The complete mitochondrial genome of an agamid lizard from the Afro-Asian subfamily agaminae and the phylogenetic position of Bufoniceps and Xenagama.
- SourceAvailable from: Jason J Head[Show abstract] [Hide abstract]
ABSTRACT: Mammals dominate modern terrestrial herbivore ecosystems, whereas extant herbivorous reptiles are limited in diversity and body size. The evolution of reptile herbivory and its relationship to mammalian diversification is poorly understood with respect to climate and the roles of predation pressure and competition for food resources. Here, we describe a giant fossil acrodontan lizard recovered with a diverse mammal assemblage from the late middle Eocene Pondaung Formation of Myanmar, which provides a historical test of factors controlling body size in herbivorous squamates. We infer a predominately herbivorous feeding ecology for the new acrodontan based on dental anatomy, phylogenetic relationships and body size. Ranking body masses for Pondaung Formation vertebrates indicates that the lizard occupied a size niche among the larger herbivores and was larger than most carnivorous mammals. Paleotemperature estimates of Pondaung Formation environments based on the body size of the new lizard are approximately 2-5°C higher than modern. These results indicate that competitive exclusion and predation by mammals did not restrict body size evolution in these herbivorous squamates, and elevated temperatures relative to modern climates during the Paleogene greenhouse may have resulted in the evolution of gigantism through elevated poikilothermic metabolic rates and in response to increases in floral productivity.Proceedings of the Royal Society B: Biological Sciences 05/2013; 280(1763):20130665. · 5.29 Impact Factor
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ABSTRACT: Abstract The complete mitogenome sequence of a mystical lizard species Phrynocephalus mystaceus was determined using polymerase chain reaction and directly sequenced with a primer walking method. The complete mitogenome was 16,660 bp in length, containing 13 protein-coding genes, 22 tRNA genes, two rRNA genes and a control region (D-loop). The gene arrangement and composition of P. mystaceus was similar to most other vertebrates, but the Proline tRNA gene was translocated to be adjacent to tRNA-Phe gene. The D-loop consisted of two parts, with part I existing between the tRNA-Thr gene and tRNA-Pro gene and another part inserting between the tRNA-Phe and 12S rRNA. In part I, one conserved sequence (CSB I) could be identified. In part II, two pair of motifs, "TACAT" and its reverted and complemented sequence "ATGTA", could be found in the domain of an extended termination-associated sequence. The mitogenome sequence of P. mystaceus could contribute to a better solution of its phylogenetic position within toad-headed agamids based on the whole mitogenomic data.Mitochondrial DNA 03/2013; · 1.70 Impact Factor
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ABSTRACT: Africa is renowned for its biodiversity and endemicity, yet little is known about the factors shaping them across the continent. African Agama lizards (45 species) have a pan-continental distribution, making them an ideal model for investigating biogeography. Many species have evolved conspicuous sexually dimorphic traits, including extravagant breeding coloration in adult males, large adult male body sizes, and variability in social systems among colorful versus drab species. We present a comprehensive time-calibrated species tree for Agama, and their close relatives, using a hybrid phylogenetic-phylogenomic approach that combines traditional Sanger sequence data from five loci for 57 species (146 samples) with anchored phylogenomic data from 215 nuclear genes for 23 species. The Sanger data are analyzed using coalescent-based species tree inference using ∗BEAST, and the resulting posterior distribution of species trees is attenuated using the phylogenomic tree as a backbone constraint. The result is a time-calibrated species tree for Agama that includes 95% of all species, multiple samples for most species, strong support for the major clades, and strong support for most of the initial divergence events. Diversification within Agama began approximately 23 million years ago (Ma), and separate radiations in Southern, East, West, and Northern Africa have been diversifying for > 10 Myr. A suite of traits (morphological, coloration, and sociality) are tightly correlated and show a strong signal of high morphological disparity within clades, whereby the subsequent evolution of convergent phenotypes has accompanied diversification into new biogeographic areas.Molecular Phylogenetics and Evolution 06/2014; 79:215-230. · 4.02 Impact Factor
Molecular Phylogenetics and Evolution 39 (2006) 881–886
1055-7903/$ - see front matter Published by Elsevier Inc.
The complete mitochondrial genome of an agamid lizard from
the Afro–Asian subfamily agaminae and the phylogenetic position
of Bufoniceps and Xenagama
J. Robert Maceya,¤, James A. Schulte IIb, Jonathan J. Fonga, Indraneil Dasc,
Theodore J. Papenfussa
a Museum of Vertebrate Zoology, 3101 Valley Life Science Building, University of California, Berkeley, CA 94720, USA
b Department of Biology, 177 Clarkson Science Center, MRC 5805, 8 Clarkson Avenue, Clarkson University, Potsdam, NY 13699-5805, USA
c Institute of Biodiversity and Environmental Conservation,Universiti Malaysia Sarawak, 94300, Kota Samarahan, Sarawak, Malaysia
Received 21 July 2005; revised 21 August 2005; accepted 30 August 2005
Squamate reptiles are traditionally divided into six
groups: Iguania, Anguimorpha, Scincomorpha, Gekkota
(these four are lizards), Serpentes (snakes), and Amphisbae-
nia (the so-called worm lizards). The Iguania is recognized
as having two major lineages the Iguanidae and Acrodonta
(Agamidae and Chamaeleonidae). Currently, there are
complete mitochondrial genomes from three Anguimorpha
(Kumazawa, 2004; Kumazawa and Endo, 2004), two from
the Scincomorpha (Kumazawa, 2004; Kumazawa and
Nishida, 1999), one from Gekkota (Macey et al., 2005) two
from Serpentes (Kumazawa, 2004; Kumazawa et al., 1998)
and 12 from Amphisbaenia (Macey et al., 2004). In addi-
tion, two representatives of the Iguanian family Iguanidae
(Janke et al., 2001; Kumazawa, 2004) have been sequenced.
Its’ sister taxon, the Acrodonta, consists of seven monophy-
letic groups-the family Chamaeleonidae and six distantly
related subfamilies of the family Agamidae (Macey et al.,
2000b). Currently, the only acrodont lineage sequenced for
the complete mitochondrial genome is Pogona vitticeps
from the Australasian agamid subfamily Amphibolurinae
(Amer and Kumazawa, 2005a).
Here, we report the complete mitochondrial genome of
Xenagama taylori, a North African representative of the
agamid subfamily Agaminae and compare it to P. vittice-
pes. The agamid lizard genus Xenagama is distributed in a
restricted region of the Horn of Africa in northwestern
Somalia and adjacent eastern Ethiopia as shown in Fig. 1,
with two species currently recognized (Moody, 1980; Wer-
muth, 1967). In addition, we report a segment of the mito-
chondrial genome of Bufoniceps laungwalansis spanning
from nad1 to cox1. The monotypic genus Bufoniceps is
restricted to the Thar Desert, Jaisalmer District, Rajasthan
State, India and adjacent Pakistan (Fig. 1).
Both Bufoniceps and Xenagama belong to the subfamily
Agaminae and are poorly understood phylogenetically.
These genera were not represented in the most recent
molecular systematic study of the Agamidae (Macey et al.,
2000b). Bufoniceps was originally described as a member of
the West Asian genus Phrynocephalus (Sharma, 1978), and
later placed in its’ own genus (Arnold, 1992) because mor-
phological data suggested it is the sister taxon to Phryno-
cephalus (Arnold, 1999). Xenagama was previously
considered part of the Agama complex before the alloca-
tion of its member species to several genera (see, Moody,
2. Materials and methods
2.1. Specimen information
The sample of Bufoniceps laungwalansis from which
DNA was extracted is deposited in the RaZes Museum of
Biodiversity Research, National University of Singapore as
ZRC 2.5681. The collection locality of this specimen is ele-
vation 192m, 26.50.26?N 70.32.24?E, vicinity of Sam, Rajas-
than State, India. The mitochondrial segment spanning
*Corresponding author. Fax: +1 510 643 8238.
E-mail address: firstname.lastname@example.org (J.R. Macey).
J.R. Macey et al. / Molecular Phylogenetics and Evolution 39 (2006) 881–886
from nad1 to cox1 is deposited in GenBank as Accession
No. DQ008214. The sample of X. taylori from which DNA
was extracted is deposited in the California Academy of
Sciences, San Francisco as CAS 225502. The collection
locality of this specimen is elevation 1140m, 9.670000?N
44.207500?E, 21km ENE of the center of Hargeysa on Ber-
bera Rd., then 4km N on dirt road, Waqooyi Galbed
Region, Somalia. The complete mitochondrial genome
sequence from this specimen of X. taylori is deposited in
GenBank as Accession No. DQ008215.
2.2. Laboratory protocols
Genomic DNA was extracted from liver using the
Qiagen QIAamp tissue kit. For B. laungwalansis, ampliWca-
tion of genomic DNA was conducted using a denaturation
at 94°C for 35s, annealing at 50°C for 35s, and extension
at 70°C for 150s with 4s added to the extension per cycle,
for 30 cycles. Primers used are described in Macey et al.
(1997a,c, 2000b). Negative controls were run on all ampliW-
cations to check for contamination. AmpliWed products
were puriWed on 2.5% Nusieve GTG agarose gels and ream-
pliWed under the conditions described above to increase
DNA yield for downstream sequencing reactions. Reampli-
Wed double-stranded products were puriWed on 2.5% acryl-
amide gels and template DNA was eluted passively over
three days with Maniatis elution buVer (Maniatis et al.,
1982) or puriWed using the QIAquick PCR puriWcation kit.
Cycle-sequencing reactions were run using the ABI Prism
Big Dye Terminator DNA Sequencing Kit (Perkin-Elmer)
with a denaturation at 95°C for 15s, annealing at 50°C for
1s, and extension at 60°C for 4min for 35–40 cycles.
Sequencing reactions were run on an ABI 373 Genetic Ana-
lyzer or MJ Research Basestation sequencers.
For X. taylori ampliWcation of the mtDNA was con-
ducted using rT th long PCR enzyme (Applied Biosystems)
with a beginning denaturation at 94°C for 45s, then fol-
lowed by 37 cycles of a denaturation at 94°C for 15s,
annealing at 50°C for 20s, and extension at 68°C for
9min, with a Wnal extension at 72°C for 12min after the
last cycle. Negative controls were run on all ampliWcations
to check for contamination. Initial ampliWcations were
conducted using primers described in Macey et al. (1997a).
Perfectly matching primers were then constructed based
on the DNA sequence of this fragment to complete the
ampliWcation of the mtDNA. AmpliWcation products were
sheared randomly into fragments of approximately 1.5kb
by repeated passage through a narrow aperture using a
Hydroshear device (GeneMachines). After end-repair, the
sheared DNA was gel puriWed and ligated into pUC18 vec-
tor to construct a library of random fragments, then trans-
formed into bacterial cells. Automated colony pickers
introduced single clones into bacterial broth in 384-well
format. These plasmid clones were processed robotically
through rolling circle ampliWcation (Dean et al., 2001;
Hawkins et al., 2002), sequencing reactions, and reaction
clean up using SPRI (Elkin et al., 2002). Sequences were
determined using ABI3730xl DNA sequencers, then
assembled to form a deep, contiguous sequence using
Phrap or Sequencher.
Fig. 1. Map showing the distribution of Bufoniceps laungwalansis and Xenagama taylori. Each taxon has a limited distribution with B. laungwalansis
restricted to the Thar Desert in western India and adjacent Pakistan. The two species of Xenagama are restricted to the Horn of Africa in Somalia and
adjacent Ethiopia. Other members of the Agaminae range from North Africa through Arabia, Southwest Asia to Central Asia, and Tibet (not shown).
I n d i a
I r a n
S a u d i
A r a b i a
E t h i o p i a
J.R. Macey et al. / Molecular Phylogenetics and Evolution 39 (2006) 881–886
2.3. Phylogenetic analysis
DNA sequences for protein- and tRNA-encoding genes
were aligned manually as in Macey et al. (2000b). Positions
encoding proteins were translated to amino acids using
MacClade 4.03 (Maddison and Maddison, 2001) for conWr-
mation of alignment. Alignments of sequences encoding
tRNAs were constructed based on secondary structural
models (Kumazawa and Nishida, 1993; Macey and Verma,
1997). Of the 1965 characters, unalignable regions totaling
561 positions were excluded from phylogenetic analyses as
in Macey et al. (2000b).
The region analyzed in Macey et al. (2000b) from nad1
to cox1 corresponds to positions 3495–5193 of the complete
mitochondrial genome of X. taylori and has a length of
1699 bases; this is the region used in phylogenetic analysis.
To align the new sequences with the 72 sequences analyzed
in Macey et al. (2000b) a total of 268 gaps are introduced in
the B. laungwalansis sequence and 266 gaps in the X. taylori
sequence. These gaps are after the following positions on
the B. laungwalansis sequence with the number of gaps
introduced in parenthesis if more than one: 86 (18), 99, 145
(2), 162 (13), 181 (2), 218 (4), 229 (11), 246 (2), 280 (4), 295
(3), 298 (12), 319 (3), 340 (3), 1105 (3), 1318 (9), 1321 (82),
1343 (3), 1369, 1380 (4), 1393 (10), 1412, 1448, 1461 (13),
1490 (3), 1507, 1515, 1528, 1553 (13), 1570 (5), 1602 (6),
1609 (6), 1625 (3), 1634, 1661 (2), and 1674 (11). These gaps
are after the following positions on the complete mitochon-
drial genome of X. taylori with the number of gaps intro-
duced in parenthesis if more than one: 3579 (19), 3592, 3638
(2), 3652 (16), 3670 (3), 3706 (5), 3718 (10), 3736, 3769 (6),
3782 (4), 3785 (9), 3809 (3), 3830 (3), 4595 (3), 4808 (9), 4812
(81), 4833 (4), 4859, 4870 (4), 4883 (10), 4900 (3), 4936, 4949
(13), 4978 (3), 4995, 5003, 5016, 5041 (13), 5057 (6), 5088 (6),
5096 (6), 5111 (4), 5120, 5147 (2), and 5160 (11). Sequence
divergences based on this alignment are reported as uncor-
rected pairwise divergences.
Phylogenetic trees were inferred by parsimony using
PAUP* ? version 4.0b8 (SwoVord, 2001) with heuristic
searches featuring 100 random additions of sequences.
Bootstrap resampling (Felsenstein, 1985a) was applied to
assess support for individual nodes using 500 heuristic
searches featuring 100 random additions of sequences per
replicate. Decay indices (D‘branch support’ of Bremer,
1994) were calculated for all internal branches using heuris-
tic searches featuring 100 random additions of sequences in
searches that retained suboptimal nodes.
To test speciWc, alternative phylogenetic hypotheses, we
Wrst built incompletely resolved constraint trees using
MacClade (Maddison and Maddison, 2001). These were
provided as input into PAUP* (SwoVord, 2001) for heuris-
tic searches featuring 100 random additions of sequences to
determine the most parsimonious tree compatible with each
alternative hypothesis. We then compared these to the
unconstrained most parsimonious tree using Wilcoxon
signed-ranks tests (Templeton, 1983). This test determines
whether the most parsimonious tree is signiWcantly shorter
than each alternative or whether their diVerences in length
are statistically indistinguishable. Wilcoxon signed-ranks
tests were conducted as one-tailed tests (Felsenstein, 1985b)
using PAUP* (SwoVord, 2001), which incorporates a cor-
rection for tied ranks. Felsenstein (1985b) showed that one-
tailed probabilities are close to the exact probabilities for
this test but not always conservative. The two-tailed proba-
bilities are simply double the one-tailed probabilities and
the two-tailed test is always conservative (Felsenstein,
3.1. Mitochondrial genomic structure
The complete mitochondrial genome of X. taylori is
16,220bp in length. This genome contains the same 37
genes common among animals but diVers from the order
as is most commonly found for vertebrates (Boore, 1999).
As previously reported for Acrodonta (Agamidae and
Chamaeleonidae), both B. laungwalansis and X. taylori
have trnI and Q switched to yield the order nad1, trnQ, I,
and M (Macey et al., 1997c, 2000a). These taxa lack any
duplicated genes or secondary structure of non-coding
sequences between trnQ and I as has been identiWed in
some species of the agamid genus Uromastyx (Amer and
Kumazawa, 2005b). Both taxa have trnC that encodes a
transfer RNA which lacks a D-stem and instead contains
a D-arm replacement loop (Macey et al., 1997b), as is typ-
ical for the Acrodonta (Macey et al., 2000a). In addition,
these taxa have atypical stem–loop structures between
trnN and trnC where light-strand replication is thought to
usually initiate for vertebrate mtDNAs, which is also
observed in other members of the Agaminae clade (Macey
et al., 2000a). In particular, the 3?-GCC-5? heavy strand
template sequence identiWed as the point of light-strand
elongation in mouse (Brennicke and Clayton, 1981) is not
present in these structures. B. laungwalansis has a Wve base
stem with a 16 base loop, whereas X. taylori has a nine
base stem with an eight base loop. As observed in most
other vertebrates, the mt-genome of X. taylori has a large
noncoding region presumed to be the Control Region
(CR) of 1449bp in length. This diVers from that of the
amphibolurine P. vitticeps which has a near identical sec-
ond noncoding region inserted between nad5 and nad6
(Amer and Kumazawa, 2005a). Unlike previously
reported mt-genomes among vertebrates (but see, McK-
night and ShaVer, 1997) the noncoding region in X. taylori
is between trnT and trnP and not between trnP and trnF,
as is typical and observed for one CR copy in P. vitticeps
(Amer and Kumazawa, 2005a). An additional 26 bases
separate trnP and trnF in X. taylori. Therefore, X. taylori
has the complete mt-order of trnF, rrnS, trnV, rrnL,
trnL(taa), nad1, trnQ, I, M, nad2, trnW, A, N, C, stem–
loop, trnY, cox1, trnS(tga), D, cox2, trnK, atp8, atp6,
cox3, trnG, nad3, trnR, nad4L, nad4, trnH, S(tct), L (tag),
nad5, nad6, trnE, cob, trnT, CR, and trnP.
J.R. Macey et al. / Molecular Phylogenetics and Evolution 39 (2006) 881–886
3.2. Phylogenetic relationships
Phylogenetic analysis of the 1434 aligned positions (1046
informative) for the 72 taxa in Macey et al. (2000b) and the
two newly reported sequences from nad1 to cox1 produces
seven equally most parsimonious trees (Fig. 2). The tree is
largely the same as that reported by Macey et al. (2000b)
with the exception of a few weak nodes that are collapsed in
this analysis because of the four additional equally parsi-
monious trees. Here, we concentrate on the phylogenetic
relationships of the Agaminae, which is monophyletic with
a bootstrap of 100% and decay index of 27. The strict con-
sensus tree yields nine lineages in the Agaminae labeled A–I
in Fig. 2. The African genus Agama (sensu stricto) appears
monophyletic with strong support (A in Fig. 2, bootstrap
100%, decay index 97). Pseudotrapelus sinaitus of Arabia,
Egypt, and Libya groups with X. taylori from the Horn of
Africa and is well-supported (B in Fig. 2, bootstrap 99%,
decay index 17). B. laungwalansis, restricted to the Indian
Subcontinent, forms the sister taxon to the wide-ranging
genus Trapelus with considerable support (C in Fig. 2,
bootstrap 100%, decay index 22). Laudakia sacra of Tibet
forms its own deep-lineage (D in Fig. 2). Laudakia nupta of
the Iranian Plateau groups weakly with Laudakia tubercu-
lata of the Himalaya (E and F in Fig. 2, bootstrap 57%,
decay index 2). Laudakia stellio of Anatolia and the Levant
forms its own deep-lineage (G in Fig. 2). The Asian genus
Phrynocephalus is well supported (H in Fig. 2, bootstrap
100%, decay index 41). A well supported clade of Laudakia
ranging from the Iranian Plateau to Mongolia is present (I
in Fig. 2, bootstrap 100%, decay index 20).
The Wilcoxon-signed-rank test (Felsenstein, 1985b;
Templeton, 1983) is applied to compare the most parsimo-
nious tree from these nucleotide sequences with alternative
hypotheses. The genus Bufoniceps has been previously sug-
gested to be either in the genus Phrynocephalus or the sister
taxon to Phrynocephalus. The seven shortest alternative
trees that unite Bufoniceps with Phrynocephalus require 55
extra steps and are rejected in favor of the unconstrained
shortest trees (P<0.0046). The genus Xenagama has been
previously suggested to be related to Agama. The two
shortest alternative trees that unite Xenagama with Agama
require 30 extra steps and are not rejected in favor of the
unconstrained shortest trees (P<0.0861).
4.1. Biogeography and the breakup of Gondwana
Acrodont lizards (Agamidae and Chamaeleonidae) are
of Gondwanan origin (Macey et al., 2000b). Clades of
agamid lizards rafted with alternative fragments of Gondw-
ana which collided with the southern margin of Asia. The
subfamily Agaminae arrived in Asia either with the Indian
Subcontinent 50 MYBP (million years before present) or
with Afro-Arabia 18 MYBP.
Bufoniceps is found to be the sister taxon to Trapelus and
is statistically rejected as the sister taxon to Phrynocephalus
as previously suggested (Arnold, 1999). Note that Bufoni-
ceps and Trapelus share the morphological character of an
open ear, and it is Phrynocephalus that has the derived fea-
ture of a closed ear (Arnold, 1999). The genus Trapelus
ranges from North Africa across Arabia, through the Ira-
nian Plateau and Caspian Basin to the western edge of the
Indian Subcontinent in the vicinity of the range of Bufoni-
ceps. Bufoniceps is restricted to a small region of the Thar
Desert on the western edge of the Indian Subcontinent. The
tree presented in Macey et al. (2000b) was suggestive of an
Afro-Arabian origin for the Agaminae but the alternative
of an origin in the Indian Subcontinent could not be
rejected. The analysis presented here is equivocal for an ori-
gin in either region because of a basal polytomy. The fact
that the sister taxon to a major clade of the Agaminae is an
Indian endemic raises the question of a possible origin of
the Agaminae in the Indian Subcontinent. Further work is
needed to resolve this issue, perhaps with phylogenetic
analysis of complete mitochondrial genomes.
Fig. 2. The strict consensus of seven most parsimonious trees resulting
from analysis of the 1434 (1046 informative) aligned sites which is 12,236
steps in length. Bootstrap values appear above branches and decay indices
are presented below. Note the analysis includes all 72 taxa from Macey
et al. (2000b) and the two newly reported sequences but only taxa in the
subfamily Agaminae are shown here. The new taxa, Bufoniceps laungwa-
lansis and Xenagama taylori are depicted in bold as is the support for their
placement in the Agaminae. The nine major lineages of the Agaminae are
delineated to the right as A–I.
J.R. Macey et al. / Molecular Phylogenetics and Evolution 39 (2006) 881–886
Xenagama is found to be the sister taxon to Pseudotrape-
lus and not to the African genus Agama. Pseudotrapelus
occurs in Arabia and adjacent regions of Egypt and Libya.
Xenagama is restricted to a small region of the Horn of
Africa in Somalia and Ethiopia directly across the Red Sea
from Arabia where Pseudotrapelus is found. The Afro-Ara-
bian Plate began to divide along the Red Sea rift 40 MYBP
but accelerated 5–10 MYBP and there have been periodic
connections between the Horn of Africa and Arabia (Gir-
4.2. Sequence divergences and age
The region of mitochondrial DNA examined here span-
ning from nad1 to cox1 has been shown to evolve at a rate
of 1.3% per million years for uncorrected pairwise compari-
sons in agamine lizards (Macey et al., 1998). This calibra-
tion has been shown to be robust across numerous
amphibian and reptile taxa (reviewed in Weisrock et al.,
2001). In addition, we calibrated a transversional rate of
0.98% per million years by comparing divergences of Lau-
dakia from the Pamir (L. lehmanni, L. himalayana, and L.
stoliczkana) with those from the Iranian Plateau (L. micro-
leps, L. caucasia, and L. erythrogastra). The rise of the
Pamir–Tien Shan is well dated at 10 MYBP (million years
before present; Abdrakhmatov et al., 1996).
Pairwise sequence divergences are presented in Table 1.
Using straight uncorrected pairwise distances we calculate
a 17.6 MYBP separation of Xenagama and Pseudotrapelus,
and using the transversional pairwise distances a 19.2
MYBP separation. Although mitochondrial DNA is
known to begin to accumulate multiple substitutions at the
same site beyond 10 million years (Moritz et al., 1987) sug-
gesting that a linear relationship between sequence diver-
gence and time may not be expected, our two estimates are
quite similar. These dates Wt well with earlier vicariant sepa-
ration of Africa and Arabia across the Red Sea rift and do
not Wt with the more recent activity 5–10 MYBP.
Using straight uncorrected pairwise distances we calcu-
late a 15.7 MYBP separation of Bufoniceps and Trapelus,
and using the transversional pairwise distances a 16.8
MYBP separation. No obvious geologic barrier currently
exists between Bufoniceps and Trapelus. Indeed, the Trape-
lus agilis complex has been found in sympatry with B. laun-
gwalansis at Sam, Rajasthan, India. Perhaps along the
western margin of the Indian Subcontinent, tectonic activ-
ity, coupled with the second phase of Tibetan uplifting 20
MYBP (Le Fort, 1998; Searle, 1991), caused the speciation
event that divided these taxa. This would have been fol-
lowed by subsequent dispersal of Trapelus back into the
Indian Subcontinent. Clearly, Miocene events are responsi-
ble for divergences of the Agaminae.
Karen Klitz prepared Fig. 1. We thank Michelle Koo
for information on morphological character states.
This work is LBNL-57501 and was performed under the
auspices of the US Department of Energy’s OYce of
Science, Biological and Environmental Research Program
and by the University of California, Lawrence Livermore
National Laboratory under Contract No. W-7405-Eng-
48, Lawrence Berkeley National Laboratory under
Contract No. DE-AC03-76SF00098, and Los Alamos
National Laboratory under Contract No. W-7405-
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Sequence divergences across taxaa
aValues above the dashed line are uncorrected pairwise distances and those below are uncorrected transversional distances. Note genera are abbreviated
as P., Pseudotrapelus; X., Xenagama; B., Bufoniceps; and T., Trapelus.
1. P. sinaitus
2. X. taylori
3. B. laungwalansis
4. T. ruderatus
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