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Taxonomic revision of the tarantula genus Aphonopelma Pocock, 1901 (Araneae, Mygalomorphae, Theraphosidae) within the United States

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Abstract

This systematic study documents the taxonomy, diversity, and distribution of the tarantula spider genus Aphonopelma Pocock, 1901 within the United States. By employing phylogenomic, morphological, and geospatial data, we evaluated all 55 nominal species in the United States to examine the evolutionary history of Aphonopelma and the group's taxonomy by implementing an integrative approach to species delimitation. Based on our analyses, we now recognize only 29 distinct species in the United States. We propose 33 new synonymies (A. apacheum, A. minchi, A. rothi, A. schmidti, A. stahnkei = A. chalcodes; A. arnoldi = A. armada; A. behlei, A. vogelae = A. marxi; A. breenei = A. anax; A. chambersi, A. clarum, A. cryptethum, A. sandersoni, A. sullivani = A. eutylenum; A. clarki, A. coloradanum, A. echinum, A. gurleyi, A. harlingenum, A. odelli, A. waconum, A. wichitanum = A. hentzi; A. heterops = A. moderatum; A. jungi, A. punzoi = A. vorhiesi; A. brunnius, A. chamberlini, A. iviei, A. lithodomum, A. smithi, A. zionis - = A. iodius; A. phanum, A. reversum = A. steindachneri), 14 new species (A. atomicum sp. n., A. catalina sp. n., A. chiricahua sp. n., A. icenoglei sp. n., A. johnnycashi sp. n., A. madera sp. n., A. mareki sp. n., A. moellendorfi sp. n., A. parvum sp. n., A. peloncillo sp. n., A. prenticei sp. n., A. saguaro sp. n., A. superstitionense sp. n., and A. xwalxwal sp. n.), and seven nomina dubia (A. baergi, A. cratium, A. hollyi, A. mordax, A. radinum, A. rusticum, A. texense). Our proposed species tree based on Anchored Enrichment data delimits five major lineages: a monotypic group confined to California, a western group, an eastern group, a group primarily distributed in high- elevation areas, and a group that comprises several miniaturized species. Multiple species are distributed throughout two biodiversity hotspots in the United States (i.e., California Floristic Province and Madrean Pine-Oak Woodlands). Keys are provided for identification of both males and females. By conducting the most comprehensive sampling of a single theraphosid genus to date, this research significantly broadens the scope of prior molecular and morphological investigations, finally bringing a modern understanding of species delimitation in this dynamic and charismatic group of spiders.
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... Whilst the latter was valid at the time of the publication of (later synonymised with Holothele longipes (L. Koch, 1875) by Guadanucci et al., 2017) A. cochise was not a valid species because this taxon was described (together with other invalid putative species) in Albert Jung's unpublished thesis and was thus, sadly, unavailable (Prentice, 1993;Hamilton et al., 2016;Sherwood, 2019). This can be rectified by reference here to the correct identification of this specimen: Aphonopelma vorhiesi (Chamberlin & Ivie, 1939). ...
... The vast majority of modern taxonomic works on theraphosid spiders utilise genital organ morphology for species and generic level delineation (e.g. Bertani, 2001;Fukushima et al., 2005;Yamamoto et al., 2007;Rodríguez-Manzanilla & Bertani, 2010;Lucas et al., 2010;Kaderka, 2015Kaderka, , 2016Hamilton et al., 2016;Gabriel, 2016;Fabiano-da-Silva et al., 2019;, 2020a, 2020b with many New World taxa, in particular, being strongly defined on male palpal bulb morphology (see previous references, and references therein), which has been typified and standardised since the start of the millennium (Bertani, 2000). The anatomical features used by a species to reproduce, which are often highly specialised, can hardly be described as "superficial". ...
... Morphological homogeneity is recognised within theraphosid spiders, particularly in the subfamily Theraphosinae, and morphology alone may not be sufficient to fully understand species boundaries (Hamilton et al., 2016;Ortiz et al., 2018). Nonetheless, it is clear that many morphological features have been used in countless studies to define many congruent boundaries and have been found to be informative. ...
... In modern taxonomy, it is generally accepted that the use of multiple lines of evidence, such as morphology, ecology, reproduction, and biogeography (integrative taxonomy), are more effective in the diagnosis, delimitation, and even the description of species (Valdez-Mondragón 2020). Several recently developed molecular delimitation methods have highlighted the extensive inconsistency in classical morphological taxonomy used on different groups of arachnids (Hamilton et al. 2011(Hamilton et al. , 2014(Hamilton et al. , 2016Ortiz & Francke 2016;Tahami et al. 2017;Cruz-López et al. 2019;Valdez-Mondragón et al. 2019;Navarro-Rodríguez & Valdez-Mondragón 2020). However, the molecular delimitation methods using mitochondrial markers VALDEZ-MONDRAGÓN A. & CORTEZ-ROLDÁN M.R., New Mexican species of Pseudocellus 3 have never been used in ricinuleids so far. ...
... Modern taxonomy is changing, and currently uses multiple lines of evidence for species identification and delimitation ( tasks to which DNA barcodes markers are being deeply applied in modern systematics and taxonomy, first distinguishing between 'sibling' species (equivalent to species identification or species diagnosis), and second, the use of molecular data to discover underestimated biodiversity (equivalent to species delimitation and species description). How species boundaries are fixed is a central debate in modern systematics, which has generated multiple molecular delimitation methods based on heuristic and parametric methods tending to solve the inconsistencies in classical taxonomy (Hamilton et al. 2011(Hamilton et al. , 2014(Hamilton et al. , 2016Ortiz & Francke 2016;Rannala & Yang 2020). The use of the mitochondrial cytochrome c oxidase subunit 1 (CO1) barcode gene has become useful as the standard animal barcoding marker in many taxonomic studies, including arachnids (Hebert et al. 2003(Hebert et al. , 2004Astrin et al. 2006;Correa-Ramírez et al. 2010;Hamilton et al. 2011Hamilton et al. , 2014Hamilton et al. , 2016Graham et al. 2015;Planas & Ribera 2015;Ortiz & Francke 2016;Tahami et al. 2017;Cruz-López et al. 2019;Valdez-Mondragón et al. 2019;Valdez-Mondragón 2020). ...
... How species boundaries are fixed is a central debate in modern systematics, which has generated multiple molecular delimitation methods based on heuristic and parametric methods tending to solve the inconsistencies in classical taxonomy (Hamilton et al. 2011(Hamilton et al. , 2014(Hamilton et al. , 2016Ortiz & Francke 2016;Rannala & Yang 2020). The use of the mitochondrial cytochrome c oxidase subunit 1 (CO1) barcode gene has become useful as the standard animal barcoding marker in many taxonomic studies, including arachnids (Hebert et al. 2003(Hebert et al. , 2004Astrin et al. 2006;Correa-Ramírez et al. 2010;Hamilton et al. 2011Hamilton et al. , 2014Hamilton et al. , 2016Graham et al. 2015;Planas & Ribera 2015;Ortiz & Francke 2016;Tahami et al. 2017;Cruz-López et al. 2019;Valdez-Mondragón et al. 2019;Valdez-Mondragón 2020). However, the use of mitochondrial markers alone may lead to the overestimation of taxonomic diversity, as has been demonstrated in some tarantula genera (Ortiz & Francke 2019). ...
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A new species of epigean ricinuleid of the genus Pseudocellus Platnick, 1980 from El Triunfo Biosphere Reserve, Chiapas, Mexico is described. DNA barcoding utilizing mitochondrial cytochrome c oxidase subunit 1 (CO1) and morphology were used for species delimitation. Molecular analyses and species delimitation included four methods: 1) General Mixed Yule Coalescent model (GMYC), 2) Automatic Barcode Gap Discovery (ABGD), 3) Bayesian Poisson Tree Process (bPTP), and 4) Assemble Species by Automatic Partitioning (ASAP). All molecular methods and morphology were consistent in delimiting and recognizing the new species described herein. The average interspecific genetic distance (p-distance) among analyzed species of Pseudocellus was 11.6%. The species is described based on adult males and females: Pseudocellus giribeti sp. nov. This is the seventh species described from Chiapas, which holds the highest number of ricinuleids species for the country. The total number of described species of Pseudocellus from Mexico increases to 21, having the highest species diversity of known ricinuleids worldwide.
... In this work, we provide a catalogue of all known specimens of the family Theraphosidae Thorell, 1869 Remarks: This single male of A. chiricahua represents the first specimen record, to our knowledge, of this species in a museum outside the United States. This specimen was collected by the notable arachnologist Vincent Roth at the Southwest Research Station (SWRS), a field station of the American Museum of Natural History (where Roth worked for many years) and known locality of A. chiricahua (see Hamilton et al. 2016 Remarks: The MMUE's Register Book contains no records for 1897, and the Museum's Report (1897)(1898) has no mention of H. Brazenor or this specimen. Therefore, it is enigmatic, and we can only present the data found on the original data label; the specimen was re-accessioned on 11 November 2021 (G7669). ...
... The female does not appear to be conspecific to the male, the latter of which belongs to the group of Aphonopelma species which occur only in the United States and Northern Mexico (including Baja California). Without a robust locality it is difficult to determine the male at the species level based only on its palpal bulb morphology, as this is very homogenous within this group (Hamilton et al. 2016). Based on the Museum's Report (1911-1912: 31), it is possible to assume that this female could be the "specimen of Mygale from California" donated by J. R. Hardy, yet with no reference on whether it was used for making such an educational exhibit. ...
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An annotated catalogue of all specimens of the family Theraphosidae Thorell, 1869 held in the collections of the Manchester Museum is presented.
... We are well aware of the fact that accurate identification of mygalomorph spiders based only on females is complicated due to their highly conserved morphology (Raven 1985;Bailey 2010;Ferretti et al. 2019). The most reliable character among females is the shape of spermathecae, but in some cases, it can also vary considerably (see Hamilton, Hendrixson & Bond 2016). In his revision of the spider genus Bolostromus, Indicatti (2007: 10, fig. 3) showed drawings of the spermathecae from seven species and diagnosed the females of B. panamanus by having a "espermateca com um ramo interno reto e mais elevado que ramo o externo, que é projetado anteriormente" [spermathecae with an internal branch straight and longer than the external branch, that is projected anteriorly] (Fig. 3A). ...
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The poorly known species of wafer-lid trapdoor spider Bolostromus panamanus (Petrunkevitch, 1925) (Mygalomorphae: Cyrtaucheniidae) is formally reported from Costa Rica for the first time, based on a female collected in Puerto Viejo de Talamanca. The specimen is described and the first pictures of a living specimen, diagnostic traits, and spermathecae of the species are provided
... Inevitably, the few studies on ants, ground beetles and spiders have often been opportunistic, largely reflecting the specific interests of the few authors who have ventured to explore the potential of SDMs in terrestrial arthropod research. For example, this is evident when looking at a sample of papers on spiders-most studies focused on large-sized, taxonomically unique and/or charismatic species (Decae et al., 2019;Hamilton et al., 2016;Jiménez-Valverde et al., 2011;Wang et al., 2018), taxa of medical importance (Planas et al., 2014;Taucare-Ríos et al., 2018;Wang et al., 2018) or taxa inhabiting peculiar habitats that are the interest of certain authors, such as caves Pavlek & Mammola, 2021). ...
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... These loci have been shown to possess sufficient variation for resolving both shallow and deepscale evolutionary relationships throughout the Araneae, e.g., [14,15]. These data have also been used to recover inter-and intrageneric relationships, as well as inter-and intraspecies relationships within a range of spider families [6,[16][17][18][19][20][21]. Library preparation, enrichment, sequencing, assembly, alignment and phylogeny construction from AHE data followed the procedures described in Kuntner et al. [6]. ...
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Reconstructing biogeographic history is challenging when dispersal biology of studied species is poorly understood, and they have undergone a complex geological past. Here, we reconstruct the origin and subsequent dispersal of coin spiders (Nephilidae: Herennia Thorell), a clade of 14 species inhabiting tropical Asia and Australasia. Specifically, we test whether the all-Asian range of Herennia multipuncta is natural vs. anthropogenic. We combine Anchored Hybrid Enrichment phylogenomic and classical marker phylogenetic data to infer species and population phylogenies. Our biogeographical analyses follow two alternative dispersal models: ballooning vs. walking. Following these assumptions and considering measured distances between geographical areas through temporal intervals, these models infer ancestral areas based on varying dispersal probabilities through geological time. We recover a wide ancestral range of Herennia including Australia, mainland SE Asia and the Philippines. Both models agree that H. multipuncta internal splits are generally too old to be influenced by humans, thereby implying its natural colonisation of Asia, but suggest quite different colonisation routes of H. multipuncta populations. The results of the ballooning model are more parsimonious as they invoke fewer chance dispersals over large distances. We speculate that coin spiders’ ancestor may have lost the ability to balloon, but that H. multipuncta regained it, thereby colonising and maintaining larger areas.
... The Californian toad, Anaxyrus boreas halophilus (Baird & Girard, 1853) is a wide-ranging amphibian found across California (Stebbins & McGinnis, 2012). Similarly, the theraphosid spider Aphonopelma steindachneri (Ausserer, 1875) displays a wide distribution in Southern and Central California, likely also ranging into Northern Baja California (Hamilton et al., 2016;DS pers. obs.). ...
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