Habitat complexity, spatial interference, and "minimum risk distribution": a framework for population stability

In the past century, the debate over whether or not density-dependent factors regulate populations has generally focused on changes in mean population density, ignoring the spatial variance around the mean as unimportant noise. In an attempt to provide a different framework for understanding population dynamics based on individual fitness, this paper discusses the crucial role of spatial variability itself on the stability of insect populations. The advantages of this method are the following: (1) it is founded on evolutionary principles rather than post hoc assumptions; (2) it erects hypotheses that can be tested; and (3) it links disparate ecological schools, including spatial dynamics, behavioral ecology, preference-performance, and plant apparency into an overall framework. At the core of this framework, habitat complexity governs insect spatial variance. which in turn determines population stability. First, the minimum risk distribution (MRD) is defined as the spatial distribution of individuals that results in the minimum number of premature deaths in a population given the distribution of mortality risk in the habitat (and, therefore, leading to maximized population growth). The greater the divergence of actual spatial patterns of individuals from the MRD, the greater the reduction of population growth and size from high, unstable levels. Then, based on extensive data from 29 populations of the processionary caterpillar, Ochrogaster lunifer, four steps are used to test the effect of habitat interference on population growth rates. (1) The costs (increasing the risk of scramble competition) and benefits (decreasing the risk of inverse density-dependent predation) of egg and larval aggregation are quantified. (2) These costs and benefits, along with the distribution of resources, are used to construct the MRD for each habitat. (3) The MRD is used as a benchmark against which the actual spatial pattern of individuals is compared. The degree of divergence of the actual spatial pattern from the MRD is quantified for each of the 29 habitats. (4) Finally, indices of habitat complexity are used to provide highly accurate predictions of spatial divergence from the MRD, showing that habitat interference reduces population growth rates from high, unstable levels. The reason for the divergence appears to be that high levels of background vegetation (vegetation other than host plants) interfere with female host-searching behavior. This leads to a spatial distribution of egg batches with high mortality risk, and therefore lower population growth. Knowledge of the MRD in other species should be a highly effective means of predicting trends in population dynamics. Species with high divergence between their actual spatial distribution and their MRD may display relatively stable dynamics at low population levels. In contrast, species with low divergence should experience high levels of intragenerational population growth leading to frequent habitat-wide outbreaks and unstable dynamics in the long term. Six hypotheses, erected under the framework of spatial interference, are discussed, and future tests are suggested.

Determination of the hemispheric character (i.e. southern or northern latitudinal position) of equatorial ionosonde stations as deduced from long-term recordings of the minimum height of the F-region (h ' F)

The virtual (or minimum) height of the F-region (h'F), recorded over a number of solar cycles for I I equatorial and mid-latitude ionosonde stations, was used to deduce the hemispheric (i.e. southern or northern hemisphere) character of equatorial stations. The semi-annual median monthly height (h'F) variations consist of two components: major local summer maximum and winter sub-maximum (about 5 percent of the summer maximum). This hemispheric pattern was most consistently observed for equatorial stations (within 5degrees of the geomagnetic equator) in a period centred on the local midnight (21-03 LT) but was also present, to a lesser extent, at mid-latitude stations and at other time intervals. It is evident that the physical parameter h'F defines the hemispheric character of an equatorial station which has different (sometimes opposite) geographic and geomagnetic latitudes. There is a sharp transition in the latitudinal character of the stations on both sides of the equator leading to hypothesis that the equal maxima in h'F in December and June solstices are observed at a near-equator position labelled as ionosonde deduced equator (IDE). Such a signature was observed for an American equatorial (both geographic and geomagnetic) station Talara (Peru) which is an experimental support of the hypothesis. The IDE can be another useful parameter characterising the equatorial ionosphere. This finding reveals a new application of the standard ionosonde data in defining the geophysical character of equatorial stations, being an important contribution to space climatology. (C) 2002 Elsevier Science Ltd. All rights reserved.

Comparative influence of forest management and habitat structural factors on the abundances of hollow-nesting bird species in subtropical Australian eucalypt forest

We examined the impact of single-tree selective logging and fuel reduction bums on the abundance of hollow-nesting bird species at a regional scale in southeastern Queensland, Australia. Data were collected on species abundance and habitat structure of dry sclerophyll production forest at 36 sites with known logging and fire histories. Sixteen bird species were recorded with most being resident, territorial, obligate hollow nesters that used hollows that were either small (18 cm diameter). Species densities were typically low, but combinations of two forest management and three habitat structural variables influenced the abundances of eight bird species in different and sometimes conflicting ways. The results suggest that habitat tree management for biodiversity in production forests cannot depend upon habitat structural characteristics alone. Management histories appear to have independent influence (on some bird species) that are distinguishable from their impacts on habitat structure per se. Rather than managing to maximize species abundances to maintain biodiversity, we may be better off managing to avoid extinctions of populations by identifying thresholds of acceptable fluctuations in populations of not only hollow-nesting birds but other forest dependent wildlife relative to scientifically valid forest management and habitat structural surrogates.

Disease, habitat fragmentation and conservation

Habitat loss and the resultant fragmentation of remaining habitat is the primary cause of loss of biological diversity. How do these processes affect the dynamics of parasites and pathogens? Hess has provided some important insights into this problem using metapopulation models for pathogens that exhibit 'S-I' dynamics; for example, pathogens such as rabies in which the host population may be divided into susceptible and infected individuals. A major assumption of Hess's models is that infected patches become extinct, rather than recovering and becoming resistant to future infections. In this paper, we build upon this framework in two different ways: first, we examine the consequences of including patches that are resistant to infection; second, we examine the consequences of including a second species of host that can act as a reservoir for the pathogen. Both of these effects are likely to be important from a conservation perspective. The results of both sets of analysis indicate that the benefits of corridors and other connections that allow species to disperse through the landscape far outweigh the possible risks of increased pathogen transmission. Even in the commonest case, where harmful pathogens are maintained by a common reservoir host, increased landscape connectance still allows greater coexistence and persistence of a threatened or endangered host.

Spatial structure and habitat variation in a grasshopper hybrid zone

A hybrid zone between the grasshoppers Chorthippus brunneus and C. jacobsi (Orthoptera: Acrididae) in northern Spain has been analyzed for variation in morphology and ecology. These species are readily distinguished by the number of stridulatory pegs on the hind femur. Both sexes are fully winged and inhabit disturbed habitats throughout the study area. We develop a maximum-likelihood approach to fitting a two-dimensional cline to geographical variation in quantitative traits and for estimating associations of population mean with local habitat. This method reveals a cline in peg number approximately 30 km south of the Picos de Europa Mountains that shows substantial deviations in population mean compared with the expectations of simple tension zone models. The inclusion of variation in local vegetation in the model explains a significant proportion of the residual variation in peg number, indicating that habitat-genotype associations contribute to the observed spatial pattern. However, this association is weak, and a number of populations continue to show strong deviations in mean even after habitat is included in the final model. These outliers may be the result of long-distance colonization of sites distant from the cline center or may be due to a patchy pattern of initial contact during postglacial expansion. As well as contrasting with the smooth hybrid zones described for Chorthippus parallelus, this situation also contrasts with the mosaic hybrid zones observed in Gryllus crickets and in parts of the hybrid zone between Bombina toad species, where habitat-genotype associations account for substantial amounts of among-site variation.

Mapping glider habitat in dry eucalypt forests for Montreal Process indicator 1.1: Fragmentation of forest types

Accurate habitat mapping is critical to landscape ecological studies such as required for developing and testing Montreal Process indicator 1.1e, fragmentation of forest types. This task poses a major challenge to remote sensing, especially in mixedspecies, variable-age forests such as dry eucalypt forests of subtropical eastern Australia. In this paper, we apply an innovative approach that uses a small section of one-metre resolution airborne data to calibrate a moderate spatial resolution model (30 m resolution; scale 1:50 000) based on Landsat Thematic Mapper data to estimate canopy structural properties in St Marys State Forest, near Maryborough, south-eastern Queensland. The approach applies an image-processing model that assumes each image pixel is significantly larger than individual tree crowns and gaps to estimate crown-cover percentage, stem density and mean crown diameter. These parameters were classified into three discrete habitat classes to match the ecology of four exudivorous arboreal species (yellowbellied glider Petaurus australis, sugar glider P. breviceps, squirrel glider P. norfolcensis , and feathertail glider Acrobates pygmaeus), and one folivorous arboreal marsupial, the greater glider Petauroides volans. These species were targeted due to the known ecological preference for old trees with hollows, and differences in their home range requirements. The overall mapping accuracy, visually assessed against transects (n = 93) interpreted from a digital orthophoto and validated in the field, was 79% (KHAT statistic = 0.72). The KHAT statistic serves as an indicator of the extent that the percentage correct values of the error matrix are due to ‘true’ agreement verses ‘chance’ agreement. This means that we are able to reliably report on the effect of habitat loss on target species, especially those with a large home range size (e.g. yellow-bellied glider). However, the classified habitat map failed to accurately capture the spatial patterning (e.g. patch size and shape) of stands with a trace or sub-dominance of senescent trees. This outcome makes the reporting of the effects of habitat fragmentation more problematic, especially for species with a small home range size (e.g. feathertail glider). With further model refinement and validation, however, this moderateresolution approach offers an important, cost eff e c t i v e advancement in mapping the age of dry eucalypt forests in the region.

Enriched behavioral prediction equation and its impact on structured learning and the dynamic calculus

This theoretical note describes an expansion of the behavioral prediction equation, in line with the greater complexity encountered in models of structured learning theory (R. B. Cattell, 1996a). This presents learning theory with a vector substitute for the simpler scalar quantities by which traditional Pavlovian-Skinnerian models have hitherto been represented. Structured learning can be demonstrated by vector changes across a range of intrapersonal psychological variables (ability, personality, motivation, and state constructs). Its use with motivational dynamic trait measures (R. B. Cattell, 1985) should reveal new theoretical possibilities for scientifically monitoring change processes (dynamic calculus model; R. B. Cattell, 1996b), such as encountered within psycho therapeutic settings (R. B. Cattell, 1987). The enhanced behavioral prediction equation suggests that static conceptualizations of personality structure such as the Big Five model are less than optimal.

Microhabitat use by the rainbowfish Melanotaenia duboulayi in a subtropical Australian stream

Microhabitat use and feeding behavior of the rainbowfish Melanotaenia duboulayi (Castelnau) were investigated in a slow-flowing stream adjacent to riparian forest in south-eastern Queensland, Australia. Fish were more abundant in vegetated areas, but did not enter dense Vallisneria beds, where predators were observed. In sunny conditions shoals of juveniles occurred near the water surface feeding floating material on the surface, but larger fish tended to occur at the bottom near submerged vegetation, often utilizing the overhanging aquatic plant community as a refuge and food source. In the middle of the day, juveniles and small fish seemed to show behavioral thermoregulation at the surface in the warmest site. Under cloudy conditions, however, fish of all sizes preferred deeper water. The present study suggests that in still and sunny pools thermal change caused by sunlight influences the microhabitat choice of small fish. A field experiment using a kingfisher model implies that fish swimming at the surface could escape from aerial predators in sunlit conditions by responding to moving shadows, but could not do so under cloudy conditions.

Habitat choice and vigilance behaviour of brush-tailed rock-wallabies (Petrogale penicillata) within their nocturnal foraging ranges

Habitat choice by brush-tailed rock-wallabies (Petrogale penicillata) in south-east Queensland was investigated by comparing the attributes of the nocturnal foraging locations that they selected with those of random locations within a radius of 50 m. Brush-tailed rock-wallabies were shown to select foraging locations on the basis of forage quality and/or their ability to see predators, rather than protection from predators amongst vegetation that could conceal them. Habitat choice may have been affected by limited food availability, as this study was conducted in the winter dry season. The attributes of foraging locations that brush- tailed rock-wallabies perceived as increasing their predation risk were assessed by recording the proportion of time that brush- tailed rock-wallabies spent vigilant while foraging. To measure vigilance, focal animals were observed with a night- vision scope for two minutes and the proportions of time spent vigilant and feeding were recorded. No measured feature of foraging locations was related to higher vigilance levels, suggesting that brush- tailed rock-wallabies did not alter their vigilance whether sheltered amongst grass tussocks or in open habitat, or whether feeding on good quality or poorer quality vegetation. Vigilance levels significantly declined as overnight temperatures decreased, which may have resulted from higher energy requirements of brush- tailed rock-wallabies during winter. The only factors that were found to significantly increase vigilance levels were high winds and moonlit nights. On bright nights, brush- tailed rock-wallabies were very unsettled and during high winds they often did not emerge to feed. More information is needed about how macropods detect predators at night before the effects of wind and light intensity upon vigilance can be fully understood.

Habitat-predator association and avoidance in rainbowfish (Melanotaenia spp.)

The ability to recall the location of a predator and later avoid it was tested in nine populations of rainbowfish (Melanotaenia spp.), representing three species from a variety of environments. Following the introduction of a model predator into a particular microhabitat, the model was removed, the arena rotated and the distribution of the fish recorded again. In this manner it could be determined what cues the fish relied on in order to recall the previous location of the predator model. Fish from all populations but one (Dirran Creek) were capable of avoiding the predator by remembering either the location and/or the microhabitat in which the predator was recently observed. Reliance on different types of visual cues appears to vary between populations but the reason for this variation remains elusive. Of the ecological variables tested (flow variability, predator density and habitat complexity), only the level of predation appeared to be correlated with the orientation technique employed by each population. There was no effect of species identity, which suggests that the habitat that each population occupies plays a strong role in the development of both predator avoidance responses and the cues used to track predators in the wild.

Occurrence of immature Culicoides molestus (Skuse) (Diptera : Ceratopogonidae) in relation to habitat characteristics

Analysis of beach sites on the Gold Coast, Australia, found that 14 physical and chemical habitat characteristics differed significantly between sites where numerous immatures of the canal biting midge, Culicoides molestus (Skuse), were found and sites where no midge immatures occurred. Five of the chemical factors found to reliably distinguish C. molestus habitat are major components of seawater, while another, electrical conductivity, is related to the concentration of seawater components. Calcium was the only one of the six primary components of seawater that was not a statistically significant correlate of C. molestus habitation by sand analysis. It is likely that a causative variable in occurrence of immatures is the concentration of seawater present in canals, because larvae are found where seawater component concentration is low in relation to uninhabited sites of similar appearance.

Low-density koala (Phascolarctos cinereus) populations in the mulgalands of south-west Queensland. III. Broad-scale patterns of habitat use

To date there have been few quantitative studies of the distribution of, and relative habitat utilisation by, koalas in the mulgalands of Queensland. To examine these parameters we applied habitat-accessibility and relative habitat-utilisation indices to estimates of faecal pellet density sampled at 149 sites across the region. Modelling the presence of pellets using logistic regression showed that the potential range of accessible habitats and relative habitat use varied greatly across the region, with rainfall being probably the most important determinant of distribution. Within that distribution, landform and rainfall were both important factors affecting habitat preference. Modelling revealed vastly different probabilities of finding a pellet under trees depending on the tree species, canopy size, and location within the region.

Heritability of adult body height: A comparative study of twin cohorts in eight countries

A major component of variation in body height is due to genetic differences, but environmental factors have a substantial contributory effect. In this study we aimed to analyse whether the genetic architecture of body height varies between affluent western societies. We analysed twin data from eight countries comprising 30,111 complete twin pairs by using the univariate genetic model of the Mx statistical package. Body height and zygosity were self-reported in seven populations and measured directly in one population. We found that there was substantial variation in mean body height between countries; body height was least in Italy (177 cm in men and 163 cm in women) and greatest in the Netherlands (184 cm and 171 cm, respectively). In men there was no corresponding variation in heritability of body height, heritability estimates ranging from 0.87 to 0.93 in populations under an additive genes/unique environment (AE) model. Among women the heritability estimates were generally lower than among men with greater variation between countries, ranging from 0.68 to 0.84 when an additive genes/shared environment/unique environment (ACE) model was used. In four populations where an AE model fit equally well or better, heritability ranged from 0.89 to 0.93. This difference between the sexes was mainly due to the effect of the shared environmental component of variance, which appears to be more important among women than among men in our study populations. Our results indicate that, in general, there are only minor differences in the genetic architecture of height between affluent Caucasian populations, especially among men.

Padrões de uso de habitat do boto-cinza (Sotalia guianensis) na região da foz do rio Doce, costa norte do Espírito Santo, Sudeste do Brasil

Entre março de 2011 e dezembro de 2012 foram realizadas 30 saídas de campo para avaliar os padrões de uso de habitat e estrutura de grupo de Sotalia guianensis na região da foz do rio Doce, costa norte do Espírito Santo. A coleta de dados foi realizada com auxílio de uma embarcação a partir de rotas pré-determinadas distribuídas homogeneamente na área de estudo. Informações sobre as variáveis ambientais foram coletadas a cada hora de amostragem e a cada grupo de golfinhos avistado. Através da análise de Cluster foram identificados três diferentes habitats baseados na similaridade entre variáveis ambientais. Durante 242hs48min de amostragem foram percorridos 1.523km e 119 grupos de botos-cinza foram avistados. Grupos de Sotalia guianensis, assim como indivíduos imaturos, foram observados ao longo de todo o ano na área de estudo, não apresentando diferença sazonal entre as estações seca (abril a setembro) e chuvosa (outubro a março). Os grupos apresentaram uma média de 9,9 indivíduos. Animais imaturos estavam presentes na maioria dos grupos (71%), apresentando uma relação direta com o número de adultos. Os botos-cinza apresentaram o uso quase exclusivo da região marinha aberta adjacente à desembocadura do rio apontando um padrão de uso de habitats heterogêneo, onde as áreas de maior concentração de golfinhos não corresponderam às áreas preferenciais de alimentação. Os resultados obtidos neste trabalho refletem como S. guianensis responde a complexidade dos fatores bióticos e abióticos da área de estudo, apontando um padrão de uso de habitat diferenciado para a espécie.