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.     

Density-dependent population dynamics in larvae of the dragonfly Pachydiplax longipennis: a field experiment

Summary Several features of dragonfly population biology suggest that population regulation occurs in the larval stage. This study was designed to determine if density-dependent interactions among larval odonates can affect survival, growth and emergence. First-instar larvae of the dragonfly Pachydiplax longipennis were raised in outdoor experimental ponds at initial densities of 38, 152, and 608 larvae · m^-2, under two levels of food availability. Food availability was supplemented in half the pools by volumetric addition of zooplankton every other day. Pools in the low food treatment did not receive the zooplankton supplement.There was a strong negative effect of density on the mean growth rate of survivors, which included both emerging tenerals and individuals overwintering in the larval stage. A higher proportion emerged from low density than high density pools. Metamorphs from high density populations were smaller and emerged slightly later than those from low density, but the absolute number of metamorphs did not differ significantly among density treatments. Food supplementation significantly increased the proportion of overwintering larvae. There were no significant food-by-density interactions, indicating that food and density acted independently on larval population dynamics. Density-dependent mechanisms can clearly contribute to odonate population regulation, especially by controlling the number of larvae which emerge and the average age at reproduction. Population-level responses to density may be a result of interference among larvae.     

Predation drives morphological convergence in the Gambusia panuco species group among lotic and lentic habitats

Fish morphology is often constrained by a trade‐off between optimizing steady vs. unsteady swimming performance due to opposing effects of caudal peduncle size. Lotic environments tend to select for steady swimming performance, leading to smaller caudal peduncles, whereas predators tend to select for unsteady swimming performance, leading to larger caudal peduncles. However, it is unclear which aspect of performance should be optimized across heterogeneous flow and predation environments and how this heterogeneity may affect parallel phenotypic evolution. We investigated this question among four Gambusia species in north‐eastern Mexico, specifically the riverine G. panuco, the spring endemics G. alvarezi and G. hurtadoi, and a fourth species, G. marshi, found in a variety of habitats with varying predation pressure in the Cuatro Ciénegas Basin and Río Salado de Nadadores. We employed a geometric morphometric analysis to examine how body shapes of both male and female fish differ among species and habitats and with piscivore presence. We found that high‐predation and low‐predation species diverged morphologically, with G. marshi exhibiting a variable, intermediate body shape. Within G. marshi, body morphology converged in high‐predation environments regardless of flow velocity, and fish from high‐predation sites had larger relative caudal peduncle areas. However, we found that G. marshi from low‐predation environments diverged in morphology between sub‐basins of Cuatro Ciénegas, indicating other differences among these basins that merit further study. Our results suggest that a morphological trade‐off promotes parallel evolution of body shape in fishes colonizing high‐predation environments and that changing predation pressure can strongly impact morphological evolution in these species.     

<Emphasis Type="Italic">Daphnia</Emphasis> distribution in Andean Patagonian lakes: effect of low food quality and fish predation

Food quality in terms of carbon (C):phosphorus (P) ratio can constrain the success of highly demanding P herbivores as Daphnia. North Andean Patagonian lakes are ultraoligotrophic with low nutrient concentrations and well-developed euphotic zones. We investigated the distribution of the large Daphnia commutata in relation with food quality (sestonic C:P ratio) and predation risk in these lakes. The predation risk was estimated based on the fish species present and their relative eye diameter and transparency of the lake. The C:P ratios in the lakes were high, varying from 350 to >1,200. The lakes with D. commutata had significantly lower C:P ratio than those without these daphnids. On the other hand, those lakes where Daphnia is present have the lower predation risk than those were Daphnia is absent. In addition, we carried out growth experiments with neonates and natural seston of three lakes with different C:P ratio. The growth rates were inversely related with C:P of the food. Food quality and predation risk together determined the success or failure of large Daphnia populations in these Andean clear ultraoligotrophic lakes.     

Predator-induced reaction patterns of landlocked <Emphasis Type="Italic">Galaxias maculatus</Emphasis> to visual and chemical cues

Anti-predator behaviour often represents a trade-off between the benefits of reducing predation risk and the drawbacks of limiting access to resources (e.g. food availability, mating and nesting sites). The effectiveness of avoidance behaviour relies on the ability to detect predator cues, which may provide reliable information on predation risk. Using controlled laboratory experiments, we studied the relative importance of visual and chemical cues in the triggering of anti-predator responses in Galaxias maculatus, where Oncorhyncus mykiss was used as the predator. Metabolic cost was also estimated, measured as oxygen consumption. Exposure to different types of predator cue induced diverse behavioural responses in G. maculatus. Detection of the exotic predator, using both visual and chemical stimuli, resulted in reduced G. maculatus swimming activity and changes in respiratory rate.     

The predator-prey relationship of perch,Perca fluviatilis,larvae and zooplankton in two Scottish lochs

Synopsis Food consumption of perch larvae and the impact of this on zooplankton were examined in two adjacent shallow Scottish lochs. Maximum annual abundance of zooplankton occurred in mid-May at L. Kinord with minimum values in mid-June. Copepods were prominent in spring but were followed by a multi-species community of cladocerans and rotifers in summer. At L. Davan zooplankton biomass remained high through summer with cladocerans dominating andDaphnia longispina the most frequent species. Availability of food items was a principal factor governing feeding behaviour of larvae. Copepodite stages were initially the most common item in the diet in L. Kinord in 1976 and 1977 and rotifers the principal food in June 1977, reflecting the dominance of these items in the zooplankton. Cladocerans were dominant in the plankton community in L. Davan and constituted the greater part of food intake. Overlying this general pattern there was an increase in the size of food items taken by larvae with time and also a definite pattern of food selection for copepods, with initially selection for smaller copepodite stages and later for larger stages and adults. On most occasions larvae selected forCyclops strenuus abyssorum andPolphemus pediculus and selected againstDaphnia longispina. The reduction in the total zooplankton biomass attributed to perch larvae was minimal, with the exception of mid-June at L. Kinord in 1976. However, predation on particular species and copepodite stages was occasionally intense and may have impacted the zooplankton populations.     

Differences in growth of perch (Perca fluviatilis L.) in two small forest lakes

Growth patterns and food composition of perch, Perca fluviatilis L., was studied in two small forest lake populations in southern Finland. Size and morphometry of the lakes and physical and chemical properties of water are similar. There is a clear difference in the growth rates of perch between the two lakes. The difference in growth is highly significant in all age groups. In the first lake there is a perch population of 2 000 (1750 ind · ha^–1) adult fishes. In the second lake there is a small population of pike, that keeps the perch population down: 200 adult perch (530 ind · ha^-1). The main food items of perch are crustacean zooplankton, Asellus aquaticus L. and Trichoptera larvae in the first lake and zooplankton, Odonata larvae, Ephemeroptera larvae and Heteroptera in the second.It is concluded that the main reason for the growth difference of studied perch populations is the different population density. There are also differences in species composition of bottom fauna of the lakes, maybe owing to the floating Sphagnum peat moss vegetation in the second lake. This can also affect the growth difference between the two populations of perch.     

Effect of starvation on growth rate,muscle growth and energy density of puyen,Galaxias maculatus

Seasonal growth of Galaxias maculatus in Tierra del Fuego, at the southern limit of its distribution, can be explained by the variation in extrinsic factors, and experiments with fasting in summer conditions would allow researchers to focus on the potential feeding/fasting conditions independent of other seasonal cues, and thus distinguish the effect of fasting from other winter factors influencing fish growth. The study presents information on muscle growth in G. maculatus as well as the results of an experiment evaluating the effect of fasting on the growth rate, total muscle growth and energy density in this species under summer‐like conditions. A uniform pattern of muscle fibres suggests the absence of mosaic hyperplastic growth in G. maculatus. Hyperplasic growth was completed before puyen reached 65 mm total length, and hyperplasia cessation was correlated to the transition from the juvenile to the adult stage. In the experimental fish, where hypertrophic growth was the unique process responsible for adult muscle growth, fasting decreased mass, condition and energy density of the individuals, and stopped hypertrophic growth. These results are proof that reduced food consumption in southerly puyen populations during winter might contribute to the seasonality of the growth previously reported for this species. A provisional summer growth rate of 2–3 mm/month is suggested for populations inhabiting the southern limits of this species' distribution. G. maculatus is thus proposed as a useful model species to study hypertrophic muscle growth.     

Experimental analysis of the predation impact of the larvae of pond smelt (<Emphasis Type="Italic">Hypomesus transpacificus nipponensis</Emphasis>) on zooplankton populations established in mesocosms

The predation impact of the larvae of pond smelt Hypomesus transpacificus nipponensis on a zooplankton community was studied using mesocosms. The fish significantly depressed the abundances of copepod nauplii and rotifers, especially Hexarthra mira. The vulnerabilities of these prey might be determined by their swimming behavior and population density, suggesting that larval fish selectively prey on zooplankton that have a high encounter rate with the predator. The larvae did not have a negative effect on the densities of cladocerans, but fish predation altered the cladoceran community structure from the dominance of B. longirostris to that of B. fatalis. This result suggests that larval fish predation is an important factor that shifts the species composition of Bosmina in some lakes, the shift occurring in the season when fish larvae are abundant. Our results have shown that predation by the larval fish would control not only the abundance, but also the community structure of the small-sized zooplankton prey.     

Effects of perceived predation risk and social environment on the development of three‐spined stickleback (Gasterosteus aculeatus) morphology

Phenotypically plastic changes in response to variation in perceived predation risk are widespread, but little is known about if and how social environment modulates induced responses to predation risk. We investigated the influence of perceived predation risk (i.e. chemical cues from a predator) and social environment (i.e. one, two or 20 individuals reared together) on three‐spined stickleback (Gasterosteus aculeatus) morphology in a factorial common garden experiment. We found that exposure to chemical cues from potential predators did not influence growth or body condition or induce more robust morphological defences (i.e. lateral plate numbers and dorsal spine lengths). However, sticklebacks exposed to predator cues developed longer caudal peduncles and larger eyes as compared with fish from the control treatment. As these responses may improve sticklebacks’ ability to avoid piscine predation, they might be adaptive. Social environment/density also influenced expression of some traits, but these effects were independent of predation‐risk treatment effects. In general, these results suggest that apart from the classic morphological defence structures, which appear mostly constitutive, three‐spined sticklebacks are capable of expressing potentially adaptive morphological responses to chemical cues from potential predators.     

Are zooplankton food resources poor in the vegetated littoral zone of shallow lakes?

1. The distribution of zooplankton in shallow lakes is negatively related to macrophyte density. However, the abundance of their food along density gradients of macrophytes is unknown. A common but untested assumption is that food quantity and quality for pelagic zooplankton is poor in the littoral zone owing to the deleterious influence of macrophytes on phytoplankton. 2. We tested this assumption with a combination of a field survey and laboratory experiments. We collected seston samples from the littoral and pelagic zones of four shallow temperate lakes and related food quantity (phytoplankton biovolume) and quality to macrophyte abundance (per cent volume infested). Seston food quality was assessed in three ways: N/C and P/C ratios, polyunsaturated fatty acid content and phytoplankton community composition. In the laboratory, we measured the growth and reproduction of Daphnia pulicaria on diets consisting of seston from the littoral and pelagic zones in one lake. 3. In our four study lakes, food quantity was not significantly influenced by macrophyte abundance, and food quality was generally high. Laboratory experiments showed increased juvenile growth, but no significant change in D. pulicaria reproduction, when feeding on littoral resources compared to pelagic resources. 4. Our results suggest that there is no nutritional cost to pelagic zooplankton inhabiting the littoral zone. Therefore, it is likely that other factors (e.g. predation, abiotic factors) are involved in determining zooplankton habitat use.     

Seasonal Cycles of Zooplankton and Related Phytoplankton Development in Three Shallow,Mesotrophic Lakes in Northern Canada

The primary goal of this study was to assess the importance of food in regulating densities of zooplankton in 3 northern Canadian lakes, where algal availability was low compared to temperate zone water bodies. Collections were made every 2 weeks during the summer and monthly during the winter from April 1978 to April 1979. Since the lakes were similar in most respects, including nannoplankton density, net phytoplankton density, temperature, depth, oxygen concentration and phosphorus levels, the seasonal cycles of the main species (Keratella cochlearis, Kellicottia longispina, Polyarthra remata, Polyarthra vulgaris, Cyclops spp., Diaptomus spp.) were generally similar throughout the study area. Changes in the densities of herbivorous species were poorly correlated with fluctuations in nannoplankton and net plankton availability, implying that food did not limit development. Although predatory copepods, particularly Cyclops spp., were abundant, they also had no measurable impact on the main species. It was therefore concluded that temperature controlled the seasonal cycles and the ultimate population size of most zooplankters.     

Predation of perch on vendace larvae: diet composition in an oligotrophic lake and digestion time of the larvae

The diet of perch Perca fluviatilis was studied to reveal possible predation on vendace Coregonus albula larvae in an oligotrophic lake. Perch diet changed with the size of the fish: small perch ate mainly zooplankton and the diet shifted more to benthic invertebrates and fishes in larger perch. There were also annual and spatial differences in the diet, probably reflecting differences in the availability of prey animals. Perch predation on vendace larvae was only observed in the area with high availability of the larvae. The result suggested strengthened predation when the density of the larvae increases. According to bioenergetics modelling, the perch population increased natural mortality of vendace larvae only marginally. Food intake of spawning female perch was slightly reduced, whereas spawning males fed similarly to non-spawning males. Hence, the spawning period of perch was only a minor refuge for vendace larvae. Laboratory experiments of perch digestion rate demonstrated that, due to rapid digestion of the small fish larvae, diet sampling interval should not be >2 h in the field.     

Behaviour and physiology shape the growth accelerations associated with predation risk, high temperatures and southern latitudes in Ischnura damselfly larvae

1.?To better predict effects of climate change and predation risk on prey animals and ecosystems, we need studies documenting not only latitudinal patterns in growth rate but also growth plasticity to temperature and predation risk and the underlying proximate mechanisms: behaviour (food intake) and digestive physiology (growth efficiency). The mechanistic underpinnings of predator-induced growth increases remain especially poorly understood. 2.?We reared larvae from replicated northern and southern populations of the damselfly Ischnura elegans in a common garden experiment manipulating temperature and predation risk and quantified growth rate, food intake and growth efficiency. 3.?The predator-induced and temperature-induced growth accelerations were the same at both latitudes, despite considerably faster growth rates in the southern populations. While the higher growth rates in the southern populations and the high rearing temperature were driven by both an increased food intake and a higher growth efficiency, the higher growth rates under predation risk were completely driven by a higher growth efficiency, despite a lowered food intake. 4.?The emerging pattern that higher growth rates associated with latitude, temperature and predation risk were all (partly or completely) mediated by a higher growth efficiency has two major implications. First, it indicates that energy allocation trade-offs and the associated physiological costs play a major role both in shaping large-scale geographic variation in growth rates and in shaping the extent and direction of growth rate plasticity. Secondly, it suggests that the efficiency of energy transfer in aquatic food chains, where damselfly larvae are important intermediate predators, will be higher in southern populations, at higher temperatures and under predation risk. This may eventually contribute to the lengthening of food chains under these conditions and highlights that the prey identity may determine the influence of predation risk on food chain length.     

High predation is of key importance for dominance of small-bodied zooplankton in warm shallow lakes: evidence from lakes, fish exclosures and surface sediments

The mean body size of limnetic cladocerans decreases from cold temperate to tropical regions, in both the northern and the southern hemisphere. This size shift has been attributed to both direct (e.g. physiological) or indirect (especially increased predation) impacts. To provide further information on the role of predation, we compiled results from several studies of subtropical Uruguayan lakes using three different approaches: (i) field observations from two lakes with contrasting fish abundance, Lakes Rivera and Rodó, (ii) fish exclusion experiments conducted in in-lake mesocosms in three lakes, and (iii) analyses of the Daphnia egg bank in the surface sediment of eighteen lakes. When fish predation pressure was low due to fish kills in Lake Rivera, large-bodied Daphnia appeared. In contrast, small-sized cladocerans were abundant in Lake Rodó, which exhibited a typical high abundance of fish. Likewise, relatively large cladocerans (e.g. Daphnia and Simocephalus) appeared in fishless mesocosms after only 2 weeks, most likely hatched from resting egg banks stored in the surface sediment, but their abundance declined again after fish stocking. Moreover, field studies showed that 9 out of 18 Uruguayan shallow lakes had resting eggs of Daphnia in their surface sediment despite that this genus was only recorded in three of the lakes in summer water samples, indicating that Daphnia might be able to build up populations at low risk of predation. Our results show that medium and large-sized zooplankton can occur in subtropical lakes when fish predation is removed. The evidence provided here collectively confirms the hypothesis that predation, rather than high-temperature induced physiological constraints, is the key factor determining the dominance of small-sized zooplankton in warm lakes.     

Latitudinal and ecological correlates of morphological variation in Galaxias platei (Pisces, Galaxiidae) in Patagonia

We examined the morphological variability (23 morphometric traits) among individual Galaxias platei ( N  = 380) collected from 20 postglacial lakes in the southern Andes. The lakes were chosen to cover the latitudinal range of the species in Patagonia. Diet examined for a subset of these fish ( N  = 261) collected during the summer months (January through March), differed among river basins but there were no clear latitudinal trends in the consumption of any prey type. Diet may, however, have partially shaped morphology, as pelvic measures were negatively correlated with consumption of amphipods. Substantial differentiation among populations was observed, primarily in the shape and dimensions of the head, in caudal morphology and in fin length. Our results indicate that the morphology of G. platei varies with latitude, and may be related to risk of predation and diet.  © 2006 The Linnean Society of London, Biological Journal of the Linnean Society , 2006, 87 , 69–82.     

Growth of Odontesthes bonariiensis (Atherinidae) larvae feeding on suboptimal zooplankton densities

Synopsis In this study we investigate the effect of food availability (zooplankton biomass) on the growth of Odontesthes bonariensis (Atherinidae) larvae. The larvae were stocked in four 45 m² outdoor tanks at relatively high densities (100 and 200 larvae m^–2). Because of the high stocking densities, the zooplankton biomass was depleted in all tanks. However, the patterns of food limitation, and particularly periods of severe food shortage, differed in tanks stocked at different densities. We could therefore, observe the effect of food limitation in larvae that differed in weight and age. The effects of variables suspected to influence O. bonariensis growth rates (age and weight of larvae, available zooplankton biomass, mean individual weight of available preys, total ingested prey weight, and mean weight of ingested preys) were investigated using standard multiple regression methods, and a model assuming: (1) an allometric relationship between maximum growth rates and weight of larvae, and (2) an inverse relationship between growth depression and the available zooplankton biomass. Both methods were consistent in showing that only the weight of larvae, and the availability of zooplankton prey had significant effects on the growth of O. bonariensis. The model's results additionally suggest that, if the observed growth rates are scaled by the maximum growth rate corresponding to the larva weight, the effect of zooplankton biomass is largely independent of age and weight of larvae.     

Horizontal distribution of pelagic zooplankton in relation to predation gradients

An investigation was conducted examining the horizontal and vertical distribution of zooplankton in Lake Miramar, a southern California reservoir. Daphnia and Mesocyclops populations were most abundant offshore and in deeper water during the day but appeared to move toward shore and upward at night. The results of inshore zooplankton sampling provided no evidence chat the diel horizontal migration pattern was a result of sampler avoidance by zooplankton. Inshore-offshore differences in Daphnia and Mesocyclops abundance and diel migrations were reduced during winter and early spring. Rotifer zooplankters exhibited less seasonal variation in their horizontal distributions than did the large crustacean zooplankters at all times of the year. It is hypothesized that the spatial distribution of zooplankton is related to predation gradients in Lake Miramar. The dominant planktivore in the reservoir, young-of-the-year Micropterus salmoides. was abundant from late May through December and much less so from January to early May. They were largely restricted to the littoral zone and this produced horizontal gradients of planktivory which varied in strength seasonally and from day lo night. It appears that crustacean zooplankton in Lake Miramar avoid areas with abundant planktivores during the day but migrate into these areas at night when the intensity of planktivory is reduced. Rotifers exhibit less horizontal heterogeneity and no significant diel migrations, which is attributed to the reduced risk of predation that rotifers experience relative to crustacean zooplankters. A graphical model is proposed to integrate our understanding of diel vertical and horizontal migrations of zooplankton. In this model, gradients of predation are completely vertical in offshore areas and strongly horizontal in near shore areas. Gradients of food availability are roughly similar to those of predation intensity. Plankiers respond to these gradients by migrating in a path parallel to gradients of predation at dawn and parallel to gradients of food availability after dark.     

Presumptive environmental effects on body shape of Aplochiton zebra (Pisces, Galaxiidae) in northern Patagonian lakes

The family Galaxiidae exhibits a marked tendency toward morphological variation, particularly in characteristics related to feeding and caudal propulsion. The body shape of Aplochiton zebra from six Andean water bodies was examined and related to diet and to environmental characteristics such as transparency and predation risk. Although adults and larger juveniles showed no inter‐lake dependence in their morphology, smaller juveniles (SL < 40 mm) did show differences in eye diameter and dorsal fin length. Aplochiton zebra juveniles from Lake Puelo, where transparency showed the minimum value, have the largest eyes; those from Lake Futalaufquen with high transparency values have the smallest eyes. No clear relationship to predation risk was established. In Futalaufquen, Puelo and Rivadavia lakes the relationship between the principal components for morphology and for diet indicates, at least in part, that variation in body shape – and particularly traits related to swimming ability – could be related to diet.     

Clay-Turbid Interactions May Not Cascade—A Reminder for Lake Managers

Food web management is a frequently used lake restoration method, which aims to reduce phytoplankton biomass by strengthening herbivorous zooplankton through reduction of planktivorous fish. However, in clay‐turbid lakes several factors may reduce the effectivity of food web management. Increasing turbidity reduces the effectivity of fish predation and weakens the link between zooplankton and phytoplankton. Therefore, the effects of fish stock manipulations may not cascade to lower trophic levels as expected. Additionally, in clay‐turbid conditions invertebrate predators may coexist in high densities with planktivorous fish and negate the effects of fish reductions. For instance, in the stratifying regions of the clay‐turbid Lake Hiidenvesi, Chaoborus flavicans is the main regulator of cladocerans and occupies the water column throughout the day, although planktivorous Osmerus eperlanus is very abundant. The coexistence of chaoborids and fish is facilitated by a metalimnetic turbidity peak, which prevents efficient predation by fish. In the shallow parts of the lake, chaoborids are absent despite high water turbidity. We suggest that, generally, the importance of invertebrate predators in relation to vertebrate predators may change along turbidity and depth gradients. The importance of fish predation is highest in shallow waters with low turbidity. When water depth increases, the importance of fish in the top‐down regulation of zooplankton declines, whereas that of chaoborids increases, the change along the depth gradient being moderate in clear‐water lakes and steep in highly turbid lakes. Thus, especially deep clay‐turbid lakes may be problematic for implementing food web management as a restoration tool.