European Journal of Taxonomy 162: 1-78

http://dx.doi.org/! 0.5852/ej t .2015.162

ISSN 2118-9773

www.europeanjournaloftaxonomy.eu

2015 • Huber B.A. et al.

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

Monograph

urn:lsid:zoobank.org:pub:BC89C4DA-4346-4B84-8A54-976F9741636B

Revision, phylogeny, and microhabitat shifts in the Southeast Asian

spider genus Aetana (Araneae, Pholcidae)

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

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

2 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: [email protected]

2 Email: [email protected]

3 Email: charllmikAsarawak.gov,my

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

2 um:lsid:zoobank.org:author:6D14FCCC-F148-4293-9F23-0EB5A4119305

3 urn:lsid:zoobank.org:author:8738F336-D1CC-4AD3-97E4-82BC1729A123

Abstract. The previously poorly known Southeast Asian spider genus Aetana Huber, 2005 is revised.

Fifteen species are newly described, and the first SEM data and a first phylogenetic analysis of the genus

are presented. Four species groups are well supported, one restricted to Borneo, two restricted to the

Philippines, and one ranging from the Philippines to Fiji. The cladistic analysis and field observations

suggest that the ancestor of Aetana built its web close to the ground, in confined spaces among and under

rocks and logs. In at least two cases, evolutionary shifts of microhabitat resulted in species being adapted

to life in higher forest strata, with correlated morphological and behavioral changes (lighter coloration;

longer abdomen; additional sheet in web or more strongly domed web). The following species are

newly described: A. abadae Huber, sp. nov., A. baganihan Huber, sp. nov., A. banahaw Huber, sp. nov.,

A. kiukoki Huber, sp. nov., A. libjo Huber, sp. nov., A. loboc Huber, sp. nov., A. lozadae Huber, sp. nov.,

A. manansalai Huber, sp. nov., A. ocampoi Huber, sp. nov., A. paragua Huber, sp. nov. and A. pasambai

Huber, sp. nov. from the Philippines; A. gaya Huber, sp. nov., A. indah Huber, sp. nov., A. lambir Huber,

sp. nov. and A. poring Huber, sp. nov. from northern Borneo. The female of A. kinabalu Huber, 2005 is

newly described. A potential case of female genital dimorphism is documented in A. ocampoi Huber,

sp. nov.

Keywords. Pholcidae, Aetana , Southeast Asia, cladistic analysis, microhabitat shift.

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

European Journal of Taxonomy 162 : 1-78 ( 2015 )

Introduction

The genus Aetana Huber, 2005 was originally created in the context of a major effort to resolve the

taxonomic chaos of small six-eyed pholcine spiders (e.g., Huber 2003a,b,c, 2005a,b). Previously, these

pholcids were mostly subsumed in the wastebasket genus Spermophora Hentz, 1841. Most species

originally described in Spermophora have been removed, either to other ‘old’ genera like Pholcus

Walckenaer, 1805; /C7 is ana T h o re11,1898; Metagonia Simon, 1 893, Anopsicus Chamberlin & I vie, 1938;

Paramicromerys Millot, 1946; Panjange Deeleman-Reinhold & Deeleman, 1983; and Spermophorides

Wunderlich, 1992; or to newly created genera like Buitinga Huber, 2003; Zatavua Huber, 2003; and

Savarna Huber, 2005. However, despite these improvements, Spermophora and its relatives continue to

be problematic. This is at least partly due to the poor knowledge of some of the groups involved. The

genus Aetana Huber, 2005 is symptomatic of this.

When Aetana was created, only three species were known, based on a total of 19 adult specimens. They

originated from a cave on Tuzon (A. omayan Huber, 2005; 1 □, 2 □ □), a forest in Sabah (A. kinabalu

Huber, 2005; 2 □ □) and the Fiji Islands (A. fiji Huber, 2005; 4 □ □, 10 □ □). No data on ultrastructure

were available; nothing was known about their natural history beyond basic locality information on

labels; none of the material available was suitable for molecular work; and relationships to other

pholcine genera remained obscure. The present paper focuses mainly on improving this situation. It is

based on almost 500 newly collected specimens representing 17 species; we provide first SEM data on

several species representing different species groups; we performed the first formal cladistic analysis of

relationships within the genus and to other pholcine genera; and we provide the first natural history data.

An analysis of molecular data of most species will be presented in the context of a molecular phylogeny

of the entire family (A. Valdez-Mondragon, D. Dimitrov, B.A. Huber, unpublished data).

A secondary focus of this paper relates to microhabitat shifts in Pholcidae and ultimately to the

question of whether such shifts are causally related with species diversification. Multiple convergent

shifts among microhabitats may allow the testing of correlation between ecological diversification and

species diversification. A fundamental problem of such studies is the usually low sample size. Only if

an ecological shift has occurred multiple times independently is a test possible for a correlation between

shift and diversification (e.g., phytophagy in insects: Mitter et al. 1988). Pholcidae may provide such

an opportunity. Shifts among microhabitats appear to have occurred multiple times convergently in

different directions: leaf litter <-► large protected spaces <-► among low vegetation <-► green leaves.

The latest molecular phylogeny of Pholcidae (Dimitrov et al. 2013) supports several such shifts, but

morphological data suggest that many more cases exist (B.A. Huber, unpubl. data). In Aetana , for

example, the dark coloration and short abdomen of A. omayan suggested a microhabitat near the ground

while the light coloration and long abdomen of A. kinabalu suggested a different microhabitat, higher

among the vegetation. We tested these assumptions in the field and combined them with a phylogenetic

analysis, thus providing one more piece in the mosaic of pholcid diversification.

Material and methods

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

Mar. 2014) and northern Borneo (July-Aug. 2014). Further specimens came from student projects at

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

diversity in the Philippines. The material is currently deposited at the Zoologisches Forschungsmuseum

Alexander Koenig, Bonn (ZFMK), Mindanao State University - Iligan Institute of Technology, Iligan

(MSU-IIT), and Sarawak Museum, Kuching (SMK). Further material came from the California

Academy of Sciences, San Francisco (CAS) and the Netherlands Centre for Biodiversity Naturalis,

Leiden (RMNH).

HUBER B.A. et al ., Revision of Aetana spiders

Zatavua griswoldi

Spermophora estebani

7 22

—oo

1 1

1 5 32

2 1 1

7 18

— 0-0

1 1

- Khorata khammouan

6 11 15 25

>—O

KXK>4-

1113

Sa varna thaleban

10 13 27 28 35

♦ Off Q-

11111

25 35

•C

1 1

9 11 20 26

1111

r-O

25 29 36 39 44

'—moomo

2 1111

Gen.n. Bor80

Gen.n. Bor20

A. ocampoi

— A. libjo

— A. baganihan

4 6 18 41

>—•OO#-

3 111

2 15

9-C

1 1

30 31

0 1

i 1

L —O— A. lambir

A. poring

A. indah

A. abadae

A. omayan

A. manansalai

A. lozadae

A. banahaw

ocampoi group

kiukoki group

kinabalu group

omayan group

Fig. 1. Single most parsimonious cladogram resulting from analyses of the matrix in Appendix 1. For

characters and character states see Appendix 3. Only unambiguous character changes are shown. See

Cladistic analysis section for further details.

European Journal of Taxonomy 162: 1-78 (2015)

Species descriptions are ordered as in the cladogram in Fig. 1. Methods and terminology used are

as in recent revisions (Huber 2011, 2013). Measurements are in mm unless otherwise specified. Eye

measurements are ± 5 pm. Epigyna were cleared in a 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 maps were generated with ArcMap 10.0.

The following abbreviations are used in the text:

ALE

ALS

AME

a.s.l.

L/d

N.P.

PME

PMS

anterior lateral eyes

anterior lateral spinnerets

anterior median eyes

above sea level

length/diameter

National Park

posterior median eyes

posterior median spinnerets

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

The numerical cladistic analyses were done using NONA, version 2 (Goloboff 1993), Pee-Wee, version

2.8 (Goloboff 1997), and TNT, version 1.1 (Goloboff et al. 2004, 2008). The matrix is shown in

Appendix 1; terminal taxa and characters scored are given in Appendices 2 and 3. Of the 44 characters,

39 are binary; four non-binary characters are treated as non-additive (char. 4, 24, 25, 35). One character

is treated as additive (char. 1). The final matrix can be downloaded at http://www.pholcidae.de/matrices.

html . Cladogram analysis was done with Winclada, version 1.00.08 (Nixon 2002). See Cladistic analysis

section below for details of the analyses.

Results

Class Arachnida Cuvier, 1812

Order Araneae Clerck, 1757

Family Pholcidae C.L. Koch, 1851

Genus Aetana Huber, 2005

Cladistic analysis

Using NONA with hold/100, mult*200 (or hold/10; mult* 10.000), and amb- for the matrix in Appendix

1 and equal character weights resulted in a single most parsimonious cladogram with a length of 80

(Cl = 67; RI = 84) (Fig. 1). All other analyses resulted in the same cladogram: TNT using the implicit

enumeration algorithm; successive weighting in NONA (with the consistency index as weighting

function); and implied weighting in Pee-Wee (which resolves character conflict in favor of the characters

that have less homoplasy).

Taxonomy

Aetana Huber, 2005

Aetana Huber, 2005a: 72-73. Type species: A. omayan Huber, 2005.

HUBER B.A. et ah , Revision of Aetana spiders

Diagnosis

Even though the cladistic analysis identifies only two synapomorphies for Aetana , the genus is fairly

easily distinguished from the putatively closest relatives by the following characters: retrolateral

trichobothrium on leg 1 very proximal (at <5% of tibia length) and presence of curved hairs on tibiae

and / or metatarsi (both in contrast to Spermophora, Khorata Huber, 2005, Savarna , and an undescribed

genus from Sarawak, below called ‘Gen.n. Borneo’); sternum not dark, retrolateral trichobothrium on

male palpal tibia in very distal position, and presence of epiandrous spigots (all in contrast to Khorata ,

Savarna , and ‘Gen.n. Borneo’); male legs without spines and male palpal coxa unmodified (both in

contrast to ‘Gen.n. Borneo’); ALS with only two spigots, epigynal plate without external pair of pockets,

and female genitalia without unpaired posterior pocket (all in contrast to Spermophora). Most characters

previously thought to be diagnostic (Huber 2005a) are rendered invalid due to the newly described

species.

Description

Male

Measurements. Total body length -2.5M.5 (smallest species in A. kinabalu group; largest species in

A. omayan group); carapace width 0.9-1.8; leg 1 length —27—44; tibia 1 length -6.0-11.0; tibia 2/tibia

4 length 0.92-1.08; tibia 1 L/d -55-95 (the largest species, A. omayan , has the relatively thickest legs;

the smallest species, A. gaya , has the relatively thinnest legs).

Color. In life mostly ochre-gray with brown and black marks (e.g., Figs 51-56, 105-110), only A. libjo

Huber, sp. nov. and A. baganihan Huber, sp. nov. with light brown to orange prosoma and palps (Figs

8-12); sternum never dark; legs usually with indistinct darker rings on femora (subdistally) and tibiae

(proximally and subdistally); darker rings missing in A. libjo Huber, sp. nov. and A. baganihan Huber,

sp. nov.

Body. Carapace either with shallow median furrow restricted to frontal part (Figs 63-64, 198) or without

furrow (Figs 118, 130); ocular area raised, eye triads on short stalks directed toward lateral, in some

species with median process (Figs 77, 96) or with pair of processes arising from near AFE (Figs 221,

226). AME absent. Clypeus high, either unmodified, with small paired processes (Fig. 191), with large

median process (Figs 59, 77), or with indistinct lateral ridges (Figs 221, 226). Abdomen from slightly

longer than high (Figs 51-56) to almost cylindrical (Figs 8, 102), pointed at spinnerets. Male gonopore

with four epiandrous spigots in all species examined with SEM (e.g., Figs 37, 143, 162); each AES with

large widened spigot and pointed spigot, without further cylindrically shaped spigots (Figs 68, 210);

PMS with two spigots each.

Chelicerae. Very variable, distal apophyses ranging from short processes in frontal position (Figs 15,

40) to long processes in lateral position (Figs 191, 226), absent in A. kiukoki group; proximal apophyses

usually present, large in A. kiukoki group (Fig. 66), absent in A. ocampoi group (Figs 15, 40); chelicerae

without modified hairs; without stridulatory ridges.

Palps. Coxa unmodified; trochanter usually with one retrolateral to ventral process, sometimes provided

with scales or teeth (Fig. 158), in A. ocampoi group fused to femur (Figs 14, 27); trochanter in some

species with additional prolateral apophysis (Fig. 189); femur rarely simple (Fig. 27), usually with one or

more processes, inA. kinabalu group with complex set ofup to fourprocesses (e.g., Figs 150-151); patella

either triangular in lateral view or ventral side longer than usual (e.g., Figs 14, 58); tibia either of usual

shape (e.g., Figs 14, 127) or rather small and slender (A. kiukoki group; e.g., Fig. 58), with retrolateral

trichobothrium in very distal position (close to tibia-tarsus joint; Figs 27, 114); palpal tarsus small, usually

with capsular tarsal organ (Figs 134, 203), exposed in A. libjo Huber, sp. nov. (Fig. 35) (not clear in

A. ocampoi Huber, sp. nov. and A. baganihan Huber, sp. nov.); procursus usually complex with proximal

European Journal of Taxonomy 162: 1-78 (2015)

and distal parts connected by membranous hinge; procursus in A. kiukoki group unusually long (Figs 58,

76), in A. ocampoi group reduced to simple semi-transparent process (Figs 14,27); bulb either with weakly

sclerotized embolus as only process (Figs 114,159), in A. kiukoki group with additional short membranous

process (Figs 58, 75), in A. ocampoi group with additional novel processes (Figs 13-14, 26-27).

Legs. Without spines; with curved hairs on tibiae and/or metatarsi (Fig. 133); usually without or with

few vertical hairs, only in A. libjo Huber, sp. nov. and A. baganihan Huber, sp. nov. with one dense row

retrolatero-dorsally on each tibia; retrolateral trichobothrium on tibia 1 very proximal (at 2-4% of tibia

length), prolateral trichobothrium absent on tibia 1, present on other tibiae. Tarsus 1 usually with -25-30

pseudosegments, fairly distinct distally; tarsus 4 with single row of ventral comb-hairs of a modified

Belisana-tyyQ ( cf Huber & Fleclcenstein 2008; Figs 32, 69, 82, 209).

Female

Similar to male but eye triads on lower humps (Figs 64, 80, 119) and closer together than in male

(sexual dimorphism low in A. ocampoi group; in other groups, eye triads in females often less than

half as wide apart as in males); clypeus and chelicerae unmodified; legs slightly shorter than in males

(tibia 1 -4.0-9.0). Females in A. omayan group with stridulatory apparatus between prosoma (elongate

median plate on carapace posteriorly; Figs 186-188) against abdomen (indistinct hairless area frontally).

Epigynum wealdy to heavily sclerotized, sometimes with scape of variable length and shape (Figs 28,

60), never with external pair of pockets in anterior epigynal plate but in A. kinabalu and A. omayan

groups sometimes with internal sclerotized pockets originating from ventral wall of uterus externus

(Figs 111, 192-194), and in A. omayan group with pair of membranous external pockets in posterior

epigynal area (Figs 192, 194, 222). Internal genitalia with pair of pore plates, in A. libjo Huber, sp. nov.

and A. baganihan Huber, sp. nov. with unique median membranous structure (Figs 29, 44); in A. kiukoki

group with distinctive serrated ridges (Figs 61, 73, 79).

Fig. 2. Known distribution of Aetana. Shown are also three undescribed species that are known from

poorly preserved specimens only.

HUBER B.A. et al. , Revision of Aetana spiders

Monophyly and relationships

Morphologically, the monophyly of Aetana appears weakly supported. Only two characters support

this node (Fig. 1), both of them without homoplasy within the taxa included in the matrix but with

considerable homoplasy among more distant Pholcinae relatives: (1) the very proximal position of the

retrolateral trichobothrium on the leg tibiae (char. 30); and (2) the presence of curved hairs on leg tibiae

and/or metatarsi (char. 31). However, preliminary analyses of molecular data (including many more

potential close relatives) (A. Valdez-Mondragon, D. Dimitrov, B.A. Huber, unpubl. data) consistently

support the monophyly of Aetana with high support values.

The sister group of Aetana appears much better supported by morphology. Four characters suggest that

three genera together ( Khorata , Savarna , and ‘Gen.n. Borneo’) are sister to Aetana. This, however, is

in conflict with our preliminary molecular data that suggest a closer relationship between Aetana and

East Asian Spermophora (incl. S. estebani) than between Aetana and Khorata , Savarna , and ‘Gen.n.

Borneo’. Such a close relationship with East Asian Spermophora was actually proposed in the original

description (Huber 2005a), at that time without a formal cladistic analysis.

Figs 3-4. Known distributions of the Aetana ocampoi (3) and A. kiukoki (4) groups. The question mark

denotes a female specimen assigned tentatively to A. kiukoki.

European Journal of Taxonomy 162: 1-78 (2015)

Within Aetana, four species groups receive strong support both from the morphological analysis herein

(Fig. 1) and from our preliminary molecular data. Conflict exists regarding relationships among these

groups. The present analysis supports a sister group relationship between the Aetana kinabalu and

A. omayan groups, and at least one of the three characters supporting this node is a unique feature (the

long retrolateral membranous process on the procursus; char. 21). Our preliminary molecular data do not

strongly support any sister group relationships among the four species groups within Aetana.

Natural history

Most species were collected in forests, from well protected spaces close to the ground, under rocks and

logs, in small holes and cavities. Few species occur higher in the vegetation, even in places directly

Fig. 5. Known distributions of the Aetana kinabalu (Borneo) and A. omayan (Philippines) groups.

HUBER B.A. et al ., Revision of Aetana spiders

reached by the sun, most notably the closely related A. libjo Huber, sp. nov. and A. baganihan Huber,

sp. nov. and the putatively close relatives A. kinabalu and A. lambir Huber, sp. nov. An exception is the

type species A. omayan, which was mainly collected in a cave but also among rocks in a treeless ravine.

All species seem to build simple domed sheet webs (like most other pholcids studied), but the sheets

are unusually strongly domed in A. libjo Huber, sp. nov. and A. baganihan Huber, sp. nov., and in

A. kinabalu and A. lambir Huber, sp. nov. a second sheet occurs a few cm under the main sheet.

In six cases, two species were found to share a locality, sometimes in different microhabitats (e.g., at

Baganihan: A. kiukoki Huber, sp. nov. near the ground; A. baganihan Huber, sp. nov. among vegetation),

sometimes in what seemed to be identical microhabitats (e.g., at Mt. Banahaw: A. manansalai Huber,

sp. nov. and A. banahaw Huber, sp. nov., both close to the ground).

In most species, males and females were often found together sharing a web. This was never observed

in A. libjo Huber, sp. nov. and A. baganihan Huber, sp. nov., where males and females were sometimes

observed very close to each other but in separate webs.

When disturbed, Aetana spiders tend to run toward the periphery of the web rather than to vibrate in the

‘typical" pholcid way. Some do then vibrate vigorously for a very short time before becoming motionless

and pressing their body against the substrate; others stop vibrating and start to gently move the abdomen

in circles (A. libjo Huber, sp. nov. and A. baganihan Huber, sp. nov.). The closely related A. poring

Huber, sp. nov. and A. indah Huber, sp. nov. barely reacted to disturbance.

Composition

The genus now includes 18 described species. The high species turnover in the Philippines and in

northern Borneo, together with the wide distribution of one species group (A. omayan group; Philippines

to Fiji Islands) and the huge sampling gaps in eastern Indonesia, New Guinea, and east to Fiji suggest

that at least several dozen further species are likely to exist.

Distribution

Ranging from northern Borneo and the Philippines to Fiji (Fig. 2). Most gaps and missing records are

probably due to lack of adequate sampling, but the absence of records from two areas might reflect

real absence. First, our intensive collecting in western Sarawak (at seven localities ranging from the

Pueh foothills in the west to Niah in the east) did not result in a single specimen of Aetana, while many

specimens were collected at all eight localities in eastern Sarawak (east of Niah) and Sabah. Second,

northern Australia is relatively well sampled, but a revision of all the material available in collections

(Huber 2001) did not reveal any Aetana.

Aetana ocampoi Huber, sp. nov.

urn:lsid:zoobank.org:act:85B2FF02-FB08-48B3-91E9-0AB54D7866F5

Figs 6-7, 13-25

Diagnosis

Easily distinguished from closest known relatives (A. libjo Huber, sp. nov.; A. baganihan Huber, sp. nov.)

by dark coloration (Figs 6-7) and by several details of the male palp (Figs 13-14; long cylindrical

bulb; ventral process on femur), male chelicerae (Fig. 15; more proximal position of apophyses), and

epigynum (Figs 16-19; short triangular scape).

European Journal of Taxonomy 162: 1-78 (2015)

Figs 6-12. Live specimens. Aetana ocampoi group. 6-7. A. ocampoi Huber, sp. nov., P from Mt. Isarog,

Luzon. 8-9. A. libjo Huber, sp. nov., 0 and 0 from Dinagat Island, Mindanao. 10-12. A. baganihan

Huber, sp. nov., □ □ from Baganighan, Mindanao.

HUBER B.A. et al ., Revision of Aetana spiders

Etymology

Named for the Filipino artist Hernando Ruiz Ocampo (1911-1978), famous for his work reflecting the

harsh realities of his country after the Second World War, but also for his interest in depicting Philippine

flora and fauna.

Material examined

Holotype

PHILIPPINES: □, Luzon, Camarines Sur Prov., Mt. Isarog, W slope (13.664° N, 123.34-123.35° E),

-600-900 m a.s.l., forest, near ground, 23 Feb. 2014 (B.A. Huber), ZFMK (Ar 13927).

Figs 13-15. Aetana ocampoi Huber, sp. nov. 13-14. Left male palp, prolateral and retrolateral views

(asterisk: retrolateral process of bulb). 15. Male chelicerae, frontal view, b = genital bulb; p = procursus.

Scale lines: 13-14 = 0.5 mm; 15 = 0.2 mm

European Journal of Taxonomy 162: 1-78 (2015)

Other material

PHILIPPINES: Luzon, 1 □, 2 □ □ (morph A and morph B), same data as holotype, ZFMK (Ar 13928);

3 □ □ (2 □ □ morph A, 1 □ morph B), in pure ethanol, same data, ZFMK (Phi 222).

Description

Male (holotype)

Measurements. Total body length 2.7, carapace width 1.1. Leg 1: 28.3 (6.7 + 0.4 + 6.6 + 11.8 + 2.8),

tibia 2: 3.9, tibia 3: 2.7, tibia 4: 4.0; tibia 1 L/d: 59. Distance PME-PME 185 pm, diameter PME

125x120 pm, distance PME-ALE 20 pm; AME absent.

Color. Carapace pale ochre with wide median dark band including ocular area and narrower lateral

black margins; clypeus pale ochre, distal half light brown; sternum pale ochre, medially light brown; legs

light brown, with dark rings subdistally on femora and proximally on tibiae (incl. patellae); abdomen

dorsally densely covered with dark marks and some white marks, ventrally gray, small dark mark behind

gonopore, larger dark mark in front of spinnerets.

Figs 16-19. Aetana ocampoi Huber, sp. nov. Cleared female genitalia of morph A (16-17) and morph B

(18-19), in ventral (left) and dorsal (right) views. Scale line: 0.5 mm (all at same scale).

HUBER B.A. et al ., Revision of Aetana spiders

Body. Habitus as in Figs 6-7; ocular area slightly raised, each triad on low hump; carapace only

anteriorly with very shallow and narrow median furrow (rather just a dark line); clypeus unmodified;

sternum wider than long (0.60/0.55), unmodified.

Chelicerae. As in Fig. 15, with pair of frontal apophyses near median line, without proximal lateral

apophyses; without modified hairs; without stridulatory ridges.

Palps. As in Figs 13-14, coxa unmodified, trochanter on retrolatero-ventral side with large apophysis

fused to femur; femur with large retrolatero-ventral process; retrolateral trichobothrium on tibia very

distal; tarsus with semitransparent simple procursus; genital bulb large, cylindrical, with retrolateral

apophysis proximally and two distinctive processes distally: dorsal process with sclerotized black tip;

ventral process with semitransparent flap.

Fegs. Without spines, with curved hairs on tibiae 1 and metatarsi 1—4, few vertical hairs; retrolateral

trichobothrium on tibia 1 at 3%; prolateral trichobothrium absent on tibia 1, present on other tibiae.

Tarsus 1 with ~25 pseudosegments, distally distinct.

Male (variation)

Tibia 1 in other male: 7.2; this male paler but otherwise identical.

Figs 20-25. Aetana ocampoi Huber, sp. nov. Female genitalia of morph A (20-22) and morph B (23-

25); untreated in ventral view, cleared in ventral and dorsal views.

European Journal of Taxonomy 162: 1-78 (2015)

Female

In general similar to male; eye triads closer together (distance PME-PME 140 jam); tibia 1 in 2 females:

5.5, 5.7. Epigynum apparently dimorphic: three females with small triangular sclerotized scape (Figs

16, 20; morph A), two females with much wider sclerotized area of epigynum (Figs 18, 23; morph B);

internal genitalia also different, as in Figs 17, 19, 22, 25.

Natural history

The spiders were found close to the ground, apparently more deeply hidden under rocks and logs than

the sympatric A. lozadae Huber, sp. nov.

Distribution

Known from type locality only (Fig. 3).

Aetana libjo Huber, sp. nov.

urn:lsid:zoobank.org:act:91DA8483-221E-4FBB-8CD8-453CFD23B90F

Figs 8-9, 26M0, 45—47

Diagnosis

Distinguished from the very similar A. baganihan Huber, sp. nov. by shapes of bulbal processes

(compare Figs 38-39 and 41—42), by shorter male palpal tibia (2.1—2.4 x longer than wide vs. 2.9-3.2

in A. baganihan Huber, sp. nov.), and by shorter epigynum and wider scape (compare Figs 28 and 43).

Distinguished from next closest known relative (A. ocampoi Huber, sp. nov.) by light coloration and

several details of male palp and epigynum (especially shapes of bulb and scape; Figs 26-28). From all other

congeners by simple procursus and complex bulbal processes (Figs 26-27) and by narrow epigynal scape.

Etymology

Named for the type locality; noun in apposition.

Material examined

Holotype

PHIFIPPINES: □, Dinagat Isl., near Fibjo, Paragua Forest, ‘site V (10.222° N, 125.553° E), 130 m

a.s.l., forest at brook, among low vegetation, 20 Feb. 2014 (B.A. Huber), ZFMK (Ar 13929).

Other material

PHIFIPPINES: 6 3□, 9 □ □, same data as holotype, ZFMK (5D □, 8D □) (Ar 13930-31) and MSU-IIT

(1 □, 1 □); 3 juveniles in pure ethanol, same data, ZFMK (Phi 231).

Description

Male (holotype)

Measurements. Total body length 3.1, carapace width 1.1. Teg 1: 37.4 (9.0 + 0.5 + 8.8 + 16.2 + 2.9),

tibia 2: 5.1, tibia 3: 3.3, tibia 4: 4.8; tibia 1 F/d: 74. Distance PME-PME 270 pm, diameter PME 125x95

pm, distance PME-AFE 20 pm; AME absent.

Color. Carapace ochre-yellow with black lateral margins, dark mark behind ocular area and dark

median mark on posterior rim; ocular area and clypeus pale ochre; sternum bright orange; leg coxae

ochre-yellow, other leg segments greenish-ochre (especially femora) to light brown (distal segments);

abdomen grey with distinct dorsal and lateral pattern of black marks, ventrally with brown band between

gonopore and spinnerets (mid-section indistinct).

HUBER B.A. et al ., Revision of Aetana spiders

Figs 26-29. Aetana libjo Huber, sp. nov. 26-27. Left male palp, prolateral and retrolateral views (asterisk:

retrolateral process of bulb). 28-29. Cleared female genitalia, ventral and dorsal views (asterisk: median

membranous structure), b = genital bulb; p = procursus; rt = retrolateral trichobothrium. Scale lines:

26-27 = 0.5 mm; 28-29 = 0.3 mm

European Journal of Taxonomy 162: 1-78 (2015)

Figs 30-37. Aetana libjo Huber, sp. nov. 30. Processes of right genital bulb, retrolateral view. 31. Detail

of preceding. 32. Comb-hairs on female tarsus 4. 33. Processes of left genital bulb, prolateral view.

34. Detail of preceding. 35. Male palpal tarsal organ. 36. Female ALS. 37. Male gonopore. Scale lines:

30, 33 = 100 pm; 31, 37 = 40 pm; 32, 34 = 20 pm; 35-36 = 10 pm.

HUBER B.A. et al ., Revision of Aetana spiders

Body. Habitus as in Fig. 8; ocular area slightly raised, each triad on low hump directed toward lateral;

carapace only anteriorly with very shallow and narrow median furrow; clypeus barely modified,

indistinct median process near rim provided with some slightly stronger hairs; sternum wider than long

(0.70/0.55), unmodified. Gonopore with four epiandrous spigots in two pairs (Fig. 37). AFS as in female

(cf Fig. 36).

Figs 38-44. — 38-40. Aetana libjo Huber, sp. nov. 38-39. Heft genital bulb in prolateral and retrolateral

views. 40. Male chelicerae, frontal view. — 41—44. Aetana baganihan Huber, sp. nov. 41-42 Heft

genital bulb in prolateral and retrolateral views. 43-44 Cleared female genitalia, ventral and dorsal

views (asterisk: median membranous structure). Scale lines: 38-39, 41M4 = 0.5 mm; 40 = 0.3 mm.

European Journal of Taxonomy 162: 1-78 (2015)

Chelicerae. As in Fig. 40, with pair of ridge-shaped apophyses distally near median line, without

proximal lateral apophyses; without modified hairs; without stridulatory ridges.

Palps. As in Figs 26-27, coxa unmodified, trochanter on retrolateral side with large apophysis fused

to femur; femur with small retrolateral process distally; tibia length/width: 0.50/0.22; retrolateral

trichobothrium on tibia very distal; tarsus with semitransparent simple procursus directed toward

patella; tarsal organ exposed (Fig. 35); genital bulb large, with small retrolateral process proximally

and three distinctive processes distally (Figs 30-31, 33, 38-39): dorsal process with distal hook; central

hinged process with large prolateral flap and complex tip; ventral hinged process with simple pointed

tip. Location of sperm duct opening unknown.

Legs. Without spines, with curved hairs on metatarsi, with vertical hairs in higher than usual density

in one retrolatero-dorsal row on each tibia; retrolateral trichobothrium on tibia 1 at 2%; prolateral

trichobothrium absent on tibia 1, present on other tibiae. Tarsus 1 with ~30 pseudosegments, distally

distinct. Tarsus 4 comb-hairs with very dense tines (Fig. 32).

Figs 45-50. Aetana libjo Huber, sp. nov. (Figs 45^17) and Aetana baganihan Huber, sp. nov. (Figs

48-50), female genitalia; untreated in ventral view, cleared in ventral and dorsal views.

HUBER B.A. et al.. Revision of Aetana spiders

Male (variation)

Tibia 1 in 6 other males: 8.3-9.0 (mean 8.6); ratio of palpal tibia length/width: 2.1-2.4. Some specimens

with white marks on abdomen in addition to black marks.

Female

In general similar to male (Fig. 9); eye triads closer together (distance PME-PME 215 pm); clypeus

unmodified; tibiae with short vertical hairs in low density; entire median area of carapace darker; clypeus

also slightly darker; abdomen with or without white marks; tibia 1 in 9 females: 6.2-7.2 (mean 6.7).

Epigynum with large, wealdy sclerotized area, posteriorly protruding with short, narrow scape (Figs 28,

45M6); internal genitalia with large median membranous structure of unknown function (Fig. 29).

Natural history

The spiders were found in strongly domed webs among the vegetation. When disturbed, they ran a short

distance in the dome, vibrated vigorously for a short time, and then made slow circular movements with

their abdomen. Males and females were often found close to each other, but always in separate webs.

The locality is shared with the ground-dwelling A. paragua Huber, sp. nov.

Distribution

Known from type locality only (Fig. 3).

Aetana baganihan Huber, sp. nov.

urn:lsid:zoobank.org:act:AA06D026-A4F7-4EA9-9661-77707B2CCFB9

Figs 10-12, 41M4, 48-50

Diagnosis

Distinguished from very similar A. libjo Huber, sp. nov. by shapes of bulbal processes (compare Figs

38-39 and 41—42), by longer male palpal tibia (2.9-3.2 x longer than wide vs. 2.1-2.4 in A. libjo Huber,

sp. nov.), and by longer epigynum and narrower scape (compare Figs 28 and 43); from next closest

known relative (A. ocampoi Huber, sp. nov.) by light coloration and several details of male palp and

epigynum (especially shapes of bulb and scape). Distinguished from all other congeners by simple

procursus and complex bulbal processes and by narrow epigynal scape.

Etymology

Named for the type locality; noun in apposition.

Material examined

Holotype

PHIFIPPINFS: □, Mindanao, Davao del Sur Prov., Marilog Distr., Baganihan (7.469° N, 125.250° E),

1210 m a.s.l., primary forest near road, among low vegetation, 15 Feb. 2014 (B.A. Huber), ZFMK (Ar

13932).

Other material

PHIFIPPINES: Mindanao, 2 □□, 3 Dp same data as holotype, ZFMK (Ar 13933); 1 juvenile, in pure

ethanol, same data as holotype, ZFMK (Phi 258).

European Journal of Taxonomy 162: 1-78 (2015)

Description

Male (holotype)

Measurements. Total body length 3.2, carapace width 1.1. Leg 1: 39.3 (9.0 + 0.5 + 9.1 + 17.4 + 3.3),

tibia 2: 5.3, tibia 3: 3.6, tibia 4: 5.2; tibia 1 L/d: 77. Distance PME-PME 325 pm, diameter PME

135><110 pm, distance PME-ALE 20 pm; AME absent.

Color. Carapace ochre-yellow with black lateral margins, pair of small dark marks behind ocular area,

and dark median mark at posterior rim; ocular area and clypeus pale ochre; sternum bright orange;

leg coxae ochre-yellow, other leg segments greenish-ochre (especially femora) to light brown (distal

segments); abdomen grey, with distinct dorsal and lateral pattern of black marks and rows of white

marks, ventrally with brown band between gonopore and spinnerets (mid-section indistinct).

Body. Habitus as in Figs 10-12; ocular area slightly raised, each triad on low hump directed toward

lateral; carapace only anteriorly with very shallow and narrow median furrow; clypeus with indistinct pair

of processes near rim, provided with some slightly stronger hairs; sternum wider than long (0.70/0.55),

unmodified.

Chelicerae. As in A. libjo Huber, sp. nov. (cf. Fig. 40), with pair of ridge-shaped apophyses distally near

median line, without proximal lateral apophyses; without modified hairs; without stridulatory ridges.

Palps. Proximal segments very similar to A. libjo ( cf. Figs 26-27), trochanter apophysis and femur

apophysis slightly larger, tibia longer (length/width: 0.70/0.24). Genital bulb longer than in A. libjo

sp. nov., with all three processes different (Figs 41M2): dorsal process wider proximally; central hinged

process with smaller prolateral flap and different tip; ventral hinged process relatively shorter.

Legs. Without spines, with curved hairs on metatarsi 1-3, with vertical hairs in higher than usual density

in one retrolatero-dorsal row on each tibia; retrolateral trichobothrium on tibia 1 at 2%; prolateral

trichobothrium absent on tibia 1, present on other tibiae. Tarsus 1 with ~30 pseudosegments, distally

distinct.

Male (variation)

Tibia 1 in 2 other males: 8.9, 9.3; ratio of palpal tibia length/width: 2.9, 3.2. Other males without or with

indistinct white marks on abdomen.

Female

In general similar to male; eye triads closer together (distance PME-PME 230 pm); clypeus unmodified;

tibiae with short vertical hairs in low density; entire median area on carapace darker; clypeus also slightly

darker (one female with wide brown median band on carapace extending over ocular area and clypeus);

abdomen with or without white marks; tibia 1 in 3 females: 6.7, 6.8, 7.0. Epigynum with large, weakly

sclerotized area, posteriorly protruding with short, narrow scape (Figs 43, 48—49); internal genitalia with

large median membranous structure of unknown function (Fig. 44).

Natural history

The spiders were found in strongly domed sheet-webs among high grasses in well-preserved forest. As

in A. libjo Huber, sp. nov., males and females were sometimes found close to each other but in separate

webs. The locality is shared with the ground-dwelling A. kiukoki Huber, sp. nov.

Distribution

Known from type locality only (Fig. 3).

HUBER B.A. et al. , Revision of Aetana spiders

Aetana kiukoki Huber, sp. nov.

um:lsid:zoobank.org:act:23ED16FE-E6AA-40B4-B55E-A2C31CB86A22

Figs 51-52, 57-69, 87-89

Diagnosis

Distinguished from closest known relatives (A. paragua Huber, sp. nov., A. loboc Huber, sp. nov.,

A. pasambai Huber, sp. nov.) by distinctive modification of male clypeus (Figs 59, 62, 65; similar only in

A. paragua Huber, sp. nov., see Fig. 72), and long tongue-shaped posterior projection of epigynum (Figs

60, 87; very similar in A. paragua Huber, sp. nov., see Fig. 73; much shorter in A. loboc Huber, sp. nov.,

see Fig. 78; female of A. pasambai Huber, sp. nov. unknown). Distinguished from A. paragua Huber,

sp. nov. also by longer male eye stalks (Fig. 59), more strongly curved apophysis on male palpal femur

(Fig. 58), different shape of distal procursus elements (Fig. 58), and pore plates closer together (Fig. 61).

Distinguished from A. loboc Huber, sp. nov. and A. pasambai Huber, sp. nov. also by modification of

male palpal femur (only one large retrolateral process; Fig. 58) and absence of median process on male

ocular area.

Etymology

Named for Filipino painter Ang Kiukok (1931-2005).

Material examined

Holotype

PHIFIPPINES: □, Mindanao, Davao del Sur Prov., Marilog Distr., Baganihan (7.469° N, 125.250° E),

1210 m a.s.l., primary forest near road, near ground, 15 Feb. 2014 (B.A. Huber), ZFMK (Ar 13934).

Other material

PHIFIPPINES, Mindanao Isl.: 7 □□, 7 □□, 9 juvs, same data as holotype, ZFMK (Ar 13935-36); 1 □,

6 juvs, in pure ethanol, same data, ZFMK (Phi 256). - 1 □, 11 juvs from Barangay Baganihan, ‘site U

(7.438° N, 125.226° E), 1000 m a.s.l., 5 Dec. 2014 (M.A. Responte), MSU-IIT; 4 □□, 17 juvs, from

same locality, ‘site T (7.456° N, 125.239° E), 6 Dec. 2014 (M.A. Responte), MSU-IIT; 9 juvs, from same

locality, ‘site 3’ (7.470° N, 125.245° E), 7 Dec 2014 (M.A. Responte), MSU-IIT. -4 □ □, 1 juv., Marilog

Distr., Epol Spring Resort (7.456° N, 125.237° E), -1100 m a.s.l., degraded forest, near ground, 15 Feb.

2014 (B.A. Huber), ZFMK (Ar 13937); 3 □ t, in pure ethanol, same data, ZFMK (Phi 253). - 1 □, 1 □,

10 juvs, from Epol Falls, ‘site T (7.454° N, 125.239° E), 1150 m a.s.l., 2 Dec. 2014 (M.A. Responte),

MSU-IIT; 1 □, 4 juvs, from same locality, ‘site 3’ (7.451° N, 125.240° E), 1200 m a.s.l., 3 Dec. 2014

(M.A. Responte), MSU-IIT; 1 juv., from same locality, ‘site V (7.455° N, 125.237° E), 1100 m a.s.l., 1

Dec. 2014 (M.A. Responte), MSU-IIT. -6 □ □, 2 □ □, 2 juvs, Mt. Matutum, KawitForest, ‘site V (6.338°

N, 125.104° E), 950 m a.s.l., along brook, near ground, 13 Feb. 2014 (B.A. Huber), ZFMK (Ar 13938);

4 juvs, in pure ethanol, same data, ZFMK (Phi 268). - 1 □, 2 □ □, Bukidnon Prov., Barangay San Jose,

Blue Water Cave (7.706° N, 125.032° E), 200 m a.s.l., near ground at cave entrance, 16 Feb. 2014 (B.A.

Huber), ZFMK (Ar 13939); 4 □ □, 1 juv., in pure ethanol, same data, ZFMK (Phi 251). - 2 £!□, 2 fei

1 juv., Barangay San Jose, Kabyaw Cave (-7.704° N, 125.038° E), -200 m a.s.l., near ground near cave

entrance, 16 Feb. 2014 (B.A. Huber), ZFMK (Ar 13940). -4 □ 8 □ □, Barangay San Jose, doline near

Kabyaw Cave (7.703° N, 125.038° E), 220 m a.s.l., near ground, 16 Feb. 2014 (B.A. Huber), ZFMK (Ar

13941). - 3 □ □, Bukidnon Prov., CEDAR (Center for Ecological Development and Recreation) (8.251°

N, 125.034° E), 760 m a.s.l., forest along river, near ground, 16 Feb. 2014 (B.A. Huber), ZFMK (Ar

13942); 1 □, in pure ethanol, same data, ZFMK (Phi 246). - 1 □, 4 □ □, 3 juvs, from CEDAR (8.251° N,

125.027° E), 15 Nov. 2014 (E.P. Mondejar), MSU-IIT. - 3 UU,1 □ □, Bukidnon Prov., Santo Domingo

(7.782° N, 125.397° E), 560 m a.s.l., forest remnant along brook, near ground, 8-9 Feb. 2014 (B.A.

Huber), ZFMK (Ar 13943); 1 □, 6 juvs, in pure ethanol, same data, ZFMK (Phi 285). - 2 □ Ejj Bukidnon

European Journal of Taxonomy 162: 1-78 (2015)

Prov., near Santo Domingo, Penolohan (7.769° N, 125.420° E), 640 m a.s.l., forest above Salug River,

near ground, 8 Feb. 2014 (B.A. Huber) ZFMK (Ar 13944); 2 □ □ in pure ethanol, same data, ZFMK (Phi

277). - 5 □ □, 15 □ □, 3 juvs, Misamis Occidental Prov., Iligan, NPC Nature’s Park near Cristina Falls

(8.186° N, 124.192° E), 90 m a.s.l., near ground, 17 Feb. 2014 (B.A. Huber), ZFMK (Ar 13945-46); 3

□ 2 juvs, in pure ethanol, same data, ZFMK (Phi 244).

PHIFIPPINES, Camiguin Isl.: 13 □ □, 11 □ 0*1 juv., Katibawasan Falls (9.215° N, 124.720° E), 300 m

a.s.l., near ground, 19 Feb. 2014 (B.A. Huber, PN. Banaag), ZFMK (Ar 13947-48); 1 □, in pure ethanol,

same data, ZFMK (Phi 236). - 9 12 20 juvs, from Katibawasan (9.213° N, 124.718° E), 5-6

s -f

Figs 51-56. Five specimens. Aetana kiukoki group. 51-53. A. kmkoki Huber, sp. nov., □ from Baganihan

(51), □ and □ from Santo Domingo (52-53), Mindanao. 54. A. paragua Huber, sp. nov., □ from Dinagat

Island, Mindanao. 55-56. A. loboc Huber, sp. nov., □ and □ with eggsac from near Foboc, Bohol.

HUBER B.A. et al ., Revision of Aetana spiders

g«P

Figs 57-61. Aetana kiukoki Huber, sp. nov. 57-58. Left male palp, prolateral and retrolateral views.

59. Male prosoma, oblique frontal view. 60-61. Cleared female genitalia, ventral and dorsal views, b =

genital bulb; e = embolus; p = procursus. Scale lines: 0.5 mm.

European Journal of Taxonomy 162: 1-78 (2015)

Figs 62-69. Aetana kiukoki Huber, sp. nov. 62-63. Male prosoma, oblique frontal and frontal views.

64. Female prosoma, frontal view. 65. Male clypeus and chelicerae, oblique frontal view. 66. Male

clypeus modification, frontal view. 67. Epigynum, ventral view. 68. Male ALS and PMS. 69. Comb-hair

on female tarsus 4. cl = clypeus; lea = lateral cheliceral apophysis. Scale lines: 62-64 = 400 pm; 65, 67 =

200 pm; 66 = 100 pm; 68-69 = 20 pm.

HUBER B.A. et al ., Revision of Aetana spiders

May 2014 (E.R Mondejar), MSU-IIT. - 8 □ 6 □ 1 juv., Mt. Hibok Hibok (9.196° N, 124.692° E),

600 m a.s.l., near ground, 18 Feb. 2014 (B.A. Huber, RN. Banaag), ZFMK (Ar 13949-50); 2 □ □, 1 juv.,

in pure ethanol, same data, ZFMK (Phi 240).

Assigned tentatively (no male available)

PHILIPPINES, Mindanao Isl., Davao Oriental: 1 □, from Mount Hamiguitan WS (access San Isidro),

‘site E (6.720° N, 126.172° E), 490 m a.s.l., 9 Feb. 2015 (M.A. Responte), ZFMK (Ar 13951); 7 juvs,

from same locality, ‘site 3’ (6.732° N, 126.179° E), 1250 m a.s.l., 11 Feb. 2015 (M.A. Responte), MSU-

IIT.

Description

Male (holotype)

Measurements. Total body length 3.0, carapace width 1.2. Leg 1: 34.2 (7.9 + 0.5 + 8.0 + 14.8 + 3.0),

tibia 2: 5.0, tibia 3: 3.5, tibia 4: 5.1; tibia 1 L/d: 72. Distance PME-PME 355 pm, diameter PME

150x120 pm, distance PME-ALE ~35 pm; AME absent.

Color. Carapace ochre-yellow with narrow lateral marginal bands and wide dark brown median band

including ocular area and clypeus; sternum with pair of wide brown bands converging posteriorly; legs

greenish ochre with slightly darker rings on femora (subdistally, with light tip), and tibiae (proximally

and subdistally, the latter followed by light tip); abdomen ochre-gray, dorsally and laterally covered

with many black and white marks, ventrally with dark band behind gonopore and very indistinct mark

in front of spinnerets.

Body. Habitus as in Figs 51-52; ocular area raised, each triad on additional short stalk directed toward

lateral (Figs 59, 62-63), without median process; carapace with very shallow median furrow in anterior

part only (Fig. 63); clypeus with large distinctive process, strongly protruding in upper part, with two

pairs of rounded apophyses in distal part, densely covered with small scales (Figs 65, 66); sternum wider

than long (0.75/0.60), unmodified. ALS and PMS as in Fig. 68.

Chelicerae. As in Fig. 59, with pair of dark lateral apophyses proximally and pair of weakly sclerotized

humps laterally, without modified hairs; without stridulatory ridges.

Palps. As in Figs 57-58; coxa unmodified; trochanter with short retrolatero-ventral apophysis; femur

with strong retrolateral apophysis distally curved toward ventral; patella large; tibia small, dorsal

trichobothrium in very proximal position, retrolateral trichobothrium in very distal position; tarsus with

long procursus, distally complex, apparently with two hinged structures; bulb with large embolus and

smaller, semitransparent, pointed process.

Legs. Without spines; with curved hairs on metatarsi 1-3 (few curved hairs also on tibiae 1-2); 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 distinct.

Male (variation)

Tibia 1 in 48 other males: 7.0-9.8 (mean: 8.2). Ventral mark behind gonopore in most males slightly

asymmetrical.

Female

In general similar to male (Fig. 53) but clypeus unmodified; eye triads much closer together (distance

PME-PME 155 pm), not on stalks (Fig. 64); without stridulatory apparatus between carapace and

abdomen. Tibia 1 in 56 females: 4.9-7.5 (mean: 6.1); dark and light rings on legs often more distinct

than in males. Tarsus 4 comb-hairs with very dense tines (Fig. 69). Epigynum in anterior part weakly

European Journal of Taxonomy 162: 1-78 (2015)

sclerotized, internal structures visible through cuticle, with flat, tongue-shaped scape (Figs 60, 87),

without membranous pockets behind epigynum. Internal genitalia as in Figs 61 and 89; pore plates close

together; without sclerotized internal pockets; with distinct transversal sclerotized ridges provided with

many small teeth. The single female from Mt. Hamiguitan cannot be unambiguously assigned to this

species rather than to A. paragua and is therefore assigned tentatively.

Natural history

The spiders were found in domed sheet webs close to the ground, usually in well protected dark spaces

among and under large rocks and logs in forests. Males and females were sometimes found together

in one web. At Baganihan, some specimens (especially juveniles) were found among mosses in deep

furrows of trees up to 1.5 m above the ground. At both localities on Camiguin Island, the spiders were

observed to be much more agile and quick when disturbed than at other localities.

Distribution

Known from numerous localities on Mindanao Island and from Camiguin Island (Fig. 4).

Aetana paragua Huber, sp. nov.

um:lsid:zoobank.org:act:9E9FEDE8-BEB5-46D4-929C-6C2259AA7FDB

Figs 54, 70-74, 90-92

Diagnosis

Distinguished from closest known relatives (A. kiukoki Huber, sp. nov., A. loboc Huber, sp. nov.,

A. pasambai Huber, sp. nov.) by distinctive modification of male clypeus (Fig. 72; similar only in

A. kiukoki Huber, sp. nov.), and tongue-shaped posterior projection of epigynum (Fig. 73; very similar

to A. kiukoki Huber, sp. nov.; much shorter in A. loboc Huber, sp. nov.; female of A. pasambai Huber,

sp. nov. unknown). Distinguished from A. kiukoki Huber, sp. nov. also by shorter male eye stalks

(Fig. 72), weakly curved apophysis on male palpal femur (Fig. 71), different shape of distal procursus

elements (Fig. 71), and pore plates wider apart (Fig. 74). Distinguished from A. loboc Huber, sp. nov.

and A. pasambai Huber, sp. nov. also by modification of male palpal femur (only one large retrolateral

process) and absence of median process in male ocular area.

Etymology

Named for the type locality; noun in apposition.

Material examined

Holotype

PHITIPPINES: □, Dinagat Isl., near Libjo, Paragua Forest, ‘site V (10.222° N, 125.553° E), 130 m

a.s.l., forest at brook, near ground, 20 Feb. 2014 (B.A. Huber), ZFMK (Ar 13952).

Other material

PHILIPPINES: Dinagat Isl., 2 □□, 4 □□, same data as holotype, ZFMK (Ar 13953); 1 □, in pure

ethanol, same data, ZFMK (Phi 230). - 1 1 juv., Paragua Forest, ‘site 2’ (10.241° N, 125.545° E),

240 m a.s.l., forest along brook, near ground, 20 Feb. 2014 (B.A. Huber), ZFMK (Ar 13954).

Description

Male (holotype)

Measurements. Total body length 3.0, carapace width 1.2. Leg 1: 35.2 (8.0 + 0.5 + 8.2 + 15.5 + 3.0),

tibia 2: 4.9, tibia 3: 3.4, tibia 4: 4.9; tibia 1 L/d: 67. Distance PME-PME 230 pm, diameter PME

135x110 pm, distance PME-ALE ~35 pm; AME absent.

HUBER B.A. et al ., Revision of Aetana spiders

Color. Carapace ochre-yellow with narrow lateral marginal bands and wide dark brown median band

including ocular area and clypeus; sternum with pair of wide brown bands converging posteriorly, labium

proximally dark brown; legs greenish ochre with barely visible darker rings on femora (subdistally, with

light tip), and tibiae (proximally and subdistally, the latter followed by light tip); abdomen ochre-gray,

dorsally and laterally covered with many black marks, with few indistinct white marks, ventrally with

dark band behind gonopore and very indistinct mark in front of spinnerets.

Figs 70-74. Aetana paragua Huber, sp. nov. 70-71. Left male palp, prolateral and retrolateral views.

72. Male prosoma, frontal view. 73-74. Cleared female genitalia, ventral and dorsal views (arrows point

at serrated ridges), rt = retrolateral trichobothrium. Scale lines: 0.5 mm.

European Journal of Taxonomy 162: 1-78 (2015)

Body. Habitus as in A. kiukoki Huber, sp. nov. (cf Figs 51-52); ocular area raised, each triad on additional

hump directed toward lateral, without median process; carapace with very shallow median furrow in

anterior part only; clypeus with large distinctive process (Fig. 72), strongly protruding in upper part, with

two pairs of rounded apophyses set with small scales; sternum wider than long (0.80/0.55), unmodified.

Chelicerae. As in Fig. 72, with pair of dark lateral apophyses proximally, barely visible lateral humps

more distally; without modified hairs; without stridulatory ridges.

Palps. As in Figs 70-71; very similar to A. kiukoki Huber, sp. nov., but retrolateral apophysis of femur

less strongly curved, distal elements of procursus different, and semitransparent pointed process on bulb

smaller.

Legs. Without spines; with curved hairs on metatarsi 1-3 (few curved hairs also on tibiae 1-2); 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 distinct.

Male (variation)

Tibia 1 in 2 other males: 8.2, 8.3.

Female

In general similar to male but clypeus unmodified; eye triads much closer together (distance PME-PME

125 pm); without stridulatory apparatus between carapace and abdomen. Tibia 1 in 4 females: 5.5, 5.6,

5.6, 5.8; dark and light rings on legs often more distinct than in males. Epigynum and internal genitalia

very similar to those of A. kiukoki Huber, sp. nov., but tongue-shaped posterior projection slightly shorter

and wider (Figs 73, 90), and pore plates wider apart (Fig. 74).

Natural history

The spiders were found in domed webs very close to the ground. At the same locality, A. libjo Huber, sp.

nov. occurred higher among the vegetation.

Distribution

Known from two neighboring localities on Dinagat Island only (Fig. 4).

Aetana loboc Huber, sp. nov.

um:lsid:zoobank.org:act:CAD63F67-D442-43BF-ACBC-062F4B3DCA0A

Figs 55-56, 75-86, 93-95

Diagnosis

Distinguished from closest known relative (A. pasambai Huber, sp. nov.) by distinctive modification of

male clypeus (Fig. 77; longer and narrower), by median process between eye stalks larger and in more

dorsal position (Fig. 77), by longer male palpal tibia and procursus (Fig. 76), and by stronger ventral

apophysis on male palpal femur (female of A. pasambai Huber, sp. nov. unknown); from other close

relatives (A. paragua Huber, sp. nov.; A. kiukoki Huber, sp. nov.) by modifications of male palpal femur

(two processes instead of one), presence of median process between eye stalks, and very short tongue¬

shaped posterior projection of epigynum (Fig. 78).

Etymology

Named for the type locality; noun in apposition.

HUBER B.A. et al ., Revision of Aetana spiders

Figs 75-79. Aetana loboc Huber, sp. nov. 75-76. Left male palp, prolateral and retrolateral views.

77. Male prosoma, frontal view. 78-79. Cleared female genitalia, ventral and dorsal views. Scale lines:

0.5 mm

European Journal of Taxonomy 162: 1-78 (2015)

Figs 80-86. Aetana loboc Huber, sp. nov. 80. Female prosoma, frontal view. 81. Female ALS. 82.

Comb-hair on female tarsus 4. 83. Detail of female tarsus 1. 84. Detail of female tarsus 4. 85. Epigynum.

86 . Detail of preceding. Scale lines: 80, 85 = 200 pm; 81 = 10 pm; 82 = 8 pm; 83-84 = 40 pm; 86 =

100 pm.

HUBER B.A. et al ., Revision of Aetana spiders

Material examined

Holotype

PHILIPPINES: □, Bohol Isl., near Loboc, above Loboc River (9.651° N, 124.022° E), ~50 m a.s.l.,

ravine in degraded forest, 5 Mar. 2014 (B.A. Huber), ZFMK (Ar 13955).

Figs 87-95. Aetana kiukoki group. Female genitalia; untreated in ventral view, cleared in ventral and dorsal

views. — 87-89. A. kiukoki Huber, sp. nov. — 90-92. A. paragua Huber, sp. nov. — 93-95. A. loboc

Huber, sp. nov.

European Journal of Taxonomy 162: 1-78 (2015)

Other material

PHILIPPINES: Bohol Isl., 4 DfjJ 13□ □, 1 juv., same data as holotype, ZFMK (Ar 13956-57); 3 □ 3

juvs, in pure ethanol, same data as holotype, ZFMK (Phi 201). - 3 0 0,3 □ □, at Loboc River (9.651° N,

124.022° E), near ground, 20 m a.s.l., 4 Mar. 2014 (B.A. Huber), ZFMK (Ar 13958). - 1 □, 1 □, Rajah

Sikatuna (Magsaysay Park) (9.705° N, 124.123° E), 430 m a.s.l., forest, 6 Mar. 2014 (B.A. Huber),

ZFMK (Ar 13959).

Description

Male (holotype)

Measurements. Total body length 3.1, carapace width 1.2. Leg 1: 32.4 (7.5 + 0.5 + 7.6 + 14.1 + 2.7),

tibia 2: 4.5, tibia 3: 3.2, tibia 4: 4.9; tibia 1 L/d: 73. Distance PME-PME 340 pm, diameter PME

135x120 pm, distance PME-ALE ~35 pm; AME absent.

Color. Carapace ochre-yellow with narrow lateral marginal bands and wide dark brown median band

including ocular area and clypeus; sternum with pair of wide brown bands converging posteriorly,

labium proximally not dark brown; legs ochre to light brown, with indistinct darker rings on femora

(subdistally, with light tip), and tibiae (proximally and subdistally, the latter followed by light tip);

abdomen ochre-gray, dorsally and laterally covered with many black marks, with few indistinct white

marks, ventrally with small spot behind gonopore and very indistinct mark in front of spinnerets.

Body. Habitus as in Fig. 55; ocular area raised, each triad on short stalk directed toward lateral, with

distinctive median process (Fig. 77); carapace with very shallow median furrow in anterior part only;

clypeus with large distinctive process (Fig. 77), strongly protruding in upper part, with distinctive pair

of distal apophyses; sternum wider than long (0.75/0.60), unmodified.

Chelicerae. As in Fig. 77, with pair of dark lateral apophyses proximally, without lateral humps more

distally; without modified hairs; without stridulatory ridges.

Palps. As in Figs 75-76; coxa unmodified; trochanter with wide ventral apophysis; femur large,

with strong ventral and slender retrolateral apophyses; patella very large; tibia small but long, dorsal

trichobothrium in very proximal position, retrolateral trichobothrium in very distal position; tarsus with

very long procursus, distally complex, apparently with two hinged structures; bulb with large embolus

and small semitransparent pointed process.

Legs. Without spines; with curved hairs on metatarsi 1-3 (few curved hairs also on tibiae 1-2); few

vertical hairs; retrolateral trichobothrium on tibia 1 at 2%; prolateral trichobothrium absent on tibia 1,

present on other tibiae; tarsus 1 with ~25 pseudosegments, only distally distinct.

Male (variation)

Tibia 1 in 7 other males: 6.5-7.7 (mean 7.1).

Female

In general similar to male but clypeus unmodified; eye triads much closer together (distance PME-

PME 135 pm); without stridulatory apparatus between carapace and abdomen; without ventral dark

band on abdomen. Tibia 1 in 17 females: 5.0-5.7 (mean: 5.3); dark and light rings on legs often more

distinct than in males. Epigynum in anterior part weakly sclerotized, internal structures partly visible

through cuticle, with short posterior projection with submarginal transversal groove (Figs 78, 85-86,

93), without membranous pockets behind epigynum; internal genitalia as in Figs 79 and 95; without

sclerotized internal pockets; with distinct transversal sclerotized ridges provided with many small teeth.

HUBER B.A. et al ., Revision of Aetana spiders

Natural history

The spiders were found in domed webs close to the ground, hidden under rocks or logs.

Distribution

Known from two neighboring localities on Bohol Island only (Fig. 4).

Aetanapasambai Huber, sp. nov.

urn:lsid:zoobank.org:act:041DF955-4589-4503-AF5D-lEC15C107E8E

Figs 96-98

Diagnosis

Distinguished from closest known relative (A. I oboe Huber, sp. nov.) by distinctive modification of male

clypeus (Fig. 96; shorter and wider), by median process between eye stalks smaller and in more frontal

position (Fig. 96), by shorter male palpal tibia and procursus (Fig. 98), and by smaller ventral apophysis

on male palpal femur (female of A. pasambai Huber, sp. nov. unknown); from other close relatives (A.

paragua sp. nov., A. kiukoki Huber, sp. nov.) by modifications of male palpal femur (two processes

instead of one), and by presence of median process between eye stalks.

Etymology

Named for Philippine-born cellist Wilfredo Pasamba.

Material examined

Holotype

PHIFIPPINES: □, Negros Isl., Negros Oriental Prov., Casaroro Falls (9.281° N, 123.208° E), 550 m

a.s.l., forest along river below waterfall, 10 Mar. 2014 (B.A. Huber), ZFMK (Ar 13960).

Other material

None.

Description

Male (holotype)

Measurements. Total body length 3.3, carapace width 1.1. Teg 1: 32.7 (7.7 + 0.5 + 7.7 + 14.1 + 2.7),

tibia 2: 4.7, tibia 3: 3.7, tibia 4: 5.0; tibia 1 F/d: 69. Distance PME-PME 385 pm, diameter PME

135x120 pm, distance PME-AFE ~35 pm; AME absent.

Color. Carapace ochre-yellow with narrow lateral marginal bands and wide dark brown median band

including ocular area and clypeus; sternum with pair of wide brown bands converging posteriorly,

with darker triangular mark posteriorly, labium proximally not dark brown; legs greenish ochre, with

indistinct darker rings on femora (subdistally, with light tip), and tibiae (proximally and subdistally, the

latter followed by light tip); abdomen ochre-gray, dorsally and laterally covered with many black marks,

with some white marks, ventrally with dark band behind gonopore and very indistinct mark in front of

spinnerets.

Body. Habitus as in A. loboc Huber, sp. nov. ( cf. Fig. 55); ocular area raised, each triad on short stalk

directed toward lateral, with small median process in frontal position (Fig. 96); carapace with very

shallow median furrow in anterior part only; clypeus with large distinctive process (Fig. 96), protruding

in upper part, with distinctive pair of distal apophyses; sternum wider than long (0.80/0.55), unmodified.

European Journal of Taxonomy 162: 1-78 (2015)

Chelicerae. As in Fig. 96, with pair of dark lateral apophyses proximally, without lateral humps more

distally; without modified hairs; without stridulatory ridges.

Palps. As in Figs 97-98; very similar to A. loboc Huber, sp. nov., but femur apophyses of different sizes

and positions, tibia and procursus shorter.

Figs 96-98. Aetanapasambai Huber, sp. nov. 96. Male prosoma, oblique frontal view. 97-98. Left male

palp, prolateral and retrolateral views. Scale lines: 0.5 mm.

HUBER B.A. et al ., Revision of Aetana spiders

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

trichobothrium on tibia 1 at 2%; prolateral trichobothrium absent on tibia 1, present on other tibiae;

tarsus 1 with ~25 pseudosegments, only distally distinct.

Female

Unknown

Natural history

The specimen was found in a small hole in the ground.

Distribution

Known from type locality on Negros Island only (Fig. 4).

Aetana kinabalu Huber, 2005

Figs 99-103, 111-112, 163-165

Aetana kinabalu Huber, 2005a: 75-76, figs 108-109, 121-123 (XI).

Note

The original description was based on two males. Here we present data on new material from the type

locality, a description of the female, and an emended diagnosis to account for the newly described

congeners.

Diagnosis

Distinguished from closest known relative (A. lambir Huber, sp. nov.) by shape of prolatero-ventral

apophysis of male palpal femur (pointed tip and subdistal branch of approximately same length; cf

Huber 2005a: fig. 121); also by presence of prolateral apophysis on femur, by details of procursus

(shapes of sclerites on complex distal part; cf. Huber 2005a: figs 121-122), and by female genitalia (pair

of internal pockets; position of pore plates; Figs 111-112).

New material examined

MALAYSIA-BORNEO, Sabah: 1 □, 1 □, 1 juv., Mt. Kinabalu, forest along Silau Silau Trail (6.010-

6.017° N, 116.537-116.543° E), 1550-1650 m a.s.l., domed webs among vegetation, 6 Aug. 2014 (B.A.

Huber, S B. Huber), ZFMK (Ar 13961); 2UU, 1 juv., in pure ethanol, same data, ZFMK (Bor 211).

-ID, Kinabalu N.R, 1550 m a.s.l., 2-8 Apr. 1998 (C.L. Deeleman-Reinhold, R Zborowski), RMNH.

-2 □ □, 5 , 3 juvs, Crocker Range between Kota Kinabalu and Tambuan, S-slope, forest along river

(5.783° N, 116.338-116.340° E), 1430-1480 m a.s.l., domed webs among vegetation, 3 Aug. 2014 (B.A.

Huber, S.B. Huber), ZFMK (Ar 13962); 4 juvs, in pure ethanol, same data, ZFMK (Bor 171). - 2 □

3 □ in very poor condition, Tawau (4.406° N, 117.892° E), 6 Sep. 2009 (A. Floren), RMNH.

MALAYSIA-BORNEO, Sarawak: 2 □□, 6 □□, 3 juvs, Gunung Mulu N.R, forest near Deer Cave

(4.027° N, 114.818° E), 60 m a.s.l., 23-24 July 2014 (B.A. Huber, S.B. Huber), ZFMK (Ar 13963-64);

same data, 3 □ □, 5 juvs, in pure ethanol, ZFMK (Bor 182). - 5 □ □, 2 , 4 juvs, Bario, forest along

river W of town (3.736° N, 115.437-115.443° E), 1150-1250 m a.s.l., domed webs among vegetation,

30 July 2014 (B.A. Huber, S.B. Huber), ZFMK (Ar 13965). - 1 □, 1 juv., Bario, forest along river N

of town (3.765-3.771° N, 115.444-115.448° E), 1170-1250 m a.s.l., domed webs among vegetation,

29 July 2014 (B.A. Huber, S.B. Huber), ZFMK (Ar 13966); 9 juvs, in pure ethanol, same data, ZFMK

(Bor 233).

European Journal of Taxonomy 162: 1-78 (2015)

Description (amendments to Huber 2005a)

Male

Thoracic furrow absent (contra Huber 2005a); clypeus slightly protruding (more than in female). Tibia 2

slightly longer than tibia 4 (e.g., 5.1/4.9; 4.9/4.8); curved hairs on metatarsi 2; tibia 1 in 7 males: 7.6-8.8

(mean: 8.3).

Female

Eye triads much closer together than in male (distance PME-PME -235 pm vs. 375 pm), not on stalks.

Tibia 1 in 15 females: 5.9-7.3 (mean 6.4). Epigynum simple plate, darker laterally anteriorly; anterior

Figs 99-104. Live specimens. Aetana kinabalu group. — 99-103. A. kinabalu Huber, 2005. □ and □

with eggsac from Gunung Mulu, Sarawak (99, 100); □ with eggsac and Q from Crocker Range, Sabah

(101, 102); and □ from Mt. Kinabalu, Sabah (103). —104. A. gaya Huber, sp. nov., □ from Gaya Island,

Sabah.

HUBER B.A. et al ., Revision of Aetana spiders

internal arc and internal sclerotized pockets visible through cuticle (Figs 111, 163-164). Internal genitalia

as in Figs 112 and 165, with pair of sclerotized pockets and more lateral pair of membranous pockets.

Variation

In males from Gunung Mulu, the tiny cone-shaped process on the genital bulb (cf Huber 2005a: figs

121 and 122) is absent. Males from Bario with small additional process at basis of prolatero-ventral

apophysis of femur.

Natural history

Webs were in most cases made of two sheets: an upper sheet in which the spider hung, and a lower sheet

that had to be removed in order to catch the spider. At most localities, webs were found high among the

Figs 105-110. Five specimens. Aetana kinabalu group. — 105-106. A. lambir Huber, sp. nov., FI and □

with eggsac from Fambir, Sarawak. — 107-108. A. poring Huber, sp. nov., □ □ from Mt. Kinabalu, Sabah.

— 109- 110. A. indah Huber, sp. nov., adult and penultimate instar, □□ from Crocker Range, Sabah.

European Journal of Taxonomy 162: 1-78 (2015)

vegetation, even in the sunlight. At Gunung Mulu, webs were found among mosses and low vegetation

on perpendicular rock surfaces about 2 m above the ground. At several localities, A. kinabalu was

sympatric with a ground-dwelling congener: with A. poring Huber, sp. nov. at Mt. Kinabalu and Poring;

with A. indah Huber, sp. nov. at Crocker Range.

Distribution

Widely distributed in Sabah and eastern Sarawak (Fig. 5).

Aetana lambir Huber, sp. nov.

um:lsid:zoobank.org:act:EFFAC871-4657-461F-9875-807E767DB6B8

Figs 105-106, 113-125, 166-168

Diagnosis

Distinguished from closest known relative (A. kinabalu) by shape of prolatero-ventral apophysis of

male palpal femur (Fig. 113; pointed tip much longer than subdistal branch); also by details of procursus

(shapes of sclerites on complex distal part), by absence of prolateral apophysis on femur, and by female

genitalia (no internal sclerotized pockets; pore plates wider apart; Figs 116-117).

Etymology

Named for the type locality; noun in apposition.

Material examined

Holotype

MALAYSIA-BORNEO: □, Sarawak, Lambir Hills N.P. (4.198^.207° N, 114.034-114.045° E), 60-

150 m a.s.l., in domed webs among low vegetation, 22 July 2014 (B.A. Huber, S.B. Huber), ZFMK (Ar

13967).

Other material

MALAYSIA-BORNEO, Sarawak: 9 □□, 11 □□, same data as holotype, ZFMK (8 □□, 10 Ar

13968-69) and SMK (1 □, 1 □); 1 □, 3^J^4juvs, in pure ethanol, same data, ZFMK (Bor 201).-3 BUy

4 Cjjrj, Niah Cave N.P, forest near headquarters (3.820° N, 113.763° E), 40 m a.s.l., night collecting,

domed webs among vegetation, 28 July 2014 (B.A. Huber, S.B. Huber), ZFMK (Ar 13970). - 6 □ 3

,Niah Cave N.P, forest near cave (3.814° N, 113.771° E), 40 m a.s.l., among low vegetation, 28 July

Figs 111-112. Aetana kinabalu Huber, 2005. Cleared female genitalia, ventral and dorsal views (arrow

in Fig. 112 points at membranous pocket), ip = internal sclerotized pocket. Scale line: 0.5 mm.

HUBER B.A. et al ., Revision of Aetana spiders

Figs 113-117. Aetana lambir Huber, sp. nov. 113-114. Left male palp, prolateral and retrolateral views.

115. Male chelicerae, frontal view. 116-117. Cleared female genitalia, ventral and dorsal views (arrow

points at membranous pocket), b = genital bulb; e = embolus; rp = retrolatero-ventral process; rt =

retrolateral trichobothrium; vl = ventral lamina. Scale lines: 113-114 = 0.5 mm ; 115-117 = 0.3 mm

European Journal of Taxonomy 162: 1-78 (2015)

Figs 118-125. Aetana lambir Huber, sp. nov. 118-119. Male and female prosomata, frontal views. 120.

Male palpal tarsal organ. 121. Left procursus, prolateral view. 122. Detail of preceding. 123. Right

procursus (and femur apophysis), retrolateral view. 124. Female ALS. 125. Epigynum. f = femur; vl =

ventral lamina. Scale lines: 118-119, 125 = 200 pm; 120 = 8 pm; 121 =50 pm; 122 = 20 pm; 123 =

60 pm; 124 =10 pm.

HUBER B.A. et al ., Revision of Aetana spiders

2014 (B.A. Huber), ZFMK (Ar 13971). - 3 11 1 juv., Niah Cave N.R, forest along main trail

(3.814-3.821° N, 113.763-113.771° E), 2CM0 m a.s.l., domed webs among vegetation, 27 July 2014

(B.A. Huber, S.B. Huber), ZFMK (Ar 13972-73); 1 11 1 juv., same data, in pure ethanol, ZFMK

(Bor 178).

Description

Male (holotype)

Measurements. Total body length 2.5, carapace width 0.95. Feg 1: 33.1 (7.9 + 0.4 + 7.6 + 14.3 + 2.9),

tibia 2: 4.4, tibia 3: 2.8, tibia 4: 4.2; tibia 1 F/d: 87. Distance PME-PME 340 pm, diameter PME 105

pm, distance PME-AFE 25 pm; AME absent.

Color. Carapace pale ochre with black lateral margins and brown median band including ocular area.

Clypeus mostly pale ochre, at rim brown. Sternum pale ochre, laterally slightly darker. Fegs ochre to

light brown, slightly darker rings on femora (subdistally) and tibiae (proximally and subdistally); tips of

femora and tibiae whitish. Abdomen with dorsal and lateral pattern of black and white marks; ventrally

with brown mark near spinnerets.

Body. Habitus as in Fig. 105; ocular area slightly raised, each triad on short stalk directed toward lateral;

carapace without thoracic furrow; clypeus slightly more protruding than usual; sternum wider than long

(0.60/0.45), unmodified.

Chelicerae. As in Fig. 115, with pair of proximal lateral apophyses and pair of simple distal apophyses

in very lateral position; without modified hairs; without stridulatory ridges.

Palps. As in Figs 113-114, coxa unmodified; trochanter with short, rounded ventral apophysis;

femur with distinct retrolatero-ventral apophysis, long prolatero-ventral apophysis with side branch,

no prolateral process. Procursus complex, ventral transparent lamina proximally wide and slightly

sclerotized; retrolatero-ventral process indistinct, apparently fused to procursus. Bulb simple, with short

and wide embolus.

Fegs. Without spines, with curved hairs on metatarsi 2 only (single dorsal row; proximal half), with few

vertical hairs; retrolateral trichobothrium on tibia 1 at 2.5%; prolateral trichobothrium absent on tibia 1,

present on other tibiae. Tarsus 1 with >20 pseudosegments, only distally fairly distinct.

Male (variation)

Tibia 1 in 19 other males: 6.1-8.5 (mean 7.4); most males with small dark mark at gonopore.

Female

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

less protruding; no curved hairs on metatarsi. Tibia 1 in 26 females: 5.6-6.6 (mean 6.2). Epigynum very

simple plate, wider than long (Fig. 125), anterior internal arc visible through cuticle (Figs 116, 166).

Internal genitalia very simple, as in Figs 117 and 168, without internal sclerotized pockets, with pair of

lateral membranous pockets. AES as in Fig. 124.

Natural history

At the type locality (Fambir), the spiders were found among vegetation close to the ground, while at

Niah they occurred higher among the vegetation, very similar to A. kinabalu , in the same type of two¬

layered web (see above).

European Journal of Taxonomy 162: 1-78 (2015)

Distribution

Known from two localities in eastern Sarawak (Fig. 5).

Aetana gaya Huber, sp. nov.

um:lsid:zoobank.org:act:3FF192B2-3F88-4BBF-A675-CA77FF9A72Fl

Figs 104, 126-144, 169-171

Diagnosis

Distinguished from closest known relatives (A. kinabalu , A. lambir Huber, sp. nov., A. indah Huber,

sp. nov., A. poring Huber, sp. nov.) by shape of prolatero-ventral apophysis of male palpal femur (Fig.

126; single pointed process without side branch); also by apophysis on male palpal trochanter (longer

and more sclerotized than in A. kinabalu and A. lambir Huber, sp. nov.; shorter than in A. indah Huber,

sp. nov. and A. poring Huber, sp. nov.), by shapes of sclerites on procursus (Figs 126-127), and by

female genitalia (Figs 128-129, 169-171; short and wide epigynum with straight posterior margin;

distinctive internal structures).

Etymology

Named for the type locality; noun in apposition.

Material examined

Holotype

MALAYSIA-BORNEO: □, Sabah, Gaya Island, forest along small stream (6.014-6.018° N,

116.020° E), 30-80 m a.s.l., among rocks and tree buttresses, 4 Aug. 2014 (B.A. Huber, S.B. Huber),

ZFMK (Ar 13974).

Other material

MALAYSIA-BORNEO: Sabah, 12 □□, 13 DD^juvs, same data as holotype, ZFMK (11 □□, 12

Ar 13975-76) and SMK (1 □, 1 CL); same data, 7 □ □, 2 juvs, in pure ethanol, ZFMK (Bor 165).

Description

Male (holotype)

Measurements. Total body length 2.5, carapace width 0.9. Leg 1: 32.6 (7.6 + 0.4 + 7.6 + 14.2 + 2.8),

tibia 2: 4.4, tibia 3: 2.8, tibia 4: 4.2; tibia 1 L/d: 96. Distance PME-PME 370 pm, diameter PME 105

pm, distance PME-ALE 25 pm; AME absent.

Color. Carapace pale ochre with black lateral bands and wide brown median band including ocular area.

Clypeus pale ochre with pair of brown marks at rim. Sternum medially ochre, laterally slightly darker.

Legs ochre to light brown, indistinct darker rings on femora (subdistally) and tibiae (proximally and

subdistally); tips of femora and tibiae whitish. Abdomen grey with dorsal and lateral pattern of black and

white marks; ventrally with small brown marks near spinnerets and in genital area.

Body. Habitus as in Fig. 104; ocular area slightly raised, each triad on short stalk directed toward lateral

(Figs 130-131); carapace without thoracic furrow (Fig. 135; only dark line in anterior part); clypeus

slightly more protruding than usual; sternum wider than long (0.75/0.55), unmodified. Gonopore with

four epiandrous spigots (Fig. 143). Spinnerets as in Figs 141-142.

Chelicerae. As in A. lambir Huber, sp. nov. ( cf Fig. 115), with pair of proximal lateral apophyses and

pair of simple distal apophyses in very lateral position; without modified hairs; without stridulatory

ridges.

HUBER B.A. et al ., Revision of Aetana spiders

Palps. As in Figs 126-127, coxa unmodified; trochanter with short ventral apophysis with small teeth

prolaterally and small prolateral branch; femur with rounded retrolatero-ventral apophysis, long pointed

prolatero-ventral apophysis without side branch, with small ventral process. Tarsal organ capsulate (Fig.

134). Procursus complex (Figs 137-140); retrolatero-ventral process with simple tip (Fig. 136). Bulb

simple, with short and wide embolus.

Figs 126-129. Aetana gaya Huber, sp. nov. 126-127. Feft male palp, prolateral and retrolateral views.

128-129. Cleared female genitalia, ventral and dorsal views (arrow points at membranous pocket), b =

genital bulb; e = embolus; rp = retrolatero-ventral process; vl = ventral lamina. Scale lines: 126-127 =

0.5 mm ; 128-129 = 0.3 mm

European Journal of Taxonomy 162: 1-78 (2015)

Figs 130-136. Aetana gaya Huber, sp. nov. 130-131. Male prosoma, frontal and oblique frontal views.

132. Female prosoma, frontal view. 133. Detail of male metatarsus 1. 134. Male palpal tarsal organ.

135. Male carapace and ocular area. 136. Right procursus, retrolateral view, rp = retrolatero-ventral process;

vl = ventral lamina. Scale lines: 130-132, 135 = 200 pm; 133 = 80 pm; 134 = 8 pm; 136 = 100 pm.

HUBER B.A. et al ., Revision of Aetana spiders

Figs 137-144. Aetana gaya Huber, sp. nov. 137. Left procursus, prolateral view. 138-140. Details of

preceding. 141. Male spinnerets. 142. Male ALS. 143. Male gonopore. 144. Epigynum. b = genital bulb;

f= femur; p = procursus. Scale lines: 137 = 100 pm; 138, 141 = 60 pm; 139-140 = 20 pm; 142 = 10 pm;

143 = 30 pm; 144 = 200 pm.

European Journal of Taxonomy 162: 1-78 (2015)

Legs. Without spines, with curved hairs dorsally on metatarsi 1 and 2 only (mostly on proximal half),

with few vertical hairs; retrolateral trichobothrium on tibia 1 at 2.5%; prolateral trichobothrium absent

on tibia 1, present on other tibiae. Tarsus 1 with ~30 pseudosegments, distally fairly distinct.

Male (variation)

Tibia 1 in 10 other males: 7.2-8.1 (mean 7.6).

Female

In general similar to male; eye triads closer together (distance PME-PME 185 pm), not on stalks (Fig.

132); clypeus less protruding; no curved hairs on metatarsi. Tibia 1 in 11 females: 4.9-5.8 (mean 5.3).

Epigynum short and wide plate (Fig. 144), slightly protruding, with pair of distinctive curved darker

marks (Figs 128,169). Internal genitalia as inFigs 129 and 171, with sclerotized structures but apparently

without sclerotized pockets, with pair of lateral membranous pockets.

Natural history

Most specimens were found close to the ground among logs, but some (mostly juveniles) built their

webs in less protected places higher on trees.

Distribution

Known from Gaya Island only (Fig. 5).

Aetanaporing Huber, sp. nov.

um:lsid:zoobank.org:act:2D2D173E-9AFC-4CDF-877F-40C44043ElC5

Figs 107-108, 145-149, 172-174

Diagnosis

Distinguished from closest known relative (A. indah Huber, sp. nov.) by shape of prolatero-ventral

apophysis of male palpal femur (Fig. 145; pointed tip much shorter), by distal cheliceral apophyses

(Fig. 147; more pointed and gradually narrowing), and by female genitalia (Figs 148, 172; pair of dark

lines; posterior rim curved toward posterior); from all other relatives also by strong apophysis prolatero-

proximally on male palpal femur (Fig. 145; present but smaller in A. indah Huber, sp. nov.).

Etymology

Named for the type locality; noun in apposition.

Material examined

Holotype

MALAYSIA-BORNEO: □, Sabah, Mt. Kinabalu, Poring Hot Springs, forest along Kipungit River

(6.049°N, 116.712°E), 450 m a.s.l., near ground, 7 Aug. 2014 (B.A. Huber), ZFMK (Ar 13977).

Other material

MALAYSIA-BORNEO, Sabah: 2 □□, 4 □□, same data as holotype, ZFMK (Ar 13978-79); 1 □, 2

juvs, in pure ethanol, same data, ZFMK (Bor 206). - 4 Ipfi Mt. Kinabalu, forest along Silau Silau Trail

(6.010-6.017° N, 116.537-116.543° E), 1550-1650 m a.s.l., near ground, 6 Aug. 2014 (B.A. Huber,

S.B. Huber), ZFMK (Ar 13980); 2 juvs, in pure ethanol, in ZFMK (Bor 210), same data. - 2 □□, 1

□ , Mt. Kinabalu, forest above Kinabalu Mountain Lodge (6.012-6.014° N, 116.534° E), 1570-1650

m a.s.l., near ground, 5 Aug. 2014 (B.A. Huber, S.B. Huber), ZFMK (Ar 13981); 1 □, 1 juv., in pure

ethanol, same data, ZFMK (Bor 215). - 5 □ □, 2 juvs, Kinabalu N.P., 1550 m a.s.l., 3 June 1979 and 26

July 1980 (C.L. & PR Deeleman), RMNH (2 vials).

HUBER B.A. et al ., Revision of Aetana spiders

Description

Male (holotype)

Measurements. Total body length 2.6, carapace width 1.1. Leg 1: 26.9 (6.4 + 0.4 + 6.4 + 11.2 + 2.5),

tibia 2: 3.8, tibia 3: 2.7, tibia 4: 4.0; tibia 1 L/d: 63. Distance PME-PME 455 pm, diameter PME 105

pm, distance PME-ALE 35 pm; AME absent.

Color. Carapace pale ochre with black lateral bands and wide brown median band including ocular area.

Clypeus pale ochre with pair of brown marks at rim. Sternum medially ochre, laterally slightly darker.

Legs ochre to light brown, indistinct darker rings on femora (subdistally) and tibiae (proximally and

Figs 145-149. Aetana poring Huber, sp. nov. 145-146. Left male palp, prolateral and retrolateral views

(arrow points at prolateral apophysis near hinge). 147. Male chelicerae, frontal view. 148-149. Cleared

female genitalia, ventral and dorsal views (arrow points at membranous pocket). Scale lines: 145-146,

148-149 = 0.5 mm; 147 = 0.3 mm

European Journal of Taxonomy 162: 1-78 (2015)

subdistally); tips of femora and tibiae whitish. Abdomen grey with dorsal and lateral pattern of black and

indistinct white marks; ventrally with small brown mark near spinnerets and in genital area.

Body. Habitus as in Figs 107-108; ocular area slightly raised, each triad on short stalk directed toward

lateral; carapace without thoracic furrow (only dark line in anterior part); clypeus slightly more protruding

than usual; sternum wider than long (0.75/0.55), unmodified.

Chelicerae. As in Fig. 147, with pair of proximal lateral apophyses and pair of simple distal apophyses

in very lateral position; without modified hairs; without stridulatory ridges.

Palps. As in Figs 145-146; coxa unmodified; trochanter with slender ventral apophysis with very small

teeth prolaterally; femur with rounded retrolatero-ventral apophysis, long prolatero-ventral apophysis

with side branch, with two prolateral processes, one of them very close to trochanter. Procursus complex;

retrolatero-ventral process with bifid tip. Bulb simple, with short and wide embolus.

Legs. Without spines, with curved hairs dorsally on metatarsi 1 and 2 only (mostly on proximal half),

with 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, distally fairly distinct.

Male (variation)

Tibia 1 in 4 other males: 6.1, 6.9, 7.1, 7.5; most males with distinct white marks on abdomen.

Female

In general similar to male; triads closer together (distance PME-PME 200 pm), not on stalks; clypeus

less protruding; no curved hairs on metatarsi; abdomen with continuous ventral dark band between

epigynum and spinnerets. Tibia 1 in 12 females: 4.2-5.8 (mean 5.1). Epigynum large brown plate (Figs

148, 172), slightly protruding, with distinctive lighter median area bordered by dark lines (parallel or

converging anteriorly), with internal sclerites visible through cuticle. Internal genitalia as in Figs 149

and 174, apparently without sclerotized pockets, with pair of lateral membranous pockets.

Natural history

The spiders were found close to the ground in small holes and cavities. They barely reacted to disturbance

and were very easy to take from their webs. They share the locality with A. kinabalu , which lives higher

among the vegetation.

Distribution

Known from two localities in Mt. Kinabalu area only (Fig. 5).

Aetana indah Huber, sp. nov.

urn:lsid:zoobank.org:act:12E0B867-3A77-4B63-92FF-97DB7302137C

Figs 109-110, 150-162, 175-177

Diagnosis

Distinguished from closest known relative (A. poring Huber, sp. nov.) by shape of prolatero-ventral

apophysis of male palpal femur (Fig. 150; pointed tip much longer), by distal cheliceral apophyses (Fig.

152; wide and distally rounded), and by female genitalia (Figs 153, 175; without pair of dark lines;

posterior rim curved toward anterior); from all other congeners also by long apophysis on male palpal

trochanter (in A. poring Fluber, sp. nov. present but more slender and shorter).

HUBER B.A. et al ., Revision of Aetana spiders

Etymology

The species name is the Malay word for ‘beautiful’; used here as noun in apposition.

Material examined

Holotype

MALAYSIA-BORNEO: □, Sabah, Crocker Range between Kota Kinabalu and Tambuan, S-slope,

forest along river (5.783° N, 116.338-116.340° E), 1430-1480 m a.s.l., near ground, 3 Aug. 2014 (B.A.

Huber, S.B. Huber), ZFMK (Ar 13982).

Other material

MALAYSIA-BORNEO, Sabah: 3 □□, 5 □□, 3 juvs, same data as holotype, ZFMK (Ar 13983-84);

1 □, 2 juvs, in pure ethanol, same data, ZFMK (Bor 170). - 1 □, 3 □ □, Crocker Range between Kota

Kinabalu and Tambuan, N-slope, forest along river (5.834° N, 116.336° E), 1600 m a.s.l., near ground,

3 Aug. 2014 (B.A. Huber, S.B. Huber), ZFMK (Ar 13985); 2 Dj|L in pure ethanol, same data, ZFMK

(Bor 168).

Description

Male (holotype)

Measurements. Total body length 2.8, carapace width 1.2. Leg 1: 36.8 (8.8 + 0.4 + 8.8 + 15.3 + 3.5),

tibia 2: 5.2, tibia 3: 3.5, tibia 4: 5.3; tibia 1 L/d: 84. Distance PME-PME 500 pm, diameter PME 115

pm, distance PME-ALE 35 pm; AME absent.

Color. Carapace pale ochre with black lateral bands and wide brown median band including ocular area.

Clypeus pale ochre with indistinct pair of brown marks at rim. Sternum medially ochre, laterally slightly

darker. Legs ochre to light brown, indistinct darker rings on femora (subdistally) and tibiae (proximally

and subdistally); tips of femora and tibiae whitish. Abdomen grey with dorsal and lateral pattern of black

and indistinct white marks; ventrally with small brown mark near spinnerets and in genital area.

Body. Habitus as in Fig. 109; ocular area slightly raised, each triad on short stalk directed toward lateral;

carapace without thoracic furrow (only dark line in anterior part); clypeus slightly more protruding than

usual; sternum wider than long (0.65/0.50), unmodified.

Chelicerae. As in Figs 152 and 161, with pair of proximal lateral apophyses and distinctive pair of wide

distal apophyses in very lateral position, with additional pair of small processes on frontal side of distal

apophyses (Fig. 160); without modified hairs; without stridulatory ridges.

Palps. As in Figs 150-151, coxa unmodified; trochanter with long ventral apophysis with small teeth

prolaterally (Fig. 158); femur with rounded retrolatero-ventral apophysis, very long prolatero-ventral

apophysis with two side branches, with small prolateral apophysis proximally close to trochanter.

Procursus complex (Figs 155-158); retrolatero-ventral process with bifid tip. Bulb simple, with short

and wide embolus (Fig. 159), weakly sclerotized.

Legs. Without spines, with curved hairs dorsally on metatarsi 1 and 2 only (mostly on proximal half),

with few vertical hairs; retrolateral trichobothrium on tibia 1 at 2.5%; prolateral trichobothrium absent

on tibia 1, present on other tibiae. Tarsus 1 with ~35 pseudosegments, fairly distinct.

Male (variation)

Tibia 1 in 4 other males: 8.7, 8.8, 8.9, 9.0.

European Journal of Taxonomy 162: 1-78 (2015)

152

Female

In general similar to male; triads closer together (distance PME-PME 210 jam), not on stalks; clypeus

less protruding; no curved hairs on metatarsi; abdomen with continuous ventral dark band between

epigynum and spinnerets. Tibia 1 in 8 females: 6.3-7.2 (mean 6.7). Epigynum large brown plate, slightly

protruding, with internal pockets and other sclerites visible through cuticle (Figs 153, 175). Internal

genitalia as in Figs 154 and 177, with pair of membranous pockets near posterior margin.

Figs 150-154. Aetana indah Huber, sp. nov. 150-151. Feft male palp, prolateral and retrolateral views

(arrow points at prolateral apophysis near hinge). 152. Male chelicerae, frontal view. 153-154. Cleared

female genitalia, ventral and dorsal views (arrow points at membranous pocket), ip = internal sclerotized

pocket; rp = retrolatero-ventral process; vl = ventral lamina. Scale lines: 0.5 mm.

150

153

154

HUBER B.A. et al ., Revision of Aetana spiders

Figs 155-162. Aetana indah Huber, sp. nov. 155-156. Left palp, prolatero-distal and retrolatero-

distal views. 157. Left procursus, retrolatero-distal view. 158. Detail of left procursus (and trochanter

apophysis), prolateral view. 159. Left embolus (arrow points at sperm duct opening). 160-161. Left

male cheliceral apophyses, oblique frontal and frontal views. 162. Male gonopore. b = genital bulb; e

= embolus; f = femur; p = procursus; ti = tibia; tr = trochanter. Scale lines: 155-156 = 200 pm; 157 =

100 pm; 158-160 = 60 pm; 161 = 80 pm; 162 = 30 pm.

European Journal of Taxonomy 162: 1-78 (2015)

Figs 163-177. Aetana kinabalu group. Female genitalia; untreated in ventral view, cleared in ventral

and dorsal views. — 163-165. A. kinabalu Huber, 2005. — 166-168. A. lambir Huber, sp. nov. — 169-

171. A. gaya Huber, sp. nov. — 172-174. A. poring Huber, sp. nov. — 175-177. A. indah Huber, sp. nov.

HUBER B.A. et al ., Revision of Aetana spiders

Natural history

The spiders were found very close to the ground in small holes and cavities and barely reacted to

disturbance (similar only to A. poring Huber, sp. nov.; see above). They share the locality with A.

kinabahi which lives higher among the vegetation.

Distribution

Known from Crocker Range only (Fig. 5).

Aetana abadae Huber, sp. nov.

urn:lsid:zoobank.org:act:C0C92230-lA67-45FC-BCC4-3B66E2E82EC0

Figs 178-179, 186, 189-193, 195-197,213-215

Diagnosis

Distinguished from closest known relative (A. omayan) by male clypeus modification (Fig. 191;

apophyses closer together), distinct ventro-distal apophysis on male palpal femur (Fig. 190; only

indistinct hump in A. omayan ), and shape of epigynum (Fig. 213; whitish areas smaller and wider apart).

Distinguished from other congeners by bipartite retrolatero-ventral process on procursus (Fig. 190),

male palpal trochanter with prolateral apophysis (Fig. 189; other species with only ventral apophysis),

and pair of internal sclerotized pockets in female genitalia (Figs 192-193).

Etymology

Named for Philippine-born cosmopolitan artist Pacita Abad (1946-2004), famous for her vibrant,

colorful abstract work, but also for her paintings of tropical flowers and animal wildlife.

Material examined

Holotype

PHIFIPPINES: □, Negros Isl., Negros Oriental Prov., Twin Fakes N.P (9.365-9.368° N, 123.181°-

123.182° E), 850-950 m a.s.l., forest above Baliansasayao Crater Fake, 9 Mar. 2014 (B.A. Huber),

ZFMK (Ar 13986).

Other material

PHIFIPPINES, Negros Isl., Negros Oriental Prov.: 5 □ □, 13 □ □, same data as holotype, ZFMK (4 □ A f:

12 □□; Ar 13987-88) and MSU-IIT (1 □, 1 □); 1 □, 4 juvs, in pure ethanol, same data, ZFMK (Phi

193). - ID, Casaroro Falls (9.281° N, 123.208° E), 550 m a.s.l., forest along river below waterfall, 10

Mar. 2014 (B.A. Huber), ZFMK (Ar 13989); 1 □, in pure ethanol, same data, ZFMK (Phi 189).

Description

Male (holotype)

Measurements. Total body length 3.4, carapace width 1.4. Feg 1: 44.3 (10.6 + 0.6 + 10.6 + 19.2 +

3.3), tibia 2: 6.6, tibia 3: 4.5, tibia 4: 6.5; tibia 1 F/d: 80. Distance PME-PME 430 pm, diameter PME

140x155 pm, distance PME-AFE ~40 pm; AME absent.

Color. Carapace ochre-yellow with narrow lateral marginal bands and wide dark brown median band

including ocular area and clypeus; sternum ochre-yellow, with darker triangular mark posteriorly and

dark labium; legs ochre-yellow with slightly darker rings on femora (subdistally, with light tip), and

tibiae (proximally and subdistally, the latter followed by light tip); abdomen ochre-gray, dorsally and

laterally covered with many black marks, ventrally with dark mark behind gonopore.

European Journal of Taxonomy 162: 1-78 (2015)

Figs 178-188. Live specimens. Aetana omayan group. — 178-179. A. abadae Huber, sp. nov., □ and

□ with eggsac from Twin Lakes, Negros. — 180-181. A. omayan Huber, 2005, □ and □ with eggsac

from Baguio, Luzon. — 182-183. A. manansalai Huber, sp. nov., □ and □ from Mt. Banahaw, Luzon.

— 184-185. A. lozadae Huber, sp. nov.,.pD from Mt. Isarog, Luzon. — 186-188. Female prosomata,

showing stridulatory plates (arrows), in A. abadae Huber, sp. nov. (186), A. omayan Huber, 2005 (187)

and A. manansalai Huber, sp. nov. (188).

HUBER B.A. et al ., Revision of Aetana spiders

Body. Habitus as in Fig. 178; ocular area raised, each triad on additional short hump directed toward

lateral, without process below ALE (Fig. 191); carapace with very shallow median furrow in anterior

part only; clypeus with distinctive pair of apophyses (Fig. 191); sternum wider than long (0.95/0.65),

unmodified.

Chelicerae. As in Fig. 191, with pair of lateral processes proximally and pair of long lateral apophyses

distally; without modified hairs; without stridulatory ridges.

Palps. As in Figs 189-190; coxa unmodified; trochanter with ventral and prolateral apophyses; femur

with retrolateral hump, large prolateral apophysis, and ventro-distal apophysis; patella triangular in

lateral view; tibia with retrolateral trichobothrium in very distal position; proximal part of procursus with

bipartite retrolatero-ventral process, with complex and apparently partly hinged distal elements; bulb with

only one process (weakly sclerotized embolus), distally with indistinct hump, without small knobs.

Legs. Without spines; with curved hairs on metatarsi 1-3; few vertical hairs; retrolateral trichobothrium

on tibia 1 at 2%; prolateral trichobothrium absent on tibia 1, present on other tibiae; tarsus 1 with ~25

pseudosegments, only distally distinct.

Male (variation)

Tibia 1 in six other males: 9.8-10.6 (mean: 10.2). Dark spot behind gonopore absent in two males.

Abdomen with or without additional white spots in dorso-lateral rows. Male from Casaroro Falls with

large light brown mark on sternum posteriorly.

Female

In general similar to male but clypeus unmodified and more homogeneously dark brown; eye triads much

closer together (distance PME-PME 165 pm); with indistinct stridulatory apparatus between carapace

and abdomen: small modified area medially on carapace (Fig. 186) versus barely distinguishable hairless

area on abdomen. Tibia 1 in 13 females: 7.8-8.8 (mean: 8.1). Epigynum large sclerotized plate with pair

of light lateral humps (Figs 192, 195, 213), with pair of very indistinct membranous pockets behind

epigynum in weakly modified cuticle (weak transversal ridges) (Figs 192, 195-196). Internal genitalia

as in Figs 193 and 215, with distinct pair of sclerotized pockets.

Natural history

At both localities the spiders were found in domed sheet webs close to the ground, usually in well

protected dark spaces under large rocks.

Distribution

Known from two localities on Negros Island only (type locality and nearby locality; Fig. 5).

Aetana omayan Huber, 2005

Figs 180-181, 187, 194, 198-212,216-218

Aetana omayan Huber, 2005a: 73-74, figs 104-105, 110-114 (□□).

Note

The original description was based on a single male and two females. Here we present data on new

material from the type locality and a nearby locality, as well as an amended diagnosis to account for the

newly described congeners.

European Journal of Taxonomy 162: 1-78 (2015)

j, . i

Tf; Mgj|g

t , v\ I V

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m wf.

/% . v . /

\ rp /

' ^ t i/ l f

W.;mij J(i /

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Figs 189-194. — 189-193. Aetana abadae Huber, sp. nov. 189-190. Left male palp, prolateral and

retrolateral views. 191. Male prosoma, oblique frontal view. 192-193. Cleared female genitalia, ventral

and dorsal views. — 194. A. omayan Huber, 2005, cleared female genitalia, dorsal view. Arrows point

at membranous pockets, ip = internal sclerotized pocket; rp = retrolatero-ventral process; vl = ventral

lamina. Scale lines: 0.5 mm (female genitalia at same scale).

HUBER B.A. et al ., Revision of Aetana spiders

Figs 195-203. — 195-197. Aetana abadae Huber, sp. nov. 195. Epigynum, ventral view. 196. Detail of

preceding, showing opening of membranous pocket (arrow on Fig. 195). 197. Female ALS. — 198-203.

A. omayan Huber, 2005. 198. Male prosoma, frontal view. 199. Left palp, retrolateral view. 200-201.

Right procursus, prolatero-dorsal and prolateral views. 202. Detail of preceding. 203. Male palpal tarsal

organ, b = genital bulb; e = embolus; f = femur; p = procursus; rp = retrolatero-ventral process; ti =

tibia; tr = trochanter; vl = ventral lamina. Scale lines: 195, 199 = 300 pm; 196, 202 = 40 pm; 197, 203

= 10 pm; 198 = 500 pm; 200 = 200 pm; 201 = 100 pm.

European Journal of Taxonomy 162: 1-78 (2015)

Diagnosis

Distinguished from closest known relative (A. abadae Huber, sp. nov.) by male clypeus modification

(apophyses wider apart; cf Huber 2005a: fig. 112), very indistinct ventro-distal apophysis on male

palpal femur (distinct in A. abadae Huber, sp. nov., cf Fig. 190), and shape of epigynum (whitish areas

Figs 204-212. A. omayan Huber, 2005. 204. Left procursus tip (ventral lamina), retrolateral view.

205. Right procursus tip (pointed sclerite and ventral lamina), prolateral view. 206. Male gonopore.

207. Detail of male tarsus 1. 208-209. Comb-hairs on male tarsus 4. 210-211. Male and female ALS.

212. Epigynum. Scale lines: 204 = 50 pm; 205 = 40 pm; 206, 208 = 30 pm; 207 = 60 pm; 209 = 8 pm;

210-211 = 20 pm; 212 = 400 pm.

HUBER B.A. et al ., Revision of Aetana spiders

larger and closer together; Fig. 216). Distinguished from other congeners by bipartite retrolatero-ventral

process on procursus (Huber 2005a: fig. Ill), male palpal trochanter with prolateral apophysis (other

species only with ventral apophysis), and pair of internal sclerotized pockets in female genitalia (Fig.

194).

New material examined

PHIFIPPINES, Fuzon Isl., Benguet Prov.: 9 □□, 16 □□, 2 juvs, Baguio, Crystal Cave (16.396° N,

120.572° E), 1360 ma.s.l., 2 Mar. 2014 (B.A. Huber), ZFMK (8 □□, 15 Ar 13990) and MSU-IIT

(lgjg, 1 □); 1 6 juvs, in pure ethanol, same data, ZFMK (Phi 206). 3 □ Jty 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 13991); 1 □, 5 juvs, in pure ethanol, same data, ZFMK (Phi 205).

Description (amendments to Huber 2005a)

Male. Tibia 2 slightly shorter than tibia 4 (e.g., 7.1/7.3); curved hairs on all tibiae and metatarsi; tibia 1

in 8 males: 9.7-10.9 (mean: 10.4).

Female. Eye triads much closer together than in male (distance PME-PME -180-200 pm vs. 300^100

pm); indistinct stridulatory apparatus between carapace and abdomen: small modified area medially on

carapace (Fig. 187) versus light brown hairless area on abdomen; with pair of very indistinct membranous

pockets behind epigynum in unmodified cuticle (Figs 194, 218). Tibia 1 in 16 females: 7.6-8.5 (mean:

8 . 2 ).

Figs 213-218. Aetana omayan group, female genitalia, part 1 (cf. Figs 234-242); untreated in ventral

view, cleared in ventral and dorsal views. 213-215. A. abadae Huber, sp. nov. 216-218. A. omayan

Huber, 2005.

European Journal of Taxonomy 162: 1-78 (2015)

Natural history

The type locality is a highly degraded and polluted cave in the midst of a suburb of Baguio City. The

cave is actually a natural tunnel of about 50 m length, open on both sides. Within the cave, the spiders

were found in high numbers, building their typical domed sheet webs mainly along lower parts of the

cave walls near the ground, in crevices and small holes. The finding of the same species among rocks

on nearby Mt. Kabuyao suggests that the species is actually widespread in the area and not in danger of

extinction by further degradation of the cave.

Distribution

Known from two localities on Tuzon Island only (type locality and nearby locality; Fig. 5).

Aetana manansalai Huber, sp. nov.

urn:lsid:zoobank.org:act:087C21AC-F569-43EE-A817-3B0407AlClEA

Figs 182-183, 188, 219-223, 229, 234-236

Diagnosis

Distinguished from closest known relatives (A. banahaw Huber, sp. nov., A. lozadae Huber, sp. nov.)

by combination of smaller retrolatero-ventral process and smaller dorso-distal sclerite on procursus

(compare F igs 229-231) and by narrower epigynum with anterior half wider and more heavily sclerotized

than posterior half (Figs 222, 234). Distinguished from A. lozadae Huber, sp. nov. also by smaller

projections at ALE (Fig. 221). Distinguished from other congeners by presence of projections at ALE,

by long epigynum, and by posterior membranous pockets close together (Fig. 222).

Etymology

Named for the Filipino painter and illustrator Vicente Silva Manansala (1910-1981), most famous for

his ‘Madonna of the Slums’.

Material examined

Holotype

PHILIPPINES: □, Luzon Isl., Quezon Prov., between Lucban and Tayabas (14.063° N, 121.567° E),

330 m a.s.l., degraded forest along river, near ground, 26 Feb. 2014 (B.A. Huber), ZFMK (Ar 13992).

Other material

PHILIPPINES, Luzon Isl.: Quezon Prov., 5 □ □, 13 □ □, same data as holotype, ZFMK (4 □ □, 12 □ □)

(Ar 13993-94) and MSU-IIT (1 1 □); 3 □□, 1 juv., in pure ethanol, same data, ZFMK (Phi 213). -

Laguna Prov., 2 UU,2 □□jMt. Banahaw, forest near Taytay Falls (14.110° N, 121.507° E), 560ma.s.l.,

near ground, 26 Feb. 2014 (B.A. Huber), ZFMK (Ar 13995).

Description

Male (holotype)

Measurements. Total body length 3.2, carapace width 1.3. Leg 1: 36.5 (8.5 + 0.5 + 8.5 + 15.5 + 3.5),

tibia 2: 5.2, tibia 3: 3.5, tibia 4: 5.3; tibia 1 L/d: 67. Distance PME-PME 350 pm, diameter PME

125><150 pm, distance PME-ALE ~45 pm; AME absent.

Color. Carapace ochre-yellow with narrow lateral marginal bands and wide dark brown median band

including posterior part of ocular area; clypeus ochre yellow with indistinct light brown pattern; sternum

light brown to orange, laterally paler, labium darker; legs greenish ochre with slightly darker rings on

femora (subdistally, with light tip), and tibiae (proximally and subdistally, the latter followed by light

HUBER B.A. et al ., Revision of Aetana spiders

Figs 219-223. Aetana manansalai Huber, sp. nov. 219-220. Left male palp, prolateral and retrolateral

views. 221. Male prosoma, oblique frontal view. 222-223. Cleared female genitalia, ventral and dorsal

views. Arrow points at membranous pocket, rp = retrolatero-ventral process; vl = ventral lamina. Scale

lines: 0.5 mm

European Journal of Taxonomy 162: 1-78 (2015)

tip); abdomen ochre-gray, dorsally and laterally covered with many black marks, ventrally with dark

mark behind gonopore and larger less distinct mark in front of spinnerets.

Body. Habitus as in Fig. 182; ocular area raised, each triad on additional short hump directed toward

lateral, with small process below ALE (Fig. 221); carapace with very shallow median furrow in anterior

part only; clypeus medially slightly projecting, with distinctive lateral plates bordered by sclerotized

ridges (Fig. 221); sternum wider than long (0.85/0.60), unmodified.

Chelicerae. As in Fig. 221, with pair of lateral processes proximally and pair of very long lateral

apophyses; without modified hairs; without stridulatory ridges.

Palps. As in Figs 219-220; coxa unmodified; trochanter with ventral apophysis; femur with small

ventro-distal apophysis and retrolateral ridge ending in small hump; patella triangular in lateral view;

tibia with retrolateral trichobothrium in very distal position; proximal part of procursus with simple

retrolatero-ventral process, with complex and apparently partly hinged distal elements (Fig. 229); bulb

with only one process (weakly sclerotized embolus), distally with several small knobs.

Legs. Without spines; with curved hairs on metatarsi 1-3; few vertical hairs; retrolateral trichobothrium

on tibia 1 at 3%; prolateral trichobothrium absent on tibia 1, present on other tibiae; tarsus 1 with ~25

pseudosegments, only distally distinct.

Male (variation)

Tibia 1 in 5 other males: 8.0-8.8 (mean: 8.3). Abdomen with or without additional white spots in dorso¬

lateral rows.

Female

In general similar to male but clypeus unmodified and with pair of dark brown bands below ALE; eye

triads much closer together (distance PME-PME 150 pm), without processes at ALE; with indistinct

stridulatory apparatus between carapace and abdomen: modified area medially on carapace (Fig. 188)

versus barely distinguishable hairless area on abdomen. Tibia 1 in 13 females: 5.8-6.5 (mean: 6.3); dark

and light rings on legs often more distinct than in males. Epigynum long, anterior half of plate wider and

more heavily sclerotized than posterior half, anterior half with pair of low humps (Figs 222, 234); area

behind epigynum with pair of very indistinct membranous pockets in weakly modified cuticle (weak

transversal ridges). Internal genitalia as in Figs 223 and 236, without sclerotized pockets.

Natural history

At both localities the spiders were found in domed sheet webs close to the ground, usually in well

protected dark spaces under large rocks. The type locality suggests that the species does not depend on

well preserved forests and is probably widespread in the area.

Distribution

Known from two localities on Luzon Island only (type locality and nearby locality; Fig. 5).

Aetana lozadae Huber, sp. nov.

urn:lsid:zoobank.org:act:3526A886-2551-4FBE-900A-603D92A43E23

Figs 184-185, 224-228, 230, 237-239

Diagnosis

Distinguished from closest known relatives (A. manansalai Huber, sp. nov., A. banahaw Huber, sp. nov.)

by combination of long projections at ALE (Fig. 226), large retrolatero-ventral process on procursus (Fig.

HUBER B.A. et al ., Revision of Aetana spiders

230; similar to A. banahaw Huber, sp. nov.), and oval shape of epigynum (Figs 227, 237). Distinguished

from other congeners by presence of projections at ALE, by epigynum longer than wide, and by posterior

membranous pockets close together (Figs 227-228).

Etymology

Named for Filipino violinist Carmencita Lozada (1940-2006), prize winner of the Paganini International

Violin Competition in Italy.

Material examined

Holotype

PHILIPPINES: □, Luzon, Camarines Sur Prov., Mt. Isarog, W slope (13.664° N, 123.34-123.35° E),

-600-900 m a.s.l., forest, near ground, 23 Feb. 2014 (B.A. Huber), ZFMK (Ar 13996).

Other material

PHILIPPINES, Luzon Isl., Camarines Sur Prov.: 3 □□, 13 same data as holotype, ZFMK (Ar

13997-98); 1 □, in pure ethanol, same data, ZFMK (Phi 221). - 1 □, 1 □, in pure ethanol, Mt. Isarog

(13.665° N, 123.354° E), 9.3 km E ofNaga City, 920 m a.s.l., 31 May-2 June 2011 (M. Yngente et al.),

CAS (9042055).

Description

Male (holotype)

Measurements. Total body length 3.9, carapace width 1.3. Leg 1: 40.1 (9.2 + 0.5 + 9.2 + 17.7 + 3.5),

tibia 2: 5.8, tibia 3: 4.0, tibia 4: 5.8; tibia 1 L/d: 71. Distance PME-PME 390 pm, diameter PME

135x155 pm, distance PME-ALE -45 pm; AME absent.

Color. Carapace ochre-yellow with narrow dark lateral marginal bands and wide dark brown median

band including posterior part of ocular area; clypeus ochre yellow with indistinct light brown pattern;

sternum light brown to orange, labium darker; legs greenish ochre with slightly darker rings on femora

(subdistally, with light tip), and tibiae (proximally and subdistally, the latter followed by light tip);

abdomen ochre-gray, dorsally and laterally covered with many black marks, ventrally with dark mark

behind gonopore and larger less distinct mark in front of spinnerets.

Body. Habitus as in Figs 184-185; ocular area raised, each triad on additional hump directed toward

lateral, with long process at ALE (Fig. 226); carapace with very shallow median furrow in anterior part

only; clypeus medially not projecting, with distinctive lateral plates bordered by sclerotized ridges (Fig.

226); sternum wider than long (0.85/0.65), unmodified.

Chelicerae. As in Fig. 226, with pair of lateral processes proximally and pair of very long lateral

apophyses, without modified hairs; without stridulatory ridges.

Palps. As in Figs 224-225; very similar to A. manansalai Huber, sp. nov. and A. banahaw Huber,

sp. nov.; procursus as in Fig. 230.

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

on tibia 1 at 2%; prolateral trichobothrium absent on tibia 1, present on other tibiae; tarsus 1 with -25

pseudosegments, only distally distinct.

Male (variation)

Tibia 1 in 2 other males: 8.8, 9.2. Abdomen with or without additional white spots in dorso-lateral rows

(white spots seem to get partly lost or indistinct in alcohol).

European Journal of Taxonomy 162: 1-78 (2015)

Figs 224-228. Aetana lozadae Huber, sp. nov. 224-225. Left male palp, prolateral and retrolateral

views. 226. Male prosoma, oblique frontal view. 227-228. Cleared female genitalia, ventral and dorsal

views, rp = retrolatero-ventral process; vl = ventral lamina. Scale lines: 0.5 mm.

HUBER B.A. et al ., Revision of Aetana spiders

■ * •'‘X

W'Vvt,

A- °c*’ V ,*«*•»

f ‘j, ‘ »,o0 4 o',<'o s *

oP p c» - 0 ®* 0

I fl“ 1 * a % *«® c e

.: Ptf 6 «g«e« a

■ °i**VJ**o.**

*> ' ‘

~.jfe a*>

f ; -#o

A O M

Figs 229-233. Left procursi, retrolateral views (asterisk: retrolatero-distal sclerite). — 229. Aetana

manansalai Huber, sp. nov. — 230. A. lozadae Huber, sp. nov. — 231. A. banahaw Huber, sp. nov. Figs

229-231 at same scale. — 232-233. A. banahaw Huber, sp. nov., cleared female genitalia, ventral and

dorsal views, rp = retrolatero-ventral process; vl = ventral lamina. Scale lines: 0.5 mm.

European Journal of Taxonomy 162: 1-78 (2015)

Female

In general similar to male but clypeus unmodified and most females with pair of dark brown bands

below ALE; eye triads much closer together (distance PME-PME 155 pm), without processes near

ALE; with indistinct stridulatory apparatus between carapace and abdomen: modified area medially on

carapace (smaller than in A. manansalai Huber, sp. nov.) versus barely distinguishable hairless area on

abdomen. Tibia 1 in 9 females: 6.7-7.7 (mean: 7.2); dark and light rings on legs often more distinct than

in males. Epigynum oval, longer than wide (Figs 227, 237), anterior half with pair of low humps; area

behind epigynum with pair of very indistinct membranous pockets in weakly modified cuticle (weak

transversal ridges). Internal genitalia as in Figs 228 and 239, without sclerotized pockets.

Natural history

The spiders were found in domed sheet webs close to the ground, usually in well protected dark spaces

under large rocks and logs, but apparently not as deeply hidden in these cavities as the sympatric

A. ocampoi Huber, sp. nov.

Distribution

Known from type locality on Luzon Island only (Fig. 5).

Aetana banahaw Huber, sp. nov.

urn:lsid:zoobank.org:act:6D270C60-BBF0-4DFE-88BC-lF90D87C5D8F

Figs 231-233,240-242

Diagnosis

Distinguished from closest known relatives (A. manansalai Huber, sp. nov., A. lozadae Huber, sp. nov.)

by shape of procursus (large retrolatero-distal sclerite; compare Figs 229-231) and by pentagonal

epigynum (Figs 232, 240); from A. manansalai Huber, sp. nov. also by larger retrolatero-ventral process

on procursus (Fig. 231) and narrower apophysis of male palpal trochanter; from A. lozadae Huber,

sp. nov. also by smaller projections at ALE (similar to A. manansalai Huber, sp. nov.; cf Fig. 221).

Distinguished from other congeners by presence of projections at ALE, by longer than wide epigynum,

and by posterior membranous pockets close together (Fig. 232).

Etymology

Named for the type locality; noun in apposition.

Material examined

Holotype

PHILIPPINES: □, Luzon Isl., Laguna Prov., Mt. Banahaw, forest near Taytay Falls (14.110° N,

121.507° E), 560 m a.s.l., near ground, 26 Feb. 2014 (B.A. Huber), ZFMK (Ar 13999).

Other material

PHILIPPINES, Luzon Isl., Laguna Prov.: 7 CD,4 □ ^ same data as holotype, ZFMK (Ar 14000-01); 1 □,

4 juvs, in pure ethanol, same data, ZFMK (Phi 217). - l! T s 1 juv., in pure ethanol, Mt. Banahaw (14.103°

N, 121.518° E), 4.38 km W of Lucban, 790ma.sT, 16May2011 (H. Wood et al.\ CAS (9045550).

Description

Male (holotype)

Measurements. Total body length 3.7, carapace width 1.35. Leg 1: 36.7 (8.6 + 0.5 + 8.6 + 15.4 +

3.6), tibia 2: 5.5, tibia 3: 3.8, tibia 4: 5.6; tibia 1 L/d: 63. Distance PME-PME 395 pm, diameter PME

140 pm, distance PME-ALE ~70 pm; AME absent.

HUBER B.A. et al ., Revision of Aetana spiders

Figs 234-242. Aetana omayan group, female genitalia, part 2 (cf. Figs 213-218); untreated in ventral view,

cleared in ventral and dorsal views. — 234-236. A. manansalai Huber, sp. nov. — 237-239. A. lozadae

Huber, sp. nov. — 240-242. A. banahaw Huber, sp. nov.

European Journal of Taxonomy 162: 1-78 (2015)

Color. Carapace ochre-yellow with narrow dark lateral marginal bands and wide dark brown median

band including posterior part of ocular area; clypeus ochre yellow, small marks below each eye triad;

sternum monochromous light brown, labium darker; legs greenish ochre with very indistinct darker

rings on femora (subdistally, with light tip), and tibiae (proximally and subdistally, the latter followed

by light tip); abdomen ochre-gray, dorsally and laterally covered with many black marks, ventrally with

dark mark behind gonopore and larger, less distinct mark in front of spinnerets.

Body. Habitus very similar to A. lozadae Huber, sp. nov. {cf Figs 184-185); ocular area raised, each

triad on additional short hump directed toward lateral, with small process below ALE (only slightly

longer than in A. manansalai Huber, sp. nov.; cf Fig. 221); carapace with very shallow median furrow in

Figs 243-248. Male gonopores and ALS. — 243-244. Khorata khammouan Huber, 2005. — 245-

246. K. dupla Yao & Li, 2013. — 247-248. K. circularis Yao & Li, 2013. Scale lines: 243 = 50 pm; 244,

246, 248 = 10 pm; 245 = 40 pm; 247 = 60 pm.

HUBER B.A. et al ., Revision of Aetana spiders

anterior part only; clypeus with distinctive lateral plates bordered by sclerotized ridges; sternum wider

than long (0.9/0.7), unmodified.

Chelicerae. As in close relatives (cf. Figs 221, 226), with pair of lateral processes proximally and pair

of very long lateral apophyses; without modified hairs; without stridulatory ridges.

Palps. In general as in A. manansalai Huber, sp. nov. and A. lozadae Huber, sp. nov. (cf Figs 219-220,

224-225); coxa unmodified; trochanter with ventral apophysis slightly narrower than in A. manansalai

Huber, sp. nov.; femur with ventro-distal apophysis and retrolateral ridge ending in small hump; patella

triangular in lateral view; tibia with retrolateral trichobothrium in very distal position; proximal part of

procursus, with simple retrolatero-ventral process, with complex and apparently partly hinged distal

elements; bulb with only one process (weakly sclerotized embolus), distally with one small knob.

Fegs. Without spines; with curved hairs on metatarsi 1-3; few vertical hairs; retrolateral trichobothrium

on tibia 1 at 3%; prolateral trichobothrium absent on tibia 1, present on other tibiae; tarsus 1 with ~25

pseudosegments, distally fairly distinct.

Male (variation)

Tibia 1 in 7 other males: 8.2-9.4 (mean: 8.9).

Female

In general similar to male but clypeus unmodified and with pair of dark brown bands below AFE; eye

triads much closer together (distance PME-PME 185 pm), without processes at AFE; with indistinct

stridulatory apparatus between carapace and abdomen: modified area medially on carapace versus

barely distinguishable hairless area on abdomen. Tibia 1 in 4 females: 6.6, 6.8, 7.1, 7.3; dark and light

rings on legs mostly more distinct than in males. Epigynum as in Figs 232 and 240, anterior large plate

pentagonal, with transversal anterior bulge bordered posteriorly by shallow indentation; area behind

epigynum with pair of very indistinct membranous pockets in weakly modified cuticle (weak transversal

ridges). Internal genitalia as in Figs 233 and 242, without sclerotized pockets.

Natural history

The spiders were found in domed sheet webs close to the ground, usually in well protected dark spaces

under large rocks. Males and females were sometimes found together in one web. When disturbed, the

spiders ran to the rock, vibrated only for a moment and then remained motionless, pressed against the

rock surface.

Distribution

Known from the type locality only (Fig. 5).

Discussion

Our main objectives were to document the high diversity of Aetana in Southeast Asia, and to provide

first data about ultrastructure, relationships, and ecology. However, while we added substantial new data

about Philippine and Borneo species, individual records reaching as far east as Fiji suggest that a huge

diversity remains to be discovered and described. Our cladistic analysis is certainly a step forward, but

while some details are congruent with preliminary molecular data (A. Valdez-Mondragon, D. Dimitrov,

B.A. Huber, unpublished data), others are in conflict and need further study. Both analyses agree on

the monophyly of Aetana and on the four species groups proposed herein. However, the sister group of

Aetana remains dubious: the present analysis suggests a group of three Southeast Asian genera as sister

( Khorata , Savarna and an undescribed genus from northern Borneo), whereas the molecular analysis

European Journal of Taxonomy 162: 1-78 (2015)

currently suggests Southeast Asian Spermophora as sister. Likewise, relationships among species groups

within Aetana remain dubious and require further sampling and analysis.

Based on our field observations and the clear support for the four species groups in the cladistic analysis,

we conclude that at least two microhabitat shifts have occurred in Aetana. Even if the four species

groups are placed in a basal tetrachotomy, assuming a plesiomorphic microhabitat higher among the

vegetation would require a minimum of three shifts toward the ground. On the other hand, assuming that

the plesiomorphic microhabitat of Aetana is near the ground (as is the case in most species, including

the cave-dwelling A. omayan ) requires only two shifts, one in the Philippines (in the A. ocampoi group),

and one on Borneo (in the A. kinabalu group). Further support for a shift from the ground towards higher

vegetation comes from the outgroup taxa. Except for Zatavua griswoldi Huber, 2003 (for which no

microhabitat data are available), all other species have been collected in the leaf litter or very close to

the ground. The microhabitat of A.fiji has never been documented, but label data (Huber 2005a) suggest

that it might represent a third independent case of microhabitat shift within Aetana.

Finally, we document a potential further case of genital dimorphism (in A. ocampoi Huber, sp. nov.),

admitting that our data are not particularly strong. Genital dimorphism is a nuisance for invertebrate

taxonomy, as it runs counter to one of the very central pillars of the discipline: species-specificity of

genitalia (e.g., Huber & Perez-Gonzalez 2001; Jocque 2002; Mutanen & Kaitala 2006). For this reason,

any proposal for a new case should ideally be supported by convincing evidence, for example derived

from rearing experiments such as in Huber & Perez-Gonzalez (2001). We do not have this kind of

data for A. ocampoi Huber, sp. nov., our sample size is small, and we have not so far succeeded in

sequencing any of the two standard barcoding genes (16S, COl) for this species. However, we document

this potential case in the hope that it might stimulate further field work and research directed at this

particular species. We also propose that only a change of paradigm might reveal the true abundance of

species with polymorphic genitalia. Traditional invertebrate taxonomy has developed an almost absolute

reliance on genital morphology to separate species, which is a major impediment for discovering genital

polymorphism in the first place. If species are routinely distinguished by genital morphology, then

different morphs will likely end up as different species, which in turn supports the original idea about

species-specificity. This circle can only be broken if taxonomists routinely consider the possibility of

genital polymorphism, even at the risk of being rebutted.

Acknowledgements

We thank our Philippine driver Vicente Bulongon for bringing us safely to all requested places. Samuel

B. Huber, Mae A. Responte, Eddie P. Mondejar, and Philip N.O. Banaag II helped in the field and

the latter three also contributed specimens from their own research. Permits were received from the

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

Region XI and from the Forest Department, Sarawak (Permit No. NCCD.907.4.4(JLD.10)-211, Park

Permit No. 174/2014). The German Research Foundation provided financial support (DFG Project HU

980/11-1).

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Manuscript received. 10 September 2015

Manuscript accepted. 14 October 2015

Published on: 14 December 2015

Topic editor : Rudy Jocque

Desk editor: Kristiaan Hoedemakers

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.

HUBER B.A. et al ., Revision of Aetana spiders

Appendix 1

Data matrix for cladistic analysis. Taxa are arranged as in the cladogram in Fig. 1. Terminals and characters

are detailed in Appendices 2 and 3; = non-applicable; “?” = unknown or coding ambiguous. This

matrix is available online at www.pholcidae.de/matrices.html

5 10 15 20 25 30 35 40

Zatavua griswoldi Huber, 2003

Spermophora estebani Simon, 189'

Khorata khammouan Huber, 2005

Savarna thaleban Huber, 2005

Gen.n. Bor80

Gen.n. Bor20

Aetana ocampoi Huber, sp. nov.

Aetana libjo Huber, sp. nov.

Aetana baganihan Huber, sp. nov.

Aetana kiukoki Huber, sp. nov.

Aetana paragiia Huber, sp. nov.

Aetana loboe Huber, sp. nov.

Aetana pasambai Huber, sp. nov.

Aetana gay a Huber, sp. nov.

Aetana kinabalu Huber, 2005

Aetana lambir Huber, sp. nov.

Aetana poring Huber, sp. nov.

Aetana indah Huber, sp. nov.

Aetana fiji Huber, 2005

Aetana abadae Huber, sp. nov.

Aetana omayan Huber, 2005

Aetana manansalai Huber, sp. nov.

Aetana lozadae Huber, sp. nov.

Aetana banahaw Huber, sp. nov.

0005000000000100000000

0000001100000000001001

2000101100000000010000

200011--00100010000000

2004100001001000000000

2000100001001000000000

1000000010100010001100

1000000010100000001100

100301--00000000011000

1 10301--00000010011000

110301--00000010011000

0000000100001010101011

0000000100000011101011

0000000100000010101010

0000000100001111101011

1002001100000011001011

1002001100010010001011

1002001100010011001011

1011001100000010001011

--400001000010100000-?

0040000100000-100000-0

-- 00000101100 - 000000-0

--30001101100-000000-0

-- 001111011011000000-0

--110000177010000000-0

- - 21001010101 1001000-1

- - 21001017701 1001000-1

--100000101010000010-0

--100000177010000010-0

— 10000017777777777770

0000000010100 - 010000-0

0000000017700-010100-1

--00000017700-010000-1

0100000017700-010000-0

0100000010100 - 010100-0

0000777077770-0007077?

0200000017710-00010100

0200000010110-00010100

1000000017712-00000110

European Journal of Taxonomy 162: 1-78 (2015)

Appendix 2

Terminal taxa scored for the cladistic analysis, with origin and depository. Taxa are arranged as in the

cladogram in Fig. 1.

Outgroups

Zatavua griswoldi Huber, 2003: data from Huber 2003a.

Spermophora estebani Simon, 1892: data from Huber 2005a.

Khorata khammouan Huber, 2005: data from Huber 2005a.

Savarna thaleban Huber, 2005: Thailand, Thale BanN.P. (ZFMK).

Gen.n. Bor80 Huber, sp. nov.: Malaysia, Sarawak, Kapit (ZFMK).

Gen.n. Bor20 Huber, sp. nov.: Malaysia, Sarawak, Niah (ZFMK).

Aetana

A. ocampoi Huber, sp. nov.: Philippines, Luzon, Mt. Isarog (ZFMK).

A. libjo Huber, sp. nov.: Philippines, Dinagat Isl. (ZFMK).

A. baganihan Huber, sp. nov.: Philippines, Mindanao, Baganihan (ZFMK).

A. kiukoki Huber, sp. nov.: Philippines, Mindanao, Baganihan (ZFMK).

A. paragua Huber, sp. nov.: Philippines, Dinagat Isl. (ZFMK).

A. loboc Huber, sp. nov.: Philippines, Bohol, Loboc (ZFMK).

A. pasambai Huber, sp. nov.: Philippines, Negros, Casaroro Falls (ZFMK).

A. kinabalu Huber, 2005: Malaysia, Sabah, Mt. Kinabalu (ZFMK).

A. lambir Fluber, sp. nov.: Malaysia, Sarawak, Lambir Hills (ZFMK).

A. gaya Huber, sp. nov.: Malaysia, Sabah, Gaya Isl. (ZFMK).

A. poring Huber, sp. nov.: Malaysia, Sabah, Poring (ZFMK).

A. indah Huber, sp. nov.: Malaysia, Sabah, Crocker Range (ZFMK).

A.fiji Huber, 2005: Data from Huber 2005a.

A. abadae Huber, sp. nov.: Philippines, Negros, Twin Lakes N.P (ZFMK).

A. omayan Huber, 2005: Philippines, Luzon, Baguio (ZFMK).

A. manansalai Huber, sp. nov.: Philippines, Luzon, betw. Lucban and Tayabas (ZFMK).

A. lozadae Huber, sp. nov.: Philippines, Luzon, Mt. Isarog (ZFMK).

A. banahaw Huber, sp. nov.: Philippines, Luzon, Mt. Banahaw (ZFMK).

Appendix 3

Characters scored for cladistic analysis.

Prosoma

HUBER B.A. et al., Revision of Aetana spiders

1. Thoracic furrow: (0) absent (Fig. 135); (1) shallow, only anteriorly (Fig. 63); (2) distinct over entire

length of carapace. The plesiomorphic condition in Pholcinae seems to be the absence of a thoracic

furrow. However, most representatives of Aetana have a shallow furrow in the anterior part of the

carapace. The absence in the Aetana kinabalu group is here considered secondary; the same is true

for the distinct and long furrows in Khorata, Savarna , and ‘Gen.n. Borneo’.

2. Male ocular area median process: (0) absent; (1) present. A median process (Figs 77, 96) is a unique

synapomorphy of Aetana loboc Huber, sp. nov. + A. pasambai Huber, sp. nov..

3. Male ocular area, ventral processes near AFE: (0) absent; (1) present. These unique processes (Figs

221, 226) unite three species of the Aetana omayan group: A. manansalai Huber, sp. nov., A. lozadae

Huber, sp. nov., and A. banahaw Huber, sp. nov..

4. Male clypeus: (0) unmodified or slightly bulging; (1) with pair of lateral ridges (Figs 221, 226); (2)

with pair of apophyses (Fig. 191); (3) with large sclerotized process at clypeus margin (Figs 59,

77); (4) with small median process between eyes and clypeus margin; (5) with pair of bulges at rim.

An unmodified clypeus seems to be plesiomorphic for Aetana. A large sclerotized process at the

clypeus rim is a synapomorphy of the Aetana kiukoki group; a pair of rather small apophyses unites

the Aetana omayan group but is modified into a pair of ridges in A. manansalai sp. nov., A. lozadae

Huber, sp. nov., and A. banahaw Huber, sp. nov.. Character states 4 and 5 are autapomorphic for

outgroup taxa (Zatavua griswoldi; ‘Gen.n. Bor 80 Huber, sp. nov.’).

5. Sternum color: (0) at least partly light; (1) entirely dark. A dark (usually black) sternum unites

several outgroup taxa (Savarna', Khorata', ‘Gen.n. Borneo’). All Aetana species have a light or

mostly light sternum.

Male chelicerae

6. Chelicerae distal apophyses: (0) present; (1) absent or indistinct hump. The presence of distal

apophyses (e.g., Figs 15, 115, 221) is interpreted to be plesiomorphic for Aetana. A reduction

occurred twice in the present matrix, once in Savarna, and once as a synapomorphy of the Aetana

kiukoki group (Figs 59, 72, 77, 96).

7. Chelicerae distal apophyses, length: (0) short; (1) long. Short distal apophyses seem to be

plesiomorphic for Aetana. Fong apophyses have evolved convergently in Spermophora estebani,

Khorata khammouan, and as a synapomorphy of the Aetana omayan group (Figs 191, 221, 226).

8. Chelicerae distal apophyses, position: (0) frontal; (1) lateral. The plesiomorphic position is

ambiguous. A lateral position may unite the Aetana kinabalu and A. omayan groups, but occurs

independently in Spermophora estebani and Khorata khammouan.

9. Chelicerae proximal apophyses: (0) present; (1) absent. Proximal cheliceral apophyses (Figs 59,

115, 221) are a synapomorphy of Pholcinae. In the present matrix, the reduction of these apophyses

(Figs 15, 40) is a synapomorphy of the Aetana ocampoi group.

Male pedipalp

10. Coxa ventral apophysis or rim: (0) absent or indistinct; (1) present and distinct. A ventral apophysis

is a synapomorphy of Modisiminae but is rare in other subfamilies. In the present matrix, it is an

autapomorphy of‘Gen.n. Borneo’.

11. Trochanter-femur connection: (0) normal hinge; (1) segments partly fused (Figs 14, 27). A partial

fusion between trochanter and femur occurs both in Savarna and in the Aetana ocampoi group, but

is not known in any other Pholcidae. The cladistic analysis suggests that these are two independent

origins.

European Journal of Taxonomy 162: 1-78 (2015)

12. Trochanter prolateral apophysis: (0) absent; (1) present. In most Pholcinae, the palpal trochanter has

one (retrolateral to ventral) apophysis. Additional apophyses (usually retrolateral or dorsal) have

evolved frequently. In the present matrix, two species of the Aetana omayan group are united by a

unique prolateral apophysis in addition to the usual ventral apophysis (Fig. 189).

13. Trochanter apophysis: (0) smooth; (1) with small teeth or scales (Fig. 158). The retrolatero-ventral

apophysis of the trochanter is usually smooth. Teeth or scales have evolved independently in ‘Gen.n.

Borneo’ and in three species of the Aetana kinabalu group.

14. Femur apophysis near prolateral joint to trochanter: (0) absent; (1) present. This small but distinct

apophysis (Figs 145, 150) is a synapomorphy of two species within the Aetana kinabalu group. A

similar process evolved independently in the outgroup species Zatavua griswoldi.

15. Femur distal retrolatero-ventral process: (0) absent; (1) present. Th q Aetana kinabalu and A. omayan

groups are united by this process, which is large and round in the A. kinabalu group (Figs 114, 151)

but smaller and more slender in the A. omayan group (Figs 190, 220). A similar structure occurs in

Aetana ocampoi Huber, sp. nov. (Fig. 14), in A. loboc Huber, sp. nov. + A. pasambai Huber, sp. nov.,

and in Savarna thaleban.

16. Femur prolateral process: (0) absent; (1) present. A prolateral process (not close to the femur-

trochanter hinge as char. 14; e.g., Fig. 145) has apparently evolved several times independently in

the Aetana kinabalu and A. omayan groups.

17. Femur distal prolatero-ventral process: (0) absent; (1) present. All species of the A. kinabalu group

share this sometimes large process on the femur (Figs 113, 126, 150).

18. Femur retrolateral apophysis: (0) absent or indistinct; (1) present/long (Figs 58, 71, 76, 98). A

distinct long apophysis is a synapomorphy of the A. kiukoki group.

19. Tibia retrolateral trichobothrium, position: (0) normal (i.e., at about 60-90% of tibia length); (1) very

distal (>90% of tibia length). All species of Aetana share a very distal position of the retrolateral

trichobothrium on the male palpal tibia (e.g., Figs 27, 71, 114). This is shared by Spermophora

estebani (and other species of Spermophora ; see Huber 2005a). The present analysis interprets this

as a convergence, but preliminary molecular data resolve Aetana and Southeast Asian Spermophora

as sister groups (A. Valdez-Mondragon, D. Dimitrov, B.A. Huber, unpublished data).

20. Procursus: (0) complex, with sclerotized elements; (1) simple semi-transparent, ribbon-shaped

process (Figs 14, 27). A highly reduced procursus is a unique synapomorphy of the Aetana ocampoi

group.

21. Procursus ventral lamina: (0) absent; (1) present. This distinctive process (e.g., Figs 127, 190, 229-

231) unites the Aetana kinabalu and A. omayan groups.

22. Procursus retrolatero-ventral process: (0) absent; (1) present. This sclerite (e.g.. Figs 127, 190, 229-

231) unites the Aetana kinabalu and A. omayan groups, but occurs independently in Spermophora

estebani (and other species of Spermophora ; see Huber 2005a).

23. Procursus retrolatero-ventral process, position: (0) transversal; (1) pointing toward distal. In the

outgroup taxon Spermophora estebani (and even more clearly in other species of Spermophora ;

see Huber 2005a) this sclerite has a transversal position on the procursus. This is thus considered

plesiomorphic for Aetana. In three species of the A. omayan group, the sclerite points toward distal

(Figs 229-231).

24. Procursus retrolatero-ventral process tip: (0) simple; (1) bifid; (2) bipartite. A simple tip seems to be

plesiomorphic. Two species of the A. kinabalu group share a bifid tip (Figs 146, 151), while in two

species of the A. omayan group the sclerite is deeply split in two parts (Fig. 190).

25. Genital bulb distal processes: (0) only embolus; (1) embolus plus short membranous process (Figs

58, 75); (2) embolus plus two partly sclerotized processes (Figs 38-39, 41^42); (3) single complex,

sclerotized process containing sperm duct; (4) embolus plus one partly sclerotized process. A

single sclerotized process (‘appendix’) in addition to a slightly sclerotized embolus (char, state 4)

is plesiomorphic for Aetana and probably for Pholcinae. The present analysis suggests a loss of

the appendix in the ancestor of Aetana + some outgroup taxa, and many species of Aetana retain

HUBER B.A. et al ., Revision of Aetana spiders

this character state (char, state 0). Most species of the Aetana kiakoki and A. ocampoi groups have

developed a short membranous process (char, state 1), and A. libjo Huber, sp. nov. and A. baganihan

Huber, sp. nov. share even two additional processes (char, state 2). A single complex, sclerotized

process containing the sperm duct (char, state 3) is a putative autapomorphy of Savarna (B.A. Huber

& B. Petcharad, unpubl. data).

26. Genital bulb proximal retrolateral process: (0) absent; (1) present. This indistinct process (Figs 14,

27) occurs in all species of the Aetana ocampoi group.

Legs

27. Male femur 1 spines: (0) absent; (1) present. Spines on the legs are very rare in Pholcinae. In

the present matrix, they only occur in ‘Gen.n. Bor 80 Huber, sp. nov.’and ‘Gen.n. Bor 20 Huber,

sp. nov.’. They also occur in two species of Savarna (B.A. Huber & B. Petcharad, unpubl. data) but

not in S. thaleban.

28. Male femora vertical hairs: (0) absent (or very few); (1) present in high density. Short vertical hairs

in high density on the femora are rare in Pholcidae (known in some Modisiminae genera; see Huber

2000). In the present matrix, they only occur in ‘Gen.n. Bor 80 Huber, sp. nov.’and ‘Gen.n. Bor 20

Huber, sp. nov.’.

29. Tibia vertical hairs: (0) absent (or very few); (1) present in high density. Short vertical hairs on

the tibiae are slightly more widespread than on the femora (especially in Modisiminae), but rare

in Pholcinae. According to the present analysis, they evolved independently in Savarna + (‘Gen.n.

Bor 80 Huber, sp. nov.’ + ‘Gen.n. Bor 20 Huber, sp. nov.’) and in Aetana libjo Huber, sp. nov. + A.

baganihan Huber, sp. nov..

30. Tibia 1 retrolateral trichobothrium position: (0) proximal (at <5% of tibia 1 length); (1) distal (at

>5% of tibia 1 length). In the present matrix, only Aetana is characterized by a very proximal

position of the retrolateral trichobothrium.

31. Curved hairs on tibiae and/or metatarsi: (0) absent; (1) present (Fig. 133). In the present matrix, only

Aetana is characterized by curved hairs on tibiae and/or metatarsi.

Abdomen

32. Epiandrous spigots: (0) present; (1) absent. The presence of epiandrous spigots is plesiomorphic for

Pholcinae and for Aetana. In the present matrix, the loss of these spigots unites several outgroup

taxa, i.e. Savarna , Khorata (Figs 243, 245, 247), and ‘Gen.n. Borneo’.

33. AES spigots: (0) eight; (1) two. The plesiomorphic condition in Pholcinae is seven to eight spigots:

one very wide spigot, one pointed spigot, and five to six smaller cylindrically-shaped spigots. In the

present matrix, Aetana and several outgroup taxa are united by the loss of the smaller cylindrically-

shaped spigots (Figs 68, 142, 210, 244, 246, 248).

34. Female stridulatory apparatus between prosoma and abdomen: (0) absent; (1) present. A modified

median plate on the carapace (Figs 186-188) acting against an indistinct hairless field on the abdomen

characterizes all species of the Aetana omayan group (not verified in A.jiji).

35. Epigynum posterior margin: (0) more or less straight; (1) with scape; (2) deeply indented. A straight

epigynal margin is plesiomorphic for Aetana. A scape is here interpreted to have evolved convergently

in ‘Gen.n. Borneo’ and within Aetana, uniting the Aetana ocampoi and A. kiukoki groups. A deeply

indented posterior margin is a synapomorphy of three species within the A. omayan group.

36. Epigynum scape, shape: (0) wide; (1) narrow. In the Aetana ocampoi and A. kiukoki groups, a wide

scape (Figs 16, 60, 78) is plesiomorphic, and a narrow scape (Figs 28, 43) unites A. libjo Huber,

sp. nov. and A. baganihan Huber, sp. nov.. In ‘Gen.n. Borneo’ the scape originated as a narrow scape.

37. Epigynum external pair of pockets: (0) absent; (1) present. A pair of pockets on the epigynal plate

occurs in the outgroup taxa Zatavua and Spermophora , but not in Aetana or other outgroup taxa.

European Journal of Taxonomy 162: 1-78 (2015)

38. Female internal genitalia, lateral membranous pockets: (0) absent; (1) present. This pair of pockets,

that is only visible in cleared preparations (Figs 112, 117, 129, 149, 154), is a synapomorphy of the

Aetana kinabalu group.

39. Female internal genitalia, median membranous structure: (0) absent; (1) present. This unique

structure (Figs 29, 44) unites two species of the Aetana ocampoi group: A. libjo Huber, sp. nov. +

A. baganihan Huber, sp. nov..

40. Female internal genitalia, sclerotized pockets: (0) absent; (1) present. Sclerotized internal pockets

originating from the ventral wall of the uterus externus (Figs 111, 153, 192, 194) occur in the Aetana

kinabalu group and in the A. omayan group, but are here interpreted to have evolved three times

convergently.

41. Female internal genitalia, serrated ridges: (0) absent; (1) present. These highly distinctive and unique

ridges (Figs 61, 73, 79) are a synapomorphy of the Aetana kiukoki group.

42. Pair of pockets behind epigynum: (0) absent; (1) present. A membranous and poorly visible pair of

pockets in the area behind the epigynum (Figs 192-194, 222, 227, 232) is a synapomorphy of the

Aetana omayan group.

43. Pair of pockets behind epigynum, distance: (0) wide apart; (1) close together. Within the Aetana

omayan group, two species are characterized by pockets that are far apart (292-294), and three

species by pockets that are close together (222, 227, 232). The plesiomorphic condition is unclear,

as is the condition in A. fiji.

Ecology

44. Microhabitat: (0) close to ground; (1) higher among the vegetation. Most Aetana species live well

protected in dark spaces under rocks and logs. A few species occur higher among the vegetation, with

their webs among twigs and leaves and often reached by direct sun. This has apparently evolved at

least twice, once in A. libjo Huber, sp. nov. + A. baganihan Huber, sp. nov., and once in A. kinabalu

+ A. lambir Huber, sp. nov.. A third independent case might occur in A. fiji , but the available data

(sparse label information) are not conclusive.