Effects of Harvest of Nontimber Forest Products and Ecological Differences between Sites on the Demography of African Mahogany

Abstract: The demographic impacts of harvesting nontimber forest products (NTFP) have been increasingly studied because of reports of potentially unsustainable harvest. Nevertheless, our understanding of how plant demographic response to harvest is altered by variation in ecological conditions, which is critical for developing realistic sustainable‐use plans, is limited. We built matrix population models to test whether and how variation in ecological conditions affects population responses to harvest. In particular, we examined the effect of bark and foliage harvest on the demography of populations of African mahogany (Khaya senegalensis) in two contrasting ecological regions of Benin, West Africa. K. senegalensis bark and foliage harvest significantly reduced its stochastic population growth rates, but ecological differences between regions had a greater effect on population growth rates than did harvest. The effect of harvest on population growth rates (Δλ) was slightly stronger in the moist than in the drier region. Life‐table response experiments revealed that the mechanism by which harvesting reduced λ differed between ecological regions. Lowered stasis (persistence) of larger life stages lead to a reduction in λ in the drier region, whereas lowered growth of all life stages lowered λ in moist region. Potential strategies to increase population growth rates should include decreasing the proportion of individuals harvested, promoting harvester‐owned plantations of African mahogany, and increasing survival and growth by promoting no‐fire zones in gallery forests. Our results show how population responses to harvest of NTFP may be altered by ecological differences across sites and emphasize the importance of monitoring populations over the climatic range in which they occur to develop more realistic recommendations for conservation.     

Deconstructing compassionate conservation

Compassionate conservation focuses on 4 tenets: first, do no harm; individuals matter; inclusivity of individual animals; and peaceful coexistence between humans and animals. Recently, compassionate conservation has been promoted as an alternative to conventional conservation philosophy. We believe examples presented by compassionate conservationists are deliberately or arbitrarily chosen to focus on mammals; inherently not compassionate; and offer ineffective conservation solutions. Compassionate conservation arbitrarily focuses on charismatic species, notably large predators and megaherbivores. The philosophy is not compassionate when it leaves invasive predators in the environment to cause harm to vastly more individuals of native species or uses the fear of harm by apex predators to terrorize mesopredators. Hindering the control of exotic species (megafauna, predators) in situ will not improve the conservation condition of the majority of biodiversity. The positions taken by so-called compassionate conservationists on particular species and on conservation actions could be extended to hinder other forms of conservation, including translocations, conservation fencing, and fertility control. Animal welfare is incredibly important to conservation, but ironically compassionate conservation does not offer the best welfare outcomes to animals and is often ineffective in achieving conservation goals. Consequently, compassionate conservation may threaten public and governmental support for conservation because of the limited understanding of conservation problems by the general public.     

Decoupling of component and ensemble density feedbacks in birds and mammals

A component density feedback represents the effect of change in population size on single demographic rates, whereas an ensemble density feedback captures that effect on the overall growth rate of a population. Given that a population's growth rate is a synthesis of the interplay of all demographic rates operating in a population, we test the hypothesis that the strength of ensemble density feedback must augment with increasing strength of component density feedback, using long-term censuses of population size, fertility, and survival rates of 109 bird and mammal populations (97 species). We found that compensatory and depensatory component feedbacks were common (each detected in approximately 50% of the demographic rates). However, component feedback strength only explained <10% of the variation in ensemble feedback strength. To explain why, we illustrate the different sources of decoupling between component and ensemble feedbacks. We argue that the management of anthropogenic impacts on populations using component feedbacks alone is ill-advised, just as managing on the basis of ensemble feedbacks without a mechanistic understanding of the contributions made by its components and environmental variability can lead to suboptimal decisions.     

Accuracy of Short‐Term Demographic Data in Projecting Long‐Term Fate of Populations

Short‐term surveys are useful in conservation of species if they can be used to reliably predict the long‐term fate of populations. However, statistical evaluations of reliability are rare. We studied how well short‐term demographic data (1999–2002) of tartar catchfly (Silene tatarica), a perennial riparian plant, projected the fate and growth of 23 populations of this species up to the year 2010. Surveyed populations occurred along a river with natural flood dynamics and along a regulated river. Riparian plant populations are affected by flooding, which maintains unvegetated shores, while forest succession proceeds in areas with little flooding. Flooding is less severe along the regulated river, and vegetation overgrowth reduces abundance of tartar catchfly on unvegetated shores. We built matrix models to calculate population growth rates and estimated times to population extinction in natural and in regulated rivers, 13 and 10 populations, respectively. Models predicted population survival well (model predictions matched observed survival in 91% of populations) and accurately predicted abundance increases and decreases in 65% of populations. The observed and projected population growth rates differed significantly in all but 3 populations. In most cases, the model overestimated population growth. Model predictions did not improve when data from more years were used (1999–2006). In the regulated river, the poorest model predictions occurred in areas where cover of other plant species changed the fastest. Although vegetation cover increased in most populations, it decreased in 4 populations along the natural river. Our results highlight the need to combine disturbance and succession dynamics in demographic models and the importance of habitat management for species survival along regulated rivers. Precisión de Datos Demográficos de Corto Plazo en la Proyección del Destino de Poblaciones a Largo Plazo     

Conserving Slow-Growing, Long-Lived Tree Species: Input from the Demography of a Rare Understory Conifer, Taxus floridana

Abstract:  Although land preservation and promotion of successful regeneration are important conservation actions, their ability to increase population growth rates of slow-growing, long-lived trees is limited. We investigated the demography of Taxus floridana Nutt., a rare understory conifer, in three populations in different ravine forests spanning its entire geographic range along the Apalachicola River Bluffs in northern Florida (U.S.A.). We examined spatial and temporal patterns in demographic parameters and projected population growth rates by using four years of data on the recruitment and survival of seedlings and established stems, and on diameter growth from cross-sections of dead stems. All populations experienced a roughly 10-fold increase in seedling recruitment in 1996 compared with other years. The fates of seedlings and stems between 8 and 16 mm differed among populations. The fates of stems in two other size classes (the 2- to 4-mm class and the 4- to 8-mm class) differed among both populations and years. Individual stems in all populations exhibited similarly slow growth rates. Stochastic matrix models projected declines in all populations. Stochastic matrix analysis revealed the high elasticity of a measure of stochastic population growth rate to perturbations in the stasis of large reproductive stems for all populations. Additional analyses also indicated that occasional episodes of high recruitment do not greatly affect population growth rates. Conservation efforts directed at long-lived, slow-growing rare plants like Taxus floridana should both protect established reproductive individuals and further enhance survival of individuals in other life-history stages, such as juveniles, that often do not appear to contribute greatly to population growth rates.     

Effect of stage-specific vital rates on population growth rates and effective population sizes in an endangered iteroparous plant

Effective population size (N(e)) determines the strength of genetic drift and can influence the level of genetic diversity a population can maintain. Assessing how changes in demographic rates associated with environmental variables and management actions affect N(e) thus can be crucial to the conservation of endangered species. Calculation of N(e) through demographic models makes it possible to use elasticity analyses to study this issue. The elasticity of N(e) to a given vital rate is the proportional change in N(e) associated with a proportional increase in that vital rate. In addition, demographic models can be used to study N(e) and population growth rate (λ) simultaneously. Simultaneous examination is important because some vital rates differ diametrically in their associations with λ and N(e). For example, in some cases increasing these vital rates increases λ and decreases N(e). We used elasticity analysis to study the effect of stage-specific survival and flowering rates on N(e), annual effective population size (N(a)), and λ in seven populations of the endangered plant Austrian dragonhead (Dracocephalum austriacum). In populations with λ ≥ 1, the elasticities of N(e) and N(a) were similar to those of λ. Survival rates of adults were associated with greater elasticities than survival rates of juveniles, flowering rates, or fecundity. In populations with λ < 1, N(e) and N(a) exhibited greater elasticities to juvenile than to adult vital rates. These patterns are similar to those observed in other species with similar life histories. We did not observe contrasting effects of any vital rate on λ and N(e); thus, management actions that increase the λ of populations of Austrian dragonhead will not increase genetic drift. Our results show that elasticity analyses of N(e) and N(a) can complement elasticity analysis of λ. Moreover, such analyses do not require more data than standard matrix models of population dynamics.     

The virtue of compassion in compassionate conservation

The role of ethics is becoming an increasingly important feature of biodiversity conservation dialogue and practice. Compassionate conservationists argue for a prohibition of, or at least a strong presumption against, the adoption of conservation policies that intentionally harm animals. They assert that to be compassionate is to care about animals and that it is antithetical to caring for animals to intentionally harm them. Compassionate conservationists thus criticize many existing conservation practices and policies. Two things together challenge the philosophical foundation of compassionate conservation. First, compassionate conservationists ground their theory in virtue ethics, yet virtue ethics permits exceptions to moral rules, so there cannot be an in-principle prohibition on adopting intentional harm-inducing policies and practices. But not all compassionate conservationists advocate for a prohibition on intentionally harming animals, only a strong presumption against it. This leads to the second point: compassion can motivate a person to adopt a harm-inducing conservation policy or practice when doing so is the best available option in a situation in which animals will be harmed no matter what policy or practice is adopted. Combining these insights with the empirical observation that conservationists regularly find themselves in tragic situations, we arrive at the conclusion that conservationists may regularly advocate for harm-inducing policies and practices from a position of compassion. Article Impact Statement: Compassionate conservationists should accept that the virtuously compassionate person may adopt harm-causing conservation policies.     

Double Allee Effects and Extinction in the Island Fox

Abstract:  An Allee effect (AE) occurs in populations when individuals suffer a decrease in fitness at low densities. If a fitness component is reduced (component AE), per capita population growth rates may decline as a consequence (demographic AE) and extinction risk is increased. The island fox ( Urocyon littoralis ) is endemic to six of the eight California Channel Islands. Population crashes have coincided with an increase in predation by Golden Eagles ( Aquila chrysaetos ). We propose that AEs could render fox populations more sensitive and may be a likely explanation for their sharp decline. We analyzed demographic data collected between 1988 and 2000 to test whether fox density (1) influences survival and reproductive rates; (2) interacts with eagle presence and affects fox fitness parameters; and (3) influences per capita fox population trends. A double component AE simultaneously influenced survival (of adults and pups) and proportion of breeding adult females. The adult survival AE was driven by predation by eagles. These component AEs led to a demographic AE. Multiple-component AEs, a predation-driven AE, and the simultaneous occurrence of both component and demographic AEs in a mammal are all previously unreported processes. Populations below 7 foxes/km² could have suboptimal population growth rates due to the demographic AE, and AEs may have contributed to the dramatic declines in three fox populations. Because fox densities in critically endangered populations are well below this level, removing Golden Eagles appears necessary to prevent a predation-driven AE. Conservationists should also be aware of AEs when planning the release of captive foxes. More generally, our findings highlight the danger of overlooking AEs in the conservation of populations of rare or threatened species.     

Incorporating diverse data and realistic complexity into demographic estimation procedures for sea otters.

Reliable information on historical and current population dynamics is central to understanding patterns of growth and decline in animal populations. We developed a maximum likelihood-based analysis to estimate spatial and temporal trends in age/sex-specific survival rates for the threatened southern sea otter (Enhydra lutris nereis), using annual population censuses and the age structure of salvaged carcass collections. We evaluated a wide range of possible spatial and temporal effects and used model averaging to incorporate model uncertainty into the resulting estimates of key vital rates and their variances. We compared these results to current demographic parameters estimated in a telemetry-based study conducted between 2001 and 2004. These results show that survival has decreased substantially from the early 1990s to the present and is generally lowest in the north-central portion of the population's range. The greatest temporal decrease in survival was for adult females, and variation in the survival of this age/sex class is primarily responsible for regulating population growth and driving population trends. Our results can be used to focus future research on southern sea otters by highlighting the life history stages and mortality factors most relevant to conservation. More broadly, we have illustrated how the powerful and relatively straightforward tools of information-theoretic-based model fitting can be used to sort through and parameterize quite complex demographic modeling frameworks.     

Demographic Forecasting in Koala Conservation

Abstract: The koala currently needs conservation intervention. There is clear evidence of decline in many populations, but the existence of other stable or expanding populations offers the possibility of a variety of creative solutions to their conservation problems. The 1998 National Koala Conservation Strategy emphasizes the need to obtain demographic information and to use this information to assess management options for koalas. We need accurate diagnoses of the status of koala populations and forecasts of their demographic future with and without particular management actions. In a qualitative fashion, this process has been undertaken many times on a local and national scale. Quantitative demographic forecasting tools are increasingly available, and koala management could benefit from their application both at the scale of individual populations and more broadly. There is already a considerable body of suitable data on the dispersal, effects of normal and catastrophic environmental variation on reproduction and survival, and on the effects of habitat change. Demographic forecasting, however, is hampered because the full suite of information is rarely available from a single population. In two Queensland populations, retrospective population viability analyses provided forecasts that were in agreement with observed population trends. Work is needed to determine whether data from one population can be applied to other populations. Models can then be developed to make projections at a multipopulation level on the basis of local population dynamics and dispersal. Certain koala populations, because of their long history of study, offer the opportunity to test demographic models retrospectively. These tests will not only aid in fine-tuning the models for koala biology and data but will also assist with the more general process of validating the models.     

Emotion as a source of moral understanding in conservation

Recent debates around the meaning and implications of compassionate conservation suggest that some conservationists consider emotion a false and misleading basis for moral judgment and decision making. We trace these beliefs to a long-standing, gendered sociocultural convention and argue that the disparagement of emotion as a source of moral understanding is both empirically and morally problematic. According to the current scientific and philosophical understanding, reason and emotion are better understood as partners, rather than opposites. Nonetheless, the two have historically been seen as separate, with reason elevated in association with masculinity and emotion (especially nurturing emotion) dismissed or delegitimated in association with femininity. These associations can be situated in a broader, dualistic, and hierarchical logic used to maintain power for a dominant male (White, able-bodied, upper class, heterosexual) human class. We argue that emotion should be affirmed by conservationists for the novel and essential insights it contributes to conservation ethics. We consider the specific example of compassion and characterize it as an emotional experience of interdependence and shared vulnerability. This experience highlights conservationists’ responsibilities to individual beings, enhancing established and widely accepted beliefs that conservationists have a duty to protect populations, species, and ecosystems (or biodiversity). We argue compassion, thus understood, should be embraced as a core virtue of conservation.     

Conservation Implications of Drastic Reductions in the Smallest and Most Isolated Populations of Giant Pandas

Abstract: In conservation biology, understanding the causes of endangerment is a key step to devising effective conservation strategies. We used molecular evidence (coalescent simulations of population changes from microsatellite data) and historical information (habitat and human population changes) to investigate how the most‐isolated populations of giant pandas (Ailuropoda melanoleuca) in the Xiaoxiangling Mountains became highly endangered. These populations experienced a strong, recent demographic reduction (60‐fold), starting approximately 250 years BP. Explosion of the human population and use of non‐native crop species at the peak of the Qing Empire resulted in land‐use changes, deforestation, and habitat fragmentation, which are likely to have led to the drastic reduction of the most‐isolated populations of giant pandas. We predict that demographic, genetic, and environmental factors will lead to extinction of giant pandas in the Xiaoxiangling Mountains in the future if the population remains isolated. Therefore, a targeted conservation action—translocation—has been proposed and is being implemented by the Chinese goverment.     

Population inertia and its sensitivity to changes in vital rates and population structure

Because the (st)age structure of a population may rarely be stable, studies of transient population dynamics and population momentum are becoming ever more popular. Yet, studies of "population momentum" are restricted in the sense that they describe the inertia of population size resulting from a demographic transition to the stationary population growth rate. Although rarely mentioned, inertia in population size is a general phenomenon and can be produced by any demographic transition or perturbation. Because population size is of central importance in demography, conservation, and management, formulas relating the sensitivity of population inertia to changes in underlying vital rates and population structure could provide much-needed insight into the dynamics of populations with unstable (st)age structure. Here, we derive such formulas, which are readily computable, and provide examples of their potential use in studies of life history and applied arenas of population study.     

The moral residue of conservation

Should conservationists use lethal management to control introduced wildlife populations? Should they kill individual animals to protect endangered species? Are trade-offs that prioritize some values at the expense of others morally appropriate? These sorts of ethical questions are common in conservation. In debating such questions, conservationists often seem to presume 1 of 2 possible answers: the act in question is right or it is wrong. But morality in conservation is considerably more complex than this simple binary suggests. A robust conservation ethic requires a vocabulary that gives voice to the uncertainty and unease that arise when what seems to be the best available course of action also seems to involve a measure of wrongdoing. The philosophical literature on moral residue and moral dilemmas supplies this vocabulary. Moral dilemmas arise when one must neglect certain moral requirements to fulfill others. Under such circumstances, even the best possible decision leaves a moral residue, which is experienced emotionally as some form of grief. Examples of conservation scenarios that leave a moral residue include management of introduced rabbits in Australia, trophy hunting in Africa, and forest management trade-offs in the Pacific Northwest. Moral residue is integral to the moral experience of conservationists today, and grief is an appropriate response to many decisions conservationists must make. Article impact statement: Defensible conservation decisions may neglect moral requirements, leaving a moral residue; conservationists should respond with grief.     

Reliability of Conservation Actions Based on Elasticity Analysis of Matrix Models

Abstract: Matrix population models have entered the mainstream of conservation biology, with analysis of proportional sensitivities (elasticity analysis) of demographic rates becoming important components of conservation decision making. We identify areas where management applications using elasticity analysis potentially conflict with the mathematical basis of the technique, and we use a hypothetical example and three real data sets (Prairie Chicken [   Tympanuchus cupido ], desert tortoise [ Gopherus agassizii ], and killer whale [ Orcinus orca ]) to evaluate the extent to which conservation recommendations based on elasticities might be misleading. First, changes in one demographic rate can change the qualitative ranking of the elasticity values calculated from a population matrix, a result that dampens enthusiasm for ranking conservation actions based solely on which rates have the highest elasticity values. Second, although elasticities often provide accurate predictions of future changes in population growth rate under management perturbations that are large or that affect more than one rate concurrently, concordance frequently fails when different rates vary by different amounts. In particular, when vital rates change to their high or low values observed in nature, predictions of future growth rate based on elasticities of a mean matrix can be misleading, even predicting population increase when the population growth rate actually declines following a perturbation. Elasticity measures will continue to be useful tools for applied ecologists, but they should be interpreted with considerable care. We suggest that studies using analytical elasticity analysis explicitly consider the range of variation possible for different rates and that simulation methods are a useful tool to this end.     

Use of Traditional Ecological Knowledge in Marine Conservation

Abstract:  Traditional ecological knowledge (TEK) represents multiple bodies of knowledge accumulated through many generations of close interactions between people and the natural world. TEK and its application via customary ecological management plans can be useful in modern conservation programs. I disaggregate the term TEK into its constituent parts and show several ways in which TEK can strengthen research designs by increasing locality-specific knowledge, including environmental linkages occurring in those localities. Examples of the uses of TEK in conservation include folk taxonomy in systematics in Micronesia, species knowledge for conservation in Kiribati, and fishers' knowledge of ecological interactions for reserve design in Belize. When conservationists recognize the utility of TEK, they can engage in an equitable exchange of knowledge and foster shared responsibility with indigenous people. These types of exchanges can also provide an opportunity for indigenous people to develop a scientific infrastructure.     

Demography of central and marginal populations of monkeyflowers (Mimulus cardinalis and M. lewisii)

Every species occupies a limited geographic area, but how spatiotemporal environmental variation affects individual and population fitness to create range limits is not well understood. Because range boundaries arise where, on average, populations are more likely to go extinct than to persist, range limits are an inherently population-level problem for which a demographic framework is useful. In this study, I compare demographic parameters and population dynamics between central and marginal populations of monkeyflowers, Mimulus cardinalis and M. lewisii, along an elevation gradient spanning both species' ranges. Central and marginal populations of both species differed in survival and fecundity. For M. lewisii, these components of fitness were higher in central than in marginal populations, but for M. cardinalis the converse was true. To assess spatiotemporal variation in population dynamics, I used transition matrix models to estimate asymptotic population growth rates (lambda) and found that population growth rates of M. lewisii were highest at the range center and reduced at the range margin. Population growth rates of M. cardinalis were highest at the range margin and greatly reduced at the range center. Life table response analysis decomposed spatiotemporal variation in lambda into contributions from each transition between life stages, finding that transitions from large nonreproductive and reproductive plants to the seed class and stasis in the reproductive class made the largest contributions to spatial differences in lambda. These transitions had only low to moderate sensitivities, indicating that differences in projected population growth rates resulted mainly from observed differences in transition matrix parameters and their underlying vital rates.     

Estimating the functional form for the density dependence from life history data

Two contrasting approaches to the analysis of population dynamics are currently popular: demographic approaches where the associations between demographic rates and statistics summarizing the population dynamics are identified; and time series approaches where the associations between population dynamics, population density, and environmental covariates are investigated. In this paper, we develop an approach to combine these methods and apply it to detailed data from Soay sheep (Ovis aries). We examine how density dependence and climate contribute to fluctuations in population size via age- and sex-specific demographic rates, and how fluctuations in demographic structure influence population dynamics. Density dependence contributes most, followed by climatic variation, age structure fluctuations and interactions between density and climate. We then simplify the density-dependent, stochastic, age-structured demographic model and derive a new phenomenological time series which captures the dynamics better than previously selected functions. The simple method we develop has potential to provide substantial insight into the relative contributions of population and individual-level processes to the dynamics of populations in stochastic environments.     

Balancing making a difference with making a living in the conservation sector

Goals play important roles in people's lives because they focus attention, mobilize effort, and sustain motivation. Understanding conservationists’ satisfaction with goal progress may provide insights into real-world environmental trends and flag risks to their well-being and motivation. We asked 2694 conservationists working globally how satisfied they were with progress toward goals important to them. We then explored how this satisfaction varied among groups, including demographic and occupational. Finally, we looked at respondents' experiences associated with goal-progress satisfaction. Many (94.0%) indicated that making a meaningful contribution to conservation was an important goal for them, and over half were satisfied or very satisfied in this area (52.5%). However, respondents were generally dissatisfied with progress on collective conservation goals (e.g., stopping species loss). Some groups were more likely to report dissatisfaction than others. For instance, those in conservation for longer tended to be less satisfied with collective goal progress (log odds –0.21, 95% credibility interval [CI] –0.32 to –0.10), but practitioners reported greater satisfaction (log odds 0.38, 95% CI 0.15–0.60). Likewise, those who were more optimistic in life (log odds 0.24, 95% CI 0.17–0.32), male (log odds 0.25, 95% CI 0.10–0.41), and working in conservation practice (log odds 0.25, 95% CI 0.08–0.43) reported greater satisfaction with individual goal progress. Free-text responses suggested widespread dissatisfaction with livelihood goals, particularly related to job security and adequate compensation. Although contributing to conservation appeared to be a source of satisfaction, slow goal progress in other areas––particularly around making a living––looked to be a source of distress and demotivation. Employers, funders, professional societies, and others should consider ways to help those in the sector make a difference while making a satisfactory living by, for example, prioritizing conservationists′ well-being when allocating funding. This support could include avoiding exploitative practices, fostering supportive work environments, and celebrating positive outcomes.     

Demographic Side Effects of Selective Hunting in Ungulates and Carnivores

Abstract:  Selective harvesting regimes are often implemented because age and sex classes contribute differently to population dynamics and hunters show preferences associated with body size and trophy value. We reviewed the literature on how such cropping regimes affect the demography of the remaining population (here termed demographic side effects ). First, we examined the implications of removing a large proportion of a specific age or sex class. Such harvesting strategies often bias the population sex ratio toward females and reduce the mean age of males, which may consequently delay birth dates, reduce birth synchrony, delay body mass development, and alter offspring sex ratios. Second, we reviewed the side effects associated with the selective removal of relatively few specific individuals, often large trophy males. Such selective harvesting can destabilize social structures and the dominance hierarchy and may cause loss of social knowledge, sexually selected infanticide, habitat changes among reproductive females, and changes in offspring sex ratio. A common feature of many of the reported mechanisms is that they ultimately depress recruitment and in some extreme cases even cause total reproductive collapse. These effects could act additively and destabilize the dynamics of populations, thus having a stronger effect on population growth rate than first anticipated. Although more experimental than observational studies reported demographic side effects, we argue that this may reflect the quite subtle mechanisms involved, which are unlikely to be detected in observational studies without rigorous monitoring regimes. We call for more detailed studies of hunted populations with marked individuals that address how the expression of these effects varies across mating systems, habitats, and with population density. Theoretical models investigating how strongly these effects influence population growth rates are also required.