Dispersal costs set the scene for helping in an atypical avian cooperative breeder

The ecological constraints hypothesis is suggested to explain the evolution of cooperative breeding in birds. This hypothesis predicts that the scene for cooperative breeding is set when ecological factors constrain offspring from dispersal. This prediction was tested in the atypical cooperative breeding system of the long-tailed tit, Aegithalos caudatus, by comparing the degree of philopatry and cooperation in an isolated and a contiguous site whilst experimentally controlling for confounding aspects of reproduction. No difference was found between the two sites in the survival of offspring but a greater proportion were found to remain philopatric in the isolated site. This difference was caused by greater philopatry of normally dispersive females suggesting, as predicted, that dispersal costs were greater from this site. Furthermore, a greater proportion of males and females cooperated following breeding failure in the isolated site than in the contiguous site. Thus, as has been suggested for typical avian cooperative breeders, dispersal costs, relative to philopatric benefits, appear to set the scene for cooperative breeding in long-tailed tits.     

The effect of woodland area on avian community composition in a fragmented southern UK landscape and associated management recommendations

ABSTRACT

Capsule: Smaller woodlands not only support fewer species but also show different avian community composition due to loss of woodland interior and an increase in edge habitat.

Aims: To use observed community composition changes, rather than traditional total species richness-area relationships, to make area-specific management recommendations for optimizing woodland habitat for avian communities in fragmented landscapes.

Methods: 17 woodlands were selected in Oxfordshire, UK, with areas between 0.2 and 120 ha. Three dawn area searches were conducted in each woodland between 1st April and 28th May 2016 to record encounter rates for each species. The impact of internal habitat variation on woodland comparability was assessed using habitat surveys.

Results: Woodlands with area less than 3.6 ha showed a significant positive relationship between total avian species richness and woodland area. Woodlands with area over 3.6 ha were all consistent with a mean (± se) total richness of 25.4?±?0.6 species, however the number of woodland specialists continued to increase with woodland area. Woodland generalists dominated the total encounter rate across the area range, however the fractional contribution of woodland specialists showed a significant positive correlation with woodland area, while the fractional contribution of non-woodland species significantly decreased. Non-woodland species numbers peaked in mid-sized woodlands with enhanced habitat heterogeneity.

Conclusions: Community composition analysis enabled more targeted recommendations than total species richness analysis, specifically: large woodlands (over 25?ha) in southern UK should focus conservation efforts on providing the specific internal habitats required by woodland specialists; medium-sized woodlands (between approximately 4 and 25?ha) should focus on promoting internal habitat variety, which can benefit both woodland species and non-woodland species of conservation concern in the surrounding landscape; small woodlands (under 4?ha) should focus on providing nesting opportunities for non-woodland species and on improving connectivity to maximize habitat for woodland generalists and facilitate movement of woodland specialists.     

The response of carabids to landscape simplification differs between trophic groups

We studied the response of carabid species richness and density to landscape simplification (measured as percentage cover of non-crop habitat surrounding each study site) in 36 wheat fields using pitfall traps. Carabids were divided to trophic groups following the literature. The number of species from different trophic groups declined with increasing landscape simplification in the order: carnivores > phytophages > omnivores. Density compensation of both carnivores and phytophages suggests that species decline is caused by the loss of specific resources rather than by an overall reduction in food availability. Increasing evenness indicates that a greater share of phytophagous species contributes to density compensation at poorer sites. A comparison with data from complementing studies shows that marked differences in species numbers (carnivores > omnivores > phytophages) are due to a different sensitivity of trophic groups to agricultural management. Since our findings seem to be partly due to increasing sensitivity to landscape changes with trophic rank, and partly to decreasing sensitivity of depauperate communities to local environmental stress, species loss can best be explained by the co-action of factors at local and regional scales. Species richness decline might significantly alter the role of carabids as biocontrol agents.     

The relative importance of landscape and community features in the invasion of an exotic shrub in a fragmented landscape

Although invasive plants are recognized as a major ecological problem, little is known of the relative importance of plant community characteristics versus landscape context in determining invasibility of communities. We determined the relative importance of community and landscape features of 30 woodlots in influencing the invasion of Lonicera maackii. We sampled woodlots using the point‐quarter method and calculated canopy openness and basal areas and densities of shrub, sapling and tree species, as well as woody species richness. We used aerial photos and ArcView GIS to calculate landscape parameters from the same woodlots using a buffer distance of 1500 m. We used logistic and linear regression analyses to determine the community and landscape factors that best explain L. maackii presence and density. We also tested whether woodlot invasion by L. maackii begins at woodlot edges. Presence of L. maackii was significantly explained only by distance from the nearest town (logistic regression, p=0.017); woodlots nearer town were more likely to be invaded. Among invaded woodlots, density of L. maackii was positively related to the amount of edge in the landscape (partial R²=0.592) and negatively related to total tree basal area (partial R²=0.134), number of native woody species (partial R²=0.054), and sapling shade tolerance index (partial R²=0.054). Lonicera maackii in woodlot interiors were not younger than those on the perimeters, leading us to reject the edge‐first colonization model of invasion. Our findings reveal that landscape structure is of primary importance and community features of secondary importance in the invasion of L. maackii. This shrub is invading from multiple foci (towns) rather than an advancing front. Connectivity in the landscape (i.e. the number of corridors) did not promote invasion. However, edge habitat was important for invasion, probably due to increased propagule pressure. The community features associated with L. maackii invasion may be indicators of past disturbance.     

The presence of an avian co-forager reduces vigilance in a cooperative mammal

Many animals must trade-off anti-predator vigilance with other behaviours. Some species facilitate predator detection by joining mixed-species foraging parties and ‘eavesdropping’ on the predator warnings given by other taxa. Such use of heterospecific warnings presumably reduces the likelihood of predation, but it is unclear whether it also provides wider benefits, by allowing individuals to reduce their own vigilance. We examine whether the presence of an avian co-forager, the fork-tailed drongo (Dicrurus adsimilis), affects rates of vigilance (including sentinel behaviour) in wild dwarf mongooses (Helogale parvula). We simulate the presence of drongos—using playbacks of their non-alarm vocalizations—to show that dwarf mongooses significantly reduce their rate of vigilance when foraging with this species. This is, to our knowledge, the first study to demonstrate experimentally that a mammal reduces vigilance in the presence of an avian co-forager.     

Testing models of parental investment strategy and offspring size in ants

Parental investment strategies can be fixed or flexible. A fixed strategy predicts making all offspring a single ‘optimal’ size. Dynamic models predict flexible strategies with more than one optimal size of offspring. Patterns in the distribution of offspring sizes may thus reveal the investment strategy. Static strategies should produce normal distributions. Dynamic strategies should often result in non-normal distributions. Furthermore, variance in morphological traits should be positively correlated with the length of developmental time the traits are exposed to environmental influences. Finally, the type of deviation from normality (i.e., skewed left or right, or platykurtic) should be correlated with the average offspring size. To test the latter prediction, we used simulations to detect significant departures from normality and categorize distribution types. Data from three species of ants strongly support the predicted patterns for dynamic parental investment. Offspring size distributions are often significantly non-normal. Traits fixed earlier in development, such as head width, are less variable than final body weight. The type of distribution observed correlates with mean female dry weight. The overall support for a dynamic parental investment model has implications for life history theory. Predicted conflicts over parental effort, sex investment ratios, and reproductive skew in cooperative breeders follow from assumptions of static parental investment strategies and omnipresent resource limitations. By contrast, with flexible investment strategies such conflicts can be either absent or maladaptive. Electronic Supplementary Material Supplementary material is available for this article at     

Contrasting heterozygosity-fitness correlations between populations of a self-compatible shrub in a fragmented landscape

The mechanisms underlying heterozygosity-fitness correlations (HFCs) are subject of intense debates, especially about how important population features such as size or degree of isolation influence HFCs. Here, we report variation in HFCs between Large and Small populations of a self-compatible shrub (Myrtus communis) occurring within an extremely fragmented landscape. In each of the five study populations, we obtained data on both heterozygosity and fitness for 9-12 maternal families (i.e. offspring from the same mother plant). Whereas heterozygosity explained most of the variance (60-86?%) in growth rate of seedling families within Large populations, this relationship was absent within Small populations. Our results suggest that inbreeding may explain the observed HFCs within Large populations, and that different genetic processes (such as genetic drift and/or selection) could have overridden HFCs within Small populations. While it is difficult to draw general conclusions from five populations, we think our results open new research perspectives on how different genetic processes underlie variation in HFCs under different population contexts. Our study also points to a need for further attention on the complex relationships between heterozygosity in self-compatible plants and their progeny in relation to mating system variation. Finally, our results provide interesting new insights into how population genetic diversity is maintained or lost in a highly fragmented landscape.     

Distribution of an increasing roe deer population in a fragmented Mediterranean landscape

This paper describes roe deer Capreolus capreolus distribution in central Spain, where the species has spread from the mountains into peripheral agricultural areas In this region, it is more abundant in forest fragments near the mountains than in those further away, in pine and oak woodlands than in sclerophyllous forests, and in forests with open water than in those lacking this resource The paper also analyzes whether habitat quality and geographic location of forests with respect to roe deer source areas are the two basic causes of its distribution, as predicted by some models of the species distribution in fragmented landscapes The results corroborate this hypothesis revealing that roe deer abundance is linked negatively to sclerophyllous forest cover and distance from mountains We conclude that this type of farmland seems to be sub-optimal for roe deer in comparison with forested, moist mountains     

The evolution of avian parental care

A stage model traces key behavioural tactics and life-history traits that are involved in the transition from promiscuity with no parental care, the mating system that typifies reptiles, to that typical of most birds, social monogamy with biparental care. In stage I, females assumed increasing parental investment in precocial young, female choice of mates increased, female-biased mating dispersal evolved and population sex ratios became male biased. In stage II, consortships between mating partners allowed males to attract rare social mates, provided a mechanism for paternity assessment and increased female ability to assess mate quality. In stage III, relative female scarcity enabled females to demand parental investment contributions from males having some paternity certainty. This innovation was facilitated by the nature of avian parental care; i.e. most care-giving activities can be adopted in small units. Moreover, the initial cost of care giving to males was small compared with its benefit to females. Males, however, tended to decline to assume non-partitionable, risky, or relatively costly parental activities. In stage IV, altriciality coevolved with increasing biparental care, resulting in social monogamy. Approaches for testing behavioural hypotheses are suggested.     

The frequency of metapopulations, metacommunities and nestedness in a fragmented landscape

Metapopulation and metacommunity theory may be extremely useful for understanding community composition and species distributions in fragmented landscapes. However, how well particular spatial ecological models represent natural systems is not known and so the general importance of those models is unclear. In three naturally fragmented landscapes in Tasmania, Australia, I sampled beetles from 67 sites, including continuous forest, small forest patches in a sedgeland matrix, streamside forest, and the matrix. Of forty commonly captured beetle species, at least 37% do not form metapopulations of any kind because they do not disperse far enough, disperse too far, or occur in the matrix. Only 7.5% of the species showed distributions consistent with a mainland‐island metapopulation, which was surprising given the mainland‐island structure of the landscape. These patterns were also consistent with a classic metapopulation. Deterministic metapopulations were probably common, involving at least 32% of species. Most predicted metacommunity patterns were not observed because only subsets of the fauna followed a particular metacommunity model. Seven, mostly flying, species were influenced only by habitat quality, supporting the Species‐sorting or Mass‐effects models, and the strong distinction between the beetle fauna of Eucalyptus patches and the matrix suggested an over‐riding influence of a Species‐sorting process. Seven, mostly flightless, species showed a negative relationship with increasing patch isolation, while another six, mostly flying, species showed the opposite pattern, with higher frequency of occurrence in more isolated patches. Both groups were strongly nested. The flying species were probably excluded from less isolated sites through interactions with poor dispersers , providing a novel mechanism leading to a nested distribution. The Patch‐dynamics and Species‐sorting processes were equally consistent with the pattern, emphasizing that these models are relatively simplistic when confronted with real community data. Additional complexity needs to be built into the models to accommodate the diverse patterns observed in natural communities.     

Habitat fragmentation shapes natal dispersal and sociality in an Afrotropical cooperative breeder

It remains poorly understood how effects of anthropogenic activity, such as large-scale habitat fragmentation, impact sociality in animals. In cooperatively breeding species, groups are mostly formed through delayed offspring dispersal, and habitat fragmentation can affect this process in two opposite directions. Increased habitat isolation may increase dispersal costs, promoting delayed dispersal. Alternatively, reduced patch size and quality may decrease benefits of philopatry, promoting dispersal. Here, we test both predictions in a cooperatively breeding bird (placid greenbul, Phyllastrephus placidus) from an Afrotropical cloud forest archipelago. Males born in fragmented forest dispersed about 1 year earlier than those born in continuous forest. Contrary to females, males also started to reproduce earlier and mostly settled within their natal patch. Females only rarely delayed their dispersal for more than 1 year, both in fragmented and continuous forests. Our results suggest that early male dispersal and reproduction is jointly driven by a decrease in the value of the natal territory and an increase in local breeding opportunities in fragmented forest. While plasticity in dispersal strategies of cooperative breeders in response to anthropogenic change is believed to optimize reproduction-survival trade-offs, to what extent it shapes the ability of species to respond to rapid environmental change remains to be studied.     

A land snail's view of a fragmented landscape

Habitat fragmentation may influence the genetic structure of populations, especially of species with low mobility. So far, these effects have been mainly studied by surveying neutral markers, and much less by looking at ecologically relevant characters. Therefore, we aimed to explore eventual patterns of covariation between population structuring in neutral markers and variation in shell morphometrics in the forest‐associated snail Discus rotundatus in relation to habitat fragment characteristics. To this end, we screened shell morphometric variability and sequence variation in a fragment of the mitochondrial 16S rDNA gene in D. rotundatus from the fragmented landscape of the Lower Rhine Embayment, Germany. The 16S rDNA of D. rotundatus was highly variable, with a total of 118 haplotypes (384 individuals) forming four clades and one unresolved group. There was a geographic pattern in the distribution of the clades with the river Rhine apparently separating two groups. Yet, at the geographic scale considered, there was no obvious effect of fragmentation on shell morphometrics and 16S rDNA variation because G_ST often was as high within, as between forests. Instead, the age of the habitat and (re‐)afforestation events appeared to affect shell shape and 16S rDNA in terms of the number of clades per site. The ecologically relevant characters thus supported the presumably neutral mitochondrial DNA markers by indicating that populations of not strictly stenecious species may be (relatively) stable in fragments. However, afforestation after large clearcuts and habitat gain after the amendment of deforestation are accompanied by several, seemingly persistent peculiarities, such as altered genetic composition and shell characters (e.g. aperture size). © 2009 The Linnean Society of London, Biological Journal of the Linnean Society, 2009, 98 , 839–850.     

Host-parasitoid extinction and colonization in a fragmented prairie landscape

Few field studies of natural populations have examined the factors influencing local extinctions and colonization of empty habitat patches for a prey species and its predator. In this study, I carried out a census of planthopper (Prokelisia crocea; Hemiptera: Delphacidae) and egg parasitoid (Anagrus columbi; Hymenoptera: Mymaridae) incidence and densities in 147 host-plant patches (Spartina pectinata; Poaceae) over seven planthopper generations in a tall-grass prairie landscape. For both species, the likelihood of going extinct in a patch was related to a number of patch-specific variables: density, temporal variability in density, proportion of hosts parasitized (planthopper only), host-plant density, patch size, patch isolation, and composition of the surrounding matrix. Colonization likelihood was related only to the physical attributes of the patch. There was high patch turnover in this prairie landscape. On average, planthoppers went extinct in 23% of the patches and A. columbi went extinct in 51% of the patches in each generation. For the planthopper, extinction likelihood increased with a decrease in patch size and the proportion of the matrix composed of mudflat. Parasitism of eggs had no effect on the extinction likelihood of local P. crocea populations, suggesting that A. columbi may not play a major role in the patch dynamics of its host. The likelihood of extinction for A. columbi was dependent on factors that spanned three trophic levels. An increase in plant density, decrease in host density and decrease in parasitoid density all increased the likelihood of A. columbi extinction within a patch. The dependency on multiple trophic levels may explain the higher extinction risk for the parasitoid than its host. A. columbi extinction was also affected by the matrix habitat surrounding the patch—the effect was the opposite of that for P. crocea. Finally, vacant patches were colonized at rates of 53% and 34% per generation for the planthopper and parasitoid, respectively. For both species, colonization probabilities decreased with an increase in patch isolation. High host densities in a patch also favored high rates of colonization by A. columbi. I discuss how anthropogenic changes to the prairie landscape can affect the metapopulation dynamics and persistence time of this host-parasitoid interaction.     

Migration, coherence and persistence in a fragmented landscape

The chance of local extinction is high during periods of small population size. Accordingly, a metapopulation made of local communities that support internal population cycling may face the threat of regional extinction if the local dynamics is coherent (synchronized). These systems achieve maximum sustainability at an intermediate level of migration that allows recolonization but prevents synchronization. Here we implement an individual-based simulation technique to examine the maximum persistence condition for a system of patch habitats connected by passive migration. The models discussed in this paper take into consideration realistic elements of metapopulations, such as migration cost, disordered spatial structure, frustration and environmental noise. It turns out that the state with maximum anti-correlation between neighboring patches is the most sustainable one, even in the presence of these complications. The results suggest, at least for small systems, a model independent conservation strategy: coherence between neighboring local communities has, in general, a negative impact, and population will benefit from intervention that increases anti-correlations.     

The oxidative costs of reproduction are group-size dependent in a wild cooperative breeder

Life-history theory assumes that reproduction entails a cost, and research on cooperatively breeding societies suggests that the cooperative sharing of workloads can reduce this cost. However, the physiological mechanisms that underpin both the costs of reproduction and the benefits of cooperation remain poorly understood. It has been hypothesized that reproductive costs may arise in part from oxidative stress, as reproductive investment may elevate exposure to reactive oxygen species, compromising survival and future reproduction and accelerating senescence. However, experimental evidence of oxidative costs of reproduction in the wild remains scarce. Here, we use a clutch-removal experiment to investigate the oxidative costs of reproduction in a wild cooperatively breeding bird, the white-browed sparrow weaver, Plocepasser mahali. Our results reveal costs of reproduction that are dependent on group size: relative to individuals in groups whose eggs were experimentally removed, individuals in groups that raised offspring experienced an associated cost (elevated oxidative damage and reduced body mass), but only if they were in small groups containing fewer or no helpers. Furthermore, during nestling provisioning, individuals that provisioned at higher rates showed greater within-individual declines in body mass and antioxidant protection. Our results provide rare experimental evidence that reproduction can negatively impact both oxidative status and body mass in the wild, and suggest that these costs can be mitigated in cooperative societies by the presence of additional helpers. These findings have implications for our understanding of the energetic and oxidative costs of reproduction, and the benefits of cooperation in animal societies.