this study, open circles are
locations reported in
earlier literature.
cheliceral furrow with a retrolateral scopula and a prolaterally a row of strong triangular
teeth. The recognition of the tribe Nemesiini (Raven 1985 p. 47) seems to make sense, not
only from a phylogenetic point of view, but also in terms of biogeography. The distributions
of the three genera are continuous and locally overlapping. Brachythele is the eastern genus
reported from south-western Asia, Anatolia and south-eastern Europe where it overlaps with
Nemesia. The range of Nemesia extends from south-eastern Europe southward and westward
through northern Africa and southern Europe to the shores of the Atlantic Ocean. On the
Iberian Peninsula 24 the range of Nemesia fully overlaps with that of Iberesia where several
Iberesia species, described and undescribed, occur sympatrically and even syntopically with
different Nemesia species.
Discussion of Nemesia hispanica sensu Frade & Bacelar 1931
The specimens described here as I. machadoi have long been incorrectly identified as
Nemesia hispanica. The confusion originated from a discussion of N. hispanica by Frade &
24 While we were preparing the final version of this text, G. Caranhac reported unnamed Nemesiids from
Algeria missing the PMS in Simon’s collection (MNHN), indicating Iberesia is not restricted to the Iberian
Peninsula.

Bacelar 1931 based on their study of museum specimens collected in Spain and Portugal.
There is no mention of spinnerets in this discussion (in fact they almost totally ignored the
spinnerets in the entire paper except for noting that they are present on the end of the
abdomen of Nemesia fagei, p. 232) and, apparently, Frade & Bacelar did not check their
material against the holotype. They did consult the original description (Ausserer 1871)
however, on which they made some interesting remarks that might illuminate the origin
of the long-lasting misidentification of N. hispanica. Frade & Bacelar unjustly disqualified
one of Ausserer’s diagnostic characters; the double row of cuspules on the ventral proximal
maxillae (Fig. 8). They justified their disqualification of the diagnostic value of the double
row of maxillary cuspules as follows: ‘Le nombre et la disposition de ces denticules sont loin
d’être des caractères fixes permettant la séparation des espèces (p. 234)’ (the number and
arrangement of these denticles is not sufficiently constant for the distinction of species). From
recently collected material in our private collection (8 specimens) and observations on the
collection of the MNHN however, it is clear that there exists a species in central and southern
Spain of which the female normally posseses double rows of cuspules on the maxillae (Fig.
8), a character otherwise very rare in Nemesia. Moreover, our specimens share the double row
of cuspules with the possession of PMS as also present in the type of N. hispanica in the
BMNH (Hillyard, pers. comm.). Therefore, we conclude that this combination of characters
(double row of cuspules and PMS present) is diagnostic for N. hispanica L. Koch in Ausserer
1871. N. hispanica sensu Frade & Bacelar apparently is a different species. That Frade &
Bacelar (1931, pp. 233-234, fig. 19) actually discussed an Iberesia species under the name of
N. hispanica, we conclude from their description of the spine pattern on tibia III – avec
l’épine sub-apicale interne, robuste et courbée (with a sub-apical robust and curved
retrolateral spine) – in which we recognise the ‘super-spine’ mentioned here as indicative for
Iberesia. The retrolateral spines on tibia III of both sexes in N. hispanica sensu Ausserer
1871, are normal spines, not usually strongly built (Fig. 5). Given the wide distribution in
Spain and Portugal that Frade & Bacelar reported for their N. hispanica it seems likely that
these authors grouped a number of different Iberesia species under this one species name,
probably including I. machadoi and I. castillana and some, as yet, undescribed species of
which we have so far seen insufficient material. Preliminary research on specimens in
Simon’s collection reported as N. hispanica by Frade & Bacelar (1931, p. 233-234) appears to
support this hypothesis (Caranhac, pers. comm.), although further study is necessary to clarify
this. Finally, Frade and Bacelar 1931 may hint at another long standing problem concerning
N. hispanica. Roewer 1942, followed by Platnick 2005, indicated that Ausserer’s description
of Koch’s type of N. hispanica is based of a juvenile specimen and that Frade & Bacelar 1931
first described the adult female. This is very unlikely however, because Ausserer reported that
the spider was –Etwas grösser und stämmiger gebaut als N. caementaria – (somewhat larger
and sturdier built than N. caementaria). We consider that this statement
(by an expert in the field in 1871) could only have been made on the basis of an adult
specimen.
Acknowledgements
We wish to thank Gilbert Caranhac for providing a most helpful database called “What’s in
What Pot” containing information on the mygalomorph collection of the MNHN, Paris, for
answering many questions and sending photographs of morphological details in numerous
specimen, Paul Hillyard for his study of the spinneret morphology in the type material of I.
brauni and N. hispanica, Robert Raven for his valuable comments and editorial advice,
Nikolaj Sharff, Rogerio Bertani and José Guadanucci for reading and commenting on the

manuscript. Furthermore we are grateful to all the people that helped in field collections of the
new material, especially in Arrábida (Alexandre Inácio, João Bento, Marco Marujo, Nuno
Cabrita) and to the Portuguese Institute for Nature Conservation (ICN) allowing collection in
the Nature Parks of Arrábida, Douro Internacional, Serras de Aire e Candeeiros and Vale do
Guadiana and also the Nature Reserve of Paúl do Boquilobo. Nollie Hallensleben helped
correcting the typescript.

Chapter 5
Sub-generic diversity and distribution in the Mediterranean trapdoor spider
genus Nemesia Audouin 1826 (Araneae, Mygalomorphae, Nemesiidae).
Arthur E. Decae and Dries Bonte
Research 2010
Zootaxa (submitted)

Abstract
A pitfall trapped sample of 74 male nemesiid trapdoor spiders collected from locations
throughout their Mediterranean distribution range was studied for generic and sub-generic
diversity. The sample was found to contain representatives of all three established
Mediterranean nemesiid genera; Brachythele, Iberesia and Nemesia. Within Nemesia two
levels of sub-generic diversity were recognized. A first broad division revealed the existence
of two sub-generic groups that might deserve formal sub-generic or even generic status after a
more thorough revision of Nemesia. These two large groups are here denoted as Pronemesia
and Holonemesia. A second level of higher, supra-specific diversity was recognized within
both Nemesia groups to reveal the existence of several distinct evolutionary lineages.
Pronemesia was found to contain at least two major species-groups distributed in specific
locations around the western Mediterranean: the Algeria-Group in northern Algeria and the
Caementaria-Group in the western Atlantic regions and the French-Italian Riviera.
Holonemesia was found to contain three major species-groups: the Manderstjernae-Group
around the western Mediterranean, the Badia-Group in central Italy and on Corsica and the
Pannonica-Group, an essentially eastern Mediterranean group with disputable extensions into
the western Mediterranean. The diversity and distribution patterns here found seem to fit a
more general longitudinally organized pattern of diversity and distribution existing in
Eurasian Nemesiidae.
Introduction
Nemesia Audouin 1826 is one of the earliest discovered and scientifically studied genera of
Mediterranean mygalomorph spiders (e.g. Sauvages 1763, Dorthes 1794, Latreille 1799,
Moggridge 1873, 1874). The genus Nemesia is morphologically homogeneous and
taxonomically well established (Ausserer 1871, Simon 1892, 1914). It is distinguished from
its putative sister genus Brachythele Ausserer 1871 primarily by a set of macro-morphological
differences (Table 1) that pertain to features of the spinnerets, the maxillae, the rastellum and
the male tibial I spur (see Raven 1985, Decae 2005). With over fifty species described
(Platnick 2010) and numerous species undescribed (Decae 2005) Nemesia is by far the most
diverse mygalomorph spider genus in the Mediterranean region. Notwithstanding the general
morphological homogeneity, on which the classical taxonomy of the genus is based, some
evidence for sub-generic diversity within Nemesia exists. Simon (1892, 1914) was the first to
recognize this and to distinguish different species-groups within Nemesia that, in a modern
taxonomical context, would signify the existence of independently evolving lineages within
the genus. Recently Decae & Cardoso (2005) confirmed the reality of this idea by splitting off
a new genus (Iberesia Decae & Cardoso 2005) from traditional Nemesia stock on grounds of
the absence of posterior median spinnerets (PMS) in Iberesia. PMS, as functional spinning
organs, are present in all Nemesia species. Iberesia, based on the supposedly apomorphic
absence of PMS, is not only a distinct morphological and evolutionary lineage within the
Mediterranean Nemesiidae, it is also a natural geographic unit restricted in its distribution to
the Iberian Peninsula, the Balearic Islands and Morocco where it was only very recently
discovered (Opatova pers.comm). As such the distribution of Iberesia appears to indicate the
existence and location of an independent centre of nemesiid evolution in the far western
Mediterranean-Atlantic Region. This realization is important to our understanding of the
historical development of Mediterranean biodiversity. The general phenomenon of diverse,
but in their geographic distribution restricted taxa (see mygalomorph listings in World Spider
Catalog, Platnick 2010), has been proposed to be a special quality of mygalomorph spiders
making them near ideal bioindicators for studies of local biodiversity (Decae 2005). A

precondition for extracting valuable biological information from any group of organisms
however is a well developed taxonomy at all levels of evolutionary diversity. Mygalomorph
taxonomy in general is in its infancy and large areas of the world remain partly or totally
unexplored. Even in a relatively well studied region as the Mediterranean, the taxonomy of
most mygalomorph genera is insufficiently developed. Poor availability of material and low
scientific attention have been key problems. A new and extensive collection of pitfall trapped
nemesiid spiders containing all three currently recognized Mediterranean genera
(Brachythele, Iberesia, Nemesia) captured at geographical locations throughout the
distribution range (Fig. 1) however has provided a new opportunity to improve taxonomical
knowledge of this animal group. In the here presented study the genus level taxonomy of
Mediterranean Nemesiidae is reconsidered and new information on the sub-generic diversity
within the genus Nemesia is presented.
Brachythele Nemesia
posterior lateral spinnerets (PLS) long & slender short & broad
distal segment PLS digitiform dome shaped
posterior median spinnerets (PMS) wide apart close together
PMS digitiform not digitiform
maxillary cuspules In groups in rows/absent
maxillary distal process in males absent present/absent
tibia I spur in males
with twin hooks with single hook
rastellum weak/absent composed of strong spines
Table 3 diagnostic characters distinguishing the genera Brachythele Ausserer 1871 from Nemesia
Audouin 1826.
Fig.1 Sample locations of pitfall trapped nemesiid spiders referred to in this study. Blue
squares indicate capture sites for Brachythele, green circles for Nemesia and orange triangles
for Iberesia.

Materials and Methods.
From a collection of over a thousand pitfall-trapped Mediterranean nemesiid trapdoor spiders
a sample of 74 male specimens was selected on grounds of their different and wide-spread
Mediterranean origins (Fig. 1). The sample contained seven Brachythele specimens all of
supposedly different species, four Iberesia specimens of four different (described and
undescribed) species and 63 Nemesia specimens (see Appendix Table 3). Because most
Nemesia specimens in the sample represent provisionally named, unidentified and/or
undescribed species, the number of different Nemesia species here discussed can only be
approximated to about fifty. Thirty-three morphological characters (see appendix Character
List 1) of established diagnostic value in Nemesiinae (Decae 2005; Decae & Di Franco 2005;
Decae et.al 2007) were studied in all 74 specimens. All characters were scored as either
present (1) or absent (0) (see appendix Table 4). To investigate generic diversity and to
identify characters diagnostic at different (generic, sub-generic) taxonomical levels, the datamatrix
(Table 4) was statistically analyzed using multidimensional scaling (MDS).
After identifying and eliminating the character scores that are primarily diagnostic at the
genus level from the data-set (characters 7,8,12,13,20,25, appendix Character List 1, character
definitions shown in italics), relationships within the genus Nemesia were studied on the basis
of a reduced data-matrix of 64 taxa and 27 characters (see appendix Table 5). Tests were run
against four different reference groups (Brachythele denieri, Iberesia machadoi and two
hypothetical groups with all character settings either to zero or to one) using the TNTparsimonious-analysis
program (Goloboff et. al. 2008). The analysis was carried out with
program settings for TNT’s ‘traditional search option’ using implied weighting (default
setting, strength K=300000), TBR swapping algorithm active, variable random seed settings.
Parameter settings and resulting search information of these analyses are given in Table 2.
Strict consensus cladograms produced with TNT’s ‘synapomorhy ⇒ map common
characters’ option are reproduced and shown in the Appendix Figs. 18-21. Sub-generic
diversity within Nemesia was read from these cladograms to reveal the existence of two subgeneric
clades. To study further, higher level, supra-specific diversity within these two clades,
they were analyzed independently using TNT with similar program settings as described
above (Fig. 4&6). The distribution of sub-generic groups and higher level species-groups
were mapped using DIVA-GIS (Hijmans et. al. 2005). All morphological characters (see
below) were studied with the aid of a CETI-MEDO 2 stereo-microscope with the specimen
fully submerged in 70% ethanol.
SEM photographs of bulb and embolus structures were prepared by Dr. R.F. Foelix (Figs. 7,
9, 10, 11) operating a ZEISS DSM 950 scanning electron microscope at the Neue
Kantonsschule Aarau, Switzerland and Dr. P. Cardoso (Fig. 8) at the Zoological Museum of
the University of Copenhagen, Denmark (for details see Decae et.al. 2007).

Results
MDS analysis of full data-set (74 taxa, 33 characters appendix, Table 5).
MDS-analysis of the primary data-matrix of 33 characters (Appendix List 1) in the studysample
of 74 Nemesiinae males resulted in the finding of four distinct groups. The genera
Brachythele, Iberesia were found to be distinct from each other and from the genus Nemesia.
Within Nemesia two separate sub-generic groups (Figs 2 & 3) were found. Iberesia (Fig. 2)
was found to cluster around the absence of PMS (Table 4 char.12). Brachythele (Fig. 2)
clusters around a set of characters that are related to the general morphology of the PLS
(Table 4 char.7), the shape of the apical segment of the PLS (Table 4 char. 8), the general
morphology of the PMS (Table 4 char.13), the grouping of maxillary cuspules (Table 4
char.20) and the presence of twin hooks on the tibia I spur in males (Table 4 char.25).
Fig. 2 graphical representation of MDS-analysis showing the presence of four
morphological distinct groups, representing one Brachythele, one Iberesia and two
distinct Nemesia groups. MDS is based on 10000 iterations using Sörensen distances.
graphical representation of MDS-analysis showing the presence of four morphological

TNT-analysis of reduced data-set (64 taxa, 27 characters, appendix Table 6)
Four runs of cladistic analyses (against four different reference groups, see above) on a
reduced data-matrix (Appendix Table 5) using the TNT-program confirmed our MDS results
by showing a basic division of Nemesia in two distinct sub-generic clades (Appendix Figs.18-
21). Detailed information on parameter settings for TNT-analysis and results produced are
summarized in Table 2.
For convenience, and based on Simon’s 1914 terminology for subdividing Nemesia, the two
sub-generic lineages are provisionally named Pronemesia and Holonemesia (Holonemesia for
Simon’s rather impractical term Nemesia sensu stricto). Morphologically the two sub-generic
groups are primarily distinguished on characters of the PMS; characters 14, 15, 16, 17,
(Appendix Character List 1) in our data-matrix Table 4 (see also Figs. 3, 15, 16). The species
composition of Pronemesia and Holonemesia found was to be constant in all four TNT-runs
(see Appendix Table 6). The distribution of Pronemesia and Holonemesia, as far as it can be
read from the limited information in our data-set, is shown in Fig. 5. Pronemesia appears to
have its centre of distribution on the Iberian Peninsula and around the western Mediterranean.
Holonemesia seems primarily distributed in the Central Mediterranean extending both far to
the East into Greece and to the West into southeastern Spain. Although not fully resolved, all
four cladograms studied (Figs. 18-21 give strict consensus cladograms of 10 to 40 shortest
trees retained by the program) show further subdivisions of Pronemesia and Holonemesia in
several distinct and coherent clades.
Outgroup Best
Hit
Total
Rearrangements
TBR
Score
Tree
Length
Random
Seed
n.Min
Steps
n.Max
Steps
I. machadoi 1x 12173886 7.772315 93 7252 24 303
B. denieri 4x 9461300 8.07192 97 195 26 304
Hypothetical 000 7x 1288030 7.66961 93 2491 26 303
Hypothetical 111 1x 13764388 8.99643 100 18487 24 310
Table 4 Score results of traditional search analysis (TNT) using implied weighting (default
settings) on four different reference groups and variable basic parameters settings.
Pronemesia Simon 1914 (Figs. 3, 4, 5, 7, 8 & 16)
The subgenus Pronemesia is reestablished on the basis of characteristic spinneret morphology
(Figs. 3 & 16) after Simon (1914: pg. 3). Simon used spine-patters and characters of the
scopulae, embolus and burrow structure to arrive at a more restricted characterization of
Pronemesia that pertained only to the Caementaria-Group (see below). As diagnosed here
Pronemesia relates to a highly diverse subgenus of Nemesia occurring around the western
Mediterranean only.
Within Pronemesia two coherent supra-specific groups appear to be consistently present in all
four cladograms (Fig. 4, see also Appendix figs. 18-21). One apparently stable group of four
species (terminals) is present in all analysis results. The clade is here denoted as the Algeria-
Group. The key, and supposedly apomorphic, character defining the Algeria-Group
(Character List 1, char. [5]) is the presence of two serrated ridges placed on opposite sites on
the embolus (Figs. 5 & 7). The fact that no discriminating characters were found between N.
sp.bejaia and N. sp.boumerdes might indicate that these are conspecific and that the Algeria-
Group in our sample is present with three species.

Brachythele & Raveniola
Holonemesia
7
8 9 13
Nemesia
10
714 15 16
11
Pronemesia
Iberesia
Fig. 3 Cladogram based on spinneret morphology (Figs. 13-16) indicating hypothesized phylogenetic
relationships within Mediterranean Nemesiidae at the genus and subgenus level. Character numbers are
explained in Character List 1 (appendix). White squares = present character state, black squares = absent
character state.
A second clade, here referred to as the Caementaria-Group (after N. caementaria) is
represented with a maximum of nine species in our sample. It is not consistently occurring in
all four analysis results however (Figs. 18-21). In its ‘full composition’ of nine species it
shows up in two out of four analysis results (runs of Nemesia versus I. machadoi and the
Hypothetical all zero outgroup). In the run of Nemesia versus the Hypothetical all one
reference group, the Caementaria-Group appears as two separate clades (Fig. 21), one of
seven species and one of two species. In the run of Nemesia versus B. denieri it appears as
one clade of six species and three unplaced species (Fig. 18). Here we will consider the
Caementaria-Group as a distinct clade represented in our sample by nine constituent species.
It is not consistently occurring in all four analysis results however (Figs. 18-21). In its ‘full
composition’ of nine species it shows up in two out of four analysis results (runs of Nemesia
versus I. machadoi and the Hypothetical all zero outgroup). In the run of Nemesia versus the
Hypothetical all one reference group, the Caementaria-Group appears as two separate clades
(Fig. 21), one of seven species and one of two species. In the run of Nemesia versus B. denieri
it appears as one clade of six species and three unplaced species (Fig. 18). Here we will
consider the Caementaria-Group as a distinct clade represented in our sample by nine
constituent species. The key−, and supposedly apomorphic, character defining the
Caementaria-Group (Character List 1, char. [4]) is the ornamented tip of the embolus (Figs. 5
& 8) that renders these members of the Caementaria-Group immediately obvious in any
sample of Mediterranean nemesiids. In some members of the clade however, particularly in
species occurring in Portugal, the embolus modifications appear largely lost (see Figs. 25-34
in Decae et.al. 2007).
A third possible clade in Pronemesia, composed of only two species, (N.sp.tunesia.2 & N.
sp.sardinia.2) exists in three out of our four analyses. We have left this clade unnamed

ecause it rests for its distinction on character [2], a character that also defines the
Manderstjernae-Group in Holonemesia. On these grounds of apparently conflicting
diagnostic characters in spinneret morphology and those of the embolus we regard this clade
as incerta sedis until more information becomes available. All four of our TNT-runs show the
group N. latina.1 and N. latina.2 indicating the probable conspecific status of the two
specimens. N. latina appears to be a somewhat aberrant Nemesia species (also combining
characters of Pronemesia (spinneret morphology) with those of Holonemesia (character [0])
that currently is the subject of closer study (Decae & Bosmans in prep.). Finally N. simoni is
keying out separately in all our studies. N. simoni also is a distinctly aberrant Nemesia species
and was recognized as such already by Simon (1914) who placed it, with N. crassimana (a
species for which the existence needs to be confirmed), in a separate class he labeled
Haplonemesia.
Fig. 4 Cladogram of Pronemesia indicating the positions of clade defining characters of
the two supra-specific species groups here distnguished. Algeria-Group = characterised by
fine double serrated ridges on opposite sides of the embolus — Ch[5]. The Caementaria-
Group = characterized by strong modifications of the embolus tip — Ch[4]. These are
apparently lost again higher-up in the cladogram in some Portuguese species (N. ungoliant,
N. bacelarae, N. fagei). The N. latina clade, at the top of the cladogram appears to be
monospecific and might be incorrectly placed in Pronemesia on grouds of the striated
emboli — Ch[0]. The N. sardinia – N.tunesia clade based on —Ch[2] may indicate the
presence of a third supra-specific group in Pronemesia, it may also indicate a
misplacement in this group. N. simoni appears an aberrant Nemesia species as such already
recognized by Simon (1914) who placed it in a separate group he named Haplonemesia.
Scale line in bulb drawings = 1mm.

Holonemesia Simon 1914 (Figs. 3, 5, 6, 9, 10, 11 & 15)
The subgenus Holonemesia is here reestablished after Simon’s Nemesia s. str. (1914: pgs. 2 &
3) on the basis of characteristic spinneret morphology (Figs. 3 & 15). Simon used spinepatters
and characters of the scopulae, embolus and burrow structure to arrive at a more
restricted characterization that pertained only to the Manderstjernae-Group and the Badia-
Group (see below). As diagnosed here Holonemesia relates to a highly diverse subgenus of
Nemesia to which all species occurring in the Central and Eastern Mediterranean belong.
Fig. 5 Distribution of two sub-generic groups found within our Nemesia sample.
Orange triangles = Pronemesia, blue squares = Holonemesia.
Within Holonemesia three prominent and coherent supra-specific groups appear to be
consistently present in all four cladograms (Fig. 6, see also appendix Figs. 18-21). The clade,
here denoted as the Pannonica-Group (after N. pannonica from the Dalmatian coast), has a
constant number of 20 terminals. The cladogram is not fully resolved and some of the
terminals may be found conspecific on closer inspection (e.g. N. sp.friuli & N.sp.karina). The
key, and supposedly apomorphic, character defining the Pannonica-Group (Character List 1,
char. [0]) is the presence of fine longitudinal ridges at the basis of the embolus (Figs. 6 &
11).The character seems lost in a small sub-clade containing N. sp.asti and N. blanca in which
the embolus is smooth and abruptly narrowing near the tip (beaked embolus of Character List
1, char [6]). The character has also been described by Simon (1914 Figs. 37 & 38) for N.
raripila, a species not present in our sample. Interestingly the beaked embolus is also present
in combination with longitudinal ribs in N. maculatipes. Nevertheless, we are inclined to
regard the asti-blanca clade as incerta sedis on grounds of distributional arguments. A

second larger clade within Holonemesia we provisionally named the Manderstjernae-Group
(after N. manderstjernae from the French-Italian Riviera). In our sample the Manderstjernae-
Group is represented by 8 supposedly different species (although not all consensus
cladograms are fully resolved e.g. see Fig. 21). The key, and supposedly apomorphic,
character defining the Manderstjernae-Group (Character List 1, char. [2]) is the strong curved
or crescent shaped embolus that might or might not have small ornamentations such as small
denticles or other protuberances near the tip (Figs. 6 & 9).
The third clade consistently present in our analyses we denoted as the Badia-Group (after N.
badia originally described from Corsica and later incorrectly placed in synonymy with N.
meridionalis by Pérez & Zárate, 1947). The Badia-Group is present with 16 terminals in our
sample several of which will be found to be conspecific on closer study (see Figs. 18-21).
The key, and supposedly apomorphic, character defining the Badia-Group (Character List 1,
char [1]) is the strongly elongated, smooth and flexible embolus (Figs. 6 & 10). Only one
species keying out consistently in the Holonemesia clade, N. sp.blida from Algeria cannot be
fitted in any of the here recognized species groups and remains unplaced.

Fig. 6 Cladogram of Holonemesia indicating the positions of clade defining characters of
the 3 supra-specific species groups here distnguished. Pannonica-Group is characterised
by fine longitudinal ribs or striae on the proximal embolus — Ch [0]. The Manderstjernae-
Group is characterized by strong rigid curved emboli with or without small modifications
at the tip — Ch [2]. The Badia-Group is characterised by long smoothly flexible and
sigmoid emboli — Ch [1]. N.sp.blida from Algeria is the only species that appears difficult
to place in any of the recognized supra-specific groups. Scale line in bulb drawings =
1mm.
Fig. 7-8 Close-up SEM images of
Pronemesia bulbs. 7 = Algeria-
Group (N. sp.bouira), note
serrations along embolus. 8 =
Ceamentaria-Group (N. athiasi),
note ornamentation of the embolus
tip.

Fig. 9-11 Close-up SEM images
of Holonemesia bulbs. 9 =
Manderstjernae-Group (N.
manderstjernae), arrow
indicates small distal
protuberance. 10 = Badia-Group
(N. badia), note smooth, slender
elongated embolus. 11 =
Pannonica-Group (N.
pannonica), note longitudinal
striae (arrow indication) on
embolus base.
Geography
The geographical distribution of all groups discussed and distinguished above seems to follow
a distinct pattern (Fig. 12). The Algeria-Group and the Badia-Group have narrow
distributions in Algeria and Italy/Corsica respectively. The Caementaria-Group and the
Manderstjernae-Group seem to have largely parallel distributions although the Caemenaria-
Group has strong Atlantic affiliations, while the Manderstjernae-Group is strictly
Mediterranean. The Pannonica-Group is also strictly Mediterranean, and an inhabitant of the
Adriatic, the southern Tyrrhenian region and the Ionian region. The Pannonica-Group is also
spread further southeast into the Peloponnesus, the southern Greek Island arc and even
reaching Cyprus (Decae personal observation, no material in our sample) and the Nile Delta,
type locality for N. cellicola (Audouin 1826).
The Nemesia fauna of the western Mediterranean is conspicuously more complex and more
diverse than the eastern Mediterranean Nemesia fauna. All species classified as incerta sedis
are western Mediterranean taxa probably indicating the insufficiency of our data-set for
resolving all supra-specific diversity in our sample.
Key to Mediterranean Nemesiidae (males generic and sub-generic level)
1. PLS long and slender with digitiform distal segment … Brachythele (Fig. 13)
PLS short and thick, distal segment dome-shaped…….. 2
2. PMS absent ……………………………………………. Iberesia (Fig. 14)
PMS present …………………………………………… 3
3. PMS knob-shaped with numerous apical spigots ……… Holonemesia (4) (Fig. 15)
PMS reduced, cone-shaped with few apical spigots …… Pronemesia (5) (Fig. 16)
4. Embolus strong, curved with or without small ornaments Manderstjernae-Group
Embolus elongated, flexible and sigmoid………………. Badia-Group
Embolus furnished with longitudinal ribs or striae ……. Pannonica-Group
5. Embolus with opposite serrated ridges ………………… Algeria-Group
Embolus distinctly ornamented at the tip ……………… Ceamentaria-Group (part)
Embolus very short and curved ………………………... Caementaria-Group (part)
Embolus with distinct fish-hook tooth ………………… Caementaria-Group (part)

Fig. 12 Geographic distribution of the clades found within Pronemesia and Holonemesia. Algeria-Group
(yellow triangles) is restricted to northern-central Algeria. Caementaria-Group (orange squares) is
distributed in the Atlantic Region and around the western Mediterranean. Mandersjernae-Group (red
triangles) is strictly distributed around the western Mediterranean. Badia-Group (blue circles) is restricted to
northern and central Italy and Corsica. Pannonica-Group (green squares) is distributed around the Adriatic
Sea with extensions into the southern Greek island arc, Sicily and Sardinia. Question marks indicate
locations of species here classified as incerta sedis with respect to sub-generic and supra-specific
Figs. 13-16 Generic and sub-generic diagnostic characters in spinneret morphology
of Mediterranean nemesiid spiders (ventral view). Note differences in spigot patterns.
13 = Brachythele, 14 = Iberesia, 15 = Holonemesia, 16 = Pronemesia. Scale line =
5mm.

Conclusions
The family Nemesiidae Raven 1985 is represented by three genera within the Mediterranean
Region. The regional distribution of these genera shows a distinct longitudinal pattern with
Brachythele in the northeast and Iberesia in the far west. Nemesia inhabits the central
Mediterranean overlapping in distribution with Brachythele in the east and with Iberesia in
the west (Fig. 1). Within Nemesia, on the sub-generic level of taxonomy, a further
longitudinally organized distribution pattern exists (Fig. 5) with Pronemesia distributed in the
Atlantic West and the western Mediterranean and Holonemesia extending its range from
southern and eastern Spain to Greece and beyond. Along the shores of the western
Mediterranean Pronemesia and Holonemesia coexist with each other, and in Spain and on the
Balearics also with Iberesia.This co-occurrence often exists in the form of syntopic
populations with representatives of the different taxa inhabiting trapdoor nests only
centimeters apart (personal observation). On a higher, supra-specific, taxonomical level the
distribution pattern of Nemesia lineages is again longitudinally organized (Fig.12). The
Caemenatria-Group occupies the Atlantic West and the western shores of the Mediterranean.
The Manderstjernae-Group shares the western Mediterranean distribution range with the
Caemenataria-Group, but is absent from the Atlantic regions. The Algeria-Group is only
found on the southcoast of the western Mediterranean. The Badia-Group occurs further to the
northeast in central and northern Italy and on Corsica. The Pannonica-Group finally has
extensions into the western Mediterranean on Sardinia, other Thyrrenian islands (personal
observation) and Sicily, but has its centre of diversity in the Eastern Mediterranean and on the
Balkan Peninsula. At all levels of taxonomy, the Nemesia fauna of the western Mediterranean
is much more diverse than that of the eastern Mediterranean. We found two nemesiid genera
and members of all recognized sub-generic and supra-specific groups in the western
Mediterranean. This is in sharp contrast with the finding of two genera and only one subgeneric
group in the eastern Mediterranean. It has been suggested (Decae in press) that the
remarkable high species diversity in Mediterranean Nemesiidae might be explained from
periodic contraction and segregation of ancestral relict populations in response to the waxing
and waning of Pleistocene glacial periods. This maybe true at the species level, but the here
found generic, sub-generic and supra-specific diversity seems to indicate the existance of
older, probably geologic and/or palaeogeographic, processes shaping nemesiid distribution.
Discussion
In his cladistic and systematic revision of the mygalomorph spiders Raven (1985) established
the family Nemesiidae in which he recognized six subfamilies. The family Nemesiidae has a
virtually cosmopolitan distribution with four subfamilies in the southern hemisphere, one
cross equatorial subfamily (Bemmerinae) and one exclusively northern subfamily
(Nemesiinae). Within the northern hemisphere the Nemesiinae occur in the warmer temperate
regions between approximately 25 0 N and 50 0 N. In this relatively narrow latitudinal zone the
Nemesiinae are widely spread throughout Eurasia from the Japanese Ryukyu Islands and
Taiwan through central Asia and the Middle East to the westernmost parts of the
Mediterranean where they occur also in the Maghreb Countries (Fig.17). Nemesiinae are
furthermore known from the western USA (genus Calisoga Chamberlin 1937) and from
Mexico (genus Mexentypesa Raven 1987), but they seem to be absent from eastern North
America where, on grounds of fossil finds in Dominican amber (Penney 1999; Selden &
Penney 2010), they might be expected to occur (or have occurred).
At the time of Raven’s 1985 revision the Nemesiinae contained only three genera: Nemesia
Audouin 1826 and Brachythele Ausserer 1871 in the Mediterranean and Calisoga in
California. Since that time, knowledge of the Eurasian Nemesiinae in particular has rapidly
increased with the descriptions of three new genera (Raveniola Zonstein 1987, Sinopesa

Raven & Schwendinger 1995, Iberesia Decae & Cardoso 2005) and 23 new species (Hu & Li
1987; Zonstein 1987, 2000, 2007, 2009; Dunin 1988; Decae 1995, 2005; Raven &
Schwendinger 1995; Zhu, Zhang & Zhang 1999; Shimojana & Haupt 2000; Song, Zhu &
Chen 2001; Xu & Yin 2002; Decae & Cardoso 2005; Lazarov 2005; Decae, Cardoso &
Selden 2007; Kovblyuk & Ponomarev 2008). All this taxonomic activity has produced
important new information on the patterns of diversity and distribution of the Nemesiinae.
The picture emerging is that, at the genus level, the subfamily is primarily distributed in a
longitudinal geographic diversity pattern with the genus Sinopesa in the Far East 25 , Raveniola
in eastern and Central Asia, Brachythele in Asia Minor and Eastern Europe, Nemesia in the
Mediterranean and Iberesia in the Iberian and Atlantic zones of the distribution range
(Fig.17). Future discoveries and descriptions have to reveal how accurate and complete this
longitudinal diversity pattern is, and if it is also visible at higher sub-generic and supraspecific
taxonomic levels. Recently Zonstein (1987, 2009) and Decae & Di Franco (2005),
following Simon (1892, 1914), have reported the existence of distinct species groups in
Raveniola and Nemesia. Here the existence of a longitudinally organized diversity pattern at
the sub-generic and supra-specific in Nemesia is confirmed.
Fig. 17 The known distribution of Nemesiinae genera based on published distribution
records. Lilac circles = Brachythele, blue squares = Iberesia, orange triangles = Nemesia,
green squares = Raveniola, red circles = Sinopesa.
Acknowledgements
25 The one Nemesia record in Eastern China (Fig. 17) is Nemesia sinensis Pocock 1901. The genetic identity of
this species needs verification.

We thank Rainer Foelix and Pedro Cardoso for providing the SEM-photos of Nemesia bulbs
and Rudy Jocqué and Jan Bosselaers for their constructive critical comments on earlier
versions of the document. This study would not have been possible without the cooperation of
Pedro Cardoso, Paolo Pantini, Marco Isaia, Alberto Chiarle, Maria Chatzaki, Rob Bosmans,
Siegfied Huber, Johan van Keer, Francesca di Franco, Fulvio Gasparo, Peter Jäger, Wolfgang
Nentwig and Jiri Kral who all provided specimens, sometimes in large collections, for this
study.
Appendix
Table 3 List of specimens with indications of genera, species and locations of origin. The
prefix ‘sp.’ preceding a species name indicates a provisional unidentified and/or undescribed
species.
Genus Species Country Region Genus Species Country Region
Brachythele sp.chios Greece Aegean Nemesia sp.sardinia Italy Sardinia
Brachythele sp.kos Greece Kos Nemesia sp.corsica.2 France Corsica
Brachythele sp.crete Greece Crete Nemesia spec-sardinia Italy Sardinia
Brachythele icterica Greece Halkidiki Nemesia maculatipes Italy Sardinia
Brachythele denieri Greece Halkidiki Nemesia sp.tunesia.2 Tunesia Kesra
Brachythele sp.croatia Croatia Karinia Nemesia sp.corsica.1 France Corsica
Brachythele media Croatia Istria Nemesia sp.tunesia.1 Tunesia Sidi Bouzid
Iberesia machadoi Portugal Algarve Nemesia sp.emilia.sig.1 Italy Emilia.Rom.
Iberesia sp.tarifa Spain Andalucia Nemesia sp.toscana Italy Toscana
Iberesia castiliana Spain Madrid Nemesia sp.emilia.elo Italy Emilia.Rom.
Iberesia sp.murcia Spain Murcia Nemesia latina.2 Italy Lazio
Nemesia athiasi Portugal Tras os Montes Nemesia sp.emilia.sig.2 Italy Emilia.Rom.
Nemesia bacelarae Portugal Tras os Montes Nemesia sp.emilia.sig.3 Italy Emilia.Rom.
Nemesia sp.tolox Spain Malaga Nemesia sp.emilia.elo Italy Emilia.Rom.
Nemesia sp.mazarron Spain Murcia Nemesia sanzoi Italy Sicily
Nemesia sp.blanca Spain Murcia Nemesia sp.umbria Italy Umbria
Nemesia simoni France Pyr. Atlantique Nemesia latina.1 Italy Lazio
Nemesia sp.valenciae Spain Murcia Nemesia spec.gran.sasso Italy Abruzzo
Nemesia sp.saida Algeria Saida Nemesia sp.marche.str Italy Marche
Nemesia sp.barcelona Spain Catalonia Nemesia sp.capestrano Italy Abruzzo
Nemesia sp.blida Algeria Blida Nemesia sp.marche.elo Italy Marche
Nemesia sp.boumerdes Algeria Boumerdes Nemesia sp.abruzzo.elo Italy Abruzzo
Nemesia sp.bouira Algeria Bouira Nemesia sp.friuli Italy Friuli
Nemesia caementaria France Herault Nemesia sp.abruzzo.str Italy Abruzzo
Nemesia spec.tizi.ouzou.b Algeria Tizi Ouzou Nemesia sp.molise Italy Molise
Nemesia spec.tizi.ouzou.a Algeria Tizi Ouzou Nemesia sp.casacalendra Italy Umbria
Nemesia sp.bejaia Algeria Bejaia Nemesia sp.acquavene Italy Campania
Nemesia carminans France Vaucluse Nemesia sp.puglia Italy Puglia
Nemesia sp.setif Algeria Setif Nemesia sp.karina Croatia Karinia
Nemesia manderstjernae France Alpes H. Prov Nemesia sp.minervino Italy Puglia
Nemesia sp.asti Italy Piemonte Nemesia sp.bari Italy Puglia
Nemesia sp.savona Italy Liguria Nemesia pannonica Bosnia Hercegovina
Nemesia sp.finale Italy Liguria Nemesia sp.pindos Greece Pindos
Nemesia badia France S-Corsica Nemesia sp.rethimnon Greece Crete
Nemesia sp.liguria Italy Liguria Nemesia sp.iraklio Greece Crete
Nemesia sp.desulo Italy Sardinia Nemesia sp.karpathos Greece Karpathos

Character List 1.
All characters are used in MDS genus level analysis. Characters in italics are not used in
TNT-cladistic analyses of sub-generic level taxonomy. Character numbers given in
square brackets are used in TNT-analyses only.
Character 0. Embolus of palpal bulb with/without longitudinal ribs (striae). With striae = 1;
without striae =0. Striae on the embolus, although not uncommon in Nemesiidae (e.g. see
figures of Nemesiidae in Raven 1985 and Goloboff 1995), have not been observed in
Brachythele or Iberesia. They are present in several, but not all, Nemesia species and are
therefore regarded as prospectively diagnostic at the sub-generic level in Nemesia.
Character 1 [1]. Embolus clearly elongated generally curved or sigmoid in shape gradually
narrowing from the base to the tip without any trace of striae or other ornaments. Smooth
elongated embolus present = 1; smooth elongated embolus absent = 0.
Character 2 [2]. Embolus not elongated otherwise as described under character 1. Shorter
smooth embolus present = 1; shorter smooth embolus absent = 0.
Character 3 [3]. Embolus without a trace of striae but ornamented with small denticles in the
distal half usually close to the embolus tip. Presence of embolus denticles = 1; absence of
denticles =0.
Character 4 [4]. Embolus not gradually narrowing towards the tip, instead the embolus tip is
strongly modified and ornamented with larger teeth, combs, ridges, bifurcations etc.
Presence of strongly modified embolus = 1; absence of strongly modified embolus = 0.
Character 5 [5]. Embolus with two serrated ridges placed on opposite sides. Presence of
opposed embolus serrations = 1; absence of such serrations = 0.
Character 6 [6]. Embolus beaked or abruptly narrowing at the tip. The proximal part of the
narrowing usually bears a little tooth. Presence of an abruptly narrowing embolus tip =1;
absence of beaked embolus tip = 0.
Character 7. Posterior lateral spinnerets (PLS) long and slender, all three segments (seen in
ventral view) of (sub)equal length = 1; thick short PLS with segments of distinctly different
lengths = 0.
Character 8. Distal segment of the PLS digitiform. This appears to be the plesiomorphic state
in Nemesiinae present in Brachythele, Raveniola and Sinopesa. Distal segment of PLS
digitiform =1; distal segment PLS not digitiform = 0.
Character 9 [7]. Distal segment of the PLS short and domed. Distal segment PLS domeshaped
= 1; distal segment PLS not dome-shaped = 0.
Character 10 [8]. Spigots on proximal segment of the PLS spread over the whole ventral
surface of the segment =1; spigots restricted to the distal half of this segment =0.
Character 11 [9]. Posterior median spinnerets (PMS) present = 1; PMS absent = 0. The
supposed phylogenetic importance of this character is given extra weight by also scoring
the reverse condition in character 12.
Character 12. The absence of PMS is considered the key diagnostic character at the genus
level for Iberesia (Decae & Cardoso 2005). PMS absent = 1; PMS present = 0.
Character 13. PMS digitiform = 1; PMS not digitiform = 0.
Character 14 [10]. PMS somewhat inflated and knob-shaped =1; PMS not inflated or knobshaped
= 0.
Character 15 [11]. PMS small conical reduced in development = 1; PMS not reduced or
conical = 0.
Character 16 [12]. PMS with many spigots = 1; PMS with few spigots = 0.
Character 17 [13]. Spigots on PMS indistinct = 1; Spigots on PMS distinct = 0.
Character 18 [14]. Keel on the ventral prolateral fang with serrations = 1; keel serrations
absent is = 0.
Character 19[15]. Maxillary cuspules present =1; cuspules absent = 0.

Character 20. Many maxillary cuspules in distinct groups = 1; cuspules not grouped = 0.
Character 21 [16]. Rastellum present = 1; rastellum absent = 0.
Character 22 [17]. Femur IV > Tibia IV = 1; Femur IV ≤ Tibia IV = 0.
Character 23 [18]. Tibia IV > Metatarsus IV = 1; Tibia IV ≤ Metatarsus IV = 0.
Character 24 19]. Maxillary process present = 1; maxillary process absent = 0.
Character 25. Palp Tibia with rake present = 1; tibial rake absent = 0.
Character 26 [20]. Scopula on ventral metatarsus I with rasp of concentrated knobby
structures. Rasp present =1; rasp absent =0.
Character 27[21] . Scopula on ventral metatarsus with concentrated spiky structures (instead
of knobby hairs). Spiky hairs present =1; spiky hairs absent =0.
Character 28 [22]. Distal spur on Tibia I with twin hooks present = 1; twin-hooks absent = 0.
Character 29 [23]. Prolateral patella III with 3 or more spines = 1; less spines = 0.
Character 30 [24]. Prolateral patella III with 2 or more spines = 1; less spines = 0.
Character 31 [25]. Vague maculae present on outer leg segments and/or proximal segment
PLS = 1; no traces of maculae = 0.
Character 32 [26]. Distinct maculae on outer leg-segments and proximal segment of PLS = 1;
maculae vague or absent = 0.
Table 4 Full data-matrix used for multidimensional scaling (MDS) analysis.
74 taxa, 33 character scores per taxon.
No (provisional) species embolus spinnerets other characters
00 Brachythele sp.chios 0100000 11011010010 111010000111100
01 Brachythele sp.kos 0100000 11011010010 111010000111100
02 Brachythele sp.crete 0100000 11011010010 111010000011100
03 Brachythele icterica 0100000 11011010010 110110000011100
04 Brachythele denieri 0000100 11011010010 111011000011100
05 Brachythele sp. croatia 0100000 11011010010 111010000011100
06 Brachythele media 0100000 11011010010 111000000010100
07 Nemesia ungoliant 0000000 00101000101 110100111000100
08 Iberesia machadoi 0001000 00100100000 010010110100100
09 Nemesia fagei 0000000 00101000101 100101111000000
10 Nemesia athiasi 0000100 00101000101 100100111000110
11 Nemesia bacelarae 0001000 00101000101 110101111000100
12 Iberesia sp.tarifa 0001000 00100100000 000100110100000
13 Nemesia sp.tolox 0010000 00111001010 010100110000100
14 Iberesia castiliana 0001000 00100100000 010100110100000
15 Iberesia sp.murcia 0001000 00100100000 010100110100100
16 Nemesia sp.mazaron 0010000 00111001010 010101110101100
17 Nemesia sp.blanca 0000001 00111001010 010101111000000
18 Nemesia simoni 0001000 00101000101 000100100001100
19 Nemesia valenciae 0000100 00101000101 000101110000100
20 Nemesia sp.saida 0000100 00101000101 000100110000000
21 Nemesia sp.barcelona 0010000 00101001010 000101110001100
22 Nemesia sp.blida 0000000 00101001010 000101100000100
23 Nemesia sp.boumerdes 0000010 00101000101 000100110000100
24 Nemesia sp.bouira 0000010 00101000101 000110110001100
25 Nemesia caementaria 0000100 00101000101 110110101000100
26 Nemesia sp.tizi.ouzou.b 0010000 00101001010 000100110000100
27 Nemesia sp.tizi.ouzou.a 0000010 00101000101 000110110000100
28 Nemesia sp.bejaia 0000010 00101000101 000100110000100

29 Nemesia carminans 0000100 00101000101 110100101000100
30 Nemesia sp.setif 0010000 00111001010 000101110001100
31 Nemesia manderstjernae 0010000 00111001010 010111110001100
32 Nemesia sp.asti 0000001 00111001010 000101110100100
33 Nemesia sp.savona 0000100 00101000101 110100100100000
34 Nemesia sp.finale 0010000 00111001010 010101110001110
35 Nemesia badia 0100000 00111001010 000100110000100
36 Nemesia sp.liguria 0100000 00101001010 000101110100100
37 Nemesia sp.desulo 1000000 00111001010 110101110101110
38 Nemesia sp.sardinia 1000000 00111001010 110111110101100
39 Nemesia sp.corsica.2 0100000 00111001010 000110110100100
40 Nemesia sp.sardinia.2 0010000 00101000101 000100110100000
41 Nemesia maculatipes 1000001 00111001010 000111110101110
42 Nemesia sp.tunesia.2 0010000 00111000101 000101110000100
43 Nemesi sp.corsica.1 0100000 00111001010 000101110100100
44 Nemesia sp.tunesia.1 0010000 00111001010 010101110000100
45 Nemesia sp.emilia.sig 0100000 00111001010 000101110000100
46 Nemesia sp.toscana 0100000 00111001010 000100110100100
47 Nemesia sp.emilia.elo 0100000 00111001010 000001110100101
48 Nemesia sp.latina.2 1000000 00101000101 010101110100110
49 Nemesia sp.emilia.sig.2 0100000 00111001010 000100110001100
50 Nemesia sp.emilia.sig.3 0100000 00111001010 000101110100100
51 Nemesia sp.emilia.elo.1 0100000 00111001010 000111110000100
52 Nemesia sanzoi 1000000 00111001010 000101110101110
53 Nemesia sp.umbria 0100000 00111001010 000101110100101
54 Nemesia sp.latina.1 1000000 00101000101 010101110000110
55 Nemesia sp.gran.sasso 0100000 00111001010 000101110000100
56 Nemesia sp.marche.str 1000000 00111001010 000111110100100
57 Nemesia sp.capistrano 1000000 00111001010 000100110101100
58 Nemesia sp.marche.elo 0100000 00111001010 000101110100000
59 Nemesia sp.abruzzo.elo 0100000 00111001010 000111110000110
60 Nemesia sp.friuli 1000000 00111001010 110101110100100
61 Nemesia sp.abruzzo.str 1000000 00101001010 010101110101100
62 Nemesia sp.molise 0100000 00111001010 010100110100100
63 Nemesia sp.casacalendra 0100000 00111001010 010100110100110
64 Nemesia sp.acquavene 1001000 00111001010 110110110101100
65 Nemesia sp.puglia 1000000 00111001010 110100110000101
66 Nemesia sp.karina 1000000 00111001010 110101110100100
67 Nemesia sp.minervino 1000000 00111001010 110100110101110
68 Nemesia sp.bari 1000000 00111001010 110110110101110
69 Nemesia pannonica 1000000 00111001010 110101110000101
70 Nemesia sp.pindos 1000000 00111001010 100100110100110
71 Nemesia sp.rethimnon 1000000 00111001010 010101110101100
72 Nemesia sp.iraklio 1000000 00111001010 000100110001100
73 Nemesia sp.karpathos 1000000 00111001010 010101110100100

Table 5 Reduced data-matrix used for TNT cladistic analysis. 64 taxa, 27 character scores per
taxon.
No (provisional) Nemesia species embolus spinnerets other characters
00 Outgroup - - - - - - - - - - - - - - - - - - - - - - -
01 Nemesia ungoliant 0000000 1010101 1110011100100
02 Nemesia fagei 0000000 1010101 1010111100000
03 Nemesia athiasi 0000100 1010101 1010011100110
04 Nemesia bacelarae 0001000 1010101 1110111100100
05 Nemesia sp.tolox 0010000 1111010 0110011000100
06 Nemesia sp.mazaron 0010000 1111010 0110111011100
07 Nemesia sp.blanca 0000001 1111010 0110111100000
08 Nemesia simoni 0001000 1010101 0010010001100
09 Nemesia valenciae 0000100 1010101 0010111000100
10 Nemesia sp.saida 0000100 1010101 0010011000000
11 Nemesia sp.barcelona 0010000 1011010 0010111001100
12 Nemesia sp.blida 0000000 1011010 0010110000100
13 Nemesia sp.boumerdes 0000010 1010101 0010011000100
14 Nemesia sp.bouira 0000010 1010101 0011011001100
15 Nemesia caementaria 0000100 1010101 1111010100100
16 Nemesia sp.tizi.ouzou.b 0010000 1011010 0010011000100
17 Nemesia sp.tizi.ouzou.a 0000010 1010101 0011011000100
18 Nemesia sp.bejaia 0000010 1010101 0010011000100
19 Nemesia carminans 0000100 1010101 1110010100100
20 Nemesia sp.setif 0010000 1111010 0010111001100
21 Nemesia manderstjernae 0010000 1111010 0111111001100
22 Nemesia sp.asti 0000001 1111010 0010111010100
23 Nemesia sp.savona 0000100 1010101 1110010010000
24 Nemesia sp.finale 0010000 1111010 0110111001110
25 Nemesia badia 0100000 1111010 0010011000100
26 Nemesia sp.liguria 0100000 1011010 0010111010100
27 Nemesia sp.desulo 1000000 1111010 1110111011110
28 Nemesia sp.sardinia 1000000 1111010 1111111011100
29 Nemesia sp.corsica.2 0100000 1111010 0011011010100
30 Nemesia sp.sardinia.2 0010000 1010101 0010011010000
31 Nemesia maculatipes 1000001 1111010 0011111011110
32 Nemesia sp.tunesia.2 0010000 1110101 0010111000100
33 Nemesi sp.corsica.1 0100000 1111010 0010111010100
34 Nemesia sp.tunesia.1 0010000 1111010 0110111000100
35 Nemesia sp.emilia.sig 0100000 1111010 0010111000100
36 Nemesia sp.toscana 0100000 1111010 0010011010100
37 Nemesia sp.emilia.elo 0100000 1111010 0000111010101
38 Nemesia sp.latina.2 1000000 1010101 0110111010110
39 Nemesia sp.emilia.sig.2 0100000 1111010 0010011001100
40 Nemesia sp.emilia.sig.3 0100000 1111010 0010111010100
41 Nemesia sp.emilia.elo.1 0100000 1111010 0011111000100
42 Nemesia sanzoi 1000000 1111010 0010111011110
43 Nemesia sp.umbria 0100000 1111010 0010111010101
44 Nemesia sp.latina.1 1000000 1010101 0110111000110
45 Nemesia sp.gran.sasso 0100000 1111010 0010111000100
46 Nemesia sp.marche.str 1000000 1111010 0011111010100

47 Nemesia sp.capistrano 1000000 1111010 0010011011100
48 Nemesia sp.marche.elo 0100000 1111010 0010111010000
49 Nemesia sp.abruzzo.elo 0100000 1111010 0011111000110
50 Nemesia sp.friuli 1000000 1111010 1110111010100
51 Nemesia sp.abruzzo.str 1000000 1011010 0110111011100
52 Nemesia sp.molise 0100000 1111010 0110011010100
53 Nemesia sp.casacalendra 0100000 1111010 0110011010110
54 Nemesia sp.acquavene 1001000 1111010 1111011011100
55 Nemesia sp.puglia 1000000 1111010 1110011000101
56 Nemesia sp.karina 1000000 1111010 1110111010100
57 Nemesia sp.minervino 1000000 1111010 1110011011110
58 Nemesia sp.bari 1000000 1111010 1111011011110
59 Nemesia pannonica 1000000 1111010 1110111000101
60 Nemesia sp.pindos 1000000 1111010 1010011010110
61 Nemesia sp.rethimnon 1000000 1111010 0110111011100
62 Nemesia sp.iraklio 1000000 1111010 0010011001100
63 Nemesia sp.karpathos 1000000 1111010 0110111010100
Table 6 Species composition of Pronemesia (top of the table) and Holonemesia (bottom of
the table), based on species and provisional species) present in our sample.
Pronemesia N. ungoliant 39.6 0 N 8.8 0 W Pronemesia N.caementaria 43.6 0 N 3.9 0 E
Pronemesia N.fagei 37.2 0 N 7.5 0 W Pronemesia N.sp.tizi.ouzou.a 36.5 0 N 4.1 0 E
Pronemesia N.athiasi 41.4 0 N 7.4 0 W Pronemesia N.sp.bejaia 36.6 0 N 5.3 0 E
Pronemesia N.bacelarae 41.2 0 N 6.7 0 W Pronemesia N carminans 43.8 0 N 5.5 0 E
Pronemesia N.simoni 43.0 0 N 1.0 0 W Pronemesia N.sp.savona 44.3 0 N 8.0 0 E
Pronemesia N. valenciae 39,0 0 N 0.1 0 W Pronemesia N.sp.sardinia.2 39,6 0 N 9.2 0 E
Pronemesia N.sp.saida 34.6 0 N 0.2 0 E Pronemesia N.sp.tunesia.2 35.8 0 N 9.4 0 E
Pronemesia N.sp.boumerdes 36.8 0 N 3.6 0 E Pronemesia N.latina.2 42.4 0 N 12.1 0 E
Pronemesia N.sp.bouira 36.1 0 N 3.7 0 E Pronemesia N.latina.1 41.5 0 N 12.8 0 E
Holonemesia N. sp.tolox 36,7 0 N 4,9 0 W Holonemesia N.sp.emilia.sig.3 43.9 0 N 12.4 0 E
Holonemesia N.mazaron 37.6 0 N 1.3 0 W Holonemesia N.emilia.elo.2 43.9 0 N 12.6 0 E
Holonemesia N.sp.blanca 38.0 0 N 1.0 0 W Holonemesia N.sanzoi 38.1 0 N 12.8 0 E
Holonemesia N.sp.barcelona 41.4 0 N 2.2 0 E Holonemesia N.sp.umbria 43.1 0 N 12.8 0 E
Holonemesia N.sp.blida 36,6 0 N 3.2 0 E Holonemesia N.sp.gran.sasso 42.6 0 N 13.2 0 E
Holonemesia N.sp.tizi.ouzou/b 36,3 0 N 4.0 0 E Holonemesia N.sp.marche.str 43.3 0 N 13.4 0 E
Holonemesia N.sp.setif 36,5 0 N 5.5 0 E Holonemesia N.sp.capistrano 42.5 0 N 13.5 0 E
Holonemesia N.manderstjernae 44.0 0 N 6.2 0 E Holonemesia N.sp.march.elo 42.9 0 N 13.6 0 E
Holonemesia N. sp.tolox 36,7 0 N 4,9 0 W Holonemesia N.sp.abruzzo.elo 42.5 0 N 13.7 0 E
Holonemesia N.sp.finale 44.2 0 N 8.6 0 E Holonemesia N.sp.friuli 45.6 0 N 13.8 0 E
Holonemesia N.badia 41.9 0 N 8.7 0 E Holonemesia N.sp.abruzzo.str 42.3 0 N 13.8 0 E
Holonemesia N.sp.liguria 44.4 0 N 8.9 0 E Holonemesia N.sp.molise 41.7 0 N 14.9 0 E
Holonemesia N.sp.desulo 40.2 0 N 9.0 0 E Holonemesia N.sp.casacalendra 41.5 0 N 15.0 0 E
Holonemesia N.sardinia.1 40.5 0 N 9.0 0 E Holonemesia N.sp.acquavene 40.0 0 N 15.5 0 E
Holonemesia N.corsica.2 42.6 0 N 9.0 0 E Holonemesia N.sp.puglia 41.9 0 N 15.6 0 E

Holonemesia N.maculatipes 40.3 0 N 9.3 0 E Holonemesia N.sp.karina 44.2 0 N 15.8 0 E
Holonemesia N.sp.corsica.1 42.7 0 N 9.5 0 E Holonemesia N.sp.minervino 41.0 0 N 16.7 0 E
Holonemesia N.sp. tunesia.1 35.0 0 N 9.5 0 E Holonemesia N.sp.bari 41.1 0 N 16.8 0 E
Holonemesia N.sp.emilia.sig.1 44.0 0 N 11.5 0 E Holonemesia N.pannonica 42.9 0 N 17.6 0 E
Holonemesia N.sp.toscana 43.5 0 N 11.9 0 E Holonemesia N.sp.pindos 39.4 0 N 20.9 0 E
Holonemesia N.sp.emilia.elo.1 44.1 0 N 12.1 0 E Holonemesia N.sp.rethimnon 35.4 0 N 24.5 0 E
Holonemesia N.sp.emilia.sig.2 44.0 0 N 12.2 0 E Holonemesia N.sp.iraklio 35.3 0 N 25.1 0 E
Holonemesia N.sp.karpathos 35.6 0 N 27.2 0 E
Fig. 18 Strict consensus cladogram based on 30 retained shortest trees found by
TNT’s traditional search option with settings as given in Table 2, analysis of
Nemesia versus Brachythele denieri

Fig. 19 Strict consensus cladogram based on 10 retained shortest trees found by
TNT’s traditional search option with settings as given in Table 2, analysis of
Nemesia versus Iberesia machadoi

Fig. 20 Strict consensus cladogram based on 40 retained shortest trees found by
TNT’s traditional search option with settings as given in Table 2, analysis of
Nemesia versus Hypothetical-Group 000 (all zero)

Fig. 21 Strict consensus cladogram based on 10 retained shortest trees found by
TNT’s traditional search option with settings as given in Table 2, analysis of
Nemesia versus Hypothetical-Group 111 (all one)

Chapter 6
The genus Ummidia Thorell 1875 in the western Mediterranean, a review
(Araneae, Mygalomorphae, Ctenizidae).
Arthur Decae
Research 2009
Journal of arachnology (in press)

Abstract.
The presence and origin of the mygalomorph spider genus Ummidia Thorell 1875 in the
western Mediterranean region is reconsidered. The traditional idea, expressed in the works of
Walckenaer and Simon, that Ummidia is a recent American import in the Mediterranean
region is opposed by the observation that at least four distinct Ummidia species inhabit
different geographical areas within the western Mediterranean. The taxonomical revision of
the Mediterranean Ummidia fauna presented here results in the description of one new species
(Ummidia algarve n. sp.), the removal of U. picea Thorell 1875 and U. algeriana (Lucas
1846) from synonymy with U. aedificatoria (Westwood 1840) and the placing of U.
occidentalis (Simon 1909) in synonymy with U. aedificatoria (Westwood 1840).
Introduction
The trapdoor spider genus Ummidia Thorell 1875 is taxonomically grouped with the genus
Conothele Thorell 1878 in the Ctenizidae subfamily Pachylomerinae (Raven 1985), recently
renamed Ummidiinae (Ortiz 2007) which name is here now used. The genus Hebestatis
Simon 1903, traditionally also included in the Ummidiinae (Simon 1903; Raven 1985), is
here excluded on grounds discussed below (see discussion). The Ummidiinae, as understood
here, are distinguished from other ctenizids on the basis of a pronounced and unique
combination of macro morphological characters (see Fig.1) that include a proximal dorsal
glabrous depression or saddle on tibia III, a sharp apophysis on the dorsal-prolateral
trochanter III, clavate trichobothria on the proximal dorsal tarsi, curvy short spines on the
lateral faces of the distal segments of the palps and anterior legs and a compact eye-group
placed on and around a distinct ocular tubercle (pers. obs.). Furthermore, spiders of the
Ummidiinae show a remarkable sexual dimorphism in the texture of the carapace. In females,
the carapace is smooth and shiny as if polished; in males, the carapace surface is dull and
typically rugose or granulated (Figs. 2-3). Finally, females of the Ummidiinae differ from
other ctenizid genera by the possession of three-partite spermathecae with a distinctly
sclerotized central section connecting the proximal and distal membranous sections (Figs. 16-
19). Geographically, the ranges of the genera Ummidia and Conothele are separated (Fig. 4)
although the presence of U. gandjinoi Andreeva 1968 in Tajikistan (see also Zonstein 2007)
appears to be a bridgehead of Ummidia in Conothele territory. The genus Ummidia, with
around 20 described and many undescribed species (Bond & Hendrixson 2005), has a
predominantly American
distribution and Conothele, with
18 recorded species (Platnick
2009) is widely distributed in the
Orient and Australasian region.
Both genera, contrary to most
trapdoor spiders, are not only
found in continental regions but
also occur on oceanic and
volcanic islands which suggests a
relatively strong capacity for
dispersal either natural or man -
aided. Conothele has been
reported from several Pacific
Islands (Pocock 1898; Berland
1938; Roewer 1963) and from
the Seychelles (Saaristo 2002).
Sd
Ap
CS
OT
Fig. 1 Right lateral view of a Ummidia (female) highlighting diagnostic
characters for the subfamily; Sd. saddle depression on dorsal tibia III; Ap.
apophysis on dorsal trochanter III; CT. clavate trichobothria; CS. curly
spines in dense spine fields; OT. ocular tubercle.
Ct

Ummidia is reported from several Caribbean islands including volcanic St. Vincent (Simon
1891) and from Bermuda (Whitehead unpub.). If this last record is correct Ummidia inhabits
an Atlantic island over a thousand kilometers off the American east coast. The ability for
aerial dispersal in Ummidia, originally reported by Bearg 1928 and recently confirmed by
Coyle 1985 and Eberhard 2005, might have played a key role in reaching such far out
locations. The presence of a geographically isolated Ummidia population in the western
Mediterranean (extreme NW Africa and southern parts of the Iberian Peninsula) is of special
interest in this respect. Is it a product of eastward cross Atlantic dispersal as Simon believed,
is it a relict of a former pan-Eurasian Ummidia/Conothele distribution, or does it have an
endemic identity of its own To solve these questions more suitable material for study, more
advanced research techniques and more coordinated research efforts will be necessary (e.g. to
establish the phylogenic relations within and between geographically isolated species), but a
taxonomical review of the currently available data on the western Mediterranean Ummidia
fauna, as presented here, is believed to be an enlightening first step.
2 3
Figs. 2-3 Ummidia algarve n. sp. 2. male, note the dull granulated carapace. 3. female, note the shiny polished
carapace.
Fig. 4 World distribution of Ummidiinae based on currently available data. Squares Ummidia,
triangles Conothele (Ummidia and Conothele are here regarded synonyms. See below).

Material & Methods
The material studied consisted of a sample of 36 Ummidia specimens (23 female + 13 male).
14 females and 11 males were recently collected from southern parts of the Iberian Peninsula
both in Spain and in Portugal. Nine females from North Africa and one male from Spain
(Cartagena) were found in Simon’s collection at the Museum National d’ Histoire Naturelle
(MNHN) in Paris and a single male from Spain was found in the collection of the British
Museum Natural of History (BMNH) London. Although not explicitly stated on the tube
labels, type specimens of Actinopus (Ummidia) algerianus (Lucas 1846) and Pachylomerus
(Ummidia) occidentalis (Simon 1909) were probably among the material studied in Paris.
Further relevant information was obtained through the kind cooperation of the Oxford
University Museum (OUM) that provided photographs of the dried type specimens of
Actinopus (Ummidia) aedificatoria (Westwood 1840). Specimens described here as U.
algarve n. sp. and U. picea Thorell 1875 are placed in the collection of the Natural History
Museum Rotterdam (NHMR).
Morphological studies were carried out with the aid of several different stereomicroscopes (as
available in the above mentioned institutions) all equipped with camera lucida drawing
devices and ocular micrometers. Photographs were taken with an Olympus E-500 reflex
camera equipped with a 50mm macro- lens and a ring-flash. Methods of measurement and
abbreviations are as given in Figures 5-9.
Abbreviations: BL = total body length, CL = carapace length, CW = carapace width, Cap =
caput length, EL = length eye-group, EW = width eye-group, SL = sternum length, SW =
sternum width, LL = labium length, LW = labium width, ML = maxillum length, MW =
maxillum width, Tar = tarsus, Met = metatarsus, Tib = tibia, Pat = patella, Fem = femur, l =
length, w = width.
All linear measures are given in millimeters.
Length/width ratios of sclerotized body parts (carapace, sternum, labium, maxillae) are given
in all descriptions. The length/width ratio of the ocular quadrangle (Fig. 6) is of important
diagnostic value. The location of the fovea is indicated by its position relative to the anterior
edge of the carapace expressed as Cap/CL (Fig. 5).
Taxonomy
Genus Ummidia Thorell 1875
Ummidia Thorell 1875, p.102.
Type species Ummidia picea Thorell (1875 p. 102) by original designation
Synonymy. All characters, morphological or behavioral, that have been proposed to
distinguish Ummidia from Conothele (Simon 1892; Roth 1982; Raven 1985; Haupt 2005)
have proved to be insubstantial (personal observation). Therefore Main’s (1982, 1998)
postulate that Ummidia and Conothele are synonyms is followed here. The name Ummidia is
retained on grounds of priority to indicate a nearly cosmopolitan genus with representative
species on all inhabitable continents and several oceanic islands.
Species list, Western Med.. The following four Ummidia species are regarded taxonomically
valid and indigenous to the Western Mediterranean region: U. algarve n.sp.; U. picea Thorell
1875; U. algeriana (Lucas 1846); U. aedificatoria (Westwood 1840). All these species are
diagnosed and described below.

CW
5 EW
6
CL
Cap
EL
BL
ML
MW
7
LL
LW
SL
8
Tib
Pat
Met
Fem
SW
Pat
Tib
9
Tar
Fem
Met
Tar
Figs 5-9 Methods of measurement and abbreviations used. 5. dorsal body parts, BL - total length of
body, CL - carapace length, CW – carapace width, Cap – length caput; 6. ocular quadrangle, EL –
length eye group, EW – width eye group; 7. ventral body parts, SL – sternum length, SW – sternum
width, LL – labium length, LW – labium width, ML – maxillum length, MW – maxillum width; 8.
anterior legs and palps length only measured along retrolateral face, Tar – tarsus, Met – metatarsus, Tib
– tibia, Pat – patella, Fem – femur; 9. posterior leg length measured along prolateral face abbreviations
as in 8.

Ummidia algarve new species Figs. 2, 3, 10, 11, 17, 21, 23.
Misidentifications:
Pachylomerus aedificatorius: O.Pickard-Cambridge 1907:818-819, Pl L. Figs. 1-6.
P. piceus: Frade & Bacelar 1931:510, Figs. 3-4: Bacelar 1937:1568-1571, Figs. 1-2.
Type specimens. All specimen collected in southern Portugal: 1 ♂ holotype, 22 March 2007
by S. Huber at Quelfes Algarve 37.217N-7.839W, slope along a field road. 1 ♀ paratype, 22
October 2006, S. Huber, east of Alte, Algarve at Pena da Rocha 37.250N−8.098W, southern
slope along walking trail.
Other material studied. 1 ♂13 August 1996 leg. P. Selden, Praia da Marinha Algarve 37.14N-
8.45W. 4 ♂♂ October 2003 leg. P. Cardoso, Ribeira de Limas Mertola Beja Alentejo
37.82N-7.62W. 4 ♂♂ October - November 2003 leg. P. Cardoso, Corredura Beja Alentejo
37.75N-7.64W. 3 ♀♀ October 2003, leg. P. Cardoso, Ribeira de Limas Mertola, Beja
Alentejo 37.82N-7.62W. 2 ♀♀ 22 August 1996, leg P. Selden, Praia da Oura Algarve
37.08N-8.24W. 1 ♀ 16 August 1996, leg. P. Selden, Belem-Monchique Algarve 37.31N-
8.59W. 1 ♀ 15 August 1986, leg. P. Selden, Senhora de Rocha Algarve 37.10N-8.37W.
Etymology. The species is named after the region and former Moorish kingdom Algarve in
South Portugal where it was first discovered (O. Pickard-Cambridge 1907). The
geographically inspired name was chosen because it is regarded appropriate for a trapdoor
spider species since these species tend to be local endemics. An earlier suggestion by Amelia
Bacelar (1937: 1569) to name the Portuguese Ummidia species after O.Pickard-Cambridge
who first reported it in scientific literature is not followed because of potential confusion with
Conothele cambridgei Thorell 1890 upon future revision of the Ummidiinae.
Diagnosis. Differs from all other western Mediterranean Ummidia species by the small
straight mushroom shaped spermathecae (Fig. 17) and the warty texture of the abdominal
cuticle. Differs from U. piceus by the relatively short, strong and smoothly curved embolus
with sub-apical fishhook tooth (Fig. 23) and low ocular quadrangle ratio (lw = 0.58).
Measurements. Male holotype: BL = 14.5; CL = 6.8; CW = 6.7; Cap = 4.8; EL = 3.0; EW =
4.2; SL = 4.2. SW = 3.7; LL = 0.8; LW = 1.3; ML = 2.7; MW= 1.6.
Tar Met Tib Pat Fem Total
Palp 1.4 ⎯ 3.3 2.1 4.4 11.2
Leg 1 1.2 2.5 3.3 2.8 5.3 17.6
Leg 2 1.2 2.2 3.2 2.8 4.9 16.4
Leg 3 1.6 2.1 2.6 2.1 4.1 14.6
Leg 4 1.8 3.5 2.3 2.8 5.5 18.6
Description. Male holotype (Fig. 2): Carapace: (l/w = 1.0) black with shades of dark red,
cephalic area slightly darker than thorax part, few bristles on clypeus and on cephalic area
crest, cuticle strongly granulated with thicker rim around edges Clypeus: narrow. Cephalic
area: moderately elevated. Eye-group: (l/w = 0.6) eight eyes compactly grouped in two rows
on and around low ocular process, anterior row strongly procurved, posterior row straight
(Fig. 10). Fovea: position Cap/CL = 0.7, deep, smoothly procurved. Chelicerae: strong,
dorsally black, cuticle granulated, few setae mainly in apical zone; ventrally warm orange
brown, cheliceral furrow lined with rows of teeth on either side, 5 prolateral, 6 retrolateral.
Rastellum: tight group of strong teeth on well developed apical process. Fangs: with distinct
serrated ventral ridges. Maxillae: (l/w = 1.7) trapezoid, orange brown, cuspules groups
strongly reduced both in sizes of cuspules and in numbers. Palp Trochanters: without ventral
cuspules. Labium: (l/w = 0.6) triangular shape, dark grey-brown, cuspules reduced. Sternum:

(l/w = 1.1) brown, grading to lighter shades posterior, setae concentrated in lateral zones
around glabrous central zone. Anterior Legs & Palps: dark brown, tarsi and metatarsi legs I &
II light yellow brown, cuticle proximal segments ribbed and granulate, spines absent from
palps and arranged in ventral lateral groups on tibiae, metatarsi and tarsi legs I&II; strong
distal spines on ventral patellae I & II; filiform trichobothria present on all metatarsi and
tibiae, filiform + clavate trichobothria on all dorsal tarsi in dense disordered groups; dense
scopula of short hairs on ventral tarsi and metatarsi I & II; paired claws with variable number
of strong lateral teeth (sometimes fused into an irregular comb); 3 rd claw very small. Leg III:
trochanter apophysis reduced armed with a strong spine, femur bent and ventrally enlarged
(Fig. 9: Fem), with spines distributed on dorsal and prolateral faces, patella short with group
of sharp spines along dorsal prolateral side, tibia with shallow saddle and spines along distal
edge and retrolaterally, metatarsus narrowing distally with strong spines
along distal edge, tarsus cylindrical with numerous spines ventrally and distally, paired claws
with one large and one small tooth. Leg IV: lighter in color than other legs, femur finely
ribbed with few short spines dorsally, patella with elliptical glabrous patch dorsally lined with
fine denticles proximally, distally segments unmodified. Abdomen: with strongly developed
wart-like sockets for individual bristles as in female. Spinnerets: PMS digitiform, proximally
light brown, distally creamy white with numerous small spigots and one apical macro-spigot,
PLS three equally short segments all proximally light brown and distally creamy white fields
with numerous fine spigots and few macro-spigots. Bulb: (Figs. 21-23) as described in
diagnosis.
Measurements. Female paratype (Fig. 3): BL = 14.5; CL = 6.9; CW = 6.2; Cap = 4.8; EL =
2.8; EW = 5.5; SL = 4.5; SW = 3.9; LL = 1.0; LW =1.6; ML = 2.9; MW = 1.6.
Tar Met Tib Pat Fem Total
Palp 1.9 ⎯ 2.0 2.1 3.7 9.7
Leg 1 1.0 1.7 2.4 2.6 4.1 11.7
Leg 2 1.0 1.4 2.0 2.4 3.7 10.6
Leg 3 1.2 1.4 1.7 2.1 3.6 9.9
Leg 4 1.4 2.2 2.5 2.6 4.7 13.5
Description. Female paratype. Carapace: (l/w = 1.1) smooth, shining, with bristles only
around eye-group, short crest-row with two lateral bristle rows reduced to only one pair of
bristles. Clypeus: protracted onto membranous connection between carapace and chelicerae.
Cephalic area: smoothly elevated. Eye-group: (l/w = 0.6) eight eyes placed in two rows near
anterior edge of carapace and compactly set around small ocular process, anterior row
strongly procurved, posterior row slightly recurved. Fovea: (Cap/CL = 0.7) deep, strongly
procurved with distinct light colored anterior tips. Chelicerae: massive, black contrasting with
color of carapace, bristles concentrated along dorsal crests, ventrally orange, cheliceral furrow
with 5 prolateral and 7 retrolateral denticles, rastellum of compactly set short teeth on strongly
developed process. Fangs: strong, blunt with serrated inner ridge. Maxillae (l/w = 1.8) subrectangular,
anterior light orange brown with greyish scopula, cuspules strongly developed,
organized in two groups; one with 25 larger cuspules more proximal and anterior and one
with 21 smaller cuspules distal and posterior, anterior apical maxillary process indistinct. Palp
trochanter: with distinct group of cuspules. Labium: (l/w = 0.6) semi-dome shaped, posterior
sloping steeply to labial furrow; distinctly bi-colored with anterior light crescent carrying an
oval-shaped group of 11 strong cuspules. Sternum (l/w = 1.2) smooth, with large glabrous
central area (fused sigilla) and evenly set setae along lateral zones. Anterior Legs & Palps:
dense lateral fields of short curvy and curved spines on tarsus, metatarsus and tibia (absent
from retrolateral tarsus, metatarsus and tibia of leg II), anterior patellae and femora without

spines with the exception of one distal prolateral spine on palp patella. Leg III: blunt pointed
apophysis on prolateral dorsal trochanter, femur curved and ventrally enlarged (Fig. 9: Fem),
patella short strong with prolateral field of short straight spines, tibia with dorsal proximal
dark colored, glabrous, saddle flanked on either side by narrow membranous slits, distal field
of short curved spines on distal upward curved part of tibia (Fig. 9: Tib), metatarsus short
with dorsal field of strong short spines along full length of segment, tarsus short with dense
prolateral spine field along full length of segment and retrolateral spine field distally
restricted. Leg IV: trochanter and femur unmodified, patella dorsal glabrous patch with
prolateral dense fields of fine cuspules, tibia unmodified without prolateral spines, metatarsus
unmodified with dorsal and ventral prolateral rows of 2 spines, tarsus unmodified with apical
prolateral group of strong short spines. Trichobothria: large groups of filiform trichobothria
and small groups of clavate trichobothria dorsal on all tarsi, few filiform trichobothria in
disordered row on dorsal metatarsi; two small rows of filiform trichobothria in proximal half
of dorsal tibiae. Abdomen: egg-shaped with evenly distributed bristles set in strongly
developed wart-like sockets. Spinnerets: as described for male. Spermathecae: (Fig. 17) short,
distally converging mushroom-shaped, proximal part tubular, lightly glandular, medial part
sclerotized, distal part donut-shaped lightly glandular.
Variation. Morphological variation in this species is small, the bulb structure and the structure
of the spermathecae were found to be constant in all specimens studied. Total body sizes vary
between 12.6mm and 18.2mm in males (n = 10) and between 12.4mm and 19.6mm in females
(n = 8). Carapace shape as judged by the CL/CW ratios was found to be quite constant in
females (CL/CW 1.2 – 1.3: n = 8) and somewhat more variable in males (CL/CW 1.0 – 1.2; n
= 10).
Natural History. U. algarve is reported to be very common and occurring in quite dense
populations often in close association with nemesiid trapdoor spiders (S. Huber pers. comm.).
The burrow structure, with a trapdoor at the entrance of the burrow and a second up-side
down trapdoor in the bottom of the burrow has attracted much attention in literature
(O.Pickard-Cambridge 1907; Bacelar 1937; Buchli 1962). This type of burrow distinguishes
U. algarve from all other Mediterranean Ummidia species that all construct simple types of
trapdoor burrows with no other internal structures than a dense silken lining of the burrow
walls. Buchli 1962 reported that the inverted trapdoor at the bottom of the burrow might only
be built by female spiders. Although this curious type of burrow is presently not known from
any other Ummidia species a similar type of burrow has recently been reported from
Conothele varvarti in eastern India (Siliwal et. al. 2009).
Ummidia picea Thorell 1875 Figs. 12, 13, 16, 20, 22.
Ummidia picea Thorell 1875a:102.
U. picea: Thorell 1875b:121.
U. piceus: Frade & Bacelar 1931:511, Fig. 4bis.
Pachylomerus aedificatorius: Simon 1909:42. Misidentification.
Diagnosis. differs from all other western Mediterranean Ummidia species by double bent
central sclerotized section of the spermathecae (Fig. 16). Differs from U. algarve by the long
slender curved embolus, proximal sclerite with distal denticles (Fig. 22).
Material studied. 1 ♂ (described) collected as juvenile 6 April 2007, adult 11 September
2008, leg Decae, Barranco de Rio Higueron, Frigliana, Andalucia 36.802N-3.875W. 1 ♀
(described) 4-6 April 1989 leg Decae, Nerja, Andalucia 36.764N-3.865W.1 ♂ MNHN Coll
Simon (undated) Cartagena, 37.61N-1.00W. 1 ♂ MNHN 18 September 1919 BMNH
Cartagena, 37.61N-1.00W. 3 ♀♀ 4-6 April 2007 leg Decae, Barranco de Rio Higueron,

Frigliana, Andalucia 37.80N-3.83W. 2 ♀♀ 4-6 April 1989 leg Decae, Nerja, Andalucia
36.764N-3.865W.
Measurements. Male: BL =13.3; CL = 5.6; CW = 5.2; Cap = 3.9; EL = 2.7; EW = 4.1; SL =
3.2; SW = 2.7. LL = 1.0; LW = 1.2; ML = 2.1; MW = 1.2.
Tar Met Tib Pat Fem Total
Palp 1.2 ⎯ 2.8 1.7 3.9 9.6
Leg 1 1.2 2.3 2.9 2.3 4.7 13.5
Leg 2 1.2 2.2 2.5 2.2 4.2 12.4
Leg 3 1.4 2.2 2.1 1.9 3.5 11.0
Leg 4 1.7 3.0 2.7 2.0 4.5 13.8
Description. Male: Carapace: (l/w = 1.1) glabrous, surface finely striated and granulated.
Clypeus: width as longest diameter of ALE with two brownish lines running from the base of
ALE to clypeus edge, setae fully absent. Cephalic area: elevated, laterally not delineated from
thorax part of carapace. Fovea: (Cap/CL = 0.7) regularly procurved. Eye-group: (l/w = 0.6) on
dome-shaped process, anterior row strongly procurved, posterior row straight, ALEs largest,
PMEs teardrop shaped. Chelicerae: basal segment striated and granulated as carapace, black
dorsally grading into warm brown ventrally, distal sharp bristles evenly spaced around very
short strong teeth of the rastellum. Rastellar process, ventrally pronounced, fangs brown, long,
sharp and slightly translucent with ventral retrolateral serrated ridge. Cheliceral furrow warm
brown, bordered with rows of teeth 5 prolateral, 6 retrolateral. Maxillae: (l/w = 1.8) subrectangular,
dark brown, proximally lighter, with distinct light colored anterior edges and
silvery white scopulae, cuspules reduced and irregularly spread along ventral surface,
proximally more concentrated, distally absent. Labium: (l/w = 0.8) relatively long striated,
with contrasting color zones, proximal dark brown, distal light brown, small group of distal
cuspules. Sternum: (l/w = 1.2) greyish brown, lateral zones lighter than central zone, centrally
fussed sigilla, widely spaced sharp bristles predominantly in finely striated lateral zones.
Palps: long and slender, femur is longest segment, tegument structure, color and setae as
described for legs, spines absent from all segments. Legs: dorsally black striated, ventrally
greyish, tarsi and metatarsi I & II ventrally light colored and scopulate. Leg III: trochanter
with small dorsal apophysis, femur slightly thickened with group of 4 short strong apical
spines dorsally, patella with short curved spines along dorsal prolateral face, tibia with
transverse striated saddle and few sharp spines irregularly placed, metatarsus with numerous
irregularly placed sharp spines, tarsus with numerous ventral spines. Leg IV: trochanter
unmodified, femur with few apical dorsal short spines, patella with longitudinal dorsal
glabrous zone flanked on either side by short spiny bristles, tibia with ventral longitudinal row
of sharp spines, metatarsus and tarsus with numerous sharp ventral spines, dorsal patella IV
with central brown zone. Filiform trichobothria on all dorsal tibiae, metatarsi and tarsi.
Clavate trichobothria in small groups on proximal dorsal surfaces of all tarsi. Abdomen:
evenly covered with fine, spiky bristles, dorsally purplish brown with irregular creamy
blotches, ventrally yellowish brown, integument not warty. Spinnerets: PMS short, light
colored with few apical spigots, PLS three segmented with groups of spigots on ventral distal
parts of proximal and medial segment and dense apical spigot field on domed distal segment.
Bulb (Figs. 20-22).
Measurements. Female: BL = 26.0; CL = 9.5; CW = 8.5; Cap = 6.8; EL = 4.8; EW = 7.0; SL
= 6.5; SW = 5.4; LL = 1.2; LW = 1.9; ML = 3.5; MW = 2.1.

Tar Met Tib Pat Fem Total
Palp 2.9 ⎯ 3.7 3.3 5.0 14.9
Leg 1 1.2 2.4 3.5 3.8 5.6 16.5
Leg 2 0.7 2.4 3.3 3.4 5.5 15.3
Leg 3 1.7 1.7 2.7 2.6 4.5 13.1
Leg 4 2.1 3.4 3.1 3.2 6.1 17.8
Description. Female: Carapace: (l/w = 1.1) smooth, shining and shaded brown, darkest zones
around fovea and above coxa III; crest-line narrow, dark contrasting with lighter crest-zone;
crest-bristles strongly developed in straight line and only in anterior half of crest-zone, few
finer bristles lateral of crest-zone; no setae along carapace edge. Clypeus: mottled brown, with
small group of setae on protracted semi-circular process anterior of eye-formation. Cephalic
area: smoothly elevated. Eye-group: (l/w = 0.6) on distinct ocular process, anterior row
strongly procurved, posterior row straight; ALE largest, AME slightly wider than their
diameter apart, PME pearly and caudally protracted projecting caudally beyond PLE. PLE
distinctly smaller than AME; groups of strong setae both anterior and posterior of AME on
ocular process. Fovea: (Cap/Cl = 0.7) deep, strongly and smoothly procurved. Chelicerae:
basal segment strong, dark brown distally grading to black and contrasting in color with
carapace, dorsally slightly lighter in color than laterally, ventrally bright orange brown in and
along the cheliceral furrow, glabrous between three distinct longitudinal zones with bristles.
Cuticle dorsally smooth, distally (in rastellar zone) striated; furrow lined with two irregular
rows of 7 or 8 strong stubby teeth, no denticles on furrow bottom, rastellum dense group short
strong teeth on distinct process. Fangs: strong, short, and blunt. Fang ridge: smooth. Maxillae:
(l/w = 1.7) cuspules spiky spread in two size-classes over ventral surface, about 35 larger
cuspules anterior, about 33 small more posterior. Palp trochanters: with short cusp like setae.
Labium: (l/w = 0.6) somewhat diamond shaped, 10 strong distal cuspules in distal halve.
Labial furrow: glabrous, shallow, with two distinct elliptical sigilla. Sternum: (l/w = 1.2) light
brown with dark edge, setae mainly in peripheral zone, sigilla fused in central glabrous field.
Legs: dorsally dark brown, ventrally lighter, ventral femora III & IV light yellowish brown,
anterior coxae darker than posterior coxae; (spine patterns) dense fields short curvy and
curved spines on lateral tibiae, metatarsi and tarsi I & II, short straight spines on all patellae
and on tibiae, metatarsi and tarsi III & IV, no spines on femora; patella III with transverse row
of short strong spines along the distal edge, tibia III with dense transverse group of short
strong spines along full dorsal width distal of black saddle depression, metatarsus III with
group of short strong spines along full dorsal length of segment (apical spines strongest),
tarsus III with few prolateral spines and dense spine groups ventrally around apical claw
implant; leg IV with dense ‘rasp-like’ field of very small short spines on dorsal patella and
only few fine spines in distal halves of tibia, metatarsus and tarsus; trochanter III with anterior
dorsal apophysis. Tarsi with proximal groups of clavate trichobothria (reduced or absent in
posterior legs), surrounded by irregularly placed filiform trichobothria; metatarsi with central
dorsal longitudinal row of very fine filiform trichobothria (absent from leg I), tibiae with two
distally converging rows of very fine filiform trichobothria in proximal quarter; leg scopulae
absent; paired claws with one long proximal tooth & one much smaller more distal tooth, 3rd
claw vestigial. Abdomen: evenly covered with fine setae, mottled purplish grey with irregular
light colored blotches, ventrally overall lighter color. Spinnerets: PMS: digitiform, with
distinct lighter colored apical spinneret field with few micro-spigots and one macro-spigot,
PLS all three segments short and distally light colored, proximal and medial segment with
transverse distal rows of macro-spigots, distal segment with apical spigot field with
exclusively micro-spigots. Spermathecae: see diagnosis.

Variation.The emboli of the 2 male spiders found one in the BMNH, London and one in
MNHN, Paris were not as elongated as the embolus in the specimen here described and
figured. This could well be the result of handling damage over a long period of time. The
material available however was insufficient to test this hypothesis and the possibility that the
Spanish Ummidia population is more diverse at the species level than presently conceived
cannot be ruled out.
Remarks. Thorell’s original description of the male of U. picea is very short and inadequate
(Thorell 1875: 102), the female described as U. picea by Frade & Bacelar 1931 actually was a
specimen of U. algarve. Therefore, both sexes are fully described here.
Natural History. U. picea was found to inhabit steep banks along trails, creeks and canyons in
Andalusia, particularly in shady locations.Different from U. algarve (see above), U. picea
does not form aggregations of nests. Single burrows were found throughout the area.
Specimens were collected from a number of burrows for further study of morphology
(taxonomy) and behavior in the laboratory. The trapdoors were typically, as reported for
several Ummidia species, placed sideways or even upside down with respect to the slope. A
remarkable difference in behavior between U. picea and U. algarve is the immediate strong
holding down of the trapdoor in the first species upon disturbance, and the absence of this
behavior in the second species. U. picea burrows may be found close to burrows of both
nemesiid Nemesia and Cyrtauchenius trapdoor spiders.
Ummidia algeriana (Lucas 1846) comb. nov. Figs. 14, 18.
Actinopus algerianus Lucas 1846: 96-97, Pl. 1, fig 5.
Cteniza algeriana: Ausserer 1871: 155.
Pachylomerus aedificatorius: Simon 1892 Fig. 86; 1903:887, Fig. 1048; 1909:42-43.
P. aedificatorius: Frade & Bacelar 1931:509 Figs. 1-2. Misidentifications.
Remarks. Lucas’(1846: 96-97) description and figures are very accurate and complete, here
only further observations on the morphology of the spermathecae and measurements are
given. The measurements are taken from a specimen in Lucas’ type series, the spermathecae
are drawn after one of the larger specimens in Simon’s collection.
Material studied. 9 ♀♀ including a probable type specimen collected by Lucas and found in
Simon’s collection at the MNHN, Paris.
Diagnosis. Differs from all other Mediterranean Ummidia species in the possession of a
twisted central sclerotized section in the spermathecae (Fig. 18). Differs from U. aedificatoria
in the higher l/w-ratio of the ocular quadrangle (Figs.14-15) the strong development of the
rastellar process, the deep labial furrow with distinct elliptical sigilla and the texture of the
abdominal cuticle.
Measurements. Female: BL = 24.4; CL = 8.6; CW = 7.9; Cap = 5.0; EL = 3.7; EW = 6.1; SL
= 5.5; SW = 5.0; LL = 1.0; LW = 1.7; ML = 2.7; MW = 1.4.
Tar Met Tib Pat Fem Total
Palp 2.4 ⎯ 2.9 2.8 4.8 12.9
Leg 1 1.1 2.3 2.9 3.5 5.2 15.0
Leg 2 1.3 2.2 2.4 3.2 4.6 13.7
Leg 3 1.4 1.9 2.6 2.4 4.3 12.7
Leg 4 1.7 3.0 3.1 3.2 5.7 16.8

Variation. Total sizes (BL) in the sample of 9 ♀♀ varied between 29.1mm and 17.0mm.
Variation in the shape of the carapace (CL/CW = 1.1-1.2; n =9), the position of the fovea
(CL/Cap = 1.4-1.5; n = 9) appeared to be very low. Differences found in the length/width
ratio of the ocular quadrangle (l/w = 0.6 – 0.8; n=9) suggest some local geographical
variability or cryptic diversity.
Natural History. Simon (1888) reported this species to be widespread in the Tell region of
Algeria and western Tunisia where its burrows were all dug in steep to vertical surfaces along
roads and rivers. The burrows were reported to be shallow (6-10cm), internally lined with
dense white silk and closed at the entrance by a stiff thin trapdoor.
Ummidia aedificatoria (Westwood 1840) Figs. 15,19.
Actinopus aedificatorius Westwood 1840:175 Pl. 10.
Sphodros aedificatorius: Walckenaer, 1842: 438.
Cteniza aedificatoria: Ausserer 1871: 155.
Pachylomerus occidentalis Simon 1909:8 NEW SYNONYM.
Pachylomerus occidentalis: Frade & Bacelar 1931:512, figs 5-6.
Remark. The type of U. aedificatoria is a dried specimen in the Oxford University Museum
collection, of which photographs by Ray Gabriel were seen, but diagnostic details of the
sexual organs could not be studied. Westwood’s original description fortunately is very
detailed and extensive and is furthermore accompanied by a good set of illustrations
(Westwood 1840: Plate 10). Two characters of supposedly diagnostic value − the relatively
short ocular quadrangle and the reduced rastellar process− are sufficiently clear in the dried
specimen and the illustrations to synonymize Westwood’s type of U. aedificatoria with
Simon’s type of U. occidentalis that originated from roughly the same type location (Tangiers
province in northwestern Morocco. Fig. 24). Because Simon’s spider at the MNHN Paris is
the only specimen available for closer study, the limited additional diagnostic and descriptive
information given here is based on that specimen.
Material studied. 2 ♀♀ on photographs in the Westwood collection, preserved in dry
condition in the Oxford University Museum. 1 ♀ found in Simon’s collection at the MNHN,
Paris, probable type specimen of U. occidentalis
that is here synonymised with U. aedificatoria.
Diagnosis. Differs from all other Mediterranean
Ummidia species by the short bent central
section of the spermathecae (Fig. 19), the low
length/width ration of the ocular quadrangle
(Fig. 15) and the reduced rastellar process.
Measurements. Female: BL = 18.1; CL = 8.2;
CW = 6.7; Cap = 5.8; EL = 0.4; EW = 0.8.
Variation. Only three female specimens are
currently known, two of which have been
preserved in dried condition for almost 170
years. Nevertheless it is clear that U.
aedificatoria falls in the same size range as the
other western Mediterranean Ummidia species.
Total body lengths of adult females in the small
sample range from 18.1 to 29.0mm.
Natural History. Neither Westwood 1840 nor
Simon 1909 provides any information about the
natural conditions in which U. aedificatoria is
10 11
12
14
13
15
Figs. 10 -15 Eye-formations in dorsal view of western
Mediterranean Ummidia species. 10. U. algarve male
holotype; 11. U. algarve female paratype; 12. U. picea
male; 13. U. picea female 14. U. algeriana female; 15. U.
aedificatoria female.

found. Westwood, however, received the spiders he described alive in their natural burrows.
From Westwood’s descriptions and figures, and also from photographs of the preserved
burrow material in the Oxford University Museum, it might be concluded that U.
aedificatoria builds the typical shallow silk lined trapdoor burrow that is very similar to the
burrows of U. algeriana and U. picea.
16 17
20 21
18 19
22 23
Figs. 16 -19 Studies of the spermathecae of western
Mediterranean Ummidia species in ventral view. Note
the sclerotized central sections of the spermathecae:
16. U. picea (note double bent central sections); 17. U.
algarve (note mushroom type and cup-shaped central
sections); 18. U. algeriana (note twisted central
sections); 19. U. aedificatoria (note short bent central
sections). Scale-line = 1mm
Figs. 20 - 23 Studies of the right bulb in Iberian Ummidia
species: 20. U. picea prolateral; 21. U. algarve prolateral; 22.
U. picea retrolateral (arrow indicates denticles); 23. U.
algarve retrolateral (arrow indicates fish-hook). Scale-line =
1mm
Fig. 24 Currently known distribution of the genus Ummidia in the
western Mediterranean Region based on specimens seen in this
study: black circle = U. aedificatoria, grey square = U. algarve,
grey triangle = U. algeriana, grey circle = U. picea.

Discussion
Simon (1903 pp. 887-888) included three genera (Ummidia, Conothele and Hebestatis) in his
Pachylomereae based on the shared absence of lateral sternal sigilla. Raven 1985 followed
this classification but used the eye tubercle and saddle tibia. However, the probable type
specimen in Simon’s collection (AR12317 MNHN examined) labelled Hebestatis theveneti
actually possesses lateral sternal sigilla and furthermore shows sufficient morphological
differences from both Ummidia and Conothele, (dorsal saddle on tibia III not pronounced and
not glabrous, absence of curvy spines, absence of tarsal clavate trichobothria, absence of
centrally sclerotized spermathecae) to exclude the genus Hebestatis from the Ummidiinae.
The taxonomy of the genus Ummidia in the western Mediterranean has been disputed from
the start. Shortly after Westwood 1840 had presented the discovery and description of his
Actinopus aedificatorius (now Ummidia aedificatoria) from Morocco to the Entomological
Society of London, Walckenaer 1842 explicitly expressed his doubts about these findings.
Walckenaer suspected that Westwood was mistaken in either the origin or the identity of the
species described (see also Westwood 1840: 181). Westwood had obtained the specimens he
presented and described from Mr Drummond Hay, H.M.’s agent and Consul-general at
Tangiers in the far northwest of Morocco. Furthermore, he classified this newly discovered
species as belonging “to the same genus as Mr. Sell’s Jamaica species, to which it is so
closely allied as scarcely to present any specific distinction beyond that of size” (Westwood
1840: 174-175). Mr. Sell’s Jamaican species is now known as Ummidia nidulans (Fabricius
1787) and the fact that Westwood regarded his Moroccan species so closely allied with a
Caribbean species caused much skepticism among the leading arachnologists at that time. The
discovery of several other Ummidia species in the Americas in the 19 th century (e.g. Ausserer
1871: 146-147) reinforced the idea that Ummidia was naturally restricted in its distribution to
the New World and that the incidental finding of Ummidia east of the Atlantic must be the
result of human mediated introduction. This idea was most vividly expressed in the work of
Simon:
“le genre Pachylomerus, qui est assez nombreux, est américain, il a cependant un
représentant au Japon (d’après Dönitz) et un autre dans la région
méditerranéenne occidentale (Algérie et Espagne), mais ce dernier paraît y avoir
été introduit en même temps que les Opuntia et les Agave d’origine américaine”
(Simon 1892: p.86).
In short, Simon regarded Pachylomerus (Ummidia) in the western Mediterranean as an
American species that was introduced probably with imports of ornamental plants. Central in
Simon’s opinion was his conviction that only one Ummidia species (U. aedificatoria
Westwood 1840), inhabits the western Mediterranean. In forming his opinion about Ummidia
in the western Mediterranean, Simon had apparently overlooked the work of Lucas who had
described a second Ummidia species in North Africa, this time from eastern Algeria. Lucas
(1846: 96-97) had collected this new species near the town of Bône (now Annaba) over
1200km east of the locality from where Westwood had obtained U. aedificatoria. Lucas
furthermore was well aware of the diagnostic differences between his new species, that he
described as Actinopus algerianus (now Ummidia algeriana) and Westwood’s U.
aedificatoria. He distinguished the two species on the grounds of differences in the
morphology of the rastellum, the curved anterior edge of the sternum, the different texture of
the abdominal cuticle and the leg-formula (Lucas 1846). Lucas does not mention any
differences in the configuration of the eyes (the most commonly used diagnostic feature in
classical species level Ummidia taxonomy), but in the excellent figures that both Lucas (1846:
Pl. 1-5) and Westwood (1840: Pl. 10) produced of their type specimens, a difference in the
configuration of the eyes between U. aedificatoria Westwood and U. algeriana Lucas is
obvious.

It is therefore unclear why Simon, who collected Ummidia in Algeria, never mentioned the
differences that Lucas had found between U. algeriana and U. aedificatoria. Probably as the
result of a preconceived conviction that Ummidia must be a recent American import in the
western Mediterranean Simon, until 1909, regarded all Old World reports of Ummidia as
reports of U. aedificatoria (Westwood 1840). It was Simon 1889 who declared Thorell’s U.
picea from Spain to be the male of U. aedificatoria and who convinced O.Pickard-Cambridge
(although not wholeheartedly, see Pickard-Cambridge 1907: 818 and Frade & Bacelar 1931:
507) that his newly found Ummidia species from Portugal also was U. aedificatoria. When
Simon 1909 worked on the spiders collected by Martínez de la Escalera in Morocco he
encountered a species of Ummidia that clearly differed from the Algerian Ummidia species
that he knew so well. That this new species was collected near Tangiers, roughly the typelocation
of U. aedificatoria, did not spark his realization that, for the first time, he might
actually see Westwood’s species under the microscope, so he proceeded to describe a new
species Pachylomerus (Ummidia) occidentalis Simon 1909. Simon’s description of U.
occidentalis is brief and without illustrations and he states that the new species is sub similar
−cui subsimilis est− to U. aedificatoria from which it only differs in the shape of the eyeformation,
the spine pattern on the tibia of the palp and the number of spines on the prolateral
face of tibia IV. Simon concludes his description of U. occidentalis with the somewhat casual
and vague remark: remplace probablement le P. aedificatorius au Maroc (Simon 1909: 8).
The discovery of a second Ummidia species in North Africa apparently did not change
Simon’s opinion about the origin of the Mediterranean Ummidia species and a year after his
description of U. occidentalis Simon notes: le genre Pachylomerus (read Ummidia) dont tous
les autres représentants sont américains (Simon 1910: 266).
Frade & Bacelar 1931 in their revision of the Mediterranean Ummidia species found that the
spine patterns Simon used are were strongly variable in Ummidia at the species level and even
in individual specimens rendering them unfit for use in species level taxonomy. Furthermore
Frade and Bacelar 1931 found that at least three different Ummidia species inhabit the
Mediterranean, indicating that Simon had underestimated the species diversity in the region.
Nevertheless, Simon’s arachnological influence has proved to be far reaching and today, a
hundred years after his slip of not recognizing U. aedificatoria in the Moroccan spider
material he examined, Simon’s classification of western Mediterranean Ummidia species is
still reflected in the World Spider Catalog (Platnick 2009).
The species descriptions given above however show that at least four different Ummidia
species inhabit the western Mediterranean. Moreover, these species each appear to inhabit
distinct geographical regions. U. algarve is common in southern Portugal and probably
extends into southwestern Spain where the great delta (las marismas) and the alluvial plains of
the Guadalquivir may separate it from U. picea. U. picea is widespread in southern Spain
between Valencia and Malaga and probably beyond towards Gibraltar. U. aedificatoria has
until now only been reported from Tangiers in the far northwest of Morocco. U. algeriana is
widespread in the Algerian Tell region as Simon 1888 reported from Bougie (Bejaia) in the
west to Kroumirie (Ain Drahan) in the east and probably even further east into Tunisia. There
are some indications (variation in configuration of the eyes, spermathecae morphology and
some other morphological traits) that further geography related cryptic diversity is present in
the Mediterranean Ummidia populations, but the still very limited availability of well
documented and usefully preserved material for study currently prevents further conclusions.
The idea that Ummidia is a recent and probably human aided introduction in the western
Mediterranean appears to be unlikely on grounds of the here presented observations. The
question if the Mediterranean Ummidia species complex is more closely related to the
American Ummidia fauna or to the Ummidia (Conothele) fauna of Australasia remains to be
investigated.

Acknowledgements
This study would not have been possible without the collection efforts and generous provision
of U. algarve n. sp. specimen by Pedro Cardoso, Siegfried Huber and Paul Selden. Jason
Bond kindly provided some American Ummidia specimens that importantly helped to develop
an understanding of Ummidia morphology and aided in my search for diagnostic characters.
Janet Beccaloni and Christine Rollard kindly allowed me to search the collections of the
BMNH, London and the MNHN, Paris for Ummidia and provided all the working space and
equipment necessary for study of those collections. Gilbert Caranhac developed the
invaluable database ‘What’s in What Pot’ to make Simon’s mygalomorph collection
accessible, Sudhikumar Ambalaparambil Vasu and Frederik Hendrickx helped to start-up my
Ummidia research. Danny Boomsma copied and sent all requested reference literature needed
from the Library of the Dutch Entomological Society in Amsterdam. Ray Gabriel provided
the photographs of Westwood’s type specimens in the collection of the Oxford University
Museum. Dries Bonte, Rudy Jocqué and Nollie Hallensleben and an anonymous referee
helped to improve the paper by critically reading the manuscript. Robert Raven is finally
thanked for expert advice and enlightening comments. Jean Pierre Maelfait is specially
thanked and remembered for his generous support and for opening up the possibility for a
research project at the University of Gent of which this Ummidia study is only the first part.

Chapter 7
Systematics of the trapdoor spider genus Cyrtocarenum Ausserer 1871
(Araneae, Mygalomorphae, Ctenizidae).
Arthur Decae
Research 1979-1996
Bulletin of the British arachnological Society (1996) 10(5): 161-170.

Abstract
The genus Cyrtocarenum Ausserer 1871 contains two species: Cyrtocarenum cunicularium
(Olivier 1811) and Cyrtocarenum grajum (C. L. Koch 1836). Both species are common in
Greece. Their range outside Greece is largely unknown. Four species attributed to the genus
— C. hellenum Doleschall in Ausserer 1871, C. ionicum (Saunders 1842), C. lapidarium
(Lucas 1853) and C. werneri Kulczynski 1903 — are placed in synonymy with C.
cunicularium. The synonymy of C. tigrinum (L. Koch 1867) and Cteniza orientalis Ausserer
1871 with C. cunicularium is confirmed. Males and females of C. cunicularium and C.
grajum are redescribed on the basis of recently collected material from the respective type
localities; diagnostic characters are given and illustrated. The male of C. grajum is described
for the first time. Keys for Mediterranean Ctenizinae and Cyrtocarenum species are included.
Introduction
Between 1811 and 1903, eight species were placed in the genus Cyrtocarenum Ausserer 1871.
No new species have been attributed to the genus since. Seven species were reported from the
north-eastern Mediterranean, and one species, C. rufidens Ausserer 1871, from Southern
Africa. Simon 1903 erected the genus Stasimopus to contain the African species, thus
restricting the range of Cyrtocarenum to Greece and western Anatolia. The original
descriptions of the seven Mediterranean species of Cyrtocarenum are generally brief, based
on one or few, sometimes immature, specimens and always restricted to one sex only.
Ausserer 1871 and Simon 1884 provided keys and descriptions for the six Cyrtocarenum
species recognised in their time. Both authors inevitably had to base their work on the limited
material and sparse information then available. Between 1979 and 1994 Cyrtocarenum has
been extensively collected in Greece. Comparison of this new material with types and other
specimens present in major European museum collections shows that C. hellenum Doleschall
in Ausserer 1871, C. ionicum (Saunders 1842), C. lapidarium (Lucas 1853), C. tigrinum (L.
Koch 1867), C. werneri Kulczynski 1903 and Cteniza orientalis Ausserer 1871 are all
synonyms of C. cunicularium (Olivier 1811) and that C. grajum (C.L. Koch 1836) is a
separate species. The brief and incomplete references in the older literature do not fit modern
standards in systematic biology. The redescriptions and keys presented here aim at providing
an improved taxonomic basis for further study of the Mediterranean ctenizid fauna.
The problematic status of the genus Cyrtocarenum Ausserer 1871
Ausserer 1871 described Cyrtocarenum as distinct from, although closely related to Cteniza
Latreille 1829 and Aepycephalus Ausserer 1871. In an additional note Ausserer (1871: 152)
emphasised the close relationship between Cteniza, Aepycephalus and Cyrtocarenum. The
general shape of the cephalic region, the configuration of the eyes and the relative sizes of the
eyes were presented as characters to distinguish the three genera, but without quantification
that would make verification possible. Moreover, there are good grounds to doubt the separate
status of the three genera in Ausserer's work, particularly from Ausserer's own apparent
confusion in this respect when he described Cteniza orientalis Ausserer 1871, recognised by
later workers (Roewer 1942; Bonnet 1956) as a synonym of C. lapidarium (now C.
cunicularium). I was able to confirm this synonymy by examination of Ausserer's type of C.
orientalis var .mannii. Simon 1892 also recognised Cteniza, Aepycephalus and Cyrtocarenum
as three distinct genera, using differences in the general configuration of the eyes as key
characters (not the relative sizes of the eyes as Ausserer did). Additionally Simon (1892: 93,
figs. 93-94) used the morphology of the rastellum to distinguish Cyrtocarenum from both
Cteniza and Aepycephalus. In preparation for the present study (Decae et al. 1982) I found

that the characters Simon used are highly constant within Cyrtocarenum and therefore are
potentially good diagnostic characters. The morphology of the rastellum in Cyrtocarenum
however, is virtually identical to what I have seen in Cteniza and therefore cannot be used to
distinguish the three genera. On the other hand, from the sparse material of Cteniza and
Aepycephalus that I have seen, the differences in configuration of the eyes between the three
genera seem distinct (Figs. 1-3). Whether these differences should be considered as definite
diagnostic characters at the genus level remains to be investigated. Although the question of
the generic identity of Cyrtocarenum lies largely outside the scope of the study presented
here, I will give a preliminary key for Mediterranean Ctenizinae (sensu Raven 1985). Raven
(1985: 142) questions the separate identity of the three genera by stating that it is only the
poor availability of material that prevents him from proposing the synonymy of Cteniza,
Aepycephalus and Cyrtocarenum. Notwithstanding the differences in eye configuration, I am
inclined to agree, although awaiting a more detailed study of the taxonomy in this group of
spiders I will use Cyrtocarenum for the Ctenizidae (sensu Raven 1985) occurring in Greece 26 .
Measurements, abbreviations and terminology
Measurements of the carapace, sternum and eye-group were performed with the specimen in a
horizontal dorsal or ventral position under the microscope. Measurements of leg and palp
segments were made along the retrolateral surface of detached appendages placed in a
horizontal position as illustrated in Figs. 6-7. A1l measurements were made using a Wild
stereomicroscope equipped with an eyepiece micrometer and are accurate to 0.1 mm.
Abbreviations used (see also Figs. 1-7, 16, 17): REF=width/height ratio of eye formation;
RPT = TL/TW (see Fig. 6) ratio of male palpa1 tibia; RBE = bw/el (see Fig. 6) ratio of palpal
organ; CL = carapace length; CW = carapace width; SL = sternum length; SW = sternum
width; FL = femur length; PL = patella length; TL = tibia length; TW = tibia width; ML =
metatarsus length; TaL = tarsus length; p = pa1p; I = leg I; II = leg II, III = leg III; IV = leg
IV; ax = axis; bc =bursa copulatrix; bw = bulbus width; co = constriction; cv = closure valve;
ef = epigastric furrow; el = embolus length; pl = prolateral; r = receptaculum; rl = retrolateral.
BMNH = Natura1 History Museum, London; FSF = Forschungsinstitut Senckenberg,
Frankfurt; MNHN = Muséum Nationa1 d' Histoire Naturelle, Paris; NHMW =
Naturhistoriches Museum Wien; NNML = Nationaal Natuurhistorisch Museum, Leiden. The
pattern, presence and absence of various types of setae is important in the description and
recognition of the species of Cyrtocarenum. The following terms are used: hook = massive
setiform structure associated with the leg claspers in males (Figs. 22-23); spines = macro
setae that occur on all appendages (Fig. 13); short spines = setiform structures as illustrated in
Fig. 14; spiny setae = conspicuously long strong setae that occur on some 1egs (e.g. on femur
III and IV of the spiders illustrated in Figs. 18-19) and on the cymbium of C. grajum (Fig.
21); setae = hair-1ike cover of most body parts (Fig. 15); teeth = strong and rigid structures in
the rastellum and much smaller structures lining the cheliceral furrow; denticles = knob-like
structures within the cheliceral furrow; cuspules = knob-like structures on maxillae and
labium.
Material examined
The Cteniza and Aepycephalus material examined is all in the collection of the MNHN and
consists of 4 specimens labelled Cteniza sauvagesi, (Rossi 1788), 7 specimens labelled
Cteniza moggridgei O. P. Cambridge 1874, one specimen labelled Aepycephalus brevidens
26 On the basis of new data (Decae 2010) the distinct generic status of Cteniza and Cyrtocarenum is confirmed on the
clasping organs on the male tibia I absent in Ctenzia, present in Cyrtocarenum.

Doleschall 1871 from Sardinia and one undetermined trapdoor spider also from Sardinia that
obviously belongs to the same genus. The Cyrtocarenum material came from the extensive
sample described below. The species-level taxonomy of Cyrtocarenum presented here is
based on a sample of 224 spiders newly collected throughout southern Greece between 1979
and 1994. The sample was found to contain two different species.170 females and 3 males
were preliminarily classified as "species A ", 45 females and 6 males were preliminarily
classified as "species B". Species A was found to be identical to C. cunicularium, species B
was found to be C. grajum. Because the original descriptions of both species were based on
females, one female of each species, newly collected at the respective type locality (Naxos for
C. cunicularium and Argolis for C. grajum), was selected for redescription. Adult males occur
for only a very short period each year. Therefore they are difficult to find in nature and
extremely rare in museum collections. The males described here were collected as juveniles
and reared in captivity. The described male of C. cunicularium originated from the island of
Tinos, that of C. grajum from the island of Kythira. Type material was studied thanks to the
co-operation of the BMNH (C. grajum, C. ionicum, C. tigrinum) and the NHMW ( C.
hellenum, C. lapidarium var .mannii = Cteniza orientalis var. mannii, C werneri). The type of
C. cunicularium could not be located. Further material was made available by the BMNH
(C. cunicularium 4♀ from Corfu and the Ionian Islands, C. grajum l ♀ from an unknown
locality), NHMW (C. cunicularium 5♀ from Kalamos, Tinos and Crete, C. grajum 2♀ from
Kalamos), MNHN (C. cunicularium 3♀ from Crete and Ithea, C. grajum 1♀ from Corfu),
FSF (C. cunicularium 13♀ from Crete and Attica, C. grajum 7♀ from Attica and Skopelos).
The material collected between 1979 and 1989 by myself and Gilbert Caranhac (including all
spiders of both sexes described here) is deposited in the NNML. Rhodos can be included in
the distribution range of C. cunicularium thanks to one specimen collected by Dr C. L.
Deeleman-Reinhold and kept in her collection. Material collected by myself after 1989 is at
present kept in my private collection and will eventually be placed in an institutional
collection. Map 1 shows the geographical distribution of Cyrtocarenum specimens included in
this study.
Figs. 1-3: Eye formations of European Ctenizinae ( dorsal views). REF= ratio describing shape of eye-formation: width
(w)/ height (h), measured as indicated. 1 Cteniza sauvagesi (Rossi 1788), REF=I.9; 2 Aepycephalus sp., REF=2.2; 3
Cyrtocarenum cunicularium (Olivier 1811), REF=3.6.
Procedure and Conclusions
The aim of this study is to clarify the species-level taxonomy of Cyrtocarenum. The first
approach has been to obtain a sufficiently large sample of specimens accompanied by a set of
reliable field data and collected at locations that together form a cross-section of the
distribution range of the genus in Greece (from the Ionian island of Sakynthos to the island of
Rhodos near the Anatolian coast). Within this sample of 215 female and 9 male spiders a

search for diagnostic characters at the species level was carried out. Two species could be
distinguished by the presence or absence (in both sexes) of a double row of trichobothria on
the palpal tibia (Figs. 8-9) combined with the presence or absence of concentrations of spigots
on the lateral spinnerets (Figs.10-11). These two characters were found to be stable
throughout the sample and to correlate fully with a distinctly different morphology of the
spermathecae in the females (Figs.16-17) and the palp, palpal organ and clasper morphology
in the males (Figs.18-23). On this basis 170 females and 3 males were classified as members
of "species-A " (single row of trichobothria on palpal tibia combined with concentrations of
spigots on the spinnerets) and 45 females and 6 males as members of "species-B" (double row
of trichobothria on palpal tibia combined with the absence of concentrations of spigots on the
spinnerets). Comparison of "species-A" and "species-B" with the available type material and
other specimens in museum collections led to the following conclusions:
(1) all type specimens and other specimens in museum collections fitted either
"species-A " or "species-B" and no other morphs (species) were discovered.
(2) the types of C. hellenum, C. ionicum, C. lapidarium var. mannii ( = Cteniza
orientalis var. mannii), C. tigrinum and C. werneri all conform to "species-A";
(3) the type of C. grajum conforms to "species-B".
The holotype of C. cunicularium could not be located. The type locality for this species
however, is the island of Naxos where, as on all islands in the Cyclades archipelago, only
"species-A " was found and "species-B" does not occur. This observation and the fact that
Ausserer's (1871: 157-158) description of C. arianum (= C. cunicularium) clearly indicates
"species-A " (spigot concentrations are mentioned) provides sufficient grounds to regard C.
cunicularium as "species-A " and to designate a neotype (newly collected specimen from
Naxos). The general conclusion therefore must be that the genus Cyrtocarenum in Greece
contains two known species, C. cunicularium ("species-A ") and C. grajum ("species-B").
Severa1 other morphologica1 structures were found to be unique to either C. cunicularium or
C. grajum but restricted to one sex only. The morphology of the spermathecae (Figs.16-17),
the presence or absence of particular spines on the tibia and metatarsus of leg IV (Figs.12-13)
and the pattern of setiform structures dorso-distally on tibia II (Figs.14-15) are good
diagnostic characters in females. The relative length of the palps (Figs.18-19), the
morphology of the palpal organ (Figs. 20-21) and the structure of the clasper on leg I (Figs.
22-23) are diagnostic characters in males. Other characters such as colour variation, relative
lengths of appendages, variations in spine pattern and measurement ratios of different body
parts are valuable in the study of geographical variation within the two species, but are not
further considered here.

Figs.4-7: Methods of measurements. All measurements made with the specimen or
appendage in horizontal position under the microscope with both points of
measurement simultaneously in focus. Legs and palps measured along retrolateral
side after removing them from the spider. 4 Carapace, CL=carapace length,
CW=carapace width; 5 Sternum, SL=sternum length, SW=sternum width; 6 Distal
end of male palp, TL= tibia length, TW=tibia width, el=embolus length, bw=bulbus
width; RPT (TL/TW)=ratio describing shape of male palpal tibia, RBE (bw/el)=ratio
describing shape of palpal organ; 7 Leg segments, FL=femur length, PL=patella
length, TL=tibia length, MT=metatarsus length, TaL=tarsus length.

European Ctenizidae
Raven's (1985) reclassification of mygalomorph spiders of the family Ctenizidae leaves four
representative genera in the Mediterranean: Ummidia Thorell 1875 in Spain and Cteniza,
Aepycephalus and Cyrtocarenum in a more or less disrupted curved zone from the extreme
south-east of France via the islands of Corsica, Sardinia and Sicily into Greece and Anatolia.
Cteniza occupies the north-west part of this area, Aepycephalus the centre and Cyrtocarenum
the south-east. Ummidia is placed in the subfamily Pachylomerinae and will not be further
discussed here. Cteniza, Aepycephalus and Cyrtocarenum represent the European Ctenizinae
(Raven 1985).
Key to the European Ctenizinae
1. REF>2.5 (Fig. 3) ……………………………………….Cyrtocarenum
REF2.5). Description: Females are squat, short-legged spiders that inhabit "cork type",
fully silk-lined trapdoor burrows of various design and complexity; with or without linear
litter (Main 1957), with or without an inverted trapdoor at the bottom of the burrow (Saunders
1842). Colour of sclerotised parts in alcohol varies from maroon to a light yellowish brown
(geographical variation). Abdomen purplish to greyish. Carapace length (CL) of reproducing
females ranges from 6.0 to 11.7. Leg formula: 4132 or 4312 (geographical variation in C.
cunicularium). Males (CL: 5.8 to 8.0) are robust, long-legged spiders. Colour of sclerotised
part in alcohol dark- to light brown. Abdomen greyish. Distal end of tibia I and proximal end
of metatarsus I modified to form a strong clasper (Figs. 22-23).

Figs. 8-15: 8-9 Right palp tibia, dorsal view. 8 C. cunicularium, one row of trichobothria (pl to longitudinal ax); 9 C.
grajum, two rows of trichobothria. 10-11 Spinnerets, ventral view. 10 C. cunicularium, spigots in concentrations on lateral
spinnerets' (arrowed) distal segment domed; 11 C. grajum, spigot concentrations lacking, distal segment digitiform. 12-13
Female, metatarsus IV and distal end oftibia IV,prolateral view. 12 C. cunicularium, note reduced number of spines on
metatarsus (compared with C.grajum, Fig. 13) and vestigial spines (often absent) on tibia; 13 C. grajum, the spine dorsodistally
on metatarsus IV (sp, arrowed) always present in this species but never in C. cunicularium. 14-15 Female, tibia II
dorsodistally. 14 C. cunicularium, field of short spines (s. sp.); 15 C. grajum, ordinary setae (set). T=tibia, M=metatarsus.
Scale lines=1.0mm.
Cyrtocarenum cunicularium (Olivier 1811) (Figs. 3, 8, 10, 12, 14, 16, 18, 20, 22)
Mygale ariana Walckenaer 1805: 6 (n. nud.); Walckenaer 1837: 239; Latreille 1818: 126.
Mygale cunicularia Olivier 1811: 85-86 (= ariana), type not located, from Naxos.
Mygale ionica Saunders 1842: 160, two female syntypes from Ionian Islands, at BMNH (examined).
syn. nov.
Mygalodonta ariana: Simon 1864: 75.
Cteniza tigrina L. Koch 1867: 882, male holotype from Syros, at BMNH (examined).
Cteniza ariana Erber 1868: 905; Moggridge 1873: 131, 135, 141, 143.
Cteniza ionica Kirby 1871: 67; Moggridge 1873: 131, 143; 1874: 210.
Cteniza orientalis Ausserer 1871: 154, var. mannii, three female syntypes from Brussa, at NHMW
(examined); Simon 1892: 96.
Cyrtocephalus hellenus Doleschall (ms) 1852: 26, female holotype at NHMW (examined). syn. nov.
Cyrtocephalus lapidarius Lucas 1853; 514,. type lost, from Crete. syn. nov.
Cyrtocephala lapidaria Simon 1864: 81.
Cyrtauchenius corcyroeus Thorell 1870: 166.

Cyrtauchenius lapidarius Thorell
1870: 165.
Cyrtocarenum arianum Ausserer
1871: 158; Moggridge 1873: 143.
Cyrtocarenum ionicum Ausserer
1871: 161; Moggridge 1873: 131,
143; Pavesi 1877: 327; 1878: 381;
Simon 1880: 115 (= corcyraeum);
1884: 346; 1892: 96; Carlini 1901: 79;
Bristowe 1935: 739; Drensky 1936b:
9; Roewer 1942: 158 (jonicum);
Bonnet 1956: 1351.
Cyrtocarenum lapidarium Ausserer
1871: 161; Pavesi 1876: 68; Simon
1884: 346, 348; 1892: 96; Fage 1921:
99; Caporiacco 1929: 223; Bristowe
1935: 738; Drensky 1936a: 110;
1936b; 9; Hadjissarantos 1940: 19;
Roewer 1942: 158; Bonnet 1956:
1352; Platnick 1993: 84.
Figs. 16-17: Spermathecae (dorsal view). 16 C. cunicularium; 17
C. grajum. Ax = axis line (see text), bc = bursa copulatrix, co =
constriction, cv = central valve, ef = epigastric furrow, r =
receptaculum.
Cyrtocarenum tigrinum Ausserer 1871: 158; Moggridge 1873: 143; Pavesi 1877: 327; 1878: 381.
Cyrtocarenum hellenum Ausserer 1871: 159; Pavesi 1877: 327; 1878: 381; Simon 1892: 96; Bristowe
1935: 739; Drensky 1936b: 9; Roewer 1942: 158; Bonnet 1956: 1351.
Cyrtocarenum cunicularium Pavesi 1877: 327; 1878: 380; Simon 1884:347; 1892: 75, 96; Bristowe
1935: 739; Drensky 1936b: 9; Gerhardt & Kästner 1938: 586; Roewer 1942: 157; Bonnet 1956: 1351;
Buchli 1969: 182; Glatz 1973: 47; Decae, Caranhac & Thomas 1982: 410-419, figs. 1-8; Decae (ms)
1983: 1-59; Coyle 1986a: 294; Decae 1986: 39-43; 1993: 75-82.
Cyrtocarenum werneri Kulczynski 1903: 627,632; Roewer 1942: 158; Bonnet 1956: 1352. syn. nov.
Diagnosis: Trichobothria on dorsal palp tibia in one row (Fig. 8); spigots concentrated in
distinct groups apically on ventral surfaces of median and terminal segments of lateral
spinnerets (Fig.10). Females: CL of reproducing females 6.0-9.5. Spermathecae "mushroom
shaped" and with a thick, annular wall of glandular tissue; lines drawn through central axes of
receptacula converge on ef (Fig. 16); tibia II with a field of short spines dorso-distally (Fig.
14); pl tibia IV without spines or with vestigial spines (Fig.12); pl metatarsus IV with only
one spine ventro-distally (Fig.12); leg formula 4132. Males: CL 6.2-8.0. Palps long and
slender, reaching beyond tibial metatarsal joint of leg I (Fig.18); tibia/metatarsus of leg I
modified to form a "clasper" (Coyle 1986b) with one strong hook distally on rl tibia (Fig. 22);
embolus terminating in a bent pointed tip (Fig. 20); leg formula 4123.
Description: Female, Neotype (no. N 81-10 NNML) col1ected in May 1981 on Naxos (type
locality of C. cunicularium). Measurements; CL=8.3; CW=7.1; SL=5.1; SW=4.1; FLp=4.3;
PLp=2.6; TLp=2.5; TaLp=3.2; FL 1=5.1; PL 1=3.6; TL 1=3.0; ML 1=2.8; TaL 1=1.3; FL
II=4.5; PL II=3.5; TL II=2.6; ML II=2.6; TaL II=1.3; FL III=3.9; PL III=3.6; TL III=2.2; ML
III=3.3; TaL III=2.0; FL IV=5.6; PL IV=4.0; TL IV=3.5; ML IV=4.4; TaL IV=2.2.
Carapace: Brown. Caput steeply arched with concentrations of setae on clypeus and posterior
to eye-formation (Fig. 3). Fovea deep, procurved. Thoracic region glabrous. Eye-formation;
REF=3.6. Eyes on low tubercle marked by dark pigmentation of integument. Median eyes
smaller than laterals, anterior row procurved, posterior row recurved (Fig. 3). Chelicerae;

Basal segment dark reddish brown. Setae concentrated along dorsal margin, merging distally
with rastellum and in a narrow longitudinal zone of thin setae on retrolateral surface.
Promargin of cheliceral furrow with 7 or 8 teeth (distals stronger). Retromargin with scopula
and row of 5 teeth (proximals stronger), small denticles on furrow bottom. Rastellar process
in ventral view triangular with apically two (paired) teeth and one (singular) tooth placed
more retrolaterally (Fig. 5). Maxillae; With distinct distal anterior lobe (Fig. 5). Few cuspules
along proximal margin anterior to labium. Labium; Trapezoidal, wider than high. Small group
of cuspules near anterior margin. Separated from sternum by a wide shallow groove. Sternum;
Flat, widest between coxae II and III. Posterior projection between coxae IV. Two large
central sigilla and two pairs of indistinct sub-marginal sigilla. Setae: All sclerotised parts,
except carapace, more or less evenly covered with black setae. Palps: One row of
trichobothria dorsally on proximal half of tibia, pl to longitudinal ax (Fig. 8). Spination:
patella pl 2-2, tibia pl 3-3 rl 5-5, tarsus pl 9-l0 rl 16-l6. Spines on pl patella and pl tibia in
longitudinal rows, those on rl tibia and both sides of tarsus in longitudinal groups in which
more ventrally placed spines are stronger. Tarsal claw with two teeth, proximal largest.
Leg I: Spination: tibia pl 13-12 rl 7-5, metatarsus pl 12-14 rl 15-l2, tarsus pl 8-8 rl 7-7. Spines
on pl tibia in a longitudinal row, other spines concentrated in longitudinal groups in which
more ventrally placed spines are stronger. Paired claws with one tooth, 3rd claw smooth.
Leg II: Spination: tibia pl 2-2 rl 4-3,metatarsus pl l0-8 rl l0-10, tarsus pl 8-9 rl 4-3. Group of
short spines on dorsal distal tibia (Fig. 14). Setting of spines and claws as leg I. Leg III:
Spination: patella pl 3-3, tibia pl 1-1 rl 4-3, metatarsus pl 6-8, tarsus pl 5-5. Spiny setae
distally on dorsal femur and dorsal patella. Dense concentrations of short spines on d tibia and
metatarsus. Claws as leg I. Thin tarsal scopula present. Leg IV: Spination: metatarsus pl 5-4 v
2-0, tarsus pl 8-8. Concentrations of short spines dorsally on both sides of femur/patella joint;
rl of paired claws largest, with one tooth, pl smaller and smooth, 3 rd claw smooth. Thin tarsal
scopula present. Spinnerets (Fig.10): Laterals ventrally three-segmented (dorsally the basal
segment is diagonally divided giving the spinnerets a four-segmented appearance); basal
segment twice as long as two terminal segments together; apical segment very short and
domed. Spigot concentrations distally on median and apical segments. Median spinnerets
small, one-segmented, with few spigots. Spermathecae (Fig.16): Membranous bursa
copu1atrix (bc) forms a continuous slit in anterior wall of epigastric furrow (et) with, on each
side of longitudinal body axis, a valve (cv) to close off entrance to a receptaculum (r) that
consists of a proximal and a distal part separated by a constriction (co) in the receptaculum
wall. A pronounced thickening in glandular tissue of the distal part of the receptaculum gives
the structure a peculiar mushroom shape.
Male (no.GR80-11, NNML): collected as a juvenile in July 1980 on Tinos and reared in
captivity. Measurements: CL=7.1; CW=6.1; SL=4.0; SW=3.4; FLp=5.7; PLp=3.5; TLp=5.1;
TaLp= 1.3; FL 1=6.9; PL 1=3.6; TL 1=4.1; ML 1=3.8; TaL 1=2.2; FL II=6.1; PL II=3.2; TL
II=3.8; ML II=4.2; TaL II=2.8; FL III=5.0; PL III=3.0; TL III=2.8; ML III=4.7; TaL III=3.0;
FL IV=7.3; PL IV=3.4; TL IV=5.0; ML IV=6.7; TaL IV=3.0. Carapace: Uniformly golden
brown with a sharp line of darker pigmentation along edges of cephalic region. Caput less
steep than in female, with setae concentrations on clypeus and directly posterior to eye
formation. Fovea procurved and deep. Thoracic region glabrous. Eye-formation: REF=3.4.
Eyes on low tubercle marked by black pigmentation of integument. Anterior row slightly
procurved; posterior row recurved. Chelicerae: Slightly darker in colour than carapace,
otherwise as female. Maxillae: As female. Labium: Approximately twice as wide as high,
anteriorly more rounded than in female. Cuspules absent. Separated from sternum by wide but
shallow groove. Sternum and Setae: as female. Palps: RPT=4.6. Long and slender, when
extended reaching beyond tibial-metatarsal joint of leg I (Fig. 18). Trochanter, femur, patella
and tibia conspicuously elongated. One row of trichobothria on dorsal tibia as in female.

Spines absent. Few spiny setae dorso-distally on femur. Cymbium apically bilobed. Embolus
with narrow tip (Fig. 20). Palpal organ, RBE = 0,96. Leg I: Tibialmetatarsal junction modified
and strongly sclerotised to form a "clasper", with one strong hook retrolaterally on enlarged
distal end of tibia (Fig. 22). Spines concentrated on pl and rl patella, and pl and ventral tibia,
Two spines, one pl and one rl, distally on ventral metatarsus, No metatarsal spines associated
with clasper (Fig. 22). Tarsal spines absent. Spiny setae on dorsal and pl faces of femur.
Paired claws with single comb of teeth, 3rd claw absent (or vestigial). Scopula only on tarsus.
Leg II: Concentrations of spines on pl and ventral patella and tibia, and rl metatarsus and
tarsus. One central spine on rl patella and tibia. Two distal spines on pl metatarsus. Strong
spiny setae on dorsal and pl femur. Claws and scopula as leg I. Leg III: Concentrations of
spines and spiny setae on all leg segments. Claws and scopula as leg I. Leg IV: Spines and
spiny setae on all except rl face of femur, on ventro-distal patella, on all faces of tibia and
metatarsus, and on pl tarsus. On rl tarsus spines form small distal group. Paired claws as leg I,
3rd claw present. Tarsal scopula absent. Spinnerets: As female (Fig. 10).
Cyrtocarenum grajum C. L. Koch 1836 (Figs. 9, 11, 13, 15, 17, 19, 21, 23)
Cteniza graja C. L. Koch 1836: 39, female holotype at BMNH (examined); 1851: 71.
Mygalodonta graja: Simon 1864: 75.
Cyrtocarenum grajum: Ausserer 1871: 158; Pavesi 1877: 327; 1878: 381; Simon 1884: 347- 348;
1892: 96; Bristowe 1935: 739; Drensky 1936b: 9; Roewer 1942: 158; Bonnet 1956: 1351; Decae (ms)
1983: 1-59; 1986: 39-43; 1993: 75-82.
Diagnosis: Trichobothria on dorsal palp tibia in two rows (Fig. 9); spigots evenly distributed
over ventral surfaces of median and terminal segments of lateral spinnerets (Fig.11). Females:
CL of reproducing females 6.3-11.7. Spermathecae "bottle shaped", evenly covered with
glandular tissue; lines drawn through central axis of receptacula diverge on epigastric furrow
(Fig.17); tibia II lacks short spines dorso-distally (Fig.15); pl surface of tibia IV with some
well-developed spines (Fig. 13); pl
metatarsus IV distally with both a
ventral and a dorsal spine (Fig.13); leg
formula 4132. Males: CL 5.8-7.1; palp
stout, not elongated, and not reaching
beyond tibialmetatarsal joint of leg I
(Fig. 19); clasper on tibia I with three
strong hooks distally (Fig. 23);
embolus with spatulate tip (Fig. 21);
leg formula 4123.
Description: Female (no. 27/10/92-1,
NNM); collected at Ahladokabos (23
km west of Nauplion along road to
Tripolis), province Argolis (type
locality of C. grajum).
Figs. 18-19: Dorsal habitus, male. 18 C. cunicularium, note elongated,
pa1ps; 19 C. grajum. Spiny setae on dorsa1 femur III and IV (see text).
Scale line= 10.0 mm.

Measurements: CL=8.1;
CW=7.9; SL=6.6; SW=4.8;
FLp=4.8; PLp=2.8; TLp=2.8;
TaLp=3.5; FL I=5.7; PL I=3.8;
TL I=3.5; ML I=3.0; TaL I=
1.4; FL II=4.8; PL II=3.5; TL
II=2.7; ML II=2.9; TaL II=1.5;
FL III=4.4: PL III=3.5: TL III
=2.6: ML III=3.6: TaL III=1.7;
FL IV=6.8; PL IV=3.9; TL
IV=3.7; ML IV=4.8; TaL
IV=2.1. Carapace: maroon.
Caput steeply arched with
concentration of setae around
eye-formation. Fovea deep,
procurved, thoracic region
glabrous. Eye-formation:
REF=3.0, otherwise as C.
cunicularium. Chelicerae: setae
as C. cunicularium. Promargin
of chelicera1 furrow with 8
teeth (distals stronger);
retromargin with scopula and
row of 7 teeth (proximals
stronger); numerous denticles
on furrow bottom. Maxillae:
As C. cunicularium. Labium:
trapezoidal, wider than high.
Cuspules absent. Sternum and
Figs.20-23: 20-21 Cymbium and palpal organ, prolateral view. 20
C. cunicularium, embolus with narrow tip, bulbus width/ embolus
length, RBE=0.96 (see also Fig. 6); 21 C. grajum, embolus with
spatulate tip, RBE= 1.20. Note different orientation of emboli
which may be related to different orientation of spermatheca in
females (Figs. 16-17), and spiny setae on cymbium of C. grajum.
Scale line = 0.5 mm. 22-23 Leg claspers on tibia I and metatarsus I
of male. 22 C. cunicularium, left leg, one hook (arrow); 23 C.
grajum, right leg, three hooks (arrows). Scale line = 1.0mm.
Setae: as C. cunicularium. Palps: two rows of trichobothria (one on either side of longitudinal
central axis) dorsally on proximal half of tibia (Fig. 9). Spination: patella pl 2-1, tibia pl 6-9, rl
9-7, tarsus pl 11-10 rl 20-21. Spine setting as C. cunicularium. Tarsal claw with one tooth.
Leg I: Spination: patella ventral 1-1, tibia ventral l-l pl 4-4 rl 13-13, metatarsus pl 14-l5 rl 20-
18, tarsus pl 7-8 rl 8-8. Spine setting (except for ventral spines on patella and tibia that are
absent in C. cunicularium) as C. cunicularium. Claws as C. cunicularium. Leg II: Spination:
patella ventral 1-l, tibia ventral l-l pl 4-4 rl 8-9, metatarsus pl l3-12 rl 4-4, tarsus pl 5-7 rl 6-6.
Dorsal short spines absent (Fig. 7b). Spine setting and claws as leg I. Leg III: Spination:
patella pl 8-5, tibia pl 3-2, metatarsus ventral 2-2 pl 11-l2 rl 3-4, tarsus pl 6-6. Few spiny setae
distally on dorsal femur. Concentrations of spiny setae on d patella, tibia, metatarsus and
tarsus. Thin scopula on tarsus. Claws as leg I. Leg IV: Spination: tibia pl 6-5, metatarsus pl l6-
16 rl 2-2, tarsus rl 10-8. Concentrations of spiny setae and short spines distally on dorsal
femur and d patella. Scattered spiny setae on dorsal tibia, metatarsus and tarsus. Paired claws
with two teeth on pl claw; rl paired claw and 3rd claw smooth. Spinnerets (Fig.11):
Segmentation of laterals as in C. cunicularium, basal segment slightly longer than two distal
segments together; apical segment digitiform. Spigots more or less evenly distributed over
ventral surfaces of all three segments. Median spinnerets small, one-segmented, with few
spigots. Spermathecae (Fig. 17): Membranous bursa copulatrix (bc) and valve (cv) similar in
structure but larger than in C. cunicularium. Receptacula large, somewhat "bottle-shaped",
with wide proximal part and narrower distal part both evenly covered with glandular tissue.

Male (no. Car. 8/82-1, NNML): collected as a juvenile in August 1982 on Kythira by Gilbert
Caranhac and reared in captivity. Measurements: CL=7.1; CW=6.4; SL=4.2; SW=3.8;
FLp=4.6; PLp=2.2; TLp=3.8; TaLp= 1.9; FL I=7.1; PL I=3.5; TL I=4.1; ML I=6.0; TaL
I=2.9; FL II=6.6; PL II=3.3; TL II=3.8; ML II=5.1; TaL II=2.6; FL III=5.1; PL III=2.6; TL
III=3.2; ML III=4.6; TaL III=2.6; FL IV=7.2; PL IV=3.1; TL IV=4.4; ML IV=6.2; TaL
IV=2.8. Carapace: caput low, few setae around eye-formation. Fovea procurved. Thoracic
region glabrous. Eyeformation: REF=3.3, general features as C. cunicularium. Chelicerae:
setae as C. cunicularium. Promargin of cheliceral furrow with 9 teeth, retromargin with
scopula and row of 8 teeth, numerous denticles on furrow bottom. Few, but conspicuously
long, spiny setae in rastellar area. Maxillae: Cuspules absent, otherwise as C. cunicularium.
Labium and Sternum: as C. cunicularium. Palps: RPT=2.6. Not elongated. Two rows of
trichobothria dorsally on proximal half of tibia as in female. Spiny setae concentrated on
cymbium, embolus with spatulate tip (Fig. 21). Palpal organ, RBE= 1.20. One dorso-distal
spine on tibia. Spiny setae on dorsal and ventral femur, ventro-distal patelIa and ventral tibia.
Leg I: Clasper with three hooks retrodistally on tibia and two ventral spines on metatarsus
(Fig. 23), other spines on ventral patella, rl, ventral and pl tibia (spines on ventral tibia very
strong, fitting description of hooks). No distal spines on metatarsus. Spiny setae on dorsal
femur. Paired claws as C. cunicularium male, 3rd claw present. Scopula extending over
ventral tarsus and distal1/3 of metatarsus. Leg II: Spines concentrated on pl tibia, proximal
metatarsus, v and rl tibia, and rl metatarsus. One ventro-distal spine on rl patella and one
on ventro-distal pl metatarsus. Spiny setae on dorsal and pl femur. Claws and scopula as leg I.
Leg III: As C. cunicularium male. Leg IV: Spines and claws as C, cunicularium male.
Scopula present. Abdomen: lateral and dorsal almost black, ventral brown, cover as C.
cunicularium. Spinnerets: as female (Fig.11).
Discussion and distribution
A common problem in mygalomorph taxonomy is the limited availability of specimens and
good collection data. Obtaining a workable sample of Cyrtocarenum spiders took some time,
but finally yielded interesting and important information on the taxonomic diversity and
biogeography of the genus. Although much work remains to be done, particularly on the
geographical variation in morphology and behaviour, and on the relationship of
Cyrtocarenum to Cteniza and Aepycephalus, I think that much confusion about the specieslevel
taxonomy of the group is clarified here. A more detailed study on the behaviour and
biogeography of Cyrtocarenum is currently in preparation. Preliminary notes on the
distribution following from the study presented here are given below.
Both species, C. cunicularium and C. grajum, occur syntopically on the Ionian islands and on
Kythira and probably in some mainland areas (e.g. Attica). Samples from Sakynthos and
Kythira, collected on the same roadside bank or hill slope, produced members of both species
in approximately equal numbers. On the nearby Peloponnesos however, the two species
exclude each other in most regions. Although one specimen of C. cunicularium was collected
near the town of Gythion (province Laconia), this region and most of the Peloponnesos is
exclusively C. grajum territory. The exception is the north-eastern province of Argolis where
C. grajum is replaced by C. cunicularium (Map 1). Misleading in this respect is the type
locality of C. grajum which is the town of Nauplion in Argolis. Much effort has been invested
in looking for C. grajum in the immediate vicinity of Nauplion with negative results. An
abrupt change in the Cyrtocarenum fauna was found on the slopes of the Parnon mountains 23
km west of Nauplion. Here the boundary between C. cunicularium and C. grajum territory
was found to be very sharp. On the Greek mainland, Cyrtocarenum is currently known only
from Attica. Here, as on the Ionian Islands and Kythira, both species seem to occur

syntopically (sample in the Senckenberg collection from Moni Penteli contains 4 females of
C. cunicularium and 6 females of C. grajum). From Crete only C. cunicularium is reported
with the exception of one specimen of C. grajum in the Senckenberg collection labelled
"Lakkos, Crete". C. grajum is completely absent from the Cyclades, where C. cunicularium is
very common on most islands. All specimens hitherto reported from Anatolia and Rhodos are
C. cunicularium. One spider in the Senckenberg collection from the Sporades island of
Skopelos was found to be C. grajum. Map 1 shows that the distribution of the two
Cyrtocarenum species cannot be readily understood from the present geographical or
climatological configuration. Interspecific competition or predation are apparently not forces
shaping the distribution of these species, given their close cohabitation on the Ionian islands
and Kythira. Because trapdoor spiders in general are an evolutionarily extremely conservative
group, that combine very poor abilities for dispersal with great qualities for survival, a
possible fruitful approach would be to search for correlations between the present distribution
of the two species and the paleogeographic development of this tectonically tumultuous
region of the Mediterranean. Such an approach seems promising in furthering our
understanding of both the evolution of the European Ctenizinae and of the region in which
they occur.

Acknowledgements
I thank Dr P. J. van Helsdingen for his expert advice, provision of facilities and never-ceasing
support and reviews of the earlier drafts of this paper. I preserve the best memories of the
early field trips to the Cyclades with Gilbert Caranhac who collected and donated much of the
material for this study. Dr C. L. Deeleman- Reinhold gave me the opportunity to study
valuable material from her private collection. Dr M. Grasshoff, Dr J. Gruber, Mr M. Hubert
and Mr F. R. Wanless sent type- and other specimens from their respective museum
collections indispensable for this study. Special thanks go to Nollie Hallensleben for her
constant encouragement in the course of this study and her help and assistance in both
fieldwork and the preparation of the manuscript.

Chapter 8
General Conclusions.
Reworked, updated and extended version of an original presentation at the 24 th European
Congress of Arachnology, Bern, Switzerland, August 2008.
Original title: Patterns of Distribution and Diversity in European Mygalomorph Spiders.
Arthur Decae
Research summer 2008
Contributions to Natural History, Bern (in press)
Ummidia algarve Decae 2010 female

Preamble
As stated in the opening paragraphs of this text (Chap 1 page 3), the central goal of this thesis
is to gain insight in the historical build-up of Mediterranean mygalomorph diversity in the
expectation that this will be in aid of a more general understanding of growing faunal
complexity as it has occurred over eons of geological time. In the course of this study the
central aim has been pursued, in more detail, only in four out of thirteen mygalomorph taxa
living in the Mediterranean Region; Nemesia, Iberesia, Ummidia and Cyrtocarenum. With the
results of these studies in mind, the Mediterranean mygalomorph diversity is discussed at a
more general level of genera and families in this concluding chapter. The chapter is a
reworked, updated and extended version of an oral presentation at the 24 th European Congress
of Arachnology, Bern Switzerland August 2008.
Introduction
Thanks to collection efforts of mainly young arachnologists from southern Europe (see
acknowledgements), a coherent picture of the Mediterranean mygalomorph fauna is now
emerging for the first time. The Mediterranean mygalomorph spider fauna as discussed here
includes seven families, thirteen genera and well over a hundred species that have only partly
been formally described (Table 1). Until recently, almost all the knowledge of Mediterranean
Mygalomorphae was based on isolated taxonomic work mainly conducted in the late 19 th and
early 20 th centuries. The problem was that nearly all this work was done as idiosyncratic
descriptions of small haphazardly collected samples, from which it was very difficult to
develop an overall view of the Mediterranean mygalomorph fauna. Recently a number of
larger and more systematically conducted collection programs have been carried out, mainly
in relation to biodiversity assessments and conservation studies. The here presented view on
the European mygalomorph fauna was developed from the results of these studies.
Table 5. List of all mygalomorph spider families and genera currently known to occur in the
Mediterranean Region with an indication of their expected species diversity and local distribution.
Family Genus n. species Distribution
1 Atypidae Atypus 3 Europe + NW Africa
2 Ctenizidae Cteniza >2 Tyrrhenian Region
Ctenizidae Cyrtocarenum >2 Aegean Region
Ctenizidae Ummidia 4 NW Africa + South Iberian Peninsula
3 Cyrtaucheniidae Cyrtauchenius 16 West Mediterranean
4 Hexathelidae Macrothele 2 South Iberian Peninsula + West Crete
5 Idiopidae Idiops >3 Near East
6 Nemesiidae Brachythele >10 NE Mediterranean
Nemesiidae Iberesia >3 Iberian Peninsula + Balearics + NW Africa
Nemesiidae Nemesia >50 Pan-Mediterranean
Nemesiidae Raveniola >3 Anatolia
7 Theraphosidae Chaetopelma 3 East Mediterranean
Theraphosidae Ischnocolus 11 West Mediterranean
7 families 13 genera >113 in total

Material and Methods
In order to evaluate the origin and affinities of the Mediterranean mygalomorph fauna, two
basic questions have to be answered: 1) which species do exist in the Mediterranean and 2)
where are they to be found To answer these questions at, the species level, extensive and
detailed taxonomical revision is necessary for all Mediterranean genera although the
information presented in the forgoing chapters on Nemesia, Iberesia, Ummidia and
Cyrtocarenum may be regarded as promising first steps. Much work remains to be done even
at a basic level of species discovery and description. At the family and genus levels some
insight is given here in the form of a quick identification key to Mediterranean genera and a
set of distribution maps (see below). The necessary data on the distribution and diversity were
collected in a survey of the extant literature, museum collections and large private and
institutional collections from regions all over the area of interest. A total of 1430 locations
were thus far recorded (Table 2) to produce the distribution maps (Figs.1-13). Over 600
specimens of this sample, including representatives of all Mediterranean genera, were
morphologically studied with the aid of a Ceti-Medo 2 binocular microscope in order to
assess diagnostic taxonomical characters necessary for taxonomic revision. The result of this
survey is summarized in the key given below. The collected information on the geographic
origin of all specimens studied was amassed in a Microsoft Access Data Base and analyzed
for distribution and diversity with the aid of DIVA-GIS geographical computer program
(Hijmans et. al. 2005). The resulting distribution maps are shown in Figs.1-13. In order to
study the affinities of the Mediterranean mygalomorph fauna with that of distributions of
related Mygalomorphae the mapped results were compared with biogeographical information
as it is reported in the Word Spider Catalog (Platnick 2010). Inset maps indicate distributions
of related taxa outside the Mediterranean. The hatched regions only indicate the countries for
which the relevant taxa are recorded in the WSC (Platnick 2010).
Results
Key to Mediterranean Mygalomorph Genera
1) Spinneret number six………………………………………………. Atypus
two……………………………………………… Iberesia
four…………………………………………… 2)
2) Claw tufts present, 3 rd claw absent (paired claws present) ……... 3)
Claw tufts absent, 3 rd claw present ……………………..……………... 4)
3) Occular process present, male tibia I unmodified …….………………. Ischnocolus
Occular proces absent, male tibia I male with spur …..………………. Chaetopelma
4) Dense scopula on anterior tarsi and metatarsi …………………….. 5)
Scopulae anterior tarsi and metatarsi absent (or very thin in males only) 6)
5) Occular process absent ............................................................ Cyrtauchenius
Occular process present ……………………………………… 7)
6) Dense spinefields on anterior tarsi and metatarsi, spinnerets short 9)
Dense spinefields absent, PLS very long PMS wide apart
Macrothele
7) PLS short, distal segment domed, PMS close together Nemesia
PLS long, distal segment digitiform, PMS wide apart 8)
8) Males tibia I with distal spur carrying twin hooks Brachythele
Male tibia without spur (only distal spines)
Raveniola
9) Eye-group split, ALE placed separately and far forward Idiops
Eye-group not split, much wider than long, occular process absent, Cyrtocarenum
Eye-group compact roughly rectangular, slight occular proces present 10)
10) Tibia III with saddle depression ……………………………………… Ummidia
Tibia III without saddle ………………………………………. Cteniza

Table 6. Numbers of recorded geographical locations per genus.
Genus records
1 Atypus 534
2 Brachythele 53
3 Chaetopelma 47
4 Cteniza 19
5 Cyrtauchenius 71
6 Cyrtocarenum 106
7 Iberesia 50
8 Idiops 4
9 Ischnocolus 31
10 Macrothele 68
11 Nemesia 406
12 Raveniola 5
13 Ummidia 36
Atypus
The genus Atypus, probably thanks to its capacity for aerial dispersal, has a broad distribution
range in Europe being distributed all over the continent with exception of the northern most
parts (Fig. 1). Its known Mediterranean distribution is restricted to those regions that have
their origin as parts of the Neoeuropean Archipelago as described in Chapter 1. Atypus has
three species in Europe; all three are found in the Mediterranean climate zone and distributed
in a broadly East-West running pattern (A. muralis in eastern Europe, A. piceus in Central
Europe and A. affinis in western Europe and the Maghreb). The distribution of all three
species reaches far into the latest Pleistocene permafrost coverage of the continent (Hewitt
1999) and can therefore be best explained as the result of a relatively recent Holocene
dispersal event. Atypus’ current European distribution has been suggested to be the result of
Pleistocene survival in Mediterranean and SW Asian refuge populations. These might have
been located in the Iberian Peninsula and/or Maghreb for A. affinis, the Southern Balkan for
A. piceus and a region near the Caspian Sea for A. muralis respectively (Schwendinger 1990).
The low species diversity (three species only) indicates that Pleistocene refuge populations
have not been sufficiently fragmented and isolated for separate Atypus populations to speciate,
or that speciation in Atypus is too slow for the duration of Pleistocene cycles to have effect at
this level of diversity growth. Population level research using DNA-identification techniques
as currently in progress at the University of Barcelona (Arnedo pers. comm.) is expected to
throw new light of the level of homogeneity of European Atypus species. Outside Europe,
Atypus is widely distributed in the Palaearctic (Fig. 1 inset) having its highest diversity in
eastern Asia (25 species described, Platnick 2010). The one Atypus species known from the
eastern USA (A. snetsingeri Sarno 1973) is difficult to explain on biogeographical grounds. If
the placing of this species in Atypus is correct it might indicate that the genus is older than the
Atlantic Ocean (some 170 million years). The only presence of Atypus in Neogondwanan
territory, A. sutherlandi from India, might be explained as the result of a more recent dispersal
event.
Nemesiidae
The Nemesiidae with four distinct genera, two subgenera and around 70 described species is
the most diverse mygalomorph spider family in the Mediterranean. All four Mediterranean

genera are classified in the subfamily Nemesiinae (Raven 1985) and they represent the
western most taxa of a more or less continuous Palaearctic longitudinal distribution range
running from the Pacific in the East to the Atlantic in the West (see Chapter 5). They are
absent from the northern permafrost affected regions of Eurasia indicating a low dispersal
capacity when compared with Atypus. The Mediterranean diversity at the genus level is fully
restricted to regions that originated in the Neoeuropean Archipelago (Figs. 2-5) and only the
genus Nemesia has one species (N. cellicola Audouin 1826) on the African continental plate.
The general Palaearctic longitudinal diversity pattern of Nemesiinae is continued within the
Mediterranean Region with Raveniola in eastern Anatolia (Fig. 2), Brachythele in
southeastern Europe (Fig. 3) and Iberesia in the western, Atlantic regions (Fig. 4). Only
Nemesia has a much broader Mediterranean distribution running from the Atlantic coast to the
eastern Mediterranean. Nemesia has a very different pattern of diversity in the western and
eastern parts of its distribution range that reflects palaeogeographic configurations in its
present distribution rather than the present geography (Chapter 5). A strong positive effect on
building Nemesia diversity should therefore be attributed to situations of geographical
fragmentation and isolation during the millions of years of existence of the Neoeuropean
Archipelago. The idea that Nemesiidae in Europe have been repeatedly ‘pushed back’ into
small fragmented and isolated refuge populations in a series of Pleistocene glaciations, and
that these ongoing cycles of fragmentation and isolation have lasted sufficiently long for
further diversification and speciation cannot be ruled out however. Thorough species level
revision of all nemesiid genera is needed to clarify these propositions.
Cyrtauchenius
Raven 1985 grouped the genera Cyrtauchenius and Homostola in an all African subfamily
Cyrtaucheniinae, with a disjunct distribution to the North and South of the tropics (Fig. 6
inset). Tropical African cyrtaucheniids are placed in another subfamily, Aporoptychinae,
together with the Australian and American taxa. The Asian genus Anemesia (WSC Platnick
2010) is probably wrongly placed in the Cyrtaucheniidae and should be moved to Nemesiidae
(probably genus Raveniola). The grouping with Homostola of Cyrtauchenius suggests an
African origin of the Mediterranean cyrtaucheniids. From biogeography this seems difficult to
explain and from this perspective the Cyrtaucheniinae are expected to be paraphyletic. On the
other hand however the Mediterranean cyrtaucheniids have clearly southern affinities,
although the comparatively high species diversity seems to be related to a long history of
fragmentation and isolation of populations in the western Neoeuropean Archipelago. A
problem with the interpretation of the currently known distribution of Cyrtaucheniinae is the
complete absence of information from sub-Saharan North Africa and from southwestern
Africa. Information on the mygalomorph fauna of the Sahel region is expected to aid
importantly to the information presented here.
Ummidia
The Mediterranean Ummidia population is revised in Chapter 6. It was found to contain at
least four species all distributed in the Iberian-Maghreb region (Fig. 7). Traditionally
Ummidia is grouped with Conothele in the subfamily Ummidiinae (formerly Pachylomerinae)
with a remarkably wide distribution in the Northern Hemisphere and the Indo-Australian
Region. In the revision presented in Chapter 6 Ummidia and Conothele are placed in
synonymy in which the name Ummidia has priority. Ummidia therefore has an almost
worldwide distribution, mainly although not entirely north of the equator. The East Asian
genus Latouchia probably is the closest living relation of Ummidia (personal observation).
The Mediterranean Ummidia population appears to be a remnant of a much wider European
distribution as is evidenced by the finding of fossil Ummidia specimens in Baltic amber

(Wunderlich 2004). The findings of U. algerianus in the southwestern Algerian Sahara (Fig.7)
might indicate the possible existence of Ummidia in sub-Saharan North Africa, but no data are
available. The western Mediterranean Ummidia diversity (four described species) probably
relates to local speciation in the western Neoeuropean Archipelago. This hypothesis would
gain supported if indications that the Ummidia populations of the Spanish Andalusia and
Valencia Regions differ at the species level could be confirmed.
Cteniza & Cyrtocarenum
The genus Cteniza has a very narrow distribution in the northwestern Tyrrhenian region only
(Fig. 8). Taxonomically Cteniza is placed in the subfamily Ctenizinae together with its
putative sister-genus Cyrtocarenum that occurs in the Aegean region only (Fig. 9). It is very
difficult, on basis of present knowledge, to find the closest relatives of these two
Mediterranean genera. The homology of the family Ctenizidae has never been established
(Raven 1985, Goloboff 1993) and a revision of Raven’s Domiothelina (Actinopidae +
Ctenizidae + Idiopidae + Migidae) is likely to reveal new relationships. On arguments of
biogeography the Ctenizidae seem to be divided in southern and northern taxa. The South
African ctenizid genus Stasimopus might be more closely related to the Gondwana families
(Actinopidae, Migidae, Idiopidae). The closest relative of the Mediterranean Ctenizidae might
be found either in California (Bothriocyrtum or Hebestatis) or in the eastern USA and eastern
Asia (Cyclocosmia). All Ctenizidae appear to be concentrated in relic populations and both
Cteniza and Cyrtocarenum confirm this observation. The known diversity in Cteniza and
Cyrtocarenum is low (2 species each), but more species might await discovery. Whatever
their origin and relationships both genera contribute to the Mediterranean biodiversity as
inhabitants of the Neoeuropean Archipelago and as such seem to support the thesis that it is
particularly this geological region that has contributed to the Mediterranean mygalomorph
diversity.
Ischnocolus & Chaetopelma
The genera Ischnocolus and Chaetopelma are the members of the family Theraphosidae in the
Mediterranean Region. The family Theraphosidae with 935 recorded species (approx. 35% of
all mygalomorph spiders) has the highest species diversity of all mygalomorpf spider families.
This might partly be due to the fact that all the very large and spectacular bird-spiders are
members of this family, and partly because some Theraphosidae live in more exposed
situations than most other mygalomorph spiders. Notwithstanding their large size and
complex morphology the taxonomy of Theraphosidae is particularly complex and unresolved.
The two Mediterranean genera are generally placed in the subfamily Ischnocolinae. In
Raven’s 1985 revision of Mygalomorphae all member genera of this subfamily are regarded
as Theraphosidae incertae sedis. It is therefore difficult to indicate the close relatives of
Ischnocolus and Chaetopelma living outside the Mediterranean. What is clear however is that
both genera have African affinities and are probably of African (Afro-Arabian for
Chaetopelma) origin. A conspicuous difference between the two genera is their species
diversity. A recent revision of Chaetopelma (Guandanucci & Galton 2008) shows that the
species diversity in this genus is low (three or four species) and the distribution range,
extending all over the Middle East and as far south as the Sudan, is comparatively large.
Chaetopelma is an inhabitant of the southeastern Mediterranean (Fig. 10) where the passive
Afro-Arabian continental front has not been fragmented over geological time. Recently
Chaetopelma has also been reported from the northeastern Mediterranean and Crete, maybe as
a result of man aided introductions. Ischnocolus inhabits the southwestern Mediterranean
(Fig. 11), an area that has seen extensive fragmentation and island forming throughout the
Tertiary period and this might explain the higher species diversity in Ischocolus (Table 1).

Knowledge of possible Saharan and sub-Saharan populations of these Theraphosidae incertae
sedis is badly missing.
Macrothele
The Hexathelidae are represented in Europe with one genus, Macrothele Ausserer 1871 and
just two species. The world distribution of Macrothele shows some peculiar disjunctions. The
centre of diversity for this genus, with 20 recognized species, is in SE Asia. A second area of
distribution with four recorded species exists in W. Africa. The third are of distribution is in
S. Europe where two isolated species occur; one on the Iberian Peninsula and one on Western
Crete. Very recently the possible existence of a third species was discovered in southern
Anatolia (Kunt pers. comm.). It is not clear however if this represents an authentic new
population or if Macrothele is here transported by man as it is to several other locations in
Europe (Fig. 12). It is very difficult to explain the Mediterranean distribution of Macrothele in
terms of geological, geographical, historical or biological arguments as used above. A recent
molecular study of Macrothele calpeiana (Arnedo & Ferràndez 2006) has indicated that this
species is a very old Iberian endemic. The peculiar distribution of two isolated Macrothele
species in Mediterranean coastal areas however, rather seems to suggest a recent man aided
import of the genus into the European fauna. This idea is further supported by findings of M.
calpeiana far away from the Iberian Peninsula in northern Italy (Pantini & Isaia in press) and
in Belgium (personal observation) and Switzerland (Jemenez pers. comm.). These findings
show the likelihood of M. calpeiana to be transported by man, possibly as an unintentional
‘stowaway’ with the export of garden material or plants such as ornamental olive trees (as is
reported for the Italian records, Pantini pers. comm.). Another fact that might explain recent
outlaying records of M. calpeiana is that the species has attracted some general publicity as
the only formally protected spider species in Europe. This might have encouraged intentional
transport by collectors of illegal animal species. In general however, the question of
Macrothele being an old endemic element in the European mygalomorph fauna or a recent
human aided introduction awaits further investigation. If the Mediterranean Macrothele
populations are indeed endemic (see Arnedo & Ferràndez 2006) it needs further research to
discover if their closest relatives are to be found among the African or Asian Hexathelidae.
Idiops
The genus Idiops seems to be the southern hemisphere counterpart of Ummidia. It also has a
virtually cosmopolitan distribution but almost exclusively on southern continents (Fig. 13
inset). In the Mediterranean it occurs only in the east as an inhabitant of the Afro-Arabian
continental plate. There must be a sharp borderline that might coincide with the Turkish-
Syrian frontier that marks the northern limit of the Idiops distribution in the Mediterranean
Region. Nothing is currently known of the species diversity of Mediterranean Idiops although
there is some indication that at least three different species inhabit the area (personal
observations). This might be just an extension of the high species diversity (58 species
known) Idiops has on the African continent. The wide Gondwanan distribution (S. America,
Africa, India) of Idiops indicates that the genus is at least 150 million years old.
Conclusions
From reading the distribution maps of all 13 genera composing the Mediterranean
mygalomorph spider fauna it is clear that fauna elements of Africa, Asia and Europe all have
contributed to the species diversity of the region. The Nemesiidae and the Atypidae are likely
to be Asian in origin or have at least strong affinities with the Asian mygalomorph fauna. A
remarkable difference between Atypidea and Nemesiinae is the very high diversity in the last
taxon versus a moderate diversity in Atypidae. The difference may partly be explained from

differences in powers for dispersal (Atypus has aerial dispersal, Nemesiinae not) and hence in
possibilities for maintaining species cohesion over larger or smaller ranges of distribution.
Another possible explanation may result from the somewhat more southerly homeland of
Nemesiinae in the geologically fragmented collision front of southern Eurasia. Increased
collection efforts in Asia along the northern front of the Himalayas, between the Caucasus and
the Pacific Ocean, is expected to yield a wealth of unknown Nemesiinae species.
Macrothele might also be an Asian contribution to the Mediterranean fauna but it could also
be African in origin. DNA studies may solve this question of the whereabouts of the closest
relatives of Mediterranean Macrothele.
Cyrtauchenius, Ischnocolus, Chaetopelma and Idiops all appear to be African contributions to
the Mediterranean mygalomorph fauna. The two eastern genera Chaetopelma and Idiops seem
not to have responded to the Mediterranean environment with increased speciation. The
western genera Ischnocolus and Cyrtauchenius in contrast seem to show extensive speciation
in the geologically fragmented western Mediterranean (compare species numbers in Table 1).
The Ctenizinae (Cteniza and Cyrtocarenum) appear to be European contributions to the
Mediteranean fauna. They seem to represent small, but very old relic populations and in that
respect should be regarded as particularly important targets for conservational attention in the
region. Notwithstanding their homeland in the Neoeuropean pre-Alpine archipelago they
show low species diversity, but local geographical variation (e.g. between Cteniza populations
on Corsica and Sardinia or Cyrtocarenum populations on Crete and the Cyclades , personal
observations) indicate much cryptic diversity. Finally the presence of Ummidia in the
southwestern Mediterranean seems also a relic, although of a genus that is widely distributed
in North America, East Asia and the Australian Region. If the closest relatives of the
Mediterranean Ummidia population are to be found in America or in Asia remains to be
investigated.
In general, it seems that the highest species diversity exists along the tectonic collision fronts,
particularly the fragmented southern fringes of the northern continents (Europe and Asia).
Incidentally these are also the geographical zones in which refuge populations might have
survived and speciated during Pleistocene glacial oscillations. However, the apparently
conservative mode of evolution in mygalomorph spiders seems to be reflected in their
distributions predominantly following palaeogeographic archipelagos and coastal fronts. The
recent findings of Chaetopelma in Greece and of Macrothele in several locations in Europe
and Anatolia might indicate the first human induced disturbances of the natural distribution
patterns of Mediterranean mygalomorph spiders. In a recent study Jiminez-Valverde et.al
(submitted) estimates the potential range extensions of human aided introductions of M.
calpeiana in Europe.
Acknowledgements
This study would not have been possible without the cooperation of Rop Bosmans, Gilbert
Caranhac, Pedro Cardoso, Maria Chatzaki, Alberto Chiarle, Marco Colombo, Herman De
Koninck, Francesca di Franco, Fulvio Gasparo, Siegfried Huber, Marco Isaia, Peter Jäger,
Rudy Jocqué, Luka Katusic, Jiri Kral, Kadir Boğaç Kunt, Iris Musli, Wolfgang Nentwig, Vera
Opatová, Paolo Pantini, Johan van Keer and Ersen Aydin Yağmur who all provided
specimens, sometimes in large collections, and/or basic information for this study. My thanks
also go to the several people who have sent, shown or given me individual specimen for
identification, and to the organizing committee of the 24 th European Congress of Arachnology
for giving me the opportunity to present the data here published. I thank Jean-Pierre Maelfait
for providing the necessary facilities for this study.

Summary
All chapters contained in this thesis discuss aspects of the Mediterranean mygalomorph spider
fauna. Chapter 1 outlines the motivation for writing the thesis and the aims pursued. The
thesis is focused on finding historical explanations for the growth of biodiversity in the
Mediterranean Region, lately recognized as one of the biodiversity hotspots in the world.
Because of the historical perspective of the thesis, the formation of the Mediterranean as a
distinct geographical region is discussed in terms of tectonic events. This discussion starts
with the breaking-up of the super-continent Pangaea in early Mesozoic times. It follows the
formation of a fragmented archipelago along the southern front of the European continent
(here called the Neoeuropean Archipelago) in which mygalomorph diversity developed
during in Tertiary times. The different events that controlled the formation of the eastern
Mediterranean (mainly stretching and subduction) and the western Mediterranean (mainly
compression and orogenesis) in the Neogene (see Chapter 1 Fig. 4) still seem to be reflected
in the composition of the mygalomorph faunae of the respective geographical zones. Having
been formed at the crossroads of Africa, Arabia, Asia and Europe the Mediterranean
mygalomorph fauna shows influences from all these continental regions and this partly
explains the remarkably rich biodiversity presently observed. Because the Mediterranean has
retained a beneficial climate during Pleistocene glaciations it has probably served as a refuge
zone for northern species, and part of the recent biodiversity may have originated in isolated
relic populations. It is not clear however if this phenomenon has attributed to the diversity of
mygalomorph spiders in the region and this possibility remains to be investigated.
In the arachnological section of the general introduction a theory is discussed that has been
proposed in a slightly different context by Decae (1984). It is stated that the ancestral lifestyle
for all spiders has been the construction of underground burrows, and that all mygalomorph
spiders and all liphistiomorph spiders are morphologically adapted to a tunneling way of life.
From this observation an alternative phylogeny at the level of taxonomical Orders is
proposed. It is argued here that a basic separation of spiders in Orthognatha and Labidognatha
is more realistic than the conventional Order level classification in Mesothelae and
Opisthothelae. The classification here proposed is based on functional synapomorphies in the
chelicerae and morphology of the hind legs in primitive spiders, with additional arguments on
the function of the pedicel and the positioning of the spinnerets. The evolution of spiders as
seen from this perspective results in envisioning two very successful waves of adaptive
radiation; the first one in two-dimensional space producing the Orthognatha., the second one
in three-dimensional space producing the Labidognatha.
Chapter 2 provides the descriptions of six new species in the genus Nemesia found on
Majorca and Ibiza. The descriptions are accompanied by field observations on apparent
habitat preferences of the different species and on behavior. Arguments for the proposition of
an alternative higher classification of spider (as outlined above) are partly based on
observations of burrow construction behavior in all these species. In Chapter 3 the attention is
focused on the Portuguese Nemesia fauna. The revision of the Portuguese Nemesia fauna led
to the description of two new species and the abolishment of one species. The revision also
led to the identification of a new genus (Iberesia) and a new species that was split-off from
traditional Nemesia stock. Iberesia is characterized by the total absence of posterior median
spinnerets (always present in Nemesia) and by a number of several different species (of which
only three have so far been scientifically described) that are widely distributed in the western
Mediterranean. In Chapter 4 the description of the new genus Iberesia and its new type
species I. machadoi is formalized. In this chapter the diagnostic characters that distinguish the
diverse Mediterranean nemesiid genera, Iberesia, Nemesia and Brachythele are discussed in
detail. Chapter 5 takes a cladistic approach to survey the sub-generic diversity in Nemesia

over its full Mediterranean distribution range. This led to the recognition of two sub-genera
(Holonemesia and Pronemesia) that, in their distributions, are considered to reflect different
centers of origin in the western and eastern Mediterranean. These different centers of
evolutionary diversity might conform to palaeogeographic configurations within the
Neoeuropean Archipelago. The palaeogeographic patterns appear also evident in further
subdivisions, in which cladistically related species groups in both subgenera are recognized.
In general the nemesiid fauna of the western Mediterranean appears to differ qualitatively
from that of the eastern Mediterranean. Also, at a sub-generic level, the Nemesia fauna of the
western Mediterranean exhibits particularly high levels of species diversity.
In Chapter 6 the attention is shifted to the genus Ummidia. This genus has long be regarded a
human aided import from the Americas. The here presented revision shows that Ummidia
probably is endemic to the western Mediterranean and that the species diversity is likely to be
higher than currently known. Four Ummidia species have here been described or redescribed
of which one (U. Algarve) is new to science.
In Chapter 7 the genus Cyrtocarenum is revised. This genus was formerly regarded to contain
seven different species within the Aegean Region alone. Revision of the genus showed the
presence of only two species and five previously recognized species were all found to be
synonyms to C. cunicularium. Geographical variation within C. cunicularium is evident in the
macro-morphology of populations from different parts of the Greek archipelagos. New
species of Cyrtocarenum may still be discovered in the northern parts of Greece, on the
Balkans or in Anatolia. Nevertheless, Cyrtocarenum is an important local endemic of the
Mediterranean mygalomorph fauna.
Chapter 8 presents a collection of the conclusions to be drawn from the above research efforts
and from reading the distribution maps of genera not here revised. The general conclusion is
that all four continental regions that border the Mediterranean have contributed importantly to
the building of its biodiversity, in which the fragmentation of the Tertiary Neoeuropaen
Archipelago seems to have played a major role. Recent records on the distributional
extensions of Macrothele calpeiana and Chaetopelma olivaceum, the two largest, least
secretive and most spectacular mygalomorph spider species, seem to indicate the first human
induced faunal disturbance within the Mediterranean mygalomorph spider fauna.
Samenvatting
Alle hoofdstukken in dit proefschrift gaan over de mygalomorfe spinnen fauna van het
Middellandse Zeegebied. Hoofdstuk 1 bevat de motivatie voor het schrijven van dit
proefschrift en een uitleg van de beoogde doelstelling. Het proefschrift is gericht op het
vinden van historische verklaringen voor de ontwikkeling van de bijzonder rijke biodiversiteit
in het Middellandse Zeegebied die heeft geleid tot de bijzondere status van ‘biodiversity
hotspot’ van het gebied. Vanuit de historische benadering van het onderwerp wordt een
uiteenzetting gegeven over het ontstaan van de Middellandse Zee in termen van tektonische
ontwikkelingen die beginnen met het uiteenvallen van het supercontinent Pangea in het
vroege Mesozoïcum. De tekst beschrijft het ontstaan van een gefragmenteerde archipel voor
de zuidkust van het Europese continent (hier aangeduid als Neoeuropa) waarin de diversiteit
van mygalomorfe spinnen zich kon ontwikkelen in de loop van het Tertiair. Vanuit geologisch
perspectief hebben de oostelijke en de westelijk Middellandse Zee zich heel verschillend
ontwikkeld en de gevolgen daarvan lijken tot op de dag van vandaag zichtbaar in de
verspreiding van de verschillende groepen mygalomorfe spinnen. Omdat de Middellandse Zee
is ontstaan op een plaats waar Arabië, Afrika, Azië en Europa elkaar raken zijn er invloeden
van al die continenten zichtbaar in de fauna van de Middellandse Zee waardoor de bijzondere
rijkdom aan soorten gedeeltelijk wordt verklaard. Omdat de Middellandse Zee bovendien een
leefbaar klimaat behield tijdens de grote ijstijden hebben veel noordelijke soorten planten en

dieren er een onderkomen gevonden waarin ze konden overleven. De scheiding van die
onderkomens op lokaal gunstige plaatsen kan de soortsvorming verder in de hand hebben
gewerkt waardoor de fauna van het Middellandse Zeegebied mogelijk extra is verrijkt. Het
staat overigens te bezien of de isolatie van de ijstijd ook heeft kunnen bijdragen aan de
ontwikkeling van mygalomorfe spinnensoorten.
In de arachnologische sectie van de algemene inleiding wordt voorts een theorie besproken
die door Decae (1984) naar voren is gebracht. Die theorie stelt dat spinnen van huis uit
gravende dieren zijn zoals te zien aan de bouw van alle primitieve spinnen (mygalomorfe
spinnen en liphistiomorfe spinnen). De realisatie dat spinnen van afkomst gravende dieren
zijn heeft consequenties voor de taxonomische indeling van spinnen zoals die momenteel
wordt gezien. De huidige taxonomie deelt de spinnen basaal in twee groepen in op basis van
de anatomische locatie van de spintepels (Mesothelae hebben de spintepels ongeveer
halverwege de onderkant van het achterlijf en Opisthothelae hebben de spintepels aan de
achterste punt van het achterlijf (zie schema’s op voorblad hoofdstuk 1). In de functionele
classificatie die hier naar voren gebracht wordt is het beter de speciaal aan graafgedrag
aangepaste kaken (cheliceren) en de speciaal aan graven aangepaste achterpoten te gebruiken
als eerste classificatiekenmerk. In dat geval is het te prefereren om alle primitieve spinnen te
groeperen in de Orthognatha en alle ‘echte spinnen’ te classificeren als Labidognatha (zie
schema’s op voorblad hoofdstuk 1). Deze indeling visualiseert bovendien de evolutie van
spinnen als het gevolg van twee succesvolle adaptieve radiatie golven die het bestaan van de
huidige spinnenfauna (met primitieve spinnen naast ‘geavanceerde’ spinnen) in een
aannemelijk perspectief plaatst.
Hoofdstuk 2 geeft de beschrijvingen van zes nieuwe soorten die op Majorca en Ibiza
gevonden werden. Naast de taxonomische beschrijvingen worden veldobservaties
gerapporteerd waarbij zowel ecologische gegevens als gedragsgegevens worden
gepresenteerd. De observaties van het graafgedrag van al deze nieuwe soorten heeft de
formulering van de hierboven uiteengezette theorie over primitieve spinnen versus
geavanceerde spinnen sterk positief beïnvloed. In hoofdstuk 3 wordt de Portugese Nemesia
fauna behandeld. Hier zijn twee niet eerder bekende soorten beschreven en is één soort
opgeheven. De revisie van de Portugese Nemesia fauna gaf voorts aanleiding tot het afsplitsen
van een nieuw geslacht (Iberesia) van de Nemesia stamboom op grond van het geheel
ontbreken van de achterste middelste spintepels die bij alle Nemesia soorten wel aanwezig
zijn. Iberesia blijkt een geslacht met verschillende soorten die allemaal voorkomen in het
uiterste westen van het Middellandse Zeegebied. Hoofdstuk 4 beschrijft het nieuwe geslacht
Iberesia en de nieuw ontdekte typesoort I. machadoi. Voorts worden in hoofdstuk 4 de
diagnostische verschillen tussen de genera Iberesia, Nemesia en Brachythele (allen familie
Nemesiidae) besproken en geïllustreerd. Hoofdstuk 6 geeft een cladistische analyse van het
geslacht Nemesia in het gehele verspreidingsgebied van de Middellandse Zee. Op grond van
de resultaten worden binnen Nemesia twee sub-genera onderscheiden (Holonemesia en
Pronemesia). De verspreiding van deze twee sub-genera vertoont markante verschillen die
duiden op verschillende oorsprongsgebieden in de oostelijke en westelijke Middellandse Zee.
Ook binnen de twee sub-genera zijn aparte groepen van soorten te onderscheiden die in hun
verspreiding de locaties van oude centra van Nemesia evolutie indiceren. Hoofdstuk 6
bespreekt de Mediterrane populatie van het geslacht Ummidia waarvan lange tijd werd
geloofd dat het een met planten ingevoerd Amerikaans geslacht zou zijn. De hier
gepresenteerde analyses tonen aan dat Ummidia waarschijnlijk een oud, endemisch, west
Mediterraan geslacht is waarvan tot op heden vier verschillende soorten bekend zijn. Eén van
die soorten U. algarve is in hoofdstuk 6 voor het eerst wetenschappelijk beschreven.
Hoofdstuk 7 beschrijft de revisie van het geslacht Cyrtocarenum dat in het Aegeisch gebied
voorkomt. Aanvankelijk waren in dit geslacht zeven soorten beschreven. De hier

gepresenteerde revisie brengt dat aantal tot twee soorten terug. Geografische variatie binnen
één van deze soorten (C. cunicularium) laat echter zien dat er mogelijk evolutionaire
ontwikkelingen gaande zijn waarbij verschillende eilandpopulaties grote verschillen vertonen.
Het is bovendien mogelijk dat er nog nieuwe soorten binnen het geslacht Cyrtocarenum te
ontdekken zijn met de beste kansen in de zuidelijke Balkan en in Anatolië.
Hoofdstuk 8 tenslotte geeft een verzameling conclusies die uit de bovenstaande onderzoeken
naar voren komen. Het interpreteren van de verspreidingskaartjes van alle Mediterrane
mygalomorfe spinnen geslachten laat zien dat alle continentale gebieden die aan het
Middellandse Zeegebied grenzen een bijdrage hebben geleverd aan het verrijken van de
Mediterrane spinnenfauna, waarbij de gefragmenteerde zuidkust van Europa het belangrijkste
aandeel heeft gehad. Nieuwe registraties van het voorkomen van twee soorten, Macrothele
calpeiana en Chaetopelma olivaceum (de twee grootse, meest opvallende en spectaculaire
spinnen soorten in het gebied) wijzen op de eerste tekenen van locale faunavervalsing door
mensen.

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Epilogue
Around the turn of the century Konrad Thaler (2000) published a paper discussing the
Mediterranean spider fauna and the work of Paolo Brignoli. He concluded that,
notwithstanding the enormous productivity of Brignoli and the fact that his research was
largely focused on the Mediterranean our current knowledge of Mediterranean spiders still is
very poor. This is particularly true for Mediterranean mygalomorph siders he noted. In this
thesis ten new mygalomorph spider species and one new genus of Mediterranean
mygalomorph spiders are named and described. More important however, the revisionary and
biogeographical work, presented here, indicates that this is only a small fragment of the taxa
yet to be discovered in this region. Large parts of the Mediterranean, including most of
Morocco, Libya, Egypt, Israel, Syria, Turkey and the Balkan States are hardly known at all
for their mygalomorph spider faunae and other regions such as Greece, Italy and Spain are
only partly studied. Even knowledge on the mygalomorph spider fauna of southern France,
where trapdoor spiders were already studied in the late 18 th century, is badly confused.
In a time when scientists are trying to track the limits of the Universe, spend fortunes chasing
subatomic particles and made giant investments to decode the human genome it is almost
cynical to note how little we know about the creatures that live among us. Since knowledge of
our world is a direct measure for the level of human culture, much cultural progress is still to
be made by studying the smaller creatures around us.