TERMS OF USE This pdf is provided by Magnolia Press for private/research use. Commercial sale or deposition in a public library or website is prohibited. Zootaxa 2392: 1–32 (2010) www.mapress.com / zootaxa/ ISSN 1175-5326 (print edition) Article Copyright © 2010 · Magnolia Press ZOOTAXA ISSN 1175-5334 (online edition) Analytic taxonomy and notes on marine, brackish-water and estuarine Gastrotricha WILLIAM D. HUMMON1 & M. ANTONIO TODARO2,3 1 Department of Biological Sciences, Ohio University, Athens, Ohio 45701 USA. E-mail: hummon@ohio.edu Dipartimento di Biologia Animale, Università di Modena e Reggio Emilia, via Campi 213/d, I-41100 Modena, Italy. E-mail: antonio.todaro@unimore.it 3 Corresponding author. E-mail: antonio.todaro@unimore.it 2 Abstract Marine Gastrotricha, both Macrodasyida and Chaetonotida, are the subject of an analytic review, citing taxonomic status of names, authorships of taxa, and those responsible for changes, in accordance with the International Code of Zoological Nomenclature, 4th ed. (1999). Notes are included with regard to taxonomic usage so as to guide workers in the future. Among the proposed novelties are: within Macrodasyida, to restrict the family Lepidodasyidae Remane, 1927 to the genus Lepidodasys Remane, 1926, and to establish a new family, Cephalodasyidae with Cephalodasys Remane, 1926 as its type-species to house the remaining genera and species that have been contained in the polyphyletic family Lepidodasyidae. Hemidasys agaso Claparède, 1867 is considered extinct, and the new name Tetranchyroderma antenniphorum is proposed for Tetranchyroderma antennatum Luporini, Magagnini & Tongiorgi, 1973; in addition, five species are here considered to be species inquirendae: Dactylopodola weilli d'Hondt, 1965, Paradasys nipponensis Sudzuki, 1976, Macrodasys indicus Kutty & Nair, 1969. Tetranchyroderma forceps d’Hondt & Balsamo, 2009 and Turbanella plana (Giard, 1904b). Among Chaetonotida: the Xenotrichula velox-species group Ruppert, 1979 and the Xenotrichula intermedia-species group Ruppert, 1979 were given each the rank of subgenus. Chaetonotus pleuracanthus Remane, 1926 is rejected as a synonym for Chaetonotus marinus Giard, 1904; Chaetonotus somniculosus Mock, 1979 is transferred to the genus Halichaetonotus, the new name Halichaetonotus euromarinus is proposed for Halichaetonotus spinosus Mock, 1979, and Xenotrichula carolinensis Ruppert, 1979 is re-established. Heteroxenotrichula variocirrata d'Hondt, 1966 is here considered to be species inquirenda. Key words: Gastrotricha, Macrodasyida, Chaetonotida, new family, new subgenus, new species, taxonomy, marine, brackish-water, estuarine Introduction With the field of meiofauna research in a mode of rapid change and an exponential increase of electronically disseminated nominal lists of the world biota, we felt that it was time for an authoritative review of the taxonomic status of marine and brackishwater/estuarine Gastrotricha, along with the author that was responsible for changes, and reasons why changes have been made; our intent is not to provide a complete taxonomic history from the 18th century onward, but to establish correct 21st century taxonomic usage. Each of us has produced lists before (see Todaro 2008, Hummon 2009b), but with greater responsibility on us for our published stand, this list is probably more conservative than either of us would otherwise be, as it has been drafted using the International Code of Zoological Nomenclature (ICZN) recommendations as a guideline. With more than 550 taxa and over 110 papers to consider, several of which belong to the “grey” literature and some of uncertain publication date, absolute perfection would be difficult to reach. However, we strive for at least ninety-five percent accuracy. Where we have given a source or date, we welcome friendly changes and hope to incorporate them as we continue forward; we also welcome those who differ with us on facts or interpretation to correct or update us by use of print medium, which should meet the criteria of scientific literature (i.e., peer reviewed papers). Accepted by W. Sterrer: 11 Jan. 2010; published: 8 Mar. 2010 1 TERMS OF USE This pdf is provided by Magnolia Press for private/research use. Commercial sale or deposition in a public library or website is prohibited. Recently, a work somewhat comparable to ours has been proposed for the freshwater taxa (Balsamo, d’Hondt, Pierboni & Grilli, 2009). We regard this as an important and useful taxonomic tool, though we have reservations on some points. We hold that new hypotheses for the re-systematization of well-established taxa should be based on new, unambiguous evidence (e.g. molecular data, additional morphological data aquired by novel techniques etc.) rather than on the rescoring of well-known traits. Consequently, for purposes of this paper the taxon Hystrichochaetonous will be considered as a valid subgenus within Chaetonotus, according to Schwank (1990) and Kisielewski (1997). Likewise, the recent paper of d’Hondt & Balsamo (2009) has been answered with respect to each of the cases on which well-established systematization has been challenged. Our work will hopefully prove useful to a wide audience, including senior and expecially junior researchers who work on gastrotrichs, but also to marine zoologists and ecologists who find these abundant metazoans in the course of research on interstitial meiobenthos and who want to refer to them correctly. Obviously, the paper may be used as a reference source to be checked against for validation by editors and keepers of electronically disseminated lists devoted to these animals. Systematic account Phylum GASTROTRICHA Metschnikoff, 1865:458 Class There are no classes within the Gastrotricha, as it was itself at one time considered a class within the phylum Aschelminthes! Order Macrodasyida Remane, 1925:125. —Includes 32 genera and 297 species, of which 30 genera and 295 species are marine or brackish [The name as given by Remane had an -oidea ending, that of a superfamily (ICZN 29.2), which was first altered to a preferred ordinal –ida ending by Brunson (1950:328) and later to be formally so designated by Rao, 1970: 109 and Rao & Clausen, 1970:80, and has almost universally been accepted as emended] Family Cephalodasyidae —new family—Includes 6 genera and 31 species [Established in this publication to include Cephalodays, Dolichodasys, Megadasys, Mesodasys and Paradasys, previously affiliated with the family Lepidodasyidae. The rationale for our choice rests on the growing body of evidence that concurs in showing the unnatural grouping of the traditional Lepidodasyidae (e.g. Ruppert 1978; Hochberg & Litvaitis 2000; Guidi et al., 2004; Todaro et al 2006) and on the vast consensus among workers in indicating that the major source of polyphyly in the family lies in the difference between Lepidodasys and the remaining constituents of the former family. The name derives from Cephalodasys, the first genus in the group to be named, while the data for the family diagnosis have been gathered from published work. Note that a split of the former Lepidodasyidae somewhat comparable to ours had been proposed by d’Hondt (1975:657) at the tribal level. However, that division was based only on the presence or absence of scales, which in our judgment was not of sufficient importance to warrant such a high-ranking division, but most of all it did not address the evident polyphyly of the former taxon] Cephalodasyidae diagnosis [In this publication] Elongate Macrodasyida, flattened ventrally and vaulted dorsally in transverse section with small to medium terminal or slightly subterminal mouth opening and little to marked head delimitation. Cuticle naked. Monociliated or multiciliated epidermal cells; ventral cilia in two longitudinal rows, often united fore and aft. Epidermal glands vary in size and number. Anterior adhesive tubes (TbA) in two groups behind the mouth, either inserting directly on the cuticle (Dolichodasys, Megadasys, Mesodasys and Paradasys) or inserting on fleshy hands (Cephalodasys and Pleurodasys); lateral, dorsolateral and ventrolateral adhesive tubes (TbL/TbDL/TbVL) arranged in columns along the body, occasionally absent (Paradasys) or in form 2 · Zootaxa 2392 © 2010 Magnolia Press HUMMON & TODARO TERMS OF USE This pdf is provided by Magnolia Press for private/research use. Commercial sale or deposition in a public library or website is prohibited. of adhesive papillae (Dolichodasys); posterior adhesive tubes (TbP) arranged marginally around the blunt to pointed posterior end. Sphincter muscle developed around mouth opening; well-developed striated radial pharyngeal musculature; pharyngeal pores basal. Circular muscles present in lateral regions of the body. Y-cells absent. Gonads paired; male anterior, female posterior; male gametes mature posterior to anterior, female gametes posterior to anterior (Megadasys, Mesodasys, Paradasys, Pleurodasys) or anterior to posterior (Cephalodasys, Dolichodasys); male pores, when present, single (Cephalodasys, Megadasys, Mesodasys), or paired (Dolichodasys, Pleurodasys); frontal and/or caudal organ usually present. Intertidal or subtidal in distribution; fine to coarse sand. Type-Genus Cephalodasys Remane, 1926b:681 [Includes =Psammodasys d’Hondt, 1974b:675] Syn. Psammodasys d’Hondt, 1974b:675 [Hummon, 2008b:121. d’Hondt and Balsamo (2009:273) reject the synonymy, considering Psammodasys to be a valid genus based on hypothetical differences on the arrangement of the accessory sexual organs; according to them Psammodasys species have bursa copulatrix (=bourse copulatrice) and seminal receptacle (=réceptacle séminal) topographically separated (bursa being much posterior) whereas Cephalodasys species have the seminal receptacle that immediately precedes the bursa. In our opinion these apparent differences are due to the nomenclatural misuse of the gastrotrichs’ accessory sexual organs, which in the past has plagued the descriptions of several taxa, and derived from early misunderstanding about the function of these organs. As later pointed out (e.g. Ruppert 1978, 1991), bursa and seminal receptacle are synonymous and constitute the frontal organ, which is part of the female sexual apparatus, in that it receives and stores allosperm from cross fertilization. Where present, shape and size of the frontal organ vary across the taxonomic spectrum and may be species specific. In some cases, the functional regions of the frontal organ may be clearly discernable morphologically (e.g., in Macrodasys, see Ruppert 1978); in others the frontal organ is visible only in specimens that have copulated (i.e. containing allosperm, e.g. Turbanella). In Gastrotricha, the male counterpart of the frontal organ is the caudal organ. Physiologically, it functions as a penis. However, because of the anatomical discontinuity between testis and the caudal organ, which characterize the phylum, this peculiar penis must first be charged with autosperm before its copulatory function can be performed (e.g. Ruppert 1991). Presence/absence of the caudal organ varies across the taxonomic spectrum, whereas the shape and size may be species specific. In general, where both frontal organ and caudal organ are present, they appear to be topographically separated, or next to each other but always without luminal continuity; a notable exception are, perhaps, the gastrotrichs Thaumastodermatinae (Ruppert 1978). With these premises it appears clear that in Cephalodasys/Psammodasys what is called a seminal receptacle actually is the frontal organ, a structure female in function, whereas the “bourse copulatrice” corresponds to the caudal organ (penis). Although in all the species in which a bursa has been reported it has been considered part of the female sexual apparatus, and in C. caudatus Rao, 1981 the author clearly stated that the species lacks a penis (see Rao, 1981, not 1961, as mistakenly reported in d’Hondt & Balsamo, 2009). That said, it is obvious that the two organs (frontal and caudal organs =seminal receptacle and “bourse copulatrice”), even when they appear to be next to each other, cannot have their lumina in anatomical continuity; consequently, it is functionally irrelevant whether these organs are topographically separated or adjacent to each other. Therefore, in our opinion the establishment of a genus (i.e. Psammodasys) to allocate species with characteristics of Cephalodasys but having caudal and frontal organ topographically separated appears not to be justified. Our position is strengthened by knowing that, within the genus,several species for which only the frontal organ is reported (e.g. C. pacificus and C. palavensis) and species which seem to be parthenogenetic (e.g., C. hadrosomus) provide a complex situation, which is matched within the Macrodasyida by the genus Urodasys] Type-species Cephalodasys maximus Remane, 1926b:681 Cephalodasys cambriensis (Boaden, 1963a):404 TAXONOMY OF MARINE GASTROTRICHA Zootaxa 2392 © 2010 Magnolia Press · 3 TERMS OF USE This pdf is provided by Magnolia Press for private/research use. Commercial sale or deposition in a public library or website is prohibited. Syn. Paradasys cambriensis Boaden, 1963a:404 [Hummon, 1974a:195] Syn. Psammodasys cambriensis d’Hondt, 1974b:675 [Hummon, 2008b:121] Cephalodasys caudatus Rao, 1981:137 Cephalodasys hadrosomus Hummon, Todaro & Tongiorgi, 1993:112 Cephalodasys littoralis Renaud-Debyser, 1964:115 [D’Hondt & Balsamo (2009:274) consider the specimens studied by Renaud-Debyser (1964) as juveniles of C. turbanelloides (=Paradasys turbanelloides = Psammodasys turbanelloides); it is obvious that this cannot be the case as the drawing provided by Renaud-Debisser depicts a fully mature specimen and the sexual condition, along with morphometry of the studied animal, is clearly stated in her article. Furthermore, mature C. littoralis have been reported subsequently from France, Italy and the east coast of North America (Ruppert 1977 {figures}; Todaro et al. 2001, 2003; and Hummon 2009b {figures, N Amer and NW Europe databases}] Cephalodasys miniceraus Hummon, 1974c:401 Cephalodasys pacificus Schmidt, 1974:10 Cephalodasys palavensis Fize, 1963:713 Cephalodasys swedmarki Hummon, 2008b:119 Cephalodasys turbanelloides (Boaden, 1960):404 Syn. Paradasys turbanelloides Boaden, 1960:404 [Hummon, 1974a:195] Syn. Psammodasys turbanelloides d’Hondt, 1974b:675 [Hummon, 2008b:121] Genus Dolichodasys Gagne, 1977:20 Type-species Dolichodasys elongatus Gagne, 1977:20 Dolichodasys carolinensis Ruppert & Shaw, 1977:186 Dolichodasys delicatus Ruppert & Shaw, 1977:192 Syn. Paradasys hexadactylus Karling, 1954:14 sensu Thane-Fenchel, 1970:119 [Ruppert & Shaw, 1977:192] Genus Megadasys Schmidt, 1974:26 Syn. Thiodasys Boaden, 1974:371 [Hummon, 1982:861] Type-species Megadasys pacificus Schmidt, 1974:26 Megadasys minor Kisielewski, 1987:840 Megadasys sterreri (Boaden, 1974):371 Syn. Thiodasys sterreri Boaden, 1974:371 [Hummon, 1982:861] Genus Mesodasys Remane, 1951:102 Type-species Mesodasys laticaudatus Remane, 1951:102 Syn. Mesodasys lobocercus (Boaden, 1960):397 [W.D. Hummon in Jouk, Hummon, Hummon & Roidou, 1992:6] Syn. Cephalodasys lobocercus Boaden, 1960:397 [Boaden, 1963a:487] Mesodasys adenotubulatus Hummon, Todaro & Tongiorgi, 1993:114 Mesodasys britanicus Hummon, 2008b:121. [Note that the the gender ending should be changed from M. brittanica to M. brittanicus (ICZN 34.2) and with a justified emendation (ICZN 19.1) the spelling of the specific epithet is changed from M. brittanicus to M. britanicus, both in this publication] Mesodasys hexapodus Rao & Ganapati, 1968:37 Mesodasys ischiensis Hummon, Todaro & Tongiorgi, 1993:115 Mesodasys littoralis Remane, 1951:105 Mesodasys rupperti Hummon, 2008b:123 Genus Paradasys Remane, 1934:473 Type-species Paradasys subterraneus Remane, 1934:473 Paradasys bilobocaudus Hummon, 2008b:125 4 · Zootaxa 2392 © 2010 Magnolia Press HUMMON & TODARO TERMS OF USE This pdf is provided by Magnolia Press for private/research use. Commercial sale or deposition in a public library or website is prohibited. Paradasys hexadactylus Karling, 1954:14 Paradasys lineatus Rao, 1980:213 Paradasys littoralis Rao & Ganapati, 1968:35 Paradasys nipponensis Sudzuki, 1976:10 [See discussion of this species in Hummon, 2008b:127; although Paradasys nipponensis is described and figured sufficiently to meet the the minimum criteria for a species description (Hummon 2008), the information available is insufficient for reliable identification of the species; therefore we consider the taxon to be a species inquirenda] Paradasys pacificus Schmidt, 1974:13 Genus Pleurodasys Remane, 1927:208 [Transferred from Macrodasyidae to Lepidodasyidae by Hummon, 1974b:197] Type-species Pleurodasys helgolandicus Remane, 1927c:208 sensu Boaden, 1963a:496 Syn. Pleurodasys megasoma Boaden, 1963a:496 [W.D. Hummon in Jouk, Hummon, Hummon & Roidou, 1992:6; the synonym has been accepted and discussed further by Marotta, Todaro & Ferraguti, 2008:111] Pleurodasys glandulifera Boaden, 1963a:487 [In this publication. The species should be considered nomen nudum because the species appeared in the literature only as a name] Family Dactylopodolidae Strand, 1929:5. Includes 3 genera and 13 species [The nominal type genus Dactylopodola was published two months before the name Dactylopodalia, and so becomes the basis of the family name (ICZN 35.3); Dactylopodellidae Remane, 1927d:41 thus becomes a junior synonym of Dactylopodolidae by Strand, 1929:5 and Dactylopodaliidae Remane, 1929:176 becomes a junior synonym of Dactylopodolidae by Blake, 1933: 606] Type-Genus Dactylopodola Strand, March 1929:5 Syn. Dactylopodella Remane, 1926b:664 [Preoccupied, renamed by Strand, 1929:5] Syn. Dactylopodalia Remane, May 1929:176 [The name Dactylopodalia was published two months later than Dactylopodola, and so becomes a junior synonym of Dactylopodola, as stated by Blake, 1933:606] Type-species Dactylopodola baltica (Remane, 1926b):664 Syn. Dactylopodella baltica Remane, 1926b:664 [Strand, 1929:5] Syn. Dactylopodalia baltica (Remane, 1929):176 [Blake, 1933:606] Dactylopodola agadasys Hochberg, 2003:41 Dactylopodola australiensis Hochberg, 2003:39 Dactylopodola cornuta (Swedmark, 1956):45 Syn. Dactylopodalia cornuta Swedmark, 1956:45 [d’Hondt, 1971b:164] Syn. nov. Dactylopodola brevis d’Hondt & Balsamo, 2009:270 Syn. nov. Dactylopodalia cornuta var. brevis d'Hondt 1966a:4 [In a previous work (Hummon, 2007c), it was proposed that this infrasubspecific category be synonymized as a subadult stage under D. cornuta; d’Hondt & Balsamo (2009:270, with Figs. 4, 5, heretofore unavailable in published form) have rejected the proposed synonymy and have raised the infraspecific taxon to the species level (D. brevis), based on the shorter body, and location and orientation of the cephalic appendanges; while they have said that the body is shorter than in adults, never has any length been given. Figures shown in d’Hondt 1971 (Fig. 9 a, b) illustrate his case; the upper one (Fig. 9 a) is as it says from Swedmark’s original description; the lower one being from d’Hondt (1966a), wherein the appendages are shorter and are inserted relatively more posterior and oriented obliquely toward the posterior. Unfortunately, they appear to be drawn to the same scale, though no scales are given. If one looks at the figure of D. cornuta on the web site Hummon (2009b: Figures, D. cornuta), one sees that the original drawing of Swedmark erred in having the appendages too far forward and too perpendicular to the body; when this is considered, it is easy to see that d’Hondt’s ‘brevis’ specimen TAXONOMY OF MARINE GASTROTRICHA Zootaxa 2392 © 2010 Magnolia Press · 5 TERMS OF USE This pdf is provided by Magnolia Press for private/research use. Commercial sale or deposition in a public library or website is prohibited. is a subadult of about 190 µm in length of the body, or mid-way between the adult and the subadult of the Hummon (2009b) figure, with cephalic appendages midway between those of the two specimens; we thus consider ‘brevis’ to be a developmental stage of D. cornuta and reject ‘brevis’ as referring to any distinct infraspecific category or species taxon] Dactylopodola indica (Rao & Ganapati, 1968):45 Syn. Dactylopodalia indica Rao & Ganapati, 1968:45 [d’Hondt, 1971b:164] Dactylopodola mesotyphle Hummon, Todaro, Tongiorgi & Balsamo, 1998:109 Dactylopodola roscovita (Swedmark, 1967):325 Syn. Dactylopodalia roscovita Swedmark, 1967:325 [d’Hondt, 1971b:164] Dactylopodola typhle (Remane, 1927):213 [Strand, 1929:5] Syn. Dactylopodella typhle Remane, 1927:213 [Strand, 1929:5] Syn. Dactylopodalia typhle (Remane, 1927):213 [Blake, 1933:606] Dactylopodola weilli (d'Hondt, 1965:6) [Luporini, Magagnini & Tongiorgi, 1970:15 considered this species to be a junior synonym of D. typhle; the synonym was rejected by Kisielewski 1987:847, the species listed as Dactylopodola weilli (d'Hondt, 1965):6; the synonym was then reinstated by Todaro, Balsamo & Tongiorgi, 1992:473. Recently the synonym has been rejected once again by d’Hondt & Balsamo 2009:271, and though the latter authors do not bring any new evidence to sustain their statement, they reiterate as discriminatory traits between the two species features whose taxonomic unreliability has already been discussed by authors favorable to the synonymy. Recent DIC micrographs and drawings by Kinieke et al. (2008: Figs. 1, 4) of D. typhle, collected near the type locality, show a different arrangement of the anterior and lateral adhesive tubes than those shown by Remane, 1927: Fig. 6; indeed, the head shape and anterior adhesive tubes of putative D. weilli specimens studied by Kisielewski (1987) differ from those reported for the typical D. weilli studied by d’Hondt (1965), both from Bassin d’Arcachon. Neither d’Hondt or Kisielewski have reported seeing D. typhle; nor has any other worker ever reported seeing the presumptive D. weilli, suggesting that further work needs to be carried out before the status of D. weilli can be ascertained. This would include investigation of both type locales, and inclusion of other traits such as anatomy of the reproductive system. In the meantime, it is preferable to treat D. weilli as a species inquirenda] Genus Dendrodasys Wilke, 1954:507 Type-species Dendrodasys gracilis Wilke, 1954:507 Dendrodasys affinis Wilke, 1954:511 sensu Hummon, Todaro, Tongiorgi & Balsamo, 1998:112 Dendrodasys pacificus Schmidt, 1974:31 Dendrodasys ponticus Valkanov, 1957:385 Genus Dendropodola Hummon, Todaro & Tongiorgi, 1993:111 Type-species Dendropodola transitionalis Hummon, Todaro & Tongiorgi, 1993:111 Family Lepidodasyidae Remane, 1927d:41—Emended family—Includes 1 genus and 5 species [Emended in this publication; the data have already been published and there is a consensus among workers that a major source of polyphyly in the family lies in the difference between Lepidodasys and the remaining constituents of the former family Lepidodasyidae; thus it only remains to make the formal taxonomic separation. The reasons for separating Lepidodasys from the others are contained in the respective diagnoses. Lepidodasyidae emended diagnosis, modified from Ruppert, 1978a Elongate Macrodasyida, most nearly circular in transverse section, with small, nearly terminal mouth opening and no head delimitation. Cuticle elaborated as flat-scales, ribbed-scales or hat-shaped scales, often organized in crossed helical patterns. Multiciliated epidermal cells; ventral cilia in two longitudinal rows, often partially reduced. Epidermal glands with granular or fibrous inclusions, sometimes extremely 6 · Zootaxa 2392 © 2010 Magnolia Press HUMMON & TODARO TERMS OF USE This pdf is provided by Magnolia Press for private/research use. Commercial sale or deposition in a public library or website is prohibited. well developed. Anterior adhesive tubes (TbA) in two groups or a semicircle behind the mouth; lateral, dorsolateral and ventrolateral adhesive tubes (TbL/TbDL/TbVL) arranged in columns along the body; posterior adhesive tubes (TbP) arranged marginally around the blunt posterior end. Sphincter muscle developed around mouth opening; powerfully developed radial pharyngeal musculature; lacking striations; pharyngeal pores absent. Circular muscles absent from lateral regions of the body. Y-cells contain myofilaments. Gonads paired; male anterior, female posterior; male gametes mature posterior to anterior, female gametes posterior to anterior; ova enter the central body region, dorsal to the gut, immediately anterior to the seminal receptacle. Male pores paired or single, located anterior to the caudal organ, where present; caudal organ with two canals sharing common ventral opening in front of the anus; one weakly developed canal leads into the seminal receptacle; the other is a blind, glandulomuscular sac. An oviduct is present. Mostly subtidal in distribution; fine to coarse sand. Type-Genus Lepidodasys Remane, 1926b:684 Type-species Lepidodasys martini Remane, 1926b:684 Lepidodasys arcolepis Clausen, 2004b:428 Lepidodasys castoroides Clausen, 2004b:431 Lepidodasys platyurus Remane, 1927c:209 Lepidodasys unicarenatus Balsamo, Fregni & Tongiorgi, 1994:218 Family Macrodasyidae Remane, 1926b:723—Includes 2 genera and 43 species Type-Genus Macrodasys Remane, 1924:290 Type-species Macrodasys buddenbrocki Remane, 1924:290 Macrodasys achradocytalis Evans, 1994:243 Macrodasys acrosorus Hummon & Todaro, 2009:50 Macrodasys affinis Remane, 1936:169 Macrodasys africanus Remane, 1950:35 Macrodasys africanus ssp. ponticus Valkanov, 1957:389 [In this publication. The trinomial was described as a variety (Macrodasys africanus var. ponticus), but with the date coming before 1960, and not being proposed as an infrasubspecific category, it must be treated as a subspecies (ICZN 45.6.4)] Macrodasys ancocytalis Evans, 1994:240 Macrodasys andamanensis Rao, 1993:26 Macrodasys balticus Roszczak, 1939:6 [This species thus far has been found only in brackish-waters] Macrodasys blysocytalis Evans, 1994:250 Macrodasys caudatus Remane, 1927:204 Macrodasys celticus Hummon, 2008b:128 Macrodasys cephalatus Remane, 1927c:206 Macrodasys cunctatus Wieser, 1957:374 Macrodasys deltocytalis Evans, 1994:245 Macrodasys digronus Hummon & Todaro, 2009:51 Macrodasys dolichocytalis Evans, 1994:248 Macrodasys fornerisae Todaro & Rocha, 2004:1619 [Change of gender ending from M. fornerise to M. fornerisae, in this publication (ICZN 34.2)] Macrodasys gerlachi Papi, 1957:179 Macrodasys hexadactylis Rao, 1970:109 Macrodasys indicus Kutty & Nair, 1969:632 [The proper name is M. indicus, as in the text, and not M. idicus, as occurs in Fig. 7; this is as determined by the first reviser, Hummon, 2008b:127 (ICZN 24.2.2), where its morphological relationships were discussed; however, the information available is insufficient for reliable identification of the species; therefore we consider the taxon to be a species inquirenda] TAXONOMY OF MARINE GASTROTRICHA Zootaxa 2392 © 2010 Magnolia Press · 7 TERMS OF USE This pdf is provided by Magnolia Press for private/research use. Commercial sale or deposition in a public library or website is prohibited. Macrodasys lakshadweepensis Hummon, 2008b:127 [Change of gender ending from M. lakshadweepense to M. lakshadweepensis, in this publication (ICZN 34.2)] Syn. Macrodasys indica Rao, 1991:63 [When gender corrected to M. indicus, this name became a junior primary homonym, which was replaced in accordance with ICZN 23.3.5 and 60.1] Macrodasys meristocytalis Evans, 1994:247 Macrodasys neapolitanus Papi, 1957:181 Macrodasys nobskaensis Hummon, 2008b:130 Macrodasys pacificus Schmidt, 1974:16 Macrodasys plurosorus Hummon, 2008b:131 Macrodasys remanei Boaden, 1963a:499 Macrodasys stenocytalis Evans, 1994:252 Macrodasys thuscus Luporini, Magagnini & Tongiorgi, 1973:284 Macrodasys waltairensis Rao & Ganapati, 1968:39 [The proper name is M. waltairensis, as in the text, and not M. waltaironsis, as occurs in Fig. 7; this is as determined by the first reviser Hummon, 2008:127 (ICZN 24.2.2)] Genus Urodasys Remane, 1926b:687 Type-species Urodasys mirabilis Remane, 1926b:687 Syn. Urodasys roscoffensis Kisielewski, 1987:843 [W.D. Hummon in Jouk Hummon, Hummon & Roidou, 1992:6] Urodasys acanthostylis Fregni, Tongiorgi & Faienza, 1998:378 Urodasys anorektoxys Todaro, Bernhard & Hummon, 2000:468 Urodasys apuliensis Fregni, Faienza, Grimaldi, Tongiorgi & Balsamo, 1999:186 Urodasys bucinastylis Fregni, Faienza, Grimaldi, Tongiorgi & Balsamo, 1999:188 Urodasys calicostylis Schoepfer-Sterrer, 1974:241 Urodasys cornustylis Schoepfer-Sterrer, 1974:234 Urodasys elongatus Renaud-Mornant, 1969:384 Urodasys nodostylis Schoepfer-Sterrer, 1974:239 Urodasys remostylis Schoepfer-Sterrer, 1974:243 Urodasys spirostylis Schoepfer-Sterrer, 1974:236 Urodasys uncinostylis Fregni, Tongiorgi & Faienza, 1998:378 Urodasys viviparus Wilke, 1954:502 Family Planodasyidae Rao & Clausen, 1970:78—Includes 2 genera and 7 species Type-Genus Planodasys Rao & Clausen, 1970:75 Type-species Planodasys marginalis Rao & Clausen, 1970:75 Planodasys littoralis Rao, 1993:38 Genus Crasiella Clausen, 1968:59 Type-species Crasiella diplura Clausen, 1968:59 Crasiella azorensis Hummon, 2008b:135 Crasiella indica Rao, 1981 (for 1980):3 Crasiella oceanica d’Hondt, 1974b:667 Crasiella pacifica Schmidt, 1974:40 Family Thaumastodermatidae Remane, 1927c:237—Includes 8 genera and 145 species Subfamily Diplodasyinae Ruppert, 1978:113 Type-Genus Diplodasys Remane, 1927:228 8 · Zootaxa 2392 © 2010 Magnolia Press HUMMON & TODARO TERMS OF USE This pdf is provided by Magnolia Press for private/research use. Commercial sale or deposition in a public library or website is prohibited. Type-species Diplodasys platydasyoides Remane, 1927c:228 Diplodasys ankeli Wilke, 1954:517 Diplodasys caudatus Kisielewski, 1987:865 Diplodasys meloriae Todaro, Balsamo & Tongiorgi, 1992:475 Diplodasys minor Remane, 1936:178 sensu Todaro, 1992:324 Diplodasys pacificus Schmidt, 1974:50 [Raised from subspecies Diplodasys ankeli ssp. pacificus Schmidt, 1974:50 to species level by Clausen 2004:435] Diplodasys remanei Rao & Ganapati, 1968:49 Diplodasys sanctimariae Hummon & Todaro, 2009:56 Diplodasys swedmarki Kisielewski, 1987:867 Genus Acanthodasys Remane, 1927c:212 [transferred from Lepidodasyidae to Thaumastodermatidae by Hummon, 1974b:198] Type-species Acanthodasys aculeatus Remane, 1927c:212 Acanthodasys algarvensis Hummon, 2008b:136 [Change of gender ending from A. algarvense to A. algarvensis, in this publication (ICZN 34.2)] Acanthodasys arcassonensis Kisielewski, 1987:862 Acanthodasys carolinensis Hummon, 2008b:138 Acanthodasys diplodasyoides Ruppert, 1978:113 [The species should be considered nomen nudum as stated by Kisielewski, 1988: 864] Acanthodasys fibrosus Clausen, 2004a:137 Acanthodasys flabellicaudus Hummon & Todaro, 2009:53 Acanthodasys lineatus Clausen, 2000:364 Acanthodasys platydasyoides Ruppert, 1978:113 [The species should be considered nomen nudum as stated by Kisielewski, 1988: 864] Acanthodasys silvulus Evans, 1992:316 Acanthodasys tetranchyrodermatoides Ruppert, 1978:94 [The species should be considered nomen nudum as stated by Kisielewski, 1988: 864] Acanthodasys thrinax Ruppert, 1978:98 [The species should be considered nomen nudum as stated by Kisielewski, 1988: 864] Acanthodasys vermiformis Ruppert, 1978:94 [The species should be considered nomen nudum as stated by Kisielewski, 1988: 864] Subfamily Thaumastodermatinae Remane, 1927c: ICZN 36 [Ruppert, 1978:109] Type-Genus Thaumastoderma Remane, 1926b:671 Type-species Thaumastoderma heideri Remane, 1926b:671 Thaumastoderma appendiculatum Chang, Lee & Clausen, 1998a:332 Thaumastoderma arcassonense d’Hondt, 1965:10 Thaumastoderma bifurcatum Clausen, 1991:158 Thaumastoderma cantacuzeni Lévi, 1958:204 Thaumastoderma clandestinum Chang, Kubota & Shirayama, 2002:774 Thaumastoderma copiophorum Chang, Lee & Clausen, 1998b:317 Thaumastoderma coronarium Chang, Lee & Clausen, 1998a:330 Thaumastoderma mediterraneum Remane, 1927c:229 [Change of gender ending from T. mediterranea to T. mediterraneum, following Luporini, Magagnini & Tongiorgi, 1973: 275 (ICZN 34.2)] Thaumastoderma minancrum Hummon, 2008b:153 Thaumastoderma moebjergi Clausen, 2005:441 Thaumastoderma natlanticum Hummon, 2008b:155 Thaumastoderma ramuliferum Clausen, 1965c:24 TAXONOMY OF MARINE GASTROTRICHA Zootaxa 2392 © 2010 Magnolia Press · 9 TERMS OF USE This pdf is provided by Magnolia Press for private/research use. Commercial sale or deposition in a public library or website is prohibited. Thaumastoderma renaudae Kisielewski, 1987:859 Thaumastoderma swedmarki Lévi, 1950:34 Thaumastoderma truncatum Clausen, 1991:162 Genus Oregodasys Hummon, 2008b:140 Syn. Platydasys Remane, 1927c:222 [Preoccupied; renamed by Hummon 2008b:140] Type-species Oregodasys maximus (Remane, 1927c):222 Syn. Platydasys maximus Remane, 1927c:222 [Hummon, 2008b:140] Oregodasys maximus ssp. celticus (Swedmark, 1955b):12 [In this publication; the trinomial was described as a variety (Platydasys maximus var. celticus), but the date coming before 1960 and not being proposed as an infrasubspecific category, it must be treated as a subspecies (ICZN 45.6.4)] Oregodasys itoi (Chang, Kubota & Shirayama, 2002):77 [Hummon, 2008b:140] Oregodasys mastigophorus (Clausen, 1965c):27 [Hummon, 2008b:140] Oregodasys kurnowensis Hummon, 2008b:140 Oregodasys ocellatus (Clausen, 1965c):31 [Hummon, 2008b:140] Oregodasys pacificus (Schmidt, 1974):55 [Hummon, 2008b:140] Oregodasys phacellatus (Clausen, 1965c):30 [Hummon, 2008b:140] Oregodasys rarus (Forneris, 1961):216 [Hummon, 2008b:140] Oregodasys ruber (Swedmark, 1956a):80 [Hummon, 2008b:140] Oregodasys styliferus (Boaden, 1965):221 [Hummon, 2008b:140] Oregodasys tentaculatus (Swedmark, 1956a):83 [Hummon, 2008b:140] Platydasys schultzi Remane, 1940:xx [The species should be considered nomen nudum as stated by d’Hondt, 1971:173] Genus Pseudostomella Swedmark, 1956b:53 Type-species Pseudostomella roscovita Swedmark, 1956b:53 Pseudostomella andamanica Rao, 1993:36 Pseudostomella cataphracta Ruppert, 1970:128 Pseudostomella cheraensis Pryialakshmi, Menon & Todaro, 2007:61 Pseudostomella etrusca Hummon, Todaro & Tongiorgi, 1993:117 Pseudostomella faroensis Clausen, 2004b:444 Pseudostomella indica Rao, 1970:115 Pseudostomella klauserae Hochberg, 2003a:572 Pseudostomella koreana Lee & Chang, 2002:209 Pseudostomella longifurca Lee & Chang, 2002:208 Pseudostomella malayica Renaud-Mornant, 1967b:209 Pseudostomella megapalpator Hochberg, 2003a:575 Pseudostomella plumosa Ruppert, 1970:123 Pseudostomella triancra Hummon, 2008b:142 Genus Ptychostomella Remane, 1926b:678 Type-species Ptychostomella pectinata Remane, 1926b:678 Ptychostomella bergensis Clausen, 1996:126 Ptychostomella brachycephala (Levi, 1954):39 Syn. Platydasys brachycephalus Levi, 1954:39 [Clausen, 2004b:447] Ptychostomella helana Roszczak, 1939:9 [This species thus far has been found only in brackish-waters] Ptychostomella higginsi Clausen, 2004b:450 Ptychostomella jejuensis Lee, Hwang & Chang, 2009:26 Ptychostomella lepidota Clausen, 2000:375 Ptychostomella mediterranea Remane, 1927c:226 10 · Zootaxa 2392 © 2010 Magnolia Press HUMMON & TODARO TERMS OF USE This pdf is provided by Magnolia Press for private/research use. Commercial sale or deposition in a public library or website is prohibited. Ptychostomella ommatophora Remane, 1927c:225 Ptychostomella orientalis Lee & Chang, 2003:482 Ptychostomella papillata Lee & Chang, 2003:485 Ptychostomella tyrrhenica Hummon, Todaro & Tongiorgi, 1993:118 Genus Tetranchyroderma Remane, 1926b:676 Syn. Echinodasys Remane, 1926 [Remane, 1936:178] Type-species Tetranchyroderma hystrix Remane, 1926b:676 [Tetranchyroderma hystrix was described as the type species for a genus having tetrancres by Remane, 1926, when he separated species that had tetrancres (Tetranchyroderma) from species that had pentancres (Echinodasys). A decade later, Remane (1936:178) realized that all species known at that time, including both T. hystrix and T. apus, had pentancres, and so synonymized Echinodasys with Tetranchyroderma] Tetranchyroderma adeleae Hochberg, 2009:730 Tetranchyroderma anomalopsum Hummon, Todaro, Balsamo & Tongiorgi, 1996:73 Tetranchyroderma antenniphorum Hummon & Todaro—new species. [In this publication] Syn. Tetranchyroderma antennatum Luporini, Magagnini & Tongiorgi, 1973a:114 [Name not available, because it was previously used as an infrasubspecific trinomial (Tetranchyroderma hystrix var. antennata d'Hondt, 1968b:395). By treating the trinomial as a variety after 1960, this name is rendered an infrasubspecific category, and hence is not available to be named a species (ICZN 45.6.3)] Tetranchyroderma aphenothigmum Hummon, Todaro, Tongiorgi & Balsamo, 1998:114 Tetranchyroderma apum Remane, 1927c:231 [Change of gender ending from T. apus to T. apum, following Todaro, Hummon, Fregni, Balsamo & Tongiorgi, 2001:124 (ICZN 34.2)] Tetranchyroderma arcticum Clausen, 1999:369 Tetranchyroderma australiense Nicholas & Todaro, 2006:254 Tetranchyroderma boadeni Schrom, in Riedl 1970:217 sensu Schrom 1972:301 Tetranchyroderma boreale Clausen, 2000:370 Tetranchyroderma bulbosum Clausen, 2000:371 Tetranchyroderma bunti (Thane-Fenchel, 1970):124 Syn. Thaumastoderma bunti Thane-Fenchel, 1970:124 [d’Hondt, 1974:232] Tetranchyroderma canariense Todaro, Ancona, Marzano, D’Addabbo & De Zio Grimaldi, 2003:192 Tetranchyroderma cirrophorum Lévi, 1950:33 [Change of gender ending from T. cirrophora to T. cirrophorum, following Todaro, Hummon, Fregni, Balsamo & Tongiorgi, 2001:124 (ICZN 34.2)] Tetranchyroderma coeliopodium Boaden, 1963b:375 Tetranchyroderma dendricum Saito, 1937:263 Tetranchyroderma dragescoi Swedmark, 1967:324 Tetranchyroderma enallosum Hummon, 1977:119 [Change of gender ending from T. enallosa to T. enallosum, in this publication (ICZN 34.2)] Tetranchyroderma esarabdophorum Tongiorgi & Balsamo, 1984:335 Tetranchyroderma faroense Clausen, 2004b:438 Tetranchyroderma forceps d’Hondt & Balsamo 2009:276. [The species has recently been described from a single specimen found in the 1960s. By admission of the authors, the description is incomplete. The main autoapomorphy of the taxon would be two pairs of prehensile cephalic appendances mimicking two tongs: a trait unique within Gastrotricha, which makes its existence doubtful without verifiable documentation. Unfortunately, the specimen examined was destroyed during the study, no drawings have been provided and the only picture supplied is of very poor quality, to an extent that it is questionable whether it can be assigned to a thaumastodermatid gastrotrich, without reading the legend. Consequenty we consider the taxon to be a species inquirenda and reject its assignation to the subgenus created for it] Tetranchyroderma gausancrum Hummon, 2008b:144 Tetranchyroderma gracilium Chang, Lee & Clausen, 1998b:315 TAXONOMY OF MARINE GASTROTRICHA Zootaxa 2392 © 2010 Magnolia Press · 11 TERMS OF USE This pdf is provided by Magnolia Press for private/research use. Commercial sale or deposition in a public library or website is prohibited. Tetranchyroderma heterotentaculatum Chang & Lee, 2001:188 Tetranchyroderma heterotubulatum Hummon, Todaro & Tongiorgi, 1993:120 Tetranchyroderma hirtum Luporini, Magagnini & Tongiorgi, 1973a:272 Tetranchyroderma hoonsooi Chang & Lee, 2001:191 Tetranchyroderma hyponiglarum Hummon & Todaro, 2009:59 Tetranchyroderma hypopsilancrum Hummon, Todaro & Tongiorgi, 1993:122 Tetranchyroderma inaequitubulatum Todaro, Balsamo & Tongiorgi, 2002:253 Tetranchyroderma indicum Rao & Ganapati, 1968:46 [Change of gender ending from T. indica to T. indicum, in this publication (ICZN 34.2)] Tetranchyroderma insulare Balsamo, Fregni & Tongiorgi, 1994:221 Tetranchyroderma interstitiale Hummon, 2008b:146 [Change of gender ending from T. interstitialis to T. interstitiale, in this publication (ICZN 34.2)] Tetranchyroderma kontosomum Hummon, Todaro, Balsamo & Tongiorgi, 1996:75 Tetranchyroderma korynetum Hummon & Todaro, 2009:61 Tetranchyroderma lameshurense Hummon, 2008b:148 [Change of gender ending from T. lameshurensis to T. lameshurense, in this publication (ICZN 34.2)] Tetranchyroderma littorale Rao, 1981a:140 [Change of gender ending from T. littoralis to T. littorale, in this publication (ICZN 34.2)] Tetranchyroderma longipedum Hummon, 2008b:151 Tetranchyroderma massilense Swedmark, 1956a:77 Tetranchyroderma megastomum (Remane, 1927c):234 Syn. Echinodasys megastoma Remane, 1927c:234 [Remane, 1936:178. Change of gender ending from T. megastoma to T. megastomum, following Todaro, Hummon, Fregni, Balsamo & Tongiorgi, 2001:126 (ICZN 34.2)] Tetranchyroderma monokerosum Lee & Chang, 2007:477 Tetranchyroderma multicirratum Lee & Chang, 2007:475 Tetranchyroderma norvegicum Clausen, 1996:124 Tetranchyroderma oligopentacrum Hummon & Todaro, 2009:64 Tetranchyroderma pachysomum Hummon, Todaro & Tongiorgi, 1993:124 Tetranchyroderma pacificum Schmidt, 1974:45 Tetranchyroderma papii Gerlach, 1953:205 Syn. Tetranchyroderma bunti (Thane-Fenchel, 1970):124 [d’Hondt, 1974:232; rejected by Hummon, 1974:212] Tetranchyroderma paradoxum Thane-Fenchel, 1970:122 [Change of gender ending from T. paradoxa to T. paradoxum, in this publication (ICZN 34.2)] Tetranchyroderma paralittorale Rao, 1991:67 [Change of gender ending from T. paralittoralis to T. paralittorale, in this publication (ICZN 34.2)] Tetranchyroderma parapapii Hummon, 2009a:123 Tetranchyroderma pentasperum Nicholas & Todaro, 2006:250 [Change of gender ending from T. pentasperus to T. pentasperum, in this publication (ICZN 34.2)] Tetranchyroderma polyacanthum (Remane, 1927c):232 Syn. Echinodasys polyacanthus Remane, 1927c:232 [Remane, 1936:178. Change of gender ending from T. polyacanthus to T. polyacanthum, in this publication (ICZN 34.2)] Tetranchyroderma polypodium Luporini, Magagnini & Tongiorgi, 1971:436 Tetranchyroderma polyprobolostomum Hummon, Todaro, Balsamo & Tongiorgi, 1996:76 Tetranchyroderma psilotopum Hummon, Todaro, Tongiorgi & Balsamo, 1998:116 Tetranchyroderma pugetense Wieser, 1957:380 Tetranchyroderma quadritentaculatum Todaro, Balsamo & Tongiorgi, 1992:479 Tetranchyroderma sanctaecaterinae Todaro, Balsamo & Tongiorgi, 1992:480 Tetranchyroderma sardum Todaro, Balsamo & Tongiorgi, 1988:69 12 · Zootaxa 2392 © 2010 Magnolia Press HUMMON & TODARO TERMS OF USE This pdf is provided by Magnolia Press for private/research use. Commercial sale or deposition in a public library or website is prohibited. Tetranchyroderma schizocirratum Chang, Kubota & Shirayama, 2002:770 Tetranchyroderma suecicum Boaden, 1960:402 [Change of gender ending from T. suecica to T. suecicum, following Luporini, Magagnini & Tongiorgi, 1973: 275 (ICZN 34.2)] Tetranchyroderma swedmarki Rao & Ganapati, 1968:48 Tetranchyroderma symphorochetum Hummon, Todaro, Tongiorgi & Balsamo, 1998:117 Tetranchyroderma tanymesatherum Hummon, Todaro, Balsamo & Tongiorgi, 1996:78 Tetranchyroderma tentaculatum Rao, 1993:31 [Change of gender ending from T. tentaculata to T. tentaculatum, in this publication (ICZN 34.2)] Tetranchyroderma thysanogaster Boaden, 1965:219 Tetranchyroderma thysanophorum Hummon, Todaro & Tongiorgi, 1993:125 Tetranchyroderma tribolosum Clausen, 1965a:10 Tetranchyroderma verum Wilke, 1954:513 [Change of gender ending from T. vera to T. verum, following Todaro, Hummon, Fregni, Balsamo & Tongiorgi, 2001:128 (ICZN 34.2)] Tetranchyroderma weissi Todaro, 2002:558 Genus Hemidasys Claparède, 1867:18 Type-species Hemidasys agaso Claparède, 1867:19 [The monospecific genus Hemidasys agaso was found but once, abundant in the finest mud of the harbor at Naples, as both a free-living and an epizootic with the polychaete Neanthes caudatus (Delle Chiaje, 1828), the latter being found widely in Italy today (Castelli, Bianchi, Cantone, Cinar, Gambi, Giangrande, Iraci Sareri, Lanera, Licciano, Musco, Sanfilippo & Simonini 2008), but never again with a gastrotrich symbiont. Boaden’s brief examination (1965a:224) of the harbor mud failed to yield either the gastrotrich or its epizootic polychaete. Extensive work in the gulf of Naples from the 1920’s through the 1990’s has failed to find this species; in our opinion if the Claparède species ever existed it presently should be considered extinct owing to the repeated efforts to find it being unsuccessful and the known degradation of its habitat] Family Turbanellidae Remane, 1926b:723—Includes 6 genera and 52 species Type-Genus Turbanella Schultze, 1853:243 Syn. Zelinkia Giard, 1904b:1063 [Remane, 1925b:312] Type-species Turbanella hyalina Schultze, 1853:243 Turbanella ambronensis Remane, 1943:239 Syn. Turbanella italica Gerlach, 1953:203 [Todaro, Hummon, Fregni, Balsamo & Tongiorgi 2001:132] Syn. Turbanella cirrata Papi, 1957:176 [Schmidt & Teuchert, 1969:15] Syn. Turbanella digitifera d'Hondt, 1965:8 [Hummon, 1969:90. [After describing the binomial as a distinct species in 1965, the author treated it in partial synonymy as T. ambronensis var. digitifera d’Hondt, 1970:167, where the trinomial becomes a variety of its senior binomial, though the variety coming after 1960 renders this name an infrasubspecific category, and hence unavailable except as a junior synonym (ICZN 45.6.3)] Turbanella aminensis Rao, 1991:71 Turbanella bengalensis Rao & Ganapati, 1968:40 Turbanella bocqueti Kaplan, 1958:31 sensu Boaden, 1974:369 Syn. Turbanella thiophila Boaden, 1974:369 [W.D. Hummon in Jouk, Hummon, Hummon & Roidou, 1992:6; the synonymy was accepted and discussed further in Todaro et al., 2001: 132] Turbanella brusci Hochberg, 2002:313 Turbanella caledoniensis Hummon, 2008b:167 Turbanella corderoi Dioni, 1960:121 Turbanella cornuta Remane, 1925b:309 Turbanella indica Rao, 1981:139 Turbanella lutheri Remane, 1952:62 TAXONOMY OF MARINE GASTROTRICHA Zootaxa 2392 © 2010 Magnolia Press · 13 TERMS OF USE This pdf is provided by Magnolia Press for private/research use. Commercial sale or deposition in a public library or website is prohibited. Turbanella lutheri ssp. scanica Wieser, 1954:152 [The trinomial was described as a variety (Turbanella lutheri var. scanica), but the date coming before 1960 and not being proposed as an infrasubspecific category, it must be treated as a subspecies (ICZN 45.6.4)] Turbanella mikrogada Hummon, 2008b:168 Turbanella multidigitata Kisielewski, 1987:850 Turbanella mustela Wieser, 1957:377 Turbanella ocellata Hummon, 1974a:433 Turbanella otti Schrom, 1972:293 Turbanella pacifica Schmidt, 1974:33 Turbanella palaciosi Remane, 1953:272 Turbanella petiti Remane, 1952:64 Turbanella plana (Giard, 1904b):1063 Syn. Zelinkia plana Giard, 1904b:1063 [Remane, 1925b:312. D’Hondt, 1971b:166, considered the species a synonym of T. cornuta; this was reiterated by d’Hondt & Balsamo 2009, since T. cornuta was the only species in the genus with cephalic horns that d’Hondt (1968a) found anywhere near Ambleteuse (the type-locality of T. plana). We disagree for the following reasons: more recent records by Hummon & Roidou (Aug. 1979), showed T. cornuta and T. bocqueti, both members of the genus having cephalic horns, at Ambleteuse and Wimereux, the two locations referred to by d’Hondt, yielding not just one but two potential candidates for synonymy (see Hummon 2009b). Furthermore, d’Hondt & Balsamo (2009) say that in absence of type material they cannot be sure about their proposed synonymy (sic!); given these uncertainties, there seems to be no choice here but to consider T. plana as a species inquirenda] Turbanella pontica Valkanov, 1957:385 Turbanella reducta Boaden, 1974:367 Turbanella remanei Forneris, 1961:313 Turbanella scilloniensis Hummon, 2008b:170 Turbanella subterranea Remane, 1934:475 Turbanella varians Maguire, 1976a:186 Turbanella veneziana Schrom, 1972:296 Genus Desmodasys Clausen, 1965:17 Type-species Desmodasys phocoides Clausen, 1965:17 Desmodasys abyssalis Kieneke & Zekely, 2007:1 Desmodasys borealis Clausen, 2000:362 Genus Dinodasys Remane, 1927c:217 Type-species Dinodasys mirabilis Remane, 1927c:217 Dinodasys delawarensis Hummon, 2008b:157 Genus Paraturbanella Remane, 1927c:220 Type-species Paraturbanella dohrni Remane, 1927c:220 Paraturbanella aggregotubulata Evans, 1992:318 Paraturbanella armoricana (Swedmark, 1954):469 Syn. Turbanella armoricana Swedmark, 1954b:469 [Wieser, 1957: 378] Paraturbanella boadeni Rao & Ganapati, 1968:42 Paraturbanella brevicaudata Rao, 1991:73 Paraturbanella cuanensis Maguire, 1976b:407 Paraturbanella eireanna Maguire, 1976b:405 Paraturbanella intermedia Wieser, 1957:379 Paraturbanella manxensis Hummon, 2008b:160 14 · Zootaxa 2392 © 2010 Magnolia Press HUMMON & TODARO TERMS OF USE This pdf is provided by Magnolia Press for private/research use. Commercial sale or deposition in a public library or website is prohibited. Paraturbanella mesoptera Rao, 1970:112 Paraturbanella pacifica Schmidt, 1974:37 [Raised from subspecies Paraturbanella pallida ssp. pacifica Schmidt, 1974:37 to species level by Hummon, 2008b:159] Paraturbanella pallida Luporini, Magagnini & Tongiorgi, 1971:444 Paraturbanella palpibara Rao & Ganapati, 1968:43 Paraturbanella pediballetor Hummon, 2008b:162 Paraturbanella scanica Clausen, 1996:120 Paraturbanella solitaria Todaro, 1995:554 Paraturbanella stradbroki Hochberg, 2002a:313 [The proper name is P. stradbroki, as in the text, and not P. helicostoma, as occurs in Figs. 2 and 3; this is as determined by the first reviser: Hummon, 2008b:160, ICZN 24.2.2] Paraturbanella teissieri Swedmark, 1954a:47 Syn. Paraturbanella microptera Wilke, 1954:507 [Schmidt & Teuchert, 1969:12] Genus Prostobuccantia Evans & Hummon, 1991:323 Type-species Prostobuccantia brocha Evans & Hummon, 1991:323 Genus Pseudoturbanella d'Hondt, 1968b:401 Type-species Pseudoturbanella stylifera d’Hondt, 1968b:401 Family Xenodasyidae Todaro, Guidi, Leasi & Tongiorgi, 2006:1006—Includes 2 genera and 4 species Type-Genus Xenodasys Swedmark, 1967:327 Syn. Chordodasys Schöpfer-Sterrer, 1969:392 [Hummon, 1982:858; the synonymy was accepted and discussed further in Todaro, Guidi, Leasi & Tongiorgi., 2006: 1012] Type-species Xenodasys sanctigoulveni Swedmark, 1967:327 sensu Kisielewski, 1984:855[Though Swedmark hyphenated the specific epithet, it should be rendered as unhyphenated, X. sanctigoulvini, in accordance with ICZN 32.5.2.3; the synonymy was accepted and discussed further in Todaro, Guidi, Leasi & Tongiorgi, 2006: 1012] Xenodasys eknomios Todaro, Guidi, Leasi & Tongiorgi, 2006:1006 Xenodasys riedli (Schoepfer-Sterrer, 1969):392 Syn. Chordodasys riedli Schoepfer-Sterrer, 1969:392 [Hummon, 1982:858; confirmed by Kisielewski, 1987:857 and Todaro, Guidi, Leasi & Tongiorgi, 2006c:1014 with further discussion] Genus Chordodasiopsis Todaro, Guidi, Leasi & Tongiorgi, 2006:1015 Type-species Chordodasyopsis antennatus (Rieger, Ruppert, Rieger & Schoepfer-Sterrer, 1974):220 [Todaro, Guidi, Leasi & Todaro, 2006:1015] Syn. Xenodasys antennatus (Rieger, Ruppert, Rieger & Schoepfer-Sterrer, 1974):220 [Todaro, Guidi, Leasi & Tongiorgi, 2006:1015] Syn. Chordodasys antennatus Rieger, Ruppert, Rieger & Schoepfer-Sterrer, 1974:220 [Hummon, 1982:858, but see Todaro, Guidi, Leasi & Tongiorgi, 2006: 1005 for further discussion] Order Chaetonotida Remane, 1925a:125—Includes 13 genera that are wholly or partially marine and 133 marine or brackish-water and estuarine species [The name as given by Remane had an -oidea ending, that of a superfamily (ICZN Article 29.2), which was first altered to a preferred ordinal –ida ending by Brunson (1950:328) and later to be formally so designated by Rao, 1970: 109 and Rao & Clausen, 1970:80, and has almost universally been accepted as emended] Suborder Multitubulatina d’Hondt, 1971:145 [The name as given by d’Hondt was Multitubulata, which was first emended to an -ina ending by Hummon, TAXONOMY OF MARINE GASTROTRICHA Zootaxa 2392 © 2010 Magnolia Press · 15 TERMS OF USE This pdf is provided by Magnolia Press for private/research use. Commercial sale or deposition in a public library or website is prohibited. 1982:862, i.e. from an order to a sub-order ending, and the name in this latter form has been almost universally accepted] Family Neodasyidae Remane, 1929:171— Includes 1 genus and 3 species [Transferred from Macrodasyida to Chaetonotida by Remane, 1936:181] Type-Genus Neodasys Remane, 1927c:216 Type-species Neodasys chaetonotoideus Remane, 1927c:216 Neodasys cirritus Evans, 1992:321 [With a justified emendation (ICZN 19.1) in this publication, the spelling of the specific epithet is changed from N. ciritus to N. cirritus, which in Latin refers to the tendrils (the small beard of elongate cilia) on the ventral surface that Evans was using to name the species (ICZN 33.2); the emended spelling was first used by Hochberg (2005:317), though he did not note that it was a change from the original] Neodasys uchidai Remane, 1961:85 Suborder Paucitubulatina d’Hondt, 1971:148 [The name as given by d’Hondt was Paucitubulata, which was first emended to an -ina ending by Hummon, 1982:862, i. e. from an order to a sub-order ending, and name in this latter form has been almost universally accepted] Family Chaetonotidae Gosse, 1864:392 sensu Leasi & Todaro, 2008:396. Includes 7 genera with 101 marine or brackish-water species. Syn. Ichthydina Ehrenberg, 1830:64 (ICZN 23.9.3) [Gosse somewhat arbitrarily inserted the name Chaetonotidae for the family because he thought it “desirable that the family should be named after the most characteristic and populous genus”. As noted by Weiss, 2001: 312 (ICZN 23.3) Chaetonotidae when proposed was invalid for the taxon it now represents, but in accordance with ICZN 23.9.3 Chaetonotidae becomes valid as a nomen protectum, replacing Ichthydina, which becomes a nomen oblitum] Subfamily Chaetonotinae Gosse, 1864: ICZN 36.1 [Kisielewski, 1991:10] Type-Genus Chaetonotus Ehrenberg, 1830:64 Type-subgenus Chaetonotus Ehrenberg, 1830:64 (ICZN 44.1) sensu stricto Syn. Euchaetonotus Schwank, 1990:147 [Weiss, 2001:313 first pointed out that the subgenus Euchaetonotus is invalid; because this subgenus contains the type species of the genus, it must bear the same name as the genus, ICZN 44.1] Type-species Chaetonotus (Chaetonotus) larus (Müller, 1773):95 [Ehrenberg 1930:64. Subgenus assigned by Kisielewski, 1997:148] (This is a freshwater taxon that has occasionally been reported from brackish-water, e. g. Galliford & Williams 1951) 18th & 19th century synonyms. Trichoda acarus, T. anas and Trichoda larus Müller, 1776; Leucophra larus Bory de St.Vincent, 1824; Diceratella larus Bory de St.Vincent, 1826 (see Ehrenberg, 1838:390 for details); Chaetonotus maximus, C. larus and C. brevis Metschnikoff, 1865; Ichthydium larus Ludwig, 1875 (see Zelinka, 1889:340 for details). These names have not been used for species identification in the 20th or 21st centuries)] Chaetonotus (Chaetonotus) aegilonensis Balsamo, Todaro & Tongiorgi, 1992:488 [Subgenus assigned by Kisielewski, 1997:148] Chaetonotus (Chaetonotus) angustus Schrom, 1972:310 [Subgenus assigned by Kisielewski, 1997:148] Chaetonotus (Chaetonotus) apechochaetus Hummon, Balsamo & Todaro, 1992:502 [Subgenus assigned by Kisielewski, 1997:148] Chaetonotus (Chaetonotus) condensus Mock, 1979:37 [Subgenus assigned by Kisielewski, 1997:148] 16 · Zootaxa 2392 © 2010 Magnolia Press HUMMON & TODARO TERMS OF USE This pdf is provided by Magnolia Press for private/research use. Commercial sale or deposition in a public library or website is prohibited. Chaetonotus (Chaetonotus) fenchelae d’Hondt, March 1974a:233 [Subgenus assigned by Kisielewski, 1997:148; change of gender ending from C. fencheli to C. fenchelae, in this publication (ICZN 34.2)] Syn. Chaetonotus pusillus Thane-Fenchel, 1970:130 [Preoccupied by C. pusillus Daday, 1905:76] New syn. Chaetonotus acareus Hummon, May 1974a:196 [In this publication; the species, being described two months later, should be considered a junior synonym of C. (C.) fenchelae] Chaetonotus (Chaetonotus) schoepferae Thane-Fenchel, 1970:132 [Subgenus assigned by Kisielewski, 1997:148; change of gender ending from C. schoepferi to C. schoepferae, in this publication (ICZN 34.2)] Chaetonotus (Chaetonotus) ichthydioides Tongiorgi, Fregni & Balsamo, 1999:587 [This species thus far has been found only in brackish waters] Chaetonotus (Chaetonotus) linguaeformis Voigt, 1902:116 [Transferred from subgenus Bifasciculatella by Kisielewski, 1997:148] (This is mostly a freshwater taxon, occasionally found in estuarine waters, e. g. Boaden 1976) Chaetonotus (Chaetonotus) magnificus Balsamo, Hummon, Todaro & Tongiorgi, 1997:84 [Subgenus assigned in this publication] Chaetonotus (Chaetonotus) maximus Ehrenberg, 1831:153. [Subgenus assigned by Kisielewski, 1997:148. This is mostly a freshwater taxon, sometimes found in estuarine and brackish-waters, e. g. Boaden 1976] 19th century synonyms. Chaetonotus gracilis Gosse, 1864:399; Chaetonotus larus and C. brevis Metschnikoff, 1865; Ichthydium maximum Ludwig, 1876; Ichthydium maximus Imhof, 1885 (See Zelinka, 1889:313 for details). These names have not been used for species identification in the 20th or 21st centuries)] Chaetonotus (Chaetonotus) mediterraneus Balsamo, Hummon, Todaro & Tongiorgi, 1997:87 [Subgenus assigned in this publication] Chaetonotus (Chaetonotus) napoleonicus Balsamo, Todaro & Tongiorgi, 1992:492 [Subgenus assigned by Kisielewski, 1997:148] Chaetonotus (Chaetonotus) siciliensis Hummon, Balsamo & Todaro, 1992:508 [Subgenus assigned by Kisielewski, 1997:149] Chaetonotus (Chaetonotus) similis Zelinka, 1889:317 [Subgenus assigned by Kisielewski, 1997:149. This is mostly a freshwater taxon, sometimes found in estuarine waters, e. g. Rudescu 1966] Chaetonotus (Chaetonotus) tempestivus Mock, 1979:40 [Subgenus assigned by Kisielewski, 1997:149] Chaetonotus (Chaetonotus) triacanthus Todaro, 1994:16 [Subgenus assigned by Kisielewski, 1997:149] Subgenus Hystricochaetonotus Schwank, 1990:217 [Balsamo, d’Hondt, Pierboni & Grilli, 2009 rejected the subgenus and included its species in the subgenus Chaetonotus (Chaetonotus), though we are following the established Schwank (1990) and Kisielewski (1997) systematization instead] Type-species Chaetonotus (Hystricochaetonotus) hystrix (Metschnikoff, 1865):451 [Subgenus assigned by Schwank, 1990:220. This is mostly a freshwater taxon, occasionally found in brackish-waters, e. g. Roszczak 1939] Chaetonotus (Hystricochaetonotus) lacunosus (Mock, 1979):42 [Subgenus assigned by Balsamo & Tongiorgi, 1995:6, accepted by Kisilewski 1997:150] Chaetonotus (Hystricochaetonotus) persetosus (Zelinka, 1889):337 [Subgenus assigned by Schwank, 1990:222. [This is mostly a freshwater taxon, occasionally reported from brackish-waters, e. g. Roszczak 1939] Chaetonotus (Hystricochaetonotus) variosquamatus (Mock, 1979:45) [Subgenus assigned to by Balsamo & Tongiorgi, 1995:6; subgeneric position unclear according to Kisielewski, 1997:151] Subgenus Marinochaetus Kisielewski, 1997:151 Type-species Chaetonotus (Marinochaetus) mariae (Todaro, 1992):327 [Subgenus assigned by Kisielewski, TAXONOMY OF MARINE GASTROTRICHA Zootaxa 2392 © 2010 Magnolia Press · 17 TERMS OF USE This pdf is provided by Magnolia Press for private/research use. Commercial sale or deposition in a public library or website is prohibited. 1997:151] Chaetonotus (Marinochaetus) aequispinosus (Schrom, 1972):312 [Subgenus assigned by Kisielewski, 1997:151] Chaetonotus (Marinochaetus) antipai (Rodewald, 1938):78 [Subgenus assigned by Kisielewski, 1997:151] Chaetonotus (Marinochaetus) apolemmus (Hummon, Balsamo & Todaro, 1992):504 [Subgenus assigned by Kisielewski, 1997:151] Chaetonotus (Marinochaetus) chicous (Hummon, 1974c):404 [Subgenus assigned by Kisielewski, 1997:151] Chaetonotus (Marinochaetus) oceanides (d'Hondt, 1971a):223 [Subgenus assigned by Kisielewski, 1997:151] Chaetonotus (Marinochaetus) oligohalinus (Hummon, 1974b):442 [Subgenus assigned by Kisielewski, 1997:151] (This species thus far has been found only in brackish waters) Chaetonotus (Marinochaetus) sagittarius (Evans, 1992):323 [Subgenus assigned by Kisielewski, 1997:151] Subgenus Schizochaetonotus Schwank, 1990:131 Type-species Chaetonotus (Schizochaetonotus) schultzei (Metschnikoff, 1865):451 [Subgenus assigned by Schwank, 1990:131] (This is a freshwater taxon) th 19 century synonyms. Chaetonotus maximus Gosse, 1864; Chaetonotus Schultzii Metschnikoff, 1865; Ichthydium Schultzii Ludwig, 1876 (See Zelinka, 1889:313 for details). These names have not been used for species identification in the 20th or 21st centuries)] Chaetonotus (Schizochaetonotus) atrox (Wilke, 1954) [Subgenus assigned by Schwank, 1990:131] Chaetonotus (Schizochaetonotus) dispar (Wilke, 1954) [Subgenus assigned by Schwank, 1990:131] Chaetonotus (Schizochaetonotus) hilarus (Schrom, 1972) [Subgenus assigned by Schwank, 1990:131] Chaetonotus (Schizochaetonotus) inaequidentatus Kisielewski, 1988:192 [Subgenus assigned by Balsamo, Fregni & Tongiorgi, 1995:281, though Tongiorgi, Fregni & Balsamo, 1999:588 later placed it in Euchaetonotus (which has become Chaetonotus s.s.)] Chaetonotus (Schizochaetonotus) luporinii Balsamo, Fregni & Tongiorgi, 1996:178 Chaetonotus (Schizochaetonotus) modestus (Schrom, 1972):319 [Subgenus assigned by Schwank, 1990:131] Chaetonotus (Schizochaetonotus) neptuni (Wilke, 1954):529 [Subgenus assigned by Schwank, 1990:131] Chaetonotus (Schizochaetonotus) schromi Hummon, 1974a:196 [Subgenus assigned in this publication] Syn. Chaetonotus jucundus Schrom, 1972:314 [Name preoccupied by C. jucundus d’Hondt, 1971; this name became a junior primary homonym, and was not available as a name within the genus Chaetonotus, even if the previous name-bearer was transferred to another genus; thus the name was replaced by Hummon, 1974a:196 in accordance with ICZN 23.3.5 and 60.1] Chaetonotus (Schizochaetonotus) serenus (Schrom, 1972:317) [Subgenus assigned by Schwank, 1990:131] Chaetonotus (Schizochaetonotus) woodi (Thane-Fenchel, 1970:128) [Subgenus assigned by Schwank, 1990:131] INCERTAE SEDIS Chaetonotus Ehrenberg, 1830:64 Chaetonotus balticus Remane, 1926a:247 [Subgeneric position unclear according to Kisielewski, 1997:151] Chaetonotus marinus Giard, 1904a:1062 [Not treated by Kisielewski, 1997, but subgeneric position unclear, as treated in this publication] Syn. Chaetonotus pleuracanthus Remane, 1926a:243 [d’Hondt, 1971:177, but considered unverified by Mock 1979:52 and rejected in this publication, see below] Chaetonotus parthenopeius Wilke, 1954:532 [Subgeneric position unclear according to Kisielewski, 1997:151] Genus Aspidiophorus (Voigt, 1903):90 [Not Voigt, 1904:128, as was noted by Weiss, 2001:312] Syn. Aspidonotus Voigt, 1902:676 [Voigt, 1903:90] Type species Aspidiophorus paradoxus (Voigt, 1902):676 [This is strictly a freshwater taxon] Syn. Aspidonotus paradoxus Voigt, 1902:676 [Voigt, 1903:90] 18 · Zootaxa 2392 © 2010 Magnolia Press HUMMON & TODARO TERMS OF USE This pdf is provided by Magnolia Press for private/research use. Commercial sale or deposition in a public library or website is prohibited. Aspidiophorus bisquamosus Mock, 1979:74 Aspidiophorus lamellophorus Balsamo, Hummon, Todaro & Tongiorgi, 1997:83 Aspidiophorus marinus Remane, 1926a:247 Aspidiophorus mediterraneus Remane, 1927d:44 sensu Hummon, 1974d:437 Aspidiophorus multitubulatus Hummon, 1974d:441 Aspidiophorus oculatus Todaro, Dal Zotto, Maiorova & Adrianov, 2009:298 Aspidiophorus paramediterraneus Hummon, 1974b:439 Aspidiophorus polystictos Balsamo & Todaro, 1987:147 Aspidiophorus tentaculatus Wilke, 1954:535 Genus Caudichthydium Schwank, 1990:61 Type-species Caudichthydium hummoni (Ruppert, 1977):2 Syn. Ichthydium hummoni Ruppert, 1977:2 [Schwank, 1990:61] Caudichthydium rupperti (Mock, 1979):62 Syn. Ichthydium rupperti Mock, 1979:62 [Schwank, 1990:61] Caudichthydium supralitorale (Mock, 1979):64 Syn. Ichthydium supralitorale Mock, 1979:64 [Schwank, 1990:61] Genus Halichaetonotus Remane, 1936:190 [Raised from subgenus to genus level by Schrom, 1972:330] Type-species Halichaetonotus pleuracanthus (Remane, 1926a):243 Syn. Chaetonotus pleuracanthus Remane, 1926a:243 [Schrom, 1972:326. D’Hondt (1968a) reported finding Chaetonotus marinus, a species of the “Maximus” group previously described by Giard, 1904, in the sand “a diatomées d’Ambleteuse”. He later (d’Hondt 1971) considered that specimens of Chaetonotus (now Halichaetonotus) pleuracanthus must correspond to C. marinus because this was the only species of that he found in the type-locality; this interpretation is now reinforced by d’Hondt & Balsamo (2009); we cannot accept this synonymy by virtue of the clear difference that exists between the dorsal cuticular covering of the two species that d’Hondt considers to be synonymous: spinated scales in C. marinus vs keeled scales in H. pleuracanthus (e.g., see Fig. 1 of Giard, 1904 vs. Fig. 1 of Remane, 1926)] Halichaetonotus aculifer (Gerlach, 1953):208 Syn. Chaetonotus aculifer Gerlach, 1953:208 [Schrom, 1972:326] Halichaetonotus aculifer forma adriatica Schrom 1972:331 [By treating the trinomial as a form after 1960, this name is rendered an infrasubspecific category, a rank which is not recognized by the ICZN 45.6.3] Halichaetonotus arenarius (d’Hondt, 1971a):218 Syn. Chaetonotus arenarius d’Hondt, 1971a:218 [In this publication] Halichaetonotus atlanticus Kisielewski, 1988:200 Halichaetonotus australis Nicholas & Todaro, 2005:974 Halichaetonotus balticus Kisielewski, 1975:20 Halichaetonotus bataceus Evans, 1992:325 Halichaetonotus batillifer (Luporini & Tongiorgi, 1972):302 Syn. Chaetonotus batillifer Luporini & Tongiorgi, 1972:302 [Mock, 1979:51] Halichaetonotus clavicornis Balsamo, Fregni & Tongiorgi, 1995:281 Halichaetonotus decipiens (Remane, 1929):167 Syn. Chaetonotus (Halichaetonotus) decipiens [Subgenus raised to genus rank by Schrom, 1972:330; the species epithet remained decipiens, since a junior secondary homonym, dubius/m, that is replaced before 1961 is permanently invalid ICZN 59.3; see Hummon, Balsamo & Todaro, 1992:514 for discussion] Syn. Chaetonotus decipiens Remane, 1929:167 [Assigned to the subgenus Halichaetonotus by Remane, 1936:190] TAXONOMY OF MARINE GASTROTRICHA Zootaxa 2392 © 2010 Magnolia Press · 19 TERMS OF USE This pdf is provided by Magnolia Press for private/research use. Commercial sale or deposition in a public library or website is prohibited. Syn. Heterolepidoderma dubium Remane, 1926a:250 [Remane, 1929:167; transfer of the species to Chaetonotus would have created a junior homonym, for the name C. dubius Daday, 1905:77 was already occupied, hence Remane renamed the species C. decipiens] Halichaetonotus etrolomus Hummon, Balsamo & Todaro, 1992:510 Halichaetonotus euromarinus Hummon & Todaro n. sp. [In this publication] Syn. Halichaetonotus spinosus Mock, 1979:54 [Name not available, because it was previously used as an infrasubspecific trinomial, and hence cannot be used as a specific epithet, ICZN 45.6.3] Syn. Chaetonotus decipiens forma spinosus d’Hondt 1971a:220 [By treating the trinomial as a form after 1960, this name is rendered an infrasubspecific category, and hence is not available to be named a species (ICZN 45.6.3)] Halichaetonotus genatus Balsamo, Fregni & Tongiorgi, 1995:282 Halichaetonotus italicus Balsamo, Hummon, Todaro & Tongiorgi, 1997:87 Halichaetonotus jucundus (d'Hondt, 1971a):219 Syn. nov. Chaetonotus jucundus d'Hondt, 1971:219 [Balsamo, Fregni & Tongiorgi, 1995:284 refer to a species of d’Hondt (1971) and of Mock (1979) as H. jucundus, but without indicating a synonymy between C. jucundus d’Hondt and H. jucundus (d’Hondt); we do so formally in this publication] Halichaetonotus lamellatus Kisielewski, 1975:16 Halichaetonotus littoralis (d’Hondt, 1971a):216 Syn. Chaetonotus littoralis d’Hondt, 1971a:216 [Mock, 1979:51] Halichaetonotus margaretae Hummon, Balsamo & Todaro, 1992:511 Halichaetonotus marivagus Balsamo, Todaro & Tongiorgi, 1992:494 Halichaetonotus paradoxus (Remane, 1927d):44 Syn. Chaetonotus paradoxus Remane, 1927d:44 [Schrom, 1972:330] Halichaetonotus parvus (Wilke, 1954):531 Syn. Chaetonotus parvus Wilke, 1954:531 [Schrom, 1972:330] Halichaetonotus polonensis Hummon, 2008a:268 [Change of gender ending from H. polonense to H. polonensis, in this publication (ICZN 34.2)] Halichaetonotus riedli Schrom, 1972:327 Halichaetonotus schromi Kisielewski, 1975:18 Halichaetonotus somniculosus (Mock, 1979):47 [Genus transfer in this publication, based on an observation by M. A. Todaro of this species in Sweden (Willems, Curini-Galletti, Ferrero, Fontaneto, Heiner, Huys, Ivanenko, Kristensen, Kånneby, MacNaughton, Martínez Arbizu, Todaro, Sterrer & Jondelius, 2009:20, as Chaetonotus somniculosus) and also on a video by Hummon from Salisbury, Massachusetts (Aug 2000, also listed in Hummon (2009b) as Chaetonotus somniculosus).] Syn. Chaetonotus somniculosus Mock, 1979:47 [Subgeneric position within Chaetonotus unclear according to Kisielewski, 1997:151] Halichaetonotus swedmarki Schrom, 1972:328 Halichaetonotus tentaculatus (d’Hondt, 1971a):222 Syn. Chaetonotus tentaculatus d’Hondt, 1971a:222 [In this publication] Halichaetonotus testiculophorus (Hummon, 1966):451 Syn. Chaetonotus testiculophorus Hummon, 1966:451 [Hummon, 2007a:268; previously, Kisielewski 1997:151 had assigned the species to the subgenus Marinochaetus) Halichaetonotus thalassopais Hummon, Balsamo & Todaro, 1992:513 Genus Heterolepidoderma Remane, 1927b:287 [Note, the name Heterolepidoderma appeared first in Remane, 1926a:248, however the name with this date should be considered nomen nudum as at that time the diagnosis of the genus was not provided; likewise nomem nudum should be considered for Heterolepidoderma in Remane 1926b:287] Type-species Heterolepidoderma marinum Remane, 1926a:248 [Note the attribution of the type species for this genus is debated, some consider the type species to be H. ocellatum (Metschnikoff, 1864) 20 · Zootaxa 2392 © 2010 Magnolia Press HUMMON & TODARO TERMS OF USE This pdf is provided by Magnolia Press for private/research use. Commercial sale or deposition in a public library or website is prohibited. crediting as designator Remane 1936 (see e.g . Balsamo et al. 2009:14); we disagree for the following reasons: in 1927, in a paper on freshwater gastrotrichs, Remane (1927a:287) differentiated species that should be retained in the genus Lepidoderma from those which should be separated into a new genus, Heterolepidoderma n. g. Within the latter group he listed, in sequence, four species, naming two of them as new: “majus n. sp., gracile n. sp.”, while transferring one: “ocellatum Metschnikoff” from the genus Lepidoderma, and included a species that he had described in this genus the year before: “marinum Remane”. Later in the article (pp. 313–317) Remane cited H. ocellatum again after describing two new freshwater species in the genus: H. majus and H. gracile. It is our contention that because the date was before 1931, as soon as the genus Heterolepidoderma Remane, 1927 was validly published as an “available” name (ICZN 10), the first species in the first publication that used the generic name, i. e. Heterolepidoderma marinum Remane, 1926, automatically became its type-species by “indication”, even without being so designated (ICZN p. 107, as was noted by Weiss, 2001:312), and that this has precedence to new species that were described or a species that was transferred to that genus from another at the time the genus was named. With respect to Remane, 1936:193, it is debatable whether he had anything more than a chronological listing of species; certainly he nowhere in the article refers to the typespecies concept] Heterolepidoderma arenosum Kisielewski, 1988:204 Heterolepidoderma armatum Schrom, 1966a:41 Heterolepidoderma axi Mock, 1979:69 Heterolepidoderma caudosquamatum Grilli, Kristensen & Balsamo, 2009:49 Heterolepidoderma clipeatum Schrom, 1972:340 Heterolepidoderma contectum Schrom, 1972:342 Heterolepidoderma foliatum Renaud-Mornant, 1967b:161 Heterolepidoderma grandiculum Mock, 1979:66 Heterolepidoderma hermaphroditum Wilke, 1954:533 Heterolepidoderma istrianum Schrom, 1972:333 Heterolepidoderma loricatum Schrom, 1972:337 Heterolepidoderma ocellatum (Metschnikoff, 1865):451 sensu Kisielewski, 1981:65 [This is mostly a freshwater taxon, sometimes found in estuarine water, e. g. Rudescu 1966] th 19 century synonyms. Ichthydium ocellatum Metschnikoff, 1865 (see Zelinka, 1889:296 for details). This name has not been used for species identification in the 20th or 21st centuries] Syn. Lepidoderma ocellatum Zelinka, 1889:307 [Remane, 1927b:316] Genus Ichthydium Ehrenberg, 1830:64 Type-subgenus Ichthydium Ehrenberg, 1830:64 (ICZN 44.1) sensu stricto Syn. Euichthydium Schwank, 1990:94 [Kånneby, Todaro & Jondelius, 2009:30 first pointed out that the subgenus Euichthydium is invalid; because this subgenus contains the type species of the genus, it must bear the same name as the genus, see ICZN Article 44.1] Type-species Ichthydium podura (Müller, 1773):66 [Ehrenberg, 1830:64 This is mostly a freshwater taxon, sometimes found in marine water, e. g. Hummon 2009b {Figure, E Med Database}] 19th century synonyms. Cercaria podura Müller, 1773:66; Furcocerca podura Lamarck, 1815; Enchelys podura Nitsch, 1817; Diurella podura Hemprich & Ehrenberg, 1831 (See Ehrenberg, 1838:390 for details); Chaetonotus podura Stokes, 1887(See Zelinka, 1889:296 for details. None of these names have been used for species identification in the 20th or 21st centuries)] Ichthydium (Ichthydium) cyclocephalum (Grünspan, 1910):208 [Subgenus assigned to Euichthydium by Schwank, 1990:94; reassigned to Ichthydium s.s. in this publication] Ichthydium (Ichthydium) tergestinum (Grünspan, 1908):233 [Subgenus assigned to Euichthydium by Schwank, 1990:94; reassigned to Ichthydium s.s. in this publication] TAXONOMY OF MARINE GASTROTRICHA Zootaxa 2392 © 2010 Magnolia Press · 21 TERMS OF USE This pdf is provided by Magnolia Press for private/research use. Commercial sale or deposition in a public library or website is prohibited. Genus Lepidodermella Blake, 1933:606 Syn. Lepidoderma Zelinka, 1889:300 [Preoccupied; name replaced by Blake, 1933:606] Type-species Lepidodermella squamata (Dujardin, 1841):569 (This is mostly a freshwater taxon, sometimes found in brackish-water, e. g. Roszczak 1939) Syn. Lepidoderma squamatum (Dujardin, 1841):569 [Blake, 1933:606] Syn. Chaetonotus squammatus Dujardin, 1841:569 [Zelinka, 1889:300. Dujardin used the species name Chaetonotus squammatus in the text, and Chaetonotus squamatus in the figure legend (though not in the figure per se). As first reviser, Weiss, 1988:369 clarified the issue of ‘mm’ versus ‘m’ (ICZN 24.2.2) and the gender ending, variously spelled in the literature (ICZN 30)] 19th century synonyms. Chaetonotus squamosus Schultze, 1853; Chaetonotus squamatus Pritchard, 1864; Chaetonotus tesselatus Metschnikoff, 1864; Ichthydium maximum Ludwig, 1875; Chaetonotus loricatus Stokes, 1887 (See Zelinka, 1889:300 for details; none of these names have been used for species identification in the 20th or 21st centuries) Lepidodermella limogena Schrom, 1972:345 [Change of gender ending from L. limogenum to L. limogena, was made in Weiss, 1988:375 (ICZN 34.2)] Family Muselliferidae Leasi & Todaro, 2008:396—Includes 2 genera and 5 species Type-Genus Musellifer Hummon, 1969:282 Type-species Musellifer sublitoralis Hummon, 1969:283 Musellifer delamarei (Renaud-Mornant, 1968):142 Syn. Polymerurus delamarei Renaud-Mornant, 1968:142 [Hummon, Balsamo & Todaro, 1992:515] Musellifer profundus Vivier, 1974:183 sensu Leasi & Todaro, 2009:3 Genus Diuronotus Todaro, Balsamo & Kristensen, 2005:1391 Type-species Diuronotus aspetos Todaro, Balsamo & Kristensen, 2005:1392 Diuronotus rupperti Todaro, Balsamo & Kristensen, 2005:1395 Family Xenotrichulidae Remane, 1927d:294—Includes 3 genera and 24 species Subfamily Draculiciterinae Ruppert, 1979:12 Type-Genus Draculiciteria Hummon, 1974b:203 Syn. Xenotrichuloides d’Hondt, 1971:148 [It should be considered nomen nudum as stated by Hummon, 1974b:203] Type-species Draculiciteria tesselata (Renaud Mornant, 1968):142 Syn. Polymerurus tesselatus Renaud Mornant, 1968:142 [Hummon, 1974a:203] Syn. Xenotrichula (Xenotrichuloides) mirabilis d’Hondt, 1967:207, 1971:181 [It should be considered nomen nudum as stated by Hummon, 1974a:204] Subfamily Xenotrichulinae Remane, 1927d: ICZN 36 [Ruppert, 1979:12] Type-Genus Xenotrichula Remane, 1927a:289 Type-subgenus Velox Remane, 1927c:289 (ICZN 44.1) sensu stricto— new subgenus [In his monography of Xenotrichulidae Ruppert (1979:19) divided the genus Xenotrichula into two groups: Xenotrichula intermedia-species group and Xenotrichula velox-species group. In our opinion the morphological characteristics of the species involved support the subdivision of the Xenotrichula species into two distinct taxonomic ranks proposing the establishment of two subgenera Xenotrichula (Velox) and Xenotrichula (Xenotrichula) the diagnoses of which are the same as in Ruppert 1979. The rationale 22 · Zootaxa 2392 © 2010 Magnolia Press HUMMON & TODARO TERMS OF USE This pdf is provided by Magnolia Press for private/research use. Commercial sale or deposition in a public library or website is prohibited. for the establishment of the two subgenera is two-fold: first to eliminate taxonomic confusion deriving from the current subdivision into species groups that have no taxonomic standing and second to stress the kin relationship of members of the two lineages] Type-species Xenotrichula (Velox) velox Remane, 1927a:289 Xenotrichula (Velox) cornuta Wilke, 1954:520 Xenotrichula (Velox) guadelupensis Kisielewski, 1984:37 [Assigned in this publication; change of gender ending from X. guadelupense to X. guadelupensis, in this publication (ICZN 34.2)] Xenotrichula (Velox) tentaculata Rao & Ganapati, 1968:51 Subgenus Xenotrichula —new subgenus [In this publication. In his monograph of Xenotrichulidae Ruppert (1979:19) divided the genus Xenotrichula into two groups: Xenotrichula intermedia-species group and Xenotrichula velox-species group. In our opinion the morphological characteristics of the species involved support the subdivision of the Xenotrichula species into two distinct taxonomic ranks proposing the establishment of two subgenera Xenotrichula (Velox) and Xenotrichula (Xenotrichula) the diagnosis of which are the same as in Ruppert 1979. The rationale for the establishment of the two subgenera is twofold: first to eliminate taxonomic confusion deriving from the current subdivision into species groups that have no taxonomic standing and second to stress the kin relationship of members of the two lineages] Type-species Xenotrichula (Xenotrichula) intermedia Remane, 1934:477 Syn. Xenotrichula beauchampi Lévi, 1950:39 [Ruppert, 1979:20. D’Hondt & Balsamo (2009) consider X. beauchampi to be a good species, by virtue of differences regarding the insertion along the ventral trunk region of the two groups of locomotor cirri, the second of which in the orginal description of X. intermedia seems to have been omitted. This in our opinion was an error of omission made by Remane, since the belly tufts are characteristic of all species in this subfamily. Moreover, all specimens found world-wide and affiliated with this taxon match the description of X. beauchampi (cf. Todaro et al., 1996), including the specimens found by one of us in the Baltic Sea, not too far from Kiel, the type locality (cf. Hummon, 2008). Ironically, Remane (1936: 183) in discussing the characteristics of the Xenotrichulidae indicated “….an additional isolated tuft of cilia [group of cirri] is present in the middle of the gut region in X. subt., X. aff. and X. pygm.” but said nothing about the presence of these locomotor structures in X. intermedia or X. velox. Responding to d’Hondt & Balsamo 2009, we note that Ruppert (1979: 20) indicates both presence and location of the rear tuft of cirri when he writes “.intermedia-type stalked scales ventral to the trunk in 9 longitudinal rows on each side of the ventral midline; these scales extend across the ventral midline in the region of the posterior group of cirri”] Syn. Xenotrichula beauchampi var. angusta d'Hondt 1966b:6 [In treating the trinomial as a variety after 1960, this name is rendered an infrasubspecific category, a rank which is not recognized by the ICZN 45.6.3. By contrast with d’Hondt & Balsamo (2009), we reconfirm the synonymy considering the metric variability within the range of the species] Syn. Xenotrichula beauchampi var. maritima d'Hondt 1970:10 [In treating the trinomial as a variety after 1960, this name is rendered an infrasubspecific category, a rank which is not recognized by the ICZN 45.6.3. By contrast with d’Hondt & Balsamo (2009), we reconfirm the synonymy considering the metric variability within the range of the species] Syn: Xenotrichula carolinensis ssp. syltensis Mock, 1979:21 [Synonymized with X. intermedia in this publication because the morphological characters fit within the scope of the species as reported from around the world] Syn: Xenotrichula punctata of Swedmark 1956b:86 [Ruppert, 1979:20] Xenotrichula (Xenotrichula) bispina Roszczak, 1939:10 Xenotrichula (Xenotrichula) carolinensis Ruppert, 1979:22 [Synonymized with X. intermedia by Balsamo, Todaro & Tongiorgi, 1992:496; synonymy rejected in this publication, X carolinensis having a TAXONOMY OF MARINE GASTROTRICHA Zootaxa 2392 © 2010 Magnolia Press · 23 TERMS OF USE This pdf is provided by Magnolia Press for private/research use. Commercial sale or deposition in a public library or website is prohibited. smaller size than X. intermedia and being thus far restricted to the North American coastline from Beaufort, North Carolina to St. Petersburg, Florida (Hummon 2009 {N.America database})] Xenotrichula (Xenotrichula) floridana Thane-Fenchel, 1970:126 [Change of gender ending from X. floridanus to X. floridana, in this publication (ICZN 34.2)] Xenotrichula (Xenotrichula) laccadivensis Rao, 1991:76 Xenotrichula (Xenotrichula) lineata Schrom, 1972:307 Xenotrichula (Xenotrichula) micracantha (Remane, 1926a):246 Syn. Chaetonotus micracanthus Remane, 1926b:246 [Remane, 1929:178] Xenotrichula (Xenotrichula) paralineata Hummon & Todaro, 2007:298 [Assigned in this publication] Xenotrichula (Xenotrichula) punctata Wilke, 1954:519 Xenotrichula (Xenotrichula) quadritubulata Kisielewski, 1988:189 [Assigned in this publication] Xenotrichula (Xenotrichula) soikai Schrom, 1966:37 Xenotrichula gymnocephala Remane, 1940:xx [It should be considered nomen nudum as stated by d’Hondt, 1971:181] Genus Heteroxenotrichula Wilke, 1954:522 Type-species Heteroxenotrichula squamosa Wilke, 1954:522 Heteroxenotrichula affinis (Remane, 1934):477 Syn. Xenotrichula affinis Remane, 1934:477 [Ruppert, 1979:13] Syn. Xenotrichula variocirrata d'Hondt, 1966b:7 [W.D. Hummon in Jouk, Hummon, Hummon & Roidou, 1992:6] Heteroxenotrichula arcassonensis Ruppert, 1979:41 Heteroxenotrichula pygmaea (Remane, 1934):477 Syn. Xenotrichula pygmaea Remane, 1934:477 [Ruppert, 1979:13] Syn. Xenotrichula flandrensis d'Hondt, 1968a:221 [Ruppert, 1979:45. D’Hondt & Balsamo (2009:267) consider the new combination of Ruppert to be incorrect, supporting d’Hondt’s (1968) original belief that this taxon is a distinct species, based on the number and disposition of cephalic bristles (soies céphaliques, figured) and perhaps the relative length of the adhesive tubes with respect to the furcal branches (not figured). We disagree and support Ruppert’s synonymization, for the following reasons. The original description was based on a single individual; the author himself had some doubt in describing it as a genuine species (d’Hondt, 1968a: 221). Specimens with head morphology matching the original description of X. flandrensis have never been found again. In describing the specimen d’Hondt (1968) noticed two illuminating details: the specimen had eggs inside (but not sperm) and its scale covering was very difficult to see. These “robust” traits are shared with Heteroxenotrichula pygmaea, which reproduces by parthenogenenesis (the only Xenotrichulinae to do so) and its scales are very weakly developed. The apparent number of cephalic ciliary tufts in d’Hondt’s specimen and those of H. pygmaea can be explained considering that in xenotriculids the cephalic ciliary tufts tend to splay apart when specimens are compressed between the slide and the coverslip, a practice that researchers tend to do when details, such as scale’s shape, are difficult to see. d’Hondt’s specimen must have been compressed because the author was unable to appreciate the details of the ventral locomotor cirri, whose shape becomes disrupted with compression. As far as the length of adhesive tube is concerned, d’Hondt (1968) indicated that the tube was 1/3 of the total furca, which is also the ratio in Ruppert’s (1979) photos and drawings of H. pygmaea. The type locality of X. flandrensis has never been sampled again] Syn. Xenotrichula sp. A of Schmidt, 1974:61 [Ruppert, 1979:45] Heteroxenotrichula simplex (Mock, 1979):23 Syn. Xenotrichula simplex Mock, 1979:23 [Hummon & Todaro, 2007:302] Heteroxenotrichula subterranea (Remane, 1934):475 Syn. Xenotrichula subterranea Remane, 1934:475 [Ruppert, 1979:13] Syn. Xenotrichula sp. of Luporini, Magagnini & Tongiorgi 1971:449 [Ruppert, 1979:31] 24 · Zootaxa 2392 © 2010 Magnolia Press HUMMON & TODARO TERMS OF USE This pdf is provided by Magnolia Press for private/research use. Commercial sale or deposition in a public library or website is prohibited. Heteroxenotrichula texana Todaro, 1994:18 Heteroxenotrichula transatlantica Ruppert, 1979:33 Heteroxenotrichula variocirrata (d’Hondt, 1966) sensu d’Hondt & Balsamo, 2009:265 Syn. Xenotrichula variocirrata d'Hondt, 1966b:7 [D’Hondt & Balsamo, 2009:265. The new morphologic information on this species recently provided by d’Hondt & Balsamo (2009) no longer justifies the synonymy of this taxon with H. affinis as proposed by Jouk et al. (1992), but in defense of Jouk et al., the original description indicated that dorsal scales were pedunculated, whereas they are now considered to be foliacious. Considering that the new information orginates from microphotographs aready available at the time of the orginal description (d’Hondt, 1966) and not from freshly collected specimens, in our opinion the discrepancies between the original description and the new data, expecially the one regarding the presence in this species of large scales (flat?) on the posterior region of the trunk need to be confirmed; consequently, we consider provisionally the taxon to be species inquirenda] Heteroxenotrichula wilkeae Ruppert, 1979:41 [Change of gender ending from H. wilkei to H. wilkeae was noted by d’Hondt & Balsamo, 2009. [Synonymyzing H. variocirrata with H. wilkeae as stated by d’Hondt & Balsamo (2009) is not substainable by virtue of differences between the two species regarding the metrics of key traits, e.g. total body length, pharynx length and furca length averaging respectively 186 µm, 41.3 µm and 54 µm in H. wilkeae (n = 19–20) vs 140–150 µm, 55–64 µm and 57–75 in H. variocirrata (see original descriptions)] Acknowledgments This research was supported by a MIUR grant to MAT (PRIN-2007 – Approccio integrato all’identificazione dei Gastrotrichi marini). We cannot sufficiently express our obligation to Mitchell Weiss and Maria Balsamo for their exhaustive reviews of this manuscript. We have accepted most, but not all, of their comments; for those on which we differ, we alone are responsible. We also thank an anonymous reviewer for many valuable suggestions. Bibliography Balsamo, M., Fregni, E. & Tongiorgi, P. 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