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Macrobenthos Macrobenthos M. Butler, M. C. Miller, and S. Mozley INTRODUCTION TO ARCTIC BENTHOS The larger benthic animals, which are mostly insects although snails and worms do occur, are extremely important to the ecology of the ponds . Not only are their biomass and productivity larger than those of any other group of animals, but they also continually affect the structure of the sediments by their feeding and burrowing . In one experiment, for example, chironomids (at twice natural density) mixed and moved the sediments so much that sand grains placed at the surface were completely covered in 10 days and 10% had reached a depth of 3 cm . This movement, plus the circulation of water through their burrows, mixes oxygen into the sediments and promotes the exchange of dissolved material . The benthic animals also eat zooplankton, microfauna, benthic algae, and each other and are, in turn, a food for insectivorous birds and even zooplankton . Macrobenthos of the ponds were investigated in two periods by different investigators . In 1971 and 1972, D . M . Bierle collected extensive data on distribution and dynamics, population densities, feeding, growth, respiration, and production . Most data were collected in Pond J and on larvae of the genus Chironomus, but comparative estimates of total benthic secondary production were obtained for three other ponds, B, D, and E. Unfortunately, the abundance of small forms and early larval instars was underestimated and species were not adequately distinguished, with the result that many population data represented mixtures of animals with different life cycles . Some estimates were too low, such as larval densities, spatial variance of larval populations, densities of emerging adults, and length of larval lifespans, but estimates of larval and adult biomass seem to have been too high. On the other hand, many data collected in the earlier years on respiration, growth, larval feeding, trends in directly measured total biomass, and occurrence of benthos or their terrestrial life stages in bird guts were apparently valid . The present authors collected further data on species composition, density, age structure, emergence, and within- and between-pond variance in 1975, 1976, and 1977. A detailed description of these studies will be included in a *S. Mozley and M . Butler 297 298 M. Butler et al. Ph . D. thesis at the University of Michigan by M . Butler. We are grateful for Dr. Bierle's permission to use his data and samples in this account . Benthic Studies . Research on arctic macrobenthos has been sparse and primarily descriptive . The earliest detailed work was Andersen's (1946) study of several shallow lakes in Greenland . He pointed out the relatively great importance of soft-bottom benthos and particularly the Chironomidae in northern waters . He observed large populations but comparatively few species, primarily Chironomidae in the groups Tanytarsini, Chironomus, Phaenopsectra, Psectrocladius, Cricotopus, Corynoneura, and Procladius, and related their abundance and distribution to environmental features such as dissolved oxygen over winter, macrophytes, development of dissolved H 2S, and depth of water . He mentioned also Acari, Turbellaria and Lepidurus . He believed most Chironomidae to have one-year life cycles. A brief survey of lakes in the mid-1950's (Livingstone et al . 1958) also emphasized the low diversity and moderate standing crops of benthos and littoral insects and the importance of chironomids in the arctic standing waters, but provided little direct data on their ecology . They found an unusual Tanytarsini preserved in lake sediments, Corynocera (or Dryadotanytarsus) and remarked on the very high density of chironomid remains in some lakes . Their higher standing crops of macrobenthos were similar to those in Barrow ponds . The classic studies of Scholander et al . (1953) on physiology of arctic animals showed how Chironomus larvae from Barrow ponds were able to freeze and return to activity with the subsequent thaw . Oliver (1968) and Danks and Oliver (1972), working in the Lake Hazen area of extreme northeast Canada, showed that metamorphosis and emergence of Chironomidae adults were extremely synchronous but varied between ponds and to a lesser extent between habitats within ponds . The Chironomidae in those studies were in the same genera but were often different species than the Barrow midges . Oliver (1968) summarized a number of other features of arctic Chironomidae, including the occurrence of copulation on surfaces rather than in aerial swarms in several species, and mechanisms controlling emergence, oogenesis, and long life cycles . Oliver found that water temperature regulated the time of emergence in shallow environments . In Cape Thompson ponds (Watson et al . 1966a), macrobenthos densities were high (38,000 m -2) and chironomids were the most numerous taxon, followed by oligochaetes . Turbellaria, Polychaeta, Isopoda, Amphipoda, Acari, and the insect orders Plecoptera, Trichoptera, Ephemeroptera, and Coleoptera were well represented, especially in beds of the macrophyte Arctophila . Unfortunately they were unable to give more taxonomic detail . Macrobenthos 299 Most species lists of arctic or cold-subarctic insects are based on collections of adults and do not discriminate among terrestrial, semi- terrestrial and truly aquatic forms (Watson et al . 1966b; McAlpine 1965 ; Rickard and Harmston 1972). The benthic studies of Holmquist (1973) have focused on coastal lakes and the presence or absence of marine relicts among the macrobenthos . The Char Lake IBP study emphasized dynamics, production, and controls on macrobenthos, particularly Chironomidae (Welch 1973, 1976, Welch and Kalff 1974, Lasenby and Langford 1972) . Chironomids shared dominance with Mysis relicta in Char Lake but there were few species ; Orthocladiinae were dominant and the variable ice cover occasionally interfered with annual reproduction . Char Lake conditions represented a more extreme polar environment than those in Barrow. Secondary productivity in Char Lake was extremely low because of low primary productivity (Welch 1976) . It was also found that chironomids have longer lifespans in the Arctic than in temperate latitudes, two or three years in the Char Lake species . Barrow thaw ponds contrast with other investigated arctic habitats in several ways . The species of Chironomidae are somewhat different from those in other areas, although genera are similar. Some of the statements and suppositions of Livingstone et al . (1958) about the absence of dytiscid beetles and widespread dominance of Corynocera in the Arctic are not borne out . Lifespans of benthic animals are longer than earlier authors report, and growth is slow in terms of calendar years . However, development is rapid in view of the low summer temperatures and when compared with daily rates at lower latitudes . A sum of less than 1 year of ice-free days is required for maturation in all the smaller species . Species Composition The principal organisms were collected from core samples of sediments as well as by sweeps of emergent vegetation, by submerged pitfalls, by quantitative entrapment of emerging adult insects, and by some aerial netting. The list (Table 7-1) is remarkable in several ways . Virtually every taxon contains fewer species than would be expected in a .temperate pond and many taxa are missing altogether . There are no Ephemeroptera, Odonata, Hemiptera or Megaloptera, no Hirudinea, lumbriculid or naidid Oligochaeta, no Pelecypoda, no Amphipoda, Isopoda, or Decapoda . Mosquitoes are absent in contrast to their notorious abundances in thaw ponds and pools farther inland. Indeed, there are few dipterans of any sort besides the abundant and (comparatively) diverse Chironomidae . The numbers of species of Coleoptera, Gastropoda, Trichoptera, and Acari are all lower than in temperate waters. Even Chironomidae are not as rich in species here as they are in permanent temperate ponds . 300 M. Butler et al . TABLE 7-1 Macrobenthos Collected From Tundra Ponds Near Barrow, Alaska Turbellaria 1 sp. A nnelida Oligochaeta Tubificidae Tubifex sp. Enchytraeidae Propappus sp . Hydracarina Lebertiidae Lebertia sp. Gastropoda Physidae Physa sp. Insecta Plecoptera Nemoura arctica Coleoptera Dytiscidae Agabus sp. Hydroporus sp. Diptera Chironomidae Podonominae Trichotanypus alaskensis Tanypodinae Procladius prolongatus P. vesus Derotanypus alaskensis D. aclines Chironominae Chironomini Chironomus pilicornis C. hyperboreus C. riparius C. (Camptochironomus) grandivalva Stictochironomus sp . Dicrotendipes lobiger Cryptochironomus sp . Tanytarsini Tanytarsus inequails T. gregarius gr. sp . 2 Paratanytarsus penicillatus Constempellina sp . Cladotanytarsus sp . Orthocladiinae Orthocladini Psectrocladius psilopterus gr. spp. (at least 2) P. dilatatus gr. sp . Cricotopus tibialis C.nr. perniger Cricotopus sp . 3 Cricotopus sp . 4 Orthocladius (Eudactylocladius) sp . O. (Pogonocladius) sp . Macrobenthos 301 TABLE 7-1 (Continued) Metriocnemini Corynoneura spp. (several) Metriocnemus spp. (at least 2) Pseudosmittia gr. sp. Parakiefferiella gracillima P. nigra Limnophyes sp. Lapposmittia sp . Mesosmittia sp . Bryophaenocladius sp . Individual species of animals tend to occur primarily either in the soft sediments in pond centers or among stems and leaves of Carex and Arctophila around the edge. These two habitats and the animals associated with them are represented in Figures 7-1 and 7-2 . In the centers, Chironomidae, led by Chironomus sp ., Tanytarsini
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