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Biogeography and Physiological Adaptations of the Brine Fly Genus Ephydra (Diptera: Ephydridae) in Saline Waters of the Great Basin

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Biogeography and Physiological Adaptations of the Brine Fly Genus Ephydra (Diptera: Ephydridae) in Saline Waters of the Great Basin Great Basin Naturalist Volume 59 Number 2 Article 3 4-30-1999 Biogeography and physiological adaptations of the brine fly genus Ephydra (Diptera: Ephydridae) in saline waters of the Great Basin David B. Herbst University of California, Mammoth Lakes and University of California, Santa Barbara Follow this and additional works at: https://scholarsarchive.byu.edu/gbn Recommended Citation Herbst, David B. (1999) "Biogeography and physiological adaptations of the brine fly genus Ephydra (Diptera: Ephydridae) in saline waters of the Great Basin," Great Basin Naturalist: Vol. 59 : No. 2 , Article 3. Available at: https://scholarsarchive.byu.edu/gbn/vol59/iss2/3 This Article is brought to you for free and open access by the Western North American Naturalist Publications at BYU ScholarsArchive. It has been accepted for inclusion in Great Basin Naturalist by an authorized editor of BYU ScholarsArchive. For more information, please contact [email protected], [email protected]. Great Basin Naturalist 59(2), ©1999, pp. 127-135 BIOGEOGRAPHY AND PHYSIOLOGICAL ADAPTATIONS OF THE BRINE FLY GENUS EPHYDRA (DIPTERA: EPHYDRIDAE) IN SALINE WATERS OF THE GREAT BASIN David B. Herbst1 ABSTRACf.-Four species of the genus Ephydra are commonly found in saline waters within the hydrologic Great Basin: E. hians, E. gracilis, E. packardi, and E. auripes. Though none of these brine flies is endemic (distributions also occur outside the Great Basin), they all inhabit distinctive habitat types and form the characteristic benthic insect fauna ofinland saline-water habitats. The affinities ofeach species for different salinity levels and chemical compositions, and ephemeral to perennial habitats, appear to form the basis for biogeographic distribution patterns. Within any habitat, changing salinity conditions over time may impose physiological or ecological constraints and further alter patterns of population productivity and the relative abundance of CO-inhabiting species. Based on the physiology of salt tolerance known for these species, high salinity conditions favor E. hians in alkaline water and E. gracilis in chloride water. At lower salinities, based on limited habitat data, E. auripes and E. packardi are often more common, again showing respective preferences for alkaline and chloride chemical conditions. Specialized adaptations for alkaline carbonate waters are found in the larval Malpighian tubule lime gland of the alkali fly E. hians, while high salt tolerance in E. gracilis appears to be conferred by high hemolymph osmolality. Adaptation to ephemeral and low salinity conditions may be accomplished by swift adult colonizing ability and rapid larval development rates. It is hypothesized that adaptive specializations in both physiology and life history and varied geochemistry of saline water habitats across the Great Basin produce the biogeographic pattern of distributions for species in this genus. This perspective on the genus Ephydra, and possibly other biota of mineral-rich Great Basin waters, suggests that intercon­ nections among pluvial lakes may be less relevant to aquatic biogeography than chemical profiles developing in remnant lakes and ponds with the progression ofarid post-pluvial climatic conditions. Key words: Ephydra, saline lakes, G1'eat Basin, osnwregulation, biogeography, salt tolerance. Ever since the studies of Hubbs and Miller within the Great Basin have been only incom­ (1948), biogeographic studies of the aquatic pletely described. Habitat affinities are poorly fauna of the Great Basin bave been dominated known, and collection records typically have by a search for vicariance patterns. Distribu­ no associated physical or chemical data. tions of fish species often revealed patterns In addition to geographic barriers, differen­ suggesting pluvial interconnections among tiation ofpopulations may also result from dif­ lake basins and proVided evidence for post­ ferences in tbe physical and chemical features pluvial hydrographic changes including isola­ of aquatic habitats without producing geo­ tion leading to extinctions and species differ­ graphic isolation and endemism. Selection for entiation (Miller 1946, Smith 1978). In addition physiological adaptation may restrict species to fish, other obligate aquatic species (e.g., distributions to certain habitat types. Great spring snails, leeches, molluscs) have received Basin aquatic habitats often include waters an inordinate amount of attention because of with varied chemistry derived from high min­ the potential for endemic distributions arising eral content. The closed-basin drainages that out of the vicariance events of pluvial lake define the Great Basin collect and evaporate drying and recession, and tbe example set by water in saline lakes, ponds, and wetlands. Huhbs and Miller (Taylor 1960, Hershler 1989, Many springs (geothermal and otherwise) con­ Hovingh 1995). Insects with poor dispersal tain high concentrations of dissolved solutes ability have also been studied in some depth and trace minerals. Streams in lower eleva­ in the Great Basin (e.g., Naucoridae; Polhemus tions ofthe basin and range also typically have and Polhemus 1994), but biogeograpbic pat­ relatively high conductivity and alkalinity. terns for most other aquatic invertebrates Physical conditions also may be quite varied ISierra Nevada Aquatic Re~earch Lahoratory, University ofCalifornia, Route 1, Box 198, Mammoth Lakes. CA 93546, and Marine Science IJI,tlhlte. Unl­ ver:sity ofCalifornia, Santa Barbara, CA 931013- 127 128 GREAT BASIN NATURALIST [Volume 59 in terms of thermal regime (hot springs) and that some Ephydra had particular chemical ephemeral and episodic filling of playas and preferences. The objective of this paper is to intermittent streams. Gradients of chemical present evidence for the hypothesis that affini­ and physical variation form a habitat template ties of each species for different salinity levels that sets the stage for adaptive syndromes and chemical compositions, and ephemeral to determining the distribution and abundance perennial habitats, form the basis for biogeo­ ofspecies (sensu Southwood 1977). graphic distribution patterns. Geochemical evolution of major solutes in closed-basin saline lakes (e.g., Eugster and METHODS Jones 1979) may be a useful model for follow­ ing progression and patterns in the biological Contrasts of physiological specializations evolution and distribution ofsaline water biota for habitat chemistry are presented here as throughout the Great Basin. Water chemistry is evidence of adaptations that determine bio­ determined primarily by the following factors: geographic patterns. Data for comparisons of • drainage basin lithology of inflowing osmoregulatory physiology and salt tolerance waters (especially igneous vs. sedimen­ in Ephydra were derived from published tary); research (Nemenz 1960, Herbst et aJ. 1988). • mineral precipitation pathways depend­ Calculations of relative costs of osmoregula­ ing on the initial ratio of bicarbonate to tion were determined by applying the known calcium and magnesium and resulting osmotic gradients to calculations of the rela­ in lake brines with differing contents of tive energy required for the work of active the major cations Na+/Ca+2/Mg+2 and transport. Data on the range of known devel­ opment times in Ephydra were also summa­ major anions Cl-/S04-2/C03-2 (high bi­ em-bonate generally evolves into alka­ rized from published sources (Ping 1921 for E. line brines and low bicarbonate into packardi, reported as E. subopaca; Collins chloride-sulfate brines, and low salinity 1980a for E. gracilis, reported as E. cinerea; and Herbst 1986 for E. hians). Distribution conditions may retain similar content of data were derived from Wirth (1971) and fur­ bicarbonate and Ca/Mg without forming ther supplemented by collection records and precipitates); habitat chemistry data of the author. • fractionation processes which enrich highly soluble (conservative) ions such as RESULTS Na and Cl and deplete others through carbonate mineral formation, release of Ecological and physiological limitations carbon dioxide gas, and biogenic reduc­ under changing salinity conditions in salt lakes tion ofsulfate for example. appear to result in varied distribution and pro­ Progressive enrichment from carbonate to ductivity of Ephydra spp. Distribution maps sulfatochloride to chloride waters appears to (Figs. 1, 2) and limited habitat data suggest occur with the concentration of inflowing high salinity conditions (usually >25 g-L-1) waters over time and distance along intercon­ favor E. hians in alkaline water and E. gracilis nected evaporation basins of varied volume in chloride water. In low salinity «25 g'L-l) (Hutchinson 1957). Spatial and temporal com­ and ephemeral waters, E. auripes and E. ponents of variation in water chemistty form a packardi are often more common, again show­ habitat template that may be the foundation ing respective preferences for alkaline and for shaping the distribution of saline water chloride chemical conditions. The general organisms. prevalence ofE. gracilis in the eastern, and E. Four species of the genus Ephydra are hians in the western, Great Basin correspond commonly found in saline waters within the to the trend in lake geochemistry being derived hydrologic Great Basin: E. hians, E. gracilis, from eastern sedimentary vs. western igneous E. packard~ and E. auripes. Though none of lithology in the generation of chemical profiles these brine flies is endemic (distributions also (Jones 1966,
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