Across the southern Andes on fin: glacial refugia,drainage reversals and a secondary contact zone revealed by the phylogeographical signal of Galaxias platei in Patagonia

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.     

Steganoderma szidati n. sp. (Trematoda: Zoogonidae) from Galaxias maculatus (Jenyns) and G. platei Steindachner in Patagonia,Argentina

Steganoderma szidati n. sp. is described from the intestine of two freshwater fishes, Galaxias maculatus and G. platei (Galaxiidae), from Andean lakes in Patagonia, Argentina. This freshwater zoogonid species fits in the genus Steganoderma (sensu Bray, 1987) because of the length of the caeca and the position of the vitelline follicles. The new species is characterised by possesing 6-13 vitelline follicles situated between the anterior margin of the ventral sucker and the poterior margin of the testis. The gonads are in the anterior hindbody and the ovary is anterior to the right testis. The cirrus has two conspicuous spines at its distal end, and the seminal vesicle always exhibits a constriction. The excretory vesicle never reaches the level of the posterior margin of the testes.     

Changes in the distribution of native fishes in response to introduced species and other anthropogenic effects

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. Methods We 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. Results The 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 conclusions We 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.     

Echoes of a distant time: effects of historical processes on contemporary genetic patterns in Galaxias platei in Patagonia

Interpreting the genetic structure of a metapopulation as the outcome of gene flow over a variety of timescales is essential for the proper understanding of how changes in landscape affect biological connectivity. Here we contrast historical and contemporary connectivity in two metapopulations of the freshwater fish Galaxias platei in northern and southernmost Patagonia where paleolakes existed during the Holocene and Pleistocene, respectively. Contemporary gene flow was mostly high and asymmetrical in the northern system while extremely reduced in the southernmost system. Historical migration patterns were high and symmetric in the northern system and high and largely asymmetric in the southern system. Both systems showed a moderate structure with a clear pattern of isolation by distance (IBD). Effective population sizes were smaller in populations with low contemporary gene flow. An approximate Bayesian computation (ABC) approach suggests a late Holocene colonization of the lakes in the northern system and recent divergence of the populations from refugial populations from east and west of the Andes. For the southern system, the ABC approach reveals that some of the extant G. platei populations most likely derive from an ancestral population inhabiting a large Pleistocene paleolake while the rest derive from a higher‐altitude lake. Our results suggest that neither historical nor contemporary processes individually fully explain the observed structure and geneflow patterns and both are necessary for a proper understanding of the factors that affect diversity and its distribution. Our study highlights the importance of a temporal perspective on connectivity to analyse the diversity of spatially complex metapopulations.     

Creating and destroying species: the 'new' biodiversity and evolutionarily significant units among New Zealand's galaxiid fishes

Morphological and genetic analyses of populations of two non-diadromous galaxiid species, the black mudfish and Neochanna heleios , the dwarf inanga Galaxias gracilis confirm the presence of a cryptic species, Neochanna heleios , within the geographic range of N. diversus , and indicate three distinct evolutionarily significant units (ESUs) in the latter species. Comparisons between G. gracilis and its parent taxon, G. maculatus , reveal that the former is not monophyletic and derives from three separate founding events. Significant morphological divergence between these two species is the result of ontogenic shifts in landlocked lacustrine populations and long-term morphological convergence in populations of G. gracilis. . Corresponding incipient changes are recognized in other younger landlocked populations of G. maculatus in New Zealand. These populations and the three groups within G. gracilis should be given the status of ESUs and managed accordingly by conservation agencies.     

A new genus of dactylogyrid from the gills of Galaxias maculatus (Osmeriformes: Galaxiidae) in Maullín Basin, Patagonia, Chile

During a parasitological survey of Galaxias maculatus (Jenyns, 1842) in the Maullín Basin (Chilean Patagonia), specimens of a new species of Monogenea were collected from the gills. This species is described as the only member of a proposed new genus, Inserotrema n. gen. (Dactylogyridae, Ancyrocephalinae), characterized by similar hooks with 2 subunits, overlapping gonads, coiled cirrus with counterclockwise rings, articulated accessory piece formed by 2 parts, a needlelike sclerite threading the distal part of the MCO, and a sclerotized midventral vagina. This new genus is proposed for dactylogyrids from gills of galaxiids (Galaxiidae). Inserotrema puyei n. sp. infects gills of G. maculatus from Llanquihue Lake, Maullín River, and Maullín Estuary. This is the first species of Ancyrocephalinae described from gills of a galaxiid.     

New metacercariae of the genus Tylodelphys (Trematoda, Diplostomatidae) in lacustrine populations of Galaxias maculatus (Teleostei, Galaxiidae)]

Metacercariae of three new species of Tylodelphys (Trematoda, Diplostomatidae) were found in the brain of Galaxias maculatus (Teleostei, Galaxiidae), at the Meliquina Lake, Neuquen Province, Argentina. These metacercariae are morphologically described as Tylodelphys argentinus n. sp., Tylodelphys barilochensis n. sp. and Tylodelphys crubensis n. sp. This is the first time that a stage of genus Tylodelphys is mentioned parasitizing Galaxias maculatus. Metacercariae of T. barilochen sis n. sp. were also found in G. maculatus brain at the Perito Moreno Lake, Rio Negro Province, Argentina.     

Reciprocal trophic niche shifts in native and invasive fish: salmonids and galaxiids in Patagonian lakes

1. Rainbow (Oncorhynchus mykiss) and brown trout (Salmo trutta) are widespread and invasive salmonids with important lethal effects as predators, although indirect effects are also possible. We used stable isotope analyses (δ¹⁵N, δ¹³C) to explore how the density of invasive trout in 25 Patagonian lakes alters the trophic niche (TN) of a widespread native fish, Galaxias platei (Galaxiidae). We also explored how the density of the galaxiid influences the TN of invasive trout. 2. We quantified two aspects of the TN: (i) the proportion of littoral carbon (PL) and (ii) trophic height (TH) (i.e. the ‘height’ at which the fish feeds in the food web). We related these measures of TN in a given species to the density of other species (as estimated by catch‐per‐unit‐effort). 3. As G. platei body size increased, their PL increased (increasing littoral feeding) in several lakes. However, none of the fish species investigated showed changes in PL with increasing density of the other fish species. TH increased with body size in all three species. In addition, the TH of large G. platei declined with increasing trout density and, reciprocally, the TH of large S. trutta decreased with decreasing G. platei density. 4. The reciprocal effects of native and the invasive fish on TH were as large as a shift of one trophic level. This pattern is consistent with an exhaustion of galaxiid prey for both piscivorous G. platei and S. trutta in lakes with high trout density. 5. These finding support the suggested management strategy of culling trout from overpopulated lakes, which should simultaneously protect native fish and enhance a lucrative sport fishery for large trout.     

Annual dynamics variation of a landlocked Galaxias maculatus (Jenyns 1842) population in a Northern Patagonian river: occurrence of juvenile upstream migration

Galaxias maculatus, broadly distributed in the southern hemisphere, presents both diadromic and landlocked populations. The upstream migration of larvae and juveniles from the sea to freshwater habitats is a characteristic phenomenon of diadromous populations, but upstream migration has never been reported in those that are landlocked. The objective of this study was to establish the population dynamics of a peculiar riverine landlocked population that presents an upstream migration at the larvae–juvenile transition from the Piedra del Águila Reservoir to the Caleufú River (Argentina). A spawning season from September to November and the arrival of shoals of metamorphic larvae and juveniles from February to April to the adult habitat were coincidental with lacustrine landlocked populations, but not with diadromous populations. Growth rate and age at migration, 147 ± 22.6 days, were also similar to other landlocked and diadromous populations. The arrival of these shoals produced a 20‐fold increase in fish density while the two cohorts of G. maculatus overlapped in time and space. No significant differences in morphology or vertebrae number were detected when riverine and reservoir adults were compared. The great life history plasticity of G. maculatus, shown to be even greater in our results, could be the key to explain the wide distribution of this species in the southern hemisphere. Likewise, discovering this juvenile potamodromous behaviour in a landlocked population will provide a new view for the analysis of the ways of this species’ dispersion in continental waters.     

Climate-induced changes to the ancestral population size of two Patagonian galaxiids: the influence of glacial cycling

Patagonia is one of the few areas in the Southern Hemisphere to have been directly influenced by Quaternary glaciers. In this study, we evaluate the influence that Quaternary glacial ice had on the genetic diversity of two congeneric fish species, the diadromous Galaxias maculatus and the nondiadromous Galaxias platei, using multilocus estimates of effective population size through time. Mid-Quaternary glaciations had far-reaching consequences for both species. Galaxias maculatus and G. platei each experienced severe genetic bottlenecks during the period when Patagonia ice sheet advance reached its maximum positions c. 1.1-0.6 Ma. Concordant drops in effective size during this time suggest that range sizes were under similar constraints. It is therefore unlikely that coastal (brackish/marine) environments served as a significant refuge for G. maculatus during glacial periods. An earlier onset of population declines for G. platei suggests that this species was vulnerable to modest glacial advances. Declines in effective sizes were continuous for both species and lasted into the late-Pleistocene. However, G. maculatus exhibited a strong population recovery during the late-Quaternary (c. 400,000 bp). Unusually long and warm interglacials associated with the late-Quaternary may have helped to facilitate a strong population rebound in this primarily coastal species.     

Genetic structuring in Andean landlocked populations of Galaxias maculatus: effects of biogeographic history

Aim To test whether the genetic diversity of diadromous and landlocked populations of the small puyen Galaxias maculatus (known as jollytail in Australia and inanga in New Zealand) follow the same structuring patterns observed for migratory and non‐migratory species of the genus Galaxias. This work also aimed to test whether the genetic structuring of a group of populations could be predicted from differences in the geomorphologic history of the region they inhabit. Location Eight landlocked populations were sampled from cold‐temperate lakes in north‐western Patagonia. The study area could be split latitudinally into two sectors that differed in their geomorphology, each of them hosting four populations. The southern sector shows evidence of a higher degree of glacial coverage, and the lakes are probably remnants of a big proglacial palaeolake. Lakes in the northern sector, on the other hand, suggest no common origin. Results Significant genetic structuring was found among the studied populations (Θ = 0.188), being the highest value reported to date for the species. Significant correlation was found between genetic diversity and lake area and perimeter. Diversity also showed a slight latitudinal variation suggesting the presence of genetically distinct groups of populations. The comparison of populations from the two geographical sectors showed that those from the north had a higher diversity, more private alleles and strong structuring, while those from the south were less diverse and much more homogeneous. Main conclusions Non‐migratory populations of G. maculatus show much higher values of genetic structuring than those reported for diadromous populations. This follows the pattern seen when comparing migratory and non‐migratory species of Galaxias. This agrees with population genetics theory which predicts that restricted gene flow would result in greater among‐population divergence. Also, differences between northern and southern populations agreed with what was predicted by the geomorphologic history of the study area. During the Last Glacial Maximum ice cover in that region may have reduced the habitat of G. maculatus to a refuge with an impoverished gene pool. When the ice receded, leaving a great proglacial lake, that former population expanded and became fragmented after water levels descended. This resulted in present day lakes harbouring homogeneous populations with reduced diversity. The northern sector, in contrast, was less affected by glaciers, resulting in more geomorphologically stable lakes holding genetically diverse populations.     

Evolution and biogeography of New Zealand's longjaw galaxiids (Osmeriformes: Galaxiidae): the genetic effects of glaciation and mountain building

1. The biological impact of glaciation in Southern Hemisphere freshwaters is poorly understood. Several large rivers of eastern South Island, New Zealand, represent a mosaic of glaciated and non-glaciated regions, and are thus well-suited for studies of post-glacial recolonization.
2. We conducted mtDNA analyses of South Island's endemic non-migratory longjaw galaxiids Galaxias prognathus and G. cobitinis (Osmeriformes: Galaxiidae) to test hypotheses of post-glacial recolonization, and to assess the vicariant effects of Pleistocene mountain building.
3. We analysed the phylogeography of longjaw cytochrome b sequences from 38 sites in central South Island ( n  = 83). On the basis of our sampling it seems that G. prognathus and G. cobitinis have a parapatric distribution in the Waitaki River system, their disjunction broadly coinciding with three large post-glacial lakes. Waitaki clades of both species are deeply divergent relative to conspecific taxa in drainages to the north and south.
4. Tests for recent population growth – predicted under post-glacial expansion of G. prognathus – do not refute recent recolonization of streams above glaciated lakes in the Waitaki River drainage. The apparent absence of potential 'source' populations from non-glaciated regions suggests a post-glacial population decline for G. prognathus below the Waitaki lakes.
5. Molecular clock calibrations based on several freshwater vicariant events elsewhere in New Zealand supported the geologically-derived hypothesis of Waitaki–Canterbury drainage isolation approximately 300 ka.     

Variation in vertebral number and its morphological implication in Galaxias platei

Variation in the vertebral number of the puyen grande Galaxias platei was examined for specimens from 22 localities that span the entire distribution range of the species (from 40° to 55° S). The mean vertebral number (N_MW) increases towards high latitudes, i.e. Jordan's rule is applicable to this species. Owing to the wide geographic variation of the species, not only in latitude but also in altitude, the most explicative variable for N_MW was mean winter air temperature, showing negative dependence. Morphological data suggest that the increment in vertebral number lies in the pre‐pelvic region of the trunk and in the caudal region, but not in the segment between pelvic‐fin insertion and the origin of the anal fin. As these alterations in body shape have important consequences for hydrodynamics and swimming performance, vertebral number variation in G. platei also holds implications for both individual and population fitness.     

Threatened native freshwater fishes in Australia–some case histories

Case histories of some of Australia's most threatened native freshwater fishes are presented. These include: six endangered species, Galaxias fontanus, G. johnsroni, G. pedderensis (Galaxiidae), Melanataenia euchamensis (Melanotaeniidae), Maccullochella sp. nov. and M. macquariensis (Percichthyidae): one vulnerable species, Galaxias tanycephalus: two potentially threatened species. Galaxias parvus and Prototroctes maraena (Prototroctidae); one indeterminate species. Maccllochella sp.; one restricted species, Macquaria australasica (Percichthyidae). Aspects of their taxonomy, distribution and reasons for their decline are discussed. Conservation management strategies that have been or are being applied to most of these species include protection of the fish and their habitats, establishment of refuge populations, and artificial propagation and re-establishment programmes.     

The relationship between fish assemblages and the helminth communities of a prey fish, in a group of small shallow lakes

Galaxias maculatus (small puyen) is an abundant native fish distributed in lakes and rivers of the Patagonia, and it is the frequent prey of other fishes, fish-eating birds, and mammals. Previous studies have shown that it is parasitized by 33 metazoan species and that the richness and composition of the parasite communities vary between lakes. The aim of the present work was to analyze the relationship between the composition of fish assemblages and the helminth component community structure of G. maculatus . Ten environmentally similar, small, shallow lakes, belonging to the Nahuel Huapi Lake basin, were chosen because of the differences in the native fish assemblages. Parasite community structure in G. maculatus varied according to the fish assemblage of each lake. The presence of the piscivorous fish Percichthys trucha regularly produced variations in the composition and richness at the component and infracommunity levels, as well as the percentage of autogenic parasite species in G. maculatus .     

Dispersal and recruitment of fish in an intermittent stream network

Animal movement is an important process connecting habitats in heterogeneous landscapes, and can play a key role in population persistence. Laboratory swim trials were conducted to determine and compare the dispersal capabilities of two native Australian fish, mountain galaxias (Galaxias olidus, Family Galaxiidae) and southern pygmy perch (Nannoperca australis, Family Nannopercidae) that maintain populations in hydrologically variable and intermittently flowing streams in south‐eastern Australia. These experiments showed that G. olidus had significantly greater swimming endurance under a range of flow velocities. Concurrent field surveys were used to establish whether swimming abilities observed in laboratory studies were consistent with patterns of inferred movement from distribution and abundance patterns observed in the field. Data collected at multiple sites from headwater to lowland reaches along multiple streams revealed substantial temporal changes in the distribution of young‐of‐year (0+) G. olidus, with spawning occurring at upland sites in winter, followed by downstream larval migration and subsequent upstream movement in late spring. Observed spatial and temporal patterns in G. olidus abundances were consistent with a source‐sink population structure, which may be disrupted by prolonged cease‐to‐flow periods during drought years. In contrast, results for N. australis suggested limited dispersal, with restricted local populations that persist at sites with permanent surface water. These field and laboratory findings complement our understanding of the spatial population structure of these two species in intermittent streams, and highlight the importance of understanding the role of dispersal in species conservation and habitat restoration.     

Acanthocephalus tumescens (Acanthocephala, Echinorhynchidae) in Galaxias maculatus (Pisces, Galaxiidae) of Lake Gutiérrez, Patagonia, Argentina

The seasonal distribution of Acanthocephalus tumescens (Acanthocephala : Echinorhynchidae) among Galaxias maculatus (Pisces : Galaxiidae) in Lake Gutiérrez was studied from March 1994 to June 1996. Acanthocephalus tumescens always occurs in the intestine, has an overdispersed frequency distribution, a similar proportion of sexes, and females are larger than males. Mean intensity and prevalence are low and increase with host length. The pattern of the infection shows seasonality, with recruitment in winter and a reproductive period during spring-summer.     

Allocreadium pichi n. sp. (Trematoda: Allocreadiidae) in Galaxias maculatus (Osteichthyes: Galaxiidae) from Lake Moreno in Patagonia (Argentina)

Allocreadium pichi n. sp. (Trematoda: Allocreadiidae) is described from the intestine of Galaxias maculatus (Osteichthyes: Galaxiidae) from Moreno Lake in Patagonia, `Argentina. This species is distinguished from A. patagonicum Shimazu, Urawa &; Coria, 2000, the other species recorded in the area, by its smaller body size, the diagonal position of the testes and different ratios of the suckers, testes and cirrus-sac in relation to body size. In addition, the synonymy of A. patagonicum withPolylekithum percai Ostrowski de Núñez, Brugni &; Viozzi, 2000 is proposed herein.     

Effects of ultraviolet radiation on the eggs of landlocked Galaxias maculatus (Galaxiidae, Pisces) in northwestern Patagonia

• 1 Ultraviolet radiation (UVR) damages early life stages of several fish species. Galaxias maculatus is a small catadromous fish, with landlocked forms occurring in many lakes within the Nahuel Huapi National Park (Patagonia, Argentina). In this work, the vulnerability of G. maculatus eggs exposed to both natural and artificial UVR was investigated in relation to water transparency.
• 2 Field experiments were performed in two lakes differing in UVR attenuation. Galaxias maculatus eggs were exposed to in situ levels of UVR in quartz tubes incubated at various depths. For laboratory experiments, the eggs were exposed to five levels of artificial UVB radiation.
• 3 Exposure to natural UVR causes various degrees of egg mortality depending on water transparency and incubation depth. In the less transparent lake (K_d320 = 3.08 m^‐1), almost complete mortality was observed near the surface. At a depth of 43 cm the observed mortality was only 22%, but was still significantly different from the dark control. In the most transparent lake (K_d320 = 0.438 m^‐1), almost total mortality was observed in tubes incubated at 2.56 m or shallower. A gradual decline in mortality was recorded from that depth to 3.78 m where the values approached those in the dark control treatments.
• 4 A monotonic relationship between mortality and UV exposure could be observed both in field and laboratory experiments. Using the results from field incubations, a LD₅₀ of 2.5 J cm^‐2 nm^‐1 was estimated. In a few mountain lakes, this value would be exceeded even if the eggs were laid at the maximum depth of the lake. Thus UVR seems a sufficient cause to explain the absence of G. maculatus populations in some mountain lakes. For most lakes, however, UVR is probably one of several important environmental factors, which together determine the habitat suitability.