SchleipWulf D.O’SheaMarkAnnotated checklist of the recent and extinct pythons (Serpentes, Pythonidae), with notes on nomenclature, taxonomy, and distributionZookeys4112010201066298010.3897/zookeys.66.683 Python Daudin, 1803Synonyms:

Aspidoboa Sauvage, 1884 – Hoser 2004

Helionomus Gray – Hoser 2004 (incorrect subsequent spelling, APP4)

Shireenhoserus Hoser, 2004 (junior synonym of Enygrus Wagler)

Distribution:

Head (2005) reported remains of an indeterminate python from Miocene-age strata of the Siwalik Group of Pakistan. From the known distribution of extant species, this is likely to be a species of Python.

Remarks:

Hoser (2004) split this genus into several genera, e.g., Aspidoboa Sauvage (for breitensteini, brongersmai, and curtus), Broghammerus Hoser (for reticulatus), and Shireenhoserus Hoser (for anchietae and regius). However, Hoser (2004) overlooked Enygrus Wagler, 1830 (also see McDowell 1979: 9–10, 28), which makes Shireenhoserus a subjective junior synonym of Enygrus Wagler. He further intended to resurrect Heleionomus Gray, 1842 (for sebae and natalensis) but spelt the name as “Helionomus”. This constitutes an incorrect subsequent spelling (APP4), although the name Helionomus was already listed in Gray 1841 but is considered a nomen nudum (see remarks for Heleionomus). Only molurus and bivittatus would have remained within Python. Evidence from genetic studies reveal that with the exception of reticulatus and timoriensis, which were placed into Broghammerus (see comments there) by Rawlings et al. (2008), no further splitting of the clade Python is indicated. Furthermore, the phylogenetic relationships of several species (e.g., regius and anchietae, molurus and bivittatus, and sebae and natalensis) have not been fully resolved (e.g., Douglas et al. 2010: fig. 4-6). Other groups (e.g. the curtus-group sensu lato) are currently under study.

SchleipWulf D.O’SheaMarkAnnotated checklist of the recent and extinct pythons (Serpentes, Pythonidae), with notes on nomenclature, taxonomy, and distributionZookeys4112010201066298010.3897/zookeys.66.683 Python anchietae Bocage, 1887Synonyms:

Shireenhoserus anchietae (Bocage) – Hoser 2004 (junior synonym of Enygrus Wagler).

Python anchietae Bocage – Henderson and Powell 2007

SchleipWulf D.O’SheaMarkAnnotated checklist of the recent and extinct pythons (Serpentes, Pythonidae), with notes on nomenclature, taxonomy, and distributionZookeys4112010201066298010.3897/zookeys.66.683 Python bivittatus (Kuhl, 1820)Distribution:

See Greene et al. (2007), Snow et al. (2007), Pyron et al. (2008), and Barker and Barker (2009) for notes on introduced populations in Florida, USA. For distribution in Nepal, see O’Shea (1998), for distribution in Asia see Pauwels et al. (2003), (Barker and Barker (2008, 2010). Barker and Barker (2010) considered records of the occurrence of bivittatus in the Sichuan Province deviant due to complete isolation from the natural range of bivittatus and therefore excluded the province from the range of occurrence. Records from Sumatra and Borneo are believed to be incorrectly identified (Haile 1958, Groombridge and Luxmoore 1991).

Remarks:

Jacobs et al. (2009) considered this taxon a valid species. Evidence for this placement was already provided by O’Shea (1998, 2007) and Barker and Barker (2008) who pointed out that isolated populations of bivittatus do exist within the distributional range of molurus along the southern Nepalese border and in north-east India as reported from Assam by O’Shea (2007). Jacobs et al. (2009) primarily referred to Barker and Barker (2008) when stating that the isolated populations appear to exist not only sympatrically but syntopically with molurus but maintain their own integrity by avoiding interbreeding. However, O’Shea (pers. obs.) has observed the species inhabiting different habitats. Python molurus appears to occur in dry sandy woodland whereas bivittatus prefers riverine forests and flooded grasslands. O’Shea had not observed the two species occurring sympatrically or syntopically. Jacobs et al. (2009: 12) stated that de Rooij (1917) had assumed the type locality of Kuhl’s (1820) concept of bivittatus, which was based on unverified pictures by Seba, to be in Indochina rather than in the Sundaland and that the populations occurring between China and Java may be considered Python molurus sondaica (sic) Werner 1899. Nevertheless, according to Jacobson et al (2009), Mertens (1930) fixed the type locality to Java without the designation of a neotype, which has led to nomenclatural problems. Mertens (1930) as well as (Werner (1909, 1930) and Pope (1935) assumed that Schlegel (1837) rather than Kuhl (1820) had introduced the name bivittatus. According to Jacobs et al. (2009), Mertens (1930) was aware that Schlegel’s (1837) composite concept of Python bivittatus included several python taxa, namely those from India (Python molurus) and from Africa (Python sebae), respectively.

SchleipWulf D.O’SheaMarkAnnotated checklist of the recent and extinct pythons (Serpentes, Pythonidae), with notes on nomenclature, taxonomy, and distributionZookeys4112010201066298010.3897/zookeys.66.683 Python bivittatus bivittatus (Kuhl, 1820)SchleipWulf D.O’SheaMarkAnnotated checklist of the recent and extinct pythons (Serpentes, Pythonidae), with notes on nomenclature, taxonomy, and distributionZookeys4112010201066298010.3897/zookeys.66.683 Python bivittatus progschai Jacobs et al., 2009[subspecies inquirenda, APP7]Holotype:

ZFMK 87481, subadult male from SW-Sulawesi.

Type locality:

Known only from the southwest of Sulawesi.

Remarks:

Jacobs et al. (2009) separated this subspecies from the nominate form by its generally smaller size (up to 240 cm in TL), up to 50% smaller egg size, and the smaller size of the neonates as well as by slightly different patterning and scale counts.

SchleipWulf D.O’SheaMarkAnnotated checklist of the recent and extinct pythons (Serpentes, Pythonidae), with notes on nomenclature, taxonomy, and distributionZookeys4112010201066298010.3897/zookeys.66.683 Python breitensteini Steindachner, 1880Synonyms:

Python breitensteini Steindachner – Keoghet al. 2001

Aspidoboa breitensteini (Steindachner) – Hoser 2004

Python breitensteini Steindachner – Henderson and Powell 2007

Remarks:

Elevated to specific rank by Keogh et al. (2001).

SchleipWulf D.O’SheaMarkAnnotated checklist of the recent and extinct pythons (Serpentes, Pythonidae), with notes on nomenclature, taxonomy, and distributionZookeys4112010201066298010.3897/zookeys.66.683 Python brongersmai Stull, 1938Synonyms:

Python brongersmai Stull – Keoghet al. 2001

Aspidoboa brongersmai (Stull) – Hoser 2004

Python brongersmai Stull – Henderson and Powell 2007

Remarks:

Elevated to specific rank by Keogh et al. (2001).

SchleipWulf D.O’SheaMarkAnnotated checklist of the recent and extinct pythons (Serpentes, Pythonidae), with notes on nomenclature, taxonomy, and distributionZookeys4112010201066298010.3897/zookeys.66.683 Python curtus Schlegel, 1872Synonyms:

Python curtus Schlegel – Keoghet al. 2001

Aspidoboa curtus (Schlegel) – Hoser 2004

Python curtus Schlegel – Henderson and Powell 2007

Remarks:

Elevated to specific rank by Keogh et al. (2001).

SchleipWulf D.O’SheaMarkAnnotated checklist of the recent and extinct pythons (Serpentes, Pythonidae), with notes on nomenclature, taxonomy, and distributionZookeys4112010201066298010.3897/zookeys.66.683 Python euboicus Römer, 1870[extinct species, considered nomen dubium by Rage 1984]Synonyms:

Python Euboicus Römer, 1870

Heteropython euboicus (Römer) – de Rochebrune 1880

Heteropython euboicus (Römer) – Kuhn 1939, 1963

Python euboicus Römer – Rage 1984

Holotype:

Fragment of the trunk portion of the vertebral column (25 vertebrae and ribs), left dentary. No accession number. According to Szyndlar (1991) the holotype is probably lost.

Type locality:

Kimi (Euboea, Greece), early Miocene (MN ?3).

Remarks:

See Szyndlar (1991) and Szyndlar and Rage (2003: 67–68) for further information.

SchleipWulf D.O’SheaMarkAnnotated checklist of the recent and extinct pythons (Serpentes, Pythonidae), with notes on nomenclature, taxonomy, and distributionZookeys4112010201066298010.3897/zookeys.66.683 Python europaeus Szyndlar & Rage, 2003[extinct species]Synonyms:

Python sp. – Rage 1982; Ivanov 2000, 2002

Python europaeus Szyndlar & Rage, 2003

Holotype:

MNHN, VCO 29. One trunk vertebra.

Type locality:

Vieux Collonges (=Mont Ceindre), France, early/middle Miocene (MN 4/5).

Remarks:

See Szyndlar and Rage (2003: 68–72), and Rage and Bailon (2005: 427–428) for further information.

SchleipWulf D.O’SheaMarkAnnotated checklist of the recent and extinct pythons (Serpentes, Pythonidae), with notes on nomenclature, taxonomy, and distributionZookeys4112010201066298010.3897/zookeys.66.683 Python molurus (Linnaeus, 1758)SchleipWulf D.O’SheaMarkAnnotated checklist of the recent and extinct pythons (Serpentes, Pythonidae), with notes on nomenclature, taxonomy, and distributionZookeys4112010201066298010.3897/zookeys.66.683 Python molurus molurus (Linnaeus, 1758)SchleipWulf D.O’SheaMarkAnnotated checklist of the recent and extinct pythons (Serpentes, Pythonidae), with notes on nomenclature, taxonomy, and distributionZookeys4112010201066298010.3897/zookeys.66.683 Python molurus pimbura (Deraniyagala, 1945)[subspecies inquirenda, APP7]Synonyms:

Python molurus molurus (Linnaeus) (part)

Python molurus molurusConstable 1949

Python molurus pimburaDeraniyagala 1955

Python molurus molurusStimson 1969

Distribution:

First reported from Nunavil (Thenmarachi), Jaffna Peninsula, Sri Lanka by Abyerami and Sivashanthini (2008).

Remarks:

Hoser (2004) resurrected this taxon from the synonymy of Python molurus molurus without providing reasons for this action. Deraniyagala (1945) separated the subspecies from Python molurus molurus based on lower subcaudal scale counts and the irregular shape of the lateral markings. Dorsal midbody scale rows and ventral scale counts overlap those of the nominate subspecies. Constable (1949: 124) did not follow this placement and synonymized this taxon with the nominate subspecies, which was followed by Stimson (1969). A second paper by Deraniyagala (1955: 6) provided a more detailed description of the subspecies. Therein, he stated that this taxon is also separated from the nominate form “in generally possessing three preoculars instead of two” or four as stated by Wall (1921: 47) for some Indian populations of the nominate form. There appears to be a range in preocular scale counts across India, from three in the northeast, to four in the north-center, and two in northwest (O’Shea pers. obs.) but this data, from only a few specimens, requires further verification. Contrary to his findings in 1945, Deraniyagala (1955) reports this taxon to have “more subcaudals” than the nominate form, obviously a typographic error according to the scale count data provided therein. It seems likely that subsequent workers overlooked this latter work, since neither Stimson (1969) nor McDiarmid et al. (1999) or Henderson and Powell (2007) cited it. Several subsequent workers accepted the placement to the synonymy of the nominate form, but no further studies have been conducted on the molurus-complex. However, besides the lower subcaudal scale counts and the higher number of preoculars, the pink surface of the head may also constitite a morphological difference. (Boulenger (1890, 1893) and MA Smith (1943) recorded two preoculars for Python molurus, while Wall (1921) records three preoculars for specimens from Ceylon. Since Sri Lanka is a known biodiversity hot spot with a high level of endemism, this allopatric population may represent a cryptic species. Because of the evidence provided by Deraniyagala (1955), these authors tentatively list this taxon as a valid subspecies and call for further research regarding its true status (APP7).

SchleipWulf D.O’SheaMarkAnnotated checklist of the recent and extinct pythons (Serpentes, Pythonidae), with notes on nomenclature, taxonomy, and distributionZookeys4112010201066298010.3897/zookeys.66.683 Python natalensis A Smith, 1840Synonyms:

Python natalensis A Smith – Broadley 1999

Helionomus natalensis (A Smith) – Hoser 2004 (nomen nudum, also see remarks on Python)

Python natalensis A Smith – Henderson and Powell 2007

Distribution:

Notes on the distribution of this species can be found in Alexander (2007).

Remarks:

McDiarmid et al. (1999) refer to A. Smith 1833. According to Branch and Bauer (2005), the name “Python Natalensis” already appeared in A. Smith (1833) as well as in A. Smith (1838) but without a description. The name appeared again in A. Smith (1840), but this time was accompanied by a plate. Gray (1842) also cites A. Smith (1840) as do Branch and Bauer (2005). Elevated to specific rank by Broadley (1999).

SchleipWulf D.O’SheaMarkAnnotated checklist of the recent and extinct pythons (Serpentes, Pythonidae), with notes on nomenclature, taxonomy, and distributionZookeys4112010201066298010.3897/zookeys.66.683 Python regius (Shaw, 1802)Synonyms:

Shireenhoserus regia (Shaw) – Hoser 2004 (junior synonym of Enygrus Wagler).

Python regius (Shaw) – Henderson and Powell 2007

Remarks:

For notes on the natural history and distribution of this species, see Barker and Barker (2006).

SchleipWulf D.O’SheaMarkAnnotated checklist of the recent and extinct pythons (Serpentes, Pythonidae), with notes on nomenclature, taxonomy, and distributionZookeys4112010201066298010.3897/zookeys.66.683 Python sardus (Portis, 1901)[extinct species, nomen dubium]Synonyms:

Paleopython sardusPortis 1901

Paleryx sardus (Portis) – Kuhl 1963

?Python sardus (Portis) – Rage 1984

Holotype:

Articulated palatine and anterior pterygoid fragment (not traced).

Type locality:

Monte Albu (=Alba?)(Sardinia) Italy, middle Miocene (MN 6 or 7+8).

Remarks:

Szyndlar and Rage (2003: 72–73) considered this name a nomen dubium as it is indistinguishable from other (extinct) Python.

SchleipWulf D.O’SheaMarkAnnotated checklist of the recent and extinct pythons (Serpentes, Pythonidae), with notes on nomenclature, taxonomy, and distributionZookeys4112010201066298010.3897/zookeys.66.683 Python sebae (Gmelin, 1788)Synonyms:

Helionomus sebae (Gmelin) – Hoser 2004 (nomen nudum, also see remarks on Python)

Python sebae (Gmelin) – Henderson and Powell 2007

Remarks:

Elevated to specific rank by Broadley (1999).

HoserRT (2004) A reclassification of the Pythoninae including the description of two new genera, two new species and nine new subspecies.Crocodilian – Journal of the Victorian Association of Amateur Herpetologists 4 (3): 31–37 and4 (4):21-40.HeadJJ (2005) Snakes of the Siwalik Group (Miocene of Pakistan): systematics and relationship to environmental change.Palaeontologia Electronica8(1): 8133. http://palaeo-electronica.org/2005_1/head18/head18.pdf [accessed 08.IX.2010]McDowellSB (1979) A catalogue of the snakes of New Guinea and the Solomons, with special reference to those in the Bernice P. Bishop Museum. Part III. Boinae and Acrochordoidea.Journal of Herpetology13 (1):1-92.GrayJE (1841) Synopsis of the Contents of the British Museum, 43. edition.Woodfall and Sons, London, 382 pp.RawlingsLHRaboskyDLDonnellanSCHutchinsonMN (2008) Python phylogenetics: inference from morphology and mitochondrial DNA.Biological Journal of the Linnean Society93:603-619.DouglasMEDouglasMRSchuettGWBeckDDSullivanBK (2010) Conservation phylogenetics of helodermatid lizards using multiple molecular markers and a supertree approach.Molecular Phylogenetics and Evolution55:153-167.HendersonRWPowellR (2007) The biology of boas and pythons: a retrospective look to the future. In: HendersonRWPowellR (Eds) Biology of the Boas and Pythons.Eagle Mountain Publishing, Utah, USA, 321.GreeneDUPottsJMDuquesnelJGSnowRW (2007) Python molurus bivittatus. Distribution Note.Herpetological Review38(3): 335.SnowRWKryskoKLEngeKMOberhoferLWarren-BradleyAWilkinsL (2007) Introduced populations of Boa constrictor (Boidae) and Python molurus bivittatus (Pythonidae) in southern Florida. In: HendersonRWPowellR (Eds) Biology of the Boas and Pythons.Eagle Mountain Publishing, Utah, USA, 417438.PyronRABurbrinkFTGuiherTJ (2008) Claims of potential expansion throughout the US by invasive python species are contradicted by ecological niche models.PLoS ONE13 (8): e2931. doi:10.1371/journal.pone.0002931.BarkerDGBarkerTM (2009) On Burmese Pythons in the Everglades questions posed and answered on the issues of pythons in South Florida and in captivity.The Occasional Papers of Vida Preciosa International1:2009, VPI Library, Boerne, Texas, USA, 117.O’SheaM (1998) Herpetological results of two short field excursions to the Royal Bardia region of western Nepal, including range extensions for Assamese/Indo-Chinese snake taxa. In: de SilvaA (Ed) Biology and Conservation of the Amphibians, Reptiles and their Habitats in South Asia. Proceeding of the Conference on the Biology and Conservation of the Amphibians and Reptiles of South Asia, Sri Lanka, August 1996, 306317.PauwelsOSGDavidPChimsunchartCThirakhuptK (2003) Reptiles of Phetchaburi Province, Western Thailand: a list of species, with natural history notes, and a discussion on the biogeography at the Isthmus of Kra.The Natural History Journal of Chulalongkorn University3 (1):23-53.BarkerDGBarkerTM (2008) The distribution of the Burmese Python, Python molurus bivittatus.Bulletin of the Chicago Herpetological Society43:33-38.BarkerDGBarkerTM (2010) The distribution of the Burmese Python, Python bivittatus, in China.Bulletin of the Chicago Herpetological Society45 (5):86-88.HaileNS (1958) The snakes of Borneo, with a key to the species.Sarawak Museum Journal8:743-771.GroombridgeBLuxmooreR (1991) Pythons in South-East Asia. A review of distribution, status and trade in three seleted species. World Conservation Monitoriing Centre, Cambridge, U.K.127 pp.JacobsHJAuliyaMBöhmeW (2009) Zur Taxonomie des Dunklen Tigerpythons, Python molurus bivittatus Kuhl, 1820, speziell der Population von Sulawesi.Sauria31 (3):5-16.O’SheaM (2007) Boas and Pythons of the World.New Holland, London, 160 pp.RooijN de (1917) The reptiles of the Indo-Australian archipelago. II. Ophidia.Brill, Leiden, 334 pp.KuhlH (1820) Beiträge zur Zoologie und vergleichenden Anatomie. Erste Abtheilung.Beiträge zur Zoologie, Frankfurt am Main, 151 pp.WernerF (1899) Allerlei aus dem Kriechtierleben im Käfig. II.Zoologischer Garten, Frankfurt am Main,40:12-24.MertensR (1930) Die Amphibien und Reptilien der Inseln Bali, Lombok, Sumbawa und Flores.Abhandlungen der Senckenbergischen Naturforschenden Gesellschaft, Frankfurt am Main,42 (3):115-344.WernerF (1909) Neue oder seltnere Reptilien aus dem Musée Royal d‘Histoire naturelle de Belgique in Brüssel.Zoologisches Jahrbuch, Abteilung Systematik,28:263-279.WernerF (1930) Boidenstudien im Wiener Naturhistorischen Museum. IV. Python molurus Gray und bivittatus Schlegel.Zoologischer Anzeiger, Leipzig,87:205-206.PopeCH (1935) The Reptiles of China.American Museum of Natural History, New York, 604 pp.SchlegelH (1837) Essai sur la physionomie des serpens. Partie Générale.Schonekat, Amsterdam, 251 pp.KeoghJSBarkerDGShineR (2001) Heavily exploited but poorly known: systematics and biogeography of commercially harvested pythons (Python curtus group) in Southeast Asia.Biological Journal of the Linnean Society73 (1):113-129.Rochebrune deAT (1880) Révision des ophidiens fossiles du Muséum d’Histoire naturelle.Nouvelles Archives du Muséum d’Histoire naturelle3:271-296.RageJ-C (1984) Serpentes.Handbuch der Palioherpetologie, Part 11. WellenhoferP (Ed) Gustav Fischer Verlag, Stuttgart-New York, 80 pp.SzyndlarZ (1991) A Review of Neogene and Quarternary Snakes of Central and Eastern Europe. Part 1: Scolecophidia, Boidae, Colubrinae.Estudios geologica47: 103126.SzyndlarZRageJ-C (2003) Non-erycine Booidea from the Oligocene and Miocene of Europe.Institute of Systematics and Evolution of Animals. Polish Academy of Sciences, Poland, 109 pp.RageJ-C (1982) Amphibia and Squamata. In: The lower Miocene Fauna of Al-Sarra (Eastern province, Saudi Arabia). Thomas H, Sen S, Khan M, Battail B, Ligabue G (Eds)Atlal Journal of Saudi Arabian Archaeology5 (3):109-136.IvanovM (2000) Snakes of the lower/middle Miocene transition at Vieux Collonges (Rhône, France), with comments on the colonisation of western Europe by colubroids.Geodiversitas22 (4): 559588.IvanovM (2002) The oldest known Miocene snake fauna from Central Europe: Merkur-North locality, Czech Republic.Acta Palaeontologica Polonica47 (3): 513534.RageJ-CBailonS (2005) Amphibians and squamate reptiles from the late early Miocene (MN 4) of Béon 1 (Montréal-du-Gers, Southwestern France).Geodiversitas27 (3):413-441.ConstableJD (1949) Reptiles from the Indian Peninsula in the Museum of Comparative Zoölogy.Bulletin of the Museum of Comparative Zoology, Harvard College103 (2):60-160.DeraniyagalaPEP (1955) Python molurus. In: A Colored Atlas of some vertebrates from Ceylon, Vol. 3, Serpentoid Reptilia.Colombo, Sri Lanka, Government Press, xix + 121 pp.StimsonAF (1969) Liste der rezenten Amphibien und Reptilien. Boidae (Boinae + Bolyeriinae + Loxoceminae + Pythoninae).Das Tierreich89:1-49.AbyeramiSSivashanthiniK (2008) Diversity of snakes from the Jaffna Peninsula, Sri Lanka.Pakistan Journal of Biological Sciences11 (16):1969-1978.DeraniyagalaPEP (1945) Some new races of the Python, Chrysopelea, Binocellate Cobra and Tith-Polonga inhabiting Ceylon and India.Spolia Zeylanica,24:103-113.WallF (1921) Ophidia Taprobanica or the Snakes of Ceylon.HR Cottle, Colombo, xxiii + 581 pp.McDiarmidRWCampbellJATouréTA (1999) Snake species of the world: a taxonomic and geographic reference. Volume 1. The Herpetologist’s League, Washington, 511 pp.BoulengerGA (1890) Reptilia and Batrachia: The Fauna of British India, including Ceylon and Burma.Taylor and Francis, London, U.K., 564 pp.BoulengerGA (1893) Catalogue of the snakes in the British Museum (Natural History). Vol. I. British Museum (Natural History), London, U.K., 448 pp.SmithMA (1943) The Fauna of British India, Ceylon and Burma, including the whole of the Indo-Chinese subregion. Reptilia and Amphibia. Vol. III Serpentes.Taylor and Francis, London, xii + 583 pp.AlexanderGJ (2007) Thermal biology of the southern African Python (Python natalensis): Does temperature limit its distribution? In: HendersonRWPowellR (Eds) Biology of the Boas and Pythons.Eagle Mountain Publishing, Utah, USA, 5175.International Commission on Zoological Nomenclature (1999) International code of zoological nomenclature. Fourth Edition.London: The International Trust for Zoological Nomenclature.BranchWRBauerAM (2005) The herpetological contributions of Sir Andrew Smith, with an introduction, concordance of names, and annotated bibliography.SSAR, Villanova, PA, iv + 80 pp.GrayJE (1842) Synopsis of the species of prehensile-tailed snakes, or Family Boidae.The Zoological Miscellany2: 4146.BroadleyDG (1999) The southern African Python, Python natalensisA. Smith 1840, is a valid species.African Herp News, Durban29:31-32.BarkerDGBarkerTM (2006) Pythons of the World, Vol. 2: Ball Pythons.The History, Natural History, Care and Breeding.VPI Library, Boerne, Texas, USA, 322 pp.PortisA (1901) II Paleopython sardus PORT. Nuovo pithonide del Miocene medio della Sardegna.Bollettio della Società Geologica Italiana20 (2):247-253.