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

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814 G.B. Wiggins and C.R. Parker<br />

brown- and gray-winged specimens. Thus females can be identified only by <strong>the</strong> brown<br />

membrane <strong>of</strong> <strong>the</strong> forewings and by association with males.<br />

In North America material <strong>of</strong> <strong>the</strong> brown-winged form was examined from <strong>the</strong> <strong>Yukon</strong><br />

(10, 12, 17; ROME, SMDV), Alaska, and British Columbia; <strong>the</strong> gray-winged form was<br />

collected only in <strong>the</strong> <strong>Yukon</strong> (4, 10, 12; ROME). Adult specimens were collected along lakes<br />

and streams and on <strong>the</strong> banks <strong>of</strong> <strong>the</strong> <strong>Yukon</strong> River. However, we received a series <strong>of</strong> <strong>the</strong><br />

brown-winged form collected near St. Petersburg, Russia, by V.D. Ivanov; genitalic structure<br />

<strong>of</strong> males is entirely consistent with our Beringian variant (Fig. 4). Consequently, we infer<br />

that two morphs may occur sympatrically over much <strong>of</strong> <strong>the</strong> range attributed to C. nigronervosa,<br />

and perhaps <strong>the</strong> two differ in preferred habitat or life cycle. The issue has to be resolved<br />

by a worker with access to populations in Europe and Asia.<br />

Note 4. Mystacides sepulchralis (Walker) (60)<br />

Although this species is common and widely distributed over much <strong>of</strong> North America<br />

including <strong>the</strong> <strong>Yukon</strong> and Alaska (Yamamoto and Wiggins 1964), its status beyond North<br />

America requires clarification. Holarctic distribution was attributed to M. sepulchralis by<br />

Yamamoto and Ross (1966), even while recognizing as valid <strong>the</strong> Siberian species M. bifidus<br />

Martynov (1924b); supporting evidence was not given, and we know <strong>of</strong> none apart from a<br />

record from Siberia attributed initially to M. sepulchralis (Martynov 1910) but later assigned<br />

to M. bifidus (Martynov 1935).<br />

There is considerable variation in genitalic characters <strong>of</strong> M. sepulchralis. Through <strong>the</strong><br />

assistance <strong>of</strong> V.D. Ivanov, we have obtained information on genitalic characters <strong>of</strong> <strong>the</strong><br />

holotype <strong>of</strong> M. bifidus in <strong>the</strong> Zoological Institute, St. Petersburg; and we have examined<br />

o<strong>the</strong>r specimens, indicating that <strong>the</strong>re is at least some variation in this species, too. The two<br />

forms are very close, and clearly are sister taxa; but Martynov affirmed his view (Betten and<br />

Mosely 1940) that <strong>the</strong> Siberian M. bifidus Martynov and <strong>the</strong> North American M. sepulchralis<br />

(Walker) are separate species. We follow that interpretation here because small morphological<br />

differences in o<strong>the</strong>r species are used here as <strong>the</strong> basis for inferring separation during<br />

glacial periods. Detailed analysis <strong>of</strong> <strong>the</strong> variation in populations within each continent might<br />

reveal evidence <strong>of</strong> a Beringian interchange; whe<strong>the</strong>r such differences would confirm status<br />

as species or as intraspecific variants would emerge from <strong>the</strong> analysis. For <strong>the</strong> present, we<br />

interpret M. sepulchralis as a strictly North American species.<br />

Larvae <strong>of</strong> M. bifidus, for which we have no information, could be relevant to this<br />

question. In recent years, distinctive larvae in some populations <strong>of</strong> M. sepulchralis from<br />

eastern North America have been found with head markings <strong>of</strong> discrete spots similar to those<br />

in M. alafimbriata, and quite unlike <strong>the</strong> largely black heads characteristic <strong>of</strong> most sepulchralis<br />

populations in North America (cf. figs. 3 and 4, Yamamoto and Wiggins 1964).<br />

Association <strong>of</strong> <strong>the</strong>se larvae with adults confirms that <strong>the</strong>y are M. sepulchralis. This larval<br />

dimorphism could have been established in populations isolated during <strong>the</strong> glacial period to<br />

<strong>the</strong> sou<strong>the</strong>ast <strong>of</strong> <strong>the</strong> continental ice mass, although it does not appear to be reflected in<br />

characters <strong>of</strong> <strong>the</strong> adults.<br />

The sister species <strong>of</strong> M. sepulchralis/bifidus is M. alafimbriata, confined to western<br />

North America including <strong>the</strong> <strong>Yukon</strong> and Alaska (Yamamoto and Wiggins 1964). Mystacides<br />

alafimbriata and sepulchralis are only slightly different morphologically; and although <strong>the</strong>re<br />

appears to be little overlap in <strong>the</strong>ir ranges, <strong>the</strong> two have been taken in <strong>the</strong> same collections<br />

in a few localities, all in <strong>the</strong> <strong>Yukon</strong> and Alaska. Possibly M. alafimbriata was derived from<br />

populations isolated to <strong>the</strong> south <strong>of</strong> <strong>the</strong> Cordilleran glacier in <strong>the</strong> mountains <strong>of</strong> western North<br />

America. Outlying montane glaciers along <strong>the</strong> Rocky and Cascade Mountain ranges served

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