European Journal of Taxonomy 225: 1-34

http://dx.doi.org/10.5852/ejt.2016.225

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.

Research article

urn:lsid:zoobank.org:pub:C 1F4C6FC-2EB6-48D9-B628-F1C8BD1FF2F8

The Philippine hair wax spiders and their relatives:

revision of the Pholcus bicornutus species group (Araneae, Pholcidae)

Bernhard A. HUBER 1 *, Olga M. NUNEZA 2 & Charles LEH MOI UNO 3

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

department of Biological Sciences, Mindanao State University - Iligan Institute of Technology,

Tibanga, Iligan City 9200, Philippines.

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

* Corresponding author: b.huber@zfmk. de

2 E-mail: [email protected]

3 E-mail: [email protected]

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

2 urn: lsid:zoobank.org: author: 6D14FCCC-F148-4293 -9F23 -0EB5 A4119305

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

Abstract. We revise the Southeast Asian Pholcus bicornutus group in which males are characterized by

a unique pair of horns on their ocular area, each of which carries at its tip a brush of hairs. In two species,

the two hair brushes are ‘glued’ or ‘waxed’ together by an unidentified substance into a very consistently

curved and pointed single median tip. In the other five species known, the hairs are unglued. We present

a first revision of ocular modifications in Pholcidae and identify twenty supposedly independent origins.

Most cases are in Pholcinae, and all but one case are limited to the male, suggesting sexual selection

as the main driving force in the evolution of ocular modifications in Pholcidae. Previously, the Pholcus

bicornutus group consisted of four species limited to the Philippines. We describe four new species,

including three species from the Philippines (P. olangapo Huber, sp. nov.; P. kawit Huber, sp. nov.;

P. baguio Huber, sp. nov.) and the first representative from outside the Philippines (P. mulu Huber,

sp. nov. from Sarawak, NE Borneo) and provide new records and SEM data for three previously

described species.

Keywords. Pholcus , taxonomy, ocular area, sexual dimorphism, ultrastructure.

Huber B.A., Nuneza O.M. & Leh Moi Ung C. 2016. The Philippine hair wax spiders and their relatives: revision

of the Pholcus bicornutus species group (Araneae, Pholcidae). European Journal of Taxonomy 225: 1-34. http://

dx.doi.org/10.5852/eit.2016.225

Introduction

When Eugene Simon returned from his expedition to Luzon in 1891, he brought with him, among

others, a highly unusual pholcid spider species from caves near Manila. Unlike any other pholcid known

at that time, males had a pair of seemingly segmented horns at their ocular region. Simon was clearly

European Journal of Taxonomy 225 : 1-34 ( 2016 )

struck by this feature, and devoted the only two figures in the original description of this new species

Pholcus bicornutus Simon, 1892 to the male ocular region (Simon 1892: figs 3—4). One year later, in

his epochal Histoire Naturelle des Araignees, he used one of these figures again (Simon 1893: fig. 455).

In both editions, the two-segmented horns are illustrated as being more or less parallel in frontal view,

with separate tips.

More than 100 years later, when preparing the first revision of Pholcus Walckenaer, 1805 (Huber

2011a), one of us (BAH) studied Simon’s type material ofP bicornutus. While one mystery of Simon’s

‘segmented horns’ could be solved, another one appeared. Scanning electron microscopy revealed that

what had looked to Simon like ‘distal segments’ were in fact brushes of hairs that originated at the tips of

the ‘basal segments’. Strikingly, however, the tips of these hair brushes were not separate as illustrated by

Simon but appeared like ‘glued’ or ‘waxed’ together at their tips (Huber 2011a: figs 1566, 1568). Simon

had clearly interpreted this as an artifact, and the latter author tended in the same direction, especially

because in all specimens of two newly described close relatives of P. bicornutus (P. arayat Huber, 2011;

P. pagbilao Huber, 2011) the hair brushes on the otherwise very similar ocular horns did not look like

they were glued together. In addition, sample sizes were small and no scanning electron images could

be made of the two new species, so Huber (2011a) did not further dwell on the mysteriously joined hair

brushes of P. bicornutus.

The present paper revises the Pholcus bicornutus group based on much larger samples than previously

available and finally presents strong evidence that the ‘glued’ or ‘waxed’ hair brushes of P. bicornutus

are not an artifact. First, in new material of P. bicornutus and of a seemingly very closely related species

(P. olangapo sp. nov.), all twelve available males have the exact same configuration, with the hair

brushes joined together and curved in a highly consistent way. Second, photographs of live males of

P. olangapo sp. nov. show that the joined tips are not an artifact of fixation and preservation. And finally,

in none of over 100 studied males of P. arayat , P. pagbilao , and three newly described species, do

the hairs appear glued together in the light microscope, and this impression is supported by scanning

electron micrographs of three of these species.

Material and methods

Most of the material studied herein was collected during recent expeditions to the Philippines (Feb-

Mar. 2014) and to northern Borneo (Jul.-Aug. 2014). Further specimens came from a student project at

Mindanao State University - Iligan Institute of Technology dealing with ecological aspects of pholcid

diversity in the Philippines. The material is currently deposited at Mindanao State University - Iligan

Institute of Technology (MSU-IIT), Philippines; Sarawak Museum, Kuching (SMK), Malaysia; and

Zoologisches Forschungsmuseum Alexander Koenig, Bo nn (ZFMK), Germany. Additional material

came from the California Academy of Sciences, San Francisco (CAS), U.S.A.; the Netherlands Centre

for Biodiversity Naturalis, Teiden (RMNH); and the Museum of Zoology, Turku University (ZMT),

Finland.

Methods and terminology used are as in Huber (2011a). Measurements are in mm unless otherwise

specified. Eye measurements are approx. ± 5 pm. Epigyna were cleared in 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.

HUBER B.A. et al., Philippine hair wax spiders

Abbreviations

ALE

ALS

AME

a.s.l.

L/d

PME

PMS

appendix

anterior lateral eye(s)

anterior lateral spinneret(s)

anterior median eye(s)

above sea level

genital bulb

embolus

femur

hinge

length/diameter

procursus

posterior median eye(s)

posterior median spinneret(s)

trochanter

uncus

Results

Class Arachnida Cuvier, 1812

Order Araneae Clerck, 1757

Family Pholcidae C.L. Koch, 1851

Pltolcus Walckenaer, 1805

Pholcus Walckenaer, 1805: 80. Type species by (assumed) monotypy: Aranea phalangioides Fuesslin,

1775.

Pholcus — Huber 2011a: 124-126.

Pholcus bicornutus species group

Diagnosis

Large, long-legged Pholcus (body length ~5-8, male leg 1 length: -40-60) with cylindrical abdomen,

eight eyes, and rather dark coloration; distinguished from similar insular Southeast Asian species groups

(P. gracillimus group; P. ancoralis group; cf. Huber 2011a) by male ocular area with unique pair of

processes, each carrying brush of hairs (Figs 12-13, 63-67, 78, 92) (P. ancoralis group also with horns

but without brushes of hairs), by male palpal tibia very large relative to femur (in contrast to P. ancoralis

species group), and by paired anterior marks in dorsal abdominal pattern (Figs 3, 36, 50, 55; unpaired in

P. gracillimus and P ancoralis groups).

Description - amendments

The original description of the species group (Huber 2011a) is still largely valid. The following are minor

updates and additions: body length -5-8, carapace width 1.3-1.8. Male distal cheliceral apophyses

provided with 2-3 modified hairs each (Figs 20, 73, 85, 96). Tibia 1 length in males -9.5-16.0, in

females -7.5-14.0; male tibia 1 L/d -65-80; legs either with curved hairs (tibiae and metatarsi 1-2;

P. bicornutus , P. olangapo sp. nov.) or without curved hairs (other species). Abdomen cylindrical, with

cuticular color pattern dorsally (also in P. bicornutus ), anterior element always paired. ALS with one

widened, one pointed, and variable number of smaller spigots: either six cylindrically-shaped spigots

European Journal of Taxonomy 225: 1-34 (2016)

of ‘usual’ size (i.e., -3-8 pm wide and -5-8 jam long; Figs 19, 21, 100, 102), or only three to five very

small conical spigots (i.e., -1.5-3 pm wide and -3-6 pm long; Figs 74, 76, 88, 90).

Composition

The eight species assigned to this group are here divided into two operational sub-groups:

(1) A sub-group of two species (P. bicornutus Simon, 1892; P. olangapo sp. nov.), sharing the following

similarities: absence of male bulbal appendix (Fig. 7); procursus with hinged distal element and with

distinctive prolatero-dorsal process (Figs 8, 17); male palpal tibia ventrally unmodified; hairs of ocular

processes ‘glued’ or ‘waxed’ together (Figs 12-13, 15); epigynum with small epigynal plate and large

membranous area in front of plate (Fig. 24); proximal lateral processes on male chelicerae large (Fig. 9);

and presence of curved hairs on tibiae and metatarsi 1-2. A third (unnamed) species occurs on Negros

Island (“sp. Phi 190” in Fig. 1; 1 §, 1 juv. from Twin Takes National Park, in ZFMK).

(2) A sub-group of six species (P. arayat Huber, 2011; P. mulu sp. nov.; P. baguio sp. nov.; P. kawit sp.

nov.; P. pagbilao Huber, 2011; P. schawalleri Yao et al., 2014), sharing the alternative character states:

presence of male bulbal appendix (Figs 40, 45); procursus without hinged distal element and without

dorsal process (Figs 41,46); male palpal tibia with small ventral cavity to accommodate proximal bulbal

sclerite in resting position (arrow in Fig. 45); hairs of ocular processes not ‘glued’ or ‘waxed’ together

(Figs 65-66, 78); epigynum with large epigynal plate (Figs 27, 30, 33); proximal lateral processes

on male chelicerae small (Figs 42, 47); no curved hairs on legs. A possible further species occurs on

Mindanao, but it is represented by two juvenile specimens only (“sp. Phi 254” in Fig. 2; from Marilog

Figs 1-2. Known distributions of the two subgroups of the Pholcus bicornutus species group.

HUBER B.A. et al., Philippine hair wax spiders

Distr., Baganihan, in ZFMK). The undescribed species from Mt. Apo mentioned in Huber (2011a: 315)

was restudied and found to belong to another species group.

Natural history

Seven of the eight named species were observed in the field. They occupied a variety of microhabitats,

ranging from caves (close to the ground; P. bicornutus ) and wet rock walls (P. olangapo sp. nov.) to

dark sheltered spaces among rocks (P. mulu sp. nov., P. baguio sp. nov., P. pagbilao ), large holes at

tree bases (P. kawit sp. nov.), abandoned buildings (P. pagbilao ), and open space among vegetation

exposed to direct sunlight (P. arayat). Webs mainly consisted of a domed sheet with a diameter of up

to ~50 cm (P. mulu sp. nov.), but were much smaller in the rock-dwelling P. olangapo sp. nov. (~15 cm).

In P. baguio sp. nov., webs were connected to each other and formed large communal structures up to

2 m in length. When disturbed, most species were observed to run toward the periphery of the web,

seeking shelter at the substrate rather than staying and whirling in the web.

Distribution

The Pholcus bicornutus group is largely restricted to the Philippines, with only one species in NE

Borneo (Figs 1-2).

Pholcus bicornutus Simon, 1892

Pholcus bicornutus Simon, 1892: 41—42, pi. 2, figs 3^1.

Pholcus bicornutus - Simon 1893: 466M68, figs 455, 458. —Huber 2011a: 315-318, figs 1509-1513,

1539-1540, 1560-1573.

Diagnosis (updated)

Distinguished from most similar known relative (P. olangapo sp. nov.) by absence of dark lateral bands

on carapace (in males and females), and by procursus shape (prolatero-dorsal process of proximal part

longer and more slender; entire distal hinged part of procursus longer and more slender; distinctive

shapes of procursus tip and of uncus; figs 1560-1561 in Huber 2011a). From other species of the

P. bicornutus group (P. pagbilao , P. arayat , P. schawalleri , P. baguio sp. nov., P. mulu sp. nov., P. kawit

sp. nov.) by hinged procursus, small epigynal plate, undivided dark band ventrally on abdomen (also in

P. kawit sp. nov. and P. mulu sp. nov.), and by presence of slightly curved hairs on legs (especially on

tibiae and metatarsi 1-2).

New material examined

PHIFIPPINES: 1 8 § ?, Fuzon, Rizal Prov., near Antipolo, Mystical Cave (14.606° N, 121.209° E),

160 m a.s.l., in cave near entrance, 11 Mar. 2014 (B.A. Huber), ZFMK (Ar 15495); 2 $$, 2 juvs, in

absolute ethanol, same data, ZFMK (Phi 185).

Note

Simon (1892) described this species from two caves, “Cueva de San Mateo” and “Cueva de Antipolo”.

The first is possibly identical to what is now called “Pamitinan Cave” (14.731° N, 121.190° E) which

in turn may be identical to what in Huber (2011a) is cited as “Montealban Cave” (correct spelling:

“Montalban Cave”). The second is presumably identical to what is now called “Mystical Cave”

(14.606° N, 121.209° E). The distance between these two caves is just 14 km. However, as noted

previously after examination of Simon’s type specimens (Huber 2011a), males from the two caves differ

slightly in details of the procursus, and the specimens from Mystical Cave are assigned tentatively to this

species (the lectotype, designated in Huber 2011a, is from “Cueva de San Mateo”).

European Journal of Taxonomy 225: 1-34 (2016)

Variation

Tibia 1 in newly examined male: 12.9; in 6 newly examined females: 10.4-12.0 (mean: 11.2).

Natural history

The spiders were found within the cave close to the cave entrance; no specimen was found deeper in the

cave. The spiders built their domed sheets among rocks close to the ground.

Distribution

Known from two caves near Manila only (Fig. 1).

Pholcus olangapo Huber, sp. nov.

urn:lsid:zoobank.org:act:lCD3A2E0-1528-4D37-9BFD-82B752CA13C5

Figs 3-26

Diagnosis

Distinguished from most similar known relative (P. bicornutus ) by dark lateral bands on carapace (in

males and females; Figs 3, 6), and by procursus shape (prolatero-dorsal process of proximal part shorter

Figs 3-6. Live specimens, Pholcus olangapo Huber, sp. nov., Pamulaklakin Forest Trail. 3-4. $ and $

in their natural habitat. 5-6. $ and $ photographed on a dead leaf.

HUBER B.A. et al. , Philippine hair wax spiders

and wider; entire distal hinged part of procursus shorter and wider; distinctive shapes of procursus

tip and of uncus; Figs 7-8). From other species of the P. bicornutus group (P. pagbilao , P. arayat ,

P. schawalleri , P. baguio sp. nov., P. mulu sp. nov., P. kawit sp. nov.) by hinged procursus, small epigynal

plate (Fig. 24), undivided dark band ventrally on abdomen (also in P. kawit sp. nov. and P. mulu sp. nov.),

and by presence of slightly curved hairs on legs (especially on tibiae and metatarsi 1-2).

Figs 7-11. Pholcus olangapo Huber, sp. nov., ZFMK Ar 15497-98. 7-8. Feft male palp, prolateral and

retrolateral views (asterisk marks prolatero-dorsal process of procursus). 9. Male chelicerae, frontal view.

10-11. Cleared female genitalia, ventral and dorsal views. Scale lines: 7-8 = 1 mm; 9-11 = 0.5 mm.

European Journal of Taxonomy 225: 1-34 (2016)

Etymology

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

Material examined

Holotype

PHILIPPINES: S, Luzon, Bataan Prov., near Olangapo, along river near Pamulaklakin Forest Trail

(14.798° N, 120.338° E), 80 m a.s.l., on wet rock wall, 28 Feb. 2014 (B.A. Huber), ZFMK (Ar 15496).

Figs 12-15. Pholcus olangapo Huber, sp. nov., ZFMK Ar 15497-98.12. Male prosoma, frontal view. 13,

15. Male ocular horns with modified and ‘glued’ or ‘waxed’ hairs, frontal and lateral views. 14. Female

prosoma, frontal view. Scale lines: 12, 14 = 400 pm; 13 = 50 pm; 15 = 60 pm.

HUBER B.A. et al., Philippine hair wax spiders

Other material

PHILIPPINES: 9 SS, 10 $$, 3 juvs, ZFMK (Ar 15497-98) and 1 1 $, MSU-IIT, same data as

holotype; 2 $ ?, 5 juvs, in absolute ethanol, same data, ZFMK (Phi 207).

Figs 16-23. Pholcus olangapo Huber, sp. nov., ZFMK Ar 15497-98. 16. Right genital bulb, prolateral

view. 17. Left tarsus and procursus, dorsal view (asterisk marks prolatero-dorsal process of procursus).

18. Tip of left procursus, dorsal view. 19. Female spinnerets. 20. Male cheliceral apophysis (arrows

point at modified hairs). 21. Male ALS. 22. Male gonopore. 23. Epigynum. Scale lines: 16,23 = 200 pm;

17 = 300 pm; 18 = 100 pm; 19 = 70 pm; 20-21 = 20 pm; 22 = 40 pm.

European Journal of Taxonomy 225: 1-34 (2016)

Figs 24-35. Epigyna, ventral views, and cleared female genitalia, ventral and dorsal view. 24-26. Pholcus

olangapo Huber, sp. nov., ZFMK Ar 15498. 27-29. Pholcus mulu Huber, sp. nov., ZFMK Ar 15502.

30-32. Pholcus kawit Huber, sp. nov., ZFMK Ar 15505. 33-35. Pholcus baguio Huber, sp. nov., ZFMK

Ar 15508.

HUBER B.A. et al., Philippine hair wax spiders

Assigned tentatively

PHILIPPINES: 1 $, Luzon, Laguna Prov., Mt. Banahaw, forest near Taytay Falls (14.110° N, 121.507° E),

560 m a.s.l., 26 Feb. 2014 (B.A. Huber), ZFMK (Ar 15499).

Description

Male (holotype)

Measurements. Total body length 6.0, carapace width 1.6. Leg 1: 42.7 (10.1 + 0.7 + 10.4 + 19.0 +

2.5), tibia 2: 7.2, tibia 3: 4.8, tibia 4: 6.7; tibia 1 L/d: 64. Distance PME-PME 370 pm, diameter PME

150x180 pm, distance PME-ALE ~40 pm, distance AME-AME 30 pm, diameter AME 100 pm.

Color. Carapace pale ochre to light brown, with wide median dark brown mark including ocular area

and connecting posteriorly to wide lateral marginal bands (Fig. 3); clypeus pale ochre; sternum brown,

with three pairs of light ochre marginal marks at bases of coxae 2-4; legs light brown to ochre, dark rings

subdistally on femora and tibiae and in patella area, tips of femora and tibiae lighter; abdomen ochre-

gray, dorsally with several pairs of indistinct cuticular marks, with distinct internal darker marks visible

through cuticle dorsally and laterally; ventrally with undivided wide brown band between gonopore and

spinnerets.

Body. Habitus as in Figs 3 and 5; ocular area raised, each eye triad on additional short hump directed

towards lateral (Fig. 12), with pair of long processes between eye triads, each with distal brush of hairs

that are ‘glued’ or ‘waxed’ together to form consistently curved pointed tip (Figs 13, 15); carapace

without median furrow; clypeus unmodified; sternum wider than long (1.00/0.75), unmodified. ALS

with one widened, one pointed, and six cylindrically shaped spigots of variable sizes (Fig. 21). Gonopore

with four epiandrous spigots (Fig. 22).

Chelicerae. As in Fig. 9, with large lateral apophyses and low frontal humps proximally, dark distal

apophyses near median line provided with two small modified (cone-shaped) hairs each (Fig. 20);

without stridulatory ridges.

Palps. As in Figs 7-8; coxa unmodified; trochanter with retrolateral pointed process; femur with distinct

ventral process and indistinct retrolateral hump proximally; tibia with dark dorsal band, without ventral

cavity; procursus complex and hinged, proximal part with distinctive prolatero-dorsal process (Fig. 17),

distal part with prolateral membranous flap and distinctive tip (Fig. 18); bulb with long, distinctively

curved uncus, wealdy sclerotized embolus distally curved, without appendix (Fig. 16).

Legs. Without spines; with weakly curved hairs on tibiae and metatarsi 1-2; few vertical hairs; retrolateral

trichobothrium on tibia 1 at 4%; prolateral trichobothrium absent on tibia 1, present on other tibiae;

tarsus 1 with >30 pseudosegments, only distally about 10 fairly distinct.

Male (variation)

Tibia 1 in 10 other males: 9.6-11.4 (mean: 10.4).

Female

In general similar to male but eye triads closer together (distance PME-PME 210 pm), without processes

between eye triads (Fig. 14). Tibia 1 in 11 females from type locality: 8.6-10.5 (mean: 9.2). Epigynum

mostly weakly sclerotized, small posterior plate with small ‘knob’ (Figs 23-24); some females with

strongly protruding epigynal area (apparently expandable); internal genitalia as in Figs 11 and 26.

Spinnerets as in Fig. 19, ALS with one widened, one pointed, and six cylindrically shaped spigots of

variable sizes, PMS with two spigots each. The single female specimen from Mt. Banahaw shares the

carapace pattern and is thus tentatively assigned to this species rather than to P. bicornutus (in which the

European Journal of Taxonomy 225: 1-34 (2016)

female genitalia appear indistinguishable; compare figs 1539 and 1564 in Huber 201 la with Figs 11 and

24 herein); tibia 1: 9.7.

Natural history

At the type locality, specimens were collected from small webs attached to a perpendicular, dripping wet

rock wall beside a small stream (Figs 3—4).

Distribution

Known from two localities in Luzon only (single female specimen from Mt. Banahaw assigned

tentatively, see above; Fig. 1).

Pholcus mulu Huber, sp. nov.

um:lsid:zoobank.org:act:36311195-45A6-4B19-BFD2-31C0687775FE

Figs 27-29, 36-37, 40M4

Diagnosis

Easily distinguished from most similar known relatives (species with horns between eye triads carrying

brushes of unglued hairs and with simple, unhinged procursi: P. arayat , P. pagbilao , P. schawalleri ,

P. baguio sp. nov., P. kawit sp. nov.) by large round uncus with pointed process (Fig. 40) and by large

prolateral sclerite distally on procursus (Fig. 40); from most species (except P. kawit sp. nov.) also by

undivided median dark band ventrally on abdomen (Fig. 37). From other species of the P. bicornutus

group (P. bicornutus and P. olangapo sp. nov.) by unhinged procursus, presence of appendix, large

epigynal plate, and by absence of curved hairs on legs.

Etymology

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

Material examined

Holotype

MALAYSIA-BORNEO: S, Sarawak, Gunung Mulu National Park, forest near Deer Cave (4.027° N,

114.818° E), 60 m a.s.l., among rocks, 23 Jul. 2014 (B.A. Huber, S.B. Huber), ZFMK (Ar 15500).

Other material

MALAYSIA-BORNEO: 2 SS, 10 55, ZFMK (Ar 15501-02) and 1 <?, 1 5, SMK, same data as

holotype; 1 $, 1 $, 1 juv., in absolute ethanol, same data, ZFMK (Bor 238); 2 4^5,1 juv., Gunung

Mulu National Park, forest near Lagang Cave (4.051° N, 114.822° E), 60 m a.s.l., domed webs among

rocks, 24 Jul. 2014 (B.A. Huber, S.B. Huber), ZFMK (Ar 15503).

Description

Male (holotype)

Measurements. Total body length 6.8, carapace width 1.7. Leg 1: 62.9 (14.4 + 0.8 + 15.1 + 28.5 + 4.1),

tibia2: 9.6, tibia 3: 6.0, tibia4: 8.4; tibia 1 L/d: 94. Distance PME-PME 480 pm, diameter PME 180 pm,

distance PME-ALE ~40 pm; distance AME-AME 50 pm, diameter AME 80 pm.

Color. Carapace pale ochre, with wide median brown mark including ocular area, without lateral marks

(Fig. 36); clypeus not darkened; sternum dark brown; legs in live specimens bluish, in alcohol ochre to

light brown, tips of femora and tibiae whitish; abdomen pale gray, dorsally with dark cuticular marks

that are fused above spinnerets, with indistinct internal darker marks visible through cuticle dorsally and

laterally; ventrally with undivided wide brown band between gonopore and spinnerets.

HUBER B.A. et al., Philippine hair wax spiders

Body. Habitus as in Figs 36-37; ocular area raised, each eye triad on additional short hump directed

towards lateral, with pair of processes between eye triads, each with distal brush of hairs; carapace

without median furrow; clypeus unmodified; sternum wider than long (1.00/0.85), unmodified.

Chelicerae. As in Fig. 42, with small lateral and frontal apophyses proximally and dark distal apophyses

near median line provided with two small modified (cone-shaped) hairs each; without stridulatory ridges.

Palps. As in Figs 40^41; coxa unmodified; trochanter with retrolateral process and S-shaped ventral

apophysis; femur with small dorsal hump proximally, finger-shaped retrolateral process proximally, and

Figs 36-39. Five specimens. 36-37. Pholcus mulu Huber, sp. nov., c?, Gunung Mulu. 38-39. Pholcus

kawit Huber, sp. nov., $ and 5, Mt. Matutum.

European Journal of Taxonomy 225: 1-34 (2016)

distinct ventral process; tibia large, with small but distinct ventral cavity (for proximal bulbal sclerite);

procursus complex distally, with distinctive prolateral sclerite and membranous structures; bulb with

large rounded uncus with pointed process, weakly sclerotized and distally widening embolus, appendix

with small prolateral spine, distally curved towards retrolateral.

Legs. Without spines or curved hairs; few vertical hairs; retrolateral trichobothrium on tibia 1 at 3.5%;

prolateral trichobothrium absent on tibia 1, present on other tibiae; tarsus 1 with >30 pseudosegments,

only distally about 20 fairly distinct.

Figs 40^4. Pholcus mulu Huber, sp. nov., ZFMK Ar 15501-02. 40^1. Left male palp, prolateral and

retrolateral views. 42. Male chelicerae, frontal view. 43-44 Cleared female genitalia, ventral and dorsal

views. Scale lines: 40M1 = 1 mm; 42M4 = 0.5 mm

HUBER B.A. et al., Philippine hair wax spiders

Male (variation)

Internal dark abdominal marks more distinct in other males. Tibia 1 in 5 other males: 13.9-16.0 (mean

14.9).

Female

In general similar to male but ocular area less elevated, eye triads closer together (distance PME-PME

320 pm), without processes between eye triads. Tibia 1 in 13 females: 12.3-14.0 (mean 13.3). Epigynum

large trapezoidal plate (Figs 27, 43) with anterior ‘knob’; internal genitalia as in Figs 29 and 44, anterior

sclerite medially distinctively curved backwards. ALS with one widened, one pointed, and five very

small conical spigots (~ 1.5-3 pm wide and ~3-6 pm long).

Natural history

Specimens were found in large domed sheet webs (diameter up to ~50 cm) among rocks near the ground.

When disturbed they moved very rapidly towards the periphery of the web.

Distribution

Known from type locality in Sarawak only (Fig. 2).

Pholcus kawit Huber, sp. nov.

um:lsid:zoobank.org:act:92029EAl-F343-406C-8D5F-AD6D9F5820C3

Figs 30-32, 38-39, 45-19

Diagnosis

Easily distinguished from most similar known relatives (species with horns between eye triads carrying

brushes of unglued hairs and with simple, unhinged procursi: P. arayat , P. pagbilao , P. schawalleri ,

P. bagnio sp. nov., P. mulu sp. nov.) by distal modifications of procursus (dorsal pointed process;

Fig. 46); from most species also by female genitalia (epigynum large trapezoidal plate - Fig. 30; similar

in P. pagbilao and P. mulu sp. nov.); from most species (except P. mulu sp. nov.) also by undivided dark

median band ventrally on abdomen (Fig. 39); from P. pagbilao also by simpler appendix (Fig. 45);

from P. arayat also by wider abdomen (Figs 38-39); from P. schawalleri also by more slender uncus

(Fig. 45). From other species of the P. bicornutus group (P. bicornutus and P. olangapo sp. nov.) by

unhinged procursus, presence of appendix, large epigynal plate, and by absence of curved hairs on legs.

Etymology

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

Material examined

Holotype

PHIFIPPINES: $, Mindanao, Mt. Matutum, Kawit Forest, ‘site V (6.338° N, 125.104° E), 950 m a.s.l.,

along brook, among rocks and tree roots, 13 Feb. 2014 (B.A. Huber), ZFMK (Ar 15504).

Other material

PHIFIPPINES: 2 1 5,3 juvs, ZFMK (Ar 15505), same data as holotype; 3 juvs, in absolute

ethanol, ZFMK (Phi 267), same data.

Description

Male (holotype)

Measurements. Total body length 8.0, carapace width 1.8. Feg 1: 52.5 (12.7 + 0.7 + 12.7 + 23.7 + 2.7),

tibia 2: 8.4, tibia 3: 5.3, tibia 4: 7.5; tibia 1 F/d: 78. Distance PME-PME 370 pm, diameter PME 140x170

pm, distance PME-AFE ~50 pm, distance AME-AME 30 pm, diameter AME 90 pm.

European Journal of Taxonomy 225: 1-34 (2016)

Color. Carapace pale ochre to orange, with wide median dark brown mark including ocular area and

small brown submarginal marks laterally (Fig. 38); clypeus light brown; sternum monochromous light

brown to orange, labium darker; legs light brown to ochre, dark rings subdistally on femora and tibiae

and in patella area, tips of femora and tibiae lighter; abdomen ochre-gray, dorsally with several pairs

of dark cuticular marks that are fused above spinnerets, with distinct internal darker marks visible

through cuticle dorsally and laterally; ventrally with undivided wide brown band between gonopore and

spinnerets.

Figs 45^9. Pholcus kawit Huber, sp. nov., ZFMK Ar 15505. 45-46. Left male palp, prolateral and

retrolateral views (arrow points at ventral cavity of tibia). 47. Male chelicerae, frontal view. 48-

49. Cleared female genitalia, ventral and dorsal views. Scale lines: 45M6 = 1 mm; 47M9 = 0.5 mm.

HUBER B.A. et al., Philippine hair wax spiders

Body. Habitus as in Fig. 38; ocular area raised, each eye triad on additional short hump directed towards

lateral, with pair of short processes between eye triads, each with distal brush of hairs; carapace without

median furrow; clypeus unmodified; sternum wider than long (1.05/0.95), unmodified. ALS with one

widened, one pointed, and three to four very small conical spigots (~ 1.5-3 pm wide and ~3-6 pm long).

Chelicerae. As in Fig. 47, with small lateral and frontal apophyses proximally and dark distal apophyses

near median line provided with two small modified (cone-shaped) hairs each; without stridulatory ridges.

Palps. As in Figs 45—46; coxa unmodified; trochanter with short conical retrolateral process and longer

ventral apophysis with distinctive tip; femur with finger-shaped retrolateral process proximally and

distinct ventral apophysis; tibia with dark dorsal band, also ventrally darkened, with small but distinct

ventral cavity (for proximal bulbal sclerite); procursus complex distally, with distinctive pointed process

dorsally (more sclerotized and at different angle than in P. baguio sp. nov.); bulb with large uncus,

weakly sclerotized wide embolus, simple appendix with main branch curved towards retrolateral and

shorter prolateral side-branch.

Fegs. Without spines or 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 >30 pseudosegments,

only distally about 10 fairly distinct.

Male (variation)

Tibia 1 in other male: 13.5 (missing in third male).

Female

In general similar to male but sternum dark brown, eye triads closer together (distance PME-PME

230 pm), without processes between eye triads. Tibiae 1 missing. Epigynum large trapezoidal plate

surrounded by whitish cuticle (Fig. 30); with anterior ‘knob’; internal genitalia as in Figs 32 and 49.

Natural history

Adult specimens were only found in large holes at tree bases near a brook in the forest. Juveniles were

more co mm on in any dark sheltered spaces along the brook.

Distribution

Known from type locality in Mindanao only (Fig. 2).

Pholcus baguio Huber, sp. nov.

urn:lsid:zoobank.org:act:753EAC47-AFE0-4704-A564-696C4DD0A14F

Figs 33-35, 50-52, 58-76

Diagnosis

Distinguished from most similar known relatives (species with horns between eye triads carrying brushes

of unglued hairs and with simple, unhinged procursi: P. arayat , P. pagbilao , P. schawalleri , P. kawit

sp. nov., P. mulu sp. nov.) by distal modifications of procursus (dorsal pointed process; large prolatero-

ventral process; Fig. 59), by relatively wide and short palpal femur (Fig. 59), and by female genitalia

(epigynum with pair of dark lateral areas; internal genitalia with large transversal sclerite; Figs 33-35);

from P. pagbilao also by simpler appendix (Figs 58, 71-72); from P. arayat also by relatively wider

abdomen (Figs 50-52); from P. kawit sp. nov. and P. mulu sp. nov. also by divided median band ventrally

on abdomen (Fig. 51); from P. schawalleri also by much more slender uncus (Fig. 58). From other

species of the P. bicornutus group (P. bicornutus ; P. olangapo sp. nov.) by unhinged procursus, presence

of appendix, large epigynal plate, and by absence of curved hairs on legs.

European Journal of Taxonomy 225: 1-34 (2016)

Figs 50-53. Live specimens. 50-52. Pholcus baguio Huber, sp. nov., §, and $ with egg-sac and

emerging juveniles, Mt. Kabuyao. 53. Pholcus arayat Huber, 2011, S, Pamulaklakin Forest Trail.

HUBER B.A. et al., Philippine hair wax spiders

Figs 54-57. Live specimens. 54. Pholcus arayat Huber, 2011, $, Pamulaklakin Forest Trail. 55-57.

Pholcuspagbilao Huber, 2011, S from Loboc (55), $ from Mt. Banahaw (56), and $ with egg-sac from

Loboc (57).

European Journal of Taxonomy 225: 1-34 (2016)

Etymology

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

Material examined

Holotype

PHILIPPINES: Luzon, Benguet Prov., near Baguio, Mt. Kabuyao, N slope (16.374° N, 120.557° E),

1200-1400 m a.s.l., among rocks, 2 Mar. 2014 (B.A. Huber), ZFMK (Ar 15506).

Figs 58-62. Pholcus baguio Huber, sp. nov., ZFMK Ar 15507-08. 58-59. Left male palp, prolateral and

retrolateral views. 60. Male chelicerae, frontal view. 61-62. Cleared female genitalia, ventral and dorsal

views. Scale lines: 58-59 = 1 mm; 60-62 = 0.5 mm.

HUBER B.A. et al., Philippine hair wax spiders

Figs 63-70. Pholcus baguio Huber, sp. nov., ZFMK Ar 15507. 63. Male prosoma, frontal view. 64. Male

ocular area, frontal (slightly dorsal) view. 65-66. Male ocular horns and modified hairs. 67. Male ocular

area, dorsal view. 68. Male left palpal tarsal organ. 69. Right male palpal trochanter and procursus,

retrolatero-dorsal view. 70. Left procursus, distal view. Scale lines: 63 = 500 pm; 64 = 300 pm; 65 =

80 pm; 66, 70 = 100 pm; 67, 69 = 200 pm; 68 = 30 pm.

European Journal of Taxonomy 225: 1-34 (2016)

Other material

PHILIPPINES: 17 S3, 10 55, 1 juv., ZFMK (Ar 15507-08) and 1 3, 1 5, MSU-IIT, same data as

holotype, among rocks and vegetation; 1 §, 3 juvs, in absolute ethanol, same data, ZFMK (Phi 204);

7 33 , 16 55 , Benguet, “Tuba, Asin road, 10 km S of Baguio” [-16.37° N, 120.62° E], rock wall at

waterfall, 12 Nov. 1979 (P. Lehtinen), ZMT.

Description

Male (holotype)

Measurements. Total body length 7.7, carapace width 1.8. Leg 1: 51.2 (12.5 + 0.8 + 12.4 + 22.8 + 2.7),

tibia 2: 8.3, tibia 3: 5.3, tibia 4: 7.5; tibia 1 L/d: 67. Distance PME-PME 380 pm, diameter PME

140x160 pm, distance PME-ALE -40 pm, distance AME-AME 30 pm, diameter AME 100 pm.

Color. Carapace ochre to orange, with wide median brown mark including ocular area and clypeus (except

rim), connecting posteriorly with lateral submarginal brown bands (Fig. 50); sternum monochromous

light brown to orange, labium darker; legs light brown, dark rings subdistally on femora and tibiae and

in patella area, tips of femora and tibiae lighter; abdomen ochre-gray, dorsally with several pairs of dark

cuticular marks that are fused above spinnerets, laterally with indistinct internal darker marks visible

through cuticle; ventrally with wide brown band divided into anterior and posterior parts separated by

light V-shaped space.

Body. Habitus as in Fig. 50; ocular area raised, each eye triad on additional short hump directed towards

lateral (Fig. 63), with pair of long processes between eye triads directed towards posterior, each with

distal brush of hairs curved upwards, hairs not ‘glued" together (Figs 64-67); carapace without median

furrow (Fig. 64); clypeus unmodified; sternum wider than long (1.10/0.85), unmodified. ALS with one

widened, one pointed, and three very small conical spigots (Fig. 74). Gonopore with four epiandrous

spigots.

Chelicerae. As in Fig. 60, small lateral and frontal apophyses proximally and dark distal apophyses near

median line provided with two small modified (cone-shaped) hairs each (Fig. 73); without stridulatory

ridges.

Palps. As in Figs 58-59; coxa unmodified; trochanter with short conical retrolateral process and longer

ventral apophysis with distinctive tip (Fig. 69); femur with distinct retrolateral process proximally,

small dorsal hump, and distinct darkened ventral apophysis; tibia with dark dorsal band, also ventrally

darkened, with very small and indistinct ventral cavity; tarsal organ capsulate (Fig. 68); procursus with

distinctive pointed process dorsally and large process prolatero-ventrally (Figs 69-70); bulb with large

uncus (Figs 58, 71), weakly sclerotized wide embolus, simple appendix with long main branch curved

towards prolateral and shorter retrolateral side-branch (Figs 71-72).

Legs. Without spines or 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 about 15 fairly distinct.

Male (variation)

Most males with distinct internal dark marks dorsally and laterally on abdomen. Tibia 1 in 19 other

males: 10.9-14.3 (mean 12.3).

Female

In general similar to male but sternum dark brown, eye triads closer together (distance PME-PME 250

pm), without processes between eye triads. Tibia 1 in 16 females: 9.6-11.6 (mean 10.7). Epigynum

HUBER B.A. et al ., Philippine hair wax spiders

Figs 71-76. Pholcus baguio Huber, sp. nov., ZFMK Ar 15507-08. 71-72. Left genital bulb, prolateral

and prolatero-distal views. 73. Male distal cheliceral apophysis (arrows point at modified hairs). 74. Male

ALS. 75. Epigynum. 76. Female ALS and PMS. Scale lines: 71-72, 75 = 200 pm; 73-74, 76 = 20 pm.

European Journal of Taxonomy 225: 1-34 (2016)

relatively small, roughly oval plate (Figs 33, 75) with pair of dark brown areas and anterior ‘knob’

wealdy sclerotized in most specimens; entire epigynal area strongly protruding in some specimens;

internal genitalia as in Figs 35 and 62. Spinnerets and spigots as in male (Fig. 76).

Natural history

The spiders were found in partly extremely high densities in a dry brook bed near the road, among rocks

and shaded vegetation. The individual domed sheets were interconnected to form large communal webs

of up to 2 m length.

Distribution

Known from Baguio area in Tuzon only (Fig. 2).

Pholcus arayat Huber, 2011

Figs 53-54, 77-90

Pholcus arayat Huber, 2011a: 318-320, figs 1514-1515, 1541-1542, 1574-1578.

Diagnosis (updated)

Distinguished from most similar known relatives (species with horns between eye triads carrying brushes

of unglued hairs and with simple, unhinged procursi: P. pagbilao , P. schawalleri , P. baguio sp. nov.,

P. kawit sp. nov., P. mulu sp. nov.) by bifid process ventro-distally on procursus (arrows in Figs 82-84;

cf. fig. 1575 in Huber 2011a) and by shapes of uncus and appendix (cf. fig. 1574 in Huber 2011a); from

P. pagbilao also by more slender abdomen and shorter epigynum (cf. fig. 1541 in Huber 2011a); from

P. baguio sp. nov. also by more slender male palpal femur (cf. fig. 1575 in Huber 2011a) and different

shape of epigynum (not with two distinct dark lateral areas); from P. kawit sp. nov. and P. mulu sp.

nov. also by divided median band ventrally on abdomen. From other species of the P. bicornutus group

(P. bicornutus ; P. olangapo sp. nov.) by unhinged procursus, presence of appendix, large epigynal plate,

and by absence of curved hairs on legs.

Material examined

New material

PHITIPPINES, Luzon: 14 SS, 12 $ $, ZFMK (Ar 15509) and 1 S, 1 $, MSU-IIT, Bataan Prov., near

Olangapo, along river near Pamulaklakin Forest Trail (14.798° N, 121.338° E), 80 m a.s.l., among

rocks and vegetation, 28 Feb. 2014 (B.A. Huber); 2 §§, 1 juv., in absolute ethanol, same data, ZFMK

(Phi 208); 2 Laguna Prov., Los Banos, Makiling Forest (14.145° N, 121.230° E), 20 Mar. 2014

(S. Huber), ZFMK (Ar 15510).

Assigned tentatively

PHILIPPINES, Bohol Isl.: 3 $$, Rajah Sikatuna (Magsaysay Park) (9.705° N, 124.123° E), 430 m

a.s.l., forest, 6 Mar. 2014 (B.A. Huber), ZFMK (Ar 15511); 1 <S, Valencia, Barangay Marawis, ‘site 3’

(9.723° N, 124.202° E), 440 m a.s.l., 11 Jun. 2015 (M.R.B. Dacar), ZFMK (Ar 15512).

Note

The single male specimen from “Baybay, VISCA complex”, misidentified as “ Pholcus phalangioides ”

by Barrion & Litsinger (1995) was tentatively assigned to Pholcus arayat in Huber (2011a). Recently,

Yao et al. (2014) described Pholcus schawalleri from “Visca, N. of Baybay” (which should be ViSCA,

Visayas State College of Agriculture, now Visayas State University, approx. 8 km N of Baybay,

~10°45' N, 124°49' E - not 10°39' N, 124°51' E as in Yao et al. 2014). These authors did not mention

Barrion & Litsinger’s (1995) record, but the identical locality and the shape of the uncus (large and

HUBER B.A. et al., Philippine hair wax spiders

Figs 77-84. Pholcus arayat Huber, 2011, ZFMK Ar 15509. 77. Male prosoma, frontal view. 78. Male

ocular horns and modified hairs. 79. Female prosoma, frontal view. 80. Male ocular area, dorsal view.

81. Right genital bulb, prolateral view. 82. Feft male palpal trochanter and procursus, dorsal view (arrow

points at bifid process). 83-84. Tip of right procursus, prolateral and prolatero-distal views (arrows point

at bifid process). Scale lines: 77, 79 = 300 pm; 78 = 50 pm; 80-82 = 200 pm; 83-84 = 100 pm.

European Journal of Taxonomy 225: 1-34 (2016)

roundish) strongly suggest that Barrion & Litsinger’s dubious specimen is P. schawalleri rather than

P. arayat.

Description (amendments)

As noted in Huber (2011a), the holotype is apparently artificially darkened. The new material supports

this assumption: carapace in males and females light brown with darker median band (sometimes

indistinct; in females medially divided) and lateral bands (sometimes barely visible); sternum light brown

to orange in both males and females. Hairs on male ocular processes not ‘glued’ together (Figs 77-78,

80). Tip of procursus extremely complex (Figs 83-84). Male palpal tibia with small but distinct ventral

Figs 85-90. Pholcus arayat Huber, 2011, ZFMK Ar 15509. 85. Male distal cheliceral apophysis (arrows

point at modified hairs). 86. Male gonopore. 87. Male tarsus 4 comb-hairs. 88. Male ALS. 89. Epigynum.

90. Female ALS. Scale lines: 85, 87-88, 90 = 20 pm; 86 = 50 pm; 89 = 200 pm.

HUBER B.A. et al., Philippine hair wax spiders

cavity. Distal male cheliceral apophyses with two modified (cone-shaped) hairs each (Fig. 85). Male

gonopore with four epiandrous spigots (Fig. 86). Tarsus 4 comb-hairs as in Fig. 87. AFS in both sexes

with one widened, one pointed, and three very small conical spigots (Figs 88, 90).

Variation

Tibia 1 in 15 newly examined males: 10.1-13.5 (mean 11.9); in 12 newly examined females: 9.2-11.1

(mean: 10.3). The specimens from Bohol are overall very similar to those from Fuzon but the male

differs in some details of the palp: tip of trochanter apophysis not so strongly directed proximad; distal

semitransparent process on procursus not clearly bifid; distance between tip of uncus and subterminal

process larger. They are thus assigned tentatively.

Natural history

The spiders were found both in sheltered spaces (among rocks and logs) and among vegetation, with

their domed webs sometimes directly exposed to the sun.

Distribution

Known from several localities in central Fuzon (Fig. 2); specimens from Bohol are assigned tentatively

(see above).

Pholcus pagbilao Huber, 2011

Figs 55-57, 91-102

Pholcus pagbilao Huber, 2011a: 320, figs 1516-1521, 1543-1544, 1579-1584.

Diagnosis (updated)

Distinguished from most similar known relatives (species with horns between eye triads carrying

brushes of unglued hairs and with simple, unhinged procursi: P. arayat, P. schawalleri , P. baguio

sp. nov., P. kawit sp. nov., P. mulu sp. nov.) by slender semitransparent process distally on procursus

(cf. fig. 1580 in Huber 2011a; arrow in Fig. 97) and by shapes of uncus and complex appendix (cf. fig.

1579 in Huber 2011a; see also Fig. 98); from P. arayat also by wider abdomen and longer, trapezoidal

epigynum (Fig. 101; see also fig. 1543 in Huber 2011a); from P. baguio sp. nov. also by more slender

male palpal femur (cf. fig. 1580 in Huber 2011a); from P. kawit sp. nov. and P. mulu sp. nov. also by

divided dark median band ventrally on abdomen (Fig. 57). From other species of the P. bicornutus group

(P. bicornutus , P. olangapo sp. nov.) by unhinged procursus, presence of appendix, large epigynal plate,

and by absence of curved hairs on legs.

Material examined

New material

PHIFIPPINES, Fuzon: 3 3 55, 1 juv., Quezon Prov., between Fucban and Tayabas (14.063° N,

121.567° E), 330 m a.s.l., degraded forest along river, among rocks, 26 Feb. 2014 (B.A. Huber), ZFMK

(Ar 15513); 1 S, in absolute ethanol, same data, ZFMK (Phi 212); 6 6 55> 1 juv., Faguna Prov.,

Mt. Banahaw, forest near Taytay Falls (14.110° N, 121.507° E), 560 m a.s.l., among rocks, 26 Feb. 2014

(B.A. Huber), ZFMK (Ar 15514); 3 juvs, in absolute ethanol, same data, ZFMK (Phi 218); 17

6 5 5> ZFMK (Ar 15515-16) and 1 $, 1 5, MSU-IIT, Camarines Sur Prov., Mt. Isarog, forest and (most

specimens) abandoned buildings near park entrance (13.663° N, 123.335° E), 500 m a.s.l., 23 Feb. 2014

(B.A. Huber); 2 $$, 1 5, 3 juvs, in absolute ethanol, same data, ZFMK (Phi 220).

PHIFIPPINES, Bohol Isl.: 1 & 2 5 5 , Rajah Sikatuna (Magsaysay Park) (9.705° N, 124.123° E), 430 m

a.s.l., forest, 6 Mar. 2014 (B.A. Huber), ZFMK (Ar 15517); 1 5, 1 juv., in absolute ethanol, same data,

European Journal of Taxonomy 225: 1-34 (2016)

Figs 91-98. Pholcus pagbilao Huber, 2011, ZFMK Ar 15515-16. 91. Male prosoma, frontal view.

92. Male ocular horns and modified hairs. 93. Female prosoma, frontal view. 94-95. Male ocular area,

dorsal and latero-dorsal views. 96. Male distal cheliceral apophysis (arrows point at modified hairs).

97. Left procursus, dorsal view (arrow points at semitransparent process). 98. Left genital bulb, prolateral

view. Scale lines: 91, 93 = 400 pm; 92 = 60 pm; 94-95, 98 = 200 pm; 96 = 20 pm; 97 = 100 pm.

HUBER B.A. et al., Philippine hair wax spiders

ZFMK(Phi 196); 13 8 $ nearLoboc, above Loboc River (-9.655° N, 124.015° E), ~250 m a.s.l.,

in caves near entrances, 5 Mar. 2014 (B.A. Huber), ZFMK (Ar 15518); 5 $$, 6 5¥ ? near Loboc, at

Loboc River (9.651° N, 124.022° E), at rocks, 20 m a.s.l., 4 Mar. 2014 (B.A. Huber), ZFMK (Ar 15519);

2 55,1 juv., in absolute ethanol, same data, ZFMK (Phi 203).

PHILIPPINES, Cebu Isl.: 1 1 5= Moalboal, Busay Cave (9.916° N, 123.437° E), cave entrance,

4 Apr. 2014 (S. Huber), ZFMK (Ar 15520).

Assigned tentatively

PHILIPPINES, Negros Isl.: 6 $$, 7 5 5 , Negros Oriental Prov., Casaroro Falls (9.281° N, 123.208° E),

550 m a.s.l., at rocks along river below waterfall, 10 Mar. 2014 (B.A. Huber), ZFMK (Ar 15521); 2 5 5>

1 juv., in absolute ethanol, same data, ZFMK (Phi 186).

PHILIPPINES, Bohol Isl.: 2 55 , 14 juvs, MSU-IIT and 1 5 , ZFMK (Ar 15522), Bilar, Barangay

Riverside (9.70° N, 124.12° E), 300M50 m a.s.l., 15 Jun. 2015 (M.R.B. Dacar); 15,3 juvs, Garcia

Hernandez, Barangay Datag (9.72° N, 124.26° E), 530 m a.s.l., 8 Jun. 2015 (M.R.B. Dacar), ZFMK

(Ar 15523); 2 5 5 , Sierra Bullones, Barangay Nan-od (9.746° N, 124.262° E), 610 m a.s.l., 5 Jun. 2015

Figs 99-102. Pholcuspagbilao Huber, 2011, ZFMK Ar 15515-16. 99. Male gonopore. 100. Male ALS.

101. Epigynum. 102. Female ALS. Scale lines: 99 = 50 pm; 100, 102 = 20 pm; 101 = 200 pm.

European Journal of Taxonomy 225: 1-34 (2016)

(M.R.B. Dacar), ZFMK (Ar 15524); 2 12 juvs, MSU-IIT and 1 ?, ZFMK (Ar 15525), Valencia,

Barangay Marawis (9.724° N, 124.201° E), 450 m a.s.l., 11 Jun. 2015 (M.R.B. Dacar).

Description (amendments)

Hairs on male ocular processes not ‘glued’ together (Figs 91-92, 94-95). Distal male cheliceral

apophyses with three modified (cone-shaped) hairs each (Fig. 96). Male palpal tibia with small and

indistinct ventral cavity. Male gonopore with four epiandrous spigots (Fig. 99). ALS in both sexes with

one widened, one pointed, and six cylindrically-shaped spigots of varying sizes (Figs 100, 102).

Variation

As noted in Huber (2011a), the tip of the procursus differs slightly between males from Negros Island

and males from Luzon. Interestingly, males from Bohol Island resemble those from Luzon much more

closely than those from neighboring Negros. In addition, there is variation among males from Negros

Island: males from Mabinay (see Huber 2011a) differ slightly from the newly collected males from

Casaroro Falls. However, all these specimens share the distinctive slender semitransparent process

distally on the procursus and the shapes of uncus and appendix (cf. figs 1579-1580 in Huber 2011a).

Tibia 1 in 46 newly examined males: 9.6-15.3 (mean 12.7) (42 of these males: 11.2-14.0); in 38 newly

examined females: 9.3-12.8 (mean: 11.2). The black ventral band of the abdomen is always clearly

divided into anterior and posterior parts but in some specimens (of both sexes) the tip of the posterior

part has a narrow connection to the anterior part. Some of the newly collected specimens from Bohol

are assigned tentatively because they are not accompanied by males and because the ventral abdominal

pattern is partly intermediate between that of P. pagbilao and P. arayat. Some of these specimens may

actually be conspecific with the specimens assigned tentatively to P. arayat.

Natural history

The spiders were found in sheltered spaces close to the ground, usually among rocks. In the abandoned

buildings near the park entrance of Mt. Isarog, the spiders were found in high numbers in all corners at

any height from the ground.

Distribution

Apparently widely distributed in the Philippines (Fig. 2).

Discussion

The male ocular horns with brushes of hairs described herein represent a unique synapomorphy of the

Pholcus bicornutus group (Huber 2011a), but ocular area modifications in general are widespread in

Pholcidae and come in a wide range of shapes (Table 1). Two aspects are particularly remarkable about

the overview in Table 1. First, with only one exception (the ‘pseudo-lenses’ accompanying the secondary

eyes in Smeringopus Simon, 1890 and Smeringopina Kraus, 1957) all modifications are restricted to the

male, i.e., represent sexual dimorphisms. Second, these modifications appear very unevenly distributed

among major taxonomic groups. No case is known in Ninetinae; only six cases are known in Arteminae,

Modisiminae, and Smeringopinae together. All other 14 cases are in Pholcinae.

Each of the twenty cases listed in Table 1 is supposed to represent an independent origin. This is derived

from available phylogenies (reviewed in Huber 2011b; see also Huber 2013; Dimitrov et al. 2013;

Huber & Nuneza 2015; Huber et al. 2015) and from partly fundamental differences in the quality of

the modifications. We admit that some particular modifications may have originated more than once

(e.g., the small median elevations in Pholcus youngae Huber, 2011 and Pholcus schwendingeri Huber,

2011; or the pair of pointed processes near the PME in the Pholcus minang group and in Panjange

Deeleman-Reinhold & Deeleman, 1983), w hil e others may not represent independent origins (e.g., the

HUBER B.A. et al, Philippine hair wax spiders

Table 1. Ocular area modifications in Pholcidae.

Taxon

Ocular area modification

References

Ninetinae

—

Arteminae

Trichocyclus septentrionalis

Male AME on median elevation

Huber 2001

Modisiminae

Modisimus (part)

Male eye turret higher than female turret

Huber et al. 2010

Modisimus culicinus

Male eye turret with hairy cuticular lobe and

glands

Huber 1997

Smeringopinae

Smeringopina & Smeringopus

Secondary eyes with ‘pseudo-lenses’ (males and

females)

Huber 2009, 2012

Smeringopina cornigera group

Males with pair of large pointed horns

Huber 2013

Smeringopina cornigera group, part

Males with additional pair of smaller horns

Huber 2013

Pholcinae

Male eye triads on short to long stalks; extremes

in, e.g.,: Panjange casaroro, Panjange camiguin ,

Pholcus schwendingeri, Pholcus minang group

Huber 2011a;

Huber & Nuneza 2015

Aetana (A. lozadae, A. manansalai,

A. banahaw )

Males with pair of processes near ALE

Huber et al. 2015

Aetana (A. loboc, A. pasambai)

Males with median process/hump

Huber et al. 2015

Hantn kapit

Males with two pairs of long horns and one pair

of small processes

Huber 2016

Pehrforsskalia

Males with pair of small cone-shaped processes

near PME

Huber 2009, 2011a

Pholcns group of genera (part)

Males with higher density of hairs

Huber 2011a

Pholcus ancoralis group (part)

Males with pair of pointed horns (without hairs)

between triads

Huber 2011a

Pholcus bicornutus group

Males with pair of horns with brushes of hairs

between/behind triads

Huber 2011a; this

study

Pholcns calligaster group (part)

Males with pair of sculptured horns (without

hairs) between triads

Huber 2011a

Pholcns halabala group

Males with pair of humps near PME with curved

spines

Huber 2011a;

Huber et al. 2016

Pholcns spp. ( minang group,

ethagala group) and Panjange

(part)

Males with pair of pointed processes near PME;

sometimes forming apparent functional unit

(Panjange casaroro , Panjange camiguin )

Huber 2011a; Huber

& Nuneza 2015;

Huber & Leh Moi

Ung 2016

Pholcns quinquenotatus

Males with median hooked process above AME

Huber 2011a

Pholcus yoimgae,

Pholcus schwendingeri

Males with small median elevation

Huber 2011a

Spermophora , some East African

species ( S. bukusu , S. man,

S. berlandi , S. maathaiae)

Males with pair of rounded anterior horns

Huber & Warui 2012

European Journal of Taxonomy 225: 1-34 (2016)

pointed horns in the Pholcus ancoralis group and the sculptured horns in the Pholcus calligaster group).

An exact count will require phylogenies with denser taxon sampling and better resolution (which is

currently in preparation; J. Eberle, A. Valdez-Mondragon, D. Dimitrov & B. A. Huber, unpubl. data), but

it seems very unlikely that the number of independent origins will change dramatically.

Unfortunately, we know much less about the function of pholcid ocular modifications. In fact, there

is only one study (Huber 1997) that deals in some detail with a sexually dimorphic ocular region.

In Modisimus culicinus (Simon, 1893), the male ocular turret is provided anteriorly with a cuticular

lobe that is densely set with hairs and with pores through which large glands in the male ocular area

discharge their products. During copulation, the female mouthparts contact this cuticular lobe, suggesting

gustatorial courtship (Huber 1997). At first sight it would seem that such a gustatorial courtship function

might be a valid hypothesis for further cases: first, the copulatory position of Pholcidae consistently

brings the male ocular area close to the female mouthparts; and secondly, dense sets of hairs occur on the

male ocular areas in many other Pholcidae (actually in most Pholcus species; Huber 2011a). However,

glandular pores have never been found on the male ocular areas in any other species, even though many

were examined in detail with the scanning electron microscope. This leaves a wide range of possible

explanations, including mechanical functions in an intersexual selection context as well as intrasexual

selection (male-male contest) scenarios. Obviously, this is a rich field for future observational and

experimental studies.

Acknowledgements

We thank Seppo Koponen, Karen van Dorp, and Charles Griswold for the loan of specimens; Maria

Dacar for contributing specimens from her own research; Samuel Huber for accompanying us in the

field; Siegfried Huber for donating specimens; our Philippine driver Vicente Bulongon for bringing

us safely to all requested places; the Sarawak Museum Department for assistance in field logistics; the

Sarawak Forest Department (Permit No. NCCD.907.4.4 JLD. 10-211, ParkPermitNo. 174/2014) and the

Philippine Department of Environment and Natural Resources (DENR) CARAGA, DENR Region X,

and DENR Region XI for issuing the relevant permits; two anonymous reviewers for useful comments;

and the German Research Foundation for financial support (DFG Project HU 980/11-1).

References

Barrion A.T. & Litsinger J.A. 1995. Riceland Spiders of South and Southeast Asia. CAB International,

Wallingford, U.K.

Brown B.V. 1993. A further chemical alternative to critical-point-drying for preparing small (or large)

flies. Fly Times 11: 10.

Dimitrov D., Astrin J.J. & Huber B.A. 2013. Pholcid spider molecular systematics revisited, with new

insights into the biogeography and the evolution of the group. Cladistics 29: 132-146. http://dx.doi.

org/10,1111/i, 1096-0031.2012,00419.x

Huber B.A. 1997. Evidence for gustatorial courtship in a haplogyne spider (. Hedypsilus

culicinus : Pholcidae: Araneae). Netherlands Journal of Zoology 47 (1): 95-98. http://dx.doi.

org/10,1163/156854297X00265

Huber B.A. 2001. The pholcids of Australia (Araneae; Pholcidae): taxonomy, biogeography, and

relationships. Bulletin of the American Museum of Natural History 260: 1-144. http://dx.doi.

org/10.1206/0003-0090(200 D260%3C0001:TPOAAP%3E2.0.CQ:2

Huber B.A. 2009. Life on leaves: leaf-dwelling pholcids of Guinea, with emphasis on Crossopriza

cylindrogaster Simon, a spider with inverted resting position, pseudo-eyes, lampshade web, and

HUBER B.A. et al., Philippine hair wax spiders

tetrahedral egg-sac (Araneae: Pholcidae). Journal of Natural History 43: 2491-2523. http://dx.doi.

org/10.1080/00222930903207876

Huber B.A. 2011a. Revision and cladistic analysis of Pholcus and closely related taxa (Araneae,

Pholcidae). Bonner zoologische Monographien 58: 1-509.

Huber B.A. 2011b. Phylogeny and classification of Pholcidae (Araneae): an update. Journal of

Arachnology 39: 211-222.

Huber B.A. 2012. Revision and cladistic analysis of the Alfotropical endemic genus Smeringopus

Simon, 1890 (Araneae: Pholcidae). Zootaxa 3461: 1-138.

Huber B.A. 2013. Revision and cladistic analysis of the Guineo-Congolian spider genus Smeringopina

Kraus (Araneae, Pholcidae). Zootaxa 3713: 1-160. http://dx.doi.org/10.11646/zootaxa.3713.1.1

Huber B.A. 2016. A new genus of ground and litter-dwelling pholcine spiders from Sarawak (Araneae,

Pholcidae). European Journal of Taxonomy 186: 1-15. http://dx.doi.org/10.5852/ejt.2016.186

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

high diversity in Sarawak, apparent absence in Sabah. European Journal of Taxonomy 184: 1-32. http://

dx.doi.org/10.5852/ejt.2016.184

Huber B.A. & Nuneza O.M. 2015. Evolution of genital asymmetry, exaggerated eye stalks, and extreme

palpal elongation in Panjange spiders (Araneae: Pholcidae). European Journal of Taxonomy 169: 1—46.

http://dx.doi.org/10.5852/ejt.2015.169

Huber B.A. & Warm C.M. 2012. East African pholcid spiders: an overview, with descriptions of eight

new species (Araneae, Pholcidae). European Journal of Taxonomy 29: 1-44. http://dx.doi.org/10.5852/

eit.2012.29

Huber B.A., Fischer N. & Astrin J.J. 2010. High level of endemism in Haiti’s last remaining forests: a

revision of Modisimus (Araneae: Pholcidae) on Hispaniola, using morphology and molecules. Zoological

Journal of the Linnean Society 158: 244-299. http://dx.doi.org/10. Ill 1/i. 1096-3642.2009.00559.x

Huber B.A., Nuneza O.M. & Leh Moi Ung C. 2015. Revision, phylogeny, and microhabitat shifts in the

Southeast Asian spider genus Aetana (Araneae, Pholcidae). European Journal of Taxonomy 162: 1-78.

http://dx.doi.org/10.5852/eit.2015.162

Huber B.A., Petcharad B., Leh Moi Ung C., Koh J.K.H. & Ghazali A.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

Simon E. 1892. Etude sur les arthropodes cavernicoles de Pile de Luzon. Annales de la Societe

entomologique de France 61: 35-52.

Simon E. 1893. Histoire Naturelle des Araignees. 2nd edition, 1 (2): 256-488. Librairie Encyclopedique

de Roret, Paris, http://dx.doi.org/10.5962/bhl.title.51973

Walckenaer C.A. 1805. Tableau des araneides ou caracteres essentiels des tribus, genres, families et

races que renferme le genre Aranea de Linne, avec la designation des especes comprises dans chacune

de ces divisions. E. Dentu, Paris.

Yao Z., Li S. & Jager P. 2014. Four new species of pholcine spiders (Araneae: Pholcidae) from Southeast

Asia. Zootaxa 3793: 331-349. http://dx.doi.Org/10.11646/zootaxa.3793.3.2

Manuscript received: 20 January 2016

Manuscript accepted: 24 March 2016