European Journal of Taxonomy 59: 1-44

http://dx.doi.org/10.5852/ejt.2013.59

ISSN 2118-9773

WWW. europeanj oumaloftaxonomy. eu

2013 • Bernhard A. Huber & Peter Kwapong

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

Research article

um:lsid:zoobank.org:pub:F3B32952-A769-4A41-92EB-3EBF52AD7F7F

West African pholcid spiders: an overview,

with descriptions of five new species (Araneae, Pholcidae)

Bernhard A. HUBER' & Peter KWAPONG"

^ Alexander Koenig Researeh Museum of Zoology, Adenauerallee 160, 53113 Bonn, Germany

Email: [email protected] (eorresponding author)

^ Department of Entomology & Wildlife - International Stingless Bee Centre (ISBC),

Sehool of Biologieal Seienees, University of Cape Coast, Cape Coast, Ghana

Email: [email protected]

^ um:lsid:zoobank.org:author:33607F65-19BF-4DC9-94FD-4BB88CED455F

^ um:lsid:zoobank.org:author:DA9A306D-7C9B-4DF 1-9529-004516F24AE7

Abstract. This paper summarizes eurrent knowledge about West Alfiean pholeids. West Alfiea is here

defined as the area south of 17°N and west of 5°E, ineluding mainly the Upper Guinean subregion of the

Guineo-Congolian eenter of endemism. This ineludes all of Senegal, The Gambia, Guinea Bissau, Guinea,

Sierra Feone, Fiberia, Ivory Coast, Ghana, Togo and Benin. An annotated list of the 14 genera and 38 speeies

reeorded from this area is given, together with distribution maps and an identifieation key to genera. Five

speeies are newly deseribed: Anansus atewa sp. nov., Artema bunkpurugu sp. nov, Leptopholcus kintampo

sp. nov., Spermophora akwamu sp. nov, and S. ziama sp. nov. The female of Quamtana kitahurira is newly

deseribed. Additional new reeords are given for 16 previously deseribed speeies, ineluding 33 new eountry

reeords. Distribution patterns of West Alfiean pholeids are diseussed, as well as possible explanations for

relatively low West Alfiean pholeid speeies diversity as eompared to Central and East Alfiea.

Keywords. Pholeidae, West Afriea, taxonomy, identifieation key, diversity.

Huber B.A. & Kwapong P. 2013. West African pholcid spiders: an overview, with descriptions of five new species

(Araneae, Pholcidae). European Journal of Taxonomy 59: 1-44. http://dx.doi.org/10.5852/eit.2013.59

Introduction

In a recent overview on African rain forest ecology and conservation (Weber et al. 2001), one of the

major “strikingly similar principles” emerging from studies of “lesser known taxa” was the lack of

basic information such as species lists, species distributions, and basic ecology. These “lesser known

taxa” were not invertebrates though, but birds, fishes, amphibians, and reptiles (in contrast to mammals

and plants). Invertebrates are barely mentioned in this otherwise superb book, probably because our

knowledge of most groups is not even close to sufficient to track basic questions of rain forest ecology

and conservation. While the lack of data is beyond question and may sometimes even appear hopelessly

overwhelming, it is also worth to focus on the progress that has been made. The present paper adopts the

latter position, concentrating on pholcid spiders and showing that many of the patterns found in plants

and vertebrates are replicated in this particular taxon.

European Journal of Taxonomy 59: 1-44 (2013)

While Pholcidae admittedly make up just a small fraetion of the megadiverse spiders (about 3% of

deseribed speeies; Platniek 2013), our knowledge about this family has inereased substantially over the

last deeade. Advanees inelude alpha taxonomy (speeies numbers for Afriean taxa have more than tripled

sinee 2002, from 88 to now 287; Huber 2013, herein), but also phylogenetie data (summary in Huber

2011a), distribution patterns (e.g., Huber 2011b, 2012,2013) andbasie eeology (e.g., Huber 2009,2013;

Huber et al. 2013). Pholeidae are most diverse in well-preserved tropieal forests, but they also oeeur in

relatively arid regions (e.g., Huber 2001; Huber & Breseovit 2003); they range from sea level to about

4000 m (Huber 2000, 2012); they inelude both widespread speeies with substantial dispersal eapabilities

(even though ballooning does not seem to oeeur in Pholeidae) and many small-seale endemies (Huber

2011b, 2013); and numerous taxa within the family inelude eeologieally very different representatives

(whieh is refleeted in a wide range of body shapes and eolors; ef Figs 1-22), suggesting multiple

independent evolutionary shifts among mierohabitats (Dimitrov et al. 2013).

Considering the preferenee of Pholeidae for tropieal forests, a high diversity in Alfiean rainforests was

expeeted. Within the Guineo-Congolian eenter of endemism. West Alfiean forests were predieted to be less

diverse than Central Alfiean forests, both for historieal and eurrent reasons (only about 12% of the original

tropieal moist forests remain in West Alfiea, as opposed to 59% in Central Alfiea; Naughton-Treves & Weber

2001). West Alfiea was also predieted to be less diverse than East Alfiea with its wider range of altitudes,

its mosaie of different vegetation zones, and its highly diverse Eastern Are. However, with pholeid material

in existing eolleetions it seemed impossible to even roughly estimate basie data like speeies numbers and

distribution patterns. For this reason, a series of six expeditions was undertaken to East Alfiea (Kenya,

Uganda), Central Alfiea (Cameroon, Gabon), and West Alfiea (Guinea, Ghana) in order to obtain eomparable

data aeross the Alfiean eontinent. The present paper summarizes the data on West Alfiea, eomplementing

revisions of all major taxa in the area (Huber 2011b, 2012, 2013) and a previous summary on East Alfiean

Pholeidae (Huber & Wami 2012). The flnal part of this trilogy (Central Alfiea) is in preparation.

Material and methods

A large part of the material studied herein was eolleeted during expeditions to Guinea (2008) and Ghana

(2013). This material is eurrently deposited at Zoologisehes Forsehungsmuseum Alexander Koenig,

Bonn (ZFMK). Further material was borrowed from the following institutions:

AMNH = Ameriean Museum of Natural History, New York

CAS = California Aeademy of Seienees, San Franeiseo

MCZ = Museum of Comparative Zoology, Cambridge

MNHN = Museum national d’Histoire naturelle, Paris

MRAC = Musee royal de FAfrique Centrale, Tervuren

SMF = ForsehungsinstitutundNaturmuseum Senekenberg, Frankfurt

USNM = National Museum of Natural History, Washington D.C.

Methods and terminology are as in reeent revisions (Huber 2011b, 2012, 2013). Measurements are in

mm unless otherwise noted. Eye measurements are +/- 5 pm. Epigyna were eleared in a warm NaOH

solution and stained with ehlorazol blaek. Eoeality eoordinates are in round braekets when eopied from

labels and original publieations or when reeeived direetly from eolleetors, in square braekets when

originating Ifom some other souree (sueh as online gazetteers, Google Earth, MRAC database, ete.). The

following abbreviations are used:

AEE = anterior lateral eyes

AES = anterior lateral spinnerets

AME = anterior median eyes

a.s.l. = above sea level

E/d = length/diameter

PME = posterior median eyes.

HUBER B.A. & KWAPONG R, West African pholcid spiders

Figs 1-13.1. Pholcus doucki Huber, 2011 from Doucki, Guinea. 2-3. Pholcus kakum Huber, 2009 from

Foret Classee de Ziama, Guinea (2) and Kakum N.P., Ghana (3). 4. Leptopholcus kintampo sp. nov.

from Kintampo, Ghana. 5. Leptopholcus tipula (Simon, 1907) from Kakum N.P, Ghana. 6. Nyikoa

limhe Huber, 2007 from Kakum N.P, Ghana. 7-8. Pehrforsskalia conopyga Deeleman-Reinhold & van

Harten, 2001 from Atewa, Ghana. 9-10. Anansus atewa sp. nov. from Atewa, Ghana. 11-13. Spermophora

akwamu sp. nov. from Kakum N.P, Ghana (13: web). Photos BAH.

European Journal of Taxonomy 59: 1-44 (2013)

Figs 14-22. 14-15. Artema bunkpurugu sp. nov. from near Bunkpumgu (14) and from Bunkpumgu

(15), Ghana. 16. Crossopriza lyoni (Blackwall, 1867) from Conakry, Guinea. 17-18. Physocyclus

glohosus (Taezanowski, 1874) from near Mpraeso, Ghana. 19. Smeringopus cylindrogaster (Simon,

1907) from Ankasa N.P., Ghana. 20. Smeringopina pulchra (Millot, 1941) from Kakum N.P., Ghana.

21-22. Smeringopina bomfobiri Huber, 2013 from Atewa, Ghana. Photos BAH.

HUBER B.A. & KWAPONG R, West African pholcid spiders

The geographic area considered here (south of 17°N, west of 5°E; Fig. 23) is somewhat arbitrary,

especially regarding the eastern limit of “West Africa”, i.e. its border to “Central Africa”. In contrast to

the major textbook on African spiders (Dippenaar-Schoeman & Jocque 1997), West Africa as defined

here is largely congruent with the usage in African Rain Forest Ecology & Conservation (Weber et al.

2001; e.g., p. 31, 119, 430). Biologically it encompasses mainly the Upper Guinean subregion of the

Guineo-Congolian center of endemism (White 2001). The eastern limit of this subregion is not clear

since the main current gap (Dahomey Gap; ~0-3°E) is not always congruent with distribution limits

of taxa, which in many cases are further east (Cross River Basin of eastern Nigeria, ~8°E) (Dowsett-

Eemaire & Dowsett 2001; Eawson & Klemens 2001; Maley 2001). For the present overview, however,

the exact eastern limit (western vs. eastern Nigeria) is fairly irrelevant since the pholcid fauna of Nigeria

remains largely unknown.

Results

Class Arachnida Cuvier, 1812

Order Araneae Clerck, 1757

Family Pholcidae C.F. Koch, 1851

Annotated list of West African genera and species

Pholcidae described from West Africa as defined herein, with countries, references, and new records.

Anansus Huber, 2007

Anansus is widely distributed in tropical Africa (Huber 2007; Huber & Wami 2012) with at least three

species endemic to West Africa (Fig. 23).

1. Anansus aowin Huber, 2007. Ivory Coast (Huber 2007).

2. Anansus atewa sp. nov. (see page 21). Ghana.

3. Anansus ewe Huber, 2007. Ghana (Huber 2007); newly recorded for Guinea and Ivory Coast (Ivory

Coast records tentative, no males available).

New records

GHANA: Central Region: 20 SS 14 $ $ 3 juvs (3 vials) in ZFMK (Ar 10450-52), Kakum National Park

(5°20.9’N, 1°23.0’W), 160 m a.s.L, forest near entrance, day collecting, 19-20 Feb. 2013 (B.A. Huber);

2 $ in pure ethanol, in ZFMK (Gha 150), same data.

GUINEA: 4 SS 5 $ $ in pure ethanol in MRAC (236926), Mt. Nimba, Zougue valley, near Gbakore

mine camp [7°42’N, 8°24’W], young secondary gallery forest, canopy fogging, 780 m a.s.L, 5 Oct. 2011

(D. Van den Spiegel, A. Hemard); 2 5 ? in pure ethanol in MRAC (236932), Mt. Nimba, near Gouan

camp or “Station de Pompage Zie” [7°40’N, 8°22’W], sieving of litter under “matete” (high grass), open

area near road, 1250 m a.s.L, 11 Oct. 2011 (D. Van den Spiegel, A. Hemard).

IVORY COAST: I $ (assigned tentatively) in MRAC (230172), Foret de Tai, station du Centre de

Recherche Ecologique (CRE) [5°50’N, 7°21’W], sieving litter, 29 Feb. 2010 (R. Jocque, D. Van den

Spiegel); 1 $ (assigned tentatively) in MRAC (230397), Adiopo Doume, Centre Suisse de Recherche

Scientifique (CSRS) [5°20’N, 4°20’W], winkler extraction, 15 Feb. 2010 (R. Jocque, D. Van den

Spiegel).

European Journal of Taxonomy 59: 1-44 (2013)

Walckenaer, 1837

The natural distribution of Artema ranges from the Sudanian region of West Afriea (Fig. 24) to Central

Asia. Millot (1941) was the first to reeord the genus from West Afriea, and he noted that most of his

material of “A. mauriciana Walekenaer, 1837” {=A. atlanta Walekenaer, 1837) did not agree well with

previous illustrations of this pantropieal speeies. After eomparing numerous West Afriean speeimens

with A. atlanta from all over the World, we agree on these substantial morphologieal differenees and

formally deseribe the West Afriean speeies below.

Figs 23-24. Known distributions of the gonovdi Anansus Huber, 2007 and Artema Walekenaer, 1837 in

West Afriea. Question marks denote uneertain identifieations (Anansus ewe\ two loealities with females

only) or impossible identifieations [“?G)juveniles only].

HUBER B.A. & KWAPONG R, West African pholcid spiders

4. Artema atlanta Walckenaer, 1837. Guinea (Millot 1941: only the variant male from Kouroussa);

newly recorded for Togo and Benin.

New records

GUINEA: 1 (J in pure ethanol, in ZFMK (Gui 82), Conakry, in building (9°3UN, 13°43’W), ~30 m

a.s.l., 18 Nov. 2008 (B.A. Huber). Guinee Forestiere: 1 $ in ZFMK (Ar 10453), Nzerekore (~7°45.3’N,

8°49.2’W), 480 m a.s.l., in building, 28 Nov. 2008 (B.A. Huber). Moyenne-Guinee: 1 (? in pure ethanol,

in ZFMK (Gui 111), Pita, at building (11°03.6’N, 12°23.7’W), 1010 m a.s.l., 25 Nov. 2008 (B.A. Huber).

Kankan: 1 (? in MNHN, Kouroussa [10°39’N, 9°53’W], Aug. 1937 (J. Millot) [this is probably Millot’s

(1941) variant male of his 'Artema mauriciana''].

TOGO: 2 (?(? 5 ?? + juvs in MRAC (136023), Niamtougou [9°46’N, 1°07’E], 2F/24. Jul. 1969 (F.

Puylaert).

BENIN: 3 4 $$ in MRAC (215041), Nikki [9°56’N, 3°13’E], Jul. 2003 (Djaouga).

Note

Millot’s (1941) specimens from Kindia, Guinea (2 juvs; $ missing), Ferkessedougou, Ivory Coast (1

juv.), and Gao, Mali (2 juvs) are in MNHN but juveniles of A. atlanta and A. bunkpurugu sp. nov.

are indistinguishable so their identity is unclear. The same is true for juvenile specimens from Dakar,

Senegal in AMNH, from Freetown, Sierra Feone in SMF, and from Bandiagara (70 km E Mopti), Mali

in CAS.

5. Artema bunkpurugu sp. nov. (see page 24). Ghana, Burkina Faso, Mali, Guinea, Togo.

Crossopriza Simon, 1893

Apart from the pantropical synanthropic C. lyoni (Blackwall, 1867), only C. soudanensis Millot, 1941

is known to occur in West Africa (Fig. 25).

6. Crossopriza lyoni (Blackwall, 1867). Mali, Nigeria (Huber et al. 1999); newly recorded for Senegal,

The Gambia, Guinea, Sierra Feone, Ivory Coast, Ghana, Burkina Faso, Benin, and Niger.

New records

SENEGAF: Kaolack Prov.: 1 $ 1 juv. in ZFMK (Ar 5207), Sonkoron, Kaymor Region [~13°47’N,

15°33’W], Jul. 1993 (M. Sarr.).

THE GAMBIA: 3 ?? in ZFMK (Ar 5404), Banjul (13°27.6’N, 16°34.7’W), on wall, 20 Oct. 2008

(Yu.M. Marusik); 1 c?' 2 in MRAC (205485), Serekunda [13°26’N, 16°41’W], in grass, 20 Aug.

1996 (G. Beyens).

GUINEA: 1 (J 2 $ ? in pure ethanol in ZFMK (Gui 103), Conakry, in building (9°34.1’N, 13°39.7’W),

40 m a.s.l., 18 Nov. 2008 (B.A. Huber). Guinee Forestiere: 1 (? in ZFMK (Ar 10454), Nzerekore

(~7°45.3’N, 8°49.2’W), 480 m a.s.l., in building, 28 Nov. 2008 (B.A. Huber); 3 (?(? 4 in ZFMK

(Ar 10455), Kissidougou, in building (9°11.2’N, 10°05.6’W), 540 m a.s.l., 26 Nov. 2008 (B.A. Huber);

2553 juvs in pure ethanol, in ZFMK (Gui 90), same data. Basse-Guinee: 1 1 ? in ZFMK (Ar

10456), Kindia, in building (10°03’N, 12°51’W), 400 m a.s.l., 20 Nov. 2008 (B.A. Huber).

SIERRA FEONE: 1 $ in MRAC (174657), Makeni [8°53’N, 12°03’W], “maison”, 19 Jan. 1993 (F.

Rensonnet).

IVORY COAST: 2 $$ 1 juv. in CAS, Bouake [7°42’N, 5°02’W], 10 Jan. 1991 (W.J. Pulawski); 1 $

in SMF, Comoe National Park, Research Station Univ. of Wurzburg [~9.1°N, 3.7°W], 23 Sep. 2002 (S.

Otto); 5 (?(? 11 ?? 9 juvs in CAS, Korhogo [9°28’N, 5°38’W], 12 Jan. 1991 (W.J. Pulawski).

MAFI: 1 (? in CAS, 10 km E Sevare (14°30’N, 4°00’W), 15 Jul.-l Sep. 1977 (WH. Settle); 1 ? in

MRAC (136760), Kogoni, 60 km N Mono [~14°50’N, 6°00’W], Dec. 1969 (G. Pierrard); 2 8 ? ?

European Journal of Taxonomy 59: 1-44 (2013)

in MRAC (136761, 138808), MTesoba [Mpessoba, 12°40’N, 5°43’W], Sep.-Oct. 1969 and Sep. 1970

(G. Pierrard).

GHANA: Northern Region: 1 5 in ZFMK (Ar 10457), Bunkpumgu (10°31.rN, 0°05.5’E), 230 m

a.s.L, in buildings, 7 Mar. 2013 (B.A. Huber); 1 (5 1 $ in pure ethanol, in ZFMK (Gha 127), same data;

5 $ $ in ZFMK (Ar 10458), Bunkpumgu (10°31.3’N, 0°05.8’E), 250 m a.s.L, in building, 6 Mar. 2013

(B.A. Huber); 2 SS 2 5 ? in ZFMK (Ar 10459), near Bunkpumgu (10°32.0’N, 0°02.2’E), 210 m a.s.L,

under eonerete bridge, 8 Mar. 2013 (B.A. Huber); 1 $ 1 juv. in pure ethanol, in ZFMK (Gha 142),

same data; 1 (? 3 $$ in ZFMK (Ar 10460), near Bimbaga (10°34.5’N, 0°03.8’W), 330 m a.s.L, under

eonerete bridge, 8 Mar. 2013 (B.A. Huber); 1 5 2 juvs in ZFMK (Ar 10461), near Gambaga (10°31.9’N,

0°25.3’W), 350 m a.s.L, under eonerete bridge, 8 Mar. 2013 (B.A. Huber); \ S in ZFMK (Ar 10462),

near Gambaga (10°29.3’N, 0°28.9’W), 300 m a.s.L, under eonerete bridge, 8 Mar. 2013 (B.A. Huber);

1 $ in ZFMK (Ar 10463), Tamale (9°24.4’N, 0°50.4’W), 195 m a.s.L, in building, 5 Mar. 2013 (B.A.

Huber). Greater Aeera Region: 1 5 in ZFMK (Ar 10464), Aeera (5°34.4’N, 0°13.EW), 15 m a.s.L, in

building, 1 Mar. 2013 (B.A. Huber). Ashanti Region: 1 (5 1 $ in ZFMK (Ar 10465), Kumasi (6°43.4’N,

1°38.EW), 310 m a.s.L, in building, 4 Mar. 2013 (B.A. Huber). Brong-Ahafo Region: 1 $ in ZFMK (Ar

10466), Teehiman (7°34.5’N, 1°57.5’W), 420 m a.s.L, in building, 4 Mar. 2013 (B.A. Huber). Eastern

Region: 2 5? in ZFMK (Ar 10467), Suhum (6°02.4’N, 0°27.rW), 215 m a.s.L, in building, 10 Mar.

2013 (B.A. Huber).

BURKINA FASO: ~11 (?(? 18 ?? (3 vials) in MRAC (128066, 068-069), Ouagadougou [12°22’N,

1°3UW], Apr.-May 1965 (B. Roman); \ S in MCZ (34049), same loeality, “bathroom wall”, 16 Jul.

1977 (Brinekerhoff).

BENIN: 4 (?(? 7 ? ? in MRAC (215041 part), Nikki [9°56’N, 13°12’E], Jul. 2003 (Djaouga); 1 (? 5 ? ?

in MRAC (212766 part), Adjohoun [6°42’N, 2°30’E], in house, 23 Dee. 2002 (S. Tehibozo).

NIGER: 4 (?(? 6 ? ? (2 vials) in MCZ (34036-37), “Aoulajouara”, 20 km S Niamey [~13°2rN, 2°08’E],

on wall of eement house, 17-23 Jul. 1971 (R. Dube); ~3 15 $$ (3 vials) in MRAC (137727, 733,

734), Niamey [13°30’N, 2°07’E], Sep.-Oet. 1970 (E. van Impe).

NIGERIA: Osun: 1 $ in MRAC (174602 part), Ile-Ife, Obafemi Awolowo Univ. eampus [7°28’N,

4°34’E], 1991 (H. Segers).

7. Crossopriza soudanensis Millot, 1941. Mali, Burkina Faso (Millot 1941).

Leptopholcus Simon, 1893

Of the ten speeies of Leptopholcus eurrently known from mainland Afriea (Huber 2011b; herein), two

are endemie to West Afriea {L. guineensis Millot, 1941; E. kintampo sp. nov.); the third [L. tipula (Simon,

1907)] oeeurs throughout the entire Guineo-Congolian rainforest (Huber 2011b) (Fig. 26).

8. Leptopholcus guineensis Millot, 1941. Senegal, The Gambia, Guinea (Millot 1941; Huber 2009);

newly reeorded for Ghana.

New records

GUINEA: 7 SS 5 $ $ in pure ethanol, in MRAC (236931), Mt. Nimba, Foret de Zie, near Gouan eamp

or “Station de Pompage Zie” [7°40’N, 8°26’W], 1250 m a.s.L, 3 Oet. 2011 (D. van den Spiegel, A.

Hemard); 1 $ in pure ethanol, in MRAC (236927 part), Mt Nimba, Zougue valley, near Gbakore mine

eamp [7°42’N, 8°24’W], young seeondary gallery forest, eanopy fogging, 780 m a.s.L, 5 Oet. 2011 (D.

Van den Spiegel, A. Henrard).

GHANA: Ashanti Region: 4 SS 1 juv. in ZFMK (Ar 10468), Bomfobiri Wildlife Sanetuary (6°57.3’N,

Ull.UW), 160 m a.s.L, forest near river, 3 Mar. 2013 (B.A. Huber); 1 $ 3 juvs in pure ethanol, in

ZFMK (Gha 131), same data. Eastern Region: 1 f 1 juv. in ZFMK (Ar 10469), near Mpraeso (6°34.1 ’N,

0°43.9’W), 370 m a.s.L, degraded forest near road, 10 Mar. 2013 (B.A. Huber); 2 juvs in pure ethanol, in

HUBER B.A. & KWAPONG R, West African pholcid spiders

ZFMK (Gha 152), same data. Volta Region: 2 33 4 $$ 7juvs inZFMK (Ar 10470), Agumatsa Wildlife

Sanctuary, Wli waterfall (7°06.2’N, 0°36.0’E), -300 m a.s.L, forest near waterfall, 27 Feb. 2013 (B.A.

Huber); 5 juvs in pure ethanol, in ZFMK (Gha 164), same data; 1 (? 1 $ 1 juv. in ZFMK (Ar 10471),

Tagbo waterfall (7°00.7’N, 0°34.4’E), -500 m a.s.L, forest near waterfall, 28 Feb. 2013 (B.A. Huber).

Central Region: 1 $ in MRAC (217306), Kakum Forest, primary forest, 11 Nov. 2005 (R. Jocque, D.

de Bakker, F. Baert).

Figs 25-26. Known distributions of the genera Crossopriza Simon, 1893 md Leptopholcus Simon, 1893

in West Africa. Question marks denote uncertain identifications [“?G)”: juveniles only].

European Journal of Taxonomy 59: 1-44 (2013)

9. Leptopholcus kintampo sp. nov. (see page 28). Ghana.

10. Leptopholcus tipula (Simon, 1907). Guinea, Ghana, Benin (Huber 2009, 2011b); newly reeorded

for Ivory Coast.

New records

IVORY COAST: 1 ? in MNHN, Lamto [6°13’N, 5°02’W?], 26 Aug. 1963 (eolleetor not given); 1 juv.

in MRAC (230358 part), Adiopo Doume, Centre Suisse de Reeherehe Seientifique (CSRS) [5°20’N,

4°20’W], small dry forest, beating, 25 Feb. 2010 (D. van den Spiegel, R. Joeque).

GHANA: Central Region: 1 $ in pure ethanol in ZFMK (Gha 143), Kakum National Park (5°20.9’N,

1°23.0’W), 160 m a.s.L, forest near entranee, day eolleeting, 20 Feb. 2013 (B.A. Huber). Western

Region: 1 (5^ 3 $$ in ZFMK (Ar 10472), Ankasa National Park (5°13.0’N, 2°39.rW), 180 m a.s.L,

forest near entranee, night eolleeting, 22 Feb. 2013 (B.A. Huber); 1 $ 3 juvs in pure ethanol, in ZFMK

(Gha 156), same data.

Micropholcus Deeleman-Reinhold & Prinsen, 1987

Only the synanthropie M. fauroti (Simon, 1887) oeeurs in West Afriea (Huber 2011b) (Fig. 27).

11. Micropholcus fauroti (Simon, 1887). Senegal, The Gambia, Sierra Leone, Guinea (Millot 1941;

Huber 2011b); newly reeorded for Ivory Coast and Ghana.

New records

IVORY COAST: 1 (? in MRAC (227392 part), Bouake [7°4rN, 5°02’W], WARDA station, weed

management experiment, upland riee, 13 Aug. 1995 (A. Russell-Smith).

GHANA: Central Region: 2 2 ?? in ZFMK (Ar 10473), Cape Coast (5°07.rN, 1°17.4’W),

15 m a.s.L, in building, 19 Feb. 2013 (B.A. Huber); 1 5 in ZFMK (Ar 10474), Cape Coast (5°06.3’N,

1°14.8’W), 15 m a.s.L, in building, 23 Feb. 2013 (B.A. Huber). Greater Aeera Region: I f I ^ in ZFMK

(Ar 10475), Aeera (5°34.4’N, 0°15.2’W), 45 m a.s.L, in building, 17 Feb. 2013 (B.A. Huber). Ashanti

Region: 2 in ZFMK (Ar 10476), Kumawu (6°54.3’N, 1°15.4’W), 380 m a.s.L, in building, 3 Mar.

2013 (B.A. Huber). Eastern Region: 2 (?(? in ZFMK (Ar 10477), Suhum (6°02.4’N, 0°27.rW), 215 m

a.s.L, in building, 10 Mar. 2013 (B.A. Huber). Volta Region: 1 1 $ in ZFMK (Ar 10478), Hohoe

(7°09.3’N, 0°29.5’E), 200 m a.s.L, in building, 27 Feb. 2013 (B.A. Huber).

Modisimus Simon, 1893

The pantropieal M. culicinus (Simon, 1893) is the only representative of this otherwise New World

genus in Afriea. It is probably not rare but due to its small size and eryptie lifestyle it has previously

been reeorded only twiee from Afriea (Congo DR and Kenya; Eessert 1938 - sub Hedypsilus lawrencei

Eessert, 1938 - and Huber & Wami 2012). It is here newly reeorded for West Afriea (Fig. 28).

12. Modisimus culicinus (Simon, 1893). Newly reeorded for West Afriea (Sierra Eeone).

New record

SIERRA EEONE: 1 (? in MRAC (148459 part), Freetown, ML Aureol [8°28.7’N, 13°13.3’W], Sep.

1976 (D. Olu-Pitt).

Nyikoa Huber, 2007

Nyikoa is a monotypie genus with the leaf-dwelling N. limbe Huber, 2007 eovering the entire Guineo-

Congolian rainforest (Huber 2007, 2009).

HUBER B.A. & KWAPONG R, West African pholcid spiders

13. Nyikoa limbe Huber, 2007. Ghana, Guinea (Huber 2007, 2009); newly recorded for Ivory Coast.

New records

IVORY COAST: I (? in MRAC (233400), Tai Forest [5°50’N, 7°2rW], beating, 12 Oct. 2010 (D. van

den Spiegel, A. Kablan).

GHANA: Western Region: 4 SS 9 $$ in ZFMK (Ar 10479), Ankasa National Park (5°13.0’N,

2°39.FW), 180 m a.s.L, forest near entrance, day collecting, 22 Feb. 2013 (B.A. Huber); 8 SS 10 $ $

in ZFMK (Ar 10480), same data but night collecting; 1 (5^ 1 $ in pure ethanol, in ZFMK (Gha 155),

Figs 27-28. Known distributions of the genera Micropholcus Deeleman-Reinhold & Prinsen, 1987,

Modisimus Simon, 1893, Nyikoa Huber, 2007, and Quamtana Huber, 2003 in West Africa.

European Journal of Taxonomy 59: 1-44 (2013)

same data; \ 3 in ZFMK (Ar 10481), Ankasa National Park, forest along Big Tree Trail (~5°15.rN,

2°38.4’W), -100 m a.s.L, 23 Feb. 2013 (B.A. Huber). Central Region: 6 33 8 $$ (2 vials) in ZFMK

(Ar 10482-83), Kakum National Park (5°20.9’N, 1°23.0’W), 160 m a.s.L, forest near entranee, day

eolleeting, 19-20 Feb. 2013 (B.A. Huber); \ 3 in ZFMK (Ar 10484), same data but night eolleeting;

1 6' 2 5 ? in pure ethanol, in ZFMK (Gha 146), same data.

Pehrforsskalia Deeleman-Reinhold & van Harten, 2001

Pehrforsskalia ineludes three deseribed speeies (Huber 2011b). Only the widely distributed P conopyga

Deeleman-Reinhold & van Harten, 2001 oeeurs in West Afriea (Fig. 29).

14. Pehrforsskalia conopyga Deeleman-Reinhold & van Harten, 2001. Guinea, Sierra Leone, Ivory

Coast, Nigeria (Deeleman-Reinhold & van Harten 2001; Huber 2009); newly reeorded for The Gambia

and Ghana.

New records

THE GAMBIA: 2 $$ in ZFMK, Banjul, Abuko Nature Reserve (13°23.5’N, 16°39.0’W), primary

gallery forest, 17 Oet. 2008 (Yu.M. Marusik).

IVORY COAST: 1 (? 7 ?? in MRAC (227392 part), Bouake [7°4rN, 5°02’W], WARDA station,

weed management experiment, upland riee, 13 Aug. 1995 (A. Russell-Smith); 5 33 2 5 ? in MRAC

(225855), Gagnoa [6°08’N, 5°57’W], upland riee, 10 Apr. 1995 (A. Russell-Smith); 1 (? 2 $ ? in ZFMK

(Ar 5171), Bobayo near Gagnoa [6°01.6’N, 6°01.9’W], in upland riee, 26 Aug. 1992 (A. Russell-Smith);

\ 3 i ^ 2 Juvs in ZFMK (Ar 5170), “Kroutla” m. Oume [~6°23’N, 5°25’W], 26 Aug. 1992 (A. Russell-

Smith).

GHANA: Eastern Region: 1 3 $$ in ZFMK (Ar 10485), Atewa Hills, Atewa Atwirebu Reserve

(6°13.8’N, 0°33.5’W), 740 m a.s.L, 25 Feb. 2013 (B.A. Huber); 2 $ ? 1 juv. in pure ethanol, in ZFMK

(Gha 136), same data; \ 3 i ^ in ZFMK (Ar 10486), Atewa Hills, Atewa Atwirebu Reserve (6°13.8’N,

0°32.4’W), 500 m a.s.L, degraded forest along road, 24 Feb. 2013 (B.A. Huber); 1 (? 7 $$ in ZFMK

(Ar 10487), near Mpraeso (6°34.1’N, 0°43.9’W), 370 m a.s.L, degraded forest near road, 10 Mar. 2013

(B.A. Huber). Volta Region: 3 33 6 $$ in ZFMK (Ar 10488), Agumatsa Wildlife Sanetuary, Wli

waterfall (7°06.2’N, 0°36.0’E), -300 m a.s.L, forest near waterfall, 27 Feb. 2013 (B.A. Huber); 2 $$

in pure ethanol, in ZFMK (Gha 163), same data; 5 33 4 5 ? in ZFMK (Ar 10489), Tagbo waterfall

(7°00.7’N, 0°34.4’E), -500 m a.s.L, forest near waterfall, 28 Feb. 2013 (B.A. Huber). Ashanti Region:

1 4 5? in ZFMK (Ar 10490), Bomfobiri Wildlife Sanetuary (6°57.3’N, Hll.l’W), 160 m a.s.L,

forest near river, 3 Mar. 2013 (B.A. Huber); 1 $ 1 juv. in pure ethanol, in ZFMK (Gha 132), same data.

Brong Ahafo Region: 1 (? 1 $ in ZFMK (Ar 10491), Booyem (7°39.9’N, 1°57.4’W), 450 m a.s.L, at

large roeks, 4 Mar. 2013 (B.A. Huber); 1 6' 2 $$ in ZFMK (Ar 10492), Kintampo Falls (8°05.3’N,

1°41.9’W), 280 m a.s.L, degraded forest along stream, 5 Mar. 2013 (B.A. Huber).

Walekenaer, 1805

In the area eonsidered here, Pholcus is mainly represented by the five speeies of the West Afriean

endemie guineensis group (Huber 2011b; Fig. 30). Pholcus berlandi Millot, 1941 from Senegal seems

to belong in a speeies group more diverse in northern Afriea (Huber 2011b); Pholcus kakum Huber, 2009

eovers the entire Guineo-Congolian rainforest (Huber 2011b).

15. Pholcus berlandi MiWoi, 1941. Senegal (Millot 1941).

16. Pholcus bourgini Millot, 1941. Guinea (Millot 1941; Huber 2011b).

17. Pholcus chattoni Huber, 2011. Guinea, Ivory Coast (Millot 1941; Huber 2011b).

HUBER B.A. & KWAPONG R, West African pholcid spiders

New records

GUINEA: 1 (5^1 $ in pure ethanol, in MRAC (236923), Mt. Nimba, Chateau [7°40’N, 8°23’W], dry

mine adit, on wall, 10 Oct. 2011 (D. Van den Spiegel, A. Henrard); 3 $$ in pure ethanol, in MRAC

(236924), Mt. Nimba, Pierre Richaud, entrance of disaffected humid mine, adit, on beams, 7 Oct. 2011

(D. Van den Spiegel, A. Hemard).

berlandi

doucki

bourgini

\ kakum \

guineensis

Pholcus

Figs 29-30. Known distributions of the genera Pehrforsskalia Deeleman-Reinhold & van Harten, 2001

and Pholcus Walckenaer, 1805 in West Africa.

European Journal of Taxonomy 59: 1-44 (2013)

18. Pholcus doucki Huber, 2011. Guinea (Huber 2011b).

19. Pholcus guineensis Millot, 1941. Guinea, Sierra Leone (Millot 1941; Huber 2011b).

20. Pholcus kakum Huber, 2009. Guinea, Ivory Coast, Ghana (Huber 2009).

New records

GUINEA: 1 $ 1 juv. in pure ethanol, in MRAC (236927 part), Mt Nimba, Zougue valley, near Gbakore

mine eamp [7°42’N, 8°24’W], young seeondary gallery forest, eanopy fogging, 780 m a.s.L, 5 Get. 2011

(D. Van den Spiegel, A. Henrard).

IVORY COAST: 1 ? in MRAC ^30510), Tai Forest, Centre de Reeherehe Eeologique (CRE) [5°50’N,

7°2EW], beating in forest near Chimpanzee Camp, 22 Feb. 2010 (R. Joeque, M. Diarassouba); 4 $ $ in

MRAC (230295), same loeality but forest aeross river, forest on elayey soil, beating, 20 Feb. 2010; 1 $

in MRAC (230245), same loeality but forest E of eamp, “layon Gerard”, inundated forest, sieved litter,

20 Feb. 2010 (R. Joeque, E. Oulai); \ S in MRAC (233345), Tai' Forest, beating, 1 Sep. 2010 (D. Van

den Spiegel, A. Kablan); 1 $ in MRAC (233747), Foret des Marais-Tanoe [5°10’N, 2°50’W], Aboisso,

Dohouan, beating, 25 Oet. 2010 (A. Kablan).

GHANA: Central Region: 5 SS 8 $$ (2 vials) in ZFMK (Ar 10493-94), Kakum National Park

(5°20.9’N, 1°23.0’W), 160 m a.s.L, forest near entranee, day eolleeting, 19-20 Feb. 2013 (B.A. Huber);

\ S in ZFMK (Ar 10495), same data but night eolleeting; 2 $ ? in pure ethanol, in ZFMK (Gha 147),

same data; 2 $ ? 1 juv. in MRAC (217296 part), Kakum Forest, seeondary forest, beating, 23 Nov. 2005

(R. Joeque, D. de Bakker, E. Baert); I f I ^ (2 vials) in MRAC (217260, 276), same data but beating

between primary and seeondary forest, 11-12 Nov. 2005. Western Region: 2 1 $ in ZFMK (Ar

10496), Ankasa National Park (5°13.0’N, 2°39.1 ’W), 180 m a.s.L, forest near entranee, night eolleeting,

22 Feb. 2013 (B.A. Huber); 2 $ ? in ZFMK (Ar 10497), Ankasa National Park, forest along Big Tree

Trail (~5°15.rN, 2°38.4’W), -100 m a.s.L, 23 Feb. 2013 (B.A. Huber). Eastern Region: 1 $ 2 juvs in

ZFMK (Ar 10498), Atewa Hills, Atewa Atwirebu Reserve (6°13.8’N, 0°33.5’W), 740 m a.s.L, 25 Feb.

2013 (B.A. Huber); 1 $ in pure ethanol, in ZFMK (Gha 135), same data.

21. Pholcus kindia Huber, 2011. Guinea (Huber 2011b).

Physocyclus Simon, 1893

Physocyclus is a New World genus. Only the pantropieal synanthropie P. globosus (Taezanowski, 1874)

oeeurs in West Afriea (Fig. 31).

22. Physocyclus globosus (Taezanowski, 1874). Guinea, Ivory Coast (Millot 1941); newly reeorded for

Sierra Eeone, Eiberia, Ghana, Togo, and Benin.

New records

GUINEA: Guinee Forestiere: 2 SS 3 $$ 1 juv. in ZFMK (Ar 10499), Eola, in building (7°48.5’N,

8°30.9’W), 490 m a.s.L, 30 Nov. 2008 (B.A. Huber).

SIERRA EEONE: -25 (?c? 35 ?? (5 vials) in MRAC (146489, 148459, 148462, 148541, 159145),

Freetown, Mount Aureol [8°28.7’N, 13°13.3’W], Sep. 1976-Nov. 1977 (D. Olu-Pitt).

EIBERIA: Montserrado Co.: 4 SS 6 $ $ + juvs (6 vials) in USNM, Monrovia (6°19’N, 10°48’W), Jun.

1894-Mar. 1895 (O.F. Cook); 1 ? 2 juvs in USNM, Mt. Coffee [6°31.7’N, 10°33.4’W], Apr. 1897 (O.F.

Cook). Maryland Co.: 2 $ ? + juvs in USNM, Muhlenberg Mission (4°28’N, 7°35’W), Feb. 1895 (G.P

Goff); 1 $ 1 juv. in USNM, same loeality, Jun. 1892 (O.F. Cook); 1 $ in USNM, unidentified loeality:

Eiberia, “N.Y. settlemenf’. Mar. 1895 (J.S. Sharp).

IVORY COAST: 1 $ in MRAC (201102), AdiopoDoume [5°20’N, 4°20’W], Orstom, around buildings,

12 Nov. 1994 (R. Joeque); 1 S in MRAC (177559), Appouesso, Foret elassee de laBossematie [6°35’N,

HUBER B.A. & KWAPONG R, West African pholcid spiders

3°28’W], Nov. 1993 (R Jocque); 4 33 2 $$ 1 juv. in CAS, Parc National de la Maraoue [~7°05’N,

6°05’W], 8 Jan. 1991 (W.J. Pulawski); 1 $ in MRAC (230443), Tai Forest, Centre de Recherche

Ecologique (CRE) [5°50’N, 7°21’W], no further data.

GHANA: Greater Accra Region: 2 5? 2juvs in MRAC (131433), Accra [~5°35’N, 0°12’W], 2-8 Oct.

1966 (D. Thys van den Audenaerde); 1 (? in MRAC (127241), same locality, 20 Jul. 1964 (G. Marlier);

1 $ in MRAC (142391), Eegon [5°39’N, OHO’W], 14 Mar. 1972 (J. Edmunds); 2 33 7 +juvs (6

Figs 31-32. Known distributions of the gQnQra Physocyclus Simon, 1893 md SmeringopinaKxms, 1957

in West Africa.

European Journal of Taxonomy 59: 1-44 (2013)

vials) in MCZ (34030-35), Achimota [5°37’N, 0°14’W], in and around house, Mar. 1961-Mar. 1962

(C.P. Hinckley). Western Region: \ S \ juv. in ZFMK (Ar 10500), Ankasa National Park (5°13.0’N,

2°39.rW), 180 m a.s.L, buildings near entrance, 22 Feb. 2013 (B.A. Huber). Central Region: 1 $ in

ZFMK (Ar 10501), Cape Coast (5°06.3’N, 1°14.8’W), 15 m a.s.L, in building, 23 Feb. 2013 (B.A.

Huber). Eastern Region: 5 SS 3 $ $ 1 juv. in ZFMK (Ar 10502), near Mpraeso (6°34.EN, 0°43.9’W),

370 m a.s.L, degraded forest near road, at rocks, 10 Mar. 2013 (B.A. Huber); 3 $ $ 1 juv. in pure ethanol,

in ZFMK (Gha 151), same data; 1 ? 1 juv. in CAS, Nkawkaw [6°33’N, 0°46’W], 31 Jan. 1991 (W.J.

Pulawski); \ S in MRAC (127253), Akosombo [6°16’N, 0°03’E], 12 Jul. 1964 (G. Marlier). Ashanti

Region: 4 SS 3 $ $ in ZFMK (Ar 10503), Kumawu (6°54.3’N, 1°15.4’W), 380 m a.s.L, in building, 3

Mar. 2013 (B.A. Huber); 1 $ in MRAC (131424), Kumasi [6°4rN, 1°37’W], 24-25 Sep. 1966 (D. Thys

van den Audenaerde).

TOGO: several (?(??? (3 vials) in MRAC (136122-123, 150069), Kolokope [7°28’N, 1°19’E], 26-31

Jul. 1969 (F. Puylaert); 1 $ 2 juvs in MRAC (131453), “Doteleope”, 6-7 Oct. 1966 (D. Thys van den

Audenaerde); 1 $ in MRAC (212772), Dzogbegan [7°14’N, 0°41’E], Jan. 2003 (S. Tchibozo).

BENIN: 1 S in MRAC (212766 part), Adjohoun [6°42’N, 2°30’E], in house, 23 Dec. 2002 (S. Tchibozo);

2 $ 5 1 ill MRAC (212769), Godomey [6°25’N, 2°19’E], in house, Dec. 2002 (S. Tchibozo); 2 $ $

in ZFMK (Ar 10504), Niaouli (6°44’N, 2°08’E), 20 Feb. 2009 (S. Tchibozo).

NIGERIA: Osun: \ S \ ^ \ juv. in MRAC (174602 part), Ile-Ife, Obafemi Awolowo Univ. campus

[7°28’N, 4°34’E], 1991 (H. Segers).

Quamtana Huber, 2003

Quamtana is most diverse in southern Africa, but a few species occur as far north as Cameroon, Congo

DR, Uganda (Huber 2003c), Rwanda [“Burundi” in Huber & Wami 2012 is a lapsus; Cyamudongo is in

Rwanda, 2°33.5’S, 28°59.5’E] and Kenya (Huber & Wami 2012). The new records below include the

first for West Africa, and indicate that Q. kitahurira Huber, 2003 (previously known from Uganda and

Rwanda only) is apparently a widespread species.

23. Quamtana kitahurira Huber, 2003. newly recorded for West Africa (Guinea) (see page 38).

Smeringopina Kraus, 1957

The Guineo-Congolian genus Smeringopina is highly diverse in Central Africa but only eight species

are known from West Africa (Huber 2013; Fig. 32). Most species belong to one of the two endemic West

African species groups, the guineensis group and the ankasa group (Huber 2013); only S. beninensis

Kraus, 1957 is a notable outlier of a group otherwise restricted to Central Africa.

24. Smeringopina ankasa Huber, 2013. Ghana, Ivory Coast (Huber 2013).

25. Smeringopina beninensis Kraus, 1957. Benin, Nigeria (Kraus 1957; Huber 2013).

26. Smeringopina bineti (Millot, 1941). Guinea (Millot 1941; Huber 2013).

27. Smeringopina bomfobiri Huber, 2013. Ghana (Huber 2013).

28. Smeringopina fon Huber, 2013. Benin (Huber 2013).

29. Smeringopina guineensis (Millot, 1941). Guinea, Eiberia (Millot 1941; Huber 2013).

30. Smeringopina ibadan Huber, 2013. Nigeria (Huber 2013).

HUBER B.A. & KWAPONG R, West African pholcid spiders

31. Smeringopina pulchra (Millot, 1941). Guinea(?), Ivory Coast, Ghana, Togo (Millot 1941; Huber

2013).

Smeringopus Simon, 1890

Smeringopus is currently the most species-rich African pholcid genus, but only two species occur in

West Africa (Fig. 33): the pantropical synanthropic S. pallidus (Blackwall, 1858) and the widespread

Guineo-Congolian S. cylindrogaster (Simon, 1907) which covers the entire Guineo-Congolian rainforest

(Huber 2012).

32. Smeringopus cylindrogaster (Simon, 1907). Guinea Bissau, Guinea, Liberia, Ivory Coast, Ghana

(Simon 1907; Huber 2009, 2012).

New records

GUINEA: 1 $ 1 juv. in pure ethanol, in MRAC (236933), Mt. Nimba, Gouan Forest Centre [7°42’N,

8°24’W], beating trees and shrubs with hanging litter at E5-3 m above ground, 1250 m a.s.L, 8 Oct.

2011 (D. Van den Spiegel, A. Henrard).

IVORY COAST: 1 5 in MRAC (230346 part), Tai Forest, Centre de Recherche Ecologique (CRE)

[5°50’N, 7°21 ’W], beating in forest near Chimpanzee Camp, 22 Feb. 2010 (R. Jocque, M. Diarassouba);

1 5 in MRAC (230146 part), same locality, at foot of trees, by hand, 22 Feb. 2010 (R. Jocque); 1 $

1 juv. in MRAC (230281), same data but forest near river, 22 Feb. 2010 (R. Jocque, M. Diarassouba);

1 5 in MRAC (230244), same locality, forest E of camp, “layon Gerard”, inundated forest, sieved litter,

20 Feb. 2010 (R. Jocque, L. Oulai); 2 in MRAC (230296), same locality, forest across river, forest

on clayey soil, beating, 20 Feb. 2010 (R. Jocque, M. Diarassouba); 1 $ in MRAC (233643), Tai Forest,

beating, 1 Sep. 2010 (D. Van den Spiegel, A. Kablan); 2 juvs in MRAC (230360 part), Adiopo Doume

[5°20’N, 4°20’W], Centre Suisse de Recherche Scientifique (CSRS), small dry forest, sieving litter, 26

Feb. 2010 (D. van den Spiegel, R. Jocque).

GHANA: Central Region: 1 $ in pure ethanol, in ZFMK (Gha 149), Kakum National Park (5°20.9’N,

1°23.0’W), 160 m a.s.L, forest near entrance, day collecting, 19 Feb. 2013 (B.A. Huber); 1 (J 1 $ 1 juv.

in MRAC (217296 part), Kakum Forest, beating in secondary forest, 23 Nov. 2005 (R. Jocque, D. de

Bakker, L. Baert). Eastern Region: 3 $ $ in ZFMK (Ar 10505), Atewa Hills, Atewa Atwirebu Reserve

(6°13.8’N, 0°33.5’W), 740 m a.s.L, 25 Feb. 2013 (B.A. Huber); 1 ? in ZFMK (Ar 10506), near Mpraeso

(6°34.UN, 0°43.9’W), 370 m a.s.L, degraded forest near road, 10 Mar. 2013 (B.A. Huber). Western

Region: 1 S in ZFMK (Ar 10507), Ankasa National Park (5° 13.0’N, 2°39.UW), 180m a.s.L, forest near

entrance, day collecting, 22 Feb. 2013 (B.A. Huber); 3 SS 5 $$ 1 juv. (2 vials) in ZFMK (Ar 10508-

09), same data but night collecting; 1 c? 1 juv. in pure ethanol, in ZFMK (Gha 157), same data.

33. Smeringopus pallidus (Blackwall, 1858). Guinea, Ivory Coast (Millot 1941); newly recorded for

Senegal, The Gambia, Sierra Leone, and Ghana.

New records

SENEGAL: 1 (? in MRAC (161825), Dakar [14°46’N, 17°15’W], in house, 14 Nov. 1983 (E. Tybaert);

4 SS 4 $ $ (4 vials) in AMNH, same locality, May-Jul. 1945, no further data.

THE GAMBIA: 4 (?(? 15 ? ? in ZFMK (Ar 5242), Bijilo Forest Park (13°26.3’N, 16°43.5’W), coastal

forest, 22 Oct. 2008 (Yu.M. Marusik); 1 (J 5 $ $ in ZFMK (Ar 5244), same data, litter, mostly under

palms; 1 6^ 1 juv. in CAS, Baccau [13°29’N, 16°40’W], 22 Oct. 1981 (W. Settle); 2 (?(? 3 ? ? in SMF,

same locality, in building, no date (J. Wunderlich).

GUINEA: 1 (5^ 3 $ $ (2 vials) in pure ethanol, in ZFMK (Gui 84, 102), Conakry, in building (9°34.UN,

13°39.7’W), 40 m a.s.L, 18 Nov. 2008 (B.A. Huber). Moyenne-Guinee: 1 $ in ZFMK (Ar 10510),

Dalaba (10°41.4’N, 12°15.7’W), in building, 1260 m a.s.L, 22 Nov. 2008 (B.A. Huber). Moyenne-

Guinee: 2 SS 6 $$ in pure ethanol, in ZFMK (Gui 85), Doucki, in and at buildings (10°59.UN,

European Journal of Taxonomy 59: 1-44 (2013)

12°36.0’W), 1050 m a.s.L, 23 Nov. 2008 (B.A. Huber). Basse-Guinee: \ S \ juv. in ZFMK (Ar 10511),

Kindia, in building (10°03’N, 12°5rW), 400 m a.s.L, 20 Nov. 2008 (B.A. Huber).

SIERRALEONE: 10 SS 6 $ $ +juvs (3 vials) inMRAC (148460 part, 159113,159145 part), Freetown,

Mount Aureol [8°28.7’N, 13°13.3’W], Sep. 1976-Jun. 1977 (D. Olu-Pitt); 5 9 ?? (4 vials) in

MRAC (146506, 148525, 530, 542), Freetown, Dee. 1974-Jan. 1977 (D. Olu-Pitt).

Figs 33-34. Known distributions of the genera Smeringopus Simon, 1890 and Spermophora Hentz, 1841

in West Afriea.

HUBER B.A. & KWAPONG R, West African pholcid spiders

IVORY COAST: 1 (? 3 ? ? 3 juvs (3 vials) in MRAC (131436-438), Toulepleu [6°35’N, 8°25’W], 28

Jul. 1966 (W. Verheyen, D. Thys van den Audenaerde).

GHANA: Ashanti Region: 1 juv. in ZFMK (Ar 10512), Kumawu (6°54.3’N, 1°15.4’W), 380 m a.s.L, in

building, 3 Mar. 2013 (B.A. Huber). Greater Accra Region: 1 (? in MRAC (142403), Legon [5°39’N,

0°10’W], 14 Mar. 1972 (J. Edmunds). Eastern Region: 1 $ in MRAC (127252), Akosombo [6°16’N,

0°03’E], 12 Jul. 1964 (G. Marlier).

SpermophoraYlQntz, 1841

Spermophora is diverse in sub-Saharan Africa but relationships among African taxa and between

African and Asian taxa remain unclear (Huber 2003b, 2003c, 2005; Dimitrov et al. 2013). Five species

are known from West Africa (Fig. 34). Of these, S. tonkoui Huber, 2003 and S. ziama sp. nov. share

several derived characters but their relationship to other conspecifics remain dubious (see under S. ziama

sp. nov. description below). Another pair of putatively close relatives is S. dieke Huber, 2009 and S.

akwamu sp. nov. (see under S. akwamu sp. nov. description below). Avery similar undescribed species

occurs in Congo DR (in MRAC). Finally, S. kyambura is a widespread Guineo-Congolian species but

West African specimens were assigned tentatively (Huber & Wami 2012).

34. Spermophora akwamu sp. nov. (see page 31). Ghana.

35. Spermophora dieke Huber, 2009. Guinea (Huber 2009).

36. Spermophora kyambura Huber & Wami, 2012. Ghana (Huber & Wami 2012).

37. Spermophora tonkoui Huber, 2003. Ivory Coast (Huber 2003b). Newly recorded for Guinea.

New record

GUINEA: 2 SS 5 $ $ in pure ethanol, in MRAC (236928), Mt. Nimba, Foret de Zie, near Gouan camp

or “Station de Pompage Zie” [7°40’N, 8°26’W], 1250 m a.s.L, beating trees and shmbs with hanging

litter, at 1.5-3m above ground, 11 Oct. 2011 (D. Van den Spiegel, A. Hemard); 2 SS 6 $$ 2 juvs (3

vials) in SMF, Mt. Nimba, Crete de Nion, 1100-1160 m a.s.L, 11 Feb.-12 May 1957 (M. Famotte).

38. Spermophora ziama sp. nov. (see page 34). Guinea.

Identification key to West African pholcid genera

References to figures in upper case (“Fig.”) refer to illustrations herein; figures in lower case (“fig.”)

have been published previously and are freely accessible online (at http://www.uni-bonn.de/~bhuberl/l .

1. Abdomen worm-shaped (>10 x longer than wide; Figs 4-5); male chelicerae with one pair of small

projections in latero-distal position (Fig. 74). Leptopholcus Simon, 1893

- Abdomen not worm-shaped (<10 x as long as wide); male chelicerae different.2

2. Six eyes (AME absent).3

- Eight eyes.6

3. Sternum with characteristic pattern of radiating lines (Fig. 40); abdomen dorsally with dark heart-

mark (Fig. 35). Anansus Huber, 2007

- Sternum without pattern; abdomen dorsal pattern different.4

European Journal of Taxonomy 59: 1-44 (2013)

4. Carapace with 2-3 pairs of dark lateral spots; oeular area (espeeially in males) strongly elevated,

in male with frontal hairy poeket; male ehelieerae without proximal lateral projeetions (figs 2-4 in

Huber 1996). Modisimus culicinus (Simon, 1893)

- Carapaee without lateral spots; oeular area not elevated; male ehelieerae with proximal lateral

projeetions.5

5. Proeursus with highly eomplex system of transparent lamellae (fig. 14 in Huber 2007); male

ehelieerae narrowing distally, with pair of simple frontal apophyses elose to median line (fig. 15 in

Huber 2007). Nyikoa limbe Huber, 2007

- Proeursus and male ehelieerae different. Spermophora Hentz, 1841

6. Carapaee with median indentation (furrow or pit).7

- Carapaee evenly domed, without median indentation.11

7. Abdomen globular, oval, or higher than long.8

- Abdomen elongated.9

8. Proeursus with prominent distal spine; female earapaee with posterior median eone aeting against

frontal plate on abdomen; epigynum with median anterior proeess.

. Physocyclus globosus (Taezanowski, 1874)

- Proeursus short, without distal spine (Figs 57-58); female earapaee without posterior eone; epigynum

without median anterior proeess (Fig. 54). Artema Walekenaer, 1837

9. Legs with many small blaek marks; abdomen angular in lateral view (posteriorly high; Fig. 16).

. Crossopriza Simon, 1893

- Legs without small blaek marks, abdomen not angular in lateral view (posteriorly tapering).10

10. Male ehelieerae with proximal lateral projeetions, either without or with several modified hairs on

eaeh side. Smeringopina Kraus, 1957

- Male ehelieerae without proximal lateral projeetions, with single modified hair on eaeh distal

apophysis (fig. 47 in Huber 2009). Smeringopus Simon, 1890

11. Abdomen globular or oval.12

- Abdomen elongated.13

12. Proeursus with long hinged dorsal proeess, epigynum weakly selerotized, internal U-shaped strueture

visible through eutiele anteriorly (figs 83-89 in Huber 2011b).

. Micropholcus fauroti (Simon, 1887)

- Proeursus widely eurved, without long dorsal proeess (figs 204-205 in Huber 2003e); epigynum

weakly selerotized, no internal strueture visible through eutiele (Fig. 107).

. Quamtana kitahurira Huber, 2003

13. Abdomen drawn into eone dorso-posteriorly (Fig. 8; fig. 29 in Huber 2011b); male ehelieerae with

pair of lateral unselerotized projeetions in distal position (fig. 130 in Huber 2009); epigynum weakly

selerotized. Pehrforsskalia conopyga Deeleman-Reinhold & van Harten, 2001

- Abdomen rounded posteriorly; male ehelieerae with pair of lateral unselerotized projeetions in

proximal position; epigynum strongly selerotized. Pholcus Walekenaer, 1805

HUBER B.A. & KWAPONG R, West African pholcid spiders

Taxonomy

Anansus atewa sp. nov.

um:lsid:zoobank.org:act:9EC8FF7E-F56B-4FAD-9B48-73QCE2795474

Figs 9-10, 23, 35-47

Diagnosis

Easily distinguished from known congeners by distinct pair of projections on male clypeus (Figs 35,

39; other known species with single tiny median process); also by male cheliceral apophyses (more

proximal, wider apart, and more pointed than in other known species; Figs 39,45), by shape of procursus

(large ventral curved sclerite; without retrolateral sclerite provided with brush of long hairs; without

ventral pointed projection; Figs 38, 44), and by more anterior position of epigynal pockets (Figs 41, 46).

Etymology

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

Type material

Holotype 3, in ZFMK (Ar 10513).

Type data

GHANA: Eastern Region: Atewa Hills, Atewa Atwirebu Reserve (6°13.8’N, 0°33.5’W), 740 m a.s.L,

25 Feb. 2013 (B.A. Huber).

Other material examined

GHANA: Eastern Region: 4 $ $ together with holotype; 2 33 2 $ $ in ZFMK (Ar 10514), same data,

specimens taken alive and died from heat; 2 $ $ 5 juvs in pure ethanol in ZFMK (Gha 137), same data.

Description

Male (holotype)

Measurements. Total body length 1.5, carapace width 0.6. Feg 1: 5.2 (1.3 + 0.2 + 1.4 + 1.6 + 0.7),

tibia 2: 0.9, tibia 3: 0.7, tibia 4: 1.1; tibia 1 F/d: 22. Distance PME-PME 25 pm, diameter PME 80 pm,

distance PME-AFE 20 pm, no AME.

CoEOR. Carapace ochre-grey, slightly darker medially, sternum with four pairs of dark lines extending

from behind labium to bases of leg coxae, legs pale ochre-yellow, without rings, abdomen ochre-gray

with distinct dark median dorsal mark.

Body. Habitus as in Figs 35-36; ocular area barely elevated; carapace without median furrow; clypeus

with distinct pair of projections at rim (Fig. 35); sternum wider than long (0.40/0.35), unmodified.

Chelicerae as in Fig. 45, with pair of lateral processes proximally and pointed frontal apophyses, without

stridulatory ridges.

Paeps. As in Figs 37-38 and 43-44, coxa unmodified, trochanter with indistinct ventral projection, femur

small relative to tibia, with small retrolatero-ventral process and larger prolatero-ventral process set

with tubercles, procursus complex, entire distal part apparently hinged against basis, without elements

distinctive for other known species (retrolateral sclerite provided with brush of long hairs; ventral

pointed projection), bulb with single weakly sclerotized process (embolus).

European Journal of Taxonomy 59: 1-44 (2013)

Legs. Without spines and curved hairs, few vertical hairs; retrolateral trichobothrium on tibia 1 at 25%;

prolateral trichobothrium absent on tibia 1, present on other tibiae; tarsus 1 with about ten indistinct

pseudosegments.

Variation. Tibia 1 in 2 other males: 1.3, 1.4.

Female

In general similar to male; tibia 1 in 6 females: 1.2-1.4 (mean 1.3). Epigynum consisting of large anterior

plate provided with pair of pockets and narrow posterior plate (Figs 40, 41, 46); internal genitalia as in

Figs 42 and 47.

Natural history

The spiders were found in the leaf-litter under leaves that are curved in a way to produce protected space

on the underside. Two eggsacs contained 13 and 15 eggs respectively.

Figs 35-42. Anansus atewa sp. nov. 35-36. dorsal and lateral views. 37-38. Feft f palp, prolateral

and retrolateral views. 39. f prosoma and right palp, oblique frontal view. 40. $, ventral view. 41-

42. Cleared $ genitalia, ventral and dorsal views. Scale lines: 35-36, 40 = 1 mm; 37-39, 41-42 =

0.5 mm.

HUBER B.A. & KWAPONG R, West African pholcid spiders

Figs 43-41 . Anansus atewa sp. nov. 43-44. Left 3 palp, prolateral andretrolateral views. 45. 3 chelicerae,

frontal view. 46-47. Cleared $ genitalia, ventral and dorsal views. Scale lines: 43-44, 46-47 = 0.3 mm;

45 = 0.2 mm.

European Journal of Taxonomy 59: 1-44 (2013)

Distribution

Known from type locality only (Fig. 23). Further juvenile specimens from near Mpraeso and from

Tagbo Falls (Ghana; in ZFMK) might also belong to this species.

Artema bunkpurugu sp. nov.

um:lsid:zoobank.org:act:Q5986623-C648-495E-AA88-C708FEFlA926

Figs 14-15, 24, 48-62

'Artema mauriciand" (misidentification) - Millot 1941: 3-5, figs lA-E, G-1 (fig. IF is the true A.

mauriciana = A. atlanta).

Diagnosis

Distinguished from the widespread A. atlanta by shape of procursus (arrows in Figs 57-58), bulbal

processes (arrow in Fig. 58), relatively longer male palpal tibia, less prominent male cheliceral

projections, more straight posterior epigynal margin (Figs 54, 61), and more prominent anterior epigynal

projections (Fig. 49). Middle East and Central Asian species have a very different epigynum (cf fig. 10

in Spassky 1934).

Etymology

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

Type material

Holotype S, in ZFMK (Ar 10515).

Type data

GHANA: Northern Region: Bunkpurugu (10°31.EN, 0°05.5’E), 230 m a.s.L, in building, 7 Mar. 2013

(B.A. Huber).

Other material examined

GHANA: Northern Region: 1 $ together with holotype; 2 $ $ 1 juv. in pure ethanol in ZFMK (Gha

126), same data; \ S 2 in ZFMK (Ar 10516), between Bunkpurugu and Nakpanduri (10°32.9’N,

0°00.2’E), 270 m a.s.L, under concrete bridge, 8 Mar. 2013 (B.A. Huber); 1 $ in pure ethanol in

ZFMK (Gha 128), same data; 1 ? in ZFMK (Ar 10517), near Bimbaga (10°34.5’N, 0°03.8’W), 330 m

a.s.L, under concrete bridge, 8 Mar. 2013 (B.A. Huber); 2 5? in ZFMK (Ar 10518), near Nakpanduri

(10°35.0’N, 0°06.4’W), 345 m a.s.L, under concrete bridge, 8 Mar. 2013 (B.A. Huber); 1 5 2 juvs in

pure ethanol in ZFMK (Gha 140), between Gambaga and Nakpanduri (10°34.0’N, 0°17.5’W), 400 m

a.s.L, under large rocks, 8 Mar. 2013 (B.A. Huber).

TOGO: 3 (?(? 2 6 juvs inMRAC (136024), Namoundjoga [10°53.4’N, 0°23.4’E], 27/29 Jul. 1969

(F. Puylaert).

BURKINA FASO: Sud-Ouest: 1 S inMRAC (128076), Ouagadougou [12°22’N, 1°32’W], Apr.-May

1965 (B. Roman); 1 $ in MNHN, same locality, Sep. 1937 (J. Millot); 3 SS 9 $ $ 13 juvs (2 vials) in

MNHN, Batie [9°52’N, 2°55’W], Sep. 1937 (J. Millot).

MAEl: 1 $ 3 juvs (?, abdomens missing in two specimens) in MNHN, Bamako [12°38’N, 8°00’W],

Oct. 1937 (J. Millot); 1 ? in AMNH, Gao [16°16’N, 0°03’W], 23-25 Nov. 1948 (B. Malkin).

GUINEA: \ S {\ palp and chelicerae missing), 2 $ $ (1 epigynum missing), 2 juvs in MNHN, Kouroussa

[10°39’N, 9°53’W], Oct. 1937 (J. Millot).

HUBER B.A. & KWAPONG R, West African pholcid spiders

Description

Male (holotype)

Measurements. Total body length 9.0, carapace width 4.2. Leg 1: 77.1 (20.3 + 1.9 + 21.1 +29.1 + 4.7),

tibia 2: 16.3, tibia 3: 12.7, tibia 4: 16.1; tibia 1 L/d: 54. Distance PME-PME 230 pm, diameter PME

220 pm, distance PME-ALE 115 pm, distance AME-AME 60 pm, diameter AME 210 pm.

Figs Artema bunkpurugu sp. nov. 48. (J, dorsal view. 49. Female genitalia and spinnerets, lateral

view. 50-52. Left S palp, prolateral, dorsal, and retrolateral views. 53. $ abdomen, ventral view. 54-

55. Cleared $ genitalia, ventral and dorsal views. Scale lines: 48 = 2 mm; 49-55 = 1 mm.

European Journal of Taxonomy 59: 1-44 (2013)

Color. Carapace pale oehre-grey with light brown median band and small brown spots laterally, elypeus

with pair of narrow light brown bands below AME, legs pale oehre-gray to light brown, with dark rings on

femora subdistally, patellae + tibiae proximally, and tibiae subdistally, tips of femora and tibiae whitish,

sternum pale gray with narrow dark brown margins, abdomen pale gray with large dark marks dorsally.

Body. Habitus as in Fig. 48; oeular area slightly elevated; earapaee with large median pit and distinet

median furrow posteriorly; elypeus unmodified; sternum wider than long (2.5/2.0), unmodified.

Chelieerae as in Figs 59-60, with heavily selerotized frontal projeetions provided with about ten modified

(eone-shaped) hairs eaeh, without stridulatory ridges. Gonopore with five epiandrous spigots.

Palps. As in Figs 50-52 and 56-58, eoxa unmodified, troehanter with short ventral projeetion, femur with

distinet retrolateral proeess proximally, large ventral membranous area proximally bordered on both

sides by heavily selerotized ridges, and small dorsal projeetion proximally; femur-patella hinges elose

together dorsally; patella very short; proeursus with proximal dorsal apophysis and weakly developed

ventral poeket, with distinetive ventral projeetion distally (arrow in Fig. 57), bulb with two apophyses

fianking membranous proeess (putative embolus).

Legs. Without spines, with some very weakly eurved hairs on tibiae and metatarsi, few vertieal hairs

mainly on distal leg segments; retrolateral triehobothrium on tibia 1 at 5.5%; prolateral triehobothrium

present on all tibiae (also tibia 1); pseudosegmentation not visible.

Variation. Tibia 1 in 8 other males: 15.5-23.1 (mean 19.8). Lateral spots on earapaee sometimes fused

to three pairs of large marks. Sternum posteriorly with variably distinet pattern of light brown marks.

Gonopore with variable number of epiandrous spigots (4-6).

Female

In general similar to male but ehelieerae with distinet selerotized patehes laterally provided with

stidulatory files (ef fig. IG in Millot 1941); tibia 1 in 20 females: 12.0-18.9 (mean 16.1). Lpigynum

eonsisting of trapezoidal anterior plate and simple posterior plate, with pair of large distinetive projeetions

in front of epigynum (Figs 49, 53-54, 61); internal genitalia as in Figs 55, 62.

Natural history

The spiders were found both in natural habitats (among large roeks) and in human-made habitats (under

low eonerete bridges, in buildings; see also Millot 1941). When disturbed, the spiders try to hide in

ereviees, running short distanees and then vibrating at low amplitude but high frequeney.

Distribution

Widely distributed in Sudanian West Afriea (Fig. 24).

Note

Millot (1941) was aware of the faet that most of his speeimens from West Afriea did not agree well

with published figures of A. mauriciana (=A. atlanta). He tentatively interpreted this as intraspeeifie

variation. However, eomparison of numerous speeimens of this new speeies with speeimens of A. atlanta

from all over the World shows that the two speeies differ in several eharaeters (see above) and that no

intermediate speeimens seem to oeeur.

Millot’s (1941) speeimens from Kindia, Guinea (2 juvs; $ missing), Ferkessedougou, Ivory Coast

(1 juv.), and Gao, Mali (2 juvs) are in MNHN but juveniles of A. atlanta andH. bunkpurugu sp. nov. are

indistinguishable so their identity is unelear.

HUBER B.A. & KWAPONG R, West African pholcid spiders

Figs 56-62. Artema bunkpurugu sp. nov. 56-57. Left 3 palp, prolateral and retrolateral views. 58. Left

procursus and genital bulb, dorsal view (e: embolus). 59-60. (S chelicerae, frontal and lateral views. 61-

62. Cleared $ genitalia, ventral and dorsal views. Arrows point at distinctive structures (as compared

withy4. atlanta). Scale lines: 1 mm.

European Journal of Taxonomy 59: 1-44 (2013)

Leptopholcus kintampo sp. nov.

um:lsid:zoobank.org:act:AB6C8FC0-A0FB-4EFA-82A5-A622435F19CF

Figs 4, 26, 63-76

Diagnosis

Distinguished from similar close relatives (species with median process on male clypeus: L. signifer

Simon, 1893; L. debakkeri Huber, 2011; L. guineensis; L. dschang Huber 2011; L. gracilis Berland,

1920; L. budongo Huber, 2011) by shapes of bulbal processes (Figs 68, 72).

Etymology

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

Type material

Holotype S, in ZFMK (Ar 10519).

Figs 63-71. Leptopholcus kintampo sp. nov. 63-64. S, dorsal and lateral views. 65-66. S prosoma,

oblique frontal and dorsal views. 67. Epigynum, ventral view. 68-69. Eeft S palp, prolateral and

retrolateral views. 70-71. Cleared $ genitalia, ventral and dorsal views. Scale lines: 63-64 = 2 mm;

66 = 1 mm; 65, 67-71 = 0.5 mm.

HUBER B.A. & KWAPONG R, West African pholcid spiders

Type data

GHANA: Brong-Ahafo Region: Kintampo Falls (8°05.3’N, 1°41.9’W), 280 m a.s.L, degraded forest

along stream, 5 Mar. 2013 (B.A. Huber).

Other material examined

GHANA: Brong-Ahafo Region: 1 (? 3 $$ 2 juvs in ZFMK (Ar 10520), same data as holotype; 2 SS

2 $ $ 2 juvs in ZFMK (Ar 10521), same data but 9 Mar. 2013; 2 $ $ 2 juvs in pure ethanol in ZFMK

(Gha 161), same data as holotype.

Description

Male (holotype)

Measurements. Total body length 7.5, carapace width 1.1. Feg 1: 33.0 (8.3 + 0.5 + 7.9 + 13.0 + 3.3),

tibia 2: 5.9, tibia 3:3.8, tibia 4: 6.0; tibia 1 F/d: 89. Distance PME-PME 345 pm, diameter PME 95 pm,

distance PME-AFE 25 pm, diameter AME 25 pm, distance AME-AME 25 pm.

CoEOR. Carapace pale ochre-yellow with large triangular brown mark (Fig. 66), ocular area and clypeus

also dark, sternum pale gray with dark margins and small dark spots, legs pale ochre-yellow with dark rings

in patella area and at tibia-metatarsus joints, abdomen ochre-yellow with numerous black dorsal marks.

Body. Habitus as in Figs 63-64; ocular area slightly elevated, each triad on short stalk directed laterally

(Fig. 65); carapace without median furrow; clypeus with distinct median process about 160 pm long;

sternum wider than long (0.65/0.55), unmodified. Chelicerae as in Fig. 74, with pair of tiny lateral

processes in rather distal position, without stridulatory ridges.

Paeps. As in Figs 68-69 and 72-73, coxa unmodified, trochanter with long ventral apophysis and short

projection at retrolateral trochanter-femur joint, femur with finger-shaped retrolateral process and large

ventral bulge distally, procursus with two membranous ventral processes, one arising from ventral ‘knee’,

the other more distally, bulb with large uncus with fiat curved process and bifid appendix, embolus

membranous with many fringes distally.

Fegs. Without spines and curved hairs, few vertical hairs; retrolateral trichobothrium on tibia 1 at

3.5%; prolateral trichobothrium absent on tibia 1, present on other tibiae; tarsus 1 with >30 indistinct

pseudosegments.

Variation. Tibia 1 in 3 other males: 7.5, 7.8, 8.0. Sternum variably dark.

Female

In general similar to male but only weak V-mark on carapace, ocular area and clypeus not darkened,

triads not on stalks and closer together (distance PME-PME 240 pm), clypeus unmodified, abdominal

marks indistinct. Tibia 1 in 5 females: 6.2-7.1 (mean 6.8). Epigynum weakly sclerotized, anterior plate

simple with median incision posteriorly (Figs 67, 70, 75); internal genitalia as in Figs 71, 76.

Natural history

The spiders were found on the undersides of large leaves, with their bodies fiat against the leaf

Distribution

Known from type locality only (Fig. 26).

European Journal of Taxonomy 59: 1-44 (2013)

Figs 72-76. Leptopholcus kintampo sp. nov. 72-73. Left S palp, prolateral and retrolateral views. 74.

S ehelieerae, frontal view. 75-76. Cleared $ genitalia, ventral and dorsal views. Seale lines: 72-73 =

0.5 mm; 74-76 = 0.3 mm.

HUBER B.A. & KWAPONG R, West African pholcid spiders

Spermophora akwamu sp. nov.

um:lsid:zoobank.org:act:D0DB384E-4A3F-4EBl-80D6-48278C31EB82

Figs 11-13, 34, 77-91

Diagnosis

Distinguished from the very similar S. dieke and from a similar undescribed species from Congo DR

(in MRAC) by pair of processes on male clypeus (Fig. 80; only one median process in S. dieke\ bifid

process in species from Congo DR); from other congeners by armature of male chelicerae (only one

pair of weakly sclerotized proximal processes; Fig. 89), by shapes of procursus (ventral sclerotized fiap,

distal fiagellum; Figs 87-88) and bulb (shape of hooked apophysis; Fig. 87), and by shape of epigynum

(rectangular plate without pockets; Figs 85, 90).

Etymology

The name is a noun in apposition, derived from the Akwamu (or Akuambo), an Akan kingdom in the 17^*’

and 18* centuries whose founders settled in the area of modern-day Kakum National Park.

Type material

Holotype S, in ZFMK (Ar 10522).

Type data

GHANA: Central Region: Kakum National Park (5°20.9’N, 1°23.0’W), IbOma.s.L, forest near entrance,

day collecting, 19 Feb. 2013 (B.A. Huber).

Other material examined

GHANA: Central Region: 15 SS 25 $$ 8 juvs in ZFMK (Ar 10523-24), Kakum National Park, same

data as holotype; 7 SS 7 ? ? 2 juvs in ZFMK (Ar 10525), same data but 20 Feb. 2013; 1 8 $ $ 2 juvs

in pure ethanol in ZFMK (Gha 145), same data; 6 SS 6 $ $ in ZFMK (Ar 10526-27), same data but

night collecting, 20 Feb. 2013; 11 SS 28 $$ in MRAC (217694, 698, 712, 725, 737), Kakum Forest,

fogging in primary forest, 16-25 Nov. 2005 (R. Jocque, D. De Bakker, F. Baert); 8 SS 20 $ $ in MRAC

(217687, 701, 703, 708), same data but secondary forest, 12-19 Nov. 2005; 1 juv. in MRAC (217292),

same data but beating in secondary forest, 23 Nov. 2005; 3 SS 2 5? in MRAC (217261, 277), same

data but beating between primary and secondary forest, 11-12 Nov. 2005; 1 (? 2 5 ? 1 juv. in MRAC

(217173), same data but sieving of forest litter, 10 Nov. 2005. Eastern Region: 2 SS 1 ? 1 juv. in ZFMK

(Ar 10528), Atewa Hills, Atewa Atwirebu Reserve at 500 m a.s.l. (6°13.8’N, 0°32.4’W), degraded forest

along road, 24 Feb. 2013 (B.A. Huber); 1 $ 5 juvs in ZFMK (Ar 10529), Atewa Atwirebu Reserve at

740 m a.s.l. (6°13.8’N, 0°33.5’W), 25 Feb. 2013 (B.A. Huber); 1 juv. in pure ethanol in ZFMK (Gha

139), same data. Western Region: 5 SS 4 5? 2 juvs in ZFMK (Ar 10530), Ankasa National Park

(5°13.0’N, 2°39.1’W), 180 m a.s.l., forest near entrance, day collecting, 22 Feb. 2013 (B.A. Huber);

2 SS 2 juvs in pure ethanol in ZFMK (Gha 159), same data; 1 (5^ 2 $ ? in ZFMK (Ar 10531), Ankasa

National Park, forest along Big Tree Trail (~5°15.UN, 2°38.4’W), -100 m a.s.l., 23 Feb. 2013 (B.A.

Huber).

Description

Male (holotype)

Measurements. Total body length 2.1, carapace width 0.7. Feg 1: 22.3 (5.3 + 0.4 + 5.4 + 9.2 + 2.0),

tibia 2: 3.1, tibia 3: 2.0, tibia 4: 3.0; tibia 1 F/d: 76. Distance PME-PME 115 pm, diameter PME 95 pm,

distance PME-ALE 35 pm, no AME.

European Journal of Taxonomy 59: 1-44 (2013)

Figs 77-86. Spermophora akwamu sp. nov. 77-79. S, dorsal, lateral, and ventral views. 80. S prosoma,

oblique frontal view. 81-82. $, ventral and dorsal views. 83-84. Left S palp, prolateral and retrolateral

views. 85-86. Cleared $ genitalia, ventral and dorsal views. Seale lines: 77-79, 81-82 = 1 mm; 80, 83-

84 = 0.5 mm; 85-86 = 0.2 mm.

HUBER B.A. & KWAPONG R, West African pholcid spiders

Figs 87-91. Spermophora akwamu sp. nov. 87-88. Left S palp, prolateral and retrolateral views (hp:

hinged process). 89. S chelicerae, frontal view. 90-91. Cleared $ genitalia, ventral and dorsal views.

Scale lines: 87-88 = 0.5 mm; 90-91 = 0.3 mm; 89 = 0.2 mm.

European Journal of Taxonomy 59: 1-44 (2013)

Color. Carapace pale gray with narrow blaek margins and indistinet median line (Fig. 77), oeular

area and elypeus dark brown to blaek, sternum pale gray (Fig. 79), legs oehre-yellow, with indistinet

dark rings subdistally on femora and tibiae, abdomen mostly pale gray, with blaek transversal mark at

posterior tip, further blaek marks posteriorly above spinnerets and near gonopore.

Body. Habitus as in Figs 77-79; oeular area slightly elevated, eaeh triad on low hump; earapaee without

median furrow (only dark line); elypeus with distinetive pair of dark proeesses, about 100 pm long

(Fig. 80); sternum wider than long (0.55/0.45), unmodified. Chelieerae as in Fig. 89, with pair of weakly

selerotized proeesses proximally, distally without modifieation, without stridulatory ridges.

Palps. As in Figs 83-84 and 87-88, eoxa with indistinet ventral proeess, troehanter with long ventral and

shorter retrolateral apophyses, proeursus with ventral selerotized fiap, prolatero-dorsal hinged proeess

and thin distal fiagellum; bulb with hooked apophysis and long weakly selerotized embolus.

Legs. Without spines and eurved hairs, few vertieal hairs; retrolateral triehobothrium on tibia 1 at

8.5%; prolateral triehobothrium absent on tibia 1, present on other tibiae; tarsus 1 with >20 indistinet

pseudosegments.

Variation. Tibia 1 in 35 other males: 4.3-5.6 (mean 4.8). Some males from type loeality with additional

small blaek marks dorsally on abdomen; in both males from Atewa Hills, these marks are present and

larger; in males from Ankasa N.P. the abdomen is almost monoehromous, without ventral marks, without

marks above spinnerets, only mark at posterior tip.

Female

In general similar to male but sternum in most females dark brown or blaek (Fig. 81), abdomen dorsally

with more and larger blaek marks (Fig. 82), elypeus unmodified, earapaee with small median proeess

aeting against indistinet selerotized plate above pedieel. Tibia 1 in 42 females: 3.5-4.4 (mean 3.9).

Epigynum simple reetangular plate, anteriorly with dark transversal internal strueture, without poekets

(Figs 85, 90); internal genitalia as in Figs 86, 91.

Relationships

Judging from the very similar morphology, this speeies is probably elosely related to S. dieke from

Guinea (ef figures in Huber 2009) as well as to a further very similar (undeseribed) speeies from Congo

DR (in MRAC).

Natural history

The spiders were found both under green leaves where they built slightly domed sheets that extended

beyond the leaves (Fig. 13; with the spider resting under a leaf), and in the leaf litter, under eurved dead

leaves, in small webs apparently not extending beyond the leaf

Distribution

Known from several loealities in southern Ghana (Fig. 34).

Spermophora ziama sp. nov.

um:lsid:zoobank.org:aet:0ACE7F4F-6D69-4F46-AlD3-ED54AAE3846D

Figs 34, 92-105

Spermophora ef tonkoui - Dimitrov et al. 2013 (DNA data).

HUBER B.A. & KWAPONG R, West African pholcid spiders

Diagnosis

Distinguished from S. tonkoui (the only similar species and probably closest known relative; see below)

by relatively shorter palpal elements (femur, tibia, bulb; Figs 100-101), by large hooked apophysis

on bulb (Fig. 100; in S. tonkoui pair of small apophyses distally on long bulb), by shorter proximal

frontal apophyses on male chelicerae (Fig. 103), and by pair of small pockets in posterior epigynal plate

(Fig. 104).

Etymology

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

Type material

Holotype (?, in ZFMK (Ar 10532).

Type data

GUINEA: Guinee Forestiere: Foret Classee de Ziama (8°24.2’N, 9°19.3’W), 640 m a.s.L, 2 Dec. 2008

(B.A. Huber).

Other material examined

GUINEA: Guinee Forestiere: I (J 3 $ $ in ZFMK (Ar 10533), same data as holotype; 1 (J 1 $ in ZFMK

(Ar 10534), same data, taken alive and died from heat; 4 $ $ 1 juv. in pure ethanol in ZFMK (Gui 67),

same data.

Description

Male (holotype)

Measurements. Total body length 2.0, carapace width 0.9. Feg 1: 19.3 (4.6 + 0.4 + 5.0 + 7.1 + 2.2),

tibia 2: 3.0, tibia 3: 2.1, tibia 4: 2.8; tibia 1 F/d: 57. Distance PME-PME 345 pm, diameter PME 70 pm,

distance PME-ALE 35 pm, no AME.

CoEOR. Carapace and clypeus mostly light brown except lighter median band, sternum dark brown, legs

ochre-yellow, abdomen mostly dark except dorsal median mark and book-lung covers pale gray.

Body. Habitus as in Figs 92-93; ocular area barely elevated, each triad on short stalk directed laterally,

with pair of elongate furrows behind each stalk; carapace without median furrow; clypeus unmodified;

sternum wider than long (0.65/0.50), unmodified. Chelicerae as in Figs 102-103, with pair of lateral

processes proximally, pair of frontal apophyses proximally close together, and pair of frontal apophyses

distally provided with five tiny modified (conical) hairs each; without stridulatory ridges.

Paeps. As in Figs 94-95 and 100-101, coxa unmodified, trochanter with retrolateral apophysis curved

towards dorsally, femur with retrolatero-dorsal process proximally, widened distally, tibia very long,

dorso-distal rim slightly projecting, procursus complex, apparently with two hinges in proximal part,

with long membranous process arising from hinge retrolaterally, distal whitish part provided with many

short pseudotrichia ventrally; bulb elongated, with hooked apophysis and long weakly sclerotized

embolus.

Fegs. Without spines and curved hairs, few vertical hairs (most hairs missing); retrolateral trichobothrium

on tibia 1 at 14%; prolateral trichobothrium absent on tibia 1, present on other tibiae; tarsus 1 with >30

very indistinct pseudosegments.

Variation. Tibia 1 in 2 other males: 4.8, 5.4.

European Journal of Taxonomy 59: 1-44 (2013)

Female

In general similar to male but triads eloser together (distanee PME-PME 185 gm), furrows behind

oeular area less distinet (Fig. 96). Tibia 1 in 2 females: 4.6 (both). Epigynum large bulging area with

median internal tube-like strueture, posterior narrow plate with pair of small poekets (Figs 97, 98, 104);

internal genitalia as in Figs 99, 105.

Figs 92-99. Spermophora ziama sp. nov. 92-93. dorsal and lateral views. 94-95. Left f palp, prolateral

and retrolateral views. 96-97. $, dorsal and ventral views. 98-99. Cleared $ genitalia, ventral and

dorsal views. Seale lines: 92-93, 96-97 = 1 mm; 94-95, 98-99 = 0.5 mm.

HUBER B.A. & KWAPONG R, West African pholcid spiders

Figs 100-105. Spermophora ziama sp. nov. 100-101. Left 3 palp, prolateral and retrolateral views

(ba = bulbal apophysis; e = embolus; pr = procursus). 102-103. 3 chelicerae, frontal and lateral views.

104-105. Cleared $ genitalia, ventral and dorsal views. Arrows point at pockets in posterior epigynal

plate. Scale lines: 100-101, 104-105 = 0.5 mm; 102-103 = 0.3 mm.

European Journal of Taxonomy 59: 1-44 (2013)

Relationships

Several unique eharaeters indieate that S. ziama sp. nov. is elosely related to S. tonkoui: eurved

retrolateral apophysis on male palpal troehanter; proeursus with long transparent retrolateral proeess and

distinetive prolateral selerite between two putative hinges; distal whitish part of proeursus with many

short pseudotriehia ventrally; male ehelieeral armature (proximal frontal apophyses elose together); and

tube-like internal strueture in female genitalia.

The relationships of these two speeies to other taxa remain obseure. Spermophora tonkoui was explieitly

assigned tentatively to the genus (Huber 2003b), and reeent analyses of moleeular data of S. ziama sp.

nov. (Dimitrov et al. 2013: 12S, 18S, h3) provided eontradieting results, mostly poorly supported but

suggesting an affinity to a group eonsisting of Afriean Spermophora and the genera Buitinga Huber,

2003, Paramicromerys Millot, 1946, Spermophorides Wunderlieh, 1992, and Belisana Thorell, 1898

(Dimitrov et al. 2013: supplement fig. S2).

Natural history

The spiders were found in ereviees in the ground, between soil and roots or between soil and roeks.

Distribution

Known from type loeality only (Fig. 34).

Quamtana kitahurira Huber, 2003

Figs 106-113

Quamtana kitahurira Huber, 2003e: 513, figs 24, 204-208 {S, Uganda).

Quamtana kitahurira - Huber & Wami 2012: 12 {S, Rwanda).

New records

GUINEA: 8 SS 4 $$ 1 juv. in pure ethanol, in MRAC (236930), Mt. Nimba, Foret de Zie, near

Gouan eamp or “Station de Pompage Zie” [7°40’N, 8°26’W], 1250 m a.s.L, eanopy fogging of trees and

understory shrub layer, 3 Get. 2011 (D. Van den Spiegel, A. Hemard); \ S in pure ethanol, in MRAC

(236929), same data but beating trees and shrubs with hanging litter at 1.5-3 m above ground.

ANGOLA: I S in SMF (Ang 59.1), near Dundo [~7°23 ’S, 20°51 ’E], gallery forest of Chilanda river, in

bushes, 4 Oet. 1946 (A. de Barros Maehado).

Description

Female

In general similar to male, habitus as in Figs 106-108; entire animal pale oehre-gray, abdomen light

yellow-greenish; tibia 1: 3.4, 3.5, 3.8. Epigynum very simple externally, strongly protruding but weakly

selerotized, with pair of poekets not visible in disseeting mieroseope (Fig. 110; arrows in Fig. 112);

internal genitalia as in Figs 111, 113.

Distribution

Apparently widely distributed in tropieal Afriea, but so far only reeorded from Uganda, Rwanda, Guinea,

and Angola.

Note

The male from Angola has a slightly more slender proeursus but it otherwise identieal to speeimens from

Guinea and East Afriea; tibia 1: 4.0.

HUBER B.A. & KWAPONG R, West African pholcid spiders

Figs 106-111. Quamtana kitahurira Huber, 2003. 106-108. dorsal, ventral, and lateral views. 109.

S prosoma, oblique frontal view. 110-111. Cleared $ genitalia, ventral and dorsal views. Scale lines:

106-108 = 1 mm; 109 = 0.5 mm; 110-111 = 0.2 mm.

Figs 112-113. Quamtana kitahurira Huber, 2003. Cleared $ genitalia, ventral and dorsal views. Arrows

point at epigynal pockets. Scale line: 0.2 mm.

European Journal of Taxonomy 59: 1-44 (2013)

Discussion

Distribution patterns

West African pholcids can be divided into five groups (not monophyla) according to their distribution

patterns:

West African endemics

Most species listed above (23, i.e. 61%; or 72% if introduced species are not counted) are endemics of

the Upper Guinean subregion of the Guineo-Congolian center of endemism. About half of these species

belong to species groups that are endemic to the Upper Guinean subregion: the Pholcus guineensis

group (five species; Huber 2011b); the Smeringopina guineensis and ankasa groups (three species each;

Huber 2013); and Spermophora tonkoui + S. ziama sp. nov. Pholcus berlandi from Senegal seems to

belong in a species group more diverse in northern Africa (Huber 2011b). All other species belong in

tropical African genera and species groups: Anansus is widespread in tropical Africa (Huber 2007;

Huber & Wami 2012), with undescribed species in Ethiopia and Angola (interspecific relationships have

not been studied; B.A. Huber unpubl. data); Leptopholcus guineensis and L. kintampo sp. nov. belong

in a species group ranging all across tropical Africa (Huber 2011b); Spermophora dieke and S. akwamu

sp. nov. belong in a species group that includes an undescribed species from Congo D.R. (B.A. Huber

unpubl. data); Smeringopina beninensis is a notable outlier of a group otherwise restricted to Lower

Guinea (Huber 2013); finally, Smeringopina fan is a species with unclear relationships (closest relatives

in Upper or Lower Guinea; Huber 2013).

Sudanian species

Artema bunkpurugu sp. nov. and Crossopriza soudanensis are restricted to the northern part of West

Africa as defined herein. Both constitute the western limits of genera that are not tropical but range from

Sudanian West Africa to Central Asia and northern India (B.A. Huber unpubl. data).

Guineo-Congolian species

Six species are not restricted to the Upper Guinean subregion but cross the Dahomey Gap (and the

Cross River Basin) into Lower Guinea or even into Congolia: Nyikoa limbe (Upper and Lower Guinea;

Huber 2007, 2009); Leptopholcus tipula (entire Guineo-Gongolian region from Guinea to Albertine

Rift; Huber 2011b); Pholcus kakum which has not yet been found in Lower Guinea but ranges deeply

into Congolia (Huber 2011b); Smeringopus cylindrogaster (entire Guineo-Congolian region at least up

to Congo River; closest relatives in Lower Guinea and Congolia; Huber 2012); Spermophora kyambura

(entire Guineo-Gongolian region from Ghana to Albertine Rift, but note that specimens from Ghana

differ slightly from eastern populations; Huber & Wami 2012); finally, Quamtana kitahurira, which has

been collected in montane forests of the Albertine Rift (Uganda, Rwanda) and is here newly recorded

from Mt. Nimba in Guinea and northeastern Angola.

Widespread African species

A single West African species is widespread across several African forest blocks and transition zones:

Pehrfarsskalia conopyga, which ranges from Cape Verde to southern Africa and the Arabian Peninsula

(Huber 2011b), possibly as a result of its tolerance of severely degraded forests. The two congeners

known are both East African (Huber 2011b).

Synanthropic species

Six widespread synanthropic species originated outside West Africa or even outside Africa and have

attained pantropical distributions by human transport: Physocyclus globosus (all congeners in Mexico

and Central America; Valdez-Mondragon 2010); Modisimus culicinus (all congeners around the

Caribbean; Huber et al. 2010); Micropholcus fauroti (origin unclear, possibly Middle East; Huber

2011b); Smeringopus pallidus (African genus; closest relatives apparently in eastern Africa; Huber

HUBER B.A. & KWAPONG R, West African pholcid spiders

2012); Crossopriza lyoni (closest relatives apparently in Middle East; B.A. Huber unpubl. data); and

Artema atlanta (origin unclear, possibly Middle East or Central Asia; B.A. Huber unpubl. data).

Diversity and conservation

Until recently it was unclear if the relatively low pholcid diversity in West Africa as compared to East

and Central Africa was an artifact resulting from taxonomic neglect or a true phenomenon (Huber

2003b). In the meantime, data have accumulated that strongly suggest that the species diversity in West

Africa is indeed significantly lower than in areas of comparable size in Central and East Africa. A

recent overview of East African pholcids resulted in a total of 87 species (Huber & Wami 2012), and

an upcoming summary paper on Central African pholcids will comprise about 80 species (B.A. Huber

unpubl. data). This is in each case more than twice the number reported here for West Africa (38). Six

expeditions of comparable duration and collecting effort, two to each region, have resulted in very

similar differences among regions: only 28 species in Guinea + Ghana versus 60 species in Gabon +

Cameroon and 40 species in Kenya + Uganda. Percentages of undescribed species (as of2008) collected

at these expeditions suggest that the undiscovered diversity is much larger in Central Africa (83%) than

in East Africa (53%) and West Africa (43%).

A range of ideas have been proposed to explain why Africa in general appears to be less diverse than

South America and tropical Asia (White 2001; Eivingstone 2001; Eawson & Klemens 2001; Antonelli

& Sanmartin 2011), and some of these explanations may also apply to the low pholcid diversity of West

Africa compared to Central and East Africa.

1. Central Africa has a much larger area that is wet enough to support rain forest (about 3.1 x 10^ km^

vs. 1.2 X 10^ km^ in West Africa; Naughton-Treves & Weber 2001).

2. Geologically, West Africa is and has been more homogeneous and stable (especially compared to

East Africa), resulting in limited opportunities for speciation (Eawson & Klemens 2001).

3. Pleistocene dry periods have probably affected West Africa more severely than Central Africa,

resulting in fewer and smaller refuges of tropical moist forest (Maley 2001).

Human impact in prehistoric times has also been invoked to explain low African diversity in general

(White 2001), but whether this was more severe in West than in Central and East Africa is not known.

Another aspect of West African pholcid distribution is pertinent to conservation issues. There seems

to be a tendency for West African endemic species to have on average wider distributions than Central

African species. Relatively few species in West Africa seem to be local endemics (e.g., Leptopholcus

kintampo sp. nov.; Pholcus doucki; Spermophora ziama sp. nov., S. dieke), while this is common both in

Central Africa (e.g., in Smeringopina; Huber 2013) and East Africa (e.g., in Buitinga and Spermophora,

Huber 2003b; Huber & Wami 2012). This certainly requires denser sampling but if it turns out to be tme

it means that most West African species are not immediately threatened with extinction.

Outlook

After almost a decade of taxonomic work on African pholcids, a fairly realistic framework is emerging

about African pholcid diversity and distribution. All major and most small genera represented in Africa

have been revised (Huber 2003a, 2003b, 2003c, 2007, 2009, 2011b, 2012, 2013; Huber & El Hennawy

2007; Huber & Wami 2012), and the total number of species has more than tripled from 88 in 2002 to

287. At the level of genera, African pholcids can now be considered relatively well known.

However, the fact that the six expeditions mentioned above resulted in 74% undescribed species (87

of 117 species) clearly demonstrates that much taxonomic work remains to be done. The undiscovered

species diversity is still tremendous. Material deposited in major museums has largely been incorporated

in the above revisions. This means that if any significant progress is to be made, new collecting is

inevitable. In West Africa, the major gaps of knowledge are probably in Eiberia and Nigeria. The western

European Journal of Taxonomy 59: 1-44 (2013)

fauna of the Western Guinean forest zone (west of the Sassandra River, i.e. western Ivory Coast and

Liberia) is rieher than the eastern fauna (e.g., for mammals; Grubb 2001), but Liberia has essentially

seen no araehnologists interested in pholeid spiders. Mueh the same is true for Nigeria, a eountry almost

four times the size of Ghana but with only four pholeid speeies reeorded for the eountry (versus 18 for

Ghana). Nigeria is partieularly interesting for its position between Upper and Lower Guinea and with

respeet to the border between these subregions of the Guineo-Congolian forests (Maley 2001; Grubb

2001; Dowsett-Lemaire & Dowsett 2001).

Acknowledgements

BAH thanks all the people who kindly sent speeimens used in this study: J. Coddington, C.E. Griswold,

R Jager, R. Joeque, E.-A. Eeguin, E. Eeibensperger, Yu.M. Marusik, N. Platniek, C. Rollard, A. Russell-

Smith, S. Tehibozo, and J. Wunderlieh. BAH thanks R. Dunean and M. Sidibe for their help in aequiring

eolleetion and export permits in Guinea and J. Bosumtwe for his invaluable and friendly assistanee

during the eolleeting trip in Ghana. The German Researeh Foundation provided finaneial support (DEG,

HU 980/9-1).

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Manuscript received: 4 June 2013

Manuscript accepted: 12 September 2012

Published on: 18 October 2012

Topic editor: Rudy Jocque

Desk editor: Charlotte Thionois

Printed versions of all papers are also deposited in the libraries of the institutes that are members of

the EJT consortium: Museum National d’Histoire Naturelle, Paris, France; National Botanic Garden

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

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

Natural History Museum of Denmark, Copenhagen, Denmark.