Pheretima Kinberg 1867

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 : 4-18

Subgenus Pheretima Kinberg 1867

Type species. Pheretima montana Kinberg 1867

Generic diagnosis

Body circular in cross section, with numerous setae regularly arranged equatorially around each segment; setae absent on first and last segments. Male pores paired within copulatory bursae opening on segment xviii; one or more pairs of spermathecal pores in intersegmental furrows between 4/5 and 8/9. Clitellum annular, covering three segments (xiv to xvi). Single midventral female pore on xiv. Genital markings usually

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absent. Internally, esophageal gizzard usually originating in viii; a pair of caeca originating in xxvii, extending forward. Ovaries and funnels free in xiii. Male sexual system holandric, with paired testes and funnels enclosed in sacs in x and xi, and seminal vesicles in xi and xii. Spermathecae one pair, multiple pairs, sometimes single and located midventrally, or sometimes lacking. Nephridia present on spermathecal duct(s). One pair of prostate glands, racemose. Copulatory bursae present; secretory diverticula lacking on coelomic surface of copulatory bursae.

Pheretima acia n. sp. ( Fig. 2A , Table 2 ) urn:lsid:zoobank.org:act:9C182FE8-4413-4598-AFAF-C8A746D1AA28

Material examined: Holotype: preclitellate ( NMA 4585 ), Brgy (Barangay) Boa, municipality of Cagdianao, Dinagat Province, (10°05'53"N, 125°39'42"E), 250 m asl, Dinagat Island, Philippines, coll. N. Aspe, J. Adeva, 23-26 Apr. 2004 . Paratypes: two adults (NMA 4601), same collection data as for holotype.

Etymology: The species name is from the Latin acia (thread, yarn), referring to the small, thread-like body.

Diagnosis: Very small, brown worm with thread-like body; adult length 38-47 mm, diameter 1-1.2 mm; pigmented segmental equator; pair of spermathecal pores at intersegment 5/6; prostates small in xvii to xviii; penes present.

Description: Brown, equators pigmented. Length 33-47 mm (n = 3 adults); diameter 1.2 mm at x, 1 mm at xx; body circular in cross-section, tail tapering; 87-94 segments. First dorsal pore at 12/13, paired spermathecal pores at 5/6, spermathecal pores 0.8 mm (0.21 circumference apart ventrally). Female pore single in xiv, openings of copulatory bursae paired in xviii, distance between openings 0.8 mm (0.24 circumference apart ventrally), 3 setae between openings. Clitellum annular, from xiv to xvi. Setae unevenly distributed around equators in some segments; 26-35 setae on vii, 29-38 setae on xx, dorsal and ventral setal gaps lacking. Genital markings lacking.

Septa 4/5-7/8 and 10/11-13/14 thin, 8/9/10 lacking. Dense tufts of nephridia on anterior faces of 5/6 and 6/7; nephridia in 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 page 5 of 33

simple, originating in xxvii, extending forward to xxvi; typhlosole originating in xxvi, simple fold, 1/3 lumen diameter; intestinal wall with 38 longitudinal blood vessels. 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. Spermathecae paired, postseptal in vi, with nephridia on ducts. Each spermatheca with ovate ampulla; slender, bulbous, muscular duct; single stalked diverticulum attached to the ental portion of the right face of duct of right spermatheca, and on left face of duct of left spermatheca; stalk short, terminating in ovate to sausage-shaped receptacle. Male sexual system holandric; testes and funnels enclosed in paired sacs in x, xi; seminal vesicles xi, xii, each with digitate dorsal lobe; vesicles of xi enclosed in testes sac; vasa deferentia slender, free from body wall on way to ental end of prostatic ducts; prostates small, in xvii to xviii; each prostate a single, dense, racemose mass; short muscular duct entering anterior margin of copulatory bursa. Copulatory bursae round, in xvii-xix; coelomic surfaces muscular, secretory diverticula lacking; roof with thick pad, floor with thin pad; short conical penis present.

Remarks: Pheretima acia n. sp. belongs to the P. urceolata species group of Sims and Easton (1972), characterized by having spermathecal pores opening only on intersegment 5/6. Sims and Easton (1972) assigned P. urceolata Horst 1893 and P. baweanensis Michaelsen 1928 to this species group, but Gates (1961) synonymized these two species. Currently, there are 20 species members of the urceolata group, including the new species described here. Pheretima acia differs markedly from any member of the P. urceolata group and from the rest of the congeners in having a very small, thread-like body. Other relatively short members of P. urceolata group have body diameters two to three times that of P. acia : P. dinagatensis n. sp. (65 mm × 2.5-3 mm), P. lantapanensis n. sp. (> 58 mm × 2.5-3 mm), P. abiadai Hong and James 2008c (33-60 mm × 2.2-3 mm), P. nagaensis Hong and James 2008c (36-53 mm × 2.5-3 mm), P. viracensis Hong and James 2009a (38-61 mm × 2-2.6 mm), and P. doriae Hong and James 2009a (34-45 mm × 2-2.4 mm). Like Pheretima acia , all these species except P. lantapanensis have penes, but the spermathecal pores in P. acia are more closely spaced (0.21 circumference) than those of P. dinagatensis (0.31), P. abiadai (0.25), P. nagaensis (0.29), P. viracensis 0.32-0.36) and P. doriae

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(0.32). The male pores in P. acia are closer (0.24) than those in P. doriae (0.32) and more widely spaced compared to that of P. abiadai (0.18- 0.2). Pheretima acia has small prostates, which only extends in two segments (xvii-xviii) while the other species have relatively bigger prostates, covering three to five segments (xv-xix). Although P. lantapanensis also has prostates that extends in page 6 of 33

two segments, they are in xvi-xvii. In addition, the caeca in P. acia are very short, covering only two segments extending only from xxvii-xxvi, whereas the other species have caeca that are longer, extend from xxvii-xxv or -xxiv or -xxiii ( Table 2 ). No other species in the P. urceolata group closely resemble P. acia .

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Pheretima dinagatensis n. sp. ( Fig. 2B , Table 2 ) urn:lsid:zoobank.org:act:1ABBFFA9-4720-444B-919F-C0260B43F486

Material examined: Holotype: adult ( NMA 4586 ), Brgy Boa , municipality of Cagdianao, Dinagat Province, (10°05'53"N, 125°39'42"E), 250 m asl, Dinagat Island, Philippines, coll. N. Aspe, J. Adeva, 23-26 Apr. 2004 . Paratypes: two adults, amputee (NMA 4602); three adults, amputee (ZRC.ANN 0055), same collection data page 7 of 33

as for holotype.

Etymology: The species is named after Dinagat Island.

Diagnosis: Worm with adult length of 65 mm; reddish brown dorsal stripes at intersegmental furrows, ventrum pale; pair of spermathecal pores at intersegment 5/6; first dorsal pore 11/12; spermathecal pore distance 0.31 circumference apart ventrally; setae between male pores numerous, 12-14; penes present.

Description: Reddish brown dorsal stripes at intersegmental furrows, pale ventrum, equators

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non-pigmented. Length 65 mm (n = 1 adult); diameter 2.5 mm at x, 3 mm at xx; body circular in cross-section, tail tapering; 127 segments. First dorsal pore 11/12, paired spermathecal pores at 5/6, positioned on the lateral margins, distance between spermathecal pores 2.4 mm (0.31 circumference apart ventrally). Female pore single in xiv, openings of copulatory bursae paired in xviii, distance between openings 2.3 mm (0.24 circumference apart ventrally), 12-14 setae between openings. Clitellum annular, from xiv to xvi. Setae irregularly distributed around equator of some segments; 39 setae on vii, 43-48 setae on xx, dorsal and ventral setal gaps lacking. Genital markings lacking.

Septa 4/5-7/8 and 10/11-13/14 membranous, 8/9/10 lacking. Dense tufts of nephridia on anterior faces of 5/6 and 6/7; nephridia in 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 xiv or xv; caeca simple, originating in xxvii, extending forward to xxiii, may be folded up or tucked under intestine in xxv or xxiv; typhlosole originating in xxvi, simple fold, 1/3 lumen diameter, intestinal wall with 31- 33 longitudinal blood vessels. 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. Spermathecae paired, postseptal in vi, with nephridia on ducts; each spermatheca with round ampulla and slender, bulbous, muscular duct; single stalked diverticulum attached to the ental portion of right face of duct of right spermatheca, and on left face of duct of left spermatheca; stalks short, terminating in small, round to oval receptacle. Male sexual system holandric; testes and funnels enclosed in paired sacs in x, xi; seminal vesicles xi, xii, each with digitate dorsal lobe; vesicles of xi enclosed in testes sac; vasa deferentia slender, free from body wall on way to ental end of prostatic ducts; prostates in xv or xvi to xx; each prostate a single, dense, racemose mass with four lobes; muscular duct entering center of copulatory bursa, forming a coil. Copulatory bursae round in xvii-xix; coelomic surfaces muscular, secretory diverticula lacking; roof with two thick pads, floor pads lacking; short conical penis present.

Remarks: Pheretima dinagatensis belongs to P. urceolata group of Sims and Easton (1972). Members of P. urceolata group with body dimensions similar to P. dinagatensis n. sp. include P. lantapanensis n. sp., P. abiadai , P. nagaensis , P.

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viracensis , and P. doriae but the former differs from the latter four in having dorsal stripes. Pheretima dinagatensis is also similar to P. lantapanensis n. sp. in the location of the first dorsal pore but differs from the latter in having more setae between the male pores and more setae in the post-clitellar area (8 and 31-36, respectively in P. lantapanensis ), and in having penes. Pheretima dinagatensis has more body segments (127) and has more setae between the male pores (12-14) than P. abiadai (74-78 segments and 6-9 setae, respectively), P. nagaensis (57-76 and 7-10, respectively) P. viracensis (83-96 and 7-10, respectively), and P. doriae (77-86 and 5-8, respectively). The first dorsal pore in P. dinagatensis is on 11/ 12 in P. dinagatensis but on 12/ 13 in P. abiadai and P. nagaensis . Like P. abiadai , P. nagaensis , P. viracensis and P. doriae , the spermathecal ampulla is round, however, the spermathecal duct is slender whereas in others it is short. In addition, P. dinagatensis has longer caeca (xxvii-xxiii) compared to that of P. abiadai , P. nagaensis , P. viracensis and P. doriae , which extend from xxvii-xxiv or -xxv. No other species in the P. urceolata group closely resemble P. dinagatensis .

Pheretima enormis n. sp. ( Fig. 2C , Table 2 ) urn:lsid:zoobank.org:act:F80F164F-2EFA-47E8-BA33-DC811AD8CBC8

Material examined: Holotype: adult ( NMA 4587 ), municipality of San Isidro , Davao Occidental Province , Mt. Hamiguitan (06°44'03"N, 126°12'15"E), 1430 m asl, Mindanao Island, Philippines, coll. Nolan Aspe, J. Cantil, 7-10 May 2006 . Paratypes: one adult (NMA 4603); two adults, amputee (ZRC.ANN 0056), same collection data as for holotype.

Etymology: The species name is from the Latin ‘enormis’ (huge, enormous), referring to the large body size.

Diagnosis: Very large worm with adult dimension 305-347 mm × 13.5-16.5 mm; purplish brown dorsum, ventrum pale; equators non-pigmented, making the body appear striped; pair of spermathecal pores at intersegment 5/6; first dorsal pore in 13/14; setae in vii 45-46, setae in xx 75-80; 7 setae between male pores; penis lacking.

Description: Purplish brown dorsum, lighter ventrum; equators non-pigmented, making the body appear striped. Length 305-347 mm (n = 2 adults); diameter 13.5-15.5 mm at x, 13-16.5 mm at xx; body cylindrical in cross-section, tail

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tapering; 110-147 segments. First dorsal pore at 13/14; paired spermathecal pores at 5/6, distance between spermathecal pores 8.5 mm (0.17 circumference ventrally apart). Female pore single in xiv; openings of copulatory bursae paired in xviii, distance between openings 7.5 mm (0.14 circumference apart ventrally), 7 setae between openings. Clitellum annular, from xiv to xvi. Setae irregularly distributed around equators in some segments; 45-46 setae on vii, 75-80 setae on xx, dorsal setal gaps present, no ventral gaps. Genital markings lacking.

Septa 4/5-7/8 and 10/11-13/14 muscular, 8/9/10 lacking. Dense tufts of nephridia on anterior faces of 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 simple, originating in xxvi, extending forward to xxiii; typhlosole originating in xxvii, simple fold, 1/3 lumen diameter; intestinal wall with 38 longitudinal blood vessels. 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. Spermathecae paired, postseptal in vi, with nephridia on ducts; each spermatheca with round to ovate ampulla and bulbous, muscular duct; stalked diverticulum attached to the right face of duct of right spermatheca, and on left face of duct of left spermatheca; stalks long, convoluted, with two kinks, terminating in long, sausage-shaped receptacle, receptacle longer than the ampulla. Male sexual system holandric, testes and funnels enclosed in paired sacs in x, xi; seminal vesicles xi, xii, each with digitate dorsal lobe; vesicles of xi enclosed in testes sac; vasa deferentia slender, free from body wall on way to ental end of prostatic ducts; prostates in xvi to xix; each prostate a single, dense, racemose mass; short muscular duct from lateral margin of prostate entering center of copulatory bursa. Copulatory bursae ovate in xvii-xix, with bulge in center forming a dome; coelomic surfaces muscular, secretory diverticula lacking; roof with two pads, floor pad lacking; blocky penis present.

Remarks: Pheretima enormis n. sp. belongs to P. urceolata group of Sims and Easton (1972). While most members of P. urceolata group have body dimensions within 33-116 mm × 1.2-5.4 mm, P. enormis has an adult body size of 305-347 mm × 13-16.5 mm. Other Pheretima species with large body size include P. maculodorsalis Aspe page 9 of 33

and James 2014 (226-235 mm); P. tigris Aspe and James 2014 (230-283); P. immanis Aspe and James 2014 (365 mm); P. lago Aspe and James 2014 (223-315 mm); P. virgata James 2004 (290 mm); P. ceramensis Cognettti 1922 (140-440 mm); P. barligensis Hong and James 2011b (225-255 mm); P. callosa Gates 1937 (330); and a large earthworm that Blakemore et. al. (2007) identified as P. darnleiensis Fletcher 1887 (700 mm). Pheretima enormis differs from these species in having one pair of spermathecal pores on 5/6, whereas the others have one pair on 7/8 ( P. maculodorsalis , P. tigris , P. immanis , P. lago , P. virgata , and P. ceramensis ), four pairs on 5/6- 8/9 ( P. barligensis and P. darnleiensis ), or three pairs on 6/7-8/9 ( P. callosa ). In addition, P. enormis has the first dorsal pore at 13/14 (12/ 13 in the other species), has more setae (7) between male pores (2-4 in P. maculodorsalis , 0-4 in P. tigris , 4 in P. virgata , 5 in P. immanis , and 0-2 in P. lago ), and has a different pigmentation pattern than the other species (the dorsal intersegmental furrows have red oval dots in P. maculodorsalis , stripes in P. tigris and P. virgata , and very thick stripes in P. immanis ).

Pheretima hamiguitanensis n. sp. ( Fig. 2D , Table 2 ) urn:lsid:zoobank.org:act:37C39A3E-70FC-4075-892E-80398EC781DF

Material examined: Holotype: adult ( NMA 4588 ), municipality of San Isidro , Davao Occidental Province , Mt. Hamiguitan (06°44'03"N, 126°12'15"E), 1430 m asl, Mindanao Island, Philippines, coll. Nolan Aspe, J. Cantil, 7-10 May 2006 . Paratypes: three adults (NMA 4604), two adults, same collection data as for holotype, one adult, Brgy Baracatan, Davao City, Mt. Apo National Park (7°00'04"N, 125°21'55"E), 1524 m asl, Mindanao Island, Philippines, coll. N. Aspe, A. Solis, Dec. 11-14, 2003.

Etymology: The species is named for Mt. Hamiguitan, the type locality.

Diagnosis: Worm with adult length 81- 90 mm; dorsum brown, ventrum pale, equators non-pigmented; pair of spermathecal pores at intersegments 5/6; setae in vii 38-42; spermathecae with ovate ampulla, spermathecal diverticulum terminating in elongate receptacle that is longer than the ampulla; penis lacking.

Description: Brown dorsum, lighter ventrum, equators non-pigmented, making the body appear striped. Length 81-90 mm (n = 2 adults); diameter

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3-3.5 mm at x, 3.5-4 mm at xx; body round in cross-section, tail tapering; 108-111 segments. First dorsal pore at 12/13, paired spermathecal pores at 5/6, distance between spermathecal pores 0.7 mm (0.07 circumference apart ventrally). Female pore single in xiv; openings of copulatory bursae paired in xviii, distance between openings 1.3 mm (0.12 circumference apart ventrally), 2 setae between openings. Clitellum annular, from xiv to xvi. Setae irregularly distributed around equators in some segments, 38-42 setae on vii, 44-47 setae on xx; dorsal setal gaps present, no ventral gaps. Genital markings lacking.

Septa 4/5-7/8 membranous, 10/11-13/14 muscular, 8/9/10 lacking. Dense tufts of nephridia on anterior faces of 5/6 and 6/7; nephridia in 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 or xvii; caeca simple originating in xxvii, extending forward to xxiv; typhlosole and longitudinal blood vessels not observed due to damage. 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. Spermathecae paired, postseptal in vi, with nephridia on ducts; each spermatheca with ovate ampulla, bulbous, muscular duct, and stalked diverticulum; diverticulum attached to ectal portion of right face of duct near ampulla of right spermatheca, and on left face of duct of left spermatheca; stalks short, terminating in elongate receptacle, receptacle longer than ampulla. Male sexual system holandric, testes and funnels enclosed in paired sacs in x, xi; seminal vesicles xi, xii, each with digitate dorsal lobe; vesicles of xi enclosed in testes sac; vasa deferentia slender, free from body wall on way to ental end of prostatic ducts; prostates variably in xvii to xx, or xv to xix; each prostate a dense, racemose mass, with three lobes; muscular duct from anterior margin of prostate entering to lateral margin of copulatory bursa. Copulatory bursae ovate in xviii-xix; coelomic surface muscular, secretory diverticula lacking; roof with thick pad, floor pad lacking; penis lacking.

Remarks: Pheretima hamiguitanensis n. sp. belongs to the P. urceolata group of Sims and Easton (1972). Members of P. urceolata group with body dimensions relatively similar to P. hamiguitanensis n. sp. include P. bicolensis Hong and James 2009a (71-98 mm × 3.8-4.6 mm); P. kitangladensis James 2004 (108 mm × 3.6-3.8 mm); P. baletei James 2004 (> 78 mm

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× 4.3-4.5 mm); P. bukidnonensis James 2004 (76 mm × 3.5 mm); and P. makilingensis Hong and James 2008b (110 mm × 4- 4.2 mm). However, P. hamiguitanensis has more closely spaced spermathecal pores (0.07 circumference) than P. kitangladensis (0.28), P. baletei (0.25), P. bukidnonensis (0.38), and P. makilingensis (0.15). Its male pores are more closely spaced (0.12 circumference) than those of P. bicolensis (0.14), P. kitangladensis (0.2), P. baletei (0.25), P. bukidnonensis (0.17) but are more distantly spaced than those of P. makilingensis (0.08). It has fewer setae (2) between male pores than P. kitangladensis , P. bukidnonensis , P. makilingensis (all with 8 setae), and P. baletei (6 setae). There are more setae in the pre-clitellar area (38-47 per segment) than in the other species (29-34 in P. bicolensis , 24 in P. baletei and P. bukidnonensis , and 35 in P. makilingensis ). Pheretima hamiguitanensis also differs from P. bicolensis , P. baletei and P. makilingensis in lacking penes. In addition, the shape of the spermathecae differs from the other species (diverticulum with small oval receptacle in P. bicolensis , club-shaped receptacle in P. kitangladensis , short club-shaped receptacle in P. baletei , ovate receptacle in P. bukidnonensis , and globular to ovate receptacle in P. makilingensis ). No other species in the P. urceolata group closely resemble P. hamiguitanensis .

Pheretima lantapanensis n. sp. ( Fig. 2E , Table 2 ) urn:lsid:zoobank.org:act:8EC52E4D-0405-403B-8E80-BED7D15C1EDF

Material examined: Holotype: adult, amputee ( NMA 4589 ), Brgy Songco , municipality of Lantapan , Bukidnon Province, Mt. Kitanglad Range (8°05'47"N, 124°55'21"E), 2200 m asl, Mindanao Island, Philippines, coll. N. Aspe, 27-31 Oct. 2012 . Paratype: one adult, amputee (NMA 4605), same collection data as for holotype.

Etymology: The species is named for the municipality of Lantapan, Bukidnon Province, where this species was collected.

Diagnosis: Worm with adult diameter 2.5- 3 mm; dorsum brown, ventrum pale, equators pigmented; pair of spermathecal pores at intersegment 5/6; setae in vii 30-39; gizzard origin in ix; prostates small, xvi-xvii; penis lacking.

Description: Brown dorsum, pale ventrum, equators pigmented. Length> 58 mm; diameter 2.5 mm at x, 3 mm at xx (n = 2 adults); body circular in cross-section. First dorsal pore at

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11/12, paired spermathecal pores at 5/6, distance between spermathecal pores 3 mm (0.38 circumference apart ventrally). Female pore single in xiv; openings of copulatory bursae paired in xviii, distance between openings 2.2 mm (0.23 circumference apart ventrally), 8 setae between openings. Clitellum annular, from xiv to xvi. Setae irregularly distributed around equator in some segments; 30-39 setae on vii, 31-36 setae on xx, dorsal setal gaps present, ventral setal gaps lacking. Genital markings lacking.

Septa 4/5-7/8 and 10/11-13/14 thin, 8/9/10 lacking. Dense tufts of nephridia on anterior faces of 5/6 and 6/7; nephridia in 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 simple, originating in xxvii, extending forward to xxiii, may be folded up or tucked under the intestine in xxiv; typhlosole originating in xxvi, simple fold slightly less than lumen diameter; intestinal wall with 33 longitudinal blood vessels. 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. Spermathecae paired, postseptal in vi, with nephridia on ducts. Each spermatheca with pear-shaped ampulla; slender, bulbous, muscular duct; single stalked diverticulum attached to ectal portion of right face of duct of right spermatheca, and on left face of duct of left spermatheca; stalks short, terminating in large, ovate receptacle. Male sexual system holandric; testes and funnels enclosed in paired sacs in x, xi; seminal vesicles xi, xii, each with digitate dorsal lobe; vesicles of xi enclosed in testes sac; vasa deferentia slender, free from body wall on way to ental end of prostatic ducts; prostates small in xvi to xvii; each prostate a dense, racemose mass with two lobes; thin duct entering from anterior margin of prostate to center of copulatory bursa. Copulatory bursae round in xvii-xix; coelomic surfaces muscular, secretory diverticula lacking; roof with two thick pads, floor pads lacking; penis lacking.

Remarks: Pheretima lantapanensis n. sp. belongs to the P. urceolata group of Sims and Easton (1972). Members of the P. urceolata group with body width similar to P. lantapanensis include P. dinagatensis , P. abiadai , P. nagaensis , and P. viracensis . Pheretima lantapanensis differs from these species in having pigmented segmental equators and in lacking penes. The distance between spermathecal pores in P.

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lantapanensis is very wide (0.38) compared to that of P. dinagatensis (0.31), P. abiadai (0.25), P. nagaensis (0.29) and P. viracensis (0.32-0.36). The new species has relatively fewer setae (30- 39) on the post-clitellar segments and the extent of the prostates is only xvi-xvii compared with that of P. dinagatensis n. sp. (43-48 and xv/xvi-xix, respectively), P. abiadai (42-47 and xvii-xix, respectively), P. nagaensis (41-48 and xvii-xx, respectively), and P. viracensis (40-45 and xvii-xix, respectively). Pheretima lantapanensis also also has large pear-shaped spermathecal ampulla with slender duct and prominent ovate diverticula, whereas the others have relatively small, round ampulla with short ducts. Although the spermathecal duct in P. dinagatensis is also slender, its diverticular receptacle is round and small. No other member of the P. urceolata group closely resembles P. lantapanensis .

Pheretima timpoongensis n. sp. ( Fig. 3A , Table 3 ) urn:lsid:zoobank.org:act:99A6DF1A-3761-4E27-A49B-22B3EA7F44A5

Material examined: Holotype: adult ( NMA 4590 ), Brgy Pandan , Mambajao City, Camiguin Province, Mt. Timpoong (9°10'45"N, 124°43'15"E), 1350 m asl, Camiguin Island, Philippines, coll. N. Aspe, J. Adeva, 11-15 Oct. 2004 . Paratypes: seven adults (NMA 4606); six adults (ZRC.ANN 0059), same collection data as for holotype.

Etymology: The species is named for Mt. Timpoong, where this species was collected.

Diagnosis: Large, purplish brown worm with adult dimensions 147-225 mm x 7-9.5 mm; ventrum pale; equators non-pigmented; pair of spermathecal pores at intersegment 7/8; large gizzard in ix-x or x-xi; intestinal origin in xv or xvi; prostates extend from xvii-xx; penes present.

Description: In living animals, purplish brown dorsum, lighter ventrum, equators non-pigmented. Length 147-225 mm (n = 14 adults); diameter 7.3-9.5 mm at x, 7-8.5 mm at xx; body circular in cross-section, tail tapering; 98-119 segments. First dorsal pore at 12/13; paired spermathecal pores at 7/8, spermathecal pores 6 mm (0.20 circumference apart ventrally). Female pore single in xiv; openings of copulatory bursae paired in xviii, distance between openings 5.4 mm (0.23 circumference apart ventrally), 2-5 setae between openings. Clitellum annular, from xiv to xvi. Setae unevenly distributed around equator in some segments; more closely spaced on ventrum; 29-43

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setae on vii, 49-73 setae on xx; dorsal setal gaps present, no ventral gaps. Genital markings lacking.

Septa 4/5-6/7 membranous, 7/8 and 10/11- 13/14 muscular, 8/9/10 lacking. Dense tufts of nephridia on anterior faces of 5/6 and 6/7; nephridia in intestinal segments located mainly on body near septum/body wall junction. Large page 12 of 33

gizzard in viii-x, esophagus with low, vertical lamellae x-xiii, intestinal origin in xv or xvi, caeca simple originating in xxvii, extending forward to xxiv or xxiii; typhlosole originating in xxvii, simple fold, 1/3 lumen diameter; intestinal wall with 31-44 longitudinal blood vessels. Hearts in x to xiii; hearts esophageal.

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Ovaries and funnels free in xiii. Spermathecae paired, postseptal in viii, with nephridia on ducts; each spermatheca with ovate ampulla, stout muscular duct, and stalked diverticulum. Diverticulum attached to the ectal portion of left face of duct of right spermatheca, and on right face of duct of left spermatheca; stalks short, with one coil, terminating in ovate receptacle. Spermathecae contain two small, ovate spermatophores. Male sexual system holandric; testes and funnels enclosed in paired sacs in x, xi;

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seminal vesicles xi, xii, each with digitate dorsal lobe; vesicles in xi enclosed in testes sac; vasa deferentia slender, free from body wall on way to ental end of prostatic ducts; prostates in xvii to xx, each a dense, racemose mass with two or three lobes; thick muscular duct from lateral margin of prostate entering posterior margin of copulatory bursa. Copulatory bursae ovate-hemispherical in xvii-xix; coelomic surfaces muscular, secretory diverticula lacking; roof with two pads, floor with folding encircling the opening; penes present;

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penis sheath lacking.

Remarks: Pheretima timpoongensis n. sp. belongs to P. sangirensis species group of Sims and Easton (1972), which differs from the P. montana Kinberg 1867 group in having spermathecal pore(s) opening only at 7/8, and no penial sheaths in the copulatory bursae. Members of the sangirensis group have a holandric male system, with paired testis sacs; the copulatory bursae are simple, and some species have short, conical penes. Blakemore (2007) acknowledged three valid subspecies in P. sangirensis : P. s. sangirensis ; P. s. crassicystis Michaelsen 1896; and P. s. chica Michaelsen 1896. Currently there are 26 species members of P. sangirensis group, including the new species described here. Pheretima timpoongensis is larger (147-225 mm × 7-9.5 mm) than P. s. sangirensis (140 mm × 3.5-4.5 mm) and P. s. chica (54-120 mm) but shorter than P. s. crassicystis (240 mm × 8 mm). The spermathecal pores in Pheretima timpoongensis (0.2 circumference) are closer than in P. sangirensis (0.25-0.28) and there are fewer setae between male pores in the new species (2-5 vs 6-10 in P. sangirensis ; particularly 10 in P. s. crassicystis). Also, there is no dorsal setal gap in P. s. crassicystis, the caeca are not simple and they have stubby pockets that are deeply divided from one another, the stalk of the spermathecal diverticulum originates entally, and the copulatory bursae are fairly flat, in contrast to that of the new species. Among other species in the P. sangirensis group, P. timpoongensis is similar to P. lago Aspe and James 2014 and P. nunezae Aspe and James 2014 from Mt. Malindang (the smaller P. lago specimens (223-315 mm × 10-11 mm) overlap in size with the larger P. timpoongensis specimens; the P. nunezae specimens were amputees but width (8.5-9 mm) are similar) in the number of setae around segments, and in having penes. However, P. lago is dark brown, has fewer setae between male pores (0-2), has closer male pores (0.15 circumference apart ventrally), has stalked diverticulum originating entally and terminating in a receptacle with 2-4 chambers, the prostates are located anterior of the copulatory bursae, and the duct form the prostate enters the anterior margin of the copulatory bursa. On the other hand, P. nunezae is dark gray-brown, has more setae (9) between the male pores, has greater distance (0.28 circumference) between the spermathecal pores, has fewer blood vessels on the intestinal wall (20-23), has smaller copulatory bursae (xviii), has stalked diverticulum originating entally and page 14 of 33

terminating in sausage-shaped receptacle, has relatively smaller prostates (xvii-xix), and the duct from the prostate enters the lateral margin of the copulatory bursa. Pheretima timpoongensis is also similar to P. apoensis n. sp. (see below) from Mt. Apo in relative size and in the number of setae around equators. However, P. timpoongensis have greater distance between male pores, the stalked diverticulum originating ectally is less stout, the duct from the prostate enters the posterior margin of the copulatory bursa, and penes are present. Pherertima timpoongensis also have some relatively overlapping features with some individuals of P. camiguinensis n. sp. (described below): the color, the number of setae on the segments, the number of setae between male pores, the position of the gizzard and the intestinal origin, the length of caeca and relative size of the prostates. However, P. timpoongensis is larger, have wider space between spermathecall pores and between male pores, and penes is present.

Pheretima camiguinensis n. sp. ( Fig. 3B , Table 3 ) urn:lsid:zoobank.org:act:1602441D-DB34-4392-9173-ED5D9A2818C4

Material examined: Holotype: adult ( NMA 4591 ), Brgy Pandan , Mambajao City, Camiguin Province, Mt. Timpoong (9°10'45"N, 124°43'15"E), 1350 m asl, Camiguin Island, Philippines, coll. N. Aspe, J. Adeva, 11-15 Oct. 2004 . Paratypes: nine adults (NMA 4607); nine adults (ZRC.ANN 0060), same collection data as for holotype.

Etymology: The species is named for Camiguin Island.

Diagnosis: Reddish brown worm with adult length 63-122 mm; pair of spermathecal pores at intersegment 7/8; spermathecal pores 0.17 circumference apart, male pores 0.15-0.18 circumference apart; 32-34 setae on vii, 39-44 setae on xx; penis lacking.

Description: In living animals, reddish brown dorsum, pale ventrum, equators non-pigmented. Length 63-122 mm (n = 19 adults); diameter 3.5- 4.8 mm at x, 3.5-5.2 mm at xx; body circular in cross-section, tail tapering; 87-103 segments. First dorsal pore at 12/13, paired spermathecal pores at 7/8, distance between spermathecal pores 1.9-2.8 mm (0.17 circumference apart ventrally). Female pore single in xiv, openings of copulatory bursae paired in xviii, distance between openings 2-2.5 mm (0.15-0.18 circumference ventrally apart), 1-4 setae between openings. Clitellum

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annular, from xiv to xvi. Setae unevenly distributed around equator in some segments; 32-34 setae on vii, 39-44 setae on xx, dorsal setal gaps present, no ventral gaps. Genital markings lacking.

Septa 5/6-7/8 and 10/11/12 thick; 4/5, 8/9, and 13/14 thin; 9/10 lacking. Dense tufts of nephridia on anterior faces of 5/6 and 6/7; nephridia in 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 originating in xxvii, extending forward to xxiv, xxiii, or xxii; caeca may be folded over, or tucked under the intestine in xxiii; typhlosole originating in xxvii, simple fold slightly less than lumen diameter; intestinal wall with 26-30 longitudinal blood vessels. Hearts in x to xiii; heart esophageal.

Ovaries and funnels free in xiii. Spermathecae paired, postseptal in viii, with nephridia on ducts; each spermatheca with ovate ampulla, stout muscular duct, and stalked diverticulum attached to ectal portion of left face of duct of right spermatheca, and on right face of duct of left spermatheca; stalks short, with one coil, terminating in ovate receptacle. Spermathecae contain two small, ovate spermatophores. Male sexual system holandric; testes and funnels enclosed in paired sacs in x, xi; seminal vesicles xi, xii, each with digitate dorsal lobe; vesicles of xi enclosed in testes sac; vasa deferentia slender, free from body wall on way to ental end of prostatic ducts; prostates in xvi to xx (or xxi in some individuals); some individuals have asymmetrical prostates (e.g. xvi to xx on left and xvi to xxi on right); each prostate a dense, racemose mass, with three to five lobes; thick muscular duct from lateral margin of prostate entering posterior margin of copulatory bursa. Copulatory bursae ovate-hemispherical in xvii-xix; coelomic surfaces muscular, secretory diverticula lacking; roof with thick pad, floor pads lacking; short, blocky penis present.

Remarks: Pheretima camiguinensis n. sp. belongs to the P. sangirensis group of Sims and Easton (1972). The new species is similar in size to P. s. chica Michaelsen 1896; P. baungonensis James 2004 ; P. quincunxia James 2004 ; P. malindangensis Aspe and James 2014 ; and P. wati Aspe and James 2014 . However, the spermathecal pores (0.17 circumference in P. camiguinensis ) are farther apart in P. s. chica (0.25-0.28) and P. baungonensis (0.32) and closer in P. quincunxia (0.13). The male pores are closer in P. camiguinensis (0.15-0.18 circumference)

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than in P. baungonensis (0.19) but farther than in P. wati (0.08). Pheretima camiguinensis has more setae per segment (32-34 in vii, 39-44 in xx) than P. quincunxia (16-20; 20-30) but fewer than P. malindangensis (43-47; 50-57) and P. wati (59-71; 52-60). The prostates in P. camiguinensis (xvii-xx/ xxi) are larger than in P. baungonensis and P. quincunxia (xvii-xix for both), but both the prostates and copulatory bursae are smaller than in P. malindangensis (xvi-xxi and xvii-xx, respectively) and P. wati (xv-xxii and xv-xxi, respectively). Pheretima camiguinensis differs from P. baungonensis in that the latter has the first dorsal pore at 13/14, more setae (8) between male pores, and shorter caeca (xxvii-xxv). Pheretima camiguinensis (reddish brown) differs in color from Pheretima s. chica (purple) and P. quincunxia (non-pigmented). No other member of the P. sangirensis group closely resembles P. camiguinensis .

Although there are variations in the size of the gizzard and prostates, and in the length of the caeca, among individuals of P. camiguinensis , molecular data for the mitochondrial 16S and CO1 genes reveal that these are intraspecific variations (0-0.013 [n = 4] and 0.002 [n = 2], respectively, in terms of K2P; Aspe and James, unpublished data). Numerous parasitic nematodes were commonly observed near the caeca in individuals of P. camiguinensis .

Pheretima sibucalensis n. sp. ( Fig. 3C , Table 3 ) urn:lsid:zoobank.org:act:8FB93CE1-0FC1-47CB-B018-2767C788F24B

Material examined: Adult, amputee (NMA 4510), Brgy Sibucal, Oroquieta City, Misamis Occidental Province, Mt. Malindang Range (8°19'31"N, 123°38'02"E), 991 m asl, Mindanao Island, Philippines, coll. N. Aspe, M. Lluch, and J. Adeva, 18-23 Feb. 2004.

Etymology: The species is named after Brgy Sibucal of Oroquieta City, where this species was collected.

Diagnosis: Dark gray-brown worm, diameter 6.5-7 mm, ventrum and equators non-pigmented; pair of spermathecal pores at 7/8; spermathecal pores and male pores, 0.21 and 0.17 circumference apart, respectively; 70 setae on vii, 89 setae on xx; 9 setae between male pores; prostate extends from xvii to xxi; penes lacking.

Description: Living animals with dorsum dark gray-brown anteriorly, fading posteriorly; ventrum non-pigmented; equators non-pigmented; clitellum

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gray, darker than adjacent segments. Length> 165 mm (n = 1 adult, amputee); diameter 6.5 mm at x; 7 mm at xx; body circular in cross-section. First dorsal pore at 12/13, paired spermathecal pores at 7/8, 0.21 circumference apart; female pore single in xiv; openings of copulatory bursae paired in xviii, 0.17 circumference apart, depressed area with 9 setae between openings. Clitellum annular, extending from xiv to xvi. Setae evenly distributed, 70 setae on vii, 89 setae on xx, dorsal gap present, ventral gap absent.

Septa 4/5/6/7/8 slightly muscular, 8/9 membranous, 9/10 absent, 10/11-13/14 slightly muscular. Dense tufts of nephridia on anterior faces of 5/6 and 6/7; nephridia of intestinal segments located mainly on body wall on anterior and posterior faces of septa, at septum/body wall junction. Large gizzard in viii-x; esophagus with lamellae from x to xiii; intestinal origin in xiv; caeca originate in xxvii, extend forward to xxii, with pocketed ventral margin; typhlosole in xxvii/xxvi, simple fold of about 1/8 lumen diameter, intestinal wall with 38 longitudinal blood vessels. Hearts in x to xiii, esophageal; commissural vessels in vi, vii, and ix lateral; viii extends to gizzard.

Ovaries and funnels free in xiii; paired spermathecae preseptal in vii, with nephridia on ducts; each spermatheca with round ampulla; short, bulbous, muscular duct; stalked diverticulum attached to the ental portion of left face of duct near ampulla of the right spermatheca, and on left face of duct of left spermatheca; stalks short, terminating in bean-shaped receptacle. Each spermatheca contains two spermatophores. Male sexual system holandric; testes and funnels enclosed in paired sacs in x and xi; seminal vesicles in xi and xii, each with short, spherical dorsal lobe; vasa deferentia slender, free from body wall, passing around lateral face of copulatory bursae en route to ental end of prostatic ducts; each prostate racemose from xvii to xxi; muscular duct from lateral margin of prostate entering posterior margin of copulatory bursa. Ductlets of lobes meet vasa deferentia at common junction with muscular prostatic duct. Copulatory bursae ovate in xvii-xx; coelomic surfaces muscular, secretory diverticula lacking; roof with thick pad, floor pad lacking; penes lacking.

Remarks: Pheretima sibucalensis n. sp. belongs to the P. sangirensis group of Sims and Easton (1972). It differs from P. sangirensis in having more setae on segmental equators (70- 89 vs. 40-60+), closer spermathecal pores, the intestinal origin in xiv rather than xv, larger page 16 of 33

prostates, and no penes. Pheretima sibucalensis is similar to P. nunezae ( Aspe and James 2014) from Mt. Malindang, in size, coloration, presence of dorsal gaps and absence of ventral gaps, arrangement of the septa, origin of the gizzard, and the length of the caeca. However, P. sibucalensis has shorter distance between male pores and between spermathecal pores (0.21 and 0.17, respectively) compared to that of P. nunezae (0.28 and 0.22, respectively). The setae on the segmental equators in P. sibucalensis is more numerous (70 in vii and 89 in xx, respectively) than that of P. nunezae (46 in vii, 51 in xx, respectively). The new species also differs from P. nunezae in the intestinal origin (xiv vs. xv), in the number of blood vessels on the intestinal wall (38 vs. 28-33), in the extent of caeca (xxvii-xxii vs. xxvii-xxiv), in the extent of the prostate and copulatory bursae (xvii-xxi vs. xvii-xix and xvii-xx vs. xviii), and in lacking penes. Pheretima timpoongensis n. sp., which also closely resembles P. nunezae , differs from P. sibucalensis in having larger body diameter, fewer setae on each segment, in having a wider space between male pores, in having fewer setae between the male pores, and in lacking penes. Pheretima sibucalensis does not closely resemble any other members of the P. sangirensis group.

Pheretima apoensis n. sp. ( Fig. 3D , Table 3 ) urn:lsid:zoobank.org:act:A7D459BD-D3BC-4F29-B330-39AC7E257887

Material examined: Holotype: adult ( NMA 4592 ), Brgy Baracatan , Davao City , Mt. Apo National Park (7°00'04"N, 125°21'55"E), 1524 m asl, Mindanao Island, Philippines, coll. N. Aspe, A. Solis, Dec. 11-14, 2003 . Paratypes: seven adult (NMA 4608), five adults same collection data as for holotype, two adults, 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; three adults (ZRC.ANN 0061), same collection data as for holotype; two adults, same collection data as for the ones collected in Mt. Musuan.

Etymology: The species is named after Mt. Apo, where this species was first collected.

Diagnosis: Large worm, adult body dimension 163-350 mm × 7.2-12 mm; 102-120 segments; dorsum purplish brown, ventrum pale; equators non-pigmented; spermathecal pores lacking in athecate individuals, at 7/ 8 in thecate individuals; 37-48 setae on vii, 50-58 setae on xx; 4-5 setae

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between male pores; male pores 0.15-0.16 circumference apart; penis lacking.

Description: In living animals, purplish brown dorsum, lighter ventrum, equators non-pigmented making the worm appear striped. Length 163- 350 mm (n = 13 adults); diameter 7.2-11 mm at x, 8-12 mm at xx; body cylindrical in cross-section, tail tapering; 102-120 segments. First dorsal pore 12/13, spermathecal pores lacking in some individuals, thecate individuals with paired spermathecal pores at 7/8, distance between spermathecal pores 0.16 circumference apart ventrally. Female pore single in xiv, openings of copulatory bursae paired in xviii, distance between openings 0.15-0.16 circumference apart ventrally, 4-5 setae between openings. Clitellum annular, from xiv to xvi. Setae unevenly distributed around equators in some segments; 37-48 setae on vii, 50- 58 setae on xx, dorsal setal gaps present, ventral setal gaps lacking. Genital markings lacking.

Septa 4/5-8/9 membranous, 11/12-13/14 thick, 9/10/11 lacking. Dense tufts of nephridia on anterior faces of 5/6 and 6/7; nephridia of intestinal segments located mainly on body near septum/body wall junction. Large gizzard in ix-xi, esophagus with low vertical lamellae x-xiii, intestinal origin in xv, caeca simple originating in xxvii, extending forward to xxvi or xxiv; typhlosole originating in xxvii, simple fold, 1/3 lumen diameter, intestinal wall with 32-38 longitudinal blood vessels. 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. Spermathecae lacking in some individuals, thecate individuals with a pair of spermathecae postseptal in viii, with nephridia on ducts; each spermatheca with round ampulla, short, stout muscular duct, stalked diverticulum attached to the ental portion of the left face of duct of the right spermatheca, and on right face of duct of left spermatheca, stalks short, terminating in small, round receptacle. Male sexual system holandric, testes and funnels enclosed in paired sacs in x, xi; seminal vesicles xi, xii, each with digitate dorsal lobe; vesicles of xi enclosed in testes sac; vasa deferentia slender, free from body wall en route to ental end of prostatic ducts; prostates in xv or xvii to xix, each a single, dense, racemose mass with 3 lobes; short muscular duct entering middle area of the surface of copulatory bursa. Copulatory bursae ovate-hemispheric in xvii-xix; coelomic surfaces muscular, secretory diverticula lacking; roof with thick pad, floor pads lacking; penis lacking.

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Remarks: Pheretima apoensis n. sp., belonging to the P. sangirensis group of Sims and Easton (1972), is a large worm similar to P. ceramensis Cognetti 1922 from Indonesia, P. lago Aspe and James 2014 from Mt. Malindang, and P. timpoongensis n. sp. from Camiguin Island, in relative size and in having pigmentation over the entire dorsal area that gradually fade toward the ventrum. However, P. apoensis differs from these species in lacking penes. It has closer male pores than in P. ceramensis (0.2 circumference apart) and in P. timpoongensis (0.23 circumference apart), has closer spermathecal pores (for thecate individuals) than in P. lago and in P. timpoongensis (0.18-0.24 and 0.2 circumference apart, respectively), and has fewer setae than in P. ceramensis (60 per segment). Its prostates also are in different position compared to that of P. lago (positioned in xiv–xviii, anterior of the copulatory bursae) and its stalked diverticulum has different structure (stalked diverticulum in P. lago terminating in a receptacle with 2-4 chambers). Although P. virgata James 2004 from Mt. Kitanglad, and P. tigris Aspe and James, 2014 , P. maculodorsalis Aspe and James 2014 , and P. immanis Aspe and James 2014 from Mt. Malindang are also similar to P. apoensis in size, these species significantly differ from the new species in pigmentation pattern from P. virgata and P. tigris (with dorsal intersegmental stripes in P. virgata and P. tigris ; with mid-dorsal intersegmental oval dots in P. maculodorsalis ; and with very thick dorsal intersegmental stripes in P. immanis ).

The P. apoensis that were collected from Mt. Apo (nine adults; 163-188 × 7.2-9 mm) are smaller compared to the ones in Bukidnon Province (four adults; 324-350 × 10-12 mm) and all individuals collected from Mt. Musuan are athecate. Also, the caeca from P. apoensis collected from Mt. Apo are longer by two segments and the prostates are more extensive (xv-xix vs. xvii-xix). The P. apoensis from Mt. Musuan and the P. apoensis from Mt. Apo have a genetic divergence of 0.011 (or 1.1%) based on mitochondrial 16S gene using K2P model, which indicates that the two are conspecific. On another note, a study on cryptic species of hormogastrid earthworms showed an upper limit of 4.27% intraspecific divergence in Hormogaster elisae using K2P based on 16S ( Novo et al. 2010). It is most likely that geographic isolation (e.g. the distance between Mt. Apo and Mt. Musuan is approximately 110 km) may have caused divergence in some morphological features such as the body size and the loss of spermathecae,

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in this case, the P. apoensis from Mt. Musuan. A study on the reproductive traits of Aporrectodea trapezoides showed that isolation of an earthworm in a controlled environment can trigger the earthworm to reproduce parthenogenetically ( Férnandez et al. 2010). Pheretima apoensis was also observed in Mt. Kitanglad Range, which is close to Mt. Musuan.

Figures

Fig. 2. Schematic dorsal views of the internal morphology of the Pheretima urceolata group species described here. (A) P. acia n. sp.; (B) P. dinagatensis n. sp.; (C) P. enormis n. sp.; (D) P. hamiguitanensis n. sp., with the intestinal origin in xv and prostates in xvii-xx; (E) P. lantapanensis n. sp. Abbreviations: s, spermatheca; h, heart; p, prostate gland; cb, copulatory bursa; c, caecum. Scale bars: A, B, D, E, 5 mm; C, 15 mm.

Fig. 3. Schematic dorsal views of the internal morphology of other Pheretima species described here. (A) P. timpoongensis n. sp., with intestinal origin in xv, and caeca extending from xxvii-xxiv; (B) P. camiguinensis n. sp., with prostates in xvii-xxi, and caeca extending from xxvii-xxi; (C) P. sibucalensis n. sp.; (D) P. apoensis n. sp. with spermathecae; (E) P. (Paraph.) pandanensis n. sp.; (F) P. (Paraph.) boaensis n. sp., with intestinal origin in xiv, prostates in xvii-xix, and caeca extending from xxvii-xxiii. Abbreviations: s, spermatheca; h, heart; p, prostate gland; cb, copulatory bursa; c, caecum. Scale bars: A-F, 5 mm.

Tables

Table 2. Comparison of species in the Pheretima urceolata group

Characters	P. urceolata Horst, 1893 *	P. acia n. sp.	P. dinagatensis n. sp.
Length	70-80	33-47	65
Width x, xx	4, 4	1.2, 1	2.5, 3
Dorsal pigmentation	?	brown	Reddish brown stripes
Segment equator	?	pigmented	unpigmented
Segments	92-104	87-94	127
1st dorsal pore	12/13	12/13	11/12
Setal gaps D, V	+, -	-, -	-, -
Sperm. pore distance	0.45	0.21	0.31
Male pore distance	0.1	0.24	0.24
Setae between openings	?	3	12-14
Setae vii, xx	20, 42	26-35, 29-38	39, 43-48
Gizzard	viii-?	viii-x	viii-x
Intestinal origin	xvi	xv	xiv/xv
Caeca	xxvii-xxiii/xxi	xxvii-xxvi	xxvii-xxiii
Prostate	xvii-xviii or xviii	xvii-xviii	xv/xvi-xix
Copulatory bursae	?	xvii-xix	xvii-xix
Penes	+	+	+

Table 3. Comparison of species in the Pheretima sangirensis group and Pheretima (Parapheretima) species

Characters	P. sangirensis * ( Michaelsen, 1891)	P. timpoongensis n. sp.	P. camiguinensis n. sp.
Length	54-240	147-225	63-122
Width x, xx	4-8	7.3-9.5, 7-8.5	3.5-4.8, 3.5-5.2
Dorsal pigmentation	Variation of purple	Purplish brown	Reddish brown
Segment equators	?	unpigmented	unpigmented
Segments	?	98-119	87-103
1st dorsal pore	?	12/13	12/13
Spermathecal pores	Paired at 7/8	Paired at 7/8	Paired at 7/8
Sperm. pore distance	0.25-0.28	0.2	0.17
Male pore distance	0.17-0.25	0.23	0.15-0.18
Setae between openings	6-10	2-5	1-4
Setae vii, xx	40; 60+	29-43, 49-73	32-34, 39-44
Gizzard	viii-?	ix-x/ ix-xi	viii-x/ ix-xi/ ix-x
Intestinal origin	xv	xv/ xvi	xv
Caeca	xxvii-?	xxvii-xxiv/ xxiii	xxvii-xxiv/ xxiii/ xxii
Prostate	xvii-xix	xvii-xx	xvii-xx/ xxi
Copulatory bursae	present	xvii-xix	xvii-xix
Secretory diverticula	-	-	-
Penes	+	+	-