Macroecology of habitat choice in long-distance migratory birds

Patterns of habitat choice in ecological communities are not only influenced by present-day selective forces but also by historical processes, such as the biogeographical history of the lineages they are composed of. Nevertheless, it has been very difficult to test historical factors. The possible tropical origin of long-distance migratory birds provides an opportunity for such a test. If habitat choice of long-distance migrants is inherited from their tropical ancestors then Nearctic long-distance migrants might have acquired their habitat choice from Neotropical forest species and European long-distance migrants from African savannah species. Here we use a macroecological approach to show that this hypothesis can be confirmed. Long-distance migrants in the Nearctic are found in forested habitat types, while those in Europe are found in open ones. In comparison, the habitat choice of residents and short-distance migrants (in genera without long-distance migration) does not differ between the Nearctic and Europe. These results demonstrate that habitat choice in temperate bird communities can be explained by the tropical history of long-distance migrants. Thus, habitat choice seems to be shaped not only by local mechanisms, but also by processes acting on much larger spatial and temporal scales.     

Male collared lizards,Crotaphytus collaris (Sauria: Crotaphytidae), signal females by broadcasting visual displays

The signalling function of displays broadcast when animals are distant from conspecifics can be difficult to determine. I tested the extent to which visually transmitted broadcast displays given by free‐ranging territorial male collared lizards signalled same‐sex rivals or females. One test involved recording the frequency of broadcast displays, aggressive contests with rivals, and courtship encounters with females during ten reproductive seasons when local sex ratios varied markedly. The frequency of broadcast displays decreased as the ratio of male competitors to females increased. The frequency with which males initiated contests with rivals was not related to the ratio of competitors to females, whereas the frequency of courtship interactions decreased with sex ratio because there were fewer females to court. The behaviour of males that defended territories during two successive seasons showed a similar pattern. Broadcast display frequency was positively correlated with courtship frequency, but not with the frequency of contests with rivals. Lastly, individual males gave more broadcast displays during focal observations when they also engaged in courtship encounters with females than other observations when they engaged in aggressive conflicts with rival males. Although these results do not reject the possibility that broadcast displays may also signal male rivals, they support a major role of these displays in advertisement to females. © 2013 The Linnean Society of London     

Habitat exploration and use in dispersing juvenile flying squirrels

1. Variation in behaviours involved in habitat selection is important for several evolutionary and ecological processes. For example, habitat use during dispersal may differ from breeding habitat use, and for dispersers the scale of habitat familiarity is determined by exploratory behaviour. We studied habitat use and exploration of 56 radio-collared juvenile flying squirrels Pteromys volans L. within natal home range and during dispersal, and compared habitat use between juveniles and 37 adults within breeding home range. 2. Before dispersal, young flying squirrels actively moved around the natal site. Surprisingly, long-distance dispersers explored less than short-distance dispersers, but philopatric individuals explored similar distances as dispersers. Females explored less than males, although females are the more dispersive sex in flying squirrels. 3. For most of the individuals the settlement area was unfamiliar due to long dispersal distance. Consequently, direction and distance of exploration were not very strong predictors of settlement location. However, individuals familiar with the settlement area concentrated exploration to that area. Exploration did not correlate with short-term survival. 4. Dispersers preferred breeding habitat while dispersing, but were found more often in matrix habitat than juveniles within natal, or adults within breeding, home ranges. 5. We conclude that familiarity does not determine settlement as much as, for example, availability of the habitat for flying squirrels. Based on our results, it also seems clear that data on adult habitat use are not enough to predict habitat use of dispersing individuals. In addition, our results support the recent view that short- and long-distance dispersers may need to be analysed separately in ecological and evolutionary analyses.     

Field use of maximal sprint speed by collared lizards (Crotaphytus collaris): compensation and sexual selection

To understand how selection acts on performance capacity, the ecological role of the performance trait being measured must be determined. Knowing if and when an animal uses maximal performance capacity may give insight into what specific selective pressures may be acting on performance, because individuals are expected to use close to maximal capacity only in contexts important to survival or reproductive success. Furthermore, if an ecological context is important, poor performers are expected to compensate behaviorally. To understand the relative roles of natural and sexual selection on maximal sprint speed capacity we measured maximal sprint speed of collared lizards (Crotaphytus collaris) in the laboratory and field-realized sprint speed for the same individuals in three different contexts (foraging, escaping a predator, and responding to a rival intruder). Females used closer to maximal speed while escaping predators than in the other contexts. Adult males, on the other hand, used closer to maximal speed while responding to an unfamiliar male intruder tethered within their territory. Sprint speeds during foraging attempts were far below maximal capacity for all lizards. Yearlings appeared to compensate for having lower absolute maximal capacity by using a greater percentage of their maximal capacity while foraging and escaping predators than did adults of either sex. We also found evidence for compensation within age and sex classes, where slower individuals used a greater percentage of their maximal capacity than faster individuals. However, this was true only while foraging and escaping predators and not while responding to a rival. Collared lizards appeared to choose microhabitats near refugia such that maximal speed was not necessary to escape predators. Although natural selection for predator avoidance cannot be ruled out as a selective force acting on locomotor performance in collared lizards, intrasexual selection for territory maintenance may be more important for territorial males.     

Evidence for ecological release over a fine spatial scale in a lizard from the White Sands formation

When a population experiences relaxation of selective pressures due to reduced predation or competition, it may undergo ecological release. Ecological release often manifests as increased niche breadth and trait variation, as individuals have the opportunity to exploit a wider variety of resources. At the recently‐colonized White Sands dune formation in New Mexico, lizards from white sand habitat have fewer predators and competitors than their dark soil counterparts, and show evidence of ecological release compared to dark soil lizards. To determine whether the dynamics of ecological release also play out over an even finer habitat gradient, we studied southwestern fence lizards in the center of the White Sands dune formation and the narrow ecotone between White Sands and the surrounding dark soil habitats. We predicted that lizards from the central dunes would exhibit ecological release in terms of broadened resource use compared to lizards from the ecotone. We first conducted avian surveys in both the central dunes and ecotone habitats to measure abundance, richness and diversity of avian species that could act as lizard predators. Next, we measured microhabitat‐scale resource use by comparing perch selection of lizards in both habitats. Finally, we measured landscape‐scale resource use by quantifying home range sizes and daily distances traveled by lizards in both habitats. We found that central dunes lizards used a greater diversity of perch types, and had greater variation in both home range size and daily distances traveled, than ecotone lizards. There were fewer predatory bird species in the central dunes than on the ecotone, and there was a tendency for fewer predation events in the central dunes than on the ecotone. Our results demonstrate that ecological release is detectable over a fine habitat gradient, such as between the center of a recently‐colonized habitat and its ecotone with the surrounding ecosystem.     

Relations between microhabitat use and limb shape in phrynosomatid lizards

With the exception of the well-documented case for anoline lizards, recent studies have found few evolutionary relationships between morphology and habitat use in lizards despite clear-cut biomechanical predictions. One of the factors typically hampering these analyses is the clustering of habitat use within evolutionary lineages. In the present study, body shape was quantified for male and female lizards of 30 species of phrynosomatid lizards. This group was selected as little clustering of ecological variables seemed to be present. The results of traditional analyses indicate that evolutionary correlates of habitat use were prominent in the hindlimbs of both sexes. Species living in open habitats are characterized by longer femurs, and longer hindlimbs relative to the forelimb. Moreover, males from ground-dwelling species utilizing open habitats have longer toes on the hind foot than males from climbing species. Phylogenetic analyses indicated strong evolutionary associations between habitat use and the relative length of front and hindlimbs, with species from open terrestrial habitats having significantly shorter frontlimbs relative to their hindlimb than rock or tree climbing species. Evolutionary associations between morphology and habitat use were generally stronger for male lizards, indicating a potentially important contribution of sexual selection to the evolution of differences in limb proportions.  © 2002 The Linnean Society of London, Biological Journal of the Linnean Society , 2002, 77 , 149–163.     

Rapid change in host use of the common cuckoo Cuculus canorus linked to climate change

Parasites require synchrony with their hosts so if host timing changes with climate change, some parasites may decline and eventually go extinct. Residents and short-distance migrant hosts of the brood parasitic common cuckoo, Cuculus canorus, have advanced their phenology in response to climate change more than long-distance migrants, including the cuckoo itself. Because different parts of Europe show different degrees of climate change, we predicted that use of residents or short-distance migrants as hosts should have declined in areas with greater increase in spring temperature. Comparing relative frequency of parasitism of the two host categories in 23 European countries before and after 1990, when spring temperatures in many areas had started to increase, we found that relative parasitism of residents and short-distance migrants decreased. This change in host use was positively related to increase in spring temperature, consistent with the prediction that relative change in phenology for different migrant classes drives host-use patterns. These findings are consistent with the hypothesis that climate change affects the relative abundance of different host races of the common cuckoo.     

Chemical signalling behaviour in intrasexual communication of lizards lacking femoral pores

Signals that are used in animal communication may have multiple sensorial channels and functions. Animal communication integrates very distinct mechanisms such as behaviour, morphology and physiological secretions. Chemical signals occur in several contexts and are known to have a role in sexual selection. In many lizards, pheromones secreted through femoral pores are used to attract females and demark territory dominance. In lizard species without femoral pores, however, study of chemical signalling is negligible. Lizards of the genus Tropidurus have no ventral pores, but express melanic patches in their ventral thighs and cloacae (body regions linked to chemical communication in other lizards), which may play a role in both visual signalling and chemical signalling. Here, we describe the occurrence of pelvic rubbing—a chemical signalling behaviour—in two Tropidurus species, displayed in intraspecific agonistic intrasexual staged encounters. In addition, we compile a list of currently reported species that display this behaviour, checking for its social contexts and presence/absence of secretion pores. We analysed behaviours in conspecific trials between males of Tropidurus semitaeniatus and of T. hispidus during their breeding season. In T. semitaeniatus, displays of pelvic rubbing elicited aggressive responses from the opponents. Tropidurus hispidus, however, did not react to these conspecific displays. Pelvic rubbings were also partially linked to defecation in both species, which is likely due to secretions from internal urodeal glands expelled during defecation. Also, high rates of tongue flick behaviours during the encounters support the hypothesis that these lizards make use of chemical communication. Our observations corroborate previous suggestions that glandular scales — scales covered by generation glands produced at the epidermis across the lizards’ shedding cycles — are present in their melanic ventral patches. Our work also highlights the potential of characterizing such ventral patches as multimodal signalling badges. Further, we investigated the contexts in which pelvic rubbing is displayed adding to our understanding of the role played by chemical signalling in lizards without femoral pores.     

Male post-breeding movements and stopover habitat selection of an endangered short-distance migrant,the Little Bustard Tetrax tetrax

Migratory decisions, such as the selection of stopover sites, are critical for the success of post-breeding migratory movements and subsequent survival. Recent advances in bio-logging have revealed the stopover strategies of many long-distance migrants, but far less attention has been given to short-distance migrants. We investigated the stopover ecology of an endangered grassland bird, the Little Bustard Tetrax tetrax, a short-distance migrant in Iberia. Using high-resolution spatial GPS/GSM data, 27 male Little Bustards breeding in southern Portugal were tracked between 2009 and 2011. We studied post-breeding movements using Dynamic Brownian Bridge models to identify the main stopover sites, and generalized linear mixed models to examine habitat selection in stopovers. During their post-breeding movements, males were essentially nocturnal migrants, making frequent stopovers while maintaining a relatively fast pace to reach more productive agricultural post-breeding areas. Stopovers occurred in most post-breeding movements (83%) regardless of the total distance covered (average 64.3 km), and most stopovers (84%) lasted less than 24 h. Birds used mostly agricultural non-irrigated and irrigated croplands as stopover sites and avoided other land uses and rugged terrain. There was a negative relationship between stopovers and the proximity to roads, but not to power lines. The high frequency of stopovers during post-breeding movements, despite the short distances travelled, together with the nocturnal migratory behaviour of bustards, may impose additional risks to a bird mainly threatened by collision with power lines in non-breeding areas. We also conclude that even for short-distance migrants, habitat connectivity between breeding and post-breeding areas is likely to be a key conservation concern.     

Dragon wars: Movement‐based signalling by Australian agamid lizards in relation to species ecology

It is evident that the environment has the potential to affect animal communication strategies. Species from diverse taxonomic groups using signals from different modalities are known to generate signals that suit the structure of their habitat in order to maximize efficiency. Studies of acoustically communicating species dominate the literature, but visual signals are also tailored to local conditions. There is now increasing evidence that dynamic visual signals, in the form of movement‐based displays, are also influenced by habitat characteristics. Australia's dragon lizards (Family: Agamidae) employ such dynamic signals in a variety of contexts but are particularly common in territory defence. With a few notable exceptions, the signalling behaviour of this group has been relatively overlooked, and the knowledge that does exist is contained in scientific papers focused on other topics or unpublished accounts from herpetologists. In this review, we collated information on the signalling behaviour of these animals and determined that 34 of the 78 species use movement‐based signalling. This number is likely to be an underestimate, as knowledge of the signalling behaviour of many species is lacking. The richly contrasting environments of Australia inhabited by these lizards provide considerable variation in ecological context, so our second objective was to place known signalling behaviour in the context of species ecology. After controlling for phylogeny, we found that broad habitat classifications do not strongly influence the likelihood of motion signalling, and specific motor patterns are not more likely to occur in particular microhabitats. We conclude by suggesting that fully understanding the motion signalling behaviour of Australia's agamids will require documenting the displays of species for which there are no data, while taking into account the high variability existing within motor patterns and considering in detail the environmental context under which signalling takes place.     

Changing tactics: dominance, territoriality, and the responses of "primary" males to competition from conditional breeders in the variegated pupfish (Cyprinodon variegatus)

Using variegated pupfish, we examined the flexibility in "primary" male tactics when dealing with the aggressive costs of competition. Analogous to conditional mating strategies, we expected primary males to exhibit one of two interrelated tactics (i.e. dominance or territoriality) in response to different numbers of competitors. In the field, competitors influenced aggression. Primary males defended territories; residents facing more intruders engaged in more chases and obtained fewer spawns per female. In the laboratory, primary males showed dominance at low density, controlling most of the aquarium. With increasing competitor numbers, primary males reduced the area controlled and defended territories. Territories occurred with intermediate to high competitor numbers and only under male-biased sex ratios. During these interactions, aggression was highest and competitors were too many for dominant males to suppress the assertions made by each subordinate to increase its rank. Relinquished control of the entire domain enabled a previously subordinate male to establish a territory in the undefended portion of the aquarium. Reduction in defended area related to a reduction in territorial males' spawning success relative to dominants. These results suggested that primary males, like conditional breeders, would adopt the tactic that enabled them to spawn despite the constraints of competition.     

Intrapopulation variation in predator-avoidance performance of Galápagos lava lizards: The interaction of sexual and natural selection

Summary The interaction of sexual and natural selection in shaping variation in defensive behavior was explored via three steps. (1) Three predictions from the hypothesis that varying responses to predation account for intrapopulation variation in locomotory performance and wariness were tested. One measure of locomotory performance and two measures of wariness were compared between lava lizards (Tropidurus albemarlensis) inhabiting sparsely (1% cover) and heavily (33% cover) vegetated areas of Isla Plaza Sur in the Galapagos Archipelago. (2) Variation in morphology was examined to identify proximate mechanisms for differences seen in (1). (3) Levels of predation were compared between males and females to test a prediction from a model explaining the observed variation in defensive behavior. Male, but not female, lizards from sparsely vegetated areas were faster than those from heavily vegetated areas. Both male and female lizards from sparsely vegetated areas were significantly warier than those from heavily vegetated areas. Multiple regression, covariance, and residual analyses identify relatively longer hindlimbs of males as the proximate cause of their greater speed over that of females, rather than differences in body size, but neither body nor hindlimb size account for the microgeographic differences in speed of males. Significantly higher predation on males provided a positive test of the major prediction from our model of selective forces in which sexual selection for territory defense by males favors short approach distances (by minimizing time away from the territory caused by erroneous flight) and leads to natural selection for their increased hindlimb size and speed.     

Prey availability affects territory size,but not territorial display behavior,in green anole lizards

The availability of food resources can affect the size and shape of territories, as well as the behaviors used to defend territories, in a variety of animal taxa. However, individuals within a population may respond differently to variation in food availability if the benefits of territoriality vary among those individuals. For example, benefits to territoriality may differ for animals of differing sizes, because larger individuals may require greater territory size to acquire required resources, or territorial behavior may differ between the sexes if males and females defend different resources in their territories. In this study, we tested whether arthropod abundance and biomass were associated with natural variation in territory size and defense in insectivorous green anole lizards, Anolis carolinensis. Our results showed that both male and female lizards had smaller territories in a habitat with greater prey biomass than lizards in habitats with less available prey, but the rates of aggressive behaviors used to defend territories did not differ among these habitats. Further, we did not find a relationship between body size and territory size, and the sexes did not differ in their relationships between food availability and territory size or behavioral defense. Together, these results suggest that differences in food availability influenced male and female territorial strategies similarly, and that territory size may be more strongly associated with variation in food resources than social display behavior. Thus, anole investment in the behavioral defense of a territory may not vary with territory quality.     

On the song resumption, polyterritorial behaviour and their population context in the Sedge Warbler Acrocephalus schoenobaenus

Mating tactics in species with facultative polygyny seem to be very flexible and to depend on local environmental conditions. We analysed the habitat and population contexts of territorial behaviour, associated with polygyny, in a population of Sedge Warblers Acrocephalus schoenobaenus inhabiting natural wetlands. Nearly one-third of all breeding males (46 individuals) resumed song after completing their mating with the first female, in order to attract another one. Resuming males were usually the earliest arrivals. There was a continuity in the territorial behaviour between resuming song on first territory and polyterritorial behaviour. Fifty-nine percent of resuming males set up second territories, clearly separated from the first. The second territory was usually located close to the primary female activity area. The quality of the first and second territories were correlated, with second territories being significantly inferior. However, their quality was not significantly different from the territories of non-breeding males. The quality of the second territories was also negatively related to the distance from the first territories. Polyterritorialism was influenced by population numbers: the higher the number of territorial males, the lower the number of second territories settled and the lower their average quality. Although the polygyny frequency in the studied population was very low, polyterritorial males were significantly more likely to be polygynous. We concluded that polygyny frequency can be significantly influenced by population numbers, which might be one of the main factors responsible for the variability in the mating system in this species.     

Experimental evidence for female choice and energetic costs of male tail elongation in red-collared widowbirds

The black nuptial plumage of the highly polygynous male red‐collared widowbird (Euplectes ardens) comprises a red carotenoid‐based collar patch and a long graduated tail (c. 22 cm). Tail length was the strongest predictor of male mating success in a previous selection analysis, motivating this experimental test of the relative importance of tail plumes in male contest competition and female choice. Males were assigned to either a short (12.5 cm) or control (20 cm) tail manipulation prior to territory establishment. Male contest competition was unaffected by the tail treatments as the shortened‐ and control‐tailed males were equally successful in acquiring territories of similar size and quality. In contrast, however, although the longer‐tailed control males spent less time in flight and courtship displays, they attracted significantly more prospecting and nearly three times as many nesting females to their territories compared to the short‐tailed males. In further support of tail length as the primary mate choice cue, none of the other measured and potential female cues (e.g. body size, collar colorimetrics, territorial behaviours or territory quality) influenced male reproductive success. In addition to potentially increasing detectability (‘signal efficacy’), the long tail is also a likely indicator of male quality (‘signal content’). Despite the higher activities of short‐tailed males, control‐tailed males showed a steeper decline in condition (relative body mass) during the breeding season. Furthermore, both short‐ and control‐tailed residents lost more condition than did the short‐ and control‐treated floaters (males not establishing territories), suggesting an interaction between tail length and the costs of territory acquisition, defence and courtship displays. These results confirm the role of mate choice and honest quality advertising as the main selection pressures behind elongated tails in widowbirds. © 2005 The Linnean Society of London, Biological Journal of the Linnean Society, 2005, 86 , 35–43.     

The Costs of Neighbors for a Territorial Dragonfly, Perithemis tenera

Past researchers have often considered neighbors to be beneficial to territorial residents, particularly compared with non‐neighbor conspecific competitors. However, neighbors have the potential to be costly to residents in terms of both defensive costs and lost resources. In this study, we assessed the relative costs of defending a mating territory against neighbors and non‐neighbors for the dragonfly Perithemis tenera, comparing across males with different numbers of contiguous neighbors; we also examined the possibility that the presence of contiguous neighbors might reduce the detection of potential mates. When neighbors were present, residents experienced a greater total number of intrusions by males; this increase in intrusions was due to higher numbers of intrusions by neighbors, as the number of intrusions by non‐neighbor males did not differ. Residents with immediately adjacent neighbors also made more sorties toward neighbors than did residents whose nearest neighbors’ territories were not immediately adjacent. Interestingly, although the number of visits by females did not vary with the presence of neighbors, residents with neighbors made fewer sorties toward females than did residents without neighbors. Our results suggest that defensive costs increased when neighbors were present, that residents with neighbors may have missed opportunities to acquire mates, and thus that living with neighbors can be costly in this species.     

Individual-based spatially-explicit model of an herbivore and its resource: the effect of habitat reduction and fragmentation

We present an individual-based, spatially-explicit model of the dynamics of a small mammal and its resource. The life histories of each individual animal are modeled separately. The individuals can have the status of residents or wanderers and belong to behaviorally differing groups of juveniles or adults and males or females. Their territory defending and monogamous behavior is taken into consideration. The resource, green vegetation, grows depending on seasonal climatic characteristics and is diminished due to the herbivore's grazing. Other specifics such as a varying personal energetic level due to feeding and starvation of the individuals, mating preferences, avoidance of competitors, dispersal of juveniles, as a result of site overgrazing, etc., are included in the model. We determined model parameters from real data for the species Microtus ochrogaster (prairie vole). The simulations are done for a case of an enclosed habitat without predators or other species competitors. The goal of the study is to find the relation between size of habitat and population persistence. The experiments with the model show the populations go extinct due to severe overgrazing, but that the length of population persistence depends on the area of the habitat as well as on the presence of fragmentation. Additionally, the total population size of the vole population obtained during the simulations exhibits yearly fluctuations as well as multi-yearly peaks of fluctuations. This dynamics is similar to the one observed in prairie vole field studies.     

Mechanisms and fitness effects of interspecific information use between migrant and resident birds

Interactions with potential competitors are an important componentof habitat quality. Due to the costs of coexistence with competitors,a breeding habitat selection strategy that avoids competitorsis expected to be favored. However, many migratory birds appearto gain benefits from an attraction to the presence of residentbirds, even though residents are assumed to be competitivelydominant. Thus far the mechanisms of this habitat selectionprocess, heterospecific attraction, are unknown, and the consequencesfor resident birds of migrant attraction remain untested. Throughheterospecific attraction, migrants may gain benefits if thedensity or territory location of residents positively reflectshabitat quality, and/or they gain benefits through increasedfrequency of social interactions with residents in foragingor predator detection. In this experiment, we examined the reciprocaleffects of spatial proximity on fitness-related traits in migrantpied flycatcher (Ficedula hypoleuca) and resident great tit(Parus major) by experimentally forcing them to breed eitheralone or in close proximity to each other. Surprisingly, greattits bore all the costs of coexistence while flycatchers wereunaffected, even gaining slight benefits. In concert with anearlier study, these results suggest that flycatchers use titsas information about good-quality nest-site locations whilebenefits from social interactions with tits are possible butless important. We suggest that utilizing interspecific socialinformation may be a common phenomenon between species sharingsimilar resource needs. Our results imply that the effects ofinterspecific information use can be asymmetric and may thereforehave implications for the patterns and consequences of speciescoexistence.     

Long-distance migrating species of birds travel in larger groups

How individuals migrate over long distances is an enduring mystery of animal migration. Strong selection pressure for travelling in groups has been suggested in long-distance migrating species. Travelling in groups can reduce the energetic demands of long migration, increase navigational accuracy and favour group foraging at migratory halts. Nevertheless, this hypothesis has received scant attention. I examined evolutionary transitions in migration distance in all North American breeding species of birds. I documented 72 evolutionary shifts in migration distance in the pool of 409 species. In contrasting clades, long-distance migration, as opposed to short-distance migration, was associated with a larger travelling group size. No other transitions occurred alongside in other traits such as group size in the non-breeding season or body mass. The results suggest that larger group sizes have been beneficial in the evolution of long-distance migration in a large clade of birds.