http://dx.doi.org/10.5852/ejt.2013.47

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2013 • Jocque R.C.A.M.

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

Research article

um:lsid:zoobank.org:pub:BABA7D0E-DC98-402E-BD45-A7D9478E376A

Cyrioctea (Araneae, Zodariidae) in Africa:

temperate Gondwanaland relict, recent radiation, or both?

Rudy C.A.M. JOCQUE

Royal Museum for Central Afriea, Eeuvensesteenweg 13, 3080 Tervuren, Belgium.

E-mail: [email protected]

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

Abstract. Two new speeies of the zodariid genus Cyrioctea Simon, 1889 are deseribed: C. sawadee

sp. nov. and C. lotzi sp. nov., both only known from males. The genus now eontains seven Afrotropieal

speeies and this abundanee is diseussed in the eontext of its basal situation in the family and its apparent

temperate Gondwanaland distribution, whieh implies a mueh greater age of the Zodariidae than presently

aeeepted. Unlike most taxa with a temperate Gondwana distribution, Cyrioctea boasts a high number

of speeies with small distribution areas. This points in the direetion of a reeent radiation initiated after

a long period of stasis.

Keywords. Cyrioeteinae, distribution area, eomplexity, identifieation key.

Jocque R.C.A.M. 2013. Cyrioctea (Araneae, Zodariidae) in Africa: temperate Gondwanaland relict, recent

radiation, or both? European Journal of Taxonomy 47: 1-12. http://dx.doi.org/10.5852/eit.2013.47

Introduction

The spider genus Cyrioctea is remarkable in many ways. The spiders are eharaeterized by a transverse

row of, most often six, short but sturdy spines between the eye rows. These are assumed to eonstitute

a burrowing deviee. Cyrioctea do indeed live under ground and females, that hardly ever emerge from

their burrows, are very rarely eolleeted. Although the position of the genus has been the subjeet of some

debate, these spiders elearly belong to the Zodariidae (Joeque 1991): they laek a serrula and are provided

with long anterior lateral spinnerets. Sinee the elaw teeth are positioned in the axis of the elaw, they

were eonsidered as the sister group of all other Zodariidae, in whieh the elaws are positioned on the

side of the elaw faeing the other one. The first eladistie analysis of the family on genus level therefore

used Cyrioctea as the outgroup. A reeent unpublished eladistie analysis (Joeque 2006) on morphologieal

grounds with Amaurobius as outgroup (see Joeque & Dippenaar 2007), finds Cyrioctea in the same

position at the root of the Zodariidae, thus sister to all other genera in the family.

Beeause of its aneestral eharaeter it is not surprising to find the genus on three eontinents: Afriea, South

Ameriea and Australia (Platniek 1986; Platniek & Griffin 1988; Platniek & Joeque 1992). This is a

typieal temperate Gondwanaland distribution with all the loealities on the southern tip of the eontinents.

On the other hand, the eomplexity of the male genitalia is startling. Moreover, the speeies appear to have

European Journal of Taxonomy 47: 1-12 (2013)

very small distribution areas, whieh is also puzzling as the Gondwanaland reliets have been assumed to

be leftovers of old radiations with a large distribution.

The present paper deseribes two new speeies, gives an overview and a distribution map of Afriean

Cyrioctea speeies. A key to the males is provided.

Material and methods

All material is preserved in 70% ethanol. Speeimens were observed and measured with a Leiea MIO

stereo mieroseope. Photographs were taken with a Leiea MZ16 using the LAS automontage software.

Drawings were earried out with a eamera lueida on a WILD M5.

All measurements are in mill im etres.

Abbreviations

ALE

Anterior Eateral Eyes

AME =

Anterior Median Eyes

Central apophysis

dorsal

Distal apophysis

dw =

distal whorl

DTA

Dorsal tibial apophysis

Embolus

Femur

median apophysis

Metatarsus

NCA =

National Colleetion of Araehnids, Pretoria (A. Dippenaar)

NMBA =

National Museum Bloemfontein, Araehnids (E. Eotz)

Patella

PEE

Posterior Eateral Eyes

PME =

Posterior Median Eyes

prolateral

retrolateral

RTA

retrolateral tibial apophysis

Tibia

V =

ventral

Results

Taxonomic descriptions

Class Araehnida Cuvier, 1812

Order Araneae Clerek, 1757

Family Zodariidae Thorell, 1881

Subfamily Cyrioeteinae Joeque, 1991

Genus Cyrioctea Simon, 1889

Remarks

The genus Cyrioctea was deseribed by Simon (1889), on a speeies from South Ameriea and redeseribed

in detail by Platniek (1986) and Joeque (1991). Cyrioctea now eontains 13 speeies of whieh five oeeur in

JOQCUE R.C.A.M., Cyrioctea in Africa: relict, recent radiation or both?

South America, one in Australia (Platnick 2012) and seven in southern Africa. These African species are

C. griswoldorum Platnick & Jocque, 1992 ( 3 ), C. hirsuta Platnick & Griffin, 1988 (?), C. lotzi sp. nov.

(6'), C. marken Platnick & Jocque, 1992 ( 3 ), C. namibensis Platnick & Griffin, 1988 ((??), C. sawadee

sp. nov. (6') and C. whartoni Platnick & Griffin, 1988 ((??)•

Key to the males of African Cyrioctea (C. hirsuta only known from females)

1 Male palpal tibia with dorsal apophysis clearly separated from broad RTA which is provided with

tooth on inferior margin.2

- Male palpal tibia without dorsal apophysis; RTA without tooth on inferior margin.3

2 Dorsal apophysis much shorter than RTA, which is roughly triangular and pointed; abdomen dark

grey except for white spot above spinnerets. C. griswoldorum Platnick & Jocque, 1992

- Dorsal apophysis almost as long as RTA, which is roughly rectangular and broadly truncated; abdomen

pale with dark pattern of central spot followed by transverse bands (Fig lA). C. sawadee sp. nov.

3 RTA a single sharp, down-curved prong. C. namibensis Platnick & Griffin, 1988

- RTA bifid .4

4 Ventral tip of RTA longer than dorsal tip; distal tegular apophysis with narrow tip.

. C. whartoni Platnick & Griffin, 1988

- Ventral tip of RTA shorter than dorsal tip; distal tegular apophysis broadly truncated.5

5 Ventral tip of RTA a tiny prong, much smaller than dorsal tip; median apophysis without median

spur at base (Fig. 3E). C. lotzi sp. nov.

- Ventral tip of RTA not so small; median apophysis with median spur at base.

. C. marken Platnick & Jocque, 1992

Cyrioctea sawadee sp. nov.

um:lsid:zoobank.org:act:8C7A2AA2-79C5-4EDA-BC0F-29FF40058A66

Figs lA-C, 2A-B, 3A-C, 4

Diagnosis

The male of C. sawadee sp. nov. can be recognized by the modified third leg with dense spination and

very short tibia, and by the characters of the palp, of which the tooth on the ventral margin of the RTA is

the most conspicuous. That character is shared with C. griswoldorum, which has a shorter dorsal tibial

apophysis, the shape of the RTA is triangular rather than rectangular and the abdominal pattern consists

of one pale spot on a dark background.

Etymology

The species name is a noun in apposition taken from the type locality.

Type material

Holotype

SOUTH AFRICA: (J, Western Cape Province, Sawadee, 32°20.316’ S - 18° 49.405’ E, Oct. 2007,

pittrap, 359 m alt., Nortje E. & Kritzinger-Klopper (NCA).

Paratypes

2 SS, same data as holotype.

European Journal of Taxonomy 47: 1-12 (2013)

Other material examined

None.

Description

Male (holotype, Fig. lA-C)

Measurements. Total length 4.32; earapaee 2.14 long, 1.22 wide; Tl+Pl: 1.95.

CoEOUR. Carapaee brownish yellow with blaek fovea and dark margin; palp, ehelieerae, mouthparts

and sternum pale brown; legs yellow, femora suffused with dark grey; abdomen pale grey with dark

pattern of eentral spot followed by four transverse bands; venter medium grey, darkened towards yellow

spinnerets.

Cepelaeothorax. Clypeus eentre devoid of setae but with dense eluster of inward eurvedthiek setae on either

side. Eye region with a row of six slightly eurved spines (Figs IB, 3A), on right side with one short thiek

extra seta.Chilum poorly developed, ineonspieuous. Sternum subeireular, with rather long, thin, posterior

extension.

Eyes. AME: 0.04; AEE: 006; AME-AME: 0.08; AME-AEE: 0.01; PME: 0.06: PEE: 0.06; PME-PME:

0.05; PME-PEE: 0.07. Clypeus 0.17 or 2.9 times width of AEE.

Fig. 1. - A-C. Cyrioctea sawadee sp. nov., 3 - A. Habitus , dorsal view. B. Carapaee, dorsal view. C.

Idem, ventral view. - D-F. Cyrioctea lotzi sp. nov., S- D. Habitus, dorsal view. E. Carapaee, dorsal view.

F. Idem, ventral view. (Seale bars: 2 mm).

JOQCUE R.C.A.M., Cyrioctea in Africa: relict, recent radiation or both?

Legs. Legs III modified, provided with numerous spines; femora slightly swollen, spineless, tibiae short,

as long as patella.

Spination.

dl-lv2

rllv2

III

pl3rll-2

pll-l-ld2-15rll-lv2

7dispdw6

pl2rll

pldl-l-lrll-lv2-2-2

8dispdw6

Fig. 2. - A-B. Cyrioctea sawadee sp. nov., S. A. Palp, ventral view. B. Idem, retrolateral view. - C-D.

Cyrioctea lotzi sp. nov., S- C. Palp, ventral view. D. Idem, retrolateral view. (Scale bars: 200 pm).

European Journal of Taxonomy 47: 1-12 (2013)

100 ijm

Fig. 3. - A-C. Cyrioctea sawadee sp. nov., 3. A. Carapace, dorsal view. B. Palp, retrolateral view. C.

Idem, ventral view. - D-G. Cyrioctea lotzi sp. nov., c5'. D. Carapaee, dorsal view. E. Palp, retrolateral

view. F. Idem, ventral view. G. Idem, prolateral view. CA: eentral tegular apophysis; DA: distal tegular

apophysis; DTA: dorsal tibial apophysis; E: embolus; MA: median apophysis; RTA: retrolateral tibial

apophysis.

JOQCUE R.C.A.M., Cyrioctea in Africa: relict, recent radiation or both?

Male palp. (Figs 2A-B, 3 B-C) Tibia with short, triangular dorsal apophysis; RTA broad with dorsal

margin smoothly curved down, ventral margin straight, with small tooth at base; cymbium oval, with

sclerotized rim, slightly bulging near RTA. Tegulum strongly bulging and complex; embolus prolateral,

broad and with semitransparent proximal flange, smoothly curved outwards; median apophysis near

retrolateral margin, curved downward, ventrally concave; central apophysis (CA) with digitiform

sclerotized tip; distal apophysis (DA) with three short prongs.

Female

Unknown

Distribution

Only known from type locality in the Western Cape Province (Fig. 4).

Cyrioctea lotzi sp. nov.

um:lsid:zoobank.org:act:57043731-C0QD-47Q5-913A-5DDFE5681F97

Figs ID-F, 2C-D, 3D-G, 4

Diagnosis

The male of C. lotzi sp. nov. can be recognized by the pale abdomen with faint dorsal stripe, the strong

spines on the first leg and the details of the male palp in which the RTA is adjacent to the dorsal apophysis.

Etymology

The species name is a patronym in honour of one of the collectors.

Type material

Holotype

SOUTH AFRICA: S, Free State, Deelfontein, 27°07’ S -26° 35’ E, Aug.-Oct. 1986, preservation trap.

Museum Staff (NMBA).

Paratypes

4 cJcJ, same data as holotype (1 (? in MRAC).

Other material examined

None.

Description

Male (holotype. Fig. ID-F)

MEASUREMENTS. Total length 4.54; carapace 2.30 long, 1.31 wide; TI+PI: 2.40.

Colour. Carapace brownish yellow with black fovea and dark margin; palp, chelicerae, mouthparts,

sternum and legs yellow; femora suffused with dark grey; abdomen pale grey with faint darker dorsal

median stripe; venter and spinnerets pale.

Cepeialothorax. Clypeus provided with dense cluster of short upturned thick setae. Eye region with a

row of six slightly curved spines (Figs IE, 3D). Chilum poorly developed, inconspicuous. Sternum oval;

slightly wider than long with rather long, thin, sharp posterior extension.

Eyes. AME: 0.06; AEE: 007; AME-AME: 0.13; AME-AEE: 0.0; PME: 0.09; PEE: 0.08; PME-PME:

0.08; PME-PEE: 0.06. Clypeus 0.16 or 2.2 t im es width of AEE.

European Journal of Taxonomy 47: 1-12 (2013)

Legs. Legs 1 with numerous and strong but short spines; legs 111 with hair eover slightly denser than

other legs.

Spination.

pll-l-2rll-lvl-l-l-l-l

v2-2

pll-lrll-l-lv3

v2dw3

111

pBdlrll

6dispv2

8dispdw6

dlvl

rll

pldl-ldl-lrll-l-lv2-2-2

8dispdw6

Male palp. (Figs 2C-D, 3E-G) Tibia with short, triangular dorsolateral apophysis, adjaeent to slightly

shorter RTA; eymbium almost rounded, with selerotized rim, thiekened near RTA. Tegulum strongly

bulging and eomplex: embolus distal, provided with transparent flange; median apophysis near posterior

retrolateral margin; eentral apophysis triangular; distal part of tegulum a transverse ridge with selerotized

rim.

Female

Unknown.

Distribution

Only known from type loeality in the Free State, South Afriea (Fig. 4).

Discussion

Taxa with a temperate Gondwana distribution, also ealled an Austral distribution (Humphreys & Parenti

1999), found on the southern tip of the eontinents Australia, Afriea and South Ameriea, sometimes

ineluding New Zealand and New Caledonia, have been eonsidered to be reliets (e.g. Crisei et al. 1991;

Robertson & Holzenthal 2005). This type of distribution is often synonymous with the presenee of

aneestral eharaeters, whieh is also the ease for Cyrioctea. Other examples are the aneestral Ratites,

flightless birds (Bourdon et al. 2009), Pleurodira, primitive turtles (Sterli 2010) and beetles that are

herbivorous on Araucaria (Sequeira & Farrel 2001). Some of these have been qualified as living fossils

as they are at the base of old, large elades. Sueh groups, albeit rarely exhibiting a Gondwana-distribution,

are most often very poor in speeies, e.g., Tuatara (Hay et al. 2010), Platypus (Groves 2005), Coelaeanths

(Forey 1998) ete. However, this phenomenon eannot be qualified as a rule sinee there are remarkable

exeeptions. The “primitive” spiders with segmented abdomen belonging to the family Liphistiidae are

an exeellent example of sueh an exeeption: the family is eomposed of 5 genera and eontains not less

than 85 speeies (Platniek 2012). And in spider families for whieh a detailed phylogenetie analysis is

available, it is not the rule that basal genera are speeies poor. It might be so for Lordhowea Griswold,

2001 in Cyatholipidae (Griswold 2001) or PararchaeaV orsior, 1955 in Pararehaeidae (Rix 2006), but in

Theridiidae, for instanee, the most basal genus is the speeiose Dipoena Thorell, 1869 (Agnarsson 2004).

Also in families like Linyphiidae (Miller & Hormiga 2004), Ctenidae (Silva 2003) and Thomisidae

(Benjamin et al. 2008), the most basal genera are rieh in speeies. Thus it is eertainly not a general

phenomenon that basal taxa of a large evolutionary line tend to be reliets in the sense that they are poor

in speeies.

But also from the morphologieal point of view, eertain eharaeters and more preeisely the genitalia

of Cyrioctea, are puzzling. In the Zodariidae, the root of the large elades is very often oeeupied by

speeies with very simple genitalia. A reeurring pattern is the presenee of a short, straight, spine-like

embolus aeeompanied by a short and simple median apophysis and a simple RTA in the speeies near

JOQCUE R.C.A.M., Cyrioctea in Africa: relict, recent radiation or both?

the root, evolving to a long embolus accompanied by intricate supporting structures and the addition

of apophyses on the palpal patella and even the femur. Examples are Diores namibia Jocque, 1991 for

Diores with more than 50 species (Jocque 1991), Tenedos infrarmatus Jocque & Baert, 2002 at the

base of a large clade containing the genera Tenedos, Ishania, Epicratinus and Colima (Jocque & Baert

2005), Procydrela procursor Jocque, 1999 at the base of the Cydrelinae (Jocque 1999), and Pentasteron

simplex Baehr & Jocque, 2001 at the base of the very large Asteron complex in which more than 100

species have already been described (Baehr & Jocque 2001).

In other families where a similar phenomenon is assumed to occur, the trend is often obscured by the

creation of separate genera exactly on the basis of these additional structures on palpal patella and femur

(e.g. Wang 2002). Data on the Zodariidae, one of the few families for which there is a combination of

generic and species phytogenies, show that a somatic radiation precedes the genital one. But Cyrioctea,

does not qualify. All the species have a complex bulbus with several tegular appendages. In several

species, however, the palpal tibia shows a typical ancestral character with the presence of a dorsal

apophysis, which may be the precursor of the retrolateral tibial apophysis (RTA). But the complexity

of the bulbus is incoherent with the idea that Cyrioctea might be considered a living fossil dating back

from the split-off of Gondwanaland. It might therefore be argued that a recent radiation has occurred,

explaining the existence of closely related species with small distribution areas. Such a recent radiation

Fig. 4. Distribution of Cyrioctea species in Africa: (■) = C. griswoldorum Platnick & Jocque, 1992; (□)

= C. hirsuta Platnick & Griffin, 1988; (A) = C. lotzi sp. nov.; (A) = C. marken Platnick & Jocque, 1992;

(•) = C. namibensis Platnick & Griffin, 1988; (+) = C. sawadee sp. nov.; (o) = C. whartoni Platnick &

Griffin, 1988.

European Journal of Taxonomy 47: 1-12 (2013)

is known for the plants belonging to the Cyeadales, eonsidered a group of “living fossils” until reeently

(Nagalingum et al. 2011). Neither the Liphistiidae nor Cyrioctea have been subjeet to sueh a detailed

analysis, but may well represent other examples of old taxa with a reeent radiation. However, the most

important eonsequenee of Cyrioctea's distribution is that the family is mueh older than what is proposed

in Penney & Selden (2011). Following Miller et al. (2010a, 2010b), they eonsider Zodariidae the sister

family of Penestomidae and eonelude that the elade is not older than 30 ma. Sinee it must date baek from

before the split up of western Gondwanaland, its age must be pushed baek to at least 100 ma (Pitman et

al. 1993).

It is evident that only a detailed eladistie analysis ineluding moleeular data ean eorroborate the hypothesis

that has been formulated here.

Acknowledgements

1 am indebted to A. Dippenaar-Sehoeman (NCA, Pretoria) and L. Lotz (NMBA, Bloemfontein) for the

loan of material. 1 have appreeiated the diseussions with Mare De Meyer and Jos Snoeks eoneeming

temperate Gondwana distributions. 1 thank Alain Reygel for the drawings.

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Manuscript received: 30 January 2013

Manuscript accepted: 16 April 2013

Published on: 13 June 2013

Topic editor: Koen Martens

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; National Botanic Garden

of Belgium, Meise, Belgium; Royal Museum for Central Africa, Tervuren, Belgium; Natural History

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

Natural History Museum of Denmark, Copenhagen, Denmark.