Some taxonomic corrections to the genus Tityus C. L. Koch, 1836 (Scorpiones: Buthidae) in Hispaniola, Greater Antilles

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Some Taxonomic Corrections to the Genus Tityus C. L. Koch,
1836 (Scorpiones: Buthidae) in Hispaniola, Greater Antilles
Rolando Teruel
March 2017 – No. 242

Euscorpius
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Publication date: 15 March 2017

Euscorpius — Occasional Publications in Scorpiology. 2017, No. 242
Some taxonomic corrections to the genus Tityus C. L. Koch,
1836 (Scorpiones: Buthidae) in Hispaniola, Greater Antilles
Rolando Teruel
Centro Oriental de Ecosistemas y Biodiversidad, Museo de Historia Natural "Tomás Romay"
José A. Saco # 601, esquina a Barnada, Santiago de Cuba 90100, CUBA
E-mail: rteruel@bioeco.cu
Summary
In the present paper, the taxonomic status of several Hispaniolan members of the genus Tityus C. L. Koch, 1966 is
revised after examination of almost all primary types and abundant supplementary material. This resulted in six new
synonymies, which involve both extant and fossil species. The extant taxa herein synonymized are Tityus anasilviae
Armas et Abud, 2004 under Tityus ottenwalderi Armas, 1999, Tityus bahoruco Teruel et Armas, 2006 under Tityus
crassimanus (Thorell, 1876), Tityus ebanoverde Armas, 1999 under Tityus elii Armas et Marcano, 1992, and Tityus
septentrionalis Armas et Abud, 2004 under Tityus portoplatensis Armas et Marcano, 1992. The fossil taxa herein
synonymized are Tityus azari Lourenço, 2013† and Tityus (Brazilotityus) hartkorni Lourenço, 2009†, both under
Tityus geratus Santiago-Blay et Poinar, 1988†. Updated distribution maps are given for all extant senior synonyms.
The buthid scorpion genus Tityus C. L. Koch, 1836,
is very diverse in the Greater Antillean island of His-
paniola: the most recent catalog by Santos et al. (2016)
listed 14 nominal species, all endemic and unevenly
divided into the "crassimanus" and "quisqueyanus"
species-groups (4 and 10 species, respectively). As
expected from such diversity, it also widespread in the
island and has colonized almost all available eco-
systems. Its members occur from the seashore to the
highest peaks above 3,000 m a.s.l., and in all vegetation
types from desert to rainforest, including montane
savannas and pine forests (Armas, 1987, 1999; Teruel,
2005; Teruel & Armas, 2006; Kovařík & Teruel, 2014).
Apart from these extant taxa, three fossil species have
been described from Dominican amber (Santiago-Blay
& Poinar, 1988; Lourenço, 2009, 2013), but their correct
infra-generic assignation to subgenera or species-groups
is still pending for two of them and unsatisfactory for the
other.
Very recently, Santos et al. (2016: 16) declared that
the correct taxonomic status of three extant species was
under investigation. The present author was finally able
to study the type-series of all Tityus described from
Hispaniola and deposited in IES collection, together
with abundant supplementary material captured by him-
self and collaborators during three intensive field trips to
Dominican Republic (2005, 2014 and 2016), plus the
collections of IES, MHNHSD and MSU, plus additional
specimens kindly donated by friends and colleagues.
The detailed comparison of all this material (more than
200 specimens in total) allowed clarifying the precise
status of four extant and the three fossil taxa: six cases of
synonymy were discovered. All pertinent nomenclatural
changes are introduced and discussed in detail in the
present paper.
Abbreviations used in the text for specimen repos-
itories are as follows: Instituto de Ecología y Sis-
temática, Havana, Cuba (IES), Museo Nacional de
Historia Natural, Santo Domingo, Dominican Republic
(MNHNSD), Montana State University, Bozeman, Mon-
tana, USA (MSU), and personal collection of the author
(RTO).
Tityus anasilviae Armas et Abud Antun, 2004
Figures 1, 6
Type Data. DOMINICAN REPUBLIC, Cordillera
Central [Central Range], Monseñor Nouel Province,
Bonao Municipality, Blanco, Arroyón [= Los Guázaros],
700 m a.s.l., inside pillowcase in house, 8/Sep-
tember/2002, A. Abud, A. S. Reynoso, ♂ holotype (IES:
CZACC-3.3145, examined).
Remarks. This species was described upon a single
adult male by Armas & Abud Antun (2004) and
accepted as valid by Teruel & Armas (2006). An adult
female was described later by Armas & Abud Antun
(2015), who stated that one of the main characters
alleged as diagnostic for this taxon (dorsolateral carinae
with terminal granule much stronger on metasomal
segments II–IV), was absent in the new specimen and
more samples were needed to define if this implied
sexual dimorphism or just individual variation.
The direct comparison of the holotype of T.
anasilviae to the types of Tityus ottenwalderi Armas,

Euscorpius — 2017, No. 242
2
Figure 1: Size-related morphological variability of adult Tityus ottenwalderi, exemplified by females: a) small specimen from
El Río, La Vega Province (typical T. ottenwalderi); b) standard specimen from Pinar Quemado, La Vega Province (intermediate);
c) large specimen from El Majagual, San Cristóbal Province (typical T. anasilviae n. syn.). Scale bar in millimeters.
1999 (the only other member of the "crassimanus"
species-group known to occur in this mountain range)
and abundant additional specimens revealed that both
taxa are conspecific. Thus, the following synonymy is
herein established: Tityus ottenwalderi Armas, 1999 =
Tityus anasilviae Armas et Abud Antun, 2004, new
synonym.
The present study proved that all Hispaniolan
members of the "crassimanus" species-group are so
highly variable morphologically, that adults belonging to
different size-classes can be easily mistaken as distinct
species. The expression of sexual dimorphism follows
the same rule as in other studied species of the genus,
i.e., larger adults are the most strongly dimorphic and
smaller ones are the least (Fig. 1).
Moreover, the degree of attenuation of pedipalps
and metasoma shows a clinal variation positively
correlated to altitude: the most slender specimens (i.e.,
typical T. ottenwalderi), come from mountains above
1,400 m, the most robust ones (i.e., typical T. anasilviae
n. syn.) come from the slopes below 800 m, and inter-
mediate individuals occur all across the intervening
altitude zone (Fig. 1).
Tityus bahoruco Teruel et Armas, 2006
Figures 2, 6
Type Data. DOMINICAN REPUBLIC, Sierra de
Bahoruco [Bahoruco Range], Pedernales Province,
Pedernales Municipality, Mencía, Río Mulito [= El
Mulito, Las Agüitas], 500 m a.s.l., under tree bark,
22/August/1987, A. Abud, L. F. de Armas, ♂ holotype
(IES, examined). Sierra de Bahoruco [Bahoruco Range],
Pedernales Province, Pedernales Municipality, Las Abe-
jas, 1,290 m a.s.l., inside rotten log, 22/July/1999, M. A.
Ivie, ♀ paratype (MSU, examined).
Remarks. The direct comparison of the types of T.
bahoruco to abundant additional specimens of T.
crassimanus (the only other member of the "crassi-
manus" species-group known to occur in this mountain
range), revealed that both taxa are conspecific. Both

Teruel: Taxonomic Corrections to Tityus in Hispaniola
3
Figure 2: Size-related morphological variability of adult Tityus crassimanus, exemplified by males: a) small specimen from
road Cabo Rojo-El Aceitillar, Pedernales Province (typical T. bahoruco n. syn.); b) standard specimen from road Cabo Rojo-El
Aceitillar, Pedernales Province (intermediate); c) large specimen from road Cabo Rojo-El Aceitillar, Pedernales Province
(intermediate); d) very large specimen from Río Mulito, Pedernales Province (typical T. crassimanus). Scale bar in millimeters.
types of the former are just the smallest adults of the
latter and the comparison made in the original des-
cription was misleading: a reexamination of the two
supposedly adult males of T. crassimanus from Fondo
Paradí used for comparison by Teruel & Armas (2006),
revealed that both are actually very large females. Thus,
the following synonymy is herein established: Isometrus
crassimanus Thorell, 1876 [currently Tityus crassimanus
(Thorell, 1876)] = Tityus bahoruco Teruel et Armas,
2006, new synonym.
The present study (which included successful
captive breeding for several consecutive generations)
allowed to define that T. crassimanus is morphologically
highly variable, that sexual maturity is attained at very
different instars (from nymphs III–VI), and that
morphological variation is directly correlated to size, to
the point that adults belonging to different size-class can
be easily mistaken as distinct species. The expression of
sexual dimorphism follows the same rule as in other
studied species of the genus, i.e., larger adults are the
most strongly dimorphic and smaller ones are the least
(Fig. 2).
Tityus ebanoverde Armas, 1999
Figures 3, 7
Type Data. DOMINICAN REPUBLIC, Cordillera
Central [Central Range], La Vega Province, Constanza
Municipality, El Arroyazo, Reserva Científica "Ébano
Verde", 1,100 m a.s.l., under box in house,
27/March/1999, R. Escalante, L. F. de Armas, ♂
holotype (IES: CZACC-3.2869, examined).
Remarks. Without having studied the holotype,
Teruel (2005) concluded that this species was most
likely not valid because all of the characters utilized as
diagnostic by Armas (1999) matched exactly those
which hypothetically must possess the yet unknown
small adult male of Tityus elii Armas et Marcano
Fondeur, 1992. Moreover, Teruel (2005) argued that he
visited the type locality of T. ebanoverde and noticed
that its ecological conditions were so homogeneous that
it could hardly support the existence of three syntopic
species of Tityus, as T. elii and T. ottenwalderi were
confirmed to occur there as well.

Euscorpius — 2017, No. 242
4
Figure 3: Size-related morphological variability of adult Tityus elii, exemplified by males: a) small preserved specimen from
Ébano Verde, La Vega Province (holotype of T. ebanoverde n. syn.); b) standard live specimen from road Bonao-Casabito, La
Vega Province (intermediate); c) large live specimen from Villa Las Neblinas, La Vega Province (typical T. elii; photo courtesy
František Kovařík); d) two selected morphometric ratios of the same three specimens, showing that robustness of pedipalp manus
and metasomal segment V increases with size (expressed as length of carapace and metasoma). Scale bar and graph axes in
millimeters.
The direct comparison of the holotype of T.
ebanoverde to the types and abundant additional ma-
terial of T. elii (representing adults of three different
size-classes), confirmed that the holotype of T. ebano-
verde is indeed a small adult male of the latter. Thus, the
following synonymy is herein established: Tityus elii
Armas et Marcano Fondeur, 1992 = Tityus ebanoverde
Armas, 1999, new synonym.
The expression of sexual secondary dimorphism in
T. elii follows exactly the same rule as in other studied
species of the genus, i.e., larger adults are the most
strongly dimorphic and smaller ones are the least, while
standard males are intermediate between both extremes.
Small males of T. elii possess the pedipalp chela and
metasomal segment V only slightly swollen and more
strongly carinate, as well as pedipalp fingers with basal
lobe/notch combination weak to moderate. The expres-
sion of all these dimorphic characters increases pos-
itively correlated to size: pedipalp chelae and metasomal
segment V become progressively wider and deeper with
their carination weaker, and lobe/notch combination of
fingers becomes stronger (Fig. 3).
Tityus septentrionalis Armas et Abud Antun,
2004
Figures 4, 7
Type Data. DOMINICAN REPUBLIC, Cordillera
Septentrional [Northern Range], Duarte Province, Loma
Quita Espuela, 700 m a.s.l., feeding on centipede in
forest litter at night, 11/February/2002, E. Gutiérrez, ♀
holotype (IES: CZACC-3.3144, examined). Cordillera

Teruel: Taxonomic Corrections to Tityus in Hispaniola
5
Figure 4: Photographical comparison amongst the holotypes of Tityus portoplatensis (a) and Tityus septentrionalis n. syn. (c),
and a live adult topotype pair of the former (b: male left, female right; photo courtesy Fr. Alejandro Sánchez). Note the strong
sexual dimorphism that demonstrates the former is actually a male and not a female, as wrongly determined by Armas &
Marcano Fondeur (1992).
Septentrional [Northern Range], Espaillat Province,
Moca, Los 21, under rock, 31/January/1989, D. Lanti-
gua, ♀ paratype (IES, not examined).
Remarks. The direct comparison of the holotypes
and additional specimens (topotypes included) of T.
septentrionalis and T. portoplatensis (the only other
member of the "quisqueyanus" species-group known to
occur in this mountain range), revealed that both taxa are
conspecific. Thus, the following synonymy is herein
established: Tityus portoplatensis Armas et Marcano
Fondeur, 1992 = Tityus septentrionalis Armas et Abud
Antun, 2004, new synonym.
The present study disclosed that Armas & Marcano
Fondeur (1992) wrongly sexed the holotype of T.
portoplatensis as a female. It is actually a large adult
male, thus, all characters alleged by Armas & Marcano
Fondeur (1992) and Armas & Abud (2004) as diagnostic
merely reflect sexual dimorphism (Fig. 4).
The specimen from Los Haitises in Hato Mayor
Province, described by Armas (2005) as the adult male
of T. septentrionalis, is either incorrectly sexed or mis-
identified (the photographs included in the paper could
also refer to a small adult female). Unfortunately, the
present author has repeatedly requested it on loan from
IES collection, without success.
Tityus (Brazilotityus) hartkorni Lourenço, 2009
and Tityus azari Lourenço, 2013
Figure 5
Type Data. "Dominican Republic. Precise amber
mine not confirmed. Lower Oligocene to Upper Eocene"
[sic], juvenile ♂ holotype of T. hartkorni (private col-
lection of Joachim Hartkorn, not examined), juvenile ♂
holotype of T. azari (private collection of Jörg Wun-
derlich, not examined). Note: Lourenço (2009: 4; 2013:
4) identically declared imprecise geographic and
stratigraphic origins for both holotypes, as literally
quoted above.
Remarks. The holotypes of both species were not
available for study, but fortunately, their original des-
criptions include good-quality color photographs that
allow a satisfactory interpretation of the main diagnostic
characters. First, two of them warrant their membership
in the "crassimanus" species-group: pedipalp fingers
with 14–15 principal rows of denticles and pectines with
19–20 teeth. These counts are within the diagnostic
ranges given for this group by Teruel & Armas (2006)
and exclude them from the "quisqueyanus" species-
group as diagnosed by Armas & Abud Antun (2004):
11–15 principal rows of denticles and 19–20 pectinal
teeth, vs. 10–12 principal rows of denticles and 8–15
pectinal teeth, respectively. Moreover, the elongate-
slender habitus (especially pedipalps, legs and meta-
soma) of both T. hartkorni and T. azari holotypes is also
unambiguously diagnostic for the "crassimanus" species-
group, as opposite to the short-stocky habitus of the
"quisqueyanus" species-group (Fig 5).
As pointed out elsewhere (Kovařík et al., 2016a–c;
Lowe & Kovařík, 2016; Prendini, 2016) for other papers
authored by Wilson R. Lourenço, both original des-
criptions are flawed by false and/or incorrectly described
characters, plus drawings of an inadmissible, poor
quality and thus, must be discarded as unreliable except
for the photographs (when included). A perfect example
is found here: the main character used by Lourenço
(2009, 3013) to support these allegedly new taxa and
their infra-generic relationships resides in the pectinal
fulcra, being described as vestigial to obsolete in T.
hartkorni and absent in T. azari. Even on this basis

Euscorpius — 2017, No. 242
6
Figure 5: Photographical comparison amongst the holotypes of Tityus hartkorni n. syn. (a) and Tityus azari n. syn. (b), and the
typical representatives of the "crassimanus" and "quisqueyanus" species-groups: Tityus crassimanus (c) and Tityus quisqueyanus
(d). All four specimens are juvenile males of similar size: 16–19 mm total length including telson. Images a–b taken from
Lourenço (2009: fig. 7) and Lourenço (2013: fig. 1), respectively.
alone, Lourenço (2009) placed T. hartkorni in the strict-
ly Amazonian, controversial subgenus Tityus (Brazilo-
tityus), without giving any biogeographical argument to
support such an odd assignment. Nevertheless, when the
photographs published by Lourenço (2009: fig. 8; 2013:
fig. 2) are zoomed-in, the pectinal fulcra become clearly
visible as normally developed at least in the right-side
pecten of both T. hartkorni and T. azari holotypes.
The third fossil member of the genus known from
Hispaniola is Tityus geratus Santiago-Blay et Poinar,
1988, but it presents a very different situation. It was
described also from a juvenile male holotype of similar
size (apparently second-instars in all three cases), but as
opposite to both previous cases, it has precise geo-
graphic and stratigraphic origins (La Toca amber mine,
in Puerto Plata Province), and its description is correctly
written, accurately discussed and satisfactorily illus-
trated. Thus, the only issue that remains to be addressed
is its infra-generic assignment. Based upon the same
characters discussed in the previous paragraph (see
above), it is clearly another member of the
"crassimanus" species-group: pedipalp fingers with 12–
14 principal rows of denticles, pectines with 18–19
teeth, and elongate-slender habitus (Santiago-Blay &
Poinar, 1988).
Once the infra-generic position of these three taxa is
established, it becomes clear that they closely match
each other in all diagnostic characters such as size,
degree of attenuation, carination and intercarinal sculp-
ture of body and appendages, counts of principal den-
ticle rows on pedipalp fingers and pectinal teeth.
Therefore, the conclusion is obvious: all three holotypes
are conspecific and the valid name corresponds to the
oldest available synonym. Thus, the following synon-
ymies are herein established: Tityus geratus Santiago-
Blay et Poinar, 1988 = Tityus (Brazilotityus) hartkorni
Lourenço, 2009, new synonym = Tityus azari Lourenço,
2013, new synonym.
It is worth to mention here that this discovery does
not represent an isolate case. Recently, the amber-fossil

Teruel: Taxonomic Corrections to Tityus in Hispaniola
7
Figure 6: Map of Hispaniola, showing the updated geographical distribution of all valid extant members of the "crassimanus"
species-group: Tityus crassimanus (circles), Tityus ottenwalderi (squares) and Tityus sp. (triangles). New records depicted in
white symbols, previous records in black.
arachnid fauna of Hispaniola has been shown to be
actually overestimated and abundant synonymies have
been demonstrated after detailed studies, see for exam-
ple Penney (2001).
General Remarks
After these changes, the diversity and distribution
patterns of the genus Tityus in Hispaniola are set as
follow. First, the "quisqueyanus" species-group is typ-
ical from and well diversified in the Central Range with
six species (T. abudi, T. altithronus, T. bellulus, T. elii,
T. kindli, and T. quisqueyanus), but also has single-spe-
cies' peripheral occurrences in the Neiba (T. neibae) and
Northern Ranges (T. portoplatensis). On the other hand,
the "crassimanus" species-group has a single allopatric
species in every major mountain range: T. crassimanus
in Bahoruco Range, T. ottenwalderi in the Central
Range, and one still-undetermined species in the Nor-
thern Range (Armas & Teruel, 2006; R. Teruel,
unpublished data).
In addition, the data personally gathered in the field
by the present author revealed that their ecological
preferences also differ markedly. All members of the
"crassimanus" species-group are microhabitat-specific,
but habitat-generalist: they are always strictly arboreal,
but widespread across all altitude ranges and vegetation
types. As opposite, the members of the "quisqueyanus"
species-group are microhabitat-generalist, but habitat-
specific: they occur indistinctly in the ground and trees,
but only in particular vegetation types and altitude
zones, i.e., T. abudi, T. elii, T. neibae, and T. porto-
platensis live in lower, humid broadleaf forests (350–
2,400 m a.s.l.), while T. altithronus, T. bellulus, T. kind-
li, and T. quisqueyanus occupy mostly higher, drier pine
forests (1,800–3,185 m a.s.l.).
The Hispaniolan amber-fossil members of the genus
have now decreased to a single species, which also
belongs in the "crassimanus" species-group. This is also
consistent with its chorology, i.e., this group is wide-
spread in most main mountain ranges of the island, but
composed of highly microhabitat-specialized taxa; such
combination suggests a very ancient origin.
The present corrections set the list of extant and
fossil Tityus species from this Greater Antillean insular
territory to 10 and 1, respectively, plus another extant
taxon that remains undetermined due to unavailability of
adequate samples. It follows below, with the updated
distribution of every species in each country (HA =
Haiti, DR = Dominican Republic) and their upper-level
political divisions (Départements in Haiti, Provinces in
Dominican Republic); new records were marked in
boldface and tentative records in question mark:
• "crassimanus" species-group
1. Tityus crassimanus Armas, 1999: HA (Sud Est),
RD (Pedernales, Independencia, Barahona).
2. Tityus ottenwalderi Armas, 1999: RD (Elías
Piña, Santiago, La Vega, Monseñor Nouel, San
Cristóbal).

Euscorpius — 2017, No. 242
8
Figure 7: Map of Hispaniola, showing the updated geographical distribution of two valid extant members of the "quisqueyanus"
species-group: Tityus elii (squares) and Tityus portoplatensis (triangles). New records depicted in white symbols, previous
records in black.
3. Tityus geratus Santiago-Blay et Poinar, 1988†:
RD (fossil in amber, confirmed only from Puerto
Plata).
4. Tityus sp.: RD (Puerto Plata, Espaillat, Hato
Mayor).
• "quisqueyanus" species-group
5. Tityus abudi Armas, 1999: RD (Santiago).
6. Tityus altithronus Armas, 1999: RD (San Juan,
Santiago).
7. Tityus bellulus Armas, 1999: RD (Santiago).
8. Tityus elii Armas et Marcano Fondeur, 1992: RD
(Santiago, La Vega, San Cristóbal).
9. Tityus kindli Kovařík et Teruel, 2014: RD (Azua,
La Vega).
10. Tityus neibae Armas, 1999: RD (Independencia,
Bahoruco, Elías Piña).
11. Tityus portoplatensis Armas et Abud Antun,
1992: RD (Puerto Plata, Duarte, Espaillat, Hato
Mayor?).
12. Tityus quisqueyanus Armas, 1982: RD (Santi-
ago, La Vega).
Last, it is worth noting here that this list is not yet
definitive. A forthcoming paper by the present author
and his collaborators (R. Teruel, F. Kovařík, M. Seiter,
and G. de los Santos, in preparation), will revise the
genus in detail for Hispaniola, and bring more changes:
it will include the description of at least one new species
from Haiti, the update of geographical distribution and
ecological information of most taxa, and the correction
of more errors discovered in the published literature.
Acknowledgments
I am greatly indebted to those friends and col-
leagues that have provided me with specimens and
literature for years and thus, made this contribution pos-
sible. I especially thank František Kovařík (Prague,
Czech Republic), Michael Seiter (Pottendorf, Austria),
Jan Ove Rein (Trondheim, Norway), Gabriel de los
Santos (Museo de Historia Natural "Eugenio de Jesús
Marcano Fondeur", Santo Domingo, Dominican Repub-
lic), Antonio A. Melic (Sociedad Entomológica Ara-
gonesa, Spain), Alexander Sánchez and Leopoldo Viña
(formerly at Bioeco). Two of them (F. Kovařík and M.
Seiter) have also made possible the field trips to
Dominican Republic and have given their valuable and
kind assistance to collect specimens, together with other
friends: Pavel Kindl (Czech Republic), Johannes Nigl
(Austria), Frederic Schramm (Germany), Abraham Abud
and Diego Rivera (Santo Domingo, Dominican Repub-
lic). In addition, to the staff of the Instituto de Ecología y
Sistemática (Havana, Cuba) for allowing access to the
type-specimens deposited in this public collection, as
well as for positively answering to most of the loan
requests for other material. Thanks also to F. Kovařík
(again) and Fr. Alejandro Sánchez (Carolina, Puerto
Rico), for the high-resolution photographs kindly sup-
plied for present Figs. 3c and 4b, respectively. More-

Teruel: Taxonomic Corrections to Tityus in Hispaniola
9
over, to Sheyla Yong (Havana, Cuba) for hand-carrying
most of the loans from IES to Bioeco and her helpful
comments to an earlier draft of the text. Last, but not
least, to two anonymous referees for the peer-review of
the manuscript.
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