Preferencia de microhábitat de Pseudoscorpiones (Arachnida) en bosques de mangle del sur del golfo de Morrosquillo, Caribe colombiano
DOI:
https://doi.org/10.22201/ib.20078706e.2021.92.3378Palabras clave:
Ecología, Microhábitat, Distribución, San Antero, CórdobaResumen
Los pseudoescorpiones son arácnidos que pueden ocupar diferentes microhábitats, algunos especializados. Durante una investigación de pseudoescorpiones, se evaluó la preferencia de microhábitat en bosques de mangle costeros de Córdoba, Caribe colombiano. Un total de 1,063 individuos fueron recolectados, distribuidos en 4 familias y 8 especies. Las especies más frecuentes en cada microhábitat fueron Pachyolpium isolatum (n = 720) y Lechytia chthoniiformis (n =190) y la menos frecuente Parachernes aff. setosus (n = 6). Los estimadores no paramétricos jackknife 1, jackknife 2 y bootstrap indicaron una buena representatividad; se registró 98.9% de la fauna y se estimó un potencial de 11 especies para los fragmentos de bosque de mangle. La hojarasca fue el microhábitat que presentó el mayor promedio de individuos 56 ± 6.7 ind/mes, y el cascajo de coral presentó el menor promedio 1.8 ± 0.9 ind/mes. La salinidad del suelo, su humedad y la humedad de la hojarasca, son las variables que pueden influir significativamente el ensamblaje de pseudoescorpiones en los bosques de mangle. Las medidas de selectividad y/o preferencias establecieron una alta especificidad con cada microhábitat, contando el suelo, la hojarasca y la corteza de árboles, con mayor frecuencia de uso. Estos resultados destacan la importancia de comprender el papel de estos arácnidos en bosques de mangle y permiten incrementar el conocimiento de su ecología, ofreciendo un potencial uso como bioindicadores.
Citas
Adis, J. (1997). Estratégias de sobrevivência de invertebrados terrestres em florestas inundáveis da Amazônia Central: uma resposta à inundação de longo período. Acta Amazonica, 27, 43–54. https://doi.org/10.1590/1809-43921997271054
Adis, J., Bonaldo, A. B., Brescovit, A. D., Bertani, R., Cokendolpher, J. C., Condé, B. et al. (2002). Arachnida at ‘Reserva Ducke’, central Amazonia/Brazil. Amazoniana,17, 1–14.
Adis, J. y Junk, W. (2002). Terrestrial invertebrates inhabiting lowland river floodplains of Central Amazonia and Central Europe: a review. Freshwater Biology, 47,711–731. https:// doi.org/10.1046/j.1365-2427.2002.00892.x
Adis, J. y Mahnert, V. (1985). On the natural history and ecology of pseudoscorpiones (Arachnida) from an Amazonian backwater inundation forest. Amazoniana, 9, 297–314.
Adis, J. y Mahnert, V. (1990). Vertical distribution and abundance of pseudoscorpions species (Arachnida) in the soil of a neotropical secondary forest during the dry and the rainy season. Acta Zoologica Fennica, 190,11–16.
Adis, J. y Mahnert, V. (1993). Vertical distribution and Abundance of pseudoscorpions (Arachnida) in the soil of two different neotropical primary forests during the dry and rainy seasons. Memoirs of the Queensland Museum, 33, 431–440.
Adis, J., Mahnert, V., Moráis, J. W. y Rodríguez, J. M. (1988). Adaptation of an Amazoniaian pseudoscorpion (Arachnida) from dryland forests to inundation forests. Ecology, 69, 287–291.
Aguiar, N. O. y Bührnheim, P. F. (2003). Pseudoscorpiones (Arachnida) da vegetaca de sub-bosque da floresta primaria tropical de terra firme (Coari, Amazonas, Brasil). Acta Amazonica, 33, 515–526. https://doi.org/10.1590/
S0044-59672003000300016
Aguiar, N. O., Gualberto, T. y Franklin, E. (2006). A medium spatial scale distribution pattern of Pseudoscorpionida (Arachnida) in a gradient of topography (altitude and inclination), soil factors, and litter in a central Amazonia forest reserve, Brazil. Brazilian Journal of Biology, 66, 791–
https://doi.org/10.1590/S1519-69842006000500004
Andrade, R. y Gnaspini, P. (2002). Feeding in Maxchernes iporangae (Pseudoscorpiones, Chernetidae) in captivity. Journal of Arachnology, 30, 613–617.
Barros, Y. J., Melo, V. F., Sautter, K. D., Buschle, B., de Oliveira, E. B. et al. (2010). Indicadores de qualidade de solos de área de mineração e metalurgia de chumbo: II - Mesofauna e plantas. Revista Brasileira de Ciência do Solo, 34, 1413–1426. https://doi.org/10.1590/S0100-06832010000400037.
Bastienne, S., Gayoso, J. y Guerra, J. (2001). Manual de procedimientos para inventarios de carbono en ecosistemas forestales. Valdivia: Universidad Austral de Chile.
Battirola, L. D., Adis, J., Marques, M. I. y Silva, F. H. O. (2007). Composição da comunidade de artrópodes associada à copa de Attalea phalerata Mart. (Arecaceae), durante o período de cheia no Pantanal de Poconé, Mato Grosso, Brasil. Neotropical Entomology, 36, 640–651. https://doi.org/10.1590/S1519-566X2007000500002
Battirola, L. D., Marques, M. I., Brescovit, A. D., Rosado-Neto, G. H. y Anjos, K. C. (2010). Comunidade edáfica de Araneae (Arthropoda, Arachnida) em uma floresta sazonalmente inundável na região norte do Pantanal de Mato Grosso, Brasil. Biota Neotropica, 10, 173–183.http://dx.doi.org/10.1590/S1676-06032010000200022
Battirola, L. D., Marques, M. I., Rosado-Neto, G. H., Pinheiro, T. G. y Pinho, N. G. C. (2009). Vertical and time distribution of Diplopoda (Arthropoda: Myriapoda) in a monodominant forest in Pantanal of Mato Grosso, Brazil. Zoologia, 26, 479–487. http://dx.doi.org/10.15/S1984-46702009005000008
Battirola, L. D., Rosado-Nieto, G. H., Batistella, D. A., Mahnert, V., Brescovit, A. D. y Marques, M. I. (2017). Vertical and time distribution of Pseudoscorpiones (Arthropoda: Arachnida) in a floodplain forest in the Brazilian Pantanal.Revista de biologia tropical, 65, 445–459. http://dx.doi.
org/10.15517/rbt.v65i2.24134
Batzli, G. O. y Lesieutre, C. (1991). The influence of high quality food on habitat use by arctic microtine rodents. Oikos, 60, 299–306. https://doi.org/10.2307/3545071
Bedoya-Roqueme, E. (2015). Una nueva especie de Paraliochthonius (Pseudoscorpiones: Chthoniidae: Tyrannochthoniini) de Colombia. Revista Ibérica de Aracnología, 26, 49–54.
Bedoya-Roqueme, E. (2019). Pseudoscorpiones of the tribe Chernetini (Chernetidae) from the Colombian Caribbean. New species and an identification key. Revista Ibérica de Aracnología, 34, 21–40.
Bedoya-Roqueme, E. y Quirós-Rodríguez, J. A. (2018). A new species of Antillochernes (Pseudoscorpiones: Chernetidae: Chernetini) from Colombia. Revista Ibérica de Aracnología, 32, 93–99.
Bedoya-Roqueme, E. y Quirós-Rodríguez, J. A. (2019). Pseudoscorpiones (Arachnida) de Isla Fuerte y Tortuguilla, Caribe colombiano. Acta Biológica Colombiana, 24, 163–173. http://dx.doi.org/10.15446/abc.v24n1.70560
Bedoya-Roqueme, E., Pérez-Agudelo, M., Zaragoza, J.A. y Quirós-Rodríguez, J.A. (2017). Nuevos reportes de Pseudoescorpiones (Arachnida) de bosques de manglar en Córdoba, Caribe colombiano. Revista Ibérica de Aracnología, 30, 25–36.
Bedoya-Roqueme, E., Salleg-Pérez, G. y Quirós-Rodríguez, J. A. (2016). Sobre la ecología de pseudoescorpiones (Arachnida:Pseudoscorpiones) en bosques de manglar del sur del Golfo de Morrosquillo, Caribe colombiano. Revista Ibérica de Aracnología, 28, 65–74.
Belozerov, V. N. (2013). Seasonal Aspects of the Life Cycles of Pseudoscorpions (Arachnida, Pseudoscorpiones). Entomological Review, 93, 634–652. https://doi.org/10.1134/S0013873813050126
Benavides, L. R, Cosgrove, J. G., Harvey, M. S. y Giribet, G. (2019). Phylogenomic interrogation resolves the backbone of the Pseudoscorpiones tree of life. Molecular Phylogenetics and Evolution, 139, 106509. https://doi.org/10.1016/j.ympev.2019.05.023
Bray, R. J. y Curtis, J. T. (1975). An ordination of the upland forest communities of southern of Wisconsin. Ecological monographs, 27, 325–349. https://doi.org/10.2307/1942268
Briano, M., Ortiz, A. y Torres, R. (2010). Introducción al conocimiento de los pseudoscorpiones del estado de Aguas Calientes, México. Investigación y ciencia de la Universidad Autónoma de Aguas Calientes, 50, 5–9.
Broza, M., Poliakov, D., Weber, S. y Izhaki, I. (1993). Soil microarthropods on postfire pine forests on Mount Carmel, Israel. Water Science and Technology, 27, 533–538. https://doi.org/10.2166/wst.1993.0592
Buddle, C. (2010). Photographic key to the Pseudoscorpions of Canada and the adjacent USA. Canadian Journal of Arthropod Identification, 10, 1–77. https://doi.org/10.3752/cjai.2010.10
Calenge, C. (2006). The package adehabitat for the R software:tool for the analysis of space and habitat use by animals. Ecological Modelling, 197, 1035. https://doi.org/10.1016/j.ecolmodel.2006.03.017
Castilho, A. C. C., Marques, M. I., Adis, J. y Brescovit, A. D. (2005). Distribuição sazonal e vertical de Araneae em área com predomínio de Attalea phalerata Mart. (Arecaceae), no Pantanal de Poconé, Mato Grosso, Brasil. Amazoniana, 18,215–239.
Clarke, K. y Gorley, R. (2005). PRIMER v.6: user manual/tutorial. Plymouth, United Kingdom: PRIMER–E Ltda.
Cole, D. C., Elgar, M. A. y Harvey, M. S. (1995). Associations between Australian pseudoscorpions and ants. Psyche Cambridge, Mass, 101, 221–227. https://doi.org/10.1155/1994/23982
Colwell, R. (2013). EstimateS: Statistical estimation of species richness and shared species from samples Version 9.1.0. User’s Guide and application. Recuperado 19 de febrero de 2019 de: http://www.viceroy.eeb.uconn.edu/estimates
Cortés, D. V. y Rangel, J. O. (2011). Los bosques de mangle en un gradiente de salinidad en la Bahía de Cispatá - Boca Tinajones, departamento de Córdoba-Colombia. Caldasia, 33, 155–176.
CVS-INVEMAR (Corporación Autónoma Regional de los Valles del Sinú y del San Jorge-Instituto de Investigaciones Marinas y Costeras). (2010). Plan integral de manejo del distrito de Manejo Integrado (DMI) bahía de Cispatá - La Balsa - Tinajones y sectores aledaños del delta estuarino del río Sinú, departamento de Córdoba. Santa Marta: Serie de Publicaciones Especiales de INVEMAR
Del-Claro, K. y Tizo-Pedroso, E. (2009). Ecological and evolutionary pathways of social behavior in Pseudoscorpions (Arachnida: Pseudoscorpiones). Acta Ethologica, 12, 13–22. https://doi.org/10.1007/s10211-009-0052-y
Donovan, B. y Paul, F. (2005). Pseudoscorpions: the forgotten beneficials inside beehives and their potential for management for control of varroa and other arthropod pests. Bee World, 86, 83–87. https://doi.org/10.1080/0005772X.2005.11417322
Etchevers, J. (1988). Análisis químico de suelos y plantas. Centro de Edafología. Chapingo, Estado de México: Colegio de Posgraduados.
Félix-Angulo, A.G. (2018). Diversidad de pseudoescorpiones (Arachnida: Pseudoscorpiones) en bosque de mangle en la localidad el conchal, Culiacán, Sinaloa. Boletin de la Sociedad Mexicana de Entomología, 4, 5–8.
Francke, O. F. y Villegas-Guzmán, G. A. (2006). Symbiotic relationships between Pseudoscorpions (Arachnida) and packrats (Rodentia). Journal of Arachnology, 34, 289–298.https://doi.org/10.1636/04-36.1
Gabbutt, P. (1962). Nets of the marie falsescorpion. Nature London, 196, 97–98.
Gabbutt, P. (1970). Sampling problems and the validity of the life history analyses of pseudoscorpions. Journal of Natural History, 4, 1–15. https://doi.org/10.1080/00222937000770011
Garshelis, D. L. (2000). Delusions in habitat evaluation: measuring use, selection, and importance. En L. Boitani y T. K. Fuller (Eds.), Research techniques in animal ecology:controversies and consequences. (pp. 111–164). New York: Columbia University.
Gerlach, J., Samways, M. y Pryke, J. (2013). Terrestrial invertebrates as bioindicators: an overview of available taxonomic groups. Journal of Insect Conservation, 17, 831–850. https://doi.org/10.1007/s10841-013-9565-9
González, V. H., Mantilla, B. y Mahnert, V. (2007). A new record for Dasychernes inquilinus (Arachnida, Pseudoscorpiones, Chernetidae), with an overview of pseudoscorpion-bee relationships. Journal of Natural History, 35, 470–474. https://doi.org/10.1636/H06-62.1
González-Megias, A., Gómes, J. G. y Sánchez-Piñero, F. (2007). Diversity-habitat heterogeneity relationship at different spatial and temporal scales. Ecography, 30, 31–41. https://doi.org/10.1111/j.0906-7590.2007.04867.x
Harvey, M. S. (1992). The phylogeny and classification of the Pseudoscorpionida (Chelicerata: Arachnida). Invertebrate Taxonomy, 6, 1373–1435. https://doi.org/10.1071/IT9921373
Harvey, M. S. (2009). The first Australasian species of the genus pseudoscorpion halophilic Paraliochthonius (Pseudoscorpiones: Chthoniidae). Records of the Australian Museum, 25, 329–344. http://dx.doi.org/10.18195/issn.0312-3162.25(3).2009.329-344
Harvey, M. S. (2013). Pseudoscorpions of the World, version 2.0. Western Museum Australian, Perth. Recuperado el 21 de enero de 2020 de: http://www.museum.wa.gov.au/catalogs/pseudoscorpion
Harvey, M. S., Waldock, J., Teale, R. y Webber, J. (2007). New distribution records of the intertidal pseudoscorpion Parahya submersa (Pseudoscorpiones: Parahyidae). Records of the Australian Museum, 23, 393–395. http://dx.doi.org/10.18195/issn.0312-3162.23(4).2007.393-395
Hoff, C. C. (1959). The ecology and distribution of the pseudoscorpions of north–central New Mexico. University New Mexico Publishing Biological, 8, 1–68.
Hoff, C. C. (1964). The pseudoscorpions of Jamaica. Part 3. The suborder Diplosphyronida. Bulletin of the Institute of Jamaica. Science series, 10, 1–47.
INVEMAR (Instituto de Investigaciones Marinas y Costeras). (2012). Informe del estado de los ambientes y recursos marinos y costeros en Colombia: año 2011. Santa Marta: Serie de Publicaciones Periódicas, 8a Ed.
Judson, M. L. I. (1992). A simple, slow-diffusion method for clearing small arthropods. The Newsletter British Arachnological Society, 64, 6–7.
Judson, M. L. I. (2005). Baltic amber fossil of Garypinus electri Beier provides first evidence of phoresy in the pseudoscorpion family Garypinidae (Arachnida: Chelonethi). En D. V. Logunov y D. Penney (Eds.), European Arachnology 2003 (Proceedings of the 21st European Colloquium of Arachnology, St. Petersburg, 4-9 August 2003 (pp. 127–131). Moscú: KMK Scientific Press.
Judson, M. L. I. (2016). Pseudoscorpions (Arachnida, Chelonethi) in Mexican amber, with a list of extant species associated with mangrove and Hymenaea trees in Chiapas. Boletín de la Sociedad Geológica Mexicana, 68, 57–79. https://doi.org/10.5281/zenodo.50595
Junk, W. J., Bayley, P. B. y Sparks, R. E. (1989). The flood pulse concept in river-floodplain systems. En D. P. Dodge (Eds.),Proceedings of the International Large River Symposium (LARS) (pp.110–127). Ontario, Canadá: Department of Fisheries and Oceans.
Kensler, C. (1967). Dessication resistance of intertidal species as a factor of their zonation. Journal of Animal Ecology, 3, 391–406.
Krausman, R. P. (1999). Some basic principles of habitat use, grazing behavior of livestock and wildlife. Idaho Forest. Wildlife and Range Experiment Station Bulletin, 70, 85–90.
Lee, V. F. (1979). The maritime pseudoscorpions of Baja California, México (Arachnida: Pseudoscorpionida). Occasional papers of the California Academy of Sciences, 131, 1–38.
Lencinas, M. V., Kreps, G., Soler, R., Peri, P. L., Porta, A., Ramírez, M. et al. (2015). Neochelanops michaelseni (Pseudoscorpiones: Chernetidae) as a potential bioindicator in managed and unmanaged Nothofagus forests of Tierra del Fuego. Journal of Arachnology, 43, 406–412.
Litvaitis, J. A., Titus, K. y Anderson, E. M. (1994). Measuring vertebrate use of territorial habitats and foods. En T. A. Bookhout (Eds.), Research and management techniques for wildlife and habitats. 5th edition (pp. 254–274), Bethesda, Maryland: The Wildlife Society.
Lubin, Y., Ellner, S. y Kotzman, M. (1993). Web relocation and habitat selection in a desert widow spider. Ecology, 74, 2456–2459.https://doi.org/10.2307/1940835
Mahnert, V. (2014). Pseudoscorpions (Arachnida: Pseudoscorpiones) from the Galapagos Islands (Ecuador).Revue suisse de Zoologie, 121, 135–210.
Mahnert, V. y Adis, J. (2002). Pseudoscorpiones. En J. Adis(Eds), Amazonian Arachnida and Myriapoda (pp. 367–380). Sofía, Moscú: Pensoft Publisher.
Mahnert, V. y Schuster, R. (1981). Pachyolpium atlanticum sp. nov., ein Pseudoskorpion aus der Gezeitenzone der Bermudas-Morphologie und Ökologie (Pseudoscorpiones: Olpiidae). Revue Suisse de Zoologie, 88, 265–273. https://doi.org/10.5962/bhl.part.82371
Manly, B. F. J. (1974). A model for certain types of selection experiments. Biometrics, 30, 281–94.
Manly, B. F. J., McDonald, L. L, Thomas, D. L., McDonald, T. L. y Erickson, W. P. (2003). Resource selection by animals statistical design and analysis for field studies. Países Bajos: Springer.
Marques, M. I., Adis, J., Battirola, L. D., Santos, G. B. y Castilho, A. C. C. (2011). Arthropods associated with a forest of Attalea phalerata Mart. (Arecaceae) palm trees in the northern Pantanal of the Brazilian state of Mato Grosso. En W. J. Junk, C. J. Da-Silva, C. Nunes-da Cunha y K. M. Wantzen (Eds.), The Pantanal: ecology, biodiversity and sustainable management of a large neotropical seasonal wetland (pp. 431–468). Sofia, Moscú: Pensoft Publishers.
McGeoch, M. A., Sithole, H., Samways, M. J., Simaika, J. P., Pryke, J. S., Picker, M. et al. (2011). Conservation and monitoring of invertebrates in terrestrial protected areas. Koedoe, 53, 1–13.https://doi.org/10.4102/koedoe.v53i2.1000
Moráis, J. W., Adis, J., Mahnert, V. y Berti-Filho, E. (1997). Abundance and phenology of Pseudoscorpiones (Arachnida) from a mixedwater inundation forest in Central Amazonia, Brazil. Revue suisse de Zoologie, 104, 475–483. http://dx.doi.org/10.5962/bhl.part.80005
Montenegro, J. y Acosta, A. (2008). Programa innovador para evaluar uso y preferencia de hábitat. Universitas Scientiarum, 13, 208–217.
Morris, D. W. (2003). Toward an ecological synthesis: a case for habitat selection. Oecologia, 136, 1–13. https://doi.org/10.1007s00442-003-1241-4
Morrison, M. L., Block, W. M., Strickland, M. D., Collier, B. A. y Peterson, M. J. (2008). Wildlife study design (2a. ed.). New York: Springer Series on Environmental Management.
Morrison, M. L., Marcot, B. G. y Mannan, R. W. (1998). Wildlife habitat relationships: concepts and applications (2a. ed.). Madison, Wisconsin: The University of Wisconsin.
Muchmore, W. B. (1976). Pseudoscorpions from Florida and the Caribbean Area. 5. Americhernes, a new genus based upon Chelifer elongatus Say (Chernetidae). Florida Entomologist, 59, 151–163.
Muñoz, J., Mendoza, A., López, F., Soler, A. y Hernández, M. (2000). Manual de análisis de suelo. Edafología. México D.F.: Escuela Nacional de Estudios Profesionales Iztacala,UNAM.
Poinar, G. O., Ćurčić, J. R. B. P. M. y Cokendolpher, J. C. (1998). Arthropod phoresy involving pseudoscorpions in thepast and present. Acta Arachnologica, 47, 79–96. https://doi.org/10.2476/asjaa.47.79
Quirós-Rodríguez, J. A. y Arias, J. E. (2013). Taxocenosis de moluscos y crustáceos en raíces de Rhizophora mangle (Rhizophoracea) en la bahía de Cispatá, Córdoba, Colombia. Acta Biológica Colombiana, 18, 329–340.
R Core Team. (2020). R: a language and environment for statistical computing. R Foundation for Statistical Computing, Vienna, Austria. Recuperado el 20 de diciembre, 2019 de: http://www.R–project.org/
Ranius, T., Johansson, V. y Fahrig, L. (2011). Predicting spatial occurrence of beetles and pseudoscorpions in hollow oaks in southeastern Sweden. Biodiversity and Conservation, 20, 2027–2040. https://doi.org/10.1007/s10531-011-0072-6
Read, S., Howlett, B. G., Donovan, B. J., Nelson, W. R. y Van Toor, R. F. (2014). Culturing chelifers (Pseudoscorpions) that consume Varroa mites. Journal of Applied Entomology, 138, 260–266. https://doi.org/10.1111/jen.12096
Sánchez, L. E, Paolini, J. y Rodríguez, J. (2010). Dinámica de las propiedades del suelo en bosques de Rhizophora mangle L. (Rhizophoraceae) en Isla de Margarita, Venezuela. Revista Biología Tropical, 58, 547–564. https://doi.org/10.15517/rbt.v58i2.5228
Schuster, R. (1962). Das marine litoral als lebensraum terrestrischer kleinarthropoden. Internationale Revue der gesamten Hydrobiologie und Hydrographie, 47, 359–412. https://doi.org/10.1002/iroh.19620470304
Tews, J., Brose, U., Grimm, V., Tielborger, K., Wichmann, M. C., Schwager, M. et al. (2004). Animal species diversity driven by habitat heterogeneity/diversity: the importance of keystone structures. Journal of Biogeography, 31, 79–92. https://doi.org/10.1046j.0305-0270.2003.00994.x
Tizo-Pedroso, E. y Del-Claro, K. (2011). Is there division of labor in cooperative pseudoscorpions? An analysis of the behavioral repertoire of a tropical species. Ethology, 117, 498–507. https://doi.org/10.1111/j.1439-0310.2011.01906.x
Tooren, D. (2011). New records of olpiid pseudoscorpions (Pseudoscorpiones: Olpiidae) from the Caribbean area and Surinam, with descriptions of four new species of the genera Pachyolpium Beier, Tricholpium gen. nov. and Heterohorus gen. nov. Zoologische Mededelingen, 85, 29, 169–194.
Topp, G. (1993). Soil water content. En M. R. Carte (Ed.), Soil sampling and methods of analysis (pp. 541–557). Florida, Boca Raton: Lewis Publishers.
Underwood, A. J., Chapman, M. G. y Crowe, T. P. (2004). Identifying and understanding ecological preferences for hábitat or prey. Journal of Experimental Marine Biology and Ecology, 300, 161–187. https://doi.org/10.1016/j. jembe.2003.12.006
Weygoldt, P. (1969). The biology of pseudoscorpions. Cambridge:Harvard University.
Willard, H., Merrit, L. y Dean, A. (1974). Instrumental methods of analysis. 5th edition. New York: Van Nostrand. Yamamoto, T., Nakagoshi, N. y Touyama, Y. (2001). Ecological study of pseudoscorpion fauna in the soil organic layer in managed and abandoned secondary forests. Ecological Research, 16, 593–601. https://doi.org/10.1046/j.1440-1703.2001.00422.x
Zeh, D. W. y Zeh, J. A. (1992a). Dispersal–generated sexual selection in a beetle–riding pseudoscorpion. Behavioral Ecology and Sociobiology, 30, 135–142. https://doi.org/10.1007/BF00173949
Zeh, D. W. y Zeh, J. A. (1992b). On the function of harlequin beetleriding in the pseudoscorpion, Cordylochernes scorpioides (Pseudoscorpionida: Chernetidae). Journal of Arachnology, 20, 47–51.
Zeh, D. W. y Zeh, J. A. (1997). Sex via the substrate: mating system and sexual selection in pseudoscorpions. En J. C. Choe y B. J. Crespi (Eds.), The evolution of mating systems in insects and arachnids (pp. 329–339). Cambridge:Cambridge University.
