Genes and song: genetic and social connections in fragmented habitat in a woodland bird with limited dispersal

Understanding the processes leading to population declines in fragmented landscapes is essential for successful conservation management. However, isolating the influence of disparate processes, and dispersal in particular, is challenging. The Grey Shrike-thrush, Colluricincla harmonica, is a sedentary woodland-dependent songbird, with learned vocalizations whose incidence in suitable habitat patches falls disproportionally with decline in tree cover in the landscape. Although it has been suggested that gaps in tree cover might act as barriers to its dispersal, the species remains in many remnants of native vegetation in agricultural landscapes, suggesting that it may have responded to habitat removal and fragmentation by maintaining or even increasing dispersal distances. We quantified population connectivity of the Grey Shrike-thrush in a system fragmented over more than 120 years using genetic (microsatellites) and acoustic (song types) data. First, we tested for population genetic and acoustic structure at regional and local scales in search of barriers to dispersal or gene flow and signals of local spatial structuring indicative of restricted dispersal or localized acoustic similarity. Then we tested for effects of habitat loss and fragmentation on genetic and acoustic connectivity by fitting alternative models of mobility (isolation-by-distance [the null model] and reduced and increased movement models) across treeless vs. treed areas. Birds within -5 km of each other had more similar genotypes and song types than those farther away, suggesting that dispersal and song matching are limited in the region. Despite restricted dispersal detected for females (but not males), populations appeared to be connected by gene flow and displayed some cultural (acoustic) connectivity across the region. Fragmentation did not appear to impact greatly the dispersal of the Grey Shrike-thrush: none of the mobility models fit the genetic distances of males, whereas for females, an isolation-by-distance model could not be rejected in favor of the models of reduced or increased movement through treeless gaps. However, dissimilarities of the song types were more consistent with the model of reduced cultural connectivity through treeless areas, suggesting that fragmentation impedes song type sharing in the Grey Shrike-thrush. Our paper demonstrates that habitat fragmentation hinders important population processes in an Australian woodland bird even though its dispersal is not detectably impacted.     

Comparative social network analysis in a leaf-roosting bat

Even though social network analysis provides an important tool to characterize and compare societies, no studies have used its analytical applications to characterize patterns of sociality in bats. Here I use social network analysis to characterize and compare patterns of sociality between three populations of the leaf-roosting bat Thyroptera tricolor. Sites differed in the density of furled leaves used by T. tricolor for roosting. Finca had more leaves per hectare (77), followed by Ureña (58), and Esquinas (7). The time period over which the probability of association is halved based on fitted models was 1,086 days for Finca, 714 days for Ureña, and 303 days for Esquinas. Finca and Ureña had very similar network topologies, with several small clusters, high-clustering coefficients, short path lengths, low node betweenness, and high network robustness. Social networks at Esquinas were composed of one large cluster and several small isolated ones. Esquinas also had high-clustering coefficients, but path length and node betweenness were high. Network resilience was lower in Esquinas compared to Finca and Ureña. These results show that, unlike many other forest-dwelling bats that switch roosts regularly, T. tricolor does not exhibit a typical fission–fusion social system, and that resource availability seems to affect social networks in this bat. In addition, this study highlights the importance of emigrating individuals in maintaining social cohesion, establishing network connectedness, and determining network robustness.     

Episodic targeting aggression and the histories of Lemur social groups

Summary Periodically, members of captive Lemur social groups target others for intense aggression. Over periods of several days, weeks, and sometimes months, one to three lemurs persistently follow and attack one or two particular group mates until the targets no longer associate with their group. Episodic targeting aggression is nonrandom with regard to time of year, group size and sex ratio, and the kinship, age, and gender of targets. The vast majority of episodes observed over the past 18 years at the Duke University Primate Center (DUPC) has occurred between like-sex adolescents and adults in conjunction with estrous cycling or infant births and most often after groups have reached apparent critical sizes. When unrelated adults have shared group membership, members of one family have almost invariably first targeted members of the other. In several groups, entire matrilines have gradually been evicted by members of another across periods of several years. When non-relatives have been absent, lemurs have evicted relatives that had previously formed separate subgroups. Episodic targeting aggression has been documented at the DUPC in over a dozen different social groups, comprising three different species. The phenomenon occurs repeatedly in groups held in a variety of large outdoor runs as well as in outdoor enclosures providing naturalistic space and physical structure. More-over, an appreciable number of recent observations in Madagascar suggest that the patterns we have documented well represent the phenomenon as it occurs in the wild. Targeting aggression based on group size, sex ratio, kinship and gender has been reported for no other primate taxon. We suggest that episodic targeting aggression reflects heretofore undescribed tactics in reproductive competition that may characterize many lemurid and indriid taxa. As such, the phenomenon has broad implications for the structure of lemur social groups and populations. Provisional models of the social dynamics and histories of Lemur social groups are presented for evaluation during upcoming field work.     

Behavioural trait assortment in a social network: patterns and implications

The social fine structure of a population plays a central role in ecological and evolutionary processes. Whilst many studies have investigated how morphological traits such as size affect social structure of populations, comparatively little is known about the influence of behaviours such as boldness and shyness. Using information on social interactions in a wild population of Trinidadian guppies (Poecilia reticulata), we construct a social network. For each individual in the network, we quantify its behavioural phenotype using two measures of boldness, predator inspection tendency, a repeatable and reliably measured behaviour well studied in the context of co-operation, and shoaling tendency. We observe striking heterogeneity in contact patterns, with strong ties being positively assorted and weak ties negatively assorted by our measured behavioural traits. Moreover, shy fish had more network connections than bold fish and these were on average stronger. In other words, social fine structure is strongly influenced by behavioural trait. We assert that such structure will have implications for the outcome of selection on behavioural traits and we speculate that the observed positive assortment may act as an amplifier of selection contributing to the maintenance of co-operation during predator inspection.     

Male sperm whale (Physeter macrocephalus) acoustics in a high-latitude habitat: implications for echolocation and communication

Sperm whales (Physeter macrocephalus) are deep-diving predators foraging in meso- and bathypelagic ecosystems off the continental shelves. To investigate the ecophysiological and communicative function of various click types from male sperm whales in a high-latitude habitat, we deployed a large-aperture array of calibrated hydrophones off northern Norway (N69, E15). Data show that sperm whales in this habitat produce three click types: usual clicks, creak clicks and, occasionally, slow clicks. Usual clicks and creak clicks exhibit short duration, profound directionality and a frequency content suited for echolocation on meso- and bathypelagic fish and squids. The acoustic properties and low repetition rate of usual clicks are suited for long-range echolocation, whereas creak clicks have properties equivalent to signals in buzzes, the terminating pulse trains known from echolocating bats during prey capture. From these source parameters and the high acoustic activity during foraging dives, it is concluded that echolocation is an important sensory cue in prey location. Sound pressure levels of creak clicks and usual clicks measured off the acoustic axis suggest that sperm whales may be subjected to eavesdropping by conspecifics, thereby conveying information about food aggregations at estimated ranges of 6 km for creak clicks and 16 km for usual clicks. Slow clicks exhibit low directionality, low-frequency emphasis and a reduced repetition rate, suggesting that this click type is more suited for communication than for echolocation. Slow clicks can be detected by submerged conspecifics at ranges up to 60 km. Thus, sperm whales producing slow clicks may represent an odontocete species that utilizes long-range sound communication.     

Generating spatially optimized habitat in a trade‐off between social optimality and budget efficiency

Auctions have been proposed as alternatives to payments for environmental services when spatial interactions and costs are better known to landowners than to the conservation agency (asymmetric information). Recently, an auction scheme was proposed that delivers optimal conservation in the sense that social welfare is maximized. I examined the social welfare and the budget efficiency delivered by this scheme, where social welfare represents the difference between the monetized ecological benefit and the conservation cost incurred to the landowners and budget efficiency is defined as maximizing the ecological benefit for a given conservation budget. For the analysis, I considered a stylized landscape with land patches that can be used for agriculture or conservation. The ecological benefit was measured by an objective function that increases with increasing number and spatial aggregation of conserved land patches. I compared the social welfare and the budget efficiency of the auction scheme with an agglomeration payment, a policy scheme that considers spatial interactions and that was proposed recently. The auction delivered a higher level of social welfare than the agglomeration payment. However, the agglomeration payment was more efficient budgetarily than the auction, so the comparative performances of the 2 schemes depended on the chosen policy criterion–social welfare or budget efficiency. Both policy criteria are relevant for conservation. Which one should be chosen depends on the problem at hand, for example, whether social preferences should be taken into account in the decision of how much money to invest in conservation or whether the available conservation budget is strictly limited.     

The role of size and aggression in intrasexual male competition in a social lizard species, Egernia whitii

Competition between males is a key component of the agonistic intrasexual interactions that influence resource acquisition, social system dynamics, and ultimately reproductive success. Sexual selection theory predicts that traits that enhance success in intrasexual competition (particularly male–male competition) should be favored. In vertebrates, this often includes body size and aggression, with larger and/or more aggressive males outcompeting smaller or less aggressive conspecifics. The majority of studies consider aggression as a flexible trait which responds to local social or environmental conditions. However, aggression frequently shows considerable within-individual consistency (i.e., individuals have identifiable aggressive behavioral types). Little is known about how such consistency in aggression may influence competition outcomes. We integrated a detailed field study with a laboratory experiment to examine how a male’s aggressive phenotype and his size influence competitive interactions in Egernia whitii, a social lizard species which exhibits strong competition over resources (limited permanent shelter sites and basking sites). Individual aggression and size did not predict competition outcome in the laboratory nor did they predict home range size, overlap, or reproductive success in the field. However, winners of laboratory trial contests maintained consistent aggressive phenotypes while consistency in aggression was lost in losers. We suggest that aggression may be important in other functional contexts, such as parental care, and that alternative traits, such as fighting experience, may be important in determining competition outcome in this species.     

Size and spacing of capercaillie leks in relation to social behavior and habitat

Summary Spring territories of 17 adult capercaillie (Tetrao urogallus L.) cocks were 10–79 ha in extent; they varied inversely in size with the relative proportion of mature forest within them. The number of resident cocks at leks increased with the amount of mature forest within a 1 km radius of each lek center. Leks in two areas (n=46) were regularly spaced with mean distances to nearest neighbors of 1.98 and 2.07 km, corresponding to the territorial space occupied by adult cocks of adjoining leks. In one area with intensive logging, interlek distance increased with decreasing amounts of mature forest between them. Spring home ranges of 18 adult females averaged 51.3 ha±8.2 SE. The spatial relationships did not fit recent models of interlek spacing based on female spacing behavior. Instead, the results suggested that spacing of leks may be related to the territorial requirements of males.     

A foundation for developing a methodology for social network sampling

Researchers are increasingly turning to network theory to understand the social nature of animal populations. We present a computational framework that is the first step in a series of works that will allow us to develop a quantitative methodology of social network sampling to aid ecologists in their social network data collection. To develop our methodology, we need to be able to generate networks from which to sample. Ideally, we need to perform a systematic study of sampling protocols on different known network structures, as network structure might affect the robustness of any particular sampling methodology. Thus, we present a computational tool for generating network structures that have user-defined distributions for network properties and for key measures of interest to ecologists. The user defines the values of these measures and the tool will generate appropriate network randomizations with those properties. This tool will be used as a framework for developing a sampling methodology, although we do not present a full methodology here. We describe the method used by the tool, demonstrate its effectiveness, and discuss how the tool can now be utilized. We provide a proof-of-concept example (using the assortativity measure) of how such networks can be used, along with a simulated egocentric sampling regime, to test the level of equivalence of the sampled network to the actual network. This contribution is part of the special issue “Social Networks: new perspectives” (Guest Editors: J. Krause, D. Lusseau and R. James).     

Potential banana skins in animal social network analysis

Social network analysis is an increasingly popular tool for the study of the fine-scale and global social structure of animals. It has attracted particular attention by those attempting to unravel social structure in fission–fusion populations. It is clear that the social network approach offers some exciting opportunities for gaining new insights into social systems. However, some of the practices which are currently being used in the animal social networks literature are at worst questionable and at best over-enthusiastic. We highlight some of the areas of method, analysis and interpretation in which greater care may be needed in order to ensure that the biology we extract from our networks is robust. In particular, we suggest that more attention should be given to whether relational data are representative, the potential effect of observational errors and the choice and use of statistical tests. The importance of replication and manipulation must not be forgotten, and the interpretation of results requires care. This contribution is part of the special issue “Social Networks: new perspectives” (Guest Editors: J. Krause, D. Lusseau and R. James).     

Male mating competitiveness and age-dependent relationship between testosterone and social rank in bighorn sheep

In males, the acquisition and development of behavioral and morphological secondary sexual traits typically depends on testosterone and correlates with mating success. Testosterone level could affect competition for mates and thus be a target of sexual selection. We sought to relate testosterone levels to male mating competitiveness, by teasing apart the relationships between testosterone, behavior, and growth before the mating period. We monitored 24 adult bighorn rams (Ovis canadensis) at Ram Mountain, Alberta, from 2008 to 2011. Using linear mixed models, we tested the relationships between testosterone metabolites in feces, social rank, and both growth and size of two sexually selected traits: horns and body mass. The correlation between testosterone and social rank varied with age. Testosterone and rank were weakly and negatively correlated for young rams, positively correlated for prime-aged rams, and negatively correlated for older rams. Although testosterone had an increasingly positive effect on total horn length until 8 years of age, we could not detect any effects on annual growth rate of horns or body mass. Testosterone may be related to male’s ability to compete for mates through its relationship with behaviors determining social rank, rather than by influencing the development of morphological traits. Differences in testosterone levels among competitors may be a proximate cause of variance in fitness.     

Structure of the social network and its influence on transmission dynamics in a honeybee colony

Infectious processes in a social group are driven by a network of contacts that is generally structured by the organization arising from behavioral and spatial heterogeneities within the group. Although theoretical models of transmission dynamics have placed an overwhelming emphasis on the importance of understanding the network structure in a social group, empirical data regarding such contact structures are rare. In this paper, I analyze the network structure and the correlated transmission dynamics within a honeybee colony as determined by food transfer interactions and the changes produced in it by an experimental manipulation. The study demonstrates that widespread transmission in the colony is correlated to a lower clustering coefficient and higher robustness of the social network. I also show that the social network in the colony is determined by the spatial distribution of various age classes, and the resulting organizational structure provides some amount of immunity to the young individuals. The results of this study demonstrates how, using the honeybee colony as a model system, concepts in network theory can be combined with those in behavioral ecology to gain a better understanding of social transmission processes, especially those related to disease dynamics.     

Signature whistle variations in a bottlenosed dolphin,Tursiops truncatus

To examine whether context-specific information is superimposed upon the individual cues present in the whistling of the bottlenosed dolphin, Tursiops truncatus, parameter variations within the two most frequently emitted whistle types of a captive individual were investigated in three different behavioural contexts. The study concentrated on comparing signal features of spontaneously occurring vocalizations in two possible phases following the performance of a trained discrimination task and those occurring during isolation. Phases of the discrimination task differed according to whether the animal showed correct (reward given) or incorrect performance (no reward). Signature whistles were most common in isolation, but also represented just over half of the whistles following a choice task. Of 14 signature whistle frequency and time parameters measured 9 differed significantly between isolation and at least one of the phases following a choice task (Table 1). Three parameters also varied according to whether performance was correct or incorrect. In contrast, only one out of four parameters (start frequency) measured from the second most frequent whistle type varied significantly between contexts (isolation vs. phase following correct choice). The results indicate that not only identity but also context-related information is available in the whistles of a bottlenosed dolphin.     

Pollinator effectiveness varies with experimental shifts in flowering time

The earlier flowering times exhibited by many plant species are a conspicuous sign of climate change. Altered phenologies have caused concern that species could suffer population declines if they flower at times when effective pollinators are unavailable. For two perennial wildflowers, Tradescantia ohiensis and Asclepias incarnata, we used an experimental approach to explore how changing phenology affects the taxonomic composition of the pollinator assemblage and the effectiveness of individual pollinator taxa. After finding in the previous year that fruit set varied with flowering time, we manipulated flowering onset in greenhouses, placed plants in the field over the span of five weeks, and measured pollinator effectiveness as the number of seeds produced after a single visit to a flower. The average effectiveness of pollinators and the expected rates of pollination success were lower for plants of both species flowering earlier than for plants flowering at historical times, suggesting there could be reproductive costs to earlier flowering. Whereas for A. incarnata, differences in average seed set among weeks were due primarily to changes in the composition of the pollinator assemblage, the differences for T. ohiensis were driven by the combined effects of compositional changes and increases over time in the effectiveness of some pollinator taxa. Both species face the possibility of temporal mismatch between the availability of the most effective pollinators and the onset of flowering, and changes in the effectiveness of individual pollinator taxa through time may add an unexpected element to the reproductive consequences of such mismatches.     

Emigration mechanisms in feral house mice – a laboratory investigation of the influence of social structure,population density,and aggression

Emigration in small mammals may be strongly related to social factors, but direct observations of emigrants are rare. Feral house mice (Mus domesticus) were studied using a population cage system that allowed continuous observation of individually marked animals. Mice that left their natal cage and took up residence in cages that could only be reached by crossing a water barrier were defined as emigrants. Six pairs of house mice with their litters were placed in the system, and data on aggressive interactions, body weight, reproduction, mortality and emigration were collected daily. Both sexes emigrated, but males did so twice as often as females. Population density was not correlated with the frequency of aggression, and had no effect on the weight of emigrating individuals. Male emigrants suffered more aggression before emigration than their non-emigrant brothers of the same age; they were aggressively driven out by other males, predominantly by the father. Female emigration depended on the female’s chances of reproduction. The probability of a female reproducing decreased with increasing birth order. Females born in a late litter, who therefore had only a low chance of reproduction, dispersed earlier than those of early litters. Resident males were reproductively suppressed. Male offspring had two different strategies for attaining top rank. They could develop rapidly and reach sexual maturity early on, but face competition with the father, risking being forced to emigrate. Alternatively, they could develop slowly, stay within their family and wait for a chance to take over the dominant position. It is concluded that emigration in male and female feral house mice is caused by intrasexual competition. Received: 13 July 1995/Accepted after revision: 8 June 1996     

Male greenfinches (Carduelis chloris) with brighter ornaments have higher virus infection clearance rate

Hamilton and Zuk proposed that bright plumage in birds indicates genetic resistance to parasites, and that by selecting brighter males as mates, females can increase their offspring’s fitness due to this inherited resistance. The theory predicts a negative relationship between parasite load and plumage brightness in males. We used Sindbis virus clearance rate after an experimental infection to quantify parasite resistance in male greenfinches (Carduelis chloris) and related variation in clearance rate with variation in male plumage brightness. We found that certain aspects of brightness of the male plumage (i.e. tail-patch area) could be used to predict the virus infection clearance rate. Wing brightness was uninformative of virus clearance rate, but revealed age class. We found no clear relationship between antibody production rate and virus clearance rate or total viraemia. However, males with large tail patches tended to have a higher antibody production rate. The results suggest that the size of the male tail patch may function as an indicator of an individual male’s ability to resist parasite infections, thus supporting the Hamilton-Zuk theory for a novel taxon of parasites, a virus. Received: 11 November 1999 / Received in revised form: 13 March 2000 / Accepted: 1 April 2000     

Structure and resilience of the social network in an insect colony as a function of colony size

Social interactions are critical to the organization of worker activities in insect colonies and their consequent ecological success. The structure of this interaction network is therefore crucial to our understanding of colony organization and functioning. In this paper, I study the properties of the interaction network in the colonies of the social wasp Ropalidia marginata. I find that the network is characterized by a uniform connectivity among individuals with increasing heterogeneity as colonies become larger. Important network parameters are found to be correlated with colony size and I investigate how this is reflected in the organization of work in colonies of different sizes. Finally, I test the resilience of these interaction networks by experimental removal of individuals from the colony and discuss the structural properties of the network that are related to resilience in a social network. This contribution is part of the special issue “Social Networks: new perspectives” (Guest Editors: J. Krause, D. Lusseau, and R. James).     

Heterogeneity in social network connections is density-dependent: implications for disease dynamics in a gregarious ungulate

Behavioral Ecology and Sociobiology - Individual animals across all taxa differ consistently in behaviour, i.e. they show personality traits. This inter-individual variability has significant...