PUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORY CENTRAL PARK WEST AT 79TH STREET, NEW YORK, NY 10024 Number 3634, 12 pp., 34 figures December 31, 2008 On the First Asian Spiders of the Family Caponiidae (Araneae, Haplogynae), with Notes on the African Genus Diploglena NORMAN I. PLATNICK1 AND PETER JA¨GER2 ABSTRACT Anewgenusandspecies,Laoponiasaetosa,areestablishedtocontainthefirstmembersofthe family Caponiidae to be discovered in Asia. Only two Old World genera of caponiids have been described, both from Africa; in having only two eyes, the newly collected spiders from Laos resemblesouthernAfricanDiploglena,ratherthanthetypegenusCaponia.However,membersof Diploglena (which are poorly known, and are therefore discussed and illustrated) have an endite shape and eye pattern indicating that they may be most closely related to the Chilean genus Tisentnops. Laoponia may be most closely related instead to the endemic Californian genus Calponia; similar California/East Asiadisjuncts are known in otheroldspiderlineages. INTRODUCTION and Diploglena Purcell (1904), which is known onlyfromsouthernAfrica.Itwasthereforewith The spider family Caponiidae is much more great surprise that we studied the specimens diverse,atboththegenericandspecificlevels,in reported on below, which were newly collected theNewWorldthanintheEasternHemisphere. bythesecondauthorinLaos. Ofthedozengeneracurrentlyrecognizedinthe Thepresentpaper,thesixthinaseriesonthe family,onlytwoarefoundintheOldWorld:the spiderfamilyCaponiidae,isthefourthtodeal type genus Caponia Simon (1887), which has withmembersofthe‘‘Caponiinae’’,apresum- been reported from Ethiopia to South Africa, ably basal group whose members are easily 1PeterJ.SolomonFamilyCurator,DivisionofInvertebrateZoology,AmericanMuseumofNaturalHistory;Adjunct Professor, Department of Biology, City College, City University of New York; Adjunct Professor, Department of Entomology, Cornell University; Adjunct Senior Research Scientist, Center for Environmental Research and Conservation,ColumbiaUniversity([email protected]). 2Arachnology, Research Institute Senckenberg, Senckenberganlage 25, D-60325 Frankfurt am Main, Germany ([email protected]). CopyrightEAmericanMuseumofNaturalHistory2008 ISSN0003-0082 2 AMERICAN MUSEUMNOVITATES NO. 3634 recognizedbytheabsenceofthesubsegmented Californian and East Asian members of a tarsi and other bizarre leg features that group is certainly not unprecedented among characterize the exclusively New World sub- relatively old lineages of spiders, with the family Nopinae. Although the Nopinae is families Hypochilidae and Antrodiaetidae arguably a monophyletic group (Platnick, providing obvious examples. 1995;PlatnickandLise,2007),caponiinesseem The presence of caponiids in Asia presum- to be grouped together only because they lack ably reflects the relatively basal phylogenetic theobviouslegapomorphiesofnopines. position,andconsequentage,ofthefamily.The TheLaotianspecimens(figs. 1,2)haveentire two fossil species described from Dominican tarsi and lack the other leg specializations of amber by Wunderlich (1988) are relatively nopines. The only Old World caponiid genera advanced taxa (nopines with metatarsal exten- are also not nopines. Members of the African sions),andthefamilyasawholeprobablydates typegenus,Caponia,resembleonlythoseofthe backatleasttoPangaeantimes,ifnotearlier. endemic Californian genus Calponia Platnick The known Laotian specimens were all (1993) in retaining the plesiomorphic number taken by sieving leaf litter late in the dry ofeighteyes.TheLaotianspecimens,likemost season.Thehabitatsweresmallbuthadample caponiidspecies,haveonlytwoeyes(fig. 12);in forest cover, often on terrains too steep to be additiontheyhaveashortembolusonthemale used for agricultural purposes, near streams palp (figs. 3–5) that is unlike the long and feeding into the Nam Khan River. The leaf elaborate embolus found in male Caponia (see litter layer in these forest patches is thick, averaging20–30 cmdeep.Atthetypelocality, Purcell,1904:figs.28–35). specimens of the sparassid spider genus Ourattentionthereforefocusedinsteadonthe Pseudopoda Ja¨ger were taken; because those other known Old World genus, Diploglena, spiders apparently require high humidity, despite its great geographic separation. Purcell’s these habitats are likely to remain relatively genusismuchlesswellknownthanCaponia,and humid even at the peak of the dry season. has in fact not been illustrated or restudied in Similar habitats occur in Thailand, and detailsinceitsoriginaldescriptionin1904.Only given the substantial amount of spider col- the South African type species is known, lecting done there over recent years, it seems although Lawrence (1928) added a subspecies odd that caponiids have seemingly not been from Namibia based only on ‘‘its proportions encountered there. Their absence might indi- and larger size’’. Through the courtesy of Drs. cate that the Mekong River is a significant AnsieDippenaarandCharlesGriswold,wehave barrier, and that these are highly relictual been able to study a few modern specimens of taxa, but might also be just an artifact of Diploglena.Althoughitisnotsurprisingthatthe seasonality, since the animals are seemingly LaotianspidersdonotbelongtoDiploglena,we adult in the dry season only. weresurprisedtofindcharactersthatmayplace The specimens reported on here are depos- DiploglenaclosertotheChileangenusTisentnops itedinthecollectionsoftheAmericanMuseum Platnick(1994)thantoCaponia. of Natural History (AMNH), the California Indeed,ofthepreviouslyknowngenera,the Academy of Sciences in San Francisco (CAS), closest match to the Laotian specimens in the Research Institute Senckenberg in Frank- male palpal structure are probably the eight- furt(SMF),andthePlantProtectionResearch eyed Calponia harrisonfordi Platnick, from InstituteinPretoria(PPRI). California. Although Calponia specimens dif- fer in several characters, including eye num- SYSTEMATICS ber, sternal shape, and cheliceral and male palpal setation, it is possible that a generic- Laoponia, new genus level cladistic analysis of the family could showthatCalponiarepresentsthesistergroup TYPE SPECIES: Laoponia saetosa, new spe- oftheLaotianspecies.Suchananalysiswould cies. be premature at present, as several genera ETYMOLOGY: The generic name is a con- remain to be revised, and others need to be traction of ‘‘Laotian Caponia’’, and is femi- newly described, but a disjunction between nine in gender. 2008 PLATNICKAND JA¨GER: FIRST EAST ASIAN CAPONIID 3 Figs. 1–5. Laoponia saetosa, new species, male. 1. Cephalothorax and abdomen, dorsal view. 2. Same, ventralview. 3.Left male palp, ventral view.4.Same, prolateral view.5.Same, retrolateral view. DIAGNOSIS: Members of the genus can be show irregular pale areas, within the oval separatedfromthepreviouslyknowncaponiid ocular ring of black pigment, that may genera as follows: from Caponia, Caponina represent remnants of the other eyes, there Simon, and Notnops Platnick by the presence arenocuticulartracesoflensesinthoseareas; of only two eyes (although some specimens see fig. 12) and the much shorter male 4 AMERICAN MUSEUMNOVITATES NO. 3634 embolus, from Diploglena by the normal tarsiwiththreeclaws;pairedclawswithabout (rather than anteriorly expanded) endites and 10 teeth, most distal of which are largest; theabsenceofategularapophysisonthemale unpaired claw long, without teeth, almost palp, from Calponia by the presence of only fused to protruding onychium (fig. 18). two eyes and a row of elongated macrosetae Tarsal organ exposed (fig. 19); trichobothria on the anterior face of the cheliceral paturon, present on tibiae, metatarsi, and tarsi, their from Taintnops Platnick by the shorter male bases with semicircular rim bearing only low embolus, from Tisentnops Platnick by the ridges (fig. 17). Abdomen with two pairs of normal size and position of the eyes, and respiratory spiracles clustered around epigas- from the nopine genera (Nops MacLeay, tric groove; anterior spiracles leading to Nopsides Chamberlin, Orthonops Chamberlin, several tracheoles; posterior spiracles of each Nyetnops Platnick and Lise, and Tarsonops sideleadingtothreetrachealtrunks(twolarge Chamberlin) by having entire, rather than ones extending anteriorly into cephalothorax, subsegmented,tarsi. one much narrower extending posteriorly for DESCRIPTION: Moderate-sized caponiids most of abdominal length, as in Forster and with two eyes (fig. 1); females unknown. Platnick, 1985: fig. 889) (tracheae observed in Carapace oval, pars cephalica flattened, grad- juvenile digested with pancreatin, as per ually narrowed opposite palpal coxae, pars A´lvarez-PadillaandHormiga,2008);posterior thoracicasloping;cuticlewith raised sculptur- spiracles connected by transverse duct. Male ing outlining tiny circular to hexagonal cells, spinnerets in typical caponiid arrangement strong bristle behind each eye and pair of (fig. 20); anterior laterals with single large dorsally directed strong bristles on clypeus; major ampullate gland spigot, without piri- thoracic groove short, shallow. Eyes dark, formglandspigots(fig. 21);posteriormedians separatedbylessthantheirdiameter,setback with two or three aciniform gland spigots and from anterior margin of clypeus by about single spigot with lower base presumed to twice their diameter, surrounded by oval ring serve minor ampullate gland (figs. 22, 23); of black pigment interrupted at sides and posterior laterals with four aciniform gland alongmidline.Cheliceraepaturonwithrowof spigots and two spigots with lower bases that four long, strong bristles along anteromedian may serve minor ampullate glands (figs. 24, face, bristles of each side overlapping distally; 25). Male palpal patella and tibia short, median lamina long, with heavily sclerotized unmodified; cymbium ovoid, without distinct anteromedian tip (fig. 6), about half of space dorsal pad of short setae but with thickened between lamina and base of fang occupied by setaeonpromargin(fig. 13);ventralsurfaceof white membranous lobe; lateral surface with bulb with small circular area clearly delimited stridulatory ridges (fig. 7; pick at base of fromremainderofcuticle(figs. 14–16);embo- prolateral side of palpal femur, figs. 4, 9, 10). lus narrow, bent distally at about half its Endites convergent, acuminate, not truncated length (figs. 4, 5). or expanded distally, anterior surface distally DISTRIBUTION: Known only from Luang with strong serrula consisting of single tooth Prabang Province, Laos. row(fig. 9),proximallywiththreestrongsetae originating from enlarged bases (figs. 9, 11). Laoponia saetosa, new species Labium triangular, fused to sternum; anterior surface of labrum bearing transverse rows of Figures 1–25 tiny teeth (fig. 8). Sternum oval, cuticle with raised sculpturing as on carapace; cepha- TYPE: Male holotype taken sieving leaf lothoracic membranes with weak epimeric litter at slope of stream at dusk, at an eleva- sclerites dorsal of coxae I, II, and III plus tion of 328 m, at Houay Kho, 19u44909.10N, IV;epimericscleritesnotfusedwithtriangular 102u16937.50E, Ban Pak Bak, Nam Khan, SE sclerites extending from sternalmargin to and Luang Prabang, Luang Prabang Prov., Laos betweencoxae.Legformula4123;legswithout (Mar.23,2007;P.Ja¨ger),depositedinSMF. spines; metatarsi and tarsi entire, without ETYMOLOGY: Thespecificnameisfromthe subsegmentation or membranous processes; Latin adjective saetosus (‘‘with hairs or bris- 2008 PLATNICKAND JA¨GER: FIRST EAST ASIAN CAPONIID 5 Figs. 6–11. Laoponia saetosa, new species, male. 6. Left chelicera, distal view. 7. Right chelicera, retrolateralview.8.Labrum,anteriorview.9.Leftpalpalendite,dorsalview.10.Stridulatorypickonleft palpalfemur,prolateral view. 11.Modified setae onbaseof palpalendite, dorsalview. tles’’) and refers to the stiff bristles on the formlypaleorange;mouthpartsyellow;sternum cheliceral paturon. pale orange; abdomen uniformly white, with DIAGNOSIS: With the characters of the scattered long setae. Palpal tibia and cymbium genus and palpi as in figs. 3–5. with promarginal surfaces heavily setose; bulb MALE (HOLOTYPE): Total length 2.83 mm. circular, embolus very narrrow, abruptly bent Carapace 1.29 mm long, 1.05 mm wide, uni- distallyatabouthalfitslength(figs. 3–5). 6 AMERICAN MUSEUMNOVITATES NO. 3634 2008 PLATNICKAND JA¨GER: FIRST EAST ASIAN CAPONIID 7 FEMALE: Unknown. squared tubercles on the anterior margin of OTHER MATERIAL EXAMINED: LAOS: Luang the endites (Platnick, 1994: fig. 26). Although Prabang Prov.: Houay Tham, 19u44951.20N, the two eyes of Tisentnops are differently 102u13915.40E, between Ban Khon Why and placed (anteriorly advanced rather than un- Xieng Ngeun, Nam Khan, SE Luang Prabang, usually posterior), they are greatly reduced in Mar. 24, 2007, elev. 363 m, sieving leaf litter at size (Platnick, 1994: fig. 4), a character also - rocks along stream (P. Ja¨ger, SMF), 2 , same true for Diploglena (to a slightly lesser degree, - (AMNH),1 . fig. 26).Itispossiblethatthereductionineye DISTRIBUTION: Known only from Laos; a size is a synapomorphy uniting Diploglena juvenile that probably belongs to this species withTisentnops.Diploglenaspecimensarealso was taken sieving leaf litter outside a cave, at notable for the enlarged femora, patellae, and an elevation of 373 m, at Tham Pathok, tibiaefoundonthefirstpairoflegs(and,toa 20u33908.20N, 102u37992.50E, Nong Khiao, lesser degree, the second). The femora are Nam Ou, NE Luang Prabang, Luang laterally compressed, especially proximally, Prabang Prov., Laos, Mar. 17–18, 2007 (P. but all three leg segments are more than twice Ja¨ger, F. Steinmetz, SMF). as high as the metatarsi and tarsi. DESCRIPTION: Moderate-sized caponiids with two eyes (figs. 26, 27). Carapace oval, Diploglena Purcell pars cephalica flattened, slightly narrowed opposite palpal coxae, pars thoracica slightly Diploglena Purcell, 1904: 169 (type species by monotypy sloping; cuticle with raised sculpturing outlin- DiploglenacapensisPurcell,1904). ingirregularlyrectangularcells,withscattered DIAGNOSIS: Specimens of this nonnopine long, weak setae concentrated at rear of pars genuscaneasilyberecognizedbytheshapeof cephalica and around ocular area; thoracic the palpal endites (fig. 27), which are distally groove obsolete. Eyes dark, separated by expanded and have a sinuous anterolateral about their diameter, set back from anterior margin.Amongtheotherdescribedcaponiids, margin of clypeus by about five times their members of the Mexican genus Nopsides diameter, surrounded by oval ring of black Chamberlin and the Chilean genus Tisentn- pigment. Chelicerae paturon with clump of ops have endites that are similarly expanded strong bristles along anteromedian face, bris- (see Chamberlin, 1924: fig. 39; and Platnick, tles of each side overlapping distally, clump 1994: fig. 26). Nopsides is a nopine, and its occupying almost total length of paturon; members can easily be distinguished from median lamina long, without sclerotized ante- thoseofDiploglenabythesubsegmentedtarsi, romedian tip, about two-thirds of space but they also resemble those of Diploglena in between lamina and base of fang occupied having the anterior median eyes set farther by white membranous lobe; lateral surface back on the carapace than is typical for the with stridulatory ridges (pick at base of family. In Nopsides, though, there are four prolateral side of palpal femur, fig. 29). eyes present, with the presumptive anterior Endites expanded distally to almost twice lateral pair occurring midway between the their width at base, with laterally sinuous anterior margin of the clypeus and the distal margin, serrula not apparent, seemingly anterior median eyes (Chamberlin, 1924: fig. fused with distal endite margin, median 38), whereas in Diploglena only two eyes surface of anterior face unsclerotized. occur. Specimens of Tisentnops (which is not Labium almost diamond-shaped, not fused a nopine) can easily be distinguished from to sternum; anterior surface of labrum with thoseofDiploglenabythepresenceofarowof pair of long, longitudinal, submarginal scler- r Figs.12–19. Laoponiasaetosa,newspecies,male.12.Eyes,dorsalview.13.Rightpalp,prolateralview. 14.Rightpalpalbulb,prolateral view.15.Same,ventral view.16.Same,close-up.17. Trichobothrialbase from tarsus IV, dorsal view. 18. Claws of tarsus I, oblique lateral view. 19. Tarsal organ from tarsus IV, dorsalview. 8 AMERICAN MUSEUMNOVITATES NO. 3634 Figs. 20–25. Laoponia saetosa, new species, male. 20. Spinnerets, distal view. 21. Left anterior lateral spinneret, distalview.22.Leftanterior medianspinneret, distalview.23. Rightanterior medianspinneret, distalview. 24.Posterior lateralspinnerets, distalview. 25.Right posterior lateralspinneret, distalview. otized strips. Sternum oval but with slightly sallybetweencoxaeIandII,IIandIII,andIII produced knobs at anterolateral corners, and IV; epimeric sclerites not fused with opposite lateral sides of endite bases, cuticle triangular sclerites extending from sternal with scattered depressions; cephalothoracic margin to and between coxae. Female palpal membranes with weak epimeric sclerites dor- tarsus expanded, extremely hirsute but with- 2008 PLATNICKAND JA¨GER: FIRST EAST ASIAN CAPONIID 9 Figs. 26–30. Diploglena capensis Purcell. 26. Cephalothorax and abdomen, dorsal view. 27. Same, ventralview. 28.Left male palp, ventral view.29. Same,prolateral view.30. Same,retrolateral view. 10 AMERICAN MUSEUMNOVITATES NO. 3634 - outwell-defineddorsalpadofshortsetae.Leg ton, PPRI 98/290), 1 . Western Cape: formula 4123; legs without spines; metatarsi Saldanha Bay, Nov. 18, 1949 (B. Malkin, U and tarsi entire, without subsegmentation or CAS), 1 ; Tierberg, July 25, 1989 (R. Dean, - membranous processes; tarsi withthree claws; PPRO 91/40), 1 . paired claws with about six relatively short DISTRIBUTION: Knownonlyfromthewest- teeth, unpaired claw short, without teeth, ern parts of South Africa and Namibia. almost fused to protruding onychium. Tarsal Lawrence (1928: 225) described D. capensis organ not scanned; trichobothria present on major, as a new ‘‘variety’’, based on a female tibiae, metatarsi, and tarsi, their bases not from Kaross, Namibia; collections including scanned. Abdomen with two pairs of respira- males will be needed to help determine tory spiracles clustered around epigastric whether the Namibian population is conspe- groove; internal anatomy not investigated. cific with the specimens from the cape, or Spinnerets in typical caponiid arrangement, belongs to a diagnosable taxon. with posterior medians situated almost as far forward as anterior laterals; spigots not SUPPLEMENT scanned, but females apparently with single, largecylindricalglandspigotsituatedatcenter After the manuscript for this paper was of posterior median spinneret spigot field (no accepted, the second author returned to Laos similarly enlarged spigot evident on posterior for additional fieldwork, and was able to laterals). Male palpal patella and tibia short, collectanadultfemaleofthenewspecies.This unmodified; cymbium ovoid, with distinct specimen was taken by sieving litter along a dorsal pad of short setae; embolus accompa- streambed in secondary forest with bamboo, niedbysecondtegularapophysis(figs. 28–30). banana, and mango (Laos: Luang Prabang Female epigastric area with pair of transverse Prov.: Xieng Ngeun Distr., SE Luang sclerotized strips, almost meeting at midline. Prabang, Ban Keng Koung, N side Nam DISTRIBUTION: Known only from South Khan, 19u4099630N, 102u1894420E, elev. ca. Africa and Namibia. 372 m, Feb. 25, 2008). This specimen, 2.69 mm long, was digested with pancreatin, Diploglena capensis Purcell and shows the same arrangement of the tracheal system (figs. 31, 32) as the juveniles Figures 26–30 examined earlier. The female genitalia consist of a single receptaculum, probably globose in Diploglena capensis Purcell, 1904: 170, figs. 36–38 (three life,situated onashortstalk(figs. 33,34);the male and four female syntypes from St. Helena Bay, tiny tube-shaped structure evident on the Malmesbury Div., Western Cape Province, South Africa, should be in the South African Museum, not dorsal side of the receptaculum in those examined). figures is probably just a small segment of a tracheole that broke off during preparation DIAGNOSIS: With the characters of the and adhered to the receptaculum surface. genus and genitalia as in figs. 28–30. MALE: Described by Purcell (1904). FEMALE: Described by Purcell (1904). ACKNOWLEDGMENTS OTHER MATERIAL EXAMINED: NAMIBIA: Erongo: 10 mi W Okombahe, May 10, 1958, WethankDrs.AnsieDippenaar(PPRI)and U elev. 920 m (E. Ross, R. Leech, CAS), 1 . Charles Griswold (CAS) for providing speci- SOUTH AFRICA: Northern Cape: 20 km N mens. At AMNH, Matthias Burger, Nadine Concordia, 26u409S, 22u40E, June 2, 1997, Dupe´rre´, Mohammad Shadab, Lou Sorkin, from scorpion burrow (A. Harrington, PPRI and Steve Thurston assisted with the tracheal - 98/291), 1 ; Garies, Namaqualand, Nov. 14, system examination, illustrations, scanning U 1949(B.Malkin,CAS),1 ;Henkries,June2, electron micrographs, and plates. We are 1997, in scorpion burrow (A. Harrington, grateful to Manivone Thoummabouth, Ouane - U PPRI 98/292), 1 , 2 ; Kweekfontein, June Sirisack, Kim Valakone, Edina Ifticene, 2, 1997, from burrow of scorpion, Opisth- Chearmoua Bear Khue (‘‘La Maison du ophthalmus crassimanus Purcell (A. Harring- Patrimoine’’, Heritage House, Luang Prabang),