Biomanipulation: the next phase — making it stable

Review of the literature on biomanipulation shows that fish manipulations have occurred through the following methods: piscivore addition; piscivore catch restriction; habitat enhancement; piscivore removal; planktivore exclusion; planktivore removal by selective catch, lake emptying, fish poisons, fish diseases, winterkill, summerkill; habitat expansion or contraction; planktivore addition; and natural events. The methods can be classified as deliberate, inadvertent, or natural, all of which have successful and unsuccessful examples. However, the problem of perpetuation of successful results remains unsolved. It is proposed that this may be resolved through use of a refuge or refuges from among the following: low light intensity refuge; low temperature refuge; low dissolved oxygen refuge; physical concealment refuge; visual clutter refuge; behavior modification refuge; and predator inefficiency refuge. Perhaps through use of such mechanisms large herbivorous zooplankters can continue to exist in lakes. Examples are given. Contribution no. 383 from the Limnological Research center.     

Time allocation of a parasitoid foraging in heterogeneous vegetation: implications for host-parasitoid interactions

1. Changing plant composition in a community can have profound consequences for herbivore and parasitoid population dynamics. To understand such effects, studies are needed that unravel the underlying behavioural decisions determining the responses of parasitoids to complex habitats. 2. The searching behaviour of the parasitoid Diadegma semiclausum was followed in environments with different plant species composition. In the middle of these environments, two Brassica oleracea plants infested by the host Plutella xylostella were placed. The control set-up contained B. oleracea plants only. In the more complex set-ups, B. oleracea plants were interspersed by either Sinapis alba or Hordeum vulgare. 3. Parasitoids did not find the first host-infested plant with the same speed in the different environments. Sinapis alba plants were preferentially searched by parasitoids, resulting in fewer initial host encounters, possibly creating a dynamic enemy-free space for the host on adjacent B. oleracea plants. In set-ups with H. vulgare, also, fewer initial host encounters were found, but in this case plant structure was more likely than infochemicals to interfere with the searching behaviour of parasitoids. 4. On discovering a host-infested plant, parasitoids located the second host-infested plant with equal speed, demonstrating the effect of experience on time allocation. Further encounters with host-infested plants that had already been visited decreased residence times and increased the tendency to leave the environment. 5. Due to the intensive search of S. alba plants, hosts were encountered at lower rates here than in the other set-ups. However, because parasitoids left the set-up with S. alba last, the same number of hosts were encountered as in the other treatments. 6. Plant composition of a community influences the distribution of parasitoid attacks via its effects on arrival and leaving tendencies. Foraging experiences can reduce or increase the importance of enemy-free space for hosts on less attractive plants.     

Refuge use in sticklebacks as a function of body length and group size

A change in anti–predator strategy from hiding to grouping outside a refuge was observed in large three–spined sticklebacks. No such change in strategy was seen in small fish. The body–length dependence of this strategy change is discussed in the context of metabolic constraints.     

Plant structural complexity and mechanical defenses mediate predator–prey interactions in an odonate–bird system

Habitat‐forming species provide refuges for a variety of associating species; these refuges may mediate interactions between species differently depending on the functional traits of the habitat‐forming species. We investigated refuge provisioning by plants with different functional traits for dragonfly and damselfly (Odonata: Anisoptera and Zygoptera) nymphs emerging from water bodies to molt into their adult stage. During this period, nymphs experience high levels of predation by birds. On the shores of a small pond, plants with mechanical defenses (e.g., thorns and prickles) and high structural complexity had higher abundances of odonate exuviae than nearby plants which lacked mechanical defenses and exhibited low structural complexity. To disentangle the relative effects of these two potentially important functional traits on nymph emergence‐site preference and survival, we conducted two fully crossed factorial field experiments using artificial plants. Nymphs showed a strong preference for artificial plants with high structural complexity and to a lesser extent, mechanical defenses. Both functional traits increased nymph survival but through different mechanisms. We suggest that future investigations attempt to experimentally separate the elements contributing to structural complexity to elucidate the mechanistic underpinnings of refuge provisioning.     

Backwater habitats and their role in nature conservation on navigable waterways

In Britain, lightly trafficked canals frequently contain diverse, productive macrophyte communities. These represent important habitats for macroinvertebrates and fish while having a high intrinsic value in nature conservation terms. As recreational boat traffic increases, fragile macrophytes are progressively eliminated and the biomass of the remaining species is greatly reduced, thereby adversely affecting weed-associated animals and ultimately simplifying the structure of the whole ecosystem. From the viewpoint of aesthetics, nature conservation and fisheries management,ecological enhancement of these traffic impacted ecosystems is desirable but options are limited by channel size and the intensity and type of disturbance. Backwater areas connected to the main channel but apparently remote from traffic influences ought however, to provide a minimally-disturbed refuge for macrophytes and dependent organisms. An extensive field survey was undertaken to test this hypothesis and evaluate the potential for exploiting backwater sites as off-line nature reserves. Principal determinants of vegetation structure and species diversity are identified and discussed and are used to prescribe a set of ideal characteristics for prospective backwater nature reserves and to forecast likely management problems.     

The effect of sand and light on predation of juvenile plaice (Pleuronectes platessa) by fishes and crustaceans

Rates of predation on 0-group plaice, Pleuronectes platessa . in aquaria were compared under four different combinations of conditions to test the hypothesis that the presence of sand in which they may bury affords a refuge from predators. The effect of light and darkness on predation rate was also examined, Two crustaceans, the shrimp, Crangon crangon , and the portunid crab, Liocarcinus holatus , and two fishes, cod, Gadus morhua , and pollack, Pollachius pollachius , were used as predators. Predaton rates were significantly higher in the dark for all predators except pollack. Predation rates in the absence of sand were signifcantly greater only for pollack. The results suggest that predation rates on plaice during their juvenile nursery stage on sandy beaches will be significantly greater during darkness than during the day. Burying in sand appears to provide only a partial refuge from predation, perhaps because natural predators have evolved effective methods of foraging for buried prey.     

Kin and familiarity influence association preferences and aggression in the mangrove killifish Kryptolebias marmoratus

Association preferences and aggression intensity were investigated in relation to kin and familiarity in the self-fertilizing, clonal vertebrate, the mangrove killifish Kryptolebias marmoratus. Results indicated that fish preferentially associated with and exhibited less intense aggression towards members of their own genotype (kin), compared to members of a different genotype (non-kin). Furthermore, when fish were presented with stimulus groups of the same genotype that were familiar or unfamiliar, fish preferentially associated with and exhibited lower aggression intensity directed towards familiar groups. These results indicate that this species prefer to associate with both kin and familiar individuals and modulate aggression accordingly. These results are discussed with reference to the adaptive benefits of kin recognition and preferences for familiars, and place results within the context of current knowledge of the ecology of K. marmoratus.     

The consequence of a food refuge collapse on a guild of lake-dwelling triclads and leeches

The diets ofPolycelis nigra, P. tenuis, Dugesia polychroa, Dendrocoelum lacteum, Glossiphonia complanata, Helobdella stagnalis andErpobdella octoculata in an English lake were examined, using a serological technique, and compared between 1981–82 and 1989–90. Leech, triclad and prey abundances were also recorded. Between the two studies, snail numbers crashed whereas the abundances ofAsellus andGammarus increased.Dugesia andGlossiphonia numbers decreased substantially, whilstPolycelis tenuis andHelobdella abundances increased. In the second study, the snail component in the diet was greatly reduced resulting in a broader food niche, particularly forDugesia andGlossiphonia, and greater food overlap between the predators with the exception ofDendrocoelum andErpobdella which do not eat molluscs. It is postulated that the reduced size of the snail refuge, and consequent increase in severity of interspecific competition with other predators, particularlyPolycelis andHelobdella, led to the observed decrease in abundances ofDugesia andGlossiphonia. The decline in the last two genera, perhaps coupled with increased crustacean abundance, could have contributed to the numerical increase of the competitively superiorPolycelis tenuis andHelobdella.     

Late Pleistocene divergence and postglacial expansion in the Brazilian Atlantic Forest: multilocus phylogeography of Rhopias gularis (Aves: Passeriformes)

In the last decade, phylogeographic studies have revealed a complex evolutionary history of the Brazilian Atlantic Forest (AF) biota. Here, we investigated the evolutionary history of Rhopias gularis, an endemic bird of the AF, based on sequences of two mitochondrial genes and three nuclear introns from 64 specimens from 15 localities. We addressed three main questions: (1) Does the genetic diversity of R. gularis exhibit a distribution pattern congruent with the refuge hypothesis postulated for the AF? (2) Is the population genetic structure of R. gularis congruent with those observed in other AF species? (3) What were the possible historical events responsible for the population structure of this species? Our mtDNA data revealed two phylogroups: (1) phylogroup central‐south, with samples from the central and southern parts of the range; (2) and phylogroup north, which included individuals from southern Bahia. Nevertheless, nuclear loci did not reveal any evidence of population structure. Bottleneck tests indicated that the central‐south lineage experienced demographic expansion, starting around 20 kya, which coincides with the end of the last glacial maximum. However, there was no evidence of population growth in phylogroup north. Isolation with migration analysis indicated that these phylogroups split c.a. 304 kya, with limited gene flow among them. Palaeodistribution models indicated that R. gularis had a reduced distribution in the south and central AF during the last glacial maximum. Our results support a diversification scenario that is in accordance with proposed Pleistocene refugia. The phylogeographic results from our study exhibited spatial and temporal concordances and discordances with previous studies of organisms from the AF. Differences in habitat requirements of these species could be behind this complex scenario. Future studies correlating variables of the niche of these species with the observed phylogeographic patterns may help understand why there are congruent and incongruent results.     

Oyster Reef Restoration in the Northern Gulf of Mexico: Effect of Artificial Substrate and Age on Nekton and Benthic Macroinvertebrate Assemblage Use

In the northern Gulf of Mexico (GOM), reefs built by eastern oysters, Crassostrea virginica, provide critical habitat within shallow estuaries, and recent efforts have focused on restoring reefs to benefit nekton and benthic macroinvertebrates. We compared nekton and benthic macroinvertebrate assemblages at historic, newly created (<5 years) and old (>6 years) shell and rock substrate reefs. Using crab traps, gill‐nets, otter trawls, cast nets, and benthic macroinvertebrate collectors, 20 shallow reefs (<5 m) in the northern GOM were sampled throughout the summer of 2011. We compared nekton and benthic assemblage abundance, diversity and composition across reef types. Except for benthic macroinvertebrate abundance, which was significantly higher on old rock reefs as compared to historic reefs, all reefs were similar to historic reefs, suggesting created reefs provide similar support of nekton and benthic assemblages as historic reefs. To determine refuge value of oyster structure for benthic macroinvertebrates compared to bare bottom, we tested preferences of juvenile crabs across depth and refuge complexity in the presence and absence of adult blue crabs (Callinectes sapidus). Juveniles were more likely to use deep water with predators present only when provided oyster structure. Provision of structural material to support and sustain development of benthic and mobile reef communities may be the most important factor in determining reef value to these assemblages, with biophysical characteristics related to reef location influencing assemblage patterns in areas with structure; if so, appropriately locating created reefs is critical.     

Using mitochondrial DNA to test the hypothesis of a European post-glacial human recolonization from the Franco-Cantabrian refuge

It has been proposed that the distribution patterns and coalescence ages found in Europeans for mitochondrial DNA (mtDNA) haplogroups V, H1 and H3 are the result of a post-glacial expansion from a Franco-Cantabrian refuge that recolonized central and northern areas. In contrast, in this refined mtDNA study of the Cantabrian Cornice that contributes 413 partial and 9 complete new mtDNA sequences, including a large Basque sample and a sample of Asturians, no experimental evidence was found to support the human refuge-expansion theory. In fact, all measures of gene diversity point to the Cantabrian Cornice in general and the Basques in particular, as less polymorphic for V, H1 and H3 than other southern regions in Iberia or in Central Europe. Genetic distances show the Cantabrian Cornice is a very heterogeneous region with significant local differences. The analysis of several minor subhaplogroups, based on complete sequences, also suggests different focal expansions over a local and peninsular range that did not affect continental Europe. Furthermore, all detected clinal trends show stronger longitudinal than latitudinal profiles. In Northern Iberia, it seems that the highest diversity values for some haplogroups with Mesolithic coalescence ages are centred on the Mediterranean side, including Catalonia and South-eastern France.     

Impact of urbanization and gardening practices on common butterfly communities in France

We investigated the interacting impacts of urban landscape and gardening practices on the species richness and total abundance of communities of common butterfly communities across France, using data from a nationwide monitoring scheme. We show that urbanization has a strong negative impact on butterfly richness and abundance but that at a local scale, such impact could be mitigated by gardening practices favoring nectar offer. We found few interactions among these landscape and local scale effects, indicating that butterfly‐friendly gardening practices are efficient whatever the level of surrounding urbanization. We further highlight that species being the most negatively affected by urbanization are the most sensitive to gardening practices: Garden management can thus partly counterbalance the deleterious effect of urbanization for butterfly communities. This holds a strong message for park managers and private gardeners, as gardens may act as potential refuge for butterflies when the overall landscape is largely unsuitable.     

Training fish for restocking: refuge and predator training in the hatchery has limited benefits for a marine fish

Large numbers of hatchery-reared fish are released in stocking programmes; however, success is limited by high mortality. Predation is seen as the main cause of deaths but might be reduced by training fish before release to avoid predators and/or use refuge. In this study on a potential restocking species, yellowfin bream Acanthopagrus australis, the effects of predator training and refuge on the behaviour of fish in the hatchery were tested. In the first experiment, juvenile bream were exposed to predatory mangrove jack (Lutjanus argentimaculatus) fed exclusively on bream flesh while housed in tanks with and without refuge. Predator training altered fish behaviour when fish were re-exposed to predators, but the effects were subtle and varied between groups of fish. In contrast, refuge created strong and consistent changes in behaviour, significantly slowing down the amount of time that fish took to consume food. A second experiment focused on the effects of refuge. Bream were trained to use artificial seagrass or house bricks as refuge and then exposed to mangrove jacks in a laboratory predation experiment. When refuge was available, fish significantly slowed down their feeding rate. There was a small, transient increase in survival for fish given seagrass refuges, but this was irrespective of whether the bream were trained to use refuge. The results of this study indicate that the use of refuge may be innate and the benefits of refuge may be available to naive hatchery-reared fish or fish trained to use refuge shortly before release. This suggests that there is potential to improve post-release survival of fingerlings without time-consuming and expensive hatchery training.     

Microclimatic buffering and resource‐based habitat in a glacial relict butterfly: significance for conservation under climate change

In contrast to several organisms that have already shown range shifts to the north as a response to climate change, southern populations of relict species are trapped in isolated altitudinal habitats. Therefore, there is a growing interest to better understand their habitat use, with particular attention to the thermal aspects and associated significance for their habitat management. We address this issue by a study of larval habitat use relative to vegetation structure and microclimate in a glacial relict butterfly of peat bog ecosystems, using a functional, resource‐based habitat approach. We analysed caterpillar presence and density relative to vegetation composition (reflecting gradients of humidity, temperature, and natural succession of the peat bog) and to the availability and quality of thermal refuges for caterpillars (i.e., structures provided by Sphagnum hummocks). We also tested caterpillar survival rates under different temperature and humidity treatments. We found that (1) Boloria aquilonaris was a specialist butterfly of early successional stages with very humid zones of peat bog, (2) the lack of Sphagnum hummocks reduced larval habitat suitability, and hence the population density, and (3) a reduction of the thermal buffering ability of Sphagnum hummocks was observed in less humid, degraded parts, or late‐successional stages of peat bog. A larval rearing experiment showed a significant impact of temperature on caterpillar survival; survival being higher at lower temperature. Our field and laboratory results support the idea that the thermal environment exploited by caterpillars should be considered as a functional resource and included in a population‐specific habitat definition. Appropriate management of the peat bog habitat of this glacial relict species should not exclusively focus on the larval and adult feeding resources, but also on the quality of thermal refuges provided by Sphagnum hummocks in humid zones of the peat bog, especially in the current critical context of climate warming.     

Ultimate and proximate mechanisms underlying the occurrence of bears close to human settlements: review and management implications

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Declining populations in one of the last refuges for threatened mammal species in northern Australia

Australia has contributed a disproportionate number of the world's mammal extinctions over the past 200 years, with the greatest loss of species occurring through the continent's southern and central arid regions. Many taxonomically and ecologically similar species are now undergoing widespread decline across the northern Australian mainland, possibly driven by predation by feral cats and changed fire regimes. Here, we report marked recent declines of native mammal species in one of Australia's few remaining areas that support an intact mammal assemblage, Melville Island, the largest island off the northern Australian coast. We have previously reported a marked decline on Melville Island of the threatened brush‐tailed rabbit‐rat (Conilurus penicillatus) over the period 2000–2015, linked to predation by feral cats. We now report a 62% reduction in small mammal trap‐success and a 36% reduction in site‐level species richness over this period. There was a decrease in trap‐success of 90% for the northern brown bandicoot (Isoodon macrourus), 64% for the brush‐tailed rabbit‐rat and 63% for the black‐footed tree‐rat (Mesembriomys gouldii), but no decline for the common brushtail possum (Trichosurus vulpecula). These results suggest that populations of native mammals on Melville Island are exhibiting similar patterns of decline to those recorded in Kakadu National Park two decades earlier, and across the northern Australian mainland more generally. Without the implementation of effective management actions, these species are likely to be lost from one of their last remaining strongholds, threatening to increase Australia's already disproportionate contribution to global mammal extinctions.     

Abundance, microhabitat selection and conservation of eyed lizards (Lacerta lepida): a radiotelemetric study

The utility of radiotelemetry as a tool for estimating the size and microhabitat requirements of a population of Lacerta lepida , the largest European lacertid, was investigated in central Spain. Population density estimates based on repeated marking and recapture (3.2 lizards ha⁻¹) were much higher than those based on line transects (0.22 lizards ha⁻¹). The probability of sighting lizards before they could retreat into a refuge was largely increased by our ability to radiolocate them. Rocks were selected as refuges 96% of the times, and the locations of radiotracked lizards were much closer to rocks than randomly expected. Rocks used as retreat sites were larger and had more crevices than those available at random, which suggests that refuge selection was primarily determined by the need to find shelter from predators. Rockrose patches, which were positively selected, might be used as refuge-connecting corridors that combine shelter with opportunities to forage and thermoregulate. Our results emphasize the need for using radiotelemetry to establish baseline information on abundance and to clarify the actuality, extent and pattern of the population declines experienced by species that may function as key links in their ecosystems, but the wariness of which poses a serious problem for monitoring their conservation status.     

Does Stream Morphology Predict the Home Range Size in Burbot?

Synopsis To determine whether burbot occupy defined home range in rivers, we radio-tracked individuals in the Ohře River, Czech Republic. We also tested the hypothesis that the size of burbot home range would correlate with the fish mass. Burbot's strong attraction to suitable refuges was the basis for our second hypotheses, that its diurnal behavior would reflect refuge availability in the riverine environment. To test these hypotheses, we analyzed data on fish movements in relation to depth, velocity, substratum size and river slope. During the night, burbot preferred deeper areas with lower slope and finer substrates than during daylight hours. The home range was smaller in areas with low or zero slopes, and significantly increased with increasing river slope. There was no relationship between home range size and fish mass. River slope appeared to be the main predictor of the burbot's home range size.     

Predation and refugia: implications for Chaoborus abundance and species composition

1. Previous studies have suggested that the occurrence of larval Chaoborus in lakes may be affected by fish predation, pH, elevation, temperature, nutrient level, water transparency and interspecific competition, but so far, a detailed statistical evaluation of these findings has not been performed. 2. The aim of this study was to apply regression and ordination techniques to a large data set of 56 lakes in order to test which variables related to lake morphology, water chemistry, and fish predation determine (1) the abundance of individual Chaoborus species and (2) their species composition. 3. Individual Chaoborus species were influenced by very different sets of environmental factors. Nutrient levels positively affected the largest species, Chaoborus americanus, which was restricted to fishless lakes. Abundance of the smallest and most transparent species, C. punctipennis, seemed to be controlled more by the larger Chaoborus species than by fish. Larger chaoborids required low water clarity in order to co‐exist with fish, probably to increase refuge availability. Generally, small lakes (for C. flavicans/C. trivittatus) and shallow lakes (for C. punctipennis) supported higher abundances of Chaoborus.     

The potential of artificial refugia for maintaining a community of large-bodied cladocera against fish predation in a shallow eutrophic lake

The Norfolk Broads are a series of shallow lakes which are highly eutrophic and typified by dense populations of phytoplankton and an absence of submerged aquatic plants. The zooplankton community is subject to intense predation pressure by young fish and is dominated by small-bodied organisms which have a low potential for reducing phytoplankton populations through grazing. Various designs and densities of artificial refugia for zooplankton against fish predation were established in Hoveton Great Broad in order to enhance populations of large-bodied Cladocera. Initially some of the refuges contained higher densities and larger individuals ofDaphnia andCeriodaphnia than the surrounding open water. However, towards the end of the first season and throughout the subsequent two years, population densities and size-structure were similar both within and outside the refuges, although there was still evidence of enhanced body-size ofDaphnia within the refuges compared with the open water. The provision of habitat structures designed as refugia from fish predation did not enhance large-bodied cladoceran populations enough to promote this restoration strategy as feasible for eutrophic and shallow lakes.