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How aquatic insects live in cold climates

Published online by Cambridge University Press:  02 April 2012

H. V. Danks
H. V. Danks
Affiliation:
Biological Survey of Canada (Terrestrial Arthropods), Canadian Museum of Nature, P.O. Box 3443, Station D, Ottawa, Ontario, Canada K1P 6P4 (e-mail: hdanks@mus-nature.ca)
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Abstract

In cold climates most aquatic habitats are frozen for many months. Nevertheless, even in such regions the conditions in different types of habitat, in different parts of one habitat, and from one year to the next can vary considerably; some water bodies even allow winter growth. Winter cold and ice provide challenges for aquatic insects, but so do high spring flows, short, cool summers, and unpredictable conditions. General adaptations to cope with these constraints, depending on species and habitat, include the use of widely available foods, increased food range, prolonged development (including development lasting more than one year per generation), programmed life cycles with diapause and other responses to environmental cues (often enforcing strict univoltinism), and staggered development. Winter conditions may be anticipated not only by diapause and related responses but also by movement for the winter to terrestrial habitats, to less severe aquatic habitats, or to different parts of the same habitat, and by construction of shelters. Winter itself is met by various types of cold hardiness, including tolerance of freezing in at least some species, especially chironomid midges, and supercooling even when surrounded by ice in others. Special cocoons provide protection in some species. A few species move during winter or resist anoxia beneath ice. Spring challenges of high flows and ice scour may be withstood or avoided by wintering in less severe habitats, penetrating the substrate, or delaying activity until after peak flow. However, where possible species emerge early in the spring to compensate for the shortness of the summer season, a trait enhanced (at least in some lentic habitats) by choosing overwintering sites that warm up first in spring. Relatively low summer temperatures are offset by development at low temperatures, by selection of warm habitats and microhabitats, and in adults by thermoregulation and modified mating activity. Notwithstanding the many abiotic constraints in cold climates, aquatic communities are relatively diverse, though dominated by taxa that combine traits such as cold adaptation with use of the habitats and foods that are most widely available and most favourable. Consequently, except in the most severe habitats, food chains and community structure are complex even at high latitudes and elevations, including many links between aquatic and terrestrial habitats. Despite the complex involvement of aquatic insects in these cold-climate ecosystems, we know relatively little about the physiological and biochemical basis of their cold hardiness and its relationship to habitat conditions, especially compared with information about terrestrial species from the same regions.

Résumé

Dans les climats froids, la plupart des habitats aquatiques sont recouverts de glace pendant plusieurs mois. Néanmoins, dans ces régions, la gamme des conditions dans les divers types d'habitats, dans les diverses parties d'un même habitat et d'une année à l'autre peut varier considérablement; certains milieux aquatiques permettent même de la croissance pendant l'hiver. Le froid et la glace de l'hiver posent des problèmes aux insectes aquatiques, mais c'est le cas aussi des forts débits du printemps, des étés courts et frais et des conditions imprévisibles. Les adaptations générales pour faire face à ces contraintes comprennent, selon l'espèce et l'habitat, l'utilisation de nourriture largement disponible, l'accroissement de l'éventail alimentaire, la prolongation du développement et en particulier, la durée de plus d'un an par génération, les cycles biologiques programmés avec présence de diapause ou d'autre réaction aux signaux environnementaux (maintenant souvent un univoltinisme strict) et un développement étalé. Les conditions hivernales peuvent être anticipées non seulement par la diapause et les autres réactions de même type, mais aussi par un déplacement vers les habitats terrestres pour l'hiver, vers des habitats aquatiques moins extrêmes ou vers des sections différentes du même habitat et par la construction de refuges. L'hiver lui-même est contré par divers types de résistance au froid, y compris la tolérance au gel au moins chez certaines espèces—particulièrement chez les moucherons chironomidés—et, chez d'autres, la surfusion même lorsqu'elles sont entourées de glace. Des cocons spéciaux fournissent une protection à certaines espèces. Quelques espèces se déplacent durant l'hiver ou résistent à l'anoxie sous la glace. Les problèmes du printemps, les forts débits et l'affouillement par la glace, peuvent être endurés ou évités par le passage de l'hiver dans des habitats moins rigoureux, l'enfouissement dans le substrat ou le report des activités après le débit maximal. Cependant, lorsque c'est possible, les espèces émergent tôt au printemps pour compenser la brièveté de la saison estivale, une caractéristique qui est favorisée (au moins dans certains habitats lénitiques) par le choix d'habitats d'hivernage qui se réchauffent les premiers au printemps. Les températures relativement basses de l'été sont compensées par un développement à basse température, par la sélection comportementale d'habitats et de microhabitats chauds et, chez les adultes, par la thermorégulation et la modification des activités reproductrices. En dépit des nombreuses contraintes abiotiques des climats froids, les communautés aquatiques y sont relativement diversifiées, bien que dominées par des taxons qui possèdent des combinaisons de caractères, tels que l'adaptation au froid et l'utilisation des habitats et des nourritures qui sont les plus disponibles et les plus avantageux. En conséquence, à l'exception des habitats les plus rigoureux, les chaînes alimentaires et la structure des communautés sont complexes même aux latitudes et aux altitudes élevées et elles comprennent de nombreux liens entre les habitats aquatiques et terrestres. Malgré le rôle complexe des insectes aquatiques dans ces écosystèmes de climat froid, on connaît relativement peu de choses sur les bases physiologiques et biochimiques de leur résistance au froid et de ses relations avec les conditions de l'habitat; cela est d'autant plus vrai si on fait des comparaisons avec ce qu'on sait des espèces terrestres des mêmes régions.

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Articles
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The Canadian Entomologist , Volume 139 , Issue 4 , August 2007 , pp. 443 - 471
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Copyright © Entomological Society of Canada 2007

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