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A revision of Pachyballus Simon, 1900 and Peplometus
Simon, 1900 (Araneae, Salticidae, Ballini) with
descriptions of new species
Wanda Wesołowska1, Galina N. Azarkina2, Konrad Wiśniewski3
1Department of Biodiversity and Evolutionary Taxonomy, University of Wrocław, Przybyszewskiego 65, 51-148
Wrocław, Poland 2Laboratory of Systematics of Invertebrate Animals, Institute of Systematics and Ecology of An-
imals, Siberian Branch Russian Academy of Sciences, Frunze Street 11, Novosibirsk 630091, Russia 3Institute
of Biology and Earth Sciences, Pomeranian University in Słupsk, Arciszewskiego 22b, 76-200 Słupsk, Poland
Corresponding author: Wanda Wesołowska (helena.wesolowska@uwr.edu.pl)
Academic editor: J. Miller | Received 7 January 2020 | Accepted 6 April 2020 | Published 30 June 2020
http://zoobank.org/CBF1BE5B-D1E5-408B-8769-E8FA935D6C78
Citation: Wesołowska W, Azarkina GN, Wiśniewski K (2020) A revision of Pachyballus Simon, 1900 and Peplometus
Simon, 1900 (Araneae, Salticidae, Ballini) with descriptions of new species. ZooKeys 944: 47–98. https://doi.
org/10.3897/zookeys.944.49921
Abstract
Two genera from the tribe Ballini (Araneae, Salticidae), Pachyballus Simon, 1900 and Peplometus Simon,
1900, are remarkable for their resemblance to beetles. eir biology is, however, poorly known and tax-
onomy has hitherto been rarely analysed. irteen species are included in this taxonomic revision of the
two genera. Six of them are new to the science: Pachyballus caelestis sp. nov. (♂♀, Congo D.R.), Pachybal-
lus miniscutulus sp. nov. (♂♀, South Africa), Pachyballus mombasensis sp. nov. (♂♀, Kenya), Pachyballus
ornatus sp. nov. (♂♀, Congo D.R. and Tanzania), Peplometus congoensis sp. nov. (♂♀, Congo and Congo
D.R.), and Peplometus nimba sp. nov. (♂, Guinea). One species (Pachyballus cordiformis Berland et Millot,
1941) and a subspecies (P. avipes aurantius Caporiacco, 1949) are recognised as synonyms of Pachyballus
avipes Simon, 1910. One new combination is proposed: Peplometus oyo (Wesołowska et Russell-Smith,
2011) comb. nov. (ex Pachyballus). e previously unknown females of Pachyballus transversus Simon,
1900 and Peplometus chlorophthalmus Simon, 1900, along with the males of Pachyballus castaneus Si-
mon, 1900 and Peplometus biscutellatus (Simon, 1887) are newly diagnosed and described. Neotypes for
Pachyballus castaneus and P. avipes are designated. Numerous new data on the distribution are provided
here and a key to Pachyballus females and to the males of Peplometus is presented. Identity of one species
remains doubtful, Pachyballus gambei (Simon, 1880).
Keywords
Africa, jumping spiders, mimicry, new combination, redescription, synonyms, taxonomy
ZooKeys 944: 47–98 (2020)
doi: 10.3897/zookeys.944.49921
https://zookeys.pensoft.net
Copyright Wanda Wesołowska et al. This is an open access article distributed under the terms of the Creative Commons Attribution License (CC
BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
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Wanda Wesołowska et al. / ZooKeys 944: 47–98 (2020)
48
Introduction
Simon (1900) established two African genera, Pachyballus and Peplometus, for some species
that mimic beetles and assigned them to the Balleae group (Simon 1901). Subsequently
Petrunkevitch (1928) included them in the subfamily Magoninae. Benjamin (2004) did not
study these two genera in his revision of Ballinae. According to the most up to date systemat-
ics of the jumping spiders (Maddison 2015), Pachyballus and Peplometus belong to the tribe
Ballini Banks, 1892 in the subfamily Salticinae Blackwall, 1841. Only a few species have
been described within these genera up to now, seven as Pachyballus (with one subspecies) and
two as Peplometus (WSC 2019). After our review the species number within the two genera
is nine and ve respectively.
Pachyballus and Peplometus are closely related and share most morphological characters.
ey are small but robust spiders, with a strongly attened body (Figs 31, 35, 137), covered
with a very hard, sclerotised exoskeleton. e dorsum of their body has a characteristic pitted
microsculpture (Fig. 6). e anterior part of the abdomen is covered by posterior edge of the
carapace, so that the pedicel is invisible (Figs 99, 172). A putative morphological synapo-
morphy for these two genera is the presence of characteristic scuta on the ventral surface of
their abdomen. is ventral “armament” consists of the two scuta: a narrow one along the
anterior margin of the abdomen, which laterally extends backwards, and a posterior trap-
ezoid scutum (Figs 1, 2). ere are also numerous minute sclerotised bumps on the ventral
side of the abdomen (Fig. 1). ese structures, in combination with dorsal strong sclerotisa-
tion, make members of these genera among the most heavily armour-plated spiders. Legs
are short, and the rst pair of male legs is clearly larger than others (Figs 19, 154) and has
thickened femora and the tibiae ventrally covered with dense setae.
e conformation of genitalia in both sexes is very similar in all species. Tibial apophysis
of the male palp is thin and straight, bulb oval, tegulum has a large posterior lobe, spiralled
embolus with more than three coils on the bulb tip (Figs 3, 4). e epigyne has anterior
semi-circular depression divided by a median septum (Figs 38, 107), very long copulatory
ducts with initial short spiral followed by several loops (Fig. 150), and more or less oval sper-
mathecae (Figs 96, 171). On the sides of the epigynal depression two crevices of unknown
role can sometimes be seen (Figs 95, 153). We observed once a broken embolus, which was
blocking the copulatory opening (Fig. 5), however we did not notice any other mating plugs.
Material and methods
Specimens examined it this study are deposited in the following institutions:
BMNH British Museum (Natural History) London, United Kingdom
CAS California Academy of Sciences, San Francisco, USA
HNHM Hungarian Natural History Museum, Budapest, Hungary
MCZ Museum of Comparative Zoology, Harvard University, Cambridge, USA
MEU Museum of Evolution, Uppsala University, Sweden
Revision of Pachyballus and Peplometus 49
Figures 1–6. Some morphological characters of Pachyballus and Peplometus 1 Pach. avipes (specimen
from Gabon), male, ventral abdominal scuta 2 Pach. ornatus (specimen from Tanzania), female, ventral
abdominal scuta 3 Pach. avipes (specimen from Congo), embolus 4 Pepl. biscutellatus (specimen from
Ivory Coast), palpal organ in ventral view 5 Pach. avipes (specimen from Zimbabwe), epigyne with bro-
ken embolus in copulatory opening 6 Pach. castaneus, pitted integument of abdomen.
Wanda Wesołowska et al. / ZooKeys 944: 47–98 (2020)
50
MNHN National Museum of Natural History, Paris, France
MRAC Royal Museum for Central Africa, Tervuren, Belgium
NCA National Collection of Arachnida, Pretoria, South Africa
NMBA National Museum, Bloemfontein, South Africa
NHRS Naturhistoriska Riksmuseet, Stockholm, Sweden
NMZ Natural History Museum, Bulawayo, Zimbabwe
SMF Senckenberg Natural History Museum, Frankfurt, Germany
UCZM Zoological Museum, University of Copenhagen, Denmark
e specimens were examined in 70% ethanol. e epigynes were macerated in
cold 5% KOH for 24 hours, dehydrated with absolute ethanol, cleared in xylene and
put in clove oil in a temporary microscope slides. A reticular eyepiece attached to a
stereomicroscope was used for drawing. After examination and reverting the above
described sequence, the female genitalia were placed to microvials and stored with
specimens. Specimens were measured as in Metzner (1999); all dimensions are given
in millimetres. A Nikon Coolpix 8400 or Canon EOS 550D mounted on the ster-
eomicroscope was used to take digital photos, which were stacked using Helicon Focus
image stacking software. Male-female matching was based both on the co-occurrence
of specimens and morphological similarities between the sexes (e.g., shape and col-
ouration of body). SEM microphotographs were taken with SEM Hitachi TM–1000.
e photographed parts were dried, and then mounted on an adhesive specimen stub.
e maps were prepared using DIVA-GIS.
Distinguishing genera
Due to numerous similarities of Pachyballus and Peplometus, many collectors have
failed to distinguish the two genera and labelled the specimens simply as Pachyballus.
All previously undetermined specimens analysed in this work had been assigned to
Pachyballus before and mostly determined only to the genus level. Peplometus after the
work by Simon (1901) was only found by Berland and Millot (1941) and mentioned
by Bodner and Maddison (2012; for specimen from Ghana used in molecular phylo-
genetic analysis).
Simon (1901) dierentiated the two genera by the shape of their carapace – signi-
cantly wider than long in Pachyballus and elongate in Peplometus (Figs 7, 8). However,
Simon depicted this feature on Pachyballus transversus, which has the widest carapace
among its congeners. In most species that can be assigned to this genus the ratio of
carapace width and length is approximately 1:1 and it may overlap with the propor-
tions seen in Peplometus. A better character allowing the separation of these genera is
the shape of abdomen. It is clearly elongated in Peplometes, whereas in Pachyballus the
length of abdomen is equal to its width. e other reliable feature for telling apart the
two genera, which may be applied only to males though, is the form of the tibia I.
ese tibiae in Pachyballus are not modied (Fig. 129), but in Peplometus are always
Revision of Pachyballus and Peplometus 51
conspicuously altered. ey are strongly thickened and often attened (Figs 130–133),
or considerably elongated, but in this case the metatarsus has a basal process (Figs
134, 135). Tibia I in Pachyballus bears dense and long hairs on ventral surface (Fig.
129), whereas in Peplometus it has a “brush” of long leaf-like setae (Figs 132–135).
e structure of tibia I of males may even be used in distinguishing Peplometus spe-
cies. Identifying females in this genus is very dicult in general and the results may be
doubtful. Conversely, species recognition by Pachyballus is relatively straightforward in
females, where a combination of morphological characters and genitalia conformation
is used. Determination of the males is sometimes impossible, because the structure
of their palpal organs is extremely similar and there often lack reliable distinguishing
morphological features. As a consequence of this, we were able to construct the key
only for Pachyballus females and males of Peplometus. Most probably, many taxonomic
intricacies will be possible to solve only using molecular methods. Fresh material for
analysis of the two genera will, however, be dicult to obtain.
Figures 7, 8. Simon’s drawings of body shape. 7 Pachyballus 8 Peplometus (from Simon 1901).
Table 1. Features for distinguishing Pachyballus and Peplometus.
Genus shape of the abdomen structure of leg I in male
Pachyballusrounded or heart-shaped (width-
length ratio = ca. 1:1)
tibia not modied, with long setae ventrally
Peplometus clearly elongated (width-length
ratio ≤ 0.8)
tibia very robust with dense feather-like setae ventrally, or
elongated with same setae and metatarsus with a dorsal hump
Wanda Wesołowska et al. / ZooKeys 944: 47–98 (2020)
52
Taxonomic account
Pachyballus Simon, 1900
Pachyballus Simon 1900: 399; 1901: 486.
Type species. Pachyballus transversus Simon, 1900.
Diagnosis. Pachyballus is closely related to Peplometus. From the latter genus it can
easily be separated by the rounded abdomen (elongated in Peplometus), by the form of
leg I in males (not modied) and by the absence of the leaf-like setae on tibia I.
Description. Small to medium-sized spiders (ca. 3.0–5.0 mm length), with very
at body, covered with tough highly sclerotised integument. Body colouration in
the majority of species dark brown or black with metallic lustre, only legs (espe-
cially in females) lighter. Carapace rounded, its width usually slightly larger than
the length, eye eld clearly trapezoid. Posterior part of carapace covered with abdo-
men. Chelicerae with three (exceptionally two) teeth on promargin and two to four
teeth on retromargin, basally fused together. Abdomen short and wide, heart-shaped
or rounded, its length to width ratio is 0.8–1.1, clearly wider than carapace, with
straight anterior border. Abdominal dorsum totally covered with strongly sclerotised
scutum, its edges sloping so that the abdomen has shape of a shallow bowl. Venter
with a narrow scutum along anterior margin, its lobes extending laterally as lobes. A
second, trapezoid scutum in posterior part, numerous very small sclerotised bumps
on both sides of abdomen (Fig. 1). Legs short, rst pair slightly bigger in males, with
enlarged femora. Tibia I not modied, setae on its ventral side not so dense as in
Peplometus. Tarsus of the female palp usually black. Male palp with oval bulb, embo-
lus spirally coiled around its tip (Fig. 3). Tibial apophysis thin and straight. Epigyne
with large semi-circular depression (atrium), usually divided by epigynal septum,
copulatory ducts usually long, forming a spiral in their initial parts and with several
complex loops distally.
Key to Pachyballus (females only)
1 Ventral scutum in posterior half of the abdomen present (Fig. 2) ................ 2
– Ventral scutum in posterior half of the abdomen absent............P. variegatus
2 Ventral posterior scutum trapezoid, large, its width about half of abdomen
width .......................................................................................................... 3
– Ventral scutum small, its width not more than third of abdomen width (Fig.
61) ....................................................................................... P. miniscutulus
3 Carapace clothed in white hairs, epigynal depression wide and short
(Fig. 16) ............................................................................ P. caelestis
– White hairs absent from carapace, epigynal depression long .......................4
4 Copulatory ducts relatively short, forming no more than a single loop........5
– Copulatory ducts very long, forming several loose loops ............................. 6
Revision of Pachyballus and Peplometus 53
5 Loop of the copulatory ducts tight (Fig. 28) ..............................P. castaneus
– Loop of the copulatory ducts loose (Fig. 73) ........................ P. mombasensis
6 Abdomen with a clear, contrasting pattern (Fig. 81) .....................P. ornatus
– Abdomen uniformly coloured ..................................................................... 7
7 Palps dark, abdomen rounded (Fig. 104) ................................ P. transversus
– Palps yellow, abdomen ovoid (Fig. 33) ..........................................P. avipes
Pachyballus caelestis sp. nov.
http://zoobank.org/DD252B66-8F33-4266-B6A1-670C09BA3D3F
Figures 9–17, 193
Holotype. C D.R. • ♀; Mayombe, Bas Congo, Luki Forest Reserve; 5°37'S,
13°05'E; 23.IX.2007; D. De Bakker and J.P. Michiels leg.; canopy fogging, old sec-
ondary rainforest; MRAC 226 102.
Paratypes. C D.R. • 1♀; same locality as the holotype; 25.IX.2007; can-
opy fogging, old secondary rainforest; MRAC 226 107 • 1♀; the same locality;
1.X.2007; canopy fogging, primary rainforest; MRAC 226 114 • 1♂; the same local-
ity; 13.XI.2006; canopy fogging, primary rainforest; MRAC 226 113.
Diagnosis. is species is covered very densely with short hairs, which is the best
feature to distinguish it from congeners. Male palpal organ has a characteristic embolus
that forms a considerably high and narrow coil similar to that of P. castaneus. e coil
comprises four loops (two and a half in the latter species). e female is distinctive in
having the epigyne with broad, short ridge on posterior edge of the epigynal depres-
sion; the copulatory ducts are relatively straight (they do not form any loops).
Etymology. e specic name is from Latin, meaning “soaring” and refers to this
species living high in a forest canopy.
Description. Male. Measurements. Cephalothorax: length 1.4, width 1.5, height
0.5. Eye eld: length 0.7, anterior width 1.2, posterior width 1.5. Abdomen: length
1.8, width 2.0.
General appearance as in Fig. 9. Small, very at spider with strongly sclerotised,
pitted integument. Carapace black, clothed in dense short light hairs. Eye eld trape-
zoid, distance between anterior lateral eyes shorter than between posterior laterals. Eyes
encircled by white scales. Clypeus low, with a few white hairs. Chelicera with two teeth
on promarginal edge and four retromarginal teeth. Endites and labium light brown with
whitish tips. Abdomen heart-shaped, wider than long, blackish with white scales on sides
and posterior part. Anterior margin of abdomen covers posterior part of carapace. Venter
brown, with typical scuta. Spinnerets black. Legs I the stoutest, femur and patella brown,
tibia slightly thickened, black with long dark setae ventrally, metatarsus and tarsus yel-
lowish. Tibia with one short ventral spine distally, metatarsus with two pairs of ventral
spines. Legs II the same in colour as the rst pair. Legs III and IV brown with yellow
distal segments. Pedipalp yellowish, its structure similar to that in other species, embolus
forms a considerably high and narrow coil that comprises four loops as in Figs 12–15.
Wanda Wesołowska et al. / ZooKeys 944: 47–98 (2020)
54
Female. Measurements. Cephalothorax: length 1.1–1.2, width 1.3–1.4, height
0.5. Eye eld: length 0.6–0.7, anterior width 1.0–1.1, posterior width 1.2–1.3. Abdo-
men: length 2.0–2.2, width 2.2–2.4.
General appearance as in Figs 10, 11. Slightly larger than male, shape of body simi-
lar. Carapace covered with white hairs. Colouration of abdomen a little lighter than
in male, brown, blackish in the mid part and along edge. Palps light brown. Epigyne
oval, central depression divided posteriorly by short, wide ridge (Fig. 16). Copulatory
ducts wide, spermathecae slightly smaller than in other species, small accessory glands
opening into copulatory ducts (Fig. 17).
Distribution. Known only from the type locality (Fig. 193).
Remarks. All specimens were collected by fogging. Probably, this species lives high
in the forest canopy.
Figures 9–13. Pachyballus caelestis sp. nov. 9 male, habitus, dorsal view 10 female, habitus, dorsal view
11 female, habitus, lateral view 12 palpal organ, ventral view 13 palpal organ, lateral view.
Revision of Pachyballus and Peplometus 55
Pachyballus castaneus Simon, 1900
Figures 6, 18–28, 129, 193
Pachyballus castaneus Simon 1900: 400 (♀).
Neotype. S A • ♀; KwaZulu-Natal, Ulundi, Ophathe Game Reserve;
28°23'S, 31°24'E; 3.X.2008; C. Haddad leg.; overgrazed savanna, beating shrubs;
NCA 2008/4147.
Figures 14–17. Pachyballus caelestis sp. nov. 14 palpal organ, ventral view 15 palpal organ, lateral view
16 epigyne 17 internal structure of epigyne.
Wanda Wesołowska et al. / ZooKeys 944: 47–98 (2020)
56
Paraneotypes. S A • 1♀; together with neotype • 1♂ 1♀; the same
locality as neotype; 3.X.2008; C. Haddad leg.; overgrazed savanna, beating, shrubs;
NCA 2008/4140.
Other material examined. S A • 1♂; the same locality as neotype;
2.X.2008; NCA 2008/4167 • 1♂ 2♀ 8 imm.; the same locality; 500 m a.s.l.; rocky moun-
tainside; NCA 2008/4154 • 3♂ 6 imm.; the same locality; 1.X.2008; NCA 2008/3993
• 1♂ 3 imm.; the same data; NCA 2008/3971 • 1♂; KwaZulu-Natal, iSimangaliso Wet-
land Park, Crocodile Centre; 28°21'S, 32°25'E; 14.V.2012; J.A. Neetling and C. Luwes
leg.; canopy fogging, wetland, Breonadia salicina; NCA 2012/5736 • 2♀; the same local-
ity, St. Lucia; 28°23'S, 32°25'E; 13.V.2012; J.A. Neetling and C. Luwes leg.; canopy fog-
ging, coastal forest, Trichilia dregeana; NCA 2012/4019 and NCA 2012/5737 • 1♀; Lake
St. Lucia, Fanies Island; 28°06'S, 32°27'E; 1.VII.1993; F.J. van der Lingen leg.; 200 m
from lake campsite, W shore, in dense bush thicket; NMBA 08069 • 1♀; Mkuze, Bang-
hoek Lodge; 27°45'S, 32°08'E; 17.V.2012; J.A. Neetling and C. Luwes leg.; canopy fog-
ging, Bushveld, Acacia karroo; NCA 2012/3965 • 1♂; Ndumo Game Reserve; 26°55'S,
32°19'E; 30.VI.2009; R. Lyle leg.; sand forest, beating foliage; NCA 2009/3660 • 1♀;
Sihangwane, Tembe Elephant Park, 40 km from Kosi Bay; 27°02'S, 32°25'E; 18.XI.1988,
R. Harris leg.; NCA 94/828 • 1♀; the same locality; 6.II.2008; R. Lyle and R. Fourie
leg.; beating, afromontane forest; NCA 2008/505 • 1♂ 1♀; Hellsgate; 28°00'S, 32°48'E;
23.VIII.2004; J. Esterhuizen leg.; NCA 2010/155 and NCA 2010/156 • 1♀; Mpuma-
langa Prov., Nelspruit, Agricultural College; 25°27'S, 30°59'E; 12.XI.1999; P. Stephen
leg.; beating, citrus; NCA 2000/223 • 1♀; Wildlife College, 10 km from Orpen Gate of
Kruger National Park; 24°28'S, 31°23'E; 11.X.2000; W. Breytenbach leg.; beating Eu-
clea divinorum; NCA 2003/626 • 1♂; Limpopo Prov., Little Leigh; 22°56'S, 29°22'E;
22.XI.2005; B. van der Waal leg.; branch beating, gallery forest; NCA 2009/2232. Z-
• 3♂; Mashonaland; Workman coll.; MCZ • 1♂; Harare; 17°50'S, 31°10'E;
2.III.2012; M. Cumming leg.; suburban garden, dropped from tree; NMZ.
Diagnosis. e male is indistinguishable from the males of P. avipes and P. mom-
basensis by body shape and colouration, but its bulb is slightly narrower than in these
species and the embolic spiral is tightly convoluted; width of the basal embolic loop
equals only a half of tegulum width, whereas in both other species it is as wide as
tegulum. e female can be separated from congeners in having copulatory ducts com-
pactly arranged and not forming loose loops (see Fig. 28).
Redescription. Male. Measurements. Cephalothorax: length 0.9–1.3, width 1.3–
1.9, height 0.5–0.6. Eye eld: length 0.6–0.7, anterior width 1.1–1.4, posterior width
1.3–1.9. Abdomen: length 1.7–2.2, width 1.8–2.6.
General appearance as in Figs 18, 19. Small spider with attened body covered with
strongly sclerotised and pitted integument (Fig. 6). Carapace dark brown, eye eld on
more or less half of carapace, vicinity of eyes black. A few long bristles at anterior eyes.
Posterior edge of carapace covered by abdomen. Chelicerae with three teeth on both
margins (Fig. 22). Mouth parts brown, sternum oval, brown, clypeus extremely low.
Abdomen dark brown, heart-shaped, as wide as long, dorsal scutum turned back via
margins to venter (Fig. 19), ventral scuta typical; the anterior one narrow with lateral
Revision of Pachyballus and Peplometus 57
extensions, posterior trapezoid. Numerous small bumps on sides. Colouration dark
brown to black, iridescent, integument clearly pitted, short hairs on edges of carapace,
dense bristles near eyes. Spinnerets brown. Legs I the thickest (Fig. 19), brown (except
yellow tarsi and metatarsi), femur, patella and tibia slightly thickened, tibia slightly
attened dorsally, covered with dense setae ventrally. Tibiae and metatarsi with two
pairs of short stout spines ventrally. Other legs yellow, with brownish femora. Pedipalp
Figures 18–21. Pachyballus castaneus 18 male (specimen from South Africa), habitus, dorsal view
19male habitus, lateral view 20 female, habitus, dorsal view 21 female, neotype, habitus, ventral view.
Wanda Wesołowska et al. / ZooKeys 944: 47–98 (2020)
58
brown, its structure as in Figs 23–26, slightly narrower than in congeners. Palpal tibia
short with single thin straight apophysis, bulb oval, embolus thin, long, spirally coiled
on bulb tip.
Female. Cephalothorax: length 0.9–1.4, width 1.2–1.4, height 0.5–0.6. Eye eld:
length 0.5–0.7, anterior width 0.9–1.1, posterior width 1.2–1.4. Abdomen: length
1.7–2.0, width 1.8–2.1.
General appearance as in Figs 20, 21. Similar to male, abdomen almost round or
oval, but its anterior margin almost straight. Abdominal venter with two scuta as in
male (Fig. 21). All legs and palps dark yellow. Epigyne typical, with horseshoe-shaped
anterior depression (Fig. 27). Internal structures relatively simple, copulatory ducts
shorter than in congeners, compactly arranged, spermathecae strongly sclerotised
(Fig. 28).
Immature specimens. Similar to adults, abdomen covered dorsally with one large
scutum.
Distribution. Known from South Africa and Zimbabwe (Fig. 193).
Figures 22–28. Pachyballus castaneus 22 cheliceral dentition 23–25 palpal organ, ventral view (24 speci-
men from Zimbabwe) 26 palpal organ, lateral view 27 epigyne 28 internal structure of epigyne.
Revision of Pachyballus and Peplometus 59
Designation of neotype. Pachyballus castaneus was originally described from Natal
(South Africa) on the basis of a single female. e type specimen was lost (the collec-
tion manager informed us that the type could not be found in Simon’s collection in
MNHN). e original description is very supercial (only body size and colouration
of legs are mentioned) and insucient for identication of the species. Colouration of
legs varies within dierent Pachyballus species (see Figs 32, 33, 104–106), so it cannot
be used as a taxonomic character. e neotype, a female that originates from the same
province as the type, is herein designated to stabilise the nomenclature.
Remarks. e male of this species is described here for the rst time.
Pachyballus avipes Simon, 1910
Figures 1, 3, 5, 29–57, 194
Pachyballus avipes Simon 1910: 414 (♀); Lessert 1925: 434, f. 7–9 (♀); Wanless and
Clark 1975: 290, f. 27–28 (♀); Dawidowicz and Wesołowska 2016: 449.
P. avipes aurantius Caporiacco 1949: 464 (♀), syn. nov.
Pachyballus cordiforme Berland and Millot 1941: 396, f. 90 (♂), syn. nov.
Pachyballus cordiformis Wesołowska and Cumming 2011: 87, f. 40–45 (♂).
Neotype. C • ♀; Biniiba, Bétaré-Oya; 5°36'N, 14°05'E; 20.VII.1949; B.
Malkin leg.; CAS.
Other material examined. A • 1♀; Hulla prov., Caconda; 13°46'S, 15°05'E;
30.IX.1949; B. Malkin leg. CAS. B • 1♀; Okavango Delta, Pom-Pom Camp;
19°18'S, 22°54'E; VII.2001; E. Kassimatis leg.; sweeping; NCA 2009/5688. C
• 1♀; Faro Game Reserve; 8°30'N, 12°30'E; 25.IV.2007; R. Jocqué, K. Loosveldt, L.
Baert and M. Alderweireldt leg.; gallery forest, sieving; MRAC 221 442. C D.R.
• 1♀; South Kivu prov., Itombwe; 3°15'S, 28°50'E; 3200 m a.s.l.; XII.1958; N. Lele-
up leg.; forest with Hagenia and bamboo; MRAC 113 229 • 1♀; Semliki river val-
ley; 9.VIII.1968; R.P.M. Lejeune leg.; MRAC 135 557 • 2♂; Mayombe, Bas Congo,
Luki Forest Reserve; 5°37'S, 13°05'E; 17.IX.2007; D. De Bakker and J.P. Michiels leg;
old secondary rainforest, fogging; MRAC 226 092 • 1♂; the same data; 18.IX.2007;
MRAC 226 097 • 1♂; the same data; 20.IX.2007; MRAC 226 095 • 1♀; the same data;
23.IX.2007; MRAC 226 103 • 2♀; the same data; 18.IX.2007; MRAC 226 096 • 1♀;
the same data; 20.IX.2007; MRAC 226 094 • 2♀; the same data; 21.IX.2007; MRAC
226 093 • 1♂ 2♀ 2 subad. ♂; the same data; 17.IX.2007; MRAC 226 101 • 2♂; the
same locality; 28.IX.2007; primary rainforest, fogging; MRAC 226 109 • 1♀; the same
data; 1.X.2007; MRAC 226 116 • 1♂ 1♀; the same data; 1.X.2007; MRAC 226 120
• 3♀; the same data; 7.XI.2006; MRAC 220 945 • 3♀; the same data; 13.XI.2006;
MRAC 220 971 • 1♀; the same locality; 14.IX.2007; secondary rainforest, beating;
MRAC 223 432 • 2♂; Kivu prov., Ngoma; 4°24'S, 26°05'E; L. Burgeon leg.; MRAC 15
552/15 553 • 1♀; Kivu prov., Rutshuru, Kako; 1°11'S, 29°27'E; IX.1932; L. Burgeon
leg.; MRAC 31 267 • 1♀; Kivu prov., Sanga plateau; 4°50'S 14°58'E; N. Leleup leg.;
Wanda Wesołowska et al. / ZooKeys 944: 47–98 (2020)
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MRAC 119189. G • 1♂; Biso-Binam [Biso stream?]; 0°52'N, 11°39'E; 3.XI.1985;
A. Pauly leg.; on Borreria verticillata; MRAC 172 765. G • 1♂; Nimba Mts,
“Mare d’hivernage” (in UNESCO Biosphere reserve); 7°38'N, 8°27'W; 1650 m a.s.l.;
1.X.2008; D. van den Spiegel leg.; wet grassland with dispersal shrubs; MRAC 225978.
I C • 1♂; Man; 7°24'N, 7°33'W; VIII,1937; J. Millot leg.; type of P. cordi-
formis: only two, left palps and legs left; MNHN • 1♀; Bingerville; 5°21'N, 3°54'W;
VIII.1962; J. Decelle leg.; MRAC 122.004. K • 1♂; Kwale prov., 30 km S to Mom-
basa; 4°10'S, 39°40'E; 12.XI.1992; V. Roth leg.; CAS • 1♀; Kitale; 1°01'N, 35°00'E;
2000 m a.s.l.; 23.I.1938; MEU • 1♀; Nairobi; 1°17'S, 36°49'E; 12.I.1970; on bamboo;
SMF • 1♂; Mt Elgon, E slope; 1°07'N, 34°31'E; 2130 m a.s.l.; 6.I.1938; NHRS • 1♀;
Mt Elgon, Salt lake estate; 2100 m a.s.l.; 17.XII.1937; Acacia steppe; MEU. T
• 1♀; Kilimanjaro, Kibonoto; 3°11'S, 37°06'E; Sjöstedt leg.; NHRS. U • 1♀;
Victoria Lake, Gaya Bay, Island Buvuma; 0°13'N, 33°16'E; III.1968; E. Vertriest leg.;
MRAC 134 737. S A • 1♂; Free State, Weltevreden Nature Reserve; 28°57'S,
26°23'E; 15.VIII.2006; H. Kilion leg.; NCA 2007/3429 • 1♀; Mpumalanga Prov., Nel-
spruit, Agricultural College; 25°27'S, 30°59'E; 22.XII.1998; P. Stephen leg.; on grape-
fruit; NCA 99/160 • 1♀; Limpopo Prov., Nylsvley Nature Reserve; 24°39'S, 28°41'E;
7.III.1998; sweeping, grass; A.S. Dippenaar leg.; NCA 98/586. Z • 1♂; 30km SW
of Mkushi; 13°43'S, 29°15'E; 1390 m a.s.l.; 22.IX.2009; J.Lenz leg.; NMZ. Z
• 1♂ 2 imm.; Batoka Gore; 17°57'S, 26°04'E; 30–31.X.1990; V. Roth leg.; CAS • 1♂
2♀; Matabeleland, Doddieburn Dam; 21°24'S, 29°21'E; 12.XII.1985; J. Minshull leg.;
NMZ A4202 • 5♀ 2 subad. ♂ 8 imm.; Matetsi safari Area, Tshowe river rapids; 18°31'S,
25°52'E; 5.XII.1988; J. Minshull leg.; NMZ A6873 • 2♂; Matetsi safari Area, Kasetsheti
Weirs; 18°01'S, 25°49'E; 11.X.1988; F. Nyati leg.; NMZ A6759.
Diagnosis. e male is almost indistinguishable from that of P. mombasensis, though
it diers a little by having a protruding tibial apophysis while in the latter species tibial
apophysis is adpressed to cymbium. e female resembles females of P. castaneus and P.
mombasensis, but has very long copulatory ducts, forming several loops, whereas in two
other species these ducts are relatively short (cf. Fig. 51 with Fig. 73 and Fig. 28).
Redescription. Male. Measurements. Cephalothorax: length 1.2–1.6, width 1.4–
1.7, height 0.5–0.6. Eye eld: length 0.6–0.8, anterior width 1.1–1.4, posterior width
1.4–1.7. Abdomen: length 1.9–2.5, width 1.8–2.3.
General appearance as in Figs 29–31. Shape of body typical for Pachyballus; small,
at, with heart-shaped abdomen. Body covered with strongly sclerotised and clearly
pitted integument. Colouration of body brown to black dorsally and dark brown ven-
trally, dorsum iridescent, almost hairless. Some light hairs near anterior eyes and below
anterior median eyes. Anterior margin of abdomen covers distal part of carapace. Ven-
ter with typical scuta and small sclerotised bumps (Fig. 1). Cheliceral dentition vari-
able, with three teeth on promargin and two, three or (exceptionally) four on retromar-
gin (Figs 41–43). Tips of mouth parts whitish. First pair of legs the stoutest, femora
and patellae dark brown, tibiae black with dense long dark setae ventrally and two pairs
of very short thick spines (Fig. 31), dorsal part of tibia slightly attened, metatarsi and
tarsi creamy. Other legs with clear contrasts: brown femora, other segments light, only
Revision of Pachyballus and Peplometus 61
sometimes dark ring on tibiae proximally (Fig. 30). Palps dark, tip of cymbium light.
Structure of palpal organ as in Figs 3, 37, 44–46.
Female. Measurements. Cephalothorax: length 1.2–1.3, width 1.3–1.4, height
0.5. Eye eld: length 0.6–0.8, anterior width 1.1–1.3, posterior width 1.3–1.4. Abdo-
men: length 1.9–3.1, width 1.7–2.9.
General appearance as in Figs 32–36. Similar to male, abdomen more oval. An-
terior median eyes surrounded by short light hairs. All legs yellowish (Fig. 35), only
femora brownish (or yellow with darker streak). Palps yellow (Figs 32, 36). Epigyne as
in Figs 5, 38, 47–50, with horseshoe-shaped anterior depression. Internal structure of
epigyne as in Figs 39, 40, 51–57, inlet part of copulatory ducts wide.
Immature specimens. Similar to adults, abdomen dorsally covered with one large
scutum.
Figures 29–35. Pachyballus avipes 29 male, habitus, dorsal view 30 male, habitus, ventral view
31male, habitus, lateral view 32, 33 female, habitus, dorsal view 34 female, habitus, ventral view 35fe-
male, habitus, lateral view (29, 30, 32 specimen from Congo 31 specimen from Kenya 33–35 specimen
from Cameroon).
Wanda Wesołowska et al. / ZooKeys 944: 47–98 (2020)
62
Distribution. Species widely distributed in Africa (Fig. 194). Some of the records
from Kenya were already mentioned in Dawidowicz and Wesołowska (2016).
Synonymisation. Caporiacco (1949) described P. avipes aurantiacus from Kenya
on the basis of a single female. According to him, the epigyne of this subspecies was the
same as in P. avipes and the only dierence consisted of colouration. However, this
feature is variable. Moreover, according to Berdondini and Whitman (2003) type speci-
men of P. avipes aurantiacus kept in the Natural History Museum of Florence collec-
tion is juvenile. us, we recognise P. f. aurantiacus as a synonym of the name P. avipes.
Figures 36–40. Pachyballus avipes (specimen from Congo) 36 female, habitus, frontal view 37 palpal
organ, ventral view 38 epigyne 39 internal structure of epigyne, ventral view 40 internal structure of
epigyne, dorsal view.
Revision of Pachyballus and Peplometus 63
Type of P. cordiformis was destroyed and only two palps and three legs persisted in
the vial. Although Berland and Millot (1941) reported that they had collected only a
single male by the city Man, the two palps in the sample are left palps, so they must
have been taken from two males. Structure of palpal organ and the gure in Berland
and Millot (1941) suggest that this species is identical with P. avipes. erefore, we
recognise the name P. cordiformis as the synonym of P. avipes.
Designation of neotype. Simon (1910) described this species from Bioko (Fer-
nando Po) on the basis of a single female. Wanless and Clark (1975) compared a female
from Ivory Coast with the type and concluded their conspecicity. Unfortunately, the
type specimen was lost (we were informed that the type could not be found in Simon’s
collection in MNHN) and the female collected by the latter authors lacked epigyne.
Original description was insucient to recognise the species; only the shape of body
and colouration of legs were given. Taking this under consideration it is justied to
propose the neotype to stabilise the nomenclature. e neotype, a female collected in
Cameroon, the nearest continental country to Bioko (ca. 40 km), is herein designated.
Figures 41–46. Pachyballus avipes, male 41–43 variation of cheliceral dentition 44–46 palpal organ,
ventral view (44 specimens from Congo 45 specimen from Gabon 46 type of P. cordiformis).
Wanda Wesołowska et al. / ZooKeys 944: 47–98 (2020)
64
Remarks. is species probably lives both in the forest canopy and in the under-
story. Given this fact, i.e. the variety of preferred microhabitats, the large geographical
range and high variation of cheliceral dentition, it is possible that it consists of several
cryptic species. e sole morphology may be insucient to solve this taxonomic prob-
lem and there is a need to support further analysis with molecular methods.
Pachyballus gambeyi (Simon, 1880)
Homalattus gambeyi Simon 1880: 166 (♂).
Pachyballus gambeyi Simon 1901: 485; Prószyński 1987: g. on p. 73 (♂).
Holotype. N C • 1♂; [leg.] Bougier; MNHN 3646; not examined.
Remarks. e holotype was examined by Prószyński (1987). His drawings show
that this specimen has palpal organ typical for Pachyballus, but morphological charac-
ters do not allow any specic identication. Simon described this species from Canala
(21°32'S, 165°57'E), central New Caledonia. All species of Pachyballus are restricted to
Africa, which is thousands kilometers from New Caledonia. is distribution pattern
Figures 47–50. Pachyballus avipes, female, epigyne (47, 48 specimens from Congo 49 specimen from
Kilimanjaro 50 specimen from Cameroon).
Revision of Pachyballus and Peplometus 65
is highly improbable, and the situation might have resulted simply by mislabelling the
sample, which is not an uncommon problem in collections. As the provenience of the
specimen is doubtful we conclude that the status of P. gambeyi remains unclear.
Pachyballus miniscutulus sp. nov.
http://zoobank.org/3DC50D0E-AC8C-4810-827C-CD04B0674B80
Figures 58–67, 195
Holotype. S A • ♂; Free State, Bloemfontein, National Botanical Gardens;
29°02'S, 26°12'E; 12.X.2012; C. Haddad leg.; sweeping, vegetation along stream;
NCA 2019/1444.
Figures 51–57. Pachyballus avipes, female, internal structure of epigyne 51, 53, 55, 56 ventral view 52,
54, 57 dorsal view (51, 52 specimens from Cameroon 53–55 specimen from Congo 56, 57 specimen
from Zimbabwe).
Wanda Wesołowska et al. / ZooKeys 944: 47–98 (2020)
66
Paratypes. S A • 3♀; together with holotype • 1♀; Free State, Bloem-
fontein, National Botanical Gardens; VII.2012; L. de Jager and J. van der Merwe leg.;
karree litter (Searsia lancea), streamside; NCA 2019/1446 • 1♀; the same locality;
19.XII.2012; C. Haddad grassland leg.; pitfall traps; NCA 2013/1635 • 2♀; the same
locality; 19.XI.2012; C. Haddad leg.; sweeping, open grassland; NCA 2013/1604 •
4♀; the same locality; 12.X.2012; C. Haddad leg.; sweeping, vegetation along stream;
NCA 2012/5707 • 2♀; KwaZulu-Natal, Ithala Game Reserve, picnic site; 27°33'S,
31°19'E; 29.I.2014; C. Haddad leg.; base of grass tussocks; NCA 2013/5098.
Diagnosis. is species is distinctive in having a unique size of ventral posterior
scutum (Figs 59, 61) that is clearly smaller than in other species. Its width is equal to
spinnerets area (2–3 times larger in the congeners). e female has the epigyne similar
to that in Pachyballus mombasensis, but the copulatory ducts are longer (cf. Fig. 74 with
Figs 66, 67).
Etymology. e specic name is derived from the Latin words “mini-” and “scu-
tum”, meaning “small” and “shield” correspondingly, and refers to the small size of
ventral posterior scutum.
Description. Male. Measurements. Cephalothorax: length 1.3, width 1.25, height
0.6. Eye eld: length 0.7, anterior width 1.0, posterior width 1.2. Abdomen: length
1.7, width 1.7.
General appearance as in Figs 58, 59. Colouration of carapace dark brown, with
black rings around eyes, some long bristles at rst row of eyes. Chelicerae dark brown.
Clypeus and cheeks dark brown, covered with sparse white hairs. Labium and endites
yellowish brown, paler apically. Sternum yellowish brown. Abdomen heart-shaped,
dark brown dorsally. Venter brownish grey, with a small posterior scutum, ranging at
one fth of abdomen length (Fig. 59). Book-lung covers yellow. Spinnerets yellowish
brown. First pair of legs brown with yellow tarsi. Legs II-IV light brown. Leg hairs
brown. Structure of palpal organ as in Figs 62, 63, embolic coil wide, comprises 2.5
loops, palpal tibia with protruding apophysis.
Female. Measurements. Cephalothorax: length 1.0–1.1, width 1.1–1.2, height 0.5–
0.6. Eye eld: length 0.5–0.6, anterior width 0.9–1.0, posterior width 1.1–1.2. Abdo-
men: length 1.8–1.9, width 1.5–1.8.
General appearance as in Figs 60, 61. Similar to male. Posterior ventral scutum small,
as in male. All legs and palps yellow. Epigyne as in Figs 64, 65, with spade-like or round
central part in semi-circular depression. Internal structure of epigyne as in Figs 66, 67.
Distribution. Known from South Africa only (Fig. 195).
Pachyballus mombasensis sp. nov.
http://zoobank.org/1A041DD3-803F-49E4-AA4B-F279265DC748
Figures 68–74, 195
Holotype. K • ♀; Diani Beach, 30 km S to Mombasa; 4°19'S, 39°34'E; 5–19.
III.1970; T. Palm leg.; MEU.
Revision of Pachyballus and Peplometus 67
Figures 58–61. Pachyballus miniscutulus sp. nov. 58 male, holotype, habitus, dorsal view 59 male, habi-
tus, ventral view 60 female, habitus, dorsal view 61 female, habitus, ventral view.
Wanda Wesołowska et al. / ZooKeys 944: 47–98 (2020)
68
Paratype. K • 1♂; together with the holotype; +3 imm.
Diagnosis. e male is indistinguishable from the males of P. avipes and P. castaneus
by body colouration; it diers from P. castaneus in having a wider bulb and wide and low
embolic coil (cf. Fig. 70 with Fig. 25); from P. avipes it can be distinguished by the pal-
pal tibial apophysis adpressed to cymbium, (protruding in P. avipes) .e latter character
requires verication though, as only single male of P. mombasensis is known. e female
has relatively short copulatory ducts, forming only a single loop (Fig. 73).
Etymology. e specic name is derived from the type locality.
Description. Male. Measurements. Cephalothorax: length 1.1, width 1.5, height
0.5. Eye eld: length 0.6, anterior width 1.1, posterior width 1.5. Abdomen: length
2.1, width 2.1.
Body attened, integument strongly sclerotised, clearly pitted. Colouration of
carapace dark brown, some bristles by the rst row of eyes. Mouth parts brownish
with lighter tips. Chelicera with three retromarginal teeth. Abdomen very at, heart-
shaped, dark brown, venter with typical scuta. First pair of legs brown, tibia short,
slightly thickened, its dorsal part a little attened, two pairs of ventral spines and dense
dark setae. Other legs yellowish, only femora brownish. Palpal organ as in Figs 70, 71,
embolic coil wide and low, tibial apophysis adpressed to cymbium.
Figures 62–67. Pachyballus miniscutulus sp. nov. 62 holotype, palpal organ, ventral view 63 palpal or-
gan, lateral view 64, 65 epigyne 66, 67 internal structure of epigyne, dorsal view.
Revision of Pachyballus and Peplometus 69
Figures 68–74. Pachyballus mombasensis sp. nov. 68 female, holotype, habitus, dorsal view 69 female,
habitus, ventralal view 70 palpal organ, ventral view 71 palpal organ, lateral view 72 holotype, epigyne
73 internal structure of epigyne, ventral view 74 internal structure of epigyne, dorsal view.
Wanda Wesołowska et al. / ZooKeys 944: 47–98 (2020)
70
Female. Measurements. Cephalothorax: length 1.3, width 1.5, height 0.5. Eye
eld: length 0.7, anterior width 1.1, posterior width 1.5. Abdomen: length 2.1,
width 2.1.
General appearance as in Figs 68, 69. Similar to male, rst pair of legs not stouter
than others, palps dark. Posterior ventral scutum large. Epigyne with semicircular de-
pression (Fig. 72). Internal structure as in Figs 73, 74, copulatory ducts clearly shorter
than in congeners, spermathecae comparatively large, strongly sclerotised.
Immature specimens. As adults, dorsum of abdomen covered with one large
scutum.
Distribution. Known only from the type locality (Fig. 195).
Pachyballus ornatus sp. nov.
http://zoobank.org/862AF941-D594-45CC-BBD5-7A38EA629BE7
Figures 2, 75–97, 195
Holotype. T • ♂; Tanga region, Usambara Mts, Amani Nature Reserve, Mbo-
mole Hill; 5°05'S, 38°37'E; 1000 m a.s.l.; 5–8.XI.1995; C. Griswold, N. Schar, D.
Ubick leg.; CAS.
Paratypes. C D.R. • 1♂; Equateur, Bokuma; 0°06'S, 18°42'E; in 1952; R.P.
Lootens leg.; MRAC 85 126 • 1♂; Mayombe, Bas Congo, Luki Forest Reserve; 5°37'S,
13°05'E; 7.XI.2006; D. De Bakker and J.P. Michiels leg.; primary rainforest, fogging;
MRAC 226 115 • 1♂; Kivu, Ibanda [Bukavu]; 2°29'S, 28°50'E; SMF. T •
2♂ 2♀ (1 of females – black form); together with the holotype; CAS • 1♂; Mundi
distr., Iringa region, Uzungwa Scarp Forest Reserve; 8°32'S, 35°54'E; 750 m a.s.l.;
8.III.1996; McKamey leg.; canopy fogging; UCZM • 1♂; the same locality, by Chita
village; 1300–1400 m a.s.l.; 26.X–14.XI.1984; N. Schar leg.; montane rain forest;
UCZM • 3♂ 4♀ (2 of females – black form); 11 km SE Masisiwe Kihanga Stream;
8°32'S, 35°58'E; 1800 m a.s.l.; 17–27V.1997; canopy; UCZM • 1♂; Uluguru Mts,
Kimboza Forest Reserve; 7°20'S, 37°47'E; 250 m a.s.l.; 18.VII.1981; M. Stoltze and
N. Schar leg.; UCZM.
Diagnosis. A characteristic body pattern allows an easy recognition of this species;
the abdomen is light with dark margins and a dark streak or central patch (Figs 78,
81). Some specimens, probably young adult females (soon after moulting), can be uni-
formly black and resemble females of P. avipes, but they are distinguishable in having
dark palps (that are yellow in the latter species).
Etymology. e specic name is Latin and means decorated, which refers to the
characteristic colour pattern.
Description. Male. Measurements. Cephalothorax: length 1.2–1.7, width 1.5–
1.8, height 0.5–0.6. Eye eld: length 0.8–1.0, anterior width 1.1–1.4, posterior width
1.5–1.8. Abdomen: length 1.7–2.3, width 1.8–2.4.
General appearance as in Figs 75–80. Body at, integument strongly sclerotised,
clearly pitted. Carapace wider than long, eye eld trapezoid (distance between anterior
lateral eyes shorter than between posterior laterals), occupies about half of carapace.
Revision of Pachyballus and Peplometus 71
Colouration of carapace brown, black around eyes, in some specimens with two large
yellowish patches in eye eld (Figs 76, 78, 80). Dense bristles at eyes of rst row and
at lateral edges of carapace. Chelicera with three short teeth on promargin and saw-
shaped tooth with four tips on retromargin. Labium and endites brownish with lighter
tips, sternum rounded, brown. Abdomen very at, triangular to heart-shaped, wider
than long, its anterior margin almost straight. Colouration of abdomen yellow, brown
on the edges, with brown longitudinal median streak (Fig. 77), which is reduced to
round patch positioned centrally in some specimens (Fig. 79). Variation of pattern as
presented in Figs 76–80. Abdomen with typical ventral scuta (Figs 2, 75). Spinnerets
yellow. First pair of legs slightly stouter than other legs, femora enlarged, tibiae slightly
thickened, brown, only tarsi light. Other legs yellowish to light brown, femora darker.
Palps as in Figs 86, 87, embolic coil low, but basal loop high.
Female. Measurements. Cephalothorax: length 1.3–1.5, width 1.4–1.8, height
0.5–0.6. Eye eld: length 0.6–0.8, anterior width 1.1–1.2, posterior width 1.4–1.5.
Abdomen: length1.9–2.1, width 2.0–2.2.
General appearance as in Figs 81–84. Body shape and colouration similar to that
in male. First legs not enlarged. Palps dark. Epigyne with anterior semi-circular de-
Figures 75–80. Pachyballus ornatus sp. nov., male, habitus 75 ventral view 76–80 dorsal view (all speci-
mens from Tanzania).
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72
pression divided by a median septum (Figs 88–91). Copulatory ducts very long, form
several loose loops, spermathecae oval (Figs 92–97).
Distribution. Tanzania and Congo D.R. (Fig. 195).
Remarks. Some specimens, probably young females (soon after the moulting),
were dark, almost black (Fig. 84). ey appear to become brighter with age due to
guanine accumulation (light spots are formed by guanine crystals stored under integu-
ment). Black females were collected together with light specimens. Internal structure
of epigynes similar in light, Figs 92–94, and black, Figs 85, 95–97 specimens. Col-
ouration seems to turn brighter in the course of life also in males.
Pachyballus transversus Simon, 1900
Figures 98–122, 191, 192, 196
Pachyballus transversus Simon 1900: 399 (♂); 1901: 482, f. 570–571; 1910: 414; Ber-
land and Millot 1941: 397.
Figures 81–85. Pachyballus ornatus sp. nov., female, 81–84 habitus, dorsal view 85 epigyne (all speci-
mens from Tanzania).
Revision of Pachyballus and Peplometus 73
Syntypes. C • 2♂; Mayombe; E. Simon coll.; MNHN 7545; examined.
Other material examined. C • 1♀; Maroua; 10°36'N, 14°19'E; col.
C.F. Roewer (nr 12 678); SMF • 1♀; without precise locality; in 1950; J. Birket-Smith
leg.; UCZM. C D.R. • 1♀; Mayombe, Bas Congo, Luki Forest Reserve; 5°37'S,
13°05'E; 28.IX.2007; D. De Bakker and J.P. Michiels leg.; primary rainforest, fogging;
MRAC 226.109A • 3♂ 2 imm; Kivu N prov., Kaisola, Ruindi plain; 0°47'S, 29°17'E;
1100 m a.s.l.; 3.VII.1972; M. Lejune leg.; MRAC 144 487 • 1♂ 2♀ 1 imm.; Ruindi
plain, Ndimo Hill; 1100 m a.s.l.; 28.VI.1972; P.M. Lejune leg.; MRAC 144 670 •
1♂; Kivu N prov., Rutshuru; 1°11'S, 29°27'E; III.1937; J. Ghesquière leg.; MRAC
30 583 • 1♂; Goma; 0°34'S, 28°42'E; 10.II.1952; E. Bertrand leg.; MRAC 78 975.
E • 3♂ 2 imm.; Gorgora; 12°14'N, 37°18'E; in 1961; F. Hartman leg.; MRAC
131 203 • 1♂; Yayu coe forest plantation; 8°10'N, 36°00'E; in 2004; N. Aklilu leg.;
MRAC 231 209 • 1♀; the same locality; beating; MRAC 230 736 • 1 imm.; the same
locality; secondary forest, beating; MRAC 229 396. I C • 1♀; Bingerville;
5°21'N, 3°54'W; VIII.1962; J. Decelle leg.; MRAC 122 004. M • 1♀;
‘N Mozambique’; col. C.F.Roewer (nr 9715); SMF. S • 1♀; Sinandogo; in
1946; R. Accigliaro leg.; MRAC 131 175 • 1♀; Giumbo; 0°14'S, 42°37'E; in 1946;
Figures 86–91. Pachyballus ornatus sp. nov., copulatory organs 86 palpal organ, ventral view 87 palpal
organ, lateral view 88–91 epigyne (89, 91 black form 89, 90 specimens from Usambara Mts).
Wanda Wesołowska et al. / ZooKeys 944: 47–98 (2020)
74
R. Accigliaro leg.; MRAC 131 223. T • 2♂; ‘Zanguebar’ (=eastern coast of
tropical Africa, probably Zanzibar); E Simon coll.; MNHN 7021. S A •
1♂; KwaZulu-Natal, Ulundi, Ophathe Game Reserve; 28°23'S, 31°24'E; 3.X.2008;
C. Haddad leg.; beating, shrubs, overgrazed savanna; NCA 2008/4154 • 1♂; the same
locality; 1.X.2008; NCA 2008/3971 • 9♂; the same locality; NCA 2019/1448 • 1♀;
Mpumalanga Prov., Nelspruit, Agricultural College; 25°27'S, 30°59'E; 12.XI.1999; P.
Stephen leg.; beating, citrus; NCA 2000/223.
Diagnosis. e body proportions of this species are dierent than in other Pachy-
ballus spp., namely width of carapace and width of abdomen are clearly greater than
their length. Sigilla are strongly marked. Shape of the eye eld is more trapezoid than
in congeners, its width at posterior eye row is a quarter larger than anterior width.
Redescription. Male. Measurements. Cephalothorax: length 1.5–1.7, width 1.9–
2.0, height 0.6. Eye eld: length 0.8–1.0, anterior width 1.3–1.5, posterior width
1.9–2.0. Abdomen: length 2.4–2.9, width 2.8–3.2.
General appearance as in Figs 98–102. Slightly larger than P. avipes, body attened,
covered with hard integument, pitted. Carapace wide, dark brown to blackish, vicinity
of eyes black. Eye eld strongly trapezoid, more than in other Pachyballus spp. Its width
at last row of eyes is a quarter larger than its anterior width. A few hairs and long bristles
at anterior eyes. Clypeus low. Chelicerae with short fang, three small teeth on promargin
and four teeth with fused base on retromargin (Fig. 110). Mouth parts brown, sternum
Figures 92–97. Pachyballus ornatus sp. nov., internal structure of epigyne 92, 94, 95, 97 ventral view,
93, 96 dorsal view (specimens 94–96 from Usambara Mts, lower row – black form).
Revision of Pachyballus and Peplometus 75
oval, dark. Abdomen heart-shaped, wide, clearly wider than long. Colouration of abdo-
men usually light, orange brownish with blackish edge, sometimes yellowish or brown.
Sigilla clearly visible. Venter brown, with typical scuta (Fig. 98). First pair of legs brown,
slightly larger than other pairs, with femora and tibiae thickened, tibiae slightly attened
dorsally. All legs brownish with lighter tarsi. Palps dark, structure as in Figs 103, 111, 112.
Female. Measurements. Cephalothorax: length 1.4–1.6, width 1.6–2.0, height
0.7. Eye eld: length 0.7–0.8, anterior width 1.2–1.5, posterior width 1.8–2.0. Abdo-
men: length 2.4–2.9, width 2.8–3.2.
General appearance as in Figs 104–106. Similar to male, eye eld strongly trap-
ezoid. Abdomen wide, rounded, with visible sigilla. All legs yellow, sometimes femora,
patellae and tibiae brownish. Palps dark. Epigyne typical, with horseshoe-shaped an-
terior depression (Figs 107–109, 113–116). Copulatory ducts long, forming several
loops, spermathecae strongly sclerotised (Figs 118–122).
Immature specimens. Shape of body as in adults, abdomen with two oval scuta,
close to each other on dorsum (Figs 191, 192). Ventral scuta absent.
Figures 98–103. Pachyballus transversus, male 98 habitus, ventral view 99–102 habitus, dorsal view,
103 palpal organ, ventral view (all specimens from Congo).
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Distribution. Widely distributed in Africa (Fig. 196).
Remarks. Simon (1900) described only the female of P. transversus. He recorded
this species from Congo, Transvaal (South Africa) and Zanzibar (Simon 1900, 1901).
He found it also (Simon 1910) later in Guinea Bissau and noted in this publication that
this species had been described from Congo. is suggests that the Congolese specimens
constitute the “type material”. However, the two specimens from Congo, personally la-
belled by Simon and kept in the MNHN collection, are males. e fact was also men-
tioned by Berland and Millot (1941). Simon most probably described these specimens
and apparently published the wrong data on sex of specimens. is description is very
supercial and the structure of copulatory organs is not depicted. ere are also some
other males from Zanzibar in Simon’s collection in MNHN (but no female). Simon’s
specimens from Natal, kept in MZC (examined) are immature and probably misidenti-
ed. us, it should be concluded that Simon’s description concerned the male and the
female of this species is described here for the rst time. e gure in Simon (1901)
shows characteristic body proportions of P. transversus (width of carapace greatly exceeds
its length, eye eld is clearly trapezoid with long distance between eyes in the posterior
row, and abdomen is rounded), which allows the proper species recognition.
Figures 104–109. Pachyballus transversus, female 104–106 habitus, dorsal view 107–109 epigyne (dis-
sected from specimens visible above) (104, 106 specimens from Congo 105 specimen from Ethiopia).
Revision of Pachyballus and Peplometus 77
Pachyballus variegatus Lessert, 1925
Figures 123–128, 193
Pachyballus variegatus Lessert 1925: 437, f. 10–14 (♂♀).
Syntypes. T • 1♂ 1♀; Kilimanjaro, Kibonoto; 3°11'S, 37°06'E; Sjöstedt leg.;
NHRS; examined.
Diagnosis. is species is the only one in the genus that does not have the poste-
rior ventral scutum on abdomen. Its other diagnostic feature is the pattern on abdo-
men; the anterior one-fourth of the abdomen is bright, the posterior part dark with a
wide, light median patch (Figs 123, 127).
Redescription. Male. Measurements: Cephalothorax: length 1.4, width 1.3,
height 0.4. Eye eld: length 0.7, anterior width 1.1, posterior width 1.3. Abdomen:
length 1.8, width 1.7.
General appearance as in Fig. 123. Body attened, integument strongly sclerotis-
ed, clearly pitted. Eye eld light, yellowish orange, blackish rings around eyes, thoracic
Figures 110–116. Pachyballus transversus 110 cheliceral dentition 111 palpal organ, ventral view 112
palpal organ, lateral view 113–116 epigyne (114 specimen from Ethiopia 115 specimen from Somalia
113, 116 specimens from Congo).
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part light brown anteriorly, posterior slope yellowish. A few delicate hairs at anterior
eyes. Chelicerae brown, with three teeth on promargin and a tooth with two tips on
retromargin (Fig. 124). Labium and endites basally brownish, with light tips. Sternum
light brown. Abdomen heart-shaped, widest anteriorly and tapering, anteriorly orange,
sides brown with wide orange area in the middle (Fig. 123). Venter yellowish, without
scutum. Spinnerets short, yellow. Legs relatively short, dark yellow (rst pair absent in
the syntype, only coxae extant, brown). Pedipalp (only right present) brown. Palpal
organ as in Figs 125, 126, embolic coil wide, with three loops.
Female. Measurements: Cephalothorax: length 1.5, width 1.4, height 0.5. Eye eld:
length 0.8, anterior width 1.2, posterior width 1.4. Abdomen: length 2.2, width 2.1.
Colouration as in male (Fig. 127). Abdomen rounded. Palp light, with brown api-
cal part. Epigyne weakly sclerotised with shallow central depression (Fig. 128). Internal
structure not studied, spermathecae probably large (visible through integument).
Remarks. Lessert (1925) wrote in the original description that the basic colour
of body was black. e specimens must have bleached heavily, however the outline of
lighter patches has been preserved.
Distribution. Known only from the type locality (Fig. 193).
Peplometus Simon, 1900
Peplometus Simon 1900: 399; 1901: 486.
Type species. Homalattus biscutellatus Simon, 1887.
Figures 117–122. Pachyballus transversus, internal structure of epigyne 117–120 ventral view 121,
122 dorsal view (117 specimen from Ethiopia 118 specimen from Somalia, other specimens from Congo).
Revision of Pachyballus and Peplometus 79
Diagnosis. Peplometus is closely related to Pachyballus. From the latter genus it can
easily be separated by the elongated abdomen (rounded in Pachyballus). It diers also in
the form of leg I in males having the leaf-like setae on tibia I (absent in the other genus).
Description. Small spiders (ca. 3.0–4.0 mm length), with very at body, cov-
ered with hard strongly sclerotised integument. Carapace almost square-shaped, pitted
dorsally, its length only slightly exceeds its width. Chelicerae with two (exceptionally
three) small teeth on promargin and on retromargin a saw-shaped tooth with four to
ve denticles. Abdomen slightly elongated, ratio of its length to width 1.3–1.5 (only
in female of P. biscutellatus 1.2). In males, width of abdomen almost equal to width
of carapace, abdomen of females wider and rather oval. Anterior margin of abdomen
straight. Hard scutum covers dorsum of abdomen, venter also strongly armoured, with
scuta as in Pachyballus, namely the narrow scutum with backwards extending “horns”
at anterior edge, and a large trapezoid scutum in posterior half of abdomen. Males
Figures 123–128. Pachyballus variegatus, syntypes 123 male, habitus124 cheliceral dentition 125 pal-
pal organ, ventral view 126 palpal organ, lateral view 127 female, colouration of abdomen 128 epigyne.
Wanda Wesołowska et al. / ZooKeys 944: 47–98 (2020)
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with a thinner sclerotised plate in front of epigastric furrow, its thick posterior margin
forms narrow wedge-shaped bar (Figs 138, 185). Numerous sclerotised bumps on sides
of the abdominal venter (Fig. 138). Legs short, in males the rst pair slightly thicker,
with enlarged femora and modied tibiae. In some species conspicuously thickened,
usually attened, in other species elongated (in this case the metatarsus I with a dorsal
process). Tibiae I black, with ventral brush of long dense attened setae, contrasting
with other segments. Copulatory organs similar to those in Pachyballus.
Key to Peplometus males
1 Metatarsus I with a dorsal process, tibia not thickened ................................ 2
– Metatarsus I without a process, tibia strongly thickened..............................3
2 Tibia I with a distal process, metatarsus shorter than tibia (Fig. 134) ...P. oy o
– Tibia I without process, length of metatarsus equal to length of tibia (Fig.
135) ................................................................................................... P. nimba
Figures 129–135. First leg of male 129 Pachyballus castaneus 130 Peplometus biscutellatus 131, 132 P.
chlorophthalmus 133 P. congoensis 134 P. oyo 135 P. nimba 132 retrolateral view, others prolateral view.
Revision of Pachyballus and Peplometus 81
3 Tibia I not attened (Fig. 133) ................................................ P. congoensis
– Tibia I attened .......................................................................................... 4
4 Tibia I attened dorsally (Fig. 131) ................................ P. chlorophthalmus
– Tibia I attened prolaterally (Fig. 130) .................................P. biscutellatus
Peplometus biscutellatus (Simon,1887)
Figures 4, 130, 136–153, 189, 190, 197
Homalattus biscutellatus Simon 1887: 263 (♀).
Peplometus biscutellatus Simon 1901: 486; Berland and Millot 1941: 398.
Holotype. I C • ♀; Assinie; 5°08'N, 3°16'W; Alluaud [C.] leg.; MNHN
9072; examined.
Other material examined. C • 2♀; Matube, Tiko plantation; 4°04'N,
9°21'E; 24.IV-6.V.1949; B. Malkin leg.; CAS • 1♀; Mabete Victoria [Limbe]; 4°01'N,
9°13'E; 24.V-7.VI.1949; B. Malkin leg.; CAS • 1♀; Reserve Forestiere Nyong, S
to Makak; 3°25'N, 12°47'E; 17.II.1950; J. Dahl and J. Birket-Smith leg.; UCZM.
G • 2♂; Kakum forest; 5°25'N, 1°19'W; 18.XI.2005; primary forest, R. Joc-
qué, D. De Bakker and L. Baert leg.; fogging; MRAC 217 897 • 1♀; the same data;
16.XI.2005; MRAC 217 875 • 2♀; the same data; 25.XI.2005; MRAC 217 945 • 7♂
4♀; the same locality; 15.XI.2005; secondary forest; MRAC 217 865 • 2♂; the same
data; 17.XI.2005; MRAC 217 881 • 1♂; the same data; 12.XI.2005; MRAC 217 857.
G • 1♀; Dalaba; 10°42'N, 12°15'W; VII.1937; J. Millot leg.; MNHN • 1♂;
Nimba Mts, Zougué Valley, near Gbakoré mine camp; 7°34'N, 8°28'W; 5.X.2011;
780 m a.s.l.; young secondary gallery forest, fogging canopy; D. van den Spiegel leg.;
MRAC 238 187 • 1♂; same locality, Seringbara road; 8.II.2012; beating; A. Henrard,
C. Allard, P. Bimou, M. Sidibé leg.; MRAC 239 050 • 2♂, 1♀; same locality; gallery
forest of Zié; 3.X.2011; 1250 m a.s.l.; fogging, canopy forest, understory shrub layer;
D. van den Spiegel, A. Hernard leg.; MRAC 238 066. I C • 1 imm.; together
with the holotype • 1♂; Bouaké, Foro Foro; 7°49'N, 5°01'W; 5–7.VIII.1974; pitfall;
G. Couturier leg.; MRAC 216 442 • 1♂; the same data; MRAC 216 380. N
• 1♂; Kabba; 7°50'N, 6°04'E; 18–23.II.1949; B. Malkin leg.; CAS. S • 1♂;
Dakar; 8°19'N, 0°13'W; IX.1947; L. Berland leg.; MNHN. S L • 2♂ 3♀
2 imm.; Free Town; 8°30'N, 13°15'W; col. E. Simon, MNHN 19 988. T • 1♀ 1
imm.; without precise locality; col. C.F. Roewer (nr 10861); SMF.
Diagnosis. e male of this species may be distinguished by a at area on prolat-
eral side of the tibia I (Figs 130, 140, 141). e female has a cordiform abdomen, while
other congeners have oval abdomen; the width/length ratio of abdomen in P. biscutel-
latus female is 0.8, whereas it is 0.75 in other species.
Redescription. Male. Measurements. Cephalothorax: length 1.1–1.7, width 1.2–
1.8, height 0.5–0.6. Eye eld: length 0.7–0.9, anterior width 1.0–1.3, posterior width
1.2–1.6. Abdomen: length 1.9–2.9, width 1.3–2.1.
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General appearance as in Figs 136, 137. Body attened, completely covered with
strongly sclerotised and pitted integument. Dorsum of body dark brown, clothed in
dense short white hairs (lost in some of the studied specimens). Eye eld on more than
half of carapace length, black rings around eyes, the rst row of eyes encircled with white
hairs. Long white bristles between anterior eyes. Clypeus extremely low, brown with a few
white hairs. Chelicerae with three diminutive teeth on promargin and ve retromarginal
Figures 136–142. Peplometus biscutellatus, male 136 habitus, dorsal view 137 habitus, lateral view
138venter of abdomen 139 cheliceral dentition 140 rst leg, dorsoprolateral view 141 rst leg, dorsal view
142 palpal organ, ventral view (photos – specimen from Ghana, drawings – specimen from Sierra Leone).
Revision of Pachyballus and Peplometus 83
teeth fused basally (Fig. 139). Mouthparts brown. Sternum oval, brown. Abdomen elon-
gated, widest anteriorly and tapering rearward, anterior edge almost straight. Dorsum of
abdomen totally covered by very strongly sclerotised and pitted scutum, laterally slightly
turned back to venter. Venter with typical scuta; anterior scutum narrow, extending back-
wards laterally, posterior ventral scutum largest, cordiform. In front of epigastric furrow
a thin sclerotised plate with thick posterior margin forms narrow wedge-shaped bar. Nu-
merous bumps on sides ventrally; armour-plate pattern is shown in Fig. 138. Spinnerets
brown, obscured by dorsal scutum. Legs I the biggest, femora enlarged, brownish with
darker prolateral side, three short spines on dorsum. Tibia thickened, clearly attened
prolaterally, blackish with pale dorsal streak, ventrally black brash of dense long feather-
shaped setae (Figs 130, 140), metatarsi and tarsi short, yellowish, two pairs of ventral
spines on metatarsi. Other legs pale yellowish with dark marks at bases of segments,
only femora IV brown (or yellow with brown streak on prolateral side). Palps light, their
structure as in Fig. 142. Palpal tibia short with single thin straight apophysis, bulb oval,
embolus thin, long, spirally coiled, forming three loops on bulb tip.
Female. Measurements: Cephalothorax: length 1.0–1.4, width 1.2–1.4, height
0.5–0.6. Eye eld: length 0.6–0.8, anterior width 1.0–1.2, posterior width 1.2–1.3.
Abdomen: length 2.1–2.3, width 1.7–1.9.
General appearance as in Figs 143–145. Similar to male, but abdomen wider,
heart-shaped. Scarce white hairs on body. Carapace dark brown, black near eyes. Legs
I as in males, not so thick, tibia slightly thickened, retrolateral attening hardly visible.
Palps dark, blackish. Epigyne large, rectangular, weakly sclerotised, with small semi-
circle shallow depression divided by median septum (Figs 146–149). In some females
horizontal crevices laterally, formed by microsculpture of integument, not part of epi-
gyne (Fig. 146). Copulatory ducts very long, spirally coiled behind openings, form
several loops distally, spermathecae bean-shaped (Figs 150–153).
Immature specimens. Abdomen not elongated, heart-shaped, with two oval dor-
sal scuta on abdomen, close to each other (Figs 189, 190).
Remarks. e rst description of the male is given here. Simon described P. bis-
cutellatus based on a female from Ivory Coast, however in his samples from Sierra Le-
one the two sexes were present (labelled by Simon himself). is fact has already been
mentioned by Berland and Millot (1941), but they have not described the missing
male. Material collected by Simon is in a very poor condition. Numerous specimens
have lately been collected by fogging, this species typically inhabits canopy.
Distribution. West equatorial Africa (Fig. 197).
Peplometus chlorophthalmus Simon, 1900
Figures 131, 132, 154–171, 197
Peplometus chlorophthalmus Simon 1900: 399 (♂); 1901: 482, f. 566–569.
Holotype. S A • ♂; Natal (eastern SA); C. M[artin] [leg.]; MNHN 17
385; examined.
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Other material examined. S A • 1♀ 5 imm.; together with the holotype.
C D.R. • 1♂ 1♀ 1 subad. ♂; Bas Congo, Mayombe, Luki Forest Biosphere Reserve;
5°40'S, 13°10'E; 7.XI.2006; D. De Bakker and J.P. Michiels leg.; beating; MRAC 221
505 • 1♀; the same data; 14.XI.2006; MRAC 219 997 • 1♀; the same data; 8.XI.2006;
MRAC 219 944 • 2♀; the same data; 14.XI.2006 and 19.IX.2007; MRAC 226 100 •
1♂ 1♀; the same locality; 24.IX.2007; old secondary rainforest, fogging; MRAC 226
104 • 1♀; the same locality; 29.IX.2007; primary rainforest, fogging; MRAC 226 110 •
2♀; the same data; 5.XI.2006; MRAC 226 118 •1♀; the same data; 11.XI.2006; MRAC
221 583 • 1♀; the same data;12.XI.2006; MRAC 220 954.
Figures 143–145. Peplometus biscutellatus, female 143, 144 habitus, dorsal view 145 habitus, ventral
view (143 specimen from Cameroon 144, 145 specimen from Ghana).
Revision of Pachyballus and Peplometus 85
Diagnosis. e tibia of the leg I in male is strongly attened dorsally (Figs 131,
158). e female is dicult to distinguish from P. biscutellatus, but it has a narrower
abdomen (see Diagnosis of the latter species).
Redescription. Male. Measurements. Cephalothorax: length 1.0–1.5, width 1.1–
1.3, height 0.5–0.6. Eye eld: length 0.6–0.8, anterior width 1.0–1.2, posterior width
1.2–1.3. Abdomen: length 2.2–2.3, width 1.4–1.5.
General appearance as in Figs 154–156. Small spider with attened body. Cara-
pace dark brown to blackish, pitted. White bristles around anterior median eyes and
between all eyes of rst row. Clypeus low, black, with a few white hairs. Fang of cheli-
cerae short, two small teeth on promargin, retromarginal tooth with four tips (Fig.
160). Sternum oval, dark brown. Abdomen narrow, covered with strongly sclerotised,
pitted, dark integument. Venter with typical scuta (same as by P. biscutellatus). Legs
yellowish with dark lines along femora and tibiae III and IV on prolateral surface. Legs
I the stoutest, tip of prolateral side of femur with small patch, tibia strongly thickened,
black with yellow line along dorsum and large light patch on retrolateral side, long
attened black setae ventrally (Figs 157, 159). Tibia I characteristic for having a large
attened part on dorsal side (Figs 131, 158). Pedipalp light, its structure as in Figs
161–164, embolic coil wide.
Figures 146–149. Peplometus biscutellatus, female, epigyne (upper row specimens from Sierra Leone,
lower row specimens from Ghana).
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Female. Measurements. Cephalothorax: length 1.0–1.4, width 1.1–1.3, height
0.5–0.6. Eye eld: length 0.6–0.7, anterior width 1.0–1.2, posterior width 1.2–1.3.
Abdomen: length 2.0–2.4, width 1.5–1.8.
Similar to male, general appearance as in Figs 166, 167. White hairs at anterior
eyes and on posterior carapace slope. Abdomen wider than in male, but relatively
narrow, narrower than in P. biscutellatus. Legs darker than in male. Femora of I–III
legs brown with yellow ventral surface, femora IV completely brown. Patellae IV with
pro- and retrolateral brown stripes. First leg not larger, its tibia black with black long
setae ventrally, in some specimens with narrow light streak along dorsum. Tibiae II
with prolateral brown stripe, tibiae IV brown with thin yellow longitudinal stripes
dorsally and ventrally. Other leg segments light yellow. Palps blackish. Epigyne as in
Figs 168, 169, rectangular with shallow depression. Ventral structure of epigyne simi-
lar to other species, copulatory ducts long, weakly sclerotised in initial part, forming
several loops (Figs 170, 171).
Immature specimens. Abdomen not elongated, heart-shaped, with two oval scuta
on dorsum, close to each other, not covering whole dorsum of abdomen (Fig. 165).
Remarks. e rst description of the female is given here. Simon described only
the male of P. chlorophthalmus, although the vial with a type specimen contains also an
Figures 150–153. Peplometus biscutellatus, female, internal structure of epigyne 150, 151 ventral view
152, 153 dorsal view (150, 151 specimen from Ghana 152 specimen from Cameroon 153 specimen
from Sierra Leone).
Revision of Pachyballus and Peplometus 87
undescribed female and a few immature specimens. is material is however in a very
poor condition.
Length of the apical part of embolus varies. It is very long in South African speci-
men, extending beyond the retrolateral edge of apical part of cymbium (Figs 161,
162), and short in specimen from Congo D. R. (Figs 163, 164).
Distribution. Known from Congo and South Africa (Fig. 197).
Figures 154–159. Peplometus chlorophthalmus, male 154 habitus, dorsal view 155 habitus, ventral view
156 habitus, lateral view 157 rst leg, dorsal view 158 rst leg, prolateral view 159 rst leg, retrolateral
view (specimen from Congo).
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Peplometus congoensis sp. nov.
http://zoobank.org/41E2658D-9A97-49CA-923F-E8A3935F9795
Figures 133, 172–183, 198
Holotype. C • ♂; Brazzaville, ORSTOM Park; 4°16'S, 15°17'E; 19.X.1963; J.
Balogh and A. Zicsi leg.; HNHM.
Paratypes. C • 1♀; together with the holotype. C D.R. • 1♀; May-
ombe, Bas Congo, Luki Forest Reserve; 5°37'S, 13°05'E; 28.IX.2007; D. De Bakker
and J.P. Michiel leg.; primary rainforest, canopy fogging; MRAC 226 108.
Figures 160–165. Peplometus chlorophthalmus 160 cheliceral dentition 161, 163 palpal organ, ventral
view 162, 164 palpal organ, lateral view 165 immature specimen (160–162 holotype 163, 164 speci-
men from Congo).
Revision of Pachyballus and Peplometus 89
Diagnosis. e most characteristic feature of this species is colouration and shape
of the rst pair of legs. Tibia I in male is totally black, it is not attened (Fig. 172),
whereas in congeners tibia is attened dorsally or laterally and has a light streak or
patch. Tibia I of the female is light retrolaterally, with large dark patch at its basis (Figs
179, 180). e abdomen of female is narrow, similar as in male, while females of other
species have abdomen wider than males.
Figures 166–171. Peplometus chlorophthalmus, female 166 habitus, dorsal view 167 habitus, ventral
view 168, 169 epigyne 170 internal structure of epigyne, ventral view 171 internal structure of epigyne,
dorsal view (171 specimen from Natal, other specimens from Congo).
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Etymology. e species is named after its terra typica (Congo).
Description. Male. Measurements. Cephalothorax: length 1.7, width 1.8, height
0.6. Eye eld: length 0.9, anterior width 1.3, posterior width 1.7. Abdomen: length
3.0, width 2.2.
General appearance as in Figs 172, 173. Body attened, integument strongly sclero-
tised and clearly pitted. Carapace trapezoid, widest posteriorly, dark brown, black around
eyes, a few bristles at eyes of rst row. Chelicera with two small teeth on promargin and
four-tip retromarginal tooth. Mouth parts and sternum light brown. Abdomen elongat-
ed, only slightly wider than carapace, venter also covered by scuta, posterior scutum large
(Fig. 173). Legs I stout, brown, lateral side of femur slightly darker, tip of patella dark,
tibia strongly thickened, black, with long dense black feather-shaped setae ventrally (Figs
133, 172). Other legs yellowish brown, with distal small dark brown patches on patellae
and tibiae II–III and proximal brown rings on patellae, tibiae and metatarsi IV, femora
IV brown. Palpal organ as in Figs 177, 178, apical part of embolus long.
Female. Measurements. Cephalothorax: length 1.2–1.3, width 1.1–1.3, height 0.4–
0.5. Eye eld: length 0.7, anterior width 1.0–1.1, posterior width 1.1–1.3. Abdomen:
length 2.6–3.0, width 1.8–1.9.
General appearance as in Figs 174–176. Carapace rounded, brown to black, strongly
sclerotised and clearly pitted, eye eld on half of its length. Some dark bristles at eyes of
rst row. Chelicerae yellowish brown, with short fang, two small teeth on promargin and
three-tip tooth on retromargin. Mouth parts creamy. Sternum oval, blackish. Abdomen
relatively narrow, elongated, widest anteriorly, anterior edge almost straight. Dorsum
of abdomen totally covered by very strongly sclerotised, pitted, black scutum, turned
back on its margin. Abdomen ventrally with hard scuta; anterior scutum narrow along
anterior margin of abdomen, extending backwards, posterior scutum large, trapezoid.
Numerous bumps on sides of abdomen venter (Fig. 175). Spinnerets dark. Legs gener-
ally light, creamy with blackish stains. Legs I the biggest (not so markedly as in males);
femora with black patch distally on prolateral side, tibia thickened, prolaterally black,
basal half of retrolateral part black (Figs 179, 180). Dense long black setae on ventral side
of tibia, two pairs of ventral spines. Dark line along dorsum of femora IV. Palps blackish.
Epigyne large, rectangular, with small depression divided by median ridge, hardly visible
horizontal crevices laterally (Fig. 181). Internal structure as in Figs 182, 183.
Distribution. Known only from Congo and DR Congo (Fig. 198).
Peplometus nimba sp. nov.
http://zoobank.org/30CA27D6-8CC9-4960-B63C-9517D696F20A
Figures 135, 184–188, 198
Holotype. G • ♂; Nimba Mts, Nion; 7°36'N, 8°28'W; 16.VI.1942; M. Lamotte
leg.; BMNH.
Diagnosis. Male of this species is very similar to that of Pachyballus oyo, it can be
distinguished by the lack of a conspicuous, sharp process on tibia I, which is very char-
acteristic for the latter species (compare Fig. 135 with Fig. 134). In P. nimba metatarsus
Revision of Pachyballus and Peplometus 91
and tibia of leg I are of equal length, while in P. o y o metatarsus I is clearly shorter, about
half of tibia I length.
Etymology. e specic name is a noun in apposition referring to the Nimba Mts,
type locality of this species.
Description. Male. Measurements. Cephalothorax: length 1.4, width1.2, height
0.5. Eye eld: length 0.7, anterior width 1.1, posterior width 1.2. Abdomen: length
2.0, width 1.4.
General appearance as in Fig. 184. Body very at, covered with strongly sclerotised
integument, clearly pitted, brown with blackish area around eyes. Carapace slightly trap-
Figures 172–176. Peplometus congoensis sp. nov. 172 male, holotype, habitus, dorsal view 173 male,
habitus, ventral view 174 female, habitus, dorsal view 175 female, habitus, ventral view 176 female,
habitus, lateral view.
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92
ezoid, white bristles near eyes of rst row. Clypeus very low. Chelicerae with two
teeth on promargin and ve on retromargin (apical tooth very small), fang short.
Endites, labium and sternum yellow. Abdomen elongated, shield shaped (its ante-
rior margin almost straight), dorsum covered with large strongly sclerotised scutum.
Ventrally abdomen with typical large scuta (Fig. 185), as in other species. In front of
epigastric furrow clearly visible narrow wedge-shaped sclerotised swelling. Spinnerets
yellow. First pair the stoutest, femora basically yellow, slightly darker on sides; patella
light yellow, tibia slightly thickened, brown (prolateral surface darker), with light
streak dorsally, dense long whitish (probably bleached) setae ventrally; metatarsus
long, creamy with black dorsal hump and black line along prolateral side (Fig. 187).
Legs II–IV yellow, with thin dark line along prolateral sides of patellae and tibiae.
Legs without spines, except two short ventral spines on metatarsus I. Palps yellow,
tibial apophysis very thin, bulb triangular, embolus spirally coiled on bulb tip (Fig. 188).
Figures 177–183. Peplometus congoensis sp. nov. 177 palpal organ, ventral view 178 palpal organ, lateral
view 179 rst leg of female, prolateral view 180 rst leg of female, retolateral view 181 epigyne 182 in-
ternal structure of epigyne, ventral view 183 internal structure of epigyne, dorsal view.
Revision of Pachyballus and Peplometus 93
Female unknown.
Distribution. Known only from the type locality, Nimba Mts in western Africa
(Fig. 198).
Peplometus oyo (Wesołowska & Russell-Smith, 2011), comb. nov.
Figures 134, 198
Pachyballus oyo Wesołowska and Russell-Smith 2011: 588, f. 126–135, 232–234 (♂♀).
Figures 184–188. Peplometus nimba 184 male, holotype, habitus, dorsal view 185 venter of abdomen
186 rst leg, retrolateral view 187 rst leg, prolateral view 188 palpal organ, ventral view.
Wanda Wesołowska et al. / ZooKeys 944: 47–98 (2020)
94
Diagnosis. e male of this species is similar to that of Peplometus nimba, but it can
be identied by the presence of a big apical process on the rst pair of legs and the
metatarsus of this pair shorter than tibia (Fig. 134).
Description. For description of both sexes see: Wesołowska and Russell-Smith
(2011).
Distribution. Southern Nigeria (Fig. 198).
Figures 189–192. Dorsal scuta of abdomen, immature specimens. 189, 190 Peplometus biscutellatus
191, 192 Pachyballus transversus 189, 191 younger instar 190, 192 subadult.
Revision of Pachyballus and Peplometus 95
Mimicry
e mimicry is a common phenomenon among spiders (Pocock 1909). Jumping
spiders imitate various models, mainly ants (overview in Cushing 1997, 2012),
but also pseudoscorpions (Platnick 1984), ies (Morrison 1981), wasps (Reiskind
1976, Żabka 1992), velvet wasps (Edwards 1984, Wesołowska 2009) or caterpillars
(Logunov and Obenauer 2019). Some salticid genera resemble beetles (Cloudsley-
Figures 193–198. Distribution maps of Pachyballus and Peplometus species. 193 circle – Pachyballus
caelestis, square – P. castaneus, triangle – P. variegatus 194 P. avipes 195 circle – P. miniscutulus, square–
P.mombasensis, triangle – P. ornatus 196 P. transversus 197 circle – Peplometus biscutellatus, square – P.chlo-
rophthalmus 198 circle – P. nimba, square – P. o y o , triangle – P. congoensis.
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96
ompson 1995) e.g. Cylistella Simon, 1901, Coccorchestes orell, 1881, Planiemen
Wesołowska and van Harten, 2007. e Ballini genera Pachyballus Simon, 1901
and Peplometus Simon, 1901 also resemble beetles, most probably from the family
Chrysomelidae, which is expressed in their body shape and a very strongly sclerotised
integument. e microsculpture of the integument and bright or iridescent colour-
ing emphasise this resemblance. In juveniles of all Peplometus species and Pachyballus
transversus the two dorsal scuta on the abdomen are arranged in a pattern similar to
elytra of beetles (Figs 165, 189–192), it is the unique feature among spiders. is
type of resemblance in spiders usually suggests a Batesian type of mimicry. On the
other hand, the two presented genera live in the canopy and their body type (small,
attened, colours often matching their preferred habitats) may suggest that it is a
type of camouage. e resemblance to other tree-living arthropods could be just a
result of a shared cryptic body pattern.
Acknowledgements
We wish to express our warmest thanks to curators of museum collections for giving
access to spiders used in this study. Suresh P. Benjamin, Yuri M. Marusik, and Wayne
P. Maddison are thanked for their valuable comments on the earlier draft of this article.
is work was partly supported by the Federal Fundamental Scientic Research Pro-
gramme for 2013–2020 (No. АААА-А16-116121410121-7) to GA.
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