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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 75<br />

tifications in some older literature (e.g.,<br />

Boulenger, 1896), has led to considerable<br />

uncertainty concerning differential characteristics<br />

<strong>of</strong> <strong>the</strong>se nominal taxa and <strong>the</strong>ir<br />

respective distributions.<br />

Peters’ (1960a) confusion <strong>of</strong> two Ecuadorian<br />

species under <strong>the</strong> name Dipsas ellipsifera<br />

affected his assessment <strong>of</strong> variation<br />

within this species. For example, his<br />

ventral and subcaudal counts for males<br />

and females <strong>of</strong> ‘‘Dipsas ellipsifera’’ (Peters,<br />

1960a: 87) are exceptionally broad for a<br />

species <strong>of</strong> snake having a small range in<br />

western Ecuador. None<strong>the</strong>less, Peters<br />

(1960a: 91) recognized a distinctive geographic<br />

pattern to <strong>the</strong> variation within his<br />

concept <strong>of</strong> D. ellipsifera:<br />

The material available can be divided into two<br />

groups as to provenance, <strong>the</strong> first coming from <strong>the</strong><br />

Río Mira drainage and referred to as <strong>the</strong> ‘‘typical<br />

population,’’ and <strong>the</strong> o<strong>the</strong>r from <strong>the</strong> western slopes<br />

<strong>of</strong> <strong>the</strong> Andes in Ecuador, to <strong>the</strong> south <strong>of</strong> <strong>the</strong> Río<br />

Mira.... Although <strong>the</strong> indicated differences between<br />

<strong>the</strong>se two populations suggest subspecific<br />

status, I am not assigning <strong>the</strong>m a name, because<br />

no satisfactory holotype is available.... The most<br />

striking differences between <strong>the</strong> two populations<br />

are in counts involving body segments.<br />

Peters goes on to detail differences between<br />

<strong>the</strong> Río Mira specimens and <strong>the</strong><br />

o<strong>the</strong>rs when samples are segregated by<br />

sex. However, he did not make <strong>the</strong> conceptual<br />

leap to recognizing distinct species<br />

on <strong>the</strong> basis <strong>of</strong> <strong>the</strong>se populational differences.<br />

Nor did he recognize that <strong>the</strong> ‘‘o<strong>the</strong>r’’<br />

population from <strong>the</strong> western slopes <strong>of</strong><br />

<strong>the</strong> Andes represented Dipsas elegans, believing<br />

as he did that D. elegans was a<br />

Mexican species.<br />

K<strong>of</strong>ron (1982) clarified much <strong>of</strong> <strong>the</strong><br />

confusion engendered by <strong>the</strong> uncertain origin<br />

<strong>of</strong> <strong>the</strong> holotype <strong>of</strong> Dipsas elegans. He<br />

summarized evidence that <strong>the</strong> holotype <strong>of</strong><br />

D. elegans came from western Ecuador on<br />

<strong>the</strong> basis <strong>of</strong> <strong>the</strong> known travels and contacts<br />

<strong>of</strong> Adolphe Boucard, who sold <strong>the</strong> specimen<br />

to <strong>the</strong> British Museum (K<strong>of</strong>ron, 1982:<br />

47). K<strong>of</strong>ron also reviewed <strong>the</strong> erroneous<br />

association <strong>of</strong> <strong>the</strong> name <strong>of</strong> <strong>the</strong> collector,<br />

François Sumichrast, who collected in <strong>the</strong><br />

vicinity <strong>of</strong> <strong>the</strong> Isthmus <strong>of</strong> Tehuantepec,<br />

with <strong>the</strong> type <strong>of</strong> D. elegans. The error was<br />

promulgated by Gün<strong>the</strong>r (1885–1902:<br />

141) and continued in checklists and faunal<br />

works (e.g., Peters, 1960a: 86; Smith<br />

and Taylor, 1945) until K<strong>of</strong>ron discovered<br />

<strong>the</strong> mistake. The unfortunate association<br />

<strong>of</strong> Sumichrast’s name with <strong>the</strong> specimen<br />

that ultimately became <strong>the</strong> holotype <strong>of</strong> D.<br />

elegans resulted in it being considered a<br />

Mexican species by Peters (1960a) and<br />

o<strong>the</strong>rs (e.g., Smith and Taylor, 1945).<br />

K<strong>of</strong>ron’s (1982) conclusion that <strong>the</strong> holotype<br />

<strong>of</strong> Dipsas elegans most likely came<br />

from Ecuador was a major step toward resolving<br />

<strong>the</strong> <strong>systematics</strong> <strong>of</strong> <strong>the</strong>se <strong>snakes</strong>.<br />

None<strong>the</strong>less, because <strong>the</strong> type specimens<br />

<strong>of</strong> D. ellipsifera and D. elegans have nearly<br />

identical color patterns, K<strong>of</strong>ron (1982: 48)<br />

synonymized D. ellipsifera with D. elegans,<br />

which is <strong>the</strong> earlier name. He did not thoroughly<br />

consider o<strong>the</strong>r character variation<br />

in available samples <strong>of</strong> <strong>the</strong>se <strong>snakes</strong>, particularly<br />

in <strong>the</strong> context <strong>of</strong> geographic origin<br />

and sex, even though some <strong>of</strong> <strong>the</strong>se patterns<br />

had already been elucidated by Peters<br />

(1960a; see previous quotation).<br />

Despite having examined much <strong>of</strong> <strong>the</strong><br />

same material as Peters, K<strong>of</strong>ron (1982)<br />

used Peters’ (1960a) data when comparing<br />

<strong>the</strong> scale counts <strong>of</strong> <strong>the</strong> holotype <strong>of</strong> Dipsas<br />

elegans with D. ellipsifera, apparently failing<br />

to realize that Peters had confused <strong>the</strong><br />

two species under <strong>the</strong> name ‘‘Dipsas ellipsifera’’<br />

and overlooking Peters’ more detailed<br />

discussion <strong>of</strong> geographic variation<br />

within his concept <strong>of</strong> D. ellipsifera: ‘‘The<br />

ventral and subcaudal counts <strong>of</strong> [<strong>the</strong> holotype<br />

<strong>of</strong>] D. elegans (183, 95) are within<br />

<strong>the</strong> ranges <strong>of</strong> 155–187 ventrals and 72–105<br />

subcaudals reported for D. ellipsifera by<br />

Peters (1960)’’ (K<strong>of</strong>ron, 1982: 47). Thus,<br />

K<strong>of</strong>ron (1982) accepted Peters’ (1960a)<br />

characterization <strong>of</strong> ‘‘Dipsas ellipsifera,’’<br />

which included specimens <strong>of</strong> D. elegans;<br />

not surprisingly, characteristics <strong>of</strong> <strong>the</strong> holotype<br />

<strong>of</strong> D. elegans conformed well to this<br />

composite taxon. K<strong>of</strong>ron did not reexamine<br />

<strong>the</strong> meristic data <strong>of</strong> specimens segregated<br />

by sex and locality and failed to no-

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