European Journal of Taxonomy 40: 1-49

http://dx.doi.org/10.5852/ejt.2013.4Q

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2013 • Jan Bosselaers & Rudy Joeque

Research article

um:lsid:zoobank.org:pub:99B180D2-CCD2-4171-B64Q-E3EB68F94E2B

Studies in Liocranidae (Araneae): a new afrotropical genus

featuring a synapomorphy for the Cybaeodinae

Jan BOSSELAERS' & Rudy JOCQUE^

Royal Museum for Central Alfiea, Tervuren, Belgium

2 Email: [email protected] (eorresponding author)

^ um:lsid:zoobank.org:author:D6AD7414-3540-4FQ6-8631-8873450AA9QC

" um:lsid:zoobank.org:author:CF15016C-8CDl-4C9D-9Q21-44CA7DC7A5D5

Abstract. Cteniogaster, a new genus of small ground spiders is deseribed from Kenya and Tanzania. It

eneompasses seven new speeies, three of whieh are known from both sexes: C. toxarchus sp. nov., the

type speeies, C. conviva sp. nov. and C. hexomma sp. nov. Three speeies are known from females only:

C. lampropus sp. nov, C. sangarawe sp. nov. and C. taxorchis sp. nov. and one only from males: C.

nana sp. nov. The new genus ean be reeognised by the presenee of a posterior ventral abdominal field

of strong setae and anterior lateral spinnerets with enlarged piriform gland spigots in males. A eladistie

analysis attributes the genus to Eioeranidae, Cybaeodinae. The results of the analysis performed do not

produee an unequivoeal autapomorphy for Eioeranidae, but provide a eombination of non-homoplasious

eharaeter ehanges that offers signifieant potential for reeognising genera as Eioeranidae. Moreover, robust

apomorphies are determined within Eioeranidae for the subfamilies Eioeraninae and Cybaeodinae. Based

on these findings Toxoniella Wami & Joeque, 2002 is transferred from Gallieniellidae to Eioeranidae,

Cybaeodinae. Jacaena Thorell, 1897, Plynnon Deeleman-Reinhold, 2001 and Teutamus Thorell, 1890

are transferred to Corinnidae, Phrurolithinae and Montebello Hogg, 1914 to Gnaphosidae. Itatsina

Kishida, 1930 is synonymised with Proc/zora Simon, 1886.

Keywords, phylogeny, enlarged piriform gland spigots, ventral abdominal setae, eye reduetion. Eastern

Are Mountains.

Bosselaers J. & Joeque R. 2013. Studies in Liocranidae (Araneae): a new afrotropical genus featuring a synapomorphy

for the Cybaeodinae. European Journal of Taxonomy 40: 1-49. http://dx.doi.org/10.5852/eit.2013.40

Introduction

Eioeranidae Simon, 1897 feature among the poorly defined spider families (Joeque & Dippenaar-

Sehoeman 2006) sinee not a single synapomorphy has been listed so far. Making abstraetion of the

Paratinae (Marusik et al. , 2008) whieh are only tentatively plaeed in Eioeranidae, both subfamilies that

eompose the family, Cybaeodinae Simon, 1893 and Eioeraninae Simon, 1897, are also poorly delimited.

The present paper deseribes a new genus in whieh the males are eharaeterized by a eonspieuous group of

rigid setae on the venter of the adomen. More important is that they have enlarged piriform gland spigots

on the male anterior lateral spinnerets, a eharaeter first deseribed by Platniek (1990) for some Clubionidae

Wagner, 1887, and later also diseovered in Toxoniella Wami & Joeque, 2002, a genus that was attributed

1

European Journal of Taxonomy 1-49 ( 2013 )

to Gallieniellidae Millot, 1947. This peculiar spinneret character appears to be present in most genera

(Agroeca Westring, 1861, Agraecina Simon, 1932, Cteniogaster gen. nov., Cybaeodes Simon, 1878,

Liocranoeca Wunderlich, 1999, Neoanagraphis Gertsch & Mulaik, 1936) that have been attributed to

the Cybaeodinae, but is lost in Apostenus Westring, 1851 and Scotina Menge, 1873. Nevertheless, a

cladistic analysis shows that its presence can be considered a synapomorphy for the subfamily. Apart

from the description of the new genus, the relationships between the genera in Liocranidae and the

distribution of the spinneret character on the cladogram are studied in the paper.

Material and Methods

Specimens were observed and drawn using Euromex M1C465 and Olympus SZX9 binocular

microscopes. Photographs of the habitus were taken with a Leica MZ16 binoclar microscope using the

LAS automontage software. Female epigynes were 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,

specimens or parts were dried in HMDS, gold coated and examined and photographed with a JEOL

6480 LV scanning electron microscope. Tarsal claws and details of the distal end of the male bulbus were

observed and drawn using a Wild M12 compound microscope, except for the tarsal clams of Apostenus

spinimanus (Koch & Berendt, 1854), for which a fossil in Baltic amber (CJB) was observed with an

Olympus SZX9 binocular microscope. In order to precisely position bulbi for observation under the

compound microscope (Fig. 11 A, B), the male palps were immobilized in Schwarzkopf “Freezing Gel

Extreme 5”, a transparent vinylpyrrolidone-vinylacetate copolymer (CAS 25086-89-9) / carbomer gel

(Schroder et al. 2000; Shulze zur Wiesche 2006). Leg spination is illustrated in a schematic representation

(Appendix 1) where pi, do, rl and ve sides of leg articles are flattened as a folding net (Durer 1525). The

format for leg spination in the genus description follows Platnick & Shadab (1975), amended for ventral

spine pairs according to Bosselaers & Jocque (2000). All measurements are in mm.

Abbreviations used

AE

AER

ALE

AES

AME

CO

do

LEGS

fe

ft

HMDS

ICS

EE

MA

ME

MOQ

mt

pa

PCT

PE

PER

pi

anterior eyes

anterior eye row

anterior lateral eyes

anterior lateral spinnerets

anterior median eyes

copulatory openings

dorsal

enlarged piriform gland spigots

femur

frontal

hexamethyldisilazane (CAS 999-97-3)

intercoxal sclerites

lateral eyes

median apophysis

median eyes

median ocular quadrangle

metatarsus

patella

precoxal triangles

posterior eyes

posterior eye row

prolateral

2

r

BOSSELAERS J. & JOCQEIE R., Anew afrotropical genus of Cybaeodinae (Araneae, Eioeranidae)

PEE =

posterior lateral eyes

PES =

posterior lateral spinnerets

plv =

prolateral ventral

PME =

posterior median eyes

PMS =

posterior median spinnerets

rh =

retrocoxal hymen

rl

retrolateral

rlv =

retrolateral ventral

RTA =

retrolateral tibial apophysis

STl =

spermatheca(e) 1

ST2 =

spermatheca(e) 2

ta =

tarsus

ti =

tibia

ve =

ventral

vt

ventral terminal.

Abbreviations of personal and institutional collections (curators in parentheses)

AMNH

CARS

CAS

CHK

CJB

CJK

MCN

MNHN

MRAC

MZHF

RMNH

SMF

UCR

ZMUC

American Museum of Natural History, New York (N. Platniek)

personal eolleetion Anthony Russell-Smith

California Academy of Sciences, San Fransisco (C. Griswold)

personal collection Herman De Koninck

personal collection Jan Bosselaers

personal collection Johan Van Keer

Museu de Ciencias Naturais, Porto Alegre (E. Buckup)

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

Royal Museum for Central Africa, Tervuren (R. Jocque)

Finnish Museum of Natural History, Helsinki (P Cardoso)

National Museum of Natural History, Eeiden, The Netherlands (J. Miller)

Senckenberg Forschungsinstitut und Naturmuseum, Frankfurt (P Jager)

Entomology Research Museum, University of California, Riverside (R. Vetter)

Zoological Museum, University of Copenhagen (N. Scharff)

Phylogenetic analyses were performed using the computer programmes PAUP* 4.0 beta 10 (Swofford

2002), Winclada 1.00.08 (Nixon 2002), and TNT 1.1 (Goloboff et al. 2003, 2008a). Optimisation of

character states and printing of the preferred tree was performed using Winclada. All three programmes

were 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).

Taxon choice

Based on somatic and genitalic characters, Cteniogaster gen. nov. was tentatively attributed to

Eioeranidae. In order to investigate its relationships in a more objective way, a phylogenetic analysis

was performed. The analysis included the three species of Cteniogaster gen. nov. of which males and

females are available, 23 species of Eioeranidae belonging to 11 different genera, and the two known

species of Toxoniella Wami & Jocque, 2002, a genus similar to Cteniogaster gen. nov., whose current

place in Gallieniellidae is perceived as uncertain. Where possible, more than one species was included

for each genus, in order to increase the robustness of the analysis. Two corinnid species, Corinna nitens

(Keyserling, 1891) and Creugas baiulus (Gertsch, 1942), were used as outgroup (Watrous & Wheeler

1981;Maddisone/a/. 1984).

A list of material examined for assessment of morphological characters is provided in Appendix 2.

3

Hesperocranum rothi

European Journal of Taxonomy 1-49 ( 2013 )

<0

O)

3

C

Cn

^ <pf

y m

) ^^^ 1 ^

4

Fig. 1. Strict consensus of three fittest trees obtained under implied weighting for 25 lioeranid speeies and two eorinnid outgroup speeies. State ehanges

are indieated on the tree for 47 out of the 99 eharaeters used, ambiguous eharaeter state ehanges were optimized individually (see text for details).

Non-homoplasious state ehanges are in blaek, homoplasious state ehanges in white. Nodes are numbered on the tree and Goloboff fit Bremer support

values, as reported in TNT, are indieated in italies below branehes.

r

BOSSELAERS J. & JOCQEIE R., Anew afrotropical genus of Cybaeodinae (Araneae, Eioeranidae)

Characters

A series of 99 eharaeters (77 binary and 22 multistate) was eoded for the 27 taxa ehosen. All eharaeters

are phylogenetieally informative. Where possible, eharaeters were seored with eharaeter states

hierarehieally related, as advoeated by Hawkins et al. (1997), even though this neeessitated eoding

missing entries due to eharaeter inapplieability in some instanees (Maddison 1993). Genitalie traits are

dealt with in 26 of the eharaeters, the other 73 eharaeters are somatie, 21 of them referring to leg or

palp spination. Possible problems of spine homology were addressed as in Bosselaers (2002: 143) and

Bosselaers & Joeque (2002: 244). All 99 eharaeters used in the analysis are explained below. On the

preferred tree (Fig. 1), 29 eharaeters ean be optimised ambiguously (indieated by ^ in the list below),

and 26 eharaeters are fully non-homoplastie (indieated by an asterisk). Out of the 99 eharaeters seored,

47 are shown in Fig. 1 (indieated by ^ in the list below).

1. ^Male rh: (0) absent, (1) present. The rh was first deseribed by Raven (1998) as the “retroeoxal

window”. It is a weak spot, in most eases hyaline and lens-shaped, on the rl faee of eoxa I (Bosselaers

& Joeque 2002: fig. lA).

2. ^Female retroeoxal hymen: (0) absent, (1) present.

3. ^Troehanter noteh: (0) absent, (1) present.

4. ^Rows of bristles in ve seopulae of legs I and II: (0) absent, (I) present.

5. *^Rows of bristles in ve seopulae of legs I and II: (0) small, (1) large, ereetile. The bristles mentioned

under eharaeter 4 ean be small, i.e. hardly larger than normal seopulae, or large and ereetile, and

with a peeuliar basal soeket, as deseribed by Ubiek & Platniek (1991: 2).

6. ^Patellar indentation of legs I and II: (0) narrow, (I) wide. The patellar indentation is a slit-like

membranous groove on the rl side of the pa (Simon 1892: 22; Fedoux & Canard 1991: 9, fig. 15A, B)

7. Patellar indentation of leg IV: (0) narrow, (I) wide.

8. *^Bent male tarsi IV: (0) absent, (I) present (Fig. I ID). The tarsi IV of males are bent and

subsegmented in a eharaeteristie way in a number of lioeranid genera (Ubiek & Vetter 2005: 71;

Wunderlieh 1999: 68).

9. *Temur I plv spines: (0) absent, (1) present.

10. Femur I apieal do spine: (0) absent, (1) present.

11. Femur III and IV apieal do spine: (0) absent, (1) present.

12. Patella III and/or IV rl spine: (0) absent, (1) present.

13. ^Male ti I plv spines: (0) absent, (1) one to three, (2) four to six, (3) more than six.

14. ^Female ti I plv and rlv spines: (0) absent, (1) one to three, (2) four to six, (3) more than six.

15. ^Tibia II plv spines: (0) absent, (1) one to three, (2) four to six.

16. ^Tibia II rlv spine number: (0) similar to plv spines, (1) at least two spines more than plv spine

number.

17. *Temale ti III do spines: (0) absent, (I) present.

18. *^Tibia IV do spines: (0) absent, (I) present.

19. ^^Metatarsus I and II ve spine pairs: (0) absent, (I) one, (2) more than one.

20. ^Metatarsus III plv and rlv spines: (0) none or one, (1) two, (2) three.

21. ^Metatarsus IV plv and rlv spines: (0) absent, (1) one, (2) two, (3) three.

22. ^^Male mt IV rl spines: (0) one or two, (1) three to four, (2) five or more.

23. Metatarsus III and IV vt spines: (0) absent, (I) present on both.

24. Male palpal pa pi spines: (0) absent, (1) present.

5

European Journal of Taxonomy 1-49 ( 2013 )

25. ^Male palpal ti pi spines: (0) none or one, (1) two, (2) three, (3) four.

26. Male palpal ti do spines: (0) absent, (1) one, (2) two.

27. ^Male palpal ta distal pi edge spines: (0) absent, (1) present.

28. Female palpal fe pi spines: (0) absent, (1) present.

29. Female palpal ti pi spines: (0) two, (1) three, (2) four.

30. ^Feathery hairs (Griswold 1993: fig. 61; Townsend & Felgenhauer 1998: fig. 8): (0) absent, (1)

present (Fig. 6B).

31. Metatarsal vt preening brush on tarsi 111 and IV: (0) absent, (1) present.

32. ^Metatarsal vt preening brush on tarsi 111 and IV: (0) sparse, (1) dense.

33. *^Tibia 1 and 11 ve seopulae: (0) absent, (1) present.

34. Metatarsal ve seopulae: (0) absent, (1) present.

35. Tarsal ve seopulae: (0) absent, (1) present.

36. *^Claw tufts: (0) absent, (1) present.

37. ^^Tenent hairs (Forster 1970: 18; Ubiek & Vetter 2005: 69, 71): (0) absent, (1) present (Figs 6H,

IIC-E).

38. *^^Tenent hair pairs: (0) one, (1) two, (2) four or more. The number of tenent hair pairs present varies

eonsiderably between lioeranid genera (Bosselaers 2009: 39, 49, 51), as will be diseussed below.

39. PCT: (0) absent, (1) present. PCT are small triangular selerites surrounding the sternum, their tips

faeing the bases of the eoxae (Penniman 1985: 16).

40. ICS in male: (0) absent, (1) two pairs, between eoxae 1 and 11 and between eoxae 11 and 111, (2) three

pairs. ICS are small triangular or elongated selerites surrounding the sternum, their tips penetrating

between the eoxae of the legs.

41. Gntereoxal selerites in female: (0) absent, (1) two pairs, between eoxae 1 and 11 and between eoxae

11 and 111, (2) three pairs.

42. ^Sternal border: (1) simple or with indistinet, thin border, (2) strongly rebordered.

43. ^Retromarginal ehelieeral teeth number: (0) two, (1) more than two.

44. ^Long shaggy hair in front of fang base: (0) absent, (1) present. A eonspieuous hair at the ehelieeral

promargin, as long as the fang and bent at a right angle just beyond its base (Bosselaers & Joeque

2002: fig. lM,Q;Platniek2000: 10).

45. ^Chilum: (0) absent, (1) present. The ehilum is a small subtriangular selerite at the base of the

ehelieerae, below the elypeus (Joeque 1991: 11).

46. Chilum: (0) bilateral and bipartite, (1) median and entire.

47. ^General shape of endites: (0) with external lateral noteh, (1) parallel-sided.

48. *^Serrula of endites: (0) eonspieuous, (1) redueed.

49. ^Apieal maxillar hair tuft of endites: (0) absent, (1) present.

50. *^Carapaee shape in lateral view: (0) earapaee slanting, highest at fovea, (1) earapaee fiat, (2)

earapaee bulging, highest in eephalie region.

51. Clypeus height: (0) smaller than diameter of AME, (1) equal to diameter of AME, (2) larger than

diameter of AME.

52. Male AER eurvature from front: (0) proeurved, (1) straight.

53. ^Male PER eurvature, do view: (0) proeurved, (1) straight, (2) reeurved.

54. ^AME size: (0) smaller than AEE, (1) equal to ALE, (2) larger than AEE.

55. PME size: (0) smaller than PEE, (1) equal to PEE.

6

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BOSSELAERS J. & JOCQEIE R., Anew afrotropical genus of Cybaeodinae (Araneae, Eioeranidae)

56. ^AME size: (0) smaller than PME, (1) equal to PME, (2) larger than PME.

57. ^PME shape: (0) eireular, (1) modified.

58. *^MOQ: (0) wider anteriorly, (1) wider posteriorly.

59. *^PEinterdistanee: (0) equidistant, interdistaneelargerthanPE diameter, (1) equidistant, interdistanee

smaller than PE diameter, (2) PME farther from eaeh other than from PEE, interdistanee PME-PEE

smaller than PE diameter.

60. * ^Curved strong hairs frontally on abdomen: (0) absent, (1) present. Many ground spider genera

have rows of strong eurved hairs frontally on the abdomen (Bosselaers & Joeque 2002: fig. 2D).

61. ^Partial male do abdominal seutum: (0) absent, (1) present.

62. *^Male epigastrie selerite: (0) absent, (1) present.

63. ^Male modified ve abdominal setae: (0) absent, (1) present. Several lioeranid genera have males

with a pateh of stout, modified ve abdominal setae (Ubiek & Vetter 2005: 71; Wunderlieh 1999: 68,

2004: 1624, 2008: 489, 2011: 120, fig. 17a, 2012: 128).

64. *^Male modified ve abdominal setae field: (0) large, (1) small (Figs 2P, 3E, 6D, 9B, lOB).

65. ^Male AES shape: (0) eonieal, (1) eylindrieal (Figs 7E, 9B, lOB).

66. ^Male AES separation: (0) tangent or elose, (1) widely separated.

67. Male PMS shape: (0) slender, (1) stout, subtriangular (Fig. 7F).

68. Male PES separation: (0) about half of PES length, (1) at least equal to PES length.

69. ^Enlarged piriform spigots on male AES: (0) absent, (1) present (Fig. 7E). Enlarged piriform gland

spigots on AES of males only were first deseribed by Platniek for Clubiona Eatreille, 1804 and

Elaver O. P.-Cambridge, 1898 (Platniek 1990: 35), and later by Platniek & Di Franeo (1992: 4-6)

also for Cybaeodes, Agroeca, Neoanagraphis, md Rhaeboctesis Simon, 1897. Bosselaers & Joeque

(2002: ehar. 112, fig. 2KE) also mention them fox Agraecina and Liocranoeca. Wami & Joeque

(2002: 307) mention the eharaeter for Toxoniella. Wunderlieh attributes the eharaeter to Cybaeodes

and Clubioninae Simon, 1897 (2004: 1625, 2011: 27), and wrongly mentions it fox Apostenini

Wunderlieh, 2008 (2008: 488, but eompare Ubiek & Vetter 2005: 69, fig. 15).

70. *^Female epigastrie selerite: (0) absent, (1) present.

71. ^Female PMS shape: (0) slender, (1) laterally eompressed, (2) stout, subtriangular (Fig. 7D).

72. ^^Arrangement of large spigots on female PMS: (0) a single one, (1) three in a triangle (Fig. 7A, D),

(2) five in two rows, (3) more than five in two rows.

73. Female PES separation: (0) about half of PES length, (1) at least equal to PES length.

74. Male eymbial tip: (0) wide and short, (1) narrowed and long.

75. ^Dorsal bristle mat on male palpal eymbium: (0) absent, (1) present. A do palpal bristle mat was

mentioned for Zoropsidae Bertkau, 1882 by Eevy (1990: fig. 7) and Bosselaers (2002: 145) It oeeurs

in many families, for example in several eorinnid genera.

76. * ^Dorsal palpal bristle mat size: (0) large, (1) small.

77. *^Tegularloekinglobe: (0)absent, (1)present. Deseribedby Griswold(1993: l,fig. 19)forEyeosoidea

Eehtinen, 1967 as a tegular lobe interloeking with a eorresponding lobe on the subtegulum. In

Eioeranidae, Agraeeina laeks the tegular lobe but has the subtegular one.

78. * ^Subtegular loeking lobe: (0) absent, (1) present.

79. *^Coiled sperm duet: (0) absent, (1) present. A eoiled sperm duet in the bulbus is attributed to

Corinninae Karseh, 1897 by Platniek & Baptista (1995: 5).

80. *^Conduetor: (0) absent, (1) present.

81. ^Conduetor: (0) selerotised, (1) membranous.

7

European Journal of Taxonomy 1-49 ( 2013 )

Fig. 2. A-D. Cteniogaster conviva sp. nov. A. habitus dorsal view. B. Idem, ventral view. C.

habitus, dorsal view. D. Idem, ventral view. E-H. Cteniogaster toxarchus gen. et sp. nov. E. S, habitus

dorsal view. E. Idem, ventral view. G. $, habitus, dorsal view. H. Idem, ventral view. I-P. Cteniogaster

hexomma sp. nov. I. $, prosoma, dorsal view. J. Idem, habitus, dorsal view. K. Idem, abdomen, ventral

view. L. Idem, habitus, ventral view. M. Idem, S, habitus, dorsal view. N. Idem, prosoma, dorsal view.

O. Idem, habitus, ventral view. P. Idem, abdomen, ventral view. Seale bars: A-0 = 0.5 mm; P = 0.2 mm.

8

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BOSSELAERS J. & JOCQEIE R., Anew afrotropical genus of Cybaeodinae (Araneae, Eioeranidae)

82. ^Conduetor shape: (0) simple, (1) eomplex.

83. *^MA: (0) absent, (1) present. The MA is a selerotised, artieulated tegular proeess arising from a

membranous area(Bonaldo 1997: 166; Griswold 1993: 10, ehar. 12; Sierwald 1990: 21).

84. MA shape: (0) simple hook, (1) eurled and bifid.

85. Embolus insertion: (0) apieal, (1) prolateral.

86. ^Embolus transseetion: (0) solid, (1) flattened.

87. ^Embolus shape: (0) a short point, (1) siekle-shaped, (2) platelike, (3) hook-shaped, (4) a flat ribbon,

(5) whip-shaped.

88. ^Additional apophyses: (0) none, (1) additional ones at embolus base.

89. *^Subtegulum: (0) pi, (1) pi and rl.

90. ^Basal rl bulge with thiekened rim on eymbium: (0) absent, (1) present.

91. *^Prolateral terminal lobe on male palpal ti: (0) absent, (1) present. Sueh a lobe (Bosselaers &

Joeque 2002: flg. 4H) was mentioned by Ubiek & Platniek (1991: 11) for Liocranum E. Koeh, 1866,

Mesiotelus Simon, 1897, and Hesperocmnum Ubiek & Platniek, 1991.

92. Epigynal selerotisation: (0) weak, (1) substantial.

93. Epigynal seape: (0) absent, (1) present.

94. ^Epigynal anterior hood (Bosselaers & Joeque 2002: flg. 4A, C, F): (0) absent, (1) present.

95. ^Epigynal septum: (0) absent, (1) present.

96. Eateral epigynal hooks: (0) absent, (1) present.

97. *^Epigyne CO position: (0) anterior, (1) posterior.

98. *^ST2: (0) absent, (1) present. For a diseussion of ST2, see Bosselaers & Joeque (2000: 15, ehar.

84).

99. ST2 size: (0) smaller than STl, (1) larger than STl.

Results

The matrix of eharaeter states, as well as the eharaeter parameters on the preferred tree, ean be found in

Appendix 3. All eharaeters were run unordered in the analyses performed.

In order to avoid ambiguous results introdueed by speeies with many missing entries, the data matrix

ineludes only speeies of whieh males and females were available. An equally weighted analysis of the

data matrix was performed in PAUP with hsearch addseq=random nreps=10000 (heuristie seareh

with tree biseetion and reeonneetion swapping and 10000 random addition sequenees). In order to avoid

spurious resolution due to unsupported (Coddington & Seharff 1994; Wilkinson 1995) or ambiguously

supported (Nixon & Carpenter 1996) branehes, those with a minimum length of zero were eollapsed

with condense collapse=minbrlen. Thirty-one shortest trees of length 312 were found, belonging to

two tree islands of 19 and 12 trees, respeetively. The striet eonsensus of these 31 trees is fully eompatible

with but less resolved than the preferred tree in Fig. 1.

Beeause eladograms obtained by attributing a posteriori weights to eharaeters based on their relative

degrees of homoplasy on a set of heuristie trees explain the data better (Bosselaers & Joeque 2002;

Goloboff et al. 2008b), a weighted analysis was also performed. Implied weighting (Goloboff 1993)

was preferred for this purpose. When applying implied weighting in PAUP with pset goloboff = yes,

hsearch addseq = random nreps = 5000, collapsing branehes with a minimum length of zero with

condense collapse=minbrlen, three Attest trees with At = -68.50714 (PAUP attributes a negative sign

to lit values) were found. The default value for the eoneavity constant was used, which equals 2 in

PAUP, since that programme uses a concavity constant k as defined in the fit formula f = (k + + k

9

European Journal of Taxonomy 1-49 ( 2013 )

+ 1 - mi) = (k + l)/(es + k + 1) in Goloboff (1993). An additional implied weighting seareh run in TNT

using Settings / implied weighting and Analyze / Traditional search as well as New Technology

search (eoneavity constant = 3) produced another fit value (30.49286) for the same trees, because TNT

calculates weighted homoplasy (a complement of fit) for implied weighting trees. The strict consensus

of the three fittest trees also has fit = -68.50714, length = 313, ci = 0.412, ri = 0.725, hi = 0.588, and re

= 0.299. This tree is our preferred solution (Fig. 1).

Bremer support values (Bremer 1988, 1994) for the preferred 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 and combining various numbers of replications (between 1 and 10) with various numbers of trees

saved per replication (between 3000 and 30000, inversely related to the numbers of replications) until

the solution stabilised.

Taxonomy

Class Arachnida Cuvier, 1812

Order Araneae Clerck, 1757

Family Liocranidae Simon, 1897

Subfamily Cybaeodinae Simon, 1893

Cteniogaster gen. nov.

um:lsid:zoobank.org:act:4D38E26E-FE0F-4FD9-ADB2-9B4728BE56AD

Type species

Cteniogaster toxarchus sp. nov.

Diagnosis

Cteniogaster gen. nov. differs from all other genera of Eiocranidae by the presence of a small posterior

ve abdominal field of strong setae in males, the presence of two pairs of tenent hairs on the tarsal tips and

the consistent presence of one plv and one rlv spine on mt IV. The genus differs from all other liocranid

genera QxcQpt Apostenus by the wide patellar indentations.

Etymology

The genus name is derived from the Greek ktsvlov, little comb, and yaoTrip, belly, referring to the

small array of strong, modified ventral setae on the abdomen of the males of the new genus. The gender

is feminine.

Description

Small (males 1.7-3.0, females 2.0-3.0) spiders. Carapace longer than wide, almost fiat but slightly

higher at fovea (Fig. 9F, G), smooth (Fig. 6A), yellow or brown, covered with feathery hairs (Fig. 6B),

unicoloured or with faint pattern, iridescent in some species. Distinct fovea in posterior half MOQ

widest posteriorly (Fig. 9H). Eight, six or four eyes, ringed with black. AME dark, EE pearl, PME very

pale and in most species reduced, or absent. AME smaller than EE. Both eye rows straight in frontal

view (Fig. 12E). Chilum single, small and subtriangular, indistinct or absent in the smaller species.

Chelicerae small, yellow or brown, with a knee-shaped shaggy hair in front of fang base. Promarginal

cheliceral rim with three teeth at a small distance from fang base, largest one in the middle. Retromarginal

cheliceral rim with two small teeth close to fang base. Eabium subtriangular, about as wide as long, with

a thickened anterior rim. Endites with a lateral notch and an apical serrula, no apical hair tuft. Sternum

10

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BOSSELAERS J. & JOCQEIE R., Anew afrotropical genus of Cybaeodinae (Araneae, Eioeranidae)

Fig. 3. A-B. Cteniogaster sangarawe sp. nov. A. 5 , habitus, ventral view. B. Idem, dorsal view.

C-D. Cteniogaster taxorchis sp. nov. C. $, prosoma, dorsal view. D. Idem, ventral view. E-G. Cteniogaster

nana sp. nov. E. S, habitus, ventral view. F. Idem, abdomen, ventral view. G. Idem, habitus, dorsal

view. H-J. Cteniogaster lampropus sp. nov. H. $, prosoma, dorsal view. I. Idem, habitus, dorsal view.

J. Idem, ventral view. Seale bars: A-D, F-J = 0.5 mm; E = 0.2 mm.

11

European Journal of Taxonomy 1-49 ( 2013 )

shield-shaped, smooth, unieoloured yellow or brown, in some speeies with a thin border. Three pairs of

weak PCX present, ICS absent.

Abdomen in different shades of grey, with an anterior row of eurved strong hairs in both sexes (Figs 2N,

9A), and with broad, flattened feathery hairs on ve surfaee (Fig. 6E, F). The male abdomen has a small

and weak anterior do seutum and a small ve posterior pateh of strong setae (Figs 2B, F, P, 3E, 6D, E, 9B,

lOB). Males with AES eylindrieal and separated from eaeh other, bearing apieal EPOS (Fig. 7E), PMS

and PES thin and slender, PES separated by more than their length and having a dome-shaped apieal

segment (Fig. 7E, F). Females with non-eontiguous eylindrieal AES with one major ampullate and a

small number of unmodifled piriform gland spigots, subtriangular PMS with three eylindrieal and one

minor ampullate gland spigot and non-eontiguous subeylindrieal PES about as large as AES bearing

two eylindrieal gland spigots and about a dozen aeiniform gland spigots (Fig. 7A-D). Eegs pale yellow

to brown, unieoloured, irideseent in some speeies, with triehobothria with basal soeket (Fig. 6G) and

oval tarsal organ (Fig. 61). Patellar indentation wide, length 2/3 of pa length in legs 1 to 111, 1/2 of patella

length in leg IV. Retroeoxal hymen present, but often pale and weak. Troehanter noteh present. Tarsus

IV bent with pseudoartieulations in males (Fig. IID), and slightly bent in females. Eeg formula 4123.

Tarsal tips without true elaw tufts but with two pairs of tenent hairs (Figs 6H, IIC, E). Metatarsus IV

with one plv and one rlv spine, both in proximal half of artiele (Appendix 1). Minim al leg spination for

the genus: males fe: palp do 0-1-1; 1 pi 0-0-1, do 1-1-0; 11 do 1-1-0; 111 do 1-1-2; IV do 1-1-0; pa: palp

do 0-1; ti: palp pi 2-0-1; 1 plv 1-0-1, rlv 1-0-1; 11 rlv 1-0-1; 111 pi 1-0-1, do 1-0-0, rl 0-0-1, plv 1-1-1, rlv

0-0-1; IV pi 1 -0-1, do 1 -0-0, rl 1 -0-1, plv 1 -1 -1, rlv 0-1 -1; mt: 1 plv 1 -0-0, rlv 1 -0-0; 11 plv 1 -0-0, rlv 1 -0-

0; 111 pi 0-0-1, do 0-2-2, rl 1-0-1, ve 2-0-1; IV pi 0-0-1, do 2-2-2, rl 1-0-1, ve 2-0-1; ta: palp pi 1-0-1, do

1-0-0; females fe: palp do 0-1-1; 1 pi 0-0-1, do 1-1-0; 11 do 1-1-0; 111 do 1-1-2; IV do 1-1-0; pa: palp pi

1-0, do 0-1; ti: palp pi 2-0-1, do 1-0-1; 1 plv 1-0-1, rlv 1-0-1; 11 rlv 1-0-1; 111 pi 1-0-1, do 1-0-0, rl 0-0-1,

plv 1-1-1, rlv 0-1-1; IV pi 1-0-1, do 1-0-0, rl 1-0-1, plv 1-1-1, rlv 0-1-1; mt: 1 plv 1-0-0, rlv 1-0-0; 11 plv

1-0-0, rlv 1-0-0; 111 pi 0-0-1, do 0-2-2, rl 1-0-1, ve 2-0-1; IV pi 0-0-1, do 2-2-2, rl 1-0-1, ve 2-0-1; ta:

palp pi 2-0-1, do 1-0-0, rl 0-1-0, ve 0-0-2.

Male palp with a simple, blunt or pointed RTA, a eymbium with a spine on the distal pi edge (Figs 9C,

IOC), a pi subtegulum, a short apieal embolus, a membranous or selerotised eonduetor and a simple,

subtriangular or subreetangular MA (Fig. 11 A, B). Epigyne with an anterior hood and a eentral depression

(Figs 9E, lOE, 12C-G, K-E). Vulva with isodiametrie, oval or kidney-shaped STl with internal spikes

and ST2 with a porous glandular strueture (Figs 4, 5, 8).

Distribution

The genus is known from the Taita hills in Kenya and from the East and West Usambara mountains in

Tanzania. So far no speeimens of Cteniogaster gen. nov. have been found outside the Eastern Are (Fig.

13), whieh is a fair indieation that the genus is endemie to that mountain range.

Identification Key

Females

(not known for C. nana sp. nov.)

1. Frontal hood on epigyne large, wider than half epigyne width (Fig. 5) . 2

- Frontal hood on epigyne narrow, less than one fourth epigyne width (Fig. 4). 4

2. Frontal hood as large as half epigyne width; spermatheeae 1 pear shaped, oriented diagonally,

spermatheeae 2 oriented transversally, touehing in the middle (Fig. 5E-F) . C. taxorchis sp. nov.

- Frontal hood almost as wide as epigyne; spermatheeae 1 rounded, spermatheeae 2 oriented along

longitudinal axis (Fig. 5A-D) . 3

12

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BOSSELAERS J. & JOCQEIE R., Anew afrotropical genus of Cybaeodinae (Araneae, Eioeranidae)

3. Frontal hood very wide and selerotized; spermatheeae 2 eurved, slightly separated, not touehing

(Fig. 5C-D) .C. sangarawe sp. nov.

- Frontal hood less wide and hardly selerotized; spermatheeae 2 sinuous, touehing in the middle

(Fig. 5A-B) .C. conviva sp. nov.

4. Anterior hood in larger oval depression; spermatheeae 1 piriform, widely separated (Fig. 4E-F) ....

. C. lampropus sp. nov.

- Anterior hood not surrounded by oval depression; spermatheeae 1 not piriform . 5

5. Spermatheeae 1 very large, oval, touehing in the middle (Fig. 4C-D) . C. hexomma sp. nov.

- Spermatheeae 1 mueh smaller, not touehing (Fig. 4A-B) .C. toxarchus sp. nov.

Males

(not known for C. lampropus sp. nov., C. sangarawe sp. nov. and C. taxorchis sp. nov.)

1. Cymbium with retrolateral bulge; selerotized eonduetor large and eoiled; RTA large, hookshaped,

sharp (Fig. 12A, B) .C. conviva sp. nov.

- Without eymbial bulge; eonduetor small; RTA less large or straight or blunt (Figs 9C-D, lOC-D,

12A-B) . 2

2. RTA very short, sharp and straight (Fig. lOC-D). C. hexomma sp. nov.

- RTA longer and blunt or eurved upwards (Figs 9C-D, 12H-I) . 3

3. RTA straight, obliquely truneated; eonduetor membranous (Fig. 9C-D) . C. toxarchus sp. nov.

- RTA eurved upwards, truneated tip; eonduetor selerotized (Fig. 12H-I). C nana sp. nov.

Cteniogaster toxarchus sp. nov.

um:lsid:zoobank.org:aet:CBA4D351-036Q-4F20-A6ED-CF4CCB6E7BBB

Figs 2E-H, 4A-B, 8D-F, 9, 11 A, E, 14; Appendix 1C, F

Diagnosis

Cteniogaster toxarchus sp. nov. differs from all other Cteniogaster gen. nov. speeies by the male palp

with a straight, blunt, subtrapezoidal RTA and a pointed subtriangular rl apieal MA with a pi membranous

border, and by the vulva with widely separated, kidney-shaped median STl.

Etymology

The speeies name is a noun in apposition and is derived from the Greek xo^apxog, eommander of

the arehers, referring to the array of arrow-like setae on the posterior ve part of the male abdomen in

Cteniogaster gen. nov., and to the present speeies’ status as typus of the genus.

Type material

Holotype

TANZANIA, E. Usambara Mts., Kwamkoro Forest Reserve, 5°10.9’S 38°35.8’E, 6 Nov. 1995,

950 m asl, Griswold C., Seharfif N., Ubiek D. (ZMUC).

Paratypes

4 $ 5 3 j.: same data as holotype; 17 S3, 25 $ $, 5 j.: TANZANIA, E. Usambara Mts., Amani, 5°5.7’S

38°38’E, 28 Oet.-9 Nov. 1995, 950 m asl, sifting litter, Griswold C., Seharff N., Ubiek D. (ZMUC);

3 $ 5 j.: as previous, 27 Get.-9 Nov. 1995; 18 S3 1 ? 1 j.: as previous; 1 (J 3 $ $ 1 j.: as previous,

Mbomole Hill, 5°5.7’S 38°37’E, 5-8 Nov. 1995, sifting litter, 100 m asl; 5 2 Tanga, W.

Usambara Mts., Mazumbai Forest, 4°49’S 38°30’E, 11-19 Nov. 1995, pitfall traps, 1400-1600 m asl.

13

European Journal of Taxonomy 1-49 ( 2013 )

Griswold C., ScharffN., Ubick D. (ZMUC); >100 ?: as previous, sifting litter, 1400-1800 m asl;

3 SSA $$, 15 j. Mazumbai Forest, 4°49’S 38°29.5’E, 12-20 Nov. 1995, 1800-1900 m asl, sifting litter,

Griswold C., SeharffN., Ubiek D. (ZMUC; 2 SS 2 $ $ in MRAC); 2 2 $ Tanga, W. Usambara

Mts., Mazumbai Forest, 4°49’S 38°30’E, 11-20 Nov. 1995, sifting litter, 1400-1800 m asl, Griswold C.,

SeharffN., Ubiek D. (CAS).

Description

Male

From Mazumbai (CAS)

Total length. 2.47. Carapaee length 1.10, width 0.88, yellow brown, somewhat irideseent, with a thin

border and grey mottling on sides (Figs 2E, 9A). Fovea brown, pronouneed, length 0.13, anterior end

0.66 from front end of earapaee. MOQ length 0.13, anterior width 0.11, posterior width 0.16, AER width

0.24, PER width 0.32. All eyes subeireular, EE equal in size, their diameter more than twiee the diameter

of ME, whieh are also equal in size. AME separated by less than their diameter, almost touehing AEE.

PME separated by 2.5 times their diameter, 1.5 diameters from PEE (Fig. 9A). Both eye rows reeurved

from above. Clypeus vertieal, equal to diameter of AME. Chilum pale, about the size of the AME group.

Chelieerae yellow brown. Eabium slightly wider than long, half as long as endites. Sternum yellow, with

a thin border, length 0.66, width 0.58. PCT weak and pointed.

Abdomen. Grey dorsally, eovered with long thin hairs, with brown, poorly defined, subtriangular anterior

do seutum eovering less than 10 % of the do surfaee area. Ventral side of abdomen greyish white, with

,- 1

Cteniogaster toxarchus

t -f

Cteniogaster hexomma

I-1

Cteniogaster lampropus

Fig. 4. Female genitalia, ventral view. A, C, E: eleared in methyl salieylate. A-B. Cteniogaster toxarchus

gen. et sp. nov. C-D. Cteniogaster hexomma sp. nov. E-F. Cteniogaster lampropus sp. nov. Seale

bars = 0.1 mm.

14

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BOSSELAERS J. & JOCQEIE R., Anew afrotropical genus of Cybaeodinae (Araneae, Eioeranidae)

a brown selerotised elliptie posterior pateh earrying long, strong setae (Figs 2F, 9B). AES separated

by their diameter. Eegs pale brown, somewhat irideseent. Retroeoxal hymen pronouneed, subglobular.

Tarsus IV bent. Eeg spination as in Appendix 1C.

Eeg measurements:

fe

pa

ti

mt

ta

Total

I

0.92

0.37

0.79

0.63

0.53

3.23

II

0.79

0.37

0.60

0.53

0.47

2.76

III

0.71

0.26

0.53

0.60

0.47

2.58

IV

0.95

0.39

0.87

0.60

3.68

Maee paep. As illustrated (Figs 9C-D, 11 A), with a straight, blunt, subtrapezoidal RTA, a short, pointed, pi

apieal embolus, a median, fan shaped membranous apieal eonduetor and a simple pointed subtriangular

rl apieal MA with a pi membranous border (Fig. 11 A).

Female (holotype)

Totae eength. 2.68. Carapaee length 1.16, width 0.95, brown, laterally mottled as in male (Fig. 2G).

Fovea pronouneed, length 0.16, anterior end 0.66 from front end of earapaee (Fig. 9H). MOQ length

0.11, anterior width 0.11, posterior width 0.15. AFR width 0.27, PER width 0.36. Relative eye sizes, eye

row eurvatures and elypeus as in male. Chilum as in male, but wider. Sternum brown, with a thin border,

length 0.71, width 0.66. PCX as in male.

Cteniogaster conviva Cteniogaster sangarawe Cteniogaster taxorchis

Fig. 5. Female genitalia, dorsal view. A, C, E: eleared in methyl salieylate. A-B. Cteniogaster conviva

sp. nov. C-D. Cteniogaster sangarawe sp. nov. E-F. Cteniogaster taxorchis sp. nov. Seale bars = 0.1 mm.

15

European Journal of Taxonomy 1-49 ( 2013 )

Abdomen. Coloured as in male, but without do seutum or ve modified setae. Legs pale brown, somewhat

irideseent. Retroeoxal hymen large, oval, subglobular and pearly white. Leg spination as in Appendix IF.

Leg measufrements:

fe

pa

ti

mt

ta

Total

I

1.00

0.42

0.79

0.60

0.50

3.31

II

0.89

0.34

0.68

0.53

0.47

2.92

III

0.79

0.32

0.58

0.66

0.47

2.81

IV

1.08

0.39

0.89

1.03

0.63

4.02

Epigyne. With a small, narrow anterior hood, and two posterior lateral ridges, semitransparent and

showing spermateeae in posterior half (Figs 2H, 9E). Vulva with widely separated, kidney-shaped

median STl with thin internal spikes and posterior ST2 earrying a porous glandular strueture (Figs 4A-

B, 8D-F).

Distribution

Tanzania, East and West Usambara mountains, 950-1800 m asl.

Cteniogaster hexomma sp. nov.

urn:lsid:zoobank.org:act: 1893EEE5-2CBC-43 AA-9669-4A2A293A89CD

Figs 21-P, 4C-D, 6, 7, 8G-H, lOA-E, IIB-D, 14; Appendix ID, G

Diagnosis

Cteniogaster hexomma sp. nov. differs from all other Cteniogaster gen. nov. speeies by the eonsistent

absenee of PME, by a male palp with a small, straight and pointed RTA and a simple, blunt and seoop-

shaped rl apieal MA, and by the vulva with large, touehing, oval STl with eonspieuous internal spikes

and smaller, more dorsally positioned posterior ST2 earrying an anterior porous glandular strueture.

Etymology

The speeies name is a noun in apposition and refers to the eonsistent absenee of PME in this speeies,

resulting in an array of six eyes.

Type material

Holotype

5 and paratype female: KENYA, Taita Hills, Ngangao Forest, 3°22’S 38°2LE, 15 Jul. 1998, mixed

forest, pitfall trap, Rogo E. (MRAC 208888).

Paratypes

I together with holotype; 1 S\ KENYA, Taita Hills, Ngangao Forest, 3°22’S 38°2LE, 15 Jul. 1998,

4 Dee. 1999, Winkler extraetion of forest litter, VandenSpiegel D. & Miehiels J. (MRAC 209203); 1

as previous, 11 Mar. 2004, pitfalls, Spanhove T. & Chovu M. (MRAC 222071); 2 SS\ as previous, 11

Mar. 2004 (MRAC 222412); 4 as previous, 11 Mar. 2004 (MRAC 222419); 2 as previous,

II Mar. 2004 (MRAC 222426); 1 as previous, 7 Mar. 2004 (MRAC 222431); 1 S\ as previous, 11

Mar. 2004 (MRAC 222439); 1 S\ as previous, 11 Mar. 2004 (MRAC 222444); 1 S\ as previous, 11

Mar. 2004 (MRAC 222449); 1 (?: as previous, 7 Mar. 2004 (MRAC 222076); 2 SS, 2 $ ?: as previous,

8 Feb. 2004 (MRAC 222084); 1 as previous, 8 Feb. 2004 (MRAC 222110); 1 S\ as previous, 10

Feb. 2004 (MRAC 222086); 1 as previous, 8 Feb. 2004 (MRAC 222088); 2 SS, 2 5 as previous,

6 Feb. 2004 (MRAC 222089); 1 ?: as previous, 6 Feb. 2004 (MRAC 222092); 6 Feb. 2004 (MRAC

222100); 6 Feb. 2004 (MRAC 222109); 4 as previous, 6 Feb. 2004 (MRAC 222107); 1 (?: as

16

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BOSSELAERS J. & JOCQEIE R., Anew afrotropical genus of Cybaeodinae (Araneae, Eioeranidae)

Fig. 6. Cteniogaster hexomma sp. nov. A. 3, earapaee, dorsal view. B. Detail of previous. C. setae on

venter of abdomen. D. S, group of modified setae on venter of abdomen. E. Detail of previous. F. Detail

of seta surrounding pateh of modified setae. G. triehobothrium on tibia I. H. tarsal elaws leg I.

I. tarsal organ leg I. Seale bars: A = 0.5 mm; D = 50 pm; B, H = 20 pm; C, E = 10 pm; F, G, I = 5 pm.

17

European Journal of Taxonomy 1-49 ( 2013 )

Fig. 7. Cteniogaster hexomma sp. nov. A-D. A. Spinnerets, posterior view. B. ALS. C. PLS. D. PMS.

E-F. (f. E. Spinnerets, posterior view. E. PLS + PMS. Seale bars: A, E-F = 50 pm; C-D = 20 pm;

B = 10 pm.

18

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BOSSELAERS J. & JOCQEIE R., Anew afrotropical genus of Cybaeodinae (Araneae, Eioeranidae)

previous, 10 Feb. 2004 (MRAC 222096); 2 (?(?: as previous, 8 Feb. 2004 (MRAC 222097); 3

2 as previous, 2 SS, 1 as previous, 10 Feb. 2004 (MRAC 222101); 1 S\ as previous, 10 Feb.

2004 (MRAC 222094); 1 (?: as previous, 1 ?: as previous, 7 Mar. 2004 (MRAC 222392); 1 (?, 1 ?:

as previous, 10 Apr. 2004 (MRAC 223157); 1 as previous, 8 Apr. 2004 (MRAC 223165); 1 S', as

previous, 6 Apr. 2004 (MRAC 223230).

Other material examined

1 S'. KENYA, Taita Hills, Chawia Forest, 03°29’S 038°20’E, 18-24 Mar. 1998, pitfalls, Rogo Fuey

(MRAC 208374); 1 S'- as previous, 16 Jul. 1998 (MRAC 208886); 1 as previous, 22 Feb. 2004,

pitfalls, Spanhove T. & Chovu M. (MRAC 221973); 3 SS'- as previous (MRAC 222090); 1 as

previous (MRAC 222095); 1 S'- as previous (MRAC 222004); 1 (?, 1 as previous, 23 Feb. 2004,

pitfalls (MRAC 222012); 1 as previous (MRAC 222085); 1 S'- as previous, 24 Feb. 2004 (MRAC

222091); 1 as previous, 8 Mar. 2004 (MRAC 222475); 1 S'- as previous, 22 Mar. 2004 (MRAC

222784); 1 S'- as previous (MRAC 222789); 1 S'- as previous (MRAC 222883); 1 S'- as previous,

24 Apr. 2004 (MRAC 223253); 1 ?: as previous, 22 Apr. 2004 (MRAC 223271); 1 ?: Taita Hills,

Yale Forest, 03°39’S 038°33’E, 6 Dee. 1999, Winkler extraetion of forest litter, VandenSpiegel D. &

Miehiels J. (MRAC 209188); 1 (?: Taita Hills, Maeha Forest, 03°25’S 028°2rE, 13 Feb. 2004, pitfalls,

Spanhove T. & Chovu M. (MRAC 222087); 1 6': as previous (MRAC 222093); 2 SS'- Taita Hills,

Fururu Forest, 03°26’S 038°20’E, 19 Feb. 2004, pitfalls, Spanhove T. & Chovu M. (MRAC 222098);

1 as previous (MRAC 222104); 1 S'- as previous (MRAC 222105); 1 S'- as previous, 19 Mar. 2004

(MRAC 222860); 1 S'- as previous (MRAC 222892); 1 S'- as previous, 19 Apr. 2004 (MRAC 223289);

1 S'- Taita Hills, Mwaehora Forest, 03°25’S 038°22’E, 15 Feb. 2004, pitfalls, Spanhove T. & Chovu M.

(MRAC 222108); 1 as previous, 15 Mar. 2004, pitfalls, Spanhove T. & ChovuM. (MRAC 222830);

2 SS'- as previous, 15 Mar. 2004 (MRAC 222846); 2 SS'- as previous (MRAC 222855); 1 Taita

Hills, Wundanyi Forest, 03°24’S 038°22’E, 17 Feb. 2004, pitfalls, Spanhove T. & Chovu M. (MRAC

222099); 2 (?(?: as previous (MRAC 222102); 1 (?: as previous, 17 Apr. 2004 (MRAC 223222).

Description

Male (holotype)

Total length. 2.92. Carapaee length 1.34, width 0.97, brown, somewhat irideseent with green and

purple hues, unbordered (Fig. 2M-N). Fovea pronouneed, length 0.13, anterior end 0.89 from front end

of earapaee. AER width 0.21, PER width 0.29. AME diameter 1/3 of AFE, separated by their diameter,

almost touehing AFE, PEE as large as AFE, PME absent (Fig. lOA). Clypeus vertieal, equal to 1.5 times

the diameter of AME. Chilum about the size of the AME group. Sternum brown, with a thin border,

length 0.76, width 0.63. PCT brown and pointed.

Abdomen. Pale greyish brown dorsally, eovered with flattened, irideseent hairs, provided with small,

brown, poorly deflned anterior do seutum. Ventral side of abdomen pale grey, with a brown selerotised

elliptie posterior pateh earrying long, strong setae (Figs 20-P, 6D-E, lOB). AES separated by their

diameter, PFS with slender eonieal apieal segment, separated by their length. Fegs brown, irideseent.

Sparse ve terminal preening brushes on mt 111 and IV. Retroeoxal hymen oval, white, subglobular. Tarsus

IV bent. Feg spination as in Appendix ID.

Feg measurements:

fe

pa

ti

mt

ta

Total

I

0.79

0.34

0.63

0.47

0.42

2.66

II

0.68

0.34

0.55

0.45

0.42

2.45

III

0.66

0.29

0.45

0.50

0.47

2.37

IV

0.87

0.34

0.74

0.79

0.58

3.31

19

European Journal of Taxonomy 1-49 ( 2013 )

Male palp. As illustrated (Figs lOC-D, IIB), with a small, straight and pointed RTA, a short, pointed,

pi apieal embolus, a median, fan shaped membranous apieal eonduetor and a simple, blunt and seoop-

shaped rl apieal MA (Fig. IIB).

Female

Total length. 2.97. Carapaee length 1.10, width 0.88, eoloured as in male (Fig. 21-J). Fovea pronouneed,

length 0.18, anterior end 0.76 from front end of earapaee. AER width 0.23, PER width 0.31. Relative

eye sizes, AER eurvature and elypeus as in male. PME absent. Chilum a small, yellow brown equilateral

triangle beneath AME group. Sternum yellow brown, with a thin border, length 0.68, width 0.63 (Fig.

2L). PCX pointed but weakly selerotised.

Abdomen. Coloured as in male, but without do seutum or ve modified setae. ALS separated by their

diameter, PLS with hemispherieal apieal segment, separated by their length. Legs yellow brown,

irideseent. Retroeoxal hymen as in male. Troehanter noteh more prominent in legs 111 and IV. Preening

brushes as in male, tarsus IV slightly bent. Leg spination as in Appendix IG.

Leg measurements:

fe

pa

ti

mt

ta

Total

I

0.76

0.34

0.60

0.39

0.42

2.52

II

0.71

0.29

0.53

0.42

0.39

2.34

III

0.60

0.29

0.42

0.50

0.45

2.26

IV

0.82

0.32

0.71

0.79

0.60

3.23

Epigyne. With a small, narrow anterior hood, semitransparent and showing the eopulatory duets and the

large, oval STl (Figs 2K, lOE). Vulva with large, touehing, oval STl with eonspieuous internal spikes

and smaller, more dorsally positioned posterior ST2 earrying an anterior porous glandular strueture

(Figs 4C-D, 8G-H).

Distribution

Kenya, Taita hills, 1600-1900 m asl.

Cteniogaster conviva sp. nov.

um:lsid:zoobank.org:aet:EB0512BB-1926-4CC2-88D8-A541A5EDC2F7

Figs 2A-D, 5A-B, 8A-C, 12A-E, 14; Appendix IE, H

Diagnosis

Cteniogaster conviva sp. nov. differs from all other Cteniogaster gen. nov. speeies by the large male palp

with a sharply pointed, hook-shaped RTA, a large, eoiled and selerotised eonduetor, and a eymbium with

a rl basal bulge; and by the epigyne with a thin, broad anterior hood.

Etymology

The speeies name is a noun in apposition and is derived from the Latin conviva, table eompanion,

referring to the faet that this speeies shares the same habitat with C. toxarchus sp. nov. in the West

Usambara mountains.

Type material

Holotype

(?: TANZANIA, Tanga, W. UsambaraMts., Mazumbai Forest, 4°49’S 38°30E, 11-20 Nov. 1998, 1400-

1800 m asl, sifting litter, Griswold C., Seharff N. & Ubiek D. (ZMUC).

20

r

BOSSELAERS J. & JOCQEIE R., Anew afrotropical genus of Cybaeodinae (Araneae, Eioeranidae)

Paratype

1 $: together with holotype; 4 $ $: same data as holotype (ZMUC).

Description

Male (holotype)

Total length. 2.24. Carapaee length 1.05, width 0.82, unieoloured yellow, unbordered (Fig. 2A).

Fovea brown, pronouneed, length 0.13, anterior end 0.59 from front end of earapaee. MOQ length

0.09, anterior width 0.05, posterior width 0.08. AER width 0.19, PER width 0.24. AME small, diameter

1/3 of diameter ALE. PEE slightly smaller than AEE. All eyes touehing, exeept for PME whieh are

isolated, strongly redueed and flattened, diameter 1/3 of AME diameter (Fig. 12E). AER reeurved and

PER straight from above. Clypeus vertieal, slightly higher than diameter of AME. Chilum indistinet and

very small. Sternum yellow, without a distinet border, length 0.60, width 0.53. PCX pale yellow.

Abdomen. Pale greyish brown, with a weak anterior do seutum eovering 10 % of the do surfaee area and

a ve posterior pateh of strong setae as is typieal for the genus. AES separated by 1/3 of their diameter.

Eegs yellow. Retroeoxal hymen pale and weak. Troehanter noteh present, more prominent on legs III

and IV. Tarsus IV slightly bent. Eeg spination as in Appendix IE.

Eeg measurements:

fe

pa

ti

mt

ta

Total

I

0.79

0.32

0.60

0.42

0.47

2.60

II

0.66

0.32

0.53

0.39

0.42

2.31

III

0.55

0.26

0.39

0.42

0.39

2.03

IV

0.79

0.37

0.60

0.71

0.55

3.02

Maee paep. As illustrated (Fig. 12A-B), with a hook-shaped, sharply pointed RTA, a short apieal embolus,

a simple subtriangular MA and a eomplex, eoiled, selerotised eonduetor. The eymbium has a thiekened

basal rl bulge (Fig. 12B).

Female

Totae eength. 2.58. Carapaee length 1.08, width 0.82, unieoloured yellow, unbordered (Fig. 2A).

Fovea brown, pronouneed, length 0.13, anterior end 0.71 from front end of earapaee. MOQ length 0.09,

anterior width 0.06, posterior width 0.10. AER width 0.22, PER width 0.26. AME small, diameter 1/3 of

AEE diameter. PEE diameter 2/3 of AEE. All eyes touehing, exeept for PME. PME absent or strongly

redueed. Eye row eurvatures as in male. Clypeus vertieal, slightly higher than diameter of AME. Chilum

a small, yellow brown equilateral triangle, situated beneath AME. Sternum yellow, with a weak border,

length 0.66, width 0.59. PCX pale yellow.

Abdomen. Pale greyish brown, without do seutum or ve modified setae. Eegs yellow. Retroeoxal hymen

as in male. Troehanter noteh as in male. Sparse ve preening brushes on mt III and IV. Tarsus IV slightly

bent. Eeg spination as in Appendix IH.

Eeg measurements:

fe

pa

ti

mt

ta

Total

I

0.74

0.34

0.58

0.42

2.50

II

0.66

0.26

0.50

0.39

2.21

III

0.55

0.24

0.37

0.47

2.10

IV

0.87

0.34

0.66

0.71

0.55

3.13

21

European Journal of Taxonomy 40 : 1-49 ( 2013 )

Fig. 8. SEM pictures of female genitalia. A-C. Cteniogaster conviva sp. nov. A. Dorsal view, arrow

indieates perforations. B. Detail of previous. C. Idem, seen at different angle. D-F. Cteniogaster

toxarchus gen. et sp. nov. D. Dorsal view. E. Detail of other speeimen, arrow indieates perforations.

F. Detail of previous. G-H. Cteniogaster hexomma sp. nov. G. Dorsal view, arrow indieates peforations.

H. Detail of previous. Seale bars = A, D, G-H = 50 pm; B, C = 10 pm; E = 20 pm; F = 5 pm.

22

r

BOSSELAERS J. & JOCQEIE R., Anew afrotropical genus of Cybaeodinae (Araneae, Eioeranidae)

Epigyne. With thin, wide anterior hood, partly transparent, showing median ST2 and widely separated

posterior globular STl, slightly variable (Fig. 12C-D). Vulva with widely separated spherieal posterior

STl with internal spikes and median, longitudinally oriented, touehing ST2 with an anterior, porous

glandular strueture (Figs 5A-B, 8A-C).

Distribution

Tanzania, West Usambara mountains, 1400-1800 m asl.

Cteniogaster sangarawe sp. nov.

um:lsid:zoobank.org:aet:B3900F5F-6616-4AlB-8698-BClF20748842

Figs 3A-B, 5C-D, 12F-G, 14; Appendix IK

Diagnosis

Cteniogaster sangarawe sp. nov. is elose to C. conviva sp. nov. but differs from it and from all other

Cteniogaster gen. nov. speeies by the epigyne with a thiek and selerotised, wide anterior hood, and the

vulva with median, longitudinally oriented ST2 that are slightly separated from eaeh other.

Etymology

The speeies name is a noun in apposition and refers to Sangarawe Forest, the type loeality in the East

Usambara mountains.

Type material

Holotype

TANZANIA, E. Usambara Mts., Amani, Sangarawe Forest, 5°6.5’S 38°35.7’E, 5-6 Nov. 1995, 990

m asl, sifting litter, Griswold C., SeharffN. & UbiekD. (ZMUC).

Paratypes

1 $, 1 juv: TANZANIA, E. Usambara Mts., Amani, Sangarawe Forest, 5°5.7’S 38°38’E, 28 Oet.-9Nov.

1995, 950 m asl, pitfalls, Griswold C., SeharffN. & Ubiek D. (ZMUC); 2 $ $ : same data as previous

(ZMUC).

Description

Male

Unknown.

Female (holotype)

Total length. 2.24. Carapaee length 1.05, width 0.79, yellow brown, somewhat irideseent, unbordered

(Fig. 3B). Fovea pronouneed, length 0.13, anterior end 0.68 from front end of earapaee. MOQ length

0.08, anterior width 0.07, posterior width 0.08. AFR width 0.21, PER width 0.26. AMF small, 1/4-1/3

of AFF diameter. Fateral eyes touehing, AMF 1.5 times diameter of PEE. PME very small and redueed,

diameter 1/3 of AMF diameter. Both eye rows straight from above. Clypeus vertieal, equal to 1.5 times

the diameter of AME. Chilum orange brown, selerotised. Sternum yellow, length 0.63, width 0.58. PCT

very weak, subtriangular.

Abdomen. Pale yellowish grey dorsally (Fig. 3B). AES separated by half their diameter, PFS eonieal,

separated by their length. Fegs yellow. Retroeoxal hymen small but pronouneed, subglobular, hyaline.

Troehanter noteh present but not very pronouneed. Sparse ve terminal preening brushes on mt III and

IV. Tarsus IV slightly bent. Feg spination as in Appendix IK.

23

European Journal of Taxonomy 1-49 ( 2013 )

Leg measurements:

fe

pa

ti

mt

ta

Total

I

0.76

0.39

0.55

0.39

0.42

2.52

II

0.68

0.32

0.50

0.34

0.42

2.26

III

0.55

0.24

0.34

0.45

1.92

IV

0.79

0.32

0.66

0.68

0.60

3.05

Epigyne. with thick and sclerotised, wide anterior hood, partly transparent, showing median ST2 and

widely separated posterior globular STl, slightly variable (Fig. 12F-G). Vulva with widely separated

spherieal posterior STl with internal spikes and median, longitudinally oriented and slightly separated

ST2 with an anterior, porous glandular strueture (Fig. 5C-D).

Distribution

Tanzania, East Usambara mountains, 950-990 m asl.

Cteniogaster lampropus sp. nov.

um:lsid:zoobank.org:aet:54BD025D-0757-4750-ACEB-403E5B47E984

Figs 3H-J, 4E-F, 12F, 14; Appendix 11

Diagnosis

Cteniogaster lampropus sp. nov. differs from all other Cteniogaster gen. nov. speeies by the relatively

large PME, the strongly irideseent legs and the vulva with widely separated piriform posterior STl with

internal spikes and more median and dorsally situated, broad, longitudinally oriented ST2.

Etymology

The speeies name is a noun in apposition and is derived from the Greek Lapjtpog, shining, and Jtoug,

leg, referring to the strongly irideseent legs.

Type material

Holotype

TANZANIA, Tanga, W. Usambara Mts., Mazumbai Forest, 4°49’S 38°29.5’E, 12-20 Nov. 1995,

1800-1900 m asl, Griswold C., Seharff N. & Ubiek D. (ZMUC).

Description

Male

Unknown.

Female (holotype)

Totae eength. 2.42. Carapaee length LI8, width 0.97, brown, with grey radiating striae and dark grey

border (Fig. 3H-I). Fovea pronouneed, length 0.09, anterior end 0.83 from front end of earapaee. MOQ

length 0.13, anterior width 0.11, posterior width 0.16. AFR width 0.29, PER width 0.37. AME 0.4 of

AFE diameter. PEE 0.8 of ALE diameter, separated from them by PEE diameter. PME eireular, not

redueed, 2/3 of AME diameter. Both eye rows reeurved from above (Fig. 3H). Clypeus vertieal, equal to

AME diameter. Chilum brown, small. Sternum yellow with a darker brown border (Fig. 3J), length 0.74,

width 0.68. PCT pale, subtriangular.

Abdomen. Dark greenish grey dorsally, eovered with greenish irideseent silky hairs. Ventral side of

abdomen ereamy brown, darker around spinnerets (Fig. 3J). AES separated by half their diameter, PES

short and stout, subeylindrieal, separated by 1.5 times their diameter. Eegs orange, femora olive grey

24

r

BOSSELAERS J. & JOCQEIE R., Anew afrotropical genus of Cybaeodinae (Araneae, Eioeranidae)

(Fig. 3H), irideseent with a green and blue lustre. Retroeoxal hymen subglobular, hyaline. Troehanter

noteh more pronouneed in legs III and IV. Sparse ve terminal preening brushes on mt III and IV. Tarsus

IV slightly bent. Eeg spination as in Appendix II.

Eeg measurements:

fe

pa

ti

mt

ta

Total

I

0.85

0.34

0.76

0.53

3.01

II

0.76

0.34

0.63

0.53

0.50

2.76

III

0.76

0.29

0.53

0.58

0.50

2.66

IV

1.03

0.39

0.84

0.92

0.63

3.81

Epigyne. With a narrow anterior hood-like strueture, partly transparent, showing superimposed lateral

STl and ST2 (Fig. 12E). Vulva with widely separated piriform posterior STl with internal spikes and

more median and dorsally situated, broad, longitudinally oriented ST2 with an anterior, porous glandular

strueture (Fig. 4E-F).

Distribution

Tanzania, West Usambara mountains, 1800-1900 m asl.

Cteniogaster taxorchis sp. nov.

um:lsid:zoobank.org:aet:DBF6AC89-62A0-4233-A459-D8DB204E7D0E

Figs 3C, G, 5E-F, 12K, 14; Appendix IK

Diagnosis

Cteniogaster taxorchis sp. nov. differs from all other Cteniogaster gen. nov. speeies by the epigyne with

a relatively broad, thin, are-shaped anterior hood, and the vulva with lateral, diagonally oriented anterior

STl and posterior, transverse ST2 touehing in the middle.

Etymology

The speeies name is a noun in apposition and is derived from the Greek xa^ig, order, and opxig, testiele,

referring to the orderly symmetrieal arrangement of the large, testiele-shaped STl.

Type material

Holotype

TANZANIA, E. Usambara Mts., Kwamkoro Forest Reserve, 5°I0.9’S 38°35.8’E, 6 Nov. 1995,

950 m asl, Griswold C., Seharff N. & Ubiek D. (ZMUC).

Description

Male

Unknown.

Female (holotype)

Totae eength. 2.21. Carapaee length 1.00, width 0.74, unieoloured pale yellow, unbordered (Fig. 3C).

Fovea pronouneed, length 0.09, anterior end 0.63 from front end of earapaee. MOQ length 0.06, anterior

width 0.06, posterior width 0.08. AER width 0.18, PER width 0.22. AME 1/4, PME 1/3 and PEE 0.7 of

AFE diameter. PME flattened and redueed. Both eye rows reeurved from above. Clypeus vertieal, equal

to twiee AME diameter. Chilum indistinet. Sternum pale yellowish white with a yellow border (Fig. 3G),

length 0.58, width 0.53. PCT indistinet.

25

European Journal of Taxonomy 1-49 (2013)

Abdomen. Pale greyish white dorsally. Spinnerets as for the genus in general, but shorter. Legs yellowish

white (Fig. 3G). Retroeoxal hymen small, oval, pale. Troehanter noteh more pronouneed in legs 111 and

IV. Sparse ve terminal preening brushes on mt 111 and IV. Tarsus IV elearly bent. Leg spination as in

Appendix IK.

Leg measurements:

fe

pa

ti

mt

ta

Total

I

0.74

0.34

0.58

0.42

2.50

II

0.63

0.26

0.45

0.37

0.39

2.10

III

0.53

0.24

0.37

0.42

0.39

1.95

IV

0.79

0.28

0.61

0.66

0.55

2.89

Epigyne with relatively broad, thin, are-shaped anterior hood, partly transparent, showing large oval

anterior STl and transverse posterior ST2 (Fig. 12K). Vulva with lateral, diagonally oriented anterior

STl with eonspieuous internal spikes and posterior, transverse ST2 touehing in the middle (Fig. 5E-F).

Distribution

Tanzania, East Usambara mountains, Kwamkoro Forest, 950 m asl.

Cteniogaster nana sp. nov.

um:lsid:zoobank.org:aet:B0321C9D-7120-407A-BAAF-BCF85186FDA0

Figs 3D-F, 12H-J, 14; Appendix IB

Diagnosis

Cteniogaster nana sp. nov. differs from all other Cteniogaster gen. nov. speeies by its small size and by

the male palp with a blunt, dorsally bent RTA, a relatively large and broad prolaterally inserted embolus,

a small selerotised apieal eonduetor and a small and subtriangular retrolaterally inserted MA.

Etymology

The speeies name is derived from the Eatin nanus, dwarf, and refers to the small size of the present

speeies.

Type material

Holotype

(?: TANZANIA, E. Usambara Mts., Amani, 5°5.7’S 38°38’E, 28 Oet.-9 Nov. 1995, 950 m asl, pitfalls,

Griswold C., Seharff N. & Ubiek D. (CAS)

Paratype

1 S', together with holotype.

Description

Male (holotype)

Totae eength. 1.74. Carapaee length 0.74, width 0.58, unieoloured pale yellow, unbordered (Fig. 3C).

Fovea pronouneed, length 0.08, anterior end 0.53 from front end of earapaee. MOQ length (when eight

eyes present) 0.05, anterior width 0.02, posterior width 0.06. AER width 0.11, PER width 0.15. Median

eyes with a strong tendeney towards reduetion, one speeimen with redueed median eyes, the other

having only four eyes (Fig. 12J). AME very small (1/6 of AEE diameter) or absent, PME very pale and

strongly redueed (1/8 of AEE diameter) or absent, PEE 2/3 of AEE diameter. Both eye rows (if ME are

26

r

BOSSELAERS J. & JOCQEIE R., Anew afrotropical genus of Cybaeodinae (Araneae, Eioeranidae)

present) straight from above. Clypeus vertieal, equal to 1/3 of AEE diameter. Chilum indistinet. Sternum

yellow, unbordered (Fig. 3D), length 0.47, width 0.39. PCX indistinet.

Abdomen. Pale eream dorsally (Fig. 3F) with traees of a weak, diffuse anterior do seutum and ventrally

with an oblong, selerotised pateh earrying strong modified setae in posterior half (Fig. 3 E). Spinnerets

as for the genus in general. Fegs pale yellow (Fig. 3D, F). Retroeoxal hymen pronouneed, oval and

white. Troehanter noteh indistinet in legs I and II, pronouneed in legs III and IV. Very sparse ve terminal

preening brushes on mt III and IV. Tarsus IV slightly bent. Feg spination as in Appendix IB.

Feg measurements:

fe

pa

ti

mt

ta

Total

I

0.53

0.24

0.39

0.29

0.28

1.72

II

0.45

0.22

0.37

0.28

1.59

III

0.42

0.14

0.29

0.32

0.29

1.46

IV

0.55

0.21

0.47

0.50

0.39

2.13

Maee paep. With a blunt, dorsally bent RTA, a relatively large and broad prolaterally inserted pointed

embolus, a small selerotised apieal eonduetor, and a pointed, small and subtriangular retrolaterally

inserted MA.

Female

Unknown.

Distribution

Tanzania, East Usambara mountains, 950 m asl.

Discussion

Phylogenetic analyses

The preferred eonsensus tree for 27 speeies known from both sexes, with node numbers, state ehanges

for 47 eharaeters and Goloboff fit Bremer support values (as reported in TNT) in italies below branehes

is illustrated in Fig. 1. Eaeh ambiguity on the tree was optimized in isolation, in order to avoid

seoring eharaeter states for absent struetures, and also beeause only a eombination of ACCTRAN and

DEFTRAN optimisation ean produee the most robust proposal for a supposed homology. Indeed, the use

of ACCTRAN only, as is often preferred, does not always maximize parallel loss of eomplex traits over

eonvergent gains (Agnarsson & Miller 2008). Of the five ambiguous eharaeters shown on the preferred

tree in Fig. 1, DEFTRAN optimisation was preferred for eharaeters 19, 38 and 72, ACCTRAN for

eharaeter 37, and only unambiguous ehanges are shown for eharaeter 22.

Homoplasy in the data matrix whieh produeed the preferred tree is quite aeeeptable: 22 out of 99

eharaeters are eompletely free of homoplasy. Sanderson & Donoghue (1989: 1785, fig. 1) performed a

polynomial regression analysis on data from 60 eladistie analyses, and derived the following equation

based on them: ei = 0.90 - 0.022x(number of taxa) + 0.000213^(number of taxa)^. Applying this equation,

27 taxa would yield a ei value of 0.461, quite similar to the aetual ei value of 0.412 obtained for the

eonsensus tree in the present analysis.

The ingroup (elade 1) is supported on the preferred tree by the absenee of apieal do spines on fe III

and IV (11:0, not shown in Fig. I, reversed in Cybaeodes marinae Di Franeo, 1989 and in elade 14),

the presenee of a distal spine on the male palpal pi edge [27:1, absent in Scotina palliardii (F. Koeh,

27

European Journal of Taxonomy 1-49 ( 2013 )

Fig. 9. Cteniogaster toxarchus gen. et sp. nov. A. S, habitus, dorsal view. B. Idem, abdomen, ventral view.

C. Idem, palp, ventral view. D. Idem, retrolateral view. E. Epigyne, ventral view. F. $, earapaee, lateral view.

G. S, earapaee, lateral view. H. $, earapaee, dorsal view. Seale bars: A-C, F-H = 0.5 mm, C-E = 0.25 mm.

28

r

BOSSELAERS J. & JOCQEIE R., Anew afrotropical genus of Cybaeodinae (Araneae, Eioeranidae)

1881)], absenee of true elaw tufts (36:0), the presenee of tenent hairs at the tip of tarsi (37:1, reversed

in elades 3 and 14, but present in Apostenus), a simple sternal border (42:0, reversed in Toxoniella), two

retromarginal ehelieeral teeth (43:0, more than two in Toxoniella), a shaggy hair in front of the fang base

(44:1, reversed in Toxoniella), a eonspieuous serrula on the endites (48:0), a flat earapaee (50:1, ehanged

to slanting in elade 20), modifled PME (57:1, reversed in elade 14), the MOQ widest posteriorly (58:1),

eurved strong hairs frontally on abdomen (60:1), no epigastrie selerite (62:0, 70:0), laterally eompressed

female PMS (71:1, ehanged to slender in elade 4 and to stout and subtriangular in elade 11), absenee of

a eoiled sperm duet (79:0), a membranous eonduetor (81:1, reversed in Cteniogaster conviva sp. nov.

md Agraecina lineata (Simon, 1878), unapplieable in elade 20), a simple eonduetor (82:0, eomplex in

C. conviva sp. nov., unapplieable in elade 20), MA present (83:1), subtegulum pi (89:0), anterior hood

Fig. 10. Cteniogaster hexomma sp. nov. A. cJ, habitus, dorsal view. B. Idem, abdomen, ventral view.

C. Idem, palp, ventral view. D. Idem, retrolateral view. E. Epigyne, ventral view. F. earapaee, anterior

view. Seale bars: A-B, F = 0.5 mm, C-E = 0.25 mm.

29

European Journal of Taxonomy 1-49 ( 2013 )

present in epigyne (94:1, reversed in Hesperocranum rothi Ubiek & Platniek, 1991, and in elade 15) and

anterior epigynal entranees (97:0). We eonsider elade 1 to represent the family Lioeranidae. It is divided

in two sister elades in the present analysis: elade 2 and elade 7 (Fig. 1).

Clade 2 is interpreted as subfamily Lioeraninae and is supported on the preferred tree by the presenee of

large ereetile bristles in the ve seopulae of legs 1 and 11 (5:1), ventral seopulae on ti 1 and 11 (33:1) and a

pi terminal lobe on the male palpal ti (91:1). Clade 4 (Mesiotelus Simon, 1897 and Liocranum L. Koeh,

1866) is further eharaeterised by slender female PMS earrying only a single large spigot (71:0, 72:0,

contra Wunderlieh 2008: 489).

Clade 7 is interpreted as subfamily Cybaeodinae and is supported on the preferred tree by the presenee

of a troehanter noteh (3:1, reversed in Apostenus), the presenee of bent male tarsi (8:1), presenee of

do spines on ti 111 (at least in females) and IV (17:1, 18:1), eylindrieal ALS in males (65:1, reversed

in elade 17), and presenee of EPOS in males (69:1, reversed in elade 20). Although present in some

Clubionidae as well (see diseussion of eharaeter 69, above), the presenee of EPOS seems to be an

interesting apomorphy for the subfamily Cybaeodinae. The eharaeter is lost in the gQnQY3. Apostenus and

Scotina, most probably due to their small size. Cybaeodes Simon, 1878 (elade 8) holds a basal position

within Cybaeodinae in the present analysis, supported by two rows of more than five large spigots

on female PMS (paralleled in Toxoniella) and a small dorsal bristle mat on the male palpal eymbium

(76:1). Toxoniella (elade 10) differs from all other members of elade 1 by a number of reversals,

being a rebordered sternum (42:1), more than two retromarginal ehelieeral teeth (43:1) and absenee

of a shaggy hair in front of the ehelieeral fangs (44:0), but the genus fits within the elade for all other

important eharaeters, ineluding the presenee of an epigynal anterior hood (94:1). Toxoniella shares with

Cybaeodes, Hesperocranum Ubiek & Platniek, 1991 and Sagana Thorell, 1875 the laterally eompressed

female PMS (71:1), and with Cybaeodes and Sagana a large number of tarsal tenent hairs (38:2). Both

eharaeters appear plesiomorphie within the family. Within Cybaeodinae, Toxoniella shares the presenee

of EPOS (69:1) with most other genera. Toxoniella is herewith transferred to Eioeranidae. None of the

other genera eurrently ineluded in Gallieniellidae is reported to possess EPOS in males.

Cteniogaster gen. nov. also fits well within Cybaeodinae, but differs from related genera by the wide

patellar indentations (6:1, 7:1, paralleled in Apostenus), the presenee of one mt IV plv and rlv spine

(21:1), two pairs of tenent hairs (38:1), and the presenee of ve abdominal setae in males (63:1, paralleled

mAgroeca brunnea (Blaekwall, 1833) dxid Agrocea cuprea Menge, 1873 (elade 19) and in Apostenus

fuscusWQsixing, 1851). The faet that these male ve abdominal setae oeeupy a well delimited small oval

area (64:1) is an autapomorphy for the genus. Clade 14 is supported by the absenee of mt 111 and IV vt

spines (23:0, reversed in elade 20) and the presenee of eireular PME (57:0). Clade 16 is supported by the

presenee of a subtegular looking lobe (78:1), eomplemented in elade 17 (the gQnQX?i Agroeca, Apostenus

?indScotina) by a tegular looking lobe (77:1) and eoniealAES in males (65:0). The gQnxxsAgroeca (elade

18) is supported by a bifid MA (84:1, paralleled in Sagana rutilans Thorell, 1875) and a fiat, ribbon¬

shaped embolus (87:4). Clade 20, eonsisting of small, derived Cybaeodinae, is distinguished by plv

spines on fe 1 (9:1), a slanting earapaee (50:0), and absenee of a eonduetor (80:0) and of EPOS (69:0).

Apostenus is further eharaeterised by the presenee of one pair of tenent hairs (38:0), widely separated

PME (59:2) and the absenee of ST2 (98:0). Its sister genus Scotina features equidistant PE (59:1) and a

whip-shaped embolus (87:5).

While some eharaeters (36:0,48:0, 58:1, 60:1, 62:0, 70:0, 79:0, 83:1, 89:0, 97:0) are eonstant throughout

Eioeranidae, others (44:1, 57:1, 65:1, 69:1, 94:1) show reversal in some elades.

Reversals and seeondary losses of eharaeters are partieularly eommon on the preferred tree in the

genus Toxoniella and in elade 20, whieh groups the genera Apostenus and Scotina. Apart from the three

30

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BOSSELAERS J. & JOCQEIE R., Anew afrotropical genus of Cybaeodinae (Araneae, Eioeranidae)

reversals mentioned above, Toxoniella has no plv and rlv spines on female ti I (14:0) and has male and

female PES elose together (68:0 and 73:0, paralleled in elade 3 and in elade 4, respeetively). In elade

20, apart from the already mentioned absenee of EPGS (69:0) and a eonduetor (80:0), the number of

mt III plv and rlv spines is redueed (20:0, paralleled in Cteniogaster gen. nov. and Hesperocranum), mt

Fig. 11. A. Cteniogaster toxarchus gen. et sp. nov. Tip of male palp, ventral view. B-D. Cteniogaster

hexomma sp. nov. B. Tip of male palp, ventral view. C. Tip of male tarsus IV, retrolateral view. D. Bent

male tarsus IV. E. Cteniogaster toxarchus gen. et sp. nov., tip of male tarsus IV. F. Apostenus spinimanus

(Koeh & Berendt, 1854), tip of male tarsus IV. G. Apostenus fuscus Westring, 1851, tip of male tarsus IV.

H. Arabeliapheidoleicomes Bosselaers, 2009, tip of female tarsus IV. Seale bars: D = 200 pm; A-C, E-G =

100 pm. Abbreviations: C = eonduetor; E = embolus; MA = median apophysis.

31

European Journal of Taxonomy 1-49 ( 2013 )

Fig. 12. A-E. Cteniogaster conviva sp. nov. A. S, palp, ventral view. B. Idem, retrolateral view.

C. Epigyne, ventral view. D. Idem, other speeimen. E. Eye region of male, anterior view. E-G.

Cteniogaster sangarawe sp. nov. E. Epigyne, ventral view. G. Idem, other speeimen. H-J. Cteniogaster

nana sp. nov. H. S, palp, ventral view. I. Idem, retrolateral view. J. Eye region of male, anterior view.

K. Cteniogaster taxorchis sp. nov. Epigyne, ventral view. L. Cteniogaster lampropus sp. nov. Epigyne,

ventral view. Seale bar = 250 pm.

32

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BOSSELAERS J. & JOCQEIE R., Anew afrotropical genus of Cybaeodinae (Araneae, Eioeranidae)

III and IV have ve terminal spines (23:1), feathery hairs are absent (30:0, paralleled in Hesperocranum,

Liocranoeca and Agraecina), there is no apieal maxillar hair tuft (49:0, paralleled in Cteniogaster

gen. nov., Hesperocranum and Liocranoeca), and PME and PEE are elose to eaeh other (59:1 or 59:2).

Moreover, Apostenus has no troehanter noteh (3:0) and no ST2 (98:0). The presenee of male modified

ve abdominal setae (63:1) appears to have evolved eonvergently in Cteniogaster gen. nov. and in some

Agroeca and Apostenus (Wunderlieh 2008: 489). The eharaeter was already present in the eoeene

Apostenus spinimanus. The presenee of tenent hairs (37:1) also shows a peeuliar distribution on the

preferred tree, being restrieted to the genera Sagana, Cybaeodes, Toxoniella, Cteniogaster gen. nov., and

Apostenus. While Sagana has 9 and Cybaeodes (see Bosselaers 2009) and Toxoniella 5 pairs of tenent

hairs, Cteniogaster gen. nov. has only two pairs (Fig. IIC-E) ein&XQCQnt Apostenus only one (Fig. IIG).

However, the extinet Apostenus spinimanus, most probably eongenerie with reeent speeies, has three

pairs (Fig. HE, contra Wunderlieh 2004: 1627). Based on the presenee of tenent hairs, modified male

ve abdominal setae, an epigynal anterior hood and the other eharaeters listed above for elade 1 (with

the exeeption of a selerotised eonduetor for Cteniogaster conviva sp. nov.), the new genus Cteniogaster

gen. nov. is plaeed in Eioeranidae, Cybaeodinae.

The present phylogenetie analysis does not produee an unequivoeal autapomorphy for Eioeranidae.

However, a eombination of a number of non-homoplasious eharaeter ehanges mentioned above for

elade 1 in the diseussion of phylogenetie findings, offers signifieant potential for reeognising genera

as Eioeranidae. No doubt, most of these eharaeter states (11:0, 27:1, 42:0, 43:0, 44:1, 48:0, 57:1,

58:1, 60:1, 89:0, 97:0) are not unique to Eioeranidae and probably plesiomorphie on a wide seale or

only apomorphie to a larger elade in whieh Eioeranidae will prove to be embedded. Nevertheless, we

eonsider a eombination of a sizable number of these eharaeter states as the best approaeh available to

date towards reeognising Eioeranidae: the absenee of true elaw tufts (36:0), the presenee of tenent hairs

(37:1), the absenee of extensive abdominal ve selerotisation, at least in females (70:0), the presenee of

a simple, membranous eonduetor as well as a MA in the male palp (81:1, 82:0, 83:1) and an epigyne

with an anterior hood (94:1). The attribution of the lioeranid genera ineluded in the present analysis

to subfamilies Eioeraninae and Cybaeodinae ean be eonsidered robust, as both supraspeeifie taxa are

supported by apomorphies within Eioeranidae (5:1, 91:1 and 3:1, 8:1, 17:1, 18:1, 69:1, respeetively).

The presenee of EPOS in males (69:1) is the main eharaeter on whieh the transfer of Toxoniella to

Eioeranidae, Cybaeodinae is based. The family Gallieniellidae is poorly defined: porreet ehelieerae, the

only synapomorphy found in Platniek’s eladogram (2002: 9), also oeeur in other araneomorph spider

families, sueh as Clubionidae and Theridiidae Sundevall, 1833. In addition, Haddad et al (2009: 16)

mention small male AME, a reeurved PER, eonieal AES and a short eymbium tip, but most of these

eharaeters reverse somewhere within the Gallieniellidae elade and none is unique to the family. The

presenee of tenent hairs (37:1), a eharaeter found in the majority of Eioeranidae, is also eneountered

in the gallieniellid genera Drassodella Hewitt, 1916 (Wami & Joeque 2002: 314) and Austrachelas

Eawrenee, 1938, although in the latter they are only present on the posterior two pairs of legs, similar

to what is deseribed in Raven & Stumkat (2002) for some Clubionidae. However, no other gallieniellid

has eylindrieal AES provided with EPGS in males, and this eharaeter, in eombination with the above

mentioned set of eharaeters eommonly eneountered in Eioeranidae, is judged suffieient to justify the

transfer.

As far as the other lioeranid genera listed in Platniek (2012) are eoneemed, some eonelusions ean be

drawn and a few transfers proposed. Argistes Simon, 1897 and Sphingius Thorell, 1890 most probably

belong in Eioeranidae, given the presenee of tenent hairs (Deeleman-Reinhold 2001: figs. 639; personal

observation), a simple eonduetor and a MA. As long as the eonspeeifieity of the male deseribed by

Bosmans (2011: 20, figs. 15-16) with females of Arabella Bosselaers, 2009 is not proved beyond doubt,

it seems better to keep the genus as Eioeranidae incertae sedis, due to the presenee of five pairs of tenent

hairs and an anterior epigynal hood, as well as the eomplete absenee of abdominal selerotisation. Pending

33

European Journal of Taxonomy 1-49 ( 2013 )

a generic revision, the same classification is defended for Rhaeboctesis Simon, 1897, given the absence

of true claw tufts and abdominal sclerotisation, as well as the presence of a simple conductor and a MA.

Andromma Simon, 1893 does not fit well in Liocranidae as defined here, due to the presence of true claw

tufts and the absence of a MA and anterior epigynal hood. It is likely to belong in Corinnidae, but since a

comprehensive cladistic analysis of a larger number of liocranid and corinnid genera is not yet available,

it is best to keep it in Liocranidae as incertae sedis. Paratus Simon, 1898, which lacks a MA (Marusik

et al. 2008; Zapata & Ramirez 2010) was placed in a subfamily of its own by Marusik et al, based,

apart from the general character states already mentioned above for clade 1, on an insuffient number of

characters (2008: 51), such as absence of a retrocoxal hymen (also absent in Cybaeodes, Liocranoeca,

Neoanagraphis and Sagana, see Appendix 3), embolus inserted centrally on tegulum, very simple

epigyne and abdomen with guanine spots. It seems best to keep the genus as Liocranidae incertae sedis

until a thorough analysis has been performed. Sudharmia Deeleman-Reinhold, 2001, with its almost

unsclerotised female abdomen and male palp with pi subtegulum and simple, membranous conductor

(but lacking MA) is also considered Liocranidae incertae sedis here. Literature data on Heterochemnis

F.O.R Cambridge, \9^^,LaudetiaGorisch, \9A\,Liparochrysis^imon, 1909 andSimon, 1897,

Fig. 13. Map showing all localities of georeferenced Liocranidae in the collections of MRAC:

Cteniogaster gen. nov. (•), all other Liocranidae (o) (n = 315).

34

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BOSSELAERS J. & JOCQEIE R., Anew afrotropical genus of Cybaeodinae (Araneae, Eioeranidae)

Fig. 14. Distribution maps. A. Cteniogaster hexomma sp. nov. (•), C. toxarchus gen. et sp. nov. (*).

B. Cteniogaster lampropus sp. nov. (A), C. nana sp. nov. (o). C. Cteniogaster taxorchis sp. nov. (□), C.

conviva sp. nov. (■), C. sangarawe sp. nov. (A).

35

European Journal of Taxonomy 1-49 ( 2013 )

genera that have never been thorougly diagnosed or revised, are insuffieient to judge on their affinities:

these four genera are best kept in Lioeranidae incertae sedis for the time being. Sesieutes Simon, 1897,

will be transferred to Corinnidae, Phrurolithinae Simon, 1903 by Dankittipakul & Deeleman-Reinhold

(in press), and we propose the same transfer for the genera Jacaena Thorell, 1897, Plynnon Deeleman-

Reinhold, 2001 and Teutamus Thorell, 1890, based on their infiated tegulum, absenee of MA, modified

male palpal fe, simple epigyne without anterior hood and abdominal selerotisation (Deeleman-Reinhold

2001). A eomparison of Simon’s speeimens of Prochora lycosiformis (O.P.-Cambridge, 1872) with the

published illustrations of the male and female eopulatory organs of Itatsina praticola (Rosenberg &

Strand, 1906) elearly shows that the monospeeifie genus Itatsina Kishida, 1930 is eongenerie with the

equally monospeeifie Prochora Simon, 1886, although both speeies are not identieal. Consequently,

Prochora praticola eomb. nov. is transferred to Miturgidae Simon, 1885 here, beeause of its bifid RTA,

eombined with a eymbium with a rl groove lined with a fringe of setae and a basally inserted embolus

eneireling the tegulum (Song et al. 1999: fig. 238J-L). Coryssiphus Simon, 1903 and Donuea Strand,

1932 will be transferred to other families in fortheoming publieations. The holotype of Montebello tenuis

Hogg, 1914 was studied by Ovtsharenko (personal eommunieation) and turned out to be a juvenile,

damaged speeimen with eyes and spinnerets reminiseent of Gnaphosidae Poeoek, 1898. The genus is

transferred to Gnaphosidae incertae sedis here.

Wunderlieh’s (2011: 108) proposal to inelude Lioeranidae in Zoridae O.P.-Cambridge, 1893 and transfer

Cybaeodes to Gnaphosidae is rejeeted beeause it is based on insuffieient data and is not supported by a

eladistie analysis. Moreover, the four zorid speeies studied by the authors, Zora spinimana (Sundevall,

1833), Tuxoctenus gloverae Raven, 200H, Argoctenus sp. andHestimodema sp., apart from laeking tenent

hairs and male EPGS, all share a set of eharaeters not eneountered in Lioeranidae: a PER with large eyes

in two rows, as in Ctenidae and Eyeosidae Sundevall, 1833, PME elose together, thiek elaw tufts, an

anteriorly strongly narrowed earapaee, a male palpal eymbial tip with modified thiek setae as deseribed

in Raven (2008: 352), a RTA with a fiattened basal membranous wing, an epigyne without anterior hood

and a vulva without ST2 and with posterior globular STl eonneeted to tortuous insemination duets.

The proposals formulated here limit Eioeranidae to 25 genera, ineluding Cteniogaster gen. nov.,

Toxoniella and the reeently deseribed genus Vankeeria Bosselaers, 2012, of whieh the latter ean be

eonsidered incertae sedis (Bosselaers 2012). Of these genera, four belong in Eioeraninae, nine ean be

attributed to Cybaeodinae, and twelve remain incertae sedis, stressing the need for additional revisions

and a more thourough analysis of Eioeranidae and related dionyehan ground spiders.

Acknowledgements

The authors are grateful to the museum eurators and individual eolleetors mentioned above for the loan

of speeimens. Thanks are further due to Elise-Anne Eeguin (MNHN), Julia Altmann (SMF) and Timo

Pajunen (MZHF) for help in loeating and sending speeimens from their respeetive eolleetions, and to

Pablo Goloboff for explaining differenee in fit value between programmes and for suggesting a way to

ealeulate implied weighting Bremer support in TNT. The authors are also grateful to Alexandre Bonaldo,

Vladimir Ovtsharenko, Norman Platniek, Martin Ramirez, Charles Haddad and Jorg Wunderlieh for

interesting diseussions on Eioeranidae and allied genera, the latter two persons also having refereed

the paper. Speeial thanks go to Darell Ubiek for initially drawing the first author’s attention to the

existenee of this new genus by sending four speeimens of Cteniogaster toxarchus sp. nov. from the CAS

eolleetion in 1998, and to Robert Raven for fruitful diseussions and for the generous gift of speeimens

of Australian Zoridae and Miturgidae.

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BOSSELAERS J. & JOCQEIE R., Anew afrotropical genus of Cybaeodinae (Araneae, Eioeranidae)

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1781-1795. http://dx.doi.org/10.2307/2409392

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Sehulze zur Wiesehe E. 2006. Modeme Haarpflegemittel. Praxis der Naturwissenschaften - Chemie in

der Schule 6 (55): 14-17.

Sierwald P. 1990. Morphology and homologous features in the male palpal organ in Pisauridae and other

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39

European Journal of Taxonomy 1-49 ( 2013 )

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Manuscript received: 25 October 2012

Manuscript accepted: 21 January 2012

Published on: 11 March 2012

Topic editor: Koen Martens

Desk editor: Charlotte Thionois

Printed versions of all papers are also 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 Alfiea, Tervuren, Belgium; Natural History

Museum, London, United Kingdom; Royal Belgian Institute of Natural Seienees, Brussels, Belgium;

Natural History Museum of Denmark, Copenhagen, Denmark.

40

BOSSELAERS J. & JOCQEIE R., Anew afrotropical genus of Cybaeodinae (Araneae, Eioeranidae)

Appendices

Appendix 1. Eeg spination sehematies. A. Eegend. B. Cteniogaster nana sp. nov., S- C, F. Cteniogaster

toxarchus gen. et sp. nov. D, G. Cteniogaster hexomma sp. nov. E, H. Cteniogaster conviva sp. nov.

C-E. 33- F-H. I. Cteniogaster lampropus sp. nov., $. J. Cteniogaster taxorchis sp. nov., $. K.

Cteniogaster sangarawe sp. nov., $. White dots are weak spines or spines that may be present or absent.

A

Palp

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.

I

II

III

IV

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41

European Journal of Taxonomy 1-49 ( 2013 )

Appendix 2. Taxa examined to provide exemplar data and to allow eomparison with related families,

with eolleetion data and deposition. Asterisk indieates type speeies of genus. Families in whieh taxa

are plaeed aeeording to prevailing views are added between square braekets (Ga = Gallieniellidae,

Li = Lioeranidae, Co = Corinnidae, Zo = Zoridae).

Outgroup

Corinna nitens (Keyserling, 1891) [Co]; 2 1 Brazil, Rio Grande do Sul, Novo Hamburgo, 27

Feb. 1979, T. de Lema leg. (MCN 8558); 1 f: Rio Grande do Sul, Sao Leopoldo, 27 Jun. 1987, L. de A.

Moura leg. (MCN 16884); 1 Rio Grande do Sul, Sao Franeiseo de Paula, Barragem dos Bugres, 25

Nov. 1998, L. de A. Moura leg. (MCN 30653).

Creugas bajulus (Gertseh, 1942) [Co]; 1 S'. USA, California, San Diego County, Miramar Naval Air

Station, eoastal sage serub, pitfall trap. May-Jun. 1996, T. Prentiee leg. (UCR); 1 same data, Aug.

1996 (UCR) (Prentiee et al. 1998).

Ingroup

Agraecina lineata"^ (Simon, 1878) [Li]; 5 SS'- Franee, Corsiea and Menton (MNHN 2095); 1 Franee,

Les Saintes Maries, “Colleetion Berland, entree n° 4.25.9.62” (MNHN).

Agroeca brunnea'^ (Blaekwall, 1833) [Li]; 1 S'- Belgium, Beerse, “De Sehrieken”, pitfall trap in pine

wood with braeken, 20 Feb. 1995, J. Bosselaers leg. (CJB 1003); 1 same loeality, pitfall trap in alder

marsh, 2 Dee. 1994, J. Bosselaers leg. (CJB 882).

Agroeca cuprea [Li]; 1 S'- Belgium, Hautes Fagnes, no further data (CHK); 1 Franee, Departement

Var, between Pierrefeu and Collobrieres, under wood and stones in open Quereus suber and Pinus forest,

43°13’53.4”N6°14’L6”E, alt. 100 m. 17 Mar. 2006, A Sehonhofer/eg. (SMF).

Agroecaparva Bosmans, 2011 [Li]; 1 S'- Greeee, Chios, Mount Pelinion, pitfall trap, 26 Sep.-10 Get.

2009 (CARS); 1 Greeee, Chios, Padoukios, pitfall trap, 6-22 May 2009 (CARS).

Agroecaproxima (O.P-Cambridge, 1871) [Li]; 2 SS'- Belgium, Beerse, “St. Jozef’, heathland, pitfall

trap, 4 Get. 1985 (CJB 472a); 1 Belgium, Beerse, “De Sehrieken”, Calluna heathland, pitfall trap, 20

Nov. 1994, J. Bosselaers leg., (CJB 837b)

Apostenusfuscus^ Westring, 1851 [Li]; 3 SS, 1 ?: Yugoslavia, Slavonski Pozega, among oak litter, 26

Mar. 1972, C. Deeleman-Reinhold leg. (RMNH).

Apostenus humilis Simon, 1932 [Li]; 1 S'- no further data (MNHN25305); 21 ""Apostenus humilis

E.S. in litteris et det. Banyuls 09” (MNHN25).

Cteniogaster conviva sp. nov.; 1 1 ? (holotypus and paratypus): Tanzania, Tanga, West Usambara

Mountains, Mazumbai Forest, sifting litter, 4°49’S 38°30’E, alt. 1400-1800 m. 11-20 Nov. 1998, Charles

Griswold, Nikolaj Seharff and Darrell Ubiek leg. (ZMUC).

Cteniogaster hexomma sp. nov.; 1 (5, 1 $ (holotypus and paratypus): Kenya, Taita Hills, Ngangao

forest, mixed forest, pitfall trap, 03°22’S 038°2UE, alt. 1820 m. 15 Jul. 1998, Euey Rogo leg. (MRAC

208888).

Cteniogaster toxarchus'^ sp. nov.; 17 SS, 25 $$ (paratypi): Tanzania, East Usambara Mountains,

Amani, sifting litter, 5°5.7’S 38°38’E, alt. 950 m. 28 Get.- 9 Nov. 1995, Charles Griswold, Nikolaj

Seharff and Darrell Ubiek leg. (ZMUC).

42

r

BOSSELAERS J. & JOCQEIE R., Anew afrotropical genus of Cybaeodinae (Araneae, Eioeranidae)

Cybaeodes mallorcensis Wunderlieh, 2008 [Ei]; 1 S (holotypus): Spain, Mallorea, South of Embalse

de Cuber, under stone, Aug., J. Wunderlieh leg. (SMF 60217); 1 $ (paratypus): Spain, Mallorea,

Valldemossa, JuL, J. Wunderlieh leg. (SMF 60218).

Cybaeodes marinae Di Franeo, 1989 [Ei]; 1 S'- Italy, Sieily, Nebrodi, E. slope of Monte Pagano, 550 m,

mixed shrub with Quereus suber and Q. gussonei, Sep. 1988 (AMNH); 1 (?, 1 Italy, Fazio, Rome,

Castelporziano, oakwood, 13 Oet. 1986, Bonavita leg. (AMNH).

Hesperocranum rothC Ubiek & Platniek, 1991 [Ei]; 1 S'- USA, California, Tulare County, 2.5 mi E of

Cal. Hot Springs, 15 Sep. 1959, V. Roth & W. Gertseh leg. (AMNH); 1 California, Fresno County,

Kings Cyn. Nat. Park, Cedar Grove, 13 Sep. 1959, V. Roth & W. Gertseh leg. (AMNH).

Liocranoeca striata^ (Kulezynski, 1882) [Ei]; 1 S'- Franee, Southern Franee, eoastal area (MNHN

6147); 1 Franee, Corsiea and Menton (MNHN 2095).

Liocranum giersbergi Kraus, 1955 [Ei]; 1 (J, 2 Italy, Sardinia, Villanova Strisaili, pitfall trap at

lakeside in wood, 23 May 1997, J. van Keer leg. (CJK 1757); 1 $ (holotypus): Italy, Sardinia, Bunorva,

alt. 500 m. 15 Apr. 1952, K. Sehtillbaeher leg. (SMF 8971/1).

Liocranum rupicola'^ (Walekenaer, 1830) [Ei]; 1 (J, 1 ?: Italy, Fazio, Eatina, street to summit of the

Monte Cireea, oak forest, sieving litter under wood and stones. 41°13‘58.3“N 13°04‘59.6“E, alt. 229 m.

9 Apr. 2006, A. Sehonhofer leg. (SMF “13Id”).

Mesiotelus cypriusKulezynski, 1908 [Ei]; 1 (5^, 8 $ Cyprus, Troodos Mountain, 1600 m, under stones

in pine forest, 28 Oet. 1981, PR. Deeleman leg. (RMNH); 7 $ $: Crete, Zoniana, outside Sventoni Cave,

700 m, under stones, 6 Apr. 1996, J. Bosselaers leg. (CJB 1047).

Mesiotelus tenuissimus^ (E. Koeh, 1866) [Ei]; 2 SS'- Greeee, Naxos, 16 Oet. 1984, Parashi leg. (SMF);

1 $#f Syria, Homs provinee, QaE at al-Husn, ruins and abandoned eemetery NW of the eastle. 8 Mar.

1979, R. Kinzelbaeh leg. (SMF).

Neoanagraphis chamberlinC Gertseh & Mulaik, 1936 [Ei]; 2 SS'- USA, NewMexieo, Soeorro County,

Sevilleta 20 mi N Soeorro, 1500-2200 m, 1989-1992, S Brantley leg. (UCR, now in CJB); 1 same

data (UCR).

Saganarutilans'^ TkoxeW, 1875 [Ei]; 1 (5': Bosnia and Herzegovina, Prisoje area, Eivno near Tomisiovgrad,

Camp Graboviea, eolleeted at night under stones at small stream, 43.69512°N 17.09543°E, alt. 875 m.

3-5 Sep. 2009, A. Sehonhofer leg. (SMF); 5 ? “Gallia” (MNHN 2098).

Scotina celans (Blaekwall, 1841) [Ei]; 1 (?, 4 Franee, Banyuls (MNHN); 2 Franee, Saint

Gilles (MNHN); 1 Yugoslavia, Island Cres, Jama, Eipiei Cave, 1 Apr. 1972, C.E. & PR Deeleman

leg. (RMNH).

Scotina gracilipes'^ (Blaekwall, 1859) [Ei]; 1 (?, 2 Denmark, Anholt, plantation, 11 Aug. 1946, 0.

Christensen leg. (ZMUC 14); 2 (Jc?: Belgium, Kalmthout, willow serub and Calluna heath, 15 and 29

Nov. 1976, P. Baert leg. (CJK 862).

Scotinapalliardii (E. Koeh, 1881) [Ei]; 5 SS, 2 $ $: Finland, North Karelia, Elomantsi, Heinasuo, 26E,

5 Jun. 2007, O. Eoukola and O. Autio leg. (MZHF 98274).

43

European Journal of Taxonomy 1-49 ( 2013 )

Toxoniella rogoae Warn! & Jocque, 2002 [Ga]; 1 f (holotypus): Kenya, Taita Hills, Ngangao forest,

mixed forest, pitfall trap, 03°22’S 38°2rE, alt. 1820 m. 15 Jul. 1998, Luey Rogo leg. (MRAC 209914);

1 $ (paratypus): same data (MRAC 209661).

Toxoniella taitensis"^ Wami & Joeque, 2002 [Ga]; 1 (J, 1 $ (paratypi): Kenya, Taita Hills, Ngangao

forest, 03°22’S 38°2rE, alt. 1820 m. 15 Jul. 1998, E. Rogo leg. (MRAC 208887).

Additional specimens studied

Argoctenus sp. [Zo]; 1 S'. Australia, SE Queensland, Eake Broadwater via Dalby, pitfall traps, 24 Nov.

1985-3 Jan. 1986, M. Bennie leg. (CJB).

Argoctenus sp. [Zo]; 1 Australia, SE Queensland, Eake Broadwater via Dalby, pitfall traps, 19 Feb.-

26 Mar. 1985, M. Bennie leg. (CJB).

Austrachelas natalensisEdiSNXQncQ, 1942 [Ga]; 1 (J, 2 South Afriea, Natal, St. Eueia, Fames Island,

woodland edge with grass, pitfall traps, 21-25 Feb. 1990, M. Alderweireldt & R. Joeque leg. (MRAC

171752).

Austrachelas pondoensis Haddad et al., 2009 [Ga]; 1 (J, 8 South Afriea, Eusikisiki Distriet, Transkei

Coast, Mzimhlava River mouth. Coastal evergreen Forest, 31°20’S 29°40’E, Feb. 1980, M. Baddeley

leg. (MRAC 163974).

Drassodella sp. [Ga] 1 S'- Mozambique, Inhaea, Coastal woodland, pitfalls, 26°0ES 32°54’E, 13-30

Apr. 1994, T. Steyn leg. (MRAC 209887).

Gallieniella jocquei Platniek, 1984 [Ga]; 1 (?, 1 Comoros, Moheli, Miringoni, plantations dTlang

Hang, 12°15’S 43°45’E, Mango and other trees at adge of plantation, pitfall traps, 5-13 Nov. 1983, R.

Joeque leg. (MRAC 160925).

Hestimodema sp. [Zo] 1 (J, 1 $: Australia, SE Queensland, Eake Broadwater via Dalby, pitfall traps, 17

May-24 Nov. 1985, M. Bennie leg. (CJB).

Tuxoctenus gloverae Raven, 2008 [Zo] 1 S'- Australia, SE Queensland, Burbank, Buhot Creek, riparian

forest, 27°35.5’S 153°10.3’E, 50 m, pitfall traps, 30 Jan.-l Mar. 2004 (CJB).

Tuxoctenus gloverae Raven, 2008 [Zo] 1 $: Australia, SE Queensland, summit of Mount Gayndah,

25°36’S 151°32’E, 340 m, pitfall traps, 27 Jan.-2 Jun. 1999, Montelth & Thompson leg. (CJB).

Zora spinimana'^ (Sundevall, 1833) [Zo] 3 SS, 3 $ $: Belgium, Zonhoven, Bolderberg, Gust Claesheide,

mixed forest with Calluna heath, pitfall traps, 27 Nov. 1983-21 Mar. 1984, J. Bosselaers leg. (CJB 0045).

44

BOSSELAERS J. & JOCQEIE R., Anew afrotropical genus of Cybaeodinae (Araneae, Eioeranidae)

Appendix 3. Charaeter-taxon matrix.

Golobofffit

0.33

00

o'

0.75

o

0.75

o

090

0.38

0.43

090

0.38

o

090

0.43

ri

o

m

o'

r--

o'

o

oo

o'

o

0 \

o'

o

00

o'

in

o'

o

o'

(N

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o'

o

in

o'

o

in

o'

o

o'

o

oo

o'

oo

m

o'

o

00

o'

m

o'

ci

o'

r--

o'

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m

o'

o

m

o'

o

in

o'

o

in

o'

o

m

o'

m

o

m

o'

00

m

o'

m

o'

o

in

o'

t--

o'

o

in

o'

m

o'

Steps

o

m

00

in

00

in

(N

in

m

in

(N

<o

00

Toxoniella taitensis

o

-

o

O-

o

-

o

O

o

-

o

o-

O-

-

(N

-

Toxoniella rogoae

o

-

o

o-

o

-

o

O

o

o-

o

-

o-

-

Scotina palliardii

-

o

o-

-

o

-

O-

-

o

m

(N

o

-

(N

o

Scotina gracilipes

-

o

o-

-

o

-

o

m

(N

o

-

(N

o

Scotina celans

-

o

o-

-

o

-

o

m

(N

O-

-

(N

o

Sagan a rutilans

o

-

o

m

(N

o

-

(N

Neoanagraphis chamberlini

o

-

o

-

o

-

o

-

(N

Mesiotelus tenuissimus

-

o

-

o

-

o

-

Mesiotelus cyprius

-

o

-

o

-

o

-

Liocranum rupicola

o

-

o

-

o

(N

o

-

O-

Liocranum giersbergi

o

-

o

-

o

O-

o

(N

o

-

(N

Liocranoeca striata

o

-

o

o-

o

-

o

-

(N

Hesperocranum rothi

-

o

-

o

O

o

O

Cybaeodes marinae

o

-

o

-

o

-

o

(N

-

Cybaeodes mallorcensis

o

-

o

-

o

(N

-

Cteniogaster toxarchus

-

o

O-

-

o

-

o

Cteniogaster hexomma

-

o

o-

-

o

-

o

Cteniogaster conviva

-

o

o-

-

o

-

o

Apostenus humilis

o

o-

-

o

-

o

(N

o

-

(N

o

Apostenus fuscus

-

o

o-

-

o

-

o

(N

o

-

(N

o

Agroeca proxima

-

o

o-

o

-

o

-

o

-

o

-

(N

Agroeca parva

o

-

o

o-

o

-

o

-

o

-

(N

Agroeca cuprea

-

o

o-

o

-

o

-

(N

Agroeca brunnea

-

o

o-

o

-

o

-

o

-

(N

Agraecina lineata

o

-

o

o-

o

-

o

-

o

-

(N

Creugas bajulus

o

-

o

-

o

(N

o

(N

-

Corinna nitens

-

o

-

o

-

o

-

(N

o-

-

o

(N

-

Characters

<s

«r5

'O

r-

00

0\

o

<s

in

'sO

r-

00

0\

O

<s

45

European Journal of Taxonomy 1-49 ( 2013 )

o

in

•n

m

o

in

o

m

•n

o

in

o

Golobofffit

m

in

r--

m

o

in

o

in

o

o'

t--

o

m

o

in

m

o

(N

o

(N

oo

o

r--

o

(N

ri

o

00

o

m

in

r--

in

<o

in

o

r--

o

00

r--

o'

o

m

(N

o

m

o

m

o

m

o

in

o

m

o

ci

m

in

(N

in

m

(N

in

m

o

in

m

o

(N

o

m

in

o'

Steps

(N

o

(N

m

0\

m

0\

m

in

o

m

o

(N

o

in

(N

0\

Toxoniella taitensis

(N

-

(N

-

o

-

o

-

(N

-

Toxoniella rogoae

(N

-

o

(N

-

o

-

o

-

(N

-

Scotina palliardii

(N

-

(N

o

-

o

o-

o

o-

O

o

Scotina gracilipes

(N

-

(N

-

o

o-

o

o-

o

Scotina celans

o-

(N

-

(N

-

o

o-

o

o-

o

Sagan a rutilans

-

m

-

(N

-

o

-

(N

-

Neoanagraphis chamberlini

cn

(N

o

-

m

-

(N

-

o

o-

o

-

o

o-

o

Mesiotelus tenuissimus

(N

-

o

m

-

o

(N

-

o

-

o

o-

-

o

Mesiotelus cyprius

(N

-

o

m

-

o

-

o

-

o

o-

-

o

Liocranum rupicola

o-

-

o

m

(N

-

o

(N

-

o

-

o

o-

-

o

Liocranum giersbergi

o-

-

o

m

(N

-

o

(N

-

o

-

o

o-

-

o

Liocranoeca striata

cn

(N

o

-

m

-

(N

o

o-

o

O-

o

Hesperocranum rothi

o

O

-

o

O

-

o

(N

o

o-

-

o

O-

-

Cybaeodes marinae

m

-

o

-

o

-

(N

-

Cybaeodes mallorcensis

cn

-

o

-

o

-

(N

-

Cteniogaster toxarchus

-

(N

-

(N

-

o

-

o

Cteniogaster hexomma

-

(N

-

(N

-

o

-

o

-

o

Cteniogaster conviva

-

(N

-

(N

O

-

o

-

o

-

o

Apostenus humilis

o-

O

-

(N

O

-

o

(N

o

O-

o

-

O

o

Apostenus fuscus

o-

-

m

(N

-

o

(N

o

o-

o

-

O

o

Agroeca proxima

cn

(N

o

-

m

o-

-

(N

-

o

o-

o

-

o

O-

-

Agroeca parva

cn

(N

o

-

m

(N

-

(N

-

o

o-

o

-

o

o-

o

Agroeca cuprea

cn

(N

o

-

m

(N

-

(N

-

o

o-

o

-

o

o-

o

Agroeca brunnea

cn

(N

o

-

m

-

(N

-

o

o-

o

-

o

o-

o

Agraecina lineata

cn

(N

o

-

m

-

(N

o

o-

o

o-

o

Creugas bajulus

(N

-

O

o

-

O

-

o

-

o

o-

-

(N

Corinna nitens

(N

O

-

(N

o

-

o

-

o

-

o

o-

o

(N

Characters

<s

tn

'O

r-

00

C\

o

<s

m

tn

'O

r-

00

C\

o

<S

<s

f<0

46

BOSSELAERS J. & JOCQEIE R., Anew afrotropical genus of Cybaeodinae (Araneae, Eioeranidae)

in

m

o

m

o

m

o

m

in

o

m

in

o

Golobofffit

o

in

o

in

m

r--

VO

m

r--

o

o'

o

r--

t--

o

in

o

in

o

(N

o

(N

o

ri

(N

o

in

o

oo

in

0 \

o

o'

o

m

in

o

m

in

r-

m

in

o

ci

m

in

(N

m

(N

o

(N

o

m

<N

VO

m

(N

in

o

o'

Steps

m

(N

m

o

m

(N

m

(N

in

(N

Toxoniella taitensis

-

o

-

o

-

(N

o

O

-

o

-

Toxoniella rogoae

-

o

-

o

-

(N

o

-

o

-

Scotina palliardii

o

-

o

o-

-

o

(N

o

O

o

-

o

-

Scotina gracilipes

o

-

o

o-

-

o

(N

o

-

o

-

Scotina celans

o

-

o

o-

-

o

(N

o

-

o

-

Sagan a rutilans

o

-

o

-

(N

o

(N

-

o

-

Neoanagraphis chamberlini

o

-

o

-

(N

o

-

o

-

o

-

Mesiotelus tenuissimus

o

-

o

o-

o

-

o

-

o

-

Mesiotelus cyprius

o

-

o

o-

o

-

o

-

o

-

Liocranum rupicola

o

-

o

-

o

-

o

-

Liocranum giersbergi

o

-

o

-

(N

o

(N

-

o

-

Liocranoeca striata

o

-

o

-

o

-

(N

o

O

o

-

o

-

o

-

Hesperocranum rothi

-

o

-

o

O-

o

-

O

o

-

o

-

o

-

Cybaeodes marinae

(N

o

-

o

-

(N

-

(N

o

-

O

-

o

-

Cybaeodes mallorcensis

-

o

-

o

-

(N

-

(N

o

O

-

o

-

Cteniogaster toxarchus

o

-

o

-

(N

o

-

o

-

Cteniogaster hexomma

o

-

o

-

(N

-

o-

o

o-

O-

-

O-

o-

-

Cteniogaster conviva

o

-

o

-

(N

-

o

(N

-

o

-

Apostenus humilis

o

-

o

o-

-

o

-

o

(N

o

-

O

o

-

(N

-

Apostenus fuscus

o

-

o

o-

-

o

(N

o

(N

o

-

o

-

(N

-

Agroeca proxima

-

o

-

o

-

(N

o

O

o

-

o

-

o

-

Agroeca parva

o

-

o

o-

-

o

-

(N

o

-

o

-

o

-

Agroeca cuprea

o

-

o

o-

-

o

-

(N

o

-

o

-

o

-

Agroeca brunnea

o

-

o

-

(N

o

-

o

-

o

-

Agraecina lineata

o

-

o

o-

o

-

O

-

o

-

o

-

o

-

Creugas bajulus

-

o

-

o

-

(N

o

-

o

(N

o

Corinna nitens

o

-

o

-

o

(N

o

(N

-

(N

o

Characters

<s

'O

00

0 \

o

<s

f<0

«r5

VO

r-

00

0 \

o

«r5

VO

47

European Journal of Taxonomy 1-49 ( 2013 )

o

in

o

in

o

•n

in

o

Golobofffit

VO

o

in

VO

O;

r--

VO

o

VO

o

o'

00

o

0\

o

t--

(N

o

00

VO

(N

o

ri

o

00

in

o

Ov

o

00

oo

00

o

o'

o

cn

o

in

o

m

o

r-

in

m

o

ci

cn

o

cn

o

m

<N

in

o

VO

m

o

o'

Steps

o

(N

VO

cn

o

t--

(N

t--

m

(N

o

00

(N

On

o

(N

in

Toxoniella taitensis

-

o

o-

-

o

O

o

-

o

-

m

o

-

O

O-

O

o

-

Toxoniella rogoae

-

o

o-

-

o

-

o

-

m

o

-

o

o-

O

o

-

Scotina palliardii

o

o-

o

-

o

(N

-

o

-

o

-

o

Scotina gracilipes

o

o-

o

-

o

(N

-

o

-

o

-

o

Scotina celans

o

o-

o

-

o

(N

-

o

-

o

-

o

Sagan a rutilans

o

o-

o

-

o

-

(N

-

o

O-

o

-

Neoanagraphis chamberlini

o

o-

-

o

-

o

(N

-

o

o-

o

-

Mesiotelus tenuissimus

o

o-

o

-

o

o-

o

-

Mesiotelus cyprius

o

o-

o

-

o

o-

o

-

Liocranum rupicola

o

o-

o

-

o

o-

o

-

Liocranum giersbergi

o

o-

o

-

o

o-

o

-

Liocranoeca striata

o

o-

-

o

(N

-

o

-

o

-

Hesperocranum rothi

o

o-

o

-

o

-

(N

o

o-

o

-

Cybaeodes marinae

-

o

o-

-

o

-

o

-

m

-

o

-

Cybaeodes mallorcensis

-

o

o-

-

o

-

o

-

m

-

o

-

Cteniogaster toxarchus

-

o

-

o

-

o

(N

-

o

o-

o

-

Cteniogaster hexomma

-

o

-

o

-

o

(N

-

o

o-

o

-

Cteniogaster conviva

-

o

-

o

-

o

(N

-

o

o-

o

-

Apostenus humilis

o

o-

o

-

o

(N

-

o

-

o

-

o

Apostenus fuscus

-

o

-

o

-

o

(N

-

o

-

o

-

o

Agroeca proxima

o

O-

o

-

o

(N

-

o

-

o

-

o

-

Agroeca parva

o

O-

o

-

o

(N

-

o

-

o

-

o

-

Agroeca cuprea

o

-

o

-

o

(N

-

o

-

o

-

o

-

Agroeca brunnea

o

-

o

-

o

(N

-

o

-

o

-

o

-

Agraecina lineata

o

O-

-

o

-

o

(N

-

o

O-

o

-

o

-

Creugas bajulus

-

o

O-

o

-

o

-

(N

-

o

o-

o

-

Corinna nitens

-

o

O-

o

-

o

-

(N

-

o

-

o

o-

o

-

Characters

<s

m

tn

'O

00

Cv

o

<s

m

tn

'O

r-

00

Cv

o

'O

r-

00

48

BOSSELAERS J. & JOCQEIE R., Anew afrotropical genus of Cybaeodinae (Araneae, Eioeranidae)

o

in

o

in

o

00

m

o

00

o

00

in

o

in

o

in

Golobofffit

VO

O;

in

m

VO

o

m

O;

m

r--

VO

•n

r--

o

o'

o

O

in

r--

in

o

O

in

o

m

o

00

ri

cn

in

O

in

VO

00

o

O

in

o

00

in

•n

o

00

o'

m

o

O

o

in

VO

m

o

r-

O

r--

o

m

in

o

ci

cn

in

O

in

(N

in

m

o

’-H

in

m

(N

in

o

in

o'

Steps

m

(N

in

00

(N

VO

(N

o

VO

(N

m

t--

m

(N

Os

Toxoniella taitensis

-

o

-

o

O

o

O

-

O

-

o

-

o

Toxoniella rogoae

-

o

-

o

-

o

O

-

o

-

o

-

o

Scotina palliardii

O-

-

o

-

in

o

-

O

o

-

o

Scotina gracilipes

o-

O-

-

o

-

in

o

-

o

O

o

-

o

Scotina celans

o-

-

o

-

in

o

-

o

-

o

Sagan a rutilans

-

o

-

o

-

o

-

o

-

Neoanagraphis chamberlini

-

o

-

o

-

m

o

-

o

-

o

-

o

Mesiotelus tenuissimus

-

o

-

o

-

o

-

o

-

o

-

Mesiotelus cyprius

-

o

-

o

-

o

-

o

-

o

-

Liocranum rupicola

-

o

-

o

-

o

-

o

-

o

-

Liocranum giersbergi

-

o

-

o

-

o

-

o

-

o

-

Liocranoeca striata

-

o

-

o

-

o

-

o

-

o

Hesperocranum rothi

-

o

-

o

-

(N

-

o

-

o

-

o

Cybaeodes marinae

-

o

-

o

-

o

-

o

-

o

-

o

-

o

-

Cybaeodes mallorcensis

-

o

-

o

-

o

-

o

-

o

-

o

-

o

-

Cteniogaster toxarchus

-

o

-

o

-

o

-

o

Cteniogaster hexomma

-

o

-

o

-

o

-

o

Cteniogaster conviva

o

-

o

-

o

-

o

-

o

-

o

Apostenus humilis

o-

O-

-

o

-

o

-

o

-

o

o-

Apostenus fuscus

o-

-

o

-

o

-

o

-

o

o-

Agroeca proxima

-

o

-

o

-

o

-

o

-

o

-

o

-

o

Agroeca parva

-

o

-

o

-

o

-

o

-

o

-

o

Agroeca cuprea

-

o

-

o

-

o

-

o

-

o

-

o

Agroeca brunnea

-

o

-

o

-

o

-

o

-

o

-

o

-

o

Agraecina lineata

o

-

o

-

o

-

o

-

o

-

o

-

o

Creugas bajulus

o

-

O

o-

-

o

-

o

-

o

-

o

-

Corinna nitens

o

-

O

o-

-

o

-

o

-

o

-

Characters

<s

m

tn

'O

r-

00

as

o

<s

m

tn

'O

r-

00

Os

sc

Os

49