Ecological periodic tables: in principle and practice

“Science is organized knowledge.” Immanuel Kant (1724–1804) Ecological periodic tables are an information organizing system with categorical habitat types as elements and predictably recurring (periodic) properties of a target biotic community, such as its relative species richness, abundance and biomass, as attributes. Ecological periodic tables are founded on the ecological tenet that habitats structure biotic communities and its corollary that habitats are templets for ecological strategies. They are a durable, open and flexible system that accommodates all operationally defined habitat types and biotic communities for which the periodicity of habitat usage patterns by a biotic community have been empirically substantiated. Discovering quantitative, periodic habitat usage patterns requires quantitative, representative, unbiased sampling of a biotic community across habitat types at ecologically relevant temporal and spatial scales. Like chemical periodic tables, the Linnaean system of classification and the Hertzsprung–Russell diagram in chemistry, biology and astronomy, respectively, ecological periodic tables are simple, easy to understand, exceptionally useful and they foster the expansion of scientific understanding, inquiry and theory.     

Scale dependency and functional response in moose habitat selection

Habitat selection can be influenced by the distribution of the habitat types in the landscape as well as net gain in visiting patches of resources, causing individual variation in habitat selection. Moreover, the hypothesis of functional response in habitat selection predicts that the degree of selection of a resource depends on its relative availability. We used radio-telemetry data from individual moose on an island off the coast of northern Norway to evaluate whether the selection of habitat types at the landscape scale differed from the choice of habitat types within the home range, and investigated the functional response in habitat selection by relating individual habitat selection to home range characteristics. At the landscape scale, moose selected for habitat types that provided both good forage and cover, with small differences between sex and age groups. At the home range scale, all individuals selected habitat types that were associated with cover and low human impact. Habitat selection was not modified by local moose density, but was related to home range size at both spatial scales. Larger home ranges contained larger proportions of non-preferred habitat types compared to smaller home ranges. At the home range scale, the selection for a habitat type decreased with its relative availability, indicating a functional response in habitat selection. This suggests that habitat selection is modified by home range size, which influences the availability of habitat types and shapes individual habitat selection patterns. Our results support previous suggestions that analyses of habitat or resource selection should follow a multi-scale approach. Both the relative availability of habitat types as well as individual variation in home range size should be accounted for in order to disentangle the complex mechanisms that contribute to shape patterns of resource selection in animals.     

Temperate migrants and resident bird species in Afro‐tropical savannahs show similar levels of ecological generalism

The specificity of an animal's habitat requirements will determine its ability to deal with anthropogenic climate and habitat change. Migratory birds are thought to be particularly vulnerable to such change, but theory predicts that they should be largely generalists. This prediction was tested with the aim of assessing whether migratory Palaearctic‐breeding birds wintering in the savannah biome of Africa are more or less generalist in their habitat use compared with taxonomically and ecologically similar Afro‐tropical resident species. The degree of specialization of these species groups to certain habitat characteristics was assessed and compared by calculating the relative occurrence of the species along habitat gradients, where wide occurrence indicates generalism and narrow occurrence indicates specialism. Palaearctic migrants as a group could not clearly be distinguished as generalists relative to Afro‐tropical residents with respect to habitat attributes. The only indication of greater flexibility in Palaearctic migrants was a significant tendency to use habitats over a wider latitudinal range. The results suggest that migrants are generalists, but not necessarily more generalist than taxonomically similar resident species that also occur over a wide range of habitat types within the savannah biome. The availability of specific habitat requirements on the wintering grounds in Africa is therefore unlikely to be a primary limiting factor for many Afro‐Palaearctic migratory bird species.     

The impact of habitat fragmentation on trophic interactions of the monophagous butterfly <Emphasis Type="Italic">Polyommatus coridon</Emphasis>

Theory predicts that habitat fragmentation, including reduced area and connectivity of suitable habitat, changes multitrophic interactions. Species at the bottom of trophic cascades (host plants) are expected to be less negatively affected than higher trophic levels, such as herbivores and their parasitoids or predators. Here we test this hypothesis regarding the effects of habitat area and connectivity in a trophic system with three levels: first with the population size of the larval food plant Hippocrepis comosa, next with the population density of the monophagous butterfly species Polyommatus coridon and finally with its larval parasitism rate. Our results show no evidence for negative effects of habitat fragmentation on the food plant or on parasitism rates, but population density of adult P. coridon was reduced with decreasing connectivity. We conclude that the highly specialized butterfly species is more affected by habitat fragmentation than its larval food plant because of its higher trophic position. However, the butterfly host species was also more affected than its parasitoids, presumably because of lower resource specialization of local parasitoids which also frequently occur in alternative hosts. Therefore, conservation efforts should focus first on the most specialized species of interaction networks and second on higher trophic levels.     

Isodars unveil asymmetric effects on habitat use caused by competition between two endangered species

In order for competing species to coexist, segregation on some ecological niche component is required and is often mediated by differential habitat use. When unequal competitors are involved, the dominant species tends to displace the subordinate one to its less preferred habitat. Here, we use habitat isodars, an approach which reflects evolutionary stable strategies of habitat selection, to evaluate whether interspecific competition between two competing species with distinct habitat preferences, the little bustard Tetrax tetrax and the great bustard Otis tarda, modulates their habitat use. Field data on these endangered species demonstrate that unequal competitors can coexist without completely segregating on their preferred habitats. The negatively sloped isodar of the subordinate little bustard unveils its competition with the dominant great bustard. Interference from great bustards in secondary cereal habitats reinforces use of preferred natural habitat by little bustards. Studies of density‐dependent habitat selection by a single‐species can thus aid in identifying the effects of competition on community composition, and guide the conservation of at‐risk species. Isodars, in particular, represent a promising method to gain clear knowledge on interspecific competition for species in which experimental manipulations are not feasible.     

Testing habitat copying in breeding habitat selection in a species adapted to variable environments

The habitat copying hypothesis states that animals use the reproductive performance of conspecifics to evaluate habitat quality and choose their future breeding site. We used data from Audouin's Gull Larus audouinii (1992–2003), a species adapted to unpredictable environments, to analyse subcolony (as patch) choice within a colony (small spatial scale). We also assessed the suitability of alternative hypotheses to the habitat copying hypothesis. The probability of subcolonies being reoccupied annually increased with their size (as number of nests), which suggests the existence of group adherence effects. Subcolony growth rate was related to its average reproductive success (or patch reproductive success) in the previous year: the higher the reproductive success in a colony, the higher the probability of growth the following year. However, this last result was obtained without considering the effect of colony size on the response variable because colony size is related to it. Therefore, results suggest at the population level that in this system habitat copying might either be one of the strategies used by the species in selecting its breeding habitat, or one of the possible strategies operating alone. The other strategies are group adherence mechanisms, and also the effect of conspecific attraction. At the individual level we failed to find evidence of habitat copying and only the previous success of an individual affected its fidelity to a subcolony. The importance of the lack of environmental predictability in the system is discussed, as predictability is a prerequisite of habitat copying.     

Fauna habitat modelling and mapping: A review and case study in the Lower Hunter Central Coast region of NSW

Abstract Habitat models are now broadly used in conservation planning on public lands. If implemented correctly, habitat modelling is a transparent and repeatable technique for describing and mapping biodiversity values, and its application in peri‐urban and agricultural landscape planning is likely to expand rapidly. Conservation planning in such landscapes must be robust to the scrutiny that arises when biodiversity constraints are placed on developers and private landholders. A standardized modelling and model evaluation method based on widely accepted techniques will improve the robustness of conservation plans. We review current habitat modelling and model evaluation methods and provide a habitat modelling case study in the New South Wales central coast region that we hope will serve as a methodological template for conservation planners. We make recommendations on modelling methods that are appropriate when presence‐absence and presence‐only survey data are available and provide methodological details and a website with data and training material for modellers. Our aim is to provide practical guidelines that preserve methodological rigour and result in defendable habitat models and maps. The case study was undertaken in a rapidly developing area with substantial biodiversity values under urbanization pressure. Habitat maps for seven priority fauna species were developed using logistic regression models of species‐habitat relationships and a bootstrapping methodology was used to evaluate model predictions. The modelled species were the koala, tiger quoll, squirrel glider, yellow‐bellied glider, masked owl, powerful owl and sooty owl. Models ranked sites adequately in terms of habitat suitability and provided predictions of sufficient reliability for the purpose of identifying preliminary conservation priority areas. However, they are subject to multiple uncertainties and should not be viewed as a completely accurate representation of the distribution of species habitat. We recommend the use of model prediction in an adaptive framework whereby models are iteratively updated and refined as new data become available.     

Exploitation ecosystems in heterogeneous habitat complexes II: Impact of small-scale heterogeneity on predator-prey dynamics

Summary The model of exploitation ecosystems was re-analysed, assuming that habitat patches are so small that they form only parts of the home range of an individual predator. For habitat complexes where productive patches abound, the results suggested that predation will strongly spill over from productive patches, which set the tune for population dynamics within the whole landscape, to barren ones. This result conforms to the one obtained by T. Oksanen by assuming despotic habitat choice and essentially larger patch sizes. For habitat complexes heavily dominated by the barren habitat, spillover predation was predicted to be weak, as was the case in her large patch model. Unlike in her analysis, however, predation pressure was substantially reduced also within the productive habitat. In habitat complexes where patches are so small that they are exploited in a fine-grained manner, predation pressure was always found to be more intense in the barren habitat, contrary to the predictions of the original model of exploitation ecosystems. This analysis thus suggests that their model is applicable mainly on the landscape level. On the level of individual habitats, the applicability of their results depends on the habitat configuration (at its best for the prevailing habitat of the landscape and for moderate-sized patches of an essentially more productive habitat) and generally decreases with decreasing patch sizes.     

Old-growth forest is what giant pandas really need

Giant pandas (Ailuropoda melanoleuca) are an iconic conservation species, but despite significant research effort, do we understand what they really need? Estimating and mapping suitable habitat play a critical role in conservation planning and policy. But if assumptions about ecological needs are wrong, maps with misidentified suitable habitat will misguide conservation action. Here, we use an information-theoretic approach to analyse the largest, landscape-level dataset on panda habitat use to date, and challenge the prevailing wisdom about panda habitat needs. We show that pandas are associated with old-growth forest more than with any ecological variable other than bamboo. Other factors traditionally used in panda habitat models, such as topographic slope, are less important. We suggest that our findings are disparate from previous research in part because our research was conducted over a larger ecological scale than previous research conducted over more circumscribed areas within individual reserves. Thus, extrapolating from habitat studies on small scales to conservation planning on large scales may entail some risk. As the Chinese government is considering the renewal of its logging ban, it should take heed of the panda''s dependency on old growth.     

HABITAT SELECTION AS AN EVOLUTIONARY GAME

Under habitat selection, mobile foragers may not only possess behavioral flexibility that allows them to utilize habitats selectively or opportunistically, but they may also possess heritable traits that influence their performance within each habitat. A game theoretic model is developed that investigates this evolutionary dimension of habitat selection. The model follows that of Rosenzweig (1987b) and considers a patchy environment containing two distinct habitat types. Behaviorally, foragers may be selective or opportunistic; morphologically, foragers possess traits that represent a trade-off between performance in the two habitat types. Depending on the environment's structure, one of three types of communities emerges as the ESS: (1) a single generalist species that behaves opportunistically, (2) two species that are extreme specialists on habitat 1 and 2, respectively; behaviorally, these species are selective on their respective habitat types, and (3) one generalist species that behaves opportunistically and one specialist species that behaves selectively on its preferred habitat. Community (1) emerges when habitat selection is costly, community (2) emerges when habitat selection is cost-free, and community (3) emerges when the relative abundances or productivities of the two habitat types are lopsided.     

Metapopulation extinction risk: Dispersal’s duplicity

Metapopulation extinction risk is the probability that all local populations are simultaneously extinct during a fixed time frame. Dispersal may reduce a metapopulation’s extinction risk by raising its average per-capita growth rate. By contrast, dispersal may raise a metapopulation’s extinction risk by reducing its average population density. Which effect prevails is controlled by habitat fragmentation. Dispersal in mildly fragmented habitat reduces a metapopulation’s extinction risk by raising its average per-capita growth rate without causing any appreciable drop in its average population density. By contrast, dispersal in severely fragmented habitat raises a metapopulation’s extinction risk because the rise in its average per-capita growth rate is more than offset by the decline in its average population density. The metapopulation model used here shows several other interesting phenomena. Dispersal in sufficiently fragmented habitat reduces a metapopulation’s extinction risk to that of a constant environment. Dispersal between habitat fragments reduces a metapopulation’s extinction risk insofar as local environments are asynchronous. Grouped dispersal raises the effective habitat fragmentation level. Dispersal search barriers raise metapopulation extinction risk. Nonuniform dispersal may reduce the effective fraction of suitable habitat fragments below the extinction threshold. Nonuniform dispersal may make demographic stochasticity a more potent metapopulation extinction force than environmental stochasticity.     

Surrogate Habitats Demonstrate the Invasion Potential of the African Pugnacious Ant

Many ant species are highly invasive and are a significant component of disturbed ecosystems. They can have a major suppressive effect upon indigenous invertebrates, including other ants. Despite overwhelming circumstantial evidence for the ecological resourcefulness of many ants, there appears to be no experimental evidence illustrating the habitat breadth of a potentially invasive ant species. We demonstrate here that a particularly opportunistic and locally dominant ant Anoplolepis custodiens, which is a major indigenous African pest, overrides habitat structure to maintain its population level. We compared A. custodiens activity, morphology, foraging behaviour and ant species diversity in artificially established surrogate habitats (cover crops) in a vineyard containing an ample food resource in the form of the honeydew-producing mealybug Planococcus ficus. These cover crops were chosen so as to create highly altered habitats. The ant's ability to overcome these potentially suppressive habitat conditions hinged on its tight mutualism with the mealybug, and on its chasing away mealybug parasitoids. This ant species is predicted to be a latent invasive beyond Africa. It is unlikely to be impeded once it has established a foothold in a variety of novel habitats. It could locally invade to obtain food resources in a wide range of habitat types. Furthermore, in agricultural systems, cover crops are unlikely to control such an ant. Potential invasives such as this ant should be flagged as important quarantine suspects.     

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.     

Viability analyses with habitat-based metapopulation models

Population viability analysis (PVA) models incorporate spatial dynamics in different ways. At one extreme are the occupancy models that are based on the number of occupied populations. The simplest occupancy models ignore the location of populations. At the other extreme are individual-based models, which describe the spatial structure with the location of each individual in the population, or the location of territories or home ranges. In between these are spatially structured metapopulation models that describe the dynamics of each population with structured demographic models and incorporate spatial dynamics by modeling dispersal and temporal correlation among populations. Both dispersal and correlation between each pair of populations depend on the location of the populations, making these models spatially structured. In this article, I describe a method that expands spatially structured metapopulation models by incorporating information about habitat relationships of the species and the characteristics of the landscape in which the metapopulation exists. This method uses a habitat suitability map to determine the spatial structure of the metapopulation, including the number, size, and location of habitat patches in which subpopulations of the metapopulation live. The habitat suitability map can be calculated in a number of different ways, including statistical analyses (such as logistic regression) that find the relationship between the occurrence (or, density) of the species and independent variables which describe its habitat requirements. The habitat suitability map is then used to calculate the spatial structure of the metapopulation, based on species-specific characteristics such as the home range size, dispersal distance, and minimum habitat suitability for reproduction. Received: April 1, 1999 / Accepted: October 29, 1999     

The relationship between population density, habitat position and habitat breadth within a neotropical forest bird community

We examined the relationship between local abundance, habitat position and habitat breadth across bird species in a large Atlantic forest reserve in Brazil. This appears to be the first such study for any rainforest taxon. Habitat position for a species was its mean foraging height, along with the mean scores on three principal habitat axes for census stations at which it was recorded. Habitat breadth was the standard deviation of recorded foraging heights and the standard deviations of "positive" station scores on the habitat axes. We also examined differences in habitat position and breadth between endemic and wide-ranging taxa and amongst dietary groups. Amongst 31 species for which density estimation was possible, there were no correlations between local abundance and breadth of habitat use on any of the habitat axes. Breadth of habitat used did not vary with degree of endemism, but herbivores used a greater breadth of habitats on the axis describing canopy closure than did omnivores. Habitat position did not vary with endemic status, but herbivores preferred higher-biomass habitats than faunivores, and higher foraging heights than either faunivores or omnivores. Local abundance was linked weakly to habitat position with commoner species tending to forage in the lower strata of open-canopied areas. The 31 most commonly recorded species tended to occupy "middle-range" habitat positions, while 28 rarer species occupied habitats toward one or other end of the vegetation axes. These results suggest an association between the local abundance of a species and its habitat position, and especially its preference for common or mid-range habitats, rather than with its ability to utilise a wide range of habitats.     

Reconstructing changes in the genotype,phenotype, and climatic niche of an introduced species

An introduced species must contend with enormous environmental variation in its introduced range. In this study, we use niche models and ordination analyses to reconstruct changes in genotype, phenotype, and climatic niche of Johnsongrass Sorghum halepense, which is regarded as one of the world's most threatening invasive plants. In the United States, Johnsongrass has rapidly evolved within‐ and among‐population genetic diversity; our results show that genetic differentiation in expanding Johnsongrass populations has resulted in phenotypic variation that is consistent with habitat and climatic variation encountered during its expansion. Moreover, Johnsongrass expanded from agricultural to non‐agricultural habitat, and now, despite occupying overlapping ranges, extant agricultural and non‐agricultural populations are genetically and phenotypically distinct and manifest different plastic responses when encountering environmental variation. Non‐agricultural accessions are broadly distributed in climatic and geographic space and their fitness traits demonstrate plastic responses to common garden conditions that are consistent with local specialization. In contrast, agricultural accessions demonstrate ‘general purpose’ plastic responses and have more restricted climatic niches and geographic distributions. They also grow much larger than non‐agricultural accessions. If these differences are adaptive, our results suggest that adaptation to local habitat variation plays a crucial role in the ecology of this invader. Further, its success relates to its ability to succeed on dual fronts, by responding simultaneously to habitat and climate variability and by capitalizing on differential responses to these factors during its range expansion.     

Resource Abundance and Invasiveness: A Simple Model

The 'invasiveness' of an alien species depends partly on its ability to become abundant and widespread in its new environment. While competitiveness may be an important component of this ability, so too is the abundance of resource or habitat. First, the local carrying capacity will depend on the local favourability of the habitat, hence the global density will depend on how widespread the habitat is. Second, and more subtly, the local density will also be affected by the global extent of favourable habitat, because of losses occasioned by dispersal when the population redistributes; these losses should be fewer the greater the contiguous area of favourable habitat or the more patches of such habitat across the landscape. Here we describe a model which demonstrates how habitat availability affects an invading speciesèquilibrium abundance, hence its invasiveness. The model shows that local density is likely to be an increasing function of global habitat abundance, and global density to be a non-linear, concave-up function of global habitat abundance. Examples are given to support the model's predictions, taken largely from alien species in New Zealand.     

Habitat selection and population trends in terrestrial bird species of Robinson Crusoe Island: habitat generalists versus forest specialists

Habitat loss and degradation on oceanic islands are key processes leading to population decline of endemic birds and facilitating the establishment of invasive bird species. In this study, carried out in the Robinson Crusoe Island, we assessed density and habitat selection of terrestrial bird species, including juan fernandez firecrown and juan fernandez tit-tyrant, two endemics, as well as green-backed firecrown and austral thrush, which apparently originate from the mainland. Results show that perturbed habitats contained a low density of the endemic species whereas the mainland species were significantly more abundant in perturbed scrub habitats. Bird species show different habitat selection patterns, with endemics selecting for native forest and mainland species selecting for perturbed habitats, or using them at random. Bird species experienced temporal trends in their overall population sizes, with the endemic tit-tyrant suffering a significant decline in its population size of about 63% between 1994 and 2009. Only mainland species exhibited temporal changes in habitat use, significantly reducing their densities in the preferred scrub habitats, possibly as a response to decreased habitat quality. Thrushes apparently were able to compensate the population decrease in one non native habitat type by using native forests, a habitat giving them the opportunity of preying on nests of endemic species. We conclude that endemic bird species behave as specialists whereas the mainland species must be treated as invasive generalists on Robinson Crusoe Island.     

Linking Habitat Restoration to Meaningful Units of Animal Demography

To restore habitat is to restore the functional aspects of a place where one or more species are intended to live. However, habitat restored with the right structural elements can still fail to support the species due to insufficient space and spatial contiguity (e.g., movement corridors) with nearby habitat. The spatial extent of habitat determines its capacity to support various demographic units, such as a breeding pair of individuals, a true population, or a metapopulation. In scatter‐plots of published density estimates and their corresponding study area sizes, I found consistent patterns that appeared to represent spatial scale domains, in which subpopulations, populations, and multiple populations exist for each species. These spatial scale domains can help guide habitat restoration by indicating the minimum area needed to support a population. The effectiveness of restored habitat, considered alone or in combination with existing habitat, should be judged by the habitat's capacity to sustain a functioning population, which includes the population's membership in a metapopulation. Additionally, the evidence increasingly demonstrates that habitat must include sufficient space for the population(s) to periodically relocate as resources are depleted locally. The spatial extent of proposed habitat restoration can now be linked to the likely demographic unit of the species that can be supported there. I provide two examples of proposed habitat conservation and their likely effectiveness based on their spatial areas.     

Habitat requirements of Lycaena dispar batavus and implications for re-establishment in England

This paper reports on studies of the habitat requirements of Lycaena dispar batavus in its largest remaining site, the Weerribben National Park in the Netherlands. Observations and experiments were carried out to assess requirements for male territories, egg-laying sites and larval survival. The management of the fenland was also assessed on a landscape scale. Male territories were all situated in summer cut fen meadow. Eggs were laid on Rumex hydrolapathum in a range of habitat situations, although a preference was shown for plants on fen edges. Larval survival was highest on plants on watersides and fen edges, with no survival on plants in summer cut fen meadow. The findings are discussed in relation to whether the Broadland area in eastern England is suitable as a re-establishment site. Although many elements of the habitat mosaic are present, at the landscape scale there is insufficient open fenland and areas suitable for male territories are probably too isolated; however, a fen restoration strategy is in place which may restore suitable habitat and make re-establishment feasible in the future.