Report on Population Viability Analysis Model Investigations of Threats to the Southern Resident Killer Whale Population from Trans Mountain Expansion Project

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Report on Population Viability Analysis model investigations of threats to the Southern Resident Killer Whale population from Trans Mountain Expansion Project

Prepared for: Raincoast Conservation Foundation

Prepared by:

Robert C. Lacy, PhD Senior Conservation Scientist Chicago Zoological Society

Kenneth C. Balcomb III Executive Director and Senior Scientist Center for Whale Research

Lauren J.N. Brent, PhD Associate Research Fellow, Center for Excellence in Animal Behaviour, University of Exeter Postdoctoral Associate, Centre for Cognitive Neuroscience, Duke University

Darren P. Croft, PhD Associate Professor of Animal Behaviour University of Exeter

Christopher W. Clark, PhD Imogene P. Johnson Senior Scientist, Bioacoustics Research Program, Cornell Lab of Ornithology Senior Scientist, Department of Neurobiology and Behavior, Cornell University

Paul C. Paquet Senior Scientist Raincoast Conservation Foundation

Table of Contents

Page 1.0. Introduction 1

2.0. PVA Methodology 3

2.1. The Population Viability Analysis (PVA) modeling approach 6

2.2. About the Vortex PVA software 6

2.3. Input variables used in developing the Vortex PVA model for the Southern Resident population 13

3.0 Results of modelling 13

3.1. Projections for the Southern Resident population under the status quo (baseline model) 17

3.2. Sensitivity testing of important model parameters 18

3.3. Examination of potential threats 18

4.0. Conclusions 36

5.0. Literature Cited 38

Appendix A Review of Previous Southern Resident Population Viability Analyses

Appendix B Curriculum vitae Dr. Robert C. Lacy, Ph.D.

Appendix B Biography Kenneth C. Balcomb III

Appendix C Curriculum vitae Laruen J.N. Brent, Ph.D.

Appendix D Curriculum vitae Darren P. Croft, Ph.D.

Appendix E Curriculum vitae Christopher W. Clark, Ph.D.

Appendix F Curriculum vitae Paul Paquet, Ph.D.

1.0. Introduction

This report explains the methodology and results of a population viability analysis (PVA) of the risk associated with aspects of the proposed Trans Mountain Expansion Project (Project) on the endangered Southern Resident Killer Whales (“Southern Residents”). PVA uses quantitative methods to evaluate and predict the likely future status of a population. The Project will twin an existing pipeline from Alberta to Burnaby BC on the Pacific Coast. That pipeline will carry diluted bitumen from Alberta to Burnaby, British Columbia, to be shipped overseas. The Project will therefore result in the increased shipping of diluted bitumen by oil tanker through the shipping lane in the Salish Sea. This shipping lane transects the critical habitat of the Southern Residents.

A PVA is a risk management tool, which models the risk to a population over time. In this case the model was used to estimate the increased risk to the Southern Residents from three threats associated with the marine shipping component of the Project: an oil spill, increased acoustic and physical disturbance from ships, and ship strikes.

Projecting the impacts of changing and even new threats to a wildlife population can be difficult. Intuition, even by acknowledged experts, is usually not adequate because the factors influencing wildlife populations are numerous and interact in complex ways; new threats with which we have little prior experience are emerging; and even if the intuitive assessments are complete and correct, they cannot be documented in a way that allows understanding, review, and improvement by others. Standard demographic models used in wildlife ecology can be powerful methods for predicting population dynamics, although they often assume no or simple interactions among variables and necessarily omit much detail of the breeding system, spatial or social structure, and individual or group differences. Much uncertainty exists in any population projection, because any model is necessarily (and by design) a simplified representation of the system, some input variables will be known only approximately, future conditions cannot be known precisely, and some biological processes are inherently unpredictable (e.g., the sex of each newborn). This uncertainty cannot be eliminated, and it should not be ignored, but it can be explored through testing perturbations of the data and model (“sensitivity testing”) to determine which uncertain factors have the greatest influence on our projections for the future and what confidence we can place on the results. Overall, population models provide a framework not only for analyzing complex situations, but also for documenting assumptions and methods, reviewing and improving assessments, and integrating new potential threats to populations into our understanding.

We prepared this report using a PVA modeling approach. A PVA modelling approach uses demographic modeling to assess risks to wildlife populations and evaluate the likely efficacy of protection, recovery, or restoration options (Shaffer 1990; Boyce 1992; Burgman et al. 1993; Sjögren-Gulve and Ebenhard 2000; Beissinger and McCullough 2002; Morris and Doak 2002). Our goal was to evaluate the current population status of Southern Residents and future population dynamics, including the probability of recovery and risk of extinction under existing and future human caused threats. We assessed the likely response of population growth and viability to additional cumulative disturbances and threats, including increased ocean noise resulting from additional vessel traffic and oil spills from petroleum tankers

associated with the proposed Project.

Our baseline or status quo model, which reflects current population demographics, environmental conditions, and human caused threats, projects a mean population decline of 0.2% annually, with fluctuation of approximately + 4.5% across 100 years, and no complete extinctions (i.e. either no males or no females remaining). There was however, a 9% chance of quasi-extinction1, where the population falls below 30 whales and is no longer viable. A 7% reduction in genetic diversity after 100 years is also projected. Birth rate (calves per adult female) dominated the variation in population growth, whereas, nonbreeding depression and mortality rates had moderate effects. The effects of inbreeding depression indicate that the population is small enough so that inbreeding is likely damaging to the future prospects for the population.

Reducing the abundance of Chinook salmon is projected to have a substantial negative impact on the Southern Residents population. Conversely, increasing Chinook abundance can lead to relatively robust population growth (up to 1.9% with 20% more prey) and protection from extinction or serious decline. The impact of oil spills on long-term population growth could be consequential. The probability of the population dropping to quasi-extinction (i.e. below 30 whales) due to an oil spill is more substantial, with a possibility of complete extinction within 100 years due to the potential of catastrophic decimation of the population to a small and unrecoverable number from which it could not recover. The modeled impact of noise and physical disturbance accompanying increased or added vessel traffic associated with the Project resulted in accelerated population decline, smaller mean population size, and increased probability of complete and quasi extinction. The cumulative impact of oil spills, increased noise, and incidental human caused deaths resulted in a population decline of more than 1% per year, resulting in a mean final population size of only 33 animals, a 8.6% probability of complete extinction of the Southern Residents, and more than 50% probability that the final population would be less than 30 animals and, therefore, probably on a course toward nearly inevitable extinction.

The Southern Resident population has experienced almost no population growth over the past 4 decades, and has declined in the last two decades. Based on status quo conditions, the population will most likely remain about at its current size or continue a very slow decline. It is abundantly clear that under status quo conditions the population cannot withstand additional negative pressures, recover from its current endangered status, and persist.

Our analysis shows that the Project will intensify existing threats, accelerating the rate of decline in the Southern Residents and possibly leading to a complete extinction. The factor projected to have the largest effect on depressing population size and possibly leading to extinction is a reduction of the Chinook prey base. In addition, occasional large or medium oil spills have the potential to depress long-

1 Quasi-extinction threshold is the minimum number below which extinction is very likely due to genetic or demographic risks. For this review, the quasi-extinction threshold is 30 killer whales, a level at which the Southern Residents are unlikely to persist.

term population stability and growth, leading to a considerable chance of the population declining below 30 animals. Cumulatively, the threats associated with the Project such as increased shipping of oil through critical habitat, the possibility of oil spills at the frequency estimated by Kinder Morgan, projected effects of noise from associated shipping traffic, and occasional additional mortalities due to boat strikes (one in 10 years) create the risk that the Southern Residents population becomes vulnerable to extinction (8.6% probability within 100 years), and increases the risk to more than 50% probability that the population will decline below 30 animals.

2.0. PVA Methodology

2.1. The Population Viability Analysis (PVA) modeling approach

PVA can extend standard demographic projections (Ricklefs 1979; Caswell 2001) in two important ways: (1) the impacts of forces external to the population (e.g., changing habitat quality, extent, and configuration; interactions with other species in the community; impacts of disease or contaminants; harvest, incidental killing, or other direct human impacts) on the demographic rates are explicitly considered and evaluated, and (2) uncertainty in the population trajectory caused by intrinsic (e.g., demographic stochasticity,2 limitations in local mate availability or other density dependent feedbacks, inbreeding impacts) and extrinsic (e.g., environmental variation, occasional catastrophes) stochastic factors can be explicitly modeled, usually through the use of simulation modeling.

The outputs of PVA are any desired measure of population performance, but commonly assessed metrics include projected mean (i.e. average) population size (N) over time, population growth rates (r), expected annual fluctuations in both N and r, probability of population extinction, and probabilities of quasi-extinction (the likelihood of N falling below any specified number). These outputs are used to assess risk (e.g., for listing under Canada’s Species at Risk Act and the U.S. Endangered Species Act or other protective regulations), assess vulnerability to possible threats, determine sustainable mortalities in the context of uncertainty, and determine the suites of actions that would be needed to achieve stated resource protection or restoration goals. PVA has become a cornerstone of conservation science and resource management.

Most PVA models can be classified into three general types that span top-down to bottom-up approaches:

(1) Deterministic (analytical) models calculate average population trajectories based on equations describing population growth, usually evaluated from matrix representations of the “life table” of fecundity and survival rates.

(2) Population-based simulation models add temporal and spatial variability to the demographic rates and then project a distribution of possible population trajectories based on simulations that sample

2 Demographic stochasticity is the random change in population size over time due to random variation in individual survival and reproductive success. 3

from those distributions. Thus, they use simulation of stochastic (probabilistic) population processes to generate population trends as emergent properties.

(3) Individual-based (also termed “agent-based”) simulation models use the probabilities of demographic events (including temporal, spatial, and individual variability) to simulate the life cycle of each individual and then project population level outcomes as the aggregate fates of the individuals in the population.

Any of these three kinds of PVA can be used to integrate impacts on individual animals (with respect to mortality, reproduction, and dispersal) into projections of the cumulative consequences at the level of the population, if the effects on individuals can be described adequately in the parameters at the level used in the model. However, deterministic analytical and population-based simulation approaches require that impacts on individuals be accurately captured in consequent changes to a few population- level rates, while individual-based simulations let the population-level effects emerge from more detailed descriptions of individual processes.

Deterministic models are most useful for describing dynamics of generalized populations with the typical species characteristics. Their output is usually in the form of calculated exponential rates of population growth and the stable age structure that will develop if all demographic rates remain constant. Because deterministic models assume constancy of demographic rates, they will over-estimate population growth and stability if rates fluctuate over time or space, if there is individual variation in rates, or if population sizes are small (on the order of 100s rather than 1000s).

Population-based and individual-based models provide the same mean outputs as do deterministic models (usually with lower estimates, due to the non-linear effects of fluctuations), but also add estimates of variation in population trajectories. To obtain these estimates of variation, the simulation models are repeated many times (500 iterations are usually considered sufficient to provide acceptably precise results) to generate a distribution of results, thereby allowing calculation of the probability that an outcome (a performance measure) will fall outside of an acceptable range.

Population-based simulations are most useful when individual characteristics and impacts can be averaged across the population without loss of ability to model key processes of interest, and populations are large enough so that chance variation among individuals is not a significant contributor to uncertainty.

Individual-based models are most useful for assessment of populations with a relatively small number of individuals in a specific and well-described environmental and management setting. Accordingly, the Vortex PVA model that we used is an individual-based model, with the fate of each animal tracked through its lifetime. In contrast to “population-based” models or analytical calculations of average population growth, individual-based models allow for determination of matrilines, individual variation in reproductive success, loss of genetic diversity, inbreeding, inbreeding avoidance, and any demographic consequences of these processes.

It is important to note that deterministic models, population-based models, and individual-based models all yield the same population predictions when the assumptions of the higher level models are met. Thus, the three kinds of models sequentially add accuracy, ability to capture more details of dynamic processes, and consideration of more sources of uncertainty, and they often represent the same demographic rates in different terms, but they are not generally conflicting approaches. However, some measures of viability cannot be assessed from the simpler deterministic or population-based models, because these measures describe specific individual or group statistics (such as the number of pods with more than two breeding adults) that are not represented at the higher population level. For example, deterministic models and population-based models almost always omit consideration of males, because reproduction in a polygamous species only requires sufficient availability of adult males to inseminate all potentially breeding females, and the models simplistically assume that there will always be enough males to ensure maximal reproduction. However, in reality, if social groups are so small that there is a lack of suitable males for breeding, the population size will decrease and perhaps crash.

A requirement for any PVA model to provide sufficiently accurate and robust projections to allow estimation of population performance (rather than simply to provide a modeling tool that allows exploration of a wide range of uncertainty) is the availability of detailed demographic data. Model input is required from the focal population or comparable reference populations for mortality rates, aspects of reproduction (e.g., age of breeding, inter-birth intervals, infant survival, or composite measures such as overall fecundity), population size, and habitat carrying capacity3 – as well as the natural fluctuations in these rates. The difficulty in obtaining sufficient demographic data on endangered or protected species is a common challenge to the usefulness of PVA models, and many practitioners consequently recommend that PVA models be used only to provide assessments of relative risk and relative value of management options, rather than absolute measures of population trajectories. In the case of the Southern Residents population, however, nearly complete demographic data are available from 1976 onwards. This allows highly detailed and situation-specific models to be developed and used for threat assessments. A review of previous PVAs carried out for Southern Resident killer whales is provided in Appendix A of this report.

2.2. About the Vortex PVA software

This report was prepared using the Vortex population model and software. The Vortex population model (Lacy 2000; Lacy et al. 2014a; Lacy et al. 2014b) is used by thousands of population biologists around the world, including independent researchers, government agencies, and teachers in classroom settings. Typical applications include projecting trends, threats, and restoration strategies for wildlife populations. For example, the United States Fish and Wildlife Service and other agencies have used

3 Carrying capacity is the maximum population size of the species that the environment can sustain indefinitely, given the food, habitat, water, and other life requisites available in the environment.

Vortex to set recovery goals and guide actions for a number of threatened and endangered species.4 No software program (nor any calculations) can be guaranteed to be completely free of errors, but as a widely distributed and used program, Vortex receives far more testing than similar models that are developed for only specific projects. Vortex simulations have been confirmed to produce population trajectories that are consistent with monitored wildlife populations (Brook et al. 2000a) and with other population models (Brook et al. 2000b). The overall structure of the program was published in Lacy (2000), and the compiled program and documentation are distributed freely at www.vortex10.org/Vortex10.aspx.

2.3. Input variables used in developing the Vortex PVA model for the Southern Resident population

In this section, we outline how the PVA modeling was done, including selection of the input variables. We explain the simulation settings (number of years modeled, number of times the simulation was repeated) and the tests that were run to determine the sensitivity of the model projections to the uncertainty in the input variables.

We conducted the PVA modeling of the Southern Residents population in three parts:

• First, a “baseline” model was developed to represent the best possible trajectories of the population if demographic rates continue as observed in recent decades, and no additional external threats such as oil spills, other environmental contamination, noise disturbance, decreasing food supply, increased disease, or increased ship strikes affect the population.

• Second, uncertain model parameter values were systematically altered to determine which would have large effects on the population projections. This “sensitivity analysis” helps determine the confidence we can have in projections given the necessarily incomplete knowledge of the population characteristics. It also reveals to which demographic rates the population performance would be most sensitive if subject to new or worsening threats.

4 Black-footed ferret, Mexican wolf (Carroll et al. 2014), Florida panther (Maehr et al. 2002), whooping crane (Mirande et al. 1991), Attwater’s prairie chicken, Puerto Rican crested toad, red wolf (Kelly et al. 1999), Anacapa Island deer mouse (Pergams et al. 2000), Alabama beach mouse (Reed and Traylor- Holzer 2006), Sonoran pronghorn (Hosack et al. 2002), Assateague Island pony (Ballou et al. 2008), Lower Keys marsh rabbit (Forys and Humphrey 1999), Hawaiian forest birds (Ellis et al. 1992), and many other species. Vortex is also a common tool for population analysis by governmental agencies in Canada, Mexico, Costa Rica, Panama, Brazil, Chile, Ecuador, New Zealand, Australia, Indonesia, Malaysia, China, Taiwan, Japan, India, Bangladesh, South Korea, Kenya, South Africa, Spain, Sweden, Norway, Denmark, Italy, UAE, and elsewhere. Vortex has been used before to assess threats to the Southern Resident Killer Whale population (Taylor and Plater 2001), and other species of marine mammals, such as Indo-Pacific bottlenose dolphins (Lacy and Wells 2009), manatee (Marmontel et al. 1997; Castelblanco-Martinez et al. 2012), dugongs (Heinsohn et al. 2004), and Steller’s sea cow (Turvey and Risley 2006).

• Third, estimated impacts of intensifying specified threats (Project related) to the population were added to the model to determine which threats individually could put the population at significant further risk, and what would be the risk of population decline or collapse under the cumulative effects of the several identified new threats.

Input variable 1: Primary data source for baseline demographic rates Demographic rates were calculated from the database on individual animal histories compiled by the Center for Whale Research (CWR 2015). Data from 1976 through 2014 were used. It is recognized that the Southern Resident population experienced moderate population growth from 1976 to 1993, and subsequently experienced a moderate decline. This might indicate that some aspect(s) of the habitat declined in quality recently, or the change in trajectory might be explained by random fluctuations in reproduction and survival, rather than indicating any long-term trend. In addition, the population might have peaked when it reached an ecological carrying capacity – the number of animals that can be supported by the available resource base. By including data back to 1976 for calculation of demographic rates, therefore, we could be over-estimating current conditions. However, to restrict analyses to only the more recent years could under-estimate the ability of the population to grow.

Because the sample sizes are necessarily small, the data are not sufficient to determine accurate estimates of demographic rates for each annual age class. We therefore used the same age class groupings as were used in the recent models of Vélez-Espino et al. (2014a), except that we set an upper limit for female breeding at 45 years rather than 50 years. Thus, we calculated survival and (for adult females) fecundity rates for calves (first year), juveniles (defined as from 1 year through 9 years of age), young mature females (10-30 years), older reproductive females (31-45 years), post-reproductive females (46 years and older), young mature males (10-21 year), and older males (22 years and older). Not all Southern Residents calves and juveniles have been sexed, and the data do not show a significant difference in survival between males and females in these younger age classes. Accordingly, survival was calculated with both sexes combined for calves and for juveniles. Survival rates of young adult females and young adult males have been better than older females and males, so it is appropriate to consider these age classes separately in the analyses. We set an upper limit of 45 years for females breeding because no females in the population have been documented to produce calves at older ages. Killer whales can survive many years after reproductive senescence (Ward et al. 2009a, Ward et al. 2009b; Foster 2012). Although it is not known what age killer whales can reach, and estimates of birth dates for the older whales in the Southern Resident population are uncertain, one female has been estimated to be 104 years old. The next oldest females are estimated to be 87 years and 50 years. We set an upper limit of age to 90 years in our models. However, because females stop breeding long before this, the population growth rate will not be affected by the assumed upper age limit.

We specified the pod (J, K, or L) of each animal, and assigned each newborn in the simulation to the pod of its mother. Southern Residents have not been observed to change pod affiliation, although males will mate with females from the other pods (and also, less often, with females from their natal pod) (Ford et al. 2011). The data on this small population are not sufficient to estimate demographic rates separately for each pod, and it is not known if the pods have differential survival or fecundity. Therefore, we

applied the same demographic rates for all three pods. By tracking the membership of each pod, however, we can project the probability that one or more pods will die out.

As previously noted, the Vortex PVA model that we used is an individual-based model, with the fate of each animal tracked through its lifetime, allowing for determination of matrilines, individual variation in reproductive success, loss of genetic diversity, inbreeding, inbreeding avoidance, and any demographic consequences of these processes. We started the projections of population dynamics with the actual ages and sexes of the killer whales alive as of 31 December 2014. We also specified the mother of each animal, where it had been reported (for 76 of 80 living animals) in published sources (CWR 2015; Ford et al. 2011). These designations of maternity are important because we specified in the simulation model that females would not mate with their father, a son, or a maternal half-sibling (i.e. from the same mother). For the oldest animals in the population, mothers or other close relatives are unknown (other than their own offspring). Thus, the analyses will underestimate the accumulation of inbreeding, because living animals may be related in ways not documented.

Input variable 2: Impacts of inbreeding Long-term pedigree information are not available from which the effect of inbreeding (mating between close relatives) could be determined for killer whales or for any cetacean species. The avoidance of matings between related pod members (Ford et al. 2011), however, indicates that the species has reduced fitness when inbred, as has been observed in almost all mammalian species for which data on inbreeding depression are available (Lacy 1997; O’Grady et al. 2009). Lacking data on the severity of inbreeding depression in cetaceans, we assumed in the model that the effect of inbreeding on reproductive success (birth rate and calf survival) is the same as the mean value reported (6.29 “lethal equivalents”) in a survey of impacts on wild species (O’Grady et al. 2009). Inbreeding might also affect other components of fitness – such as longevity – but data on such effects are too few to allow estimation of typical impacts, and we therefore did not include those effects in our population model. We used the option in Vortex to model inbreeding depression as being caused by the presence of recessive lethal alleles,5 and each initial animal in the model was assigned an average of 6.24 unique lethal alleles across its genome. If inbreeding is instead due in part to other mechanisms – such as deleterious (but not lethal) alleles, or to genetic loci that show heterozygote advantage – then the impacts of inbreeding in early generations projected in the simulation would be the same, but the population would be less able to adapt to inbreeding after many generations than is the case with fully recessive lethal alleles.

5 An allele is one of a number of alternative forms of the same gene. Alleles that cause an organism to die only when present in homozygous (2 identical genes) condition are called lethal alleles. The gene involved is considered an essential gene.

Input variable 3: Mortality rates Mortality rates were calculated from the tallies of deaths each year from 1976 through 20146. The mortality rate for each age-sex class was averaged across the 39 years of data to obtain the mean annual rates. The variation in mortality rates across years has two components – variation in the probability of mortality due to variable environmental conditions (“environmental variation”), and variation in the number of deaths that occur given the independent fates of animals even when exposed to the same probabilities of mortality (“demographic stochasticity”). To determine how much of the observed variation is due to environmental variation, the variance due to demographic stochasticity can be calculated from the expectation for a binomial process7, and then subtracted from the total variation across years (Lacy et al. 2014b). Calculated annual mortality rates and environmental variation (represented as a standard deviation) were as follows:

Calves: 17.48% (SD = 17.96) Juveniles: 2.15% (SD = 2.42) Young adult females: 0.97% (SD = 0; i.e., the variation in mortality across years was no greater than expected based on the demographic stochasticity if population rates were constant over time) Older adult females: 2.25% (SD = 0) Post-reproductive females: 6.29% (SD = 7.77) Young adult males: 3.03% (SD = 2.53) Older adult males: 9.80% (SD = 8.68)

The above mortality rates are marginally better (i.e., lower) than the rates presented in Vélez-Espino et al. (2014a), which is due to their use of data from 1987-2011, which omits a prior decade that had better demographic rates. The estimates of annual variation above are all considerably lower than the estimates by Vélez-Espino et al. (2014a) as a result of our removal of the expected demographic stochasticity from the annual variation in observed death rates.

Input variable 4: Reproductive system The breeding system is polygamous, with some males able to obtain multiple mates, and females mating with different males over their lifetimes. We assumed in the model that males become sexually mature (actively breeding, which may occur several years after they are physiologically capable of breeding) from 12 to 18 years of age. Thus, in the model, each male was assigned an age of sexual maturity by randomly selecting a value from 12 to 18 (assigned when the male was born). There is evidence that there is variation among males and among females in reproductive success. Genetic evidence indicates

6 Mortality was recorded if an individual’s matriline was observed in the population within a year but the individual did not appear. 7 A binomial process is a random counting system where there are n independent identical trials, each one of which has the same probability of success p, which produces s successes from those n trials. The simplest example of a binomial process is the toss of a coin.

that the effective population size is about 1/3 of the total size (Ford et al. 2011), as would occur if some animals are more successful breeders than others.8

To represent this phenomenon in the PVA model, we specified that each male be assigned a score from 0 to 1 to be used as his likelihood of being a capable breeder (i.e. with full access to breeding females) each year. The mean was set at 0.50, with a standard deviation (SD) of 0.20 in a beta-distributed distribution.9 This distribution will lead to most males having relative breeding success of 0.3 to 0.7, with a few extreme males having scores below 0.10 or above 0.90.

Similarly, we assigned an individual reproductive success score to each female, used to represent the factor by which the female’s probability of breeding each year was below (score < 1) or above (score > 1) the overall mean for females of her age. These scores for variation in female reproductive success had mean 1.0 and SD = 0.2, created by sampling from a beta distribution (mean =0.50, SD = 0.1) and doubling those values. (It might be expected that individual variation in reproductive success would follow instead a normal distribution (a “bell curve”), but the normal distribution has no limit, which would allow some animals to have probabilities of breeding that are below 0 or above 100%. The beta distribution has a similar shape to a normal distribution when the mean is 0.5, but it does not extend below 0 or above 1, so it is preferred for representing uncertainty in demographic rates.)

The variance among individual females and males in the number of offspring across their lifetimes will cause genetic diversity to be depleted faster and inbreeding to accumulate faster than would occur if mating was assumed random. With the above specification of variance in reproductive success, our models project an effective population size of 37% of the total size, close to the estimate obtained from genetic data (Ford et al. 2011).

Genetic data indicate that females usually mate with males from the other pods, but occasionally do mate within their own pod (Ford et al. 2011). However, no cases of father-daughter, mother-son, or maternal half-sibling matings were observed, even though the within-pod matings could make such inbreeding likely if mating were at random. Thus, it appears that killer whales generally avoid mating with those relatives in their own pod that can be reliably determined as close relatives. To include this inbreeding avoidance in the model, we specified that females would not mate with their fathers, sons, or brothers or half-brothers from the same mother. This will reduce the likelihood and extent of any damage caused by inbreeding in the closed Southern Residents population, but could also lead to a situation in which some females might be unable to find a suitable mate if the population becomes small enough so that no unrelated breeding males are available.

8 Effective population size is the number of breeding individuals in an idealized population that would show the same amount of dispersion of allele frequencies under random genetic drift or the same amount of inbreeding as the population under consideration. 9 In probability theory and statistics, the beta distribution is a family of continuous probability distributions.

Input variable 5: Reproductive rates

Breeding rates – expressed as the proportion of the females of an age class that produce a calf each year – were calculated to be: 12.04% for young adult females (age 10-30), with SD = 3.54 7.88% for older females (age 31-45), with SD = 4.15 0% for post-reproductive females (age > 45)

The above standard deviations expressing environmentally caused variation in rates across years were adjusted by removing the part of the observed variation that would be expected to be due to the demographic stochasticity (chance fluctuations in numbers breeding) even when population mean rates are constant.

Our estimated fecundities are all a little higher than those estimated by Vélez-Espino et al. (2014a) from the 1987-2011 data. Our estimates of annual variation are lower because we removed the demographic stochasticity.

Input variable 6: Carrying capacity

For our purposes, carrying capacity is the maximum population size of Southern Resident killer whales that the environment can sustain. For the initial tests of the importance of estimates of demographic rates to the population trajectories, we set the upper limit on population size to 300, so that the “carrying capacity” would not restrict population growth under the best conditions tested. (In the projections, the Southern Resident population never reached this limiting size.) In later analyses of the effects of various threats, we set the carrying capacity to K = 150, on the assumption that even under conditions that can allow growth, the population would not exceed 1.5x the largest size observed in recent decades.10 Only rarely in our models (when individual simulated populations experienced unusually lucky good years) did any iteration reach this limit of 150. Setting K = 150 rather than K = 300 had minimal effect on the baseline model, and in none of our models did the K of 150 restrict growth in more than a few percent of the independent simulations.

Simulation settings

We projected the Southern Residents population for 100 years with the above input values. The simulation was repeated 10,000 times (independent iterations) to obtain precise estimates of means and variances in measures of population performance. For analyses of variants on the input rates (see “sensitivity tests”, below) and for comparisons among alternative scenarios, fewer iterations are needed to obtain good estimates of the relative impact of altered input values, and tests of alternative models were run with 1000 iterations of the simulation.

10 NOAA (2014) estimates the historic population size of Southern Residents was about 140 whales.

Sensitivity tests

Sensitivity tests were done to determine the sensitivity of the model projections to the uncertainty in 11 demographic input variables described above. We ran 100 iterations each of 1,000 combinations of input parameters sampled from ranges. The sampled parameters were evenly spaced across the range set for each, with the sampling done according to a “Latin hyperspace” design so that the combinations of the 11 rates were evenly distributed. This method of sampling provides high statistical power for determination of the effect of each parameter. The ranges sampled for each variable were:

Inbreeding effects: Lethal equivalents = 3.145 to 9.435 (+ 50% around the baseline value) Male variance in Reproductive Success: SD = 0 to 0.4 Female variance in Reproductive Success: SD = 0 to 0.4 Birth rate: + 10% around the baseline values Annual variation in birth rate: + 10% around the baseline values Calf mortality rate: + 10% around the baseline values Annual variation in calf mortality: + 10% around the baseline values Juvenile mortality rate: + 10% around the baseline values Annual variation in juvenile mortality: + 10% around the baseline values Adult mortality rate: + 10% around the baseline values Annual variation in adult mortality: + 10% around the baseline values

Thus, for each basic demographic rate (life table values for fecundity and mortality), we tested a range of + 10% around the baseline value. For several model variables that describe other aspects of the population dynamics and are very uncertain, we tested wider ranges of values (see Table 1 below).

The above ranges for sensitivity tests are different from a common practice of examining the amount of change in the population growth for a constant proportional change in each survival and fecundity rate (“elasticities”). The apparent standardization of sensitivities of growth to changes in variables via such elasticities is arbitrary and an artifact of the age classes used for the demographic projections. It is arbitrary because the same proportional change in different demographic rates may not be likely or achievable for the population of interest. (E.g., it might be possible to shift an 80% calf survival rate up by 10%, but it is not possible to similarly shift an adult survival rate of 99% up by a “comparable” 10%.) For other model variables, such as the inbreeding depression, it is not clear that a 10% change in the value has the same meaning as a 10% shift in other demographic rates measured on very different and not always linear scales. The standardization achieved through elasticities is artifactual because very different results can be obtained if ages are binned into more or fewer age class categories for the calculations. Although strict comparison of the effect of proportional changes in model parameter values is usually not meaningful, the sensitivity tests can be useful guides as to which model parameters are the most important determinants of population trajectories, over the range of values that are chosen for testing. Thus, it is often useful to vary parameter values by amounts that reflect our uncertainty in each input value.

3.0. Results of modelling

3.1. Projections for the Southern Resident population under the status quo (baseline model)

With the above model input values, we can project the trajectory of the Southern Resident population under our best understanding of the conditions that have occurred over the past few decades. Across 10,000 iterations, the baseline model projects a mean population growth rate of r = -0.002 (i.e. 0.2% decline per year), with fluctuation across years of SD(r) = 0.045 (i.e. approximately + 4.5% growth). The following graphs illustrate the model with a sample of 100 iterations of the simulation – showing projections with an average slight decline but with much variation among the possible trajectories (Figures 1 and 2). Of course, the real population will experience only a single trajectory out of the range of possible futures. The model also projects a reduction in genetic diversity (measured as heterozygosity) after 100 years to 93% of the starting value. This is equivalent to the individuals in the population becoming inter-related on average to an extent a little more closely than first cousins are.

Figure 1. The baseline model projects a mean or average Southern Resident population growth rate of r = -0.002 (i.e. 0.2% decline per year).

Figure 2. The distribution of simulated Southern Resident population trajectories with means and SD of the projected population sizes.

The model predicts no complete extinctions (no animals remaining), but sometimes the population is reduced to a much smaller size. The risk of population reduction can be represented by a graph of “quasi-extinction” – the probability that the population will have declined below any given N at the end of 100 years. The following quasi-extinction graph shows, for example, about a 7% probability of the population being at N < 30 after 100 years (Figure 3). Quasi-extinction threshold is the minimum number below which extinction is very likely due to genetic or demographic risks. For this review, the quasi- extinction threshold is 30 killer whales, a level at which the Southern Resident are unlikely to persist. Graphs of this style will be used in this report to show the effects of various scenarios on the likelihood of population collapse.

Figure 3. The probability of the population becoming quasi-extinct (N < 30) after 100 year is about a 7%.

It is also sometimes useful to look at an “interval quasi-extinction” graph – which shows the probability of the population falling below any given N at any time during the 100 year simulation. For example, the graph below shows that there is about a 9% chance of the “baseline” population falling below N = 30 at any time over the next 100 years (Figure 4).

Figure 4. The chance of the “baseline” population falling below N = 30 at any time over the next 100 years is about a 9%.

Model validation

As stated above, the Vortex population viability analysis program and manual are available free from www.vortex10.org/Vortex10.aspx. The input files that we created for modeling the population are available at www.vortex10.org/SRKW-Vortex.zip, so all input values are available for review and the analyses can be repeated and extended by others.

The baseline model projects a very slight average decline, with a mean population growth rate of r = - 0.002, and a variation across years of SD(r) = 0.045. The observed mean growth rate from 1976 to 2014 was r = 0.002, with a variation of SD(r) = 0.042. The observed growth rate was higher (mean r = 0.017) before the peak of N = 94 individuals in 1993 and has subsequently been negative (mean r = -0.009). Thus, the baseline PVA model accurately reflects the population dynamics since 1976; although projections forward in time should be made with the caution that it is not known which recent time span best represents the current and future conditions if threats remain at the status quo. The mean age of living animals projected in the model is 24.2 years, while the observed current mean age is 21.2 years. This difference may reflect the consequence of animals having been removed from the population in previous decades, thus reducing the number of older animals currently in the population.

Other PVA models, based on fewer and less recent years of data have projected either lower (e.g. Taylor and Plater 2001; Vélez-Espino et al. 2014a) or higher (e.g., Olesiuk et al. 1990) population growth rates. These differences are consistent with the models that used demographic data from years of decline

projecting continuing decline, and the models relying on data from earlier years of growth projecting continued growth. We used demographic data over the full span of years starting in 1976, and our population model predicts almost no mean change in population size – as has been observed over that 39-year time span.

3.2. Sensitivity testing of important model parameters

For the model parameters that might have significant effect on population growth, and yet the values currently and in the future are uncertain, we ran simulations with higher and lower values to determine the importance of each to the population performance. For each model parameter that was tested, the Table 1 below gives the value used in the baseline model, the minimum and maximum values of the range that was tested, the population growth rates that resulted from the min value (r-min) and max value (r-max) for that parameter while all other parameters were held at their baseline, and the % of variance in population growth (as a percent of the total variation accounted for by the varied parameters) due to variation in that parameter when all other parameters were simultaneously and independently varied across their ranges of uncertainty. Cells with two or three values for the rate are for the several age categories.

Table 1. The importance of model parameters that potentially affect the population growth rate of the Southern Residents. The higher the %variance the greater the importance of the parameter. Parameter Baseline Min tested Max tested r-min r-max %variance

Inbreeding 6.29 3.145 9.435 -0.0005 -0.0029 6.1 depression Variance in 0.20 0.00 0.40 -0.0017 -0.0023 0.4 male RS Variance in 0.20 0.00 0.40 -0.0017 -0.0019 0.0 female RS Mean birth 0.1204 0.1084 0.1324 -0.0062 0.0019 83.0 rate 0.0788 0.0709 0.0867 SD in birth 0.0354 0.0319 0.0389 -0.0020 -0.0018 0.0 rate 0.0415 0.0374 0.0457 Calf 0.1748 0.1573 0.1923 -0.0012 -0.0029 3.6 mortality SD in calf 0.1796 0.1616 0.1976 -0.0017 -0.0019 0.0 mortality Juvenile 0.0215 0.0194 0.0237 -0.0011 -0.0028 3.6 mortality SD in juv. 0.0242 0.0218 0.0266 -0.0015 -0.0019 0.0 Mortality Adult various (see base * 0.9 base * 1.1 -0.0010 -0.0028 6.4 mortality text) SD in adult various (see base * 0.9 base * 1.1 -0.0018 -0.0018 0.0 mortality text)

As shown in the table above, the estimates of environmental variation (EV, expressed as SDs) for each demographic rate had no effect on the population growth rate (within the range tested). Therefore, although these estimates of annual variation in rates are uncertain, refining the estimates would not change any conclusions about threats and the viability of the population.

Variance in reproductive success of males had a very small effect. Inbreeding depression and each of the mortality rates had moderate effects, while the birth rate (calves per adult female) dominated the variation in population growth. This is counter to a commonly reported trend for sensitivity of survival rates being more important than fecundity in long-lived species. However, this difference can be ascribed simply to the difference in whether sensitivity is tested for survival vs. mortality. (e.g. a 1% change in a survival rate of 90% is the same as a 10% change in the equivalent 10% mortality rate. Thus, growth rate will often be most sensitive to a change in survival, less sensitive to the same proportional change in fecundity, and even less sensitive to the same proportional change in mortality.)

Note that the significant impact of a shift in inbreeding depression indicates that the population is small enough so that the effects of inbreeding are likely to be damaging to the prospects for the population. This effect would be exacerbated further if we did not include in the model an avoidance of very close inbreeding (within matrilines). If the effect of inbreeding depression on calf survival is removed entirely from the model, then the projected growth rate becomes slightly positive, at r = 0.0006.

3.3. Examination of potential threats

The sections below describe the modeling to examine the likely impacts of several kinds of existing or potential threats to the Southern Residents. Text descriptions of these scenarios and brief summaries of the results are presented, followed by a summary table comparing results across these various scenarios.

Modeling of impacts of Chinook salmon abundance

Several factors have modified the Southern Residents’ habitat, including changes in prey availability, contaminants, and vessel traffic. Reductions in salmon and other prey along much of the west coast of North America during the past 150 years, especially from Washington to California (Nehlson 1997, Kope and Wainwright 1998), have likely lessened the region’s carrying capacity for resident killer whales (Krahn et al. 2002). Resident whales likely forage selectively for Chinook salmon over other available salmonids because of the large size, high fat content, and year-round availability of this species in coastal waters (Ford et al. 1998, Ford and Ellis 2005). Ford et al. (2005) found that trends in the mortality rates of southern were strongly related to fluctuations in the abundance of Chinook salmon. Birth rates were also correlated with Chinook salmon abundance, but more weakly than mortalities.

Ward et al. (2009b), Ford et al. (2015), and Vélez-Espino et al. (2014b) all provide estimates of the relationship between abundance of Chinook salmon and demographic rates for Southern Residents. The details of the data used to quantify Chinook abundance, the co-factors considered, and the statistical

methodologies differ in many details between these studies. Vélez-Espino et al. examined fecundity for young and old females separately, and examined impacts on survival rates of different age classes, while the other papers assessed relationships with pooled demographic data. Ward et al. used Pacific Salmon Commission indices of Chinook abundance off the west coast of Vancouver Island; Ford et al. used Pacific Salmon Commission indices for the coastal region for coastal regions from SE Alaska to Oregon; and Vélez-Espino et al. used indices for terminal run abundance and ocean abundance for various salmon stocks and combinations of stocks in the region. All three studies examined the relationships with various time lags (0 to 2 years), although for projecting long-term population viability in simulation models, it is not important if changes in fecundity due to changing Chinook abundance occur in the year of the Chinook index or with a lag of a year or two.

All three previous studies identified large effects of Chinook salmon abundance on Southern Resident demographic rates. Ward et al. and Vélez-Espino et al. found large effects of Chinook abundance on fecundity, whereas Ford et al. and Vélez-Espino et al. reported significant relationships with mortality. However, Vélez-Espino et al. found the relationships of Chinook abundance with fecundity had greater influence on projected population growth than did the effect on survival, while Ford et al. reported a weaker (but still significant) effect on fecundity compared with mortality.

Both Ward et al. and Vélez-Espino et al. used regression models to determine the relationship between salmon abundance and logit(birth rate). The regressions provided by both studies can be rescaled so that Chinook abundance index is set to 1.0 for the average over the span of data (1976-2014 for our demographic analyses), and so that the regression predicts the observed birth rate when Chinook index = 1. After that rescaling, the relationship for overall fecundity reported in Ward et al. and for fecundity of the younger (more productive) females reported in Vélez-Espino et al. (for terminal run of Fraser late stock) are very similar and are both approximately: logit(birth rate) = -3.0 + Chinook, in which Chinook is the abundance relative to the long-term average.

This relationship leads to a predicted shift from a birth rate of 12% to 18.5% if the Chinook abundance increases by 50%, and a shift down to 7.6% if abundance decreases by 50%. The linear relationship reported by Ford et al. would lead to smaller but still significant shifts (up to 13.3% and down to 9.2%, respectively) in the birth rate.

Vélez-Espino et al. reported a stronger effect on fecundity of older females in relation to the Puget Sound terminal run, but a similar slope for Fraser + Puget abundance. If the intercept in the above regression is adjusted so that the fecundity of the older breeding females is correctly predicted at the baseline of Chinook = 1, then the relationship for that age class becomes: logit(birth rate) = -3.46 + Chinook

Ford et al. (2015) provide the relationship of mortality to Chinook abundance in a form that is easily transferred to the Vortex model, namely: Mortality index (relative to the expected value) = -2.6504 x Chinook (relative abundance index) + 4.0066 Rescaling this relationship so that the mortality index is 1 when Chinook = 1 gives:

Mortality index (relative to the baseline) = -2.6504 x Chinook (relative abundance index) + 3.6504 This relationship translates to a 26% increase in mortality for each 10% decrease in Chinook abundance.

The rescaling of the relationships above was confirmed to produce the same population growth in the Vortex model (r = -0.002, with SD = 0.045) as was projected from the baseline model, as it should. These general relationships can then be used for projecting the primary impacts of changing Chinook abundance on population viability. For purposes such as managing threats via individual stocks, or assessing also the lesser impacts on other demographic rates, the more specific regression analyses presented in Vélez-Espino et al. (2014b) would provide models more tailored to specific questions.

With the above relationships of Southern Residents demography to Chinook abundance, we modeled the fate of the Southern Residents if Chinook numbers were set to 80%, 90%, 100%, 110%, or 120% of current (baseline) levels. Muñoz et al. (2014) also projected that Chinook could suffer a catastrophic decline within the next 60 to 90 years due to climate change raising the water temperatures occasionally above the thermal tolerance of the Chinook. Although the time course of any decline in Chinook is unknown as global temperatures steadily rise, we examined scenarios in which Chinook decline gradually but steadily by 0.25% per year or 0.50% per year (resulting in a 25% or 50% loss over 100 years).

As found in other studies, reducing the abundance of Chinook is projected to have a substantial negative impact on the Southern Residents population. The combined effect on fecundity and survival result in the mean population growth falling to r = -1.4% or -2.6% for a 10% or 20% reduction in prey. Moreover, the probability of extinction rises to 4% or 39%, respectively, whereas the probability of the population declining to below 30 animals rises to 73% or over 99%. Conversely, increasing Chinook abundance can lead to relatively robust population growth (up to 1.9% with 20% more prey) and protection from extinction or serious decline. If a decline of 25% or 50% occurs gradually over the next 100 years because of climate change (or other factors), the consequences for Southern Residents population viability are about comparable to an immediate loss of about half the magnitude.

The quasi-extinction graph below demonstrates the dramatic effect that can occur with shifts in the abundance of Chinook prey (Figure 5).

Figure 5. The graph illustrates the effects that can occur with varying abundances of Chinook prey, which is the primary food for Southern Residents. Reducing the abundance of Chinook is projected to have a substantial negative impact on the Southern Residents population, whereas increasing abundance leads to robust population growth and protection from serious decline or extinction.

Modeling oil spill impacts

While the probability of either Northern or Southern Resident killer whales being exposed to an oil spill is low, the impact of such an event is potentially catastrophic. Southern Residents are at risk of an oil spill because of the existing tanker traffic that travels in and out of Puget Sound and the Strait of Georgia (Baird 2001, Grant and Ross 2002) and the proposed expansion of tanker traffic associated with Project.

Impacts of oil spills are tested in the model by adding two types of catastrophes – large oil spill and smaller oil spill. Catastrophes are assumed to occur at random infrequently (i.e., with low probability of occurrence in any given year) but to affect survival or reproduction when they occur. In the model, the impacts are deaths that occur in the year of the spill. The deaths occurring because of an actual spill may occur over several years, but presumably the females weakened by the spill would not be successful breeders before they die from the spill, so they can be removed from the population in the model at the time of the spill.

The initial frequency of a big or “worst case” spill (>16,500 m3) is based on an analysis by Foschi (2014). He used Kinder Morgan’s projections, which were calculated as return periods (in years) for oil spills of different volumes (Table 34 of their TERMPOL Report 3.15.), to derive a 10% probability of an oil spill in 50 years, corresponding to a 0.21% chance per year. The initial frequency of smaller spill (>8,250 m3)

was estimated to be 42% in 50 years, corresponding to a 1.08% chance per year. The basis for the Kinder Morgan estimates is not clear, and it is difficult to consider the threats of smaller volume spills (<8,250 m3) because the reports do not clarify the average volume or the distribution of volumes of those small spills. To encompass some of the uncertainty, we therefore also tested the projected effects of frequencies of large and smaller spills that are double those stated above.

The mortality caused by the spills is the product of the percent of the Southern Resident population that would become exposed to damaging doses of oil products and the percent that die from that exposure. Based on the percent overlap of oil coverage and critical habitat, as determined from Trans Mountain Pipelines oil spill models (Modeling the Fate and Behavior of Marine Oil Spills for the Trans Mountain Expansion Project. 2013. [NEB Exhibit No. B21-9 to B23-28, V8C TR 8C 12 TR S9 and V8C TR 8C 12 S9a APPE, A3S5G9 to A3S5H5]. EBA File: V13203022. Prepared for Trans Mountain Pipelines ULC.), we estimate that about 50% of the Southern Residents would encounter a large spill. We estimate that half this percent (25%) would encounter a smaller spill (of half the oil volume) (Figures 6 and 7)).11 We also assumed that a Southern Resident that has direct exposure to a large spill would have 100% mortality. Small spills are assumed to cause 50% mortality to those Killer Whales that are exposed to the oil. Thus, the overall mortality due to a large spill in the critical habitat would be 50%, and overall mortality due to a smaller spill would be 12.5%.

11 Whether an oil spill would directly affect the Southern Residents depends largely on traffic levels, the size of the whale population (i.e., the larger the population the greater the probability of exposure), and the distribution of whales relative to the spill. All of the Southern Residents are periodically in the same location at the same time (known as a “super pod”), making the whales very vulnerable to a catastrophic event like an oil spill during such times (NOAA 2014).

Figure 6. The figure shows the probability of oil presence and dispersion within the Salish Sea following an oil spill during summer. The overlap with critical habitat of Southern Resident killer whales is significant.

Figure 7. The figure shows the probability of oil presence and dispersion within the Salish Sea following an oil spill during the fall. The overlap with critical habitat of Southern Resident killer whales is significant.

Oil spills of the frequency estimated and with the estimated extent of overlap of oil with critical habitat would cause some depression of mean population growth (to r = -0.005 as opposed to r = -0.002 without oil spills) and the mean final population after 100 years (N = 56 vs. N = 72 with no oil spills). The probability of complete extinction would rise to 1%, but the probability of quasi-extinction (below N=30 threshold) rises to 24.3% (vs. 7.1%). If oil spills occur at a frequency twice that projected by Kinder Morgan, then the mean prospects for the population are worse (r = -0.008, final N = 46) and there becomes a 35.8% chance of quasi-extinction below the N = 30 threshold (see graph below) and a 3.5% chance of complete extinction of the Southern Residents. Thus, the impact on mean long-term population growth of rare oil spills could be moderate, because in most years no spill occurs, but the effect on probability of the population dropping to low numbers (e.g., quasi-extinction below N = 30) is more substantial, with a possibility of complete extinction within 100 years due to the potential of catastrophic decimation of the population to a small number from which it could not recover.

Figure 8. This figure shows the effects of oil spills on the mean population growth rate of Southern Resident killer whales. The rate of occurrence of various sized oil spills was determined using an analysis of spill probably frequencies provided by Kinder Morgan in the Trans Mountain Pipeline application. These are given in Table 34 of their TERMPOL Report 3.15.

Modeling impact of increased boat noise

Noise from boats near Southern Residents can disrupt feeding activity and reduce foraging efficiency by masking echolocation (Figures 9 and 10). However, the reduced feeding activity will likely have a much larger impact than the reduced foraging efficiency, and we will focus our analyses on the former.

Southern Residents reduce feeding activity by 25% around boats. Presently, boats are present an estimated 85% of the time that Southern Residents forage (there is no evidence that they have shifted foraging activity to the nighttime to avoid recreational and whale-watching boats), and they are foraging in the presence of boats an estimated 78% of that time. Thus, feeding is expected to be reduced by 16.6% (= 25% x 85% x 78%) due to disturbance by boats (Bain 2002, Williams et al. 2006, Lusseau et al 2009, Lusseau and Bejder 2007)

With increased shipping traffic associated with the Project, Southern Residents could be around boats up to 100% of the time – according to the proponent tankers would be a “near continuous” presence. Thus, feeding could potentially be reduced by up to 19.5%.

Figure 9. Noise from boats near Southern Residents can physically disrupt feeding activity and reduce foraging efficiency by masking echolocation.

Figure 10. This figure shows the amount of time Southern Residents from J, K, and L pod spent foraging, resting, traveling, and socializing with and without the presence of boats in two areas near San Juan Island. When vessels were present, the whales foraged significantly less and travelled more. Figure from NOAA (2014), Data from Lusseau et al. 2009.

If we assume that the effect on demographic rates of reduced feeding activity is the same as a comparable reduction in prey, then we can use the relationships described above for the effects of reduced Chinook abundance to estimate also the effects of reduced feeding activity due to the presence of boats. Thus, the foraging success relative to the baseline conditions with current levels of disturbance by boats would be: Proportional time spent feeding = (1 – (0.25 x 0.78 x BT)) / (1 – (0.25 x 0.78 x 0.85)) = (1 – (0.195 x BT)) / 0.83425), with BT being the proportion of the time that boats are present.

We tested values of BT = 85% (baseline), BT = 92.5%, and BT = 100% to determine if increase disturbance of feeding behavior could have a significant influence on Southern Resident demography. With boats always present (BT = 100%), the reduction in foraging is about 3.5% and this resulted in accelerated population decline (r = -0.007), smaller mean size (N = 47), 1% probability of extinction, and 25% chance of falling below 30 animals. The impact of an increase in boat presence of half as much (BT = 92.5%) caused proportionately about half as large shifts in population viability relative to the baseline (Figure 11).

Figure 11. This figure show that increasing disturbance of feeding behavior above current levels would affect Southern Resident killer whale demography, resulting in accelerated population decline, smaller mean population size, and increased chance quasi extinction.

Clearly, increased recreational, fishing, whale-watching, and shipping traffic can affect Southern Residents also in other ways, such as increased frequency of injury from boat strikes, and an increase in the small scale pollution from spilled fuel and emissions. The effect of additional boat strike mortality can be assessed with the models presented below for “Modeling population impacts of human-caused deaths”, although we have not attempted to determine how the frequency of such injuries will change. We also have not tried to assess the health effects on Southern Residents of increased pollution of the water and air.

Modeling impact of PCBs on Southern Residents

PCB burdens in Southern Residents are at harmful levels (Garrett and Ross 2010) and considered a primary ongoing threat by the Recovery Strategy in Canada (Fisheries and Oceans Canada 2011) and the U.S. (National Marine Fisheries Service). The detrimental health effects of carrying large contaminant burdens interact with other threats such as prey limitations and disease, and increase susceptibility to other threats emanating from vessel traffic, such as acoustical disturbance or oil spills (Figure 12). Our basic model of accumulation, depuration,12 and impact on calf survival of PCBs was based on the

12 The process of freeing from contaminants; in this case PCBs.

methodology of Hall et al. (2006, 2011, 2012), with modifications in rates for Southern Residents as described below.

Figure 12. Biological magnification of persistent organic pollutants in Southern Resident killer whales.

We assumed that Southern Residents obtain the initial load of PCBs from their mothers, but that most of that initial load is removed, primarily through a growth dilution, by the time that the animals reach 10 years of age (Hickie et al. 2007). The impact of PCB on population growth is being modeled only as the reduction in survival of calves due to the PCB loads of their mothers, so the initial PCB level before age 10 years is not important in our model. Note that we did not attempt to project the effect of PCBs on disease susceptibility, as was done in Hall et al. (2012).

We initially modeled the accumulation of PCBs as a linear function: PCB = 1 + 5 * Age, extrapolating back to a conservative estimate of 1 mg / kg remaining from the initial PCB load obtained from the mother,

and accumulating thereafter at a rate of 5 mg/kg/y. Breeding females are projected to have a 0.77 proportional reduction in their PCB levels through depuration when they raise a calf. The rate of accumulation that we used is higher than the mean (2.96 mg/kg/y) that has been reported in a population of bottlenose dolphins that does not have high PCB levels in adults (Wells et al. 2005), and within the range tested for cetaceans by Hall et al. (2011). This rate of accumulation leads to a mean PCB level in our simulations of 57 mg/kg in breeding adult females (9 < Age <46) and 106 mg/kg in males 10 y and older. These mean projected PCB loads are consistent with few data on Southern Residents reported by Ross et al. (2000), who found 55 + 19 mg/kg in two adult females and 146 + 33 mg/kg in four adult males.

For the impact of maternal PCB on calf survival, we assumed a linear regression on logit(survival), of the same form as used by Hall et al. (2006). We determined the slope and intercept that fit the two observed data points for Southern Residents (survival = 0.8252, mean PCB = 55.4 [Ross et al. 2000]) and Northern Resident Killer Whales (survival = 0.9218 [Vélez-Espino et al. 2014a], mean PCB = 9.3 [Ross et al. 2000]). This yielded a relationship of: logit(survival) = 2.65 – 0.02 * PCB. This relationship predicts a calf survival rate of 93.4% in the absence of PCBs, consistent with the higher estimates of calf survival reported for growing killer whale populations (such as Northern Residents).

When the accumulation, depuration, and impact on calf survival of PCBs were added to our baseline model, the mean population growth was r = -0.002. This is the same as the baseline model without consideration of PCBs, as is expected because the effect of PCBs on calf survival was scaled so as to produce the baseline survival rate with the average level of PCBs in the mothers. If the PCB level is set to 0 for all animals in the model with consideration of PCBs, then the mean population growth is projected to be r = 0.003. This suggests that PCB effects model can be a basis for examining likely effects of increased or decreased PCBs in the population.

Modeling population impacts of human-caused deaths

Various human activities have a risk of causing a fatal injury to a Killer Whale. These include accidental collisions with recreational or commercial boats, entanglement in fishing gear, and intentional harm to animals. Documented causes of death include ingested fishhook with trailing line, trauma from ship strikes, and Salmonellla septicemia (Gaydos and Raverty 2007). Although such events are likely rare, in a small population with low birth rates, even a few extra deaths might cause significant damage to the population growth, size, and even persistence. To explore the possible population consequences of additional deaths (or removals of any sort) caused by human activities, we ran simulations in which one KW was removed on average every 15 years, 10 years, or 5 years, which reflects the rate of documented human induced mortalities for the Southern Residents during the last 30 years (K. Balcomb pers. Comm., Gaydos and Raverty 2007). In the model the removals were stochastic (with constant probabilities of the loss of an animal over time), had equal probability of the removed animal being male or female, and removed adults (> 9 years of age) otherwise, without regard for their age.

The removal of one additional adult Killer Whale from the population every 10 years or every 15 years had little effect on population growth (from r = -0.002 to r = -0.003), and caused a small increase (from 7% to 10%) in the probability that the population would be below 30 animals after 100 years. Removal of an animal every 5 years had a more noticeable impact on average population growth (reducing to r = - 0.005), caused a 2% probability of extinction and 19% probability of being below 30 killer whales (Figure 13).

Figure 13. Various human activities have a risk of causing a fatal injury to a killer whale. These include accidental collisions with recreational or commercial boats, entanglement in fishing gear, and intentional harm to animals. To explore the possible population consequences of additional deaths (or removals of any sort) caused by human activities, we ran simulations in which one killer whales was removed on average every 15 years, 10 years, or 5 years as shown in the figure.

Summary and comparison of the impacts of potential threats

The table below summarizes the impacts of several levels of each of the kinds of threats that we have examined (Table 2). For each threat model, the columns show the mean population growth rate, mean population size after 100 years, standard deviation in population size, probability of complete extinction, and probability of quasi-extinction (defined as being at or below N = 30 after 100 years). Note that for the Baseline scenario, 10,000 independent iterations were run. For each alternative scenario, 1,000 iterations were run, resulting in outcomes that are less precise. Usually the last digit shown is within + 1 of the value that would be obtained with many more iterations of the scenario.

Table 2. Summary of the kinds impacts of potential threats that we examined at several levels. Threat mean r final N SD(N) Pr[extinct] Pr[N <= 30] model

Baseline -0.002 72 31 0.1% 7.1%

Chinook80 -0.026 7 7 39.4% 99.4% Chinook90 -0.014 24 15 4.2% 73.1% Chinook110 0.008 135 19 0% 0% Chinook120 0.019 145 8 0% 0% ChinookCC25 -0.016 20 11 6.4% 84.6% ChinookCC50 -0.027 8 4 50.6% 99.9%

PCB90 -0.002 71 32 0.1% 8.9% PCB110 -0.003 63 28 0.5% 11.0%

OilSpills -0.005 56 31 1.0% 24.3% OilSpills2x -0.008 46 28 3.5% 35.8%

Noise92.5 -0.004 59 26 0.3% 13.8% Noise100 -0.007 47 23 1.0% 24.5%

Removal15y -0.003 68 31 0% 10.3% Removal10y -0.003 65 31 0.6% 10.9% Removal5y -0.005 58 30 2.0% 19.1%

Cumulative -0.012 33 23 8.6% 53.5%

Threat models tested (see text for further explanation of each): • Baseline = no additional threats • Chinook80 = Chinook reduced to 80% of current • Chinook90 = Chinook reduced to 90% of current • Chinook110 = Chinook increased to 110% of current • Chinook120 = Chinook increased to 120% of current

• ChinookCC25 = Chinook projected to decline by 0.25% per year due to climate change • ChinookCC50 = Chinook projected to decline by 0.50% per year due to climate change • PCB90 = PCB accumulation rate set to 90% of baseline • PCB110 = PCB accumulation rate set to 110% of baseline • OilSpills = Large and smaller oil spills at annual probabilities of 0.21% and 1.0835%, respectively • OilSpills2x = frequency of spills doubled over the above • Noise92.5 = Feeding time reduced because of boat presence 92.5% of the time • Noise100 = Feeding time reduced because of boat presence 100% of the time • Removal15y = remove on average an adult every 15 years • Removal10y= remove on average an adult every 10 years • Removal5y= remove on average an adult every 5 years • Cumulative = OilSpill + Noise100 + Removal10y

From the above results, it is clear that even small changes in the abundance of Chinook could have substantial impacts on the Southern Residents population. A 10% increase in prey changes the population to a positive population growth, and eliminates the chance of extinction or dropping to very small size. A 10% decrease in prey results in a 73% (in contrast to 7% for the baseline) probability of declining below 30 animals. A 20% reduction in prey leads to almost a 100% chance that the Southern Residents population will fall below 30 animals. If Chinook decline gradually down to 75% or 50% of current numbers due to climate change over the next century, then the effects on measures of population growth and viability are about half as severe as when a comparable decrease occurs immediately.

A change of 10% in the rate of accumulation of PCBs has a relatively small effect.

Adding the expected possibility of oil spills to the model moderately decreases long-term average population growth, considerably increases the probability of a major decline to below 30 animals, and leads to a low but not insignificant possibility of complete extinction.

Noise disturbance at the 100% level, because of adverse effects on foraging, decreases long-term average population size, accelerates the probability of population decline, and increases the chance of quasi extinction.

Removing an additional animal every 10 to 15 years caused only slight decreases in population performance, as expected considering that 1 animal every 10 years removes, on average, a little more than 0.1% of the population. Removing an animal every 5 years is enough to accelerate decline noticeably and cause some chance of extinction.

Note that the amounts of change tested above do not necessarily represent projections of the changes to expected threats, because often data on which to make such predictions are not available. However,

the ranges tested do allow comparison of the magnitude of impacts with several plausible levels of each threatening process.

Modelling the cumulative impacts of threats

The above tests examine each threat on its own, but the Southern Resident population may be facing increasing pressures from multiple threats. We tested one scenario that combines some threats likely to intensify if there is an increase in shipping of oil through the critical habitat of the Southern Resident population and no adequately compensatory actions are taken to offset threats. In this cumulative impacts model, we imposed the effects of the forecast frequency of oil spills (as in the “OilSpills” model above), the effects of noise on foraging efficiency if boat traffic increases substantially (“Noise100”), and the impact of the death of one additional animal per 10 years due to boat strikes (“Removal10y”).

The cumulative impact of those three threats together resulted in a population decline of more than 1% per year, resulting in a mean final population size of only 33 animals, a 8.6% probability of complete extinction of the Southern Residents, and more than 50% probability that the final population would be less than 30 animals and, therefore, probably on a course toward nearly inevitable extinction. The following 3 graphs present the results of this cumulative effects model on various measures of population viability, in comparison with the models of the 3 threats individually and the baseline “status quo” model (Figures 14, 15, 16).

Figure 14. The graph shows the mean population size averaged across 1,000 iterations of each simulation.

Figure 15. The graph shows the distribution of population size in the 1,000 iterations of each simulation, and the range of final population size that was generated by each of the models.

Figure 16. The graph of quasi-extinction shows the probabilities for each model of the population size being below 30 killer whales at 100 years.

4.0. Conclusions

The Southern Resident population has experienced almost no population growth over the past 4 decades, and has declined in the last two decades. Models projecting population changes based on average demographic rates and the fluctuations in those rates across time project that under the status quo conditions the population will most likely remain about at its current size or continue a very slow decline. Potential population growth is inhibited in part by the very small size of the population and the consequent lack of available unrelated mates for breeding females. Thus, even without modeling new or increased external threats associated with the Project, it is clear that the population cannot withstand additional negative pressures.

In our modeling, we projected the impacts of several kinds of threats that may become worse over time through the approval of the Project. The factor projected to have the largest effect on depressing population size and possibly leading to extinction was a reduction of the Chinook prey base. Noise disturbance that affects foraging efficiency for Chinook has a moderate but significant influence on population viability. Large or medium oil spills occurring at the frequency suggested in Kinder Morgan’s own oil spill analysis could depress long-term average population growth and lead to a considerable chance of the population declining below 30 animals (quasi-extinction). Progressively smaller impacts were seen with disturbance of feeding by boats, removal (human-cause death) of an animal every 5 years, and small changes in the rate of accumulation of PCBs (used as proxy for environmental contamination) acquired from the environment. However, it should be noted that all of the threats

tested caused some reduction in population viability. Reduction of some of the existing pressures on the population is necessary for population growth to resume and for there to be a reasonable expectation of recovery to a more viable level.

In our models the probability of complete extinction (no remaining animals) within 100 years in our analysis of Project effects is low. However, because individual killer whales can live to be almost 100 years old, and a demographically collapsing killer whale population will often have at least a few individuals still alive 100 years from now, this appearance can be misleading. The probability of the population dropping below 30 animals (about the size of one current pod) is the more relevant measure of threat to population persistence. That probability is unacceptably high in all of the scenarios to which we added new threats associated with the Project.

We combined into one model the three threats that are projected to result from the marine shipping of oil through the critical habitat associate with the Project: the possibility of oil spills at the frequency estimated by Kinder Morgan, projected effects of noise if there becomes high frequency of shipping traffic, and occasional additional mortalities due to boat strikes (one in 10 years). This cumulative effects model indicates that the Southern Residents population becomes vulnerable to extinction (8.6% probability within 100 years), and faces more than 50% probability of decline below 30 animals. We have not yet examined the many other possible combinations of cumulative impacts of threats occurring together, including the effect of the above three threats if compounded or offset by changes in other pressures such as changes to the abundance of Chinook salmon. Avoidance or full mitigation of these threats will be required to give the Southern Residents population a good chance to persist, and improvements to current conditions for the population will be required to permit the growth that is necessary to meet recovery criteria.

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Appendix A. Review of Previous Southern Resident Population Viability Analyses

In recent years, numerous Population Viability Analyses (PVAs) have been used to estimate growth rates and extinction risk of the Southern Residents (Plater 2001, Taylor and Plater 2001, and Krahn et al. 2002, 2004, Vélez-Espino et al. 2013, 2014). Although methodologically similar, these PVAs differ somewhat in biological details, time frames, and suppositions as appropriate for the specific objectives of the various analyses. As would be expected, in all analyses extinction risk increased when the frequency and magnitude of catastrophes such as oil spills were elevated, or carrying capacity1 declined (Taylor and Plater 2001, and Krahn et al. 2002, 2004). Similarly, declines in prey availability (Vélez-Espino et al. 2013, 2014) were shown to adversely affect population persistence. Based on these analyses and assuming that no attempt is made to alleviate or lessen current threats to the population’s viability, a slow decline and likely extinction of the Southern Resident population can be regarded as inevitable – this is the trajectory that protection under the Species at Risk Act is intended to halt. Unless wholly mitigated, the addition of disturbances such as increasing shipping traffic and catastrophic events such as oil spill will hasten the demise of the species.

1.0 2001 PVAs

In the first comprehensive PVA of Southern Residents, Plater (2001) used demographic data from 1974– 2000 to estimate mortality and fecundity rates. Depending on model assumptions regarding the maximum population size that the environment could sustain indefinitely (i.e. carrying capacity), sex ratio at birth, and potential effects of risk factors on reproduction and survival, the time to extinction ranged from an estimated 113 to greater than 300 years.

A second series of PVAs incorporated a much wider range of assumptions and risk levels (Taylor and Plater 2001), using data from the 1996–2000 decline to estimate mortality and fecundity rates. These models included the multiple impacts of higher inbreeding, oil spills, epizootics (disease events), and reduced food supply. Addition of these threats resulted in the prediction of a greatly increased risk of extinction - a 33-100% chance of extinction in 100 years. Regardless of the assumptions made in any given PVA in this series of PVAs, each calculated that the population would go extinct within 200 years and the median predicted time to extinction ranged from 54 to 112 years, depending on the model assumptions.

The estimated risk of extinction in this second series of PVAs was very sensitive to assumptions regarding the average probability of survival and reproduction (i.e. survival rates) of Southern Residents in the future. When a PVA assumed that the previous 29 years were a good indicator of future population trends, then the estimated probability of extinction was much lower than if a more pessimistic assumption was made that the previous 10 years, during which the population declined, were a good indicator of the future. Under the pessimistic assumption, uncertainties with respect to the level of carrying capacity, and the rate and magnitude of catastrophes, had relatively little effect on the extinction probability, which was high regardless. In contrast, under the more optimistic prediction using the 29-year series of survival rates, the risk of extinction was strongly influenced by the assumption made about carrying capacity and was to a lesser extent influenced by assumptions about the level and magnitude of catastrophes.

1 The carrying capacity is the maximum population size of the species that the environment can sustain indefinitely, given the food, habitat, water, and other life requisites available in the environment. 1

2.0 2002 and 2004 PVAs

Two associated PVAs of Southern Residents were conducted by the National Marine Fisheries Service’s Biological Recovery Team to synthesize the potential biological consequences of a small population size, a slowly increasing or a declining population trend, and potential risk factors (Krahn et al. 2002, 2004). In both analyses, the probability of the population going extinct was estimated using demographic information from the yearly census of the population. Notably, the original PVA (Krahn et al. 2002) was adjusted and updated (Krahn et al. 2004) to incorporate data concerning the Southern Resident population collected through 2003. Because the two PVAs are nearly identical, we only describe the results of the modified and most recent PVA.

The recovery team determined both the probability of extinction (defined as <1 male or female) and the probability of “quasi-extinction” (defined as < 10 males or females) were determined, because the recovery team believed that a population at the quasi-extinction level would be destined to extinction, even though literal extinction might still take decades given killer whales’ long lifespans.

When the team assumed that growth rates in the future would be most accurately predicted by the full 29-year time series of available data on the population (the most optimistic scenario considered), the model predicted the probability that Southern Resident would become extinct was <0.1–3% in 100 years and 2–42% in 300 years. If the quasi-extinction threshold was used instead of actual extinction, the predicted probability of meeting the threshold ranged 1–15% in 100 years and 4–68% in 300 years. For both scenarios, the higher percentages in each range were associated with assumptions of higher probability and higher magnitude of catastrophic mortality events (e.g. oil spills), and associated with a smaller carrying capacity of 100 whales.

When it was assumed that the population survival for the previous 10 years would best predict the future (the most pessimistic model), the analysis predicted a probability of extinction of 6–19% in 100 years and 68–94% in 300 years. If a quasi-extinction threshold was used instead of actual demographic extinction, the predicted probability of meeting the threshold ranged from 39–67% in 100 years to 76– 98% in 300 years.

3.0 2014 PVA

Vélez-Espino et al. (2014) assessed the population status of Southern Residents and the population’s potential responses to non specific management actions that putatively would improve specific vital rates (i.e., improve sex- and age-specific survival probability and fecundity rates). The authors applied status-quo conditions, and assumed no environmental or demographic stochasticity or future catastrophic events. Given these conditions and assumptions, and an initial population size of 88, the authors estimated the Southern Residents’ expected population size to be 84 in 10 years, 78 in 20 years, 71 in 30 years, and 68 in 35 years,. Only a small fraction of the projected populations exceeded the initial population size of 88 at 10 and 20 years, but at 35 years all projections were below 88 and some were as low as 55-60.

Projections of population size under demographic stochasticity showed that the probability of falling below 30 individuals was greater than zero at about 55 years in the future. However, inclusion of both environmental and demographic stochasticity increased the probability of falling below 30 individuals, which was greater than zero at 10 years, 50% at 47 years and approximately 80% at 100 years. Even without the inclusion of demographic stochasticity, there was a zero probability of Southern Residents reaching 120 individuals in 14 years and thus meeting the U.S. target to “downlist” the Southern

Residents from their endangered status (National Marine Fisheries Sevice 2008). Moreover, when the time horizon was fixed, rather than fixing the carrying capacity, there was an extinction risk of 49% in 100 years with an expected abundance of 15 individuals during this timeframe.

Robert C. Lacy -- Curriculum vitae 1

ROBERT C. LACY

Conservation, Education, & Training Office phone: (708) 688-8432 Chicago Zoological Society email: [email protected] Brookfield, Illinois 60513 USA http://www.vortex10.org

Home office: 192 Ocean View Road Phone and FAX: (207)434-2710 Jonesboro, Maine, USA email: [email protected]

Education B.A., summa cum laude, Biology, Wesleyan University 1977 Secondary school teaching certification for general science and biology M.A., Biology, Wesleyan University 1977 Thesis: Evolutionary strategies for temperature regulation in Mus musculus: A quantitative genetic approach Ph.D., Evolutionary Biology (minors: Genetics, Ecology), Cornell University, August1982 Thesis: Population biology of mycophagous Drosophilidae in the eastern U.S.

Positions Held 1985 to present Senior Conservation Scientist, Chicago Zoological Society 2003 to 2011 Chairman, IUCN Species Survival Commission (SSC) Conservation Breeding Specialist Group 1992 to 1993 Chairman, Dept. of Conservation Biology, Chicago Zoological Society 1982 to 1985 Assistant Professor of Biology, Franklin & Marshall College

Academic Appointments/Graduate Advisory Committees/Postdoctoral Advisees 1985 to present Chicago Zoological Society, Department of Conservation Biology (Supervised 4 post-doctoral research associates.) 1991 to present University of Chicago, Lecturer, Committee on Evolutionary Biology (Served on PhD advisory committees for 7 students.) 1999 to present University of Illinois, Chicago, Adjunct Professor, Department of Biology (Served on PhD advisory committees for 4 students.) various External committee member for graduate students at University of Illinois- Urbana, Univ of Maryland, Univ of Wisconsin-Milwaukee, Macquarie Univ, Univ of New South Wales, Monash Univ, South Dakota State Univ, Univ Missouri-St Louis, Univ Montana, Purdue Univ

Current Research Interests Interaction among genetic, demographic, and environmental causes of extinction Modeling the dynamics of linked systems affecting wildlife – including population biology, epidemiology, wildlife harvest, habitat fragmentation, and changes in human populations Genetic management of captive populations Robert C. Lacy -- Curriculum vitae 2

Inbreeding and outbreeding depression

Teaching Experience Franklin and Marshall College Genetics, Vertebrate Biology, Biosocial and Environmental Problems University of Chicago Conservation Biology graduate seminar Chicago Zoological Society Lectures on evolution and conservation Professional schools of the American Zoo and Aquarium Association Population Management (demography and genetics sections) Advanced Training Program in the Conservation of Biodiversity Program coordination, lectures, and mentor for biologists from tropical countries Escola Superior de Conservação Ambiental e Sustentabilidade (ESCAS, Brazil) Introduction to Conservation Decision-making Numerous other workshops on genetic analysis and population management taught to wildlife biologists, zoo managers, and conservation biologists

Professional Societies Association of Zoos and Aquariums American Genetic Association Society for Conservation Biology Society for the Study of Evolution

Professional Service Journal advisory boards: Zoo Biology, Conservation Genetics, International Zoo Yearbook Prioritization Task Force, IUCN Species Survival Commission (SSC) Conservation Breeding Specialist Group, IUCN SSC (Chair, 2003-2011) Recent activities include advising US Fish and Wildlife Service, state wildlife agencies, wildlife agencies of other nations (Australia, Canada, Spain, Brazil, Kenya, Indonesia, Malaysia, India, Chile, Ecuador, South Africa) and international conservation organizations on the management of Florida panther, whooping crane, Sumatran rhinoceros, lion tamarins, lion-tailed macaque, black rhinoceros, Iberian lynx, Humboldt penguin, African penguin, grizzly bear, lowland tapir, and many other species. Member of IUCN SSC Reintroduction Specialist Group Member of AZA Small Population Management Advisory Group Advisor to AZA Field Conservation Committee Conservation Fellow, St Louis Zoo WildCare Institute Robert C. Lacy -- Curriculum vitae 3

Honors and Grants Peirce Award for Excellence in the Sciences, Wesleyan University, 1977 Phi Beta Kappa, 1976 Sigma Xi, 1978 Predoctoral Fellowship. NSF, 1977 - 1980. Doctoral Dissertation - Research in Population Biology. NSF, 1979 - 1981, $4845. Faculty research grants. Franklin & Marshall College, 1982 - 1985. Studies of inbreeding depression in Peromyscus mice. Institute of Museum Services (IMS), 1985 - 1987, $22,775. Electrophoretic analysis of zoo populations. IMS, 1986 - 1988, $24,995. Studies of outbreeding depression in Peromyscus mice. IMS, 1987 - 1989, $25,000. Electrophoretic analyses of endangered species. IMS, 1988 - 1990, $25,000. Outstanding Service Awards, American Zoo and Aquarium Association, 1988, 1989, 2001, 2011 Chromosomal analysis of endangered species. IMS, 1989 - 1991, $101,347. Predictability of inbreeding depression in insular and mainland populations. NSF, 1991-1994, $182,683. Population Management 2000 software development. AZA Conservation Endowment Fund, 1999, $20,540. Biocomplexity: Models and meta-networks for interdisciplinary research in biodiversity risk assessment. NSF, 2000-2002, $98,000 (with P Nyhus, F Westley, P Miller, and G Ness) An experimental test of the effects of breeding strategies used in AZA conservation programs. AZA Conservation Endowment Fund, 2001, $42,926. Experimental tests of the effects of captive breeding of wildlife. IMLS, 2002-2005, $75,000. Pedigree reconstruction to sustain populations. IMLS, 2005-2007, $200,293 (with J. Dubach) Meta-models as an approach to understanding biocomplexity. Private donor to Chicago Zoological Society, 2006-2010, $100,000. President’s Award, Chicago Zoological Society, 2007. IUCN Species Survival Commission Chair’s Citation of Excellence Award, 2008. Linking behavioral types and animal "job performance" with population management in zoos. 2009 IMLS National Leadership Planning Grant, $22,535 (with J. Watters and D. Powell) Incorporating mate choice into breeding recommendations. 2009 IMLS National Leadership Planning Grant, $48,997 (with C. Asa and K. Traylor-Holzer) RCN: Using metamodels to enable transdisciplinary research for the study of dynamic biological systems under global change. NSF, 2012-2017, $490,905 (with H R Akcakaya, Stony Brook University). George Rabb Award for Conservation Innovation, IUCN Species Survival Commission, 2012 Ulysses S Seal Award for Innovation in Conservation, IUCN Conservation Breeding Specialist Group, 2012 LCP NRDA Dolphin Assessment, sub-contract with Industrial Economics on contract from NOAA. 2014-2015. $118,000 (co-PI with R. Wells). Building capacity in population modeling for species conservation. Chicago Board of Trade Endangered Species Fund, 2014, $3,000 Assessing conservation strategies for the Panamanian Golden frog. Chicago Board of Trade Robert C. Lacy -- Curriculum vitae 4

Endangered Species Fund, 2014, $4,250

Presentations and international workshop participation in 2010 Symposium on Threatened Species of the World. Presentation on “Metamodels for the integration of multiple sources of information into species risk assessments.” Caracas. IUCN Species Survival Committee (SSC) Steering Committee, Choroní, Venezuela. Association of Zoo and Aquarium Docents, annual conference, Chicago (invited presenter). Instituto de Pesquisas Ecológicas, Nazaré Paulista, Brazil (seminar speaker). CBSG Strategic Committee, Cologne, Germany (organizer) CBSG Annual Meeting, Germany (organizer) World Assoc. of Zoos and Aquariums (WAZA) Annual Conference, Germany (presenter) Tools for Species Conservation Strategic Planning, CBSG, Abruzzo, Italy.

Presentations and international workshop participation in 2011 CBSG Strategic Committee, Gland, Switzerland (convener) CBSG Annual Meeting, Prague (organizer) World Assoc. of Zoos and Aquariums (WAZA) Annual Conference, Prague (presenter) Association of Zoos and Aquariums (AZA) Annual Conference, session organizer Workshop on “Future of international studbooks”, WAZA, Gland, Switzerland Workshop on “Defining species for conservation breeding programs”, St Louis (organizer) Workshop on “Prioritization methods in species conservation”, Caracas University of Maine Colby College Akron Zoo Foundation

Presentations and international workshop participation in 2012 Workshop on population management, European Assoc of Zoos and Aquariums, Copenhagen CBSG Strategic Committee, Melbourne, Australia CBSG Annual Meeting, Melbourne, Australia (presenter) Workshop on “Use of gamete banks in breeding programs”, St Louis (co-organizer) Research Coordination Network: Metamodels for transdisciplinary research, Steering Committee PHVA for Yellow-Shouldered Blackbird (3 workshops), Puerto Rico (co-organizer) Workshop on metamodels to assess biocontrol of rabbits, S Australia Biosecurity, Adelaide Workshop on reintroduction of bison to Illinois, Nature Conservancy (presenter) Advanced short-course on pedigree analysis and management, Brookfield, IL (instructor) Workshop on integrating spatial models of animal movements into PVA, Colby College Purdue University

Presentations and international workshop participation in 2013 Workshop on using Vortex software for PVA, Chicago Zoological Society Dallas Zoo Foundation International Congress on Conservation Biology (presenter, and co-author on 3 other papers) Norwegian Polar Institute, Tromsø (several presentations while a visiting researcher) Robert C. Lacy -- Curriculum vitae 5

International Conference on Behaviour, Physiology and Genetics of Wildlife, Berlin (plenary speaker, contributed paper, and poster) European Association of Zoos and Aquariums, Edinburgh (presentation) Panamanian Golden Frog Species Conservation Strategic Planning workshop, El Valle, Panama

Presentations and international workshop participation in 2014 Course on Vortex 10 software, Chicago Zoological Society (instructor) North American Congress on Conservation Biology, Missoula, MT. Presented paper on “How to consider genetic depletion and rescue within Population Viability Analyses” Workshop on “PVA to Policy”, Longyearbyen, Norway. Presentation on “Linking models for integrated analysis of populations facing multiple, interacting threats”. Symposium on “Assessing vulnerability of flora and fauna in polar areas”, Tromsø, Norway. Presented paper on “Linking PVA models to explore the impacts of declining polar ice on interconnected species in the arctic ecosystem”. Workshop on “Molecular genetics for species management in zoos and aquaria”, Antwerp, Belgium. Presentation on “Some challenges in the use of molecular genetics in population management” Workshop on “Adaptive management for species conservation along the captive-wild spectrum”, Melbourne, Australia. Workshop on “Integrating models of animal social systems within PVA”, Blacksburg, VA. CBSG Strategic Committee, New Delhi, India CBSG Annual Meeting, New Delhi, India

Publications (most are available for downloading from www.vortex10.org/lacypubs.html ) Lacy, R.C., C.B. Lynch and G.R. Lynch. 1978. Developmental and adult acclimation effects of ambient temperature on temperature regulation of mice selected for high and low levels of nest-building. Journal of Comparative Physiology B 123:185-192. Lacy, R.C. 1978. Dynamics of t-alleles in Mus musculus populations: Review and speculation. The Biologist 60:41-67. Lacy, R.C. 1979. The adaptiveness of a rare male mating advantage under heterosis. Behavior Genetics 9:51-54. Lacy, R.C. and C.B. Lynch. 1979. Quantitative genetic analysis of temperature regulation in Mus musculus. I. Partitioning of variance. Genetics 91:743-753. Lacy, R.C. 1980. The evolution of eusociality in termites: A haplodiploid analogy? American Naturalist 116:449-451. Lacy, R.C. 1981. Taxonomic and distributional notes on some fungus-feeding North American Drosophila (Diptera, Drosophilidae). Entomological News 92:59-63. Lacy, R.C. 1982. Niche breadth and abundance as determinants of genetic variation in populations of mycophagous drosophilid flies (Diptera:Drosophilidae). Evolution 36:1265-1275. Lacy, R.C. 1983. Structure of genetic variation within and between populations of mycophagous Drosophila. Genetics 104:81-94. Lacy, R.C. and P.W. Sherman. 1983. Kin recognition by phenotype matching. American Robert C. Lacy -- Curriculum vitae 6

Naturalist 121:489-512. Lacy, R.C. 1984. Ecological and genetic responses to mycophagy in Drosophilidae (Diptera). Pages 286-301 in Q. Wheeler and M. Blackwell (eds.), Fungus/Insect Relationships: Perspectives in Ecology and Evolution. Columbia University Press, New York. Lacy, R.C. 1984. Predictability, toxicity, and trophic niche breadth in fungus-feeding Drosophilidae (Diptera). Ecological Entomology 9:43-54. Lacy, R.C. 1984. The evolution of termite eusociality: Reply to Leinaas. American Naturalist 123:876-878. Hayssen, V. and R.C. Lacy. 1985. Basal metabolic rates in mammals: Taxonomic differences in the allometry of BMR and body mass. Comparative Biochemistry and Physiology 81A:741-754. Hayssen, V., R.C. Lacy and P.J. Parker. 1985. Metatherian reproduction: Transitional or transcending? American Naturalist 126:617-632. Lacy, R.C. 1985. Evidence that group selection counters the evolution of sexual dimorphism. Evolutionary Theory 7:173-177. Lacy, R.C. 1985. Some genetic considerations for the management of captive populations suggested by computer simulations. AAZPA 1985 Annual Proceedings, 627-630. Lacy, R.C. and C.E. Bock. 1986. The correlation of range size and local abundance of some North American birds. Ecology 67:258-260. Lacy, R.C. 1987. Loss of genetic diversity from managed populations: Interacting effects of drift, mutation, immigration, selection, and population subdivision. Conservation Biology 1:143-158. Lacy, R.C. 1987. Further genetic and demographic analyses of small rhino populations. Pachyderm (Newsletter of the African Elephant and Rhino Specialist Group) No. 9, pp. 16-19. Lacy, R.C. 1988. A report on population genetics in conservation. Conservation Biology 2:245-247. Lacy, R.C. 1988. Genetic variability in captive stocks: Assessing past loss, present status, and future outlook. AAZPA 1988 Annual Proceedings 113-121. Lacy, R.C., M.L. Foster, and the Primate Department Staff. 1988. Determination of pedigrees and taxa of primates by protein electrophoresis. International Zoo Yearbook 27:159-168. Lacy, R.C. 1988. Conservation genetics at Brookfield Zoo and the Brookfield-Melbourne genetics research programme. Bulletin of Zoo Management 26:27-29. Lacy, R.C. and T.W. Clark. 1989. Genetic variability in black-footed ferret populations: Past, present, and future. Pages 83-103 in U.S. Seal, E.T. Thorne, M.A. Bogan, and S.H. Anderson (eds.), Conservation Biology and the Black-Footed Ferret. Yale University Press, New Haven. Lacy, R.C. 1989. How many pairs are needed on the ark? Bison 4:24-28. Lacy, R.C. 1989. Analysis of founder representation in pedigrees: Founder equivalents and founder genome equivalents. Zoo Biology 8:111-124. Lacy, R.C, Flesness, N.R., and Seal, U.S. 1989. Puerto Rican parrot population viability analysis. Report to the U.S. Fish and Wildlife Service. IUCN SSC Captive Breeding Specialist Group, Apple Valley, Minnesota. Robert C. Lacy -- Curriculum vitae 7

Seal, U.S. and R.C. Lacy. 1989. Florida panther population viability analysis. Report to the U.S. Fish and Wildlife Service. IUCN SSC Captive Breeding Specialist Group, Apple Valley, Minnesota. Paine, F.L., J.D. Miller, G. Crawshaw, B. Johnson, R. Lacy, C.F. Smith III, and P.J. Tolson. 1990. Status of the Puerto Rican crested toad. International Zoo Yearbook 28:53-58. Maguire, L.A. and R.C. Lacy. 1990. Allocating scarce resources for conservation of endangered subspecies: Partitioning zoo space for tigers. Conservation Biology 4:157-166. Brewer, B.A., R.C. Lacy, M.L. Foster, and G. Alaks. 1990. Inbreeding depression in insular and central populations of Peromyscus mice. Journal of Heredity 81:257-266. Maguire, L.A., R.C. Lacy, R.J. Begg, and T.W. Clark. 1990. An analysis of alternative strategies for recovering the eastern barred bandicoot in Victoria. Pages 147-164 in T.W. Clark and J.H. Seebeck (eds.), The Management and Conservation of Small Populations. Chicago Zoological Society, Brookfield, Illinois. Lacy, R.C. and T.W. Clark. 1990. Population viability assessment of the eastern barred bandicoot in Victoria. Pages 131-146 in T.W. Clark and J.H. Seebeck (eds.), The Management and Conservation of Small Populations. Chicago Zoological Society. George, G.G., J. Dixon, G. Challis, and R.C. Lacy. 1990. Taxonomy and palaeontology of the eastern barred bandicoot. Pages 33-46 in T.W. Clark and J.H. Seebeck (eds.), The Management and Conservation of Small Populations. Chicago Zoological Society. Seal, U.S. and R.C. Lacy. 1990. Florida Key Deer (Odocoileus virginianus clavium) population viability assessment. Report to the U.S. Fish and Wildlife Service. IUCN SSC Captive Breeding Specialist Group, Apple Valley, Minnesota. Lindenmayer, D.B., V.C. Thomas, R.C. Lacy, and T.W. Clark. 1991. Population Viability Analysis (PVA): The concept and its applications, with a case study of Leadbeater's Possum, Gymnobelideus leadbeateri McCoy. Report to the Forest and Timber Inquiry (Resource Assessment Commission), Consultancy Series No. FTC91/18, Canberra, Australia. 170 pp. Clark, T.W., G.N. Backhouse, and R.C. Lacy. 1991. The population viability assessment workshop: A tool for threatened species management. Endangered Species Update 8:1-5. Clark, T.W., G.N. Backhouse, and R.C. Lacy. 1991. Report of a workshop on population viability assessment as a tool for threatened species management and conservation. Australian Zoologist 27:28-35. Lacy, R.C. 1991. Zoos and the surplus problem: An alternative solution. Zoo Biology 10:293-297. Johnston, L.A. and R.C. Lacy. 1991. Utilization of sperm banks to maintain genetic diversity in captive populations of wild cattle. Pages 107-118 in D.L. Armstrong and T.S. Groves (eds.), Wild Cattle Symposium Proceedings. Henry Doorly Zoo, Omaha, Nebraska. Seal, U.S., R.C. Lacy, K. Medley, R. Seal, and T.J. Foose. 1991. Tana River Primate Reserve Conservation Assessment Workshop Report. IUCN SSC Captive Breeding Specialist Group, Apple Valley, Minnesota. Mirande, C., R. Lacy, and U. Seal. 1991. Whooping crane (Grus americana) conservation viability assessment workshop report. IUCN SSC Captive Breeding Specialist Group, Apple Valley, Minnesota. Robert C. Lacy -- Curriculum vitae 8

Foose, T.J., R.C. Lacy, R. Brett, and U.S. Seal. 1991. Kenya black rhinoceros metapopulation workshop report. IUCN SSC Captive Breeding Specialist Group, Apple Valley, Minnesota. Lacy, R.C. 1992. The effects of inbreeding on isolated populations: Are minimum viable population sizes predictable? Pages 277-296 in P.L. Fiedler and S.K. Jain (eds.), Conservation Biology: The Theory and Practice of Nature Conservation, Preservation and Management. Chapman and Hall, New York. Lacy, R.C. and T. Kreeger. 1992. VORTEX Users Manual. A stochastic simulation of the extinction process. IUCN SSC Captive Breeding Specialist Group, Apple Valley, Minnesota. Seal, U.S., R.C. Lacy, et al. 1992. Genetic management strategies and population viability of the Florida panther (Felis concolor coryi). Report to the U.S. Fish and Wildlife Service. IUCN SSC Captive Breeding Specialist Group, Apple Valley, Minnesota. Ellis, S., K. Hughes, C. Kuehler, R. Lacy, and U. Seal. 1992. `Alala, Akohekohe, and Palila Population and Habitat Viability Assessment Reports. IUCN SSC Captive Breeding Specialist Group, Apple Valley, Minnesota. Ellis, S., C. Kuehler, R. Lacy, K. Hughes, and U. Seal. 1992. Hawai`ian forest birds conservation assessment and management plan. IUCN SSC Captive Breeding Specialist Group, Apple Valley, Minnesota. Lacy, R.C. 1993. Impacts of inbreeding in natural and captive populations of vertebrates: Implications for conservation. Perspectives in Biology and Medicine 36:480-496. Lacy, R.C. 1993. VORTEX: A computer simulation model for Population Viability Analysis. Wildlife Research 20:45-65. Lacy, R.C. 1993. GENES: A computer program for the analysis of pedigrees and genetic management of populations. Chicago Zoological Society, Brookfield, Illinois. Lacy, R.C. and T.W. Clark. 1993. Simulation modeling of American marten populations: Vulnerability to extinction. Great Basin Naturalist 53:282-292. Lacy, R.C., A.M. Petric, and M. Warneke. 1993. Inbreeding and outbreeding depression in captive populations of wild species. Pages 352-374 in N.W. Thornhill (ed.), The Natural History of Inbreeding and Outbreeding. University of Chicago Press. Lindenmayer, D.B., R.C. Lacy, V.C. Thomas, and T.W. Clark. 1993. Predictions of the impacts of changes in population size and environmental variability on Leadbeater's Possum, Gymnobelideus leadbeateri McCoy (Marsupialia: Petauridae) using Population Viability Analysis: an application of the computer program VORTEX. Wildlife Research 20:67-86. Lindenmayer, D.B., T.W. Clark, R.C. Lacy, and V.C. Thomas. 1993. Population viability analysis as a tool in wildlife conservation policy: With reference to Australia. Environmental Management 17:745-758. Lindenmayer, D.B. and R.C. Lacy. 1993. Using a computer simulation package for PVA to model the dynamics of sub-divided populations: An example using hypothetical metapopulations of the mountain brushtail possum. International Congress on Modelling and Simulation. Proceedings. 2:615-620. Lacy, R.C. 1993/1994. What is Population (and Habitat) Viability Analysis? Primate Conservation 14/15:27-33. Robert C. Lacy -- Curriculum vitae 9

Lacy, R.C. 1994. Review of Hartl, G.B. and Markowski, J. (eds.) Ecological genetics in mammals. Journal of Mammalogy 75:1090-1093. Jiménez, J.A., K.A. Hughes, G. Alaks, L. Graham, and R.C. Lacy. 1994. An experimental study of inbreeding depression in a natural habitat. Science 266:271-273. Lacy, R.C. 1994. Managing genetic diversity in captive populations of animals. Pages 63-89 in M.L. Bowles and C.J. Whelan (eds.), Restoration and Recovery of Endangered Plants and Animals. Cambridge University Press, Cambridge. Lacy, R.C. 1995. A sibling is as valuable as an offspring: Reply to Xia. American Naturalist 145:480-482. Lindenmayer, D.B. and R.C. Lacy. 1995. Metapopulation viability of Leadbeater's Possum, Gymnobelideus leadbeateri, in fragmented old-growth forests. Ecological Applications 5:164-182. Lindenmayer, D.B. and R.C. Lacy. 1995. Metapopulation viability of arboreal marsupials in fragmented old-growth forests: comparison among species. Ecological Applications 5:183-199. Lacy, R.C., J.D. Ballou, F. Princée, A. Starfield, and E. Thompson. 1995. Pedigree analysis. Pages 57-75 in J.D. Ballou, M. Gilpin, and T.J. Foose (eds.), Population Management for Survival & Recovery. Analytical Methods and Strategies in Small Population Conservation. Columbia University Press, New York. Ballou, J.D. and R.C. Lacy. 1995. Identifying genetically important individuals for management of genetic diversity in pedigreed populations. Pages 76-111 in J.D. Ballou, M. Gilpin, and T.J. Foose (eds.), Population Management for Survival & Recovery. Analytical Methods and Strategies in Small Population Conservation. Columbia University Press, New York. Johnston, L.A. and R.C. Lacy. 1995. Genome resource banking for species conservation: Selection of sperm donors. Cryobiology 32:68-77. Lacy, R.C. 1995. Culling surplus animals for population management. Pages 187-194 in B.G. Norton, M. Hutchins, E.F. Stevens, and T.L. Maple (eds.) Ethics on the ark: Zoos, animal welfare, and wildlife conservation. Smithsonian Institution Press, Washington, DC. Lindenmayer, D.B. and R.C. Lacy. 1995. A simulation study of the impacts of population subdivision on the mountain brushtail possum, Trichosurus caninus Ogilby (Phalangeridae: Marsupialia), in south-eastern Australia. I. Demographic stability and population persistence. Biological Conservation 73:119-129. Lacy, R.C. and D.B. Lindenmayer. 1995. A simulation study of the impacts of population subdivision on the mountain brushtail possum, Trichosurus caninus Ogilby (Phalangeridae: Marsupialia), in south-eastern Australia. II. Loss of genetic variation within and between subpopulations. Biological Conservation 73:131-142. Lindenmayer, D.B., M.A. Burgman, H.R. Akçakaya, R.C. Lacy, and H.P. Possingham. 1995. A review of the generic computer programs ALEX, RAMAS/space and VORTEX for modelling the viability of wildlife metapopulations. Ecological Modelling 82:161- 174. Lacy, R.C. 1995. Clarification of genetic terms and their use in the management of captive Robert C. Lacy -- Curriculum vitae 10

populations. Zoo Biology 14:565-577. Lacy, R.C. 1995. Conservation geneticists make their case. (Book review.) Ecology 76:1684-1685. Lacy, R.C., K.A. Hughes, and P.S. Miller. 1995. VORTEX Version 7 users manual. A stochastic simulation of the simulation process. IUCN/SSC Conservation Breeding Specialist Group. Apple Valley, Minnesota. Altmann, J., S.C. Alberts, S.A. Haines, J. Dubach, P. Muruthi, T. Coote, E. Geffen, D.J. Cheesman, R.S. Mututua, S.N. Saiyalel, R.K. Wayne, R.C. Lacy, and M.W. Bruford. 1996. Behavior predicts genetic structure in a wild primate group. Proc. Nat. Acad. Sci. USA 93:5797-5801. Lacy, R. 1996. Further population modelling of northern white rhinoceros under various management scenarios. Appendix 3 in T.J. Foose (ed.). Summary – Northern White Rhinoceros Conservation Strategy Workshop. International Rhino Foundation, Cumberland, Ohio. Lacy, R.C. and B.E. Horner. 1996. Effects of inbreeding on skeletal development of Rattus villosissimus. Journal of Heredity 87:277-287. Lacy, R.C., G. Alaks, and A. Walsh. 1996. Hierarchical analysis of inbreeding depression in Peromyscus polionotus. Evolution 50:2187-2200. Hedrick, P.W., R.C. Lacy, F.W. Allendorf, and M.E. Soulé. 1996. Directions in conservation biology: Comments on Caughley. Conservation Biology 10:1312-1320. Lacy, R.C. 1996. Review of J.C. Avise and J.L. Hamrick (eds.). Conservation Genetics: Case histories from nature. Quarterly Review of Biology 71:566. Lacy, R.C. 1997. Importance of genetic variation to the viability of mammalian populations. Journal of Mammalogy 78:320-335. Lacy, R.C. and B.E. Horner. 1997. Effects of inbreeding on reproduction and sex ratio of Rattus villosissimus. Journal of Mammalogy 78:877-887. Ballou, J.D., R.C. Lacy, D. Kleiman, A. Rylands, and S. Ellis. 1998. Leontopithecus II. The second Population and Habitat Viability Assessment for Lion Tamarins (Leontopithecus). IUCN/SSC Conservation Breeding Specialist Group, Apple Valley, MN. Lindenmayer, D.B., R.C. Lacy, and K.L. Viggers. 1998. Modelling survival and capture probabilities of the mountain brushtail possum (Trichosurus caninus) in the forests of south-eastern Australia using trap-recapture data. Journal of Zoology 245:1-13. Lacy, R.C. and J.D. Ballou. 1998. Effectiveness of selection in reducing the genetic load in populations of Peromyscus polionotus during generations of inbreeding. Evolution 52:900-909. Miller, P.S. and R.C. Lacy. 1999. VORTEX Version 8 users manual. A stochastic simulation of the simulation process. IUCN/SSC Conservation Breeding Specialist Group. Apple Valley, Minnesota. Brook, B.W., J.R. Cannon, R.C. Lacy, C. Mirande, and R. Frankham. 1999. Comparison of the population viability analysis packages GAPPS, INMAT, RAMAS and VORTEX for the whooping crane (Grus americana). Animal Conservation 2:23-31. Araya, B., D. Garland, G. Espinoza, A. Sanhuesa, A. Simeone, A. Teare, C. Zavalaga, R. Lacy, Robert C. Lacy -- Curriculum vitae 11

and S. Ellis. (eds.) 1999. Taller Análisis de la Viabilidad del Hábitat y de la Población del Pinguino Humboldt (Spheniscus humboldti). Informe final. IUCN/SSC Conservation Breeding Specialist Group, Apple Valley, MN. Cooper, J., D. Oschadleus, L. Shannon, M. Thornton, P. Whittington, R. Lacy, and S. Ellis (eds.) 1999. African Penguin Population and Habitat Viability Assessment. IUCN/SSC Conservation Breeding Specialist Group, Apple Valley, MN. Lindenmayer, D.B., R.C. Lacy, H. Tyndale-Biscoe, A.C. Taylor, K.L. Viggers, and M.L. Pope. 1999. Integrating demographic and genetic studies of the Greater Glider Petauroides volans in fragmented forests: predicting movement patterns and rates for future testing. Pacific Conservation Biology 5:2-8. Ellis, S., R.C. Lacy, S. Kennedy-Stoskopf, D.E. Wildt, J. Shillcox, O. Byers, and U.S. Seal. 1999. Florida panther population and habitat viability assessment and genetics workshop report. IUCN/SSC Conservation Breeding Specialist Group, Apple Valley, MN. Lindenmayer, D.B., R.C. Lacy, and M.L. Pope. 2000. Testing a simulation model for Population Viability Analysis. Ecological Applications 10:580-597. Lacy, R.C. 2000. Management of limited animal populations. Pages 75-93 in D. Duffield and T. Robeck (eds.). Bottlenose dolphin reproduction workshop. Report. AZA Marine Mammal Taxon Advisory Group, Silver Spring, MD. Pergams, O.R.W., R.C. Lacy, and M.V. Ashley. 2000. Conservation and management of Anacapa Island Deer Mice. Conservation Biology 14:819-832. Penn, A.M., W.B. Sherwin, G. Gordon, D. Lunney, A. Melzer, and R.C. Lacy. 2000. Demographic forecasting in koala conservation. Conservation Biology 14:629-638. Lacy, R.C. 2000. Should we select genetic alleles in our conservation breeding programs? Zoo Biology 19:279-282. Land, E.D., and R.C. Lacy. 2000. Introgression level achieved through Florida Panther genetic restoration. Endangered Species Update 17:100-105. Lacy, R.C. 2000. Considering threats to the viability of small populations. Ecological Bulletins 48:39-51. Lacy, R.C. 2000. Structure of the VORTEX simulation model for population viability analysis. Ecological Bulletins 48:191-203. Pollak, J. P., R. C. Lacy and J. D. Ballou. 2000. Population Management 2000, version 1.175. Chicago Zoological Society, Brookfield, IL. Lacy, R.C., and J.D. Ballou. 2001. Population Management 2000 User’s Manual. Chicago Zoological Society, Brookfield, IL. Lindenmayer, D.B., and R.C. Lacy. 2002. Small mammals, habitat patches and PVA models: a field test of model predictive ability. Biological Conservation 103:247-265. Lacy, R.C., and P.S. Miller. 2002. Incorporating human populations and activities into population viability analysis. Pages 490-510 in S.R. Beissinger and D.R. McCullough (eds.), Population Viability Analysis. University of Chicago Press, Chicago. Maehr, D.S., R.C. Lacy, E.D. Land, O.L. Bass, and T.S. Hoctor. 2002. Evolution of Population Viability Assessments for the Florida panther: A multiperspective approach. Pages 284- 311 in S.R. Beissinger and D.R. McCullough (eds.), Population Viability Analysis. University of Chicago Press, Chicago. Robert C. Lacy -- Curriculum vitae 12

Hosack, D.A., P.S. Miller, J.J. Hervert, and R.C. Lacy. 2002. A population viability analysis for the endangered Sonoran pronghorn, Antilocapra americana sonoriensis. Mammalia 66:207-229. Nyhus, P.J., F.R. Westley, R.C. Lacy, and P.S. Miller. 2002. A role for natural resource social science in biodiversity risk assessment. Society and Natural Resources 15:923-932. Jones, K.L., T.C. Glenn, R.C. Lacy, J.R. Pierce, N. Unruh, C.M. Mirande, F. Chavez-Ramirez. 2002. Refining the whooping crane studbook by incorporating microsatellite DNA and leg banding analyses. Conservation Biology 16:789-799 Maehr, D.S., and R. C. Lacy. 2002. Avoiding the lurking pitfalls in Florida panther recovery. Wildlife Society Bulletin 30:971-978. Ryan, K.K., R.C. Lacy, and S.W. Margulis. 2002. Impacts of inbreeding on components of reproductive success. Pages 82-96 in: W. V. Holt, A. R. Pickard, J. C. Rodger, and D. E. Wildt, eds. Reproductive Science and Integrated Conservation. Cambridge University Press, Cambridge, UK. Ryan, K.K., and R.C. Lacy. 2003. Monogamous male mice bias behaviour towards females according to very small differences in kinship. Animal Behaviour 65: 379-384. Lindenmayer, D. B., H. P. Possingham, R. C. Lacy, M. A. McCarthy and M. L. Pope. 2003. How accurate are population models? Lessons from landscape-scale tests in a fragmented system. Ecology Letters 6:41-47. Miller, P.S., and R.C. Lacy. 2003. VORTEX: A Stochastic Simulation of the Extinction Process. Version 9 User’s Manual. Conservation Breeding Specialist Group (SSC/IUCN), Apple Valley, Minnesota. Miller, P.S., and R.C. Lacy. 2003. Integrating the human dimension into endangered species risk assessment. Pages 41-63 in: F.R. Westley and P.S. Miller, eds. Experiments in Consilience: Integrating Social and Scientific Responses to Save Endangered Species. Island Press, Washington, DC. Miller, P.S., and R.C. Lacy. 2003. Metamodels as a tool for risk assessment. Pages 333-351 in: F.R. Westley and P.S. Miller, eds. Experiments in Consilience: Integrating Social and Scientific Responses to Save Endangered Species. Island Press, Washington, DC. Westley, F., P.S. Miller, and R.C. Lacy. 2003. Far from land: Further explorations in consilience. Pages 352-361 in: F.R. Westley and P.S. Miller, eds. Experiments in Consilience: Integrating Social and Scientific Responses to Save Endangered Species. Island Press, Washington, DC. Singleton, I., S. Wich, S. Husson, S. Stephens, S. Utami Atmoko, M. Leighton, N. Rosen, K. Traylor-Holzer, R. Lacy, and O. Byers (eds.). 2004. Orangutan Population and Habitat Viability Assessment: Final Report. IUCN/SSC Conservation Breeding Specialist Group, Apple Valley, MN. Lacy, R.C., and A. Vargas. 2004. Informe sobre la gestión genética y demográfica del programa de cría para la conservación del lince ibérico: escenarios, conclusiones y recomendaciones. Ministerio de Medio Ambiente, Madrid, Spain. Heinsohn, R., R. C. Lacy, D. B. Lindenmayer, H. Marsh, D. Kwan, and I.R. Lawler. 2004. Unsustainable harvest of dugongs in Torres Strait and Cape York (Australia) waters: two case studies using population viability analysis. Animal Conservation 7:417-425. Robert C. Lacy -- Curriculum vitae 13

Lacy, R.C., M. Borbat, and J.P. Pollak. 2005. VORTEX: A Stochastic Simulation of the Extinction Process. Version 9.57. Chicago Zoological Society, Brookfield, Illinois, USA. Margulis, S.W., M. Nabong, G. Alaks, A. Walsh, and R.C. Lacy. 2005. Effects of early experience on subsequent parental behaviour and reproductive success in oldfield mice, Peromyscus polionotus. Animal Behaviour 69:627-634. Traylor-Holzer, K., R. Lacy, D. Reed, and O. Byers (eds.). 2005. Alabama Beach Mouse Population and Habitat Viability Assessment: Final Report. IUCN/SSC Conservation Breeding Specialist Group, Apple Valley, MN. Mendelson, J.R. III, et al. (50 co-authors, including R.C. Lacy). 2006. Confronting amphibian declines and extinctions. Science 313:48. Mendelson, J.R. III, et al. (18 co-authors, including R.C. Lacy). 2006. Responding to amphibian loss. Response. Science 314:1541-1542. Matamoros, Y., H. Vargas, R. C. Lacy, O. Byers, E. Travis, G. Montoya. (Editores). 2006. Taller para Anàlisisde Viabilidad de Poblaciòny Hàbitatpara el Pingüino de Galápagos. Informe Final. Parque Nacional Galápagos, Puerto Ayora, Santa Cruz, Galápagos, Ecuador. 8-11 de febrero, 2005. Zippel, K., R. Lacy, and O. Byers (eds.) 2006. CBSG/WAZA Amphibian Ex Situ Conservation Planning Workshop Final Report. IUCN/SSC Conservation Breeding Specialist Group, Appl e V al l ey, M N. Vargas, F.H., R.C. Lacy, P.J. Johnson, A. Steinfurth, R.J.M. Crawford, P.D. Boersma, and D.W. MacDonald. 2007. Modelling the effect of El Niño on the persistence of small populations: The Galápagos penguin as a case study. Biological Conservation 137:138- 148. Miller, P.S., F.R. Westley, A.P. Byers, and R.C. Lacy. 2007. An experiment in managing the human animal: The PHVA process and its role in conservation decision-making. Pages 173-188 in T.S. Stoinski, H.D. Steklis, and P.T. Mehlman. Conservation in the 21st century: Gorillas as a case study. Springer, New York. Nyhus, P.J., R. Lacy, F.R. Westley, P.S. Miller, H. Vredenburg, P. Paquet, and J. Pollak. 2007. Tackling biocomplexity and meta-models for species risk assessment. Ecology & Society 12: 31 [online] URL: http://www.ecologyandsociety.org/vol12/iss1/art31/ Lacy, R.C. 2007. Understanding inbreeding depression: 20 years of experiments with Peromyscus mice. Pages 327-329 in F.W. Allendorf and G. Luikart. Conservation and the Genetics of Populations. Wiley-Blackwell, New York. Pergams, O.R.W. and R.C. Lacy. 2008. Rapid morphological and genetic change in Chicago-area Peromyscus. Molecular Ecology 17:450-463. Dasmahapatra, K.K., R.C. Lacy, and W. Amos. 2008. Estimating levels of inbreeding using AFLP markers. Heredity 100:286-295. Wich, S.A., E. Meijaard, A.J. Marshall, S. Husson, M. Ancrenaz, R.C. Lacy, C.P. van Schaik, J. Sugardjito, T. Simorangkir, K. Traylor-Holzer, M. Doughty, J. Supriatna, R. Dennis, M. Gumal, C.D. Knott, and I. Singleton. 2008. Distribution and conservation status of the orang-utan (Pongo spp.) on Borneo and Sumatra: How many remain? Oryx 42:329-339. Rudnick, J.A., and R.C. Lacy. 2008. The impact of assumptions about founder relationships on Robert C. Lacy -- Curriculum vitae 14

the effectiveness of captive breeding strategies. Conservation Genetics 9:1439-1450. Lacy, R.C. 2008. Conservation breeding – a global view. Pages 381-398 in B.R. Sharma, N. Akhtar, B.K. Gupta (eds.). India’s Conservation Breeding Initiative, Central Zoo Authority, New Delhi. Lacy, R.C. 2008. Conservation breeding – challenges and protocols. Pages 263-289 in B.R. Sharma, N. Akhtar, B.K. Gupta (eds.). India’s Conservation Breeding Initiative, Central Zoo Authority, New Delhi. IUCN/SSC. 2008. Strategic Planning for Species Conservation: A Handbook. Version 1.0. Gland, Switzerland: IUCN Species Survival Commission. 104pp. (R Lacy was one of the writers and co-editors, and chaired the task force that developed the handbook.) Lacy, R.C. 2009. Stopping evolution: Genetic management of captive populations. Pages 58-81 in: G. Amato, R. DeSalle, O.A. Ryder, and H.C. Rosenbaum. Conservation genetics in the age of genomics. Columbia University Press, New York. Lacy, R.C. and R. Wells. 2009. Population Viability Analysis (PVA). Pages 25-29 in R.R. Reeves, and R.L. Brownell, Jr., eds. Indo-Pacific bottlenose dolphin assessment workshop report: Solomon Islands case study of Tursiops aduncus. Occasional Paper of the Species Survival Commission, No. 40, IUCN, Gland, Switzerland. 53 pp. Marshall, A.J., R. Lacy, M. Ancrenaz, O. Byers, S.J. Husson, M. Leighton, E. Meijaard, N. Rosen, I. Singleton, S. Stephens, K. Traylor-Holzer, S.S.U. Atmoko, C.P. van Schaik, and S.A. Wich. 2009. Orangutan population biology, life history, and conservation. Pages 311-326 in: S.A. Wich, S.S.I. Atmoko, T.M. Setia, and C.P. van Schaik, eds. Orangutans. Oxford University Press, Oxford, UK. Ivy, J.A., A, Miller, R.C. Lacy, and J. A. DeWoody. 2009. Methods and prospects for using molecular data in captive breeding programs: an empirical example using parma wallabies (Macropus parma). Journal of Heredity 100:441-454. Leus, K., and R.C. Lacy. 2009. Genetic and demographic management of conservation breeding programs oriented towards reintroduction. Pages 74-84 in: A. Vargas, C. Breitenmoser, and U. Breitenmoser (eds.) Iberian Lynx Ex Situ Conservation: An Interdisciplinary Approach. Fundación Biodiversidad / IUCN Cat Specialist Group, Madrid. Ivy, J.A., and R.C. Lacy. 2010. Using molecular methods to improve the genetic management of captive breeding programs for threatened species. Pages 267-295 in: J.A. DeWoody, J.W. Bickham, C.H. Michler, K.M. Nicols, O.E. Rhodes, and K.E. Woeste, eds. Molecular Approaches in Natural Resource Conservation and Management. Cambridge University Press, Cambridge. Malo, A.F., F. Martinez-Pastor, G. Alaks, J. Dubach, and R.C. Lacy. 2010. Effects of genetic captive-breeding protocols on sperm quality and fertility in the white-footed mouse. Biology of Reproduction 83:540-548. Ballou, J.D., R.C. Lacy, and J.P. Pollak. 2010. PMx: software for demographic and genetic analysis and management of pedigreed populations. Chicago Zoological Society, Brookfield, Illinois, USA. Lacy, R.C. 2011. Re-thinking ex situ vs. in situ species conservation. Pages 25-29 in: G. Dick, Robert C. Lacy -- Curriculum vitae 15

ed. Biodiversity is Life. Proceedings of the 65th Annual Conference of the World Association of Zoos and Aquariums. WAZA, Gland, Switzerland. Asa, C.S., K. Traylor-Holzer, and R.C. Lacy. 2011. Can conservation-breeding programmes be improved by incorporating mate choice? International Zoo Yearbook 45:203-212. Leus, K., K. Traylor-Holzer, and R.C. Lacy. 2011. Genetic and demographic population management in zoos and aquariums: recent developments, future challenges and opportunities for scientific research. International Zoo Yearbook 45:213-225. Asa, C.S., K. Traylor-Holzer, and R.C. Lacy. 2011. Mate choice as a potential tool to increase population sustainability. WAZA Magazine 12: 23-25. Baker, A.M., R.C. Lacy, K. Leus, and K. Traylor-Holzer. 2011. Intensive management of populations for conservation. WAZA Magazine 12: 40-43. Frankham, R., J.D. Ballou, M.D. Eldridge, R.C. Lacy, K. Ralls, M.R. Dudash, and C.R. Fenster. 2011. Predicting the probability of outbreeding depression: critical information for managing fragmented populations. Conservation Biology 25:465-475. Jamieson, I.G., and R.C. Lacy. 2012. Managing genetic issues in reintroduction biology. Pages 441-475 in: J.G. Ewen, D.P. Armstrong, K.A. Parker, & P.J. Seddon, eds. Reintroduction Biology: Integrating Science and Management. Wiley-Blackwell, Oxford, UK. Lacy, R.C., J.D. Ballou, & J.P. Pollak. 2012. PMx: Software package for demographic and genetic analysis and management of pedigreed populations. Methods in Ecology & Evolution 3:433-437. Bradshaw, C.J.A., C.R. McMahon, P.S. Miller, R.C. Lacy, M.J. Watts, M.L. Verant, J.P. Pollak, D.A. Fordham, T.A.A. Prowse, and B.W. Brook. 2012. Novel coupling of individual- based epidemiological and demographic models predicts realistic dynamics of tuberculosis in alien buffalo. Journal of Applied Ecology 49:268-277. Ivy, J.A., and R.C. Lacy. 2012. A comparison of strategies for selecting breeding pairs to maximize genetic diversity retention in managed populations. Journal of Heredity 103:186-196. Lacy, R.C. 2012. Extending pedigree analysis for uncertain parentage and diverse breeding systems. Journal of Heredity 103:197-205. Lacy, R.C., and C. A. Beuchat. 2012. M anagi ng the geneti c heal th of the Basenj i . The Basenj i May/June 2012:10-13. Desbiez, A., K. Traylor-Holzer, R. Lacy, et al. 2012. Population Viability Analysis of jaguar populations in Brazil. In: Jaguar in Brazil. CATnews (Special Issue) 7:35-37. Frankham, R., J.D. Ballou, M.R. Dudash, M.D.B. Eldridge, C.B. Fenster, R.C. Lacy, J.R. Mendelson III, I.J. Porton, K. Ralls, and O.A. Ryder. 2012. Implications of different species concepts for conserving biodiversity. Biological Conservation 153:25-31. Lacy, R.C., J.P. Pollak, P.S. Miller, L. Hungerford, and P. Bright. 2012. Outbreak version2.0. IUCN SSC Conservation Breeding Specialist Group. Apple Valley, MN, USA. Lacy, R.C. 2013. Understanding inbreeding depression: 25 years of experiments with Peromyscus mice. Pages 491-492 in F.W. Allendorf, G. Luikart, and S.N. Aiken. Conservation and the genetics of populations. 2nd ed. John Wiley & Sons, Oxford, UK. Lacy, R.C. 2013. Achieving true sustainability of zoo populations. Zoo Biology 32:19-26. Lacy, R.C. and G. Alaks. 2013. Effects of inbreeding on skeletal size and fluctuating asymmetry Robert C. Lacy -- Curriculum vitae 16

of Peromyscus polionotus mice. Zoo Biology 32:125-133. Prowse, T.A.A., C.N. Johnson, R.C. Lacy, C.J.A. Bradshaw, J.P. Pollak, M.J. Watts, and B.W. Brook. 2013. No need for disease: testing extinction hypotheses for the thylacine using multi-species metamodels. Journal of Animal Ecology 82:355-364. Meile, R., R.C. Lacy, F.H. Vargas, and P.G. Parker. 2013. Modeling the potential effects of Plasmodium infection on the Galapagos penguin (Spheniscus mendiculus). Auk 130: 440- 448. Lacy, R.C., K. Traylor-Holzer, and J.D. Ballou. 2013. Managing for true sustainability of species. WAZA Magazine 14:10-14. Lacy, R.C., G. Alaks, and A. Walsh. 2013. Evolution of Peromyscus leucopus mice in response to a captive environment. PLoS ONE 8(8): e72452. doi:10.1371/journal.pone.0072452. Lacy, R.C., P.S. Miller, P.J. Nyhus, J.P Pollak, B.E. Raboy, and S. Zeigler. 2013. Metamodels for transdisciplinary analysis of population dynamics. PLoS ONE 8(12): e84211. doi:10.1371/journal.pone.0084211. Pollak, J.P., R.C. Lacy. 2013. MetaModel Manager. Version 1.0. Chicago Zoological Society, Brookfield, Illinois. Available: http://www.vortex10.org/MeMoMa.aspx. Raboy, B.E., and R.C. Lacy. 2013. MetaModel Manager user’s manual. Chicago Zoological Society, Brookfield, Illinois. Available: http://www.vortex10.org/MeMoMa.aspx. Byers, O., and R.C. Lacy. 2014. From captivity to conservation: Conservation Breeding Specialist Group and the evolving role of zoos. Pages 107-111 in G. McGregor-Reid and G. Moore, eds. History of Zoos and Aquariums: From Royal Gifts to Biodiversity Conservation. North of England Zoological Society, Chester. Carroll, C., R.J. Frederickson, and R.C. Lacy. 2014. Developing metapopulation connectivity criteria from genetic and habitat data to recover the endangered Mexican wolf. Conservation Biology 28:76-86. Frankham, R., R.C. Lacy, J.D. Ballou, M.R. Dudash, M.D.B. Eldridge, C.B. Fenster, J.R. Mendelson III, I.J. Porton, K. Ralls, and O.A. Ryder. 2014. Species concepts for conservation: Reply to Russello and Amato. Biological Conservation 170:334-335. Lacy, R.C., and J.P. Pollak. 2014. VORTEX: A Stochastic Simulation of the Extinction Process. Version 10.0. Chicago Zoological Society, Brookfield, Illinois, USA. Lacy, R.C., P.S. Miller, and K. Traylor-Holzer. 2014. Vortex 10 User’s Manual. IUCN SSC Conservation Breeding Specialist Group, and Chicago Zoological Society, Apple Valley, Minnesota, USA. Lacy, R.C., J.P. Pollak, P.S. Miller, L. Hungerford, and P. Bright. 2014. Outbreak version 2.1. IUCN SSC Conservation Breeding Specialist Group. Apple Valley, Minnesota, USA. Shoemaker, K.T., R.C. Lacy, M.L. Verant, B.W. Brook, T.M. Liveri, P.S. Miller, D.A. Fordham, and H.R. Akcakaya. 2014. Effects of prey metapopulation structure on the viability of black-footed ferrets in plague-impacted landscapes: a metamodelling approach. Journal of Applied Ecology 51:735-745. Hoffman, J.I., F. Simpson, P. David, J.M. Rijks, T. Kuiken, M.A.S. Thorne, R.C. Lacy, K.K. Dasmahapatra. 2014. High-throughput sequencing reveals inbreeding depression in a natural population. Proceedings of the National Academy of Sciences 111:3775-3780. Hedrick, P.W., and R.C. Lacy. 2015. Measuring relatedness between inbred individuals. Journal Robert C. Lacy -- Curriculum vitae 17

of Heredity 106:20-25. Willoughby, J.R., N.B. Fernandez, M.C. Lamb, J.A. Ivy, R.C. Lacy, and J.A. DeWoody. 2015. The impacts of inbreeding, drift, and selection on genetic diversity in captive breeding populations. Molecular Ecology 24:98-110. Wells, K., B.W. Brook, R.C. Lacy, G.J. Mutze, D.E. Peacock, R.G. Sinclair, N. Schwensow, P. Cassey, R.B. O’Hara, and D.A. Fordham. 2015. Timing and severity of immunizing diseases in rabbits is controlled by seasonal matching of host and pathogen dynamics. Journal of the Royal Society Interface 12:2014184 http://dx.doi.org/10.1098/rsif.2014.1184 Heinsohn, R., M. Webb, R.C. Lacy, and D. Stojanovic. 2015. A severe predator-induced population decline predicted for endangered, migratory swift parrots (Lathamus discolor). Biological Conservation (in press). Ballou, J.D., R.C. Lacy, K. Traylor-Holzer, K. Bauman, J.A. Ivy, P. Siminski, and C. Asa. Strategies for establishing and using genome resource banks in endangered species conservation. (submitted) Miller, P.S., R.C. Lacy, R. Medina-Miranda, R. López-Ortiz, and H. Díaz-Soltero. Confronting the invasive species crisis with PVA: An explicit, two-species metamodel of an endangered bird and its brood parasite in Puerto Rico. (submitted) Ueda, Y., R.C. Lacy, G. Alaks, J. Chen, S. Kajigaya, and N.S. Young. Molecular cloning and comparison of ribosomal protein, large P0 (RPLP0) of Peromyscus leucopus. (submitted) Manlik, O., J.A. McDonald, J. Mann, H.C. Smith, L. Bejder, M. Krützen, R.C. Connor, M.R. Heithaus, R.C. Lacy, and W.B. Sherwin. Feasibility and effectiveness of wildlife management actions are in stark contrast to the findings of conventional sensitivity analysis: lessons learned from two dolphin populations. (submitted) Prowse, T.A.A., C.J.A. Bradshaw, S. Delean, P. Cassey, R.C. Lacy, K. Wells, M. Aiello-Lammens, H.R. Akçakaya, and B.W. Brook. An efficient protocol for the sensitivity analysis of complex ecological models. (submitted) Canessa, S., G. Guillera-Arroita, J. Lahoz-Monfort, D.M. Southwell, S.J. Converse, D.P. Armstrong, I. Chadès, R.C. Lacy, P.S. Miller, and M.A. McCarthy. When do we need further research? A primer on calculating the value of information for applied ecology. (submitted) Canessa, S., S. Converse, D. Southwell, G. Guillera-Arroita, J. J. Lahoz-Monfort, D. Armstrong, R.C. Lacy, P.S. Miller, and I. Chades. Adaptive management for species conservation along the captive-wild spectrum. (in prep.) Malo, A.F., N. Wielebnowski, G. Alaks, A. Bellem, J. Bryant, and R.C. Lacy. Baseline corticosterone levels are positively associated with fitness. (in prep.) Lacy, R.C., A. F. Malo, and G. Alaks. Maintenance of genetic variation in quantitative traits of Peromyscus leucopus mice during generations of captive breeding. (in prep.)

Software developed and distributed for professional use PartInbr: A program providing calculation of inbreeding coefficients and partial inbreeding coefficients partitioned into the components contributed by each founder. Robert C. Lacy -- Curriculum vitae 18

PM2000: Software for demographic and genetic analysis and management of pedigreed populations. (Developed jointly with J. Ballou and J.P. Pollak). Used to guide management of captive populations of 100s of species globally. PMx: Software for demographic and genetic analysis and management of populations. (Developed jointly with J. Ballou and J.P. Pollak). Vortex: Simulation of interacting genetic, demographic, and environmental causes of extinction in small, isolated populations interconnected by occasional migration. Used by conservation and wildlife biologists to assist in the analysis and management of wild populations of 100s of species in more than 70 countries. Outbreak: Epidemiological simulation for modeling infectious disease. (Developed with J.P. Pollak, P.S. Miller, et al.) MetaModel Manager: Flexible modeling platform for linking simulation models representing diverse processes (such as species interactions, habitat change, climate change, disease, and social systems) to provide more holistic risk assessments for wildlife populations. (Developed with J.P. Pollak.) MMMacro: A dll library providing a macro script interpreter for use within system models. (MMMacroEntry provides a stand-alone program for writing and testing macro scripts.) Kenneth C. Balcomb, III – Brief biography

Ken graduated with a Bachelor’s degree in Zoology from the University of California at Davis in 1963, and was immediately employed by the Anatomy Department of the UC Davis veterinary school while taking graduate courses in Zoology and Oceanography. He was particularly interested in marine mammal anatomical specialization in the aquatic environment, and made frequent visits to the whaling stations in Richmond, California to obtain specimen materials for study. During one of these visits he learned of a planned whale-tagging expedition off the California coast to be conducted by the Marine Mammal Laboratory, Bureau of Commercial Fisheries, US Fish and Wildlife Service. Without thinking things through, Ken volunteered to be dishwasher on the expedition, and was ultimately hired to be the government whale- tagger at sea and the whale biologist at the whaling station. He found that he loved being at sea taking photos of whales, and it has been all downhill from that time. During the Vietnam war, Ken joined the US Navy and was trained as an Aviator and anti-submarine warfare specialist before being assigned to operations at several classified SOSUS (Sound Surveillance System) facilities in the Pacific. He trained himself to listen for and track whales in addition to the military mission of tracking Soviet submarines, which were much easier to detect. The trick is in thinking underwater while applying the laws of Physics. In 1975, Ken was honorably discharged from the Navy and went back to work under contract for his previous boss at the Marine Mammal Laboratory, now with NOAA National Marine Fisheries Service. His first assignment was to conduct a one-season census in 1976 of the killer whales in the Pacific Northwest, but that project has turned into a lifetime of fascinating research on a population segment now called Southern Resident Killer Whales (SRKW). Ken lives on San Juan Island and his specialty is development of benign techniques, particularly photo-identification, for study of marine mammals world-wide. He is the founder, Executive Director and senior scientist of the Center for Whale Research.

Curriculum vitae LAUREN J. N. BRENT

Centre for Research in Animal Behaviour Center for Cognitive Neuroscience University of Exeter Duke University Washington Singer Laboratories 450 Research Drive, Exeter, EX4 4QG, UK Durham, NC, 27708, USA Email: [email protected] Email: [email protected]

Nationality: Canadian and British Website: http://laurenbrent.com

CURRENT POSITIONS

Associate Research Fellow, University of Exeter School of Psychology, Centre for Research in Animal Behaviour. 2014-present.

Postdoctoral Associate, Duke University Center for Cognitive Neuroscience. 2010-present.

EDUCATION

PhD in Evolutionary Anthropology, University of Roehampton. Thesis: Investigating the causes and consequences of sociality in adult female rhesus macaques using a social network approach. Awarded March 2010.

MA in Anthropology, University of Calgary. Awarded September 2005.

BSc in Biology (Great Distinction), McGill University. Awarded June 2003.

RESEARCH GRANTS

National Institutes of Mental Health, R01-MH096875, $2.5 million (direct costs) 2012-2017 To support “Animal model of genetics and social behavior in autism spectrum disorders” Co-author, project coordinator & named postdoc.

International Primatological Society, Research Grant, $1,220 USD 2008-2009 “Social behaviour and the stress response in rhesus macaques”. Principal investigator.

The Leakey Trust UK, Research Grant, £400 GBP 2008-2009 “Social behaviour and the stress response in rhesus macaques”. Principal investigator.

University of Calgary Graduate Research Scholarship $5,000 CAD 2004-2005 “Infant handling in black and white colobus monkeys”. Principal investigator.

University of Calgary Thesis Research Grant, $1,000 CAD 2004-2005 “Infant handling in black and white colobus monkeys”. Principal investigator.

FELLOWSHIPS and AWARDS

Leverhulme Early Career Fellowship, to be held at the University of Exeter outcome pending Napier Memorial Medal for best PhD thesis winner, Primate Society of Great Britain, 2011 Duke Center for Interdisciplinary Decision Sciences Postdoctoral Fellowship 2011-2012 Natural Sciences & Engineering Research Council, Canada (NSERC) Scholarship, PhD 2006-2009 University of Roehampton Postgraduate Bursary 2006-2009 NSERC Postgraduate Scholarship, Masters 2004-2005 University of Calgary Dean’s Research Excellence Award 2004 NSERC Undergraduate Student Research Fellowship (x2) 2002, 2003

PUBLICATIONS

As Editor Brent LJN & Melin AM. The Genetic Basis of Primate Behavior – A Special Issue International Journal of Primatology (2014) Volume 35, Issue 1, ISSN: 0164-0291.

Peer-Reviewed Publications 2015 Croft DP, Brent LJN, Franks DW, Cant MA. The evolution of prolonged life after reproduction. Trends in Ecology and Evolution, in press.

Brent LJN, Franks DW, Foster EA, Balcomb KC, Cant MA, Croft DP. Ecological knowledge, leadership, and the evolution of menopause in killer whales. Current Biology, doi:10.1016/j.cub.2015.01.037.

Brent LJN. Friends of friends: Are indirect connections in social networks important to animal behaviour? Animal Behaviour, 103:211-222, doi: 10.1016/j.anbehav.2015.01.020.

Watson KK, Li D, Brent LJN, Horvath JE, Gonzalez-Martinez J, Ruiz-Lambides AV, Robinson AG, Skene JHP, Platt ML. Genetic influences on social attention in free-ranging rhesus macaques. Animal Behaviour, doi:10.1016/j.anbehav.2015.02.012.

2014 Brent LJN, Chang SW, Gariépy J, Platt ML. The neuroethology of friendship. Annals of the New York Academy of Sciences, Year in Cognitive Neuroscience, 1316: 1-17.

MacLean EL, Hare B, Nunn CL et al.ϯ The evolution of self-control. ϯ58 authors in total. Proceedings of the National Academy of Sciences, DOI:10.1073/pnas.1323533111.

Brent LJN, Semple S, MacLarnon A, Ruiz-Lambides A, Gonzalez-Martinez J, Platt ML. Personality traits are heritable in rhesus macaques (Macaca mulatta) but do not predict reproductive output. International Journal of Primatology, 35: 188-209.

Brent LJN, Melin AM. The genetic basis of primate behavior: Genetics and genomics in field- based primatology. International Journal of Primatology, 35: 1-10.

Dubuc C, Winters S, Allen WL, Brent LJN, Cascio J, Maestripieri D, Ruiz-Lambides AV, Widdig A, Higham JP. Sexually selected skin color is heritable and related to fitness in a non-human primate. Proceedings of the Royal Society, Biological Sciences, 281: 20141602.

Blomquist GE & Brent LJN. Applying quantitative genetic methods to primate social behavior. International Journal of Primatology, 35: 108-128.

2013 Brent LJN, Heilbronner SR, Horvath JE, Gonzalez-Martinez J, Ruiz-Lambides AV, Robinson A, Skene JHP, Platt ML. Genetic origins of social networks in rhesus macaques. Scientific Reports, 3: 1042, DOI 10.1038/srep01042.

Chang, SW, Brent LJN, Adams GK, Pearson J, Klein J, Watson KK, Platt ML. Neuroethology of primate social behavior. Proceedings of the National Academy of Sciences, 110: 10387- 10394.

Brent LJN, MacLarnon A, Platt ML, Semple S. Seasonal changes in the structure of rhesus macaque social networks. Behavioral Ecology and Sociobiology, 67: 349-359. Gilby IC, Brent LJN, Wroblewski EE, Rudicell RS, Hahn BH, Goodall J, Pusey, AE. Fitness benefits of coalitionary aggression in male chimpanzees. Behavioral Ecology and Sociobiology, 67: 373-381.

Dobson S & Brent LJN. On the evolution of the serotonin transporter linked polymorphic region (5-HTTLPR) in primates.Frontiers in Human Neuroscience, DOI 10.3389/fnhum.2013.00588.

2012 Buhl JS, Aure B, Ruiz-Lambides A, Gonzalez-Martinez J, Platt ML, Brent LJN. Response of rhesus macaques (Macaca mulatta) to the body of a dead group member that died from a fatal attack. International Journal of Primatology, 33:860-871.

2011 Brent LJN, Semple S, Dubuc C, Heistermann M, MacLarnon A. Social capital and physiological stress levels in free-ranging adult female rhesus macaques. Physiology and Behavior, 102:76- 83.

Brent LJN, Lehmann J, Ramos-Fernandez G. Social networks in the study of nonhuman primates: A historical perspective. American Journal of Primatology, 73: 720-730.

Higham JP, Hughes K, Brent LJN, Dubuc C, Engelhardt A, Heistermann M, Santos L, Stevens MS. Familiarity affects assessment of female facial signals of fertility by free-ranging male rhesus macaques. Proceedings of the Royal Society, Biological Sciences, 278: 3452-3458.

2010 Higham JP, Brent LJN, Dubuc C, Accamando AK, Engelhardt A, Gerald MS, Heistermann M, Stevens MS. Color signal information content and the eye of the beholder: A case study in the rhesus macaque. Behavioral Ecology, 21:739-746.

2009 Dubuc C, Brent LJN, Accamando A, Gerald MS, MacLarnon AM, Semple S, Heistermann M, Engelhardt A. Sexual skin color contains information about the timing of the fertile phase in Macaca mulatta. International Journal of Primatology, 13:777-789.

2008 Brent LJN, Teichroeb JA, Sicotte P. Preliminary assessment of natal attraction and infant handling in wild Colobus vellerosus. American Journal of Primatology, 70:101-105.

2002 Brent LJN, Drapeau P. A targeted ‘knockdown’ of channel expression in vivo with an antisense morpholino oligonucleotide. Neuroscience, 114: 275-278.

Publications in Review, Revision or Near Submission Watson KK, Madlon-Kay S, Ruiz-Lambides AV, Platt ML, Brent LJN. Adiposity is heritable in a free-ranging primate. American Journal of Primatology, in revision after 1st round of reviewer comments.

Larson SM, Ruiz-Lambides AV, Platt ML, Brent LJN. Dynamic network changes and the importance of affiliative relationships between non-relatives to group stability. In preparation to submit to Behavioural Ecology and Sociobiology.

Boyland NK, Mlynski DT, James R, Brent LJN, Croft DP. Social network structure of dairy cows, Bos taurus: From individual behaviour to group structure. In preparation to submit to Animal Behaviour.

Croft DP, Brent LJN, Franks DW, Balcomb KC, Ford J, Mazzi S, Johnstone R, Cant MA. Reproductive conflict and the evolution of menopause in killer whales. In preparation to submit to Nature.

INVITED TALKS, KEYNOTE PRESENTATIONS & SYMPOSIA ORGANISED

2015 Presidential Symposium, Animal Behavior Society, Invited speaker, Anchorage, Alaska. “What Makes a Rhesus Macaque Social Network? From Genes and Social Dynamics to Reproductive Success”

Invited seminar, University College London, Dept. of Anthropology

2014 Presidential Symposium, Animal Behavior Society, Invited speaker, Princeton, New Jersey. “Social Network Based Insights into the Evolution of Animal Behaviour”

Invited seminar, Durham University, Dept. of Anthropology

Invited seminar, University of Exeter, School of Psychology

2013 Keynote speaker, American Society of Primatology Conference, San Juan, Puerto Rico. “The Cayo Santiago Macaques: A Model for the Study of Behavioral Genetics and Human Psychiatric Disease”

2012 Symposium Organizer, International Primatological Society Congress, Cancun, Mexico. “Inside the Black Box: The Genetic Basis of Primate Behaviour”

2011 Colloquium, Dartmouth University (USA), Dept. of Anthropology

Invited seminar, Duke University, Dept. of Evolutionary Anthropology

Invited seminar, University of Pennsylvania, Dept. of Psychology

2010 Symposium Organizer, International Primatological Society Congress, Kyoto, Japan. “Social Network Theory and Analysis: Applications to the Study of Primate Sociality”

CONFERENCES ABSTRACTS (since 2011 only)

2015 Brent LJN, Franks DW, Foster EA, Balcomb KC, Cant MA, Croft DP. Ecological knowledge, leadership, and the evolution of menopause in killer whales. Animal Behavior Society, Anchorage, AL, USA.

Madlon-Kay S, Bey A, Passman R, Brent LJN, Watson KW, Skene JHP, Horvath J, Platt MLP, Jiang Y-H. SHANK3 polymorphism associated with social behavior in rhesus macaques. Society for Neuroscience, Chicago, USA.

2014 Madlon-Kay S, Watson KW, Brent LJN, Horvath J, Heilbronner SR, Skene JHP, Heller K, Platt MLP. Mapping genetic variation to morphology and behaviour in free-ranging rhesus macaque. Society for Neuroscience, Washington DC, USA.

Larson SM, Ruiz-Lambides A, Gonzalez-Martinez J, Platt ML, Fernandez-Duque E, Brent LJN. Social network dynamics of a group fission event in free-ranging rhesus macaques (Macaca mulatta). Animal Behavior Society, Princeton, NJ, USA.

Brent LJN, Semple S, MacLarnon A, Ruiz-Lambides AV, Gonzalez-Martinez J, Platt ML. Causes and reproductive consequences of social network position and personality in rhesus macaques. American Association of Physical Anthropologists, Calgary, Canada.

2013 Brent LJN, Semple S, MacLarnon A, Ruiz-Lambides AV, Gonzalez-Martinez J, Platt ML. Personality traits are heritable in rhesus macaques but do not predict reproductive output. Canadian Association of Physical Anthropologists. Toronto, Canada.

Rosati AG, Ruiz-Lambides AV, Brent LJN, Platt ML, Santos LR. The Role of Age in Looking-Time Performance: Younger Monkeys Look Longer Across Expectancy Violation and Referential Looking Tasks. Cognitive Development Society.

Ruiz-Lambides AV, Aure B, Caraballo-Cruz G, Platt ML, Brent LJN. Matrilineal overthrow followed by high mortality in free-ranging rhesus macaques. American Society of Primatologists, Puerto Rico.

2012 Brent LJN, Heilbronner SR, Horvath JE, Gonzalez-Martinez J, Ruiz-Lambides AV, Robinson A, Skene JHP, Platt ML. Genetic basis of social networks in rhesus macaques. 24th Congress of the International Primatological Society, Cancun, Mexico.

Sherwin H, Brent LJN, Horvath JE, Gonzalez-Martinez J, Ruiz-Lambides AV, Robinson A, Skene JHP, Platt ML. Polymorphic genes within the FMR1 pathway and social phenotypes in rhesus macaques. Society for Neuroscience, New Orleans, USA.

Watson KK, Li D, Skene JHP, Brent LJN, Platt ML. Serotonin-related genotypes influence social vigilance in free-ranging primates. Society for Social Neuroscience, New Orleans, USA.

Brent LJN, Heilbronner SR, Horvath JE, Gonzalez-Martinez J, Ruiz-Lambides AV, Robinson A, Skene JHP, Platt ML. Genetics of social network position in rhesus macaques. American Association of Physical Anthropologists, Portland, USA.

2011 Brent LJN, MacLarnon A, Semple S. Evidence for direct but not indirect reciprocity in the exchange of services between free-ranging rhesus macaques. Animal Behavior Society, Bloomington, USA.

Brent LJN, Heilbronner SR, Horvath JE, Gonzalez-Martinez J, Ruiz-Lambides AV, Robinson A, Skene JHP, Platt ML. Genetics of social network position in free-ranging rhesus macaques. Society for Neuroscience, Washington, DC, USA.

Gilby IC, Brent LJN, Wroblewski EE. Consequences of male chimpanzee dominance strategies: Fitness benefits of coalitionary aggression. American Association of Physical Anthropologists, Minneapolis, USA

TEACHING

Lecturing Wild Behaviour Field Course, Assistant Instructor, University of Exeter 2015 Comparative Issues in Psychology, Guest Lecturer, University of Exeter 2015 Primate Conservation, Guest Lecturer, Duke University 2013 Primate Biology, Guest Lecturer, University of Roehampton 2009 Behavioural Ecology Guest Lecturer, University of Roehampton 2009 Behaviour & Conservation, Guest Lecturer, University of Roehampton 2008 Primate Behaviour Field Course, Instructor, University of Roehampton 2008, 2009 Collecting Behavioural Data in Primates, Instructor, University of Roehampton 2008, 2009 Laboratory Practical on Enzyme Immuno-Assay, Guest Lecturer, Roehampton 2008, 2009 Primate Behaviour, Guest Lecturer, University of Calgary 2005 Primatology & Human Evolution, Guest Lecturer, University of Calgary 2005

Student Supervision Masters student co-supervisor, University of Exeter (2 students) 2014- Doctoral candidate mentor, University of Exeter (1 student) 2014- Doctoral candidate co-supervisor, University of Pennsylvania (1 student) 2012- Research assistant coordinator, manager, mentor, Duke University, (6 salaried, 7 volunteer) 2010- Doctoral candidate co-supervisor, Duke University (3 students) 2010- Masters student mentor and co-supervisor, University of Roehampton (3 students) 2010-2013 Medical student research project supervisor, Duke University (1 student) 2010-2012 Masters student co-supervisor, University of Pennsylvania (1 student) 2010-2011 Undergraduate project advisor, Duke University (6 students) 2010-2014 Undergraduate personal tutor, University of Roehampton (1 student) 2009

Teaching Assistantships Human Physiology, University of Roehampton 2009 Biostatistics, University of Roehampton 2008 Molecular Biology, University of Roehampton 2006 Primate Behaviour, University of Calgary 2004 Primatology & Human Evolution, University of Calgary 2004

Teaching Workshops Attended Teaching Genetics & Genomics Workshop: Amer. Assoc. of Anthropological Genetics 2013 Future Directions in Undergraduate Education, Duke University 2012

PROFESSIONAL CONTRIBUTIONS

Editorial Board Member: Frontiers in Behavioral and Evolutionary Ecology

Article or Grant Reviewer for: Animal Behaviour, American Journal of Physical Anthropology, American Journal of Primatology, Behavioral Ecology and Sociobiology, Behavioral Ecology, Frontiers in Ecology and Evolution, Genes Brains & Behavior, Hormones & Behavior, International Journal of Primatology, Journal of Mammalogy, Oikos, Physiology & Behavior, PLoS One, The Leakey Foundation, Scientific Reports.

Society Membership & Affiliations: Animal Behavior Society, Association for the Study of Animal Behaviour, International Primatological Society, Primate Society of Great Britain, Duke Network Analysis Center.

Academic Service: Athena Swan Working Group, University of Exeter, Psychology, Postdoc Representative, 2014-2015 Research Strategy Working Group, University of Roehampton, School of Life Sciences, Postgraduate Representative, 2008-2009

Other Contributions: Curator of Primate Tree, an academic genealogy of primate research: http://academictree.org/primate

RESEARCH OUTREACH

2015 Today’s Science, Infobase Learning for Students Conversations with Scientists Profile – “Lauren Brent: Getting to Know Animals” http://tsof.infobaselearning.com/recordurl.aspx?wid=99270&ID=32470

2014 New Scientist Contributing Author, Article – “Friendship: Friends with many benefits”. Publication date: 24/5/14.

Studium Generale Public Lecture, “The Evolution of Friendship” University of Groningen, Netherlands

2013 Duke Forward Funding Initiative. Program Leader, San Francisco. Theme: Friendship and Your Brain.

Duke Brain Awareness Week Invited Public Lecture, Motorco Music Hall. “Monkey Friends - What Primate Social Networks Teach us about Human Behavior.”

2012 TED Talent Search, New York, http://goo.gl/H6Jwn2

POPULAR PRESS COVERAGE

Television and Radio Appearances Mooney goes Wild, RTÉ Nature Programme, Ireland (interview about my killer whale research) Quirks and Quarks, Canadian Broadcasting Corporation (interview about my killer whale research) PBS Nature (an episode about animal social relationships) Primates of the Caribbean (an ARTE documentary about the research on Cayo Santiago)

Print and Online Media Brent et al. Current Biology 2015 *Altmetric score in the top 20 ever scored in Current Biology Science, Nature, New York Times, National Geographic, BBC Earth, New Scientist, Scientific American, Smithsonian Magazine, The Times, Washington Post, Forbes Magazine, more…

Brent et al. Scientific Reports 2013 *F1000Prime recommended Wall Street Journal (Column by Matt Ridley), Science Daily, Daily Mail UK

Brent et al. Physiology & Behavior 2011 Time Magazine (cover story by Carl Zimmer), Time for Kids

Interview about my research on social bonds TED blog

Gilby, Brent, et al. Behav Ecol & Sociobiol 2013 Science Daily, WUNC Radio - The State of Things

Higham et al. Proc Roy Soc B 2011 New York Times

Croft CV, May 2015

Dr Darren P. Croft Curriculum Vitae

Centre for Research in Animal Behaviour College of Life and Environmental Sciences  +44 1392 725649 University of Exeter Email: [email protected] Exeter EX4 4QG, UK

Education

2000 – 2004 PhD in Behavioural Ecology, University of Leeds. The social organisation of the Trinidadian guppy (Poecilia reticulate). 1995 – 1999 BSc Hons, Class I, Biology, University of Leeds.

Current Position

2014 – Current Associate Professor of Animal Behaviour, University of Exeter. Director of Research for Psychology.

Previous Positions

2009 – 2014 Senior Lecturer in Animal Behaviour, University of Exeter. Academic Lead for the Centre for Research in Animal Behaviour. Deputy Director for Research (Psychology). 2008 – 2009 Lecturer in Animal Behaviour, University of Exeter. 2006 – 2008 Lecturer in Animal Behaviour, Bangor University.

2005 Postdoctoral Research Fellow, University of Leeds. Extended Particle Swarms (EPSRC). 2004 Postdoctoral Research Fellow, University of Leeds. Mechanisms and Functions of Familiarity in Fishes (The Leverhulme Trust).

A. Research My research has had a major influence on our understanding of the mechanisms, constraints and functions underpinning the structure of animal societies [>60 papers in leading international journals including Science, Current Biology, Trends in Ecology and Evolution, The American Naturalist and Proceedings of the Royal Society [cited ~1500 times, web of science h-index = 20]. I have an excellent track record of securing research funding and during my time at Exeter I have been awarded >£3 M in research income (>1M as PI) from research councils (BBSRC, NERC and The Danish Council for Independent Research - Natural Sciences), charities (the Leverhulme Trust and the Nuffield Science foundation), industry (DairyCo, Pets at Home) and government (Defra, Cefas). My work has had major international inter-disciplinary impact and has been cited across disciplines including: biological sciences, veterinary sciences, psychology, behavioural ecology, ecology, evolution, mathematics and computer sciences. My international standing is reflected in 15 invitations to give international seminars and four invitations to teach on international residential PhD courses and I get regular invitations to review major international grant applications [e.g. ERC Synergy Grants and NSF]. I am currently an associate editor for Behavioural Ecology and Sociobiology and on the ethics editorial board for Animal Behaviour.

I have played a leading role in developing the application of the social network approach to non-human animal societies. In particular in 2008 I published a monograph as lead author entitled “Exploring Animal Social Networks” which draws together a disparate body of methods from across disciplines and

1 Croft CV, May 2015 demonstrates how they can be applied to non-human animals. This book has played a major role in developing the application of social network analysis to animal societies and has propelled many new studies. More recently in 2014 I co-edited a book on Animal Social Networks published by Oxford University Press which reviews the impact that social network analysis has had on the study of animal behaviour. Using social network analysis I have addressed key ecological and evolutionary questions in a range of study species. For example, using the guppy as a model system I undertook the first empirical study to investigate the link between social network structure and cooperation in real world non-human animal populations; work that features in at least one introductory animal behaviour text book. More recently I have established a project using long term social and demographic datasets for two resident killer whale populations to study the social drivers of survival and fecundity. This work aims to provide the first empirical test in a non-human animal of the general theoretical framework proposed to explain reproductive senescence in natural populations. The initial outputs from this project have been published in Science and Current Biology and I have been awarded £620k [as PI] from NERC for this work. I am also part of a large collaborative project aimed at understanding the social drivers of disease transmission in wild populations of badgers. Through the application of social network analysis this project aims to determine the causal relationship between social perturbation and the spread of tuberculosis (TB). The project has attracted £748k from NERC and has the potential to influence UK government policy on the eradication of bovine TB. Finally, taking inspiration from my work on wild animal populations I have developed a collaboration with DairyCo and Defra to apply social network theory to address key problems of welfare and productivity in the UK dairy industry which has also attracted funding from the BBSRC (£820k). The aim of this work is to transfer cutting edge concepts and methodologies from behavioural ecology and social network theory to real world issues relating to food security and animal welfare.

External Grants Awarded

2015-2018 The Evolution of Eye Salience as a Signal for Communication. The Leverhulme Trust, PI D.P. Croft, CoI S.K. Darden (£186,907).

2014-2017 Social niche construction and evolutionary implications for animal behaviour. Danish Council for Independent Research (FNU Denmark) PI T Dabelsteen (University of Copenhagen), CoIs D.P. Croft, S Darden (University of Exeter) and ON Larsen (University of Southern Denmark) (£295,000).

2014-2017 Infection In Dynamic Social Networks of a Wild Mammal NERC, PI R McDonald, CoI’s D.P. Croft, M. Boots, D.J. Hodgson (£748,000). 2014-17: Stress and Mortality in Aquarium Fish: Interventions for Improving Health and Well- being. PhD studentship funding from Pets at Home and Cefas, Joint PI’s D.P. Croft, C. Tyler (£59,000)

2013-2016 The Evolution of Prolonged Post-reproductive Lifespan in a Non-human Mammal NERC, PI D.P. Croft, CoIs, D. Franks (York) and M Cant (Exeter) (£620,000). Application scored 9 / 10 and was rated number 1 of 28 grants (panel E Dec 2012 submission).

2012-2015 Assessment of Dairy Cow Welfare through Predictive Modelling of Individual and Social Behaviour BBSRC, Joint PIs J. Amory (Wittle College) E. Codling (University of Essex) and CoI D.P. Croft (£820,330).

2012-2015 The influence of Social Networks on Welfare and Productivity in Dairy Cattle. Defra, PI D.P. Croft, CoI J. Madden (University of Exeter) (£45,000)

2012-15: Social Components of Welfare and Productivity in Dairy Cattle. PhD studentship funding from DairyCo, PI D.P. Croft, CoI J. Madden (University of Exeter) (£50,000) 2011-2013: The Evolution of Cooperation in Structured Animal Societies. The Leverhulme Trust. PI D.P Croft, CoIs C. Tyler (University of Exeter) and R James (University of Bath) (£143,813) 2011-2014: Cognitive and Physical Aspects of Animal Communication Networks. Danish Council for Independent Research (FNU Denmark). PI T Dabelsteen (University of Copenhagen), CoIs D.P. Croft, S Darden (University of Exeter) and ON Larsen (University of 2 Croft CV, May 2015

Southern Denmark) (£226,000).

2006-2010: Mechanisms and Functions of Sexual Segregation. NERC (NE/E001181/1) PI: D.P. Croft (£68,000). 2006-2010: Behavioural Syndromes: Environmental Influences in a Naturally Clonal Species of Fish. Nuffield Science Award PI: D.P. Croft (£5000). 2005 Sexual Segregation in the Trinidadian Guppy Fisheries Society of the British Isles: PI: D.P. Croft (£1,800).

Internal University Grants

2014: Evolutionary origins of the menopause Exeter Bridging the Gaps Award PI Dr Anna Murray (Exeter Medical School), CoI D.P. Croft (£2500).

2012: Bridging the Gaps Award (University of Exeter): Automated detection and classification of collective behaviour in large spatio-temporal datasets. PI A. Galton, CoIs D.P. Croft and Z. Wood (£2,482).

2011: Bridging the Gaps Award (University of Exeter): Animal Social Behaviour Based PSO. PI Ed Keedwell, CoI D.P. Croft (£4000). 2010-2011: University of Exeter Link Award to build research collaboration with DairyCo (£500).

Other research funding

2009-2012: PhD Studentship from the Fisheries Society of the British Isles (awarded to D. Jacoby) (£57120). 2008-2011: PhD Studentship from the Fisheries Society of the British Isles (awarded to M. Edenbrow) (£57120).

B. Teaching

Courses Taught Exeter (2008-Current)

ABH1101: An introduction to Animal Behaviour (15 credits). 130 students, 12 x 1-hour lectures, 3 x 1-hour tutorial classes and a 4-hour practical (note the tutorial and practical run multiple times). ABH2115: Laboratory and Field Methods in Animal Behaviour (30 credits). 30 students. Semester 1: 5 x 3-hour laboratory practical sessions on fish behavior. Semester 2: 5 x 1-hour problem based learning exercises and a 7-day residential field course (Wales). ABH2104: Evolution and Behaviour (15 credits). 30 students, 12 x 1-hour lectures, 3 x 1-hour tutorials, 1 x 4-hour practical held at Plymouth Marine Aquarium. PSY2214: Observations and Experiments in Animal Behaviour (15 credits). 30 students. 5 x 3- hour laboratory practical sessions on fish behavior. PSY2215: Problems and Projects in Animal Behaviour (15 credits). 30 students. 5 x 1-hour problem based learning exercises and a 7-day residential field course (Wales).

Courses Taught Bangor University (2006-2008)

MSc: Marine Animal Behaviour (10 credits 2006 only), 12 x 2-hour lectures and 2 x 3-hour practical classes. MSc: Ecological Interactions (20 credits) 12 x 2-hour lectures and 4 x 3-hour practical classes.

3 Croft CV, May 2015

BSc: Field Course in the Gambia (10 credits) 7 days in the field.

BSc: Behaviour and Evolution (20 credits) 8 x 3-hour practical classes in animal behaviour.

BSc: Field Study Skills (10 credits) 2-day field practical.

BSc: Ecology and Evolution (20 credits) 5 x 1-hour lectures.

MSc: Marine Animal Behaviour (10 credits 2006 only) 12 x 2-hour lectures and 2 practical classes.

Teaching Innovations and Evidence of Excellence

During my academic career I have developed a number of significant teaching innovations that have had major impact. Whilst a lecturer at Bangor University (2006-2008) I established a new degree program in Zoology with Animal Behaviour, which has rapidly become a major recruiter to the undergraduate community in Bangor. Associated with this degree I also developed two new modules which included developing a new 10-credit field course to The Gambia which has become a unique selling point of the degree program. Since arriving in Exeter in Oct 2008 I have developed three new modules. One in particular demonstrates my approach to teaching and the significant impact that my teaching innovations have had on the curriculum. In 2009 I co-developed a new module (Laboratory and Field Methods in Animal Behaviour [ABHY2114]). As part of this module I introduced a problem based learning approach combined with practical classes using live fish in which the students designed experiments to test hypotheses formulated by the class participants. The practical’s objectives are to introduce students to aspects of experimental design, data analysis and the issue of ethics in animal research. The feedback from the students’ appraisal of the practical classes has been extremely positive with comments including: “one of the most interesting practical’s I have ever taken part in” and “although most practical courses are written and prepared well in advance, I felt that the fact we influenced what we were doing made the practical’s mean more than something written ages ago”. My teaching is repeatedly rated as excellent by both peers and students which is evidenced by the student evaluation scores of my teaching. For example, for the academic year 2014-2015 my teaching scored 5 out of 5 in the student evaluation. The student comments in these evaluations included comments such as: “A truly excellent lecturer. His interactive lectures keep us interested and awake, asking questions to students throughout.”; “….he is interested in our opinions and brings anecdotes from his experience to make us laugh.”; “Never patronizing his office door is always open to ask questions” and “His enthusiasm and knowledge of his topic is never ending.”

Grants obtained for teaching purposes

2010: University of Exeter Alumni Award to organise a 3-day conference and student workshop on animal behaviour (£5000).

C. External Examining

2015: PhD examiner for Jessica Stephenson, University of Cardiff, UK.

2014: PhD examiner for Nicola Atton, University of St Andrews, UK.

2012: PhD examiner for Anne Bibost Macquarie University, Australia.

2012: PhD examiner for Kelly Davis University of Sidney, Australia.

2010: PhD examiner for Ashley Le Vin, University of Glasgow, UK.

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2010: MSc examiner for Sophie Holles, University of Bristol, UK.

D. Postgraduate Personnel

PhD students

2014-Current: PhD co-supervisor for Sylvia Dimitriadou: The evolution of cooperation among non- kin.

2014-Current: PhD supervisor for Jade Barrett: Social networks and welfare in dairy cattle.

2014-Current: PhD supervisor for Chloe Stephens: Stress and mortality in aquarium fish.

2014-Current: PhD co-supervisor for Lucy Capstick: From the Individual to the Ecosystem: Variation in Corvid Ranging, Predatory Behaviour and Trapability, and its Effects on Songbird Productivity

2013-Current: PhD co-supervisor Amy Grace Fensome: Habitat fragmentation and mammal conservation: productivity, stress and social networks.

2013-Current: PhD co-supervisor for Stephen White: Maternal effects in guppies.

2012-Current: PhD supervisor for Natasha Boyland: The influence of social networks on welfare and productivity in dairy cattle.

2012-Current: PhD supervisor for Paul Rose: Social behaviour of flamingos in captivity: implications for animal welfare and conservation.

2012-2014: PhD co-supervisor for Matthew Silk: Facebook for geese: the causes and consequences of non-random associations in a social forager.

2010-2012: PhD supervisor for Emma Foster: The structure and function of social networks in the southern resident killer whale population. 2009-2012: PhD supervisor for David Jacoby: Social networks in a marine predator.

2008-2011: PhD supervisor for Mathew Edenbrow: Behavioural phenotypes: associated life history traits and environmental effects on development. 2006-2010: PhD supervisor for Dawn Thomas (University of Bangor). The evolution of behavioural phenotypes in guppies. 2006-2010: PhD co-supervisor for Jonathan Ward (University of Leeds): Mechanisms and function of shoaling behaviour. 2005-2008 PhD co-supervisor for Chantima Piapong (University of Leeds): The mechanisms and functions of sexual segregation in guppies. 2004-2007 PhD co-supervisor for Philip Thomas (University of Leeds): The structure and function of social networks in guppies.

Post-Doctoral Assistants/Fellows

2015-2016: Robert Heathcote: The Evolution of Eye Salience as a Signal for Communication. The Leverhulme Trust. 2014-2015: Robert Heathcote: The Evolution of Cooperation in Structured Animal Societies. The Leverhulme Trust. 2014-2017: Lauren Brent: The Evolution of prolonged post-reproductive lifespan in a non-human mammal. NERC.

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2011-2013: Mathew Edenbrow: The Evolution of Cooperation in Structured Animal Societies. The Leverhulme Trust. 2011-2013: Dr Safi Darden, Social Implications of the Battle of the Sexes (Leverhulme Research Fellow) 2007-2010: Dr Safi Darden, Mechanisms and Functions of Sexual Segregation. NERC. (During this time period Dr Darden worked part time (0.5FTE) and was on maternity leave from June 2008-April 2009).

E. Administration I have demonstrated significant management, leadership and entrepreneurship in research and teaching. Since 2014 I have been the Director of Research for the School of Psychology which has 35.6 FTE academic staff. In this role I have responsibility for developing and implementing the departmental research strategy. Prior to this appointment I was the research group leader for the Centre for Research in Animal Behaviour (CRAB) which has 5.3 FTE academic staff. I have played a fundamental role in shaping the research strategy of CRAB and in establishing the group as an international centre for excellence for research and teaching in animal behaviour. I have provided mentoring to junior staff (J. Skelhorn, S. Darden and C. Ioannou) two of whom were awarded personal research fellowships within their first year at Exeter (SD and CI).

2014-Current: Director of Research (Psychology), University of Exeter.

2013-Current: Member of the Leverhulme Trust strategy group, University of Exeter.

2012-2013: Deputy Director for Research (Psychology), University of Exeter.

2011-14: Research Group Leader for the Animal Behaviour Research Group (Psychology) University of Exeter. 2011: Organised and chaired a space review of the Washington Singer Laboratories (University of Exeter) with the aim of improving the internal appearance of the building. 2010-11: Program Director for MSc in Animal Behaviour, University of Exeter.

2010-11: Deputy Research Group Leader for the Animal Behaviour Research Group (Psychology), University of Exeter.

F. External Recognition

Awards

2014: The Fisheries of the British Isles medal: awarded to younger scientists in recognition of their achievements.

Professional Bodies

2001-Current: Fisheries Society of the British Isles.

Invited Keynote Speaker

2015: 52nd Annual Conference of the Animal Behavior Society Anchorage, Alaska [I was unable to take up this invitation due to family commitments]. 2015: Animal Social Networks in Behavioural Research, Neuchâtel, Switzerland.

2014: NIMBioS Investigative Workshop, Animal Social Networks, Univ. of Tennessee, Knoxville, USA.

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2009: Complex Networks across the Natural and Technological Sciences. University of Strathclyde. 2007: International Conference of Ethology, Halifax, Canada, 2007.

2006: Social Network Analysis in Animal Behaviour, University California Davis USA.

Invited Departmental Seminars I receive many more invitations for departmental seminars (usually 3-4 per annum) than I am able to accept due to family commitments. 2015: University of Bern Switzerland.

2013: University of Bristol.

2012: University of the West Indies, Trinidad.

2012: University of Nottingham.

2011: University of Cardiff.

2011: University of Sussex.

2011: University of Copenhagen, Denmark.

2007: University of Exeter.

2007: University of the West Indies, Trinidad.

2006: University of Bangor.

2005: University of Copenhagen, Denmark.

2005: University of Edinburgh.

2004: University of Karlstad, Sweden.

Invited Lecturer 2013: 1-day MSc course at the University of Copenhagen (social networks in animals).

2011: 1-day MSc course at the University of Copenhagen (social networks in animals).

2010: 7-day PhD course at the University of Gothenburg (fish behaviour).

2008: 7-day PhD course at the University of Copenhagen (commutation networks in animals).

Editorial Responsibilities

2012-Current: Ethics committee for the Journal of Animal Behaviour. 2011-Current: Associate Editor of Behavioural Ecology and Sociobiology.

Reviewing

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Referee for BBSRC, ERC, NERC, NSF, Animal Behaviour, Behaviour, Behavioural Ecology, Behavioural Ecology & Sociobiology, Biology Letters, Ecology Letters, Fish and Fisheries, Functional Ecology, Journal of Comparative Psychology, Journal of Ethology, Journal of Fish Biology, Marine Ecology-Progress Series, Methods in Ecology and Evolution, Nature Communications, Oecologia, Proceedings of the Royal Society B, Princeton University Press, Trends in Ecology and Evolution

Conference organization

nd th th 2014: (2 -5 Sept) Organiser of the 6 International Conference of Pociliid Biologists, University of Exeter.

2012 (1-3rd Aug) Co-organiser: Animal Social Networks: Perspectives and Challenges ZiF (Centre for Interdisciplinary Research), Bielefeld, Germany

2010: Organiser of the Association for the Study of Animal Behaviour (ASAB) Easter meeting at Exeter University. 3-day conference for 150 delegates. 2005: On the organising committee of the International Ethology Congress, Budapest.

Selected media coverage I am committed to communicating my science to audiences outside the scientific community and I regularly participate in events such as UK National Science Week. I have authored media-friendly articles in popular science magazines (e.g. Planet Earth). My research is regularly featured in the national and international press (e.g. New York Times, National Geographic and TIME Magazine) and I have given a number of national and international radio interviews.

2015: “Post-menopausal whales lead the hunt After they reach menopause, female killer whales help” Nature.

2015: Granny whales 'do know best' Independent

2015 “Whales could help explain the mystery of menopause” The Times.

2014 Repeat: I'm a nice shark, not a mindless eating machine. The Daily Telegraph

2012: “Dead sons may explain killer whale menopause” New Scientist.

2012: “Not so scary now: Researchers find male killer whales are 'mummy's boys'” Daily Mail. 2012: “Significant udders: cows need their best friends”. The Daily Telegraph

2012: “Secrets of the herd mentality”. The Times.

2011: “The amphibious fish that mates with itself” New Scientist.

2009: “Sexual harassment prevents females bonding, says study” Daily Times India.

2009: “Sexual harassment in fish breaks down social structure” Practical Fish Keeping.

2009: “Guppy females take risks to avoid harassment” NERC Planet Earth.

2009 “Lemon sharks hang out with friends” NERC Planet Earth.

2008: “Female guppies risk death to avoid males on the make” The Discovery Channel.

2008: “Females risk death to avoid sexual harassment” The Independent.

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2006: “Guppies prefer death to sex” The Metro.

2006: “Fish courting 'sex pest' dangers” BBC News.

2004: “Hi Guppy!” The Hindu.

2004: “Networking in the fish tank: the humble guppy always remembers his friends” The Independent.

G. Publications

Books

74. Krause J, James R, Franks D, Croft D.P. (2014). Animal Social Networks, Oxford University Press ISBN: 9780199679058 288p 73. Croft DP, James R, Krause J (2008) Exploring Animal Social Networks. Princeton University Press, Princeton, NJ. ISBN: 0691127522 208p.

Chapters in Books

72. Krause, J., Croft, D.P. and Wilson, A.D.M. (2014). The network approach in teleost fishes and elasmobranchs. In: Krause, J., James, R., Franks, D., and Croft, D.P. (eds.) Animal Social Networks, Oxford University Press, Oxford pp. 150-159. 71. Croft D.P., Edenbrow M, Darden S.K. (2014). Assortment in social networks and the evolution of cooperation in Krause J, James R, Franks D, Croft DP (eds.) Animal Social Networks, Oxford University Press, Oxford. pp. 13-23. 70. Ioannou C.C., Couzin I.D., James R., Croft D.P. and Krause J. (2011) Social organization and information transfer in schooling fish. In: Fish Cognition and Behaviour, (Brown, C., Laland, K.N. & Krause, J. eds.) Blackwell Publishing, Cambridge, pp. 217–239. 69. Krause J., James R. and Croft D.P. 2010. Group-living in poeciliids. In: Ecology and Evolution of Poeciliid Fishes (Evans J, Pilastro A & Schlupp I eds.). University of Chicago Press, Chicago, pp.145-154. 68. Krause J, Ward A.J.W., Croft D. P. and James R (2008). Group living and social networks. In: Fish Behaviour (Magnhagen C, Braithwaite V, Forsgren E & Kapoor BG eds.). Science Publishers. pp. 485- 498. 67. Croft D. P., Griffiths, S.W., Magurran, A.E. and Krause J. (2007). The Influence of Reproductive Behaviour on the Social Organisation of Shoaling Fish . In: Fish Reproduction. Science Publishers. ISBN 978-1-57808-331-2.pp. 377-396 66. Krause J., James R. and Croft D.P. 2006. Soziale Netzwerke bei Tieren. In: Methoden der Verhaltensbiologie (Naguib M ed.), Springer Verlag, Heidelberg, pp. 148-152 65. Couzin, I.D., James, R., Croft D. P. and Krause J. (2006). Social organization and information transfer in schooling fish. In: Fish Cognition and Behaviour, (Brown, C., Laland, K.N. & Krause, J. eds.) Blackwell Publishing, Cambridge, ISBN 9781405134293, pp. 166–185 64. Croft D. P., James R., and Krause J. (2005). Predator avoidance as a factor for size assortative shoaling and its implications for sex segregation in fish. In: Sexual segregation in vertebrates: ecology of the two sexes. (K. Ruckstuhl and P. Neuhaus eds.), Cambridge University Press, Cambridge, pp. 115-126

Papers in Journals I support PhD students and post-doctoral researchers in my laboratory by not being first author on joint papers with them unless I contribute more than 80% to the research (e.g. development of ideas, undertaking the research etc.). * denotes first author papers by my students and post docs.

63. *Brent L.J.N., Franks D.W., Foster E.A., Balcomb K.C., Cant M.A., Croft D.P. ͌ (2015). Ecological knowledge, leadership, and the evolution of menopause in killer whales. Current Biology, 25(6), 746- 750. ͌ Corresponding Author

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62. Silk M.J., Jackson A.L., Croft D.P., Colhoun K., Bearhop S. (2015). The consequences of unidentifiable individuals for the analysis of an animal social network. Animal Behaviour, 104, 1-11.

61. Tranmer M., Marcum C.S., Morton F.B., Croft D.P., de Kort S.R. (2015). Using the relational event model (REM) to investigate the temporal dynamics of animal social networks. Animal Behaviour, 101, 99-105.

60 *Jacoby D.M.P., Fear L.N., Sims D.W., Croft D.P. (2014). Shark personalities? Repeatability of social network traits in a widely distributed predatory fish. Behavioral Ecology and Sociobiology, 68(12), 1995- 2003.

59. Wilson A.D.M. Krause S. James R., Croft D.P., Ramnarine I.W., Borner K.K., Clement R.J.G. & Krause J. 2014 Dynamic social networks in guppies (Poecilia reticulata). Behavioural Ecology and Sociobiology 68, 915-925. 58. * Boyland, N.K. James, R. Mlynski, D. T. and Croft D.P. (2013). Spatial proximity loggers for recording animal social networks: consequences of inter-logger variation in performance. Behavioural Ecology and Sociobiology 67: 1877-1890.

57. * Edenbrow, M. and Croft DP (2013) Environmental and genetic effects shape the development of personality traits in the mangrove killifish, Kryptolebias marmoratus. Oikos 122 667–681 56. Croft D.P., Hamilton P.B., Darden S.K., Jacoby D.M.P., James R., Bettaney E.M. and Tyler C.R. (2012) The role of relatedness in structuring the social network of a wild guppy population Oecologia 170: 955- 963 55. * Foster, E.A. Franks, D.W. Mazzi, S. Darden, S.K. Balcomb, K.C. Ford, J.K.B. and Croft D.P.͌ (2012) Adaptive benefit of prolonged female post-reproductive lifespan in a long-lived cetacean. Science 337(6100): 1313. Recommended by Faculty of 1000. ͌ Corresponding Author 54. * Jacoby, D.M.P., Sims D.W. and Croft D.P. (2012) The effect of familiarity on aggregation and social behaviour in juvenile small spotted catsharks Scyliorhinus canicula. Journal of Fish Biology 81: 1596- 1610. 53. Griffiths AM., Jacoby DMP, Casane D, McHugh M, Croft DP, Genner MJ, and Sims D.W. (2012) The first analysis of multiple paternity in an oviparous shark species, the small-spotted catshark (Scyliorhinus canicula L.). Journal of Heredity 103:166-173 52. Morley M, Keedwell EC, and Croft D.P. (2012) CTB-PSO: Continuous Trait-Based Particle Swarm Optimisation, ANTS2012, Brussels, Belgium, 12th - 14th Sep 2012, Springer LNCS 7461. 51. * Foster, E.A., Franks, D.W., Morrell, L.J., Balcomb, K.C., Parsons, K.M., van Ginneken, A., and Croft, D.P. (2012). Social network correlates of food availability in an endangered population of killer whales, Orcinus orca. Animal Behaviour, 83: 731-736 50. Bode, WFN, Franks DW Wood AJ Piercy JJB Croft D.P. and Codling EA. (2012) Distinguishing social from non-social navigation in cohesive animal groups. American Naturalist 179: 621-632 49. * Edenbrow, M. and Croft D.P. (2012) Sequential hermaphroditism and personality in a clonal vertebrate: the mangrove killifish. Behavioural processes. 90: 229-237, 48. * Edenbrow, M. and Croft D.P. (2012) Kin and familiarity influence association preferences and aggression in the mangrove killifish Kryptolebias marmoratus (Poey). Journal of Fish Biology. 80: 503- 518 47. * Brask, JB, Croft, D.P., Thompson, K. Dabelsteen, T. and Darden S.K. (2012) Social associations based on sexual attractiveness: a strategy to reduce sexual harassment from males. Proceedings of the Royal Society of London Series B-Biological Sciences 279: 1748-1753 46. Croft D.P., Edenbrow M., Darden S.K., Ramnarine I.W., van Oosterhout C. and Cable J. (2011) Effect of gyrodactylid ectoparasites on host behaviour and social network structure in guppies Poecilia reticulata.

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Behavioural Ecology and Sociobiology 65: 2219-2227

45. Kelley J.L., Morrell L.J., Inskip C., Krause J. and Croft D.P. (2011) Predation risk shapes social networks in fission-fusion populations. PLoS ONE 6(8): e24280. doi:10.1371/journal.pone.0024280 44. Mourabit S., Edenbrow M., Croft D.P. and Kudoh, T. (2011) Embryonic Development of the Self-fertilizing Mangrove Killifish Kryptolebias marmoratus. Developmental Dynamics 240: 1694–1704 43. * Edenbrow M.E. and Croft D.P. (2011) Behavioural types and life-history strategies during ontogeny in the mangrove killifish (Kryptolebias marmoratus). Animal Behaviour 82: 731-741 42. * Edenbrow M.E. Darden S.K., Ramnarine I.W. Evans J.P., James, R. and Croft D.P. (2011) Environmental effects on social interaction networks and male reproductive behaviour in guppies (Poecilia reticulata). Animal Behaviour 81: 551-558 41. Guttridge T.L. Gruber S.H. DiBattista J.D. Feldheim K.A., Krause S. Croft D.P. and Krause J. (2011) Assortative interactions and leadership in free-ranging population of juvenile lemon sharks, Negaprion brevirostris. Marine Ecology Progress Series 423: 235–245 40. Jones, K. A., Croft D. P. Ramnarine, I and Godin, J-G. J (2010) The role of active choice and population predation pressure in size assortative shoaling in the guppy. Poecilia reticulata. Ethology 116 147-154 39. * Piyapong, C., Chapman B.B., Krause J., Ramnarine I. and Croft D.P. (2010) Sex matters: A social context to boldness in guppies. Behavioural Ecology, 21: 3-8 38. Guttridge T.L., Gruber S.H., Gledhill K.S., Croft D.P., Sims D.W. and Krause, J. (2009) Social preferences in juvenile lemon sharks, Negaprion brevirostris. Animal Behaviour, 78: 543-548. 37. Croft D.P., Krause J., Darden S.K., Ramnarine I.W., Faria J.J. and James R. (2009) Behavioural Trait Assortment in Social Networks: Patterns and Implications. Behavioural Ecology and Sociobiology 63: 1495-1503. 36. * Darden S.K., James R., Ramnarine I.W. and Croft D.P. (2009) Social Implications of the Battle of the Sexes: Sexual Harassment Disrupts Female Sociality and Social Recognition. Proceedings of the Royal Society of London Series B-Biological Sciences, 276: 2651-2656. 35. Croft D.P. Darden S.K. and Ruxton G.R. (2009) Predation risk as a driving force for phenotypic assortment: a cross-population comparison. Proceedings of the Royal Society of London Series B- Biological Sciences 276: 1899-1904 34. Dyer J., Croft D.P,. Morrell, L.J. and Krause J. (2009) Shoal composition determines foraging success in the guppy. Behavioural Ecology 20: 165-171. 33. Shorrocks B. and Croft D.P. (2009) Necks and networks: a preliminary study of population structure in the reticulated giraffe (Giraffa camelopardalis reticulata Linnaeus). African Journal of Ecology, 47: 374- 381. 32. Botham M.S., Hayward R.K., Morrell L.J., Croft D.P., Ward J., Ramnarine I. and Krause J. (2008) Population differences in the response to different predators by a common prey species. Ecology, 3174– 3185. 31. * Darden S. K. and Croft D. P. (2008) Male harassment drives females to alter habitat use and leads to segregation of the sexes. Biology Letters 4: 449-451. 30. Morrell L.J, Croft D.P., Dyer R.J.G., Chapman B.B., Kelley J.L., Laland K.N. and Krause J. (2008) Association patterns and foraging behaviour in natural and artificial guppy (Poecilia reticulata) shoals. Animal Behaviour 76: 855-864. 29. Dyer J.R.G, Ioannou C., Morrell L.J., Croft D.P., Couzin I.D., Waters D.A., Wong Y. and Krause J. (2008) Consensus decision making in human crowds. Animal Behaviour 75: 461-470 28. * Thomas P.O.R., Croft D.P., Morrell L.J., Davis A, Faria J.J., Dyer J.R.G., Piyapong C., Ramnarine I., Ruxton G.D. and Krause J. (2008) Does defection during predator inspection affect social structure in

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wild shoals of guppies? Animal Behaviour 75: 43-55.

27. * Piyapong C., Morrell L.J., Croft D.P., Dyer, J.R.G., Ioannou, C.C., and Krause, J. (2007) A Cost of Leadership in Human Groups. Ethology, 113, 821-824. 26. Morrell L.J., Hunt K.L., Croft D.P. and Krause J. (2007) Diet, familiarity and shoaling decisions in guppies. Animal Behaviour, 74, 311-319. 25. Croft D. P., Morrell L. J., Wade A.S., Piyapong C., Ioannou C. C., Dyer J. R. G., Chapman B., Wong Y. and Krause J. (2006). Predation risk as a driving force for sexual segregation: A cross-population comparison. American Naturalist, 167: 867-878. Recommended by Faculty of 1000 24. Croft D. P., James R., Thomas P.O.R., Hathaway C., Mawdsley D., Laland K.N. and Krause J. (2006). Social structure and co-operative interactions in a wild population of guppies (Poecilia reticulata). Behavioural Ecology and Sociobiology 59: 644-650. 23. Croft D. P., James R., Ward A. J. W., Botham M. S., Mawdsley D. and Krause J. (2005) Assortative interactions and social networks in fish. Oecologia 143: 211-219. 22. Croft D. P., Botham M. S. and Krause J. (2004) Is sexual segregation in the guppy, Poecilia reticulata, consistent with the predation risk hypothesis? Environmental Biology of Fishes. 71:127-133. 21. Croft D. P., Krause J. and James R. (2004) Social networks in the guppy (Poecilia reticulata). Proceedings of the Royal Society of London Series B-Biological Sciences. 271: S516-S519. 20. Croft D. P., Arrowsmith B. J., Webster M. and Krause J. (2004) Intrasexual preferences for familiar individuals in male guppies (Poecilia reticulata). Journal of Fish Biology 64: 279-283. 19. Croft D. P., Albanese B., Arrowsmith B. J., Botham M., Webster M. and Krause J. (2003) Sex biased movement in the guppy (Poecilia reticulata). Oecologia 137: 62-68. 18. Croft D. P., Arrowsmith B. J., Bielby J., Skinner K., White E., Couzin I. D., Magurran A. E., Ramnarine I. and Krause J. (2003) Mechanisms underlying shoal composition in the Trinidadian guppy (Poecilia reticulata). Oikos 100: 429-438. 17. Hoare D. J., Ward A. J. W., Couzin I. D., Croft D. P. and Krause J. (2001) A grid-net technique for the analysis of fish positions within free-ranging shoals. Journal of Fish Biology 59: 1667-1672. 16. Krause J., Hoare D. J., Croft D.P., Lawrence J., Ward A., Ruxton G. D., Godin J. G. J. and James R. (2000) Fish shoal composition: mechanisms and constraints. Proceedings of the Royal Society of London Series B-Biological Sciences 267: 2011-2017. Review Articles in Journals

15. Croft D.P., Brent L.J.N., Franks D.W., Cant M.A. (2015). The evolution of prolonged life after reproduction. Trends in Ecology and Evolution. http://dx.doi.org/10.1016/j.tree.2015.04.011

14. *Rose P, Croft D.P. (2015). The potential of social network analysis as a tool for the management of zoo animals. Animal Welfare. In press

13. Kurvers R.H.J.M., Krause J, Croft D.P., Wilson ADM & Wolf M. (2014) The evolutionary and ecological consequences of animal social networks: emerging issues. Trends in Ecology and Evolution 29 : 326– 335

12 *Rose, P. E., Croft, D.P. and Lee. R. 2014 A review of captive flamingo (Phoenicopteridae) welfare: a synthesis of current knowledge and future directions. International Zoo Yearbook. 48: 139–155

11. Silk M.J., Croft D.P., Tregenza T., Bearhop S. (2014). The importance of fission-fusion social group dynamics in birds. Ibis, 156(4), 701-715.

10. Wilson, A.D.M., Croft, D.P., Krause, J. (2014). Social networks in elasmobranchs and teleost fishes. Fish and Fisheries. 15: 676–689. 12 Croft CV, May 2015

9. *Jacoby, D, Brooks, E. Croft D.P. & Sims, D. (2012) Developing a deeper understanding of animal movements and spatial dynamics through novel application of network analyses. Methods in Ecology and Evolution 3: 574-583. 8. Krause J., Wilson A.D.M. and Croft D.P. (2011) New technology facilitates study of social networks. Trends in Ecology and Evolution, 26: 5-6. 7. Krause J. James R. Croft D.P. 2010 Personality in the context of social networks. Philosophical Transactions of the Royal Society B: Biological Sciences 365: 4099-4106. 6. * Jacoby D.M.P., Croft D.P. and Sims D.W. (2011) The social behaviour of sharks and rays: patterns, implications and future research directions. Fish and Fisheries doi: 10.1111/j.1467-2979.2011.00436.x 5. Croft D.P., Madden J., Franks D. and James R. (2011) Hypothesis testing in animal social networks. Trends in Ecology and Evolution. 26: 502-507 4. James R. Croft D.P. and Krause J. (2009) Potential banana skins of analysing animal social networks. Behavioural Ecology and Sociobiology 63: 989–997 3. Krause J. Croft D.P. and James R. (2007) Social network theory in the behavioural sciences: potential applications. Behavioral Ecology and Sociobiology 62: 15 - 27. 2. Wright D, Ward A.J.W., Croft D.P. and Krause J. (2006). Social organisation, grouping and domestication in fish. Zebrafish, 3: 141-155. 1. Croft D. P., Krause J., Couzin I. D. and Pitcher T. J. (2003) When fish shoals meet: outcomes for evolution and fisheries. Fish and Fisheries 4:138-146.

Other Outputs How far will females go to avoid sexual harassment. NERC Planet Earth Autumn 2008.

Giraffe Necks and Networks by Bryan Shorrocks and Darren Croft. Mpala News, Issue 3, November, 2006.

13 Christopher W. Clark

CHRISTOPHER W. CLARK May 2015

Cornell University Born: 20 Sept. 1949 Laboratory of Ornithology PH: (607) 254-2408 159 Sapsucker Woods Rd. E-mail:[email protected] Ithaca, New York 14850

PRESENT POSITION Imogene P. Johnson Senior Scientist, Bioacoustics Research Program, Cornell Lab, Senior Scientist Department of Neurobiology and Behavior, Cornell University.

EDUCATION and EMPLOYMENT Stony Brook University, Stony Brook, NY B.Sc. 1972 Biology Stony Brook University, Stony Brook, NY B.E. 1972 Engineering Stony Brook University, Stony Brook, NY M.S. 1974 Electrical Engr. Stony Brook University, Stony Brook, NY Ph.D. 1980 Biology The Rockefeller University, NY, NY Post. Doc. 1981-83 Bio/Anim. Comm. The Rockefeller University, NY, NY Asst. Prof. 1983-87 Bio/Anim. Comm. Cornell University, Ithaca, NY Senior Sci. 1987-present CLO and NB&B

PROFESSIONAL SOCIETIES Acoustic Society of America Fellow Animal Behavior Society AAAS, IEEE Explorers Club (FN85) Society for Marine Mammalogy Sigma Xi, Tau Beta Pi

HONORS AND AWARDS Member, Tau Beta Pi, 1969 President, Tau Beta Pi, Stony Brook Chapter, 1971-1972 National Fellow, Tau Beta Pi, 1972-1973 Biomedical Research Fellowship, 1978, 1979 NIMH Postdoctoral Fellow, 1981-1983 Fellow, Acoustic Society of America, 2000

PROFESSIONAL APPOINTMENTS 1983 - 1987 Assistant Professor, The Rockefeller University, New York, NY 1985 – 2010 U.S. Delegate to the International Whaling Commission Scientific Committee 1987 - 2013 Director Bioacoustics Research Program, The Cornell Lab of Ornithology 2013 - present Senior Scientist, Cornell Lab and Dept. of Neurobiology and Behavior, Cornell University

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RECENT AND ONGOING RELEVANT RESEARCH 1996 – present-ongoing: Acoustic monitoring of large whale distributions, behaviors, and movements relative to environmental factors and man-made activities using Navy IUSS assets in the North Atlantic. DoD. 1999 – present-ongoing: Raven: Design, implementation, and distribution of bioacoustics software instrument. NSF and Cornell. 2007 – present-ongoing: Application of near-real-time auto-detection system for large whale acoustic monitoring and mitigation of Northeast Gateway Deepwater Port. Excelerate Energy and Suez Energy North America. 2010 – 2015: Portable and persistent autonomous real-time marine mammal acoustic monitoring. NOPP-NSF. 2011 – 2016: DCL System Research Using Advanced Approaches for Land-based or Ship- based Real-Time Recognition and Localization of Marine Mammals. NOPP-ONR. 2011 – 2016: CHAOZ-X - An Ocean Observing System for Monitoring and Mapping Marine Mammals and Noise in the Chukchi Sea Ecosystem. BOEM -NOAA-WHOI. 2013 – 2016: Acoustic ecology of predator-prey interactions: encoding and decoding alarm calls in multispecies communication networks. Collaborative research with Erick Greene, NSF.

PUBLICATIONS Clark, C.W., Berchok, C.L., Blackwell, S.B., Hannay, D.E., Jones, J., Ponirakis, D., and Stafford, K.M. 2015-Accepted. A year in the acoustic world of Western Arctic bowhead whales. Progress in Oceanography. Nowacek, D.P., Clark, C.W., Mann, D., Miller, P.J.O., Rosenbaum, H.C., Golden, J.S., Jasny, M., Kraska, J., and Southall, B.L. 2015 In press. Marine Seismic Surveys and Ocean Noise: Time for coordinated and prudent planning. Frontiers. Risch, D. Castellote, M., Clark, C.W., Davis, G., Dugan, P., Hodge, L.E.W., Kumar, A., Lucke, K., Mellinger, D.K., Nieukirk, S., Popescu, M., Ramp, C., Read, A.J., Rice, A.N., Silva, M.A., Siebert, U., Stafford, K.M., Verdaat, H and Van Parijs, S.M. 2015. Seasonal and geographic variation in North Atlantic minke whale pulse train occurrence: Insights into migration patterns and winter distribution. BioMed Central. Salisbury, D., Clark, C.W., and Rice, A.N. Right Whale Occurrence in Virginia Coastal Waters: Implications of Endangered Species Presence in a Rapidly Developing Energy Market. 2015 Accepted. Marine Mammal Science. Williams, R. Erbe, C., Ashe, E., and Clark, C.W. 2015 Accepted. Quiet(er) marine protected areas. Frontiers in Ecol. and Envir. Johnson, H.D., Stafford K.M., George, J.C, Ambrose W.G. Jr., and Clark, C.W. 2014. Song diversity in the Bering-Chukchi-Beaufort population of bowhead whales (Balaena mysticetus), spring 2011. Marine Mammal Science. doi: 10.1111/mms.12196. Murray, A., Rice, A., and Clark, C. 2014. Extended seasonal occurrence of humpback whales in Massachusetts Bay. Journal Marine Biological Association of the U. K. 94:1117-1125. Rice, A.N., Tielens, J.T., Palmer, K.J., Muirhead, A., and Clark, C.W. 2014. Potential Bryde’s Whale Calls (Balaenoptera edeni) Recorded in the Northern Gulf of Mexico. J. Acoust. Soc. Amer. 135:3066-3076. Rice, A. N., J. T. Tielens, B. J. Estabrook, C. A. Muirhead, A. Rahaman, M. Guerra, and C. W. Clark. 2014. Variation of ocean acoustic environments along the western North

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Atlantic coast: a case study in context of the right whale migration route. Ecological Informatics 21:89-99. Soldevilla, M. S., Rice, A. N., Clark, C. W., and Garrison, L. P. 2014. Passive acoustic monitoring on the North Atlantic right whale calving grounds. Endangered Species Research 25:115-140. Stafford, K.M., Rankin, S., Ljungblad, D., Tsuda, Y., Morse, L., Clark, C.W., and Kato, H. 2014. Acoustic studies during IWC-SOWER Antarctic cruises 1996-2009. J. Cet. Res. & Mgmt. Charif, R. A., Rahaman, A., Muirhead, C. A., Pitzrick, M. S., Warde, A. M., Hall, J. Pyc, C., and Clark, C. W. 2013. Bowhead whale acoustic activity in the southeast Beaufort Sea during late summer 2008-2010. J. Acoust. Soc. Am. 134:4323-4334. DeRuiter, S. T., Boyd, I. L., Claridge, D. E., Clark, C. W., Southall, B. L., and Tyack, P. L. 2013. Delphinid whistle production in relation to playback of simulated military sonar. Mar. Mamm. Sci.:E46-59. Risch, D., C. W. Clark, Dugan, P. J., Popescu, M., Siebert, U., and Van Parijs, S. M. 2013. Minke whale acoustics behavior and multi-year seasonal and diel vocalization patterns in Massachusetts Bay, USA. Mar. Ecol. Progr. Ser. 489: 279-295. Williams, R., Clark, C. W., Ponirakis, D. and Ashe, E. 2013. Acoustic quality of critical habitats for three threatened whale populations. Anim. Cons., Zool. Soc. London. Pp. 12. Ellison, W. T., Southall, B. L., Clark, C. W., and Frankel, A. 2012. A new context-based paradigm to assess behavioral responses of marine mammals to sound. Con. Bio. 26:21- 28. Castellote, M., Clark, C. W., and Lammers, M. O. 2102a. Fin whale (Balaenoptera physalus) population identity in the western Mediterranean Sea. Mar. Mamm. Science. 28:325-344. Castellote, M., Clark, C. W., and Lammers, M. O. 2102b. Acoustic and Behavioural Changes by Fin Whales (Balaenoptera physalus) in Response to Shipping and Airgun Noise. Biol. Cons. 147:115-122. Hatch, L. T., Clark, C. W., Van Parijs, S., Frankel, A. S., and Ponirakis, D. W. 2012. Quantifying loss of acoustic communication space for right whales in and around a U. S. National Marine Sanctuary. Con. Bio. 26:983-994. Moore, S.E., Reeves, R. R., Southall, B. L., Ragen, T. J., Suydam, R. S., and Clark, C. W. 2012. Marine mammals and anthropogenic sound in a rapidly changing Arctic. Bio. Sci. 62:289-295. Morano, J. L., Rice, A. N., Tielens, J. T., Estabrook, B. J., Murray, A., Roberts, B., and Clark, C. W. 2012a. Year-Round Presence of Right Whales in Massachusetts Bay: Acoustically Monitoring for a Critically Endangered Marine Mammal in a Highly Urbanized Migration Corridor. Con. Bio. 26:698-707. Morano, J. L., Salisbury, D. P. Rice, A. N., Conklin, K. L., Falk, K. L., and Clark, C. W. 2012b. Seasonal changes in fin whale (Balaenoptera physalus) 20-Hz song in the Western North Atlantic Ocean. J. Acoust. Soc. Am. 132:1207-1212. Parks, S. E., Hotchkin, C. F., Cortopassi, K. C., and Clark, C. W. 2012. Characteristics of gunshot sound displays by North Atlantic right whales in the Bay of Fundy. J. Acoust. Soc. Am. 131:3171-3179. Streever, W., Ellison, W. T., Frankel, A. S., Racca, R. Clark, C. W., Fleishman, E., Guerrra, M., and Sformo, T. 2012. Early progress and challenges in assessing aggregate sound exposure and associated effects on marine mammals. Soc. Petrol. Engr. Perth, AU, 11-13 Sep 2012, pp. 1-7.

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Vu, E. T., Risch, D., Clark, C. W., Gaylord, S., Hatch, L., Merrick, R., Thompson, M., Tougaard, J., Wiley, D. and Van Parijs, S. M. 2012. Humpback whale (Megaptera novaeangliae) song occurs extensively on feeding grounds in the northwestern Atlantic Ocean. Aqua. Bio. 14:175-183. Staaterman E. R., Clark, C. W., Gallagher, A. J., deVries, M. S, Claverie, T. and Patek, S. N. 2011. Rumbling in the benthos: the acoustic ecology of the California mantis shrimp. Aquat. Biol. 13:97-105. Clark, C. W., Brown, M. W. and Corkeron, P. 2010. Visual and acoustic surveys for North Atlantic right whales, Eubalaena glacialis, in Cape Cod Bay, Massachusetts, 2001-2005: Management implications. Marine Mammal Science 26: 837-854. Clark, C. W., Ellison, W. T., Southall, B. L., Hatch, L., Van Parijs, S., Frankel, A., and Ponirakis. 2009. Acoustic Masking in Marine Ecosystems: Intuitions, Analysis, and Implications. Mar. Ecol. Progr. Ser. 395:201-222. Di Iorio L., Clark C. W. 2009. Exposure to seismic survey alters blue whale acoustic communication. Biological Letters. Published online 23 September 2009. Moore, S.E., Stafford, K.M., Mellinger, D.K., and Clark, C.W. 2009. Insights into large whale ecology from broad-scale passive acoustic sampling. Mar. Ecol. Progr. Ser. 395: Parks, S., Urazghildiiev, I. and Clark, C. W. 2009. Variability in ambient noise levels and call parameters of North Atlantic right whales in three habitat areas. J. Acoust. Soc. Am., 125:1230-1239. Urazghildiiev, I. R., Clark, C. W., Krein T. P. and Parks, S. E. 2009. Detection and recognition of North Atlantic right whale contact calls in the presence of ambient noise. IEEE J. Ocean Engr. 34:358-368. Van Parijs, S., Clark, C.W., Sousa-Lima, R. S., Parks, S. E., Rankin, S., Risch, D., and Van Opzeeland, I. C. 2009. Management and research applications of real-time and archival passive acoustic sensors over varying temporal and spatial scales. Mar. Ecol. Progr. Ser. 395:21-36. Borsani, J. F., Christopher W. Clark, Barbara Nani, and Michele Scarpiniti. 2008. Fin Whales Avoid Loud Rhythmic Low-Frequency Sounds in the Ligurian Sea. Bioacoustics 17(1- 3):161-163. Hatch, L., Clark, C., Merrick, R., Van Parijs, S., Ponirakis, D., Schwehr, K., Thompson, M., and Wiley, D. 2008a. Characterizing the relative contributions of large vessels to total ocean noise fields: a case study using the Gerry E. Studds Stellwagen Bank National Marine Sanctuary. Environ. Management. 42:735-752. Hatch L, Asmussen I, Bahtiarian M, Clark C, Friedrich A, Friesch J, Hildebrand J, Hintzsche W, Kurth D, McKenna M, Metcalf K, Scovazzi T, Southall S, Stoye T, Thiele R, Weilgart L, Wittekind D Wright, AJ (ed). 2008b. International Workshop on Shipping Noise and Marine Mammals, Hamburg, Germany, 21st-24th April 2008. Okeanos - Foundation for the Sea, Auf der Marienhohe 15, D-64297 Darmstadt. Available from http://www.sound- in-the-sea.org/download/ship2008_en.pdf. Parks, S.E., Clark, C.W., Tyack, P.L. 2008. Long- and Short-term changes in right whale acoustic behaviour in increased low-frequency noise. Bioacoustics, 17:179-180. Parks, S.E., Clark, C.W. 2008. Acoustic Communication in Mysticetes. Bioacoustics,17:45- 47. Sousa-Lima, R. S. and Clark, C. W. 2008. Modeling the effect of boat traffic on the fluctuation of humpback whale singing activity in the Abrolhos National Marine Park, Brazil. Canadian Acoust. 36:174-181.

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Clark, C.W., Gillespie, D., Nowacek, D.P., and Parks, S.E. 2007. Listening to Their World: Acoustics for Monitoring and Protecting Right Whales in an Urbanized Ocean. In: The Urban Whale (eds. S. Kraus and R. Rolland). Harvard University Press, Cambridge, MA. pp. 333-357. Parks, S. E. and C. W. Clark. 2007. Acoustic Communication: Social sounds and the potential impacts of noise. In: The Urban Whale (eds. S. Kraus and R. Rolland). Harvard University Press, Cambridge, MA. pp. 310-332. Parks, S., Clark, C.W., and Tyack, P.L. 2007. Short and long-term changes in right whale calling behavior: the potential effects of noise on acoustic communication. J. Acoust. Soc. Am., 122:3725-3731. Risch, D., C. W. Clark, P. J. Corkeron, A. Elepfandt, K. M. Kovacs, C. Lydersen, I. Stirling, and S. M. Van Parijs. 2007. Vocalizations of male bearded seals, Erignathus barbatus: classification and geographical variation. Animal Behaviour 73: 747-762. Urazghildiiev, I., and C. W. Clark. 2007a. Detection performances of experienced human operators compared to a likelihood ratio based detector. J. Acoust. Soc. Am. 122 (1): 200– 204. Urazghildiiev, I., and C. W. Clark. 2007b. Acoustic detection of North Atlantic right whale contact calls using the spectrogram-based statistics. J. Acoust. Soc. Am. 122 (2):769–776. Clark, C.W., and Altman, N.S. 2006. Acoustic detections of blue whale (Balaenoptera musculus) and fin whale (B. physalus) sounds during a SURTASS LFA exercise. J. Ocean Engr. 31: 120-128. Mellinger, D. and C.W. Clark. 2006. MobySound: A reference archive for studying automatic recognition of marine mammal sounds. Appl. Acoust. 67:1226-1242. Urazghildiiev, I., and C. W. Clark. 2006. Acoustic detection of North Atlantic right whale contact calls using the generalized likelihood ratio test. J. Acoust. Soc. Am. 120 (4):1956- 1963. Van Parijs, S. and C.W. Clark. 2006. Long term mating tactics in an aquatic mating pinniped – the bearded seal, Erignathus barbatus. Animal Behavior, 72:1269-1277. Bower, J.L. and Clark, C.W. 2005. A field test of the accuracy of a passive acoustic location system. Bioacoustics 15: 1-14. Kraus, S., M. W. Brown, H. Caswell, C. W. Clark, M. Fujiwara, P. K. Hamilton, R. D. Kenney, A. R. Knowlton, S. Landry, C. A. Mayo, W. A. McLellan, M. J. Moore, D. P. Nowacek, D. A. Pabst, A. J. Read and R. M. Rolland. 2005. North Atlantic right whales in crisis. Science 309:561-562. Samuel, Y., Morreale, S.J., Clark, C.W., Greene, C.H., and Richmond, M.E. 2005. Underwater, low-frequency noise in a coastal sea turtle habitat, J. Acoust. Soc. Am. 117 (3): 1465-1472. Clark, C. W., and Clapham, P. J. 2004. Acoustic monitoring on a humpback whale (Megaptera novaeangliae) feeding ground shows continual singing into late Spring. Proceedings Roy. Soc. Lond., B. 271: 1051-1057. Clark, C. W., and Ellison, W.T. 2004. Potential use of low-frequency sounds by baleen whales for probing the environment: evidence from models and empirical measurements. Pp. 564- 582, in Echolocation in Bats and Dolphins (J. Thomas, C. Moss and M. Vater, eds.). The University of Chicago Press. Charif, RA, CW Clark, and KM Fristrup. 2004. Raven 1.2 User’s Manual. Cornell Laboratory of Ornithology, Ithaca, NY.

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George, J. C. “Craig”, Zeh, J., Suydam, R., and Clark, C. 2004. Abundance and population trend (1978-2001) of the western Arctic bowhead whales surveyed near Barrow, Alaska. Marine Mammal Science 20:755-773. Fristrup, K.M., Hatch, L.T. and Clark, C. W. 2003. Variation in humpback whale (Megaptera novaeangliae) song length in relation to low-frequency sound broadcasts. J. Acoust. Soc. Am., 113 (6): 3411-3424. Mellinger, D. K. and Clark, C. W. 2003. Blue Whale (Balaenoptera musculus) sounds from the North Atlantic. J. Acoust. Soc. Am. 114:1108-1119. Clark, C.W., Borsani, J.F. and Notarbartolo-di-Sciara, G. 2002. Vocal activity of fin whales, Balaenoptera physalus, in the Ligurian Sea. Mar. Mamm. Science 18(1): 281-285. Clark, C.W., and Gagnon, G.C. 2002. Low-frequency vocal behaviors of baleen whales in the North Atlantic: Insights from IUSS detections, locations and tracking from 1992 to 1996. J. Underwater Acoust. (USN), 52 (3):609-640. Bass, A.H. and Clark, C.W. 2002. The physical acoustics of underwater sound communication. In Acoustic Communication, edited by Simmons, Andrea M., Fay, Richard R., and Popper, Arthur N. (Springer, New York), pp. 15-64. Charif, R.A., Mellinger, D.K., Dunsmore, K.J., Fristrup, K.M. and Clark, C.W. 2002. Estimated source levels of fin whale (Balaenoptera physalus) vocalizations: adjustments for surface interference. Mar. Mamm. Science 18(1):81-98. Croll, D.A., Clark, C.W., Acevedo, A., Tershy, B., Flores, S., Gedamke, J. and Urban, J. 2002. Only male fin whales sing loud songs. Nature 417:809. Frankel, A.S. and Clark, C.W. 2002. ATOC and other factors affecting the distribution and abundance of humpback whales (Megaptera novaeangliae) off the coast of the north shore of Kauai. Mar. Mamm. Science 18 (3): 644-662. Charif, R.A., Clapham, P.J. and Clark, C.W. 2001. Acoustic detections of singing humpback whales in deep waters off the British Isles. Mar. Mamm. Science 17(4):751-768. Croll, D. A., Clark, C.W., Calambokidis, J., Ellison, W.T., and Tershy, B.R. 2001. Effect of anthropogenic low-frequency noise on the foraging ecology of Balaenoptera whales. Animal Conservation 4:13-27. Clark, C.W. and Ellison, W.T. 2000. Calibration and comparison of the acoustic location methods used during the spring migration of the bowhead whale, Balaena mysticetus off Pt. Barrow, Alaska, 1984-1993. J. Acoust. Soc. Am. 107(6):3509-3517. Frankel, A.S. and Clark, C.W. 2000. Behavioral responses of humpback whales (Megaptera novaeangliae) to full-scale ATOC signals. J. Acoust. Soc. Am. 108 (4):1930-1937. Mellinger, D .K., Carson, C.D. and Clark, C.W. 2000. Characteristics of minke whale (Balaenoptera acutorostrata) pulse trains recorded near Puerto Rico. Mar. Mamm. Science, 16(4):739-756. Mellinger, D.K. and Clark, C.W. 2000. Recognizing transient low-frequency whale sounds by spectrogram correlation. J. Acoust. Soc. Am. 107:3518-3529. Tyack, P. L. and Clark, C.W. 2000. Communication and acoustical behavior in dolphins and whales. Pp. 156-224, in Hearing by Whales and Dolphins. Springer Handbook of Auditory Research (W. W. L. Au, A. N. Popper, and R. R. Fay, eds.). Springer-Verlag, New York, 485 pp. Clark, C.W. 1998. Underwater noise. Pp. 415-419. In: 1998 McGraw-Hill Yearbook of Science and Technology, 464 pp. alternate title “Noise in the Ocean”

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Clark, C. W., Calupca, T., Charif, R., Corzilius, B., Fristrup, K. 1998. Autonomous seafloor acoustic recording systems for whale research: application to the census of bowhead whales during the spring migration off Point Barrow, Alaska. IWC 1998. The ATOC Consortium. 1998. Ocean Climate Change: Comparison of Acoustic Tomography, Satellite Altimetry and Modeling. Science 281:1327-1332. Frankel, A.S. and Clark, C.W. 1998. Results of low-frequency m-sequence noise playbacks to humpback whales in Hawai'i. Can. J. Zool. 76(3):521-535. Ljungblad, D. K. and Clark, C. W. 1998. Unique calls of the pygmy blue whale recorded off the coast of Chili in Dec 1997 and Jan 1998 during the SOWER 1997/98 program Ljungblad, D. K., Clark, C. W. and Shimada, H. 1998. A comparison of sounds attributed to pygmy blue whales (Balaenoptera musculus brevicauda) recorded south of the Madagascar Plateau and those attributed to ‘True” blue whales (Balaenoptera musculus) recorded off Antarctica. Rep. int. Whal. Commn. 49: 439-442. Clark, C.W. and Fristrup, K. 1997. Whales ‘95: A combined visual and acoustic survey of blue and fin whales off southern California. Rep. int. Whal. Commn. 47:583-600. Fristrup, K. and Clark, C.W. 1997. Combining visual and acoustic survey data to enhance density estimation. Rep. int. Whal. Commn. 47:933-936. McGregor, P.K., Dabelsteen, T., Clark, C.W., Bower, J.L., Tavares, J.P. and Holland, J. 1997. Accuracy of a passive acoustic location system: empirical studies in terrestrial habitats. Ethology Ecology and Evolution, Vol. 9 (3), pp. 269-286. Mellinger, D.K. and Clark, C.W. 1997. Methods for automatic detection of mysticete sounds. Marine and Freshwater Behaviour and Physiology. Mar. Fresh. Behav. Physiol.163-181. Clark, C. W., Charif, R., Mitchell, S., and Colby, J. 1996. Distribution and Behavior of the Bowhead Whale, Balaena mysticetus, Based on Analysis of Acoustic Data Collected During the 1993 Spring Migration off Point Barrow, Alaska. Rep. int. Whal. Commn. 46: 541-552. Spikes, C.H. and Clark, C.W. 1996. Whales '95 - Revolutionizing Marine Mammal Monitoring Technology. Sea Technology. April 1996. 49-53. Clark, C.W. 1995. Application of US Navy underwater hydrophone arrays for scientific research on whales. Annex M, Rep. int. Whal. Commn. 45:210-212. Frankel, A.S., Clark, C.W., Herman, L.M. and Gabriele, C.M. 1995. Spatial distribution, habitat utilization, movements and social interactions of humpback whales, Megaptera novaeangliae, off Hawai'i using acoustic and visual techniques. Can. J. Zool. 73:1134- 1146. Clark, C.W. 1994. Blue deep voices: Insights from the Navy’s Whales ë93 program. Whalewatcher 28 (1):6-11. Potter, J.R., Mellinger, D.K. and Clark, C.W. 1994. Marine mammal call discrimination using artificial neural networks. J. Acoust. Soc. Am. 96:1255-1262. Mellinger, D.K. and Clark, C.W. 1993. A method for filtering bioacoustic transients by spectrogram image convolution. Proc. IEEE Oceans '93:122-127. Weisburn, B.A., Mitchell, S.G., Clark, C.W. and Parks, T.W. 1993. Isolating biological acoustic transient signals. Proceedings of the Intl. Conference on Acoustics, Speech, and Signal Processing. 1:269-272. Würsig, B. and Clark, C.W. 1993. Behavior. In: J. Burns, J. Montague, and C.J. Cowles (eds.), The Bowhead Whale. Allen Press. Lawrence, Kansas. pp. 157-193. Zeh, J., Clark, C.W., George, J.C., Withrow, D., Carroll, G.M. and Koski, W.R. 1993. Current Population Size and Dynamics. In: J. Burns, J. Montague, and C.J. Cowles (eds.), The

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Bowhead Whale. Society for Marine Mammalogy, Special Publication Number 2. Allen Press. Lawrence, Kansas. pp. 409-481. Clark, C. W. 1991. Moving With the Heard. Natural History, March, pp. 38-42. Clark, C. W. 1991. Talking Heads, Natural History, March, p. 42. Clark, C.W. 1990. Acoustic behavior of mysticete whales. In: J. Thomas and R. Kastelein (eds.), Sensory Abilities of Cetaceans. Plenum Press. pp. 571-583. Tyack, P.L., Malme, C.I., Clark, C.W. and Bird, J.E. 1990. Reactions of migrating gray whales (Eschrichtius robustus) to industrial noise. J. Acoust. Soc. Am. pp.1-88. Clark, C.W. 1989. Call tracks of bowhead whales based on call characteristics as an independent means of determining tracking parameters. Report of the sub-committee on protected species and aboriginal subsistence whaling, Appendix. Rep. int. Whal. Commn. 39:111-112. Clark, C.W. and Ellison, W.T. 1989. Numbers and distributions of bowhead whales, Balaena mysticetus, based on the 1986 acoustic study off Pt. Barrow, Alaska. Rep. int. Whal. Commn. 39:297-303. George, J.C., Clark, C.W., Carroll, G.M. and Ellison, W.T. 1989. Observations on the ice- breaking and ice navigation behavior of migrating bowhead whales (Balaena mysticetus) near Point Barrow, Alaska, spring 1985. Arctic. 42:24-30. Clark, C.W. 1988. Singing in the ice. The Living Bird Quarterly. Vol., No. 4:10-16. Clark, C.W. and Ellison, W.T. 1988. Numbers and distributions of bowhead whales, Balaena mysticetus, based on the 1985 acoustic study off Pt. Barrow, Alaska. Rep. int. Whal. Commn. 38:365-370. Clark, C.W., Marler, P. and Beeman, K. 1987. Quantitative analysis of animal vocal phonology: an application to swamp sparrow song. Ethology. 76:101-115. Ellison, W.T., Clark, C.W. and Bishop, G.C. 1987a. Potential use of surface reverberation by bowhead whales, Balaena mysticetus, in under-ice navigation: preliminary considerations. Rep. int. Whal. Commn. 37:329-332. Ellison, W.T., Sonntag, R.M. and Clark, C.W. 1987b. Comparison of measured bowhead whale, Balaena mysticetus, migration parameters with results from the tracking algorithm. Rep. int. Whal. Commn. 37:309-312. Clark, C.W., Ellison, W.T., Beeman, K. 1986a. A preliminary account of the acoustic study conducted during the 1985 spring bowhead whale, Balaena mysticetus, migration of Point Barrow, Alaska, Rep. int. Whal. Commn. 36:311-316. Clark, C.W., Ellison, W.T. and Beeman, K. 1986b. Acoustic tracking of migrating bowhead whales. Oceans 86, IEEE Oceanic Eng. Soc., New York. pp. 341- 346. Ko, C., Zeh, J.E., Clark, C.W., Ellison, W.T., Krogman, B.D. and Sonntag R. 1986. Utilization of acoustic location data in determining a minimum number of spring- migrating bowhead whales unaccounted for by the ice-based visual census. Rep. int. Whal. Commn. 36:325-338. Sonntag, R.M., Ellison, W.T., Clark, C.W., Corbit, D.R. and Krogman, B.D. 1986. A description of a tracking algorithm and its implications to bowhead whale acoustic location data collected during the spring migration near Point Barrow, Alaska 1984-1985. Rep. int. Whal. Commn. 36:299-310. Watkins, W.A., Moore, K.E., Clark, C.W. and Dahlheim, M.E. 1986. The sounds of sperm whale calves. In: P.E. Nachtigall and P.W.B. Moore (eds.), Animal Sonar; Processes and Performance. Plenum Press, New York. pp. 99-107.

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Beeman, K., Clark, C.W., Miller, R. and Bunnell, T. 1985. Acoustic tracking of migrating whales. Horizon 9(2). Clark, C.W. 1984. Acoustic communication and behavior of southern right whales, Eubalaena australis. In: National Geographic Society Research Reports, vol. 17: 897-908. Clark, C.W. and Johnson, J.H. 1984. The sounds of the bowhead whale, Balaena mysticetus, during the spring migrations of 1979 and 1980. Can. J. Zool. 62:1436-1441. Clark, C.W. 1983. Acoustic communication and behavior of the southern right whale. In: Behavior and Communication of Whales. (ed. R.S. Payne), Westview Press: Boulder, CO. pp. 163-198. Clark, C.W., Dooling, R.J., and Bunnell, T. 1983. Analysis and synthesis of bird vocalizations: An FFT-based software system. Behav. Res. Methods and Instr. 15:251- 253. Dooling, R.J., Clark, C.W., Miller, R. and Bunnell, T. 1983. Program package for the analysis and synthesis of animal vocalizations. Behav. Res. Methods and Instr. 14:487. Clark, C.W. 1982. The acoustic repertoire of the southern right whale: a quantitative analysis. Anim. Behav. 30:1060-1071. Clark, C.W. 1980. A real-time direction finding device for determining the bearing to the underwater sounds of Southern Right Whales, Eubalaena australis. J. Acoust. Soc. Am. 68:508-511. Clark, C.W., and Clark, J.M. 1980. Sound playback experiments with southern right whales (Eubalaena australis). Science 207:663-665.

Submitted Papers Cholewiak, D., Cerchio, S., Jacobsen, J, Urban-Ramirez, J., and Clark, C.W. Acoustic and spatial interactions between singing humpback whales (Megaptera novaeangliae) suggest mechanisms that may mediate male competition. Anim. Behav. Clark, C.W., and Gagnon, G.C. 2015. Fin Whale Swimming Speed and Song Variability. Journal of Underwater Acoustics, USN declassified, so I can now submit to unclassified journal.

GRADUATE FIELD MEMBERSHIPS Neurobiology and Behavior Zoology

GRADUATE MAJORS (serve as chair) Total Completed John Bower (NB&B, Ph.D.2000) Bernard Brennan (NB&B, Ph.D., 2005) Renata de Sousa Lima Mobley (Zool. - DNR, Ph.D., 2007) Danielle Cholewiak (NB&B, Ph.D., 2008) Mya Thompson (NB&B, Ph.D., 2008)

GRADUATE Ph.D. MINORS (member of graduate committee) • Total Completed Peter Marchetto (2015)

9 Christopher W. Clark

Ingrid Biedron (2014) Dan Pendleton (2009) Lynn Fletcher (2008) Yianna Samuel (2007) Damian Elias (2007) Andrew Farnsworth (2006) Leila Hatch (2005) Karen Fisher (2002) Hamilton Farris (2000) Matt Weeg (2001) Paul Faure (1999) David Haskell (1998) Jessica McKibben (1998) Andrea Lee (1997) Beth Weisburn (1996) Stacey Benton (1999) David Haskell (1996) Adam Frankel (1994)

- Curriculum Vitae - Short Form PAUL C. PAQUET, Ph.D.

Address: P.O. Box 150, Meacham, SK Canada Telephone No.: (306) 376 2015 Email: [email protected] Date of Birth: 06 December 1948 Citizenship: Canadian, U.S.A.

Education

1966 - 1970 B.A., Philosophy, University of Santa Clara, Santa Clara, California. 1967 - 1968 Undergraduate, Zoology, University of California, Berkeley, California. 1970 - 1970 Graduate Philosophy, Oxford University, Oxford, England. 1970 - 1971 Post Bacc., Biology, Portland State University, Portland, Oregon. 1971 - 1972 Graduate Studies, Wildlife Management, Oregon State University, Corvallis, Oregon. 1972 - 1974 B.S., Zoology (Wildlife Biology), Arizona State University, Tempe, Arizona. Thesis topic: Distribution and movements of coyotes along the lower Colorado River, Arizona. 1976 - 1981 M.S., Biology (Ethology), Portland State University, Portland, Oregon. Thesis topic: Temporal and phenomenological aspects of social behavior in captive wolves. 1982 - 1988 Ph.D., Zoology, University of Alberta, Edmonton, Alberta. Dissertation topic: Behavioural ecology of sympatric wolves and coyotes in Riding Mountain National Park, Manitoba. 1989-1990 Postdoc, Yale University, New Haven, CT

Academic Appointments

1988/89 Assistant Professor, Department of Biology, Brandon University, Brandon, Manitoba 1989-2012 Adjunct Professor, Department of Biology, Brandon University, Brandon, Manitoba 1995-2006 Faculty Associate, Faculty of Graduate Studies, Guelph University, Guelph, Ontario 1994-2012 Adjunct Associate Professor, Faculty of Environmental Design, University of Calgary, Calgary, Alberta 1997-2001 Adjunct Professor, Department of Biology, University of Calgary, Calgary, Alberta 1997-2001 Adjunct Professor, Department of Zoology, University of Alberta, Edmonton, Alberta Paquet, P.C. 2

2003-2012 Adjunct Professor, Dept. of Environment and Geography, University of Manitoba, Winnipeg 2003-present Honorary Research Associate, Department of Biology, University of New Brunswick, Fredericton 2006-2009 Adjunct Professor, Department of Veterinary Pathology, Western College of Veterinary Medicine, University of Saskatchewan 2012 Adjunct Professor, Department of Geography, University of Victoria, Victoria, BC

Other Appointments

2000-2008 Senior Scientist, Conservation Biology Institute, U.S.A. 1995-2009 Research Fellow, World Wildlife Fund – Canada 2000-present Senior Scientist, Raincoast Conservation Society, Canada 2005-present Research Fellow, Rewilding Institute, U.S.A. 2005-present Society for Conservation Biology – International Policy Committee 2007-2009 Wildlife Society Canada – Policy Committee

Certification

1972 1977 Instructor, Outward Bound Schools; Kayaking, Snow, Ice and Rock Climbing, Mountaineering 1982 "Animal damage control in coastal forests of the Pacific Northwest." Cooperative Extension, U.S. Fish & Wildlife Service, Department of Interior. 1984 "Biotelemetry, methods and applications." Telonics Inc., Mesa, Arizona in conjunction with Lethbridge Community College, Lethbridge, Alberta. 1987 Instructor's Certification "Safety in bear country", Departments of Natural Resources, N.W.T. and Manitoba. 1988 "Wildlife Immobilization", Canadian Parks Service, Environment Canada

Professional Societies

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1974-present The Wildlife Society, Washington D.C. 1974-2010 The American Society of Mammalogists, Kansas City, Missouri 1984-1989 The Wildlife Society of Canada 1984-present Canadian Field Naturalists 1985-1999 Canadian Society of Zoologists 1989-present Society for Conservation Biology 1992-2004 New York Academy of Science 1994-present American Association for the Advancement of Science 2003-present American Institute of Biological Sciences

Committees and Boards

1986-1991 Riding Mountain Biosphere Reserve, Chairman, Manitoba 1989-2009 Central Canadian Rockies Wolf Project, Board of Directors, Canmore, AB 1991-1999 Board of Scientific Advisors, Canid Research Facility, Dalhousie University, Nova Scotia 1992-2008 Coordinator for World Wildlife Fund (Canada) Carnivore Conservation Program 1992-present Board of Directors, German Wolf Society, Cologne, Germany 1993-1999 Banff National Park Elk Advisory Committee, Banff National Park, Alberta 1993-1999 Scientific Advisor, European Wolf Federation, Liege, Belgium 1993-1999 Director of Gray Wolf Research, Tatra Mountains National Park Wolf Ecology Project, Tatra, Slovakia 1993-1999 Director of Gray Wolf Research, Slovakian Wolf Ecology Project, Slovakia 1994-1998 Advisory Board of the Environmental Research Center, University of Calgary, Calgary, AB. 1993-2001 Predator/Prey Study Steering Committee and Director of Research, Pukaskwa National Park, Pukaskwa, Ontario 1993-2005 East Slopes Steering Committee for Grizzly Bear Research in Central Canadian Rocky Mountains 1995-2000 Grizzly Bear Scientific Advisory Committee to Minister of Environment, British Columbia Provincial Grizzly Bear Strategy 1995-2000 Scientific Steering Committee, World Wild Fund for Nature - International, Pan-European Carnivore Conservation Strategy 1994-1999 Town of Canmore Wildlife Corridor Committee, Canmore Alberta. 1996 Columbia Basin Science Review Panel, U.S. Forest Service

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1995 Tongas National Forest, Alaska - Scientific Review Committee. U.S. Forest Service 1997-2008 Northwest Territories, Ecology of Wolves in the Central Arctic, N.W.T., Research Associate, Yellowknife, Northwest Territories. 1997 Nez Pearce Wolf Recovery Scientific Advisory Committee, Idaho 1997-2003 Board or Directors and Founding Member, Conservation Biology Institute, Corvallis, Oregon. 1997 U.S. Forest Service Lynx Research Advisory Committee, Missoula, Mt. 1995-1997 Riding Mountain National Park Management Plan Science Advisory Committee, Riding Mountain National Park, Manitoba. 1998-2006 Animal Care Committee, American Society of Mammalogists 1999-2005 U.S. Fish & Wildlife Service, Red Wolf Reintroduction Advisory Committee 1996-present Parks and Wilderness Specialist Group; International Union for Conservation of Nature 2000-present Conservation Breeding Specialist Group, International Union for Conservation of Nature Species Specialists, Member and Strategic Associate 2002-2009 Board of Directors, Defenders of Wildlife Canada 2003-2007 Board of Directors, Wildlands League, USA 2004-2008 Chair, Science Advisory Committee, Infectious Wildlife Disease National Strategy, Canada 2004-present Chair, National Science Advisory Committee, for Wildlife Tuberculosis, Riding Mountain, Manitoba 2008-present Board of Directors, Society for Conservation biology 2012-present Woodland Caribou Technical Committee, Province of Saskatchewan 2014 Canid Specialist Group, International Union for Conservation of Nature Species Specialists, Member and Strategic Associate

Periodicals and Reports

Paquet, P.C. 1974. Small mammals of the Colorado River islands (Arizona). Arizona Fish and Game Bull. 56.

Paquet, P.C. 1974. Coyote literature review. Special report, Arizona State University Press, Tempe, Ariz.

Paquet, P.C. 1976. Wolves. Portland Zoological Society Magazine, Portland Zoological Gardens, Portland, Oregon.

Paquet, P.C., and J.O. Sullivan. 1977. Behavior of wild canids: Social systems of wolves in large and small enclosures. Proc. Amer. Assoc. Zool. Parks & Aquar., Hills Co., Topeka, Kansas.

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Paquet, P.C., and J.O. Sullivan. 1978. Management of wolves in captivity. Proc. W. Psychol. Assoc., San Francisco State Univ., San Francisco, California.

Paquet, P.C. 1981. Behavior of Domesticated Dogs. Pamphlet, Multnomah County Animal Control, Multnomah County, Oregon, 12 pp.

Carbyn, L.N., D.M. Meleshko, and P.C. Paquet. 1983. Ecological studies of wolves and coyotes in Riding Mountain National Park, Manitoba. Canadian Wildlife Service Annual Report: 15 June 1982 to 31 March 1983. 60 pp.

Meleshko, D.M., and P.C. Paquet. 1983. Annual classified ungulate survey, Riding Mountain National Park, Manitoba. Canadian Wildlife Service Report, Large Mammal System Study Series.

Paquet, P.C., and D.M. Meleshko. 1984. Annual classified ungulate survey, Riding Mountain National Park, Manitoba. Canadian Wildlife Service Report.

Paquet, P.C., D.M. Meleshko, and L.N. Carbyn. 1985. Annual classified ungulate survey, Riding Mountain National Park, Manitoba. Canadian Wildlife Service Report.

Carbyn, L.N., Paquet, P.C., and D.M. Meleshko. 1985. Ecological studies of wolves and coyotes in Riding Mountain National Park, Manitoba. Canadian Wildlife Service Annual Report: 1 April 1983 to 31 December 1984.

Pesendorfer, J., P. C. Paquet, and L. N. Carbyn. 1986. A comparison of the 1984 and 1985 nocturnal howling surveys, RMNP, Manitoba. LMSS Series, Rep. No. 18.Carbyn, L. N., D. Meleshko, P. C. Paquet, S. Mckinlay, D. Burles, and J. Pessendorfer. 1986. Ecological studies of wolves and coyotes in Riding Mountain National Park, Manitoba. Canadian Wildlife Service Progress Report 3.

Carbyn, L.N., and P.C. Paquet. 1986. Long distance movement of a coyote from Riding Mountain National Park. Journal of Wildlife Management 50:89.

Meleshko, D.M., and P.C. Paquet. 1986. Annual classified ungulate survey, Riding Mountain National Park, Manitoba. Canadian Wildlife Service Report.

Paquet, P.C. 1986. Wolves revisited. Nature Canada 15:54.

Paquet, P.C., and L.N. Carbyn. 1986. Wolves, Canis lupus, killing denning black bears, Ursus americanus, in the Riding Mountain National Park area. Canadian

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Field-Naturalist 100:371-372.

Pesendorfer J., P.C. Paquet, and L.N. Carbyn. 1986. A comparison of the 1984 and 1985 nocturnal howling surveys, Riding Mountain National Park, Manitoba. Canadian Wildlife Service Report No. 18, Large Mammal System Study Series, 18 pp.

Carbyn, L.N., P.C. Paquet, and D.M. Meleshko. 1987. The Wolves of Riding Mountain. Canadian Wildlife Service Report, Large Mammal System Study Series.

Paquet, P.C. 1987. Canid howling survey. Riding Mountain National Park, Manitoba. Canadian Wildlife Service Report, Large Mammal System Study Series.

Paquet, P.C., Crichton, V.F.J., and Wotten. D.L. 1988. A study of cadmium levels in Manitoba wildlife. 1987 Progress Report. Prepared for Environment Canada and World Wildlife Fund Canada, Toxicology Fund.

Killiaan, H.P.L., Mamo, C., and Paquet, P.C. 1991. A coyote and badger observed together near Cypress Hills Provincial Park, Alberta. Canadian Field-Naturalist.

Gibeau, M, and Paquet, P.C. 1991. Evaluation of telezol for immobilization of black bears. Wildlife Society Bulletin 19:400-402.

Paquet, P.C. 1991. Winter spatial relationships of wolves and coyotes in Riding Mountain National Park, Manitoba. Journal of Mammalogy 72:397-401.

Paquet P.C. 1991. Black bear ecology in the Riding Mountains. Final report. Prepared for Manitoba Natural Resources and Canadian Park Service by John/Paul & Associates. 143 pp.

Paquet, P.C. 1991. Scent marking behavior of sympatric wolves (Canis lupus) and coyotes (C. latrans) in Riding Mountain National Park. Canadian Journal of Zoology 69:1721-1727.

Irwin, G.H., Purves, H., Paquet, P.C., and J.G. Woods. 1992. An intervener submission to the Natural Resource Board of Alberta on behalf of Environment Canada relating to application #9103 by Three Sisters Golf Resorts Inc. Recreational and Tourist Development in the Bow-Canmore Corridor. Environment Canada, Banff, AB. 72pp.

McCallum, M.J. and P.C. Paquet. 1992. Preliminary wildlife study and analysis of ungulate travel corridors on CADCO property, Canmore, Alberta. Prepared for Canmore Alpine Development Company Ltd. 72pp.

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Paquet, P.C. 1992. Prey use strategies of sympatric wolves and coyotes in Riding Mountain National Park, Manitoba. Journal of Mammalogy 73:337-343.

Purves, H.D., White, C.A., and P.C. Paquet. 1992. Wolf and grizzly bear habitat use and displacement by human use in Banff, Yoho, and Kootenay National Parks: a preliminary analysis. Canadian Parks Service, Banff. AB. 54pp.

McCallum, M.J. and P.C. Paquet. 1993. Stone Creek Properties Inc. wildlife management plan, Canmore, Alberta. Prepared for Stone Creek Properties by McCallum Paquet Associates. 95pp.

McTavish, C. and Paquet, P.C. 1993. Behavioral ecology of gray wolves in Kootenay National Park, British Columbia. Annual Report. Prepared for Parks Canada.

Gibeau, M.L., Paquet, P.C., Herrero, S., J. Jorgenson, and G. Green. 1994. Wildlife corridors in the Bow Valley, Alberta: an analysis. Prepared for the Inter- government Committee on Wildlife Corridors. Canmore, AB. 37pp.

Keister, R.A., Eckhardt, C., Benkman, C.W., Hansen, AJ., Houck, O. Jarvis, R.L., Lande, R., Lidicker, W.Z., McCullough, D.R., McLellan, B., Marcot, B., Mariani, J., Paquet, P.C., Pletscher, D., Powell, R., Ratti, J., Scott, J.M., Schaffer, M.L., Smith, C.C., Taber, R.D., Walters, J. 1994. Wildlife management and conservation biology on the Tongass National Forest: A synthentic review with recommendations. Pacific Northwest Research Station. USDA Forest Service, Corvallis, Ore.

Krizan, P., Neale, G., Paquet, P.C. 1994. Behavioral ecology of gray wolves in Pukaskwa National Park, Ontario, Annual Report. Prepared for Parks Canada.

Pacas, C., and Paquet, P.C. 1995. Analysis of black bear home range using a geographic information system. International Conference on Bear Research and Management. 9:419-425.

McCallum, M.J. and P.C. Paquet. 1995. Special features in Yoho National Park, MT. Prepared for Parks Canada. Yoho National Park, B.C.

McCallum, M.J. and P.C. Paquet. 1995. 1994 Wildlife Monitoring on the SilverTip property, Canmore, AB. Prepared for Stone Creek Properties by McCallum Paquet Associates, Canmore, Alberta. 47pp.

Paquet, P.C., and Hackman, A. 1995. A large carnivore conservation strategy for the Rocky Mountains. World Wildlfie Fund (Canada) and World Wildlife Fund

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(USA). 52pp.

Alexander, S., C. Callaghan, P.C. Paquet and N. Waters. 1996. GIS predictive models for habitat use by wolves (Canis lupus). Conference Proceedings: CD-ROM. GIS '96: Ten Years of Excellence. Vancouver, BC. March 17-20, 1996.

Clark, T.W., Paquet, P.C., and Purlee, P. (eds) 1996. Large Carnivore Conservation in the Rocky Mountains of the United States and Canada. Special Section. Conservation Biology 10:936-1055.

Clark, T.W., Paquet, P.C., and Curlee, P.A. 1996. Introduction: Special Section: Large Carnivore Conservation in the Rocky Mountains of the United States and Canada. Conservation Biology 10:936-939.

Clark, T.W., Paquet, P.C., and Curlee, P.A. 1996. General lessons and positive trends in large carnivore Conservation: Special Section: Large Carnivore Conservation in the Rocky Mountains of the United States and Canada. Conservation Biology 10:1055-1058.

P.C. Paquet and C. Apps. 1996. Section 3.5.3 on gray wolves. In Herrero, S. and Herrero, J. Cheviot Mine Project: Specific and cumulative environmental effects analysis for mammalian carnivores. Prepared for Cardinal River Coal Ltd. Hinton, AB. 122pp

McCallum, M.J. and P.C. Paquet. 1996. 1995 Wildlife Monitoring on the SilverTip property, Canmore, AB. Prepared for Stone Creek Properties by McCallum Paquet Associates, Canmore, Alberta. 47pp.

Noss, F. R., Quigley, H.B., Hornocker, T.M., and Paquet, P.C. 1996. Conservation biology and carnivore conservation in the Rocky Mountains. Conservation Biology 10:949-963.

P5 Study. 1996. Interim report on ecology of gray wolves and associated prey species in the Greater Pukaskwa Ecosystem, Ontario, 1994-1996. Prepared by F. Burrows, P. Krizan, G. Neale, and P. Paquet for Pukaskwa National Park, ON.

Weaver, J.L., Paquet, P.C., and L.F. Ruggierio. 1996. Resilience and Conservation of Large Carnivores in the Rocky Mountains. Conservation Biology 10:964-976.

Alexander, S., P.C. Paquet and N. Waters. 1997. Playing God with GIS: Uncertainty in Wolf Habitat Suitability Models. Conference Proceedings: GIS World '97. Vancouver, BC. February 17-20, 1997.

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Sawyer, M.D., D. Mayhood, P. Paquet, C. Wallis, R. Thomas, and W. Haskins. 1997. Southern East Slopes Cumulative Effects Assessment Project. Prepared for Morrison Petroleum Ltd. Calgary, AB. 291 pp.

Donelon, S. & P.C. Paquet. 1997. Technical report on development of a decision support model for the Lake O’hara socio-ecologic study. A Background report for the GIS decision-support model for the Lake O’Hara/McArthur Valley Socio- ecological Study. Parks Canada.

McRory, W. McTavish, C. and P.C. Paquet. 1998. Grizzly bear background research document1993-1996 for GIS bear encounter risk model, Yoho National Park, B.C. A Background report for the GIS decision-support model for the Lake O’Hara/McArthur Valley Socio-ecological Study. Parks Canada.

Paquet, P.C. 1998. Kootenay/Yoho Summary Report. Parks Canada (Title and publication date has been revised by Parks Canada; requires update)

Crichton, V. and P.C. Paquet. 1999. Cadmium in Manitoba’s Wildlife. Alces 36:205- 216.

Paquet, P.C., Noss, R.F., Carroll, C., and S. Alexander. 1999. Conservation biology primer and carnivores as conservation species. Prepared for Sierra Club of British Columbia and Central Coast LRMP.

Wilson, P. J., S. Grewal, I. D. Lawford, J. N. M. Heal, A. G. Granacki, D. Pennock, J. B. Theberge, M. T. Theberge, D. R. Voigt, W. Waddell, R. E. Chambers, P. C. Paquet, G. Goulet, D. Cluff, and B. N. White. 2000. DNA profiles of the eastern Canadian wolf and the red wolf provide evidence for a common evolutionary history independent of the gray wolf. Canadian Journal of Zoology 78:1-11.

Carroll, C., Noss, R.F., and P.C. Paquet. 1999. Carnivores as focal species for conservation planning in the Rocky Mountain region. World Wildlife Fund Canada. Toronto, Ontario, Canada. 61pp.

Carroll, C., Paquet, P.C., & R.F. Noss. 1999. Modeling carnivore habitat in the Rocky Mountain region. A literature review and suggested strategy. World Wildlife Fund Canada. Toronto, Ontario, Canada. 104pp.

Terborgh, J., J. Estes, P. C. Paquet, K. Ralls, D. Boyd-Heger, B. Miller, and R. Noss. 1999. The role of top carnivores in regulating terrestrial ecosystems. Wild Earth 9:42-56.

Darimont, C.T. and P.C. Paquet. 2000. The gray wolves (Canis lupus) of British

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Columbia’s coastal rainforests: Findings from year 2000 pilot study and conservation assessment. Prepared for Raincoast Conservation Society. Victoria, BC. 62pp.

Carroll, C., Noss, R.F., and P.C. Paquet. 2001. Carnivores as focal species for conservation planning in the Rocky Mountain Region. Ecological Applications 11:961-980.

Walton, H,. D. Cluff, P. C. Paquet, and M.A. Ramsay. 2001. Performance of two models of satellite collars for wolves. Wildlife Society Bulletin 29:180-186.

Walton, H,. D. Cluff, P. C. Paquet, and M.A. Ramsay. 2001. Movement patterns of barren-ground wolves in the Central Canadian arctic. Journal of Mammalogy 82:867-876.

Paquet, P.C., Vucetich, J.A. Phillips, M.K. and L.M. Vucetich et al. 2001. Mexican wolf recovery: three-year program review and assessment. Prepared by the Conservation Breeding Specialist Group (CBSG), Apple Valley, Minnesota; prepared for the United States Fish and Wildlife Service, Albuquerque, New Mexico. 86pp.

Fascione, N., Osborn, L.G.L., Kendrot, S.R., and P.C. Paquet. 2001. Canis soupus: Eastern wolf genetics and its implications for wolf recovery in the Northeast United States. Endangered Species Update 18:159-163.

Paquet, P.C., Burrows, F., Forshner, A., Neale, G., and K. Wade. 2002. Blastomycosis in a free ranging wolves, Canis lupus, on the North Shore of Lake Superior, Ontario - a response to P. Krizan. Canadian Field Naturalist 115:185.

Paquet, P.C. and Darimont, C.T. 2002. Yeo Island Wolf Home Site Recommendation A Proposed Solution To The Potential Conflict Between The Home Site Requirements Of Wolves And Areas Targeted For Timber Harvest Prepared For The Heiltsuk Nation, Western Forest Products, Raincoast Conservation Society.

Alexander, S.M., Duke, D.L., and P.C. Paquet. 2002. Modeling carnivore habitat-use, travel patterns and human activity around the town of Canmore. Prepared for the Wilburforce Foundation, Seattle, WA.

Carroll, C., Noss, R.F., and Paquet, P.C. 2002. Rocky Mountain carnivore project final report. Prepared For World Wildlife Fund Canada, Toronto, ON.

Hebblewhite, M., D.H. Pletscher, and P.C. Paquet. 2002. Elk population dynamics in areas with and without predation by recolonizing wolves in Banff National Park,

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Alberta. Canadian Journal of Zoology 80:789-799.

Darimont, C.T. and P.C. Paquet. 2002. The Gray Wolves, Canis lupus, of British Columbia’s Central and North Coast: distribution and conservation assessment. Canadian Field-Naturalist 116:416-422.

McCrory, W., Paquet, P., and Cross, B. 2003. Assessing conservation values for the gray wolf and sitka deer of the B.C. Central Coast Rainforest.

Cluff, D., M. Musiani, P.F. Frame, P. Paquet, and C.C. Gates. 2003. Multiple litters at den sites, Northwest Territories Wolf Notes 7:1-8.

Darimont, C., Gerwing, K, Paquet, P., Reid, B., McCrory, W. and Cross, B. 2003. Preliminary modeling of deer winter range in Heiltsuk Territory on the Central Coast of British Columbia. (Summary reference document to accompany maps) Prepared for the Heiltsuk Nation and Raincoast Conservation Society, British Columbia.

Darimont, C.T., T.E. Reimchen and P.C. Paquet. 2003. Foraging behaviour by gray wolves on salmon streams in coastal British Columbia. Canadian Journal of Zoology 81:349-353.

Hebblewhite, M.D., Paquet, P.C., and D.H. Pletscher. 2003. Development and application of a ratio-estimator to estimate wolf killing rates and variance in a multiple prey system. Wildlife Society Bulletin 31:933–946

Hebblewhite, M.D., D.H. Pletscher, and Paquet, P.C. 2003. Elk population dynamics after wolf recolonization in the Bow Valley of Banff National Park. Research Links 11:1-13

Carroll, C., Noss, R.F., Paquet, P.C., and Schumaker, N.H. 2003. Integrating population viability analysis and reserve selection algorithms into regional conservation plans. Ecological Applications 13: 1773–1789.

Musiani, M. and P.C. Paquet. 2004. Wolf Persecution, Protection, and Restoration in Canada and the United States. Bioscience, 54:50-60.

Gilbert, B., Craighead, L., Horejsi, B., Paquet, P., and W. McCrory. 2004. Scientific criteria for evaluation and establishment of grizzly bear management areas in British Columbia. Panel of Independent Scientists, Victoria, B.C. 16pp.

Paquet, P.C., C.T. Darimont, R. J. Nelson, and K. Bennett. 2004. A critical assessment of protection for key wildlife and salmon habitats under the

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proposed British Columbia Central Coast Land and Resource Management Plan. Raincoast Conservation Society. 23pp.

Carroll, C., Noss, R.F., Paquet, P.C., and Schumaker, N.H. 2004. Extinction debt of protected areas in developing landscapes. Conservation Biology 18: 1110-1120.

McLoughlin, P.D., Walton, L., Cluff, H.D., Paquet, P. 2004. Hierarchical habitat selection by tundra wolves. Journal of Mammalogy 85:576-580.

Frame, P.F., D.S. Hik, H.D. Cluff, and P.C. Paquet. 2004. Long foraging movement of a denning tundra wolf. Arctic 58: 196-203.

Forshner, S. A., P. Paquet, F. G. M. Burrows, G. K. Neale, and K. D. Wade. 2004. Demographic patterns and limitations of wolves (Canis lupus) in and near Pukaskwa National Park, Ontario, Canada. Canadian Field Naturalist 118:95- 104.

Carroll, C., Noss, R.F., and P.C. Paquet. 2004. A proposed wildlands network for carnivore conservation in the Rocky Mountains. Wild Earth, Spring: 38-41

Darimont, C. T., M. H. H. Price, N. N. Winchester, J. Gordon-Walker and P. C. Paquet. 2004. Predators in natural fragments: foraging ecology of wolves in British Columbia’s central and north coast archipelago. Journal of Biogeography 31: 1867–1877

Paquet, P. C., C. T. Darimont, C. Genovali, and F. Moola. 2004-2005. In the Great Bear Rainforest: island-hopping gray wolves give new insights into island biogeography. Wild Earth Fall/Winter: 20-25

Paquet, P., C. Darimont, C. Genovali, and F. Moola. 2005. Last of the best. Wolf Print. The UK Wolf Conservation Trust. Issue 24: 7-9.

Paquet, P. C. and S. M. Alexander. 2004-2005. Banff National Park: how helpful is highway mitigation? Wild Earth Fall/Winter: 70-71.

Paquet, P. C. and R. K. Brook. 2005. Island use as an anti-predator tactic by parturient elk and nursery herds in Riding Mountain National Park, Manitoba. Wildlife Society Bulletin.

Darimont, C. T., P. C. Paquet, T. E. Reimchen, and V. Crichton. 2005. Range expansion by moose into coastal temperate rainforests of British Columbia, Canada. Diversity and Distributions 11: 235–239

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Jickling, R., and P.C. Paquet. 2005. Wolf stories: Reflections on science, ethics, and epistemology. Environmental Ethics 27:115-134.

Alexander, S. M., P. C. Paquet, T. B. Logan, and D. J. Saher. 2005. Snow-tracking versus radio-telemetry for predicting wolf-environment relationships in the Rocky Mountains of Canada. Wildlife Society Bulletin 33: 1216-1224.

Hebblewhite, M., C. White, C. Nietvelt, J. Mckenzie, T. Hurd, J. Fryxell, S. Bayley, and P. C. Paquet. 2005. Human activity mediates a trophic cascade caused by wolves. Ecology 86: 1320–1330

Price, M. H., C. T. Darimont, N. N. Winchester, and P. C. Paquet. 2005. Facts from faeces: Prey remains in Wolf, Canis lupus, faeces revise occurrence records for mammals of British Columbia’s coastal archipelago. The Canadian Field Naturalist 119: 192-196.

Alexander, S. M., Waters, N. M. and Paquet, P. C. 2005. Traffic volume and highway permeability for a mammalian community in the Canadian Rocky Mountains. Canadian Geographer / Le Géographe Canadien 49: 321-331

Paquet, P.C., C.T. Darimont, F.M. Moola, and C. Genovali. 2005. Connectivity where the land meets the sea; preserving the last of the best. Wild Earth 14: 21-25

Paquet, P.C., C.T. Darimont, and C. Genovali. 2006. The Great Bear Rainforest; Home of island-hopping wolves. Howlings Magazine. Winter 2005/2006.

Paquet, P.C., and C.T. Darimont. 2006. Old Growth, New Concern. Monday Magazine. August 30, 2006.

Theberge, J. B., M. T. Theberge, J. A. Vucetich, and P. C. Paquet. 2006. Pitfalls of applying adaptive management to a wolf population in Algonquin Provincial Park, Ontario. Environmental Management 37: 451-460.

Alexander, S. M., T. B. Logan, P. C. Paquet. 2006. Spatio-temporal co-occurrence of cougar (Felis concolor), wolves (Canis lupus) and their prey during winter: A comparison of two analytical methods. Journal of Biogeography 33: 2001-2012.

Paquet, P. C. and E. G. Walker. 2006. Review of investigative findings relating to the death of Kenton Carnegie At Points North, Saskatchewan. Prepared for and submitted to Office of the Chief Coroner Saskatchewan Justice #920, 1801 Hamilton Street Regina, Saskatchewan S4P 4B4. 32 pp.

Bryan, H., C.T. Darimont, T.E. Reimchen, and P.C. Paquet. 2006. Early ontogenetic

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diet in wolves, Canis lupus, of coastal British Columbia. Canadian Field- Naturalist 120:61-66.

Darimont, C. T., P. C. Paquet, and T. E. Reimchen. 2007. Stable isotopic niche predicts fitness in a wolf-deer system. Biological Journal of the Linnaean Society 90: 125-137.

Stronen, A. V, Brooks, R. K., Paquet, P. C., and S. Mclachlan. 2007. Farmer attitudes toward wolves: Implications for the role of predators in managing disease. Biological Conservation 135: 1-10.

Stronen, A., P.C. Paquet, S. Herrero, S. Sharpe, N. Waters. 2007. Translocation and Recovery Efforts for the Telkwa Caribou, Rangifer tarandus caribou, Herd in westcentral British Columbia, 1997 – 2005. Canadian Field-Naturalist. 121:155- 163.

Nyhus, P. J., R. Lacy, F. R. Westley, P. Miller, H. Vredenburg, P. Paquet, and J. Pollak. 2007. Tackling biocomplexity with meta-models for species risk assessment. Ecology and Society 12(1): 31. [online] URL: http://www.ecologyandsociety.org/vol12/iss1/art31/

Musiani M., Leonard J. A., Cluff H. D., Gates C. C., Mariani S., Paquet P. C., Vilà C., Wayne R. K. 2007. Differentiation of tundra and boreal coniferous forest wolves: genetics, coat color and association with migratory caribou. Molecular Ecology 16: 4149-4170

Darimont, C.T., P.C. Paquet, and T.E. Reimchen. 2007. Stable isotopic niche predicts fitness of prey in a wolf-deer system. Biological Journal of the Linnean Society 90: 125-137

Apps, C. D., J. L. Weaver, P. C. Paquet, B. Bateman and B. N. McLellan. 2007. Carnivores in the southern Canadian Rockies: Core areas and connectivity across the Crowsnest Highway. Wildlife Conservation Society Canada Conservation Report No. 3. Toronto, Ontario, Canada.

Trapp, J.R., Beier, P., Mack, C., Parsons, D.R., and P.C. Paquet. 2008. Wolf, Canis lupus, den site selection in the Rocky Mountains. Canadian Field-Naturalist 122: 49-56.

Darimont, C. T., P. C. Paquet, and T. E. Reimchen. 2008. Spawning salmon disrupt tight trophic coupling between wolves and ungulate prey in coastal British Columbia. BMC Ecology 8: 14

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Darimont, C.T., T.E. Reimchen, H. Bryan, and P.C. Paquet. 2008. Faecal-centric approaches to wildlife ecology and conservation; methods, data and ethics. Wildlife Biology in Practice 4: 73-87

Darimont, C. T., Paquet, P. C. and T. E. Reimchen. 2009. Landscape heterogeneity and marine subsidy generate extensive intrapopulation niche diversity in a large terrestrial vertebrate. Journal of Animal Ecology 78: 126–133

Muñoz-Fuentes, V., C.T. Darimont, R.K. Wayne, P.C. Paquet, and J.A. Leonard. 2009. The genetic legacy of extirpation and re-colonization in Vancouver Island wolves. Conservation Genetics. Early edition. www.pnas.org_cgi_doi_10.1073_pnas.09007871106

Muñoz-Fuentes, V., C.T. Darimont, R.K. Wayne, P.C. Paquet, and J.A. Leonard. 2009. Ecological factors drive genetic differentiation in British Columbia gray wolves. Journal of Biogeography 36: 1516-1531.

Kinnison, M.T., E.P. Palkovacs, C.T. Darimont, S.M. Carlson, P.C. Paquet, and C.C. Wilmers. 2009. Some cautionary notes on fisheries evolutionary impact assessments. Proceedings of the National Academy of Sciences. Early edition, www.pnas.org_cgi_doi_10.1073_pnas.09007871106

Darimont, C.T., S.M. Carlson, M.T. Kinnison, P.C. Paquet, T.E. Reimchen, and C.C. Wilmers. 2009. Reply to Koons: Harvest-related trait changes in an increasingly variable world. Proceedings of the National Academy of Sciences 106: E33.

Darimont, C.T., S.M. Carlson, M.T. Kinnison, P.C. Paquet, T.E. Reimchen, and C.C. Wilmers. 2009. Human predators outpace other agents of trait change. Proceedings of the National Academy of Sciences 106: 952-954 (Research Highlight and News & Views in Nature, F1000 Biology Recommended Paper; global media coverage in popular press, e.g. New York Times)

Darimont, C.T., P.C. Paquet, and T.E. Reimchen. 2009. Landscape heterogeneity and marine subsidy generate extensive niche variation in a terrestrial carnivore. Journal of Animal Ecology 79: 126-133

McCrory, W.P. and P. Paquet. 2009. Proposed bear-viewing strategy for the K’ztim-a- deen (Khutzeymateen) Grizzly Bear Sanctuary & K’tzim-a-deen Inlet Conservancies, British Columbia. Report for the K’tzim-a-deen Management Committee & Planning Process, Prince Rupert, B.C.

Mathews, F., D.T. Blumstein, C. Darimont, K. Chan, L. Dill, D. Fraser, J. Kirkwood, K. Littin, D. Lusseau, and P. Paquet. 2010. Uniting wild animal welfare and

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conservation: guiding principles. Animal Conservation.

Bryan, H.M., Sim, K.A., Darimont, C.T., Paquet, P.C., Wagner, B.W., Muñoz-Fuentes, V., Smits, J.E., and Chilton, N.B. 2010. Identification of Parelaphostrongylus odocoilei (Nematoda: Protostrongylidae) first-stage larvae in the feces of gray wolves (Canis lupus) by molecular methods. Journal of Wildlife Diseases 46: 297-302.

Paquet, P.C., and C.T. Darimont. 2010. Wildlife conservation and animal welfare: two sides of the same coin? Journal of Animal Welfare 19: 177-190.

Stronen, A.V., Forbes, G.J., Sallows, T., Goulet, G., Musiani, M., and P. C. Paquet. 2010. Wolf body mass, skull morphology and mitochondrial DNA haplotypes in the Riding Mountain National Park region of Manitoba, Canada. Canadian Journal of Zoology 88: 496-507

Fox, C.H., Paquet, P.C., Reimchen, T.C. 2010. Active American black bear dens adjacent to a marine beach used for foraging. Ursus 21:195–197.

Darimont, C.T., Bryan, H.M., Carlson, S.M., Hocking, M.D., MacDuffee, M., Paquet, P.C., Price, M. H.H., Reimchen, T.E., Reynolds, J.D., and Wilmers, C. 2010. Salmon for terrestrial protected areas. Conservation Letters 3:379-389.

Stronen, A.V., Sallows, T. Forbes, G.J., and P.C. Paquet. 2011. Disease and parasite burden in an increasingly isolated wolf population. Journal of Wildlife Diseases 47:222–227

Bryan, H.M., Darimont, C.T., Paquet, P.C., Ellis, J.A., Goji, N., Gouix, M. and Smits, J.E. 2011. Exposure to infectious agents in dogs in remote coastal British Columbia: possible sentinels of diseases in wildlife and people. Canadian Journal of Veterinary Research 75:11-17.

Stronen, A.V., Forbes, G.J., Paquet, P. C., Goulet, G., Sallows, T., and M. Musiani. 2011. Dispersal in a plain landscape: short-distance genetic differentiation in southwestern Manitoba wolves, Canada. Conservation Genetics 13:359-371 DOI 10.1007/s10592-011-0290-1

Stronen AV, Schumaker NH, Forbes GJ, Paquet PC, Brook RK. 2011. Landscape resistance to dispersal: predicting long-term effects of human disturbance on a small and isolated wolf population in southwestern Manitoba, Canada. Environmental Monitoring and Assessment. Published Online DOI 10.1007/s10661-011-2469-9

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Nelson, M.P., Vecetich, J.A., Paquet, P.C. and J.K. Bump. 2011. An anadequate construct? North American Model: What’s flawed, what’s missing, what’s needed. The Wildlife Professional. Summer: 58-60

Daisuke Kawai Caroline H. Fox, Paul C. Paquet and Falk Huettmann. Accepted. Predictive performances of 3 models for seabird sistribution and abundance estimates; Random Forests, Boosted Regression Trees, and Generalized Additive Model. Ecological Applications

Darimont, C.T., Artelle, K., Bryan, H.M., Genovali, C., MacDuffee, M., and Paquet, P.C. Grizzlies, Salmon, and People: Traveling the Upstream Journey to a Sustainable Future. Prepared for a Bear Anthology edited by Lisa M. Kemmerer, in press August 2011.

Vander Wal, E.; Paquet, P.; and J. Andrés. 2012. Influence of landscape and social interactions on transmission of communicable disease in a social cervid. Molecular Biology 21: 271–1282 doi: 10.1111/j.1365-294X.2011.05431.x

Nelson, B., Hebblewhite, M., Ezenwa, V., Shury, T., Merrill, E.H., Paquet, P.C., Schmiegelow, F., Seip, D., Skinner, G., Webb, N. 2012. Seroprevalence of canine parvovirus and distemper in wolves in the Canadian Rocky Mountains. Journal of Wildlife Diseases 48:68-76.

Service, C., T. Nelson, W. McInnes, P.C. Paquet & C.T. Darimont. 2012. Evaluating external risks to protected areas; the proposed Enbridge Northern Gateway pipeline in British Columbia, Canada. Natural Areas Journal. 32: 500-509. DOI: http://dx.doi.org/10.3375/043.032.0404 URL: http://www.bioone.org/doi/full/10.3375/043.032.0404

Levi, T., C. T. Darimont, M.M. MacDuffee, M. Mangel, P.C. Paquet, & C.C. Wilmers. 2012. Using grizzly bears to assess economic and ecosystem tradeoffs in Pacific salmon management. PLoS Biology 10: e1001303.

Bryan, H.M. C.T. Darimont, J. Hill, P.C. Paquet, R.C.A. Thompson, B. Wagner, & J. E.G. Smits. 2012. Seasonal and biogeographic patterns of gastrointestinal parasites in large carnivores; wolves in a coastal archipelago. Parasitology 139: 781-790.

Stronen, A.V., Tessier, N., Jolicoeur, H., Paquet, P.C., He ́nault, M., Villemure, M., Patterson, B.R., Sallows, T., Goulet, G. and F.J. Lapointe. 2012. Canid hybridization: contemporary evolution in human-modified landscapes. Ecology and Evolution 1-13.

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Vander Wal, E., Paquet, P., and J. Andrés. 2012. Influence of landscape and social interactions on transmission of communicable disease in a social cervid. Molecular Biology 21: 271–1282 doi: 10.1111/j.1365-294X.2011.05431.x van Beest, F.M., Vander Wal, E., Stronen, A.V., Paquet, P.C. and R.K. Brook. 2013. Temporal variation in site fidelity: Scale-dependent effects of forage abundance and predation risk in a non-migratory large herbivore. Oecologia. 173:409–420 DOI 10.1007/s00442-013-2647-2

Vander Wal, E., Edye, I., Paquet, P.C., Coltman, D.W., Bayne, E., R.K. Brook, R.K., and J.A. Andres. 2013. Juxtaposition between an unstructured white-tailed deer population and a structured elk population: implications for disease transmission in a sympatric cervid community. Evolutionary Applications. 6:1001-1111.

Vander Wal, E., Paquet, P.C., Messier, F., and P.D. McLoughlin. 2013. Effects of phenology and sex on social proximity in a gregarious ungulate Canadian Journal of Zoology. 91:601–609 dx.doi.org/10.1139/cjz-2012-0237

Stronen, A.V. and Paquet, P.C. 2013. Perspectives on the conservation of wild hybrids. Biological Conservation. 167:390–395. http://dx.doi.org/10.1016/j.biocon.2013.09.004

Artelle, K.A., S.C. Anderson, A.B. Cooper, P.C. Paquet, J.D. Reynolds, C.T. Darimont. 2013. Confronting uncertainty in wildlife management: performance of grizzly bear management. PLoS ONE 8(11): e78041. doi:10.1371/journal.pone.0078041

MacDuffee, M., A.R. Rosenberger, R. Dixon, M.H.H. Price, and P.C. Paquet. 2013. Embroiled, Volume 1: Salmon, tankers and the Enbridge Northern Gateway Proposal. Raincoast Conservation Foundation. Sidney, British Columbia. Vers 01-13, pp. 107.

Bryan, H.M., C.T. Darimont, P.C. Paquet, K.E. Wynne-Edwards and J.E. Smits. 2014. Stress and reproductive hormones in grizzly bears reflect nutritional benefits and social consequences of a salmon foraging niche. PLOS ONE

Vander Wal, E., P.C. Paquet, F. Messier, and P.D. McLoughlin. 2013. Social proximity in an ungulate expressed as frequency and duration of interactions. Canadian Journal of Zoology.

Fox, C.H., R. El-Sabaawi, P.C. Paquet, and T.E. Reimchen. 2014. Pacific herring (Clupea pallasii) and wrack macrophytes subsidize semi-terrestrial detritivores. Marine Ecology Progress Series.

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Adams, M. S., J. Carpenter, J. A. Housty, D. Neasloss, P. C. Paquet, C. Service, J. Walkus, and C. T. Darimont. 2014. Toward increased engagement between academic and indigenous community partners in ecological research. Ecology and Society 19(3): 5. http://dx.doi.org/10.5751/ES-06569-190305

Artelle, K.A., Reynolds, J.D. Paquet, P.C. & Darimont, C.T. 2014. When science- based management isn’t. Science 343, 1311.

Darimont, C.T., C. Genovali, C. Service, and P.C. Paquet. In Press. Hunting for a trophy hunting ethic. In Animals and the Environment: Advocacy, Activism, and the Quest for Common Ground (L. Kemmerer, Ed). Routledge Press.

Bryan, H.M., Darimont, C.T., Paquet, P.C., Wynne-Edwards, K.E., and Smits, J.E.G. 2014. Stress and reproductive hormones reflect inter-specific social and nutritional conditions mediated by resource availability in a bear-salmon system. Conservation Physiology 2(1): cou010 93/conphys/cou010 doi: 10.10

Stronen, A.V., Navid, E.L., Quinn, M.S., Paquet, P.C., Bryan, H.M., and Darimont, C.T. 2014. Population genetic structure of gray wolves (Canis lupus) in a marine archipelago suggests island-mainland differentiation consistent with dietary niche. BMC Ecology 14:11. doi:10.1186/1472-6785-14-11

Christina N. Service, Megan S. Adams, Kyle A. Artelle, Laura V. Grant, Paul C. Paquet, and Chris T. Darimont. 2014. Indigenous knowledge and science unite to reveal spatial and temporal dimensions of distributional shift in wildlife of conservation concern. PLOS ON DOI: 10.1371/journal.pone.0101595

Bryan, Heather M., Judit E. G. Smits, Lee Koren, Paul C. Paquet, Katherine E. Wynne- Edwards and Marco Musiani. 2014. Heavily hunted wolves have higher stress and reproductive steroids than wolves with lower hunting pressure. Functional Ecology. DOI: 10.1111/1365-2435.12354

Brook, R.K., Cattet, M., Darimont, C.T., Paquet, P.C. and G. Proulx. 2015. Maintaining ethical standards during conservation crises. Canadian Wildlife Biology & Management 4: 72-79.

Fox, C.H., Paquet, P.C., and T.E. Reimchen. 2015. Novel species interactions: American black bears respond to Pacific herring spawn. BMC Ecology. DOI 10.1186/s12898-015-0045-9.

Best, B.D., Fox, C.H., Williams, R., Halprin, P.N., and P.C. Paquet. 2015. Updated

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marine mammal distribution and abundance estimates in British Columbia. Journal Of Cetacean Research and Management. 15: 9–26, 2015

Peer Reviewed Books and Chapters

Harrington, F.H. and P.C. Paquet (Eds.). 1982. Wolves of the world: Perspectives of behavior, ecology, and conservation. Noyes Publications, New Jersey

Paquet, P.C., S. Bragdon, and S. McCusker. 1982. Cooperative rearing of simultaneous litters in captive wolves. In F.H. Harrington and P.C. Paquet (Eds.) Wolves of the World, Noyes Publications, New Jersey, p. 223-237.

Harrington, F.H., P.C. Paquet, J. Ryon, and J.C. Fentress. 1982. Monogamy in wolves: A review of the evidence. In F.H Harrington and P.C Paquet (Eds.) Wolves of the World, Noyes Publications, New Jersey, p. 209-222.

Paquet, P.C. and W.A. Fuller. 1990. Scent marking and territoriality in wolves of Riding Mountain National Park. In Chemical signals in vertebrates 5. Eds. D.W. Macdonald, D. Muller-Schwarze, and S.E. Natynczuk. Oxford University Press, N.Y. pp 394-400.

Boyd, D.K., Paquet, P.C., Donelon S., Ream, R.R., Pletscher, D.H., and C.C. White. 1995. Transboundary movements of a recolonizing wolf population in the Rocky Mountains. Pages 135-140 In L.N. Carbyn, S.H. Fritts, and D.R. Seip, editors, Ecology and conservation of wolves in a changing world. Canadian Circumpolar Institute, Edmonton, Alberta.

Woods, J., Cornwell, L.L., Hurd, T., Kunelius, R., Paquet, P., and J. Wierzchowski 1996. Elk and Other Ungulates. Chapter 8 in: Green J., Pacas, C, Bayley, S., and Cornwell, L. (eds). A cumulative effects assessment and futures outlook for the Banff Bow Valley. Prepared for the Banff Bow Valley Study, Department of Canadian Heritage, Ottawa, On.

Paquet, P.C., Wierzchowski, J. and Callaghan, C. 1996. Summary report on the effects of human activity on gray wolves in the Bow River Valley, Banff National Park, Alberta. Chapter 7 in: Green J., Pacas, C, Bayley, S., and Cornwell, L. (eds). A cumulative effects assessment and futures outlook for the Banff Bow Valley. Prepared for the Banff Bow Valley Study, Department of Canadian Heritage, Ottawa, On.

Paquet, P.C. and Callaghan, C. 1996. Effects of linear developments on winter movements of gray wolves in the Bow River Valley of Banff National Park, Alberta. Pages 46-66 In G. Evink, D. Ziegler, P. Garret, and J. Barry, editors,

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Highways and movements of wildlife: improving habitat connections and wildlife passageways across highway corridors. Proceedings of the Florida Department of Transportation/Federal Highway Administration Transportation-Related Wildlife Mortality Seminar.

Paquet, P.C., and C. Callaghan. 1996. Effects of linear developments on winter movements of gray wolves in the Bow River Valley in Banff National Park, Alberta. Pages 46-66 In G.L. Evink, P. Garrett, D. Ziegler, and J. Berry, editors. Trends in Addressing Transportation Related Wildlife Mortality. Proceedings of the Transportation Related Wildlife Mortality Seminar. FL-ER-58-96. Florida Department of Transportation, Tallahassee, Florida, U.S.A.

Paquet, P.C. and Callaghan, C. 1997. GIS decision support models for highway planning. In G. Evink, D. Ziegler, P. Garret, and J. Barry, editors, Highways and movements of wildlife: improving habitat connections and wildlife passageways across highway corridors. Proceedings of the Florida Department of Transporation/Federal Highway Administration Transportation-Related Wildlife Mortality Seminar.

Terborgh, J., Estes, J.A., P.C. Paquet, K, Ralls, D. Boyd-Heger, B.J. Miller, and R.F. Noss. 1999. The role of top carnivores in regulating terrestrial ecosystems. Pages 39-64 in Continental Conservation, eds. M.E. Soule and J. Terborgh. Island Press, Washington, D.C. 227pp.

Carroll, C., Noss, R.F., Shoemaker, N, and P.C. Paquet. 2001. Is the restoration of wolf, wolverine, and grizzly bear to Oregon and California biologically feasible? In D. S. Maeher, R. F. Noss, and J. L. Larkin (eds.) Large Mammal Restoration, Island Press, Washington, D.C.

Duke, D.L., Hebblewhite, M., Paquet, P.C., Callaghan, C., and M. Percy. 2001. Restoration of a large carnivore corridor in Banff National Park, Alberta. Pages 261-276 in D. S. Maeher, R. F. Noss, and J. L. Larkin (eds.) Large Mammal Restoration, Island Press, Washington, D.C..

Paquet, P.C., J. R. Strittholt, N. L. Staus, P. J. Wilson, S. Grewal, and B. N. White. 2001. Feasibility of timber wolf reintroduction in Adirondack Park. Pages 47 - 64, in D. S. Maeher, R. F. Noss, and J. L. Larkin (eds.). Large Mammal Restoration, Island Press, Washington, D.C.

Paquet, P.C. 2001. Wolves. Pages 42-45, In D. Macdonald (ed) New Encyclopedia of Mammals, Oxford University Press, Oxford.

Paquet, P.C. 2001. An ongoing genetic mystery. Pages 46-47, in D. Macdonald (ed)

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New Encyclopedia of Mammals, Oxford University Press, Oxford.

Alexander, S., N. Waters, and P.C. Paquet. 2002. A Uncertainty-Based Decision Support Model for Restoring Wildlife Landscape Connectivity in Road Fragmented Environments. In G. Clarke and J. Stillwell, editors. Applied GIS and Spatial Modelling. University of Leeds, United Kingdom.

Paquet, P. C. and L. N. Carbyn. 2003. Wolf, Canis lupus and allies. Pages 482-510 in G. A. Feldhamer, B. C. Thompson, and J. A. Chapman, editors. Wild Mammals of North America: Biology, Management, and Conservation. Johns Hopkins University Press, Baltimore, MD

Alexander, S., N. Waters, and P.C. Paquet. 2004. A Probability-based GIS Model for Identifying Focal Species Linkage Zones across Highways in the Canadian Rocky Mountains. Pages 233-255 in G. Clarke and J. Stillwell, editors. Applied GIS and Spatial Modelling. University of Leeds, United Kingdom.

Paquet, P. C., S. M. Alexander, P. L. Swan, and C. T. Darimont. 2006. Pages 130-156 in Connectivity Conservation, eds K. R. Crooks and M. Sanjayan. Influence of natural landscape fragmentation and resource availability on distribution and connectivity of marine gray wolf (Canis lupus) populations on Central Coast, British Columbia, Canada. Cambridge University Press. N.Y. & England.

Paquet, P.C., S.M. Alexander, P.L. Swan, and C.T. Darimont. 2006. The influence of natural landscape fragmentation and resource availability on connectivity and distribution of marine gray wolf populations on the Central Coast, British Columbia, Canada. In Crooks, K. and M.A. Sanjayan (Eds.) Connectivity Conservation. Cambridge University Press. Cambridge, UK.

Paquet, P.C. Introduction. In McAllister, I., and C.T. Darimont. 2007. The Last Wild Wolves. Ghosts of the Great Bear Rainforest. University of California Press. Berkeley, CA.

Musiani, M., Boitani, L. and P. C. Paquet (eds.). 2009. A New Era for Wolves and People: Wolf Recovery, Human Attitudes, and Policy. University of Calgary Press, Calgary. AB Canada.

Bloch, G. , Dettling, P., Paquet, P. C., Gibeau, M. 2009. Auge in Auge mit dem Wolf. Kosmos, Koln, Germany.

Musiani, M, Boitani, L, and P. Paquet (eds). 2010. The world of wolves: new perspectives on ecology, behaviour and policy. University of Calgary Press, Calgary. AB, Canada.

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Musiani, M, Boitani, L, and P. Paquet (eds). 2010. A new era for wolves and people: wolf recovery, human attitudes and policy. University of Calgary Press, Calgary. AB, Canada.

H. Dean Cluff, H.D., Paquet, P.C., Walton, L.R., and M. Musiani. 2010. Wolf Ecology and Management in Northern Canada: Perspectives from a Snowmobile Wolf Hunt. In Musiani, M, Boitani, L, and P. Paquet (eds) The world of wolves: new perspectives on ecology, behaviour, and policy. University of Calgary Press, Calgary. AB, Canada.

Paquet, P.C., Alexander, S., Donelon, S., and C. Callaghan. 2010. Influence of anthropogenically modified snow conditions on movements and predatory behaviour of gray wolves. In Musiani, M, Boitani, L, and P. Paquet (eds) The world of wolves: new perspectives on ecology, behaviour, and policy. University of Calgary Press, Calgary. AB, Canada

Paquet, P.C., Miller, B., Kunkel, K., Reading, R.P., and M.K. Phillips. 2010. The importance of large carnivores. Pages 49-60 in Reading, R.P., B. J. Miller, A. Masching, R. Edward, and M. Phillips (eds.). Awakening Spirits: Wolves in the Southern Rockies. Fulcrum Publishing, Golden, CO.

Phillips, M.K. Miller, B., Kunkel, K., Paquet, P.C., Martin, W.W. and D.W. Smith. Potential for and implications of wolf restoration in the Southern Rockies. 2010. Pages 197-218 in Reading, R.P., B. J. Miller, A. Masching, R. Edward, and M. Phillips (eds.). Awakening Spirits: Wolves in the Southern Rockies. Fulcrum Publishing, Golden, CO.

Paquet, P.C. 2010. Foreword in Dettling, P. Will of the Land. Rocky Mountain Press. Canmore, AB.

DellaSala, D., Moola, F., Paquet, P., Noss, R. 2011. North America Coastal Rainforest: Alaska to the Redwoods. In DellaSala, D. (ed). Temperate and Lesser Known Rainforests of the World: Their Ecology and Conservation. Island Press

DellaSala, D., Paquet, P., McClellan, B., Alaback, P., Goward, T., Craighead, L. and T. Spribille. 2011. Inland Temperate Rainforests of the Interior Northwest. In DellaSala, D. (ed). Temperate and Lesser Known Rainforests of the World: Their Ecology and Conservation. Island Press

DellaSala, D.A., Alaback, P., Kirkpatrick, J., Paquet, P., von Wehrden, H., Techlin, D., Krestov, P., Noss, R., Hakon, H., Craighead, L., Nakamura, Y., Moola, F.,

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Nauman, R., and T. Spribille. 2011. Rainforests at the brink: A call for international action. In DellaSala, D. (ed). Temperate and Lesser Known Rainforests of the World: Their Ecology and Conservation. Island Press

Dominick A. DellaSala, Paul Alaback, Lance Craighead, Trevor Goward, Holien Håkon, James Kirkpatrick, Pavel V. Krestov, Faisal Moola, Yukito Nakamura, Richard S. Nauman, Reed F. Noss, Paul Paquet, Katrin Ronneberg, Toby Spribille, David Tecklin, Henrik von Wehrden. 2011. Crosscutting Issues and Conservation Strategies. In DellaSala, D. (ed). Temperate and Lesser Known Rainforests of the World: Their Ecology and Conservation. Island Press.

Paquet, P.C. in Alexander, S.M. (in press). Carnivore Management and the Role of Conservation GIS. Chapter 12 (Pages 293-317) in Resources and the Environmental Management in Practice, Ed., Dr. Bruce Mitchell. Oxford University Press. XXpp

Peer reviewing

I review ≥10 submissions per year, including those made to the following Journals: Biology Letters, Oikos, Oecologia, Animal Conservation, Aquatic Biology, Journal of Applied Ecology, Functional Ecology, Animal Behaviour, Marine Ecology Progress Series, Conservation Biology, Journal of Mammalogy, Behavioral Ecology, Biological Conservation, Rangifer, Northeastern Naturalist, Wildlife Biology.

Areas of Expertise

- Environmental Ethics - Conflict Resolution - Large Carnivores - Predator Prey Relationships - Population Ecology & Wildlife Disease - Population Dynamics of Large Mammals - Habitat Analysis - Computer Modelling - Telemetry - Remote Sensing Techniques - GIS applications - Ungulate Survey Techniques

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- Live Capture and Immobilization of Carnivores

Additional Interests and Skills

- Photography - Auto Mechanics - Mountaineering - Carpentry

Personal References

L. David Mech, Ph.D. North Central Forest Expt. Stn. 1992 Folwell Avenue St. Paul, MN, USA 55108

Rolf O. Peterson, Ph.D. Michigan Technological University Houghton, MI, USA 49931

Vince Crichton, Ph.D. Provincial Ungulate Biologist Wildlife Branch Department of Natural Resources Box 1400 1495 St. James Street Winnipeg, Manitoba R3H 0W9

Paquet, P.C. c.v. short form