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Refugia
Kinds of Refugia Terms modified by Refugia Selected AbstractsTHE COMBINED EFFECTS OF RIVERS AND REFUGIA GENERATE EXTREME CRYPTIC FRAGMENTATION WITHIN THE COMMON GROUND SKINK (SCINCELLA LATERALIS)EVOLUTION, Issue 2 2010Nathan D. Jackson Rivers can act as both islands of mesic refugia for terrestrial organisms during times of aridification and barriers to gene flow, though evidence for long-term isolation by rivers is mixed. Understanding the extent to which riverine barrier effects can be heightened for populations trapped in mesic refugia can help explain maintenance and generation of diversity in the face of Pleistocene climate change. Herein, we implement phylogenetic and population genetic approaches to investigate the phylogeographic structure and history of the ground skink, Scincella lateralis, using mtDNA and eight nuclear loci. We then test several predictions of a river,refugia model of diversification. We recover 14 well-resolved mtDNA lineages distributed east,west along the Gulf Coast with a subset of lineages extending northward. In contrast, ncDNA exhibits limited phylogenetic structure or congruence among loci. However, multilocus population structure is broadly congruent with mtDNA patterns and suggests that deep coalescence rather than differential gene flow is responsible for mtDNA,ncDNA discordance. The observed patterns suggest that most lineages originated from population vicariance due to riverine barriers strengthened during the Plio,Pleistocene by a climate-induced coastal distribution. Diversification due to rivers is likely a special case, contingent upon other environmental or biological factors that reinforce riverine barrier effects. [source] HABITAT FRAGMENTATION AND BIODIVERSITY: TESTING FOR THE EVOLUTIONARY EFFECTS OF REFUGIAEVOLUTION, Issue 6 2004Jon R. Bridle Abstract Concordant areas of endemism among taxa have important implications both for understanding mechanisms of speciation and for framing conservation priorities. Here we discuss the need for careful testing of phylogeographic data for evidence of such concordance, with particular reference to the Indonesian island of Sulawesi. This is because there are good reasons to question whether concordance between taxa is likely to be a common pattern, and because of the serious implications of incorrectly concluding that the biodiversity of a given area can be partitioned in this way. [source] TIME TO THE MOST RECENT COMMON ANCESTOR AND DIVERGENCE TIMES OF POPULATIONS OF COMMON CHAFFINCHES (FRINGILLA COELEBS) IN EUROPE AND NORTH AFRICA: INSIGHTS INTO PLEISTOCENE REFUGIA AND CURRENT LEVELS OF MIGRATIONEVOLUTION, Issue 1 2002Cortland K. Griswold Abstract We analyzed sequences from a 275-bp hypervariable region in the 5, end of the mitochondrial DNA control region in 190 common chaffinches (Fringilla coelebs) from 19 populations in Europe and North Africa, including new samples from Greece and Morocco. Coalescent techniques were applied to estimate the time to the most recent common ancestor (TMRCA) and divergence times of these populations. The first objective of this study was to infer the locations of refugia where chaffinches survived the last glacial episode, and this was achieved by estimating the TMRCA of populations in regions surrounding the Mediterranean that were unglaciated in the late Pleistocene. Although extant populations in Iberia, Corsica, Greece, and North Africa harbor haplotypes that are basal in a phylogenetic tree, this information alone cannot be used to infer that these localities served as refugia, because it is impossible to infer the ages of populations and their divergence times without also considering the population genetic processes of mutation, migration, and drift. Provided we assume the TMRCAs of populations are a reasonable estimate of a population's age, coalescent-based methods place resident populations in Iberia, Corsica, Greece, and North Africa during the time of the last glacial maximum, suggesting these regions served as refugia for the common chaffinch. The second objective was to determine when populations began diverging from each other and to use this as a baseline to estimate current levels of gene flow. Divergence time estimates suggest that European populations began diverging about 60,000 years before present. The relatively recent divergence of populations in North Africa, Italy, and Iberia may explain why classic migration estimates based on equilibrium assumptions are high for these populations. We compare these estimates with nonequilibrium-based estimates and show that the nonequilibrium estimates are consistently lower than the equilibrium estimates. [source] Mid-Holocene and glacial-maximum vegetation geography of the northern continents and AfricaJOURNAL OF BIOGEOGRAPHY, Issue 3 2000I. Colin Prentice Abstract BIOME 6000 is an international project to map vegetation globally at mid-Holocene (6000 14C yr bp) and last glacial maximum (LGM, 18,000 14C yr bp), with a view to evaluating coupled climate-biosphere model results. Primary palaeoecological data are assigned to biomes using an explicit algorithm based on plant functional types. This paper introduces the second Special Feature on BIOME 6000. Site-based global biome maps are shown with data from North America, Eurasia (except South and Southeast Asia) and Africa at both time periods. A map based on surface samples shows the method's skill in reconstructing present-day biomes. Cold and dry conditions at LGM favoured extensive tundra and steppe. These biomes intergraded in northern Eurasia. Northern hemisphere forest biomes were displaced southward. Boreal evergreen forests (taiga) and temperate deciduous forests were fragmented, while European and East Asian steppes were greatly extended. Tropical moist forests (i.e. tropical rain forest and tropical seasonal forest) in Africa were reduced. In south-western North America, desert and steppe were replaced by open conifer woodland, opposite to the general arid trend but consistent with modelled southward displacement of the jet stream. The Arctic forest limit was shifted slighly north at 6000 14C yr bp in some sectors, but not in all. Northern temperate forest zones were generally shifted greater distances north. Warmer winters as well as summers in several regions are required to explain these shifts. Temperate deciduous forests in Europe were greatly extended, into the Mediterranean region as well as to the north. Steppe encroached on forest biomes in interior North America, but not in central Asia. Enhanced monsoons extended forest biomes in China inland and Sahelian vegetation into the Sahara while the African tropical rain forest was also reduced, consistent with a modelled northward shift of the ITCZ and a more seasonal climate in the equatorial zone. Palaeobiome maps show the outcome of separate, independent migrations of plant taxa in response to climate change. The average composition of biomes at LGM was often markedly different from today. Refugia for the temperate deciduous and tropical rain forest biomes may have existed offshore at LGM, but their characteristic taxa also persisted as components of other biomes. Examples include temperate deciduous trees that survived in cool mixed forest in eastern Europe, and tropical evergreen trees that survived in tropical seasonal forest in Africa. The sequence of biome shifts during a glacial-interglacial cycle may help account for some disjunct distributions of plant taxa. For example, the now-arid Saharan mountains may have linked Mediterranean and African tropical montane floras during enhanced monsoon regimes. Major changes in physical land-surface conditions, shown by the palaeobiome data, have implications for the global climate. The data can be used directly to evaluate the output of coupled atmosphere-biosphere models. The data could also be objectively generalized to yield realistic gridded land-surface maps, for use in sensitivity experiments with atmospheric models. Recent analyses of vegetation-climate feedbacks have focused on the hypothesized positive feedback effects of climate-induced vegetation changes in the Sahara/Sahel region and the Arctic during the mid-Holocene. However, a far wider spectrum of interactions potentially exists and could be investigated, using these data, both for 6000 14C yr bp and for the LGM. [source] The role of refugia and dispersal in primary succession on Mount St. Helens, WashingtonJOURNAL OF VEGETATION SCIENCE, Issue 5 2003R.N. Fuller Taxonomy follows the Integrated Taxonomic Information System (ITIS) (http://www.itis.usda.gov) Abstract. An intense lateral blast devastated Mount St. Helens in 1980, but forest understory species survived in some north-slope ,refugia'. We explored the effects of refugia on colonization of barren pumice in 1997 and 1998, 18 yr after the eruption. The seed rain of 23 colonizers came mostly from populations that had previously established in refugia. Parachutists had small, vagile seeds, parasailors had winged seeds, and tumblers were blown along the ground. The latter two groups are heavier and dispersed more slowly, but are more likely to survive. The proportion of the vegetation represented by wind-dispersed species increased with distance from refugia. Parachutist's density declined with time and proximity to refugia. As vegetation adjacent to refugia developed, populations of parasailors and tumblers expanded, foreshadowing their dominance in more remote pumice. Refugia played a critical role in determining the rate and course of succession by providing fertile islands that permitted pioneers and dry meadow species to establish near barren pumice. Species that survived in refugia played a negligible role in colonization. This study showed that when refugia contrast sharply with new substrates, they accelerate recovery by facilitating the invasion of pioneer species. [source] Refugia, differentiation and postglacial migration in arctic-alpine Eurasia, exemplified by the mountain avens (Dryas octopetala L.)MOLECULAR ECOLOGY, Issue 7 2006INGER SKREDE Abstract Many arctic-alpine organisms have vast present-day ranges across Eurasia, but their history of refugial isolation, differentiation and postglacial expansion is poorly understood. The mountain avens, Dryas octopetala sensu lato, is a long-lived, wind-dispersed, diploid shrub forming one of the most important components of Eurasian tundras and heaths in terms of biomass. We address differentiation and migration history of the species with emphasis on the western and northern Eurasian parts of its distribution area, also including some East Greenlandic and North American populations (partly referred to as the closely related D. integrifolia M. Vahl). We analysed 459 plants from 52 populations for 155 amplified fragment length polymorphisms (AFLP) markers. The Eurasian plants were separated into two main groups, probably reflecting isolation and expansion from two major glacial refugia, situated south and east of the North European ice sheets, respectively. Virtually all of northwestern Europe as well as East Greenland have been colonized by the Southern lineage, whereas northwest Russia, the Tatra Mountains and the arctic archipelago of Svalbard have been colonized by the Eastern lineage. The data indicate a contact zone between the two lineages in northern Scandinavia and possibly in the Tatra Mountains. The two single populations analysed from the Caucasus and Altai Mountains were most closely related to the Eastern lineage but were strongly divergent from the remaining eastern populations, suggesting survival in separate refugia at least during the last glaciation. The North American populations grouped with those from East Greenland, irrespective of their taxonomic affiliation, but this may be caused by independent hybridization with D. integrifolia and therefore not reflect the true relationship between populations from these areas. [source] Speciation, hybrid zones and phylogeography , or seeing genes in space and timeMOLECULAR ECOLOGY, Issue 3 2001Godfrey M. Hewitt Abstract The origins and development of the study of speciation, hybrid zones and phylogeography are outlined using evolutionary iconography. This traces the ideas in this field from Lamarck and Darwin through to the present as represented in diagrams and figures. A ,tree of trees' summarizes this growth and current vitality. The new facility to use various DNA sequences from nuclear, mitochondrial and chloroplast genomes to determine genetic variation throughout a species range is examined particularly. There is great genomic subdivision across species distributions, which can be interpreted in the light of the recent demonstrations of severe palaeoclimatic oscillations. Refugia and postglacial colonization routes are proposed for several organisms across Europe. The role of geography in speciation through the Pleistocene is considered. These emerging principles and analyses are applied to data available on a variety of organisms in other regions of the world, such as the Arctic, North America and the Tropics, and including the progress of Homo sapiens through the last ice age. Some suggestions are made for future research directions. [source] Past and present potential distribution of the Iberian Abies species: a phytogeographic approach using fossil pollen data and species distribution modelsDIVERSITY AND DISTRIBUTIONS, Issue 2 2010Francisca Alba-Sánchez Abstract Aim, Quaternary palaeopalynological records collected throughout the Iberian Peninsula and species distribution models (SDMs) were integrated to gain a better understanding of the historical biogeography of the Iberian Abies species (i.e. Abies pinsapo and Abies alba). We hypothesize that SDMs and Abies palaeorecords are closely correlated, assuming a certain stasis in climatic and topographic ecological niche dimensions. In addition, the modelling results were used to assign the fossil records to A. alba or A. pinsapo, to identify environmental variables affecting their distribution, and to evaluate the ecological segregation between the two taxa. Location, The Iberian Peninsula. Methods, For the estimation of past Abies distributions, a hindcasting process was used. Abies pinsapo and A. alba were modelled individually, first calibrating the model for their current distributions in relation to the present climate, and then projecting it into the past,the last glacial maximum (LGM) and the Middle Holocene periods,in relation to palaeoclimate simulations. The resulting models were compared with Iberian-wide fossil pollen records to detect areas of overlap. Results, The overlap observed between past Abies refugia,inferred from fossil pollen records,and the SDMs helped to construct the Quaternary distribution of the Iberian Abies species. SDMs yielded two well-differentiated potential distributions: A. pinsapo throughout the Baetic mountain Range and A. alba along the Pyrenees and Cantabrian Range. These results propose that the two taxa remained isolated throughout the Quaternary, indicating a significant geographical and ecological segregation. In addition, no significant differences were detected comparing the three projections (present-day, Mid-Holocene and LGM), suggesting a relative climate stasis in the refuge areas during the Quaternary. Main conclusions, Our results confirm that SDM projections can provide a useful complement to palaeoecological studies, offering a less subjective and spatially explicit hypothesis concerning past geographic patterns of Iberian Abies species. The integration of ecological-niche characteristics from known occurrences of Abies species in conjunction with palaeoecological studies could constitute a suitable tool to define appropriate areas in which to focus proactive conservation strategies. [source] Regional assessment of the impact of climatic change on the distribution of a tropical conifer in the lowlands of South AmericaDIVERSITY AND DISTRIBUTIONS, Issue 6 2007Marie-Pierre Ledru Abstract For decades, palynologists working in tropical South America are using the genus Podocarpus as a climate indicator although without referring to any modern data concerning its distribution and limiting factors. With the aim to characterize the modern and past distribution of the southern conifer Podocarpus in Brazil and to obtain new information on the distribution of the Atlantic rainforest during the Quaternary, we examined herbarium data to locate the populations of three Brazilian endemic Podocarpus species: P. sellowii, P. lambertii, and P. brasiliensis, and extracted DNA from fresh leaves from 26 populations. Our conclusions are drawn in the light of the combination of these three disciplines: botany, palynology, and genetics. We find that the modern distribution of endemic Podocarpus populations shows that they are widely dispersed in eastern Brazil, from north to south and reveals that the expansion of Podocarpus recorded in single Amazonian pollen records may have come from either western or eastern populations. Genetic analysis enabled us to delimit regional expansion: between 5° and 15° S grouping northern and central populations of P. sellowii expanded c. 16,000 years ago; between 15° and 23° S populations of either P. lambertii or sellowii expanded at different times since at least the last glaciation; and between 23° and 30° S, P. lambertii appeared during the recent expansion of the Araucaria forest. The combination of botany, pollen, and molecular analysis proved to be a rapid tool for inferring distribution borders for sparse populations and their regional evolution within tropical ecosystems. Today the refugia of rainforest communities we identified are crucial hotspots to allow the Atlantic forest to survive under unfavourable climatic conditions and, as such, offer the only possible opportunity for this type of forest to expand in the event of a future climate change. [source] Patterns of isozyme variation as indicators of biogeographic history in Pilgerodendron uviferum (D. Don) FlorínDIVERSITY AND DISTRIBUTIONS, Issue 2 2002A. C. Premoli Abstract. The effects of Pleistocene glaciations on the genetic characteristics of the most austral conifer in the world, Pilgerodendron uviferum, were analysed with specific reference to the hypothesis that the species persisted locally in ice-free areas in temperate South America. It was expected that genetic variation would decrease with latitude, given that ice fields were larger in southern Patagonia and thus refugia were probably located towards the northern distributional limit of the species as suggested by the fossil record. In addition, an increase in among-population genetic divergence was expected with increasing distance to putative glacial refugia. We examined the relationship between location and within-population variability indices of 20 Pilgerodendron populations derived from isozyme analyses. We analysed possible refugia hypotheses by the distribution of allele frequencies using multivariate discriminant analysis. The degree of genetic differentiation with geographical distance between all population pairs was investigated by Mantel tests. Results indicated that Pilgerodendron populations are highly monomorphic, probably reflecting past population bottlenecks and reduced gene flow. Southernmost populations tend to be the least genetically variable and were therefore probably more affected by glacial activity than northern ones. Populations located outside ice limits seem to have been isolated during the glacial period. The presence of centres of genetic diversity, together with the lack of a significant correlation between genetic and geographical distances and the absence of geographical patterns of allelic frequencies at most analysed alleles, may indicate that Pilgerodendron did not advance southward after the last glaciation from a unique northern refugium, but spread from several surviving populations in ice-free areas in Patagonia instead. [source] Mollusk species diversity in the Southeastern Pacific: why are there more species towards the pole?ECOGRAPHY, Issue 2 2003Claudio Valdovinos The most ubiquitous and well recognized diversity pattern at large spatial scales is the latitudinal increase in species richness near the equator and decline towards the poles. Although several exceptions to this pattern have been documented, shallow water mollusks, the most specious group of marine invertebrates, are the epitome of the monotonic decline in species diversity toward higher latitudes along the Pacific and Atlantic coasts of North America. Here we analyze the geographic diversity of 629 mollusk species along the Pacific South American shelf. Our analyses are based on the most complete database of invertebrates assembled for this region of the world, consisting of latitudinal ranges of over 95% of all described mollusks between 10° and 55°S. Along this coast, mollusk diversity did not follow the typical latitudinal trend. The number of species remained constant and relatively low at intermediate latitudes and sharply increased toward higher latitudes, south of 42°S. This trend was explained by changes in shelf area, but not by sea surface temperature, unlike the pattern documented for Northern Hemisphere mollusks. Direct sampling of soft bottom communities along the gradient suggests that regional trends in species richness are produced by increased alpha diversity, and not only by artifacts produced by the increase in sampling area. We hypothesize that increased shelf area south of 42°S, geographic isolation produced by divergence of major oceanic currents, and the existence of refugia during glaciations, enabled species diversification. Radiation could have been limited by narrow continental shelves between 10°,42°. Asymmetries in latitudinal diversity trends between hemispheres show that there is not a single general factor determining large-scale diversity patterns. [source] Relationship between stream temperature, thermal refugia and rainbow trout Oncorhynchus mykiss abundance in arid-land streams in the northwestern United StatesECOLOGY OF FRESHWATER FISH, Issue 1 2001J. L. Ebersole Abstract , Warm stream temperatures may effectively limit the distribution and abundance of Pacific salmon Oncorhynchus spp. in streams. The role of cold thermal refugia created by upwelling groundwater in mediating this effect has been hypothesized but not quantitatively described. Between June 21 and September 15, 1994, rainbow trout O. mykiss abundance within 12 northeast Oregon (USA) stream reaches was inversely correlated with mean ambient maximum stream temperatures (r=,0.7, P<0.05). Some rainbow trout used thermal refugia (1,10 m2 surface area) that were on average 3,8°C colder than ambient stream temperatures. Within the warmest reaches, high ambient stream temperatures (>22°C) persisted from mid-June through August, and on average 10,40% of rainbow trout were observed within thermal refugia during periods of midday maximum stream temperatures. Frequency of cold-water patches within reaches was not significantly associated with rainbow trout density after accounting for the influence of ambient stream temperature (P=0.06; extra sum of squares F -test). Given prolonged high ambient stream temperatures in some reaches, the thermal refugia available in the streams we examined may be too small and too infrequent to sustain high densities of rainbow trout. However, these refugia could allow some rainbow trout to persist, although at low densities, in warm stream reaches. [source] Effects of fires on butterfly assemblages in lowland dipterocarp forest in East KalimantanENTOMOLOGICAL SCIENCE, Issue 2 2007Toshiya HIROWATARI Abstract The post-fire butterfly fauna in lowland dipterocarp forest of the Bukit Soeharto Education Forest (BSEF), East Kalimantan, Indonesia, was assessed during the period November 1998,April 2000 by means of consecutive Malaise trap samples, with supplementary field observations for March,April 1999. A total of 514 butterflies belonging to 61 species and representing six families were caught in the traps. Melanitis leda (Nymphalidae: Satyrinae), Charaxes bernardus (Nymphalidae: Charaxinae), and Danaus genutia (Nymphalidae: Danainae) were the species most frequently caught (60, 52 and 47 individuals, respectively), representing 31% of the total. These three species are generalists and "disturbance indicators" for tropical rainforest, being characteristic of disturbed or secondary forests, being distributed widely, and having larvae that feed on a wide range of host plants. In contrast, other species, such as Trogonoptera brookiana and Troides amphrysus, were recorded before the fires but were not recorded again afterwards. The pre- and post-fire butterfly fauna of East Kalimantan were compared on the basis of butterfly specimens deposited in the Tropical Rain Forest Research Center that were collected in and around the Bukit Soeharto Education Forest before the fires (1988,1995). On the basis of the post-fire survey, based on Malaise trap samples and field observations, only 43% of the butterfly species (not including Lycaenidae and Hesperiidae) were confirmed to have persisted. The data suggest that refugia that are not affected by fire are necessary for the conservation of specialist butterflies, as well as many other forms of wildlife. [source] An update on cyathostomins: Anthelmintic resistance and worm controlEQUINE VETERINARY EDUCATION, Issue 10 2008J. B. Matthews Summary Intestinal nematodes are an important cause of equine disease. Of these parasites, the Cyathostominae are the most important group, both in terms of their prevalence and their pathogenicity. Cyathostomin infections are complex and control is further complicated by ever-increasing levels of resistance to some of the commonly used anthelmintics. There are no new equine anthelmintics under development, so it is imperative that the efficacy of any currently-effective drug classes be maintained for as long as possible. It is believed that the proportion of refugia (i.e. the percentage of parasites not exposed to a drug at each treatment) is one of the most crucial factors in determining the rate at which anthelmintic resistance develops. It is important, therefore, that levels of refugia be taken into account when designing nematode control programmes for horses. This can be assisted by knowledge of the local epidemiology of the infection, supplemented by faecal egg count analysis to identify those animals that are making the major contribution to pasture contamination. This type of rational nematode control requires equine veterinary surgeons to get involved in designing and implementing deworming programmes. The advice given must be based on a combination of knowledge of cyathostomin biology and epidemiology as well as an awareness of the parasite population's current drug sensitivity and a sound history of husbandry at the establishment. As anthelmintic resistance will be the major constraint on the future control of cyathostomins, researchers are now actively investigating this area. Studies are underway to develop tests that will enable earlier detection of anthelmintic resistance and an assay that will help identify those horses that require anthelmintic treatments targeted at intestinal wall larvae. [source] Interpopulation Variation in the Social Organization of Female Collared Lizards, Crotaphytus collarisETHOLOGY, Issue 11 2003Troy A. Baird We tested the hypotheses that levels of intrasexual aggression and the social structure among neighboring females differed in two central Oklahoma populations of collared lizards, Crotaphytus collaris, and examined the extent to which variation in aggression might be related to differences in the availability of arthropods, elevated perches used by females to scan for prey, and crawlspace refugia. Because both the costs of aggression and access to resources may influence female fitness, we also compared growth and survival rates and the number of clutches produced. At Morningside Farms Ranch (MS), lizards occupied naturally-formed sandstone washes with naturally-sculpted irregular topographies, whereas they inhabited homogenous fields of boulders used to construct flood control spillways at the Arcadia Lake Dam (AL). The frequency of intrasexual aggression was markedly higher at MS, and groups of MS females had social hierarchies structured by size and age with older females defending territories, whereas no such social structure was apparent at AL. Moreover, experimental removal of individuals from female groups resulted in more pronounced changes by the remaining females at MS than at AL. Elevated perches and crawlspace refugia were much less abundant at MS. Arthropod availability was similar at the two sites, but at AL arthropods were clustered near the edges of rock patches where elevated perches overlooking adjacent grassy areas were particularly abundant. MS females showed lower rates of survival, and growth during the first year (when growth is highest) than AL females, whereas the number of clutches produced by females at the two sites was similar. Our results suggest that variation in the availability of perch rocks may have resulted in differences in female social structure at the two sites, and relaxed intrasexual competition for perches may have resulted in higher female fitness at AL. [source] THE COMBINED EFFECTS OF RIVERS AND REFUGIA GENERATE EXTREME CRYPTIC FRAGMENTATION WITHIN THE COMMON GROUND SKINK (SCINCELLA LATERALIS)EVOLUTION, Issue 2 2010Nathan D. Jackson Rivers can act as both islands of mesic refugia for terrestrial organisms during times of aridification and barriers to gene flow, though evidence for long-term isolation by rivers is mixed. Understanding the extent to which riverine barrier effects can be heightened for populations trapped in mesic refugia can help explain maintenance and generation of diversity in the face of Pleistocene climate change. Herein, we implement phylogenetic and population genetic approaches to investigate the phylogeographic structure and history of the ground skink, Scincella lateralis, using mtDNA and eight nuclear loci. We then test several predictions of a river,refugia model of diversification. We recover 14 well-resolved mtDNA lineages distributed east,west along the Gulf Coast with a subset of lineages extending northward. In contrast, ncDNA exhibits limited phylogenetic structure or congruence among loci. However, multilocus population structure is broadly congruent with mtDNA patterns and suggests that deep coalescence rather than differential gene flow is responsible for mtDNA,ncDNA discordance. The observed patterns suggest that most lineages originated from population vicariance due to riverine barriers strengthened during the Plio,Pleistocene by a climate-induced coastal distribution. Diversification due to rivers is likely a special case, contingent upon other environmental or biological factors that reinforce riverine barrier effects. [source] ESTIMATING A GEOGRAPHICALLY EXPLICIT MODEL OF POPULATION DIVERGENCEEVOLUTION, Issue 3 2007L. Lacey Knowles Patterns of genetic variation can provide valuable insights for deciphering the relative roles of different evolutionary processes in species differentiation. However, population-genetic models for studying divergence in geographically structured species are generally lacking. Since these are the biogeographic settings where genetic drift is expected to predominate, not only are population-genetic tests of hypotheses in geographically structured species constrained, but generalizations about the evolutionary processes that promote species divergence may also be potentially biased. Here we estimate a population-divergence model in montane grasshoppers from the sky islands of the Rocky Mountains. Because this region was directly impacted by Pleistocene glaciation, both the displacement into glacial refugia and recolonization of montane habitats may contribute to differentiation. Building on the tradition of using information from the genealogical relationships of alleles to infer the geography of divergence, here the additional consideration of the process of gene-lineage sorting is used to obtain a quantitative estimate of population relationships and historical associations (i.e., a population tree) from the gene trees of five anonymous nuclear loci and one mitochondrial locus in the broadly distributed species Melanoplus oregonensis. Three different approaches are used to estimate a model of population divergence; this comparison allows us to evaluate specific methodological assumptions that influence the estimated history of divergence. A model of population divergence was identified that significantly fits the data better compared to the other approaches, based on per-site likelihood scores of the multiple loci, and that provides clues about how divergence proceeded in M. oregonensis during the dynamic Pleistocene. Unlike the approaches that either considered only the most recent coalescence (i.e., information from a single individual per population) or did not consider the pattern of coalescence in the gene genealogies, the population-divergence model that best fits the data was estimated by considering the pattern of gene lineage coalescence across multiple individuals, as well as loci. These results indicate that sampling of multiple individuals per population is critical to obtaining an accurate estimate of the history of divergence so that the signal of common ancestry can be separated from the confounding influence of gene flow,even though estimates suggest that gene flow is not a predominant factor structuring patterns of genetic variation across these sky island populations. They also suggest that the gene genealogies contain information about population relationships, despite the lack of complete sorting of gene lineages. What emerges from the analyses is a model of population divergence that incorporates both contemporary distributions and historical associations, and shows a latitudinal and regional structuring of populations reminiscent of population displacements into multiple glacial refugia. Because the population-divergence model itself is built upon the specific events shaping the history of M. oregonensis, it provides a framework for estimating additional population-genetic parameters relevant to understanding the processes governing differentiation in geographically structured species and avoids the problems of relying on overly simplified and inaccurate divergence models. The utility of these approaches, as well as the caveats and future improvements, for estimating population relationships and historical associations relevant to genetic analyses of geographically structured species are discussed. [source] THE PHYLOGENETIC PATTERN OF SPECIATION AND WING PATTERN CHANGE IN NEOTROPICAL ITHOMIA BUTTERFLIES (LEPIDOPTERA: NYMPHALIDAE)EVOLUTION, Issue 7 2006Chris D. Jiggins Abstract Species level phylogenetic hypotheses can be used to explore patterns of divergence and speciation. In the tropics, speciation is commonly attributed to either vicariance, perhaps within climate-induced forest refugia, or ecological speciation caused by niche adaptation. Mimetic butterflies have been used to identify forest refugia as well as in studies of ecological speciation, so they are ideal for discriminating between these two models. The genus Ithomia contains 24 species of warningly colored mimetic butterflies found in South and Central America, and here we use a phylogenetic hypothesis based on seven genes for 23 species to investigate speciation in this group. The history of wing color pattern evolution in the genus was reconstructed using both parsimony and likelihood. The ancestral pattern for the group was almost certainly a transparent butterfly, and there is strong evidence for convergent evolution due to mimicry. A punctuationist model of pattern evolution was a significantly better fit to the data than a gradualist model, demonstrating that pattern changes above the species level were associated with cladogenesis and supporting a model of ecological speciation driven by mimicry adaptation. However, there was only one case of sister species unambiguously differing in pattern, suggesting that some recent speciation events have occurred without pattern shifts. The pattern of geographic overlap between clades over time shows that closely related species are mostly sympatric or, in one case, parapatric. This is consistent with modes of speciation with ongoing gene flow, although rapid range changes following allopatric speciation could give a similar pattern. Patterns of lineage accumulation through time differed significantly from that expected at random, and show that most of the extant species were present by the beginning of the Pleistocene at the latest. Hence Pleistocene refugia are unlikely to have played a major role in Ithomia diversification. [source] WHEN VICARS MEET: A NARROW CONTACT ZONE BETWEEN MORPHOLOGICALLY CRYPTIC PHYLOGEOGRAPHIC LINEAGES OF THE RAINFOREST SKINK, CARLIA RUBRIGULARISEVOLUTION, Issue 7 2004Ben L. Phillips Abstract Phylogeographic analyses of the fauna of the Australian wet tropics rainforest have provided strong evidence for long-term isolation of populations among allopatric refugia, yet typically there is no corresponding divergence in morphology. This system provides an opportunity to examine the consequences of geographic isolation, independent of morphological divergence, and thus to assess the broader significance of historical subdivisions revealed through mitochondrial DNA phylogeography. We have located and characterized a zone of secondary contact between two long isolated (mtDNA divergence > 15%) lineages of the skink Carlia rubrigularis using one mitochondrial and eight nuclear (two intron, six microsatellite) markers. This revealed a remarkably narrow (width<3 km) hybrid zone with substantial linkage disequilibrium and strong deficits of heterozygotes at two of three nuclear loci with diagnostic alleles. Cline centers were coincident across loci. Using a novel form of likelihood analysis, we were unable to distinguish between sigmoidal and stepped cline shapes except at one nuclear locus for which the latter was inferred. Given estimated dispersal rates of 90,133 m X gen,1/2 and assuming equilibrium, the observed cline widths suggest effective selection against heterozygotes of at least 22,49% and possibly as high as 70%. These observations reveal substantial postmating isolation, although the absence of consistent deviations from Hardy-Weinberg equilibrium at diagnostic loci suggests that there is little accompanying premating isolation. The tight geographic correspondence between transitions in mtDNA and those for nuclear genes and corresponding evidence for selection against hybrids indicates that these morphologically cryptic phylogroups could be considered as incipient species. Nonetheless, we caution against the use of mtDNA phylogeography as a sole criterion for defining species boundaries. [source] HISTORIC CYCLES OF FRAGMENTATION AND EXPANSION IN PARNASSIUS SMINTHEUS (PAPILIONIDAE) INFERRED USING MITOCHONDRIAL DNAEVOLUTION, Issue 1 2004Eric G. DeChaine Abstract Climate oscillations of the Quaternary drove the repeated expansion and contraction of ecosystems. Alpine organisms were probably isolated in sky island refugia during warm interglacials, such as now, and expanded their range by migrating down-slope during glacial periods. We used population genetic and phylogenetic approaches to infer how paleoclimatic events influenced the distribution of genetic variation in the predominantly alpine butterfly Parnassius smintheus. We sequenced a 789 bp region of cytochrome oxidase I for 385 individuals from 20 locations throughout the Rocky Mountains, ranging from southern Colorado to northern Montana. Analyses revealed at lease two centers of diversity in the northern and southern Rocky Mountains and strong population structure. Nested clade analysis suggested that the species experienced repeated cycles of population expansion and fragmentation. The estimated ages of these events, assuming a molecular clock, corresponded with paleoclimatic data on habitat expansion and contraction over the past 400,000 years. We propose that alpine butterflies persisted in an archipelago of isolated sky islands during interglacials and that populations expanded and became more connected during cold glacial periods. An archipelago model implies that the effects of genetic drift and selection varied among populations, depending on their latitude, area, and local environment. Alpine organisms are sensitive indicators of climate change and their history can be used to predict how high-elevation ecosystems might respond to further climate warming. [source] POSTGLACIAL DISPERSAL OF THE EUROPEAN RABBIT (ORYCTOLAGUS CUNICULUS) ON THE IBERIAN PENINSULA RECONSTRUCTED FROM NESTED CLADE AND MISMATCH ANALYSES OF MITOCHONDRIAL DNA GENETIC VARIATIONEVOLUTION, Issue 4 2002Madalena Branco Abstract Nested clade analysis was applied to cytochrome b restriction site data previously obtained on 20 natural populations of the European rabbit across the Iberian Peninsula to test the hypothesis of postglacial dispersal from two main refugia, one in the northeast and the other in the southwest. Apart from historical fragmentation that resulted in geographic discontinuity of two distinct mitochondrial DNA (mtDNA) clades A and B, patterns of haplotype genetic variability have been shaped mostly by restricted gene flow via isolation by distance. The distribution of tip versus interior haplotypes suggests that dispersal occurred from both the southwestern and northeastern groups. Dispersal from the southwest had a north and northwest direction, whereas from the northeast it had mostly a western and southern orientation, with subsequent overlap in a southeastern-northwestern axis across the Iberian Peninsula. The analysis of the pairwise mismatch distribution of a 179,181-bp fragment of the mtDNA control region, for seven of those populations, further supports the idea that major patterns of dispersal were in the direction of central Iberia. Additionally, rabbit populations do not show signs of any significant loss of genetic diversity in the recent past, implying that they maintained large population sizes and structure throughout the ice ages. This is congruent with the fact that the Iberian Peninsula was itself a glacial refugium during Quaternary ice ages. Nonetheless, climatic oscillations of this period, although certainly milder than in northern Europe, were sufficient to affect the range distributions of Iberian organisms. [source] TIME TO THE MOST RECENT COMMON ANCESTOR AND DIVERGENCE TIMES OF POPULATIONS OF COMMON CHAFFINCHES (FRINGILLA COELEBS) IN EUROPE AND NORTH AFRICA: INSIGHTS INTO PLEISTOCENE REFUGIA AND CURRENT LEVELS OF MIGRATIONEVOLUTION, Issue 1 2002Cortland K. Griswold Abstract We analyzed sequences from a 275-bp hypervariable region in the 5, end of the mitochondrial DNA control region in 190 common chaffinches (Fringilla coelebs) from 19 populations in Europe and North Africa, including new samples from Greece and Morocco. Coalescent techniques were applied to estimate the time to the most recent common ancestor (TMRCA) and divergence times of these populations. The first objective of this study was to infer the locations of refugia where chaffinches survived the last glacial episode, and this was achieved by estimating the TMRCA of populations in regions surrounding the Mediterranean that were unglaciated in the late Pleistocene. Although extant populations in Iberia, Corsica, Greece, and North Africa harbor haplotypes that are basal in a phylogenetic tree, this information alone cannot be used to infer that these localities served as refugia, because it is impossible to infer the ages of populations and their divergence times without also considering the population genetic processes of mutation, migration, and drift. Provided we assume the TMRCAs of populations are a reasonable estimate of a population's age, coalescent-based methods place resident populations in Iberia, Corsica, Greece, and North Africa during the time of the last glacial maximum, suggesting these regions served as refugia for the common chaffinch. The second objective was to determine when populations began diverging from each other and to use this as a baseline to estimate current levels of gene flow. Divergence time estimates suggest that European populations began diverging about 60,000 years before present. The relatively recent divergence of populations in North Africa, Italy, and Iberia may explain why classic migration estimates based on equilibrium assumptions are high for these populations. We compare these estimates with nonequilibrium-based estimates and show that the nonequilibrium estimates are consistently lower than the equilibrium estimates. [source] Reduction of a native fish fauna by alien species: an example from Brazilian freshwater tropical lakesFISHERIES MANAGEMENT & ECOLOGY, Issue 2 2004A. O. Latini Abstract The consequences of introducing Cichla cf. monoculus Spix & Agassiz, Astronotus ocellatus (Agassiz) and Pygocentrus nattereri Kner into lakes in the River Doce basin, Brazil, on richness, diversity and efficiency of aquatic macrophytes as natural refugia to native fishes was investigated. Samples were taken from lakes with and without alien fishes in areas with and without aquatic macrophytes. The presence of alien fishes reduced richness and diversity of the native fish community. The refugia function, which could be attributed to the clustering of aquatic macrophytes, does not exist in these lakes probably because the alien fishes exploit such habitats for reproduction. Since introductions threaten the native fish diversity of the region, studies on regional dispersion and factors that minimise the spread of alien fishes are needed. [source] The role of macroinvertebrates and fish in regulating the provision by macrophytes of refugia for zooplankton in a warm temperate shallow lakeFRESHWATER BIOLOGY, Issue 10 2010MARÍA de los ÁNGELES GONZÁLEZ SAGRARIO Summary 1.,The zooplankton often undergoes diel horizontal migration (DHM) from the open water to the littoral of shallow lakes, thus avoiding predators in the former. This behaviour has functional impacts within the lake, as it enhances zooplankton survival, increases their control of phytoplankton and tends to stabilise the clear water state. However, most of the evidence supporting this migration pattern comes from cold north temperate lakes, and more evidence from tropical and subtropical areas, as well as from southern temperate areas, is needed. 2.,We conducted a field study of the diel horizontal and vertical migration of zooplankton, and the horizontal distribution of potential predatory macroinvertebrates and fish, over two consecutive days in the summer in a temperate lake in the southern hemisphere. We took zooplankton samples at two depths, at three sampling stations (inside beds of aquatic macrophytes, at their edge and in open water) along three transects running from the centre of a bed of Ceratophyllum demersum to open water. At each sampling station, we also took samples of macroinvertebrates and fish and measured physical and chemical environmental variables. 3.,Zooplankton (pelagic cladocerans, calanoid copepods and rotifers) avoided the shore, probably because of the greater risk from predators there. Larger and more vulnerable cladocerans, such as Diaphanosoma brachyurum and Moina micrura, were two to four times more abundant in open water than at the edge of or inside beds of macrophytes, respectively, by both day and night. Less vulnerable zooplankton [i.e. of medium body size (Ceriodaphnia dubia) or with the ability to swim fast (calanoid copepods)] were distributed evenly between open water and the edge of the plant beds. Small zooplankton, Bosmina huaronensis and pelagic rotifers, showed an even distribution among the three sampling stations. Accordingly, no DHM of zooplankton occurred, although larger organisms migrated vertically inside C. demersum stands. 4.,Macrophytes contained high densities of predatory macroinvertebrates and fish. The predator assemblage, composed of large-bodied macroinvertebrates (including odonates and shrimps) and small littoral fish, was permanently associated with submerged macrophytes. None of these groups moved outside the plant beds or changed their population structure (fish) over the diel cycle. 5.,Submerged macrophyte beds do not represent a refuge for zooplankton in lakes where predators are numerous among the plants, implying a weaker top-down control of phytoplankton biomass by zooplankton and, consequently, a more turbid lake. The effectiveness of macrophytes as a refuge for zooplankton depends on the associated assemblage of predatory macroinvertebrates and fish among the plants. [source] Biodiversity of Belgian groundwater fauna in relation to environmental conditionsFRESHWATER BIOLOGY, Issue 4 2009PATRICK MARTIN Summary 1. The Pleistocene glaciations during the Quaternary appear to have resulted in an impoverished groundwater fauna in northern Europe. Re-colonisation may have occurred either through long-distance dispersal from unglaciated southern areas or from local refugia. 2. The Belgian groundwater fauna was sampled at multiple sites, and its habitats characterised, to assess whether the composition of present-day stygobiotic assemblages can be attributed to either of these mechanisms. 3. A total of 202 sampling sites were selected in four hydrogeographic units of the Meuse River catchment. Sites were equally divided among the saturated and unsaturated zones of fractured aquifers (karst) and within the hyporheic and phreatic zones of porous aquifers. Seventeen environmental parameters were determined in parallel. 4. More than 140 species were recorded, including representatives of the Amphipoda, Cladocera, Copepoda, Hydrachnidia, Isopoda, Oligochaeta, Ostracoda, Mollusca, Syncarida and Nematoda. Thirty stygobiont species were identified, of which 10 species were new to the Belgian fauna, raising the total number of stygobiotic species in Belgium to 41. 5. The frequency of occurrences of stygobiotic species was always low, with 37% of the sampled sites lacking stygobionts. A few species were exclusive to one hydrological zone, although no statistically significant differences were detected in species richness at any of the four hierarchial levels considered (Meuse catchment = region, hydrogeographic units, aquifer type and hydrological zone). 6. Overall, results suggest that the stygobiotic fauna of Belgium is species-poor and mostly comprises widely distributed species with broad ecological tolerances. This supports the view that eurytopic species re-colonised the area by long-distance dispersal from refugia in southern Europe. The virtual absence of endemic species further suggests that the scenario of an ancient fauna that survived in local refugia is of minor importance. [source] Predator,prey interactions in river networks: comparing shrimp spatial refugia in two drainage basinsFRESHWATER BIOLOGY, Issue 3 2009ALAN P. COVICH Summary 1.,Analysis of drainage networks provides a framework to evaluate the densities and distributions of prey species relative to locations of their predators. Upstream migration by diadromous shrimp (Atya lanipes and Xiphocaris elongata) during their life cycle provides access to headwater refugia from fish predation, which is intense in estuaries and coastal rivers. 2.,We postulate that geomorphic barriers (such as large, steep waterfalls >3.5 m in height), can directly limit the distribution of predatory fishes and, indirectly, affect the densities of their prey (freshwater shrimps) in headwater streams. 3.,We compared densities of shrimp in pools above and below waterfalls, in four headwater tributaries in two river basins of the Luquillo Mountains of northeastern Puerto Rico. We measured shrimp densities twice a year over 8 years (1998,2005) in Prieta, Toronja, Bisley 3 and Bisley 5 streams, which differ in drainage network positions relative to steep waterfalls in Río Espíritu Santo and Río Mameyes. 4.,Predatory fishes are absent in the Prieta and Toronja pools and present in Bisely 3 and in lower Bisley 5 pools. Atya lanipes and X. elongata rarely occur in the Bisley streams where predatory fishes are present but these shrimps are abundant in Prieta and Toronja, streams lacking predatory fishes. 5.,The mean carapace length of X. elongata is longer in pools where fish are present (Bisley 3 and lower Bisley 5) than in pools lacking fish (Prieta, Toronja, Upper Bisley 5). The increased body size is primarily due to significantly longer rostrums of individuals in stream reaches with fish (below waterfall barriers) than in those reaches lacking fish (above waterfall barriers). Rostrum length may be an adaptation to avoid predation by visually feeding fishes. 6.,Atya lanipes and X. elongata distributions and densities were predicted primarily by drainage network position relative to the presence or absence of predatory fishes. High, steep waterfalls effectively impeded fish from moving upstream and created a spatial refuge. Xiphocaris elongata may rely on size refugia (longer rostrum) to minimize predation where spatial refugia are lacking. [source] Testing species,stone area and species,bryophyte cover relationships in riverine macroinvertebrates at small scalesFRESHWATER BIOLOGY, Issue 3 2008JANI HEINO Summary 1. The species,area relationship is considered amongst the few genuine laws in ecology. Although positive species richness,stone area relationships have been found previously in stream systems, very few studies have simultaneously examined species,individuals, individuals,area, species,bryophyte biomass and individuals,bryophyte biomass relationships. We examined these relationships based on temporally replicated assessments of macroinvertebrates on stones at two river sites. 2. We found only one significant species,area relationship out of six relationship tested, and two significant individuals,area relationships. Even these significant relationships were weak, however. By contrast, we detected significant and rather strong relationships between species richness and the number of individuals at both river sites on all three sampling dates. We also found significant relationships of both species richness and the number of individuals with bryophyte biomass at both river sites on all sampling occasions. One of the river sites was disturbed by a bulldozer, and the species,bryophyte biomass relationships were somewhat stronger after the disturbance event. 3. Our findings are quite surprising, given that there were very weak species,area relationships on stream stones. By contrast, our results suggest a pivotal role for bryophyte biomass in determining the species richness and the number of individuals of stream macroinvertebrates at this small scale. The most probably origin of these relationships begins with bryophyte cover, which determines the number of individuals, and subsequently passively affects species richness. Thus, there is not necessarily a direct mechanism that determines the variability of species richness on stream stones. 4. Experimental studies are needed to disentangle the various mechanisms (e.g. passive sampling, provision of more food, more niche space, flood disturbance refugia) by which bryophyte biomass affects stream macroinvertebrates. [source] Linking ecological theory with stream restorationFRESHWATER BIOLOGY, Issue 4 2007P. S. LAKE Summary 1. Faced with widespread degradation of riverine ecosystems, stream restoration has greatly increased. Such restoration is rarely planned and executed with inputs from ecological theory. In this paper, we seek to identify principles from ecological theory that have been, or could be, used to guide stream restoration. 2. In attempts to re-establish populations, knowledge of the species' life history, habitat template and spatio-temporal scope is critical. In many cases dispersal will be a critical process in maintaining viable populations at the landscape scale, and special attention should be given to the unique geometry of stream systems 3. One way by which organisms survive natural disturbances is by the use of refugia, many forms of which may have been lost with degradation. Restoring refugia may therefore be critical to survival of target populations, particularly in facilitating resilience to ongoing anthropogenic disturbance regimes. 4. Restoring connectivity, especially longitudinal connectivity, has been a major restoration goal. In restoring lateral connectivity there has been an increasing awareness of the riparian zone as a critical transition zone between streams and their catchments. 5. Increased knowledge of food web structure , bottom-up versus top-down control, trophic cascades and subsidies , are yet to be applied to stream restoration efforts. 6. In restoration, species are drawn from the regional species pool. Having overcome dispersal and environmental constraints (filters), species persistence may be governed by local internal dynamics, which are referred to as assembly rules. 7. While restoration projects often define goals and endpoints, the succession pathways and mechanisms (e.g. facilitation) by which these may be achieved are rarely considered. This occurs in spite of a large of body of general theory on which to draw. 8. Stream restoration has neglected ecosystem processes. The concept that increasing biodiversity increases ecosystem functioning is very relevant to stream restoration. Whether biodiversity affects ecosystem processes, such as decomposition, in streams is equivocal. 9. Considering the spatial scale of restoration projects is critical to success. Success is more likely with large-scale projects, but they will often be infeasible in terms of the available resources and conflicts of interest. Small-scale restoration may remedy specific problems. In general, restoration should occur at the appropriate spatial scale such that restoration is not reversed by the prevailing disturbance regime. 10. The effectiveness and predictability of stream ecosystem restoration will improve with an increased understanding of the processes by which ecosystems develop and are maintained. Ideas from general ecological theory can clearly be better incorporated into stream restoration projects. This will provide a twofold benefit in providing an opportunity both to improve restoration outcomes and to test ecological theory. [source] Genetic diversity and historical population structure in the New Zealand mayfly Acanthophlebia cruentataFRESHWATER BIOLOGY, Issue 1 2006PETER J. SMITH Summary 1. Nucleotide sequences of a 280 base pair region of the cytochrome b gene were used to assess genetic diversity and to infer population histories in the New Zealand mayfly Acanthophlebia cruentata. 2. A hierarchial examination of populations from 19 streams at different spatial scales in the central and northern North Island of New Zealand found 34 haplotypes. A common haplotype was found in all central region streams and unique haplotypes in northern streams. Several central streams had region specific haplotypes with genetically differentiated populations at the 70,100 km scale. 3. Haplotype diversity was high (0.53,0.8) at most sites, but low (0,0.22) in some central sites. amova analyses found significant genetic diversity among regions (69%) and among catchments (58%). Most population pairwise FST tests were significant, with non-significant pairwise tests among sites in the central region and pairs of sites between neighbouring streams. 4. The levels of sequence divergence are interpreted as the result of Pleistocene divergence in multiple refugia, leading to the evolution of regionally unique haplotypes. The low diversity in some central region populations may result from recent colonisation following local extinctions, associated with volcanic events. [source] Hide, rest or die: a light-mediated diapause response in Daphnia magna to the threat of fish predationFRESHWATER BIOLOGY, Issue 1 2005Miros, lusarczyk Summary 1. In a laboratory batch culture experiment, a diapause response of Daphnia magna to a simulated threat of fish predation was tested at various light intensities, which under natural conditions determine potential vulnerability of Daphnia to visual planktivorous fish. 2. Under moderate light intensity (1.4 ,mol m,2 s,1) that allows effective predation by fish, the proportion of females producing dormant eggs was significantly higher than under dim light conditions (0.001 ,mol m,2 s,1) that are not favourable for visual detection of prey. Production of dormant eggs was not observed in complete darkness or in treatments missing fish kairomones, irrespective of tested light conditions. 3. The observed phenomenon is interpreted as a flexible response of prey to the conditional risk of predation assessed by Daphnia according to the presence of fish-derived cues on the one hand and the presence of dark refugia on the other. Irrespective of the presence of fish kairomones, Daphnia may not produce resting eggs as long as a safe, dark, bottom zone is accessible. [source] | ||||||||||||||||||||||||||||||||||||||||