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.

Food availability and predation risk have been shown to affect phenotypes during early life history of fishes. Galaxias maculatus, a small fish widely distributed around the southern hemisphere, clearly exhibits a complex trade-off between feeding and predation avoidance during growth over the larval period. We studied the effect of different environmental variables on diet, growth, mortality, and morphology through field surveys and data revision in the literature for limnetic G. maculatus larvae in five oligotrophic lakes of Patagonia. Both number of food categories and prey ingested by larvae were directly related to zooplankton density. Larval growth rate was related with zooplankton density and temperature. Lakes with high zooplankton densities and low predation risk had larvae with deeper bodies and shorter caudal peduncles, while in lakes with less food and high predation risk larvae were slender with shallower bodies and longer peduncles. Food availability and predation risk seem to operate on the swimming performance of G. maculatus larvae through the slenderness of the body and the length of the caudal peduncle. The observed phenotypic variation in growth and morphology could be a key feature that has allowed this species to successfully colonize a wide variety of environments in the southern hemisphere.

Summary 1. Understanding the influence of landscape characteristics on genetic and phenotypic intraspecific variability can yield insights into how evolutionary processes work as well as provide essential information for the conservation of biodiversity. 2. Our aim was to compare the genetic structure and phenotypic variation among Galaxias maculatus populations inhabiting two Atlantic Ocean river basins in Patagonia in relation to historical and contemporary landscape characteristics associated with latitude. 3. Population genetic analysis (based on eight microsatellite loci, 505 individuals) indicates that genetic structure is more pronounced in the Río Negro basin (RN) at 39–41°S (10 localities) than in the southernmost Santa Cruz River basin (SCR) at 49–50°S (seven localities). Spatial autocorrelation analysis showed different effects of geographic distance in shaping population differentiation patterns in the two basins. 4. While in northern Patagonia, fragmented populations could have survived the severity of Quaternary climate cycles, at higher latitudes G. maculatus populations were probably extirpated by extensive ice sheets. Extant populations in the upper reaches of the southernmost basin probably originated from refugia close to the Atlantic Ocean. 5. Based on otolith microchemistry, we have documented facultative diadromy for the first time for the species in the SCR basin, while no evidence of migration to the sea was found in individuals from the RN basin. Vertebral number increased with latitude, and within the SCR basin, the higher counts are possibly associated with a migratory life style.