European Journal of Taxonomy 184: 1-32

http://dx.doi.org/10.5852/ejt.2016.184

This work is licensed under a Creative Commons Attribution 3.0 License.

ISSN 2118-9773

WWW. europeanj oumaloftaxonomy. eu

2016 • Huber B.A. & Leh Moi Ung C.

Research article

urn:lsid:zoobank.org:pub:AC6509DC-B848-4645-BCD9-35BF0BE263Fl

The Panjange nigrifrons group in Borneo (Araneae: Pholcidae):

high diversity in Sarawak, apparent absence in Sabah

Bernhard A. HUBER '■* & Charles LEH MOI UNG ^

^ Alexander Koenig Researeh Museum of Zoology, Adenauerallee 160, 53113 Bonn, Germany.

^ Sarawak Museum, Jalan Tun Abang Haji Openg, 93566 Kuehing, Sarawak, Malaysia.

* Corresponding author: [email protected]

^ eharllmu@sarawak. gov.my

^ um:lsid:zoobank.org:author:33607F65-19BF-4DC9-94FD-4BB88CED455F

" urn:lsid:zoobank.org:author:BF48A807-815E-41E4-BCC6-F3A3FAEDBC9E

Abstract. We revise the Panjange nigrifrons group in Borneo and doeument an unexpeeted diversity

in western Sarawak forests. Five speeies oeeur within 80 km from Kuehing, eaeh speeies being known

from its type loeality only. Further speeies oeeur east until Niah, but the genus seems to be absent from

Sabah. We eontrast this with another pholeid genus (Aetana Huber, 2005), whieh is diverse in Sabah and

westward until Niah, but does not seem to oeeur in eentral and western Sarawak. Five speeies are newly

deseribed: Panjange kapit Huber, sp. nov., Panjange kubah Huber, sp. nov., Panjange niah Huber,

sp. nov., Panjangepueh Huber, sp. nov, Panjange seowi Huber, sp. nov.; Panjange tahai (Huber, 2011)

eomb. nov. is transferred from Pholcus.

Keywords. Sarawak, Sabah, Panjange, taxonomy, biogeography.

Huber B.A. & Leh Moi Ung C. 2016. The Panjange nigrifrons group in Borneo (Araneae: Pholcidae): high diversity

in Sarawak, apparent absence in Sabah. European Journal of Taxonomy 184: 1-32. http://dx.doi.org/10.5852/

eit.2016.184

Introduction

In tropical forests around the world, pholeid spiders occupy a variety of microhabitats such as leaf litter,

spaces among rocks, logs, and tree buttresses, and green leaves among the vegetation (Huber 2000,

2005). Feaf-dwelling species either attach the apex of their domed web to the underside of a leaf or

else spin a barely visible sheet of silk restricted to the (lower) surface of the leaf (Huber 2009; Huber &

Schiitte 2009; Huber et al. 2013). In Southeast Asia, leaf-dwelling pholcids are particularly diverse,

with some genera consisting entirely of leaf-dwellers (e.g., Calapnita Simon, 1892; Leptopholcus

Simon, 1893; Panjange Deeleman-Reinhold & Deeleman, 1983) and others including leaf-dwellers

and representatives in other microhabitats (e.g., Belisana Thorell, 1898; Pholcus Walckenaer, 1805)

(Deeleman-Reinhold 1986a, 1986b; Huber 2005, 2011). Some of them exhibit unique morphological or

behavioral features, such as eye stalks, extreme elongation of certain body parts, asymmetric genitalia, or

European Journal of Taxonomy 184 : 1-32 ( 2016 )

highly regular webs eonsisting of stieky and non-stieky lines in analogy to orbieularian webs (Deeleman-

Reinhold 1986a, 1986b; Huber 2005, 2011; Huber & Nuneza 2015).

Unfortunately, little is known about the relationships among leaf-dwellers and between leaf-dwellers and

their elosest relatives in other mierohabitats. In only a few eases, phylogenetie analyses of morphologieal

and moleeular data have provided a basis for reeonstrueting the direetion of evolutionary shifts among

mierohabitats. For example, the leaf-dwelling Afriean Smeringopus cylingrogaster (Simon, 1907) and

a elade of four Hispaniolan leaf-dwelling speeies in the genus Modisimus Simon, 1893 are quite elearly

derived from near-ground dwelling aneestors (Huber et al. 2010; Huber 2012; Dimitrov et al. 2013); on

the other hand, litter-dwelling Metagonia Simon, 1893 speeies in Brazil and the Afriean litter-dwelling

Pholcus kribi Huber, 2011 are apparently derived from leaf-dwelling aneestors (Dimitrov et al. 2013).

For Southeast Asian taxa, moleeular data have barely been available, and previous efforts at resolving

phylogenetie relationships using morphology have often failed to eonvineingly resolve the relevant

nodes (Huber 2011; Huber & Nuneza 2015).

The present paper foeuses on taxonomy, but is part of a major effort to resolve relationships among

Southeast Asian pholeids and to reeonstruet their evolutionary histories. It deals with Panjange, a genus

that eonsists of leaf-dwelling speeies only, but whose internal and external relationships are far from

elear. Bornean representatives of Panjange (i.e., representatives of the Panjange nigrifrons group) are

in faet more similar to speeies eurrently plaeed in Pholcus from Sumatra, the Malay Peninsula, and Sri

Lanka than to other speeies groups of Panjange in the Philippines (ineluding the type speeies) and east

of the Wallaee Line (Huber 2011; Huber & Nuneza 2015). It is not yet elear if these similarities are just

plesiomorphies or if they refleet the non-monophyly of Panjange. Preliminary moleeular data suggest

the latter (A. Valdez-Mondragon, D. Dimitrov, B.A. Huber, unpubl. data). In this eontribution we will

deal with the taxonomy of the Panjange nigrifrons group, we provide basie mierohabitat data, and we

doeument a biogeographie peeuliarity that has been known in other groups sueh as birds (e.g., Gawin

et al. 2014; Sheldon et al. 2015) but to our knowledge not in spiders: an apparent distributional limit

between Sarawak and Sabah.

Material and methods

Most of the material studied herein was eolleeted during a reeent expedition to northern Borneo (July-

August 2014). The material is eurrently deposited at Sarawak Museum, Kuehing (SMK), Malaysia

and Zoologisehes Forsehungsmuseum Alexander Koenig, Bo nn (ZFMK), Germany. Additional material

eame from the Ameriean Museum of Natural History, New York (AMNH), U.S.A. and the Netherlands

Centre for Biodiversity Naturalis, Leiden (RMNH), the Netherlands.

Speeies deseriptions are arranged as in the eladogram in Huber & Nuneza (2015). Methods and

terminology used are as in reeent revisions (Huber 2011, 2013). Measurements are in mm unless

otherwise speeifled. Eye measurements are ± 5 pm. Epigyna were eleared in warm NaOH solution

and stained with ehlorazol blaek. For SEM photos, speeimens were dried in HMDS (Brown 1993) and

photographed with a Hitaehi S-2460 seanning eleetron mieroseope. SEM data are presented within the

deseriptions but are not based on the holotype speeimens deseribed. Eoeality eoordinates are in round

braekets when eopied from labels and original publieations or when reeeived direetly from eolleetors, in

square braekets when originating from some other souree (sueh as online gazetteers, Google Earth, ete.).

The distribution maps were generated with AreMap 10.0.

The following abbreviations are used in the text:

AEE = anterior lateral eyes

AES = anterior lateral spinnerets

AME = anterior median eyes

HUBER B.A. & LEH MOIUNG C., Panjange nigrifrons group in Borneo

a.s.l. = above sea level

E/d = length/diameter

PME = posterior median eyes

In order to avoid eonfiision, Panjange is abbreviated as “Pa.” while Pholcus is never abbreviated.

Further abbreviations used only in figures are explained direetly in the figure legends.

Results

Class Araehnida Cuvier, 1812

Order Araneae Clerek, 1757

Family Pholeidae C.E. Koeh, 1851

Genus Panjange Deeleman-Reinhold & Deeleman, 1983

Panjange Deeleman-Reinhold & Deeleman, 1983: 123-124. Type speeies by original designation:

Panjange lanthana Deeleman-Reinhold & Deeleman, 1983.

Pa/ryaage - Deeleman-Reinhold 1986a: 47, fig. 6; 1986b: 220. —Huber 2001: 118-119; 2011: 108-110.

— Huber & Nuneza 2015: 3-8.

Note on species groups

Previous work on Panjange has identified three speeies groups, the nigrifrons group on Borneo (Fig. 1),

the lanthana group on the Philippines, and the widespread cavicola group (Sulawesi to northern Australia)

(Deeleman-Reinhold & Platniek 1986; Huber 2011; Huber & Nuneza 2015). From here on, we will deal

only with the nigrifrons group. A separate paper on the lanthana group has reeently been published

(Huber & Nuneza 2015), and a moleeular phylogeny of the entire genus and its elosest relatives is in

preparation (A. Valdez-Mondragon, D. Dimitrov, B.A. Huber, unpubl. data).

Panjange nigrifrons group

Diagnosis

Within Panjange, males of the nigrifrons group are easily distinguished from other speeies groups

{lanthana and cavicola groups) by presenee of distal ehelieeral apophyses (arrow in Fig. 18) and by

ventral apophysis on palpal femur (arrow in Fig. 30); also by absenee of long elongation of palpal

tarsus (eonvergently also in some speeies of lanthana group) and - females - by absent or very short

seape (relatively long in Pa. nigrifrons and Pa. kapit sp. nov.). From representatives of the very similar

Pholcus minang group by ridges ventrally on proeursus (Figs 17, 26, 34, 49; absent in Pa. tahai eomb.

nov.), by ventral apophysis on palpal femur, and possibly by wide opening of tarsal organ (figs 481

and 686 in Huber 2011). From other similar genera on Borneo {Pholcus group of genera sensu Huber

2011: Calapnita Simon, 1892; Leptopholcus Simon, 1893; Walekenaer, 1805; Uthina Simon,

1893) by eombination of male eoloration (e.g.. Figs 8, 61; male oeular area and palps blaek; similar

only in some Pholcus), by bipartite distal apophyses on male ehelieerae (Figs 35, 47; similar only in

some Calapnita), by eylindrieal rather than worm-shaped abdomen (Figs 8-15, 61-68; in eontrast to

Calapnita and Leptopholcus).

Description

Male

Measurements. Total body length ~4-5; earapaee width 0.9-1.2; leg 1 length -35-50; tibia 1

length -9-12; tibia 2/tibia 4 length 1.05-1.15; tibia 1 F/d -95-115.

European Journal of Taxonomy 184 : 1-32 ( 2016 )

Color. In life (Figs 8, 10-11, 61, 63-66) mostly pale oehre-yellow to whitish with distinetively blaek

oeular area and palps; sternum whitish; legs with dark patellae and tibia-metatarsus joints; abdomen

with darker brown to blaek marks dorsally and laterally, sometimes slightly reddish-brown.

Body. Carapaee without median furrow. Oeular area raised, eye triads on stalks of variable length

(Figs 2-7), either with or without pointed proeesses arising from near PME (Figs 2-7). AME always

absent, but sometimes with small pigment marks in AME area. Clypeus umnodified. Abdomen

eylindrieal, slightly angular above spinnerets (Figs 10, 64). Male gonopore with four epiandrous spigots

(Fig. 27); eaeh AES with large widened spigot, pointed spigot, and six eylindrieally-shaped spigots (of

varying sizes; Figs 28, 44); PMS with two spigots eaeh.

HUBER B.A. & LEH MOIUNG C., Panjange nigrifrons group in Borneo

Chelicerae. With pair of simple proximal lateral proeesses and distinetively bipartite pair of distal

apophyses (Figs 18, 35, 71); without modified hairs; without stridulatory ridges.

Paeps. Coxa unmodified; troehanter with long retrolatero-ventral apophysis (short only in Pa. nigrifrons)',

femur usually with distinetive finger-shaped ventral apophysis (e.g.. Figs 30, 70), only in Pa. pueh

sp. nov. with more eonieal ventral proeess (Fig. 17); patella triangular in lateral view; tibia with two

triehobothria; palpal tarsus sometimes with small dorsal proeess (e.g.. Fig. 70); palpal tarsal organ

eapsulate with very wide opening (ef figs 481 and 686 in Huber 2011); proeursus relatively eomplex,

only in Pa. tahai eomb. nov., with hinge dividing proximal and distal parts; parallel ridges on proeursus

usually distinet and in high number (e.g.. Figs 17, 30, 34, 49), in Pa. nigrifrons only three ridges, in Pa.

tahai eomb. nov. without ridges; proeursus sometimes with eomplex membranous proeesses (Figs 21-

23, 77-79), in some speeies with distinetive proeess in distal pit (Figs 22, 80; see also figs 478 and 683 in

Huber 2011); bulb with strong proximal selerite (e.g.. Figs 16, 29, 33), with relatively simple appendix

and weakly selerotized embolus; embolus with subterminal fringes or proeesses (Figs 24-25, 51, 75).

Fegs. Without spines, without eurved hairs, few vertieal hairs; retrolateral triehobothrium very proximal

(tibia 1: at 1.5-3% of tibia length), prolateral triehobothrium absent on tibia 1, present on other tibiae.

Tarsus 1 with -30^0 pseudosegments, very indistinet exeept distally. Tarsus 4 with single row of ventral

eomb-hairs of Pholcus-ty^Q (ef Huber & Fleekenstein 2008) (Fig. 45).

Female

Similar to male but without eye stalks; eye triads on low humps and never with pointed proeesses

(Fig. 38); ehelieerae unmodified; legs slightly shorter than in male. Epigynum weakly selerotized, with

distinet ‘knob’; either without seape (Fig. 19), with very short seape (Figs 31, 72, 84), or with medium

to long seape (Figs 36, 87). Internal genitalia with pair of pore plates of variable shape, sometimes with

very eomplex system of internal folds of unknown fimetion (Figs 59, 90, 93).

Monophyly and relationships

The eladistie analysis in Huber & Nuneza (2015) ineluded all available speeies of Panjange exeept

for Pa. tahai eomb. nov. It resolved the Pa. nigrifrons group as monophyletie, but with weak support

(only one synapomorphy: the ventral apophysis on the male palpal femur). Internal relationships

and relationships to other speeies groups in Panjange and to Pholcus also remained uneonvineing.

Preliminary moleeular data (ineluding six speeies of the Pa. nigrifrons group and six speeies of the Pa.

lanthana group; A. Valdez-Mondragon, D. Dimitrov, B.A. Huber, unpubl. data) suggest that Panjange

is not monophyletie and that the Pa. nigrifrons group is in faet more elosely related to speeies eurrently

in Pholcus than to the Pa. lanthana group. Morphologieal data partly point in the same direetion (e.g.,

the bipartite ehelieeral apophyses that oeeur in the Pa. nigrifrons group as well as in the Pholcus minang

group) but problems arise from eharaeter eonfiiet, dubious homologies of bulbal selerites, and missing

SEM data for many speeies (espeeially of the potentially elosely related Pholcus minang group).

Pholcus tahai [now Panjange tahai (Huber, 2011) eomb. nov.] from Kalimantan was not ineluded in

the eladistie analysis in Huber & Nuneza (2015). Depending on how the main bulbal proeess beside

the embolus is eoded (as unknown, as uneus, or as appendix) the speeies is either resolved as sister to

Panjange or as member of the Pa. nigrifrons group. Eaeh solution has its problems: the proeursus of

Panjange tahai eomb. nov. partly looks very different from that of representatives of the Pa. nigrifrons

group: it laeks ventral ridges and it has a hinge dividing proximal and distal parts; on the other hand,

the tip of the proeursus (eompare fig. 684 in Huber 2011 with Fig. 80 herein) and the shape of the tarsal

organ (eompare figs 481 and 685 in Huber 2011) remind strongly of other representatives of the Pa.

nigrifrons group. One further hint toward a eloser affinity with the Pa. nigrifrons group is biogeography:

Pa. tahai eomb. nov. oeeurs on Borneo like all other speeies of the Pa. nigrifrons group, while all

European Journal of Taxonomy 184 : 1-32 ( 2016 )

species of the Pholcus minang group (to which the species was previously assigned) are from Sumatra

and the Malay Peninsula (Huber 2011).

Natural history

All species observed in the field share a very similar microhabitat and web. They were consistently

found in webs among low vegetation in well preserved forests, usually at about 0.5 m above the ground

or even lower. The domed webs had a diameter of about 15-20 cm and were at their apex connected to

the underside of a leaf where the spiders rested. At some localities, large numbers of cecidomyiid files

were found hanging in the webs together with the spiders. Egg-sacs are slightly elongated and contain

approximately 20-30 eggs (e.g.. Figs 13, 62).

Composition

As construed here, the Panjange nigrifrons group now includes ten species: Panjange bako Huber, 2011;

Pa. iban Huber, 2011; Pa. kapit sp. nov.; Pa. kubah sp. nov.; Pa. niah sp. nov.; Pa. nigrifrons Deeleman-

Reinhold & Deeleman, 1983; Pa. pueh sp. nov.; Pa. sedgwicki Deeleman-Reinhold & Platnick, 1986;

Pa. seowi sp. nov.; Pa. tahai (Huber, 2011) comb. nov. Judging from known distribution patterns in

Sarawak and from the large poorly sampled forests of Kalimantan, the group is likely to contain several

times as many species. However, the absence of Panjange in northeastern Borneo may be real rather

than an artifact of poor sampling. Our own collecting at nine localities east of Niah did not produce a

single specimen of Panjange (Fig. 95; see Discussion).

Distribution

Known from Borneo only (Fig. 1).

Panjange pueh Huber, sp. nov.

um:lsid:zoobank.org:act:792671DD-F7D9-4497-BD2B-84E925032C4C

Figs 2, 8-9, 16-28, 52-54

Panjange Bor 96: Huber & Nuneza 2015: 5, 43^4.

Diagnosis

Distinguished from similar species (with pair of pointed processes arising from near PME and ventral

apophysis on male palpal femur: Pa. bako; Pa. kapit sp. nov.; Pa. niah sp. nov.; Pa. sedgwicki; Pa. tahai

comb, nov.) by details of male pedipalp (Figs 16-17; simple conical ventral femur apophysis; distinctive

prolateral membranous process on procursus; shape of procursus tip; short appendix, similar only in

Pa. niah sp. nov.); from Pa. kapit sp. nov. also by shorter palpal segments and shorter epigynal scape

(Figs 17,19). Females are difficult to distinguish from Pa. bako and Pa. niah sp. nov. (internal transversal

folds much more complex in Pa. bako, very similar in Pa. niah sp. nov.; unknown in Pa. sedgwicki).

Etymology

Named for the type locality; noun in apposition.

Type material

MAFAYSIA-BORNEO: holotype, S, Sarawak, Pueh foothills, forest along river (1.798-1.800° N,

109.708-109.718° E), 100-200 m a.s.L, 16 Jul. 2014 (B.A. Huber, S.B. Huber), ZFMK (Ar 14575).

Other material examined

MAFAYSIA-BORNEO, Sarawak: 3 SS, 5 ??, same data as holotype, ZFMK (Ar 14576-77); 1 ?,

3 juvs, in absolute ethanol, same data, ZFMK (Bor 231).

HUBER B.A. & LEH MOIUNG C., Panjange nigrifrons group in Borneo

Description

Male (holotype)

Measurements. Total body length 5.0, earapaee width 1.1. Eeg 1: 44.3 (9.9 + 0.5 + 10.2 + 21.1 + 2.6),

tibia 2: 6.3, tibia 3: 3.5, tibia 4: 5.8; tibia 1 E/d: 106. Distanee PME-PME 490 pm, diameter PME

105 pm, distanee PME-AEE ~45 pm; AME absent.

CoEOR. Carapaee pale oehre yellow to whitish, posterior mark (Fig. 8) lost in ethanol, oeular area and

elypeus dark brown (blaek in life); sternum whitish; legs oehre-orange with dark brown patellae and

tibia-metatarsus joints; abdomen oehre-gray, with blaek marks dorsally, monoehromous ventrally.

Body. Habitus as in Fig. 8; oeular area raised, eaeh triad on long stalk, with pointed straight proeess

arising from near PME (Figs 2, 18); earapaee without median furrow; elypeus unmodified; sternum

2 3 4 m

Figs 2-7. Male prosomata, oblique frontal views. 2. Panjangepueh Huber, sp. nov. (ZFMK Ar 14576).

3. Pa. niah Huber, sp. nov. (ZFMK Ar 14579). 4. Pa. kapit Huber, sp. nov. (ZFMK Ar 14583). 5. Pa.

nigrifrons Deeleman-Reinhold & Deeleman, 1983 (RMNH, Sepaku). 6. Pa. kubah Huber, sp. nov.

(ZFMK Ar 14587). 7. Pa. seowi Huber, sp. nov. (ZFMK Ar 14590).

European Journal of Taxonomy 184 : 1-32 ( 2016 )

wider than long (0.7/0.6), unmodified. Gonopore with four epiandrous spigots (Fig. 27). ALS as in

Fig. 28.

Chelicerae. Similar to elose relatives, with proximal pair of proeesses and with distinetively bipartite

distal apophyses (Fig. 18); without modified hairs; without stridulatory ridges.

Paeps. As in Figs 16-17; eoxa unmodified; troehanter with slightly eurved retrolatero-ventral apophysis

and low retrolateral hump; femur with eonieal ventral proeess; proeursus with row of about 26 ventral

ridges (Fig. 26), with distinetive prolateral membranous proeess and distal elements (Figs 21-23); bulb

with strong proximal selerite, short eurved appendix, and proximally widened and selerotized embolus

with distal proeesses and fringes (Figs 24-26).

Figs 8-15. Live speeimens. 8-9. Panjangepueh Huber, sp. nov., S and $ from Pueh. 10-14. Pa. kapit

Huber, sp. nov. from Kapit. 10-11. Adult S, 12. Penultimate instar S, 13-14. 5 with egg-sae. 15. Pa.

niah Huber, sp. nov., $ with egg-sae from Niah.

HUBER B.A. & LEH MOIUNG C., Panjange nigrifrons group in Borneo

Figs 16-20. Panjangepueh Huber, sp. nov. (ZFMK Ar 14576-77). 16-17. Eeft male palp, prolateral and

retrolateral views. 18. Male oeular area, elypeus, and ehelieerae, oblique frontal view (arrow points at

distal ehelieeral apophysis). 19-20. Cleared female genitalia, ventral and dorsal views. Abbreviations:

a = appendix; b = genital bulb; e = embolus; f = femur; p = proeursus; ps = proximal bulbal selerite; tr =

troehanter. Seale lines: 0.5 mm.

European Journal of Taxonomy 184 : 1-32 ( 2016 )

Figs 21-28. Panjange pueh Huber, sp. nov. (ZFMK Ar 14576). 21. Left proeursus and bulb, prolateral

view. 22. Left proeursus, prolateral distal view (arrow points to distinetive proeess in distal pit).

23. Prolateral membranous proeess on left proeursus. 24-25. Tip of embolus, distal views. 26. Left

proeursus and bulb, prolatero-distal view. 27. Male gonopore. 28. Male ALS. Abbreviations: a =

appendix; b = genital bulb; e = embolus; p = proeursus. Seale lines: 20 pm (28); 30 pm (25, 27); 40 pm

(23); 60 pm (22, 24); 100 pm (26); 200 pm (21).

HUBER B.A. & LEH MOIUNG C., Panjange nigrifrons group in Borneo

Eegs. Without spines and eurved hairs; few vertieal hairs; retrolateral triehobothrium on tibia 1 at

E5%; prolateral triehobothrium absent on tibia 1, present on other tibiae; tarsus 1 with > 40 indistinet

pseudosegments.

Variation

Tibia 1 in 3 other males: 8.9, 9.3, 9.6.

Female

In general similar to male but eye triads on low humps, mueh eloser together (distanee PME-PME

235 pm), without pointed proeesses. Tibia 1 in 5 females: 7.6-8.3 (mean 7.9). Epigynum weakly

selerotized plate with large weakly selerotized posterior ‘knob’ (Figs 19,52), internal areh and transversal

folds visible through eutiele; internal genitalia as in Figs 20, 53-54.

Distribution

Known from type loeality in Sarawak only (Fig. 1).

Panjange niah Huber, sp. nov.

um:lsid:zoobank.org:aet:BBDABF75-A656-4B28-AA6A-88D20FlCC113

Figs 3, 15, 29-32, 38^1, 55-57

Panjange Bor 11: Huber & Nuneza 2015: 5, 43^4.

Diagnosis

Distinguished from similar speeies (with pair of straight pointed proeesses arising from near PME and

ventral apophysis on male palpal femur: Pa. bako; Pa. kapit sp. nov.; Pa. pueh sp. nov.; Pa. sedgwicki;

Pa. tahai eomb. nov.) by shapes of proeursus and appendix (Figs 29-30; proeursus tip distinetively

bifid, similar only in Pa. bako; appendix short and eurved, similar only to Pa. pueh sp. nov.). Females

are diffieult to distinguish from Pa. bako and Pa. pueh sp. nov. (internal transversal folds mueh more

eomplex in Pa. bako, very similar in Pa. pueh sp. nov.; unknown in Pa. sedgwicki).

Etymology

Named for the type loeality; noun in apposition.

Type material

MAFAYSIA-BORNEO: holotype, S, Sarawak, Niah Cave National Park, forest near headquarters

(3.820° N, 113.763° E), 40 m a.s.l., night eolleeting, 28 Jul. 2014 (B.A. Huber, S.B. Huber), ZFMK (Ar

14578).

Other material examined

MAFAYSIA-BORNEO, Sarawak: I 6', 8 $$, I juv., same data as holotype, ZFMK (Ar 14579-80);

25 ?, same loeality, forest along main trail (3.814-3.821° N, 113.763-113.771° E), 20^0 m a.s.l.,

27 Jul. 2014 (B.A. Huber, S.B. Huber), ZFMK (Ar 14581); 3 $ $, 1 juv., same data, in absolute ethanol,

ZFMK (Bor 179).

Description

Male (holotype)

Measurements. Total body length 4.5, earapaee width 1.1. Feg 1: 41.2 (9.4 + 0.5 + 9.5 + 19.3 + 2.5),

tibia 2: 5.7, tibia 3: 3.3, tibia 4: 5.3; tibia 1 F/d: 108. Distanee PME-PME 445 pm, diameter PME

105 pm, distanee PME-AFE ~45 pm; AME absent.

European Journal of Taxonomy 184 : 1-32 ( 2016 )

Figs 29-32. Panjange niah Huber, sp. nov. (ZFMK Ar 14579-80). 29-30. Left male palp, prolateral and

retrolateral views (arrow points at ventral femur apophysis). 31-32. Cleared female genitalia, ventral

and dorsal views. Abbreviations: a = appendix; b = genital bulb; e = embolus; f = femur; p = proeursus;

ps = proximal bulbal selerite; tr = troehanter. Seale lines: 0.5 mm.

HUBER B.A. & LEH MOIUNG C., Panjange nigrifrons group in Borneo

Color. Carapace pale ochre yellow to whitish, without posterior mark, ocular area and clypeus dark

brown, with single black mark in AME area; sternum whitish; legs ochre-orange with dark brown

patellae and tibia-metatarsus joints; abdomen ochre-gray, with black marks dorsally, monochromous

ventrally.

Body. Habitus as in Pa. pueh sp. nov. (cf. Fig. 8); ocular area raised, each triad on long stalk, with pointed

straight process arising from near PME (Fig. 3); carapace without median furrow; clypeus unmodified;

sternum wider than long (0.70/0.55), unmodified.

Chelicerae. Similar to Pa. pueh sp. nov. and other close relatives (cf Fig. 18), with distinctively bipartite

distal apophyses; proximal pair of processes slightly directed towards distal; without modified hairs;

without stridulatory ridges.

Figs 33-34. Panjange Huber, sp. nov. (ZFMK Ar 14583). Feftmale palp, prolateral and retrolateral

views. Abbreviations: a = appendix; b = genital bulb; e = embolus; f = femur; p = procursus; ps =

proximal bulbal sclerite; tr = trochanter. Scale line: 1 mm.

European Journal of Taxonomy 184 : 1-32 ( 2016 )

Palps. As in Figs 29-30; coxa unmodified; trochanter with straight pointed retrolatero-ventral apophysis;

femur with curved finger-shaped ventral apophysis; procursus with row of about 12 ventral ridges, with

distinctive bifid tip; bulb with strong proximal sclerite, strongly curved short appendix, and long partly

sclerotized embolus with distinct distal fringes.

Legs. Without spines and curved hairs; few vertical hairs; retrolateral trichobothrium on tibia 1 at

2.5%; prolateral trichobothrium absent on tibia 1, present on other tibiae; tarsus 1 with > 20 indistinct

pseudosegments.

Variation

Tibia 1 in other male: 9.0.

Female

In general similar to male but eye triads on low humps, much closer together (distance PME-PME

220 pm), without pointed processes. Tibia 1 in 9 females: 6.9-7.5 (mean 7.2). Tarsus 4 comb-hairs as

in Figs 39^0. Epigynum weakly sclerotized plate with large posterior ‘knob’ (Figs 31, 41, 55), internal

anterior arch and transversal folds visible through cuticle; internal genitalia as in Figs 32, 56-57.

Figs 35-37. Panjange kapit Huber, sp. nov. (ZFMK Ar 14583-84). 35. Male ocular area, clypeus, and

chelicerae, oblique frontal view. 36-37. Cleared female genitalia, ventral and dorsal views. Scale lines:

0.5 mm.

HUBER B.A. & LEH MOIUNG C., Panjange nigrifrons group in Borneo

Figs 38^5. — 38^1. Panjange niah Huber, sp. nov. (ZFMK Ar 14580). 38. Female prosoma, frontal

view. 39-40. Comb-hairs on female tarsus 4. 41. Epigynum, ventral view. — 42-45. Panjange kapit

Huber, sp. nov. (ZFMK Ar 14583). 42. Tip of left proeursus, retrolatero-distal view. 43. Base of palpal

triehobothrium. 44. Male AES. 45. Comb-hairs on male tarsus 4. Seale lines: 10 pm (40, 43^4); 20 pm

(45); 40 pm (39); 60 pm (42); 200 pm (38, 41).

European Journal of Taxonomy 184: 1-32 (2016)

Natural history

The webs were found among the vegetation at 0.5-1 m above the ground, with the apex of the domed

sheet eonneeted to the underside of a leaf

Distribution

Known from type loeality in Sarawak only (Fig. 1).

Panjange bako Huber, 2011

Panjange bako Huber, 2011: 122-124; figs 489^93, 764, 113-11A (c??)-

Diagnosis

Distinguished from similar speeies (with ventral apophysis on palpal femur and with pointed proeesses

on eye stalks: Pa. kapit sp. nov.; Pa. niah sp. nov.; Pa. pueh sp. nov.; Pa. sedgwicki; Pa. tahai eomb.

nov.) by shapes of proeursus and appendix (proeursus tip distinetively bifid, similar only in Pa. niah

sp. nov.). Females are diffieult to distinguish from Pa. niah sp. nov. and Pa. pueh sp. nov. (internal

transversal folds mueh more eomplex in Pa. bako than in Pa. niah sp. nov.; unknown in Pa. sedgwicki).

New material examined

MALAYSIA-BORNEO, Sarawak: 2 Sf, 1 2 juvs, Bako National Park [-1.72° N, 110.45° E] (type

loeality), rainforest on slope, 28 Mar. 1985 (C.L. & P.R. Deeleman), RMNH.

Distribution

Known from type loeality in Sarawak only (Fig. 1).

Panjange Huber, sp. nov.

um:lsid:zoobank.org:aet:07838D0A-C949-4316-8B22-6892CC371A3F

Figs 4, 10-14, 33-37, 42-51, 58-60

Panjange Bor 78: Huber & Nuneza 2015: 5, 43^4.

Diagnosis

Distinguished from similar speeies (with pair of straight pointed proeesses arising from near PME and

ventral apophysis on male palpal femur: Pa. bako; Pa. niah sp. nov.; Pa. pueh sp. nov.; Pa. sedgwicki;

Pa. tahai eomb. nov.) by longer palpal segments (espeeially femur and proeursus; Figs 33-34), by

shapes of proeursus and appendix (Figs 33-34), and by longer epigynal seape (Figs 36-37; female of

Pa. sedgwicki unknown); from very similar Pa. sedgwicki (male holotype examined) by distal struetures

of proeursus, more slender male palpal tibia, and more distal ventral femur apophysis.

Etymology

Named for the type loeality; noun in apposition.

Type material

MALAYSIA-BORNEO: holotype, S, Sarawak, near Kapit, forest along river (1.937-1.944° N, 112.904-

112.910° E), 80-120 m a.s.l, 20 Jul. 2014 (B.A. Huber, S.B. Huber), ZFMK (Ar 14582).

Other material examined

MALAYSIA-BORNEO: Sarawak: 8 SS, 6 $$, 2 juvs, same data as holotype, ZFMK (Ar 14583-84;

7 SS, 5 $ $) and SMK (1 S, 1 $); 1 (?, 3 $ 1 juv., same data, in absolute ethanol, ZFMK (Bor 184).

HUBER B.A. & LEH MOIUNG C., Panjange nigrifrons group in Borneo

Figs 46-51. Panjange Huber, sp. nov. (ZFMK Ar 14583). 46. Male prosoma, frontal view. 47. Right

distal male ehelieeral apophysis. 48. Right proeursus (and bulbal proeesses), dorsal view. 49. Ridges on

left proeursus, prolateral view. 50. Tip of left appendix, prolateral view. 51. Tip of left proeursus (and

embolus), ventral view. Abbreviations: a = appendix; e = embolus; p = proeursus. Seale lines: 30 pm

(47); 60 pm (50); 100 pm (49, 51); 200 pm (48); 400 pm (46).

European Journal of Taxonomy 184: 1-32 (2016)

Figs 52-60. Female genitalia, untreated in ventral view, eleared in ventral and dorsal views. 52-

54. Panjange pueh Huber, sp. nov. (ZFMK Ar 14577). 55-57. Pa. niah Huber, sp. nov. (ZFMK Ar

14580). 58-60. Pa. Huber, sp. nov. (ZFMKAr 14584).

HUBER B.A. & LEH MOIUNG C., Panjange nigrifrons group in Borneo

Description

Male (holotype)

Measurements. Total body length 5.2, earapaee width 1.2. Eeg 1: 49.9 (11.1 + 0.5 + 11.6 + 23.8 +

2.9), tibia 2: 7.0, tibia 3: 4.1, tibia 4: 6.3; tibia 1 E/d: 110. Distanee PME-PME 520 pm, diameter PME

115 pm, distanee PME-AEE ~45 pm; AME absent.

CoEOR. Carapaee pale oehre yellow to whitish, posterior mark (Fig. 11) lost in ethanol, oeular area and

elypeus dark brown (blaek in life), without blaek spots in AME area; sternum whitish; legs oehre-orange

with dark brown patellae and tibia-metatarsus joints; abdomen oehre-gray, with blaek marks dorsally,

monoehromous ventrally.

Body. Habitus as in Figs 10-11; oeular area raised, eaeh triad on long stalk, with pointed straight proeess

arising from near PME (Fig. 4); earapaee without median furrow; elypeus unmodified; sternum wider

than long (0.75/0.55), unmodified. AES as in Fig. 44.

Cheeicerae. Similar to elose relatives, with proximal pair of proeesses and with distinetively bifid distal

apophyses (Figs 35, 46^7); without modified hairs; without stridulatory ridges.

Paeps. As in Figs 33-34; eoxa unmodified; troehanter with slightly eurved pointed retrolatero-ventral

apophysis; femur with ventral apophysis; proeursus very long, with row of about 18 ventral ridges

(Fig. 49) and isolated pateh of few prolateral ridges, with distinetive distal proeesses (Figs 48^9, 51),

apparently without proeess in distal pit (Fig. 42); bulb with strong proximal selerite, long slender appendix

with hooked tip (Fig. 50), and long partly selerotized embolus with short distal fringes (Fig. 51).

Fegs. Without spines and eurved hairs; few vertieal hairs; retrolateral triehobothrium on tibia 1 at

1.5%; prolateral triehobothrium absent on tibia 1, present on other tibiae; tarsus 1 with > 40 indistinet

pseudosegments.

Variation

Tibia 1 in 8 other males: 10.6-11.9 (mean 11.3). Some males with blaek spots in AME area; abdominal

marks variably distinet.

Female

In general similar to male but eye triads on low humps, mueh eloser together (distanee PME-PME

220 pm), without pointed proeesses. Tibia 1 in 6 females: 7.9-8.4 (mean 8.2). Epigynum weakly

selerotized plate with variably selerotized posterior ‘knob’ on extensible seape (Figs 36, 58), internal

anterior selerite and eomplex transversal folds visible through eutiele; internal genitalia as in Figs 37,

59-60.

Natural history

Webs were found among vegetation elose to the ground, with the apex of the domed sheet eonneeted

to the underside of a leaf Farge numbers of Ceeidomyiidae (deposited in ZFMK) were seen hanging

from the silk lines in most webs. Egg saes were slightly elongated, and eontained about 25-30 eggs eaeh

(n = 2).

Distribution

Known from type loeality in Sarawak only (Fig. 1).

European Journal of Taxonomy 184: 1-32 (2016)

Panjange tahai (Huber, 2011) comb. nov.

Pholcus tahai Huber, 2011: 158; figs 600-602, 616-617, 673-695 ((??)•

Diagnosis

Easily distinguished from congeners by highly distinctive procursus (very long, with hinge between

proximal and distal parts; fig. 674 in Huber 2011); females with distinctive internal genitalia (figs 617,

678 in Huber 2011) but difficult to distinguish externally from other representatives of the Pa. nigrifrons

group.

New material examined

INDONESIA-BORNEO, Central Kalimantan: 1 $, Tumbang Tahai (2°02’ S, 113°35’ E) [type locality],

primary bog forest, 3-13 Sep. 1985 (S. Djojosudharmo), RMNH; 9 SS, 12 Kaharian (2°02’ S,

113°40’ E), marshy primary forest, foliage, 2-16 Sep. 1985 (S. Djojosudharmo), RMNH; 1 (J, 4

same data but bog forest, RMNH; 5 SS, 5 $ $ (2 vials), 40 km NW Palangkaraya [-1.95° S, 113.75° E],

secondary forest, leaves, Sep. 1985 (S. Djojosudharmo), RIMNH.

Distribution

Known from three localities in Central Kalimantan (Fig. 1).

Panjange iban Huber, 2011

Figs 61-62

Panjange iban Huber, 2011: 119-121; figs 469-488, 770-772, 791-792 ((??).

Diagnosis

Distinguished from similar species (with ventral apophysis on palpal femur, without pointed processes

on eye stalks: Pa. kubah sp. nov.; Pa. nigrifrons'. Pa. seowi sp. nov.) by distinctive ventral process of

procursus (near distal end of ridges; fig. 470 in Huber 2011); females are difficult to distinguish externally

from congeners in Sarawak (distinctive internal folds arranged in anterior semicircular pattern; large

pore plates); from Pa. nigrifrons by very short scape.

New material examined

MAEAYSIA-BORNEO, Sarawak: 2 SS, 7 $$, Semengoh Arboretum, Masing Trail (1.397-1.399°

N, 110.317-110.322° E), 60-80 m a.s.L, 17 July 2014 (B.A. Huber), ZFMK (Ar 14585); 2 ?$, 2 juvs,

same data, in absolute ethanol, ZFMK (Bor 227); 6 SS, 3 $$, same locality, Apr. 1985 (C.E. & P.R

Deeleman), RMNH.

Natural history

The webs had a diameter of about 10-20 cm and were found among vegetation in primary forest close

to the ground where the domed sheets were attached to the undersides of leaves.

Distribution

Known from type locality in Sarawak only (Fig. 1).

Panjange nigrifrons Deeleman-Reinhold & Deeleman, 1983

Figs 5, 86-88

Panjange nigrifrons Deeleman-Reinhold & Deeleman, 1983: 126-128; figs 10-16 ((??)•

HUBER B.A. & LEH MOIUNG C., Panjange nigrifrons group in Borneo

Diagnosis

Distinguished from similar speeies (with ventral apophysis on palpal femur, without pointed proeesses

on eye stalks: Pa. iban; Pa. kubah sp. nov.; Pa. seowi sp. nov.) by low number of ridges ventrally on

proeursus (only three; fig. 13 in Deeleman-Reinhold & Deeleman 1983); by short male palpal troehanter

apophysis (fig. 15 in Deeleman-Reinhold & Deeleman 1983), and by distinetive female genitalia (Figs

86-88; internal lateral struetures visible through eutiele; long seape).

New material examined

INDONESIA-BORNEO, East Kalimantan: 2 33, 2 ??, Sepaku [-0.91° S, 116.77° E], 40 km N of

Balikpapan (type loeality), primary forest, 20-21 Jul. 1982 (eolleetor unknown), RMNH.

Figs 61-68. Five speeimens. 61-62. Pa. iban Huber, 2011, 3 and $ with egg-sae from Semengoh.

63-64. Pa. kubah Huber, sp. nov., 3 from Kubah. 65-68. Pa. seowi Huber, sp. nov., 3 , ?, and $ with

egg-sae from Pemissen.

European Journal of Taxonomy 184: 1-32 (2016)

Description - amendments

Male distal cheliceral apophyses elearly bipartite as in other speeies deseribed herein (this was eorreetly

noted in the original deseription but not eorreetly drawn in fig. 11). Appendix with very distinet and

unique poeket at -60% of its length. Tibia 1 L/d: 100; tibia 2/tibia 4 length: 1.09 (not 1.33 as in original

deseription); distanee PME-PME: 420 pm; retrolateral triehobothrium at 2%.

Distribution

Known from two loealities in Kalimantan only (Deeleman-Reinhold & Deeleman 1983; Fig. 1).

Panjange kubah Huber, sp. nov.

um:lsid:zoobanlvorg:aet:CF978A33-4706-4705-8245-86776DB7E34C

Figs 6, 63-64, 69-81, 89-91

Panjange Bor 109: Huber & Nuneza 2015: 5, 43^4.

Diagnosis

Easily distinguished from most eongeners by distinetive transversal selerite on proeursus (Fig. 70);

from very similar Pa. seowi sp. nov. by other details of proeursus (unique prolateral proeess. Figs 69,

77-78; distal element longer. Fig. 70; absenee of prolateral ridges on proeursus. Fig. 69). Females are

easily distinguished from Pa. nigrifrons by short seape but diffieult to distinguish externally from other

eongeners in Sarawak (distinetive shape of pore plates and pair of median selerites; similar only in Pa.

seowi sp. nov.).

Etymology

Named for the type loeality; noun in apposition.

Type material

MAEAYSIA-BORNEO: holotype, S, Sarawak, Kubah National Park, along Waterfall Trail (1.596-

1.606° N, 110.180-110.187° E), 200-300 m a.s.L, 12 Jul. 2014 (B.A. Huber, S.B. Huber), ZFMK (Ar

14586).

Other material examined

MAEAYSIA-BORNEO, Sarawak: 2 SS, 7 ??, same data as holotype, ZFMK (Ar 14587-88); 2

2 juvs, same data, in absolute ethanol, ZFMK (Bor 189).

Description

Male (holotype)

Measurements. Total body length 4.5, earapaee width 1.2. Eeg 1: 42.0 (9.3 + 0.5 + 9.6 + 20.1 + 2.5),

tibia 2: 6.1, tibia 3: 3.6, tibia 4: 5.6; tibia 1 E/d: 99. Distanee PME-PME 445 pm, diameter PME 105 pm,

distanee PME-AEE -45 pm; AME absent.

CoEOR. Carapaee pale oehre yellow to whitish, posterior mark (Fig. 63) lost in ethanol, oeular area and

elypeus dark brown (blaek in life), without blaek marks in AME area; sternum whitish; legs oehre-

orange with dark brown patellae and tibia-metatarsus joints; abdomen oehre-gray, with blaek marks

dorsally, monoehromous ventrally.

Body. Habitus as in Figs 63-64; oeular area raised, eaeh triad on medium long stalk, without pointed

proeess (ef Fig. 6); earapaee without median furrow; elypeus unmodified; sternum wider than long

(0.75/0.60), unmodified.

HUBER B.A. & LEH MOIUNG C., Panjange nigrifrons group in Borneo

■7::

V ' \ \1

\ \ 1

J\w-. a

\ ,\VvV

Figs 69-73. Panjange kubah Huber, sp. nov. (ZFMK Ar 14587-88). 69-70. Eeft male palp, prolateral and

retrolateral views (arrows point at distinetive proeesses). 71. Male oeular area, elypeus, and ehelieerae,

oblique frontal view. 72-73. Cleared female genitalia, ventral and dorsal views. Seale lines: 0.5 mm.

European Journal of Taxonomy 184: 1-32 (2016)

Figs 74-81. Panjange kubah Huber, sp. nov. (ZFMK Ar 14587-88). 74. Left bulb, prolateral (slightly

distal) view. 75. Tip of left embolus. 76. Epigynum, ventral view. 77-78. Left proeursus, dorso-distal

and prolateral views (arrows point at distinetive membranous proeess). 79. Membranous proeess of

left proeursus. 80. Tip of left proeursus, distal view. 81. Female ALS. Abbreviations: a = appendix;

b = genital bulb; e = embolus. Seale lines: 10 pm (81); 20 pm (80); 40 pm (79); 80 pm (75); 100 pm

(77-78); 200 pm (74, 76).

HUBER B.A. & LEH MOIUNG C., Panjange nigrifrons group in Borneo

Chelicerae. Similar to close relatives, with proximal pair of processes and distinctively bipartite distal

apophyses (Fig. 71); without modified hairs; without stridulatory ridges.

Paeps. As in Figs 69-70; coxa unmodified; trochanter with slightly curved pointed retrolatero-ventral

apophysis; femur with curved finger-shaped ventral apophysis; procursus with row of about 17 ventral

ridges, with distinctive transversal sclerite and long prolateral process close to large fiat distal element

(Figs 77-78), with two distinct spiny processes in distal pit (Fig. 80); bulb with strong proximal sclerite,

slightly curved appendix, and long partly sclerotized embolus with distinct distal fringes (Figs 74-75).

Fegs. Without spines and curved hairs; few vertical hairs; retrolateral trichobothrium on tibia 1 at 2%;

prolateral trichobothrium absent on tibia 1, present on other tibiae; tarsus 1 with many pseudosegments

but only ~10 distally visible in dissecting microscope.

Variation

Tibia 1 in 2 other males: 9.1, 9.3. One male with irregular black marks in AME area (Fig. 6).

Female

In general similar to male but eye triads on low humps, much closer together (distance PME-PME

230 pm). Tibia 1 in 6 females: 6.9-7.9 (mean 7.4). Epigynum weakly sclerotized plate with variably

large and variably sclerotized posterior ‘knob’ (Figs 72, 76, 89), internal arch and complex transversal

folds visible through cuticle; internal genitalia as in Figs 73, 90-91.

Natural history

Most specimens were found in a very limited area close to a waterfall. The domed webs had a diameter

of about 15-20 cm and in each case the apex of the dome was attached to the underside of a leaf where

the spider rested.

Distribution

Known from type locality in Sarawak only (Fig. 1).

Panjange seowi Huber, sp. nov.

urn:lsid:zoobank.org:act:96424104-A78F-41D3-AA64-65DA79138D21

Figs 7, 65-68, 82-85, 92-94

Panjange Bor 100: Huber & Nuneza 2015: 5, 43^4.

Diagnosis

Easily distinguished from most congeners by distinctive transversal sclerite on procursus (Fig. 83); from

the very similar Pa. kubah sp. nov. by other details of procursus (prolateral ridges on procursus. Fig. 82;

small transparent process close to transversal sclerite. Fig. 82; absence of long membranous prolateral

process; distal element shorter). Females are easily distinguished from Pa. nigrifrons by short scape but

difficult to distinguish externally from other congeners in Sarawak (distinctive shape of pore plates and

pair of median sclerites; similar only in Pa. kubah sp. nov.).

Etymology

Named for Francis Seow-Choen from Singapore, surgeon and expert on stick insects.

European Journal of Taxonomy 184: 1-32 (2016)

Figs 82-85. Panjange seowi Huber, sp. nov. (ZFMK Ar 14590-91). 82-83. Left proeursus, prolateral

and retrolateral views (arrows point at distinetive proeesses). 84-85. Cleared female genitalia, ventral

and dorsal views. Seale lines: 0.5 mm.

HUBER B.A. & LEH MOIUNG C., Panjange nigrifrons group in Borneo

Type material

MAEAYSIA-BORNEO: holotype, S, Sarawak, Mt. Penrissen near Borneo Highland Resort, Ma Gaseng

Trail (1.124-1.127° N, 110.217° E), 870-930 m a.s.L, 14 Jul. 2014 (B.A. Huber, S.B. Huber), ZFMK

(Ar 14589).

Other material examined

MAEAYSIA-BORNEO, Sarawak: 1 (?, 4 $5^ 1 same data as holotype, ZFMK (Ar 14590-91);

3 $ 1 juv., same data, in absolute ethanol, ZFMK (Bor 217).

Description

Male (holotype)

Measurements. Total body length 4.3, earapaee width 1.1. Eeg 1: 46.3 (10.4 + 0.5 + 10.5 + 22.1 + 2.8),

tibia 2: 6.5, tibia 3: 3.7, tibia 4: 5.7; tibia 1 E/d: 114. Distanee PME-PME 375 pm, diameter PME 105

pm, distanee PME-AEE ~45 pm; AME absent.

CoEOR. Carapaee pale oehre yellow to whitish, posterior mark (Fig. 65) lost in ethanol, oeular area and

elypeus dark brown (blaek in life), with pair of blaek marks in AME area; sternum whitish; legs oehre-

orange with dark brown patellae and tibia-metatarsus joints; abdomen oehre-gray, with blaek marks

dorsally, monoehromous ventrally.

Body. Habitus as in Figs 65-66; oeular area raised, eaeh triad on medium long stalk, without pointed

proeess (Fig. 7); earapaee without median furrow; elypeus unmodified; sternum wider than long

(0.70/0.60), unmodified.

Cheeicerae. Similar to elose relatives, with proximal pair of proeesses and distinetively bipartite distal

apophyses (ef Fig. 71); without modified hairs; without stridulatory ridges.

Paeps. In general very similar to Pa. kubah sp. nov. (ef Figs 69-70), but troehanter apophysis slightly

shorter and distally thinner/more pointed; proeursus with distinetive prolateral ridges (Fig. 82), with

small transparent proeess elose to transversal selerite (Fig. 82), without long membranous prolateral

proeess, entire distal element shorter (Fig. 83); bulbal proeesses apparently not distinguishable from

those in Pa. kubah sp. nov.

Fegs. Without spines and eurved hairs; few vertieal hairs; retrolateral triehobothrium on tibia 1 at 2%;

prolateral triehobothrium absent on tibia 1, present on other tibiae; tarsus 1 > 40 pseudosegments,

proximally very indistinet.

Variation

Tibia 1 in other male: 10.3; other male paler and transversal selerite of proeursus without proximal

proeess.

Female

In general similar to male but eye triads on low humps, mueh eloser together (distanee PME-PME

230 pm). Tibia 1 in 3 females: 7.8, 8.4, 8.8. Epigynum weakly selerotized plate with large weakly

selerotized posterior ‘knob’ (Figs 84, 92), internal areh and transversal folds visible through eutiele;

internal genitalia as in Figs 85, 93-94.

European Journal of Taxonomy 184 : 1-32 ( 2016 )

Figs 86-94. Female genitalia, untreated in ventral view, eleared in ventral and dorsal views. 86-

88. Panjange nigrifrons Deeleman-Reinhold & Deeleman, 1983 (RMNH, Sepaku). 89-91. Pa. kubah

Huber, sp. nov. (ZFMK Ar 14588). 92-94. Pa. seowi Huber, sp. nov. (ZFMK Ar 14591).

HUBER B.A. & LEH MOIUNG C., Panjange nigrifrons group in Borneo

Natural history

The domed webs were found under green leaves among vegetation, usually about 0.5 m above the

ground.

Distribution

Known from type loeality in Sarawak only (Fig. 1).

Discussion

We doeument high levels of diversity and endemism of Panjange in western Sarawak, but it remains

unelear to whieh extent this ean be extrapolated to other, mostly very poorly sampled regions of Borneo.

Panjange does oeeur in Kalimantan, but it might be signifieantly less diverse there than in western

Sarawak. A large part of Kalimantan is (or was originally) eovered by peat swamp forest that in general

is thought to have distinetive but limited diversity (e.g., Mohamedsaid & Holloway 1999; Posa et al.

2011). However, at least northern West Kalimantan is expeeted to have similar high diversity and

endemism as western Sarawak, but this region seems to be among the most poorly explored regions

worldwide, as far as spiders are eoneemed.

An extrapolation from Sarawak to unexplored regions in Borneo is partieularly problematie in light of

our surprising finding that Panjange seems to be absent from Sabah. Absenee is of eourse diffieult to

doeument but the available data strongly suggest a real pattern rather than an artifaet of sampling. First,

we sampled with the same methodology and diligenee at six loealities west of Niah (sea level to 930 m

a.s.l.) as we did at nine loealities east of Niah (sea level to 1650 m a.s.L). Panjange was found at all six

Fig. 95. Known distributions of the gQnQm Panjange Deeleman-Reinhold & Deeleman, 1983 md Aetana

Huber, 2005 in northern Borneo, showing the distinetness of western and eastern regions (distribution of

Aetana from Huber et al. 2015).

European Journal of Taxonomy 184 : 1-32 ( 2016 )

localities west of Niah (and in Niah), but in none of the nine loealities east of Niah. Seeond, the same

pattern is apparent in the large eolleetions made by C. Deeleman-Reinhold sinee the 1980s in Sarawak

(Bako, Semengoh) and Sabah (Mt. Kinabalu, Poring, Sepilok) (Pholeidae mostly published in Huber

2011). Third, extensive more reeent quantitative sampling by A. Floren in Sabah (Croeker Range, Mt.

Kinabalu, Tawau) also did not produee a single speeimen of Panjange (Huber 2011 and B.A. Huber,

unpubl. data).

Further evidenee suggesting a real pattern eomes from another pholeid genus, Aetana Huber, 2005.

This genus is not partieularly elose to Panjange (same subfamily but not part of the Pholcus group of

genera like Panjange\ Huber 2011) but is relevant in this eontext beeause it shows the exaetly opposite

distribution in northern Borneo. We found various speeies of Aetana at eight of nine loealities east

of Niah (and in Niah), but in none of the six loealities west of Niah (Huber et al. 2015). Figure 95

summarizes eurrent knowledge about the distributions of Panjange and Aetana in northern Borneo,

eombining all known reeords. Obviously, large sampling gaps remain, and a meaningful interpretation

will have to build on a mueh more intense sampling ineluding Kalimantan. However, data on better

studied organisms sueh as birds have also supported a biogeographieal divide between Sabah and

Sarawak despite eontinuous rainforest (Gawin et al 2014; Sheldon et al. 2015). These authors mainly

suggest historieal reasons for this pattern, sueh as early Pleistoeene rainforest refugia in Sabah versus

dry habitats in Sarawak (at that time part of eentral Sundaland). Whether the same explanations apply to

spiders as to birds remains to be established.

Acknowledgements

We thank the Sarawak Museum Department for assistanee in field logisties and Samuel B. Huber for help in

the field. The Natural History Museum in Kuehing kindly offered its faeilities as the base of the field work.

Permits were reeeived from the Forest Department, Sarawak (Permit No. NCCD.907.4.4(JLD.10)-211,

Park Permit No. 174/2014). We thank two anonymous reviewers for eommenting on a previous version

of the manuseript. The German Researeh Foundation provided finaneial support (DFG Projeet HU

980/11-1).

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