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Issue: Vol. 5711 No. 4: 21 Oct. 2025
Type: Article
Published: 2025-10-21
DOI: 10.11646/zootaxa.5711.4.3
Page range: 505-526
Abstract views: 56
PDF downloaded: 6

Ecomorphological variation in Sceloporus vandenburgianus (Squamata: Phrynosomatidae) across the sky-islands Southern California and Northern Baja California

Department of Biology; La Sierra University; 4500 Riverwalk Parkway; Riverside; California 92505; USA
Department of Biology; La Sierra University; 4500 Riverwalk Parkway; Riverside; California 92505; USA
Department of Biology; La Sierra University; 4500 Riverwalk Parkway; Riverside; California 92505; USA
Department of Biology; La Sierra University; 4500 Riverwalk Parkway; Riverside; California 92505; USA; Department of Herpetology; San Diego Natural History Museum; PO Box 121390; San Diego; California; 92112; USA
Department of Biology; La Sierra University; 4500 Riverwalk Parkway; Riverside; California 92505; USA
Reptilia Sky-islands Mountain ranges ecomorphology conservation Southern Fence Lizard

Abstract

The mountain ranges of Southern California and Northern Baja California are unique upland ecosystems composed of a series of isolated sky-islands. The habitat composition among these montane isolates varies from more woodland with a closed canopy to more rocky habitats with large granite boulders with an open canopy. Sceloporus vandenburgianus is a montane habitat generalist that ranges across all the sky-islands of Southern California and Northern Baja California and has different refugia preferences at varying localities. In this study, we developed a morphometric dataset to test the hypothesis that there are ecomorphological differences among the sky-island populations of S. vandenburgianus that correspond to refugia preferences across these sky-islands. Our data support the hypothesis of local adaptation to different habitat and demonstrate that there are statistically significant differences in head shape between populations that live in rocky habitats versus mountain tops with more forested woodland canopies.

 

References

  1. Barrows, C.W. & Fisher, M. (2014) Past, present and future distributions of a local assemblage of congeneric lizards in southern California. Biological Conservation, 180, 97–107. https://doi.org/10.1016/j.biocon.2014.09.042
  2. Barrows, C.W. (2019) Responses of a Montane Lizard Assemblage to an Extended Drought. The Southwestern Naturalist, 63 (3), 173. https://doi.org/10.1894/0038-4909-63-3-173
  3. Barrows, C.W., Sweet, L.C., Rangitsch, J., Lalumiere, K., Green, T., Heacox, S., Davis, M., Vamstad, M., Heintz, J. & Rodgers, J.E. (2020) Responding to increased aridity: Evidence for range shifts in lizards across a 50-year time span in Joshua Tree National Park. Biological Conservation, 248, 108667. https://doi.org/10.1016/j.biocon.2020.108667
  4. Brehme, C.S., Clark, D.R., Rochester, C.J. & Fisher, R.N. (2011) Wildfires Alter Rodent Community Structure Across Four Vegetation Types in Southern California, USA. Fire Ecology, 7, 81–98. https://doi.org/10.4996/fireecology.0702081
  5. Bryson, R.W., Savary, W.E., Zellmer, A.J., Bury, R.B. & McCormack, J.E. (2016) Genomic data reveal ancient microendemism in forest scorpions across the California Floristic Province. Molecular Ecology, 15, 3731–3751. https://doi.org/10.1111/mec.13707
  6. CalFire (2024) The California Department of Forestry and Fire Protection. Available from: https://www.fire.ca.gov/ (accessed 1 October 2025)
  7. Calsbeek, R., Thompson, J.N. & Richardson, J.E. (2003) Patterns of molecular evolution and diversification in a biodiversity hotspot: the California Floristic Province. Molecular Ecology, 12, 1021–1029. https://doi.org/10.1046/j.1365-294X.2003.01794.x
  8. Chan, K.O. & Grismer, L.L. (2022) GroupStruct: An R Package for Allometric Size Correction. Zootaxa, 5124, 471–482. https://doi.org/10.11646/zootaxa.5124.4.4
  9. Chan, L.M., Archie, J.W., Yoder, A.D. & Fitzgerald, L.A. (2013) Review of the systematic status of Sceloporus arenicolus Degenhardt and Jones, 1972 with an estimate of divergence time. Zootaxa, 3664 (3), 312–320. https://doi.org/10.11646/zootaxa.3664.3.2
  10. Davis, E.B., Koo, M.S., Conroy, C., Patton, J.L. & Moritz, C. (2008) The California hotspots project: identifying regions of rapid diversification of mammals. Molecular Ecology, 17, 120–138. https://doi.org/10.1111/j.1365-294X.2007.03469.x
  11. Devitt, T.J., Pereira, R., Jakkula, L., Alexandrino, J., Bardeleben, C. & Moritz, C. (2009) Isolation and characterization of 15 polymorphic microsatellites in the Plethodontid salamander Ensatina eschscholtzii. Molecular Ecology Resources, 9, 966–969. https://doi.org/10.1111/j.1755-0998.2009.02518.x
  12. Devitt, T., Devitt, S., Hollingsworth, B., McGuire, J. & Moritz, C. (2013) Montane refugia predict population genetic structure in the Large–blotched Ensatina Salamander. Molecular Ecology, 22, 1650–1665. https://doi.org/10.1111/mec.12196
  13. Dye, A.W., Gao, P., Kim, J.B., Lei, T., Riley, K.L. & Yocom, L. (2023) High-resolution wildfire simulations reveal complexity of climate change impacts on projected burn probability for Southern California. Fire Ecology, 19, 20. https://doi.org/10.1186/s42408-023-00179-2
  14. Emata, K.N. & Hedin, M. (2016) From the mountains to the coast and back again: ancient biogeography in a radiation of shortrange endemic harvestmen from California. Molecular Phylogenetics and Evolution, 98, 233–243. https://doi.org/10.1016/j.ympev.2016.02.002
  15. Feldman, C.R. & Spicer, G.S. (2006) Comparative phylogeography of woodland reptiles in California: repeated patterns of cladogenesis and population expansion. Molecular Ecology, 15, 2201–2222. https://doi.org/10.1111/j.1365-294X.2006.02930.x
  16. Freeman, B.G., Song, Y., Feeley, K.J. & Zhu, K. (2021) Montane species track rising temperatures better in the tropics than in the temperate zone. Ecology Letters, 24, 1697–1708. https://doi.org/10.1111/ele.13762
  17. García-Fernández, A., Iriondo, J.M., Escudero, A., Aguilar, J.F. & Feliner, G.N. (2013) Genetic patterns of habitat fragmentation and past climate-change effects in the Mediterranean high-mountain plant Armeria caespitosa (Plumbaginaceae). American Journal of Botany, 100, 1641–1650. https://doi.org/10.3732/ajb.1200653
  18. Goldberg, S.R. (1995) Reproduction in the California mountain kingsnake, Lampropeltis zonata (Colubridae), in southern California. Bulletin of Southern California Academy of Sciences, 3, 118–121.
  19. Gomes, V., Carretero, M. & Kaliontzopoulou, A. (2016) The relevance of morphology for habitat use and locomotion in two species of wall lizards. Acta Oecologica, 70, 87–95. https://doi.org/10.1016/j.actao.2015.12.005
  20. Grismer, L.L. (2002) Amphibians and Reptiles of Baja California, Including Its Pacific Islands and the Islands in the Sea of Cortés. University of California Press, Berkley, California, 413 pp. https://doi.org/10.1525/california/9780520224179.001.0001
  21. Grismer, L.L., Wood, P.L., Quah, E.S.H., Anuar, S., Ngadi, E.B., Izam, N.A.M. & Ahmad, N. (2018) Systematics, ecomorphology, cryptic speciation and biogeography of the lizard genus Tytthoscincus Linkem, Diesmos & Brown (Squamata: Scincidae) from the sky‐island archipelago of Peninsular Malaysia. Zoological Journal of the Linnean Society, 183 (3), 635–671. https://doi.org/10.1093/zoolinnean/zlx067
  22. Grismer, L.L., Chan, K.O., Lovich, R.E. & Grismer, J.L. (2022) A statistical reanalysis of morphological differentiation among island night lizards (Xantusia riversiana) from the California Channel Islands. Vertebrate Zoology, 72, 1–27. https://doi.org/10.3897/vz.72.e78092
  23. Grismer, L.L. & Grismer, J.L. (2017) A re-evaluation of the phylogenetic relationships of the Cyrtodactylus condorensis group (Squamata; Gekkonidae) and a suggested protocol for the characterization of rock-dwelling ecomorphology in Cyrtodactylus. Zootaxa, 4300 (4), 486–504. https://doi.org/10.11646/zootaxa.4300.4.2
  24. Grismer, L.L. & Grismer, M.S. (2024) Amphibians & Reptiles of the Anza-Borrego Desert Region: A Photographic Guide. Anza-Borrego Desert Natural History Association, Borrego Springs, California, 54 pp.
  25. Grismer, J., Scott, P., Toffelmier, E., Hinds, B., Klabacka, R., Stewart, G., White, V., Oaks, J. & Bradley Shaffer, H. (2022) Genomic data reveal local endemism in southern California rubber boas (Serpentes: Boidae, Charina) and the critical need for enhanced conservation actions. Molecular Phylogenetics and Evolution, 174, 107542. https://doi.org/10.1016/j.ympev.2022.107542
  26. Hartley, R., Clemann, N., Atkins, Z., Scheele, B.C., Lindenmayer, D.B. & Amor, M.D. (2022) Isolated on sky islands: Genetic diversity and population structure of an endangered mountain lizard. Conservation Genetics, 24, 219–233. https://doi.org/10.1007/s10592-022-01495-x
  27. Hedin, M., Starrett, J. & Hayashi, C. (2013) Crossing the uncrossable: novel trans‐valley biogeographic patterns revealed in the genetic history of low‐dispersal mygalomorph spiders (Antrodiaetidae, Antrodiaetus) from California. Molecular Ecology, 22, 508–526. https://doi.org/10.1111/mec.12130
  28. Herrel, A., Meyers, J.J. & Vanhooydonck, B. (2001) Correlations between habitat use and body shape in a phrynosomatid lizard (Urosaurus ornatus): a population-level analysis. Biological Journal of the Linnean Society, 74 (3), 305–314. https://doi.org/10.1006/bijl.2001.0579
  29. Hewitt, G.M. (2001) Speciation, hybrid zones and phylogeography - or seeing genes in space and time. Molecular Ecology, 10 (3), 537–549. https://doi.org/10.1046/j.1365-294x.2001.01202.x
  30. Holden, Z.A., Swanson, A., Luce, C.H., Jolly, W.M., Maneta, M., Oyler, J.W., Warren, D.A., Parsons, R. & Affleck, D. (2018) Decreasing fire season precipitation increased recent western US forest wildfire activity. Proceedings of the National Academy of Sciences USA, 115. https://doi.org/10.1073/pnas.1802316115
  31. Keeley, J.E. & Syphard, A.D. (2021) Large California wildfires: 2020 fires in historical context. Fire Ecology, 17, 22. https://doi.org/10.1186/s42408-021-00110-7
  32. Kitzberger, T., Falk, D.A., Westerling, A.L., Swetnam, T.W. & Rossi, S. (2017) Direct and indirect climate controls predict heterogeneous early-mid 21st century wildfire burned area across western and boreal North America. PLoS ONE, 12, e0188486. https://doi.org/10.1371/journal.pone.0188486
  33. Kuchta, S.R., Parks, D.S. & Wake, D.B. (2009) Pronounced phylogeographic structure on a small spatial scale: geomorphological evolution and lineage history in the salamander ring species Ensatina eschscholtzii in central coastal California. Molecular Phylogenetics and Evolution, 50, 240–255. https://doi.org/10.1016/j.ympev.2008.10.019
  34. Losos, J.B. (2011) n.k. In: Lizards in an evolutionary tree: Ecology and adaptive radiation of anoles. University of California Press, Berkeley, California, pp. 97–142.
  35. Jombart, T., Devillard, S. & Balloux, F. (2010) Discriminant analysis of principal components: a new method for the analysis of genetically structured populations. BMC Genetics, 11, 94. https://doi.org/10.1186/1471-2156-11-94
  36. Jombart, T. & Collins, C. (2015) A tutorial for discriminant analysis of principal components (DAPC) using adegenet 2.0.0. Available from: http://adegenet.r-forge.r-project.org/files/tutorial-dapc-pdf (accessed 1 October 2025)
  37. Lapointe, F.J. & Rissler, L.J. (2005) Congruence, consensus, and the comparative phylogeography of codistributed species in California. American Naturalist, 166, 290–299. https://doi.org/10.1086/431283
  38. Lattanzio, M.S., McCann, M.J. & Manion, M. (2020) Geographic Variation in the Ecomorphology and Thermal Ecology of a Widespread Lizard. Herpetologica, 76 (1), 31–42. https://doi.org/10.1655/Herpetologica-D-19-00032
  39. Leaché, A.D., Banbury, B.L., Linkem, C.W. & de Oca, A.N.-M. (2016) Phylogenomics of a rapid radiation: Is chromosomal evolution linked to increased diversification in North American spiny lizards (Genus Sceloporus)? BMC Evolutionary Biology, 16. https://doi.org/10.1186/s12862-016-0628-x
  40. Lleonart, J., Salat, J. & Torres, G.J. (2000) Removing allometric effects of body size in morphological analysis. Journal of Theoretical Biology, 205, 85–93. https://doi.org/10.1006/jtbi.2000.2043
  41. McCoy, M.W., Bolker, B.M., Osenberg, C.W., Miner, B.G. & Vonesh, J.R. (2006) Size correction: comparing morphological traits among populations and environments. Oecologia, 148, 547–554. https://doi.org/10.1007/s00442-006-0403-6
  42. McGurty, B.M. (1988) Natural history of the California mountain kingsnake, Lampropeltis zonata. In: Delisle, H.F., Brown, P.R., Kaufman, B. & McGurty, B.M. (Eds.), Proceedings of the Conference on California Herpetology. Vol. 4. Southwestern Herpetological Society, Van Nuys, California, pp. 73–88. https://doi.org/10.2307/1446020
  43. Mendelsohn, M.B., Brehme, C.S., Rochester, C.J., Stokes, D.C., Hathaway, S.A. & Fisher, R.N. (2008) Responses in Bird Communities to Wildland Fires in Southern California. Fire Ecology, 4, 63–82. https://doi.org/10.4996/fireecology.0402063
  44. Minnich, R.A. & Franco-Vizcaíno, E. (2005) Baja California’s enduring Mediterranean vegetation: early accounts, human impacts, and conservation status. In: Cartron, J.L.E. & Ceballos, G. (Eds.), Biodiversity, Ecosystems, and Conservation in Northern Mexico. Oxford University Press, New York, New York, pp. 370–386. https://doi.org/10.1093/oso/9780195156720.003.0019
  45. Minnich, R.A. (2007b) Southern California Conifer forests. In: Barbour, M., Keeler-Wolf, T. & Schoenherr, A.A. (Eds.), Terrestrial Vegetation of California. University of California Press, Berkeley, California, pp. 502–538. https://doi.org/10.1525/california/9780520249554.003.0018
  46. Oksanen, J., Blanchet, F.G., Friendly, M., Kindt, R., Legendre, P., McGlinn, D., Minchin, P.R., O’Hara, R.B., Simpson, G.L., Solymos, P., Henry, M., Stevens, H., Szoecs, E. & Wagner, H. (2020) R package for community ecologists: popular ordination methods, ecological null models & diversity analysis. Available from: https://github.com/vegandevs/vegan (accessed 1 October 2025)
  47. Pelegrin, N., Mesquita, D.O., Albinati, P., Caldas, F.L.S., De Queiroga Cavalcanti, L.B., Borges, T.B., Costa, Falico, D.A., Galdino, J.Y.A., Tucker, D.B. & Garda, A.A. (2017) Extreme specialization to rocky habitats in Tropidurus lizards from Brazil: Trade-offs between a fitted ecomorph and autoecology in a harsh environment. Austral Ecology, 42 (6), 677–689. https://doi.org/10.1111/aec.12486
  48. Putman, B.J., Gasca, M., Blumstein, D.T. & Pauly, G.B. (2019) Downsizing for downtown: Limb lengths, toe lengths, and scale counts decrease with urbanization in western fence lizards (Sceloporus occidentalis). Urban Ecosystems, 22, 1071–1081. https://doi.org/10.1007/s11252-019-00889-z
  49. Raxworthy, C.J., Pearson, R.G., Rabibisoa, N., Rakotondrazafy, A.M., Ramanamanjato, J.-B., Raselimanana, A.P., Wu, S., Nussbaum, R.A. & Stone, D.A. (2008) Extinction vulnerability of tropical montane endemism from warming and upslope displacement: a preliminary appraisal for the highest massif in Madagascar. Global Change Biology, 14, 1703–1720. https://doi.org/10.1111/j.1365-2486.2008.01596.x
  50. Reist, J.D. (1986) An empirical evaluation of coefficients used in residual and allometric adjustment of size covariation. Canadian Journal of Zoology, 64, 1363–1368. https://doi.org/10.1139/z86-203
  51. Rissler, L.J., Hijmans, R.J., Graham, C.H., Moritz, C. & Wake, D.B. (2006) Phylogeographic lineages and species comparisons in conservation analyses: a case study of California herpetofauna. American Naturalist, 167, 655–666. https://doi.org/10.1086/503332
  52. Román-Palacios, C. & Wiens, J.J. (2020) Recent responses to climate change reveal the drivers of species extinction and survival. Proceedings of the National Academy of Sciences USA, 117, 4211–4217. https://doi.org/10.1073/pnas.1913007117
  53. Rochester, C.J., Brehme, C.S. & Clark, D.R., Stokes, D.C., Hathaway, S.A., & Fisher, R.N. (2010) Reptile and amphibian responses to large-scale wildfires in Southern California. Journal of Herpetology, 44, 333–351. https://doi.org/10.1670/08-143.1
  54. Rurik, A.J., Wilmoth, S.C., Dayton, K.E. & Sparkman, A.M. (2022) Rock Refuges Are Strongly Associated with Increased Urban Occupancy in the Western Fence Lizard, Sceloporus occidentalis. Diversity, 14 (8), 655. https://doi.org/10.3390/d14080655
  55. Schoenherr, A.A. (1992) A Natural History of California. University of California Press, Berkeley, California, 772 pp.
  56. Skalski, J.R., Richins, S.M. & Townsend, R.L. (2018) A statistical test and sample size recommendations for comparing community composition following PCA. PLOS ONE, 13 (10), e0206003. https://doi.org/10.1371/journal.pone.0206033
  57. Stebbins, R.C. (2003) A Field Guide to Western Reptiles and Amphibians. 3rd Edition. Houghton Mifflin Company, New York, New York, 533 pp.
  58. Stein, B.A., Kutner, L.S. & Adams, J.S. (2000) Precious Heritage: The Status of Biodiversity in the United States. Oxford University Press, Oxford, 416 pp. https://doi.org/10.1093/oso/9780195125191.001.0001
  59. Stephenson, J.R. & Calcarone, G.M. (1999) Southern California mountains and foothills assessment: habitat and species conservation issues. In: General Technical Report GTR-PSW-175. Pacific Southwest Research Station, Forest Service, U.S. Department of Agriculture, Albany, California, pp. 1–402.
  60. Stockman, A.K. & Bond, J.E. (2007) Delimiting cohesion species: extreme population structure and the role of ecological interchangeability. Molecular Ecology, 16, 3374–3392. https://doi.org/10.1111/j.1365-294X.2007.03389.x
  61. Syphard, A.D., Butsic, V., Bar-Massada, A., Keeley, J.E., Tracey, J.A. & Fisher, R.N. (2016) Setting priorities for private land conservation in fire-prone landscapes: Are fire risk reduction and biodiversity conservation competing or compatible objectives? Ecology and Society, 21 (3), Art. 2. [published online] https://doi.org/10.5751/ES-08410-210302
  62. Thorpe, R.S. (1975) Quantitative handling of characters useful in snake systematics with particular reference to intraspecific variation in the Ringed Snake Natrix natrix (L.). Biological Journal of the Linnean Society, 7, 27–43. https://doi.org/10.1111/j.1095-8312.1975.tb00732.x
  63. Thorpe, R.S. (1983) A review of the numerical methods for recognizing and analyzing racial differentiation. In: Felsenstein, J. (Ed.), Numerical Taxonomy. NATO ASI Series (Series G: Ecological Sciences). Springer-Verlag, Berlin, pp. 404–423. https://doi.org/10.1007/978-3-642-69024-2_43
  64. Toshima, R. (2011) Phylogeography of the Rubber Boa, Charina bottae. Masters Thesis, California State Polytechnic University, Pomona, California. [unknown pagination]
  65. Tracey, J.A., Rochester, C.J., Hathaway, S.A., Preston, K.L., Syphard, A.D., Vandergast, A.G., Diffendorfer, J.E., Franklin, J., MacKenzie, J.B., Oberbauer, T.A., Tremor, S., Winchell, C.S., Fisher, R.N. & Zang, R. (2018) Prioritizing conserved areas threatened by wildfire and fragmentation for monitoring and management. PLoS ONE, 13, e0200203. https://doi.org/10.1371/journal.pone.0200203
  66. Turan, C. (1999) A note on the examination of morphometric differentiation among fish populations: The Truss System. Turkish Journal of Zoology, 23, 259–263.
  67. Vandergast, A.G., Bohonak, A.J., Hathaway, S.A., Boys, J. & Fisher, R.N. (2008) Are hotspots of evolutionary potential adequately protected in southern California? Biological Conservation, 141, 1648–1664. https://doi.org/10.1016/j.biocon.2008.04.009
  68. Vitt, L.J., Caldwell, J.P., Zani, P.A. & Titus, T.A. (1997) The role of habitat shift in the evolution of lizard morphology: evidence from tropical Tropidurus. Proceedings of the National Academy of Sciences of the United States of America, 94 (8), 3828–3832. https://doi.org/10.1073/pnas.94.8.3828
  69. Wiens, J.J., Camacho, A., Goldberg, A., Jezkova, T., Kaplan, M.E., Lambert, S.M., Miller, E.C., Streicher, J.W. & Walls, R.L. (2019) Climate change, extinction, and Sky Island biogeography in a montane lizard. Molecular Ecology, 28, 2610–2624. https://doi.org/10.1111/mec.15073
  70. Westerling, A.L., Gershunov, A., Brown, T.J., Cayan, D.R. & Dettinger, M.D. (2003) Climate and wildfire in the western United States. Bulletin of the American Meteorological Society, 5, 595–604. https://doi.org/10.1175/BAMS-84-5-595
  71. Westerling, A.L., Cayan, D.R., Brown, T.J., Hall, B.L. & Riddle, L.G. (2004) Climate, Santa Ana winds and autumn wildfires in southern California. Eos Transactions American Geophysical Union, 31, 289–296. https://doi.org/10.1029/2004EO310001
  72. Westerling, A.L., Hidalgo, H.G., Cayan, D.R. & Swetnam, T.W. (2006) Warming and earlier spring increase western US forest wildfire activity. Science, 313 (5789), 940–943. https://doi.org/10.1126/science.1128834
  73. Westerling, A.L. & Bryant, B.P. (2008) Climate change and wildfire in California. Climate Change, 87 (S1), 231–249. https://doi.org/10.1007/s10584-007-9363-z
  74. Westerling, A.L., Bryant, B.P., Preisler, H.K., Holmes, T.P., Hidalgo, H.G., Das, T. & Shrestha, S.R. (2011) Climate change and growth scenarios for California wildfire. Climate Change, 1, 445–463. https://doi.org/10.1007/s10584-011-0329-9
  75. Westerling, A.L. (2016) Increasing western US forest wildfire activity: Sensitivity to changes in the timing of spring. Philosophical Transactions of the Royal Society B, 371 (1696), 20150178. https://doi.org/10.1098/rstb.2015.0178
  76. Westerling, A.L. (2018) Wildfire simulations for California’s fourth climate change assessment: Projecting Changes in Extreme Wildfire Events with a Warming Climate. California’s Fourth Climate Change Assessment. Rep. CCCA4-CEC-2018-014. California Energy Commission, Sacramento, California, 57 pp. Available from: http://www.climateassessment.ca.gov/techreports/docs/20180827-Projections_CCCA4-CEC-2018-014.pdf (1 October 2025)
  77. Williams, A.P., Abatzoglou, J.T., Gershunov, A., Guzman-Morales, J., Bishop, D.A., Balch, J.K. & Lettenmaier, D.P. (2019) Observed Impacts of Anthropogenic Climate Change on Wildfire in California. Earth’s Future, 7, 892–910. https://doi.org/10.1029/2019EF001210

How to Cite

Gregory, J.J., Murdoch, M., Chau, E., Grismer, L.L. & Grismer, J.L. (2025) Ecomorphological variation in Sceloporus vandenburgianus (Squamata: Phrynosomatidae) across the sky-islands Southern California and Northern Baja California. Zootaxa, 5711 (4), 505–526. https://doi.org/10.11646/zootaxa.5711.4.3