Ninetine spiders in Brazilian Caatinga and Cerrado: revision of Kambiwa and description of Sertana gen. nov. (Araneae, Pholcidae), with analyses of predicted range shifts due to climate change

  • Bernhard A. Huber Zoological Research Museum Alexander Koenig, LIB, Bonn, Germany https://orcid.org/0000-0002-7566-5424
  • Guanliang Meng Zoological Research Museum Alexander Koenig, LIB, Bonn, Germany
  • Jiří Král Department of Genetics and Microbiology, Faculty of Science, Charles University, Prague, Czech Republic
  • Ivalú M. Ávila Herrera Department of Genetics and Microbiology, Faculty of Science, Charles University, Prague, Czech Republic
  • Leonardo S. Carvalho Campus Amílcar Ferreira Sobral, Universidade Federal do Piauí, Floriano, Piauí, Brazil
DOI: https://doi.org/10.5852/ejt.2026.1054.3276
Keywords: CO1 barcodes, climate change, karyotype, niche modeling, taxonomy

Abstract

Among daddy long-legs spiders (Pholcidae), Ninetinae is a distinctive subfamily that comprises short-legged, fast-running spiders. Most species are small or tiny, lead reclusive lives, and are largely restricted to semiarid regions, which together has made them poorly collected and poorly known. Here, we build on focused recent collections in the Brazilian Cerrado and Caatinga biomes, two of the World’s richest tropical savanna, xeric shrubland and thorn forest regions. Our focus is on the taxonomy of the genus Kambiwa Huber, 2000 that previously contained only two nominal species, each known from a single locality. Combining morphological and molecular (CO1 barcode) data, we describe six new species in Kambiwa (K. brumado Huber sp. nov.; K. coribe Huber sp. nov.; K. ibo Huber sp. nov.; K. itacarambi Huber sp. nov.; K. maracas Huber sp. nov.; K. mucuge Huber sp. nov.), redescribe the type species K. neotropica (Kraus, 1957), and synonymize the monotypic genus Pemona Huber, 2019 with Kambiwa, resulting in the new combination K. sapo (Huber, 2019) comb. nov. In addition, we describe a new genus of superficially Kambiwa-like spiders from the same geographic region: Sertana Huber gen. nov., with five new species (S. bumba Huber gen. et sp. nov.; S. capivara Huber gen. et sp. nov.; S. igapora Huber gen. et sp. nov.; S. lapa Huber gen. et sp. nov.; S. sagarana Huber gen. et sp. nov.). In line with previous efforts to explore the processes underlying the geographical distribution of Ninetinae, we also evaluate the potential effects of future climate change on the environmental niche occupied by three selected species of Kambiwa. Our results corroborate previous findings that demonstrate an altitude-mediated response to climate change. For a highland species, areas of high habitat suitability almost disappear under more severe climate change scenarios. For two species with lowland records, the areas with high habitat suitability increase significantly. Finally, we analyze the male karyotype of K. ibo which consists of 28 chromosomes including a X1X2X3Y system. All chromosomes are biarmed except for the Y chromosome. This contribution concludes a series of publications on the subfamily Ninetinae. We use this opportunity to summarize current knowledge about the subfamily, to discuss open questions and knowledge gaps, and to suggest further research topics focusing on these tiny but exceptional pholcids.

References

Aharon S., Huber B.A. & Gavish-Regev E. 2017. Daddy-long-leg giants: revision of the spider genus Artema Walckenaer, 1837 (Araneae, Pholcidae). European Journal of Taxonomy 376: 1–57. https://doi.org/10.5852/ejt.2017.376

Álvares E.S.S., Machado E.O., Azevedo C.S. & De-Maria M. 2004. Composition of the spider assemblage in an urban forest reserve in southeastern Brazil and evaluation of two sampling method protocols of species richness estimates. Revista Ibérica de Aracnología 10: 185–194. Available from https://dialnet.unirioja.es/servlet/articulo?codigo=1088718 [accessed 22 Oct. 2025].

Araujo H.F.P., Canassa N.F., Machado C.C.C. & Tabarelli M. 2023. Human disturbance is the major driver of vegetation changes in the Caatinga dry forest region. Scientific Reports 13: 18440. https://doi.org/10.1038/s41598-023-45571-9

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

Ávila Herrera I.M., Král J., Pastuchová M., Forman M., Musilová J., Kořínková T., Šťáhlavský F., Zrzavá M., Nguyen P., Just P., Haddad C.R., Hiřman M., Koubová M., Sadílek D. & Huber B.A. 2021. Evolutionary pattern of karyotypes and meiosis in pholcid spiders (Araneae: Pholcidae): implications for reconstructing chromosome evolution of araneomorph spiders. BMC Ecology and Evolution 21: 75. https://doi.org/10.1186/s12862-021-01750-8

Bachtrog D. 2013. Y-chromosome evolution: emerging insights into processes of Y-chromosome degeneration. Nature Reviews Genetics 14: 113–124. https://doi.org/10.1038/nrg3366

Barbet-Massin M., Jiguet F., Albert C.H. & Thuiller W. 2012. Selecting pseudo-absences for species distribution models: how, where and how many? Methods in Ecology and Evolution 3: 327–338. https://doi.org/10.1111/j.2041-210X.2011.00172.x

Bengtson P. 1988. Open nomenclature. Palaeontology 31: 223–227.

Beuchle R., Grecchi R.C., Shimabukuro Y.E., Seliger R., Eva H.D., Sano E. & Achard F. 2015. Land cover changes in the Brazilian Cerrado and Caatinga biomes from 1990 to 2010 based on a systematic remote sensing sampling approach. Applied Geography 58: 116–127. https://doi.org/10.1016/j.apgeog.2015.01.017.

Boyce M.S., Vernier P.R., Nielsen S.E. & Schmiegelow F.K.A. 2002. Evaluating resource selection functions. Ecological Modelling 157: 281–300. https://doi.org/10.1016/S0304-3800(02)00200-4

Brown B.V. 1993. A further chemical alternative to critical-point-drying for preparing small (or large) flies. Fly Times 11: 10.

Carvalho L.S. & Avelino M.T.L. 2010. Composição e diversidade da fauna de aranhas (Arachnida, Araneae) da Fazenda Nazareth, Município de José de Freitas, Piauí, Brasil. Biota Neotropica 10: 21–31. https://doi.org/10.1590/S1676-06032010000300001

Carvalho L.S., Sanches Ruiz G.R., Saturnino R. & Carvalho L.S. 2026. Optimizing sampling protocols: spider assemblages are robust to pitfall trap spacing in Neotropical savannas. Community Ecology 27: 149–160. https://doi.org/10.1007/s42974-025-00287-w

Castanheira P., Pérez-González A. & Baptista R.L.C. 2016. Spider diversity (Arachnida: Araneae) in Atlantic Forest areas at Pedra Branca State Park, Rio de Janeiro, Brazil. Biodiversity Data Journal 4: e7055. https://doi.org/10.3897/BDJ.4.e7055

Churchill T.B. 1993. Effects of sampling method on composition of a Tasmanian coastal heathland spider assemblage. Memoirs of the Queensland Museum 33: 475–481.

Colli G.R., Vieira C.R. & Dianese J.C. 2020. Biodiversity and conservation of the Cerrado: recent advances and old challenges. Biodiversity and Conservation 29: 1465–1475. https://doi.org/10.1007/s10531-020-01967-x

Costa-Pinto A.L. da, Bovendorp R.S., Heming N.M., Malhado A.C. & Ladle R.J. 2024. Where could they go? Potential distribution of small mammals in the Caatinga under climate change scenarios. Journal of Arid Environments 221: 105133. https://doi.org/10.1016/j.jaridenv.2024.105133

Dantas L.G., dos Santos C.A.C., Santos C.A.G., Martins E.S.P.R. & Alves L.M. 2022. Future changes in temperature and precipitation over northeastern Brazil by CMIP6 Model. Water 14: 4118. https://doi.org/10.3390/w14244118

de Paula Y.R.A.A., Bustamante A.A., Saturnino R. & Carvalho L.S. 2026. Influence of habitat complexity on the ground spider community in gallery forests of the Cerrado biome. Acta Oecologica 131: 104165. https://doi.org/10.1016/j.actao.2026.104165

Dederichs T.M., Huber B.A. & Michalik P. 2022. Evolutionary morphology of sperm in pholcid spiders (Pholcidae, Synspermiata). BMC Zoology 7: 52. https://doi.org/10.1186/s40850-022-00148-3

Dolejš P., Kořínková T., Musilová J., Opatová V., Kubcová L., Buchar J. & Král J. 2011. Karyotypes of central European spiders of the genera Arctosa, Tricca and Xerolycosa (Araneae: Lycosidae). European Journal of Entomology 108: 1–16. https://doi.org/10.14411/eje.2011.001

Eberhard W.G. 1992. Web construction by Modisimus sp. (Araneae, Pholcidae). Journal of Arachnology 20: 25–34.

Eberle J., Dimitrov D., Valdez-Mondragón A. & Huber B.A. 2018. Microhabitat change drives diversification in pholcid spiders. BMC Evolutionary Biology 18: 141. https://doi.org/10.1186/s12862-018-1244-8

Eyring V., Bony S., Meehl G.A., Senior C., Stevens B., Stouffer R.J. & Taylor K.E. 2015. Overview of the Coupled Model Intercomparison Project Phase 6 (CMIP6) experimental design and organisation. Geoscientific Model Development 9: 1937–1958. https://doi.org/10.5194/gmd-9-1937-2016

Fick S.E. & Hijmans R.J. 2017. WorldClim 2: new 1-km spatial resolution climate surfaces for global land areas. International Journal of Climatology 37: 4302–4315. https://doi.org/10.1002/joc.5086

Fielding A.H. & Bell J.F. 1997. A review of methods for the assessment of prediction errors in conservation presence/absence models. Environmental Conservation 24: 38–49. https://doi.org/10.1017/S0376892997000088

Firpo M.Â.F., Guimarães B. dos S., Dantas L.G., Silva M.G.B. da, Alves L.M., Chadwick R., Llopart M.P. & Oliveira G.S. de. 2022. Assessment of CMIP6 models’ performance in simulating present-day climate in Brazil. Frontiers in Climate 4: 948499. https://doi.org/10.3389/fclim.2022.948499

Forman M., Nguyen P., Hula P. & Král J. 2013. Sex chromosome pairing and extensive NOR polymorphism in Wadicosa fidelis (Araneae: Lycosidae). Cytogenetic and Genome Research 141: 43–49. https://doi.org/10.1159/000351041

Gonçalves G.S.R., Cerqueira P.V., Silva D.P., Gomes L.B., Leão C.F., Andrade A.F.A. de & Santos M.P.D. 2023. Multi-temporal ecological niche modeling for bird conservation in the face of climate change scenarios in Caatinga, Brazil. PeerJ 11: e14882. https://doi.org/10.7717/peerj.14882

Guindon S., Dufayard J.-F., Lefort V., Anisimova M., Hordijk W. & Gascuel O. 2010. New algorithms and methods to estimate maximum-likelihood phylogenies: assessing the performance of PhyML 3.0. Systematic Biology 59: 307–321. https://doi.org/10.1093/sysbio/syq010

Hijmans R.J. 2023a. raster: Geographic Data Analysis and Modeling. Available from https://cran.r-project.org/web/packages/raster/raster.pdf [accessed 22 Oct. 2025].

Hijmans R.J. 2023b. terra: Spatial Data Analysis. CRAN: Contributed Packages. Available from https://cran.r-project.org/web/packages/terra/index.html [accessed 22 Oct. 2025].

Hoang D.T., Chernomor O., von Haeseler A., Minh B.Q. & Vinh L.S. 2018. UFBoot2: improving the ultrafast bootstrap approximation. Molecular Biology and Evolution 35: 518–522. https://doi.org/10.1093/molbev/msx281

Hofmann G.S., Cardoso M.F., Alves R.J.V., Weber E.J., Barbosa A.A., de Toledo P.M., Pontual F.B., Salles L. de O., Hasenack H., Cordeiro J.L.P., Aquino F.E. & de Oliveira L.F.B. 2021. The Brazilian Cerrado is becoming hotter and drier. Global Change Biology 27: 4060–4073. https://doi.org/10.1111/gcb.15712

Horton T., Marsh L., Bett B.J., Gates A.R., Jones D.O.B., Benoist N.M.A., Pfeifer S., Simon-Lledó E., Durden J.M., Vandepitte L. & Appeltans W. 2021. Recommendations for the standardisation of open taxonomic nomenclature for image-based identifications. Frontiers in Marine Science 8: 620702. https://doi.org/10.3389/fmars.2021.620702

Huber B.A. 1998. Genital mechanics in some neotropical pholcid spiders (Araneae: Pholcidae), with implications for systematics. Journal of Zoology 244: 587–599. https://doi.org/10.1111/j.1469-7998.1998.tb00063.x

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. https://doi.org/10.1206/0003-0090(2000)254<0001:NWPSAP>2.0.CO;2

Huber B.A. 2001. The pholcids of Australia (Araneae; Pholcidae): taxonomy, biogeography, and relationships. Bulletin of the American Museum of Natural History 260: 1–144. https://doi.org/10.1206/0003-0090(2001)260%3C0001:TPOAAP%3E2.0.CO;2

Huber B.A. 2004. The significance of copulatory structures in spider systematics. In: Schult J. (ed.) Studien zur Theorie der Biologie, Band 5, Biosemiotik - Praktische Anwendung und Konsequenzen für die Einzeldisziplinen: 89–100. VWB-Verlag für Wissenschaft und Bildung, Berlin.

Huber B.A. 2012. Revision and cladistic analysis of the Afrotropical endemic genus Smeringopus Simon, 1890 (Araneae: Pholcidae). Zootaxa 3461: 1–138. https://doi.org/10.11646/zootaxa.3461.1.1

Huber B.A. 2018a. The South American spider genera Mesabolivar and Carapoia (Araneae, Pholcidae): new species and a framework for redrawing generic limits. Zootaxa 4395: 1–178. https://doi.org/10.11646/zootaxa.4395.1.1

Huber B.A. 2018b. Cave-dwelling pholcid spiders (Araneae, Pholcidae): a review. Subterranean Biology 26: 1–18. https://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. 2022. Revisions of Holocnemus and Crossopriza: the spotted-leg clade of Smeringopinae (Araneae, Pholcidae). European Journal of Taxonomy 795: 1–241. https://doi.org/10.5852/ejt.2022.795.1663

Huber B.A. & Carvalho L.S. 2019. Filling the gaps: descriptions of unnamed species included in the latest molecular phylogeny of Pholcidae (Araneae). Zootaxa 4546: 1–96. https://doi.org/10.11646/zootaxa.4546.1.1

Huber B.A. & Meng G. 2024. Old World Micropholcus spiders, with first records of acrocerid parasitoids in Pholcidae (Araneae). ZooKeys 1213: 95–182. https://doi.org/10.3897/zookeys.1213.133178

Huber B.A. & Meng G. 2025. Like grains of sand: Ninetis spiders on the Arabian Peninsula (Araneae: Pholcidae). Zootaxa 5563: 290–335. https://doi.org/10.11646/zootaxa.5563.1.19

Huber B.A. & Rheims C.A. 2011. Diversity and endemism of pholcid spiders in Brazil’s Atlantic Forest, with descriptions of four new species of the Atlantic Forest endemic genus Tupigea (Araneae: Pholcidae). Journal of Natural History 45: 275–301. https://doi.org/10.1080/00222933.2010.524319

Huber B.A., Nuñeza O.M. & Leh Moi Ung C. 2015. Revision, phylogeny, and microhabitat shifts in the Southeast Asian spider genus Aetana (Araneae, Pholcidae). European Journal of Taxonomy 162: 1–78. https://doi.org/10.5852/ejt.2015.162

Huber B.A., Eberle J. & Dimitrov D. 2018. The phylogeny of pholcid spiders: a critical evaluation of relationships suggested by molecular data (Araneae, Pholcidae). ZooKeys 789: 51–101. https://doi.org/10.3897/zookeys.789.22781

Huber B.A., Caspar K. & Eberle J. 2019. New species reveal unexpected interspecific microhabitat diversity in the genus Uthina Simon, 1893 (Araneae: Pholcidae). Invertebrate Systematics 33: 181–207. https://doi.org/10.1071/IS18002

Huber B.A., Meng G., Král J., Ávila Herrera I.M., Izquierdo M.A. & Carvalho L.S. 2023a. High and dry: integrative taxonomy of the Andean spider genus Nerudia (Araneae: Pholcidae). Zoological Journal of the Linnean Society 198: 534–591. https://doi.org/10.1093/zoolinnean/zlac100

Huber B.A., Meng G., Valdez-Mondragón A., Král J., Ávila Herrera I.M. & Carvalho L.S. 2023b. Short-legged daddy-long-leg spiders in North America: the genera Pholcophora and Tolteca (Araneae, Pholcidae). European Journal of Taxonomy 880: 1–89. https://doi.org/10.5852/ejt.2023.880.2173

Huber B.A., Meng G., Král J., Ávila Herrera I.M. & Izquierdo M.A. 2023c. Revision of the South American Ninetinae genus Guaranita (Araneae, Pholcidae). European Journal of Taxonomy 900: 32–80. https://doi.org/10.5852/ejt.2023.900.2301

Huber B.A., Meng G., Cabra García J. & Carvalho L.S. 2024a. Thriving in dry conditions: on the Neotropical spider genus Galapa (Araneae: Pholcidae). Zootaxa 5419: 301–347. https://doi.org/10.11646/zootaxa.5419.3.1

Huber B.A., Meng G., Dederichs T.M., Michalik P., Forman M. & Král J. 2024b. Castaways: the Leeward Antilles endemic spider genus Papiamenta (Araneae: Pholcidae). Invertebrate Systematics 38: IS23052. https://doi.org/10.1071/IS23052

Huber B.A., Meng G., Král J., Ávila Herrera I.M. & Carvalho L.S. 2024c. Diamonds in the rough: Ibotyporanga (Araneae, Pholcidae) spiders in semi-arid Neotropical environments. European Journal of Taxonomy 963: 1–169. https://doi.org/10.5852/ejt.2024.963.2687

Huber B.A., Meng G. & Valdez-Mondragón A. 2024d. Notes on Chisosa (Araneae, Pholcidae), with the description of a new species from Mexico. Zootaxa 5419: 217–244. https://doi.org/10.11646/zootaxa.5419.2.3

Huber B.A., Szymański H. & Bennett-West A. 2024e. Progress or burden? Formal description of every apparently new species available in collections is neither necessary nor useful. ZooKeys 1214: 77–90. https://doi.org/10.3897/zookeys.1214.130592

Izquierdo M.A., Dederichs T.M., Cargnelutti F. & Michalik P. 2023. Copulatory behaviour and genital mechanics suggest sperm allocation by a non-intromittent sclerite in a pholcid spider. Royal Society Open Science 10: 230263. https://doi.org/10.1098/rsos.230263

Jackson D.A. 1993. Stopping rules in principal components analysis: A comparison of heuristical and statistical approaches. Ecology 74: 2204–2214. https://doi.org/10.2307/1939574

Jackson R.R., Brassington R.J. & Rowe R.J. 1990. Anti-predator defences of Pholcus phalangioides (Araneae, Pholcidae), a web-building and web-invading spider. Journal of Zoology 220: 543–552. https://doi.org/10.1111/j.1469-7998.1990.tb04733.x

Jackson R.R., Rowe R.J. & Campbell G.E. 1992. Anti-predator defences of Psilochorus sphaeroides and Smeringopus pallidus (Araneae, Pholcidae), tropical web-building spiders. Journal of Zoology 228: 227–232. https://doi.org/10.1111/j.1469-7998.1992.tb04604.x

Jackson R.R., Jakob E.M., Willey M.B. & Campbell G.E. 1993. Anti-predator defences of a web-building spider, Holocnemus pluchei (Araneae, Pholcidae). Journal of Zoology 229: 347–352. https://doi.org/10.1111/j.1469-7998.1993.tb02641.x

Kalyaanamoorthy S., Minh B.Q., Wong T.K.F., von Haeseler A. & Jermiin L.S. 2017. ModelFinder: fast model selection for accurate phylogenetic estimates. Nature Methods 14: 587–589. https://doi.org/10.1038/nmeth.4285

Katoh K. & Standley D.M. 2013. MAFFT multiple sequence alignment software version 7: improvements in performance and usability. Molecular Biology and Evolution 30: 772–780. https://doi.org/10.1093/molbev/mst010

Kejnovsky E., Hobza R., Cermak T., Kubat Z. & Vyskot B. 2009. The role of repetitive DNA in structure and evolution of sex chromosomes in plants. Heredity 102: 533–541. https://doi.org/10.1038/hdy.2009.17

Kimura M. 1980. A simple method for estimating evolutionary rates of base substitutions through comparative studies of nucleotide sequences. Journal of Molecular Evolution 16: 111–120. https://doi.org/10.1007/BF01731581

Kořínková T. & Král J. 2013. Karyotypes, sex chromosomes, and meiotic division in spiders. In: Nentwig W. (ed.) Spider Ecophysiology: 159–171. Springer, Berlin. https://doi.org/10.1007/978-3-642-33989-9_12

Král J. 2007. Evolution of multiple sex chromosomes in the spider genus Malthonica (Araneae: Agelenidae) indicates unique structure of the spider sex chromosome systems. Chromosome Research 15: 863–879. https://doi.org/10.1007/s10577-007-1169-3

Král J., Musilová J., Šťáhlavský F., Řezáč M., Akan Z., Edwards R.L., Coyle F.A. & Ribera Almerje C. 2006. Evolution of the karyotype and sex chromosome systems in basal clades of araneomorph spiders (Araneae: Araneomorphae). Chromosome Research 14: 859–880. https://doi.org/10.1007/s10577-006-1095-9

Král J., Kořínková T., Forman M. & Krkavcová L. 2011. Insights into the meiotic behavior and evolution of multiple sex chromosome systems in spiders. Cytogenetic and Genome Research 133 (1): 43–66. https://doi.org/10.1159/000323497

Král J., Forman M., Kořínková T., Reyes Lerma A.C., Haddad C.R., Musilová J., Řezáč M., Ávila Herrera I.M., Thakur S., Dippenaar-Schoeman A.S., Marec F., Horová L. & Bureš P. 2019. Insights into the karyotype and genome evolution of haplogyne spiders indicate a polyploid origin of lineage with holokinetic chromosomes. Scientific Reports 9: 3001. https://doi.org/10.1038/s41598-019-39034-3

Král J., Ávila Herrera I.M., Šťáhlavský F., Sadílek D., Pavelka J., Chatzaki M. & Huber B.A. 2022. Karyotype differentiation and male meiosis in European clades of the spider genus Pholcus (Araneae, Pholcidae). Comparative Cytogenetics 16 (4): 185–209. https://doi.org/10.3897/compcytogen.v16i4.85059

Kraus O. 1957. Araneenstudien 1. Pholcidae (Smeringopodinae, Ninetinae). Senckenbergiana Biologica 38: 217–243.

Levan A.K., Fredga K. & Sandberg A.A. 1964. Nomenclature for centromeric position on chromosomes. Hereditas 52: 201–220. https://doi.org/10.1111/j.1601-5223.1964.tb01953.x

Lira A.F. de A., Badillo-Montaño R., Lira-Noriega A. & de Albuquerque C.M.R. 2020. Potential distribution patterns of scorpions in north-eastern Brazil under scenarios of future climate change. Austral Ecology 45: 215–228. https://doi.org/10.1111/aec.12849

Mello-Leitão C.F. de 1918. Scytodidas e pholcidas do Brasil. Revista do Museu Paulista 10: 83–144.

Mello-Leitão C.F. de 1946. Notas sobre os Filistatidae e Pholcidae. Anais da Academia Brasileira de Ciências 18: 39–83.

Meng G., Podsiadlowski L., Dimitrov D. & Huber B.A. 2025. The complex interplay between evolutionary flexibility and diversification in a family of spiders. Systematic Biology: syaf070. https://doi.org/10.1093/sysbio/syaf070

Meng G., Carvalho L.S., Podsiadlowski L. & Huber B.A. 2026. Low coverage whole genome sequencing reveals a new subfamily of daddy long-legs spiders from Brazilian Caatinga (Araneae: Pholcidae). Arthropod Systematics and Phylogeny 84: 95–121. https://doi.org/10.3897/asp.84.e174748

Minelli A. 2019. The galaxy of the non-Linnaean nomenclature. History and Philosophy of the Life Sciences 41: 31. https://doi.org/10.1007/s40656-019-0271-0

Minh B.Q., Nguyen M.A.T. & Haeseler A. 2013. Ultrafast approximation for phylogenetic bootstrap. Molecular Biology and Evolution 30: 1188–1195. https://doi.org/10.1093/molbev/mst024

Minh B.Q., Schmidt H.A., Chernomor O., Schrempf D., Woodhams M.D., von Haeseler A. & Lanfear R. 2020. IQ-TREE 2: new models and efficient methods for phylogenetic inference in the genomic era. Molecular Biology and Evolution 37: 1530–1534. https://doi.org/10.1093/molbev/msaa015

Monteverde C., De Sales F. & Jones C. 2022. Evaluation of the CMIP6 performance in simulating precipitation in the Amazon river basin. Climate 10: 122. https://doi.org/10.3390/cli10080122

Moura M.R., do Nascimento F.A.O., Paolucci L.N., Silva D.P. & Santos B.A. 2023a. Pervasive impacts of climate change on the woodiness and ecological generalism of dry forest plant assemblages. Journal of Ecology 111: 1762–1776. https://doi.org/10.1111/1365-2745.14139

Moura M.R., Oliveira G.A., Paglia A.P., Pires M.M. & Santos B.A. 2023b. Climate change should drive mammal defaunation in tropical dry forests. Global Change Biology 29: 6931–6944. https://doi.org/10.1111/gcb.16979

Myers N., Mittermeier R.A., Mittermeier C.G., da Fonseca G.A.B. & Kent J. 2000. Biodiversity hotspots for conservation priorities. Nature 403 (6772): 853–858. https://doi.org/10.1038/35002501

Neves M.O., Broennimann O., Mod H.K., Bolochio B.E., Santana D.J., Guisan A. & Moura M.R. 2025. Climate change threatens amphibians and species representation within protected areas in tropical wetlands. Journal of Applied Ecology 62: 290–302. https://doi.org/10.1111/1365-2664.14846

Oliveira J.S., Santana D.J., Pantoja D.L., Ceron K. & Guedes T.B. 2024. Climate change in open environments: revisiting the current distribution to understand and safeguard the future of psammophilous squamates of the Diagonal of Open Formations of South America. Journal of Arid Environments 220: 105117. https://doi.org/10.1016/j.jaridenv.2023.105117

Oliveira U., Paglia A.P., Brescovit A.D., de Carvalho C.J.B., Silva D.P., Rezende D.T., Leite F.S.F., Batista J.A.N., Barbosa J.P.P.P., Stehmann J.R., Ascher J.S., de Vasconcelos M.F., de Marco P., Löwenberg-Neto P., Dias P.G., Ferro V.G. & Santos A.J. 2016. The strong influence of collection bias on biodiversity knowledge shortfalls of Brazilian terrestrial biodiversity. Diversity and Distributions 22 (12): 1232–1244. https://doi.org/10.1111/ddi.12489

Oliveira U., Brescovit A.D. & Santos A.J. 2017. Sampling effort and species richness assessment: a case study on Brazilian spiders. Biodiversity and Conservation 26 (6): 1481–1493. https://doi.org/10.1007/s10531-017-1312-1

Paula-Neto E., Cella D.M., Araujo D., Brescovit A.D. & Schneider M.C. 2017. Comparative cytogenetic analysis among filistatid spiders (Araneomorphae: Haplogynae). Journal of Arachnology 45: 123–128. https://doi.org/10.1636/M14-69.1

Pebesma E. 2018. Simple features for R: standardized support for spatial vector data. The R Journal 10: 439–446. https://doi.org/10.32614/RJ-2018-009

Poggio L., de Sousa L.M., Batjes N.H., Heuvelink G.B.M., Kempen B., Ribeiro E. & Rossiter D. 2021. SoilGrids 2.0: producing soil information for the globe with quantified spatial uncertainty. SOIL 7: 217–240. https://doi.org/10.5194/soil-7-217-2021

R Core Team 2023. R: A language and environment for statistical computing, version 4.3.0. R Foundation for Statistical Computing, Vienna. Available from https://www.R-project.org/ [accessed 22 Oct. 2025].

Reis Júnior R., Oliveira M.L. de & Borges G.R.A. 2015. RT4Bio – R Tools for Biologists. https://doi.org/10.13140/RG.2.1.4532.2966

Schartl M., Schmid M. & Nanda I. 2016. Dynamics of vertebrate sex chromosome evolution: from equal size to giants and dwarfs. Chromosoma 125: 553–571. https://doi.org/10.1007/s00412-015-0569-y

Sember A., Pappová M., Forman M., Nguyen P., Marec F., Dalíková M., Divišová K., Doležálková-Kaštánková M., Zrzavá M., Sadílek D., Hrubá B. & Král J. 2020. Patterns of sex chromosome differentiation in spiders: insights from comparative genomic hybridisation. Genes 11: 849. https://doi.org/10.3390/genes11080849

Sigovini M., Keppel E. & Tagliapietra D. 2016. Open nomenclature in the biodiversity era. Methods in Ecology and Evolution 7: 1217–1225. https://doi.org/10.1111/2041-210X.12594

Silva M.C., Rowland L., Oliveira R.S., Pennington R.T. & Moonlight P. 2024. Elevation modulates the impacts of climate change on the Brazilian Cerrado flora. Diversity and Distributions 30 (5): e13832. https://doi.org/10.1111/ddi.13832

Strassburg B.B.N., Brooks T., Feltran-Barbieri R., Iribarrem A., Crouzeilles R., Loyola R., Latawiek A.E., Oliveira Filho F.J.B., Scaramuzza C.A. de M., Scarano F.R., Soares-Filho B. & Balmford A. 2017. Moment of truth for the Cerrado hotspot. Nature Ecology & Evolution 1: 0099. https://doi.org/10.1038/s41559-017-0099

Suyama M., Torrents D. & Bork P. 2006. PAL2NAL: robust conversion of protein sequence alignments into the corresponding codon alignments. Nucleic Acids Research 34: W609–W612. https://doi.org/10.1093/nar/gkl315

Suzuki S. 1954. Cytological studies in spiders. III. Studies on the chromosomes of fifty-seven species of spiders belonging to seventeen families, with general considerations on chromosomal evolution. Journal of Science of Hiroshima University, Series B 2: 23–136.

Tamura K., Stecher G. & Kumar S. 2021. MEGA11: Molecular Evolutionary Genetics Analysis Version 11. Molecular Biology and Evolution 38: 3022–3027. https://doi.org/10.1093/molbev/msab120

Thuiller W., Georges D., Gueguen M., Engler R., Breiner F., Lafourcade B., Patin R. & Blancheteau H. 2025. biomod2: ensemble platform for species distribution modeling. Available from https://cran.r-project.org/web/packages/biomod2/index.html [accessed 22 Oct. 2025].

Torres R.R., Lapola D.M. & Gamarra N.L.R. 2017. Future climate change in the Caatinga. In: Silva J.M.C., Leal I.R. & Tabarelli M. (eds) Caatinga: 383–410. Springer, Cham. https://doi.org/10.1007/978-3-319-68339-3_15

Uetz G.E. & Unzicker J.D. 1976. Pitfall trapping in ecological studies of wandering spiders. Journal of Arachnology 3: 101–111.

Velazco S.J.E., Villalobos F., Galvão F. & De Marco P. Jr 2019. A dark scenario for Cerrado plant species: effects of future climate, land use and protected areas ineffectiveness. Diversity and Distributions 25: 660–673. https://doi.org/10.1111/ddi.12886

Velazco S.J.E., Rose M.B., de Andrade A.F.A., Minoli I. & Franklin J. 2022. flexsdm: an R package for supporting a comprehensive and flexible species distribution modelling workflow. Methods in Ecology and Evolution 13: 1661–1669. https://doi.org/10.1111/2041-210X.13874

Venables W.N. & Ripley B.D. 2002. Modern Applied Statistics with S. New York, Springer.

Werneck F.P. 2011. The diversification of eastern South American open vegetation biomes: historical biogeography and perspectives. Quaternary Science Reviews 30: 1630–1648. https://doi.org/10.1016/j.quascirev.2011.03.009

World Spider Catalog 2025. World Spider Catalog. Version 26. Natural History Museum Bern. Available from http://wsc.nmbe.ch [accessed 22 Oct. 2025]. https://doi.org/10.24436/2

Zizka A., Silvestro D., Andermann T., Azevedo J., Duarte Ritter C., Edler D., Farooq H., Herdean A., Ariza M., Scharn R., Svantesson S., Wengström N., Zizka V. & Antonelli A. 2019. CoordinateCleaner: standardized cleaning of occurrence records from biological collection databases. Methods in Ecology and Evolution 10: 744–751. https://doi.org/10.1111/2041-210X.13152

Published
2026-05-14
How to Cite
Huber, B. A., Meng, G., Král, J., Ávila Herrera, I. M., & Carvalho, L. S. (2026). Ninetine spiders in Brazilian Caatinga and Cerrado: revision of Kambiwa and description of Sertana gen. nov. (Araneae, Pholcidae), with analyses of predicted range shifts due to climate change. European Journal of Taxonomy, 1054(1), 1–126. https://doi.org/10.5852/ejt.2026.1054.3276
Issue
Vol. 1054 (2026)
Section
Monograph

Copyright (c) 2026 Bernhard A. Huber, Guanliang Meng, Jiří Král, Ivalú M. Ávila Herrera, Leonardo S. Carvalho

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