8181 Croeser 1996 Ann 37 1 122

Ann. Natal Mus. Vol.
37 Pages 1-122 Pietennaritzburg October, 1996
A revision of the African huntsman spider genus Palystes L. Koch,

1875 (Araneae: Heteropodidae)l
by
P. M. C. Croeser
(Natal Museum, P. Bag 9070, Pietennaritzhurg 3200, and University of Natal, P. Bag
XOl, Scottsville 3209, South Africa)
ABSTRACT
Palystes L. Koch, 1875, type genus of Palystinae (Heteropodidae), is revised and a cladogram
proposed for the 15 species recognised, including 6 new species. A key to the species is provided.
Palystes occurs in eastern and southern Africa. Patterns of biogeography and relationships in a redefined
Palystinae, compared with sequences of hypothesised vicariance events during the evolution of the
African landscape, suggest a southern African, minimum late Miocene-Pliocene origin for the genus.
Three species groups, defined by markings and forms of male and female genitalia, are recognised:
castaneus group (western Cape Province of South Africa - 3 species), lunafus group (eastern and interior
western Cape Province of South Africa, Lesotho and Transkei - 6 species); superciliosus group (eastern
and southern Africa - 6 species). Seventeen nominal species previously attributed to Palystes, including
I I from the Indo-Australasian region, are transferred to other genera, including a new genus,
Parapalystes, which is also described. These genera are discussed; annotated lists are provided for the
African genera and for a resurrected Oceanian genus, Gnathopalystes Rainbow, 1899. Definitions of the
family Heteropodidae and three subfamilies (Palystinae, Heteropodinae and Staianinae) affected by the
revision, are reviewed.
New genus: Parapalystes (type species: Parapalystes euphorbiae sp. n. from the north-western Cape,
South Africa).
Resurrected genus: Gnathopalystes Rainbow, 1899 (type species: Gnathopalystes Jerox Rainbow,
1899, from Santa Cruz, Oceania), previously synonymised under Palystes.
Valid, named Palystes species: P. castaneus (Latreille, 1819); P. superciliosus L. Koch, 1875; P.
hoehne Ii Simon, 1890; P. lunatus Pocock, 1896; P. johnstoni Pocock, 1896; P. ellioti Pocock, 1896; P.
perornatus Pocock, 1900; P. leppanae Pocock, 1902; P. crawshayi Pocock, 1902.
New species: Palystes stilleri, P. martinfilmeri, P. karooensis, P. stuarti, P. ansiedippenaarae, P.
leroyorum and Parapalystes euphorbiae, all from South Africa.
New synonyms: Palystes chapen' Simon, 1880 = P. castaneus (Latreille, 1819); P. natalius (Karsch,
1878), P. spenceri Pocock, 1896, P. modificus Strand, 1906, and P. superciliosus var. Jasciiventris
Strand, 1907 = P. superciliosus L. Koch, 1875; P. kibonotensis Lessert, 1921, and P. bornemiszai
Caporiacco, 1947 = P. hoehneli Simon, 1897; P. amanicus Strand, 1907 and P. affinis Lessert, 1921 = P.
ellioti Pocock, 1896; Palystes incallus Thorell, 1890 = Gnathopalystes kochi (Simon, 1880).
New combinations: Palystes speciosus Pocock, 1898, P. dasyurinus Hogg, 1914 and P. ledleyi Hogg,
1922, transferred to Heteropoda Latreille, 1804; Palystes pilipodus Strand, 1913, transferred to
Anchonastus Simon, 1898; Palystes scutatus Pocock, 1902, transferred to Panaretella Lawrence, 1937;
Palystes ignicomus L. Koch, 1875, P. crucifer Simon, 1880, P. kochi Simon, 1880, P. rutilans Simon,
1899, P. nigriventer Ku\czynski, 1910 and P. nigrocornutus Merian, 1911, transferred to Gnathopalystes
Rainbow, 1899; Palystes cultrifer Pocock, 1900, P. lycosinus Pocock, 1900, P. whiteae Pocock, 1902,
and Remmius megacephalus (C. Koch, 1845), transferred to Parapalysfes gen. n.
Neotype designations: The ho1otype of Thomisus castaneus Latreille, 1819, is designated the neotype
for Palystes frenafus L. Koch, 1875, making them objective synonyms; an adult female from the type
locality (Maseru, Lesotho) is designated as neotype for Palystes crawshayi Pocock, 1902.
'This paper is based on a dissertation submitted to the University of Natal, Pietermaritzburg, in partial fulfilment of the
requirements for a M.Sc degree in the Dept. of Zoology & Entomology, under the supervision of Prof. D. J. Brothers.
2 ANNALS OF THE NATAL MUSEUM, VOL. 37 1996
Lectotype designations: Palystes superciliosus L. Koch, 1875; P. cultrifer Pocock, 1900; P. lycosinus
Pocock, 1900; P. ellioti Pocock, 1896; P. johnstoni Pocock, 1896; P. rutilans Simon, 1899, P. peromatus
Pocock, 1900; P. kibonotensis Lesser!, 1921; Gnathopalystesferox Rainbow, 1899.
Species incertae sedis: Palystes pinnotherus (Wa1ckenaer, 1837); P. fornasinii (pavesi, 1881); P.
flavidus Simon, 1897; P. reticulatus Rainbow, 1899; P. spiralis Strand, 1907 and P. convexus Strand,
1907.
CONTENTS
Introduction ................................................................................................................... 2
Materials and Methods .................................................................................................. 5
Natural History of Palystes ........................................................................................... 7
Subfamily Palystinae ................................................................................................. 10
Key to genera of Palystinae ........................................................................................ 13
Palystes L. Koch, 1875 ............................................................................................... 13
Key to species of Palystes s .s..................................................................................... 18
Palystes castaneus species group ................................................................................ 25
Palystes lunatus species group .................................................................................... 36
Palystes superciliosus species group ......................................................................... .55
Palystes species incertae sedis .................................................................................... 83
Parapalystes gen. n ..................................................................................................... 85
Panaretella Lawrence, 1937 .......................................................................................91
Anchonastus Simon, 1898 ........................................................................................... 92
Cladistics of Palystinae and Palystes ..........................................................................93
Biogeography of Palystinae and Palystes ................................................................. 102
Subfamily Heteropodinae ........... ............................................................................ 107
Gnathopalystes Rainbow, 1899, gen. resurr.............................................................. 110
Heteropoda Latreille, 1804 ....................................................................................... 113
Subfamily Staianinae .............................................................................................. 115
Acknowledgements ................................................................................................... 116
References ................................................................................................................. 117
Index to Genera and Species ..................................................................................... 122
INTRODUCTION
This is a contribution to the revision of the heteropodid subfamily Palystinae. It
concentrates on the taxonomic revision, cladistics, biogeography and natural history
of the largest genus, its type genus Palystes L. Koch, 1875, a group of large, robust
spiders (Fig. 1), previously with some 38 attributed African and Indo-Australasian
species. It also includes discussions on the three subfamilies affected by this revision:
Palystinae, Heteropodinae and Staianinae. Annotated species lists for all genera
affected by new combinations (with the exception of the very large genus
Heteropoda Latreille, 1804) are also provided. In addition to the types of Palystes
species examined during this study, a wide range of other African and Indo-
Australasian specimens, including types of other genera, were studied to determine
generic limits and affinities. Notes on this material are included in the relevant
sections.
CROESER: REVlSION OF PALYSTES (ARANEAE: HETEROPODIDAE) 3
Fig. 1. Palystes castaneus (Latreille, 1819), female (Photograph by D. R. D'Ewes).
In this study Palystes is redefined to include only those species apparently forming
a monophyletic group with the type species, Palystes castaneus (Latreille, 1819). The
generic name has been retained for only 9 of the previously described species, all
from Africa, while a further 6 new species from South Africa are added. Twelve of
these species are endemic to southern Africa south of the Limpopo River. The
remaining three are distributed north of the Limpopo, through eastern Africa into
Rwanda and Uganda. The majority of these species are restricted to relatively small
areas. It is probable that further species await description from the Cape interior, the
relatively poorly collected regions north of the Zambezi River, and the disjunct
montane forests in eastern and central Africa.
Despite the size of the family (more than 700 described species in 82 genera), and
4 ANNALS OF THE NATAL MUSEUM. VOL. 371996
the prevalence of its members among the large spider fauna of most tropical and
subtropical regions, heteropodids are still relatively poorly known. Most genera and
species were described in the last century and there is considerable confusion in the
literature over the identity of many of the older genera, including Olios Walckenaer,
1837, and Sparassus Walckenaer, 1805 (also known as Eusparassus Simon). Even
the correct name of the family has still to be determined; it is presently known under
three names: Sparassidae Bertkau, 1872, Heteropodidae Thorell, 1873 and
Eusparassidae Jarvi, 1912. The oldest name, Sparassidae, was once the most
commonly used name, but is based on an unrecognisable type genus, Sparassus
Walckenaer, 1805. This unsatisfactory situation was reviewed by Platnick and Levi
(1973), who recommended that until the identity of Sparassus had been established,
the next oldest, available name, Heteropodidae, should be used instead.
Revisory work on the Heteropodidae is long overdue, as little has been done
subsequent to the publications of Simon (1897-1903), Hogg (1903), and Jarvi (1912,
1914). Latterly, however, Australasian heteropodids have been studied by Hirst
(1989a 1989b 1990 1991a 1991b), who has revised and described Deleninae and
added a new genus, Keiiira, to the Heteropodinae, and Todd-Davies (1994), who has
revised the Heteropoda species of Australia and added a new genus, Yzinthi. The
present revision clarifies the position of the Palystinae and extends the Australian
initiative.
Whether the heteropodids form a monophyletic group has still to be settled.
Lehtinen (1967), in his notes on the evolution of araneomorphs, tentatively linked the
Clubionidae with the heteropodids in a superfamily Sparassoidea, but suggested that
they might belong in the Amaurobioidea. Lehtinen (pers. comm.) doubts that
members of the European genus Micrommata Latreille belong in the same family as
the large tropical genera allied to Heteropoda Latreille. He suggests that closer
relatives of Micrommata may be found among the philodromids and clubionids. He
also questions whether many of the species ascribed to Olios, the largest heteropodid
genus, have been correctly assigned. About half of the heteropodid species described
have been placed in Olios.
A widely used synapomorphy for the family is possession of a dorsal trilobate
membrane at the metatarsal-tarsal joint, which extends tarsal movement. A further
synapomorphy has been suggested by Homann (1971) who maintains that the nature of
the pigmentation and a split rhabdome in the secondary eyes of heteropodids are unique
to the family. He describes different structures for the clubionids and philodromids.
This has still to be tested for all genera presently included in the Heteropodidae.
In addition to the two synapomorphies mentioned, heteropodids are presently
recognised as entelegyne araneomorph hunting spiders with laterigrade leg
arrangement, sharing the following combination of morphological characters: no
colulus or its homologue; a well-developed scopular brush of modified setae
ventrally on the metatarsus and tarsus of all legs; two pectinate tarsal claws
subtended by a dense tarsal claw scopula; cheliceral fang furrow with well-developed
retro- and anterolateral rows of teeth, the retromargin with two or more teeth; a
notched trochanter; and eight eyes in two straight, or nearly straight, rows of four,
anteriorly across the caput.
CROESER: REVISION OF PALYSTES (ARANEAE: HETEROPODIDAE) 5
MATERIALS AND METHODS

Specimens were studied and drawn with the aid of a Wild M5A stereo microscope
equipped with a drawing tube. Selected specimens were also examined with a JEOL
T-200 scanning electron microscope. Descriptions and measurements (in millimetres)
and ratios given are those of type specimens, unless otherwise stated. Illustrations
(unless otherwise indicated) are of the holotype and a paratype of the opposite sex.
References in the text to 'the Code' are to the International Code of Zoological
Nomenclature (1985).
Synonymy lists for each species include all specific names and generic
combinations, as well as any other pertinent nomenclatural changes, errors or
comments. Additional citations from the literature contained in Roewer (1954),
Bonnet (1958) and Brignoli (1983) are not repeated.
Species descriptions/redescriptions are arranged alphabetically within each of the
three species groups in the following order: group-name species, other existing
species, new species. Under material examined, data entry for a specimen or series of
specimens from a particular locality ends with the depository's acronym (and
sometimes accession number) in parenthesis, and a semicolon. If entries are
separated by a comma, this means that the localities are usually suburbs of a city
(followed by a colon) or some other form of subordinate category.
Cladograms were derived using Hennig86 version 1.5 (© James S. Farris 1988).
Genital morphology:
The terminology used here for the male palpal organ (Fig. 9) largely follows that
of Lamoral (1973) for Palystes castaneus, but differs in several respects. Differences
are: 1). the use of the term 'palpal organ' for the genitalic organ on the tarsus of the
adult male pedipalp instead of 'tarsal organ' (a term reserved for the small,
apparently chemo-sensory, organs described by Blumenthal (l~5) and found on all
tarsi of both sexes of most, if not all, spiders). 2) the use of the term 'conductor' for
the membrane arising from the tegular bulb (Fig. 9b, co) and which opposes the
embolus, instead of the term 'median apophysis'. 3). the use of the phrases 'tegular
flange' or 'lateral sclerite' for the flared tegular ridge (Fig. 9a, Is) flanking the
embolus, instead of 'conductor'. This is in keeping with presently accepted
terminology (e.g. Foelix 1982, Todd-Davies 1994 & pers. comm.). Terminology used
for the epigynum (Figs 8a, 8b) follows Foelix (1982) and the translation from
German of terms used by Jarvi (1912 1914) in his comparison of heteropodid
epigyna. The typical external structure of the epigynum (Fig. 8a) comprises a well-
developed septum (= fossa) ('ems'), separated from the lateral sclerites ('els') of the
epigynum by deep lateral invaginations ('Ii'). The invagination may extend to the
anterior of the epigynum (Fig. 89), with the median portion of the septum being
produced ventrally in a posteriorly projecting 'tongue' -shaped lobe (Fig. 68) or in a
transverse 'bridge' (Figs 77,81). Internally the invaginations support the afferent and
efferent sperm ducts and spermathecae (Fig. 8b). In some groups the invaginations
form a flanged support collar (Fig. 90).
Abbreviations and terminology used:
Abbreviations used are standard, but include the following: S. S. - sensu stricto (in
the strict sense, as redefined in this study); s. l. - sensu lato - (in the broad sense, as
previously understood in the literature); AME - anterior median eye(s); ALE -
anterior lateral eye(s); PME - posterior median eye(s); PLE - posterior lateral eye(s);
AER -anterior eye row; PER - posterior eye row; MOQ - median ocular quadrangle,
measured from outer margins of AME and PME; MOQP - posterior width of MOQ;
MOQL - length of MOQ; MOQA - anterior width of MOQ; CW - carapace width at
base of leg pair II; CL - carapace length; HW - head width; SWI - widest sternum
width, between coxae II; SW2 - anterior sternum width, between coxae I. Maxillae
refers to the pedipalpal coxae.
Total length was measured from front of the AME lenses to the tip of the abdomen.
Distances separating eyes were measured between nearest lens edges. All
measurements, unless otherwise specified, are in millimetres. Heteropodids are
unusually well-endowed with leg spines on most surfaces (30-33 spines on each leg).
Leg spination in species descriptions/redescriptions only records variations from the
schematic map (Fig. 7), modified from Griswold (1987a). Variations in spination are
given in the following sequence: (anterolateral: dorsal: retrolateral). For example: left
femur 1(3:2:3). Ventral spination, although recorded, was not usually variable. Leg
measurements are of the left hand side legs (unless otherwise stated). The cheliceral
dentition formula used describes teeth of the retromargin followed by teeth of the
anteromargin (i.e. 3:2).
Specimens studied or referred to are deposited in the following institutions.
Acronyms used follow Arnett et al (1986). Material labelled with the acronym
AMGS is kept at NMSA. Curators and/or loan officers are given in parenthesis:
AMGS Albany Museum, Grahamstown, South Africa (Dr F. W. Gess, Mr A.

Weaving)
AMNH American Museum of Natural History, New York, USA (Dr N. Platnick)
AMSA Australian Museum, Sydney, Australia (Dr M. R. Gray)
BMNH The Natural History Museum, London, UK (Mr F. Wanless, Mr P. Hillyard)
CASC California Academy of Sciences, San Francisco, USA (Dr W. Pulawski)
HNHM Hungarian Natural History Museum, Budapest, Hungary
ISNB lnstitut Royal des Science Naturelles de Belgique, Brussels, Belgium (Dr
L. Baert)
MCSN Museo Civico di Storia Naturale 'Giacoma Doria', Genoa, Italy (Dr G.
Arbocco)
MCZC Museum of Comparative Zoology, Cambridge, USA (Dr H. W. Levi)
MHNG Museum d'Histoire Naturelle, Geneva, Switzerland (Dr B. Hauser)
MNHN Museum National d'Histoire Naturelle, Paris, France (Dr M. Hubert, Dr J.
Heurtault)
MRAC Musee Royal de l' Afrique Centrale, Tervuren, Belgium (Dr R. Jocque)
MWNH Museum Wiesbaden, Germany (Dr M. Geistardt)
NHMV Naturhistorisches Museum Wien, Austria (Dr J. Gruber)
NHRS Naturhistoriska Riksmuseet, Stockholm, Sweden (Dr T. Kronestedt)
NMBA National Museum, Bloemfontein, South Africa (Mr L. Lotz)
NMBZ National Museum of Natural History, Bulawayo, Zimbabwe (Ms M.
FitzPatrick, the late Ms J. Minshull)
NMSA Natal Museum, Pietennaritzburg, South Africa

OXUM Hope Entomological Collections, University Museum, Oxford, UK (Dr I.
Lansbury)
PPRI Plant Protection Research Institute, Pretoria, South Africa (Dr A. S.
Dippenaar-Schoeman)
SAMC South African Museum, Cape Town, South Africa (Dr V. B. Whitehead,
Ms C. Car, Mrs M. Cochrane)
SMFD Forschungsinstitut und Natunnuseum Senckenberg, Frankfurt, Gennany
(Dr M. Grasshoff)
SMNS Staatliches Museum ftir Naturkunde, Stuttgart, Gennany (Dr W.
Schawaller)
SMWH State Museum, Windhoek, Namibia (Ms E. Griffin)
TMSA Transvaal Museum, Pretoria, South Africa (Mr W. Haacke, Mrs L. Brown)
USNM United States National Museum of Natural History, Washington D.C., USA
(Dr J. Coddington)
UZMD Universitets Zoologiske Museum, Copenhagen, Denmark (Dr B. Petersen)
ZMHB Museum flir Naturkunde der Humboldt Universitat, Berlin, Gennany (Dr
M. Moritz)
ZMPA Instytut Zoologii PAN, Warszawa, Poland (Dr A. Slojewska)
ZMUH Zoologisches Institut und Zoologisches Museum, Universitat von
Hamburg, Gennany (Dr G. Rack)
ZSMC Zoologische Staatssammlung, Munchen, Gennany (Dr & Mrs B. Baehr)
Geographical localities:
For simplicity, South African provinces referred to in the text reflect the older
(prior to April 1994) four-province system (Cape, Natal, Transvaal and Orange Free
State) and two traditionally accepted regions within two of the provinces (Zululand-
the Natal region north of the Tugela River - and Transkei - the old Cape region
between the eastern Cape and Natal). Use is also made of geographical regions
within the provinces (e.g. eastern Cape, western Cape, north-western Cape, northern
Transvaal) for descriptive purposes.
NATURAL HISTORY OF PALYSTES

Habitat and behaviour
Like the other larger tropical heteropodids of South America, Africa, Madagascar
and Indo-Australasia, Palystes species are nocturnal hunting spiders. During the day,
adults are inactive, sheltering under loose bark or stones, or in any suitable crevice.
In the field, Palystes species are usually associated with canopy and open forest on
moist, sea-ward aspects of inland mountains, wooded gorges and river valleys, or
with low altitude forest and scrub on the coastal plains abutting the Indian Ocean. P.
superci/iosus and P. johnstoni both also occur in savannah woodland. Night-active
wandering hunters, they retire into rock, plant or tree crevices, and under loose bark
during the day, relying on immobility and cryptic colouring to escape detection. They
are particularly easily collected in plantation pockets of pine trees in indigenous
forest where they are found under the loose bark of felled logs or dead branches.
8 ANNALS OF THE NATAL MUSEUM, VOL. 371996
Label data show that three species are commonly encountered in suburban gardens
and homes in the higher rainfall areas of southern Africa: P. castaneus in the south-
western Cape, particularly the Cape Peninsula including Cape Town; P. superciliosus
in Mozambique, Swaziland, Transkei, the Transvaal, Natal, and southern and eastern
Cape provinces of South Africa; P. johnstoni in Malawi and Zimbabwe.
Reproduction
Observations of P. superciliosus mating have been recorded by Fourie (1978), and
Croeser (1979). Cayton-Boxall (1988) observed mating by Palystes but did not
identify the species. Straddling of the female by the male during mating has been
observed by myself and by Fourie.
Cayton-BoxaU noted that the male approached the female from alternate sides
during mating but did not record straddling. Fourie noted that once the male had
mounted the female, she turned her abdomen slightly to orientate the epigynum
towards the direction of the approaching palp, while the male gave its palp a slight
twist to engage it to the epigynum. The bulb contents were then rotated 'with a fast
screw-turning action', followed immediately by distension, then gradual contraction
of the bulb. The bulb was distended and contracted between 25 to 30 times before the
palp was withdrawn and preened through the chelicerae. The male then changed
position to present the other palp, the female re-orientating her epigynum in response
to light leg taps on the abdomen. The process was repeated throughout the
observation period (between six and seven hours), Contractions initially lasted 10 to
20 seconds but towards the end were slower (up to 5 minutes),
Warren (1926) estimated that a female P. superciliosus laid approximately 800
eggs during an 18-24 month life span, with about 100 eggs in the fIrst egg sac, this
number increasing to a peak of about 200 and then diminishing to about 40 per sac.
Both P. castaneus and P. superciliosus usually enclose each egg sac within fist-sized
nest envelopes of tough, papery silk reinforced with leaves, twigs and other debris
(Warren 1926, D'Ewes 1967, Croeser 1979); new nests being most often found during
summer (Croeser). The nest may be suspended by reinforced 'cables' of silk in a
shrub, in the thorny crown of a caulescent Aloe species (Liliaceae) (pers. obs.), or
below an overhanging rock. The female remains in the vicinity of the nest for some
time and will resuspend the nest if it is detached or disturbed (D'Ewes 1967, Croeser
1979, Yates 1968). The young P. superciliosus hatch some 17 days after the eggs have
been laid (Warren 1926) and work their way through the walls of the egg sac about 4
days later. Warren observed that the young remain clustered on the nest for several
weeks until their silk glands and alimentary canal are fully formed, being nourished by
yolk stored in the abdomen. During this stage, when the spiderlings will also undergo
their first ecdysis, the mother remains with the nest and 'very definitely guards her
offspring'. Sibling camivory has been observed towards the end of this period (Warren
1926, Croeser 1979), when the spiderlings begin feeding and dispersing. Aerial
dispersal of the spiderlings by ballooning has not been observed. Warren also studied
ecdysis (1925a), egg development (1929) and cellular division (1925b) in Palystes.
Adulthood in a laboratory-reared female was reached in 10 months (Warren 1926).
Diet
Specimens of P. superciliosus kept by me in the laboratory were successfully fed
on cockroaches, crickets, grasshoppers, moths and Tenebrio moUtor larvae. Prey is
caught on foot without the aid of silk, and eaten directly, without silk wrapping.
Warren (1923) twice observed P. superciliosus eating common dwarf geckos,
Lygodactylus capensis, in Pietennaritzburg. One gecko measured 70 mm in length,
the other 45 mm. All that was left of the smaller specimen after about two and a half
hours of feeding by the spider was a discarded blackish, rounded ball about 6 mm in
diameter. It contained some skin scales and connective tissue, as well as a few
vertebrae and long limb and jaw bones, the latter still with their teeth. The bones
were clean and empty of marrow and pulp.
Natural enemies
Apart from the spectrum of animal life (birds, small mammals and frogs amongst
others) which include spiders in their diet and which are likely facultative predators
of Palystes, pompilid wasps (Hymenoptera: Pompilidae) and mantispids
(Neuroptera: Mantispidae) have been identified as specialised predators and
parasitoids of Palystes species.
Gess & Gess (1980) observed that the large pompilid Tachypompilus ignitus
exclusively preyed on species of Palystes. Dr F. W. Gess (pers. comm., 1988) also
recorded predation of Palystes species by several species of another pompilid,
Cyphononyx (basalis, decipiens, andflavicornis) in the eastern Cape and Karoo.
The mantispid Climaciella erichsoni has been collected from a Palystes nest in
Port Elizabeth and several unidentified mantispids have been reared from egg sacs of
P. superciliosus collected in the field in Natal (NMSA).
Medical importance
Only a few clearly established cases of envenomation in man by species of
Palystes in South Africa are known (D'Ewes 1964, Newlands 1975, Newlands &
Martindale 1981, pers. records). The symptoms are very mild (initial burning pain for
a few minutes at the site of the bite, followed by slight swelling or barely perceptible
oedema with no further symptoms). In an exceptional case, D'Ewes (1964) reported
a woman bitten three times on the arm in Cape Town. Symptoms included dizziness,
vomiting, swelling of the arm and of the armpit glands, and inflammation of the bite
site. For some years P. superciliosus was placed on the list of potentially harmful
South African arachnids (reviewed by Newlands 1975), following a misinterpreted
experiment by Steyn (1959) in which a guinea-pig died after being bitten on the nose.
Later Newlands & Martindale (1981) established in a series of controlled
experiments that the cause of death had been shock, not envenomation, and that
while the bite of the spider could be irritant, it was not dangerous. A label with an
adult female of P. johnstoni collected in Umtali, Zimbabwe, in June 1982 bears the
following note: ' .. Office - bit man - chest pains, difficulty in breathing. Treated with
antihistamines .. ' .
TAXONOMY
SUBFAMILY PALYSTINAE
Palysteae Simon, 1897a: 62. Type genus designated: Palystes L. Koch, 1875.
Pa1ysteinae: Petrunkevitch (1928: 54); Bonnet (1958: 3308).
Pa1ystinae: Roewer (1954: 725).
Palisteinae: Petrunkevitch (1928: 159) [lapsusl.
Remarks: Simon (1897a) was the first to recognise and describe a supra-generic
group based on Palystes (Palysteae in his clubionid subfamily Sparassinae). This
included three genera: Palystes, the type genus, from Africa and Australia, Tychicus
Simon, 1880, from Malaysia, and Micrommata Latreille, 1804, from Europe.
Petrunkevitch standardised the subfamily name, converting Palysteae to Palysteinae
(1928). The spelling of the subfamily name has not been consistent. Bonnet (1958)
followed Petrunkevitch, referring to the subfamily as Palysteinae, while Roewer
(1954) cites it in its correct form as Palystinae.
Simon distinguished the Palysteae from other heteropodid genera chiefly on the
proportions of the eyes: the anterior laterals being considerably larger than the
anterior medians; the posterior eyes all sessile, similar in size to each other and
equidistant. Jarvi (1912 1914) modified Simon's Palysteae by transferring
Micrommata to his new group Micrommateae. Two other genera were subsequently
added to the Palystinae by Roewer in his catalogue (1954): Exopalystes Hogg, 1914,
from Papua New Guinea, and Palystella Lawrence, 1928, from South Africa.
Examination of type material during the present study revealed that Palystinae s .1.
contained genera with affinities elsewhere, and that the type genus, Palystes s. /., was
also paraphyletic. Consistent differences in characters between the African and Indo-
Australasian species of Palystes s. I. are summarised in Table 1.
The character differences between the African and Indo-Australasian species
previously assigned to Palystes, are recognised at generic level in this study. The
generic name Palystes is now reserved for those species here considered congeneric
with the type species, P. castaneus, of the western Cape Province of South Africa.
Palystinae has accordingly been redefined and now includes only Palystes s. s. (as
redefined and revised in this study), Parapalystes gen. n. of the western Cape,
Anchonastus Simon (transferred from the Staianinae) from central Africa, and
Panaretella Lawrence, (transferred from the Heteropodinae) of south-eastern Africa.
Most of the Indo-Australasian species now excluded from Palystes s. s. have been
transferred from the Palystinae to the Heteropodinae: to Heteropoda and to an allied
resurrected Oceanian genus, Gnathopalystes Rainbow, 1899. Also transferred from
the Palystinae are the southern African genus Palystella Lawrence (which belongs in
a group with Orchestrella Lawrence) and the Indo-Australasian genera Exopalystes
and Tychicus. Subfamily placement of these genera, which do not appear to belong in
either Palystinae or Heteropodinae, must await further revisory work.
Palystinae and Heteropodinae appear to be closely related. Apart from similarities
in size, behaviour, habitat choice and general body proportions, the two subfamilies
also share characters that include the arrangement and proportions of the eyes
(differing only in the position and orientation of the AME, and in the PLE which are
raised and larger than the PME in some genera of Heteropodinae), the shape of the
tarsal slit sense organs (Fig. 15), and the basic arrangement of leg spines and the
shield-like marking of the venter. For this reason earlier workers placed a number of
heteropodine species in Palystinae. Heteropodinae (discussed later) differ in a
number of characters, however, and all members have microdenticles on the fang
furrow floor, which appears to be a synapomorphy for Heteropodinae.
TABLE 1
Differences between Gnathopalystes Rainbow, 1899, gen. resurr. and Palystes s. s. Koch, 1875.
Gnathopalystes Palystes s. s.
Female epigynum:
• Median septum lobe covered. : • Median septum lobe well exposed.
• Lateral sclerites contiguous anteriorly. :I • Lateral sclerites well separated anteriorly.
Carapace:
• Domed posterior to PER. : • Flat posterior to PER.
• Slopes downward anterior to fovea, I • Slopes downward posterior to fovea,
fovea steeply inclined. : fovea horizontal.

I
Eye group:
• AER procurved (AME with upper rims • AER straight (AME on a common
on a line with those of ALE). I median line to ALE).
I
• AME strongly tilted upwards. I • AME only slightly tilted upwards.
I
Leg spination:
• Patellae with one, (retrolateral) spine. : • Patellae with two (retro- and antero-lateral)
: spines.
• Tibiae dorsally with no spines. : • Tibiae dorsally with two spines.
I
Leg length:
• Sexual dimorphism of leg length - males : . No sexual dimorphism of leg length
with proportionately longer legs than females.: apparent.
Cheli~erae:
• Retromargin of fang furrow with five • Retromargin with three teeth.
teeth (the fifth very small basal tooth
sometimes missing).
• Anteromargin with two teeth. • Anteromargin with three teeth.
• Fang furrow with pad of microdenticles • Fang furrow smooth.
(median teeth).
Ster urn:
• Uniform brown. : • Black or with one or more dark
I
I
transverse bands.
Labium:
• Slightly wider (1.1 X) than long. • Usually much wider (1.2-1.5 X) than long.
• Bluntly conical apex. • Widely truncate apex.
• High waist. • Low waist.
Diagnosis: Palystinae s. s. appears to be monophyletic, defined by the synapomorphy

of the unique arrangement and proportions of the AER (Fig. 2). The AER is straight
with the AME (the smallest eyes) on a common median line with the ALE (the
largest eyes). The AME are directed forwards and only slightly tilted upwards. In
their apparent closest relatives, the Heteropodinae, the AER is procurved with the
upper lens margins of the AME on a common line with those of the ALE, with the
AME strongly tilted upwards. Palystinae are further defined by common possession
of the following characters:
AME
ef _--<-.--",-
4
Figs 2-6, Palystes castaneus (Latreille, 1819), female, 2. Head from front (AME = anterior median eye;
PME = posterior median eye; PLE = posterior lateral eye; ALE = anterior lateral eye; cJb =
cJypeal band; Ie = lateral condyle). 3. Sternum from below (rna = maxilla; la = labium; co =
coxa; ste = sternum). 4. Carapace. dorsal view (tf = thoracic fovea). 5. Abdomen from below
(ef = epigastric furrow). 6. Carapace from side (showing leg bases I-IV). (SAMC B9129, Cape
Town). Scale = 4 mm.
Leg spination pattern (Fig. 7), including one or two dorsal spines on tibiae I-IV of
both sexes; ventral banding of tibiae I-IV (absent in Panaretella); cheliceral
dentition (Figs 11, 94) 3:3 with three equal sized retromarginal teeth, three subequal
anteromarginal teeth (median largest, basal smallest) and with fang furrow floor
smooth, without microdenticles (Heteropodinae 4:3, with microdenticles on fang
furrow floor); ALE»PLE=PME>I=AME; PER straight or slightly recurved, eyes

equidistant, sessile; pigmented sternum either black, black with a posterior pale to
yellow patch or with 1-3 dark to black transverse bars, latter sometimes coalescing to
cover most of sternum (except Panaretella with an unmarked sternum); labium wider
than long, low-waisted; epigynum with ventrally exposed median septum (Figs 8, 71,
81, 110); epigynum internally with sperm ducts straight or hooked, but not coiled
more than 360 on their axes; embolus (Figs 9, 41, 80, 112) straight or reflexed less
than 275, not coiled.
Distribution: Central, eastern and southern Africa.
Key to genera of Palystinae s. s.

Sternum and coxae ventrally with fine black cuticular spots against pale white
to yellow-brown background; ALE only slightly larger than other eyes; PER
1.6 X wider than AER; carapace only slightly (1.2 X) longer than wide;
abdomen posteriorly with a black marking on either side (east coast and interior
of South Africa) ................................................................. Panaretella Lawrence
Sternum and coxae marked otherwise; ALE much (1.6 X) larger than other
eyes; PER 1.1-1.2 X wider than AER; carapace much (at least 1.4 X) longer
than wide; abdomen posteriorly without lateral black markings ......................... 2
2(1) Caput domed posterior to PER; sternum black except for white to yellow
posterior or anterior third (Fig 115); coxae each ventrally with 2-5 round, dark
marks (Fig 115) (eastern and western Cape Provinces of South Africa,
including Namaqualand) ..................................................... Parapalystes gen. n.
Caput sunken or flat; sternum entirely black or with one or more transverse
dark bands; coxae ventrally uniform in colour or partially invaded by the solid
dark lateral markings of the coxae ....................................................................... 3
3(2) Carapace sunken posterior to PER; carapace, abdomen and legs with
pilose tufts of setae; posterior spinnerets removed from, and much longer
than, anterior spinnerets (ZaIre and surrounding regions of central and west
Africa) ................................................................................... Anchonastus Simon
Carapace flat posterior to PER (Fig. 6); without tufts of pilose setae; posterior
and anterior spinnerets adjacent, and similar in length (eastern and southern
Africa) .......................................................................................... Palystes L. Koch
Palystes L. Koch, 1875

Helicopis L. Koch, 1874: 495 [preoccupied].
Palystes L. Koch, 1875: 701. Type species: Palystes Jrenatus L. Koch, 1875 [by subsequent designation
of Simon (l897a: 66).
Etymology: Koch (1875) did not give a derivation for Palystes. The name is probably
a mis-transliteration of the masculine Latin noun palaestes or Greek palaistes
(meaning wrestler) in view of the spiders' laterigrade arrangement of legs which
Koch noted were long and powerful.
Remarks: L. Koch (1874) proposed the name Helicopis for a new genus in his
introductory key to the Philodromidae. In 1875 he noted that the name was
Ti Pa Fe
pm
Ta Me Ti Fe
:~ §I :" ~:I
• • • •
• • • •
• •
•• •• ••
• • • •
• • • •
• • - II
•
~ •• ••
• •
• • III
•
• •
• • IV
Fig. 7. Schematic map of insertion of leg spines in Palystes. Left hand legs of a typical female P.
castaneus (Latreille, 1819) are mapped from the anterior to the posterior (I-IV). Leg surfaces
are divided in four planes (all = anterolateral; d = dorsal; pn = retrolaterai; v = ventral) for
consecutive segments (Fe = femur; Pa = patella; Ti = tibia; Me = metatarsus; Ta = tarsus).
Spination for left hand pedipalps of male (pm) and female (pO are mapped at the top.
[Mapping method modified from Griswold (l987a).]
preoccupied in Lepidoptera and replaced it with Palystes. He included four species in

the new genus: P. pinnotherus (Walckenaer, 1837) from Oceania, and three new
species: P. ignicomus from New Ireland (Oceania), and P. jrenatus and P.
superci/iosus from South Africa. Koch did not designate a type species for Palystes.
Simon (1880) recognised one of the originally included species, P. jrenatus, as a
junior synonym of Thomisus castaneus Latreille, 1819 and later (1897a) designated a
type species: 'TYPUS: P. castaneus Latr.(P. frenatus L. Koch).' Under Article 69a (v)
of the Code this constituted designation of P. Jrenatus as the type species of the genus
(and not Thomisus castaneus which was not one of the originally included taxa). The
type material of P. Jrenatus has not been found, but the two names are clearly
synonymous based on agreement of the original descriptions, gender and provenance.
In view of the importance of the identity of the type species in this revision, the adult
female type specimen of Thomisus castaneus Latreille, 1822, is now designated as
neotype for P. Jrenatus, making the two names objective synonyms. This clearly
establishes the identity of the type species and firmly links it to that of the senior
synonym, Palystes castaneus,. This serves to stabilise the name since, following
Simon's (1897a) designation of Palystes castaneus as the type species, P. castaneus
has always been cited by subsequent authors as the type species of the genus.
Examination of all the available type material of the 38 species previously
assigned to Palystes revealed, apart from a number of misplaced species belonging in
other genera, four apparently natural groups, two African and two Indo-Australasian.
A number of morphological characters consistently separate the four groups, and the
differences are here recognised at generic level. The genus Palystes s. l. is
accordingly split into: 1) Palystes s. s., reserved for those African species considered
congeneric with the type species Palystes castaneus (Latreille); 2) Parapalystes, a
new genus described below, from the interior of the Cape Province of South Africa,
for those species considered congeneric with its type species Parapalystes
euphorbiae sp. n.; 3) Gnathopalystes Rainbow, 1899, previously synonymised by
Simon (1903) under Palystes and here resurrected to accommodate those extra-
African species considered congeneric with the type species, Gnathopalystes Jerox
Rainbow; 4) a probable new genus closely allied to Heteropoda Latreille, 1804 (at
present the species belonging here have been placed in Heteropoda pending revision
of this genus). The differences between Palystes s. s. and two of the genera,
Parapalystes and Gnathopalystes, are summarised in Tables 1 and 2.
Of the four species originally included in Palystes by L. Koch, the two southern
African species, P. Jrenatus and P. superciliosus, are retained in Palystes s. s. Of the
two included Australasian species, P. ignicomus is transferred to Gnathopalystes
while P. pinnotherus is considered incertae sedis.
Diagnosis: Two distinct longitudinal stripes of white or yellow setae present at outer
margin of each cheliceral base (Figs 2, 4, 6) ,and a broad clypeal 'moustache' of
closely adpressed white to yellow setae 'CLB' (Fig. 2) with projecting elongate
dentoid scales on their underside (Fig. 10). The 'moustache' and cheliceral markings
are synapomorphies unique to Palystes and Parapalystes. Palystes differs from
Parapalystes in the shape of the carapace (flat, not domed posteriorly), orientation of
thoracic fovea (parallel to body plane, not sloping downwards on posterior declivity),
coxal markings (no blotched-spot markings ventrally as in Fig. 115), sternum colour
(no yellow-brown patch across posterior of black sternum), the single tibial
apophysis on the male pedipalp (as opposed to two spur-like apophyses with
additional ridges as in Fig. 113), and in the dorsal spination of the tibiae of legs I-IV
(two basal and apical spines instead of a single apical spine).
Description:
Medium to large, light to dark brown hunting spiders, up to 30 mm in body length,
and with an extended leg span reaching 110 mm,
Carapace: (Figs 4, 6) Ovate, flat, sloping downwards posteriorly, posterior to
thoracic fovea; densely covered with chestnut-brown setae (ca 0.4 mm long) and
with numerous, scattered, longer, paler, attenuate setae up to 1.4 mm long in vicinity
of eyes, those above and between upper margins of ALE projecting forwards and
converging to form a loose peak of hairs extending beyond anterior margin of
carapace; head region parallel-sided, clearly delineated by striae posteriorly; fovea
(Fig, 4), from which several faint, depressed, striae radiate, a straight, longitudinal,
narrow furrow occupying approximately 0.27 X of CL, flaring into small shallow
depressions at either end, anterior to posterior declivity. Eyes (Figs 2, 4, 6): PER
straight, occupying approximately 0.6 X HW and approximately 1.2 X wider than
AER; AER recurved from above, straight from front; ALE»PLE=PME>I=AME,
with ratio range AME:ALE:PME:PLE = 1:1.3-1.7:0.9-1.2:0.9-1.2; MOQ as long as
wide to slightly longer than wide, narrowed anteriorly; AME slightly tilted upwards,
on either side of slightly raised tubercle between ALE and on common median line
with ALE, other eyes sessile; immediately anterior to each of PE a small patch of
darkened cuticle with numerous, short adpressed setae and similar patches
immediately posterior to each of anterior eyes. Clypeus (Fig. 2): Vertical, narrow,
1.1 X diameter AME in height, densely covered with short, blunt-tipped, white to
off-white adpressed setae (Fig. 10) with projecting teeth on their lateral margins
forming white band more than 0.6 X width of clypeus, tapering at either end, with 2,
irregular, rows of long, attenuate, bristle-like, pale setae among adpressed white
setae; ventral row of approximately 14 convergently directed setae, increasing in
length from near lateral margins of clypeal band to centre, 0.3-1.0 mm long, and
short, dorsal row of four longer setae 1.5-1.9 mm long. Chelicerae (Figs 2, 4, 6):
Robust, dark to black, 1.7-1.9 X longer than wide, profusely covered with long,
attenuate, dark and chestnut coloured bristle-like setae and with 2 longitudinal white
setose stripes at their external margins (Fig. 2); fang furrow (Fig. 11) smooth, without
denticles; retromargin of fang furrow with 3 equidistant large teeth, similar in size or
basal largest; anteromargin with 3 teeth, median largest, basal smallest (Fig. 11),
occasionally additional small teeth present in individual specimens (as in male
paratype of P. karooensis, Fig. 48); retromargin of fang furrow with fringe of long,
red scopular hairs external to teeth, anteromargin with shorter, denser scopula
external to teeth. Labium: 1.2-1.5 X wider than long, broadly truncate apically;
middle widest, sides convergent but apex broadly truncate; posteriorly with scattered
attenuate setae, apical margin pale with approximately 20 long, attentuate, reddish
setae up to 1.2 mm long. Sternum: 1.1-1.4 X longer than wide; shield-shaped, nearly
flat, scallops shallow, sigillae not apparent, widest between coxae II, posteriorly
tapering, blunt-tipped; densely setose and either solid black or with 1-3 dark
transverse bars of dark setae and darkly pigmented cuticle against background of
lighter cuticle and light to medium brown setae. Maxillae (Fig. 3): Approximately
1.5 X longer than wide, inner margins straight proximally, diverging distally, exterior
margins bulging outwards distally; 3 groups of red to orange scopulae with blunt-
CROESER: REVISION OF PALYSTES (ARANEAE: HETEROPODlDAE) 17
tipped bristles present (well-developed, flared, apical scapula with long setae; inner,
sub-apical scapula of shorter, stouter setae; scapula of even shorter, finer setae
forming dense mat on inner margin of maxillae); serrula present, linear, at outer
angle of apex.
sp
eco
Ii
.·.;...;.:..;..;.-els
8a
st
em
9a
Figs 8-9. Palystes castaneus (Latreille, 1819). 8a. External female epigynum (ems = median epigynal
septum; els = lateral sclerites of epigynum; Ii = lateral invagination separating septum from
lateral sclerites; msl = posteriorly produced lobes of median septum). 8b. Internal female
epigynum (goc =gonopore cavity of septum; eco =epigynal conductor of afferent sperm duct;
sp = spermatheca lying dorsal to bulbous invagination of efferent sperm duct) (SAMC 89129,
Cape Town). 9a. Anterolateral aspect, left male pedipalp (em =embolus; Is =lateral sclerite of
tegulum forming a support flange to embolus). 9b. Ventral aspect, left male pedipalp (co =
membranous conductor; st = sub-tegulum; te = tegulum; ta = tibial apophysis) (NMSA, Hout
Bay,3.v.1984).
Legs: Fonnula (12)43 with Leg I = II, or one fractionally shorter than other; well-
armed with strong, erectile black spines (mapped in Fig. 7); significant spines are
femora I-III(3:2:3), femora IV(3:2: 1 or 3:2:2), patellae I-IV(l :0: I), tibiae I-IV
dorsally with proximal and distal spine; coxae usually dark anterolaterally, unmarked
ventrally or basal half dark or completely dark, never spotted; femora ventrally with
dark brown to black markings or unmarked; tibiae ventrally black interrupted by
median and apical broad pale to white transverse band; metatarsi and tarsi ventrally
with well-developed scopulae comprising greyish, blunt-tipped setae covered for
most of their length with hooked setules (Fig. 13); otherwise all surfaces densely
covered with setulate attenuate light and dark brown setae; pair of pectinate tarsal
claws terminally, each pair sub tended by well-developed tarsal claw scopula
comprising finely setulate, flattened, spatulate, apically-bisected setae (Fig. 14);
trichobothria (Fig. 17) numerous, no discernible pattern except for those arranged in
'V' pattern distally on dorsal surface of tarsi, arms of 'V' directed distally; tarsal
organ dorsal, bulbous, recessed (Fig. IS); prominent lip-shaped slit sense organs (Fig.
15) laterally on dorso-retrolateral and dorso-anterolateral surfaces of all tarsi.
Abdomen: Dorsally with clearly outlined brown to black cardiac marking present
or absent; ventrally with dark to black transverse crescent marking immediately
posterior to epigastric groove and generally with additional black, brown or white
markings between crescent and spinnerets; posterior and anterior spinnerets two
segmented, median spinnerets with a single segment; anterior spinnerets (Fig. 20)
stout, larger, wider and longer than posterior and median spinnerets; colulus absent;
anal tubercle prominent.
Epigynum: Externally (Fig. Sa), longer than wide, lateral sclerites widely
separated, large well-developed median septum present. Internally (Fig. Sb), spenn
ducts heavily sclerotised; spennathecae prominent, produced, straight or reflexed
(Fig. 47), supported by invagination which may fonn supporting collar (Fig. 78),
which may be interrupted medially (Fig. 53).
Male pedipalp: Embolus (Fig. 9a) sturdy, as long as tegulum, straight (Fig. 9a) or
reflexed (Figs SO, 93), flattened on a vertical plane to the palp (Figs 9a, 36, 54) or
along the same axis (Figs 92, 99), recurved less than 270 distally over tegulum;
conductor (Fig. 9b) well developed, fragile, diaphanous or white, not sclerotised,
distally enfolding apical sclerite of embolus. Tibial apophysis single, trilobate (Fig.
9b), bilobate (Fig. 56), flattened (Fig. 41) or an entire spur (Fig. 91); no additional
apophyses.
Distribution: Central, eastern and southern Africa.
Key to species of Palystes s .s.

Female ............................................................................................................... 2
Male ................................................................................................................. 15
2( 1) Epigynal septum (Figs 77, S I) deeply excavate anteriorly and with rectangular
ridge posteriorly which is sometimes produced; femora ventrally dark to black
with distal broad yellow band ............................................................................ 3
Epigynal septum not as above; femora ventrally mottled or with clear white
spots ................................................................................................................... 7
CROESER: REvrSrON OF PALYSTES (ARANEAE: HETEROPODIDAE) 19
Figs 10-15. Scanning electron micrographs of cuticular detail and setae of a female Palysres casraneus
(Latreille, 1819), from Newlands, Cape Town. 10. Clypeal setae (750 X). II. Left chelicera!
teeth and fang (25 X). 12. Dorsal metatarsal setae above and ventral scopular setae below (155
X). 13. Enlargement, showing hook-tipped serules of scopular setae (475 X). 14. Tarsal
scopular setae (1365 X). 15. Slit-sense organ on dorso-retrolateral surface of tarsus (1050 X).
3(2) Sternum with dark transverse bar between coxae I and II (sometimes with 2
fainter bars posterior to it); abdomen ventrally with faint grey to black bell-
shaped mark between black crescent and spinnerets; septum ridge (Fig. 77)
not produced posteriorly; spermathecae and penultimate sperm ducts (Fig. 78)
dorsal to supporting collar, lobes of collar encircle pre-penultimate ducts
(Africa south of Cunene and Limpopo rivers) ................. superciliosus L. Koch
Sternum with 2 or more dark transverse bars, or entirely black; abdomen
ventrally with large to small dark brown spots between dark brown to black
crescent and spinnerets; septum ridge (Fig. 81) produced posteriorly;
spermathecae and penultimate ducts ventral or internal to supporting collar
lobes, lobes not encircling ducts ....................................................................... .4
4(3) Sternum with 2 chevron-like dark transverse bars; spermathecae (Fig. 98)
slender, long, unattached to supporting collar and lying ventral to collar lobes
(not visible dorsally); penultimate sperm ducts external to collar lobes
(Malawi and Ziml'1abwe) .........................................................johnstoni Pocock
Sternum with 3 dark transverse bars (Fig. 85), posterior bar strongly recurved,
or sternum entirely black; spermathecae short, embedded in supporting collar
lobes; penultimate sperm ducts internal or anterior to collar lobes ................... 5
5(4) Abdomen ventrally (Fig. 88) with brown spots coalescing medially to form a
solid, laterally spotted, irregular mark between crescent and spinnerets;
spermathecae (Fig. 82) short, stout and embedded internally in lateral lobes of
supporting collar; afferent sperm ducts immediately posterior to spermathecae
and internal to lobes of supporting collar, visible dorsally (eastern ZaIre,
Rwanda, Uganda, Tanzania and northern MalaWi) .......................ellioti Pocock
Abdomen ventrally spotted but spots not coalescing to form mesal marking as
above; epigynum internally (as in Fig. 105) similar to above but afferent
sperm ducts anterior to spermathecae and anterior or external to lobes of
supporting collar, usually not visible dorsally ................................................... 6
6(5) Sternum entirely black; afferent sperm duct external to lobe of supporting
collar (Fig. 105); lobe strongly produced anteriorly to spermatheca
and penultimate sperm ducts (Warburton and Ermelo districts, south-
easternTransvaal, to Ndumu, northern Natal, South Africa) .............................. .
ansiedippenaarae sp. n.
Sternum with dark brown to black bars (Fig. 95); afferent sperm duct (Fig. 90)
embedded anteriorly on lobe of supporting collar; lobe not produced as above
(Kenya and Tanzania) ................................................................ hoehneli Simon
7(2) Sternum entirely black; femora I-II mottled without distinct markings;
abdomen laterally mottled, without distinct markings, ventrally with black
bell-shaped mark (Fig. 5) or black-framed yellow panel between black
crescent and spinnerets; septum (Figs 8a, 23 & 32) posteriorly produced
laterally (western Cape, South Africa) .............................................................. 8
Sternum (Fig. 45) usually with two mesally interrupted transverse dark bars
(sometimes faint, sometimes with an additional short longitudinal bar
mesally); femora I-II ventrally with clear white spots; abdomen laterally with
clear white spots, ventrally (Fig. 45) rich red to orange-red with distinct clear
white spots between black crescent and spinnerets (indistinct in specimens

from Amatola Mountains, eastern Cape, South Africa); septum (Figs 39 and
68) with transverse or posteriorly produced median lobe .............................. 10
8(7) Abdomen ventrally with broad, transverse, pale brown mark between black
crescent posterior to epigastric groove and dark patch anterior to spinnerets;
septum (Fig. 32) convex, posteriorly flattened but not divided mesally, with
pair of lateral swellings; spermathecae (Fig. 33) elongated, inserted
immediately posterior to supporting bulb and directed internally (Hottentots
Holland Mountains, western Cape, South Africa) .......................... stilleri sp. n.
Abdomen ventrally with solid dark brown to black shield marking between
epigastric groove and spinnerets; epigynum not as above ............................... 9
9(8) Epigynum externally with septum (Fig. 23) short, wide, laterally excavated, with
pair of lateral swellings immediately anterior to it; internally, efferent sperm
ducts (Fig. 24) inserted ventrally below bulbous lobes of flexible, anteriorly
interrupted, collar; spermathecae inserted externally to collar (Cederberg,
Piketberg and Clanwilliam districts, Cape, South Africa) ..... martinfilmeri sp. n.
Epigynum externally with septum (Fig. 8a) longer than wide, anteriorly
concave without lateral swellings, posteriorly raised and divided mesially;
internally efferent sperm ducts (Fig. 8b) without flexible collar; spermathecae
short, stout, inserted dorsally on small supporting bulb (Cape Town,
Stellenbosch, Somerset West and Bredasdorp districts, western Cape, South
Africa) ................................................................................ castaneus (Latreille)
10(7) Median lobe of septum transverse and not produced (Fig. 39) (Maseru district,
Lesotho) ................................................................................. crawshayi Pocock
Median lobe of septum produced posteriorly (similar to Fig. 52) ................... 11
11(10) Spermathecae inserted lateral to supporting lobes and produced anteriorly
(Fig. 72) (Nieuwoudtville, western Cape, South Africa) ................ stuarti sp. n.
Spermathecae not as above .............................................................................. 12
12( 11) Median lobe of septum more than 2 X longer than wide (Fig. 37); penultimate
loops of efferent sperm duct strongly produced towards each other (Fig. 38)
(possibly interior of eastern Cape, South Africa) ...................... .Iunatus Pocock
Median lobe of septum much less than 2 X longer than wide; penultimate
loops of efferent sperm duct not produced as above ...................................... 13
13(12) Median lobe of septum (Fig. 63) longer than wide; spermathecae stout,
inserted internally on supporting lobes and on plane horizontal to median line
(Fig. 64), not hooked apically (Port Edward district (southern Natal),
Transkei, Queenstown district and Amatola Mountains, eastern Cape, South
Africa) .................................................................................. perornatus Pocock
Median lobe of septum wider than long to very much wider than long;
spermathecae not as above .............................................................................. 14
14(13) Median lobe of septum (Fig. 57) not much wider than long, with obliquely
sloping neck and shoulders anteriorly; spermathecae (Fig. 58) stout, straight,
inserted externally on supporting lobes, produced anteriorly at angle of 45° or
less to longitudinal median line (Grahamstown and Alicedale districts, eastern
22 ANN ALS OF THE NATAL MUSEUM. VOL. 37 1996
Cape, South Africa) ............................................................... .leppanae Pocock

Median lobe of septum (Fig. 46) much wider than long, with well-defined
neck and horizontal shoulders anteriorly; spermathecae (Fig. 47) hooked
apically, inserted internally on supporting lobes and produced laterally at an
angle of 90° to longitudinal median line (Cradock, Graaff-Reinet and Beaufort
West districts, Cape Province and Kokstad and Cedarville districts, south-
western Natal, South Africa) .................................................... karooensis sp. n.
15(1) Sternum entirely black (western Cape, South Africa) .................................... 16
Sternum with one or more dark transverse bars on pale background ............. 18
16(15) Tibial apophysis (Fig. 9b) three-lobed; embolus straight; conductor elongate,
straight (Cape Town, Stellenbosch, Somerset West and Bredasdorp districts,
western Cape, South Africa) .............................................. castaneus (Latreille)
Tibial apophysis entire; embolus recurved through 90° over bulb; conductor
short, bowl-shaped ........................................................................................... 17
17(16) Abdomen ventrally with a solid dark brown to black shield marking
between epigastric groove and spinnerets; embolus (Fig. 29) columnar
proximally, flaring distally and hooked apically; conductor sessile (Cederberg,
Piketberg and Clan william districts, western Cape, South Africa) .................... .
martintilmeri sp. n.
Abdomen ventrally with broad transverse pale brown marking between black
crescent marking posterior to epigastric groove and black patch anterior to
spinnerets; embolus (Fig. 28) with a broad shallow channel proximally, obtuse
apically; conductor distinctly stalked (Hottentots Holland Mountains, western
Cape, South Africa) ........... ;.............................................................stilleri sp. n.
18(15) Abdomen ventrally with dark brown spots or with a faint grey to black bell-
shaped marking between crescent and spinnerets; femora I-IT ventrally dark
brown to black with broad yellow band apically; embolus relatively narrow,
inserted proximally, reflexed distally, with vertical lamellar processes apically
(as in Fig. 92) or entire (as in Fig. 80) ............................................................. 19
Abdomen ventrally with clear white spots against a light yellow to rich
orange-red background between crescent and spinnerets; femora I-IT ventrally
with white spots; embolus a horizontally flattened, broad, inclined blade
inserted medially (as in Fig. 54) ..................................................................... .23
19(18) Embolus distally entire (Figs 79, 101), without vertical lamellate
processes) ......................................................................................................... 20
Embolus distally with verticallamellate processes (Figs 93, 100) .................. 21
20(19) Sternum with transverse dark bar at coxae IT (sometimes with additional,
fainter, bars posterior to transverse bar); abdomen ventrally unmarked or with
a faint grey to black bell-shaped mark between crescent and spinnerets;
embolus (Fig. 80) apically aciculate and recurved through 270° (Africa south
of Cunene and Limpopo rivers) ....................................... superciliosus L. Koch
Sternum with 2-3 transverse dark bars (Figs 85-87), posterior bar strongly
recurved; abdomen ventrally with large brown spots coalescing medially to
form a large, irregular mark; embolus (Fig. 84) apically obtuse and recurved
through less than 180 0 (eastern ZaIre, Rwanda, Uganda, Tanzania and
northern MalaWi) ........................................................................... ellioti Pocock
21(19) Sternum with transverse black bar at coxae II (and a widely interrupted second
bar at coxae ill) (Fig. 103), abdomen ventrally with wide black bar and followed
by fine, intense black spots, embolus apically without distal carination but a
broad, flat, wide lamellate flange (Fig 101) internally to apical embolus (Orange
Free State and western Transvaal, South Africa) ......................... .leroyorum sp. n.
Sternum with 2 or more transverse dark bars, abdomen ventrally with large or
small dark brown (not black) spots or blotches, embolus with a distal lamellar
process apically ................................................................................................ 22
22(21) Sternum with 2 chevron-like transverse dark bars; embolus (Fig. 100) with
prominent lateral ridge crossing over to ventral face of blade distally
and blunt-hooked apically, proximal process distal to upper ridge, neck
angle wider than width of proximal process (MalaWi and Zimbabwe) ............. .
johnstoni Pocock
Sternum (Fig. 95) with 2-3 transverse dark bars; embolus with distal ridge not
crossing over to ventral face of blade, proximal process opposite to distal
process, neck angle narrower than width of proximal process ........................ 23
23(22) Distal process of embolus (Fig. 93) pronounced, short, flat; embolus column
distinctly bulbous medially (Kenya and Tanzania) ................... hoehneli Simon
Distal process of embolus (Fig. 108) reduced, elongate and reflexed apically;
embolus column slender medially, nearly straight (Warburton and Errnelo
districts, south-eastern Transvaal, to Ndumu, northern Natal, South Africa) .....
ansiedippenaarae sp. n.
24(18) Bulb with distinct hooked spine adjacent to base of embolus (Fig. 50) .......... 25
Bulb without spine ........................................................................................... 27
25(24) Spine at base of embolus (Fig. 73) extremely long, more than half length of
embolus (Nieuwoudtville, western Cape, South Africa) ................ stuarti sp. n.
Spine at base of embolus short, less than quarter length of embolus .............. 26
26(25) Tibial apophysis (Fig. 66) entire, slightly convex dorsally and with a ventral
lobe; embolus (Fig. 65) flared distally, lateral margin slightly sinuate but
flattened proximally; conductor recurved distally and laterally over embolus
(Port Edward district, south-eastern Natal, Transkei, Queenstown district and
Amatola Mountains, eastern Cape, South Africa) ............... perornatus Pocock
Tibial apophysis (Figs 51, 55, 56) bifurcate to trilobate; lateral margin of
embolus (Fig. 49, 54) straight to strongly sinuate, shouldered proximally
(Cradock, Graaff-Reinet and Beaufort West districts, Cape Province and Kokstad
and Cedarville districts, south-western Natal, South Africa) ...... karooensis sp. n.
27(24) Tibial apophysis (Fig. 44) short, flat, distinctly trilobate; bulb with an
elongate, raised ridge (Fig. 42) adjacent to base of embolus; lateral margin of
embolus (Fig. 41) flattened proximally and without a median channel (Maseru
district, Lesotho) .................................................................... crawshayi Pocock
Tibial apophysis (Fig. 60) elongate, with a proximal lobe; bulb without a ridge
as above; lateral margin of embolus ridged proximally and with deep, wide,
median channel (Figs 61-62) (Grahamstown and Alicedale districts, eastern

Cape, South Africa) ........ ,................ " ...................................... Ieppan ae Pocock
Figs 16-21. Scanning electron micrographs of Paiysles castaneus (Latreille, 1819). 16-20. Surface
structures and setae of a female from Newlands, Cape Town. 16-18. Dorsal surface of tarsus
II. 16. Organ near tarsal claw base (670 X). 17. Fluted trichobothrial base (1125 Xl. 18. Dorsal
organ (10535 X). 19. Dentate and finely setulate setae on abdomen near epigynum (700 Xl.
20. Spinning field of right anterior spinneret, showing two ampulJate and numerous piriform
spigots (145 X). 21. Papillate sensory setae at apex of pedipalp tarsus of a male from
Plumstead. Cape Town (595 Xl.
Palystes castaneus species group

Characterised by: Solid black sternum, epigyna with long, broad, flat, lightly to
deeply excavated median septum ('ems', Figs 8a, 23, 32). Embolus supported by
well-developed lateral tegular sclerite Cis', Fig. 9a).
Included species: P. castaneus (Latreille, 1819), P. martinfilmeri sp. n., P. stilleri sp. n.
Distribution: South-western Cape Province, South Africa (Fig. 22).
Palystes castaneus (Latreille, 1819)

Figs 1-22
Thomisus castaneus Latreille, 1819: 30. Holotype: sex unspecified [adult'" ], 'Cap de Bonne-Esperance'
[= Cape of Good Hope], de Lalande (MNHN) [examined].
Olios castaneus; Walckenaer 1837: 571.
O.[lios] castanea; Simon, 1864: 410.
Palystes castaneus; Simon, 1880: 262. [Designated as type species of Palystes L. Koch, 1875, through
synonymy with Palystesfrenatus L. Koch, 1875, by Simon 1897a: 63, 66].
Palistes [sic] castaneus; Petrunkevitch, 1928: 159.
Olios fuscus Walckenaer, 1837: 573. Holotype: sex [adult '" I and provenance not stated ('elle est
certainement estrangere a I'Europe') (MNHN) [examine<1l. [Synonymised by Simon 1880:
262-263].
Ocypete melanogaster C. Koch, 1845: 37, pI. CCCCIV, fig. 975. Holotype: adult 0', 'Das Vorgebirg der
guten Hoffnung' (location of type unknown). [Synonymised by Simon 1897a: 63].
Ocypete nobilis sensu C. Koch, 1845: 37, pI. CCCCV, fig. 979 [misidentification, nec Aranea nobilis
Fabricius, 1798]. [Synonymised by Simon 1880: 262].
Palystes frenatus L. Koch, 1875: 705, pI. LX, figs 4a-b. Syntypes: 3"', 'Sudafrika' (lost, not in
Tubingen). [Synonymised by Simon 1880: 262-263] Neotype: holotype '" of Thomisus
castaneus Latreille, 'Cap de Bonne Esperance' (MNHN), here designated.
Palystes chaperi Simon, 1880: 263-265. Holotype: adult 0', 'Cap', M.Chaper (MNHN 3.242)
[examined]. Syn. n.
Remarks: Latreille's (1819) description of Thomisus castaneus was based on a
specimen 35 mm long, sex unspecified, from the Cape of Good Hope, given to him
by Gattoire. Simon (1880: 263) stated that he had seen Latreille's type (,Le type de
Latreille ... '), an adult female, with the label 'Philodromus castaneus Latreille, M.
Lalande, cap de Bonne Esperance' at MNHN. He noted that the specimen was badly
faded and the epigynum was missing. Two adult females from the Cape, both
identified as Palystes castaneus and both labelled 'type', are in MNHN, but only one
has the label data cited by Simon (recognised here as the type specimen for P.
castaneus). The unusual shape of the epigynum in this species, with the plate
forming a deep, sunken basin, may well have led Simon to believe it was absent in
the specimen he examined. The length given for the specimen in Latreille's
description was 35 mm, a full centimetre longer than that of the specimen accepted
here as the type. The discrepancy may due to typographical error.
Designation of the holotype specimen of Thomisus castaneus as the neotype for
Palystes frenatus is necessary to firmly fix the identity of the latter as it is the name
of the type species of the genus. No type material for P. frenatus is known to exist. L.
Koch did not designate a type specimen for P. frenatus, but in the original description
(1875: 705-706) referred to three specimens [at least one of which was an adult
female as PI. LX Fig. 4b depicts an epigynum] in the 'zoologischen Kabinet der
UniversiHit Tiibingen'. None of the syntypes for P. frenatus can be found in Tiibingen
(Prof. H. M. Peters, pers. comm.), and they have not been located elsewhere. Koch's
illustration of an epigynum (PI. LX, Fig. 4b), although stylised, is clearly that of P.
castaneus. The neotype specimen here selected, like that of the specimen illustrated
by Koch, is an adult female. The locality given by Koch ('Stidafrika') is vague, but
includes the type locality for Thomisus castaneus ('Cap de Bonne-Esperance').
C. Koch (1845) was the ftrst to describe the male (as Ocypete melanogaster).
Pocock (1896) synonymised O. melanogaster with P. castaneus, saying: 'The male
[of castaneusl, in fact, agrees so closely with the description of Ocypete
melanogaster of C. Koch ... that I have not hesitated to regard the latter as the male
of castaneus.' Koch's description includes a reference to the forked spur of the
trilobate tibial apophysis of the pedipalp, a character unique to P. castaneus. The type
of Ocypete melanogaster has not been found, but Pocock's synonymy is undoubtedly
correct. Simon (1897a) listed this synonymy, but (p. 63, Fig. 58) incorrectly
identifted a drawing of a male palp of P. superciliosus as that of a male P. castaneus.
The mistake was pointed out by Strand (1907b: 676).
Olios fuscus Walckenaer, 1837, was based on a badly preserved specimen, sex
unspecifted, of unknown provenance. Walckenaer separated it from Thomisus
castaneus Latreille, chiefly on the basis of what he saw as strongly recurved eye
rows in O. fuscus. The type specimen in MNHN is damaged and missing most of its
legs (except for right leg IV and left femur I), but the epigynum is intact and is
clearly that of P. castaneus. The damaged carapace gives the impression that the PER
is recurved.
Ocypete nobilis was the name used by C. Koch (1845) for a female from the Cape.
The specimen has not been found in any of the collections known to have C. Koch
types. In his description he referred to Aranea nobilis Fabricius, 1798 [described
from 'India']. The Fabricius name was listed by Zimsen (1964), but the specimen
described by Fabricius was not found by Zimsen and is not in Copenhagen which
contains much of the extant Fabrician collections. Simon (1880) regarded Ocypete
nobilis sensu Koch as a possible misidentiftcation of P. castaneus; Pocock (1896)
concurred, adding that it was probably not nobilis sensu Fabricius because of its type
locality. Koch's description (p. 37), closely matches that given by Latreille for P.
castaneus and the illustration (PI. CCCCV, Fig. 979) is clearly that of a Palystes
species.
Palystes chaperi was the name given by Simon (1880) to a male collected by
Chaper at the Cape. There are two specimens determined by Simon as P. chaperi in
MNHN. One, the identifted male type cited by Simon, is an adult P. castaneus. P.
chaperi is accordingly here recognised as a junior synonym. The other, a female,
belongs in the new genus Parapalystes. Although Simon provides a detailed
description of the male pedipalp (unmistakably that of P. castaneus), his description
of the rest of the body is clearly based on the female. This is evident from his
reference to markings (white spot anterior to fovea, spot markings of the coxae) and
leg spination (a single dorsal tibial spine) present in the female, which are typical of
Parapalystes but absent in the male type specimen. He also notes that the two white
line markings at the outer edge of the chelicerae (characteristic of Palystes) are
absent. The cheliceral markings are present in the male type, but absent in the
female.
1:2000000
•
• -Piketberg
•
_Stellenbosch
• Hottentots Swellendam
•
D Nature
Holiand
Reserve
•
Bredasdorp
Fig. 22. Distribution of Palystes castaneus species group in southern and western Cape, South Africa.
• = P. castaneus (Latreille, 1819); • = P. martinfilmeri sp. n.; = 0 P. stilleri sp. n.
Diagnosis: Both the trilobate tibial apophysis (Fig. 9b) of the adult male pedipalp and
the large, basin-shaped epigynum (Fig. 8a) of the adult female readily separate this
species from its congeners.
Distribution: South-western Cape Province, South Africa (Fig. 22): Cape Peninsula,
Cape Town, Stellenbosch. Anomalous specimens from Brandfontein and Springfield
in the Orange Free State, Delagoa Bay (Mozambique), and Harare and Bulawayo in
Zimbabwe, were probably accidentally transported by man.
Redescription of holotype female:
The specimen is encrusted with white sediment, and a number of the legs are
detached but it is otherwise in reasonable condition. There is a pinhole through the
carapace, just left of the fovea, and the cuticle at the end of the abdomen is
damaged.
Total length: 22.26 mm. The figured female (Figs 8a, 8b) is from Cape Town
(SAMC B9129).
Colour and markings: Abdomen ventrally (Fig. 5) with black 'X' -shaped marking
straddling epigynum anterior to epigastric groove, and with large, black bell-shaped
marking opening from transverse black crescent posterior to epigynal groove and
tapering posteriorly at spinnerets; sternum (Fig. 3) entirely black, densely covered
with short, attenuate black setae against dark red-brown to black-pigmented cuticle;
femora without ventral markings; coxae dark anterolaterally and on much of their
ventral surface.
Carapace: CL 10.38, CW 8.13, HW 5.13; thoracic fovea approximately 0.28 wide,
2.50 long, occupying 0.24 X of CL and 0.31 X of CWO Eyes: Eye diameters: AME
0.42, ALE 0.65, PME 0.52, PLE 0.52; PER 3.25 wide, wider than AER by 1.27 X
and occupying 0.63 X of HW and 0.40 X of CW; eye ratio AME:ALE:PME:PLE
1: 1.56: 1.26: 1.26; distances separating eyes: AME-ALE 0.12, AME-AME 0.31,
AME-PME 0.85, ALE-PLE 0.69, PME-PLE 0.54, PME-PME 0.54; MOQA 1.08,
MOQP 1.54, MOQL 1.54; MOQ 1.00 X longer than wide. narrowed anteriorly. with
MOQP 1.43 X wider than MOQA. Clypeus 0.55 high, 1.33 X diameter AME.
Chelicerae: 3.75 long. 2.19 wide. 1.71 X longer than wide. Sternum: 4.69 long, SWI
3.44. 1.36 X longer than wide. SW2 2.50. Labium: 1.69 wide. 1.08 long. 1.57 X
wider than long. occupying 0.68 X of SW2. Maxillae: 2.52 long, 1.63 wide, 1.55 X
longer than wide.
Abdomen: 11.88 long, 8.13 wide.
Legs: Spination as in Fig. 7. Measurements:
PaJpus I II III IV
Femur 3.13 10.00 10.38 8.75 9.75
Patella 1.88 5.00 4.75 4.25 3.88
Tibia 2.25 9.13 8.75 6.50 7.75
Metatarsus 8.13 8.00 6.13 7.50
Tarsus 4.38 2.25 2.25 1.88 2.25
Total: 11.64 34.51 34.13 27.51 31.13
Epigynum: Externally (Fig. 8a) a large, strongly sclerotised dark red to black,
sunken rectangular basin, with median septum ('ems') produced posteriorly to form 2
large lobes ('msI') divided by median incision at posterior margin; size of incision
between lobes variable in specimens examined. ranging from short and narrow to
deep and wide; median septum invaginated laterally ('Ii'). Internally (Fig. 8b)
posterior lobes of septum ('goc') enfold exit of efferent sperm ducts ('eco' ) formed
by invagination of septum; spermathecae ('sp') simple, short, closely attached to
bulbous anterior invagination.
Description of male:
Total length: 17.9 mm. Based on an adult male (Figs 9a-b) from Hout Bay
(34°02'S 18°21'E), 3.v.1984. J.Visser (NMSA).
Colour and markings: As in female.
Carapace: CL 9.7, CW 7.4, HW 3.9; thoracic fovea 0.1 wide and 3.2 long,
occupying 0.33 X of CL and 0.43 X of cwo Eyes: Eye diameters: AME 0.45, ALE
0.78, PME 0.55, PLE 0.53; PER 3.1 wide and 1.19 X wider than AER, 0.8 X of HW
and 0.42 X of CW; eye ratio AME:ALE:PME:PLE 1:1.72:1.22:1.17; distances

separating eyes: AME-ALE 0.08, AME-AME 0.28, AME-PME 0.80, ALE-PLE
0.58, PME-PLE 0.50, PME-PME 0.50; MOQA 1.18, MOQP 1.53, MOQL 1.70,
MOQ 1.12 X longer than wide, narrowed anteriorly, with MOQP 1.30 X wider than
MOQA. Clypeus: 0.50 high and 1.11 X diameter AME. Sternum: 4.20 long, SWI
3.25, 1.29 X longer than wide, SW2 2.75. Labium: 1.50 wide, 1.20 long, 1.25 X
wider than long, occupying 0.55 X of SW2. Maxillae: 2.50 long, 1.40 wide, 1.79 X
longer than wide.
Legs: Spination as in Fig. 7. Left leg I abnormally short (regrowth after autotomy).
Measurements:
PaJpus I (R) II III IV
Femur 3.70 10.40 10.30 9.20 10.40
Patella 1.80 4.90 4.90 4.00 3.80
Tibia 1.80 9.40 9.00 7.20 8.10
Metatarsus 8.80 8.80 6.40 8.10
Tarsus 4.50 2.20 2.20 2.00 2.00
Total: 11.80 35.60 35.20 28.80 32.30
Pedipalp: Tibial apophysis (Fig. 9b, 'ta') large, single, situated distally on
retrolateral surface, with 3 major prominences: dorsal, short finger-like projection
directed dorsad; median, long, finger-like, sharp-tipped main prominence directed
ventrad; ventral, swollen base crested by undulating ridge. Embolus (Figs 9a-b) long,
straight ('em'), with short, small, sickle-shaped sclerite apically, inserted medially in
radix region of tegulum ('te'), flanked anterolaterally by long embolar gulley and
lateral sclerite ('1s') and retrolaterally by conductor ('co'); conductor membranous,
long, crinkled, inserted retrolateral to embolus; tegulum ('te') large, heavily
sclerotised, without other apophyses.
Additional material examined: SOUTH AFRICA: Cape: Bredasdorp, Elim, 1 9 ,
1900, R. Baur (SAMC C8447); Cape Town: museum grounds, 19, xii.I926, A. J.
Hesse (SAMC B7148), Highlands Estate, 3318CD, 19, lO.ii.1981, Mrs Krego
(SAMC C249), 19, i.1966, P. L. G. Benoit (MRAC 130.327), University of Cape
Town, 10"19 , 1964-1965 (NMSA), 19 , 1896, F. Purcell (SAMC 643), 29 (SAMC
CI893), Bergvliet: 1 subad.9 and wasp (wasp pulling paralysed spider over half-
door), 18.ii.1986, 10", iii.1984, 10", iii.1986, E. Seekings (NMSA), Camp's Bay: 19,
xii. 1922, Prof. Thoday (SAMC B5867), 20"19, 1896, A. Jeffery & B. Wallace
(SAMC 3189), Claremont: 19, 12.ix.1963, B. Lamoral (NMSA), 19, 1938, Mrs
Barnard (SAMC B9374), 10", in house, 7.iii.1981, Mr Servator (SAMC C254),
Clifton, 20" 19, 3.v.1918, R. Wallace (SAMC B4180-B4181), Gardens, 10",
14.v.1991, N. Larsen, no. 165 (SAMC C2080), Kenilworth, 19 , xi.1897, 19 , 1898
(SAMC 2904 & 2910); Kirstenbosch, Table Mountain, Skeleton Gorge forest,
700-1000 ft, 10" Isubad.9 2juv., under bark of dry logs, 26+29.x.1985, C. E.
Griswold, J. Doyen & T. Meikle-Griswold (NMSA), Kirstenbosch, 33°58'S 18°28'E,
10", 2l.ix.1970, B. Leon, No. 42 (NMSA 11818), Newlands, 19, viii.1983, B.
Kaczmaruk (SAMC C1126), Newlands, 19 , 2l.ix.1970, B. Leon, No. 43 (NMSA),
Plumstead, 10", 25.vi.1963, B. H. Lamoral (NMSA), Signal Hill, 19, v.1896, F.
Purcell (SAMC 788), Table Mt, 3318CD, 10", ii.1982, J. Smuts (SAMC C1050),
Table View, 19, 22.vi.1991, N. Larsen, no. 174 (SAMC C2279), Vredehoek, 10"
30 ANNALS OF THE NATAL MUSEUM. VOL. 37 1996
(stung by AJropompilus ignitus which was then bitten by Palystes), i.1990, Mr Davis
(SAMC Cl729), Wynberg, 19, 1898, H. Beard (SAMC 3202), Wynberg, 19,
ix.1896, 19 , w. L. Sclater (SAMC 641-642); Cape Peninsula: 19 , K. H. Barnard
(SAMC B9129), 10", 1898, F. Purcell (SAMC 2916), Cape Flats, 19 , 1884, J. de
Soizer (SAMC 781),10", German School (SAMC 9114), Hout Bay, 40"49, in house,
14.ii.1984, 15.ii.1984, 3.v.1984, 27.vii.1984, 28.xii.1984, 6.ii.1985, 24.iv.1985,
J.Visser 2925, 2983(x2), 3076, 3319, 3695, 3703, 3825 (NMSA), Hout Bay, 10",
7.xi.1983, 10", 16.ix.1983, 29, x.1983, 10", l.xii.1983, J. V. [John Visser] 2737
(SAMC C1886, C1887, C1945, CI889), Hout Bay, Tierbos, 3.xii.1950 (AMGS),
Hout Bay, 19 Isubad.9, 9.ix.1967, E. S. Ross & A. R. Stephen (CASC), Kalk Bay,
19, xi.1896, H. Beard (SAMC 731), [Muizenberg], St James, 10", xii. 1896, F.
Purcell (SAMC 931), Newlands Forest, juv. 9 , sitting on low vegetation, ll.iii.1992,
N. Larsen, no. 331 (SAMC C2464), New1ands Forest, Paradijs, 19 , 22.vii.1991, N.
Larsen, no. 189 (SAMC C2278); Jonkershoek, 19 , x.I926, R. H. Barnard (SAMC
B7135); Marnre Mission Station [45km N. Cape Town], 19 , 1896 (SAMC 896);
Somerset West, 10", in garden, 18.iv.1984, J. Winstanley (PPRI '84/433); [Somerset
West district], on Kogelberg [34°14'S 18°52'E], 3500 ft, viii.[19]41, T. P. Stokoe
(SAMC); Stellenbosch, 29 (AMGS 2646); Stellenbosch, 19 , x.1904, R. Broom
(SAMC 14238); nr. Swellendam, 10", 1900, R. Baur (SAMC 7879); Cap d.g.H. [Cap
der guten Hoffnung], 29, 8.iii.1858, Ida Pfeiffer (NHMV). Orange Free State:
Brandfontein, 10", l1.xi.1984, A. V. Milewsky (SAMC CI890); Springfield, 19,
x.1984, A. V. Milewsky (SAMC). ZIMBABWE: Salisbury [=Harare], 19 , vii.1965
(NMSA); near Bulawayo, 19 , 1896, Dr Kolbe (SAMC 3845). MOZAMBIQUE:
Delagoa Bay, 19 , [?Eugene Simon] (MNHN 9960).
Palystes martintilmeri sp. n.

Figs 22-27, 29, 31
Type material:
Holotype: adult 0", SOUTH AFRICA: Cape: Cederberg Tourist Park, Kromrivier,
72 km SSE Clanwilliam, el. 3100ft. 17.xi. 1985, C. Griswold, J. Doyen & T. M.-
Griswold (NMSA T496).
Paratypes: SOUTH AFRICA: Cape Province: Cederberg, Kromriver, 1 9 , v.1935,
W. Nieuwoudt (SAMC B8887); Clanwilliam, Elizabeth'sfontein, 20", 1898, Miss
Bergh (SAMC 2908); Clanwilliam, Sneeukop [Cederberg Mountains], 19 , 1898, R.
Patterson (SAMC 3634); Piquetberg [Piketberg], Gutverwacht, 39 30", 1897, R.
Marks (SAMC 2162).
Type locality: Kromrivier (32° 32'S 19°17'E), 72 km SSE. of Clanwilliam, western
Cape, South Africa.
Etymology: Named in honour of Dr Martin Filmer - for many years chairman of the
Spider Club of South Africa - who has done much to promote amateur arachnology.
Diagnosis: Adult female differs from females of P. castaneus and P. stilleri in form
of epigynal septum (Fig. 23), which is short, broad, excavated anteriorly, produced
posteriorly in a transverse, laterally swollen, medially sunken bridge flanked by
bulbous lateral sclerites. In P. castaneus (Fig. 8a) septum forms sunken basin ending
24
26
Figs 23-27. Palystes martinfilmeri sp. n. 23. External female epigynum. 24. Internal epigynum (paratype
female SAMe B8887, Kromrivier). 25. Retrolateral aspect of male pedipalp. 26. Ventral
aspect. 27, anterolateral aspect (holotype male NMSA T496, Krornrivier).
posteriorly in 2 diverging elongate lobes. In P. stilleri (Fig. 32) septum swollen

medially. Adult male differs in form of embolus (Figs 25-27, 29, .31) with sturdy,
round, unexcavated column, reflexed apically, dividing (Figs 29, 31) into ventral,
laterally hooked sperm-duct spur, sub tended by sessile basin-shaped membranous
conductor (Fig. 31), with opposing dorsal, flattened, blunt-tipped 'finger' and in form
of simple tibial apophysis directed distally. In P. castaneus embolar column (Fig. 9a)
straight, not reflexed apically, with conductor elongate, running the length of
embolus, and tibial apophysis trilobate; in P. stilleri embolar column excavated
anterolaterally in form of wide valley (Fig. 28), without apical spur (Fig. 30),
conductor distinctly stalked and apex of simple tibial apophysis reflexed ventrally.
Distribution: Piketberg, Clanwilliam and Cederberg districts of the western Cape,

South Africa.
Description of holotype male:
Total length: 24.75 nun.
Colour and markings: Dorsally cryptically marked with darker and lighter
mottling, overall colouring finely mottled shades of greyish lighter brown and darker
brown; abdomen dorsally with longer pale setae rising above shorter setae that
densely clothe the body, laterally with white fleck markings continuing ventrally on
abdomen and ventral surface of femora; abdomen ventrally with large black to dark
brown shield marking between epigastric groove and spinnerets, area anterior to
epigastric groove dark brown and sparsely covered with short setae; sternum black;
coxae unmarked ventrally but darker anterolaterally.
Carapace: Dorsally 12.50 long, 10.63 wide between leg bases II. Head region 5.50
wide across PER; thoracic fovea 3.38 long, occupying 0.27 X of CL and 0.32 X of
Cw. Eyes: Eye diameters: AME 0.57, ALE 0.85, PME 0.62, PLE 0.62, PER 3.75
wide, wider than AER by 1.15 X and occupying 0.68 X of HW and 0.35 X of CW;
eye ratio AME:ALE:PME:PLE is 1: 1.49: 1.08: 1.08; distances separating eyes:
AME-ALE 0.15, AME-AME 0.34, AME-PME 0.98, ALE-PLE 0.65, PME-PLE
0.69, PME-PME 0.54; MOQ dimensions: MOQA 1.38, MOQP 1.75, MOQL 1.54,
MOQA; clypeus 0.52 high, 0.92 X diameter of AME. Chelicerae: 5.00 long, 2.56
wide, 1.95 X longer than wide. Sternum: 5.19 long, 4.06 wide between coxae II, 1.28
X longer than wide and 3.06 wide between coxae I. Labium: 1.85 wide, 1.69 long,
1.09 X wider than long, occupying 0.60 X of SW between coxae I. Maxillae: 3.14
long, 1.85 wide, 1.70 X longer than wide.
Abdomen: 12.25 long and 8.13 wide.
Pedipalp: Embolus rises on a robust, round column to recurve over bulb apically
where it divides into a horizontally curved spine carrying sperm tube and an
opposing vertically flattened blunt 'finger'. The spine is subtended by a dish-shaped
white to diaphanous membranous conductor. The tibial apophysis is large, robust,
blunt-tipped and reminiscent of the profile of a thumb cocking the hammer of a
revolver.
Legs: Spines typical of genus with femora I-III(3:2:3), femora IV(3:2:2); patellae
I-IV (1: 1); tibiae I-IV with ventral, antero- and retrolateral spines as for genus, but
with dorsal basal and apical spines present on legs I-IV. Measurements:
Palpus I II III IV
Femur 5.13 16.25 17.13 14.38 16.88
Patella 1.88 6.13 6.38 5.00 5.00
Tibia 3.25 17.75 17.75 13.38 14.63
Metatarsus 16.88 17.13 12.25 16.25
Tarsus 5.00 3.75 3.38 3.13 3.50
Total: 15.26 60.76 61.77 48.14 56.26
Description offemale (SAMC B8887):

Total length: 35.26 rnm.
Colour and markings: Abdomen dorsally with pulmonary marking outlined in
darker brown, laterally with some darker spotting, ventrally with large, dark, rich
CROESER: REVISION OF PALYSTES (ARANEAE: HETEROPODJDAE) 33
brown bell-shaped marking between epigastric groove and spinnerets. Sternum dark to
black. Coxae dark anteriorly and basally but not ventrally. Carapace dorsally with
paler median line, edged with dark brown, from PME to thoracic fovea. Femora light
yellow-brown ventrally, tibiae I-IV with 2 dark and 2 pale bands alternating ventrally.
CWo Eyes: Eye diameters: AME 0.63, ALE 1.00, PME 0.77, PLE 0.77, PER 5.00
MOQA. Clypeus height 0.82, 1.29 X diameter of AME. Chelicerae: 6.31 long, 3.69
Abdomen: 19.3810ng and 14.38 wide.
Epigynum: Externally (Fig. 23), median septum excavated, wide, ending
posteriorly in a transverse, laterally bulbous, bridge, flanked anterolaterally by
bulbous lateral epigynal sclerites. Internally (Fig. 24), spermathecae widely
separated, lateral to septum, just visible laterally at foot of anterior support collar
formed by anterior invagination of septum.
Legs: Spine pattern typical of genus, but femora I-III(3:2:3), femur IV(3:2:2);
patellae(l :0: 1); tibiae I-IV with usual spination and basal and apical spines dorsally.
Measurements:
Palpus I II III IV
Femur 5.63 17.75 18.00 15.00 16.50
Patella 2.50 7.38 7.50 5.75 5.75
Tibia 3.50 16.50 16.50 12.88 14.38
Metatarsus 15.88 16.25 11.63 15.00
Tarsus 6.25 3.38 3.38 2.75 3.13
Total: 17.88 60.89 61.63 48.01 54.76
Additional material examined: None.
Palystes stilleri sp. n.

Figs 28, 30, 32-36
Type material:
Holotype: adult cJ , SOUTH AFRICA: Cape: Hottentots Holland Mts. 3500',
i.1916, K. Barnard (SAMC B2019).
Paratype: adult 9 , SOUTH AFRICA: Cape: Jonkershoek, Stellenbosch, in rotting
log, 17.v.1975, M. Stiller (PPRI '80/193).
Type locality: Hottentots Holland Mountains, south-western Cape, South Africa.
Etymology: Named after a keen Palystes collector, Martin Stiller, who found the
paratype of this species and also the holotype and paratypes of P. ansiedippenaarae
sp. n.
Figs 28-36. Palystes species. 28-31. Comparison of apical structure of embolus in P martinfilmeri sp. n.
and P stilleri sp. n. 28-29. Anterolateral aspect. 28. P. stilleri. 29. P. martinfilmeri. 30-31.
Retrolateral aspect. 30. P. stilleri. 31. P. martinfilmeri. 32-36. P. stilleri. 32. External female
epigynum. 33. Internal epigynum (paratype female, PPRI AcAT '80/193, Jonkershoek,
Stellenbosch). 34. Retrolateral aspect, male pedipalp. 35. Ventral aspect. 36. Anterolateral
aspect (holotype male, SAMC 82019, Hottentots Holland Mts.).
Diagnosis: Epigynum (Fig. 32) with anteriorly narrowed, bowed (not medially
excavated) septum, widened and flattened posteriorly, not unlike a broad, flat nose.
Embolus excavated anterolaterally in a wide, shallow trough (Fig. 28), reflexed

apically, ending in broad, flattened, blunt-ended blade (Fig. 30) with sperm tube and
opposed by flattened opposing 'finger'; apex subtended by stalked basin-shaped
membranous conductor (Fig. 30). In P. castaneus the embolar column is straight, not
reflexed apically, and the conductor is elongate, running the length of the embolus
(Fig. 9b). In P. martinfilmeri the embolar column is not excavated anterolaterally
(Fig. 29), the vertically flattened finger-like projection opposes a large spur apically
and the conductor is sessile (Fig. 31). The ventral markings of the abdomen (a broad
fawn-brown to yellow patch transversely interrupting the median dark shield
marking) also clearly distinguish P. stilleri from the other two species which have
solid black to dark brown shield markings.
Distribution: Hottentots Holland Mountains, immediately southeast of Stellenbosch,
in the south-western Cape Province of South Africa.

Total length: 26.25 mm.
Colour and markings: Generally pale brown. Abdomen dorsally with a number of
very long pale setae rising above shorter setae that densely clothe the body; laterally
with a few white fleck markings posterior to epigastric groove; ventrally with broad
transverse pale brown marking, edged with darker brown, immediately posterior to
dark brown crescent posterior to epigastric groove, followed by small dark brown
marking anterior to spinnerets; area anterior to epigastric groove dark brown and
sparsely covered with short setae. Sternum black. Legs: coxae unmarked ventrally
but darker anterolaterally; femora ventrally with some white fleck markings.
cwo Eyes: Eye diameters: AME 0.52, ALE 0.83, PME 0.54, PLE 0.54, PER 3.56
MOQ 0.91 X longer than wide; narrowed anteriorly, with MOQP 1.38 X wider than
wide, 1.95 X longer than wide. Sternum: 5.31 long, 4.06 wide between coxae 11,1.31
Pedipalp (Figs 28, 30, 34-36): Embolus column excavated anterolaterally in a
wide, shallow trough and reflexed apically, ending in a broad, flattened, blunt-ended
blade containing sperm tubule, opposed by vertically flattened 'finger' (Fig. 30);
apex subtended by distinctly stalked, basin-shaped membranous conductor. Tibial
apophysis (Figs 34, 35) large, robust, blunt-tipped but more slender than that of P.
martinfilmeri and reflexed ventrally across base of paracymbium.
Legs: Tarsi of left legs I and II missing. Spines typical of genus with femora
I-III(3:2:3), femora IV(3:2:2); patellae I-IVO: 0; tibiae I-IV with ventral, antero-
and retrolateral spines as for genus, but with dorsal basal and apical spines present on
legs I-IV. Measurements:
Palpus I II III IV
Femur 5,13 16.13 16,00 14.13 16.13
Patella 2,00 6,50 6.75 5,63 5.25
Tibia 3.13 15.88 15.25 11,88 13.25
Metatarsus 15,75 15.00 11.25 14.00
Tarsus 6.00 3.13 3.38 2,50 2.88
Total: 16.26 57,39 56.38 45.39 51.51
Description offemale (PPRI '801193):

Colour and markings: Same as holotype male described above.
MOQA. Clypeus 0.54 high, 1.09 X diameter of AME. Chelicerae: 4.38 long, 2.44
1.59 X wider than long, occupying 0.68 X of SW between coxae L Maxillae: 2.92
Epigynum (Figs 32, 33): Epigynum externally with an anteriorly narrowed, bowed,
not medially excavated, septum, widening and flattening posteriorly, not unlike a
broad, flat nose flanked laterally by swellings. Internally, spermathecae lie
immediately posterior to foot of support collar and project inwards (Fig. 33).
Legs: Spirie pattern as for genus. Measurements:
Palpus I II III IV
Femur 4.38 13.13 13.38 11.38 12.75
Patella 2.25 5,88 6.13 4.88 4,75
Tibia 3.13 12.50 12.50 9,00 10,75
Metatarsus 12.50 12.13 8,38 10,63
Tarsus 4.75 3.00 2.63 2,13 2.63
Total: 14.51 47.01 46.77 35,77 41.51
Palystes lunatus species group

Characterised by: Abdomen ventrally with large transverse dark brown to black
crescent-shaped marking immediately posterior to epigastric groove, followed by
distinct round white spots against a tawny yellow to rich reddish-orange background,
spotting continuing ventrally on femora; well-developed 'tongue' -shaped median
lobe of epigynal septum strongly produced ventrally, projecting posteriorly; broad,
reflexed embolus of male pedipalpal bulb with vestigial to prominent hooked basal
spur (vestigial tegular sclerite). Sternum with either a single transverse band level
with anterior faces of coxae II, or first band followed by a second band level with
anterior faces of coxae III. Second band may be medially interrupted or may join up
with first band medially (varies within species).
Included species: Palystes lunatus Pocock, 1896; P. crawshayi Pocock, 1902; P.
leppanae Pocock, 1902; P. peromatus Pocock, 1900; P. karooensis sp. n; P. stuarti sp. n.
Distribution: Western and eastern Cape, Transkei, Lesotho and southern Natal (Fig.
76)
Palystes lunatus Pocock, 1896

Figs 37-38, 76
Palystes lunatus Pocock, 1896: 60, PI. vm, Fig. 5. Holotype: adult)' , oS. Africa', (Dr Quain) (BMNH
'no reg. no.') [examined].
Remarks: Pocock's description was based on an adult female donated by Dr Quain.
An adult female in BMNH (label data given above) is undoubtedly the holotype of P.
lunatus; label data, markings and form of the ventral epigynum agree with Pocock's
description (p. 60) and illustration (PI. VIII, Fig. 5). It has not been possible to
restrict the type locality further than the vague'S. Africa' given by Pocock. The
specimen was not recorded in the register when donated to BMNH and I have not
been able to find out anything about Dr Quain or where he was likely to have
collected the specimen. To add to the problem, I have found no other conspecific
females. I do not think the form of the epigynum is aberrant, as there are differences
in its basic structure compared with its relatives in the species group. Apart from the
considerable length and narrowness of the median lobe of septum ventrally (more
than twice longer than wide), the epigynum internally has a greatly produced efferent
sperm duct which folds back on itself, extending medially well beyond the base of
the spermathecae.
Diagnosis: The form of the epigynum with its posteriorly produced median septum,
and the ventral abdominal markings with distinct, large white cream blotches
posterior to the postepigynal dark brown transverse bar, place the species in the
lunatus species group. The greatly produced median septum (Fig. 37), more than
twice as long as wide, and the form of the internal efferent sperm duct (Fig. 38),
separate it from other species in the group.
Distribution: Known only from the type specimen and its labelled locality 'South
Africa'. Like several other species in this group, P. lunatus may be restricted to a
relatively small locality, probably somewhere in the montane interior of the eastern
Cape Province of South Africa.
Colour and markings: Abdomen dorsally with pulmonary marking, laterally darker
on top, ventrally with dark brown transverse bar immediately posterior to epigastric
groove and with large white to cream rough spots to blotches against a light to rich
~"".'-""~'"''
.-~ ... ,
'-.
1.-.,
i.,. ......
L.,
i
\
i
37 38
39
42
~
44
Figs 37-44. Palystes lunatus Pocock, 1896 and P. crawshayi, Pocock, 1902. 37-38. P. lunatus. 37.
External female epigynum. 38. Internal epigynum (female holotype, BMNH). 39-44. P.
crawshayi. 39. External female epigynum. 40. Internal epigynum (neotype female SAMC
14429, Lesotho). 41. Ventral aspect, male pedipalp. 42. Detail of ridge-like reduced tegular
sclerite at base of embolus. 43. Ventral aspect, tibial apophysis. 44. Retrolateral aspect, tibial
apophysis (described male, SAMC 14429).
red-brown field. Sternum with 2 dark transverse bars in line with leg bases II and III,
anterolateral faces of coxae dark, femora ventrally dark with pale patches. Tibiae
ventrally dark with a median and an apical broad pale bar.
CWO Eyes: Eye diameters: AME 0.57, ALE 0.80, PME 0.58, PLE 0.58, PER 3.88
MOQA. Chelicerae: 4.50 long, 2.50 wide, 1.80 X longer than wide. Sternum: 5.44
long, 3.81 wide between coxae II, 1.43 X longer than wide and 2.81 wide between
coxae I. Labium: 2.] 8 wide, 1.54 long, 1.42 X wider than long, occupying 0.78 X of
SW between coxae 1. Maxillae: 3.35 long, 2.12 wide, 1.58 X longer than wide.
Epigynum: Median septum of epigynum greatly produced posteriorly, more than
twice as long as wide. Interior epigynum with spermathecae dorsal to support collar
and reflexed outwards. The greatly produced efferent sperm duct folds back on itself,
extending medially well beyond base of spermathecae.
Legs: Spine pattern and markings as for genus. Measurements:
Palpus I II III IV
Femur 4.25 13.25 13.50 12.13 13.13
Patella 2.38 6.25 6.25 5.00 5.13
Tibia 2.88 12.25 12.50 9.75 10.63
Metatarsus 12.38 12.25 8.75 11.13
Tarsus 5.13 2.63 2.75 2.50 2.75
Total: 14.64 46.76 47.25 38.13 42.77
Palystes crawshayi Pocock, 1902

Figs 39-44, 76
Palystes crawshay; Pocock, 1902: 27. Syntypes: 2, at least one an adult female: 'Maseru' [Lesotho] (lost,
not in BMNH). Adult 9 neotype here designated: Lesotho, 1905, W. C. Wroughton (SAMe
14429).
Remarks: The two syntypes collected by Mr Crawshay have not been located but a
number of conspecific adult female and male specimens from Maseru, Lesotho, are
in the collections of the SAMC (donated in 1905 by W. C. Wroughton) and BMNH
(donated in 1907 and 1912 by Mrs L. Wroughton and Dr Maclean). The females all
have the characteristic transverse septum described by Pocock. At present this is the
only known locality for this species, and these specimens remain the only known
collected material. As no syntype material appears to be extant, one of the females in
SAMC collection has been designated as a neotype and is described below.
Diagnosis: The transverse, shallow 'tongue' of the median lobe of the epigynal septum
(up to 6 X wider than long) (Fig. 39) is characteristic of females of this species, as are
the anteriorly produced spermathecae inserted laterally to the internal support lobes
(Fig. 40). The short, flat, trilobate tibial apophysis (Figs 43, 44), the ridge (in place of a
spur) at the base of the embolus (Fig. 42), and the absence of a median channel on the
embolus, clearly distinguish the male from other species in the group.
Distribution: Known only from Lesotho.
Description offemale neotype:
Colour and markings: Specimen faded to a light brownish yellow (probably from
exposure to light), although remains of characteristic markings of species group are
evident (2 black bars across sternum, dark post-epigynal transverse bar and white
spotting of abdomen ventrally). Pocock's (1902: 27) description of his relatively
fresh specimens is characteristic for the lunatus species group as a whole, and is
repeated here as a guide: 'Colour of dorsal surface a fairly uniform greyish brown,
obscurely mottled like the back of tarsi; pale clypeal band on base and two superior
external pale bands on mandible; sternum yellow, with two black bands; coxae
yellow; femora greyish black below, conspicuously spotted with yellowish grey;
epigastric area yellow; a transverse black band behind the epigastric fold, the rest of
the lower surface dark red variegated with white spots.'
0.62, PME-PME 0.46; MOQ dimensions; MOQA 1.23, MOQP 1.51, MOQL 1.07,
1.58 X wider than long, occupying 0.70 X of SW between coxae L Maxillae: 2.46
Epigynum: Externally, median septum lobe transverse, not produced (Fig. 39).
Internally, spermathecae produced anteriorly and inserted laterally to supporting
lobes (Fig. 40).
Legs: Spine pattern typical of genus, but with left femur III missing a retrolateral
spine and with femora IV (left and right) both 3:2: 1 with only a single retrolateral
spine instead of the more usual two retrolateral spines. Tibial dorsal spines also
varied with left I and II and right I with both distal and proximal spines, rest of tibiae
all with only a single, distal, spine. Measurements:
Palpus I II ill IV
Femur 3.~ ll.~ Il.~ 10.13 11.38
Patella 1.88 5.00 5.00 4.13 4.00
Tibia 2.50 10.63 11.00 8.75 9.50
Metatarsus 10.63 10.50 7.50 9.38
Tarsus 4.38 2.88 2.75 2.13 2.50
Total: 12.26 40.89 41.13 32.64 36.76
Description o/male:
The description is based on an adult male in SAMC, taken from the same jar as the
neotype female, and labelled: LESOTHO, 1905, W. C. Wroughton (SAMC 14429).
Colour and markings: Faded, and abdominal cuticle split away (as in a tenera]
specimen). Otherwise as in female.
Carapace: Dorsally 9.38 long, 7.50 wide between leg bases II. Head region 4.00 wide
across PER; thoracic fovea 2.38 long, occupying 0.25 X of CL and 0.32 X of CWO Eyes:
Eye diameters: AME 0.37, ALE 0.62, PME 0.38, PLE 0.38, PER 2.69 wide, wider than
AER by 1.16 X and occupying 0.67 X of HW and 0.36 X of CW; eye ratio
AME:ALE:PME:PLE is 1:1.67:1.04:1.04; distances separating eyes: AME-ALE 0.12,
AME-AME 0.35, AME-PME 0.69, ALE-PLE 0.49, PME-PLE 0.46, PME-PME 0.46;
MOQ dimensions: MOQA 1.08, MOQP 1.38, MOQL 1.38, MOQ 1.00 X longer than
wide; narrowed anteriorly, with MOQP 1.29 X wider than MOQA; clypeus 0.38 high,
1.04 X diameter of AME. Chelicerae: 3.38 long, 1.88 wide, 1.80 X longer than wide.
Sternum: 4.06 long, 3.00 wide between coxae II, 1.35 X longer than wide and 2.50 wide
between coxae I. Labium: 1.54 wide, 1.23 long, 1.25 X wider than long, occupying 0.62
X of SW between coxae I. Maxillae: 2.06 long, 1.32 wide, 1.56 X longer than wide.
Pedipalp (Figs 41-44): Spur at base of embolus reduced to a long blade-like ridge
(Fig. 42). Tibial apophysis short, flattened, rectangular and three-lobed (Figs 41,
43-44). Embolus flattened, without median channel.
Legs: Spines typical of genus but femora IV(3:2: 1) missing a retrolateral spine on
both legs; all tibiae dorsally with both proximal and distal spines present.
Measurements:
Palpus I II III IV
Femur 2.75 11.63 11.75 10.00 11.50
Patella 1.75 4.50 4.63 3.75 3.75
Tibia 2.25 11.38 11.38 8.75 9.63
Metatarsus 11.88 11.63 8.13 9.75
Tarsus 4.25 3.00 2.88 2.25 2.50
Total: 11.00 42.39 42.27 32.88 37.13
Additional material examined: LESOTHO: [Maseru], 29 , 1905, W. C. Wroughton

(SAMC 14429); Maseru, 39 4d 2subad.d Ijuv., 1907, Dr Maclean & L.Wroughton
(BMNH 07.10.8.4-15 part); Maseru, 49 2subad.9, 1912, Mrs L. Wroughton
(BMNH 1912.7.27.4-8).
Palystes leppanae Pocock, 1902

Figs 57--62, 76
Palystes leppanae Pocock, 1902: 23, PI. III, Fig. 9. Holotype: adult 9, 'Tea Fountain, near
Grahamstown', [1901], Miss Leppan (BMNH 1901.3.13-258) [examined].
Remarks: An adult female in BMNH (labelled as above), which matches Pocock's
description and illustration is undoubtedly the female described by Pocock. The
specimen is well preserved with the legs intact; it has a small white steel pin 14 rnm
long inserted transversely through the epigastric groove (possibly inserted by Pocock
to lift the epigynal plate for drawing).
../.v ~
/'''''-
;.-
50
56~ 54
Figs 45-56. Palystes karooensis sp. n. 45-51. Holotype female (NMSA T501) and paratype male
(NMSA T502) from Stolshoek (near Beaufort West). 45. Ventral aspect, female, showing
markings of abdomen, sternum and coxae. 46. External female epigynum. 47. Internal
epigynum. 48. Cheliceral dentition, male paratype, showing additional apical teeth on antero-
and retrolateral margins of left chelicera and antero-margin of right chelicera. 49. Ventral
aspect, male pedipalp. 50. Detail of thorn-like vestigial tegular sclerite at base of embolus. 5l.
Retrolateral aspect, tibial apophysis. 52-56. Female from Cedarville (near Mvenyane) (AMGS
2344) and male from Ntolas (AMGS 2110), south-western KwaZulu-Natal. 52. External
female epigynum. 53. Internal epigynum. 54. Anterolateral aspect, left pedipalp. 55. Ventral
aspect, male tibial apophysis. 56. Retrolateral aspect, tibial apophysis.
Diagnosis: The females of P. Zeppanae and P. karooensis have a characteristic heart-

shaped, round-shouldered epigynum (Figs 46, 57) with a posteriorly produced
median lobe which varies from slightly longer than wide to slightly wider than long.
However, the two species are clearly distinguished by the structure and position of
the spermathecae internally. In P. Zeppanae the spermathecae (Fig. 58) are straight,
short and stout and project anterolaterally away from a common median line at an
angle of 45 0 , whereas in P. karooensis the spermathecae (Fig. 47) are produced
laterally, are thin and long and parallel to the epigastric groove at an angle of 90 0 to
the common median line, and are hooked apically. In the male of P. leppanae the
spur at the base of the embolus (Fig. 59) is greatly reduced to a low ridge, and the
embolus itself is deeply channelled along its anterior face (Fig. 62). In all other
species in the group, the embolus column is entire and not channelled.
Distribution: Grahamstown and Alicedale districts, eastern Cape, South Africa ..
Colour and markings: From above, a mottled greyish brown with an outlined
cardiac marking dorsally on abdomen flanked by a series of paired paler patches.
Abdomen ventrally with typical dark brown to black crescentic marking immediately
posterior to epigastric groove followed by a profusion of small round white spots
against a rich reddish brown background. White spots continue on the otherwise dark
ventral surface of femora. Femora dorsally lighter with scattered darker spots.
Sternum with 2 medially interrupted transverse black bands in line with anterior
faces of coxae II and III and a pair of dark spots in line with the anterior face of
coxae IV. Metatarsi and tarsi dark. Carapace dorsally with paler median line running
from between PME to thoracic fovea.
wide across PER; thoracic fovea 3.50 long, occupying 0.29X of CL and 0.36X of
long, 2.22 wide, 1.26 X longer than wide. Cheliceral margin teeth typical of genus (3
retromarginal and 3 anteromarginal teeth but right chelicera has a small fourth,
additional retromarginal tooth apically).
Epigynum (Figs 57-58): Externally, heart-shaped with obliquely sloping shoulders
and a relatively shallowly produced median lobe which is a little wider than long
(Fig. 57). Internally, spermathecae stout, short, straight and project anterolaterally at
45 to a common median line (Fig. 58).
Legs: Spine pattern typical of genus, Measurements:

Palpus I II III IV
Femur 4.25 12.50 13,13 11.25 12.75
Patella 1.88 5.75 5.75 5.00 5.00
Tibia 2.88 11.00 11.63 8.88 10.00
Metatarsus 10.88 10.88 7.75 10.00
Tarsus 5.00 2.75 2.63 2.13 2.38
Total: 14.01 42.88 44.02 35.01 40.13
Described reference male: Cape: Grahamstown, 6 Gilbert Street, found in house,
1O.vii.1982, Mrs Julia Stone (NMSA).
Colour and markings: As female except abdomen more orange than reddish brown
ventrally.
Carapace: Dorsally 8.88 long, 7.88 wide between leg bases II. Head region 4,44 wide
across PER; thoracic fovea 2.75 long, occupying 0.31 X of CL and 0.35 X of CWo Eyes:
AME:ALE:PME:PLE is 1: 1.81: 1.19: 1.19; distances separating eyes: AME-ALE 0.11,
wide; narrowed anteriorly, with MOQP 1.36 X wider than MOQA; dypeus 0.46 high,
X of SW between coxae L Maxillae: 2.18 long, 1.32 wide, 1.65 X longer than wide.
Cheliceral margin teeth typical of genus with 3 retromarginal and 3 anteromarginaJ teeth
except that there is a small, additional, apical retromarginal tooth on left chelicera.
Pedipalp: (Figs 59-62) Spur at base of embolus reduced to a low ridge (Fig. 59).
Stem of embolus deeply grooved (arrow, Fig. 62) anterolaterally along its length,
widening apically into a flat blade. Conductor membranous and recurved below
embolus apically. Tibial apophysis elongate with a proximal lobe forming a notched
apex from retrolateral aspect (Fig. 60).
Legs: Spines typical of genus. Measurements:
Palpus I II III IV
Femur 3.13 11.50 11.88 10.13 11.88
Patella 1.75 4.88 5.13 4.00 4.13
Tibia 1.88 10.88 10.88 7.88 9.38
Metatarsus 10.63 10.63 7.63 8.88
Tarsus 4.00 2.75 2.75 2.25 2.50
Total: 10.76 40.64 41.27 31.89 36.77
Additional material examined: SOUTH AFRICA: Cape: Alicedale: 1 9 (AMGS

2561), 19 (AMGS 1816), 10' (AMGS 2349); Grahamstown: 19 (AMGS 2924),
19 , 22.x.1905, Miss M. Pinnock (AMGS 687), 'Penesck', 1 9 (AMGS 6363),
Kowie Road, 3 miles fro Stones Hill, 10', vii.1945 (AMGS).
57 58
61
Figs 57-62. Palystes leppanae Pocock, 1902. 57. External female epigynum. 58. Internal epigynum
(holotype female, BMNH 1901.3.13-258). 59. Ventral aspect, male pedipalp. 60. Retrolateral
aspect. 61. Anterolateral aspect. 62. Detail of anterolateral aspect of embolus (described male.
NMSA, 6 Gilbert Street, Grahamstown).
Palystes peromatus Pocock, 1900

Figs 63-69, 76
Palystes peromatus Pocock, 1900: 331, PI. III, Fig, 10. Syntype series: I adult 9 (here designated as
lectotype], 2 subadult 9 and I juvenile 9 (here designated as paralectotypes], South Africa:
Cape Province: 'Queenstown', 'presented \5.v.99', Capt. C. K. Bushe (BMNH 1899.7.28.3-6)
[examined].
Remarks: Pocock briefly described an adult female and illustrated its epigynum,
from 'Queenstown, Cape Colony', donated by Captain C. K. Bushe. There is a vial
in BMNH with an adult female, 2 subadult females and a juvenile, all belonging to
the same species and so labelled above. The label data agree with information in
the description, and the adult female matches Pocock's description and illustration
of the epigynum. The total length of the adult female specimen is 22.63 mm which
is very close to the length given by Pocock (25 mm); the discrepancy is probably
the result of shrinkage of the abdomen in alcohol. Although Pocock's description is
that of an adult female, he does not specify how many specimens he saw. The vial
contents suggest that he probably sawall four specimens (since four specimens
donated by Captain Bushe were entered together in the BMNH register) and that
the four form a syntype series. For this reason I have designated the adult female,
which was illustrated by Pocock (Plate III, Fig. 10), as the lectotype. It has been
separated from the remainder of the syntype series, which now become
paralectotypes.
Diagnosis: Median lobe of female epigynal septum slightly longer than wide to
slightly wider than long (Figs 63, 68, 70) (as opposed to very much wider than long
or much longer than wide as in other species in this group). Internally spermathecae
are straight, stout, inserted internally on supporting lobes, produced laterally and lie
horizontally at 90 to the median plane (Figs 64, 69). In all other species the
spermathecae lie between 45 and 2 to the median plane or are horizontal but hooked
apically (P karooensis). In the male, the tibial apophysis projects ventrally and is an
entire, blunt-tipped attenuate spur. In other species, the apophysis is bifurcate or
trifurcate. The blade of the embolus is not grooved proximally as is P leppanae. The
membranous conductor, in addition to recurving apically over the embolus as in P.
karooensis, differs from all other species in the group in that it also recurves over the
proximal lateral margin of the embolus blade. Sternal markings of both sexes
throughout the species range is variable, ranging from a single transverse band at the
anterior faces of coxae II with a second, medially interrupted band at coxae ill, to 2
uninterrupted bands at the same position, which mayor may not be connected medially.
Distribution: From the Amatola Mountains and Queenstown district (eastern Cape)
through Transkei to Port Edward in south-eastern Natal, South Africa.
Redescription of lectotype female:

Colour and markings: Specimen somewhat faded but markings still clear.
Abdomen dorsally mottled greyish brown, with an outlined cardiac marking
medially. Abdomen ventrally with typical dark brown to black crescentic marking
immediately posterior to epigastric groove, followed by round white spots against a
faded reddish brown field. White spots continue on ventral surface of femora.
Sternum with single transverse black band in line with anterior faces of coxae II
followed by a medially interrupted bar at coxae III (similar to ventral markings of a
female from Port Edward (Fig. 67». Carapace dorsally with paler, medial line
running from between PME to thoracic fovea.
eye ratio AME:ALE:PME:PLE is 1: 1.50: 1.1 0: 1.10; distances separating eyes:
Epigynum (Figs 63-64): Externally with well developed, produced, median septum
lobe and narrow neck (Fig. 63). Externally, spermathecae (Fig. 64) straight,
horizontal (90 to median longitudinal line), inserted internally on supporting lobes
and project laterally (away from the median line). Illustrations of an additional
female from Queenstown (Figs 68-70) show minor variations in the external
epigynum and internal support collar.
Legs: Spine pattern typical of genus, but with femora IVan both sides 3:2:2. Both
proximal and distal tibial dorsal spines are present on tibiae I-III on both sides, while
on tibiae IV on both sides only distal dorsal spine present. Measurements:
Palpus I II III IV
Femur 3.75 12.00 12.13 10.63 11.88
Patella 2.25 5.25 5.38 4.38 4.38
Tibia 2.75 10.63 10.63 8.38 9.75
Metatarsus 10.00 9.75 7.25 9.25
Tarsus 4.25 2.75 2.63 2.00 2.13
Total: 13.00 40.63 40.52 32.64 37.39
Based on an adult from: Natal: Port Edward, Blencathra Farm, 31°02'S 30 0 1O'E,
5km NW. of Port Edward, 335 m, inside house, 2.iv.1983, June Stannard (NMSA).
Colour and markings: As in female except: specimen not faded, and paired paler
patches flanking cardiac marking evident down back of abdomen. Femora dorsally
mottled with black. Second transverse black band of sternum continuous and
connected medially with anterior band. Sternal bands in other males, including those
from Port Edward, vary from form described above to a single transverse band with a
medially interrupted second band at coxae III.
------~-.----- ... ----- ...

! .-,0' -... ~... , .....
............ "'.;
,
,/""
\.
,
••..
i
"
63
.•. " ........ .
68
Figs 63-70. Palystes peromatus Pocock, 1900. 63. External female epigynurn. 64. Internal epigynum
(lectotype female, BMNH 1899.7.2.8.3-6, Queenstown). 65. Ventral aspect. male pedipalp. 66.
Retrolateral aspect, tibial apophysis (AMGS 2052, Shawbury, Qumbu, Transkei). 67. Ventral
aspect of spider, showing markings (NMSA, Port Edward, April 1984). 68. External female
epigynum. 69. Internal epigynum. 70. Lateral aspect, median septum (AMGS, Queenstown, G.
Rattray).
0.55, PME-PME 0.54; MOQ dimensions: MOQA l.23, MOQP 1.54, MOQL l.69,
Pedipalp (Figs 65-66): Prominent, hooked spur at base of embolus (Fig. 65). Stem
of embolus entire, not grooved along its length, widening into a flat blade (Fig. 65).
Conductor membranous and recurved over blade of embolus apically and also over
proximal lateral margin (Fig. 65). Tibial apophysis elongate, apically blunt-pointed
but entire (Fig. 66).
Legs: Spines typical of genus but femora IV(3:2:2) on both sides. Tibiae dorsally
with both proximal and distal spines present on all legs except right tibia III with
only a distal spine. Measurements:
Palpus I II III IV
Femur 4.00 13.38 14.25 12.38 13.75
Patella 1.88 5.38 5.50 4.13 4.00
Tibia 2.50 13.25 13.38 10.38 11.50
Metatarsus 12.75 12.75 9.38 11.38
Tarsus 4.63 3.13 3.13 2.50 2.88
Total: 13.01 47.89 49.01 38.77 43.51
Additional material examined: SOUTH AFRICA: Cape: Alice: Hogsback Mountain

'One', +6100 ft, 19 , 16.x.1982, base of cliffs, SW aspect, under stone, P. G. Hawkes
(NMSA); Hogsback, 32°36'S 27°01 'E, 1 9 , 11.iii.1979, P. M. C. Croeser (NMSA);
Amatola Mountains, Hogsback Forestry camp site, 32.36S 26.56E, 19, 14.xii.1985
(on flattened, ovate, orange, unprotected egg sac, 30 X 15 mm, fastened by silk
strands to underside of protected overhanging rock), B. Londt (NMSA);
Queenstown, 19 , G. Rattray (AMGS); Queenstown, 29 2d Isubad.9 , E. T. Wells
(BMNH 01.3.12.3-7); Tarkastad, 1903, R.Broom (SAMC 13348). Transkei: 19 , W.
Roberts (AMGS 4878); Qumbu [40 km N. Umtata], Shawbury, 2d29 5juv. (AMGS
2052). Natal: Port Edward, 31°03'S 30 0 13'E, 19, viii.1985, J. Stannard (NMSA);
Port Edward, Blencathra Fann, 31°02'S 30°lO'E, 5 km NW. Port Edward, 335 m, Id
(in pot plant), ld, 25.iii.1983 (early morning in house), 2d, 16.iv.1983 (in house 11
pm - rainstorm), 19 , iv.1984 (at night in house), Id, iv.1984 (caught in same room
with d P. superciliosus), Id, iv.1986 (in house a night), J. Stannard (NMSA).
Palystes karooensis sp. n.

Figs 45-56, 76
Type material:
Holotype: Adult 9 , SOUTH AFRICA: Cape: Beaufort West, Klipbank 3222 Ad
[between 32°15'S-32°30'S & 2215'E-22°30'E], Stolshoek, 2,800 ft, l.ii.85, J. Brau
F15a (NMSA T501).
Paratypes: SOUTH AFRICA: Cape: Beaufort West, Stolshoek, Karoo National
Park, Id, 7.iv.1985, H. Braack (NMSA T502), Aberdeen [near Graaff-Reinetl,
'Leliefontein' farm, 19 , under stone, 1972-1973, M. Stiller (PPRI 'AcAT '80/168);
Cape Town environs, 19, 1967, P. L. G. Benoit (MRAC 146.161); Cradock, 19,
xii.1976 (SAMC C1170); De Wet [?De Wet: 6 Ian NW. Worcester], 19, 9.x.1940
(TMSA); Graaff-Reinet, 19 2d Isubad. d , 26.viii.1920, Rev. J. H. Whaits (AMGS
4281); 19 (AMGS 1830); Murraysburg, 31°58'S 23°45'E, 19, on wall in house,
20.vii.1987, A. van den Berg (PPRI 'AcAT '87/841); Waterford [33°04'S 25°00'E,
Klein Winterhoek Mountains, inland from Port Elizabeth], 19 , iii.1984, B. Rubidge
(NMBA097).
Type locality: Stolshoek, Karoo National Park [ca 32°18'S 22°30'E], immediately
NW. of Beaufort West, Great Karoo region of the Cape, South Africa.
Etymology: Named after the Cape's Great Karoo region, in which most specimens
have been collected.
Remarks: Further collecting is necessary to define more clearly the range of this
species which includes an outlier population in the Cedarville-Kokstad area of north-
eastern Transkei and south-eastern Natal. Further material from the outlier population
may reveal specific differences that separate it from the Karoo population.
Diagnosis: P. karooensis most closely resembles P. perornatus in general body form
and the genitalia, but the two species are relatively easily distinguished from each
other. In adult female P. karooensis the median lobe of the septum (Figs 46, 52) ranges
from wider than long to slightly longer than wide, compared with very much longer
than wide in P. perornatus (Figs 63, 68), and the spermathecae are very long, thin and
apically distinctly hooked towards the anterior (Fig. 47), as opposed to short, stout and
straight (Fig. 64). In adult male P. karooensis the tibial apophyis is bifurcate or deeply
notched apically (Figs 49,51,55,56), as opposed to entire (Fig. 66).
Distribution: Chiefly the interior montane areas of the Great Karoo including the
districts of Beaufort West, Murraysburg, Graaff-Reinet and Cradock, Cape, South
Africa. Also including the Winterhoekberge Mountains north-west of Port Elizabeth.
Description of holotype female:
Colour and markings: Dorsally mottled dark brown, black and light brown;
abdomen with a dark outlined cardiac marking flanked by paired paler patches
diminishing in size posteriorly. Ventrally, a black transverse bar immediately
posterior to epigastric groove. Posterior to the bar are numerous round white spots
against a rich dark red-brown background; the spots continue on the underside of the
otherwise dark femora. Femora ventrally becoming orange apically, dorsally lighter
and with scattered small dark spots. Sternum with 2 vestigial, widely interrupted bars
of darker setae at coxae II and III.
eye ratio AME:ALE:PME:PLE is 1:1.43:1.17:1.17; distances separating eyes:
Epigynum (Figs 46--47): Externally, median septum lobe produced but short and
much wider than long. Internally, spennathecae produced laterally, parallel to
epigastric groove. Spennathecae long, thin and distally reflexed anteriorly, following
profile of invaginated support collar (Fig. 47). Cedarville female has a slightly
narrower epigynal 'neck' externally (Fig. 52) and proportionately slightly shorter,
stouter spennathecae (Fig. 53).
Legs: Spine pattern typical of genus, but with femora IV(3:2:2) and tibiae dorsally
with proximal and distal spines on I-III but only a distal spine on IV. Measurements:
Palpus I II III IV
Femur 3.75 11.88 12.25 10.13 12.25
Patella 1.63 4.63 4.63 4.00 3.88
Tibia 2.50 10.38 10.63 8.38 9.25
Metatarsus 10.00 10.13 7.13 9.13
Tarsus 4.13 2.63 2.50 2.13 2.75
Total: 12.01 39.52 40.14 31.77 37.26
Description ojparatype male (NMSA TS02):

The abdomen is crumpled, and crease marks on the legs suggest the specimen was
newly ecdysed and preserved before the cuticle had hardened.
Colour and markings: Same as female except area posterior to epigastric groove
more orange-brown than a rich red-brown.
wide, 1.75 X longer than wide. Cheliceral teeth with an additional small apical tooth
on both retro- and anteromargins on left chelicera and on anteromargin of right
chelicera (Fig. 48), but absent in other specimens. Sternum: 3.31 long, 3.00 wide
between coxae II, 1.10 X longer than wide and 2.19 wide between coxae l. Labium:
1.38 wide, 0.92 long, 1.50 X wider than long, occupying 0.63 X of SW between
coxae I. Maxillae: 2.06 long, 1.17 wide, 1.76 X longer than wide.
/
J
\
\
'-.
i
("
\ \. .'
r-'/
.'
/
72
71
75 74
Figs 71-75. Palystes stuarti sp. n. 71. External female epigynum. 72. Internal epigynum (holotype
female, NMSA T499, Nieuwoudtville). 73. Ventral aspect, male pedipalp. 74. Detail of ventral
aspect, tibial apophysis. 75. Detail of retrolateral aspect of apophysis (paratype male, NMSA
T500, Nieuwoudtville).

Pedipa/p (Figs 49-51): Blade of embolus with sinuate lateral margin, column
entire (not grooved). Spur at base of embolus short and hooked. Tibial apophysis
bifurcate apically with lateral lobe. Ntolas male (Figs 54-56) has a slightly more
spathulate, obtusely-pointed embolus apically (Fig. 54), and the tibial apophysis is
more clearly bifurcate (Figs 55-56).
Legs: Spines typical of genus but with variations: left femora I+II(3:2:3),
III(2:2:3), IV(3:3:2); right femora 1+11(3:2:2), III(3:2:3), IV(3:2:2). Tibiae dorsally
with full complement of proximal and distal spines on I-IV, but some very small.
Measurements:
Palpus I n III IV
Femur 3.13 12.00 12.63 10.00 12.13
Patella 1.50 4.25 4.13 3.38 3.25
Tibia 2.13 11.75 12.13 9.25 10.13
Metatarsus 11.50 11.38 7.88 10.13
Tarsus 3.63 2.88 2.88 2.13 2.38
Total: 10.39 42.38 43.15 32.64 38.02
Additional material examined: SOUTH AFRICA: Transkei/Natal: Cedarville,

Mvenyane, 19 (AMGS 2344); Ntolas [30 0 35'S 29°05'E, 30 km W. Kokstad], Id
(AMGS 2110).
Palystes stuarti sp. n.

Figs 71-75, 76
Type material:
Holotype: adult 9 , SOUTH AFRICA: Cape: 25 km S. Nieuwoudtville, 31°22'S
19°06'E, Sewefontein Farm, 700 m, in house in Cederberg fynbos, ii.1990, C. T.
Stuart, collector's no. 3674 (NMSA T499).
Paratype: SOUTH AFRICA: Cape: Same data except: 1d (paralysed spider
collected with pompilid wasp on stoep [= verandah] of house in Cederberg fynbos),
25.iii.1990, C. T. Stuart, collector's no. 3677 (NMSA T500).
Type locality: Both holotype and paratype collected on Sewefontein Farm, 25 km S.
Nieuwoudtville (31 °22'S 19°06'E), western Cape, South Africa.
Etymology: Named after author-naturalist Mr Chris Stuart, the collector of the type
series.
Diagnosis: Spermathecae (Fig. 72) inserted laterally on the invaginations of the
support collar and reflexed inwards towards the median line as in P. crawshayi (Fig.
40), but median lobe of septum well developed (Fig. 71) and produced anteriorly and
not transversely as in P. crawshayi (Fig. 39). Spur at base of embolus (Fig. 73) far
longer than that of other species in P. lunatus group (spur in P. stuarti more than half
as long as embolus, compared with less than quarter as long).
Distribution: Only known from the type locality.
Description of holotype female (NMSA T499):
Colour and markings: From above entirely mottled black and brown. Abdomen
dorsally with an outlined cardiac marking medially, flanked by irregular large paired
paler patches. Abdomen ventrally with dark brown to black crescentic marking
immediately posterior to epigastric groove, followed by round white spots against a rich
reddish brown field. White spots continue on ventral surface of femora, which are
otherwise dark below but lighter above with several dark spots. Sternum with 2 medially
interrupted black transverse bars in line with anterior faces of coxae II and III. Labium
and maxillae, dark reddish black. Metatarsi and tarsi dark dorsally and ventrally.
Carapace dorsally with a paler medial line running from between PME to thoracic fovea.
wide across PER; thoracic fovea 3.50 long, occupying 0.23 X of CL and 0.29 X of CWo
Eyes: Eye diameters: AME 0.62, ALE 1.00, PME 0.77, PLE 0.69, PER 4.38 wide, wider
than AER by 1.21 X and occupying 0.64 X of HW and 0.36 X of CW; eye ratio
AME-AME 0.46, AME-PME 1.05, ALE-PLE 0.89, PME-PLE 0.75, PNIE-PME 0.66;
wide, narrowed anteriorly, with MOQP 1.33 X wider than MOQA. Clypeus 0.46 high,
between coxae 1. Labium: 2.62 wide, 1.85 long, 1.42 X wider than long, occupying 0.72
Cheliceral margin teeth typical of genus (3 retromarginal and 3 anteromarginal teeth),
but left chelicera has a small fourth, additional anteromarginal tooth apically.
Epigynum (Figs 71-72): Externally with well-developed, produced, median
septum lobe (Fig. 71). Internally, spermathecae (Fig. 72) inserted laterally on
invaginations of support collar, and reflexed inwards.
Legs: Spine pattern typical of genus, but femora IV on both sides 3:2:2 and right
femur II 1:1:3. Both proximal and distal tibial dorsal spines present on tibiae I-IV on
both sides, except for left tibia IV in which proximal spine absent (although present
on right leg IV, spine smaller and thinner than on other legs). Measurements:
Palpus I II III IV
Femur 5.00 15.63 16.25 14.00 16.50
Patella 2.50 6.88 6.88 5.63 5.63
Tibia 3.38 15.25 15.63 12.25 13.38
Metatarsus 15.00 14.38 10.63 13.25
Tarsus 5.75 3.50 3.13 2.63 3.13
Total: 16.63 56.26 56.27 45.14 51.89
Description of para type male (NMSA T500):

Colour and markings: As in female, except: abdomen ventrally darker.
wide across PER; thoracic fovea 3.25 long, occupying 0.26 X of CL and 0.34 X of CWo
than AER by 1.20 X and occupying 0.69 X of HW and 0.39 X of CW; eye ratio
AME-AME 0.43, AME-PME .1.05, ALE-PLE 0.75, PME-PLE 0.69, PME-PME
0.54; MOQ dimensions: MOQA 1.26, MOQP 1.78, MOQL 1.85, MOQ 0.97 X longer
than wide; narrowed anteriorly, with MOQP 1.41 X wider than MOQA; clypeus 0.43
high, 0.88 X diameter of AME. Chelicerae: 4.06 long, 2.25 wide, 1.81 X longer than
wide. Sternum: 5.00 long, 3.81 wide between coxae II, 1.31 X longer than wide and 3.25
wide between coxae 1. Labium: 1.97 wide, 1.14 long, 1.73 X wider than long, occupying
0.61 X of SW between coxae 1. Maxillae: 2.92 long, 1.91 wide, 1.53 X longer than wide.

Pedipalp (Figs 73-75): Elongate spur at base of embolus (arrow, Fig. 73) more
than half as long as embolus. Tibial apophysis (Figs 74, 75) with prominent median
spur laterally.
Legs: Spines typical of genus but with femora IV (3:2:2) on both sides. Tibiae
dorsally with both proximal and distal spines present on legs I-III, but only a distal
spine on legs IV. Measurements:
Palpus I II III IV
Femur 5.00 15.38 16.88 14.38 16.25
Patella 1.88 5.63 6.88 5.00 5.00
Tibia 3.13 15.63 15.88 12.63 13.38
Metatarsus 15.00 15.63 11.50 13.50
Tarsus 4.63 3.50 3.13 2.75 3.13
Total: 14.64 55.14 58.40 46.26 51.26
N
o ,.
?D
•
0.•
Q••
GR •
• "~G
? D P. lunatus
o P. stuarti
f}. P. crawshayi
• P. peromatus
• P. karooensis
~ P. leppanae
Fig 76. Map showing distribution of the Palystes lunatus species group. [CT = Cape Town, BW = Beaufort
West, D = Durban, EL = East London, G = Grahamstown, GR = Graaff-Reinett, N =
Nieuwoudtville, PE =Port Elizabeth, Q =Queenstown.]
Palystes superciliosus species group

Characterised by: Abdomen with dark transverse bar posterior to epigastric groove,
with or without large to small brown spots ventrally and lateroventrally, and
chocolate-edged cream white lateral markings; femora I-II ventrally dark brown to
black with a broad yellow band apically; epigynum exteriorly with septum reduced to
a 'bow-tie' -like bridge; embolus of male pedipalpal bulb narrow, with or without
vertical lamellae apically, and hooked.
Included species: Palystes superciliosus L. Koch, 1875; P. ellioti Pocock, 1896; P.
hoehneli Simon, 1890; P. johnstoni Pocock, 1896; P. ansiedippenaarae sp. n.; P.
leroyorum sp. n.
Distribution: Central, eastern and southern Africa (Fig. 109).
Palystes superciliosus L. Koch, 1875

Figs 77-80, 109
Palystes superciliosus L. Koch, 1875: 706-708, PI LXI, Figs I, lao Syntype: adult d, 'SUdafrika'
Godeffroy Museum (ZMUH), here designated lectotype [examined].
Heteropoda natalia Karsch, 1878: 772-773. Holotype 9 : 'Port Natal' (ZMHB) [examined]. Syn. n.
Palystes natalius; Pocock, 1896: 60; 1898: 59--60.
Palystes pulchripes Pocock, 1896: 60--61, Figs 4, 4a. Syntypes: 3 subadult 9 , 'Port Elizabeth' (BMNH)
[examined]. [Synonymised by Strand 1907b: 675.]
Palystes spenceri Pocock, 1896: 58-59, Figs 3, 3a. Syntypes: 9 d, 'Durban' (BMNH) [examined].
[Synonymised with P. natalius (Karsch) by Pocock 1898: 222.] Syn. n.
Palystes modiftcus Strand, 1906a: 39-41. Syntype: adult d, 'Kap der guten Hoffnung' (MWNH)
[examined]. Syn. n.
Palystes superciliosus var. fasciiventris Strand, 1907b: 676-677. Syntype: 9, 'Kapland' (MNHN)
[examined]. Syn. n.
Remarks: Koch (1875) described an adult male and an adult female from 'Siidafrika',
which he stated (p. 708) were in the Godeffroy Museum collection. The male is now
in ZMUH but the female has not been found. The adult male in ZMUH is well
preserved and intact except for a missing left leg IV and tarsus of right leg II. The
right pedipalp has been detached and placed in a microvial in the tube containing the
specimen. The syntypes were originally listed among L. Koch and Keyserling types
missing from the Godeffroy Museum collection by Rack (1961: 3), but the male
syntype was later found and listed as such by Rack (1971: 109, 133). There is no
doubt that this is the male described by L. Koch and I have accordingly designated it
as lectotype. The female syntype described and illustrated by Koch is clearly
conspecific.
Karsch (1878) notes that the spiders he described were donated to ZMHB by Dr
Rabl Riickhard, who had received them from Mr Aurel Schultz of Port Natal [=
Durban]. The adult female (numbered 2882) in ZMHB, although in poor condition,
agrees in all respects with the female of Palystes superciliosus L. Koch, and matches
Koch's (1875: 707, 708) description and illustrations of the whole spider and of its
unmistakable epigynum (PI. LXI, Figs 1, la), and it is undoubtedly the specimen
described by Karsch. Karsch refers only to an adult female in his description and I
have accordingly determined it as the holotype of Heteropoda natalia. ZMHB has a
second, similarly labelled specimen, a subadult female, which also belongs to P.
superciliosus, but it is here treated as not part of Karsch's type series.
Pocock suggested (1896) and confirmed (1898) the transfer of Heteropoda natalia
Karsch to Palystes but in the first (1896) paper also described specimens of Palystes
superciliosus under two different names: three subadult females as P. pulchripes
....... ~ .. ····t
..... ".,
(~ l.. "\
.... :;-..-.
......' )
~ . '.? '-.
....i \,
\....
78
80
84
79
~~~
83
85 86 87
Figs 77-88. Palystes superciliosus L. Koch, l875 and P. ellioti Pocock, 1896. 77-80. P. superciliosus.
77. External female epigynum. 78. Internal epigynum (lectotype female, Heteropoda natalia
Karsch, ZMHB, POr! Natal). 79. Ventral aspect, male pedipalp. 80. Detail of embolus apex
(lectotype male, P. superciliosus L. Koch, ZMUH, South Africa). 81-88. P. ellioti. 81.
External female epigynum. 82. Internal epigynum (lectotype female, BMNH, Uganda). 83.
Ventral aspect, male pedipalp. 84. Detail of embolus (holotype male, P. affinis de Lesser!,
Usambara, Tanzania). 85-88. Ventral markings. 85. Sternum (lectotype male). 86. Sternum,
paralectotype male. 87. Sternum, additional male (MRAC 831, W. Lake Kivu, eastern ZaIre).
88. Abdomen (paralectotype male).
(although tentatively suggesting that they were close to L. Koch's original

description and illustrations of P. superciliosus) and an unspecified ('large') number
of adult males and females as P. spenceri (suggesting that they were possibly the
same species as Heteropoda natalia Karsch). Rudimentary, externally visible, folding
of the epigynum is often seen in sub adult females of P. superciliosus and is evident in
the syntypes of P. pulchripes and Pocock's illustration (1896: PI. VIII, Fig. 3).
Pocock (1898: 222) correctly synonymised P. spenceri Pocock with P. natalius.
Strand (1907b: 674) correctly confirmed that the specimens Pocock had described as
P. pulchripes were subadult ('nicht ganz reife') females of P. superciliosus. The
syntypes of P. spenceri are also con specific with P. superciliosus.
Strand (1906a: 37-41) described one large and one small adult male from 'Kap
der guten Hoffnung' in MWNH as P. modificus, noting (p. 37) that they were
possibly the males of one of the already described species for which the male was
not known. The large adult male P. superciliosus identified as a type in the
collection of MWNH (labelled No. 391), agrees with the description of the larger
spider described in detail by Strand, and, also on account of the label data is clearly
one of the syntypes described by Strand as P. modificus. It is unmistakably the male
of P. superciliosus, and I have therefore synonymised P. modificus Strand with P.
superciliosus.
Strand (1907b: 676) described P. superciliosus var. Jasciiventris from
Grahamstown, distinguishing it on the more vivid ventral markings and a more
clearly defined epigynum. He saw 'viele Exemplare' from Grahamstown donated by
Brady. The syntype series, which included adult males and females, was deposited in
the museum at Lubeck (collections destroyed by air bombardment in 1942). An adult
female Palystes superciliosus from jar no. 1742 in MNHN and labelled 'Collection
E. Strand Societe zoologique de France Legs 1955'; 'Palystes superciliosus L.K. var.
fasciiventris Strand, 1 <I Capland' appears to be the only remnant of the syntype
series. The variation in ventral markings and clearer definition of the epigynum noted
by Strand as differentiating characters, are evident in the syntype, but are well within
the range of variation for P. superciliosus. The variations are not consistent in any
one locality, nor are they linked to any variation in internal epigynal structure. I
therefore regard Strand's described variation as an unjustified sub specific taxon and
have synonymised it with P. superciliosus.
Diagnosis: Epigynum (Fig. 77) without posterior extension found in other species of
this group (Figs 81, 89, 97, 104). Embolus distally (Fig. 80) unornamented with
lateral flanges, tapering to a fine, attenuate, recurved point. All other males in the
group have an embolus with lateral carinations distally (Figs 93, 100, 108), except P.
ellioti (Fig. 84) which has an embolus similar in shape to that of P. superciliosus
except that it is distally wider, flattened and bluntly pointed, and P. leroyorum (Fig.
101) in which is distally blade-like. Ventral abdomen with 4 longitudinal striations
between spinnerets and dark transverse bar posterior to epigastric groove, but
otherwise unmarked (some specimens with a faint greyish bell-shaped marking
ventrally). Colouring in both sexes variable, ranging from a relatively uniform pale
brown to more vividly coloured forms.
Distribution: Widely distributed throughout the eastern half of southern Africa, south
of the Limpopo, including Mozambique, Swaziland, the Transvaal, Natal, Transkei,

and eastern and south-western Cape provinces of South Africa.
Redescription of lectotype male of Palystes superciliosus:
The right anterior portion of the carapace posterior to the PER was damaged while
the specimen was still alive. The specimen, although faded, is otherwise in good
condition, but is missing left leg IV.
Colour and markings: Males and females have a range of markings varying from the
nearly uniformly pale brown abdomen of the lectotype male to specimens of both sexes
with an undulating logitudinal dark chocolate line laterally, subtended by a broader
white to creamy yellow band. Abdomen ventrally without markings except for 4
longitudinal striations between spinnerets and dark transverse bar immediately posterior
to epigastric groove. Sternum with single dark transverse bar between coxae II. A barely
visible, faint, partly formed second bar between coxae Ill. Anterolateral face of coxae
dark, femora with some darker mottling ventrally. Tibia banded as usual ventrally.
wide across PER; thoracic fovea 3.88 long, occupying 0.30 X of CL and 0.38 X of CWO
than AER by 1.22 X and occupying 69 X of HW and 0.37 X of CW; eye ratio
wide; narrowed anteriorly, with MOQP 1.14 X wider than MOQA; clypeus 0.62 high,
Pedipalp (Figs 79-80): Embolus robust basally, slender distally, recurved though
180, without any lateral carinations or ridges distally. Conductor membranous, large,
recurved over tip of embolus (Fig. 80).
Legs: Spines typical of genus with femora I-III(3:2:3), but left le~ IV missing and
spination of right femur IV missing a spine (3:2:1); otherwise as for genus.
Measurements:
Palpus I II III IV
Femur 4.63 15.00 15.38 12.13 14.00
Patella 2.13 6.25 6.25 5.00 5.00
Tibia 2.63 14.50 14.38 10.63 11.88
Metatarsus 14.00 13.50 9.00 11.50
Tarsus 4.50 3.13 3.13 2.38 2.75
Total: 13.89 52.88 52.64 39.14 45.13
Redescription of holotype female of Heteropoda natalia:

Colour and markings: Abdomen dorsally with pulmonary marking, laterally with
paler longitudinal band on either side, edged dorsally with an undulating darker line.
Abdomen ventrally unmarked between spinnerets and dark transverse band posterior
to epigastric groove, except for 4 longitudinal striations. Area between striations
slightly darker. Sternum with single, dark transverse bar continued on anterolateral
aspect of coxae II. Ventrally femora dark, tibiae I-IV with alternate 2 dark and 2 pale
bands. Because of the poor condition of the holotype, the colouring of a live female
from Natal (Munyawaneni Bush Camp, 2so0S'S 32°02'E, Hluhluwe Game Reserve,
12.i.1995, K. R. Cradock (NMSA)), is also described here: Abdomen dorsally with
deep chocolate cardiac outline marking against a greyish fawn-brown background,
becoming mottled and darker to a deep chocolate posteriorly and dorso-Iaterally;
abdomen laterally with broad cream coloured longitudinal band; abdomen ventrally
with only a thin black transverse band immediately posterior to epigastric groove,
otherwise pale fawn-brown; carapace dorsally pale and darker brown with pale
median line from between PME to thoracic fovea; chelicerae dark to black with
yellow-tipped setae and with 2 cream to white longitudinal stripes at outer edges;
white clypeal 'moustache' present; long setae of mouthparts orange rust-red, as are
short setae of tibiae and tarsi of the pedipalps; setae of sternum orange-yellow with
uninterrupted black bar in line with anterolateral aspect of coxae II; coxae I-IV
ventrally fawn tinged with yellow, anterolaterally black; femora I-II ventrally black,
proximally with scattered yellow-tipped setae and a distal broad yellow band
decorated with scattered fine black spots; femora III-IV ventrally yellowish fawn-
brown with black spots; tibiae I-IV ventrally with usual black and yellow bands; all
legs dorsally brindled fawn with some yellow, cream and black; ventral metatarsal
and tarsal scopulae laterally bordered with a longitudinal fine orange line.
MOQ 0.S9 X longer than wide, narrowed anteriorly, with MOQP 1.24 X wider than
MOQA. Chelicerae: 5.44 long, 2.94 wide, I.S5 X longer than wide. Sternum: 5.06
coxae I. Labium: 2.1S wide, 1.69 long, 1.29 X wider than long, occupying 0.70 X of
SW between coxae I. Maxillae: 3.32 long, 2.09 wide, 1.59 X longer than wide.
Abdomen: 17.3Slong and 13.50 wide.
Epigynum (Figs 77-7S): Median septum not produced posteriorly as in other
species of the group. Interior epigynum with tubules leading to spermathecae
enfolded to inside by lobes of support collar, spermathecae rising dorsally above
support collar lobes to lie fully exposed on top of collar lobes (Fig. 7S).
Legs: Spine pattern and markings typical of genus, but femora 1-11(3:2:3), left
femur III(3:2:2), right femur III(3:2:3) right femur IV(3:2: 1) (left leg missing);
patellae (1:0:1); tibiae I-III (basal and apical spines dorsally), tibia IV(dorsal apical
spine only). Measurements (of left legs, except for III where right leg is used):
Palpus I II III IV
Femur 4.63 14.13 14.00 1l.25 13.38
Patella 2.25 6.25 6.00 4.75 4.63
Tibia 2.88 13.25 13.25 9.50 10.75
Metatarsus 13.25 12.25 7.88 10.63
Tarsus 5.0 2.88 2.88 2.25 2.75
Total: 14.76 49.76 48.38 35.63 42.14
Additional material examined: MOZAMBIQUE: Delagoa Bay, 1 9 (MNHN 9.959);

Delagoa Bay, 30"39, ?1889, J. de Coster (SAMC 2880, 2881, 2885). NAMmIA:
Ovamboland: Kungveld, Tsumkwe Pan [19°35'S 20 0 30'E], 20 km E Rooidag gate,
19, 8.iv.1970, w. D. H. [= w. D. Haacke] (TMSA). SOUTH AFRICA: 'S. Africa',
19, Dr Quain (BMNH 70.26). Cape: Addo Bush, 10", vii.1919, J. Drury (SAMC
B4662); Adelaide, Commandsfontein, 19, 1900, E. Stevenson (SAMC 7794);
Alicedale, 10"29 (AMGS 1884, 1909, 2193); Ashton [33°50'S 20 0 04'E], 19,
xi.1919, R. W. Tucker (SAMC B4800); Bedford, 19 (AMGS 2236); Bonnivale
[?Bonnievale 33°56'S 20 0 05'E, 120 km E. Cape Town], 19 , vi.1991 (SAMC C2282,
Collector's No. 192); Cape Peninsula: Devil's Peak area, 19, 7.ii.1983, Mr Kalin
(SAMC CI056), Newlands Forest, 10", 18.v.1991, N. Larsen, no. 164, (SAMC
C2081); Cape Town: 10"] 9 , 1973, P. L. G. Benoit (MRAC 144.728), Table View,
] 9 , 12.vi.1983, N. McConnell (SAMC CI058); Dunbrody, 54km N. Port Elizabeth,
ld'19, 1903, Rev O'Neil (SAMC 13052 [? & 8299]); East London: 19 , Power
(SAMC), 19, 1899, Rev O'Neil (SAMC 5296), 10" (MNHN 18.523), 20"69
4subad.0" 3juv., ix.1914, R. Ellenberger (MNHN) , 10"19 (AMGS 2793), 19
(AMGS 2785); Fort Brown, Resolution, 3d , 11.xii.l925 & 12.iii.1930, Miss Walton
(AMGS 5197, 6257); George, Pacaltsdorp: 40"49 1subad.0" Isubad. 9 5imm., 1899,
Miss Leipo1dt (SAMC 5123), 10" 3 9 (SAMC 8446); George distr., 19 1subad. 9
(SAMC); Grahamstown: 29 , vi.1895, w. Frick (BMNH 99.7.24.64-65), 29 10"
lsubad.o", [24.vii.l899] (BMNH 99.7.24.66-71), ld' (det. Palystes natalius Kar. by
Pocock, 1900) (AMGS), 1920" 5juv., [5.iii.1901]' Schonland (BMNH 01.3.5.57-59),
10"19, 18.iv.1906 & 26.x.1908, Prof Schwartz (AMGS 207, 690), 19, 7.iv.1906,
Miss A. Daly (AMGS 200), 39, 7.xii.1907, 15.vii.1909, 30.xii.1909, Miss M.
Pinnock (AMGS 543, 806, 867), 29 , 6.ii.1908 & 26.iii.1909, Mr H. Babbs (AMGS
571, 769), 19 , 25.iii.1907, Master Weinick (AMGS 409), 19, 8.viii.1907, Master
McLeod (AMGS 482), 20"19 , vii.1905, vii. 1907, Miss E. M. Cherry (AMGS), 19 ,
19.iii.1906, Master J. Cherry (AMGS 174),29 , i.1967 (AMGS), 10" (AMGS 1322),
ld', lO.xi.1905, Mrs J. E. Duerden (AMGS), 10" (AMGS 1335), 19 (AMGS 1013),
10", 31.viii.1908 (AMGS 666), 10", 15.ix.1909, Miss F. Bessinger (AMGS 835), 19,
(AMGS 1348), 10", 2.iv.1906 (AMGS 194), 10", 22.x.1908, Mr A. J. W. Daly (AMGS
688), 10", 19.xi.1908, Miss A. Daly (AMGS 700), 10", 3l.iii.1981, P. M. C. Croeser
(NMSA), Brak Kloof, 4920" 1imm.9 , Mrs White (BMNH), 1subad.9 , 10.ii.1979,
Shirleen Gilbert (NMSA), Keilands, 1d'29 (AMGS 2037), Newington, 10" (AMGS
1644), Newington, 10" (AMGS 1567), Oatlands, I 9 , 7.i.1986, B. Londt (NMSA),
Penrock, 19 (AMGS 6363), Resolution Halt, 19, xii. 1929, A. Walton (SAMC
B7648), Sidbury, 10" (AMGS 2653), Slaaikraal Farm, 19 , 24.iii.1979, P. M. C.
Croeser (NMSA), Tea Fountain, 19 10" 4juv., [13.iii.1901], Miss Leppan (BMNH
01.3.13.260-266), Trapp's Valley, ld' (AMGS 2128), 10", 24.ii.1979, P. M. C. Croeser
(NMSA); Heidelberg, Grootvadersbosch For. Res., 10", xi.1985, J. Doyen (NMSA);
Hogsback Forest Reserve, 32°36'S 26°56'E, 39, 12-16.xii.1985, B. Londt (NMSA);

Humansdorp, Co1dstream, ld, 1921, Tucker (SAMC B5-678); Jansenville: I d , Miss
Leppan (BMNH), 3d, x.1900, Mr P. M. Dancel (BMNH 01.3.5.55-56); Jonkersberg
[33°55'S 22°14'E, S. Oudtshoorn], Id 69 (TMSA); Kimberley, 2subad.d 29
(SAMC); King William's Town: 29 (BMNH 76.70), 29, 1892, H. A. Spencer
(BMNH 1892.12.4.13-14), Pirie Bush, 1d29 , [12.x.1898], A. N. Stenning (BMNH
98.10.12.20-22), Pirie Bush, 59 , [1O.x.1900], A. N. Stenning (BMNH 0.10.10.3-7),
Pirie Mission, 19, 1909, R. Godfrey (SAMC B93), urban area, 19, i.1973, P.
Swanepoel (PPRI '76/1128); Klein Winterhoekberg, Perdepoort, 3325Ac, Id,
1O.vii.1980, W. R. Branch (NMSA 13748); Knysna: 19, 21.v.1912, J. H. Rex
(TMSA 1106), 1d , 21.v.1915, 1. H. Rex (TMSA 13384 old 2143), 19 (AMGS
1218), Diepwalle Forest, 19 , xi.1985, T. & C. Griswold & J. Doyen (NMSA),
Newdigate [?loc. or coil.], 19 (S AMC) , Knysna Forest, 1d, i.1911, Mrs Rex
(NMSA 2102), Knysna Forest, 19 , I. Bainbridge (SAMC 872.197), Rheenendahl,
Goudveld Forest Reserve, 19 , 3.i.198l, P. M. C. Croeser (AMGS), The Wilderness,
ld, viii. 1963, J. S. Taylor (NMSA 8845); Mosselbay: 49, i-ii.1919, J. Power
(SAMC B4593), 29 (SAMC 3185), 29 , Coster (SAMC); Pearston: W, [5.iii.1901],
Dr Broom (BMNH 01.3.5.18), 29, 27.ix.1927, B. Marais (AMGS 5679); Port
Alfred, 1d , 29.iv.1979, R. A. Jubb (NMSA); Port Elizabeth: [1890], 29 limm., H.
A. Spencer (BMNH 90.12.15.31-33), 19, L. Drege (BMNH), 19 Id limm.9
(MNHN 6.382), ld19 , xi.1897 & 1898, J. Drege (SAMC 2149, 3729), 29 (AMGS
2936, 1534), Dunbrody, 3d1 9 , 1903, Rev. O'Neil (SAMC 13052), Dunbrody, 19
(AMGS 2951), Dunbrody, Id, 5.xii.1925, Rev. K. Tasman (AMGS 5193), Dunbrody,
Blue Cliff, 19 (AMGS 1894), Bluecliff, 19, 17.xii.1899, 1. L. Drege (SAMC
5709), Redhouse, 4d, iii.1915, Mr Paterson (SAMC B1128, B3884, B4650), Sardinia
Bay, Sardinia Downs Farm, 19, 16.ii.1981, W. R. Branch (NMSA), Zwartkops
valley, 19, v.191O, Miss D. Laffan (NMSA 2097); Riversdale distr., 34°1O'S
21°20'E, [85 km W. Mossel Bay], I d , 1919, H. Hermann (SAMC B2320);
Riversdale, 19 , 1909, H. Hermann (SAMC B2321); [?Riviersonderend], 39 2d,
ix.1933, R. F. Lawrence (SAMC 8444); Somerset East district, Zuurberg, 2d,
7.ix.1969, A. E. Deacon (AMGS); Storms River Mouth, 19 , ii.1966, A. L. Capener
(NMSA 9535); Stutterheim, Amatola mountains: Evelyn Valley Forest, 1400 m, 19 ,
25.vii.1985, P. M. C. Croeser & A. C. Duckworth (NMSA), Isidenge State Forest,
19 , 25.vii.1985, 19 , 24.vii.1985, P. M. C. Croeser & A. C. Duckworth (NMSA),
Isidenge State Forest, 1d, 18.ix.1985, Mrs T. Kroger (NMSA); Worcester, 19
(AMGS 2074). Transkei: Dwesa Coastal Forest, 3d, v.1984, R. Kilburn (NMSA);
Dweza, 19, 20.xi.199l, D. A. Barraclough (NMSA); Kentani: 19, xi.1909, Miss
Pegler (SAMCB512), Id, 1914, H. P. Abernethy (SAMC B50), 2d1 9 limm., 1904,
Pyler (SAMC 13660), 19 (AMGS 1904); Kentani district, 19 , 1899, 19 , 1902, Dr
F. Kolbe (SAMC 4544 & 12461); Lusikisiki distr., Mzirnhlava River mouth, 69 ld
Isubad.9, i.1980, 1993 d , ii.1980, M. Baddeley (MRAC 163.213, 166.705);
Mbotyi, coastal grassland, 19, v.1985, D. Herbert & R. N. Kilburn (NMSA);
Mgazana Mouth, 1d, 8.xii.1975, J. R. Grindley (NMSA); Ngqeleni, 2d1 9 (AMGS
2289, 2413); Ntabanana, 3& (AMGS 4603); Port St Johns: ld, vii.1945, R. F.
Lawrence (NMSA 4564), 2d69 2suhad.9, 1907, G. Shortridge (SAMC 9924), Id,
CROESER: REVISION OF PALYSTES (ARANEAE: HETEROPOmpAE) 63
13.iv.1958, E. S. Ross & R. E. Leech (CAS C), 19, 1902, Shortridge (SAMC
13063); Umtata, 19, vii.1961, M.Courtney-Latimer (NMSA 8093). Natal: Natal,
39 , c.m. [?c. Martin] (MNHN 18.468); Natal, 90'99 6juv., C. Martin (MNHN
19.745); 29, Abrahams (NMSA 2 lOS); 29, [22.iii.1889], E. Howlettt (BMNH
89.3.22.3-4); 19 2imm., G. F. Leigh (BMNH); Drakensberg Mountains, pine forest
SE. Champagne Castle Hotel, 10', 20.i.1991, B. & V. Roth (NMSA); Drakensberg
Mountains, Cathedral Peak Forestry Station, 1360 m, 1 9 , 29.iii.1990, I. Pajor
(NMSA); Drummond, 1 9 , 24.iv.1985, June Roos (NMSA); Durban: 70'60' 2imm.,
H. A. Spencer (BMNH), 19, [1912], E. c. Chubb (BMNH 12.3.4.15), 6930',
[l4.vii.1903], J. P. Cregoe (BMNH 03-7-14.38-40), 29 10' 9imm., 10.vii.1903 L. F.
Leigh (BMNH lO.VII.03), coastal dune scrub, ] subad.9 , 17.iv.1976, F. Wanless &
A. Russell Smith (BMNH 17.4.76), Bluff, 1 9, 'Spring 1976', Fred [?Wan1essl
(BMNH), 19, 7.viii.1972, F. L. Farquharson (NMSA), ] 9 , x.1970, M. L. Jarvel
(NMSA 9840), lsubad.9 , 2.v.1968, Mrs Demont (NMSA), 19, 18.i.1984, J. Visser
No. 2837 (NMSA), Hillary, ] 9 , 19.xii.1918, F. Ducasse (NMSA 12272), Marine
Parade, 19 , 30.1ii.1983, R. Pewtress (NMSA); Estcourt: 20'40' lsubad.o' ] sub ad. 9 ,
G. Marshall (BMNH), 10', [5.iii.1907], Mrs E. J. Turner (BMNH 07.3.5.3-14[part]),
G. Marshall (BMNH 96.4.20.16-19), 20'10' Isubad.9, G. Marshall (BMNH
96.4.20.16-19); Hilton: 10', iv.1986, 1. & R. Enders (NMSA), World's View, 19,
24.ii.1986, lLondt (NMSA); Howick, 29°28'S 30 0 14'E), 20' Ijuv., 20--30.1ii.1983,
Mrs l G. Joubert (NMSA); Kokstad, 29 (AMGS 2618, 2382); 39 mi N. Kranskop,
Ekombe For., 1520 m, 10', 10.iv.1958, E. S. Ross & R. E. Leech (CASC);
KrantzkIoof, 10', 7.v.[1913], Bell-Marley (SAMC B1306); Lidgetton, 29°26'S
30 0 06'E, 40 km NE. Pietermaritzburg, 1 9 , iii.1985, G. Timms (NMSA); Lower
Umkomaas River: 10'10', [4.xi.1897], G. Marshall (BMNH 97.11.4.16-17), 50' 3juv.,
[20.vii.1903], G. F. Leigh (BMNH 03.7.20.58-65 [part]); Liineburg [27°19'S 30 0 37'E,
40 km Sw. Piet Retief], 29 (AMGS 1694 & 2019); Marina Beach [30 0 56'S
30 0 18'E], 19, xii. 1985, R. Douglas (BMSA NMBA1362); Merrivale,
TshwaiaBenyoni farm (25 km NNW. Pietermaritzburg), 19, 13.v.1983, 10',
2.v.1984, B. R. Stuckenberg (NMSA); 10 Ian SE. Muden, Mhlopeni Nature Reserve,
29°02'S 30 0 21'E, 19 , 8.iv.1983, P. M. C. Croeser, B. Londt & W. Cadman (NMSA);
nr Pennington, 30 0 15'S 30 0 45'E, 19 , 11.1i.1979, J. Hardman (NMBZ AI51O); 70 km
NE. Pietermaritzburg, side of road between Grey town and Kranskop, 1%,
12.iii.1982, R. Pewtress (NMSA); Pietermaritzburg: 19 & spiderlings, iv.1913,
Burnup (NMSA 19]4), 19, iii.1914, Burnup (NMSA 1686), 10', viii.1913,
McKellar (NMSA 1938), 10', 1923 (NMSA 948), 19 , 2l.ix.1967, Gay Bendemann
(NMSA 9591), 10', 22.x.1969, P. Lamora] (NMSA 12438), 29, l.x.1979, B.
Lamoral (NMSA 12803), 10', 20.iv.1979 (NMSA 12202), 19 (NMSA 2098), ]0',
15.v.1982, A. J. Wilby (NMSA), 10', 20.iv.1983, B. Londt (NMSA), 10', 12.viii.1984,
B. H. Lamoral (NMSA), ] 9 , xii.1984, B. Last (NMSA), 10', v.1985, P. M. C.
Croeser (NMSA), 19, 15.ix.1985, B. van Hoogdalem (NMSA), 19 , ix.1985, P. M.
C. Croeser (NMSA), 10', 25.v.1986, B. van der Merwe (NMSA), Ashburton, 10',
20.ix.1981, Dept of Entomology, University of Natal (NMSA), 19 , 21.v.1984, Elna
Venter (NMSA), Athlone, 10', 18.iv.1985, D. Barraclough (NMSA), Clarendon, ]0',
30.v.l983, R. S. Brooking (NMSA), Grange, 10', 2.v.1985, C. R. Hurt (NMSA),
Lincoln Meade, 10, 19.i.1983, S. R. Gray (NMSA 14167), Mkondeni, 10, 7.x.1969,
P. F. Tempest (NMSA 12446), Montrose, 19 , viii. 1985 (NMSA), 10, 14.xi.l985, J.
Landt (NMSA), Pelham, 19, 6.viii.1990, R. B. Graham (NMSA), Prestbury, 10,
15.v.1983, 19 , vi.1983, 19 , 6.vi.1984, 19 , 2.viii.1985, P. M. C. Croeser (NMSA),
Prestbury, 10, 7.xii.1987, W. Starega (NMSA), Scottsville, 19, 18.i.1970, 1019,
i.1973, B. Lamoral, Isubad.0 29, xi-xii. 1970, B. Joubert, 19, 23.iii.1983, P.
Croeser & R. Kilburn, 10, 26.vi.1984, R. Kilburn, 19, 5.x.1983, Miss K. Herbert,
10, 1O.vi.1983, R. du Toit, 19, 4.x.1986, S. Duff, 19, 20.vii.1984, J. Kilburn
(NMSA), Shortt's Retreat, 19 ,4.i.1984, D. M. Ward (NMSA), Town Bush, D.S.G.'s
grave, 19, 15.x.1983, P. Croeser & C. Griswold (NMSA), Town Hill, 19,
20.iii.1983, 10, 21.ii.1983, 19 & spiderlings, 17.i.1984, 19, 12.x.1985, 19,
vi.1987, P. F. Tempest (NMSA), Winterskloof, 10', vii.1985, B. Last, 10, 27.vi.1985,
B. Last (NMSA), 15 kIn NE. Pietermaritzburg, Lothlorien Farm, 10, 18.v.1983, R. S.
A. du Toit (NMSA), University of Natal, Faculty of Agriculture, 10', 8.x.1980, P.
Govender, 10, 6.x.1981, D. York (NMSA); Pinetown, 19, x.1896, J. P. Cregoe
(SAMC 972.197); Port Edward: 10', 26.iii.1983, 10', vii.1984, 1 9 , ix.1984, 1 9 ,
ix.1984, 10', caravan park, Umtamvuna River, xi.1984, 10, xii.1984, 30' 1subad.0
1subad.9, iv.1985, 19 , vii.1985, 19, viii.1985, 10', x.1985, 19 1juv., xi.1985,
2019, xii.1985, 10'19 , i.1986, 10', ii.1986, 10'19, v.1986, 19, vii.1986, 19,
x.1986, lO', x.1986, 19, i.1988, 19, [no date], June Stannard (NMSA); Port
Edward, Blencathra Farm, 31°02'S 30 0 1O'E, 5 kIn W. Port Edward, 335 m: 1 9 , 10',
5.iv.1983, 19, 7.iv.1983, 10 , 26.v.1983, 19, 12.vi.1983, 19, 17.iv.1983, 10,
25.iv.1983, 10, i.v.1983, 10', 19.v.1983, lO', 25.v.1983, 10', 15.vi.1983, 10', vii. 1983,
19, 9.vii.1983, 19, ix.1983, 19, x.1983, 19, x.1983, 19, with egg sac &
spiderlings in old hollow cane, 18.x.1983, 1 9 , with leaf nest, spiderlings and second
egg sac, 25.x.1983, 10', x.1983, 1 9 , xi.1983, 1 9 , olive-coloured, i.1984, 10, at
night, same room as 0' Palystes peromatus, iv.1984, 40'1 «, iv.1984, 20'1 «, vL1984,
20', vii.1984, 10'19 , 20.v.1985, 19 , vi.1985, 10', [no date], June Stannard (NMSA);
Port Edward, M. Walker's farm, 4 kIn NW. Port Edward: 10', 11.v.1983, 19
17.v.1983, 10, 5.vi.l983, 19, 5.vi.1983, 19, viii. 1983, 19, viii.1983, June
Stannard (NMSA); Umvoti River Mouth, lO', i.1986, J. Stannard (NMSA);
Richmond, 49 , 1899, Rev. Ward (SAMC 5338); Rietvlei distr., Blackheath Farm, 35
kIn from Grey town on Rd to Mooi River, 2019 , 16.ii.l983, Mrs F. Roos (NMSA
14166 & 14169); Salt Rock, 29°29'S 31 ° 14'E, Sheffield Beach, 1 9 , 26.xii.1985, A.
C. Duckworth (NMSA); Shafton House, 1019, iv.1921, R. E. Simons (SAMC
B5689); Umgababa, 30 0 08'S 30 0 50'E, 10, xi. 1939, J. Kent (NMSA 12197);
Umhlanga Rocks [29°43'S 31°05'E], 19, 13.i.l983, F. Reardon (NMSA 14180);
Weenen, 1019 (AMGS 3005); Weenen, Tugela Estates, 19 , 8.x.I925, G. v. Dam
(TMSA 6117); Tugela Estates, 2500 ft, 1 9 , 8.x.1925, R. E. Symons (TMSA 6116);
Winkelspniit [= Winklespruit, 300 06'S 30 0 51'E], 10'1 9, v.1917, C. Akerman
(NMSA 1933, 1932). Natal (Zululand): 1029 Isubad.0 2juv., J. C. Martin
(MNHN 19.537); Blythedale Beach [near Stanger], 10 2subad.0 Isubad.9 ,8.i.1980,
P. Reavell (NMSA 13149); Charters Creek, Lake St Lucia, 29°12'S 32°26'E: 10,
19-20.xi.1985, C. & T. Griswold, J. Doyen (NMSA), 1 9 , iv.1984, R. van der Eist
(NMSA); Dukuduku [28°22'S 32°13'E]' 19 2subad.9, viii.1905, Toppin (NMSA
2104); Empangeni [28°45'S 31°54'E]: lei'19 , xii.1977, P. E. Reavell (NMSA

12763), lei', 29.ii.1984, P. E. Reavell (NMSA), lei', 18.xi.1983 (SAMC C1882), lei',
16.i.1984, P. M. C. Croeser (NMSA), University of Zululand, 19 , P. E. Reavell
(NMSA 12116), 19 , 8.ii.1985, 19, 30.viii.1985, P. E. Reavell (NMSA); Nyala
Game Ranch, 22 kIn NW. Empangeni, 19, 27.vi.1983, B. Londt (NMSA);
Ingwavuma [27°08'S 32°01 'E], 3ei'39 , viii.1933, A. Roberts (TMSA 6485-6486);
Kosi Bay [26°57'S 35°20'E]: 19 ,xii.1905, Toppin (NMSA 2101), Kosi Bay, KBNR
campsite beside Lake Nhlange, 19 ,29.x.1989, D. G. Herbert (NMSA), Makowe via
Somkele, lei', [4.iii.1912], E. C. Chubb (BMNH 12.3.4.14), Mapula, lei', vii. 1936, R.
F. Lawrence (NMSA 162); Mseleni, 19, x.1905, Toppin (NMSA 1696); Sodwana
State Forest, 27°35'S 32°37'E, KwaMbila, 19, 25.vii.1984, A. c. Duckworth
(NMSA); Sodwana Bay, lei', 19.v.1981, C. A. Car (SAMC C273); Somkele, [28°18'S
32°10'E], ld19 , 1914, J. Hawkins & A. Loveridge (MCZC); Somkeie [?Somkele],
lei' (MCZC); St Lucia Estuary, 19 , 4-6.xii.1966, J. G. Rozen & D. J. Brothers
(AMNH). Orange Free State: Bloemfontein, I 9, E. Reimoser (MCZC).
Transvaal: 29 2subad.ei' 3subad.9 Ijuv.9, Sammlung Reimoser (NHMV); lei',
ix.l983, J. Stannard (NMSA); Acornhoek [24°36'S 31°04'E], 29 , xii.1918 [?1919],
R. w. E. Tucker (SAMC B4517 & B4522); Badplaats Post Office, via Carolina, 1 9 ,
3.xii.1932, S. Grobler (TMSA 5305); Barberton: 25°47'S 31°02'E, lei', x.I922, G.
van Dam (TMSA 13363 old 864), lei', vii.191O, 1 subad.9 Ijuv., v.191O, Isubad.9 ,
vii.191O, Miss de Beer (TMSA 13734 old 1146, 13735 old 1276, 13754 old 2446,
13731 old 1143), lei', 25.v.I923, L. C. de Beer (TMSA 14761 old 1914), Barberton,
lei', 1898, Dr Kolbe (SAMC 3738), Princetown Hill, above Agnes Mine, 19,
8.v.1943, D. H. Bowden (TMSA 8649), Nelshoogte, Eastern Transvaal Timber Co.
Plantation, 2ei'19 , 12.vi.1979, M. Stiller (PPRI '80/182, '80/192, '80/177); [Brits],
De Kroon, Crocodile River, 19 , 26.xii.191O, 29 , 25.iv.1911, G. P. F. van Dam
(TMSA 1103-1105); Carolina distr, Nelsberg, 19 , 25.v.1914, A. Roberts (TMSA
1127); Cullinan, Premier Mine, 19, 28.viii.1931, D. Uys (TMSA 5176);
Haenertsburg, 23°55'S 29°57'E, Merrick farm, 19 , (colI. 1O.xii, egg case 14.xii,
eggs hatched 29.xii), 1O.xii.1977, P. M. C. Croeser (NMSA); Haenertsburg,
Clearwaters Farm, 193ei', 4.ii.1916, Mrs C. Thompson (TMSA 1113-1116);
Johannesburg: 2ei'29 , vi-vii. 1987, 2ei'I9, vii.I987, 19, ii.I988, M. Filmer
(NMSA), 19, 13.v.1965, Zoology Dept, University of the Witwatersrand, No
X.33.(AMGS), Bryanston, 1& with egg sac, vii. 1987, M. Filmer (NMSA), Parktown
North, ld, x.1987, Jacob, 19, xii.1987, M. Filmer (NMSA), Randburg, 19 lei',
xii.1985, A. LeRoy (NMSA), Sandton, lei', xi.1983, P. Croeser (NMSA),
[Johannesburg] Zoo grounds, lei', vi.1984 [?preserved 31.xii.1986], J. Cowan (PPRI
'87/44); Kaapmuiden, Althorpe farm, 10', 15.iv.1979, M. Stiller (PPRI '80/206);
Kruger National Park, Skukuza, 19, l6.ix.1984, L. Braack (NMSA); near Lebombo
Mountains, Sabie & Sand rivers, Iei'I 9 , 1898, H. Fry (SAMC 4220, 4219); Louis
Trichardt, lei', 28.ii.1923, Dr H. C. Breyer (TMSA 1005); 35 mls E. Louis Trichardt,
Tshakoma, 19, xi.193I, G. van Son (TMSA 5187); Lydenburg, 19, Flygare
(TMSA 13774. old 2417); Lydenburg: 19, 1895 (TMSA 13777 old lL58), lei',
l.vi.1911, H. J. Gurr (TMSA 1107); Nelspruit, Rietspruit, lei', 5.vi.1916, A. Roberts
(TMSA 13369 old 2197); Nylstroom, Waterberg dist.: 19, xii.1915, G. P. F. van
Dam (TMSA 1160), 19 , xii.1923, v. Dam (TMSA13069 old 2827); Onderstepoort

[near Pretoria], 10', 14.x.1986, N. Smit (PPRI '86/415); Pienaars River, on Great
North Road, 19, vi.1971, G. de Beer (TMSA 10128); Pietersburg: 19, 20.xi.1916,
L. Beacom (TMSA 12926 old 1801), 19 , [5.v.1889], C. R. Jones (BMNH 89.5.5.4),
10', xi.1984 (NMSA); Pietersburg district, 10', Miss Czerikow (TMSA 1128); 20 mls
E. Pietersburg, 29 , 1899, Rev Daniel (SAMC 5340); Pietersburg, Smitsdrift, 19
Isubad.o', Miss Moller (AMGS); Piet Retief, 19 lsubad.9, xii.1976, L.
Klingenberg (PPRI '77/791); Potgietersrus: 19, 23.i.1920, H. B. Pretorius (TMSA
1778), 10', x.1919, Dr Melle (SAMC B4731), 10' Isubad.9 , iv.1934, R. F. Lawrence
(SAMC B8468); Pretoria: 19, 13.i.l987 (PPRI '87142), 19, 20.iii.1897, A. Smith
(TMSA 1122), 19, 23.vi.1906 (TMSA 1080), 10', 8.xi.1906 (TMSA 1081), 19,
viii.1908, G. P. F. van Dam (TMSA 1119), 19 , vii.1908, G. P. F. van Dam (TMSA
1120), 10', viii.1908, G. P. F. van Dam (TMSA 1121), 19 , vii.1909 (TMSA 1112),
29 10', x.1909, R. M. Lurcock, (TMSA 1074-76), 19, 9.iii.1911, Rev N. Roberts
(TMSA 1099), 10', 3.vii.1911, Mr Rose (TMSA 1101), 19, 19.viii.1911, Dr
Gunning (TMSA 1095), 19, 9.iii.1911, Rev N. Roberts (TMSA 1100),19, v.1912,
G. P. F. van Dam (TMSA 1098), 19 , 13.ix.1912, N. v. Tinteren (TMSA 1102), 10',
8.v.1912, I. P. Jenkins (TMSA 1125), 19, 9.v.1912, H. Brownfield (TMSA 1124),
lsubad.o', Il.xi.1913, Miss Barratt (TMSA 13130 old 2315),10' (TMSA 4995),19
(TMSA 4998), 10' (TMSA 4999), 19, 15.xi.1921, Mr Jorrison (TMSA 13132 old
1961),19 (TMSA 5209),19, 1l.ix. 1928, A. G. White (TMSA 6152),10', iv.1936,
N. von Wulff (TMSA 7465), 19, v.1941 (TMSA 8300), 19, x.1943, B. Dick
(TMSA 8653), 10', xii.1947, A. Obermeyer (TMSA 8687), 10', ix.1956, 10', x.1956,
10', 21.iii.1960, T. A. P. de Beer (TMSA), 10' lirnm., l.ix.1965 [?x.1949] (AMGS),
19, 4.xii.1969, G. Newlands (TMSA 9493), 19, 11.i.1972, F. Barrington (TMSA
10318), Isubad.9 , 14.v.1916 [?1976], D. Gunn (PPRI '76/1046), 10', 14.x.1978, M.
Grobler (PPRI '86/410), 10', 18.x.1978, Mr Fourie (PPRI '791273), 19, 16.viii.1979,
R. Kfir (PPRI '801200), 19 , 18.xi.1979, H. van Ark (PPRI '81/611), 19 ,21.xi.1979,
M. A. van den Berg (PPRI '81/618), 19, x.1980, H. van Ark (PPRI '81/581),
Isubad.9, iv.1981, G. van der Linde (PPRI '82/338), 10', 20.iii.1983, Mrs
Papadopolis (PPRI '83/195), 19, 3.v.1984, C. Eardley (PPRI '84/454), 19 , i.1985,
A. Ie Roy (NMSA), 10', 5.x.1985, E. J. Wright (NMSA), 10', 16.ix.1986, L. Vrey
(PPRI '86/409), 19, 17.ix.1987, E. S. Prentice (PPRI '87/840), 10', l.x.1987, L.
Oosthuizen (PPRI '87/839), Brooklyn, 10'19, 28.ix.1977, Mrs Harmse, (PPRI
'78/123), Faerie Glen, 10, 11.viii.1979, 1. Grobler (PPRI '801209), Faerie Glen, 20,
x.1984, Boeta Fourie (NMSA), Fountain Grove, 19, 19.iii.1901 (TMSA 1077);
Pretoria, Gezina: 19, Rev Roberts (TMSA 13143 old 2270), 19, Rev Roberts
(TMSA 13145 old 2272), 10', Rev Roberts (TMSA 13146 old 2273), 19,
13.iv.1915, Dr van Hoepen (TMSA 1202), 19, 5.vi.1930, C. 1. Fourie (TMSA
4996); Pretoria, Groenkloof, 29, xi.1973, E. de Ridder (TMSA 10815), Hazelkwist,
50'29 Ijuv.9 Isubad.9, xii.1949 (NMSA), Landbougebou, 10', 24.ix.1985, S.
Neser (PPRI '86/501), 10', 11.ix.1985, D. du Preez (PPRI '86/498), Mayville, 10',
9.x.1934, Wolswinkel (TMSA 7358), 10', 3.iii.1976, L. Harley (PPRI '76/957),
Pretoria North, 19, iii.1942 (TMSA 8614), Roodeplaat Dam, 10', 15.x.1985, P.
Giller (NMSA), Union Buildings, 10', 20.ix.1976, S. Stiemie (PPRI '76/1522),
Villieria, }d, 8.xi.1976, M. Zwiegelaar (PPRI'76/1931), 19 , xi.1977, Mrs J. F. Smit

(PPRl '77/1067), 10", 2.v.1979, Mrs Smit (PPRl '80/195), Wonderboom-Suid, 10",
1974, A. S. D. [A. S. Dippenaar-Schoeman] (PPRl '771753); Rustenburg,
Zandfontein, 10", 25.vi.1914, G. P. F. van Dam (TMSA 1123); Standerton, 19
(TMSA 7402); Sulphur Springs [probably 20 km SSE. Piet Retief], 19 , to.iv.1907,
S. C. Heymans (TMSA t073); Tshipise, 19 , 5.v.1986, S. Kamburov (PPRI '86/500);
Verwoerdburg, 18 km SE. Pretoria, Eldoraigne, 19, 27.vi.l973, J. Church (TMSA);
Waterberg, Vyeboompoort, 10", 13.x.1913, C. Truter (TMSA 13353 old 2316);
Waterpoort, 1 9 , . 20.x.1932, Maj Bronkhorst (TMSA 5279); Zebediela, 10",
14.vii.l958, 19 , 1O.x.1958, R. B. Copley (TMSA); Zoutpansberg, 10", [14.vii.1903],
J. P. Cregoe (BMNH 03.7.14.103); Zoutpansberg district, Shilowane, 19 , i.191O,
Rev Junod Maphophe (TMSA 1094). SWAZILAND: [?], Halibura, 19 , xii.1918,
Watermeyer (SAMC B4231); Henwood's Halt, 3d , v.1939, C. Major (SAMC 9842);
Mhlume, NE Swaziland, Mbuluzi Game Reserve, 10", 10.v.1986, A. LeRoy (NMSA);
Piggs Peak, 10" 1imm. 9 , 17. vi.1916, A. Roberts (TMSA 13418 old 2195).
Palystes ellioti Pocock, 1896

Figs 81-88, 109
Palystes ellioti Pocock, 1896: 58, PI. VIII, Figs 2, 2a. Syntypes: 3 adult 9 [adult 9 with pin, here
designated as lectotype, remaining specimens designated paralectotypesj, 'Uganda', G. F.
Scott Elliot (BMNH). [examined].
Palystes amallicus Strand, 1907a: 541 [amplified in Strand 1907b: 680]. Holotype: subadult d, 'Amani'
[Usambara Mtns, Tanzania] (ZMHB - Strand - not confirmed) [not examineq] Syn. n.
Palystes affillis Lessert, 1921: 396, Text Fig. 16, p. 395. Holotype: adult d, 'Usambara' [Tanzania] (body
in NHRS) [examined]; (right pedipalp and bulb of left pedipalp in MHNG) [examined] Syn. n.
Remarks: Pocock (1896: 58) described the adult female of P ellioti based on material
collected by G. F. Scott Elliot. It is most likely that it was during Scott Elliot's
Ruwenzori expedition that the syntypes of P ellioti were collected. The majority of
specimens I have determined as P ellioti come from north and north-western
Tanzania, south-western Uganda, Rwanda, Burundi and eastern ZaIre. When
describing P ellioti, Pocock did not specify how many specimens he saw, but in the
absence of any other information the three females identified as types in BMNH
obviously constitute a syntype series. I have designated the pinned female as
lectotype and the other two females as paralectotypes.
Lessert (1921: 396) described P affinis from a single adult male collected from the
Usambara Mountains (eastern Tanzania) by Dr Y. Sj6stedt's 1905-1906 Swedish
expedition. The adult male in NHRS, together with the right pedipalp and left
pedipalp bulb in MHNG (NHRS records note that the missing pedipalp and bulb are
in MHNG), agree in all respects, including label details, with Lessert's description,
and I have accordingly determined the specimen as the holotype of P affinis. I cannot
differentiate the male holotype of P affinis from males I have determined as P ellioti
from south-western Uganda, Rwanda-Burundi and eastern Zaire, and I was not able
to find any females in the various collections from the Usambara Mountains. On the
basis of the specimens examined, I have recognised P affinis as a junior synonym of
P ellioti.
Strand's (1907a, 1907b) description of P amanicus is based on a subadult male
collected by Vosseler from Amani (Usambara Mountains), but agrees in all major
respects with the description of P. affinis (holotype also from Usambara) and males
associated with P. ellioti. Strand's description does not provide any infonnation which
would justify separating P. amanicus from P. ellioti. I have not been able to establish
whether Strand's holotype is still in ZMHB where he recorded that it was originally
deposited. As I have synonymised P. affinis with P. ellioti, and in the absence of any
evidence to the contrary, I have also synonymised P. amanicus with P. ellioti.
Diagnosis: Abdomen ventrally with large dark brown blotches coalescing posterior
to epigastric groove. Epigynum internally (Fig. 82) with spenn duct dorsal to collar
and then forming a sclerotised structure on which spennathecae lie ventrally at
posterior inner angle of support collar. Collar does not enfold spennathecae (as in P.
ansiedippenaarae and P. hoehneli), nor do spennathecae lie dorsally on support
collar (as in P. superciliosus). The embolus (Fig. 84) lacks dorsal and ventral flanges
found in P. johnstoni, P. hoehneli and P. ansiedippenaarae, but is thicker and apically
obtuse and recurved through less than 1800 compared with P. superciliosus.
Distribution: Distributed from eastern ZaIre and Rwanda in the west, to south-
western Uganda in the north, and across Tanzania to northern Malawi. Although no
specimens from Burundi were seen, it doubtless occurs there as well.
Redescription of lectotype female of Palystes ellioti:

Specimen has a pin longitudinally through the cephalothorax and abdomen, and
abdominal cuticle is ruptured dorsally.
Total length 27.51 mm.
Colour and markings: Abdomen with dorsal pulmonary marking, lateral
undulating cream band with dorsal dark brown edge, ventral large dark brown
blotches coalescing posterior to epigastric groove. Sternum with 3 transverse dark
bands coalescing centrally, coxae with dark anterior margin and legs banded as in all
other congeners.
wide across PER; thoracic fovea straight-longitudinal, 2.25 long, occupying 0.19 X
of CL and 0.25 X of cwo Eyes: Eye diameters: AME 0.52, ALE 0.66, PME 0.49,
PLE 0.49; PER 3.56 wide, wider than AER by 1.21 X and occupying 0.64 X of HW
and 0.40 X of CW; eye ratio AME:ALE:PME:PLE is 1:1.26:0.94:0.94; distances
separating eyes: AME-ALE 0.18, AME-AME 0.43, AME-PME 0.60, ALE-PLE
0.52, PME-PLE 0.65, PME-PME 0.65; MOQ dimensions: MOQA 1.38, MOQP
1.66, MOQL 1.42; MOQ 1.17 X wider than long, narrowed anteriorly, with MOQP
1.20 X wider than MOQA. Chelicerae: 5.06 long, 2.56 wide, 1.98 X longer than
wide. Sternum: 4.38 long, 3.75 wide between coxae II, 1.17 X longer than wide and
3.44 X wide between coxae I. Labium: 1.94 wide, 1.29 long, 1.5 X wider than long,
occupying 0.56 X of SW between coxae I. Maxillae: 2.98 long, 2.06 wide, 1.45 X
longer than wide.
Epigynum: Internally with spenn duct dorsal to collar and then forming a
sclerotised structure on which spennathecae lie ventrally at posterior inner angle of
support collar.
Legs: Spination typical of genus. Tarsi of legs I and II missing on both sides.
Measurements:
Palpus I II III IV
Femur 4.00 11.88 12.13 9.38 11.50
Patella 2.50 5.25 5.25 4.38 4.13
Tibia 3.00 11.63 11.63 9.00 9.63
Metatarsus 10.63 10.63 7.25 9.25
Tarsus 4.75 2.13 2.38
Total: 14.25 32.14 36.89
Redescription of holotype male of Palystes affinis:

Colour and markings: As for female, but lateral abdominal marking not as distinct
and without pale longitudinal band; ventral blotches smaller. Dark bars of sternum
(Fig. 85) coalescing to cover much of sternum. In other males of P. ellioti, sternum
markings (Figs 86, 87) range from that described above to 3 dark bands coalescing
mesally, lateral abdominal markings range from form described here to those with
lateral pale bands being more yellow than white. Ventral abdominal markings (Fig.
88) are also variable, ranging from spots to dark brown blotches posterior to dark
post-epigynal bar.
Carapace: Dorsally 9.38 long, 7.00 wide between leg bases II. Head region 4.44 wide
MOQ dimensions: MOQA 1.23, MOQP 1.38, MOQL 1.38, MOQ as long as wide;
narrowed anteriorly, with MOQP 1.12 X wider than MOQA; clypeus 0.38 high, 0.83 X
diameter of AME. Chelicerae: 4.00 long, 2.06 wide, 1.94 X longer than wide. Sternum:
3.75 long, 3.31 wide between coxae II, 1.13 X longer than wide and 2.50 wide between
coxae 1. Labium: 1.63 wide, 1.14 long, 1.43 X wider than long, occupying 0.65 X of SW
between coxae I. Maxillae: 2.68 long, 1.63 wide, 1.64 X longer than wide.
Pedipalp: Embolus (Fig. 84) similar in shape to that of P. superciliosus, both
species lacking apical, lateral lamellate carinations found in other species of this
group. Distally, embolus wide, flattened, and terminating in a blunt point, less
reflexed than embolus of P. superciliosus, in which it narrows distally and attenuates
to needle thinness.
Legs: Spines typical of genus, but femora IV(3:2:2). Lessert's type is missing left
legs II+III as well as metatarsus and tarsus of legs I+II and tarsus of leg IV on both
sides. Measurements (based on remainder of legs):
Palpus I II III IV
Femur 4.13 11.88 12.25 9.38 11.88
Patella 1.50 4.50 4.00 3.50 3.50
Tibia 2.25 11.25 11.50 8.50 9.38
Metatarsus 7.13 8.63
Tarsus 3.88 2.13
Total: 11.76 30.64
Additional material examined: TANZANIA: Nyika Plateau, 6000-7000 ft, 39

8subad.9 2subad.0", H. H. Johnston (BMNH 97.4.25.29-34). MALAWI: Kondoroi to
Koronga, 2000 ft, 39 10" 1subad.0" limm. (BMNH 97.4.25.35-38); Koba Koba [Nkota
Kota], Universities Mission, jar of specimens (BMNH 96.12.4-6). RWANDA: Nyanza,
nr Astrida, 1939, 19, Lestrade (MRAC 22300); Ruhengira [?Ruhengiri], 1931, 19
with egg sac, Lestrade (MRAC 22314). UGANDA: Ruwenzori [Mokia], SE., 3400 ft,
v.1906, 1 9 , spiders from deep ravines full of thick undergrowth (BMNH 07.2.1.68-73-
part); Ruwenzori [Mubuku Valley], 7000 ft, iv.1906, 19 (BMNH 07.2.1.1-20-part);
Ruwenzori, Namwamba Valley, 9000 ft, 19, Dr J. W. Edwards, British Museum
Expedition (BMNH 1935.5.23.1). ZAIRE: Kivu: Bwido, 19 , 1934, Marlier (MRAC
12443); Bukavu, Lake Kivu, 19 ,R. Laurent (MRAC 130.326); Goma, Lake Kivu, 10",
in pitfall trap, x.1988, M. Losseau (NMSA); Lwiro River, 47 km N. Bukavo
[=Costermansville, southern edge, Lake Kivu], 1950 m, 10", 27.viii.1957, 19 10",
15.xii.1957, E. S. Ross & R. E. Leech (CASC); w. Lake Kivu, 19, 1938,
Vande1ennoit (MRAC 831); Ibanda [?Bobandana], W. Lake Kivu, 29, 1935,
Vandelennoit (MRAC 26096/ 26097); Kadjudju, W. Kivu, 19 , 1932, L. Burgeon
(MRAC 16277); Kashusha, W. Lake Kivu, 29 2subad.0", 1931, Vandelennoit (MRAC
22296/22299); Katwe, 1600 m, 19 , 20.vi.1934, Marlier (MRAC 12444); Tshibinda
[just SW of Lake Kivu], 1&, 1932, L. Burgeon (MRAC 29992).
Palystes hoehneli Simon, 1890

Figs 89-96, 109
Palystes hOhneli Simon, 1890: 125. Holotype: adult 9, 'Kilimanjaro' [Tanzania], Hohnel (MNHN
10632) [examined].
Palystes rubrioculatus Strand, 1906c: 689. Holotype: 6, 'Moschi, Kilimandscharo', Dr Widenmann
(SMNS, destroyed). [Synonymised by Strand 1907c: 91.]
Palystes kibonotensis Lessert, 1921: 393-397, Figs 14, 15, 17, 18. Lectotype 6 paralectotype 9 (here
designated): 'Kibonoto' fKilimanjaro Mountains], Prof. Sjostedt (NHRS) [examined]; 7639 ,
5 subadult 9 paralectotypes (here designated): 'Kibonoto', Collection Roger Lessert, No Cl 8,
VII-XI (MHNG) [examined]. Syn. n.
Palystes bomemiszai Caporiacco, 1947: 205-206, PI. II, Fig. 49. Syntypes: 2619, 'Kilimandjaro'
(HNHM, not yet found), 16, 'Africa Or.' [East Africa] (HNHM, not examined). Syn. n.
Remarks: The specimen described by Simon (1890) as P. hOhneli [here emended in
terms ofthe Code], was collected on Mt Kilimanjaro by M. L. von Hahnel. The adult
female in MNHN is clearly the holotype.
Undetermined specimens of P. rubrioculatus collected by Dr Widenmann, at the
type locality, are recorded in an old inventory book at SMNS (F. Renner, in. litt.
1988), but were destroyed in 1944.
Lessert (1921: 393-397) described P. kibonotensis on a syntype series from
'Kibonoto' [probably Kibongoto near Mt Kilimanjaro] collected by the 1905-1906
Swedish expedition. He described an adult male and adult female, but listed (p. 396)
material examined as: '0" 9 , types, 60", 49 ad., et plus. subad. VII-XI; 1 9 avec
cocon, V'. The adult male and female in NHRS and the two vials of specimens
in MHNG are undoubtedly part of the same syntype series. The combined number
of syntypes for P. kibonotensis in both museums is 80"49 , 5 subadult 9 (+ one egg-
sac). There thus appear to be several more syntypes than originally listed. I have no
~ o(:)C5
~
.....~.
f
94
''\ i,
,
('I
,/
/
.l
..-...,/
89
90
Figs 89-96, Palystes hoehne Ii Simon, 1890. 89. External female epigynum. 90. Internal epigynum
(holotype female, MNHN 10632, Kilimanjaro). 91. Retrolatera! aspect, male pedipalp. 92.
Ventral aspect. 93. Detail of embolus (lectotype male, P. kibonotensis de Lessert, NHRS,
Kilimanjaro). 94-96. Features of ho!otype female, P. hoehneli. 94. Left chelicera! dentition.
95. Ventral markings. 96. Lateral markings, also showing black marking on underside of
femora I-II.
doubt, however, that all the above specimens constitute the syntype series, The adult
females are all con specific with the holotype of P. hoehneli Simon, and the males are
of the same species. I have therefore recognised P. kibonotensis as a junior synonym
of P. hoehneli. I have designated the adult male in NHRS as lectotype and the
remaining specimens listed above as paralectotypes,
Caporiacco (1947) described Palystes bornemiszai on a female and two males
collected by Bornernisza from 'Kilimandjaro', and a female collected from 'Africa
Or.' [East Africa] by Kittenberger in HNHM. The specimens were part of a
collection made by Kittenberger, Kovacs, and Bornernisza in East Africa. The type
material has not yet been found, but the illustration of the male palp (Plate II, Fig.
49) clearly depicts the Palystes hoehneli embolus, with the unmistakable shape of its
distal process and its medially bulbous column. The description of the female
matches that of Simon's type specimen. Caporiacco notes that while the male palp
resembles that of Lessert's P. kibonotensis, the specimen differs in that the sternum is
darker. There is considerable variability in the intensity of colouring of the sternum
in P. hoehneli and other members of the P. superci/iosus group. Despite the absence
of the type material, in the light of the matching descriptions and figured male palp, I
have little hesitation in regarding P. bornemiszai as a junior synonym of P. hoehneli.
Diagnosis: Afferent sperm duct of epigynum (Fig. 90) embedded anteriorly on lobe
of support collar (not detached as in P. ansiedippenaarae), lobe of support collar not
anteriorly strongly produced. Distal process of embolus (Figs 92, 93) pronounced,
short and flat. Column of embolus (Fig. 93) distinctly bulbous medially,
Distribution: Kenya (Mount Kenya) and Tanzania (Mount Kilimanjaro).
Redescription of holotype female of Palystes hoehneli:

Colour and markings: Abdomen dorsally with typical pulmonary marking,
laterally (Fig. 96) with typical pale cream band, edged dorsally with undulating dark
chocolate line, ventrally with faded brown blotches posterior to epigastric groove,
Sternum (Fig. 95) with 2 dark transverse bands in line with coxae II and III. Anterior
margins of coxae dark. Legs with typical markings, basal half of femora ventrally
dark, bases of femora I-II darker than III-IV, tibiae with ventral bands.
CWO Eyes: Eye diameters: AME 0.46; ALE 0.60; PME 0.46; PLE 0.46; PER 3.19
AME-ALE 0.12; AME-AME 0.37; AME-PME 0.69; ALE-PLE 0.57; PME-PLE
0.57; PME-PME 0.54; MOQ dimensions: MOQA 1.26; MOQP 1.45; MOQL 1.45;
MOQ as long as wide, narrowed anteriorly, with MOQP 1.21 X wider than MOQA.
Chelicerae: 3.94 long, 2.19 wide, 1.8 X longer than wide. Sternum: 4.0 long, 3.44
wide between coxae II, 1.16 X longer than wide and 2.88 wide between coxae I.
Labium: 1.85 wide, 1.23 long, 1.5 X wider than long, occupying 0.64 X of SW
Epigynum: Externally, median septum (Fig. 89) 'bow-tie' shaped, with posterior
elongation found in all members of species group (P. superciliosus excepted).
Internally (Fig. 90) afferent sperm duct embedded dorsally on support collar and
spermathecae protruding ventrally and partially encircled by internal lobe of support
collar.
Legs: Spination typical of genus. Both leg III tarsi missing. Measurements:
Palpus I II III IV
Femur 3.75 10.88 11.50 9.13 11.50
Patella 2.00 4.75 4.75 3.75 4.00
Tibia 2.50 10.25 10.25 7.50 8.75
Metatarsus 9.00 9.38 6.25 8.75
Tarsus 4.25 2.63 2.63 2.25
Total: 12.5 37.51 38.51 35.25
Description of lectotype male of Palystes kibonotensis:

Colour and markings: As for female, but lateral abdominal markings faded and
femora ventrally all dark, not just basally (as with most males in this species group).
Pedipalp: Embolus with lamellate carina apically on same plane as blade, opposite
to lower, proximal, prominence, neck short and small. Conductor membranous, long
and attentuate basally.
Legs: Spines typical of genus, but femora IV(3:2:2), tibiae I-IV (left) and tibiae
III+IV (right) with distal and proximal spines dorsally, right tibia I with proximal
only and right leg II missing. Measurements (left legs):
Palpus I II III IV
Femur 3.13 10.63 10.88 8.38 10.63
Patella 1.63 4.00 4.13 3.38 3.25
Tibia 1.75 10.38 10.25 7.13 8.50
Metatarsus 9.88 7.13 6.50 8.38
Tarsus 3.50 2.75 2.38 2.13 2.25
Total: 10.01 37.64 37.27 27.52 33.01
Additional material examined: KENYA: Mt Kenia, Fort Hall, 8500 ft, 2d49
7subad.d 10 subad.9 , x.1909, Smithsonian Afr. Exped. 1909-1910 (USNM); Kapiti
Plains [south of Nairobi], Id, v.1909, Roosevelt Expedition (USNM).
Paiystes johns toni Pocock, 1896

Figs 97-100, 109
Palystes johnstoni Pocock, 1896: 57-58. Pl. VIII. Figs l-Ic. Lectotype adult 9, paralectotype adult d
(here designated): Malawi: Zomba, between 3000 + 9000 feet, A. Whyte (BMNH
1894.1.15.21,22) [examineq).
Remarks: The species description was based on an adult male and adult female
collected in Zomba (MalaWi) at altitudes between ca 900-2700 m. The specimens in
BMNH match Pocock's description and illustrations, and the label data agree with
the information provided by Pocock. I have no doubt that these are the syntypes seen
by Pocock, and I have designated the female as lectotype and the male as
paralectotype. Both specimens are intact and have all their legs attached except that
the male's right leg II is detached from the apex of the femur and has no attached
pedipalps (the right pedipalp is in a rnicrovial inside the vial containing both
specimens but the left pedipalp is missing). Pocock refers to a third specimen, 'an
immature specimen of apparently the same species', from Ugogo [probably Ugogo
River, southern highlands of Tanzania] donated by Emin Pasha. I have not found this
specimen.
Diagnosis: Sternum with 2 dark transverse bars, abdomen ventrally with dark brown
spots, spotting varying from lightly spotted to heavily blotched posterior to epigastric
groove. Epigynum internally (Fig. 98) with spermathecae attenuate and detached
from, but ventral to, internal support collar and usually hidden from view dorsally
when epigynum is dissected. Embolus (Figs 99, 100) thick, recurved though 180°,
with a prominent distal lateral ridge (Fig. 100) crossing over onto ventral face of
blade and with short, blunt-tipped hook apically.
Distribution: Palystes johnstoni is widespread in Zimbabwe and Malawi, and reaches
just east of Umtali into Mozambique (Pungwe River) and south into the Okavango
delta of Botswana. It has not been found elsewhere except for specimens collected by
P. L. G. Benoit (MRAC) in Mityana (80 km north-west of Entebbe, Uganda).
Redescription of lectotype female:

Colour and markings: Abdomen dorsally with dark-outlined pulmonary marking,
laterally with pale longitudinal cream band on either side, edged dorsally with
undulating dark chocolate line, ventrally with dark transverse band posterior to
epigastric groove. Area between band and spinnerets spotted with brown, spots faded
in type specimen but usually ranging from small to large, nearly coalescing, dark
brown blotches, often forming a rectangular darker panel between epigastric groove
and spinnerets, flanked by smaller spots. Sternum with 2 faded, rubbed, dark
transverse bars (in fresher specimens bars generally clearly defined and dark black).
eye ratio AME:ALE:PME:PLE is 1:1.47:1:1; distances separating eyes: AME-ALE
0.18, AME-AME 0.42, AME-PME 0.94, ALE-PLE 0.77, PME-PLE 0.69,
PME-PME 0.69; MOQ dimensions: MOQA 1.45, MOQP 1.78, MOQL 1.69, MOQ
0.95 X longer than wide, narrowed anteriorly, with MOQP 1.2 X wider than MOQA.
Chelicerae: 5.0 long, 2.8 wide, 1.78 X longer than wide. Sternum: 4.8 long, 4.38
wide between coxae II, 1.1 X longer than wide and 4.06 wide between coxae I.
Labium: 2.09 wide, 1.48 long, 1.42 X wider than long, occupying 0.51 X of SW
Epigynum: Epigynum fully formed but sclerotisation incomplete in type specimen,
with bulbous lobes of the well when viewed externally (Fig. 97) still pale and not
dark red-brown as is more usual (suggesting a newly adult female). Internally the
long, thin spermathecae are hidden from view behind lobes of support collar (Fig.
98).
Legs: Spine pattern and markings typical of genus, but femur IV(3:2:2).
Measurements:
Palpus I n III IV
Femur 4.38 14.38 14.75 11.50 14.63
Patella 2.13 5.63 5.75 4.75 4.50
Tibia 3.13 13.38 13.75 10.25 11.25
Metatarsus 12.50 12.13 8.38 11.00
Tarsus 5.50 3.13 2.88 2.25 2.63
Total: 15.14 49.02 49.26 37.13 44.01
Redescription of paralectotype male:

Colour and markings: As for female, but markings clearer and not as faded, and
femora ventrally all dark (as with most males in this species group).
eye ratio AME:ALE:PME:PLE is 1: 1.27:0.93:0.93; distances separating eyes:
X longer than wide and 2.25 wide between coxae 1. Labium: 1.57 wide, 0.98 long,
Pedipalp: Embolus (Figs 99, 100) thick, recurved though 180, with prominent
distal lateral ridge crossing onto ventral face of blade and with short, blunt-tipped
hook apically. Embolus neck distinct, conductor membranous, short, wide and
abruptly terminated basally (Fig. 99).
Legs: Spines of right legs typical of genus, but dorsal proximal and distal spines
missing on II and distal only on regrown III. Spine pattern otherwise as for genus.
Measurements:
Pal pus I II ill IV

Femur 3.75 12.63 12.50 9.88 12.25
Patella 1.75 4.38 4.75 3.50 3.50
Tibia 2.50 12.50 12.50 9.13 10.00
Metatarsus 11.50 11.00 7.50 9.75
Tarsus 4.25 2.50 2.50 1.88 2.25
Total: 12.25 43.51 43.25 31.89 37.75
Additional material examined: BOTSWANA: Okovango [Okavango] Delta, 4-rivers

camp, 19°03'S 23°lO'E, 10", 7.xii.1973, Falcon College & NMBZ expedition (NMBZ
NMZlA932). MALAWI: Zomba, A. Sharp (BMNH); Zomba, 19,1. R. Lennon, ca
1950 (BMNH). MOZAMBIQUE: Pungwe River, 50 mls E. Umtali [Zimbabwe],
40"69 4subad.9, 1903, D. Patrick (SAMC 13621). UGANDA: Mityana, 10"
(pedipalp only), 19, 1959, P. L. G. Benoit (MRAC 146.163). ZIMBABWE:
Birchenough Bridge, Sabi River, 19, i.1938, V. Fitzsimons (TMSA 8175);
Bulawayo: 10", 21.ix.1979, A. Thompson (NMBZ NMZlA630), 20", x.1960 (NMBZ
NMZ/AI148), Kumalo, 20 o lO'S 28°35'E, ld', 2.ix.1979, G. Allen (NMBZ
NMZ/A158), Matsheumhlope, 28°lO'S 28°35'E, 10", 30.vi.1979, D. Wheeler (NMBZ
NMZlA660), Matsheumhlope, 28°lO'S 28°35'E, 20"29 2subad.9 1subad.d',
ix.1979, D. Wheeler (NMBZ NMZlA689); Cement, Montgomery, 20 0 07'S 28°45'E,
19, 23.vii.1979, W. Black (NMBZ NMZ/A108); Chishawasha [17°47'S 31°14'E],
19, 26.x.1983, A. Mukondo (NMBZ NMZlA2018); Esigodini [30 km SE.
Bulawayo], Falcon College, 19 , 19.xi.1984, M. Bing (NMBZ NMZ/A2767); Harare
[= Salisbury]: 10", 2l.ix.1985, S. A. L. White (NMBZ NMZ/A1233), 19,
14.xi.1967, R. A. Bailey (NMBZ NMZ/AI440), Mabe1reign, 10", l1.xi.1968, M. A.
Raath (NMBZ NMZlAI549), 19, 3.ix.1967, D. J. Woodiwiss (NMBZ
NMZ/AI212), 10", 19.ix.1966, B. W. Blair (NMBZ NMZlA1300), 10", 13.x.1969, S.
Holland (NMBZ NMZ/AI556), Haig Park, Isubad.9 , 6.ii.l971, V. Bosselli (NMBZ
NMZ/A1470), Isubad.9 , ix.1967, Miss Levesque (AMOS), 19 , 17.ix.1968, Miss J.
Blowers (AMGS); Kharni, 20010'S 28°25'E, [12 km SW. Bulawayo], 19 , x.1965
(NMBZ NMZlA723); Matopos [35 km sw. Bulawayol, R. E. P. School, 20022'S
28°35'E), 30"49, 18-23.ix.1979, 7.x.1979, 16.x.1979, S. Higgins (NMBZ
NMZlA424, NMZlA319, NMZlA1238); Matopos, Maleme, 20 0 32'S 28°30'E, 10",
20.viii.1978, C. A. Car (NMBZ NMZl1l61); Mazoe, 10" Isubad.9, J. Darling
(BMNH); Mine Hill, Lalapanzi Distr., Uplands Farm, 19, 18.ii.1985, G. Allen
(NMBZ NMZ/A3709); Murambinda [137 km S. Harare], Isubad.d', ix.1984, Hindley
(NMBZ NMZ/A2931); Mutare [= Umtali]: 29 , 3.xi.1984, S. van der Pyll (NMBZ
NMZlA2214), 19 Isubad.9, 20+24.xi.1898, J. F. F. Darling (BMNH
1898.11.20.24), 1 9 (office - bit !!lan - chest pains, difficulty in breathing. Treated
with antihistamines), vi.1982, Mrs 1. Brown (NMBZ NMZ/AI562), 19 ,27.xii.1965,
D. Walton ([?NMBZ] UM/S 398), 20", ii.1903, Miss Campbell (AMGS), 19,
6.x.1961,1. Weiman (NMBZ NMZlAI429), 60"59 , ix.1916, H. Hosgood (AMGS),
Fairbridge Park, 10", 14.x.1984, P. Locke (NMBZ NMZ/A2712), hospital, 19,
30.ix.1966, Palgrave (NMBZ NMZlA1050), Penhalonga, 18°53'S 32°43'E, Hillcrest
School, 10", 16.xi.1984, S. van der Pyl (NMBZ NMZlA27 17), Penhalonga, Hillcrest
School, 19 , 1.ii.1985, Mr Ehlinger (NMBZ NMZ/A3413), Penhalonga, 19 , x.1943
(NMBZ NMZ/A1215), Yeovil, 29 , 16.xii.1966, B. Marais (AMGS), Yeovil, 19 ,
16.xii.1966, B. Marais (NMBZ NMZIA726), Baviaanskop, 30"29 1subad. 0"
\
99
{
"\
f
(
'. ,., :
\ !
i
./
,/
100
102 103
Figs 97-103. Palystesjohnstoni Pocock, 1896 and P. leroyorum sp. n. 97-100. P. johnstolli. 97. External
female epigynum. 98. Internal epigynum with dotted outline of spermathecae lying beneath
invaginated collar of septum (lectotype female, BMNH 1894.l.15.21-22, Zomba, MalaWi). 99.
Ventral aspect, male pedipalp. 100. Detail of apex of embolus (paralectotype male, BMNH
\894.1.15.21-22, Zomba, Malawi). 101-103. P. leroyorum. IOJ. Detail of embolus. 102.
Retrolateral aspect, tibial apophysis. 103. Ventral markings of spider (holotype male, NMBA
1395, Bloemfontein).
Isubad.9, vi. 1902, D.- Patrick (SAMC 12543), Queenskop, 19, i.1902, D. L.
Patrick (SAMC 12532), Greenside, 1d, 20.xi.1965, Palgrave (UMIS 315);
Nyamandhlovu, Bonisa Farm, 19°50'S 28°20'E, 2d, 26.ix.1979, 20.xii.1979, T. Gibbs
(NMBZ NMZlA30l); Tuli Road [21°S 29°E], Id Isubad.d 3subad.9 , 22.vii,1979,
D. K. B. Wheeler (NMBZ NMZ/A1552); Vumba, south of Umtali, 19 (TMSA
1415). PROVENANCE?: Id29 ,incompletely labelled (SAMC 3848).
Palystes ansiedippenaarae sp. n.

Figs 104-108,109
Type material:
Holotype: adult d, SOUTH AFRICA: Transvaal: Warburton, lessievale Forest
Reserve [26°15'S 30 0 32'E], under dry pine bark, 23.vii.1979, M. Stiller (PPRI AcAT
'801179).
Paratypes: SOUTH AFRICA: Transvaal: Warburton, lessievale Forest Reserve
[26°15'S 30 0 32'E]: under dry thin bark of pine log, l.viii.1979, M. Stiller (PPRI
AcAT '80/187), Id, under flap of cardboard box, 24.iii.1979 (PPRI AcAT '80/184),
10', same data but 22.vii.1979 (PPRI AcAT '801199) Id, same data but, 24.vii.1979
(NMSA T497), 19 , under dry pine bark, 20.vii.1979 (NMSA T498), 19 , same data
but, 23.vii.1979 (PPRI AcAT '801173),19, under dry bark on pine log, 19.vii.1979
(PPRI AcAT '80/197), 1 9 , under dry, thin bark of Pinus patula log, 13. vi.1979
(PPRI AcAT '80/211), 19, under dry bark on pine logs, 19.vii.1979 (PPRI AcAT
'80/166), 19, under dry thin bark of dead pine tree, 4.viii.1979 (PPRI AcAT
'801198), I 9 , under flap of cardboard box in plantation, 22. vii. 1979 (PPRI AcAT
'80/172); Ermelo district, Aarthol, 1 9 , 25.I.1911, M. Forbes (TMSA 1111). Natal:
Ndumu Game Reserve, 26°53'S 32°16'E, 19 ,i.1968, T. Pike (NMSA 12387).
Type locality: Warburton district, 40 km SE. Carolina, Jessievale Forest Reserve,
26°15'S 30 0 32'E, south-eastern Transvaal, South Africa.
Etymology: Named in honour of Dr Ansie Dippenaar-Schoeman who has played a
leading role in promoting contemporary arachnological research in South Africa and
was responsible for the collecting surveys that led to the discovery of this new
species.
Diagnosis: Distinguished from P. superciliosus in the markings and form of genitalia.
Generally smaller and darker than P. superciliosus, with sternum either black with no
other markings or dark with 3 black transverse bars coalescing mesially (as opposed
to single dark bar in P. superciliosus), and with dark spots ventrally on abdomen
between transverse dark bar posterior to epigastric groove (as opposed to either a
greyish bell-shaped marking or no marking but 4 longitudinal striae in P.
superciliosus). Epigynum (Fig. 104) differs externally from P. superciliosus in that it
is produced posteriorly and not contracted (as in Fig. 77), while internally (Fig. 105),
the spermathecae are enfolded by support collar lobes, but not horizontal and lying
dorsally on top of lobes (Fig. 78). Embolus (Fig. 108) with lateral carinations
distally, as opposed to no carination in P. superciliosus (Fig. 80). Adult male
distinguished from its closest relative, P. hoehne Ii from east Africa, in form of
embolus which is slender mesially (Fig. 107) and not bulbous (Figs 92-93) and in
distal carination which is reduced and elongate (Fig. 108) as opposed to pronounced,
short and flat (Fig. 93), and proximal carination which is opposite distal carination
(as opposed to closer to embolar tip). In female afferent sperm duct is external to
lobe of supporting collar (Fig. 105) as opposed to embedded (Fig. 90) .
..........: ...•.... j
108
106
107
Figs 104-108. Palystes ansiedippenaarae sp. n. 104. External female epigynum. 105. Internal female
epigynum (paratype female, PPRI AcAT '80/187, Jessievale Forest Reserve, Warburton). 106.
Retrolateral aspect, male pedipalp. 107. Ventral aspect, male pedipalp. 108. Detail of apex of
embolus (holotype male, PPRI AcAT '80/197, Jessievale Forest Reserve, Warburton).
Distribution: From western Transvaal (Nylstroom) through south-eastern Transvaal

(Ermelo and Warburton districts) to northern Zululand, Natal.
Colour and markings: As in P. hoehneli, male darker in colour than other species
in superciliosus group. Abdomen generally dark and mottled, lateral longitudinal
undulating markings only just discernible. Dark markings a rich dark brown, varying
from burnt umber to raw sienna, to black. Ventrally, a dark transverse bar
immediately posterior to epigastric groove with irregular dark liver-brown blotch
markings between bar and spinnerets, with those in centre region darker tharl the rest.
Sternum dark with a yellowish patch posteriorly. Three broad dark bars across
sternum, in line with coxae II, III and IV, coalescing at centre and occupying most of
sternum. Dark bars continue on arlterolateral faces of coxae and femora ventrally.
Much of labium and maxillae also dark ventrally.
wide across PER; thoracic fovea 2.25 long, occupying 0.24 X of CL arid 0.29 X of
wide, wider tharl AER by 1.10 X and occupying 0.66 X of HW and 0.35 X of CW;
MOQ 1.02 X longer than wide; narrowed anteriorly, with MOQP 1.58 X wider tharl
MOQA; clypeus 0.38 high, 0.86 X diameter of AME. Chelicerae: 3.69 long, l.88
X longer than wide arid 2.63 wide between coxae I. Labium: 1.66 wide, l.08 long,
long, 1.45 wide, 1.81 X longer tharl wide.
Pedipalp: Embolus (Figs 107, 108) with proximal lateral carination small arid
closer to embolus tip than to distal carination. Embolus terminates in arI elongate,
acicular point (Fig. 108).
Legs: Spines typical of genus but left femur IV(3:2:2), right femur IV(3:2:1).
Measurements:
Palpus I II III IV
Femur 4.25 13.88 14.13 10.88 13.25
Patella 1.88 5.13 5.13 4.00 4.38
Tibia 2.50 13.38 13.50 9.75 11.25
Metatarsus 12.63 12.50 8.13 10.88
Tarsus 4.38 3.00 3.00 2.25 2.75
Total: 13.01 48.02 48.26 35.01 42.51
Description of paratype female (PPRI AcAT '801187):

Colour and markings: Generally dark. Abdomen dorsally with pulmonary
marking, laterally dark, but with paler longitudinal band on either side, edged
dorsally with undulating darker line, ventrally with dark transverse barld posterior to
epigastric groove and dark liver-brown spots or blotches. Area between barld arid
spinnerets darker with larger spots. Sternum dark, ranging in other specimens from
black with no evident markings to dark with 3 black trarlsverse bars at coxae II, III
and IV, coalescing mesially. Femora I and II ventrally dark with whitish yellow patch
distally. Tibiae I-IV with alternate 2 dark and 2 pale bands ventrally.
wide across PER; thoracic fovea 2.00 long, occupying 0.20 X of CL arid 0.25 X of

MOQA. Chelicerae: 3.63 long, 2.13 wide, 1.71 X longer than wide. Sternum: 3.63
coxae I. Labium: 1.69 wide, 1.23 long, 1.38 X wider than long, occupying 0.56 X of
SW between coxae I. Maxillae: 2.49 long, 1.54 wide, 1.62 X longer than wide.
Epigynum: Externally (Fig. 104), median septum produced posteriorly. Internally
(Fig. 105), tubules leading to spermathecae (afferent sperm ducts) external to lobes
of support collar. Spermathecae enfolded by support collar lobes, but visible, not
embedded.
Legs: Spine pattern generally as for genus, but in described female some spines
missing on some legs: left femur 1(2:2:3), right femur 1(3:2:3), left femora
II-III(3:2:3), right femur II( I :2:3), right femur III(3:2:3), right and left femora
IV(3:2:1). Measurements:
Palpus I II III IV
Femur 3.88 11.63 11.88 9.13 11.38
Patella 2.00 4.75 4.88 3.88 3.88
Tibia 2.63 11.38 11.25 8.13 9.00
Metatarsus 10.25 9.75 6.63 8.75
Tarsus 4.38 2.75 2.75 2.00 2.25
Total: 12.89 40.76 40.51 29.77 35.26
Palystes leroyorum sp. n.

Figs 101-103, 109
Type materiaL:
Holotype: adult d, SOUTH AFRICA: Orange Free State: Bloemfontein,
30.i.1986, L. de Wet (NMBA 1395).
Paratypes: SOUTH AFRICA: Orange Free State: Bloemfontein, ld, in house,
2.ix.1989, L. Barkhuizen (NMBA 3268); Bloemfontein, ld, in cupboard, 26.ix.1989,
L. Barkhuizen (NMBA 3273). Transvaal: 70 km NW. Nylstroom, 1d, 6.iv.1994, M.
Filmer (NMSA); Johannesburg Zoo, found x.1986, preserved vii.1987, M. Filmer
(NMSA).
Type Locality: Bloemfontein, Orange Free State, South Africa.
Etymology: Named in honour of John and Astri LeRoy for their contribution towards
promoting arachnology to a wider audience in southern Africa, both through the
Spider Club of South Africa, and through their dedicated fieldwork and assistance to
all arachnologists.
Diagnosis: Ventral abdominal markings (Fig. 103) and form of tibial apophysis (Fig.
102) and embolus (Fig. 101) readily distinguish P. leroyorum from its closest
relatives, P. ellioti and P. superciliosus. In P. leroyorum area between black bar
immediately posterior to epigastric groove and spinnerets peppered with irregular,
fine black spots (as opposed to brown blotches coalescing medially in P. ellioti, or no
markings or faint grey shield marking in P. superciliosus). Tibial apophysis in P.

leroyorum sharply bent and projects ventrally (Fig. 102), compared with relatively
straight and directed apically in other 2 species (Figs 79, 83). Embolus in P.
leroyorum very broad and flat apically with broad, unnotched inner flange (Fig. 101).
In P. superciliosus embolus narrow and acicular apically, while in P. ellioti it is broad
and flat but without wide inner flange. Female unknown.
Distribution: Bloemfontein area (Orange Free State) and Nylstroom district, western
Transvaal.
Colour and markings: Dorsally, a dark mottled greyish brown and black. Abdomen
dorsally with a darker cardiac outline marking flanked by paired paler patches
diminishing in size and merging with background colour posteriorly. Abdomen
laterally darkly mottled, paler below with dark grey-black stippling. Ventrally (Fig.
103), a dark broad black transverse bar immediately posterior to epigastric groove,
followed by an irregular scattering of intense black spots, some coalescing, against a
pale background. Femora ventrally dark with lacework of fine, irregular white
spotted markings. Sternum with black solid bar at coxae II in line with dark
anterolateral aspects of coxae.
X longer than wide and 3.19 wide between coxae 1. Labium: 1.75 wide, 1.17 long,
1.50 X wider than long, occupying 0.55 X of SW between coxae 1. Maxillae: 2.52
Pedipalp (Figs 101-102): Embolus reflexed distally with a wide, flat, unnotched
inner flange. Tibial apophysis entire but bent sharply at base to project ventrally.
Legs: Spines typical of genus with femora I-III(3:2:3), femora IV(3:2:2); tibiae
I-IV with dorsal proximal and distal spines present. Measurements:
Pal pus I II III IV
Femur 4.13 13.13 13.63 10.88 13.38
Patella 1.63 5.00 5.13 4.13 4.00
Tibia 2.50 13.25 13.38 9.75 11.13
Metatarsus 12.25 12.25 8.75 11.00
Tarsus 4.38 3.13 3.13 2.38 2.75
Total: 12.64 46.76 47.52 35.89 42.26

Fig 109. Map showing distribution of the Palystes superciliosus species group . • = P. ellioti; = 0; P.
hoehneli; ... = P. johnstoni;. = P. superciliosus; 0 = P. leroyorum; 6. = P. ansiedippenaarae.
Palystes species incertae sedis

The following nominal species are removed from Palystes s.s. for the reasons
stated below they cannot be assigned with confidence to any genus known to me and
are therefore listed in their original combination, and treated as incertae sedis:
Palystes flavidus Simon, 1897

Palystes flavidus Simon. 1897b: 489. Holotype: subadult I( • 'Trichonopoly' [Indial. (MNHN 10907).
[examinedl.
Remarks: In addition to the subadult female, there is an adult female in the vial
together with a microvial containing the detached epigynum of the adult. Simon
probably added the adult later. The specimens appear to belong to the same species,
but neither belong in either Palystes or Gnathopalystes. The cheliceral dentition (four
teeth on retromargin, three on antero-margin with pad of microdentic1es on fang
furrow floor), the epigynal structure (with lateral lobes separated by median septum),
posterior eye row (slightly recurved, PLE raised on tubercles) and proportions of the
eyes suggest, however, that the species belongs in Heteropodinae s. s.
Gravely (1932) states that the live female is pale green and the live male more
yellowish green. His illustration (1932: 258, Fig. 14a) of the female epigynum,
however, differs considerably from that of the adult female in MNHN.
The BMNH has a number of specimens determined as Palystes flavidus from
'Allahabad', 'Chingleput', Madras and Calcutta.
Palystes fornasinii (Pavesi, 1881)

Sparassus fomasinii Pavesi, 1881: 548, holotype: adult d, 'Inhambane', Mozambique (type not MCSN)
(not examined).
Palystesfomasinii; Simon, 1903: 1028 (transferred to Palystes).
Remarks: The holotype of S. fornasinii is not in MCSN and has not been located
elsewhere. The description places it outside Palystes, considering the described eye
proportions and arrangement, and cheliceral dentition. The latter (retromargin with
four large teeth, antero-margin with one large tooth) could agree with the male of the
female illustrated and described by Jarvi (1914: 197) as Heteropoda venatoria
(Linnaeus, 1767), if it is possible that Pavesi had not observed the two very small,
teeth that flank the large tooth of the antero-margin. The rest of the description does
fit that of males identified as Heteropoda venatoria (sensu Jarvi), which have been
collected in Mozambique and Cameroon. The description, however, does not provide
sufficient specific information to justify its transferral to Heteropoda. In the absence
of the type material and being unable to place it elsewhere, I regard the name as
incertae sedis.
Palystes pinnotherus (Walckenaer, 1805)

Thomisus pinnotheres [sic] Walckenaer, 1805: 36. Provenance: 'Notasie'. [Specimen from 'Collection du
Museum de Paris' (Walckenaer, ibid.)] (?MNHN). Nomen nudum [not located, presumed lost].
Olios pinno/herus Walckenaer, 1837: 565. Syntypes: d9 , 'Monde-Maritime-Notasie-Du Port-Jackson,
dans la Nouvelle-Holland' (?MNHN) [types not located, presumed lost].
Palystes pinnotherus; L. Koch 1875: 703 (redescribed as an included species of Palystes).
?Thomisus lamarck Latreille, 1806: 113. SexJprovenance unknown (?MNHN) [not located, presumed
lost].
?Thomisus lamarckii [sic]; Walckenaer, 1837: 566 (listed as a synonym of pinnotherus).
Remarks: Walckenaer (1837: 566) refers to an illustration 'fig. 29' illustrating
Thomisus pinnotheres in Walckenaer (1805). There is no illustration of pinnotheres in
the book. Fig. 29 is an illustration of Thomisus cancerides, according to both the
description of cancerides and the legend to the plates. The name pinnotheres was
published without a description and is therefore a nomen nudum. It only became
available when Walckenaer described the species in 1837. Walckenaer (1837)
synonymised Thomisus lamarck Latreille, 1806, with Olios pinnotherus. Since only
pinnotherus sensu Walckenaer, 1837, is valid, lamarck (Latreille, 1806) would be the
senior synonym. Bonnet, however (1958: 3312 footnote 24) says the names are not
synonymous. In the absence of the type material which has not been found, and in view
of the confusion of the identity of this species, the name is considered incertae sedis.
Palystes reticulatus Rainbow, 1899

Palystes reticulatus Rainbow, 1899: 312, PI. xxiv, Fig. 4. Syntypes: 3 immature 9 , Santa Cruz, Oceania,
[AMSA K27589 I KS 9301[on reverse]] [examined].
Remarks: The type material is immature but does not belong in Palystes. It belongs
within Heteropodinae because of the following combination of characters:

ALE»AME>PME=PLE; upper rims of AME on a common line with those of ALE;
presence of microdenticles on fang furrow floor; cheliceral dentition 4(proximal very
small): 2). It does not appear to belong in Heteropoda or Gnathopalystes because of
the combination of the very shallow clypeus, the width of the labium (wider than
long), the presence of PLE tubercles and the relative straightness of PER.
Palystes spiralis Strand, 1907

Palystes spiralis Strand, 1907d: 109 [amplified in Strand 1907e: 109]. Holotype: adult d, 'Nossibe',
Madagascar. Type not located (probably destroyed in Lubeck museum during second world
war).
Remarks: Strand (1907e: 112) noted that his species was not a typical Palystes, but that
the genus had many diverse genitalic forms. He suggested a subgeneric name
Spiralifera for spiralis (the subgeneric name has no nomenclatural standing). The
description excludes the species from Palystes, not only because of the domed
cephalothorax and the larger, slightly raised posterior lateral eyes, but also because of
the th~n, multi-spiralled embolus and separate conductor sheath described by Strand. If
no existing genus can be found for this species then Spiralifera should be recognised at
generic level, with spiralis as its type species (by monotypy). This Madagascan species
is possibly conspecific with P. convexus, discussed below. Until more material is seen,
P. spiralis is removed from Palystes and is regarded as incertae sedis.
Palystes convexus Strand, 1907

Palystes convexus Strand, 1907d: 737 [amplified in Strand 190e: 112]. Holotype: subadult <;>.
'Majunga', Madagascar; type in ZMHB according to Strand, not found [not examined].
Remarks: The description corresponds in all major respects with that of P. spiralis
Strand, 1907, including the leg spination, cephalothorax shape, proportions and
arrangement of eyes, and the cheliceral dentition. This may be the female of P. spiralis.
If the two names are synonymous, then P. convexus has priority as the preliminary
diagnosis of P convexus was published earlier in 1907. P convexus, like P spiralis, is
removed from Palystes on the same grounds. For the present, in the absence of the type
material, and in view of the description which does not provide sufficient information
to place the species in any genus known to me, it is regarded as incertae sedis.
Parapalystes gen. n.
Type species: Parapalystes euphorbiae sp. n. (north-western Cape, South Africa).
Etymology: Gr. para = near, beside, by. Refers to the sister-group relationship
between Parapalystes and Palystes. Gender masculine.
Remarks: Parapalystes is the putative sister group to Palystes. Its type species is
described below, and the included species, all from the Cape and transferred from
Palystes, are listed below. Specific synonymies are not given here, but will be dealt
with in a revision of Parapalystes. One of the included species, Ocypete
megacephala C. L. Koch, was described from material collected in the Cape
Province during the last century. The holotype has not been found, but the
description clearly places the species in Parapalystes. The genitalia, which provide
critical characters for identifying species in Parapalystes, were not described. Until
the type material is located or a neotype is designated, the name will remain
unidentifiable below generic level. In anticipation that the problem of its identity will
be solved in the future, the species is transferred to Parapalystes.
TABLE 2
Comparison of differences between Parapalystes gen. n. and Palystes s. s. L. Koch, 1875.
Parapalystes gen. n. Palystes s.s
Carapace:
• Domed posterior to AER. : • Flat posterior to PER.
• Posterior slope starts anterior to fovea, : • Posterior slope starts posterior to fovea, fovea
fovea inclined. : horizontal.
• Dorsally two fine white lines, thinly : • Dorsally pale median line often present but not
divided, from middle PER to white patch : as obvious.
I
immediately anterior to fovea. I
I
Male pedipalp:
• Primary and secondary tibial apophyses : • Primary apophysis only, present.
present. :
• Embolus subtended by well-developed : • Support flange reduced or absent.
support flange. I
• Conductor of bulb robust, often sc1erotised. :. Conductor fragile, diaphanous.

I
Female epigynum:
• Median septum at least 3.0 X wider than : • Median septum longer than wide or only
long. : slightly (1.1 X) wider than long.
Leg spination:
• Tibiae with one (distal) spine dorsally. I • Tibiae with two (proximal and distal) spines
: dorsally.
I
Markings:
• Sternum black, sometimes with large yellow :. Sternum either solid black or with dark
mark posteriorly. : transverse bars.
• Coxae each with several small to large black :. Coxae dark to black antero-Iaterally, but with
spots ventrally. no spots ventrally.
• Abdomen dorsally with solid brown I • Abdomen dorsally with outlined pulmonary
pulmonary marking. : marking, not solid.
Diagnosis: Similar to Palystes with respect to eye arrangement and proportions, and
general body shape and attitude. Differs from Palystes (see Table 2) in having 2 tibial
apophyses with additional ridges and sclerotised protrusions (as opposed to a single
apophysis without additional processes), presence of an additional membrane to
conductor lateral to embolus, wider than long epigynal septum with ear-shaped
lateral excavations, presence of ventral spot markings on coxae, sternum colour
(black with a pale yellow to white patch posteriorly), I (not 2) dorsal spines on tibiae
I-IV, carapace shape (domed posterior to AER), thoracic fovea situated on posterior
declivity of carapace (not anterior to it), carapace markings (2 thin white medial
stripes running from between posterior median eyes to an oval white patch
immediately anterior to fovea), yellow clypeus and yellow stripe or stripes often
vestigial or absent at outer margin of chelicerae (not white).
Distribution: Known only from South Africa, from the north-western Cape across the
interior to the eastern Cape.
Parapalystes euphorhiae sp. n.

Figs 11 0-115
Type materal:
Holotype: adult d: SOUTH AFRICA: Cape Province: Richtersveld [or near], ca 2
kIn E. Port Nolloth, found in nest in low leafless Euphorbia sp. shrub growing on
dune hummocks, 3.ix.1989, P. M. C. Croeser (NMSA T503).
Paratypes: SOUTH AFRICA: Cape Province: 6 <;> , same data as holotype (NMSA
T504); Richtersveld, ca 5 kIn E Port Nolloth, 1 <;> , found in nest in low leafless
Euphorbia sp. shrub growing on dune hummocks, 4.ix.1989, P. M. C. Croeser, J. O.
H. Landt, B. R. Stuckenberg (NMSA T504).
Type locality: The dune hummocks immediately inland of Port Nolloth in the north-
western Cape Province.
Etymology: Named after the succulent shrub genus Euphorbia, in which the spiders
were found at the type locality.
Remarks: The type specimens were all collected in leafless, waist-high Euphorbia sp.
shrubs scattered over dune hummocks of loose sand a few kilometres inland from
Port Nolloth. No more than one occupied nest was found in a shrub, although some
shrubs were more than a metre across, and only a single spider was found in each
nest. The nests were made by binding together vertical stems of the plants. The
spiders were found in the nests, in one case with a sac of eggs. The male was found
alone in an old nest. Size, as with most species of Palystes, can be very variable. A
male of P. euphorbiae collected in 1935 from Kleinsee (50 kIn S. Port Nolloth), is 27
mm long (more than twice the length of the holotype).
Diagnosis: The shape of the embolus (Fig. 112), the form of the primary and
secondary tibial apophyses (Fig. 113), the form of the median septum lobe of the
epigynum (Fig. 110) which is widely separated but with dorsally deflected lateral
lobes, and the position of the spermathecae internally (Fig. 111), distinguish the
species from known congeners.
Distribution: Port Nolloth, north-western Cape, South Africa.
Total length 11.00 mm.
Colour and markings: Dorsally (Fig. 114) a mottled light and dark brown. Abdomen
dorsally with solid, liver-brown dorsal undulating cardiac marking along back (Fig.
114); ventrally (Fig. 115) with short, thick black transverse bar immediately posterior to
epigastric groove, bar colour being emarginated medially at its posterior edge. Second,
shorter black marking immediately anterior to spinnerets. Sternum without markings,
dark. Coxae ventrally spotted with black and some fine black spotting ventrally and
dorsally on femora. Ventral tibial banding vestigial distally, especially on ill and N.
Carapace: Dorsally domed posterior to AER, 5.00 long, 3.88 wide between leg
bases II. Head region 2.25 wide across PER; thoracic fovea straight longitudinal 1.38
long, occupying 0.28 X of CL and 0.35 X of cwo Eyes: Eye diameters: AME 0.31,
ALE 0.49, PME 0.29, PLE 0.29; PER 1.81 wide, wider than AER by 1.16 X and
occupying 0.81 X of HW and 0.47 X of CW; eye ratio AME:ALE:PME:PLE is
113
111
112
115 114
Figs 110-115. Parapalystes euphorbiae sp. n. 110. External epigynum. Ill. Internal epigynum (female
paratype, NMSA T504, Port Nolloth). 112. Anteroventral aspect, male pedipalp. 113.
Retrolateral aspect, tibial apophyses (arrow shows secondary apophysis). 114. Dorsal
markings. 115. Ventral markings (male holotype, NMSA T503).
1:1.60:0.95:0.95; distances separating eyes: AME-ALE 0.03, AME-AME 0.20,

AME-PME 0.43, ALE-PLE 0.38, PME-PLE 0.28, PME-PME 0.28; MOQ
dimensions: MOQA 0.74, MOQP 0.98, MOQL 0.98; MOQ 1.00 X wider than long,
narrowed anteriorly, with MOQP 1.33 X wider than MOQA. Chelicerae: 1.69 long,
1.06 wide, 1.59 X longer than wide. Sternum: 2.13 long, 1.94 wide between coxae II,
1.10 X longer than wide and 1.38 X wide between coxae I. Labium: 0.92 wide, 0.40
long, 2.31 X wider than long, occupying 0.67 X of SW between coxae I. Maxillae:
1.02 long, 0.65 wide, 1.57 X longer than wide.

Pedipalp: Embolus blade (Fig. 112) reflexed distally, ventral to supporting,
flanged, median sclerite, conductor stalked, spathulate. Primary tibial apophysis a
sclerotised ridge between a proximal longer spur and a distal shorter spur. Secondary
tibial apophysis a spur projecting dorsally over paracymbium.
Legs: Spination irregular: left and right femora I+II(3:2:3), left femur III(3:2:2),
left femur IV(2:3:2); right femur III(3:2:3), right femur IV(3:3:3). Tibiae all with
only a distal spine dorsally. Measurements:
Palpus I II III IV
Femur 3.38 6.75 7.13 5.50 7.13
Patella 1.75 2.63 2.50 2.00 1.63
Tibia 2.13 6.38 6.38 4.75 5.13
Metatarsus 5.63 5.63 3.88 4.63
Tarsus 5.00 1.88 1.88 1.38 1.63
Total: 12.26 23.27 23.52 17.51 20.15
Description of paratype female (NMSA T504):

Colour and markings: As for male.
Carapace: Dorsally 7.25 long, 5.25 wide between leg bases n. Head region 3.69 wide
Eye diameters: AME 0040, ALE 0.58, PME 0.43, PLE 0.40, PER 2.63 wide, wider than
AME:ALE:PME:PLE is 1: 1.46: 1.08: 1.00; distances separating eyes: AME-ALE 0.08,
MOQ dimensions: MOQA 1.08, MOQP 1.32, MOQL 1.26, MOQ as long as wide;
narrowed anteriorly, with MOQP 1.23 X wider than MOQA; clypeus 0.42 high, 1.04 X
diameter of AME. Chelicerae: 3.06 long, 1.44 wide, 2.13 X longer than wide. Sternum:
2.63 long, 2.63 wide between coxae II, 1.00 X longer than wide and 2.31 wide between
coxae I. Labium: 1.32 wide, 0.80 long, 1.65 X wider than long, occupying 0.57 X of SW
Epigynum: Widely separated posterior excavated lobes of median septum of
epigynum twisted dorsally. Sperrnathecae short, inserted on heavily sclerotised,
bulbous invaginations of median septum, and directed anteriorly.
Legs: Leg spination not as irregular as in male: left and right femora I-III(3:2:3),
left femur IV(3:3:2), right femur IV(3:4:2 - due to a small ancilliary spine rising
close to base of one of dorsal spines). Tibiae dorsally, both sides, I-IV with single,
distal spine. Measurements:
Palpus I II III IV
Femur 2.13 7.13 7.63 6.25 7.88
Patella 1.25 3.13 3.25 2.75 2.75
Tibia 1.50 6.50 6.63 5.38 6.38
Metatarsus 6.00 6.00 4.25 5.88
Tarsus 2.88 2.00 1.88 1.63 1.75
Total: 7.76 24.76 25.39 20.26 24.64

Parapalystes cultrifer (Pocock, 1900), comb. D.

Palystes eultrifer Pocock, 1900: 331, PI. III, Figs 12, 12a. Syntypes: adult d [here designated as
lectotype] [examined], adult <;1 [inferred] [here designated as paralectotype] [not examined],
'Grahamstown', Mr Tuck (BMNH 99.7.24.72-73).
Remarks: Pocock (1900: 331) briefly described P. cultrifer, which he had received
from Dr SchOnland in Grahamstown, illustrating the two tibial apophyses (PI III, Fig.
12) and also a epigynum (P] III, Fig. t2a). Although he did not refer to a female, the
epigynum depicted clearly shows that he also saw an adult female, and therefore the
male and female are syntypes. A male labelled as the type of P. cultrifer, and matching
the description and illustration, and from the locality referred to by Pocock, is in
BMNH and has been designated by me as the lectotype. Another specimen in BMNH
found with the lectotype, a subadult female labelled 'Tea Fount. Grahamstown / Miss
Leppan [handwritten in pencil]' and '01.3.13.258 [handwritten in pencil]', is not the
female syntype (which is adult and which would probably have been accessioned
before Pocock's description was published), although it belongs to Parapalystes and
possibly to Pocock's P. cultrifer. When the adult female is found, it should be labelled
as the paralectotype.
Parapalystes lycosinus (Pocock, 1900), comb. D.
Palystes lycosinus Pocock, 1900: 330. Syntypes: adult d [here designated as lectotype) and adult d
subadult d adult 9 [here designated as paraiectotypesj, 'Port Elizabeth', H. A. Spencer
(BMNH 1890.7.18.47-49) [examined) and adult <;1 paraiectotype [here designated) [not found).
Palystes megaeephalus sensu Pocock, 1896: 63 [nee Ocypete megaeepahlus C. Koch, 1845
(misidentification)).
Remarks: Pocock (1896) identified a male and female from Port Elizabeth (eastern
Cape) as Ocypete megacephalus C. Koch, transferring Koch's species to Palystes. He
noted, however, that the species was possibly generically distinct from other South
African species of Palystes. Later Pocock (1900), recognising a misidentification,
distinguished his species from that of Koch's megacephalus, and described it under
the new name lycosinus. It is a moot point whether lycosinus should be considered a
junior synonym of megacephalus sensu Pocock. In the absence of Koch's type, and
with insufficient information in Koch's description to enable the species to be
identified, Pocock's name lycosinus, which is clearly identifiable, is retained.
The vial with Pocock's types of Palystes lycosinus contains two adult males and one
subadult male. The label, handwritten in pencil, bears the following information:
'1890.7.18.47-49' 'Palystes lycosinus' 'Pocock TYPE' 'Pt. Elizabeth' 'H.A. Spencer'.
Since the label data agree with the locality cited by Pocock and the specimens with
Pocock's description, I have designated the larger male as lectotype and the other two
specimens as paralectotypes. The adult female, also described by Pocock, was not found
with the type material but, and if found, should be labelled as the third paralectotype.
Parapalystes megacephalus (C. Koch, 1845), comb. D.
Oeypete megaeephala C. Koch, 1845: 25, PI. CCCCIII, Fig. 972. Holotype: 9, 'Das Vorgebirg der
guten Hoffnung' [= Cape of Good Hope], 'Konig!. Sammlung in Munchen' [type not located
in ZSMC, presumably lost).
Olios (?) [sic] megaeephalus; Simon 1880: 298 (tentatively transferred to Olios).
Palystes megacephalus; Pocock 1896: 63.

Remmius megacephal[usJ; Simon 1897a: 6, footnote 1 (transferred to Remmius).
Remarks: Ocypete C. Koch, 1836, erected for two South American heteropodids,
was pre-occupied by Ocypete Leach, 1815 (Coleoptera). Although Koch (1845:
27) realised this, he retained the generic name. Koch's (1845: 25-27) description
of a female from the Cape Province contains a number of references to characters
which clearly place his species in Parapalystes: domed head region of
cephalothorax; median oval white marking posterior to dorsal median line and
immediately anterior to thoracic groove of cephalothorax; black sternum with
posterior yellow region containing two black spots; dark, leopard-like spots on
ventral surface of coxae. Pocock (1896) transferred Ocypete megacephala C. Koch
to Palystes and identified two specimens from Port Elizabeth as such. Later,
Pocock (1900) distinguished Palystes megacephalus sensu Pocock, 1896 (now
transferred to Panaretella Lawrence) from Palystes megacephalus sensu Koch, but
retained Koch's species in Palystes. Simon (l897a) did not agree with Pocock's
transferral of Ocypete megacephala to Palystes, and instead transferred it to
Remmius Simon, 1897, a new east African genus. Simon's opinion was unjustified
in view of Koch's description. Koch clearly stated that the anterior lateral eyes are
very large, far larger than the anterior median eyes. In Simon's account of
Remmius (1 897a) the anterior median eyes are described as larger than the anterior
lateral eyes. While Koch's megacephala belongs in Parapalaystes, its identity
cannot be fixed as the description is inadequate and the type cannot be found in
Munich where a large part of the alcohol collection was destroyed during the
second world War (Baehr of ZMSC, pers. comm.), and it has not been located
elsewhere.
Parapalystes whiteae (Pocock, 1902), comb. D.

Palystes whiteae Pocock, 1902: 24. Holotype: adult d, 'Brak Kloof, near Grahamstown', Mrs White
[BMNH 01.3.5.69] [examined].
Remarks: Pocock (1902: 24) remarked on the close relationship between P. whiteae,
P. cultrifer and P. lycosinus. The latter two species have also been transferred to
Parapalystes. The type locality near Grahamstown is in the eastern Cape, South
Africa.
Panaretella Lawrence, 1937

Paranetella [sic] Lawrence, 1937: 249. Type species (by original designation): Panaretus (1) distictus
Pocock. 1896: 61.
PanareteUa; Lawrence, 1952: 210 [emendation of typographical error].
Remarks: The genus is clearly defined and distinct from Palystes. The anterior lateral
eyes are only slightly larger than the equally-sized anterior median and posterior
eyes, the posterior eye row is recurved and much (1.6 X) wider than the anterior eye
row, the labium is obliquely truncate and has a double notch on either side, and the
sternum is raised mesally at the labium. The markings of both female and male are
distinctive: sternum and coxae ventrally with numerous, very small, black spots and a
characteristic, relatively large black mark posteriorly on either side of the lower
abdomen. The epigynum and male palp are also distinctive and clearly illustrated by
Lawrence (1938: 250, Figs 20a-b; 1952: 209, Figs 51-56).
The genus, which now comprises five nominal species, appears to be endemic to
the eastern half of southern Africa and is most often collected in indigenous valley,
montane and coastal forest. The adults are small, nearly half the size of Palystes,
despite the number of species described, little is known about the natural history of
any member of the genus. One species is transferred from Palystes to Panaretella
(see below) and an annotated list of the presently recognised species is provided.
Specific synonymies are not established as these should be dealt with in a revision.
Panaretella disticta (Pocock, 1896)

Panaretus (?) distictus Pocock, 1896: 61, PI. VIII, Figs 7, 7a. Holotype: adult <;>, 'East London'
(BMNH) lexamined].
Palystes distinctus [sic]; Simon 1897a: 65, footnote 1 (regarded as probably an abnonnal Palystes).
Paranetus [sic] distinctus [sic]; Lawrence 1937: 249 (designated as type species of Panaretella).
Panaretella disticta; Lawrence 1952: 211 (emendation of his (1937) and of Simon's (l897a)
typographical errors).
Remarks: Although the generic and specific names of the type species appear to have
been bedevilled with typographical errors in the literature, the identity of both genus
and type species are clearly established. Pocock's type, an adult female, is labelled
'Po distictus, Poe. Type / E.London. H.A.Spencer'. It clearly belongs in Panaretella
with its distinctive markings, genitalia, dentition and leg spination.
Panaretella immaculata Lawrence, 1952

Panaretella immaculata Lawrence, 1952: 211, figs 51-53. Syntypes: adult d, adult <;>, 'Mseleni,
Zululand' (NMSA) [examined].
PanareteLLa minor Lawrence, 1952

Panaretella minor Lawrence, 1952: 212, figs 54-55. Syntypes: 1 adult <;> ,2 adult d, 'Pietennaritzburg'
(NMSA) [examined].
PanareteLLa scutata (Pocock, 1902), comb. n.

Palystes scutatus Pocock, 1902: 24, PI. III, Fig. 13. Holotype: adult <;> , 'Brak Kloof, Grahamstown'
(BMNH) [examined].
Panaretella zuluana Lawrence, 1937

Paranetella [sic] zuluana Lawrence, 1937: 249, figs 20a-b. Syntypes: adult d9 , 'Hluhluwe Game
Reserve, Zululand' (NMSA) [examined].
Panaretella zuluana; Lawrence 1952: 209 (emendation of spelling).
Anchonastus Simon, 1898

Anchonastus Simon, 1898: 203. Type species (by original designation): Anchonastus caudatus Simon,
1898: 203, footnote 1.
Palystodes Pocock, 1899b: 879. Type species (by monotypy): Palystodes plumosus Pocock, 1899b: 879;
Simon, 1903: 1029 (synonymy with Anchonastus).
Remarks: When Simon (1898) described this west and central African genus with its
unusual autapomorphies (sunken cephalothorax, plumose tufts of setae on legs,

abdomen and carapace, and unusually long posterior spinnerets), he placed it in his
subfamily Staianinae. The subfamily was, he believed, intermediate between Palystes
and Sparianthis Simon, 1880, from South America. The affinity with Palystes was
also recognised by Pocock (1899b: 879) in the genus Palystodes, later synonymised
with Anchonastus by Simon (1903: 1029). The following nominal species, with the
addition of a further species transferred from Palystes, are recognised in
Anchonastus. Specific synonymies are not established here, but should be dealt with
in a revision of the genus.
Anchonastus caudatus Simon, 1898

Anchonastus caudatus Simon, 1898: 203, footnote 1, Figs 204--206, p. 202. Ho!otype: d, 'Cameron'
[Cameroon, Africa] (MNHN) [examined].
Anchonastus gertschi Lessert, 1946

Anchonastus gertschi Lessert, 1946: 207, Figs 5-7. Ho!otype (by original designation): d, 'Kivu: Katana'
[Katana, SW. Lake Kivu, Zaire] (?Geneva) [not examined].
Anchonastus pilipodus (Strand, 1913), comb. n.

Palystes pi/ipodus Strand, 1913: 403. Syntypes: 19 , 'Kiwu-See, Nordufer, Kissenji', and 2d , 'Inse!
Kwidschwi (Kiwu-See), [northern shores, Lake Kivu, Zaire] (?MNZM) lnot examined).
Remarks: I have no misgivings in assigning Palystes pilipodus to Anchonastus.
Strand's description covers all the important characters that distinguish this very
distinctive genus. I have seen other Anchonastus specimens collected in the vicinity
of the type locality, which match Strand's description. I suspect that Palystes
pilipodus is a junior synonym of Anchonastus plumosus, as the descriptions match
well.
Anchonastus plumosus (Pocock, 1899)

Palystodes plumosus Pocock, I 899b: 879, PI. LVIII, Figs 31, 31a. Syntypes: adult d, adult 9 , 'Benito
River' [Zaire] (BMNH) [examined].
Anchonastus plumosus; Simon, 1903: 1029 (transferred to Anchonastus).
CLADISTICS OF PALYSTINAE AND PALYSTES

Data were analysed using Hennig86 (version 1.5 © James S. Farris, 1988). Two
cladograms, one for Palystinae (Fig. 116) and one for Palystes (Fig. 117), are
presented and discussed separately below. In the data matrices, 0 represents a
character in the ground state. Other digits represent different character states which
treated additively in the analyses (as sequential derivations) unless otherwise
indicated in the discussion of characters below.
Cladogram for Palystinae

Polarisation of characters was rooted in terms of a hypothetical ancestor with
characters in zero state based on comparison with other heteropodids, in particular
the apparent sister group to Palystinae, the Heteropodinae. Apart from close
correspondence in eye proportions and arrangements between Palystinae and

Heteropodinae, particularly in those heteropodines in which the eyes of the PER are
equal in size and sessile (as in Gnathopalystes), the two subfamilies also share basic
leg spination patterns and the same tarsal slit sense organs (Fig. 15). Further SEM
work is necessary to test whether the latter is a potential synapomorphy for the two
subfamilies. The placement of a number of heteropodine species under Palystinae by
Simon further strengthens the suggestion that the two subfamilies may be closely
related.
The polarisation of characters, data set used for analysis, and comments appear
below:
1. Carapace flattening:
In Heteropodinae and Panaretella the carapace is domed and the thoracic fovea
lies on the posterior declivity. Carapace convex with thoracic fovea sloping
downwards on posterior declivity = O. Cephalic region domed, thoracic region
flattened with thoracic fovea horizontal to body plane = 1. Cephalic and thoracic
regions flat = 2. [Non Additive.]
2. Dorsal spination of tibiae I-IV:
In Heteropodinae, which are sexually dimorphic, females have a single spine but
males, which have longer legs, have two as do both sexes of Panaretella and
Palystes. Tibiae with 2 dorsal spines = O. Tibiae with 1 dorsal spine = 1.
3. Ventral markings of tibiae:
Heteropodids examined, including Heteropodinae and Panaretella, do not have
distinctive markings ventrally on tibiae I-IV. Tibiae ventrally brindled or dark
without distinctive markings = O. Tibiae ventrally with two broad white to yellow
bands, one medially and one distally, against a dark to black background = 1.
4. Cheliceral markings:
The two longitudinal white stripes at the outer margins of the chelicerae found in
Palystes and Parapalystes are a derived state as they were not found in other
heteropodids examined. Chelicerae dark without clearly defined markings = O.
Chelicerae with two fine white or yellow longitudinal lines at their outer margins
=1.
5. Clypeus:
The white clypeal 'moustache' found in Palystes and Parapalystes appears to be
a derived state as it was not found in any other heteropodids examined. Clypeus
of general body colour (brown to yellowish brown), setae unmodified = O.
Clypeus with a distinct white or yellow 'moustache' of modified, closely
adpressed setae = 1.
6. Posterior spinneret length:
The very long posterior spinnerets found in Anchonastus are a derived state and
have not been seen in other heteropodids examined. Base similar in length to
base of anterior spinnerets = O. Base considerably longer (more than 2 X) than
base of anterior spinnerets = 1.
7. Median septum of epigynum:
The typical pattern in many of the larger heteropodids, including Heteropodinae
and the Madagascan genera Damastes Simon, 1880 and Megaloremmius Simon,
1903, is that, posteriorly, the lateral sclerites of the epigynum are produced and
meet medially or are narrowly separated. Reduction of the lateral sclerites and
exposure and development of the median and posterior areas of the septum is
considered derived. Septum posteriorly covered, or only narrowly exposed, by
well-developed lateral sclerites of epigynum =O. Septum broadly exposed, well
developed, lateral sclerites reduced = 1. Septum very broad, overlapping lateral
sclerites =2.
8. Male tibial apophysis:
A single, spur-like or digitform tibial apophysis is common to most
heteropodids, including Heteropodinae and Palystes. The unusual compound
apophyses in Panaretella, Anchonastus and Parapalystes suggest that this is a
derived state. Single apophysis = O. Compound apophyses = 1.
9. Embolar supporting sclerite:
Most heteropodids examined, including Heteropodinae, do not appear to have a
supporting lateral sclerite (Fig 9a) to the embolus. The presence of a well-
developed lateral sclerite in Panaretella, Anchonastus and Parapalystes, and also
in the castaneus species group, suggests that this is a derivation for Palystinae. In
the lunatus group the embolus is a spur-like hook. Embolar supporting sclerite
absent = O. Embolus subtended by well-developed supporting sclerite = 1.
Supporting sclerite a spur or low ridge at the base of the embolus = 2. [Non
Additive.]
10. Embolus form:
Embolus form in heteropodids is variable but in Heteropodinae has become a
progressively elongate, whip-like spur subtended by an elongate, spathe-like
conductor. Development of this form of thin, whip-like embolus has not occurred
in Palystinae. For outgroup comparison, a starting state is postulated for the
embolus form of a shorter spur-like, distally tapered embolus as found in
Panaretella, Anchonastus and Parapalystes. Blade-like flattening of the embolus
apically is therefore considered derived. Embolus a stout spur, tapering distally =
O. Embolus apically flattened, blade-like = 1.
11. Labium width:
In Heteropodinae, Panaretella and Anchonastus the labium is longer than wide
or only slightly wider than long, the lateral notches are elongate and the sides
taper apically compared with the broad, parallel-sided labium, with short basal
notches laterally, found in Palystes and Parapalystes. Labium longer than wide
or only slightly wider than long, sides steeply inclined anteriorly, narrowly
truncate apically =O. Labium much wider than long (1.2-1.6 X), sides parallel to
=
slightly inclined anteriorly, broadly truncate apically 1.
12. Spermathecal duct length:
The duct leading to the spermatheca is coiled and becomes increasingly coiled
with progressive length of the embolus in Heteropodinae and Deleninae (see
remarks for character 10). Coiling of this nature is not found in Palystinae which
have short ducts, except for the lunatus and superciliosus groups. Increasing
length of the ducts is considered derived. Duct leading to spermatheca short, less
than 3 X longer than wide =O. Duct moderately to very elongate, more than 5 X
longer than wide = 1.
13. Coxal spotting:
Spot markings of the coxae were not found in other heteropodids examined, and
the presence of different kinds of coxal markings is considered independently
derived. Coxae without spot markings = O. Coxae speckled with fine spots = 1.
Coxae with up to five large spots =2. [Non Additive.]
14. Abdominal ventral bar marking:
A variety of ventral abdominal markings, or no markings, are found in
Heteropodinae but none examined with a distinct, transverse, dark to black bar
immediately posterior to the epigastric furrow (Fig. 95. The marking is not
present in Panaretella but its presence in the rest of the Palystinae is considered
derived. No distinct bar marking = O. Bar marking posterior to epigastric groove
=1.
15. Bell-shaped marking of venter:
Shield-like solid or divided dark to black markings of the venter are found in
some Heteropodinae, Micrommatinae (Cerbalus and Cercetius) and Palystinae.
Within Palystinae the black shield marking may be present, absent or replaced by
a red marking or by a disintegrating brown marking. These various character
states are consistent within groups but considered independently derived. Solid
or divided black marking = O. Solid red-brown to red marking = 1.
Disintegrating dark brown marking =2. No marking =3. [Non Additive.]
16. Sternum markings:
Dark transverse bar markings of the sternum occur in one of the Australian
heteropodids (Heteropoda eungeUa Todd-Davies, 1994), but the remainder have
an unmarked, normal body colour sternum. Panaratella has a normal body
colour sternum, although the bases of sternal setae are dark. The rest of the
Palystinae all have sternum markings although these are different from group to
group. The sternum may be coarsely barred to mostly black with some body
colour (Parapalystes and Anchonastus); completely black (castaneus group); or
with one or more fine transverse black bars (the lunatus and superciliosus
groups). The plesiomorphic state may either be an unpigmented sternum of
normal body colour (with barring arising independently) or a sternum with bar
markings. Since until now barring has only been observed in one heteropodine
species and does not occur in Panaretella, a normal body colour sternum is
considered plesiomorphic and the various character markings are treated as
independently derived states. Sternum normal body colour =O. Sternum coarsely
barred to mostly black = 1. Sternum completely black =2. Sternum with fme bar
markings =3. [Non Additive.]
17. Retrolateral abdominal black patch marking:
Lateral black patch markings of the abdomen are absent in Heteropodinae and
Palystinae, except for Panaretella. Absent =O. Present = 1.
18. White markings of abdomen:
Various patterns of small white fleck or spot markings of the abdomen are found
in individual species or species groups in both Heteropodinae and Palystinae.
Large white markings of the abdomen appear in two distinct forms in two of the
Palystes groups, but have not been observed in other heteropodids including
Heteropodinae and are considered independently derived. White markings in the
form of small spots or flecks, or no white markings. = O. Large white spots
ventrally = 1. Broad white band laterally =2. [Non Additive.]
19. Position of AME:
In Heteropodinae the PER appears procurved because the AME are situated high
on the carapace, and tilted strongly upwards, with their upper rims on a common
line with those of the large ALE. The lower position of the AME in the
Palystinae and their orientation (directed forwards and only slightly tilted
upwards) appear to be a derived state for Palystinae. PER procurved with upper
rims of AME on a common line with upper rims of ALE, AME strongly tilted
upwards = O. PER straight with AME on a common median line with ALE,
AME directed forwards, or only slightly tilted upwards = 1.
20. Cheliceral dentition:
Heteropodinae have a 4:3 cheliceral dentition pattern (four teeth on the
retromargin and three on the anteromargin). Eusparassinae (e.g. Damastes
Simon, 1880) Deleninae and Micrommatinae (e.g. Megaloremmius, Cebrennus
Simon, 1880 and Cercetius Simon, 1902) have 4:2 cheliceral dentition. Although
the relationships of the larger heteropodids have still to be fully investigated, it is
probable that four retromarginal teeth is a plesiomorphic state. The 3:3 cheliceral
dentition and proportions and shape of teeth found in all Palystinae appears to be
a derived state for the group. Retromargin with four teeth, basal smallest;
anteromargin with two or three elongate teeth = o. Retromargin with three equal
sized teeth, anteromargin with three triangular teeth, median largest = 1.
Character matrixfor Palystinae
Ancestor 00000 00000 00000 00000

Panaretella 00000 00110 00103 01011
Anchonastus 21100 10110 00011 10011
Parapalystes 01111 02110 10210 10011
Palystes castaneus group 10111 01011 10010 20011
P. lunatus group 10111 01021 11011 30111
P. superciliosus group 10111 01001 11012 30211
The single tree derived for 20 characters in the six groups required 33 steps
(consistency index 87, retention index 81).
The cladogram for Palystinae (Fig. 116) includes Palystes divided into three
intuitively recognised groups (the castaneus, lunatus and superciliosus groups) based
on observed patterns of markings and male and female genitalia.The species analysis
(Fig. 117), however, only confirmed two groups (the lunatus and superciliosus
groups) as being holophyletic, and indicated that the castaneus group was
paraphyletic. The apomorphy indicated for the castaneus group (character 16, state
2) is based on a non additive character state (common possession of a completely
0..
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9~2 9~O
15~1 15~2
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6~1 7~1
15~1 11~1
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Fig. 116. Cladogram proposed for Palystinae. Characters and their states indicated on the diagram are
discussed in the cladistics section. Solid filled squares indicate uniquely derived apomorphic
character states; empty squares, reversals to the plesiomorphic state; diagonals, parallel
derivations of apomorphic states.
black sternum) which was treated as a plesiomorphic character state (character 18,
state 0, black to mostly black) in the species analysis.
Cladogram for Palystes

Polarisation of characters in the cladistic analysis of Palystes (Fig. 117) was rooted
in terms of a hypothetical ancestor with characters in zero state based on outgroup
comparison with its sister genus, Parapalystes, as determined in the cladogram for
Palystinae. The two genera share as synapomorphies the markings of the chelicerae
and clypeus and shape of labium. Unless otherwise indicated, the plesiomorphic state
is present in the outgroup.
For two species, P. lunatus and P. leroyorum, only one sex is known and query
marks were inserted in the data set for sex-linked characters for the missing sexes.
The polarisation of characters, data set used for analysis, and comments appear
below:
1. Anterior epigynal septum:
In Parapalystes and in the castaneus group the epigynum fonns a broad, flat
shield anteriorly. Progressive deepening of the anterior excavation of the septum
occurs in the lunatus and superciliosus groups. Flat anteriorly = O. Deeply
excavated anteriorly =1.
2. Medial protuberance of epigynaJ septum:
In Parapalystes the septum has lateral protuberances, but is flat medially (Fig.
110), without the derived medial protuberances found in the lunatus (Fig. 37)
and supercilosus (Fig. 77) groups. Septum flat medially, no protuberance = O.
Medial 'tongue' -like protuberance overhanging septum posteriorly = 1. Medial
protuberance contracted into 'bow-tie' shape, posterior septum retracted = 2.
[Non Additive].
3. Extension of posterior margin of 'bow-tie' shaped septum protuberance:
Posterior extension of the posterior margin of the septum is not found in Palystes
except in the superciliosus group where its presence is considered derived.
Within the group the extended margin may be present (Figs 81, 89,97, 104), or
reduced (Fig. 77). Posterior extension of protuberance absent = O. Posterior
margin with fully-developed posterior extension = 1. Posterior extension reduced
=2.
4. Development of 'tongue' -like medial protuberance of septum:
The development of a 'tongue' -like medial protuberance of the septum is absent
in Palystes except for the lunatus group where its presence is considered
derived. 'Tongue-like' protuberance absent or not produced posteriorly = O.
Produced by as much as width = 1. Produced by more than twice width = 2.
5. Retrolateral excavations of epigynal septum:
In Parapalystes and in P. castaneus, the septum is an excavated, flat shield with
widely separated bulbous prominences retrolaterally (Figs 8a, 8b, 110).
Reduction of the lateral prominences appears to be a derived state. Broad septum
with widely separated retrolateral bulbous excavations = O. Septum greatly
reduced retrolaterally, without widely separated bulbous excavations = 1.
6. Spennathecal duct length:
In Parapalystes and the castaneus group, ducts leading to the spermathecae are
short. Their elongation is considered derived. Spennathecal afferent ducts short,
less than 3 X longer than wide = O. Spermathecal ducts moderately to very
elongate, more than 5 X longer than wide = 1.
7. Position of spermathecae in relation to support collar:
In Parapalystes and P. castaneus the spermathecae have short ducts and lie on
the invagination of the lateral margins of the septum (Figs 8b, 111). In P.
martinfilmeri, P. stilleri (Figs 24, 33) and the lunatus species group, the ducts are
longer and the spermathecae lie lateral to a rudimentary 'collar'. In the
superciliosus group the ducts are very much longer, and extend below the flange
of the fully developed collar (Figs 64, 90) which is formed medially by the
anterior invagination of the septum. In the superciliosus group the afferent ducts
may be hidden from view when viewed dorsally (Fig. 98), or extend below the
collar and become visible internal to the collar (Figs 82, 90, 105), or even extend
below, rising internal to the collar lobes and lying on top of them (Fig. 78).
Spermathecae lying lateral to support collar, or no support collar = O.
Spermathecae lying ventrally under support collar = 1. Spermathecae rise
internally from below collar = 2. Spermathecae rise internally from below collar
to lie dorsally on collar =3.
8. Orientation of spermathecae lateral to support collar:
Lateral insertion of the spermathecae on a support collar, when present, appears
to be derived in the lunatus group. Spermathecae not inserted lateral to support
collar = O. Inserted laterally, produced medially = 1. Inserted medially, produced
laterally = 2. Inserted medially, produced anteriorly = 3. Inserted laterally,
produced anteriorly =4. [Non Additive.]
9. Flattening of embolus:
The apex of the embolus is a stout, apically tapered spur in Parapalystes and is
elongate, rounded and whiplike in Heterodinae. The flattening of the apex, either
vertically or horizontally, is found in all Palystes species and is considered
derived. Embolus not flattened apically = O. Embolus flattened apically on a
vertical plane to the palp = 1. Embolus flattened on a plane horizontal to the palp
= 2. [Non Additive.]
10. Carination of embolus:
Lateral carination of the apex of the embolus, absent in Palystinae except for the
superciliosus group, differs in form. Progressive reduction of the carination is
considered derived. No lateral carinations apically"; O. Heavy distal carination
crossing on to ventral surface, anterior to proximal carination = 1. Light distal
carination not crossing on to ventral surface, opposite proximal carination = 2.
Carination greatly reduced and elongate = 3. Carination vestigial or absent,
embolus greatly reduced in width apically, acicular =4.
11. Embolus reflexion:
In Parapalystes and P. castaneus the embolus is straight. Increasing reflexion of
the embolus is derived. Embolus straight = O. Embolus reflexed, but less than
1200 = 1. Embolus reflexed, more than 1800 =2.
12. Conductor base:
An elongate conductor base is found in Parapalystes and P. castaneus.
Reduction to a sessile conductor, present in all other members of Palystes,
appears to be the derived state. Conductor base elongate = O. Conductor sessile =
1.
13. Embolus channel:
Channeling of the embolus is absent in P. castaneus and Parapalystes and its
presence in the remaining members of Palystes is considered derived. Embolus

without a channel = O. Embolus with a broad apical channel, but the channel not
extending to the base of the embolus = 1. Embolus with channel extending to the
base = 2. [Non Additive.]
14. Lateral selerite adjacent to embolus:
A well-developed lateral selerite is found in Palystinae and the castaneus group.
Its reduction in the lunatus group and loss in the superciliosus group are
considered derived. Embolus subtended by a well-developed lateral selerite = O.
Supporting selerite reduced = 1. Supporting selerite absent = 2.
15. Shape of reduced lateral selerite of embolus:
Remarks for character 14 above apply. Progressive reduction of the lateral
selerite within the lunatus group is considered derived. Lateral selerite well-
developed or absent = O. Reduced to long spur more than half length of embolus
(Fig. 73) = 1. Reduced to short 'rose thorn' spine at base of embolus (Fig. 50) =
2. Reduced to low ridge at base of embolus (Fig. 42) = 3.
16. Male pedipalp tibial apophyses:
Plesiomorphic within Palystinae is the presence of several complex tibial
apophyses, found in Panaratella, Anchonastus and Parapalystes. Several ridges
and two, separate, spurs are present in Parapalystes (Fig. 113). Reduction to a
single apophysis is characteristic of Palystes. Two apophyses = O. Reduction to a
single apophysis = 1.
17. Male pedipalp tibial apophysis shape:
A complex apophysis with various lobes or projections is characteristic of
Palystinae. Steady reduction within Palystes to a light entire spur is considered
derived. Single, large, apophysis with several digitiform projections (Fig. 9b)= O.
Apophysis reduced to a single, large, ridged, digitiform spur as long as embolus
(Figs 25, 34) = 1. Apophysis reduced (less than half length of embolus), ridged,
apically flattened and notched (Fig. 44), forked (Fig. 56) or entire (Fig. 66) = 2.
Apophysis reduced to a light, unridged, un flattened, rounded, apically tapered
spur (Figs 91, 102) = 3.
18. Sternum markings:
The sternum is mostly black in Parapalystes (Fig. 115) to completely black in
the castaneus group (Fig. 3). Variable fine black barring with one or more fine
transverse black bars (the lunatus and superci/iosus groups) is considered
derived. Sternum mostly black to completely black = O. Sternum with fine bar
markings = 1.
19. Abdominal ventral bell-shaped marking posterior to ventral bar marking:
Parapalystes and the castaneus group have a solid or divided black marking
ventrally on the abdomen. Red or brown markings are derived. Solid or divided
black marking = O. Solid red-brown to red marking = 1. Disintegrating brown
marking = 2. Disintegrating black marking = 3. Grey marking = 4. [Non
Additive.]
20. White markings of abdomen:
Clear white abdominal markings are not found in Palystinae with the exception
of the lunatus and superciliosus groups. White abdominal markings in the form
of large white spots ventrally (found in the lunatus group) or as lateral white
bands (in the superciliosus group) are considered derived. No white markings or
only some white flecking laterally = O. Large white spots laterally and ventrally
= 1. White spots coalescing into a broad white marking laterally = 2. [Non
Additive.]
Character matrix for Palystes
Ancestor 00000 00000 00000 00000

P. castaneus 00000 00010 00000 00000
P. martinfilmeri 00000 00110 11000 11000
P. stilleri 00001 00110 11200 11000
P.lunatus 11021 102?? ????? ??111
P. crawshayi 11001 10410 11113 12111
P. peromatus 11021 10210 11112 12111
P.leppanae 11011 10310 11213 12111
P. karooensis 11011 10210 11112 12111
P. stuarti 11011 10410 11111 12111
P. johnstoni 12101 11021 21020 13122
P. ellioti 12101 12023 21020 13122
P. hoehneli 12101 12023 21020 13122
P. ansiedippenaarae 12101 12022 21020 13122
P. superciliosus 12201 13024 21020 13142
P. leroyorum ????? ???24 21020 13132
Analysis with Hennig86 produced two equally parsimonious trees with a length of
47 steps (consistency index 93, retention index 97), differing only in one branch in
the castaneus group. The preferred tree (Fig. 117) unites P. stilleri and P.
martinfilmeri instead of placing P. martinfilmeri one node lower, but still above P.
castaneus. This provides a more useful grouping. A strict consensus tree proved less
parsimonious and was therefore not used for the cladogram.
Two clear monophyletic sister groups united at node 0, the lunatus group united at
node N, and the superciliosus group united at node I, are evident in the diagram.
They reflect 'natura1' groups identified during the study. The third group, the
castaneus group, is paraphyletic and contains P. stilleri united with P. martinfilmeri
at node P, but also includes the third western Cape species, P. castaneus. These three
western Cape species are kept together for convenience. Although P. castaneus is
clearly a member of Palystes, it possesses a number of character states considered
plesiomorphic by comparison with the outgroup.
BIOGEOGRAPHY OF PALYSTINAE AND PALYSTES

The sequence of dichotomies suggested in the cladogram for Palystinae (Fig. 116)
starts with the separation of Panaretella, an inhabitant of coastal plain woodland and
low-lying montane forest of south-eastern Africa, followed by that of Anchonastus (a
tropical lowland forest genus of west and central Africa) and finally, the separation of
the population ancestral to Parapalystes (an arid-adapted genus of the interior of the
Cape Province of South Africa) and Palystes (a chiefly montane forest and open
woodland genus of the higher rainfall areas of eastern and southern Africa).
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Fig. 117. Cladogram proposed for Palystes. Characters and their states indicated on the diagram are
discussed in the cladistics section. Solid filled squares indicate uniquely derived apomorphic
character states; empty squares, reversals to the plesiomorphic state; diagonals, parallel
derivations of apomorphic states.
It is probable that the ancestor to Palystinae was an inhabitant of tropical lowland

forest similar to the habitat in which both Anchonastus and Panaretella are found
today. Separation of the lowland forests of west and east Africa took place during the
middle to late Miocene with the accelerated development of the east African ups well
and the subsequent orogenesis, rifting and plateaux uplifting that transformed the
central and east African landscape (Laporte & Zihlman 1983). Until that time
equatorial Africa was dominated by a broad transverse band of tropical lowland

equatorial forest - a remnant of the ancient, pre-Miocene Tethyan marginal tropical
flora. The new highland and volcanic mountain spine that divided west and east
Africa resulted in the drying of the eastern third of central Africa, cut off from the
moist western Atlantic Trade winds. Three longitudinal vegetational bands,
extending into south-eastern Africa, developed down the length of eastern Africa: a
new montane tropical forest mingled with remnants of an older temperate forest
(White 1978); a drier highland open savanna woodland; and a narrow band of coastal
plain forest.
If the initial fragmentation within the Palystinae is related to this period, then it
appears probable that Panaretella was isolated in the remaining lowland forest strip
along the east African coastal plain. The population ancestral to Anchonastus left in
the still extensive lowland forest in west Africa then being affected in the east and
south-east by further changes during the mid-Miocene, gave rise to a popUlation
adapted to the high-lying open woodland (the population ancestral to Parapalystes
and Palystes). Adaptation to a subtropical, cooler climate, would have enabled the
southwards dispersal of the population into southern Africa. Axelrod & Raven (1978)
have suggested that there was a general distribution of forest in southern Africa
during the mid-Tertiary until the events of the late Miocene onwards. Already in
existence during the late Miocene were: an older established region of aridity or
semi-aridity along the south-western African coast (Tankard & Rogers 1978, Ward et
al. 1983); the old southern African Mesozoic eastern mountain escarpment (Tankard
& Rogers 1978) and an extensive south-western Cape Province flora of alternating
temperate and tropical-subtropical forests (Coetzee 1978 1980 1986, Hendey 1983b,
Scholtz 1985). Scholtz (1985) identified an early Tertiary (Palaeogene) open-
canopied, largely deciduous fairly dry forest flora at Banke in the northern Cape
interior including, as possible understorey elements (Hilton-Taylor 1987),
Proteaceae, Restionaceae and Ericaceae, which are precursors of the present day
winter rainfall fynbos flora of the southern Cape (Van Zinderen Bakker 1987).
The orographic and climatic changes of the late Miocene and Pliocene had a
marked influence on the vegetation of Africa as a whole (Axelrod & Raven 1978,
Coetzee 1981 1986) and the separation of the popUlations ancestral to Palystes,
Parapalystes and the subsequent fragmentation of Palystes into the three species
groups now recognised, may owe much to the resulting major habitat changes and
disjunctions of this period. The movement of land over the southern polar region and
the consequent development of the Antarctic ice-cap towards the end of the Miocene
appears to have had a major aridifying influence on the south-western coastlines of
southern America and southern Africa. This, and the development of a winter rainfall
regime in the south-western Cape Province of South Africa, appear to have seen the
replacement of forest and grassland with an evolving Cape macchia shrub and scrub
vegetation, accompanying increasing aridity in the south-western interior of Africa
during the Pliocene, reaching its climax during the Pleistocene (Van Zinderen Bakker
1975, Hendey 1983b, Ward et al. 1983, Tyson 1986, Hilton-Taylor 1987). With
increasing aridity in the interior of the Cape, elements of the fynbos, in tum, appear
to have given rise to the karroid scrub and succulent vegetation that today blankets
much of the Cape interior (Hilton-Taylor 1987, Van Zinderen Bakker 1987).
It is probable that the population ancestral to Parapalystes, having earlier adapted
to drier sub-tropical forests in the Cape interior, was isolated in the north-western
Cape Province during the late Miocene-early Pliocene and adapted still further to the
semi-arid conditions that developed as the forests were replaced by the karroid scrub
in which it now survives.
The 15 species recognised in Palystes are distributed in central, eastern and
southern Africa. Many of the species are endemic to relatively small regions. The
cladogram (Fig. 117) suggests two monophyletic species groups, the superciliosus
group (at node I), which extends from east to south-east Africa, and the lunatus
group (at node N), of the eastern Cape, Lesotho and Transkei. The three remaining
species, all from the south-western Cape, have for convenience been placed in a
species group, the castaneus group, despite cladistic analysis revealing that the group
is probably paraphyletic.
The cladogram suggests several, successive, isolation events, the oldest taking
place in the southern Cape, isolating the popUlation ancestral to P. castaneus (node
R).
That isolation of the ancestor to P. castaneus took place first is suggested by the
large number of plesiomorphic character states found in P. castaneus - more than in
any other Palystes species. That a continuous, suitable, habitat between the
Cederberg Mountains (where P. martinfilmeri is found) and the Stellenbosch
mountains (type locality of P. stilleri) persisted for some time after the isolation of P.
castaneus, is implied by the closer correspondence in character states between the
two species (united at node P) than either with P. castaneus. A major Pliocene marine
incursion, lasting some 1.5 million years, from the early Pliocene to the end of the
mid-Pliocene, saw sea levels off the southernmost African coast reach levels some
100 m higher than at present (Hendey 1983a). These high sea stands would have
effectively isolated the Table Mountain range from the mainland and may have been
responsible for this first fragmentation. Increasingly xeric conditions further north
during the late Pliocene-Pleistocene saw the disappearance of lower lying forest
which gave way to macchia; this process had started by the end of the Miocene
(Coetzee 1981) and would have effectively isolated the popUlation ancestral to P.
martinfilmeri (now found in the montane fynbos and karoid scrub vegetation of the
Piketberg, Cederberg and Clanwilliam districts), from that of P. stilleri, which still
survives in the cool, well-watered forests and highlands of lonkershoek and the
Hottentots Holland Mountains.
The separation of the parent population to the remainder of the Palystes species
(united at node 0) from that of the castaneus group, may be related to the
development of an end-Miocene extensive arid belt stretching from the Algoa Bay-
Uitenhage basin northwestwards deep into the Cape interior. The belt is a transition
zone between the winter rainfall region of the western Cape and the summer rainfall
region of the eastern Cape. Griswold (1985), in his revision of the mygalomorph
spider family Microstigmatidae, suggested a maximum age of late Miocene-early
Pleistocene for an eastern Cape clade with three species of Microstigmata, possibly
affected by the changes of this period. Griswold (l987b) found a similar pattern of
vicariance in Migidae with separated centres of richness, one in the western Cape and
the other in the eastern Cape. Separation of the population ancestral to the lunatus
group in the eastern Cape from that of the population ancestral to the superciliosus
group may have taken place during the Pliocene-Pleistocene when, after a series of
uplifts which had started at the end of the Miocene, the Natal mountain escarpment
was elevated by more than 1 000 m. The consequence of this was probably the
disappearance of forest above I 000 m (Stuckenberg 1962). Lewis (1988) has
reviewed evidence supporting Sparrow's (1973) suggestion about the existence of a
Pleistocene periglacial above the 2 000 m contour in the Drakensberg-Lesotho
highlands.
The eastern Cape is an ecologically unstable region where no less than five major
vegetation units meet (Gibbs-Russell & Robinson 1981): Indian Ocean Coastal Belt,
Zambezian Domain, Afromontane, Karoo-Namib and Capensis phytochoria. Despite
the active spider collecting that has taken place in the eastern Cape since shortly
before the start of this century, several species in this clade appear to be restricted to
relatively small areas and are represented by only a handful of specimens (in the case
of P. lunatus, only the holotype female).
The first suggested isolation is that of P. stuarti in the western Cape (node N),
followed by P. crawshayi (of Lesotho) and P. leppanae (of Grahamstown) (node K).
The sequence of isolations affecting the population ancestral to P. karooensis, P.
lunatus and P. peromatus (united at node L), suggests that first there was a western,
Karoo interior, isolation of P. karooensis (today found in the Beaufort West and
Graaff-Reinet districts) followed by that of P. peromatus (found from Queenstown in
the eastern Cape Border region to the Umtamvuna forest in north-eastern Transkei)
and P. lunatus (presumably from the eastern Cape interior). The isolations may be
related to the prolonged drier conditions which have affected the Karoo and eastern
Cape since at least the Pliocene and particularly during the Pleistocene. Latterly, until
some 8000 years ago (Meadows et al. 1987), cooler and moister conditions may have
enabled P. karooensis and P. peromatus to extend their range into the north-eastern
Transkei area where outlier populations of both species are found.
The superciliosus group (united at node I) extended its range northwards into the
East African highlands. Today the species with the greatest number of plesiomorphic
character states within the group, P. johnstoni, is only found north of the Limpopo
River (in Zimbabwe and MalaWi). Despite the unresolved trichotomy at node H, the
relationships between the central and east African species (P. hoehneli and P. ellioti),
and the southern African species (P. ansiedippenaarae, P. superciliosus and P.
leroyorum) suggest one or more, Pleistocene or later, longitudinal African dispersal
routes.
The male and female genitalia of P. ansiedippenaarae in the eastern Transvaal are
very close in structure to those of P. hoehneli from the Mt Kilimanjaro and Mt Kenya
ranges in east Africa, despite the gap separating the two species. The northern
population may well have followed the retreat of forest up the Tanzanian mountain
slopes during the glacial maxima of the Quaternary. The present day populations of
P. hoehneli and P. ellioti are coincident with the distribution of some forest refugia
suggested for south-western Uganda, Rwanda and Burundi in the west and of
Kilimanjaro and the Usambara Mountains in the east during the Quaternary (Van
Zinderen Bakker 1967, Van Zinderen Bakker & Coetzee 1972, Diamond & Hamilton
1980). P ellioti is distributed both in central Africa as well as in the Usumbara
Mountains to the east, separated by P hoehneli in the mountains of Kenya and
Kilimanjaro. Its closest relatives are P superciliosus and P leroyorum in southern
Africa.
This would agree with a suggested alternately open woodland (during pluvial
conditions) and arid (interpluvial), maximum Pleistocene age, interior corridor
linking the north-east of Africa with central and western southern Africa. The
periodic aridity of the hypothesised corridor has been suggested as a possible
dispersal route for arid-adapted south-western groups with affinities in north-east
Africa. It is possible that it may also have served as a wet p~riod (pluvial) dispersal
route for open woodland inhabitants of east and central Africa which have close
relatives in southern Africa.
SUBFAMILY HETEROPODINAE
Remarks: The Heteropodinae, first recognised as a subfamily by Keyserling in 1880,
is a large group of more than 170 species in some 16 genera, mainly Indo-
Australasian, with several extra-limital members from Africa and one from South
America. The type genus, Heteropoda Latreille, 1804, with well over 100 attributed
species and some 13 subspecies from the Far East (mainly the Indo-Australasian
region but including China, Japan, Celebes, Sumatra and New Guinea), is based on
Heteropoda venatoria (Linnaeus, 1767). Bonnet (1951) extensively reviews the
problems associated with the type species and its name. A type specimen has still to
be established for Heteropoda venatoria, to confirm the identity of the genus. The
type species, however, appears to be well known, and specimens in SAMC, NMSA,
BMNH and MNHN determined under this name are congeneric, if not conspecific,
with the female described and illustrated as such by Jarvi (1912, 1914: 197) in the
Berlin Museum. For a more modem illustrated description of both the male and
female of Heteropoda venatoria, see Todd-Davies (1994). The type species appears
to be southern Asian in origin, but is now tropicopolitan, probably aided by the
unwitting agency of human transport, and is not infrequently collected in harbour
cities round the world. In Africa it has been found in Mozambique (NMSA), appears
well established in west Africa (Cameroon), and is also found occasionally in Cape
Town, South Africa (SAMC).
Simon's (1897a: 31) key to, and definition of, the subfamily based on eye size and
arrangement, and little else, has resulted in a number of genera being assigned
erroneously to the subfamily by subsequent authors. Simon (l897a) differentiates
Palystinae from Heteropodinae because in the latter the posterior lateral eyes are
raised and larger than the posterior medians (as opposed to sessile and
homogeneous). The relative size of the posterior laterals, which are secondary eyes,
is variable from genus to genus (and sometimes within genera). Of far more
significance is the position of the primary eyes, the anterior medians, which appear to
be consistent from genus to genus in related groups. In those genera I consider
belong in Heteropodinae, they are tilted upwards, with their upper rims on a line with
the upper rims of the anterior laterals. In the redefined Palystinae they are tilted more
anteriorly and lie lower on the caput face, on a common median line with the anterior
laterals. The relative size of the eyes has been used in too general a manner in the
past in defining heteropodid groups. It is better used within context and in
combination with other characters, such as those included in the suggested diagnosis
below.
The discovery and suggested significance, in a number of heteropodine genera, of
micro-denticles on the fang furrow floor of the chelicerae (between the larger teeth
on the margins), has been discussed with a number of colleagues including Roth and
Todd-Davies. Hirst (1989a) refers to these additional teeth as 'median teeth' and
Todd-Davies (1994) calls them 'denticles. Neither I nor Roth have observed these
denticles in heteropodid genera other than those listed below, despite our independent
examination of much of the type material of heteropodid genera in the considerable
collections of BMNH and MNHN. Denticles between cheliceral tooth rows do occur
in other spider families, from mygalomorphs and hypochiloids up through araneoids
(Platnick, pers. comm.). Whether there is anything unique about the formation of the
denticles in the heteropodid genera discussed here, and whether the genera with this
and the other characters discussed, will prove to be more closely related to each other
than to any other taxa, however, has still to be tested. The preliminary examination
does suggest that they are related and probably form a natural (holophyletic) group.
The presence of microdenticles in this particular group of heteropodids is either a
shared reversal to a plesiomorphic state or a synapomorphy; the latter is more likely.
Microdenticles of this type are not present in Madagascan, South American or
Australian species (other than Australian heteropodine species which appear to
belong with Asian and south-western Pacific island groups and which are probably
much later arrivals in Australia), nor in any of the African genera, except for the two
Asian heteropodine genera which have also probably recently arrived in Africa
(Torania and Heteropoda). This suggests that the character is indeed a post-
Gondwanan southern Asian synapomorphy of the heteropodine genera discussed
here and listed below.
The following African genera were assigned by their authors to Heteropodinae, but
do not have the characters discussed above and probably do not belong in the
subfamily:
• Panaretella Lawrence, 1937. This genus (see above under Palystinae) was
presumably placed in Heteropodinae because of Lawrence's comment (1937:
249): 'Pocock's species is certainly not a Palystes ... ; the genus is probably related
to Panaretus Simon.' Although I agree with Lawrence that Panaretella does not
belong in Palystes, the genus also does not belong near Panaretus, lacking all the
diagnostic characters found in those genera that I suggest belong in Heteropodinae
S. S., including the type genus. It does, however, share a number of characters with
Palystes (leg spination, cheliceral dentition, eye arrangement and proportions) and
belongs near Palystes rather than in Heteropodinae. It is thus (see above) removed
from Heteropodinae and placed in Palystinae.
• Arandisa Lawrence, 1938, a monotypic genus, was based on a subadult female
from Namibia. Both Arandisa and Orchestrella Lawrence, 1965 (with two
described species from Namibia) were assigned to Heteropodinae by Lawrence

because of the large size of the posterior lateral eyes (the largest of all the eyes). I
do not yet know where Arandisa and Orchestrella belong, but they do not belong
in Heteropodinae or Palystinae, sharing none of the characters discussed for either
group.
• Damastes Simon, 1880, has been listed under various subfamily groupings,
including Heteropodinae, and presently (possibly through an oversight by the
cataloguers) is listed under SparassinaelEusparassinae by Roewer (1954) and
Bonnet (1958). This distinctive genus of large, flattened, hairy spiders with some
11 nominal species, is endemic to Madagascar. Synonymised with the Australian
genus Delena and placed by Simon (1897a) in his group Clasteae, it was
reinstated by Simon (1903) within his Sparasseae (later Sparassinae) group. Hogg
(1903) removed the Australian genera Delena, Isopeda and Australian species
attributed to the Euro-Mediterranean genus Sparassus from the Sparasseae,
placing them in a new group, Deleneae. Although confirmed by Jarvi (1914),
Hogg's Deleneae were later consolidated under the Sparassinae by Petrunkevitch
(1928) until resurrected by Hirst (1989a). Like Hogg, Jarvi (1914) had realised the
problems of combining apparently unrelated genera within the Sparassinae and
had divorced Damastes from an implied relationship with Delena, transferring
DarrUiSies to his Toraniaeformes, a sub-group of Heteropodeae, along with
Torania, on the basis of certain genitalic similarities. This re-assignation was
apparently overlooked by the cataloguers. Examination of the type material of
Damastes shows that it does not belong in Heteropodinae s. s., lacking the
characters discussed above, and also does not belong in Deleninae sensu Hogg,
Jarvi and Hirst.
No useful diagnosis for Hetropodinae exists. Until it is revised, it is suggested that the
existing poor definition of the Heteropodinae be supplemented by using the diagnostic
characters outlined by Todd-Davies (1994) for Heteropoda and Yiinthi, in combination
with the following supplemental diagnosis based on characters shared by the genera
listed below. It is further suggested that the Heteropodinae be restlicted to those genera
sharing these characters. I have also listed those genera which do not share these
character states and which I feel should be excluded from Heteropodinae s. s.
Suggested supplemental diagnosis: Presence on cheliceral fang furrow floor of
median teeth (an elongate pad, or one or more rows, of microdenticles between large
teeth of margin); usually 4 cheliceral teeth on retrolateral margin (sometimes 3 as in
Keilira Hirst; sometimes 5 as in Gnathopalystes Rainbow), usually 3 teeth on
anterior lateral margin (2 in Gnathopalystes); anterior median eyes smaller than
anterior lateral eyes but tilted upwards and with their posterior (upper) rims on a
common line with posterior (upper) rims of anterior lateral eyes; AME equal, or
nearly so, to PME; posterior eye row with eyes either equal, subequal or with PLE
larger and raised, PER straight or a little recurved; cephalothorax domed or flat
posterior to caput; thoracic fovea generally inclined posteriorly or horizontal; labium
high-waisted, apically bluntly conical and only slightly wider than long.
A character present in Heteropoda, Gnathopalystes, Torania and Yiinthi, but not
apparent in the description of Keilira, is sexual dimorphism of leg length, with the
male tibiae proportionately longer, and with additional spine sets. The structure of the
male palp may also be characteristic; conductor spathulate, subtending the embolus
which is usually attenuate (or spathulate with a subterminal flagellum as in Yzinthi
Todd-Davies), but not coiled more than a single revolution on its axis.
Suggested included genera: Suggested excluded genera:
Heteropoda Latreille, 1804 Damastes Simon, 1880
Pandercetes L.Koch, 1875 Arandisa Lawrence, 1938
Panaretus Simon, 1880 Panaretella Lawrence, 1937
Torania Simon, 1886 Orchestrella Lawrence, 1965
Gnathopalystes Rainbow, 1899
Keilira Hirst, 1989
Yzinthi Todd-Davies, 1994
Gnathopalystes Rainbow, 1899, gen. resurr.

Gnathopalystes Rainbow, 1899: 313. Type species (by monotypy): GnathopalystesJerox Rainbow. 1899.
Palystes (in part); Simon, 1903: 1028 (recognised as junior synonym of Palystes).
Remarks: Gnathopalystes appears to be the only available candidate genus for a
number of species being transferred out of Palystes, and which I consider congeneric
with its type species Gnathopalystes ferox. This is not an ideal state of affairs since
the type of Gnathopalystes ferox is a subadult female collected from the eastern-most
part of the known range of the genus, and the male of the species is not yet known.
Until revisory work is done, Gnathopalystes is accordingly resurrected. The
subsequent nominal species are transferred from Palystes to Gnathopalystes. Specific
synonymies are not here established although pertinent literature is cited. However,
examination of the types of P. kochi and P. incanus show that the latter is a junior
synonym of the former, which is recognised here.
The species transferred from Palystes to Gnathopalystes are readily distinguished
from species belonging in Palystes s. s. by the characters summarised in Table 1
(p. 11 in text).
Species belonging in Gnathopalystes are distinguished from species presently
placed in the closely related genus, Heteropoda, by cheliceral dentition and the form
and proportions of the posterior eye row. In Gnathopalystes there are usually five
teeth on the retromargin of the cheliceral fang furrow (the fifth, basal, tooth very
small and sometimes absent) compared with four teeth in Heteropoda, and there are
only two teeth on the antero-margin (compared with three in Heteropoda). In
Heteropoda the PLE are larger than the PME and are raised as opposed to PLE=PME
and sessile in Gnathopalystes. I have already discussed my doubts about the validity
of these characters which may prove of little use when revising Heteropoda.
Nonetheless, I have used PER characters to help separate Heteropoda and
Gnathopalystes until such time that the two genera can be revised.
Diagnosis: Members of the genus have the following combination of characters:
Cephalothorax: Carapace domed posterior to PER, fovea inclined downwards to rear.
Eyes: PER from above straight, wider (approx. 1.3 X) than anterior eye row, PER
eyes homogeneous, equidistant, PLE sessile; AER from the front procurved.(upper
margins of AME in line with upper margins of ALE), heterogeneous, ALE larger
than AME, AME similar in size to all 4 eyes of PER. Chelicerae: Fang furrow floor
with numerous, sharp micro-denticles; retromargin usually with 5 teeth, subapical
largest, basal very small and sometimes missing; anteromargin with 2 teeth, apical
much larger than basal. Labium: Bluntly wedge-shaped apically, high-waisted
mesally. Sternum: Raised mesally at labium, without markings.
Legs: Spination I-IV similar to that given for Palystes except patellae have only
one, posterior, lateral spine and tibiae have no dorsal spines.
Abdomen: Ventrally usually with black marking posterior to epigynal groove.
Epigynum without externally produced septum, lateral lobes contiguous, externally
of same pattern illustrated by Hirvi (1914, PI. 6, Fig. 1) for Palystes ignicomus (now
transferred to Gnathopalystes). In the few males seen, embolus relatively short, thick
and coarsely attentuate, and arises close to point of origin of conductor which
subtends it.
Distribution: Malaysia and Oceanian islands of the Melanesian western Pacific,
including New Guinea, Papua New Guinea and Vanuatu.
GnathopalystesJerox Rainbow, 189Q

Gnathopalystes ferox Rainbow, 1899: 314, PI. XXV, Figs 5, 5a-c. Syntypes: 2 subadult 9 [the larger
here designated as lectotype, the smaller as paralectotype], 'Santa Cruz' [Vanuatu, Oceania]
(AMSA K27590 / KS 6630) [examined].
Palystesferox; Simon 1903: 1028 (transferred to Palystes).
Remarks: There are two, apparently conspecific, specimens in AMSA with the above
label data. One is a subadult female with a barely formed, faintly grooved,
unsclerotised, epigynal plate. It has similar dimensions to the single, female,
specimen described by Rainbow, allowing for some shrinkage in alcohol, and
matches his description although much of the colouring has been lost. This is clearly
the specimen referred to by Rainbow, who notes (p. 315) 'Epigyne obscured by hairs,
but the form is oblong, flat, grooved laterally and posteriorly, and somewhat U-
shaped.' In view of this, I have designated this specimen as the lectotype. The other
specimen is a smaller, teneral, juvenile/subadult female with ballooned, newly
formed cuticle around the abdomen and with no sign of an epigynal plate forming.
Although no other specimens are referred to in the original description, the second
female is probably part of a syntype series. It is therefore designated as a
paralectotype. The descriptive notes given above for the genus apply to the lectotype,
further supplemented by Rainbow's description.
Gnathopalystes crucifer (Simon, 1880), comb. n.

Palystes crucifer Simon, 1880: 266. Holotype: subadult 9 , 'Port-Said' [later emended (Simon, 1897a:
65, note 2) to 'de la peninsule Malaise ou de Java'] (MNHN) [examined].
Remarks: The specimen in Simon's type collection in MNHN is clearly the holotype
juvenile female. The cheliceral dentition, microdenticles on the floor of the fang
furrow, shape' of the carapace, the labium, and leg spination, all indicate that it
belongs in Gnathopalystes.
Gnathopalystes ignicomus (L. Koch, 1875), comb. D.

Palystes ignicomus L. Koch, 1875: 701, PI. LX, Figs 2, 2a. Holotype: 9 , 'Neu-lrland' (in a Mr Bradley's
collection) [type not located, presumed lost].
Sarotes vulpinus O. P. Cambridge, 1877: 286. Holotype: [subadult 9 - Pocock, 1899a], ['Duke of York
Island or on the adjacent coasts of New Ireland or New Britain' - Pocock, 1899al, (?OXUM)
[not examined].
?Heteropoda vulpina; Simon, 1880: 271 (tentative suggestion that species might belong in Heteropoda).
Palystes vulpinus; Pocock 1899a: 114 (recognised as synonym of ignicomus).
Remarks: Pocock (1899a) examined the type of vulpinus sent to him by Cambridge,
which Pocock (ibid.) said had been collected either on the Duke of York Island or on
the adjacent coasts of New Ireland or New Britain. He said the type, although a
subadult female, was clearly a synonym of ignicomus. Both Bonnet (1958: 2202) and
Roewer (1954: 2b: 1651) list Sarotes vulpinus Cambridge, 1877, as being transferred
to Heteropoda. This is based on Simon (1880: 271) who transferred vulpinus
tentatively to Heteropoda ('H. ? vulpina') but added that it might, in fact belong to
Tychicus. Simon does not mention seeing the Cambridge type. Until Heteropoda is
revised, Pocock's definite, and later, opinion that ignicomus and vulpinus are
synonymous is followed.
I have provisionally placed this species under Gnathopalystes where I think it may
belong, based on Koch's description and on Pocock's opinion and interpretation of
Palystes.
Gnathopalystes kochi (Simon, 1880), comb. D.
Palystes kochi Simon, 1880: 265. Holotype: adult 9 , 'Singapore' (MNHN 1.133) [examined].
Palystes melanichnys Thorell, 1890: 53. Holotype: 9 'ins. Nias ad Bawo Lowalani' [Sumatra] (?MCSN)
[not examine<j].
Palystes melanichnys; Thorell, 1895: 268 (synonymy with Palystes kochi) (BMNH).
Palystes incanus Thorell, 1890: 146. Holotype: adult 9, 'Sarawak, Borneo', Doria & Beccari (MCSN)
[examined]. Syn. n.
Palystes incanus; Thorell, 1892: 37 (?redescription).
Remarks: Thorell (1895: 268), in recognising Palystes melanichnys as a junior
synonym of Palystes kochi, also describes the male. The original description of
Palystes incanus is a 10-line synopsis which is later repeated (Thorell 1892) at the
head of a much amplified description. The original description nonetheless meets the
requirements of the Code and fixes the official date of publication of the name (see
Roewer 1954). The 1892 date for the species name given by Bonnet (1958) is not
correct. The examined types of Palystes incanus and Palystes kochi, both adult
females, are clearly synonymous. It is probable that the distribution of this species in
Singapore, Sumatra and Borneo has been aided by the agency of man. The dentition,
including the microdenticles of the fang furrow, descriptions and examined material
clearly place kochi in Gnathopalystes.
Gnathopalystes nigrocornutus (Merian, 1911), comb. D.

Palystes nigrocomutus Merian, 1911: 267. Holotype: 9, 'Wald bei Kema', Celebes (probably in
NHMB) [not examined].
Remarks: Merian's description is short, but agrees in all major respects with
Gnathopalystes. Until the type can be examined, his species is placed under
Gnathopalystes. His reference to its agreement with the retromarginal dentition

provided by Simon for P. crucifer is puzzling as Simon (1880) does not describe the
dentition of P. crucifer in the original description.
Gnathopalystes nigriventer (Kulczynski, 1910), comb. n.

Palystes nigriventer Ku1czynski, 1910: 400, PI. 17, Fig. II. Syntypes: adult 9, '(Ins. Shortland)
Poperang' [Solomon Islands] (NHMW), unspecified number of juvenile and subadult 9 , 'Ins.
Bougainville' [Papua New Guinea] (I specimen in NHMW, rest not located, not in ZMPA).
Remarks: The specimens described by Kulczynski were collected by Dr Kar]
Rechniger during his expedition to Samoa, New Guinea and the Solomon islands in
1905. NHMW has two specimens determined as Palystes nigriventer which were
donated by Kulczynski; one from Shortland Island and the other from Bougainville,
both collected by 'Rechinger', with the date 1906 (1. Gruber, in. litt.). Apart from the
date discrepancy, these are undoubtedly specimens from the syntype series. The rest
of the syntype series has not yet been located, but is not in Warsaw. Kulczynski's
description and illustration of the external epigynum indicate that his species almost
certainly belongs in Gnathopalystes.
Gnathopalystes rutilans (Simon, 1899), comb. n.

Palystes rutilans Simon, 1899: 99. Lectotype subadult female [here designated], 'Sumatra' 'Weyers'
(MNHN 11184) [examined], 5 juvenile paralectotypes [here designated] same locality (ISNB)
[examined].
Remarks: The subadult female lectotype from the Simon type collection in Paris is
clearly the subadult described by Simon. The five juvenile paralectotypes in
Brussels, collected at the same time as the lectotype, are labelled with the same data
as the lectotype and determined by Simon (who added '(Typus)' to the label) and are
obviously those referred to by Simon at the foot of his description. It is probable that
the species is con specific with G. kochi but this may never be determined since the
lectotype is subadult. The species belongs in Gnathopalystes.
Heteropoda Latreille, 1804

Heteropoda Latreille, 1804: 135.
Remarks: The urgently needed revision of this genus, to which a great many species
from eastern Asia and the Indo-Australasian region have been assigned, would be a
formidable challenge. Examination of material assigned to Heteropoda suggests that
the genus is polyphyletic and will eventually be divided into several genera. One
problem is resolving the identity of the type species, a subject which has been
debated at length by several authors and reviewed comprehensively by Bonnet (1951
1958). At present, since none of the type material for the various candidate names for
the type species is still extant, identification of the genus, and continued assignment
of species to it, rests largely on the interpretation of Simon (1897a) and Jarvi (1912
1914), and more recently Todd-Davies (1994) who provides a useful diagnosis and
definition for Australian species of Heteropoda.
The following three species clearly belong within Heteropodinae, but not in
Gnathopalystes as presently defined. I have already discussed the problem of
recognising Heteropoda on eye arrangement alone (following Simon) and place will
have to be found for those heteropodids, a number of which have already been
assigned to Heteropoda, with a Heteropoda-like eye arrangement (PLE large and
similar in size to ALE but raised above carapace on tubercles), but with a nearly
straight PER and a domed carapace rather than a more steeply procurved PER and
the flattened carapace usually associated with Heteropoda. Nonetheless, this remains
the only named candidate genus of which I am aware, and since it is unwise to risk
duplicating other revisory work on the Indo-Australasian genera by erecting a new
genus, the following three species are transferred to Heteropoda until they can be
placed with their congeners in a more natural group.
Heteropoda dasyurinus (Hogg, 1914), comb. n.

Palystes dasyurinus Hogg, 1914: 57. Holotype: adult 9 , Dutch New Guinea, 1912, Wollaston Exp
(BMNH 1921.3.24.119 & 1921. 65.) [examined].
Palystes dasyurinus; Hogg, 1915: 471-472, Figs 33a-c (amplified, illustrated description).
Remarks: Hogg's original description was a brief 6-line paragraph. This and the
amplified description published the following year agree with the specimen
examined, an adult female in BMNH.
The specimen has the typical heteropodine cheliceral tooth arrangement: a thick
pad of micro-denticles on the cheliceral fang furrow floor, flanked by three teeth in
the anterior row (middle tooth largest) and four in the posterior row (with the apical
tooth a little larger than the rest). The PER is nearly straight with the PLE large, on
raised tubercles, and similar in size to ALE (ALE=PLE»PME>AME), This
separates the species from those assigned to Gnathopalystes in which the PLE are
sessile and the eye proportions are: ALE»PLE=PME>AME. The carapace is domed
posterior to the caput and the fovea is inclined to the rear. For the reasons given
above, this species is transferred to Heteropoda.
Heteropoda ledleyi (Hogg, 1922), comb. n.

Palystes ledleyi Hogg, 1922: 296, Figs 6a-c. Syntypes: 3 adult 9 , I subadult d, 'hills in South Annam',
South Annam, Langbian Province. Dran, 3 000 ft., March-May 1918, C Boden Klass (BMNH
1924.III. 1.79 1-794 K52) [examined].
Remarks: The syntypes examined at BMNH have the cheliceral dentition which is
typical for many heteropodines (3 teeth in the anterior row, the apical tooth very
small, and four in the posterior row) and the pad of micro-denticles on the fang
furrow floor. The eye arrangement and proportions (PLE larger than PME and raised
on tubercles) separate this species from those assigned to Gnathoplaystes, and it is
transferred to Heteropoda for the reasons stated above.
Heteropoda speciosus (Pocock, 1898), comb. n.

Palystes speciosus Pocock, 1898: 468, PI. XIX, Figs 8, 8a. Syntypes: d9 , New Georgia and Ugi,
26.x.1894, officers of H.M.S. Penguin ['specimens also collected by Mr C.M. Woodford,
probably in Shortland Island'] (BMNH) [examined].
Remarks: Pocock's species is clearly a member of the Heteropodinae, possessing the
characters I have suggested are useful for diagnosing the subfamily, including the
microdenticles of the cheliceral fang furrow. Although close to Gnathopalystes, the

eye arrangement and proportions (including the PLE raised and larger than the PME)
place the species in Heteropoda, although, as I have commented earlier, this is a
variable character. The species also possesses the genitalic diagnostic characters used
by Todd-Davies (1994) for Australian species of Heteropoda, including the pattern
and form of conductor and embolus, but without the sub-terminal flagellum of the
conductor found in Yiinthi Todd-Davies. It may later be found to belong in a new
genus allied to the three genera. For the present I have placed Pocock's species in
Heteropoda where it more closely belongs than to any other genus I have examined
thus far.
SUBFAMILY STAIANINAE
Remarks: Simon (1897a) erected a tribe Staianeae within his clubionid subfamily
Sparassinae for Staianus acuminatus Simon, 1888 (based on the subadult female
holotype MNHN 9328 from Madagascar), adding Anchonastus (based on the adult
male holotype MNHN 19310 of Anchonastus caudatus Simon, 1898, from
Cameroon) in 1898 and Hoedillus (based on the subadult female holotype BMNH
1901.3.3-132 of Hoedillus sexpunctatus Simon, 1898, from Guatemala) in 1903.
Since then two further species have been added to Anchonastus. Petrunkevitch
(1928) recognised Simon's Staianeae at subfamily level as Staianinae.
The examined type material of the three genera placed by Simon in Staianeae
reveals that while there is some superficial similarity in carapace flattening and eye
arrangement, the relative proportions of their eyes and their general morphology are
very different. Hoedillus has an unusual double row of spine-like setae ventrally on
femur IV, and four large dorsal spines on the pedipalp tibia. It has a short, apically
bluntly wedge-shaped labium, carapace slightly concave posterior to the PER, an
unpigmented rounded sternum, and two, small, widely spaced, retromarginal teeth
and three anterior marginal teeth on the sides of the cheliceral fang furrow. Staianus
is lightly, but irregularly spined on legs I-IV, but without the ventral double row of
spines in Hoedillus, has a short, apically rounded labium, flat but slightly convex
carapace, pigmented, elongate, posteriorly acuminate, sternum, and two small
retromarginal teeth and a single small tooth on the anterior margin of the cheliceral
fang furrow. Although Anchonastus has some unusual apomorphic characters
(remarkably flattened carapace, tufts of plumose setae and posterior spinnerets with a
first segment twice as long as the first segment of the anterior spinnerets), it also
shares with Palystes and Parapalystes the same cheliceral dentition (3:3), leg
spination and eye arrangement and proportions, and in addition, the same sternal
banding pattern found in Palystes. This closer relationship with Palystes was
observed earlier by Pocock when describing Palystodes Pocock, 1899b (later
synonymised with Anchonastus by Simon in 1903). Pocock (1899b: 879) suggested
Palystodes was closely related to Palystes, separated only by the shape of its
carapace. The examined type material of Anchonastus and Palystodes clearly reveals
their synonymy, but the placement of Anchonastus in the Staianinae is unjustifiable. I
cannot find any characters which firmly establish a close relationship between the
three genera placed in the Staianeae by Simon. In view of the above observations,
Anchonastus is transferred to Palystinae. Hoedillus also does not appear to belong in

the Staianinae, but its placement must await revisory work on the genus.
ACKNOWLEDGEMENTS
Partial financial support for overseas travel from the Foundation for Research
Development and the Natal Museum is gratefully acknowledged. I am sincerely
grateful for the continued and protracted support of the Museum Council. I am
deeply indebted to the previously listed curators who went out of their way to find
type specimens and other material for this study and who have been so patient about
delays in the return of material. The following also provided invaluable information
and help: Dr H. M. Peters (Universitat Tubingen, Germany); Dr L. Kirkendall
(University of Oslo, Norway); the late Dr B. Petersen (Universitets Zoologiske
Museum, Copenhagen); Dr A. Nebois (Museum of Victoria, Australia); Dr V. Todd-
Davies and Dr R. Raven (Queensland Museum, Brisbane, Australia); Dr C. C. L.
Deeleman-Reinhold (Ossendrecht, The Netherlands); Dr V. D. Roth (Portal,
Arizona). For hospitality, help and laboratory space I am grateful to Dr Yves Coineau
and Dr Jacqueline Heurtault (MNHN), and Mr Fred Wanless and Mr Paul Hillyard
(BMNH), Dr Ansie Dippenaar-Schoeman (PPRI) , Mrs Eryn Griffin (SMWH), Mr
Wulf Haacke and Mrs Lomi Brown (TMSA), Dr Vin Whitehead, Mrs Cathy Car and
Mrs Margie Cochrane (SAMC), Dr Fred Gess and Mr Alan Weaving (AMGS), and
Dr Gerry Newlands (formerly of the South African Institute for Medical Research). I
would also like to thank the following for hospitality and for readily collecting
specimens and comparative material for me in southern Africa: Sally and Martin
Filmer (Johannesburg); John and Astri LeRoy (Roodepoort); Fred and Sarah Gess,
Peter Hawkes, Mike Bruton, Roy Lubke and Shirleen Gilbert (Grahamstown); John
Visser and ElOise Seekings (Cape Town); Patrick Reavell (Empangeni); June
Stannard (Port Edward); Chris and Tilde Stuart (Nieuwoudtville); Harold and Toni
Braack (whilst at Beaufort West); Per and Ruth Forchhammer (Serowe, Botswana);
Joe and Inge Henschel (Gobabeb, Namibia); Eryn and Mike Griffin (Windhoek). The
following are also gratefully acknowledged for their help: Tony Bruton, Veejay
Bandu and the late Jerry Kundulran of the Natal University Electron Microscopy
Unit, the late Dr Oliver Davies and Fiona Croeser for help with French and Latin
translations; Iris Bornman, for her active support and loyal friendship; Karin Herbert,
for locating and checking references; Annette Kunz and Bianca Lawrence, for
technical assistance; and Jason Landt and David Barraclough for their patience, wise
advice and help in preparing the manuscript for publication. lowe much to
discussions with Drs Michel Emerit, Otto Kraus, Valerie Todd-Davies, Pekha
Lehtinen, Brian Stuckenberg, John Poynton and Dick Kilburn, and advice and
comment on earlier drafts of sections of the manuscript by Drs Norm Platnick, Herb
Levi, Charles Griswold and Ansie Dippenaar-Schoeman. Prof. Denis Brothers
(University of Natal, Pietermaritzburg) gave unstinting support as a supervisor for
the thesis upon which this paper was based; the cladistics section received substantial
input from him. I am also deeply grateful for support and encouragement from my
wife, Fiona, my parents, Shirley and the late Michael Croeser, my sons Michael and
David, and my other sons, Phakamanimafrika Xaba, Matthews Thabethe, Bongani
Shelembe and Sabelo Msane, who so lovingly and supportively helped me and kept
the home running, thus allowing me to complete this study. I am also deeply in debt
to my mentors, the late Dr Reg Lawrence, who fIrst stimulated my fascination with
spiders, Prof. Denis Brothers, who so patiently and sympathetically led me into the
intricacies of biological systematics, and Dr Charles Griswold who never gave up on
me. And finally, my gratitude to Dr Ansie Dippenaar-Schoeman who has done much
to help me and foster a new generation of southern African arachnologists.
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INDEX TO GENERA AND SPECIES

Valid names are in italics and synonyms in roman. Generic names are in bold.
affinis Lessert ..................................... 67 lycosinus (Pocock) .............................90
amanicus Strand ................................. 67 martinfilmeri sp. n ............................. 30
Anchonastus Simon .......................... 92 megacephalus (C. Koch) ................... 90
ansiedippenaarae sp. n ...................... 78 megacephalus (sensu Pocock) ........... 90
Arandisa Lawrence .......................... 108 Micrommata Latreille ........................ 10
bomemiszai Caporiacco ..................... 70 minor Lawrence ................................. 92
castaneus (Latreille) ........................... 25 modificus Strand ................................ 56
caudatus Simon .................................. 93 natalius (Karsch) ............................... 56
chaperi Simon ................................... .25 nigriventer (Kulczynski) ................. 113
convexus Strand ................................. 85 nigrocornutus (Merian) ................... 112
crawshayi Pocock .............................. 39 Orchestrella Lawrence .................... 108
crucifer (Simon) .............................. 111 Palystelia Lawrence ........................... 10
cultrifer (Pocock) .............................. 90 Palystes L. Koch ................................ 13
Damastes Simon .............................. 109 Palystodes Pocock ............................. 92
dasyurinus (Hogg) ........................... 114 Panaretella Lawrence ........................91
disticta (Pocock) ............................... 92 Pandercetes L. Koch ........................ l1O
ellioti Pocock ..................................... 67 Parapalystes gen. n ............................ 85
euphorbiae sp. n................................. 87 perornatus Pocock ............................ .46
Exopalystes Hogg ................................ 9 pi/ipodus (Strand) .............................. 93
ferox Rainbow .................................. 111 pinnotherus (Walckenaer) ................. 84
flavidus Simon ................................... 83 plumosus (Pocock) ............................ 93
fornasinii (Pavesi) ............................. 84 Remmius C. Koch ..............................91
frenatus L. Koch ................................. 25 reticulatus Rainbow ........................... 84
gertschi Lessert .................................. 93 rubrioculatus Strand ........................... 70
Gnathopalystes Rainbow ................. 110 rutilans (Simon) .............................. 113
Helicopis L. Koch .............................. 13 scutata (Pocock) ................................ 92
H eteropoda Latreille ........................ 113 speciosus (Pocock) .......................... 114
hoehneli Simon .................................. 70 spenceri Pocock ................................. 56
ignicomus L. Koch ........................... 112 spiralis Strand .................................... 85
immaculata Lawrence ........................ 92 stilleri sp. n ........................................33
incanus (Thorell) ............................. 112 stuarti sp. n. .. .................................... 53
johnstoni Pocock ................................ 74 superci/iosus L. Koch ........................ 56
karooensis sp. n .................................50 superciliosus var. fasciiventris
Keilira Hirst ..................................... 110 Strand ............................................ 56
kibonotensis Lessert ........................... 70 Torania Simon ................................. 110
kochi (Simon) .................................. 112 venatoria (Linn.) ............................ .107
ledleyi (Hogg) ................................. 114 vulpinus (0. P. Cambridge) ............. 112
leppanae Pocock ............................... .41 whiteae (Pocock) ............................... 91
leroyorum sp. n ................................. 81 Yiinthi Todd-Davies ......................... 110
lunatus Pocock ................................... 37 zuluana Lawrence .............................. 92

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Ann. Natal Mus. Vol.

37 Pages 1-122 Pietennaritzburg October, 1996

A revision of the African huntsman spider genus Palystes L. Koch,

Fig. 1. Palystes castaneus (Latreille, 1819), female (Photograph by D. R. D'Ewes).

MATERIALS AND METHODS

AMGS Albany Museum, Grahamstown, South Africa (Dr F. W. Gess, Mr A.

NMSA Natal Museum, Pietennaritzburg, South Africa

NATURAL HISTORY OF PALYSTES

fovea steeply inclined. : fovea horizontal.

Diagnosis: Palystinae s. s. appears to be monophyletic, defined by the synapomorphy

furrow floor); ALE»PLE=PME>I=AME; PER straight or slightly recurved, eyes

Key to genera of Palystinae s. s.

Palystes L. Koch, 1875

preoccupied in Lepidoptera and replaced it with Palystes. He included four species in

Key to species of Palystes s .s.

white spots between black crescent and spinnerets (indistinct in specimens

Cape, South Africa) ............................................................... .leppanae Pocock

median channel (Figs 61-62) (Grahamstown and Alicedale districts, eastern

Palystes castaneus species group

Palystes castaneus (Latreille, 1819)

and 0.42 X of CW; eye ratio AME:ALE:PME:PLE 1:1.72:1.22:1.17; distances

Palystes martintilmeri sp. n.

posteriorly in 2 diverging elongate lobes. In P. stilleri (Fig. 32) septum swollen

Distribution: Piketberg, Clanwilliam and Cederberg districts of the western Cape,

Description offemale (SAMC B8887):

Palystes stilleri sp. n.

Embolus excavated anterolaterally in a wide, shallow trough (Fig. 28), reflexed

Description of holotype male:

Description offemale (PPRI '801193):

Additional material examined: None.

Palystes lunatus species group

Palystes lunatus Pocock, 1896

Additional material examined: None.

Palystes crawshayi Pocock, 1902

Additional material examined: LESOTHO: [Maseru], 29 , 1905, W. C. Wroughton

Palystes leppanae Pocock, 1902

Diagnosis: The females of P. Zeppanae and P. karooensis have a characteristic heart-

Legs: Spine pattern typical of genus, Measurements:

Additional material examined: SOUTH AFRICA: Cape: Alicedale: 1 9 (AMGS

Palystes peromatus Pocock, 1900

Redescription of lectotype female:

------~-.----- ... ----- ...

.•. " ........ .

Additional material examined: SOUTH AFRICA: Cape: Alice: Hogsback Mountain

Palystes karooensis sp. n.

Description ojparatype male (NMSA TS02):

Abdomen: 7.13 long and 5.13 wide.

Additional material examined: SOUTH AFRICA: Transkei/Natal: Cedarville,

Palystes stuarti sp. n.

Description of para type male (NMSA T500):

Abdomen: 12.25 long and 7.50 wide.

Additional material examined: None.

Palystes superciliosus species group

Palystes superciliosus L. Koch, 1875

(although tentatively suggesting that they were close to L. Koch's original

of the Limpopo, including Mozambique, Swaziland, the Transvaal, Natal, Transkei,

Redescription of holotype female of Heteropoda natalia:

Additional material examined: MOZAMBIQUE: Delagoa Bay, 1 9 (MNHN 9.959);

Hogsback Forest Reserve, 32°36'S 26°56'E, 39, 12-16.xii.1985, B. Londt (NMSA);

2104); Empangeni [28°45'S 31°54'E]: lei'19 , xii.1977, P. E. Reavell (NMSA

Dam (TMSA 1160), 19 , xii.1923, v. Dam (TMSA13069 old 2827); Onderstepoort

Villieria, }d, 8.xi.1976, M. Zwiegelaar (PPRI'76/1931), 19 , xi.1977, Mrs J. F. Smit

Palystes ellioti Pocock, 1896