Victor E. Cussac

Climate control on ancestral population dynamics: insight from Patagonian fish phylogeography

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 δ18O (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.

Phylogeography of the Percichthyidae (Pisces) in Patagonia: roles of orogeny, glaciation, and volcanism

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.

Freshwater fishes of Patagonia in the 21st Century after a hundred years of human settlement, species introductions, and environmental change

We review the status of the freshwater fish fauna of Patagonia, an assemblage with 26 native species, comprising fishes of Gondwanan origin, marine dispersants, and oceanic elements of local origin. Several processes, old and new, have shaped the landscape of Patagonia and its fauna: a Gondwanan heritage, the Andes uplifting, Pleistocene ice, volcanic activity, introduction of exotic fishes, mostly Salmonids, and climate change. While there is a significant tradition of taxonomic work on native fish species, research on life history, trophic relationships, and community structure has started to emerge only in the last 15 years. Most studies were conducted in oligotrophic lakes of the Andes; while fauna of streams remains poorly observed. While documentation of impacts by salmonids is scarce, there is some compelling evidence indicating that freshwater communities have been significantly shaped by exotic fish. Impacts by exotic species appear to be dependent on temperature on the east side of the Andes, and land use and watershed perturbation on the west side. In general, freshwater habitat conditions and how they affect fishes are poorly studied. In lakes, habitat complexity and its specialized use by native fishes may have ameliorated the impact by introduced salmonids. Although impacts on rivers abound, led by dam construction, the relationship between stream habitat integrity and native species health is still poorly understood. The future of freshwater resources will largely depend on how able we are to inform managers, the general public and colleagues about their value and the costs of not taking action. But current research capacity is insufficient to deal with most demands because of limitations in people, resources and baseline information. To support our claims, we need to promote regional assessments of freshwater resources and of major threats to their integrity, the building blocks of a regional agenda for their sustainable use.

Spatial and temporal distribution of landlocked Galaxias maculatus and Galaxias platei (Pisces: Galaxiidae) in a lake in the South American Andes

Abstract Galaxiids are present in many of the Andean lakes in southern South America. We studied landlocked Galaxias maculatus (Jenyns) and Galaxias platei Steindachner populations in a deep oligotrophic lake (Lake Gutiérrez, Patagonia, Argentina). Their temporal and spatial distribution, intralacustrine movements (horizontal and vertical), and spawning periods were analysed using several sampling techniques (ichthyoplankton net, seine net, gill net, and baited benthic taps). We identified the early life stages of both species based on their morphology and otolith shape. The free embryos of both species migrate to the limnetic zone, where they coexist as larvae, facing the same food availability and probably the same predation risk. Each species then moves on to its own juvenile and adult habitat: the littoral and benthic zone for G. maculatus and only the deeper benthic zone for G. platei. Their adult habitats and part of their spawning periods partly overlap.

Distribution of introduced and native fish in Patagonia (Argentina): patterns and changes in fish assemblages

The interaction between native fishes and salmonids introduced in Patagonia at the beginning of the 20th Century, developed at the same time as the environmental change. The phenomenon of global warming has led to the formulation of predictions in relation to changes in the distribution of species, in the latitudinal dimension, both at intralacustrine, or small streams levels. The aim of the present work includes three main objectives: a) to compose a general and updated picture of the latitudinal distribution range of native and alien fishes, b) to analyze the historical changes in the relative abundance of Percichthys trucha, Odontesthes sp., and salmonids in lakes and reservoirs, and c) to relate the diversity and relative abundance of native and salmonid fishes to the environmental variables of lakes and reservoirs. We analysed previous records and an ensemble of data about new locations along the northern border of the Patagonian Province. We compared current data about the relative abundance of native fishes and salmonids in lakes and reservoirs, with previous databases (1984–1987). All samplings considered were performed during spring-summer surveys and include relative abundance, as proportions of salmonids, P. trucha, and Odontesthes sp. For the first time, we found changes in fish assemblages from twenty years back up to the present: a significant decline in the relative abundances of salmonids and an increase of P. trucha. We studied the association between the diversity and relative abundance of native and salmonid fishes and the environmental variables of lakes and reservoirs using Canonical Correspondence Analysis. Relative abundance showed mainly geographical cues and the diversity relied largely on morphometric characteristics. Relative abundance and diversity seem to have a common point in the lake area, included into the PAR concept. Native abundance and alien diversity were negatively related with latitude. Greater native diversity was observed in lakes with high PAR compared with salmonids. Historical changes such as southward dispersion, relative abundance changes, and geographical patterns for relative abundance and diversity are basic concepts needed not only in future research but also in management design for Patagonian fish populations.

Intralacustrine movements of Galaxias maculatus (Galaxiidae) and Ondontesthes microlepidotus (Atherinidae) during their early life history

SynopsisGalaxias maculatus and Odontesthes microlepidotus undertook significant habitat shifts in a small Araucanian lake during their early life history. After hatching in the littoral zone, free embryos migrate to the limnetic zone. Later, larvae return to the littoral. A third movement, littoral-limnetic, is suggested through acoustic records of fish in the limnetic zone which correspond with the growth curve of littoral sampled juveniles.

Ontogenetic shifts in the diet ofGalaxias maculatus (Galaxiidae) andOdontesthes microlepidotus (Atherinidae)

SynopsisThe feeding habits ofGalaxias maculatus (Galaxiidae) andOdontesthes microlepidotus (Atherinidae) larvae and juveniles were studied in an Araucanian lake of Argentina. Fish specimens were grouped, on the basis of the percentage of each food category found in their gut, using multivariate methods. These groups of fishes showed significant differences in average total length. Positive correlations between prey size and fish length were also detected. Diet overlap between juvenileG. maculatus and juvenileO. microlepidotus was observed. We discuss the interaction between the two species and the role of functional capabilities for prey capture during these early life periods.

Fishes of southern South America: a story driven by temperature.

The latitudinal extension of southern South America imposes a thermal gradient that affects the structure of marine and freshwater fish assemblages and the biology of the species through direct exposure to the temperature gradients or by means of a web of historical and ecological relationships. We have reviewed biological and ecological data of marine and freshwater fishes from the southern Neotropics, including Patagonia, and report several examples of dependence on temperature, from glacial times to today’s climate change. We were able to identify historic and present effects on the diversity of fish assemblages, isolation, southern limits for the distribution of species, and morphological variation among populations. There is a wide range of characteristics that exemplify an adaptation to low temperatures, including biochemical peculiarities, physiological adjustments, and alternative life history patterns, and these appear in both freshwater and marine, and native and exotic fishes. The consequences of stable temperature regimes in both the ocean and thermal streams deserve special mention as these shape specialists under conditions of low selective pressure. At present, habitat use and interactions among species are being subject to changes as consequences of water temperature, and some of these are already evident in the northern and southern hemispheres.

Body shape variation of three species of Percichthys in relation to their coexistence in the Limay River basin, in northern Patagonia

We describe morphological variation within and among three species of Percichthys (Perciformes, Percichthyidae) inhabiting two reservoirs within a single river drainage (Limay River) in Argentina. Two small-mouthed percas’, P. vinciguerrai and P. trucha were found in both reservoirs (Alicura and Piedra del Aguila), and a third species, the large-mouthed perca, P. colhuapiensis, was found in Piedra del Aguila. There was some specialization in diet where the three species coexisted. The large-mouthed perca ate a higher proportion of large prey (>25 mm) and a smaller proportion of small prey (< 15 mm) than did the small-mouthed species, suggesting some diet specialization based on prey size. Percichthys trucha and P. vinciguerrai had smaller gapes, as indicated by shorter heads and jaws, and narrower mouths, in Piedra del Aguila where they coexisted with the large-mouthed species, than in Alicura where the competitor was virtually absent. Thus the direction of morphological changes in trophic-related characters in P. trucha and P. vinciguerrai is consistent with the hypothesis of character shift due to interspecific competition.

Lethal temperatures of a Neotropical fish relic in Patagonia, the scale-less characinid Gymnocharacinus bergi

Southern South America has a rather low fish species diversity. Gymnocharacinus bergi, the southernmost characid fish of the world, is the only member of Characoidei in the Argentine Patagonia. The isolation of this species in an endorheic stream has been linked to the thermal conditions of its habitat, the head-waters of the Valcheta Stream, which is the only site where this species occurs. We provide information on the distribution and thermal habitat of this species and other fishes in the Valcheta Stream. The responses of G. bergi to high and low temperatures were assessed in the laboratory under different temperatures and heating and cooling rates. Our results suggest that G. bergi is unable to extend its distribution to the colder waters nearby, as well as to waters with greater temperature fluctuations. We discuss the implications of our experimental data, the habitat of G. bergi, and the known responses of a few other paranensean fishes to temperature, within the framework of the thermal ecology of freshwater fishes.