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Reticulated python (Malayopython reticulatus) is a widely distributed snake covering throughout Southeast Asia and almost all of Indonesia archipelago and divided into several subspecies based on morphological variation and its locality. Morphological variation data of M. reticulatus from Indonesia population has never been done thoroughly. This study aims to determine the morphological variations based on 21 meristic and 3 morphometric characters from several populations in Indonesia. The data was collected from the Museum Zoologicum Bo-goriense (MZB) Cibinong, Indonesia and other additional collections that are carried out from June to July 2018 and then analyzed by Principal Component Analysis (PCA) to determine the population grouping. The result showed a high variation on the scale range of anterior prefrontal, posterior prefrontal, frontal, parie-tal, preocular, postocular, loreal, temporal, upper labial, lower labial, and dorsal scales. Ventral and subcaudal scales in male and female specimens show high variation in the total scale count, the ratio comparison of tl: SVL measurement, indicated the sexual dimorphism. Prefrontal (anterior-posterior) and frontal scales show high variation and difficult to distinguish between each locality at the sub-species level and suggest it to be intra-specific variation. There is no significant grouping were found between populations from data on morphological variations.
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JOURNAL OF TROPICAL LIFE SCIENCE
2019, Vol. 9, No. 3, 259 266
http://dx.doi.org/10.11594/jtls.09.03.07
How to cite:
Septiadi L, Fathoni M, Hanifa BF, Hamidy A (2019) Morphological Variation of Malayopython reticulatus (Schneider, 1801)
from Several Population in Indonesia 9 (3): 259 266. doi: 10.11594/jtls.09.03.07
Research Article
Morphological Variation of Malayopython reticulatus (Schneider, 1801) from
Several Population in Indonesia
Luhur Septiadi 1, M. Fathoni 2, Berry Fakhry Hanifa 1, Amir Hamidy 3
1 Department of Biology, State Islamic University of Maulana Malik Ibrahim, Malang 65145, Indonesia
2 Department of Biology, Faculty Mathematics and Natural Sciences, Brawijaya University, Malang 65145,
Indonesia
3 Laboratory of Herpetology, Zoology Division, Research Center for Biology, Indonesian Institute of Sciences,
Cibinong 16912, Indonesia
Article history:
Submission April 2019
Revised August 2019
Accepted August 2019
ABSTRACT
Reticulated python (Malayopython reticulatus) is a widely distributed snake cov-
ering throughout Southeast Asia and almost all of Indonesia archipelago and di-
vided into several subspecies based on morphological variation and its locality.
Morphological variation data of M. reticulatus from Indonesia population has
never been done thoroughly. This study aims to determine the morphological var-
iations based on 21 meristic and 3 morphometric characters from several popula-
tions in Indonesia. The data was collected from the Museum Zoologicum Bo-
goriense (MZB) Cibinong, Indonesia and other additional collections that are car-
ried out from June to July 2018 and then analyzed by Principal Component Anal-
ysis (PCA) to determine the population grouping. The result showed a high vari-
ation on the scale range of anterior prefrontal, posterior prefrontal, frontal, parie-
tal, preocular, postocular, loreal, temporal, upper labial, lower labial, and dorsal
scales. Ventral and subcaudal scales in male and female specimens show high var-
iation in the total scale count, the ratio comparison of tl: SVL measurement, indi-
cated the sexual dimorphism. Prefrontal (anterior-posterior) and frontal scales
show high variation and difficult to distinguish between each locality at the sub-
species level and suggest it to be intra-specific variation. There is no significant
grouping were found between populations from data on morphological variations.
Keywords: Malayopython reticulatus, morphological variation, intra-specific
variation, Indonesia
*Corresponding author:
E-mail: luhur.septiadi@gmail.com
Introduction
Reticulated python (Malayopython reticula-
tus, Schneider, 1801) is a large constrictor snake
of Pythonidae family that’s widely distributed
throughout Southeast Asia and almost all of Indo-
nesia archipelago [1]. The widespread of this spe-
cies give rises to morphological and genetic varia-
tions given by many factors such as population
isolation, geographical barrier, evolutionary pro-
cess and also the influence of the environment and
its habitat. The variation is then used as a compar-
ing reference (diagnostic character) for distin-
guishing a species [2]. Morphological variations
are observed based on phenetic characters includ-
ing color patterns, specific characteristics of spe-
cies, patterns, and body size ratios, and meristical
characters such as scale ranges [3]. These varia-
tions are further analyzed, to determine the exist-
ence of significant differences between species or
sub-species.
M. reticulatus is divided into 3 subspecies
based on its locality, namely M. reticulatus retic-
ulatus (Greater Sunda), M. reticulatus saputrai
(Selayar Island), and M. reticulatus jampeanus
(Tanahjampea Island). Two subspecies recently
recognized (M. reticulatus saputrai and M. reticu-
latus jampeanus) by having a few specific mor-
phological variations that only subspecies posse-
L Septiadi, M Fathoni, BF Hanifa, A Hamidy, 2019 / Morphological Variation of Malayopython reticulatus
JTLS | Journal of Tropical Life Science 260 Volume 9 | Number 3 | September | 2019
ssed from certain locality [4]. These differences
are based on the shape of the anterior-posterior
prefrontal scales, yet are still confused to inclu-
sively categorizes it as geographical or intraspe-
cific variations, also as diagnostic characters.
Thus, examining more specimens of M. reticula-
tus from several population in Indonesia is needed.
Therefore, we present an overview of morpho-
logical characters of M. reticulatus to address sev-
eral problems, to wit: 1) the tendency of other
morphological characters from several of M. retic-
ulatus populations, 2) to verifying the diagnostic
characters of M. reticulatus especially on subspe-
cies level, 3) the grouping of M. reticulatus popu-
lation in Indonesia.
Material and Methods
Specimens observation
All of 38 specimens of M. reticulatus from
several populations in Indonesia were obtained
from the collection of the Museum Zoologicum
Bogoriense (MZB) that are observed during July-
September 2018 and few additional specimens
that are collected on July - Agustus 2013 and April
- May 2014 (Table 1). Morphological variations
were observed by meristic and morphometric
Table 1. M. reticulatus specimens observed from several populations in Indonesia, collection of Museum
Zoologicum Bogoriense (MZB) LIPI and several additional specimens
No.
ID. Number
Locality
1.
MZB. Ophi. 5287
Dalas, Southern Lampung
2.
MZB. Ophi. 201
F. Krakatau Island, Lampung
3.
MZB. Ophi. 4680
F. Krakatau Island, Lampung
4.
MZB. Ophi. 2412
Saraya, Southeast Aceh
5.
MZB. Ophi. 1692
Tiwi River, Kota Tua, Riau
6.
MZB. Ophi. 2192
Bengkalis, Riau
7.
MZB. Ophi. 2936
Pangandaran, West Java
8.
MZB. Ophi. 197
Cikopo, West Java
9.
MZB. Ophi. 4681
Cikopo, West Java
10.
MZB. Ophi. 5286
Malang, East Java
11.
MZB. Ophi. 2152
Hilir Village, Southern Barito, Central Kalimantan
12.
MZB. Ophi. 1906
Pontianak, West Kalimantan
13.
MZB. Ophi. 1457
Banjarmasin, South Kalimantan
14.
MZB. Ophi. 5884
Lumbis River, Nunukan, North Kalimantan
15.
MZB. Ophi. 3020
TNKM Kragan, Nunukan, East Kalimantan
16.
MZB. Ophi. 3121
Marang, Sangkulirang, East Kalimantan
17.
MZB. Ophi. 5785
Kawinda Toi, Sumbawa Island
18.
MZB. Ophi. 5589
Saunulu, Central Maluku
19.
MZB. Ophi. 1951
Saunulu, Central Maluku
20.
MZB. Ophi. 4481
North Halmahera, North Maluku
21.
MZB. Ophi. 4482
North Halmahera, North Maluku
22.
MZB. Ophi. 194
West Seram Island, Maluku
23.
MZB. Ophi. 200
Timor Island, Maluku
24.
MZB. Ophi. 1558
Anca Danau Lindu, Central Sulawesi
25.
MZB. Ophi. 3221
Dampala River, Morowali, Central Sulawesi
26.
MZB. Ophi. 2895
Goa Keramat, Tawaeli, Donggala, Central Sulawesi
27.
MZB. Ophi. 1762
Tomodo, Lindu Valley, Central Sulawesi
28.
MZB. Ophi. 2297
Marowo, Tojo Una-Una, Central Sulawesi
29.
MZB. Ophi. 1657
Jompi, Raha, Muna Island, Southeast Sulawesi
30.
MZB. Ophi. 4682
Jompi, Raha, Muna Island, Southeast Sulawesi
31.
MZB. Ophi. 2041
Latimojong Mountain, Luwu, South Sulawesi
32.
MZB. Ophi. 3436
Selayar Island, Southern Sulawesi Arhipelago
33.
MZB. Ophi. 3223
Tanahjampea Island, Southern Sulawesi Archipelago
34.
MZB. Ophi. 3437
Kalaotoa Island, Southern Sulawesi Archipelago
35.
S.Voucher. 020
Karompa Island, Southern Sulawesi Archipelago
36.
S.Voucher. 021
Kalaotoa Island, Southern Sulawesi Archipelago
37.
S.Voucher. 022
Kalaotoa Island, Southern Sulawesi Archipelago
38.
S.Voucher. 023
Madu Island, Southern Sulawesi Archipelago
L Septiadi, M Fathoni, BF Hanifa, A Hamidy, 2019 / Morphological Variation of Malayopython reticulatus
JTLS | Journal of Tropical Life Science 261 Volume 9 | Number 3 | September | 2019
characters.
Measurement of meristic, morphometric charac-
ters, and analysis
Morphometric characters are observed by 3
characters, as follows: a total of length (ToL), tail
length (tl), and snout-vent length (SVL). Then, 21
meristic characters observed by scales counting, as
follows: rostral (r), internasal (i), anterior prefron-
tal (apf), posterior prefrontal (ppf), frontal (f), su-
praocular (so), parietal (p), interparietal (ip), pre-
ocular (pro), postocular (pso), loreal (l), temporal
(t), upper labial (la), upper labial contacting orbit
(lao), lower labial (la'), anterior upper thermal pits
(tp), posterior lower thermal pits (tp'), mental (m),
dorsal midbody scales (d), ventral (v), and sub-
caudal (sc) (Figure 1), the characters used are ac-
cording to Auliya [4] with standard measurement
and scale counting following Lilywhite & Wray
[5]. Male and female specimens were differenti-
Figure 3. Meristical character on head scalation of M. reticulatus specimens that are observed. Abbrevation as
follows: rostral (r), internal (i), anterior prefrontal (apf), posterior prefrontal (ppf), frontal (f), supraoc-
ular (so), parietal (p), interparietal (ip), preocular (pro), postocular (pso), loreal (l), temporal (t), upper
labial (la), upper labial contacting orbit (lao), lower labial (la'), anterior upper thermal pits (tp), poste-
rior lower thermal pits (tp'), mental (m), ventral (v). Scale bars = 1 cm. (illustrated by Luhur Septiadi).
L Septiadi, M Fathoni, BF Hanifa, A Hamidy, 2019 / Morphological Variation of Malayopython reticulatus
JTLS | Journal of Tropical Life Science 262 Volume 9 | Number 3 | September | 2019
ated to avoid bias due to sexual dimorphism on
snakes, ToL, tl, SVL are converted into ratio. The
results of morphometric and meristic character
measurement were compared with previous stud-
ies. The data were further analysed using Principal
Component Analysis (PCA), which compared the
biggest variance (PC) from the data available.
Results and Discussion
Morphological characters of M. reticulatus from
several population
The analysis showed the high variation on the
scale range of anterior prefrontal, posterior pre-
frontal, frontal, parietal, preocular, postocular, lo-
real, temporal, upper labial, lower labial, and dor-
sal, ventral and subcaudal scales (Table 2). The
obtained meristic character shows the intraspecific
variations of the total number of scales that vary
between specimens from several populations. The
varying number of scales might be correlated to
habitat use and environmental conditions that
cause adaptation. This adaptation is done to pre-
vent excess water loss in the body. A large number
of scales might indicate habitats with high humid-
ity, whereas fewer scales might indicate a drier
habitat [6]. Reduction of dorsal scales might also
be affected by the development of the cutaneous
muscle system and cost-cutaneous muscles lo-
cated below the skin as a result of the evolution of
rectilinear locomotion [7].
Meristic characters do not show the sexual di-
morphism based on the number of ventral and sub-
caudal scales due to the high variation. Male spec-
imens have a range of 84-96 on subcaudal scales,
and a range of 284-343 on ventral scales. Whereas
in the female specimens, the total range is 83-100
on subcaudal scales and the total range is 294-341
on ventral scales. The measurement of the mor-
phometric ratio tail length (tl): snout-vent length
(SVL) from each population shows that the aver-
age ratio of tail length of the male specimens is
longer than the average ratio of tail length of fe-
male specimens (Table 2). According to Kerfoot
[7], ventral scales attach to the cost-cutaneous
Table 2. Measurement of meristic characters and morphometric ratios of M. reticulatus (=14, =24).
Characters
Population
Sumatera
Jawa
Kalimantan
NTB
Maluku
Sulawesi
(n=1)
(n=5)
(n=1)
(n=3)
(n=2)
(n=4)
(n=1)
(n=4)
(n=2)
(n=5)
(n=10)
r
1
1
1
1
1
1
1
1
1
1
1
i
2
2
2
2
2
2
2
2
2
2
2
apf
2
2-4
2
2
2
2
2
2
2
2-3
2
ppf
7
6-10
9
7-8
8
7-8
8
5-9
6
1-8
4-9
f
1
1-2
2
1-2
2
1-2
1
1-2
1
1
1-2
so
2
2
2
2
2
2
2
2
2
2
2
p
2
2-3
2
2
2
2
2
2
2-3
2
2
ip
1
2-3
1
3-5
2
2-4
2
1-3
2-3
1-2
1-3
pro
2
2
2
2
2
2-3
2
2
2
2
2
pso
2-4
4
3
4
3-5
3-4
4
2-4
3
2-4
2-4
l
3
3-5
5
4-5
2-4
2-6
4
3-4
4
2-6
3-4
t
4-5
6-8
7
6-7
7-8
6-8
6-7
5-7
5-7
5-7
4-8
La
14
13-14
13
13-14
14
12-13
13
12-14
13-14
12-15
11-13
Lao
7
7
7
7
7-8
7
7
7
7
7-8
6-8
la'
22
21-23
21-22
21-23
22-24
21
22
21-22
22
19-25
19-24
Tp
5
5-6
5
5
5
5
5
5
5
4-6
5
tp'
6
5-6
5
5
6
5
5
5
5
5-7
5-7
m
1
1
1
1
1
1
1
1
1
1
1
d
66
65-76
66
61-77
73-74
63-72
69
67-77
78-69
67-90
66-87
v
319
310-
319
320
301-
311
312-
320
316-
318
314
284-
322
320-
333
300-
343
294-
341
Sc
89
87-
100
94
87-93
89-95
87-96
96
84-91
90-95
87-94
83-94
tl:Tol
0.17:1
0.14:1
0.14:1
0.12:1
0.14:1
0.13:1
0.14:1
0.14:1
0.12:1
0.13:1
0.12:1
SVL:Tol
0.83:1
0.86:1
0.86;1
0.88:1
0.86:1
0.87:1
0.86:1
0.86:1
0.88:1
0.86:1
0.87:1
tl:SVL
0.20:1
0.16:1
0.16:1
0.14:1
0.16:1
0.15:1
0.16:1
0.16:1
0.13:1
0.15:1
0.14:1
L Septiadi, M Fathoni, BF Hanifa, A Hamidy, 2019 / Morphological Variation of Malayopython reticulatus
JTLS | Journal of Tropical Life Science 263 Volume 9 | Number 3 | September | 2019
muscle system that connected to the ribs and ver-
tebrae, so the number of ventral scales reflects the
number of vertebrae of the specimens. Individuals
with a large number of vertebrae have a possibility
to grew longer than individuals who have fewer
vertebrae, affecting the number of existing scales.
Differences in the shape and number of frontal
scales were also found with the presence of a sin-
gle, divided, anterior-posteriorly notch, and ante-
riorly notch frontal scales (Figure 2). It was stated
that this variation is due to geographical variations
expressed in the form of divided or single scales,
different from one region to another [8, 9]. Kluge
[10] states that there are several morphological
characters that are different from some of the spec-
imens studied in Kalimantan (Borneo) with a var-
iation of almost dividing from posterior, anterior,
or both, as well as singles and divided scales. Un-
derwood and Stimson, also Mark Auliya et al. [11,
4] state that the formation of the frontal scales is a
type of intraspecific variation. These findings
strongly support the existence of intraspecific var-
iations in frontal scales in the population studied,
and cannot be categorized as geographic varia-
tions based on frontal scales of reticulated python.
The diagnostic characters of M. reticulatus sbp.
problems
Prefrontal scales show inconsistencies in a
variation of M. reticulatus specimens in each pop-
ulation (Table 3). Mark et al. [4], states that there
were specific variations in the row of prefrontal
scales in each M. reticulatus sbp. The species of
M. reticulatus reticulatus has 2 rows of posterior
prefrontal scales with slightly short anterior pre-
frontal scales, the species of M. reticulatus jam-
peanus has only 1 row of prefrontal posterior
scales, with elongated anterior prefrontal scales.
Whereas M. reticulatus saputrai has 2 rows of
posterior prefrontal scales, and slightly enlarged
anterior prefrontal scales. Direct observations
showed specimens from Sulawesi has 1-row of
fragmented posterior prefrontal scales and anterior
prefrontal scales on varying size (ID. Number:
MZB.Ophi. 1762, MZB.Ophi. 2297, MZB.Ophi.
1558, MZB.Ophi. 3221) resembles M. reticulatus
jampeanus. Specimens from Maluku also showed
the presence of 1 row posterior prefrontal scales
with elongated anterior prefrontal scales (ID.
Number: MZB.Ophi. 1951, MZB.Ophi.4481,
MZB.Ophi. 4482). However, the specimen ob-
served from Tanahjampea, Kalaotoa, Karompa,
Figure 2. Comparisons of the Frontal Scales of several specimens from various localities, Single (top-left), Di-
vided (top-right), Anterior-Posteriorly Notch (bottom-left), and Anteriorly Notch Frontal Scales (bot-
tom-right) (blue lines indicate the notch).
L Septiadi, M Fathoni, BF Hanifa, A Hamidy, 2019 / Morphological Variation of Malayopython reticulatus
JTLS | Journal of Tropical Life Science 264 Volume 9 | Number 3 | September | 2019
and Madu Island (ID. Number: MZB. Ophi. 3223,
MZB. Ophi. 3437, S.Voucher.020,
S.Voucher.021, S.Voucher.022, S.Voucher.023)
shows the tendency of 1 row posterior prefrontal
scales with elongated anterior prefrontal scales, re-
sembling the diagnostic character of the M. retic-
ulatus jampeanus. In accordance Hanifa et al.
[12], which suggest that the Karompa, Kalaotoa,
and Madu islands were acting as stepping stones
on the dispersal patterns of this species, though
Table 3. Comparison of dorsal, ventral, prefrontal scales to few literature of M. reticulatus
Population
Mid- Dorsal
Scale
Ventral
Scales
Prefrontal Scales
Resemblance
Source
(Sunda land)
M. r. reticulatus
68-78
304-325
2 rows ppf, apf small
-
[4]
(Selayar)
M r. jampeanus
64-68
290-301
1 row of ppf, apf slightly
longer than Sunda retic
-
[4]
(TanahJampea)
M. r. saputrai
77-81
330-334
2 rows of ppf, apf very
long
-
[4]
M. reticulatus
69-79
297-330
Variated
Sulawesi Popula-
tion
[13]
M. reticulatus
71-75
300-322
-
Java Population
[1]
Sumatera (n=6)
65-76
310-319
Variated
to M. r. reticulatus
This study
Kalimantan
(n=6)
63-74
312-320
2 rows of ppf
to M. r. reticulatus
This study
Jawa (n=4)
61-77
301-320
2 rows of ppf
to M. r. reticulatus
This study
Sulawesi (n=13)
66-90
294-343
Variated
-
This study
Maluku (n=6)
67-78
284-333
Variated
to Sulawesi Popula-
tion
This study
NTB (n=1)
69
314
2 rows of ppf
to M. r. reticulatus
This study
Figure 3. Two-dimensional Scatter plot of 38 M. reticulatus specimens from several population
(Sumatra: Brown, Java: Yellow, Kalimantan: Green, NTB: Red, Sulawesi: Blue, and
Maluku: Turqoise)
L Septiadi, M Fathoni, BF Hanifa, A Hamidy, 2019 / Morphological Variation of Malayopython reticulatus
JTLS | Journal of Tropical Life Science 265 Volume 9 | Number 3 | September | 2019
share similarities to the M. reticulatus jampeanus
based on morphological and molecular data.
However, these variations are still difficult to
distinguish morphologically based on their local-
ity (reticulatus: Greater Sunda-land; saputrai: Se-
layar; jampeanus: Tanahjampea), given the high
morphological variations in these specimens, alt-
hough phylogenetically have proven to be distinct
subspecies [4]. This underlines that the diagnostic
character of subspecies based on morphological
differences in the prefrontal scales of M. reticula-
tus is still ambiguous and needs deep observation
in other characters such as anatomical and mor-
phological features. Specifically, a measurement
of prefrontal scales (Anterior-Posterior) are sug-
gested in the future analysis to distinguish this spe-
cies.
Population grouping of M. reticulatus from In-
donesia
Scatter plots show an overlap in each observed
population. The population of Sulawesi has a wide
plot width, due to the strong number of Ventral
and Dorsal mid characters, where the number of
Ventral Scales in the Sulawesi Population and
some specimens from Sumatra and Kalimantan
have a higher range than other populations.
Based on the analysis, there is no significant
differences were found between populations from
data on morphometric variations and prominent
signs of character variation among the islands.
Characters that are owned by a population are still
owned by other populations. Geographic isolation
has not made any morphological variations that
can be used as distinguishing characters. This pos-
sible in regards to the varied landscape, habitats,
and ecosystems in Indonesia, making it difficult to
select a new adaptive character.
Conclusion
The analysis showed the highly variation on
the scale range of anterior prefrontal, posterior
prefrontal, frontal, parietal, preocular, postocular,
loreal, temporal, upper labial, lower labial, and
dorsal scales. Ventral and subcaudal scales in male
and female specimens showed highly variation in
the scale count, but ratio comparison of tl:SVL
length indicated the sexual dimorphism. Anterior-
posterior prefrontal and frontal scales show high
variation and are difficult to distinguish between
each locality at the subspecies level and suggest it
to be intra-specific variation. There is no signifi-
cant grouping were found between populations
from data on morphological variations.
Acknowledgment
We would like to thank Irvan Sidik from La-
boratory of Herpetology, Zoology Division, Re-
search Center for Biology, Indonesian Institute of
Sciences LIPI, who have been very helpful in dis-
cussing and criticizing our research idea. We are
also very grateful to Herpetology LIPI Internship
team 2018 from various which is Pakuan Univer-
sity, Brawijaya University, also State Islamic Uni-
versity of Maulana Malik Ibrahim Malang,
who’ve helped in the process of documenting the
specimens. We would also thank to Maliki Herpe-
tology Society Study Club which provides a place
to study and discuss. This study partially sup-
ported by The Mohamed bin Zayed Species Con-
servation Fund, project number 13057088 and
IDEA WILD in providing equipment in docu-
menting specimens so the illustration process is
supported.
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Twenty-four extant species of snakes, usually referred to as pythonines (sensu Underwood, 1976), are compared in terms of 121 behavioural and external and internal morphological characters. A cladistic analysis of 194 synapomorphies confirms the monophyly of the group, and provides a partially resolved, well-corroborated hierarchy of lineage relationships. That hypothesis obtains without regard to assumptions of additivity or nonadditivity, and only those synapomorphies which delimit clades unambiguously are used to diagnose taxa. Aspidites is demonstrated to be the sister lineage of all other pythonines, and the remaining Australia-New Guinea taxa constitute a paraphyletic assemblage. The South-east Asia-Africa Python forms a highly derived clade. The following binominal monophyletic taxonomy is proposed: Antaresia childreni, A. maculosus, A. perthensis, A. stimsoni, Apodora papuana (n.gen.), Aspidites melanocephalus, A.
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The Central American Loxocemus and the pythons are assigned to sister subfamilies Loxoceminae and Pythoninae, within the weakly characterized family Boidae. They share the character of the left cerebral artery foramen being larger than the right. Within the Henophidia, paired subcaudal scales associate these subfamilies with Xenopeltis and the Uropeltidae. Twenty-nine discrete variables were coded as 52 binary characters. Nine continuous (meristic) variables were examined. A method is described by which 18 binary characters were selected from seven of these variables, for inclusion in the data set. With Loxocemus and Xenopeltis as outgroups, the data were analysed by a compatibility method. The Pythoninae are resolved into tribes Pythonini and Moreliini on the basis of complementary synapomorphies. The Pythonini have only the genus Python, for the African and Asiatic species, including reticulatus and timoriensis. The Moreliini have all of the Australasian species, in two genera: Aspidites and Morelia. A phylogenetic analysis of the two tribes is presented, with incomplete resolution of the Moreliini. The two tribes overlap in the areas of the Moluccas and Timor. It is postulated that: Loxocemus plus Pythoninae represent a Laurasian stock; south-east Asia was the primary centre of radiation of the Pythoninae; by the Miocene dispersal to Australasia had occurred, where there was a second radiation.
The Snakes of Java, Bali and Surrounding Islands
  • De Lang
De Lang R (2017) The Snakes of Java, Bali and Surrounding Islands. Ohio, Chimaira Edition.