Migratory connectivity of North American waterfowl across administrative flyways.

Management of waterfowl that migrate seasonally across North America occurs within four flyways that were delineated in the early 1900s to include the annual movements of populations. Movements may have changed over the past century since the administrative flyways were established, and may do so while management plans are in use, so information about transitions among flyways through time can illustrate how management assumptions may change. Today there are more than 12 million records from 60 years of migratory waterfowl band recoveries to assess adaptive management approaches that will be most effective when they account for movements within and between flyways. We examined how much the movement of North American waterfowl occurs between flyways, whether those movements have changed through time, and whether movements of mallards are representative of multiple species, as suggested by current harvest management strategies. We estimated the probability a duck would transition from one flyway to another and the strength of migratory connectivity (MC) for each species within and among flyways. We used capture-mark-recovery models to estimate population-specific movement within and among flyways (transition probabilities) for 15 migratory waterfowl species that were banded during breeding and recovered during winter. We developed new functionality in the R package MigConnectivity to estimate the species-specific strength of MC using transition probability samples from the capture-mark-recovery models. We found the regular movement of duck populations among flyways, overall weak MC, and no consistent change in migratory movements through time. Mallard movements were median among all duck species, but significantly different from many species, particularly diving ducks. Despite the significant movement between flyways, our work suggests flyway management of waterfowl matches many of the seasonal movements of these species when considering mid-continent flyway management. We recommend models accounting for all transition probabilities between populations and regularly estimating harvest derivations, transition probabilities, and MC metrics to verify that the current movements match model assumptions.

A hidden Markov model for estimating age-specific survival when age and size are uncertain.

Estimates of age-specific survival probabilities are needed for age-structured population models and to inform conservation decisions. However, determining the age of individuals in wildlife populations is often problematic. We present a hidden Markov model for estimating age-specific survival from capture-recapture or capture-recapture-recovery data when age is unknown and indicators of age, such as size and growth layer counts, are imprecise. The model is evaluated through simulations, and its implementation is illustrated with maximum likelihood and Bayesian approaches in commonly used software. The model is then applied to genetic capture-recapture data of Florida manatees to estimate age- and time-variant survival probabilities. The approach is broadly applicable to studies aiming to quantify age-specific effects of environmental change and management actions on population dynamics, including studies that rely on minimally invasive methods such as genetic and photo identification.

Monitoring abundance of aggregated animals (Florida manatees) using an unmanned aerial system (UAS).

Imperfect detection is an important problem when counting wildlife, but new technologies such as unmanned aerial systems (UAS) can help overcome this obstacle. We used data collected by a UAS and a Bayesian closed capture-mark-recapture model to estimate abundance and distribution while accounting for imperfect detection of aggregated Florida manatees (Trichechus manatus latirostris) at thermal refuges to assess use of current and new warmwater sources in winter. Our UAS hovered for 10 min and recorded 4 K video over sites in Collier County, FL. Open-source software was used to create recapture histories for 10- and 6-min time periods. Mean estimates of probability of detection for 1-min intervals at each canal varied by survey and ranged between 0.05 and 0.92. Overall, detection probability for sites varied between 0.62 and 1.00 across surveys and length of video (6 and 10 min). Abundance varied by survey and location, and estimates indicated that distribution changed over time, with use of the novel source of warmwater increasing over time. The highest cumulative estimate occurred in the coldest winter, 2018 (N = 158, CI 141-190). Methods here reduced survey costs, increased safety and obtained rigorous abundance estimates at aggregation sites previously too difficult to monitor.

Reconstructing population dynamics of a threatened marine mammal using multiple data sets.

Models of marine mammal population dynamics have been used extensively to predict abundance. A less common application of these models is to reconstruct historical population dynamics, filling in gaps in observation data by integrating information from multiple sources. We developed an integrated population model for the Florida manatee (Trichechus manatus latirostris) to reconstruct its population dynamics in the southwest region of the state over the past 20 years. Our model improved precision of key parameter estimates and permitted inference on poorly known parameters. Population growth was slow (averaging 1.02; 95% credible interval 1.01-1.03) but not steady, and an unusual mortality event in 2013 led to an estimated net loss of 332 (217-466) manatees. Our analyses showed that precise estimates of abundance could be derived from estimates of vital rates and a few input estimates of abundance, which may mean costly surveys to estimate abundance don't need to be conducted as frequently. Our study also shows that retrospective analyses can be useful to: (1) model the transient dynamics of age distribution; (2) assess and communicate the conservation status of wild populations; and (3) improve our understanding of environmental effects on population dynamics and thus enhance our ability to forecast.

Cold-related Florida manatee mortality in relation to air and water temperatures.

Many tropical and subtropical species are sensitive to sudden temperature changes, especially drops in temperature. During winters 2009-2010 and 2010-2011, unusually cold temperatures occurred in many parts of Florida, USA, resulting in increased mortality of Florida manatees, sea turtles, fish, corals, and other species. The Florida manatee, in particular, is highly susceptible to cold stress and death when water temperatures drop below 20°C. We sought to characterize the magnitude and timing of reports of cold-related manatee carcasses in relation to fluctuations in water and air temperatures in central-east and central-west Florida during the six winters from 2008 to 2014. We used a generalized linear model to predict counts of manatee carcasses with a cold-related cause of death reported over 7-day bins in relation to various short-term (two weeks or less) and cumulative (incrementally summed from the start of the winter) heating-degree-day effects (HDD; < 20°C) and a categorical winter variable. Using water temperature data, the top-ranked model in both regions included a short-term temperature effect (14-day HDD sum) that preceded increases in reports of cold-related manatee carcasses by 7 days. Cumulative exposure to cold weather over the winter amplified effects on mortality in the central-east region. Quantifying the relationship between cold events and manatee mortality helps us prepare for rescue and salvage operations when extremely cold weather is forecast. This is especially important because anticipated loss or degradation of warm-water refuges due to human activities and sea level rise could potentially impact the manatee population in the future. These methods could also be applied to other species susceptible to cold-related mortality.

Accuracy, precision, and error in age estimation of Florida manatees using growth layer groups in earbones.

Ages of Florida manatees (Trichechus manatus latirostris) can be estimated by counting annual growth layer groups (GLGs) in the periotic dome portion of the tympanoperiotic complex of their earbones. The Florida Fish and Wildlife Conservation Commission manages an archive of more than 8,700 Florida manatee earbones collected from salvaged carcasses from 1989 to 2017. Our goal was to comprehensively evaluate techniques used to estimate age, given this large sample size and changes to processing protocols and earbone readers over time. We developed new standards for estimating ages from earbones, involving two independent readers to obtain measurements of within- and between-reader precision. To quantify accuracy, precision, and error, 111 earbones from manatees with approximately known ages (first known as calves: "KAC") and 69 earbones from manatees with minimum known ages ("MKA," based on photo-identification sighting histories) were processed, and their ages were estimated. There was greater precision within readers (coefficient of variation, CV: 2.4-8.5%) than between readers (CV: 13.1-13.3%). The median of age estimates fell within the true age range for 63.1% of KAC cases and was at least the sighting duration for 75.0% of MKA cases. Age estimates were generally unbiased, as indicated by an average raw error ± SD of -0.05 ± 3.05 years for the KAC group. The absolute error (i.e., absolute value of raw error) of the KAC data set averaged 1.75 ± 2.50 years. Accuracy decreased and error increased with increasing known age, especially for animals over 15 years old, whose ages were mostly underestimated due to increasing levels of resorption (the process of bone turnover that obscures GLGs). Understanding the degree of uncertainty in age estimates will help us assess the utility of age data in manatee population models. We emphasize the importance of standardizing and routinely reviewing age estimation and processing protocols to ensure that age data remain consistent and reliable.

Temporal and demographic variation in partial migration of the North Atlantic right whale.

Animal movement plays a fundamental role in the ecology of migratory species, and understanding migration patterns is required for effective management. To evaluate intrinsic and environmental factors associated with probabilities of endangered North Atlantic right whales Eubalaena glacialis migrating to a wintering ground off the southeastern United States (SEUS), we applied a multistate temporary emigration capture-recapture model to 22 years of photo-identification data. Migration probabilities for juveniles were generally higher yet more variable than those for adults, and non-calving adult females were the least likely group to migrate. The highest migration probabilities for juveniles and adult males coincided with years of relatively high calving rates, following years of higher prey availability in a fall feeding ground. Right whale migration to the SEUS can be classified as condition-dependent partial migration, which includes skipped breeding partial migration for reproductive females, and is likely influenced by tradeoffs among ecological factors such as reproductive costs and foraging opportunities that vary across individuals and time. The high variability in migration reported in this study provides insight into the ecological drivers of migration but presents challenges to right whale monitoring and conservation strategies.

Spatial and temporal drivers of avian population dynamics across the annual cycle.

Untangling the spatial and temporal processes that influence population dynamics of migratory species is challenging, because changes in abundance are shaped by variation in vital rates across heterogeneous habitats and throughout the annual cycle. We developed a full-annual-cycle, integrated, population model and used demographic data collected between 2011 and 2014 in southern Indiana and Belize to estimate stage-specific vital rates of a declining migratory songbird, the Wood Thrush (Hylocichla mustelina). Our primary objective was to understand how spatial and temporal variation in demography contributes to local and regional population growth. Our full-annual-cycle model allowed us to estimate (1) age-specific, seasonal survival probabilities, including latent survival during both spring and autumn migration, and (2) how the relative contribution of vital rates to population growth differed among habitats. Wood Thrushes in our study populations experienced the lowest apparent survival rates during migration and apparent survival was lower during spring migration than during fall migration. Both mortality and high dispersal likely contributed to low apparent survival during spring migration. Population growth in high-quality habitat was most sensitive to variation in fecundity and apparent survival of juveniles during spring migration, whereas population growth in low-quality sites was most sensitive to adult apparent breeding-season survival. These results elucidate how full-annual-cycle vital rates, particularly apparent survival during migration, interact with spatial variation in habitat quality to influence population dynamics in migratory species.

Demography of population recovery: survival and fidelity of peregrine falcons at various stages of population recovery.

Factors influencing vital demographic rates and population dynamics can vary across phases of population growth. We studied factors influencing survival and fidelity of peregrine falcons in south Scotland-north England at two stages of population growth: when the population was recovering from pesticide-related declines and density was low, and when it had largely recovered from pesticide effects and density was high. Fidelity was higher for: adults and subadults than for juveniles, females than for males, and juveniles and adults during the low-density than during the high-density study period. Survival was age specific, with lower survival for juveniles than for older birds (juveniles, 0.600 ± SE 0.063; subadults, 0.811 ± 0.058; adults, 0.810 ± 0.034). Furthermore, there was some evidence that survival was generally lower for all age classes during the low-density period than during the high-density period, possibly due to a chronic, persistent effect of organochlorine pesticides as the population recovered. Evidence for a density-dependent effect on survival was weak, but a negative effect of density on fidelity of juveniles (dispersing age class) during the recovery phase suggests density-dependent dispersal when the population was increasing. Our results show how population density can influence demographic parameters differently and how such influences can vary across phases of population growth.

Estimating migratory connectivity of birds when re-encounter probabilities are heterogeneous.

Understanding the biology and conducting effective conservation of migratory species requires an understanding of migratory connectivity - the geographic linkages of populations between stages of the annual cycle. Unfortunately, for most species, we are lacking such information. The North American Bird Banding Laboratory (BBL) houses an extensive database of marking, recaptures and recoveries, and such data could provide migratory connectivity information for many species. To date, however, few species have been analyzed for migratory connectivity largely because heterogeneous re-encounter probabilities make interpretation problematic. We accounted for regional variation in re-encounter probabilities by borrowing information across species and by using effort covariates on recapture and recovery probabilities in a multistate capture-recapture and recovery model. The effort covariates were derived from recaptures and recoveries of species within the same regions. We estimated the migratory connectivity for three tern species breeding in North America and over-wintering in the tropics, common (Sterna hirundo), roseate (Sterna dougallii), and Caspian terns (Hydroprogne caspia). For western breeding terns, model-derived estimates of migratory connectivity differed considerably from those derived directly from the proportions of re-encounters. Conversely, for eastern breeding terns, estimates were merely refined by the inclusion of re-encounter probabilities. In general, eastern breeding terns were strongly connected to eastern South America, and western breeding terns were strongly linked to the more western parts of the nonbreeding range under both models. Through simulation, we found this approach is likely useful for many species in the BBL database, although precision improved with higher re-encounter probabilities and stronger migratory connectivity. We describe an approach to deal with the inherent biases in BBL banding and re-encounter data to demonstrate that this large dataset is a valuable source of information about the migratory connectivity of the birds of North America.

A cat's tale: the impact of genetic restoration on Florida panther population dynamics and persistence.

1. Genetic restoration has been suggested as a management tool for mitigating detrimental effects of inbreeding depression in small, inbred populations, but the demographic mechanisms underlying population-level responses to genetic restoration remain poorly understood. 2. We studied the dynamics and persistence of the endangered Florida panther Puma concolor coryi population and evaluated the potential influence of genetic restoration on population growth and persistence parameters. As part of the genetic restoration programme, eight female Texas pumas P. c. stanleyana were released into Florida panther habitat in southern Florida in 1995. 3. The overall asymptotic population growth rate (λ) was 1.04 (5th and 95th percentiles: 0.95-1.14), suggesting an increase in the panther population of approximately 4% per year. Considering the effects of environmental and demographic stochasticities and density-dependence, the probability that the population will fall below 10 panthers within 100 years was 0.072 (0-0.606). 4. Our results suggest that the population would have declined at 5% per year (λ = 0.95; 0.83-1.08) in the absence of genetic restoration. Retrospective life table response experiment analysis revealed that the positive effect of genetic restoration on survival of kittens was primarily responsible for the substantial growth of the panther population that would otherwise have been declining. 5. For comparative purposes, we also estimated probability of quasi-extinction under two scenarios - implementation of genetic restoration and no genetic restoration initiative - using the estimated abundance of panthers in 1995, the year genetic restoration was initiated. Assuming no density-dependence, the probability that the panther population would fall below 10 panthers by 2010 was 0.098 (0.002-0.332) for the restoration scenario and 0.445 (0.032-0.944) for the no restoration scenario, providing further evidence that the panther population would have faced a substantially higher risk of extinction if the genetic restoration initiative had not been implemented. 6. Our results, along with those reporting increases in population size and improvements in biomedical correlates of inbreeding depression, provide strong evidence that genetic restoration substantially contributed to the observed increases in the Florida panther population.

Stochastic population dynamics of a montane ground-dwelling squirrel.

Understanding the causes and consequences of population fluctuations is a central goal of ecology. We used demographic data from a long-term (1990-2008) study and matrix population models to investigate factors and processes influencing the dynamics and persistence of a golden-mantled ground squirrel (Callospermophilus lateralis) population, inhabiting a dynamic subalpine habitat in Colorado, USA. The overall deterministic population growth rate λ was 0.94±SE 0.05 but it varied widely over time, ranging from 0.45±0.09 in 2006 to 1.50±0.12 in 2003, and was below replacement (λ<1) for 9 out of 18 years. The stochastic population growth rate λ(s) was 0.92, suggesting a declining population; however, the 95% CI on λ(s) included 1.0 (0.52-1.60). Stochastic elasticity analysis showed that survival of adult females, followed by survival of juvenile females and litter size, were potentially the most influential vital rates; analysis of life table response experiments revealed that the same three life history variables made the largest contributions to year-to year changes in λ. Population viability analysis revealed that, when the influences of density dependence and immigration were not considered, the population had a high (close to 1.0 in 50 years) probability of extinction. However, probability of extinction declined to as low as zero when density dependence and immigration were considered. Destabilizing effects of stochastic forces were counteracted by regulating effects of density dependence and rescue effects of immigration, which allowed our study population to bounce back from low densities and prevented extinction. These results suggest that dynamics and persistence of our study population are determined synergistically by density-dependence, stochastic forces, and immigration.

Does genetic introgression improve female reproductive performance? A test on the endangered Florida panther.

Genetic introgression has been suggested as a management tool for mitigating detrimental effects of inbreeding depression, but the role of introgression in species conservation has been controversial, partly because population-level impacts of genetic introgressions are not well understood. Concerns about potential inbreeding depression in the endangered Florida panther (Puma concolor coryi) led to the release of eight female Texas pumas (P. c. stanleyana) into the Florida panther population in 1995. We used long-term reproductive data (1995-2008) collected from 61 female Florida panthers to estimate and model reproduction probability (probability of producing a litter) and litter size, and to investigate the influence of intentional genetic introgression on these parameters. Overall, 6-month probability of reproduction (±1SE) was 0.232 ± 0.021 and average litter size was 2.60 ± 0.09. Although F(1) admixed females had a lower reproduction probability than females with other ancestries, this was most likely because kittens born to F(1) females survive better; consequently, these females are unavailable for breeding until kittens are independent. There was no evidence for the effect of ancestry on litter size or of heterozygosity on probability of reproduction or litter size. In contrast, earlier studies have shown that genetic introgression positively affected Florida panther survival. Our results, along with those of earlier studies, clearly suggest that genetic introgression can have differential effects on components of fitness and highlight the importance of examining multiple demographic parameters when evaluating the effects of management actions.

Synergistic influences of phase, density, and climatic variation on the dynamics of fluctuating populations.

Although ecologists have long recognized that certain mammalian species exhibit high-amplitude, often multiannual, fluctuations in abundance, their causes have remained poorly understood and the subject of intense debate. A key contention has been the relative role of density-dependent and density-independent processes in governing population dynamics. We applied capture-mark-recapture analysis to 25 years of monthly trapping data from a fluctuating prairie vole Microtus ochrogaster population in Illinois, USA, to estimate realized population growth rates and associated vital rates (survival and recruitment) and modeled them as a function of vole density and density-independent climatic variation. We also tested for phase dependence and seasonality in the effects of the above processes. Variation in the realized population growth rate was best explained by phase-specific changes in vole density lagged by one month and mean monthly temperatures with no time lags. The underlying vital rates, survival and recruitment, were influenced by the additive and interactive effects of phase, vole density, and mean monthly temperatures. Our results are consistent with the observation that large-scale population fluctuations are characterized by phase-specific changes in demographic and physiological characteristics. Our findings also support the growing realization that the interaction between climatic variables and density-dependent factors may be a widespread phenomenon, and they suggest that the direction and magnitude of such interactive effects may be phase specific. We conclude that density-dependent and density-independent climatic variables work in tandem during each phase of density fluctuations to drive the dynamics of fluctuating populations.

Effects of prescribed fire, supplemental feeding, and mammalian predator exclusion on hispid cotton rat populations.

Predation and food resources can strongly affect small mammal population dynamics directly by altering vital rates or indirectly by influencing behaviors. Fire may also strongly influence population dynamics of species inhabiting fire-adapted habitats because fire can alter food and cover availability. We used capture-mark-recapture and radio-telemetry studies to experimentally examine how supplemental feeding, mammalian predator exclusion, and prescribed fire affected survival, abundance, and reproduction of hispid cotton rats (Sigmodon hispidus) in southwestern Georgia, USA. Prescribed fire reduced survival, abundance, and rates of transitions to reproductive states. Food supplementation increased survival, transitions to reproductive states, and abundance, but was not sufficient to prevent post-fire declines in any of these parameters. Mammalian predator exclusion did not strongly affect any of the considered parameters. Our results show that fire strongly influenced cotton rat populations in our study site, primarily by reducing cover and increasing predation risk from non-mammalian predators.

Intentional genetic introgression influences survival of adults and subadults in a small, inbred felid population.

1. Inbreeding and low genetic diversity can cause reductions in individual fitness and increase extinction risk in animal populations. Intentional introgression, achieved by releasing genetically diverse individuals into inbred populations, has been used as a conservation tool to improve demographic performance in endangered populations. 2. By the 1980s, Florida panthers (Puma concolor coryi) had been reduced to a small, inbred population that appeared to be on the brink of extinction. In 1995, female pumas from Texas (P. c. stanleyana) were released in occupied panther range as part of an intentional introgression programme to restore genetic variability and improve demographic performance of panthers. 3. We used 25 years (1981-2006) of continuous radiotelemetry and genetic data to estimate and model subadult and adult panther survival and cause-specific mortality to provide rigorous sex and age class-specific survival estimates and evaluate the effect of the introgression programme on these parameters. 4. Genetic ancestry influenced annual survival of subadults and adults after introgression, as F(1) generation admixed panthers ( = 0·98) survived better than pre-introgression type panthers ( = 0·77) and other admixed individuals ( = 0·82). Furthermore, heterozygosity was higher for admixed panthers relative to pre-introgression type panthers and positively influenced survival. 5. Our results are consistent with hybrid vigour; however, extrinsic factors such as low density of males in some areas of panther range may also have contributed to higher survival of F(1) panthers. Regardless, improved survival of F(1) subadults and adults likely contributed to the numerical increase in panthers following introgression, and our results indicate that intentional admixture, achieved here by releasing individuals from another population, appears to have been successful in improving demographic performance in this highly endangered population.

Genetic Introgression and the Survival of Florida Panther Kittens.

Estimates of survival for the young of a species are critical for population models. These models can often be improved by determining the effects of management actions and population abundance on this demographic parameter. We used multiple sources of data collected during 1982-2008 and a live recapture-dead recovery modeling framework to estimate and model survival of Florida panther (Puma concolor coryi) kittens (age 0 - 1 year). Overall, annual survival of Florida panther kittens was 0.323 ± 0.071 (SE), which was lower than estimates used in previous population models. In 1995, female pumas from Texas (P. c. stanleyana) were released into occupied panther range as part of an intentional introgression program to restore genetic variability. We found that kitten survival generally increased with degree of admixture: F(1) admixed and backcrossed to Texas kittens survived better than canonical Florida panther and backcrossed to canonical kittens. Average heterozygosity positively influenced kitten and older panther survival, whereas index of panther abundance negatively influenced kitten survival. Our results provide strong evidence for the positive population-level impact of genetic introgression on Florida panthers. Our approach to integrate data from multiple sources was effective at improving robustness as well as precision of estimates of Florida panther kitten survival, and can be useful in estimating vital rates for other elusive species with sparse data.

Genetic restoration of the Florida panther.

The rediscovery of remnant Florida panthers (Puma concolor coryi) in southern Florida swamplands prompted a program to protect and stabilize the population. In 1995, conservation managers translocated eight female pumas (P. c. stanleyana) from Texas to increase depleted genetic diversity, improve population numbers, and reverse indications of inbreeding depression. We have assessed the demographic, population-genetic, and biomedical consequences of this restoration experiment and show that panther numbers increased threefold, genetic heterozygosity doubled, survival and fitness measures improved, and inbreeding correlates declined significantly. Although these results are encouraging, continued habitat loss, persistent inbreeding, infectious agents, and possible habitat saturation pose new dilemmas. This intensive management program illustrates the challenges of maintaining populations of large predators worldwide.

Determinants of partial bird migration in the Amazon Basin.

1. Little is known about mechanisms that drive migration of birds at tropical latitudes. Because most migratory bird species in South America have populations that are present year-round, partial migration (in which only some individuals of a given population migrate at the end of the breeding season) is likely to be common, providing an opportunity to assess proximate mechanisms of migration. 2. Two non-mutually exclusive hypotheses explaining intraspecific variation in migratory behaviour were tested in a Tropical Kingbird Tyrannus melancholicus population in the southern Amazon Basin, where a dramatic dry season decrease in the abundance of insect food for kingbirds may promote migration of some individuals. 3. The Dominance hypothesis predicts sub-dominant individuals migrate at the end of the breeding season and dominant individuals do not, whereas the Body Size hypothesis predicts smaller individuals migrate and larger individuals do not. 4. Based on 4 years of data on individually-marked birds, strong support was found for occurrence of partial migration in the study population. 5. In the best model, the largest males (which are typically older and dominant to younger individuals) had the highest probability of migrating. Younger females (which are the smallest individuals in the population) were also more likely to migrate than other kingbirds, except the largest males. Thus, an individual's probability of migrating was associated with a more complex interaction of size, age and sex than predicted by current hypotheses. 6. These results suggest that determinants of migratory behaviour differ between North temperate and tropical latitudes. Most tests of partial migration theory have been conducted on granivores (e.g. emberizids) or omnivores (e.g. turdids and icterids) at North temperate latitudes, where seasonality is primarily defined by temperature cycles. In tropical South America, however, the most common long-distance migrants are primarily insectivorous (i.e. tyrannids), and seasonality is largely defined by rainfall cycles. 7. We propose a hypothesis based on seasonal food limitation to explain partial migration of primarily insectivorous species in seasonal tropical habitats.

Benefits for plants in ant-plant protective mutualisms: a meta-analysis.

Costs and benefits for partners in mutualistic interactions can vary greatly, but surprisingly little is known about the factors that drive this variation across systems. We conducted a meta-analysis of ant-plant protective mutualisms to quantify the effects of ant defenders on plant reproductive output, to evaluate if reproductive effects were predicted from reductions in herbivory and to identify characteristics of the plants, ants and environment that explained variation in ant protection. We also compared our approach with two other recent meta-analyses on ant-plant mutualisms, emphasizing differences in our methodology (using a weighted linear mixed effects model) and our focus on plant reproduction rather than herbivore damage. Based on 59 ant and plant species pairs, ant presence increased plant reproductive output by 49% and reduced herbivory by 62%. The effects on herbivory and reproduction within systems were positively correlated, but the slope of this relationship (0.75) indicated that tolerance to foliar herbivory may be a common plant response to absence of ant guards. Furthermore, the relationship between foliar damage and reproduction varied substantially among systems, suggesting that herbivore damage is not a reliable surrogate for fitness consequences of ant protection. Studies that experimentally excluded ants reported a smaller effect of ant protection on plant reproduction than studies that relied upon natural variation in ant presence, suggesting that study methods can affect results in these systems. Of the ecological variables included in our analysis, only plant life history (i.e., annual or perennial) explained variation in the protective benefit of mutualistic ants: presence of ants benefitted reproduction of perennials significantly more than that of annuals. These results contrast with other quantitative reviews of these relationships that did not include plant life history as an explanatory factor and raise several questions to guide future research on ant-plant protection mutualisms.