-» T AMERICAN MUSEUM
JSIovitates
PUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORY
CENTRAL PARK WEST AT 79TH STREET, NEW YORK, NY 10024
Number 3502, 32 pp., 45 figures, 4 tables January 12, 2006
New South African Flat Rock Scorpions
(Liochelidae: Hadogenes)
LORENZO PRENDINI 1
ABSTRACT
Two new flat rock scorpions, both endemic to South Africa, are described in the bicolor
group of Hadogenes Kraepelin, 1894: H. polytrichobothrius n.sp.; H. soutpansbergensis n.sp.
Both occupy discrete distributional ranges, allopatric with the other three species in the bicolor
group: H. bicolor Purcell, 1899; H. longimanus Prendini, 2001; H. newlandsi Prendini, 2001.
The distributions of the five species in the group are mapped, and a key provided for their
identification.
INTRODUCTION
Flat rock scorpions of the genus Hadoge¬
nes Kraepelin, 1894 represent an intriguing
group of mostly large, extremely dorsoven-
trally compressed scorpions. All described
species are obligate lithophiles, inhabiting
the narrow cracks, crevices and spaces be¬
neath exfoliations of weathered rock out¬
crops from South Africa to Tanzania. Besides
dorsoventral compression, other ecomor-
phological adaptations facilitating existence
in this specialized habitat include elongation
of the metasoma and pedipalps, perhaps to
aid with prey capture in confined spaces;
greatly enlarged lateral ocelli relative to the
median ocelli, to aid in anterior light percep¬
tion; pronounced superciliary carinae to pro¬
tect the median ocelli from abrasion; stout,
spiniform setae on the ventral surfaces of the
telotarsi, and highly curved telotarsal ungues,
to provide a vicelike grip on rock surfaces
(Newlands, 1972a, 1972b, 1978; Newlands
and Prendini, 1997; Prendini, 2001a). The
tarsal adaptations of Hadogenes facilitate lo¬
comotion on rock but hinder locomotion
across other substrata. These scorpions are
thus restricted to regions of rugged, moun¬
tainous topography and subject to allopatric
speciation when mountain ranges become
1 Division of Invertebrate Zoology, American Museum of Natural History (lorenzo@amnh.org).
Copyright © American Museum of Natural History 2006
ISSN 0003-0082
2
AMERICAN MUSEUM NOVITATES
NO. 3502
separated through erosion (Newlands, 1972a;
Prendini, 2001b). The distributional ranges
of Hadogenes species are, with few excep¬
tions, allopatric or parapatric (Newlands,
1980; Prendini, 2001a, 2005a).
Several investigations addressed the sys-
tematics of Hadogenes in recent years (e.g.,
Newlands and Prendini, 1997; Lourengo,
1999; Prendini, 2000, 2001a, 2005b). In the
present contribution, two new species are de¬
scribed in the bicolor group, which is endem¬
ic to South Africa: H. polytrichobothrius
n.sp.; H. soutpansbergensis n.sp. The exis¬
tence of both species has been known for
some time, but the paucity of material pre¬
vented their description until now. For ex¬
ample, Prendini (2001a: 158, 159) provision¬
ally assigned five specimens, designated here
as paratypes of H. polytrichobothrius n.sp.,
to H. longimanus Prendini, 2001, but sug¬
gested that they may represent an unde¬
scribed species:
[F]our specimens from Steelpoort (Lydenburg dis¬
trict), ca 100 km northeast of the northernmost local¬
ity record [of H. longimanus ] in the Groblersdal dis¬
trict and the 8 specimen from Doornkop (Carolina
district), described by Hewitt (1918), differ from the
typical form [of H. longimanus ] in several respects.
The pedipalps, especially of 8, are proportionally
shorter and broader, the carapace, post-tergites and
metasoma are slightly more granular, and the tricho-
bothrial counts are higher (total number of tricho-
bothria per pedipalp, 201—225). Morphometric ratios
of the pedipalps of a 8 and $ from Steelpoort that
differ from typical specimens are as follows: femur
length 57% greater than width in 8, 55% greater in
2; patella length 41% greater than width in 8 and
2; chela length along ventroexternal carina 39%
greater than chela width in 8, and 35% greater in 2;
length of movable finger 1 % greater than length along
ventroexternal carina in 8 , and 2% greater in 2. Un¬
fortunately, the absence of any specimens from the
area between these localities prevented an assessment
of whether this variation is continuous or discrete.
Further investigation, including the collection of ad¬
ditional material, will be required to determine if this
variation has an ecological basis, or if these speci¬
mens represent yet another cryptic species in this
complex.
Recent fieldwork in the region between
Steelpoort and the northernmost locality rec¬
ord of H. longimanus demonstrated that the
distribution of populations of Hadogenes as¬
signed to H. longimanus and those described
here as H. polytrichobothrius n.sp. is discon¬
tinuous. Morphological data obtained from
the five original specimens and fifteen addi¬
tional adult specimens (3d, 12 9) confirmed
that the differences between these species are
consistent, discrete, and diagnostic.
The collection of five additional adult
specimens (Id, 49) of H. soutpansbergensis
n.sp., for which a pair of adults (collected by
the author in 1990) were the only adult spec¬
imens previously available, has also facili¬
tated its description.
Both new species occupy discrete distri¬
butional ranges, allopatric with the other
three species in the bicolor group (fig. 1): H.
bicolor Purcell, 1899; H. longimanus ; H.
newlandsi Prendini, 2001. Recognition of the
two new species raises the number of cur¬
rently accepted species of Hadogenes to 18
(table 1).
MATERIAL AND METHODS
Personally collected specimens were lo¬
cated at night using a portable ultraviolet
(UV) lamp, comprising two mercury-vapor
tubes attached to a chromium parabolic re¬
flector and powered by a rechargeable 7 amp/
hr, 12 V battery, or by inspecting rock crev¬
ices and exfoliations during the day. A por¬
table Garmin® GPS II Plus device was used
for recording the geographical coordinates of
collection localities in the field.
Material examined is deposited in the fol¬
lowing collections: Albany Museum, Gra-
hamstown, South Africa (AMGS); American
Museum of Natural History, New York
(AMNH), some bearing accession numbers
from the Alexis Harington Collection (AH);
California Academy of Sciences, San Fran¬
cisco (CAS); Natal Museum, Pietermaritz¬
burg, South Africa (NMSA); National Col¬
lection of Arachnida, Plant Protection Re¬
search Institute, Pretoria, South Africa
(NCA); South African Museum, Cape Town
(SAMC), some bearing accession numbers
from the John Visser Collection (JV); Trans¬
vaal Museum, Pretoria, South Africa
(TMSA); Zoologisches Museum, Humboldt-
Universitat, Berlin, Germany (ZMHB); Zool¬
ogisches Institut und Zoologisches Museum,
Universitat Hamburg, Germany (ZMUH).
Tissue samples of the new species have been
stored (in the vapor phase of liquid nitrogen
at — 150°C) in the Ambrose Monell Collec-
2006
PRENDINI: SOUTH AFRICAN FLAT ROCK SCORPIONS
3
Fig. 1. Map showing the known distribution of species in the bicolor group of Hadogenes : H.
bicolor Purcell, 1899 (square), H. longimanus Prendini, 2001 (cross), H. newlandsi Prendini, 2001 (star),
H. polytrichobothrius n.sp. (triangle) and H. soutpansbergensis n.sp. (circle). Contour interval, 500 m.
4
AMERICAN MUSEUM NOVITATES
NO. 3502
TABLE 1
Currently Accepted Species of Hadogenes Kraepelin, 1894 (Scorpiones: Liochelidae), with
Countries of Distribution Compiled from Prendini (2005a)
Hadogenes bicolor Purcell, 1899
Hadogenes gracilis Hewitt, 1909
Hadogenes granulatus Purcell, 1901
Hadogenes gunningi Purcell, 1899
Hadogenes hahni (Peters, 1862)
Hadogenes lawrencei Newlands, 1972
Hadogenes longimanus Prendini, 2001
Hadogenes minor Purcell, 1899
Hadogenes newlandsi Prendini, 2001
Hadogenes paucidens Pocock, 1896 a
Hadogenes phyllodes Thorell, 1877 b
Hadogenes polytrichobothrius n.sp.
Hadogenes soutpansbergensis n.sp.
Hadogenes tityrus (Simon, 1888) b
Hadogenes trichiurus (Gervais, 1843) b
Hadogenes troglodytes (Peters, 1861)
Hadogenes zuluanus Lawrence, 1937
Hadogenes zumpti Newlands and Cantrell, 1985
South Africa
South Africa
Botswana, Mozambique, Zambia, Zimbabwe
South Africa
Angola, Namibia
Na mi bia
South Africa
South Africa
South Africa
Democratic Republic of Congo, ?Tanzania
Namibia, South Africa
South Africa
South Africa
Namibia, South Africa
South Africa
Botswana, Mozambique, South Africa, Zimbabwe
South Africa, Swaziland
Namibia, South Africa
a Species of dubious validity.
b Species complexes.
tion for Molecular and Microbial Research
(AMCC) at the AMNH.
Photographs were taken in visible light as
well as under long wave UV light using a
Microptics® ML 1000 digital imaging sys¬
tem. Measurements were recorded with Mi-
tutoyo® digital calipers and counts taken us¬
ing a Nikon® SMZ-1500 stereomicroscope.
Color designation follows Smithe (1974,
1975, 1981), trichobothrial notation follows
Vachon (1974), and sternum terminology fol¬
lows Soleglad and Fet (2003). Morphological
terminology and mensuration follows previ¬
ous papers on Hadogenes (e.g., Newlands
and Prendini, 1997; Prendini, 2000, 2001a,
2005b).
A distribution map was produced using
ArcView GIS Version 3.2 (Environmental
Systems Research Institute, Redlands, CA),
by superimposing point locality records on
coverages depicting the topography (contour
intervals of 500 m) and political boundaries
of southern Africa. A topographic contour
coverage was created from the GTOPO30
raster-grid coverage, obtained from the web¬
site of the U.S. Government Public Infor¬
mation Exchange Resource: http://
edcdaac .usgs. gov/gtopo3 0/gtopo3 0 .html.
All records of sufficient accuracy were
isolated from the material examined to create
a point locality geographical dataset for map¬
ping distributional ranges. Only a small pro¬
portion of the records were accompanied by
geographical coordinates or quarter-degree
squares (QDS), usually entered by the col¬
lector or subsequently added by the curator
or collection manager. These were checked
for accuracy and an attempt was made to
trace coordinates for as many of the remain¬
ing records as possible, by reference to gaz¬
etteers, the official 1:250,000 and 1:500,000
topocadastral maps of South Africa pub¬
lished by the Government Printer, and the
GEOnet Names Server: http://164.214.2.59/
gns/html/cntry_hles.html. Names of South
African provinces and magisterial districts
listed in the material examined follow the
most recent system (post-1994).
Spatial analyses were conducted using
ArcView, in order to ascertain whether the
distribution of the new species is related to
present environmental variables, to deter¬
mine the specific ecological correlates of
their distributional ranges, and to calculate
statistics that could be used to define their
conservation status. Coverages representing
the topography, mean annual rainfall, bi-
omes, and vegetation types in South Africa
2006
PRENDINI: SOUTH AFRICAN FLAT ROCK SCORPIONS
5
were used for these analyses. The GTOPO30
raster-grid coverage was used for spatial
analysis of topography. A raster-grid cover¬
age of mean annual rainfall was obtained
from the South African Atlas of Agrohydrol¬
ogy and -climatology (Schulze, 1997), pro¬
duced by the Computing Centre for Water
Research of the South African Water Re¬
search Commission: http://www.wrc.org.za/
wrcpublications/wrcreports/hydroclimatology.
htm. A polygon coverage of Low and Re-
belo’s (1998) Vegetation of South Africa, Le¬
sotho and Swaziland, incorporating the bi-
omes defined by Rutherford and Westfall
(1994), was obtained from the National Bo¬
tanical Institute of South Africa.
The new species of Hadogenes were cat¬
egorized according to their conservation pri¬
ority using coverages representing national
and provincial protected areas in South Af¬
rica, obtained from the Western Cape Nature
Conservation Service of South Africa, su¬
perimposed on the mapped distributional
ranges to determine whether any point local¬
ity records fell within the boundaries of pro¬
tected areas. Both species were then assigned
one of the IUCN Red List Categories (IUCN,
2001) on the basis of the number of known
locality records, extent of the distributional
range, occurrence inside and outside of pro¬
tected areas, and prevailing land uses that
might be construed as threats to their future
survival.
SYSTEMATICS
FAMILY LIOCHELIDAE FET AND BECHLY, 2001
GENUS HADOGENES KRAEPELIN, 1894
Key to the identification of species in the
bicolor group of Hadogenes.
1. Pedipalp chela with pronounced lobe, distal
to notch in fixed finger; metasoma 51-57%
of total length (d); metasomal segment V,
lateral surfaces, sparsely granular to
smooth, telson smooth (d) .2
Pedipalp chela without pronounced lobe, dis¬
tal to notch in fixed finger; metasoma 57-
68% of total length (<S); metasomal seg¬
ment V, lateral surfaces, and (usually) tel¬
son densely granular (d).4
2. Pedipalp chela with 2 trichobothria in i series
. Hadogenes bicolor
Pedipalp chela with 5-8 trichobothria in i se¬
ries .3
3. Pedipalp chela width 46-55% of length along
ventroexternal carina (d), 59-60% (9);
lobe on chela movable finger (<3) fitting
evenly into corresponding notch in fixed
finger, i.e. not overlapping when fingers
closed; telson same color as metasomal
segments . Hadogenes longimanus
Pedipalp chela width 60-73% of length along
ventroexternal carina (6, 9); lobe on chela
movable finger (d) fitting unevenly into
corresponding notch in fixed finger, i.e. dis¬
placed to external surface and overlapping
when fingers closed; telson pale in com¬
parison with metasomal segments.
. Hadogenes polytrichobothrius
4. Pedipalp chela width 60-69% of length along
ventroexternal carina (<3, 9); patella width
57-64% of length; femur width 43-48% of
length; metasoma 57-58% of total length
(d), 48% (?) .... Hadogenes newlandsi
Pedipalp chela width 49-53% of length along
ventroexternal carina (6), 57-58% (9); pa¬
tella width 51-56% of length; femur width
33-38% of length; metasoma 62-68% of
total length (<?), 52-54% (9) .
. Hadogenes soutpansbergensis
Hadogenes bicolor Purcell, 1899
Hadogenes bicolor Purcell, 1899: 437-438.
Hadogenes bicolor: Lawrence, 1955: 251 (part);
Lamoral and Reynders, 1975: 538 (part); New-
lands, 1980 (unpublished): 99-105 (part), figs.
48 (part), 49-53; Newlands and Cantrell, 1985:
40, 42, 44 (part); Kovarrk, 1998: 132; Fet,
2000: 387; Prendini, 2001a: 149-156, figs. 1-
10, 33, 36, tab. 2; Prendini, 2001b: 137; Pren¬
dini, 2005a: 67, appendix 1.
Type Material: SOUTH AFRICA: Lim¬
popo Province: Pietersburg District : Lec-
totype S (SAMC 4062), 20 miles east of Pe¬
tersburg [23°54'S 29°47'E], 1899, J.W. Da-
neel. Paralectotypes: same data as lectotype,
29 1 subad. 9 1 juv. S 1 juv. 9 (SAMC
4062).
Additional Material: SOUTH AFRI¬
CA: Limpopo Province: Letaba District :
Serala Wilderness Area, near Tzaneen
[24°00'S 30°04'E], 30.viii.1980, M. Stiller,
ex flat rocks, under overhang on steep moun¬
tainside, grass, rocks, 19 (SAMC C1602), 1
juv. 9 (SAMC 0613). Lekgalameetse Na¬
ture Reserve: Haffenden Heights [Farm Haf-
fenden Heights 35], The Downs, 24°07'S
30°07'E, 26.vi.1977, B.P.W. Fratscher, 69
(TMSA 17794, 17795, 17797-17800); Leop-
6
AMERICAN MUSEUM NOVITATES
NO. 3502
ard’s Crag, 50 km W of Haffenden Heights
[24°09'S 30°13'E], I.H. Davidson, 19
(TMSA 17449); same data, except
“9.vi.l977, I.H. Davidson”, Id (TMSA
18004), 19 (TMSA 18005). Phalaborwa
District : Farm Lillie, near Mica, 24°02'S
30°50'E, l.xii.1977,1. Davidson, 19 (TMSA
17147 [old 1258]); Hoedspruit, 32 km W
[24°23'S 30°40'E], 19 (AMNH [AH 4301]);
Jongmansspruit, on Blyde River, near Swa-
dini [24°30'S 30°47'E], 3-8.i.l999, I. Engel-
brecht and D. Eagan, in crevices in granite
rocks, 1 S 19 (SAMC C4585); Peninsula
trail, Blyderivierspoort Dam, Blyde River
Canyon Nature Reserve, 24°33'S 30°48'E,
13.vii.2000, L. Prendini, M. MacFarlane, and
K.M.A. Prendini, mixed bushveld, crevices
in quartz, 1 S 1 9 1 subad. S (AMNH), 1 juv.
9 (AMCC 138994). Pietersburg District:
Pietersburg, 19 mi E [23°54'S 29°47'E],
26.xii.1967, G. Newlands, 2 juv. S (TMSA
17456, 17459 [old 770]), 1 juv. 9 (TMSA
17457 [old 769]). Sekhukhuneland District:
Burgersfort, ca. 20 km E on R555 to Oh-
rigstad, 24°34'S 30°30'E, 5.L2005, I. Engel-
brecht, 1255 m, ridges with valleys, broad-
leaf savanna, moderate N slope, 1 9 (AMCC
144144); Farm Perkoe [Perkeo] near Olifants
River [24°28'S 30°36'E], xi.1927, H. Lang,
1 juv. S (TMSA 6086); Penge, 24°20'S
30°25'E, 1980, N. du Toit, 19 (AMNH [AH
1251]). Mpumalanga Province: Belfast
District : Patattanek, ca. 1.5 km E on R539
[25°29'S 30°22'E], ii-x.2003, A. Mathie, un¬
der stones and small rocks on large flat rocks,
1 juv. 9 (AMCC 138993). Lydenburg Dis¬
trict: 1879, T. Ayres, 1 dry specimen [lost,
not examined] (SAMC 432); Lydenburg, 31
km N, on the road to Burgersfort [24°49'S
30°21'E], 30.xii.2002, I. Engelbrecht, 1 juv.
9 (AMCC 138991); Ohrigstad, 1 km NE
[24°45'S 30°34'E], 4.viii.l984, G. Newlands,
2 subad. 9 (AMNH [AH 3449, 3450]), 2 juv.
9 (AMNH [AH 3451]); same data, except
“21.viii.1984, C. Owen”, 1 juv. S (AMNH
[AH 3787]), 1 juv. 9 (AMNH [AH 3788]);
Ohrigstad, 4 km NE, 6 km from turnoff
[24°43S 30°32'E], 6.x. 1984, C. Owen, IS
(AMNH [AH 3577]), 19 (AMNH [AH
3574]); Farm Klipheuvel 549, 1 km S R533
turnoff to Pilgrim’s Rest from R36 Lyden¬
burg to Ohrigstad, on Verraaier’s Nek
[24°54'S 30°34'E], 14.xii.2003, A. Mathie,
1326 m, crevices, 19 (NCA Ac AT 2005/
595). Pilgrim’s Rest District: Blyde River,
Lydenburg [24°38'S 30°47E], 14.i.1971,
N.H.G. Jacobsen, IS (TMSA 10100); Blyde
River Canyon Nature Reserve [24°35'S
30°49'E], 8.v.1974, N.H.G. Jacobsen, 1 juv.
S (TMSA 12515); Bourke’s Luck Potholes,
Blyde River Canyon Nature Reserve,
24°40'S 30°49'E, 12.vii.2000, L. Prendini
and M. MacFarlane, grassland, with mixed
bushveld at edge of canyon, under sandstone,
19 (AMNH), 2 juv. 6 2 juv. 9 (AMCC
138992); Dientje G.M., Vaalhoek, near Pil¬
grim’s Rest [24°39'S 30°47'E], Miss S. Prell-
er, IS (AMGS); Dientje P.O., Vaalhoek
[24°43'S 30°47'E], S. Preller, 1 subad. 9
(AMGS 4704).
Hadogenes longimanus Prendini, 2001
Hadogenes bicolor Purcell, 1899: Hewitt, 1918:
160, 161 (part), pi. 30, figs. 88, 89; Newlands,
1980 (unpublished): 99-105 (part), fig. 48
(part); Newlands and Cantrell, 1985: 40, 42, 44
(part).
Hadogenes longimanus Prendini, 2001a: 156—
159, figs. 1, 11-21, 34, 37, table 2; Prendini,
2001b: 137.
Hadogenes longimanus: Prendini, 2005a: 67, ap¬
pendix 1.
Type Material: SOUTH AFRICA:
Mpumalanga Province: Groblersdal Dis¬
trict: Holotype S (SAMC C4602), 20 km S
of Groblersdal on road to Middelburg,
25°20.30'S 29°22.85'E, 13.1.2000, L. Pren¬
dini and I. Engelbrecht, 1077 m, mixed bush¬
veld, crevices in granite rocks. Paratypes:
same data as holotype, 2d 29 2 subad. 9 1
juv. S (SAMC C4603). Bronkhorstspruit
District: Bundu Inn, on road from Bronk¬
horstspruit to Groblersdal [25°29'S 29°01'E],
18.xii.1980, M. Stiller, on hill, under large
granite rock lying on rock face, many milli¬
pede (Juliform) and beetle remains, 19
(NMSA 13931); same data, except
“20.xii.1980”, 19 (SAMC C1600); Farm
B oekenhoutskloofdrift 286 [25°18'S
29°01'E], 20.ix. 1982, E. Voigt, Id (TMSA
12507). Middelburg District: 55 km S of
Groblersdal on road to Middelburg,
25°32.27'S 29°28.67'E, 134.2000, L. Pren¬
dini and I. Engelbrecht, 1509 m, grassland
and mixed bushveld, crevices in sandstone,
Id 59 1 juv. d (SAMC C4600), Id 19
2006
PRENDINI: SOUTH AFRICAN FLAT ROCK SCORPIONS
7
(CAS), 2 juv. 9 (AMCC 138995, 138996);
Farm Noupoort 16, Selons River [25°25'S
29°28'E], 1933, 2d (TMSA 17458, 17513);
Fort Merensky, Botshabelo Nature Reserve,
25°41.82'S 29°24.87'E, 144.2000, L. Pren-
dini and I. Engelbrecht, 1410 m, grassland,
with mixed bushveld along banks of Olifants
River, under flat stones and in crevices (sand¬
stone), 26 129 (SAMC C4601), 16 19
(AMNH). Witbank District: 2-D Ranch [Los-
kop Dam Nature Reserve], 25°22.101'S
29°18.409'E, x.1989, L. Prendini and M.R.
Filmer, 1070 m, in crevices, 29 (SAMC
C4596); same data, except “x.1994, I. Eng¬
elbrecht”, 19 ljuv. 9 (SAMC C4595), 19
(SAMC C4598); same data, except “N.
MacLean”, 19 (SAMC C4599); same data,
except “J. Laing”, 1 subad. 6 (SAMC
C4594); Amaphi Nature Reserve, on road
from Loskop Dam to Verena [25°21.66'S
29°18.69E], 144.2000, L. Prendini and I.
Engelbrecht, 1102 m, mixed bushveld, in
crevices in granite, 19 1 subad. 6 (SAMC
C4597). Gauteng Province: Bronkhorst-
spruit District : Farm Zusterstroom 447,
25°35'S 29°01'E, 4.xi.l977, G. Newlands,
Id (TMSA 17452), 29 (TMSA 17453,
17460).
Additional Material: SOUTH AFRI¬
CA: Gauteng Province: Bronkhorstspruit
District : Farm Zusterstroom 447, ca. 11 km
E Vaalplaas turnoff on R25 from Bronk¬
horstspruit to Groblersdal, 25°36.290'S
28°59.875'E, 19.X.2004,1. Engelbrecht, 1235
m, ridges with valleys, broadleaf savanna on
red loamy soil with grass and litter layer,
moderate S slope, 19 1 subad. 6 1 juv. 9
(NCA AcAT 2005/504); Renosterpoort
[Farm 498, 25°45'S 28°57'E], 10 km E of
Bronkhorstspruit, l.iv.1984, G. Newlands,
19 (AMNH [AH 3350]). Mpumalanga
Province: Witbank District : Ezemvelo Na¬
ture Reserve, 25°42'S 29°0UE, 26-
304.2004, K. Manamela, 1 subad. 6 (TMSA
22859); Ezemvelo Nature Reserve, 25°43'S
28°57'E, l.iii.2005, GDACE, 1 subad. 9
(NCA AcAT 2005/513); Marulani Lodge,
just S Loskop Dam, 25°26.366'S
29°24.517'E, 3.X.2004, I. Engelbrecht, 1050
m, sympatric with Parabuthus transvaalicus
Purcell, 1899, 1 subad. 6 (NCA AcAT 2005/
506).
Hadogenes newlandsi Prendini, 2001
Hadogenes bicolor Purcell, 1899: Hewitt, 1918:
160, 161 (part); Lamoral and Reynders, 1975:
538 (part); Newlands, 1980 (unpublished): 99-
105 (part), fig. 48 (part); Newlands and Can¬
trell, 1985: 40, 42, 44 (part).
Hadogenes newlandsi Prendini, 2001: 159, 162-
169, figs. 1, 22-32, 35, 38, tab. 2; Prendini,
2001b: 137.
Hadogenes newlandsi: Prendini, 2005a: 67, ap¬
pendix 1.
Type Material: SOUTH AFRICA: Lim¬
popo Province: Soutpansberg District: Ho-
lotype 6 (SAMC C4589), Ben Lavin Nature
Reserve, 23°07.544'S 29°56.573'E,
31.xii.1999, L. Prendini and E. Scott, 850 m,
in crevices in granite rocks, mixed bushveld.
Paratypes: same data as holotype, 3d 5 9 1
subad. 9 (SAMC C4593), Id 19 (AMNH);
Tabajwane Koppie, Ben Lavin Nature Re¬
serve [23°09'S 29°58'E], xii.1990, L. Pren¬
dini and K.M.A. Prendini, 970 m, in crevices
in granite rocks, Id 19 (SAMC C4588);
Bandelierkop, 23°18'S 29°51'E, iv.1988, L.
Prendini, M.R. Filmer, A.M. Smith and V.
Hull-Williams, in crevices in granite koppie,
2 9 (SAMC C4587); same data, except
“1995, I. Engelbrecht”, Id 19 1 juv. d
(SAMC C4586); Louis Trichardt, 10 mi S
[23°08'S 29°54'E], 25.iii.1958, E.S. Ross and
R.E. Leech, 1000 m, 1 subad. 6 (CAS);
same data, except “Louis Trichardt, 18 mi S
[23°17'S 29°49'E]”, 19 3 juv. d 1 juv. 9
(CAS); Mailaskop, 23°13.43'S 29°56.63'S,
30.xii.1999, L. Prendini and E. Scott, 1124
m, in crevices in dolerite rocks on koppie,
19 2 juv. 9 (SAMC C4590). Letaba Dis¬
trict: Letsitele, Tzaneen [23°53'S 30°24'E],
2Fix. 1964, R.D. Faul, 1 subad. 6 (TMSA
12565); Mooketsi [23°36'S 30°06'E],
iv.1924, G.P.F. van Dam, 2d (TMSA 114,
117), 109 (TMSA 112, 113, 115, 116, 118—
122, 125), 2 juv. 9 (TMSA 127-128). Pe¬
tersburg District: Clearwaters, Haenertsburg
[23°51'S 29°57'E], 44U916, G.A. Thomp¬
son, 1 subad. d (TMSA 1057); Farm Mun-
niks [23°37'S 29°57'E], 164.1914, Pienaar, 1
subad. 9 (TMSA 1058); Pietersburg area
[23°54'S 29°27'E], 1 9 (AMNH); Woodbush
[23°47'S 29°54'E], xii.1907, D. Gough, 19
(TMSA 1055); The Woodbush, 1 subad. 6
(AMGS 3990). Potgietersrus District: Ma-
kapan Caves [24°09'S 29°11'E], 441.1911, A.
AMERICAN MUSEUM NOVITATES
NO. 3502
©
©
04
©
S3
<u
Sh
s
Is
S ©
ar:
i-
CS o
- 04
iH '—'
© •«
© S3
•3 S3
M 2
0> ©
> 3
CLh ei
K /—s
2 r-
s ©
Q ©
II
■S'g
sn' 3
on
on 12
00 5
rH St
s*
£1
V. £
£ £
11
"5 o
2 ^
& s
J?s
Si
S
3
l §
t-
.©
3
Q
S-
©
u
co
0
©
©
<N
©
©
©
©
pt
04
CO
pt
00
t
00
Pt
00
00
00
©
pt
987
Pt
©
'B
0 +
%
<
©
in
V I"
2
c3
©
©
00
NO
NO
NO
<N
in
00
oi
NO
l>
NO
l>
2/2
CO
CN
co
CO
to
©
04
in
in
in
Pt
04
CO
©
CO
CO
Pt
©
NO
in
>n
0 ^
00
cd
CO
©
On
cd
04
cd
00
04
©
pt
©
■3
u
CO
CO
CO
©
04
O'
pt
3
sp
00
(X
u
N O
O'
cn
m
©
(N
<N
©
©
Pt
©
Pt
00
,_l
04
04
©
©
co
O'
00
00
pt
7.26
©
04
04
"3
0 +
%
<
00
in
ctf
S-H
C3
Pt
On
NO
in
in
in
't
©
00
pf
pt;
00
CN
in
in
pt
2/2
CO
CO
CO
CO
NO
06
in
pt
in
co
in
04
CO
©
Pt
NO
l>
CO
Pt
0 ^
04
04
Pt
04
Pt
On
NO
cd
00
04
©
04
Pt
cd
CO
©
s
u
Dm
CN
CO
CO
©
CO
©
Pt
-S
2
00
to
u
v.D
04
m
r-H
in
©
©
©
CN
©
0
O'
©
©
©
00
©
©
O'
©
00
00
in
Pt
©
00
s
%
<
00
00
cd
On
^t
00
(N
rn
NO
in
in
CN
CN
CO
CO
©
©
CO
in
in
NO
Pt
04
©
©
©
00
04
00
©
00
&5
*0
in
1 1
c5
00
^t
^t
On
pf
NO
CO
CO
CN
©
00
Pt
CO
00
©
NO
CO
CO
©
©
©
©
pt
04
©
04
cd
cd
to
e
to
00
u
Pm
CN
CO
CO
Pt
04
©
Pt
.2
O
On
©
<n
m
(N
CO
m
m
CO
Pt
in
©
00
©
©
00
00
Pt
©
04
00
©
CO
,_ 1
©
pt
04
co
04
5
H
S
00
pt
(N
hH
CN
On
©
CJ
co
CO
to
O;
p-
04
l>
04
0 ^
©
©
04
00
1—1
NO
04
©
00
04
ttj
*o
in
O
On
in
NO
©
>n
r-H
NO
NO
cl
0
CO
06
Pt
in
3h
©
ND
Pt
in
00
©
04
©
pt
cd
04
cd
pt
NO
<
00
u
CO
CO
CO
Pt
04
©
©
ffi
2
(D
m
©
0
©
Pt
00
00
00
©
c^-
©
s
00
04
©
O'
00
©
©
04
©
CO
©
'3
0 +
£
©
00
r--
©
CO
in
CO
m
CO
CO
©
00
04
©
CO
©
©
00
0 ^
O'
Pt
NO
Pt
04
-3
§
03
P4
06
(N
ri
00
pf
in
CO
CN
NO
©
00
CO
04
Pt
CO
in
CO
CO
NO
NO
©
0 ^
04
04
in
04
cd
0 ^
s
3
<
’ 1
1—1
<N
1—1
1—1
CO
CO
1—1
©
Pt
1—1
©
CO
’ 1
(X
u
CO
00
't-
©
©
CO
©
00
CO
00
©
©
04
©
CO
©
Pt
©
©
©
©
Pt
©
00
O'
Pt
65
a
a
0 +
%
<
0
NO
1 1
ctf
c3
Pt
On
in
in
cn
in
r-
On
©
>n
NO
CO
NO
CN
NO
5/5
0
CO
04
00
06
NO
Pt
in
in
©
04
CO
CO
CO
NO
©
Pi¬
in
NO
©
©
04
04
04
04
NO
00
©
cd
©
04
in
NO
in
04
pt
cd
Pt
06
«
u
Ph
CO
Pt
Pt
©
CO
O'
Pt
00
Sc
a
X
©
©
©
m
©
Pt
CN
C"
©
©
©
00
pt
CO
CO
©
co
l>
O'
O'
©
©
CO
00
pt
co
Pt
-2
*o
03
&-H
O'
cn
©
■^t
On
00
l>
NO
CO
Pt
in
co
©
CO
in
00
©
©
Pt
Pt
CO
pt
>n
§
<
C3
0 ^
ci
ci
CO
in
CO
CN
©
co
04
pf
in
CO
CO
06
0 ^
©
00
cd
04
NO
04
cd
00
§
§>
Ph
CN
CO
©
CO
©
CO
0
04
O'
©
t"-
(N
©
in
©
©
©
O'
©
©
04
©
©
©
©
©
©
©
©
00
O'
co
pt
"a
S
<
00
0
jD
©
00
in
NO
>*H
Pt
hH
m
CO
co
©
in
©
©
CO
in
hH
hH
CO
CO
H
©
pt
H
in
00
O';
04
£
*0
NO
1 1
O
On
>n
in
00
in
Pt
00
00
NO
of
On
pt
0^
©
oc
NO
Pt
©
©
Pt
04
On
cd
cd
06
04
cd
04
•U
u
43
CO
CO
CO
©
04
04
O'
Pt
<
in
(N
00
©
©
00
(N
m
©
00
©
CO
Pt
©
©
Oh
O'
©
©
CO
pt
00
co
00
pt
©
0 +
00
0
00
©
(N
©
in
H
i>
l>
CJ
CO
CO
04
H
04
n
04
in
04
©
Pt
00
©
in
04
©
pt
04
04
—
2
'—*
00
06
ON
pf
On
in
Pt
ol
Pt
00
Pt
Pt
co
©.
in
CO
Pt
s
06
©
06
cd
04
NO
04
cd
0^
33
H
1-1
<N
1-1
CO
CO
©
04
1-1
CO
l -1
In
P
u
O
Oh
(NJ
O
00
0
in
00
©
pf
in
©
©
©
©
©
©
©
pt
©
Pt
©
,_1
©
r-H
00
00
CO
©
O'
0 +
vo
Jh
04
©
cn
>n
<N
CO
NO
CO
CO
00
©
hH
uo
CO
NO
CO
©
iH
O'
04
04
t-H
04
00
04
-2
<
00
O
00
rn
cn
00
Pt
pt
CO
cl
nB
04
0 ^
Pt
CO
©
in
CO
Pt
©
NO
©
00
cd
04
NO
04
04
0^
"0
w
Ctf
1 —H
H
CN
I —<
CO
Pt
H
©
CO
©
CO
•2
a
~C)
<
s
04
in
<T)
00
©
©
00
©
m
O'
©
©
04
00
©
pt
00
04
O'
©
O'
co
©
©
©
©
©
65
sj
1/5
1-1
00
(T)
!"•
©
CN
CN
CO
CO
O'
00
©
CO
CO
NO
04
00
NO
©
©
CO
0^
Pt
Pt
'■H
©
in
a
*o
s
©
00
cn
<ri
I>
CO
in
CO
CN
in
0^
CO
CO
04
in
CO
in
CO
cd
On
0^
04
04
U
04
04
On
to
00
CN
CO
CO
Pt
CO
©
co
H
1—1
JS
u
CM
NO
0
Pt
00
©
©
©
©
Pt
©
CO
00
©
00
O'
00
©
©
Pt
00
CO
O'
00
©
O'
00
04
EC!
S
<
0
in
in
(N
00
NO
00
04
CO
04
00
04
©
CO
04
©
in
CO
04
NO
00
04
Pt
pt
04
CO
in
*0
0
0 ^
ci
<N
CO
in
CO
CN
CN
©
0 ^
CO
CO
0 :
p 2
in
04
pt
00
Pt
00
0^
cd
04
04
cd
On
Pt
<D
r^H
CN
H
hH
CO
CO
©
CO
hH
©
CO
00
P
fl
"w
Sh
P
O
'p
c
0
5h
0
00
p
©
W)
1
©
04)
•n
©
04)
1
’W)
1
’W)
1
QJ
c
p
©
1
©
X
0
0
©
<D
©
-S
£
<L)
0
3
©
©
©
©
"ol)
3
3
jp
©
op
a
p
>
cd
p
p
©
04)
p
3
04)
©
3
04)
3
04)
2
0
■3
B
-a
£
Sh
s
'3
s
p
0
>
4h
2 'S
S 6
«« 0
"C
©
O
©
■y
©
0
©
s
'©
©
a
*c
0
X)
•c
©
0
X)
B
a
'©
©
3
04)
3
04)
B
a
'©
a
B
a
'©
a
0
in
i-5
©
rH
&
0
‘C
O
s
p
e
p
e
■3
O
*3
0
3
O
©
O
'B
0
©
0
0
©
0
p
e
p
3
0
-g
0
©
©
p
3
p
s
^©
P
in
3
3
3
3
3
3
3
3
£
X
0
'o
S
P
©
(-H
CZ)
M
P
- x
p
'x
p
00
p
00
p
00
p
B
‘C
X
p
p
00
p
'3
•3
p
'x
p
04)
C
"p
'p
©
04)
c
p
p
04)
3
p
p
04)
3
00
U
£
M
3
VV
Ph
O
S
3
0
0
0
ki
s
s
©
s
s
©
O
0
B
©
s
3
©
3
3
©
■ •
HH
Specimen:
Carapace:
Chela:
Patella:
Femur:
Pedipalp:
Mesosoma:
Sternite VI
Metasoma
Metasoma
Hadogenes bicolor Purcell Hadogenes longimanus Prendini Hadogenes newlandsi Prendini
2006
PRENDINI: SOUTH AFRICAN FLAT ROCK SCORPIONS
u
CO
ON
CO
ON
CO
m
l>
CO
o
r-'
CN
CN
CN
o
CN
00
NO
<
ON
VO
2
3
9
cn
o
st
o
o
in
CN
0-
cd
o
9
CN
9
cd
9
2.9
in
cd
9
CN
CN
On
Os
On
st
9
00
as
vd
st
a
t-H
i-H
in
CN
00
U
u
ND
CO
CN
m
st
On
00
l>
t>
St
CO
st
O
m
CN
o
VO
<
00
3
VO
vo
co
CN
On
CD
9
m
CN
as
Os
t>
vo
9
t-H
VO
3
CN
cd
On
CN
CN
o
CN
CN
o
CN
CN
q
00
st
st
t>
vd
00
st
a
m
00
U
1-1
u
VO
m
co
CO
CO
vo
00
CN
t>
vo
On
cn
vO
O
00
<
00
3
l>
oo
9
CO
CN
st
CN
On
00
st
CN
9
9
9
st
t—i
nj
VO
&
CN
cd
St
CN
cd
vd
CN
CN
vd
CN
cd
00
00
st
00
00
cd
st
a
t-H
l>
cn
CN
GO
U
o
Ov
CN
CO
ON
CO
00
co
CN
VO
ON
in
00
CO
00
On
l>
>n
CN
00
j0
ON
vo
CN
i>
00
st
>n
CN
as
9
o
st
CN
is
CN
INI
VO
1 1
o
cn
St
NO
CN
cd
U
CN
cd
U
CN
cd
q
o
ON
vO
o
o
q
43
00
in
t-H
CN
00
U
PC
2
m
c
00
CO
CN
On
1^-
in
in
St
st
in
st
On
vo
<
i—i
o
CN
o
i>
in
O)
CO
9
O
m
C^
O
ON
00
>n
t-H
S3
a
cn
cd
S
CN
ni
S
CN
CN
CN
vo
CN
st
o
ON
vd
vd
VO
U
CO
st
(N
CO
co
00
r-
o
i>
O
CN
VO
CN
in
On
st
On
9
3
st
st
i>
st
On
CN
9
00
m
in
00
On
-H
CN
st
cn
^H
o
3
CN
cd
00
CN
CN
On
CN
CN
o
CN
CN
q
00
q
ON
00
ON
<
st
a
m
o
00
U
X
r-
ON
o
CN
00
m
l>
o
CN
st
CN
in
00
CN
3
co
o
CN
t-H
9
On
Ov
in
vo
CO
00
9
t—1
00
00
cn
St
CN
*°i
%
CN
cd
ON
CN
CN
On
CN
o
CN
CN
q
q
q
On
00
<N
a
in
o
CN
<
u
CN
On
00
On
vO
>n
CN
On
On
On
C--
r-.
l>
00
o
O
ON
CO
<,
O
jD
VO
>n
9
9
-h
st
st
9
00
co
o
c~
t—i
CN
CN
o
1 1
o
CN
cd
CN
CN
cd
cd
CN
cd
st
CN
CO
CN
ON
o
o
q
o
id
U
o
CO
CN
00
<
VO
St
CO
CN
CO
c~
CO
in
CO
vo
ON
cn
ON
VO
vo
00
O
in
On
in
O
CN
vo
St
00
00
>n
vo
9
9
CN
H
O
c
00
o
cn
cd
t>
CN
ci
l>
CN
00
CN
CN
fd
5
On
S
vd
id
f\)
d
o
St
CO
st
CN
CN
CN
00
,-H
O
CN
vO
st
o
cn
vo
Of ^
VO
o
00
CO
o
t"
q
o
st
00
CN
vo
9
in
o
in
CN
o
q
<
00
”3
S3
CN
CN
l>
CN
CN
00
CN
CN
00
CN
CN
vd
st
st
On
S
vd
VO
a
<
s
O
00
00
O
t>
CN
00
On
00
On
00
00
st
st
CN
CO
O
oo
00
On
On
00
in
°°
9
o
t-H
9
On
9
Os
t-H
CN
o
CN
CN
On
q
CN
o
CN
CN
CN
q
o‘
vo
cd
00
00
Q
00
in
o
CN
H
o
(N
so
o
ON
VO
VO
VO
st
m
vO
On
00
i>
st
l>
m
On
CO
00
o
o
CO
ON
On
00
m
o
IN
1-H
CO
in
00
r->
st
st
CN
vo
I-H
d
CN
CO
On
rH
CN
o
CN
CN
:
CN
CN
q
i>
vo
cd
00
S
On
<c
1)
t-H
in
o
00
3
'&
cd
3
e
E
3
43
£
43
£
£
£
£
Q
i x>
o
05
3
3
3
13
00
3
00
|
■3
%
op
'3
43
V
W)
'S
43
'%
op
'3
43
%
op
'3
43
50
3
0
S
+
cd
s
o
yi
o
3
o
S
e
E
s
s
s
s
s
O
00
00
B
W)
jP
3
u
3
3
3
3
3
3
3
CO
3
3
3
3
cd
2
3
O
d
3
Q
s
'i
e
!
d
s
E
E
E
B
E
E
O
2h
s
o
6
2h
3
Sex:
Coll
9
00
9
X
M
9
w
9
X
3
3
■J'.
+
3
00
S3
cd
3
3
cd
3
3
cd
3
3
cd
— 5
o
3
d
z
H
s
E
0)
E
S
CD
s
s
CD
s
s
3
O
O
a
O
3
o
B
>
>
3
3
3
3
OJ)
s
S
6
e
s
3
co
a
o
o
o
y:
3
o
od
3
3
3
o
B
B
B
o
y±
3
B
<D
3
3
(0
3
3
Dh
C/3
s
s
s
e2
P
Ph
a Measured from base of condyle to tip of fixed finger.
b Sum of metasomal segments I—V and telson.
10
AMERICAN MUSEUM NOVITATES
NO. 3502
Roberts, 1 juv. 9 (TMSA 708); Makapansgat
[24°09'S 29°11'E], I.H. Davidson, 19
(TMSA 17451), 1 juv. 9 (TMSA 17450),
same data, except “31.viii.1973, R. Clark”,
1 juv. d (TMSA 10781); Makapansgat
World Heritage Site, iv.2000, I. Engelbrecht,
19 (AMCC 138997); Maribashoek [24°13'S
29°08'E], xii.1924, G.P.F. van Dam, 1 juv. 9
(TMSA 6108); Percy Fife Nature Reserve
[24°02'S 29°11'E], ll.v.1972, N.H.G. Jacob¬
sen, Id (TMSA 10484), 3 9 (TMSA 10481-
10483), juv. d (TMSA 10485); Potgietersrus
Nature Reserve [24°09'S 28°59'S],
ll.v.1972, N.H.G. Jacobsen, 1 subad. 9
(TMSA 20393); Potgietersrus [24°11'S
29°01'E], 27.iii.1919, H.B. Pretorius, 1 juv.
9 (TMSA 2184). Sekgosese District : Fouis
Trichardt, 35 mi S [23°31'S 29°43'E],
26.iii.1958, E.S. Ross and R.E. Feech, 1000
m, 2 juv. 9 (CAS); Mphakane, south, granite
koppies 1 km from turnoff to Munnik,
23°32.20'S 29°42.42'E, 29.xii.1999, F. Pren-
dini and E. Scott, 1000 m, in crevices in
rock, 19 1 juv. 9 (SAMC C4591); St. Bren¬
dan’s Catholic School, Mission Matok,
23°25.63'S 29°43.28'E, 29.xii.1999, F. Pren-
dini and E. Scott, 980 m, mixed bushveld,
granite outcrops, in crevices, 2d 9 9 1 juv.
d 1 juv. 9 (SAMC C4592), Id 19 (CAS),
1 juv. d 1 juv. 9 (AMCC 138998).
Additional Material: SOUTH AFRI¬
CA: Limpopo Province: Pietersburg Dis¬
trict: Mpome, 1893, Bartels, 19 (ZMUH [ex
ZMHB]). Potgietersrus District: Potgieters¬
rus [24°11'S 29°01'E], iv.1934, R.F. Fawr-
ence, 1 specimen [lost, not examined]
(SAMC B8224).
Erroneous Record: SOUTH AFRICA:
KwaZulu-Natal Province: Estcourt Dis¬
trict: Estcourt, 5 km NE towards Weenen
[29°00'S 29°53'E], 24.viii.1980, A. Haring-
ton, in rock crack on crest of very high hill,
Id, rehydrated (AMNH [AH 3439]).
Hadogenes polytrichobothrius , new species
Hadogenes bicolor Purcell, 1899: Hewitt, 1918:
160, 161 (part), pi. 30, figs. 88, 89.
Hadogenes longimanus Prendini, 2001: 158, 159
(part), 161, fig. 1 (part), tab. 2 (part).
Type Material: SOUTH AFRICA:
Mpumalanga Province: Lydenburg Dis¬
trict: Holotype d (AMNH), Steelpoort, 9 km
SE on road to Lydenburg [Farm Olifantspo-
ortjie 319, 24°45'S 30°18'E], i.2003, I. Eng¬
elbrecht and B. Watkins, in rock cracks.
Paratypes: same data as holotype, Id, 69
(AMNH), 1 juv. (AMCC 138999); Steelpoort
[24°43'S 30°12'E], J. Visser, 2d (SAMC
C4275 [JV 1844], C4276 [JV 1850]), 19
(SAMC C4281 [JV 1846]), 1 juv. d (SAMC
C3901 [JV 1836]); same data, except
“i.2003, I. Engelbrecht and B. Watkins, in
rock crack”, 19 (AMNH); Farm Steelpoort-
park 362, turn-off to Lydenburg, koppies on
left side of road 555, 24°52.510'S
30°02.812'E, 5.ii.2005, L. Prendini and
K.M.A. Prendini, 840 m, in crack of granite
rock in very dry, mixed bushveld, 1 juv. d
(AMNH); Farm Olifantspoortjie 319, en¬
trance gate, 24°46.435'S 30°15.951'E,
5.11.2005, L. Prendini and K.M.A. Prendini,
969 m, in crevices of granite outcrops in very
dry, mixed bushveld, 1 subad. d 1 juv. d
(AMNH), 1 juv. d (AMCC 145221). Belfast
District: Doornkop [Doringkop, on Farm
Doomkop 356, 25°30'S 29°55'E], near Bel¬
fast, R. Gerhardt, Id [damaged] (AMGS).
Middelburg District: Mapoch’s Grotte on
Farm 500, at bottom of hill near entrance
gate, 25°10.675'S 29°57.139'E, 5.ii.2005, L.
Prendini and K.M.A. Prendini, 1407 m, in
crevices of granite rocks at base of koppie,
mixed bushveld, drier than upslope, 3 9
(AMNH). Limpopo Province: Sekhukhune-
land District: Potlake Nature Reserve (Farm
Jagdlust 418), 24°15.160'S 29°54.649'E,
6.11.2005, L. Prendini and K.M.A. Prendini,
844 m, in crevices in sandstone at base of
southern slope of mountains, mixed bush¬
veld, 1 subad. d (AMNH); same data, except
“Potlake Nature Reserve (Farm Wintersveld
417), 24°15.845'S 29°57.021'E, 791 m”, Id
29 2 subad. d 1 subad. 9 (AMNH), 1 sub¬
ad. 9,1 juv. d, 10 juv., disarticulated chela
(AMCC 145222).
Diagnosis: Hadogenes polytrichobothrius
is placed in the bicolor group on account of
the shape of metasomal segment I, which is
wider than it is high posteriorly. It appears
to be the sister species of H. longimanus:
both are characterized by the presence of five
or more trichobothria on the internal surface
of the pedipalp chela, a pronounced lobe, dis¬
tal to the notch in the fixed finger of the ped¬
ipalp chela of adult males and females, and
2006
PRENDINI: SOUTH AFRICAN FLAT ROCK SCORPIONS
11
a relatively short metasoma in the adult male
(ca. 53-57% of the total length), compared
with most other Hadogenes species (includ¬
ing H. newlandsi and H. soutpansbergensis ).
The latter two characters are shared with H.
bicolor, suggesting that the three species col¬
lectively form a monophyletic group, to the
exclusion of H. newlandsi and H. soutpans¬
bergensis.
Hadogenes polytrichobothrius is distin¬
guished from H. bicolor, H. newlandsi, and
H. soutpansbergensis by the presence of five
or more internal trichobothria on the pedi-
palp chela; in the other species, there are
only two internal trichobothria. It is distin¬
guished from H. longimanus by its shorter,
broader pedipalps (especially the chelae),
displaced lobe on the movable finger of the
pedipalp chela of the adult male, and higher
trichobothrial counts. This species has the
greatest number of trichobothria thus far re¬
corded in Hadogenes (Newlands, 1980; L.
Prendini, unpublished data): up to 237 tri¬
chobothria per pedipalp.
Etymology: The species name refers to
the high trichobothrial counts of this species,
which are the highest recorded in the genus.
Description: The following description,
based on the holotype male (AMNH; figs. 2,
3) and a paratype female from 9 km SE of
Steelpoort (AMNH; figs. 4, 5), complements
previous descriptions (Hewitt, 1918; Prendi¬
ni, 2001a) of the first specimen collected,
from Doornkop, near Belfast (AMGS), and
four specimens from Steelpoort (SAMC
C3901, C4275, C4276, C4281). Measure¬
ments and counts are recorded from 4 6 (ta¬
ble 4) and 13 9 (table 3).
Color. Chelicerae, pedipalp chelae, legs,
and distal third of tergites slightly paler, and
contrasting with carapace, rest of pedipalps,
tergites, and metasoma. Telson (d) distinctly
paler than metasomal segments I-V. Stemites
distinctly paler than tergites and metasoma.
Chelicerae, pedipalp chelae, legs, and telson
(9), Clay Color 26; carapace, pedipalp pa¬
tellae and femora, Raw Sienna 136; tergites
I-VI, proximal two-thirds, stemite VII, me¬
tasoma, Brownish Olive 29; tergites I-VI,
distal third, pectines, genital operculum, and
sternites, Raw Umber 123; telson (d), Buff
124.
Carapace : Three pairs of lateral ocelli,
slightly smaller than median ocelli (figs. 6,
7). Median ocular tubercle with superciliary
carinae well developed, protruding above
ocelli, and interocular sulcus distinct. Ante¬
rior margin of carapace with median notch
well developed, triangular inset situated far
back, with frontal lobes protruding anterior¬
ly. Anteromedian sulcus deep, suturiform,
furcating anteriorly around triangular inset.
Median longitudinal suture distinct, contin¬
uous from anterior furcated sutures, through
ocular tubercle to posterior furcated sutures,
which converge on ocular tubercle from pos¬
terior carapace margin. Posterior furcated su¬
tures obsolete, discontinuous. Posteromedian
and posteromarginal sulci distinct, shallow.
Paired median lateral and posterolateral sulci
also distinct, shallow. Carapace entirely
granular, except for surfaces of frontal lobes,
median lateral, posterolateral and postero¬
marginal sulci, which are smooth. Granula¬
tion almost uniformly fine, becoming coarser
on anterocular and anterolateral surfaces.
Chelicerae : Movable finger with distal in¬
ternal tooth slightly smaller than distal ex¬
ternal tooth, and opposed. Ventral aspect of
fingers and manus with long, dense macro-
setae.
Pedipalps : Femur with four distinct cari¬
nae (fig. 12); ventroexternal carina obsolete,
reduced to a few granules proximally; dor-
soexternal carina, dorsointernal and ventroin-
ternal carinae costate granular, composed of
large, heavily sclerotized granules; externo-
median carina comprised of spiniform gran¬
ules; dorsal, dorsoextemal and ventral inter-
carinal surfaces finely and uniformly granu¬
lar; internal intercarinal surfaces smooth, ex¬
cept for a few scattered spiniform granules.
Femur width 39.5% (33-46%) of length (ta¬
bles 3, 4).
Patella with six distinct carinae (figs. 13-
15); dorsoextemal carina obsolete; dorsoin¬
ternal and ventrointemal carinae costate to
costate granular; internal carinae costate
granular, composed of very large, heavily
sclerotized spiniform granules, converging at
apex of anterior process; externomedian and
ventroexternal carinae granular; dorsoexter-
nal and ventral intercarinal surfaces finely
and uniformly granular, becoming granulo-
reticulate on ventral surfaces; internal inter¬
carinal surfaces smooth, except for a few
12
AMERICAN MUSEUM NOVITATES
NO. 3502
Figs. 2-5. Hadogenes polytrichobothrius n.sp., habitus of holotype 8 and paratype 9 (AMNH). 2.
Dorsal aspect, 8. 3. Ventral aspect, 8. 4. Dorsal aspect, 9. 5. Ventral aspect, 9. Scale bars =10 mm.
scattered granules; anterior process strongly
developed. Patella width 61.5% (56-67%) of
length.
Chela with three distinct carinae; dorsal
secondary and digital carinae obsolete (figs.
16, 17); external secondary carina strongly
developed, costate granular; ventroexternal
carina strongly developed, crenulate, aligned
parallel to longitudinal axis of chela, with
distal edge disconnected from external mov¬
able finger condyle and directed toward a
point between external and internal movable
2006
PRENDINI: SOUTH AFRICAN FLAT ROCK SCORPIONS
13
finger condyles, but closer to external con¬
dyle (fig. 19); ventromedian carina obsolete,
reduced to a vestigial granule proximally;
ventrointernal carina also obsolete; interno¬
median and dorsointernal carinae weakly de¬
veloped, each comprised of a series of iso¬
lated spiniform granules; dorsomedian carina
strongly developed, composed of a continu¬
ous double row of spiniform granules; dorsal
intercarinal surfaces smooth to weakly retic¬
ulate, becoming finely granular externally
(d), smooth to uniformly finely granular (9);
external intercarinal surfaces coarsely gran¬
ular (fig. 18); dorsointernal intercarinal sur¬
faces with scattered spiniform granules, be¬
coming finely granular on internal surface of
fixed finger; ventrointernal intercarinal sur¬
faces smooth. Chela with pronounced, coni¬
cal lobe on movable finger, fitting unevenly
into corresponding notch in fixed finger of
adult 8, i.e. displaced to external surface and
overlapping, when fingers closed; fixed fin¬
ger additionally with pronounced, conical
lobe distal to notch, and smaller, rounded
lobe proximally. Dentate margins of chela
fingers with double row of denticles, fused
proximally at lobe/notch and also distally.
Chela height 47% (38-56%) of width; chela
width 66.5% (60-73%) of length along ven-
troexternal carina; length movable finger
104% (97-111%) of length along ventroex-
ternal carina.
Trichobothria : Neobothriotaxic major,
type C (figs. 12-19; tables 3, 4), with the
following segment totals: femur, 3 (1 d; 1 i;
1 e), patella, 82-121 (2 d; 1 i; 28-43 v; 51-
75 e) and chela, 75-113 (66-101 manus; 13-
16 fixed finger, including 5-8 /). Total num¬
ber of trichobothria per pedipalp, 160-237.
Only femoral trichobothria, trichobothria in
the d and i series of the patella, and tricho¬
bothria in the D, d, and e series of the chela
are stable in number and distribution. Exter¬
nal and ventral trichobothria of the chela and
patella are numerically and distributionally
too variable for diagnostic purposes. This
species is characterized by the presence of 5-
8 accessory trichobothria in the i series of
the chela.
Me so soma : Tergites each with paired sub¬
median depressions and obsolete median ca¬
rina. Pre-tergites of 8 and 9 smooth and
shiny. Post-tergites I-VI smooth and shiny in
8 , except for very fine and even granulation
on anteromedial surfaces (excluding median
carina and submedian depressions, which are
smooth) and lateral surfaces; VII sparsely
and coarsely granular anteriorly, becoming
smooth posteriorly in 8; post-tergites of 9
smooth and shiny. Sternites smooth and
shiny, each with paired longitudinal depres¬
sions internal to spiracles; VII with pair of
shallow posterolateral oval depressions in 8 ,
and pair of obsolete carinae, converging dis¬
tally towards shallow notch in distal apex
(figs. 10, 11). Sternite VII length 92.5% (85-
100%) of width in <?, 76.5% (66-87%) in 9
(tables 3, 4).
Pectines: Mesial margin of first proximal
median lamella of each pecten angular, with
pectinal teeth present along entire posterior
margin in 8 (fig. 8); mesial margin of first
proximal median lamella shallowly curved,
with proximal fifth of posterior margin de¬
void of teeth in 9 (fig. 9). Pectinal teeth (left/
right): 21-24/20-23 (8), 16-21/16-20 (9).
Sternum : Subpentagonal, type 2. Median
longitudinal furrow deep and narrow along
entire length (figs. 8, 9).
Genital operculum : Suboval, completely
divided longitudinally, with genital papillae
present (<?). Subcordate, partially connected
by a membrane in anterior two-thirds, with
distinct distal lobes in posterior third, and
with genital papillae absent (9).
Legs : Femora each with paired granular
carinae on ventral surface, becoming less de¬
veloped on posterior legs. Basitarsi each with
few spiniform setae on prolateral and retro-
lateral margins, decreasing in number from
anterior to posterior legs. Telotarsi each with
two rows of three ventrosubmedian spini¬
form setae and basal row of ventromedian
spinules. Telotarsal laterodistal lobes truncat¬
ed; median dorsal lobes extending to ungues.
Telotarsal ungues short, distinctly curved,
and equal in length. Retrolateral pedal spurs
absent.
Metasoma and telson: Metasomal segment
I posterior height 88.5% (78-99%) of width
(tables 3, 4). Metasomal segments I-V pro¬
gressively increasing in length, and decreas¬
ing in width, segment V width 68.5% (57—
78%) of segment I width. Metasoma slender,
width percentage of length for segment I,
30% (23-37%) in 8, 42% (32-52%) in 9;
14
AMERICAN MUSEUM NOVITATES
NO. 3502
d
m
rx
&
. o
a®
«5 (N
fl ^
« s
Jl
1 s
^ Ph
2 -d
"5* fl
•2 3
tg
a-2
SU ._
m s fl
3 II
m *§ |
<1 d >
H Ph
- fl
4 §
c-8
s 2 J
fl £
fl z
Q £
.a ©
'M S
.83 ,2
Ll ^
S 2
2 fl
o>
s
2
3
a
0)
§
PC
z
X
z
X
z
PC
z
PC
z
PC
z
PC
z
PC
z
PC
z
PC
z
PC
z
PC
Z 2
I a
u ^
- i S S3
S ft
oo U
fl
3 s
.J| o
3 U I
Oh
00
-h r- o
00 On O
oo co in
in o co no
O 1C H 00
o a-’
O O 00
00 q Tt
On d d
— ex ue
o fl; fl;
odd
fl- H H (S
NO O ce t"
o a-' i> d
ON CO O h
fl- On O NO
o fl-' h d
hH (fl hH
00 eX On
in oo fl;
On in in
ex ro oo
on q o
o h oo
fl- ON ON
O oo m
odd
On NO 00
fl; >n o
On d d
On NO On
NO A; in
o d d t-h d ex
fl- 00 rH
NO in rH
d 00 On
i!
y o
.2 'C ^
Cod
B S g>
c= O g
U
d
d
2
3
e
'x
fl
2
j =
u
q q
d
rH 00 NO fl" 00 (fl (fl
o ex o h (N ex o
odd 3 ^ q 2
On in 00 (fl
rH Ifl (fl in
o fl-' o d
O l~H rH lfl
o h o\ rj
<4 d d oo
no in h fl
(fl: On (fl in
q fl- q in
no m
<fl >n rH
r-. <fl o (fl
On (fl fl; o
o ici o d
rH (fl H
fl fl- h in fl
on ce oo oo o
d q d q d
On oo fl- NO
no o fl fl
q d (fl d
rH (fl —(
O
ON flr
d no
H fl oo
rH fl; in q
(A d d oo
o
On
O
b
£
Q
3 O
2 2 d -c
'§ g> 3
goo
flr
o
00
O
ex
>n
00
ex
On
i>
ue
On
it
NO
fl-
ue
ee
00
ue
ex
(fl
in
00
ex
ee
e-
ee
NO
fl;
00
fl;
NO
q
fl;
q
ue
o
q
O)
ex
00
fl-
d
o
On
ue
ce
d
ON
ue
o
00
q
q
q
q
q
oo
fl-
in
1—1
r-
ee
NO
fl¬
(fl
00
o
00
ee
o
NO
ex
ON
NO
00
ue
o
ee
00
T _
ue
t"-
fl-
d
flf
NO
ex
H
e-
ee
NO
o
o
fl;
fl;
fl;
q
q
00
fl;
fl;
ON
o
d
ON
in
d
o
d
d
fl¬
q
00
00
ex
00
q
q
q
q
00
q
fl-
fl-
ee
ue
fl¬
00
NO
(fl
o
o
00
ex
00
ue
ex
fl-
]
00
o
t>
e-
(fl
d
On
o
o
NO
ee
ue
ON
fl-
re
o
q
q
rH
On
o
fl;
ee
fl-
00
(fl
On
d
in
d
On
d
(fl
q
On
q
q
00
ce
q
q
q
q
On
(fl
fl-
NO
ee
NO
fl-
i- 1
fl-
NO
ex
00
in
00
NO
o
ue
NO
e-
o
00
ue
e-
o
o
ee
(fl
in
On
rH
o
in
ee
On
rH
On
o
ex
q
ON
q
ON
q
°°
ON
d
d
On
in
in
d
d
d
fl-
ue
On
d
q
ON
q
q
q
q
q
ON
m
in
>n
ee
NO
fl-
<fl
NO
re
NO
fl-
fl¬
ue
NO
o
NO
fl-
ee
On
i>
ue
00
00
NO
d
d
O
i>
O)
NO
ee
i>
ue
ue
fl-
q
q
On
q
00
fl-
ue
ce
ee
fl-
(fl
o
fl-
fl-
cl
d
d
re
ue
q
A-
q
00
q
q
q
q
q
On
fl-
fl -
1—1
’ 1
NO
ee
1—1
NO
fl¬
- 1
m
o
00
fl-
e~
in
NO
ee
,
Cr
ex
ex
ee
fl-
ee
ex
ue
o
ee
d
m
fl;
ON
in
(fl
e]
q
fl-
o
ue
ON
00
rH
<s
q
00
ex
o
q
d
o
in
d
d
ue
q
fl¬
d
fl-’
On
q
o
fl"
q
00
q
fl-
o
fl-
(fl
- 1
>n
ee
- 1
r^
fl-
- 1
m
(fl
o
(fl
ee
in
ue
o
ee
0
,
NO
ue
,
,
NO
00
NO
On
00
fl-
fl-
NO
00
NO
we
ee
00
o
o
q
e-
fl;
q
ON
ue
fl;
re
rH
rH
fl-
_
On
fl-
fl"
ee
d
d
fl¬
d
00
q
q
q
q
q
q
q
q
ON
fl-
fl-
NO
ee
1—1
NO
fl-
i
o
On
m
NO
in
NO
00
fl-
ee
NO
NO
o
e~
ue
0
00
0
ue
ee
ex
d
fl-
O
ex
fl;
in
ex
ex
q
00
q
re
ee
o
On
o
l>
q
q
C3
fl-
On
in
>n
ex
ee
q
fl-
ue
ON
00
q
On
q
q
q
q
q
00
fl-
fl-
- 1
NO
ee
- 1
NO
fl¬
- 1
00
,
fl-
00
o
o
fl¬
,
e-
ON
O
ex
A-
00
re
ue
On
ex
ee
(fl
5
in
On
(fl
NO
ee
NO
rH
q
On
q
00
rH
e-
fl;
ee
rH
fl-
ON
d
On
fl¬
in
00
ee
d
fl"
ue
q
d
q
On
q
q
q
q
q
On
(fl
m
>n
ee
NO
fl¬
ON
in
00
ee
o
e-
,
ee
.
00
_
ee
_
fl-
ex
00
On
On
r-
00
(fl
(fl
00
ce
NO
ce
00
ex
O)
rH
q
q
00
q
>—i
o
ue
q
00
o
d
On
in
<n
d
ON
d
fl-
ue
O
fl-
q
00
q
q
q
q
q
00
fl¬
fl¬
in
(N
l>
fl¬
ex
ex
ex
m
ex
fl¬
00
o
fl-
ex
ue
NO
fl¬
o
00
On
ue
NO
00
fl;
NO
ee
re
q
o
00
fl;
ue
q
°°
00
q
NO1
q
00
On
d
fl-
d
d
q
fl¬
d
o
00
q
o
q
q
d
q
q
oo
(fl
cfl
in
ee
(H-
fl-
in
On
fl¬
NO
NO
fl¬
On
o
ue
NO
ue
NO
On
ON
fl-
e-
ex
ee
o
(fl
fl
ee
fl;
fl;
ue
ex
o
q
ON
o
NO
On
o
fl;
00
ex
q
q
q
Cfl
d
o
<n
in
ex
q
q
fl-
ue
q
q
q
On
q
q
q
q
q
00
fl-
(fl
ue
ee
Cr
A-
fl-
On
NO
On
fl-
o
ee
On
o
NO
NO
ue
NO
o
00
fl-
NO
ue
d
d
(fl
iq
NO
o
ee
o
<—i
o
Cr
rH
00
o
ON
q
ue
00
o
00
_
NO
o
in
q
ON
ee
00
ue
00
o
ue
q
q
q
q
00
q
fl-
o
fl-
fl"
rH
rH
NO
ee
rH
00
ue
rH
rH
rH
d d d
,+o i+h i+h
(t>
oo
3
d
o ® o o
3 w
>
o
2 _
, o
el
00 -2
3 d
^ C*3
d d
00 OX)
■§ 3
q q
0) o
3 o
•fl
d
o
o
d
o
q
■s d
o o
3 3
2 2
d
o
M M S -g
X X OX) -fl
3 3 3d
2 22 o
d §
1 B
2 2
3 3
3 3 ^
X X 0X)
3 3 3
2 22
3 Bn 3 q
d 0 d
2
o
q
3
q
•B d
3 3 W)
§ j5
3 ET
2 >
o o
3 d
o S
to C
<u ox
2
3
2 d
2
3
2 d
X X OX)
3 3 3
2 22
2
o
m
B
<D
X X OX)
3 3 3
2 22
2
o
5
(D
Hadogenes polytrichobothrius n.sp.
2006
PRENDINI: SOUTH AFRICAN FLAT ROCK SCORPIONS
W 1
_i -S
m 'S
< r §
H O
c,
% %
.. ’§ «i
l—H P O |
«5a U I
Cu
DO
PC
d
NO
o
d
c-
c-
o
00
ON
d
cn
NO
NO
NO
d
m
d
_ z
a
On
3f
On
d
co
o
00
o
o
00
q
o
q
q
q
d
3f
q
cd
. —i
d
00
, —i
ci
On
, —I
d
On
^ —i
d
ci
00
in
d
in
NO
CL,
>n
o
<
X
cn
3f
3f
NO
00
m
d
3t
3f
3t
3f
d
d
00
ON
z
cd
d
o
Cl
ON
in
00
m
d
cn
q
m
d
o
d
l-H
cd
ci
cn
00
q
ci
00
d
ci
On
ci
ci
q
00
o
00
ON
d
ON
Ph
>n
o
<c
3t
3f
On
00
NO
3t
On
O
NO
d
cn
in
cn
NO
d
On
z
cd
in
Cj
cn
ON
NO
ON
On
q
CO
d
3f
q
d
cn
00
"I
cd
o4
cn
On
ci
ci
ON
ci
ci
cd
ci
00
NO
3f
On
d
d
Ph
m
d
<
1—1
t>
in
d
o
NO
d
00
00
cn
NO
NO
NO
NO
00
in
3t
o
z
cd
cn
in
On
Cl
o
d
o
d
d
3t
q
°°
d
o
CO
'—i
o
d
°*i
cd
d
cd
On
ci
cd
o
d
ci
q
ci
ci
q
00
00
3T
On
00
ON
P<
m
<
1-1
On
00
NO
d
NO
cn
m
in
d
d
00
s
NO
NO
_(
ON
z
cd
3|-
'—1
3f
CO
i>
NO
cc
q
o
3"_
q
3t
NO
°9
d
CO
cd
d
cd
On
ci
ci
On
d
d
ci
ci
d
3f
00
On
00
ON
a
m
o
<
ac
in
>n
d
On
in
,_i
d
NO
o
NO
cn
NO
d
0
3t
ON
On
NO
z
para
d
On
d
NO
3t
00
d
3t
cn
o
3f
CO
o
31"
00
q
d
cd
o
ci
cd
o
d
cd
ci
cd
cd
ci
00
8
o
d
On
d
NO
<
PC
o
3t
d
d
d
00
d
3f
o
in
in
d
d
z
a
3t
CJ
3j-
ON
NO
00
o
NO
N-1
d
3f
q
d
00
3f
On
rH
<*1
3
ci
cd
On
ci
On
d
ci
ci
ci
00
in
d
00
NO
00
m
o
<
pc
cn
On
NO
in
m
3f
0
3t
cn
d
3f
cn
>n
On
>n
3t
00
z
a
3t
3t
On
d
o
o
cn
o
d
NO
00
00
o
d
O
On
d.
1—1
<*1
3
d
cd
00
ci
cd
On
d
cd
—
d
d
q
00
ci
00
d
NO
ft
m
<
PC
d
m
o
l>
d
NO
o
cn
3t
d
d
cn
00
NO
d
o
On
d
z
a
d
d
o
d
d
o
NO
d
m
ON
On
ON
cn
cn
q
— i
"I
3
ci
cd
ON
ci
ci
On
d
ci
O
ci
ci
—
d
cd
o
On
d
00
Oh
>n
<
1-1
PC
NO
00
ci
m
NO
3t
m
3f
NO
m
On
d
O
d
z
3
CO
3f
3f
in
00
cc
3f
d
in
q
<—<
00
q
d
i—i
o
00
'—1
"i
3
d
cd
On
ci
d
On
d
d
o
d
cd
q
00
cd
00
On
NO
d
a
un
o
<
PC
00
d
NO
On
On
NO
m
3t
o
d
00
o
ON
NO
cn
d
z
3
NO
d
00
r— 1
o
3t
NO
q
3t
00
3t
3l"
cn
00
t-H
3
(N
cd
00
ci
cd
o
ci
cd
ON
ci
cd
d
d
ci
o
On
NO
d
a
m
<
1-1
d
d
o
in
m
On
On
d
cn
cn
NO
d
NO
cn
in
d
z
3
3f
NO
On
cl
'■H
d
3T
On
ON
00
d
3f
On
o
cn
00
On
3
<N
cd
00
ci
cd
On
ci
d
ON
ci
cd
ci
d
d
o
On
d
NO
a
un
<
1-1
u ^
00
NO
d
On
3t
00
cn
00
cn
NO
00
d
cn
31"
d
O
3f
00
^ 00
Of ^ <N
<r T d _
r \
3
00
1— 1
>—i
q
in
o
q
cn
q
>—i
3t
ON
cn
q
00
in
00
i-H
3
Ci
3f
rd
ci
cd
d
ci
cd
d
cd
cd
d
o
in
ON
On
d
00
a
T—i
*—i
NO
cn
y—l
33
33
q a
£
33
op
'B
bp
-S .SP
33
bp
'S
43
$
'S
33
•3 0)
> 33
$
'S
33
a
3
B 33
B
3
B 33
B
3
B 33
X X W)
3 3 3
a a «
X X bD
3 3 3
3 £ £
S “ M t)D
3 3
.V N\J .'-J »4 }_i
EES EES
1 .1 1
X X ^
ctf cd P
a a £
3
o
3 3
E I
4- t/i
^ a
3 "5
o ®
o +
a
B
3
33
bo
•3
33 ^
>
3
a
o
co
a
3
a
o
GO
a
<D
35
M w>
a «
§ ia
C/3
cd rd
£ £
15
Measured from base of condyle to tip of fixed finger.
Sum of metasomal segments I-V and telson.
16
AMERICAN MUSEUM NOVITATES
NO. 3502
Figs. 6-11. Hadogenes polytrichobothrius n.sp., carapace, pectines and stemites of 6 and 9 para-
types (AMNH). 6. Carapace, 6. 7. Carapace, 9. 8. Pectines and basal piece, 6. 9. Pectines and basal
piece, 9. 10. Sternite VII, S. 11. Stemite VII, $. Scale bars = 5 mm.
2006
PRENDINI: SOUTH AFRICAN FLAT ROCK SCORPIONS
17
Figs. 12-15. Hadogenes polytrichobothrius
n.sp., carinae, trichobothria and macrosetae on
dextral pedipalpal segments of paratype 9
(AMNH). 12. Femur, dorsal aspect. 13. Patella,
dorsal aspect. 14. Patella, external aspect. 15. Pa¬
tella, ventral aspect. Scale bar = 5 mm.
for II, 19.5% (17-22%) in S, 27.5% (23—
32%) in 9; for III, 18% (15-21%) in S, 26%
(22-30%) in 9; for IV, 16.5% (14-19%) in
6, 23.5% (20-27%) in 9; and for V, 15.5%
(12-19%) in S, 21% (17-25%) in 9. Telson
vesicle width 112.5% (103-122%) of meta-
somal segment V width; globose in S , oval
in 9, with flattened dorsal surface and round¬
ed ventral surface (fig. 33), height 33% (25-
41%) of length. Aculeus short, 25% (19-
31%) of vesicle length, and sharply curved.
Total length of metasoma 115.5% (111—
120%) of combined length of prosoma and
mesosoma in S, but 85% (78-92%) in 9.
Eight carinae on segment I, six carinae on
segments II-IV, and five carinae on segment
V (figs. 20, 21). Dorsosubmedian carinae of
segment I becoming obsolete distally, but
distinct throughout length of segments II-V.
Median lateral carinae fully developed on
segment I, but absent from segments II-V.
Segments I-IV with closely paired ventro-
submedian carinae, fused into a single ven-
tromedian carina on segment V. Median lat¬
eral and dorsosubmedian carinae costate on
segment I, dorsosubmedian carinae costate to
costate granular on segments II-V (9), com¬
posed of spiniform granules on segments II-
V (c3). Dorsosubmedian carinae of metaso-
mal segments II and III each terminating dis¬
tally with enlarged, spiniform granule; dor¬
sosubmedian carinae of other metasomal seg¬
ments without spiniform granules distally.
Ventrosubmedian and ventrolateral carinae
costate on segment I, costate to costate gran¬
ular on segments II-IV. Ventrolateral and
ventromedian carinae of segment V com¬
posed of spiniform granules. Intercarinal sur¬
faces smooth, except for lateral surfaces of
segments III-V in 6 . Telson smooth, sparse¬
ly covered in long macrosetae.
Hemispermatophore : Doubled hook near
base of distal lamella; distal crest truncate
(figs. 22, 23).
Geographic variation : Specimens collect¬
ed at the northernmost locality record, Pot-
lake Nature Reserve, display longer, narrow¬
er pedipalps and a more slender metasoma
than specimens collected further south in the
distributional range (tables 3, 4).
Ontogenetic variation : The presence of a
lobe on the movable finger of the pedipalp
chela and a corresponding notch in the fixed
finger (figs. 16, 17) are indicative of sexual
maturity in most species of Hadogenes
(Lawrence, 1966; Newlands and Prendini,
1997; Prendini, 2001a). The lobe and corre¬
sponding notch are absent from the fingers
of the pedipalp chela in subadults and juve¬
niles, developing in the final instar of spe¬
cies, such as H. polytrichobothrius, in which
these characters are present in adults.
In the specimens of Hadogenes examined
for this study, sexual maturity was assessed
by the presence of a lobe and notch in males
and females, and by the presence of fully de¬
veloped paraxial organs in males, and ovari-
18
AMERICAN MUSEUM NOVITATES
NO. 3502
<3
Oh
CB
fl ^
•2 £
S? IT)
§ o
ft
-ft
05 >4
ft §
I§
Is
“3
a: S
-d £
S3 ^
3 -d
. fl
a 3
5/3 y—V
C £
sS
•C ~
^‘S
ft *3
-ft *d
ft fl
-ft ft
.ft Si
C &H
fl
ft, «
05 05
ft “
§ §
II
85 *
3 i
*a
2
fl
ft
s
Si
£
3
ft
is
ft
ft
«2
3
AS
Q
ft
X
q
CN
ON
(N
tn
3"
o
00
t>
(N
e'¬
tn
tn
o
CN
<N
On
o
q
ON
C'
o
o
cn
z
2
00
t-
NO
00
q
<n
On
r-
(N
tn
tn
en
q
NO
cn
i
q
O
(N
q
q
o
On
q
q
q
<
3
o
NO
NO
On
3"
X
NO
NO
?5
cl
00
in
ON
d
NO
tn
NO
On
X
X
o
tn
CN
00
CN
cn
X
Oft
&
tn
tn
tn
in
tn
NO
E
t>
o
t-
O
^t
O
00
tn
o
(N
NO
NO
tn
tn
00
(N
NO
00
(N
00
cn
tn
00
t"
cn
z
3
On
no
cn
On
q
q
rh
O)
nj
l>
q
q
q
tn
q
bO
m
NO
q
00
cn
o
q
q
H
q
q
3
O
NO
t>
On
(N
i>
NO
cl
tn
o
00
in
in
tn
NO
o
00
X
On
cn
CN
00
CN
cn
X
a
tn
^j-
fH
NO
tn
t>
cf
d
c«
<
d
X
00
00
m
in
00
tn
00
in
00
tn
On
O
tN
NO
ON
o
00
CN
m
(N
tn
NO
cn
tn
•2
z
2
3;
t-
On
t-
<n
°°
tn
q
<N
tn
tn
q
O)
q
NO
cn
O
q
q
o
q
o
00
o
ON
o
t>
q
xe
’5
<
3
_'
c-
c-
o
in
tn
00
X
cl
cl
d
00
3’
in
Pf
NO
d
X
tn
00
cn
o
tn
CN
On
CN
tn
CN
§
£?
ft
a
tn
■^t
tn
NO
cn
r-
cf
-ft
<<
X
cn
<N
tn
(N
tn
NO
3"
tn
00
tn
00
NO
00
CN
o
o
m
cn
O
00
O
On
00
s
z
3
q
cn
q
NO
in
q
On
CJ
tn
n
q
°°
q
in
tn
q
q
00
o
q
q
C'
q
O
NO
q
NO
0 >|
3
_3
00
00
o
in
in
00
u
cl
00
cl
00
NO
00
o
NO
tn
NO
NO
CN
On
cn
cn
On
CN
cn
o
S-
a
fH
<
H
tn
fH
<n
tn
H
in
tn
c-
cf
-H
s
ft
to
<!
*5
X
CN
tn
NO
On
NO
00
t>
(N
(N
c-
m
in
o
o
NO
00
o
00
NO
-tt
On
00
e-
00
ON
On
ft
£?
z
2
O
no
00
NO
tn
m
00
o
(N
tn
tn
q
q
m
in
cn
cn
00
H
q
q
q
q
H
00
NO
00
H
ft
3
CN
00
00
O
in
NO
00
00
cl
ON
fH
00
3 :
NO
X
d
NO
tn
00
X
00
tn
X
cn
cn
00
CN
tn
CN
00
so
tn
tn
tn
in
<N
e'¬
■cf
"ft
ft
<
&3
X
I-
o
CN
-tt
in
o
CN
CN
00
>n
c-
<N
(N
NO
On
en
CN
NO
CN
o
ON
00
cn
z
3
On
1 —
ON
o
q
O)
nj
tn
't
o
o
NO
cn
q
q
q
cn
q
o
c-
00
q
o
q
NO
*°i
<
3
o
q
in
00
tn
00
in
cl
P=
NO
cl
X
in
cn
in
CN
cn
o
o
CN
CN
X
CN
cn
in
a
CN
<n
tn
NO
cn
NO
X
o
O
m
■St
r-
ON
NO
tn
(N
(N
00
tn
tn
NO
<N
On
m
c-
o
NO
NO
00
On
n -
00
o
00
-1
JO
in
cn
On
n-
°°
tn
q
(N
tn
00
q
q
in
tn
q
00
q
cf
00
00
NO
q
q
q
00
'o
00
t>
On
n 3
NO
ON
00
cl
o
cl
00
cf"
X
00
cl
NO
cn
00
On
CN
CN
tn
tn
NO
CN
tn
43
tn
*—1
tn
>n
cn
<—1
00
cf
CN
*o
U >n
q r- g
d ^ p.
oo U
ffi
Z
*o
E
z
JJ..
.. w fl _
o Zh C r>.
u O 3 £
02 U Z H
Oh
00
On
s
tn
in
e'
r-
in
NO
>n
NO
ft-
(N
o
NO
00
<N
s
cn
X
00
o
'—i
q
o
m
NO
q
q
q
q
e'¬
cn
NO
o
00
00
On
^6
NO
o
in
CN
NO
NO
d
ON
00
in
•ri
en
d
NO
vn
00
NO
**
cn
X
tn
r-
cn
i— 1
NO
X
O cn cn On
cn in q
l> t- q
cn cn <N
m
00
,_1
CN
o
in
tn
X
On
ON
O
o
X
CN
CN
O
tn
cn
On
t—
tN
00
__i
CN
tn
q
CN
q
00
NO
q
NO
q
in
ON
q
X
q
q
o
q
fH
o
in
o
o
o
q
q
q
NO
C-
in
1 —
NO
00
r—
d
00
o
in
On
o
cn
r-
q
X
in
q
CN
o
cn
cn
tn
CN
cn
q’
cn
fH
--1
X
--1
t—
(N
00
in
--1
--1
© ON q
o vo no
CN vo 00
O On cn
o vo t>
<N m i— m
NO VD to O;
c-4 UO CO 00
h to m 'O
no >n h oo in oo
q
00
m
00
O
q
in
q
tn
tn
t—
00
in
On
O
i—
NO
i
ON
cn
tn
' —1
00
q
NO
cn
1 -H
q
q
q
q
CN
CN
00
CN
cn
On
o
ON
CN
o
in
in
d
d
X
q
X
q
in
X
X
X
X
o
X
q
tn
q
ii
NO
cn
’ 1
i—
q
' 1
o
r-
On
cn
o
in
ON
On
t-
CN
NO
00
t-
q
q
q
cn
cn oo h o in to
o\ in no no >n
h in to
.3
o
1 ll
O tfl £
X
S S
Ph Oh
<D q <D <D
X X
bn bn
II
h in
£
X
o r-
<n o
3 ;
to to o
o
q
o
3
-g
X
<D
o
X
QJ
X
X
o
q
B -fl
O
X
X
X
X
d
T3
%
d
TO
bp
cS
>
cd
.2
.2
X
_bp
cd
c3
B Sp
B
X
_bp
_bp
T5
'$
%
S
d
X
s
fH
<u
>
t+H
o
E
<+H
*c
O
|
o
X
X
o
X
%
S
‘S
X
S
‘C
o
n
‘d
B
o
n
■>
5 X
S £
bO
B
bn
%
S
'3
X
S
%
S
'3
X
S
u
o
3
3
o
o
o
o
3
3
o
Q
3 3
3
3
3
3
3
3
O
‘d
*C
(D
E
E
X
'B
o
-c
o
X
o
X
o
E
E
O
X
o
E E
B
JD
O
X
■5
E
g
B
E
g
■S
OJ
W)
X
X
W)
bO
on
‘C
•fH
X
X
bi
X X
bn
3
bn
X
'x
bn
X
'x
bn
r —1
C/J
Xn
a
a
g
a
HH
HH
C3
3
is
3
3 3
3
X
3
3
3
3
3
3
3
C-
cd
ftX
a
E
s
o
o
X
X
E
s
j3
0-
E £
CD
o
o
£
CD
E
E
CD
E
E
CD
U
43
u
a
3
a
x
0) *3
Oh rZ 00
2006
PRENDINI: SOUTH AFRICAN FLAT ROCK SCORPIONS
W 1
q -S
m 'S
< r §
H O
X
vo
CN
3d
in
3d
CO
m
in
VO
O
CO
00
ON
o
O
(N
VO
Z
2
CO
in
CN
a \
Ov
CO
00
r--
O
rH
q
q
VO
q
00
q
c3
a
cn
CO
z
CN
CN
CN
CN
CN
CO
00
s
CO
CN
00
<6
sc
00
CN
VO
CO
(N
e-
vo
<N
O
00
CO
ON
CO
vo
z
3
o
in
co
1—1
in
in
q
00
q
q
q
q
q
o
q
i
ch|
c3
cn
CO
q
CN
CN
q
CN
CN
q
CN
CN
q
00
CO
q’
On
00
a
t-H
VO
CN
d
<
d
PC
3d
3d
d^
3d
o
o
r-H
vo
CO
00
On
00
'tt
00
d'
•S
z
CO
o
CO
CO
o
q
CO
m
q
in
1—1
00
o
q
q
q
i-i
*£5
CN
3d
CN
CN
CO
oi
<N
CN
CN
<N
CO
q
On
On
00
00
U
VO
si
a
vo
CN
^5
3
"3l
65
PC
O
vO
CN
CN
VO
00
00
o
00
CO
vo
O
CO
oC
d'
a
z
3
CO
On
CO
o
o
q
q
o
q
q
q
CN
o
in
o
>—i
§
3
CN
CO
q
CN
CO
CO
q
CO
CO
CN
CO
q
ON
iri
On
ON
NO
S'
a
vO
(N
s
<
t—i
0
&3
to
CO
CN
d~
vo
00
00
vo
o
vo
o
vo
ON
VO
NO
O
l>
^5
~
z
2
NO
on
On
in
q
co
q
q
q
NO
q
ON
■—i
q
H
Nj
CN
CO
o
CN
CO
CO
q
CO
CO
CN
CO
o
ON
00
ON
On
00
d'
bo
a
VO
CN
§
<
SC
ON
CN
vo
VO
On
CN
^t
00
o
l>
00
CN
CO
CN
CN
in
o
z
3
00
3d
q
q
q
00
oo
vq
m
q
i-H
q
q
CO
d'
o|-
<N
CN
^1
3
CO
vo
q
CN
00
q
CN
VO
CN
CO
q
00
t>
o
o
o
o
a
'■H
00
"Cf
CN
' t O
in
ON
NO
00
in
CO
NO
o
in
CO
On
On
in
00
CO
,—i
in
00
o
o
q
CN
00
x —1
NO
q
q
CN
o
o
CO
CO
CN
’o
CN
CO
in
CN
CO
d^
CN
CN
■sd
CN
CO
q
o
00
On
CN
r-H
o
<N
CN
CN
CN
00
x —I
CN
U <n
*o
*0
*o
d- g
9 &
M Q,
O a
o Jo ..
.. « d «i
g 0 3 S
$ u £ H
a
cn
00 (N ^
3d q q
<N co cN
3d q
CN CO
CO CN CO CO
O O t**
q o q
CN CO NO
O O 00
o q co
CN CN 05
VO O ON
q o q
CN 3d co
CN CO CO CN CO CO
oo t" o\
h 00 \t
cn cn 3d’
00 3 d q
(N CO 05
d~ oo cm i-h
Tf o h >
<N co cn oo
<N CN CN o in
d co o oo
d co d M 05 3f
ON CO O
o no d
cS A
q
q
cS ^
q
q
q .SP
q
op
q .sp
q
op
•3 05
> q
'$
- S
q
■3 05
^ q
- S
q
bi)
c
0(5
q
£
£
£
£
£
£
£
£
-2
oo
3
3
3
3
3
3
3
3
c/5
3
3
£
£
£
£
£
£
£
£
£
£
3
05
<U
in
X
oi
X
X
00
X
X
oi
X
X
'3
3
3
3
3
3
3
3
3
3
3
3
£
£
05
£
£
05
£
£
05
£
£
O
o
£
o
l/l
o
@
05 03
£
o
c/1
B
05
£
o
c/1
B
05
£
o
4- so
^ 5
S "5
£ g
o
03 _L_
O +
w
a
y OB
O -H
-H (M
3
u
2 H
5 q
a >
05 V4H
t5 o
W) U
I §
3 O
bO *3
a O
05 O
B
£
05
too
a
43
ft t3
d £3
33
S3
O
£
00
«S 3
^ £
£
3
00
19
20
AMERICAN MUSEUM NOVITATES
NO. 3502
Figs. 16-19. Hadogenes polytrichobothrius n.sp., carinae, trichobothria and macrosetae on dextral
pedipalpal segments of $ and $ paratypes (AMNH). 16. Chela, dorsal aspect, 8. 17. Chela, dorsal
aspect, 9. 18. Chela, external aspect, $. 19. Chela, ventrointernal aspect, 9. Scale bar = 5 mm.
2006
PRENDINI: SOUTH AFRICAN FLAT ROCK SCORPIONS
21
Figs. 20-21. Hadogenes polytrichobothrius n.sp., metasoma and telson of holotype 8 and paratype
9 (AMNH). 20. Lateral aspect, $. 21. Lateral aspect, 8. Scale bar =10 mm.
uterus or the gravid condition in females.
The elongated metasoma (longer than the
combined length of prosoma and metasoma),
a secondary sexual characteristic only ac¬
quired in the final instar male (Lamoral,
1979; Newlands, 1980; Prendini, 2001a), is
a further indication that male specimens are
adult (fig. 21, cf. female, fig. 20). In all spe¬
cies of Hadogenes, juvenile males and fe¬
males resemble each other, and adult fe¬
males, very closely in general morphological
features (besides the absence of a lobe and
notch on the pedipalp chela fingers) until the
final instar. The metasoma of the juvenile
male is also shorter than the combined length
of the prosoma and mesosoma.
Sexual dimorphism : The characters of pri¬
mary external sexual dimorphism are the un¬
divided genital operculum, which opens in a
single flap, in the female (fig. 9), compared
with the two unconnected sclerites, which
open independently and cover a pair of gen¬
ital papillae, in the male (fig. 8). Secondary
sexual characters observed in adult males,
compared with adult females and juveniles of
both sexes, are as follows (tables 3, 4): more
pronounced lobes on the fixed and movable
fingers of the chela, and a more pronounced
notch in the fixed finger; slightly flatter pedi¬
palp chelae; metasoma elongated, longer
than the combined length of the prosoma and
mesosoma (fig. 21); increased granulation of
the carapace (fig. 6), tergites and metasoma;
reduced granulation on the pedipalp chelae;
greater number of pectinal teeth (fig. 8).
Remarks : The male specimen from Doorn-
kop near Belfast (AMGS), described as H.
bicolor by Hewitt (1918), and provisionally
22
AMERICAN MUSEUM NOVITATES
NO. 3502
Figs. 22-23. Hadogenes polytrichobothrius n.sp., hemispermatophore of paratype $ (AMNH). 22.
Ectal aspect. 23. Ental aspect. Scale bar = 1 mm.
assigned to H. longimanus by Prendini
(2001a), is conspecific with H. polytricho¬
bothrius, as are the four specimens from
Steelpoort (SAMC C3901, C4275, C4276,
C4281), also provisionally assigned to H.
longimanus by Prendini (2001a). The correct
georeference for Doomkop [Doringkop, on
Farm Doornkop 356], in the Belfast District,
has been established as 25°30'S 29°55'E. The
georeference assigned by Prendini (2001a:
159), i.e. 25°55'S 30°16'E, refers to another
Doomkop, in the Carolina District. That par¬
ticular locality was visited during 2001 and
no Hadogenes were found there.
Distribution: Hadogenes polytrichoboth¬
rius is endemic to rocky outcrops and ridges
in the Steelpoort River valley and the middle
Olifants River valley, of the Limpopo Prov¬
ince (Sekhukhuneland District) and Mpu¬
malanga Province (Belfast, Lydenburg, Mid-
delburg districts), South Africa (fig. 1). The
southernmost locality record, Doornkop
[25°30'S 29°55'E], is approximately 150 km
southwest of the northernmost record in the
Potlake Nature Reserve [24° 15.160'S
29°54.649'E].
The known locality records of H. polytri¬
chobothrius fall within the following range
of altitudes (percentage of locality records
indicated in parentheses): 500-1000 m
(62.5%), 1000-1500 m (25%), 1500-2000 m
(12.5%).
The distributional range falls mostly
(87.5%) within the Mixed Bushveld vegeta¬
tion zone (Van Rooyen and Bredenkamp,
1998a) of the Savanna biome (Rutherford
and Westfall, 1994; Low and Rebelo, 1998).
However, the record from Doomkop falls
close to the boundary of the Moist Sandy
Highveld Grassland vegetation zone (Bre¬
denkamp and Van Rooyen, 1998) in the
Grassland biome (Rutherford and Westfall,
1994; Low and Rebelo, 1998).
The annual rainfall in the region inhabited
by this species varies from less than 650 mm
in the north of the distributional range, to 950
mm in the south, and is received mostly in
the summer (December to May). Tempera¬
tures range from — 10°C to 40°C (Breden¬
kamp and Van Rooyen, 1998; Van Rooyen
and Bredenkamp, 1998a).
Ecology: In common with all other spe¬
cies of Hadogenes, H. polytrichobothrius is
an obligate lithophile (Prendini, 2001b). This
species inhabits the narrow cracks and crev¬
ices of weathered granite and sandstone out-
2006
PRENDINI: SOUTH AFRICAN FLAT ROCK SCORPIONS
23
crops on gently sloping hillsides at moderate
altitudes in the Steelpoort River valley and
the middle Olifants River valley. Its distri¬
butional range is allopatric with that of its
closest relative, H. longimanus, which inhab¬
its rocky outcrops along the upper reaches of
the Olifants River valley, running parallel,
further to the south and west (Prendini,
2001a). It is also allopatric with H. bicolor,
which occurs at higher altitudes along the
Drakensberg escarpment, further north and
east.
Another liochelid, Opisthacanthus validus
Thorell, 1876, was collected in sympatry
with H. polytrichobothrius at Mapoch’s Grot-
te but the two species were not syntopic. Op¬
isthacanthus validus occupied humid habitats
in the shade of dense vegetation upslope,
whereas H. polytrichobothrius occupied dri¬
er, unshaded habitats at the base of the slope.
A scorpionid, Opistophthalmus glabrifrons
Peters, 1861, is the only other scorpion spe¬
cies thus far recorded in sympatry with H.
polytrichobothrius (I. Engelbrecht, personal
commun.).
Conservation Status: Hadogenes poly¬
trichobothrius is presently known from eight
localities, falling within seven QDS. As with
many other species of Hadogenes in southern
Africa, this species is threatened by habitat
destruction. Steelpoort, the type locality of
the species, is an open-cast vanadium mine.
At least six other large mines and several
granite quarries are situated in the Steelpoort
River valley, between Doomkop and Steel¬
poort. Most of the remaining land is privately
owned and cattle farming (rangeland) is the
dominant land-use. Much of the rangeland in
the northern part of this species’ range (part
of the former “Independent Homeland” of
Lebowa) is severely degraded by overgrazing
and soil erosion. Although one population of
H. polytrichobothrius is protected in the Pot-
lake Nature Reserve, the threat of mining,
quarrying, and other forms of habitat degra¬
dation, together with the restricted distribu¬
tional range of this species, which otherwise
falls entirely outside of existing protected ar¬
eas, warrants its assignment to the Vulnera¬
ble IUCN Red List Category. This species is
characterized by an acute restriction in both
its area of occupancy and number of known
localities. It would thus be prone to the ef¬
fects of human activities (or stochastic
events, the impact of which is increased by
human activities) within a very short period
of time in an unforeseeable future, and is ca¬
pable of becoming Critically Endangered or
even Extinct in a very short period.
Hadogenes soutpansbergensis , new species
Type Material: SOUTH AFRICA: Lim¬
popo Province: Soutpansberg District : Ho-
lotype S (AMNH), Vancoller’s Pass, Sout¬
pansberg [Farm Waterpoort 695, 22°55'S
29°37'E], xii.1990-i.1991, L. Prendini and
K.M.A. Prendini, in deep rock crevice. Para-
types: same data as holotype, 1 9 (AMNH);
same data, except “i.1996, L. Prendini and
J. Laing”, 1 juv. S (AMNH); same data, ex¬
cept “iii.2003, I. Engelbrecht and B. Wat¬
kins, removed from rock crevices at night,
located with UV detection”. Id 49 1 subad.
6 2 subad. 9 (AMNH), 1 subad. 9 (AMCC
139000).
Diagnosis: Hadogenes soutpansbergensis
is placed in the bicolor group on account of
the shape of metasomal segment I, which is
wider than it is high posteriorly. It appears
to be more closely related to H. newlandsi
than to the other three species in the bicolor
group. Adult males of both species exhibit
dense granulation on the telson and lateral
surfaces of metasomal segment V. This char¬
acter occurs in only a few other Hadogenes
species (e.g., H. granulatus Purcell, 1901 and
H. minor Purcell, 1899), and is potentially
synapomorphic for H. newlandsi, H. sout¬
pansbergensis and H. granulatus. Hadogenes
minor does not appear to be closely related
to these species and is presumed to have ac¬
quired this state independently.
Unlike the other the three species in the
bicolor group, the pedipalp chela of H. sout¬
pansbergensis and H. newlandsi lacks a pro¬
nounced lobe, distal to the notch in fixed Un¬
ger, and the metasoma of the adult male is
relatively longer (more than 55% of the total
length). Hadogenes soutpansbergensis and
H. newlandsi are further distinguished from
H. longimanus and H. polytrichobothrius by
the presence of only two trichobothria on the
internal surface of the pedipalp chela. Both
H. longimanus and H. polytrichobothrius ex-
24
AMERICAN MUSEUM NOVITATES
NO. 3502
hibit five or more internal trichobothria on
the chela.
Hadogenes soutpansbergensis is distin¬
guished from H. newlandsi by its longer me¬
tasoma in the adult male, longer, narrower,
flatter pedipalps, and higher trichobothrial
counts.
Etymology: The species name refers to
the Soutpansberg mountain range, where the
species is probably endemic.
Description: Measurements and counts in
the following description are recorded from
26 and 5$ (table 4).
Color. Uniformly dark in color, with ped¬
ipalps, legs, sternites, and telson only slightly
paler than the rest (figs. 24-27). Carapace,
chelicerae, tergites, and metasoma, Sepia
219; pedipalps and legs (dorsal surface), Ma¬
roon 31; legs (ventral surface), pectines, gen¬
ital operculum, sternites, and telson, Warm
Sepia 221 A.
Carapace : As for H. polytrichobothrius,
except median notch in anterior margin
weakly developed, and frontal lobes almost
entirely granular (figs. 28, 29).
Chelicerae : As for H. polytrichobothrius.
Pedipalps: As for H. polytrichobothrius,
but differing in the following respects. Femur
width 35.5% (33-38%) of length (table 5).
Patella ventroexternal carina granular to cos¬
tate granular (fig. 37). Patella width 53.5%
(51-56%) of length. Chela with weak, round¬
ed lobe on movable finger and correspond¬
ingly shallow notch in fixed finger; fixed fin¬
ger additionally with small, rounded lobe
proximal to notch, but without pronounced,
conical lobe distally (figs. 38, 39); dorsal and
ventrointernal intercarinal surfaces finely
granular, reticulate in 6, becoming smooth,
reticulate in 9. Chela height 49.5% (47-
52%) of width; chela width 51% (49-53%)
of length along ventroexternal carina in 6,
and 57.5% (57-58%) in 9; length movable
finger 97% (95-99%) of length along ven-
troextemal carina.
Trichobothria: Neobothriotaxic major,
type C (figs. 34-41; table 5), with the fol¬
lowing segment totals: femur, 3 (1 d\ 1 i; 1
e), patella, 80-106 (2 d; 1 i; 26-36 v; 51-67
e) and chela, 79-95 (69-85 manus; 10 fixed
finger, including 2 i). Total number of tricho¬
bothria per pedipalp, 162-204. Only femoral
trichobothria, trichobothria in the d and / se¬
ries of the patella, and trichobothria in the D,
d, e and i series of the chela are stable in
number and distribution. External and ventral
trichobothria of the chela and patella are nu¬
merically and distributionally too variable for
diagnostic purposes.
Me so soma: As for H. polytrichobothrius,
except as follows. Post-tergites of 6 covered
with very fine and even granulation, impart¬
ing matt appearance to all surfaces, except
median carina and submedian depressions,
which are smooth; post-tergites of 9 smooth
and shiny, becoming finely granular laterally.
Stemite VII length equal (100%) to width in
6, 83% (76-90%) in 9 (table 5; figs. 32, 33).
Pectines: As for H. polytrichobothrius
(figs. 30, 31), except pectinal teeth (left/
right): 20/20-21 (6), 16-17/16-17 (9).
Sternum: As for H. polytrichobothrius.
Genital operculum: As for H. polytricho¬
bothrius.
Legs: As for H. polytrichobothrius.
Metasoma and telson: As for H. polytri¬
chobothrius, except as follows. Dorsosub-
median carinae costate to costate granular on
segments II-V (9), costate granular on seg¬
ments II-IV, becoming granular on V (d),
II-IV each terminating distally with small
spiniform granule. Metasomal segment V,
lateral surfaces, and telson densely granular
in 6, smooth to sparsely granular in 9 (figs.
42, 43).
In addition, metasomal segments of adult
6 longer than in H. polytrichobothrius, with
morphometric differences as follows. Meta¬
somal segment I posterior height 92.5% (87-
98%) of width (table 5). Metasomal seg¬
ments I-V progressively increasing in length,
and decreasing in width, segment V width
66% (56-76%) of segment I width. Meta¬
soma slender, width percentage of length for
segment I, 24.5% (23-26%) in 6, 36.5%
(34-39%) in 9; for II, 12% (11-13%) in 6,
21.5% (19-24%) in 9; for III, 10.5% (10-
11%) in 6, 21% (18-24%) in 9; for IV, 9%
in 6, 17% (15-19%) in 9; and for V, 10%
(9-11%) in 6, 16.5% (15-18%) in 9. Telson
vesicle width 122% (111-133%) of metaso¬
mal segment V width; distinctly elongated in
6, oval in 9, with flattened dorsal surface
and rounded ventral surface, height 34.5%
(32-37%) of length. Aculeus short, 14.5%
(10-19%) of vesicle length, and sharply
2006
PRENDINI: SOUTH AFRICAN FLAT ROCK SCORPIONS
25
Figs. 24-27. Hadogenes soutpansbergensis n.sp., habitus of 8 and 9 paratypes (AMNH). 24. Dorsal
aspect, 8. 25. Ventral aspect, 8. 26. Dorsal aspect, $. 27. Ventral aspect, $. Scale bars = 10 mm.
curved. Total length of metasoma 168.5%
(148-190%) of combined length of prosoma
and mesosoma in 8 , but approximately equal
(97-104%) to combined length of prosoma
and mesosoma in 9.
Hemispermatophore : Doubled hook near
base of distal lamella; distal crest truncate
(figs. 44, 45).
Geographic variation : No significant var¬
iation.
Ontogenetic variation : As for H. polytri-
chobothrius.
26
AMERICAN MUSEUM NOVITATES
NO. 3502
Figs. 28-33. Hadogenes soutpansbergensis n.sp., carapace, pectines, and stemites of 6 and 9 para-
types (AMNH). 28 . Carapace, 6. 29 . Carapace, $. 30 . Pectines and basal piece, 6. 31 . Pectines and
basal piece, $. 32 . Sternite VII, 6. 33 . Stemite VII, $. Scale bars = 5 mm.
2006
PRENDINI: SOUTH AFRICAN FLAT ROCK SCORPIONS
27
Figs. 34 - 37 . Hadogenes soutpansbergensis
n.sp., carinae, trichobothria, and macrosetae on
dextral pedipalpal segments of paratype 9
(AMNH). 34. Femur, dorsal aspect. 35. Patella,
dorsal aspect. 36. Patella, external aspect. 37. Pa¬
tella, ventral aspect. Scale bar = 5 mm.
Sexual dimorphism : As for H. polytricho-
bothrius, except in this species no lobe is
present distal to the notch in the fixed finger
of the pedipalp chela, and the lobe on the
movable finger and notch in the fixed finger
are slightly less developed in the adult male
and than in the adult female (figs. 38, 39). In
addition, the pedipalp chelae are more gran¬
ular in the adult male, and the metasoma is
considerably more elongated, with segment
V and telson granular (fig. 43).
Distribution: Hadogenes soutpansber¬
gensis is probably endemic to the Soutpans-
berg mountain range in the Soutpansberg
District, Limpopo Province, South Africa
(fig. 1). It is currently known only from a
single locality, Vancoller’s Pass, a deep gorge
incised in the northern slopes of the Sout¬
pansberg by the Sand River, a tributary of
the Limpopo River. The species is expected
to be more widespread in the Soutpansberg,
however, particularly on the mesic southern
slopes of the mountain range. The xeric
northern slopes, which have been better sur¬
veyed for scorpions, are dominated by the
larger Hadogenes troglodytes (Peters, 1861)
and the two species are unlikely to occur in
sympatry. The occurrence of H. soutpans¬
bergensis at Vancoller’s Pass is presumed to
be an extension through the mountain range
associated with the Sand River. Uroplectes
flavoviridis Peters, 1861, otherwise observed
only on the southern side of the Soutpans¬
berg, is also found there.
The only known locality record of H. sout¬
pansbergensis occurs at an altitude between
700-800 m, in the Soutpansberg Arid Moun¬
tain Bushveld vegetation zone (Van Rooyen
and Bredenkamp, 1998b) of the Savanna bi-
ome (Rutherford and Westfall, 1994; Low
and Rebelo, 1998).
This species inhabits a moderately arid re¬
gion, where annual rainfall varies from 300
mm on the hot, dry northern slopes of the
Soutpansberg, to more than 500 mm on the
higher plateaux. Most rainfall is received in
the summer (December to May). Tempera¬
tures vary between 3°C and 44°C, with an
average of 23°C (Van Rooyen and Breden¬
kamp, 1998b).
Ecology: Hadogenes soutpansbergensis
is an obligate lithophile (Prendini, 2001b).
This species inhabits very deep cracks and
crevices in weathered sandstone outcrops, of¬
ten on sheer cliff faces. One specimen was
collected about 2 m above ground level in a
crack in a tree trunk (I. Engelbrecht, personal
commun.). The distributional range of H.
soutpansbergensis is allopatric with that of
its closest relatives, H. newlandsi, which in¬
habits scattered inselbergs south of the Sout¬
pansberg, and H. bicolor, which occurs along
the Drakensberg escarpment, southeast of the
Soutpansberg (Prendini, 2001a). Hadogenes
soutpansbergensis has not been collected in
sympatry with H. troglodytes, which is very
common in drier habitats along the northern
slopes of the Soutpansberg (e.g., at Farm
28
AMERICAN MUSEUM NOVITATES
NO. 3502
Figs. 38-41. Hadogenes soutpansbergensis n.sp., carinae, trichobothria, and macrosetae on dextral
pedipalpal segments of holotype 8 and paratype 9 (AMNH). 38. Chela, dorsal aspect, 8. 39. Chela,
dorsal aspect, 9. 40 . Chela, external aspect, 9. 41 . Chela, ventrointemal aspect, 9. Scale bar = 5 mm.
2006
PRENDINI: SOUTH AFRICAN FLAT ROCK SCORPIONS
29
Figs. 42-43. Hadogenes soutpansbergensis n.sp., metasoma and telson of $ and $ paratypes
(AMNH). 42 . Lateral aspect, $. 43 . Lateral aspect, $. Scale bar =10 mm.
Figs. 44-45. Hadogenes soutpansbergensis n.sp., hemispermatophore of paratype 6 (AMNH). 44 .
Ectal aspect. 45 . Ental aspect. Scale bar = 1 mm.
30
AMERICAN MUSEUM NOVITATES
NO. 3502
Rochdale 700), and north of the mountain
range.
At Vancoller’s Pass, H. soutpansbergensis
has been collected in sympatry with several
other scorpion species: the buthids, Hotten-
totta trilineatus (Peters, 1861), Parabuthus
transvaalicus, Uroplectes planimanus
(Karsch, 1879), U. flavoviridis, and U. vit-
tatus (Thorell, 1876); the liochelid, Opistha-
canthus asper (Peters, 1861); and the scor-
pionid, Opistophthalmus lawrencei New-
lands, 1969. Uroplectes vittatus and O. asper
are strictly corticolous, whereas H. trilinea¬
tus, U. planimanus, and O. lawrencei prefer
flatter rocky areas.
Conservation Status: Unlike many other
species of Hadogenes, H. soutpansbergensis
does not appear to be threatened by habitat
destruction. This species is presently known
from only a single locality, but may be wide¬
spread on the southern slopes of the Sout-
pansberg, a region of low agricultural poten¬
tial where there are several provincial and
private nature reserves. The following pro¬
tected areas should be surveyed for the pres¬
ence of H. soutpansbergensis , which is sus¬
pected to occur there: Happy Rest Nature Re¬
serve, Lesheba Wilderness Area, Medike Pri¬
vate Nature Reserve. Until more data
become available on the distribution, ecology
and abundance of Hadogenes soutpansber¬
gensis, this species is assigned to the Data
Deficient category of the IUCN Red List.
ACKNOWLEDGMENTS
Financial support for this research was
provided by the Foundation for Research De¬
velopment, South Africa, and the Richard
Lounsbery Foundation, USA. The following
people and institutions provided permission
to collect scorpions in South Africa: Fim-
popo Province Environmental Affairs and
Tourism (permit issued by Deon von Wiel-
ligh); Mpumalanga Parks Board (permit is¬
sued by Koos de Wet); the staff of the Ben
Favin, Blyde River Canyon, Botshabelo and
Potlake Nature Reserves. The following peo¬
ple participated on the expeditions and per¬
sonally collected some of the specimens: Ian
Engelbrecht and Bronwyn Watkins; the late
Martin Filmer; Marco MacFarlane; Ken
Prendini; Elizabeth Scott. Additional speci¬
mens were collected by John Faing, Alistair
Mathie, and Nic MacFean. The following
people assisted with the loan of types and
additional specimens and/or allowed access
to the collections under their care during my
visits: Sarah Gess (AMGS); Charles Gris¬
wold and Darrell Ubick (CAS); Ansie Dip-
penaar-Schoeman and Annette van den Berg
(NCA); Debbie Jennings (NMSA); Margie
Cochrane (SAMC); Klaas Manamela and
Barbara Dombrowsky (TMSA); Jason Dun¬
lop and Shahin Nawai (ZMHB); Hieronymus
Dastych (ZMUH). This is the eighth paper
that includes material from the Alexis Har-
ington scorpion collection and I reiterate my
appreciation to those involved in its transfer-
ral to the AMNH, listed by name in previous
papers. The following people and institutions
provided GIS coverages and/or permission to
use them in the spatial analyses: the National
Botanical Institute of South Africa; the Water
Research Commission (Department of Water
Affairs and Forestry); Dion Marais (Depart¬
ment of Environmental Affairs and Tourism,
South Africa); Mark Horan, Steve Fynch and
Roland Schulze (Computing Centre for Wa¬
ter Research, University of Natal); Helen de
Klerk (Western Cape Nature Conservation).
Randy Mercurio, Connie Cai and Melanie
Ng (AMNH) took the photographs and re¬
corded the morphometric and meristic data
in this paper, and Steve Thurston (AMNH)
prepared the photographic plates. Warren E.
Savary and W. David Sissom provided con¬
structive criticisms of an earlier draft of the
manuscript.
REFERENCES
Bredenkamp, G.J., and N. van Rooyen. 1998.
Moist sandy high veld grassland. In A.B. Low
and A.G. Rebelo (editors), Vegetation of South
Africa, Lesotho and Swaziland: 42. Pretoria:
Department of Environmental Affairs and Tour¬
ism.
Fet, V. 2000. Family Ischnuridae Simon, 1879. In
V. Fet, W.D. Sissom, G. Lowe, and M.E.
Braunwalder, Catalog of the scorpions of the
world (1758-1998): 383-408. New York: New
York Entomological Society.
Hewitt, J. 1918. A survey of the scorpion fauna
of South Africa. Transactions of the Royal So¬
ciety of South Africa 6: 89-192.
International Union for the Conservation of Na¬
ture (IUCN). 2001. IUCN red list categories
2006
PRENDINI: SOUTH AFRICAN FLAT ROCK SCORPIONS
31
and criteria. Version 3. 1. Gland, Switzerland:
IUCN Species Survival Commission, IUCN.
Available at: http://www.iucn.org/themes/ssc/
redhsts/RLcats200 lbooklet.html.
Kovarfk, F. 1998. Stfri [Scorpions]. Jihlava: Mad-
agaskar, 175 pp. [in Czech]
Lamoral, B.H. 1979. The scorpions of Namibia
(Arachnida: Scorpionida). Annals of the Natal
Museum 23: 498-783.
Lamoral, B.H., and S.C. Reynders. 1975. A cat¬
alogue of the scorpions described from the
Ethiopian faunal region up to December 1973.
Annals of the Natal Museum 22: 489-576.
Lawrence, R.F. 1955. Solifugae, scorpions and Pe-
dipalpi, with checklists and keys to South Af¬
rican families, genera and species. Results of
the Lund University Expedition in 1950-1951.
In B. Hanstrom, P. Brinck, and G. Rudebeck
(editors), South African Animal Life 1: 152-
262. Uppsala: Almqvist and Wiksells.
Lourengo, W.R. 1999. Considerations taxono-
miques sur le genre Hadogenes Kraepelin,
1894; creation de la sous-famille des Hadogen-
inae n. subfam., et description d’une espece
nouvelle pour F Angola (Scorpiones, Scorpion-
idae, Hadogeninae). Revue Suisse de Zoologie
106: 929-938.
Low, A.B., and A.G. Rebelo. 1998. Vegetation of
South Africa, Lesotho and Swaziland, 2 nd edi¬
tion. Pretoria: Department of Environmental
Affairs and Tourism, 83 pp.
Newlands, G. 1972a. Ecological adaptations of
Kruger National Park scorpionids (Arachnida:
Scorpiones). Koedoe 15: 37-48.
Newlands, G. 1972b. A description of Hadogenes
lawrencei sp. nov. (Scorpiones) with a checklist
and key to the South West African species of
the genus Hadogenes. Madoqua (II) 1: 133-
MO.
Newlands, G. 1978. Arachnida (except Acari). In
M.J.A. Werger (editor), Biogeography and
ecology of southern Africa: 677-684. The
Hague: W. Junk.
Newlands, G. 1980. A revision of the scorpion
genus Hadogenes Kraepelin 1894 (Arachnida:
Scorpionidae) with a checklist and key to the
species. Unpublished M.Sc. Thesis. Potchefst-
room: Potchefstroom University for C.H.E.,
189 pp.
Newlands, G., and A.C. Cantrell. 1985. A re-ap-
praisal of the rock scorpions (Scorpionidae:
Hadogenes ). Koedoe 28: 35-45.
Newlands, G., and L. Prendini. 1997. Redescrip¬
tion of Hadogenes zumpti Newlands and Can¬
trell 1985: an unusual rock scorpion (Scorpi¬
ones, Ischnuridae) from the Richters veld, South
Africa. South African Journal of Zoology 32:
76-81.
Prendini, L. 2000. Phylogeny and classification of
the Superfamily Scorpionoidea Latreille 1802
(Chelicerata, Scorpiones): an exemplar ap¬
proach. Cladistics 16: 1-78.
Prendini, L. 2001a. Two new species of Hado¬
genes (Scorpiones, Ischnuridae) from South Af¬
rica, with a redescription of Hadogenes bicolor
and a discussion on the phylogenetic position
of Hadogenes. Journal of Arachnology 29:
146-172.
Prendini, L. 2001b. Substratum specialization and
speciation in southern African scorpions: the
Effect Hypothesis revisited. In V. Fet and PA.
Selden (editors), Scorpions 2001. In Memoriam
Gary A. Polis: 113-138. Burnham Beeches,
UK: British Arachnological Society.
Prendini, L. 2005a. Scorpion diversity and distri¬
bution in southern Africa: pattern and process.
In B.A. Huber, B.J. Sinclair, and K.-H. Lampe
(editors), African biodiversity: molecules, or¬
ganisms, ecosystems: 25-68. New York:
Springer.
Prendini, L. 2005b. On Hadogenes angolensis Lou¬
rengo, 1999 syn. n. (Scorpiones, Liochelidae),
with a redescription of H. taeniurus (Thorell,
1876). Revue Suisse de Zoologie 112: 1-28.
Purcell, W.F. 1899. New South African scorpions
in the collection of the South African Museum.
Annals of the South African Museum 1: 433-
438.
Rutherford, M.C., and R.H. Westfall. 1994. Bi-
omes of southern Africa: an objective catego¬
rization. 2nd edition. Memoirs of the Botanical
Survey of South Africa 63: 1-94.
Schulze, R.E. (with M. Maharaj, S.D. Lynch, B.J.
Howe, and B. Melvil-Thomson). 1997. South
African atlas of agrohydrology and -climatology.
Pretoria: Water Research Commission Report.
Smithe, F.B. 1974. Naturalist’s color guide sup¬
plement. New York: American Museum of Nat¬
ural History, xiii+229 pp.
Smithe, F.B. 1975. Naturalist’s color guide. New
York: American Museum of Natural History,
unpaginated.
Smithe, F.B. 1981. Naturalist’s color guide. Part
III. New York: American Museum of Natural
History, 37 pp.
Soleglad, M.E., and V. Fet. 2003. The scorpion
sternum: structure and phylogeny (Scorpiones:
Orthosterni). Euscorpius 5: 1-34.
Vachon, M. 1974 “1973”. Etude des caracteres
utilises pour classer les families et les genres
de scorpions (Arachnides). 1. La trichobothri-
otaxie en arachnologie. Sigles trichobothriaux
et types de trichobothriotaxie chez les scorpi¬
ons. Bulletin du Museum National d’Histoire
Naturelle (Paris), ser. 3, 140: 857-958.
Van Rooyen, N., and G.J. Bredenkamp. 1998a.
32
AMERICAN MUSEUM NOVITATES
NO. 3502
Mixed Bush veld. In A.B. Low and A.G. Rebelo
(editors), Vegetation of South Africa, Lesotho
and Swaziland: 26. Pretoria: Department of En¬
vironmental Affairs and Tourism.
Van Rooyen, N., and G.J. Bredenkamp. 1998b.
Soutpansberg Arid Mountain Bushveld. In A.B.
Low and A.G. Rebelo (editors), Vegetation of
South Africa, Lesotho and Swaziland: 21. Pre¬
toria: Department of Environmental Affairs and
Tourism.
Complete lists of all issues of the Novitates and the Bulletin are available at World Wide Web
site http://library.amnh.org/pubs. Inquire about ordering printed copies via e-mail from
scipubs@amnh.org or via standard mail from: American Museum of Natural History, Library—
Scientific Publications, Central Park West at 79th St., New York, NY 10024. TEL: (212) 769-
5545. PAX: (212) 769-5009.
® This paper meets the requirements of ANSI/NISO Z39.48-1992 (Permanence of Paper).
Live Music Archive
Librivox Free Audio
Metropolitan Museum
Cleveland Museum of Art
Internet Arcade
Console Living Room
Books to Borrow
Open Library
TV News
Understanding 9/11