Abstract
The Neotropical freshwater crab, Pseudothelphusa americana de Saussure, 1857, has historically been an unstable taxon. New collections from the published range of P. americana indicate a great deal of morphological variation within this taxon, suggesting that it may be a species complex. In the present study we describe three new species within the P. americana complex based on a phylogenetic analysis, on a study of the morpho-geometric variation of the gonopod, and on a thorough morphological revision of somatic and sexual structures. Three new species (P. chilapaensis sp. nov., P. hermosorum sp. nov., and P. queretana sp. nov.) are described, and P. americana sensu stricto is redescribed, a neotype designated and its revised geographic distribution mapped.
References
- Adams, D., Collyer, M., Kaliontzopoulou, A. & Baken, E. (2025) Geomorph: software for geometric morphometric analyses. R package Version 4.0.10. Available from: https://cran.r-project.org/package=geomorph (accessed 11 May 2023)
- Álvarez, A. & Villalobos, J.L. (1997) Pseudothelphusidae y Trichodactylidae (cangrejos). In: Soriano, R., Dirzo, R. & Vogt, R.C. (Eds.), Historia Natural de Los Tuxtlas. Instituto de Ecología, Ciudad de México, pp. 415–418.
- Álvarez, F., Ojeda, J.C., Souza-Carvalho, E., Villalobos, J.L., Magalhães, C., Wehrtmann, I.S. & Mantelatto, F.L. (2020) Revision of the higher taxonomy of neotropical freshwater crabs of the family Pseudothelphusidae, based on multigene and morphological analyses. Zoological Journal of the Linnean Society, 19, 973–1001. https://doi.org/10.1093/zoolinnean/zlaa162
- Álvarez, F., Villalobos, J.L., Armendáriz, G. & Hernández, C. (2012a) Relación biogeográfica entre cangrejos dulceacuícolas y acociles a lo largo de la zona mexicana de transición: revaluación de la hipótesis de Rodríguez (1986). Revista Mexicana de Biodiversidad, 83, 1073–1083.
- Álvarez, F., Villalobos, J.L. & Moreno, E. (2012b) Pseudothelphusa zongolicae (Decapoda, Pseudothelphusidae), a new species of freshwater crab from Veracruz, Mexico. Crustaceana, 85, 1541–1547. https://doi.org/10.1163/15685403-00003132
- Álvarez, F. & Villalobos, J.L. (1996) Especie nueva de cangrejos de agua dulce del género Pseudothelphusa (Brachyura: Pseudothelphusidae) de Guerrero, México. Anales del Instituto de Biología, Serie Zoología, 67, 297–302.
- Arnqvist, G. & Martensson, T. (1998) Measurement error in geometric morphometrics: empirical strategies to assess and reduce its impact on measures of shape. Acta Zoologica Academiae Scientiarum Hungaricae, 44, 73–96.
- Baken, E., Collyer, M., Kaliontzopoulou, A. & Adams, D. (2021) Geomorph v4.0 and gmShiny: enhanced analytics and a new graphical interface for a comprehensive morphometric experience. Methods in Ecology and Evolution, 12, 2355–2363. https://doi.org/10.1111/2041-210X.13723
- Barbour, C.D. (1973) A biogeographical history of Chirostoma (Pisces: Atherinidae): a species flock from the Mexican Plateau. Copeia, 1973, 533–556. https://doi.org/10.2307/1443118
- Baylac, M., Villemant, C. & Simbolotti, G. (2003) Combining geometric morphometrics with pattern recognition for the investigation of species complexes. Biological Journal of the Linnean Society, 80, 89–98. https://doi.org/10.1046/j.1095-8312.2003.00221.x
- Bookstein, F.L. (1991) Morphometric tools for landmark data. Cambridge University Press, Cambridge, 435 pp. https://doi.org/10.1017/CBO9780511573064
- Bookstein, F.L. (1996) Biometrics, biomathematics, and the morphometric synthesis. Bulletin of Mathematical Biology, 58, 313–365. https://doi.org/10.1007/BF02458311
- Bott, R. (1968) Fluß-Krabben aus dem östlichen Mittel-Amerika und von den Großen Antillen (Crustacea, Decapoda). Senckenbergiana Biologica, 49, 39–49.
- Bott, R. (1970) Über die Entwicklungsgeschichte und Verbreitung der Süsswasser-Krabben nach der Sammlung des Naturhistorischen Museums in Genf/Schweiz. Revue Suisse de Zoologie, 77, 327–344. https://doi.org/10.5962/bhl.part.75900
- Buser, T.J., Sidlauskas, B.L. & Summers, A.P. (2018) 2D or not 2D? Testing the utility of 2D vs. 3D landmark data in geometric morphometrics of the sculpin subfamily Oligocottinae (Pisces; Cottoidea). The Anatomical Record, 301, 806–818. https://doi.org/10.1002/ar.23752
- Cardini, A. (2014) Missing the third dimension in geometric morphometrics: how to assess if 2D images really are a good proxy for 3D structures? Hystrix-Italian Journal of Mammalogy, 25, 73–81.
- Cardini, A. & O'Higgins, P. (2004) Patterns of morphological evolution in Marmota (Rodentia, Sciuridae): geometric morphometrics of the cranium in the context of marmot phylogeny, ecology and conservation. Biological Journal of the Linnean Society, 82, 385–407. https://doi.org/10.1111/j.1095-8312.2004.00367.x
- Chace, F.A. & Hobbs, H.H. (1969) The freshwater and terrestrial decapod crustaceans of the West Indies with special reference to Dominica. Bulletin of the United States National Museum, 292, 1–258. https://doi.org/10.5479/si.03629236.292.1
- Coifmann, I. (1939) Potamondi della Guina Inglesa raccolti dal Prof. Nello Beccari. Archivio Zoologico Italiano, 27, 93–116.
- Colgan, D.J., McLauchlan, A., Wilson, G.D.F., Livingston, S.P., Edgecombe, G.D., Macaranas, J., Cassis, G. & Gray, M.R. (1998) Histone H3 and U2 snRNA DNA sequences and arthropod molecular evolution. Australian Journal of Zoology, 46, 419–437. https://doi.org/10.1071/ZO98048
- Cumberlidge, L. (1991) The respiratory system of Globonautes macropus (Rathbun, 1898), a terrestrial freshwater crab from Liberia (Gecarcinucoidea, Gecarcinucidae). Crustaceana, 61, 69–80. https://doi.org/10.1163/156854091X00524
- Darriba, D., Taboada, G.L., Doallo, R. & Posada, D. (2012) jModelTest 2: more models, new heuristics and parallel computing. Nature Methods, 9, 772. https://doi.org/10.1038/nmeth.2109
- Davis, M.A., Douglas, M.R., Collyer, M.L. & Douglas, M.E. (2016) Deconstructing a species-complex: geometric morphometric and molecular analyses define species in the Western Rattlesnake (Crotalus viridis). PLoS ONE, 11, e0146166. https://doi.org/10.1371/journal.pone.0146166
- de Saussure, H. (1857) Diagnosis de quelques Crustacés nouveaux des Antilles et du Mexique. Revue et Magasin de Zoologie Pure et Appliquée, Series 2, 304–306.
- de Saussure, H. (1858) Mémoire sur divers Crustacés nouveaux des Antilles et du Mexique. Mémoires de la Société de Physique et d'Histoire Naturelle de Genève, 14 (2), 417–496.
- Díaz, H. & Rodríguez, G. (1977) The branchial chamber in terrestrial crabs: a comparative study. The Biological Bulletin, 153, 485–504. https://doi.org/10.2307/1540602
- Domínguez-Domínguez, O., Pedraza-Lara, C., Gurrola-Sánchez, N., Perea, S., Pérez-Rodríguez, R., Israde-Alcántara, I., Garduño-Monroy, V.H., Doadrio, I., Pérez-Ponce de Léon, G. & Brooks, D.R. (2010) Historical biogeography of the Goodeinae (Cyprinodontiforms). Viviparous Fishes, 2, 13–30.
- Evin, A., Baylac, M., Ruedi, M., Mucedda, M. & Pons, J.M. (2008) Taxonomy, skull diversity and evolution in a species complex of Myotis (Chiroptera: Vespertilionidae): a geometric morphometric appraisal. Biological Journal of the Linnean Society, 95, 529–538. https://doi.org/10.1111/j.1095-8312.2008.01076.x
- Fang, F., Ji, Y., Zhao, Q., Wang, Y., Gao, W., Chu, K. & Sun, H. (2015) Phylogeography of the Chinese endemic freshwater crab Sinopotamon acutum (Brachyura, Potamidae). Zoologica Scripta, 44, 653–666. https://doi.org/10.1111/zsc.12131
- Fruciano, C. (2016) Measurement error in geometric morphometrics. Development Genes and Evolution, 226, 139–158. https://doi.org/10.1007/s00427-016-0537-4
- Geospiza, Inc. (2012) Available from: https://www.geospiza.com/finchTV (accessed 11 May 2023)
- Grinang, J., Das, I. & Ng, P.K.L. (2019) Geometric morphometric analysis in female freshwater crabs of Sarawak (Borneo) permits addressing taxonomy-related problems. PeerJ, 7, e6205. https://doi.org/10.7717/peerj.6205
- Guindon, S. & Gascuel, O. (2003) A simple, fast and accurate method to estimate large phylogenies by maximum-likelihood. Systematic Biology, 52, 696–704. https://doi.org/10.1080/10635150390235520
- Halffter, G. & Morrone, J.J. (2017) An analytical review of Halffter's Mexican transition zone, and its relevance for evolutionary biogeography, ecology and biogeographical regionalization. Zootaxa, 4226 (1), 1–46. https://doi.org/10.11646/zootaxa.4226.1.1
- Hollier, J. (2020) The Decapoda (Crustacea) described by Henri de Saussure. Revue Suisse de Zoologie, 125, 291–298.
- Hurvich, C.M. & Tsai, C.L. (1989) Regression and time series model selection in small samples. Biometrika, 76, 297–307. https://doi.org/10.1093/biomet/76.2.297
- International Commission on Zoological Nomenclature (1999) International Code of Zoological Nomenclature. 4th Edition. International Trust for Zoological Nomenclature, London, 156 pp.
- Jesse, R., Schubart, C.D. & Klaus, S. (2010) Identification of a cryptic lineage within Potamon fluviatile (Herbst) (Crustacea: Brachyura: Potamidae). Invertebrate Systematics, 24, 348–356. https://doi.org/10.1071/IS10014
- Kassambara, A. & Mundt, F. (2020) Factoextra: extract and visualize the results of multivariate data analyses. R Package Version 1.0.7. Available from: https://CRAN.R-project.org/package=factoextra (accessed 11 May 2023)
- Katoh, K., Kuma, K.I., Toh, H. & Miyata, T. (2005) MAFFT version 5: improvement in accuracy of multiple sequence alignment. Nucleic Acids Research, 33, 511–518. https://doi.org/10.1093/nar/gki198
- Katoh, K., Rozewicki, J. & Yamada, K.D. (2019) MAFFT online service: multiple sequence alignment, interactive sequence choice and visualization. Briefings in Bioinformatics, 20, 1160–1166. https://doi.org/10.1093/bib/bbx108
- Kumar, S., Stecher, G. & Tamura, K. (2016) MEGA7: molecular evolutionary genetics analysis version 7.0 for bigger datasets. Molecular Biology and Evolution, 33, 1870–1874. https://doi.org/10.1093/molbev/msw054
- Lanfear, R., Frandsen, P.B., Wright, A.M., Senfeld, T. & Calcott, B. (2017) PartitionFinder 2: new methods for selecting partitioned models of evolution for molecular and morphological phylogenetic analyses. Molecular Biology and Evolution, 34, 772–773. https://doi.org/10.1093/molbev/msw260
- Latreille, P.A. (1819) Thelphuse, Thelphusa. Nouveau Dictionnaire d'Histoire Naturelle, Appliquée aux Arts, a l'Agriculture, à l'Économie Rurale et Domestique, à la Médicine, etc. Nouvelle Édition, 33, 500–504.
- Latreille, P.A. (1831) Cours d'Entomologie, ou de l'histoire naturelle des Crustacés, des Arachnides, des Myriapodes et des Insects à l'usage des élèves de l'école du Muséum d'Histoire Naturelle (accompagné d'un Atlas). Librairie Encyclopédique de Roret, Paris, 568 pp. https://doi.org/10.5962/bhl.title.4854
- Maddison, W.P. & Maddison, D.R. (2019) Mesquite: a modular system for evolutionary analysis. Version 3.61. Available from: http://www.mesquiteproject.org (accessed 11 May 2023)
- Maderbacher, M., Bauer, C., Herler, J., Postl, L., Makasa, L. & Sturmbauer, C. (2008) Assessment of traditional versus geometric morphometrics for discriminating populations of the Tropheus moorii species complex (Teleostei: Cichlidae), a Lake Tanganyika model for allopatric speciation. Journal of Zoological Systematics and Evolutionary Research, 46, 153–161. https://doi.org/10.1111/j.1439-0469.2007.00447.x
- Mateos, M., Sanjur, O.I. & Vrijenhoek, R.C. (2002) Historical biogeography of the livebearing fish genus Poeciliopsis (Poeciliidae: Cyprinodontiformes). Evolution, 56, 972–984. https://doi.org/10.1111/j.0014-3820.2002.tb01409.x
- Maechler, M., Rousseeuw, P., Struyf, A., Hubert, M. & Hornik, K. (2024) Cluster: cluster analysis basics and extensions. R package Version 2.1.8. Available from: https://CRAN.R-project.org/package=cluster (accessed 11 May 2023)
- Mejía‐Madrid, H.H., Vázquez‐Domínguez, E. & Pérez‐Ponce De León, G. (2007) Phylogeography and freshwater basins in central Mexico: recent history as revealed by the fish parasite Rhabdochona lichtenfelsi (Nematoda). Journal of Biogeography, 34, 787–801. https://doi.org/10.1111/j.1365-2699.2006.01651.x
- Meyer, C.P., Geller, J.B. & Paulay, G. (2005) Fine scale endemism on coral reefs: archipelagic differentiation in turbinid gastropods. Evolution, 59, 113–125. https://doi.org/10.1111/j.0014-3820.2005.tb00899.x
- Miller, M.A., Pfeiffer, W. & Schwartz, T. (2010) Creating the CIPRES Science Gateway for inference of large phylogenetic trees. Proceedings of the Gateway Computing Environments Workshop (GCE), New Orleans, Louisiana, 14 November 2010, 1–8. https://doi.org/10.1109/GCE.2010.5676129
- Moreno-Juárez, E.G., Villalobos, J.L. & Álvarez, F. (2022a) Revision of the freshwater crabs of the genus Tehuana Rodríguez & Smalley in Smalley 1970 (Decapoda, Pseudothelphusidae), with the descriptions of two new species. ZooKeys, 1117, 1–35. https://doi.org/10.3897/zookeys.1117.85362
- Moreno-Juárez, E.G., Villalobos, J.L. & Álvarez, F. (2022b) Two new genera and one new species of freshwater crabs of the subfamily Pseudothelphusinae (Decapoda: Brachyura: Pseudothelphusidae) from southwestern Mexico. Zootaxa, 5200 (1), 24–36. https://doi.org/10.11646/zootaxa.5200.1.2
- Mullin, S.K. & Taylor, P.J. (2002) The effects of parallax on geometric morphometric data. Computers in Biology and Medicine, 32, 455–464. https://doi.org/10.1016/S0010-4825(02)00037-9
- Ng, P.K.L, Guinot, D. & Davie, P. (2008) Systema Brachyorum: part I. An annotated checklist of extant brachyuran crabs of the world. Raffles Bulletin of Zoology, 17, 1–128
- Ojeda-Escoto, J.C., Villalobos, J.L. & Álvarez, F. (2017) Three new species of freshwater crabs of the genus Pseudothelphusa (De Saussure) (Crustacea: Decapoda: Pseudothelphusa) from Mexico. Zootaxa, 4216 (6), 559–571. https://doi.org/10.11646/zootaxa.4216.6.3
- Oksanen, J., Simpson, G., Blanchet, F., Kindt, R., Legendre, P., Minchin, P., O'Hara, R., Solymos, P., Stevens, M., Szoecs, E., Wagner, H., Barbour, M., Bedward, M., Bolker, B., Borcard, D., Carvalho, G., Chirico, M., De Caceres, M., Durand, S., Evangelista, H., FitzJohn, R., Friendly, M., Furneaux, B., Hannigan, G., Hill, M., Lahti, L., McGlinn, D., Ouellette, M., Ribeiro-Cunha, E., Smith, T., Stier, A., Ter-Braak, C., Weedon, J. & Borman, T. (2025) Vegan: community ecology package. R package. Version 2.6-10. Available from: https://github.com/vegandevs/vegan (accessed 27 May 2025)
- Ortmann, A. (1893) Die Decapoden-Krebse des Strasbourg Museums, mit besonderer bërucksichtingung der von Herrn Dr. Döderlein bei Japan und bei den Liu-Kiu-Inseln gesammelten und zur Zeit im Strassburger Museum aufbewahrten Formen. VII. Theil. Abtheilung: Brachyura (Brachyura genuina Boas) II. Unterabtheilung: Cancroidea, 2. Section: Cancrinea, 1. Gruppe: Cyclometopa. Zoologische Jahrbücher Abtheilung für Systematik, Geographie und Biologie der Thiere, 7, 411–495. https://doi.org/10.5962/bhl.part.24064
- Ortmann, A. (1897) Carcinologishe Stuiden. Zoologische Jahrbücher Abtheilung für Systematik, Geographie und Biologie der Thiere, 10, 258–372.
- Palumbi, S.R. & Benzie, J. (1991) Large mitochondrial DNA differences between morphologically similar penaeid shrimp. Molecular Marine Biology and Biotechnology, 1, 27–34.
- Pečnerová, P., Moravec, J.C. & Martinkova, N. (2015) A skull might lie modeling ancestral ranges and diet from genes and shape of tree squirrels. Systematic Biology, 64, 1074–1088. https://doi.org/10.1093/sysbio/syv054
- Pocock, R.I. (1889) Contributions to our knowledge of the crustacea of Dominika. The Annals and Magazine of Natural History, Series 6, 3 (13), 6–21. https://doi.org/10.1080/00222938909460288
- Pretzmann, G. (1965) Vorläufiger Bericht über die Familie Pseudothelphusidae. Anzeiger der mathematisch-naturwissenschaftlichen Klasse der Österreichischen Akademie der Wissenschaften, 1965, 1–10.
- Pretzmann, G. (1968) Neue südamerikanische Süsswasserkrabben (Vorläufige Mitteilung). Entomologisches Nachrichtenblatt, Wien, 15, 1–15.
- Pretzmann, G. (1971) Fortschritte in der Klassifizierung der Pseudothelphusidae. Aus den Sitzungsberichten der Mathematisch-naturwissenschaftlichen Klasse der Österreichischen Akademie der Wissenschaften, Wien, Abteilung I, 179 Bd, 1. bis 4. Heft, 16–24.
- Pretzmann, G. (1972) Die Pseudothelphusidae (Crustacea Brachyura). Zoologica, Stuttgart, 42, 1–182.
- Rambaut, A., Drummond, A.J., Xie, D., Baele, G. & Suchard, M.A. (2018) Posterior summarization in bayesian phylogenetics using Tracer 1.7. Systematic Biology, 67, 901–904. https://doi.org/10.1093/sysbio/syy032
- Rathbun, M.J. (1893) Descriptions of new species of American freshwater crabs. Proceedings of the United States National Museum, 16, 649–661. https://doi.org/10.5479/si.00963801.16-959.649
- Rathbun, M.J. (1898) A contribution to a knowledge of the fresh-water crabs of America. The Pseudothelphusinae. Proceedings of the United States National Museum, 21, 507–537. https://doi.org/10.5479/si.00963801.21-1158.507
- Rathbun, M.J. (1905) Les Crabes d’eau douce (Potamonidae). Nouvelles Archives du Museum d’Histoire naturelle, 4, 159–321.
- Rodriguez, G. (1982) Les crabes d'eau douce d'Amérique: famille des Pseudothelphusidae. Faune Tropicale XXII. Office de La Recherche Scientifique et Technique Outre-Mer, Fort-de-France, Martinique, 223 pp.
- Rodríguez, G. & Smalley, A. (1972) Los cangrejos de agua dulce de México de la familia Pseudothelphusidae (Crustacea, Brachyura). Anales del Instituto de Biología Universidad Nacional Autónoma de México, 40, 69–112.
- Rohlf, F.J. (2006a) Tps-DIG, Digitize landmarks and outlines. Fersion 2.10. Software and manual. Department of Ecology and Evolution, State University of New York, Stony Brook, New York. [program]
- Rohlf, F.J. (2006b) Tps-UTIL, File utility program. Version 1.38. Software and Manual). Department of Ecology and Evolution, State University of New York, Stony Brook, New York. [program]
- 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
- Roth, V.L. (1993) On three-dimensional morphometrics, and on the identification of landmark points. Contributions to Morphometrics. Museo Nacional de Ciencias Naturales, 41, 61.
- Santamaría, J., Cordón, O., Damas, S. & Ibánez, O. (2009) Tackling the coplanarity problem in 3D camera calibration by means of fuzzy landmarks: a performance study in forensic craniofacial superimposition. In: Shi, J., Wang, H., Mordohai, P., Swaminathan, R., Ying, L. & Basu, A. (Eds.), 2009 IEEE 12th International Conference on Computer Vision Workshops, ICCV Workshops. IEEE, New York, New York, pp. 1686–1693. https://doi.org/10.1109/ICCVW.2009.5457486
- Schlager, S. (2017) Morpho and Rvcg – shape analysis in R. In: Zheng, G., Li, S. & Szekely, G. (Eds.), Statistical Shape and Deformation Analysis. Academic Press, London, pp. 217–256. https://doi.org/10.1016/B978-0-12-810493-4.00011-0
- Scrucca, L., Fraley, C., Murphy, T.B. & Raftery, A.E. (2023) Model-based clustering, classification, and density estimation using mclust in R. Chapman and Hall, CRC Press, New York, New York, 268 pp. https://doi.org/10.1201/9781003277965
- Smith, S.I. (1870) Notes on American crustacea. No. 1. Ocypodoidea. Transactions of Connecticut Academy of Arts and Sciences, 2, 113–176.
- Stamatakis, A. (2014) RAxML version 8: a tool for phylogenetic analysis and post-analysis of large phylogenies. Bioinformatics, 30, 1312–1313. https://doi.org/10.1093/bioinformatics/btu033
- Swainson, W. (1839) The natural history and classification of fishes, amphibians, & reptiles, or monocardian animals. Longman’s, London, 448 pp. https://doi.org/10.5962/bhl.title.62140
- Taylor, H.H. & Greenway, P. (1979) The structure of the gills and lungs of the arid-zone crab, Holthusiana (Astrothelphusa) transversa (Brachyura: Sundathelphusidae) including observations on arterial vessels within the gills. Journal of Zoology, 189, 359–384. https://doi.org/10.1111/j.1469-7998.1979.tb03969.x
- Theska, T., Sieriebriennikov, B., Wighard, S.S., Werner, M.S. & Sommer, R.J. (2020) Geometric morphometrics of microscopic animals as exemplified by model nematodes. Nature Protocols, 15, 2611–2644. https://doi.org/10.1038/s41596-020-0347-z
- Toro, M.V., Manriquez, G. & Suazo, I. (2010) Morfometría geométrica y el estudio de las formas biológicas: de la morfología descriptiva a la morfología cuantitativa. International Journal of Morphology, 28, 977–990. https://doi.org/10.4067/S0717-95022010000400001
- Torres, M.V., Collins, P.A. & Giri, F. (2014) Morphological variation of freshwater crabs Zilchiopsis collastinensis and Trichodactylus borellianus (Decapoda, Trichodactylidae) among localities from the middle Paraná river basin during different hydrological periods. ZooKeys, 457, 171–186. https://doi.org/10.3897/zookeys.457.6726
- Vilaclara, G., Martinez-Mekler, G., Cuna, E. & Ugalde, E. (2010) Diatom-inferred palaeoenvironmental changes of a Pliocene lake disturbed by volcanic activity. Journal of Paleolimnology, 44, 203–215. https://doi.org/10.1007/s10933-009-9397-5
- Villalobos-F., A. (1982) Decapoda. In: Hurlbert, S.H. & Villalobos-Figueroa, A. (Eds.), Aquatic biota of Mexico, Central America and the West Indies. San Diego State University, San Diego, California, pp. 215–239.
- Villalobos, J.L. & Alvarez, F. (2010) Phylogenetic analysis of the Mexican freshwater crabs of the tribe Pseudothelphusini (Decapoda: Brachyura: Pseudothelphusidae). Zoological Journal of the Linnean Society, 160, 457–481. https://doi.org/10.1111/j.1096-3642.2009.00606.x
- Villalobos-Hiriart, J.L. (2005) Sistemática de los cangrejos de agua dulce de México, Tribu Pseudotelphusini Ortman, 1897, (Crustacea: Decapoda: Brachyura: Pseudothelphusidae). Análisis filogenético, biogeográfico y descripción de nuevas especies. Ph.D Thesis, Instituto de Biología, Universidad Nacional Autónoma de México, Mexico City, 391 pp.
- Villalobos-Hiriart, J.L. & Álvarez, F. (2008) Los cangrejos de agua dulce de la familia Pseudothelphusidae (Decapoda: Brachyura: Eubrachyura) de México. Con un apéndice de las especies citadas para América hasta el 2006. In: Álvarez, F. y Rodríguez, G. (Eds.), Crustáceos de México: Estado actual de su conocimiento. UANL-PROMEP-SEP. Dirección General de Publicaciones, Universidad Autónoma de Nuevo León, Monterrey, Nuevo León, pp. 239–299.
- Villalobos-Hiriart, J.L., Cantú Díaz-Barriga, A. & Lira-Fernández, E. (1993) Los crustáceos de agua dulce de México. Revista de la Sociedad de Mexicana de Historia Natural Especial, 44, 267–290.
- Villalobos-Hiriart, J.L., Kamanli, S.A., Álvarez, F., Garbout, A. & Clark, P.F. (2019) Lobithelphusa mexicana Rodriguez, 1982 (Decapoda: Pseudothelphusidae): a reassessment of key characters and systematics. Zootaxa, 4586 (1), 151–161. https://doi.org/10.11646/zootaxa.4586.1.8
- Webb, S.A., Graves, J.A., Macías-García, C., Magurran, A.E., Foighil, D.Ó. & Ritchie, M.G. (2004) Molecular phylogeny of the livebearing Goodeidae (Cyprinodontiformes). Molecular Phylogenetics and Evolution, 30, 527–544. https://doi.org/10.1016/S1055-7903(03)00257-4
- Webster, M. & Sheets, H.D. (2010) A practical introduction to landmark-based geometric morphometrics. The Paleontological Society Papers, 16, 163–188. https://doi.org/10.1017/S1089332600001868
- Young, C.G. (1900) The stalked-eyed Crustacea of British Guiana, West Indies and Bermuda. John M. Watkins, London, 514 pp. https://doi.org/10.5962/bhl.title.10670
- Zelditch, M.L., Bookstein, F.L. & Lundrigan, B.L. (1992) Ontogeny of integrated skull growth in the cotton rat Sigmodon fulviventer. Evolution, 46, 1164–1180. https://doi.org/10.2307/2409763
- Zelditch, M.L., Swiderski, D.L. & Sheets, H.D. (2004) Geometric morphometrics for biologists: a primer. Elsevier Academic Press, San Diego, California, 443 pp.