East African pholcid spiders: an overview, with descriptions of eight new species (Araneae, Pholcidae)

Annotated of East African genera and species Identi ﬁ cation key to East African pholcid genera Abstract. This paper summarizes current knowledge about East African pholcids. East Africa is de ﬁ ned as the area from 12°S to 5°N and from 28° to 42°E, including all of Uganda, Kenya, Burundi, Rwanda, and Tanzania. An annotated list of the 15 genera and 87 species recorded from this area is given, together with distribution maps and an identi ﬁ cation key to genera. Most East African species (90%) belong to one of only six genera: Buitinga Huber, 2003 (21 species); Smeringopus Simon, 1890 (18); Pholcus Walckenaer, 1805 (17); Spermophora Hentz, 1841 (12); Leptopholcus Simon, 1893 (5) and Quamtana Huber, 2003 (4). Eight species for which DNA sequence data have been published recently are newly described: Buitinga batwa sp. nov., B. wataita sp. nov., Spermophora mau sp. nov., S. maathaiae sp. nov., S. bukusu sp. nov., S. kirinyaga sp. nov., S. kyambura sp. nov. and Quamtana nyahururu sp. nov. Crossopriza johncloudsleyi Deeleman-Reinhold & van Harten, 2001, previously only known from Yemen, is redescribed based on specimens from Kenya. Additional new records are given for 21 previously described species.


Introduction
The exploration of east African pholcid spiders started late and progress was slow for almost a century.Strand (1906aStrand ( , 1906bStrand ( , 1907Strand ( , 1913) ) and Tullgren (1910) reported the first six species for the area considered here, and until the end of the 20 th century, the number increased to a mere 19 currently valid species.That this is in stark contrast to the actual diversity of the region was first demonstrated by a spider inventory made in the Tanzanian Uzungwa Mountains (Sørensen et al. 2002;Sørensen 2003).Pholcids were not only the most abundant spider family but proved to be represented in the area by a rich and highly endemic fauna.Most of this material and of other material available in collections has since been studied (Huber 2003a(Huber , 2007(Huber , 2009(Huber , 2011(Huber , 2012)), and at the level of genera our knowledge about east African pholcids can probably be considered as fairly complete.At the level of species, the situation is very different.Several lines of evidence (unpublished material in collections; distribution patterns; poorly sampled areas; data from recent expeditions focusing on pholcids) indicate that many species remain undescribed and even more may remain undiscovered.The present paper is an attempt to summarize current knowledge about east African pholcids, complementing recent revisionary work that concentrated on major genera (Huber 2003a(Huber , 2011(Huber , 2012)).Rather than closing the subject it is meant to facilitate access to the available information, to guide future collecting expeditions, and to point out some biologically interesting open questions.
Methods and terminology are as in recent revisions (Huber 2011(Huber , 2012)).Measurements are in mm unless otherwise noted.eye measurements are +/-5 µm.epigyna were cleared in a warm NaOH solution and stained with chlorazol black.Locality coordinates are in round brackets when copied from labels and original publications or when received directly from collectors, in square brackets when originating from some other source (such as online gazetteers, Google Earth, MRAC database, etc.).The following abbreviations are used: ALe: anterior lateral eyes; ALS: anterior lateral spinnerets; AMe: anterior median eyes; a.s.l.: above sea level; L/d: length/diameter; PMe: posterior median eyes.

Notes
Some of the newly collected specimens above are assigned tentatively because they have a shorter abdomen and a slightly different color pattern.The MrAC has a very similar undescribed species from Burundi (Mt.Musumba), with slightly different procursus tip and longer proximal projections on the male chelicerae (MRAC 226400).(Tullgren, 1910).Tanzania (Tullgren 1910).

Note
No material of this species was available to Huber (2003a), and the type material seems to be lost.New material from Kenya (2 ♂♂, 3 ♀♀ from Kirimeri Forest, near Runyenyere, near Mt.Kenya), in MRAC (215277, 293) might belong to this species but topotypical material (from Kilimanjaro) is needed to properly redescribe this species and to evaluate the Kenyan specimens.(Tullgren, 1910).Tanzania (Tullgren 1910;Berland 1920;Huber 2003a sub Buitinga nigrescens).

Note
The Tanzanian specimens described as B. nigrescens in Huber (2003a) are very probably B. globosa (see above).Females of the two species appear indistinguishable, but newly collected males (and females) from Ol Donyo Sabuk National Park and Hell's Gate National Park (in ZFMK,Ken 114) indicate that B. nigrescens might in fact be a valid species (with distinctively hooked bulbal apophysis).However, since no topotypical males are known of B. nigrescens (Mt.Kenya, Naro Moru Track), the identity of these newly collected specimens remains dubious.17.Buitinga ruhiza Huber, 2003.Uganda (Huber 2003a).

Modisimus Simon, 1893
Modisimus is a New World genus; only the synanthropic M. culicinus (Simon, 1893) has a pantropical distribution.This tiny species is probably not rare, but it has previously been recorded only once from Africa (Congo DR; Lawrence 1938; sub Hedypsilus lawrencei).It is here newly recorded for East Africa (Fig. 20).(Simon, 1893).Newly recorded for East Africa (Kenya).

Quamtana Huber, 2003
Quamtana is most diverse in southern Africa, but a few species occur as far north as Cameroon, Congo DR, Uganda (Huber 2003b), and Kenya (herein).Four species are currently known from East Africa (Fig. 25).The new species Q. nyahururu represents the first record of the genus for Kenya and Tanzania.Huber, 2003. Congo DR (Huber 2003b).Huber, 2003. Uganda (Huber 2003b).

Spermophora Hentz, 1841
Spermophora is in fact an Asian genus (Huber 2005) and African representatives have tentatively been assigned to it for lack of a better hypothesis (Huber 2003a).Recent molecular data (Dimitrov et al. 2012) suggest that at least the six Kenyan species (Fig. 27) are closely related to the East African genus Buitinga.Twelve east African species are currently assigned to Spermophora.83.Spermophora minotaura Berland, 1920.Kenya (Berland 1920;Simon & Fage 1922;Huber 2003a).

Note
This species shows slight morphological variation (male cheliceral apophyses) and conspicuous color variation.The position of the frontal male cheliceral apophyses varies from almost parallel to strongly diverging.Huber (2003a) mentioned color variation with respect to the epigynum but it also occurs on carapace and abdomen in both males and females.Below, specimens are divided into three morphs: light morph (median dark band on carapace, light sternum, two large marks on abdomen or monochromous light abdomen; Figs 55, 56); medium morph (pair of narrow dark stripes on carapace do not reach lateral margins, dark sternum; abdomen pattern in principle as in dark morph but less developed), and dark European Journal of Taxonomy 29: 1-44 ( 2012) morph (wide black stripes on carapace reaching lateral margins, dark sternum; complex dark pattern on abdomen; Figs 53, 54).In Chogoria Forest at 2460 m, light and medium morphs were found together, but in different microhabitats: the light morph among vegetation, with the apex of the domed web attached to the underside of a green leaf (Fig. 8); the medium morph close to the ground, with the same type of web among mosses and in small cavities.However, in other localities, light and medium/dark morphs were collected by fogging (see below), suggesting that medium/dark morphs also occur higher in the vegetation (on mosses of trees?).Low molecular distances (Dimitrov, Astrin & Huber 2012) and indistinguishable genitalia suggest that this might be a case of ecological speciation at an early stage.

Diagnosis
Easily distinguished from known congeners by male clypeus modification (shapes of pair of apophyses;

Female
In general similar to male but femur 1 without spines, clypeus unmodified, chelicerae without stridulatory ridges; tibia 1 in 6 females from Kenya: 4.6-5.2(mean 4.8); in 3 females from Yemen: 3.4-3.5.Without stridulatory apparatus between prosoma and abdomen.epigynum with large and complex frontal plate with pair of long lateral apophyses, pair of deep pits, and posterior elevated transversal ridge; simple posterior plate; internal genitalia as in Figs 63 and 74.Paratypes with slightly smaller epigyna with slightly shorter projections, otherwise identical (also in dorsal view).ALS with only two spigots each.

Distribution
Known from Yemen and from several localities in Kenya, Rift Valley Province (Fig. 19).

Diagnosis
Distinguished from known congeners by combination of male cheliceral armature (Fig. 77), male palp (shapes of procursus and bulbal process; Figs 75, 76), color pattern on prosoma and abdomen , and epigynum with straight scape in anterior position and distinctive posterior structure (Fig. 78).

Etymology
The species is named after the Taita people (also Wataita or Wadawida), a Kenyan ethnic group located in the Taita-Taveta District; noun in apposition.
Body.Habitus as in Figs 32-34; ocular area slightly elevated; no thoracic furrow, with small median cone on carapace posteriorly; clypeus unmodified.Chelicerae as in Fig. 77, with proximal lateral apophyses, pair of small apophyses proximally near median line, and frontal apophyses in relatively proximal position provided with four modified hairs each (Fig. 80).Sternum as wide as long (0.65), unmodified.
Variation.The color patterns are very constant but two elements may be less distinct or even missing: the anterior dorsal abdominal mark and the anterior lateral pair of marks on the carapace.Tibia 1 in 35 other males: 5.8-6.9(mean 6.4).

Female
In general similar to male but ventral abdominal pattern slightly different (anterior paired element shorter, median element longer; Fig. 35), triads closer together (distance PME-PME 140 µm), and carapace without median cone.Tibia 1 in 25 females: 4.5-5.7 (mean 5.2).Epigynum a simple plate with straight scape in anterior position, with distinctive structure posteriorly (Fig. 78); internal genitalia as in Fig. 79, with small pore plates scattered over wide area.

Etymology
The species is named after the Batwa pygmies of south-western uganda; noun in apposition.

Diagnosis
Distinguished from very similar S. maathaiae sp.nov.by shape of female epigynal scape (longer; tip pointed; Fig. 90) and by details of procursus tip (slightly different shapes of sclerites; Figs 87, 88).From other species by male cheliceral armature (shapes of frontal apophyses; Fig. 89) and by pointed tip of female epigynal scape.

Etymology
The species name is derived from the type locality; noun in apposition.

Body. Habitus as in Figs
PalPs.As in Figs 87 and 88, coxa with retrolatero-ventral apophysis, trochanter with retrolateral apophysis, femur barely modified (except dorsal apophysis proximally), procursus with ventral flap and several complex distal elements that appear hinged to proximal part, distinctive spine-like process visible in retrolateral view, bulb with three cone-shaped projections and long embolus.

Distribution
Known from type locality only (Fig. 27).93).From other species by male cheliceral armature (shapes of frontal apophyses; Fig. 94) and by widened and sclerotized tip of female epigynal scape.

Etymology
The species is named in honor of Prof. Wangari Maathai, founder of the Green Belt Movement and first African woman to win the Nobel peace prize.

Diagnosis
Distinguished from the similar S. minotaura Berland, 1920

Etymology
The species is named after the Bukusu, a Kenyan tribe located mainly on the foothills of Mount elgon; noun in apposition.
Color.Prosoma ochre-yellow with large lateral black marks and Y-mark behind ocular area, ocular area laterally darkened, sternum monochromous black, legs ochre-yellow, with darker rings on femora (subdistally) and tibiae (proximally, subdistally), abdomen grey with distinct black pattern, also ventrally.49; ocular area elevated, with pair of horns between triads (Fig. 45); carapace only anteriorly with shallow thoracic furrow, clypeus barely modified, only large membranous area that appears collapsed (in all males examined).Chelicerae as in Fig. 99, each distal frontal apophysis with straight row of six cone-shaped modified hairs, proximal frontal apophyses slightly flattened (more slender in lateral view).Sternum wider than long (0.86/0.70), unmodified.
Variation.Tibia 1 in 8 other males: 4.3-6.1 (mean 5.1).Males from Kitum Cave and from Kitale differ slightly in several respects (arrows in Figs 97 and 98): pointed process on ventral flap of procursus larger and more curved; shapes of distal procursus structures slightly different; bulbal pointed process slightly longer and more curved.

Etymology
Kirinyaga is a Kikuyu and embu name for Mount Kenya; noun in apposition.
Body.Habitus as in Fig. 50; ocular area slightly elevated, without horns; shallow but distinct thoracic furrow; clypeus unmodified.Chelicerae as in Fig. 104, with proximal lateral apophyses, large frontal apophyses (without modified hairs) and three long modified hairs on each side, the distal one accompanied by small black cone.Sternum wider than long (0.72/0.64), unmodified.
PalPs.As in Figs 102 and 103, coxa with retrolatero-ventral apophysis, trochanter with retrolateral apophysis, femur barely modified, procursus with ventral flap and several complex distal elements that appear hinged with proximal part, bulb with only one distally bifid process and small rounded projection on opposite side.legs.Without spines and curved hairs, few vertical hairs; retrolateral trichobothrium on tibia 1 at 14%; prolateral trichobothrium absent on tibia 1, present on other tibiae; tarsus 1 with ~20 pseudosegments.

Distribution
Known from type locality only (Fig. 27).
Males from Ghana seem to have indistinguishable palps but slightly longer and more widely spaced cheliceral apophyses and slightly shorter legs: tibia 1 in 21 males from Ghana: 2.0-2.3 (mean 2.1); they are therefore assigned tentatively.
Female in general similar to male.Tibia 1 in 34 females from uganda, Congo Dr and Cameroon: 1.6-2.0(mean 1.8).Epigynum extremely simple, no external modification visible in dissecting microscope; only internal frontal sac-like structure visible through cuticle; pair of pockets only visible in compound microscope (Fig. 110).In females from Ghana, these pockets are barely visible and slightly wider apart; tibia 1 in 20 females from Ghana: 1.4-1.6 (mean 1.55).

Distribution
Widely distributed in tropical Africa (Uganda, Congo DR, Cameroon; possibly also Ghana).Figure 27 shows the only record from east Africa.

Diagnosis
Distinguished from known congeners by combination of male cheliceral armature (Fig. 113), male palp (shapes of procursus and bulbal process; Figs 111, 112), and absence of AME.The South African Quamtana ciliata (Lawrence, 1938) and Q. mbaba Huber, 2003 are similar but both have AMe and are much larger (total body length ~3.0).
Color.Prosoma pale whitish with wide dark median band on carapace including clypeus but not ocular area, legs pale ochre-yellow, without darker rings, abdomen monochromous pale grey.legs.Without spines and curved hairs, few vertical hairs (most hairs missing); retrolateral trichobothrium on tibia 1 at 9%; prolateral trichobothrium absent on tibia 1, present on other tibiae; pseudosegments not visible in dissecting microscope.

Distribution
Known from several localities in Kenya and Tanzania (Fig. 25).

Body.
Habitus as in Figs 59 and 60; ocular area barely elevated, each triad on low hump; no thoracic furrow; clypeus unmodified.Chelicerae as in Fig. 113, with proximal lateral apophyses and distal frontal apophyses provided with two long modified hairs each (Fig. 114), without stridulatory ridges.Sternum wider than long (0.48/0.40), unmodified.PalPs.As in Figs 111 and 112, coxa unmodified, trochanter with short retrolatero-ventral apophysis, femur barely modified, procursus simple except distally, bulb with T-shaped process and transparent straight embolus arising from bulb next to T-shaped process. .