Robert J. Blakemore

Japanese earthworms (Annelida: Oligochaeta): a review and checklist of species

Abstract The current revision provisionally lists 77 valid earthworm taxa in seven families from Japan, with approximately 80 further names (ca. 50% of the total) either in synonymy – including 40 new synonyms – or retained as species incertae sedis. Generic placement of the species yielded 17 new combinations. About 30 species are known introductions and another ten are possibly more widespread, thus the probable number of wholly endemic Japanese earthworms is around 40 species (ca. 50% of the total valid species). However, a definitive work on the systematics of Japans earthworms is pending, and the current revision aims only to provide a status quo and to track changes from the last comprehensive revision by Easton (1981) that listed 74 taxa. Subsequently, 60 or so new pheretimoid names were added by Ishizuka in 1999–2001, but only a few are considered valid taxa, the remainder being synonyms or species incertae sedis. The substitute name ‘Pheretima’ palarva, nom. nov., is provided for the junior homonym Pheretima parvula Ishizuka et al., 2000. While much of Eastons synopsis is supported, Pontodrilus is now placed in Megascolecidae sensu Blakemore (2000) rather than Acanthodrilidae sensu Gates (1959); Amynthas carnosus (Goto & Hatai, 1899) is removed from synonymy with Amynthas gracilis ; and an informal Amynthas corticis species-complex is established to accommodate the various morphs of this widely distributed species group. Pheretima (Parapheretima) koellikeri Michaelsen, 1928 is considered synonymous with Metaphire vesiculata (Goto & Hatai, 1899), thereby removing the genus Pheretima sensu stricto from the Japanese list. Polypheretima is also removed from Japanese indigeny, as the original description of Polypheretima iizukai (Goto & Hatai, 1899) failed to report intestinal caeca, and inspection of fresh material allows its placement in synonymy with Amynthas fuscatus (Goto & Hatai, 1898). Easton (1981) had listed this taxon as Metaphire fuscata , but further demonstration of superficial male pores qualifies it for Amynthas . Conversely, the discovery of copulatory pouches results in transfer from Amynthas to Metaphire for M. hilgendorfi (Michaelsen, 1892), comb. nov., Metaphire communissima (Goto & Hatai, 1898), comb. nov., and Metaphire megascolidioides (Goto & Hatai, 1899), comb. nov. The Metaphire hilgendorfi/Amynthas tokioensis species-complex ( Amynthas hilgendorfi species-complex sensu Easton 1981) remains one of the most intractable and pressing problems for comprehension of the Japanese fauna, as most of the component taxa, e.g. Metaphire agrestis (Goto & Hatai, 1899), are parthenogenetically degraded morphs as yet unaffiliated with their ancestral and biparental populations. Resolution may be sought employing combinations of morphological and molecular (RNA, DNA) techniques to determine specific affinities while also complying with requirements of the International Code of Zoological Nomenclature (ICZN 1999).

Saga of Herr Hilgendorf’s worms…

Abstract Herr Dr. Franz Hilgendorf, who first introduced Darwin’s evolutionary theory to Tokyo in 1873, collected ‘Vermes’ for Dr. Wilhelm Michaelsen in Hamburg. The Metaphire hilgendorfi (Michaelsen, 1892)/Amynthas tokioensis (Beddard, 1892) parthenogenetic/clonal spp-complex has since snowballed into >60 names, and its resolution remains the hottest yet seemingly intractable problem in Oriental (and Cosmopolitan) earthworm systematics. Reproductive structures, morphometrics, colouration or intestinal caeca characterizations are largely defunct. Molecular ‘solutions’ are meaningless without DNA analysis of types under the strict ICZN Principle of Typification in chronological order under its Principle of Priority. A revised diagnosis now accepts Metaphire spp from morphs having non-superficial male pores. Both Amynthas tokioensis (syns. ?M. levis; ?A. paiki syns. nov.) as a new record from USA and A. agrestis (Goto & Hatai, 1899) (syn. ?A. minjae Hong in Hong, Lee & Kim, 2001 syn. nov.) from Japan/Korea are reviewed. Metaphire soulensis (Kobayashi, 1938) and ?M. koellikeri (Michaelsen, 1928) are restored separately but the dubious A. defectus (Gates, 1930) (syn. A. jacita) is newly added to the group. Work is urgently needed to separate Metaphire Sims & Easton, 1972 from Amynthas Kinberg, 1867 and to sort degraded morphs under their respective types. More than a generation ago, Gates (1972) said naming intermediates is “ridiculous”. Despite this, names continue to be added by workers in Japan or Korea who mutually ignore earlier work in either country: Dozens of ‘nationalistic’ Japanese ‘Pheretima’ synonyms have been added as have Korean taxa with manicate caeca e.g., A. yongshilensis, A. alveolatus, A. geomunensis, A. eastoni, A. boletiformis, A. odaesanensis, A. righii, A. fasciiformis, A. sanchongensis, A. songnisanensis, A. ephippiatus and A. multimaculatus. A degraded digestive ‘tube’ from Korea named as Amynthas dageletensis Hong & Kim, 2005, plus A. sonjaesiki Hong & James, 2009 (syns. novae of A. tokioensis), have the lowest priority within this 118-year-old saga. Critical conditions of the intestinal caeca are briefly considered, and the emerging 117-year-old synonymy saga of Pheretima urceolata (Horst, 1893) is flagged as a new taxonomic ‘housekeeping’ concern. For all these issues, molecular resolution via DNA anaylsis of types is advocated.

Reports of Drawida (Oligochaeta: Moniligastridae) from far East Asia

The Moniligastridae Claus, 1880 presumably arose in a region now occupied by the Malay Archipelago sufficiently long enough ago to have widely colonized and speciated in the Far East and to have radiated westwards even as the Himalayas formed to occupy the Indian subcontinent as far as Sri Lanka (Michaelsen, 1909; 1922; Gates, 1972). A less popular alternative is the family has “recent Indian origin” colonizing Asia after collision of the Indian Plate (Easton, 1981: 34), although most authors agree on a southern Indian homeland concentration for genus Drawida speciation. Along with holarctic Lumbricidae in the Northern Hemisphere, Acanthodrilidae in the Southern Ocean and Octochaetidae in Indo-Australasia, it is considered one of the most naturally widespread of earthworm families (Gates, 1972; Blakemore, 2013c). It currently has approximately 215 valid species in six genera (Csuzdi, 2012), mainly in South Asia and southeast Asia. The family diminishes further East, or rather its taxonomic study does; for example, Kobayashi (1940: 311) reporting on Manchurian worms listed less than a dozen East Asian species although these are still important in agricultural and ecological studies to this day. Only a few are peregrine cosmopolitans (Blakemore, 2012b), in particular Drawida barwelli (Beddard, 1893) is widely transported in the tropics, e.g. first Australasian moniligastrid report by Blakemore (1994), albeit many records of this small worm are likely mistaken as it was confused even in its earliest descriptions. A major difficulty with moniligastrid description, apart Journal of Species Research 3(2):127-166, 2014

A comprehensive checklist of earthworm species and subspecies from Vietnam (Annelida: Clitellata: Oligochaeta: Almidae, Eudrilidae, Glossoscolecidae, Lumbricidae, Megascolecidae, Moniligastridae, Ocnerodrilidae, Octochaetidae)

The comprehensive checklist of earthworms in Vietnam is presented here listing 24 genera and 212 species arranged in eight families. For each species, bibliographic citations are given, including the original descriptions as well as notable citations affecting taxonomic status or distributional reports of a species. Distributions of each species are also provided. The bibliography contains all relevant papers for Vietnamese earthworms. Of 212 species, 114 have been recorded only in Vietnam, and 25 are cosmopolitan. New combinations are made for Amynthas acalifornicus (Do & Huynh, 1991) comb. nov., A. binhgiaensis (Le, 1994) comb. nov., Metaphire catbaensis (Thai & Le, 1993) comb. nov., M. khoii (Do & Tran, 1994) comb. nov., M. phaluongana (Do & Huynh, 1992) comb. nov., and M. mangophila (Nguyen, 2011) comb. nov., M. tripidoporophoratus (Thai & Nguyen, 1993) comb. nov. Additionally, Pheretima paraalexandri Nguyen, 2011 is treated as a junior synonym of Amynthas polychaetiferus (Thai, 1984).

Barcoding Eophila crodabepis sp. nov. (Annelida, Oligochaeta, Lumbricidae), a Large Stripy Earthworm from Alpine Foothills of Northeastern Italy Similar to Eophila tellinii (Rosa, 1888)

A new Italian earthworm morphologically close to the similarly large and anecic Eophila tellinii (Rosa, 1888) is described. Distribution of Eophila crodabepis sp. nov. extends over 750 km2 from East to West on the Asiago Plateau and Vittorio Veneto Hills, from North to South on mounts Belluno Prealps (Praderadego and Cesen), Asiago, Grappa and onto the Montello foothills. This range abuts that of Eophila tellinii in northern Friuli Venezia Giulia region. Known localities of both E. tellinii and E.crodabepis sp. nov. are mapped. mtDNA barcoding definitively separates the new western species from classical Eophila tellinii (Rosa, 1888).

Account of montane and insular speciation in some Korean megadriles (Annelida: Oligochaeta)

In the spirit of Kobayashi (1936; 1937; 1938; 1941) and Easton (1981), the current work contributes to the first author’s current Japanese and Korean studies, e.g. Blakemore (2003; 2008; 2012a-f; 2013a; 2013b; 2014), and attempts to use genetic barcodes of primary types or neotypes for definitive species identity (Hebert et al., 2003; Blakemore, 2013c). Current specimens are from mountains such as Seoraksan (alt. 1,708 m) the highest mountain in the 500 km Taebaek Range that edges the east coast as far down as Busan and the third highest after Hallasan and Jirisan in the south. Further collection was at nearby Bangtaesan (alt. 1,444 m) and Jeombongsan (alt. 1,424 m). Islands were also sampled, and the briefest of surveys on the isle of Muuido near Incheon unearthed two new species: a megascolecid Amynthas muuido sp. nov. and a lumbricid Eisenia muuido sp. nov., the latter comparable to endemic Eisenia koreana (Zicsi, 1972) that was surprisingly overlooked in Hong’s (2000) incomplete review of Korean Lumbricidae (compare his nine or ten spp. to the twenty listed in Blakemore, 2014). The current paper complements an update of moniligastrid Drawida spp. in Blakemore et al. (2014).

Restating scope of genus Metaphire Sims & Easton, 1972 40 years on

Abstract Clarification of correct placement of species in genus Metaphire Sims & Easton, 1972 is required as differentiation from Amynthas Kinberg, 1867 hinges on condition of male pores in their respective type-species. Confusion is due in part to misconception of rules of ICZN and partly because of problems of parthenogenetic degradation of male organs. In summary, Amynthas is the default genus of pheretimoids with superficial or absent male pores, Metaphire species differs by having non-superficial male pores and in Pheretima Kinberg, 1867, which is not known in East Asia, taxa further acquire nephridia on spermathecal ducts. The type of Metaphire, M. javanica (Kinberg, 1867), was misidentified in Australia but M. californica (Kinberg, 1867), its possible synonym, is widespread and is now confirmed in Korea. This latter cosmopolitan is still often confused with Japanese Duplodicodrilus schmardae (Horst, 1883) that has eversible intromittent organs developed much more so than in Metaphire. Figures are provided.

Call for a Census of Soil Invertebrates (CoSI)

Abstract A case is argued for urgent reappraisal of biodiversity of soils in order to mitigate their rapid global decline (MEA). A first step is to compile a “stock inventory” of soil fauna thought to number around 210,000 species. Basic knowledge is yet wanting and even earthworms are poorly known despite being the major component as well as key “environmental-engineers” and vital links in all terrestrial food webs (including in waterways, hence their excellence yet trivial use as bait). That biodiversity of earthworms is disproportionately underappreciated is surprising as, with 10,000 already named and many more expected, they are no less species-rich than marine polychaetes, for example, that number ca. 8,000 valid taxa. A model for CoSI is the 10 yr,

Biodiversity of Earthworms in Taiwan: a Species Checklist with the Confirmation and New Records of the Exotic Lumbricids Eisenia fetida and Eiseniella tetraedra

1 billion global Census of Marine Life (CoML) that concluded with 250,000 total ocean taxa, but since 2 million species are already catalogued and estimates of diversity are of 10 million, this represents 12.5% of described species and just 2.5% of a probable total. Even claims that oceans occupy two thirds of the planet overlooks that land is hilly and the relative surface areas are perhaps 50:50. Socio-economic arguments flounder in context of 99% of the total worldwide human food supply produced on land, whereas oceans and other aquatic ecosystems provide a paltry 0.6% (FAO). Thus it seems timely and appropriate to advocate a sea change to firmly ground eco-taxonomic studies on our diminishing soils that support all life on Earth and, via runoff, provision or pollute the oceans too.