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The Earthworm Genus Pleionogaster (Clitellata: Megascolecidae) in Southern Luzon, Philippines

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The Earthworm Genus Pleionogaster (Clitellata: Megascolecidae) in Southern Luzon, Philippines Org. Divers. Evol. 6, Electr. Suppl. 8: 1 - 20 (2006) © Gesellschaft für Biologische Systematik URL: http://www.senckenberg.de/odes/06-08.htm URN: urn:nbn:de:0028-odes0608-9 The earthworm genus Pleionogaster (Clitellata: Megascolecidae) in southern Luzon, Philippines Samuel W. James Natural History Museum and Biodiversity Research Center, Dyche Hall, 1345 Jayhawk Drive, University of Kansas, Lawrence, KS 66045, USA e-mail: [email protected] Received 2 December 2004 • Accepted 11 August 2005 Abstract An earthworm biodiversity survey of the Philippines has yielded 14 new species of the perichaetine megascolecid genus Pleionogaster, previously known from only a few species from scattered Philippine locations. Bicol, the southern peninsula of Luzon, has intact forests on several isolated volcanic peaks and other remote areas. Collections made in these forests yielded the following new species, here presented by type location: Mt. Malinao, Pleionogaster albayensis, P. bicolensis, P. castilloi, P. malinaoensis, P. tiwiensis; Mt. Isarog, P. ffitchae, P. isarogensis; Mt. Bulusan, P. bulusanensis, P. hongi, P. sorsogonensis; Catanduanes Island, P. nautsae, P. viracensis; Caramoan Peninsula, P. caramoanensis, P. nillosae. Most of the species were found only in the neighborhood of the type locality, but P. bicolensis occurs in two locations in northern Bicol. Intraspecific variation in P. castilloi was observed between northern and southern flanks of Mt. Malinao. The impor- tance of several previously overlooked Pleionogaster traits is demonstrated by their homogeneity within species reported here. Keywords: Pleionogaster; Megascolecidae; Clitellata; Philippines; Luzon; Bicol Introduction The perichaetine megascolecid genus Pleionogaster about its placement in that group. Its members have Michaelsen, 1892 has long been known from a few iso- a reduced esophageal gizzard in viii, all the anterior lated collections made in the Philippines during the last septa are present, and these are generally muscular an- two centuries (Easton 1979; James 2004), but its range terior of 9/10 or 10/11. There are paired meganephri- and diversity remained unknown. In early 2001 the au- dia in the intestinal segments, but not elsewhere; these thor and a field team set out to conduct the first organi- are connected to paired longitudinal ducts flanking the zed collecting of earthworm material from the Philip- dorsal blood vessel. In addition there are regular ranks pines, and discovered many species of Pleionogaster of micronephridia opening to corresponding rows of in all parts of the island of Luzon, including Catandu- nephridiopores, again in the intestinal segments. Mi- anes, an island united with Luzon at Pleistocene low cronephridia are present in the anterior segments but sea levels (Heaney 1985; 1993). This paper is one of are placed differently. The intestine is supplied with a series planned on new earthworm species discove- several gizzards posterior to xxiv, followed by an aty- red by the Philippines Terrestrial Annelid and Gastro- phlosolate section of intestine up to 25 segments long, pod Survey (PTAGS). Here I report the Pleionogaster at the end of which is a constriction and the typhlosolar species found in Bicol, a long southern peninsula of origin, if a typhlosole is present. The supra-esophageal Luzon,plus Catanduanes Island, composed of lowland vessel extends posteriorly past the heart segments to plains dotted with dormant and active volcanoes. xvii in most cases, and the vessels returning blood from Pleionogaster is an interesting group of earthworms, the clitellar region (posterior latero-parietals) are from from the standpoint of the systematics of the Phereti- one to four pairs in segments xiv–xvii. In some cases ma complex (sensu Sims and Easton 1972) of the Me- the posteriormost pair of these vessels is connected to gascolecidae. Several of its characteristics are unique paired longitudinal vessels extending many segments among the genera of the complex, raising questions back on the body wall. Most of these features have Org. Divers. Evol. 6, Electr. Suppl. 8 (2006) James: The earthworm genus Pleionogaster (Clitellata: Megascolecidae) in S Luzon, Philippines 2 been noted in older works on the genus, but some are molecular phylogenetic studies, and were archived at reported here for the first time. –20 ºC. All material was scored for external characters, No other genera in the Pheretima complex share any drawings made by way of a drawing tube, and inter- of the unusual features of Pleionogaster, and neither nal anatomy studied by dorsal dissection. Setal counts do any other megascolecid genera found in the regi- were made over 1 mm each of a dorsal, lateral and on comprising the Philippine and Indonesian archi- ventral segmental equator arc, averaging these counts, pelagoes and mainland Southeast Asia west through and multiplying the average by the estimated circum- Thailand. This poses a biogeographic and systematic ference of the segment. This was necessary due to the puzzle, the solving of which will require additional ex- extremely large numbers of setae (up to 300) crowded ploration. For now, the most likely location of related into the anterior setal rings. genera seems to be Australia, where Anisogaster Blake- Collecting site data include the acronym PTAGS more, 2000, Eastoniella Jamieson, 1977, Gastrodrilus (Philippine Terrestrial Annelid and Gastropod Survey) Blakemore, 2000, Hickmaniella Jamieson, 1974, Pseu- followed by a location number that uniquely identifies a docryptodrilus Jamieson, 1972, and Pseudonotoscolex location in the survey. Latitude and longitude are given Jamieson, 1971 share some characteristics with Pleio- in degrees and decimal minutes. Elevations were read nogaster (Jamieson 2000). The Indian genus Lampito from a GPS unit (Magellan Map410) if sufficient sa- Kinberg, 1866 has a similar arrangement of post-clitel- tellites were detectable, or from an altimeter. The map lar meganephridia and ducts (Gates 1972) datum used in the GPS readings was Luzon. Collectors Ecologically, members of Pleionogaster are mostly listed are those working at the respective site. endogeic, with generally long and slender, rarely pig- Holotypes of the new species are deposited in the mented bodies, and always found in the soil, never in National Museum of the Philippines Annelid Collec- litter or arboreal habitats. Some of the larger species in tion (NMA), P. Burgos St., Manila, Philippines. Ad- which the anterior ends are darkly pigmented may be ditional depositories: KUNHM = Kansas University anecic. Natural History Museum, Lawrence, Kansas USA; UPLBMNH = University of the Philippines Los Baños Methods and material Museum of Natural History, College, Laguna, Philip- On the upper slopes of the Bicol volcanoes lie the re- pines. maining primary montane and mossy forests of the area, and these were given priority in collecting. Collections Taxonomic section were also carried out in two low-elevation sectors in Clitellata: Megascolecidae Rosa, 1891 some relatively undisturbed forest on Catanduanes Is- Genus Pleionogaster Michaelsen, 1892 land and the nearby Caramoan Peninsula. Earthworms Perichaeta Beddard, 1886: 298 (in part). were collected by digging and handsorting soil, sear- Pleionogaster Michaelsen, 1892: 247; Beddard ching organic soil and root mats from the tops of boul- (1895: 433), Michaelsen (1896: 198), Easton (1979: ders, logs, roots and branches, dissecting epiphytes, 114), James (2004). primarily arboreal ferns, and searching the leaf axils of Plionogaster Michaelsen, 1900: 210; Stephenson palms and Pandanaceae. All specimens examined for (1930: 840), Stephenson (1933: 923), Gates (1943: this paper were recovered from mineral soil, not from 105), Jamieson (1971: 82). organic soil horizons, organic mats or other suspended soils, or plants. Type species. Pleionogaster jagori Michaelsen, Photographs were taken live of as many different 1892 (Easton 1979). species as could be distinguished in the field by in- Diagnosis. Perichaetine Megasolecidae with larger spection with the unaided eye. Additional photomic- numbers of setae in the head segments than in post–cli- rographs were taken in the laboratory using a Nikon tellate segments, a reduced esophageal gizzard in viii, Coolpix 995 digital camera fitted to an adapter moun- intestinal gizzards in the region xxiv–xxx, paired ente- ted on the phototube of a Leica M5 stereomicrosco- roic stomate meganephridia and regular ranks of exoic pe. Specimens were killed in 50% ethanol and fixed in micronephridia in post-clitellate segments, and a single 10% formaldehyde in the field. After at least 48 hours pair of racemose prostates whose ducts are united with fixation they were rinsed in three changes of tap water the vasa deferentia near the ental end of the duct. All with at least 12 hours between changes, and transferred known species have paired spermathecae in segments to 80% ethanol. This removes residual formaldehyde, viii and ix. reduces health hazards to those working with the speci- Remarks. James (2004) discussed synonymy issues mens, and may reduce DNA degradation. Duplicate raised by Easton (1979) and made some changes to the sets of specimens were preserved in 95% ethanol for genus definition in Easton (1979). Org. Divers. Evol. 6, Electr. Suppl. 8 (2006) James: The earthworm genus Pleionogaster (Clitellata: Megascolecidae) in S Luzon, Philippines 3 Pleionogaster tiwiensis n. sp. se attached to tubules adherent to either side of dorsal (Fig. 1A, B) vessel. Ovaries and funnels
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