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<Echiura:Echiuridae) JapaneseJapaneseSociety Society ofSystematicZoologyof Systematic Zoology Species Diversity, 2004, 9, 109-123 Synonymy of the West-Pacific Echiuran Listriolobus sorbillans <Echiura: Echiuridae), with Taxonomic Notes Towards a Generic Revision Teruaki Nishikawa The A]dgqya Uitiversity Museum, Chikusa-ku, IVtlgQya, 464-860I Jtipan E-mail: nishikawa@'num.nagaya-u.ac,.tp (Received 10 June 2oo3; Accepted 4 December 2003) Examination of the syntypes of Listriolobus sorbillans (Lampert, 1883) from the Philippines and new specimens from Moreton Bay, eastern Aus- tralia, and the Ryukyu Islands, Japan, together with a literature survey, re- veals that it is a senior synonym of L. billitonensis Fischer, 1926 recorded flrom thc Java Sea and L. riuiciuensts Sato, 1939 from southwestern Japan. Observations on individual variation in the ratio of proboscis to trunk length, the number of longitudina] musele bands, and the number and arrangement of gonoducts are presented. A tabular comparison of all the known species of the genus Listriotobus is given, together with some new taxonomic information about the congeners and allies. Key Words: Listriolobus, Echiura, taxonomy, synonymy, revision, Introduction The echiuran genus Listriotobus Spengel, 1912, belonging to the family Echi- uridae (for family-group nomenclature see Nishikawa 1998), is distinguished from the closely related Ochetostonea Leuckart and RUppel, 1828 by the inner-most oblique muscle layer of the trunk wall, which is continuous where it is visible be- tween the longitudinal muscle bands in the former genus but separated into fasci- cles in the latter (see, e.g., Spenge] 1912; Fisher 1946, 1949; Stephen and Edmonds 1972; Edmonds 1987; Biseswar and Moodley 1989; Nishikawa 1992; fbr historic ob- jections to the separate status of these genera see Bock 1942). The eriginal deseriptions of many species referred to these two genera [includ- ing Listriolobus sorbiltans (Lampert, 1883) in the present study] lack information about the above-stated diagnostic feature of the oblique muscle layer, and, there- fbre, the generic affiliation of such species needs a critical reexamination. For ex- ample, Biseswar and Moodley's (1989) reexamination of the holotype of Oche- tostoma capense Jones and Stephen, 1955 resulted in the transfer of this species to Listriolobus. During my preparation for this critical study, I had the opportunity to inspect the syntypes of Thatassema sorbtltans Lampert, 1883 from the Philippines and dis- covered its conspecificity with Listriolobus riukiuensis Sato, 1939, so far recorded from the Ryukyu Islands of southwestern Japan (Sato 1939; Nishikawa 1992). The first redescription of the syntypes of T, sorbillans since its establishment is given herein to confirm its affiliation to the genus Listriolobus. Furthermore, some taxo- NII-Electronic Library Service JapaneseJapaneseSociety Society ofSystematicof Systematic Zoology 11O Teruaki Nishikawa nomic, nomenclatural, and biogeographical information is presented on the basis of newly collected specimens from Japan and Australia, as well as a literature sur- vey, to provide a fbundation fbr a thorough taxonomic revision of the genus Listri- olobus and allies in the future, Historical Background Establishment of the genus Listriolobus Spengel (1912) pointed out that the anterior-most pair of nephridia (=gonod- ucts in the present study) is always situated anterior to the ventral setae, and the "Interbasa]muskel" (-interbasal muscle) is always absent, in such specimens of the genus Thalassema that have the longitudinal muscle layer of the trunk sepa- rated into bundles and also have three or more pairs of gonoducts. He thereupon proposed a taxonomic division of such specimens into two groups on the basis of `tThalassema his actual reexamination of the echiurans then described as erlythro- grammon", including its holotype housed at the Forschungsinstitut Senckenberg (stil1 there: D. Fiege, pers. comm.). "Septalleisten" One group was composed of the specimens with (derived from the fascicules of the oblique muscle layer between the longitudinal muscle bands) and included some of the syntypes of Thalassema caudex Lampert, 1883 (those col- lecteci from the Red Sea, while those from the Indian Ocean were then unexam- ined), The genus name Ochetostoma Leuckart and RUppell, 1828 was revived by Spengel (1912) as valid for this group, its type species being Ochetostoma erlythro- grammon Leuckart and RUppell, 1828. The other group included the specimens of `"71 e)lythrQgrammon" in the sense of Sluiter (1884) and Wilson (1900), as well as "Septalleisten". Spengel's (1912) own specimens, all lacking the For these speci- mens a new genus Listriolobus was erected by Spengel (1912) without type designa- tion and any nominal species included therein (see below). Later, Fischer (1926) es- tablished Listriolobus billitonensis Fischer, 1926 for Sluiter's (1884) many speci- mens firom coastal sand of Billiton Island, Indonesia, and L. bahamensts Fischer, 1926 for Wilson's (1900) one specimen from off Great Abaco Island, Bahamas, and also for Spengel's (1912) single specimen from Florida, These two species are obvi- ously available, since they were published before 1931 and accompanied by biblio- graphic references (ICZN 4th ed., Art, 12); the whereabouts of the name-bearing types of these two species are now unknown. Confused definition of genus Listriolobus As shown above, it is clear that Spengel (1912) recognized the genus Listri- otobus as lacking the interbasal muscle, as rightly pointed out by Bock (1942). How- ever, Fisher (1946: 234) reversed the definition of this genus in terms of the inter- "interbasal basal muscle, stating muscle of setae [is] present", probably based on the occurrence ef the muscle in L. pelodes Fisher, 1946, This misunderstanding on the part of Fisher has been fbllowed by most subsequent authors (e.g., Stephen and Edmonds 1972; Edmonds 1987; Biseswar and Moodley 1989). Fisher (1946) trans- ferred 71halassenza sorbillans Lampert, 1883 to the genus Listriotobus. Thus, L. sor- bittans has been in¢ orrectly believed to have the interbasal muscle, while L. riuki- uensis lacks it, as is clearly shown by its original description, This represents the NII-Electronic Library Service JapaneseJapaneseSociety Society ofSystematicof Systematic Zoology Synonymy ofaWest-Pacific echiuran 111 "L. reason why claimed, not Fisher (1946:234) riukiuensis .,, isprobably a Listri- olobus.'i Soon afterwards, Fisher (1949) rightly revised his previous definition of the genus Listrioiobus, explicitly stating that the interbasal muscle is present or ab- sent in species of the genus; this definition is revived in the present study, So far as the prevailing echiuran classification is concerned, it seems unnecessary to divide this genus into two according to the presence or absence ef the muscle. For exam- ple, the genera Thalassema (s. str.) and Ochetostoma both include species with and without the interbasal muscle (see Stephen and Edmonds 1972; Biseswar 1988). Taxonomy Genus Listriolobus Spengel, 1912 Listriolobus Spengel, 1912: 316 <type species: L. bahamensis Fischer, 1926, by Fisher's (1946) subsequent designation); Fisher 1949: 482, Diagnosis Eemended by Fisher (1949) and in the present study]. Longitudinal muscles of body wall grouped into bands (at least in adults); inner-most oblique muscles continuous, never separated into fascicles between longitudinal muscle bands; one to three pairs (often with some anomalies) of gonoducts with their gonostomes located proximally; gonostomal lips elongated and often coiled spi- rally; interbasal muscle present or absent. Remarks. Fisher (1946, 1949) and Stephen and Edmonds (1972) regarded the original description of the genus Listriolobus as a nomen nudum, and Fisher attrib- uted its authorship and date te Fischer, 1926. However, this seems incorrect ac- cording to the ICZN 4th ed. (International Commission on Zoological Nomencla- ture 2000). Spengel (1912) gave a definition of the genus; though he failed to make a type designation and did not assign any nominal species to the new genus, this does not make the genus a nomen nudum, since it was proposed before 1931 (ICZN 4th ed., Art. 13.3), Thus, this genus takes the authorship and date of Spengel, 1912, as was rightly designated by Stephen and Edmonds (1972) and fo11owed by subse- quent authors. Since Fischer (1926) is the first to include expressly the nominal species (Listri- olobus biltitonensis Fischer, 1926 and L. bahamensis Fischer, 1926) in the genus "originally Listriotobus Spengel, 1912, these two species are deemed to be the in- cluded nominal species" eligible fbr type fixation (ICZN 4th ed., Art. 67.2,2). The type species is L. bahamensis Fischer, 1926 by Fisher's (1946) subsequent designa- tion (ICZN 4th ed,, Art. 69,1), Listriolobus sorbillans (Lampert, 1883) [Japanese name: Tatejima-yumushi] (Figs 1-2, Tables 1-2) Thatassenza sorbillans Lampert, 1883: 340-341; Shipley 1899: 352 (no new record); Augener 1903: 349; Wharton 1913: 243-247, text-fig, 1. NII-Electronic Library Service JapaneseJapaneseSociety Society ofSystematicof Systematic Zoology 112 TeruakiNishikawa Listriolobus sorbillans: Fisher 1946: 234; Stephen and Edmonds 1973: 425-426 (no new record). Listriolobus billitonensis Fischer, 1926: 110 (by bibliographical reference to Tha- lassema erlythrogrammon of Sluiter, 1884: 58-79, pl. 3) . Syn. nov. Listriolobus riukiuensis Sato, 1939: 359-362, text-figs 10-13; Stephen and Edmonds 1973: 425 (no new record), Syn. nov.
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