A T AMERICAN MUSEUM
JSovitates
PUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORY
CENTRAL PARK WEST AT 79TH STREET, NEW YORK, NY 10024
Number 3564, 44 pp., 77 figures, 2 tables May 16, 2007
The genus Brachistosternus (Scorpiones: Bothriuridae)
in Chile, with Descriptions of Two New Species
ANDRES A. OJANGUREN AFFILASTRO, 1
CAMILO I. MATTONI, 2 AND LORENZO PRENDINI 3
ABSTRACT
We review the taxonomy of the Brachistosternus Pocock, 1893 scorpions of Chile, providing
revised diagnoses, comprehensive distribution maps (based on all known locality records), and an
illustrated key to all Chilean species of the genus. Two new species, Brachistosternus (Leptosternus)
chango, n.sp., and Brachistosternus ( Leptosternus ) kamanchaca, n.sp., are described from northern
Chile. The phylogenetic affinities of B. chango are unclear. Some characters suggest that this
species may be related to Brachistosternus ( L .) artigasi Cekalovic, 1974 but others suggest that it
may be related to Brachistosternus ( L .) roigalsinai Ojanguren Affilastro, 2002. Brachistosternus
kamanchaca, in contrast, appears to be closely related to Brachistosternus ( L .) donosoi Cekalovic,
1974 and other species from the plains of northern Chile and southern Peru.
RESUMEN
Se revisa la taxonomia de los escorpiones del genero Brachistosternus Pocock, 1893 de Chile, se
brindan diagnosis revisadas, mapas de distribution completos (basados en todos los registros
conocidos) y una clave ilustrada de todas las especies. Se describe a Brachistosternus (. Leptosternus)
chango, n.sp., y a Brachistosternus ( Leptosternus ) kamanchaca, n.sp., del norte de Chile. Las
relaciones filogeneticas de B. chango son poco claras. Algunos caracteres de esta especie sugieren
que puede estar relacionada con Brachistosternus ( L .) artigasi Cekalovic, 1974, aunque otros
parecerian relacionarla con Brachistosternus (L .) roigalsinai Ojanguren Affilastro, 2002.
Brachistosternus kamanchaca por otro lado, se encuentra estrechamente relacionada con
1 Division Aracnologia, Museo Argentino de Ciencias Naturales ‘Bernardino Rivadavia’ (CONICET), Av. Angel
Gallardo 470, 1405 DJR Buenos Aires, Argentina (andres.ojanguren@gmail.com and ojanguren@macn.gov.ar).
2 Division of Invertebrate Zoology, American Museum of Natural History (cmattoni@amnh.org).
3 Division of Invertebrate Zoology, American Museum of Natural History (lorenzo@amnh.org), corresponding author.
Copyright © American Museum of Natural History 2007
ISSN 0003-0082
2
AMERICAN MUSEUM NOVITATES
NO. 3564
Brachistosternus ( L .) donosoi Cekalovic, 1974, asi como con otras especies de llanura del norte de
Chile y sur del Peru.
INTRODUCTION
Although the scorpion fauna of Chile is
among the most diverse in South America, it
received little attention until recent years.
Brachistosternus Pocock, 1893, the most di¬
verse genus in the country, comprises numer¬
ous extremely range-restricted and microhab-
itat-specific species, many of which have been
described in the past 5 years (Ochoa, 2002;
Ochoa and Acosta, 2002; Ojanguren
Affilastro, 2002b, 2004b, 2005a; Agusto et
al., in press).
During recent fieldwork in central and
northern Chile, we collected several unde¬
scribed species of Brachistosternus, two of
which, Brachistosternus chango, n.sp., and
Brachistosternus kamanchaca, n.sp., are de¬
scribed below. The description of these species
raises the number of species in the genus to 38
(table 1). Brachistosternus is the second most
speciose scorpion genus in South America,
after Tityus C.L. Koch, 1836 (Buthidae C.L.
Koch, 1837; Fet and Lowe, 2000), and has
evidently undergone a radiation similar to the
mostly fossorial genera Opistophthalmus C.L.
Koch, 1837 (Scorpionidae Latreille, 1802),
Urodacus Peters, 1861 (Urodacidae Pocock,
1893), and Paruroctonus Werner, 1934
(Vaejovidae Thorell, 1876) in the arid zones
of southern Africa, Australia, and North
America, respectively. Presumably, the radia¬
tion of Brachistosternus, like those of
Opistophthalmus, Urodacus, and Paruroct¬
onus, can in large part be explained as
a consequence of allopatric speciation pro¬
moted by stenotopic substratum requirements
(Prendini, 2001).
The patterns of distribution and diversity
observed among the Brachistosternus species
inhabiting the arid to semi-arid regions of
Chile contrast starkly with those among
species inhabiting the arid plains of
Argentina, most of which are widely distrib¬
uted and somewhat less microhabitat specific
(Maury, 1974; Roig Alsina and Maury, 1981,
1984; Ojanguren Affilastro, 2001, 2002a,
2002b, 2003a, 2004a, 2005b, Ojanguren
Affilastro and Roig Alsina, 2001). This
observation is somewhat paradoxical because
most of the Chilean Brachistosternus species
appear to be closely related to those from
Argentina. However, the Brachistosternus spe¬
cies from the central arid areas (Coquimbo
Region) of Chile appear to be more closely
related to the Andean species of the genus,
most of which are also very range restricted
and microhabitat specific.
Of the two new species described in the
present contribution, B. kamanchaca is among
the most widespread Chilean species of the
genus, occupying most of the Atacama Region
and probably part of the neighboring
Antofagasta and Coquimbo Regions. In
contrast, B. chango is known from only two
localities.
Several poorly known Chilean species of
Brachistosternus were also collected during
our fieldwork in central and northern Chile.
We take the opportunity to report new records
and observations for these species in the
present contribution, as well as providing
revised diagnoses, comprehensive distribution
maps (based on all known locality records),
and an illustrated key for all Chilean species of
the genus.
MATERIALS AND METHODS
Descriptive terminology follows Maury
(1974) for the hemispermatophores (figs. 27
and 30); Vachon (1974) for the trichobothria;
Cekalovic (1973) for the androvestigia (a pair
of glands situated on the dorsal surface of
metasomal segment V in the adult male); Roig
Alsina and Maury (1981) for the male telson
gland; Stahnke (1970) for the metasomal
carinae, abbreviated as follows: DL: dorsolat¬
eral, LIM: lateral inframedian, LSM: lateral
supramedian, PL: paramedian, PM: paralat-
eral, VL: ventrolateral, VSM: ventrosubme-
dian, and VM: ventromedian; and Francke
(1977) for the pedipalpal carinae, abbreviated
as follows: DI: dorsal internal, DE: dorsal
external, and VI: ventral internal.
Abbreviations of collections from which ma¬
terial was studied are as follows: AMNH:
American Museum of Natural History, New
York; ARA: Arturo Roig Alsina personal
collection, Buenos Aires, Argentina; CDA:
2007 OJANGUREN AFFILASTRO ET AL.: CHILEAN BRACHISTOSTERNUS SCORPIONS
3
Catedra de Diversidad Animal I, Universidad
de Cordoba, Argentina; LEULS: Laboratorio
de Entomologia, Universidad de La Serena,
Chile; MACN-Ar: Museo Argentino de
Ciencias Naturales ‘Bernardino Rivadavia’,
National Arachnological Collection,
Argentina; MNHN: Museum National
DHistoire Naturelle, Paris, France; MUSM:
Museo de historia Natural, Universidad
Nacional Mayor de San Carlos, Lima, Peru;
MZUC: Museo Zoologico del Instituto de
Biologia de la Universidad de Concepcion,
Chile; and ZMH: Zoologisches Institut und
Zoologisches Museum, Universitat Hamburg,
Germany. Scorpion tissue samples are stored
(in the vapor phase of liquid nitrogen at
— 150°C) in the Ambrose Monell Collection
for Molecular and Microbial Research
(AMCC) at the AMNH. Illustrations were
produced using a Leitz Wetzlar stereomicro¬
scope and camera lucida. Measurements,
taken using an ocular micrometer, are re¬
corded in millimeters. Hemispermatophores
were dissected from surrounding tissues
and observed in 80% ethanol. Distribution
maps were generated using ArcMap 9.0
(Enviromental Systems Research Institute
[ESRI], Redlands, California), by superimpos¬
ing the point locality records of species on
coverages depicting the political boundaries
and topography of Chile. The topographic
coverage was generated from a digital eleva¬
tion model file (1 arc degree resolution) from
the United States Geological Survey (USGS)
website (http://edcdaac.usgs.gov:80/gtopo30/
gtopo30.asp). Point locality records were
georeferenced in the field with a portable
Global Positioning System (Garmin® Etrex
and Garmin® II) or retrospectively using
the GeoNet Names Server (GNS, http://
gns www .nga .mil/geonames/GN S/index .j sp).
SYSTEMATICS
FAMILY BOTHRIURIDAE SIMON, 1880
GENUS BRACHISTOSTERNUS POCOCK, 1893
Key to identification of the Chilean species
OF BRACHISTOSTERNUS POCOCK, 1893:
1. Pedipalp patella with 5-7 ventral trichobothria
(fig. 42); hemispermatophore, cylindrical apo¬
physis dorsoventrally compressed (fig. 36)
Brachistosternus (Brack istos tern us) ehrenbergii
- Pedipalp patella with 3 ventral trichobothria
(fig. 43); hemispermatophore, cylindrical
apophysis terete (figs. 1, 28, 37-41).
.... Brachistosternus (Leptosternus ) ... 2
2. Basitarsi I and II, prolateral pedal spurs
vestigial or absent (fig. 52); hemispermato¬
phore, internal structures absent. 3
- Basitarsi I, prolateral pedal spur well devel¬
oped, equal to retrolateral pedal spur
(fig. 53); hemispermatophore, internal struc¬
tures well developed.4
3. Metasomal segment V, VM carina present in
posterior half of segment; androvestigia
absent. Brachistosternus (L. ) sciosciae
- Metasomal segment V, VM carina extending
entire length of segment; androvestigia pres¬
ent, small. Brachistosternus sp. 1
4. Pedipalp femur with two macrosetae associated
with d and e trichobothria (fig. 8). 5
- Pedipalp femur with one macroseta associated
with d and e trichobothria (fig. 9). 6
5. Metasomal segments I-IV, ventral surface with
pair of VSM stripes; segment V, androvesti¬
gia (male) very large, occupying >50% of
dorsal surface (fig. 4).
. Brachistosternus ( L .) chango, n.sp.
- Metasomal segments I-IV, ventral surface with
single VM stripe; segment V, androvestigia
medium-sized, occupying <50% of dorsal
surface (fig. 46).
. Brachistosternus ( L .) artigasi
6. Metasomal segment V, VM carina conspicu¬
ously elevated, extending entire length of
segment; androvestigia very large, occupying
>50% of dorsal surface; hemispermatophore,
distal lobe elongated (fig. 37).
. Brachistosternus (L .) roigalsinai
- Metasomal segment V, VM carina, if present,
not conspicuously elevated, seldom extending
entire length of segment; androvestigia small
to medium-sized, occupying <50% of dorsal
surface (figs. 31, 49); hemispermatophore,
distal lobe not elongated (fig. 28).7
7. Metasomal segments I-IV, ventral surface with
pair of VSM stripes (fig. 54). 8
- Metasomal segments I-IV, ventral surface
unpigmented or with single VM stripe
(figs. 55, 56). 9
8 Metasomal segment V, VM carina extending
entire length of segment (fig. 45).
. Brachistosternus (L .) cekalovici
- Metasomal segment V, VM carina absent. . .
. Brachistosternus (L.) negrei
9. Metasomal segment IV densely granular (male)
(fig. 47), slightly granular (female); segment
V, androvestigia narrow and elongated
4
AMERICAN MUSEUM NOVITATES
NO. 3564
(fig. 49); hemispermatophore, internal spines
well developed (fig. 40). 10
- Metasomal segment IV smooth or slightly
granular (male) (fig. 48), smooth (female);
segment V, androvestigia medium-sized
(fig. 31); hemispermatophore, internal spines
poorly developed (fig. 30) or absent
(fig. 41). 11
10. Metasoma unpigmented, tergites unpigmented
or with lateral margins faintly spotted;
pedipalp chela, length/height ratio 2.9-3.11
(male), 3.13 (female).
. Brachistosternus (L .) mattonii
- Metasomal segment V pigmented, with single
VM and paired VL stripes ventrally, and two
posterolateral and single antero-median spots
dorsally; tergites I-VI densely pigmented,
with transverse stripe extending most of
segment; pedipalp chela, length/height ratio
3.25-3.6 (male), 3.02-3.21 (female).
. Brachistosternus ( L .) ochoai
11. Metasoma unpigmented; tergites unpigmented
or with lateral margins faintly spotted....
. Brachistosternus ( L .) donosoi
- Metasoma with single VM and pair of VL
stripes, at least on segments III-V; tergites I-
VI with transverse stripe or two to three dark
spots. 12
12. Metasomal segments I-IV, VM stripe very
narrow, not joining VL stripes (fig. 56) . . .
. Brachistosternus (L .) chilensis
- Metasomal segments I-IV, VM stripe broad,
joining VL stripes, at least on segment IV
(fig. 55). 13
13. Metasomal segment V with 18 or more ventral
macrosetae. 14
- Metasomal segment V with 17 or fewer ventral
macrosetae. 15
14. Metasomal segment V, VM carina absent;
telson gland well developed, divided into
two separate halves (fig. 50).
. Brachistosternus (L .) perettii
- Metasomal segment V, VM carina restricted to
posterior half of segment; telson gland poorly
developed, not divided into two separate
halves. Brachistosternus (L .) prendinii
15. Metasomal segment V, VM carina absent. . .
. Brachistosternus sp. 2
- Metasomal segment V, VM carina present 16
16. Metasomal segment V, VM carina restricted to
posterior half of segment (fig. 32); hemisper¬
matophore, internal spines reduced to small
group of granules (fig. 30).
. Brachistosternus (L .) kamanchaca
- Metasomal segment V, VM carina extending
entire length of segment; hemispermato¬
phore, internal spines absent (fig. 41) . . 17
17. Metasomal segments I—III, VL and VM pig¬
mentation stripes joining at posterior mar¬
gin . 18
- Metasomal segments I—III, VL and VM pig¬
mentation stripes not joining at posterior
margin. 19
18. Pedipalp chela, length/width ratio 3.7-3.9
(male). . . . Brachistosternus (L.) quiscapata
- Pedipalp chela, length/width ratio 4.8-5.09
(male). Brachistosternus sp. 3
19. Metasomal segment V, length/width ratio 1.7-
1.8. Brachistosternus (L .) piacentinii
- Metasomal segment V, length/width ratio 1.8-
1.95. Brachistosternus (L .) kovariki
SUBGENUS BRACHISTOSTERNUS POCOCK, 1893
Brachistosternus ( B .) ehrenbergii
(Gervais, 1841)
figures 36, 42, 57
Scorpio ehrenbergii Gervais, 1841: 282.
Scorpio glaber Gervais, 1841: 285 (synonymized by
Simon, 1880: 397).
Telegonus politus L. Koch, 1867: 234-235 (synon¬
ymized by Kraepelin, 1894: 216).
Brachistosternus ( Brachistosternus ) ehrenbergii :
Lowe and Fet, 2000: 48 (complete reference list
until 1998); Ochoa, 2002: 55; Ojanguren
Affilastro, 2003a: 327-328, 2005b: 145-146;
Ochoa, 2005: 51, 57.
Brachistosternus ehrenbergii : Prendini, 2000: 41;
Soleglad and Fet, 2003a: 5, 2003b: 6; Acosta
and Ochoa, 2002: 18; Prendini, 2003a: 154-156,
169, 2003b: 242; Ojanguren Affilastro, 2005a:
191; Prendini and Wheeler, 2005: 454.
Type Material: Syntypes (MNHN RS.
0615, RS.0616), PERU, Callao [12°04'S
77°09'W], M. Gaudichaud.
New Records: CHILE: Region I (Tara-
paca): Arica Province : Quebrada Vitor,
18°49'41.4"S 70°08'17.6"W, 17.L2005, C.
Mattoni, A. Ojanguren, and J. Ochoa,
435 m, 1 juv. (AMNH). Iquique Province :
Quebrada Camarones, 2-10 km from coast,
19°10'48.1"S 70°13'46.8"W, 17.L2005, 5-50 m,
C. Mattoni, A. Ojanguren, and J. Ochoa, 1 2,
2 juv. (CDA), 6 juv. (MACN), 12, 7 juv.
(AMNH), 1 juv. $, 1 juv. 2 (AMCC 159656).
Diagnosis: Brachistosternus ( B .) ehrenber¬
gii is the only Chilean species of subgenus
Brachistosternus and may be distinguished
from all other Chilean Brachistosternus species
on the basis of its trichobothrial pattern,
which exhibits five to seven ventral trichobo-
2007 OJANGUREN AFFILASTRO ET AL.: CHILEAN BRACHISTOSTERNUS SCORPIONS
5
TABLE 1
The Described and Accepted Species of Brachistostemus Pocock, 1893 (Scorpiones: Bothriuridae),
with Known Countries of Distribution
Subgenus Brachistostemus Pocock, 1893
Brachistostemus (B.) ehrenbergii (Gervais, 1841)
Brachistostemus ( B .) pegnai Cekalovic, 1969
Subgenus Ministernus Francke, 1985
Brachistostemus (M.) andinus Chamberlin, 1916
Brachistostemus (M.) ferrugineus (Thorell, 1876)
Brachistostemus (M.) peruvianus Toledo Piza, 1974
Brachistostemus (M.) simonae Lourengo, 2000
Chile, Peru
Ecuador
Peru
Argentina, Bolivia, Brazil, Paraguay
Peru
Brazil
Subgenus Leptosternus Maury, 1973
Brachistostemus (L.) alienus Lonnberg, 1898
Brachistostemus (L.) angustimanus Ojanguren Affilastro & Roig Alsina, 2001
Brachistostemus (L.) artigasi Cekalovic, 1974
Brachistostemus ( L .) castroi Mello-Leitao, 1941
Brachistostemus (L.) cekalovici Ojanguren Affilastro, 2005
Brachistostemus ( L .) chango, n.sp.
Brachistostemus (L.) chilensis Kraepelin, 1911
Brachistostemus (L.) donosoi Cekalovic, 1974
Brachistostemus (L.) galianoae Ojanguren Affilastro, 2002
Brachistostemus (L.) intermedius Lonnberg, 1902
Brachistostemus ( L .) kamanchaca, n.sp.
Brachistostemus (L.) kovariki Ojanguren Affilastro, 2003
Brachistostemus (L.) mattonii Ojanguren Affilastro, 2005
Brachistostemus ( L .) montanus Roig Alsina, 1977
Brachistostemus (L.) multidentatus Maury, 1984
Brachistostemus ( L .) negrei Cekalovic, 1975
Brachistostemus (L.) ninapo Ochoa, 2004
Brachistostemus (L.) ochoai Ojanguren Affilastro, 2004
Brachistostemus (L.) paulae Ojanguren Affilastro, 2003
Brachistostemus ( L .) pentheri Mello-Leitao, 1931
Brachistostemus (L. ) perettii Ojanguren Affilastro & Mattoni, 2006
Brachistostemus (L.) piacentinii Ojanguren Affilastro, 2003
Brachistostemus (L.) prendinii Ojanguren Affilastro, 2003
Brachistostemus (L.) quiscapata Ochoa & Acosta, 2002
Brachistostemus (L.) roigalsinai Ojanguren Affilastro, 2002
Brachistostemus (L.) sciosciae Ojanguren Affilastro, 2002
Brachistostemus (L.) telteca Ojanguren Affilastro, 2000
Brachistostemus ( L .) titicaca Ochoa & Acosta, 2002
Brachistostemus (L.) turpuq Ochoa, 2002
Brachistostemus (L.) weijenberghi (Thorell, 1876)
Brachistostemus (L.) zambrunoi Ojanguren Affilastro, 2002
Argentina
Argentina
Chile
Chile
Chile
Chile
Chile
Chile
Bolivia
Argentina, Bolivia
Chile
Chile
Chile
Argentina
Argentina
Chile
Peru
Chile
Argentina
Argentina
Chile
Bolivia, Chile
Chile
Chile, Peru
Chile
Chile
Argentina
Bolivia, Peru
Peru
Argentina
Argentina
Species not placed in a subgenus
Brachistostemus holmberghi Carbonell, 1923
Argentina?
thria on the pedipalp patella (fig. 42). All
other Chilean Brachistostemus species possess
only three ventral trichobothria in this series
(fig. 43). Brachistostemus ehrenbergii may also
be distinguished from other Chilean species of
the genus according to the shape of the
cylindrical apophysis of the hemispermato-
phore, which is dorsoventrally flattened in B.
ehrenbergii (fig. 36), but cylindrical in other
Chilean species of the genus (figs. 1, 3, 28, 30,
6
AMERICAN MUSEUM NOVITATES
NO. 3564
37-41). The basal triangle of the hemisperma-
tophore of B. ehrenbergii is more developed
than those of other Chilean species of the
genus. Only B. roigalsinai possesses a similarly
elongated and sclerotized but less developed
basal triangle. Brachistosternus ehrenbergii is
the only Brachistosternus species known to
possess a stridulatory organ, similar to that
described by Acosta and Maury (1990) in
Timogenes Simon, 1880, and formed by a small
scattered group of granules on the medial
surface of each pretergite. Brachistosternus
ehrenbergii is the second largest bothriurid;
specimens may reach up to 10 cm in total
length. Only Timogenes elegans (Mello-Leitao,
1931) is slightly larger, some female specimens
reaching about 12 cm (Ojanguren Affilastro,
2005b).
Distribution: Although B. ehrenbergii has
been reported from Argentina, Chile, Bolivia,
Peru, and Ecuador, it is actually restricted to
the Pacific desert of northern Chile and
southern Peru (Ochoa, 2005; fig. 57).
Notes: Gervais (1841) described two spe¬
cies in the same paper: Scorpio ehrenbergii and
S. glaber. Simon (1880), the first revisor,
established the synonymy of both names and
selected the first as senior synonym. A detailed
redescription with a revised synonymy of this
species is in preparation by Jose Ochoa and
the first author.
SUBGENUS LEPTOSTERNUS MAURY, 1973
Brachistosternus (L. ) artigasi Cekalovic, 1974
figures 46, 58
Brachistosternus ( Leptosternus ) artigasi Cekalovic,
1974: 248-250; Ojanguren Affilastro, 2005a: 187.
Brachistosternus artigasi: Cekalovic and Artigas,
1981: 81; Cekalovic, 1983a: 54, 1983b: 188;
Ko varik, 1998: 100; Lowe and Fet, 2000: 49;
Ojanguren Affilastro, 2002b: 38, 2004b: 74,
2005a: 176, 178, 191.
Type Material: Holotype $ (MZUC
528), CHILE: Region IV (Coquimbo): Elqui
Province : La Serena, Lomas de Penuelas
[29°54'S 71°15'W], 5.vii.l968, T. Cekalovic.
New Records: CHILE: Region IV
(Coquimbo): Elqui Province : Las Cardas
[30°17'S 71°16'W], 1978, M. Moreno (pit-
falls), \$, 1? (AMNH). Limari Province :
Parque Nacional Fray Jorge, upslope from
campsite, 30°40'34.1"S 71°38'09.5"W,
4.xi.2003, L. Prendini, C. Mattoni, and J.
Ochoa, 167 m, UV detection, cold night, full
moon, Matorral with bare gravelly patches, 3
juv. (AMCC 159657), \$ (AMCC 159658).
Diagnosis: Brachistosternus artigasi is
closely related to B. chango. These are the
only species of the genus that possess two
macrosetae associated with the d and e
trichobothria of the pedipalp femur (fig. 8).
Both species display the typical trichobothrial
pattern of subgenus Leptosternus. Their hemi-
spermatophores are also similar: The distal
lamina is equal to or slightly shorter than the
basal portion, the internal spines are absent,
the basal spines and row of spines are well
developed, and the basal triangle is medium¬
sized. The VM carina of metasomal segment V
extends the entire length of the segment in
both species.
The two species may be separated from one
another as follows. Brachistosternus chango
exhibits larger androvestigia, almost twice the
size of those exhibited by B. artigasi (figs. 4,
46). The two species may also be distinguished
on the basis of pigmentation pattern.
Brachistosternus chango possesses paired
VSM and VL stripes on metasomal segments
I IV, whereas B. artigasi possesses a single
VM and paired VL stripes. Telson glands are
absent in B. chango, but well developed in B.
artigasi.
Brachistosternus artigasi occurs in sympatry
with B. cekalovici. However, the two species
may be separated by means of the pigmenta¬
tion pattern of metasomal segments I-IV,
which display paired VSM and VL stripes in
B. cekalovici, compared with a broad, faint
VM stripe and paired VL stripes in B. artigasi.
They may also be distinguished on the basis
of setation. Brachistosternus artigasi possesses
two macrosetae associated with the d and e
trichobothria of the pedipalp femur, whereas
B. cekalovici possesses only one.
Distribution: This species inhabits the
Pacific coastline, in Coquimbo Province, central
Chile (fig. 58). All localities where it has been
collected fall within the Matorral Estepario
botanical subregion, of the Matorral y Bosque
Esclerofilo botanical region (Gajardo, 1993).
Notes: See Ojanguren Affilastro (2005a)
for a redescription of this species.
2007 OJANGUREN AFFILASTRO ET AL.: CHILEAN BRACHISTOSTERNUS SCORPIONS
7
Brachistosternus ( L .) cekalovici Ojanguren
Affilastro, 2005
figures 45, 60
Brachistosternus (L.) cekalovici Ojanguren
Affilastro, 2005a: 175-192.
Type Material: Holotype $ (MACN-Ar
10243), CHILE: Region IV (Coquimbo):
Elqui Province : Tres Cruces [29°22'24"S
70°56'02"W], January 10, 1984, E. Maury.
New Records: CHILE: Region IV
(Coquimbo): Choapa Province : Playa
Amarilla, 31°51'36.03"S 71°30'31.55"W,
3.xi.2003, L. Prendini, C. Mattoni, and J.
Ochoa, 10 m, UV detection, cool, humid
night, cloud cover (fairly dark), light sea
breeze off Pacific, coastal Matorral, semi-
degraded coastal sand dunes on south-facing
side of rocky ridge, 1 2 , 1 juv. (CDA), 2 £, 1 ?
(AMNH), 22 juv. (AMCC 159659), 1 juv.
(AMCC 159660). Elqui Province : Punta
Teatinos, ca. 10 km N La Serena,
29°49'20.28"S 71°17'23.49" W, 7.xi.2003, L.
Prendini, C. Mattoni, J. Ochoa, J. Pizarro,
and P. Agusto, 0 m, UV detection, cool, still
night, full moon, high humidity near beach,
sand dunes behind beach, scattered grasses
and forbs, 2 juv. (CDA), 3c?, 5$, 6 juv.
(AMNH), 9 juv. (AMCC 159661).
Diagnosis: Brachistosternus cekalovici is
closely related to B. negrei. The two species
share the following combination of characters.
Carapace densely pigmented except for intero¬
cular surface. Tergites with two well-devel¬
oped spots laterally and a small spot medially.
Metasoma with paired VSM and VL stripes
on segments I-IV, and a single VM and paired
VL stripes on segment V. Telson glands
divided into two separate halves. Hemis-
permatophore with distal lamina shorter than
or equal to basal portion, without internal
spines, with the row of spines unbranched,
and with a medium-sized basal triangle.
Brachistosternus cekalovici may be distin¬
guished by the presence of a well-developed
VM carina on metasomal segment V (fig. 45),
which is absent in B. negrei.
The pigmentation pattern of B. chango is
similar to that of B. cekalovici. However, B.
chango may be distinguished by the presence
of two macrosetae associated with the d and e
trichobothria of the pedipalp femur; only
a single seta is present in B. cekalovici.
Brachistosternus cekalovici occurs in sym-
patry with B. artigasi. Both species may be
separated by the pigmentation pattern of
metasomal segments I IV, which possess
paired VSM and VL stripes in B. cekalovici,
compared to a broad, faint VM and paired VL
stripes in B. artigasi. The two species may be
further distinguished on the basis of setation.
Brachistosternus artigasi possesses two macro-
setae associated with the d and e trichobothria
of the pedipalp femur, whereas B. cekalovici
possesses only one.
Distribution: Previously, this species was
known only from the type locality. We provide
two new records in the present contribution.
This species occurs in the same habitat as B.
artigasi, near the Pacific coast, in Coquimbo
Province, central Chile (fig. 60). All localities
where this species has been collected fall
within the Matorral Estepario botanical sub-
region, of the Matorral y Bosque Esclerofilo
botanical region (Gajardo, 1993).
Notes: Specimens from Playa Amarilla
and Punta Teatinos differ morphologically
from the typical form. In the original de¬
scription, the male androvestigia were de¬
scribed as very small, occupying less than
25% of the dorsal surface of metasomal
segment V (Ojanguren Affilastro, 2005a).
However, the specimens from Playa Amarilla
exhibit larger androvestigia, in some cases
occupying ca. 50% of the dorsal surface. We
presently consider this variation to be in¬
traspecific, but specimens from more popula¬
tions must be collected to assess the range of
variation in the species. This is the first
bothriurid species in which such variation in
the size of the androvestigia has been ob¬
served.
Brachistosternus (L.) chango, new species
figures 1-8, 10-13, 18-21, 53, 61, table 1
Type Material: Holotype $ (AMNH),
CHILE: Region IV (Coquimbo): Choapa
Province : 3.5 km N turnoff to Salamanca,
5 km N Los Vilos, 31°52'13.6"S 71°27'27.1"W,
3.xi.2003, L. Prendini, C.I. Mattoni, and J.A.
Ochoa, 30 m, UV light detection on still, cool,
cloudy, humid, night, in coastal matorral with
cacti on steep, rocky, north-facing slope,
clayey-loam soil, syntopic with Caraboctonus
AMERICAN MUSEUM NOVITATES
NO. 3564
2007 OJANGUREN AFFILASTRO ET AL.: CHILEAN BRACHISTOSTERNUS SCORPIONS
9
keyserlingi Pocock, 1893. Paratypes: 3 S, 7 ?, 7
juv. (MACN-Ar 10930), Los Vilos, 30 km S
(near Pichidangui), 32°06'19.8"S 71°30'
25.1"W, 27.ii.2006, A. Ojanguren, L. Com-
pagnucci, C. Cuezzo, and P. Augusto, 27 m,
UV light, IS, 1? (MZUC), 2 juv. (AMCC
159687), same data.
Etymology: The specific epithet is a noun
in apposition referring to the Changos Indian
tribe that formerly inhabited the coastal
region from southern Peru to central Chile,
including the area where this new species was
collected.
Diagnosis: Brachistosternus chango is re¬
lated to B. artigasi and B. roigalsinai. It may
be distinguished from B. artigasi by the size of
the androvestigia. The androvestigia are very
narrow in B. artigasi but very broad, occupy¬
ing almost the entire dorsal surface of
metasomal segment V, in B. chango.
Brachistosternus chango may be distin¬
guished from B. roigalsinai according to the
setation on the dorsal surface of the pedipalp
femur. Brachistosternus chango possesses two
macrosetae associated with the d and e
trichobothria of the pedipalp femur, whereas
B. roigalsinai possesses only one. The two
species may be further separated by the
morphology of their hemispermatophores. In
B. roigalsinai, the distal lamina of the hemi-
spermatophore is almost straight, with a very
broad apex, the distal lobe is very well
developed, and the cylindrical apophysis is
very broad apically (see Ojanguren Affilastro,
2002b). In B. chango, the distal lamina is
slightly curved, the apex and distal lobe are
weakly developed, and the cylindrical apoph¬
ysis is almost the same width throughout its
length.
Description: The following description is
based on the holotype male (AMNH) and
paratypes (MACN, MZUC). Measurements
of the holotype male and a paratype female
(MACN-Ar 10930) are recorded in table 2.
Color. Base color: pale yellow, with a dusky
pattern (figs. 18-21). Chelicerae: unpigmen-
ted. Carapace: median ocular tubercle and
area around lateral ocelli infuscated; two
narrow, dark stripes extending from median
ocular tubercle, along anterior longitudinal
sulcus, to anterior carapace margin; two
broad, dark stripes extending from posterior
longitudinal sulcus to lateral ocelli; posterior
carapace margin with dark band on each side
poster olaterally; remainder unpigmented.
Tergites: I-VI each with dark band on either
side, joining medially; VII with pair of dark
bands posterolaterally, and some reticulate
pigmentation laterally, creating a pale unpig¬
mented stripe medially. Sternites: unpigmen¬
ted. Metasoma: segments I—III, dorsal sur¬
faces each with narrow median band in
anterior half of segment, and pair of dark
bands posterolaterally, lateral surfaces unpig¬
mented, ventral surfaces each with paired,
faint VSM and VL stripes, which may be
absent in some specimens; segment IV similar
to segments I—III, except dorsal bands faint
and ventral surface with paired, narrow VL
stripes and broad, faint VM stripe, in some
specimens partially divided into pair of VSM
stripes connected by reticulate pigmentation;
segment V dorsal and lateral surfaces unpig¬
mented, ventral surface with single narrow
VM and paired, broad VL stripes. Telson:
vesicle with pale reticulate pigmentation;
aculeus dark brown. Pedipalps: femur faintly
pigmented near articulation with patella;
patella faintly pigmented along anterior and
posterior margins; chela unpigmented. Legs:
femur pigmented near articulation with patel¬
la; patella pigmented near articulation with
femur; remaining segments unpigmented.
Chelicerae: Two subdistal teeth.
Figs. 1-9. Diagnostic characters of Chilean Brachistosternus Pocock, 1893. 1-8. Brachistosternus (L .)
chango, n.sp. 1. Sinistral hemispermatophore, internal aspect. 2. Sinistral hemispermatophore, external
aspect. 3. Sinistral hemispermatophore, detail of lobe region. 4. Metasomal segment V, male, dorsal aspect.
5. Metasomal segment V, male, ventral aspect. 6. Telson, male, lateral aspect. 7. Telson, female, lateral
aspect. 8. Dextral pedipalp femur, male, external aspect. 9. Brachistosternus ( L .) kamanchaca, n.sp., dextral
pedipalp femur, male, external aspect. Scale bars = 1 mm.
10
AMERICAN MUSEUM NOVITATES
NO. 3564
2007 OJANGUREN AFFILASTRO ET AL.: CHILEAN BRACHISTOSTERNUS SCORPIONS 11
Carapace : Anterior margin with weak me¬
dian projection. Anterior longitudinal sulcus
well developed. Ocular tubercle well devel¬
oped, situated anteromedially, interocular
sulcus weakly developed, ocelli one diameter
apart. Lateral, posterior longitudinal and
posteromarginal sulci well developed.
Carapacial surfaces smooth, except for poste¬
rior margin, which is slightly granular.
Tergites : Tergites I-VI, surfaces sparsely
granular (male) or smooth (female). Tergite
VII, surface slightly granular; two weakly
developed PM and PL carinae, evident only
in posterior half of segment.
Sternites : Median surfaces and posterior
margin, coarsely granular, remainder smooth
(male) or entirely smooth (female). Spiracles
narrow, medium-sized.
Pectines: Pectinal tooth count, male, 30-32
(N = 5; median = 31), female, 28-33 ( N = 8;
median = 29).
Metasoma: Segment I: dorsal, lateral, and
ventral surfaces slightly granular (males) or
smooth (female); DL and LSM carinae weakly
developed, extending entire length of segment;
LIM carinae present in posterior half of
segment only. Segments II and III similar to
segment I except LIM carina restricted to
posterior third of segment. Segment IV: dorsal
and lateral surfaces slightly granular; ventral
surface smooth, covered by scattered macro-
setae; DL and LSM carinae weakly developed,
extending entire length of segment; LIM
carinae absent. Segments I-IV each with pair
of dorsolateral macrosetae. Segment V: an-
drovestigia (male) very large, occupying al¬
most 75% of dorsal surface (fig. 4); dorsal and
lateral surfaces slightly granular; ventral sur¬
face densely granular; ventral macrosetae
arranged in three or four rows, the first with
four macrosetae, the rest with one or two
macrosetae; DL and LSM carinae weakly
developed, extending entire length of segment;
VM and VL carinae well developed, extending
entire length of segment (fig. 5).
Telson : Vesicle with rounded ventral sur¬
face. Vesicle surface slightly granular. No
telson gland evident. Aculeus slightly (male;
fig. 6) to strongly curved (females; fig. 7),
same length as vesicle.
Pedipalps: Femur DI, DE, and VI carinae
weakly developed; remaining surfaces smooth;
two macrosetae (Ml and M2) associated with
d and e trichobothria (fig. 8). Patella DI and
VI carinae weakly developed, represented only
by scattered granules; remaining surfaces
smooth. Chela narrow; surfaces smooth; in¬
ternal apophysis (male) well developed
(figs. 11, 12); VM carina weakly developed
(figs. 10, 11); fingers each with median row of
denticles and seven or eight pairs of accessory
denticles.
Trichobothria : Trichobothrial pattern typi¬
cal of subgenus Leptosternus: Neobothriotaxic
Major Type C, with one accessory trichobo-
thrium in V series of chela. Femur with 3
trichobothria (1 d, 1 i, and 1 e ). Patella with 19
trichobothria (3 V, 2 d, 1 i, 3 et, 1 est, 2 em, 2
esb, and 5 eb). Chela with 27 trichobothria (1
Est, 5 Et, 5 V, 1 Esb, 3 Eb, 1 Dt, 1 Db, 1 et, 1
est, 1 esb, 1 eb, 1 dt, 1 dst, 1 dsb, 1 db, 1 ib, 1 it)
(figs. 10-13).
Legs: Surfaces of all segments smooth.
Telotarsi not elongated; I and II each with
inner unguis ca. 5% shorter than external
(fig. 53).
Hemispermatophore: Distal lamina broad
and curved medially, similar in length to basal
portion (fig. 2). Cylindrical apophysis well
developed, longer than laminar apophysis
(fig. 1). Basal triangle well developed, com¬
prising three crests. Internal spines absent.
Basal spines and row of spines well developed
and aligned along same axis (fig. 3).
Figs. 10-17. Diagnostic characters of Chilean Brachistosternus Pocock, 1893. 10-13. Brachistosternus
(E) chango, n.sp. 10. Dextral pedipalp chela, female, ventral aspect. 11. Dextral pedipalp chela, male, ventral
aspect. 12. Dextral pedipalp chela, male, internal aspect. 13. Sinistral pedipalp chela, female, external aspect.
14-17. Brachistosternus (L .) kamanchaca, n.sp. 14. Dextral pedipalp chela, female, ventral aspect. 15. Dextral
pedipalp chela, male, ventral aspect. 16. Dextral pedipalp chela, male, internal aspect. 17. Sinistral pedipalp
chela, female, external aspect. Scale bars = 1 mm.
12
AMERICAN MUSEUM NOVITATES
NO. 3564
Figs. 18-21. Brachistosternus ( L .) chango, n.sp., habitus of male and female. 18. Dorsal aspect, male.
19. Ventral aspect, male. 20. Dorsal aspect, female. 21. Ventral aspect, female. Scale bars = 1 cm.
2007 OJANGUREN AFFILASTRO ET AL.: CHILEAN BRACHISTOSTERNUS SCORPIONS 13
Variation : Pedipalp chela length/height ra¬
tio, male, 3.3-3.41 (N = 5; mean = 3.37),
female, 3.04-3.26 (N = 8; mean = 3.13);
pedipalp chela length/width ratio, male, 4.01-
4.47 (N = 5; mean = 4.34), female, 4-4.26 (N
= 8; mean = 4.16); metasomal segment V,
length/width ratio, male, 1.75-1.83 (N = 5;
mean = 1.78), female, 1.71-1.79 (N = 8; mean
= 1.75); metasomal segment V, ventral macro-
setal count, 8-11 (N = 14; median = 10);
metasomal segment V, ventrolateral macro-
setal count, 8-10 (N = 14; median = 9);
telotarsus III, dorsal macrosetal count, 11-13
(N = 14; median = 12); telotarsus III,
ventrointernal macrosetal count, 8-10 (N =
14; median = 9); telotarsus III, ventroexternal
macrosetal count, 5-6 (N = 14; median = 5);
basitarsus III, dorsal macrosetal count, 6-7 (N
= 14; median = 7); total length, male, 41.5-50
(N = 5; mean = 46.3), female, 47-58 (N = 8;
mean = 51.43).
Distribution: Brachistosternus chango is
known from only two localities near Los
Vilos, close to the coast, in the Choapa
Province of the Coquimbo Region of Chile
(fig. 61). Both localities occur in the Matorral
Estepario botanical subregion, of the
Matorral y Bosque Esclerofilo botanical re¬
gion (Gajardo, 1993).
Ecology: The holotype male was collect¬
ed at an elevation of 30 m, by UV light
detection on a still, cool, cloudy, humid
evening (before midnight), in coastal
Matorral with cacti on a steep, rocky, north¬
facing slope. The specimen was sitting still in
the open on the surface of clayey-loam soil.
Several specimens of an iurid scorpion,
Caraboctonus keyserlingi Pocock, 1893, were
collected at the same locality but appeared to
prefer more densely vegetated areas with some
leaf litter. The paratypes were collected at
a nearby locality with a similar habitat
(fig. 22).
Notes: Several characters obscure the
phylogenetic relationship of B. chango to
other species of the genus. This is one of the
few species of Brachistosternus in which two
macrosetae (Ml and M2), associated with the
d and e trichobothria, occur on the pedipalp
femur. Only B. artigasi shares this character,
all other known species of Brachistosternus
possessing a single macroseta in this position
(fig. 9), suggesting that the additional macro-
seta might be synapomorphic for the two
species. The shape of the hemispermatophore
of B. chango also resembles that of B. artigasi,
providing a second potential synapomorphy.
However, the large size of the androvestigia
and shape of the VM carina of metasomal
segment V of B. chango are similar to those of
B. roigalsinai, providing potential synapomor-
phies with this species, which is related to
others from the plains of Chile and Peru
(Ochoa, 2002; Ojanguren Affilastro, 2005a).
Brachistosternus (L.) chilensis Kraepelin, 1911
figures 56, 62
Brachistosternus intermedius chilensis: Kraepelin,
1911: 87.
Brachistosternus ( Leptosternus ) chilensis: Maury,
1973: 249; Cekalovic, 1983a: 55; Masnu de
Moreno, 1991: 185.
Brachistosternus chilensis: Kovarik, 1998: 100;
Lowe and Fet, 2000: 49, 50; Ojanguren
Affilastro, 2005a: 191.
Type Material: Holotype ? (MZIT-Sc
64), CHILE: Region V (Valparaiso): Petorca
Province: La Ligua [32°27'07"S 71°13'41"W].
New Records: CHILE: Region V
(Valparaiso): Quillota Province : Parque
Nacional La Campana: Palmas de Ocoa,
Sendero Quillay trail from campsite,
32°56'02.53"S 71°04'33.43"W, 12.xi.2003, L.
Prendini, C. Mattoni, and J. Ochoa, 494 m,
UV detection, warm, still night, no moon, dry
savanna vegetation with palms, dense brush
on slopes, large boulders in places, dry leaf
litter layer, specimens collected along trail, on
roads and roadsides, and in degraded open
sandy flats, 8c?, 7?, 1 juv. (AMNH), 12 juv.
(AMCC 159662), 4$, 1?, 1 juv. (CDA); same
locality, 14.ii.2005, A. Ojanguren and P.
Korob, palm forest, 2$, 3?, 3 juv.
(MACN), 6 juv. (AMCC 159663).
Diagnosis: Brachistosternus chilensis is
most closely related to an undescribed species
(referred to in this paper as Brachistosternus
sp. 2). The hemispermatophore of these
species is similar: The distal lamina is shorter
than or equal to the basal portion, the internal
spines are absent, the basal spines well de¬
veloped, the row of spines unbranched, and
the basal triangle medium-sized. Both species
Figs. 22-23. Collection localities for Brachistosternus Pocock, 1893 in Chile. 22. Los Vilos, 30 km S (near
Pichidangui), habitat of Brachistosternus (L .) chango, n.sp. 23. Pan de Azucar National Park, Sector Las
Lomitas, habitat of Brachistosternus ( L .) kamanchaca, n.sp.
2007 OJANGUREN AFFILASTRO ET AL.: CHILEAN BRACHISTOSTERNUS SCORPIONS 15
also exhibit a similar pigmentation pattern on
the tergites, with two well-developed spots
laterally and a small spot medially that is
usually absent. Telson glands are absent in
both species.
Brachistosternus chilensis may be separated
from Brachistosternus sp. 2 as follows: the VM
stripe of metasomal segments I-IV is narrow
and does not join the VL stripes in B. chilensis
(fig. 56) whereas in Brachistosternus sp. 2 it is
broad and joins the VL stripes (fig. 55).
Brachistosternus chilensis may be further
distinguished by the presence of a well-de¬
veloped VM carina on metasomal segment V.
The VM carina of metasomal segment V is
absent in Brachistosternus sp. 2.
Brachistosternus chilensis is also related to
B. negrei, with which it shares a similar
hemispermatophore and pigmentation pattern
on the carapace and tergites. Both species may
be separated on the basis of the pigmentation
pattern of metasomal segments I-IV, which
display a single, narrow VM and paired VL
stripes in B. chilensis, compared with paired
VSM and VL stripes in B. negrei (fig. 54).
Distribution: Brachistosternus chilensis
occurs in Chile, in Region V “Valparaiso”
and in “Region Metropolitana, de Santiago”
(fig. 62). All localities where this species has
been collected fall into the Bosque Esclerofilo
botanical subregion, of the Matorral y Bosque
Xerofilo botanical region (Gajardo, 1993).
Notes: Kraepelin (1911) briefly described
this species, in a key, as a variety of B.
intermedins. No type material was designated,
but the description was based on a female
specimen from la Ligua, in Valparaiso Region,
Chile. Emilio Maury studied the scorpion
collection of the Museo ed Istituto di
Zoologia di Torino (MIZT) and found a scor¬
pion from the type locality labeled: “5. inter-
medius chilensis n. var. La Ligua; Aconcagua;
Chile (Sc. 64)”. Maury presumed that this was
the specimen described by Kraepelin (E.
Maury, unpublished data). Thereafter, Maury
implicitly accepted the validity of B. chilensis,
when mentioning it in a list of the species of
subgenus Leptosternus (Maury, 1973). We
studied this specimen at MIZT, as well as
several apparently conspecific specimens from
central Chile, and accept the validity of B.
chilensis based on this examination. A re¬
description of the species is in preparation by
Cristina Scioscia and the first author.
Brachistosternus (L.) donosoi Cekalovic, 1974
figures 48, 63
Brachistosternus ( Leptosternus ) donosoi Cekalovic,
1974: 250-252, 1983a: 55; Ojanguren Affilastro,
2004b: 74, 2005a: 183, 184.
Brachistosternus donosoi : Cekalovic and Artigas,
1981: 81; Cekalovic, 1983b: 188; Kovarik, 1998:
100; Ojanguren Affilastro, 2004b: 69, 2005a: 186,
191.
Type Material: Holotype $ (MZUC
530), CHILE: Region I (Tarapaca): Iquique
Province : Pampa del Tamarugal, 10 km E
Pica, 20°30'S 69°21'W, ii.1969, R. Donoso.
New Records: CHILE: Region I
(Tarapaca): Iquique Province : Pampa del
Tamarugal National Park, Salar de Pintados,
near ranger’s office and camp, 20°26'16.1"S
69°45'55.2"W, 1812005, C. Mattoni, A.
Ojanguren, and J. Ochoa, 1014 m, UV sam¬
pling in Prosopis tamarugo forest and UV
under salt plates, 1 $, 1 ?, 6 juv. (MACN),
\$, 1$ (CD A), \ $, 1?, 14 juv. (AMNH), 3
subad. ?, 7 juv. (AMCC 159688); Oficina
Humber stone, near Iquique, 20°12'36.7"S
69°47'29.8"W, 181.2005, C. Mattoni, A.
Ojanguren, and J. Ochoa, 1048 m, under
stones, 1?, 1 subad. $, 1 juv. $, 1 juv. ?
(AMCC 159689); La Tirana, 2 km W,
20°19'59.8"S 69°40'07,6"W, 191.2005, 999 m,
C. Mattoni, A. Ojanguren, and J. Ochoa, 2$,
3 juv. (AMNH), \$, 1?, 3 juv. (MACN), 1
juv. (CDA); road to Mamina, 20°15'18.1"S
69°41'36.7"W, 191.2005, C. Mattoni, A.
Ojanguren, and J. Ochoa, 1^,1$ (MACN),
1? (CDA), 1?, 4 juv. (AMNH).
Diagnosis: Brachistosternus donosoi is
most closely related to B. kamanchaca. These
are the only Chilean species in which the
internal spines of the hemispermatophore are
reduced to a small group of granules (fig. 30).
Both species also exhibit similar, medium¬
sized androvestigia situated medially on me¬
tasomal segment V (fig. 31).
The two species may be distinguished on
the basis of their pigmentation pattern.
Brachistosternus donosoi is usually unpigmen-
ted; the most pigmented specimens display
a pair of faint spots on the tergites laterally. In
16
AMERICAN MUSEUM NOVITATES
NO. 3564
contrast, the carapace and tergites are densely
pigmented, and the metasomal segments
display dark VL and VM stripes, at least on
segments III-V, in B. kamanchaca. The two
species also differ in the length/width ratio of
metasomal segment V. This ratio varies from
1.95 to 2.57 in B. donosoi and from 1.63 to 1.9
in B. kamanchaca. In addition, B. kamancha¬
ca. displays a flatter pedipalp chela. The
length/height ratio of the pedipalp chela
manus of B. kamanchaca. varies from 3.34 to
3.61 in males, and from 3.19 to 3.71 in females
whereas, in B. donosoi, it varies from 2.87 to
2.97 in males, and from 2.85 to 3.15 in
females.
Brachistosternus donosoi is also related to
two species from the plains of northern Chile,
B. mattonii and B. ochoai, with which it shares
the following characters. The hemispermato-
phore distal lamina is equal to the basal
portion in length, internal spines are present,
basal spines well developed, the row of spines
well developed and usually branched, and the
basal triangle well developed and strongly
sclerotized. The inner unguis of telotarsi I and
II is 15% longer than the external unguis. The
carapace, tergites, and sternites are densely
granular in the male.
Brachistosternus donosoi may be distin¬
guished from B. mattonii and B. ochoai as
follows. The androvestigia of B. donosoi are
medium-sized, compared to the elongated,
narrow androvestigia of the other species
(fig. 49). The internal spines of the hemisper-
matophore of B. donosoi are reduced to a small
group of granules, compared with those of the
other species, which are well developed
(fig. 40). The ventral surface of metasomal
segment IV is smooth in the male of B. donosoi
(fig. 48), but granular in males of the other
species (fig. 47).
Distribution: All localities where this
species has been collected fall within the
Desierto Absoluto botanical subregion
(fig. 63), of the Desierto botanical region
(Gajardo, 1993).
Ecology: This species inhabits forests of
Prosopis tamarugo, which is one of the few
trees adapted to the extremely xeric conditions
in the Atacama desert.
Notes: See Ojanguren Affilastro (2005a)
for a redescription of this species.
Brachistosternus ( L .) kamanchaca, new species
figures 9, 14-17, 24-35, 64, table 2
Type Material: CHILE: Region III
(Atacama): Chaharal Province : Holotype $
(MZUC), Pan de Azucar National Park,
Sector Las Lomitas, 26°00'32.6"S
70°36'26.2"W, 2412005, C.I. Mattoni and
A.A. Ojanguren Affilastro, 830 m, UV sam¬
pling, “lomas” formation, wet with fog,
syntopic with Bothriurus dumayi Cekalovic,
1974. Paratypes: 1 $, 1 $, 1 juv. (AMNH), 1 $,
1 subad. 2 (CDA), 2$, 1? (MACN-Ar
10931), 1?, 3 juv. (MZUC), 1 subad. 2, 5
juv. (AMCC 159664), same data as holotype.
Material Examined: CHILE: Region ITT
(Atacama): Copiapo Province : Copiapo, 10 km
S [27°27'30"S 70°22'30"W], xii.2001, P. Korob
and A.A. Ojanguren Affilastro, 12, 9 juv.
(MACN); Copiapo, 70 km S [27°57'S
70°33'W], 4.x. 1983, A. Roig Alsina, 3 juv.
(ARA); same data, except “E.A. Maury”, 1
juv. (MACN); Los Sapos, 2 km from turnoff
to Cerro Blanco from road (729) Vallenar-
Copiapo, 28°01'25.19"S 70°33'00.4"W, L.
Prendini, C. Mattoni, and J. Ochoa,
8.ix.2003, 532 m, UV detection, cool, still
night, full moon eclipsed, start of Atacama
desert, bare sandy ground with occasional
shrubs against low, rocky hills, becoming
harder upslope, specimens found on harder
ground, syntopic with B. dumayi, Bra¬
chistosternus sciosciae collected nearby on
sandy ground, 2$, 62, 7 juv. (AMNH), 2$,
22, 4 juv. (CDA), 34 juv. (AMCC 159665), 1
juv. (AMCC 159666). Huasco Province : La
Herradura, W of Parque Nacional Llanos de
Challe, 28°06'02.02"S 71°09'15.29"W, L.
Prendini, C. Mattoni, and J. Ochoa,
10.xi.2003, 35 m, UV detection on cool, dark,
breezy night, very humid near beach, coastal
sand flats, rocky hill and soils of intermediate
hardness between, arid chaparral with cacti
and other succulents, specimens on rocky
ground, syntopic with Bothriurus dumayi,
Brachistosternus roigalsinai, and Carabo-
ctonus keyserlingi, B. sciosciae collected near¬
by on sandy ground, 12 (AMNH), 3 2,3 juv.
(AMCC 159667); Llanos de Challe National
Park, plains on road to Cerro Negro,
28°11'16.4"S 71°06'06.7"W, 303 m, UV, full
moon, 25.L2005, C. Mattoni, A. Ojanguren.
12 (CDA), \$ (MACN), \$, 2 juv.
2007 OJANGUREN AFFILASTRO ET AL.: CHILEAN BRACHISTOSTERNUS SCORPIONS 17
(AMNH), 2 juv. (MZUC). Domeyko, 18 km
S, on road to observatorio La Silla [29°07'30"S
70°56'W], 3.x.1983, E.A. Maury, \$, 9 juv.
(MACN).
Etymology: The specific epithet is a noun
in apposition referring to the sea fog occurring
along the arid Pacific coastline of central and
northern South America, known locally as the
“Kamanchaca” or “Camanchaca”. The fog
exerts a strong influence on the environment
and biota of the coastal slopes by increasing
their humidity (Rundel et al., 1990).
Diagnosis: Brachistosternus kamanchaca
is closely related to B. donosoi. The two
species are readily distinguished by means of
pigmentation. Brachistosternus donosoi is usu¬
ally unpigmented or lightly pigmented, where¬
as B. kamanchaca is darkly pigmented. There
are also differences in the length/width ratio of
metasomal segment V. In B. donosoi , this ratio
varies from 1.95 to 2.57 whereas in B.
kamanchaca , it varies from 1.63 to 1.9. In
addition, B. kamanchaca displays a flatter
pedipalp chela. The length/height ratio of the
pedipalp chela manus of B. kamanchaca varies
from 3.34 to 3.61 in males and from 3.19 to
3.71 in females, whereas in B. donosoi, it varies
from 2.87 to 2.97 in males and from 2.85 to
3.15 in females.
Brachistosternus kamanchaca is also closely
related to B. artigasi and B. cekalovici. It may
be distinguished from both species on the basis
of the VM carinae of metasomal segment V.
Compared to B. artigasi and B. cekalovici, in
which the VM carinae are well developed and
extend the entire length of the segment, the
VM carinae of B. kamanchaca are weakly
developed, and usually restricted to the
posterior half of the segment. Brachisto¬
sternus kamanchaca may also be distinguished
from these species by the presence of internal
spines, represented by a small group of
granules, on the hemispermatophore, which
are absent in B. artigasi and B. cekalovici.
Brachistosternus kamanchaca may be further
distinguished from B. artigasi according to the
setation on the dorsal surface of the pedipalp
femur. Brachistosternus kamanchaca possesses
only one macroseta associated with the i and e
trichobothria of the pedipalp femur, whereas
B. artigasi possesses two. Brachistosternus
kamanchaca may be further distinguished
from B. cekalovici according to the setation
on the ventral surface of metasomal segment
V. Brachistosternus kamanchaca possesses 5-7
rows of macro setae on the ventral surface of
segment V, whereas B. cekalovici possesses
only 3-4 rows of macrosetae.
Description: The following description is
based on the holotype (MZUC) and paratypes
(AMNH, CDA, and MACN) from Pan de
Azucar National Park. Measurements of the
holotype male (MZUC) and a paratype female
(MACN-Ar 10931) are recorded in table 2.
Color. Base color: pale yellow, with a dusky
pattern (figs. 24-27). Chelicerae: anterior mar¬
gin and movable finger with reticulate pig¬
mentation. Carapace: median ocular tubercle
and area around lateral ocelli infuscated;
inter ocular surface unpigmented; remainder
densely pigmented. Tergites: I-VI each with
dark stripe in anterior two-thirds (in some
specimens, stripe divided into three dark
spots, usually connected by reticulate pigmen¬
tation); VII with dark triangle anteromedially,
and two dark bands posterolaterally.
Sternites: unpigmented. Metasoma: segments
I and II, dorsal surfaces each with narrow
band medially and pair of dark bands poster¬
olaterally, lateral surfaces unpigmented, ven¬
tral surfaces each with single broad VM and
paired narrow VL stripes (absent or faint in
most specimens, well developed only in very
pigmented specimens) joining at posterior
margin of segment; segment III similar to
segments I and II, except VM stripe usually
present; segment IV similar to segment III,
except VM stripe always present and well
developed; segment V, dorsal surface with
faint reticulate pigmentation near anterior and
posterior articulations, remainder unpigmen¬
ted, lateral surfaces unpigmented, ventral
surface with paired broad VL stripes and
single well-developed, narrow VM stripe,
joining in posterior third of segment. Telson:
vesicle ventral surface with pale reticulate
pigmentation; aculeus dark brown.
Pedipalps: femur and patella pigmented on
anterior and posterior margins, and near
articulations; chela with pale reticulate pig¬
mentation on external surface of manus and
near articulation of movable finger, fingers
unpigmented. Legs: femur pigmented near
articulation with patella; patella pigmented
18
AMERICAN MUSEUM NOVITATES
NO. 3564
Figs. 24-27. Brachistosternus (L .) kamanchaca, n.sp., habitus of male and female. 24. Dorsal aspect,
male. 25. Ventral aspect, male. 26. Dorsal aspect, female. 27. Ventral aspect, female. Scale bars = 1 cm.
2007 OJANGUREN AFFILASTRO ET AL.: CHILEAN BRACHISTOSTERNUS SCORPIONS 19
on anterior margin and near articulations;
remaining segments unpigmented.
Chelicerae : Two subdistal teeth.
Carapace : Anterior margin with weak me¬
dian projection. Anterior longitudinal sulcus
well developed. Ocular tubercle well devel¬
oped, situated anteromedially, interocular
sulcus weakly developed, ocelli almost three
diameters apart. Lateral, posterior longitudi¬
nal and posteromarginal sulci well developed.
Carapacial surfaces slightly granular.
Tergites : Tergites I VI, surfaces entirely
smooth, or smooth in anterior two-thirds,
and slightly granular in posterior third.
Tergite VII, surface slightly granular in
posterior half; two PL carinae evident.
Sternites: Surfaces granular (male) or
smooth (female). Spiracles narrow, medium¬
sized.
Pectines : Pectinal tooth count, male, 28-35
(V = 10; median = 30), female, 24-31 (V =
10; median = 28).
Metasoma : Segments I—III: dorsal and
lateral surfaces slightly granular; ventral sur¬
faces granular (male) or smooth (female); DL
and LSM carinae weakly developed, extending
entire length of segment; LIM carinae weakly
developed, present in posterior third of
segment only. Segment IV: ventral surface
smooth, covered by scattered macrosetae; DL
carinae weakly developed, extending entire
length of segment; LIM carinae absent; LSM
carinae evident at anterior and posterior
margins of segment only. Segments I-IV each
with pair of dorsolateral macrosetae. Segment
V: androvestigia medium-sized, situated sub-
medially (fig. 31); dorsal and lateral surfaces
slightly granular (male) or smooth (female);
ventral surface sparsely granular in posterior
two-thirds of segment (fig. 32); ventral macro¬
setae usually comprising five to seven rows,
the first two or three rows with four to six
macrosetae, the rest with one to three macro¬
setae; DL carinae weakly developed (male) or
absent (female); VL carinae well developed,
extending entire length of segment; VM carina
weakly developed, present in posterior two-
thirds of segment only.
Telson : Vesicle with rounded ventral sur¬
face. Vesicle surface sparsely granular. Telson
gland well developed, triangular, partially
divided at posterior apex (fig. 35) or, rarely,
completely divided. Aculeus slightly curved,
same length as vesicle (figs. 33, 34).
Pedipalps : Femur DI, DE, and VI carinae
absent or weakly developed; remaining sur¬
faces smooth; one macroseta (Ml) associated
with d and e trichobothria (fig. 9). Patella DI
and VI carinae absent or weakly developed;
remaining surfaces smooth. Chela: narrow;
surfaces smooth; internal apophysis (male)
well developed (figs. 15, 16); VM carina
weakly developed; fingers each with median
row of denticles and four to six pairs of
accessory denticles.
Trichobothria : Trichobothrial pattern typi¬
cal of subgenus Leptosternus: Neobothriotaxic
Major Type C, with one accessory trichobo-
thrium in V series of chela. Femur with 3
trichobothria (1 d, 1 i, and 1 e ). Patella with 19
trichobothria (3 V, 2 d, 1 i, 3 et, 1 est, 2 em, 2
esb, and 5 eb). Chela with 27 trichobothria (1
Est, 5 Et, 5 V, 1 Esb, 3 Eb, 1 Dt, 1 Db, 1 et, 1
est, 1 esb, 1 eb, 1 dt, 1 dst, 1 dsb, 1 db, 1 ib, 1 it’,
figs. 14-17).
Legs: Surfaces of femur and patella smooth.
Telotarsi very elongated; I and II each with
inner unguis ca. 10% shorter than external.
Hemispermatophore : Distal lamina broad,
slightly curved, equal to or slightly shorter
than basal portion (figs. 28, 29). Cylindrical
apophysis well developed, longer than laminar
apophysis. Basal triangle well developed,
comprising two or three crests. Internal spines
reduced to small group of tiny granules. Basal
spines and row of spines well developed and
aligned along same axis (fig. 30).
Geographic variation : Different populations
of B. kamanchaca display some variation in
pigmentation and granulation. This is common
among Brachistosternus species with broad
distributional ranges, and we do not consider
this variation diagnostic at the species level.
Additionally, the following meristic variation
is recorded in B. kamanchaca : pedipalp chela
length/height ratio, male, 3.34-3.61 (N = 10;
mean = 3.49), female, 3.19-3.71 (N = 8; mean
= 3.38); pedipalp chela length/width ratio,
male, 4.12-5.03 (V = 10; mean = 4.59),
female, 4.07-5 (N = 8; mean = 4.42);
metasomal segment V, length/width ratio,
male, 1.7-1.9 (N = 10; mean = 1.79), female,
1.63-1.86 ( N = 8; mean = 1.73); metasomal
segment V, ventral macrosetal count, 13-23 (N
20
AMERICAN MUSEUM NOVITATES
NO. 3564
Figs. 28-35. Diagnostic characters of Brachistosternus (L .) kamanchaca, n.sp. 28. Sinistral hemisperma-
tophore, internal aspect. 29. Sinistral hemispermatophore, external aspect. 30. Sinistral hemispermatophore,
detail of lobe region. 31. Metasomal segment V, male, dorsal aspect. 32. Metasomal segment V, male, ventral
aspect. 33. Telson, female, lateral aspect. 34. Telson, male, lateral aspect. 35. Telson, male, dorsal aspect.
Scale bars = 1 mm.
2007 OJANGUREN AFFILASTRO ET AL.: CHILEAN BRACHISTOSTERNUS SCORPIONS 21
= 14; median = 15); metasomal segment V,
ventrolateral macrosetal count, 8-11 (N = 12;
median = 10); telotarsus III, dorsal macrosetal
count, 9-12 (N = 15; median = 10); telotarsus
III, ventrointernal macrosetal count, 6-8 (N =
15; median = 6); telotarsus III, ventroexternal
macrosetal count, 4-5 (N = 15; median = 5);
basitarsus III, dorsal macrosetal count, 6-7 (N
= 15; median = 6); total length, male, 34-49
(N = 9; mean = 41.5), female, 39^15 (N = 7;
mean = 41.35).
Distribution: Brachistosternus kamancha¬
ca inhabits the Atacama Region of northern
Chile, from 0 to 700 m. Some specimens have
been collected in coastal localities, but others
have been collected as far as 100 km inland
(fig. 64). All localities where this species has
been collected fall within the Desierto Costero
and Desierto Florido botanical subregions of
the Desierto botanical region (Gajardo, 1993).
Ecology: At the type locality in Pan de
Azucar National Park and another locality,
La Herradura, B. kamanchaca was only
collected on high, seaward-facing slopes
(fig. 23). These slopes have a Lomas
(Spanish for small hills) environment
(Rundel et al., 1990), with a microclimate that
is markedly more humid than the surrounding
desert, because it is under the influence of the
sea fog (Kamanchaca). This species was not
found in other habitats in Pan de Azucar
National Park. Outside the park, where the
species was collected in habitats not associated
with Lomas, it always occupied areas with
some (sparse) vegetation, mostly small shrubs,
loose soil, and only a few rocks.
Brachistosternus kamanchaca was syntopic
with the bothriurid Bothriurus dumayi at Pan
de Azucar, La Herradura, and Los Sapos. At
La Herradura, it was also syntopic with B.
roigalsinai and the iurid Caraboctonus key-
serlingi. Brachistosternus sciosciae was also
collected at La Herradura and Los Sapos, but
this species was restricted to sandier substrata
than B. kamanchaca, and the two species
appear to be allotopic.
Notes: Some specimens of B. kamanchaca
have been collected near Copiapo, the type
locality of B. castroi (fig. 59), raising the
possibility that the new species described here
might be a junior synonym of the latter. We
examined the holotype of B. castroi, a poorly
preserved juvenile female with few diagnostic
characters, and observed differences in the
granulation and carination of metasomal
segment V, in the number of macrosetae on
metasomal segment V and telotarsus III, and
in the pigmentation pattern, prompting our
decision to regard both as distinct species. The
type material of B. castroi closely resembles B.
intermedius, which has in the past created
confusion regarding its identity (Ringuelet,
1953; Abalos, 1963; Lowe and Fet, 2000).
More material must be collected in the vicinity
of Copiapo to clearly establish the identity of
B. castroi.
Brachistosternus (L. ) kovariki Ojanguren
Affilastro, 2003
figure 65
Brachistosternus (L .) kovariki Ojanguren Affilastro,
2003b: 23-36; Ochoa, 2004: 139.
Type Material: Holotype $ (MACN-Ar
10347), CHILE: Region II (Antofagasta): El
Loa Province : San Pedro de Atacama, 30 km
E, 22°56'09"S 67°56'02"W, 3600 m, 30.L2003,
A. Ojanguren Affilastro and P. Korob.
New Records: CHILE, Region III
(Atacama): El Loa Province : San Pedro de
Atacama, 27 km E, 22°54'49.2"S 67°55'
35.5"W, 25.i.2005, C. Mattoni and A. Ojang¬
uren, 3650 m, UV on rocky plains, 3 juv.
(MACN), \$, n (CDA), \$, 1$, 1 juv.
(AMNH), 2 juv. $, 3 juv. ?, 5 juv. (AMCC
159668).
Diagnosis: Brachistosternus kovariki is
parapatric with B. prendinii, both species
inhabiting the same region at different alti¬
tudes. Brachistosternus prendinii occurs from
2800 to 3200 m, and B. kovariki from 3200 to
3600 m. The two species may be distinguished
by the setation of metasomal segment V.
Brachistosternus prendinii possesses 19-26
ventral setae, whereas B. kovariki possesses
8-13 macrosetae. The two species may be
further distinguished on the basis of the VM
carina of metasomal segment V, which extends
the entire length of the segment in B. kovariki,
but is restricted to the second half of the
segment in B. prendinii.
Brachistosternus kovariki is closely related to
B. piacentinii. Both species exhibit a similar
pigmentation pattern: tergites with paired spots
22
AMERICAN MUSEUM NOVITATES
NO. 3564
TABLE 2
Measurements (mm) of Brachistosternus ( Leptosternus ) kamanchaca , n.sp. (Holotype and Paratype) and
Brachistosternus ( Leptosternus ) chango, n.sp. (Holotype and Paratype)
B. ( L .) kamanchaca
B. (L.)
chango
Holotype $
MZUC
Paratype ?
MACN
Holotype $
AMNH
Paratype 2
MACN
Total length
43.00
37.28
44.92
54.59
Carapace, length
5.17
4.44
5.25
6.46
Carapace, anterior width
3.64
3.15
3.88
4.60
Carapace, posterior width
5.58
4.53
5.82
6.87
Mesosoma, total length
12.71
13.86
10.59
15.80
Metasoma, total length
19.79
14.78
22.86
25.30
Metasomal segment I, length
3.23
2.42
4.00
3.80
Metasomal segment I, width
3.31
2.51
3.40
4.44
Metasomal segment I, height
2.75
2.02
2.80
3.55
Metasomal segment II, length
3.64
2.66
4.16
4.44
Metasomal segment II, width
3.07
2.18
3.24
4.04
Metasomal segment II, height
2.75
2.02
2.80
3.55
Metasomal segment III, length
3.79
2.83
4.24
4.69
Metasomal segment III, width
2.91
2.10
3.08
3.88
Metasomal segment III, height
2.59
1.94
2.80
3.39
Metasomal segment IV, length
4.28
3.15
4.80
5.66
Metasomal segment IV, width
2.75
2.00
2.93
3.64
Metasomal segment IV, height
2.51
1.78
2.68
3.07
Metasomal segment V, length
4.85
3.72
5.66
6.71
Metasomal segment V, width
2.83
2.02
3.08
3.39
Metasomal segment V, height
2.26
1.54
2.58
2.83
Telson, length
5.33
4.20
6.22
7.03
Vesicle, length
3.07
2.26
3.72
3.64
Vesicle, width
2.10
1.59
2.08
2.34
Vesicle, height
1.86
1.37
1.84
2.26
Aculeus, length
2.26
1.94
2.50
3.39
Femur, length
4.20
3.07
3.60
3.72
Femur, width
1.13
0.93
1.04
1.29
Patella, length
3.79
3.07
3.60
3.80
Patella, width
1.45
1.21
1.48
1.65
Chela, length
7.19
5.17
6.14
6.78
Chela, width
1.78
1.13
1.53
1.70
Chela, height
2.18
1.45
1.86
2.18
Movable finger, length
4.12
2.83
3.72
4.04
laterally and a single spot medially, usually
connected by reticulate pigmentation; metaso-
mal segments I—III with a single VM and paired
VL stripes, densely pigmented but not joining
at the posterior margin of the segments;
metasomal segment IV with a single, broad
VM and paired, narrow VL stripes, joining in
the posterior third of the segment; metasomal
segment V with a narrow VM and paired,
broad VL stripes, joining in the posterior third
of the segment. The hemispermatophore of
both species is also similar: Internal spines are
absent; basal spines are well developed; the row
of spines is well developed but unbranched; the
basal triangle is medium-sized. However, they
differ in the shape of the distal lamina, which is
narrow and equal to the basal portion in length
in B. kovariki, but broad and shorter than the
basal portion in B. piacentinii. Both species may
be further separated by means of the dimen¬
sions of metasomal segment V. In B. kovariki,
the length/width ratio of metasomal segment V
2007 OJANGUREN AFFILASTRO ET AL.: CHILEAN BRACHISTOSTERNUS SCORPIONS 23
varies from 1.8 to 1.95 whereas, in B. piacenti-
nii, it varies from 1.7 to 1.8. Finally, the VM
carina of metasomal segment V extends the
length of the segment in B. kovariki but is
restricted to the posterior half in B. piacentinii.
Distribution : Brachistosternus kovariki has
only been collected in a small area of western
Antofagasta (fig. 65), between 3200 and 3600 m
(Ojanguren Affilastro, 2003b). All localities
where this species has been collected fall within
the Altiplano y Puna botanical subregion of
the Estepa Alto-Andina botanical region (Ga-
jardo, 1993). At the same latitude in neighbor¬
ing Argentina, this species is replaced by B.
intermedins in similar habitats and altitudes
(Ojanguren Affilastro, 2004a).
Brachistosternus (L.) mattonii Ojanguren
Affilastro, 2005
figures 49, 66
Brachistosternus (L.) mattonii Ojanguren Affilastro,
2005a: 175-192.
Type Material: Holotype $ (MACN-Ar
10235), CHILE: Region II (Antofagasta):
Antofagasta Province : Hornitos [22°75'S
70°18'W], 2.x.1983, E. Maury (MACN-Ar
10235).
Diagnosis: Brachistosternus mattonii is
closely related to B. ochoai. Both species are
characterized by hemispermatophores with
well-developed internal and basal spines,
a well-developed, usually branched row of
spines, and a strongly sclerotized basal tri¬
angle (fig. 40); elongated, narrow androvesti-
gia (fig. 49); absence of the telson gland; inner
unguis of telotarsi I and II 15% longer than
external unguis; carapace, tergites, sternites,
and metasomal segment IV densely granular
in males (fig. 47); DI, VI, and DE carinae of
pedipalp femur, and DI and VI carinae of
pedipalp patella very well developed and
extending the entire length of the segment.
The two species may be distinguished as
follows. Brachistosternus mattonii is almost
completely unpigmented, the most pigmented
specimens displaying only very faint reticulate
pigmentation around the ocular tubercle and
along the lateral margins of the tergites. In
constrast, B. ochoai is densely pigmented on
the carapace, tergites, and metasomal segment
V, the ventral surface of which displays a single
VM and paired VL stripes, whereas the
dorsal surface displays an antero-median
and two posterolateral spots. Both species
may also be distinguished according to the
dimensions of the pedipalp chela. The length/
height ratio varies from 2.9 to 3.11 in the
males of B. mattonii and is 3.13 in the female,
whereas it varies from 3.25 to 3.6 in the males
of B. ochoai and from 3.02 to 3.21 in the
females.
Distribution: This species inhabits coast¬
al areas of northern Chile, in the Iquique and
Antofagasta regions (fig. 66). All localities
where it has been collected fall within the
Desierto Costero de Tocopilla, in the Desierto
costero botanical subregion of the Desierto
botanical Region (Gajardo, 1993).
Brachistosternus (L.) negrei Cekalovic, 1975
figures 54, 67
Brachistosternus ( Leptosternus ) negrei Cekalovic,
1975: 69-72. Cekalovic and Artigas, 1981: 81;
Cekalovic, 1983a: 56, 1983b: 188. Ojanguren
Affilastro, 2005a: 188.
Brachistosternus negrei : Kovarik, 1998: 101; Lowe
and Fet, 2000: 51. Ojanguren Affilastro, 2005a:
189-191.
Type Material: Holotype $ (MZUC 546,
lost), CHILE: Region VII (Maule): Talca
Province : Talca, 22 miles N [35°17'S
71°38'W], 22.xii.1950, Ross and
Michelbacher.
New Records: CHILE: Region VII
(Maule): Cauquenes Province : Reserva
Nacional Los Ruiles, NW Cauquenes,
35°49'30"S 72°30'30"W, 13.xi.2003, L.
Prendini, C. Mattoni, and J. Ochoa, 146 m,
UV detection on cool, still, dark, humid night
in Nothofagus forest on steep west-facing
slope, small river at base of slope, specimens
taken on open sandy ground near river and
around campsite, 5 juv. (CDA), 10 juv.
(AMCC 159669). Talca Province : Parque
Nacional Altos del Lircay [35°38'S 71°10'W],
16.ii.2005, A. Ojanguren and P. Korob,
1500 m, sub-Andean forest, 3$, 2?, 2 juv.
(MACN), 2 juv. (AMCC 159671). Region
Metropolitana de Santiago: Melipilla
Province : El Membrillo [34°00'S 71°06'W],
15.xii.2004, J.S. Ascher and A. Kawahara, UV
light detection at night, 1$, 3?, 14 juv.
(AMCC 159670).
24 AMERICAN MUSEUM NOVITATES NO. 3564
Figs. 36^3. Diagnostic characters of Chilean Brachistosternus Pocock, 1893. 36, 42. Brachistosternus
(B.) ehrenbergii (Gervais, 1841). 36. Sinistral hemispermatophore, internal aspect. 42. Sinistral pedipalp
2007 OJANGUREN AFFILASTRO ET AL.: CHILEAN BRACHISTOSTERNUS SCORPIONS 25
Diagnosis: Brachistosternus negrei is
closely related to B. cekalovici, with which it
shares the following combination of charac¬
ters: carapace densely pigmented except for
interocular surface, tergites with two well-
developed spots laterally and a small spot
medially, metasomal segments I-IV with
paired VSM and VL stripes, metasomal
segment V with a single VM and paired VL
stripes (fig. 54); telson glands divided into two
separate halves; hemispermatophore with dis¬
tal lamina shorter than or equal to basal
portion, internal spines absent, basal spines
well developed, with row of spines un¬
branched, and basal triangle medium-sized.
The two species may be separated by the
presence, in B. cekalovici, of a well-developed
VM carina on metasomal segment V. The VM
carina is absent in B. negrei.
Brachistosternus negrei is also related to B.
chilensis. The two species share similar hemi-
spermatophores and pigmentation pattern,
except for the ventral surface of the metaso¬
ma, which displays paired VSM and VL
stripes on segments I-IV in B. negrei
(fig. 54), compared with a single VM and
paired VL stripes in B. chilensis (fig. 56).
Brachistosternus chilensis may be further
distinguished by the presence of a well-de¬
veloped VM carina on metasomal segment V.
The VM carina is absent in B. negrei.
Distribution: Brachistosternus ( L .) negrei
is the southernmost species of Brachistosternus
in Chile. Previously, this species was recorded
only from the Maule and Bio-Bio Regions. In
the present contribution, we provide records
from the southern part of Region
Metropolitana de Santiago (fig. 67). All local¬
ities where this species has been collected fall
within the Matorral y Bosque Esclerofilo
and Bosque Caducifolio botanical regions
(Gajardo, 1993).
Brachistosternus (L.) ochoai Ojanguren
Affilastro, 2004
figures 40, 47, 68
Brachistosternus ( L .) ochoai Ojanguren Affilastro,
2004b: 69-74.
Type Material: Holotype $ (MACN-Ar
10517), CHILE: Region III (Atacama):
Chaharal Province : Pan de Azucar National
Park [26°09'S 70°39'W], ii.2003, A. Ojanguren
Affilastro and P. Korob.
New Records: CHILE: Region III
(Atacama): Chaharal Province : Parque
Nacional Pan de Azucar, near Chanaral:
Caleta Pan de Azucar (dunes behind beach
camp), 26°09'04.55"S 70°40'00.36"W,
9.xi.2003, C. Mattoni, L. Prendini, and J.
Ochoa, 20 m, UV detection, cool, still night,
moon completely obscured by clouds, coastal
flats and shrub-coppice dunes 300 m from
beach, specimens in open ground, 1 juv.
(CDA), 3 juv. (AMCC 159672); Quebrada
Pan de Azucar, 8 km from Pan de Azucar,
26°06'47.49" S 70°34'08.42"W, 9.xi.2003, L.
Prendini, C. Mattoni, and J. Ochoa, 229 m,
UV detection on cool, still night, moon partly
obscured by clouds, Atacama desert with steep
scree slopes and alluvial flats at the base,
specimen on rocks at base of slope, 1 ?
(AMCC 159673).
Diagnosis: Brachistosternus ochoai is most
closely related to B. turpuq from southern
Peru (Ochoa, 2002, 2005). Both species share
the following combination of characters. The
hemispermatophores possess well-developed
internal and basal spines, a very well-de-
patela, ventral aspect. 37. Brachistosternus {L.) roigalsinai Ojanguren Affilastro, 2002. Sinistral
hemispermatophore, internal aspect. 38. Brachistosternus (L .) sciosciae Ojanguren Affilastro, 2002.
Sinistral hemispermatophore, internal aspect. 39. Brachistosternus (L .) sp. 1. Sinistral hemispermatophore,
internal aspect. 40. Brachistosternus ( L .) ochoai Ojanguren Affilastro, 2004. Sinistral hemispermatophore,
internal aspect, detail of lobe region. 41. Brachistosternus (L .) sp. 3. Sinistral hemispermatophore, internal
aspect, detail of lobe region. 43. Brachistosternus ( L .) perettii Ojanguren Affilastro and Mattoni, 2006.
Sinistral pedipalp patela, ventral aspect. Scale bars = 1 mm. Abbreviations: L = distal lamina; Bp = basal
portion; lr = lobe region; dl= distal lobe; ca = cylindrical apophysis; la = laminar apophysis; bt = basal
triangle; is = internal spines; rs = row of spines; bs: basal spines.
Figs. 44-49. Diagnostic characters of Chilean Brachistosternus Pocock, 1893. 44 . Brachistosternus (L.)
sp. 2. Metasomal segment V, ventral aspect. 45 . Brachistosternus ( L .) cekalovici Ojanguren Affilastro, 2005.
Metasomal segment V, ventral aspect. 46 . Brachistosternus ( L .) artigasi Cekalovic, 1974. Metasomal segment
Y, male, dorsal aspect. 47 . Brachistosternus ( L .) ochoai Ojanguren Affilastro, 2004. Metasomal segment IV,
male, ventral aspect. 48 . Brachistosternus ( L .) donosoi Cekalovic, 1974. Metasomal segment IV, male, ventral
aspect. 49 . Brachistosternus ( L .) mattonii Ojanguren Affilastro, 2005. Metasomal segment V, male, dorsal
aspect. Scale bars = 1 mm.
2007 OJANGUREN AFFILASTRO ET AL.: CHILEAN BRACHISTOSTERNUS SCORPIONS 27
veloped, usually branched row of spines, and
a strongly sclerotized basal triangle (fig. 40).
The androvestigia are narrow and elongated.
The carapace, tergites, sternites, and metaso¬
ma, including segment V (which are smooth in
most species), are densely granular in males
(fig. 47). The inner unguis of telotarsi I and II
is 15% longer than the external unguis. The
telson gland is absent. The DI, VI, and DE
carinae of the pedipalp femur and DI and VI
carinae of the pedipalp patella are well
developed and extend the entire length of the
segment.
The two species may be separated as
follows. The metasoma of B. turpuq is
completely unpigmented, compared to that
of B. ochoai, in which a triangular spot is
evident at the anterior margin of the dorsal
surface of each segment (restricted to segment
V in some specimens), and a single VM and
paired VL stripes are evident on metasomal
segment V. Both species may also be separated
according to the dimensions of metasomal
segment V. The length/width ratio varies from
1.8 to 1.9 in B. turpuq and from 1.92 to 2.03 in
B. ochoai.
Brachistosternus ochoai is also related to B.
mattonii, from which it may be distinguished
on the basis of pigmentation. In B. ochoai, the
carapace and tergites are densely pigmented,
and metasomal segment V displays a single
VM and paired VL stripes whereas in B.
mattonii, the metasoma and, in most speci¬
mens, also the carapace and tergites are
unpigmented, the most strongly pigmented
specimens displaying only faint reticulate
pigmentation in the central area of the
carapace and near the lateral margins of the
tergites. Both species may be further distin¬
guished by the dimensions of the pedipalp
chela. The length/height ratio of the chela of
B. ochoai varies from 3.25 to 3.6 in males, and
from 3.02 to 3.21 in females whereas in B.
mattonii, it varies from 2.9 to 3.11 in males
and is 3.13 in the female.
Distribution: This species inhabits plains
along the Pacific coastline, in the Atacama
Region of northern Chile (fig. 68). All local¬
ities where it has been collected fall within
the Desierto Costero botanical subregion,
of the Desierto botanical region (Gajardo,
1993).
Brachistosternus ( L .) perettii Ojanguren
Affilastro and Mattoni, 2006
figures 43, 50, 69
Brachistosternus (L .) perettii Ojanguren Affilastro
and Mattoni, 2006: 79-84.
Type Material: Holotype $ (MZUC-
UCCC 28634), CHILE: Region IV
(Coquimbo): Elqui Province : Pastos Largos,
“El Indio” Gold Mine, 3600 m [29°49'S
70°03'W], pitfall, i.1993, H. Vasquez C.
New Records: CHILE: Region IV
(Coquimbo): Elqui Province : Quebrada el
Negro, Mina El Indio [29°47'S 70°00'W],
6.xi.2003, C. Mattoni, L. Prendini, J.
Pizarro, and J. Ochoa, 3800 m, UV detection
at night, alpine vegetation with low bushes
and grass tufts on hard, rocky ground with
bare patches and stones in places, very cold
and windy, full moon, specimens in open areas
and under vegetation, 1 ?, 1 juv. (AMNH), 1
juv. (CDA), 5 juv. (AMCC 159674); Cancha
de Sky (Sky Camp), Mina El Indio, 3300 m
[29°51'S 70°03'W], 6.xi.2003, C. Mattoni, L.
Prendini, J. Pizarro, and J. Ochoa, UV
detection, 1 ? (AMNH), 1 juv. (CDA); Pista
de Aterrizaje (Aeropuerto), airport landing
strip, Mina El Indio [29°51'S 70°03'W], ca.
4000 m, 6.xi.2003, C. Mattoni, L. Prendini, J.
Pizarro, and J. Ochoa, UV detection, 1?
(AMNH), 1 juv. (CDA); road to Mina el
Indio, 3 km from entrance [29°53'30"S
70°03'W], 26.ii.2004, C. Mattoni, J. Ochoa,
and J. Pizarro, 2850 m, 3 juv. (AMCC
159675). Elqui Province : Paso del Agua
Negra, between Juntas and international
border, 30°16'14.5"S 69°58'27.9"W, 27.L2005,
C. Mattoni and A. Ojanguren, 3295 m, UV on
“vega”, extremely wet, \ $, 1? (MACN-Ar),
1 $ (CDA), 1 $ (AMNH), 1 $ (AMCC
159676).
Diagnosis: Brachistosternus perettii is
a typical Andean species of the genus
(Ojanguren Affilastro, 2003b). It is medium¬
sized and densely pigmented and exhibits
the trichobothrial pattern of subgenus
Leptosternus (fig. 43). The tergites display
a broad longitudinal stripe, and metasomal
segments I-V display a single VM and paired
VL stripes that join in all segments. The distal
lamina of the hemispermatophore is similar to
the basal portion in length and slightly curved
28
AMERICAN MUSEUM NOVITATES
NO. 3564
Figs. 50-56. Diagnostic characters of Chilean Brachistosternus Pocock, 1893. 50. Brachistosternus (L. )
perettii Ojanguren Affilastro and Mattoni, 2006. Telson, male, dorsal aspect. 51. Brachistosternus ( L .) sp. 3.
Telson, male, dorsal aspect. 52. Brachistosternus ( L .) sp. 1. Telotarsus I, dextral leg, internal aspect. 53.
Brachistosternus ( L .) chango, n.sp. Telotarsus I, dextral leg, internal aspect. 54. Brachistosternus (L. ) negrei
Cekalovic, 1975. Metasoma, pigmentation pattern, ventral aspect. 55. Brachistosternus ( L .) sp. 2. Metasoma,
pigmentation pattern, ventral aspect. 56. Brachistosternus ( L .) chilensis Kraepelin, 1911. Metasoma,
pigmentation pattern, ventral aspect. Scale bars = 1 mm.
medially, the internal spines are absent, the
basal spines, row of spines, and basal triangle
are well developed. The male androvestigia are
medium-sized.
Brachistosternus perettii may be distin¬
guished from other Andean species of the
genus by means of the telson gland, which is
completely divided into two separate halves
(fig. 50), compared with other Andean species
in which it is undivided (fig. 51).
Brachistosternus perettii is sympatric with
an undescribed species of Brachistosternus,
2007 OJANGUREN AFFILASTRO ET AL.: CHILEAN BRA CHISTOSTERNUS SCORPIONS 29
Figs. 57-59. The known distributions of Brachistosternus Pocock, 1893 in central-northern Chile
(contour interval 500 m). 57. Brachistosternus ehrenbergii (Gervais, 1841). 58. Brachistosternus artigasi
Cekalovic, 1974. 59. Brachistosternus castroi Mello-Leitao, 1941.
Figs. 60-62. The known distributions of Brachistosternus Pocock, 1893 in central-northern Chile
(contour interval 500 m). 60. Brachistosternus cekaloviki Ojanguren Affilastro, 2005. 61. Brachistosternus
chango, n.sp. 62. Brachistosternus chilensis Kraepelin, 1911.
2007 OJANGUREN AFFILASTRO ET AL.: CHILEAN BRACHISTOSTERNUS SCORPIONS 31
referred to here as Brachistosternus sp. 3. The
two species may be separated by the presence,
in Brachistosternus sp. 3, of a complete VM
carina on metasomal segment V, which is
absent in B. perettii.
Distribution: This species occurs at alti¬
tudes between 3200 and 3600 m in the central
Andes and the Dona Ana Mountain range in
the western part of the Coquimbo Region,
Chile (fig. 69). These localities occur within
the Andes Mediterraneos botanical subregion
of the Desierto Florido botanical region
(Gajardo, 1993).
Brachistosternus (L. ) piacentinii Ojanguren
Affilastro, 2003
figure 70
Brachistosternus ( L .) piacentinii Ojanguren
Affilastro, 2003b: 23-36; Ochoa, 2004: 139.
Type Material: Holotype $ (MACN-Ar
10349), BOLIVIA: Oruro Department: Sajama
Province : Sajama National Park, near
Huanacota lake, 18°02'53"S 68°56'08"W,
4500 m, 23-24.xi.2001, A. Ojanguren
Affilastro and P. Korob.
New Records: CHILE: Region I
(Tarapaca): Parinacota Province : Parinacota,
Lauca National Park, 18°12'01.3"S
69°16'01.4"W, 16.i.2005, C. Mattoni, A.
Ojanguren, and J. Ochoa, 4431 m, under
stones, \$, 1?, 12 juv. (AMNH), \$, 1?, 3
juv. (CDA), 2 juv. $, 1 juv. ?, 5 juv. (AMCC
159677).
Diagnosis: Brachistosternus piacentinii is
closely related to B. kovariki. The two species
display a similar pigmentation pattern, in
which the tergites display one spot medially
and two spots laterally, usually connected by
reticulate pigmentation, metasomal segments
I—III display a single VM and paired VL
stripes, densely pigmented but not joining at
the posterior margin of the segments, metaso¬
mal segment IV exhibits a broad VM and
paired, narrow VL stripes, joining in the
posterior third of the segment, metasomal
segment V displays a narrow VM and paired,
broad VL stripes, joining in the posterior
third of the segment. The hemispermatophore
of both species is also similar: The internal
spines are absent, the basal spines well de¬
veloped, the row of spines well developed but
unbranched, and the basal triangle medium¬
sized.
The two species differ in the shape of the
distal lamina, which is narrow and equal to
the basal portion in length in B. kovariki, but
broad and shorter than the basal portion in B.
piacentinii. Both species may be further
separated according to the length/width ratio
of metasomal segment V, which varies from
1.8 to 1.95 in B. kovariki and from 1.7 to 1.8 in
B. piacentinii. The VM carina of metasomal
segment V extends the entire length of the
segment in B. kovariki, but is restricted to the
posterior half in B. piacentinii.
Brachistosternus piacentinii is also closely
related to B. titicaca from central Bolivia and
southern Peru (Ochoa, 2002; Ojanguren
Affilastro, 2003b). The VM carina of metaso¬
mal segment V extends the entire length of the
segment in both species. Brachistosternus
piacentinii and B. titicaca also possess similar
hemispermatophores. However, in B. titicaca,
the distal lamina is slightly curved medially
and almost equal to the basal portion in
length, compared with that of B. piacentinii,
which is almost straight and shorter than the
basal portion. The two species display a similar
pigmentation pattern except that, in B. titica¬
ca, the VM stripe of the metasoma joins with
the VL stripes in metasomal segment V only,
whereas in B. piacentinii, it joins in metasomal
segments IV and V. Both species may be
further separated according to the length/
width ratio of metasomal segment V, which
varies from 1.5 to 1.65 in B. titicaca and from
1.7 to 1.8 in B. piacentinii.
Distribution: Brachistosternus piacentinii
occurs at altitudes between 4300 and 4500 m,
in the Tarapaca Region of northern Chile,
and in Sajama Province of eastern Bolivia
(fig. 70). This is the only Brachistosternus
species recorded on both sides of the Andes.
At Sajama National Park (Bolivia), B. pia¬
centinii is sympatric with B. galianoae
(Ojanguren Affilastro, 2002c), but the latter
is absent in Chile. Both species occur at
the maximum altitudes recorded for
Brachistosternus. All localities where B. pia¬
centinii has been collected fall within the
Altiplano y Puna botanical subregion of the
Estepa Alto-Andina botanical Region
(Gajardo, 1993).
Figs. 63-65. The known distributions of Brachistosternus Pocock, 1893 in central-northern Chile
(contour interval 500 m). 63. Brachistosternus donosoi Cekalovic, 1974. 64. Brachistosternus kamanchaca,
n.sp. 65. Brachistosternus kovariki Ojanguren Affilastro, 2003.
2007 OJANGUREN AFFILASTRO ET AL.: CHILEAN BRACHISTOSTERNUS SCORPIONS 33
Brachistosternus (L.) prendinii Ojanguren
Affilastro, 2003
figure 71
Brachistosternus ( L .) prendinii Ojanguren
Affilastro, 2003b: 24-27; Ochoa, 2004: 139.
Type Material: Holotype $ (MACN-Ar
10345), CHILE: Region II (Antofagasta): El
Loa Province : San Pedro de Atacama, 20 km
E [22°56'S 68°0rW], 2800 m, 3112003, A.
Ojanguren Affilastro and P. Korob.
New Records: CHILE: Region III
(Atacama): El Loa Province : San Pedro de
Atacama, ~20 km E, 22°54'38.8"S
67°59'44.2"W, 2512005, C. Mattoni and A.
Ojanguren, 3117 m, UV on rocky plains with
some shrubs, 1$ (AMCC 159678).
Diagnosis: Brachistosternus prendinii may
be distinguished from most other species of
Brachistosternus by the large number of
macrosetae on the ventral surface of metaso-
mal segment V (19-26). Most other
Brachistosternus species possess fewer (<17)
macrosetae on this surface. Only B. perettii
possesses a similar number of macrosetae (18-
22), but the telson gland of this species is
divided into two separate halves (fig. 50),
which is not the case in B. prendinii.
Furthermore, B. prendinii possesses a VM
carina in the posterior half of metasomal
segment V. The VM carina of metasomal
segment V is absent in B. perettii.
Brachistosternus prendinii is most closely
related to B. quiscapata. Both species display
a similar pigmentation pattern in which the
carapace is almost completely pigmented, the
tergites display a broad transverse stripe, and
the metasomal segments a single VM and
paired VL stripes, joining in the posterior
margin of all segments. Brachistosternus pre¬
ndinii and B. quiscapata possess similar hemi-
spermatophores in which the distal lamina is
of moderate length, equal to the basal portion,
and slightly curved, internal spines are absent,
the row of spines and basal spines well
developed, and the basal triangle medium¬
sized. The two species also exhibit well-
developed telson glands, and their androves-
tigia are medium-sized and similar, albeit
slightly more elongated in B. prendinii than
in B. quiscapata. Besides differences in setation
on the ventral surface of metasomal segment
V, the two species may be separated by means
of the VM carina of metasomal segment V,
which extends the entire length of the segment
in B. quiscapata, but is restricted to the
posterior half in B. prendinii.
Distribution: Brachistosternus prendinii
occurs at altitudes between 2800 and 3200 m,
in the western part of the Antofagasta Region
(Ojanguren Affilastro, 2003b). At higher
altitudes, it is replaced by B. kovariki. All
localities where this species has been collected
(fig. 71) occur in the Altiplano y Puna
botanical subregion of the estepa Alto-
Andina botanical region (Gajardo, 1993).
Brachistosternus (L.) quiscapata Ochoa and
Acosta, 2002
figure 72
Brachistosternus ( L .) quiscapata Ochoa and Acosta,
2002: 1-13; Ochoa, 2002: 55, 2004: 139; 2005: 54,
57. Ojanguren Affilastro, 2003b: 25, 34.
Brachistosternus quiscapata : Ojanguren Affilastro,
2003b: 24, 35, 2005a: 191; Ochoa, 2005: 61.
Type Material: Holotype $ (MUSM),
PERU: Tacna Department: mountains around
Tarata, 3370 m, 17°28'S 70°01'W, 2012000,
J. Ochoa.
New Records: CHILE: Region I
(Tarapaca): Parinacota Province : Putre, 6 km
W, 18°13'26.8"S 69°32'47.T'W, 1612005, C.
Mattoni, A. Ojanguren, and J. Ochoa,
3732 m, UV and under stones, 1 $, 1 $, 2
juv. (AMNH), 1 juv. (CDA), 1?, 1 juv. $
(AMCC 159679).
Diagnosis: Brachistosternus quiscapata is
most closely related to an undescribed species
of Brachistosternus from the central Andes of
Chile, referred to here as Brachistosternus sp.
3. Both species display a similar pigmentation
pattern, in which the carapace is almost
completely pigmented, the tergites display
a broad transverse stripe, and the metasomal
segments a single VM and paired VL stripes,
joining at the posterior margin of all segments.
Both species possess similar hemispermato-
phores, in which the distal lamina is slightly
curved, of moderate length, and equal to the
basal portion, the internal spines are absent,
the basal spines and row of spines well
developed, and the basal triangle medium¬
sized. Both species also exhibit a well-de-
Figs. 66-68. The known distributions of Brachistosternus Pocock, 1893 in central-northern Chile
(contour interval 500 m). 66. Brachistosternus mattonii Ojanguren Affilastro, 2005. 67. Brachistosternus
negrei Cekalovic, 1975. 68. Brachistosternus ochoai Ojanguren Affilastro, 2004.
2007 OJANGUREN AFFILASTRO ET AL.: CHILEAN BRACHISTOSTERNUS SCORPIONS 35
veloped telson gland and similar, medium¬
sized androvestigia. The VM carina of meta-
somal segment V extends the entire length of
the segment in both species. Brachistosternus
sp. 3 may be separated from B. quiscapata by
means of the narrower pedipalp chela of the
male, the length/width ratio of which varies
from 4.8 to 5.09, compared with B. quiscapata ,
in which it varies from 3.7 to 3.9.
Brachistosternus quiscapata is also related to
B. montanus, from the central Andes of
Argentina. Both species possess similar hemi-
spermatophores, androvestigia, and telson
glands, and the VM carina extends the full
length of metasomal segment V. The pigmen¬
tation pattern of the two species is similar, but
B. quiscapata may be distinguished because
the VM stripe joins the VL stripes in all
metasomal segments whereas in B. montanus,
the VM and VL stripes join in segments IV
and V only.
Distribution: This species occurs on the
Andean slopes of the Tarapaca Region,
northern Chile, and the Tacna Department,
southern Peru, between 3000 and 3400 m
(fig. 72). The localities where it has been
collected occur in the Serrania Esteparia
ecoregion (Brack, 1986) and the Estepa Alto-
Andina botanical region (Gajardo, 1993).
Brachistosternus (L.) roigalsinai Ojanguren
Affilastro, 2002
figures 37, 73
Brachistosternus (L .) roigalsinai Ojanguren
Affilastro, 2002b: 37^46; Ojanguren Affilastro,
2004b: 70, 72, 74, 2005a: 183.
Brachistosternus roigalsinai : Ojanguren Affilastro,
2005a: 191.
Type Material: Holotype $ (MACN-Ar
10239), CHILE: Region IV (Coquimbo): Elqui
Province". Llano de La Higuera, 29°30'S
71°17'W, 11.x.1983, L. Pena.
New Records: CHILE: Region III
(Atacama): Chaharal Province : Parque
Nacional Pan de Azucar, near Chanaral:
Agua Salada, in Quebrada Pan de Azucar,
26°08'23.2"S 70°37'48.11"W, 9.xi.2003, L.
Prendini, C. Mattoni, and J. Ochoa, 82 m,
UV detection on cool, still night, moon
completely obscured by clouds, Atacama
desert with steep scree slopes and alluvial flats
at the base, specimen at base of rocky slope,
1 ? (AMCC 159681). Parque Nacional Pan de
Azucar: Mirador, 26°06'53.4"S 70°38'21.9"W,
23.L2005, UV, A. Ojanguren and C. Mattoni,
262 m, 1 juv. (CDA), 1 juv. (MACN), 2 juv.
(AMNH); Parque Nacional Pan de Azucar:
Sector Las Lomitas, 26°00'32.6"S
70°36'26.2"W, 833 m, “lomas” formation,
wet, with fog, 24.L2005, C. Mattoni and A.
Ojanguren, 2 juv. (AMNH), 1 juv. (CDA).
Huasco Province: La Herradura, W of Parque
Nacional Llanos de Challe, 28°06'02"S
71°09'15.5"W, 10.xi.2003, L. Prendini, C.
Mattoni, and J. Ochoa, 35 m, UV detection
on cool, dark, breezy night, very humid near
beach, coastal sand flats, rocky hill and soils
of intermediate hardness between, arid mator-
ral with cacti and other succulents, 5 juv.
(AMCC 159682); Parque Nacional Llanos de
Challe, 5.5 km from Administration building,
28°06'56.13"S 71°05'55.41"W, 10.xi.2003, L.
Prendini, C. Mattoni, and J. Ochoa, 65 m, UV
detection on cool, breezy night, moon not yet
risen, steep scree slope with cacti and bushes,
specimen on sandy loam ground at base of
slope, 1 juv. (AMCC 159683), 1 juv. (CDA);
Parque Nacional Llanos de Challe, Cerro
Negro, 28°44'16.4"S 71°06'07"W, 25.L2005,
UV, Ojanguren and Mattoni, 2 juv.
(MACN); Parque Nacional Llanos de
Challe, plains on road to Cerro Negro,
28°11'16.4"S 71°06'06.7"W, 303 m, UV, full
moon, 25.L2005, C. Mattoni, A. Ojanguren,
1$, 1 juv. (AMNH), 1? (CDA). Parque
Nacional Llanos de Challe, Administration
building, 28°09'39.8"S 71°03'20"W, 25.L2005,
A. Ojanguren and C. Mattoni, \$, 1 juv.
(MACN). Region IV (Coquimbo): Elqui
Province : Punta Teatinos, ca. 10 km N La
Serena, 29°49'20.28"S 71°17'23.49"W,
7.xi.2003, J. Ochoa, C. Mattoni, and L.
Prendini, 0 m, UV detection, cool, still night,
full moon, high humidity near beach, rocky
hill near beach, comprising granite boulders
with sandy loam soil, Matorral vegetation, 1 ?
(AMCC 159684), 1 juv. (AMCC 159680).
Diagnosis: Brachistosternus roigalsinai is
most closely related to B. ehrenbergii. The VM
carina extends the full length of metasomal
segment V and is more strongly developed in
both species than in other Chilean species of
Brachistosternus. Both species possess similar
Figs. 69-71. The known distributions of Brachistosternus Pocock, 1893 in central-northern Chile
(contour interval 500 m). 69. Brachistosternus perettii Ojanguren Affilastro and Mattoni, 2006.
70. Brachistosternus piacentinii Ojanguren Affilastro, 2003. 71. Brachistosternus prendinii Ojanguren
Affilastro, 2003.
2007 OJANGUREN AFFILASTRO ET AL.: CHILEAN BRACHISTOSTERNUS SCORPIONS 37
hemispermatophores, the distal lamina of
which is almost straight (figs. 36, 37), but
narrow medially and lobate apically, the basal
triangle is very elongated (more developed in
B. ehrenbergii ), the internal spines absent, and
the basal spines and row of spines (usually
branched), well developed. Both species also
display a very well-developed distal lobe, but
this is more elongated in B. roigalsinai than in
B. ehrenbergii. The cylindrical apophysis is
more strongly developed in the two species than
in other species of the genus but is dorsoven-
trally flattened in B. ehrenbergii (fig. 36) and
cylindrical in B. roigalsinai (fig. 37).
Brachistosternus roigalsinai and B. ehren¬
bergii may be separated according to the
number of trichobothria on the ventral surface
of the pedipalp patella: B. ehrenbergii pos¬
sesses five to seven trichobothria, whereas B.
roigalsinai possesses only three. The two
species may be further distinguished according
to the size of the androvestigia, which are very
large, occupying almost the entire dorsal
surface of metasomal segment V in B.
roigalsinai, compared with B. ehrenbergii, in
which they are medium-sized.
Brachistosternus roigalsinai is also related to
B. chango. Both species display the largest
androvestigia in the genus, occupying almost
the entire surface of metasomal segment V
(fig. 4). The two species may be distinguished
according to the setation of the pedipalp
femur: B. roigalsinai possesses a single macro-
seta associated with the d and e trichobothria
of the pedipalp femur, whereas B. chango
possesses two macrosetae (fig. 8). Both species
may also be distinguished by means of the
pigmentation pattern of the metasoma: B.
chango displays paired VSM and VL stripes
on the ventral surface of metasomal segments
I-IV, but these segments are unpigmented or
display a faint VM and paired VL stripes on
metasomal segments III and IV only, in B.
roigalsinai.
Brachistosternus roigalsinai may be distin¬
guished from all other Brachistosternus by the
very elongated distal lobe of the hemisperma-
tophore (fig. 37), which is much longer than
the distal lobe of the hemispermatophore of
other species (figs. 1, 28, 38, 39).
Distribution: This species, which inhabits
the plains of southern Antofagasta, Atacama,
and northern Coquimbo regions (fig. 73), is
one of the most widely distributed
Brachistosternus species in Chile (Ojanguren
Affilastro, 2002b, 2005a). All localities where
it has been collected occur in the Desierto
Costero botanical subregion of the Desierto
botanical region.
Brachistosternus (L.) sciosciae Ojanguren
Affilastro, 2002
figures 38, 74
Brachistosternus (L .) sciosciae Ojanguren Affilastro,
2002b: 37^16, 2004b: 72, 2005a: 183.
Brachistosternus sciosciae: Ojanguren Affilastro,
2005a: 191.
Type Material: Holotype $ (MACN-Ar
10237), CHILE: Region III (Atacama):
Copiapo Province : Caldera [27°04'S
70°17'W], 4.x.1983, E. Maury.
New Records: CHILE: Region III
(Atacama): Copiapo Province : Los Sapos,
2 km from turnoff to Cerro Blanco from road
Vallenar-Copiapo (Km 729 on Route 5),
28°01'25.19"S 70°33'06.36"W, 8.xi.2003, L.
Prendini, C. Mattoni, and J. Ochoa, 532 m,
UV detection, cool, still night, full moon
eclipsed, start of Atacama desert, bare sandy
ground with occasional shrubs against low,
rocky hills, becoming harder upslope, speci¬
mens on soft sand against hills, syntopic with
Bothriurus dumayi, B. kamanchaca collected
nearby on harder ground, 5 ? (CDA), 1 $,
18? (AMNH), 21 juv. (AMCC 159686).
Huasco Province : La Herradura, W of
Parque Nacional Llanos de Challe,
28°06'02.02"S 71°09'15.29"W, 10.xi.2003, L.
Prendini, C. Mattoni, and J. Ochoa, 35 m, UV
detection on cool, dark, breezy night, very
humid near beach, coastal sand flats, rocky
hill and soils of intermediate hardness be¬
tween, arid Matorral with cacti and other
succulents, specimens on soft sandy ground,
Bothriurus dumayi, B. kamanchaca, B. roigal¬
sinai, and Caraboctonus keyserlingi collected
nearby on harder, rocky ground, 16?, 18 juv.
(CDA), 15?, 1 juv. (AMNH), 82juv. (AMCC
159685).
Diagnosis: Brachistosternus sciosciae may
be distinguished from all other Brachis¬
tosternus species by the absence of androves¬
tigia, which are present in all other species of
Figs. 12-1 A. The known distributions of Brachistosternus Pocock, 1893 in central-northern Chile
(contour interval 500 m). 72. Brachistosternus quiscapata Ochoa and Acosta, 2002. 73. Brachistosternus
roigalsinai Ojanguren Affilastro, 2002. 74. Brachistosternus sciosciae Ojanguren Affilastro, 2002.
2007 OJANGUREN AFFILASTRO ET AL.: CHILEAN BRACHISTOSTERNUS SCORPIONS 39
the genus. Brachistosternus sciosciae is most
closely related to an undescribed species
from central Chile (referred to here as
Brachistosternus sp. 1). The two species may
be separated from other Brachistosternus
species by the weak development of the
internal structures of their hemispermato-
phores: The internal spines, basal spines, and
row of spines are absent, and the basal
triangle vestigial, that is, reduced to a small,
smooth bulge without crests or spines, in
both species. The two species are small in size
(on average less than 40 mm in total length)
and usually almost completely unpigmented.
The inner unguis of telotarsi I and II is
almost 15% longer than the external unguis,
and the internal pedal spur of telotarsi I and
II is vestigial or absent. The pedipalps and
metasoma of both species exhibit poorly
developed carinae, and the telson is slightly
compressed dorsoventrally, especially in
males.
The two species may be separated from one
another by the presence, in Brachistosternus
sp. 1, of small androvestigia, which are absent
in B. sciosciae. The two species may be further
distinguished as follows: The distal lamina of
the hemispermatophore is curved medially in
Brachistosternus sp. 1 (fig. 39) but almost
straight, and slightly curved distally, in B.
sciosciae (fig. 38); the VM carina of metaso-
mal segment V extends the entire length of the
segment in Brachistosternus sp. 1, but is
restricted to the second half of the segment
in B. sciosciae.
Distribution: This species inhabits the
Atacama Region of northern Chile (fig. 74).
All localities where it has been collected occur
in the Desierto Costero botanical subregion of
the Desierto Botanical region (Gajardo, 1993).
Although this species was previously known
only from coastal localities (Ojanguren
Affilastro, 2002b, 2005a), a population was
recently discovered in sympatry with B.
kamanchaca, n.sp. at Los Sapos, almost
100 km inland.
Notes: The specimens from Los Sapos
are remarkable in possessing a brownish color¬
ation with faint dark spots, quite different from
“typical” specimens, which are completely un¬
pigmented, with a pale yellow color. This
difference in pigmentation is presumably related
to differences in the color of the substratum
inhabited by these scorpions, which is slightly
darker at Los Sapos than at the coastal localities
where this species was collected previously.
Brachistosternus (L.) sp. 1
figures 39, 52, 75
Material Examined: CHILE: Region IV
(Coquimbo): Elqui Province : 5 km from Punta
Choros, near the road to Choros, 29°14'29.0"S
71°25'45.9"W, 22.ii.2006, A.A. Ojanguren
Affilastro, L. Compagnucci, and A.C.
Cuezzo, 12 m, UV sampling, in dunes, synto-
pic with B. roigalsinai, 25$, 5?, 4 juv.
(MACN-Ar); Choros, 29°17'10.3"S
71°19'7.2"W, 22.ii.2006, A.A. Ojanguren
Affilastro, L. Compagnucci, and A.C.
Cuezzo, UV sampling, in dunes with some
shrubs, syntopic with B. roigalsinai, and B.
coriaceus, 1 $ (MACN-Ar).
Diagnosis: Brachistosternus sp. 1 is closely
related to B. sciosciae. Both species are small
in size (less than 40 mm on average) and
almost completely unpigmented. The inner
unguis of telotarsi I and II is approximately
15% longer than the external unguis, and the
internal pedal spur of telotarsi I and II is
vestigial or absent in both species (fig. 52).
The pedipalpal and metasomal carinae are
weakly developed in these species, and the
telson slightly compressed dorsoventrally (es¬
pecially in males). Furthermore, the internal
spines, basal spines, and row of spines of
the hemispermatophore are absent, and the
basal triangle vestigial, reduced to a small
smooth bulge without crests or spines, in both
species.
The two species may be distinguished by the
presence, in Brachistosternus sp. 1, of small
androvestigia, which are absent in B. sciosciae.
Both species may also be distinguished ac¬
cording to the following characters. In
Brachistosternus sp. 1, the distal lamina of
the hemispermatophore is curved medially
(fig. 39), whereas in B. sciosciae, it is almost
straight and only slightly curved distally
(fig. 38). The VM carina of metasomal seg¬
ment V extends the entire length of the
segment in Brachistosternus sp. 1, but is
restricted to the second half of the segment
in B. sciosciae.
Figs. 75-77. The known distributions of Brachistosternus Pocock, 1893 in central-northern Chile
(contour interval 500 m). 75. Brachistosternus sp. 1. 76. Brachistosternus sp. 2. 77. Brachistosternus sp. 3.
2007 OJANGUREN AFFILASTRO ET AL.: CHILEAN BRACHISTOSTERNUS SCORPIONS 41
Distribution: Brachistosternus sp. 1 has
only been collected in coastal dunes near
Punta Choros in Coquimbo region, Chile
(fig. 75). These localities fall within the
Desierto Costero del Huasco botanical sub-
region of the Desierto botanical region
(Gajardo, 1993).
Notes: A description of this species is in
preparation by Jaime Pizarro, Pablo Agusto,
and the first two authors.
Brachistosternus (L.) sp. 2
figures 44, 55, 76
Material Examined: CHILE: Region V
(Valparaiso): Los Andes Province : between
Rio Blanco and Juncal [32°54'S 70°11'W],
6.L1984, A. Roig Alsina, 1950 m, 1 $
(MACN-Ar 10767); Guardia Vieja [32°54'S
70°17'W], 1911984, E. Maury, 1600 m, 1?,
1 juv. (MACN-Ar 10768); Juncal [32°52'S
70°10'W], 5.i. 1984, E. Maury, 1950 m, 3?,
3 juv. (MACN-Ar 10769); 20 km from
Portezuelo [32°53'S 70°13'W], 13.ii.2005,
Ojanguren and Korob, 2100 m, 1 juv.
(MACN-Ar 10770).
Diagnosis: Brachistosternus sp. 2 is closely
related to B. chilensis. Both species possess
similar hemispermatophores, in which the
distal lamina is shorter than or equal to the
basal portion, the internal spines are absent,
the basal spines well developed, the row of
spines unbranched, and the basal triangle
medium-sized. The pigmentation pattern of
the tergites is also similar, displaying two well-
developed spots laterally and a small spot
medially that is often absent. Telson glands
are absent in both species.
The two species may be separated by means
of pigmentation pattern. In B. chilensis, the
VM stripe on metasomal segments I-IV is
narrow and does not join the VL stripes
(fig. 56), whereas in Brachistosternus sp. 2, it is
broad and joins the VL stripes (fig. 55).
Additionally, B. chilensis exhibits a well-de¬
veloped VM carina on metasomal segment V,
which is absent in Brachistosternus sp. 2
(fig. 44).
Distribution: Brachistosternus sp. 2 is the
southernmost Andean species in Chile. It has
been collected at intermediate altitudes (1600-
2100 m) in the central Andes of the Valparaiso
region (fig. 76). These localities belong to the
Andes Mediterraneos botanical subregion, of
the Estepa altoandina botanical region
(Gajardo, 1993).
Notes: A description of this species is in
preparation by the first author and Cristina
Scioscia.
Brachistosternus (L.) sp. 3
figures 41, 51, 77
Material Examined: CHILE: Region IV
(Coquimbo): Elqui Province : between Juntas
and Paso del Agua Negra [30°14'S 70°02'W],
6.iii.2006, A.A. Ojanguren Affilastro, L.
Compagnucci, and A.C. Cuezzo, 3200 m,
UV sampling, syntopic with B. perettii and
an undescribed species of Orobothriurus, 5 $
(MACN-Ar); 2 km to entrance to “El Indio”
gold mine [29°54'S 70°03'W], 26.ii.2004,
2850 m, UV sampling, C.I. Mattoni, J.A.
Ochoa, and J. Pizarro Araya, 1 juv. (CDA);
Cancha Sky, “El Indio” gold mine [29°5LS
70°03'W], ii. 1992, Vasquez, 3300 m, under
stones, 1 $ (LEULS); Sancarron, “El Indio”
gold mine [29°45'S 70°00'W], ii.1993, J.
Cepeda-Pizarro, 3200 m, pitfall, syntopic with
B. perettii, 1 juv. (LEULS).
Diagnosis: Brachistosternus sp. 3 is closely
related to B. montanus, which occurs at the
same latitude and in similar habitats on the
eastern slopes of the Andes, in Argentina. Both
species possess similar hemispermatophores, in
which the distal lamina is slightly curved,
medium-sized, and similar in length to the
basal portion, the internal spines are absent, the
basal spines and row of spines well developed,
and the basal triangle weakly developed
(fig. 41). The two species also exhibit a well-
developed telson gland (fig. 51), similar,
medium-sized androvestigia, and a VM carina
extending the entire length of metasomal
segment V. Both species display a similar
pigmentation pattern in which the carapace is
almost completely pigmented, the tergites
possess a broad transverse stripe or three dark
spots, connected by dense reticulate pigmenta¬
tion, and a single VM and paired VL stripes on
metasomal segments I-V.
The two species may be distinguished as
follows. The VM stripe of metasomal seg¬
ments I-IV is broad and faint, joining the VL
42
AMERICAN MUSEUM NOVITATES
NO. 3564
stripes in all segments of Brachistosternus sp.
3, but narrower, more densely pigmented, and
does not join the VL stripes in segments I—III,
in B. montanus. Both species may also be
distinguished by the dimensions of the pedi-
palp chela of the male (the female of
Brachistosternus sp. 3 is presently unknown).
Brachistosternus sp. 3 exhibits a narrower
pedipalp chela, with a length/width ratio of
4.8-5.09, whereas the chela of B. montanus is
broader, varying from 4.01 to 4.37.
Brachistosternus sp. 3 is sympatric with B.
perettii, which may be distinguished by the
absence of a VM carina on metasomal segment
V, and a telson gland divided into two separate
halves (fig. 50). Brachistosternus sp. 3 exhibits
a well-developed VM carina, extending the
entire length of metasomal segment V, and the
telson gland is entire (fig. 51).
Distribution: Brachistosternus sp. 3 has
been collected in the Central Andes of
Coquimbo, Chile (fig. 77), between 2850 and
3300 m. The localities where it has been
collected fall within the Andes Mediterraneos
botanical subregion of the Desierto Florido
botanical region (Gajardo, 1993).
Notes: A description of this species is in
preparation by Jaime Pizarro, Pablo Agusto,
and the first two authors.
ACKNOWLEDGMENTS
We are grateful to Ivan Benoit
(Corporation Nacional Forestal del
Gobierno de Chile, CONAF) for assistance
with obtaining permits to collect scorpions in
Chilean National Parks, to the CONAF staff
at Pan de Azucar National Park and Llanos
de Challe National Park for assistance during
our visits; to Paula Korob, Jaime Pizarro
Araya, Pablo Agusto, Carolina Cuezzo, Luis
Compagnucci, and Jose Ochoa for their
assistance in the field; to John Ascher for
donating specimens from Chile; and to the
following curators for the loan of material
from collections in their care: Alfredo Peretti
(CDA); Cristina Scioscia (MACN-Ar); Jorge
Artigas (MZUC); Adriano Kury (Museu
Nacional, Universidade Federal do Rio de
Janeiro). This research was partially sup¬
ported by a postgraduate grant from the
Consejo Nacional de Investigaciones
Cientificas y Tecnicas (CONICET) and an
AMNH Collections Study Grant to A.A.O.A.,
and an AMNH Postdoctoral Research
Fellowship to C.I.M. Fieldwork in Chile was
financially supported by the following sources:
L.P. and C.I.M. (2003) from NSF grant EAR
0228699 to L.P.; C.I.M. and Jose Ochoa
(2004) from Idea Wild (www.ideawild.org);
C.I.M. and A.A.O.A. (2005) from the
AMNH; A.A.O.A. (2006) from CONICET
grant PIP 6502. We thank Lee Herman
(AMNH), Oscar Francke (Universidad
Nacional Autonoma de Mexico), and an
anonymous reviewer for their comments on
a previous version of the manuscript.
REFERENCES
Acosta, L.E., and J.A. Ochoa. 2002. Lista de
los escorpiones bolivianos (Chelicerata:
Scorpiones), con notas sobre su distribution.
Revista de la Sociedad Entomologica
Argentina 61(3^4): 15-23.
Abalos, J.W. 1963. Scorpions of Argentina. In H.L.
Keegan and W.V. MacFarlane (editors),
Venomous and Poisonous Animals and
Noxious Plants of the Pacific Region:
111-117. New York: MacMillan.
Acosta, L.E., and E. Maury. 1990. Estridulacion en
Timogenes elegans (Mello-Leitao) (Scorpiones,
Bothriuridae). Bole tin de la Sociedad de
Biologia de Concepcion 61: 29-37.
Agusto, P., C.I. Mattoni, J. Pizarro-Araya, J.
Cepeda-Pizarro, and F. Lopez-Cortes. In press.
Comunidades de escorpiones (Arachnida:
Scorpiones) del desierto costero transicional
de Chile. Revista Chilena de Historia Natural.
Brack, A. 1986. Ecologia de un pais complejo. In C.
Penaherrera del Aguila (editor), Gran geografia
del Peru, naturaleza y hombre, t. 2: 175-319.
Lima: Manfer-Mejia Baca.
Cekalovic, K.T. 1973. Nuevo caracter sexual
secundario en los machos de Brachistosternus
(Scorpiones, Bothriuridae). Boletin de la
Sociedad de Biologia de Concepcion 46:
99-102.
Cekalovic, K.T. 1974. Dos nuevas especies del
genero Brachistosternus (Scorpiones, Bothri¬
uridae). Boletin de la Sociedad Biologica de
Concepcion 47: 247-257.
Cekalovic, K.T. 1975. Brachistosternus ( Leptos -
terms) negrei n.sp. de escorpion de Chile
(Scorpiones, Bothriuridae). Brenesia 6: 69-75.
Cekalovic, K.T. 1983a. Catalogo de los escorpiones
de Chile (Chelicerata, Scorpiones). Boletin de la
Sociedad Biologica de Concepcion 54: 43-70.
2007 OJANGUREN AFFILASTRO ET AL.: CHILEAN BRACHISTOSTERNUS SCORPIONS 43
Cekalovic, K.T. 1982 [1983b]. Estado actual de la
coleccion aracnologica del Museo de Zoologla
de la Universidad de Concepcion (MZUC),
parte Scorpiones. Memorias do Instituto
Butantan 46: 187-192.
Cekalovic, K.T., and J.N. Artigas. 1981. Catalogo
de los tipos depositados en la coleccion del
departamento de zoologia de la Universidad de
Concepcion, Chile. (Parte 3). Bole tin de la
Sociedad Biologica de Concepcion 51(2):
75-107.
Fet, V., G. Lowe, and Family Buthidae C.L. Koch.
1837. In V. Fet, W.D. Sissom, G. Lowe and
M.E. Braunwalder (editors), Catalog of the
scorpions of the world (1758-1998): 55-286.
New York: New York Entomological Society.
Francke, O.F. 1977. Scorpions of the genus
Diplocentrus from Oaxaca, Mexico
(Scorpionida, Diplocentridae). Journal of
Arachnology 4(3): 145-200.
Gajardo, R. 1993. La vegetation natural de Chile,
clasificacion y distribucion geografica. Santiago
de Chile: Editorial Universitaria, 165 pp.
Gervais, P.M. 1841. Arachnides. In Eydoux and
Souleyet (editors), Voyage autour du monde
execute pendant les annes 1836 et 1837 sur la
corvette La Bonite, commandee par M.
Vaillant. Publie par ordre du roi sous les
auspices du Departement de la Marine.
Zoologie. Apteres 1: 281-285. Paris: Arthus
Bertrand.
Koch, L. 1867. Beschreibungen neuer Arachni-
den und Myriapoden. Verhandlungen der
Kaiserlich-Koniglichen Zoologisch-Botanis-
chen Gesellschaft in Wien 17: 173-250.
Kovarik, F. 1998. Stiff (Scorpions). Madagaskar,
Jihlava, 175 pp.
Kraepelin, K. 1894. Revision der Scorpione. II.
Scorpionidae und Bothriuridae. Beiheft zum
Jahrbuch der Hamburgischen Wissenscha-
ftlichen Anstalten 11: 1-248.
Kraepelin, K. 1910 [1911]. Neue Beitrage zur
Systematik der Gliederspinnen. Mitteilungen
aus dem Naturhistorischen Museum (2 Beiheft
zum Jahrbuch der Hamburgischen
Wissenschaftlichen Anstalten) 28(2): 59-107.
Lowe, G., and V. Fet. 2000. Family Bothriuridae
Simon, 1880. In V. Fet, W.D. Sissom, G. Lowe
and M.E. Braunwalder (editors), Catalog of the
scorpions of the world (1758-1998): 17-53.
New York: New York Entomological Society.
Masnu de Moreno, S.J. 1991. Aportes al estudio de
la escorpiofauna mendocina. Revista del
Museo de Historia Natural de San Rafael
11(4): 169-200.
Maury, E.A. 1973. Las tricobotrias y su importan-
cia en la sistematica del genero Brachistosternus
Pocock, 1894 (Scorpiones, Bothriuridae).
Physis (Buenos Aires), Sec. C 32: 247-254.
Maury, E.A. 1974. Escorpiofauna Chaquena. 1. La
verdadera identidad de Brachistosternus
(Microsternus) ferrugineus (Thorell 1876)
(Bothriuridae). Physis (Buenos Aires), Sec. C
33: 73-84.
Ochoa, J.A. 2002. Nueva especie de Brachis¬
tosternus Pocock (Scorpiones: Bothriuridae)
del sur del Peru. Revista Peruana de Biologia
9: 55-63.
Ochoa, J.A. 2004. Brachistosternus ninapo, una
nueva especie (Scorpiones: Bothriuridae) de
los Andes occidentales en el sur del Peru.
Revista Peruana de Biologia 11(2): 139-148.
Ochoa, J.A. 2005. Patrones de distribucion de
escorpiones de la region andina en el sur
Peruano. Revista Peruana de Biologia 12(1):
49-68.
Ochoa, J.A., and L. Acosta. 2002. Two new
Andean species of Brachistosternus Pocock
(Scorpiones: Bothriuridae). Euscorpius 2: 1-13.
Ojanguren Affilastro, A.A. 2001. Sistematica y
distribucion de Brachistosternus alienus
Lonnberg (Scorpiones, Bothriuridae). Revista
del Museo Argentino de Ciencias Naturales
‘Bernardino Rivadavia’ (Buenos Aires) 3:
169-174.
Ojanguren Affilastro, A.A. 2002a. Sistematica y
distribucion de Brachistosternus weyenberghii
Thorell. Revista del Museo Argentino de
Ciencias Naturales ‘Bernardino Rivadavia’
(Buenos Aires) 4: 203-208.
Ojanguren Affilastro, A.A. 2002b. Nuevos aportes
al conocimiento del genero Brachistosternus en
Chile, con la descripcion de dos nuevas especies
(Scorpiones, Bothriuridae). Boletin de la
Sociedad de Biologia de Concepcion 73: 37-46.
Ojanguren Affilastro, A.A. 2002c. Brachistosternus
galianoae (Scorpiones, Bothriuridae) una nueva
especie de Bolivia. Revista del Museo
Argentino de Ciencias Naturales ‘Bernardino
Rivadavia’ (Buenos Aires) 4: 105-109.
Ojanguren Affilastro, A.A. 2003a. The genus
Brachistosternus in Argentina, with the de¬
scription of a new species from Patagonia
(Scorpiones, Bothriuridae). Journal of
Arachnology 31: 317-331.
Ojanguren Affilastro, A.A. 2003b. Las especies
andinas de Brachistosternus (Leptosternus ), con
la descripcion de tres nuevas especies
(Scorpiones, Bothriuridae). Revista Iberica de
Aracnologia 8: 23-36.
Ojanguren Affilastro, A.A. 2004a. Sistematica y
distribucion de Brachistosternus ( Leptosternus )
intermedius Lonnberg. (Scorpiones,
Bothriuridae). Physis (Buenos Aires), Sec. C
59: 29-35.
44
AMERICAN MUSEUM NOVITATES
NO. 3564
Ojanguren Affilastro, A.A. 2004b. Un nuevo
Brachistosternus del norte de Chile
(Scorpiones, Bothriuridae). Revista Iberica de
Aracnologia 10: 69-74.
Ojanguren Affilastro, A.A. 2005a. Notes on the genus
Brachistosternus (Scorpiones, Bothriuridae) in
Chile, with the description of two new species.
Journal of Arachnology 33: 175-192.
Ojanguren Affilastro, A.A. 2005b. Estudio mono-
grafico de los escorpiones de la Republica
Argentina. Revista Iberica de Aracnologia 11:
75-241.
Ojanguren Affilastro, A.A., and C.I. Mattoni.
2006. A new species of Brachistosternus from
the Chilean central Andes (Scorpiones: Both¬
riuridae). Studies on Neotropical Fauna and
Environment 41(1): 79-85.
Ojanguren Affilastro, A.A., and A.H. Roig Alsina.
2001. Brachistosternus angustimanus, una nueva
especie del norte de la Patagonia, Argentina
(Scorpiones, Bothriuridae). Physis (Buenos
Aires), Sec. C 58: 15-22.
Prendini, L. 2000. Phylogeny and classification of
the superfamily Scorpionoidea Latreille 1802
(Chelicerata, Scorpiones): An exemplar ap¬
proach. Cladistics 16: 1-78.
Prendini, L. 2001. Substratum specialization and
speciation in southern African scorpions: The
Effect Hypothesis revisited. In V. Fet and P.A.
Selden (editors), Scorpions 2001. In Memoriam
Gary A. Polis: 113-138. Burnham Beeches,
UK: British Arachnological Society.
Prendini, L. 2003a. A new genus and species of
bothriurid scorpion from the Brandberg
Massif, Namibia, with a reanalysis of bothriur¬
id phylogeny and a discussion of the phyloge¬
netic position of Lisposoma Lawrence.
Systematic Entomology 28: 149-172.
Prendini, L. 2003b. Revision of the genus
Lisposoma Lawrence, 1928 (Scorpiones:
Bothriuridae). Insect Systematics and
Evolution 34(3): 241-264.
Prendini, L., and W.C. Wheeler. 2005. Scorpion
higher phylogeny and classification, taxonomic
anarchy, and standards for peer review in
online publishing. Cladistics 21: 446^494.
Ringuelet, R.A. 1953. Geonemia de los escorpiones
en la Argentina y las divisiones zoogeograficas
basadas en su distribucion. Revista del Museo
de La Plata (N. S.), Zoologia 4: 277-284.
Roig Alsina, A.H., and E.A. Maury. 1981.
Consideraciones sistematicas y ecologicas sobre
Brachistosternus ( Leptosternus ) borellii
Kraepelin, 1911 (Scorpiones, Bothriuridae).
Physis (Buenos Aires), Sec. C 39: 1-9.
Roig Alsina, A.H., and E.A. Maury. 1984.
Sistematica y distribucion geografica de
Brachistosternus ( L .) pentheri Mello-Leitao,
1931 (Scorpiones, Bothriuridae). Physis
(Buenos Aires), Sec. C 42: 17-21.
Rundel, P., M.O. Dillon, B. Palma, H.A. Mooney,
S.L. Gulmon, and J.R. Ehleringer. 1990. The
phytogeography and ecology of the coastal
Atacama and Peruvian deserts. Aliso 13(1):
1-50.
Simon, E. 1880. Etudes arachnologiques. 9e
Memoire (1). XVIII. Descriptions de genres et
especes de l’ordre des Scorpiones. Annales de la
societe entomologique de France 5(10):
377-398.
Soleglad, M.E., and V. Fet. 2003a. The scorpion
sternum: structure and phylogeny (Scorpiones:
Orthosterni). Euscorpius 5: 1-34.
Soleglad, M.E., and V. Fet. 2003b. High level
systematics and phylogeny of the extant scor¬
pions (Scorpiones: Orthosterni). Euscorpius 11:
1-172.
Stahnke, H.L. 1970. Scorpion nomenclature and
mensuration. Entomological News 83:
121-133.
Vachon, M. 1973 [1974]. Etude des caracteres
utilises pour classer les families et les genres
de scorpions (Arachnides). 1. La trichobothrio-
taxie en arachnologie. Sigles trichobothriaux et
types de trichobothriotaxie chez les scorpions.
Bulletin du Museum National d’Histoire
Naturelle (Paris), Ser. 3 140: 857-958.
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. FAX: (212) 769-5009.
@ This paper meets the requirements of ANSI/NISO Z39.48-1992 (Permanence of Paper).