Ontogenetic habitat shifts by Galaxias gracilis (Galaxiidae) between the littoral and limnetic zones of Lake Kanono, New Zealand

Synopsis Diel and spatial differences in distribution were determined for the larvae, juveniles, and adults of Galaxias gracilis (Galaxiidae) in a New Zealand dune lake during summer months. Larvae (mostly 10–25 mm TL) and juveniles (25–40 mm TL) inhabited shallow (0–3 m) waters of the limnetic zone and fed predominantly on two limnetic zooplankton species; Bosmina meridionalis and a calanoid copepod. At about 40 mm TL, fish moved from the limnetic to the littoral zone and expanded dietary breadth from two to over seven main prey species, including five species of littoral invertebrates. After reaching a size of about 60 mm TL, most fish moved back offshore to the deeper waters (5–15 m) of the limnetic zone during the day, moving back to the littoral zone at night to feed on invertebrates. The selection of different intra-lacustrine habitats by the various size groups of G. gracilis, and the movements between them, are interpreted as adaptive responses to the interaction between ontogenetic changes in feeding requirements and predation risk.     

Distribution of perch (Perca fluviatilis L.) during their first year of life in Lake Constance

The distribution and behaviour of larval and juvenile perch (Perca fluviatilis L.) were studied for two years in large, deep Lake Constance. After hatching larvae were transported by water currents to the open water. The majority of larvae remained in the pelagic zone for about one month. In both years, their return to the littoral zone coincided with the decline of pelagic zooplankton abundance. After returning to the littoral zone, juveniles stayed among submerged macrophytes within 5 m depth and lived apart from larger perch which lived at depths of 6–20 m. By late summer, juveniles changed their distribution pattern: during the day they stayed intensively close to piers and ports, but increased their swimming activity at dusk, cruising among shallow and deep waters and feeding on zooplankton, and rested on the bottom at night. This behaviour appears to be related to the decrease of inshore food resources and to the presence of predators in deeper water. 0⁺ perch left the littoral zone and moved into deep waters when autumnal mixing began in late October. They overwintered near the bottom at depths of more than 30 m. During most of the year, juvenile and adult perch were separated from each other. But as soon as they occupied the same habitat, the occurrence of cannibalism increased.     

The ecology of ruffe,Gymnocephalus cernuus (Pisces: Percidae) introduced to Mildevatn,western Norway

Synopsis Diet, habitat use, diel and seasonal activity and a number of population parameters were studied on ruffe,Gymnocephalus cernuus, introduced to Mildevatn, western Norway. This lake is sited outside the natural range of the ruffe and has a lower fish diversity and a different fish species composition than within its native range. From June through September the ruffe was planktivorous and mainly caught at 4 to 6 m depth in the benthic zone. At other times of year ruffe was feeding on zoobenthos and caught deeper in the benthic zone. Ruffe was mainly day active. Zooplankton feeding during summer is the clearest difference compared to ruffe populations living within its natural range. Presence of large zooplankton organisms available for ruffe is suggested as the main reason for the difference found in food choice. The availability of large zooplankton is probably due to community structure caused by a predator and lack of interspecific competition for zooplankton in the deeper parts of the lake. Piscivorous brown trout.Salmo trutta, restrict the habitat of threespined stickleback,Gasterosteus aculeatus, to the zone of littoral vegetation, allowing high densities of larger zooplankton species likeBythotrephes longimanus to be present in the lake. Brown trout is present only in the upper light and well oxygenated parts of the lake, leaving a refuge for the ruffe, where they can feed on the rich zooplankton community.     

Diel horizontal migration and swarm formation in Daphnia in response to Chaoborus

Dense swarms of Daphnia longispina (up to 4000 animals l^–1) were recorded along the littoral zone in a lake where Chaoborus flavicans is considered the main predator. D. longispina coexisted with D. pulex, but there were no D. pulex in the littoral swarms. Swarms were less dense at night (about 1/10 the density), and D. longispina exhibited diel horizontal migrations by aggregating in the littoral during the day and spreading out at night. Laboratory experiments showed that Chaoborus capture efficiency on juvenile daphnids was higher in the light compared to darkness, and that Daphnia exhibited a behavioural response to water that had previously contained Chaoborus. We conclude that predation from Chaoborus can be an important factor affecting the distribution patterns of Daphnia observed in this lake. The behavioural experiments indicated that this influence might be partly mediated by chemical agents.     

Combined effects of fish and macroinvertebrate predation on zooplankton in a littoral mesocosm experiment

Whether macrophytes offer an effective refuge for zooplankton in all shallow lakes is subject to debate. To explore potential constraints between different predator threats and the related habitat choice by zooplankton, we conducted a mesocosm experiment in 12 large-sized pools mimicking the nearshore environment with part of its length being covered by submersed macrophytes (Egeria densa) and holding a mixed zooplankton community. Four treatments were used: (i) young zooplanktivorous fish (3 silverside, Odontesthes bonariensis) in the “open-water” zone; (ii) macroinvertebrate predator (31 grass shrimp, Palaemonetes argentinus) in the vegetated zone; (iii) both, fish in the open-water and shrimp in the vegetated zones; and (iv) control with no predators. Our results show specific effects of each predator on the abundance, composition, and size of cladocerans. Regarding distribution, in control and shrimp mesocosms, no differences were found between the two zones, while cladocerans were clearly more abundant in the vegetated side in the presence of fish. When both fish and shrimp were present, cladocerans preferred the vegetated zone too, but in a smaller proportion, and their abundance was less. The presence of predatory macroinvertebrates in vegetated littoral zone reduces the refuge value of this habitat, at least for cladocerans.

     

Effects of ship‐induced waves on littoral benthic invertebrates

1. Ship‐induced waves can affect the physical characteristics of lake and river shorelines, and laboratory studies have shown effects on littoral invertebrates. Here, we explored whether these effects could be observed under field conditions along a natural lake shore affected by wave sequences (trains) produced by boats. 2. Individuals of five invertebrate species (Bithynia tentaculata, Calopteryx splendens, Dikerogammarus villosus, Gammarus roeselii, Laccophilus hyalinus) were exposed to waves with increasing shear stress in five habitats differing in structural complexity. 3. Detachment of invertebrates increased with increasing shear stress and was best modelled using sigmoid response curves. Habitat structural complexity mitigated the effects of shear stress, and detachment rate was influenced more by habitat type than by species. A threshold (90% of the individual invertebrates unaffected) stress level of 0.64 N m^?2 was found for a structurally complex reed habitat, compared to 0.37 N m^?2 for a simple sand habitat. 4. Shear stress associated with wave trains created by recreational boating at a distance of 35 m from the shore and at a speed of 11 km h^?1 resulted in 45% detachment of littoral invertebrates. Decreasing the boat‐to‐shore distance to 20 m increased wave shear stress by 30% and invertebrate detachments up to 75%. 5. Disturbance of littoral habitats and invertebrate assemblages are widespread in inland waters used for recreational and/or commercial navigation. Our findings show that the integrity of littoral zones of navigable surface waters could be much improved by implementing management measures such as physically protecting complex habitats with dense reed belts and tree roots, and reducing boat speeds and increasing their minimum shoreline distance.     

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

Synopsis Galaxias 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.     

Movement of plankton through lake-stream systems

1. River plankton are often assumed to come from upstream lakes, but the factors controlling the movement of plankton between lakes and rivers into outflow streams are unclear. We tested the possibility that the physical structure of the littoral zone near the lake outlet (depth, presence of macrophytes) and diurnal differences in plankton composition at the lake surface influence the movement of plankton from the lake into the stream and determine their persistence downstream. 2. Zooplankton and phytoplankton biomass, community composition and mean body size were compared between two deep lakes without macrophytes at the lake edge and two shallow lakes with macrophytes at the lake edge. Samples were collected day and night on three dates, in the lake centre, in the littoral zone adjacent to the lake outlet, at the outlet and at two sites downstream in Algonquin Park, Ontario, Canada. 3. The morphology of lake edges clearly affects the movement of lake zooplankton into outlet streams. Outlets draining deeper littoral zones had higher zooplankton biomass than shallow littoral outlets (P < 0.0001), but these differences disappeared within 50 m downstream of the lake. There was no difference in mean zooplankton body size among lake outlets or between littoral and outlet samples. However, shallow littoral zones were dominated by cyclopoid copepods and deeper littoral zones were dominated by Bosmina longirostris. In contrast, phytoplankton biomass entering the outlet was similar to that found within the lake and did not vary with lake outlet morphology. These effects were consistent across several sampling weeks and were not affected by surface zooplankton biomass changes associated with diurnal vertical migration in the lake centre. 4. A comparison with published river zooplankton data suggests that zooplankton are rapidly eliminated from shallow outlet streams (≤1 m deep) but persist in most deeper outlet rivers (≥2 m deep). Because the depth of an outlet river determines downstream zooplankton community development, the contribution of lakes to river plankton communities may be influenced by the location of each lake within the drainage basin. These findings suggest that lake and outflow physical structure influences connection strength between spatially successive habitats.     

Lake morphometry predicts the degree of habitat coupling by a mobile predator

Habitat coupling is an ecosystem process whereby semi-discontinuous habitats are connected through the movement of energy and nutrients by chemical, physical or biological processes. One oft-cited example is that of littoral–pelagic coupling in lakes. Theory has argued that such habitat coupling may be critical to food web dynamics, yet there have been few empirical studies that have quantified ecological factors that affect the degree of habitat coupling in ecosystems. Specifically, the degree to which habitat coupling occurs across important physical gradients has largely been ignored. To address this, we investigate the degree of littoral habitat coupling (i.e. the degree to which a top predator lake trout, Salvelinus namaycush, derives energy from the littoral zone) along a gradient of lake shape, where lake shape modifies the relative quantity of coupled epilimnetic benthic and pelagic habitats within each lake. Herein we demonstrate that littoral habitat coupling is intensified in simple circular lakes compared to their reticulate counterparts in seven Canadian Shield lakes. Although the more reticulate lakes had larger areas of epilimnetic benthic habitat, littoral food sources comprised 11% compared to 24% of lake trout diet in reticulate and circular lakes, respectively. This heightened interaction in circular lakes also appears to translate into increased omnivory in more circular lakes compared to reticulate lakes such that lake trout of circular lakes have a significantly lower trophic position than lake trout of reticulate lakes (F_1,5=6.71 p=0.05). These results suggest that it is the accessibility of littoral production via thermal refugia, and not the amount of littoral production, that determines the degree to which lake trout couple littoral and pelagic habitats in lakes.     

Seasonal habitat selection by lake trout (<Emphasis Type="Italic">Salvelinus namaycush</Emphasis>) in a small Canadian shield lake: constraints imposed by winter conditions

The need for cold, well-oxygenated waters significantly reduces the habitat available for lake trout (Salvelinus namaycush) during stratification of small temperate lakes. We examined the spatial and pelagic distribution of lake trout over two consecutive summers and winters and tested whether winter increased habitat availability and access to littoral regions in a boreal shield lake in which pelagic prey fish are absent. In winter, lake trout had a narrowly defined pelagic distribution that was skewed to the upper 3 m of the water column and spatially situated in the central region of the lake. Individual core areas of use (50% Kernel utilization distributions) in winter were much reduced (75%) and spatially non-overlapping compared to summer areas, but activity levels were similar between seasons. Winter habitat selection is in contrast to observations from the stratified season, when lake trout were consistently located in much deeper waters (>6 m) and widely distributed throughout the lake. Winter distribution of lake trout appeared to be strongly influenced by ambient light levels; snow depth and day length accounted for up to 69% of the variation in daily median fish depth. More restricted habitat use during winter than summer was in contrast to our original prediction and illustrates that a different suite of factors influence lake trout distribution between these seasons.     

Invertebrate zooplankton predator composition and diversity in tropical lentic waters

Invertebrate zooplankton predators are generally less diverse in average species numbers in tropical than in temperate lakes and reservoirs. Predatory Copepoda which comprise the majority of limnetic predators are particularly low in species numbers in the tropics. Predatory Cladocera are confined to the North Temperate zone. Chaoborus appears to be cosmopolitan. Among Rotifera, only the cosmopolitan predator Asplanchna occurs in tropical waters while the other common limnetic carnivorous genus Ploesoma is restricted to higher latitudes. Hydracarina, and insects besides Chaoborus, are generally restricted to the littoral and appear to be more diverse in the tropics. Lakes Awasa and Zwai, Ethiopia, are almost devoid of predators in the limnetic, which is invaded by a littoral chydorid Alona diaphana. Low diversity of lake types and low production of tropical zooplankton could restrict predator diversity too. Very low diversity of invertebrate predators in the limnetic and higher diversity in the littoral may characterize tropical lakes in contrast to temperate lakes, which have more invertebrate predators in the limnetic and perhaps relatively less in the littoral. Tropical zooplankton in freshwaters, appears to be a very immature community. Hence opportunistic species can readily invade the limnetic and even dominate in isolated situations as has been shown for Alona davidi, Hydracarina and some other unconventional forms.     

The role of the littoral zone in the functioning of a shallow tropical lake ecosystem

Lake Chilwa (Malawi) is a shallow tropical lake surrounded by an extensive zone of littoral swamp mainly composed of Typha domingensis. The data produced show that the physical and chemical conditions are different and more varied in the littoral region when compared with the open lake. Primary production in the littoral is confined almost entirely to Typha domingensis, which also acts as a nutrient pump by absorbing nutrients from the soils and eventually releasing them into the lake waters. Long term and short term exchanges in organic and inorganic materials between the littoral swamps and the open lake are described. These, as well as available evidence on the distribution, reproductive habits and diet of certain faunal groups, indicate that the littoral region of Lake Chilwa has an important regulating influence on the functioning of the whole lake.     

Comparison of Day and Night Fyke Netting, Electrofishing and Snorkelling for Monitoring a Population of the Threatened Golden Galaxias (Galaxias auratus)

The littoral zone of small off-stream water storage containing a translocated population of Galaxias auratus was sampled fortnightly at day and night with fyke nets, electrofishing and snorkelling over 3 months. Variation in population data provided by each method, including relative abundance indices, size structure, and habitat preferences, were examined. Aspects of behaviour and activity patterns were also investigated. Night sampling using all methods consistently yielded larger catches than day sampling. The size structure of catches varied, with electrofishing at night and fyke netting during the day having higher proportions of juveniles, whilst snorkelling at night and electrofishing during the day had higher proportions of adults. Fyke netting at night yielded by far the largest catches (~3-fold more than other methods) and also captured balanced proportions of juveniles and adults. Galaxias auratus had a strong diel activity pattern and were most active at night. The majority of the population migrated into the littoral zone during the night and back into deeper water during the day. A small number of juveniles remained in the littoral zone and some adults sheltered in the dense cover of species-rich littoral vegetation during the day. Shores with shallow depth profiles appeared to be preferred due to higher catches in these areas using all methods. Based on the results of this study, fyke netting at night in littoral habitats is recommended for monitoring populations of G. auratus. Fyke netting is likely to be an effective method for monitoring other lacustrine galaxiid species; however, further work is required to investigate the effects of habitat variables and fish community structure on activity patterns of galaxiids, and hence their catchability with various methods, in more extensive lentic environments.     

Maximum entropy model: Estimating the relative suitability of cetacean habitat in the northern Savu Sea,Indonesia

An understanding of cetacean distribution is necessary to gain insights into crucial ecological processes for species conservation management. However, cetacean habitat preference and distribution in the northern Savu Sea, Indonesia, are still poorly understood. We use maximum entropy modeling with five environmental predictors to describe habitat preference and distribution of seven cetacean species in the northern Savu Sea. Our study confirms that static predictors are the most important variables in explaining habitat preferences of seven cetacean species in the northern Savu Sea. Seasonally, each cetacean species has a different environmental preference. Globicephala macrorhynchus prefers the open nearshore areas adjacent to deep waters. Stenella attenuata is associated with nearshore habitats and higher productivity areas. Stenella longirostris prefers nearshore habitat during coastal upwelling events. Tursiops truncatus prefers nearshore sheltered habitat. Grampus griseus habitats are highly related to depth and steep gradients around the isobaths of 200–1,000 m. Feresa attenuata are associated with deep waters and cooler sea temperature. Pseudorca crassidens prefers protective bays with productive waters. This study provides important information for reviewing marine spatial plans of the northern Savu Sea, specifically on managing species fisheries interaction, optimizing surveillance, and regulating marine traffic.     

Littoral habitat loss caused by multiyear drought and the response of an endemic fish species in a deep desert lake

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Long‐term impacts of invasive species on a native top predator in a large lake system

1. Declining abundances of forage fish and the introduction and establishment of non‐indigenous species have the potential to substantially alter resource and habitat exploitation by top predators in large lakes. 2. We measured stable isotopes of carbon (δ¹³C) and nitrogen (δ¹⁵N) in field‐collected and archived samples of Lake Ontario lake trout (Salvelinus namaycush) and five species of prey fish and compared current trophic relationships of this top predator with historical samples. 3. Relationships between δ¹⁵N and lake trout age were temporally consistent throughout Lake Ontario and confirmed the role of lake trout as a top predator in this food web. However, δ¹³C values for age classes of lake trout collected in 2008 ranged from 1.0 to 3.9‰ higher than those reported for the population sampled in 1992. 4. Isotope mixing models predicted that these changes in resource assimilation were owing to the replacement of rainbow smelt (Osmerus mordax) by round goby (Neogobius melanostomus) in lake trout diet and increased reliance on carbon resources derived from nearshore production. This contrasts with the historical situation in Lake Ontario where δ¹³C values of the lake trout population were dominated by a reliance on offshore carbon production. 5. These results indicate a reduced capacity of the Lake Ontario offshore food web to support the energetic requirements of lake trout and that this top predator has become increasingly reliant on prey resources that are derived from nearshore carbon pathways.     

Effect of vertebrate predation on the spatio-temporal distribution of cladocerans in a temperature eutrophic lake

We analysed the spatio-temporal distribution of zooplankton along a profile of 10 stations from the shore to the pelagic zone from April to September 1988, the period when the larvae and juveniles Rutilus rutilus, the most abundant species in the Lake, are in the littoral zone. The digestive tracts of the young roach were analysed. They fed essentially on rotifers and on cladocerans. For comparison, zooplankton was also analysed at one littoral area without fish fry. There was an increase of cladoceran density from the vegetated nearshore zone to the offshore zone. Considering the density of Bosmina longirostris, Daphnia longispina, Chydorus sphaericus and Ceriodaphnia quadrangula, we observed a different distribution pattern in the course of the year. In the nearshore zone, the relative abundance of small species, Bosmina and Chydorus, was much higher than that of the larger Daphnia. From April to September, predation pressure mainly affected the smallest species: in contrast to the inshore station without fish fry, the density of Bosmina decreased in May in the littoral with fish. Chydorus was concentrated in the littoral between February and April, then grew into the pelagic zone, where predation pressure obviously was low during the warm season. The number of Daphnia, which was eaten by the fish fry at any time, remained low in the nearshore zone, which suggests that the presence of fish may cause Daphnia to avoid this zone. Ceriodaphnia which was not affected by this predation, was scarce in the nearshore zone during mid-summer. The low density of the cladocerans in the nearshore zone is likely associated with vertebrate predation by roach fry and juveniles, the result of such a process being either a depletion in density of the prey, or an avoidance behaviour.     

Environmental gradients and the horizontal distribution of microcrustaceans in lakes

1. The assemblage of suspension‐feeding microcrustaceans in lakes changes along a habitat gradient from nearshore to offshore. This gradient of microcrustaceans was explored in relation to differences in macrophytes and the associated changes in water chemistry, food resources and types of predators.
2. Some microcrustacean species were littoral or limnetic specialists, while others changed their distribution along this horizontal habitat gradient on a diel or seasonal basis. Distribution patterns were similar in a lake and a pond which differed in extent of macrophyte habitat.
3. There was a large shift in the composition of sestonic food, indicating heterotrophic seston nearshore and more autotrophic seston offshore. Sit‐and‐wait predators of microcrustaceans (e.g. Enallagma spp.) dominated nearshore and cruising predators (e.g. Leptodora kindtii Focke) were more common offshore.
4. A reciprocal transplant experiment revealed that littoral specialists could survive equally well when fed littoral or limnetic seston, while limnetic specialists performed poorly when fed littoral seston. Food resources may be important in determining where some microcrustacean species live along this horizontal habitat gradient.     

The benthic macroinvertebrate community in a coastal sand dune lake relative to habitat and changing lake levels

The benthic macroinvertebrate community (BMI) in a freshwater coastal dune lake without a surface outlet was investigated in May and October, 1986. Fifty-three invertebrate taxa were identified from Carter Lake, including three euryhaline crustacean species (Corophium spinicorne, Gnorimosphaeroma oregonensis lutea, and Acanthomysis awatchensis). Corophium spinicorne dominated the BMI communities of the littoral zones and sphaeriid clams dominated the deepwater community.The lake level dropped about 2.5 m between April and October. Based upon this decline, the lake bottom was divided into four major habitats: a sandy temporarily submerged littoral zone (A); a sandy submerged littoral zone (B); and mid-depth zone of mixed mud and sand and the macrophyte, Nitella (C); and a deep zone (D) with soft mud. The average density of BMI was highest in the littoral zones (A and B) in May and in zone B in October (zone A was dry). The lowest density occurred in zone D. In May, BMI biomass was highest in the littoral zones, but the biomass was highest in the mid-depth zone in October. The mid-depth zone in October. The mid-depth zone had the most diverse community.The two most abundant species in the temporarily submerged area, Corophium spinicorne and Juga plicifera, were found in greater numbers deeper in the lake after the water level dropped, suggesting migration by these species in response to changing water levels.     

Habitat use of arctic charrSalvelinus alpinus in Thingvallavatn,Iceland

Synopsis Habitat use by four morphs of arctic charr,Salvelinus alpinus, was investigated in Thingvallavatn, Iceland, by sampling with pelagic and benthic gill nets. Sampling was done in May/June and August/September. Greatest abundance of fish was recorded in the littoral and epipelagic zone in early autumn. Catches were low in early summer. The four morphs are partly segregated in habitat. Small (SB-) and large benthivorous (LB-) charr have a more restricted spatial distribution than piscivorous (PI-), and especially planktivorous (PL-) charr. Both benthivorous morphs are mainly found in the littoral zone, and occur in largest numbers in stony shallows at depths between 0 and 10 m. PL-charr usually dominates in numbers in all habitats. PI-charr is most abundant in epibenthic habitats, although numbers are always low. All morphs are caught in higher numbers at night than during the day, but the diurnal activity difference is highest among SB-charr. The habitat use by different morphs is as may be expected from their morphology and diets. Within the population of PL-charr, young and small fish are more abundant on the bottom than in the pelagic zone, and there is a surplus of females in the pelagic zone. Along the benthic profile, young, small and immature PL-charr are more abundant in deep than in shallow waters. The results are discussed in relation to food supply, competition and predation. Possible reasons for the occurrence of four arctic charr morphs are also discussed.Contribution from the Thingvallavatn project.