Scale Microstructures of Pygopodid Lizards (Gekkota: Pygopodidae): Phylogenetic Stability and Ecological Plasticity

Tatjana Nikolayevna Dujsebayeva, Natalia Ananjeva, Aaron M. Bauer

Abstract


The skin, as the interface of the body with the outside world, is directly exposed to the impacts of the environment. We have examined the microstructure of scale surfaces and the numerical distribution and morphology of skin sensory organs (SSO) in Australian limbless lizards of the family Pygopodidae. We have shown that the hairy sensory organs, as complex morphological structures, are a stable characteristic of the scale integument of pygopodids. This feature reflects their relationship to geckos and is shared homoplastically with some iguanian families (Dactyloidae, Leiosauridae, Opluridae, Chamaeleonidae). At the same time, scale micro-ornamentation as an elementary morphological structure is more plastic and, although the basic spinulate pattern is dominant, other variants occur on the scales of the serpentine body of pygopodids. We accept the spinules of MiO and the hairs of SSO as homologous structures at the cellular level since they are both derivatives of the Oberhäutchen cell surface. We propose to characterize the hair-bearing SSO of gekkotan and iguanian lizards as Oberhäutchen hairy sensory organs (ObHSO). Domination of SP MiO and presence of ObHSO in the integument of Gekkota and several families of Iguania, and sporadic occurrence of SP MiO in autarchoglossan taxa provide justification for regarding these characters as plesiomorphic. We characterize the high abundance (iterative state) of SSO in the scales of the head of pygopodids as representing the phenomenon of «overiteration», in which the phylogenetically established condition is enhanced by functional demands on the organism.

Keywords


Delma; Pygopus; scale micro-ornamentation; sensory organs; morphology; numerical distribution; phylogeny; functional implication

Full Text:

PDF

References


Alibardi L. (1996), «Scale morphogenesis during embryonic development in the lizard Anolis lineatopus», J. Anat., 188, 713 – 725.

Alibardi L. (1999), «Epidermal differentiation in the developing scales of embryos of the Australian scincid lizard Lampropholis guichenoti», J. Morphol., 241, 139 – 152.

Alibardi L. (2020), «Adhesive pads of gecko and anoline lizards utilize corneous and cytoskeletal proteins for setae development and renewal», J. Exp. Zool. B. Mol. Dev. Evol., 334(5). DOI: 10.1002/jez.b.22976

Alibardi L. and Bonfitto A. (2019), «Morphology of setae in regenerating caudal adhesive pads of the gecko Lygodactylus capensis (Smith, 1849)», Zoology, 133, 1 – 9. DOI: 10.1016/j.zool.2019.01.003

Alibardi L. and DeNardo D. F. (2013), «Ultrastructural and immunocytochemical features of the epidermis of the lizard Heloderma suspectum indicate richness in lipids and lack of a specialized shedding complex», Acta Zool. Stockholm, 94, 35 – 43. DOI: 10.1111/j.1463-6395.2011.00528.x

Ananjeva N. B. (2019), «Current state of the problems in the phylogeny of squamate reptiles (Squamata, Reptilia)», Biol. Bull. Rev., 9(2), 119 – 128.

Ananjeva N. B. and Matveyeva-Dujsebayeva T. N. (1996), «Some evidence of Gonocephalus species complex divergence basing on skin sense organ morphology», Russ. J. Herpetol., 3(1), 82 – 88.

Ananjeva N. B. and Dujsebayeva T. N. (1997), «SEM study of skin sense organs in two Uromastyx species (Sauria: Agamidae) and Sphenodon punctatus (Rhynchocephalia: Sphenodontidae)», Russ. J. Herpetol., 4(1), 46 – 49.

Ananjeva N. B., Dilmukhamedov M. E., and Matveyeva T. N. (1991), «The skin sensory organs of some iguanian lizards», J. Herpetol., 25(2), 186 – 199.

Ananjeva N. B., Dujsebayeva T. N., and Joger U. (2001), «Morphological study of the squamate integument: more evidence for the metataxon status of Leiolepidinae», J. Herpetol., 35(3), 507 – 510.

Aota Sh. (1939 – 1940), «An histological study of the integument of a Blind Snake, Typhlops braminis (Daudin), with special reference to the sense organs and nerves», J. Sci. Hirosima Univ. Ser. B. Div. 1. Zool., VII, 193 – 209.

Arnold E. N. (2002), «History and function of scale microornamentation in lacertid lizards», J. Morphol., 252, 145 – 169. DOI: 10.1002/jmor.1096

Arnold E. N. and Poinar G. A. (2008), «100 million year old gecko with sophisticated adhesive toe pads, preserved in amber from Myanmar», Zootaxa, 1847, 62 – 68.

Arrigo M. I., De Oliveira Vilaca L. M., Fofonjka A., Srikanthan A. N., Debry A., and Milinkovitch M. C. (2019), «Phylogenetic mapping of scale nanostructure diversity in snakes», BMC Evol. Biol., 19, 91. DOI: 10.1186/s12862- 019-1411-6

Bauer A. M. (2019), «Gecko adhesion in space and time: a phylogenetic perspective on the scansorial success story», Integ. Comp. Biol., 59(1), 117 – 130. DOI: 10.1093/icb/ icz020

Bauer A. M. and Russell A. (1988), «Morphology of gekkonid cutaneous sensilla, with comments on function and phylogeny in the Carphodactylini (Reptilia: Gekkonidae)», Can. J. Zool., 66(7), 1583 – 1588.

Berthé R. A., Westhoff G., Bleckmann H., and Gorb S. N. (2009), «Surface structure and frictional properties of the skin of the Amazon tree boa Corallus hortulanus (Squamata, Boidae)», J. Comp. Physiol. A, 195, 311 – 318. DOI: 10.1007/s00359-008-0408-1

Beyerlein P. (1998), «Studies on the significance of microdermatoglyphics in viperid systematics. I. The microdermatoglyphics of desert vipers», Herpetozoa, 11(1/2), 79 – 86.

Breyer H. (1929), «Über Hautsinnesorgane und Häutung der lacertilien», Zool. Jahrb. Anat., 51, 549 – 581.

Burstein N., Larsen K. R., and Smith H. M. (1974), «A preliminary survey of dermatoglyphic variation in the lizard genus Sceloporus», J. Herpetol., 8(4), 359 – 369. DOI: 10.2307/1562907

Carroll R. L. (1977), «The origin of lizards», in: S. M. Andrews, R. S. Miles, and A. D. Walker (eds.), Problems in Vertebrate Evolution, Linnean Society Symposium. Series No. 4, pp. 359 – 396.

Cogger H. G. (2014), Reptiles and Amphibians of Australia. 7th Edition, CSIRO Publishing.

Cole C. J. and Devender T. R. (1976), «Surface structure of fossil and recent epidermal scales from North American lizards of the genus Sceloporus (Reptilia, Iguanidae)», Bull. Am. Mus. Nat. Hist., 156(4), 451 – 514.

Crowe-Riddell J. M., Williams R., Chapuis L., and Sanders K. L. (2019), «Ultrastructural evidence of a mechanosensory function of scale organs (sensilla) in sea snakes (Hydrophiinae)», R. Soc. Open Sci., 6(4), 182022. DOI: 10.1098/rsos.182022

Darwish S. T. (2012), «Comparative light and ultrastructural studies of skin in Stenodactylus petrii and Ptyodactylus guttatus (Reptilia: Gekkonidae)», Egypt. J. Exp. Biol. (Zool.), 8(1), 9 – 14.

Daza J. D. Bauer A. M., and Snively A. (2013), «Gobekko cretacicus (Reptilia: Squamata) and its bearing on the interpretation of gekkotan affinities», Zool. J. Linn. Soc., 167, 430 – 448. DOI: 10.1111/zoj.12012

DeMar Jr. D. G., Conrad J. L., Head J. J., Varricchio D. J., and Wilson G. P. (2017), «A new Late Cretaceous iguanomorph from North America and the origin of New World Pleurodonta (Squamata, Iguania)», Proc. R. Soc. B, 284, 20161902. DOI: 10.1098/rspb.2016.1902

Dhouailly D. and Maderson P. F. A. (1984), «Ultrastructural observations on the embryonic development of the integument of Lacerta muralis (Lacertilia, Reptilia)», J. Morphol., 179, 203 – 228. DOI: 10.1002/jmor.1051790302

Dogel V. A. (1954), Oligomerization of Homologous Organs as One of the Main Pathways of Animal Evolution, Izd. LGU, Leningrad [in Russian].

Dujsebayeva T. N. (1994), The Skin Sense Organs of the Iguanian and Gekkonian lizards (Morphology, Topography). PhD Thesis, Almaty [in Russian with English Abstract].

Dujsebayeva T. N. (1995), «The microanatomy of the regenerated bristled receptors in two gecko species, Cyrtopodion fedtschenkoi and Sphaerodactylus rooseveltii (Reptilia: Gekkonidae)», Russ. J. Herpetol., 2(1), 58 – 64.

Dujsebayeva T. N. (2008), «The skin development in the Pallas’ Coluber, Elaphe dione (Pallas, 1773) (Serpentes, Colubridae)», Russ. J. Herpetol., 15(1), 44 – 54.

Dujsebayeva T. N., Ananjeva N. B., and Iohanssen L. K. (2004), «Reduced state of skin sense organs in Sphenodon punctatus (Rhynchocephalia: Sphenodontidae) and its phylogenetical value», Russ. J. Herpetol., 11(2), 106 – 110.

Estes R. (1983), «The fossil record and early distribution of lizards», in: A. G. J. Rhodin and K. Miyata (eds.), Advances in Herpetology and Evolutionary Biology, Museum of Comparative Zoology, Harvard University, Cambridge, pp. 365 – 398.

Estes R., de Queiroz K., and Gauthier J. (1988), «Phylogenetic relationships within Squamata», in: R. Estes and G. Pregill (eds.), Phylogenetic Relationships of the Lizard Families, Stanford University Press, Stanford, pp. 119 – 281.

Etheridge R. and de Queiroz K. (1988), «A phylogeny of Iguanidae», in: R. Estes and G. Pregill (eds.), Phylogenetic Relationships of the Lizard Families, Stanford University Press, Stanford, pp. 283 – 367.

Evans S. E. (1993), «Jurassic lizard assemblages», Rev. Paléobiol. Suisse, 7, 55 – 65.

Frost D. and Etheridge R. (1989), «A phylogenetic analysis and taxonomy of iguanian lizards (Reptilia: Squamata)», Misc. Publ. Univ. Kansas Mus. Nat. Hist., 81, 1 – 65.

Gamble T., Greenbaum E., Jackman T. R., and Bauer A. (2015), «Into the light: diurnality has evolved multiple times in geckos», Biol. J. Linn. Soc., 115, 896 – 910.

Gans C. and Baic D. (1977), «Regional specialization of reptilian scale surfaces: relation of texture and biologic role», Science New Ser., 195(4284), 1348 – 1350.

Gauthier J., Estes R., and de Queiroz K. (1988), «A phylogenetic analysis of Lepidosauromorpha», in: R. Estes and G. Pregill (eds.), Phylogenetic Relationships of the Lizard Families, Stanford Univ. Press, Stanford, California, pp. 15 – 98.

Gauthier J. A., Kearney M., Maisano J. A., Rieppel O., and Behlke A. D. B. (2012), «Assembling the squamate tree of life: perspectives from the phenotype and the fossil record», Bull. Peabody Mus. Nat. Hist., 53(1), 3 – 308.

Harvey M. B. (1993), «Microstructure, ontogeny and evolution of scale surfaces in xenosaurid lizards», J. Morphol., 216, 161 – 177.

Harvey M. B. and Gutberlet R. L. Jr. (1995), «Microstructure, evolution and ontogeny of scale surface in cordylid and gerrhosaurid lizards», J. Morphol., 226, 121 – 139.

Hedges S. B. and Vidal N. (2009), «Lizards, snakes, and amphisbaenians (Squamata)», in: S. B. Hedges and S. Kumar (eds.), The Timetree of Life, Oxford Univ. Press, pp. 383 – 389.

Hiller U. (1971), «Form und Funktion der Hautsinnesorgane bei Gekkonidae. 1. Licht- und Rasterelektronenmikroskopische Untersuchungen», Forma Functio, 4, 240 – 253.

Hiller U. (1978), «Morphology and electrophysiological properties of cutaneous sensilla in agamid lizards», Pflüger’s Arch., 377, 189 – 191.

Irish F. J., Williams E. E., and Seling E. (1988), «Scanning electron microscopy of changes in epidermal structure occurring during the shedding cycle in squamate reptiles», J. Morphol., 197, 105 – 126.

Ivakhnenko M. F. (1988), «Permian Parareptilians of USSR», Tr. Paleontol. Inst., 223, 1 – 160 [in Russian].

Jackson M. K. (1977), «Histology and distribution of the cutaneous touch corpuscles in some leptotyphlopid and colubrid snakes (Reptilia, Serpentes)», J. Herpetol., 1(1), 7 – 15.

Jackson M. K. and Sharawy M. (1980), «Scanning electron microscopy and distribution of specialized mechanoreceptors in the Texas Rat Snake Elaphe obsoleta lindheimeri (Baird and Girard)», J. Morphol., 163, 59 – 67.

Joger U. (1984), «Die Radiation der Gattung Tarentola in Makaronesien (Reptilia: Sauria: Gekkonidae)», Cour. Forschungsinst. Senckenb., 71, 91 – 111.

Kalyabina S. A., Ananjeva N. B., and Dujsebayeva T. N. (1998), «Numerical distribution of skin sensory organs in scaled integument of Phrynocephalus guttatus (Sauria; Agamidae)», Vestnik KazNU. Biol. Ser., 4, 70 – 78 [in Russian with English summary].

Kluge A. G. (1987), «Cladistic relationships in the Gekkonoidea (Squamata, Sauria)», Misc. Pubs. Univ. Michigan Mus. Zool., 173, 1 – 54.

Koppetsch T., Böhme W., Büsse S., and Gorb S. N. (2020), «Comparative epidermal microstructure anatomy and limb and tail osteology of eyelid geckos (Squamata: Eublepharidae): Implications of ecomorphological adaptations», Zool. Anz., 287, 45 – 60. DOI: 10.1016/j.jcz.2020.05.005

Kunitzky J. (1904), «Die Entstehung und Entwicklung der Cuticularhärchen auf den Pfoten von Platydactylus mauritanicus», Tr. S.-Peterburg. Obshch. Estestvoisp. Otd. Zool. Fiziol., 34(2), 19 [in Russian with German abstract].

Landmann L. (1975), «The sense organs in the skin of the head of Squamata (Reptilia)», Isr. J. Zool., 24, 99 – 135.

Lauff R. F., Russell A. P., and Bauer A. M. (1993), «Topography of the digital cutaneous sensilla of the tokay gecko, Gekko gecko (Reptilia, Gekkonidae), and their potential role in locomotion», Can. J. Zool., 71, 2462 – 2472.

Leydig F. (1868), «Über Organe eines sechsten Sinnes», Nova Acta Acad. Carol. Dresden, 34, 1 – 108.

Macey J., Schulte J. A., Larson A., Ananjeva N. B., Wang Y., Pethiyagoda R., Rastegar-Pouyani N., and Papenfuss T. (2000), «Evaluating Trans-Tethys migration: an example using acrodont lizard phylogenetics», Syst. Biol., 49(2), 233 – 256.

Maclean S. (1980), «Ultrastructure of epidermal sensory receptors in Amphibolurus barbatus (Lacertilia: Agamidae)», Cell Tissue Res., 210, 435 – 445.

Maderson P. F. A. (1965), «The embryonic development of the squamate integument», Acta Zool., 46, 275 – 295.

Maderson P. F. A. (1985), «Some developmental problems of the reptilian integument», in: C. Gans, F. Billett, and P. F. A. Maderson (eds.), Biology of the Reptilia. Vol. 14. Development B, John Wiley and Sons, New York, pp. 525 – 598.

Maderson P. F. A., Rabinowitz T., Tandler B., and Alibardi L. (1998), «Ultrastructural contributions to an understanding of the cellular mechanisms involved in lizard skin shedding with comments on the function and evolution of a unique lepidosaurian phenomenon», J. Morphol., 236, 1 – 24. DOI: 10.1002/(SICI)1097-4687(199804)236:1<1:: AID-JMOR1>3.0.CO; 2-B

Mamkaev Ju. V. (1981), «Comparison of taxa by intragroup diversity as a method of evolutionary morphology», Recent Problems of Evolutionary Morphology of Animals. Abstrs. Int. Symp., Nauka, Moscow, pp. 58 – 59 [in Russian].

Martoja R. and Martoja-Pierson U. (1967), Initiation aux Techniques de l’Histologie Animali, Masson, Paris.

Matveyeva T. N. and Ananjeva N. B. (1995), «The distribution and number of skin sensory organs of agamid, iguanid and gekkonid lizards», J. Zool. Lond., 235(2), 253 – 268. DOI: 10.1111/j.1469-7998.1995.tb05142.x

Nikitina N. G. and Ananjeva N. B. (2005), «The skin sensory organs of lizards of Teratoscincus genus (Squamata: Sauria: Gekkonidae)», in: N. Ananjeva and O. Tsinenko (eds.), Herpetologia Petropolitana, St. Petersburg, Russia, pp. 291 – 295.

Olivier P. M., Couper P., and Amey A. (2010), «A new species of Pygopus (Pygopodidae; Gekkota; Squamata) from north-eastern Queensland», Zootaxa, 2578, 47 – 61.

Oreyas-Miranda B., Zug G. R., Garcia D. Y. E., and Achaval F. (1977), «Scale organs on the head of Leptotyphlops (Reptilia, Serpentes): a variational study», Proc. Biol. Soc. Wash., 90, 209 – 213.

Pauwels O. S. G., David P., Maderson P. F. A., Dereck W., and Kumps Ch. (2000), «Dorsal scale microsculpture of Xenopelis unicolor (Serpentes, Xenopeltidae): description and position among the ophidian microdermatoglyphic patterns», Dumerilia, 4(2), 99 – 111.

Peterson J. A. (1984a), «The microstructure of the scale surface in iguanid lizards», J. Herpetol., 18(4), 437 – 467.

Peterson J. A. (1984b), «The scale microarchitecture of Sphenodon punctatus», J. Herpetol., 18(1), 40 – 47.

Peterson J. A. and Bezy R. L. (1985), «The microstructure and evolution of scale surface in xantusiid lizards», Herpetologica, 41(3), 298 – 324.

Povel D. and van der Kooji J. (1997), «Scale sensillae of the File Snake (Serpentes: Acrochordidae) and some other aquatic and burrowing snakes», Neth. J. Zool., 47(4), 443 – 456.

Price R. M. (1982), «Dorsal snake scale microdermatoglyphics: ecological indicator or taxonomic tool?» J. Herpetol., 16(3), 294 – 306. DOI: 10.2307/1563721

Price R. and Kelly P. (1989), «Microdermatoglyphics: basal patterns and transition zones», J. Herpetol., 23(3), 244 – 261.

Pyron R. A., Burbrink F. T., and Wiens J. J. (2013), «A phylogeny and revised classification of Squamata, including 4161 species of lizards and snakes», BMC Evol. Biol., 13, 93. DOI: 10.1186/1471-2148-13-93

Remane A. (1952), Die Grundlagen des natürlichen Systems, der vergleichenden Anatomie und der Phylogenetik. Theoretische Morphologie und Systematik I, Geest & Portig, Leipzig.

Riedel J., Vucko M. J. Blomberg S. P., Robson S. K. A., and Schwarzkopf L. (2019), «Ecological associations among epidermal microstructure and scale characteristics of Australian geckos (Squamata: Carphodactylidae and Diplodactylidae)», J. Anat., 234, 853 – 874. DOI: 10.1111/joa.12969

Rieppel O. (1997), «The Lepidosauromorpha: an overview with special emphasis on the Squamata», in: N. C. Fraser and H.-D. Sues (eds.), In the Shadow of the Dinosaurs: Early Mesozoic Tetrapods, Cambridge University Press, Cambridge – New York – Port Chester – Melbourne – Sydney, pp. 23 – 37.

Rocha-Barbosa O. and Moraes e Silva R. B. (2009), «Analysis of the microstructure of Xenodontinae snake scales associated with different habitat occupation strategies», Braz. J. Biol., 69(2), 631 – 637. DOI: 10.1590/S1519- 69842009000400021

Röll B. (1995), «Epidermal fine structure of the toe tips of Sphaerodactylus cinereus (Reptilia, Gekkonidae)», J. Zool. Lond., 235, 289 – 300.

Ruibal R. (1968), «The ultrastructure of the surface of lizard scales», Copeia, 1968(4), 698 – 703.

Russell A. P. and Bauer A. M. (1987), «Caudal morphology of the knob-tailed geckos, genus Nephrurus (Reptilia, Gekkonidae) with special reference to the tail tip», Austral. J. Zool., 35, 541 – 551.

Russell A. P., Lai E. K., Powell G. L., and Higham T. E. (2014), «Density and distribution of cutaneous sensilla on tails of Leopard Geckos (Eublepharis macularius) in relation to caudal autotomy», J. Morphol., 275, 961 – 979. DOI: 10.1002/jmor.20269

Sammartano F. (1980), «Microornamentazioni delle squame del Gekkonidae italiani (Reptilia, Squamata)», Animalia, 7(1 – 3), 151 – 162.

Schleich H.-H. and Kästle W. (1979), «Hautstrukturen als Kletteranpassungen bei Chamaeleo und Cophotis (Reptilia: Sauria: Chamaeleonidae, Agamidae)», Salamandra, 15(2), 95 – 100.

Schleich H.-H. and Kästle W. (1982), «Hautstrukturen an Zehen und Schwanzen einiger Agamidae (Reptilia: Sauria: Agamidae)», Salamandra, 18(3/4), 322 – 329 [with English summary].

Schmidt W. J. (1910), «Das Integument von Voeltzkowia mira Bttgr. Ein Beitrag zur Morphologie und Histologie der Eidechsenhaut», Z. wiss. Zool., 94, 605 – 720.

Schmidt W. J. (1913), «Studien am Integument der Reptilien. IV. Uroplatus fimbriatus (Schneid) und die Gekkoniden», Zool. Jb., 36, 377 – 464.

Schmidt W. J. (1918), «Studien am Integument der Reptilien, VIII. Allgemeine Bemerkungen über die Hautsinnesorgane der Schlangen», Zool. Jb. Anat. Ontog., 40, 185 – 200.

Schmidt W. J. (1920), «Einiger über die Hautsinnesorgane der Agamiden insbesondere von Calotes, nebst Bemerkungen über diese Organe bei Gekkoniden und Iguaniden», Anat. Anz., 53, 113 – 139.

Scortecci G. (1940), «Recettori degli iguanidi e di altri sauri», Atti Soc. Ital. Sci. Nat., 79, 1 – 10.

Scortecci G. (1941), «Recettori degli agamidi», Mem. Soc. Ital. Sci. Nat., 10(3), 207 – 326.

Shea G. M. (1987), «Two new species of Delma (Lacertilia: Pygopodidae) from northeastern Queensland and a note on the status of the genus», Aclys. Proc. Linn. Soc. N.S.W., 109, 203 – 212.

Shea G. M. (1993), «Family Pygopodidae», in: C. J. Glasby, G. J. B. Ross, and P. L. Beesley (eds.), Fauna of Australia. Vol. 2A. Amphibia and Reptilia, Australian Government Publishing Service, Canberra, pp. 234 – 239.

Sherbrooke W. C. and Nagle R. B. (1996), «Phrynosoma intraepidermal receptor: a dorsal intraepidermal mechanoreceptors in horned lizards (Phrynosoma; Phrynosomatidae; Reptilia)», J. Morphol., 228, 145 – 154.

Shishkin M. A. (2019), «Search for organizational equilibrium as a driving force of evolution and its manifestations in phylogeny», in: S. V. Rozhnov (ed.), Proc. of the Paleontol. Soc. Vol. 2, PIN RAS, Moscow, pp. 164 – 174 [in Russian with English abstract].

Simőes T. R. (2018), Diapsid Phylogeny and the Origin and Early Evolution of Squamates. PhD Thesis, Department of Biological Sciences, University of Alberta.

Spinner M., Westhoff G., and Gorb S. N. (2014), «Subdigital setae of chameleon feet: Friction-enhancing microstructures for a wide range of substrate roughness», Sci. Rep., 4, 5481. DOI: 10.1038/srep05481

Spinner M., Gorb S. N., and Westhoff G. (2013), «Diversity of functional microornamentation in slithering geckos Lialis (Pygopodidae)», Proc. R. Soc. B, 280, 20132160. DOI: 1098/rspb.2013.2160

Spinner M., Bleckmann H., and Westhoff G. (2015), «Morphology and frictional properties of scales of Pseudopus apodus (Anguidae, Reptilia)», Zoology (Jena), 118(3), 171 – 175. DOI: 10.1016/j.zool.2014.11.002

Stewart G. R. and Daniel R. S. (1972), «Scales of the lizard Gekko gecko: surface structure examined with the scanning electron microscope», Copeia, 1972, 252 – 257.

Stewart G. R. and Daniel R. S. (1973), «Scanning electron microscopy of scales from different body regions of three lizard species», J. Morphol., 139, 377 – 388.

Stewart W. R. and Daniel R. S. (1975), «Microornamentation of lizard scales: some variations and taxonomic correlations», Herpetologica, 31(1), 117 – 130.

Streicher J. W. and Wiens J. J. (2016), «Phylogenomic analyses reveal novel relationships among snake families», Mol. Phylogen. Evol., 100, 160 – 169.

Tatarinov L. P. (1976), Morphological Evolution of Theriodonts and General Problems of Phylogeny, Nauka, Moscow [in Russian].

Uetz P., Freed P., and Hošek J. (2020), The Reptile Database. http://www.reptile-database.org [accessed December 17, 2020].

Underwood G. (1957), «On lizards of the family Pygopodidae.A contribution to the morphology and phylogeny of the Squamata», J. Morphol., 100, 207 – 268.

Vorobyeva E. (1992), The Problem of the Terrestrial Vertebrate Origin, Nauka, Moscow [in Russian with English summary].

Walzthöny D. and Ziswiler V. (1979), «Vergleichend-morphologische Untersuchungen an den Hautsinnesorganen der Blindschleiche, Anguis fragilis Anguidae) und der Würfelnatter, Natrix tessellata (Colubridae)», Rev. Suiss. Zool., 86(3), 705 – 712.

Williams E. E. (1988), «A new look at the Iguania», in: P. E. Vanzolini and W. R. Heyer (eds.), Proceedings of the Workshop on Neotropical Distribution Patterns, Academia Brasileira de Ciencias, Rio de Janeiro, pp. 429 – 488.

Williams E. E., Peterson J. A. (1982), «Convergent and alternative designs in the digital adhesive pads of scincid lizards», Science, 215, 1509 – 1511.

Zavarzin A. A. (1986), Works on the Theory of Parallelism and Evolutionary Dynamics of Tissues, Nauka, Leningrad [in Russian].

Zheng Y. and Wiens J. J. (2016), «Combining phylogenomic and supermatrix approaches, and a time-calibrated phylogeny for squamate reptiles (lizards and snakes) based on 52 genes and 4162 species», Mol. Phylogen. Evol., 94, 537 – 547.




DOI: https://doi.org/10.30906/1026-2296-2021-28-5-291-308

Refbacks

  • There are currently no refbacks.



You can subscribe to the print or electronic version of the journal on the site of EastView Company. If you have any questions, please write to the email sales@ivis.ru