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BULLETIN OF THE AMERICAN MUSEUM OF NATURAL HISTORY
SYSTE M AT I C REVISION OF THE ARBOREAL
NEOTROPICAL “THORELLII” CLADE O F
CENTRUROIDES MARX, 1890, BARK SCORPIONS
(BUTHIDAE C.L. KOCH, 1837) WITH
DESCRIPTIONS OF SIX NEW SPECIES
AARON M. GOODMAN, LORENZO PRENDINI,
OSCAR F. FRANCKE AND LAUREN A. ESPOSITO
BULLETIN OF THE AMERICAN MUSEUM OF NATURAL HISTORY
Number 452, 92 pp., 43 gures, 10 tables
Issued September 16, 2021
Copyright © American Museum of Natural History 2021 ISSN 0003-0090
SYSTEMATIC REVISION OF THE ARBOREAL
NEOTROPICAL “THORELLII” CLADE OF
CENTRUROIDES MARX, 1890, BARK SCORPIONS
(BUTHIDAE C.L. KOCH, 1837) WITH
DESCRIPTIONS OF SIX NEW SPECIES
AARON M. GOODMAN
Graduate School and University Center, City University of New York;
Division of Invertebrate Zoology, American Museum of Natural History;
Institute for Biodiversity Science and Sustainability, California Academy of Sciences
LORENZO PRENDINI
Arachnology Lab and Scorpion Systematics Research Group,
Division of Invertebrate Zoology, American Museum of Natural History
OSCAR F. FRANCKE
Colección Nacional de Arácnidos, Departamento de Zoología,
Instituto de Biología, Universidad Nacional Autónoma de México
LAUREN A. ESPOSITO
Graduate School and University Center, City University of New York;
Division of Invertebrate Zoology, American Museum of Natural History;
Institute for Biodiversity Science and Sustainability, California Academy of Sciences
2
CONTENTS
Abstract ............................................................................3
Introduction.........................................................................3
Materials and Methods ...............................................................4
Systematics ..........................................................................6
Family Buthidae C.L. Koch, 1837 ....................................................6
Subfamily Centruroidinae Kraus, 1955 ............................................6
Centruroides Marx, 1890 ......................................................6
Key to Identication of the Species of the “thorellii” clade of Centruroides Marx, 1890 .. 7
Centruroides berstoni, sp. nov................................................8
Centruroides catemacoensis, sp. nov. .........................................15
Centruroides chanae, sp. nov. ...............................................22
Centruroides cuauhmapan, sp. nov ..........................................33
Centruroides hamadryas, sp. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .41
Centruroides homanni Armas, 1996 ........................................53
Centruroides rileyi Sissom, 1995 ............................................56
Centruroides schmidti Sissom, 1995..........................................61
Centruroides yucatanensis, sp. nov...........................................70
Acknowledgments...................................................................76
References..........................................................................77
Appendix 1.........................................................................84
Appendix 2.........................................................................86
Appendix 3.........................................................................91
3
ABSTRACT
e arboreal Neotropical “thorellii” clade of Centruroides Marx, 1890, bark scorpions (Buthidae
C.L. Koch, 1837) is revised, using a novel approach to species delimitation. A phylogenetic analysis,
based on 112 morphological characters and 1078 aligned DNA nucleotides from the mitochondrial
Cytochrome c Oxidase Subunit I (COI) gene, provided the framework for placing singletons from
geographically disparate localities (and oen with suboptimal preservation) using COI minibar-
codes, thereby enlarging the taxon sample for diagnosis and delimitation of morphological species.
Six new species are described, tripling the known diversity in the clade to nine: Centruroides berstoni,
sp. nov.; Centruroides catemacoensis, sp. nov.; Centruroides chanae, sp. nov.; Centruroides cuauhma-
pan, sp. nov.; Centruroides hamadryas, sp. nov.; Centruroides yucatanensis, sp. nov. Revised diagnoses
are presented for Centruroides homanni Armas, 1996, Centruroides rileyi Sissom, 1995, and Cen-
truroides schmidti Sissom, 1995. Comparative images, a key and distribution maps for all species of
the clade are provided, along with a summary of available data for their ecology.
INTRODUCTION
The Yucatan/Chortis “thorellii” clade is
one of three clades recently identified in the
diverse Neotropical bark scorpion genus Cen-
truroides Marx, 1890 (Buthidae C.L. Koch,
1837), based on a phylogenetic analysis of
morphological characters and DNA sequences
from two nuclear and three mitochondrial
gene loci (Esposito and Prendini, 2019). All
species of the clade are arboreal and cortic-
olous (Prendini, 2001a), some having been
collected 3–15 m above ground in the forest
canopy (Goodman and Esposito, 2020; fig. 1).
These cryptic scorpions are characterized by
relatively small size, mottled coloration and
an elongated metasoma and telson, more pro-
nounced in the adult male.
ree species of the clade, i.e., Centruroides
homanni Armas, 1996, Centruroides rileyi Sis-
som, 1995, and Centruroides schmidti Sissom,
1995, were originally assigned together with
Centruroides thorellii (Kraepelin, 1891) and
three other species, Centruroides chamulaensis
Homann, 1932, Centruroides tuxtla Armas,
1996, and Centruroides sissomi Armas, 1996, to
the informal thorellii species group of Centru-
roides based on their similar morphology, habi-
tat and distribution in the forests of southern
Mexico and northern Central America (Armas,
1996; Homann, 1932; Martin-Frías et al., 2005;
Sissom, 1995, Ponce-Saavedra and Moreno-
Barajas, 2005). Esposito and Prendini (2019)
demonstrated that the thorellii species group
was a paraphyletic assemblage, however. Mem-
bers of the former thorellii group dissociated
into three clades, in two cases grouping with
morphologically disparate species, suggesting
the apparent morphological similarity among
species of the thorellii group is convergent. Cen-
truroides tuxtla grouped with a clade of pre-
dominantly striped species, including the type
species of Centruroides, Centruroides exilicauda
Wood, 1863, whereas C. chamulaensis and C.
“thorellii” grouped with a clade of large, dark
species, e.g., Centruroides gracilis Latreille,
1804. e remaining species of the thorellii
group formed a distinct clade.
The limits of several species formerly
assigned to the thorellii group of Centruroides
are similarly confused. For example, the rede-
scription of C. homanni by Martin-Frías et al.
(2005) appears to be based on specimens that
are not conspecic with the types but rather
with a sympatric species of Centruroides
(Esposito, 2011), whereas most reports of C. sis-
somi that appeared aer the original description
(Vázquez, 1999; Teruel et al., 2015a; Ponce-Saa-
vedra and Francke, 2019) are misidentications
of a hitherto undescribed, sympatric species of
the “thorellii” clade, described herein.
Problems concerning the taxonomy of these
scorpions may, in part, be explained by the com-
4 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 452
parative scarcity of material in collections. Many
collection records are represented by singletons,
oen females or immatures, and the diculty of
collecting these canopy-dwelling scorpions
appears to have resulted in severe undersampling
of their known diversity and distribution.
Six operational taxonomic units (OTUs)
were previously identied within the “thorellii”
clade on the basis of DNA sequence data
(Esposito, 2011). However, insucient material
prevented a decision as to whether the OTUs
were distinct species or variable populations of
more widespread species. A larger sample of
specimens from across the geographical distri-
bution, needed for a robust assessment of spe-
cies limits within the genus, was hampered by
degraded genetic material within museum
specimens. In order to overcome this limita-
tion, a novel approach to species delimitation
was applied by Goodman et al. (2021). A phy-
logenetic analysis, based on 112 morphological
characters and 1078 aligned DNA nucleotides
from the mitochondrial cytochrome c oxidase
subunit I (COI) gene, provided the framework
for placing singletons from geographically dis-
parate localities (and oen suboptimally pre-
served) using COI minibarcodes, thereby
enlarging the taxon sample for adequate diag-
nosis and delimitation of morphological
species.
e present contribution implements the tax-
onomic discoveries of Goodman et al. (2021).
e six OTUs originally identied by Esposito
(2011) are described as new species, tripling the
known diversity in the clade to nine (table 1):
Centruroides berstoni, sp. nov.; Centruroides
catemacoensis, sp. nov.; Centruroides chanae, sp.
nov.; Centruroides cuauhmapan, sp. nov.; Centru-
roides hamadryas, sp. nov.; Centruroides yucata-
nensis, sp. nov. Revised diagnoses are presented
for Centruroides homanni Armas, 1996, Centru-
roides rileyi Sissom, 1995, and Centruroides
schmidti Sissom, 1995. Comparative images, a
key and distribution maps for all species of the
clade are provided, along with a summary of
available data for their ecology.
MATERIAL AND METHODS
Specimens collected by the authors were
located at night using ultraviolet (UV) light detec-
tion, preserved by submersion in 95% ethanol and
subsequently injected with ethanol to improve
internal preservation. Additional material, in most
cases singletons, was sourced from natural history
collections. Material is deposited in the American
Museum of Natural History (AMNH), New York;
the California Academy of Sciences (CAS), San
Francisco; the Colección Nacional de Arácnidos
(CNAN), Instituto de Biología, Universidad
Nacional Autónoma de México, Mexico City; the
Field Museum of Natural History (FMNH), Chi-
cago; the Florida State Collection of Arthropods
(FSCA), Gainesville; Northern Arizona University
(NAU), Flagsta; the Oxford University Museum
of Natural History (OUMNH), U.K.; and the
United States National Museum of Natural His-
tory (USNM), Smithsonian Institution, Washing-
ton, DC. Tissue samples used for DNA isolation
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, and the Center for Comparative Genom-
ics at the CAS (appendix 1).
Specimen provenance data from the material
examined were digitized. All records of sucient
accuracy were isolated from the material exam-
ined and published literature to create a point-
locality geographical dataset for mapping
distributional ranges. Distribution maps were
produced using ArcView GIS Version 10.4 (Envi-
ronmental Systems Research Institute, Redlands,
CA). Raster-based elevational maps, and vector-
based political boundaries of countries were
acquired from DIVA-GIS v1.4.
Morphological examination of specimens was
conduced using a Leica M125 stereomicroscope.
Measurements (in millimeters) were recorded using
an ocular micrometer calibrated at 10× magnica-
tion. Measurements follow Stahnke (1970), Lam-
oral (1979), and Prendini (2001b). Nomenclature of
general anatomy follows Hjelle (1990) and Sissom
(1990), except for carination of the carapace, ter-
2021 GOODMAN ET AL.: REVISION OF “THORELLII” CLADE OF CENTRUROIDES 5
gites and metasoma, which follows Vachon (1952),
trichobothria, which follows Vachon (1974), pedi-
palp carination, which follows Prendini (2000a),
book lung structure, which follows Kamenz and
Prendini (2008), and ovariuterine anatomy, which
follows Volschenk et al. (2008).
Photographs were taken in visible light using
a Canon EOS Camera with MP-E 65 mm or 100
mm EF macrolenses, and under long wave UV
light using a MicropticsTM ML-1000 digital imag-
ing system, or Leica SMZ-800 stereomicroscope
equipped with a Leica MZ16 A camera, and
LED-6WD UV spotlight. Focal planes of image
stacks were fused using Leica LAS image stack-
ing program and edited with Adobe Photoshop.
Genomic DNA was extracted from muscle tis-
sue from the fourth leg of the best preserved
specimens using the spin column extraction pro-
tocol of the Qiagen DNeasy Blood and Tissue Kit
(Valencia, CA). PCR amplication of complete
mitochondrial and nuclear genes proved impos-
sible for many older museum samples, due to
degradation and fragmentation of the DNA.
erefore, a 125 base-pair (bp) hypervariable
region of the cytochrome c oxidase subunit I
(COI) gene was partially amplied (Meusnier et
al., 2008). Newly generated sequences were
edited, forward and reverse primers removed,
and complementary strands assembled into con-
sensus sequences using Geneious v. 11.0.4
(Kearse et al., 2012), by reference to a 1078 bp
fragment of COI for C. rileyi from GenBank
(Esposito and Prendini, 2019). Sequences less
than 150 base pairs in length were deposited in
the Dryad digital repository (doi: 10.5061/dryad.
tdz08t2), the rest in GenBank (appendix 1).
Adults of specimens from which DNA
sequences were generated, were scored using rel-
evant characters from an unpublished morpho-
logical character matrix by Esposito (2011),
comprising characters from published (Esposito
et al., 2017) and unpublished sources (appendix
2). e data matrix comprises 112 characters
comprising 43 (35%) from the prosoma, 38
(33%) from the mesosoma, and 30 (29%) from
the metasoma; 41% of the characters were
derived from carination and surface macrosculp-
ture, 25% from shape and morphometrics, 24%
from coloration, and 7% from other character
systems, e.g., macrosetae, trichobothria, and
internal anatomy (appendix 3). e matrix is
deposited in Morphobank (http://morphobank.
org/permalink/?P4047).
Sequences were aligned using the ClustalW
method (Larkin et al., 2007; ompson et al.,
1994) in Mesquite v. 3.51 (Maddison and Mad-
dison, 2018), and checked by eye. Evolutionary
relationships were inferred with a simultane-
TABLE 1
Species of the arboreal Neotropical scorpion “thorellii” clade of Centruroides Marx, 1890, bark scorpions,
with countries, and departments or states in which recorded
Species Country State/Department
Centruroides berstoni, sp. nov. Guatemala Izabal
Centruroides catemacoensis, sp. nov. Mexico Veracruz
Centruroides chanae, sp. nov. Mexico Guerrero, Michoacán
Centruroides cuauhmapan, sp. nov. Mexico Oaxaca, Veracruz
Centruroides hamadryas, sp. nov. Mexico Chiapas
Centruroides homanni Armas, 1996 Mexico Chiapas
Centruroides rileyi Sissom, 1996 Mexico Puebla, Tamaulipas, San Luis Potosí, Veracruz
Centruroides schmidti Sissom, 1995 Guatemala El Progreso, Zacapa
Honduras Atlántida, Cortés, Francisco Morazán, Islas del Bahía
Centruroides yucatanensis, sp. nov. Mexico Quintana Roo, Yucatán
6 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 452
ous phylogenetic analysis of the molecular and
morphological datasets using Bayesian inference
(BI) in MrBayes v. 4.3.6 (Ronquist and Huelsen-
beck, 2003) and maximum likelihood (ML) using
RAxML v. 8.0.0 (Stamatakis, 2014) on the CIPRES
supercomputing cluster (Miller et al., 2009).
e limits of putative species were evaluated
on the majority-rule consensus tree from the
Bayesian 95% posterior probability of the concat-
enated dataset of morphological characters and
aligned DNA sequences using the species delimi-
tation plugin in Geneious (Kearse et al., 2012;
Masters et al., 2011). Clusters of sequences were
evaluated based on their association with indi-
viduals collected at the type locality. Specimens
that were monophyletic with those collected at
the type locality or within its vicinity with greater
than 0.95 posterior probability were considered
conspecic and assigned to the species in ques-
tion. Several metrics were investigated for assess-
ing species delimitation in Geneious, including
P (AB) for reciprocal monophyly (Rosenberg,
2007), P (RD) which measures the probability
of an observed clade’s degree of distinctiveness
(Rodrigo et al., 2008), and values for the prob-
ability of population identication of a hypothet-
ical sample based on the groups being tested, P
ID (Strict), and P ID (Liberal).
Phylogenetic analysis with BI and ML pro-
duced almost identical tree topologies with
terminal nodes well supported, but basal and
internal nodes weakly supported. Nine well-
supported, reciprocally monophyletic species-
level clades were recovered in both analyses,
but relationships within each clade received low
support. Centruroides homanni, C. rileyi, and C.
schmidti were each monophyletic and received
high posterior probabilities and likelihood val-
ues (0.99/100, 0.95/1.00, and 0.99/100 respec-
tively). Centruroides thorellii and C. tuxtla were
not monophyletic with the rest of the “thorellii”
clade (g. 2), conrming the results of Esposito
and Prendini (2019). The six new species
described herein were monophyletic and well
supported: C. berstoni (0.99/99); C. catemacoen-
sis (1.00/100); C. chanae (1.00/100); C. cuauh-
mapan (0.99/100), C. hamadryas (0.98/82); C.
yucatanensis (1.00/100). Species-delimitation
analysis identied nine distinct species, all of
which received values for Rosenberg’s P (AB) less
than 0.01, supporting the results of the BI and
ML analyses. A more detailed explanation of the
methods and results of the phylogenetic analyses
is provided by Goodman et al. (2021).
SYSTEMATICS
Family Buthidae C.L. Koch, 1837
Subfamily Centruroidinae Kraus, 1955
Centruroides Marx, 1890
Figures 1–43; tables 1–10
Centrurus (nec Ehrenberg, 1829): orell, 1876a:
9; 1876b: 83; Karsch, 1879a: 18; Pocock,
1890: 120, 121, 127; Kraepelin, 1891: 119–
124 (part); Pocock, 1893: 375, 385, 386;
Laurie, 1896: 131; Lönnberg, 1897: 196, 197,
208; Kraepelin, 1899: 87 (part); Banks, 1900:
425; Borelli, 1909: 222; Comstock, 1912: 25,
27, g. 31; Birula, 1917a: 164; 1917b: 54,
107; Ochoterena, 1920: 223; Mello-Campos,
1924a: 246; 1924b: 312; Comstock, 1940: 27,
g. 31 (lapsus calami); Millot and Vachon,
1949: 427; Díaz Nájera, 1966: 110, 111, pl. 1;
1970: 113.
Centruro: Karsch, 1879b: 120 (lapsus calami).
Centruroides Marx, 1890: 211, type species Buthus
exilicauda Wood, 1863 (= Centruroides exili-
cauda (Wood, 1863), by monotypy; Pocock,
1902a: 19, 20; 1902b: 365; Kraepelin, 1912:
69–71; 1914: 22; Homann, 1932: 244, 245;
Mello-Leitão, 1932: 27; 1934: 4, 5; Franganillo,
1936: 158; Homann, 1937: 201–203; Moreno,
1939: 63; Comstock, 1940: 25, 27; Moreno,
1940: 164; Mello-Leitão, 1942: 126; 1945: 240,
250–252; Scorza, 1954:190; Bücherl, 1964: 59;
1967: 113; Muma, 1967: 2, 4; Bücherl, 1969:
767; Aguilar and Meneses, 1970: 3; Bücherl,
1971: 327; Stahnke, 1971: 282; 1972: 125, g.
9; Armas, 1974a: 25; Vachon, 1974: 906, 908;
2021 GOODMAN ET AL.: REVISION OF “THORELLII” CLADE OF CENTRUROIDES 7
1975: 1598; Francke, 1977: 75; Stahnke and
Calos, 1977: 111; Vachon, 1977: 294; Lou-
renço, 1979: 214; Williams, 1980: 2, 4; Armas,
1984: 2; González-Sponga, 1984: 64, 65;
Francke, 1985: 7, 15; Francke and Stockwell,
1987: 7; Armas, 1988: 44, 91, 95; Stockwell,
1988: 3; Sissom, 1990: 94, 101; Nenilin and
Fet, 1992: 9, 12–14; Stockwell, 1992: 412, 419;
González-Sponga, 1996: 118, 119, 124, 125,
gs. 285–287, 289, 292; Armas, 1998: 50;
Kovařík, 1998: 106; Fet and Lowe, 2000:
98–122; Towler et al., 2001: 161–163; Armas et
al., 2004: 167–171, tables 1–3; Prendini and
Wheeler, 2005: 481, table 10; Kamenz and
Prendini, 2008: 6, 8, 22, 40, tables 1, 2, pl.
13–17; Volschenk et al., 2008: 654, 656, 658,
659, 663, 664, 674, g. 1C, tables 1, 2; Armas
et al., 2012: 106, 112; Cupitra Vergara et al.,
2014: 207–215, gs. 1–4, tables 1, 2; Kovařík
and Teruel, 2014: 1, 7, 15, 25, 26; Loria and
Prendini, 2014: 3, 9, 10, 24, 25, g. 2D, table 5;
Lourenço, 2014: 60, 63; Miller et al., 2014: 301,
306, 307, table 1; Ponce-Saavedra and Francke,
2014: 54, gs. 11, 14; Quintero Arias and
Esposito, 2014: 373–382, gs. 1–6, table 1;
Teruel and Rodríguez-Cabrera, 2014: 131;
Armas and Ávila, 2015: 66, 67, 69, 70, 71;
Ponce-Saavedra et al., 2015: 81–89, gs. 1–5,
tables 1, 2; Quijano-Ravell and Ponce-Saave-
dra, 2015: 35–44, gs. 1–5, tables 1–3;
Riquelme et al., 2015: 2, 16, 18, 19; Teruel et
al., 2015a: 3–14, gs. 1–37, table 1; 2015b:
1–18, gs. 1–35, tables 1–5; 2015c: 13–33, gs.
1–75, tables 1–4; Baldazo-Monsivaiz et al.,
2016: 76–79, g. 2A, table 1; de los Santos et
al., 2016: 1–7, 14, 16–19, 22, gs. 2A–G, 3A,
table 1; Kovařík et al., 2016a: 1, 11, 18, 19;
Lourenço and Velten, 2016: 61, 65; Quijano-
Ravell and Ponce-Saavedra, 2016a: 49–61, gs.
1–9, tables 1–4; 2016b: 85–90, gs. 1–4; Ter-
uel, 2016: 91–93, gs. 1–3; Baldazo-Monsivaiz,
2017: 21–27, g. 2A; Esposito et al., 2017:
2–15, 29, 30, 35–37, 41, 61, 71, 81, 85, 101,
116, 122, 124, 126, 128–130, g. 13; McWest et
al., 2015: 3, 9, 10, 25, 28, 30–34, 38, gs. 2A,
3A, 4A; Teruel and Myers, 2017: 1–14, gs.
1–11, table 1; Teruel et al., 2017: 6; Ubinski et
al., 2018: 475–486, gs. 1–8, tables 1, 2;
Esposito and Prendini, 2019: 1–10, gs. 1, 2, 3,
tables 1–4; Crews and Esposito, 2020: 8, 9, 12,
17, 18; Fet and Kovařík, 2020: 8, 10, 12, 14, 18,
19, 22–26, 28, 29, 31; Howard et al., 2019: 74,
84, g. 3, table 1; Miranda and Armas, 2020:
5, 7; Ponce-Saavedra and Francke, 2019: 1–17,
gs. 1–11, tables 1–4; Quijano-Ravell et al.,
2019: 31–48, gs. 1–32, tables 1–5; Teruel and
Questel, 2020: 12, 14; Trujillo and Armas,
2020: 1–8, gs. 1–4, table 1; Wendru et al.,
2020: 2, 5, 6, g. 2B.
Centruroides (Centruroides): Werner, 1934: 273.
Centruoroides: Díaz Nájera, 1970: 117 (lapsus
calami).
Centruriodes: Lourenço and Eickstedt, 1988: 7
(lapsus calami).
Key to Identication of the Species of the
“thorellii” Clade of Centruroides Marx, 1890
1. Carapace posterosubmedian carinae ............2
– Carapace posterosubmedian carinae absent ...6
2. Pedipalp chela manus, retrodorsal carina
smooth, dorsomedian carina weakly granu-
lar; mesosomal sternite VII, ventrolateral
and ventrosubmedian carinae obsolete to
absent; metasomal segments I–III, ventro-
submedian and ventrolateral carinae vesti-
gial, smooth (gs. 18, 19D); telson vesicle
surfaces smooth (♀) (gs. 23–25D).............
....................Centruroides rileyi Sissom, 1995
– Pedipalp chela manus, retrodorsal carinae
present and dorsomedian carinae present or
restricted to proximal third or half; meso-
somal sternite VII, ventrolateral and ventro-
submedian carinae distinct; metasomal
segments I–III, ventrolateral and ventrosub-
median carinae complete, granular (gs. 18L,
R, 19L, R); telson vesicle surfaces granular
(♀) (gs. 23L, R, 24L, R, 25L, R) .....................3
8 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 452
3. Pedipalp chela manus, prodorsal carina
absent.......................................4
– Pedipalp chela manus, prodorsal carina
present....................................5
4. Pectinal tooth counts, 15–17 (♂), 14 (♀)
(table 9); mesosomal tergites I–VII, dorso-
median carinae present on I–III, absent on
IV–VII; mesosomal sternite VII, ventrolat-
eral and ventrosubmedian carinae distinct;
metasoma length greater than 3× meso-
soma length; metasomal segments I–IV,
ventrolateral and ventrosubmedian
carinae finely granular on segments I–III
(figs. 18I, L, 19I, L), smooth on IV.......
....................Centruroides chanae, sp. nov.
– Pectinal tooth counts, 14 (♂), 12–14 (♀)
(table 10); mesosomal tergites I–VII, dor-
somedian carinae absent on I and II, ves-
tigial on III–VI, absent on VII; mesosomal
sternite VII, ventrolateral and ventrosub-
median carinae weakly developed; meta-
soma length less than 3× mesosoma
length; metasomal segments I–IV, ventro-
lateral and ventrosubmedian carinae pres-
ent on segments I–IV (figs. 18O, R, 19O,
R).......Centruroides yucatanensis, sp. nov.
5. Mesosomal tergites I–VII, dorsomedian
carinae distinct, complete; total length (♂,
♀), 44–60 mm (table 8); pectinal tooth
counts, 13–15 (♂), 13 (♀)...............................
...Centruroides hoffmanni Armas, 1996
– Mesosomal tergites I–VII, dorsomedian
carinae restricted to posterior two-thirds of
segments I–VI, absent on VII; total length
(♂, ♀), 38–52 mm (table 7); pectinal tooth
counts, 14–16 (♂), 12–14 (♀).......................
..............Centruroides schmidti Sissom, 1995
6. Pedipalp chela manus, prodorsal carina
complete, granular......................................
.......Centruroides cuauhmapan, sp. nov.
– Pedipalp chela manus, prodorsal carina
absent........................................7
7. Pedipalp chela manus, retrodorsal carina
complete, granular; mesosomal sternite VII,
ventrolateral and ventrosubmedian carinae
obsolete to absent (gs. 13, 14A); telson ves-
icle surfaces (♀) smooth (gs. 23–25E)......
..................Centruroides hamadryas, sp. nov.
– Pedipalp chela manus, retrodorsal carina weakly
granular; mesosomal sternite VII, ventrolateral
and ventrosubmedian carinae weakly devel-
oped; telson vesicle surfaces (♀) granular.......8
8. Total length, 39–51 mm (table 6); pedipalp
chela manus, dorsomedian carina weakly
granular, restricted to distal half; pectinal
tooth counts, 13–14 (♂, ♀); mesosomal ter-
gites I–VII, dorsomedian carinae vestigial;
metasoma length less than 3.2× greater than
mesosoma length; metasomal segments I–
IV, ventrolateral and ventrosubmedian cari-
nae absent (gs. 18M, P, 19M, P)................
.............Centruroides catemacoensis, sp. nov.
– Total length, 48–55 mm (table 5); pedipalp
chela manus, dorsomedian carina absent;
pectinal tooth counts, 14–16 (♂), 13–15
(♀); mesosomal tergites I–VII, dorsomedian
carinae weakly granular, restricted to poste-
rior half; metasoma length greater than 3.2×
mesosoma length; metasomal segments I–
IV, ventrolateral and ventrosubmedian cari-
nae present (gs. 18H, K, 19H, K)..........
........................Centruroides berstoni, sp. nov.
Centruroides berstoni, sp. n ov.
Figures 1B, D, 2, 4, 6C, D, 9C, D, 13B, 14B, 17H,
K, 18H, K, 19H, K, 20H, K, 21H, K, 22H, K,
23H, K, 24H, K, 25H, K, 34, 35, tables 1, 5, 10
Centruroides schmidti: Sissom, 1995: 94–96; gs.
10–18, table 1 (misidentication: paratypes
from Escobas, Izabal, Guatemala).
T M: GUATEMALA: Departa-
mento Izabal: Município Livingston: Holotype ♂
(CASENT 9073325), paratype ♂ (CASENT
9073298), Río Dulce, Hotel Tijax, 15°39′51.2″N
89°00′14.6″W, 17 m, 24.ix.2019, A.M. Goodman,
collected along gravel road of Hacienda Tijax
Parking Lot, anked by bamboo groves, and live
fencing; 2 ♂ paratypes (CASENT 9073312,
9073326), 4 ♀ paratypes (CASENT 9073297,
9073313, 9073324, 9073368), Biotopo Chocón
2021 GOODMAN ET AL.: REVISION OF “THORELLII” CLADE OF CENTRUROIDES 9
FIGURE 1. Representative species of the arboreal Neotropical “thorellii” clade of Centruroides Marx, 1890,
bark scorpions (Buthidae C.L. Koch, 1837), habitus in life. A, C. C. schmidti Sissom, 1995, A. ♀, El Progreso,
Finca El Olvido, Las Minas, Guatemala, C. ♂, Zacapa, Guatemala. B, D. C. berstoni, sp. nov., B. ♀, D. ♂,
Izabal, Rio Dulce, Hotel Tijax, Guatemala. Note elongated metasoma in males. Both species were collected in
vegetation, 3–4 m above ground level.
10 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 452
Machacas, 15°44′05.3″N 88°54′57.2″W, 15 m,
25.ix.2019, A.M. Goodman.
E: e species name is a patronym
honoring the late Hyman Maxwell Berston, the rst
author’s grandfather (1930–2021), who inspired his
interest in natural history during childhood visits
to the California Academy of Sciences.
D: Centruroides berstoni is most closely
related to C. hamadryas, from which it diers as
follows. e carapace is sparsely granular, more
densely so on the interocular triangle, in the female
of C. berstoni but densely granular, with distinct
lateral ocular carinae, in the female of C. hama-
dryas (g. 6B, D). e pedipalp chela manus of the
male is proportionally less incrassate in C. berstoni
(gs. 13–15B) than C. hamadryas (g. 13–15A,
tables 4, 5). e retrodorsal carina of the chela
manus is weakly granular and the dorsomedian
carina absent in the male of C. berstoni (g. 13B),
whereas the retrodorsal carina is complete and the
dorsomedian carina weakly developed and
restricted to the distal half in the male of C. hama-
dryas (g. 13A). e retrodorsal carina of the
manus is nely granular and the prodorsal carina
absent in the female of C. berstoni (g. 14B),
whereas the retrodorsal carina is complete and the
prodorsal carina restricted to the distal third in the
female of C. hamadryas (g. 14A). e rst leg of
the male is greater than 2× the length of the cara-
pace in C. berstoni but less than 2× its length in C.
hamadryas (table 10). e dorsomedian carinae of
the mesosomal tergites are weakly developed and
restricted to the posterior half in C. berstoni but
distinct and complete in C. hamadryas. e ventro-
lateral carinae of mesosomal sternite VII are weakly
developed and the ventrosubmedian carinae absent
FIGURE 2. Phylogeny of the arboreal Neotropical “thorellii” clade of Centruroides Marx, 1890, bark scorpions
(Buthidae C.L. Koch, 1837), obtained by simultaneous phylogenetic analysis of 112 morphological characters
and DNA sequences of the mitochondrial cytochrome oxidase c subunit I gene. Posterior probabilities,
obtained with Bayesian inference, and nonparametric bootstraps, obtained with maximum likelihood, indi-
cated above and below branches for corresponding nodes.
2021 GOODMAN ET AL.: REVISION OF “THORELLII” CLADE OF CENTRUROIDES 11
in C. berstoni, whereas the ventrolateral and ventro-
submedian carinae are weakly developed to absent
in C. hamadryas. e telson vesicle is sparsely
setose in C. berstoni (gs. 23H, K, 24H, K, 25H, K),
but densely setose in C. hamadryas (gs. 23B, E,
24B, E, 25B, E).
Additional dierences between C. berstoni and
other species of the clade are as follows. e ventro-
submedian and ventrolateral carinae of metasomal
segments I–IV in the male are weakly developed,
smooth in C. berstoni (gs. 18H, 19H) but absent in
C. catemacoensis (gs. 18, 19M), well developed in
C. cuauhmapan (gs. 18, 19G), weakly developed
on segments I–III, absent on IV in C. hamadryas
(gs. 18–19B, 21–22B), and vestigial, smooth in C.
rileyi (gs. 18, 19A, 21–22A). e metasomal cari-
nae of the female are nely granular in C. berstoni
(gs. 17–22K) but well developed, granular in C.
catemacoensis (gs. 17–22P). e telson vesicle is
short and not posteriorly bilobed in the male of C.
berstoni (gs. 23–25H), unlike in C. cuauhmapan
(gs. 23–25G) and C. rileyi (gs 23–25A), and with
surfaces sparsely granular in the female of C. ber-
stoni (gs. 23–25K), but smooth in the female of
C. hamadryas (gs. 23–25E) and densely granular
in the female of C. catemacoensis (gs. 23–25P),
C. cuauhmapan (gs. 23–25J), and C. rileyi (gs.
23–25D).
D: e following description is
based on the holotype male, with dierences
among other material noted in the section on
variation.
Coloration: Base color pale yellow, with exten-
sive infuscation, creating mottled or marbled pat-
tern. Carapace with uniformly infuscate marbling,
more densely infuscate medially. Pedipalp chela
ngers and manus, dorsal and retrolateral intercari-
nal surfaces with moderately infuscate marbling;
prolateral and ventral intercarinal surfaces mostly
immaculate. Legs retrolateral surfaces with infus-
cate marbling; prolateral surfaces pale, immaculate.
Tergites with unformly infuscate mottling, pale
stripe medially, blackish spots submedially, and
faint, narrow bands laterally. Sternites pale, with
faintly infuscate triangular to trapezoidal marking
at posterior margin of III, fading to infuscate mot-
tling on VII. Metasomal segments uniformly, faintly
marbled; segment V and telson markedly infuscate,
noticeably darker than preceding segments.
Carapace: Shape trapezoidal; anterior width
four-hs of posterior width (table 5); anterome-
dian sulcus moderately deep, oval; posterome-
dian sulcus shallow anteriorly, deeper posteriorly;
median ocular tubercle weakly granular; carinae
moderately developed, comprising small to
medium-sized granules (g. 6C).
Pedipalps: Orthobothriotaxic, Type A; femur
dorsal trichobothria with α conguration; pedi-
palp chela xed nger, trichobothrium db situ-
ated slightly distal to et. Femoral carinae serrate;
retromedian carinae comprising spiniform gran-
ules; dorsal intercarinal surface moderately gran-
ular; prolateral intercarinal surface with series of
large spiniform granules. Patella carinae strongly
developed, granular; prolateral intercarinal sur-
face with ve or six large, subspiniform granules.
Chela manus prodorsal and dorsomedian cari-
nae absent; retrodorsal carina weakly developed,
granular. Fixed nger, median denticle row com-
prising eight oblique subrows, each anked by
pro- and retrolateral supernumerary denticles.
Movable nger, median denticle row with short
terminal row comprising four denticles preceded
by eight oblique subrows, each anked by pro-
and retrolateral supernumerary denticles.
Legs: Leg I length 2× carapace length (table
10). Telotarsi ventral surfaces densely covered
with short setae; ungues markedly curved.
Pectines: Pectinal plate 1.65× wider than long;
posterior margin distinctly rounded; pectinal
tooth count 16/15 (♂) (g. 9C, table 5).
Mesosoma: Tergites width similar to carapace
posterior width; I and II slightly narrower (table
5). Pretergites surfaces smooth to nely granular.
Posttergites surfaces weakly granular; I–VI with
dorsomedian carinae weakly granular, restricted
to posterior half of each segment; VII surface
weakly granular, dorsomedian carinae moder-
ately granular, dorsosubmedian carinae weakly
granular, dorsolateral carinae absent. Sternites
III–VI, surfaces smooth; VII surface smooth,
ventrolateral carinae reduced to few granules.
12 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 452
Metasoma: Metasoma length 3.23× mesosoma
length (table 10). Segments longer than wide;
increasing in length posteriorly, segment V 2×
length of I; carinae weakly developed, smooth on
segments I–IV (gs. 17–21H), absent or obsolete
on V (figs. 20–22H); intercarinal surfaces
sparsely granular.
Tels o n : Vesicle elongate, ovoid; ventral surface
shallowly convex; subaculear tubercle narrow
and angular in lateral aspect, directed toward
midpoint of aculeus. Aculeus angled ventrally at
slightly less than 90° (gs. 23–25H).
Variation: Base color varies from pale yellow to
light orange with considerable variation in infus-
cation of the carapace and mesosoma, despite
localities being less than 50 km apart. Adult males
and females dier as follows. e mesosoma is
proportionally longer and slenderer, the meta-
soma up to 3× longer, with segment V markedly
longer, and the telson more elongate, with the
vesicle more rounded and bilobed posteriorly, in
males (gs. 20H, K, 21H, K, 22H, K, table 5). e
tegument is more densely infuscate, the pectinal
plate produced into a rounded lobe posteriorly,
which is punctate and slightly infuscate, and the
telson shorter and narrower with the vesicle sur-
faces less granular, in females (gs. 9C, D, 23H, K,
24H, K, 25H, K, 34–35A, B).
D: Centruroides berstoni is
endemic to the Izabal Department of Guatemala
and has been recorded from several localities
around Morales and Río Dulce (gs. 1B, D, 4).
FIGURE 3. Map of central Mexico, plotting known locality records of four species of the arboreal Neotropical
“thorellii” clade of Centruroides Marx, 1890, bark scorpions (Buthidae C.L. Koch, 1837): C. catemacoensis, sp.
nov. (triangles); C. chanae, sp. nov. (circles); C. cuauhmapan, sp. nov. (diamonds); C. rileyi Sissom, 1995
(squares).
2021 GOODMAN ET AL.: REVISION OF “THORELLII” CLADE OF CENTRUROIDES 13
FIGURE 4. Map of southeastern Mexico and northwestern Central America, plotting known locality records of
ve species of the arboreal Neotropical “thorellii” clade of Centruroides Marx, 1890, bark scorpions (Buthidae
C.L. Koch, 1837): C. berstoni, sp. nov. (circles); C. hamadryas, sp. nov. (diamonds); C. homanni Armas, 1996
(triangles); C. schmidti Sissom, 1995 (squares); C. yucatanensis, sp. nov. (crosses).
E: e localities at which C. berstoni
was recorded range in altitude from 15–17 m and
occur in a region of humid tropical lowland rain-
forest. Specimens were collected at night with UV
light detection, mostly on trees, sitting on bark or
branches, and large shoots of bamboo. e habitat
and habitus are consistent with the arboreal, cor-
ticolous ecomorphotype (Prendini, 2001a).
R: The paratype female of C.
schmidti, from Morales, Guatemala, was mis-
identified by Sissom (1995). Differences
between the paratype and other females that
species, evident from the illustrations (Sissom,
1995: 94–95 gs. 16–18), include faint reticulate
infuscation on the chelicerae, less infuscation of
the carapace and tergites, less granulation and
setation of the metasomal segments, and a
bilobed telson vesicle.
M E: GUATEMALA: Depar-
tamento Izabal: Município Livingston: Biotopo
Chocón Machacas, 15°44′05.3″N 88°54′57.2″W,
15 m, 25.ix.2019, A.M. Goodman, 8 ♂, 1 ♀, 1
juv. ♂ (CASENT 9073271); Río Dulce, Hotel
Tijax, 15°40′12.2″N 89°00′27″W, 49 m, 8.vii.2006,
J. Hu, C. Víquez, and D. Ortíz, collected along
trails through old secondary growth tropical for-
est using UV at night, 1 ♂ (AMNH [LP 5984]),
15°39′51.2″N 89°00′14.6″W, 17 m, 24.ix.2019, A.M.
Goodman, collected along gravel road of Hacienda
Tijax Parking Lot, anked by bamboo groves and
live fencing, 1 ♂ (CASENT 9073272), 15°44′05.3″N
88°54′57.2″W, 15 m, 25.ix.2019, A.M. Goodman, 8
14 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 452
TABLE 2
Meristic data for Centruroides rileyi Sissom, 1995
Material deposited in the Colección Nacional de Arácnidos (CNAN), Universidad Nacional Autónoma de
México, Mexico City. Measurements follow Stahnke (1970), Lamoral (1979), and Prendini (2001b).
♂ ♀
CNAN CNAN
SC4002 SC4003 SC3999 SC4003
Total length135.1 30 32.5 31.4 37.2 33.3 30.9
Carapace length 2.8 2.6 2.9 2.7 3.5 3.4 3.3
ant. width 1.5 1.3 1.4 1.1 1.6 1.7 1.8
post. width 3.1 2.9 3.0 2.9 3.7 3.7 3.8
Median ocelli diameter 0.3 0.2 0.3 0.2 0.3 0.3 0.3
Interocular length20.3 0.3 0.3 0.3 0.4 0.4 0.4
Pedipalp length317.4 14.3 15.1 14.9 18.3 17.6 17.6
Trochanter length 1.3 1.1 1.3 1.3 1.3 1.4 1.4
Femur length 3.2 2.6 2.8 2.6 3.3 3.2 3.2
width 0.5 0.5 0.5 0.5 0.7 0.7 0.8
height 0.8 0.6 0.8 0.7 1.0 1.0 1.0
Patella length 3.5 3.0 3.1 3.0 3.7 3.7 3.5
width 0.7 0.7 0.7 0.7 1.0 0.9 1.1
height 1.3 1.1 1.2 1.1 1.6 1.5 1.5
Chela length45.7 4.6 4.8 4.8 5.8 5.7 5.5
Manus length 2.4 2.0 1.8 2.2 2.2 2.1 2.0
width 1.1 1.0 1.0 1.1 1.2 1.2 1.2
height 1.1 0.9 1.0 1.1 1.3 1.1 1.2
Mov. nger length 3.7 3.0 3.1 3.2 4.2 3.6 4.0
Leg I length 5.8 5.0 5.3 5.1 6.3 6.6 6.2
Pectines length 4.7 4.1 4.3 4.0 4.5 4.6 4.5
tooth count 13/13 14/13 13/14 13/14 12/13 13/13 13/13
Mesosoma length57.8 9.0 9.0 8.9 12.5 9.6 9.3
Sternite VII length 2.2 2.4 2.6 2.5 3.0 2.5 2.3
width 2.6 2.5 2.9 2.8 4.0 3.7 3.8
Metasoma length624.5 18.4 20.6 19.8 21.2 20.3 19.8
Metasoma I length 3.0 2.2 2.3 2.4 2.5 2.4 2.4
width 1.1 1.1 1.2 1.2 1.6 1.6 1.4
height 1.2 1.2 1.3 1.3 1.5 1.6 1.5
Metasoma II length 3.8 2.8 3.1 3.0 3.2 3.1 3.0
width 1.1 1.1 1.2 1.2 1.4 1.5 1.3
height 1.1 1.1 1.2 1.2 1.4 1.4 1.3
2021 GOODMAN ET AL.: REVISION OF “THORELLII” CLADE OF CENTRUROIDES 15
♂ ♀
CNAN CNAN
SC4002 SC4003 SC3999 SC4003
Metasoma III length 4.2 3.2 3.5 3.4 3.5 3.4 3.3
width 1.1 1.2 1.2 1.3 1.5 1.4 1.4
height 1.0 1.1 1.1 1.1 1.4 1.4 1.4
Metasoma IV length 4.8 3.6 4.0 3.7 3.9 3.6 3.6
width 1.1 1.1 1.2 1.2 1.4 1.4 1.4
height 1.0 1.0 1.0 1.1 1.4 1.3 1.3
Metasoma V length 5.1 3.8 4.6 4.3 4.5 4.2 4.0
width 1.2 1.1 1.2 1.3 1.5 1.5 1.5
height 1.1 1.0 1.1 1.2 1.4 1.4 1.5
Tels on length 3.6 2.8 3.1 3.0 3.6 3.6 3.5
Vesicle length 2.5 1.8 2.1 2.0 2.4 2.1 2.3
width 1.1 0.8 1.0 1.0 1.2 1.2 1.2
height 1.0 0.8 0.9 0.9 1.2 1.0 1.1
Aculeus length 1.3 1.1 1.1 1.0 1.5 1.6 1.5
1 Sum of carapace, tergites I–VII, metasomal segments I–V, and telson; 2 distance between median ocelli; 3 sum of trochanter,
femur, patella, and chela; 4 measured from base of condyle to tip of xed nger; 5 sum of tergites I–VII; 6sum of metasomal
segments I–V and telson.
TABLE 2 continued
♂, 1 ♀, 1 juv. ♂ (CASENT 9073271). Município
Morales: Morales, Finca Fiymeza, Sendero Anbio,
15°24′24.1″N 88°41′46.8″W, 595 m, 17.viii.2017,
D. Barrales and R. Monjaraz, 1 juv. ♀ (CNAN
SC3968).
Centruroides catemacoensis, sp. nov.
Figures 2, 3, 5E, F, 8E, F, 11C, 12C, 17M, P,
18M, P, 19M, P, 20M, P, 21M, P, 22M, P, 23M, P,
24M, P, 25M, P, 30, 31, tables 1, 6, 10
T M: MEXICO: Veracruz:
Município Catemaco: Holotype ♂ (CNAN
T01424), 4 ♂ paratypes (CASENT 9073286,
9073287, 9073366, CNAN T01421), 4 ♀ para-
types (CASENT 9073309, 9073314, CNAN
T01422, T01423), Estacion Biología Los Tuxtlas,
Universidad Nacional Autónoma de México
(UNAM), 18°34′54″N 95°04′54.6″W, 74–416 m,
17.vii–25.vii.2018, A.M. Goodman, J. Gorneau,
and M.K. Lippey; paratype ♂ (CNAN T01420),
Estacion Biología Los Tuxtlas, UNAM, Cerro El
Vigia Reserva de la Biosfera Los Tuxtlas,
18°34′47.9″N 95°04′29.2″W, 421–429 m, 27.
vii.2005, O.F. Francke, M. Cordóva, A. Jaimes, A.
Valdez, and H. Montaño.
E: e species name refers to the
town of Catemaco, nearby the type locality, in
the state of Veracruz, Mexico.
D: Centruroides catemacoensis diers
from the closely related species, C. berstoni, C.
cuauhmapan, C. hamadryas, and C. rileyi as fol-
lows. e base coloration of C. catemacoensis var-
ies from dark yellow to orange, with considerable
infuscation on the pedipalps and metasoma
whereas the base coloration of C. berstoni, C.
cuauhmapan, and C. hamadryas is pale yellow and
of C. rileyi, yellowish orange. e carapace pos-
terosubmedian carinae are reduced or absent in C.
catemacoensis (g. 5E, F), absent in C. berstoni
(g. 6C, D), C. cuauhmapan (g. 5C, D), and C.
hamadryas (g. 6A, B), and present in C. rileyi
(g. 5A, B). e pedipalps are longer, and the
16 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 452
chela manus dorsoventrally compressed, in the
male, whereas the manus is shorter and slenderer
in the female of C. catemacoensis than the other
species (g. 11C). Additionally, the chela movable
nger is 2.5–3× (male) or 2.5× (female) the length
of the manus in C. catemacoensis (gs. 11, 12C,
table 6) but 1.5–1.7× (male) or 1.3× (female) the
length of the manus in C. berstoni (gs. 13, 14B,
table 5), C. cuauhmapan (gs. 11, 12B, table 3), C.
hamadryas (gs. 13, 14A, table 4), and C. rileyi
(gs. 11, 12A, table 2). e pedipalp chela manus
is two-thirds the length of the patella in C.
catemacoensis (gs. 11, 12C, table 6), but half its
length in C. berstoni (gs. 13, 14B, table 5), C.
cuauhmapan (gs. 11, 12B, table 3), C. hamadryas
(gs. 13, 14A, table 4), and C. rileyi (gs. 11, 12A
table 2). e prodorsal carina is restricted to the
distal half of the chela manus in C. catemacoensis
(gs. 11, 12C), whereas it is complete in C. cuauh-
mapan (gs. 11, 12B), C. hamadryas (gs. 13,
14A), and C. rileyi (gs. 11, 12A) and absent in C.
chanae (gs. 15, 16B) and C. yucatanensis (gs. 15,
16C). e rst leg is less than 2× the length of the
carapace in the male of C. catemacoensis but
greater than 2× its length in C. cuauhmapan, C.
hamadryas, and C. rileyi (table 10). e ventro-
submedian and ventrolateral carinae of sternite
VII of the male are weakly developed in C.
catemacoensis but well developed, granular in C.
cuauhmapan and obsolete to absent in C. berstoni
and C. rileyi. e metasoma of the male is greater
than 3× the length of the mesosoma in C. cuauh-
mapan but less than 3× its length in C. berstoni, C.
hamadryas, and C. rileyi (table 10). e ventrolat-
eral and ventrosubmedian carinae of metasomal
segments I–IV in the male are absent in C.
catemacoensis (gs. 18, 19M), weakly developed,
smooth in C. berstoni (gs. 18H, K, 19H, K, 21H,
K, 22H, K), well developed in C. cuauhmapan
(gs. 18G, J, 19G, J, 21G, J, 22G, J), weakly devel-
oped on segments I–III and absent on IV in C.
hamadryas (gs. 18B, E, 19B, E), and vestigial,
smooth in C. rileyi (gs. 18A, D, 19A, D). e
metasomal carinae of the female are well devel-
oped, granular in C. catemacoensis (gs. 17–22P),
but nely granular in C. berstoni (gs. 17–22K), C.
cuauhmapan (gs. 17–22J), C. hamadryas (gs.
17–22E), and C. rileyi (gs. 17–22D).
D: e following description is
based on the holotype male, with dierences
among other material noted in the section on
variation.
Coloration: Base color yellow to orange, with
extensive infuscation, creating mottled or mar-
bled pattern. Carapace with uniformly infuscate
marbling, more densely infuscate medially. Pedi-
palp chela ngers and manus, dorsal and retro-
lateral intercarinal surfaces with moderately
infuscate marbling; prolateral and ventral inter-
carinal surfaces mostly immaculate. Legs retro-
lateral surfaces with infuscate marbling;
prolateral surfaces pale, immaculate. Tergites
with unformly infuscate mottling, pale stripe
medially, blackish spots submedially, and dis-
tinct, narrow bands laterally. Sternites moder-
ately infuscate, with faintly infuscate triangular
marking at posterior margin of III, fading to
infuscate mottling on VII. Metasomal segments
uniformly, faintly marbled; segment V and telson
markedly infuscate, noticeably darker than pre-
ceding segments.
Carapace: Shape trapezoidal; anterior width
four-hs of posterior width (table 6); anterome-
dian sulcus moderately deep, narrow; postero-
median sulcus shallow anteriorly, deep, narrow
posteriorly; carinae moderately developed, com-
prising small to medium-sized granules (g. 5E).
Pedipalps: Orthobothriotaxic, Type A;
femur dorsal trichobothria with α configura-
tion; pedipalp chela fixed finger, trichoboth-
rium db situated slightly distal to et. Femoral
carinae moderately to strongly developed, ser-
rate; retromedian carinae comprising spini-
form granules; dorsal intercarinal surface
moderately granular; prolateral intercarinal
surface with series of large spiniform granules.
Patella dorsomedian, retrodorsal, prodorsal,
and proventral carinae moderately developed,
serrate; retromedian carina strongly devel-
oped, serrate; retroventral carina incomplete,
serrate; prolateral intercarinal surface with five
or six large, subspiniform granules. Chela
2021 GOODMAN ET AL.: REVISION OF “THORELLII” CLADE OF CENTRUROIDES 17
FIGURE 5. Centruroides Marx, 1890, carapace, dorsal aspect. A, B. C. rileyi Sissom, 1995, A. ♂ (CNAN
SC4002), B. ♀ (CNAN SC4003). C, D. C. cuauhmapan, sp. nov., C. holotype ♂ (CNAN T01396), D. paratype
♀ (CNAN T01399). E, F. C. catemacoensis, sp. nov., E. holotype ♂ (CNAN T01424), F. paratype ♀ (CNAN
T01423). Scale bars = 1 mm.
18 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 452
FIGURE 6. Centruroides Marx, 1890, carapace, dorsal aspect. A, B. C. hamadryas, sp. nov., A. holotype ♂
(CNAN T01408), B. paratype ♀ (CNAN T01415). C, D. C. berstoni, sp. nov., C. holotype ♂ (CASENT
9073325), D. paratype ♀ (CASENT 9073313). E, F. C. schmidti Sissom, 1995, E. ♂ (CASENT 9073316), F. ♀
(CASENT 9073317). Scale bars = 1 mm.
2021 GOODMAN ET AL.: REVISION OF “THORELLII” CLADE OF CENTRUROIDES 19
FIGURE 7. Centruroides Marx, 1890, carapace, dorsal aspect. A, B. C. homanni Armas, 1996, A. ♂ (CNAN
SC3996), B. ♀ (CNAN SC3996). C, D. C. chanae, sp. nov., C. holotype ♂ (CNAN T01403), D. paratype ♀
(CNAN T01405). E, F. C. yucatanensis, sp. nov., E. holotype ♂ (CNAN T01416), F. paratype ♀ (CNAN
T01417). Scale bars = 1 mm.
20 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 452
TABLE 3
Meristic data for type material of Centruroides cuauhmapan, sp. nov.
Material deposited in the Colección Nacional de Arácnidos (CNAN), Universidad Nacional Autónoma de
México, Mexico City. Measurements follow Stahnke (1970), Lamoral (1979), and Prendini (2001b).
Holotype Paratypes Paratypes
♂ ♂ ♀
CNAN CNAN CNAN
T01396 T01397 T01398 T01402 T01399 T01400 T01401
Total length133.8 31.9 30.9 32.8 32.4 28.2 33.5
Carapace length 2.9 2.7 2.8 2.8 3.1 2.7 3.1
ant. width 1.4 1.5 1.4 2.6 1.6 1.5 1.6
post. width 2.9 3.1 2.9 2.8 3.4 3.0 3.5
Median ocelli diameter 0.3 0.2 0.3 0.2 0.3 0.2 0.3
Interocular length20.3 0.3 0.3 0.3 0.3 0.3 0.4
Pedipalp length316.6 15.4 14.9 15.3 16.2 14.7 15.8
Trochanter length 1.3 1.2 1.1 1.3 1.1 1.3 1.3
Femur length 3.1 2.9 2.7 2.8 3.0 2.7 2.8
width 0.5 0.6 0.5 0.5 0.6 0.6 0.6
height 0.7 0.7 0.7 0.7 0.8 0.7 0.9
Patella length 3.5 3.2 3.1 3.1 3.3 3.0 3.3
width 0.7 0.9 0.7 0.7 0.8 0.8 0.9
height 1.3 1.3 1.2 1.3 1.4 1.2 1.3
Chela length45.3 4.9 4.8 4.8 5.2 4.6 4.9
Manus length 2.1 2.1 1.7 2.2 2.0 1.8 1.9
width 1.0 1.1 1.0 1.0 1.0 0.9 1.0
height 2.2 1.1 1.0 1.0 1.0 0.8 1.0
Mov. nger length 3.4 3.2 3.2 3.3 3.6 3.1 3.5
Leg I length 5.5 5.0 4.7 5.0 5.8 5.1 5.5
Pectines length 2.1 2.1 2.1 2.1 2.2 2.1 1.9
tooth count 13/13 15/14 14/13 14/14 12/12 14/14 10/11
Mesosoma length58.0 8.2 8.0 9.6 10.5 9.0 10.6
Sternite VII length 2.3 2.1 2.3 2.5 2.5 2.2 2.7
width 2.8 2.7 2.7 2.7 3.6 3.1 3.7
Metasoma length622.9 21.0 20.1 20.4 18.8 16.5 19.8
Metasoma I length 3.0 2.5 2.3 2.6 2.3 2.0 2.2
width 1.2 1.2 1.2 1.2 1.4 1.2 1.4
height 1.2 1.3 1.3 1.3 1.5 1.3 1.5
Metasoma II length 3.4 3.2 3.0 3.2 3.0 2.5 2.8
width 1.2 1.2 1.2 1.1 1.4 1.2 1.4
height 1.1 1.2 1.1 1.1 1.4 1.2 1.4
2021 GOODMAN ET AL.: REVISION OF “THORELLII” CLADE OF CENTRUROIDES 21
Holotype Paratypes Paratypes
♂ ♂ ♀
CNAN CNAN CNAN
T01396 T01397 T01398 T01402 T01399 T01400 T01401
Metasoma III length 3.8 3.7 3.4 3.5 3.2 2.6 3.1
width 1.2 1.2 1.2 1.1 1.4 1.2 1.4
height 1.1 1.2 1.1 1.1 1.3 1.2 1.3
Metasoma IV length 4.5 4.1 4.0 3.8 3.6 3.0 3.5
width 1.2 1.2 1.1 1.1 1.4 1.2 1.4
height 1.1 1.2 1.1 1.1 1.3 1.1 1.3
Metasoma V length 4.8 4.7 4.4 4.3 3.7 3.6 4.3
width 1.2 1.3 1.2 1.2 1.4 1.2 1.4
height 1.1 1.2 1.1 1.1 1.3 1.1 1.3
Tels on length 3.4 2.8 3.0 3.0 3.0 2.8 3.9
Vesicle length 2.2 1.7 2.0 2.0 2.0 1.6 2.0
width 0.9 0.9 1.0 1.1 1.0 0.8 1.0
height 0.9 0.9 0.8 1.0 1.0 0.7 1.1
Aculeus length 1.3 1.2 1.2 0.9 1.4 1.2 1.5
1 Sum of carapace, tergites I–VII, metasomal segments I–V, and telson; 2 distance between median ocelli; 3 sum of trochanter,
femur, patella, and chela; 4 measured from base of condyle to tip of xed nger; 5 sum of tergites I–VII; 6 sum of metasomal
segments I–V and telson.
TABLE 3 continued
manus proventral carina moderately devel-
oped, comprising few rounded granules; other
carinae weakly developed, granular. Fixed fin-
ger, median denticle row comprising eight
oblique subrows, each flanked by pro- and ret-
rolateral supernumerary denticles. Movable
finger, median denticle row with short termi-
nal row comprising four denticles preceded by
eight oblique subrows, each flanked by pro-
and retrolateral supernumerary denticles.
Legs: Leg I length 1.94× greater than carapace
length (table 10). Telotarsi ventral surfaces
densely covered with short setae; ungues mark-
edly curved.
Pectines: Pectinal plate 1.9× wider than long;
posterior margin distinctly rounded; pectinal
tooth count 14/14 (♂) (g. 8E, table 6).
Mesosoma: Tergites width similar to carapace
posterior width; I and II slightly narrower (table
6). Pretergites surfaces smooth to nely granular.
Posttergites surfaces weakly granular; I–VI with
dorsomedian carinae vestigial, reduced to several
small granules; VII surface weakly granular, dor-
somedian carina vestigial, reduced to several
small granules, dorsosubmedian and dorsolateral
carinae nely serrate. Sternites III–VI, surfaces
smooth; VII surface weakly granular, ventrolat-
eral carinae serrate.
Metasoma: Metasoma length 2.83× mesosoma
length (table 6). Segments longer than wide;
increasing in length posteriorly, segment V 2×
length of I; dorsolateral and dorsosubmedian
carinae weakly serrate on segments I–III, other
carinae absent or obsolete; intercarinal surfaces
sparsely granular (gs. 17–22M).
Tels o n : Vesicle elongate, ovoid, length 1.5×
width (table 6); ventral surface shallowly convex;
ventromedian carina granular, terminating at
subaculear tubercle; subaculear tubercle narrow
and angular in lateral aspect, directed toward
22 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 452
midpoint of aculeus. Aculeus angled ventrally at
slightly less than 90° (g. 25M).
Variation: Base coloration varies from light
yellow to orange with considerable variation in
infuscation of the carapace and mesosoma
(figs. 30, 31A, B). Adult males and females dif-
fer as follows. The mesosoma is proportionally
longer and slenderer, the metasoma up to 3×
longer, with segment V markedly longer, and
the telson more elongate, with the vesicle more
rounded and bilobed posteriorly, in males
(figs. 17–22M, table 6). The tegument is more
densely infuscate, the pectinal plate is pro-
duced into a rounded lobe posteriorly, which
is punctate and slightly infuscate, and the tel-
son is shorter and narrower, with the vesicle
surfaces less granular, in females (figs.
23–25P). The pectinal tooth count is similar in
both sexes (fig. 8E, F, table 6).
D: Centruroides catemacoensis is
endemic to the Los Tuxtlas Biosphere Reserve
in the state of Veracruz, eastern Mexico. Most
of the known material was collected in the
vicinity of the Estación Biologica Los Tuxtlas,
UNAM (g. 3).
E: The localities at which C.
catemacoensis has been recorded range in alti-
tude from 74 to 493, in an area of tropical moist
broadleaf forest, dominated by thorny palms,
Astrocaryum mexicanum Leibm. (Arecaceae
Berch and J. Presl,), and buttress trees including
Siparuna andina Tul. (Sipuranaceae A. DC). and
Vochysia guatemalensis Donn. Sm. (Vochysiaceae
A. St.-Hil). is arboreal scorpion has been col-
lected from 3 to 15 m above ground in the forest
canopy, oen in bromeliads, Aechmea bracteata
Grisebach (Bromeliaceae Juss). A few individuals
were collected in the leaf litter. e habitat and
habitus are consistent with the arboreal, cortico-
lous ecomorphotype (Prendini, 2001a).
M E: MEXICO: Veracruz:
Município Catemaco: Estacion Biología Los
Tuxtlas, UNAM, 18°35′05.6″N 95°04′29.9″W,
162 m, 27.vi.1968, C.R.B., in A. bracteata at 15
m, 2 ♂, 1 ♀ (CNAN SC3975), 25.i.1969, C.R.B.,
15 m above ground in A. bracteata, 1 ♀, 1 juv.
♀ (CNAN SC3976), 22.v.1969, C.R.B., A. brac-
teata at 12 m, 1 juv. ♂ (CNAN SC3972), 14.
vi.1969, C.R.B., in A. bracteata at 10 m, 1 ♂
(CNAN SC3974), 15.viii.1997, G. Pérez II, 1 ♂
(CNAN SC3971), 134 m, 18.iii.1998, D.E.
González Manuel, habitación, 1 ♀ (CNAN
SC3969), 162 m, 16.iv.1998, in leaf litter, 1 ♀,
2 juv. ♀ (CNAN SC3970), iii.2001, M. López,
15 m above ground in A. bracteata, 1 ♀ (CNAN
SC3977), 134 m, 26.viii.2005. O.F. Francke, M.
Córdova, A. Jaimes, A. Valdez, and H. Mon-
taño, 1 ♂ (AMNH [LP 5231]), 18°34′54″N
95°04′54.6″W, 134 m, 19.vii.2002, J. Ponce and
O.F. Francke, 1 ♀ (AMNH [LP 2070]), 74–416
m, 17.vii–25.vii.2018, A.M. Goodman, J. Gor-
neau, and M.K. Lippey, 2 ♂ (CASENT 9073270,
9073427), 2 juv. ♂ (CASENT 9073315,
9073409), 4 juv. ♀ (CASENT 9073408,
9073410, 9073426, 9073428); surroundings of
Estacion Biología, Reserva de la Biosfera Los
Tuxtlas, 18°30′03.5″N 95°04′29.5″W, 134–493
m, 26.viii.2005, O.F. Francke, M. Cordova, A.
Jaimes, A. Valdez, H. Montaño, 2 ♀, 1 juv. ♀
(CNAN SC3973).
Centruroides chanae, sp. nov.
Figures 2, 3, 7C, D, 10C, D, 15B, 16B, 17I, L,
18I, L, 19I, L, 20I, L, 21I, L, 22I, L, 23I, L, 24I,
L, 25I, L, 40,41, tables 1, 9, 10
T M: MEXICO: Guerrero:
Município Copala: Holotype ♂ (CNAN T01407),
Microondas Fogos, 16°33′59.5″N 98°53′18.1″W,
14 m, 2.xi.2007, O.F. Francke, H. Montaño, and
A. Ballesteros; 2 ♂ paratypes (CNAN T01403,
T01404), 2 ♀ paratypes (CNAN T01405,
T01406), same data except: 103 m, 22.vi.2007,
O.F. Francke, M. Escalante, H. Montaño, and J.
Ballesteros.
E: e species name honors Ken-
dra Chan, a friend of the rst author, who passed
away in 2018.
D: Centruroides chanae is most closely
related to C. homanni, from which it diers as fol-
lows. A dark line along the lateral margins of the
2021 GOODMAN ET AL.: REVISION OF “THORELLII” CLADE OF CENTRUROIDES 23
FIGURE 8. Centruroides Marx, 1890, pectines and sternum, ventral aspect. A, B. C. rileyi Sissom, 1995, A. ♂
(CNAN SC4002), B. ♀ (CNAN SC4003). C, D. C. cuauhmapan, sp. nov., C. holotype ♂ (CNAN T01396), D.
paratype ♀ (CNAN T01399). E, F. C. catemacoensis, sp. nov., E. holotype ♂ (CNAN T01424), F. paratype ♀
(CNAN T01423). Scale bars = 1 mm.
24 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 452
FIGURE 9. Centruroides Marx, 1890, pectines and sternum, ventral aspect. A, B. C. hamadryas, sp. nov., A.
holotype ♂ (CNAN T01408), B. paratype ♀ (CNAN T01415). C, D. C. berstoni, sp. nov., C. holotype ♂
(CASENT 9073325), D. paratype ♀ (CASENT 9073313). E, F. C. schmidti Sissom, 1995, E. ♂ (CASENT
9073316), F. ♀ (CASENT 9073317). Scale bars = 1 mm.
2021 GOODMAN ET AL.: REVISION OF “THORELLII” CLADE OF CENTRUROIDES 25
FIGURE 10. Centruroides Marx, 1890, pectines and sternum, ventral aspect. A, B. C. homanni Armas, 1996,
A. ♂ (CNAN SC3996), B. ♀ (CNAN SC3996). C, D. C. chanae, sp. nov., C. holotype ♂ (CNAN T01403), D.
paratype ♀ (CNAN T01405). E, F. C. yucatanensis, sp. nov., E. holotype ♂ (CNAN T01416), F. paratype ♀
(CNAN T01417). Scale bars = 1 mm.
26 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 452
TABLE 4
Meristic data for type material of Centruroides hamadryas, sp. nov.
Material deposited in the Colección Nacional de Arácnidos (CNAN), Universidad Nacional Autónoma de
México, Mexico City. Measurements follow Stahnke (1970), Lamoral (1979), and Prendini (2001b).
Holotype Paratypes Paratypes
♂ ♂ ♀
CNAN CNAN CNAN
T01408 T01409 T01411 T01412 T01414 T01410 T01413 T01415
Total length144.5 35.9 35 39 37.7 29.1 31.2 34.8
Carapace length 3.5 2.9 3.0 3.3 3.1 3.0 3.2 3.3
ant. width 1.8 1.5 1.5 1.7 1.5 1.5 1.6 1.8
post. width 3.6 3.2 3.2 3.5 3.3 3.3 3.4 3.6
Median ocelli diameter 0.3 0.3 0.3 0.3 0.3 0.3 0.3 0.3
Interocular length20.4 0.3 0.3 0.3 0.3 0.3 0.3 0.3
Pedipalp length320.6 16.7 16.6 18.7 16.7 15.3 14.0 17.2
Trochanter length 1.6 1.3 1.3 1.2 1.1 1.2 1.3 1.4
Femur length 4.0 3.2 3.1 3.6 3.3 2.8 3.0 3.0
width 0.6 0.6 0.5 0.6 0.5 0.6 0.6 0.7
height 1.0 0.8 0.8 0.8 0.7 0.8 0.9 0.8
Patella length 4.5 3.5 3.5 3.9 3.6 3.2 3.5 3.4
width 0.8 0.7 0.8 0.8 0.8 0.9 0.9 1.0
height 1.6 1.3 1.3 1.4 1.3 1.3 1.4 1.5
Chela length46.5 5.4 5.3 6.2 5.4 4.8 5.2 5.6
Manus length 2.9 2.4 2.5 2.8 2.3 2.0 2.0 2.1
width 1.3 1.2 1.1 1.2 1.2 1.1 1.2 1.1
height 1.1 1.1 1.0 1.1 1.1 1.1 1.1 1.1
Mov. nger length 4.0 3.3 3.4 3.8 3.3 3.3 1.0 3.8
Leg I length 6.6 5.5 5.5 5.8 5.8 5.4 6.0 5.6
Pectines length 5.2 4.6 4.5 5.0 4.5 4.2 3.8 5.8
tooth count 13/14 15/16 14/15 14/13 14/14 13/13 12/12 14/14
Mesosoma length510.2 8.6 8.4 8.5 8.7 8.5 8.8 10.2
Sternite VII length 3.0 2.6 2.6 2.5 2.4 2.9 2.4 2.4
width 3.3 2.8 2.7 2.8 2.8 3.3 3.3 3.4
Metasoma length630.8 24.4 23.6 27.2 25.9 17.6 19.2 21.3
Metasoma I length 3.9 3.0 3.1 3.6 3.2 2.3 2.5 2.8
width 1.4 1.2 1.2 1.3 1.3 1.3 1.4 1.4
height 1.4 1.2 1.3 1.3 1.3 1.4 1.6 1.6
Metasoma II length 4.9 3.8 3.3 4.3 3.9 2.7 3.0 3.2
width 1.3 1.3 1.2 1.3 1.3 1.3 1.4 1.4
height 1.3 1.2 1.2 1.2 1.2 1.3 1.4 1.4
2021 GOODMAN ET AL.: REVISION OF “THORELLII” CLADE OF CENTRUROIDES 27
Holotype Paratypes Paratypes
♂ ♂ ♀
CNAN CNAN CNAN
T01408 T01409 T01411 T01412 T01414 T01410 T01413 T01415
Metasoma III length 5.5 4.5 4.2 4.8 4.5 2.9 3.2 3.5
width 1.3 1.3 1.3 1.3 1.3 1.4 1.4 1.4
height 1.2 1.1 1.2 1.1 1.2 1.3 1.3 1.3
Metasoma IV length 6.2 4.7 4.8 5.2 5.0 3.3 3.5 3.9
width 1.3 1.3 1.2 1.2 1.3 1.3 1.4 1.3
height 1.2 1.2 1.2 1.2 1.2 1.3 1.3 1.3
Metasoma V length 6.5 5.3 5.1 5.7 5.6 3.7 4.0 4.4
width 1.4 1.3 1.3 1.3 1.4 1.4 1.5 1.4
height 1.3 1.2 1.3 1.3 1.3 1.3 1.3 1.3
Tels on length 3.8 3.1 3.1 3.6 3.7 2.7 3.0 3.5
Vesicle length 2.7 2.2 2.0 2.6 2.5 1.4 1.6 2.2
width 1.2 1.0 1.0 1.1 1.1 1.0 1.5 1.0
height 1.2 1.0 1.0 1.1 1.3 0.9 1.0 1.0
Aculeus length 1.3 1.1 1.2 1.2 1.3 1.4 0.9 1.5
1 Sum of carapace, tergites I–VII, metasomal segments I–V, and telson; 2 distance between median ocelli; 3 sum of trochanter,
femur, patella, and chela; 4 measured from base of condyle to tip of xed nger; 5 sum of tergites I–VII; 6 sum of metasomal
segments I–V and telson.
TABLE 4 continued
carapace and mesosomal tergites I–III, and pale
stripe medially on the carapace and tergites, present
in C. chanae, are absent in C. homanni (g. 7A,
D). e carapace, pedipalps, tergites, and meta-
soma are less infuscate, creating a less mottled
appearance, in C. chanae than C. homanni. More
reticulate infuscation is present on the chelicerae of
C. chanae than those of C. homanni. e interocu-
lar triangle is less darkly infuscate in C. chanae than
C. homanni. e marbled infuscation of the
mesosomal sternites is faint or absent in C. chanae
but pronounced in C. homanni. e carapace is
shorter, its length and width similar, in C. chanae,
but longer, its length greater than its width, in C.
homanni. e carapace surfaces are more nely
granular, the carinae less developed and the sulci
broader and shallower in C. chanae than C. ho-
manni. e pedipalp chela manus of the male is less
incrassate in C. chanae (gs. 15, 16A) than C. ho-
manni (gs. 15, 16B). e ventral surfaces of the
telotarsi of leg I are more coarsely and densely
setose in C. chanae than C. homanni. e pectinal
tooth count of the male is higher in C. chanae, usu-
ally 17 (g. 10A) than C. homanni, usually 15 (g.
10C, tables 8, 9). e ventrolateral carinae of meso-
somal sternite VII are distinct, granular and the
ventrosubmedian carinae weakly developed, granu-
lar in C. chanae, whereas the ventrolateral carinae
are granular, and the ventrosubmedian carinae
weakly granular and restricted to the posterior half
of the segment in C. homanni. Although the
metasomal segments of the male are shorter and
broader in C. chanae than C. homanni, the meta-
soma is more than 3× (up to 3.3×) the length of the
mesosoma in C. chanae, but less than 3× its length
in C. homanni (table 10). e ventrolateral and
ventrosubmedian carinae of metasomal segments
I–IV are less pronounced in C. chanae, being nely
28 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 452
granular to subserrate on I–III and obsolete,
smooth on IV (gs. 18I, L, 19I, L), compared with
slightly serrate on I–IV in C. homanni (g. 18C, F,
19C, F). e ventrosubmedian carinae of meta-
somal segments I and II are absent or obsolete in
C. chanae but very pronounced in C. homanni.
e telson of the male is shorter, the vesicle rounded
posteriorly, in C. chanae (gs. 23–25I) whereas the
telson is elongate and the vesicle bilobed posteriorly
in C. homanni (gs. 23–25C).
D: e following description is based
on the holotype male, with dierences among other
material noted in the section on variation.
Coloration: Base color light yellow, with exten-
sive infuscation, creating mottled or marbled pat-
tern. Carapace with uniformly infuscate marbling,
more densely infuscate medially. Pedipalp chela
ngers and manus, dorsal and retrolateral inter-
carinal surfaces with moderately infuscate mar-
bling; prolateral and ventral intercarinal surfaces
mostly immaculate. Legs retrolateral surfaces with
infuscate marbling; prolateral surfaces pale,
immaculate. Tergites with unformly infuscate
mottling, pale stripe medially, blackish spots sub-
medially, and faint, narrow bands laterally. Ster-
nites pale, mostly immaculate. Metasomal
FIGURE 11. Centruroides Marx, 1890, pedipalp chela, retrolateral aspect. A. C. rileyi Sissom, 1995, ♂ (CNAN
SC4002). B. C. cuauhmapan, sp. nov., holotype ♂ (CNAN T01396). C. C. catemacoensis, sp. nov., holotype ♂
(CNAN T01424). Scale bars = 1 mm.
2021 GOODMAN ET AL.: REVISION OF “THORELLII” CLADE OF CENTRUROIDES 29
segments uniformly marbled; segment V and tel-
son markedly infuscate, noticeably darker than
preceding segments.
Carapace: Shape trapezoidal; anterior width
four-hs of posterior width (table 9); anterome-
dian sulcus moderately deep, oval; posterome-
dian sulcus shallow anteriorly, deep posteriorly;
median ocular tubercle weakly granular; carinae
moderately developed, comprising small to
medium-sized granules; lateral ocular and pos-
terosubmedian carinae distinct; intercarinal sur-
faces nely and evenly granular (g. 7D).
Pedipalps: Orthobothriotaxic, Type A; femur
dorsal trichobothria with α conguration; pedi-
palp chela xed nger, trichobothrium db situated
slightly distal to et. Femoral carinae strongly
developed, serrate; retromedian carinae compris-
ing spiniform granules; prolateral intercarinal sur-
face with series of large spiniform granules. Patella
carinae strongly developed, granular; prolateral
intercarinal surface with ve or six large, subspini-
form granules. Chela manus dorsomedian and
retrodorsal carinae complete, granular; prodorsal
carina absent. Fixed nger, median denticle row
comprising eight oblique subrows, each anked by
pro- and retrolateral supernumerary denticles.
Movable nger, median denticle row with short
terminal row comprising four denticles preceded
FIGURE 12. Centruroides Marx, 1890, pedipalp chela, retrolateral aspect. A. C. rileyi Sissom, 1995, ♀ (CNAN
SC4003). B. C. cuauhmapan, sp. nov., paratype ♀ (CNAN T01399). C. C. catemacoensis, sp. nov., paratype ♀
(CNAN T01418). Scale bars = 1 mm.
30 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 452
by eight oblique subrows, each anked by pro-
and retrolateral supernumerary denticles.
Legs: Leg I length 1.79× greater than carapace
length (table 10). Telotarsi ventral surfaces densely
covered with short setae; ungues markedly curved.
Pectines: Pectinal plate 1.65× wider than long;
posterior margin distinctly rounded; pectinal
tooth count 17/17 (♂) (g. 10C, table 9).
Mesosoma: Tergites width similar to carapace
posterior width; I and II slightly narrower (table
9). Pretergites surfaces smooth to nely granular.
Posttergites surfaces weakly granular; I–VI with
dorsomedian carinae nely granular on I–III,
absent on IV–VI; VII surface weakly granular,
dorsomedian carina absent, dorsosubmedian
and dorsolateral carinae smooth. Sternites III–
VI, surfaces smooth; VII surface, ventrolateral
and ventrosubmedian carinae smooth.
Metasoma: Metasoma length 3.1× mesosoma
length (table 9). Segments longer than wide;
increasing in length posteriorly, segment V 2×
length of I; carinae nely granular on segments
I–III, smooth on IV, absent on V; intercarinal
surfaces sparsely granular (gs. 17–22I).
Tels o n : Vesicle elongate, ovoid; ventral surface
shallowly convex; ventromedian carina granular,
terminating at subaculear tubercle; subaculear
tubercle narrow and angular in lateral aspect,
directed toward midpoint of aculeus. Aculeus
angled ventrally at slightly less than 90° (g. 25I).
Variation: Base coloration varies from light
yellow to orange with considerable variation in
infuscation of the carapace and mesosoma (gs.
40, 41A, B). Adult males and females dier as
follows. e prodorsal carina of the pedipalp
chela manus is absent, the pectinal tooth count
FIGURE 13. Centruroides Marx, 1890, pedipalp chela, retrolateral aspect. A. C. hamadryas, sp. nov., holotype
♂ (CNAN T01408). B. C. berstoni, sp. nov., holotype ♂ (CASENT 9073325). C. C. schmidti Sissom, 1995, ♂
(CASENT 9073316). Scale bars = 1 mm.
2021 GOODMAN ET AL.: REVISION OF “THORELLII” CLADE OF CENTRUROIDES 31
higher (16 or 17), the mesosoma proportionally
longer and slenderer, the metasoma up to 3×
longer, with segment V markedly longer, and the
telson more elongate, with the vesicle more
rounded and smoother, in males (gs. 23I, L, 24I,
L, 25I, L, table 9). e prodorsal carina is granu-
lar and restricted to the distal half of the chela
manus, the tegument more densely infuscate, the
pectinal plate produced into a rounded lobe pos-
teriorly, which is punctate and slightly infuscate,
the pectinal tooth count lower (13 or 14), and
the telson shorter and narrower, with the vesicle
surfaces weakly granular, in females (gs. 10C,
D, 12B, C, 15–16B, 20I, L, 22I, L, 23I, L, 24I, L,
25I, L, table 9).
D: Centruroides chanae is
endemic to the states of Guerrero and Micho-
acán, in southwestern Mexico. The known
records extend from eastern Michoacán, near the
border of Colima, to western Guerrero, south of
the Sierra Madre del Sur and east of the Sierra
Madre Occidental (g. 3).
E: e localities at which C. chanae has
been recorded range in altitude from 8 to 221 m.
e habitat at these localities varies from low to
medium-height deciduous tropical forest and
FIGURE 14. Centruroides Marx, 1890, pedipalp chela, retrolateral aspect. A. C. hamadryas, sp. nov., paratype
♀ (CNAN T01415). B. C. berstoni, sp. nov., paratype ♀ (CASENT 9073313). C. C. schmidti Sissom, 1995, ♀
(CASENT 9073317). Scale bars = 1 mm.
32 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 452
TABLE 5
Meristic data for type material of Centruroides berstoni, sp. nov.
Material deposited in the California Academy of Sciences (CASENT), San Francisco. Measurements follow
Stahnke (1970), Lamoral (1979), and Prendini (2001b).
Holotype Paratypes Paratypes
♂ ♂ ♀
CASENT CASENT CASENT
9073325 9073312 9073326 9073298 9073313 9073324 9073368 9073297
Total length149.9 50 43.7 48.6 37.7 32.7 36.6 36.3
Carapace length 3.9 3.7 3.4 3.6 3.4 3.1 3.4 3.4
ant. width 1.9 1.8 1.7 1.9 1.8 1.5 1.8 1.7
post. width 3.9 3.9 3.5 3.9 3.9 3.3 3.7 3.8
Median ocelli diameter 0.3 0.3 0.3 0.3 0.3 0.3 0.3 0.3
Interocular length20.4 0.3 0.3 0.4 0.4 0.3 0.4 0.3
Pedipalp length323.6 22.8 20.5 22.7 19.9 16.9 19.7 19.1
Trochanter length 1.9 1.8 1.6 1.7 1.5 1.3 1.5 1.5
Femur length 4.5 4.4 4.0 4.3 3.4 2.8 3.5 3.5
width 0.7 0.7 0.7 0.8 0.6 0.5 0.7 0.7
height 1.0 1.0 1.0 1.0 1.0 0.9 1.0 1.0
Patella length 5.1 4.7 4.3 4.7 4.0 3.2 4.1 3.8
width 1.0 1.0 0.9 1.0 1.0 0.9 1.0 1.0
height 1.5 1.6 1.5 1.5 1.6 1.4 1.6 1.5
Chela length47.5 7.3 6.4 7.5 6.5 5.1 6.3 6.0
Manus length 3.5 3.2 2.8 3.3 2.4 1.9 2.4 2.2
width 1.6 1.6 1.5 1.6 1.2 1.0 1.2 1.2
height 1.4 1.4 1.4 1.5 1.1 1.0 1.1 1.0
Mov. nger length 4.6 4.6 4.2 4.5 4.5 4.5 4.3 4.3
Leg I length 7.7 7.6 6.5 7.2 6.5 5.3 6.4 6.6
Pectines length 6.6 6.0 6.0 6.0 5.2 4.8 5.1 4.8
tooth count 16/15 14/14 16/16 15/15 13/14 15/14 13/13 13/13
Mesosoma length510.7 12.5 10.4 12.0 11.3 10.0 10.5 11.3
Sternite VII length 3.2 3.4 3.0 3.3 2.9 3.2 2.7 2.7
width 3.3 3.3 3.3 3.4 4.2 2.5 3.8 3.8
Metasoma length635.3 33.8 29.9 33.0 23.0 19.6 22.7 21.6
Metasoma I length 4.5 4.2 3.7 4.2 3.1 2.3 2.6 2.6
width 1.4 1.4 1.5 1.6 1.5 1.3 1.4 1.5
height 1.4 1.4 1.4 1.5 1.7 1.5 1.6 1.7
Metasoma II length 5.6 5.3 4.6 5.2 3.4 2.9 3.3 3.2
width 1.4 1.4 1.4 1.5 1.5 1.4 1.5 1.5
height 1.3 1.3 1.3 1.4 1.4 1.3 1.5 1.4
2021 GOODMAN ET AL.: REVISION OF “THORELLII” CLADE OF CENTRUROIDES 33
Holotype Paratypes Paratypes
♂ ♂ ♀
CASENT CASENT CASENT
9073325 9073312 9073326 9073298 9073313 9073324 9073368 9073297
Metasoma III length 6.5 6.0 5.2 5.8 3.7 3.1 3.7 3.5
width 1.4 1.4 1.3 1.5 1.6 1.4 1.5 1.7
height 1.4 1.3 1.4 1.4 1.4 1.3 1.4 1.5
Metasoma IV length 7.1 6.5 5.7 6.5 4.2 3.5 4.2 3.9
width 1.4 1.4 1.4 1.5 1.5 1.4 1.5 1.6
height 1.4 1.3 1.5 1.4 1.4 1.2 1.4 1.4
Metasoma V length 7.1 7.1 6.2 6.7 4.6 4.2 4.8 4.6
width 1.5 1.5 1.5 1.6 1.6 1.3 1.6 1.6
height 1.5 1.5 1.4 1.6 1.4 1.2 1.5 1.4
Tels on length 4.5 4.7 4.5 4.6 4.0 3.6 4.1 3.8
Vesicle length 3.3 3.3 3.1 3.1 2.4 2.0 2.4 2.8
width 1.3 1.6 1.3 1.3 1.2 1.6 1.2 1.1
height 1.4 1.4 1.2 1.4 1.1 1.0 1.0 1.1
Aculeus length 1.6 1.3 1.7 1.6 1.8 1.9 1.9 1.6
1 Sum of carapace, tergites I–VII, metasomal segments I–V, and telson; 2 distance between median ocelli; 3 sum of trochanter,
femur, patella, and chela; 4 measured from base of condyle to tip of xed nger; 5 sum of tergites I–VII; 6 sum of metasomal
segments I–V and telson.
TABLE 5 continued
savanna to mangroves and oaks near the coastline.
Specimens from Microondas Fogos were collected
on fence poles in rangeland at night. e habitat
and habitus are consistent with the arboreal, cor-
ticolous ecomorphotype (Prendini, 2001a).
M E: MEXICO: Guerrero:
Município Copala: Microondas Fogos,
16°33′59.5″N 98°53′18.1″W, 103 m, 22.vi.2007, O.F.
Francke, M. Escalante, H. Montaño, and A. Balles-
teros, 1 ♂ (AMNH [LP 7032]), 1 juv. ♂ (CNAN
SC3983), 14 m, 2.xi.2007, O.F. Francke, H. Mon-
taño, and A. Ballesteros, 3 ♂ (CNAN SC3978), 1 ♀
(AMNH [LP 8582]). Michoacán: Município Aquila:
Faro de Bucerias, 18°21′08.3″N 103°30′20.9″W, 13
m, 10.iii.2002, J. Ponce, low deciduous forest, 1 juv.
♀ (CNAN SC3982), 13.iv.2002, J. Ponce, low decid-
uous forest, 2 ♂ (CNAN SC3979, SC3980), 3 ♂
(CNAN SC4005), 18°35′50.5″N 103°30′04.3″W,
221 m, 14.iv.2002, J. Ponce and E. González, low
deciduous forest, 1 ♂ (AMNH [LP 2009]).
Município Aquila: La Llorona, el Faro, 18°20′17.2″N
103°29′49.2″W, 8 m, 6.v.2000, E. Miranda, beach
gap, 1 ♂ (CNAN SC3981).
Centruroides cuauhmapan, sp. nov.
Figures 2, 3, 5A, B, 8A, B, 11A, 12A, 17A, D,
18A, D, 19A, D, 20A, D, 21A, D, 22A, D, 23A,
D, 24A, D, 25A, D, 28, 29, tables 1, 3, 10
Centruroides schmidti: Francke, 2007: 69, 71, 72,
g. 1 (misidentication); Armas and Mar-
tín-Frías, 2008: 7–10, 12, 17, 19, 20, gs.
2–4, table XIV (misidentication).
T M: MEXICO: Oaxaca:
Município San Juan Bautista Tuxtepec: Holotype
♂ (CNAN T01396), 4 ♀ paratypes (CNAN
T01399–T01402), 17 km from San Juan Bautista
Tuxtepec, Cerro del Oro Dam, 17°59′55″N
96°15′47.2″W, 74 m, 23.v.1990, E. Barrera and A.
Cadena. Veracruz: Município Actopan: 2 ♂ para-
34 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 452
TABLE 6
Meristic data for type material of Centruroides catemacoensis, sp. nov.
Material deposited in the California Academy of Sciences (CASENT), San Francisco and the Colección Nacional
de Arácnidos (CNAN), Universidad Nacional Autónoma de México, Mexico City. Measurements follow Stahnke
(1970), Lamoral (1979), and Prendini (2001b).
Holotype Paratypes Paratypes
♂ ♂ ♀
CNAN CASENT CNAN CASENT CNAN
T01424 9073286 9073287 T01420 T01421 9073309 9073314 9073366 T01422 T01423
Total length146.6 36.4 35.7 41.8 40.7 31.8 33.1 30.1 29.6 32.4
Carapace length 3.6 2.9 2.8 3.4 3.3 3.0 3.0 3.0 2.7 3.2
ant. width 1.8 1.4 1.4 1.8 1.6 1.6 1.7 1.5 1.5 1.7
post. width 3.9 3.3 3.1 3.8 3.5 3.6 3.5 3.5 3.1 3.9
Median ocelli diameter 0.3 0.3 0.3 0.3 0.4 0.3 0.3 0.3 3.0 0.3
Interocular length20.3 0.3 0.3 0.3 0.3 0.3 0.3 0.3 2.9 0.3
Pedipalp length316.8 11.3 12.3 15.9 14.0 12.0 12.2 12.0 11.4 12.8
Trochanter length 1.5 1.1 1.2 1.6 1.3 1.3 1.3 1.3 1.2 1.3
Femur length 4.0 2.8 3.0 3.8 3.5 2.8 2.7 2.7 2.6 3.0
width 0.7 0.5 0.5 0.7 0.6 0.5 0.6 0.6 0.6 0.6
height 1.0 0.7 0.7 1.0 0.8 0.9 0.9 0.9 0.7 0.9
Patella length 4.6 3.3 3.3 4.2 3.7 3.2 3.2 3.2 3.0 3.5
width 1.0 0.7 0.7 0.8 0.7 0.8 0.8 0.8 0.8 0.9
height 1.6 1.1 1.1 1.3 1.3 1.4 1.3 1.3 1.3 1.4
Chela length46.7 4.1 4.8 6.3 5.5 4.7 5.0 4.8 4.6 5.0
Manus length 3.0 2.0 2.0 2.8 2.5 1.9 1.7 1.8 1.6 2.0
width 1.4 1.0 1.0 1.3 1.2 1.0 0.9 1.0 0.9 1.0
height 1.2 0.9 1.0 1.1 1.0 0.9 1.0 0.9 0.8 1.0
Mov. nger length 4.1 2.9 3.2 4.0 3.5 3.2 3.5 3.2 3.3 3.4
Leg I length 7.0 5.3 5.0 6.6 6.0 5.5 5.8 5.2 5.2 5.6
Pectines length 5.7 4.5 4.5 5.1 4.5 4.1 4.5 4.2 3.8 4.6
tooth count 14/13 14/14 14/14 13/14 14/14 14/13 14/14 13/13 13/13 14/14
Mesosoma length511.2 10.4 9.7 9.5 11.0 10.1 11.3 8.6 9.6 9.1
Sternite VII length 3.0 2.8 3.0 2.7 3.1 2.7 2.8 2.3 2.6 2.5
width 3.6 2.9 3.0 3.2 3.0 3.4 3.6 3.4 3.4 3.4
Metasoma length631.8 23.1 23.2 28.9 26.4 18.7 18.8 18.5 17.3 20.1
Metasoma I length 4.0 3.0 3.1 3.6 3.3 2.5 2.2 2.5 2.3 2.5
width 1.4 1.2 1.2 1.4 1.3 1.4 1.4 1.3 1.4 1.5
height 1.4 1.2 1.3 1.3 1.3 1.5 1.5 1.5 1.4 1.7
Metasoma II length 5.1 3.5 3.5 4.6 4.2 2.8 2.8 2.9 2.5 3.0
width 1.4 1.2 1.3 1.4 1.3 1.4 1.5 1.5 1.3 1.6
height 1.3 1.2 1.2 1.2 1.2 1.4 1.3 1.3 1.4 1.5
2021 GOODMAN ET AL.: REVISION OF “THORELLII” CLADE OF CENTRUROIDES 35
Holotype Paratypes Paratypes
♂ ♂ ♀
CNAN CASENT CNAN CASENT CNAN
T01424 9073286 9073287 T01420 T01421 9073309 9073314 9073366 T01422 T01423
Metasoma III length 5.5 4.0 4.1 5.1 4.7 3.1 3.1 3.0 2.8 3.4
width 1.4 1.2 1.2 1.4 1.3 1.5 1.5 1.5 1.4 1.7
height 1.3 1.2 1.2 1.3 1.2 1.3 1.4 1.3 1.2 1.6
Metasoma IV length 6.2 4.5 4.4 5.5 5.3 3.4 3.6 3.4 3.0 3.7
width 1.4 1.2 1.2 1.3 1.3 1.4 1.5 1.4 1.3 1.7
height 1.4 1.1 1.2 1.2 1.3 1.4 1.4 1.3 1.2 1.6
Metasoma V length 6.7 4.7 4.6 5.9 5.3 3.7 3.7 3.7 3.6 4.0
width 1.5 1.3 1.2 1.4 1.3 1.4 1.5 1.4 1.3 1.6
height 1.4 1.2 1.2 1.3 1.2 1.3 1.4 1.3 1.2 1.6
Tels on length 4.3 3.4 3.5 4.2 3.6 3.2 3.4 3.0 3.1 3.5
Vesicle length 3.1 2.3 2.3 2.9 2.5 1.9 2.0 1.9 1.8 2.2
width 1.4 1.0 1.0 1.1 1.1 1.4 1.0 1.0 1.0 1.0
height 1.3 1.0 0.9 1.1 1.0 1.0 1.0 1.1 0.9 1.2
Aculeus length 1.2 1.2 1.0 1.3 1.2 1.0 1.5 1.3 1.3 1.5
1 Sum of carapace, tergites I–VII, metasomal segments I–V, and telson; 2 distance between median ocelli; 3 sum of trochanter,
femur, patella, and chela; 4 measured from base of condyle to tip of xed nger; 5 sum of tergites I–VII; 6 sum of metasomal
segments I–V and telson.
TABLE 6 continued
types (CNAN T01397, T01398), Los Idolos,
19°25′44.9″N 96°32′12.4″W, 112 m, 5.v.2006,
O.F. Francke, P. Berea, and J. Ballesteros, col-
lected with UV detection.
E: e species name is a noun in
apposition, taken from the Nahuatl word mean-
ing “up in a tree” and alludes to the arboreal
habitat of species in the genus.
D: Centruroides cuauhmapan is most
closely related to C. rileyi, from which it diers as
follows. e posterosubmedian carinae on the
carapace are absent in C. cuauhmapan (g. 5D)
but weakly developed in C. rileyi (g. 5A). e
retrodorsal carina of the pedipalp chela manus is
nely granular, the dorsomedian carina distinct,
granular, and the prodorsal carina distinct, granu-
lar and complete in the male of C. cuauhmapan
(gs. 11, 12B), whereas the retrodorsal carina is
smooth, the dorsomedian carina weakly granular,
and the prodorsal carina weakly granular and
restricted to the distal half of the segment in the
male of C. rileyi (gs. 11, 12A). e ventrolateral
and ventrosubmedian carinae of mesosomal ster-
nite VII are distinct, granular and the intercarinal
surfaces finely granular in C. cuauhmapan,
whereas the ventrolateral and ventrosubmedian
carinae are obsolete to absent and the intercarinal
surfaces smooth in C. rileyi. e metasoma and
telson are longer in the male and more robust,
proportionally longer and broader, in the female
of C. cuauhmapan than C. rileyi. e ventrolateral
and ventrosubmedian carinae are distinct, granu-
lar on metasomal segments I–V in the male and
female of C. cuauhmapan (gs. 18G, J, 19G, J,
21G, J, 22G, J), but granular on segment I in the
female, vestigial on I–III, and smooth on IV and
V in the male of C. rileyi (gs. 18A, D, 19A, D,
21A, D, 22A, D). e surfaces of the telson vesicle
of the female are granular in C. cuauhmapan (gs.
23–25J), but smooth in C. rileyi (gs. 23–25D).
36 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 452
D: e following description is
based on the holotype male, with dierences
among other material noted in the section on
variation.
Coloration: Base color pale yellow, with exten-
sive infuscation, creating mottled or marbled
pattern. Carapace with uniformly infuscate mar-
bling, more densely infuscate medially. Pedipalp
chela ngers and manus, dorsal and retrolateral
intercarinal surfaces with moderately infuscate
marbling; prolateral and ventral intercarinal sur-
faces immaculate. Legs retrolateral surfaces with
infuscate marbling; prolateral surfaces pale,
immaculate. Tergites with unformly infuscate
mottling, pale stripe medially, blackish spots sub-
medially, and distinct, narrow bands laterally.
Sternites slightly infuscate posteriorly, with
faintly infuscate triangular marking at posterior
margin of sternite III, fading to infuscate mot-
tling on sternite VII. Metasomal segments uni-
formly, faintly marbled; segment V and telson
markedly infuscate, noticeably darker than pre-
ceding segments.
Carapace: Shape trapezoidal; anterior width
four-hs of posterior width (table 3); anterome-
dian sulcus moderately deep, oval; posterome-
dian sulcus shallow anteriorly, deeper posteriorly;
carinae moderately developed, comprising small
to medium-sized granules (g. 5C).
Pedipalps: Orthobothriotaxic, Type A; femur
dorsal trichobothria with α conguration; pedi-
palp chela xed nger, trichobothrium db situ-
ated slightly distal to et. Femoral carinae
granular; dorsal intercarinal surface moderately
FIGURE 15. Centruroides Marx, 1890, pedipalp chela, retrolateral aspect. A. C. homanni Armas, 1996, ♂
(CNAN SC3996). B. C. chanae, sp. nov., holotype ♂ (CNAN T01403). C. C. yucatanensis, sp. nov., holotype
♂ (CNAN T01416). Scale bars = 2 mm.
2021 GOODMAN ET AL.: REVISION OF “THORELLII” CLADE OF CENTRUROIDES 37
granular; pro- and retrolateral intercarinal sur-
faces each with series of large spiniform gran-
ules. Patella carinae granular; prolateral
intercarinal surface with eight to 10 large subspi-
niform granules. Chela manus slightly incrassate;
dorsal secondary carina well developed, nely
serrate; digital and retrolateral secondary carinae
moderately developed, nely crenulate; retrodor-
sal carina well developed, coarsely crenulate; ret-
roventral carina well developed, nely crenulate;
proventral carina moderately developed, com-
prising few rounded granules; prodorsal carina
well developed, coarsely serrate. Fixed and mov-
able ngers each shallowly curved proximally.
Fixed nger, median denticle row comprising
eight oblique subrows, each anked by pro- and
retrolateral supernumerary denticles. Movable
nger, median denticle row with short terminal
subrow comprising four denticles preceded by
eight oblique subrows, each anked by pro- and
retrolateral supernumerary denticles.
Legs: Leg I length 1.92× greater than carapace
length (table 10). Telotarsi ventral surfaces densely
covered with short setae; ungues strongly curved.
Pectines: Pectinal plate 1.8× wider than long;
posterior margin distinctly rounded; pectinal
tooth count 14/14 (♂) (g. 8C, table 3).
Mesosoma: Tergites width similar to carapace
posterior width; I and II slightly narrower (table
3). Pretergites surfaces smooth to nely granular.
Posttergites surfaces weakly granular; I–VI with
dorsomedian carinae vestigial and reduced to
FIGURE 16. Centruroides Marx, 1890, pedipalp chela, retrolateral aspect. A. C. homanni Armas, 1996, ♀
(CNAN SC3996). B. C. chanae, sp. nov., paratype ♀ (CNAN T01405). C. C. yucatanensis, sp. nov., paratype
♀ (CNAN T01418). Scale bars = 1 mm.
38 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 452
FIGURE 17. Centruroides Marx, 1890, metasomal segments I and II, dorsal aspect. A, D. C. rileyi Sissom,
1995, A. ♂ (CNAN SC4002), D. ♀ (CNAN SC4003). B, E. C. hamadryas, sp. nov., B. holotype ♂ (CNAN
T01408), E. paratype ♀ (CNAN T01415). C, F. C. homanni Armas, 1996, C. ♂, F. ♀ (CNAN SC3996). G,
J. C. cuauhmapan, sp. nov., G. holotype ♂ (CNAN T01396), J. paratype ♀ (CNAN T01399). H, K. C. berstoni,
sp. nov., H. holotype ♂ (CASENT 9073325), K. paratype ♀ (CASENT 9073313). I, L. C. chanae, sp. nov., I.
holotype ♂ (CNAN T01403), L. paratype ♀ (CNAN T01405). M, P. C. catemacoensis, sp. nov., M. holotype
♂ (CNAN T01424), P. paratype ♀ (CNAN T01423). N, Q. C. schmidti Sissom, 1995, N. ♂ (CASENT 9073316),
Q. ♀ (CASENT 9073317). O, R. C. yucatanensis, sp. nov., O. holotype ♂ (CNAN T01416), R. paratype ♀
(CNAN T01417. Scale bars = 2 mm.
2021 GOODMAN ET AL.: REVISION OF “THORELLII” CLADE OF CENTRUROIDES 39
FIGURE 18. Centruroides Marx, 1890, metasomal segments I and II, ventral aspect. A, D. C. rileyi Sissom,
1995, A. ♂ (CNAN SC4002), D. ♀ (CNAN SC4003). B, E. C. hamadryas, sp. nov., B. holotype ♂ (CNAN
T01408), E. paratype ♀ (CNAN T01415). C, F. C. homanni Armas, 1996, C. ♂, F. ♀ (CNAN SC3996). G,
J. C. cuauhmapan, sp. nov., G. holotype ♂ (CNAN T01396), J. paratype ♀ (CNAN T01399). H, K. C. berstoni,
sp. nov., H. holotype ♂ (CASENT 9073325), K. paratype ♀ (CASENT 9073313). I, L. C. chanae, sp. nov., I.
holotype ♂ (CNAN T01403), L. paratype ♀ (CNAN T01405). M, P. C. catemacoensis, sp. nov., M. holotype
♂ (CNAN T01424), P. paratype ♀ (CNAN T01423). N, Q. C. schmidti Sissom, 1995, N. ♂ (CASENT 9073316),
Q. ♀ (CASENT 9073317). O, R. C. yucatanensis, sp. nov., O. holotype ♂ (CNAN T01416), R. paratype ♀
(CNAN T01417. Scale bars = 2 mm.
40 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 452
FIGURE 19. Centruroides Marx, 1890, metasomal segments I and II, lateral aspect. A, D. C. rileyi Sissom, 1995,
A. ♂ (CNAN SC4002), D. ♀ (CNAN SC4003). B, E. C. hamadryas, sp. nov., B. holotype ♂ (CNAN T01408),
E. paratype ♀ (CNAN T01415). C, F. C. homanni Armas, 1996, C. ♂, F. ♀ (CNAN SC3996). G, J. C. cuauh-
mapan, sp. nov., G. holotype ♂ (CNAN T01396), J. paratype ♀ (CNAN T01399). H, K. C. berstoni, sp. nov.,
H. holotype ♂ (CASENT 9073325), K. paratype ♀ (CASENT 9073313). I, L. C. chanae, sp. nov., I. holotype
♂ (CNAN T01403), L. paratype ♀ (CNAN T01405). M, P. C. catemacoensis, sp. nov., M. holotype ♂ (CNAN
T01424), P. paratype ♀ (CNAN T01423). N, Q. C. schmidti Sissom, 1995, N. ♂ (CASENT 9073316), Q. ♀
(CASENT 9073317). O, R. C. yucatanensis, sp. nov., O. holotype ♂ (CNAN T01416), R. paratype ♀ (CNAN
T01417). Scale bars = 2 mm.
2021 GOODMAN ET AL.: REVISION OF “THORELLII” CLADE OF CENTRUROIDES 41
several small granules; VII surface nely granu-
lar, dorsomedian and dorsosubmedian carinae
present, dorsolateral carinae nely serrate. Ster-
nites III–VI, surfaces smooth; VII surface weakly
granular, ventrolateral carinae serrate.
Metasoma: Metasoma length 2.86× mesosoma
length (table 3). Segments longer than wide;
increasing in length posteriorly, segment V 2×
length of I; carinae distinct, granular; intercari-
nal surfaces sparsely granular (gs. 20–22G).
Tels o n : Vesicle elongate, ovoid; ventral surface
shallowly convex, moderately granular; ventro-
median carina granular, terminating at subacu-
lear tubercle; subaculear tubercle narrow and
angular in lateral aspect, directed toward mid-
point of aculeus. Aculeus angled ventrally at
slightly less than 90° (gs. 23–25G).
Variation: Adult males and females dier as
follows. e dorsomedian carinae of the pedi-
palp patella are absent, the pectinal tooth count
slightly higher (13 or 14), the mesosoma propor-
tionally longer and slenderer, and the metasoma
longer, in males (gs. 28A, B, 29A, B, table 3).
e rst pair of legs are shorter and stouter, the
pectinal tooth count slightly lower (11–13), and
the metasomal carinae more developed and
nely serrate in females (gs. 8C, D, 17G, J, 18G,
J, 19G, J, 20G, J, 21G, J, 22G, J, 23G, J, 24G, J,
25G, J, table 3).
D: Centruroides cuauhmapan is
endemic to eastern Mexico and recorded from
two localities in the state of Veracruz and a third,
approximately 200 km south, in the state of Oax-
aca (g. 3).
E: e localities at which C. cuauh-
mapan has been recorded range in altitude from
74 to 555 m. e habitat at these localities varies
from subtropical highland forest to humid sub-
tropical forest. One individual was collected in a
coee plantation in lowland rainforest. e habi-
tat and habitus are consistent with the arboreal,
corticolous ecomorphotype (Prendini, 2001a).
R: Specimens from Córdoba, Vera-
cruz, were misidentied as C. schmidti by Armas
and Frías (2008). is species has not been
recorded from Mexico.
M E: MEXICO: Oaxaca:
Município San Juan Bautista Tuxtepec: Cerro del
Oro Dam, 17 km from San Juan Bautista Tuxte-
pec, 17°59′55″N 96°15′47.2″W, 74 m, 23.v.1990,
E. Barrera and A. Cadena, 1 ♂ (CNAN SC4001).
Veracruz: Município Amatlán de los Reyes:
Cañada Blanca, 18°55′43.5″N 96°51′26″W, 555
m, 18.vii.2002, E. González, found in coee plan-
tation in lowland rainforest, collected at night
with UV light, 1 ♂ (AMNH [LP 2073]).
Centruroides hamadryas, s p. nov.
Figures 2, 4, 6A, B, 9A, B, 13A, 14A, 17B, E,
18B, E, 19B, E, 20B, E, 21B, E, 22B, E, 23B, E,
24B, E, 25B, E, 32, 33, tables 1, 4, 10
T M: MEXICO: Chiapas:
Município Ocosingo: Holotype ♂ (CNAN
T01408), paratype ♂ (CNAN T01412), paratype
♀ (CNAN T01413), La Galleta, 16°48′18.5″N
90°54′25″W, 103 m, 2.v.2005 A. Valdez, O.F.
Francke, and A. Ballesteros, collected with UV
light detection; paratype ♂ (CNAN T01414),
paratype ♀ (CNAN T01415), same data, except:
2.v.1992, E. Barrera; 2 ♂ paratypes (CNAN
T01409, T01411), paratype ♀ (CNAN T01410),
same data, except: 114 m, 28.iv.2005, urban area
toward blue water bridge.
E: e species name is noun in
apposition, taken from the Greek nymph Hama-
dryas, mother of the hamadryads, tree-dwelling
nymphs with lifelong bonds to the trees.
D: Centruroides hamadryas is most
closely related to C. berstoni, from which it dif-
fers in the following respects. e carapace is
densely granular, with distinct lateral ocular cari-
nae, in the female of C. hamadryas (g. 6B) but
sparsely granular, more densely so on the
interocular triangle, in the female of C. berstoni
(g. 6D). e pedipalp chela manus of the male
is proportionally more incrassate in C. hama-
dryas (g. 13A) than C. berstoni (g. 13B). e
legs of the male are less than 2× the length of the
carapace in C. hamadryas but greater than 2× the
length of the carapace in C. berstoni (table 10).
42 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 452
TABLE 7
Meristic data for Centruroides schmidti Sissom, 1995
Material deposited in the California Academy of Sciences (CASENT), San Francisco. Measurements follow
Stahnke (1970), Lamoral (1979), and Prendini (2001b).
♂ ♀
CASENT 9073316 CASENT 9073317
Total length147.9 47.4 35.3 37.7 34.8 33.3 34.9 33.8
Carapace length 3.6 3.6 3.0 2.9 3.3 3.2 3.3 3.2
ant. width 1.8 1.8 1.5 1.5 1.7 1.7 1.7 1.7
post. width 3.9 3.9 3.3 3.2 3.7 3.6 3.8 3.7
Median ocelli diameter 0.3 0.3 0.3 0.3 0.3 0.3 0.3 0.3
Interocular length20.3 0.4 0.3 0.3 0.4 0.4 0.3 0.3
Pedipalp length317.0 16.3 13.2 13.3 14.2 13.8 14.1 13.8
Trochanter length 1.6 1.5 1.3 1.3 1.4 1.3 1.4 1.3
Femur length 4.1 3.9 3.0 3.1 3.3 3.1 3.2 3.2
width 0.7 0.7 0.6 0.5 0.7 0.6 0.7 0.7
height 1.0 1.0 0.9 0.8 1.0 1.0 1.0 1.0
Patella length 4.5 4.2 3.7 3.5 3.7 3.6 3.7 3.5
width 1.0 1.0 0.8 0.8 1.0 1.0 0.9 1.0
height 1.6 1.6 1.3 1.3 1.5 1.5 1.5 1.5
Chela length46.8 6.7 5.2 5.4 5.8 5.8 5.8 5.8
Manus length 2.6 2.7 2.2 2.3 2.2 2.1 2.1 2.3
width 1.5 1.4 1.0 1.1 1.1 1.2 1.2 1.2
height 1.3 1.3 1.0 1.1 1.2 1.1 1.2 1.1
Mov. nger length 4.2 4.3 3.5 3.1 4.0 4.0 4.1 4.0
Leg I length 7.0 6.8 5.6 5.6 6.1 6.0 6.0 6.1
Pectines length 6.2 6.2 4.6 4.7 4.3 4.5 5.0 5.0
tooth count 15/15 16/15 14/14 15/15 12/12 13/13 14/14 14/14
Mesosoma length511.8 11.6 8.8 10.1 11.7 10.4 11.2 11.6
Sternite VII length 3.4 3.3 2.4 2.7 3.0 2.5 3.0 2.6
width 3.5 3.4 2.9 2.9 4.1 3.8 4.1 4.1
Metasoma length632.5 32.2 23.5 24.7 19.8 19.7 20.4 19.0
Metasoma I length 4.2 4.1 3.0 3.1 2.1 2.4 2.3 2.2
width 1.4 1.4 1.4 1.2 1.7 1.7 1.8 1.6
height 1.4 1.5 1.4 1.3 1.5 1.5 1.6 1.5
Metasoma II length 5.1 5.0 3.7 3.8 3.0 2.9 3.1 3.0
width 1.4 1.4 1.3 1.2 1.6 1.5 1.6 1.5
height 1.3 1.3 1.2 1.2 1.5 1.5 1.5 1.3
Metasoma III length 5.7 5.7 4.2 4.0 3.3 3.2 3.2 3.2
width 1.3 1.4 1.3 1.2 1.4 1.4 1.6 1.5
height 1.2 1.3 1.2 1.1 1.6 1.4 1.4 1.4
2021 GOODMAN ET AL.: REVISION OF “THORELLII” CLADE OF CENTRUROIDES 43
♂ ♀
CASENT 9073316 CASENT 9073317
Metasoma IV length 6.5 6.2 4.5 4.7 3.8 3.7 3.8 3.7
width 1.3 1.3 1.2 1.1 1.4 1.4 1.5 1.4
height 1.2 1.2 1.1 1.1 1.5 1.4 1.4 1.3
Metasoma V length 7.0 6.7 4.7 5.5 4.1 4.0 4.4 3.7
width 1.3 1.3 1.2 1.2 1.4 1.4 1.5 1.4
height 1.3 1.3 1.2 1.1 1.5 1.5 1.4 1.3
Tels on length 4.0 4.5 3.4 3.6 3.5 3.5 3.6 3.2
Vesicle length 3.0 3.2 2.3 2.5 2.2 2.1 2.2 2.1
width 1.3 1.2 0.9 1.0 0.9 1.0 1.1 1.3
height 1.2 1.3 0.9 0.9 1.0 1.0 1.2 1.1
Aculeus length 1.3 1.5 1.1 1.3 1.5 1.5 1.4 1.0
1 Sum of carapace, tergites I–VII, metasomal segments I–V, and telson; 2 distance between median ocelli; 3 sum of trochanter,
femur, patella, and chela; 4 measured from base of condyle to tip of xed nger; 5 sum of tergites I–VII; 6 sum of metasomal
segments I–V and telson.
TABLE 7 continued
e dorsomedian carinae of the mesosomal ter-
gites are distinct and complete in C. hamadryas
but weakly developed and restricted to the pos-
terior half of the segments in C. berstoni. e
ventrolateral and ventrosubmedian carinae of
mesosomal sternite VII are weakly developed to
absent in C. hamadryas, whereas the ventrolat-
eral carinae are weakly developed and the ven-
trosubmedian carinae absent in C. berstoni. e
telson vesicle is densely setose in C. hamadryas
but sparsely setose in C. berstoni.
Additional dierences between C. hamadryas
and other species of the clade are as follows. e
retrodorsal carina of the chela manus is complete
and the dorsomedian carina weakly developed
and restricted to the distal half, in the male of C.
hamadryas (g. 13A), whereas the retrodorsal
carina is weakly granular and the dorsomedian
carina absent in the male of C. berstoni (g. 13B),
C. catemacoensis (g. 11B), C. cuauhmapan (g.
11C), and C. rileyi (g. 11A). e retrodorsal
carina of the manus of the female is complete
and the prodorsal carina restricted to the distal
third in the female of C. hamadryas (g. 13B),
whereas the retrodorsal carina is nely granular
and the prodorsal carina absent in the female of
C. berstoni (g. 14B), C. catemacoensis (g. 12B),
C. cuauhmapan (g. 12C), and C. rileyi (g.
12A). e pedipalp chela ngers bear short,
dense setation in C. hamadryas but sparse seta-
tion in C. berstoni. e dorsosubmedian and
dorsolateral carinae of metasomal segments I–III
are well developed and granular in the male of C.
hamadryas (gs. 17–19E) but weakly developed
to absent in the male of C. berstoni (gs. 17–19K).
e telson vesicle of the male is not posteriorly
bilobed in C. hamadryas (gs. 23–25B), unlike C.
C. cuauhmapan (gs. 23–25G) and C. rileyi (gs.
23–25A). e vesicle of the female is shorter and
more robust, with intercarinal surfaces smooth
in C. hamadryas (gs. 23–25E), but sparsely
granular in C. berstoni (gs. 23–25K) and densely
granular in C. catemacoensis (gs. 23–25P), C.
cuauhmapan (gs. 23–25J), and C. rileyi (gs.
23–25D).
D: e following description is
based on the holotype male, with dierences
among other material noted in the section on
variation.
Coloration: Base color pale yellow, with exten-
sive infuscation, creating mottled or marbled
pattern. Carapace with uniformly infuscate mar-
44 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 452
FIGURE 20. Centruroides Marx, 1890, metasomal segment V, dorsal aspect. A, D. C. rileyi Sissom, 1995, A.
♂ (CNAN SC4002), D. ♀ (CNAN SC4003). B, E. C. hamadryas, sp. nov., B. holotype ♂ (CNAN T01408), E.
paratype ♀ (CNAN T01415). C, F. C. homanni Armas, 1996, C. ♂, F. ♀ (CNAN SC3996). G, J. C. cuauh-
mapan, sp. nov., G. holotype ♂ (CNAN T01396), J. paratype ♀ (CNAN T01399). H, K. C. berstoni, sp. nov.,
H. holotype ♂ (CASENT 9073325), K. paratype ♀ (CASENT 9073313). I, L. C. chanae, sp. nov., I. holotype
♂ (CNAN T01403), L. paratype ♀ (CNAN T01405). M, P. C. catemacoensis, sp. nov., M. holotype ♂ (CNAN
T01424), P. paratype ♀ (CNAN T01423). N, Q. C. schmidti Sissom, 1995, N. ♂ (CASENT 9073316), Q. ♀
(CASENT 9073317). O, R. C. yucatanensis, sp. nov., O. holotype ♂ (CNAN T01416), R. paratype ♀ (CNAN
T01417). Scale bars = 2 mm.
2021 GOODMAN ET AL.: REVISION OF “THORELLII” CLADE OF CENTRUROIDES 45
FIGURE 21. Centruroides Marx, 1890, metasomal segment V, ventral aspect. A, D. C. rileyi Sissom, 1995, A.
♂ (CNAN SC4002), D. ♀ (CNAN SC4003). B, E. C. hamadryas, sp. nov., B. holotype ♂ (CNAN T01408), E.
paratype ♀ (CNAN T01415). C, F. C. homanni Armas, 1996, C. ♂, F. ♀ (CNAN SC3996). G, J. C. cuauh-
mapan, sp. nov., G. holotype ♂ (CNAN T01396), J. paratype ♀ (CNAN T01399). H, K. C berstoni, sp. nov.,
H. holotype ♂ (CASENT 9073325), K. paratype ♀ (CASENT 9073313). I, L. C. chanae, sp. nov., I. holotype
♂ (CNAN T01403), L. paratype ♀ (CNAN T01405). M, P. C. catemacoensis, sp. nov., M. holotype ♂ (CNAN
T01424), P. paratype ♀ (CNAN T01423). N, Q. C. schmidti Sissom, 1995, N. ♂ (CASENT 9073316), Q. ♀
(CASENT 9073317). O, R. C. yucatanensis, sp. nov., O. holotype ♂ (CNAN T01416), R. paratype ♀ (CNAN
T01417). Scale bars = 2 mm.
46 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 452
FIGURE 22. Centruroides Marx, 1890, metasomal segment V, lateral aspect. A, D. C. rileyi Sissom, 1995, A.
♂ (CNAN SC4002), D. ♀ (CNAN SC4003). B, E. C. hamadryas, sp. nov., B. holotype ♂ (CNAN T01408), E.
paratype ♀ (CNAN T01415). C, F. C. homanni Armas, 1996, C. ♂, F. ♀ (CNAN SC3996). G, J. C. cuauh-
mapan, sp. nov., G. holotype ♂ (CNAN T01396), J. paratype ♀ (CNAN T01399). H, K. C. berstoni, sp. nov.,
H. holotype ♂ (CASENT 9073325), K. paratype ♀ (CASENT 9073313). I, L. C. chanae, sp. nov., I. holotype
♂ (CNAN T01403), L. paratype ♀ (CNAN T01405). M, P. C. catemacoensis, sp. nov., M. holotype ♂ (CNAN
T01424), P. paratype ♀ (CNAN T01423). N, Q. C. schmidti Sissom, 1995, N. ♂ (CASENT 9073316), Q. ♀
(CASENT 9073317). O, R. C. yucatanensis, sp. nov., O. holotype ♂ (CNAN T01416), R. paratype ♀ (CNAN
T01417). Scale bars = 2 mm.
2021 GOODMAN ET AL.: REVISION OF “THORELLII” CLADE OF CENTRUROIDES 47
bling, more densely infuscate medially. Pedipalp
chela ngers and manus, dorsal and retrolateral
intercarinal surfaces with moderately infuscate
marbling; prolateral and ventral intercarinal sur-
faces mostly immaculate. Legs retrolateral sur-
faces with infuscate marbling; prolateral surfaces
pale, immaculate. Tergites with unformly infus-
cate mottling, pale stripe medially, blackish spots
submedially, and faint, narrow bands laterally.
Sternites pale, with faintly infuscate triangular to
trapezoidal marking at posterior margin of ster-
nite III, fading to infuscate mottling on sternite
VII. Metasomal segments uniformly, faintly mar-
bled; segment V and telson markedly infuscate,
noticeably darker than preceding segments.
Carapace: Shape trapezoidal; anterior width
four-hs of posterior width (table 4); anterome-
dian sulcus moderately deep, oval; posterome-
dian sulcus shallow anteriorly, deeper posteriorly;
median ocular tubercle moderately granular;
carinae weakly developed, comprising small to
medium-sized granules (g. 6A).
Pedipalps: Orthobothriotaxic, Type A;
femur dorsal trichobothria with α configura-
tion; pedipalp chela fixed finger, trichoboth-
rium db situated slightly distal to et. Femoral
carinae serrate; retromedian carinae compris-
ing spiniform granules; dorsal intercarinal sur-
face moderately granular; prolateral surface
with series of large spiniform granules. Patella
prodorsal, dorsomedian, retrodorsal and pro-
ventral carinae moderately developed, serrate;
retromedian carina well developed, serrate;
retroventral carina incomplete, serrate; prolat-
eral intercarinal surface with five or six large,
subspiniform granules. Chela manus proven-
tral carina moderately developed, comprising
few rounded granules; other carinae weakly
developed, granular. Fixed finger, median den-
ticle row comprising eight oblique subrows,
each flanked by pro- and retrolateral supernu-
merary denticles. Movable finger, median den-
ticle row with short terminal row comprising
four denticles preceded by eight oblique sub-
rows, each flanked by pro- and retrolateral
supernumerary denticles.
Legs: Leg I length 1.88× greater than carapace
length (table 10). Telotarsi ventral surfaces
densely covered with short setae; ungues mark-
edly curved.
Pectines: Pectinal plate 1.9× wider than long;
posterior margin distinctly rounded; pectinal
tooth count 14/14 (♂) (g. 6A, table 4).
Mesosoma: Tergites width similar to carapace
posterior width; I and II slightly narrower (table 4).
Pretergites surfaces smooth to nely granular. Post-
tergites surfaces weakly granular; I–VI with dorso-
median carinae moderately granular; VII surface
weakly granular, dorsomedian carina moderately
granular, dorsosubmedian carinae serrate, dorsolat-
eral carinae well developed. Sternites III–VI, sur-
faces smooth; VII surface weakly granular,
ventrolateral carinae reduced to few granules.
Metasoma: Metasoma length 3.01× mesosoma
length (table 4). Segments longer than wide;
increasing in length posteriorly, segment V 2×
length of I; carinae complete, granular on segments
I–III, other carinae absent or obsolete; intercarinal
surfaces sparsely granular (gs. 17–22E).
Tels o n : Vesicle elongate, ovoid; ventral surface
shallowly convex, sparsely granular posteriorly;
ventromedian carina granular, terminating at
subaculear tubercle; subaculear tubercle narrow
and angular in lateral aspect, directed toward
midpoint of aculeus. Aculeus angled ventrally at
slightly less than 90° (g. 25B, E).
Variation: Adult males and females dier as
follows. e pectinal tooth count is slightly
higher (14 or 15), the mesosoma proportionally
longer and slenderer, and the metasoma up to
3× longer, with segment V also roughly 1.5 mm
longer, in males (gs. 23B, E, 24B, E, 25B, E,
table 4). e tegument is more densely infus-
cate, the pectinal plate produced into a rounded
lobe posteriorly, which is punctate and slightly
infuscate, the pectinal tooth count slightly
lower (12 or 13), and the telson shorter and
narrower, in females (gs. 9A, B, 23B, E, 24B, E,
25B, E, 32A, B, 33A, B, table 4).
D: Centruroides hamadryas is
known only from the state of Chiapas in south-
eastern Mexico, but may extend across the Usu-
48 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 452
TABLE 8
Meristic data for Centruroides homanni Armas, 1996
Material deposited in the Colección Nacional de Arácnidos (CNAN), Universidad Nacional Autónoma de
México, Mexico City. Measurements follow Stahnke (1970), Lamoral (1979), and Prendini (2001b).
♂ ♀
CNAN CNAN
SC3993 SC3996 SC3997 SC3998 SC3993 SC3996
Total length137.1 55.9 38.4 40.5 42.7 35.5 33.1 34.8 33.7
Carapace length 3.1 3.9 3.3 3.2 3.3 3.4 3.4 3.3 3.4
ant. width 1.5 1.9 1.7 1.5 1.6 1.8 1.9 1.8 1.8
post. width 3.4 4.0 3.6 3.3 3.3 3.8 3.9 3.9 3.7
Median ocelli diameter 0.3 0.3 0.3 0.3 0.3 0.3 0.3 0.3 0.3
Interocular length20.3 0.4 0.4 0.3 0.3 0.3 0.3 0.4 0.4
Pedipalp length312.8 18.1 14.0 14.2 15.1 14.0 14.2 14.6 14.3
Trochanter length 1.1 1.6 1.3 1.4 1.3 1.4 1.4 1.5 1.4
Femur length 2.6 4.2 3.2 3.4 3.6 3.2 3.3 3.3 3.3
width 0.5 0.7 0.6 0.6 0.6 0.8 0.8 0.7 0.9
height 0.9 1.1 0.9 0.9 1.0 1.0 1.0 1.0 1.0
Patella length 3.6 4.9 3.6 3.8 4.0 3.7 3.7 3.8 3.9
width 0.8 1.0 0.8 0.9 0.8 1.0 1.0 1.0 1.2
height 1.2 1.7 1.4 1.4 1.3 1.4 1.5 1.5 1.6
Chela length45.5 7.4 6.0 5.7 6.2 5.7 5.8 6.0 5.7
Manus length 2.4 3.4 2.5 2.5 2.8 2.1 2.3 2.3 2.4
width 1.1 1.5 1.2 1.1 1.3 1.1 1.3 1.1 1.2
height 1.0 1.5 1.2 1.1 1.2 1.0 1.1 1.1 1.3
Mov. nger length 3.5 4.7 3.6 3.4 3.8 4.0 4.0 4.0 4.0
Leg I length 5.6 7.5 5.8 6.0 6.0 6.1 6.4 6.2 5.8
Pectines length 4.8 6.5 5.0 4.7 4.8 4.7 4.5 4.5 4.5
tooth count 14/14 15/15 14/14 15/15 13/13 15/14 13/13 13/14 13/13
Mesosoma length510.1 13.1 10.4 9.8 10.6 11.7 9.4 11.4 10.5
Sternite VII length 2.8 4.0 3.0 3.1 2.9 2.8 2.6 2.9 2.6
width 3.0 3.9 3.1 3.0 3.1 4.3 4.0 4.3 4.1
Metasoma length623.9 38.9 24.7 27.5 28.8 20.4 20.3 20.1 19.8
Metasoma I length 3.1 4.8 3.1 3.5 3.5 2.5 2.7 2.5 2.3
width 1.3 1.6 1.3 1.3 1.2 1.7 1.8 1.8 1.8
height 1.3 1.5 1.3 1.2 1.2 1.5 1.5 1.5 1.5
Metasoma II length 3.3 6.8 3.9 4.2 4.4 3.3 3.1 3.2 3.2
width 1.2 1.4 1.2 1.2 1.2 1.5 1.6 1.5 1.6
height 1.2 1.4 1.2 1.1 1.1 1.5 1.5 1.5 1.7
2021 GOODMAN ET AL.: REVISION OF “THORELLII” CLADE OF CENTRUROIDES 49
♂ ♀
CNAN CNAN
SC3993 SC3996 SC3997 SC3998 SC3993 SC3996
Metasoma III length 4.2 7.1 4.1 5.0 5.2 3.6 3.4 3.4 3.4
width 1.2 1.4 1.2 1.1 1.2 1.5 1.5 1.5 1.5
height 1.2 1.3 1.2 1.1 1.2 1.4 1.5 1.5 1.5
Metasoma IV length 4.8 7.7 5.1 5.5 5.8 4.0 3.8 3.7 3.8
width 1.2 1.3 1.2 1.1 1.1 1.4 1.4 1.3 1.6
height 1.2 1.3 1.1 1.1 1.1 1.3 1.4 1.3 1.5
Metasoma V length 5.2 8.2 5.4 5.7 6.2 4.1 4.3 4.3 4.3
width 1.2 1.5 1.2 1.2 1.2 1.5 1.4 1.3 1.5
height 1.2 1.4 1.2 1.1 1.1 1.3 1.3 1.4 1.4
Tels on length 3.3 4.3 3.1 3.6 3.7 2.9 3.0 3.0 2.8
Vesicle length 2.3 3.1 2.1 2.6 2.7 1.7 1.8 1.7 1.7
width 1.0 1.4 1.1 0.9 1.0 1.2 1.1 1.0 1.0
height 1.0 1.4 1.0 0.9 1.0 1.0 1.0 1.0 1.0
Aculeus length 1.2 1.3 1.2 1.1 1.0 1.2 1.2 1.3 1.3
1 Sum of carapace, tergites I–VII, metasomal segments I–V, and telson; 2 distance between median ocelli; 3 sum of trochanter,
femur, patella, and chela; 4 measured from base of condyle to tip of xed nger; 5 sum of tergites I–VII; 6 sum of metasomal
segments I–V and telson.
TABLE 8 continued
macinta River into Guatemala. The known
records occur in the Lacondón Forest, on the
northern edge of the Montes Azules Biosphere
Reserve (g. 4).
E: e localities at which C. hama-
dryas has been recorded range in altitude from
103 to 153 m, all situated in a lowland tropical
rainforest. Most specimens of this strictly arbo-
real species were located with UV light detection
at night and captured by holding an insect net
beneath the branch on which they were sitting
and tapping the branch with a stick; their escape
reaction is to drop immediately to the leaf litter
below, where they invariably disappear. e habi-
tat and habitus are consistent with the arboreal,
corticolous ecomorphotype (Prendini, 2001a).
R: Specimens from Frontera Corozal,
Chiapas, Mexico, were misidentied as C. schmidti
by Francke (2007). Teruel and Stockwell (2002)
and Francke (2007) noted dierences among the
pectinal counts of specimens from Chiapas, with
13–16 (♂) and 13–14 (♀), and Honduras, with
12–15 (♂) and 13–15 (♀). Although this slight
variation in pectinal tooth counts does not pro-
vide sucient evidence to distinguish between C.
hamadryas and C. schmidti, additional morpho-
logical dierences, outlined in their respective
diagnoses, together with genetic divergence
among samples from the two areas, conrmed the
distinction between them.
M E: MEXICO: Chiapas:
Município Ocosingo: La Galleta, 2 km SE of
Frontera Corozal, 16°48′12.7″N 90°52′11.1″W,
132–150 m, 28.iv.2004, R. Paredes and J.L.
Castelo, collected with UV light detection, 2
♂ (AMNH [LP 2948]), 1 ♀ (CNAN SC3987),
16°49′55″N 90°56′08″W, 146 m, 7.iv.2005, A. Val-
dez, O.F. Francke, and A. Ballesteros, collected at
night with UV lamp, 1 juv. ♂ (CNAN SC3986),
16°48′18.5″N 90°54′25″W, 114 m, 28.iv.2005, A.
Valdez, O.F. Francke, and A. Ballesteros, urban
area toward blue water bridge, collected with UV
light detection, 2 ♂, 1 ♀, 1 juv. ♂, 1 juv. (CNAN
SC3988).
50 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 452
FIGURE 23. Centruroides Marx, 1890, telson, dorsal aspect. A, D. C. rileyi Sissom, 1995, A. ♂ (CNAN
SC4002), D. ♀ (CNAN SC4003). B, E. C. hamadryas, sp. nov., B. holotype ♂ (CNAN T01408), E. paratype
♀ (CNAN T01415). C, F. C. homanni Armas, 1996, C. ♂, F. ♀ (CNAN SC3996). G, J. C. cuauhmapan, sp.
nov., G. holotype ♂ (CNAN T01396), J. paratype ♀ (CNAN T01399). H, K. C. berstoni, sp. nov., H. holotype
♂ (CASENT 9073325), K. paratype ♀ (CASENT 9073313). I, L. C. chanae, sp. nov., I. holotype ♂ (CNAN
T01403), L. paratype ♀ (CNAN T01405). M, P. C. catemacoensis, sp. nov., M. holotype ♂ (CNAN T01424),
P. paratype ♀ (CNAN T01423). N, Q. C. schmidti Sissom, 1995, N. ♂ (CASENT 9073316), Q. ♀ (CASENT
9073317). O, R. C. yucatanensis, sp. nov., O. holotype ♂ (CNAN T01416), R. paratype ♀ (CNAN T01418).
Scale bars = 1 mm.
2021 GOODMAN ET AL.: REVISION OF “THORELLII” CLADE OF CENTRUROIDES 51
FIGURE 24. Centruroides Marx, 1890, telson, ventral aspect. A, D. C. rileyi Sissom, 1995, A. ♂ (CNAN
SC4002), D. ♀ (CNAN SC4003). B, E. C. hamadryas, sp. nov., B. holotype ♂ (CNAN T01408), E. paratype
♀ (CNAN T01415). C, F. C. homanni Armas, 1996, C. ♂, F. ♀ (CNAN SC3996). G, J. C. cuauhmapan, sp.
nov., G. holotype ♂ (CNAN T01396), J. paratype ♀ (CNAN T01399). H, K. C. berstoni, sp. nov., H. holotype
♂ (CASENT 9073325), K. paratype ♀ (CASENT 9073313). I, L. C. chanae, sp. nov., I. holotype ♂ (CNAN
T01403), L. paratype ♀ (CNAN T01405). M, P. C. catemacoensis, sp. nov., M. holotype ♂ (CNAN T01424),
P. paratype ♀ (CNAN T01423). N, Q. C. schmidti Sissom, 1995, N. ♂ (CASENT 9073316), Q. ♀ (CASENT
9073317). O, R. C. yucatanensis, sp. nov., O. holotype ♂ (CNAN T01416), R. paratype ♀ (CNAN T01418).
Scale bars = 1 mm.
52 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 452
FIGURE 25. Centruroides Marx, 1890, telson, lateral aspect. A, D. C. rileyi Sissom, 1995, A. ♂ (CNAN
SC4002), D. ♀ (CNAN SC4003). B, E. C. hamadryas, sp. nov., B. holotype ♂ (CNAN T01408), E. paratype
♀ (CNAN T01415). C, F. C. homanni Armas, 1996, C. ♂, F. ♀ (CNAN SC3996). G, J. C. cuauhmapan, sp.
nov., G. holotype ♂ (CNAN T01396), J. paratype ♀ (CNAN T01399). H, K. C. berstoni, sp. nov., H. holotype
♂ (CASENT 9073325), K. paratype ♀ (CASENT 9073313). I, L. C. chanae, sp. nov., I. holotype ♂ (CNAN
T01403), L. paratype ♀ (CNAN T01405). M, P. C. catemacoensis, sp. nov., M. holotype ♂ (CNAN T01424),
P. paratype ♀ (CNAN T01423). N, Q. C. schmidti Sissom, 1995, N. ♂ (CASENT 9073316), Q. ♀ (CASENT
9073317). O, R. C. yucatanensis, sp. nov., O. holotype ♂ (CNAN T01416), R. paratype ♀ (CNAN T01418).
Scale bars = 1 mm.
2021 GOODMAN ET AL.: REVISION OF “THORELLII” CLADE OF CENTRUROIDES 53
Centruroides homanni Armas, 1996
Figures 2, 4, 7A, B, 10A, B, 15A, A, 17C, F, 18C,
F, 19C, F, 20C, F, 21C, F, 22C, F, 23C, F, 24C, F,
25C, F, 38, 39 tables 1, 8, 10
Centruroides homanni Armas, 1996: 29–32, gs.
5–9; 1999: 47, 51; Beutelspacher-Baigts,
2000: 123, 126, 139, 144, 155, map 106 (in
part: records from Arriaga, Chiapas);
Kovařík, 1998: 107; Fet and Lowe, 2000:
109; González-Santillán, 2001: 573; Armas
et al., 2002: 94, 95; 2003: 94 (misidentica-
tion); 2004: 170, table 1 (misidentication);
Martín-Frías et al., 2005: 1–6, gs. 1–13
(misidentication); Teruel et al., 2006: 223;
Santibáñez-López and Ponce-Saavedra,
2009: 321, 323, 326, 328–231, gs. 12–15,
8–11, 16 (misidentication); Santibáñez-
López and Contreras-Felix, 2013: 131, 138,
g. 7 (misidentication); Teruel et al.,
2015a: 3, 6, 7, gs. 35–37 (misidentica-
tion); Kovařík et al., 2016b: 11 (misidenti-
cation); Esposito et al., 2017: 13; Esposito
and Prendini, 2019: 4, 7, g. 2.
T M: MEXICO: Chiapas: Muni-
cípio Arriaga: Holotype ♀, (CNAN 71), La Glo-
ria 16°08′39.4″N 94°06′04.7″W, 11.xii.1974. J.L.
Garcia, R. Ruiz, and J. Luis M.G., household
collection.
D: Centruroides homanni is most
closely related to C. chanae, from which it diers
as follows. A dark line along the lateral margins of
the carapace and mesosomal tergites I–III, and
pale stripe medially on the carapace and tergites,
absent in C. homanni (g. 7A, B) are present in
C. chanae (g. 7C, D). e carapace, pedipalps,
tergites, and metasoma are more infuscate, creat-
ing a more mottled appearance, in C. homanni
(gs. 38A, B, 39A, B) than C. chanae (gs. 40A, B,
41A, B). Less reticulate infuscation is present on
the chelicerae of C. homanni than C. chanae. e
interocular triangle is more darkly infuscate in C.
homanni than C. chanae. e marbled infusca-
tion of the mesosomal sternites is pronounced in
C. homanni, but faint or absent in C. chanae. e
carapace is longer, its length greater than its width,
in C. homanni, but shorter, its length and width
similar, in C. chanae (tables 8, 9). e carapace
surfaces are more coarsely granular, the carinae
more pronounced, and the sulci narrower and
deeper in C. homanni (g. 7A, B) than C. chanae
(g. 7C, D). e pedipalp chela manus of the male
is more incrassate in C. homanni than C. chanae
(g. 15A, B). e ventral surfaces of the telotarsi
of leg I are more nely and sparsely setose in C.
homanni than C. chanae. e pectinal tooth
count of the male is lower in C. homanni, usually
15, than C. chanae, usually 17 (table 8). e ven-
trolateral carinae of mesosomal sternite VII are
granular, and the ventrosubmedian carinae weakly
granular and restricted to the posterior half of the
segment in C. homanni, whereas the ventrolat-
eral carinae are distinct, granular, and the ventro-
submedian carinae weakly developed, granular in
C. chanae. Although the metasomal segments of
the male are longer and narrower in C. homanni
than C. chanae, the metasoma is less than 3× the
length of the mesosoma in C. homanni but
greater than 3× (up to 3.3×) its length in C. chanae
(table 10). e ventrolateral and ventrosubmedian
carinae of the metasomal segments are more pro-
nounced in C. homanni, being slightly serrate on
segments I–IV, compared with nely granular to
subserrate on I–III and obsolete, smooth on IV in
C. chanae. e ventrosubmedian carinae of seg-
ments I and II are very pronounced in C. ho-
manni (gs. 18C, F, 19C, F) but absent or obsolete
in C. chanae (gs. 18I, L, 19I, L). e telson of the
male is elongate, the vesicle bilobed posteriorly in
C. homanni (g. 25C, F), whereas the telson is
shorter, the vesicle rounded posteriorly in C. cha-
nae (g. 25I, L).
D: Centruroides hoffmanni is
endemic to the state of Chiapas in southeastern
Mexico. e known records are restricted to the
Central Depression, bounded by the Central
Highlands, to the north, and the Sierra Madre de
Chiapas, to the south (g. 4), an area which
exhibits high levels of endemism (Reyes-García
and Sousa, 1995).
54 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 452
FIGURE 26. Centruroides rileyi Sissom, 1995, ♂ (CNAN SC4002), habitus. A. Dorsal aspect. B. Ventral aspect.
Scale bars = 5 mm.
E: e localities at which C. ho-
manni has been recorded range in altitude from
529 to 1513 m and are situated in subtropical dry
forest. e habitat and habitus are consistent
with the arboreal, corticolous ecomorphotype
(Prendini, 2001a).
R: Confusion has surrounded this
species since its original description. Armas
(1996) described the holotype as an adult
female, but later (Armas, 1999; Armas et al.
2003) stated it was immature. Beutelspacher-
Baigts (2000) confused C. hoffmanni with Cen-
truroides nigrovariatus Pocock, 1898, and C.
tuxtla, erroneously listing the species from the
Mexican state of Oaxaca, an error repeated by
Armas et al. (2003, 2004). Martín-Frías et al.
(2005) redescribed C. hoffmanni from material
originating in Oaxaca that is evidently hetero-
specific with the holotype based on characters
of the female: the pectinal plate of the material
described is not distinctly lobed or posteriorly
rounded, the pectinal tooth count is higher,
the metasoma and telson markedly are granu-
lar, and the subaculear tubercule is not elon-
gate and angular. Santibáñez-López and
Ponce-Saavedra (2009) again misidentified C.
hoffmanni, presenting photographs of female
specimens in which the pectinal plate is not
distinctly lobed or posteriorly rounded, along
with measurements and pectinal tooth counts
inconsistent with the holotype, once more
erroneously listing the species from Oaxaca.
2021 GOODMAN ET AL.: REVISION OF “THORELLII” CLADE OF CENTRUROIDES 55
FIGURE 27. Centruroides rileyi Sissom, 1995, ♀ (CNAN SC4003), habitus. A. Dorsal aspect. B. Ventral aspect.
Scale bars = 5 mm.
The error was repeated by Santibáñez-López
and Contreras-Félix (2013), and yet again by
Teruel et al. (2015a), who erroneously associ-
ated C. hoffmanni with the “nigrovariatus
group” of Centruroides. Kovařík et al. (2016b)
followed previous authors in misidentifying
material from Guerrero and Oaxaca as C.
hoffmanni.
M E: MEXICO: Chi-
apas: Município Angel Albino Corzo: 8 km
from Siltepec, 18°48′33″N 92°40′30.6″W,
663 m, 17.viii.2007, C. Mayorga, G. Ortega,
and L. Cervantes, 1 juv. ♀ (CNAN SC3990).
Município Comitan: Parque Nacional Lagu-
nas de Montebelo, 16°17′17″N 91°56′16″W,
1473 m, 3.ix.2005, O.F. Francke, M. Córdova,
A. Jaimes, A. Valdez, and H. Montaño, 1 juv.
♂ (CNAN SC3992). Município La Concordia:
Villa Corzo La Tigrilla, San Julián, Revolu-
ción Mexicana, 16°00′00″N 92°50′47″W, 544
m, 17.iii.2007, C. Mayorga, G. Ortega, and
L. Cervantes. 1 ♂, 1 ♀ (CNAN SC3998).
Município Tuxtla Gutiérrez: Las Delicias,
16°45′31.1″N 93°06′26.4″W, 529 m, 2.iii.2005,
O.F. Francke, M. Córdova, A. Jaimes, A. Val-
dez, and H. Montaño, 1 ♀ (AMNH [LP 5224]);
Gutiérrez, San Julián, Revolución Mexicana,
16°11′41″N 93°01′16″W, 544 m, 16.iii.2007,
G. Ortega and A. Cervantes, 2 ♂, 3 ♀ (CNAN
SC3997). Município Tzimol: Carretera [Hwy]
Comitán–Tzimol Santa Rosa, 16°11′03.4″N
92°16′59.3″W, 632–730 m, 2.ix.2005, O.F.
Francke, M. Córdova, A. Jaimes, A. Valdez, and
H. Montaño, 1 ♂, 1 ♀ (AMNH [LP 5249]), 1
♀, first instar juvs (CNAN SC3994), 1 ♀, first
instar juvs (CNAN SC3995), 3 ♂, 9 ♀, 2 juv.
♂, 14 juv. ♀ (CNAN SC3996). Município Vil-
laflores: Reserva de La Biosfera La Sepultura,
1 km SE of Ejido California, 16°15′14.2″N
93°35′46.4″W, 1009–1132 m, 30.viii.2005, O.F.
Francke, M. Córdova, A. Jaimes, A. Valdez,
and H. Montaño, 1 ♂ (AMNH [LP 5350]), 1
♂, 2 ♀, 4 juv. ♂ (CNAN SC3993), 1 ♀, first
instar juvs (CNAN SC3991).
56 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 452
FIGURE 28. Centruroides cuauhmapan, sp. nov., ♂ (CNAN T01396), habitus. A. Dorsal aspect. B. Ventral
aspect. Scale bars = 5 mm.
Centruroides rileyi Sissom, 1995
Figures 2, 3, 5C, D, 8C, D, 11B, 12B, 17G, J,
18G, J, 19G, J, 20G, J, 21G, J, 22G, J, 23G, J,
24G, J, 25G, G, 26, 27, tables 1, 2, 10
Centruroides rileyi Sissom, 1995: 96–99, gs.
19–27; Armas et al., 2002: 1; 2003: 95;
Cancino and Blanco, 2002: 71; Sissom and
Hendrixson, 2005: 126, 127, 134, 475;
Esposito et al., 2017: 14, 30, g: 14; 2018:
97, 116; Esposito and Prendini, 2019: 4, g.
2; Ponce-Saavedra and Francke, 2019: 3;
Crews and Esposito, 2020: 14, g. 11; Good-
man and Esposito, 2020: 1–9, g. 1C
(misidentication).
T M: MEXICO: Tamaulipas:
Município Gómez Farías: Holotype ♂, paratype
♀ (USNM), Bocatoma, 7 km SSE of Gómez
Farías, 22°56′30.2″N 99°06′19.3″W, 25–30.
iii.1978, E.G. Riley; paratype ♀ (FSCA), Gómez
Farías, 16.iii.1977, R. Schmidt. San Luis Potosí:
Município Tamazunchale: Paratype ♀ (NAU), 5
km N of Tamazunchale o Hwy 85, 21°18′13.6″N
98°47′58.7″W, 1.viii.1987, J.A. Nilsson.
2021 GOODMAN ET AL.: REVISION OF “THORELLII” CLADE OF CENTRUROIDES 57
D: Centruroides rileyi is most closely
related to C. cuauhmapan, from which it diers as
follows. e posterosubmedian carinae of the
carapace are weakly developed in C. rileyi (g. 5A,
B), but absent in C. cuauhmapan (g. 5C, D). e
retrodorsal carina of the pedipalp chela manus is
smooth, the dorsomedian carina weakly granular,
and the prodorsal carina weakly granular and
restricted to the distal half of the segment, in the
male of C. rileyi (gs. 11, 12A) whereas the retro-
dorsal carina is nely granular, the dorsomedian
carina distinct, granular, and the prodorsal carina
distinct, granular and complete in the male of C.
cuauhmapan (gs. 11, 12B). e ventrolateral and
ventrosubmedian carinae of mesosomal sternite
VII are obsolete to absent and the intercarinal sur-
faces smooth in C. rileyi, whereas the ventrolateral
and ventrosubmedian carinae are distinct, granu-
lar and the intercarinal surfaces nely granular in
C. cuauhmapan. e metasoma and telson are
shorter in the male and slenderer, proportionally
shorter and narrower, in the female of C. rileyi
(gs. 17A, D, 18A, D, 19A, D, 20A, D, 21A, D,
22A, D, 23A, D, 24A, D, 25A, D, table 2) than C.
cuauhmapan (gs. 17G, J, 18G, J, 19G, J, 20G, J,
21G, J, 22G, J, 23G, J, 24G, J, 25G, J, table 3). e
ventral carinae are granular on metasomal seg-
ment I in the female, vestigial on segments I–III
and smooth on IV and V in the male of C. rileyi
(gs. 18A, D, 19A, D, 20A, D, 21A, D, 22A, D) but
distinct, granular on segments I–V in the male
and female of C. cuauhmapan (gs. 18G, J, 19G, J,
20G, J, 21G, J, 22G, J). e surfaces of the telson
vesicle of the female are smooth in C. rileyi (gs.
24A, D, 25A, D) but granular in C. cuauhmapan
(gs. 24G, J, 25G, J).
FIGURE 29. Centruroides cuauhmapan, sp. nov., ♀ (CNAN T01399), habitus. A. Dorsal aspect. B. Ventral
aspect. Scale bars = 5 mm.
58 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 452
TABLE 9
Meristic data for type material of Centruroides chanae, sp. nov., and Centruroides yucatanensis, sp. nov.
Material deposited in the Colección Nacional de Arácnidos (CNAN), Universidad Nacional Autónoma de
México, Mexico City. Measurements follow Stahnke (1970), Lamoral (1979), and Prendini (2001b).
Centruroides chanae Centruroides yucatanensis
Holotype Paratypes Paratypes Holotype Paratype Paratypes
♂ ♂ ♀ ♂ ♀ juv. ♀
CNAN CNAN CNAN CNAN CNAN CNAN
T01403 T01404 T01405 T01406 T01407 T01416 T01417 T01418 T01419
Total length142.4 42.6 38 34.4 33.2 48.7 29.9 27.8 24.8
Carapace length 3.4 3.5 3.3 2.9 3.3 3.7 3.4 2.7 2.9
ant. width 1.8 1.7 1.6 1.4 1.8 1.7 1.6 1.3 1.5
post. width 3.8 3.8 3.4 3.0 3.8 3.7 3.5 2.8 3.1
Median ocelli diameter 0.3 0.3 0.3 0.2 0.3 0.3 0.3 0.2 0.3
Interocular length20.3 0.4 0.3 0.3 0.4 0.4 0.3 0.3 0.3
Pedipalp length315.1 16.0 16.7 12.7 14.3 16.5 14.7 10.8 11.5
Trochanter length 1.5 1.5 3.5 1.1 1.3 1.7 1.4 1.0 1.1
Femur length 3.7 3.8 3.6 3.0 3.3 3.8 3.3 2.4 2.6
width 0.7 0.6 0.6 0.5 0.8 0.7 0.8 0.5 0.9
height 1.0 1.0 0.9 0.8 1.0 1.0 1.0 0.9 0.6
Patella length 3.9 4.2 3.9 3.6 3.9 4.5 3.9 2.9 3.1
width 0.9 1.0 0.9 0.8 0.8 1.0 1.0 0.8 0.9
height 1.5 1.5 1.3 1.2 1.5 1.5 1.6 1.3 1.4
Chela length46.0 6.5 5.7 5.0 5.8 6.5 6.1 4.6 4.8
Manus length 2.9 2.7 3.5 3.0 3.5 3.6 3.7 1.6 1.8
width 1.1 1.3 1.0 1.0 1.1 1.4 1.2 0.8 0.9
height 1.0 1.0 1.0 0.9 0.8 1.3 1.2 0.9 0.9
Mov. nger length 4.4 4.3 4.0 3.5 4.2 4.1 4.3 3.4 3.5
Leg I length 6.1 6.6 6.2 5.5 6.0 6.6 6.1 4.6 5.1
Pectines length 5.2 6.2 5.7 5.0 4.7 5.5 4.5 3.5 3.7
tooth count 15/15 17/17 17/15 17/17 14/14 14/14 14/14 12/12 13/13
Mesosoma length59.5 9.4 8.0 8.2 8.8 13.0 8.7 8.8 7.6
Sternite VII length 3.5 2.7 2.6 2.5 3.0 2.8 2.2 2.3 2.0
width 3.3 3.5 3.0 2.8 3.8 3.1 3.5 3.3 3.0
Metasoma length629.5 29.7 26.7 23.3 21.1 32.0 17.8 16.3 17.1
Metasoma I length 3.7 3.7 3.5 3.1 2.6 4.1 2.5 2.1 2.0
width 1.6 1.5 1.3 1.3 1.6 1.5 1.7 1.4 1.5
height 1.5 1.5 1.3 1.3 1.5 1.5 1.5 1.2 1.3
2021 GOODMAN ET AL.: REVISION OF “THORELLII” CLADE OF CENTRUROIDES 59
Centruroides chanae Centruroides yucatanensis
Holotype Paratypes Paratypes Holotype Paratype Paratypes
♂ ♂ ♀ ♂ ♀ juv. ♀
CNAN CNAN CNAN CNAN CNAN CNAN
T01403 T01404 T01405 T01406 T01407 T01416 T01417 T01418 T01419
Metasoma II length 4.7 4.7 4.2 3.7 3.2 5.1 3.2 2.4 2.5
width 1.4 1.3 1.2 1.2 1.4 1.2 1.4 1.1 1.3
height 1.4 1.4 1.3 1.3 1.5 1.1 1.5 1.3 1.2
Metasoma III length 5.5 5.2 4.8 4.1 3.5 5.8 3.6 2.7 2.7
width 1.4 1.3 1.2 1.2 1.3 1.4 1.4 1.1 1.2
height 1.5 1.4 1.3 1.1 1.5 1.3 1.6 1.2 1.3
Metasoma IV length 5.8 5.7 5.3 4.5 4.0 6.1 4.0 3.1 3.2
width 1.3 1.2 1.2 1.2 1.3 1.3 1.3 1.0 1.1
height 1.4 1.4 1.2 1.1 1.5 1.3 1.5 1.1 1.2
Metasoma V length 6.0 6.4 5.7 4.8 4.5 6.9 4.5 3.5 3.9
width 1.4 1.3 1.2 1.1 1.3 1.4 1.3 1.0 1.0
height 1.5 1.3 1.3 1.3 1.5 1.3 1.5 1.1 1.2
Tels on length 3.8 4.0 3.2 3.1 3.3 4.0 - 2.6 2.8
Vesicle length 2.7 2.7 2.2 2.0 1.6 2.9 - 1.5 1.6
width 1.3 1.1 1.0 0.9 0.9 1.2 - 0.7 0.7
height 1.2 1.2 1.0 1.0 1.1 1.2 - 0.8 0.8
Aculeus length 1.3 1.6 1.3 1.2 1.6 1.3 - 1.3 1.3
1 Sum of carapace, tergites I–VII, metasomal segments I–V, and telson; 2 distance between median ocelli; 3 sum of trochanter,
femur, patella, and chela; 4 measured from base of condyle to tip of xed nger; 5 sum of tergites I–VII; 6 sum of metasomal
segments I–V and telson.
TABLE 9 continued
V: Adult males and females dier as
follows. e dorsomedian carinae of the pedipalp
patella are absent, the mesosoma proportionally
longer and slenderer, and the metasoma longer, in
males (gs. 26A, B, 27A, B, 28A, B, 29A, B, table
2). e rst pair of legs are longer in males and
the metasomal carinae more pronounced and
nely serrate in females (gs. 17A, D, 18A, D,
19A, D, 20A, D, 21A, D, 22A, D, table 2).
D: Centruroides rileyi is endemic
to northern Mexico, east of the Sierra Madre
Oriental. e species is fairly widespread, with
records from the states of Puebla, Tamaulipas,
San Luis Potosí, and Veracruz (g. 3).
E: e localities at which C. rileyi has
been recorded range in altitude from 100 to 2554
m. e habitat at these localities varies from sub-
tropical and semi-deciduous forests near the El
Cielo Biosphere Reserve, Tamaulipas, to tropical
and subtropical moist broadleaf forest at La Sierra
Gorda, San Luis Potosí, and tropical humid moist
forest in Veracruz (Mendoza-Villa et al., 2018). e
habitat and habitus are consistent with the arboreal,
corticolous ecomorphotype (Prendini, 2001a).
R: Centruroides rileyi has never been
confused with other species of the “thorellii” clade
nor with C. thorellii, perhaps due to its occurrence
in northern Mexico. Furthermore, the small size
and distinctive mottling pattern contrast with other
buthids, such as Centruroides gracilis (Latreille,
1804) and Centruroides vittatus (Say, 1821), that
occur in sympatry (Shelley and Sissom, 1995).
60 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 452
FIGURE 30. Centruroides catemacoensis, sp. nov., ♂ (CNAN T01424), habitus. A. Dorsal aspect. B. Ventral
aspect. Scale bars = 5 mm.
M E: MEXICO: Puebla:
Município Cuetzalan el Progreso: Cuetzalan,
Santiago Yancuitlalpan, 18°54′42.2″N
98°35′15.3″W, 2554 m, 19.v.1995, G. Oclogaig
Barrzia, juvs (CNAN SC3999). San Luis Potosí:
Município Axtlan de Terrazas: Axtlan de Terra-
zas, 21°25′34.9″N 98°52′42″W, 100 m, 28.iv.2006,
O.F. Francke, A. Valdez, G. Villegas and R. Pare-
des, 1 ♂ (AMNH [LP 6445]), 1 ♀, 2 juv. ♀
(CNAN SC4003). Veracruz: Município Papantla:
Papantla, 20°27′24.1″N 97°18′56.1″W, 2197 m,
iii.2000, J.L. Castelo, 1 ♂ (CNAN SC4000).
Município Tamiahua: Moralillo, Cerro Azul,
21°11′03.8″N 97°44′49.6″W, 153 m, 27.ii.2007, E.
Barrera and L. Cervantes, 2 ♀ (CNAN SC3985),
1 juv. ♀ (CNAN SC4002).
2021 GOODMAN ET AL.: REVISION OF “THORELLII” CLADE OF CENTRUROIDES 61
FIGURE 31. Centruroides catemacoensis, sp. nov., ♀ (CNAN T01423), habitus. A. Dorsal aspect. B. Ventral
aspect. Scale bars = 5 mm.
Centruroides schmidti Sissom, 1995
Figures 1A, C, 2, 4, 6E, F, 9E, F, 13C, 14C, 17N,
Q, 18N, Q, 19N, Q, 20N, Q, 21N, Q, 22N, Q,
23N, Q, 24N, Q, 25Q, B, 36, 37, tables 1, 7, 10
Centruroides schmidti Sissom, 1995: 94–96, 98,
gs. 10–18; Armas, 1996: 22–24, table I
(misidentication, part); 1999: 30; Vázquez,
1999: 53, 60–62, g. 7 (misidentication,
part); Armas and Maes, 2000: 27; 2001: 16;
Fet and Lowe, 2000: 118; Armas et al., 2002:
169–171 (misidentication); Teruel and
Stockwell, 2002: 111–127, gs. 6, 20, tables
II, III; Armas et al., 2003: 95–96 (misidenti-
cation); Armas and Martín-Frías, 2003:
205, 209; 2008: 7–10, 12, 17, 19, 20, gs.
2–4, table XIV (misidentication); Armas
and Trujillo, 2010: 235, 238, 240; Borges et
al., 2012: 131, table I; Teruel et al., 2015a: 7;
Delfín-González et al., 2017: 284 (misidenti-
cation), table I; Esposito et al., 2018: 97,
table 5; Esposito and Prendini, 2019: 4, g.
2; Crews and Esposito, 2020: 14, g. 11.
T M: HONDURAS: Departa-
mento Cortés: Município Choloma: Holotype ♂
(FMNH), Coloma [Choloma], Lake Ticamaya
15°32′41.5″N 87°53′06.1″W, 26.iv.1923, K.
Schmidt and L. Walters (Capt. Field Mus.
Exped.), found on bones of crocodile skull.
GUATEMALA: Departamento Izabal: Município
Morales: Paratype ♀ (FMNH), Escobas, Izabal
15°24′12″N 89°08′24.5″W 27.xi.1933, K.P. and
P.J. Schmidt, Leon Mandel Guatemala Exped.
D: Centruroides schmidti diers from
the closely related species, C. berstoni, C.
catemacoensis, C. cuauhmapan, C. hamadryas,
62 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 452
FIGURE 32. Centruroides hamadryas, sp. nov., ♂ (CNAN T01408), habitus. A. Dorsal aspect. B. Ventral
aspect. Scale bars = 5 mm.
and C. rileyi, as follows. e pattern of infuscation
of C. schmidti is unlike that of the other species
and includes a pronounced pale stripe medially
on the carapace and mesosomal tergites, anked
on the tergites by a pair of orange stripes, which
are more infuscate in females (gs. 36A, B, 37A,
B). e cheliceral manus is entirely dark with
reticulate infuscation in C. schmidti, whereas the
infuscation is lighter and restricted to the distal
half of the chelicerae in the other species. e
2021 GOODMAN ET AL.: REVISION OF “THORELLII” CLADE OF CENTRUROIDES 63
telotarsi of the rst pair of legs possess short,
dense setae in C. schmidti, unlike in C. cuauhma-
pan and C. rileyi. e posterosubmedian and lat-
eral ocular carinae on the carapace are present in
C. schmidti (g. 6E, F) but reduced or absent in C.
catemacoensis (g. 5E, F) and absent in C. berstoni
(g. 6C, D), C. cuauhmapan (g. 5C, D), and C.
hamadryas (g. 6A, B). e posterior margin of
sternite III is pale and setose in C. schmidti, unlike
in the other species. e dorsomedian carinae are
restricted to the posterior two-thirds of tergites
I–VI, and absent on VII in C. schmidti, whereas
the dorsomedian carinae are vestigial on tergites
I–VII in C. catemacoensis and weakly granular on
I–VII in C. berstoni. Ventrosubmedian and ven-
trolateral carinae are present on sternite VII in C.
schmidti, unlike in C. catemacoensis and C. rileyi.
Metasomal segment V is more than 2× the length
of the carapace in C. schmidti, but less than 2× its
length in the other species (table 7). e telson
surfaces of the female are very granular and the
ventromedian carina well developed in C. schmidti
(gs. 23–25D), whereas the telson surfaces of
females are smooth in C. hamadryas (gs. 23–25E)
and C. rileyi (gs. 23–25Q). e subaculear tuber-
cle is strongly angled toward the aculeus in C.
schmidti (g. 25N, Q)
Centruroides schmidti diers from C. homanni
as follows. e mottled infuscation of the cara-
pace, pedipalps, tergites, and metasoma is less
pronounced in C. schmidti than C. homanni, but
the carapace is more infuscate, with a darker bor-
der around the margins, in C. schmidti. e cara-
pacial sulci are broad and shallow in C. schmidti
FIGURE 33. Centruroides hamadryas, sp. nov., ♀ (CNAN T01415), habitus. A. Dorsal aspect. B. Ventral
aspect. Scale bars = 5 mm.
64 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 452
FIGURE 34. Centruroides berstoni, sp. nov., ♂ (CASENT 9073325), habitus. A. Dorsal aspect. B. Ventral aspect. Scale bars = 5 mm.
2021 GOODMAN ET AL.: REVISION OF “THORELLII” CLADE OF CENTRUROIDES 65
FIGURE 35. Centruroides berstoni, sp. nov., ♀ (CASENT 9073313), habitus. A. Dorsal aspect. B. Ventral
aspect. Scale bars = 5 mm.
but narrow and deep in C. homanni. e dorso-
median carina is restricted to the posterior two-
thirds of tergites I–VI, and absent on VII in C.
schmidti, whereas the dorsomedian carina is com-
plete on tergites I–VII in C. homanni.
V: Carapace surface granulation varies
from sparse granules to uniform, moderate granu-
lation. Specimens from Honduras exhibit variation
in granulation of the lateral ocular carinae (g. 6E,
F). e dorsomedian carina of the pedipalp chela
manus of the male is well developed in material
from Guatemala but weakly developed to absent in
material from Honduras.
Adult males and females dier as follows. e
pedipalp chela of the male is incrassate unlike the
female. e prodorsal carina on the chela manus
comprises a row of spiniform granules in the male
but is nely granular in the female (g. 13C, D). e
mesosoma is proportionally longer and slenderer,
the metasoma up to 3× longer, with segment V
markedly longer, and the telson more elongate, with
the vesicle more rounded, in males (gs. 17N, Q,
18N, Q, 19N, Q, 20N, Q, 21N, Q, 22N, Q, 23N, Q,
24N, Q, 25N, Q, table 2).
D: Centruroides schmidti is the
most widespread species of the clade. It appears
66 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 452
FIGURE 36. Centruroides schmidti Sissom 1995, ♂ (CASENT 9073278), habitus. A. Dorsal aspect. B. Ventral aspect. Scale bars = 5 mm.
2021 GOODMAN ET AL.: REVISION OF “THORELLII” CLADE OF CENTRUROIDES 67
to be endemic to Guatemala, where it has been
recorded from the Izabal and Zacapa depart-
ments, and Honduras, where it has been recorded
from the Atlántida, Cortés, Francisco Morazán,
and Islas de la Bahía departments. e known
localities extend along the Caribbean coasts of
both countries, including the Bay Islands of
Honduras, and inland to the Sierra de Las Minas
of Guatemala (g. 4).
E: e localities at which C. schmidti has
been recorded range in altitude from 12 to 773 m.
is species occurs in a broader range of habitats
than other species of the “thorellii” clade. e habi-
tat at localities near Zacapa, Guatemala is semiarid
savannah, dominated by scrub forest and cacti. In
this area, specimens were found on the bark of
large oaks at night. e habitat at San Antonio de
Oriente, Honduras, is open deciduous broadleaf
forest/savannah, interspersed with grassland. e
habitat near Las Minas, Guatemala, is moist mon-
tane subtropical pine-oak forest; specimens were
collected on Pinus oocarpa Schiede ex Schltdl. and
various oak species (g. 2A, C). e habitat on the
northern coast and Bay Islands of Honduras is low-
land tropical rainforest. e habitat and habitus are
consistent with the arboreal, corticolous ecomor-
photype (Prendini, 2001a).
R: Previous records of C. schmidti from
Costa Rica and the Mexican states of Chiapas,
Quintana Roo, and Veracruz are misidentica-
tions of other species, including some of those
described herein. Armas (1996) included speci-
mens from Quintana Roo, probably conspecic
with C. yucatanensis, in a redescription of C.
schmidti. Armas et al. (2002) described an indi-
vidual from northern Costa Rica as C. schmidti,
FIGURE 37. Centruroides schmidti Sissom 1995, ♀ (CASENT 9073317), habitus. A. Dorsal aspect. B. Ventral
aspect. Scale bars = 5 mm
68 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 452
FIGURE 38. Centruroides homanni Armas 1996, ♂ (CNAN SC3996), habitus. A. Dorsal aspect. B. Ventral
aspect. Scale bars = 5 mm.
2021 GOODMAN ET AL.: REVISION OF “THORELLII” CLADE OF CENTRUROIDES 69
FIGURE 39. Centruroides homanni Armas 1996, ♀ (CNAN SC3996), habitus. A. Dorsal aspect. B. Ventral
aspect. Scale bars = 5 mm.
without dierentiating it from C. thorellii, a spe-
cies erroneously recorded from the country by
Francke and Stockwell (1987). Armas et al. (2003)
included Mexico (Chiapas, Quintana Roo, and
Yucatán) in the distribution of C. schmidti.
Vázquez (1999) again included C. schmidti in the
fauna of Quintana Roo and erroneously suggested
the species is endemic to the Yucátan Peninsula.
Photographs and measurements of individuals
identied as C. schmidti from the Sian Ka’an Bio-
sphere Reserve, Quintana Roo, by Armas and
Martín-Frías (2008), are consistent with the color-
ation and morphology of C. yucatanensis. Addi-
tionally, Armas and Martín-Frías (2008) listed C.
schmidti from Veracruz. Delfín-González et al.
(2017) listed C. schmidti from Chiapas and Vera-
cruz. Finally, Armas and Martín-Frías (2003)
erroneously suggested that the distribution of C.
schmidti extends from southeastern Mexico to
Costa Rica, following Armas et al. (2002).
M E: GUATEMALA:
Departamento El Progreso: Município Rio
Hondo: San Francisco Zapotitlan: Finca El
Olvido, Las Minas, 15°02′04.8″N 89°52′26.7″W,
1214 m, 17.ix.2019, A.M. Goodman, UV hand
collection, found on oak and pine trees, 2–3
m high, 4 ♀, 4 juv. ♀ (CASENT 9073317),
18.ix.2019, A.M. Goodman, L.A. Esposito, and
70 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 452
L. Allen, 5 ♀, 1 juv. ♂, 1 juv. ♀ (CASENT
9073402). Departamento Zacapa: Município
Rio Hondo: Bosque Pino, Guadalupe, Manta
de Golpeo, 14°58′04.7″N 89°24′47″W, 751 m,
20.ix.2019, A.M. Goodman, M. Barrios, and M.
van Dam, UV hand collection, found on oak
and pine trees, 2–3 m high, 7 ♂, 1 ♀, 1 juv. ♂,
1 juv. ♀ (CASENT 9073278); Aldea Casas de
Pinto, near turno for Zacapa at Rio Hondo,
15°01′38.2″N 89°36′57.2″W, 77 m, 13.vii.2006,
J.H. Hu, semiarid region with scrub forest and
cacti, collected under rocks in shaded areas and at
night using UV, 1 ad. (AMNH [LP 5985]). HON-
DURAS: Departamento Atlántida: Município
La Ceiba: Parque Nacional Pico Bonito, Pico
Bonito, trails from Visitor Centre and park
entrance, 14°43′30.6″N 86°44′11.5″W, 184 m,
30.viii.2013, S. Longhorn, dense wet lowland
tropical forest near large river, sweeping and
beating, may have been on or under wood,
day search, 1 juv. ♂ (AMNH [LP 13416]).
Departmento Francisco Morazán: Município
San Antonio de Oriente: E.A.P. Zamorano,
Monte Redondo, Acuacultura, 13°39′59.6″N
86°59′21″W, 773 m, 23.ix.2008, C. Víquez,
UV at night, 1 ♀ (AMNH [LP 9172]). Depar-
tamento Islas de la Bahía: Município Roatán:
Cayos Cochinos, Cayos Menor, forest trails,
15°57′26.9″N 86°30′03.3″W, 101 m, 2.viii.2012,
S. Longhorn, scrub oak forest, 1 ♀ (AMNH [LP
13411]); Isla Utila, 16°06′22.1″N 86°54′08.1″W,
12 m, 21.vii.2012. S. Longhorn, scrub forest/wet
Savannah, 1 ♀ (AMNH LP [13417]).
Centruroides yucatanensis, sp. nov.
Figures 2, 4, 7E, F, 10E, F, 15C, 16C, 17O, R,
18O, R, 19O, R, 20O, R, 21O, R, 22O, R, 23O,
R, 24O, R, 25R, O, 42, 43, tables 1, 9, 10
Centruroides schmidti: Armas, 1996: 25–29, 98,
gs. 1–4 (misidentication), table I.
Centruroides sissomi: Vázquez, 1999: 53, 60–62,
g. 7 (misidentication); Armas et al., 2003:
95 (misidentication, part); Armas, 2006: 4,
7, g. 5 (misidentication, part); Teruel et
al., 2015a: 8 (misidentication); Delfín-
González et al., 2017: 283, 285, table 2
(misidentication); Esposito and Prendini,
2019: 4, g. 2 (misidentication); Ponce-
Saavedra and Francke, 2019: 3, table 1
(misidentication); Crews and Esposito,
2020: 14, g. 11 (misidentication).
T M: MEXICO: Quintana Roo:
Município Benito Juárez: Holotype ♂ (CNAN
T01416), paratype ♀ (CNAN T01417), 2 juv. ♀
paratypes (CNAN T01418, T01419), Puerto
Morelos, Jardín Botánico Alfredo Barrera,
20°50′42.1″N 86°54′12.9″W, 23 m, 4.vii.2007, G.
Montiel, R. Paredes, M. Ramírez, D. Chibras, and
G. Bonilla.
E: e species name refers to the
Yucatán Peninsula of southeastern Mexico,
where the species occurs.
D: Centruroides yucatanensis diers
from the closely related species, C. chanae and C.
homanni, as follows. e carapace, pedipalps,
tergites, and metasoma are less infuscate, creat-
ing a less mottled appearance, in C. yucatanensis
than C. chanae. Less reticulate infuscation is
present on the chelicerae of C. yucatanensis than
C. chanae. e interocular triangle is less darkly
infuscate in C. yucatanensis than C. homanni.
e carapace is shorter, its length and width
similar, in C. yucatanensis (g. 7E, F, table 10)
but longer, its length greater than its width, in C.
homanni (g. 7A, B, table 10). e carapace
surfaces are more nely granular, the carinae less
developed and the sulci broader and shallower in
C. yucatanensis than C. homanni. e pedipalp
chela manus of the male is less incrassate in C.
yucatanensis (g. 15C) than C. homanni (g.
15B), with fewer spiniform granules on its pro-
lateral surfaces than in C. chanae and C. ho-
manni (g. 15A, B). e ventral surfaces of the
telotarsi of leg I are more nely and sparsely
setose in C. yucatanensis than C. chanae. e
pectinal tooth count of the male is lower in C.
yucatanensis, usually 13 or 14 (g. 9E, table 10)
than C. chanae, usually 17 (g. 9A, table 10) and
C. homanni, usually 15 (g. 9D, table 10), and
2021 GOODMAN ET AL.: REVISION OF “THORELLII” CLADE OF CENTRUROIDES 71
the pectinal teeth are more ovoid. e ventrolat-
eral and ventrosubmedian carinae of mesosomal
sternite VII are vestigial, weakly granular in C.
yucatanensis, whereas the ventrolateral carinae
are distinct, granular, and the ventrosubmedian
carinae weakly developed, granular in C. chanae,
and the ventrolateral carinae granular, and the
ventrosubmedian carinae weakly granular and
restricted to the posterior half of the segment in
C. homanni. e ventrolateral and ventrosub-
median carinae of the metasomal segments are
more pronounced in C. yucatanensis, being
slightly serrate on segments I–IV, compared with
nely granular to subserrate on I–III and obso-
lete, smooth on IV in C. chanae. e ventrosub-
median carinae of segments I and II are absent
in C. yucatanensis (gs. 18O, R, 19O, R, 20O, R,
21O, R, 22O, R), absent or obsolete in C. chanae
(gs. 18I, L, 19I, L, 20I, L, 21I, L, 22I, L), and
very pronounced in C. homanni (gs. 18C, F,
19C, F, 20C, F, 21C, F, 22C, F).
D: e following description is
based on the holotype male, with dierences
among other material noted in the section on
variation.
Coloration: Base color yellow, with extensive
infuscation, creating mottled or marbled pattern.
Carapace with uniformly infuscate marbling,
more densely infuscate medially. Pedipalp chela
ngers and manus, dorsal and retrolateral inter-
carinal surfaces with moderately infuscate mar-
bling; prolateral and ventral intercarinal surfaces
mostly immaculate. Legs retrolateral surfaces
with infuscate marbling; prolateral surfaces pale,
immaculate. Tergites with unformly infuscate
mottling, pale stripe medially, blackish spots sub-
medially, and faint, narrow bands laterally. Ster-
nites with faintly infuscate marbling. Metasomal
segments uniformly, faintly marbled; segment V
and telson markedly infuscate, noticeably darker
than preceding segments.
Carapace: Shape trapezoidal; anterior width
four-hs of posterior width (table 10); antero-
median sulcus moderately deep, oval; postero-
median sulcus shallow anteriorly, deep
posteriorly; median ocular tubercle weakly gran-
ular; carinae moderately developed, comprising
small to medium-sized granules; lateral ocular
and posterosubmedian carinae weakly devel-
oped; intercarinal surfaces nely and evenly
granular (g. 10E).
Pedipalps: Orthobothriotaxic, Type A;
femur dorsal trichobothria with α configura-
tion; pedipalp chela fixed finger, trichoboth-
TABLE 10
Diagnostic ratios (mean/median/mode) for species of the arboreal Neotropical “thorelli” clade of
Centruroides Marx, 1890, bark scorpions
Leg I length:
carapace length
Metasoma V length:
mesosoma length
Metasoma V length:
carapace length
Centruroides berstoni, sp. nov. 1.9/1.9/1.9 2.5/2.4/2.0 1.6/1.6/1.9
Centruroides catemacoensis, sp. nov. 1.9/1.9/1.9 2.3/2.2/2.2 1.5/1.5/1.7
Centruroides chanae, sp. nov. 1.9/1.9/1.9 3.0/3.1/3.1 1.7/1.8/1.8
Centruroides cuauhmapan, sp. nov. 1.8/2.1/1.9 2.2/2.1/1.8 1.5/2.1/1.7
Centruroides hamadryas, sp. nov. 1.8/1.9/1.8 1.8/2.8/2.8 1.6/1.7/1.8
Centruroides homanni Armas, 1996 1.8/1.8/1.8 2.3/2.4/1.7 1.6/1.7/1.3
Centruroides rileyi Sissom, 1995 1.9/1.9/1.9 2.2/2.1/2.1 1.5/1.5/1.2
Centruroides schmidti Sissom, 1995 2.0/1.9/1.9 2.2/2.2/2.8 1.5/1.4/1.9
Centruroides yucatanensis, sp. nov. 1.8/1.8/1.8 2.1/2.1/2.0 1.5/1.4/1.3
72 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 452
FIGURE 40. Centruroides chanae, sp. nov., ♂ (CNAN T01407), habitus. A. Dorsal aspect. B. Ventral aspect. Scale bars = 5 mm.
2021 GOODMAN ET AL.: REVISION OF “THORELLII” CLADE OF CENTRUROIDES 73
FIGURE 41. Centruroides chanae, sp. nov., ♀ (CNAN T01405), habitus. A. Dorsal aspect. B. Ventral aspect.
Scale bars = 5 mm.
rium db situated slightly distal to et. Femoral
carinae strongly developed, serrate; dorsal
intercarinal surface moderately granular; pro-
lateral intercarinal surface with series of large
spiniform granules. Patella carinae strongly
developed, granular; prolateral intercarinal
surface with five or six large subspiniform
granules. Chela manus dorsomedian and
retrodorsal carinae complete, granular;
prodorsal carina absent. Fixed finger, median
denticle row comprising eight oblique sub-
rows, each flanked by pro- and retrolateral
supernumerary denticles. Movable finger,
median denticle row with short terminal row
comprising four denticles preceded by eight
oblique subrows, each flanked by pro- and ret-
rolateral supernumerary denticles.
Legs: Leg I length 1.78× greater than carapace
length (table 10). Telotarsi ventral surfaces
sparsely covered with short setae; ungues mark-
edly curved.
Pectines: Pectinal plate 1.61× wider than long;
posterior margin distinctly rounded; pectinal
tooth count 13/14 (♂) (g. 8D, table 10).
Mesosoma: Tergites width similar to carapace
posterior width; I and II slightly narrower (table
10). Pretergites surfaces smooth to nely granu-
lar. Posttergites surfaces weakly granular; I–VI
with dorsomedian carinae absent on I and II,
vestigial, granular on III–VI; VII surface weakly
74 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 452
FIGURE 42. Centruroides yucatanensis, sp. nov., ♂ (CNAN T01416), habitus. A. Dorsal aspect. B. Ventral
aspect. Scale bars = 10 mm.
2021 GOODMAN ET AL.: REVISION OF “THORELLII” CLADE OF CENTRUROIDES 75
FIGURE 43. Centruroides yucatanensis, sp. nov., ♀ (CNAN T01417), habitus. A. Dorsal aspect. B. Ventral
aspect. Scale bars = 10 mm.
granular, dorsomedian carina vestigial, granular,
dorsosubmedian and dorsolateral carinae
smooth. Sternites III–VI, surfaces smooth; VII
surface smooth, ventrolateral and ventrosubme-
dian carinae smooth.
Metasoma: Metasoma length 2.45× mesosoma
length (table 10). Segments longer than wide;
increasing in length posteriorly, segment V 2×
length of I; ventral carinae vestigial, weakly gran-
ular on segments I–IV, other carinae absent or
obsolete; lateral intercarinal surfaces sparsely
granular on segments I–III, other surfaces
smooth (gs. 17–22O).
Tels o n : Vesicle elongate, ovoid; ventral sur-
face shallowly convex; ventromedian carina
granular, terminating at subaculear tubercle;
subaculear tubercle narrow and angular in lat-
eral aspect, directed toward midpoint of acu-
leus. Aculeus angled ventrally at slightly less
than 90° (g. 25O, R).
Variation: Base coloration varies from light yel-
low to orange. Adult males and females dier as
follows. e pedipalp chela manus is incrassate,
with the prodorsal carina spinose, the mesosoma
proportionally longer and slenderer, the meta-
soma 2× longer, with segment V markedly longer
(1.86× carapace length), and the telson more elon-
gate, with the vesicle more rounded and bilobed
posteriorly, in males (gs. 23O, R, 24O, R, 25O, R,
table 10). e tegument is more densely infuscate,
the prodorsal carina of the pedipalp chela manus
nely granular, the pectinal plate produced into a
rounded lobe posteriorly, which is punctate and
slightly infuscate, metasomal segment V shorter
(1.3× carapace length) and the telson shorter and
narrower, with the vesicle surfaces less granular, in
females (gs. 10E, F, 15–16C, 17O, R, 18O, R,
19O, R, 20O, R, 21O, R, 22O, R, 23O, R, 24O, R,
25O, R, table 10). e pectinal tooth count is simi-
lar in both sexes (table 9).
D: Centruroides yucatanensis is
endemic to the Yucatán Peninsula. Although pres-
ently known from only three localities, two in the
state of Yucatán in the north of the peninsula, and
a third on the western coast, in the state of Quin-
tana Roo, the distribution of this species was prob-
ably more extensive before much of its habitat was
destroyed for agriculture and rangeland (g. 4).
76 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 452
E: e localities at which C. yucata-
nensis has been recorded range in altitude from
23–38 m. e habitat at these localities varies
from low or medium semideciduous broadleaf
forest to tall, moist evergreen broadleaf forest,
oen with a dense understory. Much of the origi-
nal habitat has been cleared for agriculture and
rangeland across the Yucatán Península, and this
species appears to be confined to remnant
patches of forest. e habitat and habitus are
consistent with the arboreal, corticolous ecomor-
photype (Prendini, 2001a).
R: Armas (1996) identied 11 indi-
viduals from the Jardín Botánico Alfredo Bar-
rera, Puerto Morelos, Quintana Roo, as C.
schmidti. Meristic data recorded by Armas (1996)
for an adult male and three females diered
greatly from comparative data for the holotype of
C. schmidti collected at Lake Tickamaya, Hondu-
ras (Sissom, 1995). Armas (1996) also described
C. sissomi, and assigned it to the thorellii group
due to its small size. Illustrations and photo-
graphs of the holotype female of C. sissomi in the
original description indicate that the pectinal
plate is not lobed, and subsequent photographs
of the holotype by Armas (2006) indicate a
densely granular tegument, characters inconsis-
tent with the “thorellii” clade. Material examined
during the present study conrmed the presence
of two sympatric species of Centruroides in the
Jardín Botánico Alfredo Barrera, Puerto More-
los. One species, determined to be conspecic
with the material previously misidentied as C.
schmidti by Armas (1996), based on meristic
data, is a hitherto undescribed species, described
herein as C. yucatanensis. e other is presumed
to be C. sissomi, based on the fact that the pec-
tinal plate is not lobed and the tegument is
densely granular, as well as its dark orange color-
ation and larger size (40 mm). Other records of
C. schmidti and/or C. sissomi from the Yucatán
Península (Vázquez, 1999; Teruel et al., 2015a;
Ponce-Saavedra and Francke, 2019) are misiden-
tications of C. yucatanensis.
M E: MEXICO: Quintana
Roo: Município Benito Juárez: Puerto Morelos,
Jardín Botánico Alfredo Barrera, 20°50′42.1″N
86°54′12.9″W, 38 m, 4.vii.2007, G. Montiel, R.
Paredes, M. Ramírez, D. Chibras, and G. Bonilla,
1 ♂ (AMNH [LP 7597]), 2 juv. ♂ (CNAN
SC3984). Yu catán: Município Felipe Carrillo
Puerto: Cenote Chac-ha, 3.5 km N and 3 km E
of Kalacmul, 20°04′40.3″N 88°08′27.9″W, 23 m,
9.vii.2007, R. Paredes, D. Chibras, and G. Mon-
tiel, 1 ♀ (CNAN SC4004).
ACKNOWLEDGMENTS
We thank the following for assisting with eld-
work or donating material used in the study: L.
Allen, A.J. Ballesteros, D. Barrales-Alcalá, M.A.
Barrios-Izás, P. Berea, G. Bonilla, J.L. Castelo, D.
Chibras, M. Cordóva, M. Escalante, T. Gearheart,
E. González-Santillán, J. Gorneau, J.H. Hu, A.
Jaimes, M.K. Lippey, S. Longhorn, R. Monjaraz-
Ruedas, H. Montaño, G. Montiel-Parra, D. Ortíz,
R. Paredes, J. Ponce-Saavedra, M. Ramírez, M.E.
Soleglad, A. Tietz, A. Valdez, M. van Dam, G. Vil-
legas, C. Víquez, and H. Yamaguti; R. Coates Lutes
for use of the Los Tuxtlas Field Station; P. Sierwald
and S. Ware (FMNH) for providing photographs
of the holotype of C. schmidti; E.S. Volschenk for
providing unpublished morphological characters;
D. Casellato, P. Rubi, and T. Sharma for generating
some of the DNA sequence data at the AMNH; A.
Lam, M. van Dam, L. Bonomo, and S.F. Loria for
assistance with DNA sequencing and phylogenetic
analysis at the CAS; P. Colmenares for logistical
support with collections at the AMNH; S. ur-
ston (AMNH) for assisting with UV digital pho-
tomicrography and preparation of the plates for
this contribution; and two anonymous reviewers
for constructive comments on a previous dra of
the manuscript. Research presented herein com-
prised part of the M.S. thesis of A.M. at San Fran-
cisco State University and the CAS, partially
supported by a grant from the Vincent Roth Fund
for Systematic Research of the American Arach-
nological Society, and a Collections Study Grant
from the AMNH. Some data presented herein
emanated from the Ph.D. dissertation of L.A.E. at
the City University of New York (CUNY) and the
2021 GOODMAN ET AL.: REVISION OF “THORELLII” CLADE OF CENTRUROIDES 77
AMNH, supported by a U.S. National Science
Foundation (NSF) GK-12 Fellowship, a CUNY/
NSF AGEP Grant, a CUNY Presidential Fellow-
ship, a CUNY College Now Fellowship, and an
NSF Postdoctoral Fellowship (1003087). Addi-
tional funding for this research was provided by a
grant from the eodore Roosevelt Memorial
Fund of the AMNH to L.A.E., an Ernst Mayr
Award from the Museum of Comparative Zool-
ogy, Harvard University, to L.A.E., an NSF Doc-
toral Dissertation Improvement Grant (DEB
0910147) to L.P. and L.A.E., NSF grant DEB
0413453 to L.P., and a grant from the Richard
Lounsbery Foundation to L.P.
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APPENDIX 1
Tissue samples, base-pair lengths, localities, and GenBank accession codes of DNA sequences from
the mitochondrial cytochrome c oxidase subunit I gene used for phylogenetic analysis of the arboreal
Neotropical “thorellii” clade of Centruroides Marx, 1890, bark scorpions (Buthidae C.L. Koch, 1837) and
outgroup species of Centruroides and Heteroctenus junceus (Herbst, 1800). Material deposited in the
following collections: Ambrose Monell Cryocollection (AMCC) at the American Museum of Natural
History, New York; Colección Nacional de Arácnidos, Instituto de Biología (CNAN), Universidad Nacio-
nal Autónoma de México, Mexico City; California Academy of Sciences (CASENT), San Francisco.
Oxford University Museum of Natural History (OUMNH), U.K. Sequences less than 150 base pairs in
length deposited in the Dryad digital repository (doi: 10.5061/dryad.tdz08t2).
Species Collection Locality L ength GenBank Code
Outgroup
H. junceus AMCC [LP 12613] Cuba: Guantánamo: Humboldt N. P. 1078 KY982192.1
C. bani AMCC [LP 3302] Puerto Rico: Isla Mona 1078 MK479164.1
C. exilicauda AMCC [LP 1692] Mexico: Baja California Sur: Cabo San Lucas 1078 KY982179.1
C. gracilis AMCC [LP 1550] Mexico: Veracruz: Los Idolos 1078 MK479175.1
C. hentzi AMCC [LP 1673] United States of America: Florida 1078 MK479177.1
C. infamatus AMCC [LP 1822] Mexico: Guanajuato: Acámbaro 1078 KY982181.1
C. ochraceus AMCC [LP 7666] Mexico: Morelos: Puerto Morelos Bot. Gard. 1078 MK479194.1
C. thorellii AMCC [LP 5983] Guatemala: Sacatepéquez: Parque Alux 1078 MK479208.1
OUMNH [MID166] Honduras: Cortés: San Pedro Sula 642 MZ366335
658 MZ366336
C. tuxtla AMCC [LP 3709] Mexico: Chiapas: La Vuelta de alacran 1078 MK479209.1
Ingroup
C. berstoni CASENT 9073271 Guatemala: Izabal: Biotopo Chocón Machacas 430 MZ366346
CASENT 9073272 Guatemala: Izabal: Hotel Tijax 621 MZ366345
CNAN SC3968 Guatemala: Izabal: Morelos 658 MZ366344
C. catemacoensis AMCC [LP 2070] Mexico: Veracruz: Los Tuxtlas 1078 MZ429054
AMCC [LP 5231] 1078 MZ429055
CASENT 9073270 659 MZ366343
CASENT 9073408 648 MZ366342
CASENT 9073410 658 MZ366341
CASENT 9073427 659 MZ366340
CASENT 9073428 648 MZ366339
C. chanae AMCC [LP 2009] Mexico: Michoacán: Faro de Bucerias 1078 MZ429056
AMCC [LP 7032] 1078 MZ429057
AMCC [LP 8582] Mexico: Guerrero: Microondas Fogos 1078 MZ429058
2021 GOODMAN ET AL.: REVISION OF “THORELLII” CLADE OF CENTRUROIDES 85
Species Collection Locality L ength GenBank Code
C. cuauhmapan AMCC [LP 2073] Mexico: Veracruz: Cañada Blanca 1078 MZ429059
CNAN SC4001 Mexico: Oaxaca: Cerro del Oro 40 –
CNAN T01397 Mexico: Veracruz: Los Idolos 127 –
C. hamadryas AMCC [LP 2948] Mexico: Chiapas: La Galleta 1078 MZ429060
CNAN SC3986 127 –
CNAN SC3988 127 –
C. homanni AMCC [LP 5224] Mexico: Chiapas: Las Delicias 1078 MZ429061
AMCC [LP 5249] Mexico: Chiapas: Santa Rosa 1078 MZ429062
AMCC [LP 5350] Mexico: Chiapas: Res. Biosfera Sepultura 1078 MK479178.1
CNAN SC3990 Mexico: Chiapas: Siltepec 134 –
CNAN SC3992 Mexico: Chiapas: P. N. Lagunas de Montebelo 137 –
CNAN SC3997 Mexico: Chiapas: Gutierrez 134 –
CNAN SC3998 Mexico: Chiapas: Villa Corzo 138 –
C. rileyi AMCC [LP 6445] Mexico: San Luis Potosí: Axtlan de Terrazas 1078 KY982183.1
CNAN SC3985 Mexico: Veracruz: Cerro Azul 127 –
CNAN SC3999 Mexico: Puebla: Cuetzalan 127 –
CNAN SC4000 Mexico: Veracruz: Papantla 127 –
CNAN SC4002 Mexico: Veracruz: Cerro Azul 127 –
C. schmidti AMCC [LP 13416] Honduras: Atlántida: Pico Bonito 127 –
AMCC [LP 13411] Honduras: Isla del Bahía: Cayos Menor 1078 MZ429064
AMCC [LP 13417] 1078 MZ429065
AMCC [LP 5985] Guatemala: Zacapa: Aldea casas de Pinto 1078 MZ429063
AMCC [LP 9172] Honduras: Francisco Morazán: E.A.P. Zamorano 1078 KY982184.1
CASENT 9073316 Guatemala: Zacapa: Guadalupe 606 MZ366338
CASENT 9073402 Guatemala: Zacapa: Las Minas 620 MZ366337
C. yucatanensis AMCC [LP 7597] Mexico: Quintana Roo: Puerto Morelos 1078 MK479201.1
CNAN SC3984 127 –
CNAN SC4004 Mexico: Yucatán: Cenote Chac-ha 127 –
APPENDIX 1 continued
86 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 452
APPENDIX 2
Morphological characters and character states used in phylogenetic analysis of the arboreal Neo-
tropical “thorellii” clade of Centruroides Marx, 1890, bark scorpions (Buthidae C.L. Koch, 1837) and
outgroup species of Centruroides, and Heteroctenus junceus (Herbst, 1800). Morphological terminol-
ogy follows Hjelle (1990) and Sissom (1990), except for carapace and metasomal carination, which
follows Vachon (1952), trichobothria, which follows Vachon (1974), tergite and pedipalp carination,
which follows Prendini (2000a), book lung structure, which follows Kamenz and Prendini (2008), and
ovariuterine anatomy, which follows Volschenk et al. (2008).
Carapace
1. Lateral ocular carinae: 0, present, distinct; 1, reduced to several granules; 2, absent (E.S. Volschenk
and L. Prendini, unpublished data; Esposito et al., 2017, 2018).
2. Centrolateral carinae: 0, present; 1, absent (E.S. Volschenk and L. Prendini, unpublished data;
Esposito et al., 2017, 2018).
3. Anterior centrosubmedian carinae: 0, present; 1, absent (E.S. Volschenk and L. Prendini, unpub-
lished data; Esposito et al., 2017, 2018).
4. Posterior centrosubmedian carinae: 0, present; 1, absent (E.S. Volschenk and L. Prendini, unpub-
lished data; Esposito et al., 2017, 2018).
5. Anteromedian notch: 0, present; 1, absent (Esposito, 2011).
6. Surface granulation density: 0, smooth; 1, sparsely granular; 2, densely granular medially; 3,
densely granular throughout (Esposito, 2011).
7. Surface granulation texture: 0, weakly granular, shagreened; 1, large, rounded granules; 2, large,
conical granules (Esposito, 2011).
8. Anterior median ocular sulcus: 0, absent; 1, wide; 2, narrow, deep (Esposito, 2011).
9. Median ocular sulcus: 0, absent; 1, wide; 2, narrow, deep (Esposito, 2011).
10. Posteromedian sulcus: 0, absent; 1, wide; 2, narrow, deep (Esposito, 2011).
11. Posteromarginal sulci: 0, absent; 1, present (Esposito, 2011).
12. Posterolateral sulci: 0, absent; 1, present (Esposito, 2011).
13. Anterior margin, carina: 0, absent; 1, smooth; 2, granular (Esposito, 2011).
14. Lateral margins, carina: 0, absent; 1, smooth; 2, granular (Esposito, 2011).
15. Posterior margin, carina between posterior centrosubmedian carinae: 0, absent; 1, smooth; 2,
granular (Esposito, 2011).
Pedipalps
16. Chela prodorsal carina: 0, granular; 1, smooth; 2, absent (E.S. Volschenk and L. Prendini, unpub-
lished data; Esposito, 2011; Esposito et al., 2017, 2018).
17. Chela retrodorsal carina: 0, granular; 1, smooth; 2, absent (E.S. Volschenk and L. Prendini, unpub-
lished data; Esposito, 2011).
18. Chela retromedian carinae: 0, granular; 1, smooth; 2, absent; 3, vestigial, reduced to several sparse
granules; ?, unknown (E.S. Volschenk and L. Prendini, unpublished data; Esposito, 2011; Esposito
et al., 2017, 2018).
19. Chela retroventral accessory carina: 0, complete; 1, reduced; 2, absent (E.S. Volschenk and L.
Prendini, unpublished data; Esposito, 2011).
20. Chela retroventral carinae: 0, granular; 1, smooth; ?, unknown (Esposito, 2011; Esposito et al.,
2017, 2018).
2021 GOODMAN ET AL.: REVISION OF “THORELLII” CLADE OF CENTRUROIDES 87
21. Chela retrodorsal accessory carinae: 0, present; 1, absent (Esposito, 2011; Esposito et al., 2017,
2018).
22. Chela prodorsal accessory carinae: 0, present; 1, absent (Esposito, 2011; Esposito et al., 2017,
2018).
23. Patella dorsomedian carina: 0, present; 1, absent (Esposito, 2011).
24. Femur retromedian carinae: 0, small granules; 1, large, conical granules (Esposito, 2011).
25. Fixed nger, number of median denticle subrows: 0, eight; 1, nine; 2, 10 or more; 3, seven plus
fused proximal subrow; 4, six plus fused proximal subrow (Esposito, 2011; Esposito et al., 2017,
2018).
26. Fixed and movable ngers, supernumary granules; 0, absent; 1, present (Soleglad and Fet, 2003;
Esposito, 2011; Esposito et al., 2017, 2018).
27. Movable nger, number of median denticle subrows: 0, eight; 1, nine; 2, eleven; 3, thirteen or
more; 4, seven plus fused proximal subrow (Soleglad and Fet, 2003; Prendini, 2004; Esposito, 2011;
Esposito et al., 2017, 2018).
28. Chela shape (♂): 0, incrassate (bulbous or swollen); 1, slender (Prendini, 2001b, 2004; Esposito,
2011; Esposito et al., 2017, 2018).
29. Chela shape (♀): 0, incrassate (bulbous or swollen); 1, slender (Prendini, 2001b; Esposito, 2011;
Esposito et al., 2017, 2018).
30. Chela movable nger, proximal lobe (♂): 0, present; 1, absent (Prendini, 2001b; Esposito, 2011;
Esposito et al., 2017, 2018).
31. Patella prolateral surface, setation: 0, long, dense setae; 1, short, sparse setae (E.S. Volschenk and
L. Prendini, unpublished data; Esposito, 2011).
Legs
32. Leg I, tarsal setation: 0, short, dense setae; 1, long, dense setae; 2, sparse setae (E.S. Volschenk and
L. Prendini, unpublished data; Esposito, 2011; Esposito et al., 2017, 2018).
33. Leg IV, tarsal setation: 0, short, dense setae; 1, long, dense setae; 2, sparse setae (Esposito, 2011;
Esposito et al., 2017, 2018).
34. Legs I–IV, trochanter lateral margin carina: 0, absent; 1, smooth; 2, granular (Esposito, 2011).
35. Telotarsal ungues: 0, long, slightly curved; 1, hooked (Esposito, 2011).
Pectines
36. Pectinal tooth shape: 0, elongate; 1, rounded, spadelike (Esposito, 2011; Esposito et al., 2017,
2018).
37. Proximal teeth, nodules on dorsal surface: 0, one; 1, multiple; 2, absent (Esposito, 2011; Esposito
et al., 2017, 2018).
38. Dorsal fulcra: 0, present; 1, reduced (Esposito, 2011).
39. Tympanumlike expansion between lamellae and teeth: 0, absent; 1, present (Esposito, 2011).
40. Proximal fulcra, setal count: 0, one; 1, two; 2, three; 3, four; 4, six or more; 5, none; ?, unknown.
(Esposito, 2011; Esposito et al., 2017, 2018).
41. Pectinal plate anterior margin, sulcus (♂): 0, present; 1, absent (Esposito, 2011; Esposito et al.,
2017, 2018).
42. Pectinal plate, posterior margin (♂): 0, straight; 1, convex; 2, concave (Esposito, 2011; Esposito et
al., 2017, 2018).
43. Median pectinal plate depression (♂): 0, present; 1, absent (Esposito, 2011; Esposito et al. 2017,
2018).
88 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 452
44. Lateral pectinal plate depression (♂): 0, present; 1, absent (Esposito, 2011).
45. Pectinal plate shape (♂): 0, square; 1, rectangular; 2, trapezoidal (Esposito, 2011).
46. Pectinal plate anterior margin, sulcus (♀): 0, present; 1, absent; ?, unknown (Esposito, 2011).
47. Pectinal plate posterior margin (♀): 0, straight; 1, slightly convex; 2, prominently rounded; 3,
concave; ?, unknown (Esposito, 2011).
48. Pectinal plate depressions (♀): 0, absent; 1, single wide median depression; 2, paired lateral
depressions; 3, single small, deep median depression (pinhole); ?, unknown (Esposito, 2011).
Sternites
49. Sternite VI, ventromedian carina: 0, absent; 1, granular; 2, smooth (E.S. Volschenk and L. Pren-
dini, unpublished data; Esposito, 2011; Esposito et al., 2017, 2018).
50. Sternite V, setation (♂): 0, absent; 1, present, setal base not situated in pits (surface smooth); 2,
present, setal base situated in pits (surface punctate) (Esposito, 2011).
51. Sternite VI, ventrolateral carinae: 0, absent; 1, reduced to single granule; 2, present, more than one
granule (E.S. Volschenk and L. Prendini, unpublished data; Esposito, 2011; Esposito et al., 2017, 2018).
Tergites
52. Tergites III–VI, dorsolateral carinae: 0, present; 1, absent (E.S. Volschenk and L. Prendini, unpub-
lished data; Esposito, 2011; Esposito et al., 2017, 2018).
53. Tergites III–VI, dorsosubmedian carinae: 0, absent; 1, vestigial; 2, distinct (Prendini, 2004;
Esposito, 2011; Esposito et al., 2017, 2018).
54. Tergite VII, median carina: 0, narrow, granular carina; 1, broad, granular carina; 2, broad, smooth
carina; 3, vestigial (E.S. Volschenk and L. Prendini, unpublished data; Esposito, 2011; Esposito et
al., 2017, 2018).
Metasoma
55. Segment II, median lateral carinae: 0, complete; 1, posteriorly restricted; 2, absent (Esposito, 2011;
Esposito et al., 2017, 2018).
56. Segment III, median lateral carinae: 0, complete; 1, posteriorly restricted; 2, absent (Esposito,
2011; Esposito et al., 2017, 2018).
57. Segment III, dorsolateral carinae, posterior granules: 0, similar to preceding granules; 1, larger
than preceding granules, acuminate (E.S. Volschenk and L. Prendini, unpublished data; Esposito,
2011; Esposito et al., 2017, 2018).
58. Segment IV, median lateral carinae: 0, absent or obsolete; 1, present (Esposito, 2011; Esposito et
al., 2017, 2018).
59. Segment V, posterior margin (anal rim) granulation: 0, present; 1, absent (Esposito, 2011; Esposito
et al., 2017, 2018).
60. Segment V, dorsolateral carinae: 0, present; 1, absent (Esposito, 2011; Esposito et al., 2017, 2018).
61. Segment V, median lateral carinae: 0, present; 1, absent (Esposito 2011; Esposito et al., 2017, 2018).
62. Segment V, ventrolateral carinae: 0, present; 1, absent; ?, unknown (Esposito, 2011).
63. Segment V, ventromedian carinae: 0, absent or obsolete; 1, present (Esposito, 2011; Esposito et al.,
2017, 2018).
64. Segment V, ventrosubmedian carinae: 0, absent or obsolete; 1, present (Esposito, 2011; Esposito
et al., 2017, 2018).
65. Segment V, ratio of segment length to width (♂): 0, slightly elongated, length less than 2× width;
1, moderately elongated, length 2.5–3× width; 2, markedly elongated, length more than 3× width
(Esposito, 2011; Esposito et al., 2017, 2018).
2021 GOODMAN ET AL.: REVISION OF “THORELLII” CLADE OF CENTRUROIDES 89
66. Segments I–IV, width: 0, narrowing posteriorly, segment I slightly wider than IV; 1, slightly widen-
ing posteriorly, segment I slightly narrower than IV; 2, markedly widening posteriorly, segment I
much narrower than IV (Esposito, 2011; Esposito et al., 2017, 2018).
67. Metasoma length relative to length of prosoma and mesosoma (♂): 0, similar or slightly greater;
1, 1.5–2×; 2, more than 2× (Esposito, 2011; Esposito et al., 2017, 2018).
68. Metasoma length relative to length of prosoma and mesosoma (♀): 0, similar or slightly greater;
1, 1.5–2×; 2, more than 2×; ?, unknown (Lamoral, 1978; Prendini 2001b, 2003; Esposito, 2011;
Esposito et al., 2017, 2018).
Tel s on
69. Telson shape and length (♂): 0, spherical, length similar to width; 1, slightly ovate, length approxi-
mately 1.5× width; 2, ovate, length more than 2× width (Esposito, 2011; Esposito et al., 2017,
2018).
70. Telson vesicle, width in relation to width of metasomal segment V (♂): 0, similar; 1, narrower; 2,
much narrower, less than half (Lamoral, 1978; Prendini 2001b, 2003; Esposito, 2011; Esposito et
al., 2017, 2018).
71. Telson vesicle, ventromedian carinae: 0, present; 1, absent (E.S. Volschenk and L. Prendini, unpub-
lished data; Esposito, 2011; Esposito et al., 2017, 2018).
72. Telson vesicle, ventrolateral carinae: 0, present; 1, absent (E.S. Volschenk and L. Prendini, unpub-
lished data; Esposito, 2011; Esposito et al., 2017, 2018).
73. Telson aculeus angle: 0, angled approximately 90° to vesicle; 1, angled less than 90° to vesicle
(Esposito, 2011).
74. Telson vesicle, proximal margin, notch: 0, absent; 1, present, unmodied; 2, present, projecting
vertically (Esposito, 2011).
75. Telson vesicle, lateral lobes: 0, absent; 1, present (Esposito, 2011).
76. Telson ventral surface: 0, granular; 1, smooth (Esposito, 2011).
77. Telson subaculear tubercle: 0, distinct, singular; 1, distinct, bifurcate; 2, obsolete, slight protuber-
ance; 3, absent (Lamoral, 1980; Stockwell, 1989; Prendini, 2001b, 2003; Esposito, 2011; Esposito
et al., 2017, 2018).
Total length
78. Total length, sexual dimorphism: 0, male shorter than or similar to female; 1, male much longer
than female; ?, unknown (Esposito, 2011; Esposito et al., 2017, 2018).
Ovariuterus
79. Number of loops (“cells”): 0, eight; 1, nine; ?, unknown (Volschenk et al., 2008; Esposito, 2011;
Esposito et al., 2017, 2018).
80. Loop shape: 0, simple; 1, complex bridged; ?, unknown (Volschenk et al., 2008; Esposito, 2011;
Esposito et al., 2017, 2018).
Book lungs
81. Lamellar surface: 0, slender venation; 1, ribbed venation; ?, unknown (Kamenz and Prendini,
2008; Esposito, 2011; Esposito et al., 2017, 2018).
82. Lamellar edge: 0, thorns; 1, smooth or wrinkled; ?, unknown (Kamenz and Prendini, 2008;
Esposito, 2011; Esposito et al., 2017, 2018).
83. Spiracle, posterior margin: 0, hilocks; 1, subconical; ?, unknown (Kamenz and Prendini, 2008;
Esposito, 2011; Esposito et al., 2017, 2018).
90 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 452
Color and infuscation
84. Cheliceral manus, reticulate infuscation: 0, present; 1, laterally restricted; 2, absent (Esposito,
2011).
85. Pedipalp segments, color pattern: 0, chela manus and patella similar to femur; 1, chela manus
darker than femur; 2, chela manus and patella darker than femur; 3, chela manus paler than femur
(Esposito, 2011).
86. Pedipalp femur, mottled infuscation: 0, present; 1, absent (Esposito, 2011).
87. Pedipalp patella, mottled infuscation: 0, present; 1, absent (Esposito, 2011).
88. Pedipalp chela manus, mottled infuscation: 0, present; 1, absent (Esposito, 2011).
89. Pedipalp chela ngers, mottled infuscation: 0, present; 1, absent (Esposito, 2011).
90. Tergites I–VI, mottled infuscation: 0, present; 1, absent (Esposito, 2011).
91. Tergites I–VI, lateral band of infuscation, shape: 0, absent; 1, narrow; 2, distinct, rectangular; 3,
wide, almost touching lateral margins (Esposito, 2011).
92. Tergites I–VI, lateral band of infuscation, intensity: 0, absent; 1, faint; 2, mottled; 3, distinct
(Esposito, 2011).
93. Tergites I–VI, infuscation: 0, eye-shaped pattern; 1, absent (Esposito, 2011).
94. Tergites I–VI, lateral margins infuscation: 0, distinct black line; 1, absent; ?, unknown (Esposito,
2011).
95. Tergites I–VI, median stripe of infuscation: 0, absent; 1, present (Esposito, 2011).
96. Tergite VII, color pattern: 0, similar to other tergites; 1, paler than other tergites (Esposito, 2011).
97. Carapace, mottled infuscation: 0, present; 1, absent (Esposito, 2011).
98. Carapace, bands of infuscation: 0, absent; 1, two broad bands; 2, four narrow lines (Esposito,
2011).
99. Carapace, interocular triangle infuscation: 0, present; 1, absent (Esposito, 2011).
100. Metasomal segments I–V, ventral surfaces, mottled infuscation: 0, present; 1, absent (Esposito,
2011).
101. Metasomal segments I–V, lateral surfaces, mottled infuscation: 0, present; 1, absent (Esposito,
2011).
102. Metasomal segments I–V, ventromedian stripe of infuscation: 0, present; 1, absent (Esposito,
2011).
103. Metasomal segment V, color pattern: 0, similar to preceding segments; 1, darker than preceding
segments (Esposito, 2011; Esposito et al., 2017, 2018).
104. Telson, lateral bands of infuscation: 0, present; 1, absent (Esposito, 2011).
105. Telson, median stripe of infuscation: 0, complete; 1, posteriorly conned; 2, absent (Esposito,
2011).
106. Telson, mottled infuscation: 0, present; 1, absent (Esposito, 2011).
107. Sternites III–VI, mottled infuscation: 0, present; 1, absent (Esposito, 2011).
108. Sternite V, pale surface (♂): 0, present; 1, absent; ?, unknown (Esposito, 2011).
109. Sternite V, pale surface (♀): 0, present; 1, absent; ?, unknown (Esposito, 2011).
110. Sternite VII, color pattern: 0, similar to other sternites; 0, paler than preceding sternites; 2, darker
than preceding sternites (Esposito, 2011).
111. Legs I–IV, dorsal surfaces, mottled infuscation: 0, present; 1, absent (Esposito, 2011).
112. Legs I–IV, ventral surfaces, mottled infuscation: 0, present; 1, absent (Esposito, 2011).
2021 GOODMAN ET AL.: REVISION OF “THORELLII” CLADE OF CENTRUROIDES 91
APPENDIX 3
Distribution of 112 morphological characters scored for phylogenetic analysis of the arboreal Neo-
tropical “thorellii” clade of Centruroides Marx, 1890, bark scorpions (Buthidae C.L. Koch, 1837) and
outgroup species of Centruroides and Heteroctenus junceus (Herbst, 1800). Character states scored 0–5,
unknown (?), or polymorphic [].
Outgroup
Heteroctenus junceus
2101131122 1120211321 0100011011 1212101013 0001112120 2000121000
0111020011 01120021?? 0101021011 1011001001 1111211110 11
Centruroides arctimanus
1010010011 0120221221 0000010?1? 111111210? 00111???20 2121000010
0110102?10 0111000??? ???1100000 1210000100 0110011110 00
Centruroides bani
0110021222 1120011121 0001010001 1011112000 0110011222 2112110000
0110101010 01110031?? ???1101001 1[01]10000 0011001111 11201
Centruroides exilicauda
0100030202 1120200010 0000010111 0000200000 0010113110 2110220011
0000201010 11110130?? 0111021111 1011101010 1101211110 11
Centruroides gracilis
1000011112 1120201321 0100111111 1212202000 0110003120 2120220011
0110201020 1102100100 0111001111 101?001011 1110011012 11
Centruroides hentzi
2010030122 0120200020 0000014110 1010102000 0101101112 2111010010
0111202020 00110000?? ???1101011 1210000000 1101211000 01
Centruroides infamatus
1000031212 1120200011 0000010111 1010102000 0010111111 2110121010
0110100010 01010020?? ???1001111 1210101011 1101011?12 11
Centruroides ochraceus
0000031202 1100211111 01000101?1 0000102000 00102???22 2110221011
0110101?21 01010100?? ???1101111 1010011011 1101211110 11
Centruroides thorellii
0000021101 1120200000 0100010110 0101102000 0010112301 2100120011
0110201020 01011001?? ???1100011 1301100010 0110000002 01
Centruroides tuxtla
0100131222 1120100021 0000010111 1011102005 0110202322 2111120011
0101102010 1112010??? ???1000011 1210100110 0010000012 01
Ingroup
Centruroides berstoni
1110010111 1000000010 00010101?0 1010012115 0110002111 2112220011
0?00202?10 01110100?? ???1100000 0110000000 01110101?0 01
Centruroides catemacoensis
1110010222 1021200010 0001010110 1010010115 0010202111 2110020010
0110202120 00100100?? ???1100000 0110000000 0111210000 01
92 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 452
Centruroides chanae
1110010111 1021200000 0001314110 1111010005 00112???12 2102220011
0?10202120 0111010??? ???1110000 0010000000 0111211010 01
Centruroides cuauhmapan
1110010110 1120200010 0000314110 1010012010 0111202121 2103120011
0?10202010 01110100?? ???1100000 0010000000 0111010000 01
Centruroides hamadryas
1110010111 1000000010 00010101?0 1010012115 0110002111 2112220011
0?00202?10 01110100?? ???1100000 0110000000 01110101?0 01
Centruroides homanni
0100010111 1121000010 0000010110 1111012005 1110112110 2103220011
0?00202020 01110100?? ???1100000 0210000010 0110211010 01
Centruroides rileyi
1110010110 1120200010 0000314110 1010012010 0111202121 2103120011
0?10202010 01110100?? ???1100000 0010000000 0111010000 01
Centruroides schmidti
1110010111 0120200010 0101010110 1010012010 0100012122 2102220011
0000202120 11110101?? 0111100000 0010000000 0101211010 00
Centruroides yucatanensis
1110010111 1120000010 0001014110 1010012105 0010012021 2103020011
0?10201120 0111010??? ???1100000 0010000000 0110010112 01
Scientific Publications of the American Museum of Natural History
American Museum Novitates
Bulletin of the American Museum of Natural History
Anthropological Papers of the American Museum of Natural History
Publications Committee
Robert S. Voss, Chair
Board of Editors
Jin Meng, Paleontology
Lorenzo Prendini, Invertebrate Zoology
Robert S. Voss, Vertebrate Zoology
Peter M. Whiteley, Anthropology
Managing Editor
Mary Knight
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On the cover: Centruroides berstoni, sp. nov., ♂, from the type
locality, Izabal, Rio Dulce, Hotel Tijax, Guatemala.
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is paper meets the requirements of ANSI/NISO Z39.48-1992 (permanence of paper).
