systematics of snakes of the dipsas oreas complex - BioOne
systematics of snakes of the dipsas oreas complex - BioOne
systematics of snakes of the dipsas oreas complex - BioOne
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Dipsas <strong>oreas</strong> Complex in Ecuador and Peru • Cadle 85<br />
vertical markings separated by a narrow<br />
whitish strip. In some specimens <strong>the</strong> contralateral<br />
markings meet in <strong>the</strong> vertebral<br />
region, forming more or less complete<br />
bands, although <strong>the</strong> vertebral region tends<br />
to be invested with dark pigment so that<br />
<strong>the</strong> pale central regions <strong>of</strong> <strong>the</strong> markings<br />
are interrupted. In o<strong>the</strong>rs (e.g., MCZ<br />
8431; Fig. 7), contralateral markings fail to<br />
meet middorsally, forming lateral bars with<br />
pale centers that are closed <strong>of</strong>f on <strong>the</strong>ir<br />
dorsal edges by black pigment. The markings<br />
have vertical edges and extend ventrally<br />
to <strong>the</strong> first scale row or <strong>the</strong> outer<br />
edge <strong>of</strong> <strong>the</strong> ventrals. Contralateral markings<br />
are frequently <strong>of</strong>fset virtually <strong>the</strong> entire<br />
length <strong>of</strong> <strong>the</strong> body (e.g., Fig. 7) or on<br />
<strong>the</strong> posterior body only (most specimens).<br />
The markings (i.e., each black–white–<br />
black triplet) are 5–6 scale rows in width<br />
anteriorly but narrow to about 3 rows by<br />
midbody. Interspaces anteriorly are narrower<br />
than <strong>the</strong> bands or bars (approximately<br />
3 scale rows), whereas <strong>the</strong>y are<br />
wider than <strong>the</strong> markings posteriorly (5–6<br />
scale rows). The venter is dull grayish<br />
brown with a dense covering <strong>of</strong> dark<br />
brown squarish markings that tend to be<br />
concentrated toward <strong>the</strong> outer edges <strong>of</strong><br />
<strong>the</strong> ventrals, sometimes forming longitudinal<br />
arrays (Figs. 2–4).<br />
The top and sides <strong>of</strong> <strong>the</strong> head are heavily<br />
marked with irregular dark markings on<br />
a pale brown ground color (Fig. 6). The<br />
markings are so extensive in some specimens<br />
that most <strong>of</strong> <strong>the</strong> top <strong>of</strong> <strong>the</strong> head is a<br />
solid dark brown, with just occasional light<br />
areas <strong>of</strong> <strong>the</strong> ground color showing through<br />
as fine reticulations or vermiform marks.<br />
Upper and lower labials are pale brown<br />
with dark brown spotting, but <strong>the</strong> dark<br />
stippling <strong>of</strong>ten is not especially concentrated<br />
along suture lines as in many species<br />
<strong>of</strong> <strong>snakes</strong>.<br />
Two small juveniles (MCZ 8431, 210<br />
mm SVL; BMNH 1946.1.21.27, 138 mm<br />
SVL) have patterns identical to those <strong>of</strong><br />
adults, but more contrasting (Fig. 7; K<strong>of</strong>ron,<br />
1982: fig. 1). The ground color is grayish<br />
white, and <strong>the</strong> dark markings are dark<br />
chocolate brown to blackish. The dorsal<br />
markings have pale centers and appear as<br />
blackish vertical bars (or narrow ellipses)<br />
enclosing whitish centers. Interspaces between<br />
<strong>the</strong> markings are whitish, but individual<br />
scales are speckled with tiny irregular<br />
dark brown flecks. The top <strong>of</strong> <strong>the</strong><br />
head <strong>of</strong> MCZ 8431 is whitish with heavy<br />
reticulations and irregular spots so that<br />
much <strong>of</strong> <strong>the</strong> top and sides <strong>of</strong> <strong>the</strong> head are<br />
dark. The gular regions and venter are<br />
whitish with a pattern <strong>of</strong> bold irregular<br />
spots and blotches; on <strong>the</strong> venter <strong>the</strong>se<br />
tend to form bold longitudinal streaks. If<br />
<strong>the</strong>se two specimens are representative <strong>of</strong><br />
very small individuals <strong>of</strong> Dipsas ellipsifera,<br />
this species appears already to have <strong>the</strong><br />
adult color pattern even as small juveniles,<br />
a contrast with <strong>the</strong> developmental pattern<br />
in D. elegans and D. <strong>oreas</strong> (see <strong>the</strong> following<br />
species accounts for discussion). In <strong>the</strong><br />
last two species, smallest juveniles have<br />
solid bands that develop pale centers during<br />
early juvenile ontogeny, thus acquiring<br />
<strong>the</strong> adult patterns sometime after hatching.<br />
Hemipenis<br />
The following description is based on<br />
<strong>the</strong> inverted organ <strong>of</strong> UMMZ 83697 examined<br />
in situ. The organ had previously<br />
been slit along its medial edge and <strong>the</strong> retractor<br />
muscle had been cut distal to its<br />
point <strong>of</strong> division.<br />
Total length <strong>of</strong> <strong>the</strong> organ is 18 mm and<br />
it is slightly bilobed distally. The retractor<br />
muscle is divided proximally for 4 mm.<br />
The sulcus spermaticus (in <strong>the</strong> lateral wall<br />
<strong>of</strong> <strong>the</strong> organ) divides 9 mm from <strong>the</strong> base<br />
<strong>of</strong> <strong>the</strong> organ within <strong>the</strong> capitulum; <strong>the</strong> tips<br />
<strong>of</strong> its branches end approximately 1 mm<br />
short <strong>of</strong> <strong>the</strong> distal tips <strong>of</strong> <strong>the</strong> lobes. The<br />
basal region <strong>of</strong> <strong>the</strong> organ is sparsely ornamented<br />
with minute spines. The midsection<br />
has a battery <strong>of</strong> enlarged hooked<br />
spines encircling <strong>the</strong> organ just proximal<br />
to <strong>the</strong> capitulum; <strong>the</strong> battery is about 3<br />
spines across around <strong>the</strong> entire organ. The<br />
capitulum is set <strong>of</strong>f by a distinct overhang<br />
and is completely ornamented distally with