Drawida japonica ( Michaelsen, 1892 )

Blakemore, Robert J., Lee, Seunghan & Seo, Hong-Yul, 2014, Reports of Drawida (Oligochaeta: Moniligastridae) from far East Asia, Journal of Species Research 3 (2), pp. 127-166 : 132-135

2. Drawida japonica ( Michaelsen, 1892)

Moniligaster japonicus Michaelsen, 1892: 232-233 . [From Japan. Syntype in Hamburg Museum: 403, but originally stated by Michaelsen as in Zoological Museum, Berlin: Verm. 2122 and this syntype also listed by Hartwich & Kilias (1989: 268 as “ Japan; HILGENDORF leg.” just after Allolobophora japonica (Verm. 2117) . Both were inspected by Blakemore in Blakemore & Kupriyanova (2010: 9), they were collected by Herr Dr Franz Hilgendorf, the latter definitely at Enoshima on 29.III.1875 ‾ thus this is possibly a type-locality of the D. japonica syntype although my searches of the island have thus far proved fruitless for this species; yet other Berlin specimens from the Hilgendorf series were definitely not collected there (e.g. Michaelsen’s record of M. sieboldi that must be from further south). Gates (1939: 411-413) did not mention #403 but he inspected three specimens from Hamburg labeled “ V 1194. Drawida japonicus Mich. f. Typ. Dr Chen F. Wu c. Dr Michaelsen a. Nanking, China ” and other materials none of which were types, saying (on page 413) that the original two types ‾ described by Michaelsen as “ Diese Art ist durch ein geschlechtstreifes und ein unreifes Exemplar vertreten ” ‾ were sectioned and are no longer available for study. This is incorrect as #2122 is a dissected specimen 26 mm long with the last 24 of its 95 segments regenerated that, although the internal organs are mostly removed, is nevertheless in good condition. Note the “ Typ.” in the China label refers to D. japonicus f. TYPica not to a true type specimen].

[? Moniligaster bahamensis Beddard, 1893: 690 , figs. 1- 5. Included by Michaelsen (1910: 50) as a D. japonica subspecies, it is now held in D. barwelli synonymy].

Drawida japonica : Michaelsen, 1900: 115; 1910: 48; Stephenson, 1922: 119, figs. 1-6; Chen, 1933: 189, fig.?; Gates, 1935: 3 (maintaining D. graham and questioning Chen’s ‘ D. japonica ’); Chen, 1936: 291 (syn. grahami ); Kobayashi, 1937, 1938: 94, fig. 1; Gates, 1939: 411 ( cf. grahami ); Kobayashi, 1940: 263 (part. + propatula ); 1941: 458, 515; Chen, 1959: 15, fig. 22; Ohfuchi, 1965: 546 (plus fig. apparently copied from Chen); Easton, 1981: 37 (part. excluding grahami and with China not included in distribution range); Blakemore,

August 2014 BLAKEMORE ET AL.-REPORTS OF DRAWIDA 133

5 GM 7/8rhs 9/10rhs 10 10 GM Nps 12 15 15 14 ba ab cd ba ab 1 mm

Fig. 3. Drawida ashiuranoeri”), with in situ spermatheca and male organs (testis sac, vas deferens and prostates with adjacent GM gland) and gizzards in 12-14 plus ovisacs from 12-18; [boxed enlargement of 8rhs spermathecal atrium], also nephridium from 14/15rhs. It is remarkably similar superficially to both D. koreana austri and D. koreana shindo , apart from the lack of blue colour, although the DNA data is unequivocable that they are all separate taxa (cf. D. companio ).

2003; 2005/2007 (syn. grahami from China).

Drawida japonica typica : Michaelsen, 1910: 49; 1927: 85; Stephenson, 1917: 366, fig. 1; 1922: 126; 1923: 142, fig. 52 (part. syn. bahamensis and explaining Michaelsen’s initial incomplete description).

Drawida japonicus typicus : Michaelsen, 1931: 7.

Drawida japonicus : Michaelsen, 1931: 523 (part. syn. siemsseni that is sometimes mispelt “siemmseni”).

Drawida grahami Gates, 1935: 3 ; 1939: 408. [From Suifu, Szechuan (Sichuan). Type (s) USNM: 20093. Said to have more ventral spermathecal pores in mid-bc otherwise cf. D. japonica ; three gizzards in 12-14 (as in three of Gates’ five subsequent D. japonica specimens); one of seven of Gates’ types lacked ‘genital markings’, described in just a dozen lines on a dozen characters].

Drawida propatula Gates, 1935: 449 . [From China. Types USNM 20179. Said by Kobayashi (1938: 94, 1940: 263) to be similar although Kobayashi (1940: 265) maintain- ed them separately only on extents of the ovisacs]. Syn. nov. Blakemore in Blakemore & Kupriyanova (2010: 9) cf. D. siemsseni .

Drawida japonica : Blakemore, 2005/2007 (syn. grahami ); 2008; Blakemore et al., 2010: 1; Blakemore & Kupriyanova, 2010: 8, figs. 3.1-3.7, tabs. 1-3 (syn. propatula ).

Note. Drawida calebi Gates, 1945: 211 from India is a similar species, and should probably be combined. Here is it briefly summarized including information from Julka (1976 Mitt. Zool. Mus. Berlin, 52(2): 322): Unpigment- ed. Length 35-83 by 2-4.5 mm. Segments 115-170. Dorsal line present but no dorsal pores. Spermathecal pores just median to c-lines. Male pores slits at 10/ 11 in mid-bc. Nephropores close to d lines. GMs small, unpaired and median or closely paired in aa, and widely paired in bc, presetal and postsetal variously in 7-13. Gizzards two or three or four of in some of 12-17 (as shown in a table). Vas deferens short, in a small column of loops in 9, straight in 10 and entering the prostate directly. Prostates are “not quite spheroidal”, sessile, smooth. Short penis present internally. Ovaries in conjoined 10/11 and 11/12 with ovisacs to 20. Spermathecal ampulla is small, conical or nearly spheroidal on duct four to five mm long to apex of atrium that may have glands on its anterior wall like those associated with the GMs. From Jubbulpore, Madhya Pradesh (type locality) and Nowgong, Satna, Manikpur and Tanda Falls (Gates) also Jharkhand, Orissa, Uttar Pradesh and Karnataka districts found in forests, pastures, croplands and compost pits.

Material examined. Berlin Kat. Nr. 2122 (“ Drawida japonica Syntype! Japan. Hilgendorf ”) a dissected specimen 26 mm long with the last 24 of its 95 segments regenerated and probably Michaelsen’s ‘25 segmenten regeneriert Hinterende ’- RJB pers. obs. IV.2010 ; the internal organs are mostly removed although the specimen is otherwise in good condition. Hamburg # 403 syntype label states collected by Hilgendorf in Japan but is only a desiccated posterior portion (ca. 8 mm and 25 segments comprising mostly intestinal soil). Five specimens, two from “ Aichi ” [kanji for the prefecture] (one with dorsal pores and GMs and one without either) plus three from fields at Nagura-shi near Nagoya, Aichi-ken (one with dorsal pores but no GMs, the other two without either) all stored in Kyorin Uni, Hachioji collection (RJB inspected 20.IX. 2002); Watarase one specimen without dorsal pores but with GMs in 9lhs posteriorly and 10rhs anteriorly, and male pores in 10/11 on eversible penes with gizzards in 12-14 (coll. IV.2003 by Dr Takafumi Kamitani of YNU and RJB inspected 9.IV.2004); one specimen collected from Kamakura Daibutsu shrine 13.VI.2004 by RJB (identified tentatively); four specimens from rice fields in Hikone (collected 19.VI.2009 by RJB) ‾ one dissected and figured plus host for mtDNA COI sample (GenBank GQ500902 ); plus three others, all lacking dorsal pores but having GMs as detailed below (these latter specimen registrations are LBM 1380000085 2009-13-3) .

A mature, grey specimen 80 mm long with 133 segments, dissected and figured here obtained beside suiden at Ashiura-cho, Kusatsu, Shiga-ken en route to Kanon complex, collected 18 th March, 2011 by RJB and registered LBM FY2010-22.3 providing DNA tissue for JET 101-11 initially identified as “ Drawida ashiuranoeri” (COI results in Appendix).

Not found in current Korean studies (cf. genetic barcodes of “ Drawida cf. japonica ”, WO 21, and D. koreana shindo, WO 27 below and in Fig. 1).

Distribution. Japan, from around Tokyo to Nagasaki and the Ryukyus, southern China, Taiwan, Korea (including Quelpart/ Jeju-do), and south-east Asia. Inclusion of D. propatula adds central and northeast China. Stephenson (1923: 143), Gates (1939: 413 six specimens also from Murree) and Paliwal & Julka (2005-http://www.zoosprint. org) list it outside the normal Drawida domain from the western Himalayas, India; and it is reported (as “ Drawida japonica Michaelsen, 1917 ”) from the famous Punjab beer-district of Solan, Himachal Pradesh (Dhiman & Battish, 2005-http://www.zoosprint.org/). [Note. Stephenson’s (1923) description included bahamensis that is now included in barwelli as it typically lacks ‘genital markings’; cf. Japanese D. eta specimens lacking markings that have a male pore on extractible penis rather than external and ‘flap-like’ and cf. Gate’s (1982: 18) D. bahamensis held separately].

A recent unpublished report is from Kansas, USA by Dr M. Callam (pers. comm. June, 2013 at SES meeting New Jersey, USA), unconfirmed due to current revisions.

Remarks. Full description and distribution is in Blakemore & Kupriyanova (2010) updated by Blakemore (2012b) where it is noted that male pore is superficial lateral of b on porophore on 10 that overhangs 10/11. Both Stephenson (1923) and Gates (1935) mistook a GM in 10 as male pore even though Michaelsen (1892: 232; 1900: 115) had them in 10/11 and overhanging segment 11 after a pair (or unilateral) markings in 10, this repeated by Gates, 1939. Types and current specimens have male pores on 10 posteriorly on protruded ‘flaps’ overhanging 10/11 and just lateral to b lines (pers. obs. and from types as per Michaelsen and Kobayashi) or as eversible penes in the Watarase specimen that may thus be suspect]; Hikone specimens have ventral region of segments 10 and 11 flared around the male pores to present a flattened area but lack the markings found in D. impertusa .

Correct identification is elusive as Fig. 1 phylogram shows “ Drawida japonica ” from KOBIC Korea that agrees with “ Drawida japonica ” from China although both differ from Drawida japonica from Japan as listed in Blakemore & Kupriyanova (2010). In Blakemore et al. (2010), it was also noted that COI barcodes (by Huang et al. 2007 and Chang et al. 2008) from Chinese source material differed by ~17% from a Lake Biwa sample and that conspecificity of these Chinese entities needs to be reexamined.

Results of examination of D. japonica types were presented in Blakemore & Kupriyanova (2010) and the London types of D. barwelli were reexamined by the author in June, 2013. Resolution yet requires checking of Chinese specimens, at least those still available, for comparison with synonymic Drawida grahami Gates, 1935 types (Smithsonian #20093) also with erstwhile sub-species Moniligaster bahamensis Beddard, 1893 , plus its synonym, D. propatula Gates, 1935: 449 from China. The Drawida nepalensis ( Michaelsen, 1907) species-complex that is reported from Nepal, India, Pakistan, Andamans, Myanmar, China (Yunnan), Java and Sumatra; and D. ramdadana ( Michaelsen, 1907) is a reminiscent taxon. If D. burchardi Michaelsen, 1903 from Andamans and Sumatra is indeed synonymous with D. nepalensis ‾ as suggested by Gates 1962: 331; 1972: 244, 256 ‾ then taxonomic priority is another question.

species, from which:

!o D. japonica differs from D. barwelli mostly in the form of male pores posteriorly on 10 and on its GMs (when present).

!o D. koreana differs from D. japonica mostly in its blue colour and penile details.

!o D. keikiensis differs from D. barwelli by its accessory spermathecal glands.

!o D. moriokaensis differs from D. japonica mostly on forms of male pores.

Kobayashi (1938: 106) remarked that length of spermathecal duct appeared to separate D. japonica from his D. koreana ‾ in the former it is long and coiled whilst in the latter it is shorter and thicker (see Fig. 11 ). Drawida impertusa differs from D. barwelli in its lack of dorsal pores, and GMs in 10. Lake Biwa’s Drawida eda Blakemore, 2010 , has eversible male pores on penes, five gizzards in 13-17, lacks both pigment and genital markings, and in has an accessory genital gland near spermathecal atria internally in 7. The same sort of gland is also report- ed for D. nemora (only in some specimens!?), D. keikiensis , D. tairaensis and D. jeholensis . Chinese Drawida jeholensis Kobayashi, 1940 is unpigmented, has irregular markings in 7-11, male pores on blunt poropores overhanging 10/11, and two or three gizzards in 11,12-13; thus, the only major differences from D. japonica is presence of a penis rather than a flap and of the accessory gland in 7 near its spermathecal atrium.

Subspecies Drawida japonica siemsseni ( Michaelsen, 1910) is redescribed below as D. siemsseni ; other names by Oishi, 1934 (minuta and gigantic) are absorbed in the nominal species.

Figures

Fig. 11. Drawida koreana austri sub-sp. nov. A ‾ Holotype, B ‾ Paratype dissected to show spermatheca, male organs and gizzards in 12- 14. Boxed are Kobayashi’s (1938: figs. 1 & 3) comparing spermathecae of C ‾ Drawida japonica and D ‾ D. koreana koreana that differ from those of the current specimens.