A
RECLASSIFICATION OF THE PYTHONINAE INCLUDING THE DESCRIPTIONS OF TWO NEW
GENERA, TWO NEW SPECIES AND NINE NEW SUBSPECIES
Raymond
Hoser
488 Park Road, Park Orchards, Victoria, 3134, Australia.
Phone:
+61 3 9812 3322 Fax: 9812 3355 E-mail: adder@smuggled.com
E-mail:
adder@smuggled.com
Originally
published in Crocodilian - Journal of the Victorian Association of Amateur
Herpetologists
4(3)
November 2003:31-37 and 4(4) June 2004:21-40.
ABSTRACT
As
a result of a reclassification of the Pythoninae, this paper now splits
the group into sixteen genera, including all those used by Hoser (2000)
and two resurrected from the synonymy of the catch-all genus "Python",
namely Aspidoboa from Asia and Helionomus from Africa. Furthermore
two new genera have been erected for the reticulatus and anchietae
groups, that had also been formerly placed within the genus Python.
These
are Broghammerus gen. nov. and Shireenhoserus gen. nov. respectively.
This
paper redescribes taxa described from Australia as named in Hoser (2003),
citing them here as "sp. nov." although the proper designation should be
"Hoser 2003". These are two new species of Morelia from South Australia,
bringing the known total of python species to 42.
There
are six new regional subspecies of Broghammerus reticulatus (previously
known as Python reticulatus) formally described and named, as well
as new subspecies of Chondropython viridis, Antaresia maculosus
and Katrinus fuscus formally described and named. It is anticipated
that further subspecies of Broghammerus in particular as well as
other pythons await diagnosis and formal description.
This
paper also lists all 42 recognised species as of 2003, including those
most recently described by this and other authors.
INTRODUCTION
AND
MATERIALS AND METHODS
The
"true" pythons of Africa, Asia and Australasia are generally placed in
the sub-family Pythoninae.
The
classification of the pythoninae as defined by Romer (1956) p. 572, has
in recent years been the subject of intense debate. The pythoninae are
separated from the boinae by the presence of a supraorbital bone and other
characters (McDowall 1975).
There
have also been numerous published studies detailing the morphology and
biochemistry of the pythoninae and related snakes.
Notwithstanding
this vast body of available evidence the taxonomic arrangements used in
the past by most authors have been generally inconsistent across the group.
By contrast, this paper consistently applies the same principals across
the entire group as best as is possible. It does not attempt to rehash
other researchers findings of fact.
Due
to a number of factors including the commercial significance of these snakes,
their generally large size and popularity among hobbyist keepers, there
have been numerous studies into the taxonomy of these snakes.
Furthermore,
there has also been a vast amount of previously unreported or relatively
little-known information about the pythoninae published in other scientific
and popular literature, including in such publications as: Banks (1974,
1980), Barker and Barker (1994a, 1994b, 1995, 1999), Barnett (1979, 1987,
1993, 1999), Broghammer (2001), Bullian (1994), Chiras (1982), Comber (1999),
Covacevich and Limpus (1973), Cox (1991), David and Vogel (1996), Dunn
(1979), Ehmann (1992), Fearn (1996), FitzSimmons (1970), Gharpurey (1962),
Gow (1977, 1981, 1989), Greer (1997), Heijden (1988), Hoser (1981a, 1981b,
1981c, 1982, 1988, 1989, 1990, 1991a, 1991b, 1992, 1993a, 1993b, 1995,
1996, 1999a, 1999b, 1999c, 1999d), Kend (1992, 1997), Kend and Kend (1992),
Kortlang (1989), Krauss (1995), Maguire (1995), Martin (1973), Maryan (1984),
Maryan and George (1998), Mattison (1980), Mavromichalis and Bloem (1994),
McDowell (1984), McLain (1980), Mirtschin and Davis (1992), Murdoch (1999),
O’Shea (1996), Reitinger (1978), Romer (1956), Rooyendijk (1999), Ross
(1973, 1978), Ross and Marzec (1990), Schwaner and Dessauer (1981), Sheargold
(1979), Shine (1991), Shine, Ambariyanto, Harlow, Mumpuni, (1998), Smith
(1981a, 1981b, 1985), Sonneman (1999), Storr, Smith and Johnstone (1986),
Stull (1932, 1935), Thomson (1935), Webber (1978), Weigel (1988), Wells
and Wellington (1983, 1985), Williams (1992), Wilson and Knowles (1988),
Worrell (1951, 1970) and the many further sources of information referred
to directly in these publications.
This
forms an enormous database of information on these snakes.
As
a result of this continual inflow of new information, there have been numerous
taxonomic arrangements proposed for these snakes at both the genus and
species levels.
As
a result, calls by the ICZN for stability of nomenclature as per their
rules (Ride et. al. 1999 and earlier ICZN publications) have effectively
been ignored in terms of this group of reptiles.
Thus
it is appropriate for a classification to be adopted that more accurately
reflects the true phylogeny of the pythoninae, even if it results in previously
unused names being used and there being some short-term difficulty by other
herpetologists in terms of getting used to the newer and more accurate
nomenclature.
In
recent times (the last 20 years) and at it’s most conservative, the pythoninae
has been deemed to consist of just three genera, namely Aspidites,
Morelia and Python, by authors such as Ingram and Raven (1991)
and Welch (1994).
However,
most herpetologists (e.g. Greer (1997) or Wells and Wellington (1985)),
including this author, find the above arrangement untenable and this paper
accepts the alternative arrangement, which must involve an unbiased splitting
of the genera Morelia and Python as recognised by Ingram
and Raven (1991) and Welch (1994). Hence an expansion here of the earlier
arrangement of Hoser (2000).
The
placement of the Woma and Black-headed Pythons (Aspidites ramsayi
and Aspidites melanocephalus) into the relatively distinct genus
Aspidites, has been uncontroversial and accepted by all modern herpetologists
and is not further discussed in this paper.
It’s
generally agreed that this group diverged from the other pythons a long
way back in the geological past and well before the rest of the pythoninae
diverged. Aspidites are thought to be the most primitive of the
pythoninae due to their lacking labial pits.
Further
reference to Australasian pythons in this paper should be taken to ignore
that genus and instead refer to all the other species (unless otherwise
stated).
Other
classification systems adopted have in turn split the latter two genera
(Morelia and Python) into numerous genera each consisting
of one or more closely related species.
Recent
taxonomic studies invoking DNA techniques have indicated that it is in
fact more reasonable to split the latter two genera into the various species
groups as recognised by earlier authors such as McDowall (1975), Wells
and Wellington (1985) and more recently Hoser (2000).
Most
of the various taxonomic arrangements for the pythoninae have invariably
looked at these snakes on a regional basis only (like Wells and Wellington
(1985) or Hoser (2000)), and while generally consistent in their approaches,
have by their nature ignored taxa found outside their target area.
The
relatively few publications dealing with pythoninae taxonomy on a world-wide
basis have either tended to (in hindsight) be overly conservative in lumping
most species into the catch-all genus "Python", or been generally
inconsistent in terms of the criteria used to define a genus and then assign
species to them.
This
same pattern has been particularly evident in the popular literature, which
adopts a "splitting" (sensu stricto) approach in terms of the Australasian
species by subdividing the species into various genera including such as
Liasis, Bothrochilus and Morelia, while simultaneously
ignoring similar splits of the African and Asian pythons from the single
"Python" (sensu lato) into the formally named subgroups (genera)
of Aspidoboa, Helionomus and Python, (e.g. Stafford
1986), or even a simple and obvious splitting of the "Python" group
into the molurus and reticulatus groups as so eloquently
identified by McDowall (1975).
This
paper is the first paper to propose a classification system for all the
pythoninae, reflecting more modern views of what should be included in
a given genus and taking into account recent studies on the taxonomy of
the pythoninae as a whole and the various well-defined species groups.
This
paper is also consistent in that it also follows on the trend of recent
splits of other reptile genera which had formerly been regarded as single
into two or more genera, based on distinct groups of reptiles within a
genera, separated easily on the basis of size class (big versus small)
and obvious and very different differences in morphology, scalation and
biology.
There
are numerous such examples including: Couper, Covacevich and Moritz (1993),
who split the gekkonid genus Phyllurus into two based on the above
criteria, Hoser (1998) who split the elapid genus Cannia into two
also based on similar criteria. Then there is the most obvious and now
widely accepted splitting of "Liaisis" into more than one genus
including the new genus Antaresia (Wells and Wellington 1983), which
is identified popularly as the "small Liasis" and separated most
easily from the others on the basis of their radically different adult
size.
Greer
(1997) and others have accepted this proposition and the name Antaresia
is now in general usage.
Among
the better known taxonomic studies in recent times dealing specifically
with the Pythoninae are Harvey, Barker, Ammerman and Chippendale. (2000),
Keogh, Barker and Shine (2001), Kluge (1993), McDowall (1975) and Underwood
and Stimson (1990).
This
paper draws on the findings of fact by these papers and the other publications
cited at the rear of this paper to make the classification given below,
but as a rule does not rehash these findings here as the facts themselves
are generally not in dispute.
The
classification system proposed is consistent with that of Hoser (2000),
however unlike that paper, it deals with the status of all pythons and
not just those from the Australia/New Guinea region.
This
paper also builds on the taxonomy of Harvey, Barker, Ammerman and Chippendale.
(2000) for the genus Australiasis, which was omitted from the paper
Hoser (2000) following an earlier request from one of the authors, David
Barker.
The
taxonomy of Harvey, Barker, Ammerman and Chippendale. (2000) is accepted
almost in toto at the species level, save for the resurrection of the species
duceboracensis (Gunther, 1879) from the synonymy of amethistina
(Schneider, 1801). Furthermore, unlike Harvey, Barker, Ammerman and
Chippendale. (2000), this author regards Australian Australiasis
as subspecific to southern New Guinea Australiasis amethistina.
The correct terminology for that taxon is Australiasis amethystinus
clarki (Barbour, 1914), not "kinghorni", which is effectively
a junior synonym.
The
central thrust of this paper is that the allocation of given species to
given genera should be consistent across the pythoninae, regardless of
which region the species occurs.
As
a result, in the classification that follows, one will see the resurrection
of old and/or little-known generic names and the formal naming of two new
python genera.
This
paper does not purport to be a comprehensive review of all pythoninae species
and subspecies. The author believes that further taxa will be formally
described at both the species and subspecies level in years to come and
that taxa treated here as subspecies may ultimately be generally accepted
as full species, as has already been seen for some of the taxa described
by Hoser (2000) by people such as Wells (2002).
Diagnostic
information for the Australasian genera as named in Hoser (2000), namely
Aspidites Peters 1876, Antaresia Wells and Wellington 1983,
Australiasis Wells and Wellington 1983, Bothrochilus Fitzinger
1843, Chondropython Meyer 1874, Katrinus Hoser 2000, Leiopython
Hubrecht 1879, Lenhoserus Hoser 2000, Liasis Gray 1840, Morelia
Gray 1842, Nyctophilopython Wells and Wellington 1985 is not repeated
in this paper. That paper is on the internet at the url http://www.smuggled.com/pytrev1.htm
and is also available from that site as pdf in the same format as originally
published.
For
detailed diagnostic information about the genera named in the previous
paragraph, readers are hereby directed to the original descriptions as
cited and/or the popular literature which more than adequately separates
the best known component species as cited at the rear of this paper. This
paper does not amend the taxonomy used in the paper Hoser (2000), save
for the addition of more recently described species and/or subspecies.
The taxonomy used in Hoser (2000) has been widely adopted in the two years
since publication. Examples include: Clark 2002, Kuroski 2001 and 2002
(all for Morelia harrisoni), and Schleip 2001 (for the various subspecies
of L. albertisi) as named formally by Hoser (2000).
In
this paper is a list of all currently recognised genera and species and
subspecies of python, including the two genera formally named for the first
time, namely Shireenhoserus gen. nov. and Broghammerus gen.
nov. and the newly named species and subspecies.
RELEVANT
DISCUSSION AND COMMENTS
The
following is directly relevant to the formal descriptions that follow,
the general taxonomy used and forms a part of the descriptions and this
paper.
The
list as follows in itself shows the taxonomic conclusions made by this
author.
However
a few other comments in this regard are warranted.
Two
genera of snakes, namely Loxocemus and Calabaria are not
closely related to other python genera and were removed from the Pythoninae
in 1976 by Underwood. They are now placed in different subfamilies or in
a different family altogether.
More
recent evidence (including Heise, et. al. (1995) p. 261, Fig. 1.) confirms
this move by Underwood and those two genera have been effectively ignored
for the purposes of this paper.
Calabaria
are readily separated from all true pythons (excluding Aspidites
from Australia) by their more-or-less cylindrical body shape and the fact
that their head is not distinct from the neck as in true pythons (again
excluding Aspidites).
Aspidites
can be readily separated by their yellowish brown body colour and dorsal
pattern with a tendency towards distinct or indistinct transverse banding.
By contrast for Calabaria the dorsal colour in Calabaria
is a more dark and reddish brown and the pattern is not tending towards
transverse banding in any way.
The
results of Heise, et. al. (1995) also suggests that the old-world Pythoninae
should in fact be elevated to the level of family, thereby excluding the
boids from the new world, who in turn should be placed into a separate
family. Likewise for the Calabariinae from Africa and Loxoceminae from
North America.
The
relationships of the subfamilies Bolyeriinae, and Erycinae with respect
to the other "boids" remains generally uncertain, but it is obvious that
they do not form a part of the Pythoninae.
This
author agrees with Stimson (1969) p. 28, in designating molurus
the type species for the genus Python.
The
past moves by Kluge (1993) to make Katrinus mackloti and Katrinus
fuscus synonymous were rejected by this author in Hoser (2000) and
have been corroborated by other authors (again see Hoser (2000)). Likewise
for Kluge’s (1993) erection of the genus Apodora to accommodate
the species Liasis papuana (see also below).
The
evidence does not support the position of Kluge (1993).
Neither
move is to be taken as a personal attack against Arnold Kluge in any way.
The
results and data as published by Underwood and Stimson (1990) p. 592 top
and elsewhere in the same paper and several more recent critiques of the
same paper give unequivocal support for the erection of the genus Katrinus
by Hoser (2000) and to a lesser extent give support to the erection of
the genus Lenhoserus by Hoser (2000) if one is to accept the proposition
that the species viridis should be separated from the other Morelia
and placed into it’s own genus Chondropython, and/or that the Scrub
Pythons (Australiasis) should be placed in a genus on their own
apart from Morelia and Chondropython.
Harvey,
Barker, Ammerman and Chippendale (2000) provided sufficient evidence for
the formal recognition of Australiasis duceboracensis (Gunther,
1879) from New Ireland in the Bismark Archipelago to be recognised as a
full species (as opposed to being merely a local variant of Australiasis
amethistina), but in the end of the paper failed to make this obvious
move.
Hence
it’s inclusion in the list here.
In
further explanation of this move, this author notes that an analysis of
the cytochrome b mitochondrial DNA sequence by Harvey, Barker, Ammerman
and Chippendale (2000) for the Bismark Islands form showed a 5% divergence
from the nominate New Guinea form.
In
a later paper by Keogh, Barker and Shine (2001), two other python species
(namely breitensteini and curtus) were confirmed as being
distinct at the species level with a mere 3% divergence of the same cytochrome
b mitochondrial DNA sequence using the same test.
It
didn’t escape this author’s notice that David Barker was a co-author of
both papers and hence one finds it hard to otherwise reconcile this inconsistency.
No
disrespect is implied here, however it is important that the inconsistency
be appropriately corrected.
Harvey,
Barker, Ammerman and Chippendale (2000) also called for the designation
of a neotype for Australiasis amethistina. That call is agreed by
this author. The data provided by Harvey, Barker, Ammerman and Chippendale
(2000) and obtained independently by this author confirms that the "Australiasis
amethistina" from north of the central divide in New Guinea are different
to the southern form, at least to subspecies level, which is a situation
mirrored in the snakes of the genera Acanthophis and Leiopython
who have similar distributions in this region (see Hoser (1998a) and Hoser
(2000) respectively).
This
author was going to formally name this form of Australiasis at the
subspecies level, but has deferred doing so, pending advice that others
were independently working on doing this. Hence this taxa will be ignored
for the purposes of this paper and the list that follows.
In
the absence of a neotype and any evidence to the contrary, this author
assumes that the species name "dipsadides" from south of the central
divide in New Guinea is merely a junior synonym of amethistina.
The
African species name "saxuloides" (Miller and Smith 1979) is merely
a junior synonym for the species sebae. Also see Broadley (1984).
The
obvious physical character differences between the smaller species of Asiatic
and African pythons from the larger species (herein listed as: Python,
Helionomus and Broghammerus gen. nov.) and the lack of any
recent evidence of common ancestry make a compelling case for the resurrection
of and creation of a total of two new genera to accommodate the Asian and
African species respectively.
Again
refer to the examples of: Couper, Covacevich and Moritz (1993), Hoser (1998),
Wells and Wellington (1983) to show that the move indicated by this author
is consistent with the modern scientific viewpoint.
The
species within each of the five relevant genera Python, Aspidoboa,
Helionomus, Broghammerus gen. nov. and Shireenhoserus gen.
nov. can all be separated from one another by the differences in the following
character states in combination: number of and position of labial pits,
average adult size, general build, typical head markings and body colouration,
breeding biology and average relative egg size and number, head and body
scalation. A cursory examination of the relevant component species will
more than adequately establish this fact.
These
characteristics for each species are detailed in general regional texts,
including those cited at the end of this paper.
NOTES
ON THE DESCRIPTIONS OF NEW
SUBSPECIES
IN THIS PAPER
Should
at some future stage any herpetologist choose for their own purposes to
recognize some but not all of the subspecies listed for the first time
in this paper and/or elevate some to the status of full species, thereby
perhaps making others junior synonyms at the species level, then the names
that take precedence should be those that are given first in this paper
(by alphabetical order).
Specifically
in relation to the regional variants of Broghammerus gen. nov.,
these are in many ways like those of Morelia or Australiasis
in that most specimens of a given species or subspecies can fairly easily
be referred to a given taxa on appearance alone, once a person is familiar
with them. But due to the huge degree of variation within a single population,
this is not always the case, hence the need to retain good locality data
for specimens and/or the need for relevant genetic tests if necessary.
In
terms of the subspecies listed below, this author’s investigations have
led to the inescapable conclusion that all are valid and hence the descriptions
published here.
For
the Broghammerus taxa described, it is likely that sensu stricto
many are in fact species, rather than merely subspecies as described here.
All are known to be reproductively isolated from one another in the wild
state. This was used by Harvey, Barker, Ammerman and Chippendale. (2000)
as an important factor in terms of the defining Australiasis species
in their paper.
However
this author has for the present time taken the conservative approach with
respect to naming these new taxa.
It
is anticipated that most of these taxa will inevitably be elevated to full
species status at a later time as more data becomes available.
NOTES
ABOUT THE LIST THAT FOLLOWS
Following
is a list of all species or subspecies of pythoninae known. It is in alphabetical
order.
For
some groups and taxa there is a more detailed account in line with the
descriptions that form the most important parts of this paper.
As
a rule, after each species or subspecies name is a brief indication of
the known distribution.
Synonyms
as listed in Cogger et. al. (1983), Harvey, Barker, Ammerman and Chippendale.
(2000), Kluge (1993) and Romer (1956) are not repeated here. In combination,
those lists are believed to be comprehensive. Names used in those papers
with reference to given taxa, either as species in their own right or as
junior synonyms are all regarded as junior synonyms to the names used here
unless otherwise indicated.
Aspidoboa
and Helionomus are both valid names as per the relevant sections
of the ICZN’s Code (fourth edition) and are therefore used here even though
both are very old and currently little-known.
Below
and where appropriate, the nominate subspecies is only indicated by the
binomial rather than as a trinomial. Distribution information given in
these instances for the nominate form only applies to this subspecies and
not all others. The list runs alphabetically and newly described taxa are
placed in this list where appropriate.
PYTHONINAE
SPECIES AND SUBSPECIES OF THE WORLD
GENUS
ANTARESIA WELLS AND WELLINGTON 1983
Antaresia
childreni (Gray 1842) (N. Australia)
Antaresia
maculosus (Peters 1873) (NE Australia along coast and ranges south
of about Cairns, Qld)
ANTARESIA
MACULOSUS BRENTONOLOUGHLINI subsp. nov.
HOLOTYPE
A specimen
at the Australian Museum from 16 km east of Coen, Queensland (R16772) (Lat.
13° 55' S, Long. 143° 11' E).
DIAGNOSIS
Known
in herpetoculture as "Blonde maculosus" this subspecies is the (usually)
large light coloured form from far north Queensland.
In
the absence of reliable locality data and DNA data, both of which separate
Antaresia maculosus brentonoloughlini subsp. nov. from Antaresia
maculosus maculosus this newly described subspecies can be readily
separated from other Antaresia maculosus by it's greater preponderance
of light colouration relative to dark blotches on the dorsal surface. Normal
Antaresia maculosus maculosus have roughly half to half (50%:50%)
dark versus light blotches. For Antaresia maculosus brentonoloughlini
subsp. nov. the ratio is generally at least 60% light colour to 40% or
less darker blotches.
Antaresia
maculosus brentonoloughlini subsp. nov. is also on average a larger
subspecies, with specimens attaining 150 cm being fairly common. This size
is relatively rare in normal Antaresia maculosus.
Antaresia
maculosus brentonoloughlini subsp. nov. is known only from Cape York
in Queensland to about as far south as just north of Cairns. In the region
from about Cairns to Townsville and inland from here, specimens are in
many respects intermediate in form between Antaresia maculosus brentonoloughlini
subsp. nov. and Antaresia maculosus maculosus, but are probably
best assigned to the form Antaresia maculosus maculosus on the basis
of their smaller adult maximum sizes and the fact that their dorsal patterning
is generally more like that of normal Antaresia maculosus maculosus.
Antaresia
maculosus brentonoloughlini subsp. nov. is a hardy captive and common
in captivity in Australia. A number of NSW breeders of snakes breed large
numbers of Antaresia maculosus brentonoloughlini subsp. nov. (as
of 2001-2002). Husbandry requirements for all Antaresia maculosus subspecies
appears to be identical in terms of how cages are set-up, incubation of
eggs and treatment of common ailments.
ETYMOLOGY
Named
in honour of Mr Brenton O'Loughlin of Access Lawyers in Melbourne, who
with his legal partner Mr. Gabriel Kuek, have taken on a number of important
public interest legal cases on a "pro bono" basis at a time when not one
other Melbourne lawyer would take on such matters for fear of reprisals
from the authorities. These two men have also been harassed by police and
others as a result of their taking a stand on behalf of truth, honesty
and against corrupt and dishonest people in the state Police, parliament
and their mates they improperly appointed to the judiciary..
Honest
and decent lawyers appear to be rare on the ground and these two men have
made an important contribution to Australia and have been improperly attacked
by others with vested and corrupt interests in the legal system. As the
Australian government's honours system has been so severely debased in
that corrupt and vested interests seem more pre-occupied with honouring
drug dealers and pedophiles rather than decent people, this author takes
the opportunity here to honour someone who should be honoured and recognised.
Antaresia
perthensis (Stull 1932) (Pilbara, West Australia)
Anteresia
saxacola Wells and Wellington 1985 (Central Australia)
Antaresia
saxacola campbelli Hoser 2000 (SE Inland Australia)
Antaresia
saxacola stimsoni (Smith 1985) (Western Australia)
GENUS
ASPIDITES PETERS 1876
Aspidites
melanocephalus (Krefft, 1864) (NE Australia)
Aspidites
melanocephalus adelynensis Hoser 2000 (Kimberleys, West Australia)
Aspidites
melanocephalus daviei Hoser 2000 (Pilbara, West Australia)
Aspidites
ramsayi (Macleay, 1882) (SE inland Australia)
Aspidites
ramsayi panoptes Hoser 2000 (SW West Australia)
Aspidites
ramsayi richardjonesi Hoser 2000 (NW West Australia)
GENUS
ASPIDOBOA SAUVAGE 1884
Aspidoboa
breitensteini (Steindachner 1880) (Borneo)
Aspidoboa
brongersmai (Stull 1938) (Malay Peninsula and Sumatra)
Aspidoboa
curtus (Schlegel 1872) (West Sumatra)
Genus
Australiasis Wells and Wellington 1983
Australiasis
amethistina (Schneider 1801) (Southern New Guinea)
Australiasis
amethystinus clarki (Barbour 1914) (NE Australia)
Australiasis
clastolepis (Harvey et. al. 2000) (Mollucan islands, Indonesia)
Australiasis
duceboracensis (Gunther 1879) (Bismark Archipelago)
Australiasis
nauta (Harvey et. al. 2000) (Tanimbar Islands, Indonesia)
Australiasis
timorensis (Peters 1877) (Timor)
Australiasis
tracyae (Harvey et. al. 2000) (Halmahera island, Indonesia)
Genus
BOTHROCHILUS FITZINGER 1843
Bothrochilus
boa (Schlegel 1837) (Bismark Islands, PNG)
GENUS
BROGHAMMERUS GEN. NOV.
Type
species: Boa reticulata Schneider 1801
DIAGNOSIS
A group
of extremely large pythons from the south-east Asian region.
Up
until now, this genus has comprised just one known species, namely the
Reticulated Python (Broghammerus reticulatus).
It’s
known distribution is the Indo-Malay archipelago from the small islands
just west of the Island of New Guinea west to the Indochina countries of
Thailand, Burma, Vietnam and Laos and including the Philippines (see the
more detailed listing below). However it appears that several similar taxa
have in fact been lumped within this species under the single name "reticulatus".
The regional variants now known as Broghammerus reticulatus, will
be further subdivided in the future at least to the subspecies level as
has now been done for some of the better-known regional variants (see formal
descriptions below).
These
snakes are separated from one another by a host of general traits including
physical morphology, DNA traits, colour patterns, colours, eye and/or iris
colour and so on, as well as distribution on different land masses.
There
has not been a detailed analysis of scale-count variation between the regional
subspecies and preliminary evidence suggests much overlap in this character
between various populations.
Due
to the fact that "reticulatus" as a whole is listed by the IUCN
as "vulnerable" and the increasing rate of harvesting the species for the
skin trade and general habitat destruction throughout the relevant region,
the conservation status of several regional variants may be precarious
and hence the sooner that all regional races are formally delineated, named
and properly assessed, the better.
The
identification of new subspecies of Broghammerus reticulatus in
this paper does not purport to be a comprehensive listing of all subspecies.
It is anticipated that as more specimens are collected from other parts
of the range of Broghammerus, including those areas currently subject
to civil unrest, further variants will be formally named.
In
any event, due to the fact that all the previously used names for the species,
namely Boa reticulata (Schneider, 1801), Boa rhombeata (Schneider,
1801), Boa phrygia (1802) and Python schneideri (Merrem 1820),
and since made synonymous lack anything remotely resembling accurate locality
data, save perhaps for Seba (1734), pl. 62, it is clear that all must therefore
be taken now to identify the same single race of the species.
Based
on the original description of the nominate species and the only remotely
accurate location data given, namely: "Orient", Seba (1734), it must be
deemed that the typical race of Broghammerus reticulatus is that
found in the general area of Singapore, mainland south-east Asia, including
peninsula Malaysia or immediately adjacent Islands, to which the name "Orient"
is usually referred.
This
author calls for a neotype to be designated for the species, preferably
of the "typical" Malay Peninsula form, such as that found in the general
vicinity of Singapore. There are numerous such specimens in Museum Collections.
This
is the largish regional race with a brownish head, much the same colour
as the lighter dorsal body markings, although light-headed specimens are
known and several colour variants and distinct colour mutations are also
known.
The
only snakes with which this genus (Broghammerus gen. nov.) could
likely be confused with are those of the genus Python which as defined
here in this paper only includes the species molurus.
They
are readily separated by a host of characters including the fact that they
usually have four pitted supralabials versus just two in Python molurus
(and Helionomus sebae).
McDowall
(1975), pages 50-51 separated Broghammerus gen. nov. from all other
Afro/Asian Pythons, which he put into the so-called "molorus group",
with a suite of characteristics that in themselves largely diagnosed the
former genus (Broghammerus) at least in as much as separating it
from other Afro/Asian python genera Python, Aspidoboa, Helionomus
and Shireenhoserus gen. nov..
To
repeat some of McDowall’s information here, this included the fact that
the supralabial pits in Broghammerus gen. nov. are diagonal slits
and less deeply impressed than the square pits of the more posterior infralabials;
the infralabial pits are set in a distinct groove, defined ventrally by
a longitudinal fold (similar to Leiopython albertisi). By contrast
in the so-called "molorus group" (which included all of genera Python,
Aspidoboa, Helionomus and Shireenhoserus gen. nov.) the infralabial
pits are much more shallow than the supralabial pits and are not set in
a groove; the supralabial pits are square or triangular.
The
dorsal colouration in Broghammerus is also radically different to
all other pythons as seen by comparative photos, including those published
in Stafford (1986).
In
Broghammerus the dorsal pattern in wild specimens is virtually always
that which gives the markings a reticulated appearance, hence the common
name "Reticulated Python". With the possible exception of some Python
molurus and Helionomus sebae, which to an unexperienced person
may appear to have a reticulated pattern, no other African or Asian pythons
have a dorsal patterning resembling anything like Broghammerus.
Broghammerus
are different to and readily separated from all Python, Shireenhoserus
and Helionomus by the following trait. In Broghammerus there
is generally a dark mid dorsal line running from the rear to the front
of the head and bordered on either side by distinctly lighter scales.
None
of the other three genera share this trait. Furthermore there is no line,
border or blotch running flat between the eye and the snout along the upper
side region of the head as is seen in the three other genera.
A number
of breeders have successfully propagated a number of unusual (and often
patternless) mutations, which do not necessarily conform to the colour
information just given. These are now common in herpetoculture. However
these are still easily placed within their given species (and genus) due
to the other parameters outlined in this paper.
Broghammerus
can be further separated from Python and Helionomus and
Shireenhoserus gen. nov. by it’s iris colour, being red or orange,
versus brownish.
Aspidoboa
are readily separated from Broghammerus by their smaller adult
size and distinctly stout build.
In
any event, Aspidoboa are separated from all other pythons by the
presence of a naso-preocular groove, which is a diagonal division that
passes between the large facial scales from the upper posterior of the
nasal scale posterior to the lower prefrontals. The groove is bounded above
by the prefrontals and upper preocular and below by 1-3 large loreals and
the upper margins of the supralabials. The groove itself is scaled with
a series of small to granular loreal scales. Aspidoboa pythons also
have a significantly shorter tail-length to head-length ratio than all
other pythons (1.6 for Aspidoboa). See Keogh, Barker and Shine (2001)
for further details.
In
terms of cytochrome b mitochondrial DNA sequence data, Keogh, Barker
and Shine (2001) found a 10.3% divergence between Aspidoboa brongersmai
and Broghammerus reticulatus in their tests.
The
only other pythons remotely likely to confused with Broghammerus
are those of the genus Australiasis. Australiasis species
can be separated from Broghammerus where they occur more-or-less
together (Flores and Timor) by the fact that they are essentially patternless
dorsally. They are also of generally slighter build and smaller adult size.
Other
diagnostic information for the genus Broghammerus can be readily
gleaned from the literature as cited at the end of this paper, or the excellent
list of sources specific for Broghammerus as cited by Uetz (2002).
The
modern day distribution of Broghammerus suggests that it is a recently
evolved species group that has expanded it’s distribution within a very
short period of time.
This
may be due to the fact that Broghammerus appears to have crossed
(by swimming or rafting) some sizeable water barriers, to almost cross
the divide from Asia to Australasia (assuming that it evolved on the Asian
side).
However
although found on Islands within the Australasian region, Broghammerus
hasn’t yet made it to the Australia/New Guinea mainlands.
Corroboration
of this hypothesis comes from the fact that the species is known to have
"rafted" or swam to the Island of Krakatoa after life there was extinguished
by a volcanic eruption.
However
in partial rebuttal of this theory was the findings by Harvey, Barker,
Ammerman and Chippendale. (2000) that there was little if any gene flow
evident between the main island populations of Austaliasis, whose
distribution and mobility across bodies of extant water appears to be less,
but had previously been assumed to be greater.
These
findings indicated that present day distributions of most Australian and
Asian pythons may also reflect in large part earlier distributions of land
masses at times of lower sea levels.
Based
on this most recent finding by Harvey, Barker, Ammerman and Chippendale.
(2000) and the fact that the water barriers separating populations of Broghammerus
are often huge, it seems inevitable that the various major populations
of Broghammerus will ultimately be recognised as full species rather
than as subspecies as indicated here.
However
as already indicated, in this paper, the conservative approach has been
taken with the known variants merely being named as subspecies.
ETYMOLOGY
The
genus is named in honour of German herpetologist Stefan Broghammer for
his general contributions to the science of herpetology and in particular
his work on python conservation, particularly through his publications
on captive breeding of these snakes.
KNOWN
MAXIMUM SIZES OF BROGHAMMERUS GEN. NOV.
Measurements
as quoted in the literature claiming lengths in excess of 33 feet (10 metres)
should be treated with skepticism.
Specimens
over 20 feet (more than 6 metres) are seen from time to time.
On
the internet site: http://www.bobclark.com/d_learn.asp?id=71&cat=pythons
run by Bob Clark is a picture of an adult Broghammerus. The text
on the site, cited here as Clark (2002), reads as follows:
‘Fluffy,
the 310lb. reticulated python.
Fluffy
may be the largest snake in captivity. She’s 12 years old, captive born
and tame. She weighs 310lbs, actual weight, and is somewhat over 22' long.
She continues to grow at a fairly rapid rate gaining about 40lbs in each
of the last two years.’
Taronga
Zoo in Sydney, Australia also claimed specimens in excess of 20 feet (more
than 6 metres) (Chris Williams, personal communication).
The
Pittsburgh Zoo in the United States claimed a specimen in the past at 28
feet.
There
is little doubt that the longest verified living snake was a Broghammerus.
KNOWN
DISTRIBUTION OF BROGHAMMERUS GEN. NOV.
According
to the internet site at:
http://www.nature-conservation.or.id/pythonidae.html
put together
by Ed Colijn the distribution for Broghammerus is listed as including:
India
(including Nicobar Islands north of Sumatra), Bangladesh, Myanmar, Cambodia,
Laos, Vietnam, Thailand, Peninsular Malaysia, Singapore, Weh, Simeulue,
Babi, Nias, Banyak, Mentawai, Riau, Natuna and Anambas Islands, Sumatra,
Enggano, Bangka, Belitung, Krakatau Islands, Kalimantan, Sarawak, Sabah,
Brunei, Java, Nusa Barung, Lombok, Sumbawa, Flores, Alor, Pantar, Lomblen,
Sumba, Timor, Wetar, Leti, Romang, Banda and Tanimbar Islands, Selayar,
Kayadi, Tanah Jampea, Sulawesi, Buton, Sula Islands, Bacan, Ternate, Halmahera,
Obi, Buru, Seram, Ambon, Boano, Haruku, Saparua and Philippines
This information
cited here as Colijn (2002) is believed to be accurate, although obviously
many of the islands within this general ambit that have this genus are
inadvertently omitted.
Similar
information appears in regional herpetology guides (e.g. David and Vogel
(1996)) and is reflected in the databases of 26 Museums in North America
and several others in Europe and South-east Asia.
BROGHAMMERUS
RETICULATUS DALEGIBBONSI SUBSP. NOV.
HOLOTYPE
A specimen
at the Field Museum of Natural History, 1400 S. Lake Shore Drive, Chicago,
IL 60605-2496. The specimen, number: 142320 is from Ambon Island in the
Moluccas in Indonesia, Lat. 3° S, Long. 128° E. It was collected
in 1963 by A.M.R. Wegner.
PARATYPE
A specimen
at the Field Museum of Natural History, 1400 S. Lake Shore Drive, Chicago,
IL 60605-2496. The specimen, number: 142093 is from Ambon Island in the
Moluccas in Indonesia, Lat. 3° S, Long. 128° E. It was collected
in 1963 by A.M.R. Wegner.
DIAGNOSIS
It
appears that this is a generally smaller race of Broghammerus than
the typical race from further west in South-east Asia. Size and colouration
as a trend separate this form from the nominate race reticulatus.
Their
colouration is also often darker than those from further west. It rarely
has a head lighter than the body as in some other variants of Broghammerus,
such as those from Bali or parts of Thailand (see Broghammerus reticulatus
euanedwardsi subsp. nov. below). This race generally has a pugnacious
disposition in captive settings.
This
subspecies is only definitively known from Ambon at this stage, although
it is safe to say that the Broghammerus reticulatus from nearby
Ceram are assignable to this subspecies.
This
subspecies is best separated from all other Broghammerus by DNA
analysis and/or accurate distribution information.
The
subspecies co-exists with Australiasis clastolepis.
ETYMOLOGY
Named
after Australian herpetologist Dale Gibbons for various contributions to
wildlife conservation in the Australian state of Victoria.
BROGHAMMERUS
RETICULATUS EUANEDWARDSI SUBSP. NOV.
HOLOTYPE
A specimen
at the Field Museum of Natural History, 1400 S. Lake Shore Drive, Chicago,
IL 60605-2496. The specimen, number: 180232 is from Nakhon Ratchasima,
Central Thailand. Lat. 14° 58' N, Long. 102° 07' E. It was collected
on 10 August 1969 by W Ronald Heyer.
PARATYPE
A specimen
at the Field Museum of Natural History, 1400 S. Lake Shore Drive, Chicago,
IL 60605-2496. The specimen, number: 178660 is from Khorat, Central Thailand.
Lat. 14° 58' N, Long. 102° 7' E. It was collected in October 1957.
DIAGNOSIS
This
is a large race of Broghammerus reticulatus, with specimens known
to exceed 6 metres. Although it is touted as a yellow-headed and docile
variant, not all specimens of this subspecies have this trait. However
as general trends, these factors separate this subspecies from the nominate
race.
Specimens
are often docile in temperament, especially as adults and make good pets,
provided one makes sure that they don’t handle them after cleaning out
rat or rabbit cages.
This
subspecies is known only from parts of Thailand, but probably occurs elsewhere
including the westernmost parts of the Broghammerus reticulatus
range.
It
is best separated from other Broghammerus reticulatus by either
good locality information and/or DNA analysis.
ETYMOLOGY
Named
in honour of herpetologist Euan Edwards.
BROGHAMMERUS
RETICULATUS HAYDN MACPHIEI SUBSP. NOV.
HOLOTYPE
A specimen
at the Field Museum of Natural History, 1400 S. Lake Shore Drive, Chicago,
IL 60605-2496. The specimen, number: 148968 is from, the Kapit District,
Sarawak, (Borneo), Malaysia. It was collected by F. Wayne King on 9 August
1963.
PARATYPE
A specimen
at the Field Museum of Natural History, 1400 S. Lake Shore Drive, Chicago,
IL 60605-2496. The specimen, number: 67265 is from Sarawak, (Borneo), Malaysia.
It
was collected by Tom Harrisson on 16 Jun 1951.
DIAGNOSIS
This
is a large race of Broghammerus reticulatus, with specimens known
to exceed 6 metres. It is restricted to the Island of Borneo, although
similar specimens have been seen from parts of Sulawesi and may ultimately
be referable to this taxa.
Specimens
are often snappy in temperament, even as adults and do not necessarily
make good captives.
As
a generalization, larger average adult size can be used to separate this
subspecies from "normal" reticulatus.
Yellow-headed
specimens do occur, but are not generally common.
The
subspecies has been separated by some people from other Broghammerus
by it’s iris colour, although this author has failed to find it to be a
reliable indicator.
It
is best separated from other Broghammerus reticulatus by either
good locality information and/or DNA analysis.
ETYMOLOGY
Named
in honour of Victorian (Australia) herpetologist Hayden McPhie for various
contributions to wildlife conservation.
BROGHAMMERUS
RETICULATUS NEILSONNEMANI SUBSP. NOV.
HOLOTYPE
A specimen
at the Field Museum of Natural History, 1400 S. Lake Shore Drive, Chicago,
IL 60605-2496. The specimen, number: 53272 is from Davao Province, Mindanao
Island, the Phillippine Islands. Lat. 7°04' N, Long. 125° 40' E.
It was collected by Donald Heyneman on 27 September 1946.
PARATYPES
A specimen
at the Field Museum of Natural History, 1400 S. Lake Shore Drive, Chicago,
IL 60605-2496. The specimen, number: 53281 is from Davao Province, Mindanao
Island, the Phillippine Islands. Lat. 7°04' N, Long. 125° 40' E.
It was collected by a local Philippine native on 14 January 1947.
A specimen
at the Field Museum of Natural History, 1400 S. Lake Shore Drive, Chicago,
IL 60605-2496. The specimen, number: 53287 is from Davao Province, Mindanao
Island, the Phillippine Islands. Lat. 7°04' N, Long. 125° 40' E.
It was collected by a local Philippine native on 24 November 1946.
A specimen
at the Field Museum of Natural History, 1400 S. Lake Shore Drive, Chicago,
IL 60605-2496. The specimen, number: 53273 is from Davao Province, Mindanao
Island, the Phillippine Islands. Lat. 7°04' N, Long. 125° 40' E.
It was collected by a local Philippine native on 9 October 1946.
A specimen
at the Field Museum of Natural History, 1400 S. Lake Shore Drive, Chicago,
IL 60605-2496. The specimen, number: 53283 is from Davao Province, Mindanao
Island, the Phillippine Islands. Lat. 7°04' N, Long. 125° 40' E.
It was collected by Harry Hoogstraal on 17 January 1947.
DIAGNOSIS
It
appears that this is a large and generally aggressive race Broghammerus.
Quiet and easily tamed specimens are relatively unusual.
It
rarely has a head lighter than the body as in some other variants of Broghammerus,
such as those from Bali or parts of Thailand, although light-headed specimens
are known.
This
subspecies is only definitively known from Mindanao and adjacent Philippine
Islands this stage and is best separated from all other Broghammerus
by comparative DNA analysis and/or accurate distribution information.
ETYMOLOGY
Named
in honour of the long-term reptile breeder, Neil Sonneman, from Murmungie,
in Northern Victoria, Australia.
BROGHAMMERUS
RETICULATUS PATRICKCOUPERI SUBSP. NOV.
HOLOTYPE
A specimen
at the Museum of Comparative Zoology at Harvard, MCZ number: R-25266. It
was collected in 1924 at "Djamplong", South Timor, Lat. 4° S, 125°
E. The person who collected the specimen in 1924 was M. Smith.
DIAGNOSIS
This
is the only Broghammerus found on Timor.
It
is a smaller than average race and is of variable temperament.
Broghammerus
reticulatus patrickcouperi subsp. nov. is usually a brightly coloured
subspecies and this trait alone generally separates the subspecies from
"typical" reticulatus..
The
subspecies is best separated from other Broghammerus reticulatus
subspecies by DNA properties and/or accurate locality information.
It
is a little-known and rarely kept subspecies.
It
co-exists on Timor with Australiasis timorensis.
ETYMOLOGY
Named
after Queensland-based herpetologist Patrick Couper for his contribution
to herpetology.
BROGHAMMERUS
RETICULATUS STUARTBIGMOREI SUBSP. NOV.
HOLOTYPE
A specimen
at the Museum of Comparative Zoology at Harvard, MCZ number: R-8003. It
was collected in 1906 from Buitenzore, Java, Indonesia, Lat. 3°4’S,
Long. 128°12’E. It was collected by T. Barbour in December 1906.
DIAGNOSIS
This
is a subspecies which usually has an exaggerated yellowish hue all over
it’s dorsal surface as compared to other Broghammerus reticulatus.
It
is of variable size (but generally largish) and within the constraints
of being yellowish all over has several distinct colour variations, even
in a single group of young.
It
is usually (but not always) separated from other Broghammerus reticulatus
by the following suite of characteristics: white blotches along the sides
of the body, a relative lack of head markings on a light brown or yellowish
head. The black line seen in most Broghammerus reticulatus that
usually runs from the temple to the eye, usually fails to reach the eye
in this subspecies. They also usually have a relatively light coloured
eye.
Broghammerus
reticulatus stuartbigmorei is readily distinguished from Broghammerus
reticulatus from Sumatra and Borneo, indicating that the population
has been separated for quite some time.
This
same subspecies is believed to occur on the island of Bali.
The
subspecies stuartbigmorei is best separated from others by distribution
and/or DNA properties.
ETYMOLOGY
Named
after Stuart Bigmore of Victoria, Australia for his contributions to herpetology
over two or more decades, in particular varanid taxonomy.
GENUS
CHONDROPYTHON MEYER 1874
Chondropython
viridis (Schlegel 1872) (Aru Islands, Indonesia)
PREAMBLE
TO THE DESCRIPTION OF Chondropython viridis shireenae subsp.
nov.
Hoser
(2000) did not make any new taxonomic arrangements or name changes for
any Green Pythons (Chondropython viridis) including Australian specimens,
for several reasons including pending further research on this species
and sighting of further specimens from both New Guinea, Aru and other Islands
and North East Australia.
Since
Hoser (2000) was published, this author has been fortunate enough to make
cursory and detailed observations of preserved specimens from both New
Guinea and Australia held at both the Queensland Museum (QM) and National
Museum of Victoria (NMV) as well as numerous live specimens in private
collections in Australia.
Furthermore
this author has obtained photos and other data for specimens held in captivity
outside of Australia.
Notwithstanding
this, a major problem for research into this species has been the relative
paucity of specimens from within Australia to compare with specimens from
elsewhere as well as generally unreliable locality data for many of the
specimens seen.
There
has also been an increased interest by other herpetologists in terms of
separating Australian specimens from those from elsewhere.
This
interest has also been shared by the relevant wildlife authorities, in
particular the Queensland National Parks and Wildlife Service (QNPWS) who
have launched at least one major prosecution against a herpetologist who
alleged he had Queensland C. viridis, when the QNPWS in turn alleged
that his snakes derived from stock smuggled to Australia and of origins
outside Queensland.
That
case, involving, Queensland snake breeder Bob Buckley has now been resolved.
The early stages of that case were covered in detail in the book Smuggled-2:
Wildlife Trafficking, Crime and Corruption in Australia (Hoser 1996).
It
was a reluctance to be drawn into the Buckley case as a witness or to prejudice
proceedings and be in "contempt of court" that also made this author decide
to postpone formally naming the Australian subspecies of C. viridis
until after the conclusion of the case.
The
Australian subspecies of C. viridis formally named and recognised
here for the first time has already been recognized by many herpetologists
both within Australia and elsewhere as different and this paper merely
formalizes that arrangement.
It
had originally been hoped to publish a complete revision of the C. viridis
species group with detailed information on all regional variants, however
the scope of this ongoing project is exceptionally large and may take an
indeterminate time frame due to the inherent difficulties in this project
and other competing time and resource demands, so this partial reclassification
of the species group, incorporating Australian specimens only has been
published now in this paper.
One
of the reasons for the earlier publication of this paper is the ongoing
interest by local herpetologists and authorities in maintaining the genetic
purity of Australian specimens in captivity in Australia.
A similar
situation occurred for the other three Australian taxa formally named in
this paper and those descriptions are published for the same reasons. Further
data has confirmed their status as distinct from related forms.
GENUS
CHONDROPYTHON MEYER 1874
This
is the Green Python (the common name applied to the snakes of this genus).
There is only one species within the genus. That is Chondropython viridis.
The
type locality is the Aru Islands, Indonesia, south of New Guinea. Some
recent workers, including Kluge (1993) and Underwood and Stimson (1990)
have made synonymous this genus and Morelia, the latter name taking
precedence. This author did not accept that arrangement in Hoser (2000)
and still does not accept it. While it is clear that the two genera derived
from the same ancestral stock, it is believed that the two have been separated
long enough to warrant being placed in separate genera.
Furthermore,
this author views the placement of Chondropython into Morelia
as an inconsistent move, bearing in mind the widespread splitting off of
other similar genera viz Antaresia and Leiopython, both names
of which were in general usage prior to publication of Hoser (2000).
The
lack of a distinct dorsal pattern of blotches and stripes that typifies
all Morelia (except spilota) or a black and yellow dorsal
pattern as in spilota separates Chondropython from all snakes
in Morelia. There are no iridescent green Morelia. This is
the usual dorsal colouration for adult Chondropython. The absence
of labial pits in Chondropython is frequently cited as a characteristic
that separates the genera Chondropython and Morelia. That
is not so. In fact both genera have distinct labial pits. See the photos
published on page 118 bottom left for M. variegata and page 123
bottom left for C. viridis in Barker and Barker (1994), or photos
published in Hoser (1989) and O’Shea (1996) to view the labial pits in
both genera.
Green
Pythons are separated from all Morelia by their far greater number
of small scales on the dorsal surface of the head, giving the snake an
almost granular appearance. Comparative photos of the heads as shown in
Barker and Barker (1994) and Hoser (1989) readily illustrate this point.
The
Green Pythons are readily distinguished from all other Australian pythons.
Refer to Hoser (1981a) McDowall (1975) and O’Shea (1996) for further diagnostic
information. Australian Green Pythons (as cited by Thomson (1935)) are
more likely than the New Guinea specimens to have markings in a thin line
along the spine to form some sort of vertebral line or pattern.
This
is corroborated by other authors including the photos in Greer (1997).
However the same trait is also seen commonly in south New Guinea and Aru
Island specimens and thus the trait cannot be seen to be consistent in
terms of identification. Also see the subspecies description below.
Specimens
from the north of New Guinea are likely to have spots in a more irregular
pattern.
Specimens
from around the high country of Wamena in Irian Jaya are often a very dark
green with buttercup yellow spots on the back. The dark yellow ventral
scales are commonly a grey/black in colour. As with Morelia, Chondropython
is a species with considerable variation in colour, not only between locations,
but even within a single location and even within a single litter of young.
Numerous
other colour variants are known, including blue adults and "mite phase"
which as adults are green with lots of black flecks as well as sometimes
having other markings such as scattered white scales. One such animal ("Mite
phase") was depicted in a post by Scott (?) on March 24, 2002 at 19:22:14:
at: http://www.kingsnake.com/forum/gtpython/messages/22444.html on http://www.kingsnake.com.
Photos
of Australian Green Pythons in life with exact locality data are shown
in Barker and Barker (1994) and Greer (1997) and other publications.
Photos
of New Guinea Green Pythons in life with exact locality data are shown
by O’Shea (1996) and other publications.
Chondropython
viridis shireenae
subsp. nov.
HOLOTYPE
An
adult specimen (number D51862) held at the National Museum of Victoria
(NMV), from "Cape York, Queensland" (The general Lat. Long. for Cape York
is: 15°00' S, 143°00' E). The snake was not accurately measured
due to the fact it was hardened in tight coils, however it was about 1
metre in total length.
Dorsally
the snake was dark in colour (relative for the species C. viridis)
and a generally uninterrupted colour dorsally, save for a thin line of
lighter scales along the dorsal vertebra, having been a generally dark
green in life.
Ventrally
the holotype of Chondropython viridis shireenae subsp. nov. was
light in colour, (having been a yellow colour in life) and with some dark
flecks on some scales (probably grey, black or green flecks in life).
The
sex of the holotype of Chondropython viridis shireenae subsp. nov.
was not ascertained.
There
were six pits in the lower labials and three on the upper scales of the
mouth, towards the front of the head (these counts being for each side
of the head).
The
holotype of Chondropython viridis shireenae subsp. nov. had 233
ventrals, divided anal and 65 (all paired) subcaudals.
Most
of the scales on the dorsal surface of the head were small and almost granular
in appearance and somewhat irregular.
The
teeth were long and sharp.
The
holotype of Chondropython viridis shireenae subsp. nov. had been
in captivity prior to being lodged at the museum as evidenced by the one
or more domestic mice (Mus musculus) found inside the stomach of
the snake (undigested).
Based
on the fact that the snake was collected on 20 June 1973, the age of the
specimen certainly predates the period when numbers of the species were
smuggled into Australia from the USA and Europe having themselves been
sourced from snakes derived from New Guinea/Irian Jaya stock.
DIAGNOSIS
Chondropython
viridis shireenae
subsp. nov. are the only Green Pythons (C. viridis) found
on mainland Australia and can be separated from all other C. viridis
on this basis.
In
the absence of good locality data, the subspecies is best separated from
other C. viridis by comparative DNA analysis, which has already
been successfully used to separate this subspecies.
Contrary
to statements by many Australian and overseas herpetologists that all Australian
C. viridis have white or other markings along the vertebra, whereas
those from elsewhere do not, claiming this to be diagnostic for Australian
specimens of C. viridis, this is not the case. Notwithstanding this,
it is fair to say that a very thin line or line of dots along the spine
is generally a diagnostic trait for adults of this subspecies.
This
author has seen C. viridis from Australia both with and without
such markings (e.g. several captives seen from Cape York which lack very
distinctive vertebral markings and those from Iron Range National Park
on page 115, top, of Kend (1997) or another specimen from the same area
as depicted in Barker and Barker (1994) which have such markings) and likewise
for those from New Guinea (e.g. NMV specimens D9747 and D9748 both young
specimens from "Uinea" PNG or the juvenile D54438 from Wuroi, PNG, all
of which lack vertebral markings and a small adult specimen from PNG at
the Queensland Museum (QM J35376) with very thick and distinct white vertebral
markings).
As
a trend, vertebral markings decline with age. General dorsal markings,
sometimes in the form of blotches and mid-dorsal markings are generally
far more common in juvenile C. viridis of all subspecies.
The
range of C. viridis shireenae subsp. nov. is only in very
wet habitats of the lowlands and nearby range areas on the east side of
Cape York from about the MacMillan River drainage in the north to the area
around the Normanby River drainage in the south. This includes the Sir
William Thompson and McIlwraith Ranges both in the general vicinity of
the Iron Range National Park.
The
known distribution of C. viridis in Australia is only on very restricted
parts of the Cape York Peninsula and does not include the dry savannah
habitats that adjoin Torres Strait or the islands within Torres Strait
itself.
There
is believed to be over 50 km (straight line measurement) between the northernmost
C. viridis shireenae subsp. nov. and the northern tip of Cape
York, bordering the Torres Strait, which includes generally unsuitable
habitat for the subspecies.
In
view of the lack of evidence of these snakes inhabiting this area within
recent historical or recent prehistorical times, it is reasonable to infer
that the Australian C. viridis have been separated from other populations
for many thousands of years.
Furthermore,
in view of the fact that other species of python inhabit these intervening
areas and presumably compete with this species in the areas they coexist,
it is reasonable to infer that there is not, nor has been any gene flow
between the Australian and other populations of this species within recent
historical or recent prehistorical times and perhaps as far back as or
before Aboriginal settlement of Australia an estimated 40,000 years ago.
The
40,000 year date is significant as it is thought that habitats throughout
much of Australia (including the north) may have changed significantly
(become generally more open and drier) with the arrival of humans, due
to the increased incidence of bushfires. Before the arrival of humans on
the Australian continent, habitat may have allowed for gene flow between
the Australian C. viridis and those populations to the north.
A search
of likely habitats in the far north of Cape York and Torres Strait should
be undertaken to confirm current distribution data.
CAPTIVE
HUSBANDRY
From
a keeper’s point of view there appears to be no known differences in terms
of keeping C. viridis shireenae subsp. nov. and C. viridis
from elsewhere.
These
snakes require an enclosure with at least one horizontal tree branch or
fork in which to perch and they need a humid environment when sloughing.
Eggs
usually take from 38-60 days to hatch (extremes given) (Barker and Barker
1994).
Detailed
husbandry information for this species can be found in Barker and Barker
(1994), Ross (1978) and Ross and Marzec (1990).
Papers
on breeding the species have been published by numerous authors including
definitive papers by Murdoch (1999), Rundquist (1993), Walsh (1979) and
Zulich (1990). Copies of the papers by Murdoch, Walsh and Zulich are available
in full on the internet and can be found using any decent search engine,
such as "www.yahoo.com".
ETYMOLOGY
Chondropython
viridis shireenae
subsp. nov. is named in honour of my long suffering wife.
Shireen
Vanessa Hoser has had to put up with long periods without me as I have
conducted the research necessary for this and other publications and done
an excellent job in terms of looking after and caering for our children
in my absence.
GENUS
HELIONOMUS GRAY 1842
Helionomus
sebae (Gmelin 1789) (Africa, mainly Sth of Sahara, excl. Sthn Africa))
Helionomus
natalensis (Smith 1840) (Sth Africa and adjacent areas)
GENUS
KATRINUS HOSER 2000
Katrinus
fuscus (Peters 1873) (NE Australia)
Katrinus
fuscus cornwallisius (Gunther, 1879) (Southern New Guinea)
KATRINUS FUSCUS JACKYAE SUBSP. NOV.
HOLOTYPE
A specimen
in the Western Australian Museum number 13882 from Kalumburu WA, Lat. 14°18'
S, Long 126°39' E. This is a smooth-scaled Python.
PARATYPE
A specimen
in the Western Australian Museum number 42796 from Kalumburu WA, Lat. 14°18'
S, Long 126°39' E. This is a smooth-scaled Python.
DIAGNOSIS
Katrinus
fuscus jackyae is readily identified by the following suite of characters:
It is a medium to large python, averaging 2 metres in length, with occasional
specimens attaining up to nearly three metres. It is an olive greyish green
dorsally with no discernable pattern. Ventrally the snake is usually a
bright yellow in colour, visible when the snake is viewed side-on, but
this may range from cream or occasionally orangeish. The colour is most
intense at the anterior part of the body. The scales are smooth and shiny.
The
snakes have large teeth on the premaxilla. The head is covered by large
symmetrical shields and there are pits in some of the labial scales. Katrinus
fuscus jackyae, like others in the genus are separated from Antaresia
by having a single loreal rather than two or more. Katrinus fuscus
jackyae, like others in the genus are separated from Leiopython
by having two pairs of prefrontals as opposed to having a pair. Katrinus
fuscus jackyae, like others in the genus are separated from Liasis
by usually having 55 or less mid-body rows (Liasis usually has over
60) as well as usually having a more intense colour than Liasis.
Katrinus
fuscus jackyae and Liasis olivaceous are sympatric at the
type locality (Kalumburu, WA).
Katrinus
fuscus jackyae is separated from Katrinus fuscus fuscus
several characteristics including the upper lips.
In
Katrinus fuscus fuscus (from coastal Queensland) the upper lips
are pale with a little brown peppering. However in K. fuscus jackyae
(from the NT and WA) the lips are usually darker with more dark brown peppering
or even blotches.
The
two subspecies intergrade in the region of the Gulf of Carpentaria.
This
subspecies (Katrinus fuscus jackyae) is herein restricted
to the region encompassed by the Kimberley Ranges of Western Australia,
the Northern Territory and adjacent areas.
The
subspecies Katrinus fuscus fuscus is herein restricted to the east
coast of Queensland and adjacent areas.
The
subspecies Katrinus fuscus cornwallisius is restricted to the landmass
of New Guinea and immediately adjacent islands.
Katrinus
fuscus jackyae, like others in the genus are invariably associated
with watercourses and are commonly known as ‘Water Pythons’. The population
at Fogg Dam in the Northern Territory is particularly large and has been
well-studied by students and academics at the University of Sydney, under
the guidance of Dr. Richard Shine.
Shine
and his team of researchers found Katrinus fuscus jackyae to
be one of the major predators on the river floodplains in northern
Australia. They fed mainly on small mammals, in particular native Dusky
Rats (Rathus colletti), as well as bandicoots, flying foxes, and
other vertebrates.
The
movements of the snakes did in part correlate with that of their primary
food source/s.
In
the wild state, the snakes may be either ambush predators or active hunters.
In
the Northern Territory mating takes place in June-August, egg-laying (usually
9-16 eggs) in August-September and hatching in November-December.
Nesting
occurs in abandoned burrows and among the roots of paperbark trees. For
reasons not completely known a large number of clutches are laid in unsuitable
places and therefore fail to hatch.
Queensland
populations of Katrinus fuscus fuscus have a different lifestyle
in terms of foods eaten and breeding activity in that mating and egg laying
occurs an average 8 or so weeks later than their top-end counterparts.
Anecdotal
reports within Australia suggest that Queensland Katrinus are more
placid (less snappy) than those from the NT and WA.
However
this author’s experiences suggest that the allegedly snappy nature that
these snakes have, is often over-rated. Most individuals may be snappy
in cages when they expect food, but usually become reasonably docile when
handled. Within this profile, there are some obvious exceptions.
When
photographing Katrinus (from any location) this author has found
the snakes to be placid and after some cajoling, they tend to stay put.
The only time they try to snap is usually when the author has forced them
to crawl over open areas in order to tire them out, whereupon the snakes
may turn and hold their ground (sometimes snapping). This is usually when
the snakes are most easily moved to their ‘stage’ and made to sit in an
appropriate photographic position. In captivity these snakes are hardy,
breed readily and usually present few husbandry problems.
ETYMOLOGY
Named
after Jacky Hoser, this author’s second daughter.
Katrinus
mackloti (Dumeril and Bibron 1844) (Lesser Sunda Islands, Indonesia)
Katrinus
mackloti dunni (Stull 1932) (Wetar, Indonesia)
Katrinus
savuensis (Brongersma 1956) (Sawu Island)
GENUS
LEIOPYTHON HUBRECHT 1879
Leiopython
albertisi (Gray 1842) (Eastern Irian Jaya)
Leiopython
albertisi barkeri Hoser 2000 (Mussau, PNG)
Leiopython
albertisi bennetti Hoser 2000 (NE PNG)
Leiopython
hoserae Hoser 2000 (Southern New Guinea and Southern Irian Jaya)
GENUS
LENHOSERUS HOSER 2000
Lenhoserus
boeleni (Brongersma 1953) (New Guinea and Irian Jaya)
GENUS
LIASIS GRAY 1840
Liasis
olivaceus (Gray 1842) (Nth Australia)
Liasis
olivaceus barroni (Smith, 1981) (Pilbara, West Australia)
Liasis
papuana (Peters and Doria 1878) (New Guinea and Irian Jaya)
GENUS
MORELIA GRAY 1842.
Morelia
bredli (Gow 1981) (Central Australia)
Morelia
carinata (Smith 1981) (Kimberleys North West Australia)
Morelia
cheynei Wells and Wellington 1983 (NE Queensland, Australia)
MORELIA
MIPPUGHAE SP. NOV.
HOLOTYPE
A specimen
at the South Australian Museum (SAM), Adelaide, SA, Specimen number: R1665
from Moolooloo, North Flinders Ranges, South Australia, Lat: 30°59’
Long: 138°35’.
PARATYPE
A specimen
at the South Australian Museum (SAM), Adelaide, SA, Specimen number: R14261
from Iron Dutchess, Middleback Ranges, South Australia, Lat: 33°15’
Long: 137°07’.
DIAGNOSIS
A medium
to large python similar in most respects to the others in the genus Morelia.
It is separated from one of its closest relatives Morelia macburniei
sp. nov. (see below) by a suite of characteristics including a lower
incidence of scale anomalies particularly with regards to ventral scales
in the from of longitudinally split ventrals, half ventrals, transversely
divided ventrals or incompletely transversely divided ventrals, remnant
or partially inserted ventrals or incompletely formed ventrals (such as
in two halves).
This
species is differentiated from Morelia macburniei sp. nov. from
St. Francis Island by having more rhomboidal-shaped dorsal scales as opposed
to having lanceolate-shaped dorsal scales.
Morelia
mippughae sp. nov. is separated from the closely related Morelia
metcalfei (the Murray/Darling form) by a suite of characteristics including
it’s dorsal colour pattern.
Morelia
mippughae sp. nov. is pale reddish brown dorsally, with broad transverse
black-edged patches on the top of the back and a wide pale lateral zone
for about a third of its length.
Morelia
mippughae sp. nov. is restricted to the Flinders and Middleback Ranges
areas of South Australia. No other Morelia occurs here.
Numbers
of this species have declined sharply since European settlement, (Ian Renton
and Ted Mertens personal communications).
This
author hereby calls for specimens of Morelia mippughae sp. nov.
to be taken into captivity and bred in numbers in order to secure the survival
of this taxa.
ETYMOLOGY
Named
in honour of Mrs Mip Pugh of Breakwater, Victoria for her long term contributions
to herpetology. She is part of the husband and wife team, the other half
being Mick Pugh (whom this species is not named in honour). Mip has over
the last few decades given free of charge many hundreds of hours of useful
advice and guidance to reptile keepers in Victoria, especially in relation
to her favorite reptiles, which are lizard species such as Bearded Dragons
(Pogona spp.) and other commonly kept species. Her house has often
been a defacto hotel suite for countless other herpetologists who have
enjoyed her hospitality.
Morelia
harrisoni Hoser 2000 (Southern New Guinea and Southern Irian Jaya)
Morelia
imbricata (Smith 1981) (South West Australia)
MORELIA
MACBURNIEI SP. NOV.
HOLOTYPE
A specimen
at the South Australian Museum (SAM), Adelaide, SA, Specimen number: R13994
from St. Francis Island, in the Nuyts Archipelago near Ceduna in South
Australia, Lat: 32°31’ Long: 133°18’.
PARATYPE
A specimen
at the South Australian Museum (SAM), Adelaide, SA, Specimen number: R19072
from St. Francis Island, in the Nuyts Archipelago near Ceduna in South
Australia, Lat: 32°31’ Long: 133°18’.
DIAGNOSIS
A medium
to large python similar in most respects to the others in the genus Morelia.
It is separated from its closest relative Morelia imbricata by a
suite of characteristics including a higher incidence of scale anomalies
particularly with regards to ventral scales in the from of longitudinally
split ventrals, half ventrals, transversely divided ventrals or incompletely
transversely divided ventrals, remnant or partially inserted ventrals or
incompletely formed ventrals (such as in two halves).
This
species is differentiated from Morelia from the South Australian
mainland by having lanceolate-shaped dorsal scales as opposed to more rhomboidal-shaped
dorsal scales.
This
same characteristic also separates Morelia imbricata from other
southern Australian Morelia.
Morelia
macburniei sp. nov. is separated from Morelia imbricata by distribution
(believed to be several hundred kilometers).
While
Morelia macburniei is highly variable in individual colouration
and pattern, the colouration of the species tends to look more like Morelia
from the South Australian mainland as opposed to Morelia imbricata,
even though Morelia macburniei’s dorsal scales are more like those
of M. imbricata.
This
species cannot be definitively separated from other Morelia on the
basis of scalation alone as these properties (ventral counts and the like)
may overlap with other Morelia.
Morelia
macburniei sp. nov. is separated from all other Morelia by distribution.
It is the only species to occur on St. Francis Island.
It
is assumed that the total population for this species is less than 1,000
individual specimens, subjected to seasonal variations. At present there
are no known threats to the species, but because it is a small island population,
it must be regarded as potentially vulnerable, particularly if a feral
species becomes established on the Island.
This
author hereby calls for specimens of Morelia macburniei sp. nov.
to be taken into captivity and bred in numbers in order to secure the survival
of this taxa.
ETYMOLOGY
Named
in honour of Victorian herpetologist Cameron McBurnie for services to herpetology,
including through his role with the Victorian Association for Amateur Herpetologists
(VAAH).
Morelia
macdowelli Wells and Wellington 1983 (Eastern Australia, along coast)
Morelia
metcalfei Wells and Wellington 1985 (Inland Eastern Australia)
Morelia
spilota (Lacepede 1804) (Coastal NSW and nearby areas, Australia)
Morelia
variegata (Gray 1824) (Northern Australia)
Genus
Nyctophilopython Wells and Wellington 1985
Nyctophilopython
oenpelliensis (Gow 1977) (Arnhem Land Escarpment, Australia)
GENUS
PYTHON DAUDIN 1803
Python
molurus (Linnaeus 1758) (Indian subcontinent)
Python
molurus bivittatus Kuhl 1820 (Indochina to Indonesia)
Python
molurus pimbura Deraniyagala 1945 (Ceylon)
GENUS
SHIREENHOSERUS GEN. NOV.
TYPE
SPECIES: PYTHON ANCHIETAE, BOCAGE 1887
DIAGNOSIS
A group
of relatively small (under 2 metres) pythons restricted to continental
Africa.
They
are separated from all other African pythoninae (namely Helionomus sebae
(Gmelin 1789)) by a vast suite of characteristics that more than adequately
separate the two genera.
This
includes their smaller adult size (generally well under 2000 mm (with over
1500 mm being unusual) versus an average of 3600-4500 mm in Helionomus
sebae).
The
two genera can also be separated by their totally different dorsal colouration,
which is best seen from a perusal of photos of the relevant species, including
as seen in Stafford (1986).
In
Helionomus sebae (Gmelin 1789) the head colouration is characterized
by a dark arrowhead blotch on the top, bordered on either side by a pale
stripe. This paler stripe which runs through the top of the eye, is more
or less straight and continuous and if there is a break, it is well posterior
to the eye itself. If the pale arrowhead blotch has a dark separation at
the snout it is only relatively narrow.
By
contrast in Shireenhoserus, there is also a dark dorsal surface
of the head bordered by a paler stripe running across the top of the eye.
However instead the line gives the appearance of two or more linear blotches
that tend to form a line, rather than as a single line as in Helionomus
sebae.
In
Shireenhoserus the lighter blotches do not meet at the snout; there
is a distinct and wide gap and the gap between the lighter markings on
each side is far wider than ever seen in Helionomus sebae.
Also
in Shireenhoserus this line of blotches tends to have a break above
the center of the eye, the break often being the topmost supracilliary
scale, which is instead the brownish or blackish colour seen on the flat
dorsal area of the head. No such break is seen in Helionomus sebae.
In
Shireenhoserus the mid-body scale row count ranges from 53-63, while
in Helionomus sebae the mid-body row scale count is 71-83. The ventral
count for Shireenhoserus ranges from 191-267 (versus 265-286 in
Helionomus sebae). The subcaudal count for Shireenhoserus ranges
from 28-57 (versus 60-80 in Helionomus sebae).
In
Shireenhoserus the sensory pits are only present on the upper lip
for the first 4-5 upper labials, versus just two in Helionomus sebae.
For
the lower labials, if sensory pits are present in Shireenhoserus,
they are only very slight, as opposed to the small, but distinct pits on
the lower labials in Helionomus sebae.
Their
breeding biology is also markedly different, the most obvious difference
being the number of eggs laid by the female. In Shireenhoserus the
number of eggs laid is generally well under ten (with a rare record of
11 for Shireenhoserus regia, in de Vosjoli, et. al. (1994)); in
the genus Helionomus the number is generally well above ten.
Furthermore
the relative size of the eggs laid is proportionately larger in Shireenhoserus
than Helionomus sebae, with egg sizes for all species being similar
in spite of the much smaller adult sizes of Shireenhoserus.
Shireenhoserus
cannot be confused with Asiatic or Australasian pythons, or West African
Calabaria.
The
genus Shireenhoserus comprises two known species, namely the Angola
Python (Shireenhoserus anchietae) which is the type species and
the Ball Python (Shireenhoserus regia).
These
are separated from one another by a whole suite of characters, including
of course distribution (refer to Pitman (1974) and other relevant regional
texts for details of distribution).
They
can also be readily separated by the fact that dorsally between the eyes
in Shireenhoserus anchietae the scales tend to be small and irregular,
while in Shireenhoserus regia they tend to be large and distinct
shields.
Shireenhoserus
anchietae usually has from 263-267 ventrals versus the much lower 191-207
in Shireenhoserus regia. Shireenhoserus anchietae has 45-57 subcaudals
versus the much lower 28-47 seen in Shireenhoserus regia.
The
two species can also be separated by their markedly different dorsal colour
patterns, which can be seen best in comparative photos of the species,
including the pair shown on page 47 of Stafford (1996).
Both
species typically have a dorsal pattern consisting of alternating dark
and light brown patches. However in regia the light patches are
relatively large and cross-body and give the appearance of having smooth
edges.
By
contrast in anchietae, the light patches are much smaller as in
smallish blotches and the edges give a jagged appearance. (Also see Pitman
1974 and other regional texts for further differences between the two species).
Further
diagnostic information for the genus Shireenhoserus can be readily
gleaned from the literature as cited at the end of this paper.
ETYMOLOGY
Named
in honour of the author’s wife, Shireen Hoser, who coincidentally is a
native of Africa, which is where the genus occurs.
Shireenhoserus
regia (Shaw 1802) (Western sub-Sahara Africa)
FINAL
CONCLUSIONS
It
is anticipated that in spite of expected resistance from some quarters,
including the campaigns of lies and misinformation likely to be peddled
by persons adversely named in this author’s corruption exposing texts Hoser
(1993), Hoser (1996) and Hoser (2001), the classification adopted in this
paper and Hoser (2000) will become widely used in the future as it presents
the only viable alternative to the patently untenable lumping of all pythoninae
into just three large and badly composed genera as per Underwood and Stimson
(1990) and others.
By
way of example, Hummell’s placement of the species timorensis somewhere
between reticulatus and amethistina (as per Hoser (1982)
and Hoser (2000)), as a result of his 2001 analysis of sequences for python
cytochrome b from 17 different species when combined with the results
of Harvey, Barker, Ammerman and Chippendale. (2000) vindicates the erection
by Wells and Wellington of the genus Australiasis to accommodate
these species as indicated by Hoser (2000).
By
way of further example, Hoser (2000) took the generally unpopular step
of making Kluge’s genus Apodora synonymous with Liasis. This
was at variance with other recent classifications by other authors (e.g.
O’Shea 1996) who had also followed Kluge. However Hummell’s 2001 analysis
of sequences for python cytochrome b from 17 different species, showed:
‘The
Apodora (Liasis) paupuana on the other hand was in
a clade with other Liasis, namely Liasis olivacea, and therefore
may not be a distinct genus separate from Liasis.’
Hummell
made further similar comments indicating his similar views against Apodora
and raising the possibility that Kluge’s results may have in fact been
inadvertently flawed.
This
is as Hoser (2000) had also found a year earlier, when relying on the obvious
morphological and biological data that was widely available and cited in
the paper.
Hummell
also made the valid point of noting how recent DNA studies had yielded
wildly varying results and thought that DNA gathering procedures and within
species variance may have been the reason.
Either
way, this fact means that the push by some herpetologists to rely solely
on DNA analysis based on just a limited number of substances and tests
to decide inter-relationships between species and genera should be avoided
if this is to mean forsaking more traditional (and to date, generally reliable)
methods involving analysis of morphological, biological, behavioral and
other traits.
Finally,
and with regards to the species and subspecies levels, this paper provides
the only recent and consistent approach of the same principals across the
whole Pythoninae, and hence the names used here (as those available under
the ICZN’s code) should be those used in the immediate future, unless and
until compelling evidence to the contrary arises.
ACKNOWLEDGMENTS
Numerous
people assisted over the previous three decades with the provision of live
and dead specimens for study, access to and copies of relevant published
papers and unpublished data.
Included
here were the author’s immediate family (parents and wife), various employers
who allowed the author leave to seek information and the countless herpetologists,
curators, etc., all of whom gave the author unfettered access to both specimens
and data.
Thanks
are also due to the many people who inspected specimens on this author’s
behalf and supplied relevant data.
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Hard
copy originally published in Nov 2003/June 2004 in two parts.
Placed
online a week after publication at:
http://www.smuggled.com/pytrev2.htm
PDF Version of the original paper scanned from the journal.
Placed
online a week after publication at:
http://www.smuggled.com/pdf1/PytRev3.pdf