Annotated checklist of the recent and extinct pythons (Serpentes, Pythonidae), with notes on nomenclature, taxonomy, and distribution

Wulf D Schleip; Mark O’Shea

doi:10.3897/zookeys.66.683

. 2010 Nov 4;(66):29–80. doi: 10.3897/zookeys.66.683

Annotated checklist of the recent and extinct pythons (Serpentes, Pythonidae), with notes on nomenclature, taxonomy, and distribution

Wulf D Schleip ¹, Mark O’Shea ^2,³

PMCID: PMC3088416 PMID: 21594030

Abstract Abstract

McDiarmid et al. (1999) published the first part of their planned taxonomic catalog of the snakes of the world. Since then, several new python taxa have been described in both the scientific literature and non-peer-reviewed publications. This checklist evaluates the nomenclatural status of the names and discusses the taxonomic status of the new taxa, and aims to continue the work of McDiarmid et al. (1999) for the family Pythonidae, covering the period 1999 to 2010. Numerous new taxa are listed, and where appropriate recent synonymies are included and annotations are made. A checklist and a taxonomic identification key of valid taxa are provided.

Keywords: snakes, ICZN, Antaresia, Apodora, Aspidites, Bothrochilus, Broghammerus, Leiopython, Liasis, Morelia, Python, taxa

Introduction

Pythons (family Pythonidae) represent a family of non-venomous basal snakes within the superfamily Pythonoidea Fitzinger, 1826 (sensu Vidal et al. 2007, Vidal and Hedges 2009). Although present in Europe during the Miocene, and probably since the late Eocene (Szyndlar and Rage 2003), pythons are now restricted to the warmer regions of the Old World, ranging from Africa through South and Southeast Asia, Indo-Malaysia and New Guinea, to Australia (Kluge 1993, Scanlon 2001, Rawlings and Donnellan 2003, Rawlings et al. 2008). More than two thirds of the currently recognized extant species are found in the Australo-Papuan region (Kluge 1993, Scanlon 2001, Rawlings et al. 2008), where they have the greatest level of morphological and genetic diversity (Heads 2002, Rawlings and Donnellan 2003), and a high degree of endemism (Harvey et al. 2000, Rawlings et al. 2004). Whereas two Asian species (Python molurus and Python bivittatus) range north of the Tropic of Cancer, an African and at least seven Australian species extend their ranges south of the Tropic of Capricorn. Pythons occur in a variety of habitats, from desert and savanna, to subtropical and tropical rainforest (Kluge 1993) and into seasonally flooded grasslands and paddifields. Most species are terrestrial, some are arboreal (Kluge 1993) and a few are semi-aquatic. The 40 recognized extant species range in maximum adult length from 0.61 m to 10.0 m, and include the longest extant snake species.

Taxonomic changes since 1999

McDiarmid et al. (1999) has become a standard reference for snake taxonomists. Since then python systematics has received considerable attention as new phylogenetic and geographical evidence has become available. Aside from the descriptions of new genera, species, and subspecies (Table 1), the most noteworthy action was the split of the genus Python by Rawlings et al. (2008), placing two Asian taxa, reticulatus and timoriensis, into Broghammerus.

Table 1. New and resurrected taxa from 1999 until 2010. Numbers in parentheses represent the number of taxa deemed unavailable.

Year	New genera	Resurrected genera	New species	Resurrected species	Elevated to species rank	New subspecies	Resurrected subspecies
1999	0	0	0	0	2	0	0
2000	2(1)	2	5	0	8	7	2
2001	0	0	0	0	3	0	0
2002	0	0	1	0	0	2	0
2003	0	(1)	1(2)	(1)	0	(3)	0
2004	2	4	2	1	0	9	0
2005	0	0	0	0	0	0	0
2006	0	0	0	0	0	0	0
2007	0	0	0	0	0	0	0
2008	0	0	3	0	3	0	0
2009	(2)	0	0	(2)	(1)1	(4)1	0
2010	0	0	0	0	0	0	0
Total^*	4	6	12	1	17	19	2

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* Total figures exclude those taxa considered unavailable.

One author, the amateur herpetologist Raymond T. Hoser of Victoria, Australia, has caused considerable confusion in python taxonomy over the last decade by describing numerous taxa (6 new genera and subgenera, 4 new species, and 19 new subspecies) in the non-peer-reviewed literature without providing adequate descriptions for his proposed new taxa (for discussions see Aplin 1999, 2002, Wüster et al. 2001, Williams et al. 2006, 2008, Schleip 2008). Hoser rarely included important taxonomic information or data on scale counts, numbers of specimen examined, statistics, or the results of DNA analysis. Moreover, Hoser designated several types without ever having apparently examined them. Although not mandatory, the International Code of Zoological Nomenclature (ICZN 1999), hereafter termed the “Code”, recommends that only specimens personally examined by the author should be designated as types (Recommendation 73B). Furthermore, Hoser himself (1996, 1997) considered failing to examine type specimens “sloppy taxonomy”. Another recommendation (Recommendation 73C), which states which data should be provided with the holotype, is often not followed by this author. In general, inadequate descriptions inevitably lead to problems in clearly assigning specimens to established taxa, and are expensive and time-consuming for subsequent workers who have to re-examine the type material in order to make taxonomic decisions, instead of being able to rely on adequate original descriptions.

In general, the professional herpetological community has rarely accepted Hoser's taxa (Wüster et al. 2001, Aplin 2002, Williams et al. 2006, 2008, Schleip 2008, Zaher et al. 2009) unless one of his numerous names turns out to be valid and a senior synonym based on more exacting scientific work carried out by professional researchers, as was the case with Broghammerus.

Approach and Scope

The primary objective of this taxonomic checklist is to provide an overview of the taxa in the family Pythonidae, and to establish their nomenclatural status under the provisions of the Code and their current taxonomical status based on published works and knowledge. It is, however, beyond the scope of this list to propose re-classifications or re-arrangements of genera that lack fully resolved phylogenetic relationships. Although this checklist can only be a snapshot in time, it is intended to continue the work of McDiarmid et al. (1999) for the family Pythonidae over the past decade and provide updates to the list compiled by Henderson and Powell (2007). For taxa described during the past decade type species (for generic names) or type specimens (for specific names) are provided along with their type localities. Recently designated neotypes are also provided. Where new distributional information is available, this is included with the relevant citation. However, in contrast to the work of McDiarmid et al. (1999) and Henderson and Powell (2007), this checklist also contains extinct taxa.

Taxa are hierarchically arranged by indentation, and are presented in alphabetical order at the level of genera, species, and subspecies, although, in the case of subspecies, the nominate subspecies precedes other subspecies, which then are listed in alphabetical order. Annotations are made directly below the relevant taxon, unless otherwise stated. Synonyms before the year 1999, and remarks on valid taxa, unless new data are available, can be found in McDiarmidet al. (1999). A key to the extent genera, species and subspecies recognized within the family of Pythonidae is provided in Appendix 2.

Interpretation and application of the Code

The Code rules on issues regarding nomenclatural acts and works, and aims to “provide the maximum universality and continuity in the scientific names of animals compatible with the freedom of scientists to classify animals according to taxonomic judgments” (ICZN 1999). Due to its universality, the wording of the Code leaves considerable room for interpretation. For the assessment of the nomenclatural status of published names, and for the purpose of nomenclatural stability, the Code was here strictly applied to all names. In any case of ambiguous wording, the authors have consulted the glossary of the Code as suggested in the Code’s “Explanatory Note”, and as stated in article 89. In the checklist we use the abbreviation “APP” (application):

APP1.

“Characters”: To be available a name must “be accompanied by a description or definition that states in words characters that are purported to differentiate the taxon” (ICZN 1999: Art. 13.1.1). A description in the meaning of the Code is “a statement in words of taxonomic characters of a specimen or a taxon” (ICZN 1999: glossary entry for “description”), and a definition is “a statement in words that purports to give those characters which, in combination uniquely distinguish a taxon” (ICZN 1999: glossary entry for “definition”). The glossary defines the word taxon as a “taxonomic unit, whether named or not: i.e., a population, or group of populations of organisms which are usually inferred to be phylogenetically related and which have characters in common which differentiate (…) the unit (e.g., a geographic population, a genus, a family, an order) from other such units” (ICZN 1999: glossary entry for “taxon”). This latter statement clearly excludes distribution itself as a character to differentiate taxa and that complies with article 13.1.1, since it requires characters to differentiate a “geographic population” from other such units. Many taxonomists are likely to accept a geographic population, especially an insular population, only separated from other such populations by distribution, at subspecific rank. However, the Code does not distinguish between specific and subspecific rank in its requirements (Arts. 45.1, 45.2), and therefore subspecies must also be distinguishable by characters other than by their isolated locality or distribution.

APP2.

“Generalized statements”: Generalized statements such as “separated by distribution” or “separated by analysis of DNA” or relative statements such as “usually (but not always) has” do not constitute a character in the sense of article 13.1.1 (APP1). Analysis of DNA clearly describes a method although genomic differences are of diagnostic value, and distribution itself is not a character, as it is not intrinsic to any specimen within the taxon. Therefore, these are not attributes of an organism (see glossary for character). Moreover, strictly following the glossary definition of the word description, the Code would require that a taxon must be uniquely distinguished from other taxa and generalized statements do not imply uniqueness.

APP3.

“Priority”: Article 23.3.5 requires the replacement of an unavailable name with the oldest available synonym (senior synonym).

APP4.

“Incorrect subsequent spelling”: Article 33.3 states that “any subsequent spelling of a name different from the correct original spelling, other than a mandatory change or an emendation, is an “incorrect subsequent spelling”; it is not an available name and, like an incorrect original spelling (…), it does not enter into homonymy and cannot be used as a substitute name” (ICZN 1999). For species-group names article 11.9.3.2 states that they are “deemed to have been published in combination with the correct original spelling of the generic name, even if it was actually published in combination with an emendation or incorrect spelling of the generic name” (ICZN 1999). Therefore, incorrect subsequent spellings are corrected to the original spelling.

APP5.

“Nomen dubium”(pl. nomina dubia): According to the glossary of the Code a nomen dubium is “a name of unknown or doubtful application” (ICZN 1999). This glossary definition leaves a wide scope for applying the term. A nomen dubium may be a lost type specimen or a type that lacks important diagnostic features so that a name cannot be applied to a specimen with clarity. (Melville (1980, 1984) noted that considering a name as nomen dubium is a matter of taxonomic decision and not a nomenclatural one. Moreover, Mones (1989) revealed that this term was first used for a taxon which was accompanied by an insufficient description. He states that the term “(…) denotes ignorance, incapability to interpret the facts, insufficient diagnosis, or actual poorness of the type specimen” (Mones, 1989: 232). We agree with the above mentioned views and, hence, insufficient information on the holotype (Recommendations 72E, 73A, see Introduction) that obviously was randomly chosen from an online database of a natural history museum and was not examined by the author (Recommendation 73B) along with an insufficient diagnosis or definition of taxonomic characters (see Art. 13.1.1, Recommendation 13A) may make a name be considered a nomen dubium. However, the name remains available, and a subsequent revision or re-description of the taxon may establish its validity.

APP6.

“Nomen nudum” (pl. nomina nuda):For generic names to be available, the Code requires “the fixation of a type species in the original publication” (ICZN 1999: Art. 13.3). All names must be “explicitly indicated as intentionally new” (ICZN 1999: Art. 16.1). Generic names, as well as specific and subspecific names, to which no characters were provided that comply with article 13.1.1 (see APP1, APP2), are deemed a nomen nudum, and therefore considered unavailable.

APP7.

“Species inquirenda”(pl. species inquirendae): This is “a Latin term meaning a species of doubtful identity requiring further investigation” (ICZN 1999: glossary entry).

APP8.

“Unavailable name”: A name is regarded as unavailable under the provisions of the Code, if either the requirements for publication or the requirements for availability are not met. This seems to be the case for names published by Hoser in his self-published Australasian Journal of Herpetology. Although the journal’s website states that several hard copies were placed in libraries to comply with the Code, these authors were unable to locate hard copies from any major European or North American library, or obtain such from the publisher when first issued (also see Recommendations 8B–D). An order form for hard copies (http://www.smuggled.com/AJHHCO1.htm, accessed 17 May 2009) was added to the publisher’s website on 7 May 2009. The National Library of Australia (NLA), the only library that lists this journal in their catalog, also has no hard copy (enquiry # NLAref21927, 16 April 2009) and only the PDF of the second issue (Hoser 2009) of the journal as of 17 May 2009 (NLA copy request CDC-10117150, 9.V.2009 could not be processed). Articles 8.1.2 and 8.1.3 of the Code state that to be regarded as published works they “must be obtainable, when first issued (…)”, and “must have been produced in an edition containing simultaneously obtainable copies by a method that assures numerous identical and durable copies”. Neither requirements was fulfilled. Later (20 May 2009), a colleague requesting original printed hard copies directly from the publisher only receiving single-sided, black and white versions of the online papers, printed on a domestic laser printer and bound by a large staple on the upper left hand corner (V. Wallach, pers. comm.). On his website, the publisher states “both print (first print run) and online are identical including use of color”. Therefore, the hard copy received by our colleague was apparently “printed on demand”. Article 9.7 states that “copies obtained on demand of an unpublished work [Art. 8], even if previously deposited in a library or other archive” do not constitute published work. The publisher disseminates the articles via the internet as PDFs downloadable from the journal’s website, and appears to rely on the trust of subsequent workers, that paper copies do exist (e.g., Zaher et al. 2009). However, the dissemination of PDFs over the internet does not currently constitute “published works” (Art. 9.6). Since no hard copies of the relevant second issue (Hoser 2009) were obtainable when first issued, and requested hard copies were “printed on demand”, this work must be regarded as “not published” under the provisions of the Code (Arts. 8.1.2, 8.1.3, 8.6, 9.6, 9.7) and the names therein are deemed unavailable (also see Wallach et al. 2009). The names, however, are listed for the completeness of the list but are not part of the formal synonymy.

Abbreviations for Depositories of type material

AMNH

BPBM

CAS

FMNH

MCZ

NMV

MNHN

RMNH

SAMA

UTA

WAM

ZFMK

ZMUC

Hoser (2000) resurrected the name Antaresia saxacola Wells & Wellington, 1985, but, contrary to Hoser’s (2000) claims that Antaresia stimsoni Smith, 1985 (Hoser provided an incorrect date: Smith 1995) would be a subjective junior synonym of Antaresia saxacola orientalis Smith, 1985 (see Shea and Sadlier, 1999), the date of publication for Smith (1985) preceded Wells and Wellington (1985) as stated by Wells (2009), which makes Antaresia saxacola a subjective junior synonym of Antaresia stimsoni. Nevertheless, because Wells and Wellington did not provide a description for Antaresia saxacola, the name was considered a nomen nudum by Underwood and Stimson (1990) and Shea and Sadlier (1999). We agree with these authors in considering “saxacola” a nomen nudum (APP6), and orientalis replaces it (APP3). Antaresia stimsoni orientalis was not listed by Cogger (1992), Barker and Barker (1994), Cogger (2000) and Henderson and Powell (2007), but was recognized by others (e.g., Ehmann 1992, Kend 1997, Walls 1998, Torr 2000, Wilson and Swan 2008).

Genus. Apodora

Kluge, 1993

Remarks:

Kluge (1993) established this monotypic genus for the species papuana, which until then was included in the genus Liasis Gray as Liasis papuanus. Because of the lack of clarity concerning the phylogenetic position of this taxon (Liasis, Apodora (Morelia, Python)) (see Kluge 1993: fig. 28), Kluge (1993: 53) characterized it with the term “sedis mutabilis” (of changing phylogenetic position, sensu Wiley, 1981, convention 4). However, recent researchers found some of the anatomical and morphological characters used in previous phylogenetic studies unsuitable due to misidentification (e.g., Scanlon 2001) or homoplasy (e.g., Wilcox et al. 2002, for detailed discussion see Rawlings et al. 2008). Scanlon (2001) analyzed a modified version of the dataset used by Kluge (1993) and found the genus Liasis to be paraphyletic (see Scanlon 2001: fig. 26). Later Rawlings et al. (2004) placed Apodora papuana as the sister clade to all other species of Liasis, which supports the separation of Apodora papuana as proposed by Kluge (1993), but this position was not well supported by their data. More recently, Rawlings et al. (2008) indicated a relationship between Apodora papuana and Liasis olivaceus within the monophyletic genus Liasis, both taxa forming a sister clade to the Liasis fuscus/mackloti complex, but again this position was not well supported. Hence, in anticipation of more robust data, we retain the current placement of this genus.

Apodora papuana

(Peters & Doria, 1878)

Synonyms:

Aspidites ramsayi richardjonesii Hoser, 2000

Synonyms:

Python reticulatus reticulatus (Schneider) – Auliya et al. 2002

Broghammerus reticulatus dalegibbonsi Hoser, 2004

Broghammerus reticulatus euanedwardsi Hoser, 2004

(Peters, 1876)

Synonyms:

Austroliasis timorensis (Peters) – Hoser 2000 (incorrect subsequent spelling of Australiasis Wells & Wellington and of timoriensis Peters, APP4)

Australiasis timorensis (Peters) – Hoser 2004 (incorrect subsequent spelling of timoriensis, APP4)

Python timorensis (Peters) – Henderson and Powell 2007 (incorrect subsequent spelling of timoriensis, APP4)

Broghammerus timoriensis (Peters)– Rawlings et al. 2008

Remarks:

Doubts were casted in literature that this species occurs on Timor (e.g., Barker and Barker 1996, McDiarmid et al. 1999, O’Shea 2007) because no records other than the type specimen are known from Timor, and this reported occurrence is likely incorrect.

Genus. Chondropython

Meyer, 1874 [synonym of Morelia]

Remarks:

Hoser (2000) resurrected this genus for viridis, and later added two new subspecies viridis shireenae Hoser 2004 and viridis adelynhoserae Hoser 2009 (APP8). Rawlings et al. (2008) demonstrated that two lineages (“northern” and “southern” lineage) along with the taxon Morelia carinata, form asubclade within the clade that represented Morelia. Although Chondropython is the oldest available name for this subclade, we see no advantage in resurrecting taxa as subgenera in a low-diversity genus.

Chondropython azureus

(Meyer, 1874)

Synonyms:

Chondropython viridis (Schlegel, 1872) – Hoser 2000 (part)

Main Quarry, Allingham Formation (early Pliocene), Bluff Downs Station, northeastern Queensland.

Remarks:

Remarks:

We accept the subspecies proposed by Smith (1981) and supported by molecular genetic evidence from Rawlings et al. (2004).

Liasis olivaceus olivaceus

Gray, 1842

Liasis olivaceus barroni

LA Smith, 1981

Genus. Montypythonoides

Smith & Plane, 1985 [subjective junior synonym of Morelia]

Type species:

Montypythonoides riversleighensis Smith & Plane, 1985 [extinct species]

Remarks:

Smith and Plane (1985: 194) stated that this genus “…shows strong relationship with species of Morelia”. Also see Morelia riversleighensis.

Genus. Morelia

Gray, 1842

Synonyms:

Lenhoserus Hoser, 2000

Chondropython Meyer, 1874 – Hoser 2000

Nyctophilopython Wells & Wellington, 1985 – Hoser 2000

Montypythonoides Smith & Plane, 1985 – Scanlon 2001

Australiasis Wells & Wellington, 1984 – Hoser 2004

Remarks:

Hoser (2000) proposed the splitting of this genus into several genera. He created a new genus, Lenhoserus Hoser (see comments there) (for Morelia boeleni), and resurrected Australiasis Wells & Wellington (for Morelia amethistina and Broghammerus timoriensis), but created an unavailable name (APP6) (“Austroliasis”) by incorrect subsequent spelling (APP4). Later, Hoser (2004) used the correct spelling Australiasis Wells & Wellington, added the species recognized by Harvey et al. (2000) and additionally resurrected the taxon duceboracensis Günther 1879 (see comments there). Furthermore, he resurrected Chondropython Meyer 1874 (for Morelia viridis) and Nyctophilopython Wells & Wellington (for Morelia oenpelliensis). However, phylogenetic studies (Rawlings et al. 2008) revealed that this taxonomic action is unwarranted. Although Rawlings et al.’s (2008) maximum parsimony analysis showed Morelia to be diphyletic (but monophyletic in Bayesian analysis), the separation of the amethistina-complex (sensu Harvey et al. 2000) and of oenpelliensis from the bredli/spilota-clade would in any case be unwarranted and would nullify the monophyly of this grouping. The resurrection of Chondropython would only be warranted at subgeneric rank with the inclusion of the two recognized lineages of the green tree python (sensu Rawlings and Donnellan 2003) and of Morelia carinata. However, Rawlings and Donnellan (2003) and Rawlings et al. (2008) avoided such placement because the phylogeny was not fully resolved (see comments for Chondropython azureus) The placement of Morelia boeleni as a separate monotypic genus is also unwarranted. We do not see any value in dividing such a small genus, and in the interests of nomenclatural stability, we place Australiasis, Lenhoserus, Chondropython, and Nyctophilopython in the synonymy of Morelia.

Morelia amethistina

(Schneider, 1801)

Synonyms:

Austroliasis amethistina (Schneider) – Hoser 2000 (incorrect subsequent spelling, APP4)

Australiasis amethistina (Schneider) – Hoser 2004

Australiasis amethystina clarki (Barbour, 1914) – Hoser 2004

Australiasis duceboracensis (Günther, 1879) – Hoser 2004

Australiasis amethistina (Scheider) – Hoser 2009 (APP8, see introduction)

(Brongersma, 1953)

Synonyms:

Lenhoserus boeleni (Brongersma) – Hoser 2000

Morelia boeleni (Brongersma) – Henderson and Powell 2007; Flagle and Stoops 2009

Remarks:

Austin et al. (2009) found little genetic divergence within specimens across the species’ range. A single specimen out of 98 examined using the cytochrome b gene, from the eastern Morobe Province, PNG showed about 1.1% genetic divergence from specimens from West Papua. This demonstrates reduced genetic diversity within this taxon.

Morelia bredli

(Gow, 1981)

Remarks:

Fyfe (2007) lists this species as subspecies Morelia spilota bredli.

Morelia carinata

(Smith, 1981)

Morelia clastolepis

Harvey et al., 2000

Synonyms:

Synonyms:

Australiasis tracyae (Harvey et al.) – Hoser 2004, 2009 (APP8, see introduction)

Morelia tracyae Harvey et al. – Henderson and Powell 2007

Holotype:

UTA 44473.

Type locality:

Tobelo, Halmahera, Maluku (=Moluccas), Indonesia.

Morelia viridis

(Schlegel, 1872)

Synonyms:

Chondropython viridis (Schlegel) – Hoser 2000 (part)

Chondropython viridis viridis (Schlegel) – Hoser 2004 (part)

Chondropython viridis shireenae Hoser, 2004

Morelia viridis (Schlegel) – Henderson and Powell 2007

Distribution:

see Wilson and Heinsohn (2007)

Remarks:

Rawlings and Donnellan (2003) found molecular evidence for cryptic diversity within Morelia viridis, resulting in two genetically distinct races. The type locality for Morelia viridis is Aru Island, which applies to the “southern lineage”, including Australian specimens (Rawlings and Donnellan 2003) (also see Chondropython azureus). However, Rawlings and Donnellan (2003: 42) noted that “(…) the east/west limits of the distribution of the two lineages may not necessarily be at the extreme ends of the central cordillera or the island”, and hence, there may be even more lineages present. Finally, due to the absence of molecular genetic data from the holotype, the type locality Aru Island cannot definitely be confirmed.

Genus. Python

Daudin, 1803

Synonyms:

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.

Python anchietae

Bocage, 1887

Synonyms:

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

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

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.

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.

Python molurus

(Linnaeus, 1758)

Python molurus molurus

(Linnaeus, 1758)

Python molurus pimbura

(Deraniyagala, 1945) [subspecies inquirenda, APP7]

Synonyms:

Python molurus molurus (Linnaeus) (part)

Python molurus molurus – Constable 1949

Python molurus pimbura – Deraniyagala 1955

Python molurus molurus – Stimson 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).

Python natalensis

A Smith, 1840

Synonyms:

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).

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).

Python sardus

(Portis, 1901) [extinct species, nomen dubium]

Synonyms:

Paleopython sardus – Portis 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.

Python sebae

(Gmelin, 1788)

Synonyms:

Supplementary Material

XML Treatment for Antaresia

zookeys.66.683-treatment1.xml^{(25.5KB, xml)}

XML Treatment for Apodora

zookeys.66.683-treatment10.xml^{(10.9KB, xml)}

XML Treatment for Aspidites

zookeys.66.683-treatment12.xml^{(24.6KB, xml)}

XML Treatment for Aspidoboa

zookeys.66.683-treatment20.xml^{(4.4KB, xml)}

XML Treatment for Australiasis

zookeys.66.683-treatment21.xml^{(11.8KB, xml)}

XML Treatment for Austroliasis

zookeys.66.683-treatment25.xml^{(2.8KB, xml)}

XML Treatment for Bothrochilus

zookeys.66.683-treatment26.xml^{(3.4KB, xml)}

XML Treatment for Broghammerus

zookeys.66.683-treatment28.xml^{(33.5KB, xml)}

XML Treatment for Chondropython

zookeys.66.683-treatment40.xml^{(16.3KB, xml)}

XML Treatment for Heleionomus

zookeys.66.683-treatment45.xml^{(3.9KB, xml)}

XML Treatment for Helionomus

zookeys.66.683-treatment46.xml^{(3KB, xml)}

XML Treatment for Jackypython

zookeys.66.683-treatment47.xml^{(2.3KB, xml)}

XML Treatment for Katrinus

zookeys.66.683-treatment48.xml^{(13.5KB, xml)}

XML Treatment for Leiopython

zookeys.66.683-treatment52.xml^{(20.4KB, xml)}

XML Treatment for Lenhoserus

zookeys.66.683-treatment60.xml^{(3.9KB, xml)}

XML Treatment for Liasis

zookeys.66.683-treatment61.xml^{(27.4KB, xml)}

XML Treatment for Montypythonoides

zookeys.66.683-treatment71.xml^{(2.4KB, xml)}

XML Treatment for Morelia

zookeys.66.683-treatment72.xml^{(69.7KB, xml)}

XML Treatment for Python

zookeys.66.683-treatment97.xml^{(68.4KB, xml)}

XML Treatment for Rawlingspython

zookeys.66.683-treatment114.xml^{(2.9KB, xml)}

XML Treatment for Shireenhoserus

zookeys.66.683-treatment115.xml^{(3.1KB, xml)}

Acknowledgements

The authors are grateful to P. Doughty (WAM) for tracking down a type specimen accession number. We would like to thank L. MacKenzie of the Australian National Library and V. Wallach (MCZ) for providing important information on the status of the Australasian Journal of Herpetology, D.G. Barker (VPI), R.W. Henderson (Milwaukee Public Museum), G. Shea (University of Sydney) and two anonymous reviewers for valuable advice on the manuscript, and A.F. Stimson (BMNH, now retired) for reflecting on the status of Python molurus pimbura.

Appendix 1

A list of valid taxa of pythons recognized in this study. Doubtful names (nomina dubia) are not included.

Antaresia Wells & Wellington, 1984

Antaresia childreni (Gray, 1842)

Antaresia maculosa (Peters, 1873)

Antaresia perthensis (Stull, 1932)

Antaresia stimsoni(LA Smith, 1985)

Antaresia stimsoni stimsoni (LA Smith, 1985)

Antaresia stimsoni orientalis (LA Smith, 1985)

Apodora Kluge, 1993

Apodora papuana(Peters & Doria, 1878)

Aspidites Peters, 1877

Aspidites melanocephalus (Krefft, 1864)

Aspidites ramsayi (Macleay, 1882)

Bothrochilus Fitzinger, 1843

Bothrochilus boa Fitzinger, 1843

Broghammerus Hoser, 2004 fide Rawlings et al. 2008

Broghammerus reticulatus(Schneider, 1801)

Broghammerus reticulatus reticulatus (Schneider, 1801)

Broghammerus reticulatus jampeanus (Auliya et al., 2002)

Broghammerus reticulatus saputrai (Auliya et al., 2002)

Broghammerus timoriensis(Peters, 1876)

Leiopython Hubrecht, 1879

Leiopython albertisii (Peters & Doria, 1878)

Leiopython bennettorum Hoser, 2000

Leiopython biakensis Schleip, 2008

Leiopython fredparkeri Schleip, 2008

Leiopython hoserae Hoser, 2000

Leiopython huonensis Schleip, 2008

Liasis Gray, 1842

Liasis dubudingala Scanlon and Mackness, 2002 [extinct species]

Liasis fuscus Peters, 1873

Liasis mackloti (Duméril and Bibron, 1844)

Liasis mackloti mackloti (Duméril and Bibron, 1844)

Liasis mackloti dunni Stull, 1932

Liasis mackloti savuensis (Brongersma, 1956)

Liasis olivaceus Gray, 1842

Liasis olivaceus olivaceus Gray, 1842

Liasis olivaceus barroni LA Smith, 1981

Morelia Gray, 1842

Morelia azurea (Meyer, 1874)

Morelia amethistina (Schneider, 1801)

Morelia boeleni (Brongersma, 1953)

Morelia bredli (Gow, 1981)

Morelia carinata (LA Smith, 1981)

Morelia clastolepis Harvey et al., 2000

Morelia kinghorniStull, 1933

Morelia nauta Harvey et al., 2000

Morelia oenpelliensis(Gow, 1977)

Morelia riversleighensis (Smith and Plane, 1985) [extinct species]

Morelia spilota(Lacépède, 1804)

Morelia spilota spilota (Lacépède, 1804)

Morelia spilota cheynei Wells & Wellington, 1984

Morelia spilota harrisoni Hoser, 2000

Morelia spilota imbricata LASmith, 1981

Morelia spilota mcdowelli Wells & Wellington, 1984

Morelia spilota metcalfei Wells & Wellington, 1984

Morelia spilota variegata Gray, 1842

Morelia tracyae Harvey et al., 2000

Morelia viridis(Schlegel, 1872)

Python Daudin, 1803

Python anchietae Bocage, 1887

Python bivittatus(Kuhl, 1820)

Python bivittatus bivittatus (Kuhl, 1820)

Python bivittatus progschai Jacobs et al., 2009

Python breitensteini Steindachner, 1880

Python brongersmai Stull, 1938

Python curtus Schlegel, 1872

Python europaeus Szyndlar & Rage, 2003 [extinct species]

Python molurus (Linnaeus, 1758)

Python molurus molurus (Linnaeus, 1758)

Python molurus pimbura Deraniyagala, 1945

Python natalensis A Smith, 1840

Python regius (Shaw, 1802)

Python sebae (Gmelin, 1788)

Appendix 2

A. Key to the genera of the Pythonidae

1.	Visible sensory pits absent	Aspidites
–	Visible sensory pits present	2
2.	Rostral unpitted	3
–	Rostral pitted	6
3.	No visible black pigmentation between the scales	4
–	Black pigmentation visible between the scales	Apodora
4.	Number of loreals fewer than 3	5
–	Number of loreals more than 3	Antaresia
5.	Head color not black, head distinct from neck, two pairs of prefrontals	Liasis
–	Head color black, head bearly distinct from neck, one pair of prefrontals	Bothrochilus
6.	Body unpatterned	Leiopython
–	Body patterned	7
7.	Lateroposterior margin of nasal exposed, plane of ventral position of postorbital is directed anterolaterally, neck is markedly narrower than the head in adults	Morelia
–	Lateroposterior margin of nasal is covered by prefrontal, plane ventral position of postorbital is directed anteriorly, neck is slightly narrower than the head in adults	8
8.	Well defined square or triangular supralabial pits, infralabials less well developed and not set in a groove	Python
–	Less well defined diagonal supralabial pits, infralabials placed in a longitudinal groove and ventrally in a fold	Broghammerus

Open in a new tab

B. Key to the species and subspecies of the genera of Pythonidae

Antaresia

1.	Body color pale yellowish-brown to dark purplish-brown	2
–	Head and neck color yellowish to reddish-brown	3
2.	Midbody scale rows 35 or fewer, ventrals fewer than 250, 34–45 subcaudals	Antaresia perthensis
–	Midbody scale rows 35 or more, ventrals more than 250, 38–57 subcaudals	Antaresia childreni
3.	Dorsal pattern of ragged-edged dark blotches	Antaresia maculosa
–	Dorsal pattern of smooth-edged blotches	4
4.	Ventrals 260–302	Antaresia stimsoni stimsoni
–	Ventrals 243–284	Antaresia stimsoni orientalis

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Aspidites

1.	Head and neck color glossy black, numerous dark brown crossbands, ventrals more than 310	Aspiditesmelanocephalus
–	Head and neck color yellowish to reddish-brown, black markings above the eyes, ventrals fewer than 305	Aspidites ramsayi

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Apodora

Black skin pigmentation visible between head scales, rostral and (at least) second supralabial with shallow pits, prefrontals small or absent, ventrals 358–374, 83–88 subdaudals, low number of teeth on the maxilla

Apodora papuanus

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Bothrochilus

Uniform brownish-black head barely distinct from the head, orange color body ring pattern that fades with age, lack of rostral and supralabial pits, low number of dentary teeth

Bothrochilus boa

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Broghammerus

1.	Iris color olive-golden, midbody scale rows fewer than 64, ventrals fewer than 290	Broghammerus timoriensis
–	Iris color bright yellow to golden-orange, midbody scale rows 64 or more, ventrals more than 290	2
2.	Ventrals more than 330	Broghammerus reticulatus saputrai
–	Ventrals fewer than 330	3
3.	Ventrals fewer than 304	Broghammerus reticulatus jampeanus
–	Ventral more than 304	Broghammerus reticulatus reticulatus

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Leiopython

1.	Dorsal color dark gray or blackish-blue fading to white on the flanks	Leiopython hoserae
–	Dorsal color yellow to brownish-violet fading to yellowish on the flanks	2
2.	One pair of enlarged parietals	Leiopython huonensis
–	Two pairs of enlarged parietals	3
3.	Whitish postocular spot absent	Leiopython fredparkeri
–	Whitish postocular spot present	4
4.	Two prefrontals, two or more loreals present	Leiopython bennettorum
–	One prefrontal, one loreal present	5
5.	Subocular absent, three labials enter the orbit	Leiopython albertisii
–	Subocular present, only two labials enter the orbit	Leiopython biakensis

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Liasis

1.	Body unpatterned	2
–	Body patterned	4
2.	Midbody scale rows fewer than 50, ventrals fewer than 300	Liasis fuscus
–	Midbody scale rows more than 50, ventrals more than 300	3
3.	Midbody scale rows 61–72, 355–377 ventrals	Liasis olivaceus olivaceus
–	Midbody scale rows 58–63, 374–411 ventrals	Liasis olivaceus barroni
4.	Eyes pale or white	Liasis mackloti savuensis
–	Eyes silvery or dark	5
5.	Chin and infralabials yellowish of color, brownish ground color, females larger than male	Liasis mackloti mackloti
–	Chin and infralabials of white color, grayish to reddish-brown ground color, males larger than females	Liasis mackloti dunni

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Morelia

1.	Dorsal scales rough or keeled, large round frontal scale	Morelia carinata
–	Dorsal scales smooth, frontal of different shape partly fragmented	2
2.	Two or more enlarged well-defined pairs of parietals	11
–	Small granular or fragmented head scales	3
3.	Body ground color shiny green with unpatternd head	Morelia azurea, Morelia viridis
	(Note: Morelia azurea is a cryptic species, only distinguishable by genetic markers)
–	Body ground color pale cream; red or brown with head pattern	4
4.	Loreal scales fewer than 28	5
–	Loreal scales more than 28	Morelia bredli
5.	Body pattern of speckled appearance	Morelia spilota spilota
–	Body pattern of pale and dark elements	6
6.	Lack of partial structure in the posterior margin of the nasal scale	7
–	Presence of partial structure in the posterior margin of the nasal scale	8
7.	Nostril not in contact with the internasals	Morelia spilota imbricata
–	Nostril in contact with the internasals	Morelia spilota metcalfei
8.	Dorsal color dark	9
–	Dorsal color pale cream with diagonal pale bars and lighter pattern, head pattern smudgy appearance	Morelia spilota mcdowelli
9.	Body ground color dark brown or blackish	Morelia spilota cheynei
–	Body ground color shade of brown or reddish-brown	10
10.	Body pattern consists of 60–70 pale rings	Morelia spilota variegata
–	Body pattern with pale rings but connected by two lateral pale lines	Morelia spilota harrisoni
11.	Ventrals fewer than 400, subcaudals fewer than 125, infralabials fewer than 22, parietal scales not fragmented	12
–	Ventrals more than 400, subcaudals more than 155, infralabials more than 22, parietal scales fragmented	Morelia oenpelliensis
12.	Overall glossy blackish head and body color with white or yellowish bars in the labials	Morelia boeleni
–	Overall head and body color variable	13
13.	Neck bar pattern absent	14
–	Neck bar pattern present	16
14.	Postocular stripe absent	15
–	Postocular stripe present	Morelia nauta
15.	Suboculars absent, single supraocular	Morelia kinghorni
–	Suboculars present, 2–3 supraoculars	Morelia clastolepis
16.	Iris color golden, 0–2 interparietals	Morelia amethistina
–	Iris color red, 2–3 interparietals	Morelia tracyae

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Python

1.	Small or fragmented head scales	2
–	Large, well-developed head scales	4
2.	Midbody scale rows fewer than 75, subcaudal scale counts fewer than 50	3
–	Midbody scale rows more than 75, subcaudal scale counts more than 60	Python natalensis
3.	Ventral scale counts fewer than 210, subcaudals fewer than 38	Python regius
–	Ventral scale counts more than 250, subcaudals more than 46	Python anchietae
4.	Ventral scale counts fewer than 200	5
–	Ventral scale counts more than 200	7
5.	Ventral scale counts fewer than 167	6
–	Ventral scale counts more than 167	Python brongersmai
6.	Anterior pair of parietals not in contact or are only weakly contacting	Python curtus
–	Anterior pair of parietals in broad contact at the medial structure	Python breitensteini
7.	Dorsal midbody scale rows fewer than 75	8
–	Dorsal midbody scale rows more than 75	Python sebae
8.	Suboculars absent	9
–	Suboculars present, separating the supralabials from the orbit	10
9.	Two preoculars present, subcaudals 66–70	Python molurus molurus
–	Three preoculars present, subcaudals 57–65	Python molurus pimbura
	(Additional diagnostic information: longitudinal pink marking above the eyes, fewer dark blotches that also invade the ventral scutes)
10.	Pale centered saddles, pale-centered brown blotches	Python bivittatus bivittatus
	(Additional diagnostic information: attains larger size up to 5m in length)
–	Prevalent saddles with pale margins, increased incidence of ocellic blotches (more molurus-like)	Python bivittatus progschai
	(Additional diagnostic information: does not exceed 2.5 m in total length)

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Associated Data

This section collects any data citations, data availability statements, or supplementary materials included in this article.

Supplementary Materials

XML Treatment for Antaresia

zookeys.66.683-treatment1.xml^{(25.5KB, xml)}

XML Treatment for Apodora

zookeys.66.683-treatment10.xml^{(10.9KB, xml)}

XML Treatment for Aspidites

zookeys.66.683-treatment12.xml^{(24.6KB, xml)}

XML Treatment for Aspidoboa

zookeys.66.683-treatment20.xml^{(4.4KB, xml)}

XML Treatment for Australiasis

zookeys.66.683-treatment21.xml^{(11.8KB, xml)}

XML Treatment for Austroliasis

zookeys.66.683-treatment25.xml^{(2.8KB, xml)}

XML Treatment for Bothrochilus

zookeys.66.683-treatment26.xml^{(3.4KB, xml)}

XML Treatment for Broghammerus

zookeys.66.683-treatment28.xml^{(33.5KB, xml)}

XML Treatment for Chondropython

zookeys.66.683-treatment40.xml^{(16.3KB, xml)}

XML Treatment for Heleionomus

zookeys.66.683-treatment45.xml^{(3.9KB, xml)}

XML Treatment for Helionomus

zookeys.66.683-treatment46.xml^{(3KB, xml)}

XML Treatment for Jackypython

zookeys.66.683-treatment47.xml^{(2.3KB, xml)}

XML Treatment for Katrinus

zookeys.66.683-treatment48.xml^{(13.5KB, xml)}

XML Treatment for Leiopython

zookeys.66.683-treatment52.xml^{(20.4KB, xml)}

XML Treatment for Lenhoserus

zookeys.66.683-treatment60.xml^{(3.9KB, xml)}

XML Treatment for Liasis

zookeys.66.683-treatment61.xml^{(27.4KB, xml)}

XML Treatment for Montypythonoides

zookeys.66.683-treatment71.xml^{(2.4KB, xml)}

XML Treatment for Morelia

zookeys.66.683-treatment72.xml^{(69.7KB, xml)}

XML Treatment for Python

zookeys.66.683-treatment97.xml^{(68.4KB, xml)}

XML Treatment for Rawlingspython

zookeys.66.683-treatment114.xml^{(2.9KB, xml)}

XML Treatment for Shireenhoserus

zookeys.66.683-treatment115.xml^{(3.1KB, xml)}

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PERMALINK

Annotated checklist of the recent and extinct pythons (Serpentes, Pythonidae), with notes on nomenclature, taxonomy, and distribution

Wulf D Schleip

Mark O’Shea

Abstract Abstract

Introduction

Taxonomic changes since 1999

Approach and Scope

Interpretation and application of the Code

APP1.

APP2.

APP3.

APP4.

APP5.

APP6.

APP7.

APP8.

Abbreviations for Depositories of type material

Checklist of the Pythonidae

Genus. Antaresia

Remarks:

Antaresia childreni

Antaresia maculosa

Synonyms:

Distribution:

Antaresia maculosa brentonoloughlini

Holotype:

Type locality:

Remarks:

Antaresia perthensis

Antaresia stimsoni

Antaresia stimsoni stimsoni

Antaresia stimsoni campbelli

Holotype:

Type locality:

Remarks:

Antaresia stimsoni orientalis

Synonyms:

Remarks:

Genus. Apodora

Remarks:

Apodora papuana

Synonyms:

Genus. Aspidites

Remarks:

Aspidites melanocephalus

Synonyms:

Aspidites melanocephalus adelynensis

Holotype:

Type locality:

Remarks:

Aspidites melanocephalus davieii

Holotype:

Type locality:

Remarks:

Aspidites melanocephalus rickjonesii

Holotype:

Type locality:

Remarks:

Aspidites ramsayi

Synonyms:

Holotype:

Type species:

Remarks:

Aspidites ramsayi panoptes

Synonyms:

Holotype:

Type locality:

Remarks:

Aspidites ramsayi richardjonesii

Holotype:

Type locality:

Remarks:

Genus. Aspidoboa

Remarks:

Genus. Australiasis

Synonyms:

Remarks:

Australiasis amethystinus

Australiasis amethystinus clarki