Measuring rodent incisors from scats can increase accuracy of predator diet studies. An illustration based on island cats and rats

Non-invasive diet studies, which are a simple but important tool to understand trophic interactions inside ecosystems, need to be as detailed as possible. Determining the precise biomass of ingested prey is a key to obtaining not only a better understanding of the amount of food really ingested but also the predator-prey interactions. It is particularly relevant in the case of rodents, because they are often a predominant prey in carnivores' diet and can differ widely in biomass. This study demonstrates how an original and simple method for measuring rat incisors found in cat scats produces measurements which can be correlated with rat weight. This correlation, used in a field application, made it possible to: (i) calculate a more accurate biomass of rats in cat diets and thus obtain a better estimation of the proportion of rats compared to other prey in cat diets; (ii) show that cats preferentially ate smaller rats, indicating that the use of the mean weight of rodents sampled by trap-lines may induce a significant bias in the biomass calculation. Likewise, a correlation between rat lens weight and incisor measurements was found. Using this correlation, it should be possible to estimate the age of the rats eaten by cats and obtain a better understanding of the impact of predators on prey population dynamics. To cite this article: E. Bonnaud et al., C. R. Biologies 331 (2008).     

Hidden costs to an invasive intraguild predator from chemically defended native prey

Studies of invasive species tend to concentrate on the reasons for their success, rather than on the potential costs that they face. Invasive predators are potentially vulnerable to chemical defences of native prey that they have not previously encountered, and evolved to tolerate, within their natural range. While novel chemical defences have been cited as a factor facilitating invasion by exotic prey, the potential costs of chemical novelty to invasive predators have been ignored or remained hidden. We here consider one such example. Although a dominant intraguild predator in interactions with native ladybird beetles, the invasive ladybird Harmonia axyridis finds the defensive alkaloids of certain native ladybirds either distasteful or toxic, although laboratory and field evidence suggests it readily attacks these prey; these natives appear to possess novel alkaloids not previously encountered by H. axyridis . This effect has hitherto remained hidden, due to the methodological approaches used in earlier studies.     

Low functional richness and redundancy of a predator assemblage in native forest fragments of Chiloe island, Chile

1. Changes in land use and habitat fragmentation are major drivers of global change, and studying their effects on biodiversity constitutes a major research programme. However, biodiversity is a multifaceted concept, with a functional component linking species richness to ecosystem function. Currently, the interaction between functional and taxonomic components of biodiversity under realistic scenarios of habitat degradation is poorly understood. 2. The expected functional richness (FR)-species richness relationship (FRSR) is positive, and attenuated for functional redundancy in species-rich assemblages. Further, environmental filters are expected to flatten that association by sorting species with similar traits. Thus, analysing FRSR can inform about the response of biodiversity to environmental gradients and habitat fragmentation, and its expected functional consequences. 3. Top predators affect ecosystem functioning through prey consumption and are particularly vulnerable to changes in land use and habitat fragmentation, being good indicators of ecosystem health and suitable models for assessing the effects of habitat fragmentation on their FR. 4. Thus, this study analyses the functional redundancy of a vertebrate predator assemblage at temperate forest fragments in a rural landscape of Chiloe island (Chile), testing the existence of environmental filters by contrasting an empirically derived FRSR against those predicted from null models, and testing the association between biodiversity components and the structure of forest fragments. 5. Overall, contrasts against null models indicate that regional factors determine low levels of FR and redundancy for the vertebrate predator assemblage studied, while recorded linear FRSR indicates proportional responses of the two biodiversity components to the structure of forest fragments. Further, most species were positively associated with either fragment size or shape complexity, which are highly correlated. This, and the absence of ecological filters at the single-fragment scale, rendered taxonomically and functionally richer predator assemblages at large complex-shaped fragments. 6. These results predict strong effects of deforestation on both components of biodiversity, potentially affecting the functioning of remnants of native temperate forest ecosystems. Thus, the present study assesses general responses of functional and taxonomic components of biodiversity to a specific human-driven process.     

Reproduction in an endemic bird of a semiarid island: a food-mediated process

We analyse the effects of rainfall, temperature, food availability and nest predation on the between-year variation in reproduction of the Canary Islands stonechat Saxicola dacotiae , a bird species endemic of the semiarid island of Fuerteventura (Canary Islands, Spain). We monitored the reproductive performance and output of the species across its whole distribution area during three consecutive breeding periods, also measuring rainfall, temperature and food (arthropod) availability on the ground monthly. Rainfall varied from 27.3 mm in 2000–2001 (dry year) to 124.5 mm and 125.1 mm (average years), respectively, in the 2001–2002 and 2002–2003 breeding seasons. The onset of breeding closely matched variation in the onset of the autumn-winter rains among years. Arthropod availability was strongly and positively related to rainfall one month before. Reproductive investment (number of clutches and clutch size) was correlated to arthropod availability both among years and among sites within years. Stonechat pairs bred once or did not breed at all in the dry year whereas they bred twice in the other two years. Clutch size was smaller in the dry year and larger in the second as compared with the first in the other two years. Reproductive investment largely determined reproductive output (number of fledglings) as there were no significant spatial or temporal variation in hatching success (90% on average) or nest predation (29% on average, mostly due to feral cats Felis catus ). Within- and among years variability in temperature or predation did not match variability in the onset, length or reproductive investment and output recorded throughout the study years. These results suggest that variability in annual fecundity in the Canary Islands Stonechat was mainly driven by rainfall through a food-mediated process.     

Extremely reduced dispersal and gene flow in an island bird

The Réunion grey white-eye, Zosterops borbonicus, a passerine bird endemic to Réunion Island in the Mascarene archipelago, represents an extreme case of microgeographical plumage colour variation in birds, with four distinct colour forms occupying different parts of this small island (2512 km²). To understand whether such population differentiation may reflect low levels of dispersal and gene flow at a very small spatial scale, we examined population structure and gene flow by analysing variation at 11 microsatellite loci among four geographically close localities (<26 km apart) sampled within the distribution range of one of the colour forms, the brown-headed brown form. Our results revealed levels of genetic differentiation that are exceptionally high for birds at such a small spatial scale. This strong population structure appears to reflect low levels of historical and contemporary gene flow among populations, unless very close geographically (<10 km). Thus, we suggest that the Réunion grey white-eye shows an extremely reduced propensity to disperse, which is likely to be related to behavioural processes.     

Passive recovery of an island bird community after rodent eradication

The number and scale of island invasive species eradications is growing, but quantitative evidence of the conservation efficacy of passive recovery is limited. We compare relative abundances of breeding birds on Hawadax Island (formerly named Rat island), Aleutian Archipelago, Alaska, pre- and post- rat eradication to examine short-term (<1 year post-eradication) changes due to rodenticide application, and medium-term (5 years post-eradication) changes due to the absence of invasive rats. In the short term, Bald Eagle (Haliaeetus leucocephalus) numbers decreased from 24 individuals pre-eradication to two individuals <1 year post-eradication, but recovered to 10 individuals (42 % of pre-eradication) 5 years post-eradication, with all individuals nesting (63 % of the pre-eradication nesting). Five years post-eradication relative abundances of most terrestrial birds surveyed using point counts either significantly increased [Gray-crowned Rosy Finch (Leucosticte tephrocotis), Lapland Longspur (Calcarius lapponicus), Snow Bunting (Plectrophenax nivalis), Song Sparrow (Melospiza melodia)] or did not differ [Pacific Wren (Troglodytes troglodytes)]. Shorebirds also increased 5 years post-eradication with Black Oystercatchers (Haematopus palliates) increasing fivefold, and Rock Sandpiper (Calidris ptilocnemis) nesting increasing from one to five nests. We confirmed two species of ground nesting seabirds [Tufted Puffin (Fratercula cirrhata) and Leach’s Storm-petrel (Oceanodroma leucohoa)] as nesting (puffin) or engaged in courtship behavior (Storm-petrel) 5 years post-eradication. Our results indicate that despite the short-term impact on Bald Eagles, and without further human intervention, most terrestrial and marine birds have newly-colonized, re-colonized, or increased in abundance following the eradication of invasive rats.     

Crying wolf to a predator: deceptive vocal mimicry by a bird protecting young

Animals often mimic dangerous or toxic species to deter predators; however, mimicry of such species may not always be possible and mimicry of benign species seems unlikely to confer anti-predator benefits. We reveal a system in which a bird mimics the alarm calls of harmless species to fool a predator 40 times its size and protect its offspring against attack. Our experiments revealed that brown thornbills (Acanthiza pusilla) mimic a chorus of other species'' aerial alarm calls, a cue of an Accipiter hawk in flight, when predators attack their nest. The absence of any flying predators in this context implies that these alarms convey deceptive information about the type of danger present. Experiments on the primary nest predators of thornbills, pied currawongs (Strepera graculina), revealed that the predators treat these alarms as if they themselves are threatened by flying hawks, either by scanning the sky for danger or fleeing, confirming a deceptive function. In turn, these distractions delay attack and provide thornbill nestlings with an opportunity to escape. This sophisticated defence strategy exploits the complex web of interactions among multiple species across several trophic levels, and in particular exploits a predator''s ability to eavesdrop on and respond appropriately to heterospecific alarm calls. Our findings demonstrate that prey can fool predators by deceptively mimicking alarm calls of harmless species, suggesting that defensive mimicry could be more widespread because of indirect effects on predators within a web of eavesdropping.     

Context-dependent effects of predator removal from experimental microcosm communities

The loss of a predator from an ecological community can cause large changes in community structure and ecosystem processes, or have very little consequence for the remaining species and ecosystem. Understanding when and why the loss of a predator causes large changes in community structure and ecosystem processes is critical for understanding the functional consequences of biodiversity loss. We used experimental microbial communities to investigate how the removal of a large generalist predator affected the extinction frequency, population abundance and total biomass of its prey. We removed this predator in the presence or absence of an alternative, more specialist, predator in order to determine whether the specialist predator affected the outcome of the initial species removal. Removal of the large generalist predator altered some species' populations but many were unaffected and no secondary extinctions were observed. The specialist predator, though rare, altered the response of the prey community to the removal of the large generalist predator. In the absence of the specialist predator, the effects of the removal were only measurable at the level of individual species. However, when the specialist predator was present, the removal of the large generalist predator affected the total biomass of prey species. The results demonstrate that the effect of species loss from high trophic levels may be very context-dependent, as rare species can have disproportionately large effects in food webs.     

Consumptive and non-consumptive effects of an invasive marine predator on native coral-reef herbivores

Invasive predators typically have larger effects on native prey populations than native predators, yet the potential roles of their consumptive versus non-consumptive effects (CEs vs. NCEs) in structuring invaded systems remains unclear. Invasive lionfish (Pterois volitans) may have ecosystem-level effects by altering native fish grazing on benthic algae that could otherwise displace corals. Lionfish could reduce grazing by decreasing the abundance of herbivorous fishes (CEs), and/or the predation risk posed by lionfish could alter grazing behavior of fishes (NCEs). To test for these CEs, we manipulated lionfish densities on large reefs in The Bahamas and surveyed fish populations throughout June 2009–2011. In July 2011, NCEs of lionfish were measured by observing fish grazing behavior on algal-covered substrata placed in microhabitats varying in lionfish presence at different spatial scales, and quantifying any resulting algal loss. Lionfish reduced small herbivorous fish density by the end of the 2010 summer recruitment season. Grazing by small and large fishes was reduced on high-lionfish-density reefs, and small fish grazing further decreased when in the immediate presence of lionfish within-reefs. Lionfish had a negative indirect effect on algal loss, with 66–80 % less algae removed from substrata in high-lionfish-density reefs. Parrotfishes were likely driving the response of herbivorous fishes to both CEs and NCEs of lionfish. These results demonstrate the importance of considering NCEs in addition to CEs of invasive predators when assessing the effects of invasions.     

Experimental evidence for latent developmental plasticity: intertidal whelks respond to a native but not an introduced predator

Animals with highly inducible traits may show no inducible response when exposed to a related but wholly novel cue. This appears to be true for the intertidal whelk Nucella lamellosa faced with a voracious introduced predator. In the laboratory, we exposed whelks to effluent from two species of predatory crab, the native red rock crab Cancer productus and the invasive European green crab Carcinus maenas. Nucella and Cancer have a long shared history in the northeast Pacific, whereas potential interaction with Carcinus began here less than 10 years ago. Although Nucella responded adaptively to Cancer effluent by increasing shell thickness and decreasing somatic growth, there was no such response to Carcinus. Furthermore, thicker shelled Nucella were less likely to be eaten by Carcinus. Because Nucella produces thicker shells when exposed to Cancer cues, its ability to respond similarly to Carcinus depends only on the coupling of the Carcinus cue to the existing developmental pathways for adaptive changes in shell form. Such coupling of latent plasticity to a novel cue -- via genetic changes or associative learning -- could explain many cases of rapid phenotypic change following a sudden shift in the environment.     

Cranial morphological scaling and relative prey size limitations for a native predator in an invaded system

Over evolutionary time, predator-prey interactions have shaped and constrained functional and behavioral traits of piscivorous fishes. The endangered Colorado Pikeminnow Ptychocheilus lucius, a large endemic piscivore of the Colorado River Basin, encounters a substantially altered prey base that differs in behaviors and morphologies compared to the historical suite of native prey. To assess physical limitations of Colorado Pikeminnow predation, we conducted a feeding experiment with two species of nonnative prey (spined and despined Channel Catfish Ictalurus punctatus and Red Shiner Cyprinella lutrensis) and quantified scaling of cranial morphology in this predator. In our predation experiments, Colorado Pikeminnow (215–312 mm total length) consumed both spined and despined Channel Catfish as well as Red Shiner but only consumed prey less than 20% of the predator’s total length. Previous feeding trials using smaller Colorado Pikeminnow, with native and nonnative prey species, indicated they consumed prey up to 35% of their total length, suggesting relative prey size limits may decrease as this predator grows. Morphological measurements also suggested relative prey size suitability may decrease as Colorado Pikeminnow become larger, with head depth and width demonstrating isometric scaling at small sizes and shifting to negative allometry as fish get larger. Together, these data suggest an ontogenetic shift in the head morphology of Colorado Pikeminnow may decrease the relative size of prey available to these predators. In severely altered systems, understanding trophic characteristics that limit overall predator resource availability will be critical for conservation of piscivorous fishes.     

Urban bird declines and the fear of cats

The role of domestic cats Felis catus in the troubling, on-going decline of many urban bird populations in the UK is controversial. Debate, in the UK and elsewhere, has centred on the level of avian mortality directly imposed by cats, and on whether this is principally compensatory (the 'doomed surplus' hypothesis) or additive (the 'hapless survivor' hypothesis). However, it is well established that predators also have indirect, sub-lethal effects on their prey where life-history responses to predation risk affect birth and death rates. Here, using a simple model combining cat predation on birds with a sub-lethal (fear) effect of cat density on bird fecundity, we show that these sub-lethal effects may be substantial for urban songbirds. When cat densities are as high as has been recorded in the UK, and even when predation mortality is low (e.g. <1%), a small reduction in fecundity due to sub-lethal effects (e.g. <1 offspring year⁻¹ cat⁻¹) can result in marked decreases in bird abundances (up to 95%). Thus, low predation rates in urban areas do not necessarily equate with a correspondingly low impact of cats on birds. Sub-lethal effects may depress bird populations to such an extent that low predation rates simply reflect low prey numbers.     

Anti-predator behaviour of native prey (Daphnia) to an invasive predator (Bythotrephes longimanus) is influenced by predator density and water clarity

Hydrobiologia - Bythotrephes longimanus is an invasive zooplankton predator, negatively impacting zooplankton abundance and diversity in North American lakes. Previous studies have shown that...     

Multiple predator defence strategies in Daphnia pulex and their relation to native habitat

Daphnia may respond with an array of anti-predator defences(behavioural, morphological and life history) to a chemicalcue (kairomone) exuded by its predators: fish and Chaoborus.Given the wide array of potential responses, it is an interestingquestion whether anti-predator defences are coupled or independentof each other. Since anti-predator responses are costly andeven possessing the genetic information to respond to a certainpredator might involve a cost, clones may only react to predatorsthey co-occur with in nature. In this study, we provide evidencefor an uncoupling of responses by Daphnia pulex in several anti-predatordefences against Chaoborus. We were unable to detect a correlationbetween behavioural (migration), morphological (neck-spine induction)and life history [growth rate, neonate size and size at firstreproduction (SFR)] responses. Furthermore, anti-predator responsesdid not always comply with what is commonly believed. We foundthat Daphnia clones can migrate up or down when exposed to fishor Chaoborus kairomone and that population growth rate, neonatesize and SFR can increase or decrease in response to Chaoboruskairomone. We also show patterns in anti-predator defences thatseem to relate to the habitat from which clones were derived.Daphnia clones that were collected in habitats with Chaoborusas the dominant predator tended to react strongly to Chaoboruskairomone by migrating upward and producing neck-spines. Themigration behaviour against fish kairomone in these clones wasoften an unexpected upward migration. The Daphnia clone thatco-existed with fish predators showed a downward migration inthe presence of fish as well as Chaoborus kairomone. Clonesthat had occurred with either both or no predators had mixedresponses. We sometimes found an upward migration in combinationwith smaller body size as a response to Chaoborus kairomone.This may be interpreted as a behavioural defence against Chaoborusand a life-history defence against fish. Daphnia seem not toexhibit defence behaviour against predators they do not co-occurwith. It might be costly for Daphnia to maintain genetic informationto respond to these predators and protect that information fromgenetic drift.     

The impact of short-term predator removal on vole dynamics in an arctic-alpine landscape

During 1991–95, mammalian predators (weasel, Mustela nivalis , stoat, M. erminea , mink, M. vison , and red fox, Vulpes vulpes ) were excluded in late summers from a 2 ha piece of a north Norwegian mountain slope. The exclosure extended from an outpost of luxuriant sub-arctic birch forest to a typical arctic–alpine habitat complex, including productive willow scrublands, tundra heaths, dry ridges and snow-beds. The exclosure thus encompassed the entire range of habitat conditions encountered in a typical north Fennoscandian mountain and tundra landscapes. During 1991–95, the exclosure was predator-proof from late July to late September. In wintertime and in early summer, the exclosure was accessible to mammalian predators. Vole dynamics in the short-term exclosure were compared to dynamics in five reference areas with similar habitat conditions.
In 1991, when vole densities were rising in the area, neither collective vole densities nor densities of individual vole species differed significantly between the exclosure and the replicated controls. Spring densities of voles were never significantly different between the exclosure and the controls. With respect to autumnal densities of voles, however, the exclosure was a statistical outlier in the peak year 1992 and throughout the gradual decline phase of 1993–95. In the peak year, the difference in collective vole densities was modest (30%), but increased to two-fold during the first two decline years and was almost four-fold in the crash year of 1995. The strongest response was displayed by field voles ( Microtus agrestis ), hypothesized to be the pivotal prey species of weasels, especially by females and young individuals, i.e. by the functional categories especially sensitive to mammalian predation. These results are consistent with the hypothesis that predation plays a pivotal role for the regulation of herbivorous mammals in relatively productive arctic–alpine habitats.     

Diversity and composition of cultivable gut bacteria in an endemic island bird and its mainland sister species

The study of island biodiversity has inspired many advances in evolutionary biology. However, whether patterns of microorganism diversity are influenced by insularity is poorly understood. In particular, microorganisms that live in symbiotic association, such as the microbiota that inhabit the gastrointestinal tract of bigger animals, are subjected to demographic and coevolutionary processes that may add complexity to the common expectation of impoverished diversity on oceanic islands. Here, we explore this topic by studying the cultivable gut bacteria of two sister species of birds, from São Tomé island and nearby mainland Gabon, the endemic São Tomé thrush Turdus olivaceofuscus and the African thrush Turdus pelios. We found no differences in the diversity of cultivable gut bacteria between these thrushes, suggesting that, unlike what is commonly found for macrofauna, insularity might not represent a strong constraint for gut bacterial diversity. Although further research on complete gut bacterial communities and a broader range of species and areas is needed, our initial results suggest that the cultivable gut microbial community may bypass the diversity loss associated with island colonization. This could arise from intrinsic factors such as their large population sizes within hosts and low rates of extinction. Furthermore, as gut communities are composed mainly by mutualistic bacteria, diversifying selection (against an impoverished bacterial community), may counteract the diversity loss brought about by the stochastic and demographic effects of the founder process.     

Invasive plant removal increases insect herbivory pressure on a native tree due to an increase in resource quality

Insect predator densities and plant nutritional status can affect insect herbivore densities, but these effects have not yet been assessed in the context of an invasive species management. This study investigated the influence of (i) altered predatory arthropod abundance and community composition (top-down-effects); and (ii) altered leaf nitrogen and phosphorus levels (bottom-up-effects) as possible causes of fluctuations in herbivore abundance and herbivory on the native tree Brabejum stellatifolium in riparian sites that differ in invasion status (near pristine, heavily invaded by Acacia mearnsii, and restored). Species richness, abundance and composition of herbivorous and predatory arthropods were compared between categories. Herbivore and predator abundance, species richness and assemblage composition were significantly influenced by invasion status. As expected, herbivore abundance was significantly and positively correlated to levels of leaf damage. Leaf nitrogen levels were the highest at restored sites where herbivorous arthropods were also most abundant and the abundance of predators was intermediate. These results suggest that altered leaf nitrogen content is likely a key influence for the increased herbivore pressure on B. stellatifolium at sites under pressure from invasive A. mearnsii and its status. Bottom-up influences therefore may be more important than top-down effects on controlling herbivore abundances in these altered riparian environments.