Weintrauboa , a new genus of pimoid spiders from …spiders/content/publications...Hormiga (Hormiga, 2002; Hormiga, Arnedo & Gillespie, 2003). From a modern perspective, it looks as

Zoological Journal of the Linnean Society

2003

139

261ndash281 With 12 figures

copy 2003 The Linnean Society of London


2003

139

261ndash281

261

Blackwell Science Ltd

Oxford UK

ZOJZoological Journal of the Linnean Society

0024-4082The Lin-nean Society of London 2003 2003

139

2

261281Original Article

THE SPIDER GENUS WEINTRAUBOA (PIMOIDAE)G HORMIGA

E-mail hormigagwuedu

Weintrauboa

a new genus of pimoid spiders from Japan and adjacent islands with comments on the monophyly and diagnosis of the family Pimoidae and the genus

Pimoa

(Araneoidea Araneae)

GUSTAVO HORMIGA

Department of Biological Sciences George Washington University Washington DC 20052 USA

Received July 2002 accepted for publication February 2003

The spider genus

Weintrauboa

new genus (Araneae Pimoidae) is described to place two species of pimoids fromJapan and adjacent islands that were formerly classified in the linyphiid genus

Labulla

Weintrauboa contortipes

(Karsch) new comb the type species and

W chikunii

(Oi) new comb are redescribed Parsimony analysis of mor-phological characters provides robust support for the monophyly of the genus

Weintrauboa

and corroborates themonophyly of

Pimoa

Pimoidae and the clade Linyphiidae plus Pimoidae New diagnoses for

Pimoa

and Pimoidaeare provided copy 2003 The Linnean Society of London


2003

139

261ndash281

ADDITIONAL KEYWORDS cladistics ndash Linyphiidae ndash morphology ndash phylogeny ndash systematics ndash taxonomy

INTRODUCTION

lsquoLinyphioidrsquo spiders (Linyphiidae plus Pimoidae) areone of the largest lineages of the order Araneae andinclude the most speciose family-level clade of web-weaving spiders the family Linyphiidae (Platnickrsquos(2003) on-line catalogue lists 559 valid linyphiid gen-era grouping more than 4200 species) Pimoidae thesister lineage of Linyphiidae is a relictual group thatcontains a single genus and 22 described speciesUnderstanding pimoid systematics is critical for thestudy of phylogenetic relationships within theLinyphiidae because of the role pimoids can poten-tially play in character polarization by outgroup com-parison (Hormiga 1993 1994a) The discovery of asecond genus of pimoids provides a modest increase inspecies richness (from 22 to 24 species) but a dramaticincrease in the diversity of the male genital morphol-ogy Because genital characters especially those of themale are an astonishingly rich source of phylogeneticdata it seems appropriate to study in close detail themorphology of these new pimoids Although the spe-cies included in the new pimoid genus had alreadybeen described (one of them more than a century ago)

new descriptions are necessary to understand genitalhomologies and to reconstruct their phylogeneticplacement

Eugegravene Simon (1884) erected the linyphiid genus

Labulla

to accommodate the European species

Linyphia thoracica

Wider Since then

Labulla

hascontained a diverse array of linyphiid and pimoid spe-cies most of them not closely related to the type spe-cies (

L thoracica

) As a result there have beennumerous generic transfers of which the most recenthave been the two lsquo

Labulla

rsquo species described bySimon (1900) from Hawaii currently placed in thelinyphiid genus

Orsonwelles

Hormiga (Hormiga 2002Hormiga Arnedo amp Gillespie 2003) From a modernperspective it looks as if the genus

Labulla

had beenused as a dumping ground for morphologically deviantlinyphioids As presently delimited the genus

Labulla

is polyphyletic (Hormiga 1994b 2002)

Labullanepula

Tikader from India is not congeneric with thetype species Only two species of the six currently clas-sified in

Labulla

seem to be congeneric with

L thoracica

namely

L flahaulti

Simon (from France)and

L impudica

Denis (from Algeria) (Hormiga2002) Detailed examination of the two lsquo

Labulla

rsquo spe-cies from Japan and adjacent islands

L

contortipes

(Karsch) and

L insularis

(Saito) (

=

L chikunii

) shows

262

G HORMIGA



2003

139

261ndash281

that these taxa belong into the family Pimoidae Inthis paper these Japanese species will be redescribedand its phylogenetic affinities explored using a cladis-tic approach In light of this new evidence Hormigarsquos(1994a) diagnoses for Pimoidae and

Pimoa

are alsorevised

MATERIAL AND METHODS

Morphological methods are described in detail in Hor-miga (2000 2002) Taxonomic descriptions follow theformat of Hormiga (1994a 2002) Specimens wereexamined and illustrated using a Leica MZAPO ste-reoscopic microscope with a camera lucida Furtherdetails were studied using a Leica DMRM compoundmicroscope with a drawing tube Some microscopeimages were recorded using a Leica DC-200 digitalcamera and edited using the software package Auto-Montage A LEO 1430VP scanning electron micro-scope was also used to study and photograph morpho-logical features Left structures (eg palps legs etc)are depicted unless otherwise stated Most hairs andmacrosetae are usually not depicted in the final palpdrawings All morphological measurements are givenin millimetres Somatic morphology measurementswere taken using a scale reticle in the dissectingmicroscope Eye diameters are taken from the span ofthe lens The cephalothorax length and height weremeasured in lateral view and its width was taken atthe widest point Similarly the length and height ofthe abdomen were measured in lateral view and thewidth as the widest point as seen from a dorsal viewThe measurements of the abdomen are only approxi-mations because the abdomen size changes moreeasily in preserved specimens than do other moresclerotized parts (eg the chelicerae) The total lengthwas measured in lateral view and is also an approxi-mation because it involves the size of the abdomen andits relative position Approximate leg article lengthswere measured in lateral view without detaching thelegs from the animal by positioning the article beingmeasured perpendicularly The position of the meta-tarsal trichobothrium is expressed as in Denis (1949)and Locket amp Millidge (1953) Female genitalia wereexcised using surgical blades or sharpened needlesThe specimen was then transferred to methyl salicy-late (Holm 1979) for examination under the micro-scope temporarily mounted as described inCoddington (1983) Male palps examined with theSEM were first excised and transferred to a vial with70 ethanol and then cleaned ultrasonically for1ndash3 min The specimen was then transferred to abso-lute ethanol and left overnight After critical pointdrying the specimens were glued to rounded alumin-ium rivets using an acetone solution of polyvinyl resinand then AuPd coated for examination at the SEM

A

NATOMICAL

ABBREVIATIONS

USED

IN

THE

TEXT

AND

FIGURES

Male palp

BH basal haematodochaC conductorCB cymbiumCP cymbial process (CDP in Hormiga 1994a)E embolusEF embolic flapm membrane (or membranous)MA median apophysisP paracymbiumPCS pimoid cymbial scleritePEP pimoid embolic processST subtegulumT tegulum

Epigynum

CD copulatory ductCO copulatory openingFD fertilization ductS spermatheca

Somatic morphology

AC aciniform gland spigot(s)AG aggregate gland spigot(s)ALS anterior lateral spinneretAME anterior median eye(s)CY cylindrical gland spigot(s)FL flagelliform gland spigot(s)MAP major ampullate gland spigot(s)mAP minor ampullate gland spigot(s)PI piriform gland spigot(s)PLE posterior lateral eye(s)PLS posterior lateral spinneretPME posterior median eye(s)PMS posterior median spinneret

Institutional abbreviations used in text are given inthe lsquoAcknowledgementsrsquo section

C

LADISTIC

ANALYSIS

Taxa

The taxonomic sample used in the cladistic analysiswas designed to test the monophyly of

Weintrauboa

and

Pimoa

and builds on my previous phylogeneticwork on the higher level phylogenetics of lsquolinyphioidsrsquo(Hormiga 1994a b 2000 2002 Hormiga

et al

2003)The monophyly of

Pimoa

had been previously testedby Hormiga (1994a) in the present study

Pimoa

is rep-resented by four species (out of 22 described species)including the two most basal taxa in Hormigarsquos(1994a fig 442) preferred cladogram

P rupicola

THE SPIDER GENUS

WEINTRAUBOA

(PIMOIDAE)

263



2003

139

261ndash281

(Simon) and

P breuili

(Fage) The goal of this analysiswas not to reconstruct intrageneric relationships in

Pimoa

but to test the monophyly of

Weintrauboa

andto study how the inclusion of this latter genus affectsthe monophyly and diagnosis of

Pimoa

Outgrouptaxon selection follows the taxonomic sample of Hor-miga (2000) The Linyphiidae sample attempts to rep-resent morphological diversity at the subfamilial levela reduced sample of Erigoninae was used becauseidentifying erigonine intergeneric relationships is notone of the goals of this analysis (nonetheless 13 gen-era out of the 31 used by Hormiga with emphasis onthe basal erigonine lineages follow Hormigarsquos pre-ferred optimal cladogram (fig 38))

Labulla thoracica

was included in the analysis to test the conjecture thatthe Japanese taxa are only distantly related to thetype species of

Labulla

Representatives of three ara-neoid families (Tetragnathidae Theridiosomatidaeand Theridiidae) were used to root the lsquolinyphioidsrsquo

Characters

A total of 75 characters were scored (Appendices 1 and2 Figs 1ndash10) 39 male and nine female genital char-acters 24 somatic morphological characters and threebehavioural characters (see Appendix 1) These char-acters are those of Hormiga (2000) with some changesto accommodate increased representation of pimoidsand the addition of

Labulla thoracica

A total of sevencharacters are parsimony uninformative in this taxo-nomic context but were kept in the analysis becausethey are potentially useful to reconstruct lsquolinyphioidrsquorelationships

Analyses

The parsimony analyses were performed using thecomputer programs Hennig86 version 15 (Farris1988) NONA version 20 (Goloboff 1993) WinCladaversion 10008 (Nixon 1999) and Nexus Data Editor049 (Page 2001) were used to study character opti-mizations on the cladograms and to build and edit thecharacter matrix respectively Ambiguous characteroptimizations were usually resolved so as to favourreversal or secondary loss over convergence (Farrisoptimization or ACCTRAN) if not the optimizationscheme is discussed in the text The 14 multistatecharacters were treated as non-additive (unordered orFitch minimum mutation model Fitch 1971) NONA(Goloboff 1993) was used to calculate Bremer supportindices (BS lsquodecay indicesrsquo) (Bremer 1988 1995Donoghue

et al

1992)

RESULTS

Heuristic searches under lsquoamb-rsquo using 1000 replicatesof mult TBR

+

TBR (multmax) resulted in four min-

imal length trees of 172 steps with CI and RI of 056and 074 respectively (the strict consensus cladogramis presented in Fig 11) Exclusion of the seven parsi-mony uninformative characters reduces tree length to164 steps and the CI to 054 Successive characterweighting in Hennig86 which weights characters bytheir rescaled consistency index produces stableresults (eight trees) in the second iteration with astrict consensus cladogram that is topologically iden-tical to that of the consensus cladogram under equalweights Using the command lsquoamb

=

rsquo in NONA to inter-pret clade support produces eight topologies ofminimal length summarized by the same strictconsensus cladogram than results from the analysisunder lsquoamb-rsquo (ie Fig 11)

Topological conflict among the most parsimonioustopologies resides in two areas The first one is theresolution of the trichotomy

Stemonyphantes

Myno-gleninae (

Haplinis

plus

Novafroneta

) and Erigoni-nae The second involves a distal erigonine cladethat includes the genera

Drepanotylus

Sciastes

anda clade containing

Islandiana

Erigone

and

Walcke-naeria

None of these areas of conflict is particu-larly relevant for the problem at hand (pimoidhigher level systematics) and they have alreadybeen discussed in Hormiga (2000) One of the fourminimal length trees (under equal weight andlsquoamb-rsquo) has been arbitrarily selected to optimizecharacter changes and to discuss character evolu-tion (Fig 12)

DISCUSSION

Detailed examination of the Japanese

Labulla

spe-cies shows that they are not linyphiids but mem-bers of the family Pimoidae In the absence of ageneric circumscription to include these two speciesa new genus is created to place them The results ofthe cladistic analysis show that

Weintrauboa contor-tipes

and

W chikunii

are sister species and that

Weintrauboa

is the sister lineage of

Pimoa

The addi-tion of these two species to the family Pimoidaechanges the diagnosis of both Pimoidae and

Pimoa

Until now

Pimoa

was the only member of Pimoidaeand thus the diagnosis and composition of the genusand the family were identical Inclusion of

Weintrau-boa

in the family indicates that the monophyly ofPimoidae is supported by the following unambigu-ous putative synapomorphies a dorsoectal cymbialprocess presence of cymbial cuspules a retrolateralcymbial sclerite (PCS) and the embolic process(PEP) Under Farris optimization the cheliceral strid-ulatory striae are interpreted as secondarily absent(lost) in

Weintrauboa

The male genital morphology of

Weintrauboa

israther different in the details from that of

Pimoa

264

G HORMIGA



2003

139

261ndash281

Figure 1


(Karsch)

n comb

male palp AndashF from Doi Sakuto-Cho Okayama Pref JapanEndashJ from Japan (MNHN 3837) partially expanded A ventral B caudoectal C ectal D mesal E dorsal F cymbium api-cal G apical HndashJ details of tegular division Scale bars 05 mm

THE SPIDER GENUS

WEINTRAUBOA

(PIMOIDAE)

265



2003

139

261ndash281

although it has the basic elements of pimoid palpalanatomy The

Weintrauboa

palp has a large cymbialectal process (with cuspules located at the base of thecymbium not in the process itself as in the palp of

Pimoa

) a cymbial sclerite continuous with the para-cymbium (as in

Pimoa breuili

and

P rupicola) and anembolic process whose morphology is quite differentfrom the homologous structure in Pimoa The presencein Weintrauboa of a paracymbium attached to thecymbium by means of a membrane (lsquointersegmentalparacymbiumrsquo Figs 1BC 2B 5A 6A) allows for theinterpretation of this paracymbium condition as asymplesiomorphy in Linyphiidae The intersegmental

paracymbium of linyphiids has been classicallyregarded as one of the synapomorphies of the family(eg Hormiga 2000) In the sister lineage of lsquoliny-phioidsrsquo the so-called lsquoSpineless femur cladersquo (sensuGriswold et al 1998) which includes cyatholipoidsplus theridioids the paracymbium is continuous withthe cymbium without a membranous connection(lsquointegral paracymbiumrsquo) The cladistic results suggestthat the intersegmental paracymbium could haveevolved in the common ancestor of pimoids andlinyphiids and that in Pimoa the paracymbium maybe secondarily integral In summary the presence ofan intersegmental paracymbium is now interpreted as

Figure 2 Weintrauboa chikunii (Oi) n comb (AndashD) W contortipes (Karsch) n comb (EndashF) AndashF male palp fromJuzhno-Sakhalinsk Sakhalin Island Russia EF epigynum Mt Takiyama Nagi-cho Katsuta-gun Okayama Pref JapanA mesal B ectal C dorsal D cymbium apical E ventral F lateral Scale bars 05 mm (AndashD) 025 (EF)

266 G HORMIGA

copy 2003 The Linnean Society of London Zoological Journal of the Linnean Society 2003 139 261ndash281

Figure 3 Weintrauboa contortipes (Karsch) n comb (AndashH) W chikunii (Oi) n comb (IJ) A epigynum (cleared)dorsal B epigynum (cleared) ventral C epigynum (cleared) dorsal (schematic) D prosoma dorsal E prosoma ventralF prosoma anterior G male first leg lateral H male first metatarsus dorsal I male first leg lateral J male first meta-tarsus dorsal Scale bars 01 mm (AndashC) 05 (DndashF HJ) 10 (GI)

THE SPIDER GENUS WEINTRAUBOA (PIMOIDAE) 267


a synapomorphy of lsquolinyphioidsrsquo (Pimoidae plusLinyphiidae) rather than as a synapomorphy ofLinyphiidae Analysis of these new data suggests thatthe monophyly of Linyphiidae is supported by the fol-lowing six synapomorphies the typical linyphiid para-cymbium (usually U or J shaped) presence of asuprategulum absence of a median apophysis andconductor presence of a radix and presence a columnmembrane between the embolic and tegular division

Little is known about the epiandrous fusules ofpimoids and linyphiids Hormiga (2002) documentedthese male fusules in eight species of Orsonwelles anddiscussed variation in the morphology of the fusules inLinyphiidae In the few studied pimoid species thearrangement of the epiandrous fusules varies fromsingles (Pimoa altioculata (Keyserling) Fig 9CD) toclusters of 2ndash5 fusules (P breuili (Fage) Fig 9EFWeintrauboa contortipes Figs 6EF 9AB) InP breuili the epiandrous spigots are grouped but theydo not come out from a common pit as they do in Wein-trauboa contortipes (eg Fig 9B) and in some linyphi-ids (eg in Labulla thoracica) Although Marples(1967 221) had reported substantial intraspecificvariation in the number of fusules we still lack a solidunderstanding of the levels of intraspecific variationwhich is essential to assess the phylogenetic informa-tion content of this character system

Pimoids are a relictual group distributed in thewestern coast of North America Europe (the Alps andthe Apennines and the Cantabrian Mountains ofnorthern Spain) and the Himalayas Their disjunctpattern suggests an ancestral widespread Holarcticdistribution with subsequent extinction in the inter-

vening areas (Hormiga 1994a) Hormiga (1994a) hadsuggested that new species of pimoids probablyexisted in the Himalayas The recently described newspecies from the Gaoligong Mountains of China (Gris-wold Long amp Hormiga 1999) confirms that predictionThe Japanese species described in this paper signifi-cantly expand the geographical range of the familyPimoidae

SYSTEMATICS

PIMOIDAE WUNDERLICH 1986

Pimoinae Wunderlich (1986 16)Pimoidae Hormiga (1993) type genus by monotypyPimoa Chamberlin and Ivie

DiagnosisMale palpus with a retrolateral cymbial sclerite a dor-soectal cymbial process and cuspules on either thecymbial process (Pimoa Hormiga 1994a figs 11 68)or the dorsal surface of the cymbium (WeintrauboaFig 1E 5E) Conductor and median apophysis presentin most species Embolus continuous with the tegulum(the typical linyphiid embolic division is absent) withan embolic process of varying morphology The epigy-num is protruding with a dorsal to lateral fold orgroove with the copulatory opening at the distal end(Hormiga 1994a figs 14 414 Fig 3A) fertilizationducts are anteriorly (Pimoa) or mesally (Weintrauboa)orientated As in linyphiids pimoids have stridulatorystriae on the ectal side of the chelicerae (but the striae

Figure 4 Weintrauboa contortipes (Karsch) Male (A) and female (B) from Doi Sakuto-Cho Okayama Pref (Japan)Carapace width 215 (male) 225 (female)

268 G HORMIGA


are absent in Weintrauboa) build sheet-webs andexhibit autospasy at the patella-tibia junction

DescriptionSee Hormiga (1994a) for family description

PhylogeneticsThe monophyly of Pimoidae is supported by the

following four unambiguous putative synapomor-phies a dorsoectal cymbial process cymbial cuspulesa retrolateral cymbial sclerite (PCS) and the embolicprocess (PEP)

CompositionTwo genera Pimoa Chamberlin amp Ivie and Weintrau-boa new genus

Figure 5 Weintrauboa contortipes (Karsch) A paracymbium ectal B tegular division ectal C tegular divisiondetail D cymbial process E cymbial cuspules F male PLS

AA BB

CC DDT

PEP

AC

ST

BH

MA

E

EF

C

EE FF

P

PEP

MA

C m

T

EF

E

m



WEINTRAUBOA NEW GENUS

Type species Linyphia contortipes Karsch 1881

EtymologyNamed after Professor Dr Robert L Weintraub (1912ndash1996) benefactor of the systematics programme at

The George Washington University Weintrauboa is anundeclinable proper name and masculine in gender

DiagnosisWeintrauboa differs from Pimoa by the following com-bination of characters male palpal tibia lacking dorsal

Figure 6 Weintrauboa chikunii (Oi) (AndashD) W contortipes (Karsch) (EF) A paracymbium and pimoid cymbial scleriteectal B detail tegular division ectal C cymbial process D cymbial cuspules E male epiandrous fusules F male epiandrousfusules detail

AA BB

CC DD

BH

EE FF

P

MAC

T

PCS

270 G HORMIGA


process distal end of cymbium finger-like dorsoectalcymbial process heavily sclerotized and without cus-pules cuspules arranged in rows on dorsal surface ofthe cymbium absence of an alveolar sclerite on theventral side of the cymbium a small elongated mem-

brane the embolic flap arising from the area wherethe embolic process (PEP) connects to the embolusPEP embolic process compact (filiform or lamelliformin Pimoa) of varying morphology enlarged and sinu-ous metatarsal I base in males cheliceral stridulatory

Figure 7 Weintrauboa contortipes (Karsch) male first metatarsus A ectoventral B ventral C dorsal D dorsal

AA BB

CC DD



Figure 8 Weintrauboa contortipes (Karsch) female spinnerets A left group B ALS C PMS D PLS

272 G HORMIGA


striae absent fertilization ducts mesally orientatedand PMS with at least two and PLS with at least fiveaciniform spigots

DescriptionSmall- to medium-sized pimoid spiders total length49ndash60 in males and 47ndash80 in females Carapacelonger than wide 22ndash29 long in males and 21ndash31 in

females with a conspicuous thoracic fovea (Fig 4)Colour photographs of Weintrauboa contortipes (Kar-sch) and W chikunii (Oi) are published in Chikuni(1989 figs 11 12) and Shinkai amp Takano (1984 27)Sternum longer than wide pointed in the posteriorregion slightly projecting between coxae IV (Fig 3E)AME very close together (Fig 3F) c 017ndash029 theirdiameter apart larger than the rest which areroughly of the same diameter ALE and PLE juxta-

Figure 9 Pimoid epiandrous fusules AB Weintrauboa contortipes CD Pimoa altioculata EF Pimoa breuili

AA BB

CC DD

EE FF



posed Clypeus height 138ndash15 times an AME diame-ter Chelicerae large (Fig 3F) with three prolateraland three or four retrolateral teeth stridulatory striaeabsent Legs slightly longer and slender in the adultmale femur through metatarsus usually with dark

annuli Femur I 11 in males and 10 in females timesthe length of cephalothorax Male metatarsus I prox-imal third enlarged and sinuous with a row orenlarged macrosetae (Figs 3GndashJ 7AndashD) Tibia IndashIVwith two dorsal spines prolateral and retrolateral tib-

Figure 10 Weintrauboa contortipes (Karsch) A epigynum lateral B male tarsal claws first leg (mesal)

AA BB

Figure 11 Strict consensus cladogram of the four minimal length trees of 172 steps that result from the analysis of thedata matrix presented in Appendix 2 (L = 174 CI = 055 RI = 074) Numbers next to nodes denote Bremer support values

TetragnathaTheridiosoma

Steatoda

Pimoa rupicolaPimoa breuiliPimoa lihengae

Pimoa altioculata

Weintrauboa contortipesWeintrauboa chikunii

Labulla thoracica

MicrolinyphiaLinyphia

BolyphantesTenuiphantes

StemonyphantesHaplinis

Novafroneta

AsthenargusGongylidiellum

SisicusTibioploides

OsteariusLeptorhoptrum

HilairaLaminacauda

DrepanotylusSciastes

IslandianaErigone

Walckenaeria

Pimoa

Weintrauboa

PIMOIDAE

LINYPHIIDAE

LINYPHIOIDS

1

11

1

1

1

1

1

1

1

1

3

2

22

3

4

3

3

4

1

11

1

274 G HORMIGA


ial spines present varying from one (male W chikunii)to two (male and female W contortipes) ventral tibialspines varying from four to six femur IndashIV with dorsalspines in variable numbers femur IndashIV with one dor-sal spine femur IndashII with one prolateral spine femurIIIndashIV without prolateral spines Metatarsal spinespresent in all legs Trichobothrium metatarsus I c 02trichobothrium metatarsus IV present Leg autospasyat the patellandashtibia junction Abdomen ovoid longerthan wide dark browngrey with lighter marks andsome guanine spots (see photographs in Chikuni1989) Colulus large and fleshy with setae Spinnerets

(studied in one male and one female W contortipes)typical of a lsquolinyphioidrsquo (Hormiga 1994a) (Fig 8AndashD)ALS with c 25 piriform spigots PMS have two acini-form spigots between the cylindrical and the minorampullate spigot PLS with 4ndash5 aciniform spigotsbetween the two cylindrical spigots base of theperipheral cylindrical spigot larger than the base ofthe distal one The flagelliform and aggregate spigots(araneoid lsquotripletrsquo) are well developed in the females(Fig 8D) reduced to nubbins in the adult males(Fig 5F) Epiandrous fusules (studied in one maleW contortipes Figs 6EF 9AB) arranged linearly

Figure 12 One of the four minimal length trees of 172 steps that result from the analysis of the data matrix presented inAppendix 2 (CI = 056 RI = 074) Exclusion of parsimony uninformative characters decreases the tree length to 164 stepsand the ensemble consistency index to 054 Most of the ambiguous character changes are resolved under lsquoFarris optimi-zationrsquo Closed rectangles represent non-homoplasious character changes The three nodes that collapse in the strict con-sensus cladogram of the four most parsimonious trees are marked with a closed circle The basal trichotomy has beenresolved according to the araneoid topology presented in Griswold et al (1998) (see text for details)

Tetragnatha

Theridiosoma

Steatoda

Pimoa rupicola

Pimoa breuili

Pimoa lihengae

Pimoa altioculata


Weintrauboa chikunii

Labulla thoracica

Microlinyphia

Linyphia

Bolyphantes

Tenuiphantes

Stemonyphantes

Haplinis

Novafroneta

Asthenargus

Gongylidiellum

Sisicus

Tibioploides

Ostearius

Leptorhoptrum

Hilaira

Laminacauda

Drepanotylus

Sciastes

Islandiana

Erigone

Walckenaeria

64

1

56

1

47

1

39

1

33

1

21

1

11

5

10

265

1

64

2

58

1

57

4

39

3

23

1

11

457

3

39

1

38

3

55

0

14

1

61

1

57

0

48

2

37

2

24

0

20

1

9

2

45

1

24

039

2

51

158

170

2

67

1

45

1

24

0

11

246

1

44

1

42

1

39

2

38

041

1

19

064

1

56

1

16

1

44

1

42

1

38

2

17

0

52

2

56

1

47

1

42

148

0

39

2

38

0

37

049

1

36

152

2

51

1

49

1

30

0

48

0

19

0

16

1

66

1

46

0

16

131

4

64

1

57

0

51

1

32

0

55

2

31

2

23

1

66

0

65

1

53

0

31

1

23

0

15

1

46

1

32

1

66

0

31

3

24

0

52

1

48

1

30

1

23

1

65

1

55

2

23

1

36

1

15

1

39

3

58

1

37

1

51

1

50

1

17

0

13

1

66

0

18

1

74

1

73

1

72

1

71

1

44

0

43

1

34

1

23

1

42

1

12

0

55

1

41

144

1

36

1

35

1

33

1

29

1

22

1

21

1

17

1

11

3

60

1

54

0

39

3

27

1

25

1

9

1

8

2

4

1

70

2

40

1

28

1

11

1

8

1

7

1

57

2

6

0

70

1

67

1

57

0

48

2

38

0

37

2

10

0

1

1

66

0

55

1

48

1

39

2

26

1

5

1

3

1

2

1

75

2

69

1

59

1

54

1

24

1

10

1

75

1

68

0

38

1

12

1

LINYPHIOIDS

PIMOIDAE

Pimoa

Weintrauboa

LINYPHIIDAE

9

2



along the posterior margin of the epigastric furrowmost of them in groups of 2ndash5 fusules emerging from acommon circular pit The fusules have a long shaft andare somewhat shorter than the surrounding macrose-tae Tracheal system morphology unknown Femalepedipalp with tarsal claw Male pedipalp patellashort with several thick dorsal macrosetae (Fig 1C)Tibia approximately as short as wide its distal marginwith a row of thick macrosetae (Fig 1C) one prolat-eral and one retrolateral trichobothrium On dorsalview cymbium widest at it median region (Figs 1E2C) with a sclerotized ectal process of varying degreesof complexity that has a conspicuous concavity facingventrally (Figs 1C 2B) alveolus occupying the basaltwo-thirds of the cymbium the distal third of the lat-ter much narrower and finger-like (Figs 1E 2C) Cus-pules present in rows on the proximal region of thedorsal part of the cymbium but absent from the cym-bial ectal process (Figs 1E 2C) Paracymbium inter-segmental and linguiform broadly attached to theectal margin of the cymbium base by means of a mem-brane its proximal region continuous with the pimoidcymbial sclerite (Fig 1B) Paracymbial apophysesabsent a sclerotized fold can be seen in the middlepart of the paracymbium (Figs 1B 2B) Pimoid cym-bial sclerite connected to the cymbium by a relativelyless sclerotized region (but not fully membranous)continuous with the paracymbium (Figs 1BC 2B)Tegulum with a conspicuous suture delimiting anapical tegular region less sclerotized and lighter incolour than the rest of the tegulum Conductor asmall longer than wide membrane without papillae(Fig 5C) Median apophysis a small process on tegu-lum adjacent to the base of the conductor (Fig 5C)Embolus widest at its base continuous with tegularmargin (Fig 1D) in ventral view arising from the pos-terior region of the tegulum curving clockwise (in leftpalp) around the anterior tegular area and shortlyafter curving anticlockwise to an almost full circle(Fig 1D) distal end of embolus located near the embo-lic process (PEP) PEP arises in ventral view frominner region of embolus (Fig 1DJ) PEP compact (notfiliform or lamelliform) with various processes(Fig 1J) in the membranous area where the PEP con-nects to the embolus there is a small elongated mem-branous flap (embolic flap Figs 1C 2B 5B) Epigynumprotruding less than its width (internal genitaliaunknown for W chikunii) copulatory ducts longmaking several turns before reaching elongated sper-mathecae fertilization ducts mesally orientated(Fig 3AndashC)

PhylogeneticsThe monophyly of Weintrauboa is supported by atleast the following putative synapomorphies narrow

(finger-like) distal end of cymbium embolic flap a sin-gle retrolateral trichobothrium in male pedipalpaltibia absence of cheliceral stridulatory striae and thepresence of an enlarged and sinuous metatarsal I basein males

Natural historyThe web of W contortipes is illustrated in Shinkai ampTakano (1984 27)

CompositionTwo species Weintrauboa contortipes (Karsch) andW chikunii (Oi)

DistributionJapan and adjacent islands

WEINTRAUBOA CONTORTIPES (KARSCH) NEW COMBINATION

Linyphia contortipes Karsch 1881 39Labulla contortipes Boumlsenberg amp Strand 1906 170

pl 5 figure 49 pl 11 figures 234 237 Yaginuma1971 43 figure 414 Shinkai amp Takano 1984 27Yaginuma 1986 72 figure 382 Chikuni 1989 48figure 11

TypesMale and female syntypes from Saga Japan (Doumlnitzleg) at Institut fuumlr Systematische Zoologie Univer-sitaumlt zu Berlin (ZMB3545) examined Although thesyntype label states lsquoSaga Japan leg Doumlnitzrsquo thislocality seems incorrect as the Japanese spiders thatKarsch described between 1879 and 1881 were col-lected by Doumlnitz around Tokyo where he was living atthe time before moving to Saga (H Ono unpubl data)

DiagnosisMales of W contortipes differ from those ofW chikunii in the shape of the ectal cymbial processand paracymbium the number of cymbial cuspules(more numerous in the latter species) themorphology of the PEP (larger and more complexin the former species) and in the degree of sinuosityof the metatarsus I base (less sinuous in W chikunii)The female genital morphology of the two Weintrau-boa species differs significantly in ventral view inW contortipes the epigynum appears as a protrusionof the abdomen a subtle bulge whereas inW chikunii it has a strongly marked cordiformappearance darker in the medial region

276 G HORMIGA


DescriptionMale (from Doi Sakuto-Cho Okayama Pref Fig 4A)Total length 543 Cephalothorax 254 long 217 wide161 high Sternum 155 long 127 wide Abdomen264 long 183 wide AME diameter 019 Clypeusheight 143 times one AME diameter Chelicerae withthree prolateral and three retrolateral teeth (Fig 3F)Femur I 282 long 111 times the length of cephalotho-rax Metatarsus proximal third enlarged and sinuouswith a row of enlarged macrosetae (Fig 3GH) Meta-tarsus I trichobothrium 019 Pedipalp as inFigures 1AndashJ 5AndashE Pedipalpal tibia with one prolat-eral and one retrolateral trichobothria

Female (same locality as male Fig 4B) Total length688 Cephalothorax 295 long 217 wide 146 highSternum 155 long 140 wide Abdomen 422 long 322wide AME diameter 021 Clypeus height 138 timesone AME diameter Chelicerae with three prolateraland four retrolateral teeth Femur I 281 long 095times the length of cephalothorax Metatarsus I tri-chobothrium 022 Epigynum as in Figures 2EF 3AndashC 10A

VariationMale cephalothorax ranges in length from 226 to 281(n = 4) Female cephalothorax ranges in length from214 to 312 (n = 8) Male total length ranges from 488to 577 (n = 3) Female total length ranges from 465 to800 (n = 8)

Natural historySheet web illustrated in Shinkai amp Takano (1984 27)the photograph shows an inclined (c 45infin) sheet webbuilt against a substrate with little if any scaffoldingabove or below the web Epigynal plugs are common

DistributionKy sh and Honsh islands (Japan)

Material examinedJapan Okayama Pref Doi Sakuto-cho Aida-gun26i1990 K ichi Nojima 1 male 1 female (YIPC)Okayama Pref Mt Takiyama Nagi-cho Katsuta-gun14iii1992 Kocircichi Nojima 1 male (YIPC) HiroshimaPref Mt Gokurakuji Hatsukaichi-shi 3v1991 1female Yoh Ihara (YIPC) Hiroshima Pref Taishaku-kyo Gorge Tojo-cho Hiba-gun 19vi1996 1 femaleYoh Ihara (YIPC) Okayama Pref Mt TakiyamaNagi-cho Katsuta-gun 4vi1989 K ichi Nojima 2female (YIPC) Hiroshima Pref Mt Gokurakuji Hat-sukaichi-shi 26i1992 Yoh Ihara 1 female (YIPC)

u- u- u-

o-

o-

Japan Rakusan Shimane 24iv1965 S Kageyama 1female (ZMUT) Mie Pref R Hashimoto no 6 1 male(ZMUT) lsquoJapanrsquo 3837 c St 1 male 1 female(MNHN)

WEINTRAUBOA CHIKUNII (OI) NEW COMBINATION

Labulla contortipes chikunii Oi 1979 330 figures 9ndash12 Chikuni 1989 48 figure 12

Labula chikunii Eskov 1992 53 (lapsus calami ele-vated from subspecies of Labulla contortipes)

NB Marusik et al (1993 75) synonymized Labullachikunii Oi 1979 with Lepthyphantes insularis SaitoThis synonymy is rejected here Tanasevitch amp Eskov(1987 194) had stated that Lepthyphantes insularisSaito described after a single female specimen fromthe Sakhalin Island did not belong in Lepthy-phantes based on Saitorsquos (1935 fig 1b) epigynumillustration (Tanasevitch amp Eskov did not provideany new illustrations or redescription of insularisnor did they examine the type) In their view Lepthy-phantes insularis should probably be transferred toLabulla and it could be a (junior) synonym ofL thoracica though they did not formalize anytransfer or new synonymy Oi (1979) described andillustrated both sexes of Labulla contortipes chikuniiSaitorsquos type material formerly at the University ofHokkaido is presumably lost (H Ono unpubl data)I have not been able to study any female specimensof chikunii (none was available for study) but Oirsquosillustration of the chikunii epigynum as well asChikunirsquos (1989 fig 12) excellent colour photographshows substantial differences with Saitorsquos (1935) epi-gynum illustration In Lepthyphantes insularis theventral scape is very short and wide and the poste-rior edges of the ventral epigynal wall curve posteri-orly The comparable structure in chikunii is muchlonger and thinner and the posterior edges of theventral epigynal wall curve first anteriorly beforedescending towards the epigastric furrow In addi-tion the leg formula in insularis is 4123 whereas inchikunii it is 1243 As Marusik et al (1993) did notbase their synonymy on examination of types andthe descriptions of these two species are quitedifferent on the basis of the available evidence it isunjustified to synonymize Labulla contortipeschikunii with Lepthyphantes insularis

TypesOirsquos types (male holotype and one female paratype)are apparently lost (H Ono unpubl data)

DiagnosisSee diagnosis of W contortipes



DescriptionMale (from Juzhno-Sakhalinsk Sakhalin Island)Cephalothorax 293 long 199 wide 195 high Ster-num 164 long 133 wide Abdomen (poorly preservedno accurate width measurement possible) 374 longAME diameter 020 Clypeus height 150 times oneAME diameter Chelicerae with three prolateral andfour retrolateral teeth Femur I 308 long 105 timesthe length of cephalothorax Metatarsus proximalthird enlarged and sinuous with a row of enlargedmacrosetae (Fig 3IJ) Metatarsus I trichobothrium020 Pedipalp as in Figures 2AndashD 6AndashD Pedipalpaltibia with one prolateral and one retrolateral tricho-bothria

Female No specimens were available for study Theepigynum has been illustrated by Oi (1979 fig 12)and photographed by Chikuni (1989 fig 12)

VariationMale cephalothorax ranges in length from 222 to 293(n = 2) An additional male specimen measured 491 intotal length The male in illustrated in Chikuni (1989fig 12) shows a metatarsus I with a more subtle basalprocess than the specimens studied here and a con-spicuous dark median band (very subtle in the speci-mens I examined) The Sakhalin Island specimensthat I have studied have shorter legs than thosedescribed in Oi (1979 Femur I is 148 times the lengthof cephalothorax) and Chikuni (1989 Femur I is 18times the length of cephalothorax)

DistributionRecorded from Horigane (Nagano-ken Pref) (Oi 1979)and Sakhalin island (Marusik et al 1993)

Material examinedRussia Sakhalin Island Juzhno-Sakhalinsk Touristvalley 18x1985 AM Basarukin 2 males (handwrit-ten label in Russian KEPC)

PIMOA CHAMBERLIN amp IVIE 1943

Type species (by original designation) Labullahespera Gertsch amp Ivie 1936

DiagnosisSee Weintrauboa diagnosis Male palpus (Hormiga1994a figs 9ndash11) with a retrolateral cymbial scleritewith denticles or cuspules (Hormiga 1994a figs 1168 absent in P edenticulata Hormiga) and an alveolarsclerite (Hormiga 1994a figs 54 46 156 190 233ndash

234 257 370) Pimoa males differ from those of Wein-trauboa in having a dorsal palpal tibial process (usu-ally round but conical in some species) theparacymbium continuous with the base of the cym-bium (in Weintrauboa the paracymbium is connectedto the cymbium by means of a membrane) having anelongated filiform or lamelliform embolic process andlacking an embolic flap Stridulatory striae present onthe ectal side of the chelicerae

DescriptionSee Hormiga (1994a)

PhylogeneticsThe monophyly of Pimoa is supported by at least thefollowing putative synapomorphies alveolar scleriteintegral paracymbium presence of a dorsal tibial pro-cess on the male palp two prolateral trichobothria onmale pedipalpal tibia epigynum with anteriorly ori-entated fertilization ducts and aciniform spigotsreduced to one or absent in the female PMS and PLS

CompositionTwenty-two species 21 are monographed in Hormiga(1994a) an additional species is described in Griswoldet al (1999) Additional new species have beenrecently discovered in China (CE Griswold and GHormiga unpubl data)

ACKNOWLEDGEMENTS

I would like to thank the following individuals andinstitutions for making specimens available for study(collection abbreviations used in text are given inparentheses) Jonathan Coddington (SmithsonianInstitution Washington DC USNM) Jason Dunlop(Institut fuumlr Systematische Zoologie Universitaumlt zuBerlin ISZ) Kirill Eskov (Moscow private collectionKEPC) Charles Griswold (California Academy of Sci-ences San Francisco CAS) Yoh Ihara (Hiroshimaprivate collection YIPC) Norman Platnick (AmericanMuseum of Natural History New York AMNH)Carles Ribera (Departament de Biologia Animal Uni-versitat de Barcelona UB) Christine Rollard(Museum National drsquoHistoire Naturelle ParisMNHN) Michael Saaristo (Zoological MuseumUniversity of Turku ZMUT) and Nikolaj Scharff (Zoo-logical Museum University of Copenhagen ZMUC)Dr Hirotsugu Ono (National Science Museum Tokyo)provided valuable information on specimens and col-lections sent me a copy of an obscure but importantreference and put me in contact with Mr Yoh Iharawhose private collection was instrumental in carrying

278 G HORMIGA


out this research Yura Marusik (Magadan) helped mein obtaining specimens of W chikunii deposited in Kir-ill Eskovrsquos personal collection Nikolaj Scharff helpedwith references from the ZMUC library H Don Cam-eron (University of Michigan Ann Arbor) providedexpert etymological advice Svetlana Maslakovahelped with Russian translations Nikolaj ScharffCharles Griswold Hirotsugu Ono and an anonymousreviewer provided very helpful comments on an ear-lier version of this paper Funding for this researchhas been provided by grants from the US National Sci-ence Foundation (DEB-9712353 and EAR-0228699)and the Research Enhancement Fund from TheGeorge Washington University

REFERENCES

Boumlsenberg W Strand E 1906 Japanische Spinnen Abhan-dlungen herausgegeben von der senckenbergischen naturfor-schenden Gesellschaft 30 93ndash422

Bremer K 1988 The limits of aminoacid sequence data inangiosperm phylogenetic reconstruction Evolution 42 795ndash803

Bremer K 1995 Branch support and tree stability Cladistics10 295ndash304

Chikuni Y 1989 Pictorial encyclopedia of spiders in JapanTokyo Kaisei-sha Publishers Co

Coddington JA 1983 A temporary slide mount allowing pre-cise manipulation of small structures In Kraus O ed Tax-onomy biology and ecology of Araneae and MyriapodaVerhandlungen des Naturwissenschaftlichen Vereins inHamburg 26 291ndash292

Denis J 1949 Notes sur les eacuterigonides XVI Essai sur ladeacutetermination des femelles drsquoeacuterigonides Bulletin de laSocieacuteteacute drsquoHistoire Naturelle de Toulouse 83 129ndash158

Donoghue MJ Olmstead RG Smith JF Palmer JD 1992Phylogenetic relationships of Dipsacales based on rbcLsequence data Annals of the Missouri Botanical Garden 79333ndash345

Eskov KY 1992 New data on fauna of spider family Linyphi-idae (Aranei) of the Soviet Far East Trudy zoologischeskiiInstitute Leningrad 226 51ndash59

Farris JS 1988 Hennig86 Version 15 Computer programdistributed by its author

Fitch WM 1971 Toward defining the course of evolution min-imum change for a specific tree topology Systematic Zoology20 406ndash416

Goloboff PA 1993 NONA Version 20 Program available athttpwwwcladisticscom

Griswold CE Coddington JA Hormiga G Scharff N1998 Phylogeny of the orb web building spiders (Araneo-morphae Orbiculariae Deinopoidea Araneoidea) Zoologi-cal Journal of the Linnean Society 123 1ndash99

Griswold CE Long C Hormiga G 1999 A new spider ofthe Genus Pimoa from the Gaoligongshan Mts YunnanChina (Araneae Pimoidae) Acta Botanica Yunnanica 1191ndash97

Holm Aring 1979 A taxonomic study of European and East Afri-can species of the genera Pelecopsis and Trichopterna (Ara-neae Linyphiidae) with descriptions of a new genus and twonew species of Pelecopsis from Kenya Zoologica Scripta 8255ndash278

Hormiga G 1993 Implications of the phylogeny of Pimoidaefor the systematics of linyphiid spiders (Araneae Arane-oidea Linyphiidae) Memoirs of the Queensland Museum 33533ndash542

Hormiga G 1994a A revision and cladistic analysis of thespider family Pimoidae (Araneoidea Araneae) SmithsonianContributions to Zoology 549 1ndash104

Hormiga G 1994b Cladistics and the comparative morphol-ogy of linyphiid spiders and their relatives (Araneae Arane-oidea Linyphiidae) Zoological Journal of the LinneanSociety 111 1ndash71

Hormiga G 2000 Higher level phylogenetics of erigonine spi-ders (Araneae Linyphiidae Erigoninae) Smithsonian Con-tributions to Zoology 609 1ndash160

Hormiga G 2002 Orsonwelles a new genus of giant linyphiidspiders (Araneae) from the Hawaiian islands InvertebrateSystematics 16 1ndash80

Hormiga G Arnedo MA Gillespie RG 2003 Speciation ona conveyor belt sequential colonization of the HawaiianIslands by Orsonwelles spiders (Araneae Linyphiidae) Sys-tematic Biology 52 70ndash88

Karsch F 1881 Diagnoses Arachnoidarum Japoniae Berlinerentomologische Zeitschrift 25 35ndash40

Locket GH Millidge AF 1953 British spiders Vol 2 Lon-don Ray Society

Marples BJ 1967 The spinnerets and epiandrous glands ofspiders Journal of the Linnean Society (Zoology) 46 209ndash223

Marusik YM Eskov KY Logunov DV Basarukin AM1993 A check-list of spiders (Arachnida Aranei) from Sakha-lin and Kurile Islands Arthropoda Selecta 1 73ndash85

Nixon K 1999 Winclada Version 10008 Program availableat httpwwwcladisticscom

Oi R 1979 (1980) New linyphiid spiders of Japan I (Linyphi-idae) Baika Literary Bulletin 16 325ndash341

Page RDM 2001 Nexus data editor Version 049 Available athttptaxonomyzoologyglaacukrodNDEndehtml

Platnick NI 2002 The world spider catalog Version 25American Museum of Natural History online at httpresearchamnhorgentomologyspiderscatalog81-87indexhtml

Saito S 1935 Further notes on spiders from southern Sagha-lin with descriptions of three new species Annotationes zoo-logicae Japonenses 15 58ndash61

Shinkai E Takano S 1984 A field guide to the spiders ofJapan Tokyo Tokai University Press

Simon E 1884 Les arachnides de France Paris 5 180ndash885Simon E 1900 Arachnida Fauna Hawaiiensis 2 443ndash519Tanasevitch AV Eskov KY 1987 Spiders of the genus Lep-

thyphantes (Aranei Linyphiidae) in the Siberian and Far-Eastern fauna Zoologiske Zhurnal 66 185ndash197

Wunderlich J 1986 Spinnenfauna Gestern und Heute Wies-baden Erich Bauer Verlag bei Quelle amp Meyer



Yaginuma T 1971 Spiders of Japan in color (enlarged andrevised edition) Osaka Hoikusha Publishers Co

Yaginuma T 1986 Spiders of Japan in color (new edition)Osaka Hoikusha Publishers Co

APPENDIX 1

Most of the characters presented here are discussed indetail and illustrated in Hormiga (2000) The majorityof characters relevant to pimoids are taken from Hor-miga (1994a) Only those characters that are new orthat have been recoded are discussed in this section

MALE PALP

1 Alveolar sclerite (0) absent (1) present (Hormiga1994a fig 54 the sclerite between the paracym-bium and the pimoid cymbial sclerite) This is asmall sclerite located on the ventral side of thecymbium anteroectal to the distal margin of thealveolus This sclerite appears as a dark sclerotizedplate that lies between the distal end of the alve-olus and the attachment of the pimoid cymbialsclerite (Hormiga 1994a p 6) The presence of thealveolar sclerite is a synapomorphy of Pimoa

2 Cymbium morphology (ectal region) (0) smooth(no process) (1) with ectal cymbial process Hor-miga (1994a) suggested that the presence of a dor-sal denticulate process was a synapomorphy ofpimoids The study of Weintrauboa required rede-fining this character because only in the genusPimoa are the denticles (cuspules) located on thecymbial process

3 Cymbial cuspules (0) absent (1) present4 Cymbial cuspules (0) on cymbial process itself (1)

on dorsal surface of cymbium (but not on processFigs 1E 5E)

5 Pimoid cymbial sclerite (PCS) (0) absent (1)present

6 Pimoid cymbial sclerite (0) with membranousflap (1) without membranous flap

7 PCS (0) continuous with paracymbium (1) sepa-rate from paracymbium The plesiomorphic condi-tion in pimoids is to have the PCS continuous withthe paracymbium as found in Weintrauboa(Fig 1B) and in Pimoa breuili and P rupicola(Hormiga 1994a figs 17 and 45)

8 PCS-cymbium connection (0) sclerotized andrigid (1) membranous (2) intermediate MostPimoa species (eg P hespera) have the PCS con-nected to the cymbium by means of a membraneIn P breuili and P rupicola the connection is scle-rotized and rigid (Hormiga 1994a figs 17 and 45)Weintrauboa species have an intermediate condi-tion with the PCS connected by means of a rigidmembranous area to the cymbium but at the same

time continuous with the paracymbium bothbeing heavily sclerotized

9 Distal end of cymbium (0) rounded (1) elongated(2) conical In Weintrauboa the distal end of thecymbium is distinctly narrowed finger-like(Fig 1E) In the mynoglenine genera Haplinis andNovafroneta and in the linyphiine genus Labullathe distal end of the cymbium is also somewhatelongated but not as distinctly narrowed as inWeintrauboa

10 Paracymbium attachment (0) integral (1) inter-segmental (2) articulated

11 Paracymbium morphology (0) linguiform (1) tri-angular (2) Stemonyphantes type (3) U or J (4)hook (5) straight and narrow

12 Paracymbium apophyses (0) present (1) absent13 Mynoglenine tegular apophyses (0) absent (1)

present14 Tegular suture (0) conspicuous (1) subtle or

absent15 Protegulum (0) absent (1) present16 Protegular papillae (0) absent (1) present17 Suprategulum (0) absent (1) present18 Suprategulum (0) continuous with tegulum (1)

articulated19 Suprategular distal apophysis (0) absent (1)

present20 Suprategular marginal apophysis (0) absent (1)

present21 Median apophysis (MA) (0) present (1) absent

Weintrauboa species have on their tegulum nearthe base of the conductor a small pointed protu-berance (Figs 5C 6B) that I have interpreted as ahomologue of the MA

22 Conductor (0) present (1) absent23 Embolus length (0) long (1) short24 Embolic membrane (0) absent (1) present25 Embolic flap (0) absent (1) present Weintrauboa

species have a small membranous flap that arisesfrom the area where the embolic process connectsto the embolus (Figs 1C 5B)

26 Pimoid embolic process (PEP) (0) absent (1)present

27 PEP (0) elongated (1) compact The PEP of Wein-trauboa species is a complex sclerotized structurewith several processes connected to a membra-nous area of the embolus (Fig 1DJ) The homolo-gous structure in Pimoa is either filiform orlamelliform and located ectally in reference to theembolus

28 Shape of elongated PEP (0) bifurcated (1) simple29 Radix (0) absent (1) present30 Radical tail piece (0) absent (1) present31 Radical tail piece (0) straight (1) spiralled (2)

curved ectally (3) curved mesally (4) anteriorlydirected

280 G HORMIGA


32 Anterior radical process (0) absent (1) present33 Column (0) absent (1) present34 Fickertrsquos gland (0) absent (1) present35 Terminal apophysis (TA) (0) absent (1) present

Pimoids and the non-linyphiid outgroup taxa havebeen coded as lacking a TA One could argue thatthese taxa need to be coded as inapplicable(because they lack the radix to which the TA con-nects see comments for next character)

36 Lamella characteristica (LC) (0) absent (1)present Pimoids and the non-linyphiid outgrouptaxa have been coded as lacking a LC Again onecould argue that these taxa need to be coded asinapplicable (because they lack the radix to whichthe LC connects) Coding characters 34 and 35 asinapplicable for pimoids produces the same topo-logical results than coding those cells as lsquoabsentrsquo

37 Male pedipalp tibial apophysis (0) absent (1) ret-rolateral (erigonine) (2) dorsal

38 Prolateral trichobothria on male pedipalpal tibia(0) two (1) one (2) zero (3) three

39 Retrolateral trichobothria on male pedipalpaltibia (0) two (1) four (2) three (3) one Steatodagrossa (Theridiidae) has one prolateral and tworetrolateral trichobothria on male pedipalpal tibia(I Agnarsson pers comm) and had been coded inerror in Griswold et al (1998) and Hormiga(2000)

EPIGYNUM

40 Degree of epigynal protrusion (0) protruding lessthan its width (1) protruding more than itswidth External epigynal morphology for Wein-trauboa chikunii has been coded after the illus-tration of Oi (1979 fig 12) and the photographsof Chikuni (1989 fig 12) No information is avail-able about the internal epigynal morphology ofW chikunii and no female specimens were avail-able for study

41 Dorsal plate scape (0) absent (1) present42 Ventral plate scape (0) absent (1) present43 Ventral plate scape (0) straight (1) sigmoid44 Atrium (0) absent (1) present45 Copulatory duct (0) separate from fertilization

duct (1) spirals fertilization duct46 Copulatory duct encapsulation (0) absent (1)

present47 Spermathecae (0) two (1) four48 Fertilization duct orientation (0) posterior (1)

mesal (2) anterior

SOMATIC MORPHOLOGY

49 Male post-PME lobe (0) absent (1) present50 Subocular clypeal sulci (0) absent (1) present

51 Male cephalothoracic cuticular pores (0) absent(1) present

52 Tracheal system (0) haplotracheate (1) desmitra-cheate (2) intermediate This character (and thenext one) could not be coded for Weintrauboabecause insufficient material was available fordissection

53 Taenidia in tracheoles (0) absent (1) present54 Ectal side of male chelicerae (0) smooth (1) with

stridulatory striae Weintrauboa lacks the cheli-ceral striae typical of Pimoa and most linyphiidsThis is interpreted as secondary absence

55 Cheliceral stridulatory striae (0) ridged (1) scaly(2) imbricated

56 Dorsal spur on male chelicerae (0) absent (1)present

57 Retrolateral teeth on female chelicera (0) three(1) four or more (2) two (3) zero (4) one

58 Female pedipalpal tarsus (0) with claw (1) with-out claw

59 Patella-tibia leg autospasy (0) absent (1) present60 Male Metatarsus I base diameter (proximal third

in dorsal view) (0) approximately the same diam-eter as the base or less (1) increasing in diameterup to twice or more the diameter of the proximalend (Fig 7C)

61 Male Metatarsus I proximal enlargement (0) sub-tle sinuosity (Fig 3I) (1) more sinuous (Fig 3G)

62 Dorsal spines on tibia I (0) two or more (1) one(2) zero

63 Dorsal spines on tibia II (0) two or more (1) one(2) zero

64 Dorsal spines on tibia III (0) two or more (1) one(2) zero

65 Dorsal spines on tibia IV (0) two or more (1) one(2) zero

66 Trichobothrium on metatarsus IV (0) present (1)absent

67 Aciniform spigots in female PMS (0) one or more(1) absent

68 PMS mAP nubbins (0) absent (1) one69 PLS mesal CY base (0) same size as other CY (1)

enlarged70 Aciniform spigots in female PLS (0) two or more

(1) one (2) absent71 PLS aggregate in male (0) absent (1) present72 PLS flagelliform in male (0) absent (1) present73 Male position during construction of sperm web

(0) above sperm web (1) below sperm web74 Male position during ejaculation (0) above sperm

web (1) below sperm web75 Web architecture (0) orb (1) tangle (2) sheet See

comments in Griswold et al (1998) for difficultiesin coding this behavioural suite (note that infig 5D of Griswold et al (1998) the web of Bathy-phantes pallidus was incorrectly labelled as



lsquoFrontinella pyramitelarsquo) Weintrauboa contortipeshas been coded as building a sheet web based onthe web photograph of Shinkai amp Takano

(1984 27) No information is available onW chikuniirsquos web Pimoa lihengae also buildssheet webs (Griswold et al 1999)

APPENDIX 2

CHARACTER STATE MATRIX

Tetragnatha 000-0---02500-0-0---100000--0---10000010------1ndash000-0-11000-001010100000Theridiosoma 000-0---00410-0-0---001000--0-ndash-000001300000000000-0-04100-002110100000Steatoda 000-0---0---0-0-0---000000--0-ndash-000001000000000000-0-03000-000010000001Pimoa rupicola 111010000001000-0---000-01000---0000202000000020000ndash102010ndash00000101100Pimoa breuili 111010000001000-0---000-01000---0000231000-000020000ndash110010ndash0000001100Pimoa lihengae 111011110011000-0---000-01010---0000202100-00002001100010ndash0000010122Pimoa altioculata 111011110011010-0---000-01010---0000202100-00002000ndash100010ndash000001012002Weintrauboa

contortipes011111021101000-0---000-111-0---0000013000ndash00001000ndash010111000000010002

Weintrauboachikunii

011111021101000-0---000-111-0---00000130-000ndash01100000

Labulla thoracica 000-0---21300-0-1011110000--10ndash01011210001010002000-1000010-000010010002Microlinyphia 000-0---01310-0-1010110000--10-01011010010-110000000-1101010ndash00001001000002Linyphia 000-0---01310-0-1010110100--10-01011012010-100000000-1101010ndash00001001000002Bolyphantes 000-0---01300-0-1010111100--10-011110100011000000010-1001010-00001001000Tenuiphantes 000-0---01300-0-1010111100--10-011110100011000000000-1001110-00001001000002Stemonyphantes 000-0---01210-0-1110110000--10--1000010000-010000000-100010ndash00000101211Haplinis 000-0---21311-0-0---110100--10-010000020010101000110-1001010ndash0000000101111Novafroneta 000-0---21311-0-0---110100--10-01000010010-000000110ndash1001010ndash00000001011Asthenargus 000-0---01310-101000111100--10-01001113000ndash00000000-01110ndash000111011Gongylidiellum 000-0---01310-111010111100--10-01001113000-000000000-1211110-00111001011Sisicus 000-0---01310-100-ndash110100--10-01001123001100000000-1001110-00001001011Tibioploides 000-0---01310-0-1010110100--10-01000113000-00000000211001110-0000100011Ostearius 000-0---01310-0-1010111100--110010001100010000110000-1011110-000010010112Leptorhoptrum 000-0---01310-0-1010111000--11301000002000ndash000000011101110ndash0000001011Hilaira 000-0---01310-0-1010111000--1-01001110000-00001100111001110-00000001011Laminacauda 000-0---01310-0-1010111100--11011000110000-00101001211001110-000010010112Drepanotylus 000-0---01310-101010110100--10-11000110000-00101100111001110-00001001011Sciastes 000-0---01310-111000111100--11011000110000-00100000111001110ndash00001001011Islandiana 000-0---01310-111010111100--11211000110000-00001000101201110ndash00011001011Erigone 000-0---01310-101010111100--11411000110000-00101000101201110ndash00010001011112Walckenaeria 000-0---01310-101010110100-11101000110000-00101001101000110-001100010112

262

G HORMIGA



2003

139

261ndash281

that these taxa belong into the family Pimoidae Inthis paper these Japanese species will be redescribedand its phylogenetic affinities explored using a cladis-tic approach In light of this new evidence Hormigarsquos(1994a) diagnoses for Pimoidae and

Pimoa

are alsorevised

MATERIAL AND METHODS

Morphological methods are described in detail in Hor-miga (2000 2002) Taxonomic descriptions follow theformat of Hormiga (1994a 2002) Specimens wereexamined and illustrated using a Leica MZAPO ste-reoscopic microscope with a camera lucida Furtherdetails were studied using a Leica DMRM compoundmicroscope with a drawing tube Some microscopeimages were recorded using a Leica DC-200 digitalcamera and edited using the software package Auto-Montage A LEO 1430VP scanning electron micro-scope was also used to study and photograph morpho-logical features Left structures (eg palps legs etc)are depicted unless otherwise stated Most hairs andmacrosetae are usually not depicted in the final palpdrawings All morphological measurements are givenin millimetres Somatic morphology measurementswere taken using a scale reticle in the dissectingmicroscope Eye diameters are taken from the span ofthe lens The cephalothorax length and height weremeasured in lateral view and its width was taken atthe widest point Similarly the length and height ofthe abdomen were measured in lateral view and thewidth as the widest point as seen from a dorsal viewThe measurements of the abdomen are only approxi-mations because the abdomen size changes moreeasily in preserved specimens than do other moresclerotized parts (eg the chelicerae) The total lengthwas measured in lateral view and is also an approxi-mation because it involves the size of the abdomen andits relative position Approximate leg article lengthswere measured in lateral view without detaching thelegs from the animal by positioning the article beingmeasured perpendicularly The position of the meta-tarsal trichobothrium is expressed as in Denis (1949)and Locket amp Millidge (1953) Female genitalia wereexcised using surgical blades or sharpened needlesThe specimen was then transferred to methyl salicy-late (Holm 1979) for examination under the micro-scope temporarily mounted as described inCoddington (1983) Male palps examined with theSEM were first excised and transferred to a vial with70 ethanol and then cleaned ultrasonically for1ndash3 min The specimen was then transferred to abso-lute ethanol and left overnight After critical pointdrying the specimens were glued to rounded alumin-ium rivets using an acetone solution of polyvinyl resinand then AuPd coated for examination at the SEM

A

NATOMICAL

ABBREVIATIONS

USED

IN

THE

TEXT

AND

FIGURES

Male palp

BH basal haematodochaC conductorCB cymbiumCP cymbial process (CDP in Hormiga 1994a)E embolusEF embolic flapm membrane (or membranous)MA median apophysisP paracymbiumPCS pimoid cymbial scleritePEP pimoid embolic processST subtegulumT tegulum

Epigynum

CD copulatory ductCO copulatory openingFD fertilization ductS spermatheca

Somatic morphology

AC aciniform gland spigot(s)AG aggregate gland spigot(s)ALS anterior lateral spinneretAME anterior median eye(s)CY cylindrical gland spigot(s)FL flagelliform gland spigot(s)MAP major ampullate gland spigot(s)mAP minor ampullate gland spigot(s)PI piriform gland spigot(s)PLE posterior lateral eye(s)PLS posterior lateral spinneretPME posterior median eye(s)PMS posterior median spinneret

Institutional abbreviations used in text are given inthe lsquoAcknowledgementsrsquo section

C

LADISTIC

ANALYSIS

Taxa

The taxonomic sample used in the cladistic analysiswas designed to test the monophyly of

Weintrauboa

and

Pimoa

and builds on my previous phylogeneticwork on the higher level phylogenetics of lsquolinyphioidsrsquo(Hormiga 1994a b 2000 2002 Hormiga

et al

2003)The monophyly of

Pimoa

had been previously testedby Hormiga (1994a) in the present study

Pimoa

is rep-resented by four species (out of 22 described species)including the two most basal taxa in Hormigarsquos(1994a fig 442) preferred cladogram

P rupicola

THE SPIDER GENUS

WEINTRAUBOA

(PIMOIDAE)

263



2003

139

261ndash281

(Simon) and

P breuili


Pimoa


Weintrauboa


Pimoa


Labulla thoracica


Labulla


Characters


Labulla thoracica


Analyses


et al

1992)

RESULTS


+



=



Stemonyphantes

Myno-gleninae (

Haplinis

plus

Novafroneta


Drepanotylus

Sciastes


Islandiana

Erigone

and

Walcke-naeria


DISCUSSION


Labulla



and

W chikunii


Weintrauboa


Pimoa


Pimoa

Until now

Pimoa


Weintrau-boa


Weintrauboa


Weintrauboa


Pimoa

264

G HORMIGA



2003

139

261ndash281

Figure 1


(Karsch)

n comb


THE SPIDER GENUS

WEINTRAUBOA

(PIMOIDAE)

265



2003

139

261ndash281


Weintrauboa


Pimoa


Pimoa breuili

and




266 G HORMIGA









SYSTEMATICS





268 G HORMIGA








AA BB

CC DDT

PEP

AC

ST

BH

MA

E

EF

C

EE FF

P

PEP

MA

C m

T

EF

E

m









AA BB

CC DD

BH

EE FF

P

MAC

T

PCS

270 G HORMIGA





AA BB

CC DD




272 G HORMIGA






AA BB

CC DD

EE FF






AA BB



Steatoda


Pimoa altioculata


Labulla thoracica




Novafroneta


SisicusTibioploides


HilairaLaminacauda


IslandianaErigone

Walckenaeria

Pimoa

Weintrauboa

PIMOIDAE

LINYPHIIDAE

LINYPHIOIDS

1

11

1

1

1

1

1

1

1

1

3

2

22

3

4

3

3

4

1

11

1

274 G HORMIGA





Tetragnatha

Theridiosoma

Steatoda

Pimoa rupicola

Pimoa breuili

Pimoa lihengae

Pimoa altioculata



Labulla thoracica

Microlinyphia

Linyphia

Bolyphantes

Tenuiphantes

Stemonyphantes

Haplinis

Novafroneta

Asthenargus

Gongylidiellum

Sisicus

Tibioploides

Ostearius

Leptorhoptrum

Hilaira

Laminacauda

Drepanotylus

Sciastes

Islandiana

Erigone

Walckenaeria

64

1

56

1

47

1

39

1

33

1

21

1

11

5

10

265

1

64

2

58

1

57

4

39

3

23

1

11

457

3

39

1

38

3

55

0

14

1

61

1

57

0

48

2

37

2

24

0

20

1

9

2

45

1

24

039

2

51

158

170

2

67

1

45

1

24

0

11

246

1

44

1

42

1

39

2

38

041

1

19

064

1

56

1

16

1

44

1

42

1

38

2

17

0

52

2

56

1

47

1

42

148

0

39

2

38

0

37

049

1

36

152

2

51

1

49

1

30

0

48

0

19

0

16

1

66

1

46

0

16

131

4

64

1

57

0

51

1

32

0

55

2

31

2

23

1

66

0

65

1

53

0

31

1

23

0

15

1

46

1

32

1

66

0

31

3

24

0

52

1

48

1

30

1

23

1

65

1

55

2

23

1

36

1

15

1

39

3

58

1

37

1

51

1

50

1

17

0

13

1

66

0

18

1

74

1

73

1

72

1

71

1

44

0

43

1

34

1

23

1

42

1

12

0

55

1

41

144

1

36

1

35

1

33

1

29

1

22

1

21

1

17

1

11

3

60

1

54

0

39

3

27

1

25

1

9

1

8

2

4

1

70

2

40

1

28

1

11

1

8

1

7

1

57

2

6

0

70

1

67

1

57

0

48

2

38

0

37

2

10

0

1

1

66

0

55

1

48

1

39

2

26

1

5

1

3

1

2

1

75

2

69

1

59

1

54

1

24

1

10

1

75

1

68

0

38

1

12

1

LINYPHIOIDS

PIMOIDAE

Pimoa

Weintrauboa

LINYPHIIDAE

9

2














276 G HORMIGA








u- u- u-

o-

o-






















ACKNOWLEDGEMENTS


278 G HORMIGA



REFERENCES






































APPENDIX 1


MALE PALP


























280 G HORMIGA








EPIGYNUM





mesal (2) anterior

SOMATIC MORPHOLOGY





























APPENDIX 2




Weintrauboachikunii

011111021101000-0---000-111-0---00000130-000ndash01100000


THE SPIDER GENUS

WEINTRAUBOA

(PIMOIDAE)

263



2003

139

261ndash281

(Simon) and

P breuili


Pimoa


Weintrauboa


Pimoa


Labulla thoracica


Labulla


Characters


Labulla thoracica


Analyses


et al

1992)

RESULTS


+



=



Stemonyphantes

Myno-gleninae (

Haplinis

plus

Novafroneta


Drepanotylus

Sciastes


Islandiana

Erigone

and

Walcke-naeria


DISCUSSION


Labulla



and

W chikunii


Weintrauboa


Pimoa


Pimoa

Until now

Pimoa


Weintrau-boa


Weintrauboa


Weintrauboa


Pimoa

264

G HORMIGA



2003

139

261ndash281

Figure 1


(Karsch)

n comb


THE SPIDER GENUS

WEINTRAUBOA

(PIMOIDAE)

265



2003

139

261ndash281


Weintrauboa


Pimoa


Pimoa breuili

and




266 G HORMIGA









SYSTEMATICS





268 G HORMIGA








AA BB

CC DDT

PEP

AC

ST

BH

MA

E

EF

C

EE FF

P

PEP

MA

C m

T

EF

E

m









AA BB

CC DD

BH

EE FF

P

MAC

T

PCS

270 G HORMIGA





AA BB

CC DD




272 G HORMIGA






AA BB

CC DD

EE FF






AA BB



Steatoda


Pimoa altioculata


Labulla thoracica




Novafroneta


SisicusTibioploides


HilairaLaminacauda


IslandianaErigone

Walckenaeria

Pimoa

Weintrauboa

PIMOIDAE

LINYPHIIDAE

LINYPHIOIDS

1

11

1

1

1

1

1

1

1

1

3

2

22

3

4

3

3

4

1

11

1

274 G HORMIGA





Tetragnatha

Theridiosoma

Steatoda

Pimoa rupicola

Pimoa breuili

Pimoa lihengae

Pimoa altioculata



Labulla thoracica

Microlinyphia

Linyphia

Bolyphantes

Tenuiphantes

Stemonyphantes

Haplinis

Novafroneta

Asthenargus

Gongylidiellum

Sisicus

Tibioploides

Ostearius

Leptorhoptrum

Hilaira

Laminacauda

Drepanotylus

Sciastes

Islandiana

Erigone

Walckenaeria

64

1

56

1

47

1

39

1

33

1

21

1

11

5

10

265

1

64

2

58

1

57

4

39

3

23

1

11

457

3

39

1

38

3

55

0

14

1

61

1

57

0

48

2

37

2

24

0

20

1

9

2

45

1

24

039

2

51

158

170

2

67

1

45

1

24

0

11

246

1

44

1

42

1

39

2

38

041

1

19

064

1

56

1

16

1

44

1

42

1

38

2

17

0

52

2

56

1

47

1

42

148

0

39

2

38

0

37

049

1

36

152

2

51

1

49

1

30

0

48

0

19

0

16

1

66

1

46

0

16

131

4

64

1

57

0

51

1

32

0

55

2

31

2

23

1

66

0

65

1

53

0

31

1

23

0

15

1

46

1

32

1

66

0

31

3

24

0

52

1

48

1

30

1

23

1

65

1

55

2

23

1

36

1

15

1

39

3

58

1

37

1

51

1

50

1

17

0

13

1

66

0

18

1

74

1

73

1

72

1

71

1

44

0

43

1

34

1

23

1

42

1

12

0

55

1

41

144

1

36

1

35

1

33

1

29

1

22

1

21

1

17

1

11

3

60

1

54

0

39

3

27

1

25

1

9

1

8

2

4

1

70

2

40

1

28

1

11

1

8

1

7

1

57

2

6

0

70

1

67

1

57

0

48

2

38

0

37

2

10

0

1

1

66

0

55

1

48

1

39

2

26

1

5

1

3

1

2

1

75

2

69

1

59

1

54

1

24

1

10

1

75

1

68

0

38

1

12

1

LINYPHIOIDS

PIMOIDAE

Pimoa

Weintrauboa

LINYPHIIDAE

9

2














276 G HORMIGA








u- u- u-

o-

o-






















ACKNOWLEDGEMENTS


278 G HORMIGA



REFERENCES






































APPENDIX 1


MALE PALP


























280 G HORMIGA








EPIGYNUM





mesal (2) anterior

SOMATIC MORPHOLOGY





























APPENDIX 2




Weintrauboachikunii

011111021101000-0---000-111-0---00000130-000ndash01100000


264

G HORMIGA



2003

139

261ndash281

Figure 1


(Karsch)

n comb


THE SPIDER GENUS

WEINTRAUBOA

(PIMOIDAE)

265



2003

139

261ndash281


Weintrauboa


Pimoa


Pimoa breuili

and




266 G HORMIGA









SYSTEMATICS





268 G HORMIGA








AA BB

CC DDT

PEP

AC

ST

BH

MA

E

EF

C

EE FF

P

PEP

MA

C m

T

EF

E

m









AA BB

CC DD

BH

EE FF

P

MAC

T

PCS

270 G HORMIGA





AA BB

CC DD




272 G HORMIGA






AA BB

CC DD

EE FF






AA BB



Steatoda


Pimoa altioculata


Labulla thoracica




Novafroneta


SisicusTibioploides


HilairaLaminacauda


IslandianaErigone

Walckenaeria

Pimoa

Weintrauboa

PIMOIDAE

LINYPHIIDAE

LINYPHIOIDS

1

11

1

1

1

1

1

1

1

1

3

2

22

3

4

3

3

4

1

11

1

274 G HORMIGA





Tetragnatha

Theridiosoma

Steatoda

Pimoa rupicola

Pimoa breuili

Pimoa lihengae

Pimoa altioculata



Labulla thoracica

Microlinyphia

Linyphia

Bolyphantes

Tenuiphantes

Stemonyphantes

Haplinis

Novafroneta

Asthenargus

Gongylidiellum

Sisicus

Tibioploides

Ostearius

Leptorhoptrum

Hilaira

Laminacauda

Drepanotylus

Sciastes

Islandiana

Erigone

Walckenaeria

64

1

56

1

47

1

39

1

33

1

21

1

11

5

10

265

1

64

2

58

1

57

4

39

3

23

1

11

457

3

39

1

38

3

55

0

14

1

61

1

57

0

48

2

37

2

24

0

20

1

9

2

45

1

24

039

2

51

158

170

2

67

1

45

1

24

0

11

246

1

44

1

42

1

39

2

38

041

1

19

064

1

56

1

16

1

44

1

42

1

38

2

17

0

52

2

56

1

47

1

42

148

0

39

2

38

0

37

049

1

36

152

2

51

1

49

1

30

0

48

0

19

0

16

1

66

1

46

0

16

131

4

64

1

57

0

51

1

32

0

55

2

31

2

23

1

66

0

65

1

53

0

31

1

23

0

15

1

46

1

32

1

66

0

31

3

24

0

52

1

48

1

30

1

23

1

65

1

55

2

23

1

36

1

15

1

39

3

58

1

37

1

51

1

50

1

17

0

13

1

66

0

18

1

74

1

73

1

72

1

71

1

44

0

43

1

34

1

23

1

42

1

12

0

55

1

41

144

1

36

1

35

1

33

1

29

1

22

1

21

1

17

1

11

3

60

1

54

0

39

3

27

1

25

1

9

1

8

2

4

1

70

2

40

1

28

1

11

1

8

1

7

1

57

2

6

0

70

1

67

1

57

0

48

2

38

0

37

2

10

0

1

1

66

0

55

1

48

1

39

2

26

1

5

1

3

1

2

1

75

2

69

1

59

1

54

1

24

1

10

1

75

1

68

0

38

1

12

1

LINYPHIOIDS

PIMOIDAE

Pimoa

Weintrauboa

LINYPHIIDAE

9

2














276 G HORMIGA








u- u- u-

o-

o-






















ACKNOWLEDGEMENTS


278 G HORMIGA



REFERENCES






































APPENDIX 1


MALE PALP


























280 G HORMIGA








EPIGYNUM





mesal (2) anterior

SOMATIC MORPHOLOGY





























APPENDIX 2




Weintrauboachikunii

011111021101000-0---000-111-0---00000130-000ndash01100000


THE SPIDER GENUS

WEINTRAUBOA

(PIMOIDAE)

265



2003

139

261ndash281


Weintrauboa


Pimoa


Pimoa breuili

and




266 G HORMIGA









SYSTEMATICS





268 G HORMIGA








AA BB

CC DDT

PEP

AC

ST

BH

MA

E

EF

C

EE FF

P

PEP

MA

C m

T

EF

E

m









AA BB

CC DD

BH

EE FF

P

MAC

T

PCS

270 G HORMIGA





AA BB

CC DD




272 G HORMIGA






AA BB

CC DD

EE FF






AA BB



Steatoda


Pimoa altioculata


Labulla thoracica




Novafroneta


SisicusTibioploides


HilairaLaminacauda


IslandianaErigone

Walckenaeria

Pimoa

Weintrauboa

PIMOIDAE

LINYPHIIDAE

LINYPHIOIDS

1

11

1

1

1

1

1

1

1

1

3

2

22

3

4

3

3

4

1

11

1

274 G HORMIGA





Tetragnatha

Theridiosoma

Steatoda

Pimoa rupicola

Pimoa breuili

Pimoa lihengae

Pimoa altioculata



Labulla thoracica

Microlinyphia

Linyphia

Bolyphantes

Tenuiphantes

Stemonyphantes

Haplinis

Novafroneta

Asthenargus

Gongylidiellum

Sisicus

Tibioploides

Ostearius

Leptorhoptrum

Hilaira

Laminacauda

Drepanotylus

Sciastes

Islandiana

Erigone

Walckenaeria

64

1

56

1

47

1

39

1

33

1

21

1

11

5

10

265

1

64

2

58

1

57

4

39

3

23

1

11

457

3

39

1

38

3

55

0

14

1

61

1

57

0

48

2

37

2

24

0

20

1

9

2

45

1

24

039

2

51

158

170

2

67

1

45

1

24

0

11

246

1

44

1

42

1

39

2

38

041

1

19

064

1

56

1

16

1

44

1

42

1

38

2

17

0

52

2

56

1

47

1

42

148

0

39

2

38

0

37

049

1

36

152

2

51

1

49

1

30

0

48

0

19

0

16

1

66

1

46

0

16

131

4

64

1

57

0

51

1

32

0

55

2

31

2

23

1

66

0

65

1

53

0

31

1

23

0

15

1

46

1

32

1

66

0

31

3

24

0

52

1

48

1

30

1

23

1

65

1

55

2

23

1

36

1

15

1

39

3

58

1

37

1

51

1

50

1

17

0

13

1

66

0

18

1

74

1

73

1

72

1

71

1

44

0

43

1

34

1

23

1

42

1

12

0

55

1

41

144

1

36

1

35

1

33

1

29

1

22

1

21

1

17

1

11

3

60

1

54

0

39

3

27

1

25

1

9

1

8

2

4

1

70

2

40

1

28

1

11

1

8

1

7

1

57

2

6

0

70

1

67

1

57

0

48

2

38

0

37

2

10

0

1

1

66

0

55

1

48

1

39

2

26

1

5

1

3

1

2

1

75

2

69

1

59

1

54

1

24

1

10

1

75

1

68

0

38

1

12

1

LINYPHIOIDS

PIMOIDAE

Pimoa

Weintrauboa

LINYPHIIDAE

9

2














276 G HORMIGA








u- u- u-

o-

o-






















ACKNOWLEDGEMENTS


278 G HORMIGA



REFERENCES






































APPENDIX 1


MALE PALP


























280 G HORMIGA








EPIGYNUM





mesal (2) anterior

SOMATIC MORPHOLOGY





























APPENDIX 2




Weintrauboachikunii

011111021101000-0---000-111-0---00000130-000ndash01100000


266 G HORMIGA









SYSTEMATICS





268 G HORMIGA








AA BB

CC DDT

PEP

AC

ST

BH

MA

E

EF

C

EE FF

P

PEP

MA

C m

T

EF

E

m









AA BB

CC DD

BH

EE FF

P

MAC

T

PCS

270 G HORMIGA





AA BB

CC DD




272 G HORMIGA






AA BB

CC DD

EE FF






AA BB



Steatoda


Pimoa altioculata


Labulla thoracica




Novafroneta


SisicusTibioploides


HilairaLaminacauda


IslandianaErigone

Walckenaeria

Pimoa

Weintrauboa

PIMOIDAE

LINYPHIIDAE

LINYPHIOIDS

1

11

1

1

1

1

1

1

1

1

3

2

22

3

4

3

3

4

1

11

1

274 G HORMIGA





Tetragnatha

Theridiosoma

Steatoda

Pimoa rupicola

Pimoa breuili

Pimoa lihengae

Pimoa altioculata



Labulla thoracica

Microlinyphia

Linyphia

Bolyphantes

Tenuiphantes

Stemonyphantes

Haplinis

Novafroneta

Asthenargus

Gongylidiellum

Sisicus

Tibioploides

Ostearius

Leptorhoptrum

Hilaira

Laminacauda

Drepanotylus

Sciastes

Islandiana

Erigone

Walckenaeria

64

1

56

1

47

1

39

1

33

1

21

1

11

5

10

265

1

64

2

58

1

57

4

39

3

23

1

11

457

3

39

1

38

3

55

0

14

1

61

1

57

0

48

2

37

2

24

0

20

1

9

2

45

1

24

039

2

51

158

170

2

67

1

45

1

24

0

11

246

1

44

1

42

1

39

2

38

041

1

19

064

1

56

1

16

1

44

1

42

1

38

2

17

0

52

2

56

1

47

1

42

148

0

39

2

38

0

37

049

1

36

152

2

51

1

49

1

30

0

48

0

19

0

16

1

66

1

46

0

16

131

4

64

1

57

0

51

1

32

0

55

2

31

2

23

1

66

0

65

1

53

0

31

1

23

0

15

1

46

1

32

1

66

0

31

3

24

0

52

1

48

1

30

1

23

1

65

1

55

2

23

1

36

1

15

1

39

3

58

1

37

1

51

1

50

1

17

0

13

1

66

0

18

1

74

1

73

1

72

1

71

1

44

0

43

1

34

1

23

1

42

1

12

0

55

1

41

144

1

36

1

35

1

33

1

29

1

22

1

21

1

17

1

11

3

60

1

54

0

39

3

27

1

25

1

9

1

8

2

4

1

70

2

40

1

28

1

11

1

8

1

7

1

57

2

6

0

70

1

67

1

57

0

48

2

38

0

37

2

10

0

1

1

66

0

55

1

48

1

39

2

26

1

5

1

3

1

2

1

75

2

69

1

59

1

54

1

24

1

10

1

75

1

68

0

38

1

12

1

LINYPHIOIDS

PIMOIDAE

Pimoa

Weintrauboa

LINYPHIIDAE

9

2














276 G HORMIGA








u- u- u-

o-

o-






















ACKNOWLEDGEMENTS


278 G HORMIGA



REFERENCES






































APPENDIX 1


MALE PALP


























280 G HORMIGA








EPIGYNUM





mesal (2) anterior

SOMATIC MORPHOLOGY





























APPENDIX 2




Weintrauboachikunii

011111021101000-0---000-111-0---00000130-000ndash01100000








SYSTEMATICS





268 G HORMIGA








AA BB

CC DDT

PEP

AC

ST

BH

MA

E

EF

C

EE FF

P

PEP

MA

C m

T

EF

E

m









AA BB

CC DD

BH

EE FF

P

MAC

T

PCS

270 G HORMIGA





AA BB

CC DD




272 G HORMIGA






AA BB

CC DD

EE FF






AA BB



Steatoda


Pimoa altioculata


Labulla thoracica




Novafroneta


SisicusTibioploides


HilairaLaminacauda


IslandianaErigone

Walckenaeria

Pimoa

Weintrauboa

PIMOIDAE

LINYPHIIDAE

LINYPHIOIDS

1

11

1

1

1

1

1

1

1

1

3

2

22

3

4

3

3

4

1

11

1

274 G HORMIGA





Tetragnatha

Theridiosoma

Steatoda

Pimoa rupicola

Pimoa breuili

Pimoa lihengae

Pimoa altioculata



Labulla thoracica

Microlinyphia

Linyphia

Bolyphantes

Tenuiphantes

Stemonyphantes

Haplinis

Novafroneta

Asthenargus

Gongylidiellum

Sisicus

Tibioploides

Ostearius

Leptorhoptrum

Hilaira

Laminacauda

Drepanotylus

Sciastes

Islandiana

Erigone

Walckenaeria

64

1

56

1

47

1

39

1

33

1

21

1

11

5

10

265

1

64

2

58

1

57

4

39

3

23

1

11

457

3

39

1

38

3

55

0

14

1

61

1

57

0

48

2

37

2

24

0

20

1

9

2

45

1

24

039

2

51

158

170

2

67

1

45

1

24

0

11

246

1

44

1

42

1

39

2

38

041

1

19

064

1

56

1

16

1

44

1

42

1

38

2

17

0

52

2

56

1

47

1

42

148

0

39

2

38

0

37

049

1

36

152

2

51

1

49

1

30

0

48

0

19

0

16

1

66

1

46

0

16

131

4

64

1

57

0

51

1

32

0

55

2

31

2

23

1

66

0

65

1

53

0

31

1

23

0

15

1

46

1

32

1

66

0

31

3

24

0

52

1

48

1

30

1

23

1

65

1

55

2

23

1

36

1

15

1

39

3

58

1

37

1

51

1

50

1

17

0

13

1

66

0

18

1

74

1

73

1

72

1

71

1

44

0

43

1

34

1

23

1

42

1

12

0

55

1

41

144

1

36

1

35

1

33

1

29

1

22

1

21

1

17

1

11

3

60

1

54

0

39

3

27

1

25

1

9

1

8

2

4

1

70

2

40

1

28

1

11

1

8

1

7

1

57

2

6

0

70

1

67

1

57

0

48

2

38

0

37

2

10

0

1

1

66

0

55

1

48

1

39

2

26

1

5

1

3

1

2

1

75

2

69

1

59

1

54

1

24

1

10

1

75

1

68

0

38

1

12

1

LINYPHIOIDS

PIMOIDAE

Pimoa

Weintrauboa

LINYPHIIDAE

9

2














276 G HORMIGA








u- u- u-

o-

o-






















ACKNOWLEDGEMENTS


278 G HORMIGA



REFERENCES






































APPENDIX 1


MALE PALP


























280 G HORMIGA








EPIGYNUM





mesal (2) anterior

SOMATIC MORPHOLOGY





























APPENDIX 2




Weintrauboachikunii

011111021101000-0---000-111-0---00000130-000ndash01100000


268 G HORMIGA








AA BB

CC DDT

PEP

AC

ST

BH

MA

E

EF

C

EE FF

P

PEP

MA

C m

T

EF

E

m









AA BB

CC DD

BH

EE FF

P

MAC

T

PCS

270 G HORMIGA





AA BB

CC DD




272 G HORMIGA






AA BB

CC DD

EE FF






AA BB



Steatoda


Pimoa altioculata


Labulla thoracica




Novafroneta


SisicusTibioploides


HilairaLaminacauda


IslandianaErigone

Walckenaeria

Pimoa

Weintrauboa

PIMOIDAE

LINYPHIIDAE

LINYPHIOIDS

1

11

1

1

1

1

1

1

1

1

3

2

22

3

4

3

3

4

1

11

1

274 G HORMIGA





Tetragnatha

Theridiosoma

Steatoda

Pimoa rupicola

Pimoa breuili

Pimoa lihengae

Pimoa altioculata



Labulla thoracica

Microlinyphia

Linyphia

Bolyphantes

Tenuiphantes

Stemonyphantes

Haplinis

Novafroneta

Asthenargus

Gongylidiellum

Sisicus

Tibioploides

Ostearius

Leptorhoptrum

Hilaira

Laminacauda

Drepanotylus

Sciastes

Islandiana

Erigone

Walckenaeria

64

1

56

1

47

1

39

1

33

1

21

1

11

5

10

265

1

64

2

58

1

57

4

39

3

23

1

11

457

3

39

1

38

3

55

0

14

1

61

1

57

0

48

2

37

2

24

0

20

1

9

2

45

1

24

039

2

51

158

170

2

67

1

45

1

24

0

11

246

1

44

1

42

1

39

2

38

041

1

19

064

1

56

1

16

1

44

1

42

1

38

2

17

0

52

2

56

1

47

1

42

148

0

39

2

38

0

37

049

1

36

152

2

51

1

49

1

30

0

48

0

19

0

16

1

66

1

46

0

16

131

4

64

1

57

0

51

1

32

0

55

2

31

2

23

1

66

0

65

1

53

0

31

1

23

0

15

1

46

1

32

1

66

0

31

3

24

0

52

1

48

1

30

1

23

1

65

1

55

2

23

1

36

1

15

1

39

3

58

1

37

1

51

1

50

1

17

0

13

1

66

0

18

1

74

1

73

1

72

1

71

1

44

0

43

1

34

1

23

1

42

1

12

0

55

1

41

144

1

36

1

35

1

33

1

29

1

22

1

21

1

17

1

11

3

60

1

54

0

39

3

27

1

25

1

9

1

8

2

4

1

70

2

40

1

28

1

11

1

8

1

7

1

57

2

6

0

70

1

67

1

57

0

48

2

38

0

37

2

10

0

1

1

66

0

55

1

48

1

39

2

26

1

5

1

3

1

2

1

75

2

69

1

59

1

54

1

24

1

10

1

75

1

68

0

38

1

12

1

LINYPHIOIDS

PIMOIDAE

Pimoa

Weintrauboa

LINYPHIIDAE

9

2














276 G HORMIGA








u- u- u-

o-

o-






















ACKNOWLEDGEMENTS


278 G HORMIGA



REFERENCES






































APPENDIX 1


MALE PALP


























280 G HORMIGA








EPIGYNUM





mesal (2) anterior

SOMATIC MORPHOLOGY





























APPENDIX 2




Weintrauboachikunii

011111021101000-0---000-111-0---00000130-000ndash01100000










AA BB

CC DD

BH

EE FF

P

MAC

T

PCS

270 G HORMIGA





AA BB

CC DD




272 G HORMIGA






AA BB

CC DD

EE FF






AA BB



Steatoda


Pimoa altioculata


Labulla thoracica




Novafroneta


SisicusTibioploides


HilairaLaminacauda


IslandianaErigone

Walckenaeria

Pimoa

Weintrauboa

PIMOIDAE

LINYPHIIDAE

LINYPHIOIDS

1

11

1

1

1

1

1

1

1

1

3

2

22

3

4

3

3

4

1

11

1

274 G HORMIGA





Tetragnatha

Theridiosoma

Steatoda

Pimoa rupicola

Pimoa breuili

Pimoa lihengae

Pimoa altioculata



Labulla thoracica

Microlinyphia

Linyphia

Bolyphantes

Tenuiphantes

Stemonyphantes

Haplinis

Novafroneta

Asthenargus

Gongylidiellum

Sisicus

Tibioploides

Ostearius

Leptorhoptrum

Hilaira

Laminacauda

Drepanotylus

Sciastes

Islandiana

Erigone

Walckenaeria

64

1

56

1

47

1

39

1

33

1

21

1

11

5

10

265

1

64

2

58

1

57

4

39

3

23

1

11

457

3

39

1

38

3

55

0

14

1

61

1

57

0

48

2

37

2

24

0

20

1

9

2

45

1

24

039

2

51

158

170

2

67

1

45

1

24

0

11

246

1

44

1

42

1

39

2

38

041

1

19

064

1

56

1

16

1

44

1

42

1

38

2

17

0

52

2

56

1

47

1

42

148

0

39

2

38

0

37

049

1

36

152

2

51

1

49

1

30

0

48

0

19

0

16

1

66

1

46

0

16

131

4

64

1

57

0

51

1

32

0

55

2

31

2

23

1

66

0

65

1

53

0

31

1

23

0

15

1

46

1

32

1

66

0

31

3

24

0

52

1

48

1

30

1

23

1

65

1

55

2

23

1

36

1

15

1

39

3

58

1

37

1

51

1

50

1

17

0

13

1

66

0

18

1

74

1

73

1

72

1

71

1

44

0

43

1

34

1

23

1

42

1

12

0

55

1

41

144

1

36

1

35

1

33

1

29

1

22

1

21

1

17

1

11

3

60

1

54

0

39

3

27

1

25

1

9

1

8

2

4

1

70

2

40

1

28

1

11

1

8

1

7

1

57

2

6

0

70

1

67

1

57

0

48

2

38

0

37

2

10

0

1

1

66

0

55

1

48

1

39

2

26

1

5

1

3

1

2

1

75

2

69

1

59

1

54

1

24

1

10

1

75

1

68

0

38

1

12

1

LINYPHIOIDS

PIMOIDAE

Pimoa

Weintrauboa

LINYPHIIDAE

9

2














276 G HORMIGA








u- u- u-

o-

o-






















ACKNOWLEDGEMENTS


278 G HORMIGA



REFERENCES






































APPENDIX 1


MALE PALP


























280 G HORMIGA








EPIGYNUM





mesal (2) anterior

SOMATIC MORPHOLOGY





























APPENDIX 2




Weintrauboachikunii

011111021101000-0---000-111-0---00000130-000ndash01100000


270 G HORMIGA





AA BB

CC DD




272 G HORMIGA






AA BB

CC DD

EE FF






AA BB



Steatoda


Pimoa altioculata


Labulla thoracica




Novafroneta


SisicusTibioploides


HilairaLaminacauda


IslandianaErigone

Walckenaeria

Pimoa

Weintrauboa

PIMOIDAE

LINYPHIIDAE

LINYPHIOIDS

1

11

1

1

1

1

1

1

1

1

3

2

22

3

4

3

3

4

1

11

1

274 G HORMIGA





Tetragnatha

Theridiosoma

Steatoda

Pimoa rupicola

Pimoa breuili

Pimoa lihengae

Pimoa altioculata



Labulla thoracica

Microlinyphia

Linyphia

Bolyphantes

Tenuiphantes

Stemonyphantes

Haplinis

Novafroneta

Asthenargus

Gongylidiellum

Sisicus

Tibioploides

Ostearius

Leptorhoptrum

Hilaira

Laminacauda

Drepanotylus

Sciastes

Islandiana

Erigone

Walckenaeria

64

1

56

1

47

1

39

1

33

1

21

1

11

5

10

265

1

64

2

58

1

57

4

39

3

23

1

11

457

3

39

1

38

3

55

0

14

1

61

1

57

0

48

2

37

2

24

0

20

1

9

2

45

1

24

039

2

51

158

170

2

67

1

45

1

24

0

11

246

1

44

1

42

1

39

2

38

041

1

19

064

1

56

1

16

1

44

1

42

1

38

2

17

0

52

2

56

1

47

1

42

148

0

39

2

38

0

37

049

1

36

152

2

51

1

49

1

30

0

48

0

19

0

16

1

66

1

46

0

16

131

4

64

1

57

0

51

1

32

0

55

2

31

2

23

1

66

0

65

1

53

0

31

1

23

0

15

1

46

1

32

1

66

0

31

3

24

0

52

1

48

1

30

1

23

1

65

1

55

2

23

1

36

1

15

1

39

3

58

1

37

1

51

1

50

1

17

0

13

1

66

0

18

1

74

1

73

1

72

1

71

1

44

0

43

1

34

1

23

1

42

1

12

0

55

1

41

144

1

36

1

35

1

33

1

29

1

22

1

21

1

17

1

11

3

60

1

54

0

39

3

27

1

25

1

9

1

8

2

4

1

70

2

40

1

28

1

11

1

8

1

7

1

57

2

6

0

70

1

67

1

57

0

48

2

38

0

37

2

10

0

1

1

66

0

55

1

48

1

39

2

26

1

5

1

3

1

2

1

75

2

69

1

59

1

54

1

24

1

10

1

75

1

68

0

38

1

12

1

LINYPHIOIDS

PIMOIDAE

Pimoa

Weintrauboa

LINYPHIIDAE

9

2














276 G HORMIGA








u- u- u-

o-

o-






















ACKNOWLEDGEMENTS


278 G HORMIGA



REFERENCES






































APPENDIX 1


MALE PALP


























280 G HORMIGA








EPIGYNUM





mesal (2) anterior

SOMATIC MORPHOLOGY





























APPENDIX 2




Weintrauboachikunii

011111021101000-0---000-111-0---00000130-000ndash01100000





272 G HORMIGA






AA BB

CC DD

EE FF






AA BB



Steatoda


Pimoa altioculata


Labulla thoracica




Novafroneta


SisicusTibioploides


HilairaLaminacauda


IslandianaErigone

Walckenaeria

Pimoa

Weintrauboa

PIMOIDAE

LINYPHIIDAE

LINYPHIOIDS

1

11

1

1

1

1

1

1

1

1

3

2

22

3

4

3

3

4

1

11

1

274 G HORMIGA





Tetragnatha

Theridiosoma

Steatoda

Pimoa rupicola

Pimoa breuili

Pimoa lihengae

Pimoa altioculata



Labulla thoracica

Microlinyphia

Linyphia

Bolyphantes

Tenuiphantes

Stemonyphantes

Haplinis

Novafroneta

Asthenargus

Gongylidiellum

Sisicus

Tibioploides

Ostearius

Leptorhoptrum

Hilaira

Laminacauda

Drepanotylus

Sciastes

Islandiana

Erigone

Walckenaeria

64

1

56

1

47

1

39

1

33

1

21

1

11

5

10

265

1

64

2

58

1

57

4

39

3

23

1

11

457

3

39

1

38

3

55

0

14

1

61

1

57

0

48

2

37

2

24

0

20

1

9

2

45

1

24

039

2

51

158

170

2

67

1

45

1

24

0

11

246

1

44

1

42

1

39

2

38

041

1

19

064

1

56

1

16

1

44

1

42

1

38

2

17

0

52

2

56

1

47

1

42

148

0

39

2

38

0

37

049

1

36

152

2

51

1

49

1

30

0

48

0

19

0

16

1

66

1

46

0

16

131

4

64

1

57

0

51

1

32

0

55

2

31

2

23

1

66

0

65

1

53

0

31

1

23

0

15

1

46

1

32

1

66

0

31

3

24

0

52

1

48

1

30

1

23

1

65

1

55

2

23

1

36

1

15

1

39

3

58

1

37

1

51

1

50

1

17

0

13

1

66

0

18

1

74

1

73

1

72

1

71

1

44

0

43

1

34

1

23

1

42

1

12

0

55

1

41

144

1

36

1

35

1

33

1

29

1

22

1

21

1

17

1

11

3

60

1

54

0

39

3

27

1

25

1

9

1

8

2

4

1

70

2

40

1

28

1

11

1

8

1

7

1

57

2

6

0

70

1

67

1

57

0

48

2

38

0

37

2

10

0

1

1

66

0

55

1

48

1

39

2

26

1

5

1

3

1

2

1

75

2

69

1

59

1

54

1

24

1

10

1

75

1

68

0

38

1

12

1

LINYPHIOIDS

PIMOIDAE

Pimoa

Weintrauboa

LINYPHIIDAE

9

2














276 G HORMIGA








u- u- u-

o-

o-






















ACKNOWLEDGEMENTS


278 G HORMIGA



REFERENCES






































APPENDIX 1


MALE PALP


























280 G HORMIGA








EPIGYNUM





mesal (2) anterior

SOMATIC MORPHOLOGY





























APPENDIX 2




Weintrauboachikunii

011111021101000-0---000-111-0---00000130-000ndash01100000


272 G HORMIGA






AA BB

CC DD

EE FF






AA BB



Steatoda


Pimoa altioculata


Labulla thoracica




Novafroneta


SisicusTibioploides


HilairaLaminacauda


IslandianaErigone

Walckenaeria

Pimoa

Weintrauboa

PIMOIDAE

LINYPHIIDAE

LINYPHIOIDS

1

11

1

1

1

1

1

1

1

1

3

2

22

3

4

3

3

4

1

11

1

274 G HORMIGA





Tetragnatha

Theridiosoma

Steatoda

Pimoa rupicola

Pimoa breuili

Pimoa lihengae

Pimoa altioculata



Labulla thoracica

Microlinyphia

Linyphia

Bolyphantes

Tenuiphantes

Stemonyphantes

Haplinis

Novafroneta

Asthenargus

Gongylidiellum

Sisicus

Tibioploides

Ostearius

Leptorhoptrum

Hilaira

Laminacauda

Drepanotylus

Sciastes

Islandiana

Erigone

Walckenaeria

64

1

56

1

47

1

39

1

33

1

21

1

11

5

10

265

1

64

2

58

1

57

4

39

3

23

1

11

457

3

39

1

38

3

55

0

14

1

61

1

57

0

48

2

37

2

24

0

20

1

9

2

45

1

24

039

2

51

158

170

2

67

1

45

1

24

0

11

246

1

44

1

42

1

39

2

38

041

1

19

064

1

56

1

16

1

44

1

42

1

38

2

17

0

52

2

56

1

47

1

42

148

0

39

2

38

0

37

049

1

36

152

2

51

1

49

1

30

0

48

0

19

0

16

1

66

1

46

0

16

131

4

64

1

57

0

51

1

32

0

55

2

31

2

23

1

66

0

65

1

53

0

31

1

23

0

15

1

46

1

32

1

66

0

31

3

24

0

52

1

48

1

30

1

23

1

65

1

55

2

23

1

36

1

15

1

39

3

58

1

37

1

51

1

50

1

17

0

13

1

66

0

18

1

74

1

73

1

72

1

71

1

44

0

43

1

34

1

23

1

42

1

12

0

55

1

41

144

1

36

1

35

1

33

1

29

1

22

1

21

1

17

1

11

3

60

1

54

0

39

3

27

1

25

1

9

1

8

2

4

1

70

2

40

1

28

1

11

1

8

1

7

1

57

2

6

0

70

1

67

1

57

0

48

2

38

0

37

2

10

0

1

1

66

0

55

1

48

1

39

2

26

1

5

1

3

1

2

1

75

2

69

1

59

1

54

1

24

1

10

1

75

1

68

0

38

1

12

1

LINYPHIOIDS

PIMOIDAE

Pimoa

Weintrauboa

LINYPHIIDAE

9

2














276 G HORMIGA








u- u- u-

o-

o-






















ACKNOWLEDGEMENTS


278 G HORMIGA



REFERENCES






































APPENDIX 1


MALE PALP


























280 G HORMIGA








EPIGYNUM





mesal (2) anterior

SOMATIC MORPHOLOGY





























APPENDIX 2




Weintrauboachikunii

011111021101000-0---000-111-0---00000130-000ndash01100000







AA BB



Steatoda


Pimoa altioculata


Labulla thoracica




Novafroneta


SisicusTibioploides


HilairaLaminacauda


IslandianaErigone

Walckenaeria

Pimoa

Weintrauboa

PIMOIDAE

LINYPHIIDAE

LINYPHIOIDS

1

11

1

1

1

1

1

1

1

1

3

2

22

3

4

3

3

4

1

11

1

274 G HORMIGA





Tetragnatha

Theridiosoma

Steatoda

Pimoa rupicola

Pimoa breuili

Pimoa lihengae

Pimoa altioculata



Labulla thoracica

Microlinyphia

Linyphia

Bolyphantes

Tenuiphantes

Stemonyphantes

Haplinis

Novafroneta

Asthenargus

Gongylidiellum

Sisicus

Tibioploides

Ostearius

Leptorhoptrum

Hilaira

Laminacauda

Drepanotylus

Sciastes

Islandiana

Erigone

Walckenaeria

64

1

56

1

47

1

39

1

33

1

21

1

11

5

10

265

1

64

2

58

1

57

4

39

3

23

1

11

457

3

39

1

38

3

55

0

14

1

61

1

57

0

48

2

37

2

24

0

20

1

9

2

45

1

24

039

2

51

158

170

2

67

1

45

1

24

0

11

246

1

44

1

42

1

39

2

38

041

1

19

064

1

56

1

16

1

44

1

42

1

38

2

17

0

52

2

56

1

47

1

42

148

0

39

2

38

0

37

049

1

36

152

2

51

1

49

1

30

0

48

0

19

0

16

1

66

1

46

0

16

131

4

64

1

57

0

51

1

32

0

55

2

31

2

23

1

66

0

65

1

53

0

31

1

23

0

15

1

46

1

32

1

66

0

31

3

24

0

52

1

48

1

30

1

23

1

65

1

55

2

23

1

36

1

15

1

39

3

58

1

37

1

51

1

50

1

17

0

13

1

66

0

18

1

74

1

73

1

72

1

71

1

44

0

43

1

34

1

23

1

42

1

12

0

55

1

41

144

1

36

1

35

1

33

1

29

1

22

1

21

1

17

1

11

3

60

1

54

0

39

3

27

1

25

1

9

1

8

2

4

1

70

2

40

1

28

1

11

1

8

1

7

1

57

2

6

0

70

1

67

1

57

0

48

2

38

0

37

2

10

0

1

1

66

0

55

1

48

1

39

2

26

1

5

1

3

1

2

1

75

2

69

1

59

1

54

1

24

1

10

1

75

1

68

0

38

1

12

1

LINYPHIOIDS

PIMOIDAE

Pimoa

Weintrauboa

LINYPHIIDAE

9

2














276 G HORMIGA








u- u- u-

o-

o-






















ACKNOWLEDGEMENTS


278 G HORMIGA



REFERENCES






































APPENDIX 1


MALE PALP


























280 G HORMIGA








EPIGYNUM





mesal (2) anterior

SOMATIC MORPHOLOGY





























APPENDIX 2




Weintrauboachikunii

011111021101000-0---000-111-0---00000130-000ndash01100000


274 G HORMIGA





Tetragnatha

Theridiosoma

Steatoda

Pimoa rupicola

Pimoa breuili

Pimoa lihengae

Pimoa altioculata



Labulla thoracica

Microlinyphia

Linyphia

Bolyphantes

Tenuiphantes

Stemonyphantes

Haplinis

Novafroneta

Asthenargus

Gongylidiellum

Sisicus

Tibioploides

Ostearius

Leptorhoptrum

Hilaira

Laminacauda

Drepanotylus

Sciastes

Islandiana

Erigone

Walckenaeria

64

1

56

1

47

1

39

1

33

1

21

1

11

5

10

265

1

64

2

58

1

57

4

39

3

23

1

11

457

3

39

1

38

3

55

0

14

1

61

1

57

0

48

2

37

2

24

0

20

1

9

2

45

1

24

039

2

51

158

170

2

67

1

45

1

24

0

11

246

1

44

1

42

1

39

2

38

041

1

19

064

1

56

1

16

1

44

1

42

1

38

2

17

0

52

2

56

1

47

1

42

148

0

39

2

38

0

37

049

1

36

152

2

51

1

49

1

30

0

48

0

19

0

16

1

66

1

46

0

16

131

4

64

1

57

0

51

1

32

0

55

2

31

2

23

1

66

0

65

1

53

0

31

1

23

0

15

1

46

1

32

1

66

0

31

3

24

0

52

1

48

1

30

1

23

1

65

1

55

2

23

1

36

1

15

1

39

3

58

1

37

1

51

1

50

1

17

0

13

1

66

0

18

1

74

1

73

1

72

1

71

1

44

0

43

1

34

1

23

1

42

1

12

0

55

1

41

144

1

36

1

35

1

33

1

29

1

22

1

21

1

17

1

11

3

60

1

54

0

39

3

27

1

25

1

9

1

8

2

4

1

70

2

40

1

28

1

11

1

8

1

7

1

57

2

6

0

70

1

67

1

57

0

48

2

38

0

37

2

10

0

1

1

66

0

55

1

48

1

39

2

26

1

5

1

3

1

2

1

75

2

69

1

59

1

54

1

24

1

10

1

75

1

68

0

38

1

12

1

LINYPHIOIDS

PIMOIDAE

Pimoa

Weintrauboa

LINYPHIIDAE

9

2














276 G HORMIGA








u- u- u-

o-

o-






















ACKNOWLEDGEMENTS


278 G HORMIGA



REFERENCES






































APPENDIX 1


MALE PALP


























280 G HORMIGA








EPIGYNUM





mesal (2) anterior

SOMATIC MORPHOLOGY





























APPENDIX 2




Weintrauboachikunii

011111021101000-0---000-111-0---00000130-000ndash01100000















276 G HORMIGA








u- u- u-

o-

o-






















ACKNOWLEDGEMENTS


278 G HORMIGA



REFERENCES






































APPENDIX 1


MALE PALP


























280 G HORMIGA








EPIGYNUM





mesal (2) anterior

SOMATIC MORPHOLOGY





























APPENDIX 2




Weintrauboachikunii

011111021101000-0---000-111-0---00000130-000ndash01100000


276 G HORMIGA








u- u- u-

o-

o-






















ACKNOWLEDGEMENTS


278 G HORMIGA



REFERENCES






































APPENDIX 1


MALE PALP


























280 G HORMIGA








EPIGYNUM





mesal (2) anterior

SOMATIC MORPHOLOGY





























APPENDIX 2




Weintrauboachikunii

011111021101000-0---000-111-0---00000130-000ndash01100000
















ACKNOWLEDGEMENTS


278 G HORMIGA



REFERENCES






































APPENDIX 1


MALE PALP


























280 G HORMIGA








EPIGYNUM





mesal (2) anterior

SOMATIC MORPHOLOGY





























APPENDIX 2




Weintrauboachikunii

011111021101000-0---000-111-0---00000130-000ndash01100000


278 G HORMIGA



REFERENCES






































APPENDIX 1


MALE PALP


























280 G HORMIGA








EPIGYNUM





mesal (2) anterior

SOMATIC MORPHOLOGY





























APPENDIX 2




Weintrauboachikunii

011111021101000-0---000-111-0---00000130-000ndash01100000






APPENDIX 1


MALE PALP


























280 G HORMIGA








EPIGYNUM





mesal (2) anterior

SOMATIC MORPHOLOGY





























APPENDIX 2




Weintrauboachikunii

011111021101000-0---000-111-0---00000130-000ndash01100000


280 G HORMIGA








EPIGYNUM





mesal (2) anterior

SOMATIC MORPHOLOGY





























APPENDIX 2




Weintrauboachikunii

011111021101000-0---000-111-0---00000130-000ndash01100000






APPENDIX 2




Weintrauboachikunii

011111021101000-0---000-111-0---00000130-000ndash01100000


Documents

Weintrauboa , a new genus of pimoid spiders from …spiders/content/publications...Hormiga (Hormiga, 2002; Hormiga, Arnedo & Gillespie, 2003). From a modern perspective, it looks as