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Taxonomic revision of the tarantula genus Aphonopelma Pocock, 1901 (Araneae, Mygalomorphae, Theraphosidae) within the United States
expand article infoChris A. Hamilton, Brent E. Hendrixson§, Jason E. Bond|
‡ Department of Biological Sciences & Auburn University Museum of Natural History, Auburn, United States of America
§ Department of Biology, Millsaps College, Jackson, United States of America
| Department of Biological Sciences & Auburn University Museum of Natural History, Alabama, United States of America
Open Access

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.

Keywords

Biodiversity, New species, Conservation, Molecular systematics, DNA taxonomy, DNA barcoding, Spider taxonomy

Introduction

The family Theraphosidae (tarantulas, baboon spiders, earth tigers) is the most diverse lineage (World Spider Catalog 2015) of spiders placed in the infraorder Mygalomorphae (Raven 1985, Hedin and Bond 2006, Bond et al. 2012, Bond et al. 2014). Residing within this group is the most species-rich lineage of tarantulas, the genus Aphonopelma, with 87 nominal species distributed throughout North and Central America, 55 of which are thought to occur in the United States (World Spider Catalog 2015). Despite their systematic appeal stemming from their diversity and charismatic nature as hairy, large-bodied spiders, the systematics and taxonomy of Aphonopelma have remained problematic. In the past 75 years, only four significant descriptive or revisionary works have examined the taxonomy and range of morphological character variation of Aphonopelma in the United States (Chamberlin and Ivie 1939, Chamberlin 1940, Smith 1995, Prentice 1997), none of which employed an explicit phylogenetic approach. The goal of this partial revision is to formally resolve the group’s species-level diversity by implementing an integrative approach to species delimitation that considers genomic, morphological, ecological, and geospatial data.

Morphology-based phylogenies of mygalomorph spiders have revealed widespread patterns of homoplasy among traditional taxonomic characters (Raven 1985, Goloboff 1993, Bond and Opell 2002, Hedin and Bond 2006, Bond and Hedin 2006, Hendrixson and Bond 2009, Bond et al. 2012). Furthermore, quantitative or meristic features often used to evaluate relationships among mygalomorphs have been found to be problematic (Bond and Beamer 2006, Hendrixson and Bond 2009, but see Goloboff et al. 2006). Generally, morphological approaches to species delimitation in groups with similar patterns of homoplasy or morphological conservatism have been shown to grossly oversimplify and underestimate diversity (Locke et al. 2010, Niemiller et al. 2011).

Distributed across two major biogeographic realms (Nearctic and Neotropical), Aphonopelma species are distributed across the southern third of the United States, ranging west of the Mississippi River to California and south through Mexico and Central America (Fig. 1). The North American species of Aphonopelma, proposed to have rapidly diversified following expansion and adaptation into arid and desert environments ~5 Ma (Hamilton et al. 2011), can be found across a wide range of physical and climatic conditions. Most Aphonopelma live in silk-lined subterranean burrows and are found in nearly every habitat throughout their distribution (Fig. 2). Some species thrive in harsh environments, including hot and arid regions near Death Valley (California, USA), or in more temperate high-elevation forests along the Mogollon Rim and Madrean Archipelago (Arizona, USA) (Fig. 1). Perceived high diversity, complicated biogeography, morphological homogeneity between closely related taxa, and considerable variation within nominal species have posed significant problems for species delimitation (Prentice 1997) and higher-level classification (Raven 1985) in tarantulas. Not surprisingly, many arachnologists have expressed dismay toward the present state of theraphosid taxonomy (Raven 1985, Smith 1995, Pérez-Miles et al. 1996, Prentice 1997), with Raven (1990, p. 126) even declaring the group a “nomenclatural and taxonomic nightmare”.

Figure 1.

Breadth of diversity of Aphonopelma species habitat types across the United States. A grassland prairie, Otero Co., Colorado B high-elevation pine/conifer in Coconino Co., Arizona C mid-elevation oak woodland throughout the “sky islands” of southeastern Arizona (e.g. Madera Canyon in the Santa Rita Mountains) D grass/oak foothills of the Sierra Nevada Mountains, Mariposa Co., California E Zion National Park, Utah F Chihuahuan Desert below the Chiricahua Mountains in Cochise Co., Arizona G Sonoran Desert in Pinal Co., Arizona H Mojave Desert in San Bernardino Co., California I Tamaulipan thornscrub in Starr Co., Texas.

Figure 2.

Representation of Aphonopelma burrows in different habitats across the United States. A–C a typical “scrape” (burrow under rock) of A. hentzi in rocky habitat across their distribution D–E a turreted mound around the burrow of A. icenoglei (also A. atomicum, A. mojave, and A. prenticei) F the distinct crescent mound burrow of A. paloma G–I typical free-standing burrows of A. chalcodes, A. eutylenum, A. iodius, or A. johnnycashi in desert, grassland, or rocky habitats.

The taxonomy of Aphonopelma is beset with poorly delimited species boundaries and very few specimens can be confidently identified using published keys (e.g., Chamberlin 1940, Smith 1995) or comparisons to original descriptions. With few exceptions (e.g., Prentice 1993, 1997, Warriner 2008), much of the descriptive work on Aphonopelma has been based upon only one or a few specimens, generally lacked consideration for the wide range of intraspecific and intrasexual variation, and was often subjective with respect to how characters were evaluated (Prentice 1997). Structure and variation of male and female genitalia has been a heavily weighted character in delimitation of spider species, but is of limited use in Aphonopelma due to morphological homogeneity across species and may only be useful for higher-level taxonomic groups (Prentice 1997). Male mating claspers – modifications on the first two pairs of legs in adult male mygalomorph spiders used in holding and stimulating females during copulation – have also been effective at delimiting species of mygalomorph spiders (e.g., Bond 2012), however, these are mostly homogeneous across Aphonopelma as well. Differences in somatic morphology have played an important role in delimiting species boundaries, but only a handful of described species have distinct color patterns or unique combinations of characters that readily facilitate identification. This has resulted in species boundaries delimited by highly variable morphological characters (e.g., leg article proportions, number and position of spines on appendages, eye patterns, etc.). But even when multiple samples of a given species are available, they commonly fall into two categories: (1) all specimens from the same general area; or (2) from the same sex (usually adult males because they are commonly collected after abandoning their burrows in search of females). The first issue poses problems for an obvious reason: we cannot assess geographic variation in characters that might be useful for diagnosing species. The second issue is problematic because male tarantulas undergo considerable changes upon reaching maturity. Aphonopelma often display sexual dimorphism in characters given heavy weight for delimiting species boundaries, making it especially difficult to associate male and female specimens of the same species, particularly in areas where two or more species occur in syntopy.

Aphonopelma has a complicated nomenclatural history. Pocock (1901) erected the genus during his dismemberment of Eurypelma to accommodate E. seemanni (type species) and other species from the United States, Mexico, and Central America. In the same paper, he described Dugesiella (type species D. crinita). Aphonopelma and Dugesiella were distinguished by the absence of a plumose scopula on the prolateral surface of the palpal trochanter, and presence of spiniform and thorn-like (basally-swollen) setae on prolateral coxa I. Petrunkevitch (1939) later described Delopelma (type species Eurypelma marxi) and differentiated it from both Aphonopelma (by the presence of simple, recumbent hairs on coxae and trochanters) and Dugesiella (by the complete absence of plumose hairs). Chamberlin (1940) demoted Delopelma to a subgenus of Aphonopelma when he recognized the presence of plumose setae in all genera and the similar form of setae on prolateral coxa I; described another subgenus (Gosipelma, type species G. angusi); and established a new genus Chaunopelma (type species Delopelma radinum) that differed from both Aphonopelma and Dugesiella by the presence of fine, soft prone hairs on the anterior coxa and trochanter of leg I and on the posterior palpal trochanter. The nomenclature and composition of these genera remained unchanged for 45 years until Raven (1985) conducted a large-scale revision of mygalomorph genera. Raven considered the differences between Aphonopelma, Dugesiella, and Chaunopelma insignificant and artificial, synonymizing all of them with Rhechostica Simon, 1892 (type species Homoeomma texense). However, because the name Aphonopelma had been cited more extensively than Rhechostica, the ICZN was petitioned to give Aphonopelma priority over Rhechostica. Through Opinion 1637, Aphonopelma was given precedence (ICZN 1991). Smith (1995) subsequently described the genus Apachepelma to accommodate A. paloma Prentice, 1993, the first known miniature species in the genus. However, Prentice (1997) transferred the species back to Aphonopelma (thus synonymizing Apachepelma in the process). Presently, Aphonopelma is the only theraphosid genus native to the United States.

The discovery of species provides the crucial first step in the ongoing pursuit to understand the evolutionary patterns and processes shaping the biodiversity landscape. With over 250 years of taxonomic work behind us, ~1.2 million species have been described - with an estimated 7–10 million species on Earth still remaining to be described (Mora et al. 2011, Padial et al. 2010). Unfortunately, species are disappearing at an alarming rate (Barnosky et al. 2011, Vos et al. 2014), often without even being known to science, complicating our view of the Tree of Life. According to a new study, one-fifth of the world’s invertebrates are at risk for extinction, particularly species with limited vagility (Collen et al. 2012) – highlighting a concern for long-lived mygalomorph spiders. By conducting the most comprehensive sampling of a single theraphosid genus to date, this systematic revision aims to significantly broaden the scope of prior molecular (Graham et al. 2015, Hamilton et al. 2011, 2014, Hendrixson et al. 2013, 2015, Wilson et al. 2013) and morphological investigations (Prentice 1993, 1997, Smith 1995, Warriner 2008), finally bringing a modern understanding of the true species diversity in this dynamic and charismatic group of spiders, in particular the multiple species distributed throughout two biodiversity hotspots in the United States (i.e., California Floristic Province and Madrean Pine-Oak Woodlands) (Fig. 1B–D).

Abbreviations

Institutional

AMNH American Museum of Natural History; New York, New York

AUMNH Auburn UniversityMuseum of Natural History; Auburn, Alabama

BMNH The Natural History Museum, London; London, England

MNHN Muséum national d' Historie naturelle; Paris, France

NHMW Naturhistorisches Museum Wien; Vienna, Austria

NMNH Smithsonian National Museum of Natural History; Washington D.C.

Quantitative morphological landmarks (Fig. 3)

Cl length of the carapace

Cw width of the carapace

LBl labial length

LBw labial width

F1 femur I length (retrolateral aspect)

F1w femur I width

P1 patella I length

T1 tibia I length

M1 metatarsus I length

A1 tarsus I length

F3 femur III length (prolateral aspect)

F3w femur III width

P3 patella III length

T3 tibia III length

M3 metatarsus III length

A3 tarsus III length

F4 femur IV length (prolateral aspect)

F4w femur IV width

P4 patella IV length

T4 tibia IV length

M4 metatarsus IV length

A4 tarsus IV length

PTl palpal tibia length (retrolateral aspect)

PTw palpal tibia width

SC3 ratio of the extent of metatarsus III scopulation (length of scopulation/ventral length of metatarsus III)

SC4 ratio of the extent of metatarsus IV scopulation (length of scopulation/ventral length of metatarsus IV)

Figure 3.

Diagrammatic representation of informative quantitative measurements. A–B carapace, labium length and width C retrolateral palpal tibia length and width D retrolateral lengths of leg 1 femur, patella, tibia, metatarsus, tarsus, and width of femur E prolateral lengths of leg 3 femur, patella, tibia, metatarsus, tarsus, and width of femur F prolateral lengths of leg 4 femur, patella, tibia, metatarsus, tarsus, and width of femur G ventral length of scopulation (line), ventral length of metatarsus III (dashed line) H ventral length of scopulation (line), ventral length of metatarsus IV (dashed line).

Methods

Measurement, characterization, and illustration of morphological features

All material was preserved in 80% ethanol and assigned a unique alphanumeric voucher number (APH_#### or AUMS_####) added to each vial and can be used to cross-reference all images, measurements, and locality data. Quantitative measurements are reported in millimeters and were made with a Leica M165C stereomicroscope using the Leica Application Suite software and a digital camera, or from a Mitutoyo ABSOLUTE Digimatic handheld digital caliper. Appendage measurements were based on left appendages (unless otherwise stated); palpal tibia & leg I - retrolateral, legs III & IV - prolateral, extent of metatarsal scopulation - ventral. Lengths of leg articles were taken from the mid-proximal point of articulation to the mid-distal point of the article (sensu Coyle 1995, Bond 2012, Bond and Godwin 2013, see Fig. 3). Individuals were selected for measurement from > 2,900 specimens collected during this project and museum specimens from the AMNH and the AUMNH (Suppl. material 1).

Quantitative measurements are based on a minimum of five individuals of each sex, when a sufficient number of specimens were available, that represent the geographic, molecular (based on the CO1 data), and morphological breadth of each species across its distribution (i.e., every attempt was made to select specimens that represented the range of sizes available across the distribution). Material Examined sections were generated using the MATex Python script used in Bond (2012). Digital images of specimens were made using a Visionary Digital Imaging System (Visionary DigitalTM, Richmond, VA) where images were recorded at multiple focal planes and then assembled into a single focused image using the computer program Zerene Stacker v.1.04 (Zerene Systems LLC, Richland, WA). The female genital region was removed from the abdominal wall and tissues dissolved using trypsin, incubated overnight at 37oC in a 1.5ml tube; spermathecae were examined and photographed in the manner described above. All images were cropped and toned using Adobe Photoshop (Adobe Systems, Inc.). All morphological measurement datasets and images, for all species, have been deposited in the Dryad Data Repository (doi: 10.5061/dryad.k6c82).

Evaluation of quantitative morphological characters for species diagnoses

Morphometrics that were determined to have non-overlapping ranges were used as features for morphological diagnoses of species. The approach applied here is not without certain problems. For example, the inclusion of additional specimens, or in particular the case of species represented by only one or a few specimens (i.e., A. chiricahua or A. saguaro females), additional collecting could add specimens whose features expand the range of some characters and negate some of the measurements used to diagnose species. Building on the importance of certain morphological features found in Prentice (1997), we investigated 153 ratio combinations for males and 135 in females (Suppl. material 2).

To evaluate morphological variation, we examined the morphospace occupied by each species by plotting measurement ratios in boxplots (e.g., Fig. 4; Suppl. material 2), traditional PCA morphospace (e.g., Fig. 5; Suppl. material 2), and three-dimensional PCA morphospace (see GIF movies in Suppl. material 2). Simple R (R Core Team 2014) scripts, employing the packages “rgl”, “pca3d”, and “ggplot2”, have been created to evaluate new specimens either against our data or just the users data (Suppl. material 3). To evaluate PCA morphological space, all measurements (excluding ratios) were natural logarithmically transformed (i.e., ln(x)) to account for differences in body size. Traditional PCA morphospace was evaluated by plotting PC1~PC2, while three-dimensional PCA morphospace was evaluated by plotting PC1~PC2~PC3. All morphospace plots, associated scripts, and datasets have been deposited in the Dryad Data Repository (doi: 10.5061/dryad.k6c82).

Figure 4.

Examples of boxplots from male quantitative measurements used in species diagnosis (x-axis represents species). A, B, D are ratios: A carapace length/metatarsus III length B palpal tibia length/metatarsus III length D extent of scopulation on metatarsus IV C is carapace length (a proxy for body size), clearly showing the size differences between the miniature species and all other species (y-axis in mm). Additional boxplots can be viewed in Suppl. material 2.

Figure 5.

Examples of PCA morphospace plotted from quantitative measurements of males, used in species diagnosis. A a plot of males from all species in the United States B a plot of males in the Marxi species group.

Taxon sampling, molecular techniques, and phylogenetic analyses

Through extensive fieldwork and a citizen-based science program (in association with the American Tarantula Society, see http://www.atshq.org/articles/found.html), we acquired nearly 1,800 specimens of Aphonopelma and closely related sister taxa, for DNA. Specimens were opportunistically collected throughout the southwestern United States, with every attempt made to gather topotypic material from (or near) the type localities of all nominal species of Aphonopelma in the United States. Of these, A. phasmus Chamberlin, 1940 (type locality Phantom Ranch, Grand Canyon) was the only species for which we were unable to obtain fresh material. Legs were removed from all freshly collected material and preserved in ≥95% ethanol or RNAlaterTM (Qiagen, Valencia, CA, USA) and stored at -80oC. Specimens are deposited at the AUMNH, with select duplicate specimens of novel species deposited at the AMNH.

Phylogenetic analyses of Aphonopelma relationships were conducted using molecular datasets (mitochondrial and nuclear) employing likelihood optimality criteria. A new mitochondrial dataset drawn from the taxa in Hamilton et al. (2014) and from newly collected data represent the most heavily DNA-barcoded spider genus to date (n = 1032 specimens). Genomic DNA was extracted from muscle tissues using the Qiagen DNeasy Tissue KitTM (Qiagen, Valencia, CA, USA). The concentration quality of the extracted DNA was quantified with a spectrophotometer (NanoDrop ND-1000, Thermo Scientific, Wilmington, DE, USA) or visualized via agarose gel electrophoresis. PCR and direct sequencing primers used for the cytochrome c oxidase subunit I (CO1) barcoding fragment are listed in Hamilton et al. (2011); PCR protocols follow Hendrixson et al. (2013). PCR products were purified using ExoSAP-IT (USB Corporation; Cleveland, OH, USA) and then sequenced with an ABI 3130 Genetic Analyzer (Applied Bio-systems, Foster City, CA, USA) using the ABI Big Dye Terminator version 3.2 Cycle Sequencing Ready Reaction Kit. All CO1 sequences were manually edited using Sequencher (ver. 4.1.2, Genecodes, Madison, WI, USA). All CO1 sequences were aligned with MUSCLE version 3.6 (Edgar 2004) using default parameters, followed by minor adjustment in MESQUITE version 3.03 (Maddison and Maddison 2015) if needed. Amino acid translations of the target gene region were examined to ensure the absence of stop codons in the alignment. The alignments were unambiguous and for consistency, sequences were trimmed to 900 basepairs. The program PartitionFinder v1.1.0 (Lanfear et al. 2014) was used to determine the appropriate model of DNA substitution. Each gene region was partitioned by codon position with separate models chosen for each partition. Maximum Likelihood (ML) analyses were carried out in RAxML-7.3.1 (Stamatakis 2006) on the CASIC HPC at Auburn University. Parameters for the analyses incorporated the GTRGAMMA model of evolution based on 10,000 random addition sequence replicates; branch support values were computed via 1000 non-parametric bootstrap replicates. The tree was rooted using divergent Central American species: Sericopelma sp. (APH_3003, 3004, 3020) from Panama; Aphonopelma belindae (APH_3001, 3008) from Panama; and A. sp. xanthochromum (APH_3024) from Costa Rica. All previous CO1 sequences were deposited in GenBank and can be found in Hamilton et al. (2011, 2014), Hendrixson et al. (2013, 2015), and Graham et al. (2015); all new CO1 sequences have been deposited in GenBank (accession numbers: KU054417-KU055448). DNA sequence alignments and associated phylogenetic trees and data matrices, accompanying tree files, and scripts have been deposited in the Dryad Data Repository (doi: 10.5061/dryad.k6c82).

The analyses described above and recent smaller scale molecular studies (Hamilton et al. 2011, 2014, Hendrixson et al. 2013, 2015, Wilson et al. 2013, Graham et al. 2015) illustrate the potential that molecular data have for both separating and identifying known and unknown species within Aphonopelma. Unfortunately, all of these studies relied largely upon mitochondrial markers. The limitations of mtDNA have been documented extensively, namely, gene tree/species tree incongruence and the haploid, non-recombining nature of the molecule (Maddison 1997, Funk and Omland 2003, Knowles and Kubatko 2010). With mtDNA representing only one particular genealogy out of all possible within a genome, it is unlikely to fully resolve Aphonopelma phylogeny or demographic history accurately – though the use of mtDNA data represent an important first-step in untangling the complicated taxonomic problem that has developed for the tarantula fauna in the United States.

The use of nuclear loci generally has led to an enhanced understanding of the Tree of Life, but unfortunately, many non-model invertebrates (e.g., spiders) lack adequately developed loci for targeted sequencing. Until very recently, few genetic markers were available for inferring spider phylogenies (i.e., only 13 independent markers though 2013, see Gillespie et al. 1994, Hedin 1997, Agnarsson et al. 2007, Ayoub et al. 2007, Masta et al. 2009, Hamilton et al. 2011, Bond et al. 2012, Hendrixson et al. 2013, Satler et al. 2013). The poor resolution provided by traditional nuclear markers (e.g., 28S, 18S, H3) for inferring shallower relationships indicates many more loci need to be developed in order to overcome the stochasticity of sequence evolution and gene trees. Next-generation sequencing (NGS) has transformed molecular phylogenetics by enabling systematists to more confidently resolve major branches on the Tree of Life by gathering vast quantities of genomic data with relative ease. Given the difficulties in theraphosid systematics, novel genomic data is desperately needed to accurately delimit species and to reconstruct a robust phylogenetic framework for the group. To do this, a novel targeted sequencing approach was modified for use in spiders (Hamilton et al. in prep). Anchored Enrichment (AE) is a relatively new NGS methodology designed to recover hundreds of unique loci (i.e., single copy informative markers) from across the genome to resolve relationships at all taxonomic depths (Lemmon et al. 2012). Originally developed for vertebrates, Hamilton et al. (in prep) adapted this powerful approach to work in mygalomorph spiders by using six arachnid genomes and 17 transcriptomes to identify 585 loci. When possible, two specimens of each CO1 species were sequenced, selecting specimens that encompassed the geographic and genetic breadth of each species. Unfortunately, at the time of our AE sequencing, some CO1 species were singletons or we lacked good nuclear DNA for sequencing (i.e., old specimens whose tissue was not preserved for genomic sequencing). A total of 80 OTUs, including three outgroup taxa (Aphonopelma belindae (APH_3001), A. sp. burica (APH_3012), and Sericopelma sp. (APH_3004)) were selected for sequencing and phylogenetic inference.

Anchored Enrichment data were collected through the Center for Anchored Phylogenomics at Florida State University (http://www.anchoredphylogeny.com) following the general methods of Lemmon et al. (2012), modified in Lemmon et al. (in prep), and applied to spiders in Hamilton et al. (in prep). Tissue preparation and DNA extraction are the same as above. Briefly, each genomic DNA sample was sonicated to a fragment size of ~300-800bp, library preparation and indexing were performed following a protocol modified from Meyer and Kircher (2010). Indexed samples were then pooled at equal quantities and enrichments were performed on each multi-sample pool using an Agilent Custom SureSelect Kit (Agilent Technologies; herein referred to as the Spider Probe Kit), which contained probes designed for anchored loci from multiple spider genomes and transcriptomes. After enrichment, each set of enrichment reactions were pooled in equal quantities for sequencing on PE150 Illumina HiSeq2000 lanes. Sequencing was performed in the Translational Science Laboratory in the College of Medicine at Florida State University.

Utilizing the bioinformatics pipeline described in Lemmon et al. (2012), modified in Lemmon et al. (in prep), and applied to spiders in Hamilton et al. (in prep), paired-end sequencing reads were filtered for quality. Reads were demultiplexed and pairs were identified and merged following Rokyta et al. (2012). Reads were assembled into contigs using an assembler (Lemmon et al. in prep) that makes use of both a divergent reference assembly approach to map reads to the probe regions and a de-novo assembly approach to extend the assembly into the flanks. Reads were mapped to the spider probes, consensus bases called, contamination filtered, consensus sequences were grouped by locus (across individuals) in order to produce sets of homologs, and orthology determined. From this, 455 loci (229,854bp) were selected for use in Aphonopelma. Sequences in each orthologous set were aligned using MAFFT v7.023b (Katoh 2013), with --genafpair and --maxiterate 1000 flags utilized. All alignments were visually investigated in Geneious Pro v5.6 (http://www.geneious.com, Kearse et al. 2012) for consistency. A concatenated supermatrix of all loci was constructed to infer relationships using Maximum Likelihood (ML) phylogenetic inference in RAxML v7.3.1 (Stamatakis 2006). Parameters for the concatenated RAxML analysis incorporated the GTRGAMMA model of evolution based on 1000 random addition sequence replicates, branch support values were computed via 1000 non-parametric bootstrap replicates, and partitions were set for each locus. Aphonopelma belindae (APH_3001) was designated as the outgroup. Prior analyses (Turner et al. in review), and our preliminary analyses have inferred Sericopelma to be more closely related to the Aphonopelma in the United States and Mexico, than to the Central American Aphonopelma. Phylogenies for each individual locus were inferred using RAxML under the same parameters as above, with subsequent species tree estimation performed in ASTRAL v4.7.6 (Mirarab et al. 2014). All investigations were carried out on the CASIC HPC at Auburn University and the Florida State University HPC.

Locality data, georeferencing, generation of niche-based distribution models, and conservation status

For all newly collected samples, latitude and longitude were recorded in the field using a Global Positioning System (GPS) receiver (WGS84 datum) in Decimal Degrees (DD). For previously collected museum specimens, locality data were manually georeferenced using Google Earth (Google, Mountain View, CA) or Topo North America (DeLorme, Yarmouth, ME) in Decimal Degrees (DD). All georeferenced and field recorded locality data (latitude, longitude, elevation) were crosschecked by hand in Google Earth, Topo North America, or ArcGIS (ESRI, Redlands, CA) prior to generating distribution maps, niche-based distribution models, and database entry. Distribution maps were constructed in ArcGIS. Because older locality labels often lack sufficient locality information, many georeferenced values are imprecise and should be interpreted with caution. As well, because these older labels are often faded or handwritten, there may be slight discrepancies in the spelling of localities, collectors, etc. Data for labels that document only county and/or town information were georeferenced to the approximate geographic center of the given locality. Precision for each georeferenced point is annotated as a superscript in the material examined section for each species using the confidence value scheme employed by Murphey et al. (2004) and Bond (2012) and modified herein: 1 = exact coordinates given; 2 = exact location given, validated in Google Earth; 3 = public land survey (or herein geographic place name); 4 = within 1 km radius (~.5 mile); 5 = within 5 km radius (~3 miles); 6 = within 10 km radius (~6 miles); 7 = to county or > 10 km; 8 = to state; 9 = to project region. Detailed locality and associated GIS information can be found in Suppl. material 1. See Figure 6 for a generalized map of all collecting sites of all species across the United States.

As an approach to facilitate species discovery and delimitation, niche-based distribution models (DMs) were constructed for species for which sufficient locality data were available (i.e., at least 10 different localities separated by at least 1 km). Niche-based DMs provide estimates for the probability of finding a species at a location on the landscape given the set of correlate ecological and climatic parameters used to construct the model. Locality coordinates for each specimen were imported into ArcMap (ESRI, Redlands, CA) and converted into shapefiles. Following the procedures outlined in Bond and Stockman (2008) and as previously implemented in Aphonopelma (Hendrixson et al. 2013, Graham et al. 2015), DMs were constructed using environmental layers thought to “likely influence the suitability of the environment for the species” (Phillips et al. 2006, p. 232). We selected eight environmental layers from the WORLDCLIM data set (Hijmans et al. 2005) based largely on the arguments presented by Bond and Stockman (2008): ALT (elevation), BIO4 (Temperature Seasonality), BIO5 (Max Temperature of Warmest Month), BIO6 (Min Temperature of Coldest Month), BIO12 (Annual Precipitation), BIO15 (Precipitation Seasonality), BIO16 (Precipitation of Wettest Quarter), and BIO17 (Precipitation of Driest Quarter). All layers were clipped to the same extent, cell size, and projection. DMs were created in the program Maxent ver. 3.3.3k (Phillips et al. 2006) using default parameters.

Figure 6.

A generalized distribution map of all unique collecting localities from across the United States.

A hypothesized conservation status for each species has been included with each respective description. The designations provided herein are not based on any formal calculations and therefore should not be viewed as formal status declarations. These designations are based upon our own extensive fieldwork and the observations of fellow arachnologists (e.g., Tom Prentice and Wendell Icenogle). As such, our estimation of the conservation status for each species is likely conservative.

Species delimitation and conceptualization

The species concept used throughout this taxonomic revision utilizes the ideas of de Queiroz’s Unified Species Concept (2005). Where possible, we employ a combination of molecular phylogenetic, morphological, behavioral, and biogeographic evidence to identify independently evolving lineages. If not possible, an alternative species concept is noted. Initial species hypotheses were based on the recognized morphological species hypotheses (World Spider Catalog 2015). To test these initial species hypotheses, we evaluated the agreement of mtDNA (CO1) with the morphological species hypotheses or putative new lineages, following the methodology laid out in Hamilton et al. (2014). Morphological species boundaries were then reevaluated based on CO1 species boundaries. We then evaluated whether there was agreement of biogeographical and behavioral data with these species hypotheses. Finally, we employed the nuclear loci developed using Anchored Enrichment (Hamilton et al. in prep) to evaluate those species hypotheses and establish whether independent lineages have sorted. Following this final delimitation of species, a reciprocal illumination approach (see Lienau et al. 2006 and Padial et al. 2010) was employed to find informative morphological characters that can be used (absent of molecular data) across the group. This implementation of a tree-based approach attempts to identify specimens to species by phylogenetic association, providing evidence of common ancestry with specimens identified by other means (e.g., morphological or cohesion species criteria).

Data resources

All data (molecular, morphological, geographic, and images) used to establish these species hypotheses have been deposited in the Dryad Data Repository (https://doi.org/10.5061/dryad.k6c82). All locality data underpinning the analysis reported in this paper are deposited at GBIF, the Global Biodiversity Information Facility, http://ipt.pensoft.net/resource?r=aphonopelma. All morphological images have been deposited in Morphbank, and can be viewed by referencing the "APH-S" Specimen External Identifier. Additional specimen data, species plates, and morphological data can be found in the Suppl. materials.

Results and discussion

Summary of taxonomic diversity

Prior to our work leading up to this taxonomic revision (Hamilton et al. 2011, 2014, Hendrixson et al. 2013, 2015, Graham et al. 2015), only one other study had implemented a phylogenetic approach to understanding species diversity and evolutionary relationships in Aphonopelma (Wilson et al. 2013). A well-supported phylogeny is fundamental to addressing important questions regarding the role that biogeography, allopatry, ecological divergence, and ancestral interactions have played in the diversification of these lineages. Our results demonstrate that several species in the United States need to be synonymized (see Hamilton et al. 2011, 2014), novel species need to be named and described (Hamilton et al. 2011, 2014, Hendrixson et al. 2013, 2015, Graham et al. 2015), and a handful of species must be considered nomina dubia. Of the 55 nominal species of Aphonopelma reported from the United States prior to the present contribution, we only consider 15 of them valid (i.e., we propose 33 new synonymies and consider 7 species nomina dubia). Despite this significant reduction in the number of valid species, we also recognize 14 novel species (described herein), bringing the total diversity of Aphonopelma in the United States to 29 species. It is important to point out that geographic distribution is crucial to understanding Aphonopelma diversity within the United States.

Aphonopelma phylogeny

Following the integrative approach for delimiting species within Aphonopelma outlined in Hamilton et al. (2014), we increased CO1 taxon coverage (n = 1032) to further investigate mtDNA species boundaries in the group by including more OTUs per species and by adding new putative species to the analysis. With the exception of the newly added species, relationships are generally consistent with our previous analyses (see Hamilton et al. 2014) (e.g., the same cryptic species lineages are delimited and the same hypothesized species are highly supported with deeper support throughout the tree decreasing as more OTUs were added; Fig. 7). The mitochondrial data suggest that we should recognize 41 species in the United States (not including A. phasmus because we did not have tissue for collecting CO1 data), with 17 nominal species and 24 previously undescribed species.

Figure 7.

Maximum Likelihood inferred CO1 gene tree phylogeny for the 1,032 Aphonopelma specimen dataset. Species delimitations followed the integrative methodological approach outlined in Hamilton et al. (2014). Black circles denote bootstrap support ≥ 80%; white squares denote bootstrap support ≤ 80%. White triangles indicate species clades supported with ≥ 80% bootstrap support; grey triangles indicate lineages with putative mitochondrial introgression events. Asterisks at the tips of branches indicate undescribed diversity.

To further assess species-level diversity and evaluate the associated hypothesized species boundaries delimited by the CO1 data, we sequenced multiple specimens per putative species using Anchored Enrichment. A dataset of 455 loci (229,854 basepairs) across 80 OTUs produced a highly resolved species-level phylogeny of all the major species groups and regional clades within Aphonopelma (Fig. 8). These AE data identify 14 nominal species, 14 undescribed species, and 2 lineages that we will investigate in more depth in the future (labeled as sp. n. 1 and sp. n. 2 in Fig. 8; again, this analysis does not include A. phasmus). All species are strongly supported as representing genealogically exclusive, independently evolving lineages, with one exception, A. iodius (details below). Figure 8 summarizes the maximum likelihood inference of the AE molecular data and subsequent species tree inference, highlighting the monophyly and high support of the five major clades (see Fig. 8) of Aphonopelma within the United States: (1) a monotypic group that is confined to California; (2) a western group; (3) an eastern group; (4) a group primarily distributed in the high-elevation sky islands areas of Arizona and New Mexico; and (5) a group that comprises a diverse group of miniaturized species. The area that divides the western and eastern groups roughly corresponds with the Cochise Filter Barrier biogeographic region (see Pyron and Burbrink 2010) of southeastern Arizona and southwestern New Mexico, where the Chihuahuan and Sonoran deserts converge. The monotypic lineage includes the species A. steindachneri (herein referred to as the Steindachneri species group); the western lineage includes the Iodius species group (A. chalcodes, A. eutylenum, A. iodius, and A. johnnycashi sp. n.); the eastern lineage includes the Hentzi species group (A. anax, A. armada, and A. hentzi) and Moderatum species group (A. gabeli, A. moderatum, and A. moellendorfi sp. n.). By far, the greatest amount of novel diversity is found in the two remaining clades: the Marxi species group, a lineage that includes predominately black species (both adult males and females) and often found in montane or other high-elevation habitats, in particular the sky islands region of southwestern New Mexico and southeastern Arizona (A. catalina sp. n., A. chiricahua sp. n., A. madera sp. n., A. marxi, A. peloncillo sp. n., and A. vorhiesi); and the Paloma species group, a collection of closely-related species that have evolved a small size relative to most other species in the United States (A. atomicum sp. n., A. icenoglei sp. n., A. joshua, A. mareki sp. n., A. mojave, A. paloma, A. parvum sp. n., A. phasmus, A. prenticei sp. n., A. saguaro sp. n., A. superstitionense sp. n., and A. xwalxwal sp. n.). Each species is genealogically exclusive and strongly supported (≥80 bootstrap support) (Fig. 8).

Figure 8.

Species tree of all United States Aphonopelma, inferred from the 455 loci Anchored Enrichment dataset. Species delimitations correspond to our final integrative approach outlined herein. Black circles denote 100% bootstrap support; black squares denote bootstrap support between 99–80%; white squares denote bootstrap support less than 80%. Node support values = based on the RAxML bootstrap support from all trees and all loci. All genealogically exclusive species are identified with a grey bar; A. iodius, a paraphyletic species as presently defined, is identified by the black boxes. All major species groups are identified by colored boxes.

While our methodological approach using CO1 identified the broad effectiveness of this 900 basepair fragment of mtDNA to identify known species and help illuminate species boundaries across the most comprehensive molecular sampling of a single spider genus to date, the AE nuDNA data robustly highlight where deep mitochondrial divergences and introgression have obscured our understanding of the true evolutionary history of these lineages. It is important to point out that a number of the putative mitochondrial species corresponded to lineages that were considered cryptic species in Hamilton et al. (2014), yet when viewed in the light of the AE data, are no longer distinct. This is perhaps not very surprising given the large number of papers showing how the use of a single locus, like mtDNA, to delimit species can be very misleading (e.g., Rubinoff et al. 2006, Petit and Excoffier 2009).

Steindachneri species group

The Steindachneri species group presently includes a single species (A. steindachneri) from California. The CO1 and AE datasets both support the “basal” position of A. steindachneri as the sister taxon to all other species of Aphonopelma in the United States (Figs 7, 8). There are likely more species in northwestern Mexico that will be placed into this group, in particular the Baja Peninsula.

Western species group diversity (Iodius species group)

The western group of species (note: the western designation is only applied here in an informal sense because other lineages are distributed west of the Cochise Filter Barrier) is comprised entirely of the Iodius species group (A. chalcodes, A. eutylenum, A. iodius, and A. johnnycashi sp. n.) (Fig. 8). The AE dataset identifies A. chalcodes as the sister taxon to the remaining members of the Iodius species group, with A. eutylenum the sister lineage to the A. iodius species complex. The CO1 and AE datasets recognize A. eutylenum as a distinct and genealogically exclusive species. The recognition of A. johnnycashi sp. n. renders A. iodius paraphyletic (see below).

Unfortunately, the mitochondrial data are complex. Results from the CO1 analysis confounds our understanding of species boundaries and relationships in Aphonopelma, likely due to mitochondrial introgression or deep haplotype conservation (e.g., one mtDNA lineage of A. chalcodes is sister to A. vorhiesi, a species that belongs to the Marxi species group, while another mtDNA lineage of A. chalcodes is sister to other A. iodius lineages). Alternatively, the AE nuclear data provide support for these species (A. chalcodes, A. vorhiesi) as independently evolving and monophyletic lineages. In previous analyses (Hamilton et al. 2014), we would have considered the two mtDNA lineages of A. chalcodes as separate cryptic species. We also see that A. iodius, as presently defined, is a paraphyletic species with lineages in Utah comprising a highly supported, genealogically exclusive lineage that is sister to the new species A. johnnycashi (from the western foothills of the Sierra Nevada). The Utah lineage is separate from other A. iodius lineages that are currently lumped into one species (including the species A. iviei and A. brunnius which have been synonymized with A. iodius). Unfortunately, at this time we do not have any additional evidence that can be used to separate the Utah lineage from these other lineages. Future work will need to focus on sequencing more individuals within this group, evaluating gene flow between populations, and looking more closely at the biogeographic history of these lineages to determine if/where the geographic and genetic split occurs.

Eastern species group diversity (Hentzi and Moderatum species groups)

The eastern group of species is designated as such because the six species are largely distributed east of the Cochise Filter Barrier. This monophyletic lineage is strongly supported and includes the Hentzi species group (A. anax, A. armada, and A. hentzi) and Moderatum species group (A. gabeli, A. moderatum, and A. moellendorfi sp. n.) (Fig. 8). The three species in the Hentzi species group are phenotypically similar and are the only species in the United States that possess stout setae on the prolateral surface of coxa I; freshly molted specimens typically possess black legs, an abdomen with short black setae and longer reddish setae, and a copper, brown, or tan carapace. Within the highly supported, monophyletic Hentzi species group A. armada is inferred as the sister lineage to A. hentzi and A. anax. The Moderatum species group is composed of three species whose adult males undergo significant color changes upon reaching sexual maturity (i.e., usually becoming solid black and fading over the course of their respective breeding periods). Within the highly supported, monophyletic Moderatum species group, A. moderatum is the sister lineage to A. gabeli and A. moellendorfi.

When reviewing both the putative mitochondrial and cryptic species identified in Hamilton et al. (2014), we can see how the mtDNA analyses confound our understanding of species boundaries and relationships in Aphonopelma (e.g., several species delimited by the mtDNA data are no longer recognized by the AE data) (see Fig. 7 and Hamilton et al. 2014).

Sky islands diversity (Marxi species group)

The Marxi species group includes A. catalina sp. n., A. chiricahua sp. n., A. madera sp. n., A. marxi, A. peloncillo sp. n., and A. vorhiesi (Fig. 8). Freshly molted specimens of these six species are generally black (adult males and females) and can often be found in the montane or other high-elevation habitats throughout the Madrean sky islands of southwestern New Mexico and southeastern Arizona; exceptions include A. vorhiesi and some populations of A. peloncillo which can be located in lower-elevation grassland or desert habitats, and A. marxi which is found in the montane habitats of northern Arizona and New Mexico, as well as southwestern Colorado and southeastern Utah. Phylogenetic relationships are generally similar between the CO1 and AE datasets, although the placement of the A. marxi and A. vorhiesi lineages within the CO1 phylogeny has always been problematic (e.g., low support, suspect sister relationships). Outside of A. vorhiesi, where both mitochondrial introgression and deep mitochondrial divergence obscure an understanding of species boundaries and relationships, there are no putative mitochondrial or cryptic species identified in Hamilton et al. (2014) that were invalidated by the nuclear and morphological data.

Despite our extensive collecting throughout this region, there likely still remains undescribed diversity in this species group (Figs 78). Our lack of specimens (particularly for the Madrean sky island species) is due to the difficulty of locating these spiders (see Hendrixson et al. 2015). The terrain throughout the sky islands makes it challenging to do field work because many of the mountain ranges are incredibly remote or otherwise difficult to access (due to rough roads or private land); this is confounded by the cryptic nature of these species (i.e., burrows are incredibly difficult to find throughout the rocky and forested habitats in these regions). A thorough understanding of the diversity across this unique biodiversity hotspot is going to require future funding and collaborative collecting efforts. Several other large mountain ranges in the region (e.g., the Dragoon, Galiuro, and Pinaleño Mountains) possess appropriate habitat but have not been rigorously searched for these tarantulas.

Miniaturization and diversification in Aphonopelma (Paloma species group)

The Paloma species group includes a dozen miniaturized species primarily located in the Mojave and Sonoran deserts: A. atomicum sp. n., A. icenoglei sp. n., A. joshua, A. mareki sp. n., A. mojave, A. paloma, A. parvum sp. n., A. phasmus, A. prenticei sp. n., A. saguaro sp. n., A. superstitionense sp. n., and A. xwalxwal sp. n. (Fig. 8). The lineages within this group have a particularly interesting natural history. Wherever these species are located, they reside in syntopy with a larger species (generally from the Iodius species group), yet are rarely found in syntopy with other members of the Paloma species group. The larger species typically possess distributions that extend beyond that of individual members of the Paloma species group; furthermore, these larger species can often be found in syntopy with multiple members of the Paloma species group at different places across their distribution.

The CO1 data can be used to confidently identify the species in this group (probably due to the relatively high mtDNA divergence between species), except A. mareki and A. superstitionense due to putative mitochondrial introgression. Unfortunately, interspecific relationships are not well supported and the placement of A. paloma and A. xwalxwal is problematic (i.e., not consistently recovered in the same place in the phylogeny). When the AE data are employed, the Paloma species group is strongly supported and the placement of A. paloma and A. xwalxwal is confidently resolved. Most relationships within and between the remaining species are also strongly supported and highly resolved (Fig. 8). After reviewing the putative species identified in Hamilton et al. (2014), we see there are no mitochondrial or cryptic species that were invalidated by the nuclear and morphological data.

Body size is one of the most important determinants of an organism’s ecological role (Hanken and Wake 1993, and citations within). While the literature is replete with cases of miniaturization in sympatry due to character displacement (e.g., Bond and Sierwald 2002, Hendrixson and Bond 2005), these examples seemingly represent single events and do not appear to have led to subsequent lineage diversification within the smaller taxa. In Aphonopelma, a single origin of miniaturization is strongly supported and this event undoubtedly played a role in the diversification of these spiders in the United States (we recognize 13 species in this group, eight of which are new). We hypothesize that this miniaturization event (and associated changes in niche utilization), evolved as a consequence of ancestral character displacement. Following this shift in body size (a pre-zygotic isolating mechanism), members of the Paloma species group may have been more able to capitalize on available microhabitats where the larger species were not as abundant (Prentice 1997, pers. obs.), allowing persistence in syntopy with larger taxa and leading to subsequent lineage diversification of these smaller taxa. In addition, these miniaturized species have limited dispersal capabilities and are probably more likely to undergo allopatric divergence than their larger counterparts (see Graham et al. 2015). Future research aims to investigate the processes that led to miniaturization and increased rates of diversification within the Paloma species group.

Species considered nomina dubia

Chamberlin (1940) described the species Aphonopelma baergi from Fayetteville, Arkansas. As noted by Smith (1995) and confirmed by our own examination of the holotype (AMNH), this species was likely mislabeled by the original collector. This large and robust tarantula is noticeably different from other species in the United States based on the morphology of its spermathecae (refer to Smith 1995, fig. 141); in addition, surveys of tarantulas in the region (i.e., Warriner 2008 and our own fieldwork) have failed to locate specimens that share diagnostic features with the holotype (all other tarantulas from Arkansas and surrounding states are referable to A. hentzi (Girard, 1852)). Smith (1995) suggested that the species is most closely allied with members of the Mexican and Central American genus Brachypelma Simon, 1891 but did not propose any formal taxonomic changes (see Peters 2003, 2005, Rudloff 2008). Based on the holotype’s questionable origin, however, we do not recommend transferring this species to Brachypelma but instead consider the name A. baergi a nomen dubium.

The species Aphonopelma cratium Chamberlin, 1940 is based on a male holotype and female allotype (both AMNH – examined) from an uncertain location in California (“California ?” in the locality information on the label). These specimens are morphologically similar to other valid species in the state (e.g., A. iodius (Chamberlin & Ivie, 1939) and A. eutylenum Chamberlin, 1940), but because molecular data and/or accurate locality information are needed to distinguish these species, we consider A. cratium a nomen dubium.

Smith (1995) described Aphonopelma hollyi on the basis of a single adult male specimen from Lubbock, Texas. Efforts to locate the holotype in the Oklahoma State University collection were unsuccessful and the specimen is presumed lost (Richard Grantham 2013, pers. comm.). We were unable to locate new topotypic material from Lubbock for purposes of designating a neotype but our extensive fieldwork in surrounding parts of the Llano Estacado and Caprock Escarpment strongly suggests that only two species with thorn-like setae on the prolateral surface of coxa I can be found there: A. hentzi and A. armada (Chamberlin 1940). Our initial thinking on the identity of A. hollyi was that it should be considered a junior synonym of A. hentzi. However, the description of A. hollyi is vague, preventing one from distinguishing it from either A. hentzi or A. armada. Because the holotype has been lost and its identity is uncertain, we consider A. hollyi a nomen dubium.

The description of Eurypelma mordax Ausserer, 1871 was based on a single adult male but the holotype was likely destroyed during World War II (see Smith 1995). After several decades, Roewer (1942) eventually synonymized E. mordax with A. hentzi (then Dugesiella hentzi), but as noted by Smith (1995), the synonymy was questionable because Roewer never examined the type specimen. Based upon his own examination of material in the Koch collection at the BMNH, Smith (1995) found a female specimen from Texas that was labeled Eurypelma mordax by Ausserer and removed the species from synonymy of A. hentzi. We, however, consider the identity of Aphonopelma mordax suspect for the following reasons: (1) the collection site for the destroyed holotype male is unclear; (2) Smith (1995) did not formally designate the female specimen as a neotype; (3) the female’s collection site in Texas is also unclear; and (4) because we do not have precise locality information for either specimen, there is no compelling reason to conclude that they belong to the same species. In the absence of additional useful information, we consider Eurypelma mordax = Aphonopelma mordax a nomen dubium.

Two different type localities (Ft. Yuma and Williams, Arizona) were listed in the description of Eurypelma rusticum Simon, 1891 but it is unclear which specimen was actually used for the description (Prentice 1997). In subsequent attempts to determine the precise identity of this species, both Smith (1995) and Prentice (1997) failed to locate the specimen from Ft. Yuma. In the original description, Simon (1891) also mentioned that the species was common in northern Mexico and this prompted Smith (1995) to designate a lectotype from Simon-determined paratype material in the MNHN; curiously, Smith’s choice of a lectotype (MNHN No. 5873 – not examined) from Mazatlán, Mexico is located 1300 kilometers southeast of Ft. Yuma in strikingly different habitat (see Prentice 1997). Three years later, Prentice (1997) examined what he thought might be Simon’s original specimen from Williams in the NMNH (USNM No. 1585 – not examined). He noted that the badly fragmented specimen was not conspecific with Smith’s lectotype, but did not make any changes; instead, he stated “reexamination of all type material will be necessary before a lectotype can be objectively designated” (Prentice 1997, pg. 146). Based on this confusion, we consider Eurypelma rusticum = Aphonopelma rusticum a nomen dubium because: (1) Simon’s description does little to help diagnose the species; (2) the type specimen from Ft. Yuma is lost; (3) the type specimen from Williams is in poor condition; (4) the lectotype from Mazatlán is not conspecific with Simon’s specimen from Williams; and (5) we have serious doubts that the lectotype from Mazatlán is conspecific with material from Yuma, Arizona and surrounding areas. As also discussed by Smith (1995) and Prentice (1997), specimens of this species cited in Chamberlin (1940) from Apache Trail, Arizona were described as A. rothi Smith, 1994 (= A. chalcodes Chamberlin, 1940, see below).

The enigmatic species Delopelma radinum (Chamberlin & Ivie, 1939) was described on the basis of a single adult male collected from Manhattan Beach, California in November 1937. To our knowledge, no specimens comparable to the badly fragmented holotype (AMNH – examined; see supplemental morphology images in Suppl. material 4) have been collected or reported from the Los Angeles Basin since the original description. While it is possible that the species has gone extinct (see Bond et al. 2006 and Bond 2012 for further discussion of mygalomorph spider extinctions in California), we strongly suspect that the type locality is incorrect because the habitat is atypical for North American tarantulas (see Prentice 1997). Morphological data reported in Prentice (1997, pg. 142) and the November collection date of the holotype, suggest that the species is probably Aphonopelma mojave Prentice, 1997 or A. icenoglei sp. n. from the western Mojave Desert. However, we consider Delopelma radinum = Chaunopelma radinum = Aphonopelma radinum a nomen dubium for the following reasons: (1) despite extensive fieldwork throughout the area, we have failed to locate any specimens; (2) the type locality is highly dubious; (3) without molecular data, we cannot definitively associate the species with A. mojave or A. icenoglei.

Simon (1891) also described Homoeomma texense on the basis of a single adult male from “Texas: Rio-Grande (Geo. Marx)”. Similar to the situation with Aphonopelma baergi discussed above, this species is markedly different from other species in the United States based on the morphology of its palpal bulb (Smith 1995, fig. 799, but see Prentice 1997, Smith 2010); in addition, the type locality is vague and may include any portion of the river from its origin in Texas near El Paso to its mouth in the Gulf of Mexico (a distance of approximately 2000 kilometers that passes through numerous physiographic provinces). No other specimens with this unique palpal bulb morphology have ever been found in Texas; furthermore, Smith (2010) suspects that the holotype was mislabeled (the collector, George Marx, was notorious for mislabeling specimens) and may belong to the South American genus Paraphysa Simon, 1892. For these reasons, we consider Homoeomma texense = Rhechostica texensis = Rhechostica texense = Aphonopelma texensis = Aphonopelma texense a nomen dubium.

Taxonomy

Family Theraphosidae Thorell, 1869
Subfamily Theraphosinae Thorell, 1870

Aphonopelma Pocock, 1901

Rhechostica Simon, 1892: 162 (type species by original designation Homoeomma texense Simon, 1891). Suppressed as a senior synonym of Aphonopelma by ICZN Opinion 1637.

Dugesiella Pocock, 1901: 551 (type species by original designation Dugesiella crinita Pocock, 1901). First synonymized with Rhechostica by Raven (1985: 152).

Aphonopelma Pocock, 1901: 553 (type species by original designation Eurypelma seemanni Pickard-Cambridge, 1897). First synonymized with Rhechostica by Raven (1985: 149).

Delopelma Petrunkevitch, 1939: 567 (type species by original designation Eurypelma marxi Simon, 1891). First synonymized with Rhechostica by Raven (1985: 151).

Gosipelma Chamberlin, 1940: 4 (type species by original designation Gosipelma angusi Chamberlin, 1940). Originally described as a subgenus of Aphonopelma, but never elevated to full generic status. First synonymized with Rhechostica by Raven (1985: 153).

Chaunopelma Chamberlin, 1940: 30 (type species by original designation Delopelma radinum Chamberlin & Ivie, 1939). First synonymized with Rhechostica by Raven (1985: 151).

Apachepelma Smith, 1994: 45 (type species by original designation Aphonopelma paloma Prentice, 1992). First synonymized with Aphonopelma by Prentice (1997: 147).

Type species

Eurypelma seemanni F.O. Pickard-Cambridge, 1897; female holotype from Puerto Culebra, Pacific coast, W of Liberia, Guanacaste province, Costa Rica, coll. Dr. Seeman; deposited in BMNH. [examined]

Eurypelma seemanni F. O. Pickard-Cambridge, 1897: 26.

Aphonopelma seemanni Pocock, 1901: 553.

Aphonopelma seemanni Valerio, 1980: 274.

Rhechostica seemanni Raven, 1985: 149.

Aphonopelma seemanni Smith, 1995: 141.

Diagnosis

From Prentice (1997, p. 147)

“The genus Aphonopelma is distinguished from all other theraphosid genera by the following combination of characters: (1) no known external stridulation organs; (2) hair-like or spiniform plumose setae on the prolateral surface of the trochanter and femur of leg I and on the retrolateral surface of the coxa and trochanter of the pedipalp; (3) type I urticating setae only; (4) corresponding segments of all legs approximately the same width in females (femur III in males sometimes laterally swollen); (5) scopula of tarsus IV usually entire, if divided then only partially and narrowly by line of setae; (6) setae on the prolateral surface of coxa I hair-like and not basally swollen, spiniform and basally swollen, or distinctly stout and thorn-like; (7) metatarsus I flexing against the lower process of the tibial spur, with either the apex of the spur contacting the ventral surface of the metatarsus or the outer edge of the spur in the apical half contacting the prolateral surface of the metatarsus; (8) and the lower process of the tibial spur curving prolaterodistally and widening apically, usually equipped with at least one apical or preapical megaspine, and the upper shorter process less stout basally, relatively uniform in diameter throughout its length, and equipped on its inner surface with at least one stout, basally articulated megaspine.”

Future nomenclatural status of Aphonopelma

Due to our inability to conduct fieldwork in Mexico and Central America, a comprehensive taxonomic revision of Aphonopelma (and other closely related genera) is not feasible at this time. However, we do anticipate future genus-level nomenclatural changes for those species found in the United States. As presently defined, we have reason to believe that Aphonopelma is not monophyletic. Our choice of outgroups places the Aphonopelma species from the United States closer to the genus Sericopelma than to the Central American species of Aphonopelma (Fig. 8). This has nomenclatural implications because the type species (A. seemanni) from Costa Rica is undoubtedly grouped with the other Central American Aphonopelma material (pers. obs.). Unfortunately, it is not possible to resurrect a synonymized generic name (e.g., Dugesiella or Delopelma) for the Aphonopelma species in the United States until researchers can more thoroughly sample material from Mexico and Central America (see Prentice 1997, Smith 2010).

Aphonopelma

within the United States

Aphonopelma anax (Chamberlin, 1940)

Aphonopelma armada (Chamberlin, 1940)

Aphonopelma atomicum Hamilton, sp. n.

Aphonopelma catalina Hamilton, Hendrixson & Bond, sp. n.

Aphonopelma chalcodes Chamberlin, 1940

Aphonopelma chiricahua Hamilton, Hendrixson & Bond, sp. n.

Aphonopelma eutylenum Chamberlin, 1940

Aphonopelma gabeli Smith, 1995

Aphonopelma hentzi (Girard, 1852)

Aphonopelma icenoglei Hamilton, Hendrixson & Bond, sp. n.

Aphonopelma iodius (Chamberlin & Ivie, 1939)

Aphonopelma johnnycashi Hamilton, sp. n.

Aphonopelma joshua Prentice, 1997

Aphonopelma madera Hamilton, Hendrixson & Bond, sp. n.

Aphonopelma mareki Hamilton, Hendrixson & Bond, sp. n.

Aphonopelma marxi (Simon, 1891)

Aphonopelma moderatum (Chamberlin & Ivie, 1939)

Aphonopelma moellendorfi Hamilton, sp. n.

Aphonopelma mojave Prentice, 1997

Aphonopelma paloma Prentice, 1993

Aphonopelma parvum Hamilton, Hendrixson & Bond, sp. n.

Aphonopelma peloncillo Hamilton, Hendrixson & Bond, sp. n.

Aphonopelma phasmus Chamberlin, 1940

Aphonopelma prenticei Hamilton, Hendrixson & Bond, sp. n.

Aphonopelma saguaro Hamilton, sp. n.

Aphonopelma steindachneri (Ausserer, 1875)

Aphonopelma superstitionense Hamilton, Hendrixson & Bond, sp. n.

Aphonopelma vorhiesi (Chamberlin & Ivie, 1939)

Aphonopelma xwalxwal Hamilton, sp. n.

Key to the male Aphonopelma of the United States

Arizona A1
California B1
Colorado, Kansas, Oklahoma, Missouri, Arkansas, and Louisiana C1
Nevada and Utah D1
New Mexico E1
Texas F1

A1. Key to the male Aphonopelma of Arizona

1 Stout setae on the prolateral surface of coxa I; distribution restricted to southern Greenlee County Aphonopelma hentzi
No stout setae on the prolateral surface of coxa I; widespread distribution 2
2 Small species (Cl ≤ 9 mm); generally found in desert, grassland, and/or chaparral habitats 3
Small to medium-sized species (Cl 6–12 mm); generally distributed throughout mid- to high-elevation forested habitats; possessing a black carapace – never blonde, tan, or brown 5 1
Medium to large-sized species (Cl ≥ 9 mm) found in a variety of habitats (but seldom found in higher-elevation forested habitats); carapace blonde, tan, brown, or black 6
3 Possessing a swollen or slightly swollen femur III 4 1
Lacks a swollen femur III; distributed east of Tucson in Cochise, Graham, and Greenlee Counties Aphonopelma parvum
4 Distributed across the Mojave Desert and adjacent sections of the Sonoran Desert in western Arizona Aphonopelma prenticei
Distributed across lower-elevation sections of the Sonoran Desert in southern Arizona Aphonopelma paloma
Distributed across mid- to high-elevation chaparral and forested habitats north of the Phoenix Metropolitan Area Aphonopelma mareki
Distributed at the bottom of Grand Canyon in the vicinity of Phantom Ranch Aphonopelma phasmus
Distributed across the southern foothills and canyons of the Santa Catalina and Rincon Mountains east of Tucson Aphonopelma saguaro
Distributed east of the Phoenix Metropolitan Area in the foothills and canyons of the Superstition Mountains Aphonopelma superstitionense
5 Distributed north and east of the Phoenix Metropolitan Area across the Colorado Plateau and on isolated mountains (e.g., Four Peaks, Mount Ord) Aphonopelma marxi
Distributed across the Santa Catalina Mountains Aphonopelma catalina
Distributed across the Huachuca, Pajarito, Patagonia, and Santa Rita Mountains Aphonopelma madera
Distributed across the Chiricahua Mountains Aphonopelma chiricahua
6 Carapace blonde, tan, or brown 7
Carapace black 8
7 Distributed across the Colorado Plateau north of the Colorado River Aphonopelma iodius
Widespread throughout Arizona but never north of the Colorado River Aphonopelma chalcodes
8 PT1/M1 ≤ 0.68 Aphonopelma gabeli
PT1/M1 ≥ 0.74 Aphonopelma peloncillo and Aphonopelma vorhiesi 2

B1. Key to the male Aphonopelma of California

1 Metatarsus IV scopulation ≥ 60% 2
Metatarsus IV scopulation ≤ 60% 4
2 Possessing a black carapace; distributed across the plains and foothills of the western Sierra Nevada Mountains Aphonopelma johnnycashi
Possessing a tan or brown carapace 3 1
3 Distributed west of the Mojave Desert, from the western part of the Transverse Range and down the southern California coast, along the Peninsular Ranges Aphonopelma eutylenum
Distributed from the Bay Area south along the Coast Ranges, west of the Central Valley, across the Transverse Range and into the Mojave Desert Aphonopelma iodius
4 Possessing stout setae on the medial surface of the sternum 5
Lacks stout setae on the medial surface of the sternum 6
5 F4/T4 ≤ 1.05; distributed across the mountains and foothills west of the Coachella Valley and south to the Borrego Springs area; fall breeding period Aphonopelma xwalxwal
F4/T4 ≥1.07; distributed in and around Joshua Tree National Park; summer breeding period Aphonopelma joshua
6 Medium to large-sized species (Cl ≥ 9 mm); distributed across southwestern California but not within the Mojave Desert Aphonopelma steindachneri
Small species (Cl ≤ 9mm); distributed across the Mojave Desert 7
7 Femur III swollen or slightly swollen 8
Femur III not swollen 9 1
8 A1/F3 ≥ 0.58; distributed across the Panamint Range and eastern San Bernardino County Aphonopelma prenticei
A1/F3 ≤0.56; narrowly distributed across the Amargosa Range in southeastern Inyo County Aphonopelma atomicum
9 Distributed across the northwestern Mojave Desert in eastern Kern and northwestern San Bernardino Counties Aphonopelma mojave
Distributed throughout the southern Mojave Desert along the foothills on the northern side of the San Gabriel and San Bernardino Mountains, east to the Coxcomb Mountains and south into Joshua Tree National Park, north to Kramer Junction and Barstow Aphonopelma icenoglei

C1. Key to the male Aphonopelma of Colorado, Kansas, Oklahoma, Missouri, Arkansas, and Louisiana

1 Carapace tan, brown, or copper; possessing stout setae on the prolateral surface of coxa I; distributed east of the Rocky Mountains across Colorado, Kansas, Oklahoma, Missouri, Arkansas, and Louisiana Aphonopelma hentzi
Carapace black; lacks stout setae on the prolateral surface of coxa I; distributed west of the Rocky Mountains across southwestern Colorado Aphonopelma marxi

D1. Key to the male Aphonopelma of Nevada and Utah

1 Metatarsus IV scopulation ≥ 60%; widespread distribution Aphonopelma iodius
Metatarsus IV scopulation ≤ 60% 2
2 Distributed across the Colorado Plateau in southeastern Utah Aphonopelma marxi
Distributed across the Mojave Desert in southwestern Utah and/or southern Nevada 3 1
3 A1/F3 ≥ 0.58; widespread across the northeastern Mojave Desert Aphonopelma prenticei
A1/F3 ≤ 0.58; distribution restricted to the Amargosa Valley and Nevada Test Site near Mercury, NV Aphonopelma atomicum

E1. Key to the male Aphonopelma of New Mexico

1 Medium to large-sized species (Cl ≥ 9 mm) 2
Small species (Cl ≤ 9 mm) distributed across the extreme southwestern part of the state (Hidalgo County) Aphonopelma parvum
2 Possessing stout setae on the prolateral surface of coxa I; possessing a tan, brown, or copper carapace Aphonopelma hentzi
Lacks stout setae on the prolateral surface of coxa I; possessing a black carapace 3
3 Distributed across the northern half of the state and western part of the Rockies, inhabiting high-elevation pine forest and sagebrush steppe Aphonopelma marxi
Distributed across the southern half of the state 4
4 PT1/M1 ≤ 0.68 Aphonopelma gabeli
PT1/M1 ≥ 0.74 5 3
5 Distribution restricted to the southeastern Peloncillo Mountains; males active mostly during mid-summer Aphonopelma peloncillo
Distribution widespread across southern New Mexico; males active mostly during late summer and early fall Aphonopelma vorhiesi

F1. Key to the male Aphonopelma of Texas

1 Possessing a short and stout embolus; distributed across South Texas Aphonopelma anax
Possessing a long, thin, and tapering embolus; widespread 2
2 Possessing a tan, brown, or copper carapace 3
Possessing a black carapace 4
3 Metatarsus III scopulation ≤ 63%; stout setae on the prolateral surface of coxa I present along dorsal, posterior, and ventral margins, but lacking from the center and anterior margin Aphonopelma armada
Metatarsus III scopulation ≥ 69%; stout setae abundant across the prolateral surface of coxa I Aphonopelma hentzi
4 M1/M4 ≤ 0.74 Aphonopelma gabeli
M1/M4 ≥ 0.75 Aphonopelma moderatum and Aphonopelma moellendorfi 4

1 Species that key here are morphologically indistinguishable for the most part but can be identified based on their localities and molecular data.

2 Mature males of these two species are morphologically indistinguishable and cannot be identified using morphological criteria when they co-occur in southeastern Cochise County, but can be differentiated using molecular data. Aphonopelma vorhiesi is likely the correct determination if the specimen originates from Graham, Pima, Pinal, Santa Cruz, or western Cochise Counties.

3 Species that key here are morphologically indistinguishable for the most part but can be identified based on their localities, the timing of their breeding periods, and molecular data.

4 Mature males of these two species are morphologically indistinguishable and cannot be identified using morphological criteria when they co-occur, but can be differentiated using molecular data. Aphonopelma moderatum reaches its northernmost distribution in Val Verde County but is largely distributed to the south along the Rio Grande Valley. Aphonopelma moellendorfi reaches its easternmost distribution in Val Verde County with other specimens having been located in extreme West Texas. When these two species come into syntopy (southeastern Val Verde County), males can easily be distinguished prior to their ultimate molt due to their phenotypic differences: A. moderatum possess distinct alternating black and tan/orange bands on its legs whereas A. moellendorfi is more uniformly brown.

Key to the female Aphonopelma of the United States

Arizona A2
California B2
Colorado, Kansas, Oklahoma, Missouri, Arkansas, and Louisiana C2
Nevada and Utah D2
New Mexico E2
Texas F2

A2. Key to the female Aphonopelma of Arizona 1

1 Possessing stout setae on the prolateral surface of coxa I; distribution restricted to southern Greenlee County Aphonopelma hentzi
Lacks stout setae on the prolateral surface of coxa I; widespread 2
2 Small species (Cl ≤ 9 mm); generally found in desert, grassland, and/or chaparral habitats 3 2
Small to large-sized species (Cl 7.5–16.5 mm); generally distributed throughout mid- to high-elevation forested habitats; possessing black or gray carapace – never blonde, tan, or brown 4 2
Medium to large-sized species (Cl ≥ 9 mm) found in a variety of habitats (but seldom found in higher-elevation forested habitats); carapace blonde, tan, brown, black, or gray 6
3 Distributed east of Tucson in Cochise, Graham, and Greenlee Counties Aphonopelma parvum
Distributed across the Mojave Desert and adjacent sections of the Sonoran Desert in western Arizona Aphonopelma prenticei
Distributed across lower-elevation sections of the Sonoran Desert in southern Arizona Aphonopelma paloma
Distributed across mid- to high-elevation chaparral and forested habitats north of the Phoenix Metropolitan Area Aphonopelma mareki
Distributed across the southern foothills and canyons of the Santa Catalina and Rincon Mountains east of Tucson Aphonopelma saguaro
Distributed east of the Phoenix Metropolitan Area in the foothills and canyons of the Superstition Mountains Aphonopelma superstitionense
4 T1/T4 ≥ 1.06; distributed north and east of the Phoenix Metropolitan Area across the Colorado Plateau and on isolated mountains (e.g., Four Peaks, Mount Ord) Aphonopelma marxi
T1/T4 ≤1.02; distributed across the Madrean Sky Islands in southeastern Arizona 5
5 Distributed across the Santa Catalina Mountains Aphonopelma catalina
Distributed across the Huachuca, Pajarito, Patagonia, and Santa Rita Mountains Aphonopelma madera
Distributed across the Chiricahua Mountains Aphonopelma chiricahua
6 Metatarsus IV scopulation ≥ 55% 7
Metatarsus IV scopulation ≤ 55% 8
7 Distributed across the Colorado Plateau north of the Colorado River Aphonopelma iodius
Widespread throughout Arizona, but never north of the Colorado River Aphonopelma chalcodes
8 Possessing spermathecae with capitate bulbs Aphonopelma peloncillo and Aphonopelma vorhiesi 3
Possessing short, wide, and slightly rounded spermathecae without capitate bulbs Aphonopelma gabeli

B2. Key to the female Aphonopelma of California 4

1 Metatarsus IV scopulation ≥ 55% 2 2
Metatarsus IV scopulation ≤ 55% 3
2 Distributed across the plains and foothills of the western Sierra Nevada Mountains Aphonopelma johnnycashi
Distributed west of the Mojave Desert, from the western part of the Transverse Range and down the southern California coast, along the Peninsular Ranges Aphonopelma eutylenum
Distributed from the Bay Area south along the Coast Ranges, west of the Central Valley, across the Transverse Range and into the Mojave Desert Aphonopelma iodius
3 Large species (Cl ≥ 9 mm); distributed across southwestern California but not within the Mojave Desert Aphonopelma steindachneri
Small species (Cl ≤ 9 mm); distributed across the Mojave and portions of the Sonoran (Colorado) Deserts 4 2
4 Distributed across the northwestern Mojave Desert in eastern Kern and northwestern San Bernardino Counties Aphonopelma mojave
Distributed in and around Joshua Tree National Park; F1/M4 ≤ 1.04; L1/Cl ≤ 2.87 Aphonopelma joshua 5
Distributed across the Panamint Range and eastern San Bernardino County Aphonopelma prenticei
Distributed across the Amargosa Range in southeastern Inyo County Aphonopelma atomicum
Distributed throughout the southern Mojave Desert along the foothills on the northern side of the San Gabriel and San Bernardino Mountains, east to the Coxcomb Mountains and south into Joshua Tree National Park, north to Kramer Junction and Barstow; F1/M4 ≥ 1.07; L1/Cl ≥ 2.92 Aphonopelma icenoglei 5

C2. Key to the female Aphonopelma of Colorado, Kansas, Oklahoma, Missouri, Arkansas, and Louisiana

1 Carapace tan or brown; possessing stout setae on the prolateral surface of coxa I; distributed east of the Rocky Mountains across Colorado, Kansas, Oklahoma, Missouri, Arkansas, and Louisiana Aphonopelma hentzi
Carapace black; lacks stout setae on the prolateral surface of coxa I; distributed west of the Rocky Mountains across southwestern Colorado Aphonopelma marxi

D2. Key to the female Aphonopelma of Nevada and Utah

1 Metatarsus IV scopulation ≥ 60%; widespread distribution Aphonopelma iodius
Metatarsus IV scopulation ≤ 50% 2
2 Distributed across the Colorado Plateau in southeastern Utah Aphonopelma marxi
Distributed across the Mojave Desert in southwestern Utah and/or southern Nevada 3 2
3 Cl/M4 ≥ 1.31; widespread across the northeastern Mojave Desert Aphonopelma prenticei
Cl/M4 ≤ 1.28; distribution restricted to the Amargosa Valley and Nevada Test Site near Mercury, NV Aphonopelma atomicum

E2. Key to the female Aphonopelma of New Mexico

1 Possessing spermathecae with capitate bulbs 2
Possessing short, wide, and slightly rounded spermathecae without capitate bulbs; anterior margin of carapace broad, associated with robust chelicerae Aphonopelma gabeli
2 Possessing stout setae on the prolateral surface of coxa I; possessing a tan or brown carapace Aphonopelma hentzi
Lacks stout setae on the prolateral surface of coxa I; possessing a black or gray carapace 3
3 Distributed across the northern half of the state and western part of the Rockies, inhabiting high-elevation pine forest and sagebrush steppe Aphonopelma marxi
Distributed across the southern half of the state 4
4 Medium to large-sized species (Cl ≥ 10 mm) 5 2
Small species (Cl ≤ 9 mm); distributed in the extreme southwestern part of the state (Hidalgo County) Aphonopelma parvum
5 Distribution restricted to the southeastern Peloncillo Mountains Aphonopelma peloncillo
Distribution widespread across southern New Mexico Aphonopelma vorhiesi

F2. Key to the female Aphonopelma of Texas 6

1 Possessing spermathecae with capitate bulbs 2
Possessing short, wide, and slightly rounded spermathecae without capitate bulbs 4
2 Possessing stout setae on the prolateral surface of coxa I; all leg segments uniformly colored brown or black 3
Lacks stout setae on the prolateral surface of coxa I; legs distinctly colored with the femora and tibiae generally orange or tan and the patellae, metatarsi, and tarsi dark brown to black (if the leg segments are uniformly colored, they will be orange or tan, never brown or black) Aphonopelma moderatum
3 Stout setae on the prolateral surface of coxa I present along dorsal, posterior, and ventral margins, but lacking from the center and anterior margin; metatarsi I, II, and III distinctly flared; species with overall shiny, lustrous appearance Aphonopelma armada
Stout setae abundant across the prolateral surface of coxa I; metatarsi not distinctly flared; species with overall hirsute appearance Aphonopelma hentzi
4 Distributed across South Texas; possessing a tan or brown carapace with a large robust appearance Aphonopelma anax
Distributed across the Chihuahuan Desert and southwestern High Plains in West Texas; possessing a brownish-gray carapace; anterior margin of carapace broad, associated with robust chelicerae Aphonopelma gabeli

1 Adult females of Aphonopelma phasmus remain unknown.

2 Species that key here are morphologically indistinguishable for the most part but can be identified based on their localities and molecular data.

3 Mature females of these two species can be morphologically indistinguishable when they co-occur in southeastern Cochise County. Aphonopelma vorhiesi is likely the correct determination if the specimen originates from Graham, Pima, Pinal, Santa Cruz, or western Cochise Counties.

4 Adult females of Aphonopelma xwalxwal remain unknown.

5Aphonopelma joshua and A. icenoglei are syntopic at various locations in and around Joshua Tree National Park. Females of A. joshua generally can be diagnosed by possessing tarsi IV divided by setae (Prentice 1997) but this characteristic can be difficult to assess. Molecular data should be used to confirm the identity of specimens from this area.

6 Adult females of Aphonopelma moellendorfi remain unknown.

Aphonopelma anax (Chamberlin, 1940)

Figures 9, 10, 11, 12, 13, 14, Suppl. material 4

Dugesiella anax Chamberlin, 1940: 34; male holotype and female allotype from Kingsville, Kleberg Co., Texas, 27.515869 -97.8561095, elev. 58ft., no collecting date, coll. Prof. J.C. Cross; 3 female paratypes from Harlingen, Cameron Co., Texas, 26.190631 -97.6961035, elev. 40ft., 1939, coll. Bryce Brown; deposited in AMNH. [examined]

Rhechostica anax Raven, 1985: 149.

Aphonopelma anax Smith, 1995: 71.

Aphonopelma breenei Smith, 1995: 78; female holotype from Harlingen, Cameron Co., Texas, 26.190631 -97.6961035, elev. 40ft., 1939, coll. Bryce Brown; deposited in AMNH. [examined] syn. n.

Diagnosis

Aphonopelma anax (Fig. 9) is a member of the Hentzi species group and can be identified by a combination of morphological, molecular, and geographic characteristics. Nuclear DNA identifies A. anax as a phylogenetically distinct monophyletic lineage (Fig. 8), supported as the sister lineage to A. hentzi, and closely related to A. armada. Male A. anax can be distinguished from geographically proximate species by their unique palpal bulbs - stout and wide with distinct keels on the embolus (Fig. 10). Females can be distinguished from geographically proximate species by their unique spermathecae - short, wide, slightly rounded, without capitate bulbs (Figs 1213). Significant measurements that distinguish male A. anax from its closely related phylogenetic and syntopic species are Cl, M3, and F4. Male A. anax can be distinguished by possessing a larger F4/M4 (≥0.94; 0.94–1.04) than A. armada (≤0.92; 0.86–0.92); a larger Cl/M1 (≥1.36; 1.36–1.63) than moderatum (≤1.30; 1.13–1.30) and A. moellendorfi sp. n. (≤1.31; 1.10–1.31); and a larger F1/M3 (≥1.28; 1.28–1.43) than A. gabeli (≤1.24; 1.18–1.24). There are no significant measurements that separate male A. anax from A. hentzi. Significant measurements that distinguish female A. anax from its closely related phylogenetic and syntopic species are P1 and T3. Female A. anax can be distinguished by possessing a larger P1/T3 (≥13.88; 13.88–19.15) than A. armada (≤13.84; 9.93–13.84) and A. moderatum (≤13.12; 8.14–13.12). There are no significant measurements that separate female A. anax from A. gabeli or A. hentzi. Females of A. moellendorfi are unknown and cannot be compared.

Figure 9.

Aphonopelma anax (Chamberlin, 1940) specimens, live photographs. Female (L) - APH_0524; Male (R) - APH_3122.

Figure 10.

Aphonopelma anax (Chamberlin, 1940). A–I male specimen, APH_0924 A dorsal view of carapace, scale bar = 5mm B prolateral view of coxa I C dorsal view of femur III D ventral view of metatarsus III, scale bar = 4mm E ventral view of metatarsus IV, scale bar = 4mm F prolateral view of L pedipalp and palpal tibia, scale bar = 3mm G dorsal view of palpal bulb H retrolateral view of palpal bulb, scale bar = 1mm I prolateral view of tibia I (mating clasper), scale bar = 6mm.

Figure 11.

Aphonopelma anax (Chamberlin, 1940). A–E female specimen, APH_0857 A dorsal view of carapace, scale bar = 8mm B prolateral view of coxa I C ventral view of metatarsus III, scale bar = 4mm D ventral view of metatarsus IV, scale bar = 5.5mm E prolateral view of L pedipalp and palpal tibia.

Figure 12.

Aphonopelma anax (Chamberlin, 1940). A–H cleared spermathecae A anax allotype B breenei holotype C harlingenum paratype D APH_0056 E APH_0529 F APH_0857 G APH_0858 H APH_0859.

Figure 13.

Aphonopelma anax (Chamberlin, 1940). A–E cleared spermathecae A APH_0871 B APH_0899 C APH_0902 D APH_1278 E APH_1280.

Descriptions

Male and female originally described by Chamberlin (1940).

Redescription of male exemplar

(APH_0924; Fig. 10). Specimen preparation and condition: Specimen collected live crossing road, preserved in 80% ethanol; deposited in AUMNH; original coloration faded due to preservation. Left legs I, III, IV, and left pedipalp removed for measurements and photographs; stored in vial with specimen. Right leg III removed for DNA and stored at -80°C in the AUMNH (Auburn, AL). General coloration: Generally black or faded to brown. Cephalothorax: Carapace 19.10 mm long, 18.28 mm wide; densely clothed with brown/golden iridescent pubescence appressed to surface; fringe covered in long setae not closely appressed to surface; foveal groove medium deep and straight to slightly procurved; pars cephalica region rises gradually from foveal groove, gently arching anteriorly toward ocular area; AER slightly procurved, PER recurved; normal sized chelicerae; clypeus extends forward on a very slight curve; LBl 2.96, LBw 2.94; sternum hirsute, clothed with short length black, densely packed setae. Abdomen: Densely clothed in short black pubescence with numerous longer, lighter setae interspersed (generally red or orange in situ); dense dorsal patch of black Type I urticating bristles (Cooke et al. 1972). Legs: Thick and hirsute; densely clothed in short black/brown pubescence. Metatarsus I slightly curved. F1 18.77; F1w 4.74; P1 8.10; T1 14.69; M1 13.41; A1 9.23; F3 15.19; F3w 4.61; P3 6.75; T3 12.07; M3 13.77; A3 9.42; F4 18.58; F4w 4.8; P4 8.16; T4 14.50; M4 19.51; A4 11.15; femur III is normal - not noticeably swollen or wider than other legs. All tarsi fully scopulate. Extent of metatarsal scopulation: leg III (SC3) = 68.9%; leg IV (SC4) = 43.4%. Two ventral spinose setae on metatarsus III; six ventral spinose setae on metatarsus IV. Coxa I: Prolateral surface a mix of fine, hair-like and medium tapered setae. The interior face of the retrolateral branch of the tibial apophyses possesses a very large megaspine that projects anteriorly. Pedipalps: Hirsute; densely clothed in the same setal color as the other legs, with numerous longer ventral setae; one spinose seta at the apical, prolateral femur and three spinose setae on the prolateral tibia; PTl 9.45, PTw 2.90. When extended, embolus tapers and rapidly curves up and to the retrolateral side near apex; embolus wide and thick, with smooth dorsal and ventral keels.

Variation (11). Cl 14.371–21.97 (17.885±0.7), Cw 13.32–19.84 (16.557±0.62), LBl 1.96–2.96 (2.341±0.09), LBw 2.16–3.02 (2.711±0.08), F1 14.1–19.3 (16.839±0.55), F1w 3.52–5.1 (4.392±0.15), P1 6.12–8.1 (7.105±0.22), T1 11.32–15.71 (13.314±0.39), M1 9.69–13.43 (11.878±0.4), A1 6.9–9.6 (8.499±0.28), L1 length 48.47–65.97 (57.635±1.73), F3 11.39–16.14 (13.868±0.48), F3w 3.41–5.03 (4.132±0.16), P3 4.86–7.65 (6.035±0.27), T3 7.91–12.16 (10.428±0.43), M3 9.89–14.89 (12.572±0.49), A3 6.93–9.45 (8.291±0.27), L3 length 41.32–59.2 (51.194±1.82), F4 13.52–19.31 (16.56±0.58), F4w 3.22–4.83 (4.094±0.17), P4 5.5–8.16 (6.647±0.26), T4 11.15–15.56 (13.676±0.41), M4 13.66–19.69 (17.13±0.58), A4 8.1–11.15 (9.504±0.3), L4 length 52.79–72.08 (64.171±1.92), PTl 7.187–10.136 (8.709±0.27), PTw 2.246–3.32 (2.882±0.09), SC3 ratio 0.607–0.805 (0.713±0.02), SC4 ratio 0.351–0.524 (0.44±0.02), Coxa 1 setae = thick tapered, F3 condition = normal/slightly swollen.

Redescription of female exemplar

(APH_0857; Figs 1113). Specimen preparation and condition: Specimen collected live from burrow, preserved in 80% ethanol; deposited in AUMNH; original coloration faded due to preservation. Left legs I, III, IV, and pedipalp removed for photographs and measurements; stored in vial with specimen. Right leg III removed for DNA and stored at -80°C in the AUMNH (Auburn, AL). Genital plate with spermathecae removed and cleared, stored in vial with specimen. General coloration: Faded black and light brown, medium length, dense setae cover body. Cephalothorax: Carapace 21.37 mm long, 18.26 mm wide; densely clothed with light brown pubescence closely appressed to surface; fringe densely covered in long setae; foveal groove medium deep and straight; pars cephalica region rises from thoracic furrow more steeply than male, gently arching anteriorly toward ocular area; AER procurved, PER strongly recurved; clypeus extends forward on a slight curve; LBl 2.67, LBw 3.35; sternum hirsute, clothed with light black/dark brown, medium length dense setae. Abdomen: Densely clothed dorsally in short black setae with numerous longer, lighter setae interspersed (generally red or orange in situ); dense dorsal patch of black Type I urticating bristles (Cooke et al. 1972); ventral side with short, dense black setae. Spermathecae: Paired and separate, possessing dual bulges with the smaller near the interior, with wide bases that are not fused. Legs: Thick and hirsute; densely clothed in a mix of short black/brown pubescence setae. Coxa I: Prolateral surface a mix of fine, hair-like and thick tapered setae. F1 16.02; F1w 4.88; P1 7.53; T1 12.06; M1 9.18; A1 7.54; F3 12.72; F3w 4.56; P3 6.76; T3 9.29; M3 10.09; A3 7.54; F4 16.53; F4w 4.73; P4 7.08; T4 13.31; M4 13.26; A4 8.79. All tarsi fully scopulate. Extent of metatarsal scopulation: leg III (SC3) = 64.4%; leg IV (SC4) = 47.3%. Two ventral spinose setae on metatarsus III; seven ventral spinose setae on metatarsus IV. Pedipalps: Densely clothed in the same setal color as the other legs; one spinose seta on the apical, prolateral femur and five spinose setae on the prolateral tibia.

Variation (13). Cl 16.06–23.8 (20.27±0.79), Cw 14.9–21.73 (17.951±0.62), LBl 2.45–3.79 (2.878±0.1), LBw 2.71–4.3 (3.478±0.15), F1 13.12–18.14 (15.181±0.45), F1w 3.87–5.8 (4.945±0.18), P1 5.61–8.59 (7.196±0.26), T1 9.76–13.56 (11.616±0.35), M1 7.0–10.98 (8.911±0.31), A1 5.73–8.34 (7.138±0.19), L1 length 42.75–59.3 (50.042±1.47), F3 10.94–14.63 (12.375±0.37), F3w 3.36–5.13 (4.268±0.17), P3 5.1–7.72 (6.179±0.23), T3 7.45–10.56 (8.816±0.28), M3 8.27–12.05 (9.918±0.34), A3 6.52–8.84 (7.398±0.18), L3 length 38.67–53.7 (44.687±1.26), F4 13.27–18.66 (15.654±0.49), F4w 3.61–5.31 (4.514±0.17), P4 5.34–8.4 (6.55±0.28), T4 9.72–13.98 (12.038±0.35), M4 10.37–16.15 (13.621±0.46), A4 7.01–9.83 (8.373±0.25), L4 length 45.86–66.63 (55.723±1.74), SC3 ratio 0.644–0.763 (0.706±0.01), SC4 ratio 0.368–0.474 (0.433±0.01), Coxa 1 setae = thick tapered. Spermathecae variation can be seen in Figures 1213.

Material examined

United States: Texas: Bexar: Hollywood Park, 220 Mecca, 29.59413 -98.47946 2, 934ft., [APH_0033, 2/6/2006, 1♂, Connor Shannon, Ryan Tubbesing, AUMNH]; Cameron: 2100 W. San Marcelo Blvd #158, Brownsville, 25.95835 -97.500489 2, 21ft., [APH_0523, 20/5/2009, 1♂, Lilia Perez, AUMNH]; Brownsville, 25.901747 -97.497484 5, 26ft., [APH_2045, 4/1963, 1♀, 1♂, Ted Beimler, AMNH]; Brownsville, field NE Coffeeport Rd, 25.948055 -97.480915 1, 19ft., [APH_0459-0462, 9/4/09, 1♀, 3 juv, Brent E. Hendrixson, AUMNH]; Harlingen, 26.190631 -97.696103 5, 39ft., [APH_2043, 15/11/1939, 1♀, B. Brown, AMNH]; [APH_2044, 1939, 1♀, Bryce Brown, AMNH]; Harlingen, Dixieland Park, 26.16825 -97.72063 1, 43ft., [APH_1273-1275, 12/5/11, 3 juv, Brent E. Hendrixson, Kate Hall, Austin Deskewies, Alexis Guice, AUMNH]; Harlingen, McKelvey Park, 26.180254 -97.679291 1, 35ft., [APH_0463, 9/4/09, 1 juv, Brent E. Hendrixson, AUMNH]; [APH_1271-1272, 11/5/11, 2 juv, Brent E. Hendrixson, Kate Hall, Austin Deskewies, Alexis Guice, AUMNH]; Harlingen, S of town, 26.145617 -97.661717 5, 41ft., [APH_0924, 2006, 1♂, Dave Moellendorf, AUMNH]; Laguna Vista, 26.100864 -97.290234 5, 12ft., [AUMS_2355, 4/1998, 1♂, R.G. Breene, AUMNH]; [AUMS_2583, 4/1998, 1♂, R.G. Breene, AUMNH]; [AUMS_2609, 1998, 1♂, R.G. Breene, AUMNH]; [AUMS_2613, unknown, 1♀, R.G. Breene, AUMNH]; [AUMS_2690, 4/1998, 2♂, R.G. Breene, AUMNH]; [AUMS_2695, 4/1998, 1♀, R.G. Breene, AUMNH]; [AUMS_3268-3269, 3/1998, 2♂, R.G. Breene, AUMNH]; [AUMS_3271, 4/1998, 1♂, R.G. Breene, AUMNH]; [AUMS_3274, 4/1998, 1♂, R.G. Breene, AUMNH]; [AUMS_3315, 4/1998, 1♂, R.G. Breene, AUMNH]; Laguna Vista, Roloff Park, 26.10243 -97.291 1, 3ft., [APH_0455-0458, 9/4/09, 2♀, 2 juv, Brent E. Hendrixson, AUMNH]; Los Fresnos, 26.071744 -97.476373 5, 23ft., [AUMS_2688, 1999, 1♀, R.G. Breene, AUMNH]; South Padre Island, 26.076567 -97.16315 5, 4ft., [APH_0858, 2006, 1♀, Dave Moellendorf, AUMNH]; DeWitt: between Cuero and Westhoff on Hwy 87, 29.1221 -97.410817 1, 303ft., [APH_0859-0861, 9/2008, 3♀, Chris A. Hamilton, AUMNH]; [APH_0947, 9/2008, 1♂, Chris A. Hamilton, AUMNH]; Dimmit: 6.7 miles E Maverick County Line on US-277, 28.625443 -100.004038 1, 659ft., [APH_0473, 11/4/2009, 1 juv, Brent E. Hendrixson, AUMNH]; Chaparral Wildlife Management Area, Blocker Pond, 28.3125 -99.40694 1, 570ft., [APH_0025, 12/3/2002, 1♂, Brent E. Hendrixson, WTAMU Herp Class, AUMNH]; 3 miles NW Catarina on US-83, 28.37472 -99.64806 1, 560ft., [APH_0042, 12/3/2000, 1 juv, Brent E. Hendrixson, AUMNH]; Duval: 4.4 miles E Hwy-339 on FM-2295, 27.58835 -98.33743 1, 340ft., [APH_1270, 11/5/11, 1 juv, Brent E. Hendrixson, Kate Hall, Austin Deskewies, Alexis Guice, AUMNH]; 5.3 miles SE Webb County Line on Hwy-44, 27.907752 -98.723635 1, 497ft., [APH_0529, 3/6/09, 1♀, Brent E. Hendrixson, Courtney Dugas, Sloan Click, AUMNH]; S of San Diego, TX on Hwy-359, 27.733177 -98.262366 1, 339ft., [APH_0584, 14/6/2009, 1 juv, Brent E. Hendrixson, Courtney Dugas, Sloan Click, AUMNH]; Fayette: La Grange, 29.914417 -96.866267 1, 330ft., [APH_0804-0806, 5/2008, 3♀, Chris A. Hamilton, AUMNH]; [APH_0808, 5/2008, 1♀, Chris A. Hamilton, AUMNH]; [APH_0809, 5/2008, 1♀, Chris A. Hamilton, AUMNH]; [APH_0898, 5/2008, 1♂, Chris A. Hamilton, AUMNH]; La Grange, S side of river, 29.874417 -96.850383 1, 345ft., [APH_0811, 7/2008, 1♀, Chris A. Hamilton, AUMNH]; [APH_0810, 5/2008, 1♀, Chris A. Hamilton, AUMNH]; [APH_0899, 5/2008, 1♂, Chris A. Hamilton, AUMNH]; [APH_0901, 5/2008, 1♂, Chris A. Hamilton, AUMNH]; La Grange, on FM155, 29.859333 -96.848033 1, 348ft., [APH_0900, 5/2008, 1♂, Chris A. Hamilton, AUMNH]; La Grange, rest stop on Hwy 77, 29.843383 -96.9091 1, 377ft., [APH_0902-0903, 5/2008, 1♀, 1♂, Chris A. Hamilton, AUMNH]; Gonzales: 6.1 miles SE of Gonzales, 29.497194 -97.380048 5, 328ft., [APH_2658, 18/6/1956, 1♂, W. McAlister, AMNH]; Harris: Houston, 29.760193 -95.36939 6, 43ft., [APH_2650, 6/1959, 1♂, Mrs. Emilie Steude, AMNH]; Jim Hogg: 0.9 miles E Zapata County Line on FM-2687, 26.939665 -98.941918 1, 526ft., [APH_1131, 15/3/2010, 1 juv, Brent E. Hendrixson, Gerri Wilson, Thomas Martin, AUMNH]; 2.0 miles N Starr County Line on FM-649, 26.81392 -98.855998 1, 625ft., [APH_1130, 15/3/2010, 1 juv, Brent E. Hendrixson, Gerri Wilson, Thomas Martin, AUMNH]; Karnes: Karnes City, 28.885483 -97.902683 1, 411ft., [APH_0864-0866, 9/2008, 2♀, 1 juv, Chris A. Hamilton, AUMNH]; [APH_0952, 9/2008, 1♂, Chris A. Hamilton, AUMNH]; Kleberg: Kingsville, S Brahma Blvd, 27.4806 -97.855767 1, 77ft., [APH_0856-0857, 9/2008, 2♀, Chris A. Hamilton, AUMNH]; Kingsville, 27.515869 -97.856109 5, 56ft., [APH_2046, 27/6/1970, 1♂, Gillaspy, AMNH]; Kingsville, field at high school, along Caesar Ave across street from cemetery, 27.506531 -97.879785 1, 65ft., [APH_0579-0583, 13/6/2009, 4 juv, Brent E. Hendrixson, Courtney Dugas, Sloan Click, AUMNH]; La Salle: Cotulla, 28.436934 -99.235032 1, 437ft., [APH_3129, 2013, 1♂, Roger Birkhead, AUMNH]; Los Angeles, 28.460017 -98.9999 1, 391ft., [APH_0871, 2008, 1♀, Sky Stevens, AUMNH]; Maverick: 4.2 miles SW Zavala County Line on Hwy-57, 28.88947 -100.16539 2, 711ft., [APH_1458, 24/1/2012, 1 juv, Stanley A. Schultz, AUMNH]; 4.6 miles E/SE Eagle Pass (jct US-57) on US-277, 28.69567 -100.39303 1, 833ft., [APH_0055, 17/7/2006, 1♂, Brent E. Hendrixson, AUMNH]; [APH_0058, 17/7/2006, 1♀, Brent E. Hendrixson, AUMNH]; [APH_0084, 17/7/2006, 1♀, Brent E. Hendrixson, AUMNH]; 9.0 miles NE US-57 on FM-481, 28.928198 -100.262798 1, 787ft., [APH_1149-1150, 17/3/2010, 2 juv, Brent E. Hendrixson, Gerri Wilson, Thomas Martin, AUMNH]; McMullen: 1.25 miles N FM-3445 on TX-16, 28.520536 -98.547662 1, 300ft., [APH_1117, 14/3/2010, 1 juv, Brent E. Hendrixson, Gerri Wilson, Thomas Martin, AUMNH]; Nueces: Corpus Christi, Meldo Park, approx. 2 blocks SW Santa Fe St on Brawner Parkway, 27.747437 -97.379802 1, 41ft., [APH_0577-0578, 13/6/2009, 2 juv, Brent E. Hendrixson, Courtney Dugas, Sloan Click, AUMNH]; Starr: San Carlos, Hwy 649 ~ 2 miles N of 2686 junction, 26.720783 -98.8468 1, 411ft., [APH_0889, 18/4/2008, 1♀, Christian Cox, Corey Roelke, AUMNH]; Washington: 2301 Running Valley Lane, Washington, 30.225692 -96.160138 1, 224ft., [APH_3123, 13/6/2013, 1♂, Chuck Matula, AUMNH]; 5 miles west of Carmine, 30.140828 -96.759342 5, 433ft., [APH_2660, 27/6/1960, 2♀, W. McAlister, AMNH]; Brenham, 30.145467 -96.43435 5, 323ft., [APH_0968, 6/2006, 1♂, Dave Moellendorf, AUMNH]; Chappell Hill, 1/4 mile S of intersection of Old Chappell Hill Rd and Pulaski School Rd, 30.1577 -96.288344 1, 282ft., [APH_3121, 20/5/2013, 1♂, Todd Burch, AUMNH]; [APH_3122, 30/5/2013, 1♂, Todd Burch, AUMNH]; Webb: 0.8 miles NW Hwy-44 on US-83, 28.03711 -99.54681 1, 820ft., [APH_1285, 13/5/2011, 1♂, Brent E. Hendrixson, Kate Hall, Austin Deskewies, Alexis Guice, AUMNH]; 1.1 miles SW Duval County Line on US-59, 27.786023 -98.818769 1, 451ft., [APH_0524-0528, 3/6/09, 3♀, 2 juv, Brent E. Hendrixson, Courtney Dugas, Sloan Click, AUMNH]; 3.2 miles W FM-2895 on TX-359, 27.454855 -99.136257 1, 511ft., [APH_1125, 15/3/2010, 1 juv, Brent E. Hendrixson, Gerri Wilson, Thomas Martin, AUMNH]; 3.6 miles NW I-35 on US-83, 27.80051 -99.46291 1, 760ft., [APH_1281-1282, 13/5/2011, 2 juv, Brent E. Hendrixson, Kate Hall, Austin Deskewies, Alexis Guice, AUMNH]; 4.3 miles E Loop-20 on US-59 at Los Tios Creek, 27.56159 -99.39045 1, 470ft., [APH_1280, 13/5/2011, 1 juv, Brent E. Hendrixson, Kate Hall, Austin Deskewies, Alexis Guice, AUMNH]; 5.1 miles SE La Salle County Line on Hwy-44, 28.012456 -99.195109 1, 454ft., [APH_0530-0531, 3/6/2009, 2 juv, Brent E. Hendrixson, Courtney Dugas, Sloan Click, AUMNH]; Wharton: El Campo, 29.19405 -96.2617 1, 90ft., [APH_0800-0803, 7/2008, 4♀, Chris A. Hamilton, AUMNH]; Zapata: 2.0 miles NW FM-2687 on US-83, 26.80362 -99.142 1, 410ft., [APH_1277-1279, 13/5/2011, 1♂, 2 juv, Brent E. Hendrixson, Kate Hall, Austin Deskewies, Alexis Guice, AUMNH]; Mexico: Nuevo Leon: Cañon de la Huasteca, 25.61484 -100.47585 1, 2600ft., [APH_0056, 24/7/2006, 1♀, Brent E. Hendrixson, AUMNH].

Distribution and natural history

In the United States, A. anax is widely distributed throughout South Texas (Fig. 14). We only sampled a single individual from western Nuevo Leon but the species distribution model predicts suitable habitat for this species throughout northeastern Mexico in Coahuila, Nuevo Leon, and Tamaulipas. The vast majority of specimens were collected at elevations between sea level and 300 meters along the Western Gulf Coastal Plain (Fig. 1I) or Southern Texas Plains Level III Ecoregions, but other samples were obtained from the Texas Blackland Prairies, East Central Texas Plains, South Central Plains, and the very southern edge of the Edwards Plateau. The specimen from Nuevo Leon (at Huasteca Canyon near Monterrey) was collected at an elevation of approximately 850 meters. This species has been observed in syntopy with A. moderatum (burrows located fewer than 50 centimeters from each other) at several different locations near the Rio Grande from Eagle Pass (Maverick County) to Rio Grande City (Starr County). Aphonopelma anax may also be found alongside A. armada and A. hentzi. Burrows are typical of that for North American tarantulas (i.e., circular and generally covered by a thin veil of silk) and specimens can be readily collected by pouring a small amount of water into their burrows. In extreme South Texas, A. anax is probably active year-round; however, in more northern populations, spiders likely plug their burrows for short periods of time during the winter. The breeding period appears to be restricted to spring and early summer (April-July); adult males have been observed in large numbers at night along roads following heavy bouts of rain (United States Border Patrol 2009, pers. comm.).

Figure 14.

Aphonopelma anax (Chamberlin, 1940). A distribution of known specimens B predicted distribution; warmer colors (red, orange, yellow) represent areas of high probability of occurrence, cooler colors (blue shades) represent areas of low probability of occurrence.

Conservation status

Aphonopelma anax is very common throughout its distribution. Extensive fieldwork near Edinburg and McAllen (Hidalgo County) suggests that some local populations of A. anax have probably been extirpated due to extensive agriculture in the Lower Rio Grande Valley, but overall the species is fairly abundant throughout South Texas. These spiders appear to thrive in a variety of anthropogenic settings including golf courses, residential lawns, city parks, roadside picnic areas, and mowed highway shoulders. The status of A. anax is likely secure.

Remarks

Aphonopelma anax is one of the largest and most robust Aphonopelma within the United States. This species exhibits size variation within males and females across their distribution, with northern populations generally smaller and the southern populations representing the largest tarantulas in the United States. Other important ratios that distinguish males: A. anax possess a smaller M1/F4 (≤0.75; 0.69-0.75) than A. moderatum (≥0.80; 0.80-0.88) and A. moellendorfi (≥0.81; 0.81-0.88). No other ratios distinguish female A. anax from their syntopic or closely related phylogenetic species. For both males and females, certain morphometrics have potential to be useful though due to the amounts of variation, small number of specimens, and the small differences between species none are claimed to be significant at this time (see Suppl. material 2). During evaluation of traditional PCA morphospace, males of A. anax separate from A. moderatum, A. moellendorfi, and A. gabeli along PC1~2, but do not separate from A. armada or A. hentzi. Females separate from moderatum along PC1~2, but do not separate from A. armada, A. gabeli, A. hentzi. Females of A. moellendorfi are unknown at this time and cannot be compared. Interestingly, A. anax males separate from A. gabeli, A. moderatum, and A. moellendorfi in three-dimensional PCA morphospace (PC1~PC2~PC3), but do not separate from A. armada and A. hentzi. Aphonopelma anax females separate from A. armada and A. moderatum, but do not separate from A. gabeli and A. hentzi. PC1, PC2, and PC3 explain ≥87% of the variation in male analyses and ≥96% of the variation in female analyses.

It is important to note the tremendous amount of variation that can be observed in the shape of the spermathecae from numerous populations of A. anax (Figs 1213). Previous taxonomic work considered this amount of variation to represent differences between species (e.g., Smith 1995 differentiated his new species A. breenei from A. anax on the basis of its spermathecae) (Smith 1995) but our samples demonstrate that the shape of spermathecae is quite variable and is not useful for differentiating these two species. The spermathecae are however useful for distinguishing A. anax from other members of the Hentzi species complex (A. armada and A. hentzi). This information was used to place A. harlingenum (whose type locality is surrounded by A. anax) into synonymy with A. hentzi and not A. anax. It is also important to note that the paratype of A. harlingenum that is found in the same jar as the A. harlingenum holotype is a female of A. anax (see spermathecae in Fig. 12C). Additionally, as seen in the spermathecae, we find that palpal bulb variation is extreme both across and within Aphonopelma species - an unfortunate finding that further enforces our idea that these traditionally used morphological characters are ineffective species delimiters. Unless these characters are extreme autapomorphies (i.e. A. anax spermathcae and palpal bulbs, A. gabeli spermathecae) that can be used in conjunction with other evidence to determine species boundaries in this group of spider, they should not be regarded as taxonomically informative.

Mitochondrial DNA (CO1) identifies A. anax as a polyphyletic group with respect to A. armada and A. hentzi (Fig. 7). A second lineage of A. anax is sister to a lineage of A. hentzi; both of these lineages were previously identified as putative cryptic species (Hamilton et al. 2014). The AE data demonstrate that CO1 is not effective at accurately delimiting species boundaries within this group. Finally, we examined the holotype and freshly collected topotypic material of A. breenei. Our morphological and molecular analyses fail to recognize this species as a separate, independently evolving lineage. As a consequence, we consider A. breenei a junior synonym of A. anax.

Aphonopelma armada (Chamberlin, 1940)

Figures 15, 16, 17, 18, 19

Dugesiella armada Chamberlin, 1940: 32; female holotype from Austin, Travis Co., Texas, 30.267153 -97.7430616, elev. 461ft., ix.1909, coll. A. Petrunkevitch; deposited in AMNH. [examined]

Rhechostica armada Raven, 1985: 149.

Aphonopelma armada Smith, 1995: 71.

Aphonopelma arnoldi Smith, 1995: 74; male holotype from Hwy 82 near Crosbyton, Crosby Co., Texas, 33.660017 -101.2946445, elev. 3063ft., 17.vi.1963, coll. P. Keathley; deposited in Oklahoma State University collection. [not examined] syn. n.

Diagnosis

Aphonopelma armada (Fig. 15) is a member of the Hentzi species group and can be identified by a combination of morphological, molecular, and geographic characteristics. Nuclear and mitochondrial DNA identifies A. armada as a phylogenetically distinct monophyletic lineage (Figs 78), supported as a sister lineage to A. anax and A. hentzi. Female A. armada can be distinguished by their noticeably large urticating hair patch (black spot), a less hirsute and more lustrous appearance, and metatarsi I, II, and III that are distinctly flared and wider than other syntopic species (Fig. 15). Male A. armada are generally smaller and less hirsute than other similar looking syntopic species. Setae on the prolateral face of coxa I have a unique pattern that help distinguish A. armada from all other Aphonopelma in the United States (Figs 1617). Significant measurements that distinguish male A. armada from its closely related phylogenetic and syntopic species are F4, PTl, and extent of metatarsus III scopulation. Male A. armada can be distinguished by possessing a larger PTl/M1 (≥0.73; 0.73–0.83) than A. gabeli (≤0.68; 0.61–0.68), A. moderatum (≤0.69; 0.61–0.69), and A. moellendorfi sp. n. (≤0.69; 0.60–0.69); by possessing a smaller F4/M4 (≤0.92; 0.86–0.92) than A. anax (≥0.94; 0.94–1.04); and smaller L3 scopulation extent (48%–63%) than A. hentzi (69%–86%). Significant measurements that distinguish female A. armada from its closely related phylogenetic and syntopic species are P1, Cl, and extent of metatarsus IV scopulation. Female A. armada can be distinguished by possessing a larger Cl/A3 (≥2.48; 2.48–2.69) than A. moderatum (≤2.46; 2.25–2.46); a larger P1/F4 (≥0.47; 0.47–0.52) than A. gabeli (≤0.46; 0.42–0.46); by possessing a smaller P1/T3 (≤13.84; 9.93–13.84) than A. anax (≥13.88; 13.88–19.15); and a smaller L4 scopulation extent (28%-43%) than A. hentzi (42%-72%, with slight overlap). Females of A. moellendorfi are unknown and cannot be compared.

Figure 15.

Aphonopelma armada (Chamberlin, 1940) specimens, live photographs. Male (L) - APH_1064; Female (R) - APH_3214.

Figure 16.

Aphonopelma armada (Chamberlin, 1940). A–I male specimen, APH_0950 A dorsal view of carapace, scale bar = 6.5mm B prolateral view of coxa I C dorsal view of femur III D ventral view of metatarsus III, scale bar = 4mm E ventral view of metatarsus IV, scale bar = 3mm F prolateral view of L pedipalp and palpal tibia, scale bar = 3mm G dorsal view of palpal bulb H retrolateral view of palpal bulb, scale bar = 1mm I prolateral view of tibia I (mating clasper), scale bar = 2mm.

Figure 17.

Aphonopelma armada (Chamberlin, 1940). A–E female specimen, APH_0848 A dorsal view of carapace, scale bar = 6mm B prolateral view of coxa I C ventral view of metatarsus III, scale bar = 3mm D ventral view of metatarsus IV, scale bar = 3mm E prolateral view of L pedipalp and palpal tibia.

Description

Female originally described by Chamberlin (1940).

Description of male exemplar

(APH_0950; Fig. 16). Specimen preparation and condition: Specimen collected live crossing road, preserved in 80% ethanol; deposited in AUMNH; original coloration faded due to preservation. Left legs I, III, IV, and left pedipalp removed for measurements and photographs; stored in vial with specimen. Right leg III removed for DNA and stored at -80°C in the AUMNH (Auburn, AL). General coloration: Generally black or faded to brown. Cephalothorax: Carapace 14.59 mm long, 13.42 mm wide; Very hirsute, densely clothed with brown/golden iridescent pubescence appressed to surface; fringe covered in long setae not closely appressed to surface; foveal groove medium deep and straight; pars cephalica region rises gradually from foveal groove, gently arching anteriorly toward ocular area; AER slightly procurved, PER slightly recurved; normal sized chelicerae; clypeus extends forward on a slight curve; LBl 2.09, LBw 2.42; sternum hirsute, clothed with medium length black/brown, densely packed setae. Abdomen: Densely clothed in short black/brown pubescence with numerous longer, lighter setae interspersed (generally red or orange in situ); possessing a dense dorsal patch of black Type I urticating bristles (Cooke et al. 1972). Legs: Hirsute, densely clothed in a mix of short black/brown pubescence with longer ventral setae. Metatarsus I slightly curved. F1 15.16; F1w 3.72; P1 6.29; T1 11.1; M1 10.31; A1 7.88; F3 12.04; F3w 3.49; P3 5.26; T3 8.89; M3 11.36; A3 7.68; F4 14.12; F4w 3.58; P4 5.76; T4 13.16; M4 15.28; A4 8.03; femur III is normal - not noticeably swollen or wider than other legs. All tarsi fully scopulate. Extent of metatarsal scopulation: leg III (SC3) = 63.4%; leg IV (SC4) = 37.1%. Three ventral spinose setae on metatarsus III; six ventral spinose setae on metatarsus IV. Coxa I: Prolateral surface a mix of fine, hair-like, very thick tapered, and stout setae with a unique placement around the anterior, prolateral margin. Pedipalps: Hirsute; densely clothed in the same setal color as the other legs, with numerous longer ventral setae; two spinose setae on the prolateral tibia; PTl 8.554, PTw 2.668. When extended, embolus tapers with a gentle curve to the retrolateral side near apex; embolus very slender, no keels; distinct ventral shift from the bulb to the embolus.

Variation (5). Cl 13.19–16.38 (14.614±0.61), Cw 11.37–13.88 (12.912±0.48), LBl 1.79–2.09 (1.956±0.06), LBw 2.04–2.51 (2.28±0.09), F1 13.7–15.67 (14.656±0.4), F1w 3.27–3.97 (3.688±0.13), P1 4.61–6.91 (6.056±0.38), T1 11.1–12.48 (11.724±0.23), M1 9.28–11.36 (10.386±0.39), A1 7.13–8.51 (7.76±0.25), L1 length 46.14–53.64 (50.582±1.41), F3 10.81–12.46 (11.742±0.31), F3w 3.18–3.97 (3.546±0.14), P3 4.99–5.48 (5.236±0.08), T3 8.26–9.35 (8.806±0.22), M3 10.78–12.26 (11.414±0.29), A3 6.68–7.72 (7.25±0.21), L3 length 41.97–46.59 (44.448±1.02), F4 12.85–14.12 (13.4±0.24), F4w 3.17–3.66 (3.474±0.09), P4 4.73–5.96 (5.476±0.22), T4 10.95–13.16 (11.896±0.36), M4 14.18–15.5 (14.95±0.23), A4 7.37–8.77 (8.15±0.24), L4 length 51.3–56.35 (53.872±1.03), PTl 7.053–8.554 (8.061±0.28), PTw 2.372–2.756 (2.578±0.07), SC3 ratio 0.483–0.634 (0.566±0.03), SC4 ratio 0.292–0.371 (0.328±0.01), Coxa I setae = very thick tapered & stout, F3 condition = normal.

Redescription of female exemplar

(APH_0848; Figs 1718). Specimen preparation and condition: Specimen collected live from burrow, preserved in 80% ethanol; deposited in AUMNH; original coloration faded due to preservation. Left legs I, III, IV, and pedipalp removed for photographs and measurements; stored in vial with specimen. Right leg III removed for DNA and stored at -80°C in the AUMNH (Auburn, AL). Genital plate with spermathecae removed and cleared, stored in vial with specimen. General coloration: Faded black and brown, medium length setae cover body. Cephalothorax: Carapace 16.71 mm long, 14.08 mm wide; densely clothed with light brown pubescence closely appressed to surface; fringe densely covered in medium setae; foveal groove medium deep and slightly procurved; pars cephalica region rises from thoracic furrow more steeply than male, gently arching anteriorly toward ocular area; AER slightly procurved, PER recurved; clypeus mostly straight; LBl 2.4, LBw 2.4; sternum hirsute, clothed with brown, medium length and shorter, dense setae. Abdomen: Densely clothed dorsally in short black/brown setae with numerous longer, lighter setae interspersed (generally red or orange in situ); dense dorsal patch of black Type I urticating bristles (Cooke et al. 1972); ventral side with shorter brown setae. Spermathecae: Paired and separate, tapering to capitate bulbs, possessing a secondary bulge near the interior of the base, with wide bases that are not fused. Legs: Hirsute, particularly ventrally; densely clothed in a mix of short, brown pubescence with longer setae interspersed. F1 12.9; F1w 4.36; P1 6.08; T1 9.69; M1 7.27; A1 5.97; F3 10.68; F3w 3.65; P3 5.22; T3 6.69; M3 7.79; A3 6.21; F4 12.46; F4w 4.03; P4 5.82; T4 10.34; M4 10.63; A4 7.12. All tarsi fully scopulate. Extent of metatarsal scopulation: leg III (SC3) = 60.5%; leg IV (SC4) = 28.2%. Two ventral spinose setae on metatarsus III; seven ventral spinose setae on metatarsus IV. Coxa I: Prolateral surface a mix of fine, hair-like, very thick tapered, and stout setae with a unique placement around the anterior and ventral, prolateral margin. Pedipalps: Densely clothed in the same setal color as the other legs; one spinose seta on the apical, prolateral femur and five spinose setae on the prolateral tibia.

Figure 18.

Aphonopelma armada (Chamberlin, 1940). A–G cleared spermathecae A armada holotype B APH_0547 C APH_0548 D APH_0807 E APH_0848 F APH_1049 G APH_1068.

Variation (6). Cl 11.96–16.76 (14.852±0.78), Cw 9.64–14.64 (12.702±0.78), LBl 1.7–2.4 (2.107±0.11), LBw 1.93–2.97 (2.45±0.14), F1 9.847–13.69 (11.846±0.56), F1w 3.31–4.45 (3.908±0.19), P1 4.427–6.51 (5.481±0.33), T1 7.642–10.33 (9.015±0.4), M1 5.741–7.44 (6.824±0.27), A1 4.659–6.84 (5.905±0.3), L1 length 32.316–44.74 (39.071±1.79), F3 7.03–11.18 (9.385±0.63), F3w 2.62–3.72 (3.192±0.19), P3 3.31–5.63 (4.588±0.33), T3 5.51–7.33 (6.422±0.27), M3 5.66–8.73 (7.21±0.41), A3 4.81–6.5 (5.732±0.26), L3 length 26.32–39.37 (33.337±1.84), F4 9.11–12.55 (11.117±0.56), F4w 2.72–4.03 (3.41±0.21), P4 3.85–5.82 (4.963±0.3), T4 7.67–10.35 (9.172±0.43), M4 7.81–11.74 (10.153±0.58), A4 5.58–7.59 (6.528±0.32), L4 length 34.02–47.93 (41.933±2.09), SC3 ratio 0.566–0.737 (0.629±0.02), SC4 ratio 0.282–0.429 (0.356±0.03), Coxa 1 setae = very thick tapered & stout. Spermathecae variation can be seen in Figure 18.

Material examined

United States: Texas: Andrews: SW4001 and SW7000, 32.131369 -102.621689 1, 3152ft., [APH_1049, 6/7/2010, 1♀, Skyler Stevens, AUMNH]; SW4001, 32.113981 -102.615814 1, 3140ft., [APH_1052, 6/7/2010, 1♂, Skyler Stevens, AUMNH]; Briscoe: Caprock Canyons State Park, Honey Flat camping area (site 4), 34.419514 -101.056081 1, 2611ft., [APH_0551-0554, 7/6/2009, 4 juv, Brent E. Hendrixson, Courtney Dugas, Sloan Click, AUMNH]; Burleson: Caldwell, 30.49425 -96.6921 5, 373ft., [APH_0944, 7/2008, 1♂, Dave Moellendorf, AUMNH]; Coleman: O.H. Ivie reservoir, Coleman Lake at Hords Creek, 31.842967 -99.5696 1, 1936ft., [APH_0840, 9/2008, 1♀, Chris A. Hamilton, AUMNH]; O.H. Ivie reservoir, 31.57695 -99.66065 1, 1571ft., [APH_0950, 9/2008, 1♂, Chris A. Hamilton, AUMNH]; Crosby: 0.2 miles N US-82 on FM 2591 (E of Crosbyton), 33.669122 -101.175656 1, 2802ft., [APH_0547-0550, 7/6/2009, 4 juv, Brent E. Hendrixson, Courtney Dugas, Sloan Click, AUMNH]; DeWitt: Westhoff, Meyer Rd off Hwy 240 and County Rd 142, 29.136983 -97.497033 1, 342ft., [APH_0922-0923, 9/2008, 2♂, Dan Lewis, AUMNH]; Ector: Cowden H Ranch, 32.076633 -102.789983 5, 3320ft., [APH_0855, 2006, 1♀, Dave Moellendorf, AUMNH]; [APH_0965-0967, 2006, 3♂, Dave Moellendorf, AUMNH]; [APH_0977, 2006, 1♀, Dave Moellendorf, AUMNH]; Fayette: La Grange, 29.914433 -96.866317 1, 321ft., [APH_0807, 5/2008, 1♀, Chris A. Hamilton, AUMNH]; Glasscock: E of Midland, off Hwy 137, 31.94904 -101.72346 2, 2569ft., [APH_1468, 19/6/2012, 1♂, Darryl Burton, AUMNH]; Hall: Hulver Cemetery, 5.9 miles W US-287 on Hwy-86, 34.522756 -100.534382 2, 1955ft., [APH_1460, 27/5/2012, 1♀, Brent E. Hendrixson, AUMNH]; Howard: Big Spring, at miniature golf course, 32.200567 -101.47715 1, 2701ft., [APH_0841-0844, 9/2008, 4♀, Chris A. Hamilton, AUMNH]; [APH_0945, 9/2008, 1♀, Chris A. Hamilton, AUMNH]; Kimble: Texas Tech Field Station, Junction, 30.472222 -99.780833 1, 1718ft., [APH_1067, 24/6/2010, 1♂, Skyler Stevens, AUMNH]; [APH_1068, 16/6/2010, 1♀, Bryce Hubbell, AUMNH]; [APH_1069, 17/6/2010, 1♀, Travis Fisher, AUMNH]; Kinney: 0.59 miles E US-277 on FM-693, 29.17044 -100.673259 1, 972ft., [APH_1164, 17/3/2010, 4 juv, Brent E. Hendrixson, Gerri Wilson, Thomas Martin, AUMNH]; Midland: Midland, 31.924217 -102.0583 1, 2784ft., [APH_0953, 9/2008, 1♀, Chris A. Hamilton, AUMNH]; W County Rd 54, 32.027933 -102.206853 1, 2883ft., [APH_1056-1059, 5/7/2010, 4♂, Skyler Stevens, AUMNH]; CR60, 32.010556 -102.2275 1, 2888ft., [APH_1061-1062, 2/7/2010, 2♂, Skyler Stevens, AUMNH]; near Midland, along Hwy-158, 31.999899 -102.180216 1, 2862ft., [APH_1172-1173, 21/7/2010, 1♀, 1♂, Brent E. Hendrixson, Brendon Barnes, Nate Davis, AUMNH]; near Midland, along CR-60, 32.003836 -102.256184 1, 2890ft., [APH_1174, 21/7/2010, 1♂, Brent E. Hendrixson, Brendon Barnes, Nate Davis, AUMNH]; 0.5 miles S CR 118W on Hwy-349, 31.58789 -101.9704 2, 2793ft., [APH_1462, 13/6/2012, 1♂, Darryl Burton, AUMNH]; 0.3 miles S FM-1787 on FM-1492, 31.66265 -102.20689 2, 2876ft., [APH_1466, 17/6/2012, 1♂, Darryl Burton, AUMNH]; Reagan: Hwy 137, 31.343611 -101.495833 1, 2617ft., [APH_1064-1065, 30/6/2010, 2♂, Skyler Stevens, AUMNH]; off Hwy 67, 31.23071389 -101.7264444 2, 2715ft., [APH_1385, 19/9/2011, 1 juv, Darryl Burton, AUMNH]; Scurry: Snyder, 32.682133 -100.925483 1, 2353ft., [APH_0845-0849, 9/2008, 5♀, Chris A. Hamilton, AUMNH]; Tom Green: San Angelo, approx. 1 miles N of W Hwy-67, 31.46431 -100.49511 4, 1922ft., [APH_0010, 23/6/2005, 1♂, Kati and Martha Mayfield, AUMNH]; Upton: oil fields W of Hwy 329, 31.28048056 -102.0802 2, 2605ft., [APH_1374, 17/6/2011, 1♀, Darryl Burton, AUMNH]; [APH_1375, 16/6/2011, 1 juv, Darryl Burton, AUMNH]; [APH_1377, 19/7/2011, 1 juv, Darryl Burton, AUMNH]; [APH_1379, 6/9/2011, 1 juv, Darryl Burton, AUMNH]; [APH_1381, 10/9/2011, 1♀, Darryl Burton, AUMNH]; FM 1492, 31.56408333 -102.1742556 2, 2855ft., [APH_1384, 11/9/2011, 1♀, Darryl Burton, AUMNH]; [APH_1387, 31/8/2011, 1 juv, Darryl Burton, AUMNH]; [APH_1388, 11/9/2011, 1 juv, Darryl Burton, AUMNH]; oil fields E of Hwy 329, 31.35340556 -102.0877333 2, 2695ft., [APH_1389, 30/8/2011, 1♀, Darryl Burton, AUMNH]; FM 1492, 31.56408333 -102.1742556 2, 2855ft., [APH_1391, 23/8/2011, 1♀, Darryl Burton, AUMNH]; Val Verde: Del Rio, near River Rd, 29.35553 -100.972297 5, 885ft., [APH_0593-0594, Spring 2009, 2 juv, unknown, AUMNH]; 2.2 miles off Hwy 90 on spur 406, N of Del Rio, 29.574484, -101.041898 1, 1195ft., [APH_3124, 15/6/2014, 1♀, Dave Moellendorf, AUMNH].

Distribution and natural history

Aphonopelma armada has a wide distribution across Texas and can be found inhabiting these Level III Ecoregions: Chihuahuan Deserts, High Plains, Southwestern Tablelands, Central Great Plains, Edwards Plateau, Southern Texas Plains, Texas Blackland Prairies, and East Central Texas Plains (Fig. 19). Aphonopelma armada can be found in syntopy with a number of other species across its distribution: A. anax, A. gabeli, A. hentzi, A. moderatum, and A. moellendorfi. Mating season, when mature males emerge, is similar to the other syntopic species (i.e., males are more active during the evenings and early mornings from late spring to early summer).

Figure 19.

Aphonopelma armada (Chamberlin, 1940). A distribution of known specimens B predicted distribution; warmer colors (red, orange, yellow) represent areas of high probability of occurrence, cooler colors (blue shades) represent areas of low probability of occurrence.

Conservation status

Aphonopelma armada is very common throughout its distribution in South and West Texas. The species is likely secure.

Remarks

The type locality for A. armada is vague (Austin, Texas) and despite much fieldwork in the region, we have never been able to find specimens referable to A. armada in or around the Austin city limits (this area is dominated by A. hentzi). We have, however, found the species in counties to the south and east of Austin and note that it has a crescent-like distribution around Austin and becomes more common to the west (Fig. 19). Other important ratios that distinguish males: A. armada possess a larger CL/M1 (≥1.36; 1.36–1.47) than A. moderatum (≤1.30; 1.13–1.30), and A. moellendorfi (≤1.31; 1.10–1.31); as well as possessing a smaller F4/M4 (≤0.92; 0.86–0.92) than A. hentzi (≥0.92; 0.92–1.03). Other important ratios distinguish females: A. armada possess a smaller T3/T4 (≤0.73; 0.64–0.73) than A. gabeli (≥0.73; 0.73–0.78); as well as possessing a smaller L4/Cl (≤2.89; 2.73–2.89) than A. moderatum (≥2.89; 2.89–3.09). Due to the amounts of variation, small number of specimens, and/or the small differences between species no others are claimed to be significant at this time (see Suppl. material 2). During evaluation of traditional PCA morphospace, males of A. armada separate from A. gabeli, A. moderatum, A. moellendorfi along PC1~2, but do not separate from A. hentzi or A. anax. Females do not separate from A. anax, A. gabeli, A. hentzi, and A. moderatum. Females of A. moellendorfi are unknown and cannot be compared. Interestingly, A. armada males separate from A. gabeli, A. moderatum, and A. moellendorfi in three-dimensional PCA morphospace (PC1~PC2~PC3), but do not separate from A. anax and A. hentzi. Aphonopelma armada females separate from A. anax, but do not separate from A. gabeli, A. hentzi, and A. moderatum. PC1, PC2, and PC3 explain ≥87% of the variation in male analyses and ≥96% of the variation in female analyses.

We did not examine the holotype of A. arnoldi but we did have the opportunity to study freshly collected topotypic material of the species from Crosbyton, Texas. Our morphological and molecular analyses fail to recognize this species as a separate, independently evolving lineage. As a consequence, we consider A. arnoldi a junior synonym of A. armada.

Aphonopelma atomicum Hamilton, sp. n.

Figures 20, 21, 22, 23, 24

Aphonopelma mojave Prentice, 1997 (in part): 161.

Types

Male holotype (APH_2727-2) collected at the Nevada Testing Site, Nye Co., Nevada, 37.025579 -116.023865 6, elev. 3990ft., viii.1961, coll. Gertsch; deposited in AMNH. Paratype female (AUMS_3267-2) from 10 miles W of Mercury, Mercury Valley, Nye Co., Nevada, 36.691928 -116.174483 5, elev. 3470ft., 3.xi.1972, coll. Wendell Icenogle; deposited in AUMNH. Paratype male (AUMS_2638) from Nros Rd., 20 miles W of Mercury, Nye Co., Nevada, 36.66182 -116.368 5, elev. 2744ft., 3.xi.1972, coll. C.S.L.; deposited in AUMNH.

Etymology

The specific epithet is a neuter noun meaning “of or relating to atoms”. The name references the Nevada Test Site, constructed by the Atomic Energy Commission for the testing of nuclear devices, where this species was originally collected. The name is in homage to the famous sci-fi B movies of the 1950’s, of which Tarantula (1955) was the most entertaining, and slightly ironic given that this species is one of the smallest tarantulas in the United States.

Diagnosis

Aphonopelma atomicum (Fig. 20) is a member of the Paloma species group and can be distinguished by a combination of morphological, molecular, and geographic characteristics. Nuclear and mitochondrial DNA identifies A. atomicum as a strongly supported monophyletic lineage (Figs 78) that is a sister lineage to A. mojave, A. icenoglei sp. n., A. prenticei sp. n., and A. joshua. Aphonopelma atomicum can easily be differentiated from larger syntopic species (e.g., A. iodius) due to their much smaller size and the extent of metatarsal scopulation on legs III and IV, and from all other miniaturized species of Aphonopelma by their locality (with the exception of A. prenticei). Molecular data is needed to differentiate this species from A. prenticei (see Figs 7, 8). The most significant measurements that distinguish male A. atomicum from its closely related phylogenetic and syntopic species are F3 and the extent of metatarsus IV scopulation. Male A. atomicum can be distinguished by possessing a larger F3/M3 (≥0.99; 0.99–1.03) than A. joshua (≤0.98; 0.91–0.98) and A. xwalxwal sp. n. (≤0.95; 0.91–0.95); a smaller F3L/W (≤3.40; 2.92–3.40) than A. icenoglei (≥3.51; 3.51–3.90); and a smaller L4 scopulation extent (26%-41%) than A. iodius (62%-88%). There are no significant measurements that separate male A. atomicum from A. mojave or A. prenticei (A. atomicum males possess a swollen F3 femur like A. prenticei, whereas A. mojave males do not). The most significant measurements that distinguish female A. atomicum from its closely related phylogenetic and syntopic species are F3 and the extent of metatarsus IV scopulation. Female A. atomicum can be distinguished by possessing a smaller A1/F3 (≤0.56; 0.53–0.56) than A. mojave (>0.56; 0.56–0.67), A. icenoglei (≥0.59; 0.59–0.67), and A. joshua (≥0.57; 0.57–0.64); a smaller Cl/F3 (≤1.38; 1.31–1.38) than A. prenticei (≥1.39; 1.39–1.64); and smaller L4 scopulation extent (37%–49%) than A. iodius (59%–83%).

Figure 20.

Aphonopelma atomicum sp. n. live photograph. Female - APH_1478. Do not have a photograph of a live male specimen.

Description of male holotype

(AUMS_2727-2; Fig. 21). Specimen preparation and condition: Specimen originally preserved in unknown percentage of ethanol; original coloration faded due to preservation. Missing leg IV right side. Left legs I, III, IV, and left pedipalp removed for measurements and photographs; stored in vial with specimen. No tissue for DNA. General coloration: Brown/faded brown. Cephalothorax: Carapace 6.523 mm long, 6.382 mm wide; densely clothed with brown/faded brown pubescence, appressed to surface; fringe covered in long setae not closely appressed to surface; foveal groove medium deep and straight; pars cephalica region rises gradually from foveal groove on a straight plane towards the ocular area; AER slightly procurved, PER slightly recurved; normal sized chelicerae; clypeus mostly straight; LBl 0.918, LBw 1.107; sternum hirsute, clothed with faded brown, densely packed, short setae. Abdomen: Densely clothed in short brown pubescence with numerous longer, lighter setae interspersed (generally red or orange in situ); dense dorsal patch of black Type I urticating bristles (Cooke et al. 1972) - smaller and distinct from large species. Legs: Hirsute; densely clothed in faded brown pubescence. Metatarsus I straight/very slightly curved. F1 7.691; F1w 1.62; P1 3.012; T1 6.618; M1 5.767; A1 3.83; F3 6.665; F3w 1.958; P3 2.436; T3 5.294; M3 6.438; A3 4.207; F4 7.922; F4w 1.596; P4 2.656; T4 6.615; M4 8.407; A4 4.358; femur III is swollen/slightly swollen. All tarsi fully scopulate. Extent of metatarsal scopulation: leg III (SC3) = 61.4%; leg IV (SC4) = 37.3%. No distinct ventral spinose setae on metatarsus III; three ventral spinose setae on metatarsus IV; two prolateral spinose setae on tibia I; one megaspine present on the retrolateral tibia, at the apex of the mating clasper; three megaspines on the apex of the retrolateral branch of the tibial apophyses. Coxa I: Prolateral surface covered by fine, hair-like setae. Pedipalps: Hirsute; densely clothed in the same setal color as the other legs, with numerous longer ventral setae; one spinose seta at the apical, prolateral femur and two prolateral spinose setae on the palpal tibia; PTl 4.486, PTw 1.51. When extended, embolus tapers with a curve to the retrolateral side; embolus slender, no keels; distinct dorsal and ventral transition from bulb to embolus.

Figure 21.

Aphonopelma atomicum sp. n. A–I male holotype, APH_2727-2 A dorsal view of carapace, scale bar = 2mm B prolateral view of coxa I C dorsal view of femur III D ventral view of metatarsus III, scale bar = 3mm E ventral view of metatarsus IV, scale bar = 3mm F prolateral view of L pedipalp and palpal tibia, scale bar = 3mm G dorsal view of palpal bulb H retrolateral view of palpal bulb, scale bar = 0.5mm I prolateral view of tibia I (mating clasper), scale bar = 2.5mm.

Variation (5). Cl 6.523–7.917 (7.157±0.23), Cw 6.382–6.798 (6.64±0.09), LBl 0.832–0.988 (0.927±0.03), LBw 1.098–1.394 (1.231±0.06), F1 7.434–8.185 (7.761±0.12), F1w 1.565–1.914 (1.783±0.08), P1 2.859–3.481 (3.12±0.11), T1 6.618–7.496 (7.053±0.16), M1 5.767–6.402 (6.083±0.12), A1 3.768–4.149 (3.976±0.08), L1 length 26.653–28.926 (27.993±0.5), F3 6.665–7.388 (6.962±0.12), F3w 1.958–2.454 (2.223±0.09), P3 2.355–2.84 (2.562±0.08), T3 5.294–5.812 (5.58±0.1), M3 6.438–7.349 (6.88±0.15), A3 4.204–4.298 (4.252±0.02), L3 length 25.04–27.374 (26.236±0.39), F4 7.773–8.406 (8.073±0.11), F4w 1.596–2.045 (1.816±0.08), P4 2.621–2.9 (2.786±0.06), T4 6.615–7.533 (7.11±0.17), M4 8.197–9.072 (8.573±0.15), A4 4.358–4.937 (4.64±0.1), L4 length 29.958–32.832 (31.181±0.54), PTl 4.486–4.966 (4.689±0.08), PTw 1.433–1.736 (1.564±0.06), SC3 ratio 0.552–0.725 (0.626±0.03), SC4 ratio 0.264–0.409 (0.356±0.03), Coxa I setae = very thin tapered, F3 condition = slightly swollen/swollen.

Description of female paratype

(AUMS_3267-2; Figs 2223). Specimen preparation and condition: Specimen originally preserved in unknown percentage of ethanol; original coloration faded due to preservation. Left legs I, II, III, IV, and pedipalp removed for photographs and measurements; stored in vial with specimen. No tissue for DNA. Genital plate with spermathecae removed and cleared, stored in vial with specimen. General coloration: Brown and faded brown. Cephalothorax: Carapace 7.291 mm long, 6.814 mm wide; Hirsute, densely clothed with short faded brown pubescence closely appressed to surface; fringe densely covered in slightly longer setae; foveal groove medium deep and procurved; pars cephalica region gently rises from thoracic furrow, arching anteriorly toward ocular area; AER slightly procurved, PER very slightly recurved - mostly straight; chelicerae robust, clypeus extends forward on a slight curve; LBl 1.058, LBw 1.416; sternum hirsute, clothed with short faded brown setae. Abdomen: Densely clothed dorsally in short faded brown setae with longer, lighter setae (generally red or orange in situ) focused near the urticating patch; dense dorsal patch of black Type I urticating bristles (Cooke et al. 1972) - smaller and distinct from large species. Spermathecae: Paired and separate, short, with capitate bulbs widening towards the bases; not fused. Legs: Hirsute; densely clothed in short faded brown pubescence; F1 6.49; F1w 1.891; P1 2.912; T1 5.232; M1 3.952; A1 3.102; F3 5.539; F3w 1.789; P3 2.418; T3 4.188; M3 4.363; A3 3.418; F4 6.898; F4w 1.766; P4 2.753; T4 5.716; M4 5.993; A4 3.839. All tarsi fully scopulate. Extent of metatarsal scopulation: leg III (SC3) = 68.3%; leg IV (SC4) = 37.2%. One ventral spinose setae on metatarsus III; four ventral spinose setae on metatarsus IV, with numerous thickened setae throughout. Coxa I: Prolateral surface covered by very thin tapered and fine, hair-like setae. Pedipalps: Densely clothed in the same setal color as the other legs; one spinose seta on the apical, prolateral femur, five prolateral (three at the apical, prolateral border with the tarsus) spinose setae and one ventral spinose seta on the tibia.

Figure 22.

Aphonopelma atomicum sp. n. A–E female paratype, APH_3267-2 A dorsal view of carapace, scale bar = 2mm B prolateral view of coxa I C ventral view of metatarsus III, scale bar = 1.5mm D ventral view of metatarsus IV, scale bar = 1.5mm E prolateral view of L pedipalp and palpal tibia.

Figure 23.

Aphonopelma atomicum sp. n. A–C cleared spermathecae A APH_2727 B APH_3267 C APH_3267-2.

Variation (3). Cl 7.081–7.978 (7.45±0.27), Cw 5.951–7.105 (6.623±0.35), LBl 0.922–1.218 (1.066±0.09), LBw 1.105–1.621 (1.381±0.15), F1 5.656–7.25 (6.465±0.46), F1w 1.663–2.032 (1.862±0.11), P1 2.912–3.253 (3.026±0.11), T1 5.232–5.714 (5.394±0.16), M1 3.461–4.532 (3.982±0.31), A1 2.787–3.179 (3.023±0.12), L1 length 20.052–23.928 (21.889±1.12), F3 5.11–5.984 (5.544±0.25), F3w 1.493–1.907 (1.73±0.12), P3 2.046–2.81 (2.425±0.22), T3 3.603–4.223 (4.005±0.2), M3 3.981–4.61 (4.318±0.18), A3 3.305–3.641 (3.455±0.1), L3 length 18.045–21.268 (19.746±0.93), F4 6.234–7.413 (6.848±0.34), F4w 1.473–1.956 (1.732±0.14), P4 2.541–2.92 (2.738±0.11), T4 5.234–6.039 (5.663±0.23), M4 5.687–6.245 (5.975±0.16), A4 3.446–3.86 (3.715±0.13), L4 length 23.142–26.477 (24.939±0.97), SC3 ratio 0.683–0.755 (0.726±0.02), SC4 ratio 0.372–0.49 (0.439±0.03), Coxa I setae = thin tapered. Spermathecae variation can be seen in Figure 23.

Material examined

United States: California: Inyo: Death Valley National Park, Hwy-178, Salsberry Pass, 35.923118 -116.431747 1, 3204ft., [APH_1478, 30/7/2012, 1♀, Brent E. Hendrixson, Brendon Barnes, Austin Deskewies, AUMNH]; Death Valley National Park, ~7.5 miles S Hwy-190 on Dante’s View Rd, 36.267976 -116.665576 1, 3242ft., [APH_1479, 30/7/2012, 1 juv, Brent E. Hendrixson, Brendon Barnes, Austin Deskewies, AUMNH]; Nevada: Nye: Rock Valley Wash, off Hwy-95, 36.622963 -116.309464 1, 2784ft., [APH_1549, 24/10/2012, 1♀, Brent E. Hendrixson, AUMNH]; Rock Valley, 36.632732 -116.313933 5, 2884ft., [AUMS_2637, 20/10/1972, 1♀, E.L. Sleeper, AUMNH]; Mercury, 36.66051 -115.994475 5, 3796ft., [APH_2725-1, 6/9/1960, 2♀, 14♂, Gertsch, AMNH]; Nevada Testing Site, 37.025579 -116.023865 6, 3990ft., [APH_2725-2, 1/10/1959, 3♂, JRM, AMNH]; [APH_2727, 29/6/1961, 3♀, 15♂, Gertsch, AMNH]; [APH_2727-2, 8/1961, 1♂, Gertsch, AMNH]; [APH_2730, 16/8/1964, 1♀, unknown, AMNH]; Nros Rd., 20 miles W of Mercury, 36.66182 -116.368 5, 2744ft., [AUMS_2638, 3/11/1972, 1♂, C. S. L., AUMNH]; 10 miles W of Mercury, Mercury Valley, 36.691928 -116.174483 5, 3470ft., [AUMS_3267 & 3267-2, 3/11/1972, 2♀, W. Icenogle, AUMNH].

Distribution and natural history

Aphonopelma atomicum is only known from a handful of specimens surrounding the Amargosa Desert in southern Nye County (Nevada) and southeastern Inyo County (California), including the Amargosa Range and Nevada Test Site (Fig. 24). Specimens have been collected at elevations between 850 and 1220 meters, inhabiting the Mojave Basin and Range Level III Ecoregion. This species is likely syntopic with A. iodius throughout its range and may be found near populations of A. prenticei. Burrow entrances are generally surrounded by a distinct mound or turret made of excavated soil and silk (Fig. 2D–E). Mating most likely occurs during daylight hours in autumn (October-November).

Figure 24.

Aphonopelma atomicum sp. n. distribution of known specimens. There is no predicted distribution map due to the limited number of sampling localities and restricted distribution this species possesses.

Conservation status

Aphonopelma atomicum has a highly restricted distribution limited to the mountains and foothills surrounding the Amargosa Desert and Death Valley. While this species is not dramatically different from A. prenticei, it is genetically unique and should be considered important. The species is most likely secure.

Remarks

Aphonopelma atomicum is unique because it was quite possibly the first miniature tarantula species collected in the United States (although it was never described and sat on a shelf in the AMNH collection). It is somewhat puzzling that Gertsch never described this species given the number of individuals he collected and how radically different (i.e., small in size) specimens are from all other Aphonopelma in that region. Other important ratios that distinguish males: A. atomicum possess a smaller A1/F3 (≤0.58; 0.55–0.58) than A. mojave (≥0.58; 0.58–0.63) and A. prenticei (≥0.58; 0.58–0.65). Certain morphometrics have potential to be useful, though due to the amounts of variation, small number of specimens, and the small differences between species, no other are claimed to be significant at this time (see Suppl. material 2). During evaluation of traditional two-dimensional PCA morphospace and three-dimensional PCA morphospace (PC1~PC2~PC3), males of A. atomicum separate from A. iodius, A. joshua, and A. xwalxwal along PC1~2, but do not separate from A. mojave, A. icenoglei, or A. prenticei. Female atomicum separate from A. iodius in morphological space, but do not separate from the other miniature species (A. mojave, A. icenoglei, and A. prenticei). There are no known female A. xwalxwal at this time to compare. PC1, PC2, and PC3 explain ≥97% of the variation in all analyses.

Aphonopelma catalina Hamilton, Hendrixson & Bond, sp. n.

Figures 25, 26, 27, 28, 29

Types

Male holotype (APH_1440) from Coronado National Forest, along Bug Spring Trail, Pima Co., Arizona, 32.34544 -110.71602 4, elev. 5255ft., 17.xii.2011, coll. Brent E. Hendrixson and Thomas Martin; deposited in AUMNH. Paratype female (APH_1602) from Coronado National Forest, along Bug Spring Trail, Pima Co., Arizona, 32.34544 -110.71602 4, elev. 5255ft., 9.xi.2012, coll. Brent E. Hendrixson; deposited in AUMNH. Paratype male (APH_1439) from Coronado National Forest, along Bug Spring Trail, Pima Co., Arizona, 32.34544 -110.71602 4, elev. 5255ft., 17.xii.2011, coll. Brent E. Hendrixson and Thomas Martin; deposited in AMNH.

Etymology

The specific epithet is a noun in apposition taken from type locality, the Santa Catalina Mountains near Tucson, Arizona, where this new species appears to be endemic.

Diagnosis

Aphonopelma catalina (Fig. 25) is a member of the Marxi species group and can be distinguished by a combination of morphological, molecular, and geographic characteristics. Nuclear and mitochondrial DNA identifies A. catalina as a phylogenetically distinct monophyletic lineage (Figs 78), supported as a sister lineage to A. chiricahua sp. n. (a species endemic to the Chiricahua Mountains). The significant measurement that distinguishes male A. catalina from its closely related phylogenetic and syntopic species is F1. Male A. catalina can be distinguished by possessing a larger F1/A3 (≥2.25; 2.25–2.49) than A. chiricahua (≤1.86; 1.55–1.86), A. madera sp. n. (≤2.14; 2.02–2.14), A. parvum sp. n. (≤1.98; 1.79–1.98), A. peloncillo sp. n. (≤2.23; 1.86–2.23), A. saguaro sp. n. (1.96 ± (only 1 specimen)), and A. vorhiesi (≤2.17; 1.71–2.17). Significant measurements that distinguish female A. catalina from its closely related phylogenetic and syntopic species are Cl and M3. Female A. catalina can be distinguished by possessing a larger M3/A4 (≥1.07; 1.07–1.10) than A. chiricahua (0.80 ± (only 1 specimen)), A. madera (≤1.07; 0.96–1.07), A. parvum (≤0.92; 0.86–0.92), and A. saguaro (0.92 ± (only 1 specimen)), but smaller than A. peloncillo (≥1.11; 1.11–1.23); and by possessing a smaller Cl/F4 (≤1.18; 1.17–1.18) than A. chiricahua (1.21 ± (only 1 specimen)), A. madera (≥1.20; 1.20–1.27), A. peloncillo (≥1.22; 1.22–1.34), and A. vorhiesi (≥1.23; 1.23–1.37).

Figure 25.

Aphonopelma catalina sp. n. specimens, live photographs. Female paratype (L) - APH_1602; Male (R) - APH_1438.

Description of male holotype

(APH_1440; Fig. 26). Specimen preparation and condition: Specimen collected live crossing trail, preserved in 80% ethanol; original coloration faded due to preservation. Left legs I, III, IV, and left pedipalp removed for measurements and photographs; stored in vial with specimen. Right leg III removed for DNA and stored at -80°C in the AUMNH (Auburn, AL). General coloration: Generally black or faded black. Cephalothorax: Carapace 12.39 mm long, 11.33 mm wide; densely clothed with black/faded black pubescence, appressed to surface; fringe covered in long setae not closely appressed to surface; foveal groove medium deep and procurved; pars cephalica region rises gradually from foveal groove, gently arching anteriorly toward ocular area; AER procurved, PER recurved; normal sized chelicerae; clypeus straight; LBl 1.18, LBw 1.56; sternum hirsute, clothed with short black/brown, densely packed setae. Abdomen: Densely clothed in short black/brown pubescence with numerous longer, lighter setae interspersed (generally red or orange in situ); dense dorsal patch of black Type I urticating bristles (Cooke et al. 1972); ventral setae same as dorsal. Legs: Hirsute; densely clothed with short, similar length black/brown setae, and longer setae dorsally. Metatarsus I very slightly curved. F1 14.64; F1w 2.83; P1 5.31; T1 12.78; M1 8.91; A1 5.95; F3 10.40; F3w 2.88; P3 3.89; T3 8.63; M3 9.18; A3 5.89; F4 12.90; F4w 2.78; P4 4.54; T4 10.96; M4 12.34; A4 7.11; femur III is normal - not noticeably swollen or wider than other legs. All tarsi fully scopulate. Extent of metatarsal scopulation: leg III (SC3) = 53.0%; leg IV (SC4) = 30.6%. Five ventral spinose setae on metatarsus III; eleven ventral spinose setae, one prolateral and one retrolateral spinose seta on metatarsus IV; one large megaspine is present on the retrolateral tibia at the apex of the mating clasper - this can be seen when viewing the prolateral face of the mating clasper; the prolateral branch of the tibial apophyses possesses a very large megaspine that projects anteriorly. Coxa I: Prolateral surface a mix of fine, hair-like and tapered/thin tapered setae. Pedipalps: Hirsute; densely clothed in the same setal color as the other legs, with numerous longer ventral setae; one spinose seta at the apical, prolateral femur and two spinose setae on the prolateral tibia; PTl 8.058, PTw 2.911. When extended, embolus tapers with a gentle curve to the retrolateral side near apex; embolus slender, no keels.

Figure 26.

Aphonopelma catalina sp. n. A–I male holotype, APH_1440 A dorsal view of carapace, scale bar = 4mm B prolateral view of coxa I C dorsal view of femur III D ventral view of metatarsus III, scale bar = 3mm E ventral view of metatarsus IV, scale bar = 4mm F prolateral view of L pedipalp and palpal tibia, scale bar = 4mm G dorsal view of palpal bulb H retrolateral view of palpal bulb, scale bar = 1mm I prolateral view of tibia I (mating clasper), scale bar = 4.5mm.

Variation (4). Cl 9.57–12.39 (10.595±0.62), Cw 8.88–11.33 (9.708±0.55), LBl 0.967–1.237 (1.114±0.06), LBw 1.102–1.56 (1.354±0.11), F1 11.28–14.64 (12.765±0.7), F1w 2.18–2.83 (2.467±0.14), P1 3.73–5.31 (4.483±0.32), T1 9.89–12.78 (11.132±0.61), M1 6.42–8.91 (7.446±0.54), A1 5.0–5.95 (5.392±0.2), L1 length 36.32–47.59 (41.217±2.34), F3 7.97–10.4 (9.098±0.5), F3w 2.1–2.88 (2.465±0.16), P3 3.04–4.084 (3.551±0.26), T3 5.74–8.63 (7.039±0.6), M3 6.72–9.18 (7.882±0.51), A3 4.82–5.89 (5.37±0.23), L3 length 28.29–37.99 (32.94±2.02), F4 9.75–12.9 (11.077±0.66), F4w 2.05–2.78 (2.357±0.15), P4 3.26–4.54 (3.902±0.26), T4 7.82–10.96 (9.444±0.64), M4 9.26–12.34 (10.762±0.63), A4 5.41–7.11 (6.095±0.4), L4 length 35.5–47.85 (41.279±2.53), PTl 6.587–8.058 (7.173±0.32), PTw 2.328–2.911 (2.572±0.12), SC3 ratio 0.481–0.53 (0.514±0.01), SC4 ratio 0.228–0.359 (0.286±0.03), Coxa I setae = tapered/thin tapered, F3 condition = normal.

Description of female paratype

(APH_1602; Figs 2728). Specimen preparation and condition: Specimen collected live from burrow, preserved in 80% ethanol; original coloration faded due to preservation. Left legs I, III, IV, and pedipalp removed for photographs and measurements; stored in vial with specimen. Right leg III removed for DNA and stored at -80°C in the AUMNH (Auburn, AL). Genital plate with spermathecae removed and cleared, stored in vial with specimen. General coloration: Black/faded black and brown. Cephalothorax: Carapace 14.79 mm long, 13.74 mm wide; Hirsute, densely clothed with black/faded black, pubescence closely appressed to surface; fringe densely covered in longer setae; foveal groove medium deep and straight; pars cephalica region gently rises from thoracic furrow, arching anteriorly toward ocular area; AER very slightly procurved, PER straight; robust chelicerae, clypeus extends forward on a slight curve; LBl 1.81, LBw 1.97; sternum hirsute, clothed with shorter black/faded black setae. Abdomen: Densely clothed dorsally in short black setae with numerous longer, lighter setae interspersed (generally red or orange in situ); dense dorsal patch of black Type I urticating bristles (Cooke et al. 1972); ventral setae same as dorsal. Spermathecae: Paired and separate, tapering and slightly curving medially towards capitate bulbs, with wide bases that appear fused. Legs: Very hirsute, particularly ventrally; densely clothed in short and medium black pubescence, with longer setae colored similarly as the long abdominal setae; F1 11.87; F1w 3.84; P1 5.36; T1 9.77; M1 6.92; A1 6.04; F3 9.74; F3w 3.32; P3 4.52; T3 7.26; M3 7.29; A3 6.07; F4 12.52; F4w 3.41; P4 4.92; T4 10.00; M4 10.98; A4 6.64. All tarsi fully scopulate. Extent of metatarsal scopulation: leg III (SC3) = 68.6%; leg IV (SC4) = 46.5%. Six ventral spinose setae on metatarsus III; nine ventral spinose setae and one prolateral spinose seta on metatarsus IV. Coxa I: Prolateral surface a mix of fine, hair-like and tapered setae. Pedipalps: Densely clothed in the same setal color as the other legs; two prolateral spinose setae and one ventral spinose seta on the tibia.

Figure 27.

Aphonopelma catalina sp. n. A–E female paratype, APH_1602 A dorsal view of carapace, scale bar = 6mm B prolateral view of coxa I C ventral view of metatarsus III, scale bar = 3mm D ventral view of metatarsus IV, scale bar = 3mm E prolateral view of L pedipalp and palpal tibia.

Figure 28.

Aphonopelma catalina sp. n. cleared spermathecae. A APH_1602 B AUMS_2615.

Variation (2). Cl 14.79–16.39 (15.59±0.8), Cw 13.74–15.06 (14.4±0.66), LBl 1.81–2.03 (1.92±0.11), LBw 1.97–2.67 (2.32±0.35), F1 11.87–13.85 (12.86±0.99), F1w 3.84–4.24 (4.04±0.2), P1 5.36–5.58 (5.47±0.11), T1 9.77–11.33 (10.55±0.78), M1 6.92–7.57 (7.245±0.33), A1 6.04–6.05 (6.045±0), L1 length 39.96–44.38 (42.17±2.21), F3 9.74–11.31 (10.525±0.79), F3w 3.32–3.78 (3.55±0.23), P3 4.52–5.33 (4.925±0.41), T3 7.26–8.35 (7.805±0.55), M3 7.29–8.42 (7.855±0.57), A3 6.07–6.77 (6.42±0.35), L3 length 34.88–40.18 (37.53±2.65), F4 12.52–13.95 (13.235±0.72), F4w 3.41–3.86 (3.635±0.23), P4 4.92–5.54 (5.23±0.31), T4 10.0–11.17 (10.585±0.59), M4 10.98–11.74 (11.36±0.38), A4 6.64–7.84 (7.24±0.6), L4 length 45.06–50.24 (47.65±2.59), SC3 ratio 0.629–0.686 (0.657±0.03), SC4 ratio 0.37–0.465 (0.418±0.05), Coxa I setae = medium tapered. Spermathecae variation can be seen in Figure 28.

Material examined

United States: Arizona: Pima: Coronado National Forest, along Bug Spring Trail, 32.34544 -110.71602 4, 5255ft., [APH_0454, 28/12/2008, 1♂, Paul E. Marek, Charity Hall, AUMNH]; [APH_1438, 11/12/2011, 1♂, Jillian Cowles, Bill Savary, AUMNH]; [APH_1439-1440, 17/12/2011, 2♂, Brent E. Hendrixson, Thomas Martin, AUMNH & AMNH]; [APH_1602, 9/11/2012, 1♀, Brent E. Hendrixson, AUMNH]; .5 miles W of Catalina Hwy, 2 miles N of Molino Basin, Santa Catalina Mtns, 32.336526 -110.718179 4, 4886ft., [AUMS_2615, 15/9/1974, 1♀, W. Icenogle, AUMNH].

Distribution and natural history

Aphonopelma catalina is known from only six individuals collected within a few kilometers of each other but this species appears to be a sky island endemic found in the Santa Catalina Mountains in Pima County, Arizona at elevations above 1480 meters in oak-grassland communities (Figs 1, 29); it is possible that this species is also found in bordering sections of the Rincon Mountains but the range has not been thoroughly sampled for tarantulas. Aphonopelma catalina can be found inhabiting the Madrean Archipelago Level III Ecoregion; the upper elevational limit for A. catalina remains unknown but the species can likely be found in adjacent oak and pine-oak woodlands at higher elevations similar to its close relatives A. chiricahua and A. madera. This species has not been observed in syntopy with other Aphonopelma spp. but can probably be found alongside A. chalcodes and A. vorhiesi in oak-grassland communities along the lower elevations of the Santa Catalina Mountains. Only a single burrow (of an adult female, APH_1602, Fig. 25) of this species has been observed, located a few meters from a hiking trail surrounded by grasses and large rocks. The structure of the burrow was fairly typical for a North American theraphosid (i.e., circular and covered by a thin veil of silk) and did not appear too deep or well drained; the specimen was easily flooded from its burrow with less than a liter of water. Each adult male of A. catalina examined in this study was found during daylight hours in December, suggesting that the breeding period for this species is restricted to late autumn and early winter; one of these males was found wandering along a snow bank on a sunny afternoon (APH_0454, P. Marek 2008, pers. comm.). An unconfirmed adult male (no voucher specimen available, identification tentatively assigned based on a photograph and locality data) was found above 1760 meters in an oak woodland community during early February (P. MacDuff 2015, pers. comm.).

Figure 29.

Aphonopelma catalina sp. n. distribution of known specimens. There is no predicted distribution map due to the limited number of sampling localities and restricted distribution this species possesses.

Conservation status

It is difficult to fully assess the distribution and abundance (and therefore the conservation status) of A. catalina due to a lack of specimens and thorough sampling; however, as previously mentioned, the species appears to be narrowly endemic to the Santa Catalina Mountains, which may put the species at some risk. This mountain range is entirely contained within the Coronado National Forest (Santa Catalina Ranger District) which is afforded some degree of protection; however, increased urbanization of the Tucson Metropolitan Area (one of the most rapidly growing areas in the United States), increased recreation in the mountains, and climate change have impacted these habitats (Coronado Planning Partnership 2008, Brusca et al. 2013, Moore et al. 2013, Hendrixson et al. 2015) and pose additional threats to A. catalina.

Remarks

As noted in Hendrixson et al. (2015), A. catalina is morphologically very similar to other Madrean sky island endemics in the Marxi species group, although generally larger than A. chiricahua and A. madera and possess a more hirsute appearance than A. vorhiesi. Other important ratios that distinguish males: A. catalina possess a larger T1/M3 (≥1.39; 1.39–1.47) than A. parvum (≤1.34; 1.16–1.34) and A. saguaro (1.16 ± (only 1 specimen)); by possessing a smaller Cl/T1 (≤0.97; 0.90–0.97) than A. chiricahua (≥1.00; 1.00–1.17), A. madera (≥1.06; 1.06–1.12), A. peloncillo (≥1.01; 1.01–1.22), and A. vorhiesi (≥0.99; 0.99–1.34). Other important ratios that distinguish females: A. catalina possess a smaller A3/M4 (≤0.58; 0.55–0.58) than A. chiricahua (0.71 ± (only 1 specimen)), A. parvum (≥0.60; 0.60–0.69), and A. saguaro (0.67 ± (only 1 specimen)); by possessing a smaller Cl/M3 (≤2.03; 1.94–2.03) than A. chiricahua (2.36 ± (only 1 specimen)), A. madera (≥2.06; 2.06–2.31), and A. saguaro (2.10 ± (only 1 specimen)); by possessing a smaller Cl/Cw (≤1.09; 1.07–1.09) than A. chiricahua (1.02 ± (only 1 specimen)), A. peloncillo (≥1.11; 1.11–1.17), and A. vorhiesi (≥1.11; 1.11–1.21). For both males and females, certain morphometrics have potential to be useful, though due to the amounts of variation, small number of specimens, and the small differences between species, no other are claimed to be significant at this time (see Suppl. material 2). During evaluation of PCA morphospace, males of A. catalina separate from A. chalcodes, A. peloncillo, A. vorhiesi, and all miniature species along PC1~2, but do not separate from A. chiricahua, A. madera, and A. marxi. Females of A. catalina separate from A. chalcodes, A. chiricahua and all miniature species along PC1~2, but do not separate from A. madera, A. marxi, A. peloncillo, and A. vorhiesi. Interestingly, A. catalina males separate from A. chalcodes, A. peloncillo, A. saguaro, and A. vorhiesi in three-dimensional PCA morphospace (PC1~PC2~PC3), but do not separate from A. chiricahua, A. madera, and A. marxi. Aphonopelma catalina females separate from A. chalcodes, A. chiricahua, A. marxi, and A. saguaro but do not separate from A. madera, A. peloncillo, and A. vorhiesi. PC1, PC2, and PC3 explain ≥96% of the variation in all analyses.

Aphonopelma chalcodes Chamberlin, 1940

Figures 30, 31, 32, 33, 34, 35; Suppl. material 4

Aphonopelma chalcodes Chamberlin, 1940: 7; male holotype, male paratype, and two female paratypes from Tucson, Pima Co., Arizona, 32.221743 -110.9264796, elev. 2473ft., 27.vii.1936, coll. Prof. C.T. Vorhies; deposited in AMNH. [examined]

Rhechostica chalcodes Raven, 1985: 149.

Aphonopelma chalcodes Smith, 1995: 82.

Aphonopelma apacheum Chamberlin, 1940: 15; male holotype from Tucson, Pima Co., Arizona, 32.221743 -110.9264796, elev. 2473ft., elev. ft., no collecting date, coll. unknown; deposited in AMNH. Paratype male from the Santa Catalina Mountains, Pima Co., Arizona, 32.315500 -110.7111685, elev. 3800ft., 8-12.vii.1916, coll. Dr. F.E. Lutz; deposited in AMNH. [examined]

Rhechostica apacheum Raven, 1985: 149.

Aphonopelma apacheum Smith, 1995: 73. syn. n.

Aphonopelma minchi Smith, 1995: 121; male holotype from Usery Pass Rd., near Usery Mountain Regional Park, Maricopa Co., Arizona, 33.482543 -111.6231784, elev. 2033ft., no collecting date, coll. A. Smith and M. Sullivan; deposited in BMNH. Paratype male from western end of Apache Trail, 33.444378 -111.5104915, elev. 1937ft., no collecting date, coll. A. Smith and M. Sullivan; deposited in BMNH. [examined] syn. n.

Aphonopelma schmidti Smith, 1995: 140; male holotype and female paratype from Mineral Mountain, near Florence Junction on Hwy 60, Pinal Co., Arizona, 33.265044 -111.3484275, elev. 1916ft., 10.viii.1992, coll. A. Smith and M. Sullivan; deposited in BMNH. [examined] syn. n.

Aphonopelma stahnkei Smith 1995: 146; male holotype and male paratype from Tempe, Maricopa Co., Arizona, 33.425510 -111.9400056, elev. 1176ft., no collecting date, coll. Dr. H.L. Stahnke; deposited in BMNH. [examined] syn. n.

Diagnosis

Aphonopelma chalcodes (Fig. 30) is a member of the Iodius species group and can be identified by a combination of morphological, molecular, and geographic characteristics. Nuclear DNA identifies A. chalcodes as a strongly supported lineage that is sister to the rest of the Iodius species group (A. eutylenum, A. iodius, and A. johnnycashi sp. n.) (Fig. 8). Aside from distribution, there are no pronounced measurements or characters that help discriminate A. chalcodes from the phenotypically similar species in the Iodius species group. Females and non-mature males of A. chalcodes can easily be differentiated from A. catalina sp. n., A. chiricahua sp. n., A. madera sp. n., A. marxi, and A. vorhiesi by possessing a more lightly colored carapace (the other species are generally black or dark brown). Males and females of A. chalcodes can easily be differentiated from syntopic members of the Paloma species group (mareki sp. n., paloma, parvum sp. n., prenticei sp. n., saguaro sp. n., and superstitionense sp. n.) by their greater extent of metatarsal scopulation on legs III and IV and their much larger size. The most significant measurements that distinguish male A. chalcodes from its syntopic species are M3 and extent of metatarsus IV scopulation. Male A. chalcodes can be distinguished by possessing a larger PTl/M3 (≥0.65; 0.65–0.76) than A. gabeli (≤0.64; 0.58–0.64); by possessing a larger L4 scopulation extent (42%–74%) than A. catalina (22%–36%), A. chiricahua (33%–40%), and A. vorhiesi (20%–36%); and by possessing a smaller F1/M3 (≤1.31; 1.13–1.31) than A. catalina (≥1.57; 1.57–1.68), A. chiricahua (≥1.40; 1.40–1.75), A. madera (≥1.51; 1.51–1.69), A. marxi (≥1.58; 1.58–1.87), and A. peloncillo sp. n. (≥1.33; 1.33–1.49). The most significant measurement that distinguishes female A. chalcodes from its syntopic species is extent of metatarsus IV scopulation. Female A. chalcodes can be distinguished by possessing a larger L4 scopulation extent (56%–81%) than A. catalina (37%–46%), A. chiricahua (27% ± (only 1 specimen)), A. gabeli (39%–53%), A. madera (29%–35%), A. marxi (33%–51%), A. peloncillo (32%-39%), and A. vorhiesi (29%–37%).

Figure 30.

Aphonopelma chalcodes Chamberlin, 1940 specimens, live photographs. Female (L) - APH_0697; Male (R) - APH_0600.

Redescription of male exemplar

(APH_0954; Fig. 31). Specimen preparation and condition: Specimen collected live crossing road, preserved in 80% ethanol; deposited in AUMNH; original coloration faded due to preservation. Left legs I, III, IV, and left pedipalp removed for measurements and photographs; stored in vial with specimen. Right leg III removed for DNA and stored at -80°C in the AUMNH (Auburn, AL). General coloration: Black and faded brown. Cephalothorax: Carapace 16.33 mm long, 15.71 mm wide; Hirsute; densely clothed with light brown iridescent pubescence mostly appressed to surface; fringe covered in long setae not closely appressed to surface; foveal groove medium deep and straight; pars cephalica region rises gradually from foveal groove, gently arching anteriorly toward ocular area; AER procurved, PER recurved; normal sized chelicerae; clypeus very slightly extends forward on a curve - mostly straight; LBl 2.15, LBw 2.76; sternum hirsute, clothed with shorter black/dark brown, densely packed setae. Abdomen: Densely clothed in short black/brown pubescence with numerous longer red/orange setae interspersed; possessing a dense dorsal patch of black Type I urticating bristles (Cooke et al. 1972). Legs: Hirsute, particularly ventrally; densely clothed in a mix of short/medium length black or faded black pubescence, femurs are darker. Metatarsus I slightly curved. F1 16.51; F1w 3.99; P1 7.12; T1 14.12; M1 13.37; A1 8.19; F3 13.73; F3w 4.15; P3 5.67; T3 10.59; M3 13.16; A3 7.95; F4 16.44; F4w 3.78; P4 6.16; T4 13.74; M4 16.84; A4 8.55; femur III is slightly swollen. All tarsi fully scopulate. Extent of metatarsal scopulation: leg III (SC3) = 81.3%; leg IV (SC4) = 76.4%. Three ventral spinose setae on metatarsus III; four ventral spinose setae on metatarsus IV. Coxa I: Prolateral surface a mix of fine, hair-like and tapered/thin tapered setae. Pedipalps: Hirsute; densely clothed in the same setal color as the other legs, with numerous longer ventral setae; one spinose seta on the apical, prolateral femur and three spinose setae on the prolateral tibia; PTl 9.54, PTw 3.15. When extended, embolus tapers and curves rapidly to the retrolateral side near apex; transition area from bulb to embolus is longer and wider than other species in the Iodius group, no keels.

Figure 31.

Aphonopelma chalcodes Chamberlin, 1940. A–I male specimen, APH_0954 A dorsal view of carapace, scale bar = 5mm B prolateral view of coxa I C dorsal view of femur III D ventral view of metatarsus III, scale bar = 4mm E ventral view of metatarsus IV, scale bar = 3.5mm F prolateral view of L pedipalp and palpal tibia, scale bar = 3mm G dorsal view of palpal bulb H retrolateral view of palpal bulb, scale bar = 1.5mm I prolateral view of tibia I (mating clasper), scale bar = 4.5mm.

Variation (13). Cl 14.43–21.07 (16.98±0.51), Cw 13.23–18.32 (15.675±0.45), LBl 1.44–2.6 (2.07±0.08), LBw 1.69–2.91 (2.401±0.08), F1 14.56–19.13 (16.687±0.38), F1w 3.65–5.05 (4.119±0.12), P1 6.0–8.11 (6.852±0.16), T1 13.29–15.65 (14.136±0.19), M1 12.08–15.77 (13.313±0.3), A1 7.49–9.02 (8.219±0.14), L1 length 53.99–67.39 (59.208±1.09), F3 12.86–15.58 (14.193±0.25), F3w 3.97–5.5 (4.369±0.13), P3 5.08–6.86 (5.769±0.15), T3 10.26–12.33 (11.244±0.21), M3 12.46–14.84 (13.615±0.24), A3 7.16–9.09 (8.096±0.15), L3 length 48.04–58.45 (52.917±0.92), F4 15.25–18.59 (16.683±0.3), F4w 3.53–5.09 (4.012±0.12), P4 5.09–7.09 (6.138±0.15), T4 12.11–15.53 (13.848±0.28), M4 14.9–20.1 (17.587±0.39), A4 8.03–10.05 (8.951±0.17), L4 length 56.11–70.85 (63.208±1.17), PTl 8.835–11.026 (9.601±0.19), PTw 2.79–3.48 (3.065±0.06), SC3 ratio 0.651–0.86 (0.773±0.02), SC4 ratio 0.428–0.764 (0.647±0.03), Coxa I setae = tapered, F3 condition = normal/slightly swollen.

Redescription of female exemplar

(APH_0887; Figs 3234). Specimen preparation and condition: Specimen collected live from burrow, preserved in 80% ethanol; deposited in AUMNH; original coloration faded due to preservation. Left legs I, III, IV, and pedipalp removed for photographs and measurements; stored in vial with specimen. Right leg III removed for DNA and stored at -80°C in the AUMNH (Auburn, AL). Genital plate with spermathecae removed and cleared, stored in vial with specimen. General coloration: Faded brown/black. Cephalothorax: Carapace 20.37 mm long, 18.10 mm wide; Hirsute, densely clothed with light brown pubescence closely appressed to surface; fringe densely covered in longer setae; foveal groove medium deep and straight; pars cephalica region gently rises from thoracic furrow, arching anteriorly toward ocular area; AER slightly procurved, PER slightly recurved; large chelicerae, clypeus mostly straight; LBl 2.45, LBw 3.01; sternum very hirsute, clothed with dark brown setae. Abdomen: Densely clothed dorsally in short black setae with numerous longer, lighter setae interspersed (generally red or orange in situ); dense dorsal patch of black Type I urticating bristles (Cooke et al. 1972); ventral side with shorter black/dark brown setae. Spermathecae: Paired and separate with wide bases, tapering and slightly curving medially towards capitate bulbs, with secondary medial bulges. Legs: Very hirsute, particularly ventrally; densely clothed in medium and long light brown pubescence, femurs much darker. F1 14.74; F1w 4.24; P1 6.40; T1 10.41; M1 8.93; A1 7.36; F3 12.95; F3w 4.27; P3 5.67; T3 9.23; M3 9.96; A3 7.32; F4 15.91; F4w 4.55; P4 6.66; T4 12.45; M4 13.79; A4 7.72. All tarsi fully scopulate. Extent of metatarsal scopulation: leg III (SC3) = 89.7%; leg IV (SC4) = 64.8%. Two ventral spinose setae on metatarsus III; six ventral spinose setae on metatarsus IV. Coxa I: Prolateral surface a mix of fine, hair-like and tapered/thin tapered setae. Pedipalps: Densely clothed in the same setal color as the other legs; one spinose seta on the apical, prolateral femur and six spinose setae on the prolateral tibia.

Figure 32.

Aphonopelma chalcodes Chamberlin, 1940. A–E female specimen, APH_0887 A dorsal view of carapace, scale bar = 8mm B prolateral view of coxa I C ventral view of metatarsus III, scale bar = 4.5mm D ventral view of metatarsus IV, scale bar = 5mm E prolateral view of L pedipalp and palpal tibia.

Figure 33.

Aphonopelma chalcodes Chamberlin, 1940. A–F cleared spermathecae. A chalcodes allotype B APH_0608 C APH_0887 D APH_1485 E AUMS_2605 F AUMS_3282.

Figure 34.

Aphonopelma chalcodes Chamberlin, 1940. A–D cleared spermathecae A APH_0168 B APH_0500 C AUMS_2339 D AUMS_2692.

Variation (10). Cl 13.49-21.79 (18.548±0.72), Cw 11.72-19.71 (16.589±0.68), LBl 1.90-2.61 (2.375±0.07), LBw 2.21-3.02 (2.773±0.1), F1 11.39-16.37 (14.684±0.42), F1w 3.34–4.76 (4.397±0.14), P1 5.03–7.26 (6.554±0.21), T1 9.03–12.42 (11.224±0.31), M1 6.81–11.81 (9.35±0.41), A1 5.68–7.39 (7.012±0.16), L1 length 37.94–55.02 (48.824±1.42), F3 9.38–13.96 (12.093±0.38), F3w 3.15–4.58 (4.028±0.13), P3 3.92–6.25 (5.575±0.21), T3 6.78–9.67 (8.581±0.25), M3 7.83–11.6 (9.925±0.34), A3 5.89–7.32 (7.035±0.14), L3 length 33.8–48.7 (43.209±1.2), F4 12.15–16.3 (14.699±0.36), F4w 3.11–4.73 (4.134±0.14), P4 4.25–6.98 (6.134±0.24), T4 9.59–12.45 (11.334±0.25), M4 11.16–15.35 (13.498±0.37), A4 6.51–8.28 (7.676±0.15), L4 length 43.66–57.92 (53.341±1.24), SC3 ratio 0.728–0.926 (0.843±0.02), SC4 ratio 0.566–0.812 (0.682±0.03), Coxa I setae = medium tapered. Spermathecae variation can be seen in Figures 3334.

Material examined

United States: Arizona: Cochise: 0.6 miles E of Portal, 31.913718 -109.130089 4, 4700ft., [AUMS_2676, 28/3/1990, 1♂, T.R. Prentice, AUMNH]; 1.4 miles NW Portal Rd on FR 42B (San Simon Rd); 31.926949 -109.169118 1, 5126ft., [APH_1229, 7/8/10, 1♂, Brent E. Hendrixson, Ashley Bailey, Andrea Reed, AUMNH]; 10 miles east of Dos Cabezas, 31.93051 -109.794753 5, 4281ft., [APH_2062, 4/8/71, 1♂, A. Jung, AMNH]; 2.4 miles NW Portal Rd on FR 42B (San Simon Rd); 31.932782 -109.183094 1, 5293ft., [APH_1230, 7/8/10, 1♂, Brent E. Hendrixson, Ashley Bailey, Andrea Reed, AUMNH]; 2.6 miles NW Airport Rd on Muleshoe Ranch Rd, 32.259698 -110.103465 1, 4967ft., [APH_0710, 19/7/2009, 1♂, Brent E. Hendrixson, Nate Davis, AUMNH]; 5 miles north of Douglas, 31.427339 -109.545202 5, 4163ft., [APH_2069, 18/8/1964, 2♂, W.J. Gertsch, AMNH]; 5 miles south of Apache, 31.631987 -109.093064 5, 4528ft., [APH_2074, 22/7/1964, 2♂, W.J. Gertsch and J.A. Woods, AMNH]; 5.1 miles NW AZ/NM state line on Portal Rd, 31.91264458 -109.1192604 2, 4620ft., [APH_0386, 31/7/2008, 1♂, Alice Abela, AUMNH]; Airport Rd, W of Wilcox, 32.23908 -109.975427 1, 4424ft., [APH_0709, 19/7/2009, 1♂, Brent E. Hendrixson, Nate Davis, AUMNH]; along Montezuma Canyon Rd, 31.385646 -110.406385 1, 5650ft., [APH_1485, 2/8/12, 1♀, Brent E. Hendrixson, Brendon Barnes, Austin Deskewies, AUMNH]; along Ramsey Canyon Rd, 31.460005 -110.295399 1, 5171ft., [APH_1225, 6/8/10, 1♂, Brent E. Hendrixson, Ashley Bailey, Andrea Reed, AUMNH]; Benson, 31.967008 -110.29502 5, 3589ft., [APH_2579, 8/1/54, 1♂, George Bredt, AMNH]; Cochise Cemetery (along Hwy-191); 32.093827 -109.910145 1, 4180ft., [APH_0619-621, 11/7/09, 2♀, 1 juv, Brent E. Hendrixson, Nate Davis, AUMNH]; Courtland Rd, near Jct. Ghost Town Trail, 31.758972 -109.796338 1, 4596ft., [APH_0715, 20/7/2009, 1♀, Brent E. Hendrixson, Nate Davis, AUMNH]; Oro Rd, just NE of Hwy-80, 31.402388 -109.773026 1, 4277ft., [APH_0686, 17/7/2009, 1 juv, Brent E. Hendrixson, Nate Davis, AUMNH]; Portal, 31.913703 -109.14145 5, 4770ft., [APH_2057, 22/7/1976, 1♀, Mark Russel, AMNH]; [APH_2058, 1/9/64, 1♂, W.J. Gertsch, AMNH]; [APH_2060, 7/1/67, 1♂, W.J. Gertsch, AMNH]; [APH_2066, 27/8/1970, 1♀, 1♂, Chamberlin, AMNH]; [APH_2067-2068, 10/7/68, 2♂, W.J. Gertsch, AMNH]; [APH_2078, 8/1/65, 1♀, W.J. Gertsch, AMNH]; Rucker Canyon Rd, 31.784784 -109.557742 1, 4575ft., [APH_0713-0714, 20/7/2009, 1♀, 1 juv, Brent E. Hendrixson, Nate Davis, AUMNH]; S of Sierra Vista at mouth of Ramsey Canyon, 31.463612 -110.291459 5, 5095ft., [AUMS_2396, 14/7/1993, 1♂, V. Roth, AUMNH]; Sulphur Canyon, near Portal , 31.892604 -109.167082 5, 5860ft., [APH_2059, 26/7/1955, 1♀, Guy Miller, AMNH]; Truman Rd, just off Hwy-82, 31.700137 -110.296948 1, 4242ft., [APH_0687-0690, 17/7/2009, 1♀, 3 juv, Brent E. Hendrixson, Nate Davis, AUMNH]; Coconino: Flagstaff, 35.200524 -111.639452 6, 6929ft., [APH_2586, unknown, 1♂, unknown, AMNH]; Mayer, just S of Prescott, 34.395374 -112.237457 5, 4488ft., [AUMS_2390, 6/20/09, 1♂, Tom Roberts, AUMNH]; Schnebly Hill Road NE of Sedona, 34.887682 -111.703022 5, 5810ft., [APH_2497, 6/8/77, 1♂, Barbara and M.W. Sanderson, AMNH]; [APH_2483, 6/8/77, 1♂, Barbara and M.W. Sanderson, AMNH]; Gila: 0.55 miles N Strawberry on Hwy-87, 34.41409752 -111.4912396 4, 6046ft., [APH_0388, 2/8/08, 1♂, Alice Abela, AUMNH]; 0.75 miles E AZ-87 on Control Rd, 34.360317 -111.41347 1, 5555ft., [APH_0505-0508, 20/5/2009, 2♀, 2 juv, Brent E. Hendrixson, Bernadette DeRussy, Sloan Click, AUMNH]; 1.1 miles NW AZ-87 on Barnhardt Trail Rd (FR 419); 34.09815 -111.36165 2, 3200ft., [APH_0344, 10/5/08, 1 juv, Brent E. Hendrixson, Zach Valois, Josh Richards, AUMNH]; 1.4 miles E US-87 on Conrol Rd, 34.35949 -111.40302 2, 5430ft., [APH_0346, 10/5/08, 1 juv, Brent E. Hendrixson, Zach Valois, Josh Richards, AUMNH]; 10 miles NW of Globe, 33.527502 -110.694934 5, 4337ft., [APH_2064, 10/9/62, 1♂, Roth and Roth, AMNH]; 8 miles S of Young-dirt road Hwy 288, 33.997859 -110.953748 5, 6170ft., [AUMS_2319, 6/8/92, 1♂, T.R. Prentice, AUMNH]; along Control Rd, east of Hwy-87, 34.3602 -111.415521 1, 5855ft., [APH_1531-1532, 4/10/12, 1♀, 1 juv, Brent E. Hendrixson, AUMNH]; Along Houston-Mesa Rd, 34.29263192 -111.2868479 4, 5168ft., [APH_0387, 2/8/08, 1♂, Alice Abela, AUMNH]; Apache Trail, 33.532562 -110.92045 6, 3711ft., [APH_2456, 25/3/1936, 1♂, Edith M. Patel, AMNH]; Mazatzal Mountains, Tonto National Forest, El Oso Rd, 33.74139 -111.35831 1, 5900ft., [APH_0191, 6/9/07, 1♀, Lorenzo Prendini, Jeremy Huff, AUMNH]; Payson, .41 miles E of Nf-200, 34.203644 -110.980578 1, 5297ft., [APH_3127, 9/8/13, 1♂, Tyler P. McKee, AUMNH]; Salt River Canyon, 33.619952 -110.921017 5, 2503ft., [APH_2065, 6/1/56, 1♂, W.D. Allison, AMNH]; Tonto National Forest, near jct. FR-449 off Hwy-188, 33.61465 -111.034975 1, 2474ft., [APH_1301-1303, 26/7/2011, 3 juv, Brent E. Hendrixson, Brendon Barnes, Nate Davis, Jake Storms, AUMNH]; Graham: 11.2 miles W Hwy-191 on Hwy-266, 32.582263 -109.852113 1, 5592ft., [APH_1235, 8/8/10, 1♀, Brent E. Hendrixson, Ashley Bailey, Andrea Reed, AUMNH]; 3.6 miles SW Hwy-191 along Swift Trail (Hwy-366); 32.6896 -109.754556 1, 3950ft., [APH_1509, 7/9/12, 1♂, Brent E. Hendrixson, AUMNH]; along Ash Creek Rd, 32.51432 -110.077472 1, 4450ft., [APH_1234, 8/8/10, 1♂, Brent E. Hendrixson, Ashley Bailey, Andrea Reed, AUMNH]; along High Creek Rd, 32.570316 -110.048141 1, 4557ft., [APH_1232, 8/8/10, 1 juv, Brent E. Hendrixson, Ashley Bailey, Andrea Reed, AUMNH]; along Sunset Loop Rd, 32.512166 -110.159031 1, 4747ft., [APH_1233, 8/8/10, 1♂, Brent E. Hendrixson, Ashley Bailey, Andrea Reed, AUMNH]; Bonita-Klondyke Rd, W of Bonita, 32.600171 -109.983213 1, 4571ft., [APH_0695-0697, 18/7/2009, 3♀, Brent E. Hendrixson, Nate Davis, AUMNH]; crossing Sunset Rd, 32.506667 -110.180056 1, 4711ft., [APH_0391, 24/7/2008, 1♂, Kari, Hunter McWest, AUMNH]; Lower Canyon of Mount Graham, 32.64023 -109.931671 6, 5281ft., [APH_2071, 15/7/1956, 1♂, V. Roth and W.J. Gertsch, AMNH]; [APH_2583, 15/7/1956, 1♂, W. Gertsch and V. Roth, AMNH]; W Ash Creek Rd, 32.513833 -110.05858 1, 4420ft., [APH_0392, 24/7/2008, 1♂, Kari, Hunter McWest, AUMNH]; Greenlee: 14 miles west of Clifton, 33.06165 -109.44632 5, 3921ft., [APH_2493, 28/7/1962, 2♂, G. Bradt, AMNH]; Groom: Bradshaw Mountains near Prescott in pine forest, 34.385392 -112.368863 6, 6440ft., [APH_2486, 15/9/1964, 1♂, Lowe, Soule, Wright, AMNH]; La Paz: 6.9 miles N US-60 on Alamo Rd (near Base of Harcuvar Mtns N of Wenden); 33.92479 -113.54419 1, 2285ft., [APH_0340-0341, 9/5/08, 1♀, 1 juv, Brent E. Hendrixson, Zach Valois, AUMNH]; Alano Road, 34.08542 -113.5515 4, 1982ft., [APH_0038, 12/8/06, 1♂, Alice Abela, AUMNH]; Bill Williams River NWA, E of Parker Dam (Buckskin Mtns); 34.29172 -114.08915 1, 560ft., [APH_1003, 8/5/10, 1♀, Chris A. Hamilton, AUMNH]; S of Hwy-72 on Bouse-Quartzsite Rd, 33.86255 -114.02868 1, 1130ft., [APH_0450, 17/11/2008, 1♀, June Olberding, AUMNH]; [APH_0451, 30/11/2008, 1♂, June Olberding, AUMNH]; Maricopa: Palomas Plain, 2.5 miles N of Sentinel, 32.886722 -113.238144 5, 667ft., [AUMS_2604, 11/8/90, 1♂, T.R. Prentice, AUMNH]; Palomas Plain, 3 miles N of Sentinel, 32.90163 -113.212893 5, 662ft., [AUMS_2647, 22/11/1992, 1♂, unknown, AUMNH]; Rio Verde, 1 mile S of Forest Rd and Rio Verde Dr. jct, off Forest Rd, 33.727441 -111.670456 4, 1586ft., [AUMS_2673, 13/5/1991, 1♀, Chris Baptista, AUMNH]; Sentinel, 1.5 miles N of I-8, 32.879995 -113.223031 5, 663ft., [AUMS_2620, 7/1/92, 1♂, unknown, AUMNH]; 1.35 miles N McDowell Rd on Usery Pass Rd, 33.480479 -111.624831 1, 1989ft., [APH_1497, 5/9/12, 1 juv, Brent E. Hendrixson, AUMNH]; 3.0 miles N I-8 (Sentinel) on Agua Caliente Rd, 32.890757 -113.24223 1, 656ft., [APH_0428, 16/11/2008, 1 juv, Brent E. Hendrixson, AUMNH]; 4.65 miles N I-8 (Sentinel) on Agua Caliente Rd, 32.90721 -113.262703 1, 592ft., [APH_0429, 16/11/2008, 1 juv, Brent E. Hendrixson, AUMNH]; about 9-10 miles E AZ-87 on NF-143, 33.7285 -111.43288 4, 3719ft., [APH_0134, 15/6/2007, 1♀, Austin Spears, AUMNH]; [APH_0135, 15/6/2007, 1 juv, Gilbert Quintana, AUMNH]; along Apache Trail, 33.606639 -111.19778 1, 2372ft., [APH_1492-1496, 4/9/12, 4♂, 1 juv, Brent E. Hendrixson, AUMNH]; [APH_1502-1505, 6/9/12, 2♀, 2♂, Brent E. Hendrixson, AUMNH]; east of Tortilla Flat off of Hwy 88, 33.5336 -111.37225 1, 1376ft., [APH_0954, 6/8/14, 1♂, Ben Allen, AUMNH]; in the Tonto National Forest, off Road 78 that splits from Apache Trail (88); E of Apache Junction, 33.4701 -111.47296 1, 2082ft., [APH_3199, 15/11/2013, 1 juv, Chris A. Hamilton, Brent E. Hendrixson, AUMNH]; Mazatzal Mountains, Four Peaks, Cline Cabin Rd./F.R. 143, 33.71667 -111.45509 4, 3080ft., [APH_0298, 6/10/07, 1 juv, Zach Valois, AUMNH]; Mesa, 33.414255 -111.831101 5, 1220ft., [APH_2591, unknown, 1♂, Michael Soleglad, AMNH]; [APH_2595, unknown, 1♂, Michael Soleglad, AMNH]; N of Gila Bend along Hwy-85, 0.6 miles E of Hwy, 33.043715 -112.633476 1, 807ft., [APH_0419, 15/11/2008, 1 juv, Brent E. Hendrixson, AUMNH]; N side of Cline Cabin Rd, 33.72875 -111.43244 4, 3750ft., [APH_0360, 17/4/2008, 1♀, Zach Valois, AUMNH]; Palomas Plain, Sentinel, 32.861687 -113.21082 5, 683ft., [AUMS_2323, 18/7/1989, 1♂, T.R. Prentice, AUMNH]; Phoenix, Usery Pass Rd off N Bush Hwy, 33.5468 -111.645 1, 1346ft., [APH_0887, 6/8/14, 1♀, Ben Allen, AUMNH]; Phoenix, 33.460362 -112.039455 6, 1119ft., [APH_2075, unknown, 1♀, unknown, AMNH]; South Mountain Park, 33.35257 -112.072563 6, 1332ft., [APH_2501, 21/11/1965, 1♂, S. C. Williams, AMNH]; South of Geronimo Head Mountain, 33.498275 -111.387253 6, 2933ft., [APH_2070, 21/12/1963, 1♂, Chamberlin, AMNH]; Sunflower, Sycamore Creek Rd., 33.88512 -111.47874 4, 3526ft., [APH_0300-301, 6/10/07, 2 juv, Zach Valois, AUMNH]; E/NE of Pyramid Peak, E of New River Rd., 33.921335 -112.101997 5, 2184ft., [AUMS_2631, 18/11/2003, 1♂, C. Shipley, AUMNH]; New River, 33.915059 -112.136776 5, 2022ft., [AUMS_2687, 8/1/98, 1♂, Tom Roberts, AUMNH]; 6 miles N AZ-74 on New River Rd (near Lake Pleasant); 33.86502 -112.18967 1, 1799ft., [APH_0007, 5/10/01, 1♂, Brent E. Hendrixson, Darrin Vernier, AUMNH]; [APH_0046, 5/10/01, 1 juv, Brent E. Hendrixson, Darrin Vernier, AUMNH]; 6.0 miles N AZ-74 on New River Rd (near Lake Pleasant); 33.86307 -112.19106 1, 1773ft., [APH_0127-0130, 13/6/2007, 1♀, 1♂, 2 juv, Brent E. Hendrixson, AUMNH]; [APH_0142-0143, 13/6/2007, 1♀, 1 juv, Brent E. Hendrixson, AUMNH]; 6.6 miles W jct. US-93 on US-60, 33.956369 -112.840469 1, 2582ft., [APH_0498-0501, 18/5/2009, 1♀, 3 juv, Brent E. Hendrixson, Bernadette DeRussy, Sloan Click, AUMNH]; New River, ~0.5 miles W I-17 on AZ-74, 33.79765 -112.14307 4, 1655ft., [APH_0164-0165, unknown, 1♂, 1♀, Brandon Anderson, AUMNH]; Wickenburg, about 2.84 miles south of Interstate 60, 33.922158 -112.903511 1, 2700ft., [APH_3128, 1/9/13, 1♀, Tyler P. McKee, AUMNH]; Maricopa/Yavapai: between Prescott and Phoenix, 34.052118 -112.147482 7, 2047ft., [APH_2076, 31/12/1933, 1♂, McCanly, AMNH]; Mohave: 0.9 miles S I-40 on Hwy-93, 35.14879 -113.689479 1, 3646ft., [APH_1221-1222, 2/8/10, 2 juv, Brent E. Hendrixson, Brendon Barnes, Nate Davis, AUMNH]; 7.4 miles NE Hwy-93 on CR-25 in Dolan Springs, 35.60418 -114.253563 1, 3638ft., [APH_1309, 28/7/2011, 1 juv, Brent E. Hendrixson, Brendon Barnes, Nate Davis, Jake Storms, AUMNH]; Bullhead City, ~2 miles S Hwy-68, 35.16973 -114.53414 4, 1003ft., [APH_0048, 22/8/2001, 1♀, Tom and Tracy Vezie, AUMNH]; Cool Spring on Oatman Rd, off Rt. 66 (Black Mtns); 35.02957 -114.31629 1, 2772ft., [APH_0999-1001, 6/5/10, 3♀, Chris A. Hamilton, Rick West, AUMNH]; just N I-40 on Silver Springs Rd, 35.161716 -113.564676 1, 3927ft., [APH_1305, 28/7/2011, 1♂, Brent E. Hendrixson, Brendon Barnes, Nate Davis, Jake Storms, AUMNH]; NW of Oatman on Rt. 10, 35.07184 -114.43861 1, 2191ft., [APH_0310-0312, 11/10/07, 3♂, Rick C. West, AUMNH]; Oatman Rd., Black Mountains, 35.009671 -114.389097 6, 2420ft., [AUMS_2682, 27/3/1988, 1♀, T.R. Prentice, AUMNH]; Oatman, N of town off Rt. 66 (Black Mtns); 35.03484 -114.39156 1, 2772ft., [APH_1002, 7/5/10, 1♀, Chris A. Hamilton, AUMNH]; Navajo: Hwy 60 (77); N of Carrizo, mile marker 324, 34.070957 -110.193557 5, 5609ft., [AUMS_2477, 31/8/1986, 1♂, T.R. Prentice, AUMNH]; Pima: 0.18 miles S Hwy-86 on Hwy-386, 32.02419162 -111.5769608 2, 3251ft., [APH_0739, 23/8/2009, 1♂, Alice Abela, AUMNH]; 0.2 miles W SR-83 on Sahuarita Rd, 31.963159 -110.675585 1, 3679ft., [APH_0616, 9/7/09, 1♂, Brent E. Hendrixson, Jon Davenport, Nate Davis, AUMNH]; 0.75 miles S Ball Rd in Why along Hwy-85, 32.252063 -112.7449 1, 1787ft., [APH_0425, 15/11/2008, 1 juv, Brent E. Hendrixson, AUMNH]; 1.02 miles S Hwy-86 on Hwy-386, 32.01220214 -111.5747721 2, 3372ft., [APH_0738, 23/8/2009, 1♂, Alice Abela, AUMNH]; 1.75 miles N Sahuarita Rd on Houghton Rd, 31.987937 -110.772369 1, 3201ft., [APH_0617, 9/7/09, 1♂, Brent E. Hendrixson, Jon Davenport, Nate Davis, AUMNH]; 10560 E Wildfire Drive, Tucson, 32.2213 -110.76858 2, 2757ft., [APH_0014, 8/2/14, 1♂, Alex Binford, AUMNH]; 2 miles N AZ-86 (W Ajo Hwy) on Sandario Rd, near jct. with Dusty Mesquite Trail, 32.14446246 -111.2177859 2, 2350ft., [APH_0370, 24/7/2008, 1♂, Alice Abela, AUMNH]; 5 miles west on St.289 from Jef.289 and US 89- north of Nogales, 31.426474 -111.008278 5, 3665ft., [APH_2526, 20/7/1966, 1♂, E. Brown and J. Cole, AMNH]; along Hwy-86, 32.1992 -112.43633 5, 1980ft., [APH_0174, 8/7/14, 1♂, Alice Abela, AUMNH]; [APH_0176, 8/7/14, 1♂, Alice Abela, AUMNH]; [APH_0740-0742, 23/8/2009, 3♂, Alice Abela, AUMNH]; along Hwy-86, W of San Pedro, 32.03889549 -111.5014244 4, 2943ft., [APH_0737, 22/8/2009, 1♂, Alice Abela, AUMNH]; along Kitt Peak Rd, 31.97428845 -111.604144 4, 4625ft., [APH_0371-0372, 24/7/2008, 2♂, Alice Abela, AUMNH]; [APH_0374-0375, 25/7/2008, 2♂, Alice Abela, AUMNH]; along Madera Canyon Rd, 31.836971 -110.946796 1, 3083ft., [APH_1344-1345, 4/8/11, 2♀, Brent E. Hendrixson, Brendon Barnes, Nate Davis, Jake Storms, AUMNH]; along SR-83, 31.896859 -110.665638 1, 4045ft., [APH_0611-0615, 9/7/09, 5♂, Brent E. Hendrixson, Jon Davenport, Nate Davis, AUMNH]; AZ-86, 0.06 miles SW jct Indian Rt 30, 31.99315318 -111.7053503 2, 2924ft., [APH_0373, 25/7/2008, 1♂, Alice Abela, AUMNH]; Batamote Rd, 31.729183 -111.161558 1, 3387ft., [APH_0610, 9/7/09, 1 juv, Brent E. Hendrixson, Jon Davenport, Nate Davis, AUMNH]; Bates Well Rd., 2 miles N of OPCN monument (Organ Pipe Cactus National Monument); 32.226852 -112.890815 5, 1545ft., [AUMS_3329, 10/9/90, 1♂, W. Icenogle and G. Lowe, AUMNH]; Catalina Mountains, Bug Spring Trail, 32.341135 -110.714921 1, 3624ft., [APH_0603-0604, 9/7/09, 1♀, 1 juv, Brent E. Hendrixson, Jon Davenport, Nate Davis, Paul Marek, Charity Hall, AUMNH]; Catalina State Park, campground, 32.4238 -110.9233 2, 2700ft., [APH_1487, 2/8/12, 1♂, Brent E. Hendrixson, Brendon Barnes, Austin Deskewies, AUMNH]; Catalina State Park, trailhead parking lot and nature trail, 32.425002 -110.909336 1, 2635ft., [APH_0598-0600, 8/7/09, 1♀, 2♂, Brent E. Hendrixson, Jon Davenport, Nate Davis, AUMNH]; [APH_0602, 8/7/09, 1♂, Brent E. Hendrixson, Jon Davenport, Nate Davis, AUMNH]; Fish Canyon Rd, 31.742778 -110.707871 1, 4888ft., [APH_0691-0693, 17/7/2009, 1♀, 2 juv, Brent E. Hendrixson, Nate Davis, AUMNH]; NFWL Campsite, Cabeza Prieta Game Refuge, 32.232313 -113.608052 5, 932ft., [APH_2585, 7/1/76, 1♀, B. Woodward and R. Jones, AMNH]; Organ Pipe National Monument, 32.016369 -112.858922 5, 1988ft., [APH_2498, 5/22/09, 1♂, Vogt, AMNH]; Quail Creek-Veterans Municipal Park, 31.898272 -110.964712 1, 2794ft., [APH_0605-0609, 9/7/09, 5♀, Brent E. Hendrixson, Jon Davenport, Nate Davis, AUMNH]; Redington, 32.42157 -110.5037 5, 2890ft., [APH_2072, 25/5/1970, 1♀, Mary Smallhouse, AMNH]; Sabino Canyon, Tucson, 32.315852 -110.81935 5, 2782ft., [APH_2474, 11/9/35, 1♂, W.J. Baerg, AMNH]; [APH_2588, 9/1/35, 1♂, unknown, AMNH]; Saguaro National Park, eastern portion of park, off Cactus Forest Loop Dr. at Cactus Forest south hiking trail, 32.171204 -110.730429 1, 3061ft., [APH_3174-3175, 12/11/13, 1♂, 1 juv, Chris A. Hamilton, Brent E. Hendrixson, AUMNH]; San Vicente, 33.336256 -111.831893 5, 1220ft., [APH_2061, 19/8/1957, 2♂, V. Roth, AMNH]; Santa Catalina Mountains, 32.44193 -110.860732 6, 3806ft., [APH_2073, 8/7/16, 1♂, T.E. Luty, AMNH]; Santa Catalina Mountains, Catalina Hwy, 32.337262 -110.6959 5, 4501ft., [AUMS_3282, 26/8/1989, 1♂, T.R. Prentice, AUMNH]; SE of Ajo, mile marker 46, 32.357132 -112.826569 5, 1710ft., [AUMS_2605, 27/8/1989, 1♀, T.R. Prentice, AUMNH]; SW of Ajo, Little Ajo Mountain, Darby Well Road, 32.35451 -112.886495 4, 2090ft., [APH_0309, 15/10/2005, 1♂, Lennart Pettersson, Dave Kandiyeli, Sheri Monk, AUMNH]; Tucson, 32.221743 -110.926479 6, 2470ft., [APH_2088, 27/7/1936, 1♀, 1♂, unknown, AMNH]; [APH_2089, 7/1/35, 1♀, P. Stichler, AMNH]; [APH_2063, 23/8/1973, 1♂, A. Jung, AMNH]; [APH_2205, 9/1/35, 1♂, W.J. Baerg, AMNH]; Tucson area, near Catalina State Park, N side of Tangerine Rd and E of Tangerine Crossing, E of Marana, 32.424545 -111.03393 1, 2731ft., [APH_3173, 12/11/13, 1 juv, Brent E. Hendrixson, Chris A. Hamilton, AUMNH]; Ajo, SE on Hwy 85, 32.35653 -112.826776 5, 1711ft., [AUMS_2384, 27/8/1989, 1♀, T.R. Prentice, AUMNH]; near Sahuarita off I19 (Hwy 89); 32.002043 -110.981937 5, 2702ft., [AUMS_2324, 15/10/1992, 1♂, Barney Tomberlin, AUMNH]; Papago Indian Reservation, Hwy 86, 7.2 miles W of Indian Rte 15, 32.176101 -112.232872 4, 2418ft., [AUMS_2382, 18/11/1989, 1♂, T.R. Prentice, AUMNH]; Pinal: 0.1 miles S Apache Trail on Mountain View Rd, 33.445508 -111.502834 1, 1935ft., [APH_0503, 19/5/2009, 1 juv, Brent E. Hendrixson, Bernadette DeRussy, Sloan Click, AUMNH]; 1.5 miles S US-60 on Mineral Mountain Rd, 33.242909 -111.270678 1, 2067ft., [APH_0502, 19/5/2009, 1 juv, Brent E. Hendrixson, Bernadette DeRussy, Sloan Click, AUMNH]; 2.4 miles E Hwy-79 on Cottonwood Canyon Rd, 33.187604 -111.313416 1, 1901ft., [APH_1304, 26/7/2011, 1 juv, Brent E. Hendrixson, Brendon Barnes, Nate Davis, Jake Storms, AUMNH]; 3.4 miles SE AZ-77 on SR-76, near San Manuel, 32.628709 -110.648951 1, 3392ft., [APH_0596, 8/7/09, 1♀, Brent E. Hendrixson, Jon Davenport, Nate Davis, AUMNH]; 7 miles south of Oracle-Santa Catalina Mountains, 32.531565 -110.773332 5, 4678ft., [APH_2529, 25/7/1949, 2♂, unknown, AMNH]; in desert NW jct of Hwy-8 4and Amarillo Valley Rd (3.2 miles NW I-8); 32.856009 -112.085251 1, 1517ft., [APH_0420, 15/11/2008, 1♀, Brent E. Hendrixson, Paul Marek, Charity Hall, Kojun, AUMNH]; near Cogburn Ostrich Ranch, 32.64396 -111.39235 4, 1840ft., [APH_0369, 5/4/08, 1 juv, Shasta Michaels, AUMNH]; off Florence-Kelvin Hwy, BLM land E of Florence, 32.99965 -111.26476 1, 1970ft., [APH_3196, 15/11/2013, 1 juv, Chris A. Hamilton, Brent E. Hendrixson, AUMNH]; off Hwy 79 around 1/2 way between 77 split and Florence at Tom Mix Rest Area, 32.82084 -111.20621 1, 2333ft., [APH_3192, 15/11/2013, 1 juv, Chris A. Hamilton, Brent E. Hendrixson, AUMNH]; off Hwy 79 around 1/2 way between 77 split and Florence at Tom Mix Rest Area, 32.820504 -111.20632 1, 2370ft., [APH_3193, 15/11/2013, 1 juv, Brent E. Hendrixson, Chris A. Hamilton, AUMNH]; OHV trail area, near Kearny, 33.065521 -110.888678 1, 2163ft., [APH_0595, 8/7/09, 1♀, Brent E. Hendrixson, Jon Davenport, Nate Davis, AUMNH]; Oracle, 32.610602 -110.771115 5, 4537ft., [APH_2593, 11/7/40, 1♂, Gertsch and Hook, AMNH]; Picacho, just off Camino Adelante Dr, 32.71453 -111.4975 2, 1612ft., [APH_0716, 7/9/14, 1♀, Warren Burke, AUMNH]; Robert Engstrom’s property, 0.2 miles S East Cactus Forest Rd on North Coolidge Airport Rd, 32.97029 -111.42175 2, 1541ft., [APH_0198, 14/6/2007, 1♂, Robert Engstrom, AUMNH]; Superior, Boyce Thompson Arboretum, 33.279265 -111.160952 2, 2440ft., [APH_1370, 3/9/11, 1♂, Benjamin Curtis, Mason McWest, AUMNH]; W of Chuck’s Corner off S. Amarillo Valley Rd, off Hwy 84, N of I-8, 32.8572 -112.08641 1, 1517ft., [APH_3167, 11/11/13, 1♂, Chris A. Hamilton, Brent E. Hendrixson, AUMNH]; [APH_3169, 11/11/13, 1 juv, Brent E. Hendrixson, Chris A. Hamilton, AUMNH]; Santa Cruz: 0.65 miles W Mount Hopkins Rd on Forest Service Rd 184, 31.69646 -111.00164 2, 3512ft., [APH_0175, 8/7/14, 1♂, Alice Abela, AUMNH]; 0.95 miles W Mount Hopkins Rd on Forest Service Rd 184, 31.69671 -111.00641 2, 3497ft., [APH_0173, 8/7/14, 1♂, Alice Abela, AUMNH]; along Duquesne Rd, 31.363711 -110.792456 1, 4146ft., [APH_1340, 4/8/11, 1♀, Brent E. Hendrixson, Brendon Barnes, Nate Davis, Jake Storms, AUMNH]; along Duquesne Rd (FR-61]; 31.347952 -110.509098 1, 5057ft., [APH_1339, 4/8/11, 1♀, Brent E. Hendrixson, Brendon Barnes, Nate Davis, Jake Storms, AUMNH]; along FR-61, 31.34832 -110.508551 1, 5088ft., [APH_1224, 6/8/10, 1♀, Brent E. Hendrixson, Ashley Bailey, Andrea Reed, AUMNH]; along Montosa Canyon Rd (FR-184); 31.701063 -111.039066 1, 3181ft., [APH_1341, 4/8/11, 1♀, Brent E. Hendrixson, Brendon Barnes, Nate Davis, Jake Storms, AUMNH]; Along Mt Hopkins Rd, 31.67354048 -110.9229983 4, 5047ft., [APH_0383-0384, 28/7/2008, 2♂, Alice Abela, AUMNH]; [APH_0730, 20/8/2009, 1♂, Alice Abela, AUMNH]; Along Ruby Rd, 31.43198526 -111.188291 4, 4018ft., [APH_0376, 26/7/2008, 1♂, Alice Abela, AUMNH]; [APH_0377-0379, 27/7/2008, 3♂, Alice Abela, AUMNH]; [APH_0380-0381, 28/7/2008, 2♂, Alice Abela, AUMNH]; Coronado National Forest, along FR-61, 31.343749 -110.485365 1, 5083ft., [APH_1195-1196, 28/7/2010, 2♀, Brent E. Hendrixson, Brendon Barnes, Nate Davis, AUMNH]; Coronado National Forest, Pena Blanca Lake, Hwy 289, 31.39225 -111.13153 1, 4290ft., [APH_0193, 5/9/07, 1♂, Lorenzo Prendini, Jeremy Huff, AUMNH]; Harshaw Rd, 31.531637 -110.718517 4, 4187ft., [APH_0389, unknown, 1♂, Alice Abela, AUMNH]; [APH_0694, 17/7/2009, 1 juv, Brent E. Hendrixson, Nate Davis, AUMNH]; Oro Blanco Mountains-12 miles from Nogales, 31.333796 -111.023962 5, 4472ft., [APH_2492, 7/1/37, 3♂, Steckler, AMNH]; Ruby Road, 31.3948 -111.13614 4, 4487ft., [APH_0037, 6/8/06, 1♂, Alice Abela, AUMNH]; Ruby Road near Pena Lake, 31.400702 -111.090398 5, 3871ft., [APH_2487, 19/7/1962, 1♂, unknown, AMNH]; Ruby Road-10 miles northwest of Nogales, 31.402097 -111.027724 5, 3865ft., [APH_2505, 8/1/62, 1♂, W.J. Gerstch, AMNH]; Yavapai: 0.2 miles E I-17 on Dugas Rd, 34.39945 -112.07044 1, 3800ft., [APH_0131-0132, 13/6/2007, 1♂, 1 juv, Brent E. Hendrixson, AUMNH]; 0.8 miles E I-17, 0.9 miles S Stoneman Lake Road [near Exit 306]; 34.76055 -111.64488 1, 5472ft., [APH_0049, 3/10/01, 1 juv, Brent E. Hendrixson, Darrin Vernier, AUMNH]; 3.4 miles SE AZ-71 on US-93, 34.09027 -112.90285 1, 2717ft., [APH_0133, 15/6/2007, 1 juv, Brent E. Hendrixson, Shasta Michaels, AUMNH]; Beaver Creek Trail at trailhead, 40.4 miles S of jct I-40 and I-17 on 17, 34.674376 -111.713738 5, 3875ft., [AUMS_2339, 31/3/1990, 1♂, T.R. Prentice, AUMNH]; Beaver Creek, E of I-17, E of interstate, N of Hwy 260, 34.671522 -111.724763 5, 3920ft., [AUMS_3264, 31/3/1990, 1♀, T.R. Prentice, AUMNH]; Black Canyon City, 0.85 miles E I-17 on South Mud Springs Rd, 34.073 -112.13204 4, 2044ft., [APH_0168-0170, unknown, 1♀, 2♂, Brandon Anderson, AUMNH]; Bumble Bee Ranch Rd. (Bumble Bee, AZ Ghost Town off Co. Rd. 59); 34.152931 -112.156127 5, 2640ft., [AUMS_2325, 10/8/90, 1♂, T.R. Prentice, AUMNH]; Bumblebee Ranch Rd (Crown King Rd); 1.5 miles E of Crown King, 34.217448 -112.307279 5, 5477ft., [AUMS_2692, 30/8/1990, 2♂, T.R. Prentice, AUMNH]; Bumblebee Ranch Road, 34.227934 -112.154007 5, 2800ft., [AUMS_2383, 30/8/1990, 1♂, T.R. Prentice, AUMNH]; [AUMS_2320, 11/8/90, 1♂, T.R. Prentice, AUMNH]; [AUMS_2338, 12/8/90, 1♀, T.R. Prentice, AUMNH]; [AUMS_3272, 16/8/1994, 1♀, T.R. Prentice, AUMNH]; [AUMS_3324, 11/8/90, 1♂, T.R. Prentice, AUMNH]; [AUMS_3339, 10/8/90, 1♂, T.R. Prentice, AUMNH]; [AUMS_2379, 16/8/1994, 1♂, Enza Prentice, AUMNH]; Constellation, 34.064896 -112.585608 5, 3446ft., [AUMS_2366, 8/1/91, 1♂, T.R. Prentice, AUMNH]; Cordes Lake Development, Cordes jct, SE of Mayer, 34.307807 -112.103491 5, 3701ft., [AUMS_3342, 21/8/1998, 1♂, Stephen Roy, AUMNH]; Dugas Road, approx. 2.0 miles W I-17 on Orme Ranch Road, 34.42758 -112.07979 4, 3914ft., [APH_0013, 24/3/2002, 1♂, John Bell, AUMNH]; N of Village of Oak Creek, Courthouse Butte/Big Park Trail, 34.79569 -111.758817 1, 4233ft., [APH_1543, 22/10/2012, 1 juv, Brent E. Hendrixson, AUMNH]; near Deer Pass Crossing on Oak Creek about 8 miles southeast Sedona, 34.838954 -111.721245 5, 6411ft., [APH_2485, 15/6/1977, 1♂, M.W. Sanderson, AMNH]; Prescott, 34.53923 -112.468695 5, 5387ft., [APH_2504, 28/7/1948, 1♂, C. and P. Vaurie, AMNH]; Yarnell, 34.221331 -112.747488 5, 4806ft., [APH_2587, 4/9/61, 1♂, Roth and Roth, AMNH]; Yuma: Rd. 36 E off I-8, S side of I-8, E field, 32.679615 -114.015636 5, 358ft., [AUMS_2362, 12/8/90, 1♂, T.R. Prentice, AUMNH]; Rd. 36 E, south, W side of rd., 32.645663 -114.02637 5, 449ft., [AUMS_2363, 16/11/1986, 1♂, T.R. Prentice, AUMNH]; New Mexico: Hidalgo: Rodeo, 31.835372 -109.03117 5, 4131ft., [APH_2090, 20/1/1969, 1♀, V. Roth, AMNH].

Distribution and natural history

Aphonopelma chalcodes is widely distributed across the southern two-thirds of Arizona south of the Grand Canyon, bound to the west by the Colorado River and barely making its way into southwestern New Mexico (Fig. 35). The species is undoubtedly widespread throughout northern Sonora, Mexico as well. This species inhabits a large diversity of habitats including the following Level III Ecoregions: Arizona/New Mexico Mountains, Arizona/New Mexico Plateau, Madrean Archipelago, and the Sonoran Basin and Range (Fig. 1G). Aphonopelma chalcodes can be found in syntopy with a number of species across its distribution including: A. catalina , A. chiricahua , A. gabeli, A. madera , A. mareki , A. marxi, A. paloma, A. parvum , A. peloncillo, A. prenticei , A. saguaro , A. superstitionense , and A. vorhiesi. The breeding season, when mature males abandon their burrows in search of females, occurs during the summer (generally July–September), particularly during the summer monsoon season. Additional information about the natural history of this species can be found in Smith (1995).

Figure 35.

Aphonopelma chalcodes Chamberlin, 1940. A distribution of known specimens B predicted distribution; warmer colors (red, orange, yellow) represent areas of high probability of occurrence, cooler colors (blue shades) represent areas of low probability of occurrence.

Conservation status

Aphonopelma chalcodes is the most widespread and abundant tarantula species in Arizona. The species is secure.

Remarks

Aphonopelma chalcodes is herein considered a member of the problematic Iodius species group. Morphological and molecular data confirm that A. chalcodes is the sister lineage to the remaining species in the group (A. iodius, A. eutylenum, and A. johnnycashi). There are no major morphological features that can be used to distinguish A. chalcodes from these species so we must rely on molecular data and distributional information (A. chalcodes is largely restricted to Arizona south of the Grand Canyon). Other important ratios that distinguish males: A. chalcodes possess a larger M1/A1 (≥1.51; 1.51–1.79) than A. johnnycashi (≤1.43; 1.29–1.43). Other important ratios that distinguish females: A. chalcodes possess a smaller F1/M1 (≤1.68; 1.38–1.68) than A. catalina (≥1.71; 1.71–1.83), A. chiricahua (1.84 ± (only 1 specimen)), A. madera (≥1.73; 1.73; 1.73–2.15), and A. marxi (≥1.77; 1.77–1.88); by possessing a smaller P1/M1 (≤0.75; 0.61–0.75) than A. vorhiesi (≥0.75; 0.75–0.85). For both males and females, certain morphometrics have potential to be useful, though due to the amounts of variation, small number of specimens, and the small differences between species, no others are claimed to be significant at this time (see Suppl. material 2). During evaluation of traditional two-dimensional PCA morphospace and three-dimensional PCA morphospace (PC1~PC2~PC3), male and female A. chalcodes do not separate from the other species in the Iodius species group. Male A. chalcodes separate from syntopic species A. catalina, A. chiricahua, A. madera, and A. marxi in two-dimensional morphospace as well as all syntopic miniature tarantulas (A. mareki, A. paloma, A. parvum, A. prenticei, A. saguaro, and A. superstitionense), but do not separate from A. gabeli or A. peloncillo; when three-dimensional morphospace is evaluated, A. chalcodes separates from all of their syntopic species. Female A. chalcodes separate from their syntopic species A. chiricahua, A. madera, and A. marxi in traditional two-dimensional morphospace as well as all syntopic miniature tarantulas, but do not separate from A. catalina, A. gabeli, A. peloncillo or A. vorhiesi; when three-dimensional morphospace is evaluated, A. chalcodes separates from A. chiricahua, A. madera, A. marxi, A. peloncillo, and A. vorhiesi, but not A. catalina or A. gabeli. PC1, PC2, and PC3 explain ≥95% of the variation in all analyses. We examined the holotypes and freshly collected topotypic material of A. apacheum, A. minchi, A. schmidti, and A. stahnkei. Our morphological and molecular analyses fail to recognize these four species as separate, independently evolving lineages. As a consequence, we consider A. apacheum, A. minchi, A. schmidti, and A. stahnkei junior synonyms of A. chalcodes.

Mitochondrial DNA (CO1) identifies A. chalcodes as a polyphyletic group with some samples more closely related to specimens of A. iodius (Fig. 7); these latter samples of A. chalcodes were previously considered a putative cryptic species by Hamilton et al. (2014). The mtDNA also identifies A. vorhiesi as the sister lineage to A. chalcodes. The AE nuclear DNA on the other hand shows that A. chalcodes is a single lineage that is sister to the rest of the Iodius species group. Again, these results highlight how CO1 is not effective at accurately delimiting species boundaries within this group.

Aphonopelma chiricahua Hamilton, Hendrixson & Bond, sp. n.

Figures 36, 37, 38, 39

Types

Male holotype (APH_3191) collected 1 mile up the road (42 Forest Rd.) from the lookout trail, Cochise Co., Arizona, 31.886417 -109.173356 1, elev. 5083ft., 14.xi.2013, coll. Helen Snyder; deposited in AUMNH. Paratype female (APH_2097) from SWRS (Southwest Research Station, 5 miles W of Portal), Cochise Co., Arizona, 31.884056 -109.208261 5, elev. 5436ft., 30.xi.1965, coll. Jon Jenson; deposited in AMNH. Paratype male (APH_2105) from SWRS, Cochise Co., Arizona, 31.883356 -109.207107 5, elev. 5404ft., 31.x.1956, coll. E. Ordway; deposited in AMNH.

Etymology

The specific epithet is a noun in apposition taken from type locality, the Chiricahua Mountains outside of Portal, Arizona, where this new species appears to be endemic.

Diagnosis

Aphonopelma chiricahua (Fig. 36) is a member of the Marxi species group and can be distinguished by a combination of morphological, molecular, and geographic characteristics. Nuclear and mitochondrial DNA identifies A. chiricahua as a phylogenetically distinct lineage (Figs 78), supported as a sister lineage to A. catalina sp. n. (a species endemic to the Santa Catalina Mountains). The significant measurement that distinguishes male A. chiricahua from its closely related phylogenetic and syntopic species is A3. Male A. chiricahua can be distinguished by possessing a larger A3/M4 (≥0.65; 0.65–0.72) than A. catalina (≤0.52; 0.47–0.52), A. madera sp. n. (≤0.60; 0.54–0.60), A. parvum sp. n. (≤0.64; 0.53–0.64), A. peloncillo sp. n. (≤0.58; 0.45–0.58), and A. vorhiesi (≤0.57; 0.46–0.57). The significant measurement that distinguishes female A. chiricahua from its closely related phylogenetic and syntopic species is P1. Female A. chiricahua can be distinguished by possessing a smaller Cl/P1 (2.21 ± (only 1 specimen)) than A. catalina (≥2.75; 2.75–2.94), A. madera (≥2.71; 2.71–3.01), A. parvum (≥2.69; 2.69–3.04), A. peloncillo (≥2.71; 2.71–3.02), and A. vorhiesi (≥2.59; 2.59–2.88).

Figure 36.

Aphonopelma chiricahua sp. n. live photograph. Male holotype - APH_3191. Do not have a photograph of a live female specimen.

Description of male holotype

(APH_3191; Fig. 37). Specimen preparation and condition: Specimen collected live crossing road, preserved in 80% ethanol; original coloration faded due to preservation. Left legs I, III, IV, and left pedipalp removed for measurements and photographs; stored in vial with specimen. Right legs III & IV removed for DNA and stored at -80°C in the AUMNH (Auburn, AL). General coloration: Generally black or faded black. Cephalothorax: Carapace 11.42 mm long, 11.22 mm wide; densely clothed with black/faded black pubescence, slightly appressed to surface and longer than lower elevation species, slight iridescence; fringe covered in long setae not closely appressed to surface; foveal groove medium deep and straight; pars cephalica region rises gradually from foveal groove, gently arching anteriorly toward ocular area; AER slightly procurved, PER very slightly recurved; normal sized chelicerae; clypeus slightly extends forward on a curve; LBl 1.37, LBw 1.61; sternum hirsute, clothed with medium black, densely packed setae. Abdomen: Densely clothed in short black/brown pubescence with numerous longer, lighter setae interspersed (generally red or orange in situ), longer with a more hirsute appearance than lower elevation species; dense dorsal patch of black Type I urticating bristles (Cooke et al. 1972); ventral setae same as dorsal. Legs: Hirsute; densely clothed with medium length black/brown setae, and longer setae ventrally. Metatarsus I slightly curved. F1 12.72; F1w 3.28; P1 4.95; T1 11.37; M1 7.61; A1 6.16; F3 9.53; F3w 2.98; P3 4.11; T3 7.60; M3 7.79; A3 6.84; F4 11.41; F4w 3.20; P4 4.41; T4 9.67; M4 10.28; A4 7.78; femur III is normal - not noticeably swollen or wider than other legs. All tarsi fully scopulate. Extent of metatarsal scopulation: leg III (SC3) = 65.5%; leg IV (SC4) = 37.9%. Three ventral spinose setae and one retrolateral spinose seta on metatarsus III; nine ventral spinose setae, one prolateral spinose seta on metatarsus IV; two ventral spinose setae on tibia I. Coxa I: Prolateral surface a mix of fine, hair-like and thin/very thin tapered setae. Pedipalps: Hirsute; densely clothed in the same setal color as the other legs, with numerous longer ventral setae; one spinose seta at the apical, prolateral femur and four spinose setae on the prolateral tibia; PTl 7.34, PTw 2.82. When extended, embolus tapers with a gentle curve to the retrolateral side near apex; embolus slender, no keels.

Figure 37.

Aphonopelma chiricahua sp. n. A–I male holotype, APH_3191 A dorsal view of carapace, scale bar = 5mm B prolateral view of coxa I C dorsal view of femur III D ventral view of metatarsus III, scale bar = 3mm E ventral view of metatarsus IV, scale bar = 4mm F prolateral view of L pedipalp and palpal tibia, scale bar = 3.5mm G dorsal view of palpal bulb H retrolateral view of palpal bulb, scale bar = 1mm I prolateral view of tibia I (mating clasper), scale bar = 5mm.

Variation (7). Cl 6.837–11.42 (8.18±0.62), Cw 6.254–11.22 (8.269±0.86), LBl 0.684–1.368 (0.959±0.11), LBw 0.985–1.765 (1.292±0.11), F1 6.145–12.718 (8.731±0.77), F1w 1.898–3.281 (2.309±0.19), P1 2.859–4.947 (3.517±0.27), T1 5.851–11.372 (7.397±0.7), M1 4.09–7.61 (5.06±0.46), A1 3.572–6.165 (4.542±0.31), L1 length 22.568–42.812 (29.248±2.48), F3 5.591–9.531 (6.823±0.5), F3w 1.688–2.982 (2.147±0.18), P3 2.304–4.112 (2.896±0.23), T3 4.162–7.603 (5.286±0.43), M3 4.379–7.794 (5.317±0.45), A3 3.955–6.838 (5.003±0.35), L3 length 20.391–35.878 (25.325±1.95), F4 6.648–11.414 (8.181±0.62), F4w 1.74–3.205 (2.174±0.2), P4 2.524–4.414 (3.141±0.25), T4 5.784–9.674 (7.104±0.48), M4 5.772–10.277 (7.342±0.56), A4 4.944–7.78 (5.781±0.38), L4 length 25.672–43.559 (31.549±2.26), PTl 4.42–7.341 (5.424±0.36), PTw 1.888–2.82 (2.241±0.12), SC3 ratio 0.48–0.656 (0.556±0.02), SC4 ratio 0.33–0.404 (0.376±0.01), Coxa I setae = thin/very thin tapered, F3 condition = normal.

Description of female paratype

(APH_2097; Fig. 38). Specimen preparation and condition: Specimen collected live, preserved in unknown percentage of ethanol; original coloration faded due to preservation. Left legs I, III, IV, and pedipalp removed for photographs and measurements; stored in vial with specimen. No tissue for DNA. Genital plate with spermathecae removed and cleared, stored in vial with specimen. General coloration: Brown and faded black. Cephalothorax: Carapace 7.651 mm long, 7.479 mm wide; Hirsute, densely clothed with brown pubescence closely appressed to surface; fringe densely covered in longer setae; foveal groove medium deep and slightly procurved; pars cephalica region gently rises from thoracic furrow, arching anteriorly toward ocular area; AER slightly procurved, PER slightly recurved; robust chelicerae, clypeus extends forward on a slight curve; LBl 1.194, LBw 1.218; sternum hirsute, clothed with medium short brown setae. Abdomen: Densely clothed dorsally in brown setae with numerous longer, lighter setae interspersed (generally red or orange in situ); dense dorsal patch of black Type I urticating bristles (Cooke et al. 1972); ventral setae shorter than dorsal. Spermathecae: Paired and separate, tapering and curving medially towards capitate bulbs, with wide bases that are not fused. Legs: Hirsute; densely clothed in short and medium brown pubescence; F1 6.261; F1w 2.223; P1 3.453; T1 4.952; M1 3.398; A1 3.011; L1 length 21.075; F3 4.989; F3w 1.645; P3 2.332; T3 3.341; M3 3.231; A3 3.681; L3 length 17.574; F4 6.292; F4w 1.897; P4 3.17; T4 5.134; M4 5.114; A4 3.995; L4 length 23.705. All tarsi fully scopulate. Extent of metatarsal scopulation: leg III (SC3) = 56.2%; leg IV (SC4) = 27.7%. Two ventral spinose setae and one prolateral spinose seta on metatarsus III; six ventral spinose setae and one prolateral spinose seta on metatarsus IV. Coxa I: Prolateral surface a mix of fine, hair-like and tapered/thin tapered setae. Pedipalps: Densely clothed in the same setal color as the other legs; one spinose seta at the apical, prolateral femur, two prolateral spinose setae on the tibia.

Figure 38.

Aphonopelma chiricahua sp. n. A–F female paratype, APH_2097 A dorsal view of carapace, scale bar = 3.5mm B prolateral view of coxa I C ventral view of metatarsus III, scale bar = 2mm D ventral view of metatarsus IV, scale bar = 2mm E prolateral view of L pedipalp and palpal tibia F cleared spermathecae.

Material examined

United States: Arizona: Cochise: SWRS (5 miles W of Portal), 31.884056 -109.208261 5, 5436ft., [APH_2097, 30/11/1965, 1♀, Jon Jenson, AMNH]; Cave Creek Canyon, 31.885338 -109.175462 5, 5105ft., [APH_2101, 30/11/1963, 1♂, V. Roth, AMNH]; Upper Cave Creek Canyon, 31.900796 -109.229328 5, 5997ft., [APH_2102, 1966, 1♂, Marlene Posedly, AMNH]; South West Research Station, 31.883356 -109.207107 5, 5404ft., [APH_2105, 31/10/1956, 1♂, E. Ordway, AMNH]; Sunny Flat, 31.884963 -109.175964 4, 5108ft., [APH_2480-A, 30/10/1971, 1♂, V. Roth, AMNH]; South West Research Station-Portal, 31.883372 -109.205727 5, 5384ft., [APH_2480-B, 20/11/1971, 1♂, V. Roth, AMNH]; Chiricahua Mtns, 31.903946 -109.279016 6, 8432ft., [APH_2548, 4/11/1970, 1♂, Rustler and Long Park, Joan Harper, AMNH]; Cave Creek Canyon, on Portal Rd/42A (road to SWRS), 1 mile up the road from the lookout trail, 31.886417 -109.173356 1, 5083ft., [APH_3191, 14/11/2013, 1♂, Helen Snyder, AUMNH].

Distribution and natural history

Aphonopelma chiricahua is a sky island endemic restricted to the Chiricahua Mountains in Cochise County, Arizona at elevations ranging from 1550 to 2700 meters in oak woodland, pine-oak woodland, and mixed conifer communities (Fig. 39). Aphonopelma chiricahua can be found inhabiting the Madrean Archipelago Level III Ecoregion. Very little is known about the natural history of this elusive species. No burrows or shelters have been observed but these spiders probably seek refuge under rocks and rarely place silk around their burrow entrances. Most specimens in natural history collections have been collected near the AMNH’s Southwest Research Station. Aphonopelma chiricahua is probably the only tarantula found at higher elevations in the Chiricahua Mountains but four other species are known from the area (including A. chalcodes, A. gabeli, A. parvum, and A. vorhiesi) and might be syntopic with A. chiricahua at lower elevations near various canyon mouths. An unconfirmed adult female (no voucher specimen available, identification tentatively assigned based on a photograph and locality data) was found walking across a road in Cave Creek Canyon in August during the summer monsoon season (A. Abela 2006, pers. comm.). Vouchered adult males were collected between late October and late November; an adult male and female (no voucher specimens available, identification tentatively assigned based on video images and locality data) were found mating during daylight hours in early December along Echo Canyon Trail in Chiricahua National Monument (Chiricahua National Monument 2015, pers. comm.; https://www.facebook.com/video.php?v=785835144804065). The holotype male was collected wandering across 42 Forest Road in November, one year before this video was taken. These data indicate that the breeding season for this species is restricted to late autumn and early winter, similar to that of other high-elevation species in the region (A. catalina and A. madera).

Figure 39.

Aphonopelma chiricahua sp. n. distribution of known specimens. There is no predicted distribution map due to the limited number of sampling localities and restricted distribution this species possesses.

Conservation status

It is difficult to assess the conservation status of Aphonopelma chiricahua due to small sample sizes and the very cryptic nature of these spiders. This species does not occur outside of the Chiricahua Mountains so its narrow distribution is one factor that may threaten its future survival. These mountains have the advantage of being somewhat protected by their remoteness and management by the federal government (Coronado National Forest, Douglas Ranger District, Chiricahua National Monument); however, these habitats have also been subjected to habitat degradation from recent urban growth, human-caused forest fires, off-road driving, poorly managed livestock grazing, invasive species, recreational activities, human immigrants, and illegal drug trafficking (Coronado Planning Partnership 2008). Climate change in the sky island region (Brusca et al. 2013, Mitchell and Ober 2013, Moore et al. 2013, Hendrixson et al. 2015) also poses a potential threat to the future survival of A. chiricahua.

Remarks

Aphonopelma chiricahua is morphologically very similar to other Madrean sky island endemics in the Marxi species group, although generally smaller than A. catalina, A. madera, and A. vorhiesi. Other important ratios that distinguish males: A. chiricahua possess a larger Cl/M3 (≥1.46; 1.46–1.61) than A. catalina (≤1.42; 1.26–1.42), A. parvum (≤1.39; 1.20–1.39), A. peloncillo (≤1.40; 1.20–1.40), and A. vorhiesi (≤1.43; 1.24–1.43); by possessing a larger L3/Cl (≥2.98; 2.98–3.19) than A. madera (≤2.95; 2.71–2.95). Other important ratios that distinguish females: A. chiricahua possess a smaller M3/P4 (1.02 ± (only 1 specimen)) than A. catalina (≥1.48; 1.48–1.52), A. madera (≥1.39; 1.39–1.48), A. parvum (≥1.32; 1.32–1.54), A. peloncillo (≥1.39; 1.39–1.67), and A. vorhiesi (≥1.27; 1.27–1.64). For both males and females, certain morphometrics have potential to be useful, though due to the amounts of variation, small number of specimens, and the small differences between species, no other are claimed to be significant at this time (see Suppl. material 2). During evaluation of PCA morphospace, males of A. chiricahua separate from A. parvum and all other miniature species along PC1~2, but do not separate from A. catalina, A. madera, A. peloncillo, and A. vorhiesi. Though we only have one female of A. chiricahua, it appears to separate from all other sky island species (A. catalina, A. madera, A. peloncillo, and A. vorhiesi), grouping more closely with the miniature species in morphospace along PC1~2. Interestingly, A. chiricahua males separate from A. parvum, A. peloncillo, and A. vorhiesi in three-dimensional PCA morphospace (PC1~PC2~PC3), but do not separate from A. catalina, A. madera, A. marxi. There is only one A. chiricahua female, but she separates from all other phylogenetic sister species or syntopic species - A. catalina, A. madera, A. marxi, A. parvum, A. peloncillo, and A. vorhiesi. PC1, PC2, and PC3 explain ≥96% of the variation in all analyses.

This species was first identified as novel by Jung (1975) but was never formally described. Of particular note is the size of the holotype male and paratype female; the two specimens probably represent opposite extremes on the size spectrum for what is possible in this species. The rather large holotype male was chosen because it was a fresh specimen and could be associated with molecular data. The female, though small, is sexually mature (based on spermathecal development).

Aphonopelma eutylenum Chamberlin, 1940

Figures 40, 41, 42, 43, 44

Aphonopelma eutylenum Chamberlin, 1940: 9; male holotype and female allotype from San Diego, San Diego Co., California, 32.715738 -117.1610856, elev. 54ft., 1935, coll. unknown; deposited in AMNH. One male from San Diego, San Diego Co., California, 32.715738 -117.1610856, elev. 54ft., 20.vii.1925, coll. unknown; deposited in AMNH. One female paratype from San Diego, San Diego Co., California, 32.715738 -117.1610856, elev. 54ft., 28.v.1927, coll. unknown; deposited in AMNH. [examined]

Rhechostica eutylenum Raven, 1985: 149.

Aphonopelma eutylenum Smith, 1995: 99.

Aphonopelma chambersi Smith, 1995: 86; male holotype from Garner Valley, S of Idyllwild-Pine Cove, Riverside Co., California, 33.635459 -116.6439745, elev. 4475ft., no collecting date, coll. Aaron Chambers; deposited in BMNH. [examined] syn. n.

Aphonopelma clarum Chamberlin, 1940: 10; male holotype from mountains near Claremont, Los Angeles Co., California, 34.137812 -117.7181944, elev. 1571ft., no collecting date, coll. R.V. Chamberlin; deposited in AMNH. [examined]

Rhechostica clarum Raven, 1985: 149.

Aphonopelma clarum Smith, 1995: 89. syn. n.

Aphonopelma cryptethus Chamberlin, 1940: 16; male holotype from Los Angeles, Los Angeles Co., California, 34.052234 -118.2436856, elev. 309ft., 9.v.1908, coll. unknown; deposited in AMNH. Female allotype from Claremont, Los Angeles Co., California, 34.096676 -117.7197785, elev. 1166ft., 9.v.1908, coll. unknown; deposited in AMNH. [examined]

Rhechostica cryptethus Raven, 1985: 149.

Aphonopelma cryptethum Smith, 1995: 95. syn. n.

Aphonopelma sandersoni Smith, 1995: 138; male holotype and female allotype from San Bernardino Mountains., San Bernardino Co., California, 34.180742 -117.1646387, elev. 3137ft., x.1995, coll. R. Douglas; deposited in BMNH. [examined] syn. n.

Aphonopelma sullivani Smith, 1995: 149; male holotype from Coachella Valley, Palm Springs, Riverside Co., California, 33.767209 -116.3598686, elev. 277ft., ix.1991, coll. Michael Sullivan; deposited in BMNH. [examined] syn. n.

Diagnosis

Aphonopelma eutylenum (Fig. 40) is a member of the Iodius species group and can be identified by a combination of morphological, molecular, and geographic characteristics. Nuclear and mitochondrial DNA (CO1) identifies A. eutylenum as a strongly supported phylogenetically distinct monophyletic lineage (Figs 78), supported as the sister lineage to A. iodius and A. johnnycashi sp. n. Aside from measurements, no pronounced morphological features are useful for distinguishing A. eutylenum from closely related phylogenetic species. Significant measurements that distinguish male A. eutylenum from its closely related phylogenetic and syntopic species are T3 and the extent of scopulation on metatarsus IV. Male A. eutylenum can be distinguished by possessing a larger L4 scopulation extent (62%-77%) than A. steindachneri (21%–31%) and A. xwalxwal sp. n. (34%–48%); and by possessing a smaller T1/T3 (≤1.23; 1.16–1.23) than A. johnnycashi (≥1.25; 1.25–1.31) and A. steindachneri (≥1.28; 1.28–1.36). There are no significant measurements that separate male A. eutylenum from A. chalcodes and A. iodius. Significant measurements that distinguish female A. eutylenum from its closely related phylogenetic and syntopic species are F1 and the extent of scopulation on metatarsus IV. Female A. eutylenum can be distinguished by possessing a larger L4 scopulation extent (62%–75%) than A. steindachneri (24%–34%); and by possessing a smaller F1/M3 (≤1.52; 1.41–1.52) than A. johnnycashi (≥1.52; 1.52–1.61). There are no significant measurements that separate female A. eutylenum from A. chalcodes and A. iodius. Females of A. xwalxwal are unknown at this time and cannot be compared.

Figure 40.

Aphonopelma eutylenum Chamberlin, 1940 specimens, live photographs. Male (L) - APH_3207; Female (R) - APH_3108.

Description

Originally described by Chamberlin (1940).

Redescription of male exemplar

(APH_1088; Fig. 41). Specimen preparation and condition: Specimen collected live crossing road, preserved in 80% ethanol; deposited in AUMNH; original coloration faded due to preservation. Left legs I, III, IV, and left pedipalp removed for measurements and photographs; stored in vial with specimen. Right leg III removed for DNA and stored at -80°C in the AUMNH (Auburn, AL). General coloration: Black, grey, and faded brown. Cephalothorax: Carapace 16.30 mm long, 15.89 mm wide; Hirsute; densely clothed with light brown iridescent pubescence mostly appressed to surface; fringe covered in long setae not closely appressed to surface; foveal groove medium deep and straight; pars cephalica region rises gradually from foveal groove, gently arching anteriorly toward ocular area; AER and PER not in normal placement - looks to be developmental issues; normal sized chelicerae; clypeus extends forward on a curve; LBl 1.94, LBw 2.08; sternum hirsute, clothed with black, densely packed setae. Abdomen: Densely clothed in short black/brown pubescence with numerous longer red/orange setae interspersed; possessing a dense dorsal patch of black Type I urticating bristles (Cooke et al. 1972). Legs: Hirsute, particularly ventrally; densely clothed in a mix of black or faded black pubescence, femurs are darker. Metatarsus I slightly curved. F1 16.12; F1w 4.16; P1 6.93; T1 13.24; M1 13.41; A1 9.02; F3 14.19; F3w 4.41; P3 5.82; T3 10.82; M3 14.44; A3 8.30; F4 16.77; F4w 4.10; P4 6.71; T4 13.76; M4 18.16; A4 9.02; femur III is slightly swollen. All tarsi fully scopulate. Extent of metatarsal scopulation: leg III (SC3) = 83.5%; leg IV (SC4) = 71.6%. Two ventral spinose setae on metatarsus III; four ventral spinose setae on metatarsus IV; one large megaspine is present on the retrolateral tibia at the apex of the mating clasper - this can be seen when viewing the prolateral face of the mating clasper. Coxa I: Prolateral surface a mix of fine, hair-like and tapered setae. Pedipalps: Hirsute; densely clothed in the same setal color as the other legs, with numerous longer ventral setae; one spinose seta on the apical, prolateral femur and two spinose setae on the prolateral tibia; PTl 9.361, PTw 3.47. When extended, embolus tapers and gently curves to the retrolateral side near apex; embolus very slender, no keels; distinct ventral bulge at the shift from bulb to embolus.

Figure 41.

Aphonopelma eutylenum Chamberlin, 1940. A–I male specimen, APH_1088 A dorsal view of carapace, scale bar = 6mm B prolateral view of coxa I C dorsal view of femur III D ventral view of metatarsus III, scale bar = 5mm E ventral view of metatarsus IV, scale bar = 4.5mm F prolateral view of L pedipalp and palpal tibia, scale bar = 3mm G dorsal view of palpal bulb H retrolateral view of palpal bulb, scale bar = 1mm I prolateral view of tibia I (mating clasper), scale bar = 6mm.

Variation (8). Cl 12.01–18.42 (15.497±0.73), Cw 10.92–16.38 (14.213±0.68), LBl 1.53–2.14 (1.846±0.09), LBw 1.57–2.38 (2.113±0.11), F1 12.67–16.86 (15.34±0.5), F1w 2.85–4.51 (3.799±0.2), P1 4.94–7.22 (6.028±0.28), T1 10.52–13.62 (12.456±0.42), M1 10.08–14.57 (12.415±0.59), A1 6.71–9.18 (8.328±0.32), L1 length 45.83–61.45 (55.503±1.95), F3 10.75–14.19 (12.924±0.45), F3w 3.21–4.77 (3.944±0.19), P3 3.98–5.98 (5.186±0.28), T3 8.82–11.26 (10.247±0.36), M3 10.59–14.7 (13.179±0.67), A3 6.75–8.48 (7.771±0.26), L3 length 41.73–54.29 (49.353±2.08), F4 12.93–16.82 (15.241±0.54), F4w 2.85–4.41 (3.656±0.19), P4 4.61–6.71 (5.734±0.26), T4 10.79–14.21 (12.781±0.53), M4 13.84–18.71 (16.776±0.77), A4 7.42–9.47 (8.556±0.31), L4 length 49.65–65.06 (59.139±2.5), PTl 7.375–10.727 (8.984±0.35), PTw 2.319–3.47 (3.082±0.14), SC3 ratio 0.728–0.93 (0.827±0.02), SC4 ratio 0.626–0.772 (0.701±0.02), Coxa I setae = tapered, F3 condition = slightly swollen.

Description of female exemplar

(APH_1031; Figs 4243). Specimen preparation and condition: Specimen collected live from burrow, preserved in 80% ethanol; deposited in AUMNH; original coloration faded due to preservation. Left legs I, III, IV, and pedipalp removed for photographs and measurements; stored in vial with specimen. Right leg III removed for DNA and stored at -80°C in the AUMNH (Auburn, AL). Genital plate with spermathecae removed and cleared, stored in vial with specimen. General coloration: Faded brown/black. Cephalothorax: Carapace 16.55 mm long, 15.05 mm wide; Hirsute, densely clothed with light brown pubescence closely appressed to surface; fringe densely covered in longer setae; foveal groove medium deep and straight; pars cephalica region gently rises from thoracic furrow, arching anteriorly toward ocular area; AER procurved, PER slightly recurved; large chelicerae, clypeus extends forward on a curve; LBl 2.05, LBw 2.38; sternum hirsute, clothed with dark brown setae. Abdomen: Densely clothed dorsally in short black setae with numerous longer, lighter setae interspersed (generally red or orange in situ); dense dorsal patch of black Type I urticating bristles (Cooke et al. 1972); ventral side with shorter black/dark brown setae. Spermathecae: Paired and separate with wide bases, tapering and curving medially towards capitate bulbs, with smaller bulges laterally and medially. Legs: Very hirsute, particularly ventrally; densely clothed in medium and long brown pubescence, femurs darker. F1 13.77; F1w 4.14; P1 5.95; T1 10.15; M1 8.89; A1 7.44; F3 10.84; F3w 3.83; P3 5.29; T3 8.95; M3 9.07; A3 7.29; F4 13.70; F4w 3.86; P4 5.51; T4 11.43; M4 12.70; A4 8.14. All tarsi fully scopulate. Extent of metatarsal scopulation: leg III (SC3) = 88.2%; leg IV (SC4) = 74.7%. Two ventral spinose setae on metatarsus III; seven ventral spinose setae on metatarsus IV. Coxa I: Prolateral surface a mix of fine, hair-like and tapered setae. Pedipalps: Densely clothed in the same setal color as the other legs; one spinose seta on the apical, prolateral femur and five spinose setae on the prolateral tibia.

Figure 42.

Aphonopelma eutylenum Chamberlin, 1940. A–E female specimen, APH_1031. A dorsal view of carapace, scale bar = 8mm B prolateral view of coxa I C ventral view of metatarsus III, scale bar = 4.5mm D ventral view of metatarsus IV, scale bar = 5mm E prolateral view of L pedipalp and palpal tibia.

Figure 43.

Aphonopelma eutylenum Chamberlin, 1940. A–H cleared spermathecae. A eutylenum allotype B eutylenum paratype C APH_1018 D APH_1031 E APH_1045 F APH_2035 G AUMS_3303 H cryptethum allotype.

Variation (8). Cl 15.23–20.84 (18.248±0.72), Cw 13.33–17.78 (16.206±0.61), LBl 1.95–2.81 (2.353±0.12), LBw 2.28–3.04 (2.675±0.1), F1 12.87–15.78 (14.599±0.5), F1w 3.62–4.94 (4.357±0.2), P1 5.38–7.34 (6.479±0.25), T1 9.82–13.05 (11.243±0.42), M1 8.36–10.75 (9.565±0.33), A1 6.6–7.92 (7.36±0.17), L1 length 43.43–54.02 (49.137±1.72), F3 10.53–13.35 (12.208±0.42), F3w 3.3–4.67 (4.044±0.16), P3 5.12–7.26 (5.776±0.27), T3 8.3–9.93 (8.82±0.2), M3 8.64–11.06 (9.958±0.36), A3 6.49–7.87 (7.084±0.16), L3 length 39.24–48.1 (43.845±1.2), F4 13.21–16.57 (14.909±0.53), F4w 3.42–4.73 (4.106±0.19), P4 5.51–7.40 (6.312±0.32), T4 10.5–12.84 (11.509±0.31), M4 11.84–15.26 (13.32±0.48), A4 6.94–8.82 (7.779±0.26), L4 length 48.62–60.13 (54.192±1.91), SC3 ratio 0.772–0.906 (0.851±0.02), SC4 ratio 0.628–0.747 (0.683±0.02), Coxa I setae = tapered. Spermathecae variation can be seen in Figure 43.

Material examined

United States: California: Imperial: 1.5 miles N Gordons Well off of I-8, 32.7636 -114.966431 2, 248ft., [APH_0152, unknown, 1 juv, T.R. Prentice, AUMNH]; E of Brawley, off of Hwy 78, 32.969167 -115.259444 5, 3ft., [APH_3116, 10/1992, 1♀, T.R. Prentice, AUMNH]; East Mesa, 33.058096 -115.324147 5, 46ft., [AUMS_3332, 30/5/1995, 1♂, unknown, AUMNH]; East Mesa Exit of I-8, 2.1 miles N of I-8, 32.73013 -114.955351 4, 158ft., [AUMS_2358, 26/10/1993, 1♂, Greg Ballmer, AUMNH]; East Mesa, 1/2 mile E of Highline Canal and 3/10 mile S of Hwy 78, 32.968133 -115.290717 1, 37ft., [APH_3211, 15/11/2013, 1♂, W. Icenogle, AUMNH]; East Mesa, 5.4 miles E of Hwy 115 E of Wander Linder Exit of I-8, 32.788365 -115.207479 4, 50ft., [AUMS_3303, 30/5/1995, 1♀, unknown, AUMNH]; East Mesa, 5.4 miles E of Hwy 115 E of Wander Linder Exit, 2.1 miles N, 32.790359 -115.240569 4, 63ft., [AUMS_3341, 30/5/1995, 1♂, T.R. Prentice, AUMNH]; Hwy 78, 1.0 miles east of Highline Canal, 32.970575 -115.28374 4, 30ft., [AUMS_2356,-2357 3/11/1998, 2♂, Greg Ballmer, AUMNH]; Hwy 78, 3.35 miles E of East Highline Canal, N of 78 ~0.4 miles, 32.972797 -115.242665 4, 90ft., [AUMS_2674, unknown, 1♀, T.R. Prentice, AUMNH]; off I-8 split, W of Devils Canyon, W of Ocotillo, 32.67371 -116.10147 1, 2238ft., [APH_1028-1029, 18/5/2010, 1♂, 1♀, Chris A. Hamilton, Xavier Atkinson, AUMNH]; Pinto Wash, 32.708139 -115.715525 5, 13ft., [APH_2690, 5/3/1961, 1♂, Laurie Breese, AMNH]; 15 miles west of Glamis , 33.008844 -115.321813 5, -7ft., [APH_2154, 4/10/1961, 1♀, 1♂, Findlay E. Russell, AMNH]; Los Angeles: Claremont College - Frary Dining Hall on steps, 34.10018 -117.710527 4, 1225ft., [AUMS_3308, 11/11/1991, 1♂, Martin Ramirez, AUMNH]; Claremont, Mt. Baldy Rd and Mills Ave, 34.13823 -117.7072 1, 1660ft., [APH_1087, 8/10/2010, 1♂, Chris A. Hamilton, AUMNH]; Claremont, Thompson Creek Trail, 34.13716 -117.71389 1, 1582ft., [APH_1088-1089, 8/10/2010, 3♂, Chris A. Hamilton, Dylan Burke, Warren Burke, AUMNH]; [APH_2023-2027, 8/10/2010, 5♂, Chris A. Hamilton, AUMNH]; Glendora Ridge Rd., Angeles National Forest, 34.221163 -117.739386 6, 3666ft., [AUMS_2686, 17/11/1991, 1♂, John Adams, AUMNH]; Glendore, 34.136118 -117.865337 5, 774ft., [APH_2687, 18/10/1963, 1♂, R. Ayre, AMNH]; Morris Lake, 34.192644 -117.865749 5, 1207ft., [APH_2711, 8/1956, 1♂, Gay J. Kay, AMNH]; Palos Verde, 33.768911 -118.373032 5, 1132ft., [AUMS_2496, unknown, 1♂, C. Grover, AUMNH]; Phillips Ranch Rd., N of Freeway, 34.028759 -117.772093 5, 862ft., [AUMS_2498, 10/1992, 1♂, Dianna Hansen, AUMNH]; Pico Rivera, 33.983069 -118.096735 5, 163ft., [AUMS_3336, 4/10/1963, 1♂, unknown, AUMNH]; San Gabriel Mtns (N of Azusa), 34.2175 -117.8553 1, 1697ft., [APH_0895, 6/2007, 1♂, Chris A. Hamilton, AUMNH]; San Gabriel Mtns., near jct. Mt. Baldy Rd. and Glendora Ridge Rd., 0.8 miles up Glendora Ridge Rd. to dirt road winding around mtn., 34.228259 -117.670819 5, 4532ft., [AUMS_2490, 24/6/1989, 1♂, T.R. Prentice, AUMNH]; Orange: 21263 Mohler Drive, Anaheim, 33.853135 -117.755736 2, 617ft., [APH_2707, 1963, 2♂, F.W. Handsfield Jr., AMNH]; Brea, 33.916706 -117.899988 5, 362ft., [AUMS_2487, 15/10/1975, 1♂, Jim Lawrence, AUMNH]; Caspers Wilderness Park (San Juan Capistrano), 33.5572 -117.556783 1, 709ft., [APH_0981, 9/2009, 1♀, Kyle Dickerson, AUMNH]; El Toro (Marine Corps Air Station MCAS), 33.673927 -117.72924 5, 340ft., [AUMS_2588, 26/9/1966, 1♂, 1st Lt Jim Hardy, AUMNH]; Hidden Hills area, (Laguna Nigel) near San Juan Capistrano, 33.529839 -117.685743 5, 525ft., [AUMS_3309, 20/11/1991, 1♂, Scott Elgnam, AUMNH]; Laguna Beach , 33.542248 -117.783109 5, 16ft., [APH_2155, 26/7/1931, 1♀, R.V. Chamberlin, AMNH]; [APH_2472, 26/5/1931, 1 juv, Wilton Ivie, AMNH]; Laguna Hills, 33.599722 -117.699444 5, 372ft., [AUMS_2481, 28/10/1967, 1♂, Tom Simms, AUMNH]; Laguna Niguel, 33.522526 -117.707553 5, 407ft., [AUMS_2488, unknown, 1♀, unknown, AUMNH]; Lookout Point, Corona del Mar, 33.592518 -117.877554 5, 0ft., [APH_2708, 16/10/1962, 1♂, Jerry Parrish, AMNH]; San Clemente, E of Avenida Pico, off The Christianitos Trail, 33.455 -117.57523 1, 388ft., [APH_1035, 22/5/2010, 1♀, Chris A. Hamilton, Xavier Atkinson, AUMNH]; San Jauquin Hills, S of Lagunas Reservoir, off Hwy 133, 33.607836 -117.75685 5, 397ft., [AUMS_3316, 7/5/1989, 1♂, T.R. Prentice, AUMNH]; San Juan Capistrano, 33.501693 -117.662551 5, 127ft., [AUMS_3327, 23/8/1964, 2♀, 1♂, J.H. Menees, AUMNH]; Santa Ana, 33.745573 -117.867931 5, 112ft., [APH_2153, 17/7/1931, 2♀, R.V. Chamberlin, AMNH]; Starr Ranch, between Caspers and Clev. Nat Forest, 33.632326 -117.555835 5, 999ft., [AUMS_2399, 13/8/1997, 1♂, unknown, AUMNH]; Starr Ranch, N Perusker and Bell Canyon Rd, 33.626176 -117.556666 5, 919ft., [AUMS_2404, 4/8/1997, 1♂, unknown, AUMNH]; Riverside: 1 mile N of Winchester, 33.72247 -117.081149 4, 1558ft., [APH_2171, 13/10/1968, 1♂, W. Icenogle, AMNH]; 1 mile NW of Winchester, 33.71869 -117.096162 5, 1745ft., [AUMS_2590, 3/9/1990, 1♀, Gary Larson, AUMNH]; 2 miles N of Winchester, 33.734766 -117.076361 5, 1556ft., [AUMS_2491, 30/10/1995, 1♂, Ken Harbridge, AUMNH]; 2 miles NE of Winchester, 33.734476 -117.074678 5, 1562ft., [AUMS_3301, 13/10/1995, 1♀, W. Icenogle, AUMNH]; 7 miles E of Hemet - off Hwy 74 in San Jacinto Mtns, 33.72003 -116.7916 1, 2625ft., [APH_3114, 22/7/2012, 1 juv, Chris A. Hamilton, AUMNH]; at base of Double Butte, Washington Ave, 1/2 mile N of Winchester town center, 33.714 -117.085 1, 1492ft., [APH_3207, 28/10/2013, 1♂, Patrick Parker, AUMNH]; [APH_3208, 6/11/2013, 1♂, Kim Johnson, AUMNH]; Box Spring Mtns, 33.972945 -117.29587 6, 2351ft., [AUMS_2585, 7/1991, 2♂, T.R. Prentice, AUMNH]; Cajalco Canyon, 8.6 miles southeast of Corona at Lake Mathews, 33.829201 -117.462018 4, 1368ft., [APH_2692, 3/11/1963, 1♂, W. A. Powder, AMNH]; Chino Canyon, on road to Palm Springs Aerial Tramway (Valley Station), 3 miles W of junction with Hwy 111 San Jacinto Mtns, 33.8447 -116.60125 1, 1971ft., [APH_3138, 10/9/2013, 1♂, W. Icenogle, AUMNH]; Chino Canyon, Palm Springs Aerial Tramway (Valley Station), San Jacinto Mtns, 33.838433 -116.6132 1, 2532ft., [APH_3137, 22/9/2013, 1♂, W. Icenogle, AUMNH]; [APH_3139, 4/9/2013, 1♂, W. Icenogle, AUMNH]; [APH_3140, 5/9/2013, 1♂, W. Icenogle, AUMNH]; Deep Canyon, 33.664423 -116.372819 5, 764ft., [AUMS_2629, unknown, 1♂, unknown, AUMNH]; E of Perris Valley Airport, Perris, 33.760961 -117.202485 5, 1417ft., [AUMS_2628, 19/7/1994, 1♀, unknown, AUMNH]; East of Sun City off I-215, Sun Mtns east, 33.702126 -117.171559 5, 1849ft., [AUMS_3285, 5/2/1989, 1♂, T.R. Prentice, AUMNH]; Garner Valley, off Fobes Ranch Rd - off Hwy 74, S of Lake Hemet, 33.63184 -116.63173 1, 4501ft., [APH_3111-3113, 22/7/2012, 2♂, 1♀, Chris A. Hamilton, Amy Skibiel, AUMNH]; just S of the Garner Valley, outside Nightingale, off Piñon Flats Transfer Station Rd - off Hwy 74, Sawmill Trailhead, 33.5802 -116.45131 1, 4047ft., [APH_3106-3110, 21/7/2012, 2♂, 3♀, Chris A. Hamilton, Amy Skibiel, AUMNH]; Lake Elsinore, near J Bethlee home, 33.668144 -117.327394 5, 1293ft., [AUMS_2503, 9/1998, 1♂, J. Bethlee, AUMNH]; Lake Mathews, 33.828903 -117.437512 5, 1401ft., [AUMS_3284, 14/4/1968, 1♂, M. Cline, AUMNH]; Lake Mathews, heading N on Cajalco Rd., 0.5 mile E of junction La Sierra Ave., 33.827762 -117.451927 4, 1397ft., [AUMS_2517, 12/11/1998, 1♂, T.R. Prentice, AUMNH]; Lake Skinner, at entrance gate, 33.590905 -117.07877 5, 1408ft., [AUMS_2499, 26/11/1997, 1♂, Tom Ash, AUMNH]; Lake Skinner, below U3 on rd. above oaks, 33.604263 -117.038208 5, 1567ft., [AUMS_2681, 29/10/1992, 1♂, T.R. Prentice, AUMNH]; Lake Skinner, below U5, above lake, 33.609043 -117.037303 5, 1661ft., [AUMS_2507, 28/9/1997, 1♀, T.R. Prentice, AUMNH]; Lake Skinner, between road and B9, 33.580808 -117.088785 5, 1353ft., [AUMS_2255, 16/8/1998, 1♂, T.R. Prentice, AUMNH]; Lake Skinner, by 90 deg. turn in dirt rd., S. of U7?, S side of lake, 33.575862 -117.066618 5, 1539ft., [AUMS_2501, 3/9/1987, 1♀, T.R. Prentice, AUMNH]; Lake Skinner, found on N side of lake E of Borel rd, 33.588117 -117.028523 5, 1517ft., [AUMS_2581, 18/8/1999, 1♂, T.R. Prentice, AUMNH]; Lake Skinner, Rowson Canyon, 1.5 miles N of middle gate, 0.8 miles S of Loop Rd., 33.573703 -117.066585 4, 1519ft., [AUMS_2505, 28/9/1997, 1♀, T.R. Prentice, AUMNH]; Lake Skinner, S of B5, close to lake, 33.569155 -117.064281 5, 1534ft., [AUMS_2506, 28/9/1997, 1♂, T.R. Prentice, AUMNH]; Lake Skinner, S side, 33.578777 -117.057225 5, 1535ft., [AUMS_2518, 11/9/1997, 1♂, Adam Bucklin , AUMNH]; Lake Skinner, SE of H2O, 33.573965 -117.025814 5, 1605ft., [AUMS_3325, 3/9/1997, 1♀, Adam Backlin, AUMNH]; Lamb Canyon, 33.855805 -117.015244 5, 1590ft., [AUMS_2587, 23/7/1988, 1♂, T.R. Prentice, AUMNH]; Lamb Canyon - N of jct 79 and Gilman Springs Rd., 33.858779 -117.016279 5, 1627ft., [AUMS_2401, 14/10/1989, 1♂, T.R. Prentice, AUMNH]; Lamb Canyon (S of Beaumont), off Hwy 79, 33.85507 -117.00342 1, 1911ft., [APH_1041-1042, 27/5/2010, 2♀, Chris A. Hamilton, Tom Prentice, AUMNH]; Lamb Canyon, Hwy 79, 33.853149 -117.007918 5, 1660ft., [AUMS_2407, 27/6/1987, 1♂, T.R. Prentice, AUMNH]; Mt Vernon Park, base of Sugarloaf Mtn, 33.988098 -117.295535 5, 2078ft., [AUMS_2497, 30/9/1992, 1♀, James Adams, AUMNH]; Murrieta, 33.553914 -117.213923 5, 1103ft., [AUMS_2655, 13/10/2001, 1♂, Bill, AUMNH]; [AUMS_3297, 8/9/1996, 1♂, C. Webster, AUMNH]; P.L. Boyd Deep Canyon Reserve, 0.1 miles above reserve gate, 33.66925 -116.372791 4, 701ft., [AUMS_2337, 21/9/1995, 1♂, T.R. Prentice, AUMNH]; P.L. Boyd Deep Canyon Reserve, 0.1 miles below reserve gate, 33.672099 -116.372355 4, 670ft., [AUMS_2335, 27/9/1995, 1♂, T.R. Prentice, AUMNH]; P.L. Boyd Deep Canyon Reserve, 0.55 miles N of reserve station, 33.655658 -116.373614 4, 883ft., [AUMS_2336, 22/9/1995, 1♀, T.R. Prentice, AUMNH]; P.L. Boyd Deep Canyon Reserve, 0.6 miles below reserve gate, 33.677492 -116.370089 4, 615ft., [AUMS_2334, 27/9/1995, 1♂, T.R. Prentice, AUMNH]; [AUMS_2368, 21/9/1995, 1♂, T.R. Prentice, AUMNH]; P.L. Boyd-Deep Canyon Reserve, 1.1 miles below reserve gate, 33.571489 -116.248672 4, 540ft., [AUMS_3293, 21/9/1995, 1♂, T.R. Prentice, AUMNH]; Perris, 33.782519 -117.228648 5, 1468ft., [AUMS_2400, 20/7/1994, 1♂, T.R. Prentice, AUMNH]; [AUMS_2409, 20/7/1994, 1♂, T.R. Prentice, AUMNH]; Perris, between I-215 and Perris Valley Airport, just N of Hwy 74 exit, 33.770547 -117.208584 5, 1417ft., [AUMS_2486, 19/7/1994, 1♂, T.R. Prentice, AUMNH]; Perris, E of Perris Valley Airport, 33.774681 -117.210995 5, 1419ft., [AUMS_4196, 20/7/1994, 1♂, T.R. Prentice, AUMNH]; Perris, E of Perris Valley Airport, W of I-215, 33.766645 -117.21044 5, 1416ft., [AUMS_2403, 20/7/1993, 1♂, T.R. Prentice, AUMNH]; Perris, field between Perris Valley airport and 74G exit (north of exit), off I-215, 33.765075 -117.201682 5, 1415ft., [AUMS_3311, 24/10/1994, 1♂, T.R. Prentice, AUMNH]; Perris, Rd 11 to I-215 between 74E exit and Perris exit, W of I-215, 33.775744 -117.239759 5, 1529ft., [AUMS_2405, 20/9/1993, 1♀, W. Icenogle, AUMNH]; [AUMS_2483, 23/9/1993, 1♀, T.R. Prentice, AUMNH]; Pinyon Pines area, Palm Canyon Dr, 2 mils N of junction with Hwy 74, Santa Rosa Mtns, 33.609067 -116.471767 1, 4054ft., [APH_3142, 16/8/2013, 1♂, W. Icenogle, AUMNH]; Pinyon Pines, Piñon Rd just N of junction with Hwy 74, Santa Rosa Mtns, 33.583867 -116.455383 1, 4036ft., [APH_3141, 16/8/2013, 1♂, W. Icenogle, AUMNH]; S of Corona, off I-15 west side, Weirick Rd exit (Temescal Canyon), 1.5 miles W, 33.796868 -117.531405 5, 1732ft., [AUMS_2685, 4/9/1995, 1♀, John Hermesman, AUMNH]; S of Hemet, NE of Lake Skinner, off De Portola Rd and Crown Valley Rd, 33.64276 -116.992 1, 2307ft., [APH_1038, 27/5/2010, 1♀, Chris A. Hamilton, Tom Prentice, AUMNH]; [APH_1040, 27/5/2010, 1♀, Chris A. Hamilton, Tom Prentice, AUMNH]; S side of Lake Skinner, 33.574388 -117.054156 5, 1533ft., [AUMS_2504, 11/9/1997, 1♂, Adam Bucklin, AUMNH]; San Jacinto mountains, Palms to Pines Hwy 74, above Palm Desert, 33.664336 -116.398965 5, 1240ft., [AUMS_2480, 2/4/1988, 1♂, T.R. Prentice, AUMNH]; Santa Rosa Mtns, W side of Deep Canyon, Hwy 74, 1 mile S of jct with Carrizo Road, 33.5947 -116.422467 1, 3733ft., [APH_3206, 17/10/2013, 1♂, W. Icenogle, AUMNH]; Santa Rosa Plateau, 33.545816 -117.270534 5, 1779ft., [AUMS_2314, 25/10/1998, 1♂, Connell Dunning, AUMNH]; Santa Rosa Plateau (reserve), jct of Clinton Keith and Tenaja Rd. (bend), 33.52847 -117.273132 5, 1800ft., [AUMS_2509, 29/9/1998, 1 juv, Connell Dunning, AUMNH]; Santa Rosa Plateau Ecological Reserve, Lomas Trail 1.2 miles E of Clinton Keith Rd, 33.522687 -117.271312 4, 1949ft., [AUMS_2679, 29/9/1998, 1♂, C. Dunning, AUMNH]; Sun Mtns., Sun City, 33.717019 -117.216043 5, 1623ft., [AUMS_2493, unknown, 1♀, T.R. Prentice, AUMNH]; Temecula, 33.493639 -117.148365 5, 1047ft., [APH_2686, 8/6/1965, 1♂, Helen Wheeler, AMNH]; Temescal Canyon (Valley), N of Weirick Rd, exit off I-15, Bedford Motorway, 33.807554 -117.513595 5, 1008ft., [AUMS_3321, 24/9/1995, 1♂, John Hermesman, AUMNH]; UC Riverside - backside between Ent. Annex I and Boden Lab, 33.971765 -117.325971 5, 1091ft., [AUMS_2519, 13/12/1998, 1♂, T.R. Prentice, AUMNH]; Winchester, 33.706966 -117.084473 5, 1486ft., [APH_2167, 18/11/1967, 1♂, W. Icenogle, AMNH]; [APH_2168, 1/10/1967, 1♂, W. Icenogle, AMNH]; [AUMS_2402, unknown, 1♀, W. Icenogle, AUMNH]; [AUMS_2478, 11/9/1990, 2♂, W. Icenogle, AUMNH]; Arabian Gardens Mobile Estates, Indio, 33.72995 -116.24115 4, 9ft., [AUMS_2376, 9/1988, 1♂, Thomas R. Prentice, AUMNH]; San Bernardino: 1255 Colony Dr., Upland (base of Mt. Baldy), 34.1553 -117.672217 1, 2102ft., [APH_2035, 7/17/2011, 1♀, Jolene Stewart, AUMNH]; 620 Greenwood Avenue, Devore, 34.242532 -117.419506 2, 2690ft., [APH_0189, 25/9/2007, 1♂, Joshua Gutierrez, AUMNH]; Hwy 38, 1.8 miles W jct of Forest Falls, 34.099825 -116.98744 4, 3926ft., [AUMS_2484, 24/8/1990, 1♂, W. Icenogle, AUMNH]; N of Rancho Cucamonga, off Hermosa, 34.16862 -117.58182 1, 2565ft., [APH_1085, 8/10/2010, 1♂, Chris A. Hamilton, AUMNH]; N of Rancho Cucamonga, off Hermosa, 34.16742 -117.58117 1, 2477ft., [APH_1086, 8/10/2010, 1♂, Chris A. Hamilton, AUMNH]; San Bernardino (San Bernardino National Forest), in hills N of city, off Quail Canyon Rd and Del Rosa, 34.174 -117.252 1, 2299ft., [APH_1045, 1/6/2010, 1♀, Chris A. Hamilton, AUMNH]; San Diego: 0.5 W of most SW shore of Otay Lake, 32.608062 -116.939954 2, 491ft., [APH_0305, 27/10/2007, 1 juv, Dorian LaPaglia, AUMNH]; 12 miles south of San Diego, 32.556071 -117.075033 5, 30ft., [APH_2476, 15/10/1935, 1♂, W.J. Baerg, AMNH]; 2/10 mile W of Yaqui Pass road (S3), at the Anza-Borrego State Park sign, 3 miles S of jct with Borrego Springs Rd, 33.170767 -116.337967 1, 1207ft., [APH_3209, 8/11/2013, 1♂, W. Icenogle, AUMNH]; Anza Borrego desert, on S22 (Borrego Salton Seaway) W of Salton City, 33.30525 -116.208333 1, 957ft., [APH_3144, 9/11/2013, 1♂, Chris A. Hamilton, Brent E. Hendrixson, Molly Taylor, AUMNH]; Anza Borrego desert, W outside Borrego Springs off of Hwy S22, 33.228106 -116.410867 1, 1523ft., [APH_3143, 9/11/2013, 1♀, Chris A. Hamilton, Brent E. Hendrixson, Molly Taylor, AUMNH]; Anza-Borrego Desert State Park, Borrego Springs, 32.908056 -116.491047 6, 5115ft., [APH_2165, 31/10/1961, 1♂, Dalton E. Merkel, AMNH]; Anza-Borrego State Park, 33.039933 -116.402333 1, 2584ft., [APH_0982-0984, 9/2009, 2♀, 1♂, Kyle Dickerson, AUMNH]; [APH_2697, 12/1962, 1♂, Merkel, AMNH]; [APH_2704-2705, 12/1962, 3♂, Merkel, AMNH]; [APH_2716, 11/1962, 2♂, Merkel, AMNH]; [APH_2719, 10/1962, 1♂, Merkel, AMNH]; Anza-Borrego, Yaqui Pass, 33.151285 -116.346386 5, 1800ft., [AUMS_2680, 10/5/1987, 1♂, T.R. Prentice, AUMNH]; Borrego Valley, Borrego Springs Road, 8/10 mile SE of jct with Jaqui Pass road (S3), 33.204 -116.318417 1, 543ft., [APH_3210, 8/11/2013, 1♂, W. Icenogle, AUMNH]; Camp Pendleton, range 313A - HOLF, 33.313998 -117.314552 6, 353ft., [AUMS_2610, 20/8/unknown, 1♀, unknown, AUMNH]; Camp Pendleton, HOLF, San Mateo Creek, 33.469175 -117.475198 6, 415ft., [AUMS_2611, 20/8/1999, 1♀, unknown, AUMNH]; Camp Pendleton, plot C30, 33.313998 -117.314552 6, 353ft., [AUMS_2675, 30/5/1996, 1♂, T.R. Prentice, AUMNH]; Camp Pendleton, San Mateo Creek, 33.434796 -117.523894 6, 321ft., [AUMS_2317, 1990, 1♂, unknown, AUMNH]; [AUMS_2644, 1999, 1♂, Dan Holland, AUMNH]; [AUMS_3328, unknown, 1♂, Dan Holland, AUMNH]; Camp Pendleton, San Onofre Cr., 33.234926 -117.389044 6, 146ft., [AUMS_2582, 29/9/1999, 1♀, Dan Holland, AUMNH]; [AUMS_2258, 9/1999, 2♀, Dan Holland, AUMNH]; [AUMS_2251, 29/9/1999, 1♂, Dan Holland, AUMNH]; [AUMS_2397, 9/1999, 1♀, Dan Holland, AUMNH]; [AUMS_3348, 12/9/1999, 1♂, Dan Holland, AUMNH]; [AUMS_2315, 9/1999, 1♂, Dan Holland, AUMNH]; Chula Vista, 32.640023 -117.084004 5, 67ft., [AUMS_2502, 10/8/1997, 1♀, unknown, AUMNH]; De Luz, off De Luz Rd, NW of Fallbrook, 33.42858 -117.32127 1, 384ft., [APH_1037, 24/5/2010, 1♀, Chris A. Hamilton, Xavier Atkinson, AUMNH]; El Cajon, 32.794773 -116.962527 5, 433ft., [APH_2147, 4/5/1926, 1♀, unknown, AMNH]; Encanto, 32.711739 -117.061755 5, 230ft., [APH_2146, 18/8/1928, 1♂, unknown, AMNH]; [APH_2170, 12/11/1931, 1♂, unknown, AMNH]; Growmont, 32.799285 -116.999051 5, 715ft., [APH_2169, 17/8/1931, 1♂, unknown, AMNH]; La Mesa, 32.767829 -117.023084 5, 541ft., [APH_2152, 1/8/1938, 1♂, H. Stredwick, AMNH]; [APH_2162, 20/7/1925, 1♂, unknown, AMNH]; [APH_2163, 4/8/1925, 1♀, unknown, AMNH]; [APH_2164, 25/4/1927, 1♀, unknown, AMNH]; Laguna Beach , 33.542248 -117.783109 5, 16ft., [APH_2158, 20/7/1930, 1♂, Edward Lawrance, AMNH]; Lake Henshaw, on Hwy 76, 33.22337 -116.75292 1, 2801ft., [APH_1016-1017, 15/5/2010, 1♀, 1♂, Chris A. Hamilton, Xavier Atkinson, AUMNH]; Lake Henshaw, on Hwy 76 at Lake Henshaw campground, 33.23073 -116.76336 1, 2929ft., [APH_1014-1015, 15/5/2010, 2♀, Chris A. Hamilton, Xavier Atkinson, AUMNH]; Las Flores, 33.154632 -117.207187 5, 568ft., [APH_2145, 30/8/1936, 1♂, unknown, AMNH]; Lemon Grove, 32.742552 -117.031417 5, 453ft., [APH_2156, 10/10/1930, 1♂, unknown, AMNH]; [APH_2157, 5/8/1926, 1♀, unknown, AMNH]; off S6/Hwy 76, E of Rincon, W of S6 turnoff for Palomar Mtn, 33.29765 -116.92304 1, 2468ft., [APH_1020-1021, 15/5/2010, 2♀, Chris A. Hamilton, Xavier Atkinson, AUMNH]; Oriflamme Canyon, 2 miles W of S2, W of Anza-Borrego state park boundary, 33.004274 -116.460837 5, 2246ft., [AUMS_2612, 5/1990, 1♂, T.R. Prentice, AUMNH]; San Diego, 32.71533 -117.157253 6, 62ft., [APH_2148, 1935, 1♂, unknown, AMNH]; [APH_2149, 7/1962, 1♂, Chris Parrish, AMNH]; [APH_2159-2160, unknown, 1♀, 2♂, unknown, AMNH]; [APH_2161, 20/8/1925, 1♂, unknown, AMNH]; San Diego, Mission Trails Regional Park, 32.8402 -117.04561 1, 349ft., [APH_1031, 20/5/2010, 1♀, Chris A. Hamilton, AUMNH]; San Diego, Mission Trails Regional Park, Oak Canyon Trail and Grassland Trail intersect, 32.84467 -117.0437 1, 324ft., [APH_1009-1010, 11/5/2010, 2♀, Chris A. Hamilton, Xavier Atkinson, Kyle Dickerson, AUMNH]; Santa Ysabel, on Hwy 79, next to Santa Ysabel Open Space Preserve, 33.12607 -116.67841 1, 2938ft., [APH_1018-1019, 15/5/2010, 2♀, Chris A. Hamilton, Xavier Atkinson, AUMNH]; Santee, in hills N of where Carlton Hills Blvd ends, 32.86146 -116.99485 4, 780ft., [APH_0144, 5/2007, 1 juv, Gilbert Quintana, AUMNH]; Sweetwater Reserve near San Diego, 32.602142 -116.879255 5, 1345ft., [APH_2473, 21/9/1935, 2♀, W.J. Baerg, AMNH]; Wruck Canyon, San Ysidro, off Cactus Rd, 32.5519 -116.99642 1, 351ft., [APH_1025, 16/5/2010, 1♀, Chris A. Hamilton, Xavier Atkinson, Jordan Satler, AUMNH].

Distribution and natural history

Aphonopelma eutylenum has a distribution that stretches from the western part of the Transverse Ranges, south down the length of the California coast along the Peninsular Ranges, and west of the Mojave Desert (Fig. 44). Aphonopelma eutylenum inhabits the following Level III Ecoregions in California: Southern California/Northern Baja Coast, Sonoran Basin and Range, and Southern California Mountains. This species can be found in syntopy with a number of species across its distribution including A. iodius, A. steindachneri, and A. xwalxwal. The breeding season, when mature males abandon their burrows in search of females, occurs during the fall (generally September–November).

Figure 44.

Aphonopelma eutylenum Chamberlin, 1940. A distribution of known specimens B predicted distribution; warmer colors (red, orange, yellow) represent areas of high probability of occurrence, cooler colors (blue shades) represent areas of low probability of occurrence.

Conservation status

Aphonopelma eutylenum is widely distributed across Southern California and is very common. The species is likely secure although some localized populations in urbanized areas (e.g., Los Angeles and San Diego) are likely threatened by human encroachment and development.

Remarks

Aphonopelma eutylenum can easily be differentiated from A. steindachneri and A. xwalxwal by the extent of scopulation on legs III and IV, and from A. xwalxwal a larger body size. Female and immature male A. eutylenum can be distinguished from A. steindachneri and A. xwalxwal by body color as well. Other important ratios that distinguish males: A. eutylenum possess a smaller T1/F3 (≤1.00; 0.93–1.00) than A. steindachneri (≥1.01; 1.01–1.11) and A. xwalxwal (≥1.10; 1.10–1.17); by possessing a larger T3/A3 (≥1.30; 1.30–1.35) than A. johnnycashi (≤1.27; 1.18–1.27), but smaller than A. xwalxwal (≥1.41; 1.41–1.64). Other important ratios that distinguish females: A. eutylenum possess a larger M1/M4 (≥0.67; 0.67–0.78) than A. steindachneri (≤0.67; 0.62–0.67). For both males and females, certain morphometrics have potential to be useful, though due to the amounts of variation, small number of specimens, and the small differences between species, no others are claimed to be significant at this time (see Suppl. material 2). During evaluation of traditional two-dimensional PCA morphospace, male A. eutylenum separate from their syntopic species A. steindachneri and A. xwalxwal, but do not separate from the other species in the Iodius species group. Interestingly, when evaluating three-dimensional PCA morphospace (PC1~PC2~PC3), male A. eutylenum separates from A. johnnycashi, as well as A. steindachneri and A. xwalxwal. Female A. eutylenum separate in two-dimensional and three-dimensional morphospace from their syntopic species A. steindachneri and A. xwalxwal, but do not separate from the other species in the Iodius species group. PC1, PC2, and PC3 explain ≥95% of the variation in all analyses. We examined the holotypes and freshly collected topotypic material of A. chambersi, A. clarum, A. cryptethum, A. sandersoni, and A. sullivani. Our morphological and molecular analyses fail to recognize these five species as separate, independently evolving lineages. As a consequence, we consider A. chambersi, A. clarum, A. cryptethum, A. sandersoni, and A. sullivani junior synonyms of A. eutylenum.

Mitochondrial DNA (CO1) is problematic in the Iodius species group. While this locus identifies A. eutylenum as a monophyletic group, sister relationships are unclear. Nuclear DNA reveals the true evolutionary history of the A. eutylenum lineage and highlights the ineffectiveness of CO1 for accurately delimiting species boundaries within this group.

Aphonopelma gabeli Smith, 1995

Figures 45, 46, 47, 48, 49

Aphonopelma gabeli Smith, 1995: 100; male holotype from E of Tucson, Pima Co., Arizona, 31.956396 -110.3398177, elev. 4055ft., no collecting date, coll. Russ Gurley; deposited in BMNH. [examined]

Diagnosis

Aphonopelma gabeli (Fig. 45) is a member of the Moderatum species group and can be identified by a combination of morphological, molecular, and geographic characteristics. Nuclear and mitochondrial DNA identifies A. gabeli as a phylogenetically distinct monophyletic lineage (Figs 78), supported as the sister lineage to A. moellendorfi sp. n. and closely related to A. moderatum. Female A. gabeli can be distinguished from syntopic species by their unique spermathecae, noticeably large and robust chelicerae, and associated broad anterior carapace margin (Figs 45, 47). Male A. gabeli have an overall black body appearance with very long, thin legs. Significant measurements that distinguish male A. gabeli from its closely related phylogenetic and syntopic species are PTl and M4. Male A. gabeli can be distinguished by possessing a smaller PTl/M3 (≤0.63; 0.57–0.63) than A. anax (≥0.64; 0.64–0.76), A. armada (≥0.65; 0.65–0.75), A. hentzi (≥0.67; 0.67–0.81), A. chalcodes (≥0.65; 0.65–0.75), A. peloncillo sp. n. (≥0.71; 0.71–0.82), and A. vorhiesi (≥0.71; 0.71–0.87); and a smaller M1/M4 (≤0.74; 0.70–0.74) than A. moderatum (≥0.76; 0.76–0.81) and A. moellendorfi (≥0.75; 0.75–0.82). Significant measurements that distinguish female A. gabeli from its closely related phylogenetic and syntopic species are P1, M3, and extent of scopulation on metatarsus IV. Female A. gabeli can be distinguished by possessing a larger M3/M4 (≥0.73; 0.73–0.78) than A. moderatum (≤0.72; 0.67–0.72); a larger L4 scopulation extent (39%-53%) than A. peloncillo (32%-38%) and A. vorhiesi (26%-37%); a smaller L4 scopulation extent than A. chalcodes (63%-81%); and by possessing a smaller P1/F4 (≤0.46; 0.42–0.46) than A. armada (≥0.47; 0.47–0.51). Females of A. moellendorfi are unknown and cannot be compared.

Figure 45.

Aphonopelma gabeli Smith, 1995 specimens, live photographs. Female (L) - APH_1481; Male (R) - APH_0628.

Description

Male originally described by Smith (1995).

Redescription of male exemplar

(APH_1054; Fig. 46). Specimen preparation and condition: Specimen collected live crossing road, preserved in 80% ethanol; deposited in AUMNH; original coloration faded due to preservation. Left legs I, III, IV, and left pedipalp removed for measurements and photographs; stored in vial with specimen. Right leg III removed for DNA and stored at -80°C in the AUMNH (Auburn, AL). General coloration: Generally black or faded to brown. Cephalothorax: Carapace 16.79 mm long, 15.08 mm wide; densely clothed with black pubescence appressed to surface; fringe covered in long setae not closely appressed to surface; foveal groove deep and straight; pars cephalica region rises gradually from foveal groove, gently arching anteriorly toward ocular area; AER slightly procurved, PER recurved; normal sized chelicerae; clypeus extends slightly forward; LBl 1.90, LBw 2.05; sternum hirsute, clothed with short and medium length black, densely packed setae. Abdomen: Densely clothed in short black/brown pubescence with numerous longer red/orange setae interspersed; possessing a dense dorsal patch of black Type I urticating bristles (Cooke et al. 1972). Legs: Hirsute; densely clothed in a mix of short black/brown pubescence with longer ventral setae. Metatarsus I slightly curved. F1 16.15; F1w 3.65; P1 6.31; T1 12.7; M1 11.71; A1 7.98; F3 13.75; F3w 4.01; P3 5.53; T3 10.31; M3 13.01; A3 7.87; F4 16.26; F4w 3.75; P4 5.95; T4 12.76; M4 16.60; A4 8.55; femur III is normal - not noticeably swollen or wider than other legs. All tarsi fully scopulate. Extent of metatarsal scopulation: leg III (SC3) = 67.5%; leg IV (SC4) = 47.1%. Two ventral spinose setae on metatarsus III; five ventral spinose setae on metatarsus IV. Coxa I: Prolateral surface a mix of fine, hair-like and tapered setae. Pedipalps: Hirsute; densely clothed in the same setal color as the other legs, with numerous longer ventral setae; one spinose seta on the apical, prolateral femur and six spinose setae on the prolateral tibia; PTl 7.963, PTw 2.590. When extended, embolus tapers but quickly curves to the retrolateral side near apex; embolus very slender, no keels.

Figure 46.

Aphonopelma gabeli Smith, 1995. A–I male specimen, APH_1054 A dorsal view of carapace, scale bar = 5mm B prolateral view of coxa I C dorsal view of femur III D ventral view of metatarsus III, scale bar = 4.5mm E ventral view of metatarsus IV, scale bar = 4mm F prolateral view of L pedipalp and palpal tibia, scale bar = 5mm G dorsal view of palpal bulb H retrolateral view of palpal bulb, scale bar = 1mm I prolateral view of tibia I (mating clasper), scale bar = 5mm.

Variation (6). Cl 15.21–16.79 (15.667±0.24), Cw 13.4–15.08 (14.075±0.25), LBl 1.7–2.02 (1.872±0.06), LBw 1.91–2.27 (2.075±0.05), F1 15.63–17.21 (16.127±0.23), F1w 3.41–3.65 (3.518±0.04), P1 5.53–6.37 (6.022±0.13), T1 12.7–13.55 (13.067±0.12), M1 11.71–12.98 (12.328±0.18), A1 7.64–8.56 (7.953±0.14), L1 length 54.27–57.46 (55.497±0.44), F3 13.4–14.24 (13.665±0.13), F3w 3.36–4.01 (3.723±0.09), P3 4.67–5.53 (5.253±0.13), T3 10.02–10.89 (10.472±0.13), M3 12.86–13.88 (13.28±0.14), A3 7.33–7.91 (7.635±0.1), L3 length 48.39–51.87 (50.305±0.46), F4 14.81–17.47 (15.853±0.38), F4w 3.44–3.81 (3.592±0.06), P4 4.79–6.02 (5.423±0.2), T4 12.64–13.39 (12.985±0.12), M4 16.33–18.26 (17.022±0.28), A4 8.09–9.14 (8.673±0.15), L4 length 57.18–63.65 (59.957±0.88), PTl 7.677–8.48 (8.083±0.11), PTw 2.38–2.59 (2.525±0.03), SC3 ratio 0.591–0.721 (0.663±0.02), SC4 ratio 0.361–0.471 (0.416±0.02), Coxa I setae = tapered, F3 condition = normal.

Description of female exemplar

(APH_0680; Figs 4748). Specimen preparation and condition: Specimen collected live from burrow, preserved in 80% ethanol; deposited in AUMNH; original coloration faded due to preservation. Left legs I, III, IV, and pedipalp removed for photographs and measurements; stored in vial with specimen. Right leg III removed for DNA and stored at -80°C in the AUMNH (Auburn, AL). Genital plate with spermathecae removed and cleared, stored in vial with specimen. General coloration: Faded brown, black, and grey, medium length setae cover body; brownish-grey, with green tint following a molt (in situ). Cephalothorax: Carapace 18.34 mm long, 15.81 mm wide; densely clothed with brown pubescence closely appressed to surface; fringe densely covered in medium setae; broad anterior margin of carapace; foveal groove medium deep and straight; pars cephalica region rises from thoracic furrow more steeply than male, arching anteriorly toward ocular area; AER very slightly procurved, PER recurved; large, robust chelicerae; clypeus extends forward on a slight curve; LBl 2.38, LBw 2.44; sternum hirsute, clothed with brown, medium length setae. Abdomen: Densely clothed dorsally in short black setae with numerous longer, lighter setae interspersed (generally red or orange in situ); dense dorsal patch of black Type I urticating bristles (Cooke et al. 1972); ventral side with shorter dark brown setae. Spermathecae: Uniquely shaped; paired and separate that taper to a pocket, with wide bases that are not fused. Legs: Hirsute, particularly ventrally; densely clothed in short, brown pubescence with longer setae interspersed. F1 15.24; F1w 4.26; P1 6.26; T1 12.11; M1 10.20; A1 7.57; F3 12.72; F3w 3.58; P3 6.26; T3 8.88; M3 10.71; A3 7.73; F4 14.9; F4w 3.83; P4 6.55; T4 12.03; M4 13.75; A4 8.49. All tarsi fully scopulate. Extent of metatarsal scopulation: leg III (SC3) = 79.2%; leg IV (SC4) = 52.9%. One ventral spinose seta on metatarsus III; four ventral spinose setae on metatarsus IV. Coxa I: Prolateral surface a mix of fine, hair-like and tapered setae. Pedipalps: Densely clothed in the same setal color as the other legs; one spinose seta on the apical, prolateral femur and two spinose setae on the prolateral tibia.

Figure 47.

Aphonopelma gabeli Smith, 1995. A–E female specimen, APH_0680 A dorsal view of carapace, scale bar = 7mm B prolateral view of coxa I C ventral view of metatarsus III, scale bar = 4mm D ventral view of metatarsus IV, scale bar = 4mm E prolateral view of L pedipalp and palpal tibia.

Figure 48.

Aphonopelma gabeli Smith, 1995. A–F cleared spermathecae A APH_0044 B APH_0642 C APH_0680 D APH_0946 E APH_1338 F Jung’s “portal” paratype.

Variation (6). Cl 14.69–18.34 (16.535±0.53), Cw 12.65–15.81 (14.145±0.5), LBl 2.07–2.41 (2.29±0.06), LBw 2.29–2.82 (2.485±0.08), F1 12.13–15.24 (13.288±0.45), F1w 3.67–4.26 (3.917±0.09), P1 5.41–6.26 (5.772±0.15), T1 9.35–12.11 (10.285±0.4), M1 7.41–10.2 (8.33±0.43), A1 5.83–7.57 (6.647±0.24), L1 length 40.18–51.38 (44.322±1.6), F3 9.63–12.72 (10.783±0.44), F3w 3.44–3.76 (3.575±0.05), P3 4.27–6.26 (5.053±0.28), T3 7.08–8.88 (7.773±0.26), M3 7.78–10.71 (8.75±0.41), A3 6.09–7.73 (6.593±0.24), L3 length 35.81–46.30 (38.953±1.56), F4 12.21–14.90 (13.065±0.41), F4w 3.43–3.83 (3.642±0.06), P4 4.67–6.55 (5.553±0.27), T4 9.31–12.03 (10.187±0.4), M4 10.5–13.75 (11.663±0.48), A4 6.71–8.49 (7.168±0.28), L4 length 43.83–55.72 (47.637±1.7), SC3 ratio 0.725–0.805 (0.762±0.01), SC4 ratio 0.397–0.529 (0.471±0.02), Coxa 1 setae = tapered. Spermathecae variation can be seen in Figure 48.

Material examined

United States: Arizona: Cochise: 0.1 mi. west of Portal, 31.913699 -109.143184 4, 4780ft., [APH_2356, 2/7/1961, 1♂, J. Cole, AMNH]; 0.5 miles east of Portal, 31.914884 -109.149967 4, 4846ft., [APH_2364, 1/7/1961, 1♂, J. Cole, AMNH]; 1 mile southwest of Portal, 31.903762 -109.152806 5, 4918ft., [APH_2363, 2/9/1960, 1♀, R. Zweifel, AMNH]; 1 mile west of Portal, 31.9135 -109.158499 5, 4961ft., [APH_2365, 4/7/1963, 1♂, Steve Aaron, AMNH]; 1.7 miles northeast of Portal on San Simon Rd., 31.930194 -109.12083 4, 4570ft., [APH_2377, 21/7/1961, 1♂, J. Cole, AMNH]; 15.5 miles S I-10 on Noland Rd, 32.032719 -109.186732 1, 4478ft., [APH_1337-1338, 2/8/2011, 2♀, Brent E. Hendrixson, Brendon Barnes, Nate Davis, Jake Storms, AUMNH]; 2 miles northeast of Portal, 31.934159 -109.117117 5, 4534ft., [APH_2380, 10/6/1963, 1♀, Cazier and Mortenson, AMNH]; 2.5 miles southeast of Portal on Portal Rd., 31.902146 -109.10097 4, 4478ft., [APH_2372, 6/7/1973, 1♂, R. Zweifel, AMNH]; 2.6 miles NW AZ/NM state line on Portal Rd, 31.89302489 -109.0856873 1, 4349ft., [APH_0385, 31/7/2008, 1♂, Alice Abela, AUMNH]; 7.7 miles S I-10 on Noland Rd, 32.145844 -109.173364 1, 3872ft., [APH_1336, 2/8/2011, 1♀, Brent E. Hendrixson, Brendon Barnes, Nate Davis, Jake Storms, AUMNH]; Apache Pass Rd, S of I-10, 32.267466 -109.464574 1, 3854ft., [APH_0712, 20/7/2009, 1 juv, Brent E. Hendrixson, Nate Davis, AUMNH]; Cave Creek Canyon, 1.3 miles northeast of Ranger Station on Cave Creek Canyon Rd. near Portal, 31.896202 -109163672 4, 4961ft., [APH_2383, 11/7/1973, 1♂, A. Bush, AMNH]; Chiricahua Mtns, 31.929812 -109.382285 6, 6119ft., [APH_2367, 8/1972, 2♂, J.A.L. Cooke, AMNH]; Dos Cabezas, 32.114138 -109.920729 5, 4219ft., [APH_2378, 19/9/1954, 1♂, G. Bradt, M. Oazier, AMNH]; Fan Road, 5.5 miles NE Bowie, 32.375306 -109.447667 1, 3640ft., [APH_0393, 24/7/2008, 1♂, Kari and Hunter McWest, AUMNH]; Portal, 31.913699 -109.1414 5, 4770ft., [APH_2357, 8/7/1964, 2♂, D. Rich, AMNH]; [APH_2359, 1/8/1965, 1♂, W.J. Gerstch, AMNH]; [APH_2360, 25/6/1962, 1♂, W.J. Gerstch, AMNH]; [APH_2362, 10/7/1962, 1♀, Melinda Stebbins, AMNH]; [APH_2368, 4/7/1961, 1♂, J. Cole, AMNH]; [APH_2369, 15/8/1962, 1♀, C. Parrish and W.J. Gertsch, AMNH]; [APH_2381, 14/6/1962, 1♀, W.J. Gerstch, AMNH]; Portal Rd, 31.884644 -109.071997 5, 4250ft., [APH_0390, unknown, 1♂, Alice Abela, AUMNH]; S of I-10 on Noland Rd, 32.213729 -109.176179 1, 3645ft., [APH_0711, 20/7/2009, 1 juv, Brent E. Hendrixson, Nate Davis, AUMNH]; Sulphur Canyon, 9 miles west of Rodeo, 31.836758 -109.188226 5, 7060ft., [APH_2355, 23/7/1955, 1♂, Guy Miller, AMNH]; Wilcox, 32.252851 -109.83201 5, 4170ft., [APH_2382, 12/7/1954, 1♂, W.J. Gerstch, AMNH]; Graham: 0.25 miles E Hwy-191 on Tanque Rd, 32.605235 -109.682418 1, 3873ft., [APH_1329-1330, 1/8/2011, 2♀, Brent E. Hendrixson, Brendon Barnes, Nate Davis, Jake Storms, AUMNH]; [APH_1481-1482, 1/8/2012, 1♀, 1♂, Brent E. Hendrixson, Brendon Barnes, Austin Deskewies, AUMNH]; [APH_1489, 4/9/2012, 1♀, Brent E. Hendrixson, AUMNH]; 0.4 miles E Hwy-191 on Tanque Rd, 32.606204 -109.681524 1, 3891ft., [APH_1184, 25/7/2010, 1♀, Brent E. Hendrixson, Brendon Barnes, Nate Davis, AUMNH]; 3.6 miles E Hwy-191 on Tanque Rd, 32.619274 -109.633622 1, 3623ft., [APH_1231, 8/8/2010, 1♂, Brent E. Hendrixson, Ashley Bailey, Andrea Reed, AUMNH]; 5.8 miles E Hwy-191 on Tanque Rd, 32.621499 -109.596498 1, 3481ft., [APH_1236, 8/8/2010, 1♂, Brent E. Hendrixson, Ashley Bailey, Andrea Reed, AUMNH]; along Hwy-191, 32.662275 -109.701131 1, 3530ft., [APH_1179, 25/7/2010, 1♂, Brent E. Hendrixson, Brendon Barnes, Nate Davis, AUMNH]; dirt road S of US-70, 32.744869 -109.344099 1, 4088ft., [APH_0641-0642, 12/7/2009, 2♀, Brent E. Hendrixson, Nate Davis, AUMNH]; Hwy-366, near Hwy-191, 32.726066 -109.71822 1, 3266ft., [APH_0637-0638, 11/7/2009, 2♂, Brent E. Hendrixson, Nate Davis, AUMNH]; Klondyke Rd, SW of Hwy-70, 32.914146 -109.975734 1, 3110ft., [APH_0698-0699, 18/7/2009, 2 juv, Brent E. Hendrixson, Nate Davis, AUMNH]; [APH_0701-0708, 18/7/2009, 2♀, 6♂, Brent E. Hendrixson, Nate Davis, AUMNH]; Tanque Rd, near Hwy-191, 32.604126 -109.681695 1, 3887ft., [APH_0627-0632, 11/7/2009, 5♂, 1 juv, Brent E. Hendrixson, Nate Davis, AUMNH]; [APH_0635, 11/7/2009, 1♀, Brent E. Hendrixson, Nate Davis, AUMNH]; Greenlee: 0.4 miles N Hwy-75 on Goat Camp Rd, 32.755403 -109.110492 1, 3726ft., [APH_1351, 5/8/2011, 1 juv, Brent E. Hendrixson, Brendon Barnes, Nate Davis, Jake Storms, AUMNH]; New Mexico: Chaves: N of Roswell, 33.50677 -104.52311 5, 3626ft., [APH_0316, 10/7/2007, 1♀, Rick C. West, AUMNH]; rest area near Hagerman, near NM-249, 33.09694 -104.44167 2, 3559ft., [APH_0044, 20/6/2002, 1♀, Shasta Michaels, JJ East, AUMNH]; Dona Ana: 0.9 miles NE I-10 on CR-B19 (I-10 Exit 155), 32.130859 -106.626039 1, 4030ft., [APH_0538, 5/6/2009, 1♀, Brent E. Hendrixson, Courtney Dugas, Sloan Click, AUMNH]; 2711 Los Misioneros, Las Cruces, 32.336008 -106.749999 2, 4170ft., [APH_1461, 27/6/2012, 1♂, Jesse Ortiz, AUMNH]; Aguirre Springs Rd, 32.43067 -106.547785 1, 5258ft., [APH_0655, 13/7/2009, 1 juv, Brent E. Hendrixson, Nate Davis, AUMNH]; [APH_0662-0664, 14/7/2009, 3 juv, Brent E. Hendrixson, Nate Davis, AUMNH]; along Hwy-9, 31.79866 -106.907625 1, 4113ft., [APH_0653, 13/7/2009, 1 juv, Brent E. Hendrixson, Nate Davis, AUMNH]; Just outside Las Cruces, on Aguirre Spring Road, 32.431202 -106.54921 5, 5306ft., [APH_0002, 8/2003, 1♂, Roy Thibodeau, AUMNH]; Eddy: 0.5 miles W US-62/180 on CR 408 (Dark Canyon Rd), 32.28746 -104.28966 1, 3330ft., [APH_0366-0368, 22/6/2008, 3♂, Shasta Michaels, AUMNH]; 1.2 miles W US-62/180 on CR 408 (Dark Canyon Rd), 32.28969 -104.30144 1, 3350ft., [APH_0364-0365, 21/6/2008, 2♂, Shasta Michaels, AUMNH]; 1.7 miles W US-62/180 on CR-408, 32.293874 -104.309514 1, 3392ft., [APH_0542-0543, 6/6/2009, 2 juv, Brent E. Hendrixson, Courtney Dugas, Sloan Click, AUMNH]; 2.3 miles W US-62/180 on CR 408, 32.29345 -104.31993 1, 3434ft., [APH_0061, 14/6/2001, 1♀, Brent E. Hendrixson, AUMNH]; near jct. CR-1 and US-82, 32.830265 -104.797078 1, 4228ft., [APH_0541, 6/6/2009, 1♀, Brent E. Hendrixson, Courtney Dugas, Sloan Click, AUMNH]; Grant: Silver City, pinyon pine habitat, 32.770075 -108.280326 4, 5944ft., [APH_1293, 7/2011, 1 juv, Ken McNeil, AUMNH]; Hidalgo: 1.6 miles S I-10 on Hwy-80, 32.213781 -108.951553 1, 4233ft., [APH_1180, 26/7/2010, 1♂, Brent E. Hendrixson, Brendon Barnes, Nate Davis, AUMNH]; 15.5 miles north of Rodeo, 32.049697 -109.030541 5, 3934ft., [APH_2376, 11/7/1960, 1♂, Zweifel, AMNH]; 18 miles north of Rodeo, 32.089638 -109.036588 5, 3976ft., [APH_2361, 7/7/1956, 1♂, H. Howden, AMNH]; 19.5 miles north of Rodeo, 32.11859 -109.03127 5, 4042ft., [APH_2385, 11/7/1960, 1♂, Zweifel, AMNH]; 21.5 miles north of Rodeo, 32.141862 -109.028611 5, 4124ft., [APH_2371, 11/7/1960, 1♂, Zweifel, AMNH]; 4.1 miles E Hwy-80 on Hwy-9, 31.936704 -108.970448 1, 4187ft., [APH_0678, 16/7/2009, 1♀, Brent E. Hendrixson, Nate Davis, AUMNH]; 5 miles north of junction of Animas Rd. and rt. 80, 31.443254 -109.827763 5, 4678ft., [APH_2374, 23/7/1960, 1♂, R. Zweifel, AMNH]; 5 miles south of Road Forks, 32.275308 -108.794078 5, 4354ft., [APH_2384, 2/7/1962, 1♂, W.J. Gerstch, AMNH]; 6 miles north of Lordsburg, 32.436395 -108.707975 5, 4432ft., [APH_2358, 28/7/1962, 1♂, unknown, AMNH]; 8 miles north of Rodeo, 31.952008 -109.030121 5, 4078ft., [APH_2370, 14/7/1963, 1♂, C. Bagwell, AMNH]; along Hwy-338, 31.807343 -108.801127 1, 4675ft., [APH_1192-1194, 26/7/2010, 3♂, Brent E. Hendrixson, Brendon Barnes, Nate Davis, AUMNH]; along Hwy-80 at CR-C078, 32.101572 -108.957714 1, 4408ft., [APH_0676, 15/7/2009, 1 juv, Brent E. Hendrixson, Nate Davis, AUMNH]; along Hwy-80 at Granite Gap, 32.086958 -108.977019 1, 4495ft., [APH_1483, 1/8/2012, 1♂, Brent E. Hendrixson, Brendon Barnes, Austin Deskewies, AUMNH]; along Hwy-9, just E of Continental Divide, 31.963316 -108.673824 1, 4489ft., [APH_0680, 16/7/2009, 1♀, Brent E. Hendrixson, Nate Davis, AUMNH]; along Hwy-9, just west of Gas Line Rd, 31.935932 -108.940821 1, 4355ft., [APH_0677, 15/7/2009, 1♂, Brent E. Hendrixson, Nate Davis, AUMNH]; along Hwy-90, NE of Lordsburg, 32.469585 -108.60797 1, 5005ft., [APH_0646, 12/7/2009, 1 juv, Brent E. Hendrixson, Nate Davis, AUMNH]; along US-70, SE of Hwy-92, 32.612037 -108.984613 1, 4151ft., [APH_0643-0644, 12/7/2009, 1♀, 1 juv, Brent E. Hendrixson, Nate Davis, AUMNH]; Cieanega Lake, 15.5 miles north of Rodeo, 32.049697 -109.030541 5, 3934ft., [APH_2375, 14/7/1961, 1♂, J. Cole, AMNH]; Lordsburg, vicinity of Fraggle Rock, 32.31738 -108.81833 4, 4250ft., [APH_1294, 7/2011, 1♂, Ken McNeil, AUMNH]; Rodeo, 31.950087 -109.031176 5, 4085ft., [APH_2366, 17/7/1963, 2♂, V. Roth, AMNH]; Rt. 9, 2 miles east of juncture with US 80, 31.920498 -109.070014 4, 4301ft., [APH_2387, 3/7/1958, 2♂, Robert Chew, AMNH]; Luna: 0.5 miles E Hidalgo Co. Line along I-10, 32.208983 -108.221098 1, 4547ft., [APH_1178, 23/7/2010, 1♂, Brent E. Hendrixson, Brendon Barnes, Nate Davis, AUMNH]; along Hwy-11, N of Columbus, 31.935144 -107.670986 1, 4220ft., [APH_0650-0651, 13/7/2009, 2 juv, Brent E. Hendrixson, Nate Davis, AUMNH]; along Hwy-180, 32.431967 -107.90736 1, 4691ft., [APH_0647-0649, 13/7/2009, 3 juv, Brent E. Hendrixson, Nate Davis, AUMNH]; along Hwy-9, 31.828861 -107.320841 1, 4141ft., [APH_0652, 13/7/2009, 1 juv, Brent E. Hendrixson, Nate Davis, AUMNH]; along Hwy-9, 2.3 miles SE Grant County Line, 31.868935 -108.182075 1, 4605ft., [APH_0665, 14/7/2009, 1 juv, Brent E. Hendrixson, Nate Davis, AUMNH]; Cookes Canyon Rd, A019, 3.4 miles NW NM-26, 32.453472 -107.614333 1, 4710ft., [APH_0395, 28/7/2008, 1♂, Kari and Hunter McWest, AUMNH]; Deming, on ramp to I-10, 32.267886 -107.780667 1, 4817ft., [APH_1177, 23/7/2010, 1♂, Brent E. Hendrixson, Brendon Barnes, Nate Davis, AUMNH]; Little Florida Mtns, 1 miles SE on Bonita Rd from turnoff on Gap Rd, 32.15817 -107.58858 4, 4480ft., [APH_0194, 3/9/2007, 1♀, Lorenzo Prendini, Jeremy Huff, AUMNH]; Sierra: near Truth or Consequences, 0.2 miles W I-25 off Exit 79, 33.157129 -107.257152 1, 4491ft., [APH_1295, 24/7/2011, 1 juv, Brent E. Hendrixson, Shasta Michaels, AUMNH]; Socorro: Escondida, 34.05833 -106.89083 5, 4607ft., [APH_0034, 7/8/2006, 1♀, Kristin Greene, AUMNH]; Texas: Andrews: SW4001, 32.113981 -102.615814 1, 3139ft., [APH_1053-1054, 6/7/2010, 2♂, Skyler Stevens, AUMNH]; SW7000 and SW6601, 32.110219 -102.710939 1, 3233ft., [APH_1055, 6/7/2010, 1♂, Skyler Stevens, AUMNH]; SW8000 and SW3001, 32.111511 -102.566667 1, 3122ft., [APH_1050-1051, 6/7/2010, 2♂, Skyler Stevens, AUMNH]; Brewster: 1.85 miles N Ranch Rd 2627 on Hwy-385, 29.71843 -103.15894 2, 2759ft., [APH_1469, 22/6/2012, 1♂, Darryl Burton, AUMNH]; 14 miles N Ranch Rd 2627 on Hwy-385, 29.86536 -103.24919 2, 3215ft., [APH_1472, 24/6/2012, 1♂, Darryl Burton, AUMNH]; 14.3 miles NE jct US-90 on US-67, 30.53666 -103.39322 1, 3831ft., [APH_0029, 18/6/2001, 1♂, Jeff Owens, AUMNH]; 5.5 miles N Ranch Rd 2627 on Hwy-385, 29.76887 -103.16619 2, 2840ft., [APH_1471, 24/6/2012, 1♂, Darryl Burton, AUMNH]; Crane: off Hwy 385, N or Crane, 31.41690278 -102.3563111 2, 2550ft., [APH_1386, 29/8/2011, 1 juv, Darryl Burton, AUMNH]; Ector: Cowden H Ranch, 32.07805 -102.780783 6, 3316ft., [APH_0940, 2006, 1♀, Dave Moellendorf, AUMNH]; [APH_0943, 9/2008, 1♀, Chris A. Hamilton, AUMNH]; [APH_0946, 9/2008, 1♀, Chris A. Hamilton, AUMNH]; El Paso: 3457 Red Sails Drive, El Paso, 31.79305 -106.313143 1, 3987ft., [APH_3126, 18/7/2013, 1♂, Jackie Ortegon, AUMNH]; Rest Area, 1.4 miles SE FM-793 (SE Fabens), 31.503951 -106.116307 1, 3779ft., [APH_0537, 5/6/2009, 1♀, Brent E. Hendrixson, Courtney Dugas, Sloan Click, AUMNH]; Gaines: Seminole, 32.72695 -102.660533 1, 3327ft., [APH_0850-0854, 9/2008, 5♀, Chris A. Hamilton, AUMNH]; [APH_0888, 9/2008, 1♀, Chris A. Hamilton, AUMNH]; Seminole, 0.4 miles NW 11th St on Hwy-214, 32.729362 -102.661669 1, 3329ft., [APH_0545, 7/6/2009, 1♀, Brent E. Hendrixson, Courtney Dugas, Sloan Click, AUMNH]; Midland: CR60, 32.010556 -102.2275 1, 2888ft., [APH_1060, 2/7/2010, 1♂, Skyler Stevens, AUMNH]; FM 1788 S, 31.762925 -102.16813 2, 2895ft., [APH_1372-1373, 27/6/2011, 2♂, Darryl Burton, AUMNH]; Reeves: outside Carlsbad, 31.83806 -103.88556 1, 2780ft., [APH_0018-0021, 20/6/2002, 4♂, ATS Conference, AUMNH]; Upton: just S of Hwy 329, before FM1492 jct, 31.42758333 -102.1889667 2, 2800ft., [APH_1378, 10/8/2011, 1 juv, Darryl Burton, AUMNH]; oil fields E of Hwy 329, 31.35481944 -102.0862667 2, 2715ft., [APH_1382, 10/9/2011, 1♀, Darryl Burton, AUMNH]; [APH_1383, 30/8/2011, 1♀, Darryl Burton, AUMNH]; oil fields W of Hwy 329, 31.42420833 -102.1828 2, 2795ft., [APH_1376, 30/7/2011, 1 juv, Darryl Burton, AUMNH]; [APH_1380, 6/9/2011, 1 juv, Darryl Burton, AUMNH]; [APH_1390, 31/7/2011, 1 juv, Darryl Burton, AUMNH]; Ward: E of Pecos on I-20, 31.444196 -103.371451 1, 2583ft., [APH_1175, 22/7/2010, 1♂, Brent E. Hendrixson, Brendon Barnes, Nate Davis, AUMNH.

Distribution and natural history

Aphonopelma gabeli is distributed mostly throughout the Chihuahuan Desert in southeastern Arizona, southern New Mexico, and West Texas; this includes the northern finger-like extensions of the desert along the Rio Grande and Pecos Rivers into Socorro and Chaves Counties, New Mexico, respectively (Fig. 49A). This species is also known from adjacent sections of the High Plains and near canyon mouths of the Madrean sky islands. The species distribution model (Fig. 49B) predicts suitable habitat throughout most of Trans-Pecos Texas, the boot heel of New Mexico, and sections of southeastern Arizona, northeastern Sonora, and northwestern Chihuahua. Aphonopelma gabeli can be found inhabiting the following Level III Ecoregions: Chihuahuan Deserts, High Plains, Arizona/New Mexico Mountains, Southwestern Tablelands, and Madrean Archipelago (Fig. 1F). Specimens accompanied with precise georeferenced locality data have been collected at elevations ranging from 775 to 1620 meters in short grass prairie, desert grassland, and desert scrub communities; at higher elevations, habitats are sometimes associated with various oaks and junipers. Aphonopelma gabeli has been observed in syntopy (burrows located within a few meters of each other) with A. armada, A. hentzi, A. parvum sp. n., and A. vorhiesi and can probably be found alongside A. chalcodes in Cochise and Graham Counties, Arizona. Burrows are typical of that for North American tarantulas (i.e., circular and generally covered by a thin veil of silk) and specimens can be readily collected by pouring a small amount of water into their burrows. Burrows are plugged during the winter months. The breeding period is late spring and early summer (June-August); adult males have been observed in large numbers at night along dirt roads in Graham County, Arizona and Eddy County, New Mexico. The data in Hamilton et al. (2011) and other preliminary mtDNA data suggests that A. gabeli recently expanded its distribution into present-day portions of the Chihuahuan Desert, likely following the desert’s appearance in the United States as the climate warmed and dried during the late Holocene (see Van Devender 1990).

Figure 49.

Aphonopelma gabeli Smith, 1995. A distribution of known specimens B predicted distribution; warmer colors (red, orange, yellow) represent areas of high probability of occurrence, cooler colors (blue shades) represent areas of low probability of occurrence.

Conservation status

Landscape fragmentation due to oil and natural gas production has raised concern about the conservation status of some species in the Permian Basin of West Texas and southeastern New Mexico (Leavitt and Fitzgerald 2013). Aphonopelma gabeli, however, is one of the most common and widely distributed tarantulas in the United States and is frequently encountered in areas that have been developed for such production (D. Burton 2011, pers. comm.). This species is secure.

Remarks

Aphonopelma gabeli females and juvenile males are easily differentiated from A. armada by the flared metatarsal scopulae and prolateral coxa I setae of A. armada; from A. anax by spermathecae and palpal bulbs, as well as prolateral coxa I setae; from A. hentzi by its phenotypic appearance and prolateral coxa I setae; from A. moderatum by their unique phenotypic color and banding; from A. peloncillo by its phenotypic appearance; from A. vorhiesi by their black appearance; and from A. parvum due to the extreme small size of the miniature species. Other important ratios that distinguish males: A. gabeli possess a larger F4L/W (≥4.29; 4.29–4.60) than A. hentzi (≤4.24; 3.62–4.24); by possessing a smaller L1/L4 (≤0.95; 0.91–0.95) than A. moderatum (>0.95; 0.95–0.99) and A. moellendorfi (≥0.96; 0.96–1.00); by possessing a larger L4 scopulation extent (36%–47%) than A. vorhiesi (20%-36%) and smaller than A. chalcodes (66%–76%); by possessing a smaller F1/M3 (≤1.24; 1.18–1.24) than A. anax (≥1.28; 1.28–1.43), A. armada (≥1.26; 1.26–1.33), and A. peloncillo (≥1.33; 1.33–1.49). Other important ratios distinguish females: A. gabeli possess a larger T3/T4 (≥0.73; 0.73–0.79) than A. armada (≤0.73; 0.64–0.73) and A. moderatum (≤0.71; 0.63–0.71); by possessing a larger L3 scopulation extent (72%-80%) than A. peloncillo (58%–68%) and A. vorhiesi (49%–69%); by possessing a larger CL/CW (≥1.14; 1.14–1.20) than A. chalcodes (≤1.14; 1.09–1.14). For both males and females, certain morphometrics have potential to be useful, though due to the amounts of variation, small number of specimens, and the small differences between species, no others are claimed to be significant at this time (see Suppl. material 2). During evaluation of traditional PCA morphospace, males of A. gabeli separate from A. anax, A. armada, A. hentzi, and A. vorhiesi along PC1~2, but do not separate from A. moderatum, A. moellendorfi, A. chalcodes, or A. peloncillo. Female A. gabeli do not separate from any of their syntopic species or phylogenetic sister lineages in PCA morphospace. Females of A. moellendorfi are unknown and cannot be compared. Interestingly, A. gabeli males separate from A. anax, A. armada, and A. hentzi in three-dimensional PCA morphospace (PC1~PC2~PC3), but do not separate from A. moderatum and A. moellendorfi. Aphonopelma gabeli females separate from A. anax, but do not separate from A. armada, A. hentzi, and A. moderatum. PC1, PC2, and PC3 explain ≥87% of the variation in male analyses and ≥96% of the variation in female analyses. It is also important to note the tremendous variation in spermathecae shape that can be seen across A. gabeli populations (Fig. 48). Previous taxonomic work considered this variation enough to split and describe separate species; this is clearly not an effective character due to the large amounts of subtle variation that is possible. This species was first identified as a new species by Jung (1975) but was not formally described until Smith (1995). The type locality for A. gabeli (“east of Tucson”) is vague and is probably located closer to Willcox or Safford, Arizona.

Aphonopelma hentzi (Girard, 1852)

Figures 50, 51, 52, 53, 54, 55; Suppl. material 4

Mygale hentzii Girard, 1852: 251; male holotype from southwestern Oklahoma, 34.309094 -98.3964947, elev. 968ft., coll. unknown, 1849-1852.

Eurypelma hentzi Simon, 1891: 322.

Rhechostica hentzi Raven, 1985: 149.

Aphonopelma hentzi Smith, 1995: 107; neotype male and female exemplar from Garfield Co., Oklahoma, 36.436139 -97.8721607, elev. 1264ft., summer 1975, coll. R.L. Lardie; deposited in Oklahoma State University collection. [not examined]

Aphonopelma clarki Smith, 1995: 87; female holotype from Dallas, Dallas Co., Texas, 32.780141 -96.8004517, elev. 421ft., 1968, coll. H.J. Berman; deposited in BMNH. [examined] syn. n.

Dugesiella coloradana Chamberlin, 1940: 35; female holotype from Sugar City, Crowley Co., Colorado, 38.231949 -103.6630016, elev. 4307ft., no collecting date, coll. unknown; deposited in AMNH. [examined]

Rhechostica coloradanum Raven, 1985: 149.

Aphonopelma coloradanum Smith, 1995: 90. syn. n.

Dugesiella echina Chamberlin, 1940: 36; male holotype from Arkansas Valley, Colorado, 38.055115 -103.6217437, elev. 4182ft., 15.xi.1938, coll. unknown; deposited in AMNH. [examined]

Rhechostica echinum Raven, 1985: 149.

Aphonopelma echinum Smith, 1995: 96. syn. n.

Aphonopelma gurleyi Smith, 1995: 104; male holotype from Interstate 35, near Moss Lake, Sherman, Grayson Co., Texas, 33.781417 -97.2216335, elev. 796ft., no collection date, coll. Russ Gurley; deposited in BMNH. [examined] syn. n.

Dugesiella harlingena Chamberlin, 1940: 37; female holotype from Harlingen, Cameron Co., Texas, 26.190631 -97.6961036, elev. 41ft., 1939, coll. Bryce Brown; deposited in AMNH. [examined]

Rhechostica harlingenum Raven, 1985: 149.

Aphonopelma harlingenum Smith, 1995: 106. syn. n.

Aphonopelma odelli Smith, 1995: 126; female holotype from Beavers Bend State Resort Park McCurtain Co., Oklahoma, 34.13104 -94.690045, elev. 670ft., 13.vi.1979, coll. D.C. Arnold; deposited in Oklahoma State University collection. [not examined] syn. n.

Dugesiella wacona Chamberlin, 1940: 38; male holotype from Waco, McClennan Co., Texas, 31.549333 -97.1466706, elev. 498ft., 5.vii.1931, coll. unknown; deposited in AMNH. [examined]

Rhechostica waconum Raven, 1985: 149.

Aphonopelma waconum Smith, 1995: 156. syn. n.

Dugesiella wichitana Chamberlin, 1940: 35; male holotype from Wichita Mtns. National Wildlife Refuge, Comanche Co., Oklahoma, 34.772106 -98.6013086, elev. 1495ft., 5.vii.1928, coll. N.M. Newport; deposited in AMNH. [examined]

Rhechostica wichitanum Raven, 1985: 149.

Aphonopelma wichitanum Smith, 1995: 157. syn. n.

Diagnosis

Aphonopelma hentzi (Fig. 50) is a member of the Hentzi species group and can be identified by a combination of morphological, molecular, and geographic characteristics. Nuclear DNA identifies A. hentzi as a phylogenetically distinct monophyletic lineage (Fig. 8), supported as the sister lineage to A. anax, closely related to A. armada, and exclusive of A. moellendorfi sp. n. (a new species previously confused with A. hentzi). Aphonopelma hentzi can be distinguished from A. anax by the shapes of its spermathecae in females and palpal bulbs in males; from A. armada by the pattern of setae on coxa I and the flared metatarsal scopulae of females and juvenile males; from A. gabeli by its large chelicerae and general phenotypic appearance; from moderatum by their unique phenotypic color and banding; from parvum sp. n. due to the extreme small size; from peloncillo by its phenotypic appearance; and from vorhiesi due to its black appearance. Significant measurements that distinguish male A. hentzi from its closely related phylogenetic and syntopic species are PTl, M1, and the extent of scopulation on metatarsus III. Male A. hentzi can be distinguished by possessing a larger PTl/M1 (≥0.74; 0.74–0.88) than A. gabeli (≤0.68; 0.61–0.68), A. moderatum (≤0.69; 0.61–0.69), and A. moellendorfi (≤0.69; 0.60–0.69); a larger L3 scopulation extent (69%–86%) than A. armada (48%-63%), A. peloncillo sp. n. (52%-68%), and A. vorhiesi (44%–62%); and by possessing a smaller M1/F4 (≤0.77; 0.68–0.77) than A. moderatum (≥0.80; 0.80–0.87) and A. moellendorfi sp. n. (≥0.81; 0.81–0.88). The significant measurement that distinguishes female A. hentzi from syntopic species is the extent of scopulation on metatarsus IV. Female A. hentzi can be distinguished by possessing a larger L4 scopulation extent (42%–72%) than A. peloncillo (32%–38%) and A. vorhiesi (26%–37%). There are no significant measurements that separate female A. hentzi from A. anax, A. armada, A. gabeli, or A. moderatum. Females of A. moellendorfi are unknown and cannot be compared.

Figure 50.

Aphonopelma hentzi (Girard, 1854) specimens, live photographs. Female (L) - APH_0576; Male (R) - APH_3216.

Description

Male and female described by Girard (1852); presumed lost. Neotype designated and redescribed by Smith (1995); new female specimen designated and redescribed by Smith (1995).

Redescription of male exemplar

(APH_0921; Fig. 51). Specimen preparation and condition: Specimen collected live crossing road, preserved in 80% ethanol; deposited in AUMNH; original coloration faded due to preservation. Left legs I, III, IV, and left pedipalp removed for measurements and photographs; stored in vial with specimen. Right leg III removed for DNA and stored at -80°C in the AUMNH (Auburn, AL). General coloration: Generally black or faded to brown. Cephalothorax: Carapace 14.89 mm long, 13.47 mm wide; Hirsute; densely clothed with brown/golden iridescent pubescence appressed to surface; fringe covered in long setae not closely appressed to surface; foveal groove deep and straight; pars cephalica region rises gradually from foveal groove, gently arching anteriorly toward ocular area; AER slightly procurved, PER recurved; normal sized chelicerae; clypeus extends forward on a slight curve; LBl 1.79, LBw 1.93; sternum very hirsute, clothed with long black/brown, densely packed setae. Abdomen: Densely clothed in short black/brown pubescence with numerous longer red/orange setae interspersed; possessing a dense dorsal patch of black Type I urticating bristles (Cooke et al. 1972). Legs: Very hirsute, particularly ventrally; densely clothed in a mix of medium and long black/brown pubescence. Metatarsus I slightly curved. F1 14.11; F1w 3.69; P1 5,94; T1 12.54; M1 10.48; A1 7.60; F3 12.06; F3w 3.71; P3 5.63; T3 9.32; M3 10.85; A3 7.56; F4 14.71; F4w 4.02; P4 5.86; T4 12.17; M4 14.99; A4 8.36; femur III is normal - not noticeably swollen or wider than other legs. All tarsi fully scopulate. Extent of metatarsal scopulation: leg III (SC3) = 80.9%; leg IV (SC4) = 54.3%. Two ventral spinose setae on metatarsus III; nine ventral spinose setae on metatarsus IV. Coxa I: Prolateral surface a mix of fine, hair-like and thick tapered setae. Pedipalps: Very hirsute; densely clothed in the same setal color as the other legs, with numerous longer ventral setae; one spinose seta on the apical, prolateral femur and four spinose setae on the prolateral tibia; PTl 8.525, PTw 2.77. When extended, embolus tapers but gently curves to the retrolateral side near apex; embolus very slender, no keels.

Figure 51.

Aphonopelma hentzi (Girard, 1854). A–I male specimen, APH_0921 A dorsal view of carapace, scale bar = 6mm B prolateral view of coxa I C dorsal view of femur III D ventral view of metatarsus III, scale bar = 4mm E ventral view of metatarsus IV, scale bar = 5mm F prolateral view of L pedipalp and palpal tibia, scale bar = 4.5mm G dorsal view of palpal bulb H retrolateral view of palpal bulb, scale bar = 1mm I prolateral view of tibia I (mating clasper), scale bar = 5mm.

Variation (14). Cl 13.58–18.04 (16.008±0.38), Cw 12.45–17.28 (14.924±0.41), LBl 1.78–2.46 (2.09±0.07), LBw 1.92–2.79 (2.371±0.09), F1 13.1–17.31 (15.509±0.36), F1w 3.19–4.50 (3.991±0.1), P1 5.36–7.63 (6.494±0.18), T1 10.71–14.18 (12.986±0.28), M1 9.15–13.03 (11.118±0.29), A1 6.03–9.13 (7.836±0.26), L1 length 44.35–60.93 (53.943±1.31), F3 10.56–16.83 (13.026±0.42), F3w 3.23–4.71 (3.999±0.11), P3 4.16–7.57 (5.675±0.23), T3 7.89–12.61 (10.126±0.33), M3 9.69–13.93 (11.859±0.33), A3 5.92–8.88 (7.786±0.21), L3 length 38.72–59.23 (48.471±1.42), F4 12.88–17.50 (15.175±0.37), F4w 3.11–4.69 (3.924±0.12), P4 4.36–7.13 (6.046±0.2), T4 10.75–15.16 (12.809±0.32), M4 12.91–17.57 (15.406±0.33), A4 6.64–9.95 (8.803±0.26), L4 length 47.54–66.69 (58.239±1.42), PTl 7.762–10.81 (9.067±0.23), PTw 2.722–3.61 (3.003±0.07), SC3 ratio 0.691–0.866 (0.769±0.01), SC4 ratio 0.324–0.968 (0.579±0.05), Coxa 1 setae = stout/thick tapered, F3 condition = normal/slightly swollen.

Redescription of female exemplar

(APH_0812; Figs 5254). Specimen preparation and condition: Specimen collected live from burrow, preserved in 80% ethanol; deposited in AUMNH; original coloration faded due to preservation. Left legs I, III, IV, and pedipalp removed for photographs and measurements; stored in vial with specimen. Right leg III removed for DNA and stored at -80°C in the AUMNH (Auburn, AL). Genital plate with spermathecae removed and cleared, stored in vial with specimen. General coloration: Faded brown/black, with a very hirsute appearance. Cephalothorax: Carapace 20.89 mm long, 19.82 mm wide; densely clothed with light brown pubescence closely appressed to surface; fringe densely covered in longer setae; foveal groove medium deep and straight; pars cephalica region gently rises from thoracic furrow, arching anteriorly toward ocular area; AER slightly procurved, PER recurved; clypeus extends forward on a slight curve; LBl 2.84, LBw 3.42; sternum very hirsute, clothed with black/brown, long setae. Abdomen: Densely clothed dorsally in short black setae with numerous longer, lighter setae interspersed (generally red or orange in situ); dense dorsal patch of black Type I urticating bristles (Cooke et al. 1972); ventral side with shorter dark brown/black setae. Spermathecae: Uniquely shaped; paired and separate, taper rapidly to a medium width with a secondary bulge and strongly capitate bulbs, with wide bases that are not fused. Legs: Very hirsute, particularly ventrally; densely clothed in long brown pubescence. F1 14.90; F1w 4.85; P1 6.91; T1 11.32; M1 7.95; A1 7.60; F3 12.65; F3w 4.33; P3 6.62; T3 8.82; M3 9.39; A3 7.48; F4 14.91; F4w 4.27; P4 6.34; T4 10.97; M4 13.48; A4 8.65. All tarsi fully scopulate. Extent of metatarsal scopulation: leg III (SC3) = 74.0%; leg IV (SC4) = 54.7%. Two ventral spinose setae on metatarsus III; four ventral spinose setae on metatarsus IV. Coxa I: Prolateral surface a mix of fine, hair-like and thick tapered setae. Pedipalps: Densely clothed in the same setal color as the other legs; one spinose seta on the apical, prolateral femur, one spinose seta on the prolateral patella, and three spinose setae on the prolateral tibia.

Figure 52.

Aphonopelma hentzi (Girard, 1854). A–E female specimen, APH_0812 A dorsal view of carapace, scale bar = 6mm B prolateral view of coxa I C ventral view of metatarsus III, scale bar = 4.5mm D ventral view of metatarsus IV, scale bar = 4.5mm E prolateral view of L pedipalp and palpal tibia.

Figure 53.

Aphonopelma hentzi (Girard, 1854). A–I cleared spermathecae A APH_0052 B APH_0571 C APH_0743 D APH_0812 E APH_0813 F APH_0833 G APH_0838 H APH_0862 I APH_0867.

Figure 54.

Aphonopelma hentzi (Girard, 1854). A–F cleared spermathecae A APH_0868 B APH_0927 C APH_0934 D APH_1063 E APH_1353 F harlingenum holotype.

Variation (15). Cl 12.65–22.36 (17.892±0.68), Cw 11.57–19.87 (16.328±0.61), LBl 1.91–2.86 (2.399±0.08), LBw 2.13–3.42 (2.773±0.1), F1 10.41–17.13 (13.72±0.48), F1w 3.16–5.56 (4.413±0.17), P1 4.55–8.09 (6.4±0.25), T1 7.79–13.28 (10.669±0.32), M1 6.06–10.74 (7.945±0.3), A1 5.17–8.86 (6.671±0.23), L1 length 34.72–57.70 (45.404±1.51), F3 8.32–14.09 (11.281±0.43), F3w 2.79–4.99 (3.804±0.16), P3 3.94–8.06 (5.64±0.31), T3 6.11–10.56 (8.164±0.32), M3 6.46–11.65 (8.915±0.38), A3 5.46–9.03 (6.866±0.26), L3 length 30.84–53.39 (40.818±1.7), F4 10.43–17.15 (13.795±0.44), F4w 3.04–5.16 (4.072±0.15), P4 4.58–7.81 (5.906±0.23), T4 8.66–13.41 (10.851±0.31), M4 8.95–15.84 (11.924±0.47), A4 6.04–9.79 (7.672±0.27), L4 length 39.16–64.0 (50.132±1.68), SC3 ratio 0.671–0.953 (0.805±0.02), SC4 ratio 0.424–0.717 (0.547±0.02), Coxa 1 setae = stout/thick tapered. Spermathecae variation can be seen in Figures 5354.

Material examined

United States: Arkansas: Carroll: near Eureka Springs, 36.404273 -93.735646 5, 1201ft., [APH_2600, unknown, 1♂, H.A. Clay, AMNH]; Cleburne: 6 miles S/SW of Drasco, 35.569318 -91.998288 5, 843ft., [APH_2602, 9/1979, 1♀, D. Pearson, AMNH]; Drew: Monticello, 33.628997 -91.790964 5, 292ft., [APH_2174, 1936, 1♀, unknown, AMNH]; Lawrence: Imboden, 36.202568 -91.174573 5, 308ft., [APH_2196, 1931, 5♂, R.V. Chamberlin, AMNH]; [APH_2203, 1935, 1♀, 1♂, unknown, AMNH]; Marion: between Pyatt and Eros, near Jct Hwy 125 and MC 4054, 36.21402 -92.86224 4, 912ft., [APH_0024, 9/2002, 1♂, Lewonna Nelson, AUMNH]; Washington: Fayetteville, 36.044074 -94.169349 5, 1211ft., [APH_2172-2173, unknown, 2♂, 1♀, W.J. Baerg, AMNH]; [APH_2467, 1921, 2♀, 1♂, W.J. Baerg, AMNH]; Arizona: Greenlee: 0.4 miles N Hwy-75 on Goat Camp Rd, 32.755403 -109.110492 1, 3726ft., [APH_1352-1353, 5/8/2011, 2♀, Brent E. Hendrixson, Brendon Barnes, Nate Davis, Jake Storms, AUMNH]; Colorado: Bent: John Martin Reservoir State Park, 38.07612 -102.957421 1, 3839ft., [APH_0570, 10/6/2009, 1 juv, Brent E. Hendrixson, Courtney Dugas, Sloan Click, AUMNH]; Crowley: 1.5 miles N Hwy-96 on CR-20, S of Lake Henry (near Sugar City), 38.246723 -103.711337 1, 4371ft., [APH_0565, 9/6/2009, 1 juv, Brent E. Hendrixson, Courtney Dugas, Sloan Click, AUMNH]; Fremont: Canon City, Hwy 64, 5 miles N of Junction with Hwy 50, 38.496596 -105.106624 5, 5800ft., [AUMS_3334, 17/8/1987, 1♂, unknown, AUMNH]; Canon City, N toward Cripple Creek, 5 miles N off Hwy 50, 38.497302 -105.291414 5, 6248ft., [AUMS_3338, 17/8/1986, 1♀, T.R. Prentice, AUMNH]; Otero: 1.5 miles SW Hwy-109 on CR-802, in field and bluffs N of road, 37.794004 -103.521272 1, 4425ft., [APH_0560-0564, 9/6/2009, 5♀, Brent E. Hendrixson, Courtney Dugas, Sloan Click, AUMNH]; Commanche Grasslands, La Junta, 37.859983 -103.719267 6, 4440ft., [APH_0917, 10/2009, 1♂, Ken Suzuki, AUMNH]; Prowers: Prowers Co. Rd., S. of Lamar, on Deahl Ranch, 37.892338 -102.60625 6, 3914ft., [AUMS_2469, 4/7/1986, 1♂, T.R. Prentice, AUMNH]; S of Lamar, Deahl Ranch, 38.013237 -102.613274 6, 3804ft., [AUMS_2455, 4/7/1986, 1♀, T.R. Prentice, AUMNH]; Pueblo: 1.5 miles E Boone on Hwy-96, 38.243146 -104.233761 2, 4448ft., [APH_0749, 16/9/2009, 1♂, Jerry Smith, Mario Juvera, AUMNH]; I-25, approx. 25 miles S Pueblo, 37.93879 -104.81027 5, 5908ft., [APH_0022, 7/9/2002, 1♂, Kristin Young, AUMNH]; Piñon Truck Stop, I-25 N of Pueblo, 38.433689 -104.610331 4, 5030ft., [AUMS_2329, 28/6/1986, 1♂, T.R. Prentice, AUMNH]; Pueblo, 0.9 miles N of Hwy-47 on Baculite Mesa Rd, 38.302884 -104.553482 1, 4814ft., [APH_0566-0569, 9/6/2009, 4 juv, Brent E. Hendrixson, Courtney Dugas, Sloan Click, AUMNH]; Kansas: Elk: 1.5 miles W of Longton, 37.377606 -96.108334 4, 935ft., [AUMS_2321, 28/4/1994, 1♂, B. Cutler, AUMNH]; Gove: along Smokey Hill River just W of Hwy 23, 38.74457 -100.50428 4, 2453ft., [APH_0027, 10/2002, 1♂, Alvis Wade, AUMNH]; Wilson: Fredonia, 37.532925 -95.826937 5, 922ft., [APH_2603, 16/8/1964, 1♂, A.R. Moldenke, AMNH]; Fredonia, 2.5 miles SW Hwy 400 and Hwy 47, 37.50556 -95.83278 5, 858ft., [APH_0003, 6/7/2003, 1♀, Jordan B. Johnson, AUMNH]; Louisiana: Allen (Parish): near town of Harmony, off Hwy-1147, 30.59583 -92.91361 2, 57ft., [APH_1171, 24/4/2010, 1 juv, Roland H. Delaune, AUMNH]; Reeves, 2250 Martin Tram Rd, 30.48063 -92.9884 2, 25ft., [APH_0060, 29/10/2006, 1♂, Gary Bellon, AUMNH]; Bossier (Parish): Haughton, 400 ft. N I-20, 1/2 mile W Hwy 614, 32.53917 -93.54944 4, 230ft., [APH_0028, 20/10/2002, 1♂, Jimmy Landen, AUMNH]; Caddo (Parish): Shreveport, 32.525152 -93.750179 6, 159ft., [AUMS_2449, 16/5/1996, 1♂, R. Moore, AUMNH]; East Baton Rouge (Parish): Baton Rouge, 30.457702 -91.14032 6, 49ft., [APH_2610, 1958, 1♂, Leon Rody, AMNH]; La Salle (Parish): Tullos, 31.819261 -92.329862 5, 121ft., [APH_2612, 1/10/1950, 1♂, W.J. Gerstch, AMNH]; Natchitoches (Parish): Kisatchie National Forest, Kisatchie District, along Longleaf Vista Trail, 31.47644 -92.99622 5, 288ft., [APH_0403, 4/10/2008, 1♂, Brent E. Hendrixson, AUMNH]; Vernon (Parish): Near Fort Polk, 31.029236 -93.176675 6, 289ft., [APH_2202, 1957, 1♀, F.E. Potter, AMNH]; Missouri: Greene: Springfield, 37.208957 -93.292299 5, 1280ft., [APH_2183, 1931, 1♂, Truman Smith, AMNH]; [APH_2190, 10/1931, 2♂, Truman Smith, AMNH]; Henry: Hartwell, 38.435258 -93.934662 5, 761ft., [APH_2611, 1937, 1♂, J.K. Connolly, AMNH]; Hickory: 4 miles northwest of Wheatland, 37.969377 -93.427109 5, 991ft., [APH_2609, 2/10/1967, 1♂, W. Ivie, AMNH]; Newton: Joplin , 37.031191 -94.52842 5, 909ft., [APH_2191, 11/10/1942, 4♂, A.B. Gurney, AMNH]; Ozark: Caney Mountain Conservation Area, along Long Bald trail, 36.696831 -92.45143 1, 1193ft., [APH_0591, 25/6/2009, 1♀, Brent E. Hendrixson, AUMNH]; Phelps: Rolla, 37.948543 -91.771535 5, 1093ft., [APH_2175, 1/10/1954, 4♂, L. Bade and Walker, AMNH]; [APH_2177, 10/1960, 1♂, Lee Malone, AMNH]; Ripley: between Calm and Gatewood on Hwy 142, 36.560976 -91.117925 1, 786ft., [APH_0874, 6/2008, 1 juv, Matt Ingrasci, AUMNH]; St. Francois: Bonne Terre, near city lake, 37.912189 -90.55567 4, 866ft., [APH_0199, 30/9/2007, 1♂, Andrew Rieger, AUMNH]; Taney: Bradleyville, edge of floodplains of Breaver Creek, 36.7754 -92.920951 5, 814ft., [APH_2608, 25/9/1955, 1♂, N. and J. Crenshaw, AMNH]; near Branson, Ruth and Paul Henning Conservation Area, 36.6627 -93.293056 1, 1165ft., [APH_0586-0590, 24/6/2009, 5 juv, Brent E. Hendrixson, AUMNH]; New Mexico: Bernalillo: Albuquerque, base of Sandia Mtns at Copper Trail, 35.079247 -106.484022 1, 5948ft., [APH_1437, 12/11/2011, 1♂, Jessica Ashley, AUMNH]; Dona Ana: Aguirre Springs Campground, 32.370504 -106.561475 1, 5662ft., [APH_0539-0540, 5/6/2009, 2♀, Brent E. Hendrixson, Courtney Dugas, Sloan Click, AUMNH]; CR-72, just off I-25, 32.67635 -107.012973 1, 4348ft., [APH_0654, 13/7/2009, 1 juv, Brent E. Hendrixson, Nate Davis, AUMNH]; Eddy: 0.3 miles W US-62/180 on CR 408 (Dark Canyon Rd), 32.28694 -104.28646 1, 3320ft., [APH_0363, 21/6/2008, 1 juv, Shasta Michaels, AUMNH]; 1.7 miles W US-62/180 on CR-408, 32.293874 -104.309514 1, 3392ft., [APH_0544, 6/6/2009, 1 juv, Brent E. Hendrixson, Courtney Dugas, Sloan Click, AUMNH]; Lincoln National Forest, just off Queens Hwy, 32.20002 -104.67976 1, 5583ft., [APH_0522, 31/8/2008, 1♂, Rick C. West, AUMNH]; Hidalgo: 10 miles S I-10 on Hwy-338, 32.13267 -108.87908 2, 4210ft., [APH_0795, 22/10/2009, 1♂, Clyde Mahan, AUMNH]; 2.64 miles W Hwy-338 on Hwy-9, 31.94331765 -108.8509352 2, 4403ft., [APH_0728, 18/8/2009, 1♂, Alice Abela, AUMNH]; 4 miles west of Animas, 31.942746 -108.853622 5, 4400ft., [APH_2628, 26/8/1960, 1♂, unknown, AMNH]; along Hwy-338, 32.226309 -108.879093 1, 4078ft., [APH_1510-1511, 8/9/2012, 2♂, Brent E. Hendrixson, AUMNH]; [APH_1513-1515, 8/9/2012, 3♂, Brent E. Hendrixson, AUMNH]; along Hwy-9, W of Animas, 31.94337666 -108.8874883 2, 4407ft., [APH_0726, 18/8/2009, 1♂, Alice Abela, AUMNH]; [APH_0729, 18/8/2009, 1♂, Alice Abela, AUMNH]; along US-70, SE of Hwy-92, 32.612037 -108.984613 1, 4151ft., [APH_0645, 12/7/2009, 1 juv, Brent E. Hendrixson, Nate Davis, AUMNH]; E side of Animas Mtns, 31.560017 -108.499913 6, 4430ft., [APH_0862, 2006, 1♀, Dave Moellendorf, AUMNH]; NM-9 at Little Hatchet Mtns Rd, W of Hachita, 31.926633 -108.363083 4, 4480ft., [APH_0408-0410, 11/10/2008, 3♂, Kari McWest, Keisha Hendricks, AUMNH]; Luna: 15.5 miles NE Hwy-180 on Hwy-26, 32.430777 -107.566477 1, 4584ft., [APH_1572, 29/10/2012, 1♂, Brent E. Hendrixson, AUMNH]; Rio Arriba: along CR 55, 36.084375 -106.12193 2, 5715ft., [APH_1248, 1/11/2010, 1♂, Chis Tough, AUMNH]; Sandoval: Rt 44 (Rt 448), 3.7 miles west of Placitas, 35.269619 -106.605786 4, 5046ft., [APH_2198, 16/11/1963, 2♂, William A. Shear, AMNH]; Taos: 70 miles N of Santa Fe, 36.650692 -105.636765 7, 7832ft., [AUMS_2476, unknown, 1♂, unknown, AUMNH]; Oklahoma: Bryan: Calera Golf Center, 2.5 miles S Calera on Hwy-75, 33.9094 -96.462 4, 755ft., [APH_0156, 22/7/2007, 1♂, Julianne Smith, AUMNH]; Cimarron: 6.6 miles S Colorado State line on US-287, 36.900429 -102.520409 1, 3924ft., [APH_0555-0559, 8/6/2009, 2♀, 3 juv, Brent E. Hendrixson, Courtney Dugas, Sloan Click, AUMNH]; Oklahoma panhandle, Cimarron River, just N off Hwy 287-385, N of Boise City, 36.945676 -102.521756 6, 3933ft., [AUMS_2472, 1/6/1986, 1♂, T.R. Prentice, AUMNH]; Cleveland: Lake Thunderbird State Park, 35.21208 -97.24742 5, 1085ft., [APH_0052, 25/6/2004, 1♀, Brent E. Hendrixson, AUMNH]; [APH_0150, 25/6/2004, 1♀, Brent E. Hendrixson, AUMNH]; Comanche: In the Wichita Mtns, 3 miles north of Medicine Park, 34.798239 -98.511099 5, 1411ft., [APH_2204, 19/5/1977, 1♀, J.C. Colkendolpher, AMNH]; N side of Wichita Mtns, 34.841533 -98.727542 1, 1771ft., [APH_0571-0574, 11/6/2009, 1♀, 3 juv, Brent E. Hendrixson, Courtney Dugas, Sloan Click, AUMNH]; Wichita National Forest , 37.683896 -97.363593 5, 1302ft., [APH_2176, unknown, 2♂, unknown, AMNH]; Wichita Mtns near Mount Scott, 34.767625 -98.724761 6, 1955ft., [APH_2601, 16/9/1971, 1♀, R. Forbes, AMNH]; Wichita Mtns Wildlife Refuge, 34.709358 -98.623307 6, 1476ft., [APH_2455, 6/9/1975, 1♂, F. Fryce, AMNH]; Wichita Mtns NWR - Comanche Rd., 34.780183 -98.598183 1, 1520ft., [APH_0873, 5/2008, 1 juv, Christian Cox, AUMNH]; Wichita Mtns NWR - Taylor Ranch Rd., 34.841783 -98.728283 1, 1712ft., [APH_0939, 5/2008, 1♀, Christian Cox, AUMNH]; Cotton: in cemetery NW of Walters, approx. 0.4 miles N Hwy-5 on CR-N2610, 34.367655 -98.329618 1, 1029ft., [APH_0575, 11/6/2009, 1 juv, Brent E. Hendrixson, Courtney Dugas, Sloan Click, AUMNH]; Creek: SW of Tulsa, 35.99622 -96.33341 1, 787ft., [APH_0918, 25/4/2008, 1♂, Christian Cox, John Morse, AUMNH]; SW of Tulsa, NE OK off route 33, 35.99622 -96.33341 1, 787ft., [APH_0834, 25/4/2008, 1♀, Christian Cox, John Morse, AUMNH]; Jackson: 498 Honeysuckle ave, Altus, Altus Air Force Base, 34.664112 -99.296041 1, 1372ft., [APH_3124, 15/6/2013, 1♂, David Welch, AUMNH]; Latimer: Along US-270, E of Red Oak, 34.950086 -95.054512 1, 571ft., [APH_0585, 24/6/2009, 1 juv, Brent E. Hendrixson, AUMNH]; unknown, 34.835035 -95.31025 7, 830ft., [APH_2182, 1931, 4♂, unknown, AMNH]; [APH_2193, 6/1931, 1♀, 2♂, 1 juv, unknown, AMNH]; Le Flore: Ouachita Mtns, S of Smithville on US 259, 34.418983 -94.676 1, 736ft., [APH_0921, 10/2008, 1♂, Chris A. Hamilton, AUMNH]; Ouachita Mtns, S on Hwy 259 from Winding Stair Mountain, 34.600767 -94.6987 1, 1701ft., [APH_0920, 10/2008, 1♂, Mike Logan, AUMNH]; Ouachita Mtns, Winding Stair Mountain, 34.7489 -94.8037 1, 2185ft., [APH_0919, 10/2008, 1♂, Brian Fontenot, AUMNH]; Logan: Stillwater , 36.115607 -97.058368 5, 876ft., [APH_2181, 15/9/1969, 1♀, N.V. Horner, AMNH]; McCurtain: 1 mile east of Signal Mountain, Fort Hill, 34.333306 -95.024261 4, 1168ft., [APH_2201, 3/10/1952, 1♀, 2♂, T. Cohn , AMNH]; Oklahoma: Oklahoma City/Edmond at intersection of 150th St and Macarrthur, 35.623642 -97.620781 2, 1100ft., [APH_0402, 21/9/2008, 1♂, Natalie Ellis, AUMNH]; Osage: between Pahuska and Bartlesville (10 miles west of Bartlesville), 36.740071 -96.299699 5, 827ft., [APH_2606, 6/9/1940, 1♀, 1♂, A. Clay, AMNH]; Pontotoc: Ada, 34.773126 -96.678772 5, 1007ft., [APH_2607, 10/1981, 1♂, unknown, AMNH]; Washington: 2 miles W Bartlesville, Circle Mountain, forested area, 36.71598 -95.99562 4, 677ft., [APH_0026, 29/9/2002, 1♂, Jerry Tolbert, AUMNH]; Woodward: Alabaster Cavern- 9 miles south of Freedom, 36.698679 -99.147057 5, 1742ft., [APH_2605, 12/10/1952, 1♂, unknown, AMNH]; Texas: Bastrop: Wolf Lane and Pearce Rd, 30.140783 -97.5673 5, 555ft., [APH_0960-0964, 22/5/2006, 5♂, Dave Moellendorf, AUMNH]; Baylor: 7 miles north of Maybelle, 33.654334 -99.263208 5, 1325ft., [APH_2657, 8/11/1964, 2♂, Karl W. Haller, AMNH]; Brewster: 3.5 miles S US-90 on US-395, 30.15626 -103.23608 1, 4070ft., [APH_1288, 14/5/2011, 1♂, Brent E. Hendrixson, Kate Hall, Austin Deskewies, Alexis Guice, AUMNH]; Big Bend, 29.330733 -103.536367 6, 2592ft., [APH_0933, 5/2008, 1♂, Corey Roelke, AUMNH]; Black Gap WMA, 29.467633 -102.837333 6, 1833ft., [APH_0868, 2006, 1♀, Dave Moellendorf, AUMNH]; [APH_0973, 2006, 1♂, Dave Moellendorf, AUMNH]; [APH_0975, 2006, 1♂, Dave Moellendorf, AUMNH]; Big Bend National Park, 0.55 miles S park boundary on US-385, 29.67449 -103.17149 1, 2910ft., [APH_1289, 15/5/2011, 1♂, Brent E. Hendrixson, Kate Hall, Austin Deskewies, Alexis Guice, AUMNH]; Black Gap WMA, 29.5058 -102.884817 1, 2205ft., [APH_0932, 5/2008, 1♂, Corey Roelke, AUMNH]; [APH_0972, 6/2006, 1♂, Dave Moellendorf, AUMNH]; [APH_0974, 2006, 1♂, Dave Moellendorf, AUMNH]; Black Gap WMA, on FM2627, 29.4867 -102.862733 1, 2009ft., [APH_0931, 5/2008, 1♂, Corey Roelke, AUMNH]; Carson: Pantex Plant, 35.302672 -101.561938 5, 3534ft., [AUMS_2328, 7/10/1997, 1♀, Felix Chavez, AUMNH]; Coleman: O.H. Ivie reservoir, 31.60165 -99.5915 1, 1684ft., [APH_0832, 9/2008, 1♀, Chris A. Hamilton, AUMNH]; Collin: Anna, 2 miles E of junction of FM455 and Hwy 75, 41 miles N of Dallas, 33.344388 -96.552215 5, 659ft., [AUMS_2327, 7/1988, 1♂, SCJ, AUMNH]; [AUMS_2330, 7/1988, 1♂, SCJ, AUMNH]; Cooke: Moss Lake, Sherman, 33.781417 -97.221633 6, 777ft., [APH_3035, unknown, 1♂, Russ Gurley, BMNH]; Coryell: Hwy 84, 14 miles W of Gatesville, 31.470143 -97.977296 5, 1041ft., [AUMS_2616, 27/6/1999, 1♂, M. Buffington, AUMNH]; Dallam: Dalhart, 36.059477 -102.51325 5, 3986ft., [AUMS_3291, unknown, 1♂, unknown, AUMNH]; just NE jct Thompson Grove Ln and Tex-Top Rd, 36.41505 -102.82264 1, 4369ft., [APH_0744, 25/6/2009, 1 juv, Zach Valois, AUMNH]; Dallas: 1455 N. Joe Wilson Road, Cedar Hill, 32.62016 -96.925792 1, 731ft., [APH_3125, 18/6/2013, 1♂, Nikki Miller, AUMNH]; Dallas, 32.802955 -96.769923 6, 515ft., [APH_2200, 5/1938, 1♀, Ottys Sanders, AMNH]; Dallas, Dallas Zoo, 32.73665 -96.815917 1, 484ft., [APH_0839, 5/2008, 1♂, Julie Post, AUMNH]; Dallas, Restland cemetery, 32.924625 -96.744856 1, 593ft., [APH_0576, 12/6/2009, 1 juv, Brent E. Hendrixson, Courtney Dugas, Sloan Click, AUMNH]; [APH_0837-0838, 7/2008, 2♀, Chris A. Hamilton, AUMNH]; [APH_0915-0916, 7/2008, 2♂, Chris A. Hamilton, AUMNH]; [APH_0951, 7/2008, 1♂, Chris A. Hamilton, AUMNH]; Grand Prairie, 32.728783 -96.990383 1, 477ft., [APH_0836, 3/2008, 1♀, Stephanie Rudy, AUMNH]; Gaines: Seminole, 0.4 miles NW 11th St on Hwy-214, 32.729362 -102.661669 1, 3329ft., [APH_0546, 7/6/2009, 1 juv, Brent E. Hendrixson, Courtney Dugas, Sloan Click, AUMNH]; Gray: Pampa, approx. 12 miles S Pampa on US-70, 35.35843 -100.97436 5, 3210ft., [APH_0023, 24/9/2002, 1♂, Kyle Harrison, AUMNH]; Grayson: 6 miles southwest of Pottsboro, 33.71582 -96.73669 5, 666ft., [APH_2664, 1/11/1965, 1♂, Karl W. Haller, AMNH]; Hartley: Smith Ranch, N side of FM-767, 35.67812 -102.72925 1, 3813ft., [APH_0745, 27/6/2009, 1 juv, Zach Valois, AUMNH]; Hays: Buda, Buda High School, 30.03265 -97.89275 4, 848ft., [APH_0910, 7/2008, 1♂, Dave Moellendorf, AUMNH]; Kyle, 29.995 -97.893867 5, 801ft., [APH_0835, 7/2006, 1♀, Dave Moellendorf, AUMNH]; [APH_0872, 2006, 1♀, Dave Moellendorf, AUMNH]; unknown rd N of FM-3296, 35.82407 -102.95766 1, 4122ft., [APH_0743, 26/6/2009, 1 juv, Zach Valois, AUMNH]; Hill: 5 miles north of Itasca, 32.22622 -97.156988 5, 659ft., [APH_2185, 16/7/1958, 1♂, P. Carne Jr., AMNH]; Jeff Davis: Davis Mtns, 30.8044 -103.8923 1, 4087ft., [APH_0936, 31/9/2008, 1♂, Christian Cox, AUMNH]; Davis Mtns State Park, 30.592854 -103.940197 1, 5045ft., [APH_0536, 4/6/2009, 1♀, Brent E. Hendrixson, Courtney Dugas, Sloan Click, AUMNH]; Davis Mtns, Hwy 118 between McDonald observatory and Davis Mtns State Park, 30.6638 -104.0317 1, 6154ft., [APH_0934, 31/9/2008, 1♀, Christian Cox, AUMNH]; Davis Mtns, Hwy 118 between McDonald observatory and Hwy 166, 30.7049 -104.1118 1, 5905ft., [APH_0935, 31/9/2008, 1♂, Christian Cox, AUMNH]; Davis Mtns, Hwy 166 at the foothills of the Davis Mtns, 30.63155 -104.278117 1, 5393ft., [APH_0937, 31/9/2008, 1♂, Corey Roelke, AUMNH]; Fort Davis, 30.588211 -103.894625 5, 4895ft., [APH_2466, 5/1938, 2♂, unknown, AMNH]; Kimble: CR182, Junction, 30.470278 -99.756389 1, 1743ft., [APH_1066, 15/6/2010, 1♀, Skyler Stevens, AUMNH]; McCulloch: Brady, Brady Lake, 31.1235 -99.384517 1, 1759ft., [APH_0827-0831, 7/2008, 4♀, 1 juv, Chris A. Hamilton, AUMNH]; McLennan: Waco, near Lake Waco, 31.553483 -97.203633 5, 559ft., [APH_0914, 2006, 1♂, Dave Moellendorf, AUMNH]; Oldham: Boys Ranch area, Hwy 385, S of Canadian River, 35.500017 -102.25236 5, 3222ft., [AUMS_2474, 25/10/1986, 1♂, T.R. Prentice, AUMNH]; Parmer: 2 miles north of Friona, 34.670063 -102.72741 4, 4029ft., [APH_2184, 5/10/1961, 1♂, W.J. Gertsch, AMNH]; Presidio: Terlingua, 29.323867 -103.617117 6, 3020ft., [APH_0863, 2006, 1♀, Dave Moellendorf, AUMNH]; unknown, 29.948067 -104.100133 7, 4310ft., [APH_0870, 2006, 1♀, Dave Moellendorf, AUMNH]; Randall: Canyon, field due west of Canyon Crest Apts., 34.9703 -101.92336 2, 3602ft., [APH_0041, 24/5/2002, 1♀, Brent E. Hendrixson, AUMNH]; FM Rd 1151 W jct. Whitaker Rd near playa lake, 35.09147 -101.75591 4, 3573ft., [APH_0035, 26/9/2006, 1♂, Kari McWest, AUMNH]; San Saba: Colorado Bend State Park, 31.095817 -98.501567 1, 1185ft., [APH_0822-0826, 6/2008, 5♀, Chris A. Hamilton, AUMNH]; [APH_0909, 6/2008, 1♀, Chris A. Hamilton, AUMNH]; [APH_0976, 6/2008, 1 juv, Chris A. Hamilton, AUMNH]; Sherman: 10 miles south of Texhoma, 36.399192 -101.803752 5, 3510ft., [APH_2645, 9/1967, 1♂, W. Ivie, AMNH]; 6 miles south of Stratford, 36.284667 -102.049499 5, 3655ft., [APH_2662, 9/1967, 1♂, W. Ivie, AMNH]; Tarrant: 537 Jakmar Rd, Benbrook, 32.6549 -97.47915 2, 800ft., [APH_0361, 6/2008, 1♂, Laura Gleason, AUMNH]; Travis: 1907 Casa Grande, Austin, 30.34272 -97.86449 2, 673ft., [APH_0015, 14/9/2002, 1♂, Mike Comai, AUMNH]; 5203 Rico Cove, Austin, 30.374853 -97.77368 2, 885ft., [APH_0401, 5/10/2008, 1♂, Rhone McCall, AUMNH]; Austin, 30.303983 -97.839883 6, 991ft., [APH_0875, 7/2008, 1♀, Dave Moellendorf, AUMNH]; Austin - City Park Rd and Pence Ln, 30.354083 -97.832567 1, 843ft., [APH_0812, 6/2008, 1♀, Chris A. Hamilton, AUMNH]; [APH_0904, 6/2008, 1♂, Chris A. Hamilton, AUMNH]; Austin - Cuernavaca area in West Lake Hills, 30.34478 -97.85278 1, 623ft., [APH_0159, Summer/2007, 1♂, Chris A. Hamilton, AUMNH]; [APH_0816-0819, 6/2008, 4♀, Chris A. Hamilton, AUMNH]; Austin - Red Bud trail in West Lake Hills, 30.2984 -97.80258 1, 784ft., [APH_0158, Summer/2007, 1 juv, Chris A. Hamilton, AUMNH]; [APH_0813-0815, 6/2008, 3♀, Chris A. Hamilton, AUMNH]; [APH_0906-0907, 6/2008, 2♂, Chris A. Hamilton, AUMNH]; Austin area, along Wild Basin Ledge, 30.307721 -97.818971 2, 774ft., [APH_0748, 5/9/2009, 1♂, Claudia Cobianchi, AUMNH]; Austin Baseball Field, 30.267153 -97.743061 6, 459ft., [APH_2186, 29/5/1958, 1♀, W.H. McAlister, AMNH]; [APH_2187, 21/7/1958, 1♂, W.H. McAlister, AMNH]; Austin, Duvall and Jolleyville Rd, 30.417283 -97.750033 6, 882ft., [APH_0911, 2006, 1♂, Dave Moellendorf, AUMNH]; Austin, near 290 and 71 split, 30.2425 -97.874367 6, 941ft., [APH_0876, 7/2008, 1♀, Dave Moellendorf, AUMNH]; Jonestown - Reed Park, 30.4779 -97.94605 1, 860ft., [APH_0160, Summer/2007, 1♀, Chris A. Hamilton, AUMNH]; [APH_0820-0821, 6/2008, 2♀, Chris A. Hamilton, AUMNH]; [APH_0908, 6/2008, 1♂, Chris A. Hamilton, AUMNH]; [APH_0949, 6/2008, 1♀, Chris A. Hamilton, AUMNH]; Val Verde: Comstock, 29.685683 -101.17145 1, 1580ft., [APH_0833, 7/2008, 1♀, Chris A. Hamilton, AUMNH]; Del Rio - Lake Amistad, 29.496167 -101.04455 5, 1147ft., [APH_0927, 2006, 1♀, Dave Moellendorf, AUMNH]; Del Rio, 29.370886 -100.895867 6, 995ft., [APH_0971, 2006, 1♂, Dave Moellendorf, AUMNH]; Del Rio, on spur 454, 29.460165 -100.948547 5, 1139ft., [APH_0957, 6/2006, 1♂, Dave Moellendorf, AUMNH]; Langtry, 29.809722 -101.5525 1, 1272ft., [APH_1063, 18/6/2010, 1♀, Travis Fisher, AUMNH]; [APH_0958, 6/2006, 1♂, Dave Moellendorf, AUMNH]; Langtry, Hwy 90, 29.814633 -101.563317 1, 1372ft., [APH_0867, 5/2008, 1♀, Corey Roelke, AUMNH]; N of Langtry on Pandale Dirt Rd, 29.828883 -101.59095 6, 1425ft., [APH_0926, 2006, 1♂, Dave Moellendorf, AUMNH]; Pandale, 30.130017 -101.575733 1, 1607ft., [APH_1070, 2/6/2010, 1♂, Lynn McCutchen, AUMNH]; [APH_1071, 26/6/2010, 1♂, Roxana Leija, AUMNH]; Wichita: 4 miles southeast of Burkburnett, 34.067561 -98.541374 5, 994ft., [APH_2665, 4/7/1975, 1♂, J. Cokendolpher, AMNH]; unknown, 33.930965 -98.748117 7, 1007ft., [APH_2199, 23/4/1975, 1♀, M. Nipper, AMNH]; Wilbarger: 12 miles west of Electra, 34.025734 -99.145057 5, 1171ft., [APH_2179, 3/9/1998, 1♀, M.E. Janowski-Bell, AMNH]; [APH_2192, 3/9/1998, 1♂, M.E. Janowski-Bell, AMNH]; Williamson: Cedar Park, 30.5015 -97.847917 5, 1018ft., [APH_0905, 2006, 1♂, Dave Moellendorf, AUMNH]; Core Hole Cave- 1 mile south Georgetown, 30.607738 -97.687453 5, 787ft., [APH_2555, 3/11/1963, 1♀, J. Reddell, D. McKenzie, John Porter, AMNH]; Leander, 30.57295 -97.850467 5, 972ft., [APH_0896-0897, 2006, 2♂, Dave Moellendorf, AUMNH]; Round Rock, Fern Bluff Elementary, 30.517917 -97.719917 4, 806ft., [APH_0912, 6/2008, 1♂, Dave Moellendorf, AUMNH]; Taylor, 30.560167 -97.413333 5, 518ft., [APH_0913, 6/2008, 1♂, Dave Moellendorf, AUMNH]; Yoakum: unknown rd N of Hwy-380, 33.35231 -103.01736 1, 3857ft., [APH_0746, 17/6/2009, 1 juv, Zach Valois, AUMNH].

Distribution and natural history

Aphonopelma hentzi is the most widely distributed species in the United States. Its distribution is bound to the east by the Mississippi River but these tarantulas are found in all or parts of Missouri, Arkansas, Louisiana, Kansas, Oklahoma, Texas, Colorado, New Mexico and Arizona (Fig. 55). Aphonopelma hentzi thrives in a variety of habitats and can be found inhabiting the following Level III Ecoregions: Interior River Valleys and Hills, Ozark Highlands, Central Irregular Plains, Alluvial Plain, South Central Plains, Ouachita Mountains, Arkansas Valley, Boston Mountains, Western Gulf Coastal Plains, Flint Hills, Central Oklahoma/Texas Plains, Western High Plains, Southwestern Tablelands, Central Great Plains, Cross Timbers, East Central Texas Plains, Arizona/New Mexico Mountains, Chihuahuan Deserts, Edwards Plateau, Southern Texas Plains, Blackland Prairies, Arizona/New Mexico Plateau, and Madrean Archipelago. Aphonopelma hentzi can be found in syntopy with a number of species across its distribution including A. armada, A. gabeli, A. moderatum, A. moellendorfi sp. n., A. parvum sp. n., A. peloncillo sp. n., and vorhiesi.

Figure 55.

Aphonopelma hentzi (Girard, 1854). A distribution of known specimens B predicted distribution; warmer colors (red, orange, yellow) represent areas of high probability of occurrence, cooler colors (blue shades) represent areas of low probability of occurrence. Of note, the lone south Texas outlier specimen in A is the harlingenum holotype.

Aphonopelma hentzi exhibits significant variation in burrowing behavior across its distribution. This species is known to inhabit freestanding burrows or scrapes (burrows under rocks, wood, etc.; see Fig. 2A–C) depending on the habitat. Aphonopelma hentzi inhabit a wide range of habitats and elevations and exhibit phenotypic differences between those (e.g., large and robust in lower elevations or plains habitats and a smaller body with longer, thinner legs in rocky or higher elevation habitats). Depending on the locality, A. hentzi can exhibit two different breeding periods: a spring/summer season (April-August) and a fall season (September-October), sometimes at the same location. Additional information about the natural history of A. hentzi is provided by Smith (1995).

Conservation status

Aphonopelma hentzi would be considered rare in Arizona due to its very limited distribution in the state but this species is probably the most abundant tarantula in the United States. These spiders thrive in many different habitats and are not uncommon in major metropolitan areas. More than 40 individuals were collected by one of the authors (BEH) from a single residential lawn in suburban Fort Worth, Texas (specimens not reported). Specimens have also been collected from well-maintained grounds at a cemetery just outside Downtown Dallas, Texas near a busy freeway. This species is secure.

Remarks

Other important ratios that distinguish males: A. hentzi possess a larger L3 scopulation extent (69%-86%) than A. armada (48%-63%); by possessing a larger F4/M4 (≥0.92; 0.92–1.03) than A. armada (≤0.92; 0.86–0.92); by possessing a larger PTl/T4 (>0.65; 0.65–0.77) than A. gabeli (≤0.64; 0.58–0.64). Other important ratios distinguish females: A. hentzi possess a larger L4 scopulation extent (42%-72%) than A. armada (28%–43%, with slight overlap); by possessing a smaller L1/L3 (≤1.14; 1.07–1.14) than A. moderatum (>1.14; 1.14–1.24, with slight overlap). For both males and females, certain morphometrics have potential to be useful, though due to the amounts of variation, small number of specimens, and the small differences between species, no others are claimed to be significant at this time (see Suppl. material 2). During evaluation of traditional PCA morphospace, males of A. hentzi separate in PCA morphological space from A. gabeli, A. moderatum, and A. moellendorfi along PC1~2, but do not separate from A. anax, A. armada, A. peloncillo, or A. vorhiesi. Female A. hentzi do not separate from A. anax, A. armada, A. gabeli, A. moderatum, A. peloncillo, or A. vorhiesi. Females of A. moellendorfi are unknown and cannot be compared. Interestingly, A. hentzi males separate from A. gabeli, A. marxi, A. moderatum, A. moellendorfi, A. parvum, and A. peloncillo in three-dimensional PCA morphospace (PC1~PC2~PC3), but do not separate from A. anax, A. armada, A. chalcodes, or A. vorhiesi. Aphonopelma hentzi females separate from A. marxi and A. parvum, but do not separate from A. anax, A. armada, A. chalcodes, A. gabeli, A. moderatum, A. peloncillo, or A. vorhiesi. PC1, PC2, and PC3 explain ≥87% of the variation in male analyses and ≥96% of the variation in female analyses.

Of particular note, Warriner (2008) determined that the type locality for A. hentzi was near the Wichita Mountains.in Comanche County, Oklahoma where the Marcy Expedition likely camped in May 1852. Even though we have examined material from this area, we have chosen not to designate a new neotype at this time because Smith’s (1995) neotype is conspecific with this material and we were unable to secure adults. We examined most of the holotypes and/or freshly collected topotypic material of A. clarki, A. coloradanum, A. echinum, A. gurleyi, A. harlingenum, A. odelli, A. waconum, and A. wichitanum. Our morphological and molecular analyses fail to recognize these eight species as separate, independently evolving lineages. As a consequence, we consider A. clarki, A. coloradanum, A. echinum, A. gurleyi, A. harlingenum, A. odelli, A. waconum, and A. wichitanum junior synonyms of A. hentzi. It is also important to note that the paratype of A. harlingenum that is found in the same jar as the A. harlingenum holotype is a female of A. anax (see spermathecae in Fig. 12C).

Mitochondrial DNA (CO1) identifies A. hentzi as a polyphyletic group with respect to A. moderatum, A. anax, and A. armada (Fig. 7). A second lineage of A. hentzi is embedded within a secondary lineage of A. anax; both lineages were previously identified as putative cryptic species (Hamilton et al. 2014). Based on the AE results, it appears that CO1 is not effective at delimiting species boundaries within this group.

Aphonopelma icenoglei Hamilton, Hendrixson & Bond, sp. n.

Figures 56, 57, 58, 59, 60

Aphonopelma mojave Prentice, 1997 (in part): 161.

Types

Male holotype (APH_2396) collected 7.5 miles north of Pipes Canyon Rd., San Bernardino Co., California, 34.303363 -116.440492 5, elev. 3133ft., 14.iv.1991, coll. T.R. Prentice; deposited in AMNH. Paratype female (APH_1562) from 0.6 miles S Hwy-62 on La Contenta Rd., San Bernardino Co., California, 34.126565 -116.368852 1, elev. 3170ft., 25.x.2012, coll. Brent E. Hendrixson; deposited in AUMNH. Paratype male (AUMS_2643) from near junction of Main Rd. and past Cottonwood Road by Eagle Mtn. mine, 33.828312 -115.756448 5, elev. 2476ft., xii.1999, coll. unknown; deposited in AUMNH. Paratype female (APH_2393) from Apple Valley, 2 miles south of Highway 18, Milpas Dr., San Bernardino Co., California, 34.53717 -117.103251 4, elev. 3140ft., 5.v.1992, coll. T.R. Prentice; deposited in AMNH.

Etymology

The specific epithet is a patronym in recognition of Wendell Icenogle, an arachnologist and prolific collector of North American mygalomorph spiders. This work benefitted substantially from his help collecting specimens and his wealth of knowledge concerning tarantulas in the United States.

Diagnosis

Aphonopelma icenoglei (Fig. 56) is a member of the Paloma species group and can be distinguished by a combination of morphological, molecular, and geographic characteristics. Nuclear and mitochondrial DNA identifies A. icenoglei as a strongly supported monophyletic lineage (Figs 78) that is a sister lineage to A. mojave, A. atomicum sp. n., A. prenticei sp. n., and A. joshua. Aphonopelma icenoglei can easily be differentiated from A. iodius by its smaller size and limited extent of scopulation on metatarsus IV, and from A. atomicum and A. prenticei by locality. The most significant measurements that distinguish male A. icenoglei from its closely related phylogenetic and syntopic species are F3 and the extent of scopulation on metatarsus IV. Male A. icenoglei can be distinguished by possessing a larger F3L/W (≥3.51; 3.51–3.90) than A. atomicum (≤3.40; 2.92–3.40) and A. prenticei (≤3.27; 2.76–3.27); a larger PTl/F3 (≥0.67; 0.67–0.71) than A. joshua (≤0.61; 0.55–0.61) and A. xwalxwal sp. n. (≤0.63; 0.59–0.63); and a smaller L4 scopulation extent (31%-46%) than A. iodius (62%-88%). There are no significant measurements that separate male A. icenoglei from A. mojave. The most significant measurements that distinguish female A. icenoglei from its closely related phylogenetic and syntopic species are M4 and the extent of scopulation on metatarsus IV. Female A. icenoglei can be distinguished by possessing a larger F1/M4 (≥1.07; 1.07–1.23) than A. joshua (≤1.04; 0.98–1.04); a larger A3/M4 (≥0.63; 0.63–0.70) than A. atomicum (≤0.58; 0.57–0.58) and A. joshua (≤0.59; 0.50–0.59); a smaller Cl/F1 (≤1.16; 1.07–1.16) than A. prenticei (≥1.17; 1.17–1.33); and a smaller L4 scopulation extent (27%–42%) than A. iodius (59%–83%). There are no significant measurements that separate female A. icenoglei from A. mojave. Aphonopelma icenoglei is most similar (morphologically and geographically) to A. mojave but these two species are phylogenetically distinct and probably are not sister taxa (Figs 7, 8).

Figure 56.

Aphonopelma icenoglei sp. n. live photograph. Female - APH_3146. Do not have a photograph of a live male specimen.

Description of male holotype

(APH_2396; Fig. 57). Specimen preparation and condition: Specimen originally collected from burrow and preserved in unknown percentage of ethanol; original coloration faded due to preservation. Left legs I, III, IV, and left pedipalp removed for measurements and photographs; stored in vial with specimen. No tissue for DNA. General coloration: Faded black/brown. Cephalothorax: Carapace 8.228 mm long, 6.965 mm wide; densely clothed with faded pubescence, appressed to surface; fringe covered in long setae not closely appressed to surface, hirsute appearance; foveal groove medium deep and straight; pars cephalica region rises very gradually from foveal groove on a straight plane towards the ocular area; AER very slightly procurved, PER recurved; normal sized chelicerae; clypeus extends slightly on a curve; LBl 1.051, LBw 1.468; sternum hirsute, clothed with faded, densely packed, short setae. Abdomen: Densely clothed in short black/brown pubescence with numerous longer, lighter setae interspersed (generally red or orange in situ); dense dorsal patch of black Type I urticating bristles (Cooke et al. 1972) - smaller and distinct from large species. Legs: Hirsute; densely clothed in faded pubescence. Metatarsus I straight. F1 7.605; F1w 1.786; P1 2.994; T1 6.779; M1 5.848; A1 4.17; F3 7.202; F3w 1.851; P3 2.613; T3 5.625; M3 6.742; A3 4.464; F4 8.432; F4w 1.872; P4 2.869; T4 7.231; M4 8.368; A4 4.938; femur III is normal. All tarsi fully scopulate. Extent of metatarsal scopulation: leg III (SC3) = 62.6%; leg IV (SC4) = 35.1%. Two ventral spinose setae on metatarsus III; five ventral spinose setae on metatarsus IV; one prolateral spinose seta on tibia I; one megaspine on the apex on the retrolateral branch of the tibial apophyses. Coxa I: Prolateral surface covered by fine, hair-like setae. Pedipalps: Hirsute; densely clothed in the same setal color as the other legs, with numerous longer ventral setae; one spinose seta at the apical, prolateral femur and three prolateral spinose setae on the palpal tibia; PTl 4.938, PTw 1.77. When extended, embolus tapers with a curve to the retrolateral side; embolus slender, no keels; distinct dorsal and ventral transition from bulb to embolus.

Figure 57.

Aphonopelma icenoglei sp. n. A–I male holotype, APH_2396 A dorsal view of carapace, scale bar = 2.5mm B prolateral view of coxa I C dorsal view of femur III D ventral view of metatarsus III, scale bar = 2.5mm E ventral view of metatarsus IV, scale bar = 2.5mm F prolateral view of L pedipalp and palpal tibia, scale bar = 3mm G dorsal view of palpal bulb H retrolateral view of palpal bulb, scale bar = 0.5mm I prolateral view of tibia I (mating clasper), scale bar = 2.5mm.

Variation (5). Cl 7.326–8.228 (7.831±0.15), Cw 6.609–7.307 (6.976±0.11), LBl 1.031–1.069 (1.046±0.01), LBw 1.092–1.475 (1.271±0.08), F1 7.605–8.996 (8.292±0.24), F1w 1.703–1.938 (1.821±0.05), P1 2.909–3.301 (3.067±0.07), T1 6.679–7.727 (7.204±0.21), M1 5.848–7.032 (6.459±0.22), A1 4.012–4.818 (4.338±0.14), L1 length 27.396–31.441 (29.36±0.78), F3 6.881–7.558 (7.203±0.14), F3w 1.763–2.145 (1.963±0.08), P3 2.477–2.771 (2.579±0.05), T3 5.625–6.187 (5.88±0.12), M3 6.742–7.609 (6.986±0.16), A3 4.448–4.952 (4.569±0.1), L3 length 26.376–28.76 (27.216±0.48), F4 7.919–9.094 (8.46±0.22), F4w 1.691–1.902 (1.836±0.04), P4 2.549–2.869 (2.77±0.06), T4 6.914–7.825 (7.283±0.16), M4 8.368–9.267 (8.657±0.17), A4 4.702–5.595 (5.036±0.16), L4 length 30.775–33.887 (32.205±0.65), PTl 4.758–5.115 (4.953±0.07), PTw 1.548–1.77 (1.684±0.04), SC3 ratio 0.626–0.788 (0.704±0.03), SC4 ratio 0.312–0.456 (0.376±0.02), Coxa I setae = very thin tapered, F3 condition = normal.

Description of female paratype

(APH_1562; Figs 5859). Specimen preparation and condition: Specimen collected live from burrow, preserved in 80% ethanol; original coloration faded due to preservation. Left legs I, III, IV, and pedipalp removed for photographs and measurements; stored in vial with specimen. Right legs III & IV removed for DNA and stored at -80°C in the AUMNH (Auburn, AL). Genital plate with spermathecae removed and cleared, stored in vial with specimen. General coloration: Faded black/brown. Cephalothorax: Carapace 7.943 mm long, 6.522 mm wide; Hirsute, densely clothed with short faded black/brown pubescence closely appressed to surface; fringe densely covered in slightly longer setae; foveal groove medium deep and slightly procurved; pars cephalica region gently rises from thoracic furrow, arching anteriorly toward ocular area; AER slightly procurved, PER very slightly recurved; chelicerae robust, clypeus extends forward on a curve; LBl 1.353, LBw 1.435; sternum hirsute, clothed with short faded setae. Abdomen: Densely clothed dorsally in short faded black setae with longer, lighter setae (generally red or orange in situ) focused near the urticating patch; dense dorsal patch of black Type I urticating bristles (Cooke et al. 1972) - smaller and distinct from large species. Spermathecae: Paired and separate, with capitate bulbs widening towards the bases; not fused. Legs: Hirsute; densely clothed in short faded black/brown pubescence; F1 6.983; F1w 1.974; P1 2.993; T1 5.757; M1 4.266; A1 3.976; F3 5.884; F3w 1.834; P3 2.608; T3 4.443; M3 4.682; A3 4.092; F4 7.463; F4w 1.906; P4 2.765; T4 5.853; M4 6.269; A4 4.678. All tarsi fully scopulate. Extent of metatarsal scopulation: leg III (SC3) = 63.5%; leg IV (SC4) = 27.1%. One ventral and one prolateral spinose seta on metatarsus III; four ventral spinose setae and one prolateral spinose seta on metatarsus IV. Coxa I: Prolateral surface covered by very thin tapered and fine, hair-like setae. Pedipalps: Densely clothed in the same setal color as the other legs; one spinose seta on the apical, prolateral femur, four prolateral (two at the apical, prolateral border with the tarsus) spinose setae and one ventral spinose seta on the tibia.

Figure 58.

Aphonopelma icenoglei sp. n. A–E female paratype, APH_1562 A dorsal view of carapace, scale bar = 3mm B prolateral view of coxa I C ventral view of metatarsus III, scale bar = 2mm D ventral view of metatarsus IV, scale bar = 2.5mm E prolateral view of L pedipalp and palpal tibia.

Figure 59.

Aphonopelma icenoglei sp. n. A–E cleared spermathecae A APH_0761 B APH_0885 C APH_1562 D APH_2393 E APH_3118.

Variation (5). Cl 7.424–8.188 (7.89±0.13), Cw 6.522–7.385 (6.993±0.16), LBl 1.285–1.353 (1.322±0.01), LBw 1.368–1.615 (1.453±0.05), F1 6.565–7.644 (6.966±0.18), F1w 1.974–2.131 (2.049±0.03), P1 2.716–3.161 (2.988±0.08), T1 5.185–6.381 (5.695±0.2), M1 4.163–4.72 (4.406±0.1), A1 3.568–3.976 (3.708±0.08), L1 length 22.432–25.667 (23.763±0.54), F3 5.362–6.315 (5.807±0.16), F3w 1.766–1.979 (1.849±0.04), P3 2.217–2.711 (2.47±0.1), T3 3.995–4.755 (4.305±0.13), M3 4.216–4.864 (4.54±0.11), A3 3.90–4.389 (4.041±0.09), L3 length 20.101–22.92 (21.164±0.51), F4 6.84–8.227 (7.376±0.24), F4w 1.766–1.967 (1.897±0.03), P4 2.625–3.112 (2.836±0.09), T4 5.477–6.325 (5.799±0.14), M4 5.589–6.77 (6.14±0.19), A4 4.174–4.678 (4.436±0.1), L4 length 25.043–28.961 (26.587±0.68), SC3 ratio 0.636–0.726 (0.691±0.02), SC4 ratio 0.272–0.419 (0.365±0.03), Coxa I setae = very thin tapered. Spermathecae variation can be seen in Figure 59.

Material examined

United States: California: Los Angeles: 0.66 miles N Ft Tejon Rd on Valyermo Rd, 34.468812 -117.860377 1, 3532ft., [APH_0756-0757, 4/10/2009, 2 juv, Brent E. Hendrixson, Thomas Martin, AUMNH]; 0.67 miles S Hwy-18 on 263rd St, 34.48904 -117.661058 1, 3461ft., [APH_1202, 30/7/2010, 1♀, Brent E. Hendrixson, Brendon Barnes, Nate Davis, AUMNH]; 1.15 miles N of Valyermo, Bobs Gap Rd, 34.448937 -117.82921 4, 3920ft., [AUMS_2511, 4/10/1992, 1♂, T.R. Prentice, AUMNH]; Piñon Hills off Hwy 138, 34.448134 -117.668735 5, 4002ft., [AUMS_2515, 1993, 1♂, unknown, AUMNH]; Valyermo, 34.446108 -117.852286 5, 4403ft., [APH_2400, 28/10/1989, 1♂, T.R. Prentice, AMNH]; Riverside: Eagle Mountain area, 2 miles E of Cottonwood Springs, 33.733266 -115.784522 5, 3485ft., [AUMS_2464, 4/4/1989, 1♂, T.R. Prentice, AUMNH]; Joshua Tree National Park, 33.82491 -115.720588 6, 2423ft., [APH_0885, 2007, 1♀, Josh Richards, AUMNH]; near junction of Main Rd. and past Cottonwood Road by Eagle Mt. mine, 33.828312 -115.756448 5, 2476ft., [AUMS_2643, 12/1999, 1♂, unknown, AUMNH]; San Bernardino: 0.5 miles SE Mikiska Blvd along Hwy-247 (NW of Landers), 34.319594 -116.48102 1, 3406ft., [APH_0761-0762, 5/10/2009, 2 juv, Brent E. Hendrixson, Thomas Martin, AUMNH]; 0.6 miles S Hwy-62 on La Contenta Rd, 34.126565 -116.368852 1, 3170ft., [APH_1562, 25/10/2012, 1♀, Brent E. Hendrixson, AUMNH]; 1 miles W Ft. Irwin Rd on Irwin Rd, 34.998665 -116.945203 1, 2658ft., [APH_1320, 30/7/2011, 1♀, Brent E. Hendrixson, Brendon Barnes, Nate Davis, Jake Storms, AUMNH]; 1.1 miles NE Lucerne Valley Cutoff along Hwy-247 (Barstow Rd), 34.614513 -116.968992 1, 3239ft., [APH_0760, 5/10/2009, 1 juv, Brent E. Hendrixson, Thomas Martin, AUMNH]; 12.2 miles west of junction of 177 and Highway 62, 34.046346 -115.432997 4, 2205ft., [APH_2391, 12/11/1989, 1♀, T.R. Prentice, AMNH]; [APH_2398, 24/11/1989, 1♂, T.R. Prentice, AMNH]; [APH_2402, 24/11/1989, 1♂, T.R. Prentice, AMNH]; 2.6 miles S Phelan Rd on Baldy Mesa Rd, 34.389404 -117.454546 1, 3904ft., [APH_1577-1578, 6/11/2012, 1♀, 1 juv, Brent E. Hendrixson, AUMNH]; 3.7 miles north of Pipes Canyon Rd. Highway 247 Access Rd., 34.243922 -116.440137 4, 3484ft., [APH_2395, 18/4/1991, 1♀, T.R. Prentice, AMNH]; 7.5 miles north of Pipes Canyon Rd., 34.303363 -116.440492 5, 3133ft., [APH_2396, 14/4/1991, 1♂, T.R. Prentice, AMNH]; along Hwy-62, west of Coxcomb Mtns, 34.090719 -115.424593 1, 1767ft., [APH_1584-1585, 7/11/2012, 2♀, Brent E. Hendrixson, AUMNH]; Apple Valley, 2 miles south of Highway 18, Milpas Dr., 34.53717 -117.103251 4, 3140ft., [APH_2393, 5/5/1992, 1♀, T.R. Prentice, AMNH]; Cadiz Valley, 34.03779 -115.238314 5, 1247ft., [APH_2389, 24/11/1989, 1♂, T.R. Prentice, AMNH]; C