Ecosystem service flows from a migratory species: Spatial subsidies of the northern pintail.

Migratory species provide important benefits to society, but their cross-border conservation poses serious challenges. By quantifying the economic value of ecosystem services (ESs) provided across a species' range and ecological data on a species' habitat dependence, we estimate spatial subsidies-how different regions support ESs provided by a species across its range. We illustrate this method for migratory northern pintail ducks in North America. Pintails support over $101 million USD annually in recreational hunting and viewing and subsistence hunting in the U.S. and Canada. Pintail breeding regions provide nearly $30 million in subsidies to wintering regions, with the "Prairie Pothole" region supplying over $24 million in annual benefits to other regions. This information can be used to inform conservation funding allocation among migratory regions and nations on which the pintail depends. We thus illustrate a transferrable method to quantify migratory species-derived ESs and provide information to aid in their transboundary conservation.

Willingness to Pay for Conservation of Transborder Migratory Species: A Case Study of the Mexican Free-Tailed Bat in the United States and Mexico.

We estimated U.S. and Mexican citizens' willingness to pay (WTP) for protecting habitat for a transborder migratory species, the Mexican free-tailed bat (Tadarida brasiliensis mexicana), using the contingent valuation method. Few contingent valuation surveys have evaluated whether households in one country would pay to protect habitat in another country. This study addresses that gap. In our study, Mexican respondents were asked about their WTP for conservation of Mexican free-tailed bat habitat in Mexico and in the United States. Similarly, U.S. respondents were asked about their WTP for conservation in the United States and in Mexico. U.S. households would pay $30 annually to protect habitat in the United States and $24 annually to protect habitat in Mexico. Mexican households would pay $8 annually to protect habitat in Mexico and $5 annually to protect habitat in the United States. In both countries, these WTP amounts rose significantly for increasing the size of the bat population rather than simply stabilizing the current bat population. The ratio of Mexican household WTP relative to U.S. household WTP is nearly identical to that of Mexican household income relative to U.S. household income. This suggests that the perceived economic benefits received from the bats is similar in Mexico and the United States, and that scaling WTP by relative income in international benefit transfer may be plausible.

Quantitative tools for implementing the new definition of significant portion of the range in the U.S. Endangered Species Act.

In 2014, the Fish and Wildlife Service (FWS) and National Marine Fisheries Service announced a new policy interpretation for the U.S. Endangered Species Act (ESA). According to the act, a species must be listed as threatened or endangered if it is determined to be threatened or endangered in a significant portion of its range (SPR). The 2014 policy seeks to provide consistency by establishing that a portion of the range should be considered significant if the associated individuals' "removal would cause the entire species to become endangered or threatened." We reviewed 20 quantitative techniques used to assess whether a portion of a species' range is significant according to the new guidance. Our assessments are based on the 3R criteria-redundancy (i.e., buffering from catastrophe), resiliency (i.e., ability to withstand stochasticity), and representation (i.e., ability to evolve)-that the FWS uses to determine if a species merits listing. We identified data needs for each quantitative technique and considered which methods could be implemented given the data limitations typical of rare species. We also identified proxies for the 3Rs that may be used with limited data. To assess potential data availability, we evaluated 7 example species by accessing data in their species status assessments, which document all the information used during a listing decision. In all species, an SPR could be evaluated with at least one metric for each of the 3Rs robustly or with substantial assumptions. Resiliency assessments appeared most constrained by limited data, and many species lacked information on connectivity between subpopulations, genetic variation, and spatial variability in vital rates. These data gaps will likely make SPR assessments for species with complex life histories or that cross national boundaries difficult. Although we reviewed techniques for the ESA, other countries require identification of significant areas and could benefit from this research.

Recreation economics to inform migratory species conservation: Case study of the northern pintail.

Quantification of the economic value provided by migratory species can aid in targeting management efforts and funding to locations yielding the greatest benefits to society and species conservation. Here we illustrate a key step in this process by estimating hunting and birding values of the northern pintail (Anas acuta) within primary breeding and wintering habitats used during the species' annual migratory cycle in North America. We used published information on user expenditures and net economic values (consumer surplus) for recreational viewing and hunting to determine the economic value of pintail-based recreation in three primary breeding areas and two primary wintering areas. Summed expenditures and consumer surplus for northern pintail viewing were annually valued at $70M, and annual sport hunting totaled $31M (2014 USD). Expenditures for viewing ($42M) were more than twice as high than those for hunting ($18M). Estimates of consumer surplus, defined as the amount consumers are willing to pay above their current expenditures, were $15M greater for viewing ($28M) than for hunting ($13M). We discovered substantial annual consumer surplus ($41M) available for pintail conservation from birders and hunters. We also found spatial differences in economic value among the primary regions used by pintails, with viewing generally valued more in breeding regions than in wintering regions and the reverse being true for hunting. The economic value of pintail-based recreation in the Western wintering region ($26M) exceeded that in any other region by at least a factor of three. Our approach of developing regionally explicit economic values can be extended to other taxonomic groups, and is particularly suitable for migratory game birds because of the availability of large amounts of data. When combined with habitat-linked population models, regionally explicit values could inform development of more effective conservation finance and policy mechanisms to enhance environmental management and societal benefits across the geographically dispersed areas used by migratory species.

Monarch butterfly population decline in North America: identifying the threatening processes.

The monarch butterfly (Danaus plexippus) population in North America has sharply declined over the last two decades. Despite rising concern over the monarch butterfly's status, no comprehensive study of the factors driving this decline has been conducted. Using partial least-squares regressions and time-series analysis, we investigated climatic and habitat-related factors influencing monarch population size from 1993 to 2014. Potential threats included climatic factors, habitat loss (milkweed and overwinter forest), disease and agricultural insecticide use (neonicotinoids). While climatic factors, principally breeding season temperature, were important determinants of annual variation in abundance, our results indicated strong negative relationships between population size and habitat loss variables, principally glyphosate use, but also weaker negative effects from the loss of overwinter forest and breeding season use of neonicotinoids. Further declines in population size because of glyphosate application are not expected. Thus, if remaining threats to habitat are mitigated we expect climate-induced stochastic variation of the eastern migratory population of monarch butterfly around a relatively stationary population size.

Density estimates of monarch butterflies overwintering in central Mexico.

Given the rapid population decline and recent petition for listing of the monarch butterfly (Danaus plexippus L.) under the Endangered Species Act, an accurate estimate of the Eastern, migratory population size is needed. Because of difficulty in counting individual monarchs, the number of hectares occupied by monarchs in the overwintering area is commonly used as a proxy for population size, which is then multiplied by the density of individuals per hectare to estimate population size. There is, however, considerable variation in published estimates of overwintering density, ranging from 6.9-60.9 million ha-1. We develop a probability distribution for overwinter density of monarch butterflies from six published density estimates. The mean density among the mixture of the six published estimates was ∼27.9 million butterflies ha-1 (95% CI [2.4-80.7] million ha-1); the mixture distribution is approximately log-normal, and as such is better represented by the median (21.1 million butterflies ha-1). Based upon assumptions regarding the number of milkweed needed to support monarchs, the amount of milkweed (Asclepias spp.) lost (0.86 billion stems) in the northern US plus the amount of milkweed remaining (1.34 billion stems), we estimate >1.8 billion stems is needed to return monarchs to an average population size of 6 ha. Considerable uncertainty exists in this required amount of milkweed because of the considerable uncertainty occurring in overwinter density estimates. Nevertheless, the estimate is on the same order as other published estimates. The studies included in our synthesis differ substantially by year, location, method, and measures of precision. A better understanding of the factors influencing overwintering density across space and time would be valuable for increasing the precision of conservation recommendations.

Quasi-extinction risk and population targets for the Eastern, migratory population of monarch butterflies (Danaus plexippus).

The Eastern, migratory population of monarch butterflies (Danaus plexippus), an iconic North American insect, has declined by ~80% over the last decade. The monarch's multi-generational migration between overwintering grounds in central Mexico and the summer breeding grounds in the northern U.S. and southern Canada is celebrated in all three countries and creates shared management responsibilities across North America. Here we present a novel Bayesian multivariate auto-regressive state-space model to assess quasi-extinction risk and aid in the establishment of a target population size for monarch conservation planning. We find that, given a range of plausible quasi-extinction thresholds, the population has a substantial probability of quasi-extinction, from 11-57% over 20 years, although uncertainty in these estimates is large. Exceptionally high population stochasticity, declining numbers, and a small current population size act in concert to drive this risk. An approximately 5-fold increase of the monarch population size (relative to the winter of 2014-15) is necessary to halve the current risk of quasi-extinction across all thresholds considered. Conserving the monarch migration thus requires active management to reverse population declines, and the establishment of an ambitious target population size goal to buffer against future environmentally driven variability.

Market forces and technological substitutes cause fluctuations in the value of bat pest-control services for cotton.

Critics of the market-based, ecosystem services approach to biodiversity conservation worry that volatile market conditions and technological substitutes will diminish the value of ecosystem services and obviate the "economic benefits" arguments for conservation. To explore the effects of market forces and substitutes on service values, we assessed how the value of the pest-control services provided by Mexican free-tailed bats (Tadarida brasiliensis mexicana) to cotton production in the southwestern U.S. has changed over time. We calculated service values each year from 1990 through 2008 by estimating the value of avoided crop damage and the reduced social and private costs of insecticide use in the presence of bats. Over this period, the ecosystem service value declined by 79% ($19.09 million U.S. dollars) due to the introduction and widespread adoption of Bt (Bacillus thuringiensis) cotton transgenically modified to express its own pesticide, falling global cotton prices and the reduction in the number of hectares in the U.S. planted with cotton. Our results demonstrate that fluctuations in market conditions can cause temporal variation in ecosystem service values even when ecosystem function--in this case bat population numbers--is held constant. Evidence is accumulating, however, of the evolution of pest resistance to Bt cotton, suggesting that the value of bat pest-control services may increase again. This gives rise to an economic option value argument for conserving Mexican free-tailed bat populations. We anticipate that these results will spur discussion about the role of ecosystem services in biodiversity conservation in general, and bat conservation in particular.

When ecosystem services crash: preparing for big, fast, patchy climate change.

Assessments of adaptation options generally focus on incremental, homogeneous ecosystem responses to climate even though climate change impacts can be big, fast, and patchy across a region. Regional drought-induced tree die-off in semiarid woodlands highlights how an ecosystem crash fundamentally alters most ecosystem services and poses management challenges. Building on previous research showing how choice of location is linked to adaptive capacity and vulnerability, we developed a framework showing how the options for retaining desired ecosystem services in the face of sudden crashes depend on how portable the service is and whether the stakeholder is flexible with regard to the location where they receive their services. Stakeholders using portable services, or stakeholders who can move to other locations to obtain services, may be more resilient to ecosystem crashes. Our framework suggests that entering into cooperative networks with regionally distributed stakeholders is key to building resilience to big, fast, patchy crashes.

Salinity and light interactively affect neotropical mangrove seedlings at the leaf and whole plant levels.

We have studied the interactive effects of salinity and light on Avicennia germinans mangrove seedlings in greenhouse and field experiments. We hypothesized that net photosynthesis, growth, and survivorship rates should increase more with an increase in light availability for plants growing at low salinity than for those growing at high salinity. This hypothesis was supported by our results for net photosynthesis and growth. Net daily photosynthesis did increase more with increasing light for low-salinity plants than for high-salinity plants. Stomatal conductance, leaf-level transpiration, and internal CO(2) concentrations were lower at high than at low salinity. At high light, the ratio of leaf respiration to assimilation was 2.5 times greater at high than at low salinity. Stomatal limitations and increased respiratory costs may explain why, at high salinity, seedlings did not respond to increased light availability with increased net photosynthesis. Seedling mass and growth rates increased more with increasing light availability at low than at high salinity. Ratios of root mass to leaf mass were higher at high salinity, suggesting that either water or nutrient limitations may have limited seedling growth at high salinity in response to increasing light. The interactive effects of salinity and light on seedling size and growth rates observed in the greenhouse were robust in the field, despite the presence of other factors in the field--such as inundation, nutrient gradients, and herbivory. In the field, seedling survivorship was higher at low than at high salinity and increased with light availability. Interestingly, the positive effect of light on seedling survivorship was stronger at high salinity, indicating that growth and survivorship rates are decoupled. In general, this study demonstrates that environmental effects at the leaf-level also influence whole plant growth in mangroves.