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PubMed2004

Induced defences in an endangered amphibian in response to an introduced snake predator.

Introduced species have contributed significantly to the extinction of endemic species on islands. They also create new selection pressures on their prey that may result in modified life history strategies. Introduced viperine snakes ( Natrix maura) have been implicated in the decline of the endemic midwife toad of Mallorca ( Alytes muletensis). A comparison of A. muletensis tadpoles in natural pools with and without snakes showed that those populations subject to snake predation possessed longer tails with narrower tail fins but deeper tail muscles. Field and laboratory experiments showed that these changes in tail morphology could be induced by chemical and tactile cues from snakes. Populations of tadpoles that were subject to snake predation also displayed clear bimodal size-frequency distributions, with intermediate-sized tadpoles missing from the pools completely. Tadpoles in pools frequented by snakes developed faster in relation to their body size than those in pools without snakes. Variation in morphology between toad populations may therefore be caused by a combination of size-selective predation and tadpole plasticity. The results of this study indicate that the introduction of alien species can result in selection for induced defences, which may facilitate coexistence between predator and prey under certain conditions.

Analysis of Variance↗

PubMed2004

Reassessment of the environmental mechanisms controlling developmental polyphenism in spadefoot toad tadpoles.

Identifying the environmental mechanism(s) controlling developmental polyphenism is the first step in gaining a mechanistic and evolutionary understanding of the factors responsible for its expression and evolution. Tadpoles of the spadefoot toad Spea multiplicata can display either a "typical" omnivorous or a carnivorous phenotype. Exogenous thyroxine and feeding on conspecific tadpoles have been accepted as triggers for development of the carnivorous phenotype on the basis of a series of studies in the early 1990s. I repeated the thyroxine and conspecific-feeding assays and demonstrated that neither exogenous thyroxine nor feeding on conspecifics induces the carnivorous phenotype. Previous researchers used simple ratio statistics to argue that field-collected carnivores and thyroxine-treated tadpoles are similar, and my results supported these claims if I used the same simple ratio methodology. However, investigation of trait developmental trajectories and allometries for field-collected carnivores and thyroxine-treated and conspecific-fed tadpoles show that these phenotypes are profoundly different.

Adaptation, Physiological↗

PubMed2004

Testing the enemies hypothesis in forest stands: the important role of tree species composition.

Numerous studies conducted in agro-ecosystems support the enemies hypothesis, which states that predators and parasites are more efficient in controlling pest densities in polycultures than in monocultures. Few similar studies, however, have been conducted in forest ecosystems, and we do not yet have evidence as to whether the enemies hypothesis holds true in forests. In a 2-year study, we investigated whether the survival of autumnal moth ( Epirrita autumnata) larvae and pupae differs between silver birch monocultures and two-species mixtures of birch with black alder, Norway spruce and Scots pine. We placed young larvae on birch saplings and monitored their survival until the end of the larval period, when we checked whether they had been parasitized. After the larvae had pupated, pupal survival was tested in a field trial. In 2002, the larvae disappeared earlier and their overall survival was lower in birch-pine mixtures than in other stand types. In 2003, survival probability was lowest in birch-pine stands only during the first week and there were no differences between stands in overall survival. Larval parasitism was not affected by tree species composition. Pupal weight and pupal survival were likewise not affected by stand type. Among the predators, wood ants were more abundant on birches growing in birch-pine mixtures than in other stand types probably because colonies of myrmecophilic aphids were common on pines. In contrast, spider numbers did not differ between stand types. Ant exclusion by means of a glue ring around the birch trunk increased larval survival, indicating that ants are important predators of the autumnal moth larvae; differences in larval survival between stands are probably due to differential ant predation. Our results provide only partial support for the enemies hypothesis, and suggest that it is both tree species composition and species diversity which affect herbivore survival and predation.

Analysis of Variance↗

PubMed2004

Predator-induced transgenerational phenotypic plasticity in the cotton aphid.

Transgenerational phenotypic changes, whereby offspring have an altered trait or a distinct alternate phenotype, frequently occur in response to increased maternal predation risk. The cotton aphid, Aphis gossypii (Glover), is unique, however, as offspring consist of four distinct phenotypes ("normal" light green apterae, "normal" dark green apterae, "dwarf" yellow apterae, and alatae), all with divergent life history patterns and resulting population dynamics. Here, we show that increased predation risk induces transgenerational phenotypic changes in cotton aphids. When exposed to search tracks from larval or adult convergent ladybird beetles, Hippodamia convergens Guerin-Meneville, cotton aphids produced greater numbers of winged offspring. In a subsequent experiment, apterous and alate individuals on clean plants were found to have primarily normal and dwarf offspring, respectively. We suggest that elevated predation risk may cause phenotypic changes in aphids over multiple generations, resulting in a more precipitous decline in herbivore populations than could be explained solely by increased predation rates.

Analysis of Variance↗

PubMed2004

Predation by sparrowhawks decreases with increased breeding density in a songbird, the great tit.

Predators may regulate prey populations if predation rate increases with prey density. Alternatively, if space-limited (e.g. territorial) predators become 'satiated' when prey exceed a certain density, increased prey abundance may lead to reduced predation rate. These alternatives have been difficult to test experimentally for mobile prey in the wild. We present such a test, manipulating the density of great tits (Parus major) by adding nest boxes in territories of sparrowhawks (Accipiter nisus). Predation rate was measured for young tits after they left the nests. Although the great tit is an important prey, there was no evidence for regulation during the breeding season: the rate of hawk predation declined with increasing density of tits. This result was not confounded by changes in breeding density of alternative prey species (other songbirds). Hawk predation can therefore favour dense breeding in a territorial (solitary) bird, and conspecific attraction and aggregation reported in several territorial species may partly result from predation pressure. This result also has potential implications for conservation work.

Animals↗

PubMed2004

What limits herb biomass in grasslands: competition or herbivory?

Competition and herbivory are two of the main forces shaping plant communities. Although several studies have investigated their impact on plant populations separately, few investigations have examined how they might interact. With the purpose of clarifying the combined roles of competition and herbivory on herb biomass in a grassland, we assessed the effects of different herbivores with reduced grass competition. We conducted a field experiment in 2000-2001 in a British acid grassland (Oak Mead), where we experimentally reduced grass biomass and excluded rabbits, insects and molluscs in a factorial design. Removing the grasses from Oak Mead dramatically increased herb biomass and total above-ground biomass. Herbivore exclusions (i.e. rabbits, insects and molluscs) did not affect total above-ground biomass, but they altered relative abundance of several species. Grass removal and rabbit exclusion had positive interactions on biomass of several herb species, and there were some subtle interactions between different herbivore groups: monocots benefitted when both rabbits and molluscs were excluded, and some herb species had greater biomass when insects and rabbits were absent. We then compared the results with a 10-year experiment that manipulated similar variables in neighbouring grassland (Nash's Field). The comparison between Oak Mead and Nash's Field showed that cessation of herbicide application returns the system to its previous state of grass dominance after 3 years. Therefore, even when herbs were more abundant, they could not prevent reinvasion of the grasses once external factors were removed.

Analysis of Variance↗

PubMed2004

Limitations on reproductive success in endemic Aquilegia viscosa (Ranunculaceae) relative to its widespread congener Aquilegia vulgaris: the interplay of herbivory and pollination.

Plant reproduction can be strongly affected by herbivory and different features of pollination ecology, such as pollinator visitation rates and capacity for self-pollination. The purpose of this study is to compare the relative impact of herbivory and pollination on maternal reproductive success in endemic Aquilegia viscosa and its widespread congener Aquilegia vulgaris. We conducted herbivore exclusion experiments in two populations of each species in 2 different years and showed that the maternal fertility of A. viscosa was significantly more limited by floral predation and pre-dispersal seed predation than its widespread congener. In the absence of herbivory, A. viscosa retained significantly lower maternal fertility than A. vulgaris. Experimental pollinations in an insect-free glasshouse showed that the two species have an equal seed/ovule ratio both in the absence of pollinators and in the presence of non-limiting outcross pollination. Pollinator visitation rates were significantly higher in populations of A. vulgaris than in populations of A. viscosa. In addition, path analyses showed that spur length, an important trait for pollinator attraction in Aquilegia, and, indirectly sepal and petal width, contribute positively to the seed/ovule ratio in A. vulgaris, but not in A. viscosa. These results indicate that maternal fertility of endemic A. viscosa is strongly reduced by flower and seed predation despite low rates of pollinator visitation, and that pollen or resource limitation in the wild may further reduce maternal fertility. Finally, floral trait variation appears to be decoupled from fertility variation in endemic A. viscosa, which possibly constrains the evolution of reproductive traits in this species.

Analysis of Variance↗

PubMed2004

The persistence of ripe fleshy fruits in the presence and absence of frugivores.

A trade-off between antimicrobial defences and palatability to dispersers may place limits on fruit persistence in nature. The retention times of ripe fruits on 34 wild plant species under natural conditions (unbagged persistence) and when fruits had been bagged with nylon mesh to exclude frugivores (bagged persistence) were compared in Hong Kong, China (22 degrees N). Bagged persistence is a measure of the effectiveness of fruit defence while unbagged persistence is an inverse measure of attractiveness to vertebrate frugivores. Bagged fruits persisted significantly longer than unbagged fruits in 30 species, with half the species tested persisting for more than 2 months. There was a significant positive relationship between the median persistence times of bagged and unbagged fruits, suggesting that species with a high resistance to microbial infection are also less attractive to frugivores. Both bagged and unbagged fruits persisted significantly longer at lower temperatures. There was a significant positive relationship between bagged persistence time and fibre content of the fruit pulp, but no significant relationships between unbagged persistence and the six fruit traits tested (diameter, pulp as a percentage of fruit fresh weight, and lipid, total soluble carbohydrate, nitrogen and fibre as percentages of pulp dry weight). Mechanical damage significantly decreased the bagged persistence time for half of the species. Although some fruits decayed or dried up while attached to the plant, fruits of 53% of the species remained visually attractive until they fell off.

Animals↗

PubMed2004

Spatial avoidance of littoral and pelagic invertebrate predators by Daphnia.

Studies on spatial avoidance behaviour of predators by prey often ignored the fact that prey typically face multiple predators which themselves interact and show a spatial pattern in abundance and predation rates (PRs). In a series of laboratory experiments, we investigated predation risk (PRI) and horizontal migration of the cladoceran Daphnia magna between open water and vegetation in response to two important invertebrate predators with a contrasting spatial distribution: pelagic Choaborus and vegetation-associated Ischnura. As expected, PRI by Chaoborus was higher in open water due to higher numbers and higher PRs of Chaoborus, while for Ischnura, PRI was highest in the vegetation due to higher densities, despite lower PRs of Ischnura. In accordance with this, Daphnia moved into the vegetation in the presence of the pelagic Chaoborus alone. In the presence of Ischnura alone, however, Daphnia showed no response. We hypothesize this may be the result of a constitutive behaviour of Daphnia to avoid pelagic fish, which impedes a response to the open water. In the combined predator treatment, Daphnia migrated to the open water zone. The increased risk of predation in the vegetation, due to a facilitating effect of Chaoborus on Ischnura PRs is believed to have caused this migration of the Daphnia. This response of Daphnia declined through time and Daphnia moved toward the vegetation. A decline in the activity of the Ischnura larvae through time may have switched the risk balance in favour of the vegetation environment.

Animals↗

PubMed2004

Lichen palatability depends on investments in herbivore defence.

Lichens are well-suited organisms for experimental herbivory studies because their secondary compounds, assumed to deter grazing, can be non-destructively extracted. Thalli of 17 lichen species from various habitats were cut in two equal parts; compounds were extracted from one part by acetone, the other served as a control. These two pieces were offered as a paired choice to the generalist herbivore snail Cepaea hortensis. Control thalli of all lichens were consumed at a low rate regardless of their investments in acetone-extractable lichen compounds; naturally compound-deficient lichen species were not preferred compared to those with high contents. However, for extracted thalli, there was a highly significant positive correlation between rate of consumption and the extracted compound contents. These data imply that herbivore defence has evolved in different directions in different lichens. Studied members of Parmeliaceae, common in oligotrophic habitats, have high contents of carbon-rich acetone-soluble compounds; these lichens became highly palatable to snails subsequent to acetone rinsing. Extracted lichen compounds were applied to pieces of filter paper and fed to snails. Extracts from members of the Parmeliaceae significantly deterred feeding on paper. Such data suggest that generalist herbivores may have shaped evolution in the widespread and highly diverse Parmeliaceae towards high investments in lichen compounds. On the other hand, lichens belonging to the Physciaceae and Teloschistales, common in nutrient-enriched habitats, are deficient in, or have low concentrations of, lichen compounds. Such lichens did not become more palatable after acetone rinsing. The orange anthraquinone compound parietin, restricted to the Teloschistales, and which has previously been found to protect against excess light, did not deter grazing.

Acetone↗

PubMed2004

The direct and indirect effects of insectivory by birds in two contrasting Neotropical forests.

A goal among community ecologists is to predict when and where trophic cascades occur. For example, several studies have shown that forest birds can limit arthropod abundances on trees, but indirect effects of bird predation (i.e. decreased arthropod damage to trees) are not always observed and their context is not well understood. Because productivity is one factor that is expected to influence trophic cascades, we compared the extent to which birds indirectly limit herbivore damage to trees in two lowland Neotropical forests that differed in seasonality of leaf production and rainfall. We compared the effects of bird predation on local arthropod densities and on damage to foliage through a controlled experiment using bird exclosures in the canopy and understory of two forests. We found that birds decreased local arthropod densities and leaf damage in the canopy of the drier site during periods of high leaf production, but not in the wetter forest where leaf production was low and sporadic throughout the year. Birds had no effect on arthropod abundances and leaf damage in the understory where leaf production and turnover rates were low. In support of these experimental interpretations, although we observed that arthropod densities were similar at the two sites, bird densities and the rate at which birds captured arthropods were greater at the drier, seasonally productive site. The influence of top-down predation by birds in limiting herbivorous insects appears to be conditional and most important when the production and turnover of leaves are comparatively high.

Animals↗

PubMed2004

Herbivore attack in Casearia nitida influenced by plant ontogenetic variation in foliage quality and plant architecture.

Traits influencing plant quality as food and/or shelter for herbivores may change during plant ontogeny, and as a consequence, influence the amount of herbivory that plants receive as they develop. In this study, differences in herbivore density and herbivory were evaluated for two ontogenetic stages of the tropical tree Casearia nitida. To assess plant ontogenetic differences in foliage quality as food for herbivores, nutritional and defensive traits were evaluated in saplings and reproductive trees. Predatory arthropods were quantified and the foraging preferences of a parasitoid wasp of the genus Zacremnops were assessed. In addition, survival rates of lepidopteran herbivores (Geometridae) were evaluated experimentally. Herbivore density was three times higher and herbivory was 66% greater in saplings than in reproductive trees. Accordingly, concentrations of total foliar phenolics were higher in reproductive trees than in saplings, whereas leaf toughness, water and nitrogen concentration did not vary between ontogenetic stages. Survival rates of lepidopteran larvae exposed to natural enemies were equivalent in reproductive trees and saplings. Given the greater herbivore density on saplings, equal survival rates implied a greater foraging effort of predators on reproductive trees. Furthermore, observed foraging of parasitoid wasps was restricted to reproductive trees. I propose that herbivore density, and as a consequence, leaf damage were lower in reproductive trees than in saplings due to both traits influencing food quality, and architectural or unmeasured indirect defensive traits influencing foraging preference of natural enemies of herbivores.

Animals↗

PubMed2004

A new approach to the solution of the linear mixing model for a single isotope: application to the case of an opportunistic predator.

Mixing models are used to determine diets where the number of prey items are greater than one, however, the limitation of the linear mixing method is the lack of a unique solution when the number of potential sources is greater than the number (n) of isotopic signatures +1. Using the IsoSource program all possible combinations of each source contribution (0-100%) in preselected small increments can be examined and a range of values produced for each sample analysed. We propose the use of a Moore Penrose (M-P) pseudoinverse, which involves the inverse of a 2x2 matrix. This is easily generalized to the case of a single isotope with (p) prey sources and produces a specific solution. The Antarctic leopard seal (Hydrurga leptonyx) was used as a model species to test this method. This seal is an opportunistic predator, which preys on a wide range of species including seals, penguins, fish and krill. The M-P method was used to determine the contribution to diet from each of the four prey types based on blood and fur samples collected over three consecutive austral summers. The advantage of the M-P method was the production of a vector of fractions f for each predator isotopic value, allowing us to identify the relative variation in dietary proportions. Comparison of the calculated fractions from this method with 'means' from IsoSource allowed confidence in the new approach for the case of a single isotope, N.

Animals↗

PubMed2005

Stable isotopes show food web changes after invasion by the predatory cladoceran Cercopagis pengoi in a Baltic Sea bay.

Cercopagis pengoi, a recent invader to the Baltic Sea and the Laurentian Great Lakes, is a potential competitor with fish for zooplankton prey. We used stable C and N isotope ratios to elucidate trophic relationships between C. pengoi, zooplankton (microzooplankton, 90-200 microm, mostly copepod nauplii and rotifers; mesozooplankton, >200 microm, mostly copepods), and zooplanktivorous fish (herring, size range 5-15 cm and sprat, 9-11 cm) in a coastal area of the northern Baltic Sea. The isotope ratios in C. pengoi and fish were much higher than those of zooplankton, showing general trends of enrichment with trophic level. Young-of-the-year (YOY) herring had a significantly higher (15)N/(14)N ratio than C. pengoi, suggesting of a trophic linkage between the two species. To evaluate the possible relative importance of different food sources for C. pengoi and YOY herring, two-source isotope-mixing models for N were used, with micro- and mesozooplankton as prey for C. pengoi and mesozooplankton and C. pengoi as prey for YOY herring. These models indicate that mesozooplankton was the major food source of both species. However, microzooplankton may be important prey for young stages of C. pengoi. Comparative analyses of the herring trophic position before and after the invasion by C. pengoi showed a trophic level shift from 2.6 to 3.4, indicating substantial alterations in the food web structure. Our findings contribute to a growing body of evidence, showing that C. pengoi can modify food webs and trophic interactions in invaded ecosystems.

Animals↗

PubMed2005

Relative importance of fertiliser addition to plants and exclusion of predators for aphid growth in the field.

Herbivore dynamics and community structure are influenced both by plant quality and the actions of natural enemies. A factorial experiment manipulating both higher and lower trophic levels was designed to explore the determinants of colony growth of the aphid Aphis jacobaeae, a specialist herbivore on ragwort Senecio jacobaea. Potential plant quality was manipulated by regular addition of NPK-fertiliser and predator pressure was reduced by interception traps; the experiment was carried out at two sites. The size and persistence of aphid colonies were measured. Fertiliser addition affected plant growth in only one site, but never had a measurable effect on aphid colony growth. In both habitats the action of insect predators dominated, imposing strong and negative effects on aphid colony performance. Ants were left unmanipulated in both sites and their performance on the aphid colonies did not significantly differ between sites or between treatments. Our results suggest that, at least for aphid herbivores on S. jacobaea, the action of generalist insect predators appears to be the dominant factor affecting colony performance and can under certain conditions even improve plant productivity.

Analysis of Variance↗

PubMed2005

Sequential predator effects across three life stages of the African tree frog, Hyperolius spinigularis.

While theoretical studies of the timing of key switch points in complex life cycles such as hatching and metamorphosis have stressed the importance of considering multiple stages, most empirical work has focused on a single life stage. However, the relationship between the fitness components of different life stages may be complex. Ontogenetic switch points such as hatching and metamorphosis do not represent new beginnings--carryover effects across stages can arise when environmental effects on the density and/or traits of early ontogenetic stages subsequently alter mortality or growth in later stages. In this study, I examine the effects of egg- and larval-stage predators on larval performance, size at metamorphosis, and post-metamorphic predation in the African tree frog Hyperolius spinigularis. I monitored the density and survival of arboreal H. spinigularis clutches in the field to estimate how much egg-stage predation reduced the input of tadpoles into the pond. I then conducted experiments to determine: (1) how reductions in initial larval density due to egg predators affect larval survival and mass and age at metamorphosis in the presence and absence of aquatic larval predators, dragonfly larvae, and (2) how differences in mass or age at metamorphosis arising from predation in the embryonic and larval environments affect encounters with post-metamorphic predators, fishing spiders. Reduction in larval densities due to egg predation tended to increase per capita larval survival, decrease larval duration and increase mass at metamorphosis. Larval predators decreased larval survival and had density-dependent effects on larval duration and mass at metamorphosis. The combined effects of embryonic and larval-stage predators increased mass at metamorphosis of survivors by 91%. Larger mass at metamorphosis may have immediate fitness benefits, as larger metamorphs had higher survival in encounters with fishing spiders. Thus, the effects of predators early in ontogeny can alter predation risk even two life stages later.

Age Factors↗

PubMed2005

Predation regulation of sedimentary faunal structure: potential effects of a fishery-induced switch in predators in a Newfoundland sub-Arctic fjord.

The collapse of the cod fishery in Newfoundland has coincided with marked increases in abundances of snow crab, pandalid shrimp, and other crustaceans that prey on sedimentary infauna. A 3-year sampling program in Bonne Bay, Newfoundland indicates differences in composition and number of these predators in the two main arms of the fjord that coincide with strong differences in benthic community structure. To test whether predation pressure contributes to the observed patterns in sedimentary fauna, exclusion field experiments with full and partial cages were deployed in both arms at 30-m depth and sampled along with ambient sediments at 0-, 4-, and 8-week periods. Predation significantly influenced species composition, abundance and, in some cases, diversity. The most striking changes included increases in the polychaetes Phöloe tecta and Ophelina cylindricaudata in exclusions relative to controls, and concurrent declines in the polychaete Paradoneis lyra and the cumacean Lamphros fuscata. In laboratory experiments, fresh non-disturbed sediment cores from each experimental area were either protected or exposed to snow crab, the most abundant predator in the bay. A snow crab inclusion experiment was also carried out in the field, using cages similar to those used for exclusions. Despite differences in sedimentary faunas in the two arms, both types of experiments detected a predator effect that was very similar to that documented in exclusion experiments. Thus, despite differences in the scales associated with each type of manipulation, our results suggest that crab predation is a significant structuring force in Newfoundland sedimentary communities. Given the historical changes that have occurred in predator composition as a result of cod over-fishing, we hypothesize that broad-scale community changes may be taking place in North Atlantic benthic ecosystems.

Analysis of Variance↗

PubMed2005

Indirect cost of a defensive trait: variation in trichome type affects the natural enemies of herbivorous insects on Datura wrightii.

The costs and benefits of defensive traits in plants can have an ecological component that arises from the effect of defenses on the natural enemies of herbivores. We tested if glandular trichomes in Datura wrightii, a trait that confers resistance to several species of herbivorous insects, impose an ecological cost by decreasing rates of predation by the natural enemies of herbivores. For two common herbivores of D. wrightii, Lema daturaphila and Tupiocoris notatus, several generalized species of natural enemies exhibited lower rates of predation on glandular compared to non-glandular plants. Lower rates of predation were associated with reductions in the residence time and foraging efficiency of natural enemies on plants with glandular trichomes, but not with direct toxic effects of glandular exudate. Our results suggest that the benefit of resistance to herbivores conferred by glandular trichomes might be offset by the detrimental effect of this trait on the natural enemies of herbivores, and that the fitness consequences of this trichome defense might depend on the composition and abundance of the natural-enemy community.

Analysis of Variance↗