Macroscopic and histological investigation of guanaco footpads (Lama guanicoe,Müller 1776)

The surface of guanaco footpads is characterized by hairless skin with up to 4‐mm‐thick stratum corneum that protects from abrasion. The horny layer is pliable and elastic, and ensures firm contact with irregular ground. It is padded with a particular structure of the subcutaneous layer, the digital cushion. The flat cushions of each of the two digits are of elongated ovate shape, each about 45‐mm long, up to 20‐mm wide, and 8‐mm thick. The cushions are lined by a 1–2‐mm‐thick capsule that resembles a tunica albuginea. The capsule consists of coarse collagen fibers, with elastic fibers absent. The cushion capsule and dermis approach each other, and fuse along a line that runs parallel to the longitudinal axes of cushion and digit. Loose connective tissue rich in elastic fibers and acidic glycosaminoglycans separates dermis and cushion capsule lateral to the narrow interconnecting zone. The cushion capsule encloses cloudy yellowish, gelatinous material. Microscopy shows bundles of elastic fibers in abundant mucinous matrix. Tightly gathered elastic bundles adjoin the inner surface of the capsule. Rough cords of elastic fibers branch out from there and traverse to the opposite side. The cushion is pressed flat, and elastic fibers are stretched when bearing weight. With relief of load, elastic fibers contract and reset the cushion's shape. Contractile cells are absent. A resistant capsule and easily malleable mucinous contents establish the functioning as a gel pad. Mucinous connective tissue between elastic fiber bundles contains abundant basophilic matrix. Hyaluronan, chondroitin sulfate, and dermatan sulfate are main matrix constituents. Spindle‐shaped or stellate fibroblasts contain vimentin, S100 protein, and neuron specific enolase. Moprhology, staining characteristics and synthesis activities of these cells meet the criteria to be classified as myxoid cells. The connective tissue in guanaco digital cushions represents myxoid tissue. J. Morphol. 276:331–341, 2015. © 2014 Wiley Periodicals, Inc.     

Predator behaviour and predation risk in the heterogeneous Arctic environment

1. Habitat heterogeneity and predator behaviour can strongly affect predator-prey interactions but these factors are rarely considered simultaneously, especially when systems encompass multiple predators and prey. 2. In the Arctic, greater snow geese Anser caerulescens atlanticus L. nest in two structurally different habitats: wetlands that form intricate networks of water channels, and mesic tundra where such obstacles are absent. In this heterogeneous environment, goose eggs are exposed to two types of predators: the arctic fox Vulpes lagopus L. and a diversity of avian predators. We hypothesized that, contrary to birds, the hunting ability of foxes would be impaired by the structurally complex wetland habitat, resulting in a lower predation risk for goose eggs. 3. In addition, lemmings, the main prey of foxes, show strong population cycles. We thus further examined how their fluctuations influenced the interaction between habitat heterogeneity and fox predation on goose eggs. 4. An experimental approach with artificial nests suggested that foxes were faster than avian predators to find unattended goose nests in mesic tundra whereas the reverse was true in wetlands. Foxes spent 3.5 times more time between consecutive attacks on real goose nests in wetlands than in mesic tundra. Their attacks on goose nests were also half as successful in wetlands than in mesic tundra whereas no difference was found for avian predators. 5. Nesting success in wetlands (65%) was higher than in mesic tundra (56%) but the difference between habitats increased during lemming crashes (15%) compared to other phases of the cycle (5%). Nests located at the edge of wetland patches were also less successful than central ones, suggesting a gradient in accessibility of goose nests in wetlands for foxes. 6. Our study shows that the structural complexity of wetlands decreases predation risk from foxes but not avian predators in arctic-nesting birds. Our results also demonstrate that cyclic lemming populations indirectly alter the spatial distribution of productive nests due to a complex interaction between habitat structure, prey-switching and foraging success of foxes.     

Aggressive behavior and dominance hierarchies in female sable antelope,Hippotragus niger: Implications for captive management

Social interactions of adult female sable antelope, Hippotragus niger, were observed at the National Zoological Park's Conservation and Research Center. Aggressive behavior was quantified for two herds containing 8 and 11 females. Aggression was lowest at calving, and highest near the time of conception. Dominance hierarchies were strictly linear and stable over time. Dominance rank was highly correlated with age, with the oldest females achieving highest rank. High-ranking females initiated more aggression and were more likely to behave aggressively towards a newly introduced adult female. The newly introduced individual was the victim of prolonged, intense group chases and was permanently excluded from the herd. Low-ranking females directed more aggression at yearling herd members than did high-ranking females. Temporary absences by herd members resulted in long-lasting, substantial increases in received aggression upon their return. Because of their aggressive nature, sable antelope present a challenge to captive management. © 1993 Wiley-Liss, Inc.     

The influence of the arid Andean high plateau on the phylogeography and population genetics of guanaco (Lama guanicoe) in South America

A comprehensive study of the phylogeography and population genetics of the largest wild artiodactyl in the arid and cold‐temperate South American environments, the guanaco (Lama guanicoe) was conducted. Patterns of molecular genetic structure were described using 514 bp of mtDNA sequence and 14 biparentally inherited microsatellite markers from 314 samples. These individuals originated from 17 localities throughout the current distribution across Peru, Bolivia, Argentina and Chile. This confirmed well‐defined genetic differentiation and subspecies designation of populations geographically separated to the northwest (L. g. cacsilensis) and southeast (L. g. guanicoe) of the central Andes plateau. However, these populations are not completely isolated, as shown by admixture prevalent throughout a limited contact zone, and a strong signal of expansion from north to south in the beginning of the Holocene. Microsatellite analyses differentiated three northwestern and 4–5 southeastern populations, suggesting patterns of genetic contact among these populations. Possible genetic refuges were identified, as were source‐sink patterns of gene flow at historical and recent time scales. Conservation and management of guanaco should be implemented with an understanding of these local population dynamics while also considering the preservation of broader adaptive variation and evolutionary processes.     

Mesopredator suppression by an apex predator alleviates the risk of predation perceived by small prey

Predators can impact their prey via consumptive effects that occur through direct killing, and via non-consumptive effects that arise when the behaviour and phenotypes of prey shift in response to the risk of predation. Although predators'' consumptive effects can have cascading population-level effects on species at lower trophic levels there is less evidence that predators'' non-consumptive effects propagate through ecosystems. Here we provide evidence that suppression of abundance and activity of a mesopredator (the feral cat) by an apex predator (the dingo) has positive effects on both abundance and foraging efficiency of a desert rodent. Then by manipulating predators'' access to food patches we further the idea that apex predators provide small prey with refuge from predation by showing that rodents increased their habitat breadth and use of ‘risky′ food patches where an apex predator was common but mesopredators rare. Our study suggests that apex predators'' suppressive effects on mesopredators extend to alleviate both mesopredators'' consumptive and non-consumptive effects on prey.     

Geographic distribution and possible taxonomic distinction of Callicebus torquatus populations (Pitheciidae: Primates) in Peruvian Amazonia

Population densities of yellow-handed titi monkey (Callicebus torquatus) were estimated using transect census methods. Densities were 2.8 individuals/km(2 )in the upper Rio Itaya basin and 2.5 individuals/km(2) in the lower Rio Algodón basin. Group size varied from two to five individuals, with an average of 2.9 individuals per group at the Rio Itaya. Groups were generally composed of two adults, probably the reproductive pair, with progeny from one to three previous birth seasons. Although the Rio Itaya population is phenotypically identical to populations from the rivers Nanay and Tigre, it differs from population on the rivers Napo and Putumayo. This suggests the existence of two disjunct populations of C. torquatus in Peruvian Amazonia whose taxonomic status warrants further examination.     

Effects of predation risk on the grouping patterns of white‐bellied spider monkeys (Ateles belzebuth belzebuth) in Western Amazonia

Predation is proposed to be one of the most important factors influencing the evolution of mammalian societies. Although predation risk is thought to influence both the behavior and grouping patterns of most diurnal primates, evidence supporting this hypothesis is still limited. The spatial and temporal patterns of mineral lick use by one group of white‐bellied spider monkeys (Ateles belzebuth) were evaluated, based on the growing evidence that mineral licks are perceived as areas of relative high predation risk by Neotropical primates. The area immediately surrounding the mineral lick was the most intensively used area within the home range of the study group, particularly by large subgroups of monkeys, and there were differences in mean subgroup size on days of mineral lick visitation versus days without lick visits. Additionally, on days of mineral lick visitation, subgroup size reached its maximum specifically during the period of lick visitation. Finally, on visit days subgroups showed a greater increase in size and higher fusion rates in the 2 hr before arriving at the lick in comparison with matched time windows on non‐visit days. Together, these results provide an example of how primates employ behavioral strategies that might reduce the effects of predation. This study also demonstrates how taxa characterized by a high degree of fission‐fusion dynamics can allow us to test hypotheses regarding the effects of socioecological variables on primate grouping patterns. Am J Phys Anthropol 150:579–590, 2013. © 2013 Wiley Periodicals, Inc.     

The landscape of fear as an emergent property of heterogeneity: Contrasting patterns of predation risk in grassland ecosystems

The likelihood of encountering a predator influences prey behavior and spatial distribution such that non‐consumptive effects can outweigh the influence of direct predation. Prey species are thought to filter information on perceived predator encounter rates in physical landscapes into a landscape of fear defined by spatially explicit heterogeneity in predation risk. The presence of multiple predators using different hunting strategies further complicates navigation through a landscape of fear and potentially exposes prey to greater risk of predation. The juxtaposition of land cover types likely influences overlap in occurrence of different predators, suggesting that attributes of a landscape of fear result from complexity in the physical landscape. Woody encroachment in grasslands furnishes an example of increasing complexity with the potential to influence predator distributions. We examined the role of vegetation structure on the distribution of two avian predators, Red‐tailed Hawk (Buteo jamaicensis) and Northern Harrier (Circus cyaneus), and the vulnerability of a frequent prey species of those predators, Northern Bobwhite (Colinus virginianus). We mapped occurrences of the raptors and kill locations of Northern Bobwhite to examine spatial vulnerability patterns in relation to landscape complexity. We use an offset model to examine spatially explicit habitat use patterns of these predators in the Southern Great Plains of the United States, and monitored vulnerability patterns of their prey species based on kill locations collected during radio telemetry monitoring. Both predator density and predation‐specific mortality of Northern Bobwhite increased with vegetation complexity generated by fine‐scale interspersion of grassland and woodland. Predation pressure was lower in more homogeneous landscapes where overlap of the two predators was less frequent. Predator overlap created areas of high risk for Northern Bobwhite amounting to 32% of the land area where landscape complexity was high and 7% where complexity was lower. Our study emphasizes the need to evaluate the role of landscape structure on predation dynamics and reveals another threat from woody encroachment in grasslands.     

Effects of perceived predation risk and social environment on the development of three‐spined stickleback (Gasterosteus aculeatus) morphology

Phenotypically plastic changes in response to variation in perceived predation risk are widespread, but little is known about if and how social environment modulates induced responses to predation risk. We investigated the influence of perceived predation risk (i.e. chemical cues from a predator) and social environment (i.e. one, two or 20 individuals reared together) on three‐spined stickleback (Gasterosteus aculeatus) morphology in a factorial common garden experiment. We found that exposure to chemical cues from potential predators did not influence growth or body condition or induce more robust morphological defences (i.e. lateral plate numbers and dorsal spine lengths). However, sticklebacks exposed to predator cues developed longer caudal peduncles and larger eyes as compared with fish from the control treatment. As these responses may improve sticklebacks’ ability to avoid piscine predation, they might be adaptive. Social environment/density also influenced expression of some traits, but these effects were independent of predation‐risk treatment effects. In general, these results suggest that apart from the classic morphological defence structures, which appear mostly constitutive, three‐spined sticklebacks are capable of expressing potentially adaptive morphological responses to chemical cues from potential predators.     

Habitat use and movements of plains zebra (Equus burchelli) in response to predation danger from lions

Prey species must adapt their behavior to avoid predation. Asa key prey item for lions (Panthera leo), plains zebras (Equusburchelli) were expected to respond to immediate threats posedby lions in their area. In addition, zebras were predicted toexhibit behavior tuned to reduce the potential for encounterswith lions, by modifying their movement patterns in the timesof day and habitats of greatest lion danger. We studied a populationof approximately 600 plains zebra living in Ol Pejeta Conservancy,Kenya. We found that zebra abundance on or near a grasslandpatch was lower if lions had also been observed on that patchduring the same day. Predation danger was highest in grasslandhabitat during the night, when lions were more active. Zebrasightings and global positioning system radio collar data indicatedthat zebras also reduced their use of grassland at night, insteadusing more woodland habitat. Zebras moved faster and took sharperturns in grassland at night. It is hypothesized that these moreerratic movements assist zebras in avoiding detection or captureby lions.     

Habitat structure and association with ungulates drive habitat selection and grouping behaviour of lesser rhea (Rhea pennata subsp. pennata)

Social prey species respond to predation risk by modifying habitat selection and grouping behaviour. These responses may depend on both actual predation risk (predator probability of occurrence) and/or on perceived predation risk associated with habitat structure. Other factors like food availability and co-occurrence with other species may also affect habitat selection and group formation. We analyse habitat selection and grouping behaviour (group size and cohesion) of lesser rhea (Rhea pennata subsp. pennata), a ratite endemic of South America inhabiting steppe shrublands and grasslands, in relation to actual (puma probability of occurrence) and perceived (habitat structure: openness, visibility) predation risk, co-occurrence with other herbivore species and forage availability in the Chilean Patagonia. We used data from 9 sampling seasons in 5 years. Results show that habitat selection, group size and cohesion in lesser rhea were mainly driven by variables associated with perceived predation risk and by co-occurrence with other herbivores both during breeding and non–breeding season. As expected, lesser rhea preferred open habitats (vegas and grasslands) that allow a behaviour of ‘watch and run’ to avoid predation and formed larger groups in them. Moreover, lesser rhea positively selected year-round habitats where livestock occur, forming large groups during non–breeding season there. Group size and co-occurrence with other herbivores significantly decreased group cohesion, suggesting a reduction of perceived predation risk. Therefore, lesser rhea seems to take advantage of forming mixed interspecific groups to reduce predation risk. These results suggest that lesser rhea habitat selection and grouping behaviour are preferentially driven by factors related to perceived predation risk than by actual predator occurrence or food availability.     

Social information, social feeding, and competition in group-living goats (Capra hircus)

There are both benefits (e.g., social information) and costs(e.g., intraspecific competition) for individuals foraging ingroups. To ascertain how group-foraging goats (Capra hircus)deal with these trade-offs, we asked 1) do goats use socialinformation to make foraging decisions and 2) how do they adjusttheir intake rate in light of having attracted by other groupmembers? To establish whether goats use social information,we recorded their initial choice of different quality food patcheswhen they were ignorant of patch quality and when they couldobserve others foraging. After determining that goats use socialinformation, we recorded intake rates while they fed alone andin the presence of potential competitors. Intake rate increasedas the number of competitors increased. Interestingly, lonegoats achieved an intake rate that was higher than when onecompetitor was present but similar to when two or more competitorswere present. Faster intake rates may allow herbivores to ingesta larger portion of the available food before competing groupmembers arrive at the patch. This however, does not explainthe high intake rates achieved when the goats were alone. Weprovide 2 potential explanations: 1) faster intake rates area response to greater risk incurred by lone individuals, theloss of social information, and the fear of being left behindby the group and 2) when foraging alone, intake rate is no longera trade-off between reducing competition and acquiring socialinformation. Thus, individuals are able to feed close to theirmaximum rate.     

Habitat use and social structure of an isolated population of guanacos (Lama guanicoe) in the Monte Desert, Argentina

We studied abundance, habitat use and social structure of an isolated population of guanacos in the Lihue Calel National Park in central Argentina, based on data from foot surveys during 1998–2000. Based on topography and vegetation characteristics we distinguished three types of habitat: hills, valleys and piedmonts, and lowlands. Three types of social groups were observed: solitary males, family groups and male groups. The guanacos used hills preferentially, followed by valley and piedmont. The guanacos avoided the lowlands where the vegetation is dense. Habitat selection, both on a coarse and a fine scale, could be explained by habitat quality and possibly also predator avoidance behaviour. Abundance of guanacos increased by 21% over the 3-year-study period. However, the specificity of habitat requirements of guanacos and the susceptibility of the study area to wildfires could cause guanacos to move into sub-optimal areas were they are more vulnerable to predation and human persecution.     

Human activity and habitat type affect perceived predation risk in urban white‐footed mice (Peromyscus leucopus)

Predation risk is one of the largest costs associated with foraging in small mammals. Small mammals select microhabitat features such as tree and shrub canopy cover, woody debris and vegetative ground cover that can lower the risk of detection from predators and provide greater protection if discovered. Small mammals also increase foraging activity and decrease selection for cover when cloud cover increases and moon illumination is less. Often researchers assume small mammals in urban areas respond to these cues in the same manner as in natural areas, but these cues themselves are altered in urban zones. In this study, we investigated how Amur honeysuckle (Lonicera maackii) and coarse woody debris (CWD) affected giving‐up density (GUD) in white‐footed mice (Peromyscus leucopus). Each of three habitat treatments (open flood channel, the edge and interior of the honeysuckle patch) contained cover treatments with coarse woody debris present or absent. The six treatment combinations were compared to environmental variables (temperature, humidity and illumination) and habitat variables to test their effect on GUD. Peromyscus leucopus foraged to lower densities in areas with CWD present and also under the honeysuckle canopy, using this invasive shrub to decrease predation risk, potentially increasing survivability within this urban park. Increased human presence negatively affected foraging behavior across treatments. Human presence and light pollution significantly influenced P. leucopus, modifying their foraging behavior and demonstrating that both fine‐ and coarse‐scale urban factors can affect small mammals. Foraging increased as humidity increased, particularly under the honeysuckle canopy. Changes in illumination due to moonlight and cloud cover did not affect foraging behavior, suggesting urban light pollution may have altered behavioral responses to changes in light levels. Lonicera maackii seemed to facilitate foraging in P. leucopus, even though it adversely affects the plant community, suggesting that its impact may not be entirely negative.