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A Review of the North American Freshwater Snail Genus Pyrgulopsis

A Review of the North American Freshwater Snail Genus Pyrgulopsis

A Review of the North American Freshwater Snail Genus Pyrgulopsis

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A <strong>Review</strong> <strong>of</strong> <strong>the</strong> <strong>North</strong> <strong>American</strong> <strong>Freshwater</strong><br />

<strong>Snail</strong> <strong>Genus</strong> <strong>Pyrgulopsis</strong> (Hydrobiidae)<br />

Introduction<br />

Prosobranch snails <strong>of</strong> <strong>the</strong> genus <strong>Pyrgulopsis</strong> are widespread<br />

in <strong>the</strong> United States (extending into sou<strong>the</strong>rn Canada) and<br />

nor<strong>the</strong>rn Mexico, where <strong>the</strong>y occur in lakes, springs, seeps,<br />

marshes, and diverse lotic waters. These small (about 1-8 mm<br />

shell length) animals <strong>of</strong>ten are found in densities exceeding<br />

>1000/m 2 (for example see Noel, 1954), comprising a<br />

prominent element <strong>of</strong> <strong>the</strong> benthic epifauna. <strong>Pyrgulopsis</strong>, as<br />

recognized herein, is <strong>the</strong> second most diverse genus <strong>of</strong> <strong>North</strong><br />

<strong>American</strong> freshwater gastropods (<strong>the</strong> pleurocerid Elimia ranking<br />

as first) and includes many narrowly localized species.<br />

Numerous undescribed species also are known, and large<br />

physiographic regions still await exploration by those interested<br />

in <strong>the</strong>se easily overlooked organisms. These snails are<br />

beginning to draw <strong>the</strong> attention <strong>of</strong> wildlife agencies and groups<br />

seeking to preserve and manage <strong>North</strong> <strong>American</strong> aquatic<br />

ecosystems: over half (52%) <strong>of</strong> <strong>the</strong> <strong>Pyrgulopsis</strong> species<br />

recognized herein currently are candidates for addition to <strong>the</strong><br />

Federal List <strong>of</strong> Threatened and Endangered Species (USDI,<br />

1991a) and three snails (from New Mexico and Idaho) recently<br />

were listed as endangered (USDI, 1991b, 1992,1993). At least<br />

three members <strong>of</strong> <strong>the</strong> genus have gone extinct since <strong>the</strong>ir<br />

original description during <strong>the</strong> early part <strong>of</strong> this century.<br />

Call and Pilsbry (1886) erected <strong>Pyrgulopsis</strong> for four species<br />

from <strong>the</strong> United States having a small, fairly elongate shell<br />

whose single basal carina distinguished <strong>the</strong> group from<br />

multicarinate European Pyrgula. Ancey (1888) early reviewed<br />

<strong>Pyrgulopsis</strong>, and described 13 new Central <strong>American</strong> taxa, all<br />

<strong>of</strong> which were later transferred to <strong>the</strong> hydrobiid subfamily<br />

Cochliopinae, genus Pyrgophorus, by Hershler and Thompson<br />

(1992). Wenz (1926) also provided an early review <strong>of</strong> fossil<br />

species assigned to this genus. The concept <strong>of</strong> this group varied<br />

little since that time (even recent workers such as Burch (1982)<br />

and Taylor (1985) used <strong>Pyrgulopsis</strong> in <strong>the</strong> original sense) until<br />

Robert Hershler, Department <strong>of</strong> Invertebrate Zoology, National<br />

Museum <strong>of</strong> Natural History, Smithsonian Institution, Washington,<br />

D.C. 20560.<br />

Robert Hershler<br />

Hershler and Thompson (1987) redefined <strong>the</strong> genus in a<br />

broader fashion based on penial (ra<strong>the</strong>r than shell) characters.<br />

They consequently placed five o<strong>the</strong>r <strong>North</strong> <strong>American</strong> genuslevel<br />

taxa in synonomy with our group, which as a result<br />

swelled to 24 Recent species. The relationship between<br />

<strong>Pyrgulopsis</strong> (sensu Hershler and Thompson) and two similar<br />

genera recently described from <strong>the</strong> West (Taylor, 1987) has not<br />

been addressed in <strong>the</strong> literature.<br />

The scope and content <strong>of</strong> <strong>Pyrgulopsis</strong> remain incompletely<br />

known because <strong>the</strong> anatomy <strong>of</strong> relatively few <strong>North</strong> <strong>American</strong><br />

freshwater hydrobiids has been studied—this includes some <strong>of</strong><br />

<strong>the</strong> species allocated to <strong>the</strong> genus, on a provisional basis, by<br />

Hershler and Thompson (1987) and o<strong>the</strong>r, mostly western<br />

hydrobiids with simple, ovate-conic shells that were early<br />

described as members <strong>of</strong> "catch-all" genera such as Amnicola<br />

and Paludestrina.<br />

There has been no comprehensive review <strong>of</strong> <strong>the</strong> species <strong>of</strong><br />

<strong>Pyrgulopsis</strong> since that <strong>of</strong> Ancey (1888), at which time only two<br />

<strong>of</strong> <strong>the</strong> 65 species considered valid herein had been described.<br />

The present review is intended to provide nomenclatural<br />

stability for <strong>Pyrgulopsis</strong> and its Recent species, and is based on<br />

evaluation <strong>of</strong> available type and o<strong>the</strong>r museum material as well<br />

as anatomical study <strong>of</strong> almost all Recent <strong>North</strong> <strong>American</strong><br />

hydrobiid snails potentially referable to <strong>the</strong> genus.<br />

<strong>Pyrgulopsis</strong> species usually are readily distinguished from<br />

one ano<strong>the</strong>r by combinations <strong>of</strong> qualitative characters, and thus<br />

I have not ga<strong>the</strong>red and analyzed quantitative morphological<br />

data as part <strong>of</strong> this broad review. Note, however, that analysis<br />

<strong>of</strong> such data has been previously pursued with good results in<br />

studies <strong>of</strong> local <strong>Pyrgulopsis</strong> faunas (Hershler and Sada, 1987;<br />

Taylor, 1987; Hershler and Landye, 1988), and use <strong>of</strong> this and<br />

o<strong>the</strong>r means <strong>of</strong> comparing populations (i.e., biochemical) will<br />

be necessary to better resolve <strong>the</strong> lingering species-level<br />

problems in this group.<br />

Only nominate taxa are included in this review, as my<br />

intention is to provide herein a framework for future studies<br />

describing new species. Eastern <strong>American</strong> <strong>Pyrgulopsis</strong> were<br />

included despite <strong>the</strong> fact that most <strong>of</strong> <strong>the</strong>m recently were well

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