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916
Accepted by P. Jäger: 10 Mar. 2005; published: 24 Mar. 2005 1
ZOOTAXA
ISSN 1175-5326 (print edition)
ISSN 1175-5334 (online edition)
Copyright © 2005 Magnolia Press
Zootaxa 916: 1–15 (2005)
www.mapress.com/zootaxa/
Rediscovery of Orobothriurus bivittatus (Thorell 1877) stat. n.,
comb. n. in the Sierra del Tontal, Argentina
(Scorpiones, Bothriuridae)
LUIS E. ACOSTA
CONICET — Cátedra de Diversidad Animal I, Facultad de Ciencias Exactas, Físicas y Naturales, Univer-
sidad Nacional de Córdoba, Av. Vélez Sarsfield 299, X5000JJC Córdoba, Argentina; lacosta@com.uncor.edu
Abstract
The long neglected scorpion species Orobothriurus bivittatus (Thorell) stat. n., comb. n. (Bothriu-
ridae), formerly considered a subspecies of Urophonius brachycentrus (Thorell) and more recently
placed under synonymy of the type species of Orobothriurus Maury, O. alticola (Pocock), is reval-
idated and redescribed on a male captured in the Sierra del Tontal, west Argentina. It proved to be a
close relative of O. alticola, from which it can be separated by some details of the hemispermato-
phore morphology (e.g. relative length of the lamina apex, lobular expansions at the inflexion point
of the front crest). This species is probably a high altitude endemic to the Precordillera, a range sep-
arated from the Andes (where O. alticola was collected) by a narrow valley. The type locality for O.
bivittatus stat. n., comb. n. (and also for Telegonus weijenberghi Thorell) is restricted to the upper
belts of the mentioned range, as thoroughly discussed. To ensure nomenclatural stability, it is pro-
posed to set aside the existing holotype and to designate the studied male as neotype.
Key words: Neotropics, Precordillera, systematics, nomenclature, type locality
Introduction
The scorpion species Cercophonius brachycentrus bivittatus Thorell 1877 (Bothriuridae),
currently under the synonymy of Orobothriurus alticola (Pocock 1899), is probably the
most enigmatic species of the Argentinean scorpiofauna. It was described on a single juve-
nile, as a presumable "variety" of Cercophonius brachycentrus Thorell 1877 (today in
Urophonius Pocock 1893), but remained ignored in the literature for more than 50 years
(Acosta 2002). It was Mello-Leitão (1931) who "rescued" the name, by formally assigning
to it a subspecific rank, Urophonius brachycentrus bivittatus. That change was just
nomenclatural, since no author, not even Mello-Leitão, knew what kind of scorpion bivit-
tatus was. For a further half a century the few subsequent citations of the name simply fol-
ACOSTA
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ZOOTAXA lowed uncritically Mello-Leitão's (1931) proposal. Maury (1977) first reviewed the type
and only available specimen. He concluded it does not belong to genus Urophonius, but
more probably was a synonym of Orobothriurus alticola, type species of genus Orobothri-
urus Maury 1976. It should be noted that the concept of Maury (1976) for O. alticola was
wider than accepted today (Acosta & Ochoa 2001). Maury (1977) did not formally pro-
pose the synonymy, and bivittatus remained in its old taxonomical placement, as still cited
by Acosta & Maury (1998) and Lowe & Fet (2000). Acosta (2002) formally stated the syn-
onymy bivittatus = alticola.
The mentioned synonymy would have meant that the older and long uncertain name
bivittatus had priority over the well-established name of the type species, alticola. Since
this was deemed to threaten the nomenclatural stability, Acosta (2002) also asked the
International Commission on Zoological Nomenclature to use its plenary powers, so as to
give alticola conditional precedence over bivittatus, whenever these names are considered
synonyms (Case 3213). The Commission agreed with this application, as ruled through
Opinion 2074 (ICZN 2004). The conditional precedence was considered central in this
proposal, since the type of bivittatus is a very small, not well preserved juvenile, a fact that
weakens the certitude of the taxonomical judgment. Genus Orobothriurus, with most spe-
cies from high altitude sites, is seemingly not well known, while it shows a high degree of
endemicity in elevated Andes-related ranges (Acosta & Ochoa 2001, Ochoa 2004). In
other words, it is still quite possible to discover other species anywhere, including the
actual bivittatus in its original sense. If that is the case, there is already a name available
for it. The point was then, to determine precisely what Cercophonius brachycentrus bivit-
tatus is, and where it occurs, a question that had first to be tackled by a thorough analysis
of the information on the type locality, to ascertain its actual location. The second step
would be the collecting effort to find the species there. This two-fold task was successful
and I am now able to recognize Orobothriurus bivittatus stat. n., comb. n. as a valid spe-
cies. These results, together with a redescription of Thorell's (1877a) species, based on an
adult male, are shown below. After O. alticola and O. famatina Acosta in Acosta & Ochoa
2001, O. bivittatus is the third nominal species of the genus identified from Argentina,
although, according to materials I have examined and to the known distributional patterns,
other species likely remain to be described (Acosta pers. obs.). With the removal of several
species in a separate genus (Ochoa 2004) and the inclusion of O. bivittatus, Orobothriurus
is now composed by 10 nominal species inhabiting Andean and sub-Andean sites, from
Peru, Chile and Argentina.
Materials and methods
Carinae of metasomal segments are named after Francke (1977), as follows: DL: dorsal
lateral, DSM: dorsal submedian, LIM: lateral inframedian, LM: lateral median, LSM: lateral
supramedian, VL: ventral lateral, VM: ventral median, VSM: ventral submedian. Trichobo-
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thriotaxial notation according to Vachon (1974): Eb2: basal 2, Esb: sub-basal, Est: sub-ter-
minal, Et3: terminal 3, all of the external series of the manus. In the section comparing
populations of O. alticola, those from the province of Mendoza are referred to as MZA,
and SJ for province of San Juan. Acronyms of repositories: CDA (Cátedra de Diversidad
Animal I, Facultad de Ciencias Exactas, Físicas y Naturales, Universidad Nacional de Cór-
doba), MACN (Museo Argentino de Ciencias Naturales, Buenos Aires), NRS (Naturhis-
toriska Riksmuseet, Stockholm).
Material examined of Orobothriurus alticola. ARGENTINA. Province of Mendoza.
Puente del Inca, 16 February 1977 (A. Roig), 3 males, 2 females, 2 juv. (MACN); same
loc. and coll., 18 January 1976, 3 juv. (MACN); Los Horcones, 19 January 1976 (A. Roig),
2 juv. (MACN); Complejo Fronterizo "Los Horcones", 1 km W de Puente del Inca (2800
m), 17 January 2000 (C. Mattoni), 1 female (CDA 000.060). Province of San Juan. Paso
del Agua Negra (3250 m), 17 February 1978 (A. Roig), 2 males, 2 females, 1 juv.
(MACN), 1 male, 1 female (CDA 000.061); same loc. and coll., 23 January 1982, 1 male,
2 females (MACN); Paso del Agua Negra (4050 m), 17 February 1978 (A. Roig.), 1 male,
1 juv. (MACN); "El Galpón", 15 km W Arrequintín (3500 m), 23 January 1982 (E.
Maury), 2 females (MACN).
Orobothriurus bivittatus (Thorell 1877) stat. n., comb. n.
Figs. 1–7
Cercophonius brachycentrus var. β bivittatum Thorell 1877a: 183 [recte bivittatus].
Urophonius brachycentrus bivittatus: Mello-Leitão 1931: 99–100; 1933: 35; 1934: 48, 51; 1938:
94–95; 1945: 213, 215; Acosta & Maury 1998: 559; Lowe & Fet 2000: 44.
Urophonius brachicentrus bivittatus: Mello-Leitão 1939: 612; Abalos 1959: 592; 1963: 117; Roig
Alsina 1973: 200 (incorrect subsequent spelling).
? [Orobothriurus alticola]: Maury 1977: 148 (synonymy and combination suspected, not formally
proposed)
Orobothriurus alticola: Acosta 2002: 177 (formal synonymy, here removed).
Type material. Holotype juvenile (NRS), examined. This specimen is stored in a small
vial with the label "Urophonius brachycentrus bivittatus (Thorell), E. Maury det. 1971". In
turn, this vial is placed within a larger vial labeled: "Cercophonius brachycentrus Thor.,
Córdova (pullus), S. Juan, Argentina, Weijenb.", and containing a further small vial with
one adult male and a second juvenile (these two specimens constituting the type series of
Urophonius brachycentrus). Because the original holotype is a poorly preserved juvenile,
it provides little information on the species identity, what may render it as a nomen
dubium. Therefore, an application was sent to the ICZN to set it aside, and to designate the
male described below as neotype, according to Art. 75.5 (Case 3332). Conditions qualify-
ing this material as neotype, especially the evidence that it is consistent on what is known
on the species and its probable locality, are shown below. This new specimen will have the
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ZOOTAXA advantage of being an adult male, bearing the most valuable taxonomic features (espe-
cially the hemispermatophore), and the type locality will become precise. Until a decision
is made by the ICZN, the current usage (NRS holotype) is to be maintained.
FIGURES 1–5. Orobothriurus bivittatus (Thorell 1877) stat. n., comb. n., male (CDA); 1. Metaso-
mal segment V and telson, lateral view; 2. Metasomal segment V, ventral view; 3–5. Right pedipalp
chela; 3. Ventromedial view; 4. Ventral view; 5. Lateral view. Scale bars: 1 mm.
Additional material examined. ARGENTINA. Province of San Juan. Parque Nacional
El Leoncito, Sierra del Tontal, near 'Portezuelo del Tontal' (3450 m), U.V., 25
November
2003 (L. Acosta), 1 male (CDA 000.364).
Description (adult male).
Coloration. General color reddish hazel, ventral side and legs lighter, pectines whitish
yellow. Carapace with a dense pigment reticle, leaving a clear depigmented area on the
anterior border, between the lateral eyes. Tergites I–VI with two wide paralateral pig-
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mented stripes, leaving a clear median space in between (representing 15.9% of the tergite
V width), and narrow clear areas each side; tergite VII with dense paramedian reticles,
irregularly connected in the middle. Sternites depigmented. Metasomal segments I–III
with a dark median dorsal spot, shield-like, which does not reach the posterior border, but
bears three anterior projections; this spot is vestigial on segment IV, as a tenuous reticle
with similar arrangement; dorsal side of segment V almost depigmented. Lateral side of
segments I–IV with two longitudinal well-defined stripes of dense pigment. The lateral
stripe occupies the basal half on segment I, between DSM and DL, while it is almost com-
plete on segments II–IV, extended along DL; on segment V it is faint and reticulate. The
ventrolateral stripe, of darker pigment, extends from the VL carina to the lateral side, and
is well developed along the entire metasoma; on segments I–III it is separated from the lat-
eral stripe by a narrow space, while a feeble reticle connects these stripes on segments IV–
V. Ventromedian line absent on segment I, on segments II–V it is well defined, and
remains independent from the lateroventral pigment. Vesicle with faint ventral pigment, a
narrow median stripe is separated from the lateral pigmented areas by two clear lines. Che-
licerae just with a vestigial reticle. Pedipalp femur dorsally covered by pigment, except for
an elongated clear area on the basal half; remaining sides only with faint reticles. Patella
with dense reticle on the retrolateral side, and some pigment on the prodorsal and proven-
tral borders. Chela with a large ventral, retrolateral and prolateral spot on the fingers' base,
connected to the hand base by several irregular, faint stripes. Legs spotted on the external
and dorsal sides (especially femur and patella), other sides clear.
Morphology. Small sized scorpion, with slender and delicate appearance; measure-
ments, see Table 1. Carapace depressed, with very low eye mound, front border straight.
Tegument of carapace and tergites I–VII finely granular, acarinate; tergite VII is more
granulous and bears four incomplete posterior keels. Sternites I–IV smooth; sternite V
granulous, with one pair of paralateral keels on the posterior half. Metasomal segments I–
IV. DL keels complete and granulous, in each segment granules are caudally larger and
more acute (ending in a posterior larger granule), but carinae become weaker from seg-
ment I to IV. LSM complete in segment I; only present on the caudal one third on segment
II; just as a smooth tegumentary elevation and a few isolated posterior granules on seg-
ment III; lacking on segment IV. LIM complete and conspicuous in segment I, then
decreasing suddenly on segment II to just a few posterior granules, and disappearing on
segments III–IV. Ventral surface of segment I granulous, VL and VSM well developed and
conspicuous (also a granulous area between LIM and VL); on segment II granulation is
markedly reduced, VL are just tegumentary borders with very few sparse granules and
VSM are not discernible; both VL and VSM are absent on segments III–IV (ventral surface
smooth). Metasomal segment V: DL carinae almost lacking, with just very few and incon-
spicuous basal granules. Lateral surface with LM only insinuated. VL almost complete but
somewhat dispersed in an irregular double row in the middle, with large granules alternat-
ing with smaller ones. Few sparse granules represent the otherwise lacking VSM. VM com-
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ZOOTAXA plete, more 'regular' than VL, expanded on the posterior end. Telson: vesicle elongated and
low, aculeus short. Movable finger of chelicerae with two subdistal teeth. Pedipalps slen-
der and smooth, except for the prolateral face of femur, densely granulous, and a few gran-
ules on the prodorsal and proventral borders of the patella. Chela slender, with an acute
prolateral apophysis. Trichobothriotaxy according to the genus pattern (as described by
Ochoa 2004); Et3 slightly more basal than Est, Esb situated above Eb2. Telotarsal spina-
tion on legs I–IV: 1+1, 2+2, 3+3, 3+3. Number of pectinal teeth: 19-19. Hemispermato-
phore very slender, like in other members of the "species group A" (as recognized by
Acosta & Ochoa 2001), the lamina is sharply divided in a straight basal part, and an S-
curved apical part (apex); frontal crest along the border of the basal part of the lamina with
two portions (proximal oblique and straight distal); a lobular projection each side of the
crest at the inflexion point; basal lobe as typical "tortuous" stem, tipped by a delicate distal
process (the subdistal "spoon-like" dilatation is suspected to have been damaged during
dissection).
TABLE 1. Measurements (mm) of the studied male of Orobothriurus bivittatus (Thorell 1877) stat.
n., comb. n.
Holotype, juvenile (presumably female). Damaged, only the following parts remain:
carapace, mesosoma and caudal segments I–IV, together with left pedipalp trochanter and
femur, right I–II legs (leg I without telotarsus), and pectines; metasomal segment V and
Total body length 24.4
Carapace length 3.1
Anterior / posterior width 1.9 / 3.5
Mesosoma length 6.4
Metasoma total length 15.0
Metasomal segment I length / width 1.5 / 1.9
Metasomal segment II length / width 1.8 / 1.7
Metasomal segment III length / width 1.9 / 1.6
Metasomal segment IV length / width 2.4 / 1.5
Metasomal segment V length / width / height 3.4 / 1.4 / 1.2
Telson length / width / height 4.0 / 1.3 / 1.1
Aculeus length 0.8
Pedipalp total length 11.2
Femur length / width 3.0 / 0.8
Patella length / width 2.9 / 1.0
Chela length / width / height 5.3 / 1.4 / 1.4
Movable finger length 2.9
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right pedipalp are loose; all remaining body parts or appendages missing. Pectinal teeth:
15–12 [in the latter it is evident that several teeth are lacking].
FIGURES 6–7. Orobothriurus bivittatus (Thorell 1877) stat. n., comb. n., left hemispermatophore;
6. Internal view; 7. External view. Figure 8: O. alticola (Pocock 1899), left hemispermatophore,
external view. All drawings at the same scale; scale bar: 1 mm.
Comparisons. Both the external morphology and the hemispermatophore shape place
O. bivittatus in the so-called "group A" (Acosta & Ochoa 2001), which proved to be a
monophyletic ensemble (Ochoa 2004). Within the group, it is closely related to O. alticola.
ACOSTA
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ZOOTAXA Similarities concern both the pigment pattern and the external morphology, being difficult
indeed to find a sharp morphological discrimination. Only the hemispermatophore pro-
vides reliable differences. The fact that just one male of O. bivittatus is hitherto available
– i.e. variability remains unknown – lessens to some extent the strength of these conclu-
sions, but the differences encountered, together with the apparent geographical isolation,
support the latter being regarded as an independent entity. Whether it represents a separate
species or a subspecies (in which case the name should be subordinated to O. alticola) is
probably a useless question. With the taxonomic situation being subtle, both the condi-
tional precedence of alticola and the designation of a neotype for bivittatus are considered
valuable nomenclatural tools to ensure stability at different levels. As shown in Fig. 9, O.
alticola is hitherto known from two quite separate areas on the east side of the Andes in
western Argentina (Acosta & Ochoa 2001): surroundings of Puente del Inca, province of
Mendoza (type locality), and of Paso del Agua Negra, province of San Juan (referred to as
MZA and SJ, respectively). These two populations evidence some slight external differ-
ences (see below), but the hemispermatophore morphology is identical. In comparison, the
hemispermatophore of O. bivittatus is in overall outline more slender, especially regard-
ing the basal part of the lamina (Figs. 6–8). Other differences are recorded in the partial
key below. O. bivittatus seems to be smaller, with total length (24.4 mm) below the size
range of O. alticola (MZA: 26.9–31.5 mm, mean=28.7 mm, SJ: 27.1–31.5 mm,
mean=28.9 mm); females reach 34.2 mm in MZA and 37.5 mm in SJ. To insert O. bivitta-
tus in the key provided by Ochoa (2004), the very last couple should be modified, as fol-
lows:
7. Sternite V granous, carinae not discernible. Ventral side of metasomal segment V cov-
ered by dense granulation that hides carina VM. Legs III–IV with 3+4 telotarsal spines.
Ventral pigment of metasoma: axial stripe joins distally the ventrolateral pigment on
segments I–III (not joining on segments IV–V) ...........................................O. atiquipa
– Sternite V little granulous, with 2–4 carinae. Ventral side of metasomal segment V
without dense granulation, carina VM well defined. Legs III–IV with 3+3 telotarsal
spines. Ventrolateral and ventro-axial pigment stripes independent along the entire
metasoma ......................................................................................................................8
8. Hemispermatophore: apex reduced, it represents only 38% of the lamina length; front
crest almost not surpassing its connection to the apex, and without lobular expansion
at the inflexion point. Metasomal segment II with VSM carinae. Metasomal segment
IV with six VSM macrosetae. Total length (males): 27–31.5 mm .................O. alticola
– Hemispermatophore: apex larger, 47% of the lamina length; front crest surpassing its
connection to the apex, forming in lateral view a 'spur-like' projection (notch in
between with slightly different curvature: Figs. 6–8), and with lobular expansions at
the inflexion point. Metasomal segment II without VSM carinae. Metasomal segment
IV with eight VSM macrosetae. Total length (male): 24.4 mm ................. O. bivittatus
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In other features, currently employed in the species-level taxonomy of Orobothriurus
(Acosta & Ochoa 2001), the single specimen O. bivittatus fits within the variability range
of O. alticola:
Frequencies of the number of pectinal teeth.— O. bivittatus (19-19). MZA males
(n=12 pectines): 18 teeth (2 pectines), 19 (2), 20 (6), 21 (2); females (n=11): 15 (7), 16 (4);
SJ males (n=16 pectines): 19 teeth (7 pectines), 20 (5), 21 (4); females (n=16): 15 (1), 16
(5), 17 (9), 18 (1).
Number of VSM macrosetae on metasomal segment I.— This feature shows the most
noticeable difference between the two populations of O. alticola. MZA: 3+3: 10 speci-
mens (83.3%), 3+2: 2 specimens (16.7%); SJ: 3+3: 4 specimens (25.0%), 3+2: 3 spec.
(18.7%), 2+2: 9 spec. (56.2%). The single available O. bivittatus has 3+3 setae.
Telson, length/height ratio.— O. bivittatus (3.64). MZA males (n=3): 3.1–3.8, females
(n=3): 3.1–3.2; SJ males (n=6): 3.8–4.2 (mean=3.9), females (n=8): 2.8–3.3 (mean=3.0).
Pedipalp chela, length/width ratio.— O. bivittatus (3.9). MZA males (n=3): 3.3–3.6,
females (n=3): 3.4–3.7. SJ males (n=6): 3.5–4.2 (mean=3.8), females (n=8): 3.3–3.5
(mean=3.4).
Relative width of the median clear stripe.— O. bivittatus (15.9% of the width of tergite
V). MZA: 14.4–19.3% (mean=15.9%), n=8; SJ: 11.1–16.1% (mean=14.5%), n=15.
Type locality. The type specimen is said to come from "San Juan", without further
detail. This material was sent to Thorell by the Dutch zoologist Hendrik Weyenbergh,
together with other arachnids collected 'around Córdoba and San Juan' (Thorell 1877b,
1878). In addition to "Cercophonius brachycentrus var. β bivittatum", Thorell (1877a)
described other four scorpion species on that material, and two harvestmen and one pseu-
doscorpion were added in a further publication (Thorell 1877b, 1878). The place of origin
given in some cases was questioned, and some confusion between these two localities is to
be suspected. Despite Thorell's (1877b, 1878) statement that all specimens were collected
by Weyenbergh himself, those from San Juan were actually obtained by the botanist Saile
Echegaray. As Hieronymus (1881) complained, Weyenbergh never was in San Juan, and
the mistaken statement of Thorell (1877b, 1878) was said to be a 'printer's error'. This fact
might have contributed to the confusion of labelling, but at the same time gives additional
evidence to our case, as we will see. The species with presumably wrong localities are:
Telegonus weijenberghi Thorell 1877a: 173 (Scorpiones, Bothriuridae; currently under
Brachistosternus Pocock 1893). Supposedly collected at "Córdoba". The species was
never found there (Acosta 1989, Acosta & Rosso de Ferradás 1996), while, in contrast,
it was recorded in localities from Western and Northwestern Argentina (Ojanguren
Affilastro 2002), among them several sites in province of San Juan.
Cercophonius brachycentrus Thorell 1877a: 180 (Scorpiones, Bothriuridae; currently
under Urophonius Pocock). The type series comprises two specimens: an adult male is
stated to be from "San Juan", and a juvenile from "Córdoba". The latter is correct, but
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ZOOTAXA the male is probably mislabeled. This species is widely distributed in central Argen-
tina, being quite common in Córdoba, but was never found in San Juan (Maury 1977,
Acosta & Rosso de Ferradás 1996).
Pachylus butleri Thorell 1877b: 207 (Opiliones, Gonyleptidae, currently under Pachy-
loidellus Müller 1916). From "San Juan". This species is not found there, but is very
common and frequently collected in Córdoba instead (Acosta 1993a).
Meanwhile, the type localities of Isometrus fuscus Thorell 1877 (Córdoba), Telegonus
ferrugineus Thorell 1877 (Córdoba) and Ostracidium pertyi Thorell 1877 (Córdoba) are
consistent with their known range, and thus deemed to be correct (Abalos 1953, Maury
1974, Acosta 1989, Acosta 1993b, Acosta & Rosso de Ferradás 1996). Species from Cór-
doba can easily be assumed to have been collected in the surroundings of the city, in those
times little more than a small village. Urophonius brachycentrus, Pachyloidellus butleri,
Brachistosternus ferrugineus, Neopucroliella pertyi are still occasionally collected in the
city suburbs, while Zabius fuscus can be caught in sites less than 15 km from the center
(Acosta 1989, 1993a, 1993b). In contrast, the exact collecting sites in 'San Juan' seem not
as easy to determine, since neither Brachistosternus weijenberghi nor any Orobothriurus
species exist in the 'surroundings' of the capital city (Roig Alsina & Maury 1981, Ojan-
guren Affilastro 2002, Acosta & Ochoa 2001). The right answer should be searched in the
itinerary of the collector, Saile Echegaray, in his botanical trip in this province.
Echegaray (1878) traveled to San Juan in November 1875. His excursion was targeted
to the locality of "Leoncito", today best known as "Estancia El Leoncito", which is now
partially embraced in the Parque Nacional El Leoncito. It was an extensive farm, whose
main settlement can still be seen on the base of the western slopes of the Sierra del Tontal,
at about 2300 m. This mountain belongs to the Precordillera orographic system and
reaches over 4000 m a.s.l. In his report, Echegaray (1878) indicates he visited the 'Cerro
Tontal' where he collected many plant samples. At the present day there is no practicable
road crossing this range, but surprisingly enough, in Echegaray's times the so called 'road
of Tontal' was the only available communication between the city of San Juan and El
Leoncito. This road, abandoned long ago, crosses the mountain at the pass known as
'Portezuelo del Tontal' or 'Portezuelo de la Virgen', at 3640 m. Taking all the evidence
together, the most plausible locality for Thorell's (1877a) scorpions collected in 'San Juan'
appears to be 'somewhere in the Sierra del Tontal'. Against that reasoning, Roig-Juñent et
al. (2003) stated (as a 'prediction' of their analysis) that the scorpion genus Orobothriurus
does not occur in the Precordillera; but as seen, this is not supported by the facts. There-
fore I hereby restrict the type locality of Cercophonius brachycentrus bivittatus (and also
of Telegonus weijenberghi) to a so far undetermined site along the old road on the Sierra
del Tontal, at least above 3000 m.
Zoogeographic comments. Orobothriurus bivittatus is likely a high-altitude endemic
in the Sierra del Tontal. This mountain continues southerly into the Sierra de Uspallata,
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province of Mendoza (Fig. 9), which is a part of the Precordillera, too. Both chains are
separated from the Andes by the "Uspallata-Calingasta" tectonic depression, a xeric, N-S
oriented 'valley' composed by two adjacent basins (Fig. 9: Ca, Us), with watershed at
around 2200 m. The Uspallata-Calingasta valley is covered by Monte (sub-Andean) vege-
tation (Martinez Carretero 1995). Though being quite narrow (50–100 km), it likely repre-
sents an effective barrier for low vagility high-Andean elements (Roig Juñent et al. 2003).
At the Sierra del Tontal the Monte ascends the west-faced slopes, reaching as high as
2800–3000 m (e.g., at Pampa del Jarillal); at least on the path to Portezuelo del Tontal,
pure grasslands only appear above 3200 m (Acosta, pers. obs.). In general, the upper
regions of the Tontal and Uspallata ranges should be ascribed to the Puna vegetation (Mar-
tinez Carretero 1995), but the most elevated sites of the former show many high-Andean
elements, even with a marked ecotonal character (E. Martínez Carretero, in litt. 2004).
Grasslands of the Sierra del Tontal are just 60 km away from the Andes, and this isolation
might explain the hemispermatophore differences between the close relatives O. alticola
and O. bivittatus. In contrast, the two populations of O. alticola (MZA-SJ), though sepa-
rated by 280 km, are very probably continuous along the intricate Andes eastern slope
(Fig. 9), and thus their hemispermatophores do not show any divergence. Since differences
between alticola and bivittatus are small, a relatively recent separation is supposed. Past
fluctuations of the altitudinal belts, in which the high-altitude vegetation might have low-
ered its limit in colder/more humid periods, would have connected the Andes and the Pre-
cordillera; in such case, the continuity of the ancestor range would have been enabled
along these two mountain regions, being then split during a retraction phase. Palynological
studies made by Markgraf (1983) show that a locality at 2000 m in the Uspallata-Calin-
gasta depression had a predominance of graminous vegetation between 5,000 and 4,000 yr
B.P., decreasing thereafter, to give place to present Monte elements. This scenario contra-
dicts the presumed unrelatedness of the Andes and the Precordillera, as proposed by Roig
Juñent et al. (2003). These authors mention the evidence of several tenebrionid beetles,
which are endemic to the Andes (e.g. Falsopraocis) but do not occur in the Precordillera at
the same latitude. Their results, however, might have been biased by the scarce informa-
tion on Andean and Precordilleran species considered in their analysis. Since the arthropod
fauna of the Sierra del Tontal is poorly known, it is highly advisable to re-analyze the
Andes-Precordillera links by including new information from this neglected range.
Considering the lowest records in Argentina (2450 m for O. famatina, 2700 m for O.
alticola), Orobothiurus would be expected to exist in the Sierra de Uspallata. But despite
repeated sampling efforts, no species of this genus has hitherto been found there. It is
worth noting that the Sierra de Uspallata only slightly surpasses 3000 m at its highest ele-
vation, and this altitude is not continuous with the Sierra del Tontal, but a lower area (at
~2700 m) separates the upper portions of the two ranges. All known Argentinean records
of Orobothriurus (Maury 1976, Acosta & Ochoa 2001) originate from mountains whose
maximal altitude reaches or surpasses 3500 m.
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FIGURE 9. Localities of Orobothriurus bivittatus (Thorell 1877) stat. n., comb. n. (square) and O.
alticola (Pocock 1899) (circles) in west Argentina. Dark grey: areas above 3600 m; light grey:
between 2700 and 3600 m. Dashed line: administrative provincial boundaries, thick solid line:
international boundary. ST: Sierra del Tontal, SU: Sierra de Uspallata, Ca: valley of Calingasta, Us:
valley of Uspallata. 1: collecting sites near Paso del Agua Negra, 2: Puente del Inca. The city of San
Juan is indicated with a small dot.
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Acknowledgements
For the loan of the type of Orobothriurus bivittatus I am indebted to Dr Torbjörn Kronest-
edt (NRS). The late Dr Maria Elena Galiano (MACN) made available additional materials
of O. alticola. Dr Gustavo Flores helped me in the collecting trip to the Sierra del Tontal,
and Dr Eduardo Martínez Carretero provided valuable information on the Precordillera
vegetation. Special thanks are due to the Administración Nacional de Parques Nacionales
for permission to sample in the Parque Nacional El Leoncito, and to Guardaparque Héctor
Almonacid for logistic support in the field work. Financial help was obtained from Secre-
taría de Ciencia y Tecnología, Universidad Nacional de Córdoba. I acknowledge com-
ments made by an anonymous reviewer and the subject editor, which improved the final
version of the text. The author is a researcher of the Argentinean Consejo Nacional de
Investigaciones Científicas y Técnicas.
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