TY - JOUR
AU - Francke,, Oscar
AB - Abstract 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. distribution, geography, genus revision, mitochondrial DNA, new genera, phylogenetic nomenclature, phylogenetics INTRODUCTION Tarantulas (family Theraphosidae) are the largest and heaviest spiders of the world. They can inhabit almost every terrestrial ecosystem, with the exception of polar areas, but they are mainly found in tropical, subtropical, semi-arid and arid regions around the world. They can be found from sea level to 4000 m elevation and some even can be found inside caves down to –800 m deep (Bond et al., 2012; Lüddecke et al., 2018; Mendoza, 2014a). According to the World Spider Catalog (2018), there are currently 1001 described species. Many of them are colourful in appearance and docile in captivity, making them attractive pets for collectors (West, 2005). Some of the most appreciated by enthusiasts are those of the genus Brachypelma due to their longevity, size and attractive coloration. Because of the high demand for specimens of Brachypelma smithi (Pickard-Cambridge, 1897) in the late 1980s, 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 Species (CITES). To prevent the same problem from arising with other Brachypelma species, the inclusion of the entire genus in CITES appendix II was proposed (Smith, 1994; West, 2005). Later, the World Conservation Monitoring Centre (1996) evaluated and included B. smithi as a near-threatened species in the International Union for Conservation of Nature (IUCN) Red List of Threatened Species. In recent years, the Commission for Environmental Cooperation has been working with authorities from Mexico, United States and Canada to generate and implement action plans to promote the legal, sustainable and traceable trade in selected North American species that are listed in appendix II of CITES. These actions were developed based on the information compiled from consultation with stakeholders. Sixteen tarantula species, comprising one from the genus Aphonopelma and 15 from the genus Brachypelma, were selected as ‘priority tarantula species’ and are the subject of one of the action plans (CEC, 2017). The genus Brachypelma was established by Eugene Simon in 1891, based on material that he believed to be Mygale emiliaWhite, 1856, which was deposited in the Natural History Museum of Paris (Smith, 1994), and subsequently it was designated as the type material (Rudloff, 2003). This species was originally listed from Panama, but this location is erroneous as its currently known natural distribution is north-western Mexico, specifically in the Pacific Coast region (Smith, 1994; Locht et al., 1999; Gabriel & Longhorn, 2015). Pickard-Cambridge (1897) considered Brachypelma as a junior synonym of Eurypelma and described E. smithi. This was then accepted and followed by Simon (1903). It was Pocock (1903) who divided the genus Eurypelma into a number of genera and restored the validity of Brachypelma. In his redescription of the genus, he emphasized the presence of plumose hairs on the palp trochanter and on the trochanter and femur of leg I, and he also included B. albicepsPocock 1903, B. smithi and B. vagans (Ausserer 1875) in Brachypelma. Roewer (1942) then again suspended Brachypelma and restored the genus Eurypelma. This change remained in place for several decades until Valerio (1980) re-established Brachypelma and included four additional species [B. albopilosum Valerio, 1980, B. angustum Valerio, 1980, B. fossorium Valerio, 1980 and B. mesomelas (O. P.-Cambridge, 1892)], although only B. albopilosum and B. fossorium continue currently in Brachypelma. Then, Raven (1985) in his revision of the Mygalomorphae concluded that Brachypelma was a junior synonym of the older genus Euathlus Ausserer, 1875, based on the study of a single specimen from the Koch collection in The Natural History Museum, London (NHM). Smith (1994) mentioned that an amateur arachnologist named Frank Fritzlen presented an old document in 1991 based on scientific descriptions and drawings, which could indicate that a mistake may have taken place in the collection of the NHM. Smith (1994) also described how that error could have been made in 1905 when the Koch collection was labelled, after it was acquired by the NHM. It was Schmidt (1992a) who removed it from synonymy of Euathlus and restored Brachypelma (see Brachypelma andrewi Schmidt, 1992 section for further details). Since then, nine more species have been described: B. annitha Tesmoingt, Cléton & Verdez, 1997, B. auratum Schmidt, 1992, B. baumgarteni Smith, 1993, B. boehmei Schmidt & Klaas, 1993, B. klaasi Schmidt & Krausse, 1994, B. hamorii Tesmoingt et al., 1997, B. schroederi Rudloff, 2003, B. verdezi Schmidt, 2003 and B. kahlenbergi Rudloff, 2008. Three species originally described as Eurypelma have been transferred to Brachypelma: B. aureoceps (Chamberlin, 1917), B. epicureanum (Chamberlin, 1925) and B. sabulosum (Pickard-Cambridge, 1897). There was an attempt to include B. klaasi and B. albiceps (formerly described as Brachypelmides ruhnaui Schmidt, 1997) in the genus Brachypelmides (Schmidt & Krausse, 1994) based on the presumptive presence of a small pad of slightly plumose setae on the femur IV and the sharply tapered embolous of the males. However, after examination of B. klaasi types, Smith (1994) concluded that the species was a Brachypelma, but an ‘extreme form’ of the genus, considering Brachypelmides as a junior synonym of Brachypelma. His conclusion was followed by Locht et al. (1999) who examined B. klaasi and B. ruhnaui and did not find any plumose setae on femur IV, and concluded also that, although the embolous of both species is sharper and more tapered, it does not differ significantly from the one in other Brachypelma. It was Locht et al. (2005) who concluded that Brachypelmides ruhnaui is a junior synonym of Brachypelma albiceps after revision of topotype material and specimens deposited in scientific collections. Recently, based on molecular characters, Mendoza & Francke (2017) considered that B. annitha is a junior synonym of B. smithi. The following year, Turner et al. (2018) presented a mtDNA gene tree of some tarantula spiders in which Brachypelma was recovered as biphyletic. Currently, there are 18 described species of Brachypelma ranging from Mexico to Costa Rica (Smith, 1994; Locht et al., 1999; Gabriel & Longhorn, 2015; WSC, 2018). Of these, 13 occur in Mexico, where 12 are endemic. The most northerly species is Brachypelma emilia (White, 1856) found west of the Sierra Madre Occidental mountain range in the states of Sonora, Sinaloa, Durango and Nayarit. This highly colourful species is a fossorial burrower whose habitat ranges widely from the drier coastal thorn forests and savannahs, through the palm transition forests and wetter inland tropical deciduous forests, up into the higher and cooler elevations of the oak forests (West, 2005). The southernmost species was Brachypelma embrithes (Chamberlin, 1925), which is found in Panama, but Gabriel & Longhorn (2015) transferred this species to the genus SericopelmaAusserer, 1875. Currently, the southernmost species in the genus is Brachypelma albopilosum from Costa Rica. Although correct species identification is a fundamental component of many biological investigations, taxonomic expertise is shrinking and at the same time overwhelmed by demand. The correct identification of a species based on morphological characters can sometimes take too long or can be impossible for certain life stages, like immature spiders or even for members of one sex (Greenstone et al., 2005; Blagoev et al., 2009). There are also some problems in the identification of species in groups with similar patterns of convergence or morphological conservatism (Hebert et al., 2003; Locke et al., 2010; Niemiller et al., 2011; Hamilton et al., 2014, 2016). Theraphosid spiders are known by having simple genitalia and tend to show relatively low interspecific and relatively high intraspecific variation, which have revealed widespread patterns of homoplasy among traditional taxonomic characters (Raven, 1985; Goloboff, 1993; Pérez-Miles et al., 1996; Pérez-Miles, 2000; Bertani, 2001; Bond & Opell, 2002; Bond & Hedin, 2006; Hedin & Bond, 2006; West et al., 2008; Hendrixson & Bond, 2009; Bond et al., 2012; Guadanucci, 2014; Hamilton et al., 2014, 2016; Perafán & Pérez-Miles, 2014; Ortiz & Francke, 2016; Fukushima & Bertani, 2017). An alternative for a sustainable identification is the construction of systems that employ DNA sequences as taxon barcodes (Sun et al., 2012). Molecular markers, such as COI, may provide species boundary information in certain taxonomic groups and consequently have the potential to be a rapid and efficient means to delineate and identify species (Hebert et al., 2003; Barrett & Hebert, 2005; Arnedo & Fernandez, 2007; Chen et al., 2011; Kuntner & Agnarsson, 2011; Hamilton et al., 2014; Blagoev et al., 2016). However, molecular identification is not free of problems and may have restrictions and inconsistencies. So, the use of molecular data, together with morphology, is recommended, along with collection data and field observations to allow for a better species delimitation and proposal of phylogenetic relationships (Scotland et al., 2003; Will & Rubinoff, 2004; Prendini, 2005; Bond & Stockman, 2008; Slowik & Blagoev, 2012; Hendrixson et al., 2015; Ortiz & Francke, 2016). In the particular case of tarantulas, there are a limited number of studies that have used molecular information for species delimitation, and some of them found moderate to deep inconsistencies in the morphology-based taxonomy due to homoplasy in some characters (Hamilton et al., 2011, 2014, 2016; Hendrixson et al., 2013, 2015; Wilson et al., 2013; Graham et al., 2015; Montes de Oca et al., 2015; Ortiz & Francke, 2016). To date, no taxonomic revision has been made of the genus Brachypelma,despite its popularity in the pet trade and being a priority group for conservation protected by international conventions such as CITES or the General Law of Wildlife (LGVS) in Mexico. Also, although geographical ranges have been published for Brachypelma species of Mexico (Baerg, 1958; Valerio, 1980; Smith, 1994; West, 1996, 2005; Locht et al., 1999; Reichling, 2001, 2003; Rojo, 2004; Arisqueta-Chablé et al., 2009; Shaw et al., 2011; Hijmensen, 2012; Longhorn, 2014; Mendoza & Francke, 2017), these ranges are general in scope or incomplete (Fig. 1). Furthermore, studies to determine the susceptibility, exact zoogeographical range and/or genome for each priority tarantula species have not been conducted (Garcia, 2016). The geographic distribution of Brachypelma shows a clear difference between the ‘red leg’ and ‘red rump’ species, the first ones mostly distributed through the Mexican Pacific Coast and central Mexico and the second ones distributed along the south-west of Mexico from the Gulf of Mexico to the Pacific Coast in southern Guerrero and continuing southward to northern Costa Rica (Fig. 2). Most Brachypelma studies have focused on a few taxa, as those of population interactions, ethology, morphological variation and genetic diversity in Brachypelma vagans (Ausserer, 1875) (Reichling, 2000; Longhorn, 2002; Machkour-M’Rabet et al., 2005, 2007, 2011, 2012, 2015, 2017; Shillington & McEwen, 2006; Dor et al., 2008; Dor & Hénaut, 2011, 2012, 2013; Vilchis-Nestor et al., 2013; Hénaut et al., 2015), from courtship and habitat preference of Brachypelma klaasi (Schmidt & Krause, 1994) (Yáñez et al., 1999; Yáñez & Flotaer, 2000) and only a few studies have tested the utility of molecular markers for species delimitation or identification of the priority tarantula species (Longhorn et al., 2007; Petersen et al., 2007; Machkour-M’Rabet et al., 2009; Mendoza & Francke, 2017; Turner et al., 2018). Figure 1. Open in new tabDownload slide 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. Figure 1. Open in new tabDownload slide 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. Figure 2. Open in new tabDownload slide 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. Figure 2. Open in new tabDownload slide 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. In Mexico, wild populations of Brachypelma tarantulas are in decline, due to habitat loss, because people often kill them when encountered in the wild and because large numbers of some of the more colourful species are collected, particularly from the Pacific Coast, to supply the national pet market. Field-caught tarantulas are either sold in traditional markets (with little or no law enforcement) or exported illegally for the commercial pet trade (García, 2016; CEC, 2017). The sustainable use, conservation and management of native tarantulas in Mexico is regulated via the General Law of Wildlife (LGVS for its Spanish acronym) and its by-laws that allow the creation of the named Management Units for the Conservation of Wildlife (UMA for its Spanish acronym). The UMA refers to the registered facilities that operate in accordance with an approved management plan. These are intensive and extensive breeding sites where wild flora, fauna and fungi are reproduced and propagated, and products and by-products destined to different types of use are generated. Their general objective is the conservation of natural habitat, populations and specimens of wild species. They function as centers of breeding stock, as germplasm banks, an alternative ex situ conservation for reproduction of key species or species that are in any category of risk. For environmental education, research and production of species, parts and derivatives of wildlife that can be incorporated into the different circuits of the legal market for commercialization (SEMARNAT, 2000). Under the UMA programme, qualified persons may present a request to collect a limited number of wild tarantulas to keep and breed in captivity. The resulting offspring may then be sold domestically or exported, and may also be used for reintroduction programmes. Few hobbyists, breeders or academics would be experienced in identifing each species of priority tarantula on sight. To ascertain that the objectives of protecting and conserving tarantulas, supported by international policies and those of Mexico, have better results, it is important to have an adequate description of tarantula species, to allow for the prioritization of certain species for conservation. Thus, the development of morphological guides and molecular databases allowing correct identification is of the utmost importance. In addition, it is necessary to delimit their natural distribution areas and habitat preferences in order to develop better strategies for conservation of their populations, which will also allow selection of potential sites appropriate for in situ repopulation programmes. Initially, it is the aim of this work to revise the genus Brachypelma Simon, 1891 and to test its monophyly by the use of morphological and molecular characters. Some species were originally described based solely on one sex or even using only exoskeletons as type specimens, so we reviewed and illustrated specimens of both sexes. Some new characters are introduced as a result of our comparative morphological studies of these species and their morphological variability. The redescription of Brachypelma species and the relevant taxonomic changes are made based on our results. Distribution maps for all known species are presented and general descriptions of their habitat and life cycle are provided. MATERIAL AND METHODS The general descriptive format used in the present study follows Mendoza & Francke (2017). All measurements are in millimetres and were taken using an ocular micrometre on a stereomicroscope Nikon SMZ645 for smaller structures, and a digital calliper with an error of 0.1 mm for larger ones. Leg and palp measurements were taken along the dorsal central axis of the left side. Description of tarsal scopulae follows Pérez-Miles (1994). Male palpal bulb keels and tegular apophysis terminology follows Bertani (2000). Spermatheca shape follows the general description format used with theraphosids (Pérez-Miles, 1989; Bertani, 2001; Mendoza & Francke, 2017). Description of stridulating setae follows Galleti & Guadanucci (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 to these places and to avoid the illegal collection of specimens given that these species are protected by law. Photographs (Figs 5–7, 9–10, 11C, 12, 14–16, 18–19, 20B, 21, 23–25, 27, 29–31, 33, 39C–D, 40) were taken with a Nikon Coolpix S10 VR digital camera coupled to a stereomicroscope. Descriptions of colours use the standard names of the 267 Colour Centroids of the NBS/IBCC Colour System (Mundie, 1995). The colours depicted herein are included to promote some standardization in describing coloration of living animals. To avoid differences of perception in coloration because of the calibration of the monitor, we extracted the RGB code and pantones colours from the photographs using Photoshop CS Live. A digital photograph was opened and the eyedropper tool clicked over the image to obtain the RGB colour code in the Set foreground colour and cross-checked in the colour libraries for the Pantone solid coat. Both habitat 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 background were taken in the laboratory where general illumination was provided by one fluorescent 30-W light bulb held approximately 20 cm from the specimen. Once the RGB code was obtained, it was possible to infer the real colour of the specimen using the RGB code of Colour Centroids. The range of each colour centroid as perceived by the human eye is wide enough to account for errors of observation. Abbreviations used in the text are as follows: Ocular patterns – ALE, anterior lateral eyes; AME, anterior median eyes; PLE, posterior lateral eyes; PME, posterior median eyes. Morphology – legs and palpi: d, dorsal; p, prolateral; r, retrolateral; v, ventral; Rap, retrolateral tibial apophysis; Pap, prolateral tibial apophysis. Palpal bulbs: AK, apical keel; PI, prolateral inferior keel; PS, prolateral superior keel; TA: tegular apophysis. Spermatheca: Bp, spermathecal baseplate. Spinnerets: PLS, posterior lateral spinnerets; PMS, posterior median spinnerets. In species synonymies, we follow the World Spider Catalog (2018): D = Described, f = female, m = male, T = Transferred. Institutions: CNAN = Colección Nacional de Arácnidos, MEXICO DF, MNHNP = Museum National d´Histoire Naturelle, Paris, NHM = Natural History Museum, London, PROFEPA (acronym in Spanish) = Federal Environmental Protection Agency, SNMF = Senckenberg Naturmuseum, Frankfurt, UCR = Museo de Zoología Universidad de Costa Rica; UNAM = Universidad Nacional Autónoma de MEXICO. Morphological protocol Taxon sampling The cladistic analysis was based on 24 taxa. The ingroup included 16 species of Brachypelma. The outgroups were selected based on the cladistic analyses of subfamily Theraphosinae made by Pérez-Miles et al. (1996), Pérez-Miles (2000), Bertani (2001) and Perafán & Pérez-Miles (2014). The number of external groups used was eight, belonging to the following taxa: Acanthoscurria geniculata (C. L. Koch, 1841), Aphonopelma seemanni (F. O. P.-Cambridge, 1897), Bonnetina cyaneifemur Vol, 2000, Megaphobema mesomelas (O. Pickard-Cambridge, 1892), Psalmopoeus victori Mendoza, 2014b, Sericopelma melanotarsum Valerio, 1980, Theraphosa stirmi Rudloff & Weinmann, 2010 and Xenesthis immanis (Ausserer, 1875). The trees were rooted on P. victori, selected because it belongs to subfamily Psalmopoeinae Schmidt, 2010 (formerly Aviculariinae), making it an appropriate outgroup for Theraphosinae. Character matrix and cladistics Sereno (2007) mentioned that characters are simple characteristics expressed as independent variables and that character states are the mutually exclusive conditions of a character. Together, the characters and character states make up character statements. Based on this definition, the coding of characters was carried out taking phylogenetic works carried out in other genera of the family as a starting point (Bertani, 2001; Fukushima, 2005; Pérez-Miles, 2000; Pérez-Miles & Locht, 2003; West et al., 2008; West & Nunn, 2010; Fukushima & Bertani, 2017). Intraspecific variation of each Brachypelma species was determined by reviewing between five and ten specimens of each sex for each species, with the exception of Central American species, because it was not possible to obtain additional samples to those already deposited in scientific collections. The analysed characters were only of adult specimens of both sexes, because only mature individuals present diagnostic characters and other morphological characters that allow formulation of a homology hypothesis. The character matrix comprised 103 characters of adult morphology scored for 24 taxa; eight characters were non-informative (Table 1; Supporting Information, Appendix S1). Multistate characters were optimized as non-additive (Fitch, 1971). In the absence of evidence that would support the order in the multistate characters, the heuristic searches were conducted in TNT v.1.1 (Goloboff et al., 2008). The search method applied 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 fused suboptimal trees to prevent clogging); memory with a maximum of 30 000 trees was retained. The analysis was performed using equal weighting, the length (L) of the optimal trees, consistency index (Ci) and retention index (Ri) are reported. Optimization of ambiguous characters was carried out using ACCTRAN (Accelerated Transformation), in order to disadvantage parallelisms and promote regressions. According to Agnarsson & Miller (2008), there are no grounds for preferring any algorithm to solve ambiguous optimizations, so the use of ACCTRAN is somewhat arbitrary and is only used for the sole purpose of presenting a hypothesis about a resolution of these ambiguities. The relative support for each node on the preferred hypothesis was calculated with bootstrap resampling (Felsenstein, 1985). Bootstrap support was estimated with heuristic searches of 1000 pseudoreplicates, with P = 50. Cladograms obtained by TNT were exported in Tree file, to be properly edited in WinClada Asado v.1.7 (Nixon, 1999–2004), mapping all the characters on the consensus tree, and were subsequently edited with Adobe Illustrator CS5. Table 1. Characters matrix used in cladistic analysis of Brachypelma s.l. and s.s. (–) inapplicable (?) unknown; treated as missing data in the analysis. Open in new tab Table 1. Characters matrix used in cladistic analysis of Brachypelma s.l. and s.s. (–) inapplicable (?) unknown; treated as missing data in the analysis. Open in new tab Molecular protocol Taxa Specimens were collected throughout the known distribution area of the genus Brachypelma with special attention to the type localities (where possible). Material was fixed in 80% ethanol. The third leg on the right side of each spider was stored in 96% ethanol at –20 °C. Tissue samples of 69 specimens were used for DNA extraction, representing 16 species of Brachypelma. Five tissue samples each of Brachypelma albiceps Pocock, 1903, B. auratum Schmidt, 1992b, B. baumgarteni Smith, 1993, B. emilia (White, 1856), B. epicureanum (Chamberlin, 1925), B. hamorii Tesmoingt, Cléton & Verdez, 1997, B. kahlenbergi Rudloff, 2008, B. klaasi (Schmidt & Krause, 1994), B. schroederi Rudloff, 2003 and B. smithi (F. O.-Pickard Cambridge, 1897); four each of B. boehmei Schmidt & Klaas, 1993 and B. verdezi Schmidt, 2003; three each of B. sabulosum (F. O.-Pickard Cambridge, 1897) and B. vagans (Ausserer, 1875) and one each of B. albopilosum Valerio, 1980, B. fossorium Valerio, 1980, Aphonopelma seemanni (F. O.-Pickard Cambridge, 1897), Megaphobema mesomelas (O. Pickard-Cambridge, 1892) and Sericopelma melanotarsum Valerio, 1980 were used in this study. Additionally, four sequences were retrieved from GenBank to use as outgroups for phylogenetic analyses: Psalmopoeus cambridgei Pocock, 1895 (JQ412455.1), Eupalaestrus campestratus (Simon, 1891) (JQ412446.1) (both from Briscoe et al., 2013), Lasiodora parahybana Mello-Leitão, 1917 (JN018128.1) (from Arabi et al., 2012) and Xenesthis immanis (Ausserer, 1875) (MG273518) (from Lüddecke et al., 2017). We obtained sequences of mitochondrial COI from 69 samples. Vouchers were deposited in CNAN and assigned a unique number (CNAN-Ar00xxxx). All sequences were submitted to GenBank. Accession numbers and specimen information are given in Supporting Information, Appendix S2. DNA protocols DNA isolation, PCR amplification and sequencing were performed at the Laboratorio de Sistemática Molecular, Instituto de Biología, UNAM. Muscle tissue was extracted from the leg by removing ~20 mg of tissue. Genomic DNA was extracted using the Qiagen DNeasy Tissue Kit, following the protocol of the manufacturer. The concentration of the extracted DNA was quantified with a spectrophotometer (Nanodrop 2000 Techno Scientific) or visualized via agarose gel electrophoresis. DNA amplification was performed using the polymerase chain reaction (PCR) for the mtDNA barcoding gene region COI. A single set of primers was used: LCO 1490: 5’-GGTCAACAAATCATAAAGATATTGG-3’, together with HCO 2198 5’-TAAACTTCAGGGTGACCAAAAAATCA-3’ (designed by Folmer et al., 1994). This primer set amplified a 710-bp region of the mitochondrial cytochrome oxidase subunit 1 gene. PCR reaction (100.8 µL) contained 48 µL 10× PCR-buffer, 24 µL MgCl2, 7.68 µL of forward and reverse primer each, 9.6 µL dNTP’s and 3.84 µL Taq polymerase, using 1 µL of the DNA template for each sample. PCR program for COI followed initial denaturation at 94 °C for 2 min; 30 cycles of denaturation at 94 °C for 1 min, annealing at 48 °C for 45 s, elongation at 72 °C for 2:30 min; followed by 7 min of final elongation at 72 °C. LCO 1490 and HCO 2198 primers used for single-stranded sequencing. The accuracy of sequences was verified by independently amplifying and sequencing the complementary strands of all fragments. Primer sequences were removed and complementary strands of DNA assembled into consensus sequences, edited and checked for quality using GENEIOUS R8 (Kearse et al., 2012). If complementary strands disagreed (besides minor mismatches), the sample was amplified and sequenced again to resolve the discrepancies. DNA sequence alignment and phylogenetic analysis Static alignments of COI gene fragments were generated with MAFFT online v.7 (Katoh et al., 2002, 2005). The G-INS-I strategy, which performs a global alignment based on an FFT approximation, was selected (Katoh et al., 2002). This method is suitable for large datasets comprising sequences with relatively limited variation in length, i.e. few short gaps (Katoh et al., 2005). The scoring matrix for nucleotide sequences was set to 1/PAM K = 2, gap opening penalty to 1.53 and offset value to 0. We carried out maximum likelihood inference using 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 GTR+gamma+Pinvar model of nucleotid evolution and Bootstrap support obtained by running 1000 pseudo replicates. The relative supports are showed for each clade on the best tree. RESULTS Phylogenetic analysis Morphological analysis of 103 characters finds two most parsimonious trees (MPTs) (L = 433, Ci = 0.443, Ri = 0.541). The genus Brachypelma recovers as non-monophyletic, being separated into two clades. The only difference between the two MPTs is the position of Tliltocatl verdezi as a sister-species of T. kahlenbergi, or as a sister-species to a small clade formed of T. kahlenbergi, T. epicureanum, T. vagans and T. sabulosum. The strict consensus tree shows the same topology as MPTs, except for the collapse of the internal clade that relates to T. verdezi and T. kahlenbergi due to the change of the position of T. verdezi (Fig. 3). The first clade includes a group in the ‘red leg’ Brachypelma (Brachypelma s.s.) that includes the type species of the genus B. emilia; although this group is not well supported (P = 15), the strict consensus tree shows that Brachypelma has four synapomorphic characters: tibiae colour pattern completely orange (char35), embolus apical region slightly broad (char78), apical keel developed and slightly extended to backwards (char89) and spermatheca baseplate divided and narrower than base width (char100). The other three characters that support the Brachypelma s.s. clade are homoplastic but can be used in the diagnosis of the genus: fovea shallow (char80), embolus retrolateral curvature only in apex (char75) and embolus apical shape very flattened concave/convex appearance with neck in the base (char79) (Fig. 3). Figure 3. Open in new tabDownload slide 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. Figure 3. Open in new tabDownload slide 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. The second clade includes a group in the ‘red rump’ Brachypelma, which is formally described here as the new genus Tliltocatl, even though this group is not well supported (P = 11). A small clade formed by Aphonopelma seemanni, Xenesthis immanis and Sericopelma melanotarsum appears as its sister-group. The strict consensus tree shows that Tliltocatl possesses one synapomorphy that is the embolus prolateral inferior keel longitudinal and larger than the prolateral superior keel (char86). The other nine characters that support the Tliltocatl clade are homoplastic but can be used in the diagnosis of the new genus: fovea in males and females procurved (char6), fovea in males and females pit-like (char7), distal scopula of metatarsus III with 75% of coverage relative to its length (char13), femur I prolateral face setae claviform (char29), male femur III slightly incrassate (char33), patella I with one spine (char60), embolus curved to retrolateral (char75), embolus apical region broad (char78), and embolus apical shape very flattened concave/convex appearance with neck in the base (char79) (Fig. 3). In the case of the nominal species ‘Brachypelma’ fossorium, it is shown that it does not belong to either Brachypelma or Tliltocatl, although in the strict consensus tree it appears to be closely related to Tliltocatl. ‘Brachypelma’ fossorium lacks important features present in Tliltocatl such as urticating setae type III, prolateral inferior keel longer than prolateral superior keel, femur III incrassate, percentage coverage of distal scopula of metatarsus III is not more than 50%. All these differences are enough to propose the transference of B. fossorium to another genus (see below). Maximum likelihood inference analysis of the COI sequences also strongly supports the non-monophyly of Brachypelma and is congruent with the analysis of morphological characters. The final ML optimization score is –5527.883771. The recovered topology shows two clades: the first one formed by species of Brachypelma ‘red leg complex’ that includes the type species of the genus B. emilia (Brachypelma s.s.) and a second one formed by species in the ‘red rump complex’ (here transferred formally to the new genus Tliltocatl; Fig. 4). The Brachypelma s.s. clade has a bootstrap support value of 78, it is formed by eight species with high bootstrap support. Sericopelma melanotarsum, Aphonopelma seemanni, Xenesthis immanis and Brachypelma fossorium are shown to be more closely related to the new genus Tliltocatl. The clade formed by ‘red rump species’ of Tliltocatl has a bootstrap support of 92 and is formed by seven species with relatively high support, but the internal relationships are unclear. Tliltocatl schroederi is shown to be the sister-species of the rest, but there is no support for the position of T. verdezi and T. kahlenbergi. An internal node follows with T. epicureanum as sister to T. sabulosum, T. albopilosum and T. vagans. However, no resolution is shown for interspecific relationships for the three last species. ‘Brachypelma’ fossorium is clearly shown to be excluded from either Brachypelma or Tliltocatl, being closer to Sericopelma and Aphonopelma (Fig. 4); just as in the morphological analysis, differences are enough to transfer ‘B.’ fossorium to another genus as we propose in the taxonomical section of this paper. Figure 4. Open in new tabDownload slide 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. Figure 4. Open in new tabDownload slide 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. Despite the differences between the parsimony and maximum likelihood analyses, both topologies indicate that Brachypelma s.l., as currently recognized, is not monophyletic, including two highly divergent lineages. The species of both Brachypelma s.s. and Tliltocatl can be identified with high accuracy using COI as a barcode (probably due to the relatively high mtDNA divergence between species). In both analyses inner relationships of some of these species require further resolution, and in the case of morphology, the homoplasy in some of the characters can lead to not strongly supported topologies. On the other hand, the molecular phylogenies based on mtDNA have limitations as gene tree/species tree incongruence, with COI representing only one particular genealogy out of all possible in the genome. Taxonomy Family Theraphosidae Thorell, 1869 Subfamily Theraphosinae Thorell, 1870 Brachypelma Simon, 1891: 338 (Figs 1–38) Type species: Mygale emiliaWhite, 1856: 183 (White, 1857: 406; Simon, 1892: 168; Pocock, 1903: 103; Petrunkevich, 1939: 191; Schmidt, 1986: 49, 1991: 11; Smith, 1986: 50, 1991: 17; Fritzlen, 1991: 14; Schmidt, 1993: 82; Schmidt & Klaas, 1993: 7; Smith, 1993: 14, 1994: 156; Pérez-Miles et al., 1996: 46; Tesmoingt, Cleton & Verdez, 1997: 8; Vol, 2000: 1; Locht et al., 2005: 108; Gabriel & Longhorn, 2015: 85; Mendoza & Francke, 2017: 161; Turner et al., 2018: 11) Eurypelma C. L. Koch, 1850: 73. Type species: Aranea avicularia Linnaeus, 1758 (Simon, 1903: 936; Roewer, 1942: 238). First synonymized by Pickard-Cambridge (1897: 21). Valerio (1980: 268) removed it from synonymy. EuathlusAusserer, 1875: 188. Type species: E. truculentus L. Koch in Ausserer, 1875. First synonymized by Raven (1985: 182). Schmidt (1992: 9) removed it from synonymy. BrachypelmidesSchmidt & Krause, 1994: 7. Type species: B. klaasi Schmidt & Krausse, 1994 (Schmidt, 1997a: 205, 1997b: 19; Locht et al., 1999: 196; Vol, 1999: 11; Schmidt, 2003: 137, 2004: 4). Smith (1994: 159) considered it as junior synonym of Brachypelma. Emended diagnosis The genus Brachypelma can be distinguished from all other known theraphosinae genera (except the new genus Tliltocatl described below) by the following combination of characters: (1) having just claviform stridulating setae on the prolateral face of leg I trochanter/femur and on the retrolateral face trochanter of the palp; (2) both sexes possess always urticating setae types I and III; type III are located in the dorsoposterior area and type I surrounding these; (3) the male palpal bulb distally wide and flattened (spoon-shaped) and presents prolateral superior and apical keels (Figs 6, 10, 15, 19, 24, 27A–D, 30, 33A–D) and can have a small or reduced prolateral inferior keel, except in B. albiceps, B. baumgarteni, B. emilia and B. klaasi; the apical keel can extend slightly (Fig. 6) or widely (Fig. 10) backwards; (4) females can have a divided spermatheca with each lobe as large as wide [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, 33E–H), apically narrowed (present in B. auratum, B. baumgarteni, B. boehmei, B. emilia, B. hamorii and B. smithi); (5) having a spermatheca baseplate divided and well sclerotized; (6) both sexes lack a plumose pad of setae on leg IV femur; and (7) all tarsi scopulae are undivided. Note: Stridulation can be defined as the process of sound production by friction of one rigid body part (the scraper) across a second part (the comb) (Uetz & Stratton, 1982; Vol, 2002). It plays a role in different contexts, such as in intraspecific (reproduction) and interspecific (defence) communication (Marshall et al., 1995). It was Pocock (1903) who first mentioned the plumose setae on the trochanter and femur I as a key feature for Brachypelma recognition. Since then, presence of plumose setae has been used as a diagnostic character for Brachypelma (Smith, 1994; Pérez-Miles et al., 1996; Gabriel & Longhorn, 2015; Mendoza & Francke, 2017). This feature has also been recognized as stridulatory setae by Galleti & Guadanucci (2018) and identified by them as claviform stridulating setae, so we here also consider them as stridulating. Most of the stridulatory organs in Theraphosinae are positioned on the first appendages and consist of plumose or claviform hairs found between the coxae and/or trochanters of the palps and first legs. The function of stridulation is not well understood for theraphosid spiders. The different morphologies and positions of the stridulatory apparatus amongst the theraphosid subfamilies suggest that they are not homologous. Presumably, these organs evolved independently several times in theraphosids (Bertani et al., 2008). In Brachypelma, claviform stridulating setae are used as warning when tarantulas are disturbed, but are also used during courtship to give some signals to the female (Mendoza, personal observation). Species included: Brachypelma albiceps Pocock, 1903, Brachypelma auratum Schmidt, 1992, Brachypelma baumgarteni Smith, 1993, Brachypelma boehmei Schmidt & Klaas, 1993, Brachypelma emilia (White, 1856), Brachypelma hamorii Tesmoingt, Cleton & Verdez, 1997, Brachypelma klaasi Schmidt & Krause, 1994, Brachypelma smithi (F. O. Pickard-Cambridge, 1897). Distribution and natural history The genus is endemic to Mexico and is distributed from Sonora in the north to Guerrero in the south, in the biogeographical region of the Pacific Coast, with a couple of species (B. auratum and B. albiceps) distributed in parts of Central Mexico following the biogeographical region of the Balsas Depression (Fig. 38). It is important to notice that the type of Brachypelma is B. emilia, which was originally described as a species from Panama (White, 1856). However, as was mentioned by Gabriel & Longhorn (2015) this locality could be an error, because the original collector of the species, Berthold Seemann, joined a ship in Panama and travelled north along the Pacific coast, docking in Mexico at Nayarit and Sinaloa, both within the area of distribution for B. emilia. The genus Brachypelma has a natural distribution in the tropical region of the Mexican Pacific Coast and can be found mainly in tropical deciduous forest, which is strongly influenced by dry and rainy seasons, although some populations can also be found in thorn scrub or oak forest (Smith, 1994; West, 2005; Mendoza, personal observation). All species are located between 10 and 1600 m elevation, although rarely some specimens of B. albiceps can be found as high as 1700 m (Mendoza, personal observation). Brachypelma species collected during field work were found mainly in burrows on hillsides. Some burrows were made by the tarantula, but others were clearly adapted by the spider from previous burrows. There are some specimens that made burrows under large, flat rocks or between tree roots, sometimes in open fields, but never far from taller vegetation (Locht et al., 1999). Unlike many other Theraphosinae species, the burrow entrances have no tell-tale trace of silk, indicating inhabitation of a theraphosid or spider, so there never is an indication whether a spider will found while digging the burrow until the end is excavated (West, 2005). Brachypelma albiceps Pocock, 1903 (Figs 2, 5–8, 38) Figure 5. Open in new tabDownload slide 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). Figure 5. Open in new tabDownload slide 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). Figure 6. Open in new tabDownload slide Brachypelma albiceps, male CNAN-Ar003412. Left palpal bulb: A, dorsal view; B, ventral view; C, retrolateral view; D, prolateral view. Scale = 2 mm. Figure 6. Open in new tabDownload slide Brachypelma albiceps, male CNAN-Ar003412. Left palpal bulb: A, dorsal view; B, ventral view; C, retrolateral view; D, prolateral view. Scale = 2 mm. Figure 7. Open in new tabDownload slide 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). Figure 7. Open in new tabDownload slide 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). Figure 8. Open in new tabDownload slide A–B, Brachypelma albiceps, habitus; C, habitat. A, male; B, female, C, deciduous forest in the habitat of B. albiceps. Photos: J. Mendoza. Figure 8. Open in new tabDownload slide A–B, Brachypelma albiceps, habitus; C, habitat. A, male; B, female, C, deciduous forest in the habitat of B. albiceps. Photos: J. Mendoza. Eurypelma pallidum F. O. Pickard-Cambridge, 1897: 21, plate 1, fig. 17 (female misidentified). Brachypelma albicepsPocock, 1903: 103 (D female). Schmidt (2004: 4, fig. 1) transferred to synonymized generic name. Locht et al. (2005: 108) transferred from Aphonopelma. Estrada-Alvarez (2014: 57, fig. 18), female. Teyssié (2015: 266–267), female. Aphonopelma albiceps (Pocock, 1903). Smith (1994: 70, figs 76–82) removed female from synonymy of A. pallidum. Peters (2000: 23, fig. 43), female. Peters (2003: 32, fig. 107), female. Brachypelmides ruhnauiSchmidt, 1997a: 205, fig. 1–4 (D male and female). Schmidt (1997b: 19, figs 199–201), male and female. Locht, Yáñez & Vázquez (1999: 196, fig. 3), male. Peters (2000: 76, figs 250–252), male and female. Peters (2003: 133, figs 542, 545–547), male and female. Schmidt (2003: 137, fig. 207), male and female. Locht et al. (2005: 108) considered it as junior synonym of B. albiceps. Material examined Holotype of Brachypelma albiceps: MEXICO: Guerrero: ♀, Venta de Zopilote. No more data (NHM labelled as BM1898.12.24.34-37). Holotype and paratype of Brachypelmides ruhnaui: MEXICO: Estado de MEXICO: 1♂, 1♀, Toluca. Col. Mark Ruhnau (SMF39013). Other material: MEXICO: Guerrero: 1 ♀, no more data, source E. Hijmensen, S. Longhorn (RUHZ Longhorn DNA sample 2005); 1♂, 1♀, Mpio. Copalillo, Papalutla, 14/X/2008, J. Mendoza (CNAN-Ar003412); 1 ♀, Mpio. Azoyu, El Carrizo VII/2011, A. Alcaraz (CNAN-Ar007850); 2 ♀, Mpio. Tixtla de Guerrero, Zotoltitlan, 23/IX/2012, J. Mendoza, G. Contreras, J. Cruz, D. Ortiz (CNAN-Ar007839, CNAN-Ar007843); 1 juvenile, Mpio. Chilpancingo de los Bravo, Milpillas, 22/IX/2012, J. Mendoza, G. Contreras, J. Cruz, D. Ortiz (CNAN-Ar007852); 1 ♂, Mpio. Arcelia, Teloloapan, 15/IX/1952, col. L. Vazquez (CNAN-Ar003099); 1 ♂, Mpio. Arcelia, Presa Vicente Guerrero, 05/X/1976, (CNAN-Ar003413); 1 ♂, Mpio. Arcelia, Presa Vicente Guerrero, 23/XI/1976, A. Castillo (CNAN-Ar003437); 1 ♂, Mpio. Eduardo Neri, Ahuelican, 03/X/2008, O. Francke, A. Valdez, T. Lopez (CNAN-Ar004128); 1 ♂, Mpio. Eduardo Neri, Venta Vieja, 26/V/1960, (CNAN-Ar004129); 1 ♂, Mpio. Tepecoacuilco, Ahuehuepan, 30/VIII/2009, O. Francke, A. Valdez, T. Lopez, C. Santibañez (CNAN-Ar004130). Morelos: 1 ♀, Mpio. Cuernavaca, 21/IX/1959, (CNAN-Ar003082); 1 ♂, Mpio. Tequesquitengo, 24/IX/1961, Santibañez (CNAN-Ar003113); 1 ♂, Mpio. Amacuzac, Huajintlán, 28/VIII/2009, O. Francke, A. Valdez, C. Santibañez, T. Lopez, (CNAN-Ar010574). Diagnosis Brachypelma albiceps can be distinguished from all other known Brachypelma species by its coloration, consisting of brown range setae on the carapace and red setae on the opistosoma (Fig. 8A–B). The shape of the genitalia also differs in both sexes: with the palpal bulb being sharp and tapered (Fig. 6) and the spermatheca separated and with rounded receptacles (Fig. 7E–G). Brachypelma albiceps can be identified by possesing the following character combination: male palpal bulb with sharp embolus curving to dorsal through its length (Fig. 6E–D), prolateral superior keel very reduced, apical keel developed, wider behind embolus tip (Fig. 6E–D). Embolus tip directed to retrolateral (Fig. 6A–B). Embolus similar in length to tegulum (Fig. 6). Spermatheca separated with semicircular receptacles. Spermathecal baseplate divided, oval; twice wider than its height (Fig. 7E–G). Carapace of both sexes golden yellow (Fig. 8A–B). Redescription Male (CNAN-Ar003412) (Figs 5–6): Body length 38.45 (not including chelicerae and spinnerets). Carapace 17.64 length, 16.61 width. Caput not markedly elevated; fovea straight, 3.25 wide (Fig. 5A). Eyes: anterior eye row procurved, posterior eye row recurved. Eye sizes and interocular distances: AME 0.45; ALE 0.70; PME 0.35; PLE 0.53; AME–AME 0.38; AME–ALE 0.20; PME–PME 0.98; PME–PLE 0.10; ALE–PLE 0.18. Ocular tubercle width 2.23, length 1.98; clypeus lacking (Fig. 5D). Labium length 2.43, width 2.77; with 95 cuspules. Maxilla inner corner (left–right) with approximately 167–194 cuspules. Cheliceral promargin with (left–right) ten–ten teeth. Sternum length 7.90, width 6.43. Sigilla oval, second and third pairs hardly visible, posterior sigilla once its length from the margin (Fig. 5B). Leg formula: IV, I, III, II. Length of legs and palpal segments (femur, patella, tibia, metatarsus, tarsus, total): I: 16.68, 8.70, 13.10, 12.76, 8.05, 59.29; II: 14.37, 7.54, 11.35, 11.54, 8.62, 53.42; III: 14.13, 7.07, 10.32, 13.78, 8.85, 54.15; IV: 16.88, 7.68, 14.29, 18.88, 10.60, 68.33. Palp: 10.51, 5.96, 9.53, -, 3.88, 29.88. Spinnerets: PMS, 1.90 long, 1.00 apart; PLS, 2.90 basal, 1.95 middle, 3.25 distal. Tarsi I–IV entirely scopulated. Metatarsus I densely scopulated, II densely scopulated, III scopulated 75% distally, IV scopulated 50% distally. Tibia I with two tibial apophyses normally developed which originate from a common base. Prolateral apophysis with one inner conical spine; retrolateral apophysis almost the same width throughout its length and curved to dorsal on apex (Fig. 5F–G). Metatarsus I curved (Fig. 5E). Stridulatory setae: with plumose setae on palp trochanter retrolateral face, and leg I trochanter and femur prolateral face. Chaetotaxy (left side): femora palp 1p; patellae none; tibiae II 2v; III 1r, 2v; IV 1p, 2v; palp 1p, 1v; metatarsi II 2v; III 2p, 6v; IV 2p, 9v, 1r. Palp: embolus tapering and with very reduce spoon-like shape at tip. Embolus slightly twisted to retrolateral. Prolateral superior keel very reduced and thin, prolateral inferior keel absent, apical keel extending to posterior and wider at the embolus tip. The opening of the embolus is on the prolateral side, just behind the opening is located a concavity which delimits the apical keel boundary from the remaining part of the embolus. Tegular apophysis rounded. (Fig. 6A–D). Urticating setae: types I and III arranged in one dorsoposterior patch, black in colour. Type III are located in an oval dorsomedial area extended to posterior. Type I surrounding the type III area, with intermediates between type III and I in transition areas (Fig. 5C). Female (CNAN-Ar003082) (Fig. 7A–E): Body length 53.61 (not including chelicerae and spinnerets), carapace 23.50 length, 22.40 width. Caput not markedly elevated; fovea procurved, 5.80 wide (Fig. 7A). Eyes: anterior eye row procurved, posterior eye row recurved. Eye sizes and interocular distances: AME 0.47; ALE 0.50; PME 0.30; PLE 0.40; AME–AME 0.67; AME–ALE 0.33; PME–PME 1.40; PME–PLE 0.15; ALE–PLE 0.43. Ocular tubercle width 2.80, length 2.43; clypeus lacking (Fig. 7C). Labium length 2.95, width 4.15; with 94 cuspules. Maxilla inner corner (left–right) with approximately 129–130 cuspules (Fig. 7D). Cheliceral promargin with (left–right) nine–ten teeth (proximal to distal: first–third large, fourth–sixth medium, seventh–eighth large, ninth small; first-third large, fourth small, fifth medium, sixth–tenth large). Sternum length 11.40, width 9.43. Sigilla oval, third pair hardly visible; posterior sigilla once its length from the margin (Fig. 7B). Leg formula: IV, I, II, III. Length of legs and palpal segments (femur, patella, tibia, metatarsus, tarsus, total): I: 16.52, 10.27, 13.05, 12.26, 8.36, 60.73; II: 15.35, 9.57, 10.94, 10.57, 7.94, 54.37; III: 14.40, 8.49, 9.30, 12.35, 6.92, 51.46; IV: 17.65, 9.00, 12.90, 17.39, 8.79, 65.91. Palp: 12.88, 6.12, 9.16, -, 9.12, 37.28. Spinnerets: PMS, 3.00 long, 3.50 apart; PLS, 4.75 basal, 3.10 middle, 3.65 distal. Tarsi I–IV entirely scopulated. Metatarsi I–II entirely scopulated, III scopulated 75% distally, IV scopulated 50% distally. Stridulatory setae: with claviform stridulating setae on leg I trochanter and femur prolateral face. Chaetotaxy (left side): femora palp 1p; patellae none; tibiae I 2v, II 1p, 4v; III 3v, 1r; IV 3v, 1r; palp 5v, 1r; metatarsi I 1v; II 3v; III 2p, 5v, 1r; IV 13v, 2r. Genitalia: spermatheca separated with semicircular receptacles strongly sclerotized, ventral face smooth. Spermathecal baseplate divided, oval; twice wider than its height. (Fig. 7E). Urticating setae: types I and III arranged in one dorsoposterior patch, black in colour. Type III are located in an oval dorsomedial area extended to posterior. Type I surround the type III area; with intermediates between type III and I in transition areas. Colour pattern: In living specimens, adults of both sexes have the carapace brown range (pantone 719c) in colour (Fig. 8A–B), with juveniles or subadults paler in colour; chelicerae dorsally dark blue-grey; ventral coxae, labium, maxillae and sternum brownish black; abdomen dorsally black with light orange setae, ventrally brownish black. Legs and palpi: bluish black. (Fig. 8A–B). Distribution and habitat Brachypelma albiceps is known from Central Mexico in Morelos, part of Puebla and Estado de Mexico to central Guerrero as the southern limit of its distribution. Its burrows occur in dense thickets or dry thorn forests under large rocks or tree roots (Figs 2, 8C, 38). There are no traces of silk at the burrow entrance and the interior can often be multitunnelled. The breeding season occurs during the last part of the rainy and first part of dry seasons (August to January). Egg-sacs are constructed through the drier winter months with young emerging and dispersing in late spring, just before the onset of the early summer rains. Brachypelma auratum schmidt, 1992 (Figs 2, 9–13, 36A–D, 38) Figure 9. Open in new tabDownload slide 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 9. Open in new tabDownload slide 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). Brachypelma differs from Tliltocatl by the red/orange coloration pattern on legs and/or carapace (except B. albiceps) (Figs 36, 37, 41). The shape of genitalia also differs in both sexes with the male palpal bulb apex shorter than in Tliltocatl and by the lacking of prolateral inferior keel or restricted to the apex when present (Figs 6, 10), whereas in Tliltocatl it is posteriorly extended and parallel to a prolateral superior keel (Fig 40A–C). It also differs in lacking any spination on patellae of palps or legs. Females differs by the spermatheca apex not as inwardly curved, being generally straight (except B. klaasi and B. albiceps due to divided spermatheca) (Figs 7E–G, 12E–F, 40D). Brachypelma differs also by having the spermatheca baseplate more developed and sclerotized than Tliltocatl when it is present in the last (Figs 12E–F, 16E–G, 40D). Figure 10. Open in new tabDownload slide Brachypelma auratum, male CNAN-Ar003658. Left palpal bulb: A, dorsal view; B, ventral view; C, retrolateral view; D, prolateral view. Scale = 2 mm. Figure 10. Open in new tabDownload slide Brachypelma auratum, male CNAN-Ar003658. Left palpal bulb: A, dorsal view; B, ventral view; C, retrolateral view; D, prolateral view. Scale = 2 mm. Figure 11. Open in new tabDownload slide Brachypelma auratum. A–C, holotype male SMF-38045. A, habitus, dorsal view; B, laminated spermatheca; C, spermatheca, ventral view. Figure 11. Open in new tabDownload slide Brachypelma auratum. A–C, holotype male SMF-38045. A, habitus, dorsal view; B, laminated spermatheca; C, spermatheca, ventral view. Figure 12. Open in new tabDownload slide 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). Figure 12. Open in new tabDownload slide 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). Figure 13. Open in new tabDownload slide 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. Figure 13. Open in new tabDownload slide 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. Brachypelma auratumSchmidt (1992b: 9, fig. 1, 3), D male and female. Schmidt (1993: 82, fig. 187), female. Smith (1994: 160, figs 851–866), male and female. Tesmoingt et al., (1997a: 9, plate 2, fig. 5), female. Schmidt (1997b: 19, fig. 192), female. Locht et al. (1999: 196, fig. 6), female. Peters (2000: 65, figs 210–211), male and female. Peters (2003: 110, figs 439–440, 443, 446–447, male and female. Schmidt (2003: 137, fig. 203), female. Material examined Holotype of Brachypelma auratum: MEXICO: ♂, donated 22/II/1992 by Fritzlen (SMF 38045). Female spermatheca from exuvia mounted in a permanent preparation (SMF 38045). Other material: MEXICO: Michoacán: 1 ♂, Mpio. Los Reyes, Los Reyes Salgado, 2/II/2013, Col. J. Mendoza, G. Contreras, D. Ortiz, D. Barrales (CNAN-Ar003658); 1 ♀, Mpio. San Lucas, Salguero 2/II/2013, J. Mendoza, G. Contreras, D. Ortiz, D. Barrales (CNAN-Ar007164); 2♀, Mpio. La Huacana, 5 km SE of Zicuirán, 16/IX/2015, Col. J. Mendoza (CNAN-Ar007903, CNAN-Ar010284); 1♀, Mpio. La Huacana, 1 km SE of Chau (CNAN-Ar010284); Guerrero: 1 ♀, Mpio. Arcelia, Desviación a Chacamerito, 3/II/2013, J. Mendoza, G. Contreras, J. Cruz, D. Ortiz (CNAN-Ar007136); 1 ♀, Mpio. Chilpancingo de los Bravo, carretera Coyuca de Catalán-Zihuatanejo, cols. A. Zaldívar, M. García, J. Martínez, V. Salinas (CNAN-Ar007878); Estado de MEXICO: 1 ♂, Mpio. Luvianos, 2/11/2002, E. Gonzalez-Santillan, R. Paredes, C. Durán (CNAN-Ar003592); 2 ♂ and 1♀, donation received from private collection of J. Mendoza (CNAN-Ar003660, CNAN-Ar003676. CNAN-Ar003657). Diagnosis Brachypelma auratum can be distinguished from all other known Brachypelma species by the coloration of the legs with red-orange flame-shape on patellae (Fig. 36A–D). It also differs in the shape of the genitalia in both sexes with palpal bulb straight, embolus short and broader at apex (Fig. 10C–D). The prolateral superior keel normally developed and slightly directed retrolaterally (Fig. 10A). Apical keel with a great backward development, extending almost the same length of prolateral superior keel (Fig. 10B–C). The prolateral inferior keel is joined to prolateral superior keel at the apex and slightly extended to downwards (Fig. 10D). Also differs by the spermatheca ventral face smooth, with spermathecal baseplate subrectangular (Figs 11C, 12E–F). Brachypelma auratum is identified by possesing the following character combination: male palpal bulb straight with a broad spoon-shape (Fig. 10C), prolateral superior keel normally developed, directed retrolaterally and slightly extended to backward (Fig. 10A, C); prolateral inferior keel weakly developed restricted to embolus apex, better seen dorsally (Fig. 10A); apical keel developed extending widely backwards (Fig. 10B). Embolus tip slightly directed to retrolateral (Fig. 10A–B). Embolus short and wide, similar in length to tegulum (Fig. 10). Spermatheca fused semicircular, with a single receptacle strongly sclerotized (Figs 11C, 12E–F). Spermathecal baseplate divided, more separated above (Fig. 12E–F); ovate and 2.5 wider than its height (Figs 11C, 12E–F). Both sexes possess an orange flame-shape area over the patellae (Fig. 13A–C). Redescription Male (CNAN-Ar003658) (Figs 9–10): Body length 42.09 (not including chelicerae and spinnerets), carapace 18.16 length, 18.00 width. Caput not markedly elevated; fovea straight, 4.15 wide (Fig. 9A). Eyes: anterior eye row procurved, posterior eye row recurved. Eye sizes and interocular distances: AME 0.40; ALE 0.55; PME 0.40; PLE 0.65; AME–AME 0.50; AME–ALE 0.23; PME–PME 1.13; PME–PLE 0.13; ALE–PLE 0.37. Ocular tubercle width 2.60, length 2.13; clypeus absent (Fig. 9D). Labium length 2.07, width 2.27; with 113 cuspules. Maxilla inner corner with approximately 270 (left) and 248 (right) cuspules. Cheliceral promargin with 10 (left) and 10 (right) teeth (proximal to distal: first large, second small, third large, fourth–tenth large). Sternum length 9.00, width 7.42. Sigilla oval, first to third pairs hardly visible, posterior pair once 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, 14.08, 15.17, 10.32, 66.27; II: 15.92, 9.32, 12.61, 13.91, 9.06, 60.82; III: 14.49, 8.06, 11.73, 14.15, 8.68, 57.11; IV: 17.61, 8.40, 14.79, 19.44, 9.61, 69.85. Palp: 10.88, 7.08, 10.35, -, 3.55, 31.86. Spinnerets: PMS, 2.10 long, 1.30 apart; PLS, 2.83 basal, 2.20 middle, 3.70 distal. Tarsi I–IV entirely scopulated. Metatarsus I entirely scopulated, II scopulated 75%, III scopulated 50% distally, IV scopulated 25% distally. Tibia I with two tibial apophyses normally developed, which originate from a common base. Prolateral apophysis with inner spine half its length; retrolateral apophysis same width throughout its length, apex slightly curved to prolateral (Fig. 9F–G). Metatarsus I curved (Fig. 9E). Stridulatory setae: with claviform stridulating setae on palp trochanter retrolateral face; leg I trochanter and femur prolateral face. Chaetotaxy (left side): femora I 1p; II 1p; palp 1p; patellae none; tibiae I 2p, 1r; II 2p, 3v; III 3p, 4v, 2r; IV 1p, 4v, 1r; palp 2p, 3v; metatarsi II 1p, 1v; III 3p, 7v, 2r; IV 2p, 13v, 2r. Palp: embolus almost straight with a broad spoon-shape, prolateral superior keel normally developed, directed retrolaterally and slightly extended to backward; prolateral inferior keel weakly developed restricted to embolus apex, better seen dorsally, apical keel developed extending widely backwards. Opening of the embolus is on the prolateral side, just behind the opening is located a concavity which delimits the apical keel boundary from the remaining part of the embolus. Embolus apex slightly curved to retrolateral (Fig. 10A–D). Urticating setae: types I and III arranged in one dorsoposterior patch, black in colour. Type III are located in an oval dorsomedial area extended to posterior. Type I surround the type III area; with intermediates between type III and I in transition areas (Fig. 9C). Colour pattern: in live specimens, adult males with carapace peach colour (pantone 712c) around the border, some specimens also present this colour behind the fovea. Carapace colour is black (pantone process black); chelicerae dorsally black; ventral coxae, labium, maxillae and sternum brownish black; abdomen dorsally black with sparse rose colour setae (pantone 487c), ventrally brownish black. Legs and palpi: femora black, patellae with a proximal dorsomedial signal orange (pantone 173c) flame-shape area, distal ring putty colour (pantone 728c), with some brownish pink setae laterally; tibiae proximal three-quarters process black with some sparse brownish pink setae, distal quarter brown range and metatarsi process black with brownish pink setae and a warm grey ring at the terminal end; tarsi black (Fig. 13A). Female (CNAN-Ar007878) (Fig. 12A–E): Body length 61.36 (not including chelicerae and spinnerets), carapace 26.34 length, 24.61 width. Caput not markedly elevated; fovea straight, 5.70 wide (Fig. 12A). Eyes: anterior eye row procurved, posterior eye row recurved. Eye sizes and interocular distances: AME 0.47; ALE 0.80; PME 0.37; PLE 0.70; AME–AME 0.67; AME–ALE 0.33; PME–PME 1.57; PME–PLE 0.13; ALE–PLE 0.43. Ocular tubercle width 2.97, length 2.93; clypeus lacking (Fig. 12C). Labium length 3.05, width 3.55; with 86 cuspules. Maxilla inner corner with approximately 250 (left)–276 (right) cuspules (Fig. 12D). Cheliceral promargin with 10 (left)–11 (right) teeth. Sternum length 11.39, width 9.76. Sigilla oval, first to third pairs hardly visible; posterior pair once its length from the margin (Fig. 12B). Leg formula: IV, I, II, III. Length of legs and palpal segments (femur, patella, tibia, metatarsus, tarsus, total): I: 16.56, 10.85, 13.73, 13.01, 8.64, 62.79; II: 15.65, 10.60, 11.60, 12.49, 7.59, 57.93; III: 14.59, 9.57, 10.83, 13.77, 8.52, 57.28; IV: 17.94, 10.05, 14.04, 18.78, 9.75, 70.56. Palp: 12.62, 8.57, 8.93, -, 10.35, 40.47. Spinnerets: PMS, 2.50 long, 2.05 apart; PLS, 4.15 basal, 4.15 middle, 4.25 distal. Tarsi I–IV entirely scopulated. Metatarsi I entirely scopulated, II scopulated 90%, III scopulated 70% distally, IV scopulated 40% distally. Stridulatory setae: with claviform stridulating setae on palp trochanter retrolateral face; leg I trochanter and femur prolateral face. Chaetotaxy (left side): femora palp 1p; patellae none; tibiae palp 1p, 5v; II 1p, 3v; III 3v; IV 3v; metatarsi II 2v; III 5v, 2r; IV 1p, 8v, 1r. Genitalia: fused semicircular spermatheca, with a single receptacle strongly sclerotized, four times wider than its height. Spermatheca baseplate divided, more separated above, each baseplate ovate 2.5 wider than high, outer side slightly smaller than the inner (Fig. 12E). Variation: some specimens have a wider base up to five times greater than height, young or juveniles can present the superior edge slightly inward in middle. Ventral face smooth. Baseplate division can vary in length (Figs 11C, 12F). Urticating setae: types I and III arranged in one dorsoposterior patch, black in colour. Type III are located in an oval dorsomedial area extended to posterior. Type I surround the type III area, with intermediates between type III and I in transition areas. Colour pattern: in live specimens, adult females with two carapace patterns: (1) light brown (pantone 7414c) around the border and process black in dorsomedial, juveniles or subadults same pattern (Fig. 13B); (2) light brown around the border and extended behind the fovea, with process black from the fovea to caput (Fig. 13C); chelicerae dorsally oxford blue (pantone 532c); ventral coxae, labium, maxillae and sternum brownish black; abdomen dorsally black with sparse rose colour setae (pantone 487c), ventrally brownish black. Legs and palpi: femora black, patellae with a proximal dorsomedial signal orange (pantone 173c) flame-shape area, distal ring putty colour (pantone 728c), with some brownish pink setae laterally; tibiae proximal three quarters process black with some sparse brownish pink setae, distal quarter brown range and metatarsi process black with brownish pink setae and a warm grey ring at the terminal end; tarsi black (Fig. 36A–D). Distribution and habitat: Brachypelma auratum is known from part of the Neotransvolcanic belt, being found in south-western Estado de MEXICO, north-western Guerrero, central Michoacán and the north-eastern border of Jalisco with Michoacán (Figs 2, 38) where it occurs in thorn and deciduous secondary forests (Fig. 13D). It is a fossorial species whose modified or self-excavated burrows can be found under large rocks and large tree roots amongst thorny brush. Burrows do not have any silk around the entrance. Remarks The type specimen of B. auratum (SMF38045) lacks palpal bulbs because they were removed, possibly when it was described (Fig. 11A). It was not possible to find them in the holotype jar, nor in the laminated collection at the Senckenberg Museum. One mounted spermatheca was found instead of a palpal bulb (Fig. 11B–C). The origin of that spermatheca is uncertain. However, the other features like patellae coloration and tibial apophysis in the type specimen are sufficient for identification of this species. Brachypelma baumgarteni Smith, 1993 (Figs 2, 14–17, 37A–D, 38) Figure 14. Open in new tabDownload slide 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 14. Open in new tabDownload slide 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. Open in new tabDownload slide Brachypelma baumgarteni, male CNAN-Ar010588. Left palpal bulb: A, dorsal view; B, ventral view; C, retrolateral view; D, prolateral view. Scale = 2 mm. Figure 15. Open in new tabDownload slide Brachypelma baumgarteni, male CNAN-Ar010588. Left palpal bulb: A, dorsal view; B, ventral view; C, retrolateral view; D, prolateral view. Scale = 2 mm. Figure 16. Open in new tabDownload slide Brachypelma baumgarteni. A–E, female CNAN-Ar007150. 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 CNAN-Ar007149; G, female CNAN-Ar007151. Scale = 10 mm (A–B), 3 mm (D), 1 mm (C, E–G). Figure 16. Open in new tabDownload slide Brachypelma baumgarteni. A–E, female CNAN-Ar007150. 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 CNAN-Ar007149; G, female CNAN-Ar007151. Scale = 10 mm (A–B), 3 mm (D), 1 mm (C, E–G). Figure 17. Open in new tabDownload slide 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. Figure 17. Open in new tabDownload slide 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. Brachypelma baumgarteniSmith, 1993: 15, figs 1–10 (D male); Smith (1994: 163, figs 875–883), male; Teyssié (2015: 269, fig. 1), female. Misidentified by Peters (2000: 66, figs 214–215), male and Peters (2003: 114, figs 457, 460), male, the species shown in the figures is an adult male of Brachypelma hamorii Tesmoingt, Cleton & Verdez, 1997 not B. baumgarteni. Material examined Holotype of Brachypelma baumgarteni: MEXICO: Michoacán: ♂, Sierra Madre del Sur, M. Baumgarten (BMNH-1999-122). Other material: MEXICO: Michoacán: 1 ♂, Carretera La Mira-Arteaga, 09/IX/2012, without collector (CNAN-Ar003597); 2 ♀, Mpio. Lázaro Cárdenas, Los Amates 11/XII/2013, J. Mendoza (CNAN-Ar007149) (CNAN-Ar007150); 1 ♀, Mpio. Lázaro Cárdenas, Puente Chuquiapan, 10/XII/2013 J. Mendoza (CNAN-Ar007151); 1 ♂, Mpio. Lázaro Cárdenas, Los Coyotes, 8/XII/2016, J. Mendoza, R. Ramírez (CNAN-Ar010588). Diagnosis Brachypelma baumgarteni can be distinguished from all other known Brachypelma species by the coloration of the legs with deep orange flame-shape on patellae with yellowish around, tibia and metatarsus with similar yellowish colour (Fig. 37A–D). It also differs in the shape of genitalia in both sexes with palpal bulb slightly curved to dorsal (Fig. 15C–D), embolus shorter than tegulum and broader at apex (Fig. 15A–B). The prolateral superior keel broad and short, slightly directed retrolaterally (Fig. 15A, C). Apical keel thin, extending more than the length of prolateral superior keel (Fig. 15B). Also differs by the spermatheca ventral face smooth, with spermathecal baseplate oblong (Fig. 16E–G). Brachypelma baumgarteni is identified by possesing the following character combination: male palpal bulb slightly curved to dorsal with a narrow spoon-shape at apex, prolateral superior keel short, thin and directed retrolaterally; apical keel short and thin, larger than prolateral superior keel. Embolus tip slightly directed to retrolateral. Embolus compact and thin, shorter in length than tegulum (Fig. 15A–D). Spermatheca fused trapezoidal, with a single receptacle strongly sclerotized. Spermathecal baseplate divided, more 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 seen on legs I and IV (Fig. 32A–D). Description Male (CNAN-Ar010588) (Figs 14, 15, 17A): Body length 49.63 (not including chelicerae and spinnerets), carapace 22.59 length, 21.54 width. Caput not markedly elevated; fovea recurved, 3.75 wide (Fig. 14A). Eyes: anterior eye row procurved, posterior eye row recurved. Eye sizes and interocular distances: AME 0.40; ALE 0.43; PME 0.33; PLE 0.70; AME–AME 0.40; AME–ALE 0.20; PME–PME 0.97; PME–PLE 0.10; ALE–PLE 0.23. Ocular tubercle width 2.50, length 2.13; clypeus 0.17 (Fig 14D). Labium length 2.57, width 2.90; with 80 cuspules. Maxilla inner corner with approximately 179 (left) and 148 (right) cuspules. Cheliceral promargin with ten (left) and ten (right) teeth (proximal to distal: first-third large, fourth medium, fifth large, sixth small, seventh–tenth large). Sternum length 9.60, width 7.68. Sigilla oval, second and third pairs hardly visible, posterior pair twice its length from the margin (Fig. 14B). Leg formula: IV, I, II, III. Length of legs and palpal segments (femur, patella, tibia, metatarsus, tarsus, total): I: 19.29, 9.87, 14.70, 16.21, 9.97, 70.04; II: 17.52, 9.79, 12.35, 13.39, 9.91, 62.96; III: 16.15, 9.13, 11.83, 15.25, 9.01, 61.37; IV: 18.92, 9.99, 15.38, 19.99, 11.10, 75.38. Palp: 11.95, 7.32, 11.26, -, 5.20, 35.73. Spinnerets: PMS, 2.90 long, 1.45 apart; PLS, 4.10 basal, 2.95 middle, 4.15 distal. Tarsi I–IV entirely scopulated. Metatarsus I entirely scopulated, II scopulated 75%, III scopulated 60% distally, IV scopulated 30% distally. Tibia I with two tibial apophyses normally developed, which originate from a common base. Prolateral apophysis with inner spine half its length; retrolateral apophysis same width throughout its length, apex slightly curved to prolateral (Fig. 14F–G). Metatarsus I curved (Fig. 14E). Stridulatory setae: with plumose setae on palp coxa and trochanter retrolateral face; leg I trochanter and femur prolateral face. Chaetotaxy (left side): femora I 1p; II 1p; palp 1p; patellae none; tibiae I 2p; II 3p, 4v; III 3p, 5v, 1r; IV 4v, 1r; palp 2p, 1v; metatarsi I 3v; II 4v; III 3p, 6v, 2r; IV 3p, 12v, 2r. Palp: embolus with narrow spoon-shape, slightly curved to dorsal. Embolus short and flat at base. Prolateral superior keel short, thin and directed retrolaterally. Apical keel short and thin, larger than prolateral superior. Both keels fusioned at apex and extended retrolatrally forming a well defined concave area on retrolateral. Opening of the embolus is on the prolateral side, just behind the opening is located a concavity which delimits the apical keel boundary from the remaining part of the embolus. Embolus apex slightly curved to retrolateral (Fig. 15A–D). Urticating setae: types I and III arranged in one dorsoposterior patch, black in colour. Type III are located in an oval dorsomedial area extended to posterior. Type I surround the type III area, with intermediates between type III and I in transition areas (Fig. 14C). Colour pattern: in live specimens, adult males with carapace rust colour (pantone 167c) with the border lightest. Chelicerae dorsally Del rio colour (pantone 4735c); ventral coxae, labium, maxillae and sternum brownish black; abdomen dorsally black with sparse light terra di siena setae (pantone 472c), ventrally brownish black. Legs and palpi: femora black, patellae with a proximal dorsomedial signal tomato red (pantone 485c) flame-shape area, distodorsal paramedian tierra di siena setae with some long setae laterally of the same colour; tibiae raw sienna (pantone 722c) with some sparse setae from same colour and metatarsi corvette colour (pantone 720c) with mandys pink (pantone 473c) longitudinal line just at metatarsi centre; tarsi black (Fig. 17A). Female (CNAN-Ar007150) (Figs 16, 17B, 37A–D): Body length 65.97 (not including chelicerae and spinnerets), carapace 25.67 length, 23.57 width. Caput not markedly elevated; fovea recurved, 5.10 wide (Fig. 16A). Eyes: anterior eye row procurved, posterior eye row recurved. Eye sizes and interocular distances: AME 0.53; ALE 0.60; PME 0.50; PLE 0.53; AME–AME 0.50; AME–ALE 0.20; PME–PME 1.30; PME–PLE 0.13; ALE–PLE 0.27. Ocular tubercle width 2.53, length 2.47; clypeus 0.47 (Fig. 16C). Labium length 2.85, width 3.85; with 128 cuspules. Maxilla inner corner with approximately 211 (left)–225 (right) cuspules (Fig. 16D). Cheliceral promargin with eight (left)–nine (right) teeth (all big). Sternum length 11.40, width 10.22. Sigilla oval, second and third pairs hardly visible; posterior pair once and half its length from the margin (Fig. 16B). Leg formula: IV, I, II, III. Length of legs and palpal segments (femur, patella, tibia, metatarsus, tarsus, total): I: 17.88, 11.25, 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; IV: 18.60. 10.16, 14.66, 19.72, 10.71, 73.85. Palp: 13.08, 8.25, 10.37, -, 10.14, 41.84. Spinnerets: PMS, 2.50 long, 1.65 apart; PLS, 4.05 basal, 3.10 middle, 4.15 distal. Tarsi I–IV entirely scopulated. Metatarsi I and II entirely scopulated, III scopulated 75% distally, IV scopulated 50% distally. Stridulatory setae with claviform stridulating setae on palp trochanter and femur retrolateral face; leg I coxa, trochanter and femur prolateral face. Chaetotaxy (left side): femora I 1p; II 1p; palp 1p; patellae none; tibiae palp 1p, 6v; I 3v; II 4v; III 1p, 4v, 1r; IV 1p, 3v, 1r; metatarsi I 2v; II 4v; III 2p, 5v, 1r; IV 13v, 2r. Genitalia: fused trapezoidal spermatheca with a single receptacle strongly sclerotized, ventral face looks smooth. Three times wider than its height. Spermatheca baseplate divided, oblong, almost as high as one-third the width of its base. Baseplate division very narrow and poorly sclerotized in the basal half (Fig. 16E). Variation: young or juveniles can present the superior edge slightly inward in middle. Baseplate division can vary in length, and some specimens can look more sclerotized (Fig. 16F, G). Urticating setae: types I and III arranged in one dorsoposterior patch, black in colour. Type III are located in an oval dorsomedial area extended to posterior. Type I surround the type III area, with intermediates between types III and I in transition areas. Colour pattern: in live specimens, adult females with two carapace patterns: (1) provincial pink (pantone 4685c) around the border and black (pantone 426c) in dorsomedial, juveniles or subadults same pattern (Fig. 17B); (2) Tuscany (pantone 7522c) around the border and extended behind the fovea, with black pearl (pantone black 6c) from the fovea to caput (Fig. 17C); chelicerae dorsally manatee (pantone 5285c) with two oriental pink (pantone 7521c) cheliceral bands; ventral coxae, labium, maxillae and sternum black pearl; abdomen dorsally black pearl (pantone black 6c) with rosy brown (pantone 5005c) setae, ventrally black pearl. Legs and palpi: femora black pearl, patellae with a proximal dorsomedial cinnabar (pantone 7597c) flame-shape area, distodorsal paramedian spanish white (pantone 4685c) setae with some long setae laterally of the same colour; tibiae eunry (pantone 4735c) with some sparse setae from same colour and metatarsi hemp (pantone 8021c) with mandys pink (pantone 473c) longitudinal line just at metatarsi centre; tarsi black (Fig. 17A–D). Distribution and habitat: Brachypelma baumgarteni is known only from Michoacán on the Sierra Madre del Sur Region (Figs 2, 38), where it occurs in deciduous forests (Fig. 17D). It is a fossorial species whose modified or self-excavated burrows, can be found between large tree roots amongst large trees. Burrows do not have any silk around the entrance. Brachypelma boehmei Schmidt & Klaas, 1993 (Figs 2, 18–22, 37E–H, 38) Figure 18. Open in new tabDownload slide 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). Figure 18. Open in new tabDownload slide 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). Figure 19. Open in new tabDownload slide Brachypelma boehmei, male CNAN-Ar010589. Left palpal bulb: A, dorsal view; B, ventral view; C, retrolateral view; D, prolateral view. Scale = 2 mm. Figure 19. Open in new tabDownload slide Brachypelma boehmei, male CNAN-Ar010589. Left palpal bulb: A, dorsal view; B, ventral view; C, retrolateral view; D, prolateral view. Scale = 2 mm. Figure 20. Open in new tabDownload slide Brachypelma boehmei. A–B, holotype female SMF-40590. A, exuvia used as holotypus; B, spermatheca, ventral view (was not removed from the exuvia). Figure 20. Open in new tabDownload slide Brachypelma boehmei. A–B, holotype female SMF-40590. A, exuvia used as holotypus; B, spermatheca, ventral view (was not removed from the exuvia). Figure 21. Open in new tabDownload slide Brachypelma boehmei. A–E, female CNAN-Ar007905. 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 CNAN-Ar007185; G, female CNAN-Ar007833. Scale = 10 mm (A–B), 3 mm (D), 1 mm (C, E–G). Figure 21. Open in new tabDownload slide Brachypelma boehmei. A–E, female CNAN-Ar007905. 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 CNAN-Ar007185; G, female CNAN-Ar007833. Scale = 10 mm (A–B), 3 mm (D), 1 mm (C, E–G). Figure 22. Open in new tabDownload slide 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. Figure 22. Open in new tabDownload slide 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. Brachypelma boehmeiSchmidt & Klaas, 1993: 7, figs 1–2 (D male and female); Schmidt (1993: 82, fig. 193a), male; Schmidt & Klaas (1994: 8, figs 1–2), male and female; Smith (1994: 164, figs 884–900), male and female; Tesmoingt et al., (1997a: 9, plate 2, fig. 1), female; Schmidt (1997b: 19, fig. 190), male; Bertani (2000: 30, figs 41–42), male; Peters (2000: 67, fig. 218), female; Peters (2003: 115–117, figs 461–469), male and female; Schmidt (2003: 137, figs 201, 273), male and female. Material examined Holotype of Brachypelma boehmei: MEXICO: ♀, no more data, K. Böhme (SMF 40590). Paratype. MEXICO: ♂, no more data, K. Böhme (SMF 38043). Other material: MEXICO: Guerrero: 1 ♂, 2 ♀, Mpio. La Unión Isidoro Montes de Oca, 3km NE of La Unión, 12/XII/2013, J. Mendoza (CNAN-Ar007185, CNAN-Ar007186, CNAN-Ar007833); 2 ♂, Mpio. La Unión Isidoro Montes de Oca, 1km SW of La Unión, 17/XII/2016, J. Mendoza (CNAN-Ar010589, CNAN-Ar010591); 2 ♀, Mpio. La Unión Isidoro Montes de Oca, Carretera Fed 37D, 16/IX/2015, J. Mendoza (CNAN-Ar007905, CNAN-Ar010285); Oaxaca: 1 ♂, Mpio. Salina Cruz, X/1963, E. Martín (CNAN-Ar003426) (collection data of this specimen are considered wrong due to lack of evidence of the species in Oaxaca and considering the restricted distribution of the species to a single municipality in Guerrero). Diagnosis Brachypelma boehmei can be distinguished from all other known Brachypelma species by the coloration of the carapace and legs that are orange (Fig. 22A–C). It also differs in the shape of genitalia in both sexes with palpal bulb straight (Fig. 19C–D), embolus similar in length as tegulum and broader than embolus base at apex (Fig. 19A–B). The prolateral superior keel normally developed and slightly directed retrolaterally (Fig. 19A). Apical keel with a great backward development, extending more than the length of prolateral superior keel (Fig. 19B). The prolateral inferior keel is joined to prolateral superior keel at the apex and slightly extended to backwards (Fig. 19A). The prolateral apophysis is almost half the length of the retrolateral apophysis (Fig. 18F–G). It also differs in the spermatheca ventral face striated (Fig. 20B), with spermathecal baseplate lanceolate, notoriously separated and lower than the height of the spermatheca (Fig. 21E–G). Brachypelma boehmei is identified by possesing the following character combination: male palpal bulb straight with a broad spoon-shape at apex, prolateral superior keel normally developed and slightly directed retrolaterally; prolateral inferior keel weakly developed restricted to embolus apex; apical keel developed extending to backwards more than the length of the prolateral superior keel. Embolus tip slightly directed to retrolateral. Embolus large and broad, similar in length to tegulum (Fig. 19A–D). Spermatheca fused trapezoidal, with a single receptacle strongly sclerotized. Spermathecal baseplate divided, noticeably separated; lanceolate and three times wider than its height (Figs 20B, 21E–G). Carapace and legs of both sexes are orange in general (Fig. 22A–C). Redescription Male (CNAN-Ar010589) (Figs 18, 19, 22A): Body length 41.39 (not including chelicerae and spinnerets), carapace 19.71 length, 18.82 width. Caput not markedly elevated; fovea recurved, 4.50 wide (Fig. 18A). Eyes: anterior eye row procurved, posterior eye row recurved. Eye sizes and interocular distances: AME 0.46; ALE 0.53; PME 0.46; PLE 0.56; AME–AME 0.43; AME–ALE 0.20; PME–PME 1.20; PME–PLE 0.13; ALE–PLE 0.43. Ocular tubercle width 2.47, length 2.23; clypeus absent (Fig. 18D). Labium length 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 (right) teeth (proximal to distal: first–second large, third medium, fourth large, fifth small, sixth–tenth large). Sternum length 8.70, width 8.09. Sigilla oval, first to third pairs hardly visible, posterior pair once its length from the margin (Fig. 18B). Leg formula: IV, I, II, III. Length of legs and palpal segments (femur, patella, tibia, metatarsus, tarsus, total): I: 16.94, 9.01, 13.20, 13.72, 8.47, 61.34; II: 15.56, 8.79, 11.94, 12.60, 8.48, 57.37; III: 14.40, 8.00, 11.29, 13.86, 8.43, 55.98; IV: 17.26, 8.51, 13.06, 18.04, 9.26, 66.13. Palp: 11.02, 6.54, 9.51, -, 4.51, 31.58. Spinnerets: PMS, 2.15 long, 0.95 apart; PLS, 3.80 basal, 2.55 middle, 3.25 distal. Tarsi I–IV entirely scopulated. Metatarsus I entirely scopulated, II scopulated 80%, III scopulated 60% distally, IV scopulated 40% distally. Tibia I with two tibial apophyses normally developed, which originate from a common base. Prolateral apophysis with inner spine half its length; retrolateral apophysis same width throughout its length, apex slightly curved to prolateral (Fig. 18F–G). Metatarsus I curved (Fig. 18E). Stridulatory setae: with claviform stridulating setae on palp trochanter and femur retrolateral face; leg I coxa, trochanter and femur prolateral face. Chaetotaxy (left side): femora I 1p; II 1p; palp 1p; patellae none; tibiae II 3p, 3v, 1r; III 3p, 4v, 2r; IV 1p, 4v, 1r; palp 2p, 2v; metatarsi I 2v, II 1p, 4v; III 2p, 7v, 2r; IV 2p, 15v, 1r. Palp: embolus straight with a broad spoon-shape at apex and similar in length as tegulum. Prolateral superior keel normally developed and slightly directed retrolaterally; prolateral inferior keel weakly developed restricted to embolus apex, better seen dorsally, apical keel developed extending to backwards more than the length of the prolateral superior keel. Opening of the embolus is on the prolateral side, just behind the opening is located a concavity which delimits the apical keel boundary from the remaining part of the embolus. Embolus apex slightly curved to retrolateral (Fig. 19A–D). Urticating setae: types I and III arranged in one dorsoposterior patch, black in colour. Type III are located in an oval dorsomedial area extended to posterior. Type I surround the type III area, with intermediates between type III and I in transition areas (Fig. 18C). Colour pattern: in live specimens, adult males with carapace colour persimmon (pantone 166c) and tacao (pantone 721c) around the border (Fig 22A); chelicerae dorsally tussock (pantone 729c); ventral coxae, labium, maxillae and sternum seal brown (pantone 440c); abdomen dorsally coffee bean (pantone black 2c) with sparse light raw sienna (pantone 722c) setae, ventrally seal brown. Legs and palpi: femora coffee black, patellae with a proximal dorsomedial persimmon flame-shape area, surrounding this and distally is raw sienna, with some setae laterally of same colour; tibiae and metatarsi persimmon, with some lateral raw sienna setae; tarsi proximally backer’s chocolate (pantone 732c) and distally coffee bean (Fig. 37E–H). Female (CNAN-Ar007905) (Figs 21A–E, 22B): Body length 61.36 (not including chelicerae and spinnerets), carapace 26.34 length, 24.61 width. Caput not markedly elevated; fovea straight, 5.70 wide (Fig. 21A). Eyes: anterior eye row procurved, posterior eye row recurved. Eye sizes and interocular distances: AME 0.47; ALE 0.80; PME 0.37; PLE 0.70; AME–AME 0.67; AME–ALE 0.33; PME–PME 1.57; PME–PLE 0.13; ALE–PLE 0.43. Ocular tubercle width 2.97, length 2.93; clypeus lacking (Fig. 21C). Labium length 3.05, width 3.55; with 86 cuspules. Maxilla inner corner with approximately 250 (left)–276 (right) cuspules (Fig. 21D). Cheliceral promargin with ten (left)–11 (right) teeth. Sternum length 11.39, width 10.09. Sigilla oval, first to third pairs hardly visible; posterior pair once its length from the margin (Fig. 21B). Leg formula: IV, I, II, III. Length of legs and palpal segments (femur, patella, tibia, metatarsus, tarsus, total): I: 16.56, 10.85, 13.73, 13.01, 8.64, 62.79; II: 15.65, 10.60, 11.60, 12.49, 7.59, 57.93; III: 14.59, 9.57, 10.83, 13.77, 8.52, 57.28; IV: 17.94, 10.05, 14.04, 18.78, 9.75, 70.56. Palp: 12.48, 7.51, 9.34, -, 9.32. Spinnerets: PMS, 2.50 long, 2.05 apart; PLS, 4.15 basal, 4.15 middle, 2.55 distal. Tarsi I–IV entirely scopulated. Metatarsi I entirely scopulated, II scopulated 90%, III scopulated 70% distally, IV scopulated 40% distally. Stridulatory setae: with claviform stridulating setae on palp trochanter retrolateral face; leg I trochanter and femur prolateral face. Chaetotaxy (left side): femora I 1p, II 1p; patellae none; tibiae I 1p, 3v; II 3p, 4v; III 3p, 6v, 2r; IV 1p, 4v, 1r; palp 2p, 6v; metatarsi I 3v; II 1p, 3v; III 2p, 8v, 1r; IV 2p, 15v, 1r. Genitalia: fused trapezoidal spermatheca, with a single receptacle strongly sclerotized, three times wider than its height. Spermatheca baseplate divided, widely separated in middle, each baseplate lanceolate three times wider than high, outer side slightly smaller than the inner (Fig. 21E). Variation: some specimens have a wider base up to four times wider than height, young or juveniles can present the superior edge slightly inward in middle. Ventral with variation on striation. Baseplate division can vary in length (Figs 20B, 21F–G). Urticating setae: types I and III arranged in one dorsoposterior patch, black in colour. Type III are located in an oval dorsomedial area extended to posterior. Type I surround the type III area, with intermediates between type III and I in transition areas. Colour pattern: in live specimens, adult females with two carapace patterns: (1) tango (pantone 7412c) in all the carapace with pink flare (pantone 5025c) around the border, juveniles or subadults same pattern (Fig 22B); (2) pink flare around the border and sandy brown (pantone 7411c) in dorsomedial, with two seal brown (pantone 440c) longitudinal patches from the fovea to caput (Fig. 22C); chelicerae dorsally beaver colour (pantone 4715c); ventral coxae, labium, maxillae and sternum seal brown; abdomen dorsally black (pantone 426c) with putty colour (pantone 721c) setae, ventrally black. Legs and palpi: femora coffee black, patellae with a proximal dorsomedial cinnabar (pantone 173c) flame-shape area, surrounding this and distally is dark salmon (pantone 472c), with some setae laterally of wafer colour (pantone 4745c); tibiae and metatarsi chardonnay (pantone 1355c), with some lateral wafer setae; tarsi proximally Backer’s chocolate (pantone 732c) and distally coffee bean (Fig. 37E–H). Distribution and habitat: Brachypelma boehmei is the species with the most restricted distribution being only known from the Sierra de Cumbres region, an area 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 and deciduous forests (Fig. 22D). It is a fossorial species whose burrows can be found under large angular rocks and large tree roots amongst thorny brush. Burrows do not have any silk around the entrance. Brachypelma emilia (White, 1856) (Figs 2, 23–26, 36E–H, 38) Figure 23. Open in new tabDownload slide Brachypelma emilia, male CNAN-Ar003599. 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 23. Open in new tabDownload slide Brachypelma emilia, male CNAN-Ar003599. 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 24. Open in new tabDownload slide Brachypelma emilia, male CNAN-Ar003599. Left palpal bulb: A, dorsal view; B, ventral view; C, retrolateral view; D, prolateral view. Scale = 2 mm. Figure 24. Open in new tabDownload slide Brachypelma emilia, male CNAN-Ar003599. Left palpal bulb: A, dorsal view; B, ventral view; C, retrolateral view; D, prolateral view. Scale = 2 mm. Figure 25. Open in new tabDownload slide 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). Figure 25. Open in new tabDownload slide 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). Figure 26. Open in new tabDownload slide 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. Figure 26. Open in new tabDownload slide 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. Mygale emiliaWhite, 1856: 185, pl. 43 (D male). Brachypelma emilia (Simon 1891: 338), D male and female; Smith (1986: 49, fig. 27h), Tmf from Eurypelma=Avicularia; Smith (1987: 49, plate 2, fig. 27h), male; Hancock & Hancock (1989: 46, fig. 41), female; Schmidt (1992: 10), Tmf from Euathlus per Raven; Schmidt (1993: 82, fig. 188), female; Smith (1994: 166, figs 901–915), male and female; Pérez-Miles et al. (1996: 46, figs 9–10), male and female; Tesmoingt et al. (1997a: 9, plate 2, fig. 6), female; Schmidt (1997: 19, figs 191, 193), male and female; Locht et al. (1999: 196, fig. 7), female; Peters (2000: 68, fig. 222), female; Bertani (2001: 338, figs 153–156), male and female; Peters (2003: 117, figs 473–474, 477, 480, 483), male and female; Schmidt I2003: 152, figs 274–277), male and female; Gabriel & Longhorn (2015: 100, fig. 13), female. Material examined Neotype of Brachypelma emilia: MEXICO: Durango: ♂, Ciudad, leg Mr. Forrer (BMNH-1898-12-24-32). 1 ♂ (labelled as paratype), Ciudad, leg. Forrer (OUNMH Jar 106). Note: Both specimens mentioned as neotype and paratype were designated by Smith (1994: 166). Other material: MEXICO: Nayarit: 1 ♂ 4 ♀, Mpio. Compostela, 9.xii.2012, E. Goyer, E. Hijmensen, D. Ortiz (CNAN-Ar003599, CNAN-Ar007153, CNAN-Ar007173, CNAN-Ar007178, CNAN-Ar007875); 1 ♂, Mpio. Estación Ruíz, 1.xii.1989, A. Cadena (CNAN-Ar003436); 4 ♂, Mpio. Estación Ruíz, 5.xii.2014, J. Mendoza, G. Contreras (CNAN-Ar007146, CNAN-Ar007894, CNAN-Ar007895, CNAN-Ar007899); Sinaloa: 1 ♂, Mpio. Mazatlán, vii.1959, without more data (CNAN-Ar003427); 1 ♂ 1 ♀, Mpio. Mazatlán, 3.xii.2014, J. Mendoza, G. Contreras (CNAN-Ar007898, CNAN-Ar010602); 1 ♂, Mpio. Mazatlán, without more data, Collection E. Simon (MP Ar4871A); 1 ♂, Mpio. Sinaloa de Leyva, 30.i.1965, without more data (CNAN-Ar003590); Sonora: 1 ♂, Mpio. Altar, 9.i.1970, W. Lopez Forment (CNAN-Ar003578); Jalisco: 1 ♀, Norte del Río Santiago, Godman, Salvin, without more data (BMNH-1962-2-28-1); 1 ♂, donation received from private collection of J. Mendoza (CNAN-Ar003631). Diagnosis Brachypelma emilia can be distinguished from all other known Brachypelma species by the coloration of the carapace and legs, with carapace orange except in the caput, which is black in colour (Fig. 26A–C). The legs have black femora and patellae, orange tibiae and metatarsi I–III are black and IV is orange (Fig. 36E–H). It also differs in the shape of genitalia in both sexes with palpal bulb slightly curved to dorsal having a small and narrow spoon-shape (Fig. 24C–D). The prolateral superior keel is slightly developed, thin and directed retrolaterally (Fig. 24A, C). The prolateral inferior keel is absent, the apical keel normally developed, wide but not shorter than prolateral superior keel (Fig. 24B). It also differs by the spermatheca ventral face smooth, with a single receptacle strongly sclerotized slightly notched in the middle, spermathecal baseplate oblanceolate (Fig. 25E, F). Brachypelma emilia is identified by possesing the following character combination: male palpal bulb with narrow spoon-shape embolus curving slightly to dorsal through its length, prolateral superior keel slightly developed; apical keel normally developed, wide but not shorter than the prolateral superior keel tip. Embolus tip directed to retrolateral. Embolus similar in length than tegulum (Fig. 24A–D). Spermatheca fused with single semitrapezoidal receptacle. Spermathecal baseplate divided, oblanceolate; one and half wider than its height (Fig. 25E, F). Carapace of both sexes carapace orange with black triangle in the ocular area (Fig. 26A–C). Redescription Male (CNAN-Ar003599) (Figs 23, 24, 26A): Body length 40.16 (not including chelicerae and spinnerets), carapace 20.23 length, 18.85 width. Caput not markedly elevated; fovea procurved, 3.80 wide (Fig. 23A). Eyes: anterior eye row procurved, posterior eye row recurved. Eye sizes and interocular distances: AME 0.43; ALE 0.60; PME 0.33; PLE 0.37; AME–AME 0.70; AME–ALE 0.20; PME–PME 1.23; PME–PLE 0.13; ALE–PLE 0.37. Ocular tubercle width 2.47, length 2.27; clypeus 0.23 (Fig. 23D). Labium length 2.60, width 3.40; with 109 cuspules. Maxilla inner corner with approximately 205 (left) and 179 (right) cuspules. Cheliceral promargin with ten (left) and nine (right) teeth (proximal to distal: first–third large, fourth small, fifth–seventh medium, eighth–tenth large; first–third large, fourth small, fifth–seventh medium, eighth–ninth large). Sternum length 10.10, width 9.38. Sigilla oval, fourth pair hardly visible, posterior pair one and half its length from the margin (Fig. 23B). Leg formula: IV, I, II, III. Length of legs and palpal segments (femur, patella, tibia, metatarsus, tarsus, total): I: 17.25, 9.39, 11.67, 12.22, 9.28, 59.81; II: 15.93, 9.20, 10.96, 11.89, 8.21, 56.19; III: 14.35, 8.11, 10.00, 12.08, 8.15, 52.69; IV: 16.86, 8.63, 12.81, 16.13, 9.92, 64.35. Palp: 11.83, 7.23, 9.61, -, 4.14, 32.81. Spinnerets: PMS, 2.20 long, 1.00 apart; PLS, 3.10 basal, 2.20 middle, 3.20 distal. Tarsi I–IV entirely scopulated. Metatarsus I entirely scopulated, II scopulated 75%, III scopulated 50% distally, IV scopulated 30% distally. Tibia I with two tibial apophyses normally developed, which originate from a common base. Prolateral apophysis with inner spine third its length; retrolateral apophysis wider in basal half, apex almost straight (Fig. 23F, G). Metatarsus I curved (Fig. 23E). Stridulatory setae: with claviform stridulating setae on palp trochanter retrolateral face; leg I trochanter and femur prolateral face. Chaetotaxy (left side): femora I 1p; II 1p; palp 1p; patellae none; tibiae I 1p, 2v; II 2p, 3v; III 2p, 5v, 1r; IV 1p, 4v, 1p; palp 2p, 1v; metatarsi I 1v; II 2v; III 3p, 7v, 1r; IV 1p, 18v, 1r. Palp: embolus slightly curved to dorsal having a small and narrow spoon-shape, prolateral superior keel slightly developed, thin and directed retrolaterally; prolateral inferior keel absent; apical keel normally developed, wide but not shorter than prolateral superior keel. Opening of the embolus is on the prolateral side, just behind the opening is located a concavity which delimits the apical keel boundary from the remaining part of the embolus. Embolus apex slightly curved to retrolateral (Fig. 24A–D). Urticating setae: types I and III arranged in one dorsoposterior patch, black in colour. Type III are located in an oval dorsomedial area extended to posterior. Type I surround the type III area, with intermediates between type III and I in transition areas (Fig. 23C). Female (CNAN-Ar010602) (Fig. 25A–E): Body length 61.31 (not including chelicerae and spinnerets), carapace 26.34 length, 24.33 width. Caput not markedly elevated; fovea straight, 6.40 wide (Fig. 25A). Eyes: anterior eye row procurved, posterior eye row recurved. Eye sizes and interocular distances: AME 0.65; ALE 0.70; PME 0.50; PLE 0.70; AME–AME 0.63; AME–ALE 0.33; PME–PME 1.57; PME–PLE 0.07; ALE–PLE 0.40. Ocular tubercle width 3.25, length 2.83; clypeus 0.27 (Fig. 25D). Labium length 3.05, width 3.85; with 94 cuspules. Maxilla inner corner with approximately 187 (left)–208 (right) cuspules. Cheliceral promargin with nine (left)–nine (right) teeth. Sternum length 11.60, width 10.90. Sigilla oval, fourth pair hardly visible; posterior pair twice its length from the margin (Fig. 25B). Leg formula: IV, I, II, III. Length of legs and palpal segments (femur, patella, tibia, metatarsus, tarsus, total): I: 17.87, 10.79, 13.69, 13.40, 8.75, 64.50; II: 16.35, 10.18, 12.05, 11.77, 8.94, 59.29; III: 15.69, 9.40, 10.97, 12.79, 8.50, 57.35; IV: 18.34, 10.20, 14.20, 17.91, 9.98, 70.63. Palp: 13.20, 8.35, 10.12, -, 9.92, 41.59. Spinnerets: PMS, 2.90 long, 3.00 apart; PLS, 4.30 basal, 2.90 middle, 4.35 distal. Tarsi I–IV entirely scopulated. Metatarsus I entirely scopulated, II scopulated 90%, III scopulated 50% distally, IV scopulated 40% distally. Stridulatory setae: with claviform stridulating setae on palp trochanter and femur retrolateral face; leg I trochanter and femur prolateral face. Chaetotaxy (left side): femora I 1p; palp 1p; patellae none; tibiae I 1p, 3v; II 1p, 4v; III 1p, 5v, 1r; IV 1p, 6v, 1r; palp 2p, 5v; metatarsi I 4v; II 4v; III 1p, 8v, 2r; IV 2p, 14v, 1r. Genitalia: fused semicircular spermatheca, with a single receptacle strongly sclerotized slightly notched in the middle, four times wider than its height. Spermatheca baseplate divided, widely separated above, each baseplate oblanceolate 3.5 wider than high, inner side smaller than the outer (Fig. 25E). Variation: some specimens have a wider base up to five times greater than height, young or juveniles can present the superior edge slightly inward in middle. Ventral face smooth. Baseplate division can vary in length (Fig. 25F). Urticating setae: types I and III arranged in one dorsoposterior patch, black in colour. Type III are located in an oval dorsomedial area extended to posterior. Type I surround the type III area, with intermediates between type III and I in transition areas (Fig. 25C). Colour pattern: In live specimens, adults of both sexes have the carapace orange (pantone 7412c) on almost all the carapace except in the caput, which is pantone process black in colour, and also has a longitudinal line of beaver colour (pantone 4715c) that goes from back of the eyes to the fovea (Figs. 26A–C); chelicerae dorsally blue whale colour (pantone 533c); ventral coxae, labium, maxillae and sternum black pearl colour (pantone black 6c); abdomen dorsally black with Christine colour setae (pantone 7583c), ventrally black pearl colour. Legs and palpi: femora and patellae black pearl; tibiae orange (pantone 157c); metatarsi I–III black (pantone black 7c) with proximal third rope colour (pantone 876c), IV black (pantone black 7c) with large brandy punch colour (pantone 722c) covering it almost completely; tarsi black (pantone black c) (Fig. 36E–H). With juveniles of same pattern but paler in colour. Distribution and habitat: Brachypelma emilia is known from the northern Pacific Coast region on the 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 Jalisco with Nayarit. It could also be possibly found in Durango, but there is no evidence for that yet (Figs 2, 38). It occurs in drier, coastal thorn scrub, grasslands, palm transition to deciduous forest, and into higher elevations of oak forest (Fig. 26D). It is a fossorial species whose modified or self-excavated burrows can be found under large rocks, under dense thorny thickets, large tree roots or burrows in leafy ground cover, in both forested and moderately disturbed areas. Some can be also found close to houses or human structures, but this is most likely because the spiders lived there before these buildings were constructed. Burrows do not have any silk around the entrance to indicate there is a spider inside. This species is sympatric (overlapping distributions) with a small population of Brachypelma klaasi in south-western Nayarit (Mendoza, personal observation). Brachypelma hamorii Tesmoingt et al, 1997 (Figs 2, 27–28, 35B, D, 37I–L, 38) Figure 27. Open in new tabDownload slide 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). Figure 27. Open in new tabDownload slide 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). Figure 28. Open in new tabDownload slide 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. Figure 28. Open in new tabDownload slide 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. Brachypelma hamoriiTesmoingt et al. (1997a: 9, plates 1–6), D male and female; Tesmoingt et al. (1997b: 3, plates 9–11), male; Mendoza & Francke (2017: 167, figs 24–40, 46–49, 52, 57–60), male and female. Euathlus smithi (F. O. Pickard-Cambridge): Baxter (1993: 73, figs 17–18, plate A, figs 1–6), misidentification. Brachypelma smithi (F. O. Pickard-Cambridge): Hancock & Hancock (1989: 44, fig. 39), misidentification female; Schmidt (1992a: 10), transferred from Euathlus; Schmidt (1992b: 14, figs 2, 4), male and female; Schmidt (1993: 82, fig. 190), female; Smith (1994: 170, figs 940–956), male and female; Tesmoingt et al. (1997a: 9, plate 2, fig. 2), female; Tesmoingt et al. (1997b: 4, plates 10–11), male; Schmidt (1997: 19, fig. 195), female; Peters (2000: 72, figs 235–236), male and female; Peters (2003: 125, figs 510, 512–513), male and female; Schmidt (2003: 153, figs 283–284), male and female. Misidentifications. Material examined Neotype of Brachypelma hamorii: MEXICO: Colima: ♂, Mpio. Tecoman, 5/XII/2013, D. Barrales, G. Contreras, D. Ortiz (CNAN-T0900). Designated as neotype by Mendoza & Francke (2017: 167). Other material: MEXICO: Colima: 1 ♂, Mpio. Tecoman, 1/XII/2012, E. Goyer, E. Hijmensen, D. Ortiz (CNAN-Ar003614); 1 ♀, Mpio. Colima, 30/XI/2012, E. Goyer, E. Hijmensen, D. Ortiz (CNAN-Ar004779); 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, G. Contreras (CNAN-Ar007168); 1 ♂, Mpio. Manzanillo, IV/2004, IBT (CNAN-Ar010278); 1 ♂, Mpio. Manzanillo, 4/XII/2013, D. Ortíz, D. Barrales, G. Contreras (CNAN-Ar010277); 1 ♂, donation received from private collection of J. Mendoza (CNAN-Ar003616); 1 ♂, Mpio. 3/XII/2013, D. Ortíz, D. Barrales, G. Contreras (CNAN-Ar007827); 1 ♂, Mpio. Manzanillo, 4/XII/2013, D. Ortiz, D. Barrales, G. Contreras (CNAN-Ar007171); 3 ♀, Mpio. 11 km al SE. de Colima (CNAN-Ar007870, CNAN-Ar007872, CNAN-Ar007873); 1 ♀, Mpio. Tecoman, 2/XII/2012, E. Goyer, E. Hijmensen, D. Ortiz (CNAN-Ar007871); Jalisco: 2 ♀, Mpio. Pihuamo, 30/XI/2012, E. Goyer, E. Hijmensen, D. Ortiz (CNAN-Ar007874, CNAN-Ar010279); 1 ♂, Carretera entre Colima y Jalisco, carretera Colima-Cd. Victoria, 8/VIII/2008, A. Cervantes, M. E. Olsen (CNAN-Ar003425); Michoacán: 1 ♂, Mpio. Lázaro Cardenás, 8/XI/2011, G. Vila (CNAN-Ar003340); 1 ♂, Michoacán, Mpio. Aquila, G. Vila (CNAN-Ar003659). Diagnosis Brachypelma hamorii can be distinguished from all other known Brachypelma species (except B. smithi) by the coloration of the legs with yellow-orange coloration on patellae, tibiae and metatarsi (Figs 28, 37I–L). It also differs in the shape of genitalia in both sexes with palpal bulb curved upward and short (Fig. 27C–D) and spermatheca trapezoid rounded in shape (Fig. 27E–F). It differs from B. smithi by the narrow curved upward 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 spermatheca ventral face smooth, with spermathecal baseplate elliptic (Fig. 27E–F). Although similar in coloration, B. hamorii differs from B. smithi in the patellae flame pattern not as brightly coloured as the one exhibited by B. smithi (Fig. 37I–L). The sides of the patellae are typically black rather than the yellowish-orange as is typical in B. smithi (Fig. 35C–D). The lateral setae are whitish throughout the length of legs, while in B. smithi, the same are yellowish (Fig. 37I–L). In addition, B. hamorii typically have longitudinal cheliceral lines of lighting setae, which are not exhibited by B. smithi (Fig. 35A–B). The carapace of B. hamorii is typically black, bordered with pink or orange in both sexes (Fig. 28A–B, E–F). Rare variants of females can have a black radiating pattern, bordered with pale yellow (Fig. 28D), and rare variants of males may have a carapace that is almost yellowish (Fig. 28C). Brachypelma hamorii is identified by the following character combination: male palpal bulb with narrow spoon-like embolus, prolateral superior keel shorter than in other species, thin and directed retrolaterally, prolateral inferior keel weakly developed directed from dorsal to ventral, better seen dorsally, apical 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. 27C–D). Spermatheca trapezoid rounded; ventral face is almost smooth; Spermathecal baseplate divided, elliptic; five times wider than its height (Fig. 27E–F). Carapace in general black with orange setae on border, with cheliceral longitudinal line of lighting setae (Fig. 35B). Legs and palpi with deep orange and pale orange in patellae, tibiae and metatarsi with orange yellow/white setae on distal (Fig. 37I–L). Note: Redescription of this species was done by Mendoza & Francke (2017). Brachypelma klaasi (Schmidt & Krausse, 1994) (Figs 2, 29–32, 36I–L, 38) Figure 29. Open in new tabDownload slide Brachypelma klaasi, male CNAN-Ar003333. 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 29. Open in new tabDownload slide Brachypelma klaasi, male CNAN-Ar003333. 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 30. Open in new tabDownload slide Brachypelma klaasi, male CNAN-Ar003333. Left palpal bulb: A, dorsal view; B, ventral view; C, retrolateral view; D, prolateral view. Scale = 2 mm. Figure 30. Open in new tabDownload slide Brachypelma klaasi, male CNAN-Ar003333. Left palpal bulb: A, dorsal view; B, ventral view; C, retrolateral view; D, prolateral view. Scale = 2 mm. Figure 31. Open in new tabDownload slide 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). Figure 31. Open in new tabDownload slide 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). Figure 32. Open in new tabDownload slide 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. Figure 32. Open in new tabDownload slide 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. Brachypelmides klaasiSchmidt & Krause, 1994: 7, figs 1–2, D male and female; Schmidt (1997b: 19, figs 198, 202), male and female; Locht et al. (1999: 196, figs 4, 9), male and female; Vol (1999: 11, fig. A), female; Peters (2000: 75, figs 244–247), male and female; Peters (2003: 131, figs 533–535, 537), male and female; Schmidt (2003: 137, figs 204, 208), male and female. Smith (1994: 159) transferred the species to Brachypelma. Brachypelma klaasiSmith (1994: 169, figs 926–939), male and female. Material examined Holotype of Brachypelmides klaasi (by original designation): MEXICO: Nayarit: ♀, close to Tepic, K. Böhme (SMF 40599) (only microscope slide with spermatheca). Other material: MEXICO: 1 ♀, without more data (SMF 58101–84) (only microscope slide with spermatheca); 1 ♂, without more data, Böhme (SMF 38044); Jalisco: 2 ♂, 1 ♀, Mpio. Tomatlán, 6/XII/2012, D. Ortiz, E. Goyer, E. Hijmensen (CNAN-Ar003333, CNAN-Ar003341, CNAN-Ar007831); 1 ♂, Mpio. Cihuatlán, 4/XII/2013, D. Ortiz, D. Barrales, G. Contreras (CNAN-Ar007160); 1 ♀, Mpio. La Huerta, Reserva Chamela, 6/XII/2013, D. Ortíz, E. Goyer, E. Hijmensen (CNAN-Ar007879); 1 ♂, Mpio. La Huerta, Reserva Chamela, 8–20/XI/2014, W. Maddison (CNAN-Ar007857); 1 ♂, Mpio. La Huerta, Reserva Chamela, 18/V/1981, A. Pescador (CNAN-Ar003432); 1 ♂, donation received from private collection of J. Mendoza (CNAN-Ar003344); Colima: 1 ♂, Mpio. Manzanillo, 4/XII/2013, D. Ortiz, D. Barrales, G. Contreras (CNAN-Ar007162); 1 ♂, Mpio. Colima, 30/XI/2012, D. Ortiz, E. Goyer, E. Hijmensen (CNAN-Ar007845). Diagnosis Brachypelma klaasi can be distinguished from all other known Brachypelma species in the coloration of the legs with pinkish colour on the tibiae and metatarsi (Fig. 36I–L). It also differs in the shape of the genitalia in both sexes with palpal bulb almost straight, tapering and lacking the typical spoon-shape (Fig. 30C–D). The prolateral superior keel reduced, directed retrolaterally and slightly extended to backward (Fig. 30C). Apical keel thin and reduced, shorter than prolateral superior keel (Fig. 30B). Lacking prolateral inferior keel. Also differs by the divided spermatheca widely separated, with spermathecal baseplate oblong. Ventral face smooth (Fig. 31D–F). Brachypelma klaasi is identified by possesing the following character combination: male palpal bulb with embolus almost straight, tapering and without typycal spoon-shape due to the reduced apical keel. Prolateral superior keel reduced, apical keel thin and reduced, shorter than prolateral superior keel. Embolus tip directed to retrolateral. Embolus similar in length to tegulum (Fig. 30A–D). Spermatheca separated with semitriangular receptacles. Spermathecal baseplate divided, oblong; twice wider than its height (Fig. 31D–F). Carapace of both sexes black (Fig. 32A–C). Redescription Male (CNAN-Ar003333) (Figs 29, 30, 32A): Body length 48.60 (not including chelicerae and spinnerets), carapace 22.82 length, 22.17 width. Caput not markedly elevated; fovea straight, 4.15 wide (Fig. 29A). Eyes: anterior eye row procurved, posterior eye row recurved. Eye sizes and interocular distances: AME 0.55; ALE 0.80; PME 0.40; PLE 0.70; AME–AME 0.55; AME–ALE 0.20; PME–PME 1.20; PME–PLE 0.05; ALE–PLE 0.23. Ocular tubercle width 2.80, length 2.55; clypeus 0.25 (Fig. 29D). Labium length 3.05, width 3.55; with 98 cuspules. Maxilla inner corner with approximately 175 (left) and 171 (right) cuspules. Cheliceral promargin with 11 (left) and 12 (right) teeth (proximal to distal: first–second large, third medium, fourth large, fifth small, sixth–11th large; first large, second small, third large, fourth small, fifth large, sixth small, seventh–12th large). Sternum length 10.80, width 10.04. Sigilla oval, second to third pairs hardly visible, posterior pair once its length from the margin (Fig 29B). Leg formula: IV, I, II, III. Length of legs and palpal segments (femur, patella, tibia, metatarsus, tarsus, total): I: 18.95, 10.29, 14.84, 15.15, 10.72, 69.95; II: 17.25, 9.79, 13.15, 14.09, 10.07, 64.35; III: 16.11, 8.27, 11.83, 15.10, 10.28, 61.59; IV: 18.89, 9.38, 15.53, 18.32, 10.63, 72.75. Palp: 12.28, 7.47, 11.92, -, 5.71, 37.38. Spinnerets: PMS, 2.40 long, 1.15 apart; PLS, 4.05 basal, 2.35 middle, 3.60 distal. Tarsi I–IV entirely scopulated. Metatarsus I entirely scopulated, II scopulated 90%, III scopulated 60% distally, IV scopulated 40% distally. Tibia I with two tibial apophyses normally developed, which originate from a common base. Prolateral apophysis with inner spine half its length; retrolateral apophysis tapering throughout its length, apex slightly curved to prolateral (Fig. 29F, G). Metatarsus I curved (Fig. 29E). Stridulatory setae: with claviform stridulating setae on palp trochanter and femur retrolateral face; leg I trochanter and femur prolateral face. Chaetotaxy (left side): femora none; patellae none; tibiae II 1v; III 3v; IV 2v; palp 1v; metatarsi I 1v; II 3v; III 1p, 5v; IV 9v. Palp: embolus almost straight, tapering and lacking the typical spoon-shape due to the reduced apical keel. Prolateral superior keel reduced, directed retrolaterally and slightly extended to backward; prolateral inferior keel absent. Apical keel thin and reduce, shorter than prolateral superior keel. Opening of the embolus is on the prolateral side, just behind the opening is located a concavity which delimits the apical keel boundary from the remaining part of the embolus. Embolus apex slightly curved to retrolateral. Embolus similar in length as tegulum (Fig. 30A–D). Urticating setae: types I and III arranged in one dorsoposterior patch, black in colour. Type III are located in an oval dorsomedial area extended to posterior. Type I surround the type III area, with intermediates between type III and I in transition areas (Fig. 29C). Female (CNAN-Ar007831) (Figs 31, 32B): Body length 63.26 (not including chelicerae and spinnerets), carapace 27.65 length, 24.27 width. Caput not markedly elevated; fovea straight, 6.40 wide (Figs 31A). Eyes: anterior eye row procurved, posterior eye row recurved. Eye sizes and interocular distances: AME 0.57; ALE 0.63; PME 0.43; PLE 0.67; AME–AME 0.50; AME–ALE 0.33; PME–PME 1.33; PME–PLE 0.07; ALE–PLE 0.07. Ocular tubercle width 2.80, length 2.33; clypeus 0.47. Labium length 3.65, width 5.25; with 94 cuspules. Maxilla inner corner with approximately 68 (left)–58 (right) cuspules. Cheliceral promargin with 10 (left)–11 (right) teeth. Sternum length 11.60, width 10.63. Sigilla oval, first to third pairs hardly visible; posterior pair once its length from the margin (Fig. 31B). Leg formula: IV, I, II, III. Length of legs and palpal segments (femur, patella, tibia, metatarsus, tarsus, total): I: 16.86, 10.39, 12.94, 12.74, 8.92, 61.85; II: 15.61, 10.06, 11.28, 12.49, 9.04, 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, 8.02, 9.09, -, 9.29, 38.95. Spinnerets: PMS, 2.80 long, 3.65 apart; PLS, 5.50 basal, 4.15 middle, 4.75 distal. Tarsi I–IV entirely scopulated. Metatarsus I entirely scopulated, II scopulated 80%, III scopulated 70% distally, IV scopulated 50% distally. Stridulatory setae: with claviform stridulating setae on palp trochanter and femur retrolateral face; leg I trochanter and femur prolateral face. Chaetotaxy (left side): femora none; patellae none; tibiae I 2v; II 2v; III 3v; IV 2v; palp 1p, 5v; metatarsi I 3v; II 4v; III 5v; IV 1p, 14v, 1r. Genitalia: spermatheca separated with subtriangular receptacles strongly sclerotized, each receptacle almost the same height as width. Spermatheca baseplate divided and widely separated, each baseplate oblong twice wider than high (Fig. 31D). Baseplate division can vary in length (Fig. 31E, F). Urticating setae: types I and III arranged in one dorsoposterior patch, black in colour. Type III are located in an oval dorsomedial area extended to posterior. Type I surround the type III area, with intermediates between type III and I in transition areas (Fig. 31C). Colour pattern: In live specimens, adults of both sexes have the carapace black (pantone process black c) with zinnwaldite colour (pantone 489c) around the border; despite the black carapace, the radiating thoracic sulci are clearly seen (Fig. 32A–C); chelicerae dorsally brown colour (pantone 7533c) with some large setae French beige colour (pantone 4715c); ventral coxae, labium, maxillae and sternum black pearl colour (pantone black 6c); abdomen dorsally black with corvette colour setae (pantone 720c), ventrally black pearl colour. Legs: femora black pearl; patellae black pearl with scattered setae shilo colour (pantone 488c); tibiae and metatarsi mandys pink colour (pantone 473c); tarsi black (pantone black c); palpi: femora black pearl; patellae, tibia and tarsi black pearl with scattered setae shilo colour (Fig. 36I–L). With juveniles of same pattern but paler in colour. Distribution and habitat: Brachypelma klaasi is known from the Pacific Coast region of Jalisco on the western side of Sierra Madre Occidental, with small populations in Colima and southern Nayarit (Figs 2, 38). It occurs in drier coastal thorn scrub, deciduous forest and higher elevations oak forest (Fig. 32D). It is a fossorial species whose modified or self-excavated burrows, can be found under large rocks, under thorny thickets, tree roots or burrows on hillsides, in both forested and moderately disturbed areas. Burrows do not have any silk around the entrance to indicate there is a spider inside. This species is sympatric (overlapping in range) with Brachypelma emilia in south-western Nayarit and with Brachypelma hamorii in Colima. Brachypelma smithi (F. O. Pickard-Cambridge, 1897) (Figs 2, 33, 34, 35A, C, 37M–P, 38) Figure 33. Open in new tabDownload slide 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 33. Open in new tabDownload slide 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. Open in new tabDownload slide 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. Figure 34. Open in new tabDownload slide 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. Figure 35. Open in new tabDownload slide 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. Figure 35. Open in new tabDownload slide 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. Figure 36. Open in new tabDownload slide 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 36. Open in new tabDownload slide 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. Open in new tabDownload slide 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. Figure 37. Open in new tabDownload slide 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. Figure 38. Open in new tabDownload slide Geographic distribution of the genus Brachypelma from published records and specimens collected or examined for this study. Figure 38. Open in new tabDownload slide Geographic distribution of the genus Brachypelma from published records and specimens collected or examined for this study. Eurypelma smithi F. O. Pickard-Cambridge, 1897: 20, plate 1, fig. 4, D female. Brachypelma smithi (F. O. Pickard-Cambridge, 1897): Pocock (1903: 103); Locht et al. (1999: 198, fig. 5), male; Mendoza & Francke (2017: 162, figs 3–22, 42–45, 50–51, 53–56), male and female. Brachypelma annitha Tesmoingt, Cléton & Verdez, 1997a: 9, plates 1–6, D male and female; Tesmoingt et al. (1997b: 2, plates 7–8, 11), male; Peters (2000: 64, figs 205–207), male and female; Peters (2003: 108, figs 428, 435–436), male and female. Mendoza & Francke (2017: 162) considered this as a junior synonym of B. smithi). Material examined Holotype of Brachypelma smithi: MEXICO: Guerrero: Juvenile ♂ (labelled as ♀), Dos Arroyos. H. H. Smith (BMNH 1143; also labelled BM1898.12.24.33). Other material: MEXICO: Guerrero: 1 ♂, 2 ♀, Mpio. Acapulco, Dos Arroyos. 13/XII/2013, J. Mendoza (CNAN-Ar007146, CNAN-Ar007143, CNAN-Ar007144); 1♂, Mpio. Acapulco, 04/I/1965, E. Rivapalacios (CNAN-Ar003086); 1 ♂, Mpio. Atoyac, 27/II/1984, J. G. Julio (CNAN-Ar003434); 1 ♀, Mpio. Acapulco, 10/VII/1979, M. Adams (CNAN-Ar003435); 1 ♂, Mpio. Acapulco, without additional data (CNAN-Ar003594); 2 ♀, donation received from private collection of J. Mendoza (identified in pet trade as B. annitha) (CNAN-Ar004131, CNAN-Ar003611); 3 ♂, Mpio. Coyuca de Benítez, 29/VIII/2015, J. Mendoza, R. Ramírez (CNAN-Ar010281, CNAN-Ar010275); 1 ♂, 1 ♀, Mpio. Acapulco, El Quemado 14/X/2014, A. Ortega (CNAN-Ar007897, CNAN-Ar007896); 1 ♀, Mpio. Josue Azueta, 24/VIII/2015, D. Ortiz, J. Baldazo (CNAN-Ar007904). Diagnosis Brachypelma smithi can be distinguished from all other known Brachypelma species (except B. hamorii) by the coloration of the carapace and legs, consisting of red-orange coloration on the patellae, tibiae and metatarsi (Fig. 37M–P), as well as orange-black starburst striations on the carapace in general (Fig. 34D). The shape of the genitalia also differs in both sexes with the palpal bulb being straight and broad (Fig. 33C–D) and the spermatheca shaped like a trapezoid (Fig. 33E–H). It differs from B. hamorii in the straight palpal bulb having a broad spoon-shape, a wider apical keel and the prolateral superior keel is not as elevated (Fig. 33A–D). It also differs in the spermatheca with the spermathecal baseplate being divided and subtriangular (Fig. 33E–H). The spermathecal ventral face is also striated, whereas it is smooth in B. hamorii (Fig. 33F). Although similar in coloration, B. smithi differs from B. hamorii by the patellae flame pattern brightly coloured (Fig 37M–P). The sides of the patellae are yellowish-orange rather than black as is typical in B. hamorii (Fig. 35C). The lateral setae are yellowish throughout the length of legs (Fig. 37M–P), whereas in B. hamorii the same are whitish, providing greater contrast with darker areas (Fig. 37I–L). In addition, B. smithi does not tend to show the longitudinal cheliceral lines of lighting setae typically exhibited by B. hamorii (Fig. 35A). Carapace of B. smithi females can be black bordered with orange (Fig. 34F), with a black radiating pattern bordered with orange (Fig. 34D) or, almost completely orange with black around the ocular area (Fig. 34E). Males normally have orange carapace (Fig. 34C), rare variants of males may have also an orange carapace with scant black around ocular area (Fig. 34A) or a black carapace bordered with orange (Fig. 34B). Brachypelma smithi is identified by possessing the following character combination: male palpal bulb with broad spoon-like embolus, prolateral superior keel normally developed, thin and directed retrolaterally, prolateral inferior keel weakly developed, better seen dorsally, apical keel strongly developed, wider in the middle (Fig. 33C–D). Embolus straight and wide throughout its length, one and half times longer than tegulum (Fig. 33A–B). Spermatheca semicircular or trapezoidal (Fig. 33E–H); ventral face striated (Fig. 33F). Spermathecal baseplate divided, subtriangular; four times wider than its height (Fig. 33E–H). Carapace of adult male typically orange in colour (Fig. 34C), whereas in females almost black with orange around the border, with black striated pattern and orange around or almost orange with only black caput area (Fig. 34D–F). Note: Redescription of this species was made by Mendoza & Francke (2017). 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. (Figs 39–44) urn:lsid:zoobank.org:act:EEEC09EE-B64B-4F7E-83A0-D174637009AC Figure 39. Open in new tabDownload slide 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 39. Open in new tabDownload slide 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. Open in new tabDownload slide 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). Figure 40. Open in new tabDownload slide 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). Figure 41. Open in new tabDownload slide 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 41. Open in new tabDownload slide 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. Open in new tabDownload slide 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. Figure 42. Open in new tabDownload slide 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. Figure 43. Open in new tabDownload slide 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 43. Open in new tabDownload slide 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. Open in new tabDownload slide 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. Figure 44. Open in new tabDownload slide 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. Type species: Eurypelma vagansAusserer, 1875, herein designated. Species included: Tliltocatl albopilosum (Valerio, 1980), comb. nov., Tliltocatl epicureanum (Chamberlin, 1925), comb. nov., Tliltocatl kahlenbergi (Rudloff, 2008), comb. nov., Tliltocatl sabulosum (F. O. Pickard-Cambridge, 1897), comb. nov., Tliltocatl schroederi (Rudloff, 2003), comb. nov., Tliltocatl vagans (Ausserer, 1875), comb. nov., Tliltocatl verdezi (Schmidt, 2003), comb. nov. Diagnosis The new genus Tliltocatl can be distinguished from all other known theraphosinae genera (except Brachypelma) by the following character combination: (1) having just claviform stridulating setae on the prolateral face of leg I trochanter/femur and retrolateral face trochanter of the palp; (2) both sexes possess always urticating setae types I and III – type III are located in the dorsoposterior area and 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 apex (Figs 39C, D, 40C–F), prolateral superior and prolateral inferior keels are at similar height, joined at their distal end and widely separating towards the embolus base (better seen in dorsal position) (Fig. 39F), the prolateral inferior keel is similar in length or longer than the prolateral superior (Fig 39E), the apical keel can extend widely to backwards just as the prolateral inferior keel and usually is broader on its distal half (Figs 39D, E; 40B, D–F); (4) females have a single fused spermatheca, apically narrowed (Figs 39G–H, 41A); (5) spermathecal baseplate absent or slightly developed, poorly sclerotized (Fig. 39H); (6) both sexes lack a plumose pad of setae on leg IV femur; and (7) all tarsi scopulae are undivided. It differs from Brachypelma by the coloration of legs, which are black (Figs 42, 43B–D) or have long, whitish setae (as T. albopilosum, Fig. 43A) in combination with a dark carapace and long red/yellowish setae on abdomen (Figs 42, 43B–D). The shape of genitalia also differs in both sexes with the male palpal bulb apex larger 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 larger than in Brachypelma (Figs 30D, 33D) and wider on distal half (Figs 39B, D, E, 40B, D, E). Embolus is regularly similar in length or longer than the tegulum (Figs 39C–D, 40C–F), whereas in Brachypelma it is shorter (Fig. 10D). It also differs in having spination on the patellae of palps and legs. Females differs in the spermatheca apex inwardly curved and by lacking spermathecal baseplate (Fig. 39G) or poorly sclerotized and widely separated when present (Fig. 39H). Description Carapace regularly as long as wide, caput slightly elevated (Figs 42, 43). Cephalic striae inconspicuous (Fig. 43A). Fovea deep, straight or recurve. Eye tubercle distinct and raised, wider than long. Anterior eye row procurved, posterior eye row recurved. Clypeus narrow or absent. Labium subtrapezoidal, wider than long, with 85–120 cuspules on anterior third centre. Maxilla subrectangular, anterior lobe distinctly produced into conical process, inner corner with 130–220 cuspules. Sternum longer than wide. Anterior pair of sigillae circular and slightly seen, second pair oval enlarged, posterior pair oval or rounded, generally once its length from the sternum margin. Leg formula: I, II, III, IV. Tarsi I–IV fully scopulated. All tarsi scopulae are undivided. Metatarsi I–II fully scopulated. Metatarsus III is 50% distally scopulated and metatarsus IV is 20–40% distally scopulated. The femur of leg III is slightly enlarged but not swollen as in other genera. Femora IV without retrolateral scopulae. Claviform stridulating setae on palp trochanter retrolateral face and in leg I trochanter and femur prolateral face. Patellae of the legs have at least one spine on prolateral or retrolateral side. Posterior lateral spinneret distally elongating, digitiform. Both sexes possess urticating setae type I and type III; Type III are located in the dorsoposterior area and type I surrounding these. Males possess two tibial apophysis, retrolateral apophysis slightly curved in apically. Globous bulb with small subtegulum longer than its height. Embolus flattened (Fig. 39A, B), longer than tegulum (Fig. 39C, D), with prolateral superior and apical keel large, joint at the embolus tip forming a typical spoon-shape (Figs 39C–E, 40C–F). Prolateral inferior keel well developed, parallel to prolateral superior and larger in general (Figs 39D–F, 40A, D, E). Females with a simple undivided/fused spermatheca apically narrowed and medially curved inwardly (Fig. 39G, H). Spermathecal baseplate absent or slightly developed (Fig. 39H). Spermatheca midwidth shorter than its base and well sclerotized (Fig. 39G, H). Most of the species are black with long, red setae on the opisthosoma (except T. albopilosum, T. schroederi and T. verdezi) (Figs 42–43). Juveniles are similar in colour to the adults but paler. Remarks Tliltocatl can be confused by unexperienced people with Sericopelma spp. due to the similar coloration, but they can be easily differentiated from each other, because adult males of Sericopelma lack tibial apophysis, the female spermatheca of Sericopelma is strongly sclerotized (Fig. 41B), also the spermatheca is distinctly swollen on the apex showing a P-shape (better seen laterally). In addition, Sericopelma has a distinct radiating sulcus on the carapace (Gabriel & Longhorn, 2015). Etymology The genus gender is masculine. The name is a noun in apposition comprising the Nahuatl words Tlil, which means ‘black’, and tocatl, which means ‘spider’, referring to the black coloration of species in the genus. Distribution Tliltocatl occurs in Mexico, Guatemala, Belize, Honduras, El Salvador, Nicaragua and Costa Rica (Fig. 44). The species of the genus are found in deciduous forest, evergreen rain forest and grasslands along the Pacific coast, Mexican Gulf and Atlantic Coast. Specimens live inside burrows under flat rocks, fallen logs, sidehills, tree roots and even some species dig burrows in farmlands, gardens or close to flooded land. Species considered nomina dubia Brachypelma andrewi Schmidt, 1992:Schmidt (1992a) described this species based on a single specimen located in the collection of the British 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 the description of B. andrewi comes from the L. Koch collection. Another specimen from the same Koch collection was found in the NHM containing an alleged undescribed specimen. Initially the jar that contained the specimen, of what would later be described as B. andrewi, was labelled as the holotype of Euathlus truculentusAusserer, 1875. This created a confusion that resulted in the genus Brachypelma being listed as a junior synonym of Euathlus, as was proposed by Raven (1985). Smith (1992) redescribed the specimen 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 decision that the genus Brachypelma was synonymous with Euathlus. However, shortly afterwards, Schmidt (1992a) observed that the specimen seen by Raven, and later by Smith, was not the same species as the type 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 was a junior synonym of Euathlus, while Smith, after studying the said Brachypelma, was of the opinion that Raven was right with his synonymizing. He had not noticed that only the body type of the Euathlus type, called Ausserer, deviates considerably from those of the alleged 2ndEuathlus truculentus: e.g. in the real type, the body length is 44 mm, while in the erroneously paratype Brachypelma is 55 mm. This Brachypelma is said to originate from Cuba and has not been mentioned so far – I shared the error with Smith in a letter dated 28 May 1992 and confirmed that he was wrong in the matter of synonymizing and was no longer tenable.’ After this, Schmidt (1992a) named the undescribed Brachypelma as B. andrewi after Andrew Smith. Unfortunately, because of the lack of a useful locality for this species, no new specimens can be collected for comparison. Also, after our visit to the arachnological collection of the NHM, it was not possible to locate the type specimen of Brachypelma andrewi, which is why it is considered lost to date. After reviewing, in detail, the redescription of the specimen made by Smith (1992), it is possible to know that B. andrewi is a species of the ‘red rump complex’ (vagans group) tarantulas (which were transferred to Tliltocatl in this work). So, we propose that this species be transferred as Tliltocatl andrewi comb. nov. However, without being able to make a direct observation of the specimen, it is not possible to correctly identify the species or compare it with those closely related and already described. Additionally, given the lack of data on its distribution, it is not possible to obtain new material. Therefore, based on the loss of the holotype and in its inability to be identified, we propose that the species T. andrewi be considered nomen dubium. Brachypelma aureoceps (Chamberlin, 1917,): This species was described by Chamberlin (1917) based on a single female (RVC43 – examined) from Florida, Tortugas. However, there is no evidence of native tarantulas from Florida (Smith, 1994; West, personal communication). Smith (1994) mentioned that is highly probable that the B. aureoceps specimen was accidentally imported into the Florida Keys. This specimen has morphological features, such as spermatheca without a baseplate, highly notched medially, patellae spination and claviform setae on prolateral trochanter and femur of leg I, showing that it belongs to the genus Tliltocatl. So, we propose that this species be transferred as Tliltocatl aureoceps comb. nov. Nevertheless, this species is similar to other valid species distributed in Mexico (e.g. T. kahlenbergi and T. vagans and cannot be differentiated adequately from these. In such a way that molecular data or accurate morphological features are needed to properly differentiate this species, and without the possibility to collect fresh specimens we consider T. aureoceps a nomen dubium. Citharacanthus alvarezi Estrada-Álvarez et al., 2013: This species was described by Estrada-Álvarez et al. (2013) based on a single specimen referred to as female (CNAN-T01275 – examined) from an uncertain locality in Tuxtla Gutierrez, Chiapas, that was donated by an unknown collector to the Zoológico Miguel Álvarez del Toro (ZooMAT). After examination of the holotype, we were able to determine that it does not belong to the genus Citharacanthus Pocock (1901), and it is not a female, but a subadult male belonging to Tliltocatl. The main evidence of this is the presence of type I and III urticating setae, in combination with stridulatory claviform setae on trochanter and femur I and palp trochanter. It is evident that the specimen is a subadult male just by the presence of the accessory organs. These gland-like structures can be prominent in some groups, such as Tliltocatl, and are often mistaken for paired spermatheca (as was in this case). The male accessory organs do not have direct connection with the gonopore and neither do they present uterus externus as can be observed in a spermatheca. Based on the evidence we proposed to transfer the species as Tliltocatl alvarezi, comb. nov. Unfortunately, subadult males do not show features that could be useful to identify a species. Due to the lack of morphological and/or molecular data that would allow us to make a comparison with the already described known species of Tliltocatl, as well as a more precise collection locality, we propose T. alvarezi to be considered a nomen dubium. The following has been transferred to another group: Stichoplastoris fossorius (Valerio, 1980) comb. nov. (Figs 44, 45) Brachypelma fossoriaValerio, 1980: 271, fig. 25–28, D male and female; Smith (1986: 49, fig. 28h), male; Smith (1987: 49, fig. 28h), male. Brachypelma fossorium: Schmidt (1992: 10, figs 9–12) transferred male and female from Euathlus; Rudloff (2003: 8, figs 26–29), male; Schmidt (2003: 153, figs 278–281), male and female. Misidentification:Peters (2000: 70, fig. 228), male; Peters (2003: 121, figs 486, 488–489), male and female. Peters (2000) and Peters (2003) show as B. fossorium an unidentified species of Tliltocatl, possibly pet-trade material, making it difficult to identify. Holotype and paratype of Brachypelma fossorium: COSTA RICA: Guanacaste: 1♂, Filadelfia. Col. E. Herrera (UCR-238) examined. 1♀, Liberia, finca Santo Tomas, without more data (UCR-126) not examined. Other material examined: COSTA RICA: Guanacaste: 2♀, Canton Santa Cruz, Huacas, without more data. Remarks Originally Valerio (1980) placed this species in the genus Brachypelma based only in the shape of the male palpal bulb and female genitalia. However, after the revision of the holotype and based in the cladistic analysis we conclude that this species does not belong to Brachypelma or Tliltocatl. This species has a short embolus with presence of prolateral superior, prolateral inferior and apical keels, but with a different configuration from that observed in Brachypelma and Tliltocatl (Fig. 45C–D). The keel arrangement is more similar to Stichoplastoris Rudloff (1997). The urticating setae present in B. fossorium are only type I, whereas Brachypelma and Tliltocatl possesses type I and III. Also, B. fossorium has the scopula IV divided by a strong band of setae, whereas Brachypelma and Tliltocatl have all their scopulae entire. These characteristics fit those found in some species of the genus Stichoplastoris [e.g. S. elusinus (Valerio, 1980), S. obelix (Valerio, 1980)]. The weakest point of the kinship with Stichoplastoris would be the general shape of the male palpal bulb that is wide in the apex, and the spermatheca of the female that is unilobular, whereas in Stichoplastoris the bulb looks thinner apically and the spermatheca is divided (although in species like S. obelix it is wider at its base). Despite this, there are similar examples in genera such as Brachypelma in which species such as B. klaasi and B. albiceps have thinner palpal bulbs and a divided spermatheca, unlike the rest of the species in the genus that have wider bulbs apically and fused spermatheca. General body shape and legs of males and females are not as thick as in Brachypelma and Tliltocatl (Fig. 45A–B). We, therefore, propose, based on the similarities indicated above, that B. fossorium be transferred to the genus Stichoplastoris creating the new combination Stichoplastoris fossorius. Figure 45. Open in new tabDownload slide 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. Figure 45. Open in new tabDownload slide 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. DISCUSSION Non-monophyly of Brachypelma and the use of barcodes Simon (1891) described several diagnostic characters for Brachypelma such as: femur IV lacking inner scopula (no dense pad of plumose setae), presence of a distinct scopula on the metatarsus IV, palpal bulb narrowly piriform, but with the apex wide and attenuated, very much compressed and obtuse. Pocock (1903) also distinguished the plumose setae on the prolateral face of leg I trochanter/femur and retrolateral face of the palp. These key features have been supported by subsequent authors as diagnostic for Brachypelma, as also are, no tarsal division by strong setae, the male palpal bulb distally wide and flat (commonly mentioned as spoon-shaped), two unequal apophyses on male tibia I, and the spermatheca regularly fused, semi-fused or with two separated wide lobes (Schmidt, 1992a, 1992b; Smith, 1993, 1994; Pérez Miles et al., 1996; Locht et al., 1999; Gabriel & Longhorn, 2015; Mendoza & Francke, 2017). Smith (1994) adequately explains the taxonomic history of Brachypelma, which has been complicated and has multiple taxonomic changes (Simon, 1891, 1903; Pickard-Cambridge, 1897; Roewer, 1942; Valerio, 1980; Raven, 1985; Schmidt, 1992). In the early 1990s, new Brachypelma species were described based mainly on material collected for the pet trade (e.g. B. auratum, B. baumgarteni, B. boehmei, B. klaasi, B. ruhnaui, B. hamorii and B. annitha). Although there were evident colour differences among the species from the Mexican Pacific Coast and Mexico–south-west Central America, no study had been done in depth to prove the monophyly of the genus. Rudloff (2003) was the first to propose a designation of complexes of species for Brachypelma based on colour characteristics and with no intention of making nomenlatural changes. He postulated a group called ‘vagans’ formed by the species with dark to black base colour on them (B. albopilosum, B. angustum, B. aureoceps, B. embrithes, B. epicureanum, B. fossorium, B. sabulosum, B. schroederi and B. vagans) and a group named ‘emilia’ formed by the species with red/orange banded legs (B. annitha, B. auratum, B. baumgarteni, B. boehmei, B. emilia, B. klaasi and B. smithi). He also said that the species B. ruhnaui (currently B. albiceps) cannot be classified in this scheme and that it was possibly more closely related to Aphonopelma than to Brachypelma. Petersen et al. (2007) developed a method to obtain mtDNA from Brachypelma spp. using exuvia. Their results show a phylogeny based on a cytochrome oxidase 1 (COI) gene fragment trimmed to 205bp where two Brachypelma subgroups corresponding to the ‘red rump vagans group’ and the ‘red leg emilia group’ were recovered, but were simply (mis)-grouped together, because no other related genera were used for comparison. Mendoza & Francke (2017) used molecular data for a phylogenetic analysis with COI data to clarify some relationships among red knee species and revised their nomenclature, but only included some species from the ‘red leg emilia group’, excluding the type species B. emilia, leaving the monophyly of the genus untested. Recently, Turner et al. (2018) presented a mtDNA gene tree of tarantula spiders based on the mitochondrial 16S-tRNA (leu)-NDI gene region as an initial hypothesis to clarify some taxonomic relationships of the subfamily Theraphosinae. Their recovered phylogeny in both Bayesian and maximum likelihood analyses strongly supported the non-monophyly of both Brachypelma and Aphonopelma, indicating that neither of these genera, as currently recognized, are monophyletic. This result is congruent with our results based on COI, so it is another strong support for the non-monophyly of Brachypelma. We agree with the results of Turner et al. (2018) and proposed the ‘red leg group’, which contains the type species B. emilia, as Brachypelma s.s., while the ‘red rump group’ is considered as a different genus here described as Tliltocatl with the designation of T. vagans as its type species. We also performed a morphological character-based analysis of the genus Brachypelma, using parsimony to test the monophyly of the genus and to reconcile morphological and molecular characters. Our best tree is based on the strict consensus of two most parsimonious trees obtained from the parsimony analysis of 103 characters. In this phylogeny, the ‘red leg group’ and the ‘red rump group’ were also recovered as separated genera, giving additional evidence for the non-monophyly of Brachypelma. However, morphological data in Theraphosidae tends to be homoplastic due to similarities, simplicity in sexual structures and conserved characters (Raven, 1985; Goloboff, 1993; Pérez-Miles et al., 1996; Pérez-Miles, 2000; Bertani, 2001; Bond & Opell, 2002; Bond & Hedin, 2006; Hedin & Bond, 2006; West et al., 2008; Hendrixson & Bond, 2009; Bond et al., 2012; Guadanucci, 2014; Hamilton et al., 2014, 2016; Perafán & Pérez-Miles, 2014; Ortiz & Francke, 2016; Fukushima & Bertani, 2017). Despite this shortcoming, both Brachypelma and Tliltocatl have diagnostic characteristics and synapomorphies that differentiate them from each other, the most evident, in addition to their differences in coloration, are the prolateral inferior keel absent or slightly developed and restricted to the bulb apical, the spermatheca baseplate well developed and well sclerotized in Brachypelma; whereas in Tliltocatl the prolateral inferior keel is always present and is widely extended backwards, the spermatheca do not have a baseplate or only present a small area slightly sclerotized. Within the taxonomy of tarantulas, the count of spines in the legs or the presence/absence of them does not usually present great taxonomic relevance due to its variability between individuals of the same species. However, our analysis of these characters showed differences between Brachypelma and Tliltocatl, as Tliltocatl always presents spinations on all the patellae, whereas Brachypelma never shows this characteristic, thus we consider this is a reliable character to differentiate both genera. Brachypelma fossorium is a species that is not grouped with either Brachypelma or Tliltocatl in both the morphological and molecular phylogenies (Figs 3, 4). This species was originally described by Valerio (1980) as Brachypelma based solely on the shape of the male palpal bulb and female genitalia. However, as is shown in our results, we conclude that this species does not belong to Brachypelma or Tliltocatl. Despite the similar shape of the genitalia, this species does not possess the diagnostic features of either genus: it has a short embolus with different arrangement of keels from that observed in Brachypelma or Tliltocatl. Also, the urticating setae present in B. fossorium are only type I, whereas Brachypelma and Tliltocatl possess types I and III. Brachypelma fossorium has scopula IV divided by a strong band of setae, whereas Brachypelma and Tliltocatl have all their scopulae entire. So, based on this evidence, and other arguments exposed in our morphological revision of the species above, we are certain that this species does not belong in either Brachypelma or Tliltocatl, and we transfer it to Stichoplastoris. Mendoza & Francke (2017) concluded that barcode marker COI in Brachypelma proved to be sufficient for correct species identification. They considered it also as a useful tool in preventing black market trade and in providing better strategies to reintroduce tarantulas into the correct distribution areas. DNA barcoding is a useful technique that, together with morphology, field observations and museum collections, allow for better definition and delimitation of species (Scotland et al., 2003; Chen et al., 2011; Slowik & Blagoev, 2012; Chan et al., 2014; Hendrixson et al., 2015; Pante et al., 2014). However, its use to resolve phylogenies is limited. Hamilton et al. (2016) mentioned that the limitations of mtDNA used for phylogeny are gene tree/species tree incongruence and the haploid, non-recombining nature of the molecule, with COI representing only one particular genealogy out of all possible ones in a genome. These limitations are shown in our ML tree, with the strong support for the confidence of each genus, but with low resolution on some inner clades in both Brachypelma and Tliltocatl (Fig. 4). In the case of Brachypelma, the most problematic is the relationship of B. baumgarteni, B. boehmei and B. auratum, just as is seen also in Mendoza & Francke (2017), where all these species were well-supported but the clade that includes them was collapsed. In our phylogeny, something similar happens with B. auratum and B. baumgarteni as sister-species but with low support and with poor resolution in B. boehmei (Fig. 4). For Tliltocatl, the ML tree shows no resolution in the position of T. verdezi and T. kahlenbergi, both sister of a clade formed by T. epicureanum, T. albopilosum, T. sabulosum and T. vagans. In this clade, T. epicureanum appears as a sister-species of the other three; although the clade, which shows T. vagans as sister to T. sabulosum and T. albopilosum, has no support (Fig. 4). The morphology-based tree has some similarities with the one obtained with ML for COI, showing B. emilia as the sister-species of all other Brachypelma, followed by B. klaasi and B. albiceps, which are supported by a combination of five homoplastic characters but not by bootstrap values (Fig. 3; char35, 55, 98, 100, 101). Three of these characters are related to the spermatheca, which have similarities because they are the only species with divided spermatheca. So, this is the main reason why the three species are grouped as sister-species in the morphological phylogeny, unlike the molecular where they are separated (Figs. 3, 4). The same occurs in the case of the group formed by B. baumgarteni and B. hamorii, since these species are supported by two synapomorphies (char4, 35) and six homoplastic characters (char6, 10, 13, 79, 82, 83); the synapomorphies are the coloration of the chelicerae and the similar coloration on the tibiae. Three of the six homoplastic characters are related to the male palpal bulb, since both species have a similar shape with the apical keel slightly developed and the prolateral superior keel wide and short (Fig. 3). In contrast, in the ML tree, B. baumgarteni and B. hamorii are genetically distant, with B. hamorii followed by B. smithi and this is followed by a group formed by B. boehmei, B. baumgarteni and B. auratum (Fig. 4). For the genus Tliltocatl, there are differences between both topologies. Where in the morphological tree T. albopilosum and T. schroederi are shown as sister-species supported by only three homoplastic characters and no bootstrap support (char43, 62, 71). All the characters are related to the number of spines in patellae II and III and metatarsus III (Fig. 3). The ML tree shows T. schroederi as the sister-species of all other Tliltocatl, whereas T. albopilosum is in an unresolved group with T. sabulosum and T. vagans. The other species resolution is more similar in both topologies, with T. verdezi and T. kahlenbergi with no resolution and as sister-species of an inner clade formed by T. epicureanum, T. vagans and T. sabulosum in the morphological tree, and with the inclusion of T. albopilosum in the molecular tree as was shown above (Figs 3, 4). These discrepancies between morphological and genetic phylogenies are apparently given by the grouping of some species based mainly on the similarity of their reproductive organs, which are probably convergent. Also, it is possible that the inclusion of other outgroups can help to resolve better the morphological hypothesis. We suggest that future studies must explore the use of different molecular markers, such as nuclear ones, which are more conservative and could help to resolve the inner relationships of the species. Also, the use of next-generation sequencing methods can help to resolve phylogenetic problems, evolutionary concerns or even to help in conservation strategies (Turner et al., 2018). Further, the Inter Simple Sequence Repeats (ISSR) used by Machkour-M’Rabet et al. (2009) as a molecular marker for the study of genetic diversity in populations of tarantulas proved to be a useful tool to learn about the intraspecific variation in these organisms, which can be a valuable resource to evaluate the population structure of priority tarantulas such as Brachypelma and Tliltocatl. Conservation issues As was mentioned by Turner et al. (2018), the non-monophyly of Brachypelma with some species being transferred to another genus has immediate implications for conservation. The main concern for those authors lies in the state of conservation that could be affected after the removal of the ‘red rump’ species complex from Brachypelma. This is because, as was observed, we still know very little about their diversity, distribution, ecological characteristics, habitat preferences, reproductive success and how much they are affected by habitat loss and predation (Reichling, 2000; Longhorn, 2002; Machkour-M’Rabet et al., 2005, 2007, 2011, 2012, 2015, 2017; Shillington & McEwen, 2006; Dor et al., 2008; Dor & Hénaut, 2011, 2012, 2013; Vilchis-Nestor et al., 2013; Hénaut et al., 2015). Nevertheless, CITES is aware of possible changes in nomenclature in the taxonomic groups included for its regulation and in its resolution Conf. 12.11 revised in the COP17 held in Johannesburg, it is mentioned that ‘whenever a change in the name of a taxon included in the Appendices is proposed, the Secretariat, in consultation with the Animals or Plants Committee, determine whether this change would alter the scope of protection for fauna or flora under the Convention’. This previous statement that in case where the scope of a taxon is redefined, the Animals or Plants Committee shall evaluate whether acceptance of the taxonomic change would cause additional species to be included in the appendices or listed species to be deleted from the appendices. Thus, if that is the case, the Depositary Government should be requested to submit a proposal to amend the appendices in accordance with the recommendation of the Animals or Plants Committee, so that the original intent of the listing is retained. This means that even if species that belong to the ‘red rump’ group are transferred from Brachypelma into the genus Tliltocatl, these species will not lose their protective status under CITES appendix II. However, this protective clause only concerns international trade. It is equally important to know if the taxonomic change of these species will affect their protection status in Mexico. There, it is the Ley General de Vida Silvestre (LGVS) who regulates the sustainable use, conservation and management of native wild animals and plants. It regulates the protection of species or populations that are at risk (SEMARNAT, 2000). The LGVS establishes the national policy for wildlife protection and sustainable use via the SUMA programme and the Mexican Official Standard NOM059-SEMARNAT-2010 (NOM-059) on Mexican species at risk. In addition, the LGVS regulates the creation of UMAs. To change the status of the species protected by law, it is necessary to submit a petition for evaluation with information about trade, conservation and population status. Thus, despite the nomenclatural changes proposed above, fortunately all the species formerly known as Brachypelma will remain protected in Mexico as well. In fact, a particular interest has recently emerged from the Mexican Government to protect all tarantulas and regulate their trade. In 2015, the governments of Canada, Mexico and the United States initiated a collaborative project through the Commission for Environmental Cooperation (CEC) to strengthen the conservation and sustainable 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 tarantulas. This action plan includes information on 16 priority tarantula species, comprising one from the genus Aphonopelma and 15 from Brachypelma s.l. The information was compiled for the species as a group and included: the impact of trade on conservation and livelihoods, and identification challenges for CITES’ enforcement. Currently, there are insufficient population data available for Mexican tarantulas, especially those restricted to small geographical areas that are particularly vulnerable to overexploitation (Reichling, 2003). Without such information, it would be very difficult to make sure that the impact of exporting adult or sub-adult specimens would not be detrimental to the species survival in the wild, unless there was compelling information available to show that the specimens in question were captive-bred. Preliminary field studies indicate that B. baumgarteni, B. boehmei and B. hamorii have small zoogeographical ranges and are sensitive to habitat disruption. Therefore, at least these three species should not be considered for direct capture and export until more research has been conducted on their viability (CEC, 2017). At sustainable levels of exploitation, both wildlife and people can benefit from legal trade. Granting local people an economic stake in wildlife management provides the best incentive for careful stewardship of species and habitats (Carey, 1999; Dickinson, 2002). Although, DNA barcodes are not good enough to either capture the dynamic processes of evolution or to reflect the precise phylogenetic relationships within a given group, they could provide legislators with a framework of data when enacting protection laws. This could translate into greater success in prosecuting those involved in the illegal pet-trade. It is also generally agreed that mtDNA alone is not sufficient for precise species delimitation. However, it can be useful for identification if the findings correspond with morphological evidence (Petersen et al., 2007). Because of this, we will create a genetic library of priority tarantulas (Brachypelma and Tliltocatl) as part of the Wildlife Barcode Project in Mexico (BWPM, 2014), to be used as a reference for the authorities responsible for species conservation. Juvenile tarantulas are very difficult or impossible to identify, even to genus. Therefore, the regulation of the pet-trade of juvenile tarantulas will benefit tremendously by using molecular barcode markers to easily and accurately identify them. It is essential to explore other non-subtractive economic endeavours, such as tourism, wildlife observation or sale of captive-bred animals, to provide alternative incomes for local communities and to avoid further damage to wildlife populations. [Version of permanent record, published online 4 November 2019; http://zoobank.orgurn:lsid:zoobank.org:pub:E4D09A17-444F-45A0-95DB-059ECA175569] ACKNOWLEDGEMENTS This publication is a requirement for the PhD graduate training programme of Jorge Mendoza at the Posgrado en Ciencias Biológicas of the Universidad Nacional Autónoma de MEXICO (UNAM). JM thanks the Instituto de Biología (IBUNAM) and the Posgrado en Ciencias Biológicas of the UNAM for the programme received. JM also thanks the Consejo Nacional de Ciencia y Tecnología (CONACYT) for providing financial support for his post-graduate studies, as well as the grants required for a research stay at the Muséum National d´Histoire Naturelle in Paris, France (MNHNP), the Natural History Museum in London, United Kingdom (NHM), the Senckenberg Naturmuseum in Frankfurt, Germany (SNMF) and the Museo de Zoología de Costa Rica (MZUCR). We thank Janet Beccaloni, Christine Rollard, Elise-Anne Leguin, Peter Jäger, Julia Altmann and Gilbert Barrantes for providing access to types and historical specimens. Stuart Longhorn and Ray Gabriel are thanked for their assistance and hospitality during JM’s research stay in Europe. We thank Virginia Leon for her support for travel to Europe and Andrea Jiménez for her assistance during molecular work. JM thanks the members of the laboratory of arachnology (CNAN) for their support and assistance on field trips. JM thanks R. Ramírez, J. Hinojosa, E. Hijmensen and E. Goyer for their assistance during field trips. We also thank the editor and reviewers for their critical reading and valuable comments. The specimens were collected under scientific collection permit FAUT-0175, authorized by SEMARNAT for Oscar F. Francke. The present project was supported by MEXBOL of CONACYT (project 251085) and the Barcode of Wildlife Project in Mexico, sponsored by Google´s Global Impact Awards. 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 BrachypelmaSimon 1891 and seven species in the Tliltocatl genus. P. cambridgei, E. campestratus, Lasiodora parahybana and Xenesthis immanis was used from GenBank. REFERENCES Agnarsson I , Miller JA . 2008 . Is ACCTRAN better than DELTRAN? Cladistics 24 : 1032 – 1038 . Google Scholar Crossref Search ADS WorldCat Arisqueta-Chablé C , Manrique-Saide P , Pinkus MA , Melendez B . 2009 . Noteworthy records of Brachypelma (Araneae: Theraphosidae) from Peninsula of Yucatan, Mexico . Entomological News 120 : 566 – 569 . Google Scholar Crossref Search ADS WorldCat Arnedo M , Fernández M . 2007 . Mitochondrial markers reveal deep population subdivision in the European protected spider Macrothele calpeiana (Walckenaer, 1805) (Araneae, Hexathelidae) . Conservation Genetics 8 : 1147 – 1162 . Google Scholar Crossref Search ADS WorldCat Ausserer A . 1875 . Zweiter Beitrag zur Kenntnis der Arachniden-Familie der Territelariae Thorell (Mygalidae Autor) . Verhandlungen der Kaiserlich-Königlichen Zoologisch-Botanischen Gesellschaft in Wien 25 : 125 – 206 . WorldCat Baerg WJ . 1958 . The tarantula . London : Fitzgerald Publishing , 85 . Google Preview WorldCat COPAC Barrett R , Hebert P . 2005 . Identifying spiders through DNA barcodes . Canadian Journal of Zoology 83 : 481 – 491 . Google Scholar Crossref Search ADS WorldCat Bertani R . 2000 . Male palpal bulbs and homologous features in Theraphosinae (Araneae, Theraphosidae) . Journal of Arachnology 28 : 29 – 42 . Google Scholar Crossref Search ADS WorldCat Bertani R . 2001 . Revision, cladistic analysis, and zoogeography of Vitalius, Nhandu, and Proshapalopus; with notes on other theraphosinae genera (Araneae, Theraphosidae) . Arquivos de Zoologia 36 : 265 – 356 . WorldCat Bertani R , Guadanucci JP . 2013 . Morphology, evolution and usage of urticating setae by tarantulas (Araneae: Theraphosidae) . Zoologia 30 : 403 – 418 . Google Scholar Crossref Search ADS WorldCat Bertani R , Fukushima CS , Da Silva Junior PI . 2008 . Two new species of Pamphobeteus Pocock 1901 (Araneae: Mygalomorphae: Theraphosidae) from Brazil, with a new type of stridulatory organ . Zootaxa 1826 : 45 – 58 . Google Scholar Crossref Search ADS WorldCat Blagoev G , Hebert P , Adamowicz S , Robinson E . 2009 . Prospects for using DNA barcoding to identify spiders in species-rich genera . ZooKeys 16 : 27 – 46 . Google Scholar Crossref Search ADS WorldCat Blagoev GA , deWaard JR , Ratnasingham S , deWaard SL , Lu L , Robertson J , Telfer AC , Hebert PDN . 2016 . Untangling taxonomy: a DNA barcode reference library for Canadian spiders . Molecular Ecology Resources 16 : 325 – 341 . Google Scholar Crossref Search ADS PubMed WorldCat Bond JE , Hedin M . 2006 . A total evidence assessment of the phylogeny of North American euctenizinae trapdoor spiders (Araneae, Mygalomorphae, Cyrtaucheniidae) using Bayesian inference . Molecular Phylogenetics and Evolution 41 : 70 – 85 . Google Scholar Crossref Search ADS PubMed WorldCat Bond JE , Opell B . 2002 . Phylogeny and taxonomy of the genera of southwestern North American Euctenizinae trapdoor spiders and their relatives (Araneae: Mygalomorphae, Cyrtaucheniidae) . Zoological Journal of the Linnean Society 136 : 487 – 534 . Google Scholar Crossref Search ADS WorldCat Bond JE , Stockman AK . 2008 . An integrative method for delimiting cohesion species: finding the population–species interface in a group of Californian trapdoor spiders with extreme genetic divergence and geographic structuring . Systematic Biology 57 : 628 – 646 . Google Scholar Crossref Search ADS PubMed WorldCat Bond JE , Hendrixson BE , Hamilton CA , Hedin M . 2012 . A reconsideration of the classification of the spider infraorder Mygalomorphae (Arachnida: Araneae) based on three nuclear genes and morphology . PLoS ONE 7 : e38753 . Google Scholar Crossref Search ADS PubMed WorldCat Briscoe AG , Goodacre S , Masta SE , Taylor MI , Arnedo MA , Penney D , Kenny J , Creer S . 2013 . Can long-range PCR be used to amplify genetically divergent mitochondrial genomes for comparative phylogenetics? A case study within spiders (Arthropoda: Araneae) . PLoS ONE 8 : e62404 . Google Scholar Crossref Search ADS PubMed WorldCat BWPM. 2014 . Códigos de barras de la vida silvestre MEXICO . Available online at http://bwp-mex.blogspot.mx/ (accessed 12 January 2018) . Carey JE . 1999 . Improving the efficacy of CITES by providing the proper incentives to protect Endangered Species . Washington University Law Review 77 : 1291 – 1322 . WorldCat CEC. 2017 . Sustainable trade in tarantulas: action plan for North America . Montreal : Commission for Environmental Cooperation , 52 . Google Preview WorldCat COPAC Chamberlin RV . 1917 . New spiders of the family Aviculariidae . Bulletin of the Museum of Comparative Zoology at Harvard College 61 : 25 – 75 . WorldCat Chamberlin RV . 1925 . New North American spiders . Proceedings of the California Academy of Sciences 14 : 105 – 142 . WorldCat Chan A , Chiang L , Hapuarachi H , Tan C , Pang S , Lee R , Lee K , Ng L , Lam-Phua S . 2014 . DNA barcoding: complementing morphological identification of mosquito species in Singapore . Parasites & Vectors 7 : 1 – 12 . Google Scholar Crossref Search ADS PubMed WorldCat Chen J , Li Q , Kong L , Yu H . 2011 . How DNA barcodes complement taxonomy and explore species diversity: the case study of a poorly understood marine fauna . PLoS ONE 6 : e21326 . Google Scholar Crossref Search ADS PubMed WorldCat Dickinson B . 2002 . International conservation treaties, poverty and development: the case of CITES . Natural Resource Perspectives 74 : 1 – 4 . WorldCat Dor A , Hénaut Y . 2011 . Are cannibalism and tarantula predation, factors of the spatial distribution of the wolf spider Lycosa subfusca (Araneae Lycosidae)? Ethology, Ecology and Evolution 23 : 375 – 389 . Google Scholar Crossref Search ADS WorldCat Dor A , Hénaut Y . 2012 . Silk use and spiderling behaviour in the tarantula Brachypelma vagans (Araneae: Theraphosidae) . Acta Zoologica Mexicana 28 : 1 – 12 . WorldCat Dor A , Hénaut Y . 2013 . Importance of body size and hunting strategy during interactions between the redrump tarantula Brachypelma vagans and the wolf spider Lycosa subfusca . Canadian Journal of Zoology 91 : 545 – 553 . Google Scholar Crossref Search ADS WorldCat Dor A , Machkour-M’Rabet S , Legal L , Williams T , Hénaut Y . 2008 . Chemically-mediated intraspecific recognition in the Mexican tarantula Brachypelma vagans . Naturwissenschaften 95 : 1189 – 1193 . Google Scholar Crossref Search ADS PubMed WorldCat Estrada-Alvarez JC . 2014 . New data from mygalomorph spiders (Araneae: Mygalomorphae) of Estado de Mexico, with taxonomic comments about the genus Davus O. Pickard-Cambridge, 1892 . Dugesiana 21 : 55 – 66 . WorldCat Estrada-Álvarez JC , Guadarrama RCA , Martínez OM . 2013 . Nueva especie de Citharacanthus Pocock, 1901 (Theraphosidae: Theraphosinae) para MEXICO . Dugesiana 20 : 63 – 66 . WorldCat Felsenstein J . 1985 . Confidence limits on phylogenies: an approach using Bootstrap . Evolution 39 : 783 – 791 . Google Scholar Crossref Search ADS PubMed WorldCat Fitch WM . 1971 . Toward defining the course of evolution: minimum change for a specific tree topology . Systematic Zoology 20 : 406 – 416 . Google Scholar Crossref Search ADS WorldCat Folmer O , Black M , Hoech W , Lutz R , Vrijenhoeck R . 1994 . DNA primers for amplification of mitochondrial cytochrome c oxidase subunit I from diverse metazoan invertebrates . Molecular Marine Biology and Biotechnology 3 : 294 – 299 . Google Scholar PubMed WorldCat Fukushima CS , Bertani R . 2017 . Taxonomic revision and cladistic analysis of Avicularia Lamarck, 1818 (Araneae, Theraphosidae, Aviculariinae) with description of three new aviculariine genera . ZooKeys 659 : 1 – 185 . Google Scholar Crossref Search ADS WorldCat Fukushima CS , Bertani R , Da Silva P Jr . 2005 . Revision of Cyriocosmus Simon, 1903, with notes on the genus Hapalopus Ausserer, 1875 (Araneae: Theraphosidae) . Zootaxa 846 : 1 – 31 . Google Scholar Crossref Search ADS WorldCat Gabriel R , Longhorn SJ . 2015 . Revised generic placement of Brachypelma embrithes (Chamberlin & Ivie, 1936) and Brachypelma angustum Valerio, 1980, with definition of the taxonomic features for identification of female Sericopelma Ausserer, 1875 (Araneae, Theraphosidae) . ZooKeys 526 : 75 – 104 . Google Scholar Crossref Search ADS WorldCat Galleti A , Guadanucci JP . 2018 . Morphology of setae on the coxae and trochanters of theraphosine spiders (Mygalomorphae: Theraphosidae) . Journal of Arachnology 46 : 214 – 225 . Google Scholar Crossref Search ADS WorldCat Garcia G . 2016 . Wildlife population status in the World, illegal trading, and conservation actions . Cancún : Tarántulas de MEXICO . Available at: http://www.tarantulasdemexico.com/en/statuspoblacion_en.htm (accessed 15 January 2018) . Google Preview WorldCat COPAC Goloboff PA . 1993 . A reanalysis of mygalomorph spider families (Araneae) . American Museum Novitates 3056 : 1 – 32 . WorldCat Goloboff PA , Farris JS , Nixon KC . 2008 . TNT, a free program for phylogenetic analysis . Cladistics 24 : 774 – 786 . Google Scholar Crossref Search ADS WorldCat Graham MR , Hendrixson BE , Hamilton CA , Bond JE . 2015 . Miocene extensional tectonics explain ancient patterns of diversification among turret-building tarantulas (Aphonopelma mojave group) in the Mojave and Sonoran deserts . Journal of Biogeography 42 : 1052 – 1065 . Google Scholar Crossref Search ADS WorldCat Greenstone M , Rowley D , Heimbach U , Lundgren J , Pfannenstiel R , Rehner S . 2005 . Barcoding generalist predators by polymerase chain reaction: carabids and spiders . Molecular Ecology 14 : 3247 – 3266 . Google Scholar Crossref Search ADS PubMed WorldCat Guadanucci JPL . 2014 . Theraphosidae phylogeny: relationships of the ‘Ischnocolinae’ genera (Araneae, Mygalomorphae) . Zoologica Scripta 43 : 508 – 518 . Google Scholar Crossref Search ADS WorldCat Hamilton CA , Formanowicz DR , Bond JE . 2011 . Species delimitation and phylogeography of Aphonopelma hentzi (Araneae, Mygalomorphae, Theraphosidae): cryptic diversity in North American tarantulas . PLoS ONE 6 : e26207 . Google Scholar Crossref Search ADS PubMed WorldCat Hamilton CA , Hendrixson BE , Brewer MS , Bond J . 2014 . An evaluation of sampling effects on multiple DNA barcoding methods leads to an integrative approach for delimiting species: a case study of the North American tarantula genus Aphonopelma (Araneae, Mygalomorphae, Theraphosidae) . Molecular Phylogenetics and Evolution 71 : 79 – 93 . Google Scholar Crossref Search ADS PubMed WorldCat Hamilton CA , Hendrixson BE , Bond JE . 2016 . Taxonomic revision of the tarantula genus Aphonopelma Pocock 1901 (Araneae, Mygalomorphae, Theraphosidae) within the United States . Zookeys 560 : 1 – 340 . Google Scholar Crossref Search ADS WorldCat Hancock K , Hancock J . 1989 . Sex determination of immature theraphosid spiders from their cast skins . Southminster : Published by the authors , 61 . Google Preview WorldCat COPAC Hebert PD , Cywinska A , Ball SL , deWaard JR . 2003 . Biological identifications through DNA barcodes . Proceedings of the Royal Society of London, Series B: Biological Sciences 270 : 313 – 321 . Google Scholar Crossref Search ADS WorldCat Hedin M , Bond J . 2006 . Molecular phylogenetics of the spider infraorder Mygalomorphae using nuclear rRNA genes (18S and 28S): conflict and agreement with the current system of classification . Molecular Phylogenetics and Evolution 41 : 454 – 471 . Google Scholar Crossref Search ADS PubMed WorldCat Hénaut Y , Rabet S , Weissenberger H , Rojo R . 2015 . Dimorphism and population size of the Mexican redrump tarantula, Brachypelma vagans (Araneae: Theraphosidae) . Revista Mexicana de Biodiversidad 86 : 737 – 743 . Google Scholar Crossref Search ADS WorldCat Hendrixson BE , Bond JE . 2009 . Evaluating the efficacy of continuous quantitative characters for reconstructing the phylogeny of a morphologically homogeneous spider taxon (Araneae, Mygalomorphae, Antrodiaetidae, Antrodiaetus) . Molecular Phylogenetics and Evolution 53 : 300 – 313 . Google Scholar Crossref Search ADS PubMed WorldCat Hendrixson BE , DeRussy BM , Hamilton CA , Bond JE . 2013 . An exploration of species boundaries in turret-building tarantulas of the Mojave Desert (Araneae, Mygalomorphae, Theraphosidae, Aphonopelma) . Molecular Phylogenetics and Evolution 66 : 327 – 340 . Google Scholar Crossref Search ADS PubMed WorldCat Hendrixson BE , Guice AV , Bond JE . 2015 . Integrative species delimitation and conservation of tarantulas (Araneae, Mygalomorphae, Theraphosidae) from a North American biodiversity hotspot . Insect Conservation and Diversity 8 : 120 – 131 . Google Scholar Crossref Search ADS WorldCat Hijmensen E . 2012 . The genus Brachypelma . Netherlands . Available at: http://mantid.nl/tarantula/Brachypelma.html (accessed 10 February 2018) . Google Preview WorldCat COPAC Katoh K , Misawa K , Kuma K , Miyata T . 2002 . MAFFT: a novel method for rapid multiple sequence alignment based on fast Fourier transform . Nucleic Acids Research 30 : 3059 – 3066 . Google Scholar Crossref Search ADS PubMed WorldCat Katoh K , Kuma K , Toh H , Miyata T . 2005 . MAFFT v.5: improvement in accuracy of multiple sequence alignment . Nucleic Acids Research 33 : 511 – 518 . Google Scholar Crossref Search ADS PubMed WorldCat Kearse M , Moir R , Wilson A , Stones-Havas S , Cheung M , Sturrock S , Buxton S , Cooper A , Markowitz S , Duran C , Thierer T , Ashton B , Mentjies P , Drummond A . 2012 . Geneious Basic: an integrated and extendable desktop software platform for the organization and analysis of sequence data . Bioinformatics 28 : 1647 – 1649 . Google Scholar Crossref Search ADS PubMed WorldCat Koch CL . 1841 . Die Arachniden . Nürnberg : Achter Band , 41 – 131 , 1–56. Google Preview WorldCat COPAC Kuntner M , Agnarsson I . 2011 . Biogeography and diversification of hermit spiders on Indian Ocean Islands (Nephilidae: Nephilengys) . Molecular Phylogenetics and Evolution 59 : 477 – 488 . Google Scholar Crossref Search ADS PubMed WorldCat Locht A , Yáñez M , Vázquez I . 1999 . Distribution and natural history of Mexican species of Brachypelma and Brachypelmides (Theraphosidae, Theraphosinae) with morphological evidence for their synonymy . Journal of Arachnology 27 : 196 – 200 . WorldCat Locht A , Medina F , Rojo R , Vázquez I . 2005 . Una nueva especie de tarántula del género Aphonopelma Pocock 1901 (Araneae, Theraphosidae, Theraphosinae) de MEXICO con notas sobre el género Brachypelma Simon 1891 . Boletín de la Sociedad Entomológica Aragonesa 37 : 105 – 108 . WorldCat Locke SA , McLaughlin DJ , Marcogliese DJ . 2010 . DNA barcodes show cryptic diversity and a potential physiological basis for host specificity among Diplostomoidea (Platyhelminthes: Digenea) parasitizing freshwater fishes in the St. Lawrence River, Canada . Molecular Ecology 19 : 2813 – 2827 . Google Scholar Crossref Search ADS PubMed WorldCat Longhorn SJ . 2002 . Non-lethal DNA sampling from CITES II protected ‘tarantula’ spiders of Belize . Las Cuevas Newsletter 9 : 8 – 9 . WorldCat Longhorn SJ . 2014 . On type localities of Mexican tarantulas, either missing for Bonnetina or misleading for Brachypelma, with appeal for accurate biogeographic data . Journal of the British Tarantula Society 29 : 16 – 28 . WorldCat Longhorn SJ , Nicholas M , Chuter J , Vogler A . 2007 . The utility of molecular markers from non-lethal DNA samples of the CITES II protected ‘tarantula’ Brachypelma vagans (Araneae, Theraphosidae) . Journal of Arachnology 35 : 278 – 292 . Google Scholar Crossref Search ADS WorldCat Lüddecke T , Krehenwinkel H , Canning G , Glaw F , Longhorn SJ , Tänzler R , Wendt I , Vences M . 2018 . Discovering the silk road: nuclear and mitochondrial sequence data resolve the phylogenetic relationships among theraphosid spider subfamilies . Molecular Phylogenetics and Evolution 119 : 63 – 70 . Google Scholar Crossref Search ADS PubMed WorldCat Machkour-M'Rabet S , Hénaut Y , Sepúlveda A , Rojo R , Calmé S , Geissen V. 2007 . Soil preference and borrow structure of an endangered tarantula, Brachypelma vagans (Mygalomorphae: Theraphosidae) . Journal of Natural History 41 : 1025 – 1033 . Google Scholar Crossref Search ADS WorldCat Machkour-M’Rabet S , Hénaut Y , Rojo R , Calmé S . 2005 . A not so natural history of the tarantula Brachypelma vagans: interaction of the human activity . Journal of Natural History 39 : 2515 – 2523 . Google Scholar Crossref Search ADS WorldCat Machkour-M’Rabet SM , Hénaut Y , Dor A , Perez-Lachaud G , Pelissier C , Gers C , Legal L . 2009 . ISSR (Inter Simple Sequence Repeats) as molecular markers to study genetic diversity in tarantulas (Araneae, Mygalomorphae) . Journal of Arachnology 37 : 10 – 14 . Google Scholar Crossref Search ADS WorldCat Machkour-M’Rabet SM , Hénaut Y , Winterton P , Rojo R . 2011 . A case of zootherapy with the tarantula Brachypelma vagans Ausserer, 1875 in traditional medicine of the Chol Mayan ethnic group in Mexico . Journal of Ethnobiology & Ethnomedicine 7 : 1 – 7 . Google Scholar Crossref Search ADS WorldCat Machkour-M’Rabet S , Hénaut Y , Calmé S , Legal L . 2012 . When landscape modification is advantageous for protected species. The case of synanthropic tarantula, Brachypelma vagans . Journal of Insect Conservation 16 : 479 – 488 . Google Scholar Crossref Search ADS WorldCat Machkour-M’Rabet S , Dor A , Hénaut Y . 2015 . Megaselia scalaris (Diptera: Phoridae): an opportunistic endoparasitoid of the endangered Mexican redrump tarantula Brachypelma vagans (Araneae: Theraphosidae) . Journal of Arachnology 43 : 115 – 119 . Google Scholar Crossref Search ADS WorldCat Machkour-M’Rabet S , Vilchis-Nestor C , Barriga-Sosa IA , Legal L , Hénaut Y . 2017 . A molecular approach to understand the riddle of the invasive success of the tarantula, Brachypelma vagans, on Cozumel Island, Mexico . Biochemical Systematics and Ecology 70 : 260 – 267 . Google Scholar Crossref Search ADS WorldCat Marshall SD , Thoms EM , Uetz GW . 1995 . Setal entanglement: an undescribed method of stridulation by a Neotropical tarantula (Araneae: Theraphosidae) . Journal of Zoology 235 : 587 – 595 . Google Scholar Crossref Search ADS WorldCat Mendoza MJI . 2014a . Taxonomic revision of Hemirrhagus Simon, 1903 (Araneae: Theraphosidae, Theraphosinae), with description of five new species from Mexico . Zoological Journal of the Linnean Society 170 : 634 – 689 . Google Scholar Crossref Search ADS WorldCat Mendoza MJI . 2014b . Psalmopoeus victori, the first arboreal theraphosid spider described for Mexico (Araneae: Theraphosidae: Aviculariinae) . Revista Mexicana de Biodiversidad 85 : 728 – 735 . Google Scholar Crossref Search ADS WorldCat Mendoza JI , Francke OF . 2017 . Systematic revision of Brachypelma red-kneed tarantulas (Araneae: Theraphosidae), and the use of DNA barcodes to assist in the identification and conservation of CITES-listed species . Invertebrate Systematics 31 : 157 – 179 . Google Scholar Crossref Search ADS WorldCat Miller MA , Pfeiffer W , Schwartz T . 2010 . Creating the CIPRES Science Gateway for inference of large phylogenetic trees. In: Proceedings of the Gateway Computing Environments Workshop (GCE) . New Orleans, LA : Institute of Electrical and Electronics Engineers, 1 – 8 . Montes de Oca L , D’Elía G , Pérez-Miles F . 2015 . An integrative approach for species delimitation in the spider genus Grammostola (Theraphosidae, Mygalomorphae) . Zoologica Scripta 45 : 322 – 333 . Google Scholar Crossref Search ADS WorldCat Mundie DA . 1995 . The NBS/ISCC color system / David A . Mundie Pittsburgh : Polymath Systems 535.6 dc-20 . Available at: http://www.anthus.com/Colors/NBS.html (accessed 12 January 2018) . Google Preview WorldCat COPAC Niemiller ML , Near TJ , Fitzpatrick BM . 2011 . Delimiting species using multilocus data: diagnosing cryptic diversity in the southern cavefish, Typhlichthys subterraneus (Teleostei: Amblyopsidae) . Evolution 66 : 846 – 866 . Google Scholar Crossref Search ADS PubMed WorldCat Ortiz D , Francke F . 2016 . Two DNA barcodes and morphology for multi-method species delimitation in Bonnetina tarantulas (Araneae: Theraphosidae) . Molecular Phylogenetics and Evolution 101 : 176 – 193 . Google Scholar Crossref Search ADS PubMed WorldCat Pante E , Schoelinck C , Puillandre N . 2014 . From integrative taxonomy to species description: one step beyond . Systematic Biology 64 : 152 – 160 . Google Scholar Crossref Search ADS PubMed WorldCat Perafán C , Pérez-Miles F . 2014 . The Andean tarantulas Euathlus Ausserer, 1875, Paraphysa Simon 1892 and Phrixotrichus Simon, 1889 (Araneae: Theraphosidae): phylogenetic analysis, genera redefinition and new species descriptions . Journal of Natural History 48 : 2389 – 2418 . Google Scholar Crossref Search ADS WorldCat Pérez-Miles F . 1989 . Variación relativa de caracteres somáticos y genitales en Grammostola mollicoma (Araneae, Theraphosidae) . Journal of Arachnology 17 : 263 – 274 . WorldCat Pérez-Miles F . 1994 . Tarsal scopula division in Theraphosinae (Araneae, Theraphosidae): its systematic significance . Journal of Arachnology 22 : 46 – 53 . WorldCat Pérez-Miles F . 2000 . Iracema cabocla new genus and species of a theraphosid spider from Amazonic Brazil (Araneae, Theraphosinae) . Journal of Arachnology 28 : 141 – 148 . Google Scholar Crossref Search ADS WorldCat Pérez-Miles F, Locht A. 2003. Revision and cladistic analysis of the genus Hemirrhagus Simon, 1903 (Araneae, Theraphosidae, Theraphosinae). Bulletin of the British Arachnological Society 12: 365–375. Pérez-Miles F , Lucas SM , da Silva PI Jr , Bertani R . 1996 . Systematic revision and cladistics analysis of Theraphosinae (Araneae: Theraphosidae) . Mygalomorph 1 : 33 – 68 . WorldCat Peters HJ . 2000 . Tarantulas of the world: kleiner Atlas der Vogelspinnen – Band 1 . Wegberg : Published by the author , 148 . Google Preview WorldCat COPAC Peters HJ . 2003 . Tarantulas of the world: Amerika’s Vogelspinnen . Wegberg : Published by the author , 328 . Google Preview WorldCat COPAC Petersen FT , Damgaard J , Meier R . 2007 . DNA taxonomy: how many DNA sequences are needed for solving a taxonomic problem? The case of two parapatric species of louse flies (Diptera: Hippoboscidae: Ornithomya Latreille, 1802) . Arthropod Systematics and Phylogeny 65 : 119 – 125 . WorldCat Pickard-Cambridge FO . 1897 . Arachnida – Araneida and Opiliones. Biologia Centrali-Americana . Zoology 2 : 1 – 40 . WorldCat Pickard-Cambridge O. 1892 . Arachnida. Araneida. Biologia Centrali-Americana . Zoology 1 : 89 – 104 . WorldCat Pocock RI . 1903 . On some genera and species of South-American Aviculariidae . Annals and Magazine of Natural History 11 : 81 – 115 . Google Scholar Crossref Search ADS WorldCat Prendini L . 2005 . Comment on ‘Identifying species through DNA barcodes’ . Canadian Journal of Zoology 83 : 481 – 491 . Google Scholar Crossref Search ADS WorldCat Raven RJ . 1985 . The spider infraorder Mygalomorphae (Araneae): cladistics and systematics . Bulletin of the American Museum of Natural History 182 : 1 – 180 . WorldCat Reichling SB . 2000 . Group dispersal in juvenile Brachypelma vagans (Araneae, Theraphosidae) . Journal of Arachnology 28 : 248 – 250 . Google Scholar Crossref Search ADS WorldCat Reichling SB . 2001 . Theraphosid spiders surveyed at Las Cuevas . The Newsletter of the Las Cuevas Research Station Belize 8 : 6 . WorldCat Reichling SB . 2003 . Tarantulas of Belize . Florida : Krieger Publishing Company , 127 . Google Preview WorldCat COPAC Roewer CF . 1942 . Katalog der Araneae von 1758 bis 1940. 1. Band (Mesothelae, Orthognatha, Labidognatha: Dysderaeformia, Scytodiformia, Pholciformia, Zodariiformia, Hersiliaeformia, Argyopiformia). In: Natura, Buchhandlung für Naturkunde und exakte Wissenschaften . Bremen : Paul Budy , 1040 . Google Preview WorldCat COPAC Rojo R . 2004 . Las tarántulas de MEXICO: pequeños gigantes incomprendidos . Biodiversitas 56 : 7 – 11 . WorldCat Rudloff JP . 2003 . Eine neue Brachypelma-Art aus Mexiko, Brachypelma schroederi sp. n. (Araneae: Mygalomorphae: Theraphosidae: Theraphosinae) . Arthropoda 11 : 2 – 15 . WorldCat Rudloff JP . 2008 . Eine neue Brachypelma-Art aus Mexiko (Araneae: Mygalomorphae: Theraphosidae: Theraphosinae) . Arthropoda 16 : 26 – 30 . WorldCat Rudloff JP , Weinmann D . 2010 . A new giant tarantula from Guyana . Arthropoda Scientia 1 : 21 – 40 . WorldCat Schmidt G . 1992a . Brachypelma Simon 1890 oder Euathlus Ausserer 1875? (Araneida: Theraphosidae: Theraphosinae) . Arachnologischer Anzeiger 3 : 9 – 11 . WorldCat Schmidt G . 1992b . Brachypelma auratum sp. n., die sogenannte Hochlandform von Brachypelma smithi (Araneida: Theraphosidae: Theraphosinae) . Arachnologischer Anzeiger 3 : 9 – 14 . WorldCat Schmidt G . 1993 . Vogelspinnen: Vorkommen, Lebensweise, Haltung und Zucht, mit Bestimmungsschlüsseln für alle Gattungen, vierte Auflage . Hannover : Landbuch Verlag , 151 . Google Preview WorldCat COPAC Schmidt G . 1997a . Eine zweite Brachypelmides-Art aus Mexiko: Brachypelmides ruhnaui n. sp. (Arachnida: Araneae: Theraphosidae: Theraphosinae) . Entomologische Zeitschrift 107 : 205 – 208 . WorldCat Schmidt G . 1997b . Bestimmungsschlüssel für die Gattungen der Unterfamilie Theraphosinae (Araneae: Theraphosidae) . Arachnologisches Magazin 3 : 1 – 27 . WorldCat Schmidt G . 2003 . Die Vogelspinnen: eine weltweite Übersicht . Hohenwarsleben : Neue Brehm-Bücherei , 383 . Google Preview WorldCat COPAC Schmidt G . 2004 . Die Gattung Brachypelmides Schmidt & Krause, 1994 (Araneae: Theraphosidae: Theraphosinae) . Tarantulas of the World 99 : 4 – 5 . WorldCat Schmidt G , Klaas P . 1993 . Eine neue Brachypelma-Spezies aus Mexiko (Araneida: Theraphosidae: Theraphosinae) . Arachnologischer Anzeiger 4 : 7 – 9 , 11 – 13 . WorldCat Schmidt G , Krause RH . 1994 . Eine neue Vogelspinnen-Spezies aus Mexico, Brachypelmides klaasi sp. n. (Araneida, Theraphosidae, Theraphosinae) . Studies on Neotropical Fauna and Environment 29 : 7 – 10 . Google Scholar Crossref Search ADS WorldCat Scotland RW , Hughes C , Bailey D , Wortley A . 2003 . The Big Machine and the much-maligned taxonomist . Systematics and Biodiversity 1 : 139 – 143 . Google Scholar Crossref Search ADS WorldCat SEMARNAT (Secretaria de Medio Ambiente, Recursos Naturales y Pesca). 2000 . Ley general de vida silvestre gaceta ecológica . Available at: http://www.redalyc.org/articulo.oa?id=53905505 (accessed 18 February 2018) . Sereno P . 2007 . Logical basis for morphological characters in phylogenetics . Cladistics 23 : 565 – 587 . WorldCat Shaw EM , Bennett SP , Wheater CP . 2011 . Distribution of Brachypelma vagans (Theraphosidae) burrows and their characteristics in Belize over two years . Journal of Arachnology 39 : 515 – 518 Google Scholar Crossref Search ADS WorldCat Shillington CJ , McEwen M . 2006 . Activity of juvenile tarantulas in and around the maternal borrow . Journal of Arachnology 34 : 261 – 265 . Google Scholar Crossref Search ADS WorldCat Simon E . 1891 . Liste des Aviculariides qui habitant le Mexique et l’Amérique centrale . Actes de la Société Linnéenne de Bordeaux 44 : 327 – 339 . WorldCat Simon E . 1903 . Histoire naturelle des araignées . Paris : 669 – 1080 . Available at: https://doi.org/10.5962/bhl.title.51973 WorldCat Slowik J , Blagoev GA . 2012 . A survey of spiders (Arachnida: Araneae) of Prince Wales Island, Alaska; combining morphological and DNA barcode identification techniques . Insecta Mundi 251 : 1 – 12 . WorldCat Smith AM . 1992 . In defense of Raven’s decision to make the genus Brachypelma Simon 1891 a junior synonymy [sic] of Euathlus Ausserer 1895 . British Tarantula Society Journal 7 : 14 – 19 . WorldCat Smith AM . 1993 . A new mygalomorph spider from Mexico (Brachypelma, Theraphosidae, Arachnida) Brachypelma baumgarteni N sp . British Tarantula Society Journal 8 : 14 – 19 . WorldCat Smith AM . 1994 . Tarantula spiders: tarantulas of the U.S.A. and Mexico . London : Fitzgerald , 411 . Google Preview WorldCat COPAC Stamatakis A . 2014 . RAxML v.8: a tool for phylogenetic analysis and post-analysis of large phylogenies . Bioinformatics 30 : 1312 – 1313 . Google Scholar Crossref Search ADS PubMed WorldCat Stamatakis A , Hoover P , Rougemont J . 2008 . A rapid bootstrap algorithm for the RAxML Web servers . Systematic Biology 57 : 758 – 771 . Google Scholar Crossref Search ADS PubMed WorldCat Sun Y , Li Q , Kong L , Zheng X . 2012 . DNA barcoding of Caenogastropoda along coast of China based on the COI gene . Molecular Ecology Resources 12 : 209 – 218 . Google Scholar Crossref Search ADS PubMed WorldCat Tesmoingt M , Cleton F , Verdez JM . 1997a . Description de Brachypelma annitha n. sp. et de Brachypelma hamorii n. sp. mâles et femelles, nouvelles espèces proches de Brachypelma smithi (Cambridge, 1897) du Mexique . Arachnides 32 : 8 – 20 . WorldCat Tesmoingt M , Cleton F , Verdez JM . 1997b . Description de Brachypelma annitha n. sp. et de Brachypelma hamorii n. sp. mâles et femelles, nouvelles espèces proches de Brachypelma smithi (Cambridge, 1897) du Mexique. 2ème partie . Arachnides 33 : 2 – 10 . WorldCat Teyssié F . 2015 . Tarantulas of the world . Paris : NAP Éditions , 487 . Google Preview WorldCat COPAC Turner SP , Longhorn SJ , Hamilton CA , Gabriel R , Pérez-Miles F , Vogler AP . 2018 . Re-evaluating conservation priorities of New World tarantulas (Araneae: Theraphosidae) in a molecular framework indicates non-monophyly of the genera, Aphonopelma and Brachypelma . Systematics and Biodiversity 16 : 89 – 107 . Google Scholar Crossref Search ADS WorldCat Uetz GW , Stratton GE . 1982 . Acoustic communication and reproductive isolation in spiders. In: Witt P , Rovner J , eds. Spider communication. Mechanisms and ecological significance . Princeton : Princeton University Press , 123 – 159 . Google Preview WorldCat COPAC Valerio CE . 1980 . Arañas terafosidas de Costa Rica (Araneae, Theraphosidae). I. Sericopelma y Brachypelma . Brenesia 18 : 259 – 288 . WorldCat Vilchis-Nestor CA , Machkour-M’Rabet S , Barriga-Sosa IA , Winterton P , Hénaut Y . 2013 . Morphological and colour differences between island and mainland populations in the Mexican redrump tarantula, Brachypelma vagans . Journal of Insect Science 13 : 95 . Google Scholar Crossref Search ADS PubMed WorldCat Vol F . 2002 . Stridulation in tarantulas . Journal of the British Tarantula Society 18 : 21 – 27 . WorldCat West RC . 1996 . Some natural history field notes on three Brachypelma species from Mexico . British Tarantula Society Journal 10 : 111 – 116 . WorldCat West RC . 2005 . The Brachypelma of Mexico . Journal of the British Tarantula Society 20 : 108 – 119 . WorldCat West R , Nunn C . 2010 . A taxonomic revision of the tarantula spider genus Coremiocnemis Simon 1892 (Araneae, Theraphosidae), with further notes on the Selenocosmiinae . Zootaxa 2443 : 1 – 64 . Google Scholar Crossref Search ADS WorldCat West RC , Marshall SD , Fukushima CS , Bertani R . 2008 . Review and cladistic analysis of the neotropical tarantula genus Ephebopus Simon 1892 (Araneae: Theraphosidae) with notes on the Aviculariinae . Zootaxa 1849 : 35 – 58 . Google Scholar Crossref Search ADS WorldCat White A . 1856 . Description of Mygale emilia, a spider from Panama, hitherto apparently unrecorded. Proceedings of the Zoological Society of London 24 : 183 – 185 . Will K , Rubinoff D . 2004 . Myth of the molecule: DNA barcodes for species cannot replace morphology for identification and classification . Cladistics 20 : 47 – 55 . Google Scholar Crossref Search ADS WorldCat Wilson JS , Gunnell CF , Wahl DB , Pitts JP . 2013 . Testing the species limits of the tarantulas (Araneae: Theraphosidae) endemic to California’s Southern Coast Ranges, USA . Insect Conservation and Diversity 6 : 365 – 371 . Google Scholar Crossref Search ADS WorldCat World Conservation Monitoring Centre. 1996 . Brachypelma smithi . The IUCN Red List of Threatened Species. Available at: http://dx.doi.org/10.2305/IUCN.UK.1996.RLTS.T8152A12893193.en (accessed 2 March 2018) . Google Preview WorldCat COPAC World Spider Catalog. 2018. Version 17. Natural History Museum Bern. Available at: http://wsc.nmbe.ch (accessed 25 January 2018) . Yáñez M , Floater G . 2000 . Spatial distribution and habitat preference of the endangered tarantula, Brachypelma klaasi (Araneae: Theraphosidae) in Mexico . Biodiversity & Conservation 9 : 795 – 810 . Google Scholar Crossref Search ADS WorldCat Yáñez M , Locht A , Macías-Ordóñez R . 1999 . Courtship and mating behavior of Brachypelma klassi (Araneae, Theraphosidae) . Journal of Arachnology 27 : 165 – 170 . WorldCat © 2019 The Linnean Society of London, Zoological Journal of the Linnean Society This article is published and distributed under the terms of the Oxford University Press, Standard Journals Publication Model (https://academic.oup.com/journals/pages/open_access/funder_policies/chorus/standard_publication_model)
TI - Systematic revision of Mexican threatened tarantulas Brachypelma (Araneae: Theraphosidae: Theraphosinae), with a description of a new genus, and implications on the conservation
JF - Zoological Journal of the Linnean Society
DO - 10.1093/zoolinnean/zlz046
DA - 2020-01-01
UR - https://www.deepdyve.com/lp/oxford-university-press/systematic-revision-of-mexican-threatened-tarantulas-brachypelma-UNekGUECcG
SP - 82
VL - 188
IS - 1
DP - DeepDyve
ER -