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

Aplacophoran in the Natural History Museum . An annotated catalogue of Thiele’s type specimens, with a brief review of “Aplacophora” classification

Matthias Glaubrecht*,1, Lothar Maitas1 & Luitfried v. Salvini-Plawen**,2

1 Department of Malacozoology, Museum of Natural History, Humboldt University, Invalidenstraße 43, D-10115 Berlin, 2 Institut fu¨ r Zoologie, Universita¨t 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, (¼ Neomeniomorpha), (¼ Chaetodermomorpha), Aculifera, Amphineura, Johannes Thiele, Ernst Vanho¨ ffen, 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 con- stituting a monophyletic clade called Aplacophora Von Ihering, 1876, although evidence is available to the contrary, suggest- ing the latter to be a paraphyletic grade. Accordingly, the hitherto accepted “Aplacophora” may consist of two Recent, diphy- letic 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 rather than molluscs resulted in the deposition of aplacophoran material of the ZMB (and hence the late re- discovery 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 , all from the very same type locality near the Antarctic Gaussberg volcano at 6620S, 89380E, collected by the expedition’s biologist Ernst Vanho¨ ffen.

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Introduction tection found in most other molluscs, their slen- der body is provided with a cover of a Aplacophorans are vermiform (i.e. worm- chitinous cuticle invested with numerous aragoni- shaped), shell-less, often deep-sea and bottom- tic integumental (i.e. “spicules”, “scales” feeding with a phylogenetically basal po- and “needles”). In addition, they exhibit epider- sition in the Mollusca. Instead of the shell pro- mal papillae (Solenogastres), a typical tetraneu-

* Corresponding author: e-mail: [email protected] ** e-mail: [email protected]

# 2005 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim 146 Glaubrecht, M. et al., Aplacophoran Mollusca in the Natural History Museum Berlin ral nervous system, a with different num- early evolution; for a recent discussion in a bers of teeth per row, as well as – uniquely broader molluscan framework see e.g. Salvini- among molluscs – a reproductive system with Plawen & Steiner (1996), Salvini-Plawen (1990) gonads emptying into the pericard and secundary and Lindberg & Ponder (1996). gametoducts; for detailed accounts on anatomical The systematics within aplacophoran molluscs characters in aplacophoran molluscs see the gen- is also partly unresolved. According to tradi- eral treatments e.g. by Simroth (1893b), Thiele tional knowledge the aplacophoran molluscs (1913c, 1925), Hoffmann (1929), Fischer-Piette & comprise two higher level taxa (subclasses or Franc (1960), Hyman (1967), Salvini-Plawen proper classes) with strongly distinct anatomical (1967, 1971, 1972, 1985), Scheltema et al. (1994) habitus and habits. In most classifications these and Scheltema (1998). taxa are separated as Solenogastres Gegenbaur, Although aplacophoran molluscs are a small 1878 (sensu nomine Simroth, 1893) and Caudo- group of animals ranging from 1 mm to 30 cm, foveata Boettger, 1956. The naming and separa- and are only rarely preserved even in major mu- tion of both groups – that were first discovered seum collections, they actually represent a phylo- in Scandinavian waters in 1844 (Chaetoderma ni- genetically highly important group, with constitu- tidulum Love´n, 1845) and 1875 (Neomenia cari- ent taxa possessing morphological characters nata Tullberg), respectively – refers to the pre- considered essential for the reconstruction of the sence of a small foot within a ventral groove in evolution of Mollusca; see e.g. recent reviews Solenogastres (thus “Furchenfu¨ ßer” or “Bauch- and discussions in Salvini-Plawen (1967, 1972, furcher” in German) or its absence in Caudofo- 1985, 2003a), Scheltema (1978, 1993, 1996), Iva- veata (“Schildfu¨ ßer”). Solenogastres lack a cuti- nov (1996), Salvini-Plawen & Steiner (1996) and cular oral shield and posterior ctenidia, live Haszprunar (2000). mostly cnidarivore and even epizoic on However, the curious and unique mixture of colonies (such as Octocorallia and Hydrozoa) or plesiomorphic and derived features of aplaco- benthic and are hermaphrodtites. In contrast, a phorans have contributed to long-standing dis- cuticular oral or pedal shield and paired ctenidia putes as to the phylogenetic relationships not are present in the Caudofoveata which live only to other mollusc groups but also among benthic and burrowing in marine soft sediments these shell-less taxa that apparently modified where they move by hydrostatic action; the latter many of their anatomical characteristics in dra- are micro-omnivore to micro-carnivore (feeding matical ways. Ongoing controversies include the on debris, foraminiferans and other small organ- question of the validity of the Aculifera or Am- isms) and are of separate sex. phineura concepts (the latter going back to Iher- Unfortunately, our knowledge of the biology ing (1876)), that both propose Aplacophora Iher- of aplacophorans is still fragmentary. While only ing, 1876 and Polyplacophora Gray, 1821 as about 120 species of Caudofoveata are known so adelphotaxa, or the question whether Aplaco- far, about 240 species of Solenogastres have phora are monophyletic. been named, with new ones currently under de- While molecular phylogeny point to a diphy- scription. This relation in species number is also letic origin, it was unable to propose probable more or less reflected by the extant historical sister group relationships to other molluscan material in the ZMB where Solenogastres lar- groups (for reasons given below). Most currently gely dominate (see below). Undoubtedly, the lat- used text books in zoology (e.g. Go¨ tting 1985, ter taxon is not only specious but also more di- 1996; Brusca & Brusca 2003) and some mono- verse in respect to habitats used, which can be graphic malacological accounts (e.g. Scheltema benthic, epizoic, and meiofaunal or even burrow- 1996, 1998, 2001) thus continue to perceive ing. Aplacophora and Aculifera as monophyletic In recent years several new additions come taxa. This conservative view is largely ignoring from various regions of the world, as is evident available evidence for an alternative systematiza- from the descriptions of new taxa of different tion according to cladistic analyses that suggest Solenogastres from Norway (Handl & Salvini- the latter two taxa being paraphyletic assem- Plawen 2001, 2002), from the Mediterranean Sea blages (i.e. grades) only, both representing basal, (Salvini-Plawen 2003b) from the Western Indian independent off-shoots of Mollusca (Salvini-Pla- (Todt & Salvini-Plawen 2003), from the wen & Steiner 1996; Haszprunar 2000). The un- southern hemisphere (Salvini-Plawen & Paar- resolved phylogenetic issues currently compro- Gausch 2004) and from various other regions mise our understanding of molluscan origin and (Salvini-Plawen 2004), as well as Caudofoveata

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(Prochaetodermatidae) of the Atlantic Ocean This re-examination of ZMB material resulted, and Mediterranean Sea (Scheltema & Ivanov for example, in the recent transfer and re-de- 2000) or the Indian and Pacific (Ivanov scription as type species of a new genus, as in & Scheltema 2002, 2004). Apart from the mono- the case of Thieleherpia thulensis (Thiele 1900) graph of antarctic-subantarctic Solenogastres by Salvini-Plawen (2004: 86), and in the erection with 75 new species (Salvini-Plawen 1978) an- of a new family Notomediidae, based on the re- other striking example for the recent increase in examination of the type species of the respective knowledge is the Australian region, where appar- genus Notomenia clavigera Thiele, 1897 (see Sal- ently a rich and diverse fauna exists in the south- vini-Plawen 2004: 89). The revision of earlier de- east of the continent with at least 32 species in scriptions in light of new material, collections 14 or more genera, some still awaiting descrip- and taxa described will eventually allow more tion (see Scheltema 2001). Prior to this recent accurate systematics at the generic and family survey only two species were recorded by Thiele level as shown, for example, recently in the Cla- (1897), viz. Notomenia clavigera and “Proneome- vibelonia (cf. Salvini-Plawen 2004). Again, this nia”(¼ Epimenia) australis of which the types underscores the importance of historical collec- are extant in the ZMB collection (see below). tions such as Thiele’s type material of aplaco- Irrespective of their rare representation in phorans. most collections, aplacophorans form a numeri- Therefore, the objective in the present paper cally small but consistent deep-sea faunal ele- is twofold. The Museum of Natural History of ment; some taxa are apparently common and Berlin (formerly Zoological Museum Berlin, sometimes form an abundant part of the shelf, ZMB) houses rich type material of aplacopho- bathyal, abyssal and hadal oceanic benthic rans, particularly Solenogastres, which are evi- macrofauna from depths of c. 200 m to 9000 m dently of basic importance for any systematic (see Belyaev 1966). Since representatives of both evaluation of these molluscs. This material stems aplacophoran taxa are found in deeper water on to a substantial degree from the systematic work the continental shelf and in the deep-sea down of the former curator of the Berlin Malacological to the abyssal zone and even at hydrothermal Collection, Johannes Thiele (1860–1935), who vents, they are generally not only less accessible was certainly one of the preeminent malacolo- than other molluscs; since they are often small, gists of the twentieth century. Following posi- with many being less than five millimeters (but tions as assistant in the invertebrate section of also reach exceptionally 300 mm), they certainly the Dresden Museum fu¨ r Tierkunde (1891– represent a neglected part of the Mollusca; for a 1895), Strassburg’s Zoological Institute (1895– review of the biology of aplacophorans see, for 1896) and Go¨ ttingen’s Zoological Institute example, Salvini-Plawen (1971, 1985) and Schel- (1896–1898), Thiele took a position as a lecturer tema (2001). at the Zoological Institute in Berlin in spring Not only recent discoveries of new taxa have 1898, before starting as a scientific collaborator spurred aplacophoran research, but also renewed or helper at the Zoological Museum in Berlin in interest in the phylogeny of metazoans in general spring 1899, where he became Research Assist- and mollusca in particular. Clearly, their systema- ant in 1900, then curator of the Crustacea Sec- tics with respect to the new findings is still in tion in 1903, before he finally took charge of the flux and continuously has to be adapted. How- Mollusk Section in 1905, after the death of ever, in view of the fact that traditional classifica- Eduard von Martens (1831–1904). Thiele tions did apparently neither reflect the phyloge- worked there also after his official retirement (in netic affinities among nor within the two major 1925) until his death in 1935. For a brief biogra- groups Solenogastres and Caudofoveata, the ac- phy, including a bibliography, see e.g. Bieler & tual systematics in general appears to be in ac- Boss (1989). cordance with computerized cladistic analysis The Solenogastres were among the molluscan (Salvini-Plawen 2003a). groups Thiele paid particular attention to, culmi- Nevertheless, this situation all the more nating in his revision of this taxon in Das Tier- strengthens the importance of historical museum reich (Thiele 1913c). Among the total of 291 material. The recent interest in aplacophoran genus-level taxa in the phylum Mollusca intro- classification often necessitates the re-examina- duced by Thiele, there are 11 aplacophoran tion of this archival material in particular of the names (see Boss & Bieler 1991), to which at many taxa of Solenogastres named by one of the least 24 new Solenogastres species names by former leading malacologists, Johannes Thiele. Thiele can be added according to the material

# 2005 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim 148 Glaubrecht, M. et al., Aplacophoran Mollusca in the Natural History Museum Berlin presented herein. First, we give an annotated in his final statement Thiele (1902c: 455) proposed catalogue of the ZMB type material with notes explicitly (and printed in bold) that “Solenogastres on the type specimens, including their original are a group of worms closely related to annelids status and references (i.e. author and biblio- and gordiids, connecting those – by the relation- graphic source), type localities, the nature of the ship of the uterus and the pericard as well as the extant type material and current systematic pla- inital stages of a radula organisation – to the mol- cement and taxonomic status of the taxa. Sec- luscs, among which the are particularly clo- ond, we aim to place these taxa within the fra- sely related to them through the preservation of the mework of the current systematic knowledge on lateral cords [of the nervous system]”; [translated the phylogeny of aplacophoran molluscs by re- from the German original – M.G.]. In his phyloge- viewing the development of this knowledge, in- netic tree, printed as text-figure in Thiele (1902c: cluding an outline of the results of recent cladistic 438) and following contemporay perception of analyses. This annotated catalogue of aplacophor- phylogenetics, he depicted this hypothesis by plac- an types continues earlier compilations of mollus- ing Solenogastres among the Vermes, as a late off- can material in the ZMB (see bibliography of R. shoot from a branch eventually leading to the Kilias (1929–1999) in Glaubrecht 2001). It adds Mollusca. to those catalogues of other less species-rich However, Thiele’s reservations as to the place- groups, such as Polyplacophora (Kilias 1995a), ment of Solenogastres within molluscs go back Scaphopoda (Kilias 1995b), but also Cephalopo- to his earliest accounts published in 1891 and da (Glaubrecht & Salcedo-Vargas 2000; Ko¨ hler & 1892, when he suggested that “Wurmmollusken” Glaubrecht 2004), whereas the extremely rare (i.e. worm-shaped molluscs) are the Amphineura Monoplacophora are not represented in the of H. von Ihering with the two constituent orders ZMB. Solenogastres or Aplacophora and the Polypla- cophora (Thiele 1892). He viewed Solenogastres as more primitive and the amphineurans in an intermediate and transitional position – as so- Brief review of the history of classification called “bergangsformen” (Thiele 1892) – be- in aplacophoran molluscs tween annelids and molluscs (see also Thiele 1891: 521). Later, Thiele (1895: 867) concluded Seit ich angefangen, mich mit der Anatomie from his ongoing dissections of aplacophorans von Mollusken zu bescha¨ftigen, ist es mein Be- that “Solenogastres are no molluscs because they streben gewesen, u¨ ber die Verwandtschaftsbezie- lack the typical molluscan characters” [translated hungen des ganzen Stammes wie einzelner from the German original]. In addition, Thiele Gruppen ins Klare zu kommen. (1895: 869) suggested to transfer Solenogastres Johannes Thiele, 1894: 222 to the “worms” (i.e. Annelida), thus, also break- Representatives of the two major aplacophoran ing up the Amphineura. molluscs were first published from Scandinavian He continued to stress this view for the rest of waters in 1845 and 1875, respectively, and subse- his life, as is evident, for example, from his ac- quently recognized increasingly often in dredged counts on Solenogastres in Thiele (1925: 13), material from oceanographic expeditions. For the where he perceived them – according to then first time, Heinrich Simroth (1893a) not only dis- prevailing thinking – as “phylogenetische Aus- cussed the then available morphological data on gangsform”, i.e. not only as hypothetical but still aplacophorans but added tentatively a first “Ver- extant evolutionary ancestor to the Mollusca (his such eines Systems” in listing the known genera “Elternform”, see Thiele 1891: 480), with close according to their geographical occurrence, i.e. affinity now to turbellarian worms. Thiele’s posi- as nordic and mediterranean forms. This not tion is also evident from the introductory state- being a systematization in the strict sense, it was ment in his last contribution on Solenogastres an early approach more to compile rather than (Thiele 1933: 144). He was followed in his view, to classify the described taxa; for his extended for example, by Nierstrasz (1909–1910) and systematization see Simroth (1893b: 226–233). Odhner (1921) but not Heath (1911), while e.g. Thiele (1902c) is often cited with the conclusion Hans Hoffmann (1929) expressed an opposed from his first account on the systematic placement opinion in his critical evaluation of the “Amphi- of the Solenogastres in connection with the phylo- neura” discussion; for an historical review see geny of the Mollusca that aplacophoran species e.g. Hoffmann (1929, 1937) and in particular Sal- are not molluscs (e.g. Scheltema 1996: 57). Indeed, vini-Plawen (1967, 1972).

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It was only consequent from this, albeit erro- groups but also discussed at length phylogenetic neous, hypothesis that Thiele (1929) did not in- pathways and higher classification based on com- clude aplacophorans in his Handbuch der Sys- parative investigations. Others, such as in Ger- tematischen Weichtierkunde, which started many Hans Hoffmann (1929, 1951) stated not to accordingly with the Loricata Schumacher, 1817 separate aplacophorans from the Mollusca and (¼ Polyplacophora Gray, 1821) interpreted by proposed to view the Solenogastres as most “pri- him as most primitive and true representatives of mitive” group within this phylum. It was then the phylum Mollusca. In the second volume of Boettger (1956) and later in particular Salvini- his Handbuch, Thiele (1935: 1074) for the last Plawen (1967, and subsequent publications) who time outlined the phylogeny of the Mollusca, took up, after almost half a century of neglect- stating that the Solenogastres cannot be viewed ance of this important molluscan group, again as derived from shelled animals due to, for ex- the controversies as to the monophyly and phy- ample, their peculiar hautmuskelschlauch logeny of “Aplacophora”. In the absence of cla- (p. 1073) and because of other characteristic ana- distic analyses, most authors long followed the tomical features, thus repeating that they are to general assignment of all “worm shaped” mol- be perceived as “Vermes” and excluded from luscs as two clades Solenogastres or Neomenio- the Mollusca (p. 1074). morpha (or Ventroplicida Boettger, 1956 which Two conclusions can be drawn from this. First, is a later naming) and Caudofoveata or Chaeto- evidendently, Thiele and his contemporaries dermomorpha within a monohylum Aplaco- struggled with quite similar problems of molluscan phora. In contrast to these classifications of classification as we still do over a century of zool- “Aplacophora” or “Aculifera”, Salvini-Plawen ogical systematics later, viz. monophyly vs. para- (e.g. 1972, 1985, 1991) has, early and repeatedly, phyly of the aplacophorans, their relationship to argued that aplacophorans are paraphyletic or Polyplacophora within the Amphineura and/or diphyletic, respectively (see below). Aculifera concept and the ancestry of the Mollus- Furthermore, despite revision (Salvini-Plawen ca within the metazoans, in particular their rela- 1978) the systematic relationships remained tionship with Annelida. And second, Thiele’s er- partly unresolved within Solenogastres, and the roneous assumption of Solenogastres being status of some families was poorly understood, “Vermes” resulted in the deposition of aplaco- as was recently shown, for example within the phoran material of the ZMB, and hence the late Cavibelonia by Salvini-Plawen (2004). Earlier re-discovery of it, in the “Vermes” department suggestions to separate three families most likely that initiating this annotated type catalogue. did not reflect natural phylogenetic lineages (Sal- The first half century of research on aplaco- vini-Plawen 1967). According to the systematiza- phoran classification was dominated by the prac- tion suggested by Salvini-Plawen (1978; see also tice to consider all “worm-like” molluscs as Sole- phylogram in Salvini-Plawen & Steiner 1996: 31), nogastres, thus including also those taxa later to four orders are to be distinguished within the So- be distinguished as Caudofoveata; see e.g. treat- lenogastres, viz. Pholidoskepia, Neomeniamor- ments by Simroth (1893b), Thiele (e.g. 1902c, pha (both as Aplotegmentaria) and Sterrofustia 1913c), Nierstrasz (1908) and Heath (1911). plus Cavibelonia (these two as Pachytegmenta- Thiele (1913c) distinguished four taxa among this ria). Thus, the Cavibelonia, in which 11 genus- paraphyletic group, the Chaetodermatidae, Neo- group or families are now distinguished, replaced meniidae, Proneomeniidae, and Lepidomeniidae. the Proneomeniidae s.l. and most of Neomenii- Later, Thiele (1932) suggested to separate five dae s.l. in former systems. For discussion and re- families within the aplacophorans (all considered ferences see Salvini-Plawen (1978, 2003, 2004). by him to represent Solenogastres), viz. Neome- This systematization is followed herein; see sec- diidae, Proneomeniidae, Gymnomeniidae, Lepi- tion C for a classification of the ZMB type mate- domediidae and Crystallophrissontidae. The lat- rial. ter, however, represents Chaetodermomorpha according to current knowledge which were sub- sequently separated by Boettger (1956); see e.g. Phylogenetic analyses Fischer-Piette & Franc (1960) and Salvini-Plawen (1967). For several decades now two hypotheses have Following Thiele’s era, Sigurd Hoffman (1949) been discussed as to the classification reflecting not only contributed essentially new information phylogenetic relationships of the two aplacopho- on the then widely neglected aplacophoran ran taxa, Solenogastres (or neomenioids) and

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Caudofoveata (or chaetoderms). of unresolved whether the lack of ctenidia is a ple- the aplacophorans was proposed by Salvini-Pla- sio- or apomorphic state in Solenogastres. In wen (1967, 1972) and is supported by evidence contrast, radula features (e.g. monoserial type presented by Salvini-Plawen & Steiner (1996) with median bars or two teeth with symphysis) and Haszprunar (2000). It was suggested to rank suggest the Solenogastres to be more conserva- Caudovofeata and Solenogastres as separate tive, as is suggested in cladistic analyses. How- classes, and that Caudofoveata should be distin- ever, this does not necessary imply that Soleno- guished from the remaining molluscan classes gastres actually represent the first (extant) off- (those named Adenopoda). shoot within the Mollusca. Alternatively, Caudo- In contrast, Scheltema (1978, 1993, 1996) and foveata could have just evolved more dramati- others continue – irrespective of her own sepa- cally, in concert with the development of many rate treatment of both distinct groups (e.g. in derived features. As an additional aspect, this Scheltema et al. 1994) – to view both taxa as most recent cladistic analysis suggests that Cavi- subclasses within a single monophyletic class belonia are, contrary to earlier assumptions, Aplacophora that together with the Polyplaco- monophyletic. phora form a clade Aculifera. For example, Different molecular methods have not suc- Scheltema et al. (1994: 13) considered them to ceeded in yielding reliable results beyond sup- be “specialized molluscs with a derived worm porting that aplacophoran groups are diphyletic shape accompanied by reduction or loss of the and are thus not closer related. For example, foot”, while other features “reflect a rather primi- analyses in Solenogastres suffer mainly from the tive molluscan state”. To account for these fact that food material was sequenced instead of “otherwise apparently contradictory states”, Schel- the aplacophorans itself (e.g. , see in tema (1996: 57) suggested progenesis and rapid Salvini-Plawen 2003a; Okusu & Giribet 2003; evolution. However, it should be noted here that Polychaeta for Helicoradomenia in Passamaneck this combination of derived and plesiomorphic et al. 2004; pers. comm. Dr. Hermann Dreyer, features in the grundmuster is a common biologi- Univ. Wien). Another problem is that much of cal phenomenon long known and discussed as the archived material in major museum collec- mosaic evolution without necessarily involving tions has generally been fixed in formaldehyd, progenetic or paedomorphic processes. which is not considered a suitable medium for Recent cladistic analyses by both Salvini-Pla- subsequent molecular work. wen & Steiner (1996) and Haszprunar (2000) found the Solenogastres in the most basal posi- tion suggesting it to represent the earliest mol- On the phylogeny of aplacophoran taxa luscan offshoot. At the same time the mono- phyly of all remaining molluscan classes is Although the origins and interrelationships of suggested, named Hepagastralia by Haszprunar the seven to eight clades or classes of Mollusca (2000: 125) and that way reflecting the distinct are discussed for over a century now, no end of and complex subdivision of the midgut as syna- this debate is visible; for reviews and references pomorphy. Thus, computation of all available see e.g. contributions in Taylor (1996) and also morphological data indicate that both “Aplaco- Haszprunar (2000). While the other clades are phora” and “Aculifera” should not be considered broadly agreed on to represent monophyla, most monophyletic clades, but that – following the recent disagreement centers around the systema- Solenogastres as first off-shoot – the Caudofo- tics and position of aplacophoran, polyplacopho- veata are the sister group to the remaining mol- ran and scaphopod molluscs. As reviewed above, luscs (i.e. Testaria). cladistic analyses finding aplacophorans to form Based on analyses by Hoffman (1949) and a basic, paraphyletic assemblage, also support summarised in Salvini-Plawen (2003a) it is ar- earlier suggestions to consider many conditions gued that the mantle cavity in Neomeniomorpha in their organization as plesiomorphic for mol- and Chaetodermomorpha reveals an indepen- luscs rather than derived through paedomorpho- dent evolutionary transformation of both taxa, sis. In this context Haszprunar (2000: 127) and and that the midgut sac in Caudofoveata is not Salvini-Plawen (2003a: 93) pointed out that many homologous to the so-called digestive gland in shared aplacophoran features can now be consid- Testaria (i.e. Placophora þ ) which, ered as characters typical for the so-called and thus, cannot be interpreted as synapomorphy long-thought-of “hypothetical ancestral molluscs” (see e.g. Haszprunar 2000). The question remains (HAM).

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However, many issues in molluscan phyloge- feran assemblage. In ignorance of this it may be- netics still remain unresolved. Extremely difficult come understandable that these latter authors, in is the interpretation of the scarce ontogenetic context of their new ontogenetic data, again data, not only in molluscs in general but in the stress their view that aplacophorans are “derived aplacophorans in particular. While for Caudofo- molluscs related to chitons and are perhaps pro- veata only the lecithotrophic larvae of two spe- genetic” (Scheltema & Ivanov 2002: 7). cies are known (Salvini-Plawen 1990, Gustafson In summary, recent phylogenetic analyses sug- in Nielsen 1995), development in Solenogastres gest that aplacophorans are unlikely to represent is either lecithotrophic or direct. The ciliated, a monophyletic clade. Instead, among the basal free swimming larva in some Solenogastres re- molluscan groups, Solenogastres may be viewed sembles superficially the so-called “trochophora” as the most earliest off-shoot while Caudofovea- larva in annelids; for a review of aplacophoran ta may be closer to the remaining Mollusca (yet reproduction and development see e.g. Salvini- not as “Hepagastralia”). However, interpretation Plawen (1985). The developmental stages in a of character polarity as well as systematics re- Soleogastres was described originally as so-called main controversial. “Pruvot larva” by the French malacologist G. Pruvot (1890). He reported on a late larva (Ne- matomenia banyulensis) depicting seven rows of Material and Methods scales, or “plaques” as he called it. Although sur- rounded by ambiguities and (even after a cen- Some aplacophoran type material and additional non-type material (see section B below) was found by one of us tury) still in need of further substantiation, this (L.M.), more or less accidentially, during our on-going eva- description of iterated spicules arrangement has luation (since 1997) of all molluscan type material, as it was ever since been interpreted as being homologous long misplaced among opistobranch material in cabinets of the Malacological Department in the ZMB. That way, aplaco- with a shell, thus fueling the discussion on phorans were at least in part hidden to R. Kilias who started the systematic placement of Solenogastres in re- to publish, among other major gastropod groups (see biblio- lation to polyplacophorans and other Mollusca graphy in Glaubrecht 2001), also on some minor (i.e. less speciose taxa) such as e.g. Polyplacophora (Kilias 1995a) and (e.g. Salvini-Plawen 1972, 1985; Salvini-Plawen & Scaphopoda (Kilias 1995b). Steiner 1996). As we can reconstruct from some notes accompanying this Recently, Scheltema & Ivanov (2002) reported aplacophoran material, it was long housed, for the reason of Thiele’s explicit perception of Solenogastres not being Mol- another neomenioid postlarva exhibiting six iter- lusca (see review above), in the “Vermes” department of the ated, transverse groups of spicules separated by ZMB. Upon investigation we luckily found some other parts seven regions devoid of spicules which they com- of the aplacophoran type material there which are now re- turned and housed together with Thiele’s original serial sec- pared to the shell fields in developing polyplaco- tions in the Malacological Collection as listed in the compila- phorans as well as the arrangement on tion below. the Cambrian fossils and Halkieria.The Following sorting and inventarization of this re-discovered aplacophoran material done essentially by L.M., a list of all authors suggested, also refering to recent insight constituent lots was prepared. A first draft of this catalogue from developmental biology on the function of was sent to LvS-P in Dezember 2002, who agreed to check regulatory genes, that this iteration (or serial re- and comment on the taxonomic status of these taxa which was crucial for our endeavour. The final version of the manu- petition) in morphogenesis of ectodermal skele- script was written by M.G. in August 2004, with the inclusion tol structures in these taxa is a result of pro- of the introduction and the historical parts, a review of apla- cesses already present in early pre-Cambrian cophoran classification and its role in molluscan phylogeny. This version was finally again cross-checked in October 2004 bilaterial animals and, thus, would indicate evo- by LvS-P. lutionary relationships among Neomeniomorpha, Based on all combined information available to us then, Polyplacophora and early Paleozoic fossils. They we found additional type material of Thiele’s aplacophorans, viz. of Neomania grandis, Rhopalomenia eisigi and Amphime- suggested that these rows of transverse spicules nia neapolitana, all described by Thiele (1894), but missing that appear briefly in the early development of from the ZMB collection, due to research facilitated by deci- some Solenogastres are expressions of genes that sive notes of LvS-P made 1977 in the Zoological Museum Amsterdam. It has now, after a long post-WW II odyssee, may be present generally in aplacophorans. returned to the ZMB; for more details see below under the However, the authors failed to comment on the species. fact that, when assuming these features do actu- ally represent an ancient developmental process also in early molluscs, then these ectodermal Remarks on the origin of the Berlin type material iterations – albeit homologues – should be viewed as plesiomorphic. They would then not The type material that comprises complete ani- qualify for supporting an aplacophoran or aculi- mals, histological serial sections as well as tissues

# 2005 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim 152 Glaubrecht, M. et al., Aplacophoran Mollusca in the Natural History Museum Berlin and animal parts embedded in paraffin blocks The German Deep-Sea Expedition (Deutsche has diverse origins. For example, due to his early Tiefsee-Expedition), on board of the research investigations of aplacophorans Thiele was pro- steamer “Valdivia”, was on route in the Atlantic vided, starting in the time prior to his curatorial and Southern Indic Ocean from July 31, 1898 to position at the ZMB in 1905, with material by May 1, 1899, as the first major German deep-sea various zoologists, as he reported in case of three research activity subsidized by the German Em- newly named species of Solenogastres from Nea- pire in an nationalistic attempt to keep up with pel (see Thiele 1894: 222–223), or in case of the successful deep-sea expeditions in particular Anamenia amboinensis. Later, Thiele was also a by the British, as e.g. the “Challenger” expedi- contributor of scientific descriptions based on tion 1872–1876. Lead by the biologist Carl Chun material from several of the German imperial (1852–1914) from the University of Leipzig, the expeditions around the turn of the century. In DTE was most innovative in then modern echo particular, he studied the aplacophorans from sounding and sampling techniques and achieved two major oceanographic expeditions, viz. the important results and collections in nearly all zo- German Deep-Sea Expedition on board of the ological phyla. The material on which specialists steamer “Valdivia”, 1898–1899 (with n ¼ 3 new from all over Europe worked for the following species), and in particular the First German decades – among them scientists at the ZMB South Polar Expedition, 1901–1903, on board such as Johannes Thiele (recognized, among the sailing vessel “Gauss” (with 15 new species many other taxa, n ¼ 3 new solenogastres spe- described). Thus, these German expeditions were cies) – forms today an essential part of the Ber- essential in providing new taxa from various re- lin museum’s type collection. Only one example gions of the world and taxonomic groups. How- of the rich material provided by the “Valdivia” ever, they are largely unknown today for various expedition is reflected in the type reasons and, although being of immense histori- collection housed in the ZMB which renders it cal importance, only few historians of science second only to the respective collection in The have dealt with them yet. Here, only brief men- Natural History Museum in London (formely tion will be made of those expeditions in context British Museum of Natural History); see Glau- crucial for collecting Thiele’s aplacophoran mate- brecht & Salcedo-Vargas (2000) with additions in rial, with some hints at the most important avail- Ko¨ hler & Glaubrecht (2004). A most readable – able accounts. and then very successful – travel narrative was The German “Gazelle” Expedition between given by Charl Chun (1900); the zoological re- 1874 and 1876 on board of the navy corvette sults were published in 24 volumes, edited until S.H.S “Gazelle”, lead by Georg Emil Gustav his death by Chun (1902–1914) himself. On the Freiherr von Schleinitz (1834–1910), was on occassion of the 100th anniversary of the “Valdi- route around the world with a focus on research via” expedition, short historical reviews were in the deep-sea of the Atlantic and Indic Ocean, published recently e.g. by Kabisch (1998), Cole- nearly at the same time as the British “Challen- man (1999), and more detailed by Landsberg ger” expedition. One of the expeditions’ aims (2000). Unfortunately, however, no comprehen- was to study the Venus orbit on the near-Ant- sive treatment of this most important expedition arctic Kerguela Island. As naturalist the Swiss and evaluation of its achievements is available zoologist Theophil Studer (1845–1922) took part yet. in the “Gazelle” expedition, who was instrumen- The First German South Polar Expedition tal in collecting and later describing most of its (Deutsche Su¨ dpolar-Expedition) was on route natural objects. An original expedition report from November 11, 1901 until November 25, can be found in Schleinitz & Rottok (1889); see 1903, on board the sailing vessel “Gauss”. Lead also brief accounts in Studer (1882) and Buch- by the young German polar scientist and profes- wald (1999). Only one aplacophoran molluscs, sor of geophysics Erich von Drygalski (1865– Epimenia australis (Thiele 1897) from off the 1949), it was part of the “heroic” period of polar NW coast of Australia, was described and depos- exploration and, as one of the first truely scienti- ited in the ZMB, so we refrain here from a fic endeavours, scientifically highly successful. more detailed review of this endeavour. Never- Carried out in the framework of an international theless, the “Gazelle” marks the start of German programm of geophysical observations, the expe- maritime explorations later continued by two dition was nevertheless rated a failure in Ger- other major expeditions of zoological impor- many due to comtemporary political hopes for tance. imperialistic expansion, resulting in a lack of re-

# 2005 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim Mitt. Mus. Nat.kd. Berl., Zool. Reihe 81 (2005) 2 / http://museum-zool.wiley-vch.de 153 cognition of its merits for long; for a brief histor- “Schwarzen Berg”or“Gaussberg” volcano, as it ical account along these lines see Lu¨ decke was later entered into the maps. This only area (2001) and Lu¨ decke et al. (2001). At the turn of free of ice found by the expedition had an eleva- the century most parts of the South Polar region tion of 366 m above the sea. After breaking free were completely unknown and following the from the ice in early 1903 the expedition failed “Valdivia’s” successful exploration of southern to find other access further south to the Antarc- regions south of the Kerguela Islands at 90E, it tic and returned in June 1903 to South Africa was the clear aim and explicit task of the and home via the Atlantic Ocean. Its scientific “Gauss” expedition to scientifically explore the collections were studied and analysed during the South Polar area, in contrast to the geopolitical subsequent three decades, the results being pub- endeavour of the parallel and competing British lished by Drygalski as editor in a total of 20 vol- expedition lead by Robert Falcon Scott (1868– umes and 2 atlantes (Drygalski 1905–1931); see 1912) on the “Discovery” who was successful in also Drygalski (1904) for an account of his two- reaching south at 82170S, while Drygalski only years Antarctic expedition. (Note that it took 85 made it to about 66S. years to translate into English this semi-popular On board of the “Gauss” was, as the only biol- narrative of the First German South Polar Expe- ogist officially employed, the geograph and zool- dition which partly contributed to its neglectance ogist Ernst Vanho¨ ffen (1858–1918), who had in the annales of history of science). also been a member of the former German Another addition to the knowledge of the in- Deep-Sea Expedition on the “Valdivia”. Edu- vertebrate fauna of polar regions came from the cated at the University Ko¨ nigsberg, Eastern German Ro¨ mer & Schaudinn Expedition to Arc- Prussia, under the influence of Richard Hertwig tic waters around Spitsbergen Island in 1898 on and Carl Chun, Vanho¨ ffen was lecturer, assistant board the steamer “Helgoland”; for a travel nar- and honorary professor at the University Kiel rative see Ro¨ mer & Schaudinn (1900). Basically from 1890 to 1906, when he became curator for a privat enterprise it, nevertheless, gained many Crustacea, Myriapoda and Coelenterata at the zoological objects of interest. Among them, Museum of Natural History in Berlin. He is to- Thiele (1900, 1913b; see also 1932) described day mostly remembered for his studies on cni- two new aplacophoran species. darian medusae and as member of several ex- All this material finally amounted to a total of ploring expeditions, most prominently the n ¼ 24 new aplacophoran mollusc taxa newly de- German South Polar Expedition; for a brief bio- scribed by Thiele (including here also his “kergu- graphy of Vanho¨ ffen including complete biblio- lensis”, albeit re-described only much later by graphy of his zoological writings see Lohmann Salvini-Plawen; see under the species); all are (1918). It was Vanho¨ ffen who collected most zo- now extant in the Malacozoological Collection in ological objects, among them also the many apla- the ZMB. cophorans, as is evident from the original labels and documents. Most material was brought back in particular from the so-called “Winterlager”, Designation of type material when the “Gauss” became beset by pack-ice at 6620S and 89380E in the Indian sector of the Specimens of Thiele’s aplacophoran material Antarctic, 85 km off the Antarctic shelf, and in a were stored in a sea-water/ethanol solution region later to be called “Kaiser Wilhelm II. (“Seewasseralkohol” e.g. in Thiele 1894: 222), of Land”. It is this the type location were the ma- which Thiele generally made histological serial jority of the ZMB material, i.e. a total of n ¼ 15 sections himself, using a double staining agens new solenogastres species were found (see de- (Boraxkarmin and Methylenblau); this is noted tails in Section A), rendering this expedition as “Thiele fec.” on his original labels and below most successful in bringing back Antarctic taxa in Section A; an example of Thiele’s sections is of this interesting molluscan group. The DSE illustrated in Fig. 1. spent the Antarctic winter in 1902 on the sea ice In some taxa several specimens are registered while doing research for 50 weeks. Vanho¨ ffen including intact animals as well as histological se- during this time conducted numerous nettings rial sections mounted on microscopic double- and dredges in various depths down to the bot- slides. Since the majority of Solenogastres taxa tom in –385 m. During one of the seven “in- are well described by their anatomical characters land” explorations by dog-sledges Drygalski’s ex- only, we here follow the preferred procedure of pedition discovered and explored the so-called treating as holotypes and lectotypes, respectively,

# 2005 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim 154 Glaubrecht, M. et al., Aplacophoran Mollusca in the Natural History Museum Berlin those parts of the material (the serial sections), antarctica (Thiele, 1913a) – Dorymenia on which the original description was based, with [Proneomenia] most other undissected material being para(lec- to)types. This turned out to be essentially crucial Proneomenia antarctica Thiele, 1913a: 57, pl. IV, fig. 11; in case of Dorymenia/Proneomenia antarctica pl. VII, figs 10–13; pl. VIII, figs 1–3, 23, 31. Thiele and Labidoherpia/Pruvotina spinosa Ty p e l o c a l i t y : “Antarktis: Winterstation am Thiele (for details see under the species). Lecto- Gaussberg”; Antarctic: winter station at Mount type designation are here done in accordance Gauss at 6620S, 89380E; DSE, Twist 385 m, July with the purpose of Article 74.7.3. of the current 31, 1902; leg. Vanho¨ ffen. issue of the ICZN in order to ensure the name’s Ty p e m a t e r i a l : ZMB Moll. 105.415a, part of holotype, se- proper and consistent application. rial sections on 11 slides (all were originally marked as “types”; n ¼ 6 are labelled a–f, anterior part; plus n ¼ 5 la- belled a–e, posterior part, with b–d additionally marked as Museum codes: “Q 1925 g–i”); USNM United States National Museum of Natural History, ZMB Moll. 105.415b, 4 vials with one (1. vial) containing Smithsonian Insitution, Washington, DC, USA; presumably the remaining, non-sectioned part of the holo- type. ZMA Zoological Museum Amsterdam, The Netherlands; ZMB Museum of Natural History (Museum fu¨ r Natur- 1. vial: 06. XII. 1902, Twist 385 m, part of specimen, dark col- kunde), Humboldt University, Berlin, Germany (for- oured (holotype?); merly Zoological Museum Berlin); 2. vial: 12. X. 1902, Twist 385 m, part of specimen, light col- ZMC Zoological Museum Copenhagen, Denmark. oured (could also be D. hoffmani); 3. vial: 31. XII. 1902, paratype, n ¼ 1 specimen; Expedition abbreviations: 4. vial: 12. X. 1602, n ¼ 1 specimen in ethanol, Thiele fec. ZMB Moll. 105.415c, 4 Paraffin blocks; two blocks each DSE Deutsche Su¨ dpolar-Expedition, 1901–1903; i.e. marked “grosse glatte Promeomenia antarct. 06. XII. 1902 German South Polar Expedition; Vanho¨ ffen” and two blocks marked “Promeomenia? antarcti- DTE Deutsche Tiefsee-Expedition, 1898–1899; i.e. Ger- ca 10. XII. 1902”; paratypes? (but could also be D. hoffma- man Deep Sea Expedition; ni). “Gazelle” Expedition with the vessel “Gazelle”, 1874–1876. C o m m e n t s : Re-examination and determi- nation as Dorymenia antarctica (Thiele, 1913a) by Salvini-Plawen (1978), who was then pro- Section A – Alphabetic list of Solenogastres type vided, unfortunately, with only part of the holo- material in the Malacozoological Collection of type serial sections by R. Kilias (n ¼ 3 slides, the Berlin Museum of Natural History with label “Q1925 g–i), thus his statement “von der Original-Schnittserie liegen drei Objekttra¨- No type material of the second major aplaco- ger vor (Hinterende part.)” (Salvini-Plawen phoran group, the Caudofoveata, is held in the 1978: 247). We verify herewith that the holo- ZMB (but see remark under Chaetoderma pro- type is completely extant, mentioned by Thiele ductum). However, some non-type material of originally as “untersuchtes Tier” in his publica- Chaetodermomorpha is present which is listed tion, with serial sections of the animal’s anterior together with non-type material of the Soleno- and posterior part. Re-examination by LvS-P of gastres in section B. the postradular sections reveals that Thiele’s de- scription is brief but correct. Note that some of the above specimens might actually represent D. hoffmani. Solenogastres amboinensis (Thiele, 1902a) – Anamenia [Proneomenia] arctica Thiele, 1913b –

Proneomenia amboinensis Thiele, 1902a: 733. Nematomenia arctica Thiele, 1913: 161, pl. 1, 2. Ty p e l o c a l i t y : “Amboina”; Indonesia: Am- Ty p e l o c a l i t y : “Spitzbergen; Genauer Fundort boina; leg. Richard Semon. nicht bekannt”; Spitsbergen. Ro¨ mer & Schaudinn Ty p e m a t e r i a l : ZMB Moll. 105.408a, part of holotype, se- Expedition 1898; exact locality unknown. rial sections on 11 slides (labelled a–i “vorn”, i.e. anterior Ty p e m a t e r i a l : ZMB Moll. 105.380a, part of holotype, se- part of animal, and a–e “hinten”, i.e. posterior part); ZMB rial sections on 2 slides (labelled “a vorn” and “b hinten” Moll. 105.408b, part of holotype, in ethanol. and “ad 5075”); ZMB Moll. 105.380b, 1 vial, part of holotype) in ethanol; C o m m e n t s : Type species of Anamenia Nier- ZMB Moll. 105.380c, 1 Paraffin block. (marked with as- strasz, 1908. terics for “type”).

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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-sec- tions 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) austra- lis (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 Ro¨ mer & taxa to be described within the family Epimenii- Schaudinn Expedition see “General remarks” in dae (see Scheltema 2001: 10). Revision and syno- the introduction. nymy in Salvini-Plawen (1997a).

australis (Thiele, 1897) – Epimenia austrina Thiele, 1913a – Phyllomenia [Proneomenia] Phyllomenia austrina Thiele, 1913a: 45, pl. IV, fig. 6; pl. V, Proneomenia australis Thiele, 1897: 398. figs 10–14; pl. VIII, figs 6–9. Ty p e l o c a l i t y : “NW Australien”; NW Austra- Ty p e l o c a l i t y : “Antarktis: Winterstation am lia; 60 fathom (i.e. 108 m isobath); leg. May 7, Gaussberg”; Antarctic: winter station at Mount 1875; “Gazelle” Expedition. Gauss at 6620S, 89380E; DSE, 380 m, January Ty p e m a t e r i a l : ZMB Moll. 105.407a, parts of holotype, 30, 1903; leg. Vanho¨ ffen. serial sections on 48 slides, Thiele fec. (labelled 1–36, ante- Ty p e m a t e r i a l : ZMB Moll. 105.392a, holotype, serial sec- rior part, “Q 3070a–LL; and 1–12 posterior part, “Q tions on 3 slides; Thiele fec. (labelled “a–b vorn”, “ad Q 3070 mm–xx); 1921a–b”; and “a hinten”, “ad Q 1921c”); ZMB Moll. 105.407b, 2 parts of holotype. ZMB Moll. 105.392b, 1 vial, 2 parts of animal and part of C o m m e n t s : Transferred by Nierstrasz (1908) dermis (“30. 01. 1903 WST 380 m, Phyllomenia n. gen., Cuti- cula”); as type species, by monotypy, to his new genus ZMB Moll. 105.392c, 2 Paraffin blocks (marked “Phyllo- Epimenia, which is characterized by unusually menia” and on back of label “Solenogaster, 2 nov. gen., large (up to 300 mm) and bright coloured species 30. 01. 1903”). that occur on the continental shelf and are found C o m m e n t s : Type-species, by monotypy, of the even close to shore at diving depths. Six named new genus Phyllomenia Thiele, 1913a. Re-de- congeneric species are known today, with more scription in Salvini-Plawen (1978: 84–89).

# 2005 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim 156 Glaubrecht, M. et al., Aplacophoran Mollusca in the Natural History Museum Berlin carinata Thiele, 1913a – Sandalomenia Ty p e l o c a l i t y : “Neapel”; Mediterranean Sea: Gulf of Naples. Sandalomenia carinata Thiele, 1913a: 44, pl. IV, figs 5, 25; Ty p e m a t e r i a l : ZMB Moll. 105.403a, holotype; serial sec- pl. V, figs 6–9. tions on 9 slides (labelled “CCCX a–d vorn” and “CCCX a– Ty p e l o c a l i t y : “Antarktis: Winterstation am e hinten”); Gaussberg”; Antarctic: winter station at Mount ZMB Moll. 105.403b, part of holotype. Gauss at 6620S, 89380E; DSE, 385 m, Novem- C o m m e n t s : A synonym of Rhophalomenia ber 24, 1902; leg. Vanho¨ ffen. aglaopheniae (Kowalevsky & Marion, 1887), as Ty p e m a t e r i a l : ZMB Moll. 105.385, holotype, serial sec- was already assumed by Thiele (1894: 269) and tions on 6 slides; Thiele fec. (labelled “a–d vorn”, confirmed by Nierstrasz & Stork (1940: 74) and “24. XI. 1902a, Q 1929 a–d”; and “e” þ “a hinten”, Salvini-Plawen (1967). The serial sections of the “24. XI. 1902a, Q 1929 e þ f”). holotype were given for comparative study to C o m m e n t s : Juvenile animal; re-examination Nierstrasz, later transferred to and re-discovered by Salvini-Plawen (1978: 50–51). in the Zoological Museum in Amsterdam. The material was returned in November 2004 to the ZMB; for more details see below under Amphi- cataphracta (Thiele, 1913a) – Pholidoherpia menia grandis. [Lepidomenia]

Lepidomenia cataphracta Thiele, 1913a: 38, pl. IV, figs 1, 15–18. gauszi Salvini-Plawen, 1978 – Pruvotina Ty p e l o c a l i t y : “Antarktis: Winterstation am Gaussberg”; Antarctic: winter station at Mount Pruvotina gauszi Salvini-Plawen, 1978: 144–145. Gauss at 6620S, 89380E; DSE, 385 m, December Ty p e l o c a l i t y : “Antarktis, Winterstation am 31, 1902; leg. Vanho¨ ffen. Gaussberg”; Antarctic: winter station at Mount 0 0 Ty p e m a t e r i a l : ZMB Moll. 105.386a, syntype, serial sec- Gauss at 66 2 S, 89 38 E; DSE; leg. Vanho¨ ffen. tion on 1 slide; fec. L. v. Salvini-Plawen, 19.02.1975; Ty p e m a t e r i a l : ZMB Moll. 105.394, holotype, serial sec- ZMB Moll. 105.386b, syntype; 2 vials (with 2 specimens, tion on 1 slide Thiele fec. (labelled “ad Q 1923d, Pruvotina Thiele fec., and n ¼ 15 specimens). spinosa juv. Thiele”). C o m m e n t s : Thiele (1913a) mentioned the C o m m e n t s : This material was originally de- study of 20 specimens, here treated as syntypes. scribed by Thiele (1913a) as juvenile of Pruvoti- Systematic revision in Salvini-Plawen (1978: 51), na spinosa Thiele, 1913a and determined as the with transfer as type species in new genus Pholi- above taxon by Salvini-Plawen (1978: 144–145). doherpia. Note that the serial sections (ZMB 105.394), ori- ginally labelled “d” only represent the posterior clavigera Thiele, 1897 – Notomenia part of the animal.

Notomenia clavigera Thiele, 1897: 398. Ty p e l o c a l i t y : “Torres-Straße”; Torres Strait, gaussiana Thiele, 1913a – Acanthomenia between Australia and New Guinea; 20 fathoms (36m), dredged. Acanthomenia gaussiana Thiele, 1913a: 62, pl. IV, fig. 7a; textfig. 2. Ty p e m a t e r i a l : ZMB Moll. 105.391, holotype, serial sec- tions on 6slides; Thiele fec. (labelled “CCCXLIII a–f”, with Ty p e l o c a l i t y : “Antarktis”; Antarctic: DSE, no. of “Vermes” Department “5495 a–f”). at 65270S, 80330E, in depth of –3398 m, March C o m m e n t s : Type species, by original designa- 30, 1903; leg. Vanho¨ ffen. tion, of Thiele’s new genus Notomenia. Re-exam- Ty p e m a t e r i a l : ZMB Moll. 105.404, holotype, serial sec- ination and re-description in Salvini-Plawen tions on 2 slides; Thiele fec. (labelled “Q 1928a–b, 3398 m”). (2004) leads to the classification within the new C o m m e n t s : Type-species by monotypy of family Notomeniidae due to its anatomical orga- Thiele’s (1913a) new genus Acanthomenia. Revi- nization unrelated to any other genus within the sion in Salvini-Plawen (1978: 157–158). framework of the hitherto established families.

glacialis Thiele, 1913a – Nematomenia eisigi Thiele, 1894 – Rhopalomenia Nematomenia glacialis Thiele, 1913a: 40, pl. IV, figs 3, 21–22; Rhopalomenia eisigi Thiele, 1894: 269. pl. V, fig. 1.

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Ty p e l o c a l i t y : “Antarktis, Winterstation am Gauss at 6620S, 89380E; DSE, 380 m, Jun 14, Gaussberg”; Antarctic: winter station at Mount 1902; Twist 385 m, November 09, 1902; Decem- Gauss at 6620S, 89380E; DSE, Twist 385 m, Jun ber 17, 1902; leg. Vanho¨ ffen. 16, 1902; 380–385 m, from July 31 to December Ty p e m a t e r i a l : ZMB Moll. 105.399a, part of lectotype 19, 1902; Twist 380 m, from July 01, 1902 to Jan- (by present designation), serial sections on 3 slides. Thiele uary 12, 1903; leg. Vanho¨ ffen. fec. (labelled “a–b vorn”, “1924a–b, 09. XI. 1902”; and “a hinten”, “1924c, 09. XI. 1902”); Ty p e m a t e r i a l : ZMB Moll. 105.381a, lectotype (by pre- ZMB Moll. 105.399b, part of lectotype (by present desig- sent designation), with serial section on 5 slides, Thiele fec. nation); labelled “Th., 09. XI. 1902, Twist 385 m”); (labelled “Q 1919a–c”, and “Q 1919d–e”); ZMB Moll. 105.399c, paralectotypes in 3 vials, in alcohol ZMB Moll. 105.381b, paralectotypes in 14 vials (1. vial: (1. vial: “? M. intermedia Th., Gauss-St[ation]”, one speci- 31. VII. 1902, WSt 380 m, one part of specimen; 2. vial: men; 2. vial: “? M. intermedia, 14. VI. 1902”, one specimen; 16. VI. 1902, Twist 385 m, one specimen; 3. vial: 19. XII. 3. vial: “Proneomenia intermedia, 17. XII. 1902”, one speci- 1902, 385 m, one specimen; 4. vial: 07. I. 1903, Twist 380 m; men). vial 5 –14, with a total of n ¼ 9 specimens, all without labels or date); C o m m e n t s : Type-species, by monotypy, of ZMB Moll. 105.381c, paralectotypes, 2 Paraffin block (la- Thiele’s new genus Metamenia. Re-description in belled “glacialis 31. 07. 1902”); Salvini-Plawen (1978: 149–150). 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) kerguelensis Salvini-Plawen, 2005 – clearly indicated one fully adult animal, here giv- Ocheyoherpia en as lectotype. Ocheyoherpia kerguelensis Salvini-Plawen, 2005: 100, figs 1–4. Ty p e l o c a l i t y : “Kerguelen”; South of the ant- grandis Thiele, 1894 – Neomenia arctic circle; France, Antarctic-Kerguelen Islands, DTE, station 160. Unfortunately, in the 1899 Neomenia grandis Thiele, 1894: 223. compilation of station numbers for the DTE ex- Ty p e l o c a l i t y : “Neapel”; Mediterranean Sea: pedition on board “Valdivia” no precise coordi- Gulf of Naples. nates were given, but only a note that says “Ga- Ty p e m a t e r i a l : ZMB Moll. 105.387a, part of holotype, se- zelle Basin” and “3.5–4.0 C surface tem- rial sections on 52 slides (labelled “CCCI a–z2 vorn”, Thiele Dresden 1893; “CCCII a–b”; “CCCIII a–r Hinterende”; perature”. “CCCVI a–f”); Ty p e m a t e r i a l : ZMB Moll. 105.395a, holotype, serial sec- ZMB Moll. 105.387b, part of holotype (ca. 4 cm) in etha- tions on 3 slides and one spicule slide; Thiele fec. (labelled nol. originally by Thiele as “Q 3030a–c”, plus spicule slide “Q 3030 d”); C o m m e n t s : Re-examined by H. A. Stork (in ZMB Moll. 105.395b, paratypes, Thiele fec.; 3 specimens Nierstrasz & Stork 1940: 43) who already sug- (two of them, with 0.6 and 0.5 mm juveniles, the other juve- gested not to distinguish N. grandis Thiele, 1894 nile with 1.15 mm, decalcified without calcerous bodies/spi- cules). The latter specimen is illustrated in Fig. 2, prior to from N. carinata Tullberg, 1875. They placed histological sections (on n ¼ 10 slides) done by LvS-P in Oc- grandis in synonymy of N. carinata Tullberg, as tober 2004. did Salvini-Plawen (1986, 1997b). The missing se- C o m m e n t s : This taxon was only provisionally rial sections of the holotype (and other material named kerguelensis by Johannes Thiele, as it is of Thiele) were given for comparative study to evident from the labels accompagning the ZMB Nierstrasz and later transferred together with material, but was never described in publication Amphimenia neapolitana and Rhopalomenia eisi- by this author, thus providing only a manuscript gi to the collection of the Zoological Museum in name without nomenclaturial bearings. There- Amsterdam. It was sent back to the ZMB thank- fore, the taxon was recently described as a spe- fully by Robert Moolenbeek in November 2004 cies of the genus Ocheyoherpia, now formally after uncovering the post-WW II odyssee of this named by Salvini-Plawen (2005) utilizing the material (see more under A. neapolitana). name “kerguelensis” as originally suggested by Thiele. The taxon is tentatively transferred to ordo Sterrofustia, family Phyllomeniidae. intermedia Thiele, 1913a – Metamenia

Metamenia intermedia Thiele, 1913a: 52, pl. IV, fig. 9; pl. VII, figs 1–2; pl. VIII, figs 16–21. neapolitana Thiele, 1889 – Amphimenia

Ty p e l o c a l i t y : “Antarktis, Winterstation am Proneomenia neapolitana Thiele, 1889: 429 (footnote). Gaussberg”; Antarctic: winter station at Mount Proneomenia (Amphimenia) neapolitana Thiele, 1894: 244.

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to the ZMB in November 2004 by Robert Moo- lenbeek.

papilligera Thiele, 1913a – Sandalomenia

Sandalomenia papilligera Thiele, 1913a: 41, pl. IV, figs 4, 24; pl. V, figs 2–5. Ty p e l o c a l i t y : “Antarktis, Winterstation am Gaussberg”; Antarctic: winter station at Mount Gauss at 6620S, 89380E; DSE, 385 m, from No- vember 22 to 24, 1902; Twist 380 m, January 28, 1903; leg. Vanho¨ ffen. Fig. 2. Habitus of one juvenile of Ocheyoherpia kerguelensis Ty p e m a t e r i a l : ZMB Moll. 105.384a, syntype, serial sec- Salvini-Plawen, 2005 (ZMB 105.395b; paratype); the com- tions on 7 slides; Thiele fec. (labelled “Q 1920 a–d vorn” plete material of this new taxon, including three juvenile ani- and “Q 1920 e–g hinten”); mals, was originally labelled “Pruvotina kerguelensis“by ZMB Moll. 105.384b, part of syntype; Thiele, but only recently validly named and described by Sal- ZMB Moll. 105.384c, syntype, 1 specimen in alcohol (la- vini-Plawen (2005); see text for more details. Scale ¼ 0.5 mm. belled “28. I. 1903, Twist 380 m); ZMB Moll. 105.384d, syntypes, 2 Paraffin blocks. Ty p e l o c a l i t y : “Neapel”; Mediterranean Sea: C o m m e n t s : Type-species, by subsequent des- Gulf of Naples. ignation (Salvini-Plawen 1978: 48), of the new Ty p e m a t e r i a l : ZMB Moll. 105.429, holotype, serial sec- genus Sandalomenia Thiele, 1913a. Re-descrip- tions on 14 slides, from animal (c. 3 cm in length and 1.5 mm tion in Salvini-Plawen (1978: 48–50). thick according to Thiele (1894: 244)); Thiele fec. (labelled “CCII a–o”). C o m m e n t s : Type-species, by monotypy, of Amphimenia Thiele, 1894; as new subgenus of prisca Thiele, 1906 – Archaeomenia Proneomenia Hubrecht, 1880. Currently trans- ferred to the Amphimeniidae. This taxon, de- Archaeomenia prisca Thiele, 1906: 315, pl. XXVIII. scribed together with Neomenia grandis and Ty p e l o c a l i t y : “Agulhas Bank, am su¨ dlichen Rhopalomenia eisigi by Thiele (1894), was long Teil”; southern part of Agulhas bank, off South missing from the ZMB collection as was also Africa; DTE, station 110 (3590S, 1832.80E; trawl Proneomenia vagans. Uncovering the odyssee of 564 m, October 4, 1898. all this material from the available evidence, we Ty p e m a t e r i a l : ZMB Moll. 59.910a, syntype, serial sec- reconstruct that it was given by Thiele in the late tions on 8 slides (labelled “a–c vorn”, “Q 5078/1–3, Valdi- via” and “a–e hinten”, “Q 5078/4–8, Valdivia”); 1920th/early 1930th on loan for the preparation ZMB Moll. 59.910b, syntype, serial sections on 5 slides (la- of the Naples monograph to H. F. Nierstrasz at belled “a–e Sagitalschnitte”, “Q 5078/9–13); the University Museum in Utrecht (Nierstrasz in ZMB Moll. 59.910c, syntypes, 2 specimens in veil. Nierstrasz & Stork 1940: 1). After Nierstrasz’ C o m m e n t s : Type-species, by monotypy, of death (Sept. 1937) the material remained in Thiele’s new genus Archaeomenia. Re-examined Utrecht until the university stopped its systema- in Salvini-Plawen & Paar-Gausch (2004). tic research. Thiele’s material was sent in 1956 to the then curator at the Zoological Museum in Amsterdam, T. Van Bentem-Jutting (while other protecta Nematomenia material from Utrecht was transferred at the Thiele, 1913a – same time to the Natural History Museum in Nematomenia protecta Thiele, 1913a: 39, pl. IV, figs 2, 19–20. Leiden). In search of the type-material, it was traced out in May 1977 by LvS-P together with Ty p e l o c a l i t y : “Antarktis, Winterstation am S. Van der Spoel, the then curator at the ZMA, Gaussberg”; Antarctic: winter station at Mount 0 0 on the occassion of a visit to the collection of Gauss at 66 2 S, 89 38 E; DSE, 385 m, July 9, the ZMA. LvS-P made exact notes about the 1902; 385 m, December 2–4, 1902; leg. Vanho¨f- boxes in which Thiele’s original material in the fen. ZMA were deposited. Triggered by this for- 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 warded information, the material was finally re- “9. VII. 1902, 385 [m]” and one slide labelled “2.– discovered in the ZMA and thankfully sent back 4. XII. 1902, 385 m”).

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C o m m e n t s : Note that only the organisation of belled “d” and “e–g”, respectively, were as- the scales of the mantle is known (Thiele 1913a: signed to Pruvotina gauszi Salvini-Plawen, 1978 39–40; Salvini-Plawen 1978: 45, fig. 28). and Gephyroherpia antarctica Salvini-Plawen, 1978 (for details see there). providens Thiele, 1913a – Pruvotina squamosa Thiele, 1913a – Nematomenia Pruvotina providens Thiele, 1913a: 48, pl. IV, fig. 7; pl. V, figs 15–17; pl. VI, figs 1–3; pl. VIII, figs 10–11. Nematomenia squamosa Thiele, 1913a: 40, pl. IV, fig. 23. Ty p e l o c a l i t y : “Antarktis, Winterstation am Ty p e l o c a l i t y : “Antarktis, Winterstation am Gaussberg”; Antarctic: winter station at Mount Gaussberg”; Antarctic: winter station at Mount 0 0 Gauss at 66 2 S, 89 38 E; DSE 385 m Breitnetz Gauss at 6620S, 89380E; DSE, 385 m, December and Twist, December 17, 1902; 385 m Twist, Jan- 17, 1902; leg. Vanho¨ ffen. uary 28, 1903; 350 m Twist, February 8, 1903; leg. Ty p e m a t e r i a l : ZMB Moll. 105.383, holotype, whole Vanho¨ ffen. mount on slide; Thiele fec. (“labelled 17. XII. 1902; 385 m”; Ty p e m a t e r i a l : ZMB Moll. 105.397a, part of holotype, se- one of three animals on the microscopic slide is circled with rial sections on 3 slides, Thiele fec. (labelled “a–b Vorder- black marking, indicated as “type”; the other two are given ende”, “ad Q 1922 a–b”; and “a Hinterende”, “ad Q 1922 c”); as “N. glacialis Thiele, Antarktis, Gauss Winterstation, D. ZMB Moll. 105.397b, part of holotype (labelled Su¨ dp. Exp., Vanho¨ ffen S, Q1931”). “17. XII. 1902, 385 m”); C o m m e n t s : Only the mantle scales are known ZMB Moll. 105.397c, paratypes in 6 vials (1. vial: 08. II. 1903, Twist 350 m, 1 specimen; 2. vial: without label, 1 (Thiele 1913a: 39–40; Salvini-Plawen 1978: 45). 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 speci- mens); thulensis (Thiele, 1900) – Thieleherpia ZMB Moll. 105.397d, paratype, 1 Paraffin block. [Proneomenia] C o m m e n t s : Re-description by Salvini-Plawen (1978: 119–120). Proneomenia thulensis Thiele, 1900: 111, pl. V. Ty p e l o c a l i t y : “Spitzbergen, Station 18 (8080N, 16550E), 480 m”; Spitsbergen, Hinlopen spinosa (Thiele, 1913a) – Labidoherpia Strait, at northern entrance; leg. Ro¨ mer & [Pruvotina] Schaudinn Expedition, June 1, 1898; on fine, blue mud with only few small stones in front of a gla- Pruvotina spinosa Thiele, 1913a: 50, pl. IV, fig. 8; pl. VI, cier. Locality data here added according to en- figs 4–7; pl. VIII, fig. 12–15. tries given in Ro¨ mer & Schaudinn (1900: 40) in Ty p e l o c a l i t y : “Antarktis, Winterstation am a compilation of dredge stations. Gaussberg”; Antarctic: winter station at Mount Ty p e m a t e r i a l : ZMB Moll. 105.405a, part of holotype, se- Gauss at 6620S, 89380E; DSE; 385 m; leg. Van- rial section on 14 slides (labelled “a–f” for anterior part, and ho¨ ffen. “a–f” for posterior part); ZMB Moll. 105.405b, part of holotype, in ethanol. Ty p e m a t e r i a l : ZMB Moll. 105.398a, lectotype (by pre- sent designation), serial sections on 3 slides; Thiele fec. (la- C o m m e n t s : Re-description and classification belled “a – vorn”, “ad Q 1923 a”; and “a–b”, “ad Q 1923 as Thieleherpia thulensis (Thiele, 1900) in Salvini- b–c; b ¼ 28. I. 1903”); Plawen (2004: 86–88), with the additional exam- 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, ination of another specimen from off northeast- 1 specimen; 3. vial: 12. I. 1903, 380 m, 6 specimens; 4. vial: 8 ern Iceland (ZMUC). Due to features of the specimens; 5. vial: 14 specimens; 6. vial: 1 specimen; 7. vial: ventral foregut glandular organs the species was 46 specimens; 8. vial: 12. X. 1902, 385 m, 1 specimen; 9. vial: 22. I. 1903, 2 specimens, one with long spiculae; 10. vial: stated by Salvini-Plawen (1978: 231) not to be- 09. VII. 1902, 385 m, 2 specimens); long to Proneomenia and finally placed together ZMB Moll. 105.398c, paralectotypes, 2 Paraffin blocks (la- with Rhipidoherpia within the Rhipidoherpiidae belled “10. 01. 1903 juv.” and “28. 01. 1903”). (see Salvini-Plawen 2004: 88). Comments: Pruvotina spinosa Thiele, 1913a has been assigned by Salvini-Plawen (1978) as type species of the new genus Labidoherpia. tricarinata (Thiele, 1913a) – Dorymenia Note that of the originally seven slides with seri- [Proneomenia] al sections assigned to this species by Thiele (1913a), only three (ZMB Moll. 105.398a) are Proneomenia tricarinata Thiele, 1913a: 54, pl. IV, fig. 10; here kept with spinosa, while the other slides, la- pl. VII, figs 3–9; pl. VIII, figs 22, 24–30.

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Ty p e l o c a l i t y : “Antarktis, Winterstation am C o m m e n t s : Type species of the genus. Type Gaussberg”; Antarctic: winter station at Mount locality: Banyuls-sur-mer, France, Mediterranean Gauss at 6620S, 89380E; DSE, (17. 07. 1902– coast. Known distribution: European shelf from 12. 10. 1902 Twist 385 m, 09. 11. 1902–22. und Peleponnes (Greece) to Scottland; in 50–137 m. 24. 11. 1902 Twist 385 m, 17. 12. 1902 Breitnetz The material now housed again in the ZMB has 385 m – 23. 12. 1902 Twist 385 m, 12. 01. 1903 been resent from the ZMA together with other Twist 385 m – 24. 01. 1903 Twist 380 m, parts of Thiele’s material from Neaples such as, 31. 01. 1903 Twist 380 m – 08. 02. 1903 385 m, for example, Amphimenia neapolitana (see under 15. 02. 1903 Twist 400 m), leg. Vanho¨ ffen. this species) and Rhopalomenia eisigi (Thiele, Ty p e m a t e r i a l : ZMB Moll. 105.413a, part of holotype, se- 1894) which is synonymized with Rh. agalophe- rial sections on 11 slides, Thiele fec. (n ¼ 5 slides labelled niae (see under the former species). “a–e vorn”, “Q 1925 k–o”; n ¼ 6 slides labelled “a–f hin- ten”, “Q 1925 a–f”); ZMB Moll. 105.413b, part of holotype (labelled “08. II. 1903, 385 m”); antarctica Salvini-Plawen, 1978 – 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 Gephyroherpia 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: Gephyroherpia antarctica Salvini-Plawen, 1978: 115. 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, L o c a l i t y : “Antarktis: Winterstation am Gauss- 1 specimen; 8. vial: 17. XII. 1902, 1 specimen; 9. vial: berg”; Antarctic: winter station at Mount Gauss 12. X. 1902, Twist 385 m, 1 specimen; 10. vial: 09. XI. 1902, 0 0 Twist 385 m, 1 specimen); at 66 2 S, 89 38 E; DSE; leg. Vanho¨ ffen. ZMB Moll. 105.413d, paratypes, 2 Paraffin blocks. M a t e r i a l : ZMB Moll. 105.393, serial sections on 3 slides, C o m m e n t s : Re-description and classification Thiele fec. (labellel “e–g”). as Dorymenia tricarinata (Thiele, 1913a) in Salvi- C o m m e n t s : The type material for this species ni-Plawen (1978: 257–260). is deposited in the USNM (Smithsonian Institu- tion) in Washington, DC (USA). Type locality: Ross Sea. The material in ZMB has been de- scribed as “Pruvotina spinosa juv.” by Thiele valdiviae Thiele, 1902b – Proneomenia (1913a: 52); see Salvini-Plawen (1978: 118).

Proneomenia valdiviae Thiele, 1902b: 167, pl. XXIII. Type locality: “Ku¨ ste von Ostafrika, N von banyulensis Pruvot, 1890 – Nematomenia Sansibar; am Netz ha¨ngend, Sublimat”; DTE, station 249 (370S, 4045.80E); offshore E Africa, Dondersia banyulensis Pruvot, 1890: 699–810. Myzomenia banyulensis (Pruvot, 1890) – Simroth 1893a. north of Sansibar; trawl 748 m, March 23, 1899. L o c a l i t y : Mediterranean Sea: Gulf of Neaples; Ty p e m a t e r i a l : ZMB Moll. 105.406a, part of holotype, se- rial sections on 15 slides (labelled “a–i vorn”; and “a–f hin- Italia. ten”); M a t e r i a l : ZMB Moll. 105.426, serial sections on 6 slides, ZMB Moll. 105.406b, part of holotype, in ethanol (labelled Thiele fec. (labellel “CLXXIV a–c”, anterior part of animal; “5049”, referring to a number in the “Vermes” Collection). and “CLXXII”, posterior part of animal); ZMB Moll. 105.379, specimen in ethanol; Thiele fec. (“Ply- mouth, 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-sur- Section B – The non-type material in the Mala- mer, France, Mediterranean coast. The actually cozoological Collection of the ZMB known distribution ranges from Dalmatia (Adria- tic Sea) to the Trondheimsfjord (Norway) at 31– Solenogastres 300 m depth. The material now housed again in the ZMB (105.426) has been resent from the aglaopheniae (Kowalevsky & Marion, 1887) – ZMA together with other parts of Thiele’s materi- Rhopalomenia al from Neaples such as, for example, Amphime- nia neapolitana (see under this species). Proneomenia aglaopheniae Kowalevsky & Marion, 1887: 1–76. L o c a l i t y : Mediterranean Sea: Gulf of Neaples; Italia. hoffmani Salvini-Plawen, 1978 – Dorymenia 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”). Dorymenia hoffmani Salvini-Plawen, 1978: 247.

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L o c a l i t y : “Antarktis: Winterstation am Gauss- productum Wire´n, 1892 – Chaetoderma berg”; Antarctic: winter station at Mount Gauss at 6620S, 89380E; DSE, leg. Vanho¨ ffen. Chaetoderma productum, Wire´n 1892: 86. Crystallophrisson productum (Wire´n) – Thiele (1932). M a t e r i a l : ZMB Moll. 105.420, serial section on 9 slides 0 0 (labelled by Thiele as “Q 1926 a–k”); see Fig. 1. Ty p e l o c a l i t y : Kara Sea (71 20 N, 59 58 E); C o m m e n t s : This material has been assigned 62 fathoms (¼ –114 to 115 m depth), leg. Sept. originally to Proneomenia antarctica by Thiele 9, 1883; “Dijmphna”-Expedition (Stat. 188). (1913a), but was separated from this species by Salvini-Plawen (1978: 247). The type material of Ty p e m a t e r i a l : Lectotype and paralectotype in ZMC, vide Salvini-Plawen (1975). the latter is deposited in the Zoological Museum of the University Uppsala (Schweden), as serial section on slides. 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. Wire´n’s description of vagans Kowalevsky & Marion, 1887 – a new arctic species (see Comments below). Dorymenia C o m m e n t s : There are three locations known Proneomenia vagans Kowalevsky & Marion 1887: 1–76. from the Kara Sea where C. productum was L o c a l i t y : Mediterranean Sea: Gulf of Neaples; caught with several specimens each during the Italia. Danish-International “Dijmphna” Expedition M a t e r i a l : ZMB Moll. 105.427, serial sections on 3 slides, from August 1882 to August 1883 (see Paulsen Thiele fec. (labellel “CLXVII a–c”). 1884). The three specimens in the ZMB are ap- C o m m e n t s : The species had been redescribed parently part of the material originally collected by Thiele (1894). It is known from the western at the locus typicus in the Kara Sea by this expe- Mediterranean, between Marseille and Neaples, dition, albeit they are not of the syntype series from depth down to 20–30 m. The serial sections that Wire´n based his description on. While we of the ZMB were given on loan for comparative were unable to reconstruct how the ZMB got study to Nierstrasz, later transferred to the Zool- part of the material from the Dijmphna” Expedi- ogical Museum in Amsterdam and returned in tion in the first place, Wire´n (1892: 8) explicitly November 2004 to the ZMB; for more details stated in his introduction that in summer 1891 he see below under Amphimenia neapolitana. had a chance to see this material in the collec- tions of both the Copenhagen Museum and the Berlin Museum. However, he continued that for his description in 1892 of a new species from the Caudofoveata Kara Sea, distinct from C. nitidulum, he received only the material from the Copenhagen Museum This group (also named Chaetodermomorpha), through Dr. G. M. R. Levinsen. is traditionally subdivided into three families, Li- Thiele (1902c: 273) mentioned dissections mifossoridae (with Metachaetoderma Thiele, done on a specimen from the Go¨ tting Zoological 1913), Prochaetodermatidae (with Prochaetoder- collection, ded. Ehlers, and the publication date ma Thiele, 1902) and Chaetodermatidae; see e.g. is before Thiele starting to work as curator in Salvini-Plawen (1971, 1975, 1985) and Scheltema the ZMB. However, this Go¨ tting specimens re- (1981). presents 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 col- Chaetodermatidae Ihering, 1876 lection from the “Vermes” department in De- zember 1930. The family represents a cosmopolitan taxon com- This species was originally considered by Odh- prising three genera with a total of 80 species de- ner (1921) as subspecies of Chaetoderma nitidu- scribed, mostly from the shelf regions and the lum (Love´n, 1845), which is, however, not known continental slopes. It includes with Chaetoderma from a holotype or syntype specimen (see Salvi- productum being up to 14 cm in length the great- ni-Plawen 1984; Ivanov & Scheltema 2000). For est Caudofoveate. a new characterisation see Salvini-Plawen (1975).

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Prochaetodermatidae Salvini-Plawen, 1968 A – Solenogastres

The family represents a cosmopolitan taxon Aplotegmentaria based upon the single genus Prochaetoderma, Thiele, including today six (poorly defined) gen- Ordo Pholidoskepia era or subgenera with 25 named species. They are of small size, less than 5 mm, and mostly be- Nematomenia arctica Thiele, 1913b long to the deep-sea fauna (Salvini-Plawen 1992; Nematomenia banyulensis (Pruvot, 1890) Scheltema & Ivanov 2000, 2004; Ivanov & Schel- [Dondersia] tema 2002). Thiele (1902c: 275) introduced Pro- Nematomenia glacialis Thiele, 1913 chaetoderma for Chaetoderma radulifera Kowa- Nematomenia (?) protecta Thiele, 1913 levsky (1901), according to the description by Nematomenia (?) squamosa Thiele, 1913a this author, from the Marmara Sea. Today no Sandalomeniidae original material is retained. Sandalomenia papilligera Thiele, 1913 Sandalomenia carinata Thiele, 1913a Incertae sedis Section C – List of nominal taxa housed in the Pholidoherpia cataphracta (Thiele, 1913a) Malacozoological Collection of the Museum of [Lepidomenia] Natural History Berlin (ZMB), arranged accord- Ordo Neomeniamorpha ing to current systematic affiliation Hemimeniidae Archaeomenia prisca Thiele, 1906 The present catalogue comprises a total of 31 Neomeniidae aplacophoran taxa, including six non-type taxa. Neomenia grandis Thiele, 1894 ¼ N. carinata Of these taxa n ¼ 23 were named and described Tullberg, 1875 originally by Johannes Thiele, with an additional four by Salvini-Plawen (1978, 2005), which are Pachytegmentaria partly based on material first described by Thiele. With the exception of the caudofoveate Ordo Sterrofustia Chaetoderma productum Wire´n, 1892 all aplaco- Phyllomeniidae phorans in the ZMB are Solenogastres. They are Phyllomenia austrina Thiele, 1913a assigned to 20 genera, following the classification Ocheyoherpia kerguelensis Salvini-Plawen, 2005 suggested by Salvini-Plawen (1978). For a char- Ordo Cavibelonia1 acterization of the genera see e.g. Salvini-Plawen Pararrhopaliidae2 (1967). Gephyroherpia antarctica Salvini-Plawen, 1978 Currently, 29 out of the 31 aplacohoran taxa [Pruvotina spinosa Thiele, partim] are considered valid species, two are considered Pruvotina gauszi Salvini-Plawen, 1978 [Pruvotina synonyms (i.e. Neomenia grandis of N. carinata, spinosa Thiele, partim] and Rhopalomenia eisigi of R. aglaopheniae). A Pruvotina providens Thiele, 1913 third, Ocheyoherpia kerguelensis Salvini-Plawen, Labidoherpia spinosa (Thiele, 1913a) 2005 was newly described, based on material [Pruvotina] provisionally named but not described and pub- Metamenia intermedia Thiele, 1913a lished as valid name by J. Thiele. Acanthomeniidae The following list is considered the most re- Acanthomenia gaussiana Thiele, 1913a cent classification on aplacophoran molluscs, and Notomeniidae essentially follows that suggested by Salvini-Pla- Notomenia clavigera Thiele, 1897 wen (1978) with additions in Salvini-Plawen Rhopalomeniidae (2004). Type species of genera are given in bold, Rhopalomenia aglaopheniae (Kowalevsky & synonyms are given as “¼”; original generic as- Marion, 1887) [Proneomenia] sigments are in square parentheses. Numbers re- Rhopalomenia eisigi Thiele, 1894 ¼ R. aglaophe- fer to footnotes below. niae (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)

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Strophomeniidae Caudofoveata Anamenia amboinensis (Thiele, 1902a) [Proneomenia] Limifossoridae Epimeniidae Metachaetoderma Thiele, 1913a; as new genus. Epimenia australis (Thiele, 1897) [Proneomenia] Type-species, by monotypy: Chaetoderma chal- Proneomeniidae lengeri Nierstrasz, 1903. Proneomenia valdiviae Thiele, 1902b Prochaetodermatidae Dorymenia antarctica (Thiele, 1913a) Prochaetoderma Thiele, 1902; as new genus. [Proneomenia] Type-species, by monotypy: Chaetoderma radu- Dorymenia hoffmani Salvini-Plawen, 1978 liferum Kowalevsky, 1901. [Proneomenia antarctica Thiele, partim] Dorymenia tricarinata (Thiele, 1913a) [Proneomenia] Acknowledgements Dorymenia vagans (Kowalevsky & Marion, 1887) [Proneomenia] We are most grateful to Robert Moolenbeck for altruistically searching for and locating in the ZMA three of Thiele’s apla- Amphimeniidae cophoran types from Neaples that were missing after being Amphimenia neapolitana (Thiele, 1889) sent out on loan to Nierstrasz in Utrecht before WW II. We [Proneomenia] thank Frank Ko¨ hler for his help with inventorizing the apla- cophoran type material of the ZMB and for valuable com- Rhipidoherpiidae ments on the manuscript, as well as Jeroen Goud (Naturalis, Thieleherpia thulensis (Thiele, 1900) Leiden) for additional information. We thank, as always, [Proneomenia] 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 B – Caudofoveata: care; for helping to locate aplacophoran material in the latter department we are thankful to Mrs. K. Ka¨mpf and K. Mesch- ter. We are indepted to Michael Ohl (ZMB) and an anony- Ordo Chaetodermomorpha mous referee for some valuable comments that helped to im- Chaetodermatidae prove the manuscript. Chaetoderma productum Wire´n, 1892

References

Section D – List of nominal taxa named by Belyaev, G. M. 1966. Hadal bottom fauna of the world ocean. Acad. Nauk. USSR, Inst. Okeanology: 1–199. [translated Thiele, but representatives not extant in the Mu- from Russian and published 1972 by the Israel Program seum of Natural History Berlin (ZMB) for Scientific Translations, Jerusalem]. Boettger, C. 1956. Beitra¨ge zur Systematik der Urmollusken (Amphineura). – Zoologischer Anzeiger, Suppl. 19: 223– The following taxa have been named by Thiele, 256. in addition to those listed alphabethically under Bieler, R. & Boss, K. J. 1989. Johannes Thiele and his contri- section A and B (see also systematic section C). butions to zoology. Part 1. Biography and bibliography. – Nemouria 34: 1–30. However, material representing these genus-level Boss, K. J. & Bieler, R. 1991. Johannes Thiele and his contri- taxa is not extant (i.e. not found in the ZMB col- butions to zoology. Part 2. Genus-group names (Mollus- lection). Compilation according to Boss & Bieler ca). – Nemouria 39: 1–76. Brusca, R. C. & Brusca, G. J. 2003. Invertebrates, Second (1991) who listed a total of 11 genus-level names edition. Sunderland: Sinauer Associates, Inc. introduced by Thiele. Buchwald, B. 1999. Die Forschungsreise SMS Gazelle 1874 bis 1876. Hamburg, Berlin: Elbe-Spree-Verlag. Coleman, C. O. 1999. 100 Jahre Deutsche Tiefsee-Forschung. – Naturwissenschaftliche Rundschau 52: 442–445. Solenogastres Chun, C. 1900. Aus den Tiefen des Weltmeeres. Schilderun- gen von der Deutschen Tiefsee-Expedition. Jena: G. Fischer [second revised edition 1903] Dondersiidae –– 1902–1914. Wissenschaftliche Ergebnisse der Deutschen Heathia Thiele, 1913a; as new genus. Type-spe- Tiefsee-Expedition auf dem Dampfer “Valdivia” 1898– cies, by monotypy: Ichthyomenia porosa 1899. 24 vols. Jena: G. Fischer. Drygalski, E. v. 1904. Zum Kontinent des eisigen Su¨ dens. Heath, 1911. Berlin: G. Reimer. [English translation, by M. M. Raraty, Rhopalomeniidae 1989: The Southern Ice-Continent: the German South Po- Pruvotia Thiele, 1894; as new genus. Type-spe- lar Expedition aboard the Gauss 1901–1903. Harleston: Bluntisham Books & Erskine Press. 394 pp.] cies, by monotypy: Proneomenia sopita Pruvot, Drygalski, E. v. (ed.) 1905–1931. Deutsche Su¨ dpolar-Expedi- 1891. tion 1901–1903. 20 vols., 2 atlantes. Berlin: G. Reimer.

# 2005 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim 164 Glaubrecht, M. et al., Aplacophoran Mollusca in the Natural History Museum Berlin

Fischer-Piette, E. & Franc, A. 1960. Classe des Aplacophores. –– 1995b. Scaphopoda-Typen und -Typoide (Mollusca) im In Grasse´, P.-P. (ed.), Traite´ de Zoologie. Anatomie, Sys- Zoologischen Museum in Berlin. – Mitteilungen aus dem te´matique, Biologie. Vol. 5 (2): 1655–1700. Paris: Masson Museum fu¨ r Naturkunde Berlin, Zoologische Reihe 71 et Cie. (1): 171–177. Glaubrecht, M. 2001. In memoriam Rudolf Kilias. – Archiv Ko¨ hler, F. & Glaubrecht, M. 2004. An addendum to the cata- fu¨ r Molluskenkunde 129 (1/2): 1–6. logue of cephalopod types in the Museum fu¨ r Natur- Glaubrecht, M. & Salcedo-Vargas, M. A. 2000. Annotated kunde Berlin, with remarks on Onychoteuthis taxa de- type catalogue of the Cephalopoda (Mollusca) in the Mu- scribed by Hinrich Lichtenstein. – Mitteilungen aus dem seum fu¨ r Naturkunde, Humboldt University of Berlin. – Museum fu¨ r Naturkunde Berlin, Zoologische Reihe 80 Mitteilungen Museum fu¨ r Naturkunde Berlin, Zool. Rei- (2): 275–282. he 76 (2): 269–282. Landsberg, H. 2000. „Die Wissenschaft wird streng und Go¨ tting, K.-J. 1974. Malakozoologie. : G. Fischer. nu¨ chtern richten ...“ (Carl Chun 1900). 100 Jahre –– 1985. Mollusca, Weichtiere. In Siewing, R. (ed.), Lehr- Deutsche Tiefsee-Expedition “Valdivia”. – Verhandlun- buch der Zoologie, Band 2. Systematik [3. Auflage], gen zur Geschichte und Theorie der Biologie 5: 75–90. pp. 653–700. Stuttgart, New York: G. Fischer. Lindberg, D. R. & Ponder, W. F. 1996. An evolutionary tree –– 1996. Mollusca, Weichtiere. In Westheide, W. & Rieger, for the Mollusca: branches or roots? In Taylor, J. (ed.), R. (eds), Spezielle Zoologie, Teil 1. Einzeller und wirbel- Origin and evolutionary radiation of the Mollusca, lose Tiere, pp. 276–330. Stuttgart, Jena: G. Fischer. pp. 66–75. Oxford, New York: Oxford University Press. Handl, C. & Salvini-Plawen, L. v. 2001. New records of Sole- Lohmann, H. 1918. Ernst Vanho¨ ffen. – Mitteilungen aus nogastres-Pholidoskepia (Mollusca) from Norwegian dem Zoologischen Museum in Berlin 9 (1): 71–90. fjords and shelf waters including two new species. – Sar- Lu¨ decke, C. 2001. Das Prinzip der Freiheit. Erich von Dry- sia 86: 367–381. galski leitete die erste deutsche Antarktisexpedition. – –– 2002. New records of Solenogastres-Cavibelonia (Mollus- Naturwissenschaftliche Rundschau 54 (12): 643–648. ca) from Norwegian fjords and shelf waters including Lu¨ decke, C., Brogiato, H. P. & Ho¨ nsch, I. 2001. Universitas three new species. – Sarsia 87: 423–450. Antarctica. 100 Jahre deutsche Su¨ dpolarexpedition 1901– Haszprunar, G. 2000. Is the Aplacophora monophyletic? A 1903 unter der Leitung Erich von Drygalskis. Ausstellungs- cladistic point of view. – American Malacological Bulletin heft Inst. La¨nderkunde, Leipzig (Publik Offset, Leipzig). 15: 115–130. 32 pp. Heath, H. 1911. Reports on the scientific results of the expe- Nierstrasz, H. F. 1908. Mollusca. VII. Solenogastres. – Natl. dition to the tropical Pacific, in charge of Alexander Antarctic Expedition 1901–1904 4 (7): 1–13. Agassiz, by the U.S. Fish Commission steamer “Alba- –– 1909–1910. – Ergebnisse und Fortschritte der Zoologie tross”, from August, 1899, to June, 1900. Vol. 14, The So- 1 þ 2: 239–306; 367–430. lenogastres. – Memoirs of the Museum of Comparative Nierstrasz, H. F. & Stork, H. A. 1940. – Monographie der Zoology, Harvard College 45 (1): 1–179. Solenogastren des Golfes von Neapel. Zoologica (Stutt- Hoffmann, H. 1929. Amphineura und Scaphopoda; Nach- gart) 99: 1–99. tra¨ge. Bronns Klassen und Ordnungen des Tierreichs wis- Odhner, N. 1921. Norwegian Solenogastres. – Bergens Mu- senschaftlich dargestellt in Wort und Bild. Vol. 3, Mollus- seums Aarbok 1918/19: 1–86. ca, I. Abtl. Suppl., pp. 1–368. Leipzig: Akademische Okusu, A. & Giribet, G. 2003. New 18S rRNA sequences Verlagsgesellschaft. from neomenioid aplacophorans and the possible origin –– 1937. ber die Stammesgeschichte der Weichtiere. – Zo- of persistent exogenous contamination. – Journal of Mol- ologischer Anzeiger 10 (Suppl. Verhandlungen der luscan Studies 69: 385–387. Deutschen Zoologischen Gesellschaft): 33–69. Passamaneck, Y. J., Schander, Ch. & Halanych, K. M. 2004. –– 1951. Mollusca. In Bertalanffy, L. v. (ed.), Handbuch der Investigation of molluscan phylogeny using large-subunit Biologie. Vol. 6 (6), pp. 161–208. Potsdam: Akademische and small-subunit nuclear rRNA sequences. – Molecular Verlagsgesellschaft Athenaion. Phylogenetics and Evolution 32: 25–38. Hoffman, S. 1949. Studien u¨ ber das Integument der Soleno- Paulsen, A. F. W. 1884. Re´sume´ des travaux de l’expedition gastren nebst Bemerkungen u¨ ber die Verwandtschaft internationale danoise faits a Godthaab (Groenland Oc- zwischen den Solenogastren und Placophoren. – Zoolo- cidental), 1er aout 1882–31 aout 1883, suivi d’un som- giska Bidrag fran Uppsala 27: 293–427. maire des observations me´te´orologiques faites pendant Hyman, L. 1967. Mollusca I. The Invertebrates 6: 1–792. la de´rive du Dijmphna dans la mer de Kara. Copenha- Ihering, H. v. 1876. Versuch eines natu¨ rlichen Systemes der gen. Mollusken. – Jahrbu¨ cher der Deutschen Malakozoolo- Pruvot, G. 1890. Sur le de´veloppement d’un Sole´nogastre. – gischen Gesellschaft 3: 97–148. Comptes rendus hebdomadairs de Se´ances de l’Acade´mie Ivanov, D. L. 1996. Origin of Aculifera and problems of des Sciences 111: 689–692. monophyly of higher taxa in molluscs. In Taylor, J. (ed.), Ro¨ mer, F. & Schaudinn, F. 1900. Einleitung, Plan des Werkes Origin and evolutionary radiation of the Mollusca, und Reisebericht. In Ro¨ mer, F. & Schaudinn, F. (eds), pp. 59–65. Oxford, New York: Oxford University Press. Fauna Arctica. Eine Zusammenstellung der arktischen Ivanov, D. L. & Scheltema, A. H. 2000. On the nomenclatur- Tierformen, mit besonderer Beru¨ cksichtigung des Spitz- al status of the “type series” of Chaetoderma nitidulum bergen-Gebietes auf Grund der Ergebnisse der Love´n, 1844. – Ruthenica 10:1–4. Deutschen Expedition in das No¨ rdliche Eismeer im Jahre –– 2002. Prochaetodermatidae of the Indian Ocean collected 1898, vol. 1, pp. 1–84. Jena: G. Fischer. during Soviet VITYAZ cruises 1959–1964 (Mollusca: Salvini-Plawen, L. v. 1967. Kritische Bemerkungen zum Sys- Aplacophora). – Molluscan Research 22: 183–202. tem der Solenogastres (Mollusca, Aculifera). – Zeitschrift –– 2004. Dacryomica plana, gen. et sp. nov., a prochaetoder- fu¨ r zoologische Systematik und Evolutionsforschung 5: matid Aplacophora from a Pacific seamount. – The Veli- 398–444. ger 47 (1): 1–5. –– 1971. Schild- und Furchenfu¨ sser (Caudofoveata und Sole- Kabisch, K. 1998. 100 Jahre deutsche Tiefseeforschung. – nogastres), verkannte Weichtiere am Meeresgrund. Die Deutsche Aquarien- und Terrarien-Zeitschrift 51 (10): Neue Brehm-Bu¨ cherei, vol. 441. Wittenberg Lutherstadt: 656–658. A. Ziemsen Verlag. Kilias, R. 1995a. Polyplacophora-Typen und -Typoide (Mol- –– 1972. Zur Morphologie und Phylogenie der Mollusken: lusca) im Zoologischen Museum in Berlin Polyplaco- Die Beziehung der Caudofovata und der Solenogastres phora. – Mitteilungen aus dem Museum fu¨ r Naturkunde als Aculifera, als Mollusca und als Spiralia. – Zeitschrift Berlin, Zoologische Reihe 71 (1): 155–170. fu¨ r wissenschaftliche Zoologie 184: 205–394.

# 2005 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim Mitt. Mus. Nat.kd. Berl., Zool. Reihe 81 (2005) 2 / http://museum-zool.wiley-vch.de 165

– 1975. Mollusca Caudovofeata. – of Scheltema, A. H., Tscherkassky, M. & Kuzirian, A. M. 1994. Scandinavia, 4: 1–35. Aplacophora. In Harrison, F. W. & Kohn, A. J. (eds), Mi- –– 1978. Antartische und subantarktische Solenogastres – croscopic anatomy of Invertebrates. Vol. 5, Mollusca I, Eine Monographie: 1898–1974. – Zoologica (Stuttgart) pp. 13–54. New York: Wiley-Liss. 128: 1–315. Schleinitz, F. v. & Rottok, D. 1889 (eds). Die Forschungs- –– 1984. Comments on Chaetoderma and Crystallophrisson reise S.M.S. “Gazelle” in den Jahren 1874 bis 1876 unter (Mollusca, Caudofoveata). – Zool. Zhurnal 63: 171–175 Kommando des Kapita¨n zur See Freiherr von Schleinitz. [in Russian]. 1. Theil. Der Reisebericht. Berlin: E. S. Mittler und –– 1985. Early evolution and the primitive groups. In True- Sohn. man, E. R. & Clarke, M. R. (eds), The Mollusca. Evolu- Simroth, H. 1893a. Kritische Bemerkungen u¨ ber die Syste- tion. Vol. 10, pp. 59–150. Orlando: Academic Press. matik der Neomeniiden. – Zeitschrift fu¨ r wissenschaf- –– 1990. Origin, phylogeny and classification of the phylum tliche Zoologie 56: 310–327. Mollusca. – Iberus 9: 1–33. –– 1893b. Aplacophora. In: Amphineura und Scaphopoda. –– 1992. On certain Caudofoveata from the Vema-Expedi- Bronns Klassen und Ordnungen des Tierreichs wis- tion. Proceedings of the 9th International Malacolological senschaftlich dargestellt in Wort und Bild. Vol. 3, Mollus- Congress, Edinburgh 1986 (Unitas Malacologica, Leiden): ca, I. Abtl.: 133–233. Leipzig: C. F. Winter’sche Verlags- 317–333. handlung. –– 1997a. Systematic revision of the Epimeniidae (Mollusca: Studer, T. 1882. ber einige wissenschaftliche Ergebnisse der Solenogastres). – Journal of Molluscan Studies 63: 131– Gazelle-Expedition namentlich in zoogeographischer Be- 155. ziehung. In: Verhandlungen des zweiten deutschen Geo- – 1997b. Fragmented knowledge on West-European and graphentages, pp. 1–19: Berlin: Dietrich Reimer. Iberian Caudofoveata and Solenogastres. – Iberus 15: Taylor, J. (ed.) 1996. Origin and evolutionary radiation of the 35–50. Mollusca. Oxford, New York: Oxford University Press. –– 2003a. On the phylogenetic significance of the aplaco- Thiele, J. 1891. Die Stammesverwandtschaft der Mollusken. phoran Mollusca. – Iberus, 21 (1): 67–97. Ein Beitrag zur Phylogenie der Tiere. – Jenaische Zeit- –– 2003b. Contributions to West-Mediterranean Solenogas- schrift fu¨ r Naturwissenschaft 25 (N.F. 18): 480–543. tres (Mollusca) with three new species. – Iberus 21 (2): –– 1892. Wurmmollusken. – Sitzungsberichte der naturwis- 37–60. senschaftlichen Gesellschaft ISIS in Dresden 1892:3–4. –– 2004. Contributions to the morphological diversity and –– 1894. Beitra¨ge zur vergleichenden Anatomie der Amphi- classification of the order Cavibelonia (Mollusca: Soleno- neuren. I. ber einige Neapeler Solenogastres. – Zeit- gastres). – Journal of Molluscan Studies 70: 73–93. schrift fu¨ r wissenschaftliche Zoologie 58 (2): 222–302. –– 2005. Ocheyoherpia kerguelensis spec. nov. (Mollusca: So- –– 1895. ber die Verwandtschaftsbeziehungen der Amphi- lenogastres) von der Deutschen Tiefsee-Expedition „Val- neuren. – Biologisches Centralblatt 15 (24): 859–869. divia“ 1898–1899. – Mitteilungen aus dem Museum fu¨r –– 1897. Zwei australische Solenogastres. – Zoologischer Naturkunde, Zoologische Reihe 81 (1): 99–104. Anzeiger 20: 398–400. Salvini-Plawen, L. v. & Paar-Gausch, I. 2004. Three new spe- –– 1900. Proneomenia thulensis n. sp. In Ro¨ mer, F. & Schau- cies of Neomenia (Mollusca: Solenogastres) from the dinn, F. (eds), Fauna Arctica. Eine Zusammenstellung der southern hemisphere. – New Zealand Journal Marine arktischen Tierformen, mit besonderer Beru¨ cksichtigung Freshwater Research 38: 137–162. des Spitzbergen-Gebietes auf Grund der Ergebnisse der Salvini-Plawen, L. v. & Steiner, G. 1996. Synapomorphies Deutschen Expedition in das No¨ rdliche Eismeer im Jahre and symplesiomorphies in higher classification of Mollus- 1898, vol. 1, pp. 111–116. Jena: G. Fischer. ca. In Taylor, J. (ed.), Origin and evolutionary radiation –– 1902a. Proneomenia amboinensis n. sp. – Denkschriften of the Mollusca, pp. 29–51. Oxford, New York: Oxford der Medizinisch-Naturwissenschaftlichen Gesellschaft zu University Press. Jena 8: 733–738. Scheltema, A. H. 1978. Position of the class Aplacophora in –– 1902b. Proneomenia valdiviae n. sp. Wissenschaftliche Er- the phylum Mollusca. – Malacologia 17: 99–109. gebnisse der Deutschen Tiefsee-Expedition auf dem –– 1981. Comparative morphology of the radulae and ali- Dampfer „Valdivia“, 1889–1899, 3: 167–174. mentary tracts in the Aplacophora. – Malacologia 20: –– 1902c. Die systematische Stellung der Solenogastren und 361–383. die Phylogenie der Mollusken. – Zeitschrift fu¨ r wis- –– 1993. Aplacophora as progenetic aculiferans and the co- senschaftliche Zoologie 72: 249–466. elomate origin of the mollusks as the sister taxon of Si- –– 1906. Archaeomenia prisca n. g., n. sp. Wissenschaftliche puncula. – Biological Bulletin 184: 57–78. Ergebnisse der Deutschen Tiefsee-Expedition auf dem –– 1996. Phylogenetic position of Sipuncula, Mollusca and Dampfer „Valdivia“ 1889–1899, 9: 315–324. the progenetic Aplacophora. In Taylor, J. (ed.), Origin –– 1911. Die Solenogastres der Russischen Polar-Expedition and evolutionary radiation of the Mollusca, pp. 53–58. 1900–1903. – Me´moires de l’Acade´mie inpe´riale des Oxford, New York: Oxford University Press. Sciences St. Pe´tersbourg, se´rie VIII, classe physico-mathe´- –– 1998. Class Aplacophora. In: Beesley, P. L., Ross, G. J. B. matique 29 (5): 1–4. & Wells, A. (eds), Mollusca: The Southern Synthesis, Fau- –– 1913a. Antarktische Solenogastren. Deutsche Su¨ dpolar- na of Australia, vol. 5, part A, pp. 145–159. Melbourne: Expedition 1901–1903. Vol. 14 (Zool., 6) (1): 35–65. Ber- CSIRO Publishing. lin. G. Reimer. –– 2001. Aplacophora. In: Wells, A. & Houston, W. W. K. –– 1913b. Ein neuer Solenogaster von Spitzbergen. – Sit- (eds.), Zoological Catalogue of Australia. Vol. 17.2. Mol- zungsberichte der Gesellschaft Naturforschender Freunde lusca. Aplacophora, Polyplacophora, Scaphopoda, Cepha- Berlin 1913 (2): 160–162. lopoda, pp. 1–18. Melbourne: CSIRO Publishing. –– 1913c. Solenogastres. In Schulze, F. E. (ed.), Das Tier- Scheltema, A. H. & Ivanov, D. L. 2000. Prochaetodermatidae reich. Eine Zusammenstellung und Kennzeichnung der of the eastern Atlantic Ocean and the Mediterranean Sea rezenten Tierformen. Vol. 38, pp. 1–57. Berlin: R. Fried- (Mollusca: Aplacophora). – Journal of Molluscan Studies la¨nder & Sohn. 66: 313–362. –– 1925. Solenogastres. In Ku¨ kenthal, W. & Krumbach, T. –– 2002. An aplacophoran postlarva with iterated dorsal (eds), Handbuch der Zoologie. Eine Naturgeschichte der groups of spicules and skeletal similarities to Paleozoic Sta¨mme des Tierreiches. Vol. 5 (1), pp. 1–14. Berlin, fossils. – Invertebrate Biology 121 (1): 1–10. Leipzig: Walter de Gruyter & Co. –– 2004. Use of birefringence to characterize Aplacophora –– 1929–1935. Handbuch der Systematischen Weichtier- sclerites. – The Veliger 47: 153–156. kunde. Vols. 1 þ 2. Jena: G. Fischer.

# 2005 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim 166 Glaubrecht, M. et al., Aplacophoran Mollusca in the Natural History Museum Berlin

–– 1932. Die Solenogastres des arktischen Gebietes. In Ro¨- Todt, C. & Salvini-Plawen, L. v. 2003. New mer, F. & Schaudinn, F. (eds), Fauna Arctica, Eine Zu- (Mollusca: Solenogastres) from the Mozambique Chan- sammenstellung der arktischen Tierformen, mit beson- nel, Western Indian Ocean. – The Veliger 46 (3): 252– derer Beru¨ cksichtigung des Spitzbergen-Gebietes auf 266. Grund der Ergebnisse der Deutschen Expedition in das Wire´n, A., 1892. Studien u¨ ber Solenogastres II (Chaetoderma No¨ rdliche Eismeer im Jahre 1898, vol. 6 (4), pp. 379–382. productum, Neomenia, Proneomenia acuminata). – Kungl. Jena: Frommann Druck (H.Pohl). Svenska Vetenskapsakademiens Handlingar 25 (6): 1– – 1933. Solenogastres. In Handbuch der Naturwissenschaf- 100. ten. Zweite Auflage [2nd edition]. Vol. 9, pp. 144–146. Jena. G. Fischer.

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