Spatial Patterns of Breeding Success of Grizzly Bears Derived from Hierarchical Multistate Models

Conservation programs often manage populations indirectly through the landscapes in which they live. Empirically, linking reproductive success with landscape structure and anthropogenic change is a first step in understanding and managing the spatial mechanisms that affect reproduction, but this link is not sufficiently informed by data. Hierarchical multistate occupancy models can forge these links by estimating spatial patterns of reproductive success across landscapes. To illustrate, we surveyed the occurrence of grizzly bears (Ursus arctos) in the Canadian Rocky Mountains Alberta, Canada. We deployed camera traps for 6 weeks at 54 surveys sites in different types of land cover. We used hierarchical multistate occupancy models to estimate probability of detection, grizzly bear occupancy, and probability of reproductive success at each site. Grizzly bear occupancy varied among cover types and was greater in herbaceous alpine ecotones than in low‐elevation wetlands or mid‐elevation conifer forests. The conditional probability of reproductive success given grizzly bear occupancy was 30% (SE = 0.14). Grizzly bears with cubs had a higher probability of detection than grizzly bears without cubs, but sites were correctly classified as being occupied by breeding females 49% of the time based on raw data and thus would have been underestimated by half. Repeated surveys and multistate modeling reduced the probability of misclassifying sites occupied by breeders as unoccupied to <2%. The probability of breeding grizzly bear occupancy varied across the landscape. Those patches with highest probabilities of breeding occupancy—herbaceous alpine ecotones—were small and highly dispersed and are projected to shrink as treelines advance due to climate warming. Understanding spatial correlates in breeding distribution is a key requirement for species conservation in the face of climate change and can help identify priorities for landscape management and protection. Patrones Espaciales del Éxito Reproductivo de Osos Pardos, Derivados de Modelos Jerárquicos Multi‐Estado     

Efficacy of created and restored nesting sites for the conservation of colonial Laridae in the South of France

By rapidly modifying key habitat components, habitat restoration is at risk of producing attractive cues for animals without providing habitats of sufficient quality. As such, individual fitness components, such as reproduction, could be reduced and restored habitats could become ecological traps. This risk notably appears by using artificial constructions in restoration projects, yet few studies have evaluated their efficacy in a robust way. We investigated this by analyzing 154 islets that were created or restored to improve the conservation status of 7 colonial Laridae species in the South of France. From 2007 to 2016, we compared occupancy dynamics and breeding parameters of these species between the restored sites and 846 unmanaged nesting sites. We also explored species’ preference for different nesting site characteristics and their respective effect on breeding parameters. Restored nesting sites were 2–9 times as attractive as unmanaged sites for all species except the Black-headed Gull (Chroicocephalus ridibundus). Colonization probability was up to 100 times higher in sites already used by other species the previous year and increased with distance to the shore until >0.2 when distance was over 250 m. Abandonment probability was 29–70% lower when breeding was successful the previous year in all species except the Sandwich Tern (Thalasseus sandvicensis). Productivity and breeding success probability were 2 times higher on managed sites. Distance from the shore was an important attractive characteristic of artificial nesting sites in all species. Other nesting site characteristics had species-specific effects on colonization, abandonment, and breeding success. Our results indicate that managed nesting sites are successful conservation tools for colonial Laridae in the Mediterranean and do not act as ecological traps. Our study showed that testing the ecological trap hypothesis is a robust way to evaluate the success of restoration projects of breeding habitats.     

Possible Extinction Vortex for a Population of Iberian Lynx on the Verge of Extirpation

Abstract: Theory suggests that demographic and genetic traits deteriorate (i.e., fitness and genetic diversity decrease) when populations become small, and that such deterioration could precipitate positive feedback loops called extinction vortices. We examined whether demographic attributes and genetic traits have changed over time in one of the 2 remaining small populations of the highly endangered Iberian lynx (Lynx pardinus) in Doñana, Spain. From 1983 to 2008, we recorded nontraumatic mortality rates, litter size, offspring survival, age at territory acquisition, and sex ratio. We combined these demographic attributes with measures of inbreeding and genetic diversity at neutral loci (microsatellites) and genes subjected to selection (major histocompatibility complex). Data on demographic traits were obtained through capture and radio tracking, checking dens during breeding, track surveys, and camera trapping. For genetic analyses, we obtained blood or tissue samples from captured or necropsied individuals or from museum specimens. Over time a female‐biased sex ratio developed, age of territory acquisition decreased, mean litter size decreased, and rates of nontraumatic mortality increased, but there were no significant changes in overall mortality rates, standardized individual heterozygosity declined steadily, and allelic diversity of exon 2 of class II major histocompatibility complex DRB genes remained constant (2 allelic variants present in all individuals analyzed). Changes in sex ratio and age of territory acquisition may have resulted from demographic stochasticity, whereas changes in litter size and nontraumatic mortality may be related to observed increases in inbreeding. Concomitant deterioration of both demographic attributes and genetic traits is consistent with an extinction vortex. The co‐occurrence, with or without interaction, of demographic and genetic deterioration may explain the lack of success of conservation efforts with the Doñana population of Iberian lynx.     

Effects of age and sex ratios on offspring recruitment rates in translocated black rhinoceros

Success of animal translocations depends on improving postrelease demographic rates toward establishment and subsequent growth of released populations. Short‐term metrics for evaluating translocation success and its drivers, like postrelease survival and fecundity, are unlikely to represent longer‐term outcomes. We used information theory to investigate 25 years of data on black rhinoceros (Diceros bicornis) translocations. We used the offspring recruitment rate (ORR) of translocated females—a metric integrating survival, fecundity, and offspring recruitment at sexual maturity—to detect determinants of success. Our unambiguously best model (AICω = 0.986) predicted that ORR increases with female age at release as a function of lower postrelease adult rhinoceros sex ratio (males:females). Delay of first postrelease reproduction and failure of some females to recruit any calves to sexual maturity most influenced the pattern of ORRs, and the leading causes of recruitment failure were postrelease female death (23% of all females) and failure to calve (24% of surviving females). We recommend translocating older females (≥6 years old) because they do not exhibit the reproductive delay and low ORRs of juveniles (<4 years old) or the higher rates of recruitment failure of juveniles and young adults (4–5.9 years old). Where translocation of juveniles is necessary, they should be released into female‐biased populations, where they have higher ORRs. Our study offers the unique advantage of a long‐term analysis across a large number of replicate populations—a science‐by‐management experiment as a proxy for a manipulative experiment, and a rare opportunity, particularly for a large, critically endangered taxon such as the black rhinoceros. Our findings differ from previous recommendations, reinforce the importance of long‐term data sets and comprehensive metrics of translocation success, and suggest attention be shifted from ecological to social constraints on population growth and species recovery, particularly when translocating species with polygynous breeding systems.     

Living at the Edge: Local versus Positional Factors in the Long‐Term Population Dynamics of an Endangered Orchid

Abstract: Populations at the margin of geographic ranges of distribution have been considered more vulnerable than central ones, but recent reviews have caste doubt on this generalization. We examined the reproductive and demographic performance of a rare Euroasiatic orchid (Cypripedium calceolus) at its southwesterly range limit and compared our findings with those of previous studies of nine central populations at the center of the orchid's range. We sought to test the central‐marginal model and to evaluate factors involved in long‐term performance of forest Eurosiberian species with peripheral populations in southern European mountains. We characterized (structure, temporal fluctuations, herbivory, reproductive success, and recruitment at different habitats) four Pyrenean populations of C. calceolus of different sizes (5–3500 ramets) and monitored three of them for up to 13 years. Two quantitative stochastic models (count data and matrix models) were used to assess population trends and viability and the effect of herbivory. Contrary to expectations, and despite the negative effect of sporadic events of herbivory, the peripheral populations we studied (except the smallest one) performed similarly or better than populations occurring in central part of the species’ range in terms of reproductive success and population growth rates. Landscape changes over the last 50 years suggest that natural reforestation could be involved in the success of this plant at its southern limit. Forest expansion in the mountain regions of southern Europe may provide new opportunities for plants with geographic distributions centered mainly at higher latitudes and give some hope for their recovery in future scenarios dominated by biodiversity loss.     

Impacts of tropical forest disturbance on species vital rates

Tropical forests are experiencing enormous threats from deforestation and habitat degradation. Much knowledge of the impacts of these land-use changes on tropical species comes from studies examining patterns of richness and abundance. Demographic vital rates (survival, reproduction, and movement) can also be affected by land-use change in a way that increases species vulnerability to extirpation, but in many cases these impacts may not be manifested in short-term changes in abundance or species richness. We conducted a literature review to assess current knowledge and research effort concerning how land-use change affects species vital rates in tropical forest vertebrates. We found a general paucity of empirical research on demography across taxa and regions, with some biases toward mammals and birds and land-use transitions, including fragmentation and agriculture. There is also considerable between-species variation in demographic responses to land-use change, which could reflect trait-based differences in species sensitivity, complex context dependencies (e.g., between-region variation), or inconsistency in methods used in studies. Efforts to improve understanding of anthropogenic impacts on species demography are underway, but there is a need for increased research effort to fill knowledge gaps in understudied tropical regions and taxa. The lack of information on demographic impacts of anthropogenic disturbance makes it difficult to draw definite conclusions about the magnitude of threats to tropical ecosystems under anthropogenic pressures. Thus, determining conservation priorities and improving conservation effectiveness remains a challenge.     

Sex‐Biased Inbreeding Effects on Reproductive Success and Home Range Size of the Critically Endangered Black Rhinoceros

A central premise of conservation biology is that small populations suffer reduced viability through loss of genetic diversity and inbreeding. However, there is little evidence that variation in inbreeding impacts individual reproductive success within remnant populations of threatened taxa, largely due to problems associated with obtaining comprehensive pedigree information to estimate inbreeding. In the critically endangered black rhinoceros, a species that experienced severe demographic reductions, we used model selection to identify factors associated with variation in reproductive success (number of offspring). Factors examined as predictors of reproductive success were age, home range size, number of nearby mates, reserve location, and multilocus heterozygosity (a proxy for inbreeding). Multilocus heterozygosity predicted male reproductive success (p< 0.001, explained deviance >58%) and correlated with male home range size (p < 0.01, r² > 44%). Such effects were not apparent in females, where reproductive success was determined by age (p < 0.01, explained deviance 34%) as females raise calves alone and choose between, rather than compete for, mates. This first report of a 3‐way association between an individual male's heterozygosity, reproductive output, and territory size in a large vertebrate is consistent with an asymmetry in the level of intrasexual competition and highlights the relevance of sex‐biased inbreeding for the management of many conservation‐priority species. Our results contrast with the idea that wild populations of threatened taxa may possess some inherent difference from most nonthreatened populations that necessitates the use of detailed pedigrees to study inbreeding effects. Despite substantial variance in male reproductive success, the increased fitness of more heterozygous males limits the loss of heterozygosity. Understanding how individual differences in genetic diversity mediate the outcome of intrasexual competition will be essential for effective management, particularly in enclosed populations, where individuals have restricted choice about home range location and where the reproductive impact of translocated animals will depend upon the background distribution in individual heterozygosity. Efectos de la Endogamia Sesgada por el Sexo sobre el Éxito Reproductivo y el Rango del Tamaño de Hábitat del Rinoceronte Negro, Especie en Peligro Crítico     

Use of fecundity measured directly throughout the breeding season to test a source-sink demographic model

Populations of landbirds (bird species that occupy terrestrial habitats for most of their life cycle) are declining throughout North America (north of Mexico) and Europe, yet little is known about how demography is driving this trend. A recent model of 5 geographically separated populations of Cerulean Warblers (Dendroica cerulea) that was based on within-season sampling of nest survival and fledgling success shows that all populations are sinks (annual reproduction is consistently less than annual adult mortality). I tested this indirect model by directly measuring fecundity (number of female fledglings/female) during the breeding season for 2 years in a Cerulean Warbler population occupying a mature forest in southwestern Michigan (U.S.A.) I determined territories of male birds on the basis of male plumage characters and phases of the nesting cycle (2007) and on uniquely color-banded males (2008). I transferred locations of identified males to topographic maps. I counted all fledglings in territories from May to July each year. The model I tested may apply only to single-brooded species; therefore, I searched the literature to estimate the percentage of single-brooded species in North America. The breeding season of Cerulean Warblers was short- nearly all nests were initiated from mid-May to late June. Nest predation and brood parasitism were primary and rare causes of nest failure, respectively. Significantly fewer Cerulean Warblers fledged from parasitized than from nonparasitized nests. Fledgling survival required to maintain the population size was well above previously published values for Neotropical migrants. Single-brooded species comprise 62% of North American breeding bird species for which the number of broods per year is known; I believe my results may apply to these species. The consistency between identification of populations as sources or sinks on the basis of either model estimates or direct measurements suggests that a demographic model relying on within-season sampling of fecundity is adequate to determine population status of single-brooded avian populations. In addition, on the basis of results of previous studies, annual adult survival rate of the Cerulean Warbler is typical of parulid warblers that are not declining. Thus, low fecundity, here determined with different quantitative methods, can drive status of landbird species with high-observed survival.     

Divergent effects of climate change on the egg-laying opportunity of species in cold and warm regions

Climate warming can substantially impact embryonic development and juvenile growth in oviparous species. Estimating the overall impacts of climate warming on oviparous reproduction is difficult because egg-laying events happen throughout the reproductive season. Successful egg laying requires the completion of embryonic development as well as hatching timing conducive to offspring survival and energy accumulation. We propose a new metric—egg-laying opportunity (EO)—to estimate the annual hours during which a clutch of freshly laid eggs yields surviving offspring that store sufficient energy for overwintering. We estimated the EO within the distribution of a model species, Sceloporus undulatus, under recent climate condition and a climate-warming scenario by combining microclimate data, developmental functions, and biophysical models. We predicted that EO will decline as the climate warms at 74.8% of 11,407 sites. Decreasing hatching success and offspring energy accounted for more lost EO hours (72.6% and 72.9%) than the occurrence of offspring heat stress (59.9%). Nesting deeper (at a depth of 12 cm) may be a more effective behavioral adjustment for retaining EO than using shadier (50% shade) nests because the former fully mitigated the decline of EO under the considered warming scenario at more sites (66.1%) than the latter (28.3%). We advocate for the use of EO in predicting the impacts of climate warming on oviparous animals because it encapsulates the integrative impacts of climate warming on all stages of reproductive life history.     

Land trust evaluation of progress toward conservation goals

In the United States local land trusts preserve and conserve land to protect water quality and supply, farmlands, habitat for native plants and animals, areas of cultural or historical significance, and scenic views. We surveyed the 24 active, local land trusts in North Carolina (U.S.A.) to determine how they report progress toward attaining such conservation goals. Twenty-two land trusts responded to our survey. Of these, eight reported developing specific conservation goals for all of the properties they protect, five identified conservation targets on all properties, and two reported monitoring biological indicators on all of their protected properties. On the basis of these results, we believe most of the land trusts surveyed could not determine whether they were meeting conservation goals because most did not identify explicit, measurable goals and monitor progress toward them. Instead they reported success in terms of the amount of land protected and money raised. We think this is a lost opportunity for land trusts to build sound approaches to environmental management, engage the public, entice new donors, and attain funding for additional conservation activities. We propose conservation professionals help local land trusts adopt the Open Standards for the Practice of Conservation, a framework developed by a consortium of international conservation organizations, to develop conservation goals and measure whether the goals are achieved.