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Caddisflies of the Yukon - Department of Biological Sciences ...

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o<strong>the</strong>r reasons, most Palaearctic-East Beringian species <strong>of</strong> category III have functioned<br />

nei<strong>the</strong>r as aggressive colonists nor ecological generalists in North America; but quite clear<br />

is <strong>the</strong> contrast with Europe and Asia where most <strong>of</strong> <strong>the</strong>m are widely distributed, and<br />

apparently are successful generalists. It is one <strong>of</strong> <strong>the</strong> striking paradoxes <strong>of</strong> Beringian distributions<br />

that wide-ranging, and evidently competitively successful, Eurasian species remain<br />

confined to <strong>the</strong>ir East Beringian outpost, evidently unable to disperse much beyond <strong>the</strong>ir<br />

former glacial refuge. This distributional paradox can be added to <strong>the</strong> productivity paradox<br />

(e.g. Hopkins et al. 1982) as significant questions about <strong>the</strong> biological history <strong>of</strong> Beringia.<br />

This issue focusses on species <strong>of</strong> category III because <strong>the</strong>y may differ from category II<br />

essentially in lacking <strong>the</strong> competitive ecological edge required in new communities;<br />

Palaearctic species having that competitive edge now meet <strong>the</strong> distributional criterion <strong>of</strong><br />

category II.<br />

For several <strong>of</strong> <strong>the</strong>m (Ylodes kaszabi, Arctopora trimaculata, Grammotaulius signatipennis,<br />

Limnephilus stigma, and Agrypnia obsoleta), sister-group relationships suggest<br />

intercontinental vicariance, perhaps during <strong>the</strong> Pliocene or earlier, followed later by dispersal<br />

<strong>of</strong> <strong>the</strong> Palaearctic form to East Beringia, probably across <strong>the</strong> Pleistocene land bridge.<br />

In Agraylea cognatella and Limnephilus fenestratus, morphological similarity to <strong>the</strong>ir<br />

respective Nearctic sister species is so close that dichotomy during <strong>the</strong> Pleistocene could be<br />

reasonably inferred. Subdivision <strong>of</strong> <strong>the</strong> range by glacial ice seems likely, with <strong>the</strong> Nearctic<br />

sister species originating in isolation to <strong>the</strong> south <strong>of</strong> <strong>the</strong> glaciers.<br />

Spicipalpia<br />

Hydroptilidae<br />

Agraylea cognatella McLachlan<br />

Rhyacophilidae<br />

Rhyacophila mongolica Schmid, Arefina<br />

and Levanidova<br />

Integripalpia<br />

Leptoceridae<br />

Ylodes kaszabi Schmid<br />

Hydroptilidae<br />

Agraylea cognatella McLachlan (4)<br />

The Palaearctic range <strong>of</strong> A. cognatella appears to be circumscribed by A. multipunctata<br />

Curtis which is widely distributed through Europe and western Asia (BotojAneanu and<br />

Levanidova 1988); in North America, A. cognatella is evidently circumscribed to <strong>the</strong> south<br />

<strong>of</strong> Beringia, not by A. multipunctata as has been <strong>the</strong> traditional interpretation (e.g. Ross<br />

1944), but by a sibling species A. fraterna Banks apparently now widely distributed on this<br />

continent (Vineyard and Wiggins in prep.). It seems reasonable to infer from present<br />

evidence that A. cognatella is a Palaearctic-East Beringian species now confined in North<br />

America to <strong>the</strong> refugium, and that A. fraterna passed <strong>the</strong> glacial period to <strong>the</strong> south <strong>of</strong> <strong>the</strong><br />

ice where it may have arisen.<br />

Rhyacophilidae<br />

Rhyacophila mongolica Schmid, Arefina and Levanidova (14)<br />

<strong>Caddisflies</strong> <strong>of</strong> <strong>the</strong> <strong>Yukon</strong> 841<br />

Limnephilidae<br />

Arctopora trimaculata (Zetterstedt)<br />

Dicosmoecus obscuripennis Banks<br />

Grammotaulius signatipennis McLachlan<br />

Grensia praeterita (Walker)<br />

Limnephilus diphyes McLachlan<br />

Limnephilus fenestratus (Zetterstedt)<br />

Limnephilus stigma Curtis<br />

Phryganeidae<br />

Agrypnia obsoleta (Hagen)<br />

Agrypnia sahlbergi (McLachlan)<br />

Oligotricha lapponica (Hagen)

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