Bull. Natl. Mus. Nat. Sci., Ser. A, 38(3), pp. 95–124, August 22, 2012 On Opening a Box of Worms (Oligochaeta, Megascolecidae) — Historical Earthworm Specimens Transferred to Tokyo from the Saito Ho-on Kai Museum of Natural History in Sendai Robert J. Blakemore National Institute of Biological Resources, Incheon 404–170, Korea E-mail: [email protected] (Received 18 May 2012; accepted 23 July 2012) Abstract A box discovered at the National Museum of Nature and Science, Tokyo held histori- cal specimens from Saito Ho-on Kai Museum dating from 1920–1930s coincident with author Dr. Shinkishi Hataiʼs tenure and likely identified by him or his assistant, Mr. Shinryo Ohfuchi. A syn- type is newly recognized for Amynthas maculosus (Hatai, 1930) comb. nov., no others could be unequivocally proven so. However, several are candidates for neotypification under current ICZN (1999) rules as a first step to resolving zoological complexities of parthenogenesis and of taxo- nomic confusion persisting for >112 years. Neotypes are explicitly designated herein for Amynthas vittatus (Goto and Hatai, 1898), Duplodicodrilus acinctus (Goto and Hatai, 1899) new combina- tion (synonym yezoensis Kobayashi, 1938), and for topotypes of Amynthas? yunoshimensis (Hatai, 1930) itself a probable synonym of Metaphire hilgendorfi (Michaelsen, 1892), and of M. yamadai (Hatai, 1930) that is retained separately from M. soulensis (Kobayashi, 1938). Metaphire soulensis synonyms (shinkeiensis Kobayashi, 1938 and aokii Ishizuka, 1999) are re-stated. Details are pre- sented of two recently unearthed syntypes at the University of Tokyo Museum (UMUTZ), viz. Metaphire communissima (Goto and Hatai, 1899) herein designated the lectotype, and Amynthas levis (Goto and Hatai, 1899) itself a probable synonym of A. tokioensis (Beddard, 1892). Neotypi- fication is flagged both of Amynthas micronarius (Goto and Hatai, 1898), and of Amynthas carno- sus (Goto and Hatai, 1899) that is now separated from other synonyms of the A. corticis (Kinberg, 1867) species-complex. Metaphire hilgendorfi (Michaelsen, 1892) is confirmed in its proper genus based on material labelled “Ph. glandularis.” Unfortunately, mtDNA COI gene barcoding failed to provide conclusive results on these older, formol-fixed samples. Key words : neotypes, syntypes, natives, invertebrate biodiversity, taxonomic history. Tokyo that was to become integrated as the Introduction Komaba campus of University of Tokyo in later While no Pheretima Kinberg, 1867 sensu years, purported to describe “new or imperfectly stricto are known from Japan (Sims and Easton, known species of earthworms collected from var- 1972; Easton, 1981; Blakemore, 2003a, b), the ious parts of the Japanese Empire” (Goto and first scientifically named Japanese pheretimoids, Hatai, 1898, 1899). The first publication was i.e., Pheretima auct. species now in various gen- “Printed September 30th, 1898.” Both publica- era, are shown in chronological order in Table 1. tions were flawed. Naming 28 “new” species, Thus only about ten Japanese species were their descriptions were so inadequate and/or con- reported when Prof. Seitaro Goto (1867–1928) fused that most soon went directly into synon- and his Assistant Shinkishi Hatai (1876–1963) ymy or incertae sedis in Michaelsen (1899) and working from the First High School near Ueno in in his classical review in Das Tierreich 96 Robert J. Blakemore Table 1. Original and current names of earliest Japanese pheretimoid species. No. 　　　　Original name Current combination/synonym 1 *Megascolex sieboldi Horst, 1883 Metaphire sieboldi (Horst, 1883) 2 *Megascolex japonicus Horst, 1883 Amynthas japonicus (Horst, 1883) 3 Megascolex schmardae Horst, 1883 Duplodicodrilus schmardae (Horst, 1883) 4 *Perichaeta ijimae Rosa, 1891 Amynthas corticis (Kinberg, 1867) 5 Perichaeta hilgendorfi Michaelsen, 1892 Metaphire hilgendorfi (Michaelsen, 1892) 6 Perichaeta divergens Michaelsen, 1892 Amynthas corticis (Kinberg, 1867) 7 Perichaeta rokugo Beddard, 1892 Metaphire hilgendorfi (Michaelsen, 1892) 8 Perichaeta nipponica Beddard, 1892 Amynthas corticis (Kinberg, 1867) 9 Perichaeta masatakae Beddard, 1892 Amynthas robustus (Perrier, 1872)? 10 Perichaeta tokioënsis Beddard, 1893 Amynthas tokioensis (Beddard, 1893) * The only three species that Goto and Hatai (1899: 23) later claimed not to have seen. (Michaelsen, 1900) and where many remain described yet more junior synonyms and failed to today or, at best, as species inquirendae in accept nor correct his many earlier errors. Just Michaelsen (1903: 85) (Table 2). one of several examples is Perichaeta vesiculata Prof. Goto made no further offerings and the Goto and Hatai, 1899 (with its spermathecal next earthworm publication was 25 years later by pores likely miscounted, as with several other of Hatai (1924) now in Sendai. In a subsequent his species) along with Pheretima kikuchii Hatai footnote, Hatai (1929: 271) remarked that he had and Ohfuchi, 1936 both being ostensible syn- collaborated (as assistant but designator of “new” onyms of prior Duplodicodrilus schmardae species) with Prof. Goto more than 25 years pre- (Horst, 1883). This perhaps relevant as Tube #4 viously, but that this work was discontinued specimen in the current collection labeled “Ph. (around 1900) owing to change of his residency schmardae” is a misidentification of Metaphire to the USA as a student at Chicago and Professor californica (Kinberg, 1867). Ironically, it seems in Philadelphia. Hatai (1929) said he returned Dr. Hatai retired to Kamakura where many of his permanently to Japan five years earlier (ca. 1923) “species” were from. and for three years preceding publication his col- Mr. Shinryo Ohfuchi was Hataiʼs student and lection locations were (at least): Yunoshima co-author, both working in the Zoological Island; Kominato, Aomori; Sendai, Miyagi; Department of the Saito Ho-on Kai Museum in Uwajima, Ehime (Shikoku); Oshima Island Sendai, established from a charitable trust (The (Tokyo); Kirishima yama and Sakurajima, Saito Gratitude Foundation) which funded col- Kagoshima. Hatai (1929: 274) used formalin to lection trips (e.g., Hatai, 1930) some time before preserve specimens which, although common, is the Museum opened. Later Hatai became director perhaps relevant to specimens examined herein of that Museum (possibly partly on the “strength” that have this poisonous odour. of his taxonomic work) as he continued to pub- In a later paper, unapologetically justifying his lish on earthworms in the Museum journal. initial mis-diagnosis of Metaphire sieboldi Simultaneously, Hatai became a professor at the (Horst, 1883), Hatai (1931a) remarked that Goto Biological Institute of Tohoku Imperial Univer- and Hataiʼs collections were limited to the central sity in Sendai where it appears Ohfuchi also part of Japan (and Taiwan, then part of a Japa- worked since both authors gave both addresses in nese Empire) and to “gardens, refuge piles, pas- joint papers. tures etc.” rather than mountain passes. Hatai Regarding collection localities, Ohfuchi (1931a: 401) also hoped to “straighten up most of (1937: 32, 110) said “The materials upon which the confusions” he claimed were caused by Euro- the present article is based, were collected from pean writers. Nevertheless, his later papers often the six prefectures of Northeast Honshu, Japan, On Opening a Box of Worms 97 Table 2. Most critical errors and mistakes in Goto and Hataiʼs earthworms, all initially in defunct genus Peri- chaeta Schmarda, 1861 now in Amynthas Kinberg, 1867, Metaphire Sims and Easton, 1972 or in Dupodico- drilus Blakemore, 2008; with notes on discovery of some syntypes now in UMUTZ. No. Genus Species Date: page Error statement Syntypes** Actual situation and synonymy 1 Metaphire sieboldii 1898: 65 Spermathecae in 5/6/7/8 — 6/7/8/9, see Metaphire communissima 2 Amynthas fuscatus 1898: 66 *Spermathecae in 6–9 or in 5–7 Possibly Amynthas fuscatus (Goto and Hatai, 1898) 3 Amynthas campestris 1898: 67 *Spermathecae in 8 and 9 or in 6–8; Not found Amynthas robustus (Perrier, 1872)? dorsal pores in 13/14 or 12/13 4 Amynthas kamakuren- 1898: 68 One specimen described from Not found Amynthas gracilis (Kinberg, 1867)? sis “Kamakura, Tokyo” 5 Amynthas parvulus 1898: 68 ? Amynthas gracilis (Kinberg, 1867) 6 Amynthas heteropodus 1898: 69 *Prostates absent (page 69) or ? Amynthas corticis (Kinberg, 1867) present in Table 18 7 Amynthas obscurus 1898: 70 Spermathecal pores shown as ? ? markings 8 Amynthas scholasticus 1898: 70 ? Amynthas corticis (Kinberg, 1867) 9 Amynthas decimpapil- 1898: 71 ? Amynthas gracilis (Kinberg, 1867)? latus 10 Amynthas flavescens 1898: 72 ? Amynthas gracilis (Kinberg, 1867)? 11 Amynthas productus 1898: 73 Male pore presence or absence ? Amynthas gracilis (Kinberg, 1867)? confused 12 Amynthas micronarius 1898: 74 Male pores shown with 10 setae Topotypes Amynthas micronarius (Goto and Hatai, between but said to be only 8 (?syntypes) 1898) 13 Amynthas vittatus 1898: 74 Genital markings confused with Not found Amynthas vittatus (Goto and Hatai, spermathecal pores 1898) 14 Amynthas grossus 1898: 75 Not found Aminthas fuscatus (Goto and Hatai, 1898) 15 Amynthas schizoporus 1898: 76 Genital glands confused with Not found Amynthas tokioensis (Beddard, 1892)? spermathecal diverticula 16 Amynthas takatorii 1898: 76 Glands as spermathecae Maybe Amynthas aspergillum (Perrier, 1872) 17 Amynthas candidus 1898: 77 Maybe Amynthas candidus (Goto and Hatai, 1898) 18 Amynthas irregularis 1899: 13 Setae confused as either 61, ? Amynthas tokioensis (Beddard, 1892)? or 51 on 17 19 Amynthas iizukai 1899: 14 No caeca Missing Simple caeca present; Amynthas fuscatus (Goto and Hatai, 1898) 20 Amynthas shimaensis 1899: 15 Simple caeca present Not found Amynthas fuscatus (Goto and Hatai, 1898)? 21 Amynthas carnosus 1899: 15 Spermathecae in 5/6/7/8 Not found Spermathecae in 5/6/7/8/9 (i.e., in 6–9); (i.e., in 6–8) or in 7–9 A. carnosus (Goto and Hatai, 1899) 22 Metaphire acincta 1899: 16 Clitellum absent Not found Clitellum present; D. acinctus (Goto and Hatai, 1899) 23 Amynthas agrestis 1899: 17 Goto and Hatai (1899: 23) missed Not found Amynthas agrestis (Goto and Hatai, A. agrestis manicate caeca 1899) 24 Amynthas parvicystis 1899: 18 Spermathecal pores and markings ? Amynthas tokioensis (Beddard, 1892) if confused; Goto and Hatai manicate caeca, otherwise cf. (1899: 23) ambiguous on caeca A. masatakae (Beddard, 1892) 25 Metaphire glandularis 1899: 18 Markings mid-7 (miscounted) and ? Mid-8 and doubtfully glands near those glands near spermathecal pores pores; M. hilgendorfi? and male pores 26 Amynthas levis 1899: 20 Papillae around spermathecal pores Yes Amynthas tokioensis (Beddard, 1892)? confused 27 Metaphire vesiculata 1899: 21 Spermathecal pores in 6/7/8 ? but Not found Probably synonym of Duplodicodrilus maybe a mistake schmardae (Horst, 1883) (as for Perichaeta parvicystis)? 28 Metaphire megascoli- 1899: 21 Multiple intestinal caeca Not found Metaphire megascolidioides (Goto and dioides misdescribed Hatai, 1899) 29 Metaphire communis- 1899: 23 See M. sieboldi Yes Metaphire communissima (Goto and sima Hatai, 1899) * Errors indicated by Horst (1889: 242). ** Syntypes published in Blakemore and Ueshima (2011) with several other specimens/labels deteriorated (marked “?”) 98 Robert J. Blakemore from 1934 to 1936. Besides those collected by tion by Easton (1979: 43) of a type of Perichaeta Dr. Hatai in 1923 and myself from the said iizukai Goto and Hatai, 1899 at one time being in region, many specimens from Central and West- the collection of the University of Tokyo ern Japan were also studied.” although no material remained there according to Under Ohfuchi (1937: text fig. 30) details are: Dr. Minoru Imajima, and this confirmed by Drs. Ibuki, Ibuki district, Kagoshima Prefecture, Takenori Sasaki and Rei Ueshima, curators of the March and October, 1928 (Kyushu); Hokonage, University Museum of University of Tokyo Mt. Kirishima, October 19, 1928 (collected by (UMUTZ). Investigations for types in Tohoku Hatai and Araya); Tomitaka, Miyazaki, Miyazaki University and Saito Ho-on Kai Museum, Sendai Prefecture, October 21, 1928; Agricultural were also fruitless, as reported in Blakemore et school, Izumi, Kumamoto Prefecture, October, al. (2010). This due, in part, to the transfer of the 1928; Kochi, Kochi Prefecture, October 10, 1929 earthworm collection of the Saito Ho-on Kai (Shikoku); Matuyama, Ehime Prefecture, Octo- Museum to the National Museum of Nature and ber, 1929 (Shikoku); Izumitsu, Oshima, October, Science (NSMT) in 2006 whence cataloguing of 1927; near Lake Hamana, Shizuoka Prefecture, >1,200 earthworm specimens has been under- August, 1930; Matuyama, Ehime Prefecture, taken by senior curator, Dr. Toshiaki Kuramochi. October 20, 1930 (Shikoku again); Odawara, The eponymous “Box of Worms” in question Kanagawa Prefecture, August 15, 1930; Komaba, was re-discovered on a shelf in the spirit collec- Tokyo Prefecture, June 14, 1931 (significant as tion of NSMT and passed to the author on the this may be a “Tokyo” site); Mito, Ibaraki Prefec- day a fellowship there started on 19th April, ture, August 5, 1930; Sendai, Miyagi Prefecture, 2010. Apparently it had been taken to Tokyo 30 1922–1924 (collected by Hatai?). Ohfuchi (1937: years earlier as the only label (see photo Fig. 1) 121) also noted “The eight hundred and eight stuck on a Ford motor parts box addressed to Dr. specimens examined in this study were collected Imajima at the National Science Museum (Natu- from the localities shown in Text-fig. 30, from ral History Institute) 3–23–1 Hyakunin-cho, 1922 until 1930 by Dr. S. Hatai and Mr. T. Shinjuku-ku, Tokyo 160, Japan, was written in Araya.” Ohfuchi (1938: 2) later mentioned that Japanese and read (Fig. 1): he was helped by Mr. Araya, also a curator of the Saito Ho-on Kai Museum. “1981 (Showa 61) [sic], October, 10th. Their contemporary was Shinjiro Kobayashi Transferred from Saito Ho-on Kai Museum. working from a high school in “Keijo, Korea Imajima and Ishizuka. (＝Seoul)” when it was annexed to Japan who wrote several papers on earthworms (e.g., Oligochaeta Pheretima group (futo mimizu).” Kobayashi, 1939, 1941) with some specimens Earlier inspection of many other specimens supplied by Dr. Hatai as acknowledged in several transferred in 2006 from the Saito Ho-on Kai of his papers published in the Tohoku University Museum collection by the author in 2009 (prior journal. to the discovery of this box) found most in too Search for types. Types are crucial for resolu- poor a condition to determine accurately, being tion of taxonomic confusion. Historical type mainly dried out and variably labeled. In particu- specimens are essential for revision of Japanese lar, the search was for potential syntypes of earthworms, especially the controversial species Hataiʼs later Drawida spp., plus possibly of of Goto and Hatai (1898, 1899) and Hatai Amynthas phaselus (Hatai, 1930) (synonym (1930), but searches had been fruitless until ?maculosus Hatai, 1930), Metaphire yamadai recently (see Blakemore and Ueshima, 2011). (Hatai, 1930), Amynthas? yunoshimensis (Hatai, The whereabouts of early Japanese earthworm 1930), Amynthas tappensis (Ohfuchi, 1935), material was unreported, apart from a brief men- Metaphire tosaensis (Ohfuchi, 1938), Amynthas On Opening a Box of Worms 99 bution now complements. It is entirely appropriate, in the Results below, to first deal with Metaphire communissima (Goto and Hatai, 1899: 23) that was their final species name after Goto and Hataiʼs (1898: 65) initial misidentification of it with “?Perichaeta sieboldii Horst” (＝Metaphire sieboldi), an error that Goto and Hatai (1898: 66) called “preposterous to sup- pose” due to inability on the part of European contemporaries (e.g., Horst, Rosa, Beddard and Michaelsen) to correctly identify Japanese nationalistic science and species. Materials and Ｍethods The eponymous box contained 21 sample tubes with single or pairs of worms (Fig. 1). Each tube included a label reading “The Saito Ho-on Kai Museum, Zool. No.” with some infor- mation written by hand (probably Hataiʼs or Fig. 1. The box of samples from the Saito Ho-on Ohfuchiʼs, or possibly Arayaʼs) in black ink. Most Kai Museum. had new silicone plugs but appeared otherwise unadulterated, only one, slightly larger tube had gomejimensis (Ohfuchi, 1937), Metaphire hataii an original cork plug and it had this note “Dra- (Ohfuchi, 1937) and Metaphire servina (Hatai wida hattamimizu Kanazawa,” as with all other and Ohfuchi, 1937) that were all described samples, it was preserved in formalin. This around 1930s by these authors. Preliminary undissected immature/subadult specimen it is not inspection findings (and omissions) are presented considered further as it is not a pheretimoid nor a in Appendix 1. likely syntype of D. hattamimizu Hatai, 1930, After initial work on this NSMT material was i.e., nothing indicates that it formed a part of completed and a first draft of the current paper Hataiʼs original description (see Blakemore et al., submitted in September 2010, yet another “Box 2010). of Worms” was discovered in the corner of a The remainder of 20 tubes, with the same storage room at Yokohama National University Saito Ho-on Kai Museum labels, were numbered (YNU) that had also been loaned many years ear- by me with prefix “#” and had specimens that lier to Mr. Kotaro Ishizuka and handed to Dr. were, for the most part, previously undissected. Eijiro Nishi in 2002 without any catalogue but They were allocated registration numbers said to contain no types. In fact, this box con- (NSMT-An) and many were sketched, dissected tained crucial historical material, including syn- and described in the authorʼs usual style (Blake- types of Goto and Hatai from UMUTZ that, more, 2000, 2010b–d) in order to provide more because the samples and labels were rapidly accurate identification. These then are the sub- deteriorating, required priority curation. Details jects of the current paper, augmented with data of this second box are provided in Appendix 2. on historical material now returned to UMUTZ. Some information was published in an earlier Tissue samples from non-essential posterior paper (Blakemore and Ueshima, 2011) which the segments of these historical specimens sent for work presented in this thoroughly revised contri- DNA barcoding at the iBOL project at Guelph 100 Robert J. Blakemore University (courtesy Drs. Paul Hebert, Natalia some uncertainty as to its description or position. Ivanova and Sean Prosser) failed to yield usable Another “?” after genus means tentative generic results unlike those by Blakemore et al. (2010) position. Following abbreviations are used: rhs based on fresher, Ethanol-preserved material. — right hand side, lhs — left hand side, GMs — Discussion and justification for specimen sta- genital markings. tus is mostly confined to Remarks following spe- cies and/or specimens descriptions that follow Results ICZN (1999) rules and recommendations espe- cially for explicit designation of neotypes where The box in question and sample specimens of appropriate. A “?” before a taxon name implies concern are shown in Fig. 1 and Table 3. Table 3. Samples in the Saito Ho-on Kai Museum specimen box. Tube Condition of Identification on original label NSMT Current identification Specimen status No. specimen(s) #1 Ph. communissima Goto and An 428 Mature, partly dissected Metaphire communissima Non type specimen (note: many Hatai with its gut still in jar (Goto and Hatai, 1899) other specimens in NSMT) Sendai City 1923–1924 #2 Ph. glandularis An 427 Mature Metaphire glandularis (Goto Non type specimen, synonym Sendai City 1923–1925 and Hatai, 1899) of M. hildgendorfi? #3 Ph. acincta Goto and Hatai/VIII An 429 Mature, dissected and Duplodicodrilus acinctus (Goto Neotype 1930 part of gut missing and Hatai, 1899) #4 Ph. schmardae Horst An 430 Mature Metaphire californica (Kinberg, Misidentified non type Nagasaki City 1867) specimen #5 Ph. agrestis Goto and Hatai An 431 Two matures Amynthas agrestis (Goto and Neotype plus undissected non Morioka (Iwate Pref., Tohoku) and 432 Hatai, 1899) type specimen (also see #17 and Mishima specimens) #6 Ph. hilgendorfi Michaelsen An 434 Mature Metaphire hilgendorfi Non type specimen Sendai City 1923–1925 (Michaelsen, 1892) #7 Ph. carnosa (Goto and Hatai) An 435 Mature Amynthas carnosus (Goto and Neotype. Published in Sendai City 1923–1925 Hatai, 1899) Blakemore (2012) #8 Ph. sieboldi Horst An 436 Aclitellate sub-adult Metaphire sieboldi (Horst, Non type specimen Kochi 17/X 1930 1883) #9 Ph. yunoshimaensis (sic) Hatai An 437 Mature, undissected Amynthas? yunoshimensis Topotipic neotype (possible Yunoshima Aomori Pref. 1922 (Hatai, 1930) synonym of M. hilgendorfi?) #10 Ph. irregularis Goto and Hatai An 438 Mature Amynthas irregularis (Goto and Synonym of A. tokioensis? (cf. Oarai Ibaraki Pref. Hatai, 1899) #11 and #16) #11 Ph. levis Goto and Hatai An 439 Mature Nothing of note differs from A. Non type specimen (cf. #10 and Sendai Kunimi pass 5/X 1930 irregularis #10 #16) #12 Ph. phaselus Hatai An 441 Mature Appears same as Amynthas Syntype? (synonym of A. Aomori Kominato Village 1922 maculosus #13 maculosus?). See #13 #13 Ph. maculosus (Hatai) An 442 Mature, undissected Amynthas maculosus (Hatai, Syntype (synonym of A. Aomori Yunoshima 1927 1930) phaselus?). See #12 #14 Ph. Marenzelli Cognette (sic) An 423 2 matures Amynthas marenzelli (Cognetti, Non type specimens (synonym 24/XI 1929 Aone Onsen Miyagi and 443 1906) of A. corticis) Pref. #15 Ph. vittata Goto and Hatai An 444 Mature Amynthas vittatus (Goto and Neotype (not syntype as Kanagawa, Odawara-Station/VIII Hatai, 1898) collected >1898) 1930 #16 Ph. abnormal An 440 Mature Nothing of note differs from A. Non type specimen cf. #10 and Ibaraki Pref. Kuji Cty Kuji irregularis #10 #11 Village #17 Ph. agrestis Goto and Hatai An 433 Mature Amynthas agrestis See #5 also #18 Ph. sp. An 445 Mature Amynthas micronarius (Goto Non type specimen (cf. #19) Sendai Naga Town Kamohara's and Hatai, 1898) home 16/VI 1931 #19 Ph. micronaria Goto and Hatai An 446 2 matures, both Amynthas micronarius Neotype plus a non type (no other information) and 447 undissected specimen (cf. #18) with publication pending #20 Ph. yamadai Hatai An 448 Mature, undissected Amynthas yamadai (Hatai, Neotype (possibly syntype but Tottori Pref. (collected from 1930) undissected and undated) type locality) On Opening a Box of Worms 101 Tube #1. Metaphire communissima Japan, Nakahama in Province Setsu.Types missing). (Goto and Hatai, 1899) Pheretima communissima: Michaelsen, 1900: 262 (syn- onym sieboldi var. lenzi); Oishi, 1930: 400; Ishizuka, (Fig. 2) 2001: 66, figs. 14.1–8; Minamiya et al., 2007: 56. ?Perichaeta sieboldi: Beddard, 1892: 759. Amynthas communissimus: Sims and Easton, 1972: 235; Perichaeta sieboldii: Goto and Hatai, 1898: 65; Goto and Easton, 1981: 51 (synonymy). Hatai, 1899: 23 (not of Megascolex sieboldi Horst, Pheretima florea Ishizuka, 1999b: 52 [From (Mt. Daibo- 1883). satsu-toge in Yamanashi Prefecture)]. Perichaeta communissima Goto and Hatai, 1899: 23 Pheretima commnissima (lapsus calami): Ishizuka, (Tokyo, Sendai, Tsugaru, Shizuoka, Ibaraki, Bitchū. 1999b: 53. Types unreported). Pheretima frolea (lapsus calami): Ishizuka, 2001: 66. Perichaeta sieboldi lenzi Michaelsen, 1899: 9 (Central Metaphire communissima: Blakemore, 2003b: 7, 28 (new Fig. 2. Metaphire communissima. — a, NSMT-An 428 (Tube #1, previously dissected); b, lectotype UMUTZ- Ann-Og-26, ventral view with spermathecae, prostates and caeca in situ; dorsal view of undissected paralecto- typeʼs prostomium and [boxed] X2 enlargements of non-superficial male pores (18 rhs) of both specimens. 102 Robert J. Blakemore combination, synonyms sieboldi lenzi, florea); Blake- First dorsal pore in 12/13. Setae numerous (60) more and Ueshima, 2011: 64, figs. 2a–d. and crowded ventrally. Spermathecal pores in 5/6/7/8. Genital markings absent. Male pores in Material examined. NSMT-An 428, previously small copulatory pouches (sometimes invagi- dissected around caeca; mature labeled “Ph. com- nated into lateral slits), about 14–20 setae inter- munissima Goto and Hatai, Sendai-City 1923– vene. Septa 8/9 thin or absent and 9/10 aborted 1924,” [same label as Tubes #2 (M. glandularis), around gizzard, from 10/11 onwards thin. Sper- #6 (M. hilgendorfi) and #7 (A. carnosus) speci- mathecae in 6–8 roundish ampullae often some- mens herein], but since this date is after 1899 what rugose with long, convoluted diverticula. publication it is not a syntype although from a Seminal vesicles large in 11 and 12. Ovaries in stated type-locality. Many other M. commmunis- 13 with small pseudovesicles on 12/13 just above sima specimens from the Saito Ho-on Kai the ovaries; small ovisacs on 13/14 (in NSMT- Museum are in the NSMT collection, in various An 428). Prostates in 17–20 with long, muscular states of preservation (see Appendix 1). Subse- duct to slight copulatory pouch. Last hearts in 13 quently found syntypes now in UMUTZ-Ann- (those in 10 not found in NSMT-An 428). Intes- Og-26 were described in Blakemore and tine from 15; caeca manicate with about 5–9 “fin- Ueshima (2011), see Fig. 2b and now I choose gers” from 27, a low lamellar ridge but no typh- the previously dissected specimen (figured) as losole found; gut contains mucous-enveloped lectotype (under ICZN, 1999: Article 74 and soil with a few grits. NSMT-An 428 has grega- ICZN Declaration 44) in order to enhance the rine cysts around its prostates and scattered else- stability of nomenclature; the remaining where internally. UMUTZ specimen in the same jar becomes the Remarks. In contrast to the current species, paralectotype. and contrary to Goto and Hatai (1898), M. Distribution. Japan. Ishizuka (2001) implies sieboldi is one of the most striking and easily that geographical or topographic locations of his recognized of Japanese species due to its bril- P. florea specimens coming from a Yamanashi liant, iridescent blue colouration in the adult mountain is unique, but he appears to ignore the form (possibly to deter bird predation). Other dif- distribution of M. communissima given in (Goto ferences from M. sieboldi—apart from sperma- and Hatai, 1898: 66, 1899: 23) as from Bitchū thecal pores not in 6/7/8/9—are that here the (＝Okayama Prefecture) to around Osaka, male pores are perhaps slightly wider, almost lat- through Shizuoka, Tokyo, Ibaraki and Sendai to eral, and spermathecal ducts appear longer and Aomori or, as Goto and Hatai state “that is to say more muscular. all over the Main Island.” Michaelsen (1899) Ishizuka (2001: 66) redescribes communissima gives us Tottori and Easton (1981: 51) quotes on the same page as his florea (misspelt frolea) “Ohfuchi, 1938d” and others extending the range giving their respective lengths as 90–180 and to southern Hokkaido; all this putting Yamanashi 60–80 mm, but bigger worms almost always about central within its known range. grow from smaller ones. The only other differ- Diagnosis based on NSMT-An 428 specimen ence is spermathecal ampullae stated to be and UMUTZ-Ann-Og-26 syntypes. NSMT-An “shovel-shaped” in florea as opposed to “globu- 428: 170 mm long with 116 segments. Lectotype, lar” in communissima, even though Michaelsen 125 mm with 103 segments (but lacking its pos- (1900: 262) had stated they were “flattened” and, terior tip), undissected paralectotype 130 mm; rather obviously, it is irrelevant as ampullae by Goto and Hatai (1899: 23) say up to 250 mm or their nature can be either inflated or deflated due more with 140 segments, but usually round to use and “packing.” Both his figured specimens 190 mm with about 100 segments. Pale grey with (Ishizuka, 2001: figs. 14, 15) have non-superfi- buff clitellum 14–16, or puce (NSMT-An 428). cial male pores within copulatory pouches, sup- On Opening a Box of Worms 103 porting their inclusion in Metaphire, and are by Blakemore (2012) of A. carnosus (Goto and essentially indistinguishable, supporting their Hatai, 1899), that this same label location and synonymy. Were parthenogenetic specimens of date of “Sendai 1923–1925” for Tubes #1 (M. M. communissima to lack male pores, these communissima), #2 (M. cf. glandularis), #6 (M. would presumably be similar (synonymous?) to hilgendorfi) and #7 (A. carnosus) specimens, either A. agrestis (Goto and Hatai, 1899) or M. possibly mean simply that the specimens were in hataii (Ohfuchi, 1937: 13) specimens that also the collection in Sendai at that time (see Intro- lack genital makings. Thus, the relationship of duction), and not necessarily from there. M. communissima to the prior A. agrestis and Remarks. Perichaeta glandularis was subsequent M. hataii may require resolution described by Goto and Hatai (1899: 18) as dor- should their characteristics overlap. sally banded, with spermathecal pores in 6/7/8, The NSMT specimen conforms tolerably to markings in 7 (mistake for 8?) and 17/18 and fig- the UMUTZ syntypes that both agree with Goto ured with male pores in copulatory pouches (i.e., and Hataiʼs original précis and with later descrip- Metaphire), but in other regards complying with tions, although its name may sometimes be found Michaelsenʼs prior α morph of M. hilgendorfi. misspelt as “communisima” or “communisimma.” Their original description is problematical as the authors (Goto and Hatai, 1899: 19–20) state “Sometimes the posterior borders of the sper- Tube #2. ?Metaphire glandularis matheal pores are surrounded by similar papil- (Goto and Hatai, 1899) lae . . . and a fourth group of 8–9 glands close to (as part of a Metaphire hilgendorfi species-group) each male pore” and they figure (Goto and Hatai, 1899: figs. 10, 11) these secondary capsuloge- (Fig. 3) nous glands opening near the spermathecal and Perichaeta glandularis Goto and Hatai, 1899: 18, figs. 9–11. [From “Takahashi (Prov. Bitchū),” now male pores, whereas such glands are more usu- Okayama Pref., Type unknown]. ally associated with Amynthas tokioensis (Bed- Pheretima glandularis: Michaelsen, 1900: 315 (as a pos- dard, 1892) that typically lacks the central genital sible “variety” of his P. hilgendorfi); Kobayashi, 1941: markings and has superficial male pores, or with 260; Gates, 1958: 11–13 (as possible synonym of P. A. vittatus (as redescribed herein). Possibly they hilgendorfi). relate to parasitic artefacts or, more likely, the Metaphire glandularis: Sims and Easton, 1972: 238 (in a now defunct Metaphire glandularis species-group); figures by Goto and Hatai are composite images Easton, 1981: 51 (include separately in his Amynthas of several species/specimens. On current knowl- hilgendorfi species-complex); Blakemore, 2003b: 29, edge, no one has seen such an arrangement of 2005: 108, 2007: 115, 2010a: 12 (as a junior synonym features in any subsequent worm in the last 112 of Metaphire hilgendorfi). years and neither does this specimen conform, Material examined. NSMT-An 427, a mature thus itʼs identification as “Ph. glandularis” may specimen labeled “Ph. glandularis Sendai 1923– be questioned. 1925,” previously undissected, here dissected and Specimen NSMT-An 427 is uniformly figured. So far (see Blakemore and Ueshima, coloured, 150 mm long with 114 segments, has 2011) a glandularis syntype has been elusive. spermathecae in 7/8/9 and a unilateral non-super- Distribution. The label locality, if actually ficial male pore on 18 lhs; its marking are mid- from Sendai, is far removed from the original ventral in 8 (rather than 7) and anteriorly in 18 Takahashi collection site; however, as a junior and it thus complies with prior M. hilgendorfi, as synonym of M. hilgendorfi (Michaelsen, 1892), redescribed by Blakemore (2003a, 2003b, 2005, its distribution includes Hokkaido, all of central 2010a, in prep.), to synonym Perichaeta rokugo Japan, as well as Korea and USA. Beddard, 1892, and also to Amynthas? yunoshi- It should be noted here and for the description mensis (Hatai, 1930) as mentioned below. Proba- 104 Robert J. Blakemore Fig. 3. Metaphire glandularis (a) and Metaphire cf. glandularis (＝M. hilgendorfi) (b). — a, Goto and Hatai (1899: figs. 9–11); b, NSMT-An 427 showing spermathecae, 18 rhs prostate from non-superficial male pore and intestinal caeca in situ, and genital markings glands overlain by ventral nerve cord. bly M. glandularis should be considered a misde- Tube #3. Duplodicodrilus acinctus scribed synonym of prior M. hilgendorfi, or, at (Goto and Hatai, 1899) comb. nov. best, a species incertae sedis. As noted in Discussion, also by Gates (1982: (Fig. 4) 52), this specimenʼs single, non-superficial male Perichaeta acincta Goto and Hatai, 1899: 16, fig. 6 pore exemplifies correct placement in genus (Tokyo. Types unknown). Metaphire Sims and Easton, 1972. Conversely, Pheretima acincta: Michaelsen, 1900: 252; Hatai, 1931b: 182, fig. 32; Ohfuchi, 1957b: 1360, fig. 3849; Yamagu- under some authorsʼ schemes, one half of this chi, 1962: 10 (synonym yezoensis); Kamihira, 1973: specimen would belong to one species and genus 57; Minamiya et al., 2007: 56. and the other half to another, this being clearly Amyntas acinctus: Beddard, 1900: 650. ridiculous. Amynthas acinctus: Sims and Easton, 1972: 235 [hawaya-