Resembling a Viper: Implications of Mimicry for Conservation of the Endangered Smooth Snake

The phenomenon of Batesian mimicry, where a palatable animal gains protection against predation by resembling an unpalatable model, has been a core interest of evolutionary biologists for 150 years. An extensive range of studies has focused on revealing mechanistic aspects of mimicry (shared education and generalization of predators) and the evolutionary dynamics of mimicry systems (co‐operation vs. conflict) and revealed that protective mimicry is widespread and is important for individual fitness. However, according to our knowledge, there are no case studies where mimicry theories have been applied to conservation of mimetic species. Theoretically, mimicry affects, for example, frequency dependency of predator avoidance learning and human induced mortality. We examined the case of the protected, endangered, nonvenomous smooth snake (Coronella austriaca) that mimics the nonprotected venomous adder (Vipera berus), both of which occur in the Åland archipelago, Finland. To quantify the added predation risk on smooth snakes caused by the rarity of vipers, we calculated risk estimates from experimental data. Resemblance of vipers enhances survival of smooth snakes against bird predation because many predators avoid touching venomous vipers. Mimetic resemblance is however disadvantageous against human predators, who kill venomous vipers and accidentally kill endangered, protected smooth snakes. We found that the effective population size of the adders in Åland is very low relative to its smooth snake mimic (28.93 and 41.35, respectively).Because Batesian mimicry is advantageous for the mimic only if model species exist in sufficiently high numbers, it is likely that the conservation program for smooth snakes will fail if adders continue to be destroyed. Understanding the population consequences of mimetic species may be crucial to the success of endangered species conservation. We suggest that when a Batesian mimic requires protection, conservation planners should not ignore the model species (or co‐mimic in Mullerian mimicry rings) even if it is not itself endangered. Implications of mimicry for Conservation of the endangered smooth snake     

Niche complementarity due to plasticity in resource use: plant partitioning of chemical N forms

Niche complementarity, in which coexisting species use different forms of a resource, has been widely invoked to explain some of the most debated patterns in ecology, including maintenance of diversity and relationships between diversity and ecosystem function. However, classical models assume resource specialization in the form of distinct niches, which does not obviously apply to the broadly overlapping resource use in plant communities. Here we utilize an experimental framework based on competition theory to test whether plants partition resources via classical niche differentiation or via plasticity in resource use. We explore two alternatives: niche preemption, in which individuals respond to a superior competitor by switching to an alternative, less-used resource, and dominant plasticity, in which superior competitors exhibit high resource use plasticity and shift resource use depending on the competitive environment. We determined competitive ability by measuring growth responses with and without neighbors over a growing season and then used 15N tracer techniques to measure uptake of different nitrogen (N) forms in a field setting. We show that four alpine plant species of differing competitive abilities have statistically indistinguishable uptake patterns (nitrate > ammonium > glycine) in their fundamental niche (without competitors) but differ in whether they shift these uptake patterns in their realized niche (with competitors). Competitively superior species increased their uptake of the most available N form, ammonium, when in competition with the rarer, competitively inferior species. In contrast, the competitively inferior species did not alter its N uptake pattern in competition. The existence of plasticity in resource use among the dominant species provides a mechanism that helps to explain the manner by which plant species with broadly overlapping resource use might coexist.     

Population dynamics of Müllerian mimicry under interspecific competition

We ask what the effects of mutualism on population dynamics of two competitive species are. We model the population dynamics of mutualistic interactions with positive density- and frequency-dependences. We specifically assume the dynamics of Müllerian mimicry in butterflies, where the mortality of both species is reduced depending on the relative frequency of the other species. We assume that the two species are under Lotka–Volterra density-dependent competition. The equilibria are compared with the cases of competition alone. Unlike the traditional model of positive density-dependence, population explosion does not appear in the current dynamics, but the new equilibrium is simply achieved. It is because the effects of positive density- or frequency-dependence are restricted to parts of mortality. Both positive density- and frequency-dependences do promote coexistence of the mimetic species. However, the two models show a distinctive difference for coexistence. The effects of positive density-dependence are rather limited. In contrast, positive frequency-dependence always promotes coexistence, irrespective of environmental conditions. The results may imply that the evolutionary origin of Müllerian mimicry may depend on frequency-dependence (and density-dependence), but that its current population dynamics may depend solely on density-dependence. The role of frequency- and density-dependences on evolutionary dynamics is an open question.     

Size- and food-dependent growth drives patterns of competitive dominance along productivity gradients

Patterns of coexistence among competing species exhibiting size- and food-dependent growth remain largely unexplored. Here we studied mechanisms behind coexistence and shifts in competitive dominance in a size-structured fish guild, representing sprat and herring stocks in the Baltic Sea, using a physiologically structured model of competing populations. The influence of degree of resource overlap and the possibility of undergoing ontogenetic diet shifts were studied as functions of zooplankton and zoobenthos productivity. By imposing different size-dependent mortalities, we could study the outcome of competition under contrasting environmental regimes representing poor and favorable growth conditions. We found that the identity of the dominant species shifted between low and high productivity. Adding a herring-exclusive benthos resource only provided a competitive advantage over sprat when size-dependent mortality was high enough to allow for rapid growth in the zooplankton niche. Hence, the importance of a bottom-up effect of varying productivity was dependent on a strong top-down effect. Although herring could depress shared resources to lower levels than could sprat and also could access an exclusive resource, the smaller size at maturation of sprat allowed it to coexist with herring and, in some cases, exclude it. Our model system, characterized by interactions among size cohorts, allowed for consumer coexistence even at full resource overlap at intermediate productivities when size-dependent mortality was low. Observed shifts in community patterns were crucially dependent on the explicit consideration of size- and food-dependent growth. Accordingly, we argue that accounting for food-dependent growth and size-dependent interactions is necessary to better predict changes in community structure and dynamics following changes in major ecosystem drivers such as resource productivity and mortality, which are fundamental for our ability to manage exploitation of living resources in, e.g., fisheries.     

Delayed plumage maturation in Lazuli buntings: tests of the female mimicry and status signalling hypotheses

The evolutionary importance of delayed plumage maturation (DPM) in passerines, the condition when more than 1 year is required to achieve adult-like coloration, remains highly contentious. Adaptive hypotheses propose that aggression from after 2nd-year (ASY) males or predation favors DPM in 2nd-year (SY) males, thereby increasing SY male survivorship or reproductive success. However, each hypothesis suggests a distinct selective mechanism explaining “how” this is accomplished. Alternatively, DPM may be a consequence of a nonadaptive molt constraint. We tested the female mimicry and status signalling hypotheses in territorial ASY male lazuli buntings (Passerinaamoena) using three sets of model presentation experiments. The female mimicry hypothesis proposes that dull SY male plumage deceptively mimics female plumage, and predicts that ASY males can not distinguish SY male from female plumage. The status signalling hypothesis proposes that dull SY male plumage honestly signals low competitive threat, and predicts that ASY males respond less aggressively to dull versus bright, ASY-like plumage. Contrary to the female mimicry hypothesis, ASY males distinguished between SY male and female plumage, as they were aggressive to SY male models exclusively and attempted to copulate with female models. Supporting the status signalling hypothesis, ASY males were significantly less aggressive to SY versus ASY male plumage. While DPM may result from a physiological constraint on bright SY male plumage, our results support the idea that dull plumage in an SY male's first breeding season may be maintained by selection to reduce aggression from ASY males, serving as a signal of competitive status. Received: 21 February 1997 / Accepted after revision: 16 June 1997     

Assessing the Value of the Umbrella‐Species Concept for Conservation Planning with Meta‐Analysis

Abstract: The umbrella‐species concept, which suggests that conservation strategies designed for one species may benefit co‐occurring species, has been promoted as a framework for conservation planning. Nevertheless, there has been considerable variation in the outcome of empirical tests of this concept that has led researchers to question its value, so we used data from 15 published studies in a meta‐analysis to evaluate whether conservation of putative umbrella species also conserves co‐occurring species. We tested the effectiveness of putative umbrella species categorized by taxonomic group, taxonomic similarity to co‐occurring species, body size, generality of resource use, and trophic level to evaluate criteria proposed to guide the selection of umbrella species. We compared species richness and number of individuals (by species and higher taxonomic group) between sites with and without putative umbrella species to test whether more co‐occurring species were present in greater abundances when the area or resource needs of umbrella species were met. Species richness and abundance of co‐occurring species were consistently higher in sites where umbrella species were present than where they were not and for conservation schemes with avian than with mammalian umbrella species. There were no differences in species richness or species abundance with resource generalist or specialist umbrella species or based on taxonomic similarity of umbrella and co‐occurring species. Taxonomic group abundance was higher in across‐taxonomic umbrella species schemes than when umbrella species were of the same taxon as co‐occurring species. Co‐occurring species had similar, or higher, species richness with small‐bodied umbrella species relative to larger‐bodied umbrella species. The only significant difference among umbrella species categorized by trophic level was that species richness was higher with omnivorous than it was with carnivorous avian umbrella species. Our results suggest there is merit to the umbrella‐species concept for conservation, but they do not support the use of the criteria we used to identify umbrella species.     

Why do Horsfield’s bronze-cuckoo <Emphasis Type="Italic">Chalcites basalis</Emphasis> eggs mimic those of their hosts?

The Horsfield’s bronze-cuckoo (Chalcites basalis) egg closely matches the appearance of its host fairy-wren (Malurus spp.) eggs. Mimicry of host eggs by cuckoos is usually attributed to coevolution between cuckoos and hosts, with host discrimination against odd-looking eggs selecting for ever better mimicry by cuckoos. However, this process cannot explain Horsfield’s bronze-cuckoo egg mimicry because fairy-wren hosts rarely reject odd-looking eggs from their nest. An alternative hypothesis is that cuckoos have evolved egg mimicry to disguise their eggs from other cuckoos. Female cuckoos remove one egg from the nest during parasitism and would potentially benefit by selectively removing any cuckoo egg that has already been laid in the nest because otherwise, their egg will be evicted by the first nestling cuckoo along with the host clutch. We used painted, non-mimetic eggs to test whether cuckoos selectively remove odd-looking eggs during parasitism. We found that they were no more likely to remove a non-mimetic egg from a superb fairy-wren Malurus cyaneus clutch than would be expected by chance. Thus, our study does not support the cuckoo egg replacement hypothesis to explain mimicry of host eggs by cuckoos.     

Niche and fitness differences relate the maintenance of diversity to ecosystem function

The frequently observed positive correlation between species diversity and community biomass is thought to depend on both the degree of resource partitioning and on competitive dominance between consumers, two properties that are also central to theories of species coexistence. To make an explicit link between theory on the causes and consequences of biodiversity, we define in a precise way two kinds of differences among species: niche differences, which promote coexistence, and relative fitness differences, which promote competitive exclusion. In a classic model of exploitative competition, promoting coexistence by increasing niche differences typically, although not universally, increases the "relative yield total", a measure of diversity's effect on the biomass of competitors. In addition, however, we show that promoting coexistence by decreasing relative fitness differences also increases the relative yield total. Thus, two fundamentally different mechanisms of species coexistence both strengthen the influence of diversity on biomass yield. The model and our analysis also yield insight on the interpretation of experimental diversity manipulations. Specifically, the frequently reported "complementarity effect" appears to give a largely skewed estimate of resource partitioning. Likewise, the "selection effect" does not seem to isolate biomass changes attributable to species composition rather than species richness, as is commonly presumed. We conclude that past inferences about the cause of observed diversity-function relationships may be unreliable, and that new empirical estimates of niche and relative fitness differences are necessary to uncover the ecological mechanisms responsible for diversity-function relationships.     

Disaggregated intertemporal models with an exhaustible resource and technical advance

We compare results of four organizations of an economy that is dependent upon an irreplenishable resource but may develop a new technology that releases that dependence. We find a decentralized competitive organization can replicate the behavior of a centrally planned economy, given an appropriate distribution of initial wealth. This holds whether the R&D is done by the government or by the natural resource sector, so long as borrowing from other sectors is possible. Further, the economy's behavior is independent of whether the resource sector is monopolistic or competitive if and only if the production function is Cobb-Douglas.     

Plant competition varies with community composition in an edaphically complex landscape

There is currently no consensus on how physical and biological factors affect competitive intensity. Tests of whether competitive intensity varies along axes of environmental change have commonly been conducted in systems with a single strong environmental gradient, such as productivity, a soil resource, or an environmental stress. Frequently, these same axes are associated with changes in species composition, yet few studies have asked whether shifts in the identity of competitors affect competitive intensity. We ask whether resources (nutrients, water), stressors (heavy metals, Ca:Mg ratio), productivity (aboveground biomass), or species identity (an ordination axis of plant community composition) were the best predictors of the intensity of competition in a heterogeneous grassland landscape that included multiple independent environmental gradients. The reproductive fitness of six annual plant species was measured in the presence and absence of competitors and used to calculate relative interaction intensity (RII). We found that RII was best predicted by community composition. Nutrient availability was also important, and a post hoc test showed that competitive intensity was best explained by the combined effects of community composition and nutrient availability. We argue that community composition may be the most effective metric for predicting competitive intensity in many ecosystems because it includes both the competitive effects of the local community and information about covarying environmental characteristics.     

The cost of endangered species protection: Evidence from auctions for natural resources

This paper examines the effect that endangered species regulation has on natural resource development. Specifically, we use data from competitive auctions to estimate the effect that land-use regulation protecting endangered caribou in the Canadian province of Alberta has on the price producers pay for the right to extract oil. We exploit a regression discontinuity design to evaluate how prices differ along regulation boundaries that constrain resource development. The auction format and the regulation discontinuity allow us to measure the total cost of the regulation. We find that producers pay 24% less on average for oil leases that are regulated and that the total net present value cost of the regulation exceeds $1.15 billion for leases sold between 2003 and 2012, all of which is borne by the government. In spite of these costs, the populations of endangered caribou remain in widespread decline.     

Multidimensional trade-offs in species responses to disturbance: implications for diversity in a subtropical forest

Species employ diverse strategies to cope with natural disturbance, but the importance of these strategies for maintaining tree species diversity in forests has been debated. Mechanisms that have the potential to promote tree species coexistence in the context of repeated disturbance include life history trade-offs in colonization and competitive ability or in species' ability to survive at low resource conditions and exploit the temporary resource-rich conditions often generated in the wake of disturbance (successional niche). Quantifying these trade-offs requires long-term forest monitoring and modeling. We developed a hierarchical Bayes model to investigate the strategies tree species employ to withstand and recover from hurricane disturbance and the life history trade-offs that may facilitate species coexistence in forests subject to repeated hurricane disturbance. Unlike previous approaches, our model accommodates temporal variation in process error and observations from multiple sources. We parameterized the model using growth and mortality data from four censuses of a 16-ha plot taken every five years (1990-2005), together with damage data collected after two hurricanes and annual seed production data (1992-2005). Species' susceptibilities to hurricane damage as reflected by changes in diameter growth and fecundity immediately following a storm were weak, highly variable, and unpredictable using traditional life history groupings. The lower crowding conditions (e.g., high light) generated in the wake of storms, however, led to greater gains in growth and fecundity for pioneer and secondary-forest species than for shade-tolerant species, in accordance with expectation of life history. We found moderate trade-offs between survival in high crowding conditions, a metric of competitive ability, and long-distance colonization. We also uncovered a strong trade-off between mean species fecundity in low crowding conditions, a metric of recovery potential, and competitive ability. Trade-offs in competitive and colonization ability, in addition to successional niche processes, are likely to contribute to species persistence in these hurricane-impacted forests. The strategies species employ to cope with hurricane damage depend on the degree to which species rely on sprouting, repair of adult damage, changes in demographic rates in response to enhanced resource availability after storms, or long-distance dispersal as recovery mechanisms.     

Attitudes towards risk and the optimal exploitation of an exhaustible resource

The exploitation of a nonrenewable natural resource, such as petroleum or mineral ores, is analyzed in a stochastic framework with price uncertainty. The market setting may be either monopolistic or competitive. We demonstrate that the rate of extraction varies directly with the resource owner's willingness to accept risk. Rish-preferring owners use the resource more rapidly than risk-neutral owners, who in turn deplete the resource more rapidly than risk-averse owners. It is also seen that the usual practice of increasing the discount rate to account for risk induces a more rapid rate of resource use, when in fact a slower rate of depletion is desired.     

Female mimicry and resource defense polygyny by males of a tropical rove beetle,Leistotrophus versicolor (Coleoptera : Staphylinidae)

Summary Adults of the staphylinid beetle Leistotrophus versicolor Grav. aggregate at vertebrate dung and carrion where males and females forage for adult Diptera. Some males aggressively exclude others from dung and carrion. Winners in male combat gain access to many females, which are often receptive at these foraging sites. The mating system can be categorized as resource defense polygyny. Males vary greatly in size, are larger than females on average, and have allometrically enlarged mandibles that they use in fighting. Large males consistently defeat smaller ones. Some males employ female mimicry in order to avoid aggression, remaining at dung where they forage and even obtain copulations while being courted by rival males. Female mimicry is most often practiced by males that are smaller than their rivals or by males that are unable to use their jaws aggressively because they are feeding or courting females when encountered by an opponent. Female mimicry is a conditional tactic of mature males; some individuals behave like females toward larger males but attack smaller rivals. Offprint requests to: J. Alcock     

Prevalence of sustainable and unsustainable use of wild species inferred from the IUCN Red List of Threatened Species

Unsustainable exploitation of wild species represents a serious threat to biodiversity and to the livelihoods of local communities and Indigenous peoples. However, managed, sustainable use has the potential to forestall extinctions, aid recovery, and meet human needs. We analyzed species-level data for 30,923 species from 13 taxonomic groups on the International Union for Conservation of Nature Red List of Threatened Species to investigate patterns of intentional biological resource use. Forty percent of species (10,098 of 25,009 species from 10 data-sufficient taxonomic groups) were used. The main purposes of use were pets, display animals, horticulture, and human consumption. Intentional use is currently contributing to elevated extinction risk for 28–29% of threatened or near threatened (NT) species (2752–2848 of 9753 species). Intentional use also affected 16% of all species used (1597–1631 of 10,098). However, 72% of used species (7291 of 10,098) were least concern, of which nearly half (3469) also had stable or improving population trends. The remainder were not documented as threatened by biological resource use, including at least 172 threatened or NT species with stable or improving populations. About one-third of species that had use documented as a threat had no targeted species management actions to directly address this threat. To improve use-related red-list data, we suggest small amendments to the relevant classification schemes and required supporting documentation. Our findings on the prevalence of sustainable and unsustainable use, and variation across taxa, can inform international policy making, including the Intergovernmental Science-Policy Platform on Biodiversity and Ecosystem Services, the Convention on Biological Diversity, and the Convention on International Trade in Endangered Species.     

Root plasticity buffers competition among plants: theory meets experimental data

Morphological plasticity is a striking characteristic of plants in natural communities. In the context of foraging behavior particularly, root plasticity has been documented for numerous species. Root plasticity is known to mitigate competitive interactions by reducing the overlap of the individuals' rhizospheres. But despite its obvious effect on resource acquisition, plasticity has been generally neglected in previous empirical and theoretical studies estimating interaction intensity among plants. In this study, we developed a semi-mechanistic model that addresses this shortcoming by introducing the idea of compensatory growth into the classical-zone-of influence (ZOI) and field-of-neighborhood (FON) approaches. The model parameters describing the belowground plastic sphere of influence (PSI) were parameterized using data from an accompanying field experiment. Measurements of the uptake of a stable nutrient analogue at distinct distances to the neighboring plants showed that the study species responded plastically to belowground competition by avoiding overlap of individuals' rhizospheres. An unexpected finding was that the sphere of influence of the study species Bromus hordeaceus could be best described by a unimodal function of distance to the plant's center and not with a continuously decreasing function as commonly assumed. We employed the parameterized model to investigate the interplay between plasticity and two other important factors determining the intensity of competitive interactions: overall plant density and the distribution of individuals in space. The simulation results confirm that the reduction of competition intensity due to morphological plasticity strongly depends on the spatial structure of the competitive environment. We advocate the use of semi-mechanistic simulations that explicitly consider morphological plasticity to improve our mechanistic understanding of plant interactions.     

Feeding and non-feeding aggression can be induced in invasive shore crabs by altering food distribution

Human activity can have a large impact on surrounding ecosystems. For example, humans alter resource distributions for other species, potentially modifying these species competitive dynamics. These changes in local competitive processes are frequently associated with species invasions. Here, we investigate how differences in resource distribution affect competitive behaviour using the highly invasive European shore crab (Carcinus maenas). Using a controlled laboratory experiment in combination with behaviour assays and social network analysis, we show that individuals feeding in habitats with clumped food distributions are more aggressive than individuals feeding in habitats where food is evenly dispersed, and this aggression is present even on days where crabs are not feeding. Additionally, this persistent aggression can be induced, suggesting that individuals of this invasive species possess the flexibility to modify their competitive behaviours in response to differences in food distributions. Furthermore, we show how these individual responses can lead to changes in overall organisation of aggressive interactions within a population. We discuss these results in relation to how human impacts can have long-term effects on competitive behavioural strategies, and how behavioural flexibility can allow invasive species to colonise and persist in highly impacted sites such as urban ecosystems.     

Rarity of Organisms in the Sand Pine Scrub Habitat of Florida

Naturalists conceive of "rare" organisms as those narrow in geographic range, restricted to few habitats, or small in population sizes. Because rare organisms may be rare in a variety of ways, a particular conservation strategy that purports to protect them may not do so effectively in all cases; therefore, categorization of rarity in advance could help ensure that the chosen strategy has a reasonable chance of success in a particular case. We placed plants, amphibians, and reptiles resident in the Sand Pine scrub habitat of Florida into categories of rarity. Determining if and in what way a particular organism is rare proved difficult, because the process requires exact specification of what is meant by "narrow,""restricted," and "small." Furthermore, the process requires choice of a scale for judging rarity and relies upon data that are likely to be of variable availability and reliability. We categorized rarity by several schemes, and they produced distributions of taxa among categories of rarity that largely were similar to one another but that diverged in telling ways. The distributions of taxa among categories of rarity produced by any scheme were different for scrub plants than for scrub amphibians and reptiles. Likewise, these distributions were different than those produced by previously published studies of rarity. We found that the results of our categorization schemes did not match particularly well with the official listing of scrub organisms as endangered or potentially endangered.