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Sonora annulata (BAIRD, 1859)

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Higher TaxaColubridae, Colubrinae, Colubroidea, Caenophidia, Alethinophidia, Serpentes, Squamata (snakes)
SubspeciesSonora annulata annulata (BAIRD 1859)
Sonora annulata klauberi (STICKEL 1941) 
Common NamesE: Colorado Desert Shovelnose Snake
klauberi: Tucson Shovelnose Snake 
SynonymLamprosoma annulatum BAIRD 1859
Chionactis occipitalis annulata — SMITH & HOLLAND 1971
Chionactis occipitalis saxatilis FUNK 1967
Chionactis occipitalis annulata — CROTHER 2000: 58
Chionactis occipitalis annulata — CROTHER et al. 2012
Chionactis annulata annulata — WOOD et al. 2014
Chionactis annulata — FELDMAN et al. 2015
Sonora (Chionactis) annulata — COX et al. 2018: 975
Chionactis annulata — LEMOS-ESPINAL et al. 2019
Sonora annulata — HERPMAPPER 2020

Sonora annulata klauberi (STICKEL 1941)
Sonora occipitalis klauberi STICKEL 1941
Chionactis occipitalis klauberi — CROTHER 2000: 58
Chionactis occipitalis klauberi — CROTHER et al. 2012
Chionactis annulata klauberi — WOOD et al. 2014
Sonora annulatus klauberi — COX et al. 2018 
DistributionUSA (SE California, Arizona, Baja California); Terra typica: “Colorado desert”, restricted to Holtville, Imperial County, California by SMITH & TAYLOR 1950.

klauberi: USA (SC Arizona: Sonoran Desert); Type locality: "Tucson, Pima County. Arizona”  
Reproduction 
TypesSyntypes: USNM 2105a and 2105b
Holotype: SDNHM = SDSNH 29647, adult male; paratype SDNHM = SDSNH 17115 [klauberi]
Syntype: USNM 2105, 2106 [Lamprosoma annulatum] 
DiagnosisDiagnosis: Sonora annulata can be distinguished from all other Sonora, excepting S. occipitalis and S. palarostris, by the presence of a flattened, spadelike rostrum. Wood et al. (2014); distinguish S. annulata from S. occipitalis by the presence of black crossbands (as opposed to brown crossbands in S. occipitalis) and the presence of a red secondary crossband. Sonora annulata can be distinguished from S. palarostris by having more than 23 crossbands (as opposed to less than 23 in S. palarostris) according to Wood et al. (2014) (Cox et al. 2018: 975). 
CommentSynonymy: mostly after Mahrdt et al. 2001. WALLACH et al. 2014: 160 listed this species as a synonym of Chionactis occipitalis.

Subspecies: Wood et al. (2008) concluded that “Neither molecular nor morphological data are concordant with the traditional C. occipitalis subspecies taxonomy. Mitochondrial sequences suggest specimens recognized as C. o. klauberi are embedded in a larger geographic clade whose range has expanded from western Arizona populations, and these data are concordant with clinal longitudinal variation in morphology.” Consequently, the subspecies of C. occipitalis should be rejected.
The subspecies are distinguished partly by ventral scale counts and number of dark bands encircling the body, but the most striking variation is in pattern and coloration of secondary bands (Stickel 1941; Klauber 1951).

Distribution: not in Sonora fide Lemos-Espinal et al. 2019 but in Sonora fide Cox et al. 2018 (map in Fig. 5). 
EtymologyThe name occipitalis is derived from the Latin occipit meaning the back of the head, in reference to "the occipital crescent blotch" (Baird 1859a). 
References
  • Baird, S.F. 1859. Reptiles of the Boundary. In: United States and Mexican Boundary Survey under the Order of Lieut. Col. W. H. Emory, Major First Cavalry, and United States Commisioner. 2, Rept., Pt.2. Department of the Interior, Washington, 35 pp. - get paper here
  • Banta, Benjamin H. 1953. Some herpetological notes from southern Nevada. Herpetologica 9: 75-76 - get paper here
  • Barts, M. 2009. Chionactis occipitalis annulata (BAIRD). Sauria 31 (4): 2 - get paper here
  • Bezy, R. L., P. C. Rosen, T. R. Van Devender, and E. F. Enderson. 2017. Southern distributional limits of the Sonoran Desert herpetofauna along the mainland coast of northwestern Mexico. Mesoamerican Herpetology 4(1): 138–167 - get paper here
  • Collins J T 1997. Standard Common and current scientific names for North American amphibians and reptiles, 4th edition. Herpetological Circular 25: 1-40
  • Cox, Christian L.; Alison R. Davis Rabosky, Iris A. Holmes, Jacobo Reyes-Velasco, Corey E. Roelke, Eric N. Smith, Oscar Flores-Villela, Jimmy A. McGuire & Jonathan A. Campbell 2018. Synopsis and taxonomic revision of three genera in the snake tribe Sonorini. Journal of Natural History 52: 945-988 - get paper here
  • Crother, B. I. (ed.) 2012. Standard Common and Current Scientific Names for North American Amphibians, Turtles, Reptiles, and Crocodilians, Seventh Edition. Herpetological Circular 39: 1-92
  • Friedrich, U. 2007. Lebendfutter für Amphibien und Reptilien - Geschichte und Grundlagen. Draco 7 (28): 4-21 [2006] [Sonderheft Lebendfutter] - get paper here
  • Funk, Richard S. 1967. A New Colubrid Snake of the Genus Chionactis from Arizona. Southwestern Naturalist 12 (2): 180 - get paper here
  • Goode, Matthew J.;Schuett, Gordon W. 1994. Male combat in the western shovelnose snake (Chionactis occipitalis). Herpetological Natural History 2 (1): 115-117
  • Hallowell,E. 1854. Description of new reptiles from California. Proc. Acad. Nat. Sci. Philad. 7 [1854]: 91-97 - get paper here
  • Klauber, Laurence M. 1951. The shovel-nosed snake, Chionactis with descriptions of two new subspecies. Transactions of the San Diego Society of Natural History 11 (9): 141-204 - get paper here
  • Kunz, K. 2006. Zerteilen von Beute bei Schlangen. Reptilia (Münster) 11 (59): 6-7 - get paper here
  • Kunz, K. 2012. Kleine Tiere, kleine Terarrien, großer Anspruch: Nano-Terraristik!. Reptilia (Münster) 17 (97): 22-31 - get paper here
  • Mahrdt, Clark R., Kent R. Beaman, Philip C. Rosen and Peter A. Holm 2001. Chionactis Cope Shovel-nosed Snakes. Catalogue of American Amphibians and Reptiles (730): 1-6 - get paper here
  • Mahrdt, Clark R., Kent R. Beaman, Philip C. Rosen and Peter A. Holm 2001. Chionactis occipitalis (Hallowell) Western Shovel-nosed Snake. Catalogue of American Amphibians and Reptiles (731): 1-12 - get paper here
  • Peralta-García A, Valdez-Villavicencio JH, Fucsko LA, Hollingsworth BD, Johnson JD, Mata-Silva V, Rocha A, DeSantis DL, Porras LW, and Wilson LD. 2023. The herpetofauna of the Baja California Peninsula and its adjacent islands, Mexico: composition, distribution, and conservation status. Amphibian & Reptile Conservation 17(1&2): 57–142
  • Schmidt, D. & Kunz, K. 2005. Ernährung von Schlangen. Natur und Tier Verlag, Münster, 159 pp. - get paper here
  • Smith, H. M.; Holland, R. L. 1971. Noteworthy snakes and lizards from Baja California. Journal of Herpetology 5 (1-2): 56-59 - get paper here
  • Stebbins,R.C. 1985. A Field Guide to Western Reptiles and Amphibians, 2nd ed. Houghton Mifflin, Boston
  • Stickel, W.H. 1943. The Mexican snakes of the genera Sonora and Chionactis with notes on the status of other colubrid genera. Proc. Biol. Soc. Washington 56: 109-128 - get paper here
  • Stickel, William H. 1938. The snakes of the genus Sonora in the United States and Lower California. Copeia 1938 (4): 182-190 - get paper here
  • Stickel, William H. 1941. The subspecies of the spade-nosed snake, Sonora occipitalis. Bull. Chicago Acad. Sci. 6: 135-140
  • Werning, Heiko 2012. Die Reptilien und Amphibien des Südwestens. Draco 13 (50): 18-60 - get paper here
  • Wood DA, Fisher RN, Vandergast AG 2014. Fuzzy Boundaries: Color and Gene Flow Patterns among Parapatric Lineages of the Western Shovel-Nosed Snake and Taxonomic Implication. PLoS One 9 (5): e97494. doi:10.1371/journal.pone.0097494 - get paper here
  • Wood, D.A.; Meik, J.M.; Holycross, A.T.; Fisher, R.N. & Vandergast, A.G. 2008. Molecular and phenotypic diversity in Chionactis occipitalis (Western Shovel-nosed Snake), with emphasis on the status of C. o. klauberi (Tucson Shovel-nosed Snake). Conservation Genetics 9:1489–1507 - get paper here
 
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