New earthworms of Pheretima and Pithemera (Oligochaeta: Megascolecidae) from Mt. Arayat, Luzon Island, Philippines

New earthworms of Pheretima and Pithemera (Oligochaeta: Megascolecidae) from Mt. Arayat, Luzon Island, Philippines. Three new species of Pheretima and two new species of Pithemera are described from Mt. Arayat: Pheretima arayatensis sp. n., Pheretima simsi sp. n., and Pheretima castilloi sp. n., Pithemera rotunda sp. n., and Pithemera philippinensis sp. n. Pheretima arayatensis sp. n., Pheretima simsi sp. n., and Pheretima castilloi sp. n. have spermathecae in segment ix, vi, and viviii, respectively. Pithemera rotunda sp. n., and Pithemera philippinensis sp. n. have spermathecae in segment vix. The former species has numerous circular genital markings, but the latter Pithemera species has no genital markings. Descriptions of the new species are provided, including illustrations of the ventral view, male pore region, and spermathecae. Keywords: Earthworms Pheretima Pithemera M

New earthworms of Pheretima and Pithemera (Oligochaeta:

Megascolecidae) from Mt Arayat, Luzon Island, Philippines

SamuelW JAMES\Yong HONG2 &TaeHeung KIM2

1 Departmentof Life Sciences,MaharishiUniversityofManagement,Fairfield,

IA52557, U.S.A

2Faculty of Biological Resources Science, College ofAgriculture, JeonbukNational University, Jeonju 561-756, RepublicofKorea

New earthworms of Pheretima and Pithemera (Oligochaeta:

Mega-scolecidae) from Mt Arayat, Luzon Island, Philippines - Three new

speciesof Pheretimaand two new speciesofPithemeraaredescribed from

Mt.Arayat: Pheretimaarayatensissp n.,Pheretimasimsisp n.,and Phere-timacastilloisp n.,Pithemerarotunda sp n.,and Pithemeraphilippinensis

sp n. Pheretima arayatensis sp n., Pheretima simsi sp n., andPheretima

castilloi sp.n.havespermathecaeinsegmentix, vi,andvi-viii,respectively

Pithemera rotunda sp n., and Pithemera philippinensis sp n. have sper-mathecaeinsegmentv-ix.Theformerspecieshasnumerouscirculargenital

markings, but the latter Pithemera species has no genital markings Descriptions ofthe newspecies are provided, includingillustrations ofthe ventralview,maleporeregion,and spermathecae

Key- words: Earthworms - Pheretima - Pithemera - Megascolecidae

-Oligochaeta- Mt.Arayat -Philippines- taxonomy

INTRODUCTION

Mt.Arayatisanisolatedvolcanicmountaininthe large plainsofcentralLuzon, north ofManila Its forested upperelevations have always been isolated from other mountain ecosystems, such as the central Cordillera to the north and west, the Caraballo Rangetothe north, andthesouthern SierraMadreto theeast.The remain-ing forestsofMt.Arayatareanimportant refugeforbiotawhosehabitatswould other-wise be destroyed inthe region Its biogeographic importance as an isolatedpeak is

nowaccentuatedby deforestation and conversion of landto intensive agriculture.As

partofa biotic survey oftheearthwormsofthe Philippines, specimenswerecollected

from Mt Arayat in March, 2001 In this paper, we report three new species of Pheretima Kinberg, 1867 and two new species ofPithemera Sims & Easton, 1972 James (2004) described 21 new species of Megascolecidae of which 18- were new

species ofthegenus Pheretima fromtheMt Kitanglad Range, Mindanao Island He

followed the systematics ofSims & Easton (1972) These authors assigned species

*Towhomcorrespondenceandreprintrequests shouldbeaddressed

E-mail:geoworm@hanmail.net

withtheintestinalcaeca(um)originatinginornearxxii tothisgenus,Pithemera.They alsodefined fourspeciesgroups within Pithemera,one ofwhichisthebicinctagroup, containingthegenerotypePithemerabicincta(Perrier, 1875).Wereport2newspecies

ofthebicinctagroupherein

Holotypesandparatypesare depositedintheNationalMuseumofAnnelidsof the Philippines (NMA). Paratypes are deposited in the Field Museum of Natural History(FMNH),and theMuseumof Natural History ofGeneva (MHNG).

DESCRIPTIONS

PheretimaarayatensisJames& Hong sp n. Figs 1A-B Material: Holotype: One clitellate (NMA 0003741): Philippines, Luzon Island,

PampangaProvince, Mt Arayat (15°12.29'N, 120°43.25'E), 410m, litter layers in forest, 14

March2001,Y.Hong &A.Castillo colls 3 paratypes: 1 clitellate(NMA0003746), 1 clitellate (FMNH 10012), 1 clitellate(MHNG34807): Samedataas forholotype.Othermaterial: Same data as for holotype, 4 clitellate, 18 aclitellate specimens; Mt Arayat (15°12.34'N, 120° 43.90'E),330 m,litter layers in forest, 1 clitellate, 1 semiclitellate, 13March2001

Etymology:The speciesisnamedforitstypelocality.

Diagnosis: Spermathecal pore in 8/9, brown dorsal pigment, genital marking

absent

Description:Browndorsalpigment Dimensions 88-141 mmby5.8-6.3 mmat

segmentx, 7.0-7.2mmatxxx, 5.1-5.5mm atclitellum, segments96-115; body cylin-dricalincross-section Setaeregularly distributedaroundsegmentalequators, number-ing 54 at vii, 87 at xx; 5-6between male pores, size, distance regular; setal formula

AA:AB:YZ:ZZ =2:1:2:4at xiii. Clitellumannularxiv-xvi; setae invisible externally First dorsal pore 12/13, spermathecal pore midventral deepin intersegmental furrow8/9,with thickened surroundingtissue,monothecate.Femaleporesinglein xiv, 0.3 mm oval shape, 0.5 mm openings of copulatory bursae paired in xviii, distance

betweenmalepores2.1 mm.Genitalmarkinglacking

Septa5/6-7/8thick, 8/9 absent,9/10thin,onlyasmallmembraneconnectedto 10/11,not tobodywall, 10/11-13/14thick,muscular Gizzard largein viii-x,intestine begins in xv, small paired lymph glands from xxviii along dorsal vessel; intestinal caecasimple, originatinginxxvii, andextendinganteriorlyto xxii,eachconsistingof

afinger-shaped sac; typhlosole low simplefold fromxxvii Heartsx-xiii esophageal;

ix lateral,rightsidelarge, leftvery reduced

Ovaries and funnels in xiii, single spermatheca in ix with nephridia on

sper-mathecalducts; spermathecawith small ampulla, ducts shorterthan ampulla, 2 diver-ticulaeach composedof5 separate chambers on single stalksor doublestalks termi-natingintwo chambers, club-shaped.Malesexualsystemholandric,testesandfunnels

inventralpaired sacs in x, xi. Seminalvesicles twopairs inxi, xii withdorsal lobes, pseudovesicles xiii, prostates inxviii, with stoutmuscularductentering centerofthe copulatory bursae without stalked glands Vas deferens muscular xiv-xviii. Pads or otherstructureslikepenisoninsideof copulatorybursae

Remarks: The species shares thebrowndorsalpigment, dimensions,setae, and monothecate condition with Pheretima ambonensis Cognetti, 1913 from Ambon,

Indonesia Pheretimaarayatensis has 2 spermathecaldiverticulaeachcomposedof5

Fig 1

A-B Pheretimaarayatensissp n.A: ventralview; B: spermathecae.C-D Pheretimasimsi sp.

n C: ventral view;D: spermathecae E-F. Pheretima castilloi sp n E: ventralview; F: sper-mathecae Scale bars=5mm(A,C,E),2mm(B), 1mm(D, F).

separate chambers onsingle stalks or doublestalks terminating intwochambers,but

P. ambonensishas a thickductandonly 2 diverticulaonsinglestalks.Also Pheretima arayatensis hasits spermathecalporein 8/9and spermathecainix,butP. ambonensis hasthespermathecalporein7/8and spermathecainvii.

PheretimasimsiJames& Hong sp n. Figs 1C-D Material: Holotype: One clitellate (NMA 0003742): Philippines, Luzon Island,

PampangaProvince, Mt.Arayat(15°12.41'N, 120°44.45'E), 1050m,litter layers in forest, 13

March2001,Y.Hong &O.Castillo colls 3paratypes: 1 clitellate(NMA0003747), 1 clitellate

(FMNH 10013), 1 clitellate(MHNG34808):Samedataas forholotype.Othermaterial: Same

data as forholotype,4clitellate,6aclitellatespecimens;Mt Arayat(15°12.34'N, 120°43.90'E),

330 m,litter layers in forest,9clitellate,2aclitellate,13March2001

Etymology: NamedforDrR.W. Sims, who made goodcontributions to

earth-wormsystematics

Diagnosis: Spermathecalporesin 5/6.Diverticulum,aslongas ampulla, cham-ber and stalk showing spermatozoal iridescence, nephridia covering duct-ampulla junction, partof ampulla

Description: Brown dorsalpigment Dimensions68-91 mmby 2.6-2.8 mmat

drical in cross-section.Setaeregularly distributedaroundsegmentalequators, number-ing 31-33 at vii, 39-47atxx;6-8betweenmalepores,size, distanceregular; setal

for-mula AA:AB:YZ:ZZ = 1.5:1:2:3 at xiii. Clitellum annular xiv-xvi; setae invisible externally

First dorsal pore 11/12, 12/13, spermathecal pores in 5/6, at lateral margin of ventrum, whitecolor,withslightlyprotuberantlips.Femaleporesinglein xiv,0.4mm

oval shape, 0.5 mmopenings of copulatory bursae paired in xviii, distance between malepores2.5 mm.Genitalmarkinglacking

Septa 5/6-8/9 thin, 9/10 absent, 10/11-12/13 thick, muscular, 13/14 thin.

Gizzard usual in viii, intestine begins in xv, small paired lymph glands fromxxviii along dorsal vessel; intestinal caeca simple, originating in xxvii, and extending anteriorlyaboutto xxvi,eachconsistingof a finger-shapedsac;typhlosolelowsimple fold from xxvii Hearts x-xiii esophageal; ix lateral, viii to gizzard, vii lateral, with branchestoanteriorgizzardwall

Ovaries and funnels in xiii, paired spermathecalin vi with nephridiaon

sper-mathecalducts;spermathecawith small ampulla, duct veryshort,diverticulum,aslong

as ampulla, chamberandstalkshowing spermatozoaliridescence,nephridiacovering duct-ampulla junction, part of ampulla Male sexual system holandric, testes and funnelsinventralpairedsacs inx, xi.Seminalvesiclestwopairs inxi, xiiwithnarrow

dorsal lobes, prostates in xvii- xix, with stout muscular duct entering center ofthe copulatory bursae without stalked glands Copulatory bursa opening flanked by

anterior,posteriorpads,malepore withinlargeconvolutedopeningonroughly conical protrusionfromcopulatorybursaroof

Remarks: PheretimasimsikeystotheurceolatagroupinSims&Easton(1972),

which is composed of two species, P urceolata Horst 1893 and P baweanensis Michaelsen, 1924 Gates (1961) synonymized these two species and P ditheca Michaelsen, 1928 Thespecies shares the lack ofsetaein the clitellum, 31 setaeper segment at vii, and spermathecal pores in 5/6 with P urceolata Pheretima simsi differsfromP. urceolatainhavingastheampullalongas thediverticulum,andin hav-ing short caeca The locality ofPheretima simsi is close to human disturbance and

activity, as was the collection site ofP. urceolata The present species is similarto

Pheretimakitangladensis (James, 2004), butP. kitangladensishasnodorsalorventral setalgaps, andhasverticallamellaeinthe esophagous Also,P. kitangladensisisquite

remotefromhumandisturbanceatahighelevationsite (2250m), butPheretimasimsi andP.urceolatawerecollectedfromlow and mid- elevationsites(330-1050m, 1100 m) P urceolata was collected from Sumatura, Kupang, Timor, Indonesia and P.

kitangladensisfromMt Kitanglad Range,MindanaoIsland, Philippines

PheretimacastilloiJames& Hong sp n. Figs 1E-F

Material: Holotype: One clitellate (NMA 0003743): Philippines, Luzon Island,

PampangaProvince, Mt Arayat(15°12.43'N, 120°44.38'E), 900 m, litter layers in forest, 13

March2001,Y.Hong &A.Castillo colls 3 paratypes: 1 clitellate(NMA0003748), 1 clitellate (FMNH 10014), 1 clitellate (MHNG 34809): Samedata as forholotype Othermaterial: Mt Arayat (15°12.41'N, 120°44.45'E), 1050m,litter layers in forest,4 clitellate, 1 aclitellate, 13

March2001

Etymology.Thespeciesisnamedafterthe type's collector,AugustoCastillo Diagnosis: Spermathecalporesin5/6-7/8, genitalmarkingabsent

Description: Brownish dorsalpigment Dimensions 59-81 mmby 3.1-3.3 mm

atsegmentx,3.2-3.3mmatxxx, 3.0-3.3mmatclitellum,segments64-81 body cylin-drical incross-section.Setaeregularly distributedaroundsegmentalequators,

number-ing26-28at vii,46-50atxx;7-12 between malepores,size,distance regular;setal

for-mula AA:AB:YZ:ZZ = 1:1.5:2:6 at xiii. Clitellum annular xiv-xvi; setae invisible externally

Firstdorsalpore 12/13,pairedspermathecal poresin 5/6-7/8,at lateral margin

of ventrum, distancebetween spermathecal pores 3.0mm. Femalepore singlein xiv, 0.4mm oval shape, 0.4 mm openings of copulatory bursae pairedin xviii, distance between malepores2.5 mm.Genital markinglacking

Septa 5/6-7/8 thin, 8/9 absent, 9/10 very thin, attached to 10/11, 10/11 thick, 11/12-13/14 thin. Gizzardusual in viii-x, intestine begins in xv, small paired lymph glandsfromxxviiialongdorsal vessel;intestinalcaeca simple,originatinginxxvii,and extending anteriorly about to xxiv, each consisting of a small finger-shaped sac; typhlosole verylowsimple fold fromxxvii Hearts x-xiiiesophageal; ix lateral, very smallviii togizzard,vii lateral,extraesophagealvesselpharynxtoventralesophageal, thentoclitellum areaxiii to xiv, xv

Ovariesandfunnelsinxiii,pairedspermathecainvi-viii, viiiwith nephridiaon spermathecalducts;spermathecawith smallampullapouch, ductshorterthanampulla, diverticulum golf club-shaped, shorter than ampulla Male sexual system holandric, testesandfunnelsinventral pairedsacs inx, xi. Seminal vesiclestwo pairs inxi, xii withdorsal lobes, prostates in xvi- xviii, with stout muscularduct entering centerof copulatory bursae without stalked glands Copulatory bursa opening flanked by

circularpadsanterior, posterior,padsbacked byglandularchambers,malepore within longslitonsideof conical protrusionfromcopulatorybursaroof

Remarks:InSims &Easton(1972)Pheretimacastilloi keystoacouplet offer-ingachoicebetween one andfourthecalsegments,correspondingto theurceolataand lejokana (genital marking present), darnleiensis (genital marking absent) species groups, respectively However, the present species has three pairs of spermathecal poresin5/6-7/8 ThereforePheretima castilloi isunlikeallknownPheretima

Pithemera rotunda James & Hongsp n. Figs2A-B

Material: Holotype: One duellate (NMA 0003744): Philippines, Luzon Island,

PampangaProvince, Mt.Arayat(15°12.41'N, 120°44.45'E), 1050 m,litter layers in forest, 13

March2001,Y.Hong &A.Castillo colls 3 paratypes: 1 clitellate(NMA0003749), 1 clitellate

(FMNH 10015), 1 clitellate(MHNG34810): Samedataas forholotype Othermaterial: Same data as for holotype, 3 clitellate, 15 aclitellate specimens; Mt Arayat (15°12.43'N, 120°44.38'E),900 m,litter layers in forest,7clitellate, 1 aclitellate, 13March2001

Etymology:The namerotundaisLatinforcircular,referringtotheshape ofthe genitalmarking

Diagnosis: Spermathecalporesin4/5-8/9,maleporesin xviii,on0.3mmsmall hardened slightlyprotuberantcircularporophore Genitalmarkingsnumerouscircular within thickened epidermalarea

FlG.2 A-B Pithemerarotundasp n.A:ventralview; B: spermathecae.C-D Pithemeraphilippinensis

sp n.C: ventralview;D: spermathecae Scalebars=5mm(C),2mm(A), 1mm(B,D)

Description:Wormunpigmented.Dimensions 31-44mm(aclitellates25-32)by 2.0-2.4mmatsegmentx,2.3-2.6mmatxxx, 1.9-2.0mmatclitellum,segments79-91; bodycylindrical incross-section Setaeregularly distributed around segmental equa-tors,numbering 49-54at vii, 48-61atxx;9between malepores,size,distanceregular; setalformulaAA: AB: YZ:ZZ = 2:1:1:3at xiii. Clitellum annularxiv-l/2xvi; xvi setae visible,dorsal poreinvisible externally

Firstdorsalpore 12/13, five pairsofspermathecal poresin4/5-8/9, very small whitespotin lateralmarginof ventrum.Femalepores pairedin xiv,0.3mmoval shape, maleporesinxviii,on0.3mmsmallhardenedslightlyprotuberantcircularporophore, pores 1.9 mm apart. Circularraised genital markings numerous;2 on rightsidexviii,

3 between male pores xviii, 3 presetal in xix, xx, xxi, xxii; genital markings within white thickened epidermalarea

Septa 5/6 thin, 6/7 thick, 7/8 thin, 8/9 absent, 9/10, 10/11 thin, 11/12, 12/13 somemuscularthickening, 13/14thin. Gizzard usualinviii-ix, intestinebeginsinxv; intestinalcaecasimple, originating inxxii,andextendinganteriorlyabouttoxxi,each

consistingofasmall triangle-shaped sac;typhlosoleabout 1/3 lumendiametersimple foldfromxxii.Heartsx-xii esophageal;ix lateral, viiiabsent, vi, vii lateral.

Ovariesandfunnelsin xiii,spermathecaeinv-ix;nonephridiaonspermathecal ducts; spermathecae with small strawberry-shaped ampulla, ducts shorter than ampulla, diverticulum club-shaped, shorterthan ampulla Male sexual system

holan-dric, testesandfunnelsinsingle large sacsin x, xi;sacsattachedtofrontsideofseptum

10/11, 11/12, sacs donotenclosehearts,sacs ofxienclose seminal vesicles Seminal

vesiclestwopairs in xi, xiiwithoutdorsal lobes, prostates multilobedinxvi- xx, ectal halfof ductexpanded,muscular

Remarks: Pithemera rotunda keys to the bicincta group in Sims & Easton

(1972), which is composed of two species, Pithemera bicincta (Perrier, 1875) and Pithemeraviolacea (Beddard, 1895).LikeP bicinctaandP. violacea, P. rotunda has spermathecal poresin4/5-8/9, andthe spermathecalampullais longer thanthe diver-ticulum Pithemerarotunda hasthree circular papillae insegmentsxviiitoxxii,butP.

bicinctahas pairedgenitalpapillaeextendingfrom segmentsxviii toxix,and1 pairof genitalpapillae justbehindthemaleporeson 18/19 Onlyonepairofgenitalpapillae

ispresentinP. violaceajustlateraltoand behindthemalepores(Beddard, 1895).

PithemeraphilippinensisJames& Hongsp n. Figs2C-D

Material: Holotype: One clitellate (NMA 0003745): Philippines, Luzon Island,

PampangaProvince,Mt Arayat(15°12.41'N, 120°44.45'E), 1050m,litter layers in forest, 13

March 2001,Y. Hong &A.Castillo colls. 2 paratypes: 1 clitellate(FMNH 10016), 1 clitellate (MHNG 34811): Same data as for holotype Other material: Same data as for holotype, 11

aclitellate specimens; Mt Arayat(15°12.43'N, 120°44.38'E), 900 m, litter layers in forest, 3

clitellate, 1 aclitellate, 13March2001

Etymology:Thespeciesis namedforits typelocality.

Diagnosis:Spermathecalporesin4/5-8/9,maleporesxviiionfoot print-shaped pads extendingto 17/18-18/19 Genitalmarkingabsent

Description:Lightbrowndorsalpigment.Dimensions 51-54mm(semiclitellate 41-48) by 2.3-2.5 mm at segment x, 2.8-3.3 mm at xxx, 2.5-2.6 mm at clitellum,

segments 78-90;body cylindrical in cross-section Setae regularly distributedaround segmentalequators,numbering 56-68 at vii, 49-61 atxx; 9between male pores, size, distance regular; setalformulaAA:AB:YZ:ZZ =3:1:1:2at xiii.Clitellumannular xiv-l/2xvi;xvisetaevisible, dorsalporeinvisible externally

Firstdorsalpore 12/13,fivepairsofspermathecal poresin4/5-8/9,small white spots, lateral.Femalepores pairedinxiv,0.4mmoval shape,maleporesxviiionfoot print-shaped pads extending to 17/18-18/19, 2.0 mm between male pores Genital marking lacking

Septa 5/6-7/8 thin, 8/9 absent, 9/10-11/12 thick, 12/13, 13/14 thin. Gizzard usual in viii-ix,intestinebeginsinxv;intestinal caeca simple,originating inxxii, and extending anteriorly about to xxi, each consisting of a small triangle-shaped sac; typhlosolesimplefoldabout1/4lumendiameterfromxxii.Heartsx-xiiesophageal;ix lateral, viii lateral.

Ovariesandfunnelsinxiii,spermathecaein v-ix;nonephridiaonspermathecal ducts; spermathecae with small pouch, diverticulum sausage-shaped, as long as ampulla Malesexual systemholandric, testes andfunnelsin ventralpaired sacs inx,

xi. Seminal vesicles twopairs in xi, xii,prostates in xvii- xix, two main lobes, ental halfof ducts slender,nonmuscular; ectal halfof ducts enlarged, musculartapered at both ends; body wall surface over external male pads covered with densely packed,

Remarks:The species alsokeystothe bicinctagroupinSims&Easton(1972), but unlike the other members, Pithemeraphilippinensis has foot print-shaped pads extending over 17/18-18/19

ACKNOWLEDGEMENTS

This studywassupportedbyNational ScienceFoundationgrantDEB-0072764.

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