Pheretima (Parapheretima) saba Sims & Easton, 1972

Blakemore, Robert J., Csuzdi, Csaba, Ito, Masamichi T. & Kaneko, Nobuhiro, 2007, Taxonomic status and ecology of Oriental Pheretima darnleiensis (Fletcher, 1886) and other earthworms (Oligochaeta: Megascolecidae) from Mt Kinabalu, Borneo, Zootaxa 1613, pp. 23-44 : 34-39

Pheretima (Parapheretima) saba Sims & Easton, 1972

( Fig. 6 )

Pheretima (Parapheretima) saba Sims & Easton, 1972: 265 from region of Mesilau Cave at 2,100–2,400m and ridge between Mesilau Rivers at 2,750m. Syntypes in Natural History Museum, London: BMNH 1971:19: 88–89, 90/93 (not re-inspected here).

Diagnosis: Parapheretima occurring on Mt Kinabalu with paired spermathecae in 5/6/7. Genital markings absent. Holandric. Intestinal caeca simple in 27.

Distribution: Mt Kinabalu , Sabah, Malaysia. [Type locality near Mesilau Cave].

Material examined: mature specimen, dissected and sketched by RJB – NSMT-An 370, fixed in formalin at collection and stored in 80% ethanol with a spermatheca removed to a separate vial in jar; from vicinity of Paka Cave, Mt Kinabalu , Sabah, Malaysia collected by Tatsuya Kawaguchi, 13.vii.2005 (original sample labeled “ 050713 PAKA 5-B” also contained specimens of Metaphire paka as described above).

Lengths: 60–119 mm; current specimen 60 mm, Sims & Easton (1972) had size range 87–119 mm.

Width: ca. 2–4 mm.

Segments: 107–135; current specimen 107, Sims & Easton (1972) had range 112–135.

Colour in formalin: faint brown anteriorly; clitellum yellow/buff.

Prostomium: open epilobous.

First dorsal pore: 11/12–13/14 (12/ 13 in current specimen).

Setae: 28–40 per segment; Sims & Easton (1972) had range 34–40; current specimen has 28–30 [actual data ca. 31 on 7, 28 on 20, 22–26 posteriorly; no setae between male pores (pers. obs. not previously noted by Sims & Easton)].

Nephropores: diffuse, not found.

Clitellum: annular 14–16, setae obscured.

Male pores: on 18 ca. 0.3 circumference apart on large eversible copulatory structures protruding from male pores described by Sims & Easton “ as large crenellate copulatory pouches 0.3 of the body circumference apart ” with no setae intervening (current specimen); actual male pores seem to be just anterio-median to large sucker-like pads (pers. obs. and see sketch).

Female pores: single, midventral on 14.

Spermathecal pores: paired in 5/6/7 approximately 0.3 circumference apart.

Genital markings: absent.

Septa: 4/5-7/8 slightly thickened, “ 8/9,?9/10 absent ” according to Sims & Easton and both absent in current specimen; 10/11 and subsequent thin.

Dorsal blood vessel: single (pers. obs).

Hearts: 10–13 (cf. Sims & Easton who just state “ Last lateral heart in xiii [13]”.

Gizzard: “ between 7/8–?9/10, 10/11 ” according to Sims & Easton, found to be large between 7/8–10/ 11 in current specimen.

Calciferous glands: none.

Intestine origin (caeca, typhlosole): in “ xv [15]” (Sims & Easton) or 18 where it thins and greatly dilates (pers. obs. on current specimens); caeca small, simple from 27–25,23; small spherical typhlosole present from about 27 (pers. obs. on current specimens – this not noted by Sims & Easton); gut contents not noted.

Nephridia: meroic, “ present on spermathecal ducts ” according to Sims & Easton, but in current specimen found to be few in number and scarcely present on duct as they can almost be teased back to origins on body wall; nevertheless, when a spermatheca was removed they are seen attached to the duct where it too enters the body wall..

Testes/sperm funnels: holandric, testes in 10 and 11 in testis sacs, seminal vesicles paired in 11 and 12; with pseudovesicles in 10 and 13 (pers. obs. in current specimen not noted by Sims & Easton).

Ovaries: paired, palmate in 13 with funnels posteriorly; ovisacs not found.

Prostates: large racemose in 17–20; muscular ducts pass into large copulatory pouches that each contains a large penis and externally bears a long tubular (?secretory) diverticulum that is perhaps related to eversible pads on copulatory organs.

Spermathecae: paired in 6 and 7 each with ovoid ampulla on shorter duct with small clavate diverticulum; two or three nephridia attach near the base of duct (pers. obs. as noted above).

Remarks. Sims and Easton (1972: 266) compared their species with Ph. ( Ph.) koellikeri ( Michaelsen, 1928) from Japan for which it is not surely know whether nephridia are apparent on the spermathecal ducts, thus, according to Blakemore (2006a), it ‘defaults’ to Metaphire rather than qualifying for Pheretima . Both taxa have what are called ‘secretory diverticula’ on their copulatory pouches and, at least in Ph. saba , this qualifies it for Parapheretima sub-genus. In Ph. (Pa.) saba these diverticula may relate to the large eversible copulatory organs that seem to have (?adhesive) pads whereas in M. koellikeri they exit near penial setae ( Michaelsen, 1928). The attached sketch is the first time Ph. (Pa.) saba has been figured.

Genus Polypheretima Michaelsen 1934 sensu Easton (1979)

Polypheretima everetti ( Beddard & Fedarb, 1895) Figs. 7–8

Perichaeta everetti Beddard & Fedarb, 1895: 69 ; Beddard, 1895: 428 (non Amynthas stelleri everetti: Michaelsen, 1899: 43: 1900 : 306 where P. everetti was combined with P. kinabaluensis ). Type locality stated by Beddard & Fedarb as “ Mount Kina Balu ”, but based on type labels Easton (1976: 43) redesignated it under what is now ICZN (1999: 76A) as on the adjacent Balabac Island, Palawan southern Philippines. Syntypes in Natural History Museum, London: BMNH 1904: 10:5:38–40 inspected by Easton (1976: 51). [GMs paired on 19–21; spermathecal batteries of 12–17 in segments 6 and 7].

Perichaeta papillata Beddard & Fedarb, 1895: 70 , 71; Beddard, 1895: 428 (redescription). From Merabah, Borneo. Syntypes British Museum: BMNH 1904: 10.5.1265 –70 inspected by Easton (1976: 52). [GMs paired on 19–29; spermathecae 7 per segment in 6 and 7].

Perichaeta sarawacensis Beddard and Fedarb, 1895: 71 ; Beddard, 1895: 429. From Labuan, Sarawak. Holotype British Museum: BMNH 1904.10.5.150 inspected by Sims & Easton (1972: 254). [GMs paired on 19–22; spermathecae 14 in segments 6 and 7].

? Perichaeta barami Michaelsen, 1896: 203 . From Baram River, Sarawak and/or (Minahassa) North Celebes – Michaelsen was unsure. Holotype Hamburg: v3835 inspected by Sims & Easton (1972: 254) and they only give the former locality; same specimen listed by Easton (1976: 51) as from “ Minahassa ” and Easton (1976: 52) as “ Baram River ”. [Grey, setae 42–55 per segment, GMs on 19,20, spermathecae absent (as in Po. elongata )].

Amyntas stelleri seriatus Michaelsen, 1899: 44 (originally spelt seriata). From either Uangkahulu Valley or Buol, North Celebes. Paratype Hamburg: v5197 inspected by Sims & Easton (1972: 254); and v.5198 inspected by Easton (1976: 51). [Brownish with violet shimmer, setae 42–111, GMs 19–22,23, spermathecal groups of 3–6 in segments 6 and 7].

? Amyntas stelleri klabatensis Michaelsen, 1899: 46 . From Klabat, North Celebes, 1,800m. Paratype Hamburg: v.5196 inspected by Sims & Easton (1972: 254) and claimed as “ Holotype ” by Easton (1976: 51). [Setae 72–120, no GMs, spermathecal groups of 8–11 in 6 and 7 (lack of GMs is a departure from Po. everetti diagnosis below and is therefore similar to Po. annamensis ( Stephenson, 1931) that however is bithecal].

? Amyntas stelleri bonensis Michaelsen, 1899: 45 . From Bone Valley, North Celebes. Types lost. [Blue in life, setae ca. 130, GM paired in 19, spermathecal groups of 18–26 in 6 and 7 (as in Po. stelleri s. stricto)].

Amynthas stelleri everetti (part): Michaelsen, 1899: 43 (where P. everetti was combined with P. kinabaluensis ).

Pheretima stelleri everetti (part): Michaelsen, 1900: 306.

Pheretima stelleri koroensis Michaelsen, 1910: 109 . From Koro Valley, Celebes (Sulawesi). Holotype listed on Basel Museum type website: http://www.nmb.bs.ch/NaturmuseumBasel/LinksNMB/Sammlung/Kataloge/Guertelwuermer.xls].

Pheretima stelleri mahakkami Michaelsen, 1922: 25 . From Mahakkam River, Sarawak. Holotype Leiden: 1897 inspected by Sims & Easton (1972: 254).

Pheretima (Pheretima) beranensis Michaelsen, 1928: 23 . From Birang River, Beran district, Kalimanton. Type (s) Hamburg: v.10576 inspected by Easton (1976: 52).

Pheretima (Pheretima) baritoensis Michaelsen, 1932: 9 . From Boentok on the Barito River, Kalimanton. Types lost.

Pheretima (Polypheretima) beranensis tinjarana Michaelsen, 1934: 25 . From Tinjar River, Long Lejok, Sarawak. Syntypes British Museum: BMNH 1933.10.6.12–20 and Hamburg: v.11951 inspected by Sims & Easton (1972: 254) and Easton (1976: 52).

Metapheretima everetti: Sims & Easton, 1972: 233 ; Easton, 1976: 41,fig. 5 [syns. papillata , sarawacensis , barami , stelleri seriatus (originally spelt seriata), stelleri klabatensis , stelleri bonensis , stelleri koroensis , stelleri mahakkami , beranensis , baritoensis , beranensis tinjarana ].

Polypheretima everetti: Easton, 1979: 54 ; Blakemore, 2000, 2004: 137, 2006b.

Taxonomic Note: two of Easton’s synonyms, at least, must be questioned as P. barami lacked spermathecae (as in Po. elongata ) and P. stelleri bonensis had high numbers of spermathecal batteries of 18–26 (as in Po.

stelleri ); moreover, A. stelleri klabatensis lacked genital markings as described in the diagnosis below.

Diagnosis: Polypheretima with male pores in copulatory pouches without stalked glands. Colour reddishpurple. Length up to 300 mm with up to 260 segments. Spermathecal batteries (each usually with 3–12 spermathecae) in 5/6/7. Setae numerous (up to 130 per segment in large specimens). Genital markings paired, presetal in line with male pores in some or all of 19–21,22,23. Holandric. Intestinal caeca absent. (Cf. Po. elongata , Po. stelleri and Po. kinabaluensis ).

Distribution: Balabac Island south of Palawan, Philippines (new type locality rather than Mt Kinabalu designated by Easton, 1976: 43); north and west Celebes; Borneo including Mt Kinabalu (from 1,200– 2,400m); and Lombok, Indonesia (350–450m only). Easton (1976: 43) claims that this species is unknown outside its indigenous range, although this is quite extensive and some of the island records (eg. Balabac and Lombok) may be due to transportation. It occurs on lowlands of Sabah, Borneo (current specimens) yet is sympatric with Po. kinabaluensis at higher altitude on Mt. Kinabalu (Easton, 1976) . Current specimens examined were from lowlands East of Mt Kinabalu , collected by T. Kawaguchi 01.iii.2006 (labeled: “ 060301 T-3” and “ I-2 S”), along with other samples (labeled: “ 060307 R4-2”, “ 060308 R1L”, “ 060307 R4-2”, and “ 060308 ET”), of Pontoscolex corethrurus plus other immature pheretimoids, and samples of immature specimens (labeled: “ 060305 T3-3”, “ 060302 R-1-15” and “ 060302 R-1-2-3”) that look like striped and more regularly coloured forms of Pheretima ( Ph.) darnleiensis (as noted in its description above).

Remarks. Although it is similar to Po. elongata , and sympatric with this and other members of the Po. elongata species-complex, e.g. Easton (1976: fig. 6) shows it to occur throughout Borneo and with Po. elongata on Lombok; with Po. phacellotheca and Po. stelleri in northern Celebes; and with Po. kinabaluensis on Mt Kinabalu , yet Easton (1976: 42) states that there is no indication of hybridization among the species. Easton (1976: 52) further separated off one Lombok specimen (Berlin: 7214) from the type series of Po. badia (Ude, 1932) as Po. everetti .

Figures

FIGURE 6. Pheretima (Parapheretima) saba from region of Paka Cave, Mt Kinabalu; figure shows a, prostomium; b, a spermatheca (with an enlargement to show nephridia on duct); and c, prostate gland and accessory diverticulum in situ (with section of diverticulum); d, a dorso-lateral section of intestine to show caeca and typhlosole; and e, dorsal view of posterior segments. Note: the copulatory pouches that everted on preservation would normally in life be withdrawn into body leaving only slits of male pores visible externally.

FIGURE 7. Polypheretima everetti showing ventral view and anterio-dorsal view with a - spermathecal batteries in situ, paired genital markings (GM) in 19 and b - prostate ducting to male pores (original sketch by RJB from from YNU Mt Kinabalu / north Borneo specimens).

FIGURE 8. Polypheretima everetti with more extensive genital markings (GM); Sp. pr. – position of spermathecal pore batteries (modified after Easton, 1976: fig. 5).