Systematic_revision_of_Mexican_threatened_tarantulas_Brachypelma

Zoological Journal of the Linnean Society, 2019, XX, 1–66. With 46 figures.
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Systematic revision of Mexican threatened tarantulas
Brachypelma (Araneae: Theraphosidae: Theraphosinae),
with a description of a new genus, and implications on
the conservation
JORGE MENDOZA1,2* and OSCAR FRANCKE2
1
Posgrado en Ciencias Biológicas, Universidad Nacional Autónoma de MEXICO, Av. Universidad 3000,
C.P. 04510, Coyoacán, Distrito Federal, MEXICO
2
Colección Nacional de Arácnidos, Módulo D planta baja, Departamento de Zoología, Instituto de
Biología, Universidad Nacional Autónoma de MEXICO, 3er circuito exterior, Apto. Postal 70–153, CP
04510, Ciudad Universitaria, Coyoacán, Distrito Federal
Received 11 May 2018; revised 3 April 2019; accepted for publication 11 May 2019
The tarantula genus Brachypelma includes colourful species that are highly sought after in the commercial pet trade.
They are all included in CITES appendix II. We present phylogenetic analyses using molecular and morphological
characters to revise Brachypelma, in which we include all currently known species. Our results agree with
a previous study that shows the non-monophyly of Brachypelma. Both phylogenies strongly favour the division
of Brachypelma into two smaller genera. The first clade (Brachypelma s.s.) is formed by B.albiceps, B. auratum,
B. baumgarteni, B. boehmei, B. emilia, B. hamorii, B. klaasi and B. smithi. The species included in the second clade
are transferred to the new genus Tliltocatl and is formed by T. albopilosum comb. nov., T. epicureanum comb.
nov., T. kahlenbergi comb. nov., T. sabulosum comb. nov., T. schroederi comb. nov., T. vagans comb. nov.
and T. verdezi comb. nov. Both genera can be differentiated by their coloration and the shape of the genitalia. We
transfer to Tliltocatl: T. alvarezi, T. andrewi and T. aureoceps, but should be considered as nomina dubia. In addition,
we transfer B. fossorium to Stichoplastoris. We discuss the implications of these taxonomical changes for CITES and
for the Mexican Laws for wildlife protection.
ADDITIONAL KEYWORDS: distribution – geography, genus revision – mitochondrial DNA – new genera –
phylogenetic nomenclature – phylogenetics.
INTRODUCTION 2018; Mendoza, 2014a). According to the World Spider

Catalog (2018), there are currently 1001 described
Tarantulas (family Theraphosidae) are the largest
species. Many of them are colourful in appearance and
and heaviest spiders of the world. They can inhabit
docile in captivity, making them attractive pets for
almost every terrestrial ecosystem, with the exception
collectors (West, 2005). Some of the most appreciated by
of polar areas, but they are mainly found in tropical,
enthusiasts are those of the genus Brachypelma due to
subtropical, semi-arid and arid regions around the
their longevity, size and attractive coloration. Because
world. They can be found from sea level to 4000 m
of the high demand for specimens of Brachypelma
elevation and some even can be found inside caves
smithi (Pickard-Cambridge, 1897) in the late 1980s,
down to –800 m deep (Bond et al., 2012; Lüddecke et al.,
and with the concern that they might be in danger,
authorities placed this species on appendix II of the
Convention on International Trade and Endangered
*Corresponding author. E-mail: nomeireth@hotmail.com
[Version of permanent record, published online 4 November Species (CITES). To prevent the same problem from
2019; http://zoobank.orgurn:lsid:zoobank.org:pub:E4D09A17- arising with other Brachypelma species, the inclusion
444F-45A0-95DB-059ECA175569] of the entire genus in CITES appendix II was
© 2019 The Linnean Society of London, Zoological Journal of the Linnean Society, 2019, XX, 1–66 1
2 J. MENDOZA and O. FRANCKE
proposed (Smith, 1994; West, 2005). Later, the World how that error could have been made in 1905 when
Conservation Monitoring Centre (1996) evaluated the Koch collection was labelled, after it was acquired
and included B. smithi as a near-threatened species by the NHM. It was Schmidt (1992a) who removed it
in the International Union for Conservation of Nature from synonymy of Euathlus and restored Brachypelma
(IUCN) Red List of Threatened Species. In recent (see Brachypelma andrewi Schmidt, 1992 section for
years, the Commission for Environmental Cooperation further details). Since then, nine more species have
has been working with authorities from Mexico, United been described: B. annitha Tesmoingt, Cléton & Verdez,
States and Canada to generate and implement action 1997, B. auratum Schmidt, 1992, B. baumgarteni Smith,
plans to promote the legal, sustainable and traceable 1993, B. boehmei Schmidt & Klaas, 1993, B. klaasi
trade in selected North American species that are Schmidt & Krausse, 1994, B. hamorii Tesmoingt
listed in appendix II of CITES. These actions were et al., 1997, B. schroederi Rudloff, 2003, B. verdezi
developed based on the information compiled from Schmidt, 2003 and B. kahlenbergi Rudloff, 2008. Three
consultation with stakeholders. Sixteen tarantula species originally described as Eurypelma have been
species, comprising one from the genus Aphonopelma transferred to Brachypelma: B. aureoceps (Chamberlin,
and 15 from the genus Brachypelma, were selected as 1917), B. epicureanum (Chamberlin, 1925) and
‘priority tarantula species’ and are the subject of one of B. sabulosum (Pickard-Cambridge, 1897). There
the action plans (CEC, 2017). was an attempt to include B. klaasi and B. albiceps
The genus Brachypelma was established by Eugene (formerly described as Brachypelmides ruhnaui
Simon in 1891, based on material that he believed to be Schmidt, 1997) in the genus Brachypelmides (Schmidt
Mygale emilia White, 1856, which was deposited in the & Krausse, 1994) based on the presumptive presence
Natural History Museum of Paris (Smith, 1994), and of a small pad of slightly plumose setae on the femur
subsequently it was designated as the type material IV and the sharply tapered embolous of the males.
(Rudloff, 2003). This species was originally listed from However, after examination of B. klaasi types, Smith
Panama, but this location is erroneous as its currently (1994) concluded that the species was a Brachypelma,
known natural distribution is north-western Mexico, but an ‘extreme form’ of the genus, considering
specifically in the Pacific Coast region (Smith, 1994; Brachypelmides as a junior synonym of Brachypelma.
Locht et al., 1999; Gabriel & Longhorn, 2015). Pickard- His conclusion was followed by Locht et al. (1999) who
Cambridge (1897) considered Brachypelma as a junior examined B. klaasi and B. ruhnaui and did not find any
synonym of Eurypelma and described E. smithi. This plumose setae on femur IV, and concluded also that,
was then accepted and followed by Simon (1903). It although the embolous of both species is sharper and
was Pocock (1903) who divided the genus Eurypelma more tapered, it does not differ significantly from the
into a number of genera and restored the validity of one in other Brachypelma. It was Locht et al. (2005)
Brachypelma. In his redescription of the genus, he who concluded that Brachypelmides ruhnaui is a junior
emphasized the presence of plumose hairs on the palp synonym of Brachypelma albiceps after revision of
trochanter and on the trochanter and femur of leg I, topotype material and specimens deposited in scientific
and he also included B. albiceps Pocock 1903, B. smithi collections. Recently, based on molecular characters,
and B. vagans (Ausserer 1875) in Brachypelma. Roewer Mendoza & Francke (2017) considered that B. annitha
(1942) then again suspended Brachypelma and restored is a junior synonym of B. smithi. The following year,
the genus Eurypelma. This change remained in place Turner et al. (2018) presented a mtDNA gene tree of
for several decades until Valerio (1980) re-established some tarantula spiders in which Brachypelma was
Brachypelma and included four additional species recovered as biphyletic.
[B. albopilosum Valerio, 1980, B. angustum Valerio, Currently, there are 18 described species of
1980, B. fossorium Valerio, 1980 and B. mesomelas (O. Brachypelma ranging from Mexico to Costa Rica
P.-Cambridge, 1892)], although only B. albopilosum and (Smith, 1994; Locht et al., 1999; Gabriel & Longhorn,
B. fossorium continue currently in Brachypelma. Then, 2015; WSC, 2018). Of these, 13 occur in Mexico,
Raven (1985) in his revision of the Mygalomorphae where 12 are endemic. The most northerly species is
concluded that Brachypelma was a junior synonym of Brachypelma emilia (White, 1856) found west of the
the older genus Euathlus Ausserer, 1875, based on the Sierra Madre Occidental mountain range in the states
study of a single specimen from the Koch collection in of Sonora, Sinaloa, Durango and Nayarit. This highly
The Natural History Museum, London (NHM). Smith colourful species is a fossorial burrower whose habitat
(1994) mentioned that an amateur arachnologist named ranges widely from the drier coastal thorn forests and
Frank Fritzlen presented an old document in 1991 savannahs, through the palm transition forests and
based on scientific descriptions and drawings, which wetter inland tropical deciduous forests, up into the
could indicate that a mistake may have taken place in higher and cooler elevations of the oak forests (West,
the collection of the NHM. Smith (1994) also described 2005). The southernmost species was Brachypelma
© 2019 The Linnean Society of London, Zoological Journal of the Linnean Society, 2019, XX, 1–66
SYSTEMATIC REVISION OF BRACHYPELMA 3
embrithes (Chamberlin, 1925), which is found in To date, no taxonomic revision has been made of
Panama, but Gabriel & Longhorn (2015) transferred the genus Brachypelma,despite its popularity in the
this species to the genus Sericopelma Ausserer, 1875. pet trade and being a priority group for conservation
Currently, the southernmost species in the genus is protected by international conventions such as
Brachypelma albopilosum from Costa Rica. CITES or the General Law of Wildlife (LGVS) in
Although correct species identification is a Mexico. Also, although geographical ranges have
fundamental component of many biological been published for Brachypelma species of Mexico
investigations, taxonomic expertise is shrinking and (Baerg, 1958; Valerio, 1980; Smith, 1994; West, 1996,
at the same time overwhelmed by demand. The correct 2005; Locht et al., 1999; Reichling, 2001, 2003; Rojo,
identification of a species based on morphological 2004; Arisqueta-Chablé et al., 2009; Shaw et al.,
characters can sometimes take too long or can be 2011; Hijmensen, 2012; Longhorn, 2014; Mendoza &
impossible for certain life stages, like immature spiders Francke, 2017), these ranges are general in scope or
or even for members of one sex (Greenstone et al., 2005; incomplete (Fig. 1). Furthermore, studies to determine
Blagoev et al., 2009). There are also some problems in the susceptibility, exact zoogeographical range and/
the identification of species in groups with similar or genome for each priority tarantula species have
patterns of convergence or morphological conservatism not been conducted (Garcia, 2016). The geographic
(Hebert et al., 2003; Locke et al., 2010; Niemiller distribution of Brachypelma shows a clear difference
et al., 2011; Hamilton et al., 2014, 2016). Theraphosid between the ‘red leg’ and ‘red rump’ species, the
spiders are known by having simple genitalia and tend first ones mostly distributed through the Mexican
to show relatively low interspecific and relatively high Pacific Coast and central Mexico and the second
intraspecific variation, which have revealed widespread ones distributed along the south-west of Mexico from
patterns of homoplasy among traditional taxonomic the Gulf of Mexico to the Pacific Coast in southern
characters (Raven, 1985; Goloboff, 1993; Pérez-Miles Guerrero and continuing southward to northern Costa
et al., 1996; Pérez-Miles, 2000; Bertani, 2001; Bond & Rica (Fig. 2). Most Brachypelma studies have focused
Opell, 2002; Bond & Hedin, 2006; Hedin & Bond, 2006; on a few taxa, as those of population interactions,
West et al., 2008; Hendrixson & Bond, 2009; Bond ethology, morphological variation and genetic diversity
et al., 2012; Guadanucci, 2014; Hamilton et al., 2014, in Brachypelma vagans (Ausserer, 1875) (Reichling,
2016; Perafán & Pérez-Miles, 2014; Ortiz & Francke, 2000; Longhorn, 2002; Machkour-M’Rabet et al., 2005,
2016; Fukushima & Bertani, 2017). An alternative 2007, 2011, 2012, 2015, 2017; Shillington & McEwen,
for a sustainable identification is the construction of 2006; Dor et al., 2008; Dor & Hénaut, 2011, 2012, 2013;
systems that employ DNA sequences as taxon barcodes Vilchis-Nestor et al., 2013; Hénaut et al., 2015), from
(Sun et al., 2012). Molecular markers, such as COI, courtship and habitat preference of Brachypelma
may provide species boundary information in certain klaasi (Schmidt & Krause, 1994) (Yáñez et al., 1999;
taxonomic groups and consequently have the potential Yáñez & Flotaer, 2000) and only a few studies have
to be a rapid and efficient means to delineate and tested the utility of molecular markers for species
identify species (Hebert et al., 2003; Barrett & Hebert, delimitation or identification of the priority tarantula
2005; Arnedo & Fernandez, 2007; Chen et al., 2011; species (Longhorn et al., 2007; Petersen et al., 2007;
Kuntner & Agnarsson, 2011; Hamilton et al., 2014; Machkour-M’Rabet et al., 2009; Mendoza & Francke,
Blagoev et al., 2016). However, molecular identification 2017; Turner et al., 2018).
is not free of problems and may have restrictions and In Mexico, wild populations of Brachypelma
inconsistencies. So, the use of molecular data, together tarantulas are in decline, due to habitat loss, because
with morphology, is recommended, along with collection people often kill them when encountered in the wild and
data and field observations to allow for a better because large numbers of some of the more colourful
species delimitation and proposal of phylogenetic species are collected, particularly from the Pacific
relationships (Scotland et al., 2003; Will & Rubinoff, Coast, to supply the national pet market. Field-caught
2004; Prendini, 2005; Bond & Stockman, 2008; Slowik tarantulas are either sold in traditional markets (with
& Blagoev, 2012; Hendrixson et al., 2015; Ortiz & little or no law enforcement) or exported illegally for
Francke, 2016). In the particular case of tarantulas, the commercial pet trade (García, 2016; CEC, 2017).
there are a limited number of studies that have used The sustainable use, conservation and management
molecular information for species delimitation, and of native tarantulas in Mexico is regulated via the
some of them found moderate to deep inconsistencies General Law of Wildlife (LGVS for its Spanish
in the morphology-based taxonomy due to homoplasy acronym) and its by-laws that allow the creation of
in some characters (Hamilton et al., 2011, 2014, 2016; the named Management Units for the Conservation
Hendrixson et al., 2013, 2015; Wilson et al., 2013; of Wildlife (UMA for its Spanish acronym). The UMA
Graham et al., 2015; Montes de Oca et al., 2015; Ortiz refers to the registered facilities that operate in
& Francke, 2016). accordance with an approved management plan. These
Figure 1. Previous distribution map of Mexican species of Brachypelma, from Locht et al. (1999). Note: Brachypelma
pallidum as was labelled originally belonged to Brachypelma verdezi, which here is transferred to Tliltocatl.
are intensive and extensive breeding sites where wild Mexico, have better results, it is important to have an
flora, fauna and fungi are reproduced and propagated, adequate description of tarantula species, to allow for
and products and by-products destined to different the prioritization of certain species for conservation.
types of use are generated. Their general objective Thus, the development of morphological guides and
is the conservation of natural habitat, populations molecular databases allowing correct identification is
and specimens of wild species. They function as of the utmost importance. In addition, it is necessary
centers of breeding stock, as germplasm banks, an to delimit their natural distribution areas and habitat
alternative ex situ conservation for reproduction preferences in order to develop better strategies for
of key species or species that are in any category of conservation of their populations, which will also
risk. For environmental education, research and allow selection of potential sites appropriate for in
production of species, parts and derivatives of wildlife situ repopulation programmes. Initially, it is the
that can be incorporated into the different circuits of aim of this work to revise the genus Brachypelma
the legal market for commercialization (SEMARNAT, Simon, 1891 and to test its monophyly by the use of
2000). Under the UMA programme, qualified persons morphological and molecular characters. Some species
may present a request to collect a limited number of were originally described based solely on one sex or
wild tarantulas to keep and breed in captivity. The even using only exoskeletons as type specimens, so
resulting offspring may then be sold domestically or we reviewed and illustrated specimens of both sexes.
exported, and may also be used for reintroduction Some new characters are introduced as a result of our
programmes. Few hobbyists, breeders or academics comparative morphological studies of these species
would be experienced in identifing each species of and their morphological variability. The redescription
priority tarantula on sight. To ascertain that the of Brachypelma species and the relevant taxonomic
objectives of protecting and conserving tarantulas, changes are made based on our results. Distribution
supported by international policies and those of
Figure 2. Distribution map of formerly known Brachypelma redleg (s.s.) and red rump (s.l.) species complex with more
accurate distribution areas based on museum specimens. Biogeographic regions of distribution for Mexican species is
indicated. Circles = red leg complex; squares = red rump complex; diamond = Brachypelma fossorium, which actually does
not belong to any of the mentioned groups.
maps for all known species are presented and general of stridulating setae follows Galleti & Guadanucci
descriptions of their habitat and life cycle are provided. (2018). The urticating setae distribution pattern over
the abdomen follows Bertani & Guadanucci (2013).
The GPS coordinates of the collection sites of each
specimen are omitted in order to avoid easy access
MATERIAL AND METHODS
to these places and to avoid the illegal collection of
The general descriptive format used in the present specimens given that these species are protected
study follows Mendoza & Francke (2017). All by law.
measurements are in millimetres and were taken using Photographs (Figs 5–7, 9–10, 11C, 12, 14–16, 18–
an ocular micrometre on a stereomicroscope Nikon 19, 20B, 21, 23–25, 27, 29–31, 33, 39C–D, 40) were
SMZ645 for smaller structures, and a digital calliper taken with a Nikon Coolpix S10 VR digital camera
with an error of 0.1 mm for larger ones. Leg and palp coupled to a stereomicroscope. Descriptions of colours
measurements were taken along the dorsal central use the standard names of the 267 Colour Centroids
axis of the left side. Description of tarsal scopulae of the NBS/IBCC Colour System (Mundie, 1995).
follows Pérez-Miles (1994). Male palpal bulb keels and The colours depicted herein are included to promote
tegular apophysis terminology follows Bertani (2000). some standardization in describing coloration of
Spermatheca shape follows the general description living animals. To avoid differences of perception in
format used with theraphosids (Pérez-Miles, 1989; coloration because of the calibration of the monitor,
Bertani, 2001; Mendoza & Francke, 2017). Description we extracted the RGB code and pantones colours from
the photographs using Photoshop CS Live. A digital 2010 and Xenesthis immanis (Ausserer, 1875). The
photograph was opened and the eyedropper tool trees were rooted on P. victori, selected because it
clicked over the image to obtain the RGB colour code belongs to subfamily Psalmopoeinae Schmidt, 2010
in the Set foreground colour and cross-checked in the (formerly Aviculariinae), making it an appropriate
colour libraries for the Pantone solid coat. Both habitat outgroup for Theraphosinae.
and laboratory images were taken with a Canon G12
Digital camera. The habitat shots were taken under
natural daylight conditions. The images with a white Character matrix and cladistics
background were taken in the laboratory where Sereno (2007) mentioned that characters are simple
general illumination was provided by one fluorescent characteristics expressed as independent variables
30-W light bulb held approximately 20 cm from the and that character states are the mutually exclusive
specimen. Once the RGB code was obtained, it was conditions of a character. Together, the characters
possible to infer the real colour of the specimen using and character states make up character statements.
the RGB code of Colour Centroids. The range of each Based on this definition, the coding of characters was
colour centroid as perceived by the human eye is wide carried out taking phylogenetic works carried out in
enough to account for errors of observation. other genera of the family as a starting point (Bertani,
Abbreviations used in the text are as follows: Ocular 2001; Fukushima, 2005; Pérez-Miles, 2000; Pérez-
patterns – ALE, anterior lateral eyes; AME, anterior Miles & Locht, 2003; West et al., 2008; West & Nunn,
median eyes; PLE, posterior lateral eyes; PME, 2010; Fukushima & Bertani, 2017). Intraspecific
posterior median eyes. Morphology – legs and palpi: variation of each Brachypelma species was determined
d, dorsal; p, prolateral; r, retrolateral; v, ventral; Rap, by reviewing between five and ten specimens of each
retrolateral tibial apophysis; Pap, prolateral tibial sex for each species, with the exception of Central
apophysis. Palpal bulbs: AK, apical keel; PI, prolateral American species, because it was not possible to
inferior keel; PS, prolateral superior keel; TA: tegular obtain additional samples to those already deposited
apophysis. Spermatheca: Bp, spermathecal baseplate. in scientific collections. The analysed characters were
Spinnerets: PLS, posterior lateral spinnerets; PMS, only of adult specimens of both sexes, because only
posterior median spinnerets. In species synonymies, mature individuals present diagnostic characters and
w e f o l l o w t h e Wo r l d S p i d e r C a t a l o g ( 2 0 1 8 ) : other morphological characters that allow formulation
D = Described, f = female, m = male, T = Transferred. of a homology hypothesis. The character matrix
Institutions: CNAN = Colección Nacional de comprised 103 characters of adult morphology scored
Arácnidos, MEXICO DF, MNHNP = Museum for 24 taxa; eight characters were non-informative
National d´Histoire Naturelle, Paris, NHM = Natural (Table 1; Supporting Information, Appendix S1).
History Museum, London, PROFEPA (acronym in Multistate characters were optimized as non-additive
Spanish) = Federal Environmental Protection Agency, (Fitch, 1971). In the absence of evidence that would
SNMF = Senckenberg Naturmuseum, Frankfurt, support the order in the multistate characters, the
UCR = Museo de Zoología Universidad de Costa heuristic searches were conducted in TNT v.1.1
Rica; UNAM = Universidad Nacional Autónoma de (Goloboff et al., 2008). The search method applied
MEXICO. to analyse was New Technology Search (Goloboff
et al., 2008) using Sectorial Search, Ratchet (5000
iterations and 2000 substitutions), with drifting (100
iterations) and Tree Fusing (ten rounds; dumping
Morphological protocol
fused suboptimal trees to prevent clogging); memory
Taxon sampling with a maximum of 30 000 trees was retained. The
The cladistic analysis was based on 24 taxa. The analysis was performed using equal weighting, the
ingroup included 16 species of Brachypelma. The length (L) of the optimal trees, consistency index (Ci)
outgroups were selected based on the cladistic analyses and retention index (Ri) are reported. Optimization
of subfamily Theraphosinae made by Pérez-Miles of ambiguous characters was carried out using
et al. (1996), Pérez-Miles (2000), Bertani (2001) and ACCTRAN (Accelerated Transformation), in order to
Perafán & Pérez-Miles (2014). The number of external disadvantage parallelisms and promote regressions.
groups used was eight, belonging to the following According to Agnarsson & Miller (2008), there are
taxa: Acanthoscurria geniculata (C. L. Koch, 1841), no grounds for preferring any algorithm to solve
Aphonopelma seemanni (F. O. P.-Cambridge, 1897), ambiguous optimizations, so the use of ACCTRAN
Bonnetina cyaneifemur Vol, 2000, Megaphobema is somewhat arbitrary and is only used for the sole
mesomelas (O. Pickard-Cambridge, 1892), Psalmopoeus purpose of presenting a hypothesis about a resolution
victori Mendoza, 2014b, Sericopelma melanotarsum of these ambiguities. The relative support for each
Valerio, 1980, Theraphosa stirmi Rudloff & Weinmann, node on the preferred hypothesis was calculated with
7
SYSTEMATIC REVISION OF BRACHYPELMA
Table 1. Characters matrix used in cladistic analysis of Brachypelma s.l. and s.s. (–) inapplicable (?) unknown; treated as missing data in the analysis.
bootstrap resampling (Felsenstein, 1985). Bootstrap tissue. Genomic DNA was extracted using the Qiagen
support was estimated with heuristic searches of 1000 DNeasy Tissue Kit, following the protocol of the
pseudoreplicates, with P = 50. Cladograms obtained by manufacturer. The concentration of the extracted DNA
TNT were exported in Tree file, to be properly edited was quantified with a spectrophotometer (Nanodrop
in WinClada Asado v.1.7 (Nixon, 1999–2004), mapping 2000 Techno Scientific) or visualized via agarose gel
all the characters on the consensus tree, and were electrophoresis. DNA amplification was performed
subsequently edited with Adobe Illustrator CS5. using the polymerase chain reaction (PCR) for the
mtDNA barcoding gene region COI. A single set of
primers was used: LCO 1490: 5’-GGTCAACAAATCA
Molecular protocol TAAAGATATTGG-3’, together with HCO 2198 5’-TAA
Taxa ACTTCAGGGTGACCAAAAAATCA-3’ (designed by
Specimens were collected throughout the known Folmer et al., 1994). This primer set amplified a 710-
distribution area of the genus Brachypelma with bp region of the mitochondrial cytochrome oxidase
special attention to the type localities (where possible). subunit 1 gene.
Material was fixed in 80% ethanol. The third leg on PCR reaction (100.8 µL) contained 48 µL 10× PCR-
the right side of each spider was stored in 96% ethanol buffer, 24 µL MgCl2, 7.68 µL of forward and reverse
at –20 °C. Tissue samples of 69 specimens were primer each, 9.6 µL dNTP’s and 3.84 µL Taq polymerase,
used for DNA extraction, representing 16 species of using 1 µL of the DNA template for each sample. PCR
Brachypelma. Five tissue samples each of Brachypelma program for COI followed initial denaturation at 94 °C
albiceps Pocock, 1903, B. auratum Schmidt, 1992b, for 2 min; 30 cycles of denaturation at 94 °C for 1 min,
B. baumgarteni Smith, 1993, B. emilia (White, 1856), annealing at 48 °C for 45 s, elongation at 72 °C for
B. epicureanum (Chamberlin, 1925), B. hamorii 2:30 min; followed by 7 min of final elongation at 72 °C.
Tesmoingt, Cléton & Verdez, 1997, B. kahlenbergi LCO 1490 and HCO 2198 primers used for single-
Rudloff, 2008, B. klaasi (Schmidt & Krause, 1994), stranded sequencing. The accuracy of sequences was
B. schroederi Rudloff, 2003 and B. smithi (F. O.-Pickard verified by independently amplifying and sequencing
Cambridge, 1897); four each of B. boehmei Schmidt the complementary strands of all fragments. Primer
& Klaas, 1993 and B. verdezi Schmidt, 2003; three sequences were removed and complementary strands
each of B. sabulosum (F. O.-Pickard Cambridge, of DNA assembled into consensus sequences, edited
1897) and B. vagans (Ausserer, 1875) and one each of and checked for quality using GENEIOUS R8 (Kearse
B. albopilosum Valerio, 1980, B. fossorium Valerio, 1980, et al., 2012). If complementary strands disagreed
Aphonopelma seemanni (F. O.-Pickard Cambridge, (besides minor mismatches), the sample was amplified
1897), Megaphobema mesomelas (O. Pickard- and sequenced again to resolve the discrepancies.
Cambridge, 1892) and Sericopelma melanotarsum
Valerio, 1980 were used in this study. Additionally,
DNA sequence alignment and phylogenetic
four sequences were retrieved from GenBank to use
analysis
as outgroups for phylogenetic analyses: Psalmopoeus
cambridgei Pocock, 1895 (JQ412455.1), Eupalaestrus Static alignments of COI gene fragments were
campestratus (Simon, 1891) (JQ412446.1) (both from generated with MAFFT online v.7 (Katoh et al., 2002,
Briscoe et al., 2013), Lasiodora parahybana Mello- 2005). The G-INS-I strategy, which performs a global
Leitão, 1917 (JN018128.1) (from Arabi et al., 2012) and alignment based on an FFT approximation, was
Xenesthis immanis (Ausserer, 1875) (MG273518) selected (Katoh et al., 2002). This method is suitable
(from Lüddecke et al., 2017). We obtained sequences for large datasets comprising sequences with relatively
of mitochondrial COI from 69 samples. Vouchers were limited variation in length, i.e. few short gaps (Katoh
deposited in CNAN and assigned a unique number et al., 2005). The scoring matrix for nucleotide
(CNAN-Ar00xxxx). All sequences were submitted sequences was set to 1/PAM K = 2, gap opening penalty
to GenBank. Accession numbers and specimen to 1.53 and offset value to 0.
information are given in Supporting Information, We carried out maximum likelihood inference using
Appendix S2. RAxML-HPC BlackBox on XSEDE (8.2.8) (Stamatakis
et al., 2008, Stamatakis, 2014) in the CIPRES
Science Gateway platform (Miller et al., 2010) under
DNA protocols GTR+gamma+Pinvar model of nucleotid evolution and
DNA isolation, PCR amplification and sequencing Bootstrap support obtained by running 1000 pseudo
were performed at the Laboratorio de Sistemática replicates. The relative supports are showed for each
Molecular, Instituto de Biología, UNAM. Muscle tissue clade on the best tree.
was extracted from the leg by removing ~20 mg of
Figure 3. Strict consensus of the two trees obtained by parsimony analysis of 103 morphological characters for 24 taxa
with equal weighting. Tree shows the non-monophyly of Brachypelma with the species that remain in the genus and the
ones transferred to Tliltocatl. Bootstrap support with percentages less than 100 indicated above branches. Synapomorphies
were mapped on the tree. Tree length 443, Ci: 0.443, Ri: 0.541.
RESULTS The second clade includes a group in the ‘red rump’

Brachypelma, which is formally described here as
Phylogenetic analysis
the new genus Tliltocatl, even though this group
Morphological analysis of 103 characters finds two is not well supported (P = 11). A small clade formed
most parsimonious trees (MPTs) (L = 433, Ci = 0.443, by Aphonopelma seemanni, Xenesthis immanis and
Ri = 0.541). The genus Brachypelma recovers as Sericopelma melanotarsum appears as its sister-
non-monophyletic, being separated into two clades. group. The strict consensus tree shows that Tliltocatl
The only difference between the two MPTs is the possesses one synapomorphy that is the embolus
position of Tliltocatl verdezi as a sister-species of prolateral inferior keel longitudinal and larger than
T. kahlenbergi, or as a sister-species to a small clade the prolateral superior keel (char86). The other
formed of T. kahlenbergi, T. epicureanum, T. vagans and nine characters that support the Tliltocatl clade are
T. sabulosum. The strict consensus tree shows the same homoplastic but can be used in the diagnosis of the
topology as MPTs, except for the collapse of the internal new genus: fovea in males and females procurved
clade that relates to T. verdezi and T. kahlenbergi due (char6), fovea in males and females pit-like (char7),
to the change of the position of T. verdezi (Fig. 3). The distal scopula of metatarsus III with 75% of coverage
first clade includes a group in the ‘red leg’ Brachypelma relative to its length (char13), femur I prolateral
(Brachypelma s.s.) that includes the type species of face setae claviform (char29), male femur III slightly
the genus B. emilia; although this group is not well incrassate (char33), patella I with one spine (char60),
supported (P = 15), the strict consensus tree shows embolus curved to retrolateral (char75), embolus
that Brachypelma has four synapomorphic characters: apical region broad (char78), and embolus apical shape
tibiae colour pattern completely orange (char35), very flattened concave/convex appearance with neck
embolus apical region slightly broad (char78), apical in the base (char79) (Fig. 3). In the case of the nominal
keel developed and slightly extended to backwards species ‘Brachypelma’ fossorium, it is shown that it
(char89) and spermatheca baseplate divided and does not belong to either Brachypelma or Tliltocatl,
narrower than base width (char100). The other three although in the strict consensus tree it appears to be
characters that support the Brachypelma s.s. clade are closely related to Tliltocatl. ‘Brachypelma’ fossorium
homoplastic but can be used in the diagnosis of the lacks important features present in Tliltocatl such as
genus: fovea shallow (char80), embolus retrolateral urticating setae type III, prolateral inferior keel longer
curvature only in apex (char75) and embolus apical than prolateral superior keel, femur III incrassate,
shape very flattened concave/convex appearance with percentage coverage of distal scopula of metatarsus III
neck in the base (char79) (Fig. 3). is not more than 50%. All these differences are enough
to propose the transference of B. fossorium to another species’ of Tliltocatl has a bootstrap support of 92
genus (see below). and is formed by seven species with relatively high
Maximum likelihood inference analysis of the COI support, but the internal relationships are unclear.
sequences also strongly supports the non-monophyly Tliltocatl schroederi is shown to be the sister-species
of Brachypelma and is congruent with the analysis of of the rest, but there is no support for the position
morphological characters. The final ML optimization of T. verdezi and T. kahlenbergi. An internal node
score is –5527.883771. The recovered topology follows with T. epicureanum as sister to T. sabulosum,
shows two clades: the first one formed by species T. albopilosum and T. vagans. However, no resolution
of Brachypelma ‘red leg complex’ that includes the is shown for interspecific relationships for the three
type species of the genus B. emilia (Brachypelma last species. ‘Brachypelma’ fossorium is clearly shown
s.s.) and a second one formed by species in the ‘red to be excluded from either Brachypelma or Tliltocatl,
rump complex’ (here transferred formally to the being closer to Sericopelma and Aphonopelma (Fig. 4);
new genus Tliltocatl; Fig. 4). The Brachypelma just as in the morphological analysis, differences are
s.s. clade has a bootstrap support value of 78, it is enough to transfer ‘B.’ fossorium to another genus as
formed by eight species with high bootstrap support. we propose in the taxonomical section of this paper.
Sericopelma melanotarsum, Aphonopelma seemanni, Despite the differences between the parsimony
Xenesthis immanis and Brachypelma fossorium and maximum likelihood analyses, both topologies
are shown to be more closely related to the new i n d i c a t e t h a t B r a ch y p e l m a s. l . , a s c u r r e n t l y
genus Tliltocatl. The clade formed by ‘red rump recognized, is not monophyletic, including two highly
Figure 4. Maximum likelihood phylogenetic hypothesis of 658 aligned nucleotides from barcoding gene COI of the
mitochondrial genomes of 66 samples from eight Brachypelma and seven Tliltocatl. species. Nodal support shows maximum
likelihood bootstrap. Tree shows the non-monophyly of Brachypelma with the species that remain in the genus and the ones
transferred to Tliltocatl.
divergent lineages. The species of both Brachypelma leg I trochanter/femur and on the retrolateral face
s.s. and Tliltocatl can be identified with high trochanter of the palp; (2) both sexes possess always
accuracy using COI as a barcode (probably due to the urticating setae types I and III; type III are located in
relatively high mtDNA divergence between species). the dorsoposterior area and type I surrounding these;
In both analyses inner relationships of some of these (3) the male palpal bulb distally wide and flattened
species require further resolution, and in the case of (spoon-shaped) and presents prolateral superior and
morphology, the homoplasy in some of the characters apical keels (Figs 6, 10, 15, 19, 24, 27A–D, 30, 33A–D)
can lead to not strongly supported topologies. On and can have a small or reduced prolateral inferior
the other hand, the molecular phylogenies based keel, except in B. albiceps, B. baumgarteni, B. emilia
on mtDNA have limitations as gene tree/species and B. klaasi; the apical keel can extend slightly (Fig.
tree incongruence, with COI representing only 6) or widely (Fig. 10) backwards; (4) females can have
one particular genealogy out of all possible in the a divided spermatheca with each lobe as large as wide
genome. [present in B. albiceps (Fig. 7E–G) and B. klaasi (Fig.
29D–F)], or a simple undivided/fused spermatheca
(Figs 11C, 12E–F, 16E–G, 20B, 21E–G, 25E–F, 27E–F,
Taxonomy 33E–H), apically narrowed (present in B. auratum,
Family Theraphosidae Thorell, 1869 B. baumgarteni, B. boehmei, B. emilia, B. hamorii and
B. smithi); (5) having a spermatheca baseplate divided
Subfamily Theraphosinae Thorell, 1870 and well sclerotized; (6) both sexes lack a plumose pad
Brachypelma Simon, 1891: 338 of setae on leg IV femur; and (7) all tarsi scopulae are
undivided.
(Figs 1–38) Brachypelma differs from Tliltocatl by the red/orange
Type species: Mygale emilia White, 1856: 183 (White, coloration pattern on legs and/or carapace (except
1857: 406; Simon, 1892: 168; Pocock, 1903: 103; B. albiceps) (Figs 36, 37, 41). The shape of genitalia also
Petrunkevich, 1939: 191; Schmidt, 1986: 49, 1991: 11; differs in both sexes with the male palpal bulb apex
Smith, 1986: 50, 1991: 17; Fritzlen, 1991: 14; Schmidt, shorter than in Tliltocatl and by the lacking of prolateral
1993: 82; Schmidt & Klaas, 1993: 7; Smith, 1993: 14, inferior keel or restricted to the apex when present (Figs
1994: 156; Pérez-Miles et al., 1996: 46; Tesmoingt, 6, 10), whereas in Tliltocatl it is posteriorly extended
Cleton & Verdez, 1997: 8; Vol, 2000: 1; Locht et al., and parallel to a prolateral superior keel (Fig 40A–C).
2005: 108; Gabriel & Longhorn, 2015: 85; Mendoza & It also differs in lacking any spination on patellae of
Francke, 2017: 161; Turner et al., 2018: 11) palps or legs. Females differs by the spermatheca apex
not as inwardly curved, being generally straight (except
Eurypelma C. L. Koch, 1850: 73. Type species: Aranea B. klaasi and B. albiceps due to divided spermatheca)
avicularia Linnaeus, 1758 (Simon, 1903: 936; Roewer, (Figs 7E–G, 12E–F, 40D). Brachypelma differs also by
1942: 238). First synonymized by Pickard-Cambridge having the spermatheca baseplate more developed and
(1897: 21). Valerio (1980: 268) removed it from sclerotized than Tliltocatl when it is present in the last
synonymy. (Figs 12E–F, 16E–G, 40D).
Note: Stridulation can be defined as the process of
Euathlus Ausserer, 1875: 188. Type species: sound production by friction of one rigid body part
E. truculentus L. Koch in Ausserer, 1875. First (the scraper) across a second part (the comb) (Uetz &
synonymized by Raven (1985: 182). Schmidt (1992: Stratton, 1982; Vol, 2002). It plays a role in different
9) removed it from synonymy. contexts, such as in intraspecific (reproduction) and
interspecific (defence) communication (Marshall
Brachypelmides Schmidt & Krause, 1994: 7. Type et al., 1995). It was Pocock (1903) who first mentioned
species: B. klaasi Schmidt & Krausse, 1994 (Schmidt, the plumose setae on the trochanter and femur I as a
1997a: 205, 1997b: 19; Locht et al., 1999: 196; Vol, key feature for Brachypelma recognition. Since then,
1999: 11; Schmidt, 2003: 137, 2004: 4). Smith (1994: presence of plumose setae has been used as a diagnostic
159) considered it as junior synonym of Brachypelma. character for Brachypelma (Smith, 1994; Pérez-Miles
et al., 1996; Gabriel & Longhorn, 2015; Mendoza &
Francke, 2017). This feature has also been recognized
Emended diagnosis as stridulatory setae by Galleti & Guadanucci (2018)
The genus Brachypelma can be distinguished from and identified by them as claviform stridulating
all other known theraphosinae genera (except setae, so we here also consider them as stridulating.
the new genus Tliltocatl described below) by the Most of the stridulatory organs in Theraphosinae
following combination of characters: (1) having just are positioned on the first appendages and consist
claviform stridulating setae on the prolateral face of of plumose or claviform hairs found between the
coxae and/or trochanters of the palps and first legs. never far from taller vegetation (Locht et al., 1999).
The function of stridulation is not well understood Unlike many other Theraphosinae species, the burrow
for theraphosid spiders. The different morphologies entrances have no tell-tale trace of silk, indicating
and positions of the stridulatory apparatus amongst inhabitation of a theraphosid or spider, so there never
the theraphosid subfamilies suggest that they are is an indication whether a spider will found while
not homologous. Presumably, these organs evolved digging the burrow until the end is excavated (West,
independently several times in theraphosids (Bertani 2005).
et al., 2008). In Brachypelma, claviform stridulating
setae are used as warning when tarantulas are
disturbed, but are also used during courtship to
Brachypelma albiceps Pocock, 1903
give some signals to the female (Mendoza, personal
observation). (Figs 2, 5–8, 38)
Species included: Brachypelma albiceps Pocock, 1903, Eurypelma pallidum F. O. Pickard-Cambridge, 1897:
Brachypelma auratum Schmidt, 1992, Brachypelma 21, plate 1, fig. 17 (female misidentified).
baumgarteni Smith, 1993, Brachypelma boehmei Brachypelma albiceps Pocock, 1903: 103 (D female).
Schmidt & Klaas, 1993, Brachypelma emilia (White, Schmidt (2004: 4, fig. 1) transferred to synonymized
1856), Brachypelma hamorii Tesmoingt, Cleton & generic name. Locht et al. (2005: 108) transferred from
Verdez, 1997, Brachypelma klaasi Schmidt & Krause, Aphonopelma. Estrada-Alvarez (2014: 57, fig. 18),
1994, Brachypelma smithi (F. O. Pickard-Cambridge, female. Teyssié (2015: 266–267), female.
1897). Aphonopelma albiceps (Pocock, 1903). Smith (1994:
70, figs 76–82) removed female from synonymy of
Distribution and natural history A. pallidum. Peters (2000: 23, fig. 43), female. Peters
(2003: 32, fig. 107), female.
The genus is endemic to Mexico and is distributed Brachypelmides ruhnaui Schmidt, 1997a: 205, fig. 1–4
from Sonora in the north to Guerrero in the south, (D male and female). Schmidt (1997b: 19, figs 199–
in the biogeographical region of the Pacific Coast, 201), male and female. Locht, Yáñez & Vázquez (1999:
with a couple of species (B. auratum and B. albiceps) 196, fig. 3), male. Peters (2000: 76, figs 250–252), male
distributed in parts of Central Mexico following the and female. Peters (2003: 133, figs 542, 545–547), male
biogeographical region of the Balsas Depression and female. Schmidt (2003: 137, fig. 207), male and
(Fig. 38). It is important to notice that the type female. Locht et al. (2005: 108) considered it as junior
of Brachypelma is B. emilia, which was originally synonym of B. albiceps.
described as a species from Panama (White, 1856).
However, as was mentioned by Gabriel & Longhorn
(2015) this locality could be an error, because the Material examined
original collector of the species, Berthold Seemann,
Holotype of Brachypelma albiceps: MEXICO: Guerrero:
joined a ship in Panama and travelled north along
♀, Venta de Zopilote. No more data (NHM labelled as
the Pacific coast, docking in Mexico at Nayarit and
BM1898.12.24.34-37).
Sinaloa, both within the area of distribution for
B. emilia. Holotype and paratype of Brachypelmides ruhnaui:
The genus Brachypelma has a natural distribution MEXICO: Estado de MEXICO: 1♂, 1♀, Toluca. Col.
in the tropical region of the Mexican Pacific Coast Mark Ruhnau (SMF39013).
and can be found mainly in tropical deciduous forest,
which is strongly influenced by dry and rainy seasons, Other material: MEXICO: Guerrero: 1 ♀, no more
although some populations can also be found in thorn data, source E. Hijmensen, S. Longhorn (RUHZ
scrub or oak forest (Smith, 1994; West, 2005; Mendoza, Longhorn DNA sample 2005); 1♂, 1♀, Mpio. Copalillo,
personal observation). All species are located between Papalutla, 14/X/2008, J. Mendoza (CNAN-Ar003412);
10 and 1600 m elevation, although rarely some 1 ♀, Mpio. Azoyu, El Carrizo VII/2011, A. Alcaraz
specimens of B. albiceps can be found as high as 1700 (CNAN-Ar007850); 2 ♀, Mpio. Tixtla de Guerrero,
m (Mendoza, personal observation). Brachypelma Zotoltitlan, 23/IX/2012, J. Mendoza, G. Contreras,
species collected during field work were found mainly J. Cruz, D. Ortiz (CNAN-Ar007839, CNAN-Ar007843);
in burrows on hillsides. Some burrows were made 1 juvenile, Mpio. Chilpancingo de los Bravo, Milpillas,
by the tarantula, but others were clearly adapted by 22/IX/2012, J. Mendoza, G. Contreras, J. Cruz, D. Ortiz
the spider from previous burrows. There are some (CNAN-Ar007852); 1 ♂, Mpio. Arcelia, Teloloapan,
specimens that made burrows under large, flat rocks 15/IX/1952, col. L. Vazquez (CNAN-Ar003099); 1 ♂,
or between tree roots, sometimes in open fields, but Mpio. Arcelia, Presa Vicente Guerrero, 05/X/1976,
Figure 5. Brachypelma albiceps, male CNAN-Ar003412. A, carapace, dorsal view; B, prosoma, ventral view; C, opisthosoma,
dorsal view; D, ocular tubercle, dorsal view; E, metatarsus I, prolateral view; F, tibial apophyses, prolateral view; G, tibial
apophyses, ventral view. Scale = 10 mm (A–C), 8 mm (E), 4 mm (F, G), 1 mm (D).
(CNAN-Ar003413); 1 ♂, Mpio. Arcelia, Presa Vicente Diagnosis

Guerrero, 23/XI/1976, A. Castillo (CNAN-Ar003437); Brachypelma albiceps can be distinguished from all
1 ♂, Mpio. Eduardo Neri, Ahuelican, 03/X/2008, other known Brachypelma species by its coloration,
O. Francke, A. Valdez, T. Lopez (CNAN-Ar004128); consisting of brown range setae on the carapace and
1 ♂, Mpio. Eduardo Neri, Venta Vieja, 26/V/1960, red setae on the opistosoma (Fig. 8A–B). The shape
(CNAN-Ar004129); 1 ♂, Mpio. Tepecoacuilco, of the genitalia also differs in both sexes: with the
Ahuehuepan, 30/VIII/2009, O. Francke, A. Valdez, palpal bulb being sharp and tapered (Fig. 6) and the
T. L o p e z , C. S a n t i b a ñ e z ( C N A N - A r 0 0 4 1 3 0 ) . spermatheca separated and with rounded receptacles
Morelos: 1 ♀, Mpio. Cuernavaca, 21/IX/1959, (Fig. 7E–G).
(CNAN-Ar003082); 1 ♂, Mpio. Tequesquitengo, 24/ Brachypelma albiceps can be identified by possesing
IX/1961, Santibañez (CNAN-Ar003113); 1 ♂, Mpio. the following character combination: male palpal
Amacuzac, Huajintlán, 28/VIII/2009, O. Francke, bulb with sharp embolus curving to dorsal through
A. Valdez, C. Santibañez, T. Lopez, (CNAN-Ar010574). its length (Fig. 6E–D), prolateral superior keel very
Figure 6. Brachypelma albiceps, male CNAN-Ar003412. Left palpal bulb: A, dorsal view; B, ventral view; C, retrolateral
view; D, prolateral view. Scale = 2 mm.
reduced, apical keel developed, wider behind embolus 8.62, 53.42; III: 14.13, 7.07, 10.32, 13.78, 8.85, 54.15;
tip (Fig. 6E–D). Embolus tip directed to retrolateral IV: 16.88, 7.68, 14.29, 18.88, 10.60, 68.33. Palp: 10.51,
(Fig. 6A–B). Embolus similar in length to tegulum 5.96, 9.53, -, 3.88, 29.88. Spinnerets: PMS, 1.90 long,
(Fig. 6). Spermatheca separated with semicircular 1.00 apart; PLS, 2.90 basal, 1.95 middle, 3.25 distal.
receptacles. Spermathecal baseplate divided, oval; Tarsi I–IV entirely scopulated. Metatarsus I densely
twice wider than its height (Fig. 7E–G). Carapace of scopulated, II densely scopulated, III scopulated 75%
both sexes golden yellow (Fig. 8A–B). distally, IV scopulated 50% distally. Tibia I with two
tibial apophyses normally developed which originate
from a common base. Prolateral apophysis with one
Redescription inner conical spine; retrolateral apophysis almost
Male (CNAN-Ar003412) (Figs 5–6): Body length the same width throughout its length and curved to
38.45 (not including chelicerae and spinnerets). dorsal on apex (Fig. 5F–G). Metatarsus I curved (Fig.
Carapace 17.64 length, 16.61 width. Caput not 5E). Stridulatory setae: with plumose setae on palp
markedly elevated; fovea straight, 3.25 wide (Fig. 5A). trochanter retrolateral face, and leg I trochanter and
Eyes: anterior eye row procurved, posterior eye row femur prolateral face. Chaetotaxy (left side): femora
recurved. Eye sizes and interocular distances: AME palp 1p; patellae none; tibiae II 2v; III 1r, 2v; IV 1p,
0.45; ALE 0.70; PME 0.35; PLE 0.53; AME–AME 0.38; 2v; palp 1p, 1v; metatarsi II 2v; III 2p, 6v; IV 2p, 9v,
AME–ALE 0.20; PME–PME 0.98; PME–PLE 0.10; 1r. Palp: embolus tapering and with very reduce
ALE–PLE 0.18. Ocular tubercle width 2.23, length spoon-like shape at tip. Embolus slightly twisted to
1.98; clypeus lacking (Fig. 5D). Labium length 2.43, retrolateral. Prolateral superior keel very reduced
width 2.77; with 95 cuspules. Maxilla inner corner and thin, prolateral inferior keel absent, apical keel
(left–right) with approximately 167–194 cuspules. extending to posterior and wider at the embolus tip.
Cheliceral promargin with (left–right) ten–ten teeth. The opening of the embolus is on the prolateral side,
Sternum length 7.90, width 6.43. Sigilla oval, second just behind the opening is located a concavity which
and third pairs hardly visible, posterior sigilla once delimits the apical keel boundary from the remaining
its length from the margin (Fig. 5B). Leg formula: IV, part of the embolus. Tegular apophysis rounded. (Fig.
I, III, II. Length of legs and palpal segments (femur, 6A–D). Urticating setae: types I and III arranged in
patella, tibia, metatarsus, tarsus, total): I: 16.68, 8.70, one dorsoposterior patch, black in colour. Type III
13.10, 12.76, 8.05, 59.29; II: 14.37, 7.54, 11.35, 11.54, are located in an oval dorsomedial area extended to
Figure 7. Brachypelma albiceps. A–E, female CNAN-Ar003082. A, carapace, dorsal view; B, prosoma, ventral view; C,
ocular tubercle, dorsal view; D, labial and maxillary cuspules; E, spermatheca, ventral view; F, G, spermatheca ventral view
of: F, female BMNH1898.12.24.34-37; G, female BMNH. Scale = 10 mm (A–B), 2 mm (D), 1 mm (C, E–G).
posterior. Type I surrounding the type III area, with Sigilla oval, third pair hardly visible; posterior sigilla
intermediates between type III and I in transition once its length from the margin (Fig. 7B). Leg formula:
areas (Fig. 5C). IV, I, II, III. Length of legs and palpal segments (femur,
patella, tibia, metatarsus, tarsus, total): I: 16.52, 10.27,
Female (CNAN-Ar003082) (Fig. 7A–E): Body length 13.05, 12.26, 8.36, 60.73; II: 15.35, 9.57, 10.94, 10.57,
53.61 (not including chelicerae and spinnerets), 7.94, 54.37; III: 14.40, 8.49, 9.30, 12.35, 6.92, 51.46;
carapace 23.50 length, 22.40 width. Caput not IV: 17.65, 9.00, 12.90, 17.39, 8.79, 65.91. Palp: 12.88,
markedly elevated; fovea procurved, 5.80 wide (Fig. 6.12, 9.16, -, 9.12, 37.28. Spinnerets: PMS, 3.00 long,
7A). Eyes: anterior eye row procurved, posterior eye 3.50 apart; PLS, 4.75 basal, 3.10 middle, 3.65 distal.
row recurved. Eye sizes and interocular distances: Tarsi I–IV entirely scopulated. Metatarsi I–II entirely
AME 0.47; ALE 0.50; PME 0.30; PLE 0.40; AME–AME scopulated, III scopulated 75% distally, IV scopulated
0.67; AME–ALE 0.33; PME–PME 1.40; PME–PLE 50% distally. Stridulatory setae: with claviform
0.15; ALE–PLE 0.43. Ocular tubercle width 2.80, stridulating setae on leg I trochanter and femur
length 2.43; clypeus lacking (Fig. 7C). Labium length prolateral face. Chaetotaxy (left side): femora palp 1p;
2.95, width 4.15; with 94 cuspules. Maxilla inner corner patellae none; tibiae I 2v, II 1p, 4v; III 3v, 1r; IV 3v, 1r;
(left–right) with approximately 129–130 cuspules palp 5v, 1r; metatarsi I 1v; II 3v; III 2p, 5v, 1r; IV 13v, 2r.
(Fig. 7D). Cheliceral promargin with (left–right) Genitalia: spermatheca separated with semicircular
nine–ten teeth (proximal to distal: first–third large, receptacles strongly sclerotized, ventral face smooth.
fourth–sixth medium, seventh–eighth large, ninth Spermathecal baseplate divided, oval; twice wider
small; first-third large, fourth small, fifth medium, than its height. (Fig. 7E). Urticating setae: types
sixth–tenth large). Sternum length 11.40, width 9.43. I and III arranged in one dorsoposterior patch, black
Figure 8. A–B, Brachypelma albiceps, habitus; C, habitat. A, male; B, female, C, deciduous forest in the habitat of B. albiceps.
Photos: J. Mendoza.
in colour. Type III are located in an oval dorsomedial ventrally brownish black. Legs and palpi: bluish black.
area extended to posterior. Type I surround the type (Fig. 8A–B).
III area; with intermediates between type III and I in
transition areas. Distribution and habitat
Colour pattern: In living specimens, adults of both Brachypelma albiceps is known from Central Mexico in
sexes have the carapace brown range (pantone 719c) Morelos, part of Puebla and Estado de Mexico to central
in colour (Fig. 8A–B), with juveniles or subadults Guerrero as the southern limit of its distribution. Its
paler in colour; chelicerae dorsally dark blue-grey; burrows occur in dense thickets or dry thorn forests
ventral coxae, labium, maxillae and sternum brownish under large rocks or tree roots (Figs 2, 8C, 38). There
black; abdomen dorsally black with light orange setae, are no traces of silk at the burrow entrance and the
interior can often be multitunnelled. The breeding Tesmoingt et al., (1997a: 9, plate 2, fig. 5), female. Schmidt
season occurs during the last part of the rainy and (1997b: 19, fig. 192), female. Locht et al. (1999: 196, fig. 6),
first part of dry seasons (August to January). Egg-sacs female. Peters (2000: 65, figs 210–211), male and female.
are constructed through the drier winter months with Peters (2003: 110, figs 439–440, 443, 446–447, male and
young emerging and dispersing in late spring, just female. Schmidt (2003: 137, fig. 203), female.
before the onset of the early summer rains.
Material examined
Holotype of Brachypelma auratum: MEXICO: ♂,
Brachypelma auratum schmidt, 1992 donated 22/II/1992 by Fritzlen (SMF 38045). Female
spermatheca from exuvia mounted in a permanent
(Figs 2, 9–13, 36A–D, 38) preparation (SMF 38045).
Brachypelma auratum Schmidt (1992b: 9, fig. 1, 3), D
male and female. Schmidt (1993: 82, fig. 187), female. Other material: MEXICO: Michoacán: 1 ♂, Mpio. Los
Smith (1994: 160, figs 851–866), male and female. Reyes, Los Reyes Salgado, 2/II/2013, Col. J. Mendoza,
Figure 9. Brachypelma auratum, male CNAN-Ar003658. A, carapace, dorsal view; B, prosoma, ventral view; C, opisthosoma,
dorsal view; D, ocular tubercle, dorsal view; E, metatarsus I, prolateral view; F, tibial apophyses, prolateral view; G, tibial
apophyses, ventral view. Scale = 12 mm (A–C), 8 mm (E), 4 mm (F, G), 1 mm (D).
Figure 10. Brachypelma auratum, male CNAN-Ar003658. Left palpal bulb: A, dorsal view; B, ventral view; C, retrolateral
Figure 11. Brachypelma auratum. A–C, holotype male SMF-38045. A, habitus, dorsal view; B, laminated spermatheca; C,
spermatheca, ventral view.
G. Contreras, D. Ortiz, D. Barrales (CNAN-Ar003658); CNAN-Ar010284); 1♀, Mpio. La Huacana, 1 km SE of

1 ♀, Mpio. San Lucas, Salguero 2/II/2013, J. Mendoza, Chau (CNAN-Ar010284); Guerrero: 1 ♀, Mpio. Arcelia,
G. Contreras, D. Ortiz, D. Barrales (CNAN-Ar007164); Desviación a Chacamerito, 3/II/2013, J. Mendoza,
2♀, Mpio. La Huacana, 5 km SE of Zicuirán, 16/ G. Contreras, J. Cruz, D. Ortiz (CNAN-Ar007136); 1 ♀,
I X / 2 0 1 5 , C o l . J. M e n d o z a ( C N A N - A r 0 0 7 9 0 3 , Mpio. Chilpancingo de los Bravo, carretera Coyuca
Figure 12. Brachypelma auratum. A–E, female CNAN-Ar007878. A, carapace, dorsal view; B, prosoma, ventral view; C,
ocular tubercle, dorsal view; D, labial and maxillary cuspules; E, spermatheca, ventral view; F, spermatheca ventral view of:
F, female SMF-58207. Scale = 10 mm (A–B), 2 mm (D), 1 mm (C, E, F).
de Catalán-Zihuatanejo, cols. A. Zaldívar, M. García, backward development, extending almost the same
J. Martínez, V. Salinas (CNAN-Ar007878); Estado de length of prolateral superior keel (Fig. 10B–C). The
MEXICO: 1 ♂, Mpio. Luvianos, 2/11/2002, E. Gonzalez- prolateral inferior keel is joined to prolateral superior
Santillan, R. Paredes, C. Durán (CNAN-Ar003592); keel at the apex and slightly extended to downwards
2 ♂ and 1♀, donation received from private collection (Fig. 10D). Also differs by the spermatheca ventral face
of J. Mendoza (CNAN-Ar003660, CNAN-Ar003676. smooth, with spermathecal baseplate subrectangular
CNAN-Ar003657). (Figs 11C, 12E–F).
Brachypelma auratum is identified by possesing
the following character combination: male palpal
Diagnosis bulb straight with a broad spoon-shape (Fig. 10C),
Brachypelma auratum can be distinguished from all prolateral superior keel normally developed, directed
other known Brachypelma species by the coloration of retrolaterally and slightly extended to backward (Fig.
the legs with red-orange flame-shape on patellae (Fig. 10A, C); prolateral inferior keel weakly developed
36A–D). It also differs in the shape of the genitalia in restricted to embolus apex, better seen dorsally (Fig.
both sexes with palpal bulb straight, embolus short 10A); apical keel developed extending widely backwards
and broader at apex (Fig. 10C–D). The prolateral (Fig. 10B). Embolus tip slightly directed to retrolateral
superior keel normally developed and slightly directed (Fig. 10A–B). Embolus short and wide, similar in length
retrolaterally (Fig. 10A). Apical keel with a great to tegulum (Fig. 10). Spermatheca fused semicircular,
Figure 13. A–C, Brachypelma auratum, habitus; D, habitat. A, male; B, female; C, juvenile; D, deciduous forest and
shrubland in the habitat of B. auratum, C. Photos: J. Mendoza.
with a single receptacle strongly sclerotized (Figs with 10 (left) and 10 (right) teeth (proximal to distal:
11C, 12E–F). Spermathecal baseplate divided, more first large, second small, third large, fourth–tenth
separated above (Fig. 12E–F); ovate and 2.5 wider than large). Sternum length 9.00, width 7.42. Sigilla oval,
its height (Figs 11C, 12E–F). Both sexes possess an first to third pairs hardly visible, posterior pair once
orange flame-shape area over the patellae (Fig. 13A–C). its length from the margin (Fig. 9B). Leg formula: IV,
I, II, III. Length of legs and palpal segments (femur,
patella, tibia, metatarsus, tarsus, total): I: 17.25, 9.45,
Redescription 14.08, 15.17, 10.32, 66.27; II: 15.92, 9.32, 12.61, 13.91,
Male (CNAN-Ar003658) (Figs 9–10): Body length 42.09 9.06, 60.82; III: 14.49, 8.06, 11.73, 14.15, 8.68, 57.11;
(not including chelicerae and spinnerets), carapace IV: 17.61, 8.40, 14.79, 19.44, 9.61, 69.85. Palp: 10.88,
18.16 length, 18.00 width. Caput not markedly 7.08, 10.35, -, 3.55, 31.86. Spinnerets: PMS, 2.10 long,
elevated; fovea straight, 4.15 wide (Fig. 9A). Eyes: 1.30 apart; PLS, 2.83 basal, 2.20 middle, 3.70 distal.
anterior eye row procurved, posterior eye row recurved. Tarsi I–IV entirely scopulated. Metatarsus I entirely
Eye sizes and interocular distances: AME 0.40; ALE scopulated, II scopulated 75%, III scopulated 50%
0.55; PME 0.40; PLE 0.65; AME–AME 0.50; AME–ALE distally, IV scopulated 25% distally. Tibia I with two
0.23; PME–PME 1.13; PME–PLE 0.13; ALE–PLE 0.37. tibial apophyses normally developed, which originate
Ocular tubercle width 2.60, length 2.13; clypeus absent from a common base. Prolateral apophysis with inner
(Fig. 9D). Labium length 2.07, width 2.27; with 113 spine half its length; retrolateral apophysis same
cuspules. Maxilla inner corner with approximately 270 width throughout its length, apex slightly curved to
(left) and 248 (right) cuspules. Cheliceral promargin prolateral (Fig. 9F–G). Metatarsus I curved (Fig. 9E).
Stridulatory setae: with claviform stridulating setae 13.73, 13.01, 8.64, 62.79; II: 15.65, 10.60, 11.60, 12.49,
on palp trochanter retrolateral face; leg I trochanter 7.59, 57.93; III: 14.59, 9.57, 10.83, 13.77, 8.52, 57.28;
and femur prolateral face. Chaetotaxy (left side): IV: 17.94, 10.05, 14.04, 18.78, 9.75, 70.56. Palp: 12.62,
femora I 1p; II 1p; palp 1p; patellae none; tibiae I 2p, 8.57, 8.93, -, 10.35, 40.47. Spinnerets: PMS, 2.50 long,
1r; II 2p, 3v; III 3p, 4v, 2r; IV 1p, 4v, 1r; palp 2p, 3v; 2.05 apart; PLS, 4.15 basal, 4.15 middle, 4.25 distal.
metatarsi II 1p, 1v; III 3p, 7v, 2r; IV 2p, 13v, 2r. Palp: Tarsi I–IV entirely scopulated. Metatarsi I entirely
embolus almost straight with a broad spoon-shape, scopulated, II scopulated 90%, III scopulated 70%
prolateral superior keel normally developed, directed distally, IV scopulated 40% distally. Stridulatory setae:
retrolaterally and slightly extended to backward; with claviform stridulating setae on palp trochanter
prolateral inferior keel weakly developed restricted retrolateral face; leg I trochanter and femur prolateral
to embolus apex, better seen dorsally, apical keel face. Chaetotaxy (left side): femora palp 1p; patellae
developed extending widely backwards. Opening none; tibiae palp 1p, 5v; II 1p, 3v; III 3v; IV 3v;
of the embolus is on the prolateral side, just behind metatarsi II 2v; III 5v, 2r; IV 1p, 8v, 1r. Genitalia: fused
the opening is located a concavity which delimits the semicircular spermatheca, with a single receptacle
apical keel boundary from the remaining part of the strongly sclerotized, four times wider than its height.
embolus. Embolus apex slightly curved to retrolateral Spermatheca baseplate divided, more separated above,
(Fig. 10A–D). Urticating setae: types I and III arranged each baseplate ovate 2.5 wider than high, outer side
in one dorsoposterior patch, black in colour. Type III slightly smaller than the inner (Fig. 12E). Variation:
are located in an oval dorsomedial area extended some specimens have a wider base up to five times
to posterior. Type I surround the type III area; with greater than height, young or juveniles can present the
intermediates between type III and I in transition superior edge slightly inward in middle. Ventral face
areas (Fig. 9C). Colour pattern: in live specimens, smooth. Baseplate division can vary in length (Figs
adult males with carapace peach colour (pantone 11C, 12F). Urticating setae: types I and III arranged
712c) around the border, some specimens also present in one dorsoposterior patch, black in colour. Type III
this colour behind the fovea. Carapace colour is black are located in an oval dorsomedial area extended
(pantone process black); chelicerae dorsally black; to posterior. Type I surround the type III area, with
ventral coxae, labium, maxillae and sternum brownish intermediates between type III and I in transition areas.
black; abdomen dorsally black with sparse rose colour Colour pattern: in live specimens, adult females with
setae (pantone 487c), ventrally brownish black. Legs two carapace patterns: (1) light brown (pantone 7414c)
and palpi: femora black, patellae with a proximal around the border and process black in dorsomedial,
dorsomedial signal orange (pantone 173c) flame-shape juveniles or subadults same pattern (Fig. 13B); (2) light
area, distal ring putty colour (pantone 728c), with some brown around the border and extended behind the
brownish pink setae laterally; tibiae proximal three- fovea, with process black from the fovea to caput (Fig.
quarters process black with some sparse brownish 13C); chelicerae dorsally oxford blue (pantone 532c);
pink setae, distal quarter brown range and metatarsi ventral coxae, labium, maxillae and sternum brownish
process black with brownish pink setae and a warm black; abdomen dorsally black with sparse rose colour
grey ring at the terminal end; tarsi black (Fig. 13A). setae (pantone 487c), ventrally brownish black. Legs
and palpi: femora black, patellae with a proximal
Female (CNAN-Ar007878) (Fig. 12A–E): Body length dorsomedial signal orange (pantone 173c) flame-shape
61.36 (not including chelicerae and spinnerets), area, distal ring putty colour (pantone 728c), with some
carapace 26.34 length, 24.61 width. Caput not brownish pink setae laterally; tibiae proximal three
markedly elevated; fovea straight, 5.70 wide (Fig. 12A). quarters process black with some sparse brownish
Eyes: anterior eye row procurved, posterior eye row pink setae, distal quarter brown range and metatarsi
recurved. Eye sizes and interocular distances: AME process black with brownish pink setae and a warm
0.47; ALE 0.80; PME 0.37; PLE 0.70; AME–AME 0.67; grey ring at the terminal end; tarsi black (Fig. 36A–D).
AME–ALE 0.33; PME–PME 1.57; PME–PLE 0.13;
ALE–PLE 0.43. Ocular tubercle width 2.97, length Distribution and habitat: Brachypelma auratum is
2.93; clypeus lacking (Fig. 12C). Labium length 3.05, known from part of the Neotransvolcanic belt, being
width 3.55; with 86 cuspules. Maxilla inner corner with found in south-western Estado de MEXICO, north-
approximately 250 (left)–276 (right) cuspules (Fig. western Guerrero, central Michoacán and the north-
12D). Cheliceral promargin with 10 (left)–11 (right) eastern border of Jalisco with Michoacán (Figs 2, 38)
teeth. Sternum length 11.39, width 9.76. Sigilla oval, where it occurs in thorn and deciduous secondary
first to third pairs hardly visible; posterior pair once forests (Fig. 13D). It is a fossorial species whose
its length from the margin (Fig. 12B). Leg formula: IV, modified or self-excavated burrows can be found under
I, II, III. Length of legs and palpal segments (femur, large rocks and large tree roots amongst thorny brush.
patella, tibia, metatarsus, tarsus, total): I: 16.56, 10.85, Burrows do not have any silk around the entrance.
Remarks Brachypelma baumgarteni Smith, 1993
The type specimen of B. auratum (SMF38045) lacks
(Figs 2, 14–17, 37A–D, 38)
palpal bulbs because they were removed, possibly
when it was described (Fig. 11A). It was not possible Brachypelma baumgarteni Smith, 1993: 15, figs 1–10
to find them in the holotype jar, nor in the laminated (D male); Smith (1994: 163, figs 875–883), male;
collection at the Senckenberg Museum. One mounted Teyssié (2015: 269, fig. 1), female. Misidentified by
spermatheca was found instead of a palpal bulb (Fig. Peters (2000: 66, figs 214–215), male and Peters
11B–C). The origin of that spermatheca is uncertain. (2003: 114, figs 457, 460), male, the species shown
However, the other features like patellae coloration in the figures is an adult male of Brachypelma
and tibial apophysis in the type specimen are sufficient hamorii Tesmoingt, Cleton & Verdez, 1997 not
for identification of this species. B. baumgarteni.
Figure 14. Brachypelma baumgarteni, male CNAN-Ar010588. A, carapace, dorsal view; B, prosoma, ventral view; C,
opisthosoma, dorsal view; D, ocular tubercle, dorsal view; E, metatarsus I, prolateral view; F, tibial apophyses, ventral view;
G, tibial apophyses, prolateral view. Scale = 12 mm (A–C), 8 mm (E), 4 mm (F, G), 1 mm (D).
Figure 15. Brachypelma baumgarteni, male CNAN-Ar010588. Left palpal bulb: A, dorsal view; B, ventral view; C,
retrolateral view; D, prolateral view. Scale = 2 mm.
Material examined keel (Fig. 15B). Also differs by the spermatheca ventral
Holotype of Brachypelma baumgarteni: MEXICO: face smooth, with spermathecal baseplate oblong (Fig.
Michoacán: ♂, Sierra Madre del Sur, M. Baumgarten 16E–G).
(BMNH-1999-122). Brachypelma baumgarteni is identified by possesing
the following character combination: male palpal bulb
Other material: MEXICO: Michoacán: 1 ♂, Carretera slightly curved to dorsal with a narrow spoon-shape at
La Mira-Arteaga, 09/IX/2012, without collector apex, prolateral superior keel short, thin and directed
(CNAN-Ar003597); 2 ♀, Mpio. Lázaro Cárdenas, Los retrolaterally; apical keel short and thin, larger than
Amates 11/XII/2013, J. Mendoza (CNAN-Ar007149) prolateral superior keel. Embolus tip slightly directed
(CNAN-Ar007150); 1 ♀, Mpio. Lázaro Cárdenas, to retrolateral. Embolus compact and thin, shorter
Puente Chuquiapan, 10/XII/2013 J. Mendoza in length than tegulum (Fig. 15A–D). Spermatheca
(CNAN-Ar007151); 1 ♂, Mpio. Lázaro Cárdenas, fused trapezoidal, with a single receptacle strongly
Los Coyotes, 8/XII/2016, J. Mendoza, R. Ramírez sclerotized. Spermathecal baseplate divided, more
(CNAN-Ar010588). separated below; oblong and three times wider than
its height (Fig. 16E–G). Both sexes possess a yellowish
longitudinal line just at metatarsi middle, this is better
Diagnosis seen on legs I and IV (Fig. 32A–D).
Brachypelma baumgarteni can be distinguished from
all other known Brachypelma species by the coloration
of the legs with deep orange flame-shape on patellae Description
with yellowish around, tibia and metatarsus with Male (CNAN-Ar010588) (Figs 14, 15, 17A): Body length
similar yellowish colour (Fig. 37A–D). It also differs in 49.63 (not including chelicerae and spinnerets), carapace
the shape of genitalia in both sexes with palpal bulb 22.59 length, 21.54 width. Caput not markedly elevated;
slightly curved to dorsal (Fig. 15C–D), embolus shorter fovea recurved, 3.75 wide (Fig. 14A). Eyes: anterior eye
than tegulum and broader at apex (Fig. 15A–B). The row procurved, posterior eye row recurved. Eye sizes and
prolateral superior keel broad and short, slightly interocular distances: AME 0.40; ALE 0.43; PME 0.33;
directed retrolaterally (Fig. 15A, C). Apical keel thin, PLE 0.70; AME–AME 0.40; AME–ALE 0.20; PME–PME
extending more than the length of prolateral superior 0.97; PME–PLE 0.10; ALE–PLE 0.23. Ocular tubercle
width 2.50, length 2.13; clypeus 0.17 (Fig 14D). Labium to prolateral (Fig. 14F–G). Metatarsus I curved (Fig.
length 2.57, width 2.90; with 80 cuspules. Maxilla inner 14E). Stridulatory setae: with plumose setae on palp
corner with approximately 179 (left) and 148 (right) coxa and trochanter retrolateral face; leg I trochanter
cuspules. Cheliceral promargin with ten (left) and ten and femur prolateral face. Chaetotaxy (left side): femora
(right) teeth (proximal to distal: first-third large, fourth I 1p; II 1p; palp 1p; patellae none; tibiae I 2p; II 3p, 4v;
medium, fifth large, sixth small, seventh–tenth large). III 3p, 5v, 1r; IV 4v, 1r; palp 2p, 1v; metatarsi I 3v; II 4v;
Sternum length 9.60, width 7.68. Sigilla oval, second and III 3p, 6v, 2r; IV 3p, 12v, 2r. Palp: embolus with narrow
third pairs hardly visible, posterior pair twice its length spoon-shape, slightly curved to dorsal. Embolus short
from the margin (Fig. 14B). Leg formula: IV, I, II, III. and flat at base. Prolateral superior keel short, thin
Length of legs and palpal segments (femur, patella, tibia, and directed retrolaterally. Apical keel short and thin,
metatarsus, tarsus, total): I: 19.29, 9.87, 14.70, 16.21, larger than prolateral superior. Both keels fusioned at
9.97, 70.04; II: 17.52, 9.79, 12.35, 13.39, 9.91, 62.96; III: apex and extended retrolatrally forming a well defined
16.15, 9.13, 11.83, 15.25, 9.01, 61.37; IV: 18.92, 9.99, 15.38, concave area on retrolateral. Opening of the embolus is
19.99, 11.10, 75.38. Palp: 11.95, 7.32, 11.26, -, 5.20, 35.73. on the prolateral side, just behind the opening is located
Spinnerets: PMS, 2.90 long, 1.45 apart; PLS, 4.10 basal, a concavity which delimits the apical keel boundary
2.95 middle, 4.15 distal. Tarsi I–IV entirely scopulated. from the remaining part of the embolus. Embolus apex
Metatarsus I entirely scopulated, II scopulated 75%, III slightly curved to retrolateral (Fig. 15A–D). Urticating
scopulated 60% distally, IV scopulated 30% distally. Tibia setae: types I and III arranged in one dorsoposterior
I with two tibial apophyses normally developed, which patch, black in colour. Type III are located in an oval
originate from a common base. Prolateral apophysis dorsomedial area extended to posterior. Type I surround
with inner spine half its length; retrolateral apophysis the type III area, with intermediates between type III
same width throughout its length, apex slightly curved and I in transition areas (Fig. 14C). Colour pattern: in
Figure 16. Brachypelma baumgarteni. A–E, female CNAN-Ar007150. A, carapace, dorsal view; B, prosoma, ventral view; C,
of: F, female CNAN-Ar007149; G, female CNAN-Ar007151. Scale = 10 mm (A–B), 3 mm (D), 1 mm (C, E–G).
Figure 17. A–C, Brachypelma baumgarteni, habitus; D, habitat. A, male; B, female dark carapace; C, female clear carapace;
D, deciduous forest in the habitat of B. baumgarteni. Photos: J. Mendoza.
live specimens, adult males with carapace rust colour Eye sizes and interocular distances: AME 0.53; ALE
(pantone 167c) with the border lightest. Chelicerae 0.60; PME 0.50; PLE 0.53; AME–AME 0.50; AME–ALE
dorsally Del rio colour (pantone 4735c); ventral coxae, 0.20; PME–PME 1.30; PME–PLE 0.13; ALE–PLE 0.27.
labium, maxillae and sternum brownish black; abdomen Ocular tubercle width 2.53, length 2.47; clypeus 0.47
dorsally black with sparse light terra di siena setae (Fig. 16C). Labium length 2.85, width 3.85; with 128
(pantone 472c), ventrally brownish black. Legs and palpi: cuspules. Maxilla inner corner with approximately
femora black, patellae with a proximal dorsomedial 211 (left)–225 (right) cuspules (Fig. 16D). Cheliceral
signal tomato red (pantone 485c) flame-shape area, promargin with eight (left)–nine (right) teeth (all big).
distodorsal paramedian tierra di siena setae with Sternum length 11.40, width 10.22. Sigilla oval, second
some long setae laterally of the same colour; tibiae raw and third pairs hardly visible; posterior pair once and
sienna (pantone 722c) with some sparse setae from half its length from the margin (Fig. 16B). Leg formula:
same colour and metatarsi corvette colour (pantone IV, I, II, III. Length of legs and palpal segments (femur,
720c) with mandys pink (pantone 473c) longitudinal patella, tibia, metatarsus, tarsus, total): I: 17.88, 11.25,
line just at metatarsi centre; tarsi black (Fig. 17A). 14.47, 13.15, 9.50, 66.25; II: 15.64, 10.49, 12.88, 12.58,
10.71, 62.30; III: 14.61, 10.10, 11.01, 15.58, 9.52, 60.82;
Female (CNAN-Ar007150) (Figs 16, 17B, 37A–D): Body IV: 18.60. 10.16, 14.66, 19.72, 10.71, 73.85. Palp: 13.08,
length 65.97 (not including chelicerae and spinnerets), 8.25, 10.37, -, 10.14, 41.84. Spinnerets: PMS, 2.50
carapace 25.67 length, 23.57 width. Caput not markedly long, 1.65 apart; PLS, 4.05 basal, 3.10 middle, 4.15
elevated; fovea recurved, 5.10 wide (Fig. 16A). Eyes: distal. Tarsi I–IV entirely scopulated. Metatarsi I and
anterior eye row procurved, posterior eye row recurved. II entirely scopulated, III scopulated 75% distally,
IV scopulated 50% distally. Stridulatory setae with Schmidt & Klaas (1994: 8, figs 1–2), male and female;
claviform stridulating setae on palp trochanter and Smith (1994: 164, figs 884–900), male and female;
femur retrolateral face; leg I coxa, trochanter and femur Tesmoingt et al., (1997a: 9, plate 2, fig. 1), female;
prolateral face. Chaetotaxy (left side): femora I 1p; II 1p; Schmidt (1997b: 19, fig. 190), male; Bertani (2000: 30,
palp 1p; patellae none; tibiae palp 1p, 6v; I 3v; II 4v; III figs 41–42), male; Peters (2000: 67, fig. 218), female;
1p, 4v, 1r; IV 1p, 3v, 1r; metatarsi I 2v; II 4v; III 2p, 5v, Peters (2003: 115–117, figs 461–469), male and female;
1r; IV 13v, 2r. Genitalia: fused trapezoidal spermatheca Schmidt (2003: 137, figs 201, 273), male and female.
with a single receptacle strongly sclerotized, ventral
face looks smooth. Three times wider than its height.
Spermatheca baseplate divided, oblong, almost as high Material examined
as one-third the width of its base. Baseplate division Holotype of Brachypelma boehmei: MEXICO: ♀, no
very narrow and poorly sclerotized in the basal half more data, K. Böhme (SMF 40590). Paratype. MEXICO:
(Fig. 16E). Variation: young or juveniles can present ♂, no more data, K. Böhme (SMF 38043).
the superior edge slightly inward in middle. Baseplate
division can vary in length, and some specimens Other material: MEXICO: Guerrero: 1 ♂, 2 ♀, Mpio.
can look more sclerotized (Fig. 16F, G). Urticating La Unión Isidoro Montes de Oca, 3km NE of La
setae: types I and III arranged in one dorsoposterior Unión, 12/XII/2013, J. Mendoza (CNAN-Ar007185,
patch, black in colour. Type III are located in an oval CNAN-Ar007186, CNAN-Ar007833); 2 ♂, Mpio.
dorsomedial area extended to posterior. Type I surround La Unión Isidoro Montes de Oca, 1km SW of La
the type III area, with intermediates between types Unión, 17/XII/2016, J. Mendoza (CNAN-Ar010589,
III and I in transition areas. Colour pattern: in live CNAN-Ar010591); 2 ♀, Mpio. La Unión Isidoro
specimens, adult females with two carapace patterns: Montes de Oca, Carretera Fed 37D, 16/IX/2015,
(1) provincial pink (pantone 4685c) around the border J. Mendoza (CNAN-Ar007905, CNAN-Ar010285);
and black (pantone 426c) in dorsomedial, juveniles Oaxaca: 1 ♂, Mpio. Salina Cruz, X/1963, E. Martín
or subadults same pattern (Fig. 17B); (2) Tuscany (CNAN-Ar003426) (collection data of this specimen are
(pantone 7522c) around the border and extended considered wrong due to lack of evidence of the species
behind the fovea, with black pearl (pantone black in Oaxaca and considering the restricted distribution
6c) from the fovea to caput (Fig. 17C); chelicerae of the species to a single municipality in Guerrero).
dorsally manatee (pantone 5285c) with two oriental
pink (pantone 7521c) cheliceral bands; ventral coxae,
Diagnosis
labium, maxillae and sternum black pearl; abdomen
dorsally black pearl (pantone black 6c) with rosy brown Brachypelma boehmei can be distinguished from all
(pantone 5005c) setae, ventrally black pearl. Legs and other known Brachypelma species by the coloration of
palpi: femora black pearl, patellae with a proximal the carapace and legs that are orange (Fig. 22A–C). It
dorsomedial cinnabar (pantone 7597c) flame-shape also differs in the shape of genitalia in both sexes with
area, distodorsal paramedian spanish white (pantone palpal bulb straight (Fig. 19C–D), embolus similar in
4685c) setae with some long setae laterally of the same length as tegulum and broader than embolus base at
colour; tibiae eunry (pantone 4735c) with some sparse apex (Fig. 19A–B). The prolateral superior keel normally
setae from same colour and metatarsi hemp (pantone developed and slightly directed retrolaterally (Fig.
8021c) with mandys pink (pantone 473c) longitudinal 19A). Apical keel with a great backward development,
line just at metatarsi centre; tarsi black (Fig. 17A–D). extending more than the length of prolateral superior
keel (Fig. 19B). The prolateral inferior keel is joined to
Distribution and habitat: Brachypelma baumgarteni is prolateral superior keel at the apex and slightly extended
known only from Michoacán on the Sierra Madre del to backwards (Fig. 19A). The prolateral apophysis is
Sur Region (Figs 2, 38), where it occurs in deciduous almost half the length of the retrolateral apophysis
forests (Fig. 17D). It is a fossorial species whose (Fig. 18F–G). It also differs in the spermatheca ventral
modified or self-excavated burrows, can be found face striated (Fig. 20B), with spermathecal baseplate
between large tree roots amongst large trees. Burrows lanceolate, notoriously separated and lower than the
do not have any silk around the entrance. height of the spermatheca (Fig. 21E–G).
Brachypelma boehmei is identified by possesing

Brachypelma boehmei Schmidt & Klaas, 1993 the following character combination: male palpal
bulb straight with a broad spoon-shape at apex,
(Figs 2, 18–22, 37E–H, 38)
prolateral superior keel normally developed and
Brachypelma boehmei Schmidt & Klaas, 1993: 7, figs 1–2 slightly directed retrolaterally; prolateral inferior keel
(D male and female); Schmidt (1993: 82, fig. 193a), male; weakly developed restricted to embolus apex; apical
Figure 18. Brachypelma boehmei, male CNAN-Ar010589. A, carapace, dorsal view; B, prosoma, ventral view; C, opisthosoma,
dorsal view; D, ocular tubercle, dorsal view; E, metatarsus I, prolateral view; F, tibial apophyses, ventral view; G, tibial
apophyses, prolateral view. Scale = 12 mm (A–C), 8 mm (E), 4 mm (F, G), 1 mm (D).
keel developed extending to backwards more than carapace 19.71 length, 18.82 width. Caput not
the length of the prolateral superior keel. Embolus markedly elevated; fovea recurved, 4.50 wide (Fig.
tip slightly directed to retrolateral. Embolus large 18A). Eyes: anterior eye row procurved, posterior eye
and broad, similar in length to tegulum (Fig. 19A–D). row recurved. Eye sizes and interocular distances:
Spermatheca fused trapezoidal, with a single receptacle AME 0.46; ALE 0.53; PME 0.46; PLE 0.56; AME–AME
strongly sclerotized. Spermathecal baseplate divided, 0.43; AME–ALE 0.20; PME–PME 1.20; PME–PLE
noticeably separated; lanceolate and three times wider 0.13; ALE–PLE 0.43. Ocular tubercle width 2.47,
than its height (Figs 20B, 21E–G). Carapace and legs length 2.23; clypeus absent (Fig. 18D). Labium length
of both sexes are orange in general (Fig. 22A–C). 2.40, width 3.25; with 85 cuspules. Maxilla inner
corner with approximately 248 (left) and 187 (right)
cuspules. Cheliceral promargin with ten (left) and ten
Redescription (right) teeth (proximal to distal: first–second large,
Male (CNAN-Ar010589) (Figs 18, 19, 22A): Body third medium, fourth large, fifth small, sixth–tenth
length 41.39 (not including chelicerae and spinnerets), large). Sternum length 8.70, width 8.09. Sigilla oval,
Figure 19. Brachypelma boehmei, male CNAN-Ar010589. Left palpal bulb: A, dorsal view; B, ventral view; C, retrolateral
Figure 20. Brachypelma boehmei. A–B, holotype female SMF-40590. A, exuvia used as holotypus; B, spermatheca, ventral
view (was not removed from the exuvia).
first to third pairs hardly visible, posterior pair once scopulated, II scopulated 80%, III scopulated 60%
its length from the margin (Fig. 18B). Leg formula: IV, distally, IV scopulated 40% distally. Tibia I with two
I, II, III. Length of legs and palpal segments (femur, tibial apophyses normally developed, which originate
patella, tibia, metatarsus, tarsus, total): I: 16.94, 9.01, from a common base. Prolateral apophysis with inner
13.20, 13.72, 8.47, 61.34; II: 15.56, 8.79, 11.94, 12.60, spine half its length; retrolateral apophysis same
8.48, 57.37; III: 14.40, 8.00, 11.29, 13.86, 8.43, 55.98; width throughout its length, apex slightly curved to
IV: 17.26, 8.51, 13.06, 18.04, 9.26, 66.13. Palp: 11.02, prolateral (Fig. 18F–G). Metatarsus I curved (Fig.
6.54, 9.51, -, 4.51, 31.58. Spinnerets: PMS, 2.15 long, 18E). Stridulatory setae: with claviform stridulating
0.95 apart; PLS, 3.80 basal, 2.55 middle, 3.25 distal. setae on palp trochanter and femur retrolateral face;
Tarsi I–IV entirely scopulated. Metatarsus I entirely leg I coxa, trochanter and femur prolateral face.
Figure 21. Brachypelma boehmei. A–E, female CNAN-Ar007905. A, carapace, dorsal view; B, prosoma, ventral view; C,
of: F, female CNAN-Ar007185; G, female CNAN-Ar007833. Scale = 10 mm (A–B), 3 mm (D), 1 mm (C, E–G).
Chaetotaxy (left side): femora I 1p; II 1p; palp 1p; Colour pattern: in live specimens, adult males with
patellae none; tibiae II 3p, 3v, 1r; III 3p, 4v, 2r; IV carapace colour persimmon (pantone 166c) and tacao
1p, 4v, 1r; palp 2p, 2v; metatarsi I 2v, II 1p, 4v; III (pantone 721c) around the border (Fig 22A); chelicerae
2p, 7v, 2r; IV 2p, 15v, 1r. Palp: embolus straight with dorsally tussock (pantone 729c); ventral coxae, labium,
a broad spoon-shape at apex and similar in length as maxillae and sternum seal brown (pantone 440c);
tegulum. Prolateral superior keel normally developed abdomen dorsally coffee bean (pantone black 2c) with
and slightly directed retrolaterally; prolateral inferior sparse light raw sienna (pantone 722c) setae, ventrally
keel weakly developed restricted to embolus apex, seal brown. Legs and palpi: femora coffee black,
better seen dorsally, apical keel developed extending patellae with a proximal dorsomedial persimmon
to backwards more than the length of the prolateral flame-shape area, surrounding this and distally is
superior keel. Opening of the embolus is on the raw sienna, with some setae laterally of same colour;
prolateral side, just behind the opening is located tibiae and metatarsi persimmon, with some lateral
a concavity which delimits the apical keel boundary raw sienna setae; tarsi proximally backer’s chocolate
from the remaining part of the embolus. Embolus apex (pantone 732c) and distally coffee bean (Fig. 37E–H).
slightly curved to retrolateral (Fig. 19A–D). Urticating
setae: types I and III arranged in one dorsoposterior Female (CNAN-Ar007905) (Figs 21A–E, 22B): Body
patch, black in colour. Type III are located in an length 61.36 (not including chelicerae and spinnerets),
oval dorsomedial area extended to posterior. Type carapace 26.34 length, 24.61 width. Caput not
I surround the type III area, with intermediates markedly elevated; fovea straight, 5.70 wide (Fig. 21A).
between type III and I in transition areas (Fig. 18C). Eyes: anterior eye row procurved, posterior eye row
Figure 22. A–C, Brachypelma boehmei, habitus; D, habitat. A, male; B, female with black around ocular area; C, femalewith
black lateral areas on ocular regions; D, deciduous forest and shrubland in the habitat of B. boehmei. Photos: A, B, D,
J. Mendoza; C, E. Goyer.
recurved. Eye sizes and interocular distances: AME scopulated, II scopulated 90%, III scopulated 70%
0.47; ALE 0.80; PME 0.37; PLE 0.70; AME–AME 0.67; distally, IV scopulated 40% distally. Stridulatory setae:
AME–ALE 0.33; PME–PME 1.57; PME–PLE 0.13; with claviform stridulating setae on palp trochanter
ALE–PLE 0.43. Ocular tubercle width 2.97, length retrolateral face; leg I trochanter and femur prolateral
2.93; clypeus lacking (Fig. 21C). Labium length 3.05, face. Chaetotaxy (left side): femora I 1p, II 1p; patellae
width 3.55; with 86 cuspules. Maxilla inner corner with none; tibiae I 1p, 3v; II 3p, 4v; III 3p, 6v, 2r; IV 1p, 4v,
approximately 250 (left)–276 (right) cuspules (Fig. 1r; palp 2p, 6v; metatarsi I 3v; II 1p, 3v; III 2p, 8v, 1r; IV
21D). Cheliceral promargin with ten (left)–11 (right) 2p, 15v, 1r. Genitalia: fused trapezoidal spermatheca,
teeth. Sternum length 11.39, width 10.09. Sigilla oval, with a single receptacle strongly sclerotized, three
first to third pairs hardly visible; posterior pair once times wider than its height. Spermatheca baseplate
its length from the margin (Fig. 21B). Leg formula: IV, divided, widely separated in middle, each baseplate
I, II, III. Length of legs and palpal segments (femur, lanceolate three times wider than high, outer side
patella, tibia, metatarsus, tarsus, total): I: 16.56, slightly smaller than the inner (Fig. 21E). Variation:
10.85, 13.73, 13.01, 8.64, 62.79; II: 15.65, 10.60, 11.60, some specimens have a wider base up to four times
12.49, 7.59, 57.93; III: 14.59, 9.57, 10.83, 13.77, 8.52, wider than height, young or juveniles can present the
57.28; IV: 17.94, 10.05, 14.04, 18.78, 9.75, 70.56. Palp: superior edge slightly inward in middle. Ventral with
12.48, 7.51, 9.34, -, 9.32. Spinnerets: PMS, 2.50 long, variation on striation. Baseplate division can vary
2.05 apart; PLS, 4.15 basal, 4.15 middle, 2.55 distal. in length (Figs 20B, 21F–G). Urticating setae: types
Tarsi I–IV entirely scopulated. Metatarsi I entirely I and III arranged in one dorsoposterior patch, black
in colour. Type III are located in an oval dorsomedial Material examined
area extended to posterior. Type I surround the type Neotype of Brachypelma emilia: MEXICO: Durango:
III area, with intermediates between type III and I in ♂, Ciudad, leg Mr. Forrer (BMNH-1898-12-24-32). 1 ♂
transition areas. Colour pattern: in live specimens, (labelled as paratype), Ciudad, leg. Forrer (OUNMH
adult females with two carapace patterns: (1) tango Jar 106).
(pantone 7412c) in all the carapace with pink flare
(pantone 5025c) around the border, juveniles or Note: Both specimens mentioned as neotype and
subadults same pattern (Fig 22B); (2) pink flare paratype were designated by Smith (1994: 166).
around the border and sandy brown (pantone 7411c) Other material: MEXICO: Nayarit: 1 ♂ 4 ♀, Mpio.
in dorsomedial, with two seal brown (pantone 440c) Compostela, 9.xii.2012, E. Goyer, E. Hijmensen,
longitudinal patches from the fovea to caput (Fig. D. Ortiz (CNAN-Ar003599, CNAN-Ar007153,
22C); chelicerae dorsally beaver colour (pantone CNAN-Ar007173, CNAN-Ar007178, CNAN-Ar007875);
4715c); ventral coxae, labium, maxillae and sternum 1 ♂, Mpio. Estación Ruíz, 1.xii.1989, A. Cadena
seal brown; abdomen dorsally black (pantone 426c) (CNAN-Ar003436); 4 ♂, Mpio. Estación Ruíz,
with putty colour (pantone 721c) setae, ventrally 5.xii.2014, J. Mendoza, G. Contreras (CNAN-Ar007146,
black. Legs and palpi: femora coffee black, patellae CNAN-Ar007894, CNAN-Ar007895, CNAN-Ar007899);
with a proximal dorsomedial cinnabar (pantone 173c) Sinaloa: 1 ♂, Mpio. Mazatlán, vii.1959, without more
flame-shape area, surrounding this and distally is data (CNAN-Ar003427); 1 ♂ 1 ♀, Mpio. Mazatlán,
dark salmon (pantone 472c), with some setae laterally 3.xii.2014, J. Mendoza, G. Contreras (CNAN-Ar007898,
of wafer colour (pantone 4745c); tibiae and metatarsi CNAN-Ar010602); 1 ♂, Mpio. Mazatlán, without
chardonnay (pantone 1355c), with some lateral wafer more data, Collection E. Simon (MP Ar4871A); 1 ♂,
setae; tarsi proximally Backer’s chocolate (pantone Mpio. Sinaloa de Leyva, 30.i.1965, without more data
732c) and distally coffee bean (Fig. 37E–H). (CNAN-Ar003590); Sonora: 1 ♂, Mpio. Altar, 9.i.1970,
W. Lopez Forment (CNAN-Ar003578); Jalisco: 1 ♀,
Distribution and habitat: Brachypelma boehmei is the Norte del Río Santiago, Godman, Salvin, without more
species with the most restricted distribution being only data (BMNH-1962-2-28-1); 1 ♂, donation received from
known from the Sierra de Cumbres region, an area private collection of J. Mendoza (CNAN-Ar003631).
characterized by low complex and rolling hills with
plains, belonging to the subprovince Costas del Sur in
south-western Guerrero (Figs 2, 38). It occurs in thorn Diagnosis
and deciduous forests (Fig. 22D). It is a fossorial species Brachypelma emilia can be distinguished from all
whose burrows can be found under large angular rocks other known Brachypelma species by the coloration of
and large tree roots amongst thorny brush. Burrows the carapace and legs, with carapace orange except in
do not have any silk around the entrance. the caput, which is black in colour (Fig. 26A–C). The
legs have black femora and patellae, orange tibiae and
Brachypelma emilia (White, 1856) metatarsi I–III are black and IV is orange (Fig. 36E–
H). It also differs in the shape of genitalia in both sexes
(Figs 2, 23–26, 36E–H, 38) with palpal bulb slightly curved to dorsal having a small
Mygale emilia White, 1856: 185, pl. 43 (D male). and narrow spoon-shape (Fig. 24C–D). The prolateral
Brachypelma emilia (Simon 1891: 338), D male superior keel is slightly developed, thin and directed
and female; Smith (1986: 49, fig. 27h), Tmf from retrolaterally (Fig. 24A, C). The prolateral inferior keel
Eurypelma=Avicularia; Smith (1987: 49, plate 2, is absent, the apical keel normally developed, wide
fig. 27h), male; Hancock & Hancock (1989: 46, fig. 41), but not shorter than prolateral superior keel (Fig.
female; Schmidt (1992: 10), Tmf from Euathlus per 24B). It also differs by the spermatheca ventral face
Raven; Schmidt (1993: 82, fig. 188), female; Smith smooth, with a single receptacle strongly sclerotized
(1994: 166, figs 901–915), male and female; Pérez-Miles slightly notched in the middle, spermathecal baseplate
et al. (1996: 46, figs 9–10), male and female; Tesmoingt oblanceolate (Fig. 25E, F).
et al. (1997a: 9, plate 2, fig. 6), female; Schmidt (1997:
19, figs 191, 193), male and female; Locht et al. (1999: Brachypelma emilia is identified by possesing the
196, fig. 7), female; Peters (2000: 68, fig. 222), female; following character combination: male palpal bulb with
Bertani (2001: 338, figs 153–156), male and female; narrow spoon-shape embolus curving slightly to dorsal
Peters (2003: 117, figs 473–474, 477, 480, 483), male through its length, prolateral superior keel slightly
and female; Schmidt I2003: 152, figs 274–277), male developed; apical keel normally developed, wide but not
and female; Gabriel & Longhorn (2015: 100, fig. 13), shorter than the prolateral superior keel tip. Embolus
female. tip directed to retrolateral. Embolus similar in length
Figure 23. Brachypelma emilia, male CNAN-Ar003599. A, carapace, dorsal view; B, prosoma, ventral view; C, opisthosoma,
than tegulum (Fig. 24A–D). Spermatheca fused with 0.70; AME–ALE 0.20; PME–PME 1.23; PME–PLE
single semitrapezoidal receptacle. Spermathecal 0.13; ALE–PLE 0.37. Ocular tubercle width 2.47,
baseplate divided, oblanceolate; one and half wider length 2.27; clypeus 0.23 (Fig. 23D). Labium length
than its height (Fig. 25E, F). Carapace of both sexes 2.60, width 3.40; with 109 cuspules. Maxilla inner
carapace orange with black triangle in the ocular area corner with approximately 205 (left) and 179 (right)
(Fig. 26A–C). cuspules. Cheliceral promargin with ten (left) and
nine (right) teeth (proximal to distal: first–third large,
fourth small, fifth–seventh medium, eighth–tenth
Redescription large; first–third large, fourth small, fifth–seventh
Male (CNAN-Ar003599) (Figs 23, 24, 26A): Body medium, eighth–ninth large). Sternum length 10.10,
length 40.16 (not including chelicerae and spinnerets), width 9.38. Sigilla oval, fourth pair hardly visible,
carapace 20.23 length, 18.85 width. Caput not posterior pair one and half its length from the margin
markedly elevated; fovea procurved, 3.80 wide (Fig. (Fig. 23B). Leg formula: IV, I, II, III. Length of legs and
23A). Eyes: anterior eye row procurved, posterior eye palpal segments (femur, patella, tibia, metatarsus,
row recurved. Eye sizes and interocular distances: tarsus, total): I: 17.25, 9.39, 11.67, 12.22, 9.28, 59.81;
AME 0.43; ALE 0.60; PME 0.33; PLE 0.37; AME–AME II: 15.93, 9.20, 10.96, 11.89, 8.21, 56.19; III: 14.35,
Figure 24. Brachypelma emilia, male CNAN-Ar003599. Left palpal bulb: A, dorsal view; B, ventral view; C, retrolateral
8.11, 10.00, 12.08, 8.15, 52.69; IV: 16.86, 8.63, 12.81, I and III arranged in one dorsoposterior patch, black
16.13, 9.92, 64.35. Palp: 11.83, 7.23, 9.61, -, 4.14, 32.81. in colour. Type III are located in an oval dorsomedial
Spinnerets: PMS, 2.20 long, 1.00 apart; PLS, 3.10 basal, area extended to posterior. Type I surround the type
2.20 middle, 3.20 distal. Tarsi I–IV entirely scopulated. III area, with intermediates between type III and I in
Metatarsus I entirely scopulated, II scopulated 75%, transition areas (Fig. 23C).
III scopulated 50% distally, IV scopulated 30%
distally. Tibia I with two tibial apophyses normally Female (CNAN-Ar010602) (Fig. 25A–E): Body length
developed, which originate from a common base. 61.31 (not including chelicerae and spinnerets),
Prolateral apophysis with inner spine third its length; carapace 26.34 length, 24.33 width. Caput not
retrolateral apophysis wider in basal half, apex almost markedly elevated; fovea straight, 6.40 wide (Fig. 25A).
straight (Fig. 23F, G). Metatarsus I curved (Fig. 23E). Eyes: anterior eye row procurved, posterior eye row
Stridulatory setae: with claviform stridulating setae recurved. Eye sizes and interocular distances: AME
on palp trochanter retrolateral face; leg I trochanter 0.65; ALE 0.70; PME 0.50; PLE 0.70; AME–AME 0.63;
and femur prolateral face. Chaetotaxy (left side): AME–ALE 0.33; PME–PME 1.57; PME–PLE 0.07;
femora I 1p; II 1p; palp 1p; patellae none; tibiae I 1p, ALE–PLE 0.40. Ocular tubercle width 3.25, length
2v; II 2p, 3v; III 2p, 5v, 1r; IV 1p, 4v, 1p; palp 2p, 1v; 2.83; clypeus 0.27 (Fig. 25D). Labium length 3.05,
metatarsi I 1v; II 2v; III 3p, 7v, 1r; IV 1p, 18v, 1r. Palp: width 3.85; with 94 cuspules. Maxilla inner corner
embolus slightly curved to dorsal having a small and with approximately 187 (left)–208 (right) cuspules.
narrow spoon-shape, prolateral superior keel slightly Cheliceral promargin with nine (left)–nine (right)
developed, thin and directed retrolaterally; prolateral teeth. Sternum length 11.60, width 10.90. Sigilla oval,
inferior keel absent; apical keel normally developed, fourth pair hardly visible; posterior pair twice its
wide but not shorter than prolateral superior keel. length from the margin (Fig. 25B). Leg formula: IV,
Opening of the embolus is on the prolateral side, I, II, III. Length of legs and palpal segments (femur,
just behind the opening is located a concavity which patella, tibia, metatarsus, tarsus, total): I: 17.87, 10.79,
delimits the apical keel boundary from the remaining 13.69, 13.40, 8.75, 64.50; II: 16.35, 10.18, 12.05, 11.77,
part of the embolus. Embolus apex slightly curved 8.94, 59.29; III: 15.69, 9.40, 10.97, 12.79, 8.50, 57.35;
to retrolateral (Fig. 24A–D). Urticating setae: types IV: 18.34, 10.20, 14.20, 17.91, 9.98, 70.63. Palp: 13.20,
Figure 25. Brachypelma emilia. A–E, female CNAN-Ar010602. A, carapace, dorsal view; B, prosoma, ventral view; C,
opisthosoma, dorsal view; D, ocular tubercle, dorsal view; E, spermatheca, ventral view; F, spermatheca ventral view of: F,
female BMNH1962.2.28.1. Scale = 10 mm (A–C), 1 mm (D–F).
8.35, 10.12, -, 9.92, 41.59. Spinnerets: PMS, 2.90 long, area extended to posterior. Type I surround the type
3.00 apart; PLS, 4.30 basal, 2.90 middle, 4.35 distal. III area, with intermediates between type III and I in
Tarsi I–IV entirely scopulated. Metatarsus I entirely transition areas (Fig. 25C).
scopulated, II scopulated 90%, III scopulated 50%
distally, IV scopulated 40% distally. Stridulatory setae: Colour pattern: In live specimens, adults of both
with claviform stridulating setae on palp trochanter sexes have the carapace orange (pantone 7412c) on
and femur retrolateral face; leg I trochanter and femur almost all the carapace except in the caput, which
prolateral face. Chaetotaxy (left side): femora I 1p; is pantone process black in colour, and also has a
palp 1p; patellae none; tibiae I 1p, 3v; II 1p, 4v; III 1p, longitudinal line of beaver colour (pantone 4715c)
5v, 1r; IV 1p, 6v, 1r; palp 2p, 5v; metatarsi I 4v; II 4v; III that goes from back of the eyes to the fovea (Figs.
1p, 8v, 2r; IV 2p, 14v, 1r. Genitalia: fused semicircular 26A–C); chelicerae dorsally blue whale colour
spermatheca, with a single receptacle strongly (pantone 533c); ventral coxae, labium, maxillae
sclerotized slightly notched in the middle, four times and sternum black pearl colour (pantone black
wider than its height. Spermatheca baseplate divided, 6c); abdomen dorsally black with Christine colour
widely separated above, each baseplate oblanceolate setae (pantone 7583c), ventrally black pearl colour.
3.5 wider than high, inner side smaller than the outer Legs and palpi: femora and patellae black pearl;
(Fig. 25E). Variation: some specimens have a wider tibiae orange (pantone 157c); metatarsi I–III black
base up to five times greater than height, young or (pantone black 7c) with proximal third rope colour
juveniles can present the superior edge slightly inward (pantone 876c), IV black (pantone black 7c) with
in middle. Ventral face smooth. Baseplate division large brandy punch colour (pantone 722c) covering it
can vary in length (Fig. 25F). Urticating setae: types almost completely; tarsi black (pantone black c) (Fig.
I and III arranged in one dorsoposterior patch, black 36E–H). With juveniles of same pattern but paler in
in colour. Type III are located in an oval dorsomedial colour.
Figure 26. A–C, Brachypelma emilia, habitus; D, habitat. A, male; B, juvenile female; C, adult female; D, deciduous forest
in the habitat of B. emilia. Photos: J. Mendoza.
Distribution and habitat: Brachypelma emilia is a small population of Brachypelma klaasi in south-
known from the northern Pacific Coast region on the western Nayarit (Mendoza, personal observation).
western side of Sierra Madre Occidental, being found
in southern Sonora, Sinaloa, Nayarit and a small
area in north-western Jalisco, just on the border of Brachypelma hamorii Tesmoingt et al, 1997
Jalisco with Nayarit. It could also be possibly found in
Durango, but there is no evidence for that yet (Figs 2, (Figs 2, 27–28, 35B, D, 37I–L, 38)
38). It occurs in drier, coastal thorn scrub, grasslands, Brachypelma hamorii Tesmoingt et al. (1997a: 9,
palm transition to deciduous forest, and into higher plates 1–6), D male and female; Tesmoingt et al. (1997b:
elevations of oak forest (Fig. 26D). It is a fossorial 3, plates 9–11), male; Mendoza & Francke (2017: 167,
species whose modified or self-excavated burrows figs 24–40, 46–49, 52, 57–60), male and female.
can be found under large rocks, under dense thorny E u a t h l u s s m i t h i ( F. O. P i ck a r d - C a m b r i d g e ) :
thickets, large tree roots or burrows in leafy ground Baxter (1993: 73, figs 17–18, plate A, figs 1–6),
cover, in both forested and moderately disturbed misidentification.
areas. Some can be also found close to houses or Brachypelma smithi (F. O. Pickard-Cambridge):
human structures, but this is most likely because Hancock & Hancock (1989: 44, fig. 39), misidentification
the spiders lived there before these buildings were female; Schmidt (1992a: 10), transferred from Euathlus;
constructed. Burrows do not have any silk around Schmidt (1992b: 14, figs 2, 4), male and female; Schmidt
the entrance to indicate there is a spider inside. This (1993: 82, fig. 190), female; Smith (1994: 170, figs 940–
species is sympatric (overlapping distributions) with 956), male and female; Tesmoingt et al. (1997a: 9, plate 2,
Figure 27. A–F, Brachypelma hamorii, neotype male CNAN-Ar007828. Left palpal bulb: A, dorsal view; B, ventral view; C,
retrolateral view; D, prolateral view. E, F spermatheca ventral view of: E, female CNANAr010279; F, female CNAN-Ar010280.
Scale = 2 mm (A–F).
fig. 2), female; Tesmoingt et al. (1997b: 4, plates 10–11), Other material: MEXICO: Colima: 1 ♂, Mpio.
male; Schmidt (1997: 19, fig. 195), female; Peters (2000: Tecoman, 1/XII/2012, E. Goyer, E. Hijmensen, D. Ortiz
72, figs 235–236), male and female; Peters (2003: 125, (CNAN-Ar003614); 1 ♀, Mpio. Colima, 30/XI/2012,
figs 510, 512–513), male and female; Schmidt (2003: E. Goyer, E. Hijmensen, D. Ortiz (CNAN-Ar004779);
153, figs 283–284), male and female. Misidentifications. 2 ♂, Mpio. Tecoman, 5/XII/2013, D. Ortíz, D. Barrales,
G. Contreras (CNAN-Ar007163, CNAN-Ar007826);
1 ♂, Mpio. Tecoman, 3/XII/2013, D. Ortíz, D. Barrales,
Material examined
G. Contreras (CNAN-Ar007168); 1 ♂, Mpio. Manzanillo,
Neotype of Brachypelma hamorii: MEXICO: Colima: ♂, IV/2004, IBT (CNAN-Ar010278); 1 ♂, Mpio. Manzanillo,
Mpio. Tecoman, 5/XII/2013, D. Barrales, G. Contreras, 4/XII/2013, D. Ortíz, D. Barrales, G. Contreras
D. Ortiz (CNAN-T0900). Designated as neotype by (CNAN-Ar010277); 1 ♂, donation received from private
Mendoza & Francke (2017: 167). collection of J. Mendoza (CNAN-Ar003616); 1 ♂,
Figure 28. A–F, Brachypelma hamorii, habitus. A, neotype male CNAN-Ar007828, in life; B, female CNANAr007874, in
life; C, male, in habitat (Michoacán); D, female, in habitat (Michoacán); E, female, in habitat (Colima); F, female, in habitat
(Colima). Photos: (A, B, C, F) J. Mendoza; (E) E. Goyer, (D) G. Vila.
Mpio. 3/XII/2013, D. Ortíz, D. Barrales, G. Contreras elliptic; five times wider than its height (Fig. 27E–F).
(CNAN-Ar007827); 1 ♂, Mpio. Manzanillo, 4/XII/2013, Carapace in general black with orange setae on border,
D. Ortiz, D. Barrales, G. Contreras (CNAN-Ar007171); with cheliceral longitudinal line of lighting setae (Fig.
3 ♀, Mpio. 11 km al SE. de Colima (CNAN-Ar007870, 35B). Legs and palpi with deep orange and pale orange
CNAN-Ar007872, CNAN-Ar007873); 1 ♀, Mpio. in patellae, tibiae and metatarsi with orange yellow/
Tecoman, 2/XII/2012, E. Goyer, E. Hijmensen, white setae on distal (Fig. 37I–L).
D. Ortiz (CNAN-Ar007871); Jalisco: 2 ♀, Mpio.
Pihuamo, 30/XI/2012, E. Goyer, E. Hijmensen, Note: Redescription of this species was done by
D. Ortiz (CNAN-Ar007874, CNAN-Ar010279); 1 ♂, Mendoza & Francke (2017).
Carretera entre Colima y Jalisco, carretera Colima-Cd.
Victoria, 8/VIII/2008, A. Cervantes, M. E. Olsen
(CNAN-Ar003425); Michoacán: 1 ♂, Mpio. Lázaro Brachypelma klaasi (Schmidt & Krausse, 1994)
Cardenás, 8/XI/2011, G. Vila (CNAN-Ar003340); 1 ♂,
Michoacán, Mpio. Aquila, G. Vila (CNAN-Ar003659). (Figs 2, 29–32, 36I–L, 38)
Brachypelmides klaasi Schmidt & Krause, 1994:
7, figs 1–2, D male and female; Schmidt (1997b: 19,
Diagnosis
figs 198, 202), male and female; Locht et al. (1999:
Brachypelma hamorii can be distinguished from all 196, figs 4, 9), male and female; Vol (1999: 11, fig.
other known Brachypelma species (except B. smithi) by A), female; Peters (2000: 75, figs 244–247), male and
the coloration of the legs with yellow-orange coloration female; Peters (2003: 131, figs 533–535, 537), male and
on patellae, tibiae and metatarsi (Figs 28, 37I–L). It female; Schmidt (2003: 137, figs 204, 208), male and
also differs in the shape of genitalia in both sexes with female. Smith (1994: 159) transferred the species to
palpal bulb curved upward and short (Fig. 27C–D) and Brachypelma.
spermatheca trapezoid rounded in shape (Fig. 27E–F). Brachypelma klaasi Smith (1994: 169, figs 926–939),
It differs from B. smithi by the narrow curved upward male and female.
embolus (Fig. 27C–D), the prolateral superior keel
shorter and dorsally wider (Fig. 27C) and the apical
keel slightly developed (Fig. 27B). It also differs in the Material examined
spermatheca ventral face smooth, with spermathecal Holotype of Brachypelmides klaasi (by original
baseplate elliptic (Fig. 27E–F). Although similar in designation): MEXICO: Nayarit: ♀, close to Tepic,
coloration, B. hamorii differs from B. smithi in the K. Böhme (SMF 40599) (only microscope slide with
patellae flame pattern not as brightly coloured as the spermatheca).
one exhibited by B. smithi (Fig. 37I–L). The sides of the
patellae are typically black rather than the yellowish- Other material: MEXICO: 1 ♀, without more data (SMF
orange as is typical in B. smithi (Fig. 35C–D). The 58101–84) (only microscope slide with spermatheca);
lateral setae are whitish throughout the length of legs, 1 ♂, without more data, Böhme (SMF 38044); Jalisco:
while in B. smithi, the same are yellowish (Fig. 37I– 2 ♂, 1 ♀, Mpio. Tomatlán, 6/XII/2012, D. Ortiz, E. Goyer,
L). In addition, B. hamorii typically have longitudinal E. Hijmensen (CNAN-Ar003333, CNAN-Ar003341,
cheliceral lines of lighting setae, which are not exhibited CNAN-Ar007831); 1 ♂, Mpio. Cihuatlán, 4/XII/2013,
by B. smithi (Fig. 35A–B). The carapace of B. hamorii is D. Ortiz, D. Barrales, G. Contreras (CNAN-Ar007160);
typically black, bordered with pink or orange in both 1 ♀, Mpio. La Huerta, Reserva Chamela, 6/XII/2013,
sexes (Fig. 28A–B, E–F). Rare variants of females can D. Ortíz, E. Goyer, E. Hijmensen (CNAN-Ar007879);
have a black radiating pattern, bordered with pale 1 ♂, Mpio. La Huerta, Reserva Chamela, 8–20/
yellow (Fig. 28D), and rare variants of males may have XI/2014, W. Maddison (CNAN-Ar007857); 1 ♂, Mpio.
a carapace that is almost yellowish (Fig. 28C). La Huerta, Reserva Chamela, 18/V/1981, A. Pescador
Brachypelma hamorii is identified by the following (CNAN-Ar003432); 1 ♂, donation received from
character combination: male palpal bulb with narrow private collection of J. Mendoza (CNAN-Ar003344);
spoon-like embolus, prolateral superior keel shorter Colima: 1 ♂, Mpio. Manzanillo, 4/XII/2013, D. Ortiz,
than in other species, thin and directed retrolaterally, D. Barrales, G. Contreras (CNAN-Ar007162); 1 ♂, Mpio.
prolateral inferior keel weakly developed directed Colima, 30/XI/2012, D. Ortiz, E. Goyer, E. Hijmensen
from dorsal to ventral, better seen dorsally, apical (CNAN-Ar007845).
keel smaller than in other species, short and thin (Fig.
27A–D). Embolus tapered throughout its length but
turning wide at apex, and slightly curved dorsally (Fig. Diagnosis
27C–D). Spermatheca trapezoid rounded; ventral face Brachypelma klaasi can be distinguished from all
is almost smooth; Spermathecal baseplate divided, other known Brachypelma species in the coloration of
Figure 29. Brachypelma klaasi, male CNAN-Ar003333. A, carapace, dorsal view; B, prosoma, ventral view; C, opisthosoma,
the legs with pinkish colour on the tibiae and metatarsi shorter than prolateral superior keel. Embolus tip
(Fig. 36I–L). It also differs in the shape of the genitalia directed to retrolateral. Embolus similar in length to
in both sexes with palpal bulb almost straight, tegulum (Fig. 30A–D). Spermatheca separated with
tapering and lacking the typical spoon-shape (Fig. semitriangular receptacles. Spermathecal baseplate
30C–D). The prolateral superior keel reduced, directed divided, oblong; twice wider than its height (Fig. 31D–F).
retrolaterally and slightly extended to backward Carapace of both sexes black (Fig. 32A–C).
(Fig. 30C). Apical keel thin and reduced, shorter than
prolateral superior keel (Fig. 30B). Lacking prolateral
inferior keel. Also differs by the divided spermatheca Redescription
widely separated, with spermathecal baseplate oblong. Male (CNAN-Ar003333) (Figs 29, 30, 32A): Body
Ventral face smooth (Fig. 31D–F). length 48.60 (not including chelicerae and spinnerets),
Brachypelma klaasi is identified by possesing the carapace 22.82 length, 22.17 width. Caput not markedly
following character combination: male palpal bulb with elevated; fovea straight, 4.15 wide (Fig. 29A). Eyes:
embolus almost straight, tapering and without typycal anterior eye row procurved, posterior eye row recurved.
spoon-shape due to the reduced apical keel. Prolateral Eye sizes and interocular distances: AME 0.55; ALE
superior keel reduced, apical keel thin and reduced, 0.80; PME 0.40; PLE 0.70; AME–AME 0.55; AME–ALE
Figure 30. Brachypelma klaasi, male CNAN-Ar003333. Left palpal bulb: A, dorsal view; B, ventral view; C, retrolateral
0.20; PME–PME 1.20; PME–PLE 0.05; ALE–PLE 0.23. Stridulatory setae: with claviform stridulating setae
Ocular tubercle width 2.80, length 2.55; clypeus 0.25 on palp trochanter and femur retrolateral face; leg
(Fig. 29D). Labium length 3.05, width 3.55; with 98 I trochanter and femur prolateral face. Chaetotaxy
cuspules. Maxilla inner corner with approximately 175 (left side): femora none; patellae none; tibiae II 1v; III
(left) and 171 (right) cuspules. Cheliceral promargin 3v; IV 2v; palp 1v; metatarsi I 1v; II 3v; III 1p, 5v; IV 9v.
with 11 (left) and 12 (right) teeth (proximal to distal: Palp: embolus almost straight, tapering and lacking
first–second large, third medium, fourth large, fifth the typical spoon-shape due to the reduced apical keel.
small, sixth–11th large; first large, second small, third Prolateral superior keel reduced, directed retrolaterally
large, fourth small, fifth large, sixth small, seventh– and slightly extended to backward; prolateral inferior
12th large). Sternum length 10.80, width 10.04. Sigilla keel absent. Apical keel thin and reduce, shorter than
oval, second to third pairs hardly visible, posterior pair prolateral superior keel. Opening of the embolus is on
once its length from the margin (Fig 29B). Leg formula: the prolateral side, just behind the opening is located
IV, I, II, III. Length of legs and palpal segments (femur, a concavity which delimits the apical keel boundary
patella, tibia, metatarsus, tarsus, total): I: 18.95, 10.29, from the remaining part of the embolus. Embolus apex
14.84, 15.15, 10.72, 69.95; II: 17.25, 9.79, 13.15, 14.09, slightly curved to retrolateral. Embolus similar in
10.07, 64.35; III: 16.11, 8.27, 11.83, 15.10, 10.28, 61.59; length as tegulum (Fig. 30A–D). Urticating setae: types
IV: 18.89, 9.38, 15.53, 18.32, 10.63, 72.75. Palp: 12.28, I and III arranged in one dorsoposterior patch, black
7.47, 11.92, -, 5.71, 37.38. Spinnerets: PMS, 2.40 long, in colour. Type III are located in an oval dorsomedial
1.15 apart; PLS, 4.05 basal, 2.35 middle, 3.60 distal. area extended to posterior. Type I surround the type
Tarsi I–IV entirely scopulated. Metatarsus I entirely III area, with intermediates between type III and I in
scopulated, II scopulated 90%, III scopulated 60% transition areas (Fig. 29C).
distally, IV scopulated 40% distally. Tibia I with two
tibial apophyses normally developed, which originate Female (CNAN-Ar007831) (Figs 31, 32B): Body length
from a common base. Prolateral apophysis with 63.26 (not including chelicerae and spinnerets),
inner spine half its length; retrolateral apophysis carapace 27.65 length, 24.27 width. Caput not
tapering throughout its length, apex slightly curved to markedly elevated; fovea straight, 6.40 wide (Figs 31A).
prolateral (Fig. 29F, G). Metatarsus I curved (Fig. 29E). Eyes: anterior eye row procurved, posterior eye row
Figure 31. Brachypelma klaasi. A–D, female CNAN-Ar007831. A, carapace, dorsal view; B, prosoma, ventral view; C,
opisthosoma, dorsal view; D, spermatheca, ventral view; E–F, spermatheca ventral view of: E, female SMF-40599; F, female
SMF-58101. Scale = 10 mm (A–C), 1 mm (D–F).
recurved. Eye sizes and interocular distances: AME femur prolateral face. Chaetotaxy (left side): femora
0.57; ALE 0.63; PME 0.43; PLE 0.67; AME–AME none; patellae none; tibiae I 2v; II 2v; III 3v; IV 2v;
0.50; AME–ALE 0.33; PME–PME 1.33; PME–PLE palp 1p, 5v; metatarsi I 3v; II 4v; III 5v; IV 1p, 14v, 1r.
0.07; ALE–PLE 0.07. Ocular tubercle width 2.80, Genitalia: spermatheca separated with subtriangular
length 2.33; clypeus 0.47. Labium length 3.65, width receptacles strongly sclerotized, each receptacle almost
5.25; with 94 cuspules. Maxilla inner corner with the same height as width. Spermatheca baseplate
approximately 68 (left)–58 (right) cuspules. Cheliceral divided and widely separated, each baseplate oblong
promargin with 10 (left)–11 (right) teeth. Sternum twice wider than high (Fig. 31D). Baseplate division
length 11.60, width 10.63. Sigilla oval, first to third can vary in length (Fig. 31E, F). Urticating setae: types
pairs hardly visible; posterior pair once its length I and III arranged in one dorsoposterior patch, black
from the margin (Fig. 31B). Leg formula: IV, I, II, III. in colour. Type III are located in an oval dorsomedial
Length of legs and palpal segments (femur, patella, area extended to posterior. Type I surround the type
tibia, metatarsus, tarsus, total): I: 16.86, 10.39, 12.94, III area, with intermediates between type III and I in
12.74, 8.92, 61.85; II: 15.61, 10.06, 11.28, 12.49, 9.04, transition areas (Fig. 31C).
58.48; III: 14.82, 9.59, 10.07, 13.65, 9.09, 57.22; IV:
17.26, 9.68, 12.32, 18.29, 9.53, 67.08. Palp: 12.55, Colour pattern: In live specimens, adults of both
8.02, 9.09, -, 9.29, 38.95. Spinnerets: PMS, 2.80 long, sexes have the carapace black (pantone process
3.65 apart; PLS, 5.50 basal, 4.15 middle, 4.75 distal. black c) with zinnwaldite colour (pantone 489c)
Tarsi I–IV entirely scopulated. Metatarsus I entirely around the border; despite the black carapace, the
scopulated, II scopulated 80%, III scopulated 70% radiating thoracic sulci are clearly seen (Fig. 32A–C);
distally, IV scopulated 50% distally. Stridulatory setae: chelicerae dorsally brown colour (pantone 7533c)
with claviform stridulating setae on palp trochanter with some large setae French beige colour (pantone
and femur retrolateral face; leg I trochanter and 4715c); ventral coxae, labium, maxillae and sternum
Figure 32. A–C, Brachypelma klaasi, habitus; D, habitat. A, male; B, female from coast line; C, female from inland; D,
deciduous forest in the habitat of B. klaasi. Photos: J. Mendoza.
black pearl colour (pantone black 6c); abdomen thickets, tree roots or burrows on hillsides, in both
dorsally black with corvette colour setae (pantone forested and moderately disturbed areas. Burrows
720c), ventrally black pearl colour. Legs: femora black do not have any silk around the entrance to indicate
pearl; patellae black pearl with scattered setae shilo there is a spider inside. This species is sympatric
colour (pantone 488c); tibiae and metatarsi mandys (overlapping in range) with Brachypelma emilia in
pink colour (pantone 473c); tarsi black (pantone black south-western Nayarit and with Brachypelma hamorii
c); palpi: femora black pearl; patellae, tibia and tarsi in Colima.
black pearl with scattered setae shilo colour (Fig.
36I–L). With juveniles of same pattern but paler in
colour. Brachypelma smithi (F. O. Pickard-Cambridge,
1897)
Distribution and habitat: Brachypelma klaasi is (Figs 2, 33, 34, 35A, C, 37M–P, 38)
known from the Pacific Coast region of Jalisco on the
western side of Sierra Madre Occidental, with small Eurypelma smithi F. O. Pickard-Cambridge, 1897: 20,
populations in Colima and southern Nayarit (Figs 2, plate 1, fig. 4, D female.
38). It occurs in drier coastal thorn scrub, deciduous Brachypelma smithi (F. O. Pickard-Cambridge, 1897):
forest and higher elevations oak forest (Fig. 32D). It Pocock (1903: 103); Locht et al. (1999: 198, fig. 5),
is a fossorial species whose modified or self-excavated male; Mendoza & Francke (2017: 162, figs 3–22,
burrows, can be found under large rocks, under thorny 42–45, 50–51, 53–56), male and female.
Figure 33. A–H, Brachypelma smithi, male CNAN-Ar007832. Left palpal bulb: A, dorsal view; B, ventral view; C, retrolateral
view; D, prolateral view. E–H, spermatheca ventral view of: E, female CNAN-Ar003611; F, female CNAN-Ar004131; G,
female CNAN-Ar007896; H, female CNAN-Ar007904. Scale = 2 mm (A–H).
Figure 34. A–F, Brachypelma smithi, habitus. A, male in habitat (Guerrero); B, male of black carapace in habitat
(Guerrero); C, male CNAN-Ar007832 in life; D, female CNAN-Ar007144 in life; E, female CNANAr007901 in life; F, female
CNAN-Ar007147. Photos: J. Mendoza.
Brachypelma annitha Tesmoingt, Cléton & Verdez, male; Peters (2000: 64, figs 205–207), male and
1997a: 9, plates 1–6, D male and female; female; Peters (2003: 108, figs 428, 435–436),
Tesmoingt et al. (1997b: 2, plates 7–8, 11), male and female. Mendoza & Francke (2017:
Figure 35. A–D, Brachypelma smithi and Brachypelma hamorii comparisons. A–B, comparison of the chelicerae in B. smithi
and B. hamorii, the arrow shows the typical longitudinal lighter lines over chelicerae of B. hamorii (B), which are absent in
B. smithi (A); C–D, comparison of the patellae in B. smithi and B. hamorii, the arrow point to the sides of the patellae, which
are typically black in B. hamorii (D), while in B. smithi they are orange (A). Photos: J. Mendoza.
162) considered this as a junior synonym of CNAN-Ar010275); 1 ♂, 1 ♀, Mpio. Acapulco, El

B. smithi). Quemado 14/X/2014, A. Ortega (CNAN-Ar007897,
CNAN-Ar007896); 1 ♀, Mpio. Josue Azueta, 24/VIII/2015,
D. Ortiz, J. Baldazo (CNAN-Ar007904).
Material examined
Holotype of Brachypelma smithi: MEXICO: Guerrero:
Juvenile ♂ (labelled as ♀), Dos Arroyos. H. H. Smith Diagnosis
(BMNH 1143; also labelled BM1898.12.24.33). Brachypelma smithi can be distinguished from all
other known Brachypelma species (except B. hamorii)
Other material: MEXICO: Guerrero: 1 ♂, 2 ♀, Mpio. by the coloration of the carapace and legs, consisting
Acapulco, Dos Arroyos. 13/XII/2013, J. Mendoza of red-orange coloration on the patellae, tibiae and
(CNAN-Ar007146, CNAN-Ar007143, CNAN-Ar007144); metatarsi (Fig. 37M–P), as well as orange-black
1♂, Mpio. Acapulco, 04/I/1965, E. Rivapalacios starburst striations on the carapace in general (Fig.
(CNAN-Ar003086); 1 ♂, Mpio. Atoyac, 27/II/1984, 34D). The shape of the genitalia also differs in both
J. G. Julio (CNAN-Ar003434); 1 ♀, Mpio. Acapulco, 10/ sexes with the palpal bulb being straight and broad
VII/1979, M. Adams (CNAN-Ar003435); 1 ♂, Mpio. (Fig. 33C–D) and the spermatheca shaped like a
Acapulco, without additional data (CNAN-Ar003594); 2 ♀, trapezoid (Fig. 33E–H). It differs from B. hamorii
donation received from private collection of J. Mendoza in the straight palpal bulb having a broad spoon-
(identified in pet trade as B. annitha) (CNAN-Ar004131, shape, a wider apical keel and the prolateral superior
CNAN-Ar003611); 3 ♂, Mpio. Coyuca de Benítez, 29/ keel is not as elevated (Fig. 33A–D). It also differs
VIII/2015, J. Mendoza, R. Ramírez (CNAN-Ar010281, in the spermatheca with the spermathecal baseplate
Figure 36. A–D, Brachypelma auratum, female (CNAN-Ar007878) A, leg I; B, leg II; C, leg III; D, leg IV; E–H, Brachypelma
emilia, female (CNAN-Ar010602) E, leg I; F, leg II; G, leg III; H, leg IV. I–L, Brachypelma klaasi, female (CNAN-Ar007831)
I, leg I; J, leg II; K, leg III; L, leg IV.
Figure 37. A–D, Brachypelma baumgarteni, female (CNAN-Ar007150) A, leg I; B, leg II; C, leg III; D, leg IV; E–H,
Brachypelma boehmei, female (CNAN-Ar007905) E, leg I; F, leg II; G, leg III; H, leg IV; I–L, Brachypelma hamorii, female
(CNAN-Ar007874) I, leg I; J, leg II; K, leg III; L, leg IV; M–P, Brachypelma smithi, female (CNAN-Ar007144) M, leg I; N, leg
II; O, leg III; P, leg IV.
being divided and subtriangular (Fig. 33E–H). The Brachypelma smithi is identified by possessing
spermathecal ventral face is also striated, whereas it the following character combination: male palpal
is smooth in B. hamorii (Fig. 33F). Although similar bulb with broad spoon-like embolus, prolateral
in coloration, B. smithi differs from B. hamorii by the superior keel normally developed, thin and directed
patellae flame pattern brightly coloured (Fig 37M–P). retrolaterally, prolateral inferior keel weakly
The sides of the patellae are yellowish-orange rather developed, better seen dorsally, apical keel strongly
than black as is typical in B. hamorii (Fig. 35C). The developed, wider in the middle (Fig. 33C–D).
lateral setae are yellowish throughout the length of Embolus straight and wide throughout its length,
legs (Fig. 37M–P), whereas in B. hamorii the same one and half times longer than tegulum (Fig.
are whitish, providing greater contrast with darker 33A–B). Spermatheca semicircular or trapezoidal
areas (Fig. 37I–L). In addition, B. smithi does not tend (Fig. 33E–H); ventral face striated (Fig. 33F).
to show the longitudinal cheliceral lines of lighting Spermathecal baseplate divided, subtriangular; four
setae typically exhibited by B. hamorii (Fig. 35A). times wider than its height (Fig. 33E–H). Carapace
Carapace of B. smithi females can be black bordered of adult male typically orange in colour (Fig. 34C),
with orange (Fig. 34F), with a black radiating pattern whereas in females almost black with orange around
bordered with orange (Fig. 34D) or, almost completely the border, with black striated pattern and orange
orange with black around the ocular area (Fig. 34E). around or almost orange with only black caput area
Males normally have orange carapace (Fig. 34C), rare (Fig. 34D–F).
variants of males may have also an orange carapace
with scant black around ocular area (Fig. 34A) or a Note: Redescription of this species was made by
black carapace bordered with orange (Fig. 34B). Mendoza & Francke (2017).
Figure 38. Geographic distribution of the genus Brachypelma from published records and specimens collected or examined
for this study.
Identification key for species of Brachypelma Simon 1891

Adult males
1. With any other colour besides black in any segment(s) of the legs (generally orange or reddish) (Figs 31, 32)
�� …………………………2
With all segments of the legs black in colour, carapace golden yellowish (Fig. 8A)�� B. albiceps
2. All patellae with any other colour besides black (generally pink or reddish) (Figs 31A–D, I–L, 32)�� 3
All patellae totally black in colour, with orange coloration in all tibiae and metatarsus IV, male
palpal bulb with narrow spoon-shape, prolateral superior keel slightly developed (Figs 24, 26A,
31E–H)…………………….B. emilia
3. All patellae with a central flame pattern orange or reddish, male palpal bulb with a distinct spoon-shape,
apical keel curved outward (Figs 10, 15, 32A–H, 32)�� 4
All patellae without central flame pattern, with pinkish colour on patellae, tibiae and metatarsi, male
palpal bulb without distinct spoon-shape, apical keel curved inward (Figs 28, 30A)��B. klaasi
4. Embolus equal or slightly larger than tegulum and broad at apex with large spoon-shape, prolateral
superior keel narrow and widely extended backwards (Fig. 10)�� 5
Embolus shorter than tegulum and broad at apex with short spoon-shape, prolateral superior keel wide
and short, not extended widely to backwards (Fig. 15)�� 7
5. All patellae orange or yellowish with central flame-shape area reddish in colour, without distal white
rings on patellae or tibiae; large embolus base and slightly narrower than the apical spoon-shape
(Figs 19, 22A)�� 6
All patellae black with a central flame-shape area reddish or orange, with a distal white ring on patellae,
tibiae and metatarsi; short embolus base and similar in width to the apical spoon-shape (Figs 10, 13A)
�� B. auratum
6. Carapace orange or black, patellae orange with central flame-shape reddish area, tibiae with orange in
distal half, metatarsi with yellowish ring distally; ventral apophysis wide through its length, apex slightly
curved to dorsal; apical keel very wide and larger than prolateral superior keel (see Mendoza & Francke,
2017: figs 3, 7, 8, 19, 42–45)�� B. smithi
Carapace always orange, patellae, tibiae and metatarsi completely orange in colour; ventral apophysis
wide in base and tapering, apex straight; apical keel wide and similar in length or slightly larger than
prolateral superior keel (Figs 18F, 19, 22A)�� B. boehmei
7. Carapace always orange, patellae yellowish with central flame-shape reddish area, tibiae and metatarsi
yellowish, with a distinctive diagonal yellow line on metatarsi; tegulum swollen, embolus base very short,
apical keel wide and longer than prolateral superior keel (Figs 15, 17A)�� B. baumgarteni
Carapace black or yellowish with only ocular area with some black, yellowish colour around the border,
patellae dorsally orange slightly expanded laterally, with central flame-shape reddish area, tibiae distal
half yellowish, metatarsi with distal white ring; tegulum globose but not swollen, embolus base short and
thin, apical keel slightly wide and similar in length to prolateral superior, apex very curved to retrolateral
(see Mendoza & Francke, 2017: figs 24, 28, 29, 37, 46–49)�� B. hamorii
Adult females
1. With any other colour besides black in any segment(s) of the legs (generally orange or reddish) (Figs 31, 32)
�� 2
With all segments of the legs black in colour, carapace golden yellowish; spermatheca separated and with
rounded receptacles (Figs 8B, 7E–G)�� B. albiceps
2. All patellae with any other colour besides black (generally rose or reddish) (Figs 31A–D, 31I–L, 32)�� 3
All patellae totally black in colour, with orange coloration in all tibiae and metatarsus IV, spermatheca
with a single receptacle strongly sclerotized slightly notched in the middle, spermathecal baseplate
oblanceolate (Figs 25E, F, 26B, C, 31E–H)�� B. emilia
3. All patellae with a central flame pattern orange or reddish, spermatheca fused with single receptacle (Figs
12E, F, 21E–G, 32A–H)�� 4
All patellae without central flame pattern, with pinkish colour on patellae, tibiae and metatarsi,
spermatheca separated with semitriangular receptacles, spermathecal baseplate oblong (Figs 29D–F, 30B,
C, 31I–L)��B. klaasi
4. Spermathecal baseplate lower than the seminal receptacle, spermatheca ventral face striated (Figs 12E–F,
21E–G)�� 5
Spermathecal baseplate higher than the seminal receptacle, spermatheca ventral face smooth (Fig. 16E–G)
�� 7
5. All patellae orange or yellowish with central flame-shape area reddish in colour, without distal white
rings on patellae or tibiae; spermatheca ventral face with clearly defined striatation (Figs 21E–G, 22B, C,
32E–H)�� 6
All patellae black with a central flame-shape area reddish or orange, with a distal white ring in patella,
tibiae and metatarsi; spermatheca ventral face with slight striatation, spermatheca baseplate more
separated above, each baseplate ovate (Figs 12E, F, 13B, C, 31A–D)�� B. auratum
6. Carapace could be yellowish around the border and behind the fovea with starburst black pattern from
the fovea to caput; or yellowish around the border and black in dorsomedial; or yellowish pink in almost
all carapace except by two longitudinal black areas in the caput; all patellae orange with central flame-
shape reddish area, tibiae with orange in distal half, metatarsi with yellowish ring distally; spermatheca
baseplate subtriangular, decreasing the upper side toward the outer side (see Mendoza & Francke, 2017:
figs 14–18, 20–22, 50, 51, 53–56)�� B. smithi
Carapace could be completely orange or orange in almost all carapace except by two longitudinal black
areas in the caput; patellae, tibiae and metatarsi completely orange in colour; spermatheca baseplate
widely separated in middle, each baseplate lanceolate, outer side slightly smaller than the inner (Figs 20B,
21E–G, 22B, C)�� B. boehmei
7. Carapace could be light orange around the border and black in dorsomedial or light orange around the
border with orange extended behind the fovea and black from the fovea to caput; patellae yellowish with
central flame-shape reddish area, tibiae and metatarsi yellowish, with a distinctive diagonal yellow line
on metatarsi; spermatheca baseplate oblong, baseplate division narrow, baseplate poorly sclerotized in the
basal inner corner (Figs 16E–G, 17B, C)�� B. baumgarteni
Carapace could be yellowish around the border and black in dorsomedial or pale orange around the border
and behind the fovea with starburst black pattern from the fovea to caput; patellae dorsally orange slightly
expanded laterally, with central flame-shape reddish area, tibiae distal half yellowish, metatarsi with
distal white ring; spermatheca baseplate elliptic, outer side slightly smaller than the inner (see Mendoza
& Francke, 2017: figs 34–36, 38–40, 52, 57–60)�� B. hamorii
Tliltocatl Mendoza & Francke, gen. nov. length or longer than the prolateral superior (Fig 39E),
the apical keel can extend widely to backwards just
(Figs 39–44) as the prolateral inferior keel and usually is broader
urn:lsid:zoobank.org:act:EEEC09EE-B64B- on its distal half (Figs 39D, E; 40B, D–F); (4) females
4F7E-83A0-D174637009AC have a single fused spermatheca, apically narrowed
(Figs 39G–H, 41A); (5) spermathecal baseplate absent
Type species: Eurypelma vagans Ausserer, 1875, herein or slightly developed, poorly sclerotized (Fig. 39H); (6)
designated. both sexes lack a plumose pad of setae on leg IV femur;
and (7) all tarsi scopulae are undivided. It differs
Species included: Tliltocatl albopilosum (Valerio, 1980), from Brachypelma by the coloration of legs, which are
comb. nov., Tliltocatl epicureanum (Chamberlin, 1925), black (Figs 42, 43B–D) or have long, whitish setae (as
comb. nov., Tliltocatl kahlenbergi (Rudloff, 2008), comb. T. albopilosum, Fig. 43A) in combination with a dark
nov., Tliltocatl sabulosum (F. O. Pickard-Cambridge, carapace and long red/yellowish setae on abdomen
1897), comb. nov., Tliltocatl schroederi (Rudloff, 2003), (Figs 42, 43B–D). The shape of genitalia also differs
comb. nov., Tliltocatl vagans (Ausserer, 1875), comb. in both sexes with the male palpal bulb apex larger
nov., Tliltocatl verdezi (Schmidt, 2003), comb. nov. than in Brachypelma and by the presence of prolateral
inferior keel well developed and posteriorly extended
(Figs 39A, D–F, 40A, D, E). The apical keel is also
Diagnosis larger than in Brachypelma (Figs 30D, 33D) and wider
The new genus Tliltocatl can be distinguished on distal half (Figs 39B, D, E, 40B, D, E). Embolus is
from all other known theraphosinae genera (except regularly similar in length or longer than the tegulum
Brachypelma) by the following character combination: (Figs 39C–D, 40C–F), whereas in Brachypelma it is
(1) having just claviform stridulating setae on shorter (Fig. 10D). It also differs in having spination
the prolateral face of leg I trochanter/femur and on the patellae of palps and legs. Females differs in
retrolateral face trochanter of the palp; (2) both the spermatheca apex inwardly curved and by lacking
sexes possess always urticating setae types I and III spermathecal baseplate (Fig. 39G) or poorly sclerotized
– type III are located in the dorsoposterior area and and widely separated when present (Fig. 39H).
type I surrounding these; (3) the male palpal bulb
distally wide and flattened (spoon-shaped) and has
prolateral superior and apical keels united at the Description
apex (Figs 39C, D, 40C–F), prolateral superior and Carapace regularly as long as wide, caput slightly
prolateral inferior keels are at similar height, joined elevated (Figs 42, 43). Cephalic striae inconspicuous
at their distal end and widely separating towards (Fig. 43A). Fovea deep, straight or recurve. Eye tubercle
the embolus base (better seen in dorsal position) distinct and raised, wider than long. Anterior eye row
(Fig. 39F), the prolateral inferior keel is similar in procurved, posterior eye row recurved. Clypeus narrow
Figure 39. A–G, Tliltocatl vagans; H, Tliltocatl albopilosum. A–F, male CNAN-Ar004123. Left palpal bulb:
A, dorsal view; B, ventral view; C, retrolateral view; D, prolateral view; E, embolus prolateral view; F, embolus dorsal
view. G, spermatheca ventral view of female CNAN-Ar010575. H, spermatheca ventral view of female allotype UCR-504.
Scale = 2 mm (A–H).
Figure 40. A–D, Tliltocatl epicureanum CNAN-Ar010578. E–F, Tliltocatl albopilosum holotype UCR-258. Left
palpal bulb: A, dorsal view; B, ventral view; C, retrolateral view; D, prolateral view. Right palpal bulb: E, prolateral view, F,
retrolateral view. Photos: J. Mendoza. Scale = 2 mm (A–F).
or absent. Labium subtrapezoidal, wider than long, retrolateral scopulae. Claviform stridulating setae on palp
with 85–120 cuspules on anterior third centre. Maxilla trochanter retrolateral face and in leg I trochanter and
subrectangular, anterior lobe distinctly produced into femur prolateral face. Patellae of the legs have at least one
conical process, inner corner with 130–220 cuspules. spine on prolateral or retrolateral side. Posterior lateral
Sternum longer than wide. Anterior pair of sigillae spinneret distally elongating, digitiform. Both sexes
circular and slightly seen, second pair oval enlarged, possess urticating setae type I and type III; Type III are
posterior pair oval or rounded, generally once its length located in the dorsoposterior area and type I surrounding
from the sternum margin. Leg formula: I, II, III, IV. Tarsi these. Males possess two tibial apophysis, retrolateral
I–IV fully scopulated. All tarsi scopulae are undivided. apophysis slightly curved in apically. Globous bulb
Metatarsi I–II fully scopulated. Metatarsus III is 50% with small subtegulum longer than its height. Embolus
distally scopulated and metatarsus IV is 20–40% distally flattened (Fig. 39A, B), longer than tegulum (Fig. 39C, D),
scopulated. The femur of leg III is slightly enlarged with prolateral superior and apical keel large, joint at the
but not swollen as in other genera. Femora IV without embolus tip forming a typical spoon-shape (Figs 39C–E,
40C–F). Prolateral inferior keel well developed, parallel the Pacific coast, Mexican Gulf and Atlantic Coast.
to prolateral superior and larger in general (Figs 39D– Specimens live inside burrows under flat rocks, fallen
F, 40A, D, E). Females with a simple undivided/fused logs, sidehills, tree roots and even some species dig
spermatheca apically narrowed and medially curved burrows in farmlands, gardens or close to flooded land.
inwardly (Fig. 39G, H). Spermathecal baseplate absent
or slightly developed (Fig. 39H). Spermatheca midwidth
shorter than its base and well sclerotized (Fig. 39G, H). Species considered nomina dubia
Most of the species are black with long, red setae on the Brachypelma andrewi Schmidt, 1992: Schmidt
opisthosoma (except T. albopilosum, T. schroederi and (1992a) described this species based on a single
T. verdezi) (Figs 42–43). Juveniles are similar in colour to specimen located in the collection of the British
the adults but paler. Museum of Natural History (now The Natural History
Museum, London). The history of this specimen was
elaborated on by Smith (1992); the holotype used for
Remarks the description of B. andrewi comes from the L. Koch
Tliltocatl can be confused by unexperienced people collection. Another specimen from the same Koch
with Sericopelma spp. due to the similar coloration, collection was found in the NHM containing an alleged
but they can be easily differentiated from each undescribed specimen. Initially the jar that contained
other, because adult males of Sericopelma lack tibial the specimen, of what would later be described as
apophysis, the female spermatheca of Sericopelma is B. andrewi, was labelled as the holotype of Euathlus
strongly sclerotized (Fig. 41B), also the spermatheca truculentus Ausserer, 1875. This created a confusion
is distinctly swollen on the apex showing a P-shape that resulted in the genus Brachypelma being listed
(better seen laterally). In addition, Sericopelma has a as a junior synonym of Euathlus, as was proposed by
distinct radiating sulcus on the carapace (Gabriel & Raven (1985). Smith (1992) redescribed the specimen
Longhorn, 2015). in the NHM labelled as the Euathlus truculentus
holotype believing it to be the one used by Ausserer
(1875). He used his description to support Raven’s
Etymology decision that the genus Brachypelma was synonymous
The genus gender is masculine. The name is a noun with Euathlus. However, shortly afterwards, Schmidt
in apposition comprising the Nahuatl words Tlil, (1992a) observed that the specimen seen by Raven, and
which means ‘black’, and tocatl, which means ‘spider’, later by Smith, was not the same species as the type
referring to the black coloration of species in the genus. of Euathlus truculentus, but, was in fact a new species
of Brachypelma. Schmidt stated as follows: ‘Raven
had almost come to the conclusion that Brachypelma
Distribution was a junior synonym of Euathlus, while Smith, after
Tliltocatl occurs in Mexico, Guatemala, Belize, studying the said Brachypelma, was of the opinion
Honduras, El Salvador, Nicaragua and Costa Rica (Fig. that Raven was right with his synonymizing. He had
44). The species of the genus are found in deciduous not noticed that only the body type of the Euathlus
forest, evergreen rain forest and grasslands along
Figure 41. A, Tliltocatl epicureanum, B, Sericopelma melanotarsum. A, spermatheca ventral view of female
CNAN-Ar007880. B, spermatheca ventral view of female UCR-without voucher number. Scale = 2 mm (A–B).
Figure 42. A–F, Tliltocatl spp, habitus. A–B, Tliltocatl epicureanum: A, male from type locality (Yucatán); B, female
from type locality (Yucatán). C–D, Tliltocatl kahlenbergi: C, female (Veracruz); D, male (Oaxaca); E–F, Tliltocatl
schroeder: E, female (Oaxaca); F, male (Oaxaca). Photos: J. Mendoza.
type, called Ausserer, deviates considerably from those were transferred to Tliltocatl in this work). So, we
of the alleged 2nd Euathlus truculentus: e.g. in the real propose that this species be transferred as Tliltocatl
type, the body length is 44 mm, while in the erroneously andrewi comb. nov. However, without being able to
paratype Brachypelma is 55 mm. This Brachypelma is make a direct observation of the specimen, it is not
said to originate from Cuba and has not been mentioned possible to correctly identify the species or compare
so far – I shared the error with Smith in a letter dated it with those closely related and already described.
28 May 1992 and confirmed that he was wrong in the Additionally, given the lack of data on its distribution,
matter of synonymizing and was no longer tenable.’ it is not possible to obtain new material. Therefore,
After this, Schmidt (1992a) named the undescribed based on the loss of the holotype and in its inability to
Brachypelma as B. andrewi after Andrew Smith. be identified, we propose that the species T. andrewi be
Unfortunately, because of the lack of a useful locality considered nomen dubium.
for this species, no new specimens can be collected for
comparison. Also, after our visit to the arachnological Brachypelma aureoceps (Chamberlin, 1917): This
collection of the NHM, it was not possible to locate the species was described by Chamberlin (1917) based
type specimen of Brachypelma andrewi, which is why it on a single female (RVC43 – examined) from Florida,
is considered lost to date. After reviewing, in detail, the Tortugas. However, there is no evidence of native
redescription of the specimen made by Smith (1992), it tarantulas from Florida (Smith, 1994; West, personal
is possible to know that B. andrewi is a species of the communication). Smith (1994) mentioned that is highly
‘red rump complex’ (vagans group) tarantulas (which probable that the B. aureoceps specimen was accidentally
Figure 43. A–D, Tliltocatl spp, habitus. A, Tliltocatl albopilosum, female (Upala); B, Tliltocatl verdezi, male
(Guerrero). C–D, Tliltocatl vagans: C, male (Campeche); D, female (Campeche). Photos: J. Mendoza.
Figure 44. Geographic distribution of the described species of the genus Tliltocatl from published records and specimens
collected or examined for this study from Mexico to Costa Rica. Also, is recorded de distribution of Stichoplastoris fossorius
in Costa Rica.
imported into the Florida Keys. This specimen has were able to determine that it does not belong to the
morphological features, such as spermatheca without a genus Citharacanthus Pocock (1901), and it is not a
baseplate, highly notched medially, patellae spination female, but a subadult male belonging to Tliltocatl.
and claviform setae on prolateral trochanter and femur The main evidence of this is the presence of type I and
of leg I, showing that it belongs to the genus Tliltocatl. III urticating setae, in combination with stridulatory
So, we propose that this species be transferred as claviform setae on trochanter and femur I and palp
Tliltocatl aureoceps comb. nov. Nevertheless, this trochanter. It is evident that the specimen is a subadult
species is similar to other valid species distributed in male just by the presence of the accessory organs. These
Mexico (e.g. T. kahlenbergi and T. vagans and cannot gland-like structures can be prominent in some groups,
be differentiated adequately from these. In such a way such as Tliltocatl, and are often mistaken for paired
that molecular data or accurate morphological features spermatheca (as was in this case). The male accessory
are needed to properly differentiate this species, and organs do not have direct connection with the gonopore
without the possibility to collect fresh specimens we and neither do they present uterus externus as can be
consider T. aureoceps a nomen dubium. observed in a spermatheca. Based on the evidence we
proposed to transfer the species as Tliltocatl alvarezi,
Citharacanthus alvarezi Estrada-Álvarez et al., 2013: comb. nov. Unfortunately, subadult males do not show
This species was described by Estrada-Álvarez et al. features that could be useful to identify a species. Due
(2013) based on a single specimen referred to as female to the lack of morphological and/or molecular data that
(CNAN-T01275 – examined) from an uncertain locality would allow us to make a comparison with the already
in Tuxtla Gutierrez, Chiapas, that was donated by an described known species of Tliltocatl, as well as a more
unknown collector to the Zoológico Miguel Álvarez del precise collection locality, we propose T. alvarezi to be
Toro (ZooMAT). After examination of the holotype, we
considered a nomen dubium. The following has been an unidentified species of Tliltocatl, possibly pet-trade
transferred to another group: material, making it difficult to identify.
Holotype and paratype of Brachypelma fossorium:

Stichoplastoris fossorius (Valerio, 1980) comb. COSTA RICA: Guanacaste: 1♂, Filadelfia. Col.
nov. E. Herrera (UCR-238) examined. 1♀, Liberia, finca
Santo Tomas, without more data (UCR-126) not
(Figs 44, 45)
examined.
Brachypelma fossoria Valerio, 1980: 271, fig. 25–28,
D male and female; Smith (1986: 49, fig. 28h), male; Other material examined: COSTA RICA: Guanacaste:
Smith (1987: 49, fig. 28h), male. 2♀, Canton Santa Cruz, Huacas, without more data.
Brachypelma fossorium: Schmidt (1992: 10, figs 9–12)

transferred male and female from Euathlus; Rudloff Remarks
(2003: 8, figs 26–29), male; Schmidt (2003: 153, Originally Valerio (1980) placed this species in the
figs 278–281), male and female. genus Brachypelma based only in the shape of the
male palpal bulb and female genitalia. However,
Misidentification: Peters (2000: 70, fig. 228), male; after the revision of the holotype and based in the
Peters (2003: 121, figs 486, 488–489), male and female. cladistic analysis we conclude that this species does
Peters (2000) and Peters (2003) show as B. fossorium not belong to Brachypelma or Tliltocatl. This species
Figure 45. A–D, Stichoplastoris fossorius: A–B, habitus; C–D embolus holotype UCR-238. A, female in habitat (Guanacaste);
B, male in habitat (Guanacaste); C–D, right palpal bulb: C, prolateral view; D, retrolateral view. Scale = 2 mm (C, D). Photos:
J. Mendoza.
has a short embolus with presence of prolateral has multiple taxonomic changes (Simon, 1891, 1903;
superior, prolateral inferior and apical keels, but Pickard-Cambridge, 1897; Roewer, 1942; Valerio, 1980;
with a different configuration from that observed in Raven, 1985; Schmidt, 1992). In the early 1990s, new
Brachypelma and Tliltocatl (Fig. 45C–D). The keel Brachypelma species were described based mainly on
arrangement is more similar to Stichoplastoris Rudloff material collected for the pet trade (e.g. B. auratum,
(1997). The urticating setae present in B. fossorium B. baumgarteni, B. boehmei, B. klaasi, B. ruhnaui,
are only type I, whereas Brachypelma and Tliltocatl B. hamorii and B. annitha). Although there were
possesses type I and III. Also, B. fossorium has the evident colour differences among the species from the
scopula IV divided by a strong band of setae, whereas Mexican Pacific Coast and Mexico–south-west Central
Brachypelma and Tliltocatl have all their scopulae America, no study had been done in depth to prove
entire. These characteristics fit those found in some the monophyly of the genus. Rudloff (2003) was the
species of the genus Stichoplastoris [e.g. S. elusinus first to propose a designation of complexes of species
(Valerio, 1980), S. obelix (Valerio, 1980)]. The weakest for Brachypelma based on colour characteristics
point of the kinship with Stichoplastoris would be and with no intention of making nomenlatural
the general shape of the male palpal bulb that is changes. He postulated a group called ‘vagans’
wide in the apex, and the spermatheca of the female formed by the species with dark to black base colour
that is unilobular, whereas in Stichoplastoris the on them (B. albopilosum, B. angustum, B. aureoceps,
bulb looks thinner apically and the spermatheca is B. e m b r i t h e s , B. e p i c u r e a n u m , B. fo s s o r i u m ,
divided (although in species like S. obelix it is wider B. sabulosum, B. schroederi and B. vagans) and a group
at its base). Despite this, there are similar examples in named ‘emilia’ formed by the species with red/orange
genera such as Brachypelma in which species such as banded legs (B. annitha, B. auratum, B. baumgarteni,
B. klaasi and B. albiceps have thinner palpal bulbs and B. boehmei, B. emilia, B. klaasi and B. smithi). He also
a divided spermatheca, unlike the rest of the species said that the species B. ruhnaui (currently B. albiceps)
in the genus that have wider bulbs apically and fused cannot be classified in this scheme and that it was
spermatheca. General body shape and legs of males possibly more closely related to Aphonopelma than to
and females are not as thick as in Brachypelma and Brachypelma.
Tliltocatl (Fig. 45A–B). We, therefore, propose, based Petersen et al. (2007) developed a method to obtain
on the similarities indicated above, that B. fossorium mtDNA from Brachypelma spp. using exuvia. Their
be transferred to the genus Stichoplastoris creating results show a phylogeny based on a cytochrome
the new combination Stichoplastoris fossorius. oxidase 1 (COI) gene fragment trimmed to 205bp
where two Brachypelma subgroups corresponding to
the ‘red rump vagans group’ and the ‘red leg emilia
DISCUSSION group’ were recovered, but were simply (mis)-grouped
together, because no other related genera were used
Non-monophyly of Brachypelma and the use of for comparison. Mendoza & Francke (2017) used
barcodes molecular data for a phylogenetic analysis with COI
Simon (1891) described several diagnostic characters data to clarify some relationships among red knee
for Brachypelma such as: femur IV lacking inner species and revised their nomenclature, but only
scopula (no dense pad of plumose setae), presence included some species from the ‘red leg emilia group’,
of a distinct scopula on the metatarsus IV, palpal excluding the type species B. emilia, leaving the
bulb narrowly piriform, but with the apex wide and monophyly of the genus untested. Recently, Turner
attenuated, very much compressed and obtuse. et al. (2018) presented a mtDNA gene tree of tarantula
Pocock (1903) also distinguished the plumose setae spiders based on the mitochondrial 16S-tRNA (leu)-
on the prolateral face of leg I trochanter/femur and NDI gene region as an initial hypothesis to clarify
retrolateral face of the palp. These key features have some taxonomic relationships of the subfamily
been supported by subsequent authors as diagnostic Theraphosinae. Their recovered phylogeny in both
for Brachypelma, as also are, no tarsal division by Bayesian and maximum likelihood analyses strongly
strong setae, the male palpal bulb distally wide and flat supported the non-monophyly of both Brachypelma
(commonly mentioned as spoon-shaped), two unequal and Aphonopelma, indicating that neither of these
apophyses on male tibia I, and the spermatheca genera, as currently recognized, are monophyletic.
regularly fused, semi-fused or with two separated This result is congruent with our results based on COI,
wide lobes (Schmidt, 1992a, 1992b; Smith, 1993, 1994; so it is another strong support for the non-monophyly
Pérez Miles et al., 1996; Locht et al., 1999; Gabriel & of Brachypelma. We agree with the results of Turner
Longhorn, 2015; Mendoza & Francke, 2017). Smith et al. (2018) and proposed the ‘red leg group’, which
(1994) adequately explains the taxonomic history contains the type species B. emilia, as Brachypelma
of Brachypelma, which has been complicated and s.s., while the ‘red rump group’ is considered as a
different genus here described as Tliltocatl with B. fossorium are only type I, whereas Brachypelma
the designation of T. vagans as its type species. We and Tliltocatl possess types I and III. Brachypelma
also performed a morphological character-based fossorium has scopula IV divided by a strong band
analysis of the genus Brachypelma, using parsimony of setae, whereas Brachypelma and Tliltocatl have
to test the monophyly of the genus and to reconcile all their scopulae entire. So, based on this evidence,
morphological and molecular characters. Our best and other arguments exposed in our morphological
tree is based on the strict consensus of two most revision of the species above, we are certain that this
parsimonious trees obtained from the parsimony species does not belong in either Brachypelma or
analysis of 103 characters. In this phylogeny, the Tliltocatl, and we transfer it to Stichoplastoris.
‘red leg group’ and the ‘red rump group’ were also Mendoza & Francke (2017) concluded that
recovered as separated genera, giving additional barcode marker COI in Brachypelma proved to be
evidence for the non-monophyly of Brachypelma. sufficient for correct species identification. They
However, morphological data in Theraphosidae tends considered it also as a useful tool in preventing black
to be homoplastic due to similarities, simplicity in market trade and in providing better strategies to
sexual structures and conserved characters (Raven, reintroduce tarantulas into the correct distribution
1985; Goloboff, 1993; Pérez-Miles et al., 1996; Pérez- areas. DNA barcoding is a useful technique that,
Miles, 2000; Bertani, 2001; Bond & Opell, 2002; together with morphology, field observations and
Bond & Hedin, 2006; Hedin & Bond, 2006; West museum collections, allow for better definition and
et al., 2008; Hendrixson & Bond, 2009; Bond et al., delimitation of species (Scotland et al., 2003; Chen
2012; Guadanucci, 2014; Hamilton et al., 2014, et al., 2011; Slowik & Blagoev, 2012; Chan et al.,
2016; Perafán & Pérez-Miles, 2014; Ortiz & Francke, 2014; Hendrixson et al., 2015; Pante et al., 2014).
2016; Fukushima & Bertani, 2017). Despite this However, its use to resolve phylogenies is limited.
shortcoming, both Brachypelma and Tliltocatl have Hamilton et al. (2016) mentioned that the limitations
diagnostic characteristics and synapomorphies that of mtDNA used for phylogeny are gene tree/species
differentiate them from each other, the most evident, tree incongruence and the haploid, non-recombining
in addition to their differences in coloration, are the nature of the molecule, with COI representing only
prolateral inferior keel absent or slightly developed one particular genealogy out of all possible ones in
and restricted to the bulb apical, the spermatheca a genome. These limitations are shown in our ML
baseplate well developed and well sclerotized in tree, with the strong support for the confidence of
Brachypelma; whereas in Tliltocatl the prolateral each genus, but with low resolution on some inner
inferior keel is always present and is widely extended clades in both Brachypelma and Tliltocatl (Fig. 4).
backwards, the spermatheca do not have a baseplate In the case of Brachypelma, the most problematic
or only present a small area slightly sclerotized. is the relationship of B. baumgarteni, B. boehmei
Within the taxonomy of tarantulas, the count of and B. auratum, just as is seen also in Mendoza &
spines in the legs or the presence/absence of them Francke (2017), where all these species were well-
does not usually present great taxonomic relevance supported but the clade that includes them was
due to its variability between individuals of the same collapsed. In our phylogeny, something similar
species. However, our analysis of these characters happens with B. auratum and B. baumgarteni as
showed differences between Brachypelma and sister-species but with low support and with poor
Tliltocatl, as Tliltocatl always presents spinations on resolution in B. boehmei (Fig. 4). For Tliltocatl, the ML
all the patellae, whereas Brachypelma never shows tree shows no resolution in the position of T. verdezi
this characteristic, thus we consider this is a reliable and T. kahlenbergi, both sister of a clade formed by
character to differentiate both genera. Brachypelma T. epicureanum, T. albopilosum, T. sabulosum and
fossorium is a species that is not grouped with either T. vagans. In this clade, T. epicureanum appears as a
Brachypelma or Tliltocatl in both the morphological sister-species of the other three; although the clade,
and molecular phylogenies (Figs 3, 4). This species which shows T. vagans as sister to T. sabulosum
was originally described by Valerio (1980) as and T. albopilosum, has no support (Fig. 4). The
Brachypelma based solely on the shape of the male morphology-based tree has some similarities with the
palpal bulb and female genitalia. However, as is one obtained with ML for COI, showing B. emilia as
shown in our results, we conclude that this species the sister-species of all other Brachypelma, followed
does not belong to Brachypelma or Tliltocatl. Despite by B. klaasi and B. albiceps, which are supported
the similar shape of the genitalia, this species does by a combination of five homoplastic characters but
not possess the diagnostic features of either genus: not by bootstrap values (Fig. 3; char35, 55, 98, 100,
it has a short embolus with different arrangement 101). Three of these characters are related to the
of keels from that observed in Brachypelma or spermatheca, which have similarities because they
Tliltocatl. Also, the urticating setae present in are the only species with divided spermatheca. So, this
is the main reason why the three species are grouped Conservation issues
as sister-species in the morphological phylogeny, As was mentioned by Turner et al. (2018), the non-
unlike the molecular where they are separated (Figs. monophyly of Brachypelma with some species
3, 4). The same occurs in the case of the group formed being transferred to another genus has immediate
by B. baumgarteni and B. hamorii, since these implications for conservation. The main concern for
species are supported by two synapomorphies (char4, those authors lies in the state of conservation that
35) and six homoplastic characters (char6, 10, 13, 79, could be affected after the removal of the ‘red rump’
82, 83); the synapomorphies are the coloration of the species complex from Brachypelma. This is because,
chelicerae and the similar coloration on the tibiae. as was observed, we still know very little about their
Three of the six homoplastic characters are related diversity, distribution, ecological characteristics,
to the male palpal bulb, since both species have a habitat preferences, reproductive success and how
similar shape with the apical keel slightly developed much they are affected by habitat loss and predation
and the prolateral superior keel wide and short (Fig. (Reichling, 2000; Longhorn, 2002; Machkour-M’Rabet
3). In contrast, in the ML tree, B. baumgarteni and et al., 2005, 2007, 2011, 2012, 2015, 2017; Shillington
B. hamorii are genetically distant, with B. hamorii & McEwen, 2006; Dor et al., 2008; Dor & Hénaut,
followed by B. smithi and this is followed by a group 2011, 2012, 2013; Vilchis-Nestor et al., 2013; Hénaut
formed by B. boehmei, B. baumgarteni and B. auratum et al., 2015). Nevertheless, CITES is aware of possible
(Fig. 4). For the genus Tliltocatl, there are differences changes in nomenclature in the taxonomic groups
between both topologies. Where in the morphological included for its regulation and in its resolution Conf.
tree T. albopilosum and T. schroederi are shown as 12.11 revised in the COP17 held in Johannesburg, it
sister-species supported by only three homoplastic is mentioned that ‘whenever a change in the name of
characters and no bootstrap support (char43, 62, 71). a taxon included in the Appendices is proposed, the
All the characters are related to the number of spines Secretariat, in consultation with the Animals or Plants
in patellae II and III and metatarsus III (Fig. 3). The Committee, determine whether this change would
ML tree shows T. schroederi as the sister-species of alter the scope of protection for fauna or flora under
all other Tliltocatl, whereas T. albopilosum is in an the Convention’. This previous statement that in case
unresolved group with T. sabulosum and T. vagans. where the scope of a taxon is redefined, the Animals or
The other species resolution is more similar in Plants Committee shall evaluate whether acceptance
both topologies, with T. verdezi and T. kahlenbergi of the taxonomic change would cause additional species
with no resolution and as sister-species of an inner to be included in the appendices or listed species to
clade formed by T. epicureanum, T. vagans and be deleted from the appendices. Thus, if that is the
T. sabulosum in the morphological tree, and with case, the Depositary Government should be requested
the inclusion of T. albopilosum in the molecular tree to submit a proposal to amend the appendices in
as was shown above (Figs 3, 4). These discrepancies accordance with the recommendation of the Animals
between morphological and genetic phylogenies are or Plants Committee, so that the original intent of the
apparently given by the grouping of some species listing is retained. This means that even if species that
based mainly on the similarity of their reproductive belong to the ‘red rump’ group are transferred from
organs, which are probably convergent. Also, it is Brachypelma into the genus Tliltocatl, these species
possible that the inclusion of other outgroups can will not lose their protective status under CITES
help to resolve better the morphological hypothesis. appendix II. However, this protective clause only
We suggest that future studies must explore the use concerns international trade. It is equally important
of different molecular markers, such as nuclear ones, to know if the taxonomic change of these species will
which are more conservative and could help to resolve affect their protection status in Mexico. There, it is the
the inner relationships of the species. Also, the use Ley General de Vida Silvestre (LGVS) who regulates
of next-generation sequencing methods can help to the sustainable use, conservation and management
resolve phylogenetic problems, evolutionary concerns of native wild animals and plants. It regulates the
or even to help in conservation strategies (Turner protection of species or populations that are at risk
et al., 2018). Further, the Inter Simple Sequence (SEMARNAT, 2000). The LGVS establishes the
Repeats (ISSR) used by Machkour-M’Rabet et al. national policy for wildlife protection and sustainable
(2009) as a molecular marker for the study of genetic use via the SUMA programme and the Mexican
diversity in populations of tarantulas proved to be a Official Standard NOM059-SEMARNAT-2010 (NOM-
useful tool to learn about the intraspecific variation 059) on Mexican species at risk. In addition, the
in these organisms, which can be a valuable resource LGVS regulates the creation of UMAs. To change the
to evaluate the population structure of priority status of the species protected by law, it is necessary
tarantulas such as Brachypelma and Tliltocatl. to submit a petition for evaluation with information
about trade, conservation and population status. be used as a reference for the authorities responsible
Thus, despite the nomenclatural changes proposed for species conservation. Juvenile tarantulas are
above, fortunately all the species formerly known as very difficult or impossible to identify, even to genus.
Brachypelma will remain protected in Mexico as well. Therefore, the regulation of the pet-trade of juvenile
In fact, a particular interest has recently emerged from tarantulas will benefit tremendously by using
the Mexican Government to protect all tarantulas and molecular barcode markers to easily and accurately
regulate their trade. identify them. It is essential to explore other non-
In 2015, the governments of Canada, Mexico and the subtractive economic endeavours, such as tourism,
United States initiated a collaborative project through wildlife observation or sale of captive-bred animals, to
the Commission for Environmental Cooperation provide alternative incomes for local communities and
(CEC) to strengthen the conservation and sustainable to avoid further damage to wildlife populations.
trade of the 16 tarantulas that are included in
appendix II of CITES. The result of this was the
creation of an action plan for the sustainable trade in
ACKNOWLEDGEMENTS
tarantulas. This action plan includes information on
16 priority tarantula species, comprising one from the This publication is a requirement for the PhD graduate
genus Aphonopelma and 15 from Brachypelma s.l. The training programme of Jorge Mendoza at the Posgrado
information was compiled for the species as a group en Ciencias Biológicas of the Universidad Nacional
and included: the impact of trade on conservation Autónoma de MEXICO (UNAM). JM thanks the
and livelihoods, and identification challenges for Instituto de Biología (IBUNAM) and the Posgrado en
CITES’ enforcement. Currently, there are insufficient Ciencias Biológicas of the UNAM for the programme
population data available for Mexican tarantulas, received. JM also thanks the Consejo Nacional de
especially those restricted to small geographical areas Ciencia y Tecnología (CONACYT) for providing
that are particularly vulnerable to overexploitation financial support for his post-graduate studies, as
(Reichling, 2003). Without such information, it would well as the grants required for a research stay at
be very difficult to make sure that the impact of the Muséum National d´Histoire Naturelle in Paris,
exporting adult or sub-adult specimens would not France (MNHNP), the Natural History Museum in
be detrimental to the species survival in the wild, London, United Kingdom (NHM), the Senckenberg
unless there was compelling information available Naturmuseum in Frankfurt, Germany (SNMF) and
to show that the specimens in question were the Museo de Zoología de Costa Rica (MZUCR). We
captive-bred. Preliminary field studies indicate that thank Janet Beccaloni, Christine Rollard, Elise-
B. baumgarteni, B. boehmei and B. hamorii have Anne Leguin, Peter Jäger, Julia Altmann and Gilbert
small zoogeographical ranges and are sensitive to Barrantes for providing access to types and historical
habitat disruption. Therefore, at least these three specimens. Stuart Longhorn and Ray Gabriel are
species should not be considered for direct capture thanked for their assistance and hospitality during
and export until more research has been conducted JM’s research stay in Europe. We thank Virginia
on their viability (CEC, 2017). At sustainable levels of Leon for her support for travel to Europe and Andrea
exploitation, both wildlife and people can benefit from Jiménez for her assistance during molecular work. JM
legal trade. Granting local people an economic stake thanks the members of the laboratory of arachnology
in wildlife management provides the best incentive (CNAN) for their support and assistance on field trips.
for careful stewardship of species and habitats (Carey, JM thanks R. Ramírez, J. Hinojosa, E. Hijmensen and
1999; Dickinson, 2002). E. Goyer for their assistance during field trips. We
Although, DNA barcodes are not good enough to also thank the editor and reviewers for their critical
either capture the dynamic processes of evolution or reading and valuable comments. The specimens were
to reflect the precise phylogenetic relationships within collected under scientific collection permit FAUT-
a given group, they could provide legislators with a 0175, authorized by SEMARNAT for Oscar F. Francke.
framework of data when enacting protection laws. This The present project was supported by MEXBOL
could translate into greater success in prosecuting of CONACYT (project 251085) and the Barcode of
those involved in the illegal pet-trade. It is also Wildlife Project in Mexico, sponsored by Google´s
generally agreed that mtDNA alone is not sufficient Global Impact Awards.
for precise species delimitation. However, it can be
useful for identification if the findings correspond with
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SUPPORTING INFORMATION
Additional Supporting Information may be found in the online version of this article at the publisher's web-site.
Appendix 1. Most characters were codified based on Bertani (2001), West et al. (2008) and Fukushima & Bertani
(2017). New ones or modified characters are indicated, as also are explained the ones based on Brachypelma and
Tliltocatl characteristics.
Appendix 2. Genbank accession codes for tissue samples, deposited in the Laboratorio de Sistemática Molecular
(Zoología) at the Instituto de Biología, UNAM, Mexico City, from which DNA was extracted and sequenced for
phylogenetic analyses of 8 species in the genus Brachypelma Simon 1891 and seven species in the Tliltocatl genus.
P. cambridgei, E. campestratus, Lasiodora parahybana and Xenesthis immanis was used from GenBank.

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Zoological Journal of the Linnean Society, 2019, XX, 1–66. With 46 figures.

INTRODUCTION 2018; Mendoza, 2014a). According to the World Spider

RESULTS The second clade includes a group in the ‘red rump’

(CNAN-Ar003413); 1 ♂, Mpio. Arcelia, Presa Vicente Diagnosis

G. Contreras, D. Ortiz, D. Barrales (CNAN-Ar003658); CNAN-Ar010284); 1♀, Mpio. La Huacana, 1 km SE of

Brachypelma boehmei is identified by possesing

162) considered this as a junior synonym of CNAN-Ar010275); 1 ♂, 1 ♀, Mpio. Acapulco, El

Identification key for species of Brachypelma Simon 1891

Holotype and paratype of Brachypelma fossorium:

Brachypelma fossorium: Schmidt (1992: 10, figs 9–12)

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