Polypheretima Michaelsen 1934

Aspe, Nonillon M. & James, Samuel W., 2016, NeW species OF Pheretima, Amynthas, Polypheretima, and Pithemera (Clitellata: MegascOlecidae) FrOm MindanaO and assOciated islands, Philippines, Zoological Studies 55 (8), pp. 1-33 : 24-27

Genus Polypheretima Michaelsen 1934

Type species: Perichaeta stelleri Michaelsen

1892.

Generic Diagnosis

Body cylindrical; setal arrangement perichaetine; annular clitellum covering segments xiv-xvi; pair of male pores in xviii on circular porophores that may be within copulatory bursae; ventral genital markings present or absent; esophageal gizzard in viii; intestine begins in xv or xvi; nephridia on spermathecal ducts lacking; caeca lacking; male sexual system usually holandric, with testes and funnels enclosed in paired sacs in x and xi; seminal vesicles in xi and xii; spermathecal pores small, spermathecal diverticula simple and usually ectal in origin;

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prostates racemose; copulatory bursae may or may not be present; ovaries free in xiii; oviducts lead to single or closely paired opening ( Easton 1979).

Polypheretima bukidnonensis n. sp. ( Fig. 5A, 5B , Table 4 ) urn:lsid:zoobank.org:act:A39D1ED9-69B3-4E63-9A90-F7A6D258AA6C

Material examined: Holotype: adult ( NMA 4598 ), municipality of Maramag , Bukidnon Province, Mt. Musuan Range (7°52'53.8"N, 125°03'57.6"E), 400 m asl, Mindanao Island, Philippines, coll. N. Aspe, 26 Oct. 2012 . Paratype: one adult, amputee (NMA 4614), same collection data as for holotype.

Etymology: The species is named for the province of Bukidnon.

Diagnosis: Brown worm, adult length 131 mm; paired spermathecal battery pores at intersegments 5/6/7; 7-11 pyriform-shaped spermathecae in each battery; male pores 0.22 circumference apart; 39-45 setae on vii, 46-58 setae on xx; genital markings widely paired on xix to xxi, in line with the male pores; gizzard origin in viii; copulatory bursae lacking.

Description: Brown dorsum, pale ventrum, equators pigmented. Length 131 mm (n = 1 adult); diameter 5-6.5 mm at x, 5.5-6.5 mm at xx; body circular in cross-section, tail tapering; 147 segments. First dorsal pore at 12/13, paired spermathecal battery pores at intersegments 5/6/7. Female pore single in xiv, openings of male pores paired in xviii, distance between pores 3.8- 4.5 mm (0.22 circumference apart ventrally), 6-7 setae between pores. Clitellum annular, from xiv to xvi. Setae unevenly distributed around equators in some segments; 39-45 setae on vii, 46-58 setae on xx, dorsal setal gaps present, ventral setal gaps lacking. Paired genital markings widely spaced on xix to xxi, in line with male pores.

Septa 4/5 membranous, 5/6-7/8 and 10/11- 13/14 thin, 8/9/10 lacking. Dense tufts of nephridia on anterior faces of 4/5, 5/6, and 6/7; nephridia of intestinal segments located mainly on body near septum/body wall junction. Large gizzard in viii-x, esophagus with low vertical lamellae x-xiii, intestinal origin in xv, caeca lacking; typhlosole originating in xxvi, simple fold, 1/3 lumen diameter. Hearts in x to xiii, esophageal; commissural vessels in vi, vii, and ix lateral; those in viii extend to gizzard.

Ovaries and funnels free in xiii. Spermathecal

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batteries paired, postseptal in vi and vii. One adult individual with eight spermathecae closely aligned on left side and nine spermathecae closely aligned on right side in vi, and eight spermathecae closely aligned on left side and 10 spermathecae closely aligned on right side in vii. Another adult individual with eight spermathecae closely aligned on left side and 11 spermathecae closely aligned on right in vi, and seven spermathecae on left and 11 on right in vii. Spermathecae small, ampulla pyriform, spermathecal duct short, slender; diverticulum stalk long and slender, attached ectally to duct, terminating in short, sausage-shaped receptacle; no nephridia on spermathecal ducts. Male sexual page 25 of 33

system holandric, testes and funnels enclosed in paired ventral sacs in x and xi; seminal vesicles in xi and xii; pseudovesicles in xiii; vasa deferentia slender, free from body wall on way to ental end of prostatic ducts; prostates in xvii to xix, each prostate racemose, compact; muscular duct from lateral margin of prostate widens towards body wall, then narrows slightly just before body wall; copulatory bursae lacking.

Remarks: Polypheretima bukidnonensis n. sp. belongs to the Po. elongata Perrier 1872 group of Easton (1979), characterized by having paired genital markings in xix and successive segments in line with the male pores, paired batteries of

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up to 28 spermathecae in vi and/or vii, and a shallow copulatory bursae with no stalked glands. Easton (1979) included five species in the group: Po. elongata ; Po. everetti Beddard and Fedarb 1895 ; Po. kinabaluensis Beddard and Fedarb 1985 ; Po. phacellotheca Michaelsen 1899 ; and Po. stelleri Michaelsen 1892 . Polypheretima mindanaoensis Aspe and James 2015 from Mt. Malindang was recently added to this species group. Among members of Po. elongata group, Po. bukidnonensis is similar to Po. elongata and Po. everetti in having paired genital markings widely spaced on xix to xxi or xxi, in line with the male pores. However, Po. elongata and Po. everetti are large worms (355 mm and 300 mm, respectively); the male pores are more widely spaced (0.25 circumference apart ventrally); there are many more setae per segment (up to 130), whereas Po. bukidnonensis has only 39-58 setae; and these species have copulatory bursae, whereas Po. bukidnonensis lacks them. Polypheretima elongata is also non-pigmented and rarely has more than three spermathecae per battery in 5/6/7, while Po. everetti has 6-17 spermathecae per battery in 5/6/7, and there are 16 setae between male pores. Polypheretima bukidnonensis is similar to Po. mindanaoensis in size (90-118 mm × 5.1-7 mm) and in lacking copulatory bursae, but the latter is non-pigmented and the genital markings extend from xix to xxvi. In addition, Po. mindanaoensis has fewer spermathecae (0-5) per battery in 5/6/7, and its gizzard originates in viii.

Polypheretima zamboangensis n. sp. ( Fig. 5C, 5D , Table 4 ) urn:lsid:zoobank.org:act:CC5B1AC9-260E-4E39-BE08-01A417ED5776

Material examined: Holotype: adult ( NMA 4599 ), Brgy Timolan , municipality of Tigbao , Zamboanga del Sur Province, Mt. Timolan (7°48'00.0"N 123°15'00.0"E), 1000 m asl, Mindanao Island, Philippines, coll. N. Aspe, 18-20 Dec. 2012 . Paratypes: five adults (NMA 4615); four adults (ZRC.ANN 0068), same collection data as for holotype.

Etymology: The species is named for the province of Zamboanga del Sur.

Diagnosis: Slender worm, adult body dimension 223-306 mm × 3.5-5; body non-pigmented; 227-248 segments; spermathecal pores paired and inconspicuous at intersegments 5/6/7, or lacking; 60-84 setae on vii, 69-70 setae on xx; male pores 0.31 circumference apart;

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gizzard in x; metandric male sexual system.

Description: Body non-pigmented, equators non-pigmented. Length 2230- 306 mm (n = 10 adults); diameter 3.5-4 mm at x, 4-5 mm at xx; body circular in cross-section, tail tapering; 227- 248 segments. First dorsal pore at 12/13; paired spermathecal pores at intersegments 5/6/7, inconspicuous, lacking in some specimens. Female pore single in xiv, openings of male pores paired in xviii, distance between openings 3.9 mm (0.31 circumference apart ventrally), 8-9 setae between openings. Clitellum annular, from xiv to xvi. Setae irregularly distributed around equators in some segments; 60-84 setae on vii, 69-70 setae on xx; two first ventral setal lines prominent; dorsal and ventral setal gaps present. Paired genital markings widely spaced on xix to xxii, in line with male pores.

Septa 4/5-9/10 muscular, 11/12-13/14 thin, 10/11 lacking. Dense tufts of nephridia on anterior faces of 5/6-8/9; nephridia of intestinal segments located mainly on body near septum/body wall junction. Gizzard in x-xi, esophagus with low vertical lamellae x-xiii, intestinal origin in xvii, caeca lacking; typhlosole originating in xx, simple fold, about 1/5 lumen diameter. Hearts in xi to xiii, esophageal; commissural vessels in vi, vii, and ix lateral; those in viii extend to gizzard.

Ovaries and funnels free in xiii. In one specimen, spermathecae two pairs, postseptal in vi and vii, spermathecal ampulla pyriform, spermathecal duct short, slender; stalked diverticulum attached to ental portion of left face of duct near ampulla of right spermatheca, and on left face of duct of left spermatheca; stalks short, terminating in small ovate receptacle; no nephridia on spermathecal ducts. Spermathecae lacking in other specimens. Male sexual system metandric, testes and funnels enclosed in paired ventral sacs in xi; seminal vesicles in xii and xiii; pseudovesicles in xiii; vasa deferentia slender, free from body wall on way to ental end of prostatic ducts; prostates in xvi to xix, each prostate racemose, compact, with five minor lobes; stout U-shaped muscular duct from lateral margin of prostate widens towards body wall, then narrows slightly just before body wall; copulatory bursae lacking.

Remarks: A metandric male sexual system and paired spermathecal pores in 5/6/7 indicate that Po. zamboangensis n. sp. belongs to the Po. huoensis Easton 1979 species group. However, members of this species group are confined to Papua New Guinea ( Easton 1979) and have a different genital marking pattern (genital markings

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present on vii-ix, xvii, and xix). Ten species in Polypheretima have been reported from the Philippines: Polypheretima monticola Beddard 1912 from Benguet Province, Luzon Island; Po. elongata Perrier 1872 and Po. everetti Beddard and Fedarb 1895 , both from Balabac Island, Palawan Province; Po. fruticosa Hong and James 2008a , Po. perlucidula Hong and James 2008a , and Po. bannaworensis Hong and James 2008a , from Banaue, Luzon Island; Po. pagudpudensis Hong and James 2011a from Kalbaryo, Luzon Island; and Po. mindanaoensis Aspe and James 2015 , Po. bukidnonensis n. sp. (this study), and Po. zamboangensis n. sp. (this study), from Mindanao Island. Among these species, Po. zamboangensis is similar to Po. perlucidula and Po. pagupdudensis in having a pair of spermathecae in vi and vii and in lacking body pigmentation. However, the latter two species are smaller ( Po. perlucidula , 38-56 mm x 1.3-1.5 mm; Po. pagudpudensis , 56-63 mm x 2.1-2.2 mm); Po. perlucidula lacks genital markings, and there are only epidermal thickenings on segment xviii in Po. pagudpudensis . In addition, in the latter two species the pre-clitellar setae are more numerous than the post-clitellar setae (50 and 23, respectively, in Po. perlucidula ; 63 and 27 in Po. pagupudensis ), the origin of gizzard is in viii, and the male sexual system is holandric. Polypheretima zamboangensis has more segments (227-248) than all other Philippine Polypheretima species ( Po. fruticosa , 105-107; Po. perlucidula , 117-131; Po. bannaworensis , 67-95; Po. pagudpudensis , 126-136; Po. mindanaoensis M, 140-141; Po. bukidnonensis n. sp., 147; Po. everetti , 126) except for Po. elongata (136-297). But the male pores in Po. zamboangensis are more distant (0.31 circumference apart ventrally) compared to that of Po. elongata (0.25 circumference apart ventrally), the gizzard is bigger (x-xi vs. ix in Po. elongata ), the intestinal origin is in xvii (vs. xv in Po. elongata ), and the prostates are smaller (xvi-xix vs. xvi-xxi in Po. elongata ). The other species have holandric male sexual system and either have spermathecal batteries in 5/6/7 ( Po. bukidnonensis n. sp., Po. mindanaoensis , Po. elongata , and Po. everetti ), have four pairs of spermathecae in 5/6- 8/9 ( Po. monticola and Po. bannaworensis ), or have only one pair in 5/6 ( Po. fruticosa ).

Figures

Fig. 5. Schematic views of the internal and external morphology of the Polypheretima and Pithemera species described here. (A, B) Polypheretima bukidnonensis; (A) dorsal view of internal morphology; (B) external ventral view; (C, D) Polypheretima zamboangensis n. sp.; (C) dorsal view of internal morphology; (D) external ventral view; (E) Pithemera nolani n. sp., dorsal view of internal morphology, with prostates in xv-xix. Abbreviations: s, spermatheca; h, heart; p, prostate gland; cb, copulatory bursa; c, caecum; spbp, spermathecal batteries pores; cl, clitellum, mp, male pores, gm, genital markings. Scale bars: A-E, 5 mm.