Cutting the Gordian Knot: Phylogenetic and ecological diversification of the Mesalina brevirostris species complex (Squamata, Lacertidae)
Author
Jiří Šmíd
Author
Jiří Moravec
Author
Václav Gvoždík
Author
Jan Štundl
Author
Daniel Frynta
Author
Petros Lymberakis
Author
Paschalia Kapli
Author
Thomas Wilms
Author
Andreas Schmitz
Author
Mohammed Shobrak
Author
Saeed Hosseinian Yousefkhani
Author
Eskandar Rastegar-Pouyani
Author
Aurora M. Castilla
Author
Johannes Els
Author
Werner Mayer
text
Zoologica Scripta
2017
2017-10-31
46
6
1
30
journal article
10.1111/zsc.12254
44b12099-59c0-4c00-b34c-c4f14d37ec56
1037920
Mesalina
saudiarabica
Moravec
, Šmíd
, Schmitz
, Shobrak
, Wilms
– sp. n.
Mesalina
brevirostris
– Kapli
et al.
(2015)
Mesalina
sp. – Kapli
et al
. (
2015
)
Holotype
.
ZFMK 91912
, subadult male,
Mahazat
asSayd,
Makkah Province
,
Saudi Arabia
,
22.237 N
,
41.843 E
,
1000 m
a.s.l.
, collected in
October 2006
by
T. Wilms. MorphoBank
pictures: M407306–M407312.
Paratype
.
ZFMK 86583
, subadult male,
Mahazat
asSayd, near
Al Muwayh
,
Makkah Province
,
Saudi Arabia
,
22.395 N
41.753 E
,
960 m
a.s.l.
, collected in
October 2006
by
T
. Wilms. MorphoBank
picture: M410851.
Referred material not included in the
type
series.
NMP
6
F
2930 (photovoucher), adult male, observed on the
type
locality in
October 2006
by
T
. Wilms.
Diagnosis.
A
species of
Mesalina
and a member of the
M
. brevirostris
species complex as revealed by the genetic analyses and characterized by the following combination of characters: (1) genetic (uncorrected) distance of 2.0% from
M
. brevirostris
, 2.7% from
M
. bernoullii
, and 2.8% from
M
. microlepis
for the
12
S
(after Gblocks); 2.9% from
M
. brevirostris
, 3.3% from
M
. bernoullii
, and 4.0% from
M. microlepis
for the
16S
(after Gblocks); 9.5% from
M. brevirostris
, 7.5% from
M. bernoullii
, and 10.8% from
M. microlepis
for the
cytb
; (2) low number of dorsal scales (41–42); (3) low number of collar plates (6–8); (4) low number of preanal scales (2–3); (5) low number of femoral pores in males (12–13); (6) having 1–2 large semitransparent scales in the lower eyelid window; (7) in life, dorsum light cinnamon brown with a pattern of small whitish and larger dark cinnamon spots arranged in more or less regular longitudinal rows. Most of the whitish spots are not edged with dark brown color. The dark cinnamon brown spots predominate on flanks where they form a characteristic longitudinal lateral row that continues onto the tail. Ventral side bright white, sharply contrasting with the coloration of the dorsum.
Comparisons.
Mesalina
saudiarabica
sp. n.
can primarily be distinguished from other species of the complex by its genetic differentiation at both mtDNA and nDNA level. Genetic distances in the mtDNA genes are given above in the diagnosis and in
Table 1
. The differentiation in the nDNA is obvious from the allele networks (
Fig. 2
) that show that the species does not share alleles of any gene with any other species.
Mesalina
saudiarabica
sp. n.
is also geographically isolated from the rest of the complex. The nearest localities of
M. bernoullii
lie
680 km
to the east or 860 to the north. Moreover it can be distinguished morphologically from
M. microlepis
by having 1–2 large semitransparent scales in the lower eyelid (several roughly equal semitransparent scales in the latter), lower number of collar plates (6–8 vs. 10–13;
t
test t = 5.01,
p
<0.001), lower number of dorsal scales (41–42 vs. 48– 61;
t
test t = 3.78,
p
<0.005), lower number of preanal scales (2–3 vs. 4;
t
test t = 8.14,
p
<0.001), and lower number of femoral pores in males (12–13 vs. 15–20;
t
test t = 5.12,
p
<0.001). The lower number of collar plates differentiates the new species also from
M
. bernoullii
(6–8 vs. 8–13;
t
test t = 3.14,
p
<0.005) and
M
. brevirostris
s. s.
, although not significantly after Bonferroni correction (6–8 vs. 8–10;
t
test t = 2.67,
p
<0.05) (
Tables
S
4
,
S
5
).
Description of the
holotype
. Subadult male (Fig. 4). Body slender, slightly depressed; snout short with prominent elevated nostrils, occipital shield absent, two large semitransparent scales in the lower eyelid; snoutvent length 31.0 mm, tail length 56.0 mm, head length 6.9/8.0 mm (to the anterior/posterior edge of the ear), head width
5.1 mm
, head depth
3.5 mm
. Upper labials (left/right) anterior the centre of eye 5/5 (smaller fifth upper labial separating the subocular from the mouth included), gulars 25, plates in collar 8, dorsals across midbody 41, ventrals across belly 12, transverse rows of ventrals 32, preanals in straight median series 3, subdigital lamellae 23/24, femoral pores 13/13. In alcohol, dorsum light brown with a pattern of small whitish and larger dark brown spots. Most of the whitish spots are not edged with dark brown color. The spots are arranged in more or less regular longitudinal rows. The dark brown spots predominate on flanks where they form a longitudinal lateral row that continues onto the tail. The lateral row of large brown spots is bordered by a dorsolateral row of small whitish spots and a narrow inconspicuous whitish ventrolateral line. Ventral side white.
Variation. The
paratype
generally corresponds in morphology with the
holotype
. Apart from several minute differences in scalation described in the paragraph Diagnosis it differs also in the presence of a distinct occipital shield and the presence of only one large semitransparent scale in the lower eyelid. The
paratype
lacks the left hind leg and the tail.
Distribution and ecology. All eight so far known localities of
Mesalina
saudiarabica
sp. n.
as well as its range of suitable conditions are located in centralwestern
Saudi Arabia
on the central plateau of the Arabian Peninsula at elevations of
900–1050 m
a.s.l. The region is characterized by hot and semiarid to arid climate with mean summer temperatures up to 30°C and mean annual precipitation of
50–100 mm
with rain typically occurring between March and May (Edgell 2006; Shobrak 2011). The terrain consists mostly of flat gravel plains known as ‘regs’, occasionally intersected by dry sandy wadis and dominated by sparse vegetation of perennial grasses including
Stipagrostis
sp.,
Panicum
turgidum
and
Lasiurus
scindicus
and small trees, mainly
Acacia
sp. (Mandaville 1990).
The
type
locality is located in the Mahazat asSayd Nature Reserve, approximately
170 km
ENE of Taif. The reserve is Saudi Arabia’s only completely fenced wildlife reserve and is a reintroduction site for MacQueen’s bustard (
Chlamydotis
macqueenii
), Arabian oryx (
Oryx leucoryx
) and Sand gazelles (
Gazella
subgutturosa
). At the reserve,
Acacia
tortilis
is the most common tree species,
Fagonia
indica
and
Indigofera
spinosa
are the most common herbs, and
Panicum
turgidum
and
Stipagrostis
spp. are the prevailing grasses.
Etymology. The specific epithet is derived from the name of the country where the species occurs. Proposed English name – Arabian shortnosed desert lizard.