Content uploaded by Brent Hendrixson
Author content
All content in this area was uploaded by Brent Hendrixson on Feb 09, 2016
Content may be subject to copyright.
A preview of the PDF is not available
Taxonomic revision of the tarantula genus Aphonopelma Pocock, 1901 (Araneae, Mygalomorphae, Theraphosidae) within the United States
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.
Supplementary resources (33)
Data
February 2016
Data
February 2016
Data
February 2016
Data
February 2016
Data
February 2016
Data
February 2016
Data
February 2016
Data
February 2016
Data
February 2016
Data
February 2016
Data
February 2016
Data
February 2016
Data
February 2016
Data
February 2016
Data
February 2016
Data
February 2016
Data
February 2016
Data
February 2016
Data
February 2016
Data
February 2016
Data
February 2016
Data
February 2016
Data
February 2016
Data
February 2016
Data
February 2016
Data
February 2016
Data
February 2016
Data
February 2016
... However, this character has only been treated as a double state character (presence or absence) instead of classifying in relation to the extension of coxae surface and type of modified spinule. Currently there is no standardised terminology for these structures, finding various terms in literature such as ''spike setae'', ''spiniform setae '', ''thorn-like'', and ''stout setae'' (Pérez-Miles 2000;Bertani and Carla-da-Silva 2003;Ferretti et al. 2011;Gallon and Wendt 2015;Hamilton et al. 2016). For this reason, and in relation to the use of the terms spiniform and stout setae for other modified setae, herein coxal spinules is proposed as a unified term with the following description: short spiniform setae with a notorious wider basal section almost pear-shaped and extremely short barbs (see Ferretti et al. 2011: fig. ...
... nov., and potentially in Grammostola doeringi (Holmberg, 1881) ( Fig. 8A; Ferretti et al. 2011: fig. 18) or central and apical section could be shorter or almost differentiable and basal section more wider (e.g., developed coxal spinules in G. schulzei, Thrixopelma spp., and Aphonopelma spp.; Ferretti et al. 2011;Hamilton et al. 2016;pers. obs.). ...
Three new species of Cymbiapophysa Gabriel & Sherwood, 2020 are described from south, central, and northwestern Ecuador, showing the wide range of distribution that this genus has in Ecuador and its biogeographical provinces. These three new species are easily differentiated from other congeners based on keel morphology of the male pal-pal bulb. Supplementary information about the locality of C. magna Sherwood, Gabriel, Brescovit & Lucas, 2021 is provided, alongside additional data on morphology and some commentaries about the general distribution and biogeography of Cymbiapophysa. Additionally, a taxonomic key for males of Cymbiapophysa species is presented, based on the palpal bulb morphology.
... R. C. West] for N. d. dorsalis Cresson from Warsaw, Benton County, Missouri (Table 1; B. Eldridge, pers. comm.), a significant range extension for the wasp (Kurczewski et al. 2020b) and just beyond the northernmost extent in Missouri of the tarantula genus Aphonopelma [hentzi (Girard)] (Theraphosidae) (Hamilton et al. 2016), its natural host. All other host records for this species, as N. dorsalis arizonicus Townes, are for Aphonopelma chal codes Chamberlin juveniles (Minch 1979;Simons 1989;Byrne 2016;Kurczewski et al. 2020b). ...
... (Halonoproctidae) by Notocyphus d. dorsalis in Missouri is easily explained. Its usual host genus Aphonopelma (Mygalomorphae: Theraphosidae) does not occur as far north geographically as the wasp (Hamilton et al. 2016;Kurczewski et al. 2020b). In this case, the wasp attacked a trapdoor, instead of cursorial species of Mygalomorphae, and used it as a substitute host. ...
Host records and nesting behavior of the Nearctic trapdoor spider-hunting spider wasps (Hymenoptera: Pompilidae) Calopompilus Ashmead and Priocnemissus Haupt (Pepsinae: Pepsini) and Aporus Spinola and Psorthaspis Banks (Pompilinae: Aporini) are reviewed, investigated, compared, and discussed. First time incidental trapdoor spider host records for Priocnemis (Priocnemissus) minorata Banks (Pepsinae: Pepsini), Anoplius (Lophopompilus) carolina (Banks) (Pompilinae: Pompilini), and Notocyphus dorsalis
dorsalis Cresson (Notocyphinae: Notocyphini) are included, although they are not typical trapdoor spider-hunting spider wasp species. The Palearctic Aporus (Aporus) unicolor Spinola, A. (Aporus) bicolor Spinola and A. (Aporus) planiceps (Latreille) are referenced for comparison with Nearctic Aporus sensu stricto. Early 20th century papers on species of Aporus and Psorthaspis are revived. New information on nesting behavior of Nearctic trapdoor spider-hunting spider wasps is described and first host trapdoor spider records for Psorthaspis formosa (Smith), P. legata (Cresson) and P. mariae (Cresson) are documented. Potential Pompilidae species in the genera Calopompilus and Aporus are suggested for host trapdoor spider remains found in burrows with spider wasp eggs, larvae and cocoons (pupae) based on geographic distribution, habitat, spider species, trapdoor and burrow structure, wasp cocoon size, and wasp congeneric host records.
... The total body length, including cephalothorax, abdomen, and appendage, was measured using digital vernier calipers to two decimal places in millimeters (mm). Measurements were obtained along the central axis of the structures, beginning on the left side, from the mid-proximal articulation point to the mid-distal articulation point, and were recorded according to the method outlined by Hamilton et al. (2016). The measurement process was repeated three times, and the results were averaged. ...
The enchanting phenomenon of blue coloration in animals arises from the fact that blue is one of the rarest colors found in nature, and it is a structural color that is produced by the arrangement of biological photonic nanostructures, rather than pigments. This unique coloration has evolved independently in many different species, adding to the fascination and diversity of coloration patterns in the animal kingdom. This study describes a new species of Chilobrachys Karsch, 1892 from southern Thailand that exhibits a blue-violet hue resembling the color of electrical sparks. Photographic illustrations, a morphological description, and the natural habitat of the new species are given. The diagnosis, palpal-bulb structures, spermathecae, and stridulatory organ morphology of related species are discussed.
... The deceptive morphology of tarantulas, characterized by a blend of substantial intraspecific variability and a limited array of features that appear to have evolved independently on multiple occasions, has historically rendered these spiders as taxonomically conflictive (Hamilton et al., 2016;Raven, 1985), have led to unstable classifications and have discouraged the study of other aspects of their biology. Pioneering morphology-based attempts to understand tarantula's deep phylogeny (Bertani, 2001;Guadanucci, 2014;Pérez-Miles et al., 1996), biogeography (West et al., 2012) and trait evolution (Bertani, 2000;Bertani et al., 2013;Hsiung et al., 2015;Kaderka et al., 2019), have provided essential insights into the group's biology and evolution but have often been hindered by the lack of solid support for their phylogenetic hypotheses. ...
Although tarantulas are a widespread, highly diversified, and charismatic spider group, our understanding of their evolution remains limited. A recent transcriptome-based approach, including 16% of the nominal tarantula genera (25), provided the first robust phylogenetic hypothesis of deep tarantula relationships. However, transcriptomics has practical downsides, making it less than ideal for phylogenomic analyses. Ultraconserved Elements (UCE) sequencing, a cost-effective alternative, allows for the simultaneous reading of hundreds of loci and utilizing samples with degraded DNA, including museum material, but its utility for resolving deep tarantula relationships remains to be tested. In this study, I recovered up to ~850 unique UCE loci from publicly available individual tarantula transcriptomes. Phylogenetic estimation using these loci resulted in topologies identical to those obtained from the published complete transcriptomic datasets, with limited phylogenetic uncertainty restricted to two branches in the tarantula tree of life. I conclude that UCE information has great potential for resolving at least moderate to deep tarantula relationships and that numerous UCE loci can be harvested from tarantula transcriptomic data. Finally, I provide databases of transcripts and UCE loci for the 27 currently available tarantula genera, facilitating their inclusion in future phylogenomic studies with deeper taxon coverage.
... from 2016-2018 within the Chiricahua and Peloncillo Mountains region. Following taxonomy from Hamilton et al. (2016), we found 60 records across seven species of Aphonopelma (Margalef's S richness = 1.534; Fig. 2). ...
American Arachnology: Newsletter of the American Arachnology Society 85: 13–16. Available at https://www.americanarachnology.org/fileadmin/documents/am_arachnol_newsletter/AmerArachnol85.pdf
Abstract–Tarantula road ecology is understudied and largely unknown. We observed an Aphonopelma gabeli (Smith, 1995) survive an encounter by a passing recreational vehicle’s slipstream that upheaved the tarantula. During the process, the spider tucked in its legs ventrally to protect its abdomen while rotating violently through the air before ricocheting off the pavement to a stop. The tarantula was dazed but survived the encounter.
Resumen–La ecología de las tarantulas es no ha sido ampliamente estudiada y es poco conocida. Observamos como un individuo de Aphonopelma gabeli (Smith, 1995) sobrevivió a la corriente de aire generada por un vehiculo recreativo en una carretera. Durante el proceso, la araña dobló ventralmente sus piernas para proteger su abdomen mientras giraba violentamente por el aire antes de car el pavimento. Se observó que la tarántula estuvo aturdida por unos moentos pero sobrevivió el impacto.
... Targeted enrichment approaches (e.g., Ultraconserved Elements and Anchored Hybrid Enrichment) (Faircloth et al., 2012;Lemmon and Lemmon, 2013) have been useful for inferring robust phylogenetic relationships of the non-model groups, such as Mygalomorphae spiders, such as Aphonopelma Pocock, 1901, Aptostichus Simon, 1891, Aliatypus Smith, 1908, Antrodiaetus Ausserer, 1871, Hexurella Gertsch and Platnick, 1979, Mecicobothrium Holmberg, 1882, Megahexura Kaston, 1972, Porrhothele Simon, 1892(Faircloth et al., 2012Hamilton et al., 2016b;Starrett et al., 2016;Yeates et al., 2016;Branstetter et al., 2017;Hedin et al., 2018Hedin et al., , 2019Derkarabetian et al., 2019;Zhang et al., 2019;Kulkarni et al., 2020;Xu et al., 2021). To date, there has been only one study using either of these phylogenomic approaches in tarantulas, the North American genus Aphonopelma (Hamilton et al., 2016a). Kulkarni et al. (2020) designed a spider-specific UCE probe set, which has been recovering a high number of loci, contributing to several questions in different phylogenetic studies. ...
Theraphosinae is the most diverse subfamily of Theraphosidae spiders, but their evolutionary history remains unresolved to date. This problem is common in taxonomic groups with phylogenetic hypotheses that have often been based only on qualitative morphological characters and, rarely, on molecular analyses. Phylogenomics has significantly contributed to the understanding of the evolution of many non-model groups, such as spiders. Herein, we employed ultraconserved elements (UCEs) phylogenomics to propose a new hypothesis for a group of Theraphosinae genera, namely Lasiodoriforms: Vitalius, Lasiodora, Eupalaestrus, Pterinopelma, Proshapalopus , and Nhandu . We propose three genera and their respective morphological diagnoses are provided. Our phylogeny supports the transfer of species from the genus Vitalius to Pterinopelma and Proshapalopus and from Proshapalopus to Eupalaestrus . Finally, we describe a new species of Vitalius from Southern Brazil. Based on these three new generic descriptions and transferred species, the Lasiodoriform tarantulas comprise nine genera from Argentina, Brazil, Paraguay, and Uruguay, and the genus Vitalius now includes seven species.
... formed to log-normal values, and a principal component analysis was conducted using the prcomp function in the R package stats (R Core Team, 2022) and visualized in ggplot2(Wickham, 2016), followingHamilton et al. (2016).We conducted two genetic clustering analyses. Variational AutoEncoder (VAE), an unsupervised machine learning approach derived from Bayesian probability theory, was used to visualize clustering of these lineages(Figure 4; for details seeDerkarabetian et al., 2019). ...
Species delimitation is an imperative first step toward understanding Earth's biodiversity, yet what constitutes a species and the relative importance of the various processes by which new species arise continue to be debatable. Species delimitation in spiders has traditionally used morphological characters; however, certain mygalomorph spiders exhibit morphological homogeneity despite long periods of population-level isolation, absence of gene flow, and consequent high degrees of molecular divergence. Studies have shown strong geographic structuring and significant genetic divergence among several species complexes within the trapdoor spider genus Aptostichus, most of which are restricted to the California Floristic Province (CAFP) biodiversity hotspot. Specifically, the Aptostichus icenoglei complex, which comprises the three sibling species, A. barackobamai, A. isabella, and A. icenoglei, exhibits evidence of cryptic mitochondrial DNA diversity throughout their ranges in Northern, Central, and Southern California. Our study aimed to explicitly test species hypotheses within this assemblage by implementing a cohesion species-based approach. We used genomic-scale data (ultraconserved elements, UCEs) to first evaluate genetic exchangeability and then assessed ecological interchangeability of genetic lineages. Biogeographical analysis was used to assess the likelihood of dispersal versus vicariance events that may have influenced speciation pattern and process across the CAFP's complex geologic and topographic landscape. Considering the lack of congruence across data types and analyses, we take a more conservative approach by retaining species boundaries within A. icenoglei.
... Although some names currently fashionable in modern-day taxonomy carry little or no biological information regarding the species itself, they can help to give visibility to taxonomic and species-discovery research. For example, homages to celebrities -from Johnny Cash (Aphonopelma johnnycashi Hamilton et al., 2016) to Angelina Jolie (Aptostichus angelinajolieae Bond, 2008in Bond & Stockman, 2008 to David Bowie (Heteropoda davidbowie Jäger, 2008 and Spintharus davidbowiei Agnarsson & Chomitz, 2018in Agnarsson et al., 2018 to Arnold Schwarzenegger (Predatoroonops schwarzeneggeri Brescovit, Rheims & Ott, 2012in Brescovit et al., 2012 -often attract huge scientific and media attention. This raises the important question: can this attention be harnessed to spotlight problems surrounding the taxonomy crisis? ...
There is a secret pleasure in naming new species. Besides traditional etymologies recalling the sampling locality, habitat or morphology of the species, names may be tributes to some meaningful person, pop culture references and even exercises of enigmatography. Using a dataset of 48 464 spider etymologies, we tested the hypothesis that species names given by taxonomists are deeply influenced by their cultural background. Specifically, we asked whether naming practices change through space or have changed through time. In absolute terms, etymologies referring to morphology were the most frequently used. In relative terms, references to morphology peaked in 1850–1900 and then began to decline, with a parallel increase in etymologies dedicated to people and geography. We also observed a dramatic increase in etymologies referring to pop culture and other cultural aspects in 2000–2020, especially in Europe and the Americas. While such fashionable names often carry no biological information regarding the species itself, they help give visibility to taxonomy, a discipline currently facing a profound crisis in academia. Taxonomy is among the most unchanged disciplines across the last centuries in terms of tools, rules and writing style. Yet, our analysis suggests that taxonomists remain deeply influenced by their living time and space.
... Therefore, an integration of several independent lines of evidence and broad geographic sampling is usually needed to achieve a reliable outcome (Edwards & Knowles, 2014). The presence of overlooked diversity seems to be the rule rather than an exception in Theraphosidae (Hendrixson et al., 2013) and integrative approaches have proven to be useful for detecting cryptic species within many genera (Hamilton, 2014(Hamilton, , 2016bMontes de Oca et al., 2016;Ortiz & Francke, 2016;Candia-Ramírez & Francke, 2020). In the case of the opportunistic valentinus clade, the deep genetic divergence between its two main lineages was further corroborated by genitalic as well as ecological differences. ...
Theraphosidae is the most diversified family of mygalomorph spiders, commonly known as tarantulas. Two genera inhabit the Mediterranean region: Chaetopelma in the east and Ischnocolus mostly in the western part of the Basin. Their phylogenetic position and the validity of some Ischnocolus species remain unclear. We implemented a multilocus target approach to shed new light on the position of both genera and further integrated molecular data with additional lines of evidence (morphology and ecology) to explore species boundaries in western Mediterranean Ischnocolus. Our results reveal that Ischnocolus and Chaetopelma are not closely related. Chaetopelma formed a clade with the African subfamily Eumenophorinae and Ischnocolus was recovered in a clade comprising all remaining theraphosids. The western Mediterranean Ischnocolus comprises two deeply divergent clades that separated during the Early Miocene and differ in both morphology and lifestyle. We found molecular, morphological and ecological evidence to restore the name Ischnocolus mogadorensis and revalidate this species. We also uncovered distinct allopatric lineages in Ischnocolus elongatus. However, the lack of males, the uniform morphology of females and low within-clade support hampered the assessment of their status and boundaries. Finally, our data support that I. elongatus should be considered a senior synonym of Ischnocolus hancocki and Harpactirella insidiosa.
Seven new species of Schismatothele Karsch, 1879 (Araneae, Theraphosidae) are described, almost doubling the diversity of the genus: S. caeri sp. nov.; S. caiquetia sp. nov.; S. merida sp. nov.; S. moonenorum sp. nov.; S. quimbaya sp. nov.; S. timotocuica sp. nov. and S. wayana sp. nov. An identification key for all species of Schismatothele (except S. kastoni) is presented, as well as a complementary diagnosis for the genus. Also, a standardized nomenclature is proposed to describe the prolateral keels of male palpal bulbs of species of Schismatothele.
Aim
Phylogeographical studies in the Mojave and Sonoran deserts often find genetic discontinuities that pre‐date the Pleistocene. A recent synthesis of phylogeographical data, called the Mojave Assembly Model, provides a hypothesis for the historical assembly of these desert biotas but does not adequately capture the complexity of pre‐Pleistocene vicariance events. We tested this model and assessed pre‐Pleistocene divergences by exploring the phylogeography of the Aphonopelma mojave group, which is composed of turret‐building tarantula species from the Mojave and Sonoran deserts.
Location
Mojave and Sonoran deserts, south‐western USA .
Methods
We augmented the sampling from a previous study by sequencing mitochondrial DNA ( COI ) from new material of the A. mojave group. We used phylogenetic and network analyses to identify clades and a molecular clock and lineages‐through‐time plots ( LTT plots) to explore the timing and tempo of diversification. We tested for demographic expansion using neutrality tests and mismatch distributions. Species distribution models ( SDM s) were constructed to compare current suitable habitat to that at the Last Glacial Maximum ( LGM ).
Results
Phylogenetic, network and molecular‐clock analyses identified six major clades that probably diverged during the late Miocene. The rate of diversification appears to have slowed during the Pliocene. Most clades exhibit signals of recent demographic expansion. SDM s predicted that suitable habitat shifted south and to lower elevations during the LGM .
Main conclusions
Phylogeographical analyses suggest that the A. mojave group experienced a burst of diversification in the late Miocene, followed by population expansions during the Pleistocene. Six major clades with origins in the late Miocene cannot be adequately explained by the Mojave Assembly Model. We propose the novel hypothesis that Miocene extensional tectonics caused populations to diverge in allopatry by producing low‐elevation habitat barriers. Geological models, such as kinematic reconstructions, provide an ideal but underutilized framework for testing biogeographical hypotheses in these deserts and the wider Basin and Range Province.
Motivation: Species trees provide insight into basic biology, including the mechanisms of evolution and how it modifies biomolecular function and structure, biodiversity and co-evolution between genes and species. Yet, gene trees often differ from species trees, creating challenges to species tree estimation. One of the most frequent causes for conflicting topologies between gene trees and species trees is incomplete lineage sorting (ILS), which is modelled by the multi-species coalescent. While many methods have been developed to estimate species trees from multiple genes, some which have statistical guarantees under the multi-species coalescent model, existing methods are too computationally intensive for use with genome-scale analyses or have been shown to have poor accuracy under some realistic conditions.
Results: We present ASTRAL, a fast method for estimating species trees from multiple genes. ASTRAL is statistically consistent, can run on datasets with thousands of genes and has outstanding accuracy—improving on MP-EST and the population tree from BUCKy, two statistically consistent leading coalescent-based methods. ASTRAL is often more accurate than concatenation using maximum likelihood, except when ILS levels are low or there are too few gene trees.
Availability and implementation: ASTRAL is available in open source form at https://github.com/smirarab/ASTRAL/. Datasets studied in this article are available at http://www.cs.utexas.edu/users/phylo/datasets/astral.
Contact:
warnow@illinois.edu
Supplementary information:
Supplementary data are available at Bioinformatics online.
The Madrean Sky Islands of northern Mexico and the south-western United States are located in the Madrean pine–oak woodlands ‘biodiversity hotspot’. This area is renowned for its exceptional flora and fauna, but comparatively little is known about arthropod diversity in this region. We obtained specimens of the tarantula genus Aphonopelma from five different mountain ranges (Huachucas, Pajaritos, Peloncillos, Santa Catalinas, and Santa Ritas) to assess patterns of diversity within the group.Species delimitation in tarantulas is difficult because they possess few informative morphological features. We implemented a rigorous methodological framework for delimiting species that incorporated multiple lines of evidence (molecules, distributional information, morphology, and breeding period) in an explicitly hypothesis-driven manner.Our integrative taxonomic approach delimited three species (all undescribed): one endemic to the Peloncillos, one endemic to the Santa Catalinas, and one that is more widely distributed in the Huachucas, Santa Ritas, and Pajaritos. These patterns of diversity and endemicity are comparable to other arachnids in the region and suggest that species-level diversity of stenotypic/dispersal-limited arthropods throughout the Madrean Sky Island Region may be underestimated.Ecological data are not widely available for Madrean Sky Island tarantulas and it is difficult to evaluate their conservation status. Several intrinsic (patterns of endemicity, limited vagility, and life history) and extrinsic (habitat degradation, commercial exploitation, and climate change) factors, however, suggest that these spiders may be of conservation concern. The approach taken in this study highlights the importance of considering multiple lines of evidence when species delimitation has direct implications on conservation.
The genus Aphonopelma (Theraphosidae) is marked by a dearth of distributional data and a problematic taxonomy. To develop occurrence information for the genus in Arkansas, I conducted a citizen-science based survey augmented by field work and examination of museum collections. Results of these efforts yielded a significant amount of data that enabled the construction of a county level map of Aphonopelma distribution in Arkansas; a resource hitherto unavailable. Three Aphonopelma species have been described as occurring in Arkansas: A. baergi, A. hentzi, and A. odelli. Specimens were collected from Arkansas and Oklahoma to evaluate taxonomic and historical issues concerning the three species. Morphologically, the specimen series examined comprised a relatively homogeneous group most similar to descriptions of A. hentzi. In light of comparison with specimens examined for this study, species descriptions of A. baergi and A. odelli are marked by such taxonomic uncertainty as to challenge their validity and presence in Arkansas.