Components of breeding performance in two competing species: habitat heterogeneity, individual quality and density-dependence

Density‐dependent breeding performance due to habitat heterogeneity has been shown to regulate populations of territorial species, since the progressive occupation of low quality territories as breeding density increases may cause a decline in the mean per capita fecundity of a population while variation in fecundity increases. Although the preemptive use of sites may relegate low quality individuals to sites of progressively lower suitability, few studies on density dependence have tried to separate the effects of territory quality from individual quality, and none have simultaneously considered the effects of heterospecific competitors. Using two long‐term monitored populations, we assessed the relative contribution of habitat heterogeneity and bird quality (in terms of age) on the productivity of sympatric golden Aquila chrysaetos and Bonelli's eagles Hieraaetus fasciatus under different scenarios of intra‐ and inter‐specific competition. Productivity (number of offspring fledged) varied among territories and average annual productivity was negatively related to its variability in both species and populations, thus giving some support to the habitat heterogeneity hypothesis. However, the effect of habitat heterogeneity on productivity became non‐significant when parental age and local density estimators were included in multivariate analyses. Therefore, temporal changes in bird quality (age) combined with intra‐ and interspecific competition explained variability in territory productivity rather than habitat heterogeneity among territories per se. The recruitment of subadult breeders, a surrogate of mortality in eagles, strongly varied among territories. Habitat heterogeneity in productivity may thus arise not because sites differ in suitability for reproduction but because of differences in factors affecting survival. Territories associated with high mortality risks have a higher probability of being occupied by young birds, whose lower quality, interacting with the density competitors, leads to a reduction of productivity. Site‐dependent variability in adult survival and interspecific competition may be extensive, but so far largely overlooked, factors to be seriously considered for the site‐dependent population regulation framework.     

Short- and long-term consequences of individual and territory quality in a long-lived bird

Site-quality is a major determinant of fitness but its effect can be confounded by individual quality, a relationship that has been little studied in large, long-lived vertebrates. The fitness effects of quality estimates depend on the assumption of co-variation between individual and territory quality and can be framed as five working hypotheses: no effect on fitness, exclusive effect of individual quality, exclusive effect of site quality, and independent or interactive effects of the two. We explored such a framework using a medium-sized raptor, the black kite Milvus migrans, as a model species. Individual and territory quality co-varied, but the strength of the relationship varied across different estimates of individual quality (age, body size, or mass residuals). Short-term production of fledglings was related to the independent effects of both individual and territory quality. However, longer-term production of recruits was related solely to territory quality. The disappearance of individual quality effects over the long-term may be caused by antagonistic selective pressures acting during different stages of the life cycle. Our results contribute to a growing appreciation of the long-term fitness-benefits of advantages experienced in early life and highlight the importance of a long-term perspective in studies assessing the effects of individual and territory quality. In our case study, prioritizing sites for conservation on the basis of territory quality may be a feasible pathway to maintain the viability of the population. However, scenarios where such a method could be inefficient have been previously reported, suggesting caution in its application. More studies are needed to understand the generality of the efficiency of priority-setting approaches based on site quality.     

Tasmania revisited: rotifer communities and habitat heterogeneity

The results of four field surveys for Rotifera in Tasmania are summarized. Most new species and records in a 1987 survey were from acid waters (pH < 4.0) of dune lakes on the west coast (42° S). Marked intra- and interhabitat differences in rotifer communities of lakes and ponds were demonstrated by cluster analysis and related to habitat heterogeneity.     

Site-dependent population dynamics: the influence of spatial habitat heterogeneity on individual fitness in the sedge warbler Acrocephalus schoenobaenus

During nine years of study, we detected a mechanism corresponding to site dependence operating in a population of the sedge warbler Acrocephalus schoenobaenus inhabiting a natural wetland of high within-habitat heterogeneity. Sites with a larger share of cover by tall wetland vegetation were preempted during spring settlement; they were occupied in more breeding seasons and by more experienced males. The fitness of males occupying these sites was higher in terms of local recruit production. The total area occupied by the population expanded or contracted depending on its population size. This mechanism increased or decreased the mean site quality, influencing local recruitment of young, although mainly by changing the share of unproductive sites in the total pool of occupied sites. The results indicate that population demographic rates may depend on the spatial heterogeneity of resources at the level of individuals.     

Access to mates in a territorial ungulate is determined by the size of a male's territory, but not by its habitat quality

1. Territoriality is commonly associated with resource defence polygyny, where males are expected to gain access to females by anticipating how resources will influence female distribution and competing for resource-rich sites to establish their zone of dominance. 2. We tested this hypothesis in European roe deer (Capreolus capreolus) by simultaneously assessing the influence of resources on female distribution and the influence of female distribution on male distribution and breeding success using paternity analyses. 3. Females did not fully distribute themselves among male territories in relation to resources. As a result, relative female abundance in a male's territory depended on territory size, but not on its habitat quality. In turn, relative female abundance in a male's territory determined, at least partially, his breeding success. 4. Interestingly, male territory size, and hence access to females, was partly determined by male body mass (all males) and by residual antler size (subadults only). The latter result suggests that large antlers may be important to young males for establishing their first territory, which is then usually retained for all subsequent reproductive seasons. 5. To conclude, although territoriality of male roe deer has certainly evolved as a tactic for ensuring access to mates, our results suggest that it does not really conform to a conventional resource defence polygyny strategy, as males seem to gain no obvious benefit from defending a territory in an area of high habitat quality in terms of enhanced access to mates. 6. This may explain the stability of male territories between years, suggesting that male territoriality conforms to an 'always stay' and 'low risk-low gain' mating strategy in roe deer.     

Small-scale heterogeneity in soil quality influences photosynthetic efficiency and habitat selection in a clonal plant

BACKGROUND AND AIMS: In clonal plants, internode connections allow translocation of photosynthates, water, nutrients and other substances among ramets. Clonal plants form large systems that are likely to experience small-scale spatial heterogeneity. Physiological and morphological responses of Fragaria vesca to small-scale heterogeneity in soil quality were investigated, together with how such heterogeneity influences the placement of ramets. As a result of their own activities plants may modify the suitability of their habitats over time. However, most experiments on habitat selection by clonal plants have not generally considered time as an important variable. In the present study, how the foraging behaviour of clonal plants may change over time was also investigated. METHODS: In a complex of environments with different heterogeneity, plant performance was determined in terms of biomass, ramet production and photosynthetic activity. To identify habitat selection, the number of ramets produced and patch where they rooted were monitored. KEY RESULTS: Parent ramets in heterogeneous environments showed significantly higher maximum and effective quantum yields of photosystem II than parents in homogeneous environments. Parents in heterogeneous environments also showed significantly higher investment in photosynthetic biomass and stolon/total biomass, produced longer stolons, and had higher mean leaf size than parents in homogeneous environments. Total biomass and number of offspring ramets were similar in both environments. However, plants in homogeneous environments showed random allocation of offspring ramets to surrounding patches, whereas plants in heterogeneous environments showed preferential allocation of offspring to higher-quality patches. CONCLUSIONS: The results suggest that F. vesca employs physiological and morphological strategies to enable efficient resource foraging in heterogeneous environments and demonstrate the benefits of physiological integration in terms of photosynthetic efficiency. The findings indicate that short-term responses cannot be directly extrapolated to the longer term principally because preferential colonization of high-quality patches means that these patches eventually show reduced quality. This highlights the importance of considering the time factor in experiments examining responses of clonal plants to heterogeneity.     

Functional diversity versus species diversity: relationships with habitat heterogeneity at multiple scales in a subtropical evergreen broad-leaved forest

Understanding the relationship between functional and species diversity as well as their association with habitat heterogeneity can help reveal the mechanisms of species coexistence in ecological communities. However, these interactions have been poorly studied in subtropical forests. In this paper, we evaluated functional diversity (as measured by Rao’s Q) and traditional species diversity (based on Simpson’s index) in a 24 ha forest plot in a subtropical evergreen broad-leaved forest (EBLF) in China. We compared the sensitivities of functional and species diversity to topographic variables (elevation, convexity, slope and aspect) at multiple spatial scales based on 10 × 10, 20 × 20, 40 × 40 and 50 × 50 m quadrats. Functional and species diversity were found to have different distribution patterns along a topographical gradient, with functional diversity better explained by topography than was species diversity using a spatial autocorrelation regression error model. Furthermore, functional diversity had a significantly greater association with topographic variables than species diversity in both adult and young trees; in both cases, the strength of the diversity-habitat association increased with quadrat size. We conclude that functional diversity reflects a greater diversity-habitat association in EBLF than does species diversity, and that the association depends on the spatial scale and life stages of the woody plants under evaluation.     

Diversity and abundance of birds in relation to forest fragmentation,habitat quality and heterogeneity

This study investigated the importance of habitat quality and habitat heterogeneity for the abundance and diversity of breeding birds in continuous forest and in forest fragments surrounded by farmland in central Sweden. Positive correlations were found between species number and area, volume of Aspen Populus tremula and habitat heterogeneity. Spatial segregation of habitats at a relatively fine-grained scale is suggested to allow for the co-occurrence of more species. The abundance of at least 18 of the species in this study was influenced by fragmentation, and nine of these species preferred fragments to forest sites. The total density of birds was higher in fragments than in forest sites, probably because several fragment species forage in farmland surrounding the sites and a few also forage at edges. Nine species were more common in forest sites than in fragments, but only one species was restricted to continuous forest. However, several fragments were relatively close to forests (150 m) and forest was common in larger scale contexts. The abundance of most species (25 of 33 species) in this study was correlated with habitat quality variables (i.e. variables measuring the size, volume and diversity of ‘tree species’). Among these habitat variables the most important was the occurrence of deciduous trees which seemed to be important for 14 species. The second most important habitat factor seemed to be the diameter of trees, which was positively correlated with the abundance of eight species of which five are hole-nesters. Among coniferous trees, six species were positively correlated with the volume of Norway Spruce Picea abies, whereas no species seemed to be correlated with the volume of Pine Pinus sylvestris.     

Effects of individual condition and habitat quality on natal dispersal behaviour in a small rodent

1. Individuals should benefit from settling in high-quality habitats, but dispersers born under favourable conditions have a better physical condition and should therefore be more successful at settling in high-quality habitats. 2. We tested these predictions with root voles (Microtus oeconomus) by a manipulation of individual condition through litter-size enlargement and reduction during lactation combined with a manipulation of habitat quality through degradation of the vegetation cover. We accurately monitored movements of 149 juveniles during a settlement and breeding period of 3 months. 3. The litter size treatment had long-lasting effects on body size, life-history traits and home range size, but did not influence dispersal behaviour. 4. Different stages of dispersal were influenced by habitat quality. In low-quality patches, females dispersed earlier, spent more time prospecting their environment before settling, and settlers had a smaller adult body size than in high-quality patches. Preference and competition for high-quality patches is likely adaptive as it increased fitness both in terms of survival and reproduction. 5. We found no interactive effect of individual condition and habitat quality on natal dispersal and habitat selection. 6. These findings suggest that immediate conditions are more important determinants of dispersal decisions than conditions experienced early in life.     

Point patterns of tree distribution determined by habitat heterogeneity and dispersal limitation

Understanding processes underlying spatial distribution of tree species is fundamental to studying species coexistence and diversity. This study modeled point patterns of tree distribution, expressed by Cartesian coordinates of individual trees within a mapped forest stand, for the purpose of identifying processes that may generate spatial patterns of tree communities. We used four primary point pattern processes (homogeneous Poisson process, inhomogeneous Poisson process, homogeneous Thomas process, and inhomogeneous Thomas process) to model tree distribution in two stem-mapped forests in Taiwan, Republic of China. These four models simulate spatial processes of habitat association and seed dispersal, allowing us to evaluate the potential contribution of habitat heterogeneity and dispersal limitation to the formation of spatial patterns of tree species. The results showed that the inhomogeneous Thomas process was the best fit model and described most of the species studied, suggesting that spatial patterns of tree species might be formed by the joint effects of habitat associations and dispersal limitation. The homogeneous Thomas process that models the effect of dispersal limitation was the second best model. We also found that the best fit models could be predicted by species attributes, including species abundance and dispersal mode. The significant traits, however, differed between the two study plots and demonstrated site-specific patterns. This study indicated that the interactive operation of niche-based (habitat heterogeneity) and neutral-based (dispersal limitation) may be important in generating spatial patterns of tree species in forest communities.     

Consequences for host-parasitoid interactions of grazing-dependent habitat heterogeneity

1. Environmental heterogeneity can produce effects that cascade up to higher trophic levels and affect species interactions. We hypothesized that grazing-dependent habitat heterogeneity and grazing-independent host plant heterogeneity would influence directly and indirectly a host-parasitoid interaction in a woodland habitat. 2. Thistles were planted randomly in 20 birch woodlands, half of which are grazed by cattle. The abundances of two species of seed herbivore and their shared parasitoid were measured, and related to habitat and host-plant heterogeneity. 3. The presence of cattle grazing created a structurally and compositionally distinct plant assemblage from the ungrazed seminatural situation. Grazing did not affect the number or dispersion of the host plant underpinning the host-parasitoid interaction. 4. The density of one insect herbivore, Tephritis conura, and its parasitoid Pteromalus elevatus was significantly increased by the presence of cattle; but another herbivore, Xyphosia miliaria, was unaffected. The percentage of parasitism of T. conura was increased in grazed habitat occurring at twice the rate found in ungrazed habitat. 5. The increase in T. conura abundance was correlated with increased species richness and cover of forbs in grazed sites. This effect of grazing-dependent habitat variation on host insect density cascaded up to parasitoid density and percentage of parasitism. Habitat heterogeneity had a further direct, positive effect on parasitoid density and percentage of parasitism after controlling for host-insect density. 6. Independent of grazing, heterogeneity in host-plant flowering, architecture and stature further affected T. conura and its parasitoid's densities. Parasitoid density was also affected by the dispersion of the host plant. 7. A combination of habitat and host-plant scale environmental heterogeneity influenced a host-parasitoid interaction indirectly and directly, providing a rare example of an anthropogenic disturbance positively affecting a tertiary trophic level. This finding highlights the need to consider not only the importance of bottom-up effects for top-down processes, but also the role of environmental heterogeneity arising from anthropogenic disturbance for trophic interactions such as parasitism.     

A process of pair formation leading to assortative mating: passive age-assortative mating by habitat heterogeneity

We present an individual-based model dealing with mating as a process of pair formation. Model simulations, based on data from a 19-year study of Spanish imperial eagles, Aquila adalberti, showed that the mating pattern of a population is not necessarily a direct consequence of the mating preferences of individuals; positive age-assortative mating, by which individuals of similar age are more likely to become paired, does not necessarily indicate homotypic mating preferences. For example, individuals of similar, young age may be constrained to a few low-quality territories, leading to passive assortative mating, independent of individual preferences. Confounding factors such as territory quality can affect the encounters between a male and a female available for mating, generating an age-assortative mating totally independent of mate preferences. Such a process may apply to many territorial species when spatial variation in territory quality is pronounced. Copyright 2003 Published by Elsevier Science Ltd on behalf of The Association for the Study of Animal Behaviour.      

Enhancement of habitat heterogeneity and species richness on rocky shores inundated by sand

Summary Many rocky shores are subject to periodic inundation by sand, which is often thought to reduce species richness by eliminating organisms intolerant of sand scour or sand smothering. However, regular disturbance (e.g. inundation) should promote richness by preventing the development of low diversity climax communities. A study of faunal richness on 10 regularly inundated shores showed that inundation does promote richness, but by increasing habitat heterogeneity. Some species are excluded from parts of the shore by sand, but because of the patchiness of sand deposits they are rarely excluded from the entire shore. Other species are found only on rocks associated with sand, while typically sandy shore animals occur in the sand deposits themselves. Total richness (281 species) was greater than for local noninundated shores and sandy beaches combined.     

Microclimate and habitat heterogeneity through the oil palm lifecycle

The rapid expansion of oil palm cultivation and corresponding deforestation has invoked widespread concern for biodiversity in Southeast Asia and throughout the tropics. However, no study explicitly addresses how habitat characteristics change when (1) forest is converted to oil palm, or (2) through the dynamic 25–30-year oil palm lifecycle. These two questions are fundamental to understanding how biodiversity will be impacted by oil palm development.Our results from a chronosequence study on microclimate and vegetation structure in oil palm plantations surrounding the Pasoh Forest Reserve, Peninsular Malaysia, show dramatic habitat changes when forest is converted to oil palm. However, they also reveal substantial habitat heterogeneity throughout the plantation lifecycle. Oil palm plantations are created by clear-cutting forests and then terracing the land. This reduces the 25 m-tall forest canopy to bare ground with a harsh microclimate. Eight-year-old oil palm plantations had 4 m open-canopies; 22-year-old plantations had 13 m closed-canopies. Old plantations had significantly more buffered microclimates than young plantations. Understory vegetation was twice as tall in young plantations, but leaf litter depth and total epiphyte abundance were double in old plantations. Nonetheless, leaf litter coverage was patchy throughout the oil palm life cycle due to the stacking of all palm fronds. Overall, oil palm plantations were substantially hotter (+2.84 °C) and drier (+0.80 hPa vapor pressure deficit), than forests during diurnal hours. However, there were no nocturnal microclimate differences between forests and plantations. Finally, we describe how the variable retention of old palm trees during crop rotation can retain habitat features and maintain more stable microclimate conditions than clear-cutting senescent plantations. We discuss the implications of habitat changes for biodiversity and introduce three methods to utilize temporal habitat heterogeneity to enhance the quality of the oil palm landscape matrix.     

The influence of habitat heterogeneity on host-pathogen population dynamics

The influence of spatial heterogeneity on the population dynamics of a naturally occurring invertebrate host-pathogen system was experimentally investigated. At ten week intervals over a two year period, I quantified the spatial distribution of natural populations of the terrestrial isopod crustacean Porcellio scaber infected with the isopod iridescent virus (IIV). During the seasonally dry periods of summer and early fall in central California, isopod populations were highly aggregated and the degree of patchiness and distance between inhabited patches was greatest. Coincident with increased patchiness and patch spacing was an increase in isopod density within patches. During the wet seasons of winter and spring, isopod population patchiness, inter-patch spacing, and within-patch density was low. Seasonal changes in virus prevalence were negatively correlated with within-patch density, patchiness, and inter-patch spacing. The influence of the spatial distribution of isopods on virus prevalence was also tested in field experiments. The virus was introduced into arrays of artificial habitat patches colonized by isopods in which interpatch distance was varied. The prevalence of resulting infections was monitored at weekly intervals. In addition, dispersal rates between artificial patches and natural patches were quantified and compared. The results showed that isopods in treatments with the smallest inter-patch spacing had the highest virus prevalence, with generally lower prevalence among isopods in more widely spaced patches. The spacing of experimental patches significantly affected virus prevalence, although the experiments did not resolve a clear relationship between patch spacing and virus prevalence. Rates of dispersal between patches decreased with increased patch spacing, and these rates did not differ significantly from dispersal between natural patches. The results suggest that rates of dispersal between isopod subpopulations may be an important component of the infection dynamics in this system. I discuss the consequences of these findings for host-pathogen dynamics in fragmented habitats, and for other ecological interactions in spatially heterogeneous habitats.     

Freshwater snail diversity: effects of pond area,habitat heterogeneity and isolation

Summary A large number of eutrophic ponds were surveyed for the presence of freshwater gastropods. Factors thought to influence the distribution of the snails were evaluated. As the investigated area has a homogeneous geological background physicochemical factors probably have a low effect on the local distribution of snails. There was a significant, positive regression between pond area and the number of gastropod species in the pond, but the regression only explained a minor part of the variation in species numbers. Multiple regression of an extended number of variables, associated with habitat complexity and dispersal, indicated that, in addition to area, macrophyte diversity and the mean number of gastropods in the 5 closest ponds (S₅) were important in explaining the distribution of gastropods. An increase in the number of macrophyte species increases the number of available microhabitats and refuges from predators. S₅ probably influences the dispersal rates between ponds. The gastropods in this area are thought to have additional dispersal modes, besides aerial dispersal with birds, and this probably increases the immigration rates and/or decreases the extinction rate.