European Journal of Taxonomy 186: 1-15

http://dx.doi.org/10.5852/ejt.2016.186

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ISSN 2118-9773

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2016 • Huber B.A.

Research article

urn:lsid:zoobank.org:pub:B73C6A06-1245-41CB-9175-C33BC2273245

A new genus of ground and litter-dwelling pholcine spiders

from Sarawak (Araneae, Pholcidae)

Bernhard A. HUBER

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

Email: b.huber@zfmk de

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

Abstract. Two small, ground and litter-dwelling pholcid species from northern Borneo are described

as representatives of a new genus, Hantu gen. nov.: H. kapit gen. et sp. nov. and H. niah gen. et sp. nov.

Previous cladistic analyses suggested a closer relationship with the genera Savarna Huber, 2005 and

Khorata Huber, 2005 (mainland Southeast Asia) than with the geographically closer genus Aetana Huber,

2005 (Borneo and Philippines to Fiji). Since the two species do not share any of the synapomorphies of

Khorata and Savarna while having several synapomorphies on their own (ventral apophysis on male

palpal coxa; male palpal trochanter apophysis with small teeth or scales; spines on male femora 1; high

density of vertical hairs on male femora; presence of scape on epigynum), they are here proposed as

representing a new genus.

Keywords. Borneo, Sarawak, endemism, taxonomy, Pholcidae.

Huber B.A. 2016. Anew 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/eit.2Q16.186

Introduction

The two species newly described herein were initially to be included in a revision of the Southeast

Asian pholcid genus Aetana Huber, 2005 (Huber et al. 2015a). Superficially, they resembled ground

and litter-dwelling representatives of that genus, and their geographic origin also pointed towards

Aetana (ranging from Borneo and the Philippines to Fiji) rather than to other similar genera on the

Southeast Asian mainland ( Savarna Huber, 2005; Khorata Huber, 2005). However, closer examination

and cladistic analyses revealed that these two species (under the names “Gen.n. Bor80” and “Gen.n.

Bor20” in Huber et al. 2015a) were not only different from Aetana in many details, but, in fact, closer

to Khorata and Savarna. At the same time, the two species do not share any of the distinctive characters

of Khorata (Huber 2005b) and Savarna (Huber 2005b; Huber et al. 2015b). As a consequence, they are

here described as representatives of a new genus.

Material and methods

The material studied herein was collected during a recent expedition to northern Borneo (Jul.-Aug.

2014). It is currently deposited at the Zoologisches Forschungsmuseum Alexander Koenig, Bonn

(ZFMK), and the Sarawak Museum, Kuching (SMK).

European Journal of Taxonomy 186 : 1-15 ( 2016 )

Methods and terminology are as in recent revisions (Huber 2011, 2013). Measurements are in mm

unless otherwise noted. Eye measurements are ± 5 jam. Epigyna were cleared in warm NaOH solution

and stained with chlorazol black. For SEM photos, specimens were dried in 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. The distribution map was 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

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

Results

Class Arachnida Cuvier, 1812

Order Araneae Clerck, 1757

Family Pholcidae C.L. Koch, 1851

Hantu gen. nov.

urn:lsid:zoobank.org:act:9DBCF7B6-8A03-4346-BFA7-9DE99CD065D7

Type species

Hantu kapit gen. et sp. nov.

Diagnosis

Small, six-eyed, dark, ground and litter-dwelling pholcids with dark (usually black) sternum, with

thoracic furrow extending over entire length of carapace, and without epiandrous spigots. Distinguished

from putatively closest known relatives ( Khorata , Savarna ) by ventral apophysis on male palpal coxa

(arrows in Figs 9, 19, 27), by scales or teeth on male palpal trochanter (Fig. 16), by spines on male

femora 1 (—10-25 in single ventral row), by short vertical hairs in high density on male femora (in two

dorsal rows), and by scape on female external genitalia (Figs 40M5); from Savarna also by presence

of distal cheliceral apophyses (Figs 11, 28) and by male palpal trochanter apophysis not fused to femur

(Figs 9, 27); from Khorata also by absence of sclerotized ledges laterally on male chelicerae (Figs 11,

28) and by absence of retrolateral process on male palpal femur (Figs 9, 27).

Etymology

Named for the Hantu Rimba, deep-forest ghosts in traditional Malaysian mythology. Gender masculine.

Description

Male

Measurements. Total body length -2.0-2.5; carapace width 0.9-1.0; leg 1 length -20-25; tibia 1

length -4.5-6.0; tibia 2/tibia 4 length 1.0-1.1; tibia 1 L/d -55-75.

Color. In life (Figs 1-6) mostly ochre-gray with black marks, sternum black, legs without dark rings;

abdomen with dark brown to black marks dorsally, laterally, and ventrally.

Body. Carapace with distinct median furrow (Figs 14,31,34); ocular area raised, eye triads on short stalks

directed toward lateral (Figs 14, 31), with complex modification in H. kapit gen. et sp. nov. (Figs 10,

HUBER B.A., New spider genus from Sarawak

14, 15, 17). AME absent. Clypeus high, either unmodified or with small median process (Figs 15, 17).

Abdomen slightly longer than high, pointed at spinnerets (Figs 1-6). Male gonopore without epiandrous

spigots (Figs 23, 32). AES with large widened spigot and pointed spigot, without further cylindrically

shaped spigots (Figs 24, 38); PMS with two spigots each.

Chelicerae. With two pairs of processes, one proximal lateral, one distal frontal (Figs 11, 28); without

modified hairs; without stridulatory ridges.

Palps. Coxa with distinct ventral process (Figs 8, 19, 27); trochanter with retrolateral process of varying

length provided with small scales or teeth (Fig. 16); femur large, without processes; patella relatively

long ventrally (not triangular in lateral view; Figs 9, 27); tibia rather small and slender, with retrolateral

trichobothrium in ‘normal’ (not very distal) position (at -60% of tibia length); palpal tarsus small, with

pit-shaped rather than capsular tarsal organ (Figs 21, 33); procursus complex, proximal and distal parts

connected by membranous hinge; bulb with weakly sclerotized embolus as only process (Figs 9, 27).

Fegs. With spines on femora 1; without curved hairs; vertical hairs in two dorsal rows on all femora and

tibiae; retrolateral trichobothrium on tibia 1 at -6-8% of tibia length; prolateral trichobothrium absent

on tibia 1, present on other tibiae. Tarsal organ not examined. Tarsus 1 with -15-20 pseudosegments

fairly distinct distally; tarsus 4 with single row of ventral comb-hairs of Belisana-type (cf. Huber &

Fleclcenstein 2008) (Figs 25, 39).

Female

Similar to male but eye triads on low humps (Fig. 34); clypeus and chelicerae unmodified; legs slightly

shorter than in male (tibia 1 -4.0-5.0); without spines on femora 1. Epigynum weakly sclerotized, with

distinct scape directed toward posterior (Figs 12, 29, 40M5). Internal genitalia with pair of oval pore

plates.

Monophyly and relationships

In a recent morphological cladistic analysis of Aetana and putatively close relatives (Huber et al. 2015a),

the monophyly of Hantu gen. nov. was consistently supported by five synapomorphies: (1) presence of

ventral apophysis on male palpal coxa (Figs 9, 19, 27); (2) male palpal trochanter apophysis with small

teeth or scales (Fig. 16); (3) presence of spines on male femora 1; (4) high density of vertical hairs on

male femora; (5) presence of scape on epigynum (Figs 40—45).

Three characters suggested a closer relationship with the mainland genera Khorata and Savarna rather

than with Aetana: (1) thoracic furrow extending over entire length of carapace (Figs 14, 31, 32); (2) dark

(usually black) sternum; and (3) absence of epiandrous spigots (Figs 23,32). A single character supported

a sister-group relationship with Savarna : high density of vertical hairs on tibiae. Preliminary molecular

data (A. Valdez-Mondragon, D. Dimitrov, B.A. Huber, unpubl. data) are not conclusive: while H. niah

gen. et sp. nov. is placed in a clade together with Khorata and Savarna (in agreement with morphology),

H. kapit gen. et sp. nov. is currently misplaced in Belisana Thorell, 1898 for unknown reasons.

Natural history

Both species were found in small protected spaces close to the ground (in small holes; among and under

rocks, logs, and large leaves) where they build their domed webs.

Composition

Only the two species newly described below.

Distribution

Known from Sarawak only (Fig. 7).

European Journal of Taxonomy 186: 1-15 (2016)

Hantu kapit gen. et sp. nov.

um:lsid:zoobank.org:act:C86E7783-5E91-4FB9-AB7E-A7F32FDA9431

Figs 1-2, 8-25, 40^12

Gen.n. Bor80 - Huber et al. 2015a: 73-74.

Diagnosis

Easily distinguished from H niah gen. et sp. nov. (and from other putatively close relatives in the

genera Khorata, Savarna, and Aetana) by two pairs of horns on male ocular area (Figs 10, 14, 15, 17),

Figs 1-6. Five specimens. 1-2. Hantu kapit gen. et sp. nov., $ and $ from Kapit, Sarawak. 3-6. Hantu

niah gen. et sp. nov., 3 and $ from Niah, Sarawak.

HUBER B.A., New spider genus from Sarawak

by extremely long apophysis on male palpal trochanter (longer than femur; Fig. 9), and by scape on

epigynum (Fig. 40) less sclerotized than in H. niah gen. et sp. nov. and with fine transversal folds.

Etymology

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

Type material

MAFAYSIA-BORNEO: holotype, S, Sarawak, nearKapit, forest along river, 1.937-1.944° N, 112.904-

112.910° E, 80-120 m a.s.l., 20 Jul. 2014 (B.A. Huber, S.B. Huber), ZFMK (Ar 15068).

Other material examined

MAFAYSIA-BORNEO, Sarawak: 7 4 $$, same data as holotype, ZFMK (Ar 15069-70; 6 <$<$,

3 5$) and SMK (1 S, 1 ?); 2 $ ?, 4 juvs, same data, in pure ethanol, ZFMK (Bor 183).

Description

Male (holotype)

Measurements. Total body length 2.4, carapace width 0.9. Teg 1: 24.5 (5.8 + 0.4 + 5.7 + 9.3 + 3.3),

tibia2: 3.4, tibia 3: 2.4, tibia 4:3.1; tibia 1 F/d: 72. Distance PME-PME 500 pm, diameter PME 105 pm,

distance PME-ALE 25 pm, no AME.

Color. Carapace pale ochre with large dark lateral marks and median mark posteriorly, dark median line,

ocular area and clypeus dark brown. Sternum dark brown to black. Legs ochre to light brown, without

dark rings. Abdomen ochre-grey with large dorsal and lateral marks; ventral side largely covered by

large dark mark and smaller dark mark at spinnerets.

European Journal of Taxonomy 186: 1-15 (2016)

Body. Habitus as in Fig. 1; ocular area highly modified (Figs 10, 14, 15, 17), with two pairs of large

horns arising from near PME and PLE and pair of smaller processes arising from near ALE; each triad

on short stalk directed toward lateral; carapace with distinct thoracic furrow; clypeus with small median

Figs 8-13. Hantukapit gen. et sp. nov., ZFMKAr 15069-70.8-9. Left male palp, prolateral andretrolateral

views (arrow points at ventral process on coxa). 10. Male prosoma, oblique frontal view. 11. Male

chelicerae, frontal view. 12-13. Cleared female genitalia, dorsal and ventral views. Abbreviations: b =

genital bulb; e = embolus; p = procursus; tr = trochanter. Scale lines: 0.5 mm (8-10); 0.3 mm (11-13).

HUBER B.A., New spider genus from Sarawak

process (Figs 15, 17); sternum wider than long (0.60/0.50), unmodified. Chelicerae as in Fig. 11, with

pair of proximal lateral apophyses in rather frontal position and pair of small, distal apophyses near

lamellae; without modified hairs.

Palps. As in Figs 8-9, coxa with sclerotized protruding rim ventro-distally; trochanter with distinctive

long retrolateral apophysis widening distally and provided with small teeth at tip (Fig. 16); femur large.

Figs 14-19. Hantu kapit gen. et sp. nov., ZFMK Ar 15069. 14-15. Male prosoma, frontal and slightly

oblique frontal views. 16. Tip of right male palpal trochanter apophysis. 17. Male prosoma, oblique

frontal view. 18. Feft procursus and genital bulb, prolateral view. 19. Right procursus and bulb,

retrolateral view (arrow points at ventral process on coxa). Abbreviations: b = genital bulb; e = embolus;

p = procursus. Scale lines: 50 pm (16); 200 pm (14-15, 17-19).

European Journal of Taxonomy 186: 1-15 (2016)

without processes; patella unusually long; tibia long and slender. Procursus highly complex, apparently

with several hinges between distinctive sclerites. Bulb simple, weakly sclerotized embolus with subdistal

branch.

Legs. With single ventral row of ~25 spines on femora 1; without curved hairs; with short vertical hairs

in two dorsal rows on all femora and tibiae; retrolateral trichobothrium on tibia 1 at 6%; prolateral

Figs 20-25. Hantu kapit gen. et sp. nov., ZFMK Ar 15069. 20. Tip of right procursus, retrolateral view.

21. Male palpal tarsal organ. 22. Trichobothrium base on male palpal tibia. 23. Male gonopore. 24. Male

ALS. 25. Comb-hairs on male tarsus 4. Scale lines: 8 pm (21); 10 pm (22, 24-25); 40 pm (20, 23).

HUBER B.A., New spider genus from Sarawak

trichobothrium absent on tibia 1, present on other tibiae. Tarsus 1 pseudosegments indistinct, only

distally ~15 visible in dissecting microscope.

Male (variation)

Tibia 1 in 7 other males: 5.6-5.9 (mean 5.7). Spines on femora variable in thickness and number (~20-30).

Female

In general similar to male (Fig. 2); ocular area unmodified, triads closer together (distance PME-PME

220 pm), not on stalks; clypeus unmodified; no spines on femora; fewer vertical hairs on femora and

tibiae (but especially on tibiae also with higher than usual density). Tibia 1 in 4 females: 4.7, 4.8, 4.9,

4.9. Epigynum large dark plate, wealdy sclerotized, slightly protruding and with many fine transversal

folds, especially in posterior half (Figs 13, 40, 41); with distinctive scape, pale whitish distally; internal

genitalia as in Figs 12 and 42.

Natural history

The spiders were found in domed webs in small holes in the ground along a river in well-preserved

forest.

Distribution

Known from type locality near Kapit only (Fig. 7).

Hantu niah gen. et sp. nov.

um:lsid:zoobank.org:act:0C378095-994C-427A-8989-E119A644A7EC

Figs 3-6, 26-39, 43-15

Gen.n. Bor20 - Huber et al. 2015a: 73-74.

Diagnosis

Easily distinguished from H. kapit gen. et sp. nov. by absence of horns in male ocular area (Fig. 31),

by highly distinctive male palp (Figs 26-27; square-shaped trochanter apophysis; dorsal process on

procursus), by strong distal apophyses on male chelicerae (Fig. 28), and by scape on epigynum (Figs 29,

43) more sclerotized and without fine transversal folds. From other putatively close relatives in the

genera Khorata and Savarna by male procursus shape (dorsal process) and epigynal scape.

Etymology

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

Type material

MAFAYSIA-BORNEO: holotype, Sarawak, Niah Cave N.R, forest near cave, 3.814° N, 113.771° E,

40 m a.s.l., 28 Jul. 2014 (B.A. Huber), ZFMK (Ar 15071).

Other material examined

MAFAYSIA-BORNEO, Sarawak: 10 $$, 15 §§, 1 juv., same data as holotype, ZFMK (Ar 15072-73;

9 <$<$, 14 55) and SMK (1 1 §); 4 same data, in pure ethanol, ZFMK (Bor 226); 2 5?, Niah

Cave N.R, forest along main trail, 3.814-3.821° N, 113.763-113.771° E, 20^10 m a.s.l., 27 Jul. 2014

(B.A. Huber, S.B. Huber), ZFMK (Ar 15074); 1 §, 1 juv., same data, in pure ethanol, ZFMK (Bor 177).

European Journal of Taxonomy 186: 1-15 (2016)

■

Figs 26-30. Hantu niah gen. et sp. nov., ZFMK Ar 15072-73. 26-27. Left male palp, prolateral and

retrolateral views (arrow points at ventral process on coxa). 28. Male chelicerae, frontal view. 29-

30. Cleared female genitalia, ventral and dorsal views. Abbreviations: b = genital bulb; e = embolus;

p = procursus; tr = trochanter. Scale lines: 0.5 mm (26-27, 29-30); 0.3 mm (28).

HUBER B.A., New spider genus from Sarawak

Figs 31-39. Hantu niah gen. et sp. nov., ZFMK Ar 15072-73. 31. Male prosoma, anterior view. 32. Male

gonopore. 33. Male palpal tarsal organ. 34. Female prosoma, frontal view. 35. Tip of right procursus,

retrolateral view. 36. Famellae of left procursus, prolateral view. 37. Epigynum, ventral view. 38. Male

AES. 39. Comb-hairs on male tarsus 4. Scale lines: 8 pm (33); 10 pm (38-39); 30 pm (32, 35); 80 pm

(36); 200 pm (31,34,37).

European Journal of Taxonomy 186: 1-15 (2016)

Description

Male (holotype)

Measurements. Total body length 2.3, carapace width 1.0. Leg 1: 20.5 (5.0 + 0.4 + 5.1 + 7.7 + 2.3),

tibia 2: 3.0, tibia 3: 2.2, tibia 4: 2.9; tibia 1 L/d: 58. Distance PME-PME 420 pm, diameter PME 90 pm,

distance PME-ALE 25 pm, no AME.

Color. Carapace light ochre with narrow lateral dark margins and large median dark mark including

ocular area. Clypeus not darkened. Sternum dark brown. Legs ochre to light brown, without dark rings.

Abdomen grey with large dorsal and lateral marks; ventrally with three large dark marks (in genital area,

at spinnerets, and in-between).

Body. Habitus as in Figs 3—4; ocular area slightly raised, each triad on short stalk directed toward lateral

(Fig. 31); carapace with shallow but distinct thoracic furrow; clypeus unmodified; sternum wider than

long (0.65/0.55), unmodified. Chelicerae as in Fig. 28, with pair of proximal lateral apophyses and pair

of strong distal apophyses curved toward median line; without modified hairs.

Figs 40-45. Female genitalia; untreated in ventral view, cleared in ventral and dorsal views. 40-42 Hantu

kapit gen. et sp. nov., ZFMK Ar 15070. 43^5. Hantu niah gen. et sp. nov., ZFMK Ar 15073. At varying

scales.

HUBER B.A., New spider genus from Sarawak

Palps. As in Figs 26-27, coxa with sclerotized protruding rim ventro-distally; trochanter with short but

very large, square-shaped ventral apophysis with very small teeth distally; femur very large, without

processes; patella unusually long; tibia long and slender. Procursus very long; with distinctive dorsal

process and complex tip. Bulb oval, with simple weakly sclerotized embolus.

Legs. With single ventral row of ~10 spines on femora 1; without curved hairs; with short vertical hairs

in two dorsal rows on all femora and tibiae; retrolateral trichobothrium on tibia 1 at 7.5%; prolateral

trichobothrium absent on tibia 1, present on other tibiae. Tarsus 1 pseudosegments indistinct, only

distally ~20 visible in dissecting microscope.

Male (variation)

Tibia 1 in 8 other males: 4.8-5.3 (mean 5.0). Margins of carapace sometimes not dark; dorsal pattern

on abdomen sometimes indistinct. Number and thickness of spines on femora variable, from few barely

thicker hairs (small males) to ~20 distinct spines (large males).

Female

In general similar to male (Figs 5-6); eye triads closer together (distance PME-PME 220 pm), not on

stalks (Fig. 34); no spines on femora; fewer vertical hairs on femora and tibiae (but especially on tibiae

also with higher than usual density). Tibia 1 in 16 females: 4.3M.7 (mean 4.5). Entire epigynal area

protruding, posteriorly sclerotized with distinctive scape (Figs 29, 43, 44); internal genitalia as in Figs

30 and 45.

Natural history

The spiders were found close to the ground in small domed webs among and under rocks, under dead

leaves, and in little cavities. They were extremely efficient at escaping through the closed hand, then

dropped to the ground and remained motionless, becoming essentially invisible.

Distribution

Known from Niah Cave N.P only (Fig. 7).

Discussion

Pholcid spiders are widely known for their long-legged representatives, some of which are synanthropic,

but a large number of species in a range of genera are actually relatively short-legged ground and litter-

dwellers. About half of all currently recognized genera either include or consist entirely of such short¬

legged species (e.g., Huber 2005a, 2005b, 2011, 2013, 2015; Huber et al. 2005). This suggests multiple

convergent shifts among microhabitats; in fact, molecular data support the notion that such shifts have

occurred repeatedly in various directions (Huber et al. 2010; Dimitrov et al. 2013; see also Huber &

Dimitrov 2014).

Ground and litter-dwelling pholcids share a similar habitus to a degree that allows reasonable predictions

even for museum specimens without microhabitat information. They are small (body size ~1—3 mm),

relatively short legged (leg 1 length <30 mm), rather dark (brown), and have a globular or oval abdomen.

This combination seems to be extremely rare in pholcids living in other microhabitats. The only apparent

exception known to me are West and Central African representatives of the genus Anansus Huber, 2007

that were collected by canopy fogging (Huber 2007).

In Southeast Asia, at least seven pholcid genera other than Hantu gen. nov. include ground and litter¬

dwelling representatives: Aetana Huber, 2005; Belisana Thorell, 1898; Holocneminus Berland, 1942;

Pholcus Walckenaer, 1805; Savarna Huber, 2005; Spermophora Hentz, 1841; and Wugigarra Huber,

2001 .

European Journal of Taxonomy 186: 1-15 (2016)

Acknowledgements

I thank Charles Leh Moi Ung and the Sarawak Museum Department for assistance in the request of

permits and in field logistics, and Samuel B. Huber for help in the field. The Natural History Museum

in Kuching kindly offered its facilities as the base of the field work. Permits were received from the

Forest Department, Sarawak (Permit No. NCCD.907.4.4(JLD.10)-211, Park Permit No. 174/2014). I

thank Facundo M. Labarque and an anonymous reviewer for valuable comments on the manuscript. The

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

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Manuscript received: 12 October 2015

Manuscript accepted. 10 December 2015

Published on: 1 April 2016

Topic editor: Rudy Jocque

Desk editor: Kristiaan Hoe demakers

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, De nm ark