Jonathan M. Jeschke

PREDATOR FUNCTIONAL RESPONSES: DISCRIMINATING BETWEEN HANDLING AND DIGESTING PREY

We present a handy mechanistic functional response model that realistically incorporates handling (i.e., attacking and eating) and digesting prey. We briefly review current functional response theory and thereby demonstrate that such a model has been lacking so far. In our model, we treat digestion as a background process that does not prevent further foraging activities (i.e., searching and handling). Instead, we let the hunger level determine the probability that the predator searches for new prey. Additionally, our model takes into account time wasted through unsuccessful attacks. Since a main assumption of our model is that the predators hunger is in a steady state, we term it the steady-state satiation (SSS) equation. The SSS equation yields a new formula for the asymptotic maximum predation rate (i.e., asymptotic maximum number of prey eaten per unit time, for prey density approaching infinity). According to this formula, maximum predation rate is determined not by the sum of the time spent for handling and digesting prey, but solely by the larger of these two terms. As a consequence, predators can be categorized into two types: handling-limited predators (where maximum predation rate is limited by handling time) and digestion-limited predators (where maximum predation rate is limited by digestion time). We give examples of both predator types. Based on available data, we suggest that most predators are digestion limited. The SSS equation is a conceptual mechanistic model. Two possible applications of this model are that (1) it can be used to calculate the effects of changing predator or prey characteristics (e.g., defenses) on predation rate and (2) optimal foraging models based on the SSS equation are testable alternatives to other approaches. This may improve optimal foraging theory, since one of its major problems has been the lack of alternative models.

Are invaders different? A conceptual framework of comparative approaches for assessing determinants of invasiveness

What determines invasiveness of alien organisms is among the most interesting and urgent questions in ecology. In attempts to answer this question, researchers compare invasive alien species either to native species or to non-invasive alien species, and this is done in either the introduced or native ranges. However, inferences that can be drawn from these comparisons differ considerably, and failure to recognize this could hamper the search for determinants of invasiveness. To increase awareness about this issue, we present a framework of the various comparisons that can be used to test for determinants of invasiveness, and the specific questions each comparison can address. Moreover, we discuss how different comparisons complement each other, and therefore should be used in concert. For progress in invasion biology, it is crucial to realize that different comparisons address different biological questions and that some questions can only be answered unambiguously by combining them.

Usefulness of Bioclimatic Models for Studying Climate Change and Invasive Species

Bioclimatic models (also known as envelope models or, more broadly, ecological niche models or species distribution models) are used to predict geographic ranges of organisms as a function of climate. They are widely used to forecast range shifts of organisms due to climate change, predict the eventual ranges of invasive species, infer paleoclimate from data on species occurrences, and so forth. Several statistical techniques (including general linear models, general additive models, climate envelope models, classification and regression trees, and genetic algorithms) have been used in bioclimatic modeling. Recently developed techniques tend to perform better than older techniques, although it is unlikely that any single statistical approach will be optimal for all applications and species. Proponents of bioclimatic models have stressed their apparent predictive power, whereas opponents have identified the following unreasonable model assumptions: biotic interactions are unimportant in determining geographic ranges or are constant over space and time; the genetic and phenotypic composition of species is constant over space and time; and species are unlimited in their dispersal. In spite of these problematic assumptions, bioclimatic models often successfully fit present‐day ranges of species. Their ability to forecast the effects of climate change or the spread of invaders has rarely been tested adequately, however, and we urge researchers to tie the evaluation of bioclimatic models more closely to their intended uses.

A unified classification of alien species based on the magnitude of their environmental impacts.

We present a method for categorising and comparing alien or invasive species in terms of how damaging they are to the environment, that can be applied across all taxa, scales, and impact metrics.

Consumer-food systems: why type I functional responses are exclusive to filter feeders

The functional response of a consumer is the relationship between its consumption rate and the abundance of its food. A functional response is said to be of type I if consumption rate increases linearly with food abundance up to a threshold level at which it remains constant. According to conventional wisdom, such type I responses are more frequent among filter feeders than among other consumers. However, the validity of this claim has never been tested. We review 814 functional responses from 235 studies, thereby showing that type I responses are not only exceptionally frequent among filter feeders but that they have only been reported from these consumers.

Defining the Impact of Non-Native Species

Non-native species cause changes in the ecosystems to which they are introduced. These changes, or some of them, are usually termed impacts; they can be manifold and potentially damaging to ecosystems and biodiversity. However, the impacts of most non-native species are poorly understood, and a synthesis of available information is being hindered because authors often do not clearly define impact. We argue that explicitly defining the impact of non-native species will promote progress toward a better understanding of the implications of changes to biodiversity and ecosystems caused by non-native species; help disentangle which aspects of scientific debates about non-native species are due to disparate definitions and which represent true scientific discord; and improve communication between scientists from different research disciplines and between scientists, managers, and policy makers. For these reasons and based on examples from the literature, we devised seven key questions that fall into 4 categories: directionality, classification and measurement, ecological or socio-economic changes, and scale. These questions should help in formulating clear and practical definitions of impact to suit specific scientific, stakeholder, or legislative contexts. Definiendo el Impacto de las Especies No-Nativas Resumen Las especies no-nativas pueden causar cambios en los ecosistemas donde son introducidas. Estos cambios, o algunos de ellos, usualmente se denominan como impactos; estos pueden ser variados y potencialmente dañinos para los ecosistemas y la biodiversidad. Sin embargo, los impactos de la mayoría de las especies no-nativas están pobremente entendidos y una síntesis de información disponible se ve obstaculizada porque los autores continuamente no definen claramente impacto. Discutimos que definir explícitamente el impacto de las especies no-nativas promoverá el progreso hacia un mejor entendimiento de las implicaciones de los cambios a la biodiversidad y los ecosistemas causados por especies no-nativas; ayudar a entender cuáles aspectos de los debates científicos sobre especies no-nativas son debidos a definiciones diversas y cuáles representan un verdadero desacuerdo científico; y mejorar la comunicación entre científicos de diferentes disciplinas y entre científicos, administradores y quienes hacen las políticas. Por estas razones y basándonos en ejemplos tomados de la literatura, concebimos siete preguntas clave que caen en cuatro categorías: direccionalidad, clasificación y medida, cambios ecológicos o socio-económicos, y escala. Estas preguntas deberían ayudar en la formulación de definiciones claras y prácticas del impacto para encajar mejor con contextos científicos, de las partes interesadas o legislativos específicos.

No saturation in the accumulation of alien species worldwide

Although research on human-mediated exchanges of species has substantially intensified during the last centuries, we know surprisingly little about temporal dynamics of alien species accumulations across regions and taxa. Using a novel database of 45,813 first records of 16,926 established alien species, we show that the annual rate of first records worldwide has increased during the last 200 years, with 37% of all first records reported most recently (1970–2014). Inter-continental and inter-taxonomic variation can be largely attributed to the diaspora of European settlers in the nineteenth century and to the acceleration in trade in the twentieth century. For all taxonomic groups, the increase in numbers of alien species does not show any sign of saturation and most taxa even show increases in the rate of first records over time. This highlights that past efforts to mitigate invasions have not been effective enough to keep up with increasing globalization.

Advancing impact prediction and hypothesis testing in invasion ecology using a comparative functional response approach

Invasion ecology urgently requires predictive methodologies that can forecast the ecological impacts of existing, emerging and potential invasive species. We argue that many ecologically damaging invaders are characterised by their more efficient use of resources. Consequently, comparison of the classical ‘functional response’ (relationship between resource use and availability) between invasive and trophically analogous native species may allow prediction of invader ecological impact. We review the utility of species trait comparisons and the history and context of the use of functional responses in invasion ecology, then present our framework for the use of comparative functional responses. We show that functional response analyses, by describing the resource use of species over a range of resource availabilities, avoids many pitfalls of ‘snapshot’ assessments of resource use. Our framework demonstrates how comparisons of invader and native functional responses, within and between Type II and III functional responses, allow testing of the likely population-level outcomes of invasions for affected species. Furthermore, we describe how recent studies support the predictive capacity of this method; for example, the invasive ‘bloody red shrimp’ Hemimysis anomala shows higher Type II functional responses than native mysids and this corroborates, and could have predicted, actual invader impacts in the field. The comparative functional response method can also be used to examine differences in the impact of two or more invaders, two or more populations of the same invader, and the abiotic (e.g. temperature) and biotic (e.g. parasitism) context-dependencies of invader impacts. Our framework may also address the previous lack of rigour in testing major hypotheses in invasion ecology, such as the ‘enemy release’ and ‘biotic resistance’ hypotheses, as our approach explicitly considers demographic consequences for impacted resources, such as native and invasive prey species. We also identify potential challenges in the application of comparative functional responses in invasion ecology. These include incorporation of numerical responses, multiple predator effects and trait-mediated indirect interactions, replacement versus non-replacement study designs and the inclusion of functional responses in risk assessment frameworks. In future, the generation of sufficient case studies for a meta-analysis could test the overall hypothesis that comparative functional responses can indeed predict invasive species impacts.

Density-dependent effects of prey defences.

Abstract In this study, we show that the protective advantage of a defence depends on prey density. For our investigations, we used the predator-prey model system Chaoborus-Daphnia pulex. The prey, D. pulex, forms neckteeth as an inducible defence against chaoborid predators. This morphological response effectively reduces predator attack efficiency, i.e. number of successful attacks divided by total number of attacks. We found that neckteeth-defended prey suffered a distinctly lower predation rate (prey uptake per unit time) at low prey densities. The advantage of this defence decreased with increasing prey density. We expect this pattern to be general when a defence reduces predator success rate, i.e. when a defence reduces encounter rate, probability of detection, probability of attack, or efficiency of attack. In addition, we experimentally simulated the effects of defences which increase predator digestion time by using different sizes of Daphnia with equal vulnerabilities. This type of defence had opposite density-dependent effects: here, the relative advantage of defended prey increased with prey density. We expect this pattern to be general for defences which increase predator handling time, i.e. defences which increase attacking time, eating time, or digestion time. Many defences will have effects on both predator success rate and handling time. For these defences, the predator’s functional response should be decreased over the whole range of prey densities.

Prey swarming: which predators become confused and why?

When confronted with a swarm of their prey, many predators become confused and are less successful in their attacks. Despite the general notion that this confusion effect is a major reason for prey swarm formation, it is largely unknown how widespread it is and which predator or prey traits facilitate or impede it. We carried out experiments with four predator–prey systems: Aeshna and Chaoborus larvae, but not Libellula and Triturus larvae, became confused when confronted with high Daphnia densities. When combining this result with literature data, we found that predators became confused in 16 of the 25 predator–prey systems studied to date. Tactile predators appear to be generally susceptible, whereas visual predators are susceptible mainly when their prey is highly agile. This difference probably results from the superiority of the visual sensory system. However, while our study is an important step towards the mechanistic understanding of predator confusion, it also reveals how poor this understanding currently is.