Determinants of Aedes mosquito larval ecology in a heterogeneous urban environment- a longitudinal study in Bengaluru, India.

BACKGROUND: Aedes-borne disease risk is associated with contemporary urbanization practices where city developing structures function as a catalyst for creating mosquito breeding habitats. We lack better understanding on how the links between landscape ecology and urban geography contribute to the prevalence and abundance of mosquito and pathogen spread. METHODS: An outdoor longitudinal study in Bengaluru (Karnataka, India) was conducted between February 2021 and June 2022 to examine the effects of macrohabitat types on the diversity and distribution of larval habitats, mosquito species composition, and body size to quantify the risk of dengue outbreak in the landscape context. FINDINGS: A total of 8,717 container breeding sites were inspected, of these 1,316 were wet breeding habitats. A total of 1,619 mosquito larvae representing 16 species from six macrohabitats and nine microhabitats were collected. Aedes aegypti and Aedes albopictus were the dominant species and significantly higher in artificial habitats than in natural habitats. Breeding preference ratio for Aedes species was high in grinding stones and storage containers. The Aedes infestation indices were higher than the WHO threshold and showed significant linear increase from Barren habitat to High density areas. We found Ae. albopictus breeding in sympatry with Ae. aegypti had shorter wing length. CONCLUSIONS: A large proportion of larval habitats were man-made artificial containers. Landscape ecology drives mosquito diversity and abundance even at a small spatial scale which could be affecting the localized outbreaks. Our findings showed that sampling strategies for mosquito surveillance must include urban environments with non-residential locations and dengue transmission reduction programmes should focus on 'neighbourhood surveillance' as well to prevent and control the rising threat of Aedes-borne diseases.

Spoilt for choice: Do female mosquitoes experience choice overload when deciding where to lay eggs?

Animals live in complex natural environments. Based on the effects of natural selection, theory on animal information use says that it is optimal for animals to make "rational" decisions, i.e., to choose alternatives which maximize fitness gains, irrespective of the number of alternatives presented to them. Yet, animals commonly make seemingly "irrational" choices in the face of complex and variable stimuli that challenge their cognitive machinery. Here, we test the choice overload hypothesis - decision-making is negatively affected when animals experience an overload of choice. Using simultaneous-choice trials that varied in choice repertoire size, we examined oviposition site selection behaviour in Aedes aegypti towards larval predators, the nymphs of Bradinopyga geminata. Based on the underlying fitness trade-offs of oviposition decision-making, we predicted that female oviposition preference would be weaker and variation in this response would be higher in complex, multiple-choice trials than in binary-choice trials. In partial support of our hypothesis, oviposition preference was weaker in the complex, multiple-choice trials, but the variation in response depended on predator density, and did not depend on choice repertoire size. We suggest that information overload can negatively affect certain aspects of animal decision-making, resulting in choices appearing as "irrational" if the complexity of the decision-making context is not incorporated. Information overload can potentially lead to alternative strategies, such as bet-hedging or decision-making with reduced discrimination.

Lekking as collective behaviour.

Lekking is a spectacular mating system in which males maintain tightly organized clustering of territories during the mating season, and females visit these leks for mating. Various hypotheses-ranging from predation dilution to mate choice and mating benefit-offer potential explanations for the evolution of this peculiar mating system. However, many of these classic hypotheses rarely consider the spatial dynamics that produce and maintain the lek. In this article, we propose to view lekking through the perspective of collective behaviour, in which simple local interactions between organisms, as well as habitat, likely produce and maintain lekking. Further, we argue that interactions within the leks change over time, typically over a breeding season, to produce many broad-level as well as specific collective patterns. To test these ideas at both proximate and ultimate levels, we argue that the concepts and tools from the literature on collective animal behaviour, such as agent-based models and high-resolution video tracking that enables capturing fine-scale spatio-temporal interactions, could be useful. To demonstrate the promise of these ideas, we develop a spatially explicit agent-based model and show how simple rules such as spatial fidelity, local social interactions and repulsion among males can potentially explain the formation of lek and synchronous departures of males for foraging from the lek. On the empirical side, we discuss the promise of applying the collective behaviour approach to blackbuck (Antilope cervicapra) leks-using high-resolution recordings via a camera fitted to unmanned aerial vehicles and subsequent tracking of animal movements. Broadly, we suggest that a lens of collective behaviour may provide novel insights into understanding both the proximate and ultimate factors that shape leks. This article is part of a discussion meeting issue 'Collective behaviour through time'.

Antelope space-use and behavior indicate multilevel responses to varying anthropogenic influences in a highly human-dominated landscape.

A primary means of conserving a species or a habitat in a human-dominated landscape is through promoting coexistence with humans while minimizing conflict. For this, we should understand how wildlife is impacted by direct and indirect human activities. Such information is rare in areas with high human densities. To investigate how animals respond to altered ecological conditions in human-dominated landscapes, we focused on a wild herbivore of conservation concern in the Krishnasaar Conservation Area (KrCA) in Nepal. Here, blackbuck Antilope cervicapra, a generalist grazer, lives in refugia located with a growing human population. We studied the impacts of humans on habitat use and behavior of blackbuck. We laid 250 × 250 m grid cells in the entire KrCA and carried out indirect sign surveys with three replications for habitat use assessment. We observed herds of blackbuck for 89 h in different habitat types using scan sampling methods. Our habitat-use survey showed that habitats under intensive human use were hardly used by blackbuck, even when high-quality forage was available. Habitat openness was the major predictor of habitat use inside the core area, where levels of human activities were low. We also found a positive correlation between habitat use by blackbuck and livestock. Blackbuck were substantially more vigilant when they were in forest than in grassland, again indicating an influence of risk. Overall, blackbuck appear to be sensitive to the risk associated with both natural and anthropogenic factors. Our findings have direct implications for managing human-wildlife interactions in this landscape, specifically regarding strategies for livestock grazing in habitats highly used by blackbuck and concerning predictions of how changing land use will impact the long-term persistence of blackbuck. Our work suggests that wild herbivores may be able to persist in landscapes with high human densities so long as there are refuges where human activities are relatively low.

Habitat specialisation and matrix resistance predict responses of butterfly populations to landscape features in tropical grassland-forest complexes.

A fundamental question in ecology is to understand how a species is distributed in a landscape. In terrestrial landscapes, the size and isolation of habitat patches, and matrix properties, are thought to drive population density patterns. Yet, given the same set of landscape features, why do species from a single taxon vary so widely in their density patterns? A primary hypothesis for such variation, proposed by community-level studies, is that key ecological characteristics of species influence how they respond to landscape features. However, robust tests of this hypothesis, which require measurements of populations of multiple species in an assemblage, are still scarce. We investigated the ability of ecological specialisation and the interaction of species with the matrix (matrix resistance) to predict population responses of butterfly species to patch size and connectivity in naturally heterogeneous tropical forest-grassland complexes. We surveyed 56 habitat patches in a 65 sq. km area by laying 276 transects, along which the identity and abundances of butterfly species were recorded. We also used transects that cut across two habitats to estimate matrix resistance. We find that habitat specialisation predicted the strength of area-density and isolation-density relationships. Matrix resistance also predicted variation in area-density relationships, highlighting the importance of species interactions with the matrix. Specialists showed higher matrix resistance and stronger area and isolation effects than did generalists. Our findings suggest that investigating how traits related to ecological specialisation and matrix resistance affect demographic parameters can contribute towards understanding mechanisms underlying species distributions in heterogeneous landscapes.

The impact of leopards (Panthera pardus) on livestock losses and human injuries in a human-use landscape in Maharashtra, India.

There are many ways in which large carnivores and humans interact in shared spaces. In this study we provide insights into human-leopard relationships in an entirely modified, human-dominated landscape inhabited by dense populations of humans (266 per km2), their livestock (162 per km2) and relatively high densities of large predators (10 per 100 km2). No human deaths were recorded, and livestock losses to leopards numbered only 0.45 per km2per year (averaged over three years) despite the almost complete dependency of leopards on domestic animals as prey. Predation was not the major cause of livestock mortality as diseases and natural causes caused higher losses (80% of self-reported losses). We also found that ineffective night time livestock protection and the presence of domestic dogs increased the probability of a farmer facing leopard attacks on livestock. Resident farmers faced much lower livestock losses to leopard predation in contrast to the migratory shepherds who reported much higher losses, but rarely availed of the government compensation schemes. We recommend that local wildlife managers continue to shift from reactive measures such as leopard captures after livestock attacks to proactive measures such as focusing on effective livestock protection and informing the affected communities about safety measures to be taken where leopards occur in rural landscapes. The natural causes of livestock deaths due do diseases may be better prevented by involving animal husbandry department for timely vaccinations and treatment.

The enemy of my enemy: multiple interacting selection pressures lead to unexpected anti-predator responses.

Despite being a major selective force, predation can induce puzzling variability in anti-predator responses-from lack of predator aversion to lifelong predator-induced fear. This variability is hypothesised to result from variation in the trade-offs associated with avoiding predators. But critical information on fitness outcomes of these trade-offs associated with anti-predator behaviours is lacking. We tested this trade-off hypothesis in Aedes aegypti, by examining oviposition site selection decisions in response towards larval predation risk and comprehensively measuring the fitness implications of trade-offs of avoiding larval predators, using three fitness measures: larval survival, development time and size. In a field study, we find that adult females show a surprisingly variable response to predators, ranging from attraction to avoidance. This variation is explained by fitness outcomes of oviposition along a predation-risk gradient that we measured in the laboratory. We show that ovipositing females could gain fitness benefits from ovipositing in pools with a low density of predators, rather than in predator-free pools, as predators provide a release from negative density effects of conspecific larvae that might co-occur in a pool. Interacting selection pressures may thus explain diverse prey responses. We suggest other systems in which similarly unexpected prey behaviour is likely to occur.

Do extra-group fertilizations increase the potential for sexual selection in male mammals?

Fertilizations by males outside the social breeding group (extra-group paternity, EGP) are widespread in birds and mammals. EGP is generally proposed to increase male reproductive skew and thereby increase the potential for sexual selection, but the generality of this relationship is unclear. We extracted data from 27 mammals in seven orders and used phylogenetic comparative methods to investigate the influence of EGP and social mating system on measures of inequality in male fertilization success, which are indices of the potential for sexual selection. We find that EGP and social mating system can predict the potential for sexual selection in mammalian populations, but only when considered jointly and not individually. EGP appears to increase the potential for sexual selection but only when the degree of social polygyny is relatively low. When social polygyny is high, EGP appears to result in a more uniform distribution of reproduction and a decrease in the potential for sexual selection. A possible explanation to be investigated is that the phenotype of extra-group fathers differs systematically across social mating systems. Our findings have implications for the use of EGP and social mating system as indices of sexual selection in comparative analyses of trait evolution under sexual selection.

Alternative reproductive tactics and inverse size-assortment in a high-density fish spawning aggregation.

BACKGROUND: At high densities, terrestrial and marine species often employ alternate reproductive tactics (ARTs) to maximize reproductive benefits. We describe ARTs in a high-density and unfished spawning aggregation of the squaretail grouper (Plectropomus areolatus) in Lakshadweep, India. RESULTS: As previously reported for this species, territorial males engage in pair-courtship, which is associated with a pair-spawning tactic. Here, we document a previously unreported school-courtship tactic; where territorial males court multiple females in mid-water schools, which appears to culminate in a unique 'school-spawning' tactic. Courtship tactics were conditional on body size, local mate density and habitat, likely associated with changing trade-offs between potential mating opportunities and intra-sexual competition. Counter-intuitively, the aggregation showed a habitat-specific inverse size-assortment: large males courted small females on the reef slope while small males courted equal-sized or larger females on the shelf. These patterns remained stable across two years of observation at high, unfished densities. CONCLUSIONS: These unique density-dependent behaviours may disappear from this aggregation as overall densities decline due to increasing commercial fishing pressure, with potentially large consequences for demographics and fitness.

A statistical approach to understanding reproductive isolation in two sympatric species of tree crickets.

In acoustically communicating animals, reproductive isolation between sympatric species is usually maintained through species-specific calls. This requires that the receiver be tuned to the conspecific signal. Mapping the response space of the receiver onto the signal space of the conspecific investigates this tuning. A combinatorial approach to investigating the response space is more informative as the influence on the receiver of the interactions between the features is also elucidated. However, most studies have examined individual preference functions rather than the multivariate response space. We studied the maintenance of reproductive isolation between two sympatric tree cricket species (Oecanthus henryi and Oecanthus indicus) through the temporal features of the calls. Individual response functions were determined experimentally for O. henryi, the results from which were combined in a statistical framework to generate a multivariate quantitative receiver response space. The predicted response was higher for the signals of the conspecific than for signals of the sympatric heterospecific, indicating maintenance of reproductive isolation through songs. The model allows prediction of response to untested combinations of temporal features as well as delineation of the evolutionary constraints on the signal space. The model can also be used to predict the response of O. henryi to other heterospecific signals, making it a useful tool for the study of the evolution and maintenance of reproductive isolation via long-range acoustic signals.

Wild Ungulate Decision-Making and the Role of Tiny Refuges in Human-Dominated Landscapes.

Wildlife conservation in human-dominated landscapes requires that we understand how animals, when making habitat-use decisions, obtain diverse and dynamically occurring resources while avoiding risks, induced by both natural predators and anthropogenic threats. Little is known about the underlying processes that enable wild animals to persist in densely populated human-dominated landscapes, particularly in developing countries. In a complex, semi-arid, fragmented, human-dominated agricultural landscape, we analyzed the habitat-use of blackbuck, a large herbivore endemic to the Indian sub-continent. We hypothesized that blackbuck would show flexible habitat-use behaviour and be risk averse when resource quality in the landscape is high, and less sensitive to risk otherwise. Overall, blackbuck appeared to be strongly influenced by human activity and they offset risks by using small protected patches (~3 km2) when they could afford to do so. Blackbuck habitat use varied dynamically corresponding with seasonally-changing levels of resources and risks, with protected habitats registering maximum use. The findings show that human activities can strongly influence and perhaps limit ungulate habitat-use and behaviour, but spatial heterogeneity in risk, particularly the presence of refuges, can allow ungulates to persist in landscapes with high human and livestock densities.

Reduced hornbill abundance associated with low seed arrival and altered recruitment in a hunted and logged tropical forest.

Logging and hunting are two key direct threats to the survival of wildlife in the tropics, and also disrupt important ecosystem processes. We investigated the impacts of these two factors on the different stages of the seed dispersal cycle, including abundance of plants and their dispersers and dispersal of seeds and recruitment, in a tropical forest in north-east India. We focused on hornbills, which are important seed dispersers in these forests, and their food tree species. We compared abundances of hornbill food tree species in a site with high logging and hunting pressures (heavily disturbed) with a site that had no logging and relatively low levels of hunting (less disturbed) to understand logging impacts on hornbill food tree abundance. We compared hornbill abundances across these two sites. We, then, compared the scatter-dispersed seed arrival of five large-seeded tree species and the recruitment of four of those species. Abundances of hornbill food trees that are preferentially targeted by logging were two times higher in the less disturbed site as compared to the heavily disturbed site while that of hornbills was 22 times higher. The arrival of scatter-dispersed seeds was seven times higher in the less disturbed site. Abundances of recruits of two tree species were significantly higher in the less disturbed site. For another species, abundances of younger recruits were significantly lower while that of older recruits were higher in the heavily disturbed site. Our findings suggest that logging reduces food plant abundance for an important frugivore-seed disperser group, while hunting diminishes disperser abundances, with an associated reduction in seed arrival and altered recruitment of animal-dispersed tree species in the disturbed site. Based on our results, we present a conceptual model depicting the relationships and pathways between vertebrate-dispersed trees, their dispersers, and the impacts of hunting and logging on these pathways.

Rodent seed predation: effects on seed survival, recruitment, abundance, and dispersion of bird-dispersed tropical trees.

Tropical tree species vary widely in their pattern of spatial dispersion. We focus on how seed predation may modify seed deposition patterns and affect the abundance and dispersion of adult trees in a tropical forest in India. Using plots across a range of seed densities, we examined whether seed predation levels by terrestrial rodents varied across six large-seeded, bird-dispersed tree species. Since inter-specific variation in density-dependent seed mortality may have downstream effects on recruitment and adult tree stages, we determined recruitment patterns close to and away from parent trees, along with adult tree abundance and dispersion patterns. Four species (Canarium resiniferum, Dysoxylum binectariferum, Horsfieldia kingii, and Prunus ceylanica) showed high predation levels (78.5-98.7%) and increased mortality with increasing seed density, while two species, Chisocheton cumingianus and Polyalthia simiarum, showed significantly lower seed predation levels and weak density-dependent mortality. The latter two species also had the highest recruitment near parent trees, with most abundant and aggregated adults. The four species that had high seed mortality had low recruitment under parent trees, were rare, and had more spaced adult tree dispersion. Biotic dispersal may be vital for species that suffer density-dependent mortality factors under parent trees. In tropical forests where large vertebrate seed dispersers but not seed predators are hunted, differences in seed vulnerability to rodent seed predation and density-dependent mortality can affect forest structure and composition.

Intraspecific variation in group size in the blackbuck antelope: the roles of habitat structure and forage at different spatial scales.

The main ecological factors that are hypothesized to explain the striking variation in the size of social groups among large herbivores are habitat structure, predation, and forage abundance and distribution; however, their relative roles in wild populations are not well understood. I combined analyses of ecological correlates of spatial variation in group size with analyses of individual behaviour in groups of different sizes to investigate factors maintaining variation in group size in an Indian antelope, the blackbuck Antilope cervicapra. I measured group size, habitat structure, forage, and the occurrence of predators in ten blackbuck populations, and, at a smaller spatial scale, within an intensively studied population. To examine the processes by which these ecological factors influence group size, I used behavioural observations and an experiment to estimate the shape of the relationship between group size and potential costs and benefits to individuals. Group size varied extensively both among and within populations. Analyses of spatial variation in group size suggested that both forage and habitat structure influence group size: large-scale, among-population variation in group size was primarily related to habitat structure, while small-scale, within-population variation was most closely related to forage abundance. Analyses of individual behaviour suggested that larger groups incur greater travel costs while foraging. However, individuals in larger groups appeared to experience greater benefits, namely the earlier detection of a "predator", a reduction in vigilance, and an increase in the time spent feeding. Overall, these findings suggest that individuals in groups experience a trade-off between predation-related benefits and costs arising from feeding competition. Habitat structure and forage likely influence the nature of this trade-off; thus, variation in these ecological factors may maintain variation in group size. The role of predation pressure and other factors in explaining the remaining variation needs further exploration.

Ecological correlates of extra-group paternity in mammals.

Extra-group paternity (EGP) can form an important part of the mating system in birds and mammals. However, our present understanding of its extent and ecology comes primarily from birds. Here, we use data from 26 species and phylogenetic comparative methods to explore interspecific variation in EGP in mammals and test prominent ecological hypotheses for this variation. We found extensive EGP (46% of species showed more than 20% EGP), indicating that EGP is likely to play an important role in the mating system and the dynamics of sexual selection in mammals. Variation in EGP was most closely correlated with the length of the mating season. As the length of the mating season increased, EGP declined, suggesting that it is increasingly difficult for males to monopolize their social mates when mating seasons are short and overlap among females in oestrus is likely to be high. EGP was secondarily correlated with the number of females in a breeding group, consistent with the idea that as female clustering increases, males are less able to monopolize individual females. Finally, EGP was not related to social mating system, suggesting that the opportunities for the extra-group fertilizations and the payoffs involved do not consistently vary with social mating system.