European Journal of Taxonomy 82: 1-20

http://dx.doi.org/10.5852/ejt.2014.82

BY

ISSN 2118-9773

www. europeanj oumaloftaxonomv. eu

2014 • Henrard A., Jocque R. & Baehr B.

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

Research article

urn:lsid:zoobank.org:pub:BA6707Bl-9D AD-4448-9431-47B7E9B63980

Redescription of Tapinesthis inermis (Araneae, Oonopidae),

with detailed information on its ultrastructure

Arnaud HENRARD 125 *, Rudy JOCQUE 16 & Barbara C. BAEHR 347

1 Section Invertebrates non-insects. Royal Museum for Central Africa,

Leuvense Steenweg 13, B-3080 Tervuren, Belgium

* Corresponding author. E-mail: [email protected]

2 Earth and Life Instititute, Biodiversity Research Center, Universite Catholique de Louvain,

Place Croix du Sud 1—4, B-1348 Louvain la Neuve, Belgium

3 Queensland Museum, PO Box 3300, South Brisbane, QLD 4101, Australia

4 CSER, School of Environmental and Life Sciences, University of Newcastle,

Callaghan, NSW 2308, Australia

5 urn:lsid:zoobank.org:author:ElB02E6E-D91C-43FE-8D8C-CD102EFEE3B4

6 urn: lsid: zoobank.org: author: CF15016C-8CD1-4C9D-9021-44C A7DC7A5D5

7 um:lsid:zoobank.org:author:CFAFC574-5691-4A77-AF0C-E6B8B0563340

Abstract. Tapinesthis inermis Simon, 1882, the only species in the genus, is widely distributed in western

Europe. This redescription provides the first information on the ultrastructure of the species using SEM.

The morphology of the spinnerets, tarsal claws and tarsal organs, and the internal structure of the female

genitalia and the male palp are described and illustrated in detail. The combination of these structures

is very similar to those encountered in some dysderoid spiders and supports the basal placement of

Tapinesthis among Oonopinae. The phylogenetic relationships of the species are discussed. The only

female among the three syntypes is designated as the lectotype.

Key words. Female genitalia, male palp, organ receptors, spinnerets, systematics

Henrard A., Jocque R. & Baehr B. 2014. Redescription of Tapinesthis inermis (Araneae, Oonopidae) with detailed

infonnation on its ultrastructure. European Journal of Taxonomy 82: 1-20. http://dx.doi.org/10.5852/eit.2014.82

Introduction

The spider genus Tapinesthis Simon, 1914 is monospecific, containing the single species T. inermis

(Simon, 1882), widely distributed in Europe (Le Peru 2011; Platnick 2013). This rare species is often

found in houses or in the neighbourhood of habitations (Kraus 1967; Heimer & Nentwig 1991; Lazarov

et al. 2001; Korenko et al. 2007; Van Keer et al. 2006). Tapinesthis inermis thus appears to be strongly

synanthropic, as it is often found in urban areas. Saaristo & Marusilc (2009) even employed the term

“hemisynanthropic” to describe its habitat preference. However, it has also been collected in natural

habitats such as forests (Dalmas 1920; Le Peru 2007), garrigues and dry grasslands (Le Peru 2011) or

near cliffs with a dense cover of ivy (e.g., Rochers de Marches les Dames in Belgium, this study). In

Belgium, T. inermis appears to be abundant in litter and among ivy (Van Keer et al. 2006; pers. obs.).

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European Journal of Taxonomy 82: 1-20 (2014)

The species is often found together with Oonops domesticus and O. pulcher (Van Keer et al. 2006,

2010), and, at least in Belgium, also with Harpactea hombergi (Scopoli, 1763) (pers. obs.).

Tapinesthis inermis is easy to identify and has been reviewed by many authors (see Platnick 2013).

Originally, the species was described as Oonops inermis and only the female was known (Simon 1882).

Later, Simon (1914) created the genus Tapinesthis for this species and described both sexes.

A remarkable redescription of both males and females was provided by Kraus (1967), with detailed

drawings of the genitalia and other body parts. Other significant illustrations and photos have been

published by Korenlco et al. (2007) (female prosoma, vulva) and by Saaristo & Marusilc (2009) (male

and female habitus, male palp, female genitalia). However, this is the first detailed morphological study

of T. inermis using SEM images, providing detailed information about the ultrastructure of the female

genitalia, the spinnerets, mouthparts and legs. We also compare the genitalia with those of related taxa

and discuss the relationships of this species with other oonopids.

The results of this work have previously been used for the polarization of characters in a phylogenetic

study of Afrotropical Orchestina Simon, 1882 (Henrard & Jocque 2012). However, one particular

character, the number of tarsal organ receptors, is updated here and the consequences are dealt with in

the discussion.

Material and methods

Specimens were observed with a Zeiss Stemi 2000 stereo microscope. Measurements and photographs of

the habitus and details of the mouthparts and detached male palps and genitalia were taken with a Leica

MZ16 using the LAS automontage software (ver. 3.8). Female genitalia were dissected, digested with

pancreatin and then immersed in 75% ethanol. One male palp was detached and temporarily mounted in

a clearing mixture of methyl salicylate and cedulcol (Merck, Darmstadt), observed with a Leitz Dialux

22 microscope and subjected to automontage with the Syncroscopy software (ver. 5.03.61). For SEM

photos, specimens were dried in HMDS (36 h), gold coated, and examined and photographed with a

JEOL 6480 LV scanning electron microscope. All measurements are in mill im etres

This paper follows the terminology of Burger’s series of papers (Burger 2009, 2011a, 2011b, 2013;

Burger & Michalik 2010) with a few adaptations.

The following abbreviations are used:

ALS

= anterior lateral spinnerets

ALE

= anterior lateral eyes

ARe

= anterior receptaculum

AS

= anterior sclerite

D

= duct

Ht

= holotype

FA

= flattened apodeme

MNHN

= Museum national d’Histoire naturelle, Paris (C. Rollard)

MRAC

= Musee Royal de l’Afrique Centrale, Tervuren (R. Jocque)

PBI

= Planetary Biodiversity Inventory project

PER

= posterior eye row

PLE

= posterior lateral eyes

PLS

= posterior lateral spinnerets

PME

= posterior median eyes

PMS

= posterior median spinnerets

Pp

= posterior plate

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HENRARD A., JOCQUE R. & BAEHR B., Redescription of Tapinesthis inermis

Pr

protrusion

PRe

posterior receptaculum

Pt

paratype

RBINS =

Royal Belgian Institute of Natural Sciences, Bmssels

V

vesicle

Te

tendon

Tw =

thin-walled duct

Results

Class Arachnida Cuvier, 1812

Order Araneae Clerck, 1757

Family Oonopidae Simon, 1890

Tapinesthis Simon, 1914

Diagnosis

Soft-bodied, pale Oonopidae with strongly sloping cephalothorax provided with dark net-shaped pattern,

without leg spines, with unipectinate tarsal claws; pedicel with meshed texture; male palp with short

sclerified sperm duct and female genitalia with wide median T-shaped structure visible by transparency,

flanked by two rounded apodemes.

Description

Male

Cephalothorax. Carapace in vivo pale yellowish-orange (Fig. 1A-B) to p ink ish, in ethanol yellowish-

white to pale orange (Figs 2A-B, 3A-D), with dark net-shaped pattern and dark margins; non-marginal

pars cephalica setae present in three rows, ovoid in dorsal view, pars cephalica strongly sloping in lateral

view, reaching highest point at level of second coxa (Fig. 2E), anteriorly narrowed to 0.49 times its

maximum width or less, with rounded posterolateral corners (Fig. 2A-B), surface of elevated portion

of pars cephalica smooth, sides smooth, with radiating patches of smooth platelets (Fig. 6A-B); thorax

without depressions, fovea absent, lateral margin undulate, smooth; pars cephalica setae needle-like.

Clypeus margin curved downwards in front view (Fig. 2G), sloping forward in lateral view (Fig. 2E), high,

AFE separated from edge of carapace by their radius or more. Eyes (Fig. 2A, E): six, well developed, all

subequal, AFE oval, PME circular, PEE circular; posterior eye row recurved from both above and front;

AFE separated by their radius to diameter, AFE-PFE separated by less than AFE radius, PME touching,

PFE-PME separated by less than PME radius, AFE-PME separated by less than PME radius. Sternum

(Fig. 2C) yellowish white, with faint radiating dark stripes, longer than wide, with radial smooth furrows

between coxae I—II, II—III and HI—IV, anterior margin with semicircular depression on the middle half,

posterior margin extending posteriorly beyond anterior edges of coxae IV as single extension, distance

between coxae approximately equal; setae abundant, evenly scattered, without hair tufts. Mouthparts:

chelicerae, endites and labium yellowish white. Chelicerae (Fig. 6F) straight; without teeth on both

promargin and retromargin; fangs directed medially, paturon inner margin with scattered setae. Fabium

(Figs. 2C, 6C-D, F) rectangular, fused to sternum, anterior margin indented at middle, basal comers

with small circular depression, sclerotization as in sternum; with 6 or more setae on anterior margin,

subdistal portion with unmodified setae. Endites (Fig. 6C-G) distally not excavated, serrula present in

single row, anteromedian tip unmodified, posteromedian part unmodified, sclerotization as in sternum;

anterior margin with ventral row of spatulate setae and dorsal row of distally pectinate setae, median

margin with finely barbed setae.

Abdomen (Fig. 2A-B, D-F). Ovoid, rounded posteriorly,without scuta; dorsum soft portions yellow-

brown, without color pattern. Dorsum setae present, needle-like. Epigastric area setae uniform, needle-

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European Journal of Taxonomy 82 : 1-20 ( 2014 )

like. Postepigastric area setae present, needle-like. Pedicel with meshed texture (Fig. 6C) as in Orchestina

(see Henrard & Jocque 2012, figs. 21, 113, 114, 404), abdomen extending anteriad of pedicel. Colulus

present (Fig. 9A), with four setae. Spinnerets (Fig. 9B-D): ALS with one major ampullate gland spigot

emerging from cylindrical tubercle and two piriform gland spigots emerging from shallow tubercle;

PMS with one minor ampullate gland spigot emerging from cylindrical tubercle; PLS with two ac ini form

gland spigots emerging from cylindrical tubercle.

Legs. White, without color pattern; femur IV not thickened, same size as femora I—III, patella plus

tibia I longer than carapace, tibia I unmodified, metatarsi of at least leg II in the middle and IV at basal

quarter, with eye-shaped smooth gland outlet with single pore (Fig. 12C-E). Leg spines absent. Tarsal

claws (Fig. 10A-F) unipectinate, tarsal proclaws and retroclaws inner face smooth; tarsus I-IV superior

claws with six teeth on lateral surface of proclaw, six teeth on lateral surface of retroclaw. Tarsi claw

tuft with three carpeted setae. Tarsi I to IV without inferior claw. Trichobothria (Fig. 12D-E) with

rounded bothrium, internal texture of aperture not grate-like, hood covered by numerous low, closely

spaced ridges; sometimes accompanied by eye shaped structure with unknown function (Fig. 12C-D).

Tarsal organ (Fig. 11A-D, FT—I) pear-shaped, margin raised above surroundings, with 4 receptors on

legs I and II, with 3 receptors on legs III and IV. In a probably unusual case, the tarsal organ has only 1

receptor on both legs IV (Fig. 11H-I), whereas I—III of the same specimen have the usual 4-4-3 formula

(not illustrated). According to observations made on other oonopids the formula appears to be 3-3-2-2,

since the distalmost receptor is distally bifid, but both parts stem from a common base as in Stenoonops

Simon, 1891 (Platnick & Duperre 2010).

Fig. 1. Tapinesthis inermis photographed in vivo. A. $ (PBI 33564). B. Same. C. 5 (PBI 33267).

D. Same. E. Same. Remark color differences of cephalothorax between $ and §. For explanation, see

text.

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HENRARD A., JOCQUE R. & BAEHR B., Redescription of Tapinesthis inermis

Fig. 2. Tapinesthis inermis. A. $ (PBI 33564), habitus, dorsal view. B. Same, carapace. C. Same,

sternum, ventral view. D. $ (PBI 33267), habitus, dorsal view. E. $ (PBI 33564), habitus, lateral view.

F. Same, $ (PBI 33267). G. $ (PBI 08123), carapace, anterior view. Scale bars = 0.5 mm.

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European Journal of Taxonomy 82 : 1-20 ( 2014 )

Genitalia (Figs. 4A-E, 7A-D). Epigastric region with sperm pore small, narrow, slit-like, situated at

level of anterior spiracles. Palp normal size, not strongly sclerotized, right and left palps symmetrical,

proximal segments white; trochanter normal size, unmodified; femur normal size, two or more t im es

as long as trochanter, without posteriorly rounded lateral dilation, attaching to patella basally; patella

shorter than femur, not enlarged, without prolateral row of ridges, setae unmodified; tibia ventral margin

swollen; cymbium white, ovoid in dorsal view, not fused with bulb, not extending beyond distal tip of

bulb, plumose setae absent, without stout setae, without distal patch of setae; bulb white, 1 to 1.5 times

as wide as cymbium, stout, piriform with frontal margin slightly concave, tapering apically with long,

medially bent embolus, distal sclerotized part of seminal duct (D) short, restricted to distal part, abruptly

interrupted, internally with poorly defined vesicle (V), opening into embolus through duct with basal

part thin-walled (Tw), distal part sclerotized; tendon (Te) running from base of cymbium to distal end

of vesicle (V); embolus simple, dark, tube-shaped, flattened apically, tip blunt, grooved, appearing pale.

Female (Figs 1C-E, 2D, F-G)

As male except as noted.

Cephalothorax. Female palpal claw absent; spines absent. Tarsal organs (Fig. 11A-D) as in male.

Fig. 3. Tapinesthis inermis , $ from the Czech Republic (PBI 32902). A. Habitus, dorsal view. B. Same,

lateral view. C. Same, lateral view. D. Prosoma, ventral view.

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HENRARD A., JOCQUE R. & BAEHR B., Redescription of Tapinesthis inermis

Abdomen. Spinnerets (Fig. 9E-H): ALS with one major ampullate gland spigot emerging from cylindrical

tubercle and three piriform gland spigots emerging from shallow tubercle; PMS with minor ampullate

gland spigot and three aciniform gland spigots; PLS with six aciniform gland spigots.

Genitalia (Figs 5A-D, 8A-G). Shape: anterior epigastric margin strongly sclerotized, preceded by

T-shaped structure visible in transparency (AS), with larger, square part (ARe) in between, the latter

provided at base with fine transverse sclerite (T) ending in two faint procurved lobes (FA) extending as

far as lateral protrusions (Pr) of frontal T-shape (AS). SEM view, details: anterior sclerite (AS) short,

T-shaped, with lateral protrusions (Pr) well developed and provided with ramified ridges on anterior side

and dorsally near extremities; AS connected with anterior receptaculum (ARe) antero-ventrally; ARe

about half as wide as AS including Pr, square, provided dorsally and apically with rows of gland ducts

(GD) confined in frontal groove; ARe adjacent to transverse sclerite (T) ending laterally in well developed.

Fig. 4. Tapinesthis inermis. - A-C. $ (PBI 33564), palp. A. Retrolateral view. B. Same, frontal view.

C. Same, prolateral view. - D. $ (PBI 09021), palp, prolateral view, transmitted light. - E. $ palp,

lateral view after Saaristo & Marusik, 2009 (modified). -D: duct; Te: tendon; Tw: thin-walled duct; V:

vesicle.

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European Journal of Taxonomy 82 : 1-20 ( 2014 )

bilobed, flattened apodeme (FA), its dorsal extremity anchoring into the posterior receptaculum (PRe);

PRe well developed, egg-shaped, emerging from base of ARe. Uterus externus not visible.

Tapinesthis inermis (Simon, 1882)

Oonops inermis Simon, 1882: 240.

Tapinesthis inermis. - Simon 1914: 88, 90. —Kaston 1948: 61, figs 4-6. —Kraus 1967: 382, figs 1-10.

— Heimer & Nentwig 1991: 52, fig. 110. — Lazarov, Deltshev & Blagoev 2001: 8, figs 5-6. — Trotta

2005: 170, figs 369-370. —Van Keer 2006: 36, fig. 3. —Korenko, Rezac & Pekar 2007: 6, figs 1-3. —

Saaristo & Marusik 2009: 64, figs 11-14, 23, 27. —Le Peru 2011: 307, fig. 537.

Fig. 5. Tapinesthis inermis , genitalia of $ (PBI 33267) A. Genital area, ventral view. B. Same, after

digestion, ventral view. C. Same, dorsal view. D. Same, anterior view. - ARe: Anterior Receptaculum;

AS: Anterior Sclerite; FA: Flattened Apodeme; Pr: lateral Protrusion; PRe: Posterior Receptaculum.

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HENRARD A., JOCQUE R. & BAEHR B., Redescription of Tapinesthis inermis

Material examined

Lectotype (here designated)

5, Oonops inermis E.S. [= E. Simon], Gall. m. [= Gallia mediterranea?], MNHN AR1468 pars (PBI

OON 6262 pars).

Paralectotypes

2 SS, Oonops inermis E.S. [= E. Simon], Gall. m. [= Gallia mediterranea?], MNHN AR1468 pars (PBI

OON 6262 pars , PBI OON 6263).

Fig. 6. Tapinesthis inermis , SEM views. A. $ (PBI 08123), carapace, antero-lateral view. B. Same,

close up on the distribution of the smooth platelets. C. $ (PBI 33564), sternum, ventral view. D. Same,

mouthparts. E. Same, close up of endite setae. F. Same, cheliceral fangs. G. $ (PBI 08123), endites,

anterior view with detail of serrala. H. $ (PBI 33564), genital region, ventral view.

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European Journal of Taxonomy 82 : 1-20 ( 2014 )

Other material

BELGIUM: Antwerpen, Tumhout, forest remnant, under liana, 27 m, 51.31666° N, 4.93333° E, 19 Oct.

2006, H. De Coninck, 5 $$, RBINS I.G.32708 pars (PBI OON 8123 pars), 3 $$, RBINS I.G.32708

pars (PBI OON 8123 pars). Namur, Marche-les-Dames, 50.48250° N, 4.94917° E, 25 May 2010, A.

Henrard, 1 $, RBINS I.G.32708 pars (PBI OON 33267); Marche-les-Dames, understory, litter among

ivy, 50.48250° N, 4.94917° E, 7 Aug. 2011, A. Henrard, 1 $, RBINS I.G.32708 pars (PBI OON 33564).

CZECH REPUBLIC: Zlinsky Kraj, Lhota u Vsetina, Lhotske Paseky Sklep, building, 545 m,

49.324528° N, 17.936269° E, 8 Aug. 2010, coll. Andrej Machac, det. Jorgen Lissner, 1 $ (leg. J. Lissner,

collection (V), 6821) (PBI OON 32902).

Description

Male (PBI OON 33564)

As for the genus except as noted. Total length 2.02.

Female (PBI OON 6262)

As male except as noted. Total length 2.21.

Distribution

Austria, Belgium, Bulgaria, Czech Republic, France, Germany, Italy, Netherlands, Spain, Switzerland

(after Le Peru 2011, extended).

Remarks

It was the count de Dalmas (1916), and then Kaston (1948), who pointed out that the specimen

mentioned in Emerton’s note on the New England spiders (Emerton 1909) was not Orchestina saltitans

but Tapinesthis inermis. This single specimen is the only one ever recorded from the United States and

was found in the cellar of the Boston Society of Natural History building, probably hidden in a package

shipped from Europe to the Society (Kaston 1948). The mention ‘introduced in USA’ is therefore not

valid as no further specimens were found, which indicates that no viable population of this species

persists in the USA.

Fig. 7. Tapinesthis inermis, SEM views of $ (PBI 33564). A. Palp, frontal view. B. Same, prolateral

view. C. Tip of embolus, ventral view. D. Same, retrolateral view.

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HENRARD A., JOCQUE R. & BAEHR B., Redescription of Tapinesthis inermis

Fig. 8. Tapinesthis inermis , SEM views of $ genitalia (PBI 33267). A. Overview of genital area, dorsal

view. B. Same, close up. C. Same, view slightly anterior. D. Anterior receptaculum (ARe), close up,

arrow indicates gland ducts (GD). E. Same, anterior view. F. Same, detail of gland ducts (arrows).

G. Lateral protrusion (Pr), detail, dorsal view. - ARe: Anterior Receptaculum; AS: Anterior Sclerite; FA:

Flattened Apodeme; GD: Gland Duct; Pr: lateral Protrusion; PRe: Posterior Receptaculum; T: transverse

sclerite.

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European Journal of Taxonomy 82 : 1-20 ( 2014 )

Discussion

Tapinesthis inermis is perhaps one of the easiest oonopid species to identify. This spineless “molles”

oonopid, without abdominal scuta, can easily be recognized by the absence of ventral spines on the first

tibiae, the net-shaped pattern on the carapace, the clumped eyes with slightly recurved PER and by the

plesiomorphic aspect of the male palp and the female genital organs. Although this species is not easily

seen, it cannot be considered as “rare”. However, in the Czech Republic for instance, it is rarefy found

and only females were caught before this study (J. Krai, pers. comm.; see also Korenko et al. 2007).

Dalmas (1916) mentions the occurrence of two adult seasonal forms, one in spring and one in summer,

differing by the colouration and the hair cover. This observation is of interest since it is the only spider

Fig. 9. Tapinesthis inermis , SEM views of spinneret area of <3 (PBI 33564) and $ (PBI 33267).

A. (J, colulus, ventral view. B. Same, anterior lateral spinnerets, ventral view. C. Same, posterior lateral

spinnerets. D. Same, focus on posterior median spinnerets. E. §, posterior median spinnerets, ventral

view. F. Same, anterior lateral spinnerets. G. Same, posterior lateral spinnerets. H. Same, lateral view.

- ALS: anterior lateral spinnerets; PLS: posterior lateral spinnerets; PMS: posterior median spinnerets.

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HENRARD A., JOCQUE R. & BAEHR B., Redescription of Tapinesthis inermis

species for which seasonal forms have been recognized. However, our observations do not point in the

same direction. The illustrations of Fig. 1 clearly show the colour differences between a specimen caught

in su mm er (Fig. 1A-B) and another one photographed in spring (Fig. 1C-E). The spring specimen

(female) is pale, whereas the one from August (male) is orange. The hair cover is similar in both cases.

Since we know that the differences do not depend on the sex, we assume that the two forms are simply

specimens of different age: they become darker, possibly by incorporating pigment from their prey, and

might in some cases loose part of the hair cover. Dalmas (1916) possibly observed dark specimens that

were born in autumn and had passed the winter, which, in the Mediterranean area where he made his

observations, might have been perfectly possible.

Genitalia, male palp

The male palp of T. inermis is an excellent example of the basic plesiomorphic structure of the haplogyne

palp: a piriform bulbus with a simple tubular embolus, without appendage as found in some segestriids.

The latter character is the main difference with Oonopinus angustatus Simon, 1882, another spineless

species (Simon 1882; Saaristo & Marusik 2009; Fe Peru 2011).

A number of features of the male palp can only be seen when it is cleared. The seminal duct has apparently

two parts: a short, well sclerotized part leading into the embolus and a thin-walled part connecting to the

faintly delimited, vesicle-like reservoir. Next to the latter, a thin linear structure runs from the base of

the bulb to the distal end of the sperm reservoir. This structure is also visible in Spinestis nikita Saaristo

& Marusik, 2009 and Megaoonops avrona Saaristo, 2007 (Saaristo & Marusik 2009) and some other

oonopids. Saaristo and Marusik (2009) regard the linear structure adjacent to the sperm reservoir as a

sclerite, but Grismado & Ramirez (2013) and Grismado et al. (in press) clearly showed that this is a

tendon, which is in accordance with the findings of Huber (2004).

Figs. 10. Tapinesthis inermis , SEM views of tarsal claws of $ (PBI 33564). A. Feg I, anterior view.

B. Same, lateral view. C. Feg II, lateral view. D. Same, anterior view. E. Feg III, lateral view. F. Feg

IV, lateral view.

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European Journal of Taxonomy 82: 1-20 (2014)

Female genitalia

It is now well established that the internal genital structures of some haplogynes are highly diverse

and can be remarkably complex (Uhl 2000; Huber 2002; Burger et al. 2003, 2006; Burger 2007).

According to some authors, the complexity of the female genital organ may be correlated to its function

for postcopulatory female choice (Burger et al. 2003; Burger 2011a, 2011b, 2011c).

The female genitalia of Tapinesthis inermis are characterized by the presence of a large T-shaped

anterior sclerite (AS) based antero-ventrally to the anterior receptaculum (ARe), associated with gland

ducts dorsally and apically to the ARe, strong, flattened apodemes and a large, egg-shaped membranous

posterior receptaculum.

Fig. 11. Tapinesthis inermis , SEM, A-H dorsal views, I lateral view. - A-D. S (PBI 33882). A. Leg I.

B. Leg II. C. Leg III. D. Leg IV. - E-G. $ (PBI 33883). E. Leg I. F. Leg II. G. Leg III. - H-I $ (PBI

33564). H. Leg IV. I. Tarsal organ, leg IV.

14

HENRARD A., JOCQUE R. & BAEHR B., Redescription of Tapinesthis inermis

The combination of these structures is very similar to the one encountered in many dysderoid spiders

such as Dysderidae (Forster & Platnick 1985; Ulh 2000; Burger & Kropf 2007), Segestriidae (Brignoli

1976), Orsolobidae (Forster & Platnick 1985; Izquierdo & Labarque 2010) and some basal Oonopidae

such as Pnan chechehet Izquierdo, 2012 (Izquierdo et al. 2012) and Unicorn catleyi Platnick & Brescovit,

1995 (Izquierdo & Rubio 2011). It thus supports the basal placement of Tapinesthis among Oonopinae.

Burger’s terminology for receptacula is based on sections. If a sclerite anterior of the uterus didn’t contain

sperm in one of the different females analyzed, the use of the term “receptaculum” was considered

ambiguous and the term “anterior sclerite” was preferred over “anterior receptaculum”. Burger (2010,

2011a, 2013) assumes that in “higher” oonopids, the receptacula are reduced to sclerites and that sperm

is stored only in the uterus in order to improve the control over fertilization processes through cryptic

female choice (Burger et al. 2003; Burger 2011a, 2011b, 2011c).

In the present work, no sections were made to verily the presence of sperm in those sclerites. However,

the use of “ARe” for this sclerite was preferred to differentiate the upper part, i.e., the T-shaped structure

Fig. 12. Tapinesthis inermis S (PBI 33564), SEM views. A. Cymbium, frontal view. B. Same.

C. Metatarsus II, dorsal view, black arrow showing pore and white arrow indicating eye-shaped structure.

D. Metatarsus IV, black arrow as in C. E. Trichobothrium, dorsal view of bothrium on tibia IV. F. Dorsal

view of bothrium on metatarsus.

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European Journal of Taxonomy 82: 1-20 (2014)

called here anterior sclerite (As), which has a different function (muscle attachment, see Izquierdo &

Rubio 2011).

It was not possible to observe the uterus externus (UE). The tissue of that organ typically appears as

a very flat, membranous structure on SEMs and the opening for the UE is supposed to be somewhere

between the ARe and the PRe (M. Burger, pers. comm.). Again, cross sections are needed to certify

the position of the UE. We therefore avoid the term ‘Anterior Uterine Sclerite (AUS)’ used by Fannes

(2010) and Henrard & Jocque (2012) for what is called here the AS (e.g.). The term ‘uterine’ can indeed

be confusing, especially since we do not know the position of the uterus on SEMs.

In Tapinesthis , gland ducts are confined to a frontal groove on the apical part of the ARe. In other

oonopids, GD are located in the middle of the AS (e.g., Puan chechehet , see Izquierdo et al. 2012) or

at the base of the AS (e.g., Heteroonops spinimanus , see Burger 2011a; Oonopinus kilikus , see Burger

2013). Note that GD are also present on the AS in other dysderoid genera of the families Orsolobidae

(Izquierdo & Labarque 2010), Segestriidae (Brignoli 1976; Forster & Platnick 1985) and Dysderidae

(Uhl 2000). According to Burger (2011b), gland ducts associated with the anterior sclerites may produce

part of the secretion surrounding the spermatozoa in the posterior receptacula. This secretion may either

serve to store and fix the spermatozoa within the receptaculum or provide nutrition for the spermatozoa

(e.g., Uhl 1994, 1996).

Relationships

Simon (1893) recognized two informal groups, ‘molles’ and ‘loricatae’, based on the presence of

abdominal scuta and sclerotization of the body. Petrunkevitch (1923) and subsequent authors recognized

these groups as subfamilies, Oonopinae and Gamasomorphinae, respectively. Later, a third subfamily,

the monotypic Orchestininae, was added for the genus Orchestina (Chamberlin & Ivie 1942). In a

recent paper, Platnick et al. (2012) profoundly changed the subfamilial structure of the Oonopidae. The

main characters they used were the structure of the tarsal organs and more precisely their number of

receptors, the sclerotization of the spermduct in the male palp and the position of the eyes. On the basis

of seven characters, the family was subdivided into three subfamilies: the Sulsulinae, Oonopinae and

Orchestininae, which remained monotypic.

The three main characters on which the recent subfamilial structure is based are not straightforward in

all the oonopids and may need an additional state. In Tapinesthis , and a few other genera, it might indeed

be questioned whether the eyes are clumped and whether the male palpal bulbus lacks a sclerotized

sperm duct. The situation in Tapinesthis can be considered as intermediate between Orchestininae and

Oonopinae: the eyes are not really clumped but also not as far apart as in Orchestina and they nearly

have an H-shaped disposition with a slightly recurved PER. The absence of a clearly thick-walled,

heavily sclerotized sperm duct within the male palp leaves no doubt about the placement of Tapinesthis

among Oonopinae. The sperm duct in the bulbus of Tapinesthis has a thin-walled part, but the distal part

is still clearly sclerotized. However, this structure cannot be considered identical to the sclerotized duct

as found in Orchestina and the outgroups used in Platnick et al. (2012); hence an intermediate state for

this set-up would be necessary. The tarsal organ receptor pattern that gives the impression to be 4433

at a first glance turns out to be 3322, since the distalmost receptor appears to be bifid. Here again, this

configuration would best be considered as a separate state different from both the “simple” 3322 without

a bifid receptor and the plesiomorphic 4433 receptor formulas.

In view of these modifications, a more extensive set of characters wifi be needed to clarify the position of

Tapinesthis within the Oonopinae. However, the combination of the characters discussed above supports

a relatively basal placement of Tapinesthis in the Oonopinae. In addition, recent molecular results of De

16

HENRARD A., JOCQUE R. & BAEHR B., Redescription of Tapinesthis inermis

Busschere et al. (submitted), which place Tapinesthis relatively basally among the Oonopinae, appear to

corroborate our findings on the basis of morphology.

Acknowledgements

The present study is part of a Planetary Biodiversity Inventory (PBI) project of the family Oonopidae,

coordinated by the American Museum of Natural History (AMNH), New York. We are indebted to

Jorgen Lissner for the loan of material ( Tapinesthis inermis S from the Czech Republic). We thank Y.

Marusik for sharing the nice picture of the $ palp (Fig. 4E). M. Burger and M. Izquierdo shared their

knowledge of the Oonopidae in order to solve difficulties with the interpretation of the structure of the

female genitalia. B. Huber, P. Michalik and M. Ramirez co mm ented on the structure in the cleared male

palp. This paper is publication BRC 307 of the Biodiversity Research Center (Universite Catholique de

Louvain).

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Manuscript received: 19 December 2013

Manuscript accepted: 18 March 2014

Published on: 5 May 2014

Topic editor: Koen Martens

Desk editor: Danny Eibye-Jacobsen

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; National History Museum, London, United

Kingdom; Royal Belgian Institute of Natural Sciences, Brussels, Belgium; Natural History Museum of

Denmark, Copenhagen, Denmark.

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