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Mitt. Mus. Nat.kd. Berl., Zool. Reihe 81 (2005) 2, 145–166 / DOI 10.1002/mmnz.200510009 

Aplacophoran Mollusca in the Natural History Museum Berlin. 

An annotated catalogue of Thiele’s type specimens, with a brief review 

of “Aplacophora” classification 

Matthias Glaubrecht*, 1 , Lothar Maitas 1 & Luitfried v. Salvini-Plawen**, 2 

1 Department of Malacozoology, Museum of Natural History, Humboldt University, Invalidenstraße 43, D-10115 Berlin, 

Germany 

2 Institut für Zoologie, Universität Wien, Althanstraße 14, A-1090 Vienna, Austria 

Received January 2005, accepted April 2005 

Published online 08. 09. 2005 

With 2 figures 

Key words: Systematization, cladistic analyses, Solenogastres (¼ Neomeniomorpha), Caudofoveata (¼ Chaetodermomorpha), 

Aculifera, Amphineura, Johannes Thiele, Ernst Vanhöffen, First German South Polar Expedition, “Gauss”, “Valdivia”. 

Abstract 

Aplacophoran molluscs are a small, often neglected and still poorly known but phylogenetically important basal group, with 

taxa possessing morphological characters considered essential for the reconstruction of the basal Mollusca and their evolution. 

Currently, in most textbooks of zoology and major malacological treatise Solenogastres and Caudofoveata are viewed as constituting 

a monophyletic clade called Aplacophora Von Ihering, 1876, although evidence is available to the contrary, suggesting 

the latter to be a paraphyletic grade. Accordingly, the hitherto accepted “Aplacophora” may consist of two Recent, diphyletic 

taxa, viz. Solenogastres Gegenbaur, 1878 (sensu Simroth, 1893) or Neomeniomorpha Pelseneer, 1906 (also called 

Ventroplicida Boettger, 1955) and Caudofoveata Boettger, 1955 or Chaetodermomorpha Pelseneer, 1906. The Museum of 

Natural History Berlin (formerly Zoological Museum Berlin, ZMB) houses rich type material essentially of Solenogastres on 

which to a substantial degree the preeminent German malacologist Johannes Thiele (1860–1935), working as curator in this 

collection from 1905 on, has based his respective systematic accounts of that time. A review given here briefly outlines the 

historical development of knowledge on the systematics and phylogeny of aplacophoran molluscs allowing two conclusions: 

First, that evidendently Thiele struggled with the very same problems of molluscan classification as we still do more than a 

century of zoological systematics later; and second, that Thiele’s erroneous assumption of Solenogastres being closely related 

to annelids rather than molluscs resulted in the deposition of aplacophoran material of the ZMB (and hence the late rediscovery 

of it) in the “Vermes” department, then initiating this annotated type catalogue. Here we provide information on a 

total of 31 aplacophoran taxa in the ZMB, including notes on type specimens and localities, their original description and 

current systematic placement. The majority (i.e. 25 taxa) are represented by types, essentially being named by Thiele in 23 

cases. With the exception of one caudofoveate, all these aplacophoran molluscs in the ZMB are Solenogastres. Following 

recent classification they are assigned to 20 genera. The type material was mainly collected by German imperial expeditions, 

which are briefly reviewed, in particular the First German South Polar Expedition on board of the sailing vessel “Gauss”, 

1901–1903, with a total of 15 new aplacophoran species, all from the very same type locality near the Antarctic Gaussberg 

volcano at 66 2 0 S, 89 38 0 E, collected by the expedition’s biologist Ernst Vanhöffen. 

Introduction 

Aplacophorans are vermiform (i.e. wormshaped), 

shell-less, often deep-sea and bottomfeeding 

animals with a phylogenetically basal position 

in the Mollusca. Instead of the shell pro- 

* Corresponding author: e-mail: matthias.glaubrecht@museum.hu-Berlin.de 

** e-mail: luitfried.salvini-plawen@univie.ac.at 

# 2005 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim 

tection found in most other molluscs, their slender 

body is provided with a mantle cover of a 

chitinous cuticle invested with numerous aragonitic 

integumental sclerites (i.e. “spicules”, “scales” 

and “needles”). In addition, they exhibit epidermal 

papillae (Solenogastres), a typical tetraneu- 

# 2005 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim

146 

ral nervous system, a radula with different numbers 

of teeth per row, as well as – uniquely 

among molluscs – a reproductive system with 

gonads emptying into the pericard and secundary 

gametoducts; for detailed accounts on anatomical 

characters in aplacophoran molluscs see the general 

treatments e.g. by Simroth (1893b), Thiele 

(1913c, 1925), Hoffmann (1929), Fischer-Piette & 

Franc (1960), Hyman (1967), Salvini-Plawen 

(1967, 1971, 1972, 1985), Scheltema et al. (1994) 

and Scheltema (1998). 

Although aplacophoran molluscs are a small 

group of animals ranging from 1 mm to 30 cm, 

and are only rarely preserved even in major museum 

collections, they actually represent a phylogenetically 

highly important group, with constituent 

taxa possessing morphological characters 

considered essential for the reconstruction of the 

evolution of Mollusca; see e.g. recent reviews 

and discussions in Salvini-Plawen (1967, 1972, 

1985, 2003a), Scheltema (1978, 1993, 1996), Ivanov 

(1996), Salvini-Plawen & Steiner (1996) and 

Haszprunar (2000). 

However, the curious and unique mixture of 

plesiomorphic and derived features of aplacophorans 

have contributed to long-standing disputes 

as to the phylogenetic relationships not 

only to other mollusc groups but also among 

these shell-less taxa that apparently modified 

many of their anatomical characteristics in dramatical 

ways. Ongoing controversies include the 

question of the validity of the Aculifera or Amphineura 

concepts (the latter going back to Ihering 

(1876)), that both propose Aplacophora Ihering, 

1876 and Polyplacophora Gray, 1821 as 

adelphotaxa, or the question whether Aplacophora 

are monophyletic. 

While molecular phylogeny point to a diphyletic 

origin, it was unable to propose probable 

sister group relationships to other molluscan 

groups (for reasons given below). Most currently 

used text books in zoology (e.g. Götting 1985, 

1996; Brusca & Brusca 2003) and some monographic 

malacological accounts (e.g. Scheltema 

1996, 1998, 2001) thus continue to perceive 

Aplacophora and Aculifera as monophyletic 

taxa. This conservative view is largely ignoring 

available evidence for an alternative systematization 

according to cladistic analyses that suggest 

the latter two taxa being paraphyletic assemblages 

(i.e. grades) only, both representing basal, 

independent off-shoots of Mollusca (Salvini-Plawen 

& Steiner 1996; Haszprunar 2000). The unresolved 

phylogenetic issues currently compromise 

our understanding of molluscan origin and 


Glaubrecht, M. et al., Aplacophoran Mollusca in the Natural History Museum Berlin 

early evolution; for a recent discussion in a 

broader molluscan framework see e.g. Salvini- 

Plawen & Steiner (1996), Salvini-Plawen (1990) 

and Lindberg & Ponder (1996). 

The systematics within aplacophoran molluscs 

is also partly unresolved. According to traditional 

knowledge the aplacophoran molluscs 

comprise two higher level taxa (subclasses or 

proper classes) with strongly distinct anatomical 

habitus and habits. In most classifications these 

taxa are separated as Solenogastres Gegenbaur, 

1878 (sensu nomine Simroth, 1893) and Caudofoveata 

Boettger, 1956. The naming and separation 

of both groups – that were first discovered 

in Scandinavian waters in 1844 (Chaetoderma nitidulum 

Lovén, 1845) and 1875 (Neomenia carinata 

Tullberg), respectively – refers to the presence 

of a small foot within a ventral groove in 

Solenogastres (thus “Furchenfüßer” or “Bauchfurcher” 

in German) or its absence in Caudofoveata 

(“Schildfüßer”). Solenogastres lack a cuticular 

oral shield and posterior ctenidia, live 

mostly cnidarivore and even epizoic on animal 

colonies (such as Octocorallia and Hydrozoa) or 

benthic and are hermaphrodtites. In contrast, a 

cuticular oral or pedal shield and paired ctenidia 

are present in the Caudofoveata which live 

benthic and burrowing in marine soft sediments 

where they move by hydrostatic action; the latter 

are micro-omnivore to micro-carnivore (feeding 

on debris, foraminiferans and other small organisms) 

and are of separate sex. 

Unfortunately, our knowledge of the biology 

of aplacophorans is still fragmentary. While only 

about 120 species of Caudofoveata are known so 

far, about 240 species of Solenogastres have 

been named, with new ones currently under description. 

This relation in species number is also 

more or less reflected by the extant historical 

material in the ZMB where Solenogastres largely 

dominate (see below). Undoubtedly, the latter 

taxon is not only specious but also more diverse 

in respect to habitats used, which can be 

benthic, epizoic, and meiofaunal or even burrowing. 

In recent years several new additions come 

from various regions of the world, as is evident 

from the descriptions of new taxa of different 

Solenogastres from Norway (Handl & Salvini- 

Plawen 2001, 2002), from the Mediterranean Sea 

(Salvini-Plawen 2003b) from the Western Indian 

Ocean (Todt & Salvini-Plawen 2003), from the 

southern hemisphere (Salvini-Plawen & Paar- 

Gausch 2004) and from various other regions 

(Salvini-Plawen 2004), as well as Caudofoveata

Mitt. Mus. Nat.kd. Berl., Zool. Reihe 81 (2005) 2 / http://museum-zool.wiley-vch.de 147 

(Prochaetodermatidae) of the Atlantic Ocean 

and Mediterranean Sea (Scheltema & Ivanov 

2000) or the Indian and Pacific Oceans (Ivanov 

& Scheltema 2002, 2004). Apart from the monograph 

of antarctic-subantarctic Solenogastres 

with 75 new species (Salvini-Plawen 1978) another 

striking example for the recent increase in 

knowledge is the Australian region, where apparently 

a rich and diverse fauna exists in the southeast 

of the continent with at least 32 species in 

14 or more genera, some still awaiting description 

(see Scheltema 2001). Prior to this recent 

survey only two species were recorded by Thiele 

(1897), viz. Notomenia clavigera and “Proneomenia” 

(¼ Epimenia) australis of which the types 

are extant in the ZMB collection (see below). 

Irrespective of their rare representation in 

most collections, aplacophorans form a numerically 

small but consistent deep-sea faunal element; 

some taxa are apparently common and 

sometimes form an abundant part of the shelf, 

bathyal, abyssal and hadal oceanic benthic 

macrofauna from depths of c. 200 m to 9000 m 

(see Belyaev 1966). Since representatives of both 

aplacophoran taxa are found in deeper water on 

the continental shelf and in the deep-sea down 

to the abyssal zone and even at hydrothermal 

vents, they are generally not only less accessible 

than other molluscs; since they are often small, 

with many being less than five millimeters (but 

also reach exceptionally 300 mm), they certainly 

represent a neglected part of the Mollusca; for a 

review of the biology of aplacophorans see, for 

example, Salvini-Plawen (1971, 1985) and Scheltema 

(2001). 

Not only recent discoveries of new taxa have 

spurred aplacophoran research, but also renewed 

interest in the phylogeny of metazoans in general 

and mollusca in particular. Clearly, their systematics 

with respect to the new findings is still in 

flux and continuously has to be adapted. However, 

in view of the fact that traditional classifications 

did apparently neither reflect the phylogenetic 

affinities among nor within the two major 

groups Solenogastres and Caudofoveata, the actual 

systematics in general appears to be in accordance 

with computerized cladistic analysis 

(Salvini-Plawen 2003a). 

Nevertheless, this situation all the more 

strengthens the importance of historical museum 

material. The recent interest in aplacophoran 

classification often necessitates the re-examination 

of this archival material in particular of the 

many taxa of Solenogastres named by one of the 

former leading malacologists, Johannes Thiele. 

This re-examination of ZMB material resulted, 

for example, in the recent transfer and re-description 

as type species of a new genus, as in 

the case of Thieleherpia thulensis (Thiele 1900) 

by Salvini-Plawen (2004: 86), and in the erection 

of a new family Notomediidae, based on the reexamination 

of the type species of the respective 

genus Notomenia clavigera Thiele, 1897 (see Salvini-Plawen 

2004: 89). The revision of earlier descriptions 

in light of new material, collections 

and taxa described will eventually allow more 

accurate systematics at the generic and family 

level as shown, for example, recently in the Clavibelonia 

(cf. Salvini-Plawen 2004). Again, this 

underscores the importance of historical collections 

such as Thiele’s type material of aplacophorans. 

Therefore, the objective in the present paper 

is twofold. The Museum of Natural History of 

Berlin (formerly Zoological Museum Berlin, 

ZMB) houses rich type material of aplacophorans, 

particularly Solenogastres, which are evidently 

of basic importance for any systematic 

evaluation of these molluscs. This material stems 

to a substantial degree from the systematic work 

of the former curator of the Berlin Malacological 

Collection, Johannes Thiele (1860–1935), who 

was certainly one of the preeminent malacologists 

of the twentieth century. Following positions 

as assistant in the invertebrate section of 

the Dresden Museum für Tierkunde (1891– 

1895), Strassburg’s Zoological Institute (1895– 

1896) and Göttingen’s Zoological Institute 

(1896–1898), Thiele took a position as a lecturer 

at the Zoological Institute in Berlin in spring 

1898, before starting as a scientific collaborator 

or helper at the Zoological Museum in Berlin in 

spring 1899, where he became Research Assistant 

in 1900, then curator of the Crustacea Section 

in 1903, before he finally took charge of the 

Mollusk Section in 1905, after the death of 

Eduard von Martens (1831–1904). Thiele 

worked there also after his official retirement (in 

1925) until his death in 1935. For a brief biography, 

including a bibliography, see e.g. Bieler & 

Boss (1989). 

The Solenogastres were among the molluscan 

groups Thiele paid particular attention to, culminating 

in his revision of this taxon in Das Tierreich 

(Thiele 1913c). Among the total of 291 

genus-level taxa in the phylum Mollusca introduced 

by Thiele, there are 11 aplacophoran 

names (see Boss & Bieler 1991), to which at 

least 24 new Solenogastres species names by 

Thiele can be added according to the material 


148 

presented herein. First, we give an annotated 

catalogue of the ZMB type material with notes 

on the type specimens, including their original 

status and references (i.e. author and bibliographic 

source), type localities, the nature of the 

extant type material and current systematic placement 

and taxonomic status of the taxa. Second, 

we aim to place these taxa within the framework 

of the current systematic knowledge on 

the phylogeny of aplacophoran molluscs by reviewing 

the development of this knowledge, including 

an outline of the results of recent cladistic 

analyses. This annotated catalogue of aplacophoran 

types continues earlier compilations of molluscan 

material in the ZMB (see bibliography of R. 

Kilias (1929–1999) in Glaubrecht 2001). It adds 

to those catalogues of other less species-rich 

groups, such as Polyplacophora (Kilias 1995a), 

Scaphopoda (Kilias 1995b), but also Cephalopoda 

(Glaubrecht & Salcedo-Vargas 2000; Köhler & 

Glaubrecht 2004), whereas the extremely rare 

Monoplacophora are not represented in the 

ZMB. 

Brief review of the history of classification 

in aplacophoran molluscs 

Seit ich angefangen, mich mit der Anatomie 

von Mollusken zu beschäftigen, ist es mein Bestreben 

gewesen, über die Verwandtschaftsbeziehungen 

des ganzen Stammes wie einzelner 

Gruppen ins Klare zu kommen. 

Johannes Thiele, 1894: 222 

Representatives of the two major aplacophoran 

molluscs were first published from Scandinavian 

waters in 1845 and 1875, respectively, and subsequently 

recognized increasingly often in dredged 

material from oceanographic expeditions. For the 

first time, Heinrich Simroth (1893a) not only discussed 

the then available morphological data on 

aplacophorans but added tentatively a first “Versuch 

eines Systems” in listing the known genera 

according to their geographical occurrence, i.e. 

as nordic and mediterranean forms. This not 

being a systematization in the strict sense, it was 

an early approach more to compile rather than 

to classify the described taxa; for his extended 

systematization see Simroth (1893b: 226–233). 

Thiele (1902c) is often cited with the conclusion 

from his first account on the systematic placement 

of the Solenogastres in connection with the phylogeny 

of the Mollusca that aplacophoran species 

are not molluscs (e.g. Scheltema 1996: 57). Indeed, 



in his final statement Thiele (1902c: 455) proposed 

explicitly (and printed in bold) that “Solenogastres 

are a group of worms closely related to annelids 

and gordiids, connecting those – by the relationship 

of the uterus and the pericard as well as the 

inital stages of a radula organisation – to the molluscs, 

among which the chitons are particularly closely 

related to them through the preservation of the 

lateral cords [of the nervous system]”; [translated 

from the German original – M.G.]. In his phylogenetic 

tree, printed as text-figure in Thiele (1902c: 

438) and following contemporay perception of 

phylogenetics, he depicted this hypothesis by placing 

Solenogastres among the Vermes, as a late offshoot 

from a branch eventually leading to the 

Mollusca. 

However, Thiele’s reservations as to the placement 

of Solenogastres within molluscs go back 

to his earliest accounts published in 1891 and 

1892, when he suggested that “Wurmmollusken” 

(i.e. worm-shaped molluscs) are the Amphineura 

of H. von Ihering with the two constituent orders 

Solenogastres or Aplacophora and the Polyplacophora 

(Thiele 1892). He viewed Solenogastres 

as more primitive and the amphineurans in an 

intermediate and transitional position – as socalled 

“Ûbergangsformen” (Thiele 1892) – between 

annelids and molluscs (see also Thiele 

1891: 521). Later, Thiele (1895: 867) concluded 

from his ongoing dissections of aplacophorans 

that “Solenogastres are no molluscs because they 

lack the typical molluscan characters” [translated 

from the German original]. In addition, Thiele 

(1895: 869) suggested to transfer Solenogastres 

to the “worms” (i.e. Annelida), thus, also breaking 

up the Amphineura. 

He continued to stress this view for the rest of 

his life, as is evident, for example, from his accounts 

on Solenogastres in Thiele (1925: 13), 

where he perceived them – according to then 

prevailing thinking – as “phylogenetische Ausgangsform”, 

i.e. not only as hypothetical but still 

extant evolutionary ancestor to the Mollusca (his 

“Elternform”, see Thiele 1891: 480), with close 

affinity now to turbellarian worms. Thiele’s position 

is also evident from the introductory statement 

in his last contribution on Solenogastres 

(Thiele 1933: 144). He was followed in his view, 

for example, by Nierstrasz (1909–1910) and 

Odhner (1921) but not Heath (1911), while e.g. 

Hans Hoffmann (1929) expressed an opposed 

opinion in his critical evaluation of the “Amphineura” 

discussion; for an historical review see 

e.g. Hoffmann (1929, 1937) and in particular Salvini-Plawen 

(1967, 1972).


It was only consequent from this, albeit erroneous, 

hypothesis that Thiele (1929) did not include 

aplacophorans in his Handbuch der Systematischen 

Weichtierkunde, which started 

accordingly with the Loricata Schumacher, 1817 

(¼ Polyplacophora Gray, 1821) interpreted by 

him as most primitive and true representatives of 

the phylum Mollusca. In the second volume of 

his Handbuch, Thiele (1935: 1074) for the last 

time outlined the phylogeny of the Mollusca, 

stating that the Solenogastres cannot be viewed 

as derived from shelled animals due to, for example, 

their peculiar hautmuskelschlauch 

(p. 1073) and because of other characteristic anatomical 

features, thus repeating that they are to 

be perceived as “Vermes” and excluded from 

the Mollusca (p. 1074). 

Two conclusions can be drawn from this. First, 

evidendently, Thiele and his contemporaries 

struggled with quite similar problems of molluscan 

classification as we still do over a century of zoological 

systematics later, viz. monophyly vs. paraphyly 

of the aplacophorans, their relationship to 

Polyplacophora within the Amphineura and/or 

Aculifera concept and the ancestry of the Mollusca 

within the metazoans, in particular their relationship 

with Annelida. And second, Thiele’s erroneous 

assumption of Solenogastres being 

“Vermes” resulted in the deposition of aplacophoran 

material of the ZMB, and hence the late 

re-discovery of it, in the “Vermes” department 

that initiating this annotated type catalogue. 

The first half century of research on aplacophoran 

classification was dominated by the practice 

to consider all “worm-like” molluscs as Solenogastres, 

thus including also those taxa later to 

be distinguished as Caudofoveata; see e.g. treatments 

by Simroth (1893b), Thiele (e.g. 1902c, 

1913c), Nierstrasz (1908) and Heath (1911). 

Thiele (1913c) distinguished four taxa among this 

paraphyletic group, the Chaetodermatidae, Neomeniidae, 

Proneomeniidae, and Lepidomeniidae. 

Later, Thiele (1932) suggested to separate five 

families within the aplacophorans (all considered 

by him to represent Solenogastres), viz. Neomediidae, 

Proneomeniidae, Gymnomeniidae, Lepidomediidae 

and Crystallophrissontidae. The latter, 

however, represents Chaetodermomorpha 

according to current knowledge which were subsequently 

separated by Boettger (1956); see e.g. 

Fischer-Piette & Franc (1960) and Salvini-Plawen 

(1967). 

Following Thiele’s era, Sigurd Hoffman (1949) 

not only contributed essentially new information 

on the then widely neglected aplacophoran 

groups but also discussed at length phylogenetic 

pathways and higher classification based on comparative 

investigations. Others, such as in Germany 

Hans Hoffmann (1929, 1951) stated not to 

separate aplacophorans from the Mollusca and 

proposed to view the Solenogastres as most “primitive” 

group within this phylum. It was then 

Boettger (1956) and later in particular Salvini- 

Plawen (1967, and subsequent publications) who 

took up, after almost half a century of neglectance 

of this important molluscan group, again 

the controversies as to the monophyly and phylogeny 

of “Aplacophora”. In the absence of cladistic 

analyses, most authors long followed the 

general assignment of all “worm shaped” molluscs 

as two clades Solenogastres or Neomeniomorpha 

(or Ventroplicida Boettger, 1956 which 

is a later naming) and Caudofoveata or Chaetodermomorpha 

within a monohylum Aplacophora. 

In contrast to these classifications of 

“Aplacophora” or “Aculifera”, Salvini-Plawen 

(e.g. 1972, 1985, 1991) has, early and repeatedly, 

argued that aplacophorans are paraphyletic or 

diphyletic, respectively (see below). 

Furthermore, despite revision (Salvini-Plawen 

1978) the systematic relationships remained 

partly unresolved within Solenogastres, and the 

status of some families was poorly understood, 

as was recently shown, for example within the 

Cavibelonia by Salvini-Plawen (2004). Earlier 

suggestions to separate three families most likely 

did not reflect natural phylogenetic lineages (Salvini-Plawen 

1967). According to the systematization 

suggested by Salvini-Plawen (1978; see also 

phylogram in Salvini-Plawen & Steiner 1996: 31), 

four orders are to be distinguished within the Solenogastres, 

viz. Pholidoskepia, Neomeniamorpha 

(both as Aplotegmentaria) and Sterrofustia 

plus Cavibelonia (these two as Pachytegmentaria). 

Thus, the Cavibelonia, in which 11 genusgroup 

or families are now distinguished, replaced 

the Proneomeniidae s.l. and most of Neomeniidae 

s.l. in former systems. For discussion and references 

see Salvini-Plawen (1978, 2003, 2004). 

This systematization is followed herein; see section 

C for a classification of the ZMB type material. 

Phylogenetic analyses 

For several decades now two hypotheses have 

been discussed as to the classification reflecting 

phylogenetic relationships of the two aplacophoran 

taxa, Solenogastres (or neomenioids) and 


150 

Caudofoveata (or chaetoderms). Paraphyly of 

the aplacophorans was proposed by Salvini-Plawen 

(1967, 1972) and is supported by evidence 

presented by Salvini-Plawen & Steiner (1996) 

and Haszprunar (2000). It was suggested to rank 

Caudovofeata and Solenogastres as separate 

classes, and that Caudofoveata should be distinguished 

from the remaining molluscan classes 

(those named Adenopoda). 

In contrast, Scheltema (1978, 1993, 1996) and 

others continue – irrespective of her own separate 

treatment of both distinct groups (e.g. in 

Scheltema et al. 1994) – to view both taxa as 

subclasses within a single monophyletic class 

Aplacophora that together with the Polyplacophora 

form a clade Aculifera. For example, 

Scheltema et al. (1994: 13) considered them to 

be “specialized molluscs with a derived worm 

shape accompanied by reduction or loss of the 

foot”, while other features “reflect a rather primitive 

molluscan state”. To account for these 

“otherwise apparently contradictory states”, Scheltema 

(1996: 57) suggested progenesis and rapid 

evolution. However, it should be noted here that 

this combination of derived and plesiomorphic 

features in the grundmuster is a common biological 

phenomenon long known and discussed as 

mosaic evolution without necessarily involving 

progenetic or paedomorphic processes. 

Recent cladistic analyses by both Salvini-Plawen 

& Steiner (1996) and Haszprunar (2000) 

found the Solenogastres in the most basal position 

suggesting it to represent the earliest molluscan 

offshoot. At the same time the monophyly 

of all remaining molluscan classes is 

suggested, named Hepagastralia by Haszprunar 

(2000: 125) and that way reflecting the distinct 

and complex subdivision of the midgut as synapomorphy. 

Thus, computation of all available 

morphological data indicate that both “Aplacophora” 

and “Aculifera” should not be considered 

monophyletic clades, but that – following the 

Solenogastres as first off-shoot – the Caudofoveata 

are the sister group to the remaining molluscs 

(i.e. Testaria). 

Based on analyses by Hoffman (1949) and 

summarised in Salvini-Plawen (2003a) it is argued 

that the mantle cavity in Neomeniomorpha 

and Chaetodermomorpha reveals an independent 

evolutionary transformation of both taxa, 

and that the midgut sac in Caudofoveata is not 

homologous to the so-called digestive gland in 

Testaria (i.e. Placophora þ Conchifera) which, 

thus, cannot be interpreted as synapomorphy 

(see e.g. Haszprunar 2000). The question remains 



unresolved whether the lack of ctenidia is a plesio- 

or apomorphic state in Solenogastres. In 

contrast, radula features (e.g. monoserial type 

with median bars or two teeth with symphysis) 

suggest the Solenogastres to be more conservative, 

as is suggested in cladistic analyses. However, 

this does not necessary imply that Solenogastres 

actually represent the first (extant) offshoot 

within the Mollusca. Alternatively, Caudofoveata 

could have just evolved more dramatically, 

in concert with the development of many 

derived features. As an additional aspect, this 

most recent cladistic analysis suggests that Cavibelonia 

are, contrary to earlier assumptions, 

monophyletic. 

Different molecular methods have not succeeded 

in yielding reliable results beyond supporting 

that aplacophoran groups are diphyletic 

and are thus not closer related. For example, 

analyses in Solenogastres suffer mainly from the 

fact that food material was sequenced instead of 

the aplacophorans itself (e.g. Cnidaria, see in 

Salvini-Plawen 2003a; Okusu & Giribet 2003; 

Polychaeta for Helicoradomenia in Passamaneck 

et al. 2004; pers. comm. Dr. Hermann Dreyer, 

Univ. Wien). Another problem is that much of 

the archived material in major museum collections 

has generally been fixed in formaldehyd, 

which is not considered a suitable medium for 

subsequent molecular work. 

On the phylogeny of aplacophoran taxa 

Although the origins and interrelationships of 

the seven to eight clades or classes of Mollusca 

are discussed for over a century now, no end of 

this debate is visible; for reviews and references 

see e.g. contributions in Taylor (1996) and also 

Haszprunar (2000). While the other clades are 

broadly agreed on to represent monophyla, most 

recent disagreement centers around the systematics 

and position of aplacophoran, polyplacophoran 

and scaphopod molluscs. As reviewed above, 

cladistic analyses finding aplacophorans to form 

a basic, paraphyletic assemblage, also support 

earlier suggestions to consider many conditions 

in their organization as plesiomorphic for molluscs 

rather than derived through paedomorphosis. 

In this context Haszprunar (2000: 127) and 

Salvini-Plawen (2003a: 93) pointed out that many 

shared aplacophoran features can now be considered 

as characters typical for the so-called and 

long-thought-of “hypothetical ancestral molluscs” 

(HAM).


However, many issues in molluscan phylogenetics 

still remain unresolved. Extremely difficult 

is the interpretation of the scarce ontogenetic 

data, not only in molluscs in general but in the 

aplacophorans in particular. While for Caudofoveata 

only the lecithotrophic larvae of two species 

are known (Salvini-Plawen 1990, Gustafson 

in Nielsen 1995), development in Solenogastres 

is either lecithotrophic or direct. The ciliated, 

free swimming larva in some Solenogastres resembles 

superficially the so-called “trochophora” 

larva in annelids; for a review of aplacophoran 

reproduction and development see e.g. Salvini- 

Plawen (1985). The developmental stages in a 

Soleogastres was described originally as so-called 

“Pruvot larva” by the French malacologist G. 

Pruvot (1890). He reported on a late larva (Nematomenia 

banyulensis) depicting seven rows of 

scales, or “plaques” as he called it. Although surrounded 

by ambiguities and (even after a century) 

still in need of further substantiation, this 

description of iterated spicules arrangement has 

ever since been interpreted as being homologous 

with a chiton shell, thus fueling the discussion on 

the systematic placement of Solenogastres in relation 

to polyplacophorans and other Mollusca 

(e.g. Salvini-Plawen 1972, 1985; Salvini-Plawen & 

Steiner 1996). 

Recently, Scheltema & Ivanov (2002) reported 

another neomenioid postlarva exhibiting six iterated, 

transverse groups of spicules separated by 

seven regions devoid of spicules which they compared 

to the shell fields in developing polyplacophorans 

as well as the sclerite arrangement on 

the Cambrian fossils Wiwaxia and Halkieria. The 

authors suggested, also refering to recent insight 

from developmental biology on the function of 

regulatory genes, that this iteration (or serial repetition) 

in morphogenesis of ectodermal skeletol 

structures in these taxa is a result of processes 

already present in early pre-Cambrian 

bilaterial animals and, thus, would indicate evolutionary 

relationships among Neomeniomorpha, 

Polyplacophora and early Paleozoic fossils. They 

suggested that these rows of transverse spicules 

that appear briefly in the early development of 

some Solenogastres are expressions of genes that 

may be present generally in aplacophorans. 

However, the authors failed to comment on the 

fact that, when assuming these features do actually 

represent an ancient developmental process 

also in early molluscs, then these ectodermal 

iterations – albeit homologues – should be 

viewed as plesiomorphic. They would then not 

qualify for supporting an aplacophoran or aculi- 

feran assemblage. In ignorance of this it may become 

understandable that these latter authors, in 

context of their new ontogenetic data, again 

stress their view that aplacophorans are “derived 

molluscs related to chitons and are perhaps progenetic” 

(Scheltema & Ivanov 2002: 7). 

In summary, recent phylogenetic analyses suggest 

that aplacophorans are unlikely to represent 

a monophyletic clade. Instead, among the basal 

molluscan groups, Solenogastres may be viewed 

as the most earliest off-shoot while Caudofoveata 

may be closer to the remaining Mollusca (yet 

not as “Hepagastralia”). However, interpretation 

of character polarity as well as systematics remain 

controversial. 

Material and Methods 

Some aplacophoran type material and additional non-type 

material (see section B below) was found by one of us 

(L.M.), more or less accidentially, during our on-going evaluation 

(since 1997) of all molluscan type material, as it was 

long misplaced among opistobranch material in cabinets of 

the Malacological Department in the ZMB. That way, aplacophorans 

were at least in part hidden to R. Kilias who started 

to publish, among other major gastropod groups (see bibliography 

in Glaubrecht 2001), also on some minor (i.e. less 

speciose taxa) such as e.g. Polyplacophora (Kilias 1995a) and 

Scaphopoda (Kilias 1995b). 

As we can reconstruct from some notes accompanying this 

aplacophoran material, it was long housed, for the reason of 

Thiele’s explicit perception of Solenogastres not being Mollusca 

(see review above), in the “Vermes” department of the 

ZMB. Upon investigation we luckily found some other parts 

of the aplacophoran type material there which are now returned 

and housed together with Thiele’s original serial sections 

in the Malacological Collection as listed in the compilation 

below. 

Following sorting and inventarization of this re-discovered 

aplacophoran material done essentially by L.M., a list of all 

constituent lots was prepared. A first draft of this catalogue 

was sent to LvS-P in Dezember 2002, who agreed to check 

and comment on the taxonomic status of these taxa which 

was crucial for our endeavour. The final version of the manuscript 

was written by M.G. in August 2004, with the inclusion 

of the introduction and the historical parts, a review of aplacophoran 

classification and its role in molluscan phylogeny. 

This version was finally again cross-checked in October 2004 

by LvS-P. 

Based on all combined information available to us then, 

we found additional type material of Thiele’s aplacophorans, 

viz. of Neomania grandis, Rhopalomenia eisigi and Amphimenia 

neapolitana, all described by Thiele (1894), but missing 

from the ZMB collection, due to research facilitated by decisive 

notes of LvS-P made 1977 in the Zoological Museum 

Amsterdam. It has now, after a long post-WW II odyssee, 

returned to the ZMB; for more details see below under the 

species. 

Remarks on the origin of the Berlin type material 

The type material that comprises complete animals, 

histological serial sections as well as tissues 


152 

and animal parts embedded in paraffin blocks 

has diverse origins. For example, due to his early 

investigations of aplacophorans Thiele was provided, 

starting in the time prior to his curatorial 

position at the ZMB in 1905, with material by 

various zoologists, as he reported in case of three 

newly named species of Solenogastres from Neapel 

(see Thiele 1894: 222–223), or in case of 

Anamenia amboinensis. Later, Thiele was also a 

contributor of scientific descriptions based on 

material from several of the German imperial 

expeditions around the turn of the century. In 

particular, he studied the aplacophorans from 

two major oceanographic expeditions, viz. the 

German Deep-Sea Expedition on board of the 

steamer “Valdivia”, 1898–1899 (with n ¼ 3 new 

species), and in particular the First German 

South Polar Expedition, 1901–1903, on board 

the sailing vessel “Gauss” (with 15 new species 

described). Thus, these German expeditions were 

essential in providing new taxa from various regions 

of the world and taxonomic groups. However, 

they are largely unknown today for various 

reasons and, although being of immense historical 

importance, only few historians of science 

have dealt with them yet. Here, only brief mention 

will be made of those expeditions in context 

crucial for collecting Thiele’s aplacophoran material, 

with some hints at the most important available 

accounts. 

The German “Gazelle” Expedition between 

1874 and 1876 on board of the navy corvette 

S.H.S “Gazelle”, lead by Georg Emil Gustav 

Freiherr von Schleinitz (1834–1910), was on 

route around the world with a focus on research 

in the deep-sea of the Atlantic and Indic Ocean, 

nearly at the same time as the British “Challenger” 

expedition. One of the expeditions’ aims 

was to study the Venus orbit on the near-Antarctic 

Kerguela Island. As naturalist the Swiss 

zoologist Theophil Studer (1845–1922) took part 

in the “Gazelle” expedition, who was instrumental 

in collecting and later describing most of its 

natural objects. An original expedition report 

can be found in Schleinitz & Rottok (1889); see 

also brief accounts in Studer (1882) and Buchwald 

(1999). Only one aplacophoran molluscs, 

Epimenia australis (Thiele 1897) from off the 

NW coast of Australia, was described and deposited 

in the ZMB, so we refrain here from a 

more detailed review of this endeavour. Nevertheless, 

the “Gazelle” marks the start of German 

maritime explorations later continued by two 

other major expeditions of zoological importance. 



The German Deep-Sea Expedition (Deutsche 

Tiefsee-Expedition), on board of the research 

steamer “Valdivia”, was on route in the Atlantic 

and Southern Indic Ocean from July 31, 1898 to 

May 1, 1899, as the first major German deep-sea 

research activity subsidized by the German Empire 

in an nationalistic attempt to keep up with 

the successful deep-sea expeditions in particular 

by the British, as e.g. the “Challenger” expedition 

1872–1876. Lead by the biologist Carl Chun 

(1852–1914) from the University of Leipzig, the 

DTE was most innovative in then modern echo 

sounding and sampling techniques and achieved 

important results and collections in nearly all zoological 

phyla. The material on which specialists 

from all over Europe worked for the following 

decades – among them scientists at the ZMB 

such as Johannes Thiele (recognized, among 

many other taxa, n ¼ 3 new solenogastres species) 

– forms today an essential part of the Berlin 

museum’s type collection. Only one example 

of the rich material provided by the “Valdivia” 

expedition is reflected in the cephalopod type 

collection housed in the ZMB which renders it 

second only to the respective collection in The 

Natural History Museum in London (formely 

British Museum of Natural History); see Glaubrecht 

& Salcedo-Vargas (2000) with additions in 

Köhler & Glaubrecht (2004). A most readable – 

and then very successful – travel narrative was 

given by Charl Chun (1900); the zoological results 

were published in 24 volumes, edited until 

his death by Chun (1902–1914) himself. On the 

occassion of the 100 th anniversary of the “Valdivia” 

expedition, short historical reviews were 

published recently e.g. by Kabisch (1998), Coleman 

(1999), and more detailed by Landsberg 

(2000). Unfortunately, however, no comprehensive 

treatment of this most important expedition 

and evaluation of its achievements is available 

yet. 

The First German South Polar Expedition 

(Deutsche Südpolar-Expedition) was on route 

from November 11, 1901 until November 25, 

1903, on board the sailing vessel “Gauss”. Lead 

by the young German polar scientist and professor 

of geophysics Erich von Drygalski (1865– 

1949), it was part of the “heroic” period of polar 

exploration and, as one of the first truely scientific 

endeavours, scientifically highly successful. 

Carried out in the framework of an international 

programm of geophysical observations, the expedition 

was nevertheless rated a failure in Germany 

due to comtemporary political hopes for 

imperialistic expansion, resulting in a lack of re-


cognition of its merits for long; for a brief historical 

account along these lines see Lüdecke 

(2001) and Lüdecke et al. (2001). At the turn of 

the century most parts of the South Polar region 

were completely unknown and following the 

“Valdivia’s” successful exploration of southern 

regions south of the Kerguela Islands at 90 E, it 

was the clear aim and explicit task of the 

“Gauss” expedition to scientifically explore the 

South Polar area, in contrast to the geopolitical 

endeavour of the parallel and competing British 

expedition lead by Robert Falcon Scott (1868– 

1912) on the “Discovery” who was successful in 

reaching south at 82 17 0 S, while Drygalski only 

made it to about 66 S. 

On board of the “Gauss” was, as the only biologist 

officially employed, the geograph and zoologist 

Ernst Vanhöffen (1858–1918), who had 

also been a member of the former German 

Deep-Sea Expedition on the “Valdivia”. Educated 

at the University Königsberg, Eastern 

Prussia, under the influence of Richard Hertwig 

and Carl Chun, Vanhöffen was lecturer, assistant 

and honorary professor at the University Kiel 

from 1890 to 1906, when he became curator for 

Crustacea, Myriapoda and Coelenterata at the 

Museum of Natural History in Berlin. He is today 

mostly remembered for his studies on cnidarian 

medusae and as member of several exploring 

expeditions, most prominently the 

German South Polar Expedition; for a brief biography 

of Vanhöffen including complete bibliography 

of his zoological writings see Lohmann 

(1918). It was Vanhöffen who collected most zoological 

objects, among them also the many aplacophorans, 

as is evident from the original labels 

and documents. Most material was brought back 

in particular from the so-called “Winterlager”, 

when the “Gauss” became beset by pack-ice at 

66 2 0 S and 89 38 0 E in the Indian sector of the 

Antarctic, 85 km off the Antarctic shelf, and in a 

region later to be called “Kaiser Wilhelm II. 

Land”. It is this the type location were the majority 

of the ZMB material, i.e. a total of n ¼ 15 

new solenogastres species were found (see details 

in Section A), rendering this expedition 

most successful in bringing back Antarctic taxa 

of this interesting molluscan group. The DSE 

spent the Antarctic winter in 1902 on the sea ice 

while doing research for 50 weeks. Vanhöffen 

during this time conducted numerous nettings 

and dredges in various depths down to the bottom 

in –385 m. During one of the seven “inland” 

explorations by dog-sledges Drygalski’s expedition 

discovered and explored the so-called 

“Schwarzen Berg” or“Gaussberg” volcano, as it 

was later entered into the maps. This only area 

free of ice found by the expedition had an elevation 

of 366 m above the sea. After breaking free 

from the ice in early 1903 the expedition failed 

to find other access further south to the Antarctic 

and returned in June 1903 to South Africa 

and home via the Atlantic Ocean. Its scientific 

collections were studied and analysed during the 

subsequent three decades, the results being published 

by Drygalski as editor in a total of 20 volumes 

and 2 atlantes (Drygalski 1905–1931); see 

also Drygalski (1904) for an account of his twoyears 

Antarctic expedition. (Note that it took 85 

years to translate into English this semi-popular 

narrative of the First German South Polar Expedition 

which partly contributed to its neglectance 

in the annales of history of science). 

Another addition to the knowledge of the invertebrate 

fauna of polar regions came from the 

German Römer & Schaudinn Expedition to Arctic 

waters around Spitsbergen Island in 1898 on 

board the steamer “Helgoland”; for a travel narrative 

see Römer & Schaudinn (1900). Basically 

a privat enterprise it, nevertheless, gained many 

zoological objects of interest. Among them, 

Thiele (1900, 1913b; see also 1932) described 

two new aplacophoran species. 

All this material finally amounted to a total of 

n ¼ 24 new aplacophoran mollusc taxa newly described 

by Thiele (including here also his “kergulensis”, 

albeit re-described only much later by 

Salvini-Plawen; see under the species); all are 

now extant in the Malacozoological Collection in 

the ZMB. 

Designation of type material 

Specimens of Thiele’s aplacophoran material 

were stored in a sea-water/ethanol solution 

(“Seewasseralkohol” e.g. in Thiele 1894: 222), of 

which Thiele generally made histological serial 

sections himself, using a double staining agens 

(Boraxkarmin and Methylenblau); this is noted 

as “Thiele fec.” on his original labels and below 

in Section A; an example of Thiele’s sections is 

illustrated in Fig. 1. 

In some taxa several specimens are registered 

including intact animals as well as histological serial 

sections mounted on microscopic doubleslides. 

Since the majority of Solenogastres taxa 

are well described by their anatomical characters 

only, we here follow the preferred procedure of 

treating as holotypes and lectotypes, respectively, 


154 

those parts of the material (the serial sections), 

on which the original description was based, with 

most other undissected material being para(lecto)types. 

This turned out to be essentially crucial 

in case of Dorymenia/Proneomenia antarctica 

Thiele and Labidoherpia/Pruvotina spinosa 

Thiele (for details see under the species). Lectotype 

designation are here done in accordance 

with the purpose of Article 74.7.3. of the current 

issue of the ICZN in order to ensure the name’s 

proper and consistent application. 

Museum codes: 

USNM United States National Museum of Natural History, 

Smithsonian Insitution, Washington, DC, USA; 

ZMA Zoological Museum Amsterdam, The Netherlands; 

ZMB Museum of Natural History (Museum für Naturkunde), 

Humboldt University, Berlin, Germany (formerly 

Zoological Museum Berlin); 

ZMC Zoological Museum Copenhagen, Denmark. 

Expedition abbreviations: 

DSE Deutsche Südpolar-Expedition, 1901–1903; i.e. 

DTE 

German South Polar Expedition; 

Deutsche Tiefsee-Expedition, 1898–1899; i.e. German 

Deep Sea Expedition; 

“Gazelle” Expedition with the vessel “Gazelle”, 1874–1876. 

Section A – Alphabetic list of Solenogastres type 

material in the Malacozoological Collection of 

the Berlin Museum of Natural History 

No type material of the second major aplacophoran 

group, the Caudofoveata, is held in the 

ZMB (but see remark under Chaetoderma productum). 

However, some non-type material of 

Chaetodermomorpha is present which is listed 

together with non-type material of the Solenogastres 

in section B. 

Solenogastres 

amboinensis (Thiele, 1902a) – Anamenia 

[Proneomenia] 

Proneomenia amboinensis Thiele, 1902a: 733. 

Ty p e l o c a l i t y : “Amboina”; Indonesia: Amboina; 

leg. Richard Semon. 

Ty p e m a t e r i a l : ZMB Moll. 105.408a, part of holotype, serial 

sections on 11 slides (labelled a–i “vorn”, i.e. anterior 

part of animal, and a–e “hinten”, i.e. posterior part); ZMB 

Moll. 105.408b, part of holotype, in ethanol. 

C o m m e n t s : Type species of Anamenia Nierstrasz, 

1908. 



antarctica (Thiele, 1913a) – Dorymenia 

[Proneomenia] 

Proneomenia antarctica Thiele, 1913a: 57, pl. IV, fig. 11; 

pl. VII, figs 10–13; pl. VIII, figs 1–3, 23, 31. 

Ty p e l o c a l i t y : “Antarktis: Winterstation am 

Gaussberg”; Antarctic: winter station at Mount 

Gauss at 66 20S, 89 380E; DSE, Twist 385 m, July 

31, 1902; leg. Vanhöffen. 


sections on 11 slides (all were originally marked as 

“types”; n ¼ 6 are labelled a–f, anterior part; plus n ¼ 5 labelled 

a–e, posterior part, with b–d additionally marked as 

“Q 1925 g–i”); 

ZMB Moll. 105.415b, 4 vials with one (1. vial) containing 

presumably the remaining, non-sectioned part of the holotype. 

1. vial: 06. XII. 1902, Twist 385 m, part of specimen, dark coloured 

(holotype?); 

2. vial: 12. X. 1902, Twist 385 m, part of specimen, light coloured 

(could also be D. hoffmani); 

3. vial: 31. XII. 1902, paratype, n ¼ 1 specimen; 

4. vial: 12. X. 1602, n ¼ 1 specimen in ethanol, Thiele fec. 

ZMB Moll. 105.415c, 4 Paraffin blocks; two blocks each 

marked “grosse glatte Promeomenia antarct. 06. XII. 1902 

Vanhöffen” and two blocks marked “Promeomenia? antarctica 

10. XII. 1902”; paratypes? (but could also be D. hoffmani). 

C o m m e n t s : Re-examination and determination 

as Dorymenia antarctica (Thiele, 1913a) 

by Salvini-Plawen (1978), who was then provided, 

unfortunately, with only part of the holotype 

serial sections by R. Kilias (n ¼ 3 slides, 

with label “Q1925 g–i), thus his statement “von 

der Original-Schnittserie liegen drei Objektträger 

vor (Hinterende part.)” (Salvini-Plawen 

1978: 247). We verify herewith that the holotype 

is completely extant, mentioned by Thiele 

originally as “untersuchtes Tier” in his publication, 

with serial sections of the animal’s anterior 

and posterior part. Re-examination by LvS-P of 

the postradular sections reveals that Thiele’s description 

is brief but correct. Note that some of 

the above specimens might actually represent 

D. hoffmani. 

arctica Thiele, 1913b – Nematomenia 

Nematomenia arctica Thiele, 1913: 161, pl. 1, 2. 

Ty p e l o c a l i t y : “Spitzbergen; Genauer Fundort 

nicht bekannt”; Spitsbergen. Römer & Schaudinn 

Expedition 1898; exact locality unknown. 


sections on 2 slides (labelled “a vorn” and “b hinten” 

and “ad 5075”); 

ZMB Moll. 105.380b, 1 vial, part of holotype) in ethanol; 

ZMB Moll. 105.380c, 1 Paraffin block. (marked with asterics 

for “type”).


Fig. 1. Johannes Thiele (1860–1935) made numerous histological serial sections of aplacophoran molluscs, deposited in the 

Malacozoological Collection of the Berlin Natural History Museum today. Illustrated here are two examples of his original 

serial sections: – A –“Proneomenia” (¼ Dorymenia) hoffmani (Salvini-Plawen, 1978), two microscopic slides with cross-sections 

of Thiele’s series Q 1926 a–k (ZMB Moll. 105.420); Thiele originally assigned this material to “Proneomenia” antarctica 

(Thiele, 1913a), but later separated it from this species (see Salvini-Plawen 1978). – B –“Proneomenia” (¼ Epimenia) australis 

(Thiele, 1897), two slides with cross-sections of series 3070 (ZMB Moll. 105.407). Note Thiele’s handwriting on the original 

labels glued onto the slides. 

C o m m e n t s : For a brief note on the Römer & 

Schaudinn Expedition see “General remarks” in 

the introduction. 

australis (Thiele, 1897) – Epimenia 

[Proneomenia] 

Proneomenia australis Thiele, 1897: 398. 

Ty p e l o c a l i t y : “NW Australien”; NW Australia; 

60 fathom (i.e. 108 m isobath); leg. May 7, 

1875; “Gazelle” Expedition. 

Ty p e m a t e r i a l : ZMB Moll. 105.407a, parts of holotype, 

serial sections on 48 slides, Thiele fec. (labelled 1–36, anterior 

part, “Q 3070a–LL; and 1–12 posterior part, “Q 

3070 mm–xx); 

ZMB Moll. 105.407b, 2 parts of holotype. 

C o m m e n t s : Transferred by Nierstrasz (1908) 

as type species, by monotypy, to his new genus 

Epimenia, which is characterized by unusually 

large (up to 300 mm) and bright coloured species 

that occur on the continental shelf and are found 

even close to shore at diving depths. Six named 

congeneric species are known today, with more 

taxa to be described within the family Epimeniidae 

(see Scheltema 2001: 10). Revision and synonymy 

in Salvini-Plawen (1997a). 

austrina Thiele, 1913a – Phyllomenia 

Phyllomenia austrina Thiele, 1913a: 45, pl. IV, fig. 6; pl. V, 

figs 10–14; pl. VIII, figs 6–9. 



Gauss at 66 20S, 89 380E; DSE, 380 m, January 


Ty p e m a t e r i a l : ZMB Moll. 105.392a, holotype, serial sections 

on 3 slides; Thiele fec. (labelled “a–b vorn”, “ad Q 

1921a–b”; and “a hinten”, “ad Q 1921c”); 

ZMB Moll. 105.392b, 1 vial, 2 parts of animal and part of 

dermis (“30. 01. 1903 WST 380 m, Phyllomenia n. gen., Cuticula”); 

ZMB Moll. 105.392c, 2 Paraffin blocks (marked “Phyllomenia” 

and on back of label “Solenogaster, 2 nov. gen., 

30. 01. 1903”). 

C o m m e n t s : Type-species, by monotypy, of the 

new genus Phyllomenia Thiele, 1913a. Re-description 

in Salvini-Plawen (1978: 84–89). 


156 

carinata Thiele, 1913a – Sandalomenia 

Sandalomenia carinata Thiele, 1913a: 44, pl. IV, figs 5, 25; 

pl. V, figs 6–9. 



Gauss at 66 20S, 89 380E; DSE, 385 m, November 


Ty p e m a t e r i a l : ZMB Moll. 105.385, holotype, serial sections 

on 6 slides; Thiele fec. (labelled “a–d vorn”, 

“24. XI. 1902a, Q 1929 a–d”; and “e” þ “a hinten”, 

“24. XI. 1902a, Q 1929 e þ f”). 

C o m m e n t s : Juvenile animal; re-examination 

by Salvini-Plawen (1978: 50–51). 

cataphracta (Thiele, 1913a) – Pholidoherpia 

[Lepidomenia] 

Lepidomenia cataphracta Thiele, 1913a: 38, pl. IV, figs 1, 

15–18. 



Gauss at 66 20S, 89 380E; DSE, 385 m, December 


Ty p e m a t e r i a l : ZMB Moll. 105.386a, syntype, serial section 

on 1 slide; fec. L. v. Salvini-Plawen, 19.02.1975; 

ZMB Moll. 105.386b, syntype; 2 vials (with 2 specimens, 

Thiele fec., and n ¼ 15 specimens). 

C o m m e n t s : Thiele (1913a) mentioned the 

study of 20 specimens, here treated as syntypes. 

Systematic revision in Salvini-Plawen (1978: 51), 

with transfer as type species in new genus Pholidoherpia. 

clavigera Thiele, 1897 – Notomenia 

Notomenia clavigera Thiele, 1897: 398. 

Ty p e l o c a l i t y : “Torres-Straße”; Torres Strait, 

between Australia and New Guinea; 20 fathoms 

(36m), dredged. 


on 6slides; Thiele fec. (labelled “CCCXLIII a–f”, with 

no. of “Vermes” Department “5495 a–f”). 

C o m m e n t s : Type species, by original designation, 

of Thiele’s new genus Notomenia. Re-examination 

and re-description in Salvini-Plawen 

(2004) leads to the classification within the new 

family Notomeniidae due to its anatomical organization 

unrelated to any other genus within the 

framework of the hitherto established families. 

eisigi Thiele, 1894 – Rhopalomenia 

Rhopalomenia eisigi Thiele, 1894: 269. 



Ty p e l o c a l i t y : “Neapel”; Mediterranean Sea: 

Gulf of Naples. 

Ty p e m a t e r i a l : ZMB Moll. 105.403a, holotype; serial sections 

on 9 slides (labelled “CCCX a–d vorn” and “CCCX a– 

e hinten”); 

ZMB Moll. 105.403b, part of holotype. 

C o m m e n t s : A synonym of Rhophalomenia 

aglaopheniae (Kowalevsky & Marion, 1887), as 

was already assumed by Thiele (1894: 269) and 

confirmed by Nierstrasz & Stork (1940: 74) and 

Salvini-Plawen (1967). The serial sections of the 

holotype were given for comparative study to 

Nierstrasz, later transferred to and re-discovered 

in the Zoological Museum in Amsterdam. The 

material was returned in November 2004 to the 

ZMB; for more details see below under Amphimenia 

grandis. 

gauszi Salvini-Plawen, 1978 – Pruvotina 

Pruvotina gauszi Salvini-Plawen, 1978: 144–145. 

Ty p e l o c a l i t y : “Antarktis, Winterstation am 


Gauss at 66 2 0 S, 89 38 0 E; DSE; leg. Vanhöffen. 

Ty p e m a t e r i a l : ZMB Moll. 105.394, holotype, serial section 

on 1 slide Thiele fec. (labelled “ad Q 1923d, Pruvotina 

spinosa juv. Thiele”). 

C o m m e n t s : This material was originally described 

by Thiele (1913a) as juvenile of Pruvotina 

spinosa Thiele, 1913a and determined as the 

above taxon by Salvini-Plawen (1978: 144–145). 

Note that the serial sections (ZMB 105.394), originally 

labelled “d” only represent the posterior 

part of the animal. 

gaussiana Thiele, 1913a – Acanthomenia 

Acanthomenia gaussiana Thiele, 1913a: 62, pl. IV, fig. 7a; 

textfig. 2. 

Ty p e l o c a l i t y : “Antarktis”; Antarctic: DSE, 

at 65 270S, 80 330E, in depth of –3398 m, March 



on 2 slides; Thiele fec. (labelled “Q 1928a–b, 3398 m”). 

C o m m e n t s : Type-species by monotypy of 

Thiele’s (1913a) new genus Acanthomenia. Revision 


glacialis Thiele, 1913a – Nematomenia 

Nematomenia glacialis Thiele, 1913a: 40, pl. IV, figs 3, 21–22; 

pl. V, fig. 1.




Gauss at 66 2 0 S, 89 38 0 E; DSE, Twist 385 m, Jun 

16, 1902; 380–385 m, from July 31 to December 

19, 1902; Twist 380 m, from July 01, 1902 to January 


Ty p e m a t e r i a l : ZMB Moll. 105.381a, lectotype (by present 

designation), with serial section on 5 slides, Thiele fec. 

(labelled “Q 1919a–c”, and “Q 1919d–e”); 

ZMB Moll. 105.381b, paralectotypes in 14 vials (1. vial: 

31. VII. 1902, WSt 380 m, one part of specimen; 2. vial: 

16. VI. 1902, Twist 385 m, one specimen; 3. vial: 19. XII. 

1902, 385 m, one specimen; 4. vial: 07. I. 1903, Twist 380 m; 

vial 5 –14, with a total of n ¼ 9 specimens, all without labels 

or date); 

ZMB Moll. 105.381c, paralectotypes, 2 Paraffin block (labelled 

“glacialis 31. 07. 1902”); 

ZMB Moll. 105.381d, paralectotypes, 2 Paraffin blocks. 

C o m m e n t s : Re-description by Salvini-Plawen 

(1978: 40). Thiele (1913a: 40, Taf. V, Fig. 1) 

clearly indicated one fully adult animal, here given 

as lectotype. 

grandis Thiele, 1894 – Neomenia 

Neomenia grandis Thiele, 1894: 223. 




sections on 52 slides (labelled “CCCI a–z2 vorn”, Thiele 

Dresden 1893; “CCCII a–b”; “CCCIII a–r Hinterende”; 

“CCCVI a–f”); 

ZMB Moll. 105.387b, part of holotype (ca. 4 cm) in ethanol. 

C o m m e n t s : Re-examined by H. A. Stork (in 

Nierstrasz & Stork 1940: 43) who already suggested 

not to distinguish N. grandis Thiele, 1894 

from N. carinata Tullberg, 1875. They placed 

grandis in synonymy of N. carinata Tullberg, as 

did Salvini-Plawen (1986, 1997b). The missing serial 

sections of the holotype (and other material 

of Thiele) were given for comparative study to 

Nierstrasz and later transferred together with 

Amphimenia neapolitana and Rhopalomenia eisigi 

to the collection of the Zoological Museum in 

Amsterdam. It was sent back to the ZMB thankfully 

by Robert Moolenbeek in November 2004 

after uncovering the post-WW II odyssee of this 

material (see more under A. neapolitana). 

intermedia Thiele, 1913a – Metamenia 

Metamenia intermedia Thiele, 1913a: 52, pl. IV, fig. 9; pl. VII, 

figs 1–2; pl. VIII, figs 16–21. 



Gauss at 66 2 0 S, 89 38 0 E; DSE, 380 m, Jun 14, 

1902; Twist 385 m, November 09, 1902; December 


Ty p e m a t e r i a l : ZMB Moll. 105.399a, part of lectotype 

(by present designation), serial sections on 3 slides. Thiele 

fec. (labelled “a–b vorn”, “1924a–b, 09. XI. 1902”; and “a 

hinten”, “1924c, 09. XI. 1902”); 

ZMB Moll. 105.399b, part of lectotype (by present designation); 

labelled “Th., 09. XI. 1902, Twist 385 m”); 

ZMB Moll. 105.399c, paralectotypes in 3 vials, in alcohol 

(1. vial: “? M. intermedia Th., Gauss-St[ation]”, one specimen; 

2. vial: “? M. intermedia, 14. VI. 1902”, one specimen; 

3. vial: “Proneomenia intermedia, 17. XII. 1902”, one specimen). 

C o m m e n t s : Type-species, by monotypy, of 

Thiele’s new genus Metamenia. Re-description in 

Salvini-Plawen (1978: 149–150). 

kerguelensis Salvini-Plawen, 2005 – 

Ocheyoherpia 

Ocheyoherpia kerguelensis Salvini-Plawen, 2005: 100, figs 1–4. 

Ty p e l o c a l i t y : “Kerguelen”; South of the antarctic 

circle; France, Antarctic-Kerguelen Islands, 

DTE, station 160. Unfortunately, in the 1899 

compilation of station numbers for the DTE expedition 

on board “Valdivia” no precise coordinates 

were given, but only a note that says “Gazelle 

Basin” and “3.5–4.0 C surface temperature”. 

Ty p e m a t e r i a l : ZMB Moll. 105.395a, holotype, serial sections 

on 3 slides and one spicule slide; Thiele fec. (labelled 

originally by Thiele as “Q 3030a–c”, plus spicule slide “Q 

3030 d”); 

ZMB Moll. 105.395b, paratypes, Thiele fec.; 3 specimens 

(two of them, with 0.6 and 0.5 mm juveniles, the other juvenile 

with 1.15 mm, decalcified without calcerous bodies/spicules). 

The latter specimen is illustrated in Fig. 2, prior to 

histological sections (on n ¼ 10 slides) done by LvS-P in October 

2004. 

C o m m e n t s : This taxon was only provisionally 

named kerguelensis by Johannes Thiele, as it is 

evident from the labels accompagning the ZMB 

material, but was never described in publication 

by this author, thus providing only a manuscript 

name without nomenclaturial bearings. Therefore, 

the taxon was recently described as a species 

of the genus Ocheyoherpia, now formally 

named by Salvini-Plawen (2005) utilizing the 

name “kerguelensis” as originally suggested by 

Thiele. The taxon is tentatively transferred to 

ordo Sterrofustia, family Phyllomeniidae. 

neapolitana Thiele, 1889 – Amphimenia 

Proneomenia neapolitana Thiele, 1889: 429 (footnote). 

Proneomenia (Amphimenia) neapolitana Thiele, 1894: 244. 


158 

Fig. 2. Habitus of one juvenile of Ocheyoherpia kerguelensis 

Salvini-Plawen, 2005 (ZMB 105.395b; paratype); the complete 

material of this new taxon, including three juvenile animals, 

was originally labelled “Pruvotina kerguelensis“ by 

Thiele, but only recently validly named and described by Salvini-Plawen 

(2005); see text for more details. Scale ¼ 0.5 mm. 




on 14 slides, from animal (c. 3 cm in length and 1.5 mm 

thick according to Thiele (1894: 244)); Thiele fec. (labelled 

“CCII a–o”). 


Amphimenia Thiele, 1894; as new subgenus of 

Proneomenia Hubrecht, 1880. Currently transferred 

to the Amphimeniidae. This taxon, described 

together with Neomenia grandis and 

Rhopalomenia eisigi by Thiele (1894), was long 

missing from the ZMB collection as was also 

Proneomenia vagans. Uncovering the odyssee of 

all this material from the available evidence, we 

reconstruct that it was given by Thiele in the late 

1920th/early 1930th on loan for the preparation 

of the Naples monograph to H. F. Nierstrasz at 

the University Museum in Utrecht (Nierstrasz in 

Nierstrasz & Stork 1940: 1). After Nierstrasz’ 

death (Sept. 1937) the material remained in 

Utrecht until the university stopped its systematic 

research. Thiele’s material was sent in 1956 to 

the then curator at the Zoological Museum in 

Amsterdam, T. Van Bentem-Jutting (while other 

material from Utrecht was transferred at the 

same time to the Natural History Museum in 

Leiden). In search of the type-material, it was 

traced out in May 1977 by LvS-P together with 

S. Van der Spoel, the then curator at the ZMA, 

on the occassion of a visit to the collection of 

the ZMA. LvS-P made exact notes about the 

boxes in which Thiele’s original material in the 

ZMA were deposited. Triggered by this forwarded 

information, the material was finally rediscovered 

in the ZMA and thankfully sent back 



to the ZMB in November 2004 by Robert Moolenbeek. 

papilligera Thiele, 1913a – Sandalomenia 

Sandalomenia papilligera Thiele, 1913a: 41, pl. IV, figs 4, 24; 

pl. V, figs 2–5. 



Gauss at 66 20S, 89 380E; DSE, 385 m, from November 

22 to 24, 1902; Twist 380 m, January 28, 

1903; leg. Vanhöffen. 

Ty p e m a t e r i a l : ZMB Moll. 105.384a, syntype, serial sections 

on 7 slides; Thiele fec. (labelled “Q 1920 a–d vorn” 

and “Q 1920 e–g hinten”); 

ZMB Moll. 105.384b, part of syntype; 

ZMB Moll. 105.384c, syntype, 1 specimen in alcohol (labelled 

“28. I. 1903, Twist 380 m); 

ZMB Moll. 105.384d, syntypes, 2 Paraffin blocks. 

C o m m e n t s : Type-species, by subsequent designation 

(Salvini-Plawen 1978: 48), of the new 

genus Sandalomenia Thiele, 1913a. Re-description 


prisca Thiele, 1906 – Archaeomenia 

Archaeomenia prisca Thiele, 1906: 315, pl. XXVIII. 

Ty p e l o c a l i t y : “Agulhas Bank, am südlichen 

Teil”; southern part of Agulhas bank, off South 

Africa; DTE, station 110 (35 90S, 18 32.80E; trawl 

564 m, October 4, 1898. 

Ty p e m a t e r i a l : ZMB Moll. 59.910a, syntype, serial sections 

on 8 slides (labelled “a–c vorn”, “Q 5078/1–3, Valdivia” 

and “a–e hinten”, “Q 5078/4–8, Valdivia”); 

ZMB Moll. 59.910b, syntype, serial sections on 5 slides (labelled 

“a–e Sagitalschnitte”, “Q 5078/9–13); 

ZMB Moll. 59.910c, syntypes, 2 specimens in veil. 


Thiele’s new genus Archaeomenia. Re-examined 

in Salvini-Plawen & Paar-Gausch (2004). 

protecta Thiele, 1913a – Nematomenia 

Nematomenia protecta Thiele, 1913a: 39, pl. IV, figs 2, 19–20. 



Gauss at 66 20S, 89 380E; DSE, 385 m, July 9, 

1902; 385 m, December 2–4, 1902; leg. Vanhöffen. 

Ty p e m a t e r i a l : ZMB Moll. 105.382, syntypes, n ¼ 6 

whole mounts on 3 slides; Thiele fec. (2 slides labelled 

“9. VII. 1902, 385 [m]” and one slide labelled “2.– 

4. XII. 1902, 385 m”).


C o m m e n t s : Note that only the organisation of 

the scales of the mantle is known (Thiele 1913a: 

39–40; Salvini-Plawen 1978: 45, fig. 28). 

providens Thiele, 1913a – Pruvotina 

Pruvotina providens Thiele, 1913a: 48, pl. IV, fig. 7; pl. V, 

figs 15–17; pl. VI, figs 1–3; pl. VIII, figs 10–11. 



Gauss at 66 20S, 89 380E; DSE 385 m Breitnetz 

and Twist, December 17, 1902; 385 m Twist, January 

28, 1903; 350 m Twist, February 8, 1903; leg. 

Vanhöffen. 


sections on 3 slides, Thiele fec. (labelled “a–b Vorderende”, 

“ad Q 1922 a–b”; and “a Hinterende”, “ad Q 1922 c”); 

ZMB Moll. 105.397b, part of holotype (labelled 

“17. XII. 1902, 385 m”); 

ZMB Moll. 105.397c, paratypes in 6 vials (1. vial: 

08. II. 1903, Twist 350 m, 1 specimen; 2. vial: without label, 1 

specimen; 3. vial: 17. XII. 1902, 385 m, 1 specimen; 4. vial, 

28. I. 1903, Twist 385 m, part of specimen; 5. vial: 

17. XII. 1902, Twist 385 m, 11 specimens; 6. vial: 2 specimens); 

ZMB Moll. 105.397d, paratype, 1 Paraffin block. 

C o m m e n t s : Re-description by Salvini-Plawen 

(1978: 119–120). 

spinosa (Thiele, 1913a) – Labidoherpia 

[Pruvotina] 

Pruvotina spinosa Thiele, 1913a: 50, pl. IV, fig. 8; pl. VI, 

figs 4–7; pl. VIII, fig. 12–15. 



Gauss at 66 20S, 89 380E; DSE; 385 m; leg. Vanhöffen. 

Ty p e m a t e r i a l : ZMB Moll. 105.398a, lectotype (by present 

designation), serial sections on 3 slides; Thiele fec. (labelled 

“a – vorn”, “ad Q 1923 a”; and “a–b”, “ad Q 1923 

b–c; b ¼ 28. I. 1903”); 

ZMB Moll. 105.398b, paralectotypes in n ¼ 10 vials (1. vial: 

17. XII. 1902, 385 m, 1 specimen, 2. vial: 14. IV. 1902, 385 m, 

1 specimen; 3. vial: 12. I. 1903, 380 m, 6 specimens; 4. vial: 8 

specimens; 5. vial: 14 specimens; 6. vial: 1 specimen; 7. vial: 

46 specimens; 8. vial: 12. X. 1902, 385 m, 1 specimen; 9. vial: 

22. I. 1903, 2 specimens, one with long spiculae; 10. vial: 

09. VII. 1902, 385 m, 2 specimens); 

ZMB Moll. 105.398c, paralectotypes, 2 Paraffin blocks (labelled 

“10. 01. 1903 juv.” and “28. 01. 1903”). 

Comments: Pruvotina spinosa Thiele, 1913a 

has been assigned by Salvini-Plawen (1978) as 

type species of the new genus Labidoherpia. 

Note that of the originally seven slides with serial 

sections assigned to this species by Thiele 

(1913a), only three (ZMB Moll. 105.398a) are 

here kept with spinosa, while the other slides, la- 

belled “d” and “e–g”, respectively, were assigned 

to Pruvotina gauszi Salvini-Plawen, 1978 

and Gephyroherpia antarctica Salvini-Plawen, 

1978 (for details see there). 

squamosa Thiele, 1913a – Nematomenia 

Nematomenia squamosa Thiele, 1913a: 40, pl. IV, fig. 23. 



Gauss at 66 2 0 S, 89 38 0 E; DSE, 385 m, December 


Ty p e m a t e r i a l : ZMB Moll. 105.383, holotype, whole 

mount on slide; Thiele fec. (“labelled 17. XII. 1902; 385 m”; 

one of three animals on the microscopic slide is circled with 

black marking, indicated as “type”; the other two are given 

as “N. glacialis Thiele, Antarktis, Gauss Winterstation, D. 

Südp. Exp., Vanhöffen S, Q1931”). 

C o m m e n t s : Only the mantle scales are known 

(Thiele 1913a: 39–40; Salvini-Plawen 1978: 45). 

thulensis (Thiele, 1900) – Thieleherpia 

[Proneomenia] 

Proneomenia thulensis Thiele, 1900: 111, pl. V. 

Ty p e l o c a l i t y : “Spitzbergen, Station 18 

(80 80N, 16 550E), 480 m”; Spitsbergen, Hinlopen 

Strait, at northern entrance; leg. Römer & 

Schaudinn Expedition, June 1, 1898; on fine, blue 

mud with only few small stones in front of a glacier. 

Locality data here added according to entries 

given in Römer & Schaudinn (1900: 40) in 

a compilation of dredge stations. 


section on 14 slides (labelled “a–f” for anterior part, and 

“a–f” for posterior part); 

ZMB Moll. 105.405b, part of holotype, in ethanol. 

C o m m e n t s : Re-description and classification 

as Thieleherpia thulensis (Thiele, 1900) in Salvini- 

Plawen (2004: 86–88), with the additional examination 

of another specimen from off northeastern 

Iceland (ZMUC). Due to features of the 

ventral foregut glandular organs the species was 

stated by Salvini-Plawen (1978: 231) not to belong 

to Proneomenia and finally placed together 

with Rhipidoherpia within the Rhipidoherpiidae 

(see Salvini-Plawen 2004: 88). 

tricarinata (Thiele, 1913a) – Dorymenia 

[Proneomenia] 

Proneomenia tricarinata Thiele, 1913a: 54, pl. IV, fig. 10; 

pl. VII, figs 3–9; pl. VIII, figs 22, 24–30. 


160 



Gauss at 66 2 0 S, 89 38 0 E; DSE, (17. 07. 1902– 

12. 10. 1902 Twist 385 m, 09. 11. 1902–22. und 

24. 11. 1902 Twist 385 m, 17. 12. 1902 Breitnetz 

385 m – 23. 12. 1902 Twist 385 m, 12. 01. 1903 

Twist 385 m – 24. 01. 1903 Twist 380 m, 

31. 01. 1903 Twist 380 m – 08. 02. 1903 385 m, 

15. 02. 1903 Twist 400 m), leg. Vanhöffen. 


sections on 11 slides, Thiele fec. (n ¼ 5 slides labelled 

“a–e vorn”, “Q 1925 k–o”; n ¼ 6 slides labelled “a–f hinten”, 

“Q 1925 a–f”); 

ZMB Moll. 105.413b, part of holotype (labelled 

“08. II. 1903, 385 m”); 

ZMB Moll. 105.413c, paratypes in 9 vials in ethanol (1. vial: 

12. I. 1903, Twist 385 m, 1 specimen; 2. vial: 23. XII. 1902, Twist 

385 m, 1 specimen; 3. vial: 22. þ 24. XI. 1902, Twist 385 m, 

1 specimen; 4. vial: 31. I. 1903, Twist 380 m, 1 specimen; 5. vial: 

15. II. 1903, Twist 400 m, 2 specimens; 6. vial: 09. XI. 1902, 

Twist 385 m, 1 specimen; 7. vial: 17. XII. 1902, Breitnetz 385 m, 

1 specimen; 8. vial: 17. XII. 1902, 1 specimen; 9. vial: 

12. X. 1902, Twist 385 m, 1 specimen; 10. vial: 09. XI. 1902, 

Twist 385 m, 1 specimen); 

ZMB Moll. 105.413d, paratypes, 2 Paraffin blocks. 

C o m m e n t s : Re-description and classification 

as Dorymenia tricarinata (Thiele, 1913a) in Salvini-Plawen 

(1978: 257–260). 

valdiviae Thiele, 1902b – Proneomenia 

Proneomenia valdiviae Thiele, 1902b: 167, pl. XXIII. 

Type locality: “Küste von Ostafrika, N von 

Sansibar; am Netz hängend, Sublimat”; DTE, 

station 249 (3 70S, 40 45.80E); offshore E Africa, 

north of Sansibar; trawl 748 m, March 23, 1899. 


sections on 15 slides (labelled “a–i vorn”; and “a–f hinten”); 

ZMB Moll. 105.406b, part of holotype, in ethanol (labelled 

“5049”, referring to a number in the “Vermes” Collection). 

Section B – The non-type material in the Malacozoological 

Collection of the ZMB 

Solenogastres 

aglaopheniae (Kowalevsky & Marion, 1887) – 

Rhopalomenia 

Proneomenia aglaopheniae Kowalevsky & Marion, 1887: 1–76. 

L o c a l i t y : Mediterranean Sea: Gulf of Neaples; 

Italia. 

M a t e r i a l : ZMB Moll. 105.402a, serial sections on 12 slides, 

Thiele fec. (labellel “CCCXV a–d”, and “a–e”). 



C o m m e n t s : Type species of the genus. Type 

locality: Banyuls-sur-mer, France, Mediterranean 

coast. Known distribution: European shelf from 

Peleponnes (Greece) to Scottland; in 50–137 m. 

The material now housed again in the ZMB has 

been resent from the ZMA together with other 

parts of Thiele’s material from Neaples such as, 

for example, Amphimenia neapolitana (see under 

this species) and Rhopalomenia eisigi (Thiele, 

1894) which is synonymized with Rh. agalopheniae 

(see under the former species). 

antarctica Salvini-Plawen, 1978 – 

Gephyroherpia 

Gephyroherpia antarctica Salvini-Plawen, 1978: 115. 

L o c a l i t y : “Antarktis: Winterstation am Gaussberg”; 

Antarctic: winter station at Mount Gauss 

at 66 2 0 S, 89 38 0 E; DSE; leg. Vanhöffen. 

M a t e r i a l : ZMB Moll. 105.393, serial sections on 3 slides, 

Thiele fec. (labellel “e–g”). 

C o m m e n t s : The type material for this species 

is deposited in the USNM (Smithsonian Institution) 

in Washington, DC (USA). Type locality: 

Ross Sea. The material in ZMB has been described 

as “Pruvotina spinosa juv.” by Thiele 

(1913a: 52); see Salvini-Plawen (1978: 118). 

banyulensis Pruvot, 1890 – Nematomenia 

Dondersia banyulensis Pruvot, 1890: 699–810. 

Myzomenia banyulensis (Pruvot, 1890) – Simroth 1893a. 




Thiele fec. (labellel “CLXXIV a–c”, anterior part of animal; 

and “CLXXII”, posterior part of animal); 

ZMB Moll. 105.379, specimen in ethanol; Thiele fec. (“Plymouth, 

Marien Biological Laboratory, Thiele rev.”). 

C o m m e n t s : Ty p e l o c a l i t y : Banyuls-surmer, 

France, Mediterranean coast. The actually 

known distribution ranges from Dalmatia (Adriatic 

Sea) to the Trondheimsfjord (Norway) at 31– 

300 m depth. The material now housed again in 

the ZMB (105.426) has been resent from the 

ZMA together with other parts of Thiele’s material 

from Neaples such as, for example, Amphimenia 

neapolitana (see under this species). 

hoffmani Salvini-Plawen, 1978 – Dorymenia 

Dorymenia hoffmani Salvini-Plawen, 1978: 247.


L o c a l i t y : “Antarktis: Winterstation am Gaussberg”; 

Antarctic: winter station at Mount Gauss 

at 66 2 0 S, 89 38 0 E; DSE, leg. Vanhöffen. 

M a t e r i a l : ZMB Moll. 105.420, serial section on 9 slides 

(labelled by Thiele as “Q 1926 a–k”); see Fig. 1. 

C o m m e n t s : This material has been assigned 

originally to Proneomenia antarctica by Thiele 

(1913a), but was separated from this species by 

Salvini-Plawen (1978: 247). The type material of 

the latter is deposited in the Zoological Museum 

of the University Uppsala (Schweden), as serial 

section on slides. 

vagans Kowalevsky & Marion, 1887 – 

Dorymenia 

Proneomenia vagans Kowalevsky & Marion 1887: 1–76. 




Thiele fec. (labellel “CLXVII a–c”). 

C o m m e n t s : The species had been redescribed 

by Thiele (1894). It is known from the western 

Mediterranean, between Marseille and Neaples, 

from depth down to 20–30 m. The serial sections 

of the ZMB were given on loan for comparative 

study to Nierstrasz, later transferred to the Zoological 

Museum in Amsterdam and returned in 

November 2004 to the ZMB; for more details 

see below under Amphimenia neapolitana. 

Caudofoveata 

This group (also named Chaetodermomorpha), 

is traditionally subdivided into three families, Limifossoridae 

(with Metachaetoderma Thiele, 

1913), Prochaetodermatidae (with Prochaetoderma 

Thiele, 1902) and Chaetodermatidae; see e.g. 

Salvini-Plawen (1971, 1975, 1985) and Scheltema 

(1981). 

Chaetodermatidae Ihering, 1876 

The family represents a cosmopolitan taxon comprising 

three genera with a total of 80 species described, 

mostly from the shelf regions and the 

continental slopes. It includes with Chaetoderma 

productum being up to 14 cm in length the greatest 

Caudofoveate. 

productum Wirén, 1892 – Chaetoderma 

Chaetoderma productum, Wirén 1892: 86. 

Crystallophrisson productum (Wirén) – Thiele (1932). 

Ty p e l o c a l i t y : Kara Sea (71 20 0 N, 59 58 0 E); 

62 fathoms (¼ –114 to 115 m depth), leg. Sept. 

9, 1883; “Dijmphna”-Expedition (Stat. 188). 

Ty p e m a t e r i a l : Lectotype and paralectotype in ZMC, 

vide Salvini-Plawen (1975). 

A d d i t i o n a l m a t e r i a l : ZMB Moll. 105.377, 

n ¼ 3 specimens. Topotypical material, originally 

labelled as “C. nitidulum”, later changed to C. 

productum referring to A. Wirén’s description of 

a new arctic species (see Comments below). 

C o m m e n t s : There are three locations known 

from the Kara Sea where C. productum was 

caught with several specimens each during the 

Danish-International “Dijmphna” Expedition 

from August 1882 to August 1883 (see Paulsen 

1884). The three specimens in the ZMB are apparently 

part of the material originally collected 

at the locus typicus in the Kara Sea by this expedition, 

albeit they are not of the syntype series 

that Wirén based his description on. While we 

were unable to reconstruct how the ZMB got 

part of the material from the Dijmphna” Expedition 

in the first place, Wirén (1892: 8) explicitly 

stated in his introduction that in summer 1891 he 

had a chance to see this material in the collections 

of both the Copenhagen Museum and the 

Berlin Museum. However, he continued that for 

his description in 1892 of a new species from the 

Kara Sea, distinct from C. nitidulum, he received 

only the material from the Copenhagen Museum 

through Dr. G. M. R. Levinsen. 

Thiele (1902c: 273) mentioned dissections 

done on a specimen from the Götting Zoological 

collection, ded. Ehlers, and the publication date 

is before Thiele starting to work as curator in 

the ZMB. However, this Götting specimens represents 

C. nitidulum; its origin is unknown, but 

most likely comes from the North Sea or the 

Kattegatt. The series of C. productum (ZMB 

105.377) was returned to the Malacological collection 

from the “Vermes” department in Dezember 

1930. 

This species was originally considered by Odhner 

(1921) as subspecies of Chaetoderma nitidulum 

(Lovén, 1845), which is, however, not known 

from a holotype or syntype specimen (see Salvini-Plawen 

1984; Ivanov & Scheltema 2000). For 

a new characterisation see Salvini-Plawen (1975). 


162 

Prochaetodermatidae Salvini-Plawen, 1968 

The family represents a cosmopolitan taxon 

based upon the single genus Prochaetoderma, 

Thiele, including today six (poorly defined) genera 

or subgenera with 25 named species. They 

are of small size, less than 5 mm, and mostly belong 

to the deep-sea fauna (Salvini-Plawen 1992; 

Scheltema & Ivanov 2000, 2004; Ivanov & Scheltema 

2002). Thiele (1902c: 275) introduced Prochaetoderma 

for Chaetoderma radulifera Kowalevsky 

(1901), according to the description by 

this author, from the Marmara Sea. Today no 

original material is retained. 

Section C – List of nominal taxa housed in the 

Malacozoological Collection of the Museum of 

Natural History Berlin (ZMB), arranged according 

to current systematic affiliation 

The present catalogue comprises a total of 31 

aplacophoran taxa, including six non-type taxa. 

Of these taxa n ¼ 23 were named and described 

originally by Johannes Thiele, with an additional 

four by Salvini-Plawen (1978, 2005), which are 

partly based on material first described by 

Thiele. With the exception of the caudofoveate 

Chaetoderma productum Wirén, 1892 all aplacophorans 

in the ZMB are Solenogastres. They are 

assigned to 20 genera, following the classification 

suggested by Salvini-Plawen (1978). For a characterization 

of the genera see e.g. Salvini-Plawen 

(1967). 

Currently, 29 out of the 31 aplacohoran taxa 

are considered valid species, two are considered 

synonyms (i.e. Neomenia grandis of N. carinata, 

and Rhopalomenia eisigi of R. aglaopheniae). A 

third, Ocheyoherpia kerguelensis Salvini-Plawen, 

2005 was newly described, based on material 

provisionally named but not described and published 

as valid name by J. Thiele. 

The following list is considered the most recent 

classification on aplacophoran molluscs, and 

essentially follows that suggested by Salvini-Plawen 

(1978) with additions in Salvini-Plawen 

(2004). Type species of genera are given in bold, 

synonyms are given as “¼”; original generic assigments 

are in square parentheses. Numbers refer 

to footnotes below. 


A – Solenogastres 

Aplotegmentaria 

Ordo Pholidoskepia 

Dondersiidae 

Nematomenia arctica Thiele, 1913b 

Nematomenia banyulensis (Pruvot, 1890) 

[Dondersia] 

Nematomenia glacialis Thiele, 1913 

Nematomenia (?) protecta Thiele, 1913 

Nematomenia (?) squamosa Thiele, 1913a 

Sandalomeniidae 

Sandalomenia papilligera Thiele, 1913 

Sandalomenia carinata Thiele, 1913a 

Incertae sedis 

Pholidoherpia cataphracta (Thiele, 1913a) 

[Lepidomenia] 

Ordo Neomeniamorpha 

Hemimeniidae 

Archaeomenia prisca Thiele, 1906 

Neomeniidae 

Neomenia grandis Thiele, 1894 ¼ N. carinata 

Tullberg, 1875 

Pachytegmentaria 

Ordo Sterrofustia 

Phyllomeniidae 

Phyllomenia austrina Thiele, 1913a 

Ocheyoherpia kerguelensis Salvini-Plawen, 2005 

Ordo Cavibelonia1 Pararrhopaliidae2 Gephyroherpia antarctica Salvini-Plawen, 1978 

[Pruvotina spinosa Thiele, partim] 

Pruvotina gauszi Salvini-Plawen, 1978 [Pruvotina 

spinosa Thiele, partim] 

Pruvotina providens Thiele, 1913 

Labidoherpia spinosa (Thiele, 1913a) 

[Pruvotina] 

Metamenia intermedia Thiele, 1913a 

Acanthomeniidae 

Acanthomenia gaussiana Thiele, 1913a 

Notomeniidae 

Notomenia clavigera Thiele, 1897 

Rhopalomeniidae 

Rhopalomenia aglaopheniae (Kowalevsky & 

Marion, 1887) [Proneomenia] 

Rhopalomenia eisigi Thiele, 1894 ¼ R. aglaopheniae 

(Kowalevsky & Marion, 1887) 

1 Salvini-Plawen (1978, 2004) suggested to distinguish 11 families, of which 7 are represented here by type material extant 

in the ZMB 

2 Family also named Pruvotinidae or Perimeniidae; morphologically diverse, but ill-defined group according to Scheltema 

(2001: 16) 



Strophomeniidae 

Anamenia amboinensis (Thiele, 1902a) 

[Proneomenia] 

Epimeniidae 

Epimenia australis (Thiele, 1897) [Proneomenia] 

Proneomeniidae 

Proneomenia valdiviae Thiele, 1902b 

Dorymenia antarctica (Thiele, 1913a) 

[Proneomenia] 

Dorymenia hoffmani Salvini-Plawen, 1978 

[Proneomenia antarctica Thiele, partim] 

Dorymenia tricarinata (Thiele, 1913a) 

[Proneomenia] 

Dorymenia vagans (Kowalevsky & Marion, 

1887) [Proneomenia] 

Amphimeniidae 

Amphimenia neapolitana (Thiele, 1889) 

[Proneomenia] 

Rhipidoherpiidae 

Thieleherpia thulensis (Thiele, 1900) 

[Proneomenia] 

B – Caudofoveata: 

Ordo Chaetodermomorpha 

Chaetodermatidae 

Chaetoderma productum Wirén, 1892 

Section D – List of nominal taxa named by 

Thiele, but representatives not extant in the Museum 

of Natural History Berlin (ZMB) 

The following taxa have been named by Thiele, 

in addition to those listed alphabethically under 

section A and B (see also systematic section C). 

However, material representing these genus-level 

taxa is not extant (i.e. not found in the ZMB collection). 

Compilation according to Boss & Bieler 

(1991) who listed a total of 11 genus-level names 

introduced by Thiele. 

Solenogastres 

Dondersiidae 

Heathia Thiele, 1913a; as new genus. Type-species, 

by monotypy: Ichthyomenia porosa 

Heath, 1911. 

Rhopalomeniidae 

Pruvotia Thiele, 1894; as new genus. Type-species, 

by monotypy: Proneomenia sopita Pruvot, 

1891. 

Caudofoveata 

Limifossoridae 

Metachaetoderma Thiele, 1913a; as new genus. 

Type-species, by monotypy: Chaetoderma challengeri 

Nierstrasz, 1903. 

Prochaetodermatidae 

Prochaetoderma Thiele, 1902; as new genus. 

Type-species, by monotypy: Chaetoderma raduliferum 

Kowalevsky, 1901. 

Acknowledgements 

We are most grateful to Robert Moolenbeck for altruistically 

searching for and locating in the ZMA three of Thiele’s aplacophoran 

types from Neaples that were missing after being 

sent out on loan to Nierstrasz in Utrecht before WW II. We 

thank Frank Köhler for his help with inventorizing the aplacophoran 

type material of the ZMB and for valuable comments 

on the manuscript, as well as Jeroen Goud (Naturalis, 

Leiden) for additional information. We thank, as always, 

Mrs. Ingeborg Kilias for her help with literature research, 

and Birger Neuhaus for additional hints to the history of 

these enigmatic “vermiform” molluscs partly deposited and 

misplaced among “Vermes” in the department under his 

care; for helping to locate aplacophoran material in the latter 

department we are thankful to Mrs. K. Kämpf and K. Meschter. 

We are indepted to Michael Ohl (ZMB) and an anonymous 

referee for some valuable comments that helped to improve 

the manuscript. 

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