Acanthonyx petiverii " H. Milne Edwards 1834

New species of Pheretima, Amynthas, Polypheretima, and Pithemera (Clitellata: Megascolecidae) from Mindanao and Associated Islands, Philippines Nonillon M. Aspe1,* and Samuel W. James2 1Department of Natural History Sciences, Graduate School of Science, Hokkaido University, N10 W8, Sapporo 060-0810, Japan. E-mail: [email protected] 2Department of Biology, University of Iowa, Iowa City 52242-1324, Iowa, USA. E-mail: [email protected] (Received January 2, 2015; Accepted October 13, 2015) Nonillon M. Aspe and Samuel W. James (2016) Studies over the past decade have shown the Philippine native earthworm fauna to be highly diverse, with apparent high endemism among islands and localities. Here we describe 17 new species of pheretimoid earthworms collected from priority conservation sites on Mindanao and associated islands. Nine species of the subgenus Pheretima (Pheretima), distinguished by the absence of secretory diverticula on the coelomic surface of the copulatory bursae, were detected. Pheretima acia n. sp., P. dinagatensis n. sp., P. enormis n. sp., P. hamiguitanensis n. sp., P. lantapanensis n. sp. belong to the P. urceolata species group, with a pair of spermathecal pores in 5/6; P. timpoongensis n. sp., P. camiguinensis n. sp., P. sibucalensis n. sp., and P. apoensis n. sp. belong to the P. sangirensis group, with a pair of spermathecal pores in 7/8. Pheretima (Parapheretima) pandanensis n. sp. and P. (Parapheretima) boaensis n. sp., members of the subgenus Pheretima (Parapheretima), with secretory diverticula on the coelomic surface of the copulatory bursae, are the first Philippine records of Parapheretima. In the genus Amynthas, A. dinagatensis n. sp. is athecate; A. cagdianaoensis n. sp. belongs to the belongs to the A. supuensis species group, with a pair of spermathecal pores at 8/9; and A. talaandigensis n. sp. belongs to A. rimosus group, with intrasegmental spermathecae in 6, 7, 8, and 9. In the genus Polypheretima, P. bukidnonensis n. sp. belongs to the Po. elongata species group, with paired genital markings on xix and successive segments, and paired spermathecal batteries in vi and/or vii; Po. zamboangensis n. sp., either has spermathecal pores in 5/6/7 or lacks them, and has a metandric male sexual system. Pithemera nolani n. sp. is unique among congeners in having only one pair of spermathecal pores in 5/6 and in having copulatory bursae. The geological history of Mindanao, which is a collision complex between three arc systems, may have contributed to the high diversity of the earthworm fauna on Mindanao. Future molecular studies can help elucidate the evolutionary relationships and geographical distribution of species and populations, and shed light on mechanisms of earthworm diversification in the Philippines. Key words: Pheretima, Amynthas, Polypheretima, Pithemera, New species, Oligochaeta, Mindanao, Philippines. *Correspondance: E-mail: [email protected] BACKGROUND Mindanao is the southernmost major island group in the Philippines, consisting of Mindanao Island and small surrounding islands. It is surrounded by four seas: the Sulu Sea to the west, the Philippine Sea to the east, the Celebes Sea to the south, and the Mindanao Sea to the north. With a land area of 104,630 km2, Mindanao is the second largest island in the Philippines next to Luzon. The island is mountainous, and is home to Mount Apo, the highest mountain in the country. Mindanao initially consisted of separate islands, but these merged in the Miocene due to closing of the Eurasian and Australian plates, which affected minor plates in between (Hall 1996). Partly due its Zoological Studies 55: 8 (2016) doi:10.6620/ZS.2016.55-08 1 complex geological history, Mindanao has very high species diversity (Heaney and Rabor 1982; Heaney et al. 2006; Heaney and Tabaranza 2006; Welton et al. 2010; James 2004; Aspe and James 2014). However, as in other parts of the Philippines, the biodiversity on Mindanao is under severe threat due to habitat alteration, overexploitation of natural resource, urban development, and increasing human population. Specific threats include mining and logging operations, and land conversion for industry, agriculture, and urban development. Overall, less than six percent of the original forest of the Philippines now remains, and 491 species of animals are listed as globally threatened on the 2004 IUCN Red List of Threatened Species (CI et al. 2006). Adequate knowledge of the natural resources is crucial to enacting effective laws and designing conservation management strategies. Prior to 2004, knowledge on Philippine earthworms was very limited. Taxonomic studies over the past decade, however, have resulted in the identification of around 200 species of native earthworms representing eight genera in the Philippines (James 2004, 2005a, 2006, 2009; James et al. 2004; Hong and James 2004, 2008ac, 2009a, 2010, 2011a, b; Aspe and James 2014, 2015). These indicate high diversity and high local endemicity of earthworms in the Philippines. Most of the Philippine earthworm species detected in the last decade described were from Luzon and associated islands, with only around 20% from Mindanao, where James (2004) described 18 species from Mt. Kitanglad, and Aspe and James (2014, 2015) described 22 species from Mt. Malindang. In this study, we describe 17 new earthworm species in the genera Pheretima (Pheretima), Pheretima (Parapheretima), Amynthas, Polypheretima, and Pithemera, from various localities on Mindanao, Camiguin, and Dinagat Islands. MATERIALS AND METHODS Collecting localities Collection sites were chosen based primarily on the Key Conservation Sites identified by the Haribon Foundation (Mallari et al. 2001). These sites roughly correspond to those identified as Priority Sites for Conservation in the Philippines, which are actually or potentially managed for conservation (CI et al. 2006). Collection was conducted intermittently from December 2003 to December 2012. The collection sites (Fig. 1) include Mt. Timpoong on Camiguin Island, approximately 10 km north of the Misamis Peninsula (11-15 October 2004); Cagdianao Municipality on Dinagat Island northeast of Mindanao (23-26 April 2004); the Mt. Kitanglad Range and Mt. Musuan in Bukidnon Province in central Mindanao (26-31 October 2012); the Mt. Apo National Park; the Mt. Hamiguitan Range Wildlife Sanctuary in Davao Occidental Province in southeastern Mindanao (7-10 May 2006); the Mt. Malindang Range in Misamis Occidental Province at the base of the Zamboanga Peninsula (18-23 February 2004); and the Mt. Timolan Protected Landscape in Zamboanga del Sur Province (1820 December 2012). Since most of the collection sites are in areas protected by the Protected Areas and Wildlife Bureau (PAWB) of the Department of Environment and Natural Resources (DENR), Prior Informed Consent certificates were secured from the Protected Area Management Board for these sites before collecting was conducted. Sampling Collecting was done from soil in primary and secondary forests at the high elevations away from human settlements or trails to increase the likelihood that the earthworms collected are native species. Other habitats including tree bark, ferns, mosses, and the insides of rotten logs were also checked for earthworms. Upon collection, earthworms were rinsed in tap water, killed in 10% ethanol, and placed in Saranex sealable plastic bags filled with a volume of 10% formalin that was at least three times the total volume of the earthworms. After two days, the formalin was replaced with 80% ethanol. Elevations were read by GPS (Magellan Map 410; Luzon map datum) if a satellite signal was detectable, or with an altimeter if not. Elevation is expressed in this paper as meters above sea level (m asl). Examination and descriptions Worms were sorted in the field to putative species, using body size, coloration, and number and location of spermathecal pores as identifying characters. External and internal characters were examined later in the laboratory for a representative subset of specimens of each species. All descriptions are based on external examination and on dorsal dissection under a stereomicroscope, following the terminology and page 2 of 33Zoological Studies 55: 8 (2016) conventions of Easton (1979). Descriptions of body color are based on living specimens. Body dimensions refer to fixed material. The degree of separation between pores is expressed as a proportion of the circumference of the worm; for example, ‘spermathecal pores 0.21 circumference apart ventrally’ means the distance between the pores is 0.21 the circumference of the worm at that point, with the circumference calculated as π times segment diameter. The generic diagnoses and assignment to species groups follow Sims and Easton (1972). While the species described share many character states diagnostic for the genus, we include shared characters to facilitate information retrieval from the separate descriptions. For convenience in species comparison, we initially compared the new species with the other members of the same species group by their sizes and by the spermathecal pores spacing and the male pores spacing. However, we made sure that the combinations of all the morphological characters were used to examine and compare all the members of the species group. Line drawings were prepared with Adobe Illustrator ver. CS5. Holotypes and paratypes are deposited in the National Museum of the Philippines Annelid Collection (NMA), P. Burgos St., Manila, Philippines. Other paratypes are deposited in the Annelid Collection of the Zoological Reference Collection (ZRC. ANN) of the Lee Kong Chian Natural History Museum (formerly Raffles Museum of Biodiversity Research), Faculty of Science, National University of Singapore, Singapore. RESULTS We detected 17 new earthworm species on Mindanao and associated islands. Table 1 lists the species found, and the total number of species currently known, at each site. Nine of the 17 new species belong to the subgenus Pheretima (Pheretima) Kinberg 1867, which differs from the other subgenus, Pheretima (Parapheretima) Cognetti 1912, in lacking secretory diverticula on the coelomic surface of the copulatory bursae. Among the new Pheretima species, five (P. acia, P. dinagatensis, P. enormis, P. hamiguitanensis, Fig. 1. Map of Mindanao and associated islands. Black circles indicate collecting sites; dashed lines indicate the boundaries among three arc systems and the Zamboanga Peninsula, which coalesced between the Early Miocene and late Pliocene (20-1 Ma). N page 3 of 33Zoological Studies 55: 8 (2016) and P. lantapanensis) belong to the P. urceolata group of Sims and Easton (1972), characterized by having a pair of spermathecal pores on intersegments 5/6; three (P. timpoongensis, P. camiguinensis, P. sibucalensis and P. apoensis) belong to the P. sangirensis group of Sims and Easton (1972), characterized by having a pair of spermathecal pores on intersegments 7/8. Two new species, P. (Parapheretima) pandanensis and P. (Paraph.) boaensis, are members of the subgenus Pheretima (Parapheretima), characterized by having secretory diverticula on the coelomic surface of the copulatory bursae. Three new species are in Amynthas, characterized by lacking nephridia on the spermathecal ducts and lacking copulatory bursae. These include A. dinagatensis, which is athecate; A. cagdianaoensis, which belongs to the A. supuensis group of Sims and Easton (1972), characterized by having a pair of spermathecal pores at 8/9; and A. talaandigensis, which belongs to the A. rimosus group of Sims and Easton (1972), characterized by having intrasegmental spermathecae in 6, 7, 8, and 9. Two species are in Polypheretima: Po. bukidnonensis n. sp., which belongs to the Po. elongata group, has a pair of genital markings in xix, successive segments in line with the male pores, and paired batteries of up to 28 spermathecae in vi and/or vii; Po. zamboangensis either has spermathecal pores at intersegments 5/6/7 or lacks spermathecal pores, has 220+ segments, and has a proandric male sexual system. Finally, one new species is in Pithemera. Pithemera nolani is unique among congeners in having only a pair of spermathecal pores in 5/6 and in possessing copulatory bursae, and it is by far the largest species of Pithemera in the Philippines. TAXONOMY Family Megascolecidae Rosa 1891 Genus Pheretima Kinberg 1867 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 Table 1. The list of new species collected in each site Locality Current total no. of earthworm species New species Mt. Timpoong, Camiguin Island 3 Pheretima (Pheretima) timpoongensis n. sp. Pheretima (Pheretima) camiguinensis n. sp. Pheretima (Parapheretima) pandanensis n. sp. Cagdianao, Dinagat Island 5 Pheretima (Pheretima) acia n. sp. Pheretima (Pheretima) dinagatensis n. sp. Pheretima (Parapheretima) boaensis n. sp. Amynthas dinagatensis n. sp. Amynthas cagdianaoensis n. sp. Mt. Kitanglad Range, Bukidnon 21 Pheretima (Pheretima) lantapanensis n. sp. Amynthas talaandigensis n. sp. Pheretima (Pheretima) apoensis n. sp.* Mt. Apo 2Pheretima (Pheretima) apoensis n. sp.* Pheretima (Pheretima) hamigitanensis n. sp.* Mt. Musuan, Bukidnon 2 Pheretima (Pheretima) apoensis n. sp.* Polypheretima bukidnonensis n. sp. Mt. Hamiguitan Range, Davao Occidental 3 Pheretima (Pheretima) enormis n. sp. Pheretima (Pheretima) hamigitanensis n. sp.* Pithemera nolani n. sp. Mt. Malindang Range, Misamis Occidental 23 Pheretima (Pheretima) sibucalensis n. sp. Mt. Timolan, Zamboanga del Sur 1 Polypheretima zamboangensis n. sp. *indicates that the species is found in more than one locality. page 4 of 33Zoological Studies 55: 8 (2016) 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-AFAFC8A746D1AA28 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 crosssection, 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 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 page 5 of 33Zoological Studies 55: 8 (2016) (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 (0.180.2). 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 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. 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. page 6 of 33Zoological Studies 55: 8 (2016) Pheretima dinagatensis n. sp. (Fig. 2B, Table 2) urn:lsid:zoobank.org:act:1ABBFFA9-4720-444B-919FC0260B43F486 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 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 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 + + + Characters P. enormis n. sp. P. hamiguitanensis n. sp. P. lantapanensis n. sp. Length 305-347 81-90 > 58 Width x, xx 13.5-15.5, 13-16.5 3-3.5, 3.5-4 2.5, 3 Dorsal pigmentation Purplish brown brown brown Segment equator unpigmented unpigmented pigmented Segments 110-147 108-111 ? 1st dorsal pore 13/14 12/13 11/12 Setal gaps D, V +, - +, - +, - Sperm. pore distance 0.17 0.07 0.38 Male pore distance 0.14 0.12 0.23 Setae between openings 7 2 8 Setae vii, xx 45-46, 75-80 38-42, 44-47 30-39, 31-36 Gizzard viii-x viii-x viii-x Intestinal origin xv xv/ xvii xv Caeca xxvi-xxiii xxvii-xxiv xxvii-xxiii Prostate xvi-xix xvii-xx/ xv-xix xvi-xvii Copulatory bursae xvii-xix xvii-xix xvii-xix Penes + - - *based from type examined by Gates (1961). page 7 of 33Zoological Studies 55: 8 (2016) 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 3133 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. 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 xxviixxiv 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-BA33DC811AD8CBC8 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 nonpigmented, 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 page 8 of 33Zoological Studies 55: 8 (2016) 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 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 5/68/9 (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-4075-892E80398EC781DF 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 8190 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 page 9 of 33Zoological Studies 55: 8 (2016) 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 (7089 vs. 40-60+), closer spermathecal pores, the intestinal origin in xiv rather than xv, larger 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-B33039AC7E257887 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 page 16 of 33Zoological Studies 55: 8 (2016) 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 163350 mm (n = 13 adults); diameter 7.2-11 mm at x, 8-12 mm at xx; body cylindrical in crosssection, 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, 5058 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 ixxi, 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. 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, page 17 of 33Zoological Studies 55: 8 (2016) 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. Subgenus Parapheretima Cognetti, 1912 Type species: Pheretima aberrans Cognetti 1911. Diagnosis: Pheretima species with a secretory diverticulum discharging into each copulatory bursa. Pheretima (Parapheretima) pandanensis n. sp. (Fig. 3E, Table 3) urn:lsid:zoobank.org:act:6B9AB7F7-FEF2-43F3-89C312503816176D Material examined: Holotype: adult (NMA 4593), 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: five adults (NMA 4609); four adults (ZRC.ANN 0062), same collection data as for holotype. Etymology: The species is named for Brgy Pandan of Mambajao City, Camiguin Province, where this species was collected. Diagnosis: Dark reddish brown worm, adult length 85-150 mm; ventrum pale; pair of spermathecal pores at intersegment 7/8; male pores and spermathecal pores 0.23 and 0.23-0.25 circumference apart; 50-55 setae on xx; 4-9 setae between male pores; single secretory diverticulum projecting from lateral surface of each copulatory bursa; long, tapering penis. Description: In living animals, dark reddish brown dorsum, lighter ventrum, equators nonpigmented. Length 86-150 mm (n = 10 adults); diameter 4-5 mm at x, 4-5 mm at xx; body circular in cross-section, tail tapering; 85-104 segments. First dorsal pore at 12/13, paired spermathecal pores at 7/8, distance between spermathecal pores 3 mm (0.23 circumference apart ventrally), spermathecal pores inconspicuous in some individuals. Female pore single in xiv, openings of copulatory bursae paired in xviii, distance between openings 3-4 mm (0.23-0.25 circumference apart ventrally), 4-9 setae between openings. Clitellum annular, from xiv to xvi. Setae unevenly distributed around equators; 36-44 setae on vii, 50-55 setae on xx, dorsal setal gaps present, no ventral gaps. Genital markings lacking. Septa 6/7/8 and 10/11-12/13 muscular, 4/5/6 and 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 xiv; caeca simple, originating in xxvii, extending forward to xxv or to xxiv, margins serrated; typhlosole originating in xxvii, simple fold slightly less than lumen diameter, intestinal wall with 31-53 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 viii, with nephridia on ducts. Each spermatheca with round ampulla, very large, short, bulbous, muscular duct; single stalked diverticulum attached to ectal portion of left face of duct close to the ampulla of the right spermatheca, and on right face of duct of left spermatheca; stalks terminating in elongate receptacle, becoming bulbous abruptly midway toward duct. Spermathecae contained two small, ovate spermatophores, lacking in some individuals. 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; prostate glands in two parts, in xvii-xviii, and in xix-xx, each part dense, racemose mass with two or three lobes; long duct from each prostate joins others to form thick, coiling, muscular duct connecting to copulatory bursa. Each copulatory bursa hemispheric in xvii-xix; coelomic surface muscular, with single secretory diverticulum projecting from lateral surface; roof with two pads, floor with thin pads; long, tapering penis arises from padded roof. Remarks: Pheretima (Parapheretima) pandanensis n. sp. and P. (Paraph.) boaensis n. sp. (see next description) are the first P. (Paraph.) species reported from the Philippines. Pheretima (Paraph.) pandanensis n. sp. differs from other P. (Paraph.) species in the location of the spermathecae, which are paired in intersegment 7/8. Other P. (Paraph.) species are either monothecate (members of the P. (Paraph.) barbara Cognetti 1913 species group), or if bithecate, have the spermathecae located at intersegment 6/7 [P. (Paraph.) bernhardi Gates 1948 or members of page 18 of 33Zoological Studies 55: 8 (2016) the P. (Paraph.) beauforti Cognetti 1913 species group] or at intersegments 5/6/7 [P. (Paraph.) saba Sims and Easton 1972]. Pheretima (Paraph.) pandanensis is similar to P. camiguinensis n. sp. in size and coloration, but differs from the latter in having a wider distance between spermathecal pores and between male pores, in having more setae between the male pores, and in having more post-clitellar setae on the segmental equators (Table 3). Pheretima (Parapheretima) boaensis n. sp. (Fig. 3F, Table 3) urn:lsid:zoobank.org:act:7FC61F12-455E-4A4B-B6BFA67E884E7C35 Material examined: Holotype: adult (NMA 4594), 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 (NMA 4610); three adults (ZRC.ANN 0063), same collection data as for holotype. Etymology: The species is named after Brgy Boa ion the municipality of Cagdianao, Dinagat Province, where this species was collected. Diagnosis: Reddish brown worm, adult length 94-133 mm; ventrum pale; single spermathecal pore midventrally at intersegment 5/6; single muscular secretory diverticulum projecting from anterior margin of each copulatory bursa; 6-9 setae between male pores; penis lacking. Description: Reddish brown dorsum, lighter ventrum, equators pigmented. Length 94-133 mm (n = 6 adults); diameter 3.5 mm at x, 4-4.5 mm at xx; body circular in cross-section, tail tapering; 94146 segments. First dorsal pore at 12/13, single midventral spermathecal pore at 5/6. Female pore single in xiv, openings of copulatory bursae paired in xviii, distance between openings 2.3 mm (0.16 circumference apart ventrally), 6-9 setae between openings. Clitellum annular, from xiv to xvi. Setae unevenly distributed around equators in some segments; 41-46 setae on vii, 57-59 setae on xx, dorsal setal gaps present, no ventral gaps. Genital markings lacking. Septa 4/5/6 thin, 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 xiv or xv, caeca simple originating in xxvii, extending forward to xxiii. One individual with caeca folded upward in xxiii, extending about length of one segment. Typhlosole originating in xxvi, simple fold, 1/3 lumen diameter; intestinal wall with 35-39 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. Single midventral spermatheca positioned on right side of the body, postseptal in vi, with nephridia on duct. Spermatheca with irregularly round ampulla; bulbous, muscular duct; two stalked diverticula attached ectally to duct, terminating in beanshaped receptacle, stalk short. 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 testis sac; vasa deferentia slender, free from body wall on way to ental end of prostatic ducts; prostates in xvii to xix (one individual has the prostates in xvi to xviii); each prostate a dense, racemose mass with three lobes; muscular duct entering anterior margin of copulatory bursa. Each copulatory bursa round in xviii; coelomic surface muscular, with single, muscular, banana-shaped secretory diverticulum projecting from the anterior margin; roof with thick pad, floor pads lacking; short, blocky penis present. Remarks: Having a single midventral spermatheca indicates that Pheretima (Parapheretima) boaensis n. sp. belongs to the P. barbara group of Sims and Easton (1972). However, this species group is characterized by having a midventral spermatheca in 6/7 and 7/8. Sims and Easton (1972) assigned to this species group P. (Paraph.) barbara barbigua [replacement name for P. (Paraph.) barbara ambigua Cognetti 1913 by Blakemore (2004)]; P. (Paraph.) barbara barbara Cognetti 1913; and P. (Paraph.) hellwigiana Cognetti 1913. The new species differs from these species in having a single midvental spermatheca in 5/6. The shape of the secretory diverticula also differs; in the other species, the diverticula are more elongate, form kinks, and project from the posterior end of the copulatory bursae. Pheretima (Parapheretima) boaensis differs from P. (Paraph.) pandanensis n. sp. in the number and location of the spermathecae, distance between male pores, and projection of the secretory diverticula from the anterior end of the copulatory bursae. page 19 of 33Zoological Studies 55: 8 (2016) Genus Amynthas Kinberg 1867 Type species: Amynthas aeruginosus 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 and superficial, opening on xviii. Spermathecal pores small or large, usually paired (bithecate) but occasionally numerous (polythecate) or single (monothecate). Spermathecal pores positioned either intersegmental or intrasegmental between 4/5 and 8/9. Clitellum annular, covering three segments (xiv to xvi). Single female pore midventrally on xiv. Genital markings present or absent. If genital markings present, variable in number and forming complex pattern on segments near male pores. Esophageal gizzard usually originating in viii; esophageal bursae lacking; pair of caeca originating in xxvii, extending forward. Ovaries and funnels free in xiii. Male sexual system holandric, metandric, or proandric. Spermathecae a single pair, multiple pairs, sometimes single and located midventrally, or sometimes lacking. Nephridia on spermathecal duct lacking. One pair of prostate glands, racemose. Copulatory bursae lacking. Amynthas dinagatensis n. sp. (Fig. 4A, Table 4) urn:lsid:zoobank.org:act:3151BD49-87AF-40F3-B22B5112CBCD992D Material examined: Holotype: preclitellate (NMA 4595), 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 preclitellate individuals (NMA 4611); three preclitellate individuals (ZRC.ANN 0064), same collection data as for holotype. Etymology: The species is named for Dinagat Island. Diagnosis: Purplish brown worm, body dimension 240 mm x 7-9 mm (longest preclitellate specimen among samples); ventrum pale; equators non-pigmented, making the body appear striped; spermathecal pores and spermathecae lacking; 20 setae between male pores; intestinal origin in xiii; caeca long, from xxvii to xxi; prostate glands small, rudimentary, confined to xviii. Description: Purplish brown dorsum, lighter ventrum, equators non-pigmented, making the body appear striped. Large preclitellates with length 223-240 mm (n = 6 individuals); diameter 6.5-9 mm at x, 6-7 mm at xx; body circular in cross-section, tail tapering; 163-167 segments. First dorsal pore at 12/13, spermathecal pore Fig. 4. Schematic dorsal views of the internal morphology of the three Amynthas species described here. (A) A. dinagatensis n. sp.; (B) A. cagdianaoensis n. sp.; (C) A. talaandigensis n. sp. Scale bars: A-C: 5 mm. (D) Schematic ventral view of polythecal A. talaandigensis n. sp., showing the intersegmental spermathecal pores (sp). Abbreviations: s, spermatheca; h, heart; p, prostate gland; c, caecum; sp, spermathecal pores. page 20 of 33Zoological Studies 55: 8 (2016) absent. Female pore single in xiv, openings of male pores paired in xviii, distance between pores 6 mm (0.27 circumference apart ventrally), 20 setae between pores. Setae irregularly distributed around equators in some segments; 35-48 setae on vii, 70-73 setae on xx, dorsal setal gaps present, no ventral gaps. Genital markings lacking. Septa 5/6/7 membranous, 7/8 and 10/1112/13 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-xii, intestinal origin in xiii, caeca simple originating in xxvii, extending forward to xxi; typhlosole originating in xxvi, simple fold, about 1/5 lumen diameter, intestinal wall with 43 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 lacking. Male sexual system holandric, testes and funnels enclosed in paired sacs in x, xi; seminal vesicles xi, xii, each with short spherical dorsal lobe; vesicles of xi enclosed in testes sac; vasa deferentia slender, free from body wall on way to ental end of prostatic ducts; racemose prostates small, rudimentary, confined to xviii. Remarks: The main distinguishing characters that separate Amynthas from Pheretima are the absence of nephridia on the spermathecal ducts and the absence of copulatory bursae in Amynthas (Sims and Easton 1972). Despite having no spermathecae to determine if nephridia is present on the spermathecal ducts or not, we assign A. dinagatensis n. sp. in Amynthas on the basis of the absence of copulatory bursae. Even at preclitellate stage, the spermathecae and the copulatory bursae in thecate Pheretima species should already be evident. Prior to the last decade, seven Amynthas species had been reported from the Philippines, all from Luzon Island: A. albobrunneus Beddard 1912; A. americanorum Beddard 1912; A. orientalis Beddard 1912; A. pauaiensis Beddard 1912; A. sodalis Beddard 1912; A. robustus Vaillant 1889; and A. corticis Kinberg 1867. However, due to the lack of information on the location of the caeca on Beddard’s (1912) five Amynthas species, and because the types of these species could not be located (Reynolds and Cook 1976), Hong and James (2004) proposed that placement of these species in Amynthas be considered provisional. Furthermore, Beddard’s (1912) Amynthas species have share some features with Philippine Pithemera (James et al. 2004), leading Hong and James (2004) to suspect that the four species might actually be Pithemera. Since all the A. dinagatensis specimens collected were preclitellate, we provisionally assign the new species to the A. illotus group of Sims and Easton (1972), which includes Amynthas Table 4. Comparison of Amynthas, Polypheretima, and Pithemera species Characters A. dinagatensis n. sp. A. cagdianaoensis n. sp. A. talaandigensis n. sp. P. bukidnonensis n. sp. P. zamboangensis n. sp. P. nolani n. sp. Length 240 >52 85-110 131 223-306 143-159 Width x, xx 9, 7 2.6, 3 3.2-4.5, 3.7-4.5 5-6.5, 5.5-6.5 3.5-4, 4-5 3.5, 3.5 Dorsal pigmentation Dark brown brown brown brown unpigmented Reddish brown Segment equator unpigmented pigmented unpigmented pigmented unpigmented unpigmented Segments 163-167 ? 85-105 147 227-248 100-114 1st dorsal pore 12/13 11/12 13/14 12/13 12/13 11/12 Setal gaps D, V +, - +, - -, - +, - +, + +, + Spermathecal pores - Paired at 8/9 Paired at 6,7,8,9 Batteries at 5/6/7 -, paired at 5/6/7 Paired at 5/6 Sperm. pore distance - 0.12 0.13 - inconspicuous 0.19 Male pore distance 0.27 0.11 0.2 0.22 0.31 0.20 Setae between openings 20 3 12-13 6-7 8-9 9-11 Setae vii, xx 35-48, 70-73 40, 39 32-37, 40-45 39-45, 46-58 60-84, 69-70 34-36, 56-59 Genital markings - - - Paired in xix-xxi Paired in xix-xxii - Gizzard viii-x viii-x viii-x viii-x x-xi viii-x Intestinal origin xiii xv xvi xv xvii xiv Caeca xxvii-xxi xxvii-xxiii xxvii-xxiv - - xxiii-xxi Prostate xviii xv-xix xviii xvi/xvii-xix xvi-xix xvi-xx/ xv-xix Copulatory bursae - - - - - xvii-xix page 21 of 33Zoological Studies 55: 8 (2016) species without spermathecae. Tsai et al. (2002) acknowledged five species in this group: A. hohuanmontis Tsai et al. 2002; A. illotus Gates 1932; A. assacceus Chen 1935; A. sheni Chen 1935; and A. oyuensis Ohfuchi 1937. Blakemore (2007) eliminated this species group, with species assigned to it either considered have doubtful identity (given the status species inquirenda) and in need of further investigation, or synonymized with other species. Amynthas dinagatensis, despite having no clitellate specimen, differs markedly from other Philippine Amynthas in its relatively large size (240 mm × 79 mm; adult body dimensions of other species range from 3248 mm × 2.7-3.5 mm (A. mindoroensis Hong and James, 2004) to 84-111 mm × 4.6-5.8 mm (A. heaneyi James 2004); in having numerous setae (20) between male pores (6-9 in A. isarogensis Hong and James 2004; 4-7 in A. malinaoensis Hong and James 2004; 2-6 in A. heaneyi James 2004; and 0 in A. philippinensis Hong and James 2004, A. mindoroensis and A. halconensis Hong and James 2004); in having the intestinal origin in xiii (in xv, xvi, or xvii in the others); and in having longer caeca (xxvii-xxi) than the others (xxvii-xxv, xxiv, or xxiii). Although A. dinagatensis is similar in size to A. americanorum (200 mm × 8 mm) and A. robustus (113-244 mm × 5.6-7.5 mm), in contrast, the new species appears to have stripes because the segmental equators don’t have pigmentation, whereas the equators are pigmented in A. robustus while A. americanorum has pale brown coloration. The new species also lacks genital markings around the spermathecal pores, while the latter two have, and the genital markings are especially numerous in A. americanorum. The intestinal origin of A. dinagatensis is in xiii while it is in xiv or xv in A. robustus and A. americanorum. Also, the caeca is long, covering seven segments, extending from xvii-xxi, while it only covers three segments in the latter two. In addition, the prostate glands are small, rudimentary, and confined to xviii in A. dinagatensis, but these are larger in A. americanorum (xvi-xxii) and A. robustus (xvi-xx or xxi). Amynthas cagdianaoensis n. sp. (Fig. 4B, Table 4) urn:lsid:zoobank.org:act:BBE85AA1-A978-4733-BBCE8E5DF196C183 Material examined: Holotype: adult, amputee (NMA 4596), 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. Etymology: The species is named for the municipality of Cagdianao, Dinagat Province, where this species was collected. Diagnosis: Brown worm, body width 2.63 mm; closely paired spermathecal pores at intersegment 8/9; first dorsal pore 11/12; spermathecal pores and male pores 0.12 and 0.11, respectively; 3 setae between male pores; genital markings lacking; proandric male sexual system. Description: Brown dorsum, pale ventrum, equators pigmented. Length > 52 mm (n = 1, adult amputee); diameter 2.6 mm at x, 3 mm at xx; body circular in cross-section. First dorsal pore at 11/12, paired spermathecal pores closely paired at 8/9, distance between spermathecal pores 1 mm (0.12 circumference apart ventrally). Female pore single in xiv, openings of male pores paired in xviii, distance between pores 1 mm (0.11 circumference apart ventrally), 3 setae between pores. Clitellum annular, from xiv to xvi. Setae irregularly distributed around equators in some segments; 40 setae on vii, 39 setae on xx, dorsal setal gaps present, ventral 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 of intestinal segments located mainly on body near septum/body wall junction. Large gizzard in ix-x, esophagus with low vertical lamellae x-xiii, intestinal origin in xv; caeca simple, originating in xxvii, folding upward in xxv, extending for two segments; typhlosole originating in xxvi, simple fold, about 1/4 lumen diameter, intestinal wall with 24 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 ix, no nephridia on ducts. Each spermatheca with pyriform ampulla; short, slender, 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 long and convoluted, terminating in sausage-shaped receptacle. Male sexual system proandric, testes and funnels enclosed in paired sacs in x; seminal vesicles xi, each with short spherical dorsal lobe; vasa deferentia slender, free from body wall on way to ental end of prostatic ducts; prostates in xv to xix; each prostate a dense, racemose mass with four lobes; stout U-shaped muscular duct from lateral margin of prostate widens toward body wall, then page 22 of 33Zoological Studies 55: 8 (2016) narrows slightly just before body wall. Remarks: Amynthas cagdianaoensis n. sp. belongs to the A. supuensis group of Sims and Easton (1972), characterized by having a pair of spermathecal pores at 8/9. Four species are assigned to this group: A. antefixus Gates 1935 and A. supuensis Michaelsen 1896 (both described from Halmahera, Indonesia), and A. dinghumontis Zhang et. al, 2006 and A. liaoi Zhang et. al, 2006 (both described from Guandong, China). The new species is smaller than A. antefixus (85-120 mm × 3.5-5 mm) and A. supuensis (115 mm × 6 mm) and lacks genital markings, whereas A. antefixus has the genital markings at the midventral line on iii, iv and v, and A. supuensis has them around the spermathecal and male pores. The first dorsal pore is at 11/12 in A. cagdianaoensis, but at 12/13 in A. antefixus and A. supuensis. Also, the male pores in A. antefixus are distantly spaced and located towards the lateral margins, while in A. cagdianaoensis, the male pores are much closer, (1 mm, 0.11 circumference apart ventrally). Moreover, the spermathecal diverticulum in A. antefixus has short stalk while in the new species it is long and convoluted. Amynthas cagdianaoensis is larger than A. dinghumontis (13-60 mm x 0.62 mm) and A. liaoi (55 mm x 1.1-2 mm), and although A. dinghumontis lacks genital markings like A. cagdianaoensis, the spermathecal and male pores of the former are more widely spaced (0.25 and 0.17-0.25, respectively). Amynthas liaoi differs from A. cagdianaoensis in having genital markings around the spermathecal and male pores, and more setae (8) between male pores. No Philippine congener closely resembles A. cagdianaoensis n. sp. Amynthas talaandigensis n. sp. (Fig. 4C, 4D, Table 4) urn:lsid:zoobank.org:act:D9356AF7-E305-43E9-9D7FFEDCA398816F Material examined: Holotype: adult (NMA 4597), 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. Paratypes: one adult (NMA 4613); two adults (ZRC. ANN 0066), same collection data as for holotype. Etymology: The species is named for the Talaandig indigenous tribe of Mt. Kitanglad. Diagnosis: Brown worm, adult length 85110 mm; pair of spermathecal pores on small protrusions at intrasegments anterior of setal equators of 6, 7, 8, 9; first dorsal pore at 13/14; genital markings lacking; intestinal origin in xvi, prostate glands small, confined to xviii. Description: Brown dorsum, pale ventrum, equators non-pigmented. Length 85-110 mm (n = 4 adults); diameter 3.2-4.5 mm at x, 3.7-4.5 mm at xx; body circular in cross-section, tail tapering; 85-105 segments. First dorsal pore at 13/14, paired spermathecal pores on small protrusions at intrasegments anterior of setal equators of 6, 7, 8, 9; spermathecal pores inconspicuous on 6; distance between spermathecal pores 1.8 mm (0.13 circumference apart ventrally). Female pore single in xiv, openings of male pores paired in xviii, distance between pores 2.8 mm (0.20 circumference apart ventrally), 12-13 setae between pores. Clitellum annular, from xiv to xvi. Setae irregularly distributed around equators in some segments; 32-37 setae on vii, 40-45 setae on xx, dorsal and ventral setal gaps present. Genital markings lacking. Septa 4/5-6/7 and 12/13/14 thin, 7/8 and 10/11/12 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 xvi; caeca simple, originating in xxvii, extending forward to xxiv; typhlosole originating in xxvi, simple fold slightly less than lumen diameter, intestinal wall with 2740 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, vii, viii and ix, nephridia on ducts lacking. Each spermatheca with pyriform ampulla; long, slender, muscular duct; single stalked diverticulum attached to the ectal portion of right face of duct of right spermatheca, and on left face of duct of left spermatheca; stalk long, terminating in ovate receptacle. Male sexual system holandric, testes and funnels enclosed in paired sacs in x, xi; seminal vesicles xi, xii, each with short, spherical 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 very small, attached to wall in xviii; stout U-shaped muscular duct widens towards body wall, then narrows slightly just before body wall. Remarks: Amynthas talaandigensis n. sp. belongs to the A. rimosus group of Sims and Easton (1972), which is characterized by having page 23 of 33Zoological Studies 55: 8 (2016) intrasegmental spermathecae in 6, 7, 8, and 9. Sims and Easton assigned to this group A. rimosus rimosus (Gates 1931) and A. rimosus effeminatus (Gates 1932), both described from Myanmar. However, Blakemore (2007) suggested that the latter subspecies is probably a synonym of the nominal subspecies. The new species differs from A. r. rimosus in the location of the first dorsal pore (at 13/14, compared to 12/13 in A. r. rimosus), the absence of genital markings or distinctly demarcated porophores around the male pores (Gates in 1931 described A. r. rimosus to have a pair of thickly crescent-shaped genital markings in xviii, but in 1972, he described the species to have distinctly demarcated porophores around the male pores instead of genital markings), the origin of the intestine (in xvi, compared to xv in A. r. rimosus), and the size of the prostate glands (in xviii, compared to covering five segments from xv-xix in A. r. rimosus). Another species with ventral intrasegmental spermathecal pores is A. monstriferus (Kobayashi 1936) from Korea. This species is much larger (235 mm) than A. talaandigensis, has genital markings on viii, and has manicate intestinal caeca. Hong and James (2009b) included A. ani, also from Korea, in this species group; Amynthas ani is larger (164-171 mm × 7.8 mm) than A. talaandigensis, has more segments and more setae around segments vii and xx (125-130, 65 and 68, respectively), and has longer prostate glands (xvii-xx). Amynthas talaandigensis is the second Amynthas species described from Mt. Kitanglad Range, following Amynthas heaneyi (James 2004). 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; 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:A39D1ED9-69B3-4E63-9A90F7A6D258AA6C 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 3.84.5 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 10/1113/14 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 page 24 of 33Zoological Studies 55: 8 (2016) 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 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 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. page 25 of 33Zoological Studies 55: 8 (2016) Mindanao, Philippines. Fieldiana Zool 112:1-63. Heaney LR, Tabaranza BR. 2006. Mammal and land bird studies on Camiguin Island, Philippines: Background and conservation priorities. Fieldiana Zool 106:1-13. Heaney LR, Rabor D. 1982. Mammals of Dinagat and Siargao Islands, Philippines. Occas Pap Mus Zool Univ Mich 699:1-30. Hong Y. 2007. Some new earthworms of the genus Amynthas (Oligochaeta: Megascolecidae) with male discs from Bogildo Island, Korea. Rev Suisse Zool 114:721-728. Hong Y, James SW. 2004. New species of Amynthas Kinberg, 1867 from the Philippines (Oligochaeta: Megascolecidae). Rev Suisse Zool 111:729-741. Hong Y, James SW. 2008a. Nine new species of earthworms (Oligochaeta: Megascolecidae) of the Banaue rice terraces, Philippines. Rev Suisse Zool 115:341-354. Hong Y, James SW. 2008b. Three new earthworms of the genus Pheretima (Oligochaeta: Megascolecidae) from Mt. Makiling, Luzon Island, Philippines. Zootaxa 1695:45-52. Hong Y, James SW. 2008c. Two new earthworms of the genus Pheretima (Oligochaeta: Megascolecidae) from Mt. Isarog, Luzon Island, Philippines. J Nat Hist 42:15651571. doi:10.1080/00222930802000398. Hong Y, James SW. 2009a. New earthworms of the Pheretima urceolata species group (Oligochaeta: Megascolecidae) from southern Luzon, Philippines. Zootaxa 2059:33-45. Hong Y, James SW. 2009b. Some new Korean Megascolecoid earthworms (Oligochaeta). J Nat Hist 43:1229-1256. Hong Y, James SW. 2010. Six new earthworms of the genus Pheretima (Oligochaeta: Megascolecidae) from BalbalanBalbalasang, Kalinga Province, the Philippines. Zool Stud 49:523-533. Hong Y, James SW. 2011a. New species of Pheretima, Pithemera, and Polypheretima (Clitellata: Megascolecidae) from Kalbaryo, Luzon Island, Philippines. Raffles Bull Zool 59:19-28. Hong Y, James SW. 2011b. New earthworm species of the genus Pheretima (Clitellata: Megascolecidae) from Mountain Province, Philippines. J Nat Hist 45:1769-1788. doi:10.1080/00222933.2011.560726. Hong Y, James SW. 2013. Three new earthworm species of the genus Amynthas (Clitellata: Megascolecidae) from Mt. Chiak National Park, Korea. Zootaxa 3646:075-081. Horst R. 1893. Descriptions of earthworms. No. 7. On Malayan earthworms. Nores Leyden Mus 15:316-329. James SW. 2004. New species of Amynthas, Pheretima, Pleionogaster (Oligochaeta: Megascolecidae) of the Mt. Kitanglad Range, Mindanao Island, Philippines. Raffles Bull Zool 52:289-313. James SW, Hong Y, Kim TH. 2004. New earthworms of Pheretima and Pithemera (Oligochaeta: Megascolecidae) from Mt. Arayat, Luzon Island, Philippines. Rev Suisse Zool 111:3-10. James SW. 2005a. New genera and species of pheretimoid earthworms (Clitellata: Megascolecidae) from southern Luzon, Philippines. Syst Biodivers 2:271-279. doi:10.1017/ S1477200004001446. James SW. 2005b. Preliminary molecular phylogeny in the Pheretima group of genera (Crassiclitellata: Megascolecidae) using Bayesian analysis. In: Pop VV, Pop AA (Eds) Advances in earthworm taxonomy II (Annelida: Oligochaeta). Cluj University Press, ClujNapoca, Romania. James SW. 2006. The earthworm genus Pleionogaster (Clitellata: Megascolecidae) in southern Luzon, Philippines. Org Divers Evol 8:1-20. James SW. 2009. Revision of the earthworm genus Archipheretima Michaelsen (Clitellata: Megascolecidae), with descriptions of new species from Luzon and Catanduanes Islands, Philippines. Org Divers Evol 9:244. e1-244.e16. Kinberg JGH. 1867. Annulata nova. Ofvers K. Vetensk Academy, Ford Stockholm, 23:97-103. Kobayashi S. 1936. Earthworms from Koryo, Korea. Sci Rep Tohoku Univ 11:140-184. Mallari N, Tabaranza, BR, Crosby M. 2001. Key Conservation Sites in the Philippines. Bookmark, Philippines. Michaelsen W. 1891. Oligochaeten des Naturhistorischen Museum in Hamburg IV. Jahrb Hamburg Wiss Anst 8:1-4. Michaelsen W. 1892. Terricolen der Berliner Zoologischen Sammlung, II. Arch Naturgesch 58:209-261. Michaelsen W. 1896. Oligochaeten. In: Kükenthal W (Ed) Ergebnisse einer zoologischen Forschungsreise in den Molukken und Borneo. Abh Senckenb Naturforsch Ges 23:192-243. Michaelsen W. 1899. Terrikolen von verschiedenen Gebieten der Erde. Mitt Naturhist Mus Hamburg 16:1-122. Michaelsen W. 1928. Die Oligochaten Borneos. Ark Zool 20:160. Michaelsen W. 1934. Oligochaten von Franzosisch-Indochina. Arch Zool Exp Gen 76:493-546. Novo M, Almodovar A, Férnandez R, Trigo D, Diaz Cosin DJ. 2010. Cryptic speciation of hormogastrid earthworms revealed by mitochondrial and nuclear data. Mol Phylogenet Evol 56:507-512. Ohfuchi S. 1937. Descriptions of three new species of the genus Pheretima from north-eastern Honshu, Japan. Res Bull Saito Ho-on Kai Mus 12:13-29. Oliveros CH, Moyle RG. 2010. Origin and diversification of Philippine bulbuls. Mol Phylogenet Evol 54:822-832. Perrier E. 1872. Recherches pour servir a l’hestoire des Lombriciens terrestres. Nouv Archs Mus Hist Nat Paris 8:5-198. Perrier E. 1875. Sur les vers de terre des iles Philippines et de la Cochinchine. CR Hebd Séanc Acad Sci Paris 81:10431046. Reynolds JW, Cook DG. 1976. Nomenclatura oligochaetologica: a catalogue of names, descriptions and type specimens of the Oligochaeta. Univ. of New Brunswick, Fredericton, Canada. Rosa D. 1891. Die exotischen Terricolen des k. k. naturhistorischen Hofmuseums. Annalen des (K. K.) naturhistorischen (Hof) Museums Wein 6:379-406. Shen HP. 2012. Three new earthworms of the genus Amynthas (Megascolecidae: Oligochaeta) from eastern Taiwan with redescription of Amynthas hongyehensis Tsai and Shen, 2010. J Nat Hist 46:2259-2283. Shen HP, Chang CH, Li CL, Chih WJ, Chen JH. 2013. Four new species of the genus Amynthas (Oligochaeta: Megascolecidae) from Kinmen, Taiwan. Zootaxa 3599:471-482. Shen HP, Chang CH, Chih WJ. 2014. Five new earthworm species of the genus Amynthas and Metaphire (Megascolecidae: Oligochaeta) from Matsu, Taiwan. J Nat Hist 48:495-522. Sims R, Easton E. 1972. A numerical revision of the earthworm genus Pheretima with the recognition of new genera and an appendix on the earthworms collected by the Royal page 32 of 33Zoological Studies 55: 8 (2016) Society North Borneo Expedition. Biol J Linn Soc 4:169268. doi:10.1111/j.1095-8312.1972.tb00694.x. Sun J, Zhao Qi, Jiang J, Qiu J. 2013. New Amynthas species (Oligochaeta: Megascolecidae) from south and central Hainan Island, China and estimates of evolutionary divergence among some corticis-group species. J Nat His 47:1143-1160. Tsai CF, Shen HP, Tsai SC. 2002. A new athecate earthworm of the genus Amynthas Kinberg (Megascolecidae: Oligochaeta) from Taiwan with discussion on phylogeny and biogeography of the A. illotus species-group. J Nat Hist 36:757-765. Tsai CF, Shen HP, Tsai SC. 2010. Four new species of Amynthas earthworms (Oligochaeta: Megascolecidae) from the Central Mountain Range of Southern Taiwan. J Nat Hist 44:22-21. doi:10.1080/00222931003624788. Vaillant L. 1889. Lombriciniens, Hirudiniens, Bdellomophes, Tkrhlariens et Planariens. In Histoire naturelle des Anneles marins et d’eau douce 3:1-766. Vidal N, Azvolinsky A, Cruaud C, Hedges BS. 2008. Origin of tropical American burrowing reptiles by transatlantic rafting. Biol Letters 4:115-118. doi:10.1098/ rsbl.2007.0531. Welton LJ, Siler CD, Linkem CW, Diesmos AC, Brown RM. 2010. Philippine bent-toed geckoes of the Cyrtodactylus agusanensis complex: Multilocus phylogeny, morphological diversity, and descriptions of three new species. Herpetol Monogr 24:55-85. Zhang W, Li J, Fu S, Qiu J. 2006. Four new earthworm species belonging to Amynthas Kinberg and Metaphire Sims et Easton (Megascolecidae: Oligochaeta) from Guandong, China. Ann Zool 56:259-254. Zhao Q, Sun J, Jiang J, Qiu J. 2013. Four new species of genus Amynthas (Oligochaeta: Megascolecidae) from Hainan Island, China. J Nat Hist 47:2175-2192. page 33 of 33Zoological Studies 55: 8 (2016)