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Science of Water : Concepts and Applications

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192 The <strong>Science</strong> <strong>of</strong> <strong>Water</strong>: <strong>Concepts</strong> <strong>and</strong> <strong>Applications</strong><br />

that are hard <strong>and</strong> leathery <strong>and</strong> not used in fl ight; the membranous hindwings, which are used for<br />

fl ight, are concealed under the elytra when the organisms are at rest. Only 10% <strong>of</strong> the 350,000<br />

described species <strong>of</strong> beetles are aquatic.<br />

Beetles are holometabolous (i.e., they complete metamorphosis). Eggs <strong>of</strong> aquatic coleopterans<br />

hatch in 1 or 2 weeks, with diapause (i.e., no suspension <strong>of</strong> development) occurring rarely. Larvae<br />

undergo from three to eight molts. The pupal phase <strong>of</strong> all coleopterans is technically terrestrial,<br />

making this life stage <strong>of</strong> beetles the only one that has not successfully invaded the aquatic habitat.<br />

A few species have diapausing prepupae, but most complete transformation to adults in 2–3 weeks.<br />

Terrestrial adults <strong>of</strong> aquatic beetles are typically short-lived <strong>and</strong> sometimes nonfeeding, like those<br />

<strong>of</strong> the other orders <strong>of</strong> aquatic insects. The larvae <strong>of</strong> Coleoptera are morphologically <strong>and</strong> behaviorally<br />

different from the adults, <strong>and</strong> their diversity is high.<br />

Aquatic species occur in two major suborders, the Adephaga <strong>and</strong> the Polyphaga. Both<br />

larvae <strong>and</strong> adults <strong>of</strong> the six beetle families are aquatic. Dytiscidae (predaceous diving beetles),<br />

Elmidae (riffl e beetles), Gyrinidae (whirligig beetles), Halipidae (crawling water beetles),<br />

Hydrophilidae (water scavenger beetles), <strong>and</strong> Noteridae (burrowing water beetles). Five families—<br />

Chrysomelidae (leaf beetles), Limnichidae (marsh-loving beetles), Psephenidae (water pennies),<br />

Ptilodactylidae (toe-winged beetles), <strong>and</strong> scirtidae (marsh beetles)—have aquatic larvae <strong>and</strong><br />

terrestrial adults, as do most <strong>of</strong> the other orders <strong>of</strong> aquatic insects; adult limnichids, however,<br />

readily submerge when disturbed. Three families have species that are terrestrial as larvae <strong>and</strong><br />

aquatic as adults: Curculionidae (weevils), Dryopidae (long-toed water beetles), <strong>and</strong> Hydraenidae<br />

(moss beetles), a highly unusual combination among insects.<br />

√ Note: Because they provide a greater underst<strong>and</strong>ing <strong>of</strong> the condition <strong>of</strong> a freshwater body (i.e.,<br />

they are useful indicators <strong>of</strong> water quality), the focus <strong>of</strong> discussion here is on the riffl e beetle,<br />

water penny, <strong>and</strong> whirligig beetle.<br />

Riffl e beetle larvae (most commonly found in running waters, hence the name riffl e beetle) are up<br />

to 3/4 in. long (see Figure 6.24). Their bodies are not only long, but also hard, stiff, <strong>and</strong> segmented;<br />

they resemble small torpedoes <strong>and</strong> are usually grayish in color. They have six long segmented legs<br />

on the upper middle section <strong>of</strong> body; the back end has two tiny hooks <strong>and</strong> short hairs. Larvae may<br />

take 3 years to mature before they leave the water to form a pupa; adults return to the stream.<br />

Riffl e beetle adults are considered better indicators <strong>of</strong> water quality than larvae because they<br />

have been subjected to water quality conditions over a longer period. They walk very slowly under<br />

the water (on stream bottom), <strong>and</strong> do not swim on the surface. They have small oval-shaped bodies<br />

(see Figure 6.25) <strong>and</strong> are typically about 1/4 in. in length.<br />

Both adults <strong>and</strong> larvae <strong>of</strong> most species feed on fi ne detritus with associated microorganisms<br />

that are scraped from the substrate. Others may be xylophagous, that is, wood eating (e.g., Lara,<br />

Elmidae). Predators do not seem to include riffl e beetles in their diet, except perhaps for eggs, which<br />

are sometimes attacked by fl atworms.<br />

The adult water penny is inconspicuous; it resembles circular encrustations on rocks <strong>and</strong> is <strong>of</strong>ten<br />

found clinging tightly in a sucker-like fashion to the undersides <strong>of</strong> submerged rocks, where they<br />

feed on attached algae. The body is broad, slightly oval <strong>and</strong> fl at in shape, ranging from 4–6 mm<br />

(1/4 in.) in length. The body is covered with segmented plates <strong>and</strong> looks like a tiny round leaf<br />

FIGURE 6.24 Riffl e beetle larvae.

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