D. T. Tyler Flockhart

Tracking multi-generational colonization of the breeding grounds by monarch butterflies in eastern North America.

Insect migration may involve movements over multiple breeding generations at continental scales, resulting in formidable challenges to their conservation and management. Using distribution models generated from citizen scientist occurrence data and stable-carbon and -hydrogen isotope measurements, we tracked multi-generational colonization of the breeding grounds of monarch butterflies (Danaus plexippus) in eastern North America. We found that monarch breeding occurrence was best modelled with geographical and climatic variables resulting in an annual breeding distribution of greater than 12 million km2 that encompassed 99% occurrence probability. Combining occurrence models with stable isotope measurements to estimate natal origin, we show that butterflies which overwintered in Mexico came from a wide breeding distribution, including southern portions of the range. There was a clear northward progression of monarchs over successive generations from May until August when reproductive butterflies began to change direction and moved south. Fifth-generation individuals breeding in Texas in the late summer/autumn tended to originate from northern breeding areas rather than regions further south. Although the Midwest was the most productive area during the breeding season, monarchs that re-colonized the Midwest were produced largely in Texas, suggesting that conserving breeding habitat in the Midwest alone is insufficient to ensure long-term persistence of the monarch butterfly population in eastern North America.

Experimental evidence shows no fractionation of strontium isotopes (87Sr/86Sr) among soil, plants, and herbivores: implications for tracking wildlife and forensic science

Strontium isotopes (87Sr/86Sr) can be useful biological markers for a wide range of forensic science applications, including wildlife tracking. However, one of the main advantages of using 87Sr/86Sr values, that there is no fractionation from geological bedrock sources through the food web, also happens to be a critical assumption that has never been tested experimentally. We test this assumption by measuring 87Sr/86Sr values across three trophic levels in a controlled greenhouse experiment. Adult monarch butterflies were raised on obligate larval host milkweed plants that were, in turn, grown on seven different soil types collected across Canada. We found no significant differences between 87Sr/86Sr values in leachable Sr from soil minerals, organic soil, milkweed leaves, and monarch butterfly wings. Our results suggest that strontium isoscapes developed from 87Sr/86Sr values in bedrock or soil may serve as a reliable biological marker in forensic science for a range of taxa and across large geographic areas.

Experimental Examination of Intraspecific Density-Dependent Competition during the Breeding Period in Monarch Butterflies (Danaus plexippus)

A central goal of population ecology is to identify the factors that regulate population growth. Monarch butterflies (Danaus plexippus) in eastern North America re-colonize the breeding range over several generations that result in population densities that vary across space and time during the breeding season. We used laboratory experiments to measure the strength of density-dependent intraspecific competition on egg laying rate and larval survival and then applied our results to density estimates of wild monarch populations to model the strength of density dependence during the breeding season. Egg laying rates did not change with density but larvae at high densities were smaller, had lower survival, and weighed less as adults compared to lower densities. Using mean larval densities from field surveys resulted in conservative estimates of density-dependent population reduction that varied between breeding regions and different phases of the breeding season. Our results suggest the highest levels of population reduction due to density-dependent intraspecific competition occur early in the breeding season in the southern portion of the breeding range. However, we also found that the strength of density dependence could be almost five times higher depending on how many life-stages were used as part of field estimates. Our study is the first to link experimental results of a density-dependent reduction in vital rates to observed monarch densities in the wild and show that the effects of density dependent competition in monarchs varies across space and time, providing valuable information for developing robust, year-round population models in this migratory organism.

Patterns of parasitism in monarch butterflies during the breeding season in eastern North America

1. Migratory behaviour can result in reduced prevalence of pathogens in host populations. Two hypotheses have been proposed to explain this relationship: (i) ‘migratory escape’, where migrants benefit from escaping pathogen accumulation in contaminated environments; and (ii) ‘migratory culling’, where the selective removal of infected individuals occurs during migration.

Owned dog ecology and demography in Villa de Tezontepec, Hidalgo, Mexico.

Dog overpopulation in developing countries has negative implications for the health and safety of people, including the transmission of zoonotic diseases, physical attacks and intimidation to humans and animals, as well as impacts on canine welfare. Understanding the ecology and demographic characteristics of a dog population can help in the planning and monitoring of canine population control programs. Little data exist regarding demography and dynamics of domestic dog populations in semi-urban areas in Mexico. A cross-sectional study was carried out between October 21 and November 7, 2015, to characterize the dog ecology and demography in Villa de Tezontepec, Hidalgo, Mexico. A face-to-face survey was used to collect data from randomly selected households in four contiguous communities using stratified two-stage cluster sampling. Within each household, adults answered questions related to their dogs and their experiences with dog bites and aggression. A total of 328 households were interviewed, representing a participation rate of 90.9% (328/361) and 1,450 people. Approximately 65.2% of the households owned one or more dogs, with a mean of 1.3 (SD=1.5) and 2.0 (SD=1.5) owned dogs in all participant households and dog-owning households, respectively. The human: owned dog ratio for all participant households was 3.4:1 (1450/428), and for the dog-owning households was 2.3:1 (984/428). The owned dog male: female ratio was 1.4:1 (249/179). Approximately 74.4% (95.0% CI=69.8% - 78.7%) of the owned dogs were older than one year (mean age: 2.9 years; SD=2.5). The mean age of owned female dogs at first litter was 1.9 years (SD=1.2) and the mean litter size was 4.2 puppies (SD=2.1). Approximately 36.9% (95.0% CI=31.8% - 46.4%) of the females were spayed, and 14.1% (95.0% CI=10.7% - 19.7%) of the males were neutered. Only 44.9% (95.0% CI=40.1% - 49.7%) were always confined when unsupervised. Approximately 84.4% (95.0% CI=80.6% - 87.7%) were reported to have been vaccinated against rabies in 2015. The knowledge of owned dog demography and ecology provided by this study can inform local government planning of dog population control interventions, and could serve as a baseline for the development of agent-based models to evaluate the effects of different dog population control strategies on dog demography.

A general modeling framework for describing spatially structured population dynamics

Abstract Variation in movement across time and space fundamentally shapes the abundance and distribution of populations. Although a variety of approaches model structured population dynamics, they are limited to specific types of spatially structured populations and lack a unifying framework. Here, we propose a unified network‐based framework sufficiently novel in its flexibility to capture a wide variety of spatiotemporal processes including metapopulations and a range of migratory patterns. It can accommodate different kinds of age structures, forms of population growth, dispersal, nomadism and migration, and alternative life‐history strategies. Our objective was to link three general elements common to all spatially structured populations (space, time and movement) under a single mathematical framework. To do this, we adopt a network modeling approach. The spatial structure of a population is represented by a weighted and directed network. Each node and each edge has a set of attributes which vary through time. The dynamics of our network‐based population is modeled with discrete time steps. Using both theoretical and real‐world examples, we show how common elements recur across species with disparate movement strategies and how they can be combined under a unified mathematical framework. We illustrate how metapopulations, various migratory patterns, and nomadism can be represented with this modeling approach. We also apply our network‐based framework to four organisms spanning a wide range of life histories, movement patterns, and carrying capacities. General computer code to implement our framework is provided, which can be applied to almost any spatially structured population. This framework contributes to our theoretical understanding of population dynamics and has practical management applications, including understanding the impact of perturbations on population size, distribution, and movement patterns. By working within a common framework, there is less chance that comparative analyses are colored by model details rather than general principles.

Modeling the effect of surgical sterilization on owned dog population size in Villa de Tezontepec, Hidalgo, Mexico, using an individual-based computer simulation model

Surgical sterilization programs for dogs have been proposed as interventions to control dog population size. Models can be used to help identify the long-term impact of reproduction control interventions for dogs. The objective of this study was to determine the projected impact of surgical sterilization interventions on the owned dog population size in Villa de Tezontepec, Hidalgo, Mexico. A stochastic, individual-based simulation model was constructed and parameterized using a combination of empirical data collected on the demographics of owned dogs in Villa de Tezontepec and data available from the peer-reviewed literature. Model outcomes were assessed using a 20-year time horizon. The model was used to examine: the effect of surgical sterilization strategies focused on: 1) dogs of any age and sex, 2) female dogs of any age, 3) young dogs (i.e., not yet reached sexual maturity) of any sex, and 4) young, female dogs. Model outcomes suggested that as surgical capacity increases from 21 to 84 surgeries/month, (8.6% to 34.5% annual sterilization) for dogs of any age, the mean dog population size after 20 years was reduced between 14% and 79% compared to the base case scenario (i.e. in the absence of intervention). Surgical sterilization interventions focused only on young dogs of any sex yielded greater reductions (81% - 90%) in the mean population size, depending on the level of surgical capacity. More focused sterilization targeted at female dogs of any age, resulted in reductions that were similar to focusing on mixed sex sterilization of only young dogs (82% - 92%). The greatest mean reduction in population size (90% - 91%) was associated with sterilization of only young, female dogs. Our model suggests that targeting sterilization to young females could enhance the efficacy of existing surgical dog population control interventions in this location, without investing extra resources.

Multistate matrix population model to assess the contributions and impacts on population abundance of domestic cats in urban areas including owned cats, unowned cats, and cats in shelters

Concerns over cat homelessness, over-taxed animal shelters, public health risks, and environmental impacts has raised attention on urban-cat populations. To truly understand cat population dynamics, the collective population of owned cats, unowned cats, and cats in the shelter system must be considered simultaneously because each subpopulation contributes differently to the overall population of cats in a community (e.g., differences in neuter rates, differences in impacts on wildlife) and cats move among categories through human interventions (e.g., adoption, abandonment). To assess this complex socio-ecological system, we developed a multistate matrix model of cats in urban areas that include owned cats, unowned cats (free-roaming and feral), and cats that move through the shelter system. Our model requires three inputs—location, number of human dwellings, and urban area—to provide testable predictions of cat abundance for any city in North America. Model-predicted population size of unowned cats in seven Canadian cities were not significantly different than published estimates (p = 0.23). Model-predicted proportions of sterile feral cats did not match observed sterile cat proportions for six USA cities (p = 0.001). Using a case study from Guelph, Ontario, Canada, we compared model-predicted to empirical estimates of cat abundance in each subpopulation and used perturbation analysis to calculate relative sensitivity of vital rates to cat abundance to demonstrate how management or mismanagement in one portion of the population could have repercussions across all portions of the network. Our study provides a general framework to consider cat population abundance in urban areas and, with refinement that includes city-specific parameter estimates and modeling, could provide a better understanding of population dynamics of cats in our communities.

Effect of Capacity for Care on cat admission trends at the Guelph Humane Society, 2011–2015

ABSTRACT In recent years, there has been a growing concern regarding populations of cats who are homeless. Shelters are constantly overwhelmed by the influx of cats without caregivers and are seeking solutions to enhance positive outcomes for them. In 2014, the Guelph Humane Society implemented a population management program to expedite the movement of cats through the shelter by decreasing the average nonhuman animal’s length of stay using scheduled intakes to control for overcrowding and by implementing strategies to increase adoption rates. This study investigated the time trends in admission rates of cats to the Guelph Humane Society to assess the effectiveness of a population management program called Capacity for Care using a generalized linear autoregressive moving average model. From January 2011 to December 2015, a total of 3295 live cats were admitted to the Guelph Humane Society. When the analysis was adjusted to account for admissions of kittens, there was a significant reduction in admissions for adult cats (p < .01) following the introduction of the population management program. The results also showed a strong seasonal peak in total admissions during the summer months.

Migratory monarchs that encounter resident monarchs show life‐history differences and higher rates of parasite infection

Environmental change induces some wildlife populations to shift from migratory to resident behaviours. Newly formed resident populations could influence the health and behaviour of remaining migrants. We investigated migrant-resident interactions among monarch butterflies and consequences for life history and parasitism. Eastern North American monarchs migrate annually to Mexico, but some now breed year-round on exotic milkweed in the southern US and experience high infection prevalence of protozoan parasites. Using stable isotopes (δ2 H, δ13 C) and cardenolide profiles to estimate natal origins, we show that migrant and resident monarchs overlap during fall and spring migration. Migrants at sites with residents were 13 times more likely to have infections and three times more likely to be reproductive (outside normal breeding season) compared to other migrants. Exotic milkweed might either attract migrants that are already infected or reproductive, or alternatively, induce these states. Increased migrant-resident interactions could affect monarch parasitism, migratory success and long-term conservation.