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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

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