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systematics of snakes of the dipsas oreas complex - BioOne

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122 Bulletin Museum <strong>of</strong> Comparative Zoology, Vol. 158, No. 3<br />

and attempting to paint a scenario <strong>of</strong> what<br />

seems to occur. At <strong>the</strong> Río Zaña site, eggs<br />

<strong>of</strong> D. <strong>oreas</strong> with well developed embryos<br />

were discovered in mid-June in a well-secluded,<br />

relatively cool location deep within<br />

a crevice (details reported above). The advanced<br />

developmental stages <strong>of</strong> two embryos<br />

suggest that <strong>the</strong> eggs were laid some<br />

time prior to <strong>the</strong>ir discovery in mid-June.<br />

This observation, combined with <strong>the</strong> observed<br />

aggregation <strong>of</strong> several adult males<br />

and an adult female in late January, suggests<br />

that a mating period in late January<br />

may not be unreasonable. Moreover, <strong>the</strong><br />

condition <strong>of</strong> <strong>the</strong> oviducts in <strong>the</strong> female and<br />

<strong>the</strong> vasa deferentia in <strong>the</strong> males suggest<br />

that <strong>the</strong>se individuals were reproductively<br />

competent. The delay between <strong>the</strong> presumed<br />

mating period and egg laying would<br />

give ample time for fertilization and yolking<br />

<strong>of</strong> <strong>the</strong> ova.<br />

In a related colubrid, Sibynomorphus<br />

ventrimaculatus, mating apparently takes<br />

place several months (perhaps as many as<br />

seven) before vitellogenesis commences<br />

and egg-laying occurs (Cechin and Oliveira,<br />

2003). A similar sequence in Dipsas<br />

<strong>oreas</strong> would explain <strong>the</strong> small follicles in<br />

<strong>the</strong> female described above well before<br />

<strong>the</strong> inferred oviposition season. The strong<br />

seasonal activity pattern <strong>of</strong> D. <strong>oreas</strong> at <strong>the</strong><br />

Río Zaña Study Site, as well as <strong>the</strong> few<br />

observations on eggs and hatchings, imply<br />

that reproduction is strongly seasonal in<br />

this species, at least in <strong>the</strong> sou<strong>the</strong>rn part<br />

<strong>of</strong> its range. O<strong>the</strong>r species <strong>of</strong> Dipsas that<br />

inhabit less seasonal environments <strong>of</strong><br />

Amazonia or <strong>the</strong> Atlantic Forest <strong>of</strong> sou<strong>the</strong>astern<br />

Brazil are apparently aseasonal in<br />

<strong>the</strong>ir reproductive patterns (Zug et al.<br />

[1979] for D. catesbyi; Porto and Fernandes<br />

[1996] for D. neivai), and some o<strong>the</strong>r<br />

members <strong>of</strong> <strong>the</strong> Dipsadini have extended<br />

periods <strong>of</strong> oviposition (e.g., two species <strong>of</strong><br />

Sibynomorphus; Laporta-Ferreira et al.,<br />

1986).<br />

Aggregation behavior is best known in<br />

north temperate species <strong>of</strong> natricine colubrids<br />

and viperids, although scattered observations<br />

<strong>of</strong> scolecophidians, elapids, and<br />

o<strong>the</strong>r groups <strong>of</strong> colubrids suggest that aggregation<br />

behavior is taxonomically widespread.<br />

Aggregations have previously been<br />

reported in three o<strong>the</strong>r species <strong>of</strong> Neotropical<br />

colubrids. An aggregation <strong>of</strong><br />

Rhadinaea calligaster (Myers, 1974: 221)<br />

in a pile <strong>of</strong> decomposing thatch consisted<br />

<strong>of</strong> six females and a male. Females were<br />

apparently attracted to <strong>the</strong> site for oviposition<br />

because a communal nest was in <strong>the</strong><br />

same pile <strong>of</strong> debris. The o<strong>the</strong>r cases involve<br />

aggregations for mating functions in<br />

<strong>snakes</strong> <strong>of</strong> <strong>the</strong> clade Colubrinae (sensu Cadle,<br />

1984c, 1985): Drymobius margaritiferus<br />

(Campbell, 1998: 213) and Chironius<br />

flavolineatus (Feio et al., 1999). The observations<br />

reported here for Dipsas <strong>oreas</strong><br />

constitute <strong>the</strong> first report <strong>of</strong> aggregation<br />

behavior in any species <strong>of</strong> Dipsadini (Dipsas,<br />

Sibon, Tropido<strong>dipsas</strong>, Sibynomorphus)<br />

and one <strong>of</strong> only three cases <strong>of</strong> mating<br />

aggregations in Neotropical colubrids.<br />

Because communal nests are now known<br />

in two distantly related members <strong>of</strong> <strong>the</strong><br />

larger Neotropical clade Dipsadinae (sensu<br />

Cadle, 1984b; Zaher, 1999), D. <strong>oreas</strong> and<br />

Rhadinaea calligaster, aggregation behavior<br />

for oviposition may be widespread<br />

within this group if particular oviposition<br />

sites <strong>of</strong>fer favorable conditions for egg development.<br />

TAXONOMIC AND GEOGRAPHICAL<br />

NOTES ON DIPSAS GRACILIS,<br />

D. LATIFASCIATUS, AND<br />

D. LATIFRONTALIS<br />

While studying <strong>the</strong> <strong>systematics</strong> <strong>of</strong> species<br />

in <strong>the</strong> Dipsas <strong>oreas</strong> <strong>complex</strong> and o<strong>the</strong>rs<br />

in western Ecuador and Peru (e.g., Cadle<br />

and Myers, 2003) I had occasion to<br />

examine specimens <strong>of</strong> o<strong>the</strong>r species reported<br />

from <strong>the</strong> eastern and western versants<br />

<strong>of</strong> <strong>the</strong> Andes in this region. In a few<br />

cases, <strong>the</strong> specimens examined amplified<br />

distributional knowledge <strong>of</strong> species or<br />

raised questions concerning systematic<br />

and nomenclatural issues. I here comment<br />

on <strong>the</strong>se points for three species. Dipsas<br />

gracilis has been well known from <strong>the</strong><br />

western lowlands <strong>of</strong> Ecuador, but new in-

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