Adaptive latitudinal variation in Common Blackbird Turdus merula nest characteristics

Nest construction is taxonomically widespread, yet our understanding of adaptive intraspecific variation in nest design remains poor. Nest characteristics are expected to vary adaptively in response to predictable variation in spring temperatures over large spatial scales, yet such variation in nest design remains largely overlooked, particularly amongst open‐cup‐nesting birds. Here, we systematically examined the effects of latitudinal variation in spring temperatures and precipitation on the morphology, volume, composition, and insulatory properties of open‐cup‐nesting Common Blackbirds’ Turdus merula nests to test the hypothesis that birds living in cooler environments at more northerly latitudes would build better insulated nests than conspecifics living in warmer environments at more southerly latitudes. As spring temperatures increased with decreasing latitude, the external diameter of nests decreased. However, as nest wall thickness also decreased, there was no variation in the diameter of the internal nest cups. Only the mass of dry grasses within nests decreased with warmer temperatures at lower latitudes. The insulatory properties of nests declined with warmer temperatures at lower latitudes and nests containing greater amounts of dry grasses had higher insulatory properties. The insulatory properties of nests decreased with warmer temperatures at lower latitudes, via changes in morphology (wall thickness) and composition (dry grasses). Meanwhile, spring precipitation did not vary with latitude, and none of the nest characteristics varied with spring precipitation. This suggests that Common Blackbirds nesting at higher latitudes were building nests with thicker walls in order to counteract the cooler temperatures. We have provided evidence that the nest construction behavior of open‐cup‐nesting birds systematically varies in response to large‐scale spatial variation in spring temperatures.     

Ants on the Move: Resource Limitation of a Litter‐nesting Ant Community in Costa Rica1

The leaf litter of tropical wet forests is replete with itinerant ant nests. Nest movement may help ants evade the constraints of stress and disturbance and increase access to resources. I studied how nest relocation and environmental factors may explain the density, size, and growth of leaf litter ant nests. I decoupled the relationships among litter depth, food abundance, and nest availability in a 4‐mo manipulation of food and leaf litter in a community of litter‐nesting ants in a lowland wet forest in Costa Rica. Over 4 mo, 290 1 m² treatment and control plots were sampled without replacement. Nest densities doubled in response to food supplementation, but did not decrease in response to litter removal or stress (from litter trampling). The supplementation of food increased the utilization of less favored nesting materials. In response to food supplementation and litter trampling, arboreal ants established nests in the litter, and growth rates of the most common ants (Pheidole spp.) increased. Colony growth was independent of colony size and growth rates of the most abundant ants. In general, I conclude that litter‐nesting ant density is driven primarily by food limitation, that nest relocation behavior significantly affects access to resource and the demographic structure of this community, and that nest fission may be a method to break the growth–reproduction trade‐off.     

Intraseasonal patterns in shorebird nest survival are related to nest age and defence behaviour

Nest survival may vary throughout the breeding season for many bird species, and the nature of this temporal variation can reveal the links between birds, their predators, and other components of the ecosystem. We used program Mark to model patterns in nest survival within the breeding season for shorebirds nesting on arctic tundra. From 2000 to 2007, we monitored 521 nests of five shorebird species and found strong evidence for variation in nest survival within a nesting season. Daily nest survival was lowest in the mid-season in 5 of 8 years, but the timing and magnitude of the lows varied. We found no evidence that this quadratic time effect was driven by seasonal changes in weather or the abundance of predators. Contrary to our prediction, the risk of predation was not greatest when the number of active shorebird nests was highest. Although nest abundance reached a maximum near the middle of the breeding season, a daily index of shorebird nest activity was not supported as a predictor of nest survival in the models. Predators’ access to other diet items, in addition to shorebird nests, may instead determine the temporal patterns of nest predation. Nest survival also displayed a positive, linear relationship with nest age; however, this effect was most pronounced among species with biparental incubation. Among biparental species, parents defended older nests with greater intensity. We did not detect a similar relationship among uniparental species, and conclude that the stronger relationship between nest age and both nest defence and nest survival for biparental species reflects that their nest defence is more effective.     

Fitness consequences of ecological constraints and implications for the evolution of sociality in an incipiently social bee

Ecological constraints such as resource limitation, unfavourable weather conditions, and parasite pressure have long been considered some of the most important selective pressures for the evolution of sociality. In the present study, we assess the fitness consequences of these three ecological factors on reproductive success of solitary nests and social colonies in the socially polymorphic small carpenter bee, Ceratina australensis, based on 982 nests collected over four reproductive periods. Nest site limitation was predicted to decrease opportunities for independent nest initiation and increase the frequency of social nesting. Nest sites were not limiting in this species and the frequency of social nesting was consistent across the four brood‐rearing periods studied. Unfavourable weather was predicted to lower the frequency of female dispersal from their natal nests and to limit the brood‐rearing season; this would increase the frequency and fitness of social colonies. Daily temperature and precipitation accumulation varied between seasons but were not correlated with reproductive success in this bee. Increased parasite pressure is predicted to increase the frequency and fitness of social colonies because solitary bees must leave the nest unattended during foraging bouts and are less able to defend the nest against parasites. Severe parasitism by a chalcid wasp (Eurytoma sp.) resulted in low reproductive success and total nest failure in solitary nests. Social colonies had higher reproductive success and were never extirpated by parasites. The high frequency of solitary nests suggests that this is the optimal strategy. However, social colonies have a selective advantage over solitary nesting females during periods of extreme parasite pressure, and we suggest that social nesting represents a form of bet‐hedging against unpredictable fluctuations in parasite number. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 103 , 57–67.     

Conditional female strategies influence hatching success in a communally nesting iguana

The decision of females to nest communally has important consequences for reproductive success. While often associated with reduced energetic expenditure, conspecific aggregations also expose females and offspring to conspecific aggression, exploitation, and infanticide. Intrasexual competition pressures are expected to favor the evolution of conditional strategies, which could be based on simple decision rules (i.e., availability of nesting sites and synchronicity with conspecifics) or on a focal individual's condition or status (i.e., body size). Oviparous reptiles that reproduce seasonally and provide limited to no postnatal care provide ideal systems for disentangling social factors that influence different female reproductive tactics from those present in offspring‐rearing environments. In this study, we investigated whether nesting strategies in a West Indian rock iguana, Cyclura nubila caymanensis, vary conditionally with reproductive timing or body size, and evaluated consequences for nesting success. Nesting surveys were conducted on Little Cayman, Cayman Islands, British West Indies for four consecutive years. Use of high‐density nesting sites was increasingly favored up to seasonal nesting activity peaks, after which nesting was generally restricted to low‐density nesting areas. Although larger females were not more likely than smaller females to nest in high‐density areas, larger females nested earlier and gained access to priority oviposition sites. Smaller females constructed nests later in the season, apparently foregoing investment in extended nest defense. Late‐season nests were also constructed at shallower depths and exhibited shorter incubation periods. While nest depth and incubation length had significant effects on reproductive outcomes, so did local nest densities. Higher densities were associated with significant declines in hatching success, with up to 20% of egg‐filled nests experiencing later intrusion by a conspecific. Despite these risks, nests in high‐density areas were significantly more successful than elsewhere due to the benefits of greater chamber depths and longer incubation times. These results imply that communal nest sites convey honest signals of habitat quality, but that gaining and defending priority oviposition sites requires competitive ability.     

Testing hypotheses about the function of repeated nest abandonment as a life history strategy in a passerine bird

Nest structures are essential for successful reproduction in most bird species. Nest construction costs time and energy, and most bird species typically build one nest per breeding attempt. Some species, however, build more than one nest, and the reason for this behaviour is often unclear. In the Grey Fantail Rhipidura albiscapa, nest abandonment before egg‐laying is very common. Fantails will build up to seven nests within a breeding season, and pairs abandon up to 71% of their nests before egg‐laying. We describe multiple nest‐building behaviour in the Grey Fantail and test four hypotheses explaining nest abandonment in this species: cryptic depredation, destruction of nests during storm events, and two anti‐predatory responses (construction of decoy nests to confuse predators, and increasing concealment to ‘hide’ nests more effectively). We found support for only one hypothesis – that abandonment is related to nest concealment. Abandoned nests were significantly less concealed than nests that received eggs. Most abandoned nests were not completely built and none received eggs, thus ruling out cryptic predation. Nests were not more likely to be abandoned following storm events. The decoy nest hypothesis was refuted as abandoned nests were constructed at any point during the breeding season and some nests were dismantled and the material used to build the subsequent nest. Thus, Grey Fantails are flexible about nest‐site locations during the nest‐building phase and readily abandon nest locations if they are found to have deficient security.     

Climate change and nesting behaviour in vertebrates: a review of the ecological threats and potential for adaptive responses

Nest building is a taxonomically widespread and diverse trait that allows animals to alter local environments to create optimal conditions for offspring development. However, there is growing evidence that climate change is adversely affecting nest‐building in animals directly, for example via sea‐level rises that flood nests, reduced availability of building materials, and suboptimal sex allocation in species exhibiting temperature‐dependent sex determination. Climate change is also affecting nesting species indirectly, via range shifts into suboptimal nesting areas, reduced quality of nest‐building environments, and changes in interactions with nest predators and parasites. The ability of animals to adapt to sustained and rapid environmental change is crucial for the long‐term persistence of many species. Many animals are known to be capable of adjusting nesting behaviour adaptively across environmental gradients and in line with seasonal changes, and this existing plasticity potentially facilitates adaptation to anthropogenic climate change. However, whilst alterations in nesting phenology, site selection and design may facilitate short‐term adaptations, the ability of nest‐building animals to adapt over longer timescales is likely to be influenced by the heritable basis of such behaviour. We urgently need to understand how the behaviour and ecology of nest‐building in animals is affected by climate change, and particularly how altered patterns of nesting behaviour affect individual fitness and population persistence. We begin our review by summarising how predictable variation in environmental conditions influences nest‐building animals, before highlighting the ecological threats facing nest‐building animals experiencing anthropogenic climate change and examining the potential for changes in nest location and/or design to provide adaptive short‐ and long‐term responses to changing environmental conditions. We end by identifying areas that we believe warrant the most urgent attention for further research.     

A description of nesting behaviors,including factors impacting nest site selection,in black‐and‐white ruffed lemurs (Varecia variegata)

Nest site selection is at once fundamental to reproduction and a poorly understood component of many organisms’ reproductive investment. This study investigates the nesting behaviors of black‐and‐white ruffed lemurs, Varecia variegata, a litter‐bearing primate from the southeastern rainforests of Madagascar. Using a combination of behavioral, geospatial, and demographic data, I test the hypotheses that environmental and social cues influence nest site selection and that these decisions ultimately impact maternal reproductive success. Gestating females built multiple large nests throughout their territories. Of these, females used only a fraction of the originally constructed nests, as well as several parking locations as infants aged. Nest construction was best predicted by environmental cues, including the size of the nesting tree and density of feeding trees within a 75 m radius of the nest, whereas nest use depended largely on the size and average distance to feeding trees within that same area. Microhabitat characteristics were unrelated to whether females built or used nests. Although unrelated to nest site selection, social cues, specifically the average distance to conspecifics’ nest and park sites, were related to maternal reproductive success; mothers whose litters were parked in closer proximity to others’ nests experienced higher infant survival than those whose nests were more isolated. This is likely because nesting proximity facilitated communal crèche use by neighboring females. Together, these results suggest a complex pattern of nesting behaviors that involves females strategically building nests in areas with high potential resource abundance, using nests in areas according to their realized productivity, and communally rearing infants within a network of nests distributed throughout the larger communal territory.     

Nest reuse in arctic‐breeding shorebirds: an analysis of potential benefits and factors affecting the occurrence of this rare behavior

During a 13‐yr study near Utqia?vik (formerly Barrow), Alaska, we documented the prevalence of nest reuse in eight arctic‐breeding shorebirds. We evaluated whether nest reuse saved individuals time and energy, enhanced nest survival, or was related to nest density. We documented 208 (6.2%) cases of nest reuse among 3336 nesting attempts. Nest reuse occurred in all years but the first and in all species, with greatest reuse in semipalmated sandpiper (10.9%) and American golden‐plover (10.0%). While most cases of nest reuse occurred with conspecifics, many cases of heterospecific nest reuse were also observed, indicating high niche overlap in nest site preferences among species. We found that individuals reusing old nests may have benefited by nesting earlier, but nest reuse did not generally enhance nest survival. A significant positive relationship was also found between nest reuse and nest density at the community level and for four of the eight species, suggesting high inter‐ or intraspecific competition combined with limited suitable nest sites may force individuals to reuse old nests. Our observations also suggest that upland nesting species may be the most dependent on old nest sites. Preferential development of these sites may therefore have a previously unknown detrimental effect on these species, although further study is needed to better determine the impact of such habitat loss.     

Effect of floating nest platforms on the breeding performance of Black Terns

ABSTRACT.   In 2003 and 2004, we placed 41 floating nest platforms on Grassy Lake in southeastern Wisconsin (USA) to test the hypothesis that reproductive success of Black Terns ( Chlidonias niger ) is limited by the quality of suitable nesting habitat. Extreme differences in water levels between these 2 yr provided a natural experiment to evaluate the effectiveness of the nest platforms during a drought year (2003) when natural nesting substrate was abundant, and a flood year (2004) when natural substrate was limited during the peak nesting period. Terns nested on 27 of 41 (66%) of the platforms in 2003 and 26 of 41 (63%) in 2004. No difference in the occupancy rate of platforms and natural nests was evident in 2003, but the pattern of clutch initiations early in the season in 2004 indicated that platforms were preferred over natural substrates. In 2003, nest survival rates did not differ between nests placed on platforms and those placed on natural substrates, but platform nests had significantly higher hatching success and nest survival rates in 2004. Both the Kaplan-Meier and Apparent Nest Success methods of calculating nest survival provided similar estimates. In both years, eggs laid on platforms were significantly larger than those laid on natural substrates, suggesting that platforms were occupied by high-quality birds. Our study indicates that floating nest platforms can be an effective management tool to enhance nesting habitat for Black Terns and other aquatic birds that construct floating nests, primarily because platforms provide nest sites when natural sites are not available due to flooding. Nest platforms also may be useful for addressing questions concerning habitat selection and parental quality.     

Testing the importance of nest concealment: does timing matter?

ABSTRACT.   Nest concealment by vegetation is considered an important factor affecting predation rates for many passerines and, therefore, is frequently measured in studies examining nest predation. However, the time when concealment measurements are made may affect the results of such studies, particularly in highly seasonal ecosystems where characteristics of the vegetation later in the breeding period may differ considerably from those at the time of nest-site selection. We used artificial nests baited with quail ( Coturnix sp.) eggs in a highly seasonal tropical dry forest in Jalisco, western Mexico, to test the effects of seasonal change in concealment on nest predation. We placed 40 open-cup, artificial nests in shrubs at the end of the dry season and again at the beginning of the rainy season in 2007, and monitored the fate of the nests and the degree of concealment by vegetation during both periods. Nest concealment was significantly greater during the wet season than during the dry season. The percentage of nests predated was marginally higher during the dry (100%) than the wet (72.5%) season, and daily nest survival was lower during the dry than the wet season. Our results suggest that, in highly seasonal environments such as tropical dry forests, delayed measurement of nest concealment after nest completion rather than during nesting may constitute a significant source of error.     

Nesting habitat characteristics of Marbled Murrelets occurring in near‐shore waters of the Olympic Peninsula,Washington

Marbled Murrelets (Brachyramphus marmoratus) are listed as threatened in the portion of their range extending from British Columbia to California due to loss of nesting habitat. Recovery of Marbled Murrelet populations requires a better understanding of the characteristics of their nesting habitat in this part of their range. Our objective, therefore, was to describe their nesting habitat in Washington State and Vancouver Island, British Columbia. We captured Marbled Murrelets from 2004 to 2008, fitted them with radio transmitters, and followed them to nests (N = 20). We used Cohen's unbiased d effect size to assess differences between forest plots surrounding nest sites and nearby control sites (N = 18). Nest sites had less canopy cover of the dominant conifers and fewer, but larger, trees than control sites. Nest sites also had greater percentages of trees with platforms >10 cm diameter and >15 cm diameter, and more platforms of these sizes than control sites. The mean diameter at breast height of nest trees was 136.5 cm (range = 84–248 cm) and all but one nest was in dominant or co‐dominant tree species. At the landscape scale, we used vegetation maps derived from remotely sensed data and found greater canopy cover, higher density of mature trees, more platforms >10 cm/ha, and more old‐growth habitat at nest sites than at random sites. Our findings suggest that, at the site scale, nesting Marbled Murrelets selected the most suitable features of forest structure across expansive potentially suitable habitat. Our landscape‐scale analysis showed that habitat features in nesting stands differed from those features in available stands in the murrelet's range in Washington. We also found that stands with nests were less fragmented than available forest across murrelet range. All nest sites of radio‐tagged birds in Washington were in protected areas in mostly undisturbed forest habitat. Conservation of these areas of inland nesting habitat will be critical to the recovery of Marbled Murrelet populations.     

Nest Building in Captive Gorilla gorilla gorilla

Although various aspects of gorilla nest building have been described in wild populations, nest-building behavior of captive gorillas has not been subject to much scientific review. We observed nest building in 17 gorillas during three periods: summer baseline, winter baseline, and winter treatment, in which the amount of available nesting material was doubled. We conducted observations exclusively in the indoor holding area in the hour following evening departure of animal care staff. During baseline, gorillas engaged in nest-building on 3.1% of scans and were on a constructed nest on 27.9% of scans. Overall, gorillas spent significantly more time on elevated nests than on floor nests. There were no statistically significant sex, age class, or rearing history differences in nest building, time spent on a nest, or nest location preference. Nest building consisted of both gathering and manipulating materials. The gorillas spent significantly more time building nests in winter than in summer. Additionally, they were more often on elevated nests during winter than summer. Doubling the amount of nesting material did not increase nest building, but it increased the number of floor nests. Our results suggest that providing adequate materials to captive gorillas for nest building may facilitate performance of species-typical nest-building, minimize competition among individuals for nesting sites and materials, and permit individual thermoregulation.     

Elevation has contrasting effects on avian and mammalian nest traits in the Andean temperate mountains

Nest building is a widespread breeding strategy across taxa. Nest composition and structure can play a critical role in the breeding success and/or adult survival of nest‐building vertebrates. Although nest traits are expected to vary adaptively across elevational gradients, few studies address this relationship. We studied the variation in nest traits (composition and structure) across elevation for two taxa with two different functions in the Andean temperate forests of southern Chile: a bird (Aphrastura spinicauda, Furnariidae, 170 breeding nests) and a marsupial mammal (Dromiciops gliroides, Microbiotheriidae, 91 winter torpor nests). For A. spinicauda, we further assessed how nest traits influenced clutch size and hatching success. Both species used fewer types of nest materials (items) with increasing elevation, and a greater proportion of leaves were used in highland compared to lowland forests. Aphrastura spinicauda used feathers and hair, and D. gliroides used bryophytes more frequently in lowland forests. The mass and volume of nests decreased with increasing elevation for A. spinicauda and increased for D. gliroides. Nest traits had subsequent fitness consequences for A. spinicauda, such that: (i) greater cup volume and depth were associated with larger clutch sizes, (ii) more items used during nest building were linked to improved clutch size at high elevation only, and (iii) nest wall thickness was negatively associated with hatching success. Thus, in temperate mountain ecosystems, elevation may be an important factor influencing nest‐building behaviour for cavity‐using vertebrates. However, the direction of the elevational effects may vary among taxa and nest functions in these ecosystems.     

Waterfowl Nesting in Fall-Seeded and Spring-Seeded Cropland in Saskatchewan

Abstract: Waterfowl nesting in annual croplands has remained a little-known aspect of waterfowl nesting ecology because of the inability of many studies to systematically search this habitat through the nesting season. Where searches have been conducted, they are generally restricted to the period prior to seeding, and many nests found are destroyed by the seeding operation. Consequently, fall-seeded crops have been promoted as an alternative cropping practice that could increase nest survival of waterfowl nesting in croplands. During 1996–1999, we conducted 3–4 complete nest searches on 4,274 ha of cropland, including spring-seeded wheat and barley, winter wheat, and fall rye in southern Saskatchewan, Canada. Using suites of predictive models, we tested hypotheses regarding relative nest abundance and nest survival among crop types and tested the influence of several landscape-scale covariates on these metrics. Apparent nest densities were higher in fall-seeded crops (winter wheat: 0.39 nests/ha, fall rye: 0.25 nests/ha) than in spring-seeded crops (0.03 nests/ha), and nest density in spring-seeded croplands increased with percent cropland and percent wetland habitat in the surrounding landscape. Nest survival was higher in winter wheat (38%) than in either fall rye (18%) or spring-seeded crops (12%), and nest survival in spring-seeded crops increased with relative nest initiation date. Nest survival was unaffected by surrounding landscape characteristics but tended to be higher in years of average wetness. Based on our findings, winter wheat and fall rye have the potential to provide productive nesting habitat for ≥7 species of upland nesting ducks and fall-seeded crops are a conservation tool well suited to highly cropped landscapes.     

Merriam's Turkey Nest Survival and Factors Affecting Nest Predation by Mammals

Abstract: Nest success is an important parameter affecting population fluctuations of wild turkeys (Meleagris gallopavo). Factors influencing mammalian predation on turkey nests are complicated and not well understood. Therefore, we assessed nest hazard risk by testing competing hypotheses of Merriam's turkey (M. g. merriami) nest survival in a ponderosa pine (Pinus ponderosa) ecosystem during 2001–2003. We collected nesting information on 83 female Merriam's turkeys; annual nest success averaged 50% for adult females (range = 45–59%) and 83% for yearling females (range = 75–100%). Proportional hazard modeling indicated that precipitation increased the hazard of nest mortality. However, estimated hazard of nest predation was lowered when incubating females had greater shrub cover and visual obstruction around nests. Coyotes (Canis latrans) were the primary predator on turkey nests. We hypothesize that precipitation is the best predictor of nest survival for first nests because coyotes use olfaction effectively to find nesting females during wet periods. Temporally, as the nesting season progressed, precipitation declined and vegetation cover increased and coyotes may have more difficulty detecting nests under these conditions later in the nesting period. The interaction of concealment cover with precipitation indicated that nest hazard risk from daily precipitation was reduced with greater shrub cover. Management activities that promote greater shrub cover may partially offset the negative effects of greater precipitation events.     

Exotic shrubs as ephemeral ecological traps for nesting birds

Certain exotic plants may increase risk of nest predation, and, in this way, may act as ecological traps. We hypothesized that the greater vulnerability to predation was a consequence of either (1) reduced nest height due to architectural differences among plant species or (2) seasonal changes in the distribution of nests among forest strata. To test this, we examined temporal variation in nest survival of 888 nests of Northern Cardinal (Cardinalis cardinalis) in native substrates and two exotic shrubs (Lonicera maackii and Rosa multiflora) in Ohio, USA, 2001–2006. We evaluated evidence for an ecological trap by monitoring the annual reproductive productivity of 245 breeding pairs of cardinals. Only nests in Rosa experienced relatively constant survival rates across the season, whereas probability of survival increased over the season for nests in other substrates. Interestingly, the relative vulnerability of nests in different substrates varied across the season. Most strikingly, nests in Lonicera in early spring showed the lowest survival rates but exceeded survival rates of nests in native substrates late in the season. Nest height failed to explain seasonal changes in nest survival, as only nests in native plants significantly increased in height as the season progressed. Rather, predation risk seemed to be a function of the proportion of nests within each substrate, as illustrated by the decreased predation in Lonicera as the relative proportion of nests in native substrates increased. The patterns of temporal variation in predation risk that we detected show that impacts of Lonicera are not a function of plant architecture alone and may be related to leaf phenology, changes in nest density, nest site location, and/or nest synchrony. Examination of the reproductive productivity of cardinals showed that pairs that made their first nest attempt in Lonicera fledged 20% fewer cardinal young than birds that began the season using other substrates. Thus, we suggest that exotic plants may represent an ephemeral ecological trap for certain nesting birds, where negative effects persist only during certain periods.     

Nest predation reduces benefits to early clutch initiation in northern cardinals Cardinalis cardinalis

Life history theory and empirical studies suggest that early breeding confers higher reproductive success, but the extent to which this advantage can be generalized to human‐dominated systems and across species is less well understood. We studied the fitness consequences of clutch initiation for 181 female northern cardinals Cardinalis cardinalis and 1228 nests in forests within urban and rural landscapes of Ohio, USA between 2004–2007. Cardinals that bred earlier made significantly more nesting attempts, but cumulative number of young fledged was similar to that of later‐breeding individuals. The expected number of fledglings produced per successful nest was unrelated to date and remained ~1.8 fledglings across the season, despite the fact that nest survival rates improved dramatically as the season progressed. Because the probability of resighting breeding individuals in subsequent years was unrelated to first clutch initiation date, we have no evidence that clutch initiation affected adult survival. The absence of a clear benefit to early breeding appears to be a consequence of high rates of nest predation early in the breeding season.     

Nesting Behavior of <Emphasis Type="Italic">Gorilla gorilla diehli</Emphasis> at Kagwene Mountain,Cameroon: Implications for Assessing Group Size and Density

We recorded nesting data at 569 fresh night nest sites, comprising 7032 individual nests, of Cross River gorillas inhabiting the Kagwene Mountain in western Cameroon. The mean night nest group size was 12.4. Overall, 55% of night nests were constructed on the ground and 45% in trees. Significantly more arboreal nests were constructed in the wet season (69%), vs. the dry season (19%). Day nest construction was common at Kagwene (n = 260 nest sites, mean nest group size = 5.98) and we encountered significantly more day nest sites in the wet season. Nest site reuse was also common (35%), though not related to season. Our results of nesting habits concur with those from other western gorilla studies, in which rainfall influences arboreal nesting. However, we encountered wet season arboreal nesting, day nest construction, and overall nest site reuse more frequently than reported for other sites. Our observations have considerable implications when estimating group size and density using traditional nest count data. The gorillas at Kagwene inhabit the highest altitudinal range of all Cross River gorilla subpopulations and rainfall is also high; therefore other subpopulations may demonstrate different nesting characteristics. However, one should consider our findings when attempting to estimate Cross River gorilla density at other localities through nest site data.