Ixchela azteca (Araneae: Pholcidae), a widespread spider species from Central Mexico: Underestimated diversity or morphological and genetic variation?

Authors

DOI:

https://doi.org/10.22201/ib.20078706e.2025.96.5466

Keywords:

Delimitación de especies, Taxonomía integrativa, CO1, Ecología, Provincias biogeográficas

Abstract

We describe the morphological and genetic variation in Ixchela azteca Valdez-Mondragón & Francke from central Mexico, based on morphology and DNA barcoding of the cytochrome c oxidase subunit 1 (CO1) mitochondrial marker. Molecular analyses of species delimitation included three methods under the corrected genetic p-distances neighbor-joining (NJ) criteria: 1) Assemble Species by Automatic Partitioning (ASAP), 2) General Mixed Yule Coalescent model (GMYC), and 4) Bayesian Poisson Tree Processes (bPTP). The genetic analyses between two populations of I. azteca found a genetic p-distance of 3.5%. The molecular methods and morphology were not congruent in delimiting and recognizing two possible different species, but two allopatric populations of I. azteca are recognized. In conclusion, the incongruence of the molecular delimitation methods for species delimitations in combination with the morphology, genetic, ecological, and biogeographical evidence, corroborates to I. azteca as a widespread species in central region of Mexico, with allopatric populations in temperate but also semiarid regions.

References

Agnarsson, I. (2010). The utility of ITS2 in spider phylogenetics: Notes on prior work and an example from Anelosimus. The Journal of Arachnology, 38, 377–382. https://doi.org/10.1636/B10-01.1

Andersson, M. (1994). Sexual selection. Princeton, NJ: Princeton University Press.

Astrin, J. J., Huber, B. A., Misof, B., & Klütsch, C. F. C. (2006). Molecular taxonomy in pholcid spiders (Pholcidae: Araneae): evaluation of species identification methods using CO1 and 16S and rRNA. Zoologica Scripta, 35, 441–457. https://doi.org/10.1111/j.1463-6409.2006.00239.x

Barraclough, T. G., Harvey, P. H., & Nee, S. (1995). Sexual selection and taxonomic diversity in passerine birds. Proceedings of the Royal Society of London, Series B, 259, 211–215. https://doi.org/10.1098/rspb.1995.0031

Carstens, B. C., Pelletier, T. A., Reid, N. M., & Satler, J. (2013). How to fail at species delimitation. Molecular Ecology, 22, 4369–4383. https://doi.org/10.1111/mec.12413

Correa-Ramírez, M. M., Jiménez, M. L., & García-De León, F. J. (2010). Testing species boundaries in Pardosa sierra (Araneae: Lycosidae). Journal of Arachnology, 38, 538–554. https://doi.org/10.1636/Sh09-15.1

DeSalle, R., Egan, M. G., & Siddall, M. (2005). The unholy trinity: taxonomy, species delimitation and DNA barcoding. Philosophical Transactions of the Royal Society, London, Ser. B, 360, 1905‒1916. https://doi.org/10.1098/rstb.2005.1722

Debat, V., & David, P. (2001). Mapping phenotypes: canalization, plasticity and developmental stability. Trends in Ecology and Evolution, 16, 555‒561. https://doi.org/10.1016/S0169-5347(01)02266-2

Drummond, A. J., Suchard, M. A., Xie, D., Rambaut, A. (2012). Bayesian phylogenetics with BEAUti and the BEAST 1.7. Molecular Biology and Evolution, 29, 1969–1973. https://doi.org/10.1093/molbev/mss075

Eberhard, W. G. (1985). Sexual selection and animal genitalia. Cambridge, MA: Harvard University Press. https://doi.org/10.4159/harvard.9780674330702

Eberhard, W. G., Huber, B. A., Rodríguez, R. L., Briceno, R. D., Salas, I., & Rodríguez, V. (1998). One size fits all? Relationships between the size and degree of variation in genitalia and other body parts in twenty species of insects and spiders. Evolution, 52, 415–431. https://doi.org/10.1111/j.1558-5646.1998.tb01642.x

Fujisawa, T., & Barraclough, T. G. (2013). Delimiting species using single-locus data and the Generalized Mixed Yule Coalescent approach: a revised method and evaluation on simulated data sets. Systematic Biology, 62, 707–724. https://doi.org/10.1093/sysbio/syt033

Graham, M. R., Hendrixson, B. E., Hamilton, C. A., & Bond, J. E. (2015). Miocene extensional tectonics explain ancient patterns of diversification among turret-building tarantulas (Aphonopelma mojave group) in the Mojave and Sonoran deserts. Journal of Biogeography, 42, 1052–1065. https://doi.org/10.1111/jbi.12494

Gross, M. R. (1996). Alternative reproductive strategies and tactics: diversity within sexes. Trends in Ecology & Evolution, 11, 92–98. https://doi.org/10.1016/01695347(96)81050-0

Hall, T. A. (1999). BioEdit: a user-friendly biological sequence alignment editor and analysis program for Windows 95/98/NT. Nucleic Acids Symposium Series, 41, 95–98.

Hamilton, C. A., Formanowicz, D. R., & Bond, J. E. (2011). Species delimitation and phylogeography of Aphonopelma hentzi (Araneae, Mygalomorphae, Theraphosidae): Cryptic diversity in North American tarantulas. PLoS ONE, 6, e26207. https://doi.org/10.1371/journal.pone.0026207

Hamilton, C. A., Hendrixson, B. E., Brewer, M. S., & Bond, J. E. (2014). An evaluation of sampling effects on multiple DNA barcoding methods leads to an integrative approach for delimiting species: A case study of the North American tarantula genus Aphonopelma (Araneae, Mygalomorphae, Theraphosidae). Molecular Phylogenetics and Evolution, 71, 79–93. https://doi.org/10.1016/j.ympev.2013.11.007

Hamilton, C. A., Hendrixson, B. E., & Bond, J. E. (2016). Taxonomic revision of the tarantula genus Aphonopelma Pocock, 1901 (Araneae, Mygalomorphae, Theraphosidae) within the United States. Zookeys, 560, 1–340. https://doi.org/10.3897/zookeys.560.6264

Hebert, C. A., Ball, S. L., & Dewaard, J. R. (2003). Biological identifications through DNA barcodes. Proceedings Biological Sciences, 270, 313–321. https://doi.org/10.1098/rspb.2002.2218

Hebert, P. D. N., Ratnasingham, S., DeWaard, J. R. (2003). Barcoding animal life: Cytochrome c oxidase subunit 1 divergences among closely related species. Proceedings of the Royal Society of London, Ser. B, 270, S96–S99. https://doi.org/10.1098/rsbl.2003.0025

Hebert, P. D. N., Penton, E. H., Burns, J. M., Janzen, D. H., & Hallwachs, W. (2004). Ten species in one: DNA barcoding reveals cryptic species in the Neotropical skipper butterfly Astraptes fulgerator. Proceedings of the National Academy of Sciences of the United States of America, 101, 14812‒14817. https://doi.org/10.1073/pnas.0406166101

Huber, B. A. (2000). New World Pholcid Spiders (Araneae: Pholcidae): a revision at generic level. Bulletin of the American Museum of Natural History, 254, 1–348. http://dx.doi.org/10.1206/0003-0090(2000)254<0001:NWPSAP>2.0.CO;2

Huber, B. A. (2003). Rapid evolution and species-specificity of arthropod genitalia: fact or artifact? Organisms Diversity & Evolution, 3, 63–71. https://doi.org/10.1078/1439-6092-00059

Huber, B. A. (2011). Phylogeny and classification of Pholcidae (Araneae): an update. Journal of Arachnology, 39, 211–222. https://doi.org/10.1636/CA10-57.1

Huber, B. A. (2018). Cave-dwelling pholcid spiders (Araneae, Pholcidae): a review. Subterranean Biology, 26, 1–18. http://doi.org/10.3897/subtbiol.26.26430

Huber, B. A. (2021). Beyond size: sexual dimorphisms in pholcid spiders. Arachnology, 18, 656–677. https://doi.org/10.13156/arac.2020.18.7.656

Huber, B. A., & Pérez G. A. (2001). A new genus of pholcid spiders (Araneae: Pholcidae) endemic to western Cuba, with a case of female genitalic dimorphism. American Museum Novitates, 3329, 1–23. https://doi.org/10.1206/00030082(2001)329<0001:ANGOPS>2.0.CO;2

Kapli, P., Lutteropp, S., Zhang, J., Kobert, K., Pavlidis, P., & Stamatakis, A. (2017). Multi-rate Poisson tree processes for single-locus species delimitation under maximum likelihood and Markov chain Monte Carlo. Bioinformatics, 33, 1630–1638. https://doi.org/10.1093/bioinformatics/btx02

Katoh, K., & Toh, H. (2008). Recent developments in the MAFFT multiple sequence alignment program. MAFFT version 7. Briefings in Bioinformatics, 4, 286–298. https://doi.org/10.1093/bib/bbn013

Kearse, M., Moir, R., Wilson, A., Stones-Havas, S., Cheung, M., & Sturrock, S. (2012). Geneious Basic: an integrated and extendable desktop software platform for the organization and analysis of sequence data. Bioinformatics, 28, 1647–1649. https://doi.org/10.1093/bioinformatics/bts199

Letunic, I., & Bork, P. (2021). Interactive Tree of Life (iTOL) v5: an online tool for phylogenetic tree display and annotation. Nucleic Acids Research, 49, 293–296. https://doi.org/10.1093/nar/gkab301

Monaghan, M. T., Wild, R., Elliot, M., Fujisawa, T., Balke, M., & Inward, D. J. (2009). Accelerated species inventory on Madagascar using coalescent-based models of species delineation. Systematic Biology, 58, 298–311. https://doi.org/10.1093/sysbio/syp027

Navarro-Rodríguez, I., & Valdez-Mondragón, A. (2020). Description of a new species of Loxosceles Heineken & Lowe (Araneae, Sicariidae) recluse spiders from Hidalgo, Mexico, under integrative taxonomy: morphological and DNA barcoding data (CO1+ITS2). European Journal of Taxonomy, 704, 1–30. https://doi.org/10.5852/ejt.2020.704

Navarro-Rodríguez, C. I., & Valdez-Mondragón, A. (2024). Violins we see, species we don’t… Species delimitation of the spider genus Loxosceles Heineken & Lowe (Araneae: Sicariidae) from North America using morphological and molecular evidence. Zootaxa, 5428, 527–548. https://doi.org/10.11646/zootaxa.5428.4.4

Nolasco, S., & Valdez-Mondragón, A. (2022a). To be or not to be… Integrative taxonomy and species delimitation in the daddy long-legs spiders of the genus Physocyclus (Araneae, Pholcidae) using DNA barcoding and morphology. Zookeys, 1135, 93–118. https://doi.org/10.3897/zookeys.1135.94628

Nolasco, S., & Valdez-Mondragón, A. (2022b). Four new species of the spider genus Physocyclus Simon, 1893 (Araneae: Pholcidae) from Mexico, with updated taxonomic identification keys. European Journal of Taxonomy, 813, 173–206. https://doi.org/10.5852/ejt.2022.813.1739

Ortiz, D., & Francke, O. F. (2016). Two DNA barcodes and morphology for multi-method species delimitation in Bonnetina tarantulas (Araneae: Theraphosidae). Molecular Phylogenetics and Evolution, 101, 176‒193. https://doi.org/10.1016/j.ympev.2016.05.003

Planas, E., & Ribera, C. (2015). Description of six new species of Loxosceles (Araneae: Sicariidae) endemic to the Canary Islands and the utility of DNA barcoding for their fast and accurate identification. Zoological Journal of the Linnean Society, 174, 47–73. https://doi.org/10.1111/zoj.12226

Pons, J., Barraclough, T. G., & Gómez-Zurita, J. (2006). Sequence based species delimitation for the DNA taxonomy of undescribed insects. Systematic Biology, 55, 595–609. https://doi.org/10.1080/10635150600852011

Posada, D., & Buckley, T. R. (2004). Model selection and model averaging in phylogenetics: advantages of the AIC and Bayesian approaches over likelihood ratio tests. Systematics Biology, 50, 580–601. https://doi.org/10.1080/10635150490522304

Puillandre, N., Brouillet, S., & Achaz, G. (2021). ASAP. Assemble species by automatic partitioning. Molecular Ecology Resources, 21, 609–620. https://doi.org/10.1111/1755-0998.13281

Rambaut, A., & Drummond, A. J. (2003). TRACER, MCMC trace analysis tool. Version 1.6. Institute of Evolutionary Biology, University of Edinburgh, Edinburgh, Department of Computer Science, University of Auckland, Auckland.

Ronquist, F., & Huelsenbeck, J. P. (2003). MRBAYES 3: Bayesian phylogenetic inference under mixed models. Bioinformatics, 19, 1572‒1574. https://doi.org/10.1093/bioinformatics/btg180

Tahami, M. S., Zamani, A., Sadeghil, S., & Ribera, C. (2017). A new species of Loxosceles Heineken & Lowe, 1832 (Araneae: Sicariidae) from Iranian caves. Zootaxa, 4318, 377–387. https://doi.org/10.11646/zootaxa.4318.2.10

Tamura, K., Dudley, J., Nei, M., & Kumar, S. (2007). MEGA7: Molecular Evolutionary Genetics Analysis (MEGA) software version 4.0. Molecular Biology and Evolution, 24, 1596–1599. https://doi.org/10.1093/molbev/msm092

Valdez-Mondragón, A. (2010). Revisión taxonómica del género de arañas Physocyclus Simon, 1893 (Araneae: Pholcidae), con la descripción de especies nuevas de México. Revista Ibérica de Aracnología, 18, 3–80.

Valdez-Mondragón, A. (2013). Taxonomic revision of the spider genus Ixchela Huber, 2000 (Araneae: Pholcidae), with description of ten new species from Mexico and Central America. Zootaxa, 3608, 285–327. http://dx.doi.org/10.11646/zootaxa.3608.5.1

Valdez-Mondragón, A. (2020). COI mtDNA barcoding and morphology for species delimitation in the spider genus Ixchela Huber (Araneae: Pholcidae), with the description of two new species from Mexico. Zootaxa, 4747, 54–76. https://doi.org/10.11646/zootaxa.4747.1.2

Valdez-Mondragón, A., & Francke, O. F. (2015). Phylogeny of the spider genus Ixchela Huber, 2000 (Araneae: Pholcidae) based on morphological and molecular evidence (CO1 and 16S), with a hypothesized diversification in the Pleistocene. Zoological Journal of the Linnean Society, 175, 20‒58. https://doi.org/10.1111/zoj.12265

Valdez-Mondragón, A., Navarro-Rodríguez, C. I., Solís-Catalán K. P., Cortez-Roldán, M. R., & Juárez-Sánchez, A. R. (2019). Under an integrative taxonomic approach: the description of a new species of the genus Loxosceles (Araneae, Sicariidae) from Mexico City. Zookeys, 892, 93–133. https://doi.org/10.3897/zookeys.892.39558

World Spider Catalog (WSC). (2025). World Spider Catalog. Version 25.5. Natural History Museum Bern. http://wsc.nmbe.ch

Zhang, J., Kapli, P., Pavlidis, P., & Stamatakis, A. (2013). A general species delimitation method with applications to phylogenetic placements. Bioinformatics, 29, 2869–2876. https://doi.org/10.1093/bioinformatics/btt499

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Published

2025-02-28

How to Cite

Valdez-Mondragón, A., & Nolasco-Garduño, S. . (2025). Ixchela azteca (Araneae: Pholcidae), a widespread spider species from Central Mexico: Underestimated diversity or morphological and genetic variation?. Revista Mexicana De Biodiversidad, 96, e965466. https://doi.org/10.22201/ib.20078706e.2025.96.5466

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TAXONOMÍA Y SISTEMÁTICA