Large Refugium Research Articles

Ecological consequences of intraspecific variation in lake Daphnia

Summary1. Although populations harbour considerable diversity, most ecological studies still assume they are homogeneous. However, mounting evidence suggests that intraspecific diversity is not only common, but also important for interactions with community members. Here, intraspecific variation in Daphnia dentifera in haemoglobin content is shown to be a marker of hypolimnion use.2. Hypolimnion use differed substantially within and among D. dentifera populations. Daphnia dentifera with haemoglobin resided primarily in the hypolimnion, while D. dentifera lacking haemoglobin migrated vertically. These ‘deep’ and ‘migratory’D. dentifera had different seasonal phenologies and dynamics.3. Deep and migratory D. dentifera had qualitatively different relationships with an important competitor, Daphnia pulicaria. Deep D. dentifera density was negatively correlated with D. pulicaria density, whereas migratory density was not correlated with D. pulicaria density.4. Given that D. pulicaria tends to reside in the hypolimnion, this negative correlation probably reflects competition between D. pulicaria and the deep D. dentifera. This pattern would have been missed if only the relationship between the overall lake populations of D. dentifera and D. pulicaria had been studied.5. Abundances of deep D. dentifera and D. pulicaria were both correlated with the size of the hypolimnetic refuge from fish predation, but in opposite directions. Lakes with large refuges generally had high D. pulicaria and low deep D. dentifera densities.

Free-living amoebae isolated from water-hyacinth root ( Eichhornia crassipes)

Free-living amoebae are widely distributed in aquatic environments and their hygienic, medical and ecological relationships to man are increasingly important. The purpose of this study was to isolate free-living amoebae from water-hyacinth root ( Eichhornia crassipes) and the water of an urban lake in Mexico City. Five grams of wet root were seeded on non-nutritive agar with Enterobacter aerogenes (NNE). Water samples were concentrated by centrifugation at 1200 g for 15 min and the pellet was seeded on NNE. Of the 16 isolated genera, 10 were detected in both habitats. The most frequent were Vannella in root and Acanthamoeba and Naegleria in water. The total number of isolates and genera isolated from root was higher than that isolated from water. The differences between root and water are probably due to the morphological characteristics of water-hyacinth root, which provides a large habitat and refuge area for many organisms.

Community-level enhancements of biodiversity and ecosystem services

A common management technique for preserving and maintaining biodiversity is the establishment of large refuges and preserves. Although extensive sanctuaries can provide crucial protection for many organisms and ecosystems, they cannot fulfill all the needs of regional conservation. An alternative to a few large refuges is to create many different habitats across the landscape that enhance and improve local and regional biodiversity and provide immediate benefits to nearby communities in the form of ecosystem services. Furthermore, these can all be initiated and achieved by individuals or communities. Some key landscape enhancements can be undertaken on a local level: the creation or expansion of small wooded areas, windbreaks, or hedgerows; the construction of small wetlands; and the release of some lands from heavy pressure for the reestablishment of natural ecological processes, namely, the natural accumulation of woody and other organic materials. Newly created ecosystems can be inoculated at the outset with soil biota such as seed banks, microbes, fungi, and organic material that can accelerate ecological functioning and balance. In addition to increasing much local and regional biodiversity, locally enhanced areas can provide fuel, plant and animal food and medicinal products, and agroforestry products directly to the nearby community. These small ecological oases can serve as nesting and overwintering sites for numerous pollinators that are hugely beneficial to agricultural production. Moreover, several ecosystem enhancements may contribute positively to local and regional hydrologic cycles and prevent prolonged droughts. Enhancements to local landscapes can take on many forms. We believe that any changes that increase structural complexity in natural systems almost certainly lead to increases in local biological complexity. In addition, wider landscape level considerations, such as corridors and connectivity of populations, can be integrated on a broader scale to improve regional biodiversity and ecosystem services. Small landscape enhancements undoubtedly cannot provide for all conservation needs, but they can greatly increase widespread biodiversity, restore local ecosystem services, and can be used to complement the relatively few larger parks.

Response of a southern temperate marsupial, the Tasmanian pademelon (Thylogale billardierii), to historical and contemporary forest fragmentation

Despite only limited Pleistocene glacial activity in the southern hemisphere, temperate forest species experienced complex distributional changes resulting from the combined effects of glaciation, sea level change and increased aridity. The effects of these historical processes on population genetic structure are now overlain by the effects of contemporary habitat modification. In this study, 10 microsatellites and 629 bp of the mitochondrial control region were used to assess the effects of historical forest fragmentation and recent anthropogenic habitat change on the broad-scale population genetic structuring of a southern temperate marsupial, the Tasmanian pademelon. A total of 200 individuals were sampled from seven sites across Tasmania and two islands in Bass Strait. High mitochondrial and nuclear genetic diversity indicated the maintenance of large historical population sizes. There was weak phylogeographical structuring of haplotypes, although all King Island haplotypes and three Tasmanian haplotypes formed a divergent clade implying the mid-Pleistocene isolation of a far northwestern population. Both the mitochondrial and nuclear data indicated a division of Tasmanian populations into eastern and western regions. This was consistent with a historical barrier resulting from increased aridity in the lowland 'midlands' region during glacial periods, and with a contemporary barrier resulting from recent habitat modification in that region. In Tasmania, gene flow appears to have been relatively unrestricted during glacial maxima in the west, while in the east there was evidence for historical expansion from at least one large glacial refuge and recolonization of Flinders Island.

On the fauna of the Diptera Brachycera of Samarskaya Luka: 1. Brachycera Orthorrhapha; Cyclorrhapha: Aschiza

The presumptive data on results of the investigation of the Diptera Brachycera in the territory of Samarskaya Luka, the largest refuge in eastern European Russia, are given. Based on the published data and on examination of 18 000 specimens, a faunal list for the region is compiled, including 897 species of 395 genera of 62 families. The annotated list of Brachycera Orthorrhapha, Cyclorrhapha: Aschiza includes data on the flight periods, habitats, and number of examined specimens.

Evidence for cryptic glacial refugia from North American mountain sheep mitochondrial DNA

The separation of populations by ice sheets into large refugia can account for much of the genetic diversity found in present day populations. The evolutionary implications of small glacial refugia have not been as thoroughly explored. To examine refugial origins of North American mountain sheep Ovis spp., we analyzed a 604 bp portion of the mitochondrial DNA (mtDNA) control region from 223 O. dalli and O. canadensis. Major refugia were identified in eastern Beringia and southern North America, and we found evidence for two smaller refugia situated between the Laurentide and Cordilleran glaciers. Our results are the first to demonstrate support for survival of any organism in the latter two refugia. These refugia also appear to have conserved a genetic signal that confirms past hybridization of O. dalli and O. canadensis.

Physiologically based demographics of Bt cotton–pest interactions: II. Temporal refuges, natural enemy interactions

An holistic demographic physiologically based age–mass structured model of transgenic Bt cotton expressing one or two toxin of the bacterium Bacillus thuringiensis Berliner (Bt) and 10 of its pests as affected by generalist natural enemies is developed. The goal of the model is to assess factors favoring the development of resistance in pest species. Among the pest included in the model are the pink bollworm (a primary pest of cotton), and several secondary pests of the family Noctuidae (tobacco budworm, bollworm, fall armyworm, beet armyworm, cabbage looper, soybean looper) as well as a plant bug, boll weevil and whitefly. The pests have widely varying tolerance to the toxin. Resistance to the Bt toxin is assumed recessive, autosomal and controlled by a single diallelic gene. S patial refuges of non-Bt cotton have been mandated to increase heterozygosity in Bt cotton, but spatial refuges may exist for many pest species in non-Bt and/or sylvan hosts. In addition, temporal refuges may arise within Bt cotton due to innate pest tolerance to the toxin that increases with larval age, and variable toxin concentrations in the plant over time and with plant subunit age. Mortality rates and non-lethal effects that increase larval developmental time and decrease adult fecundity vary with Bt toxin concentration. All genotypes are affected but resistant genotypes are assumed less affected than susceptible ones. The effects of spatial and temporal refuges on the development of resistance and pest densities in one and two toxin cottons are examined over several consecutive seasons. Resistance to Bt toxin is more likely to develop in stenophagous highly susceptible pests such as pink bollworm and budworm than in polyphagous highly tolerant pests such as fall armyworm, beet armyworm and soybean looper that have large temporal refuges that slow and possibly reverse resistance development. Predator longevity feeding on Bt intoxicated prey is decreased ca. 28% [Ponsard, S., Gutierrez, A.P., Mills, N.J., 2002. Effects of Bt-toxin in transgenic cotton on the adult longevity of four heteropteran predators. Environ. Entomol. 31, 1197–1205.] creating tradeoffs between the mortality caused by the toxin and that due to reduced predation. Despite reduced natural enemy efficacy, levels of susceptible pests (budworm) are reduced, while levels of Bt tolerant and immune pests (e.g. fall armyworm, Lygus) increase. The use of pesticide for supplementary control of tolerant pests in Bt cotton may further disrupts natural enemies and increase pest levels.

Refuge effects of a cactus in grazed short‐grass steppe

Abstract. Question: Which factors influence the effectiveness of biotic refuges for harbouring grazing‐sensitive species in pastures with a long history of grazing by large herbivores? Previous research showed that spiny clumps of the cactus Opuntia polyacantha provided refuges from cattle grazing for plants and for inflorescence production on short‐grass steppe. In this paper, seven factors that may have a potential positive influence on the refuge effects of cactus at a landscape scale were assessed.Location: Short‐grass steppe of the Great Plains of North America.Methods: The study was conducted in eight long‐term grazed pastures and their respective ungrazed controls that were established 60 years ago.Results: Heavy grazing intensities were necessary for some positive effects of cactus to manifest, and some refuge effects changed to negative effects under lower grazing pressure. Refuge effects increased with plant community productivity due to greater abundances of grazing‐sensitive species, and greater grazing intensities in the more productive areas. Cover of cactus cladodes (spine‐covered pads) inside clumps appeared to be the main limiting factor for refuge effects, probably by limiting available space for grazing‐sensitive species in the clumps. Other factors such as size and density of cactus clumps, and the presence of large refuges in the proximity of clumps had minor influence on the effectiveness of cactus refuges.Conclusions: The effects of biotic refuges largely varied with ecological conditions and structural characteristics of the refuge. Refuge effects were mainly influenced by grazing intensity, plant community productivity, and structural characteristics of the biotic refuges. A conceptual model of factors influencing refuge effects at a local landscape scale in plant communities grazed by large herbivores is presented.

Surviving the ice: Northern refugia and postglacial colonization.

The contemporary distribution of biological diversity cannot be understood without knowledge of how organisms responded to the geological and climatic history of Earth. In particular, Quaternary expansions and contractions of glacial ice sheets are thought to have played an important role in shaping the distribution of biodiversity among current populations in the north-temperate region. In the central U.S., fossil and palynological data provide support for the maintenance of a large southeastern refuge during the last glacial maximum, and many temperate organisms are believed to have responded to glacial expansion by shifting their ranges to southern refugia and recolonizing northward to track the receding ice sheets. Thus, organisms are assumed to track favorable climates, and species ranges are expected to have shifted significantly. Here we present data from a deciduous forest vertebrate, the eastern chipmunk (Tamias striatus) in the central U.S., indicating the maintenance of multiple refugial sources as well as a southward expansion from a northern refugium. These results challenge the view that, during glacial maxima, organisms must have migrated south out of their ranges to track favorable climates.

Grazing refuges, external avoidance of herbivory and plant diversity

Avoidance and tolerance are the two means by which plants cope with herbivores. Avoidances internal to the plant, such as morphology, chemical repellants, thorns, etc., have received considerable attention in the plant‐herbivore literature, but relatively little consideration has been given to avoidances external to the plant. We develop a conceptual framework of external plant avoidances of herbivory based on foraging selection impedances (associational avoidances), behavioral impedances (indirect avoidances), and physical impedances (refuges) organized along axes of efficiency, degree of protection, and necessity of tolerance characteristics. Associational avoidances are uncommon for terrestrial mammalian herbivores compared to plant‐insect or marine situations. Indirect avoidances mediated through herbivore territoriality, predator avoidance, and other behaviors independent of foraging decisions are probably common in nature, but few have been formally documented. Biotic and geologic refuges providing a physical impedance are the only avoidances shown to have implications for plant biodiversity. This is particularly true for geologic refuges, where there is not a tradeoff between competition and the refuge effect. Small geologic refuges (rock outcrops, cliffs, etc.) are more likely to also positively or negatively alter associated plant microenvironments than large geologic refuges (mesas, islands, etc.). In a survey, 86% of small refuge studies reported positive effects on plant diversity compared to 50% for large refuges. Geologic refuges in more productive environments were more important in protecting diversity than refuges in less productive, semiarid environments, and the effects of protection were greater in communities with short compared to long evolutionary histories of grazing. Other characteristics of refuges such as extent across the landscape and the manner they alter or ameliorate the environment, as well as characteristics of the herbivore such as small or large, generalist or specialist may also determine the effectiveness of refuges, but there are too few studies to assess these factors.

Intraspecific allozymatic differentiation reveals the glacial refugia and the postglacial expansions of EuropeanErebia medusa (Lepidoptera: Nymphalidae)

Allozyme analysis of Erebia medusa over large regions of Europe revealed a significant population differentiation (FST: 0.149±0.016). A UPGMA-analysis showed a division into four major lineages with mean inter-group genetic distances ranging from 0.051 (±0.010) to 0.117 (±0.024). An AMOVA revealed that rather more than two-thirds of the variance between samples was being between these lineages and less than one-third within lineages. An eastern group included the samples from the Czech Republic, Slovakia and north-eastern Hungary. This genetic lineage expressed significantly higher genetic diversity than the other three. A second lineage was formed by the samples from France and Germany. The two samples from western Hungary represent a third delimited lineage and the sample from northern Italy a fourth. We suppose that this genetic differentiation took place during the last ice-age in four disjunct refugia. The genetically more diverse eastern genetic lineage might have evolved in a relatively large refugium in south-eastern Europe. We assume that the other three lineages developed in relatively small relict areas around the Alps. It is likely for the western lineage that its ice-age distribution showed at least one disjunction in late Würm with the consequence of further genetic differentiation. Most probably, the eastern lineage colonized postglacial Central Europe using two alternative routes: one north and one south of the Carpathians. Up to now, neither similar glacial refugia, nor comparable secondary disjunctions in late Würm, are reported for any other animal or plant species.

Intraspecific allozymatic differentiation reveals the glacial refugia and the postglacial expansions of European Erebia medusa (Lepidoptera: Nymphalidae)

Allozyme analysis of Erebia medusa over large regions of Europe revealed a significant population differentiation (FST: 0.149±0.016). A UPGMA-analysis showed a division into four major lineages with mean inter-group genetic distances ranging from 0.051 (±0.010) to 0.117 (±0.024). An AMOVA revealed that rather more than two-thirds of the variance between samples was being between these lineages and less than one-third within lineages. An eastern group included the samples from the Czech Republic, Slovakia and north-eastern Hungary. This genetic lineage expressed significantly higher genetic diversity than the other three. A second lineage was formed by the samples from France and Germany. The two samples from western Hungary represent a third delimited lineage and the sample from northern Italy a fourth. We suppose that this genetic differentiation took place during the last ice-age in four disjunct refugia. The genetically more diverse eastern genetic lineage might have evolved in a relatively large refugium in south-eastern Europe. We assume that the other three lineages developed in relatively small relict areas around the Alps. It is likely for the western lineage that its ice-age distribution showed at least one disjunction in late Wurm with the consequence of further genetic differentiation. Most probably, the eastern lineage colonized postglacial Central Europe using two alternative routes: one north and one south of the Carpathians. Up to now, neither similar glacial refugia, nor comparable secondary disjunctions in late Wurm, are reported for any other animal or plant species.

107 Days on the SH-7: Experiences and Events of the Last Large Refuge Transport from the <I>Reichsgbiet</I>

107 Days on the SH-7: Experiences and Events of the Last Large Refuge Transport from the <I>Reichsgbiet</I>

Reversing insect adaptation to transgenic insecticidal plants.

The refuge-high-dose strategy for delaying insect adaptation to transgenic plants produces non-transgenic plants that enable survival of susceptible individuals. Previous theoretical work has suggested three requirements for success of the refuge-high-dose strategy: a low initial frequency of the resistance allele, extensive mating between resistant and susceptible adults and recessive inheritance of resistance. In order to understand an observed decrease in resistance frequency and improve the potential for managing resistance better, we used analytical and simulation models for exploring the conditions that prevent or reverse the evolution of resistance, even when resistance is not rare initially. Assuming random mating and recessive or nearly recessive inheritance of resistance, the factors favouring reversal of resistance are non-recessive costs of resistance, low initial resistance allele frequency, large refuges, incomplete resistance and density-independent population growth in refuges.

The Quaternary vegetation and landscape evolution of Novaya Zemlya in the light of palynological records

The paper presents the results of paleoenvironmental (pollen) investigations of Novaya Zemlya throughout all the Quaternary, including Eopleistocene, Pleistocene and Holocene. Clear evidence of continuity between the modern flora and paleofloras confirms the lack of sharp and continuous interruptions in the vegetation development in high latitudes as well as the absence of gigantic ice sheets in the Eastern Arctic. During the Eopleistocene plant communities varied from Boreal meadow coenoses in the south of Novaya Zemlya to Arctic tundra in the north. During the Lower Pleistocene the climate had deteriorated considerably, but still remained warmer than today's. There were two cold (=glacial) and two warm (=interglacial) stages in the Upper Pleistocene. The older interglacial, Kumzha, was marked by rather mild climate and extensive marine transgression. During the climatic optimum southern dwarf shrub tundra grew in the Southern Island, but forest did not penetrate there. The younger interglacial, Krestyakhino, had cooler climate and more restricted transgression extent. Typical tundra existed in the Southern Island. The Upper Pleistocene glaciations were no more extensive than nowadays and large biota refuges existed in the coastal and even some inner areas of Novaya Zemlya at least between 15,000 and 10,000 BP. During the Holocene, great change took place at the end of climatic optimum when dwarf shrub tundra was replaced by more open tundra communities.

Progress Toward Lake Trout Restoration in Lake Michigan

Progress toward lake trout restoration in Lake Michigan is described through 1993. Extinction of the native lake trout fishery by sea lamprey predation, augmented by exploitation and habitat destruction, resulted in an extensive stocking program of hatchery-reared lake trout that began in 1965. Sea lamprey abundance was effectively controlled using selective chemical toxicants. The initial stocking produced a measurable wild year class of lake trout by 1976 in Grand Traverse Bay, but failed to continue probably due to excessive exploitation. The overall lack of successful reproduction lakewide by the late 1970s led to the development and implementation in 1985 of a focused inter-agency lakewide restoration plan by a technical committee created through the Lake Committee structure of the Great Lakes Fishery Commission. Strategies implemented in 1985 by the plan included setting a 40% total mortality goal lakewide, creating two large refuges designed to encompass historically the most productive spawning habitat and protect trout stocked over their home range, evaluating several lake trout strains, and setting stocking priorities throughout the lake. Target levels for stocking in the 1985 Plan have never been reached, and are much less than the estimated lakewide recruitment of yearlings by the native lake trout stocks. Since 1985, over 90% of the available lake trout have been stocked over the best spawning habitat, and colonization of the historically productive offshore reefs has occurred. Concentrations of spawning lake trout large enough for successful reproduction, based on observations of successful hatchery and wild stocks, have developed at specific reefs. Continued lack of recruitment at these specific sites suggests that something other than stotk abundance has limited success. Poor survival of lake trout eggs, assumed to be related to contaminant burden, occurred in the late 1970s and early 1980s, but survival has since increased to equal survival in the hatchery. A recent increase in lamprey wounding rates in northern Lake Michigan appears to be related to the uncontrolled build-up of lampreys in the St. Marys River a tributary of Lake Huron. If left uncontrolled, further progress toward restoration in the Northern Refuge may be limited.

Cladoceran assemblages, seasonal succession and the importance of a hypolimnetic refuge

SUMMARY. 1. We investigate the importance of a refuge from fish predation to the abundance, species composition and seasonal succession of zooplankton. Thirty lakes representing a range of depths were sampled twice in summer for physical/chemical parameters and zooplankton community structure.2. We define the refuge from centrarchid predators to be that space between the thermocline and the zone of anoxia. As lakes vary in rate of oxygen depletion from the hypolimnion. the refuge size and lake depth are independent: refuge size decreases during the summer period.3. Lake depth and refuge size independently explain variation among lakes in zooplankton species composition, but seasonal community change within lakes is best predicted by loss of refuge size.4. Refuge size also explains the substantial variation in the relative dominance of the two major daphnid species. Lakes possessing a large refuge are dominated by D. pulicaria; those with a small refuge are dominated by the smaller, D. galeata mendotae. We suggest that lakes of intermediate refuge size, which are characterized by high species diversity, represent a more equitable balance of predation and competition.

Land and labour in Latin American agriculture from the 1950s to the 1980s

A considerable amount of statistical information on the evolution of the Latin American peasantry during the last 35 years is used to show that it has become a large refuge sector that symptomises the developmental failures of the rest of the economy. While economic growth has been rapid until the debt crisis of the early 1980s, neither access to land nor employment creation has been sufficient to reduce rural poverty. As a result, the bulk of the peasantry neither competes with commercial farming nor becomes dispossessed of access to a plot of land. Through migration, poverty gradually becomes displaced to the urban sector.

On species richness and nature reserve design: An empirical study of UK woodland avifauna

Abstract The numbers and turnover of breeding bird species are compared for sets of 5, 3,2 and 1 similar UK woodland sites, each set having the same total area. Five small (8 ha) sites support the largest total number of species, though it is argued on theoretical grounds that this result may be scale-dependent. Species turnover is lowest for the set of five small sites and hence their overall species composition will be more predictable in time. An attempt is made to predict the expected number of species remaining in sites that act as refuges in the absence of an external species pool; the data indicate that one large refuge may come to retain more species than several smaller refuges as extinctions occur, provided that the refuges are effectively isolated from one another. The implications for avifaunal conservation are discussed, with emphasis on the need for an empirical approach to evaluating nature reserve design options.

Effects of insularisation on plant species richness in the prairie-forest ecotone

Data on plants from five groups of remnant prairies and forests in the prairie-forest ecotone of the midwestern United States show that: (1) Archipelagos of small sites tend to contain more species than do single large ones of equal total area. (2) No species are excluded from small sites. (3) Small sites tend to have surprisingly many species, and large sites surprisingly few, relative to a random colonisation model. (4) ‘Rare’ species (those that occur in only one site) are found more often in small sites than a random colonisation model would predict. (5) There is no evidence for any of these sites that species number is ‘relaxing’, but all these sites have been remnants for only a short time. Result (1) is consistent with results from similar studies on other taxa and suggests that there is no automatic reason to believe that species richness is maximised by single large refuges.