We present four lines of evidence that the magnitude of prey exchange (=immigration/emigration) among substrate patches has an overwhelming influence on the perceived effects of predators on prey populations. (1) An extensive review of the literature on predation effects in benthic and littoral freshwater revealed a significant relationship between prey exchange rate and observed predator impact. In streams, studies showing significant predator effects used cages with smaller mesh sizes than studies showing nonsignificant effects. Similarly, there was a highly significant correlation between cage mesh size and the magnitude of predator impact on common prey. Large—scale stream studies indicated that prey drift and colonization rate were inversely related to predator impact on benthic prey. (2) These patterns were confirmed by field experiments and observations where mesh size was directly manipulated or where exchange rates varied among taxa. In Colorado streams we saw greater predator impacts on Baetis prey when immigration/emigration was restricted vs. when the mesh size of the cage was relatively large. Similarly, the effects of trout in California stream pools were greater when prey turnover rates were low. (3) A re—analysis of Peckarsky's (1985) data shows an inverse relationship between predator impact and prey mobility within a field experiment. (4) Finally, a model that incorporates both predation and exchange of prey indicates that we ought to expect a lower magnitude of predator effects when exchange rates are high. These results suggest that some discrepancies in past studies may be explained by differences in the exchange rates of prey, and that differences in predator effects across different systems or habitats may be related to variation in the rates of prey dispersal and colonization.

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We used molecular evidence to examine the roles that vicariance mechanisms (mountain-building and drainage changes during the Pleistocene) have played in producing phylogeographical structure within and among South American fish species of the temperate perch family Percichthyidae. The percichthyids include two South American genera, Percichthys and Percilia, each containing several species, all of which are endemic to southern Argentina and Chile (Patagonia). Maximum-likelihood phylogenies constructed using mitochondrial DNA (mtDNA) control region haplotypes and nuclear GnRH3-2 intron allele sequences support the current taxonomy at the genus level (both Percichthys and Percilia form strongly supported, monophyletic clades) but indicate that species-level designations need revision. Phylogeographical patterns at the mtDNA support the hypothesis that the Andes have been a major barrier to gene flow. Most species diversity occurs in watersheds to the west of the Andes, together with some ancient divergences among conspecific populations. In contrast, only one species (Percichthys trucha) is found east of the Andes, and little to no phylogeographical structure occurs among populations in this region. Mismatch analyses of mtDNA sequences suggest that eastern populations last went through a major bottleneck c. 188 000 bp, a date consistent with the onset of the penultimate and largest Pleistocene glaciation in Patagonia. We suggest that eastern populations have undergone repeated founder-flush events as a consequence of glacial cycles, and that the shallow phylogeny is due to mixing during recolonization periods. The area of greater diversity west of the Andes lies outside the northern limit of the glaciers. mtDNA mismatch analysis of the genus Percilia which is restricted to this area suggests a long-established population at equilibrium. We conclude that patterns of genetic diversity in these South American genera have been primarily influenced by barriers to gene flow (Andean orogeny, and to a lesser extent, isolation in river drainages), and by glacial cycles, which have resulted in population contraction, re-arrangement of some watersheds, and the temporary breakdown of dispersal barriers among eastern river systems.

We employed DNA sequence variation at two mitochondrial (control region, COI) regions from 212 individuals of Galaxias platei (Pisces, Galaxiidae) collected throughout Patagonia (25 lakes/rivers) to examine how Andean orogeny and the climatic cycles throughout the Quaternary affected the genetic diversity and phylogeography of this species. Phylogenetic analyses revealed four deep genealogical lineages which likely represent the initial division of G. platei into eastern and western lineages by Andean uplift, followed by further subdivision of each lineage into separate glacial refugia by repeated Pleistocene glacial cycles. West of the Andes, refugia were likely restricted to the northern region of Patagonia with small relicts in the south, whereas eastern refugia appear to have been much larger and widespread, consisting of separate northern and southern regions that collectively spanned most of Argentinean Patagonia. The retreat of glacial ice following the last glacial maximum allowed re-colonization of central Chile from nonlocal refugia from the north and east, representing a region of secondary contact between all four glacial lineages. Northwestern glacial relicts likely followed pro-glacial lakes into central Chilean Patagonia, whereas catastrophic changes in drainage direction (Atlantic --> Pacific) for several eastern palaeolakes were the likely avenues for invasions from the east. These mechanisms, combined with evidence for recent, rapid and widespread population growth could explain the extensive contemporary distribution of G. platei throughout Patagonia.

Changes in lake and stream habitats during the growth and retreat of Pleistocene glaciers repeatedly altered the spatial distributions and population sizes of the aquatic fauna of the southern Andes. Here, we use variation in mtDNA control region sequences to infer the temporal dynamics of two species of southern Andean fish during the past few million years. At least five important climate events were associated with major demographic changes: (i) the widespread glaciations of the mid-Pliocene (c. 3.5 Ma); (ii) the largest Patagonian glaciation (1.1 Ma); (iii) the coldest Pleistocene glaciation as indicated by stacked marine delta(18)O (c. 0.7 Ma); (iv) the last southern Patagonian glaciation to reach the Atlantic coast (180 ka); and (v) the last glacial maximum (LGM, 23-25,000 years ago). The colder-water inhabitant, Galaxias platei, underwent a strong bottleneck during the LGM and its haplotype diversity coalesces c. 0.7 Ma. In contrast, the more warm-adapted and widely distributed Percichthys trucha showed continuous growth through the last two glacial cycles but went through an important bottleneck c. 180,000 years ago, at which time populations east of the Andes may have been eliminated. Haplotype diversity of the most divergent P. trucha populations, found west of the Andes, coalesces c. 3.2 Ma. The demographic timelines obtained for the two species thus illustrate the continent-wide response of aquatic life in Patagonia to climate change during the Pleistocene, but also show how differing ecological traits and distributions led to distinctive responses.

BackgroundThe dynamic geological and climatic histories of temperate South America have played important roles in shaping the contemporary distributions and genetic diversity of endemic freshwater species. We use mitochondria and nuclear sequence variation to investigate the consequences of mountain barriers and Quaternary glacial cycles for patterns of genetic diversity in the diadromous fish Galaxias maculatus in Patagonia (~300 individuals from 36 locations).ResultsContemporary populations of G. maculatus, east and west of the Andes in Patagonia, represent a single monophyletic lineage comprising several well supported groups. Mantel tests using control region data revealed a strong positive relationship when geographic distance was modeled according to a scenario of marine dispersal. (r = 0.69, P = 0.055). By contrast, direct distance between regions was poorly correlated with genetic distance (r = -0.05, P = 0.463). Hierarchical AMOVAs using mtDNA revealed that pooling samples according to historical (pre-LGM) oceanic drainage (Pacific vs. Atlantic) explained approximately four times more variance than pooling them into present-day drainage (15.6% vs. 3.7%). Further post-hoc AMOVA tests revealed additional genetic structure between populations east and west of the Chilean Coastal Cordillera (coastal vs. interior). Overall female effective population size appears to have remained relatively constant until roughly 0.5 Ma when population size rapidly increased several orders of magnitude [100× (60×-190×)] to reach contemporary levels. Maximum likelihood analysis of nuclear alleles revealed a poorly supported gene tree which was paraphyletic with respect to mitochondrial-defined haplogroups.ConclusionsFirst diversifying in the central/north-west region of Patagonia, G. maculatus extended its range into Argentina via the southern coastal regions that join the Pacific and Atlantic oceans. More recent gene flow between northern populations involved the most ancient and most derived lineages, and was likely facilitated by drainage reversal(s) during one or more cooling events of the late Pleistocene. Overall female effective population size represents the end result of a widespread and several hundred-fold increase over approximately 0.5 Ma, spanning several climatic fluctuations of the Pleistocene. The minor influence of glacial cycles on the genetic structure and diversity of G. maculatus likely reflects the access to marine refugia during repeated bouts of global cooling. Evidence of genetic structure that was detected on a finer scale between lakes/rivers is most likely the result of both biological attributes (i.e., resident non-migratory behavior and/or landlocking and natal homing in diadromous populations), and the Coastal Cordillera as a dispersal barrier.

Aim Globally, one of the major threats to the integrity of native faunas is the loss of biodiversity that can result from the introduction of exotics. Here we document recent changes in the distribution of five common fish species that are linked to introductions in Chile.Location Chile from 28°S to 54°S. MethodsWe assess the extent of changes in distribution of galaxiid species by comparing their historical and current distributions based on the results of the most extensive survey of freshwater fishes in Chile to date, a range that encompasses the full latitudinal and elevational range of the Galaxiidae in Chile. We test for relationships of the distributions and abundances of native fishes with the incidence of introduced species. ResultsThe latitudinal range of Galaxias maculatus has declined by 26%, and most of this reduction has occurred in the northern part of its range. Aplochiton taeniatus and Brachygalaxias bullocki have experienced reductions (8-17% loss) in total drainage area occupied, and they have disappeared from, or are now extremely difficult to find, in latitudes 36°to 41°S, coincidently with areas of urban growth and intense economic activities. The distribution of Galaxias platei has, instead, increased considerably. In northern basins, G. maculatus has apparently been replaced by an introduced poeciliid Gambusia sp. High-elevation systems remain dominated by native Galaxias platei, whereas systems at intermediate elevations, especially rivers, are now dominated by introduced salmonids. Within drainages, native galaxiids remain abundant where exotic salmonid abundance is low. Main conclusionsWe suggest that negative interactions between introduced and native fish are responsible for some of the range reductions among Galaxiidae in Chile. The severity of the impacts varies with latitude and altitude and is probably related to temperature. The effects of Gambusia are restricted to warmer systems. Native fish also appear to have found temperature refugia from salmonids; impacts are low in the warmer northern and coastal systems, as well as in high-altitude relatively cold systems. Native fish also appear less vulnerable to salmonids in lakes than in rivers. This study identifies watersheds critical for the conservation of biodiversity within the Galaxiidae.

Current patterns of genetic and morphological diversity are the product of historical climatic and geomorphological events, and of contemporary selection processes acting upon this diversity. Here we examine the phylogeographic and phenotypic patterns of diversity within Percichthys trucha, a widely distributed Patagonian fish species complex that inhabits Andean and steppe freshwater environments. Molecular analysis (mtDNA control region) of 21 populations distributed throughout its latitudinal range revealed little evidence of phylogeographic structure and no evidence of species-level genetic divergence east of the Andes. The complex, however, exhibits high levels of intra-and interpopulation phenotypic variation. Patterns of among-population divergence in morphology were most easily explained by differences in predation pressure among populations; dorsal fin spines (commonly a defensive characteristic) were longer in environments with greater densities of potentially piscivorous fish. Trophic characters were highly variable within populations, suggesting an important role for resources in generating within-population morphological variation. The very shallow levels of divergence shown by the molecular data most likely reflect the historical mixing of populations as a result of the climatic and landscape changes that affected Patagonia throughout the Quaternary. The phenotypic divergences, in contrast, are probably the result of differing contemporary selection regimes acting on currently disjoint populations.Los patrones de diversidad genética y morfológica que pueden observarse en poblaciones existentes son el producto de la influencia conjunta de procesos históricos (climáticos, geomorfológicos) y de la selección natural. En este trabajo examinamos los patrones de diversidad filogeográfica y fenotípica en Percichthys trucha, una especie o complejo de especies de amplia distribución en Patagonia andina y esteparia. Análisis molecular (Region de Control ADN mitocondrial) de 21 poblaciones a lo largo y ancho del rango distribucional del grupo reveló poca evidencia de estructura filogeográfica (estructura poco profunda) y ninguna evidencia de divergencia genética a nivel de especie al este de los Andes. El complejo exhibe sin embargo, altos niveles de variación fenotípica tanto intra-, como interpoblacional. Los patrones de divergencia morfológica entre poblaciones se correlacionan con diferencias interpoblacionales en la intensidad de predación; las espinas dorsales (comúnmente una característica defensiva) son más largas en ambientes con mayor densidad de peces potencialmente piscívoros. Los caracteres tróficos exhiben alta variación intrapoblacional sugiriendo que los recursos tróficos cumplen un rol importante en