Mixed-stock analysis reveals the migrations of juvenile hawksbill turtles (Eretmochelys imbricata) in the Caribbean Sea

Hawksbill turtles (Eretmochelys imbricata) migrate between nesting beaches and feeding habitats that are often associated with tropical reefs, but it is uncertain which nesting colonies supply which feeding habitats. To address this gap in hawksbill biology, we compile previously published and new mitochondrial DNA (mtDNA) haplotype data for 10 nesting colonies (N = 347) in the western Atlantic and compare these profiles to four feeding populations and four previously published feeding samples (N = 626). Nesting colonies differ significantly in mtDNA haplotype frequencies (Phi(ST) = 0.588, P < 0.001), corroborating earlier conclusions of nesting site fidelity and setting the stage for mixed-stock analysis. Feeding aggregations show lower but significant structure (Phi(ST) = 0.089, P < 0.001), indicating that foraging populations are not homogenous across the Caribbean Sea. Bayesian mixed-stock estimates of the origins of juveniles in foraging areas show a highly significant, but shallow, correlation with nesting population size (r = 0.378, P = 0.004), supporting the premise that larger rookeries contribute more juveniles to feeding areas. A significant correlation between the estimated contribution and geographical distance from nesting areas (r = -0.394, P = 0.003) demonstrates the influence of proximity on recruitment to feeding areas. The influence of oceanic currents is illustrated by pelagic stage juveniles stranded in Texas, which are assigned primarily (93%) to the upstream rookery in Yucatan. One juvenile had a haplotype previously identified only in the eastern Atlantic, invoking rare trans-oceanic migrations. The mixed-stock analysis demonstrates that harvests in feeding habitats will impact nesting colonies throughout the region, with the greatest detriment to nearby nesting populations.     

Dispersal, recruitment and migratory behaviour in a hawksbill sea turtle aggregation

We investigated the dispersal, recruitment and migratory behaviour of the hawksbill sea turtle ( Eretmochelys imbricata ), among different life-history stages and demographic segments of the large hawksbill turtle aggregation at Mona Island, Puerto Rico. There were significant differences in both mitochondrial DNA (mtDNA) haplotype diversity and haplotype frequencies among the adult males, females and juveniles examined, but little evidence for temporal heterogeneity within these same groups sampled across years. Consistent with previous studies and the hypothesis of strong natal homing, there were striking mtDNA haplotype differences between nesting females on Mona Island and nesting females in other major Caribbean rookeries. Breeding males also showed strong, albeit weaker, genetic evidence of natal homing. Overall, Bayesian mixed-stock analysis suggests that Mona Island was the natal rookery for 79% (65–94%) of males in the aggregation. In contrast, the Mona Island rookery accounted for only a small subset of the new juvenile recruits to the foraging grounds or in the population of older juvenile hawksbills turtles on Mona. Instead, both new recruits and the older juvenile hawksbill turtles on Mona more likely recruited from other Caribbean rookeries, suggesting that a mechanism besides natal homing must be influencing recruitment to feeding habitats. The difference in the apparent degree of natal homing behaviour among the different life-history stages of hawksbill turtles at Mona Island underscores the complexity of the species' life-history dynamics and highlights the need for both local and regional conservation efforts.     

Turtle groups or turtle soup: dispersal patterns of hawksbill turtles in the Caribbean

Despite intense interest in conservation of marine turtles, spatial ecology during the oceanic juvenile phase remains relatively unknown. Here, we used mixed stock analysis and examination of oceanic drift to elucidate movements of hawksbill turtles (Eretmochelys imbricata) and address management implications within the Caribbean. Among samples collected from 92 neritic juvenile hawksbills in the Cayman Islands we detected 11 mtDNA control region haplotypes. To estimate contributions to the aggregation, we performed ‘many‐to‐many’ mixed stock analysis, incorporating published hawksbill genetic and population data. The Cayman Islands aggregation represents a diverse mixed stock: potentially contributing source rookeries spanned the Caribbean basin, delineating a scale of recruitment of 200–2500 km. As hawksbills undergo an extended phase of oceanic dispersal, ocean currents may drive patterns of genetic diversity observed on foraging aggregations. Therefore, using high‐resolution Aviso ocean current data, we modelled movement of particles representing passively drifting oceanic juvenile hawksbills. Putative distribution patterns varied markedly by origin: particles from many rookeries were broadly distributed across the region, while others would appear to become entrained in local gyres. Overall, we detected a significant correlation between genetic profiles of foraging aggregations and patterns of particle distribution produced by a hatchling drift model (Mantel test, r = 0.77, P < 0.001; linear regression, r = 0.83, P < 0.001). Our results indicate that although there is a high degree of mixing across the Caribbean (a ‘turtle soup’), current patterns play a substantial role in determining genetic structure of foraging aggregations (forming turtle groups). Thus, for marine turtles and other widely distributed marine species, integration of genetic and oceanographic data may enhance understanding of population connectivity and management requirements.     

Contextualising the Last Survivors: Population Structure of Marine Turtles in the Dominican Republic

Nesting by three species of marine turtles persists in the Dominican Republic, despite historic threats and long-term population decline. We conducted a genetic survey of marine turtles in the Dominican Republic in order to link them with other rookeries around the Caribbean. We sequenced a 740_bp fragment of the control region of the mitochondrial DNA of 92 samples from three marine turtle species [hawksbill (n = 48), green (n = 2) and leatherback (n = 42)], and incorporated published data from other nesting populations and foraging grounds. The leatherback turtle (Dermochelys coriacea) in the Dominican Republic appeared to be isolated from Awala-Yalimapo, Cayenne, Trinidad and St. Croix but connected with other Caribbean populations. Two distinct nesting populations of hawksbill turtles (Eremochelys imbricata) were detected in the Dominican Republic and exhibited interesting patterns of connectivity with other nesting sites and juvenile and adult male foraging aggregations. The green sea turtle (Chelonia mydas) has almost been extirpated from the Dominican Republic and limited inference could be made from our samples. Finally, results were compared with Lagrangian drifting buoys and published Lagrangian virtual particles that travelled through the Dominican Republic and Caribbean waters. Conservation implications of sink-source effects or genetic isolation derived from these complex inter-connections are discussed for each species and population.     

Príncipe island hawksbills: Genetic isolation of an eastern Atlantic stock

The hawksbill turtle is a critically endangered species that has been extensively exploited for centuries. Príncipe Island off western Africa harbours one of the species' major nesting populations in the eastern Atlantic, as well as hosting year-round foraging aggregations of juveniles, subadults and adult males. To gain insight into the population's genetic structure and relationships with regional stocks, we analysed mitochondrial DNA (mtDNA) sequences of nesting females (N = 9), foraging adult females (N = 11), adult males (N = 32), subadults (N = 15) and juveniles (N = 80). The nesting population was found to be fixed for a single haplotype (EATL), which had been previously reported in both western and eastern Atlantic hawksbill foraging sites but had no known rookery source prior to this study. Thus it is now possible to confirm the westward transoceanic movement by hawksbills originating from Príncipe Island. Our analyses demonstrated that the Príncipe Island nesting colony is genetically distinct from breeding populations in the western Atlantic and is phylogenetically linked with Indo-Pacific hawksbill clades, suggesting that Príncipe Island was most likely colonised by migrants from the Indian Ocean via the Cape of Good Hope in southern Africa. Mixed stock analyses revealed that the eastern Atlantic appears to be the primary foraging area for Príncipe hawksbills (75%) while most of the foraging juveniles in Príncipe waters originate from the Príncipe rookery (84%). Furthermore, the presence of Caribbean haplotypes at low frequencies (< 5%) suggests that eastward transatlantic movements by juveniles to distant foraging and developmental habitats also take place. Depleted hawksbill populations in the eastern Atlantic combined with the low genetic variability and high genetic distinctiveness found in the Príncipe nesting and foraging aggregations with respect to the western Atlantic, underscore the high degree of isolation and vulnerability of this eastern Atlantic stock. These characteristics are highly relevant for the development of effective conservation programmes and highlight the urgent need to consolidate international cooperation across regional boundaries.     

Phenotypically linked dichotomy in sea turtle foraging requires multiple conservation approaches

Marine turtles undergo dramatic ontogenic changes in body size and behavior, with the loggerhead sea turtle, Caretta caretta, typically switching from an initial oceanic juvenile stage to one in the neritic, where maturation is reached and breeding migrations are subsequently undertaken every 2-3 years. Using satellite tracking, we investigated the migratory movements of adult females from one of the world's largest nesting aggregations at Cape Verde, West Africa. In direct contrast with the accepted life-history model for this species, results reveal two distinct adult foraging strategies that appear to be linked to body size. The larger turtles (n = 3) foraged in coastal waters, whereas smaller individuals (n = 7) foraged oceanically. The conservation implications of these findings are profound, with the population compartmentalized into habitats that may be differentially impacted by fishery threats in what is a global fishing hotspot. Although the protection of discrete areas containing coastal individuals may be attainable, the more numerous pelagic individuals are widely dispersed with individuals roaming over more than half a million square kilometers. Therefore, mitigation of fisheries by-catch for sea turtles in the east Atlantic will likely require complex and regionally tailored actions to account for this dichotomous behavior.     

Behavioral and energetic costs of group membership in a coral reef fish

Animals in social aggregations often spend more time foraging than solitary conspecifics. This may be a product of the relative safety afforded by aggregations: group members can devote more time to foraging and less time to antipredator behaviors than solitary animals (the “risk reduction” effect). All else being equal, risk reduction should result in higher food intake for grouped animals. However, intragroup competition may force group members to spend more time foraging in order to obtain the same food ration as solitary individuals (the “resource competition” effect). We compared these opposing explanations of foraging time allocation in a coral reef fish, bluehead wrasse (Thalassoma bifasciatum). Aggregations of juvenile bluehead wrasse experience safety-in-numbers, and preliminary observations suggested that juveniles in aggregations spent more time foraging for copepods in the water column than solitary juveniles. However, the risk reduction and resource competition hypotheses are indistinguishable on the basis of behavioral observations alone. Therefore, we collected behavioral, dietary, and growth data (using otolith growth rings) for bluehead wrasse at multiple reefs around a Caribbean island. Despite spending more time foraging in the water column, grouped fish did not capture more prey items and had slower growth rates than solitary fish. Thus, the increased foraging time of grouped fish appears to reflect resource competition, not risk reduction. This competition may limit the size and frequency of aggregations among juvenile bluehead wrasse, which have been shown to experience reduced mortality rates in larger groups. Bluehead wrasse recruits also spent less time foraging but grew faster at sites where planktonic copepod prey were more abundant. This suggests the possibility that large-scale spatiotemporal variability in the abundance of planktonic copepods over coral reefs may produce corresponding variability in the dynamics of reef fish populations. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Down but not out: marine turtles of the British Virgin Islands

We present the result of a multi-annual assessment of the spatio-temporal patterns of marine turtle nesting, and foraging in the Eastern Caribbean archipelago state of the British Virgin Islands. Despite exploitation over several centuries, three species (leatherback Dermochelys coriacea , green Chelonia mydas and hawksbill Eretmochelys imbricata turtles) are still nesting and green and hawksbill turtles are found foraging. Leatherback turtles are showing signs of a recovery co-incident with the implementation of an effective moratorium on adult take. When compared with historical data we demonstrate an apparent reduction in nesting levels in green and hawksbill turtles and a nesting range contraction in green turtles. Despite current conservation steps, it will be a decade or more before evidence of recovery can be expected in the two hard-shelled species. Coupled with wider biological knowledge, our findings offer insights into the relative resilience of the different species to exploitation pressure. Additionally, the intra-annual temporal and spatial spread of nesting demonstrated underlines the difficulties of monitoring such a multi-species assemblage in such a diffuse archipelago.     

Natal homing in juvenile loggerhead turtles (Caretta caretta)

Juvenile loggerhead turtles (Caretta caretta) from West Atlantic nesting beaches occupy oceanic (pelagic) habitats in the eastern Atlantic and Mediterranean, whereas larger juvenile turtles occupy shallow (neritic) habitats along the continental coastline of North America. Hence the switch from oceanic to neritic stage can involve a trans-oceanic migration. Several researchers have suggested that at the end of the oceanic phase, juveniles are homing to feeding habitats in the vicinity of their natal rookery. To test the hypothesis of juvenile homing behaviour, we surveyed 10 juvenile feeding zones across the eastern USA with mitochondrial DNA control region sequences (N = 1437) and compared these samples to potential source (nesting) populations in the Atlantic Ocean and Mediterranean Sea (N = 465). The results indicated a shallow, but significant, population structure of neritic juveniles (PhiST = 0.0088, P = 0.016), and haplotype frequency differences were significantly correlated between coastal feeding populations and adjacent nesting populations (Mantel test R2 = 0.52, P = 0.001). Mixed stock analyses (using a Bayesian algorithm) indicated that juveniles occurred at elevated frequency in the vicinity of their natal rookery. Hence, all lines of evidence supported the hypothesis of juvenile homing in loggerhead turtles. While not as precise as the homing of breeding adults, this behaviour nonetheless places juvenile turtles in the vicinity of their natal nesting colonies. Some of the coastal hazards that affect declining nesting populations may also affect the next generation of turtles feeding in nearby habitats.     

Artisanal Green Turtle,Chelonia mydas,Fishery of Caribbean Nicaragua: I. Catch Rates and Trends, 1991–2011

This is the first assessment of catch rates for the legal, artisanal green turtle, Chelonia mydas, fishery in Caribbean Nicaragua. Data were collected by community members, monitoring up to 14 landing sites from 1991 to 2011. We examined take levels, and temporal and spatial variability in catch rates for the overall fishery, by region, and community using General Additive Mixed Models (GAMMs). More than 171,556 green turtles were killed during the period, with a mean estimated minimum 8,169±2,182 annually. There was a statistically significant decline in catch rates overall. Catch rates peaked in 1997 and 2002, followed by a downward trend, particularly from mid-2008 to the end of the study period. Similar downward trends were evident in both study regions. Community specific catch rate trends also indicated declines with decreases ranging from 21% to 90%. Decrease in catch rates in Nicaragua is cause for concern even though the principal source rookery at Tortuguero, Costa Rica, shows an increase in nesting activity. Explanations for the apparent discrepancy between the increasing trend at Tortuguero and decreasing catch rate trends in Nicaragua include: i) an increase in reproductive output, ii) insufficient time has passed to observe the impact of the fishery on the rookery due to a time lag, iii) changes in other segments of the population have not been detected since only nesting activity is monitored, iv) the expansive northern Nicaragua foraging ground may provide a refuge for a sufficient portion of the Tortuguero rookery, and/or v) a larger than expected contribution of non-Tortuguero rookeries occurring in Nicaragua turtle fishing areas. Our results highlight the need for close monitoring of rookeries and in-water aggregations in the Caribbean. Where consumptive use still occurs, nations sharing this resource should implement scientifically based limits on exploitation to ensure sustainability and mitigate impacts to regional population diversity.     

Nest sites,breeding cycle and possible cleptoparasites of Monoeca catarina Aguiar (Hymenoptera,Apidae, Tapinotaspidini) from Santa Catarina Island,southern Brazil

ABSTRACT

The recently described solitary bee Monoeca catarina Aguiar (Apidae, Tapinotaspidini) is only known from two nest aggregations on Santa Catarina Island, state of Santa Catarina, southern Brazil. Nests are excavated in sandy soils along trails near dunes and beaches. Up to 287 nests were counted in one aggregation. Bees were active for about seven weeks from late November to mid-January suggesting a univoltine life cycle. Males perform patrolling flights over the nest area and mate with females sitting on low vegetation. Males and females visited Stachytarpheta cayennensis (Verbenaceae) for nectar, and females were observed foraging on Tetrapterys sp. or Heteropterys sp. flowers (Malpighiaceae). Pollen loads consisted almost exclusively of Malpighiaceae pollen. Potential natural enemies are the cleptoparasitic bee Protosiris Roig-Alsina sp. (Apidae, Osirini) recorded within the aggregations, and the mutillid wasp Hoplocrates specularis (Gerstaecker) (Mutillidae, Sphaeropthalmini) seen entering the Monoeca nests and mating within the aggregations. Except for the soil characteristics, our observations on nesting and foraging behavior and associated potential enemies fit well into the facts reported from other Monoeca species.     

Colonially breeding seabirds: Predators or prey?

The evolution of aggregation in seabirds is usually attributed to predation pressure, although many authors have argued for the role of foraging constraints to be considered. Discriminating between factors that result in group living and characteristics arising from group living has been regarded as an insoluble problem; yet it is possible to test the predictions of the different models proposed to explain colonial nesting. The resulting explanation suggests that seabird aggregations have evolved in order to utilize a dispersed and unpredictable food supply. As such, colonial nesting leads to increased vulnerability to predation, rather than being a strategy adopted to combat it.     

Genetic composition,population structure and phylogeography of the loggerhead sea turtle: colonization hypothesis for the Brazilian rookeries

The loggerhead sea turtle, Caretta caretta, is the most common species of sea turtle nesting in Brazil and is listed as endangered by the IUCN. Our study characterizes the genetic structure of loggerheads in Brazil based on mitochondrial DNA control region variability and presents a hypothesis for the colonization of Brazilian rookeries. We analyzed 329 samples from Brazilian rookeries and an oceanic foraging ground, and we compared our results with previously published data for other loggerhead populations. Brazilian rookeries had four haplotypes, none of which have been reported for rookeries outside Brazil. Six haplotypes were found in the foraging aggregation. The presence of the CC-A4 haplotype at all sampled sites and the low nucleotide diversity suggest a common origin for all rookeries, with CC-A4 being the ancestral haplotype of the Brazilian populations. The occurrence of three haplotypes in the foraging aggregation that are known only from rookeries outside of Brazil is consistent with the transoceanic migratory behavior of loggerheads. Our results indicated that the colonization of Brazilian rookeries probably occurred from the southern USA stock. This recent colonization most likely followed a north to south route along the Brazilian coastline, influenced by the Brazilian warm current. Our results further suggest the existence of two genetic population units of loggerheads in Brazil and corroborate natal homing behavior in loggerheads.     

Age and hunting success in the brown pelican: influences of skill and patch choice on foraging efficiency

Summary Age-related differences in the foraging efficiency of piscivorous birds may be the results of differences in foraging skill, patch usage, or both. Brown pelicans were observed while foraging around a small Caribbean island. Areas where the birds fed were subdivided into small, homogeneous subunits (patches), and the bird's foraging success and patch use were noted and analyzed using multivariate techniques. Adult birds were found to be better at capturing prey under all conditions than were juveniles, but the differences were small in some patches. The density of prey and the birds' foraging success influenced the foraging efforts of adult and juvenile pelicans to similar degrees. Both age groups utilized local enhancement in their foraging, but such behavior augmented the foraging success only of juvenile birds. Both age groups often fed in patches where their foraging success was quite low. Such behaviour was much more costly for juvenile than for adult pelicans.     

Genetic structure of an Apis dorsata population: the significance of migration and colony aggregation

Eight microsatellite loci were used to investigate the genetic structure of the giant honeybee (Apis dorsata) population in northeast India. This species migrates seasonally between summer and winter nesting sites, and queens appear to return to their previously occupied site. Furthermore, there is a strong tendency for colonies of this species to aggregate at perennially utilized nesting sites that may be shared by more than 150 colonies. These behavioral features suggest that colonies within aggregations should be more related than random colonies, but that the long-distance migration could act to minimize genetic differentiation both between geographical areas and within aggregations. Our genetic study supports these conjectures arising from natural history. A. dorsata aggregations are comprised of colonies that share more alleles than expected by chance. Although queens heading neighboring colonies are not close relatives, fixation indices show significant genetic differentiation among aggregation sites. However, there appears to be sufficient gene flow among aggregations to prevent high degrees of relatedness developing between colonies within aggregations. The results also suggest that there is significant population structuring between geographical regions, although the level of structuring caused by aggregation exceeds the differentiation attributable to geographic region.     

Modeling the emergence of migratory corridors and foraging hot spots of the green sea turtle

Environmental factors shape the spatial distribution and dynamics of populations. Understanding how these factors interact with movement behavior is critical for efficient conservation, in particular for migratory species. Adult female green sea turtles, Chelonia mydas, migrate between foraging and nesting sites that are generally separated by thousands of kilometers. As an emblematic endangered species, green turtles have been intensively studied, with a focus on nesting, migration, and foraging. Nevertheless, few attempts integrated these behaviors and their trade‐offs by considering the spatial configurations of foraging and nesting grounds as well as environmental heterogeneity like oceanic currents and food distribution. We developed an individual‐based model to investigate the impact of local environmental conditions on emerging migratory corridors and reproductive output and to thereby identify conservation priority sites. The model integrates movement, nesting, and foraging behavior. Despite being largely conceptual, the model captured realistic movement patterns which confirm field studies. The spatial distribution of migratory corridors and foraging hot spots was mostly constrained by features of the regional landscape, such as nesting site locations, distribution of feeding patches, and oceanic currents. These constraints also explained the mixing patterns in regional forager communities. By implementing alternative decision strategies of the turtles, we found that foraging site fidelity and nesting investment, two characteristics of green turtles' biology, are favorable strategies under unpredictable environmental conditions affecting their habitats. Based on our results, we propose specific guidelines for the regional conservation of green turtles as well as future research suggestions advancing spatial ecology of sea turtles. Being implemented in an easy to learn open‐source software, our model can coevolve with the collection and analysis of new data on energy budget and movement into a generic tool for sea turtle research and conservation. Our modeling approach could also be useful for supporting the conservation of other migratory marine animals.     

Modeling spatiotemporal dynamics of krill aggregations: size,intensity, persistence,and coherence with seabirds

Understanding aggregation dynamics of forage species is important for evaluating biophysical scaling in marine ecosystems and heterogeneity of trophic interactions. In particular, zooplankton aggregations are fundamental units of many pelagic systems, but are difficult to observe continuously through space and time. Using an established modeling framework that encompasses a coupled regional oceanographic and individual‐based modeling system, we test the hypothesis that persistence (duration) of krill aggregations is dependent on their size, intensity, and location of formation within the coastal upwelling region of the California Current. In support of this hypothesis, we found that aggregation size is positively related to intensity, whereas persistence has a parabolic response to aggregation size and intensity, indicating the likelihood that large and highly persistent aggregations are rare. Persistence of krill aggregations also depends on formation location within coastal upwelling areas. We found that krill aggregations were more likely to form near a major seabird colony and that some coastal upwelling areas act as sources of aggregations for other areas. Observations of seabird aggregations were used to evaluate the potential structural realism of predicted krill aggregations. Seabird aggregations displayed marked coherence with predicted krill aggregations in space, providing important criteria on the scaling and availability of krill aggregations to breeding and migratory species. Predicting scales of krill aggregation dynamics will benefit ecosystem assessments, and numerical modeling of predator foraging and marine spatial management aimed at ensuring protection of ecologically important areas.     

Conservation implications of complex population structure: lessons from the loggerhead turtle (Caretta caretta)

Complex population structure can result from either sex-biased gene flow or population overlap during migrations. Loggerhead turtles (Caretta caretta) have both traits, providing an instructive case history for wildlife management. Based on surveys of maternally inherited mtDNA, pelagic post-hatchlings show no population structure across the northern Atlantic (phi(ST) < 0.001, P = 0.919), subadults in coastal habitat show low structure among locations (phi(ST) = 0.01, P < 0.005), and nesting colonies along the southeastern coast of the United States have strong structure (phi(ST) = 0.42, P < 0.001). Thus the level of population structure increases through progressive life history stages. In contrast, a survey of biparentally inherited microsatellite DNA shows no significant population structure: R(ST) < 0.001; F(ST) = 0.002 (P > 0.05) across the same nesting colonies. These results indicate that loggerhead females home faithfully to their natal nesting colony, but males provide an avenue of gene flow between regional nesting colonies, probably via opportunistic mating in migratory corridors. As a result, all breeding populations in the southeastern United States have similar levels of microsatellite diversity (H(E) = 0.70-0.89), whereas mtDNA haplotype diversity varies dramatically (h = 0.00-0.66). Under a conventional interpretation of the nuclear DNA data, the entire southeastern United States would be regarded as a single management unit, yet the mtDNA data indicate multiple isolated populations. This complex population structure mandates a different management strategy at each life stage. Perturbations to pelagic juveniles will have a diffuse impact on Atlantic nesting colonies, mortality of subadults will have a more focused impact on nearby breeding populations, and disturbances to adults will have pinpoint impact on corresponding breeding populations. These findings demonstrate that surveys of multiple life stages are desirable to resolve management units in migratory marine species.     

A New Mass Mortality of Juvenile Protoceratops and Size-Segregated Aggregation Behaviour in Juvenile Non-Avian Dinosaurs

Background

Monodominant bonebeds are a relatively common occurrence for non-avian dinosaurs, and have been used to infer associative, and potentially genuinely social, behavior. Previously known assemblages are characterized as either mixed size-classes (juvenile and adult-sized specimens together) or single size-classes of individuals (only juveniles or only adult-sized individuals within the assemblage). In the latter case, it is generally unknown if these kinds of size-segregated aggregations characterize only a particular size stage or represent aggregations that happened at all size stages. Ceratopsians (“horned dinosaurs”) are known from both types of assemblages.

Methods/Principal Findings

Here we describe a new specimen of the ceratopsian dinosaur Protoceratops andrewsi, Granger and Gregory 1923 from Mongolia representing an aggregation of four mid-sized juvenile animals. In conjunction with existing specimens of groups of P. andrewsi that includes size-clustered aggregations of young juveniles and adult-sized specimens, this new material provides evidence for some degree of size-clustered aggregation behaviour in Protoceratops throughout ontogeny. This continuity of size-segregated (and presumably age-clustered) aggregation is previously undocumented in non-avian dinosaurs.

Conclusions

The juvenile group fills a key gap in the available information on aggregations in younger ceratopsians. Although we support the general hypothesis that many non-avian dinosaurs were gregarious and even social animals, we caution that evidence for sociality has been overstated and advocate a more conservative interpretation of some data of ‘sociality’ in dinosaurs.     

Nest Suitability,Fine-Scale Population Structure and Male-Mediated Dispersal of a Solitary Ground Nesting Bee in an Urban Landscape

Bees are the primary pollinators of flowering plants in almost all ecosystems. Worldwide declines in bee populations have raised awareness about the importance of their ecological role in maintaining ecosystem functioning. The naturally strong philopatric behavior that some bee species show can be detrimental to population viability through increased probability of inbreeding. Furthermore, bee populations found in human-altered landscapes, such as urban areas, can experience lower levels of gene flow and effective population sizes, increasing potential for inbreeding depression in wild bee populations. In this study, we investigated the fine-scale population structure of the solitary bee Colletes inaequalis in an urbanized landscape. First, we developed a predictive spatial model to detect suitable nesting habitat for this ground nesting bee and to inform our field search for nests. We genotyped 18 microsatellites in 548 female individuals collected from nest aggregations throughout the study area. Genetic relatedness estimates revealed that genetic similarity among individuals was slightly greater within nest aggregations than among randomly chosen individuals. However, genetic structure among nest aggregations was low (Nei’s G_ST = 0.011). Reconstruction of parental genotypes revealed greater genetic relatedness among females than among males within nest aggregations, suggesting male-mediated dispersal as a potentially important mechanism of population connectivity and inbreeding avoidance. Size of nesting patch was positively correlated with effective population size, but not with other estimators of genetic diversity. We detected a positive trend between geographic distance and genetic differentiation between nest aggregations. Our landscape genetic models suggest that increased urbanization is likely associated with higher levels of inbreeding. Overall, these findings emphasize the importance of density and distribution of suitable nesting patches for enhancing bee population abundance and connectivity in human dominated habitats and highlights the critical contribution of landscape genetic studies for enhanced conservation and management of native pollinators.