Statistical Power of Presence-Absence Data to Detect Population Declines

Abstract: Population declines may be inferred from a decrease in the number of sites at which a species is detected. Although such presence-absence data often are interpreted informally, it is simple to test the statistical significance of changes in the number of sites occupied by a species. I used simulations to examine the statistical power (i.e., the probability of making the Type II error that no population decline has occurred when the population actually has declined) of presence-absence designs. Most presence-absence designs have low power to detect declines of < 20–50% in populations but have adequate power to detect steeper declines. Power was greater if the population disappeared entirely from a subset of formerly occupied sites than if it declined evenly over its entire range. Power also rose with (1) increases in the number of sites surveyed; (2) increases in population density or sampling effort at a site; and (3) decreases in spatial variance in population density. Because of potential problems with bias and inadequate power, presence-absence designs should be used and interpreted cautiously.     

Population Declines and Priorities for Amphibian Conservation in Latin America

Abstract: Although dramatic amphibian population declines have been reported worldwide, our understanding of the extent of the declines in Latin America, where amphibian diversity is high, is limited to a few well-documented studies. To better understand the geographic extent of declines, their possible causes, and the measures needed to improve Latin American scientists' ability to research the phenomenon and make effective management recommendations, we convened three regional workshops with 88 Latin American herpetologists and conservationists. Population declines are widespread in Latin America. At least 13 countries have experienced declines, and in 40 cases species are now thought to be extinct or extirpated in a country where they once occurred. Declines or extinctions have affected 30 genera and nine families of amphibians. Most declines have occurred in remote highlands, above 500 m in elevation in Central America and above 1000 m in the Andes. Most documented declines occurred in the 1980s. Of the possible causes studied to date, climate change appears to be important at one site and chytrid fungal disease has been identified at sites in three countries. Although many monitoring studies are currently underway in a variety of habitats, most studies are recent and of short duration. In a signed resolution, workshop participants called for greater collaboration and communication among scientists working in Latin America to understand the geographic extent of population declines and the distribution of possible causal factors. In situ conservation is important to protect habitats, but captive-rearing programs for species subject to imminent extinction are also needed. Better understanding of the taxonomy and natural history of amphibians and more funding for research and monitoring are critical to developing a scientific basis for management action to arrest and reverse population declines.     

Population Declines and Range Contractions among Lowland Farmland Birds in Britain

We used extensive atlas and census data to assess trends in the distribution and population levels of birds on lowland farmland in Britain between the late 1960s and early 1990s. Many species of farmland birds have become less widespread or have declined in numbers, or both, but few have become more wide-spread or have increased. Of the 28 species classified as farmland birds the distributions of 24 contracted between 1970 and 1990. Of the 18 farmland species for which it was possible to assess population change, 15 were less abundant in 1990 than in 1970. Seven of the species were estimated to have undergone population decreases of at least 50%. Farmland species showing the largest population declines tended also to show substantial range contractions. Farmland species underwent an appreciably larger contraction of distribution than species associated with any other habitat. Furthermore, farmland species tended to decrease in abundance, whereas woodland species tended to increase. Population declines among farmland birds became evident in the mid- to late 1970s, a period when several fundamental changes were taking place in British agricultural practices. These included a great reduction in the spring sowing of cereals, a simplification of crop rotations, increased use of chemical pesticides and inorganic fertilizers, and more-intensive grassland management. We suggest that the declines of farmland bird species have been caused or aggravated by this pervasive intensification of agriculture. Existing research on declining farmland birds, however, indicates that there is no single mechanism underlying the population changes. We identify priorities for research, focusing mainly on relationships between bird populations and agricultural practices, but we also recognize a need for a better understanding of the role of predation.     

Association of Coloration Mode with Population Declines and Endangerment in Australian Frogs

Abstract: Successful protection of biodiversity requires increased understanding of the ecological characteristics that predispose some species to endangerment. Theory posits that species with polymorphic or variable coloration should have larger distributions, use more diverse resources, and be less vulnerable to population declines and extinctions, compared with taxa that do not vary in color. We used information from literature on 194 species of Australian frogs to search for associations of coloration mode with ecological variables. In general, species with variable or polymorphic color patterns had larger ranges, used more habitats, were less prone to have a negative population trend, and were estimated as less vulnerable to extinction compared with nonvariable species. An association of variable coloration with lower endangerment was also evident when we controlled statistically for the effects of range size. Nonvariable coloration was not a strong predictor of endangerment, and information on several characteristics is needed to reliably identify and protect species that are prone to decline and may become threatened by extinction in the near future. Analyses based on phylogenetic‐independent contrasts did not support the hypothesis that evolutionary transitions between nonvariable and variable or polymorphic coloration have been accompanied by changes in the ecological variables we examined. Irrefutable demonstration of a role of color pattern variation in amphibian decline and in the dynamics and persistence of populations in general will require a manipulative experimental approach.     

Mass Mortality and Population Declines of Anurans at an Upland Site in Western Panama

I report an episode of anuran mortality and decline in the Reserva Forestal Fortuna, Chiriquí, Panama. The symptoms of decline at this site include population reductions, the presence of dead or dying adults, and tadpole abnormalities. Streamside anurans were abundant and diverse in 1993–1995, were restricted to a few streams in December 1996–January 1997, and then became rare throughout the reserve in July–August 1997. Between December 1996 and January 1997, I found 54 dead or dying frogs belonging to 10 species, and 12% of tadpoles had abnormalities of the oral disc. In July–August 1997 I monitored nine streams 37 times and captured only six individuals, whereas 13 terrestrial surveys along five trails resulted in 18 captured individuals. No dead or dying animals were found during those two months, but 11% of tadpoles had mouthpart abnormalities. Necropsies revealed that 18 of 18 dead anurans had a fungal infection of the skin; because this fungus was the only infection shared among all dead frogs, I suggest that it killed them and contributed to the decline of these populations. The presence of mouthpart abnormalities during a period of adult mortality suggests that this symptom may also be linked to the fungus infection. Clinical signs of decline in the anurans of Fortuna are similar to those found in the anurans of Monteverde and Las Tablas, Costa Rica, and I hypothesize that this pathogen was involved in the declines at all three sites.     

Detecting Population Declines over Large Areas with Presence-Absence, Time-to-Encounter, and Count Survey Methods

Abstract:  Ecologists often discount presence-absence surveys as a poor way to gain insight into population dynamics, in part because these surveys are not amenable to many standard statistical tests. Still, presence-absence surveys are sometimes the only feasible alternative for monitoring large areas when funds are limited, especially for sparse or difficult-to-detect species. I undertook a detailed simulation study to compare the power of presence-absence, count, and time-to-encounter surveys to detect regional declines in a population. I used a modeling approach that simulates both population numbers and the monitoring process, accounting for observation and other measurement errors. In gauging the efficacy of presence-absence surveys versus other approaches, I varied the number of survey sites, the spatial variation in encounter rate, the mean encounter rate, and the type of population loss. My results showed that presence-absence data can be as or more powerful than count data in many cases. Quantitative guidelines for choosing between presence-absence surveys and count surveys depend on the biological and logistical constraints governing a conservation monitoring situation. Generally, presence-absence surveys work best when there is little variability in abundance among the survey sites, the organism is rare, and the species is difficult to detect so that the time spent getting to each survey site is less than or equal to the time spent surveying each site. Count surveys work best otherwise. I present a case study with count data on the Northern Flicker ( Colaptes auratus ) from the North American Breeding Bird Survey to illustrate how the method might be used with field-survey data. The case study demonstrates that a count survey would be the most cost-effective design but would entail reduction in the number of sites. If this site reduction is not desirable, a presence-absence survey would be the most cost-effective survey.     

Population Declines, Viable Breeding Areas, and Management Options for Flamingos in Southern Africa

Habitat rehabilitation or intervention to prevent species declines are rarely employed in Africa. I argue that despite protection in national parks, active intervention is necessary to halt declines in southern Africa's Greater ( Phoenicopterus ruber ) and Lesser ( Phoeniconaias minor ) Flamingo populations. Flamingos are long-lived species that breed sporadically at only two localities in southern Africa: the Makgadikgadi Pans in Botswana and Etosha National Park in Namibia. Despite well-publicized breeding on Etosha Pan, flamingos have experienced only three major breeding events in 40 years. Breeding failure occurs when high evaporation rates rapidly dry the pan, and up to 100,000 flightless chicks may starve. Consequently, pairs breeding in Etosha exhibit extraordinarily low recruitment (0.040 young pair/year) and extrapolations indicate that adults can replace themselves only if they breed for 38 to 50 years and all offspring survive. Because survival of offspring from fledging to adulthood (5 years) is about 46%, this breeding lifespan rises to an unrealistic 83 years, making Etosha a nonviable breeding site. Alternative, suitable flamingo habitats in Africa are being mined for soda-ash, are damaged by pollution, or are unprotected. Accordingly, continent-wide estimates and those from southern Africa alone suggest a population decline of about 40% in both species over the last 15 years. Because Namibia regularly supports 84% of the Greater and 93% of the Lesser Flamingos in southern Africa, conservation strategies are best focused there. Simple but effective management methods, based on those employed in western Europe, could reverse these downward trends. In Etosha a small island surrounded by a water-filled depression would allow up to 4000 pairs to breed annually. The benefits of enhancing the breeding of flamingos in Etosha include research opportunities, tourism revenue, and a safe haven for two Red Data species.     

Habitat Split as a Cause of Local Population Declines of Amphibians with Aquatic Larvae

Abstract:  Most amphibian species have biphasic life histories and undergo an ontogenetic shift from aquatic to terrestrial habitats. In deforested landscapes, streams and forest fragments are frequently disjunct, jeopardizing the life cycle of forest-associated amphibians with aquatic larvae. We tested the impact of habitat split—defined as human-induced disconnection between habitats used by different life-history stages of a species—on four forest-associated amphibian species in a severely fragmented landscape of the Brazilian Atlantic Forest. We surveyed amphibians in forest fragments with and without streams (referred to as wet and dry fragments, respectively), including the adjacent grass-field matrix. Our comparison of capture rates in dry fragments and nearby streams in the matrix allowed us to evaluate the number of individuals that engaged in high-risk migrations through nonforested habitats. Adult amphibians moved from dry fragments to matrix streams at the beginning of the rainy season, reproduced, and returned at the end of the breeding period. Juveniles of the year moved to dry fragments along with adults. These risky reproductive migrations through nonforested habitats that expose individuals to dehydration, predation, and other hazards may cause population declines in dry fragments. Indeed, capture rates were significantly lower in dry fragments compared with wet fragments. Declining amphibians would strongly benefit from investments in the conservation and restoration of riparian vegetation and corridors linking breeding and nonbreeding areas.     

Detecting Insect Pollinator Declines on Regional and Global Scales

Recently there has been considerable concern about declines in bee communities in agricultural and natural habitats. The value of pollination to agriculture, provided primarily by bees, is >$200 billion/year worldwide, and in natural ecosystems it is thought to be even greater. However, no monitoring program exists to accurately detect declines in abundance of insect pollinators; thus, it is difficult to quantify the status of bee communities or estimate the extent of declines. We used data from 11 multiyear studies of bee communities to devise a program to monitor pollinators at regional, national, or international scales. In these studies, 7 different methods for sampling bees were used and bees were sampled on 3 different continents. We estimated that a monitoring program with 200–250 sampling locations each sampled twice over 5 years would provide sufficient power to detect small (2–5%) annual declines in the number of species and in total abundance and would cost U.S.$2,000,000. To detect declines as small as 1% annually over the same period would require >300 sampling locations. Given the role of pollinators in food security and ecosystem function, we recommend establishment of integrated regional and international monitoring programs to detect changes in pollinator communities. Detección de Declinaciones de Insectos Polinizadores a Escalas Regional y Global     

Sea-Cage Aquaculture, Sea Lice, and Declines of Wild Fish

Abstract: A sea cage, sometimes referred to as a net pen, is an enclosure designed to prevent farm fish from escaping and to protect them from large predators, while allowing a free flow of water through the cage to carry away waste. Farm fish thus share water with wild fish, which enables transmission of parasites, such as sea lice, from wild to farm and farm to wild fishes. Sea lice epidemics, together with recently documented population‐level declines of wild salmon in areas of sea‐cage farming, are a reminder that sea‐cage aquaculture is fundamentally different from terrestrial animal culture. The difference is that sea cages protect farm fish from the usual pathogen‐control mechanisms of nature, such as predators, but not from the pathogens themselves. A sea cage thus becomes an unintended pathogen factory. Basic physical theory explains why sea‐cage aquaculture causes sea lice on sympatric wild fish to increase and why increased lice burdens cause wild fish to decline, with extirpation as a real possibility. Theory is important to this issue because slow declines of wild fish can be difficult to detect amid large fluctuations from other causes. The important theoretical concepts are equilibrium, host‐density effect, reservoir‐host effect, and critical stocking level of farmed fish (stocking level at which lice proliferate on farm fish even if wild fish are not present to infect them). I explored these concepts and their implications without mathematics through examples from salmon farming. I also considered whether the lice‐control techniques used by sea‐cage farmers (medication and shortened grow‐out times) are capable of protecting wild fish. Elementary probability showed that (where W is the abundance of wild fish, W* is the prefarm abundance, F is the abundance of farm fish, and is the ratio of lice per farm fish to lice per wild fish). Declines of wild fish can be reduced by short growing cycles for farm fish, medicating farm fish, and keeping farm stocking levels low. Declines can be avoided only by ensuring that wild fish do not share water with farmed fish, either by locating sea cages very far from wild fish or through the use of closed‐containment aquaculture systems. These principles are likely to govern any aquaculture system where cage‐protected farm hosts and sympatric wild hosts have a common parasite with a direct life cycle.     

A Strategy for Monitoring and Managing Declines in an Amphibian Community

Although many taxa have declined globally, conservation actions are inherently local. Ecosystems degrade even in protected areas, and maintaining natural systems in a desired condition may require active management. Implementing management decisions under uncertainty requires a logical and transparent process to identify objectives, develop management actions, formulate system models to link actions with objectives, monitor to reduce uncertainty and identify system state (i.e., resource condition), and determine an optimal management strategy. We applied one such structured decision‐making approach that incorporates these critical elements to inform management of amphibian populations in a protected area managed by the U.S. National Park Service. Climate change is expected to affect amphibian occupancy of wetlands and to increase uncertainty in management decision making. We used the tools of structured decision making to identify short‐term management solutions that incorporate our current understanding of the effect of climate change on amphibians, emphasizing how management can be undertaken even with incomplete information. Estrategia para Monitorear y Manejar Disminuciones en una Comunidad de Anfibios     

Declines in Woodland Salamander Abundance Associated with Non-Native Earthworm and Plant Invasions

Abstract:  Factors that negatively affect the quality of wildlife habitat are a major concern for conservation. Non-native species invasions, in particular, are perceived as a global threat to the quality of wildlife habitat. Recent evidence indicates that some changes to understory plant communities in northern temperate forests of North America, including invasions by 3 non-native plant species, are facilitated by non-native earthworm invasion. Furthermore, non-native earthworm invasions cause a reduction in leaf litter on the forest floor, and the loss of forest leaf litter is commonly associated with declines in forest fauna, including amphibians. We conducted a mark-recapture study of woodland salamander abundance across plant invasion fronts at 10 sites to determine whether earthworm or plant invasions were associated with reduced salamander abundance. Salamander abundance declined exponentially with decreasing leaf litter volume. There was no significant relationship between invasive plant cover and salamander abundance, independent of the effects of leaf litter loss due to earthworm invasion. An analysis of selected salamander prey abundance (excluding earthworms) at 4 sites showed that prey abundance declined with declining leaf litter. The loss of leaf litter layers due to non-native earthworm invasions appears to be negatively affecting woodland salamander abundance, in part, because of declines in the abundance of small arthropods that are a stable resource for salamanders. Our results demonstrate that earthworm invasions pose a significant threat to woodland amphibian fauna in the northeastern United States, and that plant invasions are symptomatic of degraded amphibian habitat but are not necessarily drivers of habitat degradation.     

Integrating the Metapopulation and Habitat Paradigms for Understanding Broad-Scale Declines of Species

Abstract:  Caughley (1994) argued that researchers working on threatened populations tended to follow the "small population paradigm" or the "declining population paradigm," and that greater integration of these paradigms was needed. Here I suggest that two related paradigms exist at the broader spatial scale, namely the metapopulation paradigm and habitat paradigm, and that these two paradigms also need to be integrated if we are to provide sound management advice. This integration is not trivial, and I outline five problems that need to be addressed: (1) habitat variables may not measure habitat quality, so site-specific data on vital rates are needed to resolve the effects of habitat quality and metapopulation dynamics; (2) measurements of vital rates may be confounded by movements; (3) vital rates may be density dependent; (4) vital rates may be affected by genotype; and (5) vital rates cannot be measured in unoccupied patches. I reviewed papers published in Conservation Biology from 1994 to 2003 and found 41 studies that analyzed data from 10 or more sites to understand the factors limiting species' distributions. Five of the analyses presented were purely within the metapopulation paradigm, 14 were purely within the habitat paradigm, 17 involved elements of both paradigms, and 7 were theoretically ambiguous (2 papers presented 2 distinct analyses and were counted twice). This suggests that many researchers appreciate the need to integrate the paradigms. Only one study, however, used data on vital rates to resolve the effects of habitat quality and metapopulation dynamics (problem 1), and this study did not address problems 2–5. I conclude that more intensive research incorporating site-specific data on vital rates and movement is needed to complement the numerous analyses of distributional data being produced.