Can settlement in natal-like habitat explain maladaptive habitat selection?

The study of habitat selection has long been influenced by the ideal free model, which maintains that young adults settle in habitat according to its inherent quality and the density of conspecifics within it. The model has gained support in recent years from the finding that conspecifics produce cues inadvertently that help prebreeders locate good habitat. Yet abundant evidence shows that animals often fail to occupy habitats that ecologists have identified as those of highest quality, leading to the conclusion that young animals settle on breeding spaces by means not widely understood. Here, we report that a phenomenon virtually unknown in nature, natal habitat preference induction (NHPI), is a strong predictor of territory settlement in both male and female common loons (Gavia immer). NHPI causes young animals to settle on natal-like breeding spaces, but not necessarily those that maximize reproductive success. If widespread, NHPI might explain apparently maladaptive habitat settlement.     

Correlation and studies of habitat selection: problem,red herring or opportunity?

With the advent of new technologies, animal locations are being collected at ever finer spatio-temporal scales. We review analytical methods for dealing with correlated data in the context of resource selection, including post hoc variance inflation techniques, ‘two-stage’ approaches based on models fit to each individual, generalized estimating equations and hierarchical mixed-effects models. These methods are applicable to a wide range of correlated data problems, but can be difficult to apply and remain especially challenging for use–availability sampling designs because the correlation structure for combinations of used and available points are not likely to follow common parametric forms. We also review emerging approaches to studying habitat selection that use fine-scale temporal data to arrive at biologically based definitions of available habitat, while naturally accounting for autocorrelation by modelling animal movement between telemetry locations. Sophisticated analyses that explicitly model correlation rather than consider it a nuisance, like mixed effects and state-space models, offer potentially novel insights into the process of resource selection, but additional work is needed to make them more generally applicable to large datasets based on the use–availability designs. Until then, variance inflation techniques and two-stage approaches should offer pragmatic and flexible approaches to modelling correlated data.     

Evolution of protein inhibitors of serine proteinases: Positive Darwinian selection or compositional effects?

Summary In at least two instances involving serine proteinase inhibitors it has been shown that functionally important sites evolve faster and exhibit more interspecific variability than functionally neutral sites. Because these phenomena are difficult to reconcile with the neutral theory of molecular evolution, it has been suggested that the accelerated rate of amino acid substitution at the reactive sites is brought about by positive Darwinian selection. We show that differences in the amino acid composition in the different regions of proteinase inhibitors can account for the differences in the rates of amino acid substitution. By using an index of protein mutability [D. Graur (1985) J Mol Evol 2253–62], we show that the amino acid composition of the reactive center in the ovomucoids andSpi-2 gene products is such that, regardless of function, they are expected to evolve more rapidly than any other polypeptide for which the rate of substitution is known. In addition, the reactive region in theSpi-2 proteins is shown to be free of compositional constraint. Positive Darwinian selection need not be invoked at the present time in these cases.     

Can models of density-dependent habitat selection be applied for trap-building predators?

Density-dependent habitat selection theory was mainly tested on active foragers and therefore its applicability to trap-building predators is poorly understood. The high sensitivity of trap-building predators to changes in their physical environment, combined with their limited movement capability, can lead to habitat specialization, reducing their utilization of alternative habitats. We studied density-dependent habitat selection in two pit-building antlions, differing in their habitat utilization spectrum. The habitat generalist, Myrmeleon hyalinus, inhabits and performs equally well in both sand- and loess-derived soils, although preferring the former, more productive coarse-grained soils. In contrast, the habitat specialist, Cueta lineosa, only inhabits fine-grained soils such as loess, while showing reduced foraging performance in coarse-grained sandy soils. We allowed larvae to select between these two soils, while manipulating con-specific density and initial stocking position. Irrespective of the initial stocking position, the number of M. hyalinus pits in the sand was positively correlated with the number of con-specific pits in the loess, implying that this species is a density-dependent habitat selector. Furthermore, these patterns of density-dependence were consistent with the expectations of ideal pre-emptive distribution (i.e., strong non-linearity in the distribution of antlions between soils with increased total density), suggesting that interference competition largely dictates habitat selection in this species. In contrast, the habitat specialist showed constant habitat selectivity, as neither con-specific density nor initial stocking positions influenced its habitat preference. Although mainly tested on active foragers, habitat selection theory can be applicable for trap-building predators, demonstrating how mechanisms operating at the individual level influence spatial distribution patterns.     

Evolution of genome size: multilevel selection, mutation bias or dynamical chaos?

In the past two years, new data on conceptual aspects of the evolution of eukaryotic genome size have appeared, including the adaptivity of genome enlargement, the mechanisms of genome size change and the relation of genome size to organismal complexity. New data on the hypotheses of "selfish DNA" and "mutational equilibrium" have been recently obtained. A relationship is emerging between the intragenomic distribution of noncoding DNA and differential gene expression, which suggests that noncoding DNA is involved in epigenetic organization of the genome and organismal complexity. The standpoint of dynamical chaos, which integrates multilevel selection and mutation biases, may provide a framework for studying the evolution of genome size.     

Use of hunters in wildlife surveys: does hunter and forest grouse habitat selection coincide?

Wildlife monitoring performed by volunteer personnel may suffer from bias with regard to their habitat use. Such errors can lead to erroneous population estimates, evidently influencing both management programmes and research that are based on the monitoring. We used a dataset on hunters’ habitat use in forest while searching for black grouse (Tetrao tetrix) and capercaillie (Tetrao urogallus) to test whether hunters’ utilisation of the habitat was independent of that of grouse or if it corresponded to the grouse habitat preference. Twenty volunteer hunters with dogs registered their tracks and all observations of capercaillie and black grouse in Østfold County, Norway, during August 2003 and 2004. We performed an ecological niche factor analysis (ENFA) and a K-select analysis with respect to hunters’ selection for habitat, described with ecogeographical variables related to forest stand characteristics, as well as the habitat preference of the observed grouse, conditional on the habitat utilisation of the hunters. Individual ENFA on the hunter's tracks revealed large variation in the habitat preferences of the hunters. The K-select indicated few overall patterns in the habitat characteristics of grouse observations, conditional on the hunters selected habitat. Accordingly, the results indicate that hunters’ observation of grouse prior to the hunt may give indicators of changes in grouse density unbiased by habitat preference due to the large between-hunter variation in habitat preference, given that a sufficient number of hunters is used. This suggests that such monitoring programmes can provide information about fluctuations in grouse population sizes valuable for both the management and research of forest grouse species.     

Background sampling for multi-scale ensemble habitat selection modeling: Does the number of points matter?

Ensemble habitat selection modeling is becoming a popular approach among ecologists to answer different questions. Since we are still in the early stages of development and application of ensemble modeling, there remain many questions regarding performance and parameterization. One important gap, which this paper addresses, is how the number of background points used to train models influences the performance of the ensemble model. We used an empirical presence-only dataset and three different selections of background points to train scale-optimized habitat selection models using six modeling algorithms (GLM, GAM, MARS, ANN, Random Forest, and MaxEnt). We tested four ensemble models using different combinations of the component models: (a) equal numbers of background points and presences, (b) background points equaled ten times the number of presences, (c) 10,000 background points, and (d) optimized background points for each component model. Among regression-based approaches, MARS performed best when built with 10,000 background points. Among machine learning models, RF performed the best when built with equal presences and background points. Among the four ensemble models, AUC indicated that the best performing model was the ensemble with each component model including the optimized number of background points, while TSS increased as the number of background points models increased. We found that an ensemble of models, each trained with an optimal number of background points, outperformed ensembles of models trained with the same number of background points, although differences in performance were slight. When using a single modeling method, RF with equal number of presences and background points can perform better than an ensemble model, but the performance fluctuates when the number of background points is not properly selected. On the other hand, ensemble modeling provides consistently high accuracy regardless of background point sampling approach. Further, optimizing the number of background points for each component model within an ensemble model can provide the best model improvement. We suggest evaluating more models across multiple species to investigate how background point selection might affect ensemble models in different scenarios.     

Tubeworm succession at hydrothermal vents: use of biogenic cues to reduce habitat selection error?

Species colonizing new deep-sea hydrothermal vents along the East Pacific Rise show a distinct successional sequence: pioneer assemblages dominated by the vestimentiferan tubeworm Tevnia jerichonana being subsequently invaded by another vestimentiferan Riftia pachyptila, and eventually the mussel Bathymodiolus thermophilus. Using a manipulative approach modified from shallow-water ecological studies, we test three alternative hypotheses to explain the initial colonization by T. jerichonana and its subsequent replacement by R. pachyptila. We show that R. pachyptila and another vestimentiferan, Oasisia alvinae, colonized new surfaces only if the surfaces also were colonized by T. jerichonana. This pattern does not appear to be due to restricted habitat tolerances or inferior dispersal capabilities of R. pachyptila and O. alvinae, and we argue the alternative explanation that T. jerichonana facilitates the settlement of the other two species and is eventually outcompeted by R. pachyptila. Unlike the classic model of community succession, in which facilitating species promote their own demise by modifying the environment to make it more hospitable for competitors, we suggest that T. jerichonana may produce a chemical substance that induces settlement of these competitors. This process of selecting habitat based on biogenic cues may be especially adaptive and widespread among later-successional species that occupy a physically variable and unpredictable environment. In these cases, the presence of weedy species implies some integrated period of environmental suitability, whereas an instantaneous assessment of physical habitat conditions, such as water temperature for vent tubeworms, provides a poorer predictor of long-term habitat suitability. Received: 13 July 1999 / Accepted: 2 November 1999     

Evolution of the ungulate dewlap: thermoregulation rather than sexual selection or predator deterrence?

Background

Dewlaps are iconic features of several ungulate species and, although a role in signalling has been postulated, their function remains largely unexplored. We recently failed to find any age-independent link between dewlap size and social status in the common eland (Tragelaphus oryx), pointing to the possibility that sexual selection may not be the primary cause of dewlap evolution in ungulates. Here I use a two-pronged approach to test hypotheses on the function of ungulate dewlaps: an interspecific comparative analysis of bovids and deer, and an intraspecific study of eland antelopes in the wild.

Results

Across species, the presence of dewlaps in males was not found to be associated with sexual size dimorphism, a commonly used measure of the intensity of sexual selection. The presence of dewlaps was, however, linked to very large male body size (>400 kg), which agrees with a thermoregulatory function as lower surface/volume-ratio counteracts heat dissipation in large-bodied species. In eland antelopes, large dewlap size was associated with higher, rather than lower, incidence of claw-marks (independently of age), a result which speaks against the dewlap as a predator deterrent and rather indicates a predation cost of the structure.

Conclusion

The findings suggest that, although an additional function in communication should not be ruled out, the dewlap of ungulates may contrast with that of lizards and birds in thermoregulation being a primary function.

     

Eurasian badger habitat selection in Mediterranean environments: Does scale really matter?

It is widely believed that spatial scale affects habitat selection, and should influence management options, especially for species with wide geographic distribution or large territories. Eurasian badger habitat selection has been well studied throughout most of its European distribution range, but never at multiple spatial scales. We used compositional analysis to assess habitat selection of Eurasian badgers in southern Portugal at four spatial scales (1, 4, 25, and 100 km²). We assessed habitat use from setts, latrines and footprints presence, and road kills. Oak woodlands with understorey were selected at all scales, being the most preferred habitat at 3 scales (1, 4, and 100 km²). Pastures were most selected at the scale of the 25 km² cell, but their use was not significantly different from oak woodland with understorey. Shrubs and pastures were also secondly important at the majority of scales. Contrary to findings at northern latitudes, deciduous forests decreased in importance as cell size increased. In the highly humanized and fragmented landscape of southern Portugal, Eurasian badgers are selecting the matrix of oak woodlands interspersed with patches of pastures, shrubs and riparian vegetation. In these oak woodlands, scale does not have a marked effect. Management for badgers should provide, for at least, 30% of oak woodland cover at all scales. Our study illustrates the across-scale importance of maintaining the historically human altered, sustainable and unique landscape and land use system – the montado.     

Prey habitat selection under shared predation: tradeoffs between risk and competition?

An individual's choice of habitat should optimize amongst conflicting demands in a way that maximizes its fitness. Habitat selection by one species will often be influenced by presence and abundance of competitors that interact directly and indirectly with each other (such as through shared predators). The optimal habitat choice will thus depend on competition for resources by other species that can also modify predation risk. It may be possible to disentangle these two effects with careful analysis of density‐dependent habitat selection by a focal prey species. We tested this conjecture by calculating habitat isodars (graphs of density assuming ideal habitat selection) of chital deer living in two adjoining dry‐forest habitats in Gir National Park and Sanctuary, western India. The habitats differed only in presence (Sanctuary) and absence (National Park) of domestic prey (cattle and buffalo). Both species are preyed on by Asiatic lions. The habitat isodar revealed at low densities, that chital live in small groups and prefer habitat co‐occupied by livestock that reduce food resources, but also reduce predation risk. At higher densities, chital form larger groups and switch their preference toward risky habitat without livestock. The switch in chital habitat use is consistent with theories predicting that prey species should trade off safety in favor of food as population density increases.     

Can pinewoods provide habitat for a deciduous forest specialist? A two-scale approach to the habitat selection of Bechstein's bat

Populations of Myotis bechsteinii in Sierra de Cazorla, Segura y Las Villas Natural Park (southwestern Spain) survive in a large, old-growth coniferous woodland, which is in contrast with known ecological preferences of the species. We tracked ten lactating females and studied patterns of habitat selection by Bechstein's bat (Myotis bechsteinii) at two spatial scales: macrohabitat and microhabitat, regarding forest essence (coniferous vs. deciduous). The tracked individuals always foraged within forested areas and did not use areas devoid of trees. At the macrohabitat level, no positive selection of deciduous stands was apparent, suggesting selection studies of coarse resolution may not be able to capture subtle selection patterns. At the microhabitat level Myotis bechsteinii selected deciduous patches within the coniferous matrix, therefore, our results corroborate the perception of this species as dependent of services provided by deciduous woodlands. Larger foraging home ranges and commuting distances as compared with other Mediterranean localities suggest that our studied population inhabits a marginal or suboptimal environment in terms of habitat quality. We argue that this population of Bechstein's bat has survived as a relict one probably as a consequence of fragmentation and transformation of deciduous forest ecosystems in the Mediterranean range.     

Testing for evidence of inefficient selection in bdelloid rotifers: do sample size and habitat differences matter?

Preliminary evidence on inefficient selection has indicated that asexual (=parthenogenetic) animals suffer from a greater accumulation of deleterious mutations compared with their closest sexual sister taxa. However, previous work on bdelloid rotifers did not completely rule out the confounding effects of sample size and habitat differences. Here, we present further evidence of inefficient selection against deleterious mutations in the bdelloid rotifers in comparison with their closest clade, the monogononts, by taking account of larger samples. However, the analysis of samples from both clades co-occurring in the same location seems to contradict the hypothesis. Both groups show evidence of the accumulation of deleterious mutations in mitochondrial DNA, but further population sampling and inclusion of additional genes is needed to resolve this issue.     

Influence of fire salamander larvae on among-pool distribution of mosquito egg rafts: oviposition habitat selection or egg raft predation?

Many amphibian populations are in decline worldwide. Surprisingly, few studies have examined how such declines may benefit mosquitoes. Amphibian larvae may compete with and prey upon mosquito larvae, and may alter oviposition habitat selection (OHS) of mosquito adults. However, often overlooked, observed among-pool egg distributions attributed to OHS may additionally or alternatively be explained by egg predation. Temporary pools of mountainous areas of the Mediterranean serve as larval habitat for both the mosquito, Culiseta longiareolata, and the salamander, Salamandra infraimmaculata. We found Culiseta larvae and egg rafts to be highly vulnerable to predation by pre-metamorphosing Salamandra larvae, but not to metamorphosing ones. In outdoor mesocosm experiments, oviposition avoidance by Culiseta females in response to caged Salamandra was not demonstrated regardless of salamander developmental stage. Egg raft abundance was significantly reduced in free-roaming, pre-metamorphosing Salamandra but not by metamorphosing ones. Thus, Salamandra larvae may have little deterrence on Culiseta oviposition. Instead, fewer egg rafts are attributed largely to egg predation. This study highlights the importance of egg raft predation in addition to OHS when interpreting the influence of predators on prey egg distributions. It also highlights that a cost of declining amphibian populations is their reduced impacts on mosquito populations.     

Evolution of cooperation: two for one?

How can cooperation thrive in a selfish world? Recent evolution experiments show how bacteria themselves can generate conditions that make cooperation a winning strategy. At least in the short term.     

Evolution of biodiversity: Hyperbola or exponent?

Two models of the evolution of biodiversity during the Phanerozoic are critically analyzed.     

Prospecting in the collared flycatcher: gathering public information for future breeding habitat selection?

Visiting breeding patches can allow prospecting individuals to gather information on local patch quality to make optimal decisions about selecting a breeding habitat in the following year. This prospecting behaviour has been described in many bird species. However, the nature of the information gathered by prospectors often remains unknown. We collected data on prospecting behaviour in the collared flycatcher, Ficedula albicollis, a small hole-nesting passerine bird, to investigate whether prospectors could gather information on their conspecifics' reproductive success, that is, ‘public information’. If they could, they would be expected to inspect conspecific nests, prospect at the time when public information is reliable, and be attracted to, and spend time at, successful sites. Prospecting at conspecific nests was frequent. Prospectors were mainly males, and prospecting activity closely matched the peak of nesting activity. The probability of observing a prospector at a nest increased with parental activity, measured by feeding rate and vigilance behaviour, but did not depend on direct measures of success of the inspected nest (nestling number and condition), or on female characteristics. Because feeding rate and vigilance behaviour were predictors of breeding success at fledging, prospectors were attracted to the most successful nests when cueing on conspicuous parental activity. Therefore, prospectors could gather accurate information on local conspecific reproductive success that may be used for breeding patch choice in the following year. We discuss alternative explanations for our results, and the need to test experimentally whether prospectors gather public information. We also discuss the role of breeding constraints on, and the origin of sex differences in, prospecting.