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8181 Croeser 1996 Ann 37 1 122
8181 Croeser 1996 Ann 37 1 122
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
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
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:
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).
CROESER: REVISION OF PALYSTES (ARANEAE: HETEROPODIDAE) 9
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 .. ' .
10 ANNALS OF THE NATAL MUSEUM. VOL. 371996
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
CROESER: REVISION OF PALYSTES (ARANEAE: HETEROPODIDAE) 11
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,
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
CROESER: REVISION OF PALYSTES (ARANEAE: HETEROPODIDAE) 13
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).]
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).
16 ANNALS OF THE NATAL MUSEUM, VOL. 371996
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).
18 ANNALS OF THE NATAL MUSEUM, VOL. 37 1996
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.
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).
20 ANNALS OF THE NATAL MUSEUM, VOL. 37 1996
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
CROESER: REVISION OF PALYSTES (ARANEAE: HETEROPODIDAE) 21
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,
24 ANNALS OF THE NATAL MUSEUM, VOL. 37 1996
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.
CROESER: REVISION OF PALYSTES (ARANEAE: HETEROPODlDAE) 25
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.
CROESER: REVISION OF PALYSTES (ARANEAE: HETEROPODIDAE) 27
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.
28 ANNALS OF THE NATAL MUSEUM, VOL. 371996
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
CROESER: REVISION OF PALYSTES (ARANEAE: HETEROPODIDAE) 29
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).
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).
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.
Carapace: Dorsally 15.88 long, 12.50 wide between leg bases II. Head region 8.00
wide across PER; thoracic fovea 4.75 long, occupying 0.30 X of CL and 0.38 X of
CWo Eyes: Eye diameters: AME 0.63, ALE 1.00, PME 0.77, PLE 0.77, PER 5.00
wide, wider than AER by 1.25 X and occupying 0.63 X of HW and 0.40 X of CW;
eye ratio AME:ALE:PME:PLE is 1: 1.63: 1.22: 1.22; distances separating eyes:
AME-ALE 0.12, AME-AME 0.38, AME-PME 1.00, ALE-PLE 0.83, PME-PLE
0.82, PME-PME 0.77; MOQ dimensions: MOQA 1.66, MOQP 2.34, MOQL 2.25,
MOQ 0.96 X longer than wide, narrowed anteriorly, with MOQP 1.41 X wider than
MOQA. Clypeus height 0.82, 1.29 X diameter of AME. Chelicerae: 6.31 long, 3.69
wide, 1.71 X longer than wide. Sternum: 6.56 long, 5.00 wide between coxae II, 1.31
X longer than wide and 3.75 wide between coxae I. Labium: 2.55 wide, 2.18 long,
1.17 X wider than long, occupying 0.68 X of SW between coxae I. Maxillae: 3.85
long, 2.65 wide, 1.45 X longer than wide.
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.
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.
CROESER: REVISION OF PALYSTES (ARANEAE: HETEROPODIDAE) 35
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
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)
~"".'-""~'"''
.-~ ... ,
'-.
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).
CROESER: REVISION OF PALYSTES (ARANEAE: HETEROPODIDAE) 39
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.
Carapace: Dorsally 12.50 long, 9.63 wide between leg bases II. Head region 5.63
wide across PER; thoracic fovea 4.13 long, occupying 0.33 X of CL and 0.43 X of
CWO Eyes: Eye diameters: AME 0.57, ALE 0.80, PME 0.58, PLE 0.58, PER 3.88
wide, wider than AER by 1.11 X and occupying 0.69 X of HW and 0.40 X of CW;
eye ratio AME:ALE:PME:PLE is 1: 1.41: 1.03: 1.03; distances separating eyes:
AME-ALE 0.18, AME-AME 0.38, AME-PME 0.83, ALE-PLE 0.77, PME-PLE
0.71, PME-PME 0.62; MOQ dimensions: MOQA 1.38, MOQP 1.78, MOQL 1.63,
MOQ 0.91 X longer than wide, narrowed anteriorly, with MOQP 1.29 X wider than
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.
Abdomen: 16.00 long and 11.25 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
(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:
Total length: 26.50 mm.
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.'
Carapace: Dorsally 10.25 long, 9.38 wide between leg bases II. Head region 5.19
wide across PER; thoracic fovea 3.63 long, occupying 0.35 X of CL and 0.39 X of
cwo Eyes: Eye diameters: AME 0.46, ALE 0.69, PME 0.46, PLE 0.46, PER 3.38
wide, wider than AER by 1.20 X and occupying 0.65 X of HW and 0.36 X of CW;
eye ratio AME:ALE:PME:PLE is 1: 1.50: 1.00: 1.00; distances separating eyes:
AME-ALE 0.11, AME-AME 0.35, AME-PME 0.77, ALE-PLE 0.74, PME-PLE
0.62, PME-PME 0.46; MOQ dimensions; MOQA 1.23, MOQP 1.51, MOQL 1.07,
MOQ 0.94 X longer than wide, narrowed anteriorly, with MOQP 1.23 X wider than
MOQA. Clypeus 0.38 high, 0.83 X diameter of AME. Chelicerae: 4.06 long, 2.38
wide, 1.71 X longer than wide. Sternum: 4.38 long, 3.63 wide between coxae II, 1.21
X longer than wide and 2.63 wide between coxae I. Labium: 1.85 wide, 1.17 long,
1.58 X wider than long, occupying 0.70 X of SW between coxae L Maxillae: 2.46
long, 1.48 wide, 1.67 X longer than wide.
Abdomen: 16.25 long and 11.25 wide.
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
CROESER: REVISION OF PALYSTES (ARANEAE: HETEROPODIDAE) 41
Description o/male:
Total length: 17.76 rnm.
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.
Abdomen: 8.38 long and 4.88 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
../.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.
CROESER: REVISION OF PALYSTES (ARANEAE: HETEROPODIDAE) 43
Description of male:
Described reference male: Cape: Grahamstown, 6 Gilbert Street, found in house,
1O.vii.1982, Mrs Julia Stone (NMSA).
Total length: 16.63 mm.
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:
Eye diameters: AME 0.42, ALE 0.75, PME 0.49, PLE 0.49, PER 3.06 wide, wider than
AER by 1.20 X and occupying 0.69 X of HW and 0.39 X of CW; eye ratio
AME:ALE:PME:PLE is 1: 1.81: 1.19: 1.19; distances separating eyes: AME-ALE 0.11,
AME-AME 0.28, AME-PME 0.85, ALE-PLE 0.55, PME-PLE 0.49, PME-PME 0.46;
MOQ dimensions: MOQA 1.11, MOQP 1.51, MOQL 1.38, MOQ 0.92 X longer than
wide; narrowed anteriorly, with MOQP 1.36 X wider than MOQA; dypeus 0.46 high,
1.11 X diameter of AME. Chelicerae: 3.13 long, 1.88 wide, 1.67 X longer than wide.
Sternum: 4.38 long, 3.31 wide between coxae II, 1.32 X longer than wide and 2.50 wide
between coxae I. Labium: 1.57 wide, 0.86 long, 1.82 X wider than long, occupying 0.63
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.
Abdomen: 7.75 long and 4.63 wide.
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
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).
46 ANNALS OF THE NATAL MUSEUM, VOL. 371996
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.
Carapace: Dorsally 10.75 long, 9.13 wide between leg bases II. Head region 5.19
wide across PER; thoracic fovea 3.00 long, occupying 0.28 X of CL and 0.33 X of
Cw. Eyes: Eye diameters: AME 0.46, ALE 0.69, PME 0.51, PLE 0.51, PER 3.31
wide, wider than AER by 1.15 X and occupying 0.64 X of HW and 0.36 X of CW;
eye ratio AME:ALE:PME:PLE is 1: 1.50: 1.1 0: 1.10; distances separating eyes:
AME-ALE 0.15, AME-AME 0.42, AME-PME 0.85, ALE-PLE 0.69, PME-PLE
0.54, PME-PME 0.49; MOQ dimensions: MOQA 1.26, MOQP 1.54, MOQL 1.69,
MOQ 1.10 X longer than wide, narrowed anteriorly, with MOQP 1.22 X wider than
MOQA. Clypeus 0.38 high, 0.83 X diameter of AME. Chelicerae: 4.06 long, 2.38
wide, 1.71 X longer than wide. Sternum: 4.69 long, 3.56 wide between coxae II, 1.32
X longer than wide and 2.81 wide between coxae I. Labium: 1.85 wide, 1.05 long,
1.76 X wider than long, occupying 0.66 X of SW between coxae I. Maxillae: 2.55
long, 1.78 wide, 1.43 X longer than wide.
Abdomen: 11.88 long and 7.13 wide.
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
Description of male:
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).
Total length: 22.25 mm.
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.
Carapace: Dorsally 11.00 long, 8.75 wide between leg bases II. Head region 4.69
48 ANNALS OF THE NATAL MUSEUM. VOL. 371996
\.
,
••..
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).
CROESER: REVISION OF PALYSTES (ARANEAE: HETEROPODIDAE) 49
wide across PER; thoracic fovea 3.38 long, occupying 0.31 X of CL and 0.39 X of
Cw. Eyes: Eye diameters: AME 0.51, ALE 0.69, PME 0.52, PLE 0.51, PER 3.13
wide, wider than AER by 1.19 X and occupying 0.67 X of HW and 0.36 X of CW;
eye ratio AME:ALE:PME:PLE is 1: 1.36: 1.03: 1.00; distances separating eyes:
AME-ALE 0.11, AME-AME 0.35, AME-PME 0.78, ALE-PLE 0.77, PME-PLE
0.55, PME-PME 0.54; MOQ dimensions: MOQA l.23, MOQP 1.54, MOQL l.69,
MOQ 1.10 X longer than wide; narrowed anteriorly, with MOQP 1.25 X wider than
MOQA; clypeus 0.54 high, 1.06 X diameter of AME. Chelicerae: 3.75 long, 2.19
wide, 1.71 X longer than wide. Sternum: 4.56 long, 3.63 wide between coxae II, 1.26
X longer than wide and 3.13 wide between coxae I. Labium: 1.75 wide, 1.08 long,
1.63 X wider than long, occupying 0.56 X of SW between coxae I. Maxillae: 2.55
long, 1.54 wide, 1.66 X longer than wide.
Abdomen: 11.25 long and 7.50 wide.
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
and with scattered small dark spots. Sternum with 2 vestigial, widely interrupted bars
of darker setae at coxae II and III.
Carapace: Dorsally 10.25 long, 8.25 wide between leg bases II. Head region 4.88
wide across PER; thoracic fovea 2.88 long, occupying 0.28 X of CL and 0.35 X of
Cw. Eyes: Eye diameters: AME 0.46, ALE 0.69, PME 0.54, PLE 0.54, PER 3.00
wide, wider than AER by 1.07 X and occupying 0.62 X of HW and 0.36 X of CW;
eye ratio AME:ALE:PME:PLE is 1:1.43:1.17:1.17; distances separating eyes:
AME-ALE 0.12, AME-AME 0.35, AME-PME 0.75, ALE-PLE 0.69, PME-PLE
0.48, PME-PME 0.48; MOQ dimensions: MOQA 1.43, MOQP 1.52, MOQL 1.54,
MOQ 0.98 X longer than wide, narrowed anteriorly, with MOQP 1.06 X wider than
MOQA. Clypeus 0.54 high, 1.17 X diameter of AME. Chelicerae: 3.88 long, 2.25
wide, 1.72 X longer than wide. Sternum: 4.19 long, 3.56 wide between coxae II, 1.18
X longer than wide and 3.00 wide between coxae I. Labium: 1.94 wide, 1.08 long,
1.80 X wider than long, occupying 0.65 X of SW between coxae I. Maxillae: 2.31
long, 1.51 wide, 1.53 X longer than wide.
Abdomen: 12.13 long and 8.25 wide.
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
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).
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
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.
Carapace: Dorsally 15.00 long, 12.25 wide between leg bases II. Head region 6.88
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:ALE:PME:PLE is 1:1.63:1.25:1.13; distances separating eyes: AME-ALE 0.15,
AME-AME 0.46, AME-PME 1.05, ALE-PLE 0.89, PME-PLE 0.75, PNIE-PME 0.66;
MOQ dimensions: MOQA 1.57, MOQP 2.09, MOQL 1.69, MOQ 0.81 X longer than
wide, narrowed anteriorly, with MOQP 1.33 X wider than MOQA. Clypeus 0.46 high,
0.75 X diameter of AME. Chelicerae: 5.63 long, 2.81 wide, 2.00 X longer than wide.
Sternum: 5.94 long, 4.69 wide between coxae II, 1.27 X longer than wide and 3.63 wide
between coxae 1. Labium: 2.62 wide, 1.85 long, 1.42 X wider than long, occupying 0.72
X of SW between coxae I. Maxillae: 3.85 long, 2.31 wide, 1.67 X longer than wide.
Cheliceral margin teeth typical of genus (3 retromarginal and 3 anteromarginal teeth),
but left chelicera has a small fourth, additional anteromarginal tooth apically.
Abdomen: 18.75 long and 13.38 wide.
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
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.]
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).
....... ~ .. ····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).
58 ANNALS OF THE NATAL MUSEUM, VOL. 37 1996
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.
Carapace: Dorsally 13.00 long, 10.50 wide between leg bases II. Head region 6.25
wide across PER; thoracic fovea 3.50 long, occupying 0.27 X of CL and 0.33 X of
CWO Eyes: Eye diameters: AME 0.63, ALE 0.91, PME 0.62, PLE 0.60, PER 4.19
wide, wider than AER by 1.20 X and occupying 0.67 X of HW and 0.40 X of CW;
eye ratio AME:ALE:PME:PLE is 1:1.44:0.98:0.94; distances separating eyes:
AME-ALE 0.12, AME-AME 0.37, AME-PME 0.69, ALE-PLE 0.60, PME-PLE
0.65, PME-PME 0.63; MOQ dimensions: MOQA 1.54, MOQP 1.91, MOQL 1.69,
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
long, 4.13 wide between coxae II, 1.23 X longer than wide and 3.13 wide between
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):
CROESER: REVISION OF PALYSTES (ARANEAE: HETEROPODIDAE) 61
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
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),
64 ANNALS OF THE NATAL MUSEUM, VOL. 37 1996
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
CROESER: REVISION OF PALYSTES (ARANEAE: HETEROPODIDAE) 65
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.
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
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
70 ANNALS OF THE NATAL MUSEUM, VOL. 37 1996
~ 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.
72 ANNALS OF THE NATAL MUSEUM, VOL. 371996
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).
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
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).
74 ANNALS OF THE NATAL MUSEUM. VOL. 371996
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
between coxae I. Maxillae: 3.2 long, 2.31 wide, 1.39 X longer than wide.
Abdomen: 14.88 long and 11.50 wide.
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
\
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).
78 ANNALS OF THE NATAL MUSEUM, VOL. 371996
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).
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).
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.
Carapace: Dorsally 9.38 long, 7.75 wide between leg bases II. Head region 4.19
wide across PER; thoracic fovea 2.25 long, occupying 0.24 X of CL arid 0.29 X of
CWO Eyes: Eye diameters: AME 0.45, ALE 0.60, PME 0.42, PLE 0.42, PER 2.75
wide, wider tharl AER by 1.10 X and occupying 0.66 X of HW and 0.35 X of CW;
eye ratio AME:ALE:PME:PLE is 1:1.34:0.93:0.93; distances separating eyes:
AME-ALE 0.09, AME-AME 0.35, AME-PME 0.57, ALE-PLE 0.51, PME-PLE
0.42, PME-PME 0.43; MOQ dimensions: MOQA 1.20, MOQP 1.32, MOQL 1.35,
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
wide, 1.97 X longer than wide. Sternum: 3.75 long, 3.25 wide between coxae II, 1.15
X longer than wide arid 2.63 wide between coxae I. Labium: 1.66 wide, l.08 long,
1.54 X wider than long, occupying 0.63 X of SW between coxae I. Maxillae: 2.62
long, 1.45 wide, 1.81 X longer tharl wide.
Abdomen: 7.50 long and 5.25 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
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.
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.
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
86 ANNALS OF THE NATAL MUSEUM, VOL. 371996
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.
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
: 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
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.
CROESER: REVISION OF PALYSTES (ARANEAE: HETEROPODIDAE) 87
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).
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.
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).
CROESER: REVISION OF PALYSTES (ARANEAE: HETEROPODIDAE) 91
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.
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
96 ANNALS OF THE NATAL MUSEUM. VOL. 37 1996
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:
CROESER: REVISION OF PALYSTES (ARANEAE: HETEROPODIDAE) 97
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.
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
98 ANNALS OF THE NATAL MUSEUM, VOL. 37 1996
0..
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a.
~ e
UI
UI eCl C.
:;,
Cl
UI
.!!!
~III ~>- UI e ~
UI
g
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~ Cl
Q)
"iii
(Il
c:
a
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...m
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rn
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c: (J "til c.
(Il c: (Il m
c.. 0:( c.. u ~ ~
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9~2 9~O
15~1 15~2
18~1 18~2
12~1
16~3
C
1~2 4~1
2~1 5~1
6~1 7~1
15~1 11~1
D
3~1
14~1
16~1
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.
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.
100 ANNALS OF THE NATAL MUSEUM. VOL. 371996
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
CROESER: REVISION OF PALYSTES (ARANEAE: HETEROPODIDAE) 101
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.]
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.
B~3
13~2 19~3
p J K
15~3
M H
15~2
N
2~2
3~1
7~1
9~2
10~1
11~2
17~3
19~2
20~2
8~1
5~1
11~1
12~1
17~1
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.
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
106 ANNALS OF THE NATAL MUSEUM. VOL. 37 1996
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
CROESER: REVISION OF PALYSTES (ARANEAE: HETEROPODIDAE) 107
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
108 ANNALS OF THE NATAL MUSEUM. VOL. 37 1996
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
CROESER: REVISION OF PALYSTES (ARANEAE: HETEROPODIDAE) 109
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
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.
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.
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.
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,
116 ANNALS OF THE NATAL MUSEUM, VOL. 37 1996
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
CROESER: REVISION OF PALYSTES (ARANEAE: HETEROPODIDAE) 117
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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