European Journal of Taxonomy 26: 1-11

http://dx.doi.org/10.5852/ejt.2012.26

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2012 • Joeque R., Bosselaers J. & Henrard A.

ISSN 2118-9773

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Research article

um:lsid:zoobank.org:pub:BB26F2C9-AC4E-4941-9A34-7CDC5ECFB63F

A new species of Hortipes (Araneae, Corinnidae), the first spider with

an insertable retrolateral tibial apophysis on the male palp

Rudy JOCQUE', Jan BOSSELAERS^ and Amaud HENRARD^

h2TRoyal Museum for Central Afriea, B 3080 Tervuren, Belgium.

Email: [email protected]

^Earth and life Institute, Biodiversity researeh Center, UCE-I7.07.04,

Batiment Camoy, Croix du Sud, 5, B 1348, Eouvain-la-Neuve, Belgium.

^ um:lsid:zoobank.org:author:CF15016C-8CDl-4C9D-9021-44CA7DC7A5D5

^ um:lsid:zoobank.org:author:D6AD7414-3540-4F06-8631-8873450AA90C

^ um:lsid:zoobank.org:author:ElB02E6E-D91C-43FE-8D8C-CD102EFEE3B4

Abstract. Hortipes gigapophysalis (Araneae, Corinnidae) is a new speeies deseribed from both sexes

from montane forest on Mt Nimba, eastern Guinea. The speeies is remarkable for its long, whip-shaped

retrolateral tibial apophysis (RTA) on the male palp. The strueture apparently has an insertable fimetion

as the epigyne of the female eontains a separate set of duets starting from a eentral eoneavity that is

unique in the genus. This duet system is apparently meant to reeeive the supple RTA. This type of

struetural arrangement has never previously been found in spiders.

Keywords, genitalia, morphology, Corinnidae, Guinea, Mt Nimba.

Joeque R., Bosselaers J. & Henrard A. 2012. A new species of Hortipes (Araneae, Corinnidae), the first spider with

an insertable retrolateral tibial apophysis on the male palp. European Journal of Taxonomy 26: 1-11. http://dx.doi.

org/10.5852/eit.2012.26

Introduction

With almost 70 known speeies, Hortipes Bosselaers & Eedoux, 1998 is one of the largest Afrotropieal

genera (Bosselaers & Joeque 2000a). Most of the speeies live in the litter layer of dense forests and

woodlands, henee its very wide distribution from West Airiea to Ethiopia in the north and to large parts

of South Afriea in the south. However, the genus is entirely absent from miombo woodland and is

therefore an exeellent example of an Airotropieal spider genus with this type of distribution. The genus

is also remarkable as an example of the range of eomplexity of genitalia that is met with in many spider

genera. In Hortipes, the genitalia range from quite simple struetures as in H. silvarum Eedoux & Emerit,

1998 to eomplex eonfigurations sueh as in H sceptrum Bosselaers & Joeque, 2000a. The genus ean

therefore be eonsidered as suitable for testing the evolution of genitalia in the eontext of sexual seleetion

hypotheses (e.g. Eberhard & Huber 1998; Huber 1995a; Joeque 2002).

The present paper deseribes a new speeies in the genus that has a remarkable male palp provided with a

very long retrolateral tibial apophysis (RTA) unlike any that has been found in spiders so far.

European Journal of Taxonomy 26\ 1-11 ( 2012 )

Material and Methods

Specimens were observed and measured with a Leica MIO stereomicroscope. Photographs of the

habitus were taken with a Leica MZ16 using the LAS automontage software. The female epigyne was

detached from the abdomen, temporarily mounted in a clearing mixture of methyl salicylate and cedukol

(Merck, Darmstadt) and observed with a Leitz Dialux 22 microscope and subject to automontage with

the Syncroscopy software. For SEM photos, a palp was dried in HMDS, gold coated and examined

and photographed with a JEOL 6480 LV scanning electron microscope. The format for leg spination

follows Platnick & Shadab (1975), amended for ventral spine pairs according to Bosselaers & Jocque

(2000b). The spine row sequences run from the base of segments towards the tip. Leg spine numbers

between brackets refer to spines present in some instances and absent in others. All measurements are

in mill im eters.

A phylogenetic analysis was performed under implied weighting (Goloboff 1993) using the computer

programmes PAUP 4.0 beta 10 (Swofford 2002), and TNT 1.1 (Goloboff et al. 2003, 2008) run on a

dual-core Intel iMac under a Windows XP virtual machine (VMware Fusion 2, VMware, Inc., 3401

Hillview Avenue, Palo Alto, CA 94304, USA; Bugnion et al. 2000). For this purpose, the characters and

data from Bosselaers & Jocque (2000a: 11-24) were used, with scores added for H. gigapophysalis (1:1,

2:1, 3:1, 4:1, 5:1, 6:1, 7:2, 8:2, 9:0, 10:0, 11:1, 12:0, 13:1, 14:0, 15:0, 16:0, 17:0, 18:0, 19:0, 20:0, 21:0,

22:1, 23:1, 24:1, 25:1, 26:0, 27:1, 28:1, 29:1, 30:1, 31:1, 32:0, 33:?, 34:0, 35:?, 36:0, 37:0, 38:2, 39:3,

40:1, 41:2, 42:0, 43:0, 44:0, 45:1, 46:?, 47:?, 48:?, 49:?, 50:?, 51:0, 52:1, 53:0, 54:7, 55:?, 56:?, 57:?,

58:?, 59:?, 60:0, 61:0, 62:0, 63:0, 64:0, 65:0, 66:0, 67:0, 68:1, 69:0, 70:0, 71:0, 72:0, 73:0, 74:1, 75:0,

76:1, 77:0, 78:0, 79:0, 80:0, 81:1, 82:0, 83:2, 84:1, 85:1, 86:1, 87:0, 88:0, 89:0, 90:2). Moreover, the

scoring for character 44, which had been inadvertently inverted in Bosselaers & Jocque (2000a: table 1),

was rectified, and an extra character state was added to character 54 in order to score H. gigapophysalis.

Bremer support values (Bremer 1988, 1994) for the preferred consensus tree, expressed as fit values,

were calculated in TNT using Analyze / suboptimal, followed by Analyze / Traditional search / tree

bisection reconnection (TBR) and Trees / Bremer Supports, retaining trees suboptimal up to 10 units

of fit (TNT values, see Jocque & Bosselaers 2011: 274) and combining various numbers of replications

(between 1 and 100) with various numbers of trees saved per replication (between 200 and 20000,

inversely related to the numbers of replications) until the solution stabilized.

Abbreviations

DTA

HMDS

MRAC

PAUP

pdv

plv

rlv

RTA

carapace length

carapace width

dorsal

distal tegular apophysis

dorso-ventral stretch of secundary tube system

embolus

hexamethyldisilazane (CAS 999-97-3)

median apophysis

Royal Museum for Central Africa

metatarsus

phylogenetic analysis under parsimony

position of dorsoventral stretch

prolateral

prolateral ventral

retrolateral

retrolateral ventral

retrolateral tibial apophysis

JOCQUE R., BOSSELAERS J. & HENRARD A., A new Hortipes (Araneae) with an insertable RTA

tibia

total length

ventral terminal

Results

Class Araehnida Cuvier, 1812

Order Araneae Clerek, 1757

Family Corinnidae Karseh, 1880

Genus Hortipes Bosselaers & Eedoux, 1998

Hortipes gigapophysalis sp. nov.

um:lsid:zoobank.org:aet:199C827C-56D9-4F42-AlQ3-FQ63ACAQQ33F

Figs 1-5

Diagnosis

Males of H. gigapophysalis are easily reeognized by the long, winding RTA; the female is eharaeterized

by the eentral depression and the separate duet systems in the epigyne. These eharaeters are unique in

the genus.

Etymology

The speeies is ealled gigapophysalis in view of the extraordinarily long retrolateral tibial apophysis on

the male palp.

Type material

Holotype

(? (MRAC 239507): GFTNEA, Mt. Nimba, Station de pompage Zie, alt. 1250 m, 07° 40’ N - 008° 22’ W,

03 Feb. 2012, hand eolleeting by A. Hemard, C. Allard, R Bimou and M. Sidibe near Zie river, forest.

Paratypes

2 SS (MRAC 239104) and, together with holotype, 1 $ (MRAC 239482): GUINEA, Mt. Nimba, Pierre

Riehaud, Kaiser 1, alt. 1480 m, 07° 39’ N - 008° 22’ W, 7 Get. 2011, hand eolleeting by A. Henrard and

D. Vandenspiegel, at the entranee (± 4 m inside) of disused humid adit, under wood log.

Description

Male

Measurements. Total length 2.21 earapaee 1.07 long, 0.87 wide; length of fe: I 0.89, II 0.92, III 0.72,

IV 1.07.

Feg spination. Fe: I plv I-I-I rlv I-I-I; II plv I-I-I rlv I-I-I; ti: I plv I-I-I-I-I-I rlv I-I-I-I-I-I; II plv

rlv l-l-l-l-l-l; IV pi 0-1-0 rl 0-1-0; mt: I plv 1-1-1 rlv 1-1-1; II plv 1-1-1 rlv 1-1-1; IV pi

1-0-1 rl 1-0-1 vt 2.

CoEORATiON. Carapaee and legs yellow, with an orange tinge; sternum pale yellow, ehelieerae, labium

and endites yellow. Abdomen pale. (Fig. lA-C).

Paep (Figs 2A-B, 4A-F). Tibia with eentral eonstrietion. RTA very long, slender and eoiled, narrowing

from base to tip; eymbium subeireular, proximally with deep retrolateral fold, with few straight setae

along retrolateral margin and group of ehemosensory setae at distal dorsal tip; sperm duet appears at

base of embolus, runs along retrolateral, posterior and prolateral margins of tegulum, fairly broad over

European Journal of Taxonomy 26\ 1-11 ( 2012 )

Fig. 1. Hortipes gigapophysalis sp. nov. A. Male habitus, dorsal view. B. Male prosoma, ventral view.

C. Male habitus, lateral view. D. Female habitus, dorsal view. E. Female prosoma, ventral view (epigyne

removed). F. Male prosoma, ventrolateral view.

JOCQUE R., BOSSELAERS J. & HENRARD A., A new Hortipes (Araneae) with an insertable RTA

Fig. 2. Hortipes gigapophysalis sp. nov. A. Male palp, lateral view. B. idem, ventral view. C. Epigyne,

ventral view. D. Epigyne, eleared, dorsal view, seeond system omitted. E. idem, primary system omitted.

F. idem, both systems eombined. E: embolus, entl: entranee of insemination duet, ent2: entranee of

seeondary duet system, dv: dorso-ventral streteh of seeondary duet system, pdv: position of dorso-

ventral streteh, RTA: retrolateral tibial apophysis, stl: spermatheeae 1, st2: spermatheea 2. Seale bars:

100 pm.

European Journal of Taxonomy 26\ 1-11 ( 2012 )

entire eourse exeept last quarter strongly narrowed before entering E; E originating at distal end of

tegulum direeted retrolaterad, very long, whiplike, looped over slightly more than some 500°. True MA,

being an artieulated tegular proeess inserted on a flexible membrane (Bonaldo 1997: 166) absent, but

tegulum distally with membranous, translueent DTA, direeted retrolaterad, eurved baek over 90° ending

near posterior margin of eymbium.

Female

Measurements. Total length 2.54; earapaee 1.15 long, 0.85 wide; length of fe: 1 0.95,11 1.00, 111 0.81,

IV 1.11.

Eeg spination. Fe: 1 plv l-l-l-(l) rlv 1-1-1; 11 plv 1-1-1 rlv l-l-l-l-(l); ti: Palp do 1-0-0; 1 plv l-l-l-l-

1-1 rlv l-l-l-l-l-l; 11 plv l-l-l-l-l-l rlv l-l-l-l-l-l; mt: 1 plv 1-1-1 rlv 1-1-1; 11 plv 1-1-1 rlv 1-1-1;

IV pi 0-0-1 rl 0-0-1 vt 2.

CoEORATiON. Carapaee, legs and ehelieerae pale orange; sternum pale. Abdomen pale (Fig. ID-F).

Genitaeia. Epigyne with a eentral eoneavity and with eonvoluted tubes visible in transpareney. Two

funnel-shaped openings leading into a system of wide, very simple, thin-walled tubes, with blind ends

(Fig. 2C-F). Thiek-walled insemination duets starting from slit-shaped lateral openings leading to two

sets of spermatheeae. Anterior pair oval, opening posteriorly into duets leading towards posterior,

globular pair. The systems are separate and there is no eonneetion between them (Figs 2C-F, 3A-B).

Variability

Male paratypes (n=2) TF: 2.16 and 2.03; CF 1.10 and 0.89; CW 0.89 and 0.85. Spination identieal.

Distribution

High altitude gallery forest (> 1200 m) on Mt Nimba, Guinea.

Fig. 3. Hortipes gigapophysalis sp. nov. A. Epigyne eleared in methyl salieylate, ventral view. B. idem,

dorsal view.

JOCQUE R., BOSSELAERS J. & HENRARD A., A new Hortipes (Araneae) with an insertable RTA

Discussion

The RTA is usually a solid, strongly selerotized stmeture. Its main fimetion was long thought to be

internally stabilising the expanded male palp during eopulation, whieh means that the RTA loeks either

speeifie selerites of the bulbus or between the bulbus and eymbium in order to arrest further rotation or

movement (Huber 1995 a: 151; Sierwald & Coddington 1988: 264). However, the RTA has been found

to fulfill many different funetions. Huber (1994) mentions four different fimetions for the RTA in one

Fig. 4. Hortipes gigapophysalis sp. nov. A. Male palp, retrolateral view. B. Male palpal patella and

tibia, dorsal view. C. Male palp, ventral view. D. idem, dorsal view. E. Male palpal tibia and patella,

dorsolateral view. F. Detail of previous. E: embolus, DTA: distal tegular apophysis, RTA: retrolateral

tibial apophysis.

European Journal of Taxonomy 26\ 1-11 ( 2012 )

single spider family (Agelenidae). In Textrix denticulata (Olivier, 1768), the RTA fixes the male palp

to the female epigyne; in Histopona torpida (C.L. Koeh, 1834) the role of the RTA is unelear, sinee the

patellar apophysis arrests the male bulbus internally; in Agelena gracilis and Agelenopsis spp. the RTA

loeks the male palp internally in two different ways, as deseribed below (Gering 1953; Osterloh 1922).

Jager (2006: 58) eonfirms this diversity in funetion, showing that the RTA anehors in the epigastrie

furrow in Sparassidae but anehors in the epigyne in Gnaphosidae. Huber (1995a) points out that the

funetion of the RTA is highly speeies speeifie and may be involved in mate ehoiee by physieal eontaet

during eopulation (1995a: 152), rather than internally stabilising the male palp. Huber (1995a, 1995b:

698, fig. 5) demonstrated that the RTA is used to fix the male pedipalp to the female epigyne for six

speeies from six different families of the RTA-elade (Coddington & Levi 1991). Huber (1995a: 159-

160) also states that most data inferring that the RTA arrests the male bulb internally during eopulation

are doubtful for methodologieal reasons. Either the authors used artifieially expanded genital bulbs,

or they studied genital bulbs fixed in eopula, but sprung away from the epigyne. The vast majority of

researeh using reliable methodology (observations of animals in copula, seetions of animals fixed in

copula) led to the eonelusion that the RTA fixes the male pedipalp to the female externally. Sueh fixation

to the female epigyne may be neeessary beeause of the laek of innervation of the male bulbus (Eberhard

& Huber 1998).

Huber eites the studies of Osterloh (1922) and Gering (1953) as exeeptions to this generalisation, and

lists them in his table 1 as an RTA that arrests the genital bulb internally in Agelena gracilens C.E. Koeh,

1841 and Agelenopsis spp. Giebel, 1869, respeetively. More speeifieally, Osterloh (1922: 348) states that

the RTA is inserted between the palpal bulbus (“Kapsel”) and the palpal terminal selerites (“Sterna”)

in eopula, and that only the terminal selerites fAlles, was zum Sterna gehorf) are introdueed into the

Coryssiphus

— Andromma

7.69?

1.75

0.45

silvarum

gigapophysalis

—salticola

0.18

0.62

0.22

0.18

0.22

0.10

0.25

fastigiensis

£— pollux

castor

0.24

ostiovolutus

exoptans

ly— scharffi

leno

0.30

cucurbita

0.11

aurora

0.91

0.30

libidinosus

delphinus

orchatocnemis

oronesiotes

contubernalis

— schoemanae

0.18

0.11

0.08

0.29

0.05

creber

zombaensis

merwei

coccinatus

_I-wimmertensi

0 27 - luytenae

0.30

hastatus

0.46

0.31

0.03

1.04

0.29

0.38

lejeunei

I- narcissus

' -falcatus

tarachodes

machaeropolion

architelones

robertus

— bosmansi

I-sceptrum

0 25 -hormigricola

0.12

0.37

Fig. 5. Cladogram obtained from the eharaeter matrix in Bosselaers & Joeque (2000) with inelusion of

H. gigapophysalis. Striet eonsensus of 18 trees of fit -53.061 (PAUP values), obtained under implied

weighting. Bremer support values are shown under the branehes.

JOCQUE R., BOSSELAERS J. & HENRARD A., A new Hortipes (Araneae) with an insertable RTA

epigyne. Eocking of the male palpus to the epigyne in these two speeies is performed by the patellar

apophysis, not the RTA (1922: 390). Gering (1953: 13) states: ''The ectoproximal margin of the cymbium

has a weakly produced depression, which normally is somewhat more heavily pigmented and sclerotized

than the surrounding area (figs 11-12, 39). The tibial process rests in this depression when the palpus

is locked. We ean eonelude that neither author elearly advoeates that the RTA is introdueed into the

internal female genital organs. In Gnaphosidae the RTA, a “primary anehoring deviee to the epigynal

poekef ’ (Senglet 2004: 87) does not take part in intromission, whieh takes plaee after the palpus is fixed

(Senglet 2004: 91).

However, in the present eontribution, an RTA is deseribed whieh has exaetly this very unusual fimetion:

intromission into the female genital organs. To the best of our knowledge, sueh a use of the RTA has

never been mentioned before. As the RTA of H. gigapophysalis sp. nov. is very long, thin and supple, it

is very unlikely that it is used as a rigid loeking deviee as deseribed above. Its analogy with the embolus,

a long whiplike, supple strueture, made us suspeet that it might be an insertable selerite. Detailed

study of the epigyne indeed reveals the presenee of two independent tubular systems. The first is the

usual insemination duet starting at a lateral slit-like entranee and leading to two sets of spermatheeae

(Fig. 2C, D). The seeond is eomposed of wide, very simple, thin-walled tubes starting from funnel-

shaped entranees in a eentral eoneavity, ending blind at the end of several bends without eonneetion with

the first system (Fig. 2E, F). It is assumed that the seeond system is meant to reeeive the RTA.

Hortipes is a genus with an amazing genitalie variability. Nevertheless, not a single speeies with an RTA

that is elearly insertable in the female genitalia has been deseribed to date. Among the known speeies

(Bosselaers & Joeque 2000a; Eedoux & Emerit 1998), only a few speeies of the orchatocnemis elade

(node 12 in Bosselaers & Joeque 2000a, fig. 4) have vulvae that, in view of their weakly selerotised

entranee ehamber (Bosselaers & Joeque 2000a: 14, eharaeter 65) show possible evidenee of additional

selerites being inserted during eopulation. H. merwei Bosselaers & Joeque, 2000, for example, has an

elaborate lobed entranee ehamber (Bosselaers & Joeque 2000a: fig. 20e) that seems espeeially suited for

insertion of the exeeptionally large, fiexible and eoiled MA of the male palp (Bosselaers & Joeque 2000a:

fig. 19a, b). Jager (2012: figs 19, 23) deseribes Sinopoda vulvae with similar thin-walled saes. In this

genus, these saes proved to be expandable, but Jager does not mention any role for them. The entranee

ehamber of H. aelurisiepae Bosselaers & Joeque, 2000, also has peeuliar thin-walled lobes, lateral in

this ease, that might reeeive an additional palpal selerite from the unknown male. However, none of the

Hortipes speeies deseribed to date have an RTA of eonsiderable length that matehes a vulvar strueture

in whieh it eould be inserted. Moreover, as a result of the faet that the Hortipes speeies of western

Afriea are very poorly sampled (Bosselaers & Joeque 2000a: map 1), no intermediate stages are known

between the genitalia of the speeies already deseribed and the insertable RTA of H. gigapophysalis

sp. nov. Based on leg spination and genitalie strueture, H. gigapophysalis sp. nov. seems elosest to

H. hesperoecius Bosselaers & Joeque, 2000 and//, silvarum. Repeating the eladistie analysis of Hortipes

speeies known from both sexes (Bosselaers & Joeque 2000a), ineluding H. gigapophysalis sp. nov.,

eonfirms this (Fig. 5): H. gigapophysalis sp. nov. turns out to be the sister speeies of H. silvarum and is

situated at the base of the Hortipes elade in the striet eonsensus of 18 trees of fit -53.061 (PAUP values),

obtained under implied weighting (Fig. 5). In the striet eonsensus of 1656 shortest trees of length 265

obtained under equal weighting, the salticola elade (node 3 in Bosselaers & Joeque 2000a, fig. 4) holds

a still more basal position within Hortipes, but the sister speeies relationship between H. silvarum and

H. gigapophysalis is eonfirmed. However, although H. silvarum has a relatively slender and apieally

eoiled RTA (Bosselaers & Joeque 2000a: fig. 12e, f), it has no vulvar tubes in whieh the RTA eould be

inserted (Bosselaers & Joeque 2000a: fig. 13d). Only more extensive sampling of the forest litter layer

of tropieal western Afriea may produee the as yet unknown Hortipes speeies allowing to reeonstruet the

phytogeny of this most unusual genitalie development.

European Journal of Taxonomy 26\ 1-11 ( 2012 )

Acknowledgements

We are indebted to Alain Reygel for the drawings of the genitalia. We thank P. Jager and an anonymous

referee for useful eomments on a previous version. This researeh was supported by Golder Assoeiates and

sub-eonsultants as a eomponent of the Nimba Projeet ESIA for Soeiete des Mines de Per de Guinee. This

paper is publieation BRC 277 of the Biodiversity Researeh Center (Universite eatholique de Louvain).

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Manuscript received: 8 September 2012

Manuscript accepted: 20 October 2012

Published on: 21 October 2012

Topic editor: Koen Martens

In eomplianee with the ICZN, printed versions of all papers are deposited in the libraries of the institutes

that are members of the EJT eonsortium: Museum National d’Histoire Naturelle, Paris, Franee; National

Botanie Garden of Belgium, Meise, Belgium; Royal Museum for Central Afriea, Tervuren, Belgium;

Natural History Museum, Eondon, United Kingdom; Royal Belgian Institute of Natural Seienees,

Brussels, Belgium; Natural History Museum of Denmark, Copenhagen, Denmark.