Monograph |
Chris A. Hamilton‡, Brent E. Hendrixson§, Jason E. Bond|
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Corresponding author: Chris A. Hamilton ( chris@8legs2fangs.com ) Academic editor: Jeremy Miller
© 2016 Chris A. Hamilton, Brent E. Hendrixson, Jason E. Bond.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Hamilton CA, Hendrixson BE, Bond JE (2016) Taxonomic revision of the tarantula genus Aphonopelma Pocock, 1901 (Araneae, Mygalomorphae, Theraphosidae) within the United States. ZooKeys 560: 1-340. doi: 10.3897/zookeys.560.6264
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Abstract
This systematic study documents the taxonomy, diversity, and distribution of the tarantula spider genus Aphonopelma Pocock, 1901 within the United States. By employing phylogenomic, morphological, and geospatial data, we evaluated all 55 nominal species in the United States to examine the evolutionary history of Aphonopelma and the group’s taxonomy by implementing an integrative approach to species delimitation. Based on our analyses, we now recognize only 29 distinct species in the United States. We propose 33 new synonymies (A. apacheum, A. minchi, A. rothi, A. schmidti, A. stahnkei = A. chalcodes; A. arnoldi = A. armada; A. behlei, A. vogelae = A. marxi; A. breenei = A. anax; A. chambersi, A. clarum, A. cryptethum, A. sandersoni, A. sullivani = A. eutylenum; A. clarki, A. coloradanum, A. echinum, A. gurleyi, A. harlingenum, A. odelli, A. waconum, A. wichitanum = A. hentzi; A. heterops = A. moderatum; A. jungi, A. punzoi = A. vorhiesi; A. brunnius, A. chamberlini, A. iviei, A. lithodomum, A. smithi, A. zionis = A. iodius; A. phanum, A. reversum = A. steindachneri), 14 new species (A. atomicum sp. n., A. catalina sp. n., A. chiricahua sp. n., A. icenoglei sp. n., A. johnnycashi sp. n., A. madera sp. n., A. mareki sp. n., A. moellendorfi sp. n., A. parvum sp. n., A. peloncillo sp. n., A. prenticei sp. n., A. saguaro sp. n., A. superstitionense sp. n., and A. xwalxwal sp. n.), and seven nomina dubia (A. baergi, A. cratium, A. hollyi, A. mordax, A. radinum, A. rusticum, A. texense). Our proposed species tree based on Anchored Enrichment data delimits five major lineages: a monotypic group confined to California, a western group, an eastern group, a group primarily distributed in high-elevation areas, and a group that comprises several miniaturized species. Multiple species are distributed throughout two biodiversity hotspots in the United States (i.e., California Floristic Province and Madrean Pine-Oak Woodlands). Keys are provided for identification of both males and females. By conducting the most comprehensive sampling of a single theraphosid genus to date, this research significantly broadens the scope of prior molecular and morphological investigations, finally bringing a modern understanding of species delimitation in this dynamic and charismatic group of spiders.
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 (
Morphology-based phylogenies of mygalomorph spiders have revealed widespread patterns of homoplasy among traditional taxonomic characters (
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.
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
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.,
Aphonopelma has a complicated nomenclatural history.
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 (
Abbreviations
Institutional
AUMNH
BMNH
The
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)
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
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
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
To evaluate morphological variation, we examined the morphospace occupied by each species by plotting measurement ratios in boxplots (e.g., Fig.
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
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
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
The analyses described above and recent smaller scale molecular studies (
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
Anchored Enrichment data were collected through the Center for Anchored Phylogenomics at Florida State
Utilizing the bioinformatics pipeline described in
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
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
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 (
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 (
Aphonopelma phylogeny
Following the integrative approach for delimiting species within Aphonopelma outlined in
Maximum Likelihood inferred CO1 gene tree phylogeny for the 1,032 Aphonopelma specimen dataset. Species delimitations followed the integrative methodological approach outlined in
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.
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
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
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.
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 (
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.
When reviewing both the putative mitochondrial and cryptic species identified in
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.
Despite our extensive collecting throughout this region, there likely still remains undescribed diversity in this species group (Figs
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.
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.
Body size is one of the most important determinants of an organism’s ecological role (
Species considered nomina dubia
The species Aphonopelma cratium Chamberlin, 1940 is based on a male holotype and female allotype (both
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
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 (
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 (
Taxonomy
Family Theraphosidae Thorell, 1869
Subfamily Theraphosinae Thorell, 1870
Aphonopelma
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
Aphonopelma
Pocock, 1901: 553 (type species by original designation Eurypelma seemanni Pickard-Cambridge, 1897). First synonymized with Rhechostica by
Delopelma
Petrunkevitch, 1939: 567 (type species by original designation Eurypelma marxi Simon, 1891). First synonymized with Rhechostica by
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
Chaunopelma
Chamberlin, 1940: 30 (type species by original designation Delopelma radinum Chamberlin & Ivie, 1939). First synonymized with Rhechostica by
Apachepelma
Smith, 1994: 45 (type species by original designation Aphonopelma paloma Prentice, 1992). First synonymized with Aphonopelma by
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
“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.
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 |
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 |
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 |
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 |
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
6 Adult females of Aphonopelma moellendorfi remain unknown.
Aphonopelma anax
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.
Aphonopelma anax (Chamberlin, 1940) specimens, live photographs. Female (L) - APH_0524; Male (R) - APH_3122.
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.
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.
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.
Descriptions
Male and female originally described by
Redescription of male exemplar
(APH_0924; Fig.
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
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
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,
Distribution and natural history
In the United States, A. anax is widely distributed throughout South Texas (Fig.
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
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
Mitochondrial DNA (CO1) identifies A. anax as a polyphyletic group with respect to A. armada and A. hentzi (Fig.
Aphonopelma armada
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.
Aphonopelma armada (Chamberlin, 1940) specimens, live photographs. Male (L) - APH_1064; Female (R) - APH_3214.
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.
Description
Female originally described by
Description of male exemplar
(APH_0950; Fig.
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
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
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.
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.
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 , sp. n.
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
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.
Description of male holotype
(AUMS_2727-2; Fig.
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
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.
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
Material examined
United States: California: Inyo:
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.
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
Aphonopelma catalina , sp. n.
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
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.
Description of male holotype
(APH_1440; Fig.
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
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.
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
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 &
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
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 (
Remarks
As noted in
Aphonopelma chalcodes
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
Diagnosis
Aphonopelma chalcodes (Fig.
Redescription of male exemplar
(APH_0954; Fig.
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
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.
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.
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
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,
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.
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
Mitochondrial DNA (CO1) identifies A. chalcodes as a polyphyletic group with some samples more closely related to specimens of A. iodius (Fig.
Aphonopelma chiricahua , sp. n.
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
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.
Description of male holotype
(APH_3191; Fig.
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.
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,
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.
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 (
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
This species was first identified as novel by
Aphonopelma eutylenum
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.
Description
Originally described by
Redescription of male exemplar
(APH_1088; Fig.
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
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.
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
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,
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.
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
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
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.
Description
Male originally described by
Redescription of male exemplar
(APH_1054; Fig.
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
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.
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
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,
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.
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 (
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
Aphonopelma hentzi
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.
Description
Male and female described by
Redescription of male exemplar
(APH_0921; Fig.
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
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.
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.
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
Material examined
United States: Arkansas: Carroll: near Eureka Springs, 36.404273 -93.735646 5, 1201ft., [APH_2600, unknown, 1♂, H.A. Clay,
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.
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.
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
Of particular note,
Mitochondrial DNA (CO1) identifies A. hentzi as a polyphyletic group with respect to A. moderatum, A. anax, and A. armada (Fig.
Aphonopelma icenoglei , sp. n.
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
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.
Description of male holotype
(APH_2396; Fig.
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
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.
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
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,