European Journal of Taxonomy 190: 1-55
http://dx.doi.org/10.5852/ejt.2016.190

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

ISSN 2118-9773
www. europeanj ournaloftaxonomy. eu
2016 • Huber B.A. et al.

Monograph

urn:lsid:zoobank.org:pub:BE92596B-62D9-46CD-8486-CF6B36C640Bl

The Southeast Asian Pholcus halabala species group (Araneae, Pholcidae):
new data from field observations and ultrastructure

Bernhard A. HUBER 1 *, Booppa PETCHARAD 2 . Charles LEH MOI UNG 3 ,

Joseph K.H. KOH 4 & Amir R.M. GHAZALI 5

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

2 Department of Biology, Faculty of Science, Prince of Songkla University, Hat Yai, Songkhla 90112, Thailand.

3 Sarawak Museum, Jalan Tun Abang Haji Openg, 93566 Kuching, Sarawak, Malaysia.

4 National Biodiversity Centre, National Parks Board, IB Cluny Road, Singapore 259598, Singapore.

5 Centre for Research and Consultancy, Unitar International University, Malaysia.

* Corresponding author: b.huber@zfmk.de

2 Email: zigzagargiope@vahoo.com

3 Email: charl 1 m u@sarawak.gov.my

4 Email: iosephkhkoh@gmail.com

5 Email: amir.ridhwan@gmail.com

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

2 um:lsid:zoobank.org:author:E1480A4E-3FA8-441C-A803-515B8AE7860D

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

4 urn:lsid:zoobank.org:author:DA9CA194-2A90-41BD-8109-D8E536F23C0C

5 urn:lsid:zoobank.org:author:EA038837-AA7B-49F0-8012-2B23C36F1605

Abstract. The Southeast Asian Pholcus halabala species group is revised and re-delimited, based mainly on
field observations (life color pattern, web design, position of egg-sac when carried by female, microhabitat)
and ultrastructure (silk spigots, modifications of male cheliceral apophyses). The core group includes six leaf¬
dwelling species that have distinctive color patterns in life specimens (black and white or yellowish abdominal
marks, dark pattern on posterior half of carapace) and build round to oval silk platforms on the undersides
of leaves. Seven further species are tentatively assigned to the group pending further study. Several species
originally assigned to the Pholcus halabala group are transferred to three newly proposed species groups, the
Ph, krabi, Ph. buatong , and Ph. andulau groups. Nine species are newly described, four in the Ph. halabala
group (Ph. khaolek Huber, sp. nov.; Ph. kuhapimuk Huber, sp. nov.; Ph. fifing Huber, sp. nov.; Ph. ubin Huber,
sp. nov.); three in the Ph. krabi group (Ph. kipungit Huber, sp. nov.; Ph. krabi Huber, sp. nov.; Ph. narathiwat
Huber, sp. nov.); one in the Ph. buatong group (Ph. buatong Huber, sp. nov.); and one in the Ph andulau group
(Ph. lambir Huber, sp. nov.). The females of Ph. satun Huber, 2011 and Ph. schwendingeri Huber, 2011 (both
members of the buatong group) are newly described.

Keywords. Pholcus , taxonomy, web design, microhabitat, ultrastructure.

Huber B.A., PetcharadB., Leh Moi Ung C., KohJ.K.H. & GhazaliA.R.M. 2016. The Southeast Asian Pholcus
halabala species group (Araneae, Pholcidae): new data from field observations and ultrastructure. European
Journal of Taxonomy 190: 1-55. http://dx.doi.org/10.5852/ejt.2016.19Q

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European Journal of Taxonomy 190 : 1-55 ( 2016 )

Introduction

With currently over 300 described species, the genus Pholcus Walckenaer, 1805 is by far the most
species-rich genus in the Pholcidae (World Spider Catalog 2015). It includes some widespread and
well-studied synanthropic species such as Ph. phalangioides (Fuesslin, 1775), but most of its diversity is
concentrated in the Old World tropics and subtropics. For more than 200 years, almost any large pholcid
with eight eyes and cylindrical abdomen resembling Ph. phalangioides was assigned to this genus. As
a result, a first attempt at revising the genus (Huber 2011) revealed some evidence for non-monophyly,
with several genera (especially those containing leaf-dwelling taxa in Southeast Asia) apparently nested
within a paraphyletic Pholcus. However, no nomenclature changes were formally proposed, partly
because taxonomic sampling in the cladistic analysis seemed too preliminary, partly because some of
the 29 species groups proposed seemed non-monophyletic themselves. In fact, the main focus of that
revision was a first tentative division of this previously chaotic genus into more manageable units, which
were explicitly proposed as “operational species groups”.

One of the most problematic species groups in Huber (2011) was the Southeast Asian Pholcus halabala
group. Only a core group of three species (Ph. halabala Huber, 2011; Ph. sabah Huber, 2011; Ph. sepaku
Huber, 2011) was supported by fairly convincing synapomorphies. Eight further species were tentatively
assigned to the group for lack of a better solution. The present paper is a first step towards resolving
relationships among these species. Our main focus is on documenting some behavioral (web design,
egg-sac position), ecological (microhabitat) and ultrastructural (spigots, male cheliceral apophyses)
characters that are likely to be informative in future cladistic re-analyses of the genus.

Material and methods

Most of the material studied herein was collected during recent expeditions to northern Borneo (Juf-
Aug. 2014) and the Malay Peninsula (Feb.-Mar. 2015). The material is currently deposited at the Princess
Maha Chakri Sirinthorn Natural History Museum, Prince of Songkla University, Hat Yai (PSUZC),
Thailand; the Sarawak Museum, Kuching (SMK), Malaysia; and the Zoologisches Forschungsmuseum
Alexander Koenig, Bonn (ZFMK), Germany. Additional material came from the Netherlands Centre for
Biodiversity Naturalis, Teiden (RMNH).

Methods and terminology used are as in Huber (2011). Measurements are in mm unless otherwise
specified. Eye measurements are approx. ± 5 pm. Epigyna were cleared in a warm NaOH solution and
stained with chlorazol black. For SEM photos, specimens were dried in hexamethyldisilazane (HMDS)
(Brown 1993) and photographed with a Hitachi S-2460 scanning electron microscope. SEM data are
presented within the descriptions but are not based on the holotype specimens described. Locality
coordinates are in round brackets when copied from labels and original publications or when received
directly from collectors, in square brackets when originating from some other source (such as online
gazetteers, Google Earth, etc.). Distribution maps were generated with ArcMap 10.0.

The following abbreviations are used in the text:

ALE = anterior lateral eyes
ALS = anterior lateral spinnerets
AME = anterior median eyes
a.s.l. = above sea level
L/d = length/diameter
PME = posterior median eyes.

In order to avoid confusion, Pholcus is abbreviated as “Ph.” while Panjange is never abbreviated.
Abbreviations used only in figures are explained directly in the figure legends.

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HUBER B.A. et al., Pholcus halabala group in Southeast Asia

Results

Class Arachnida Cuvier, 1812
Order Araneae Clerck, 1757
Family Pholcidae C.L. Koch, 1851

Pholcus Walckenaer, 1805

Pholcus Walckenaer, 1805: 80. Type species by (assumed) monotypy: Araneaphalangioides ¥uesshn, 1775.
Pholcus- Huber 2011: 124-126.

Pholcus halabala species group

Diagnosis

The core group of six species (see below) includes medium-sized, long-legged spiders (body length ~3-4,
male leg 1 length —30^40); distinguished from other species groups in Pholcus by the combination of the
following characters: elongate abdomen pointed dorso-posteriorly, with distinctive dorsal pattern of black
and whitish or yellowish marks in life specimens (Figs 1-16; similar only in Ph. sudhami Huber, 2011,
cf. Figs 54-56); eight eyes; male ocular area with conspicuous modified hairs (setae), which may appear
as stiff bristles or stout curved spines, or both (Figs 19, 23, 43); male chelicerae with proximal and distal
apophyses, distal apophyses with two cone-shaped teeth (modified hairs) each (Fig. 28); male bulb with
uncus and appendix; procursus with distinctive dorsal flap (Fig. 35; absent in Ph. erawan Huber, 2011);
epigynum weakly sclerotized, with ‘knob’.

Description - amendments

The original description of the core group (Huber 2011) is still largely valid. The following can be
added: male eye triads on low humps (Figs 18, 20,43; i.e., not on short stalks as seen in Ph. krabi group)
and only slightly wider apart than female eye triads (1.2-1.5 x); dorsal elongation of male palpal tarsus
usually short and conical (asterisk in Fig. 30), long in Ph. erawan Huber, 2011 (fig. 1467 in Huber
2011); procursus usually with dorsal flap, absent in Ph. erawan ; tibia 1 in males ~6-9, in females ~6-8;
male gonopore with four epiandrous spigots (Figs 26, 44; confirmed in Ph. halabala and Ph. erawan );
AFS with eight spigots each (one widened, one pointed, and six smaller cylindrically shaped spigots
of varying sizes, Figs 25, 45; confirmed in Ph. halabala and Ph. erawan ); web consists mostly of flat
round platform closely attached to underside of leaf; web sometimes with large numbers of small silk
tufts (Figs 4, 16; confirmed for Ph. halabala , Ph. erawan and Ph. sepaku); female holds egg-sac under
body rather than in front of it (Figs 3, 5-6, 16; confirmed for Ph. halabala , Ph. erawan and Ph. sepaku).

Composition

The 13 species assigned to this group are here divided into two operational sub-groups: a core-group of
six species for which the evidence for monophyly is considered to be strong (Ph. erawan Huber, 2011;
Ph. halabala Huber, 2011; Ph. lintang Huber, sp. nov.; Ph. sabah Huber, 2011; Ph. sepaku Huber, 2011;
Ph. ubin sp. nov.); and a sub-group of seven species whose assignment is tentative [Ph. elongatus (Yin &
Wang, 1981); Ph. exceptus Tong & Fi, 2009; Ph. khaolek Huber, sp. nov.; Ph. kuhapimuk Huber, sp. nov.;
Ph. pakse Huber, 2011; Ph. pyu Huber, 2011; Ph. sudhami Huber, 2011].

The composition of the group changes as follows: to the original core group consisting of three species
(Ph. halabala ; Ph. sabah ; Ph. sepaku ) we add two newly described species (Ph. ubin Huber, sp. nov.;
Ph. lintang sp. nov.) as well as Ph. erawan. The latter species was originally part of another problematic
species group (Pholcus quinquenotatus group; Huber 2011: 290). It lacks the distinctive dorsal flap

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European Journal of Taxonomy 190 : 1-55 ( 2016 )

on the procursus that is present in the other five species of the core group, and the hairs in the male
ocular area are not particularly strong, but it is moved into the core group because (1) it builds the
same distinctive web closely attached to the leaf surface as the five other species; (2) webs are provided
with facultative silk tufts like in Ph. halabala and Ph. sepaku (otherwise only known in the African
genus Smeringopus Simon, 1890; Huber 2012); (3) females carry the egg-sac under the body like Ph.
halabala and Ph. sepaku , which is unique among Pholcidae; (4) life specimens have an almost identical
distinctive pattern of large whitish to yellowish and black spots on the abdomen, and on the carapace
a pair of V-marks; and (5) the procursus is provided with a distinctive retrolateral distal pocket also
present in Ph. halabala (Figs 21-22, 41). Preliminary molecular data (A. Valdez-Mondragon, B.A.

Figs 1-8. Live specimens, Pholcus halabala Huber, 2011 (1-6) and Ph. ubin Huber, sp. nov. (7-8). 1. $
from MacRitchie, Singapore. 2. $ from Gunung Jerai, Malaysia. 3. $ with egg-sac from Gunung Liang,
Malaysia. 4. Juvenile in web with silk tufts, Upper Selatar, Singapore. 5. $ with egg-sac from HalaBala,
Thailand. 6. $ with egg-sac from Ulu Dong, Malaysia. 7-8. $ and $ from Pulau Ubin, Singapore.

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HUBER B.A. et al., Pholcus halabala group in Southeast Asia

Huber & D. Dimitrov unpublished data), including all species of the core group except Ph. sepaku (i.e.,
also Ph. erawan), strongly support the monophyly of this group. The RMNH has three further species
of the core group from Sabah (Fig. 17), but the specimens are poorly preserved and for this reason not
formally described here.

In addition to this core group, we include in the Ph. halabala group two further species that were
previously part of the Pholcus quinquenotatus group (Ph. sudhami Huber, 2011; Ph. pakse Huber, 2011)
as well as two newly described species (Ph. kuhapimuk sp. nov.; Ph. khaolek sp. nov.). Again, preliminary

Figs 9-16. Live specimens, Pholcus sabah Huber, 2011 (9-10), Ph. lintang Huber, sp. nov. (11-12) and
Ph. erawan Huber, 2011 (13-16). 9. S from Poring, Sabah. 10. S from Sepilok, Sabah. 11-12. S and $
from Bako, Sarawak. 13. S from Khao Nan, Thailand. 14. Male from Erawan, Thailand. 15-16. S and
$ with egg-sac from Gunung Jerai, Malaysia.

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European Journal of Taxonomy 190 : 1-55 ( 2016 )

molecular data (A. Valdez-Mondragon, B.A. Huber & D. Dimitrov unpublished data) strongly support a
close relationship of these four species with the core group, and at least Ph. sudhami and Ph. pakse share
some morphological similarities with the core group (carapace and abdomen pattern, male eye triads on
low humps, large AME, male cheliceral apophyses with modified hairs), but no putative morphological
synapomorphy is known that would support this close relationship. The microhabitat of these four
species differs from the core group (rocks and tree roots rather than foliage); egg-sacs are carried in
front of the body as in typical pholcids; webs were never seen to be provided with silk tufts. In addition,
males of Ph. sudhami and Ph. pakse lack stronger hairs in the ocular area, and both species lack the
bulbal appendix.

Three species originally assigned to the halabala group are kept in the group simply for the lack of
a better solution: Ph. pyu Huber, 2011; Ph. elongatus (Yin & Wang, 1981); and Ph. exceptus Tong &
Li, 2009. Data on natural history and ultrastructure are not available for any of them, nor is any of
them included in our preliminary molecular analysis (A. Valdez-Mondragon, B.A. Huber & D. Dimitrov
unpublished data). We speculate that Ph. pyu may in fact belong to this group, but the other two species are
quite certainly misplaced. Pholcus elongatus lacks AME; males in this species have short eye stalks and

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HUBER B.A. et al., Pholcus halabala group in Southeast Asia

pointed cheliceral apophyses without modified hairs; females have a long epigynal scape. We speculate
that the closest relatives of Ph. elongatus are the Taiwanese species Ph. pingtung Huber & Dimitrov, 2014
and Ph. chengpoi Huber & Dimitrov, 2014 (six eyes; similar cheliceral apophyses; scape directed toward
anterior). Pholcus exceptus is a mysterious species with entirely reduced distal cheliceral apophyses.

Finally, five species originally included in the Pholcus halabala group are transferred to new species
groups (see below): Ph. andulau Huber, 2011; Ph. chiangdao Huber, 2011; Ph. khene Huber, 2011; Ph.
kinabalu Huber, 2011; and Ph. satun Huber, 2011.

Natural history

All six species of the core group were observed in the field. At most localities, abundances seemed very
low. Surprisingly, several species were partly or exclusively found in degraded forests. All specimens
seen during the day were tightly pressed against the undersides of mostly large leaves. Webs consisted
mainly of round platforms with a diameter of about 10 cm; most of the platform was closely attached
to the leaf surface. Small silk tufts (Figs 4, 16) were observed in some webs of three species (Ph.
halabala ; Ph. erawan ; Ph. sepaku), but may occur in other species as well (very few specimens seen in
other species). The same applies to the peculiar position in which egg-sacs are held by females: in Ph.
halabala , Ph. erawan and Ph. sepaku (and possibly in close relatives), the egg-sac is carried under the
prosoma rather than in front of it (Figs 3, 5-6, 16). For further details, see individual species descriptions
below.

Three of the four species assigned tentatively to the halabala group based on preliminary molecular data
were observed in the field (Ph. kuhapimuk sp. nov.; Ph. khaolek sp. nov.; and Ph. sudhami ), and these
were mostly found on rocks at cave entrances, with the body flat on the rock surface. Only Ph. sudhami
at Erawan was mostly found on exposed tree roots at the riverside. Webs were either barely visible (a
few threads close to the rock surface) or consisted of very delicate small domed sheets attached to the
rock (Ph. khaolek sp. nov.). Silk tufts were not seen in any of these webs. Egg-sacs were carried in front
of the body as in typical pholcids (Fig. 51).

Distribution

The Ph. halabala group is restricted to Southeast Asia, from Myanmar and southern China to Sumatra
and Borneo (Figs 17, 57).

Pholcus halabala Huber, 2011
Figs 1-6, 18-28

Pholcus halabala Huber, 2011: 128-131, figs 495M96, 517-518, 530-535 ((??).

Diagnosis

Easily distinguished from most other species in halabala core group by simple triangular uncus and
short curved appendix with simple rounded tip (fig. 530 in Huber 2011; Fig. 24), and by distinctive
sclerite anteriorly in internal female genitalia (fig. 533 in Huber 2011; similar in Ph. lintang sp. nov, cf.
Fig. 37); from very similar Ph. sepaku only by shape of procursus (distinctive distal elements; compare
figs 535 and 536 in Huber 2011).

New material examined

THAIFAND: 1 2 juvs, in absolute ethanol, ZFMK (Mai 302, 315), Narathiwat, Hala Bala Wildlife

Sanctuary, ‘site V, forest at river near headquarters (5°47.8' N, 101°49.9' E), 90 m a.s.l., on leaves,
1-2 Mar. 2015 (B.A. Huber, B. Petcharad).

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European Journal of Taxonomy 190 : 1-55 ( 2016 )

MALAYSIA: 1 $ in ZFMK (Ar 15000), Kedah, Gunung Jerai, forest near Sri Perigi Waterfall (5°48.3' N,
100°24.6' E), 100-200 m a.s.l., on leaf, 27 Feb. 2015 (B.A. Huber); 1 juv., in absolute ethanol, ZFMK (Mai
292), same data; 1 S, 1 ZFMK (Ar 15001), Pulau Pinang, Penang National Park near Teluk Bahang
(5°27.7' N, 100°12.1' E), 10-50 m a.s.l., on leaves, 28 Feb. 2015 (B.A. Huber); 1 in absolute ethanol,
ZFMK (Mai 119), same data; 1 $, ZFMK (Ar 15002), Perak, Gunung Liang (3°47.7' N, 101°32.0' E),
250 m a.s.l., forest along river, on leaf, 22 Feb. 2015 (B.A. Huber, A.R.M. Ghazali, K.A. Braima); 2 §§,
1 juv., in absolute ethanol, ZFMK (Mai 268), same data; 1 ZFMK (Ar 15003), Pahang, Ulu Dong
(3°56.2' N, 102°01.9' E), 190 m a.s.l., forest near river, on leaf, 21 Feb. 2015 (B.A. Huber, A.R.M. Ghazali,
K.A. Braima); 1 $, in absolute ethanol, ZFMK (Mai 253), same data; 1 ZFMK (Ar 15004), Selangor,
Kemensah (3°13.3F N, 101°47.57' E), 230 m a.s.l., forest along stream, on leaf, 19 Feb. 2015 (B.A. Huber,
A.R.M. Ghazali, K.A. Braima, M. Muslimin); 2 1 §, ZFMK (Ar 15005), Johor, Gunung Ledang, forest

near Puteri Falls (2°21.2'-2°21.6' N, 102°37.8’—102°38.1' E), 100-300 m a.s.l., on leaves, 17 Feb. 2015
(B.A. Huber); 1 §, 4 juvs, in absolute ethanol, ZFMK (Mai 239), same data; 2 §?, ZFMK (Ar 15006),
Gunung Ledang, forest near Puteri Falls (2°21.3' N, 102°38.1' E), 110 m a.s.l., on leaves, 18 Feb. 2015
(B.A. Huber), night collecting; 1 juv. in absolute ethanol, ZFMK (Mai 244), same data.

SINGAPORE: 1 (?, ZFMK (Ar 15007), Upper Selatar Reservoir Park (1°24.0' N, 103°48.4' E), 20 m
a.s.l., on leaf, 15 Feb. 2015 (B.A. Huber, D. Court); 3 juvs, in absolute ethanol, ZFMK (Mai 224), same
data; 2 ZFMK (Ar 15008), Dairy Farm Nature Park (1°21.6' N, 103°46.7' E), 50 m a.s.l., on leaves,
15 Feb. 2015 (B.A. Huber, J. Koh); 1 §, 2 juvs, in absolute ethanol, ZFMK (Mai 214), same data; 2
1 J, ZFMK (Ar 15009), MacRitchie Reservoir Park (1°21.3' N, 103°48.8' E), 50 m a.s.l., on leaves,
14 Feb. 2015 (B.A. Huber, J. Koh, D. Court); 1 $, 4 juvs, in absolute ethanol, ZFMK (Mai 201), same
data.

INDONESIA: 1 (?, 1 $, 1 juv., RMNH, Sumatra, Gunung Leuser National Park at Bohorok [3.54°N,
98.12°E], 1000 m a.s.l., from leaves, 7—10 Aug. 1982, collector unknown.

Description - amendments

Male ocular area on each side with four strong spines (Figs 18-20); apparently without gland openings
(Fig. 23); tarsus 4 comb-hairs of the simplified Pholcus -type (cf. Huber & Fleckenstein 2008), with
three lateral tines (Fig. 27); procursus with retrolateral distal pocket (Figs 21-22); distal cheliceral
apophyses with two cone-shaped teeth (modified hairs) each (Fig. 28); gonopore with four epiandrous
spigots (Fig. 26); ALS with one widened, one pointed, and six smaller cylindrically shaped spigots of
varying sizes (Fig. 25).

Variation

Previously, this species was known only from southern Thailand (type locality) and Sumatra (Huber 2011).
Specimens from Sumatra were assigned tentatively because of minor differences in the procursus. The
new specimens above support the idea that this is intraspecific variation rather than an indication of species
limits. First, the ventral distal spine of the procursus varies continuously among localities, with specimens
from Singapore having a short spine like specimens from Sumatra, and specimens from Malaysia being
intermediate between those from Singapore and Thailand. Most deviating are specimens from Pulau
Pinang, where the ventral distal process of the procursus carries three spines instead of only one. Second,
the shape of the dorsal process of the procursus varies strongly with the angle at which it is viewed. If the
procursus is viewed in slightly dorsal view, the process looks as in the original description (Huber 2011,
fig. 531); if the procursus is viewed in perfect retrolateral view, the process appears slightly wider.

The usual pattern on the carapace consists of two V-marks on the posterior half (Figs 1, 6). In some
specimens, these marks are fused into larger marks or even into a single large mark that covers the entire

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HUBER B.A. et al., Pholcus halabala group in Southeast Asia

Figs 18-28. Pholcus halabala Huber, 2011, ZFMK Ar 15008. 18. Male ocular area, frontal view.
19. Modified hairs on male ocular area, dorsal view. 20. Male prosoma, frontal view. 21-22. Right
procursus (and trochanter apophysis), retrolateral and retrolatero-distal views (arrows point at distal
pocket). 23. Modified hairs on male ocular area. 24. Left bulbal processes, prolateral (slightly ventral)
view. 25. Male ALS. 26. Male gonopore. 27. Comb-hairs on male tarsus 4. 28. Male distal cheliceral
apophysis. Abbreviations: a = appendix; AME = anterior median eyes; e = embolus; p = procursus;
tr = trochanter; u = uncus. Scale lines: 10 pm (25, 27); 20 pm (28); 30 pm (23); 40 pm (26); 80 pm (24);
100 pm (19, 21-22); 200 pm (18); 300 pm (20).

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European Journal of Taxonomy 190: 1-55 (2016)

posterior half of the carapace (Fig. 5). Tibia 1 in newly collected specimens: 12 males: 7.0-9.1 (mean
8.0); 6 females: 6.4-7.7 (mean 7.0).

Natural history

Even though this species is widespread, it seemed to be extremely rare at most localities. This explains
the low specimen numbers even though it was searched for with considerable effort. At the type locality,
three days of intensive search yielded only one female and two juveniles. The ATOL Expedition in
2003 (including several experienced arachnologists using a variety of collecting techniques) also found
only one adult specimen. At some localities (Gunung Jerai; Penang), Ph. halabala was found together
with Ph. erawan, but Ph. halabala seemed to prefer dicot leaves while Ph. erawan was mostly found
on monocot leaves. Otherwise these two species are barely distinguishable in the field. Webs consisted
mainly of round platforms attached to the undersides of leaves. Small silk tufts (Fig. 4) were observed
in some webs at most localities. Egg-sacs are carried under the prosoma (Figs 3, 5-6).

Distribution

Widely distributed from southern Thailand to Singapore and Sumatra (Fig. 17).

Pholcus ubin Huber, sp. nov.

um:lsid:zoobank.org:act:5EC32BD5-FE22-45DE-A5EB-7659D61FFC03

Figs 7-8, 29-33, 90-92

Diagnosis

Easily distinguished from putatively closest known relatives (other species in the halabala core group)
by morphology of male palps (Figs 29-30; procursus with heavily sclerotized dorsal process distally;
shapes of uncus and appendix) and by female internal genitalia (Figs 32-33; triangular pore plates).

Etymology

The species name is derived from the type locality; noun in apposition.

Type material

SINGAPORE: holotype, S, ZFMK (Ar 15010), Pulau Ubin, degraded forest near park headquarters
(1°24.2' N, 103°58.2' E), 20 m a.s.l., on leaves, 16 Feb. 2015 (B.A. Huber, J. Koh, D. Court).

Other material examined

SINGAPORE: 2 S<5, 3 §§, ZFMK (Ar 15011), same data as holotype; 2 §?, 3 juvs, in absolute
ethanol, ZFMK (Mai 232), same data.

MALAYSIA-BORNEO: 1 $, RMNH, Sabah, Gaya Island (6° 00.90' N, 116° 01.17' E), 22 Aug. 2009
(A. Floren).

Description

Male (holotype)

Measurements. Total body length 4.1, carapace width 1.0. Leg 1: 30.4 (7.3 + 0.4 + 7.2 + 14.2 + 1.3),
tibia 2: 4.7, tibia 3: 2.7, tibia 4: 3.9; tibia 1 L/d: 86. Distance PME-PME 220 pm, diameter PME 95 pm,
distance PME-ALE ~35 pm; distance AME-AME 35 pm; diameter AME 60 pm.

Color. Carapace ochre-yellow with brown pattern of radiating marks posteriorly, in live specimens with
reddish color in median area between posterior marks (Fig. 7); ocular area and clypeus not darkened;
sternum with some dark marks posteriorly; legs ochre-yellow with dark brown patellae and tibia-

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HUBER B.A. et al ., Pholcus halabala group in Southeast Asia

metatarsus joints; abdomen pale ochre-gray with small black and white marks dorsally and laterally,
monochromous ventrally.

Body. Habitus as in Fig. 7; ocular area slightly raised, with brushes of ~ 10 stronger hairs behind each PME;
carapace without median furrow; clypeus unmodified; sternum wider than long (0.62/0.40), unmodified.

Figs 29-33. Pholcus ubin Huber, sp. nov., ZFMK Ar 15011. 29-30. Feft male palp, prolateral and
retrolateral views (arrow points at distinctive dorsal process; asterisk marks dorsal elongation of male
palpal tarsus). 31. Male chelicerae, frontal view. 32-33. Cleared female genitalia, ventral and dorsal
views. Abbreviations: a = appendix; b = genital bulb; e = embolus; p = procursus; u = uncus. Scale lines:
0.3 mm (31); 0.5 mm (29-30, 32-33).

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European Journal of Taxonomy 190: 1-55 (2016)

Chelicerae. As in Fig. 31, with pair of frontal apophyses provided with two modified hairs each and
rounded lateral processes.

Palps. As in Figs 29-30; coxa unmodified; trochanter with retrolatero-ventral apophysis; femur with
small retrolateral apophysis proximally; tibia very large; tarsus with dorsal elongation carrying subdistal
tarsal organ; procursus with prominent ventral ‘knee’, with distinctive heavily sclerotized dorsal process
distally (arrow in Fig. 30); bulb oval, with distinctive uncus and appendix; weakly sclerotized embolus
with subdistal fringed side branch (hidden by appendix in Fig. 29).

Legs. Without spines and curved hairs; few vertical hairs; retrolateral trichobothrium on tibia 1 at 5%;
prolateral trichobothrium absent on tibia 1, present on other tibiae; tarsus 1 with ~20 pseudosegments
(only distally ~15 fairly distinct).

Male (variation)

Tibia 1 in 2 other males: 6.1, 7.3.

Female

In general similar to male (Fig. 8), but without stronger hairs behind PME; sternum mostly dark with
some small light marks; eye triads closer together than in male (PME-PME distance: 185 pm). Tibia 1
in 3 females: 5.8, 5.9, 6.1. Epigynum wealdy sclerotized whitish plate (Fig. 90), anterior internal arch
visible through cuticle, posterior margin laterally slightly more sclerotized; with median ‘knob’ (Figs 32,
90-91); internal genitalia as in Figs 33 and 92.

Natural history

All Pulau Ubin specimens were found in a highly degraded patch of forest near park headquarters, while
no specimen was found in a well preserved forest about 3.3 km WNW. Most specimens were collected
by beating of branches, but some were observed in their flat resting position on the leaves.

Distribution

Known from two localities in Singapore and Gaya Island (Sabah) respectively (Fig. 17).

Pholcus sab ah Huber, 2011
Figs 9-10

Pholcus sahah Huber, 2011: 133, figs 497M98, 537-541 (c?$).

Diagnosis

Easily distinguished from putatively closest known relatives (other species in the halabala core group)
by ‘double’ uncus and large rounded rather than pointed flap dorsally on procursus (figs 537, 538 in
Huber 2011) and by large lateral structures in female internal genitalia (fig. 541 in Huber 2011).

New material examined

MALAYSIA-BORNEO: 1 ZFMK (Ar 15012), Sabah, MtKinabalu, PoringHot Springs (type locality),
forest near beginning of Kipungit Trail (6.048° N, 116.706° E), 450 m a.s.l., on underside of leaf, 7 Aug.
2014 (B.A. Huber, S.B. Huber); 1 RMNH, Poring Hot Springs, 28-31 Mar. 1998 (C.L. Deeleman-
Reinhold, P. Zborowski); 1 c?, 2 $?, 2 juvs, ZFMK (Ar 15013), Sepilolc, Rainforest Discovery Centre,
forest along Pitta Trail (5.875-5.878° N, 117.937-117.942° E), 30 m a.s.l., on undersides of leaves,
9 Aug. 2014 (B.A. Huber, S.B. Huber); 1 §, 5 juvs, in absolute ethanol, ZFMK (Bor 173), same data.

12

HUBER B.A. et al., Pholcus halabala group in Southeast Asia

Note

The color dimorphism observed among females from Poring in the original description also occurs
among males: while the single male from Sepilok has the ‘usual’ pattern of two V-marlcs (Fig. 10), the
newly collected male from Poring has a large black mark covering most of the carapace posteriorly
(Fig. 9). This latter pattern also occurs in one of the three females from Sepilok. In the male it is
associated with a slightly darker brown sternum that is only medially and anteriorly light; in the female
it is associated with a black sternum. Tibia 1 in two males: 7.1, 8.4; in two females: 7.1, 7.3.

Natural history

Most new specimens were collected relatively close to the ground (approximately 50 cm above the
ground). All previously known specimens (lc?, 3? $) were collected by canopy fogging (Huber 2011).

Distribution

Known from two localities in northeastern Borneo (Fig. 17).

Pholcus lintang Huber, sp. nov.

um:lsid:zoobank.org:act:2AB6CE0F-8B36-4D8F-BlED-9DFEBB5637DE

Figs 11-12, 34-38, 93-95

Diagnosis

Easily distinguished from putatively closest known relatives (other species in the halabala core group)
by morphology of male palps (Figs 34-35; procursus with rather pointed dorsal flap and without ventro-
distal sclerotized process; simple beak-shaped uncus and simple curved appendix) and by female internal
genitalia (Figs 37-38; large rounded pore plates close to each other; small lateral structures).

Etymology

The species name is derived from the type locality; noun in apposition.

Type material

MALAYSIA-BORNEO: holotype, ZFMK (Ar 15014), Sarawak, Bako National Park, along Lintang
Trail (1.713-1.722° N, 110.447-110.457° E), 10-130 m a.s.l., 11 Jul. 2014 (B.A. Huber, S.B. Huber).

Other material examined

MALAYSIA-BORNEO: 1 c?, 1 $, 2 juvs, ZFMK (Ar 15015), same data as holotype; 1 $ §, 4 juvs,

in absolute ethanol, ZFMK (Bor 192), same data.

Assigned tentatively

MALAYSIA-BORNEO: 1 $, 4 juvs, in absolute ethanol, ZFMK (Bor 221), Sarawak, Kubah National
Park, along Main Trail (1.611°N, 110.191-110.195° E), 160-200 m a.s.l., 13 Jul. 2014 (B.A. Huber, S.B.
Huber); 1 §, in absolute ethanol, ZFMK (Bor 224), Niah Cave National Park, forest near headquarters
(3.820° N, 113.763° E), 40 m a.s.l., night collecting, 28 Jul. 2014 (B.A. Huber, S.B. Huber).

Description

Male (holotype)

Measurements. Total body length 3.5, carapace width 0.95. Leg 1: 27.6 (6.6 + 0.4 + 6.6 + 12.5 + 1.5),
tibia 2: 4.3, tibia 3: 2.6, tibia 4: 3.7; tibia 1 L/d: 79. Distance PME-PME 290 pm, diameter PME 95 pm,
distance PME-ALE ~35 pm; distance AME-AME 55 pm; diameter AME 50 pm.

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European Journal of Taxonomy 190: 1-55 (2016)

Color. Carapace pale ochre-whitish with brown pattern of radiating marks posteriorly; ocular area and
clypeus not darkened; sternum whitish with dark lateral margins; legs pale whitish with brown patellae
and tibia-metatarsus joints; abdomen pale gray with black and white marks dorsally and laterally,
monochromous ventrally.

*

•K"; m

£ 34 ?

tPa r> ‘ ft* f - 0 i

Wif&h

Figs 34-38. Pholcus lintang Huber, sp. nov., ZFMK Ar 15015. 34-35. Left male palp, prolateral and
retrolateral views. 36. Male chelicerae, frontal view. 37-38. Cleared female genitalia, ventral and dorsal
views. Abbreviations: a = appendix; b = genital bulb; e = embolus; p = procursus; u = uncus. Scale lines:
0.3 mm (36-38); 0.5 mm (34-35).

14

HUBER B.A. et al., Pholcus halabala group in Southeast Asia

Body. Habitus as in Fig. 11; ocular area raised, with brushes of ~4 spines on low hump behind each PME;
carapace without median furrow; clypeus unmodified; sternum wider than long (0.66/0.50), unmodified.

Chelicerae. As in Fig. 36, with pair of frontal apophyses provided with two modified hairs each, rounded
lateral processes, and indistinct small frontal proximal humps.

Palps. As in Figs 34-35; coxa unmodified; trochanter with retrolatero-ventral apophysis; femur widened
proximally ventrally, with small retrolatero-dorsal apophysis proximally and low retrolateral hump;
tibia very large; tarsus with conical dorsal elongation carrying subdistal tarsal organ; procursus with
prominent ventral ‘knee’, with distinctive flat dorsal process and distal elements; bulb oval, with
distinctive uncus and appendix; weakly sclerotized embolus with subdistal fringed side branch.

Fegs. Without spines and curved hairs; few vertical hairs; retrolateral trichobothrium on tibia 1 at
6%; prolateral trichobothrium absent on tibia 1, present on other tibiae; tarsus 1 pseudosegments very
indistinct, only distally ~10 poorly visible in dissecting microscope.

Male (variation)

Tibia 1 in other male: 7.0; this male with slightly more pigment, clypeus with large light brown mark.

Female

In general similar to male (Fig. 12) but without spines behind PME and eye triads closer together than
in male (PME-PME distance: 230 pm). Tibia 1 in 1 female: 6.2. Epigynum mostly weakly sclerotized
except for posterior area (Fig. 93); with median ‘knob’ (Figs 37, 94); internal genitalia as in Figs 38 and
95, with roundish pore plates close to each other. Females from Kubah National Park and Niah Cave
National Park are assigned tentatively because females in this species group are not easily distinguished
externally and no males are available from these localities.

Natural history

At Bako, this species was found on the plateau, in the rather low Kerangas forest. At Kubah, most
specimens were taken from large leaves of ornamental plants close to the buildings; only one specimen
was found in well-preserved forest.

Distribution

Known from three localities in northern Borneo (Fig. 17; note that specimens from Kubah and Niah are
assigned tentatively).

Pholcus erawan Huber, 2011
Figs 13-16, 39M8

Pholcus erawan Huber, 2011: 297-298, figs 1374-1375, 1404-1405, 1466-1470 ((??).

Diagnosis

Easily distinguished from putatively closest known relatives (other species in the halabala core group)
by absence of dorsal flap on procursus, by long whitish process of male palpal tarsus (fig. 1467 in Huber
2011), by unique shapes of bulbal processes (fig. 1644 in Huber 2011), and by much longer than wide
female internal genitalia and small oval pore plates (fig. 1470 in Huber 2011).

New material examined

THAIFAND: 8 S8, 9 ? ?, ZFMK (7 88, 8 $ ?, AR 15016-17) and PSUZC (1 1 ?), Kanchanaburi,

Erawan National Park (14°22.2' N, 99°08.75' E), 85 m a.s.l., forest along stream, on leaves, 15 Mar. 2015

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European Journal of Taxonomy 190: 1-55 (2016)

Figs 39-48. Pholcus erawan Huber, 2011, ZFMK Ar 15016-17. 39. Male distal cheliceral apophysis
(arrows point at modified hairs). 40. Left procursus, prolateral view. 41. Right procursus, retrolatero-dorsal
view (arrow points at distal pocket). 42. Left bulbal processes, distal view. 43. Male ocular area, frontal
view. 44. Male gonopore. 45. Female ALS. 46. Comb-hairs on male tarsus 4. 47. Female prosoma, frontal
view. 48. Epigynum, ventral view. Abbreviations: a = appendix; e = embolus; u = uncus. Scale lines: 10 pm
(39); 20 pm (45-46); 40 pm (44); 60 pm (42); 80 pm (40); 100 pm (41, 48); 200 pm (43, 47).

16

HUBER B.A. et al., Pholcus halabala group in Southeast Asia

(B.A. Huber, B. Petcharad); 2 S3, 7 $ in absolute ethanol, ZFMK (Mai 378), same data; 1 3, 1 ?, ZFMK
(Ar 15018), Nakhon Si Thammarat, Khao Nan National Park (8°46.2' N, 98°48.1' E), 250-300 m a.s.l.,
on leaves in forest, 9 Mar. 2015 (B.A. Huber, B. Petcharad); 1 3, ZFMK (Ar 15019), same data, collected
penultimate, adult on 11 Mar. 2015; 3 juvs, in absolute ethanol, ZFMK (Mai 345), same data; 3 33, 3 $ $,
ZFMK (Ar 15020), Nakhon Si Thammarat, Khao Nan National Park (8°46.23' N, 98°48.27' E), 100 m

а. s.l., on palm leaves near park buildings, 9 Mar. 2015 (B.A. Huber, B. Petcharad); 1 3, ZFMK (Ar 15021),
same data, collected penultimate, adult on 12 Mar. 2015; 2 33, ZFMK (Ar 15022), Krabi, Khao Phanom
Bencha National Park, trails near headquarters (8° 14. P N, 98°55. V E), 150-300 m a.s.l., on leaves, 8 Mar.
2015 (B.A. Huber, B. Petcharad); 4 $ §, 4 juvs, in absolute ethanol, ZFMK (Mai 340), same data.

MAFAYSIA: 2 33, 5 ZFMK (Ar 15023), Kedah, Gunung Jerai, forest near Sri Perigi Waterfall
(5°48.3'N, 100°24.6'E), 100-200 m a.s.l., on leaves, 27 Feb. 2015 (B.A. Huber); 1 §, 6 juvs, in absolute
ethanol, ZFMK (Mai 291), same data; 4 33, 3 55, 1 juv., ZFMK (Ar 15024), Pulau Pinang, Penang
National Park near Telulc Bahang (5°27.7' N, 100° 12. V E), 10-50 m a.s.l., on leaves, 28 Feb. 2015 (B.A.
Huber); 1 3, ZFMK (Ar 15025), same data, collected penultimate, adult on 2 Mar. 2015; 1 3, 1 5 >
1 juv., in absolute ethanol, ZFMK (Mai 296), same data.

Description - amendments

Carapace pattern slightly variable, ranging from two separate V-marks (Fig. 15) to medially fused
V-marks to almost completely fused single posterior mark (Fig. 14). Females and juveniles with more
delicate V-marks. Sternum coloration also slightly variable, from almost monochromous whitish to
small black posterior marks (males) and larger black posterior marks (females). Tibia 1 in 21 males:

б. 3-8.1 (mean 7.1); in 20 females: 5.8-6.7 (mean 6.3). In most males, except those from the type locality
(Erawan), the ventro-distal sclerite of the procursus is slightly more pointed than illustrated in Huber
2011 (fig. 1467). Male ocular area with dense brush of stronger hairs, but without spines (Fig. 43); tarsus
4 comb-hairs of the simplified Pholcus -type (cf. Huber & Fleckenstein 2008), with three lateral tines
(Fig. 46); procursus with retrolateral distal pocket (Fig. 41); distal male cheliceral apophyses with two
cone-shaped teeth (modified hairs) each (Fig. 39); gonopore with four epiandrous spigots (Fig. 44); AES
with one widened, one pointed, and six smaller cylindrically shaped spigots of varying sizes (Fig. 45;
pointed spigot damaged in this spinneret).

Natural history

As noted above, Ph. erawan was sometimes found at the same localities as Ph. halabala but on monocot
rather than dicot leaves. Only at Erawan, this species seemed to occur on all kinds of leaves, preferably
(but not only) large ones. At Khao Nan, Ph. erawan was found on palm leaves both in the forest and in
the garden near the park buildings. At Penang, specimens were found both on green leaves and on dead
brown leaves still attached to the plant. Small silk tufts were observed in the webs at most localities. At
night (at Erawan), spiders were observed moving among the vegetation.

Distribution

Widely distributed on the Malay Peninsula, reaching northern Faos (Fig. 17).

Pholcus kuhapimuk Huber, sp. nov.
um:lsid:zoobank.org:act:851588EA-3FD5-4800-9F4D-33BAE6E2E0F9

Figs 49-51, 58-62, 96-98

Diagnosis

Easily distinguished from most similar known relative (Ph. khaolek sp. nov.) by morphology of male
palps (Figs 58-59; shapes of male palpal tarsus and procursus tip, small uncus, slender appendix) and by

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European Journal of Taxonomy 190: 1-55 (2016)

female internal genitalia (Figs 61-62; absence of median triangular sclerite). From other putatively close
relatives also by combination of pale coloration (Figs 49-51), shape of male palpal trochanter apophysis
(curved, proximally wide, distally pointed; Fig. 59), and small round pore plates far apart (Fig. 62).

Etymology

The species name is derived from the type locality; noun in apposition.

Type material

THAILAND: holotype, ZFMK (Ar 15026), Yala, Wat Kuhapimuk (6°31.7' N, 101°13.5' E), 40 m
a.s.l., on wall in cave entry area, 4 Mar. 2015 (B.A. Huber, B. Petcharad).

Figs 49-56. Live specimens, Pholcus kuhapimuk Fluber, sp. nov. (49—51), Ph. khaolek Fluber, sp. nov.
(52-53), and Ph. sudhami Huber, 2011 (54-56). 49-51. penultimate instar and $ with egg-sac
from Wat Kuhapimuk, Thailand. 52-53. $ with spiderlings and S from Khao Lek, Thailand. 54-55. $
from Tham Phraya Nakhon, Thailand. 56. $ from Erawan, Thailand.

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HUBER B.A. et al., Pholcus halabala group in Southeast Asia

Other material examined

THAILAND: 3 33, 6 ZFMK (2 33, 5 Ar 15027) and PSUZC (1 3, 1 $), same data as
holotype; 1 3, 4 4 juvs, in absolute ethanol, ZFMK (Mai 324), same data.

Assigned tentatively

THAILAND: 1 $,2juvs, in absolute ethanol, ZFMK (Mai 329), Satun, ThalebanNational Park (6°43.58' N,
100°09.74' E), at rocks near cave entrance, 100 m a.s.l., 5 Mar. 2015 (B.A. Huber, B. Petcharad).

Description

Male (holotype)

Measurements. Total body length 5.0, carapace width 1.4. Leg 1: 53.3 (12.9 + 0.5 + 12.9 + 25.0 + 2.0),
tibia2: 8.6, tibia 3: 5.4, tibia 4: 7.3; tibia 1 L/d: 98. Distance PME-PME 265 pm, diameter PME 115 pm,
distance PME-ALE ~35 pm; distance AME-AME 35 pm; diameter AME 60 pm.

Color. Carapace pale ochre-grey with pair of brown marks posteriorly; ocular area slightly darker,
clypeus not darkened; sternum pale grey with narrow dark margins; legs pale ochre-yellow with dark
brown patellae and tibia-metatarsus joints; abdomen monochromous pale gray.

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European Journal of Taxonomy 190: 1-55 (2016)

Body. Habitus as in Fig. 49; ocular area slightly raised, with slightly stronger hairs behind each PME;
carapace without median furrow; clypeus unmodified; sternum wider than long (1.00/0.67), unmodified.

Chelicerae. As in Fig. 60, with pair of distal frontal apophyses provided with one or two modified hairs
each, pair of rounded lateral processes, and pair of small indistinct proximal frontal humps.

Figs 58-62. Pholcus kuhapimukUubQr , sp. nov., ZFMK Ar 15027. 58-59. Left male palp, prolateral and
retrolateral views. 60. Male chelicerae, frontal view. 61-62. Cleared female genitalia, ventral and dorsal
views. Abbreviations: a = appendix; b = genital bulb; e = embolus; p = procursus; u = uncus. Scale lines:
0.3 mm (61-62); 0.5 mm (58-60).

20

HUBER B.A. et al., Pholcus halabala group in Southeast Asia

Palps. As in Figs 58-59; coxa unmodified; trochanter with large but weakly sclerotized retrolatero-
ventral apophysis and low retrolateral hump; femur proximally widened on ventral side, with small
retrolatero-dorsal apophysis; tarsus with short conical dorsal elongation carrying subdistal tarsal organ;
procursus rather simple, ventral ‘knee’ with distal process; bulb with distinctive small uncus and slender
appendix; weakly sclerotized short embolus.

Legs. Without spines and curved hairs; few vertical hairs; retrolateral trichobothrium on tibia 1 at 6%;
prolateral trichobothrium absent on tibia 1, present on other tibiae; tarsus 1 with >30 pseudosegments,
distally fairly distinct.

Male (variation)

Tibia 1 in 2 other males: 11.1, 12.6.

Female

In general similar to male (Fig. 51) but without stronger hairs behind PME; eye triads closer together
than in male (PME-PME distance: 185 pm). Tibia 1 in 7 females: 8.1-9.8 (mean 8.7). Epigynum weakly
sclerotized bulging area, only posterior margin more strongly sclerotized, apparently without ‘knob’,
or ‘knob’ hidden below membranous flap (Figs 61, 96-97); internal genitalia as in Figs 62 and 98, with
small round pore plates far from each other. The specimens from Thaleban National Park are assigned
tentatively because no male specimen is available from this locality.

Natural history

At both localities, specimens were only found on rocks at the cave entrances but not deeper in the caves.
The spiders were extremely cryptic, tightly pressed against the rock surface and barely visible even at
close distance.

Distribution

Known from two localities in southern Thailand only (Fig. 57; specimens from Thaleban National Park
assigned tentatively).

Pholcus khaolek Huber, sp. nov.

um:lsid:zoobank.org:act:2AlCAF47-9EB4-480C-8C83-DFB0162AF617

Figs 52-53, 63-78, 99-101

Diagnosis

Easily distinguished from most similar known relative (Ph. kuhapimuk sp. nov.) by morphology of male
palps (Figs 63-64; shapes of male palpal tarsus and procursus tip, larger uncus, complex appendix with
retrolateral process) and by female internal genitalia (Figs 66-67; distinctive median triangular sclerite).
From other close relatives also by combination of pale coloration (Figs 52-53), shape of male palpal
trochanter apophysis (curved, proximally wide, distally pointed; Fig. 64), and shape and position of pore
plates (Fig. 67).

Etymology

The species name is derived from the type locality; noun in apposition.

Type material

THAILAND: holotype, ZFMK (Ar 15028), Nakhon Si Thammarat, Khao Nan National Park, Tham
Khao Lek (8°46.09' N, 98°43.68' E), 95 m a.s.l., on rock walls around cave, 10 Mar. 2015 (B.A. Huber,
B. Petcharad).

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European Journal of Taxonomy 190: 1-55 (2016)

Other material examined

THAILAND: 6 88, 5 2 juvs, ZFMK (5 88, 4 $$, Ar 15029-30) and PSUZC (1 8, 1 ?), same

data as holotype; 1 c?, 2 $ 3 juvs, in absolute ethanol, ZFMK (Mai 350), same data.

Figs 63-67. Pholcus khaolek Huber, sp. nov., ZFMK Ar 15029-30. 63-64. Left male palp, prolateral
and retrolateral views. 65. Male chelicerae, frontal view. 66-67. Cleared female genitalia, ventral and
dorsal views. Abbreviations: a = appendix; b = genital bulb; e = embolus; p = procursus; u = uncus.
Scale lines: 0.5 mm

22

HUBER B.A. et al., Pholcus halabala group in Southeast Asia

Figs 68-78. Pholcus khaolek Huber, sp. nov., ZFMK Ar 15029-30. 68. Male ocular area, frontal
view. 69. Modified hairs between male eye triads. 70. Male distal cheliceral apophysis (arrows point
at modified hairs). 71. Right procursus, retrolateral view. 72. Left procursus, prolateral view. 73. Left
bulbal processes, prolateral view. 74. Comb-hairs on male tarsus 4. 75. Tip of right procursus, retrolateral
view. 76. Female prosoma, frontal view. 77. Epigynum, ventral view. 78. Male ALS. Abbreviations:
a = appendix; b = genital bulb; e = embolus; p = procursus; tr = trochanter; u = uncus. Scale lines: 20 pm
(70, 74, 78); 30 pm (69, 75); 80 pm (73); 100 pm (72); 200 pm (68, 71, 77); 400 pm (76).

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European Journal of Taxonomy 190: 1-55 (2016)

Description

Male (holotype)

Measurements. Total body length 4.6, carapace width 1.3. Leg 1: 48.6 (11.6 + 0.6 + 11.7 + 22.8 + 1.9),
tibia2: 7.9, tibia 3: 4.5, tibia4: 6.3; tibia 1 L/d: 95. Distance PME-PME 340 pm, diameter PME 125 pm,
distance PME-ALE ~35 pm; distance AME-AME 50 pm; diameter AME 55 pm.

Color. Carapace pale ochre-yellow with pair of light brown marks posteriorly; ocular area and clypeus
not darkened; sternum light brown with lighter marks and dark lateral margins; legs ochre-yellow with
dark brown patellae and tibia-metatarsus joints; abdomen monochromous ochre-gray.

Body. Habitus as in Fig. 53; ocular area slightly raised, with brushes of stronger hairs behind each
PME (Figs 68-69); carapace without median furrow; clypeus unmodified; sternum wider than long
(0.84/0.52), unmodified. ALS with one widened, one pointed, and six smaller cylindrically shaped
spigots of varying sizes (Fig. 78).

Chelicerae. As in Fig. 65, with pair of distal frontal apophyses provided with two to three modified
(cone-shaped) hairs each (Fig. 70), pair of rounded lateral processes, and pair of small indistinct proximal
frontal humps.

Palps. As in Figs 63-64; coxa unmodified; trochanter with large retrolatero-ventral apophysis; femur
proximally widened on ventral side, with small retrolatero-dorsal apophysis; tarsus without dorsal
elongation; procursus rather simple (Figs 71-72), with prolateral weakly sclerotized process and
transparent membranous structures; procursus tip as in Fig. 75; bulb with distinctive uncus and appendix
with retrolateral process (Fig. 73); wealdy sclerotized short embolus.

Legs. Without spines and curved hairs; few vertical hairs; retrolateral trichobothrium on tibia 1 at 6%;
prolateral trichobothrium absent on tibia 1, present on other tibiae; tarsus 1 with >30 pseudosegments,
distally fairly distinct. Tarsus 4 comb-hairs as in Fig. 71.

Male (variation)

Tibia 1 in 5 other males: 10.1-12.8 (mean 11.7). Some males with white marks on abdomen.

Female

In general similar to male (Fig. 52) but without stronger hairs behind PME; eye triads closer together
than in male (PME-PME distance: 220 pm). Tibia 1 in 5 females: 8.7-10.1 (mean 9.3). Epigynum
weakly sclerotized bulging area, only posterior area more strongly sclerotized, with small but distinct
‘knob’ (Figs 66, 77, 99-100); internal genitalia as in Figs 67 and 101, with anterior arch and distinctive
triangular sclerite visible through cuticle.

Natural history

This species was abundant at the type locality on vertical and slightly overhanging smooth rocks.
Specimens were observed tightly pressed against the rock surface, in some cases with a small domed
web nearby. When disturbed, the spiders dropped to the ground.

Distribution

Known from type locality only (Fig. 57).

Pholcus sudhami Huber, 2011
Figs 54-56, 79-89

Pholcus sudhami Huber, 201F 300, figs 1380-1382, 1411-1412, 1476-1480 ($$).

24

HUBER B.A. et al., Pholcus halabala group in Southeast Asia

Diagnosis

Easily distinguished from most congeners by male palpal morphology (figs 1476, 1477 in Huber 2011;
distinctive procursus with large membranous structure, absence of appendix) and by female genitalia

Figs 79-89. Pholcus sudhami Huber, 2011, ZFMK Ar 15031-32. 79-80. Male and female prosomata,
frontal views. 81. Female ALS. 82. Comb-hairs on male tarsus 4. 83. Right procursus, retrolateral
(slightly distal) view. 84. Left bulb and procursus, prolateral view. 85. Left uncus and embolus, prolateral
view. 86. Male distal cheliceral apophysis (arrows point at modified hairs). 87. Epigynum, ventral view.
88. Male gonopore. 89. Pseudotrichia on ventral side of procursus. Abbreviations: b = genital bulb;
e = embolus; p = procursus; u = uncus. Scale lines: 20 pm (81-82, 86); 30 pm (89); 50 pm (88); 60 pm
(85); 200 pm (83-84, 87); 300 pm (79-80).

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European Journal of Taxonomy 190: 1-55 (2016)

(figs 1479 and 1480 in Huber 2011; elongate pore plates in lateral position and semicircular internal
structures visible through cuticle; very similar to Ph. pakse)', from Ph. pakse only by male embolus
without spines (compare figs 1476 and 1481 in Huber 2011).

Figs 90-101. Female genitalia, untreated in ventral view, cleared in ventral and dorsal views. 90-
92. Pholcus ubin Huber, sp. nov. 93-95. Ph. lintang Huber, sp. nov. 96-98. Ph. kuhapimuk Huber, sp.
nov. 99-101. Ph. khaolek Huber, sp. nov.

26

HUBER B.A. et al., Pholcus halabala group in Southeast Asia

New material examined

THAILAND: 9 88 , 8 ? ?, ZFMK (8 88 , 7 $ ?, Ar 15031-32) and PSUZC (1 8 , 1 ?), Kanchanaburi,
Erawan National Park (14°22.2' N, 99°08.75' E), 85 m a.s.l., forest along stream, on rocks and tree roots,
15 Mar. 2015 (B.A. Huber, B. Petcharad); 2 5 ?, in absolute ethanol, ZFMK (Mai 379), same data; 1 8 ,
2 $ J, RMNH, Erawan National Park, 15-16 Mar. 1986 (C.L. & PR. Deeleman).

Assigned tentatively

THAILAND: 3 88 , 3 $ $, ZFMK (Ar 15033), Prachuap Khiri Khan, Khao Sam Roi Yot National Park,
near Tham Phraya Nakhon (12° 12.0' N, 100°00.8' E), 40 m a.s.l., at rocks in forest, 14 Mar. 2015 (B.A.
Huber, B. Petcharad); 2 §§, 3 juvs, in absolute ethanol, ZFMK (Mai 373), same data; 1 §, ZFMK
(Ar 15034), Khao Sam Roi Yot National Park, Tham Kaeo (12°12.2' N, 99°59.5' E), 60 m a.s.l., under
rock in cave entrance, 14 Mar. 2015 (B.A. Huber, B. Petcharad); 1 §, RMNH, Sam Roi Yot National
Park, no further locality information, 8 Dec. 1990 (C.L. & PR. Deeleman); 1 juv., in absolute ethanol,
ZFMK (Mai 376), Wat Huai Takaeng (13°35.23' N, 99°45.52' E) (= “Tham Phraya Prap” in Huber
2011), 30 m a.s.l., in cave, 15 Mar. 2015 (B.A. Huber, B. Petcharad).

Description - amendments

In males from Khao Sam Roi Yot, the uncus has a slightly more pointed tip and the subdistal spine on
the embolus has a more transversal position relative to the embolus. Specimens from this locality are
therefore assigned tentatively. Tibia 1 in 7 males from type locality (Erawan): 8.7-10.6 (mean 9.9); in
7 females: 7.5-8.6 (mean 8.1). Hairs on male ocular area not visibly different from those in females
(Figs 79-80); tarsus 4 comb-hairs of the simplified Pholcus- type (cf. Huber & Fleckenstein 2008), with
three lateral tines (Fig. 82); procursus with distinctive field of ventral spines (Figs 83, 89); male distal
cheliceral apophyses with two modified hairs each (Fig. 86); gonopore with four epiandrous spigots
(Fig. 88); ALS with one widened, one pointed, and six smaller cylindrically shaped spigots of varying
sizes (Fig. 81).

Natural history

At the type locality (Erawan) most specimens were found on the lower surfaces of exposed tree roots
at the riverside. At the other localities, specimens were only found on overhanging smooth rocks at the
cave entrances (Tham Kaeo; Wat Huai Takaeng) or in the forest (Phraya Nakhon). When disturbed, the
spiders dropped to the ground.

Distribution

Known from four localities in central western Thailand (Fig. 57).

Pholcus krabi species group

This species group is newly proposed to include three species previously part of the Ph. halabala
group (Ph. chiangdao Huber, 2011; Ph. khene Huber, 2011; Ph. kinabalu Huber, 2011) as well as three
newly described species (Ph. krabi Huber, sp. nov.; Ph. narathiwat Huber, sp. nov.; Ph. kipungit Huber,
sp. nov.). They share three putative synapomorphies: (1) absence of AME (Figs 116-117); (2) absence
of modified hairs on distal male cheliceral apophyses (Fig. 118); and (3) reduction of ALS spigots to two
(one widened, one pointed; Fig. 122; confirmed in Ph. kinabalu and Ph. krabi sp. nov. only). In addition,
live males of the three newly described species share highly distinctive reddish to orange palps (Figs 102,
106, 108), and females of at least Ph. chiangdao , Ph. kinabalu , and Ph. narathiwat sp. nov. share
dimorphic color patterns on the prosoma. Species newly observed in the field (Ph. kipungit sp. nov.; Ph.
narathiwat sp. nov.; Ph. krabi sp. nov.) built very similar domed webs among the vegetation (0.5-2 m
above the ground), usually with the apex of the dome attached to the underside of a leaf. They are also

27

European Journal of Taxonomy 190 : 1-55 ( 2016 )

very similar in general appearance (the three species are indistinguishable in the field; Figs 102-109).
Very low abundances and/or patchy distributions were observed in all three species. However, most
specimens known of Ph. kinabalu were collected by canopy fogging (Huber 2011), suggesting that
abundances of at least this species may be different in higher forest strata. Egg-sacs are carried in front
of the body (Figs 105, 109) as in typical pholcids. This species group is known from mainland Southeast
Asia and Borneo (Fig. 110). The RMNH has an additional species from East Kalimantan (Fig. 110) that
is not described here because only a single poorly preserved male specimen is available.

Pholcus krabi Huber, sp. nov.

urn: 1 sid: zoobank, org: act: A078E A4B - 8 9F1-4E10-8592-DF39683AFA43

Figs 102-105, 111-123, 134-136

Diagnosis

Distinguished from similar species (other species in the Ph. krabi group) by morphology of male palps
(Figs 111-112; unique shapes of small uncus and small appendix; distinctive procursus with prolateral
process ending in heavily sclerotized tip) and by distinctive pair of small dark internal structures in
internal female genitalia (arrow in Fig. 114).

Etymology

The species name is derived from the type locality; noun in apposition.

Type material

THAILAND: holotype, ZFMK (Ar 15035), Krabi, Khao Phanom Bencha National Park, trails near
headquarters (8° 14.6' N, 98°55.0' E), 110 m a.s.l., domed webs among forest vegetation, 8 Mar. 2015
(B.A. Huber, B. Petcharad).

Other material examined

THAILAND: 3 4 ¥¥> ZFMK (Ar 15036-37), same data as holotype; 2 ¥¥> 4 juvs, in absolute

ethanol, ZFMK (Mai 338), same data; 2 $$, 5 ¥¥> ZFMK (Ar 15038-39), Surat Thani, Khao Sole
National Park, forest along nature trail (8°54.8' N, 98°29.3'-98 o 30.5' E), domed webs among vegetation,
110-160 m a.s.l., 11-12 Mar. 2015 (B.A. Huber, B. Petcharad); 3 $$, in absolute ethanol, ZFMK
(Mai 357), same data; 2 SS, 3 $¥, ZFMK (Ar 15040), Ranong, Klong Nakha Wildlife Sanctuary
(9 0 27.6' N, 98°30.7' E), 40 m a.s.l., forest, domed webs among vegetation, 12 Mar. 2015 (B.A. Huber,
B. Petcharad); 3 ¥ ¥, in absolute ethanol, ZFMK (Mai 363), same data.

Description

Male (holotype)

Measurements. Total body length 4.1, carapace width 0.95. Leg 1: 34.8 (8.3 + 0.4 + 8.3 + 15.5 + 2.3),
tibia2: 5.2, tibia 3: 2.9, tibia4: 4.5; tibia 1 L/d: 86. Distance PME-PME 375 pm, diameter PME 105 pm,
distance PME-ALE ~35 pm; AME absent.

Color. Carapace pale ochre to whitish with small dark median mark, ocular area and clypeus with
light brown pattern; sternum whitish; palps reddish; legs pale ochre to whitish with brown patellae
and tibia-metatarsus joints; abdomen pale gray with very indistinct darker marks dorsally and laterally,
monochromous ventrally.

Body. Habitus as in Figs 102-103; ocular area slightly raised and each triad on short stalk directed laterad
(Fig. 116); carapace without median furrow; clypeus unmodified; sternum wider than long (0.60/0.48),
unmodified. ALS with only two spigots (one widened, one pointed; Fig. 122).

28

HUBER B.A. et al., Pholcus halabala group in Southeast Asia

Chelicerae. As in Fig. 113, with large proximal processes and pair of rounded distal apophyses without
modified hairs (Fig. 118).

Palps. As in Figs 111-112; coxa unmodified; trochanter with long retrolatero-ventral apophysis directed
first laterad then bending sharply ventrad (Fig. 119); femur with small retrolatero-dorsal process; tibia
large; procursus with distinctive prolateral process ending in heavily sclerotized tip (Fig. 120), with
sclerotized and membranous pointed processes ventrally; bulb with indistinct prolateral process, strong
proximal sclerite, small uncus and small appendix (Fig. 121), wealdy sclerotized embolus.

Fegs. Without spines and curved hairs; few vertical hairs; retrolateral trichobothrium on tibia 1 at 3%;
prolateral trichobothrium absent on tibia 1, present on other tibiae; tarsus 1 pseudosegments not seen in
dissecting microscope. Tarsus 4 comb-hairs of the simplified Pholcus- type (cf. Huber & Fleckenstein
2008), with five lateral tines (Fig. 123).

Figs 102-109. Five specimens, Pholcuskrabi Huber, sp. nov. (102-105), Ph. narathiwatHubQr, sp. nov.
(106-107), and Ph. kipungit Huber, sp. nov. (108-109). 102-104. S and $ from Khao Sole, Thailand.
105. $ with egg-sac from Phanom Bencha, Thailand. 106-107. $ and $ from Hala Bala, Thailand.
108-109. & and $ from Poring, Sabah.

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European Journal of Taxonomy 190 : 1-55 ( 2016 )

Male (variation)

Tibia 1 in 6 other males: 7.7-8.3 (mean 8.1).

Female

In general similar to male (Figs 104-105) but eye triads on low humps and closer together (Fig. 117;
PME-PME distance: 220 pm); clypeus and ocular area frontally black; sternum anterior part slightly
darker. Tibia 1 in 12 females: 6.3-7.3 (mean 6.8). Epigynum weakly sclerotized slightly bulging plate
with semi-transparent posterior ‘knob’, internal anterior arch and distinctive small sclerites visible
through cuticle (Figs 114, 134-135); internal genitalia as in Figs 115 and 136, with straight anterior
sclerite and oval pore plates.

Natural history

At Phanom Bencha, all eight specimens were collected from two bushes close to each other. Two days
of intensive search in various parts of the forest did not result in any further specimens. At Khao Sole, all
specimens were found in degraded forest close to the park road; none was found in well preserved forest.

Distribution

Known from three localities in southern Thailand (Fig. 110).

30

HUBER B.A. et al ., Pholcus halabala group in Southeast Asia

Figs 111-115. Pholcus krabi Huber, sp. nov., ZFMK Ar 15036-37. 111-112. Left male palp, prolateral
and retrolateral views. 113. Male chelicerae, frontal view. 114-115. Cleared female genitalia, ventral
and dorsal views (arrow points at distinctive small dark internal structure). Abbreviations: a = appendix;
b = genital bulb; e = embolus; p = procursus; u = uncus. Scale lines: 0.3 mm (113), 0.5 mm (111-112,
114-115).

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European Journal of Taxonomy 190 : 1-55 ( 2016 )

Figs 116-123. Pholcuskrabi Huber, sp. nov., ZFMKAr 15038-39.116-117. Male and female prosomata,
frontal views. 118. Male distal cheliceral apophysis. 119. Left male palp, dorsal view. 120. Left procursus
(and trochanter apophysis), retrolateral view. 121. Right bulbal processes, prolateral (slightly ventral)
view. 122. Male ALS. 123. Comb-hairs on male tarsus 4. Abbreviations: a = appendix; b = genital bulb;
e = embolus; f = femur; p = procursus; tr = trochanter; u = uncus. Scale lines: 8 pm (123); 10 pm (118,
122); 80 pm (121); 100 pm (120); 200 pm (116-117, 119).

32

HUBER B.A. et al., Pholcus halabala group in Southeast Asia

Pholcus narathiwat Huber, sp. nov.

urn:lsid:zoobank.org:act:0127ElC4-BABC-4C33-A92E-65744D43D8E2

Figs 106-107, 124-128, 137-139

Diagnosis

Distinguished from similar species (other species in the Ph. krabi group) by morphology of male palps
(Figs 124-125; unique shape of bifid appendix; shape of uncus; procursus with strong distal ventral
sclerite, similar only in Ph. chiangdao ) and by distinctive rounded sclerites in internal female genitalia
(Fig. 127).

Etymology

The species name is derived from the type locality; noun in apposition.

Type material

THAILAND: holotype, ZFMK (Ar 15041), Narathiwat, Hala Bala Wildlife Sanctuary, ‘site 3’
(5°48.4' N, 101°49.4' E), 180 m a.s.l., forest near road, domed webs among vegetation, 2 Mar. 2015
(B.A. Huber, B. Petcharad).

Other material examined

THAILAND: 1 5, together with holotype; 1 $, in absolute ethanol, ZFMK (Mai 312), same data but
at 220 m a.s.l.; 1 §, in absolute ethanol, ZFMK (Mai 316), same locality but ‘site E, forest at river
near headquarters (5°47.8' N, 101°49.9' E), 90 m a.s.l., 2 Mar. 2015 (B.A. Huber, B. Petcharad), night
collecting.

Description

Male (holotype)

Measurements. Total body length 3.9, carapace width 1.0. Leg 1: 30.6 (7.2 + 0.4 + 7.3 + 13.5 + 2.2),
tibia 2: 4.4, tibia 3: 2.4, tibia 4: 4.1; tibia 1 L/d: 76. Distance PME-PME 350 pm, diameter PME 95 pm,
distance PME-ALE ~35 pm; AME absent.

Color. Carapace pale ochre-yellow with light brown median line and V-mark, ocular area and clypeus
light brown; sternum whitish; palps orange; legs pale ochre-yellow with darker brown patellae and tibia-
metatarsus joints; abdomen pale gray with some indistinct marks dorsally.

Body. Habitus as in Fig. 106; ocular area slightly raised and each triad on short stalk directed laterad;
carapace without median furrow; clypeus unmodified; sternum wider than long (0.60/0.52), unmodified.

Chelicerae. As in Fig. 126, with large proximal lateral processes and pair of rounded distal apophyses
without modified hairs.

Palps. As in Figs 124-125; coxa unmodified; trochanter with retrolatero-ventral apophysis directed
first laterad then bending sharply ventrad; femur with small retrolatero-dorsal process proximally; tibia
large; procursus distally complex, with distinctive strong ventral apophysis; bulb with strong proximal
sclerite, with uncus, with distinctive bifid appendix (prolateral part with large scales), short weakly
sclerotized embolus.

Legs. Without spines and curved hairs; few vertical hairs; retrolateral trichobothrium on tibia 1 at 4%;
prolateral trichobothrium absent on tibia 1, present on other tibiae; tarsus 1 with > 20 pseudosegments,
only distally fairly distinct.

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European Journal of Taxonomy 190 : 1-55 ( 2016 )

Female

In general similar to male (Fig. 107) but eye triads on low humps and closer together (PME-PME
distance: 235 pm), ocular area and clypeus either with one large black mark (1 J) or with two transversal
black bands (2 §?)• Tibia 1: 5.9, 6.3, 6.6. Epigynum wealdy sclerotized slightly bulging plate with
posterior ‘knob’, internal anterior arch and distinctive rounded sclerites poorly visible through cuticle
(Figs 127, 137-138); internal genitalia as in Figs 128 and 139.

Figs 124-128. Pholcus narathiwat Huber, sp. nov., ZFMK Ar 15041.124-125. Left male palp, prolateral
and retrolateral views. 126. Male chelicerae, frontal view. 127-128. Cleared female genitalia, ventral
and dorsal views. Abbreviations: a = appendix; b = genital bulb; e = embolus; p = procursus; u = uncus.
Scale lines: 0.3 mm (126), 0.5 mm (124-125, 127-128).

34

HUBER B.A. et al., Pholcus halabala group in Southeast Asia

Natural history

Three of the four specimens were collected from a single bush. Three days of intensive collecting at the
type locality resulted in only one further specimen. The ATOL Expedition in 2003 did not collect this
species. The domed webs were easily visible among the vegetation, about 1—1.5 m above the ground.

Distribution

Known from type locality in southern Thailand only (Fig. 110).

Pholcus kipungit Huber, sp. nov.

urn:lsid:zoobank.org:act:374082FB-1878-493D-A8B6-CC148D8BB223

Figs 108-109, 129-133, 140-142

Diagnosis

Distinguished from similar species (other species in the Ph. krabi group) by morphology of male
palps (Figs 129-130; long trochanter apophysis; distinctive shapes of simple round uncus and small
appendix; procursus strongly bent dorsad, with distinctive prolatero-ventral pointed process and fringed
membranous distal elements) and by unique median sclerite in internal female genitalia visible through
cuticle (Figs 132, 140).

Etymology

The species name is derived from the type locality; noun in apposition.

Type material

MAFAYSIA-BORNEO: holotype, ZFMK (Ar 15042), Sabah, Mt Kinabalu, Poring Hot Springs,
forest near beginning of Kipungit Trail (6.048° N, 116.706° E), 450 m a.s.l., domed webs in vegetation,
7 Aug. 2014 (B.A. Huber, S.B. Huber).

Other material examined

MAFAYSIA-BORNEO: 1 $, together with holotype; 2 §§, in absolute ethanol, ZFMK (Bor 205),
same data; 1 RMNH, Sabah, Poring Hot Springs (6°03.467' N, 116°42.205' E), 9 Aug. 2009
(A. Floren); 1 2 juvs, RMNH, Poring Hot Springs, 500 m a.s.l., primary forest, 2 May 1991 (C.F. &

PR. Deeleman); 1 5, ZFMK (Ar 15043), Sabah, Sepilok, Rainforest Discovery Centre, forest along
Pitta Trail (5.875-5.878° N, 117.937-117.942° E), 30 m a.s.l., domed web among vegetation, 9 Aug.
2014 (B.A. Huber, S.B. Huber); 3 $ $, in absolute ethanol, ZFMK (Bor 174), same data; 3 SS, 3 $ §,
RMNH (2 vials), Sabah, Danum Valley [5.022° N, 117.747° E], primary forest, 6-16 May 1991 (C.F. &
PR. Deeleman).

Description

Male (holotype)

Measurements. Total body length 3.7, carapace width 0.9. Teg 1: 32.1 (7.5 + 0.4 + 7.4 + 14.3 + 2.5),
tibia 2: 4.5, tibia 3: 2.5, tibia 4: 4.1; tibia 1 F/d: 84. Distance PME-PME 270 pm, diameter PME 95 pm,
distance PME-AFE ~35 pm; AME absent.

Color. Carapace pale whitish to gray with pair of light brown marks, ocular area and clypeus not
darkened; sternum whitish; palps orange; legs pale ochre-yellow with dark patellae and tibia-metatarsus
joints; abdomen pale gray with some indistinct darker marks dorsally.

Body. Habitus as in Fig. 108; ocular area slightly raised and each triad on low hump; carapace without
median furrow; clypeus unmodified; sternum wider than long (0.60/0.50), unmodified.

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European Journal of Taxonomy 190: 1-55 (2016)

Chelicerae. As in Fig. 131, with small proximal processes and pair of rounded distal apophyses without
modified hairs.

Palps. As in Figs 129-130; coxa unmodified; trochanter with very long retrolatero-ventral apophysis
with retrolateral process proximally; femur with small retrolatero-ventral process proximally; tibia

Figs 129-133. Pholcus kipungit Huber, sp. nov., ZFMK Ar 15042. 129-130. Left male palp, prolateral
and retrolateral views. 131. Male chelicerae, frontal view. 132-133. Cleared female genitalia, ventral and
dorsal views. Abbreviations: a = appendix; b = genital bulb; e = embolus; p = procursus; tr = trochanter;
u = uncus. Scale lines: 0.3 mm (131-133), 0.5 mm (129-130).

36

HUBER B.A. et al ., Pholcus halabala group in Southeast Asia

large; procursus strongly bent dorsad, with distinctive prolatero-ventral pointed process and fringed
membranous distal elements; bulb with strong proximal sclerite, with small uncus and small appendix,
short weakly sclerotized embolus.

Legs. Without spines and curved hairs; few vertical hairs; retrolateral trichobothrium on tibia 1 at 3%;
prolateral trichobothrium absent on tibia 1, present on other tibiae; tarsus 1 with >30 pseudosegments,
only distally fairly distinct.

Female

In general similar to male (Fig. 109) but eye triads closer together (PME-PME distance: 195 pm), entire
clypeus and ocular area frontally black; legs pale ochre, less yellowish. Tibia 1: 6.7, 6.9 (missing in
other females). Epigynum weakly sclerotized almost flat plate with posterior ‘knob’, internal distinctive
median sclerite visible through cuticle (Figs 132, 140-141); internal genitalia as in Figs 133 and 142.

Natural history

Of the four webs seen at Poring, three had their apex connected to the underside of a leaf; the fourth (of
the only male) was freely suspended among the twigs at about 1.5 m above the ground.

Figs 134-142. Female genitalia, untreated in ventral view, cleared in ventral and dorsal views. 134-
136. Pholcus krabi Huber, sp. nov. 137-139. Ph. narathiwat Huber, sp. nov. 140-142. Ph. kipungit
Huber, sp. nov.

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European Journal of Taxonomy 190: 1-55 (2016)

Distribution

Known from two localities in Sabah (Fig. 110).

Pholcus kinabalu Huber, 2011

Pholcus kinabalu Huber, 2011: 138-141, figs 511-514, 527-528, 556-569 (c??)-

Diagnosis

Distinguished from other species in krabi group by male palpal morphology (triangular appendix
provided with scales; rather short pointed trochanter apophysis; fig. 556 in Huber 2011) and female
internal genitalia (large round pore plates far from each other; fig. 559 in Huber 2011).

New material examined

MALAYSIA-BORNEO: 1 S, RMNH, Sabah, Mt Kinabalu National Park, Poring Hot Springs (6°02' N,
116°50' E, coordinates dubious), canopy fogging, 20 Jan. 1992 (A. Floren); 1 S,2 4 juvs, RMNH,

same locality, canopy fogging, 19 Feb.-26 Mar. 1996 (A. Floren); 1 3 §§, RMNH, Poring Hot

Springs (6°05' N, 116°33' E, coordinates dubious), 16 Sep. 2006 (A. Floren).

Pholcus buatong species group

This species group is newly proposed to include one species previously part of the Ph. halabala group
(Ph. satun Huber, 2011), one species previously tentatively assigned to the Pholcus ethagala group
(Ph. schwendingeri Huber, 2011), and a newly described species (Ph. buatong Huber, sp. nov.). They
share three putative synapomorphies, (1) the complete reduction of distal anterior apophyses on the
male chelicerae (Fig. 156); (2) the very distinctive dorsal bulging of the male palpal patella (Fig. 155;
angle between femur and patella -120-125° rather than -180° as in typical pholcids); and (3) the large,
heavily sclerotized ‘knob’ on the epigynum (Figs 184, 187, 190). The group is strongly supported by
preliminary molecular data (A. Valdez-Mondragon, B.A. Huber & D. Dimitrov unpublished data).
Pholcus schwendingeri and Ph. buatong sp. nov. also share a distinctive whitish membranous process
retrolatero-distally on the procursus (arrows in Figs 155, 180). Otherwise this group appears rather
inhomogeneous: Pholcus schwendingeri males have extremely long eye stalks (Fig. 173) while males of
the other two species have short eye stalks (Fig. 155); Pholcus buatong sp. nov. is rock-dwelling while
the other two species are leaf litter dwelling; Pholcus satun has small AME, while the other two species
lack AME; Pholcus satun males have only one bulbal process (sclerotized embolus), while males of
the other two species have a membranous embolus plus an appendix. In all three species, egg-sacs are
carried in front of the body (Figs 145, 152) as in typical pholcids. This species group is known from
southern Thailand and northern mainland Malaysia (Fig. 153).

Pholcus buatong Huber, sp. nov.

um:lsid:zoobank.org:act:D2E21FAE-2ECD-414D-81E4-4C3CC7FC352F

Figs 143-146, 154-168, 184-186

Diagnosis

Easily distinguished from known congeners by morphology of male palps (Figs 154-155; unique shape
of procursus with transversal sclerotized ridges on retrolateral side and complex processes on prolateral
side; slender appendix) and by internal female genitalia (large lateral sclerites; shape and position of
pore plates; Figs 158, 186). From most congeners (except putatively closest relatives Ph. satun and Ph.
schwendingeri ) also by dorsally uniquely widened male palpal patella (Fig. 155) and by large sclerotized
‘knob’ on female external genitalia (Fig. 184).

38

HUBER B.A. et al., Pholcus halabala group in Southeast Asia

Etymology

The species name is derived from one of the localities where this species was found; noun in apposition.

Type material

THAILAND: holotype, 3 , ZFMK (Ar 15044), Krabi, near Khao Phanom Bencha National Park, Tham Khao
Phueng (8° 14.16’ N, 98°54.26’ E), 45 m a.s.l., on walls in cave, 8 Mar. 2015 (B.A. Huber, B. Petcharad).

Figs 143-152. Live specimens, Pholcus buatong Huber, sp. nov. (143-146), Ph. satun Huber, 2011
(147-149), and Ph. schwendingeri Huber, 2011 (150-152). 143-145. S3 and $ from Tham Buatong,
Thailand. 146. 3 from Tham Khao Phueng, Thailand. 147-149. Adult S, penultimate instar 3 , and
$ from Gunung Jerai, Malaysia. 150-151. 33 from Khao Sole, Thailand. 152. $ with egg-sac, from
Phanom Bencha, Thailand.

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European Journal of Taxonomy 190: 1-55 (2016)

Other material examined

THAILAND: 9 3 $ ?, 1 juv., ZFMK (8 2 $ Ar 15045) and PSUZC (1 1 ?), same data as

holotype; 2 SS, 3 $ §, in absolute ethanol, ZFMK (Mai 335), same data; 3 SS, 1 ?, ZFMK (Ar 15046),
~10 km N Krabi town, Tham Buatong (8° 10.83' N, 98°53.06' E), 50 m a.s.l., on cave walls, 7 Mar. 2015
(B.A. Huber, B. Petcharad); 3 $ $, in absolute ethanol, ZFMK (Mai 332), same data.

Description

Male (holotype)

Measurements. Total body length 4.0, carapace width 1.0. Leg 1: 44.3 (10.4 + 0.4 + 10.5 + 21.2 + 1.8),
tibia 2: 7.1, tibia 3: 4.2, tibia 4: 5.9; tibia 1 L/d: 120. Distance PME-PME 370 pm, diameter PME 90 pm,
distance PME-ALE ~35 pm; AME absent.

Color. Carapace pale ochre-grey with light brown posterior mark, ocular area with small median and
pair of lateral brown marks; clypeus not darkened; sternum pale gray with small very indistinct darker
marks; legs ochre-yellow with darker brown patellae and tibia-metatarsus joints; abdomen ochre-gray
with some black and indistinct whitish marks dorsally and laterally, monochromous ventrally.

Body. Habitus as in Figs 143-144; ocular area slightly raised and each triad on short stalk directed
obliquely dorsad (Fig. 159); carapace without median furrow; clypeus unmodified; sternum wider than
long (0.60/0.50), unmodified. Gonopore with four epiandrous spigots (Fig. 167). ALS with one widened,
one pointed, and six smaller cylindrically shaped spigots of varying sizes (Fig. 166).

Chelicerae. As in Fig. 156, barely modified, very indistinct frontal humps.

Palps. As in Figs 154-155; coxa unmodified; trochanter with long ventral apophysis and small hump
at its basis; femur with small retrolatero-dorsal process proximally (Fig. 162) and rounded protrusion
ventrally; patella dorsally characteristically widened, resulting in an angle between femur and patella
of-120° (lateral view); procursus very distinctive and complex, with transversal sclerotized ridges on
retrolateral side and complex processes on prolateral side (Figs 162-164); bulb elongate, with strong

Fig. 153. Known distributions of the Pholcus buatong species group (Malay Peninsula) and of the
Pholcus andulau species group (northern Borneo).

40

HUBER B.A. et al., Pholcus halabala group in Southeast Asia

proximal sclerite, slender and simple appendix (Fig. 161), without uncus, with long wealdy sclerotized
embolus distally transparent.

Legs. Without spines and curved hairs; few vertical hairs; retrolateral trichobothrium on tibia 1 at 4%;
prolateral trichobothrium absent on tibia 1, present on other tibiae; tarsus 1 with >20 pseudosegments,

Figs 154-158. Pholcus buatong Huber, sp. nov., ZFMK Ar 15045. 154-155. Left male palp, prolateral
and retrolateral views (arrow points at whitish membranous structure). 156. Male chelicerae, frontal
view. 157-158. Cleared female genitalia, ventral and dorsal views. Abbreviations: a = appendix;
b = genital bulb; e = embolus; p = procursus. Scale lines: 0.3 mm (156), 0.5 mm (154-155, 157-158).

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European Journal of Taxonomy 190: 1-55 (2016)

Figs 159-168. Pholcus buatong Huber, sp. nov., ZFMK Ar 15045.159-160. Male and female prosomata,
frontal views. 161. Left male palp, prolateral view. 162-164. Right male palp, retrolatero-dorsal, dorsal,
and retrolateral views. 165. Epigynum, ventral view. 166. Male ALS. 167. Male gonopore. 168. Comb-
hairs on male tarsus 4. Abbreviations: a = appendix; b = genital bulb; e = embolus; p = procursus;
pa = patella; tr = trochanter. Scale lines: 20 pm (166, 168); 40 pm (167); 200 pm (159-165).

42

HUBER B.A. et al., Pholcus halabala group in Southeast Asia

very indistinct. Tarsus 4 comb-hairs of the simplified Pholcus-typQ (cf. Huber & Fleckenstein 2008),
with four lateral tines (Fig. 168).

Male (variation)

Tibia 1 in 10 other males: 8.9-10.4 (mean 9.7).

Female

In general similar to male (Fig. 145) but eye triads on low humps and closer together (Fig. 160; PME-
PME distance: 200 pm). Tibia 1 in 5 females: 6.4-8.5 (mean 7.3). Epigynum weakly sclerotized flat
plate with large conspicuous sclerotized ‘knob’ (Fig. 165), anterior arch and lateral internal sclerites
visible through cuticle (Figs 157, 184-185); internal genitalia as in Figs 158 and 186, with large lateral
sclerites and oval pore plates in rather anterior position.

Natural history

At Buatong Cave, most specimens were found in barely visible domed webs close to the rock surface.
Only one specimen had its body tightly pressed against the rock surface. At Khao Phueng Cave,
specimens were abundant in the cave entrance area (again in very fine webs) but not deeper in the cave.

Distribution

Known from two neighboring localities in Krabi Province, southern Thailand (Fig. 153).

Pholcus satun Huber, 2011
Figs 147-149, 169-170, 187-189

Pholcus satun Huber, 2011: 144, figs 515-516, 580-582 (c?).

Diagnosis

Easily distinguished from all known congeners by long sickle-shaped bulbal process and by unique
shape of procursus (long, S-shaped, with subdistal ventral pointed process; fig. 580 in Huber 2011);
from other species in the buatong group also by longer than wide female internal genitalia (Fig. 170).

New material examined

THAIFAND: 1 $, in absolute ethanol, ZFMK (Mai 328), Satun, Thaleban National Park (6°42.6' N,
100° 10.2' E) (type locality), forest near headquarters, 110 m a.s.l., leaf litter, 5 Mar. 2015 (B.A. Huber,
B. Petcharad).

MAFAYSIA: 1 §, ZFMK (Ar 15047), Kedah, Gunung Jerai, forest near Sri Perigi Waterfall (5°48.3' N,
100°24.6' E), 100-200 m a.s.l., 27 Feb. 2015 (B.A. Huber); 2 1 juv., in absolute ethanol, ZFMK

(Mai 288), same data; 1 ZFMK (Ar 15048), same data, collected penultimate, adult on 2-3 Mar. 2015.

Description (female)

In general similar to male (Fig. 149; cf. Huber 2011), but eye triads on low humps and closer together
(distance PME-PME: 185 pm) and black mark at ocular area smaller. AME present as in male. Tibia 1 in
3 females: 5.5, 5.6, 6.1. Epigynum longer than wide, mostly wealdy sclerotized, with large dark ‘knob’
at posterior margin, anterior internal arch visible through cuticle (Figs 169, 187-188); posterior plate
laterally slightly more sclerotized. Internal genitalia as in Figs 170 and 189.

Distribution

Known from two localities in southern Thailand and northern mainland Malaysia (Fig. 153).

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European Journal of Taxonomy 190: 1-55 (2016)

Pholcus schwendingeri Huber, 2011
Figs 150-152, 171-183, 190-192

Pholcus schwendingeri Huber, 2011: 183-184, figs 761-763, 823-825 ((?).

Diagnosis

Easily distinguished from congeners by combination of very long male eye stalks (Figs 173-174), by
male palpal morphology (twisted segments; strong trochanter apophysis; procursus with large prolateral
process; simple appendix; absence of uncus; Figs 178-180; see also figs 823-824 in Huber 2011); from
other species in buatong group also by female internal genitalia (Fig. 192; wider than long in contrast to
Ph. satun ; with angular anterior ‘valve’ in contrast to Ph. buatong sp. nov.).

New material examined

THAILAND: 3 88, 9 ? ?, ZFMK (Ar 15049-50), Ranong, Klong Nakha Wildlife Sanctuary (9 0 27.6' N,
98°30.7' E) (type locality), 40 m a.s.l., forest, leaf litter, 12 Mar. 2015 (B.A. Huber, B. Petcharad); 4 $ §, in
absolute ethanol, ZFMK (Mai 362), same data; 20 SS, 12 $ §, ZFMK (Ar 15051-52), Surat Thani, Khao
Sole National Park, forest along nature trail (8 0 54.8' N, 98°29.3'-98°30.5' E), leaf litter, 110-160 m a.s.l.,
11-12 Mar. 2015 (B.A. Huber, B. Petcharad); 3 $ $, 3 juvs, in absolute ethanol, ZFMK (Mai 355), same

Figs 169-172. Cleared female genitalia, ventral and dorsal views. 169-170. Pholcus satun Huber, 2011,
ZFMK ‘Mai 328’. 171-172. Ph. schwendingeri Huber, 2011, ZFMK Ar 15050. Scale lines: 0.5 mm

44

HUBER B.A. et al., Pholcus halabala group in Southeast Asia

Figs 173-183. Pholcus schwendingeri Huber, 2011, ZFMK Ar 15051-52. 173-174. Male prosoma,
oblique and frontal views. 175. Female prosoma, frontal view. 176. Male eye triad. 177. Process between
male eye stalks. 178-179. Right genital bulb and procursus, prolateral and dorsal views. 180. Feft palp,
retrolateral view (arrow points at whitish membranous structure). 181. Male gonopore. 182. Epigynum,
ventral view. 183. Female AES. Abbreviations: a = appendix; b = genital bulb; e = embolus; p = procursus;
tr = trochanter. Scale lines: 10 pm (183); 20 pm (177); 50 pm (181); 80 pm (176); 200 pm (175, 178-
180, 182); 400 pm (173-174).

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European Journal of Taxonomy 190: 1-55 (2016)

data; 8 88, 10 ZFMK (7 88, 9 Ar 15053-54) and PSUZC (1 8, 1 ?), Krabi, Khao Phanom
Bencha National Park, trails near headquarters (8°14.1' N, 98°55.1' E), 150-300 m a.s.l., leaflitter in forest,
8 Mar. 2015 (B.A. Huber, B. Petcharad); 5 $5, 1 juv., in absolute ethanol, ZFMK (Mai 339), same data; 1
8, RMNH, Khao Phanom Bencha National Park, 15 Dec. 1990 (C.L. & PR. Deeleman).

Description

Male - amendments

Males from Khao Sok are significantly larger than males from Phanom Bencha: tibia 1 in 19 males from
Khao Sok: 7.4-8.9 (mean 8.0); in 7 males from Phanom Bencha: 6.6-7.3 (mean 7.0). This difference

Figs 184-192. Female genitalia, untreated in ventral view, cleared in ventral and dorsal views. 184-
186. Pholcus buatong Huber, sp. nov. 187-189. Ph. satun Huber, 2011. 190-192. Ph. schwendingeri
Huber, 2011.

46

HUBER B.A. et al., Pholcus halabala group in Southeast Asia

is even more pronounced in eye stalk length: in 19 males from Khao Sole: 0.70-0.78 (mean 0.74); in 8
males from Phanom Bencha: 0.53-0.60 (mean 0.55). Males from Klong Nakha seem to be intermediate
(low sample size). Angle between eye stalks varies even within localities (compare Figs 150 and 151).
All males with process between eye stalks (Fig. 177). Male tibia 2/tibia 4 length: 1.08. Patella dorsally
widened, resulting in an angle between femur and patella of-125° (lateral view; in figures 823 and 824
in Huber 2011, femora and patellae are not in perfect lateral view). Male gonopore with four epiandrous
spigots (Fig. 181). AFS as in female (see below).

Female

In general similar to male (Fig. 152), but eye triads on low humps and closer together (Fig. 175; distance
PME-PME: 185 pm); black marks in place of AME, but without lenses; clypeus unmodified. Tibia 1 in
10 females from Khao Sole: 5.6-6.3 (mean 6.0); in 9 females from Klong Nakha: 5.3-6.3 (mean: 5.7);
in 10 females from Phanom Bencha: 4.7-5.5 (mean 5.3). Epigynum wider than long, mostly weakly
sclerotized, with large dark ‘knob’ at posterior margin (Fig. 182), anterior internal arch visible through
cuticle (Figs 171, 190-191). Internal genitalia as in Figs 172 and 192. AFS with one widened, one
pointed, and several (apparently six) smaller cylindrically-shaped spigots of varying sizes (Fig. 183).

Natural history

At all three localities, the species was fairly abundant in suitable forest patches with large numbers of
large dead leaves on the ground. The poorly visible webs were closely attached to the lower leaf surface.
The spiders barely reacted to disturbance.

Distribution

Known from three localities in southern Thailand (Fig. 153).

Pholcus andulau species group

This species group is newly proposed to include one species previously included in the Ph. halabala
group (Ph. andulau Huber, 2011) and the newly described Ph. lambir Huber, sp. nov. They share three
putative synapomorphies: (1) the unique, partly sclerotized embolus with strong sclerotized pointed
processes (Figs 200, 209-212); (2) pointed male cheliceral apophyses directed toward each other and
without modified hairs (Figs 202, 214); and (3) large unsclerotized ‘knob’ on female external genitalia
directed toward anterior (Figs 203, 213). The two species are also otherwise very similar (females are
indistinguishable in the field; Ph. lambir sp. nov. males have a darker ocular area than Ph. andulau
males) and restricted to a limited geographic area in northern Borneo (Fig. 153). Preliminary molecular
data (A. Valdez-Mondragon, B.A. Huber & D. Dimitrov unpublished data) suggest a close relationship
with the Panjange nigrifrons group (which is also restricted to Borneo), but we know of no putative
morphological synapomorphy that would support this relationship. However, general habitus and
coloration are almost identical, and the same is true of web structure and microhabitat: in both groups, the
spiders build domed webs among the vegetation, with the apex of the sheet connected to the underside of
a leaf. In addition, they hang in their webs under the leaf rather than having their bodies pressed against
the leaf, and in both groups, cecidomyiid flies were often seen in large numbers hanging in the spider
webs. When disturbed, Pholcus andulau and Ph. lambir sp. nov. vibrate vigorously. Egg-sacs are carried
in front of the body (Figs 194, 196), as in typical pholcids.

Pholcus andulau Huber, 2011
Figs 193-194

Pholcus andulau Huber, 2011: 141-142, figs 502-504, 521-522, 570-574 ($$).

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European Journal of Taxonomy 190: 1-55 (2016)

Diagnosis

Easily distinguished from putatively closest known relative (Ph. lambir sp. nov.) by long dorsal process
on procursus (fig. 571 in Huber 2011), by smaller sclerotized teeth on male embolus (fig. 570 in Huber
2011), and by strongly curved anterior sclerite in internal female genitalia (fig. 573 in Huber 2011).

New material examined

MALAYSIA-BORNEO: 3 6 ??, ZFMK (Ar 15055-56), Sarawak, Gunung Mulu National Park,

Paku Waterfall Trail (4.037° N, 114.823° E), 60 m a.s.l., in domed webs under leaves, 23 Jul. 2014
(B.A. Huber, S.B. Huber); 3 $ $, in absolute ethanol, ZFMK (Bor 239), same data; 2 1 §, ZFMK

(Ar 15057), Gunung Mulu National Park, forest near Fagang Cave (4.051° N, 114.822° E), 60 m a.s.l.,
domed webs under leaves, 24 Jul. 2014 (B.A. Huber, S.B. Huber); 1 §, ZFMK (Ar 15058), Gunung
Mulu National Park, forest near Deer Cave (4.027° N, 114.818° E), 60 m a.s.l., night collecting, 24 Jul.
2014 (B.A. Huber); 1 RMNH, Gunung Mulu National Park, 12-22 Oct. 2003 (C.F. & PR. Deeleman).

BRUNEI: 1 $, RMNH, Temburong, Bukit Betoi, “87.04.23.06”.

Figs 193-199. Five specimens, Pholcus andulau group, and female genitalia of Ph. lambir Huber,
sp. nov. 193-194. Pholcus andulau Huber, 2011, S and $ with egg-sac from Gunung Mulu, Sarawak.
195-196. Ph. lambir sp. nov., $ and $ with egg-sac from Fambir, Sarawak. 197-199. Ph. lambir
sp. nov., female genitalia, untreated in ventral view, cleared in ventral and dorsal views.

48

HUBER B.A. et al., Pholcus halabala group in Southeast Asia

Distribution

Known from several localities in Brunei and northeastern Sarawak (Fig. 153).

Pholcus lamhir Huber, sp. nov.

urn:lsid:zoobank.org:act:2B969B98-2995-4325-911C-D3013371FFE3

Figs 195-218

Diagnosis

Easily distinguished from putatively closest known relative (Ph. andulau ) by very short dorsal (slightly
prolateral) process on procursus (Fig. 200), by much larger sclerotized teeth on male embolus (Figs 200,
209-212), and by less curved anterior sclerite in internal female genitalia (Figs 197-199).

Etymology

The species name is derived from the type locality; noun in apposition.

Type material

MAFAYSIA-BORNEO: holotype, ZFMK (Ar 15059), Sarawak, Fambir Hills National Park (4.198—
4.207° N, 114.034-114.045° E), 60-150 m a.s.l., in domed webs under leaves, 22 Jul. 2014 (B.A. Huber,
S.B. Huber).

Other material examined

MAFAYSIA-BORNEO: 9 $$, 13 55, 1 juv., ZFMK (8 $$, 12 55, Ar 15060-61) and SMK (1
1$), same data as holotype; 1 S, 2 5 5 , in absolute ethanol, ZFMK (Bor 200), same data; 3 SS, 8 5 ?,
ZFMK (Ar 15062-63), Sarawak, Niah Cave National Park, forest along main trail (3.814-3.821° N,
113.763-113.771° E), 20^10 m a.s.l., domed webs under leaves, 27 Jul. 2014 (B.A. Huber, S.B. Huber).

Description

Male (holotype)

Measurements. Total body length 5.0, carapace width 1.05. Teg 1: 48.4 (10.6 + 0.5 + 11.0 + 23.5 + 2.8),
tibia 2: 6.9, tibia 3: 4.0, tibia 4: 6.2; tibia 1 F/d: 100. Distance PME-PME 380 pm, diameter PME
100 pm, distance PME-AFE ~40 pm; AME absent (only small irregular internal black marks).

Color. Carapace pale ochre-yellow, ocular area and clypeus brown; sternum whitish; legs ochre-yellow
to orange, with brown patellae and tibia-metatarsus joints; abdomen ochre-gray with some darker marks
dorsally, monochromous ventrally.

Body. Habitus as in Fig. 195; ocular area slightly raised and each triad on short stalk directed obliquely
dorsad (Fig. 205); carapace without median furrow; clypeus unmodified; sternum wider than long
(0.80/0.65), unmodified. Gonopore with four epiandrous spigots (Fig. 216). AES with one widened, one
pointed, and six smaller cylindrically shaped spigots of varying sizes (Fig. 218).

Chelicerae. As in Fig. 202, with distinctive pair of pointed frontal apophyses directed toward each other
(Fig. 214); indistinct lateral processes.

Palps. As in Figs 200-201; coxa unmodified; trochanter with retrolatero-ventral apophysis; femur with
distinctive ventral process directed proximad; procursus relatively simple, with small semitransparent
prolateral process (Figs 207-209) and distinctive sclerotized and membranous distal elements; bulb
large, with strong proximal sclerite, with uncus, with proximally heavily sclerotized embolus provided
with strong teeth (Figs 209-212).

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European Journal of Taxonomy 190: 1-55 (2016)

Figs 200-204. Pholcus lambir Huber, sp. nov., ZFMK Ar 15060-61. 200-201. Left male palp, prolateral
and retrolateral views. 202. Male chelicerae, frontal view. 203-204. Cleared female genitalia, ventral
and dorsal views. Abbreviations: b = genital bulb; e = embolus; p = procursus; u = uncus. Scale lines:
0.3 mm (202-204), 0.5 mm (200-201).

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HUBER B.A. et al., Pholcus halabala group in Southeast Asia

Legs. 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 pseudosegments very
indistinct, only distally a few visible in dissecting microscope. Tarsus 4 comb-hairs of the simplified
Pholcus-typQ (cf. Huber & Fleckenstein 2008), with four lateral tines (Fig. 215).

Male (variation)

Tibia 1 in 9 other males: 9.6-11.7 (mean 10.5).

Female

In general similar to male (Fig. 196), but eye triads on low humps and closer together (Fig. 206;
PME-PME distance: 200 pm), clypeus darker than in males. Tibia 1 in 20 females: 6.8-8.8 (mean
8.0). Epigynum wealdy sclerotized flat plate with large unsclerotized ‘knob’ directed toward anterior
(Fig. 213), anterior internal sclerite visible through cuticle (Figs 197-198, 203); internal genitalia as in
Figs 199 and 204. ALS as in male (Fig. 217).

Figs 205-210. Pholcus lambir Huber, sp. nov., ZFMK Ar 15060-61. 205-206. Male and female
prosomata, frontal views. 207. Right procursus, retrolateral (slightly dorsal) view. 208. Left procursus
(and genital bulb), prolateral view. 209. Right procursus (and embolus sclerite), distal view. 210. Right
embolus sclerite, prolateral view. Abbreviations: b = genital bulb; e = embolus; p = procursus. Scale
lines: 50 pm (210); 100 pm (207-209); 200 pm (205-206).

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European Journal of Taxonomy 190 : 1-55 ( 2016 )

Natural history

Spiders were collected from domed webs among the vegetation, at approximately 1-2 m above the
ground. The apex of the sheet was connected to the underside of a leaf, but spiders hung in their webs
under the leaf rather than having their bodies pressed against the leaf In some webs, cecidomyiid flies

Figs 211-218. Pholcus lambir Huber, sp. nov., ZFMK Ar 15060-61. 211-212. Left bulbal processes,
prolateral (slightly distal) and ventral views (arrows point at sperm duct opening). 213. Epigynum,
ventral view. 214. Distal male cheliceral apophyses. 215. Comb-hairs on male tarsus 4. 216. Male
gonopore. 217-218. Female and male ALS. Abbreviations: e = embolus; u = uncus. Scale lines: 10 pm
(215, 218); 20 pm (217); 30 pm (214, 216); 60 pm (211); 80 pm (212); 200 pm (213).

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HUBER B.A. et al., Pholcus halabala group in Southeast Asia

were seen and collected in large numbers. When disturbed, the spiders vibrated vigorously. Egg-sacs
were carried in front of the body (Fig. 196), as in typical pholcids; they are slightly elongate and contain
-25-30 eggs.

Distribution

Known from two localities in northeastern Sarawak (Fig. 153).

Discussion

An unrevised genus with more than 300 described and probably hundreds of undescribed species like
Pholcus is notoriously difficult to handle: internal relationships are mostly entirely obscure; decisions
whether newly collected specimens represent new species or not are difficult or impossible to make;
and biological peculiarities cannot be interpreted in an evolutionary context. For more than 200 years,
isolated species descriptions were added to Pholcus , not only without any phylogenetic framework,
but often even without the chance to compare new species with the most similar ‘known’ congeners.
The result was more akin to a long list of biologically relatively meaningless names than to testable
hypotheses about natural groups, species limits, and evolutionary trends within the genus. This was
unfortunate not only because it led to synonyms and chaos, but mainly because Pholcus is characterized
by numerous biologically interesting phenomena whose evolution will only be understood in the context
of a phylogenetic framework. For example, (1) several different kinds of male ‘head’ modifications
have evolved within the genus, including spines, hooks, eye stalks, and curiously modified hair brushes
(Huber 2011), but the evolution of these structures is barely understood; (2) different Pholcus species
appear to specialize in specific microhabitats, such as leaf litter, dark and sheltered spaces under logs and
rocks, open space among vegetation, undersides of green leaves, and rock surfaces. Shifts among these
microhabitats and the potential effects of the underlying evolutionary ecological flexibility on species
diversification remain largely unknown (Dimitrov et al. 2013).

A first major effort to bring structure into the taxonomic chaos of Pholcus was made in Huber (2011).
Many ‘old’ and poorly known species were redescribed; first SEM data were provided for numerous
species; and the genus was divided into 29 operational species groups, based partly on synapomorphies
derived from a first cladistic analysis, partly on putative synapomorphies not derived from formal
analysis, and partly just on similarity and geographic closeness. Obviously, this preliminary analysis was
just a first step that requires substantial further refinement to approximate the true phylogenetic history
of the genus. More light could have been shed if the nine taxonomic contributions on Pholcus published
since 2011 (mostly on Chinese, Korean, and Vietnamese species) had considered anything beyond pure
alpha taxonomy, such as phylogenetic relations, distribution patterns, diversity patterns, evolution of
character systems, or any other questions of biological interest. Instead, none of them even attempted
to place new species into existing species groups. We do not claim that such purely descriptive work is
worthless, but we agree with Darlington (1971: 347) that “describing of scattered new species in poorly
known groups is often not useful” and with Mayr & Ashlock (1991: 347) that “isolated description [... ]
divorced from revisional or monographic work is the least desirable form of taxonomic publication”.

This contribution deals mainly with the Pholcus halabala group, originally considered one of the most
problematic (i.e., unnatural or non-monophyletic) species groups in Pholcus. The combination of field
observations (web design and structure, microhabitat, egg-sac holding position, life color pattern),
ultrastructure (silk spigots, male cheliceral apophyses), and general morphology (abdomen shape) have
prompted us to completely rearrange the group and to propose three new species groups. We explicitly
point out the remaining gaps that need to be addressed, and highlight a number of potentially useful
characters (those listed above) in this and related species groups for future formal cladistic analyses.
The fact that some of these characters can only be observed in life specimens in the field underlines

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European Journal of Taxonomy 190 : 1-55 ( 2016 )

the importance of continued and focused field work beyond the “accumulation of dead specimens” (cf.
Darlington 1971). Future work should especially focus on the Pholcus quinquenotatus group. Three
of the species we newly assign to the Ph. halabala group were originally (Huber 2011) proposed as
representatives of the Ph. quinquenotatus group. This leaves three species in the Ph. quinquenotatus
group, but field observations (and molecular data) may suggest that at least two of them are also part
of the Ph. halabala group (Ph. quinquenotatus Thorell, 1878; Ph. namou Huber, 2011), making the Ph.
quinquenotatus group obsolete.

Acknowledgements

We thank David Court, Venus Saksongmuang, Mustakiza Muslimin, Kamil A. Braima, and Samuel B.
Huber for accompanying us in the field; our Thai drivers Usman Hayee-Aarwae, Jeerabute Suwannatut,
and Preesha Petsout for bringing us safely to all requested places; Bjorn Rulik for identifying the flies
found in pholcid webs; Sara Bumrungsri for supporting our permit requests in Thailand; Noraishah My din
Abdul Aziz for facilitating field work in Malaysia; the Sarawak Museum Department for assistance in
field logistics; the National Research Council of Thailand (NRCT), the Singapore National Parks Board
(Permit No. NP/RP15-003a) and the Sarawak Forest Department (Permit No. NCCD.907.4.4 JTD.10-
211, Park Permit No. 174/2014) for issuing the relevant permits; two anonymous reviewers for valuable
co mm ents on a previous version of the manuscript; and the German Research Foundation for financial
support (DFG Project HU 980/11-1).

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Manuscript received: 9 October 2015
Manuscript accepted: 19 December 2015
Published on: 14 April 2016

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Topic editor: Rudy Jocque
Desk editor: Charlotte Thionois

HUBER B.A. et al., Pholcus halabala group in Southeast Asia

Printed versions of all papers are also deposited in the libraries of the institutes that are members of the
EJT consortium: Museum national d’Histoire naturelle, Paris, France; Botanic Garden Meise, Belgium;
Royal Museum for Central Africa, Tervuren, Belgium; Natural History Museum, London, United
Kingdom; Royal Belgian Institute of Natural Sciences, Brussels, Belgium; Natural History Museum of
Denmark, Copenhagen, Denmark.

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