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Taxonomic notes on giant southern African stick insects
(Phasmida), including the description of a new
Bactrododema species
P. D. Brock
‘Papillon’, 40 Thorndike Road, Slough SL2 1SR, United Kingdom
E-mail: pbrock@wexhamcourt.slough.sch.uk
BROCK, P. D., 2004. Taxonomic notes on giant southern African stick insects (Phasmida), including the description of
a new
Bactrododema
species.
Annals of the Transvaal Museum
41: 61–77
The southern African species of
Bactrododema
Stål, 1858,
Palophus
Westwood, 1859 (both Diapheromeridae:
Palophinae) and
Bactricia
Kirby, 1896 (Diapheromeridae: Diapheromerinae), are reviewed.
Bactrododema krugeri
spec. nov. is described, while
Bactrododema hecticum
(Lichtenstein, 1796) comb. nov. is redescribed and discussed.
Bactrododema brevitarsis
Stål, 1876,
Bactrododema aculiferum
Kirby, 1902,
Bactrododema lugardi
Kirby, 1902,
Palophus brevicornis
Redtenbacher, 1908,
Palophus holubi
Redtenbacher, 1908, and
Palophus transvaalensis
Redtenbacher, 1908, are synonymized with
Bactrododema tiaratum
Stål, 1858.
Phibalosoma calametum
Bates,
1865,
Hyrtacus carinatus
Kirby, 1902, and
Bactricia irregularis
Brunner, 1907, are synonymized with
Bactricia
bituberculata
(Schaum, 1857) stat. rev., which takes priority over
Bactricia trophinus
(Westwood, 1859). The genus
Palophus
is regarded as a synonym of the genus
Bactrododema. Palophus titan
Sjöstedt, 1913 is synonymized with
Bactrododema hippotaurum
(Karsch, 1896) comb. nov.
Palophus rothschildi
Bolivar, 1922 is synonymized with
Bactrododema phillipsi
(Kirby, 1896) comb. nov., and
Palophus brongniarti
Redtenbacher, 1908 is synonymized with
Bactrododema episcopalis
(Kirby, 1896) comb. nov. Keys to the adults of South African species are provided, eggs of
two
Bactrododema
species are described, and the distribution of all southern African
Bactrododema
species
mapped.
Keywords: Phasmida,
Bactricia, Bactrododema, Palophus,
Taxonomy, Southern Africa.
INTRODUCTION
In a recent catalogue of the stick insect fauna, Otte
and Brock (2003) included nine South African
species from the following genera: Bactrododema
Stål, 1858; Palophus Westwood, 1859; and
Bactricia Kirby, 1896, with three species in each
genus. A review of the South African species was
considered necessary to evaluate the fauna, partic-
ularly as many synonyms of stick insect species
exist in this unstudied insect order worldwide
(Brock, 1999). Morphological variation in phasmids
can be extreme and without examining a series of
specimens, it is sometimes difficult to observe
intraspecific variation (Brock, 1999). Even though
little is known about most of the spectacular larger
African phasmids, they have attracted some atten-
tion from entomologists. Sjöstedt (1913) included a
colour plate of his new species Palophus titan, and
Brongniart (1892) published a monograph of
the genus Palophus, listing Bactrododema as a
synonym, although it has priority over Palophus.
Redtenbacher (1908) provided keys to the species
of Bactrododema and Palophus, and regarded
these as valid genera, as did Bradley and Galil
(1977). Several species were described in various
short publications between 1908 and 1922 (these
are listed in the species group sections). Even
though this paper considerably expands our knowl-
edge of the distribution of some stick insect
species, very little is known about the biology of
these insects. Here the southern African species of
Bactrododema and Bactricia are reviewed, species
groups proposed for Bactrododema, and Bactrodo-
dema krugeri spec. nov. described.
MATERIAL AND METHODS
Material examined is deposited in the following
collections: The Natural History Museum, London,
U.K. (BMNH); Entomologisches Institut, ETH-
Zentrum, Zurich, Switzerland (ETHZ); Natur-
historiska Riksmuseet, Stockholm, Sweden
(NHRS); Naturhistorisches Museum Wien, Austria
(NHMW); Oxford University Museum, Hope Ento-
mological Collections, Oxford, U.K. (OXUM); P. D.
Brock Private Collection, Slough, U.K. (PB); South
African Museum, Cape Town, South Africa (SAMC);
National Collection of Insects, Pretoria, South Africa
(SANC); Skukuza Biological Reference Collection,
Kruger National Park, Skukuza, South Africa
(SBRC); Transvaal Museum, Pretoria, South Africa
(TMSA); and the Museum für Naturkunde der
Humboldt-Universität, Berlin, Germany (ZMHB).
This study is based on the examination of all
primary types in the aforementioned collections.
Species were compared side by side, whenever
possible. New and previous synonyms are listed in
chronological order, with a brief explanation of the
key records where significant. Notes on the genera
are given (including synonymy), with references to
the original description of species and type details,
followed by the reference to the same species.
Non-type material examined mainly refers to speci-
mens measured and examined during a visit to
South Africa in 2003. For species of Bactrododema
and Palophus not covered in detail below, the
detailed type data is included in Otte and Brock
(2003).
Key to the genera of giant southern African
Phasmida
1. Elongate, wingless insects. Head with
pair of well-spread short horns or spines.
Female with long chute at end of abdomen
(Fig. 2). Male with large overlapping spatu-
late cerci (Fig. 1). . . . . . . . . . . Bactricia Kirby
— Elongate, male with full-sized wings (Figs 4,
7, 9), female wings shortened (Figs 3, 6, 8).
Head with pair of variably sized horns, often
large and almost fused. End of abdomen in
female normal, operculum not extending
beyond endofabdomen.Cerci short . . . .
. . . . . . . . . . . . . . . . . . . . Bactrododema Stål
Family DIAPHEROMERIDAE
Subfamily DIAPHEROMERINAE
Genus
BACTRICIA
Kirby, 1896
Bactricia Kirby, 1896: 463. Bacteria trophinus
Westwood, 1859: 30, pl. 5: 5. Type species by
original designation.
Bactricia; Kirby, 1904: 327; Brunner, 1907: 330;
Brock, 1995: 86; Otte and Brock, 2003: 116.
Scaphegyna Karsch, 1898: 364, 366. Bacteria
trophinus Westwood, 1859: 30, pl. 5: 5. Type
species by original designation.
Scaphegyna; Kirby, 1904: 327 (synonym of
Bactricia Kirby); Brunner, 1907: 330; Otte and
Brock, 2003: 116.
REDESCRIPTION [following Kirby (1896)]. Apterous
in both sexes: body very long and slender. Head
usually with two strong spinous processes on vertex
in male (sometimes very short), lamellate pro-
cesses or spines in female close together. Legs
pubescent, especially tarsi; first joint of tarsi as long
or longer than rest, not appendiculate, but carinate
in female. Legs unarmed in male, armed with strong
sub-triangular spines in female, on carinae above
and below (but may be inconspicuous or absent).
Median segment one-fifth length of metanotum in
male, one-fourth in female. Cerci of male large,
broad, almost spatulate, and strongly curved,
crossing each other. Operculum of female very
long, longer than last three segments of abdomen
together and slightly expanded at extremity.
Bactricia bituberculata
(Schaum, 1857) stat.
rev., Figs 1, 2
Bacteria bituberculata Schaum, 1857: 423.
Bacteria bituberculata Schaum; Westwood,
1859: 180 (as Schaum MSS); Schaum, 1862:
558. The following records list B. bituber-
culata as a synonym of Bacteria trophinus:
Kirby, 1896: 463; Kirby, 1904: 327; Brunner,
1907: 330; Otte and Brock, 2003: 116.
Bacteria trophinus Westwood, 1859: 30, pl. 5: 5.
Bactricia trophinus (Westwood); Kirby, 1896:
463, pl. 39: 3, 3a, 4, 4a (female; as B.
trophimus [sic]; synonymized with Bacteria
bituberculata Schaum); Kirby, 1904: 327;
Brunner, 1907: 330, pl. 15: 1a–b; Otte and
Brock, 2003: 116.
Phibalosoma calametum Bates, 1865: 341; syn.
nov.
Bactricia calametum (Bates); Kirby, 1904: 327;
Brunner, 1907: 330; Otte and Brock, 2003:
116.
Scaphegyna calametum (Bates); Karsch, 1898:
367.
Hyrtacus carinatus Kirby, 1902: 46, pl. 2: 3–4;
syn. nov.
Hyrtacus carinatus Kirby; Kirby, 1904: 331; Otte
and Brock, 2003: 130.
Bactricia irregularis Brunner, 1907: 330; syn. nov.
Bactricia irregularis Brunner; Brock, 1998: 36;
Otte and Brock, 2003: 116.
BODY LENGTH. Male 120–142 mm (n= 7), female
165–200 mm (n= 10).
DISTRIBUTION. Only recorded from Mozambique,
South Africa (KwaZulu-Natal, Mpumalanga, Western
Cape) and Swaziland.
REMARKS. Brunner (1907) did not consider mor-
phological variation when he redescribed this
species. His simplified key to females distinguished
related species by the presence of a double-spined
head or lamellated process, which is a variable
feature in both sexes, as are dentations on the
legs.MM
TYPE MATERIAL EXAMINED. B. bituberculata Holo-
type 9, MOZAMBIQUE: Inhambane, Peters (ZMHB).
B. trophinus Holotype 8, SOUTH AFRICA: Port Natal
[Durban] (BMNH). P. calametum Holotype 9,
62 ANNALS OF THE TRANSVAAL MUSEUM, VOLUME 41, 2004
*As currently there is no agreement on this insect order name, the
oldest name (Phasmida) is used, other authors favour
Phasmatodea or Cheleutoptera.
BROCK: TAXONOMY OF GIANT SOUTHERN AFRICAN STICK INSECTS 63
Fig. 1
Bactricia bituberculata
male, A: dorsal view; B: head, lateral view
(after Kirby, 1896).
Fig. 2
Bactricia bituberculata
female, A: dorsal view, B: head, lateral
view (after Kirby, 1896).
SOUTH AFRICA ‘Caffraria’, M. Gerrard (OXUM) H.
carinatus Syntypes: 18,19, SOUTH AFRICA:
Barberton, Rendall, [ex. Distant collection acquired
in 1911] (BMNH). Bactricia irregularis Holotype 9,
SOUTH AFRICA: Cape of Good Hope ‘Cap. d.g.h.’,
Thorey (NHMW).
ADDITIONAL MATERIAL EXAMINED (58,89).
SOUTH AFRICA: Barberton, ix.1921, E. Howard (19
TMSA); Barberton, 1906 (19SANC); Dientje,
Graskop, ii.1918, de Celliers (19TMSA); Hazyview,
18.iv.1981, E. F. von Malitz (19nymph TMSA);
Marieps Mnt., i.1926, G. van Son (19TMSA); Port
Natal [Durban] (28,19BMNH); Port St Johns,
24–30.xi.1956 (18TMSA); Umbogantwini, Natal,
23.x i.[19]42, Marley (19TMSA); no data (19TMSA).
MOZAMBIQUE: Inhaca, Isl. P. E., 20–27.ix.1937,
G. van Son (18TMSA). SWAZILAND: M’babane,
20.xi.1929 (18TMSA).
Family DIAPHEROMERIDAE
Subfamily PALOPHINAE
Key to adult females of South African
Bactrododema
species
1. Head with a pair of large horns. Elongate or
‘stocky’, mesonotum with a pair of even
or uneven central spines . . . . . . . . . . . . . . . 2
— Head with a pair of shorter horns. Elongate.
Mesonotum with a pair of central spines
(sometimes absent, or in extreme cases,
with several spines). Body length 125–
185 mm (Fig. 3a,b) . . . . . . . . B. tiaratum Stål
2. Moderately long c. 140 mm. ‘Stocky’, some-
times elongate, mesonotum with a few
well-spread tubercles. Mid and hind femora
with a few small dentations. Body length
126–150 mm (Fig. 6). . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . B. hecticum Lichtenstein
— Elongate c. 200 mm. Mesonotum elongate,
with a pair of large central, usually uneven
spines. Conspicuous sub-basal lobes on
mid and hind femora. Body length 193–
226 mm (Fig. 8 a,b). . . B. krugeri spec. nov.
Key to adult males of South African
Bactrododema
species
1. Head with a pair of narrow horns curved
backwards. Femora lacking dentations or
foliose lobes. Hind wings tessellate dark
brown with medium-sized whitish blotches.
Body length 109–133 mm (Fig. 4a,b) . . . .
. . . . . . . . . . . . . . . . . . . . . . . B. tiaratum Stål
— Head with a pair of large horns, straight
or pointing forwards. Femora with den-
tation(s) and/or foliose lobes. Hind wings
tessellate dark brown with small or larger
whitish blotches. . . . . . . . . . . . . . . . . . . . . . 2
2. Body of medium length. Legs with only a
few short dentations on femora and tibiae.
Hind wings tessellate dark brown with
series of small whitish spots or blotches.
Femora lacking subapical lobes. Body
length 104–124 mm (Fig. 7a,b,c) . . . . . . .
. . . . . . . . . . . . . . . . B. hecticum Lichtenstein
— Body long. Femora with subapical lobes
and broad sub-basal foliose lobes, includ-
ing other large dentations/lobes. Hind
wings tessellate dark brown with mainly
large whitish spots or blotches, particularly
on inner margin. Body length 138–171 mm
(Fig. 9a,b,c) . . . . . . . . . B. krugeri spec. nov.
Key to eggs of South African
Bactrododema
species
1. Capsule length c. 5 mm, micropylar plate
large, broad-lobed (Fig. 5a,b)
. . . . . . . . . . . . . . . . . . . . . . . B. tiaratum Stål
— Capsule length c. 7 mm, micropylar plate
small, ‘lozenge-shaped’ (Fig. 10a,b)
. . . . . . . . . . . . . . . . . . . B. krugeri spec. nov.
2. Eggs unknown . . . B. hecticum Lichtenstein
Genus
BACTRODODEMA
Stål, 1858
Bactrododema Stål, 1858: 308. Bactrododema
tiarata Stål, 1858: 308. Type species by
monotypy.
Bactrododema; Stål, 1875a: 12, 14 Stål, 1875b:
32, Brongniart, 1892: 635 (synonym of
Palophus Westwood), Kirby, 1904: 366,
Redtenbacher, 1908: 391; Karny, 1923: 366;
Otte and Brock, 2003: 286.
Palophus Westwood, 1859: 90. Palophus
centaurus Westwood, 1859: 233, pl. 32: 1, 1a.
Type species by subsequent designation of
Kirby,1904: 365. Brongniart, 1892: 193, Kirby,
1896: 465; Karsch, 1898: 367; Rehn, 1904:
70, Kirby, 1904: 365; Redtenbacher, 1908:
393 (in part); Karny, 1923: 239; Otte and
Brock, 2003: 288.
Ischnopoda Grandidier, 1869 [Homonym of
Ischnopoda Stephens, 1835 (Coleoptera)].
Ischnopoda reyi Grandidier, 1869: 293. Type
species by monotypy. [Uvarov, 1940: 176,
proposed Ischnophasma, a synonym of
Palophus Westwood, as a replacement name
for Ischnopoda].
Ischnopoda; Lucas, 1869: 430; Brongniart,
1892: 25 (synonym of Palophus); Karsch,
1898: 367; Kirby, 1896: 468; Kirby, 1904: 365;
Redtenbacher, 1908: 393; Karny, 1923: 239;
Bradley and Galil, 1977: 185; Bragg, 2001:
635; Otte and Brock, 2003: 288.
64 ANNALS OF THE TRANSVAAL MUSEUM, VOLUME 41, 2004
REDESCRIPTION (based on the type species and
African species examined). Large, elongate, cylin-
drical, sometimes body ‘stout’ in female. Antennae
in male ‘hairy’, almost as long as fore legs; slightly
longer than femora in female. Head broad, back of
head spiny or tuberculate, anteriorly with a pair of
long, broad ‘tuft-like’ horns almost joined, which
can be curved forwards or backwards – sometimes
reduced in size (as in the type species), or excep-
tionally replaced by spines in female. In some
species, including the type species B. tiaratum, the
male has a pair of very narrow strands instead of
horns. Ocelli in male distinct, absent in female.
Pronotum longer than head, mesonotum elongate,
granular or occasionally spiny. Mesonotum often
with a pair of large central spines, but a variable
feature with the spines even, uneven, sometimes
absent or occasionally specimens with several
large central spines. Forewings ovate with large
round or conical tubercle. Hind wings in male large,
completely formed. In female wings variably sized,
but shortened compared with male. Legs elongate,
base of fore femora curved inwards. Femora, tibiae
and tarsi variably lobed or dentate, often conspicu-
ously so, although sometimes lobes/dentations
absent. Abdomen elongate, broadened in some
females and may have central tufts or lobes on
mid-abdominal segments. Some species, for exam-
ple B. tiaratum and B. centaurum, are much nar-
rower than others, particularly in the males. Cerci
short, subgenital plate in male rounded, operculum
in female cymbiform, not extending beyond end of
abdomen.
REMARKS. The generic status of Bactrododema
and Palophus has varied over the years. Brongniart
(1892) regarded Bactrododema as a synonym of
Palophus. However, Palophus was described after
Bactrododema Stål, which therefore has prior-
ity. Kirby (1904) listed Bactrododema as a valid
genus, as did Redtenbacher (1908), although the
latter transferred the type species to Palophus,
synonymizing Palophus with Bactrododema.
Ischnophasma (= Ischnopoda) was listed as a
synonym by Brongniart (1892), accepted by Karsch
(1898), and Otte and Brock (2003). Kirby (1896,
1904), Karny (1923), Bradley and Galil (1977) and
Bragg (2001) treated it as a valid genus.
The genus Dematobactron Karny, 1923 [Type
species: Bactrododema fuscipenne Redtenbacher,
1908, recorded from Gabon and Ghana] is morpho-
logically very similar to Bactrododema and may be
synonymous. Only the female is known, a rather
slender insect with shorter wings than usual in
Bactrododema.
Examination revealed that the female of Bactro-
dodema tiaratum Stål, 1858, is a shorter form of
Palophus phillipsi (Kirby, 1896), except for the
shorter paired horns on the head and other minor
differences, suggesting Bactrododema and
Palophus to be the same genus. The horns are a
variable feature in Bactrododema, with longer horns
mainly occurring on the longest species (often
c. 200 mm and longer in the female). There may be
some value in regarding Palophus as a subgenus of
Bactrododema, but even illustrations of the three
South African species show how variable these
species are, with long and short insects present.
Thus, taking morphological variability into account, I
consider that the species included are representa-
tives of a single, but variable genus.
Eggs of Bactrododema species are similar in
appearance and have unusual shapes, but differ in
the length and shape of the micropylar plate. The
eggs of B. tiaratum, B. krugeri and B. reyi are illus-
trated in Figs 5, 10, 14.
The following three groups of Bactrodema
species are proposed:
Bactrododema centaurum
species group
Head with a pair of large horns in both sexes,
mesonotum smooth or granular, except for two
(sometimes more, rarely none) large central spines
in females. Wings tessellate. Only B. hecticum
(Lichtenstein) is medium-sized to large (male
c. 120 mm; female ≤150 mm), all other species are
distinctly longer (male c. ≤180 mm; female
c. ≤200 mm), and can be divided into species where
males have fully tessellate hind wings: B. hecticum
(Lichtenstein, 1796) comb. nov. [= B. haworthii
(Gray, 1835) and Haplopus ceratophyllus Bur-
meister, 1838] (South Africa: Eastern, Western and
Northern Cape Provinces, also Namibia); B. krugeri
spec. nov. (Botswana, South Africa: Gauteng,
Mpumalanga, Limpopo Province); B. reyi
(Grandidier, 1869) comb. nov. (Angola (?), Kenya,
Uganda, Zambia) and species where males have
whitish wings, tessellate only towards the outer
margin: B. alldridgei Kirby, 1905 comb. nov. (Sierra
Leone); B. centaurum (Westwood, 1859) comb.
nov. (Cameroon, Central African Republic, Congo,
Gabon, Nigeria, Togo); B. episcopalis (Kirby, 1896)
comb. nov. [= Palophus brongniarti Redtenbacher,
1908 syn. nov.] (Tanzania).
In the following species only females are known,
therefore they could fall within either subgroup
mentioned above, as the wing colour of males is
unknown: B. bayeri (Schouteden, 1917) comb. nov.
(Kenya), B. leopoldi (Schouteden), 1916 comb. nov.
(Democratic Republic of Congo), B. moirae (Kirby,
1896) comb. nov. (Central Africa: Lauderdale), B.
phillipsi (Kirby, 1896) comb. nov. [= Palophus
rothschildi Bolivar, 1922 syn. nov.] (Ethiopia,
Kenya).
BROCK: TAXONOMY OF GIANT SOUTHERN AFRICAN STICK INSECTS 65
Bactrododema hippotaurum
species group
Head with a pair of large horns in both sexes,
mesonotum spiny, with many lateral spines, includ-
ing two large central spines in female. Wings tessel-
late. A single species: B. hippotaurum Karsch, 1896
comb. nov. [= Palophus titan Sjöstedt, 1913 syn.
nov.] (Tanganyika, Malawi).
Bactrododema tiaratum
species group
Head with a pair of shorter horns in both sexes
(male tufts narrow) or sometimes replaced by
spines, mesonotum with variable number of large
central spines, often two, but sometimes absent.
Wings tessellate, usually shorter in female. This
group comprises seven species (although those
from outside South Africa require further research):
B. tiaratum Stål, 1858 [= Bactrododema brevitarsis
Stål, 1876 syn. nov., Bactrododema aculiferum
Kirby, 1902 syn. nov., Bactrododema lugardi Kirby,
1902 syn. nov., Palophus brevicornis Redten-
bacher, 1908 syn. nov., Palophus holubi Redten-
bacher, 1908 syn. nov., and Palophus transvaal-
ensis Redtenbacher, 1908 syn. nov. (Botswana,
Namibia, South Africa: Gauteng, KwaZulu-Natal,
Mpumalanga, Northern Cape, Limpopo Province,
North West, Western Cape; and Zimbabwe)]; B.
aestuans (Westwood, 1859) (Congo, Gabon, Sene-
gal, São Tomé); B. minotaurum (Gerstäcker, 1883)
(Ghana); B. miliaris Bolivar, 1889 (São Tomé); B.
welwitschi Bolivar, 1889 (Angola, Gabon); B. wayi
Kirby, 1902 (Malawi); B. pectinicornis (Redten-
bacher, 1908) comb. nov. (Tanzania).
BIOLOGY. Nymphs and adults feed on Acacia
species (Mimosaceae/Leguminosae). Nymphs
resemble miniature versions of adults and gradually
develop wings. Adults have been collected
throughout the year, but mainly from November to
March.
DISTRIBUTION. The species are widespread
throughout central and southern Africa, although
often recorded from a limited range.
Southern African
Bactrododema
species
Bactrododema tiaratum
Stål, 1858, Figs 3, 4, 5
Bactrododema tiarata Stål, 1858: 308.
Bactrododema tiarata Stål; Stål, 1875b: 82; Stål,
1876: 66; Bolivar, 1889: 87; Brongniart, 1892:
203; Karny, 1923: 239.
Bactrododema tiaratum Stål; Kirby, 1902: 47;
Kirby, 1904: 366, Picker et al. 2002: 108 [as
Palophus reyi; misidentification]; Otte and
Brock 2003: 287.
Palophus tiaratus (Stål); Griffini, 1897: 11;
Redtenbacher, 1908: 396, pl. 18: 5.
Bactrododema brevitarsis Stål, 1876: 67; syn.
nov.
Ischnopoda brevitarsis (Stål); Kirby, 1904: 366.
Palophus brevitarsis (Stål); Redtenbacher, 1908:
397; Karny, 1910: 89; Otte and Brock, 2003:
288.
Bactrododema aculiferum Kirby, 1902: 47, pl. 2:
1–2; syn. nov.
Bactrododema aculiferum Kirby; Kirby, 1904:
366; Redtenbacher, 1908: 393; Taylor 1987:
157 (in Zimbabwe, as Bactrododermus [sic]);
Otte and Brock, 2003: 287.
Bactrododema lugardi Kirby, 1902: 448; syn.
nov.
Bactrododema lugardi Kirby; Kirby, 1904: 366;
Redtenbacher, 1908: 393; Otte and Brock,
2003: 287.
Palophus brevicornis Redtenbacher, 1908: 396;
syn. nov.
Palophus brevicornis Redtenbacher; Karny,
1910: 89; Otte and Brock, 2003: 288.
Palophus holubi Redtenbacher, 1908: 397; syn.
nov.
Palophus holubi Redtenbacher; Brock, 1998: 33;
Otte and Brock, 2003: 289.
Palophus transvaalensis Redtenbacher, 1908:
396; syn. nov.
Palophus transvaalensis Redtenbacher; Brock,
1998: 62; Otte and Brock, 2003: 289.
REMARKS. There is wide variation in this species,
both in size (small females only 125 mm, others up
to 185 mm) and the number of head/thoracic
spines.
BIOLOGY. A colony of this species, originating from
Zimbabwe, was maintained at the London Zoo
(United Kingdom) in the 1960s, until it was stolen.
Recent attempts to maintain and rear B. tiaratum in
South Africa on Acacia met with limited success.
Brock (2000) provided detailed notes on how to rear
phasmids in general, which can be applied to this
species.
TYPE MATERIAL EXAMINED. B. tiaratum Holotype
9, [NAMIBIA] Damara (NHRS). B. brevitarsis
Holotype 9, [NAMIBIA] Damaraland (NHRS).
B. aculiferum Syntypes: 18, SOUTH AFRICA:
Barberton [presumably collected by Rendall and
with a label attached ‘spec. figd’];1 9, Crocodile
River, Distant (BMNH, possibly also 18,19, no data,
ex. Distant collection, which may not be type mate-
rial). B. lugardi Holotype 8, [BOTSWANA]: N’Gami
country, pres. Sir F.D.J. Lugard, 1898 (BMNH). P.
brevicornis Holotype 9, SOUTH AFRICA: Zeerust
(ETHZ). P. holubi Holotype 9, [BOTSWANA]: Tati,
‘South Africa’, Dr Holub (NHMW – alcohol coll.).
66 ANNALS OF THE TRANSVAAL MUSEUM, VOLUME 41, 2004
BROCK: TAXONOMY OF GIANT SOUTHERN AFRICAN STICK INSECTS 67
Fig. 3
Bactrododema tiaratum
female, A: dorsal view, B: head, lateral
view.
Fig. 4
Bactrododema tiaratum
male, A: dorsal view, B: lateral view
(after Redtenbacher, 1908).
P. transvaalensis Syntype 19, SOUTH AFRICA:
Transvaal, Hartmann (NHMW).
ADDITIONAL MATERIAL EXAMINED (278, 399).
SOUTH AFRICA: Barberton, 19–26.viii.[19]20, G.
v[an]. D[am], A. Rob[erts] (18TMSA); Barberton,
15.i.1927, J.S. Taylor (18BMNH); Barberton, Distant
coll. (18BMNH); Barberton, P. Rendall (39BMNH);
Barberton, 25.iii.[19]12, E.L. Lounsbury (19TMSA);
Barberton, 15.iv.1910, Miss De Beer (19TMSA);
Barberton, J.C. Morgan (19TMSA); Boksberg,
Johannesburg, C.H. Pead [acquired 1907] (19
BMNH); Blouberg, Tvl., 6000–6714 feet, 10.i.1955,
Transvaal Mus. Exp. (28TMSA); D’Nyala Nature
Res., De Kroon, i.1911, G. v. Dam (19TMSA);
Ellisras, 23°45’S, 27°49’E, 850 m., 17–20.xii.1987
[at light], M.W. Mansell (18SANC); (?) Glenlyon,
Cape Prov., 11.i.1919, Hay (18SANC); Hans
Merensky Nat. Res., 23°42’S, 30°44’E,
23–25.i.1987, B. Grobbelaar (18SANC);
Hartbeesportdam, SE25 29CA, 20.iii.1982, R. Gricia
(18TMSA); Hectorspruit, Tvl., 6.vii.[19]21, J. Dea-
con (19SANC); Hluhluwe Bushlands, KwaZulu-Na-
tal, 28°08’S, 32°18’E, 4.xii.1994, Gareth Davies (18
TMSA); Kaapmuiden, 23.ii.1920, J.B. Pienaar (19
TMSA); Kiepersol, Mpumalanga, 26°10’S, 31°07’E,
20.ii.1998, J. Smith (18TMSA); Klip Koppies, Kruger
National Park, 23°51’17”S, 31°44’29”E, viii.1965, S.
Joubert (19SBRC ref. no. A784); Knysna, 34°03’S,
23°03’E, 29.x.1991, SK325 1991/180, S. Kamburov
[‘feeds on citrus leaves’] (18SANC); Komatipoort,
iii.1935, E.D. Ras (19TMSA); Lapalala Nat. Res.,
Tvl., 23°51’S, 28°17’E, 21–22.i.1987, E. Grobbelaar
(18SANC); Leeupoort, 24°53’S, 27°43’E,
27.xii.1997, C.W. Louw (28TMSA); Letaba,
25.i.1943, E. Bedford (18TMSA); Lothair, O[os]
T[rans]v[aa]l, 26°22’S, 30°29’E, 4.iv.1996, C.
Coetzee (19TMSA); Louis Trichardt, 6.ii.1952 (28
TMSA); M’dup, iv.[19]70 (19TMSA); Lydenburg, B.
Buys, v.1963 (19TMSA); Malekon, 25°28’S,
31°32’E, 12.v.1989, W.L. Pretorius (18TMSA);
Marloth Park, 25°22’S, 31°55’E, 29.iii.1989, L.G. Le
Roux (18TMSA); Mogol Nature Reserve, Ellisras
Dist., Tvl., 23°58’S, 27°45’E, 25–26.i.1982, R.G.
Oberprieler (18SANC); Naboomspruit, 5.iii.2000,
M. Picker (18PB); Ngwenya Lodge, Mpumalanga,
25°25’S, 31°52’E, 29.xi.–9.xii.1996, P.W. Bateman
(19TMSA); Noordwes, 24°54’S, 27°31’E,
24.iii.1998, F. Swanepoel (19nymph TMSA);
Nylsvley Nature Res., Tvl., 24°39’S, 28°42’E,
1095m, iii.1978 (S. Nun) (19SANC); Orpen [to
Satara] Road, Kruger National Park, 3.iv.2001, R.B.
Toms (19TMSA [and eggs]); Paardefontein, Preto-
ria, Tvl., 7.iii.1971, J.E. Prinsloo (19SANC [and
eggs]); Pelindaba, Tvl., 6.iii.1975, H. Mern (19
SANC); Pietersburg dist., v.1962, via Dr A.J.T. Janse
(19TMSA); Pretoria, G.H. (19TMSA); Sabi Sand
Wild Tuin, iii.v.[19]69, J. Herbert (19TMSA); Satara
Region, Kruger National Park, 2233b, u324,
14.iii.1984, R. Toms (19TMSA); Skukuza, Kruger
National Park, 24°59’41”S, 31°35’25”E, 13.v.1966,
T. de Vos (19SBRC ref. no. C280); Skukuza, Kruger
National Park, 24°59’41”S, 31°35’25”E, 22.i.1968, J.
Braack (19SBRC ref. no. E414); Skukuza,
11.ii.2004, P.D. Brock (18PB); Transvaal [acquired
1893] (29BMNH); Upington, 3.vii.1925, H.J.
Schmidt (19TMSA); Warmbaths, Transvaal,
30.i.1918 (19SANC); Zoutp[ans]b[erg] dist.,
6.vii.1917, H.G. Breyer (19TMSA); No data (19
TMSA), (29SANC); 25°24’S, 28°16’E, 25.iv.1996,
S.P. Schoeman (19PB); 2/15, G. v. Son (18TMSA);
No data, labelled B. aculiferum (19TMSA).
NAMIBIA: Grootfontein, 19°32’S, 18°06’E,
12.i.1976, R. Oberprieler (18SANC); 30m. N.
Okahandja, 1.iv.1969, W.D. Haacke (19SANC);
Windhoek, at light, R.G. Oberprieler (18TMSA).
ZIMBABWE: 8, Birchenough Bridge, S.Rhod.,
i.1938, G. v. Son (18TMSA); Umtali District, Rhode-
sia, ii.1932, P.A. Sheppard (19TMSA).
Bactrododema hecticum
(Lichtenstein, 1796)
comb. nov., Figs 6, 7
Phasma hecticum Lichtenstein, 1796: 78;
Lichtenstein, 1802: 39, pl. 1: 2; Redten-
bacher, 1908: 397 (synonym of Palophus
haworthii).
Plactycrana hectica (Lichtenstein); Gray, 1835:
34.
Cyphocrania ?hectica (Lichtenstein); Westwood,
1859: 109.
Palophus ?hecticus (Lichtenstein); Kirby, 1904:
365.
68 ANNALS OF THE TRANSVAAL MUSEUM, VOLUME 41, 2004
Fig. 5
Bactrododema tiaratum
egg, A: dorsal view, B: lateral view.
Ctenomorpha haworthii Gray, 1835: 41.
Palophus haworthii (Gray); Westwood, 1859: 90,
pl. 10: 5; Kirby, 1904: 365; Redtenbacher,
1908: 397, pl. 18: 6; Karny, 1910: 89.
Bactrododema haworthii (Gray); Günther, 1956:
93; Otte and Brock, 2003: 287.
Haplopus ceratophyllus Burmeister, 1838: 577.
Haplopus ceratophyllus (Burmeister); West-
wood, 1859: 90; Redtenbacher, 1908: 397
(synonym of Palophus haworthii).
REMARKS. Based on the original description
(Lichtenstein, 1796), and Lichtenstein’s (1802)
illustration of this species there is no doubt that
B. hecticum is the valid name for this species. Even
though authors tentatively linked B. hecticum and
C. haworthii, they may have had doubts because of
Lichtenstein’s recorded type locality ‘China’ for
B. hecticum. The holotype female of B. hecticum is
in Holthuisen’s collection, ex. Hamburg, possibly
sold at auction.
TYPE MATERIAL EXAMINED. P. hecticum Holotype
9(not 8) ‘China’ [in error ?], not traced. C. haworthii
Holotype 8, ‘Australia’ [in error, believed to be
SOUTH AFRICA] (OXUM). H. ceratophyllus
Syntypes: 18,29, [SOUTH AFRICA], Cape of Good
Hope, Pr[omontorium] b[onae] sp[ei] (8Alund, 19
Meyer), (ZMHB).
ADDITIONAL MATERIAL EXAMINED (138,69).
SOUTH AFRICA: G[rea]t. Fish R[iver], [acquired
1893] (19BMNH); Karoo National Park, Beaufort
West, Cape Province, 32°20’S, 22°28’E, 900 m,
21–22.xii.1989, H. & U. Aspöck, M.W. Mansell (28
SANC); Kenhardt, Driekop, 29°23’S, 21°06’E,
850 m, 19.ii.1991, M.W. Mansell (18SANC);
Kenhardt, Driekop NR, N. Cape, 29°24’S, 21°06’E,
850 m, 24.ii.1996, M.W. Mansell, H. & U. Aspöck, H.
Hölzel, P. Ohm (28SANC); Longhope, Cape
Province, i–ii.1928, Ethel Gough (18BMNH);
Longhope, Cape Province, 30.v.1927, Ethel Gough
(19BMNH); Prieska Poort, Cape Province, 29°47’S,
22°42’E, 1050 m, 18.ii.1991, M.W. Mansell (18
SANC); Willowmore, Cape, various dates, xii.1925
to 24.v.1928, Dr H. Brauns (28,49TMSA); Wolfdrif,
Pakhuispad, Western Cape Province, 5.ix.1999, M.
Picker (18PB); South Africa [no other data] (18
BMNH). NAMIBIA: Bullspoort, iv.[19]49, R.G. Strey
(18TMSA); Amichab Mts Game Res. No. 3,
27.iv.[19]66, H. Dick Brown (18SANC).
Bactrododema krugeri
spec. nov., Figs 8–12;
Tables 1, 2
DESCRIPTION. Adult male (holotype) (Fig. 9). Large
brown stick insect, various shades of brown with
BROCK: TAXONOMY OF GIANT SOUTHERN AFRICAN STICK INSECTS 69
Fig. 6
Bactrododema hecticum
female in dorsal view.
whitish mottling, including brown and white banding
on legs. Body length 163 mm (295 mm including
outstretched legs).
Head. Heavily tuberculate, slightly longer than
wide, tapering slightly towards hind part. Large
raised pair of horns (4.5 mm height) between eyes.
Compound eyes large, dark brown. Two orange
ocelli present on either side of horns. Back of head
with three ridges. Antennae long, ‘hairy’; with 31
segments; longer than fore legs. Basal segment
slightly thicker and second segment greatly
reduced in length. Apices of segments slightly darker.
Thorax. Pronotum much longer than wide, con-
spicuously longer than head, with several short
tubercles. Conspicuous ridges present, the most
prominent at two-thirds length of segment. Meso-
notum elongate (3.5 times length of pronotum), with
slightly uneven, short pair of central spines less than
1 mm high at an angle of 45° (left-hand spine further
up). A bold central line present, with series of ridges
70 ANNALS OF THE TRANSVAAL MUSEUM, VOLUME 41, 2004
Fig. 7
Bactrododema hecticum
male, A: dorsal view, B: head, lateral view (after Westwood, 1859), C: lateral view (after Redtenbacher,
1908).
on remainder of segment; slightly ‘hairy’, some
small tubercles/granules present. Underside paler.
Metanotum and median segment combined longer
than mesonotum.
Abdomen. Elongate, slightly ‘hairy’, sparsely
granular. End of segment 5 rather swollen and
ridged. Abdominal segment 8 reduced in size, less
than half length of segment 7, hind part broadened.
Segment 9 tapered towards the anal segment,
expanded towards the hind part, which is almost
triangularly excised at tip; with three black denta-
tions beneath. Cerci short, incurved, broadened
towards rounded tip. Sub-genital plate with large,
but narrow lobe, end triangular excised in centre,
reaching just over half length of abdominal segment
9 (Fig. 11).
Wings. Forewings elongate with strongly raised
tubercle; mottled green and dark brown, outer
margin whitish. Hind wings long, not quite reaching
end of the sixth abdominal segment. Preanal part of
hind wings dark brown tessellate with whitish spots
and blotches of moderate size, larger on inner part
of wings.
Legs. Fore femora with central crest. A pair of
apical spines and lobes on all femora, lobes shorter
on hind femora. Underside of fore femora with small
spine beneath crest in middle of segment. Mid and
hind femora with broad sub-basal lobe on upper
and underside; smaller central lobe on upper side,
shorter on hind femora. All tibiae with large
sub-basal crest and smaller subapical crest.
Underside of tibiae with three evenly spaced
serrations along length on hind legs, other legs with
a single serration beyond sub-basal lobe. Tarsi short.
Intraspecific variation. Paratype males (13 speci-
mens). Same as holotype, except for minor size and
colour variations in the number and size of serra-
tions/crests (some lack crests on fore femora). Most
individuals lack central spines on mesonotum. Hind
wings may vary slightly, one paratype has smaller
white spots/blotches, or spots may be yellowish
rather than white.
Adult female. (Fig. 8) Paratype females (19 adults,
1 nymph). Whitish brown with mottled legs. Body
length of adults 193–226 mm (n= 18).
Head. Heavily tuberculate, slightly longer than
wide, tapering towards back. Large raised horns
(7 mm high) present between eyes. Eyes large, dark
brown. Back of head with three ridges. Antennae
long, ‘hairy’, and with about 31 segments; longer
than fore legs. Basal segment slightly thicker.
Thorax. Pronotum much longer than wide, con-
spicuously longer than head. Several ridges pres-
ent, the largest one at two-thirds length of segment.
Mesonotum elongate (3.5 times length of pro-
notum), with an uneven, rarely even, pair of central
spines pointing outwards in most specimens, about
BROCK: TAXONOMY OF GIANT SOUTHERN AFRICAN STICK INSECTS 71
Fig. 8
Bactrododema krugeri
female, A: dorsal view, B: head, lateral
view.
72 ANNALS OF THE TRANSVAAL MUSEUM, VOLUME 41, 2004
Fig. 9
Bactrododema krugeri
holotype male, A: dorsal view, B: head, lateral view, C: hind wing.
3 mm high. Usually spine further up on right hand
side, but sometimes reversed. Metanotum and
median segment longer than mesonotum.
Abdomen. Elongate. End of the fourth abdominal
segment with swollen raised lobes, larger at end of
segment 5. Abdominal segment 8 reduced in size.
Cerci short, slightly incurved, rounded at tip. End of
anal segment triangular, excised in centre. Oper-
culum ‘boat-shaped’, tapered to almost pointed tip,
reaching about half length of anal segment 9
(Fig. 12).
Wings. Forewings rounded at tip, with strongly
elevated tubercle. Hind wings short for genus, not or
just reaching end of abdominal segment 3. Preanal
part of hind wings dark brown and yellowish tessel-
late.
Legs. As in male, but lobes/crests shorter.
Eggs (Fig. 10). Various shades of brown. Oper-
culum flat, with protrusion not quite central. Outer
ring of raised cristae. Opercular angle 40°. Capsule
sculptured and strongly raised in area around the
small ‘lozenge-shaped’ micropylar plate, which
sometimes has a darker outer rim. Capsule size:
length 7 mm; height 2.8 mm; width 2.5 mm. Egg
stuck ventrally to suitable surfaces, e.g., tree
branches.
BROCK: TAXONOMY OF GIANT SOUTHERN AFRICAN STICK INSECTS 73
Fig. 10
Bactrododema krugeri
egg, A: dorsal view, B: lateral view.
Fig. 11
Bactrododema krugeri,
end of male abdomen, A: dorsal view,
B: lateral view.
Table 1
Body measurements in millimetres of the adult type series of
Bactrododema krugeri
.
Holotype male Paratypes (13 males) Paratypes (19 females)
Body length 163 138–171 193–226
(mean 157) (mean 210)
Head 5.5 4–5.5 7–8.5
Antennae 110 100–110 62–66
Pronotum 7 6–7 10–11
Mesonotum 24 20–24 35–38
Metanotum* (incl. median segment) 28 (9) 22–28 (8–9) 34–37 (11–12)
Forewings 20 15–20 23–25
Hind wings 97 78–97 56–60
Profemora 50 40–50 54–58
Mesofemora 40 30–40 42–46
Metafemora 49 39–50 53–57
Protibiae 62 47–64 65–69
Mesotibiae 44 33–45 49–53
Metatibiae 58 47–60 64–68
Cerci 2 2 2.5–3
*The metanotum length includes the separate median segment length in brackets, as the first abdominal segment (= median segment) is fused to the
metanotum.
DISTRIBUTION. Fairly widespread in eastern South
Africa (Gauteng, Mpumalanga, Limpopo Province),
also recorded from Botswana (Fig. 13). It is likely
that B. krugeri occurs in Mozambique and possibly
elsewhere. However, it appears to have a more
restricted distribution in southern Africa than
B. tiaratum.
BIOLOGY. Known food plants in South Africa in-
clude Acacia species, such as knob thorn (A.
nigrescens, Mimosaceae/Leguminosae). Miller
(1949) recorded feeding nymphs, of what may be B.
krugeri, on Swartzia madagascariensis (Legumi-
nosae). Miller (1949) described the egg of
‘Palophus tiaratus’ from the Odzi district in Zimba-
bwe, which is similar to that of B. krugeri and may
represent the same species, but it is definitely not
B. tiaratum. Adults have been collected and seen at
lights, including those in the rest camps of the
Kruger National Park.
ETYMOLOGY. Named after the Kruger National
Park, a magnificent National Park famous through-
out the world for its diversity of wildlife.
74 ANNALS OF THE TRANSVAAL MUSEUM, VOLUME 41, 2004
Fig. 13
Distribution of
Bactrododema
species in southern Africa.
Bactrododema tiaratum
(✱);
B. krugeri
(▲);
B. hecticum
(●).
Fig. 12
Bactrododema krugeri,
end of female abdomen, A: lateral view,
B: ventral view.
REMARKS. A review of species in the Bactrodo-
dema centaurum group suggests that B. krugeri is
closest to B. phillipsi and B. reyi. While the male of
B. phillipsi is undescribed, elaborate dentations,
crests and lobes on the legs of B. krugeri readily
distinguish it from B. reyi (illustrated in Karsch,
1898), which also lacks the broad sub-basal lobes.
The female of B. leopoldi from Katanga, Democratic
Republic of Congo (near the border with Zambia) is
similar to B. krugeri, but the legs are more lobed and
measure 94–97 mm, and the hindwings are much
longer than in B. krugeri. The male of B. leopoldi is
undescribed. Key differences in the females are
included in Table 2.
TYPE MATERIAL. Holotype 8, SOUTH AFRICA:
Skukuza Rest Camp, Kruger National Park,
18.iii.1985, R. Toms, (18TMSA). Paratypes (138,
209) SOUTH AFRICA: Cape Town, iv.1912, Lord
Gladstone [in view of the known distribution of this
species, this locality is likely to be an error] (18
TMSA); Geluk, Waterb[erg] dist., vi.1916, de Villiers
(19TMSA); Giyani, Gazankula, 8.viii.1976, P.J.
Steencamp (19TMSA); Komatipoort, xii.1920, Mr
Haagner (19nymph TMSA); Linhope River,
i.viii.1917 (19SANC); Louis Trichardt, 6.ii.[19]52 (19
TMSA); Louis Trichardt, 6.viii.1968, H.S. Snyman (18
TMSA); Lydenburg, Col. Darmant (19TMSA);
Malalane, ii.1919, Elphick (19TMSA); Messina,
Northern Prov[ince], 22°37’S, 39°52’E, 13.ii.1999,
Natalie Wilson (18TMSA); Messina Nature Reserve,
22°22’S, 30°02’E, 554 m, 11–12.ii.1985, at light, H.U
& C. Aspöck, H. Hölzel, M.W. Mansell (28SANC);
Mica, viii.1934, W.C. Rogers (19TMSA, ex. Univer-
sity of Pretoria coll.); Noordelike Prov., Ellisras,
23°45’S, 27°04’E, 20.iii.1996, H. v. d. Merwe (18
TMSA); Nylstroom dist., iv.1948, Dr W. Maherk (19
TMSA); Potgietersrust, T.P., viii.1927, C. Fuller (19
SANC); Pretoria, Gauteng, 25°45’S, 28°12’E,
3.v.1996, J.A. Marais (18TMSA); Skukuza, Kruger
BROCK: TAXONOMY OF GIANT SOUTHERN AFRICAN STICK INSECTS 75
Table 2
Key differences between females of three
Bactrododema
species.
Character
B. krugeri B. phillipsi B. reyi
Head / paired horns Tuberculate. Horns broad,
c.
7 mm high. Tuberculate. Horns elongate / narrowed
towards tip,
c.
9 mm high. Rather spiny behind horns. Horns broad,
c.
8mm
high.
Mesonotum Pair of long central rarely even spines, spine
on left lower down, but sometimes higher up. Pair of central spines, either even, or spine
on left higher up; spine not long for the genus. Pair of long central spines, either even, or spine
on left higher up.
Forewings Strongly raised tubercle. Rather rounded at tip. Strongly raised tubercle. Elongate, ovate. Strongly raised tubercle. Elongate, ovate.
Hind wings Rather short, not or just reaching end of third
abdominal segment. Reaching about half length of fourth
abdominal segment. Just exceeding end of third abdominal segment.
Mesofemur and metafemur Broad sub-basal lobe on upper and underside. Base slightly broadened. Base slightly broadened.
End of tenth abdominal (anal)
segment Triangular excised in centre. Truncate / sub-truncate, slightly pointed at
either end. Emarginate, laterally lobed.
Egg Fairly elongate, length
c.
7 mm. Elongate, length
c.
10.9 mm (detailed study by
Scali and Mazzini 1990, ex. Somalia). Elongate, length
c.
10 mm (from eggs glued to
body of female illustrated by Brongniart, 1892;
Fig. 14).
Fig. 14
Bactrododema reyi
egg, A: dorsal view; B: lateral view (after
Brongniart, 1892).
AB
National Park, 24°59’41”S, 31°35’25”E, 11.ii.1966,
L. Braack (18SBRC ref. no. B598); Skukuza, Kruger
National Park, v.1993, at light, L. Braack (19SBRC);
Skukuza, Kruger National Park, 16.vii.1991, L.
Braack (19SBRC); Thabazimbi, iv.1987 (19TMSA);
Thorn Hill, Hectorspruit, 5.v.1930, H.S. Webb (19
TMSA); Tzaneen, 23°45’S, 30°10’E, 12.ii.1999, Celia
Goberz (18TMSA); VW [Vaalwater], N. Province,
24°22’S, 28°10’E, 29.iv.1996, S.J. van Niekerk (18
TMSA); Venda, Nwanedi resort, 22°38’S, 30°24’E,
550m, 5–9.ii.1994, R. Oberprieler (28SANC);
Waterberg, Augustus, 1915 (19TMSA); Waterpoort,
N. Transvaal, E.R.T. Cheales, acquired 1925 (19
BMNH); Zoutpansberg, iv.1932, Govt. Officer (19
TMSA); No data, B. tiarata (18TMSA); No data (19
SANC). BOTSWANA. Palla Road, Bechuanaland,
iv.1925, J. Reid (19TMSA); Gaberones,
Bechuanal[and] Prot[ectorate], x.1920, Terrard (19
TMSA).
ACKNOWLEDGEMENTS
I thank the following individuals for assistance and
in the case of the Transvaal Museum for the loan of
material: U. Aspöck and A. P. Kaltenbach (NHMW);
B. Dombrowsky and R. Toms (TMSA); L. Braack
(South Africa); T. Kronestedt (NHRS); G. McGavin
and D. Mann (OXUM); J. Marshall (BMNH); M. Ohl
(ZMHB); M. Picker (Zoology Department, University
of Cape Town); M. Cochrane and H. Robertson
(SAMC); I. Roesch (Hoedspruit); H. Sithole
(Scientific Services, Kruger National Park,
Skukuza); R. Urban (SANC); G. Zambatis (SBRC);
and O. Zompro (Kiel, Germany). B. Dombrowsky
kindly arranged accommodation. H. Sithole
arranged accommodation and permits. M. Burger
(TMSA) compiled the distribution map. Comments
from J. du G. Harrison (TMSA) and anonymous
referees improved this paper.
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