Population Structure in Harbour Porpoises (Phocoena Phocoena) of British Columbia and Widespread Hybridization in Cetaceans

Population structure in harbour porpoises (Phocoena phocoena) of British Columbia and widespread hybridization in cetaceans

Carla Anne Crossman

BSc, Queen’s University, 2010

A THESIS SUBMITTED IN PARTIAL FUFILLMENT OF

THE REQUIREMENTS FOR THE DEGREE OF

MASTER OF SCIENCE

The Faculty of Graduate Studies

(Zoology)

THE UNIVERSITY OF BRITISH COLUMBIA

(Vancouver)

December 2012

Abstract

Harbour porpoises (Phocoena phocoena) are one of the most abundant small cetaceans in the world and, while they are extensively studied across most of their range, little is known about their biology in British Columbia, Canada. Recent management plans have identified a need to better understand the population structure of harbour porpoises in this region. I investigated the genetic population structure of harbour porpoises in British Columbia using mitochondrial DNA (mtDNA) and eight microsatellite loci. My findings are consistent with a single population of harbour porpoises inhabiting the coastline between Haida G’waii and the southern Juan de Fuca

Strait. I also confirmed that hybridization between harbour porpoises and Dall’s porpoises (Phocoenoides dalli) has occurred over a larger geographic region than previously known and I present evidence that the resultant hybrids are reproductively viable and have the potential to successfully backcross with both parental species.

Building on these findings, I examined patterns of hybridization across the order

Cetacea. I found that species pairs that share a greater number of ecological, morphological, and behavioural traits have a higher propensity to hybridize than species pairs that do not. This trend is largely driven by behavioural and morphological traits such as vocalization frequency and body size. My study aids in understanding harbour porpoise population structure in British Columbia, and highlights the occurrence of widespread cetacean hybridization.

Preface

This work did not require ethics review as the tissue samples were donated by the

Animal Heath Centre, Fisheries and Oceans Canada, Simon Fraser University,

Vancouver Aquarium Marine Science Centre and the Whale Museum. The tissue samples were not collected for my explicit use in research and were collected from animals that died naturally.

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Table of Contents

Abstract...... ii Preface ...... iii Table of Contents ...... iv List of Tables ...... vi List of Figures ...... vii List of Equations ...... viii Acknowledgements ...... ix 1 Introduction ...... 1 1.1 Conservation genetics ...... 1 1.2 Cetacean conservation ...... 3 1.3 The harbour porpoise ...... 4 1.4 Hybridization ...... 10 1.5 Opportunistic data ...... 10 1.6 Research objectives ...... 11 2 Population Structure and Intergeneric Hybridization ...... 13 2.1 Introduction ...... 13 2.1.1 The use of molecular markers for studies of population structure ...... 13 2.1.2 Harbour porpoise population structure ...... 14 2.1.3 Porpoise hybridization ...... 16 2.2 Methods ...... 17 2.2.1 Samples ...... 17 2.2.2 DNA extraction ...... 17 2.2.3 Mitochondrial DNA ...... 18 2.2.4 Microsatellites ...... 19 2.2.5 Identifying hybrids ...... 19 2.2.6 Population structure – mtDNA ...... 20 2.2.7 Population structure – microsatellites ...... 21 2.3 Results ...... 23 2.3.1 Identifying hybrids ...... 23 2.3.2 mtDNA ...... 27

2.3.3 Microsatellites ...... 30 2.4 Discussion ...... 34 2.4.1 Hybridization ...... 39 2.4.2 Contributions ...... 40 3 An Analysis of Cetacean Hybridization ...... 42 3.1 Introduction ...... 42 3.1.1 Research objectives ...... 46 3.2 Methods ...... 46 3.2.1 Data collection ...... 46 3.2.2 Similarity index ...... 46 3.2.3 Hybridization and similarity index ...... 49 3.3 Results ...... 51 3.3.1 Un-weighted analysis ...... 51 3.3.2 Weighted analysis ...... 56 3.4 Discussion ...... 60 3.4.1 Species barriers ...... 61 3.4.2 Potential benefits of interspecific mating ...... 62 3.4.3 Conclusions ...... 63 4 Conclusions ...... 64 4.1 Summary of findings ...... 64 4.2 Studies using samples of bycatch and strandings ...... 64 4.3 Future directions ...... 65 References ...... 68 Appendices ...... 85 A.1 Appendix S1 – Chapter 2 ...... 85 A.2 Appendix S2 – Chapter 3 ...... 98

List of Tables

Table 2.1 Prior probability and delta K from STRUCTURE runs on all samples………25 Table 2.2 Nucleotide and haplotypic diversity…………………………..………………29 Table 2.3 Variation in mtDNA among and within a priori populations……………..29 Table 2.4 Pairwise ФST between two a priori populations using mtDNA………………29 Table 2.5 Variation at eight microsatellite loci assayed in harbour porpoise………..30 Table 2.6 Variation in microsatellite loci among and within a priori populations…..33 Table 2.7 Pairwise FST between two a priori populations using microsatellite loci…….33 Table 3.1 Documented cases of cetacean hybridization in captivity…………….………..43 Table 3.2 Documented cases of cetacean hybridization in the wild………………………44 Table 3.3 Results of first four principal components for all species comparisons……….53 Table 3.4 Results of first four principal components for species with 44 chromosomes..55 Table 3.5 Results of the survey to calculate weighted traits………………………………57 Table 3.6 Results of weighted principle components for all species comparisons………59 Table 3.7 Results of weighted principle components for 44 chromosome species…….60 Table S1.1 Case numbers and additional organizations associated with each sample…85 Table S1.2 Nucleotide and haplotypic diversity using a 99% threshold………………..92 Table S1.3 Variation in mtDNA among/within populations using a 99% threshold……92 Table S1.4 Pairwise ФST between two a priori populations using a 99% threshold…….92 Table S1.5 Variation at eight microsatellite loci using a 99% threshold…………..………93 Table S1.6 Variation in microsatellite loci among/within population at 99% ……………93 Table S1.7 Pairwise FST between two a priori populations using a 99% threshold….…93 Table S1.8 Posterior probability of 10 runs of Geneland using a 99% threshold………...94 Table S1.10 Coordinate data used in Geneland …...………………………….………….…95 Table S1.11 mtDNA haplotypes by region…...……………………………………………...96 Table S2.1 Species trait values……………………………………….…………………....…..98 Table S2.4 Results of 10,000 subsampled PCAs for all species comparisons…………...144 Table S2.5 Results of 10,000 subsampled PCAs for species with 44 chromosomes...... 145 l

List of Figures

Figure 1.1 Pictures of a harbour porpoise and a Dall's porpoise…………………………...5 Figure 1.2 Distribution of harbour porpoise and Dall's porpoise…………………………..6 Figure 1.3 Sightings of porpoises in British Columbia………………………………………7 Figure 2.1 Graphical output of genetic assignment from NEWHYBRIDS……………….23 Figure 2.2 Summary output of a structure run from Dall's and harbour porpoises…….24 Figure 2.3 Sampling locations of harbour porpoises and hybrid porpoises……………..26 Figure 2.4 Maximum likelihood mtDNA phylogeny of harbour porpoises……………..28 Figure 2.5 Posterior probability of population membership from STRUCTURE……..31 Figure 2.6 Estimated population assignment from Geneland……………………………..32 Figure 2.7 Proportion of alleles in each frequency class from bottleneck……………34 Figure 3.1 Similarity index for hybridizing vs non-hybridizing pairs of species………52 Figure 3.2 Similarity index for cetacean species with 44 chromosomes………………….54 Figure 3.3 Weighted similarity index for all cetacean species……………………………58 Figure S1.9 Probability of population membership from STRUCTURE..……….……….94 l

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List of Equations

Equation 3.1 Similarity of continuous traits …………….………….…...…………….…….47 Equation 3.2 Similarity in continuous traits as categorical traits …………………………48 Equation 3.3 Compiling similarity index .…………………………….….……………….....49 Equation 3.4 Applying weighted average to similarity index …………...……………….49

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Acknowledgements

First and foremost I would like to thank my supervisors Lance Barrett-Lennard and Rick Taylor. Both of you were invaluable resources for not just academic guidance, but also personal growth. Through the experiences you have given me, I was intellectually and personally challenged and I will ever be grateful for your help in my becoming more independent throughout the course of my degree. You were both perfect compliments to one another and I can't imagine having a better pair of advisors to lean on for support.

I also wish to thank my committee members Dr. Andrew Trites and Dr. John Ford for your valuable contributions, suggestions and encouragement over the course of my degree.

I would like to thank the Wild Killer Whale Adoption Program at the Vancouver Aquarium for my funding. I thank the BC Cetacean Sightings Network for providing data for my research. These two organizations are nothing without the fantastic staff and volunteers who made working with them a wonderful experience. In particular I wish to extend my deep appreciation to Meghan McKillop, Caitlin Birdsall, Heather Lord, Leticiaa Legat and Joan Lopez for directly and indirectly making contributions to both my thesis and my experiences in Vancouver.

A huge thank-you to Stephen Raverty for the hands on expereinces you gave me with harbour porpoises, and for your continued interest and encouragment toward my project. Also thanks to Allyson Miscampbell who taught me everything I needed know about doing labwork. Both Allyson and Carol Ritland were fantastic resources for troubleshooting early on in my work. I also wish to thank Anna Hall - the porpoise guru of the west coast - for all of her help, guidance and advice.

A large number of people and organizations who helped contribute samples and sample information to my project deserve thanks: Animal Health Centre, BC Marine Mammal Response Network, Cascadia Research Collective (Robin Baird, John Calambokidis, Jessie Huggins), Central Puget Sound Marine Mammal Stranding Network, Cetus Research and Conservation Society, Department of Fisheries and Oceans Canada (John Ford, Lisa Spaven), NOAA (Kristin Wilkinson), San Juan County Marine Mammal Stranding Network, Whale Museum (Amy Traxler), Strawberry Isle

Research Society, Washington Department of Fish and Wildlife, Brad Hanson, and Pam Willis.

To my lab mates, I would like to extend huge thanks: Jon Mee, JS Moore, Matt Siegle, Stefan Dick, Monica Yau, Shannan May-McNally, Jen Ruskey and Amanda Moreira for all of your advice and support.

While the research brought me to Vancouver, my experiences here have been memorable and shaped by the amazing people I have met. For academic or emotional support, and/or for just making my time in Vancouver unforgettable, I owe a great deal of thanks to Stephanie Avery-Gomm, Alistair Blachford, Gwyllim Blackburn, Gina Conte, Georgina Cox, Alex Dalton, Anne Dalziel, Rich Fitzjohn, Carling Gerlinsky, Aleeza Gerstein, Taylor Gibbons, Andy LeBlanc, Julie Lee-Yaw, Alice Liou, Andrew MacDonald, Milica Mandic, Jasmine Ono, Erin Rechsteiner, Seth Rudman, Alana Schick, Michael Scott, Andrea Stephens, Dave Toews, Brianna Wright and so many other amazing members of the Department of Zoology.

Whether it was complaining that I was getting paid to go to school, sympathizing with my struggles or just being those people you could always turn to, I need to thank my friends from afar: Emilie deGuzman, Lindsay Kwan, Malia Murphy and Jessica Sealy.

I never liked to hear “you need to make sure you are working” and “everyone goes through this”; but hese kinds of continued support were always said with the best of intentions. I want to thank Leithen M’Gonigle for his unconditional support and for giving me that extra motivation to finish. Thanks to my parents for their editorial (and sometimes financial) support and for always trying their best to understand what I was doing. Thank you for encouraging both Rodney and I to follow our hearts into, not the most lucrative of careers, but in career directions where we will always be passionate about the work we do.

Last but certainly not least, my I want to thank my true sources of continued inspiration Daisy, Jack and Theodore. They always helped me put things in perspective and allowed me to see the importance of what I was doing.

1 Introduction

1.1 Conservation genetics

The targets of conservation actions can range from ecosystems to single species.

Protecting many species over their entire ranges would often require unrealistic

financial costs and logistical organization and, therefore, conservation managers often

seek to protect only particular populations based on how they rank in terms of various

prioritization exercises (e.g. Taylor et al. 2011). Thus, in any given case, it is critical for

us to first identify what constitutes a “population” and then to understand how the

persistence of a healthy meta-population (a group of smaller separated yet interacting

populations) might be affected by conservation of only some of the constituent

populations. Defining these “populations”, their boundaries and movements is a major

goal in conservation genetics.

The pattern of genetic diversity of a species is often a consequence of how

individuals arrange themselves spatially in the habitat that is available to them.

Populations that are separated by large distances are usually more genetically distinct

than neighbouring populations (Wright 1943, 1951). This likely reflects the higher

probability of encountering individuals from the same population and may also include

a specific preference for mates from their same population. The strength of the

restrictions to inter-population matings, the length of time they have been separated

and the mutation rate determine how much neutral genetic differentiation there is

between populations.

There is a continuum in the degree to which species exhibit population subdivision or structure (Waples and Gaggiotti 2006). The low-structure end of the spectrum occurs when the likelihood of mating is equal between all members of the species. The high structure end is represented by species comprising a set of stable, reproductively isolated populations. Such a system usually requires physical or ecological barriers to constrain mating between individuals from different populations.

Typically, in nature an intermediate condition occurs where there are several smaller populations, with some gene flow between them. The proportion of the mating among versus within groups in these so-called meta-populations is a principal determinant of population structure.

The level of genetic population structure can also depend on the biology of the organism being studied. High site fidelity limits gene flow, resulting in a much higher likelihood of mating with a neighbour than with an individual in a distant population causing relatedness to be directly linked to location (Pfenninger et al. 1996). The social dynamics of a population can also influence their structure. In some killer whale

(Orcinus orca) populations, individuals within matrilineal groups are closely related to one another and genetic diversity within matrilines is maintained because most mating occurs between members of different matrilines during periods of temporary association (Barrett-Lennard 2000). The level of genetic structure in these populations is, therefore, a direct reflection of selection to mate outside of their matriline. In other instances, the landscape may influence the genetic structure. For many species of freshwater fishes, waterfalls or tiny creeks may act as an impermeable barrier to upstream migration, and two independent populations may live and thrive on either side of this physical barrier (e.g., McGlashan and Hughes 2000). Some genetic exchange may take place if a fish is swept downstream, and in this case, the upstream population

could contribute diversity to the downstream, while crossing a waterfall in the

upstream direction could be much less common. In summary, profoundly ecological,

social, and geographic characteristics of the species and movement barriers in their

environments affects population structure. For this reason it is important to study the

population structure of an organism across its entire range as local-factors may affect

smaller populations in different ways.

Studies of population structure are becoming more common as the costs

associated with genetic analyses decrease. These types of studies are important to

conservation biologists, as they enable the tailoring of local conservation plans to the

needs of smaller populations and, therefore help protect the greater genetic diversity of

the species.

1.2 Cetacean conservation

Much remains unknown about many species occupying the world’s oceans. This

lack of knowledge is not so surprising when it comes to tiny microorganisms or vast

deep sea ecosystems both of which are difficult to study. It is, however, harder to justify

the limited understanding of some of the largest creatures on earth – cetaceans (whales,

porpoises, and dolphins). More than 50% of cetacean species are classified as Data

Deficient (i.e., there are insufficient data to assign a conservation status) by the

International Union for Conservation of Nature (IUCN) and 10% are listed as

Endangered or Critically Endangered (IUCN 2012). With such scarcity of knowledge,

conservation efforts are based on little information – making protecting cetacean

populations an enormous challenge.

Marine protected areas (MPA) are increasingly being proposed to conserve

marine species. In these areas, fishing is severely restricted and vessel traffic can be

limited. One of the major problems, however, is enforcement. While MPAs inshore

could be patrolled by local coastguards or conservation officers, offshore protection is

more difficult, as no single governing body has enforceable legislation over actions in

the open ocean (Van Dyke et al. 1993). Also, MPAs may help protect small regions of

habitat for some cetacean species; however, many species occupy much larger ranges –

often off the continental shelf. The inability to actively protect offshore habitat echoes

the difficulty of enforcing regulations at sea that could reduce potential threats such as

commercial whaling or harmful fishing practices.

Another challenge to conserving marine species involves the very nature of their

habitat. Pollution, for instance, is a major problem in many marine habitats (Shahidul

Islam and Tanaka 2004). While one country may have strict regulations in place to try to

prevent marine pollution, they cannot force neighbouring countries to follow the same

guidelines. Marine pollution at a global scale is, therefore, hard to prevent. Despite this,

efforts to reduce habitat degradation must continue as pollution poses a major threat to

many long-lived wide ranging cetaceans which accumulate toxins throughout their

lives (Rowe 2008). With so many species of cetaceans possibly at risk, it is imperative to

study the movements, needs and threats of these species in order to make changes at

both a local and global scale that can help conserve the diversity of the remaining

cetaceans.

1.3 The harbour porpoise

Harbour porpoises (Phocoena phocoena) are one of the smallest oceanic cetacean

species (Hoelzel 2002, Fig. 1.1a). At birth, calves range from 0.7m to 0.9m (Gaskin et al.

1974) and are less than 2m in length at physical maturity (Gaskin et al. 1974; Baird and

Guenther 1995). The average adult weighs 50-65kg (Gaskin et al. 1974; Read and Tolley

1997) and the species exhibits sexual dimorphism for body size, with females often larger than males (Read and Tolley 1997). Harbour porpoises are typically dark grey, with a white or light grey underbelly and a dark or brown line leading from the pectoral flipper to the eye (Scheffer and Slipp 1948; Koopman and Gaskin 1994). As with other porpoise species, their bodies are robust and they lack a pronounced rostrum

(Scheffer and Slipp 1948; Gaskin et al. 1974).

Figure 1.1 Pictures of two stranded animals used in the study for comparison between species: a) a stranded harbour porpoise (AHC 12-1023) and b) a stranded Dall’s porpoise (JX475443). 5

Harbour porpoises are restricted to temperate and sub-Arctic regions of the

Northern Hemisphere, inhabiting coastal waters of North America, Europe, the Black

Sea and northeastern Asia (Gaskin et al. 1974; Baird 2003, Fig. 1.2). They exhibit a strong preference for shallow waters, i.e., less than 125m in depth (Baird and Guenther 1994,

1995; Calambokidis et al. 1997). While they are still considered a single species, the

Society for Marine Mammalogy currently recognizes four subspecies: P. phocoena vomerina in the eastern Pacific Ocean, P. p. phocoena in the Atlantic Ocean, P. p. relicta in the Black Sea and an unnamed sub-species in the western Pacific Ocean (Committee on

Taxonomy 2009).

Figure 1.2 Distribution of harbour porpoise (Phocoena phocoena, hatched lines) and Dall's porpoise (Phocoenoides dalli, solid light grey shading). Range data available from IUCN 2012. 6

In the northeastern Pacific Ocean, harbour porpoises are found along the coast of

California north to southern Alaska. In British Columbia, harbour porpoises occur all along the coast, however higher densities have been reported in the Strait of Georgia and the Strait of Juan de Fuca (Hall 2004, Fig. 1.3).

Figure 1.3 Sightings data from the BC Cetacean Sightings Network. (2012, Vancouver Aquarium Marine Science Centre and Fisheries and Oceans Canada). Harbour porpoises are represented by yellow dots, Dall's porpoises by blue dots, hybrid porpoises by green dots (southwestern Vancouver Island). Data not corrected for effort. Used with permission.

Harbour porpoises are a secretive species that seldom approach vessels and rarely exhibit aerial behaviour (Hall 2004). They are usually sighted by their characteristic ‘porpoising’ motions where their backs and dorsal fins barely break the water’s surface (Hall 2004). They exhibit high levels of variation in their individual movement patterns, and, in the western Atlantic Ocean, they have been reported to travel an average distance of 20km per day (Read and Westgate 1997). Harbour porpoises in British Columbia are typically found in groups of one to four individuals

(Hall 2004), but groups of over 200 have been reported (Hall 2011). Their size dimorphism, and large testes (Gaskin et al. 1974) suggest a polygyandrous mating system (where both males and females breed with multiple partners; Dugatkin, 2009).

Females typically produce one calf annually and nurse their calves for 8-12 months

(Read and Hohn 1995; Oftedal 1997). Harbour porpoises have been reported to live 13 years in the Atlantic Ocean (Gaskin et al. 1974), while the oldest animal on record from

British Columbia lived to be 10 years old (Baird 2003).

The greatest identified threat to small cetaceans in British Columbia is accidental net entanglement in commercial fisheries (Stacey et al. 1997; Hall et al. 2002; Fisheries and Oceans Canada 2009). Gillnets, like those used in the British Columbia’s salmon fisheries, are affecting all species of porpoises worldwide (Jefferson and Curry 1994), and the salmon gillnet management areas designated by the Department of Fisheries and Oceans fall within high density harbour porpoise habitat (Williams et al. 2008;

Department of Fisheries and Oceans 2010). Annually, almost 100 harbour porpoises are estimated to be killed by salmon gill nets in British Columbia (Hall et al. 2002; Williams et al. 2008). The significance of this mortality for harbour porpoises is still unknown, as little is known about their population size(s), demography, and distribution.

The management plan for harbour porpoises in British Columbia highlights many other potential threats to this species including pollutant contamination, habitat degradation and vessel strikes (Fisheries and Oceans Canada 2009). Because harbour porpoises are concentrated near areas that are heavily populated by humans and that experience high vessel traffic flow, anthropogenic threats pose an even greater risk to the species.

As a small marine mammal, harbour porpoises face harassment and predation from many other cetacean species. In British Columbia, Pacific white sided dolphins

(Lagenorhynchus obliquidens) and resident killer whales have been observed harassing harbour porpoises to the point of mortality (Baird 1998; Ford et al. 1998). Harbour porpoises are also at risk from predation by killer whales. While resident killer whales persist on a piscivorous diet, transient killer whales feed on smaller marine mammals and harbour porpoises are a preferred food source (Jefferson et al. 1991; Ford et al.

1998).

Harbour porpoise diet varies both seasonally and across their range, reflecting prey availability and abundances (Rae 1973; Gannon et al. 1998; Santos et al. 2004).

Fishes, principally herring (Clupea spp.) and gadids (Family Gadidae), make up the largest portion of their diet, followed by other small fish species and occasionally squid

(Rae 1965, 1973; Recchia and Read 1989; Gannon et al. 1998; Hall 2004; Santos et al.

2004). Harbour porpoise diet in British Columbia may also include a high proportion of sand lance (Ammodytes hexapterus) near Vancouver Island (Hall 2004). In British

Columbia, harbour porpoises share considerable dietary overlap with Dall’s porpoises

(Phocoenoides dalli) and where the two species overlap in range, they compete for food resources (Walker et al. 1998).

1.4 Hybridization

There are two members of the family Phocoenidae – true porpoises – present in

Canada: the harbour porpoise and their sister species the Dall’s porpoise (Barnes 1985;

Baird and Guenther 1995; Hoelzel 2002). These two species diverged from each other

≈3.5 million years ago (McGowen et al. 2009). In British Columbia, hybrids have been

reported between Dall’s and harbour porpoise (Baird et al. 1998; Willis et al. 2004). A

hybrid was first reported as a fetus in a female Dall’s porpoise in 1994 (Baird et al.

1998). The fetus was intermediate in colouration patterns and in the number of

vertebrae, and its identity as a hybrid was confirmed using genetics (Baird et al. 1998).

Over the next ten years, more hybrids were identified based on their distinctive

colouration patterns.

Most of the hybrids have been seen associating with Dall’s porpoise (Willis et al.

2004). A genetic study targeting hybrids through biopsy sampling found that all of the

hybrids were the product of a male harbour porpoise and a female Dall’s porpoise

(Willis et al. 2004). Female hybrids have been found stranded while pregnant with full-

term neonatal calves, but secondary sexual characters are not as obvious and have not

been identified in male hybrids (Willis et al. 2004, A. Traxler: San Juan County Marine

Mammal Stranding Network, Washington pers. comm.). These observations raise

questions about the fertility of the hybrid porpoises, and the viability of their offspring,

but there has been little empirical or detailed study of the extent and distribution of

interspecific hybrids.

1.5 Opportunistic data

While much sighting data exist for harbour porpoises along the eastern Pacific

coast, it is difficult to make inferences about porpoise distributions with these data,

owing to the considerable spatial and temporal biases present in the sampling effort.

For instance, most sightings reported to the BC Cetacean Sightings Network come from

eco-tourism industry, government agencies, and researchers (C. Birdsall, BC Cetacean

Sightings Network pers. comm., 2012). These activities are often constrained to regions

of high vessel traffic and popular recreational areas and consequently, do not include

observations across much of the range of the harbour porpoise in British Columbia. The

biases in sighted record can be corrected using models of observer effort (e.g.

Rechsteiner 2012) and as a result, much can be inferred about harbour porpoise

distribution in BC.

1.6 Research objectives

Pacific harbour porpoises are listed as a species of Special Concern under the

Species-at-Risk Act in Canada (COSEWIC 2003). The current species management plan

noted gaps in knowledge about the biology of this species and cited a specific need to

better understand its population structure (Fisheries and Oceans Canada 2009). I

address this need, while also using genetic tools to better understand the ongoing

hybridization between harbour and Dall’s porpoise. I also look at hybridization across

the order Cetacea and try to uncover factors that may have an influence in

hybridization. Unlike many other taxonomic groups where the mechanisms underlying

hybridization have been the focus of many studies (i.e. fish: Jones et al. 2006; birds:

Randler 2006), hybridization in cetaceans on a broader scale has received little attention

beyond documenting hybrid occurrences (Sylvestre and Tasaka 1985).

In summary, the goals of my research are:

1) To quantify the degree of population structure exhibited by harbour porpoises in

British Columbia.

2) To assess the frequency and occurrence of hybridization between harbour

porpoises and Dall’s porpoises in British Columbia.

3) To test for associations between hybridization and species traits across the order

Cetacea.

For each question, I made the following predictions:

1) Harbour porpoises in British Columbia should exhibit high levels of population

structure that would be influenced by the geography of the region. Because

harbour porpoises exhibit high levels of population structure throughout many

other parts of their range and, because this structure is often associated with the

local geography (e.g. Walton 1997; Wang and Berggren 1997), populations

inhabiting the complex system of waterways along British Columbia’s coastline

should exhibit similarly complex patterns of population structure.

2) Hybrid porpoises in the wild should be more common and more widespread

than had been previously identified. Hybrid porpoises are extremely hard to

identify in the wild, and can often be confused with juvenile Dall’s porpoises (A.

Hall, UBC Zoology, pers. comm., 2012, Fig. 1.2). This, combined with the use of

more powerful genetic analysis and potential evidence that hybrids may be

fertile, suggests hybrid porpoises may often go unrecognized and unreported.

3) Pairs of cetacean species that hybridize will be more similar in their ecological,

morphological and behavioural traits than species pairs that do not. Similar

patterns have been suggested for other taxa (e.g. Randler 2006; Whitney et al.

2010). Increased similarity between hybridizing pairs of species should be

expected as species that are more closely related should have fewer genetic

incompatibilities and species with similar ecological traits, will occur more often

in sympatry providing more opportunities for hybrid pairs to form.

2 Population Structure and Intergeneric Hybridization

2.1 Introduction

2.1.1 The use of molecular markers for studies of population structure

Methods of studying population structure have improved greatly over the past

few decades thanks to the development and efficiency of molecular markers to

differentiate between populations (Sunnucks 2000). Non-coding regions of

mitochondrial (mtDNA) or nuclear DNA can be isolated and used to determine levels

of gene flow between populations (Bataillon et al. 1996; Petit et al. 1998). In this study, I

have chosen to use the D-Loop region of the mtDNA and microsatellites for the nuclear

markers to assess population structure in harbour porpoises from British Columbia.

The D-Loop is a variable non-coding region of mtDNA located adjacent to the

12S and cytochrome b regions (Southern et al. 1988) and is commonly used in studies of

population structure (e.g. Pourkazemi et al. 1999; Hirota et al. 2004; Boyko et al. 2009).

The region varies in length across taxa, and while providing considerable variation, is

also conserved across phylogenetic groups in flanking regions making broad scale

studies possible (i.e., humans and dolphins have 86% similarity in the D-Loop; Southern

et al. 1988).

Microsatellites are nuclear markers, meaning that unlike mtDNA which is solely

inherited maternally, they are inherited from both parents. Microsatellites are simple

sequence repeats, where repeats can vary in length typically less than five base pairs

(Bruford and Wayne 1993; Charlesworth et al. 1994). Allelic variation expressed as

different number of base pair repeats can help to distinguish between individuals or

populations (Charlesworth et al. 1994). They are very common nuclear markers used in studying many fields of genetics (Zane et al. 2002), and in particular in the study of population structure in cetaceans.

Microsatellite primers are designed for use on a given species, but they will often amplify in other related species. In cetaceans, the most common primers are used to detect dinucleotide repeats and were designed for the humpback (Megaptera novaeangeliae) and sperm whale (Physeter macrocephalus) (Valsecchi and Amos 1996).

Recently, Chen and Yang (2007) designed tetranucleotide primers for the finless porpoise (Neophocaena phocaenoides). Using tetranucleotide repeat primers over dinucleotides helps mitigate errors from polymerase slippage and errors during scoring and should therefore be favoured when sufficiently polymorphic (Taberlet et al. 1999;

Morin et al. 2001). The use of both mitochondrial and nuclear markers can help identify if there is a sexual bias to gene flow and can lend additional support to the identification of genetically distinct populations (Sunnucks 2000).

2.1.2 Harbour porpoise population structure

The population structure of harbour porpoises has been studied over most parts of their range. On a global scale, four regional groups are recognized: North Pacific,

Northwest Atlantic, Northeast Atlantic and the Black Sea, with no evidence of recent genetic exchange between groups (Rosel et al. 1995). Extensive study has been conducted on the Atlantic group of populations (e.g., Wang et al., 1996; Wang &

Berggren, 1997; Rosel et al., 1999; Wiemann et al., 2010) with the majority of studies focusing on European and Baltic population groups. By contrast, only two studies have directly looked at structure in the Pacific Ocean (Chivers et al. 2002; Taguchi et al. 2010).

Spatially structured populations have been found to characterize the region around the United Kingdom and the Baltic Sea, with varying levels of subdivision between these populations (Walton 1997; Wang and Berggren 1997; Wiemann et al.

2010; De Luna et al. 2012). Further, Wang et al. (1996) provided evidence that a structured population (with four subunits) also characterizes the eastern coast of North

America. In the northeastern Pacific Ocean, harbour porpoise contaminant load varied along the coast, which provides some initial evidence of limits to the spatial distribution and movement within this population (Calambokidis and Barlow 1991). Inferences from preliminary mtDNA studies suggested that four sub-populations occupy this region, roughly corresponding to the shorelines along California, Washington, British

Columbia and Alaska; however, many haplotypes were shared across regions, suggesting high levels of gene flow. Small sample sizes from certain regions, however, may compromise the robustness of these findings (Rosel et al. 1995). More recent evidence supports the assertion that there are four sub-populations between California and Washington, and one sub-population in British Columbia (Chivers et al. 2002).

Furthermore, the subpopulation in British Columbia appeared to be even further divided, with significant differentiation between inland waters (i.e., Strait of Georgia) and outer waters (i.e., Vancouver Island) (Chivers et al. 2002). While interesting patterns have begun to emerge in British Columbia, this areahas not been the focal point of any of these studies and has not been studied with adequate sample sizes, and thus further research was needed before informed conclusions about the nature of these populations could be made.

The United States government has accepted findings from these studies and currently recognizes five Pacific populations and manages them independently

(Carretta et al. 2005). The Canadian government currently recognizes a single

population in British Columbia although the Species at Risk Act (SARA) Pacific harbour porpoise management plan (Fisheries and Oceans Canada 2009) highlights the need for a better understanding of the population structure of the species. Here, I conduct a thorough assessment of the structure of this population. This work will help address the conservation needs of this species in Canada.

2.1.3 Porpoise hybridization

Dall’s porpoises and harbour porpoises completely overlap in their range in

British Columbia (Gaskin et al. 1974; Jefferson 1988, Fig. 1.2). Dall’s porpoises are primarily black, with a white patch on their flanks that wraps around ventrally

(Jefferson 1988). They are larger and more robust than harbour porpoises and frequently approach boats and engage in aerial displays (Willis et al. 2004). The two porpoise species are members of two different sub-families; Phocoeninae and

Phocoenoidinae. While the divergence time between these two sub-families is still not known, evidence suggests they diverged over 3 million years ago (McGowen et al. 2009;

Slater et al. 2010).

There have been documented cases of hybridization between harbour porpoises and Dall’s porpoises in British Columbia (Baird et al. 1998; Willis, et al. 2004) and previous research has suggested these to be potentially reproductively viable hybrids

(Baird et al. 1998). Consequently, in addition to population structure, I investigated the presence and distribution of hybrid porpoises in my study area.

Hybrids between harbour porpoise and Dall’s porpoises most closely resemble the former species, yet they have been reported to behave much more like Dall’s porpoises (Willis et al. 2004). This makes field recognition of hybrids problematic and therefore, it is important to confirm, using more powerful genetic methods (Anderson

and Thompson 2002), whether samples collected from other so-called “pure” harbour porpoises also contain evidence of mixed ancestry. Failing to do so could lead to spurious conclusions about the population structure and demographic trends in this species. In assessing the presence and relative abundance of classes of hybrids, my data will also be able to assess whether hybrids are viable in nature.

2.2 Methods

2.2.1 Samples

All of the samples used in this study were collected along the coasts of British

Columbia and Washington with most coming from stranded carcasses of both harbour and Dall’s porpoises and donated by a number of organizations (See Appendix S1.1 for a list of sample case numbers and organizations which donated samples). Of the 247 samples, 189 were identified as harbour porpoises, 44 as Dall’s porpoises, 10 as hybrids and 4 as unidentified porpoises. A small portion of the samples (< 8%) were obtained via biopsy darting from a previous study (Willis et al. 2004). Samples were collected between and May 1992 and May 2012. When available, skin/blubber was the preferred tissue (≈ 75% of samples); however muscle and organ tissues were also used. Samples were stored at -20°C or -80°C in 20% DMSO or in 95% ethanol (EtOH). Samples are currently stored at the Vancouver Aquarium’s Cetacean Research Lab.

2.2.2 DNA extraction

I extracted DNA using standard phenol-chloroform methods (Sambrook et al.

1989). Between 25 and 50 mg of tissue were finely diced and added to a solution of 10%

SDS, proteinase K buffer (0.1M Tris, 50mM EDTA, and 100mM NaCl), and proteinase K enzyme (20ng/mL) for digestion. Solutions were placed in a rotator oven on slow speed

and incubated at 57°C overnight. Phenol:chloroform isoamyl alcohol was added, mixed for 3 minutes and centrifuged for 1 min. Supernatant was removed and the previous step was repeated twice: once with phenol:chloroform isoamyl alcohol and once with chloroform:isoamyl alcohol. 5M NaCl and 95% EtOH were added to the solution and chilled at -20°C to allow DNA to precipitate. Solutions were spun at -5°C for < 15min to allow DNA precipitate to form a pellet. EtOH was removed and pellet was washed with

70% EtOH. The DNA pellet was spun dry in a heated speed vac. DNA was resuspended in 1X TE buffer at room temperature overnight. DNA concentrations were quantified using a NanoDrop 2000 spectrophotometer (NanoDrop Technologies, Wilmington, DE) and diluted to 50ng/μl.

2.2.3 Mitochondrial DNA

A 545 bp region of the mitochondrial D-Loop region was amplified by polymerize chain reaction (PCR) using the primers RHD5MF (5’-

TACCCCGGTCTTGTAAACC-3’) and RHDint (5’-CCTGAAGTAAGAACCAGATG-3’)

(based on: Barrett-Lennard 2000; Rosel, Dizon, & Heyning 1994). The reagents for the

PCR reaction for each sample included 1.00μl of DNA (50ng/μl), 11.86μl of distilled autoclaved water, 3.00μl dNTP (2.5mM), 3.00μl of each primer (10μM), 0.50μl MgCl2

(50mM), 2.50μl 10x Paq buffer, 0.14μl Paq (5U). The PCR was conducted under the following conditions: 94°C for 3 min, 63°C for 1 min, 72°C for 3 min, 30 cycles of: 94°C for 50 sec, 61°C for 1 min, 72°C for 3 min; 72°C for 25 min. Products were cleaned and purified using a Wizard Prep Kit (Promega, Madison, WI). Purified DNA samples were sent to the Nucleic Acid Protein Service Unit (NAPS Unit, University of British

Columbia, Vancouver, BC) to be sequenced. Seven percent of the samples were also sequenced in the reverse direction to ensure proper sequencing. Sequences were

entered in GenBank under accession numbers JX475289 – JX475450, and JX477596 –

JX477621.

2.2.4 Microsatellites

Nine tetranucleotide primers (Np403, NP404, Np407, Np409, Np417, Np426,

Np427, Np428, Np430) were designed for finless porpoise in a previous study and were shown to amplify on other porpoise species (Chen and Yang 2008). Samples were prepared for PCR using QIAGEN multiplex kits (QIAGEN, Toronto, ON). The multiplex kits allowed multiple primers to be amplified and analyzed in a single reaction. Each reaction contained 1.5μl DNA (diluted to 20ng/μl), 10μl QIAGEN Master

Mix, 0.4μl of each primer – both forward and reverse (0.8μl was used for Np417 and

Np427), and distilled autoclaved water to top up solution to 20.0μl. The PCR was performed under the following conditions: 95°C for 15 min, 25 cycles of: 94°C for 30 sec, 57.3°C for 90 sec, 72°C for 60 sec; 60°C for 30 min. Distilled autoclaved water (200μl) was added to each reaction. Amplified PCR products were sized by comparison to a 400 bp size standard on a CEQ 8000 (Beckman-Coulter, Mississauga, ON). The primers were grouped into two batches (Np404, Np407, Np426, Np427 and Np403, Np409,

Np417, Np428, Np430) and each sample underwent an independent multiplexed PCR for each group of primers. One primer (Np403) was excluded from the analyses as it did not amplify under the multiplex conditions.

2.2.5 Identifying hybrids

With ongoing hybridization between Dall’s porpoises and harbour porpoises, analyses of population structure for a single species require certainty that an individual is of pure origin (e.g., is not the product of or a recent descendant of a hybridization

event). I identified hybrids from the output of NEWHYBRIDS v.1.1 beta (Anderson and

Thompson 2002) and STRUCTURE v.2.3.4 (Pritchard et al. 2000).

I used the genetics software NEWHYBRIDS v.1.1 beta (Anderson and Thompson

2002) to assign a probability that each individual belonged to one of the following classes: a pure harbour porpoise (HP x HP), a pure Dall’s porpoise (DP x DP), an F1 hybrid (HP x DP), an F2 hybrid (F1 x F1), a backcross with a harbour porpoise (F1 x

HP), or a backcross with a Dall’s porpoise (F1 x DP). From NEWHYBRIDS, those individuals that did not have a greater than 0.95 probability of being either a pure harbour porpoise or a pure Dall’s porpoise were classified as having sufficient mixed ancestry to be considered potential hybrids.

Using the program STRUCTURE v.2.3.4 (Pritchard et al. 2000), I conducted a test of population assignment on all samples of both species with 20 independent runs with the following parameters: 100,000 burn-in replicates, 500,000 MCMC replicates and assuming an admixture model with correlated allele frequencies. I tested for the number of putative populations (K) from 1 to 10; as expected the results suggested K = 2

(harbour porpoises and Dall’s porpoises). I considered those individuals that had less than a 95% membership to a single population as having sufficient mixed ancestry to be potential hybrids. I also conducted all of the population structure analyses with a 99% threshold value for pure species (n = 183) and the results followed the same patterns and trends as the 95% threshold value (n = 194) (Appendix S1.2-S1.9).

2.2.6 Population structure – mtDNA

I conducted my analysis of population structure using only those individuals that had a ≥95% probability of being a ‘pure’ harbour porpoise in NEWHYBRIDS or

STRUCTURE. I performed the analyses of mtDNA sequences in MEGA 5 (Kumar et al.

2008; Tamura et al. 2011) and aligned the sequences with ClustalW. The best model for determining phylogenetic relatedness was estimated using the Akaike Information

Criterion corrected for sample size (AICc) and a maximum likelihood tree was built using the best model and bootstrapped with 500 replicates.

To identify whether sampling location could be driving population structure, I analyzed sequences in Arlequin v.3.5 (Excoffier et al. 2005), using two sampling groups defined a priori: inside waters (Juan de Fuca Strait, Strait of Georgia, and Puget Sound), outside/northern waters (west of Vancouver Island, Johnstone Strait and north of

Vancouver Island). There were insufficient sample sizes to allow testing for three populations, and therefore based on sample size and similarity as open water areas, the northern and outside populations were combined. I chose these groups a priori because the coastline of British Columbia provided natural geographic constrictions that could prevent or discourage movement between these areas. An analysis of molecular variance (AMOVA) was used to determine whether population differentiation was greater between or within groups.

2.2.7 Population structure – microsatellites

I checked for the presence of null alleles and deviations from Hardy-Weinberg and linkage equilibrium in Microchecker v.2.2.3 (Van Oosterhout et al. 2004) and

GENEPOP v.4.1.3 (Raymond and Rousset 1995) respectively. Using the microsatellite data and STRUCTURE, I estimated the number of putative populations (K) within the sampling region using no prior information. The STRUCTURE analysis used correlated allele frequencies with a burn-in period of 100,000 replications and 500,000 MCMC replications. In 20 independent runs, K was estimated from 1 to 10.

In order to confirm the results of the STRUCTURE using sampling location as priors, a similar analysis was conducted in R v.2.12.2 (R Project for Statistical

Computing) using the package Geneland v.4.0.0 (Guillot et al. 2005). Coordinates for sampling locations were estimated using the most accurate stranding location information available (Appendix S1.10). Some samples were provided with exact stranding coordinates, while others were referenced to a nearby community or landmark. I allowed uncertainty in the coordinates based on the longest known daily range movement (similar to McAuliffe Dore, Turner, & Lorenz 2009) as reported via satellite telemetry in Read & Westgate (1997). I used 100,000 iterations, correlated allele frequencies and 10 independent runs and estimating a K from 1 to 10.

To determine if a step-wise or infinite allele model would be more appropriate, I used SPAGeDi v.1.3 (Hardy and Vekemans 2002). FST and RST were highly correlated

(R=0.981) and allele size did not have a significant contribution to population differentiation. Therefore an infinite allele model (and FST) was used throughout the analyses.

I then tested for the potential presence of a bottleneck effect using 10,000 replicates in Bottleneck v.1.2.02 (Piry et al. 1990) to ensure that there have been no large decreases in population size that could affect the genetic diversity of the current population of harbour porpoises. Finally, I assessed population structure based on the locations defined a priori using an AMOVA in ARLEQUIN and using allele frequency contingency tests for population differentiation in GENEPOP.

2.3 Results

2.3.1 Identifying hybrids

Out of 258 individuals, NEWHYBRIDS identified 205 near pure harbour porpoises, 36 near pure Dall’s porpoises,

and 17 individuals of mixed ancestry. Of these putative hybrids, NEWHYBRIDS identified many of these individuals as

F1 hybrids, whereas others were more likely F2 hybrids, or backcrosses between F1 hybrids and Dall’s porpoises (Fig 2.1).

Individual number

Figure 2.1 Graphical output of genetic assignment of individuals from NEWHYBRIDS for 263 harbour porpoises. Each individual is represented by a vertical bar, with colours depicting probability (P) of ancestry: pure harbour porpoise is blue, pure Dall’s porpoise in green, F1 hybrids in red, F1xDall’s in dark orange, F1xharbour in deep yellow, and F1xF1 in lemon yellow.

There were two putative populations, as expected when all samples (n = 258) were combined (STRUCTURE, K = 2, Dall’s and harbour porpoises) (Fig 2.2, Table 2.1).

Every individual was assessed a probability of being a Dall’s porpoise and of being a harbour porpoise. STRUCTURE identified 196 near pure harbour porpoises (> 95% harbour porpoise genome), 44 near pure Dall’s porpoises and 18 individuals with potential mixed ancestry. In addition to the hybrid parentage results I found using

NEWHYBRIDS, I was able to identify putative hybrids in STRUCTURE with a small proportion of Dall’s porpoise DNA relative to harbour porpoise DNA suggesting backcrosses may also be occurring between F1 hybrids and harbour porpoises. These hybrid porpoises are found across a wide range of coastline (Fig 2.3). Many were not identifiable morphologically as many genetically-identified hybrids were mistaken as a parental species by experienced observers and a veterinary pathologist.

Individual Figure 2.2 Summary output of 20 independent runs in STRUCTURE from Dall's and harbour porpoises (K=2, N = 258). Each individual is represented by a thin vertical line the height of which shows its admixture coefficient (Q, summing to 1.0). Individuals with pure harbour porpoise ancestry are indicated by vertical green bars and those with pure Dall's porpoise ancestry by vertical red bars. Individuals with mixed ancestry are indicated by bars with both red and green, the proportional height of each coloured section representing the proportional contribution of Dall’s (red) or harbour (green) porpoise genome.

Table 2.1 Results of 20 independent runs testing the number of putative populations (K) from STRUCTURE using both posterior probability (Mean LnP(K)) and the Evanno method (Delta K) of all samples – harbour, Dall’s and hybrid porpoises (n = 258). K Posterior Probability Delta K (Mean LnP(K)) 1 -5482.49 2 -4604.19 284.32 3 -4558.51 3.22 4 -4533.20 2.62 5 -4578.69 1.48 6 -4539.70 0.34 7 -4524.55 0.98 8 -4572.09 0.62 9 -4563.74 0.21 10 -4575.98

Figure 2.3 Sampling locations of harbour porpoises (dark circles) and hybrid porpoises (white crossed circles) along the coast of British Columbia and Washington. Hybrid locations are based on identifying hybrids throughout the analyses. A priori sampling areas (outside, inside and northern waters) are labelled and boundaries indicated by dashed lines.

2.3.2 mtDNA

I examined a 545 bp region of the mitochondrial D-Loop in 95% pure harbour porpoises (n = 147). There were 134 unique haplotypes with 168 variable sites. Few haplotypes were shared between or within the defined sampling populations

(Appendix S1.11).

The best model under the AICc to build the maximum likelihood tree was a general time-reversible model including a gamma distribution and invariant sites –

GTR+G+I with both a finless porpoise (GenBank Accession Number HQ108437.1) and a

Dall’s porpoise (GenBank JX475429, this study) as an outgroup (Fig 2.4). The tree shows little to no resolution of major groupings and results in a star phylogeny with little evidence of population structure.

Because there were very few shared haplotypes, haplotype diversity was close to

1 in each defined subpopulation. Nucleotide diversity, however, was much lower and ranged from 0.023-0.034 (Table 2.2).

The results of the AMOVA (Outside/Northern Waters: n=19, Inside Waters: n=109) suggested all of the variation was found within these geographic groupings and little to none of the variation was from between populations (Table 2.3). The pairwise

ФST value between populations was < 0.001 and non-significant (Table 2.4).

JX475329 JX475405 JX475389 JX475289 JX475386 Finless Porpoise JX475322

Dall's Porpoise JX475344 JX475304 JX475310 JX475345 JX475328 JX475328 JX475364

JX475317 JX475346 JX475415

JX475358 JX475382

JX475342 JX475293 JX475318

JX475375 JX475298

JX475360 JX475316

JX475337 JX475366 JX475291

JX475332 JX475313

JX475371 JX475393

JX475356 JX475392

JX475290 JX475367 JX475394

JX475300 JX475331

JX475300 JX475321

JX475363 JX475325 JX475300

JX475335 JX475336

JX475347 JX475326

JX475308 JX475296

JX475383 JX475323 JX475351

JX475401 JX475323

JX475348 JX475374

JX475306 JX475385 JX475348

JX475401 JX475324

JX475320 JX475414

JX475410 JX475305 JX475349 JX475299

JX475357

JX475411 JX475292

JX475416 JX475417 JX475403 JX475400

JX475400

JX475397 JX475398

JX475391 JX475354

JX475412 JX475399 JX475409 JX475301 JX475307

JX475390 JX475402 JX475312

JX475315

JX475387 JX475334 JX475314 JX475294

JX475352

JX475413 JX475295

JX475311 JX475370

JX475339 JX475406 JX475309 JX475338

JX475368

JX475303 JX475379

JX475404 JX475294 JX475408 JX475328

JX475333

JX475395 JX475353 JX475303 JX475328

JX475294

JX475396 JX475340 JX475372 JX475369

JX475377

JX475365 JX475323

JX475327 JX475302

JX475418 JX475361

JX475359 JX475343

JX475341 JX475378

JX475419 JX475362

JX475373 JX475381

JX475376 JX475294 JX475294 JX475380

Figure 2.4 A bootstrap consensus maximum likelihood mtDNA phylogeny of harbour porpoises in the northeastern Pacific Ocean based on clustering of pairwise sequence divergence estimates derived following the general time- reversible model of DNA substitution of the d-loop region (545 base pairs). No groups were supported at bootstraps values of at least 75%. Haplotypes are defined by location in Appendix S1.11. The finless porpoise (Neophocoena phoco enoides) and Dall’s porpoise were used as outgroups.

Table 2.2 Nucleotide and haplotypic diversity of 545 base pairs of mtDNA D-loop sequences from 138 harbour porpoises, 121 of known location. Nucleotide Haplotypic Sample Diversity Diversity Size (n) Three Sampling Locations Northern 0.0233±0.0149 1.0000±0.1265 5 Waters Outer Waters 0.0294±0.0157 1.0000±0.0270 14 Inner Waters 0.0269±0.0134 0.9966±0.0020 109 Two Sampling Locations Outside/North 0.0275±0.0144 1.0000±0.0171 19 Inside Waters 0.0269±0.0134 0.9966±0.0020 109 All Samples Single 0.0343±0.0169 0.9979±0.0014 147 Population

Table 2.3 Results from analysis of molecular variance apportioning variation in mtDNA d-loop sequences among and within a priori populations of harbour porpoise.

% variation among % variation within ФST P populations populations Two Sampling -0.70 100.70 -0.00705 0.68±0.016 Groups

Table 2.4 Pairwise ФST (lower diagonal) between two a priori populations based on variation in mtDNA d-loop sequences among and within a priori populations of harbour porpoise. The probabilities that the reported ФST values are significantly different from 0 are indicated in the upper diagonal. Inside Outside/North Inside - 0.66 ± 0.01 Outside/North -0.0071 -

2.3.3 Microsatellites

I analysed the microsatellite data for samples in which more than half (5+) loci amplified (n = 194). Across all samples of harbour porpoise there was general conformance to Hardy-Weinberg equilibrium (at all except one locus). There was only one locus that showed evidence of null alleles (Np426). At this locus, there appeared to be Dall’s porpoise specific alleles (which differed in length from the harbour porpoise alleles by a single base pair) present in the near pure harbour porpoises, suggesting either a possible mutation or a former hybridization event. Microsatellite variation was different for each locus (Table 2.5), and the number of alleles at each locus ranged from

1 to 12. Observed heterozygosity (Ho) values ranged from 0.36-1.0. The one monomorphic locus (Np407) was helpful in discriminating between harbour and Dall’s porpoises, as it was fixed for a single allele in harbour porpoise and other alleles were present in most of the Dall’s and hybrid individuals. Seven of twenty-eight pairwise tests between loci showed signs of linkage disequilibrium; however, only one comparison remained significant after Bonferonni correction (Np404/Np426).

Table 2.5 Variation at eight microsatellite loci assayed in harbour porpoise. Loci Number of Expected Observed Range in Alleles Heterozygosity Heterozygosity Allele Size (He) (Ho) (bp) Np404 6 0.6105 0.5926 134-150 Np407 1 0.0000 0.0000 186 Np409 3 0.4895 0.4844 221-229 Np417 12 0.7762 0.8000 128-176 Np426 6 0.4018 0.3579 103-116 Np427 7 0.6656 0.6667 178-194 Np428 8 0.7476 0.7713 110-134 Np430 3 0.0941 0.0855 144-168

The analysis in STRUCTURE suggested a single population of harbour porpoises as the highest log-likelihood was consistently associated with K = 1 (Figure 2.5).

Geneland also suggested a single putative population (n = 167, K = 1) (Figure 2.6) and there was no support for population differentiation from the analysis in GENEPOP (χ2 =

15.122, df = 14, P = 0.37). Consistent with the mtDNA results, almost all of the variation was found within rather than among any regional groupings when examined with

AMOVA (Table 2.6 and Table 2.7).

-2800

-3000

-3200 Mean LnP(K) Mean

-3400

-3600

2 4 6 8 10

Figure 2.5 Posterior probability of population membership from STRUCTURE for 1 to 10 putative populations (K) of harbour porpoise sampled within southern British Columbia. Each value is the mean of 20 STRUCTURE simulations of vartiation across eight microsatellite DNA loci (error bars are standard deviations).

Figure 2.6 Estimated population assignment from Geneland. Different colours are used to represent distinct populations and the presence of a single colour (yellow) suggests a single population based on variation in allele frequencies of harbour porpoise assayed at eight microsatellite DNA loci across various using the sampling locations (black dots). An approximate outline map of the coastline of British Columbia is overlaid for reference. Because the program is not given species range boundaries, it is unable to assess that harbour porpoises would not inhabit the land and therefore assigns land area to a population as well.

Table 2.6 Results from analysis of molecular variance apportioning variation in allele frequencies among and within a priori populations (Inside and Outside/Northern Waters) of harbour porpoise assayed at eight microsatellite DNA loci.

% variation % variation FST P among within geographic geographic groups groups Two Sampling 0.25 99.75 0.0025 0.25 ± 0.015 Groups

Table 2.7 Pairwise FST (lower diagonal) between two a priori (Inside, Outside/North) geographic groupings based on variation in allele frequencies among and within a priori populations of harbour porpoise assayed at eight microsatellite DNA loci. The probabilities of Type I error rates associated with the observed FST values are indicated in the upper diagonal. Inside (N= 134) Outside/North (N = 25)

Inside - 0.33 ± 0.02 Outside/North 0.0010 -

There was not consistent evidence of a population bottleneck using heterozygote excess under the infinite alleles model (I.A.M.) and the step wise mutation model

(S.M.M.) (I.A.M.: P = 0.33, 0.055; S.M.M.: P = 0.021, 0.99; sign test and 1-tailed Wilcoxon test respectively). The graphical output of the mode-shift fit a qualitative L-shaped distribution, providing additional evidence against a previous population bottleneck

(Figure 2.7).

0.7

0.6

0.5

0.4

0.3

Proportion of alleles of Proportion

0.2

0.1 0.0

0.0 0.2 0.4 0.6 0.8 1.0

Allelic frequency class

Figure 2.7 Qualitative L-shape distribution between allele frequency class and proportion of alleles within each class for 194 harbour porpoises assayed at eight microsatellite DNA loci.

2.4 Discussion

Unlike some other areas of the harbour porpoise range, the coastline of British

Columbia appears to contain a single population. My results reject any population

subdivision between locations that might reasonably be expected to differ (pairwise FST

< 0.005), however areas such as the Baltic have three populations of harbour porpoises

(Pairwise FST = 0.04-0.05, De Luna et al. 2012). These results of a single population are in

contrast to the findings of Chivers et al. (2002) who suggested a fairly high level of subdivision of harbour porpoises within the waters of British Columbia and northern

Washington using smaller sample sizes and a broader study area less focused on this particular region. High levels of structure in other areas are likely explained by geographic barriers which limit movements (and hence gene flow) (Fontaine et al.

2007). In view of this, it is difficult to explain how the complex geography of the British

Columbia coastline is occupied by a single population of harbour porpoises. Along the coast of France, only a single population has been detected, although it has been suggested that two pre-existing populations merged following contact caused by anthropogenic habitat disturbance (Alfonsi et al. 2012).

The combined set of samples in my study show comparable levels of genetic diversity to single populations from other studies (e.g., Rosel et al. 1999; Chivers et al.

2002; De Luna et al. 2012). Some of these previous studies used the same mtDNA region that I employed (D-Loop), and most had a similar (although slightly higher) number of microsatellite alleles. The microsatellite loci used in my study were tetranucleotide primers designed for the finless porpoise and tested on other porpoise species (Chen and Yang 2008). The microsatellite primers used in many of the other harbour porpoise studies were dinucleotide and designed for other cetacean species (Valsecchi and Amos

1996). Therefore, the choice of mtDNA region and microsatellite markers in my study should have the power to detect similar diversity levels as other studies on harbour porpoise. Based on an estimate of effective population size calculated in MLNE v1.0

(Wang 2001) of 10,000 individuals and using using PowSim v4.1 (Ryman and Palm

2006), I determined that my allele frequency data had adequate power (0.85) to detect genetic structure with FST values of >0.01, but only an estimated power of 0.13-0.26 to detect genetic structure for FST values closer to those seen in my population (FST = 0.001 -

0.0025). One way to increase power would be to boost sample sizes (e.g., my outside waters group had only 19 individuals) and/or increase the number if loci (Paetkau et al.

2004). Nonetheless, while FST values calculated with more samples or more loci might prove statistically significant, they likely would not have changed in absolute value appreciably to the levels (~0.05) reported by De Luna et al. (2012).

Population structure of non-migratory marine mammals in British Columbia is not well understood for most species. For species in which these studies have been undertaken, the patterns of population structure have a variety of driving forces. Killer whales in British Columbia belong to at least four separate populations that are maintained by differences in social dynamics and culturally-transmitted food preferences, while occupying the same geographic regions (Hoelzel et al. 1998; Barrett-

Lennard 2000; Baird 2001). Harbour seal (Phoca vitulina) population structure appears to be influenced by colonization patterns post-glaciation. Colonization after the last glaciation has resulted in two genetically distinct populations separated between Haida

G’waii and Vancouver Island (Burg et al. 1999).

At a finer resolution, the waters of British Columbia present unique environmental gradients that may influence different species to be structured in different ways. For example, unlike in harbour porpoises where the genetic diversity across British Columbia is probably maintained by the movement of adults, in Copper

Rockfish (Sebastes caurinus) population structure is influenced by larval dispersal

(Buonaccorsi et al. 2002). While larval dispersal is also considered a driving force for maintaining genetic diversity of Dungeness crab (Metacarcinus magister) in British

Columbia, their movement is influenced by the geography of each inlet and the genetic structure corresponds accordingly (Beacham et al. 2008). Population structure in the north Pacific shows similar trends for many species. In the north Pacific, there has been

separation between two genetically distinct Steller Sea Lion (Eumetopias jubatus) populations: one in the Aleutian Islands stretching over to Japan, and a second from southeastern Alaska down to northern California (Hoffman et al. 2006). Dall’s porpoises live in three populations in the western, central and northeastern Pacific (Escorza-

Treviño and Dizon 2000). At a broad scale, many species exhibit similar patterns of population structure, but these population borders are rarely identical. Harbour porpoises throughout the north Pacific come from two broader genetic populations which may overlap along the coast of British Columbia perhaps north of my study area

(Taguchi et al. 2010), and the population from British Columbia south to California exhibits more genetic structuring at a finer scale (Chivers et al. 2002). Extending the boundaries of my study region, or more samples from northern British Columbia may allow detection of more broad scale population structure.

The processes that seem to have resulted in a single genetic population of harbour porpoises in southern British Columbia are open to conjecture. Individual range size of harbour porpoises has received little attention; however, satellite tagging has recorded daily movements of nearly 60km (Read and Westgate 1997). Being relatively solitary animals, it is possible that harbour porpoises in British Columbia travel long distances along the coastline and mate arbitrarily on their way. An alternative, however, is that some aspect(s) of harbour porpoise biology and behaviour actually favour panmixia. Throughout their range, harbour porpoises are typically found in very small groups of 1-3 individuals. In British Columbia, however, large aggregations have been reported with groups of over 200 individuals. More than 60 of these aggregations of over 50 animals have been reported to the BC Cetacean Sightings

Network over the past 10 years (C. Birdsall, BC Cetacean Sightings Network, pers. comm., 2012) occurring all along the coast and at all times of the year, although they

appear to be more common in May through September (Hall 2011). The potential causes of this social behaviour are yet unexplored. Like many other cetacean aggregations, these groups could be driven by prey availability and distribution (Calambokidis et al.

2002; Canning et al. 2008; Hall 2011). Large accumulations of prey could be detectable by harbour porpoises from many regions and result in individuals from a wide range congregating for feeding. These groups could also be used as mating aggregations.

Alternatively, transient killer whales are relatively abundant in BC keeping harbour porpoise densities fairly low. As a result, harbour porpoises may roam widely in search of mates maintaining panmixia, whereas areas of high harbour porpoise density may exhibit more structure.

Relatively little is known about harbour porpoise mating patterns. They have a long (11 month) gestation period (Boness et al. 2002), and a long calving season signals a potentially long breeding season peaking in June through August (Read and Hohn

1995; Lockyer et al. 2001; Hall 2011). Both large mating aggregations and long movement distances could contribute to the maintenance of a fairly homogenous population across a large landscape. If these mating groups are the driving force maintaining a single genetic population, it would require that individual harbour porpoises are coming from different regions, aggregating to mate, and re-dispersing to different areas. Testing this idea is much harder in harbour porpoises than in many other species because individual identification is virtually impossible. Satellite tagging of multiple individuals in these groups is, therefore, an important next step to explain dispersal patterns of individuals to test this idea.

2.4.1 Hybridization

My results confirm previous hypotheses that harbour x Dall’s porpoise hybridization occurs and that reproductively viable hybrids are capable of backcrossing with either parental species, with a tendency toward backcrossing with Dall’s porpoises over harbour porpoises. While most backcrosses seem to be occurring with Dall’s porpoises, it appears that, historically, there may have been some crosses with harbour porpoises, indicating that crosses in both directions are possible. I have identified hybrids from a much larger geographic range than was covered by previous specimens and sightings. This then raises the question whether hybridization is occurring much more commonly (e.g., over the entire region of range overlap) and simply goes unnoticed, or whether there is something unusual about the coast of British Columbia that might be promoting these inter-specific matings. Further work is needed to confirm why these two species mate with each other and why the crosses appear to be primarily unidirectional.

What is causing these species to hybridize and promoting backcrosses? As will be discussed in more detail in Chapter 3, cetacean hybridization is not uncommon and similarity in certain ecological, morphological and/or behavioural traits might enable or encourage hybridization. There may be, however, certain behaviours specific to either of these parental species that facilitate their mating.

In addition to helping us understand the genetic population structure, the aggregating behaviour of harbour porpoises could provide an explanation for the production of these hybrids. In some cetacean species, there is evidence of coercive mating (Scott et al. 2005), while there is no direct evidence of this for porpoises, very little is known about porpoise mating behaviour. If harbour porpoises do practice

coercive mating, these large aggregations would offer an opportunity for many males to gather together and for these forced matings to occur. If these males were together and showed high mating motivation, a nearby female Dall’s porpoise could easily become involved in such forced matings. Such coercive mating might provide one way whereby female mate preference might be suppressed, leading to these hybridization events.

This could also explain why first generation hybrids tend to usually result from a male harbour porpoise mating with a female Dall’s porpoise. Furthermore, because these hybrid calves are then raised by a Dall’s mother, they will most likely identify with

Dall’s porpoises and thus should be more likely to mate with another Dall’s porpoise.

To address the frequency of hybridization, a study using biopsy darting of live

Dall’s porpoises would be necessary. Hybrids often face fitness costs including sterility and/or low survivorship (Stebbins 1958). Due to the high likelihood that there is some fitness disadvantage for hybrid porpoises, a stranding record could show a bias, containing a higher proportion of hybrids than are actually found in the wild. Such a bias could only be identified by sampling a large number of live animals.

2.4.2 Contributions

My study makes three main contributions to the study of porpoises in British

Columbia. First, it provides critical information for conservation managers who should now recognize a single population of harbour porpoises off the coast of southern British

Columbia. This is important because management actions or habitat issues that are localized to one portion of the range in southern BC would seem likely to have potential demographic and genetic impacts throughout the range. Important management decisions will rely on a better understanding of harbour porpoise mating strategies that are essential for maintaining the genetic diversity of the entire population. Second, the

study contributes to an understanding of porpoise hybridization and highlights the directions for future research that will be essential to determine the specific conservation needs of these hybrids. Finally, it provides baseline data that will make it possible in the future to detect long-term changes to genetic diversity in the population

(Schwartz et al. 2007).

3 An Analysis of Cetacean Hybridization

3.1 Introduction

Aggregating in groups can have both benefits and consequences. Individuals living in groups may have higher incidences of parasitism, disease or inbreeding (Côté and Poulin 1995; Loehle 1995; Pusey and Wolf 1996) than individuals living alone; however, groups can also facilitate prey capture, and group defense (Alexander 1974).

Group living is present broadly across animal taxa including insects, fish, birds and mammals (Alexander 1974). The benefits and consequences of these aggregations are not limited to groups of a single species. Mixed species assemblages form for many of the same reasons as single species groups, but they introduce a different consequence – hybridization.

Hybridization holds potential risks for both the fitness of individual hybrid offspring and for the overall future of the parental species. Most interspecific copulations do not result in offspring, and are often thought of as an expression of dominance, practice to increase the chance of success in intraspecific mating, or merely social interactions (Vasey 1995). Interspecific copulation has many obstacles to overcome in order to result in a viable hybrid (Orr 1995). For this to occur there needs to be no pre- or post-zygotic reproductive isolating barriers: genitalia need to be compatible, sperm needs to survive and be able to fuse with an egg, the species need to have a compatible number of chromosomes, and the hybrid offspring need to survive barring other genetic incompatibilities. Many hybrids that do survive until birth have sterile offspring, with higher instances of sterility in the heterogametic sex than the homogametic sex (Haldane 1922; Noor 1999). If in fact two species are able to overcome these hurdles without fitness disadvantages they may eliminate pure strains of a

parental species (Mallet 2007). If the hybrids are able to outcompete one or both the parental species, they may increase in frequency and displace them from the region of overlap (Rhymer and Simberloff 1996; Huxel 1999).

Hybridization is not uncommon in some classes of animals (e.g. birds, Grant and

Grant 1992; fishes, Scribner et al. 2001), but is rare for terrestrial mammals (Gray 1954).

Within the order Cetacea, close to 20% of species have been implicated in hybridization events (Table 3.1 and 3.2). Cetaceans are a recent radiation, where most of the cetacean diversity on Earth has arisen in the past 10 million years (McGowen et al. 2009; Slater et al. 2010). Cetaceans are also known to have slow molecular clocks (Hoelzel et al. 1991;

Schlötterer et al. 1991). In combination, these factors likely account for the fact most cetaceans have a common chromosome count (2n = 44) and karyotypic arrangement

(Árnason and Benirschke 1973; Árnason et al. 1977; Pause et al. 2006). The only exceptions are that sperm, beaked and right whales which have 42 chromosomes.

Excluding these exceptions, almost 50% of oceanic cetacean species are known to have hybridized.

Table 3.1 Documented cases of cetacean hybridization in captivity. Paternal Species Maternal Species Source Sotalia guianensis Tursiops truncatus Caballero and Baker 2010 Tursiops truncatus Steno bredanensis Dohl et al. 1974; Shallenberger and King 1977 Grampus griseus Tursiops truncatus Shimura et al. 1985; Miyazaki et al. 1992 Lagenorhynchus Tursiops truncatus Miyazaki et al. 1992 obliquidens Tursiops truncates Globicephala macrorhynchus Antrim and Cornell 1981 Delphinus capensis Tursiops truncatus Zornetzer and Duffield 2003 Pseudorca crassidens Tursiops truncatus Nishiwaki and Tobayama 1982

Table 3.2 Documented cases of cetacean hybridization in the wild. Species 1 Species 2 Source Balaenoptera physalus Balaenoptera musculus Spilliaert et al. 1991; Berube and Aguilar 1998 Delphinus capensis Lagenorhynchus obscurus Reyes 1996 (Probable) Balaenoptera Balaenoptera bonaerensis Glover et al. 2010 acutorostrata (probable) Tursiops truncates Stenella frontalis Herzing et al. 2003 Grampus griseus Tursiops truncatus Shimura et al. 1985; Miyazaki et al. 1992; Fraser 1940 Tursiops aduncas Tursiops truncatus Martien et al. 2011 (unknown) Stenella attenuata Stenella longirostiris Silva-Jr et al. 2005 Stenella clymene Stenella longirostiris Silva-Jr et al. 2005 Lissodelphis peronei Lagenorhynchus obscurus Yazdi 2002 Phocoena phocoena Phocoenoides dalli Baird et al. 1998; Willis et al. 2004; Chapter 2 (both directions) Pseudorca crassidens Tursiops truncatus Nishiwaki and Tobayama 1982 Monodon monoceros Delphinaptera leucas Heide-Jørgensen and Reeves 1993 Tursiops truncatus Sousa chienensis Karczmarski et al. 1997 (Possible)

Hybrid cetaceans have been documented both in captivity and in the wild. While hybridization in captivity does not prove that hybridization occurs in the wild, it does signify the potential for different species to create hybrid offspring. Identification of hybrid cetaceans dates back to whaling industry of the 1800s; whaling records of report catches whose size and coloration were intermediates of blue whale (Balaenoptera musculus) and fin whale (Balaenoptera physalus) (Spilliaert et al. 1991). Many hybrids are still identified in the field based on intermediate morphological features (e.g. Spilliaert et al. 1991; Herzing et al. 2003; Silva-Jr et al. 2005) instead of genetic analyses.

Morphometric analyses of dead specimens can reveal many intermediate traits that can

not be easily measured in their live counterparts; therefore, live hybrids in the wild can easily go unnoticed. Confirmation of these potential hybrids is best done using genetic techniques; however, samples are often expensive to collect. Without the ability to genetically test all potential hybrids, identification is accepted using morphological evidence.

At least seven instances of hybridization in captivity have been recorded between pairs of cetacean species (Table 3.1). At least two of these hybrids were fertile and produced backcrossed offspring (Zornetzer and Duffield 2003; Maines and Kestin

2009), and the fertility of the others is unknown. Infant mortality is high in cetaceans born in captivity (DeMaster and Drevenak 1988). Consequenty, it is difficult to determine whether high incidents of infant mortality in hybrids would be related to hybridization, or the nature of captive breeding with poor records of breeding and calving activity. While hybridization in captivity occurs under artificial conditions, many pairs of species also have natural range overlap, and are known to form mixed species assemblages (See Appendix S2.1). This suggests that interspecific matings among these pairs of species are also possible in the wild.

Unfortunately, most wild hybrids are not followed to maturity, and in many cases the sex of the hybrid is unknown. However, in both wild and captive species pairs, there is evidence of reproductively viable hybrids and successful backcrosses

(Spilliaert et al. 1991; Baird et al. 1998; Odell and McClune 1999). While fertile female hybrids have been confirmed, determining male fertility is difficult. In hybrids generally, the fertility of the heterogametic sex – males for cetaceans (Árnason 1974) – is usually the most negatively affected (Haldane 1922). It is therefore difficult to estimate the impact of hybridization at the population level.

3.1.1 Research objectives

While the long-term impacts of hybridization on cetacean populations are not well understood, it seems clear that there are relatively few post-mating, including post- zygotic, barriers to hybridization. Consequently, the question arises: what promotes or enables hybridization in cetaceans? Are there certain behavioural, morphological or ecological traits that might predispose certain species pairs to hybridize? In this study I tested for associations between trait similarity and ability to hybridize across all species pairs of marine cetaceans.

3.2 Methods

3.2.1 Data collection

I conducted a literature search to collect data on the ecological, morphological and behavioural traits of the 78 species of marine cetaceans. The following traits were examined: male body length, female body length, visible sexual dimorphism (size, colour etc.), group size, species’ range size, water depth, water temperature, prey species, predator species, parasite species, known associate species, natural range overlap between each species pair and vocalization frequency. These were chosen to depict both gross morphological characteristics of a species, as well as ecological traits that would define their specific niche (Aldridge and Rautenbach 1987; Gowans and

Whitehead 1995; Vanhooydonck et al. 2000; Bearzi 2005). When possible, information was collected from sources referencing various parts of the species’ (Appendix S2.1).

3.2.2 Similarity index

For each trait, I calculated an index of similarity for each species’ pair using similar methods to those presented by Geange et al. (2011). Traits that were not

described in the scientific literature for a given species were assigned ‘Not Available’

(NA) and were removed from the analysis for comparisons within all pairs for that species. The similarity of traits was calculated differently depending on the type of trait data being assessed. Two species that shared 100% of their traits would receive a similarity index of ‘1’ and pairs of species with no overlap in any traits would have a similarity of ‘0’.

Presence/absence – (species pair range overlap, sexual dimorphism)

The matrix for natural range overlap of species pairs was built by simply assigning ‘1’ to pairs of species’ that overlap in their ranges, and a ‘0’ where they do not.

Sexual dimorphism was examined in a similar fashion; if the two species are both sexually dimorphic or if neither is sexually dimorphic, I assigned them a similarity index of ‘1’. If one species shows dimorphism and the other does not, the pair was assigned a value of ‘0’.

Continuous traits – (male body length, female body length, water temperature)

These traits analyzed as a continuous range of body size of each sex at physical maturity and of preferred water temperature. If the ranges of trait values for the two species did not overlap, I assigned the pair a similarity of ‘0’. If the trait value of the two species overlapped, I calculated the amount they overlapped and divided this value by the size of the smaller range in trait values of the two species (Eq. 3.1). This would result in a percentage of overlap of the trait relative to the more specialized species with the smaller range in trait values.

(Smaller of Max or Max ) (Larger of Min or Min ) 1 2 1 2 Smaller of (Max1 Min1 or Max2 Min2)

Continuous traits as categorical data – (group size, species’ range size, water depth, vocalization frequency)

I took the average value of each trait for each species and grouped them into categories: group size (solitary: 1-5 individuals, medium: 5-50 individuals; social: 50+ individuals), species’ range size (small: <10,000,000 km2; medium: 10,000,000-

100,000,000 km2; large: >100,000,000 km2), water depth (shallow: 0-200 m; medium: 200-

1000 m; deep: >1000 m), and vocalization frequency (low: 0-5 kHz; medium: 5-10 kHz; high: >10 kHz). If two species occupy the same category (i.e. both solitary), I assigned the pair an index of ‘1’. If two species occupy categories at either end of the spectrum

(ie. one species with small range size and one with large range), the pair was assigned

‘0’. If one of the species occupies an intermediate value and the other does not (i.e. one species from medium water depth and one from deep water), I assigned the pair a similarity index of ‘0.5’.

Categorical traits – (prey species, predator species, parasite species, known associate species)

I examined prey, predators and parasites at the family level to help account for global variation in species’ distribution. For these traits, I calculated the number of shared families or species, and divided that by the total number of families or species encountered by both cetacean species (Eq. 3.2). I used this value as the similarity between species pairs for these categorical traits.

Number of shared prey, predator, parasite families or known associate species

( )

Total similarity index

The total similarity index was calculated by taking both an un-weighted and a weighted average of the similarities of each trait. The un-weighted index was simply the mean of all similarity indices for each trait for each species pair (Eq. 3.3).

∑ Individual trait similarity indices

Number of traits

A weighted average was calculated by conducting a survey of expert opinion regarding the relative importance each trait might have on the predisposition of species to hybridize (Appendix S2.3). These weightings were averaged across survey participants and each trait received a weight that represented a proportion of the predisposition to hybridize. I applied each the weightings to their respective traits and summed these to achieve a weighted index of similarity again from ‘0’ to ‘1’ (Eq. 3.4).

∑

3.2.3 Hybridization and similarity index

In order to assess whether species that have been known to hybridize are more or less similar in these traits than species pairs that do not hybridize, I conducted a Mantel test in R v.2.12.2 (R Project for Statistical Computing) using the Kendall method from the package vegan (Oksanen et al. 2011). I compared the matrix of trait similarity and a matrix of all possible species pairs where ‘0’ represents a non-hybridizing pair, and ‘1’ indicates a known hybridization event. I omitted the diagonal from the analyses to avoid a bias because it represents the mating between the same species and has a similarity of ‘1’ in every instance. In order to determine which traits might be driving

trends in either similarity or dissimilarity, I conducted a principal components analysis

(PCA) in R using the covariance matrix. I included all 13 species traits as factors and used the prcomp() function in R for increased numerical accuracy and because the inputted trait matrix was not raw values. I present all principal components that account for at least 10% of the variance.

With the assumption that a different number of chromosomes or chromosomal arrangement will prevent or strongly deter hybridization (Dobzhansky 1935), I repeated the analyses with only cetaceans with the same chromosomal number. Therefore, those

26 species with 42 chromosomes were removed from the analyses. The Mantel test and

PCA with all comparisons were performed twice for both all species comparisons and for species with 44 chromosomes: once with the raw similarity index, and again with the similarity index weighed according to expert opinion.

My data for the PCA were pairwise comparisons and therefore violate an assumption of the test. I included these results to allow for comparison between the relative importance of each trait, and also conducted a test to remove pairwise comparisons. This consisted of subsampling random pairs of species, so that each species was represented in a single comparison and using the similarity index of these randomly chosen pairs, I conducted a PCA. I replicated this PCA 10,000 times using different combinations of species pairs. I averaged the eigenvectors for each trait in each of the first four principal componants to account for variations across the 10,000 iterations and to obtain a single eigenvector for each trait.

3.3 Results

3.3.1 Un-weighted analysis

Pairs of species that are known to hybridize are more similar in ecological and morphological traits than species pairs that do not (r = 0.077, P = 0.001, Fig. 3.1). The r- values for the correlation are low because there are many more pairs of species that do not hybridize (n = 6048) than pairs of species that do hybridize (n = 36). A principal components analysis accounted for 68.8% of the variation in the traits examined across the first four PC axes (Table 3.3). The first principal component, which accounted for almost 25% of the variation in traits, was driven strongly by similarities in sexual dimorphism between the species, with species range size, body length of both sexes and vocalization frequencies also playing strong roles having the greatest (+ or -) eigenvectors (Table 3.3). In the next three principal components (PC2 – PC4), similarities in water temperature and natural range overlap were the more important features with body length again contributing to the similarity. Average group size was also a strong contributor to PC2.

Non-hybridizing Hybridizing species pairs species pairs

Figure 3.1 Similarity index of non-hybridizing species pairs (n = 6048) and hybridizing species pairs (n = 36) for all cetacean species comparisons.

Table 3.3 Eigenvectors of the first four principal components of variation in similarity of traits for all cetacean species comparisons. Variables that are more important for each principal component have larger values (+ or -). Trait (ALL) PC1 PC2 PC3 PC4 Male Body Length -0.276 -0.318 0.285 -0.400 Female Body Length -0.275 -0.304 0.357 -0.366 Sexual Dimorphism -0.658 0.423 -0.197 -0.047 Range Size -0.436 0.292 -0.120 -0.024 Water Depth -0.187 -0.009 -0.145 0.153 Water Temperature -0.159 -0.452 -0.452 0.436 Prey Species -0.134 -0.086 0.018 -0.030 Predator Species -0.122 -0.124 -0.159 -0.035 Parasite Species -0.153 -0.083 0.036 -0.021 Average Group Size -0.188 -0.266 0.072 0.322 Known Associate -0.124 -0.094 -0.010 -0.019 Species Natural Range Overlap 0.021 -0.387 -0.577 -0.461 Vocalization Frequency -0.239 -0.283 0.383 0.413 Proportion of Variation 24.32% 19.76% 14.63% 10.09% Accounted For

The results were consistent for the subset of species with the same chromosomal number (2n = 44) (Mantel test: r = 0.116, P = 0.001, Fig. 3.2). The principal component analysis also pointed toward similar explanatory traits where the same traits seem to be contributing to the variation in similar proportions in each of the first few principal components (Table 3.4).

Non-hybridizing Hybridizing species pairs species pairs

Figure 3.2 Similarity index of non-hybridizing species pairs (n = 2704) and hybridizing species pairs (n = 36) for cetacean species with 44 chromosomes.

Table 3.4 Eigenvectors of the first four principal components of variation in similarity of traits for all cetacean species with 44 chromosomes. Variables that are more important for each principal component have larger values (+ or -). Trait (2n = 44) PC1 PC2 PC3 PC4 Male Body Length -0.339 -0.294 0.200 0.458 Female Body Length -0.347 -0.280 0.283 0.420 Sexual Dimorphism -0.540 0.589 -0.123 0.023 Range Size -0.357 0.395 -0.071 0.019 Water Depth -0.166 0.066 -0.115 -0.134 Water Temperature -0.207 -0.323 -0.531 -0.340 Prey Species -0.152 -0.051 -0.020 0.028 Predator Species -0.134 -0.060 -0.215 0.050 Parasite Species -0.175 -0.046 0.002 -0.018 Average Group Size -0.243 -0.227 -0.023 -0.318 Known Associate -0.140 -0.065 -0.054 0.029 Species Natural Range Overlap -0.034 -0.251 -0.630 0.360 Vocalization Frequency -0.352 -0.310 0.342 -0.495 Proportion of Variation 25.03% 19.04% 15.20% 10.60% Accounted For

The 10,000 replicated PCAs with subsampled pairs of species that were found to be similar for all species comparisons revealed that the traits with the strongest influence in the first principal component were male body size, water depth and temperate and natural range overlap. Female body size and sexual dimorphism also showed large eigenvectors in PC2 (Appendix S2.4). In analyses of species with 44 chromosomes, sexual dimorphism, water depth and range size were traits with the strongest influence on PC1, with overlap and body size also contributing to the variance. Both sexual dimorphism and range size high very strong impact on PC2

(Appendix S2.5). The relative influence of each of the traits was similar to their importance from the PCA with all species pairs.

3.3.2 Weighted analysis

The survey of expert opinion elicited 41 responses, and I calculated the average weighting for each trait (Table 3.5). The new weightings had little influence on the results and the ability to hybridize was still significantly correlated with similarity in morphological and ecological traits (all species: r = 0.077, P = 0.001, Fig. 3.3; 44 chromosomes: r = 0.116, P = 0.001). The PCA accounted for 75.0% of the variation in the traits examined across the first four PC axes (Table 3.6), and for all species comparisons indicated slight differences in which traits may be driving this pattern (Table 3.6). The first PC accounted for 26% of the variation and was driven largely by natural range overlap, water temperature, sexual dimorphism and range size. In the next three PCs

(PC2-PC4), many of the driving traits were the same as in the un-weighted analysis, with more influence from natural range overlap. In PC2, the variable “known associate species” was also a large contributing factor. The analysis of 44-chromosome species only showed very similar patterns of the influence of each trait (Table 3.7)

Table 3.5 Results of the survey (N = 41) to calculate weighted traits for assessing relative influence of each trait would have on the ability of pairs of cetacean species to hybridize. Trait Range of Weightings Average Male Body Length 0-0.250 0.085 Female Body Length 0-0.250 0.080 Sexual Dimorphism 0-0.400 0.088 Range Size 0-0.273 0.067 Water Depth 0-0.200 0.086 Water Temperature 0-0.136 0.086 Prey Species 0-0.116 0.055 Predator Species 0-0.105 0.032 Parasite Species 0-0.136 0.035 Average Group Size 0-0.182 0.059 Known Associate Species 0-0.500 0.112 Natural Range Overlap 0-0.450 0.120 Vocalization Frequency 0-0.210 0.095

Non-hybridizing Hybridizing species pairs species pairs

Figure 3.3 Weighted similarity index of non-hybridizing species pairs (n =

6048) and hybridizing species pairs (n = 36) for all species comparisons.

Table 3.6 Eigenvectors of the first four principal components of variation in the weighted similarity of traits for all cetacean species comparisons. Variables that are more important for each principal component have larger values (+ or -). Trait (ALL) PC1 PC2 PC3 PC4 Male Body Length -0.064 -0.320 0.335 -0.292 Female Body Length -0.018 -0.293 0.352 -0.283 Sexual Dimorphism 0.255 -0.652 -0.520 -0.061 Range Size 0.138 -0.316 -0.255 -0.028 Water Depth -0.005 -0.198 -0.079 0.240 Water Temperature -0.305 -0.249 0.133 0.828 Prey Species -0.013 -0.090 0.036 -0.009 Predator Species -0.042 -0.047 -0.008 0.012 Parasite Species -0.009 -0.066 0.026 -0.022 Average Group Size -0.042 -0.137 0.148 0.073 Known Associate Species -0.047 -0.181 0.065 0.026 Natural Range Overlap -0.900 -0.115 -0.247 -0.283 Vocalization Frequency 0.041 -0.327 0.562 -0.026 Proportion of Variation Accounted For 26.03% 21.71% 16.64% 10.66%

Table 3.7 Eigenvectors of the first four principal components of variation in the weighted similarity of traits for cetacean species comparisons with 44 chromosomes. Variables that are more important for each principal component have larger values (+ or -). Trait (2n = 44) PC1 PC2 PC3 PC4 Male Body Length -0.068 0.373 0.249 0.408 Female Body Length -0.003 0.359 0.259 0.383 Sexual Dimorphism 0.175 0.520 -0.678 0.024 Range Size 0.094 0.254 -0.332 0.020 Water Depth -0.005 0.158 -0.128 -0.173 Water Temperature -0.317 0.228 0.073 -0.744 Prey Species -0.018 0.095 0.007 0.008 Predator Species -0.054 0.042 -0.021 -0.010 Parasite Species -0.011 0.072 0.008 0.005 Average Group Size -0.049 0.168 0.106 -0.121 Known Associate Species -0.072 0.185 0.014 -0.016 Natural Range Overlap -0.915 0.035 -0.167 0.223 Vocalization Frequency 0.088 0.490 0.489 -0.195 Proportion of Variation 24.50% 22.74% 17.66% 10.75% Accounted For

3.4 Discussion

My literature review and analysis suggest that pairs of cetacean species that have been known to hybridize are more similar in their ecological, morphological and behavioural traits than their counterparts that do not. This pattern seems to be driven largely by similarities between species in the extent of sexual dimorphism, body length, geographic range size, and vocalization frequency. Similarities in body size and state of sexual dimorphism among hybridizing pairs of species may indicate a component of poor visual discrimination where species are unable to visually identifiy conspecifics

and vocalization frequency may indicate poor acoustic discrimination between species.

The similarity in the the size of the entire species’ range could indicate behavioural similarity between the species, where individals of a species with a large range size may also have large home range sizes, which could influence their speed and distance of their daily movement patterns. These traits may, therefore, play a large role in how cetaceans discriminate between species – perhaps both acoustically and visually (based on both behaviour and appearance).

3.4.1 Species barriers

My data suggest that morphological and behavioural traits drive patterns in the similarity index. Therefore high similarity between hybridizing species, could indicate mistakes in species recognition, whereas relatively fewer similarities in ecological traits could provide the necessary opportunities for these interspecific copulations. Higher similarity of morphological and behavioural traits among hybridizing pairs of species suggest that species may be basing mating decisions on morphological and behavioural cues instead of ecological cues realted to habitat choice. This suggests that species barriers in cetaceans may be maintained primarily by morphological and behavioural traits with little influence of ecological traits.

These results are also consistent with patterns of increased hybridization between closely related or sister species that may have diverged in ecological traits, but remain similar in morphological and/or behavioural traits. To tease apart these mechanisms, genetic data are needed to support phylogenetically-independent contrasts to control for genetic distance between pairs of species. It is important to highlight that there are many more species traits that clearly help define a species’ morphology, behavioural and ecological niche that were not included my study, and

therefore many more traits should also be exmined to help better define the niche of each species. If genetic distance does not have a large influence on the results, this would further support that species barriers in cetaceans are being maintained by morphological and ecological traits.

3.4.2 Potential benefits of interspecific mating

In cetaceans, mating behaviour is not just limited to single adult male-female pairs during the breeding season. Mating behaviour is witnessed year-round (e.g.,

Shane et al. 1986) and is often seen between individuals of different age classes (e.g., reproductively immature calves, Herzing 1997), same sex pairs (e.g., male-male copulations, Mann 2006), and even interspecific pairs (e.g., Stenella and Tursiops;

Herzing et al. 2003). In many instances, these copulations offer no possibility of offspring, so why devote time and energy to these activities that have no reproductive potential? There may be some benefits to these mating attempts that outweigh the efforts.

It has been hypothesized that cetaceans might exhibit mating behaviour as a form of social play (Brown and Norris 1956; Herzing and Johnson 1997). Social play is expressed through sexual behaviour in other higher mammalian taxa such as primates

(Vasey 1995). This social play could be solely for entertainment, or it could be used to establish a dominance hierarchy between individuals (Vasey 1995). Established dominance roles can be important for daily interactions between individuals in a larger group. Alternatively, these ongoing mating attempts could be for practice (Mann 2006).

Mating success is essential for passing genes on to subsequent generations and as a result, it is extremely important that individuals are able to complete successful mating attempts in a potentially narrow fertility window. A male that is able to practice mating

with no negative consequences may have a higher chance of reproductive success during the breeding season compared to conspecific males, and may therefore experience an increase in the probability and number of offspring that season.

3.4.3 Conclusions

Could widespread hybridization in cetaceans be the result of incomplete speciation? Most of the diversity within the order Cetacea has arisen over the past 10 million years (McGowen et al. 2009; Slater et al. 2010). Slow ecological or parapatric speciation in cetaceans could have lingering hybrids and their prevalence may be the result of an incomplete speciation process. A million years from now, will there be any more natural cetacean hybrids? Without known rates of hybridization over time it is difficult to assert whether or not hybridization is increasing, or slowly fading away.

Information obtained from implementing long-term genetic monitoring programs can allow for monitoring and conservation of individual cetacean species, and can enable documentation of hybridization events and the ability to follow these occurrences over time.

The results of my study suggest that pairs of species involved in known hybridization events are more similar in their ecological, morphological and behavioural traits than those which have not. This pattern seems to be driven by traits which could contribute to species recognition via visual and acoustic measures and suggests that a poor ability to, or a disinterest in discriminating between species may lead to increased cases of hybridization.

4 Conclusions

4.1 Summary of findings

My study addresses some of the knowledge gaps for harbour porpoises in British

Columbia and contributes to a better understanding of not just porpoise hybridization, but hybridization of cetaceans as a whole. Major findings include:

1. Unlike many other parts of their range, harbour porpoises in British Columbia

show little sign of population structure and appear to belong to a single

panmictic population.

2. Harbour and Dall’s porpoises hybrids are reproductively viable and successful

backcrosses are ongoing with both parental species (although more often with

Dall’s porpoise).

3. Pairs of species of cetaceans that hybridize are more similar in their

morphological and ecological traits than non-hybridizing species pairs. Traits of

particular importance in cetacean hybrid pairs are sexual dimorphism, body

length, geographic range size and vocalization frequency.

4.2 Studies using samples of bycatch and strandings

As an increasing number of species are facing risk of extinction largely due to anthropogenic causes (Butchart et al. 2010), it is critical to not exasterbate these effects and therefore minimally invasive methods are necessary to investigate the specific needs of species at risk. Studies using stranded animals and those accidentally caught in fishing gear, such as my study, can be used to learn more about a species. While studying these incidents can provide a lot of opportunistic data, the results can also be difficult to interpret.

Many factors may contribute to the probability of a cetacean stranding or entangling in fishing gear, including parasites (Stroud and Roffe 1979), neonatal abandonment (Kirkwood et al. 1997), injuries sustained from human activities

(Kirkwood et al. 1997), etc. This means that the stranding record alone might not give a random sample of individuals from the population. For example, in a study of stomach content analysis on cetacean samples retained from by-catch, the diet could be highly skewed towards the species targeted by the fishery in which the cetaceans were caught.

In analyses of population structure, such as in Chapter 2, it has been suggested that stranding records may underestimate the levels of population structure as all sub- populations might not be represented equally (Bilgmann et al. 2011). Therefore, it is important to consider even low levels of structure as potentially having a much larger impact on the population as a whole. Low sample sizes in Chapter 2 mostly obtained from stranded carcasses could therefore be masking potentially low levels of population structure. As hybrids often have reduced fitness compared to their parental species

(Stebbins 1958), a stranding record could artificially inflate the proportion of hybrids in the population. As in Chapter 2, studies that genetically identify hybrids in the wild can be extremely useful in confirming the reproductive viability of hybrids and the directionality of hybrid crosses. Conservation efforts can be greatly aided by information from bycatch studies highlighting how parental species may be affected by increased rates of uni-directional hybridization.

4.3 Future directions

Because harbour porpoises in British Columbia are one of the most abundant cetaceans on the British Columbia coast, understanding the role they play in the ecosystem and their specific needs and threats is an important to setting conservation

priorities as the focus is switching from species-specific conservation plans to complete ecosystem based approaches to conservation (Sherman et al. 2005). Based on the results of this study, I suggest research be conducted directly on large aggregations of harbour porpoises to understand not only where they come from, but also the possible function of such aggregations. Hall (2011) suggests large aggregations could be a social phenomenon, for feeding, or for cooperative detection of predation, however additional study on these events will enhance the understanding of the behavioural changes that take place and the consequences of potential anthropogenic threats to these large aggregations.

Hybridization between harbour and Dall’s porpoises seems to be unique to

British Columbia and the adjacent waters of Washington. In order to gain an accurate assessment of ongoing levels of hybridization between harbour and Dall’s porpoises, I propose a slightly more invasive, yet commonly practiced study that would use biopsy sampling of Dall’s porpoises to complete a full genetic assessment of hybridization.

Other areas of range overlap (i.e., the United States or Japan) between the two species could also be included in a biopsy darting study or further genetic analysis of stranded and by-caught Dall’s porpoises to understand the geographic extent of these hybrids.

There is already evidence of introgression in some cetacean species (e.g., harbour and Dall’s porpoise, Chapter 2; blue and fin whale, Spilliaert et al. 1991); however, for many, very little genetic assessment has been conducted. With increasing availability and affordability of genetic analyses, it is becoming progressively more feasible to set up a genetic monitoring program for cetaceans. Such programs are important to help understand long-term trends in hybridization (and perhaps even uncover unknown hybrid crosses) as well as long-term fluctuations in genetic diversity. A better understanding of historical and present rates of hybridization could help us determine

why this pattern is so common in cetaceans. Monitoring changes to genetic diversity

(both with respect to hybrids and individual species) can help identify either sharp increases and/or decreases in diversity that may indicate changes in population size or potential threats to the populations. This can be of great aid to conservation management decisions as it may highlight changes or threats to the populations that are not always evident from field surveys.

Hybridization in marine cetaceans is far from being well understood. My study has identified correlations between similarity in traits and ability to hybridize, but this is only a starting point. More traits need to be examined and phylogenetic relationships need to be considered. A better understanding of the widespread hybridization in cetaceans might provide insight into the origins of diversity in the order and/or insights about potential threats of species collapse.

Making use of accidental mortalities, their potential biases notwithstanding, to establish a genetic monitoring program, could enable this abundant small cetacean to serve as an indicator for other marine mammals as more information could be obtained from higher rates of collected carceasses of this species than many other cetaceans in

BC. By examining harbour porpoises in the northeastern Pacific, my study helps to ensure complete range coverage of the study of this species and establishes a better understanding and precedent for monitoring the genetics of harbour porpoise.

References

Aldridge, H. D. J. N. and I. L. Rautenbach. 1987. Morphology, echolocation and resource partitioning in insectivorous bats. Journal of Animal Ecology 56:763–778.

Alexander, R. D. 1974. The evolution of social behavior. Annual Review of Ecology and Systematics 5:325–383.

Alfonsi, E., S. Hassani, F.-G. Carpentier, J.-Y. Le Clec’h, W. Dabin, O. Van Canneyt, M. C. Fontaine and J.-L. Jung. 2012. A European melting pot of harbour porpoise in the French Atlantic coasts inferred from mitochondrial and nuclear data. PLoS one 7:e44425. doi:10.1371/journal.pone.0044425.

Anderson, E. C. and E. A. Thompson. 2002. A model-based method for identifying species hybrids using multilocus genetic data. Genetics 160:1217–1229.

Antrim, J. E. and L. H. Cornell. 1981. Globicephala–Tursiops hybrid. in Abstracts of the 4th Biennial Conference on the Biology of Marine Mammals, 14–18 December. Society for Marine Mammalogy, Lawrence, KA, San Francisco, CA.

Árnason, Ú. 1974. Comparative chromosome studies in Cetacea. Hereditas 77:1–36.

Árnason, Ú. and K. Benirschke. 1973. Karyotypes and idiograms of sperm and pygmy sperm whales. Hereditas 75:67–74.

Árnason, Ú., K. Benirschke, J. G. Mead and W. W. Nichols. 1977. Banded karyotypes of three whales: Mesoplodon europaeus, M. carlhubbsi and Balaenoptera acutorostrata. Hereditas 87:189–200.

Baird, R. W. 1998. An interection between Pacific white-sided dolphins and a neonatal harbor porpoise. Mammalia 62:1291134.

Baird, R. W. 2001. Status of killer whales, Orcinus orca, in Canada. Canadian Field- Naturalist 115:676–701.

Baird, R. W. 2003. Update COSEWIC status report on the harbour porpoise Phocoena phocoena (Pacific Ocean population) in Canada, in COSEWIC assessment and update status report on the harbour porpoise Phocoena phocoena (Pacific Ocean

population) in Canada. Committee on the Status of Endangered Wildlife in Canada, Ottawa.

Baird, R. W. and T. J. Guenther. 1995. Account of Harbour Porpoise (Phocoena phocoena) strandings and bycatches along the coast of British Columbia. Reports of the International Whaling Commission. Special Issue No. 16:159–168.

Baird, R. W. and T. J. Guenther. 1994. Status of porpoises in British Columbia/Washington trans-boundary area: a Canadian perspective. Puget Sound Notes 34:5–8. Olympia, Washington.

Baird, R. W., P. M. Willis, T. J. Guenther, P. J. Wilson and B. N. White. 1998. An intergeneric hybrid in the family Phocoenidae. Canadian Journal of Zoology 76:198–204.

Barnes, L. G. 1985. Evolution, taxonomy and antitropical distributions of the porpoises (Phocoenidae, Mammalia). Marine Mammal Science 1:149–165.

Barrett-Lennard, L. G. 2000. Population structure and mating patterns of killer whales (Orcinus orca) as revealed by DNA analysis. PhD Thesis. University of British Columbia: Canada.

Bataillon, T. M., J. L. David and D. J. Schoen. 1996. Neutral genetic markers and conservation genetics: simulated gerplasm collections. Genetics 144:409–417.

Beacham, T. D., J. Supernault and K. M. Miller. 2008. Population structure of Dungeness crab (Cancer magister) in British Columbia. Journal of Shellfish Research 27:901– 906.

Bearzi, M. 2005. Habitat partitioning by three species of dolphins in Santa Monica Bay, California. Bulletin, Southern California Academy of Sciences 104:113–124.

Bilgmann, K., L. M. Möller, R. G. Harcourt, C. M. Kemper and L. B. Beheregaray. 2011. The use of carcasses for the analysis of cetacean population genetic structure: a comparative study in two dolphin species. PLoS one 6:e20103. doi:10.1371/journal.pone.0020103.

Boness, D. J., P. J. Clapham and S. L. Mesnick. 2002. Life history and reproductive strategies. Pp. 278–284 in A. R. Hoelzel, ed. Marine Mammal Biology: an evolutionary approach. Blackwell Science, Malden, MA.

Boyko, A. R., R. H. Boyko, C. M. Boyko, H. G. Parker, M. Castelhano, L. Corey, J. D. Degenhardt, A. Auton, M. Hedimbi, R. Kityo, E. A. Ostrander, J. Schoenebeck, R. J. Todhunter, P. Jones and C. D. Bustamante. 2009. Complex population structure in African village dogs and its implications for inferring dog domestication history. Proceedings of the National Academy of Sciences 106:13903–13908.

Brown, D. H. and K. S. Norris. 1956. Observations of captive and wild cetaceans. Journal of Mammalogy 37:311–326.

Bruford, M. W. and R. K. Wayne. 1993. Microsatellites and their application to population genetic studies. Current Opinion in Genetics and Development 3:939– 943.

Buonaccorsi, V. P., C. A. Kimbrell, E. A. Lynn and R. D. Vetter. 2002. Population structure of Copper Rockfish (Sebastes caurinus) relfects postglacial colonization and contemporary patterns of larval dispersal. Canadian Journal of Fisheries and Aquatic Sciences 59:1374–1384.

Burg, T. M., A. W. Trites and M. J. Smith. 1999. Mitochondrial and microsatellite DNA analyses of harbour seal population structure in the northeast Pacific Ocean. Canadian Journal of Zoology 77:930–943.

Butchart, S. H. M., M. Walpole, B. Collen, A. van Strien, J. P. W. Scharlemann, R. E. A. Almond, J. E. M. Baillie, B. Bomhard, C. Brown, J. Bruno, K. E. Carpenter, G. M. Carr, J. Chanson, A. M. Chenery, J. Csirke, N. C. Davidson, F. Dentener, M. Foster, A. Galli, J. N. Galloway, P. Genovesi, R. D. Gregory, M. Hockings, V. Kapos, J.-F. Lamarque, F. Leverington, J. Loh, M. a McGeoch, L. McRae, A. Minasyan, M. Hernández Morcillo, T. E. E. Oldfield, D. Pauly, S. Quader, C. Revenga, J. R. Sauer, B. Skolnik, D. Spear, D. Stanwell-Smith, S. N. Stuart, A. Symes, M. Tierney, T. D. Tyrrell, J.-C. Vié and R. Watson. 2010. Global biodiversity: indicators of recent declines. Science 328:1164–1168.

Bérubé, M. and A. Aguilar. 1998. A new hybrid between a blue whale, Balaenoptera Musculus, and a fin whale, B. Physalus: frequency and implications of hybridization. Marine Mammal Science 14:82–98.

Caballero, S. and C. S. Baker. 2010. Captive-born intergeneric hybrid of a Guiana and bottlenose dolphin: Sotalia guianensis x Tursiops truncatus. Zoo Biology 29:647–657.

Calambokidis, J. and J. Barlow. 1991. Chlorinated hydrocarbon concentrations and their use for describing population discreteness in Harbor Porpoises from Washington, Oregon, and California. Pp. 101–110 in J. E. Reynolds III and D. K. Odell, eds. Marine mammal strandings in the United States: proceedings of the Second Marine Mammals Stranding Workshop. NOAA Technical Report NMFS 98, Miami, FL.

Calambokidis, J., J. D. Darling, V. Deecke, P. Gearin, M. Gosho, W. Megill, C. M. Tombach, D. Goley, C. Toropova and B. Gisborne. 2002. Abundance, range and movements of a feeding aggregation of gray whales (Eschrichtius robustus) from California to southeastern Alaska in 1998. Journal of Cetacean Research and Management 4:267–276.

Calambokidis, J., S. Osmek and J. L. Laake. 1997. Survey report for the 1997 aerial surverys for harbor porpoise and other marine mammals of Oregon, Washington and British Columbia outside waters. Olympia.

Canning, S. J., M. B. Santos, R. J. Reid, P. G. H. Evans, R. C. Sabin, N. Bailey and G. J. Pierce. 2008. Seasonal distribution of white-beaked dolphins (Lagenorhynchus albirostris) in UK waters with new information on diet and habitat use. Journal of the Marine Biological Association of the United Kingdom 88:1159–1166.

Carretta, J. V, K. A. Forney, M. M. Muto, J. Barlow, J. Baker, B. Hanson and M. S. Lowry. 2005. U.S. Pacific marine mammal stock assessment: 2004. Technical Memorandum. NOAA-TM-NMFS-SWFSC-375. U.S. Department of Commerce, National Oceanic and Atmospheric Administration. Rockville, Maryland, USA.

Charlesworth, B., P. Sniegowski and W. Stephan. 1994. The evolutionary dynamics of repetitive DNA in eukaryotes. Nature 371:215–220.

Chen, L. and G. Yang. 2008. Development of tetranucleotide microsatellite loci for the finless porpoise (Neophocaena phocaenoides). Conservation Genetics 9:1033–1035.

Chivers, S. J., A. E. Dizon, P. J. Gearin and K. M. Robertson. 2002. Small-scale population structure of eastern North Pacific harbour porpoises (Phocoena phocoena) indicated by molecular genetic analyses. Journal of Cetacean Research and Management 4:111–122.

Committee on Taxonomy. 2009. List of marine mammal species and subspecies. Society for Marine Mammalogy. www.marinemammalscience.org Date Accessed [January 21, 2010].

COSEWIC. 2003. COSEWIC assessment and update status report on the harbour porpoise Phocoena phocoena (Pacific Ocean population) in Canada. Committee on the Status of Endangered Wildlife in Canada, Ottawa.

Côté, I. M. and R. Poulin. 1995. Parasitism and group size in social animals: a meta- analysis. Behavioral Ecology 6:159–165.

De Luna, C. J., S. J. Goodman, O. Thatcher, P. D. Jepson, L. Andersen, K. Tolley and A. R. Hoelzel. 2012. Phenotypic and genetic divergence among harbour porpoise populations associated with habitat regions in the North Sea and adjacent seas. Journal of Evolutionary Biology 25:674–681.

DeMaster, D. P. and J. K. Drevenak. 1988. Suvivorship patterns in three species of captive cetaceans. Marine Mammal Science 4:297–311.

Department of Fisheries and Oceans. 2010. Pacific Region integrated fisheries management plan salmon, southern B.C., June 1, 2010 to May 31, 2011. Vancouver.

Dobzhansky, T. 1935. A critique of the species concept in biology. Philosophy of Science 2:344–355.

Dohl, T. P., K. S. Norris and I. Kang. 1974. A porpoise hybrid: Tursiops × Steno. Journal of Mammalogy 55:217–221.

Dugatkin, L. A. 2009. Principles of Animal Behaviour. 2nd ed. W.W. Norton & Company, New York.

Escorza-Treviño, S. and A. E. Dizon. 2000. Phylogeography, intraspecific structure and sex-biased dispersal of Dall’s porpoise, Phocoenoides dalli, revealed by mitochondrial and microsatellite DNA analyses. Molecular Ecology 9:1049–1060.

Excoffier, L., G. Laval and S. Schneider. 2005. Arlequin (version 3.0): an integrated software package for population genetics data analysis. Evolutionary Bioinformatics Online 1:47–50.

Fisheries and Oceans Canada. 2009. Management plan for the pacific Harbour Porpoise (Phocoena phocoena) in Canada. Fisheries and Oceans Canada. Species at Risk Act Management Plan Series. Fisheries and Oceans Canada, Ottawa.

Fontaine, M. C., S. J. E. Baird, S. Piry, N. Ray, K. A. Tolley, S. Duke, A. Birkun, M. Ferreira, T. Jauniaux, A. Llavona, B. Oztürk, A. A Oztürk, V. Ridoux, E. Rogan, M. Sequeira, U. Siebert, G. A. Vikingsson, J.-M. Bouquegneau and J. R. Michaux. 2007. Rise of oceanographic barriers in continuous populations of a cetacean: the genetic structure of harbour porpoises in Old World waters. BMC Biology 5:1–16.

Ford, J. K. B., G. M. Ellis, L. G. Barrett-Lennard, A. B. Morton, R. S. Palm and K. C. Balcomb III. 1998. Dietary specialization in two sympatric populations of killer whales (Orcinus orca) in coastal British Columbia and adjacent waters. Canadian Journal of Zoology 76:1456–1471.

Fraser, F. C. 1940. Three anomalous dolphins from Blacksod Bay, Ireland. Proceedings of the Royal Irish Academy. Section B. Biological, Geological and Chemical Science 45:413–455.

Gannon, D. P., J. E. Craddock and A. J. Read. 1998. Autumn food habits of harbor porpoises, Phocoena phocoena, in the Gulf of Maine. Fishery Bulletin 96:428–437.

Gaskin, D. E., P. W. Arnold and B. A. Blair. 1974. Phocoena phocoena. Mammalian Species 42:1–8.

Geange, S. W., S. Pledger, K. C. Burns and J. S. Shima. 2011. A unified analysis of niche overlap incorporating data of different types. Methods in Ecology and Evolution 2:175–184.

Glover, K. A., N. Kanda, T. Haug, L. A. Pastene, N. Oien, M. Goto, B. B. Seliussen and H. J. Skaug. 2010. Migration of Antarctic minke whales to the Arctic. PloS One 5:e15197.

Gowans, S. and H. Whitehead. 1995. Distribution and habitat partitioning by small odontocetes in the Gully, a submarine canyon on the Scotian Shelf. Canadian Journal of Zoology 73:1599–1608.

Grant, P. R. and B. R. Grant. 1992. Hybridization of bird species. Science 256:193–197.

Gray, A. P. 1954. Mammalian hybrids: a checklist with bibliography. Technical Communication no 10. Commonwealth Bureau of Animal Breeding and Genetics, Edinburgh, Scotland.

Guillot, G., F. Mortier and A. Estoup. 2005. Geneland: a computer package for landscape genetics. Molecular Ecology Notes 5:712–715.

Haldane, J. B. S. 1922. Sex ratio and unisexual sterility in hybrid animals. Journal of Genetics 12:101–109.

Hall, A. 2011. Foraging behaviour and reproductive season habitat selection of Northeast Pacific porpoises. MSc Thesis. University of British Columbia: Canada.

Hall, A., G. Ellis and A. W. Trites. 2002. Harbour Porpoise interactions with the 2001 selective salmon fisheries in southern British Columbia and license holder reported small cetacean by-catch. Selective Salmon Fisheries Science Program Report. Fisheries and Oceans Canada.

Hall, A. M. 2004. Seasonal abundance and prey distribution of Harbour Porpoise (Phocoena phocoena) in Southern Vancouver Island waters. MSc Thesis. University of British Columbia: Canada.

Hardy, O. J. and X. Vekemans. 2002. SPAGeDi: a versitile computer program to analyse spatial genetic structure at the individual or population level. Molecular Ecology Notes 2:618–620.

Heide-Jørgensen, M. P. and R. R. Reeves. 1993. Description of an anomalous monodontid skull from West Greenland: a possible hybrid? Marine Mammal Science 9:258–268.

Herzing, D. L. 1997. The life history of free-ranging atlantic spotted dolphins (Stenella frontalis): age classes, color phases, and female reproduction. Marine Mammal Science 13:576–595.

Herzing, D. L. and C. M. Johnson. 1997. Interspecific interactions between Atlantic Spotted dolphins (Stenella frontalis) and Bottlenose dolphins (Tursiops truncatus) in the Bahamas, 1985–1995. Aquatic Mammals 23:85–99.

Herzing, D. L., K. Moewe and B. J. Brunnick. 2003. Interspecies interactions between Atlantic spotted dolphins, Stenella frontalis and bottlenose dolphins, Tursiops truncatus, on Great Bahama Bank, Bahamas. Aquatic Mammals 29:335–341.

Hirota, T., T. Hirohata, H. Mashima, T. Satoh and Y. Obara. 2004. Population structure of the large Japanese field mouse, Apodemus speciosus (Rodentia: Muridae), in suburban landscape, based on mitochondrial D-loop sequences. Molecular Ecology 13:3275–3282.

Hoelzel, A. R. (ed). 2002. Marine mammal biology: an evolutionary approach. Blackwell Publishing, Malden, MA.

Hoelzel, A. R., M. Dahlheim and S. J. Stern. 1998. Low genetic variation among Killer Whales (Orciuns orca) in the Eastern North Pacific and genetic differentiation between foraging specialists. Journal of Heredity 89:121–128.

Hoelzel, A. R., J. M. Hancock and G. A. Dover. 1991. Evolution of the cetacean mitochondrial D-loop region. Molecular Biology and Evolution 8:475–493.

Hoffman, J. I., C. W. Matson, W. Amos, T. R. Loughlin and J. W. Bickham. 2006. Deep genetic subdivision within a continuously distributed and highly vagile marine mammal, the Steller’s sea lion (Eumetopias jubatus). Molecular Ecology 15:2821– 2832.

Huxel, G. R. 1999. Rapid displacement of native species by invasive species: effects of hybridization. Biological Conservation 89:143–152.

IUCN. 2012. The IUCN Red List of Threatened Species. Version 2012.1. www.iucnredlist.org Date Accessed [September 9, 2012].

Jefferson, T. A. 1988. Phocoenoides dalli. Mammalian Species 319:1–7.

Jefferson, T. A. and B. E. Curry. 1994. A global review of porpoise (Cetacea: Phocoenidae) mortality in gillnets. Biological Conservation 67:167–183.

Jefferson, T. A., P. J. Stacey and R. W. Baird. 1991. A review of Killer Whale interactions with other marine mammals: predation to co-existence. Mammal Review 21:151– 180.

Jones, F. C., C. Brown, J. M. Pemberton and V. A. Braithwaite. 2006. Reproductive isolation in a threespine stickleback hybrid zone. Journal of Evolutionary Biology 19:1531–1544.

Karczmarski, L., M. Thornton and V. G. Cockcroft. 1997. Description of selected behaviours of humpback dolphins Sousa chinensis. Aquatic Mammals 23:127–133.

Kirkwood, J. K., P. M. Bennett, P. D. Jepson, T. Kuiken, V. R. Simpson and J. R. Baker. 1997. Entanglement in fishing gear and other causes of death in cetaceans stranded on the coasts of England and Wales. The Veterinary Record 141:94–98.

Koopman, H. N. and D. E. Gaskin. 1994. Individual and geographical variation in pigmentation patterns of the harbour porpoise, Phocoena phocoena (L.). Canadian Journal of Zoology 72:135–143.

Kumar, S., M. Nei, J. Dudley and K. Tamura. 2008. MEGA: a biologist-centric software for evolutionary analysis of DNA and protein sequences. Briefings in Bioinformatics 9:299–306.

Lockyer, C., M. P. Heide-Jørgensen, J. Jensen, C. C. Kinze and T. Buus Sørensen. 2001. Age, length and reproductive parameters of harbour porpoises Phocoena phocoena (L.) from West Greenland. ICES Journal of Marine Science 58:154–162.

Loehle, C. 1995. Social barriers to pathogen tranmission in wild animal populations. Ecology 76:326–335.

Maines, J. and S. Kestin. 2009. Database: U.S. Marine Mammal Inventory. National Marine Fisheries Service. Date Accessed [August 18, 2012].

Mallet, J. 2007. Hybrid speciation. Nature 446:279–283.

Mann, J. 2006. Establishing trust: socio-sexual behvaiour and the development of male- male bonds among Indian Ocean bottlenose dolphins. Pp. 107–130 in V. Sommer and P. L. Vassey, eds. Homosexual behaviour in animals. Cambridge University Press, New York, NY.

Martien, K. K., R. W. Baird, N. M. Hedrick, A. M. Gorgone, J. L. Thieleking, D. J. McSweeney, K. M. Robertson and D. L. Webster. 2012. Population structure of island-associated dolphins: Evidence from mitochondrial and microsatellite markers for common bottlenose dolphins (Tursiops truncatus) around the main Hawaiian Islands. Marine Mammal Science 28:E208–E232.

McAuliffe Dore, K., T. R. Turner, J. G. Lorenz and J. P. Grobler. 2009. Intergrating information into the analysis of the genetic distribution of South African vervet monkeys. Field Notes: A Journal of Collegiate Anthropology 1:112–127.

McGlashan, D. J. and J. M. Hughes. 2000. Reconciling patterns of genetic variation with stream structure, earth history and biology in the Australian freshwater fish Craterocephalus stercusmuscarum (Atherinidae). Molecular Ecology 9:1737–1751.

McGowen, M. R., M. Spaulding and J. Gatesy. 2009. Divergence date estimation and a comprehensive molecular tree of extant cetaceans. Molecular Phylogenetics and Evolution 53:891–906.

Miyazaki, N., Y. Hirosaki, T. Kinuta and H. Omura. 1992. Osteological Study of a hybrid between Tursiops truncatus and Grampus griseus. Bulletin of the National Science Museum, Tokyo, Series A. 18:79–94.

Morin, P. A., K. E. Chambers, C. Boesch and L. Vigilant. 2001. Quantitative polymerase chain reaction analysis of DNA from noninvasive samples for accurate microsatellite genotyping of wild chimpanzees (Pan troglodytes verus). Molecular Ecology 10:1835–1844.

Nishiwaki, M. and T. Tobayama. 1982. Morphological study on the hybrid between Tursiops and Pseudorca. Scientific Report Whales Research Institute 34:109–121.

Noor, M. A. F. 1999. Reinforcement and other consequences of sympatry. Heredity 83:503–508.

Odell, D. K. and K. M. McClune. 1999. False Killer Whale Pseudorca crassidens (Owen, 1846). Pp. 213–243 in S. H. Ridgway and R. Harrison, eds. Handbook of Marine Mammals: Volume 6 The second book of dolphins and the porpoises. Academic Press Inc., Toronto, ON.

Oftedal, O. T. 1997. Lactation in whales and dolphins: evidence of divergence between baleen- and toothed-species. Journal of Mammary Gland Biology and Neoplasia 2:205–230.

Oksanen, J., F. G. Blanchet, R. Kindt, P. Legendre, P. R. Minchin, R. B. O’Hara, G. L. Simpson, P. Solymos, M. H. H. Stevens and H. Wagner. 2011. vegan: Community Ecology Package. R package version 2.0-2." Режим доступа: http://CRAN. R- project. org/package= vegan (2011).

Orr, H. A. 1995. The population genetics of speciation: the evolution of hybrid incompatibilities. Genetics 139:1805–1813.

Paetkau, D., R. Slade, M. Burden and A. Estoup. 2004. Genetic assignment methods for the direct, real-time estimation of migration rate: a simulation-based exploration of accuracy and power. Molecular Ecology 13:55–65.

Pause, K. C., R. K. Bonde, P. M. McGuire, R. T. Zori and B. A. Gray. 2006. G-banded karyotype and ideogram for the North Atlantic Right Whale (Eubalaena glacialis). Journal of Heredity 97:303–306.

Petit, R. J., A. El Mousadik and O. Pons. 1998. Identifying populations for conservation on the basis of genetic markers. Conservation Biology 12:844–855.

Pfenninger, M., A. Bahl and B. Streit. 1996. Isolation by distance in a population of a small land snail Trochoidea geyeri: evidence from direct and indirect methods. Proceedings of the Royal Society B: Biological Sciences 263:1211–1217.

Piry, S., G. Luikart and J.-M. Cornuet. 1990. BOTTLENECK: A computer program for detecting recent reductions in the effective population size using allele frequency data. Journal of Heredity 90:502–503.

Pourkazemi, M., D. O. F. Skibinski and J. A. Beardmore. 1999. Application of mtDNA d- loop region for the study of Russian sturgeon population structure from Iranian coastline of the Caspian Sea. Journal of Applied Ichthyology 15:23–28.

Pritchard, J. K., M. Stephens and P. Donnelly. 2000. Inference of population structure using multilocus genotype data. Genetics 155:945–959.

Pusey, A. and M. Wolf. 1996. Inbreeding avoidance in animals. Trends in Ecology & Evolution 11:201–206.

Rae, B. B. 1973. Additional notes on the food of the common porpoise (Phocoena phocoena). Journal of Zoology, London 169:127–131.

Rae, B. B. 1965. The food of the Common porpoise (Phocaena phocaena). Journal of Zoology 146:114–122.

Randler, C. 2006. Behavioural and ecological correlates of natural hybridization in birds. Ibis 148:459–467.

Raymond, M. and F. Rousset. 1995. An exact test for population differentiation. Evolution 49:1280–1283.

Read, A. J. and A. A. Hohn. 1995. Life in the fast lane: the life history of harbor porpoises from the Gulf of Maine. Marine Mammal Science 11:423–440.

Read, A. J. and K. A. Tolley. 1997. Postnatal growth and allometry of harbour porpoises from the Bay of Fundy. Canadian Journal of Zoology 75:122–130.

Read, A. J. and A. J. Westgate. 1997. Monitoring the movements of harbour porpoises (Phocoena phocoena) with satellite telemetry. Marine Biology 130:315–322.

Recchia, C. A. and A. J. Read. 1989. Stomach contents of harbour porpoises, Phocoena phocoena (L.), from the Bay of Fundy. Canadian Journal of Zoology 67:2140–2146.

Rechsteiner, E. 2012. Resting metabolism, energetics, and seasonal distribution of Pacific White-Sided Dolphins. MSc Thesis. University of British Columbia:Canada.

Reyes, J. C. 1996. A possible case of hybridism in wild dolphins. Marine Mammal Science 12:301–307.

Rhymer, J. M. and D. Simberloff. 1996. Extinction by hybridization and introgression. Annual Review of Ecology and Systematics 27:83–109.

Rosel, P. E., A. E. Dizon and M. G. Haygood. 1995. Variability of the mitochondrial control region in populations of the harbour porpoise, Phocoena phocoena, on interoceanic and regional scales. Canadian Journal of Fisheries and Aquatic Sciences 52:1210–1219.

Rosel, P. E., A. E. Dizon and J. E. Heyning. 1994. Genetic analysis of sympatric morphotypes of common dolphins (genus Delphinus). Marine Biology 119:159–167.

Rosel, P. E., S. C. France, J. Y. Wang and T. D. Kocher. 1999. Genetic structure of harbour porpoise Phocoena phocoena populations in the northwest Atlantic based on mitochondrial and nuclear markers. Molecular Ecology 8:S41–S54.

Rowe, C. L. 2008. “The calamity of so long life”: life histories, contaminants, and potential emerging threats to long-lived vertebrates. BioScience 58:623–631.

Ryman, N. and S. Palm. 2006. POWSIM: a computer program for assessing statistical power when testing for genetic differentiation. Molecular Ecology 6:600–602.

Sambrook, J., E. F. Fritsch and H. Maniatis. 1989. Molecular cloning: a laboratory manual. 2nd ed. Cold Springs Harbor Laboratory Press, Cold Spring Harbor, NY.

Santos, M. B., G. J. Pierce, J. A. Learmouth, R. J. Reid, H. M. Ross, I. A. P. Patterson, D. G. Reid and D. Beare. 2004. Variability in the diet of Harbor Porpoises (Phocoena phocoena) in Scottish Waters 1992–2003. Marine Mammal Science 20:1–27.

Scheffer, V. B. and J. W. Slipp. 1948. The whales and dolphins of Washington State with a key to the cetaceans of the west coast of North America. American Midland Naturalist 39:257–337.

Schlötterer, C., B. Amos and D. Tautz. 1991. Conservation of polymorphic simple sequence loci in cetacean species. Nature 354:63–65.

Schwartz, M. K., G. Luikart and R. S. Waples. 2007. Genetic monitoring as a promising tool for conservation and management. Trends in Ecology & Evolution 22:25–33.

Scott, E. M., J. Mann, J. J. Watson-Capps, B. L. Sargeant and R. C. Connor. 2005. Aggression in bottlenose dolphins: evidence for sexual coercion, male-male competition and female tolerance through analysis of tooth-rake marks and behaviour. Behaviour 142:21–44.

Scribner, K. T., K. S. Page and M. L. Bartron. 2001. Hybridization in freshwater fishes: a review of case studies and cytonuclear methods of biological inference. Reviews in Fish Biology and Fisheries 10:293–323.

Shahidul Islam, M. and M. Tanaka. 2004. Impacts of pollution on coastal and marine ecosystems including coastal and marine fisheries and approach for management: a review and synthesis. Marine Pollution Bulletin 48:624–649.

Shallenberger, E. W. and I. King. 1977. Dolphin birth at Sea Life Park. Pp. 77–84 in S. M. Ridgeway and K. Bernirschke, eds. Breeding dolphins: present status, suggestions for the future. U.S. Marine Mammal Commission, Washington, DC.

Shane, S. H., R. S. Wells and B. Würsig. 1986. Ecology, behavior and social organization of the Bottlenose Dolphin: a review. Marine Mammal Science 2:34–63.

Sherman, K., M. Sissenwine, V. Christensen, A. Duda, G. Hempel, C. Ibe, S. Levin, D. Lluch-Belda, S. Seitzinger, R. Serra, H.-R. Skjoldal, Q. Tang, J. Thulin, V. Vandeweerd and K. Zwanenburg. 2005. A global movement toward an ecosystem based approach to management of marine resources. Marine Ecology Progress Series 300:275–279.

Shimura, E., K. Numachi, K. Sezaki, Y. Hirosaki, S. Watabe and K. Hashimoto. 1985. Biochemical evidence of hybrid formation between the two species of dolphin, Tursiops truncatus and Grampus griseus. Bulletin of the Japanese Society of Scientific Fisheries 52:725–730.

Silva-Jr, J. M., F. J. L. Silva and I. Sazima. 2005. Two Presumed Interspecific Hybrids in the Genus Stenella (Delphinidae) in the Tropical West Atlantic. Aquatic Mammals 31:468–472.

Slater, G. J., S. A. Price, F. Santini and M. E. Alfaro. 2010. Diversity versus disparity and the radiation of modern cetaceans. Proceedings of the Royal Society B: Biological Sciences 277:3097–3104.

Southern, Š. O., P. J. Southern and A. E. Dizon. 1988. Molecular characterization of a cloned dolphin mitochondrial genome. Journal of Molecular Evolution 28:32–42.

Spilliaert, R., G. Vikingsson, Ú. Árnason, A. Palsdottir, J. Sigurjonsson and A. Arnason. 1991. Species hybridization between a female blue whale (Balaenoptera musculus) and a male fin whale (B. physalus): molecular and morphological documentation. The Journal of Heredity 82:269–274.

Stacey, P. J., D. A. Duffus and R. W. Baird. 1997. A preliminary evaluation of incidental mortality of small cetaceans in coastal fisheries in British Columbia, Canada. Marine Mammal Science 13:321–326.

Stebbins, G. L. 1958. The inviability, weakness, and sterility of interspecific hybrids. Advances in Genetics 9:147–215.

Stroud, R. K. and T. J. Roffe. 1979. Causes of death in marine mammals stranded along the Oregon coast. Journal of Wildlife Diseases 15:91–97.

Sunnucks, P. 2000. Efficient genetic markers for population biology. Trends in Ecology & Evolution 15:199–203.

Sylvestre, J.-P. and S. Tasaka. 1985. On the intergeneric hybrids in cetaceans. Aquatic Mammals 11:101–108.

Taberlet, P., L. P. Waits and G. Luikart. 1999. Noninvasive genetic sampling: look before you leap. Trends in Ecology & Evolution 14:323–327.

Taguchi, M., S. J. Chivers, P. E. Rosel, T. Matsuishi and A. Syuiti. 2010. Mitochondrial DNA phylogeography of the harbour porpoise Phocoena phocoena in the North Pacific. Marine Biology 157:1489–1498.

Tamura, K., D. Peterson, N. Peterson, G. Stecher, M. Nei and S. Kumar. 2011. MEGA5: molecular evolutionary genetics analysis using maximum likelihood, evolutionary distance, and maximum parsimony methods. Molecular Biology and Evolution 28:2731–2739.

Taylor, E. B., P. Tamkee, E. R. Keeley and E. A. Parkinson. 2011. Conservation prioritization in widespread species: the use of genetic and morphological data to assess population distinctiveness in rainbow trout (Oncorhynchus mykiss) from British Columbia, Canada. Evolutionary Applications 4:100–115.

Valsecchi, E. and W. Amos. 1996. Microsatellite markers for the study of cetacean populations. Molecular Ecology 5:151–156.

Van Dyke, J., D. Zaelke and G. Hewison (eds). 1993. Freedom for the seas in the 21st century: ocean governance and environmental harmony. Island Press, Washington, DC.

Van Oosterhout, C., W. F. Hutchinson, D. P. M. Wills and P. Shipley. 2004. MICRO- CHECKER: Software for identifying and correcting genotyping errors in microsatellite data. Molecular Ecology Notes 4:535–538.

Vanhooydonck, B., R. Van Damme and P. Aerts. 2000. Ecomorphological correlates of habitat partitioning in Corsican lacertid lizards. Functional Ecology 14:358–368.

Vasey, P. L. 1995. Homosexual behavior in primates: a review of evidence and theory. International Journal of Primatology 16:173–204.

Walker, W. A., M. B. Hanson, R. W. Baird and T. J. Guenther. 1998. Food habits of the harbor porpoise, Phocoena phocoena, and Dall’s porpoise, Phocoenoides dalli, in the inland waters of British Columbia and Washington. Pp. 63–75 in Marine Mammal Protection Act and Endangered Species Protection Act Implementation Program 1997. AFSC Processed Report 98-10. National Marine Fisheries Service, Seattle, WA.

Walton, M. J. 1997. Population structure of harbour porpoises Phocoena phocoena in the seas around the UK and adjacent waters. Proceedings of the Royal Society London London B 264:89–94.

Wang, J. 2001. A pseudo-likelihood method for estimating effective population size from temporally spaced samples. Genetical Research 78:243–257.

Wang, J. Y. and P. Berggren. 1997. Mitochondrial DNA analysis of harbour porpoises (Phocoena phocoena) in the Baltic Sea, the Kattegat-Skagerrak Seas and off the west coast of Norway. Marine Biology 127:531–537.

Wang, J. Y., D. E. Gaskin and B. N. White. 1996. Mitochondrial DNA analysis of the harbour porpoise, (Phocoena phocoena), subpopulations in North American waters. Canadian Journal of Fisheries and Aquatic Sciences 53:1632–1645.

Waples, R. S. and O. Gaggiotti. 2006. What is a population? An empirical evaluation of some genetic methods for identifying the number of gene pools and their degree of connectivity. Molecular Ecology 15:1419–1439.

Whitney, K. D., J. R. Ahern, L. G. Campbell, L. P. Albert and M. S. King. 2010. Patterns of hybridization in plants. Perspectives in Plant Ecology, Evolution and Systematics 12:175–182.

Wiemann, A., L. W. Andersen, P. Berggren, U. Siebert, H. Benke, J. Teilmann, C. Lockyer, I. Pawliczka, K. Skóra, A. Roos, T. Lyrholm, K. B. Paulus, V. Ketmaier and R. Tiedemann. 2010. Mitochondrial control region and microsatellite analyses on harbour porpoise (Phocoena phocoena) unravel population differentiation in the Baltic Sea and adjacent waters. Conservation Genetics 11:195–211.

Williams, R., A. Hall and A. Winship. 2008. Potential limits to anthropogenic mortality of small cetaceans in coastal waters of British Columbia. Canadian Journal of Fisheries and Aquatic Sciences 65:1867–1878.

Willis, P. M., B. J. Crespi, L. M. Dill, R. W. Baird and M. B. Hanson. 2004. Natural hybridization between Dall’s porpoises (Phocoenoides dalli) and harbour porpoises (Phocoena phocoena). Canadian Journal of Zoology 82:828–834.

Wright, S. 1943. Isolation by distance. Genetics 28:114–138.

Wright, S. 1951. The genetical structure of populations. Annals of Eugenics 15:323–354.

Yazdi, P. 2002. A possible hybrid between the dusky dolphin (Lagenorhynchus obscurus) and the southern right whale dolphin (Lissodelphis peronii). Aquatic Mammals 28:211–217.

Zane, L., L. Bargelloni and T. Patarnello. 2002. Strategies for microsatellite isolation: a review. Molecular Ecology 11:1–16.

Zornetzer, H. R. and D. A. Duffield. 2003. Captive-born bottlenose dolphin × common dolphin (Tursiops truncatus × Delphinus capensis) intergeneric hybrids. Canadian Journal of Zoology 81:1755–1762.

Appendices

A.1 Appendix S1 – Chapter 2

Table S1.1 Case numbers and additional organizations associated with each sample Species DFO AHC LBL Other Case Number Other Org. Dall's Porpoise AHC 00-3075 Dall's Porpoise AHC 02-3166 Dall's Porpoise AHC 04-1780 04-SJ004 Dall's Porpoise AHC 11-3920 Dall's Porpoise DFO 1075 AHC 03-2768 LBL 03-14 AH 03OBDM06 Dall's Porpoise DFO 2362 AHC 06-0706 Dall's Porpoise DFO 2374 AHC 06-0916 Dall's Porpoise DFO 4078 Dall's Porpoise DFO 4110 Dall's Porpoise DFO 4287 AHC 02-2110 LBL 02-18 Dall's Porpoise DFO 4631 LBL 02-19 Dall's Porpoise DFO 4838 AHC 09-01909 Dall's Porpoise DFO 5563 Dall's Porpoise LBL 00-06 Ron Lewis 98-403 From AHC Dall's Porpoise LBL 00-07 Ron Lewis 98-405 From AHC Dall's Porpoise LBL 00-08 Ron Lewis 98-405, From AHC 97-05 Dall's Porpoise LBL 00-09 Ron Lewis 98-405, From AHC 97-10 Dall's Porpoise LBL 00-10 Ron Lewis 98-421 From AHC Dall's Porpoise LBL 00-11 Ron Lewis 98-422 From AHC Dall's Porpoise LBL 00-12 Ron Lewis 98-442, SWDP From AHC 97-11 Dall's Porpoise DFO 3763 LBL 00-13 Ron Lewis 98-1468 From AHC Dall's Porpoise LBL 00-14 Ron Lewis 98-1473 From AHC Dall's Porpoise DFO 4287 AHC 02-02110 LBL 02-18 Dall's Porpoise AHC 02-00609 LBL 02-19 Dall's Porpoise AHC 02-00369 LBL 02-20 Dall's Porpoise AHC 02-00248 LBL 02-21 Dall's Porpoise AHC 03-02770 LBL 03-17 030WCPPF10 Dall's Porpoise DFO 1812 AHC 04-00362 LBL 04-18 Dall's Porpoise DFO 1911 AHC 04-01444 LBL 04-22 Dall's Porpoise DFO 3056 AHC 07-3966 LBL 07-04 FOS 3056 Dall's Porpoise DFO 2910 AHC 07-2879 LBL 07-10 CR

Species DFO AHC LBL Other Case Number Other Org. Dall's Porpoise DFO 3433 AHC 08-3876 LBL 08-21 FOS 3433 Dall's Porpoise DFO 5289 AHC 09-4544 LBL 09-08 FOS 4289 Dall's Porpoise DFO 3083 AHC 09-2920 LBL 09-10 FOS 3083 Dall's Porpoise LBL 96-01 G. Ellis Dall's Porpoise LBL 98-17A Dall's Porpoise LBL 99-02 SIRS 99-07 Dall's Porpoise PW 0619-1(97) P. Willis Dall's Porpoise SWDP 95-17 P. Willis Dall's Porpoise SWDP 97-05 P. Willis Dall's Porpoise SWDP 97-07 P. Willis Dall's Porpoise SWDP 97-10 P. Willis Dall's Porpoise SWDP 97-14 P. Willis Dall's Porpoise SWDP 97-16 P. Willis Harbour Porpoise AHC 00-3866 Harbour Porpoise AHC 00-3979 Harbour Porpoise AHC 02-3164 Harbour Porpoise AHC 02-3171 LBL 02-28 Harbour Porpoise AHC 03-2356 2002SJ-006 Harbour Porpoise AHC 03-3096 Harbour Porpoise AHC 05-03200 Harbour Porpoise AHC 05-03901 A-Ppunk-03-04, 1-29045B Harbour Porpoise AHC 06-02388 060412-JST-PHPH Harbour Porpoise AHC 06-03282 Harbour Porpoise AHC 06-1962 06PpVicF-04 Harbour Porpoise AHC 06-3276 CRC-733 Harbour Porpoise AHC 07-00626 Harbour Porpoise AHC 07-2921 06Pp21NovW1-06 Vancouver Aquarium Harbour Porpoise AHC 08-1592 Harbour Porpoise AHC 09-01252 09-0SJ001 Harbour Porpoise DFO 5026 AHC 09-02921 Harbour Porpoise AHC 09-03252 CRC-966 Harbour Porpoise AHC 09-3238 CRC-929 Harbour Porpoise AHC 10-01596 CRC-1031 Harbour Porpoise AHC 10-0515 09Pp01SepWI-02 CPSMMSN Harbour Porpoise AHC 10-0516 09Pp02SeptWH-10 CPSMMSN Harbour Porpoise AHC 10-1618 Harbour Porpoise AHC 10-1907 LBL 09-07 Harbour Porpoise AHC 10-4951 DFO 5733 Harbour Porpoise AHC 10-4952 DFO 5737

Species DFO AHC LBL Other Case Number Other Org. Harbour Porpoise AHC 10-4953 Harbour Porpoise AHC 10-514 09Pp07AugSK-01 Harbour Porpoise DFO 5941 AHC 11-150 Harbour Porpoise DFO 6507 AHC 11-1945 Harbour Porpoise DFO 6509 AHC 11-1946 Harbour Porpoise DFO 6449 AHC 11-1987 Harbour Porpoise DFO 6448 AHC 11-2196 Harbour Porpoise DFO 6599 AHC 11-2197 Harbour Porpoise DFO 6600 AHC 11-2198 Harbour Porpoise DFO 6610 AHC 11-2199 Harbour Porpoise DFO 6611 AHC 11-2200 Harbour Porpoise DFO 6620 AHC 11-2201 Harbour Porpoise AHC 11-2487 11Pp01JanWI-01 Harbour Porpoise DFO 6627 AHC 11-3143 Harbour Porpoise DFO 6779 AHC 11-3921 Harbour Porpoise AHC 11-721 Harbour Porpoise DFO 6285 AHC 11-930 Harbour Porpoise DFO 6928 AHC 12-1018 Harbour Porpoise DFO 6840 AHC 12-1022 Harbour Porpoise DFO 9373 AHC 12-1023 Harbour Porpoise DFO 6290 AHC 11-1641 Harbour Porpoise DFO 6296 AHC 11-1642 Harbour Porpoise DFO 6298 AHC 11-1643 Harbour Porpoise Birch Bay 2006 B. Hanson Harbour Porpoise CRC-711 Harbour Porpoise DFO 1071 AHC 03-00171 LBL 03-20 Harbour Porpoise DFO 1081 AHC 03-02186 LBL 03-19 Harbour Porpoise DFO 1833 AHC 04-00682 LBL 04-16 Harbour Porpoise DFO 2227 AHC 05-01619 Harbour Porpoise DFO 2246 AHC 05-002074 AH 05PpVICM20 Harbour Porpoise DFO 2250 AHC 05-02076 AH 05PpSSF23 Harbour Porpoise DFO 2400 AHC 06-01964 Harbour Porpoise DFO 2787 AHC 07-01692 Harbour Porpoise DFO 2788 AHC 07-1621 Harbour Porpoise DFO 2820 AHC 07-01685 Harbour Porpoise DFO 3241 AHC 08-02855 LBL 08-25 Harbour Porpoise DFO 3340 AHC 08-02977 Harbour Porpoise DFO 3399 AHC 08-3529 LBL 08-22 Harbour Porpoise DFO 3406 AHC 08-03815 AH 08PpEsqM06 Harbour Porpoise DFO 3410 AHC 08-03531 LBL 08-18

Species DFO AHC LBL Other Case Number Other Org. Harbour Porpoise DFO 3519 AHC 02-02112 LBL 02-16 AH 02PpM02 Harbour Porpoise DFO 3539 SWDP 93-46, HP93-15 SWDP Harbour Porpoise DFO AHC 02-02115 LBL 02-12 AH 02PhF08 3599A? Harbour Porpoise DFO 3599B AHC 02-02115 LBL 02-33 AH 02PpF08 Harbour Porpoise DFO 3639 SWDP 94-08, HP94-01A SWDP Harbour Porpoise DFO 3691 AHC 01-03253 Harbour Porpoise DFO 3742 SWDP 93-54, HP93-14 SWDP Harbour Porpoise DFO 3819 SWDP 93-06, HP93-02 R. Baird Harbour Porpoise DFO 3860 SWDP 93-43, HP93-10 SWDP Harbour Porpoise DFO 3897 AHC 190 LBL 98-08 HP98-02B Harbour Porpoise DFO 3903 AHC 01-02128 Harbour Porpoise DFO 3916A AHC 01-03740 M0697 Harbour Porpoise DFO 3916B AHC 01-03740 M0697 Harbour Porpoise DFO 3919 AHC 01-04519 Harbour Porpoise DFO 3964 AHC 01-03742 M753-2 Harbour Porpoise DFO 3974 SWDP 94-09, HP94-02A SWDP Harbour Porpoise DFO 3993 AHC 01-04520 AH 01JdFM20 Harbour Porpoise DFO 4007 SWDP 94-23, HP94-04 SWDP Harbour Porpoise DFO 4024 SWDP 93-52, HP93-13 SWDP Harbour Porpoise DFO 4064 SWDP 94-41, HP94-11 SWDP Harbour Porpoise DFO 4065 AHC 98-00445 SWDP 94-43, HP94-12 SWDP Harbour Porpoise DFO 4098 SWDP 93-24, HP93-08 R. Baird Harbour Porpoise DFO 4104 AHC 02-02685 LBL 02-31 Harbour Porpoise DFO 4126 AHC 03-02766 LBL 03-18 379, AH 01HSPP002 Harbour Porpoise DFO 4239 AHC 02-03292 LBL 02-09 Harbour Porpoise DFO 4240 AHC 02-03302 LBL 02-14 Harbour Porpoise DFO 4242 AHC 02-03634 LBL 02-13 Harbour Porpoise DFO 4273 SWDP 92-50, HP92-01A, DFO Pp92-1 By-catch Harbour Porpoise DFO 4298 SWDP 93-20, HP93-07A R. Baird Harbour Porpoise DFO 4327 SWDP 94-15, HP94-03 SWDP Harbour Porpoise DFO 4345 SWDP 94-31, HP94-06 SWDP Harbour Porpoise DFO 4346 SWDP 94-32, HP94-07 SWDP Harbour Porpoise DFO 4448 SWDP 93-11, HP93-05 R. Baird Harbour Porpoise DFO 4494 SWDP 93-13, HP93-06 R. Baird Harbour Porpoise DFO 4497 SWDP 94-39, HP94-09 SWDP Harbour Porpoise DFO 4581 AHC 01-03741 Harbour Porpoise DFO 4609 SWDP 93-03, HP93-01 R. Baird Harbour Porpoise DFO 4635 AHC 01-02007 Harbour Porpoise DFO 4644 AHC 03-01246 LBL 03-21

Species DFO AHC LBL Other Case Number Other Org. Harbour Porpoise DFO 4661 AHC 01-3813 Harbour Porpoise DFO 4697 SWDP 95-01, HP95-01 SWDP Harbour Porpoise DFO 4723 SWDP 93-41, HP93-11 SWDP Harbour Porpoise DFO 4788 SWDP 93-07, HP93-03 R. Baird Harbour Porpoise DFO 4819 Harbour Porpoise DFO 5206 AHC 09-03814 RR 09-0254 Harbour Porpoise DFO 5556 Harbour Porpoise DFO 5566 AHC 04-01950 Harbour Porpoise DFO 5649 AHC 10-01948 LBL 10-12 AH 16PpRRF10, RR 10- 0084 Harbour Porpoise DFO 5651 AHC 10-01946 LBL 10-10 Harbour Porpoise DFO 5656 AHC 10-01952 LBL 10-16 AH 20PpVicF10 Harbour Porpoise DFO 5723 Harbour Porpoise DFO 5733 AHC 10-4591 Harbour Porpoise DFO 5738 Harbour Porpoise DFO 6292 AHC 02-01779 SJ057 Harbour Porpoise DFO 6443 SWDP94-?, HP94-10, SWDP 172 Harbour Porpoise DFO 6445 B. Hanson Harbour Porpoise JackVA Vancouver Aquarium Harbour Porpoise LBL 00-02 SIRS SIRS Harbour Porpoise LBL 00-16 Ron Lewis 98-406 T. Guenther Harbour Porpoise LBL 00-17 Ron Lewis 98-406, SWDP T. Guenther 97-08 Harbour Porpoise LBL 00-18 Ron Lewis 98-408 T. Guenther Harbour Porpoise DFO 4656 LBL 00-19 Ron Lewis 98-409, SWDP From AHC 94-06 Harbour Porpoise LBL 00-20 Ron Lewis 98-410 T. Guenther Harbour Porpoise LBL 00-21 Ron Lewis 98-411 From AHC Harbour Porpoise LBL 00-22 Ron Lewis 98-412 From AHC Harbour Porpoise LBL 00-23 Ron Lewis 98-413 T. Guenther Harbour Porpoise LBL 00-24 Ron Lewis 98-414 From AHC Harbour Porpoise LBL 00-25 Ron Lewis 98-415 T. Guenther Harbour Porpoise LBL 00-26 Ron Lewis 98-416 T. Guenther Harbour Porpoise LBL 00-27 Ron Lewis 98-420 T. Guenther Harbour Porpoise LBL 00-28 Ron Lewis 98-432 T. Guenther Harbour Porpoise LBL 00-30 Ron Lewis 98-441 T. Guenther Harbour Porpoise LBL 00-31 Ron Lewis 98-443, SWDP From AHC 97-13 Harbour Porpoise DFO 4565 LBL 00-33 Ron Lewis 98-417 SWDP From AHC 95-27

Species DFO AHC LBL Other Case Number Other Org. Harbour Porpoise LBL 02-09 SIRS Harbour Porpoise AHC 02-03278 LBL 02-22 Harbour Porpoise AHC 02-02896 LBL 02-29 Harbour Porpoise AHC 03-02767 LBL 03-15 VICPPM05 Harbour Porpoise AHC 05-01611 LBL 05-30 05PpVicU03 Harbour Porpoise DFO 3080 AHC 07-4080 LBL 07-07 FOS 3080 Harbour Porpoise DFO 2926 AHC 07-3212 LBL 07-08 FOS 2926 Vancouver Aquarium Harbour Porpoise DFO 2929 AHC 07-3230 LBL 07-12 FOS 2929 Harbour Porpoise DFO 3257 AHC 08-3530 LBL 08-19 FOS 3257 Harbour Porpoise DFO 3402 AHC 08-3528 LBL 08-23 Harbour Porpoise DFO 3494 AHC 09-1908 LBL 09-09 FOS 3494 Harbour Porpoise DFO 5192 AHC 09-4545 LBL 09-11 FOS 5088 Harbour Porpoise DFO 5360 AHC 10-1850 LBL 10-04 FOS 5360 Harbour Porpoise DFO 5507 AHC 10-1851 LBL 10-05 Harbour Porpoise DFO 3184 AHC 10-1852 LBL 10-06 John Ford Harbour Porpoise DFO 3185 AHC 10-1853 LBL 10-07 FOS 3185 Harbour Porpoise DFO 5633 AHC 10-1855 LBL 10-08 Harbour Porpoise DFO 5647 AHC 10-1945 LBL 10-09 Harbour Porpoise DFO 5648 AHC 10-1947 LBL 10-11 15PpRRM10 Harbour Porpoise DFO 5650 AHC 10-1949 LBL 10-13 17PpMetM10 Harbour Porpoise DFO 5652 AHC 10-1950 LBL 10-14 18PpMetM10 Harbour Porpoise DFO 5653 AHC 10-1951 LBL 10-15 19PpMetM10 Harbour Porpoise DFO 5655 AHC 10-1953 LBL 10-17 21PpSKF10 Harbour Porpoise DFO 5713 AHC 10-3026 LBL 10-18 Harbour Porpoise DFO 5726 AHC 10-3823 LBL 10-19 Harbour Porpoise DFO 5719 AHC 10-3822 LBL 10-20 Harbour Porpoise DFO 5657 AHC 10-3819 LBL 10-21 Harbour Porpoise DFO 5722 AHC 10-3820 LBL 10-22 Harbour Porpoise DFO 5731 AHC 10-3824 LBL 10-23 Harbour Porpoise DFO 5712 AHC 10-3025 LBL 10-24 Harbour Porpoise DFO 5506 AHC 10-3818 LBL 10-25 Harbour Porpoise DFO 5720 AHC 10-3821 LBL 10-26 Harbour Porpoise DFO 5730 AHC 10-3567 LBL 10-29 Harbour Porpoise LBL 95-01 J. Ford Harbour Porpoise LBL 98-08 G. Ellis Harbour Porpoise LBL 98-09 Nitnat Fishery Harbour Porpoise LBL 98-10 Harbour Porpoise LBL 98-12 Harbour Porpoise LBL 98-13

Species DFO AHC LBL Other Case Number Other Org. Harbour Porpoise LBL 98-14 Nitnat Fishery Harbour Porpoise LBL 98-15 Area 12-1 test fishery Harbour Porpoise LBL 98-16 Harbour Porpoise LBL 98-17B Ron Lewis #97/003539 Harbour Porpoise LBL 98-19 SIRS Harbour Porpoise SWDP 95-10 P. Willis Harbour Porpoise SWDP 95-13 P. Willis Harbour Porpoise SWDP 95-27 P. Willis Harbour Porpoise SWDP 96-11 P. Willis Harbour Porpoise SWDP 97-09 P. Willis Hybrid PW 51698 P. Willis Hybrid PW 52597 P. Willis Hybrid PW 61397 P. Willis Hybrid PW 628 P. Willis Hybrid PW 82897 P. Willis Hybrid PW 98-01 P. Willis Hybrid PW 98-03 P. Willis Hybrid PW 98-04 P. Willis Hybrid PW 98-05 P. Willis Hybrid AHC 08-120 LBL 07-11 WDFW 1207-04 Porpoise LBL 00-39 Ron Lewis 98-444, SWDP T. Guenther 97-16 Porpoise AHC 02-02114 LBL 02-11 AH 02PpM06 Porpoise AHC 02-02113 LBL 02-15 AH 02PpF05 Porpoise DFO AHC 07-3624 LBL 07-09 Vancouver 3027(8) Aquarium

Abbreviated Organizations Listed Above: AH – Anna Hall AHC – Animal Health Centre CPSMMSN – Central Puget Sound Marine Mammal Stranding Network CR – Cetus Research and Conservation Society CRC – Cascadia Research Collective DFO – Department of Fisheries and Oceans (also includes BC Marine Mammal Response Network) FOS – Fisheries and Oceans Service (now DFO) LBL – Lance Barrett-Lennard SIRS – Strawberry Isle Research Society SJ – San Juan County Marine Mammal Stranding Network SWDP – Stranded Whale and Dolphin Program WDFW – Washington Department of Fish and Wildlife Additional information on individual samples can be obtained through one of the related organizations

Table S1.2 Nucleotide and haplotypic diversity of harbour porpoise using a 99% threshold for pure species. Nucleotide Haplotypic Sample Size (n) Diversity Diversity Three sampling groups Northern Waters 0.024254±0.016604 1.000±0.1768 4 Outer Waters 0.027455±0. 014783 1.000±0.0302 13 Inner Waters 0.026988±0.013486 0.9965±0.0022 104 All samples Single Population 0.033900±0.016718 0.9977±0.0015 138 Two sampling groups Outside/North 0.026208±0.013845 1.000±0.0202 17 Inside Waters 0.026988±0.013486 0.9965±0.0022 104

Table S1.3 Results from analysis of molecular variance apportioning variation in mtDNA d-loop sequences among and within a priori populations of harbour porpoise using a 99% threshold for pure species.

% variation % variation ФST P among within populations populations Three sampling -1.82 101.82 -0.01818 0.82±0.011 groups Two sampling -0.99 100.99 -0.00991 0.75±0.013 groups

Table S1.4 Pairwise ФST (lower diagonal) between two a priori populations based on variation in mtDNA d-loop sequences among and within a priori populations of harbour porpoise using a 99% threshold for pure species. The probabilities that the reported ФST values are significantly different from 0 are indicated in the upper diagonal. Inside Outside/North Inside - 0.76±0.012 Outside/North -0.0099 -

Table S1.5 Variation in microsatellite loci of harbour porpoise using a 99% threshold for pure species.

Loci Number Expected Observed Range in of Alleles Heterozygosity Heterozygosity Allele Size (bp) (He) (Ho) Np404 6 0.584 0.614 134-158 Np407 1 0.000 0.000 186 Np409 3 0.481 0.487 221-229 Np417 12 0.799 0.741 128-176 Np426 6 0.346 0.392 98-116 Np427 7 0.686 0.666 150-276 Np428 8 0.786 0.750 110-142 Np430 3 0.085 0.094 144-168

Table S1.6 Results from analysis of molecular variance apportioning variation in allele frequencies among and within a priori populations of harbour porpoise assayed at eight microsatellite DNA loci using a 99% threshold for pure species.

% variation % variation FST P among within populations populations Two sampling groups 0.25 99.75 0.00247 0.25±0.015

Table S1.7 Pairwise FST (lower diagonal) between two a priori populations based on variation in allele frequencies among and within a priori populations of harbour porpoise assayed at eight microsatellite DNA loci using a 99% threshold for pure species. The probabilities that the reported FST values are significantly different from 0 are indicated in the upper diagonal.

Inside Outside/North Inside - 0.35±0.016 Outside/North 0.00078 -

Table S1.8 Posterior probability 10 runs of Geneland each converging on a single population of harbour porpoises using a 99% threshold for pure species. Number of Clusters Posterior Probability Run1 1 -4587.705 Run2 1 -4582.982 Run3 1 -4595.395 Run4 1 -4591.118 Run5 1 -4565.540 Run6 1 -4578.210 Run7 1 -4526.933 Run8 1 -4608.563 Run9 1 -4574.873 Run10 1 -4608.977

-2600

-2700

-2800

-2900 Mean LnP(K) Mean

-3000

-3100

2 4 6 8 10

K Figure S1.9 Posterior probability of population membership from STRUCTURE for 1 to 10 putative populations (K). Each value is the mean of 20 STRUCTURE simulations.

Table S1.10 Coordinate data used in Geneland Sample Latitude Longitude Sample Latitude Longitude Sample Latitude Longitude AHC 03-2356 48.454 -122.937 DFO 3539 49.008 -123.093 LBL 00-30 49.647 -126.848 AHC 06-02388 48.071 -123.027 DFO 3599B 48.426 -123.382 LBL 00-31 48.450 -122.963 AHC 06-1962 48.418 -123.415 DFO 3639 48.410 -123.326 LBL 00-33 48.419 -123.387 AHC 06-3276 46.928 -124.175 DFO 3742 48.798 -123.202 LBL 02-11 48.373 -123.715 AHC 07-2921 47.972 -122.478 DFO 3819 48.417 -123.370 LBL 02-15 48.416 -123.398 AHC 09-01252 48.454 -123.012 DFO 3897 53.654 -131.905 LBL 03-15 48.403 -123.348 AHC 09-02921 49.848 -124.534 DFO 3919 48.405 -123.355 LBL 05-30 48.420 -123.421 AHC 09-3238 47.241 -124.220 DFO 3964 48.421 -123.396 LBL 07-07 50.286 -125.350 AHC 10-01596 47.244 -124.219 DFO 3974 49.920 -125.184 LBL 07-08 49.254 -124.072 AHC 10-0514 48.318 -122.375 DFO 3993 54.138 -130.279 LBL 07-09 50.590 -127.101 AHC 10-0515 48.198 -122.354 DFO 4007 48.645 -123.803 LBL 07-12 49.021 -122.806 AHC 10-0516 48.744 -122.720 DFO 4024 49.019 -122.808 LBL 08-19 48.333 -123.547 AHC 10-1618 50.127 -125.360 DFO 4064 48.406 -123.353 LBL 08-23 48.254 -123.393 AHC 10-1907 47.607 -122.350 DFO 4065 49.804 -124.527 LBL 09-11 48.746 -123.264 AHC 10-4951 48.751 -123.413 DFO 4098 48.417 -123.317 LBL 10-04 48.798 -123.243 AHC 11-150 49.580 -124.797 DFO 4104 48.407 -123.345 LBL 10-05 48.428 -123.394 AHC 11-1945 48.412 -123.381 DFO 4126 48.417 -123.400 LBL 10-08 48.407 -123.368 AHC 11-1946 48.406 -123.362 DFO 4239 49.262 -126.230 LBL 10-09 48.405 -123.359 AHC 11-1987 48.756 -123.445 DFO 4240 49.198 -123.963 LBL 10-13 48.300 -123.534 AHC 11-2196 48.418 -123.468 DFO 4242 49.671 -124.934 LBL 10-14 48.415 -123.409 AHC 11-2197 48.431 -123.447 DFO 4273 49.179 -122.554 LBL 10-15 48.416 -123.406 AHC 11-2198 48.432 -123.429 DFO 4298 48.330 -123.617 LBL 10-17 48.373 -123.715 AHC 11-2199 48.407 -123.339 DFO 4327 48.369 -123.712 LBL 10-18 49.129 -125.904 AHC 11-2200 48.411 -123.313 DFO 4345 49.048 -125.726 LBL 10-19 49.155 -125.909 AHC 11-2201 48.425 -123.430 DFO 4346 53.313 -132.789 LBL 10-20 49.155 -125.905 AHC 11-2487 47.966 -122.476 DFO 4448 48.351 -123.531 LBL 10-21 49.020 -122.807 AHC 11-3143 48.306 -123.543 DFO 4494 48.350 -123.530 LBL 10-22 48.554 -124.420 AHC 11-3921 49.360 -124.442 DFO 4497 50.027 -125.239 LBL 10-23 48.406 -123.365 AHC 11-930 49.021 -123.100 DFO 4635 48.408 -123.336 LBL 10-24 47.651 -122.295 AHC 12-1018 49.169 -123.935 DFO 4697 48.428 -123.473 LBL 10-25 48.408 -123.372 AHC 12-1022 48.422 -123.421 DFO 4723 49.662 -124.928 LBL 10-29 49.017 -122.794 AHC 12-1023 49.008 -123.119 DFO 4788 48.417 -123.300 LBL 98-08 53.696 -131.871 AHC 2011-1641 49.357 -124.437 DFO 4819 50.779 -126.700 LBL 98-09 48.668 -124.862 AHC 2011-1642 48.378 -123.520 DFO 4838 48.408 -123.341 LBL 98-14 48.666 -124.854 AHC 2011-1643 48.408 -123.367 DFO 5206 49.276 -123.170 LBL 98-19 49.268 -126.142 Birch Bay 2006 48.915 -122.758 DFO 5566 48.299 -123.533 SWDP 96-11 48.357 -123.822 DFO 1071 49.286 -123.216 DFO 5649 48.410 -123.381 SWDP 97-09 48.352 -123.544 DFO 1081 49.273 -123.269 DFO 5651 48.811 -123.602 DFO 1833 49.247 -123.262 DFO 5738 50.029 -125.243 DFO 2227 48.414 -123.388 DFO 6445 48.235 -123.567 DFO 2246 48.419 -123.389 JackVanAqua 49.376 -123.274 DFO 2250 48.826 -123.441 LBL 00-02 49.069 -125.755 DFO 2400 48.417 -123.392 LBL 00-16 48.410 -123.341 DFO 2787 50.683 -126.695 LBL 00-17 48.410 -123.340 DFO 2788 50.684 -126.691 LBL 00-18 48.457 -123.293 DFO 2820 49.152 -125.914 LBL 00-20 48.370 -123.717 DFO 3340 49.229 -123.243 LBL 00-22 48.419 -123.419 DFO 3399 49.293 -126.037 LBL 00-23 49.069 -125.755 DFO 3406 48.413 -123.384 LBL 00-25 48.437 -123.291 DFO 3410 48.409 -123.378 LBL 00-26 48.384 -123.515 DFO 3519 48.424 -123.383 LBL 00-27 49.509 -124.245

Table S1.9 Locations of 112 mtDNA Haplotype Northern Outside Waters Inside (Genbank Waters (14) Waters haplotypes resolved by sequencing 545 Accession) (5) (109) base pairs of d-loop from 134 harbour JX475419 0 0 1 porpoises. The localities “inside”, JX475296 0 0 1 “outside” and “northern” water are JX475301 0 0 1 JX475302 0 0 1 shown in Fig. 2.1. JX475307 0 0 1 Haplotype Northern Outside Waters Inside (Genbank Waters (14) Waters JX475310 0 0 1 Accession) (5) (109) JX475312 0 0 1 JX475334 1 0 0 JX475314 0 0 2 JX475335 1 0 0 JX475315 0 0 1 JX475353 1 0 0 JX475322 0 0 1 JX475380 1 0 0 JX475325 0 0 1 JX475407 1 0 0 JX475326 0 0 1 JX475304 0 1 0 JX475327 0 0 1 JX475306 0 1 0 JX475333 0 0 1 JX475340 0 1 0 JX475336 0 0 1 JX475355 0 1 0 JX475344 0 0 1 JX475328 0 1 3 JX475361 0 0 1 JX475362 0 1 0 JX475402 0 0 1 JX475373 0 1 0 JX475410 0 0 1 JX475294 0 1 4 JX475414 0 0 1 JX475376 0 1 0 JX475348 0 0 2 JX475383 0 1 0 JX475298 0 0 1 JX475387 0 1 0 JX475299 0 0 1 JX475406 0 1 0 JX475300 0 0 3 JX475411 0 1 0 JX475303 0 0 2 JX475413 0 1 0 JX475308 0 0 1 JX475290 0 0 1 JX475309 0 0 1 JX475295 0 0 1 JX475311 0 0 1 JX475305 0 0 1 JX475316 0 0 1 JX475313 0 0 1 JX475321 0 0 1 JX475317 0 0 1 JX475331 0 0 1 JX475323 0 0 3 JX475339 0 0 1 JX475319 0 0 1 JX475346 0 0 1 JX475332 0 0 1 JX475350 0 0 1 JX475338 0 0 1 JX475352 0 0 1 JX475343 0 0 1 JX475354 0 0 1 JX475357 0 0 1 JX475356 0 0 1 JX475384 0 0 1 JX475359 0 0 1 JX475388 0 0 1 JX475369 0 0 1 JX475392 0 0 1 JX475370 0 0 1 JX475405 0 0 1 JX475372 0 0 1 JX475401 0 0 2 JX475374 0 0 1 JX475417 0 0 1 JX475377 0 0 1

Haplotype Northern Outside Waters Inside (Genbank Waters (14) Waters Accession) (5) (109) JX475378 0 0 1 JX475379 0 0 1 JX475395 0 0 1 JX475396 0 0 1 JX475404 0 0 1 JX475408 0 0 1 JX475409 0 0 1 JX475412 0 0 1 JX475418 0 0 1 JX475291 0 0 1 JX475318 0 0 1 JX475347 0 0 1 JX475349 0 0 1 JX475351 0 0 1 JX475360 0 0 1 JX475363 0 0 1 JX475371 0 0 1 JX475375 0 0 1 JX475391 0 0 1 JX475403 0 0 1 JX475415 0 0 1 JX475416 0 0 1 JX475337 0 0 1 JX475342 0 0 1 JX475394 0 0 1 JX475400 0 0 2 JX475397 0 0 1 JX475398 0 0 1 JX475399 0 0 1 JX475324 0 0 1 JX475292 0 0 1 JX475293 0 0 1 JX475320 0 0 1

A.2 Appendix S2 – Chapter 3

Table S2.1 Values for key morphological, ecological and behavioural traits in 78 species obtained from literature and literature reviews. Species Length Length Length Length Sexually Species' Water Average Prey MaleMin MalMax FemMin FemMax Dimorphic Range Depth Group Size Size Andrew's Beaked Whale 3.9 4.4 3.94 4.87 No Medium NA NA NA (Mesoplodon bowdoini)

Arnoux's Beaked Whale 8 9.34 8.84 9.33 No Medium Medium Medium NA (Berardius arnuxii)

Baird's Beaked Whale 9.5 11.9 10 12.8 Yes Medium Deep Medium Moridae, Gonatidae, Macrouridae, Cradchiidae (Berardius bairdii) Blainville's Beaked Whale 3.49 4.15 3.89 3.98 Yes Large Deep Solitary Cepolidae, Melamphaidae, Myctophidae (Mesoplodon densirostris)

Cuvier's Beaked Whale 5.5 6.93 5.27 7.54 Yes Large Deep Solitary Histioteuthidae, Gonatidae, Cranchiidae, (Ziphius cavirostris) Ommastrephidae, Vampyroteuthidae, Bolitaenidae,

Stauroteuthidae

Gervais' Beaked Whale 3.54 4.56 NA NA Yes Medium Deep Solitary Stomiidae, Octopoteuthidae, Lophogastridae (Mesoplodon europaeus)

Ginkgo-Toothed Beaked Whale NA NA NA NA Yes Large NA NA NA (Mesoplodon ginkgodens)

Gray's Beaked Whale NA NA NA NA Yes Medium Deep Solitary Merluccidae, Phosichthyidae, Myctophidae (Mesoplodon grayi)

Hector's Beaked Whale 3.65 4.34 NA NA Yes Medium NA Solitary Octopoteuthidae (Mesoplodon hectori)

Hubb's Beaked Whale 4.96 5.3 4.9 5.32 Yes Medium NA NA Gonatidae, Mastigoteuthidae, Melamphaidae, (Mesoplodon carlhubbsi) Onychoteuthidae, Histioteuthidae, Myctophidae,

Stomiidae, Octopoteuthidae Pygmy Beaked whale 3.26 3.72 NA NA Yes Medium NA Solitary Myctophidae, Nemipteridae (Mesoplodon peruvianus) Longman's Beaked Whale NA NA NA NA Yes Large Deep Solitary Cranchiidae, Onychoteuthidae, Chiroteuthidae, (Mesoplodon pacificus) Histioteuthidae

Northern Bottlenose Whale 7.3 9.8 6 8.7 Yes Medium Deep Solitary Gonatidae, Myopsidae, Oegposidae, Clupeidae, Gadidae (Hyperoodon ampullatus)

Southern Bottlenose Whale NA 6.94 5.7 7.45 Yes Medium Deep Solitary Cranchiidae, Onychoteuthidae, Enoploteuthidae, (Hyperoodon planifrons) Neoteuidae, Psychroteuthidae, Gonatidae

Sowerby's Beaked Whale 4.09 5.5 4.1 5.1 No Medium Shallow Solitary Gadidae, Merluccidae (Mesoplodon bidens)

Species Length Length Length Length Sexually Species' Water Average Prey MaleMin MalMax FemMin FemMax Dimorphic Range Depth Group Size Size Straptoothed Whale 4.87 5.5 5 6.25 Yes Medium NA Solitary Vampyroteuthidae, Chiroteuthidae, Cranchiidae, (Mesoplodon layardii) Cycloteuthidae, Gonatidae, Histioteuthidae,

Mastigoteuthidae, Octopoteuthidae, Ommastrephidae, Onchoteuthidae Stejneger's Beaked Whale 3.89 5.3 4.34 5.24 No Medium Deep Solitary Gonatidae, Cranchiidae (Mesoplodon stejnegeri)

Tasman Beaked Whale 5.96 7.35 NA NA No Medium Deep Solitary Merlucciidae, Serranidae, Bythitidae (Tasmacetus shepherdi)

True's Beaked Whale 4.6 5.3 4.87 5.18 Yes Medium NA NA Cranchiidae, Loliginidae (Mesoplodon mirus)

Atlantic Humpbacked Dolphin 2 2.48 NA 2.35 Yes Small Shallow Medium Haemulidae, Clupeidae, Mugilidae (Sousa teuszii)

Atlantic Spotted Dolphin 1.66 2.26 1.67 2.29 Yes Medium Shallow Medium Gadidae, Clupeidae, Carangidae, Sciaenidae, Congridae, (Stenella plagiodon/frontalis) Trichiyridae, Triglidae

Atlantic White-Sided Dolphin 2.44 2.75 1.94 2.43 Yes Medium Shallow Medium Loliginidae, Ammodytidae, Osmeridae, Scombridae (Lagenorhynchus acutus)

Black Dolphin 1.24 1.65 1.23 1.61 Yes Small Shallow Medium Munididae, Loliginidae (Cephalorhynchus eutropia)

Bottlenose Dolphin 2.02 3.81 1.9 3.67 Yes Large Medium Medium Engraulidae, Apongidae, Trichiuridae, Synodontidae, (Tursiops truncatus) Scaridae, Haemulidae, Merluccidae, Serranidae,

Clupeidae, Gadidae, Sparidae, Ophiidae, Congridae, Cepolidae, Caraengidae, Octopodidae, Loliginidae, Ommastrephidae, Sepiolidae, Sepiidae, Alphaeidae, Penaeideae, Grapsidae, Ophichthidae, Gerreidae, Mugilidae, Congiopodidae, Elopidae, Batrachoididae, Sciaenidae Clymene Dolphin 1.76 1.97 1.71 1.83 Yes Medium Deep Social Myctophidae (Stenella clymene)

Commerson's Dolphin 1.3 1.67 1.39 1.74 Yes Small Shallow Solitary Merlucciidae, Atherinopsidae, Lithodidae, Loliginidae, (Cephalorhynchus commersonii) Rhodomelaceae, Laminariaceae, Sphacelariaceae ,

Ceramiaceae, Sertulariidae, Nereidae, Mysidae, Euphausiidae, Sphaeromatidae, Diastylidae, Ophiomixidae, Styelidae, Clupeidae, Gnathiidae, Sphaeromatidae, Cirolanidae, Pyuridae Short Beaked Common 1.71 2.6 1.67 2.44 Yes Medium Medium Medium Argentinidae, Bathylagidae, Batrachoididae, Dolphin (Delphinus delphis) Melamphaidae, Scomberesocidae, Myctophidae, Sciaenidae, Engraulidae, Merlucciidae, Ophidiidae, Scombridae, Stromateidae, Loliginidae, Onychoteuthidae Dusky Dolphin 1.67 2.11 1.67 2.05 No Medium Shallow Medium Engraulidae (Lagenorhynchus obscurus)

Species Length Length Length Length Sexually Species' Water Average Prey MaleMin MalMax FemMin FemMax Dimorphic Range Depth Group Size Size False Killer Whale 3.96 6.1 3.4 5.06 Yes Large Deep Medium Lycoteuthidae, Ommastrephidae, Scombridae, (Pseudorca crassidens) Coryphaenidae, Sciaenidae, Lateolabracidae,

Salmonidae, Ariidae, Gadidae, Gonatidae, Cranchiidae Fraser's Dolphin 2.31 2.7 2.06 2.64 Yes Large Deep Social Onychoteuthidae, Ophichthyidae, Scomberoscocidae, (Lagenodelphis hosei) Octopoteuthidae, Diretmidae, Gempylidae,

Acropomatidae, Macrouridae, Bregmacerotidae, Melamphidae, Myctophidae, Neoscopelidae, Paralepididae, Scopelarhidae, Sparidae, Trichiuridae, Nomeidae, Argentinidae, Bathylagidae, Gonostomatidae, Sternoptychidae, Stomiidae, Enoploteuthidae, Lycoteuthidae, Histioteuthidae, Ctenopteridae, Brachioteuthidae, Ommastrephidae, Thysanoteuthidae, Chiroteuthidae, Mastigoteuthidae, Cranchiidae, Oplophoridae, Sergestidae Indo-Pacific Humpbacked 2 3.2 2.16 2.44 Yes Medium Shallow Medium Mugilidae, Engraulidae, Pristigasteridae, Haemulidae, Dolphin (Sousa chinensis) Sparidae, Callichthyidae, Sciaenidae, Trichiuridae

Heaviside's Dolphin 1.51 1.74 1.59 1.66 No Small Shallow Solitary Merlucciidae, Ophidiidae, Gobiinae, Loliginidae (Cephalorhynchus heavisidii)

Hector's Dolphin 1.17 1.38 1.28 1.53 Yes Small Shallow Solitary Mugilidae, Uranoscopidae, Moridae, Ommastrephidae, (Cephalorhynchus hectori) Arripidae

Hourglass Dolphin 1.63 1.87 1.66 1.83 No Small Deep Medium Mycophidae (Lagenorhynchus cruciger)

Irrawaddy Dolphin 2.2 2.35 2 2.32 Yes Small Shallow Solitary Cyprinidae, Teraponidae, Apogonidae, Chirocentridae, (Orcaella brevirostris) Pangasiidae, Cuttlefish, Engraulidae, Clupeidae,

Synodontidae, Hemirhamphidae, Psettodidae, Leigignathidae, Nemipteridae, Pomadasyidae, Sillaginidae, Platycephalidae Killer Whale 5.2 9.75 4.57 8.53 Yes Large Medium Medium Eschrichtiidae, Ohyseteridae, Balaenopteridae, (Orcinus orca) Balaenidae, Phocidae, Phocoenidae, Delphinidae,

Monodontidae, Ziphiidae, Dugongidae, Otariidae, Odobenidae, Mustelidae, Salmonidae, Gadidae, Alces alces, Dasyatidae, Myliobatidae, Lamnidae, Sebastidae, Anoplopomatidae, Pleuronectidae, Centrolophidae, Clupidae, Torpedinidae, Triakidae, Carcharhinidae, Cetorhinidae, Cervidae Long Beaked 2 2.4 1.9 2.2 Yes Small Shallow Social Engraulidae, Myctophidae, Phosichthyidae, Common Dolphin (Delphinus Atherinopsidae, Meriuccidae, Clupeidae, capensis) Centrolophidae, Carangidae, Scombridae, Triglidae, Scomberesocidae, Sphyraenidae, Normanichthyidae, Batrachoididae, Ophichthidae, Galatheidae

100

Species Length Length Length Length Sexually Species' Water Average Prey MaleMin MalMax FemMin FemMax Dimorphic Range Depth Group Size Size Long-Finned Pilot Whale 5 6.1 4.05 4.72 Yes Large Medium Medium Octopodidae, Gonatidae, Ommastreohidae, (Globicephala melaena) Mastigoteuthidae, Loligoinidae, Argentinidae, Gadidae, Pleuronectidae, Macrouridae, Zoarcidae, Ammoditidae, Trichiuridae, Pandalidae, Galatheidae, Chiroteuthidae, Brachioteuthidae, Sepiolidae, Cranchiidaae, Histioteuthidae Melon-Headed Whale 2.05 2.64 2.11 2.57 Yes Medium Deep Social Ommastrephidae, Histioteuthidae, Loliginidae, (Peponocephala electra) Onycoteuthidae, Chiroteuthidae, Mastigoteuthidae,

Cranchiidae, Enoploteruthidae, Myctophidae, Paralepididae, Scopelarchidae Northern Right Whale 2.11 3.1 1.95 2.3 Yes Medium Deep Social Gonatidae, Onychoteuthidae, Loligindae, Dolphin (Lissodelphis borealis) Enoploteuthidae, Histioteuthidae, Centrolophidae,

Melamphaidae, Scomberesocidae, Merlucciidae, Myctophidae, Bathylagidae, Paralepididae Pacific White-Sided Dolphin 1.7 2.5 1.7 2.36 Yes Medium Medium Medium Engraulidae, Merlucciidae, Loliginidae (Lagenorhynchus obliquidens)

Peale's Dolphin 1.6 2.18 1.63 2.1 No Small Shallow Solitary Myxinidae, Ommastrephidae, Clupeidae, Moridae, (Lagenorhynchus australis) Merlucciidae, Octopodidae, Loliginidae

Pygmy Killer Whale 2.07 2.59 2.07 2.45 No Large Medium Medium Delphinidae (Feresa attenuata)

Risso's Dolphin 2.53 3.6 2.4 3 No Small Medium Medium Sepiidae, Brachioteuthidae, Cranchiidae, (Grampus griseus) Mastigoteuthidae, Onychoteuthidae, Histioteuthidae,

Ommastrephidae, Argonautidae, octopodidae, Ascidiacea, Pyrosomidae, Salpidae, Enoploteuthidae, Chiroteuthidae, Loliginidae, Sepiolidae Rough-Toothed Dolphin 2.09 2.65 2.12 2.55 Yes Large Medium Medium Tremoctopodidae, Onychoteuthidae, Coryphaenidae, (Steno bredanensis) Trichiuridae, Atherinopsidae, Scomberesocidae,

Belonidae, Atherinopsidae, Centriscidae, Coryphaenidae, Ommastrephidae, Atherinidae, Loliginidae

Short-Finned Pilot Whale 4.24 4.91 3.34 3.92 Yes Medium Medium Social Octopodidae, Enoploteuthidae, Histioteuthidae, (Globicephala macrorhynchus) Loliginidae, Mastigoteuthidae, Chiroteuthidae,

Cranciidae, Brachioteuthidae, Lepidoteuthidae, Ommastrephidae, Melamphaidae Southern Right Whale Dolphin NA NA NA NA Yes Medium Medium Social Ommastrephidae, Mastigoteuthidae, Cranchiidae, (Lissodelphis peronii) Gonatidae, Bathylagidae, Photichthyidae, Myctophidae,

Merlucciidae, Engraulididae

101

Species Length Length Length Length Sexually Species' Water Average Prey MaleMin MalMax FemMin FemMax Dimorphic Range Depth Group Size Size Spinner Dolphin 1.36 2.35 1.29 2.04 Yes Large Deep Social Mastigotruthidae, Congridae, Stomiidae, Paralepididae, (Stenella longirostris) Bathylagidae, Ophichthyidae, Diretmidae,

Melamphidae, Bregmacerotidae, Centrolophidae, Macrouridae, Neoscopelidae, Acropomatidae, Sparidae, Gempylidae, Exocoetidae, Gonostomatidae, Myctophidae, Scopelarchidae, Trichiuridae, Nomeidae, Argentinidae, Oplophoridae, Penaeidae, Sergestidae, Enoploteuthidae, Octopoterthidae, Onychoteuthidae, histioteuthidae, Brachioteuthidae, Ommastrephidae, Chiroteuthidae, Cranchiidae Pan tropical Spotted Dolphin 1.66 2.57 1.63 1.63 Yes Large Deep Social NA (Stenella attenuata) Striped Dolphin 2.15 2.56 1.85 2.36 Yes Large Deep Medium Ateleopodidae, Myctophidae, Microstomatidae, (Stenella coeruleoalba) Melamphaeidae, Bathylagidae, Gempylidae,

Brachioteuthidae, Cranchiidae, Cycloteuthidae, Enoploteuthidae, Grimalditeuthidae, Histioteuthidae, Mastigoteuthidae, Octopoteuthidae, Ommastrephidae, Onychoteuthidae, Pholidoteuthidae, Tremoctopodidae, Sternoptychidae, Nomeidae, Paralepididae, Phosichthyidae, Scopelarchidae Guiana dolphin 1.31 1.87 1.38 2.06 No Small Shallow Solitary Clupeidae, Scianidae, Batrachoididae, Trichiuridae, (Sotalia guianensis) Loliginidae

White-Beaked Dolphin 2.51 3.1 1.74 2.78 Yes Medium Shallow Medium Gadidae, Clupeidae, Osmeridae (Lagenorhynchus albirostris)

Beluga 3.5 4.7 3.1 3.9 Yes Medium Medium Solitary Osmeridae, Clupeidae, Ammodytidae, Cyclopteridae, (Delphinapterus leucas) Characidae, Osmeridae, Salmonidae, Gadidae, Illicinae

Narwhal 4.1 4.7 3.4 4.15 Yes Small Deep Solitary Gadidae, Pleuronectidae, Salmonidae, Clupeidae (Monodon monoceros)

Franciscana 1.21 1.58 1.37 1.74 Yes Small Shallow Medium Gobiidae, Penaeidae, Ophidiiae, Cynoglossidae, (Pontoporia blainvillei) Trichiuridae, Carangidae, Phycidae, Antherinidae,

Poatmidae, Engraulidae, Sciaenidae, Loliginidae, Batrachoidiae, Gadidae, Stromatidae, Congridae, Stromateidae, Nomeidae Burmeister's Porpoise 1.51 1.75 1.53 1.85 Yes Small Shallow Solitary Merlucciidae, Engraulidae, Loliginidae, Euphausiidae, (Phocoena spinipinnis) Centrolophidae, Carangidae, Clupeidae, Congridae,

Centropomidae, Atherinopsidae, Sciaenidae, Gadidae, Sparidae

102

Species Length Length Length Length Sexually Species' Water Average Prey MaleMin MalMax FemMin FemMax Dimorphic Range Depth Group Size Size Cochito [Vaquita] 1.27 1.44 1.35 1.48 Yes Small Shallow Solitary (Phocoena sinus) Sciaenidae, Haemulidae

Dall's Porpoise 1.75 1.8 1.74 1.77 Yes Medium Medium Medium Bolitaenidae, Enoploteuthidae, Gonatidae, (Phocoenoides dalli) Paralepididae, Opisthoproctidae, Scombridae,

Clupeidae, Onychoteuthidae, Loliginidae, Carangidae, Osmeridae, Merlucciidae, Scomberesocidae, Engraulidae, Sebastidae, Myctophidae, Anomalopidae, Cranchiidae, Salmonidae, Paralichthyidae, Sepiolidae, Ommastrephidae, Octopoteuthidae, Histioteuthidae, Scopelarchidae, Pleuronectidae, Ophidiidae, Stromateidae, Gadidae, Lotidae, Macrouridae, Ammodytidae, Anoplopomatidae, Melamphaidae, Hexagrammidae Finless Porpoise 1.32 2.27 1.32 2.06 Yes Medium Shallow Solitary Loliginidae, Apogonidae, Leiognathidae, Sepiidae, (Neophocaena phocaenoides) Engraulidae

Harbour Porpoise 1.23 1.6 1.38 1.7 Yes Medium Shallow Solitary Octopodidae, Clupeidae, Gadidae, Gobiidae, (Phocoena phocoena) Anguillidae, Gonostomatidae, Marlucciidae, Sparidae,

Zoarcidae, Ammodytidae, Pleuronectidae, Sebastidae, Illicinae, Sternoptychidae, Merlucciidae, Stromateidae, Scombridae, Sepiolidae, Myxinidae, Loliginidae Spectacled Porpoise 1.89 2.24 1.74 2.04 Yes Small Deep Solitary Engrauliidae (Phocoena dioptrica)

Gray whale 11.1 14.6 11.7 15 Yes Small Shallow Solitary Pachychilidae, Pinnotheridae, Galatheidae, (Eschrichtius robustus) Nephropidae, Atylidae, Ampeliscidae

Blue Whale 20 25 21 33.6 Yes Large Medium Solitary Euphausiidae, Temoridae (Balaenoptera musculus)

Bryde's Whale 12 14.2 13.7 15.5 Yes Medium Medium Solitary Clupeidae, Engraulidae, Euphausiidae, Carangidae (Balaenoptera edeni)

Fin Whale 17.7 25 18.3 27 Yes Large Medium Medium Ommastrephidae, Euphausiidae, Osmeridae, Calanidae, (Balaenoptera physalus) Clupeidae, Gadidae

Antarctic Minke Whale 7.3 8.6 7.9 9 Yes Medium Deep Solitary Channichthyidae, Myctophidae, Paralepididae, (Balaenoptera bonaerensis) Nototheniidae, Euphausiidae, Hyperiidae Common Minke Whale 6.7 8.2 7.2 8.8 Yes Medium Shallow Solitary Euphausiidae, Gadidae, Salmonidae, Ammodytidae, (Balaenoptera acutorostrata) Clupeidae, Osmeridae, Scombridae, Anarhichadidae, Squalidae, Merlucciidae Sei Whale 12.8 15.9 13.3 16.1 Yes Large Deep Solitary Clausocalanidae, Hyperiidae, Temoridae, Euphausiidae, (Balaenoptera borealis) Eucalanidae, Metridinidae, Calanidae, Engraulidae,

Clupeidae, Scomberesocidae, Myctophidae

103

Species Length Length Length Length Sexually Species' Water Average Prey MaleMin MalMax FemMin FemMax Dimorphic Range Depth Group Size Size Humpback Whale 12 14.8 13.9 15.5 Yes Large Deep Solitary Mysidae, Euphausiidae, Pandalidae, Clupeidae, (Megeptera novaeangliae) Osmeridae, Gadidae

Bowhead Whale 11.6 15.5 14 18 Yes Small Medium Solitary Euphausiidae, Calanidae (Balaena mysticetus)

North Pacific Right Whale 15 17.1 15.5 18.3 Yes Medium Shallow Solitary Euphausiidae, Calanidae (Eubalaena japonica) North Atlantic Right Whale 11 12.9 11 18 Yes Small Shallow Solitary Euphausiidae, Calanidae (Eubalaena glacialis) Southern Right Whale 11.3 15.2 12.3 16.5 Yes Small Shallow Solitary Euphausiidae, Calanidae (Eubalaena australis)

Pygmy Right Whale 5.47 6.09 6 6.45 Yes Small NA Solitary Paracalanidae, Centropagidae, Calanidae, Acartiidae, (Caperea marginata) Clausocalanidae, Onceidae, Oithonidae, Hyperiidae

Indo-pacific bottlenose dolphin 2.09 2.43 2.01 2.38 No Medium Shallow Medium Apogonidae, Leiognathidae, Lethrinidae, (Tursiops aduncus) Cynnoglossidae, Congridae, Clupeidae, Carangidae, Argentinidae, Chlorophthalmidae, Citharoidea, Dactylopteridae, Gerreidae, Gempylidae, Gobiidae, Haemulidae, Holocentridae, Mugiloididae, Monacanthidae, Lutjanidae, Muraenesidae, Muraenidae, Myctophidae, Nemipteridae, Opichthidae, Ophidiidae, Platycephalidae, Pomacanthidae, Pomacentridae, Scorpaenidae, Serranidae, sparidae, Sternoptychidae, Synaphobranchidae, Trichiuridae, Synodontidae Sperm Whale 15.2 183 10.4 12.5 Yes Large Deep Solitary Ceratiidae, Gadidae, Macrouridae, Trachipteridae, (Physeter macrocephalus) Icosteidae, Scorpaenidae, Anoplomatidae, Hexagrammida, Nototheniidae, Gnathophausia, Majidae, Cancridae, Architeuthis, Ommastrephidae, Onychoteuthidae, Gonatidae, Pholidoteuthidae, Octopoteuthidae, Histipteuthidae, Cranchiidae, Vampyroteuthidae, Octopodidae Pygmy Sperm Whale 2.7 3.3 2.66 3.3 No Medium Deep Solitary Cranchiidae, Enoploteuthidae, Histioteuthidae, (Kogia breviceps) Lycoteuthidae, Ommastrephidae, Myctophidae, Gadidae, Gempylidae Dwarf Sperm Whale 2.19 2.34 2.1 2.34 No Medium Deep Solitary Ommastrephidae, Cranchiidae, Onychoteuthidae, (Kogia sima) Lycoteuthidae, Enoploteuthidae, Octopoteuthidae, Chiroteuthidae, Vampyroteuthidae, Gonatidae, Gonostomatidae, Macrouridae, Sternoptychidae, Congridae, Argentinidae, Loligiginidae, Sepiidae, Histioteuthidae, Octopodidae, Moridae, Myctophidae, Penaeidae, Acanthephyridae, Aristeidae, Argentinidae, Microstomatidae

104

Species Predators Min Max Vocalizatio Parasites Species Known to Associate with References Temp Temp n Frequency (and sources (°C) (°C) cited within) Andrew's Beaked Whale Dalatiidae 13.0 19 Medium NA NA 1, 3, 12, 202, (Mesoplodon bowdoini) 339

Arnoux's Beaked Whale Dalatiidae NA NA Medium NA Killer Whale (Orcinus orca), Hourglass 2, 9, 12, 46, (Berardius arnuxii) Dolphin (Lagenorhynchus cruciger), 101, 200, 204,

Peale's Dolphin (Lagenorhynchus 258, 340 australis) Baird's Beaked Whale Dalatiidae, Delphinidae 15.0 29 Medium Cyamidae, Lepadidae, Northern Right Whale Dolphin 7, 8, 9, 12, 101, (Berardius bairdii) Anisakidae, Tetrameridae, (Lissodelphis borealis) 200, 340 Monocotylidae, Phyllobothriidae, Diphyllobothriidae, Brachycladiidae, Cestodes, Trematodes, Nematodes Blainville's Beaked Whale Dalatiidae 25.0 29 Medium Anisakidae, Tetrabothriidae, NA 4, 9, 11, 12, 69, (Mesoplodon densirostris) Lepadidae 207, 258, 319,

339 Cuvier's Beaked Whale Dalatiidae, Lamnidae, 25.0 29 High Phyllobothriidae, Tetrameridae NA 9, 10, 11, 12, (Ziphius cavirostris) Delphinidae 79, 103, 131,

201, 207, 258, 304, 319, 338 Gervais' Beaked Whale Dalatiidae NA NA Medium Lepadidae, Phyllobothriidae, NA 9, 12, 69, 71, (Mesoplodon europaeus) Cyamidae 258, 314, 339

Ginkgo-Toothed Beaked Whale Dalatiidae NA NA Medium NA NA 9, 12, 69, 258, (Mesoplodon ginkgodens) 339

Gray's Beaked Whale Dalatiidae NA NA Medium NA NA 9, 12, 46, 69, (Mesoplodon grayi) 71, 258, 339

Hector's Beaked Whale Dalatiidae NA NA Medium Lepadidae, Phyllobothriidae NA 6, 9, 12, 46, 69, (Mesoplodon hectori) 258, 339

Hubb's Beaked Whale Dalatiidae NA NA Medium Pennellidae NA 1, 5, 12, 69, (Mesoplodon carlhubbsi) 339

Pygmy Beaked whale NA 18.2 19.3 Medium Anisakidae, Campulidae NA 12, 68, 69, 71, (Mesoplodon peruvianus) 312, 313, 339 Longman's Beaked Whale Dalatiidae 27.0 30 Medium NA Short-Finned Pilot Whale (Globicephala 12, 34, 69, 80, (Mesoplodon pacificus) macrorhynchus), Bottlenose Dolphin 309, 339

(Tursiops truncatus), Spinner Dolphin (Stenella longirostris) Northern Bottlenose Whale Delphinidae -1.3 -0.9 Medium Tetrabothriidae, Anisakidae, Killer Whale (Orcinus orca) 9, 12, 46, 49, (Hyperoodon ampullatus) Polymorphidae 103, 126, 300,

304, 336, 339

105

Species Predators Min Max Vocalizatio Parasites Species Known to Associate with References Temp Temp n Frequency (and sources (°C) (°C) cited within) Southern Bottlenose Whale Dalatiidae NA NA Medium Tetrameridae Killer Whale (Orcinus orca), Peale's 9, 12, 46, 70, (Hyperoodon planifrons) Dolphin (Lagenorhynchus australis), 126, 258, 336,

Hourglass Dolphin (Lagenorhynchus 339 cruciger) Sowerby's Beaked Whale Dalatiidae NA NA Medium Phyllobothriidae, NA 9, 12, 46, 69, (Mesoplodon bidens) Tetrabothriidae, Lepadidae, 71, 103, 173,

Tetrameridae, Anisakidae, 258, 339 Polymorphidae Straptoothed Whale Dalatiidae 10.0 16 Medium Lepadidae NA 9, 12, 46, 67, (Mesoplodon layardii) 69, 81, 258,

339 Stejneger's Beaked Whale Dalatiidae, Lamnidae NA NA Medium Tetrameridae, Tetrabothriidae NA 9, 12, 69, 71, (Mesoplodon stejnegeri) 201, 258, 310,

319, 339 Tasman Beaked Whale NA 13.0 19 NA NA NA 9, 12, 102, 123, (Tasmacetus shepherdi) 202, 308

True's Beaked Whale Dalatiidae 13.0 19 Medium Coronulidae, Pennellidae, NA 9, 12, 69, 71, (Mesoplodon mirus) Anisakidae 174, 202, 258,

315, 339 Atlantic Humpbacked Dolphin NA 18.8 24.5 High Cyamidae Bottlenose Dolphin (Tursiops truncatus) 25, 37, 137, (Sousa teuszii) 187, 262, 285,

306, 307, 340 Atlantic Spotted Dolphin NA 19.0 27 High Lepadidae, Coronulidae, Rough-Toothed Dolphin (Steno 13, 16, 38, 46, (Stenella plagiodon/frontalis) Cyamidae, Echeneidae, bredanensis), Bottlenose Dolphin 94, 212, 217,

Campulidae, Heterophyidae, (Tursiops truncatus), Risso's Dolphin 340, 341, 345 Brauninidae, Pseudaliidae, (Grampus griseus), Pan tropical Spotted Anisakidae Dolphin (Stenella attenuata) Atlantic White-Sided Dolphin NA 4.0 10 High Heterophyidae, Brachycladiidae, Killer Whale (Orcinus orca), Bottlenose 17, 21, 26, 27, (Lagenorhynchus acutus) Tetrabothriidae, Dolphin (Tursiops truncatus), Short 49, 51, 50, 51,

Phyllobothriidae, Beaked Common Dolphin (Delphinus 55, 340, 344, Tetrabothriidae, Pseudaliidae , delphis), Long-Finned Pilot Whale 346 Tetrameridae, Pseudaliidae, (Globicephala melas), White-Beaked Polymorphidae, Pseudaliidae, Dolphin (Lagenorhynchus albirostris), Heterophyidae Fin Whale (Balaenoptera physalus), Humpback Whale (Megeptera novaeangliae) Black Dolphin NA 11.0 14.6 Low Anisakidae, Brauninidae, Peale's Dolphin (Lagenorhynchus 17, 46, 60, 111, (Cephalorhynchus eutropia) Campulidae, Polymorphidae australis), Commerson's Dolphin 170, 188, 348

(Cephalorhynchus commersonii)

106

Species Predators Min Max Vocalizatio Parasites Species Known to Associate with References Temp Temp n Frequency (and sources (°C) (°C) cited within) Bottlenose Dolphin Dalatiidae, 15.0 29 High Brauninidae, Nasitrematidae, Guiana dolphin (Sotalia guianensis), 13, 15, 17, 154, (Tursiops truncatus) Carcharhinidae, Heterophyidae, Anisakidae, Atlantic Spotted Dolphin (Stenella 158, 159, 160,

Lamnidae, Hexanchidae Brachycladiidae, plagiodon/frontalis), Melon-Headed 186, 187, 207, Phyllobothriidae, Campulidae, Whale (Peponocephala electra), Short- 246, 265, 266, Tetrameridae, Finned Pilot Whale (Globicephala 319, 322, 326, Diphyllobothriidae, macrorhynchus), Atlantic Humpbacked 340, 341 Pseudaliidae, Polymorphidae Dolphin (Sousa teuszii), Longman's Beaked Whale (Mesoplodon pacificus), Atlantic White-Sided Dolphin (Lagenorhynchus acutus), Humpback Whale (Megeptera novaeangliae), Common Minke Whale (Balaenoptera acutorostrata), Harbour Porpoise (Phocoena phocoena), Burmeister's Porpoise (Phocoena spinipinnis), Pan tropical Spotted Dolphin (Stenella attenuata), Southern Right Whale Dolphin (Lissodelphis peronii), Risso's Dolphin (Grampus griseus), Hourglass Dolphin (Lagenorhynchus cruciger), Killer Whale (Orcinus orca), Northern Right Whale Dolphin (Lissodelphis borealis), Pacific White-Sided Dolphin (Lagenorhynchus obliquidens), Rough- Toothed Dolphin (Steno bredanensis), Gray whale (Eschrichtius robustus), White-Beaked Dolphin (Lagenorhynchus albirostris), False Killer Whale (Pseudorca crassidens), Indo-Pacific Humpbacked Dolphin (Sousa chinensis), Sperm Whale (Physeter macrocephalus), Blue Whale (Balaenoptera musculus) Clymene Dolphin Dalatiidae 20.2 28.5 High Coronulidae, Cyamidae, Pan tropical Spotted Dolphin (Stenella 13, 16, 61, 62, (Stenella clymene) Pseudaliidae, Phyllobothriidae, attenuata), Short Beaked Common 130, 169, 172,

Nasitrematidae Dolphin (Delphinus delphis), Melon- 212, 217, 340, Headed Whale (Peponocephala electra) 341, 345

107

Species Predators Min Max Vocalizatio Parasites Species Known to Associate with References Temp Temp n Frequency (and sources (°C) (°C) cited within) Commerson's Dolphin NA 1.0 16 Low Brachycladiidae, Anisakidae, Peale's Dolphin (Lagenorhynchus 17, 46, 108, (Cephalorhynchus commersonii) Brauninidae, Heterophyidae, australis), Black Dolphin 109, 110, 185,

Tetrabothriidae (Cephalorhynchus eutropia), 200, 348 Burmeister's Porpoise (Phocoena spinipinnis) Short Beaked Common Dalatiidae, 7.0 23 High Anisakidae, Pseudaliidae, Striped Dolphin (Stenella coeruleoalba), 15, 17, 23, 24, Dolphin (Delphinus delphis) Carcharhinidae, Tetrameridae, Phyllobothriidae, Risso's Dolphin (Grampus griseus), 27, 94, 95, 124, Lamnidae, Delphinidae, Campulidae, Heterophyidae Clymene Dolphin (Stenella clymene), 143, 149, 201, Sphyrnidae Atlantic White-Sided Dolphin 202, 205, 246, (Lagenorhynchus acutus), Northern 258, 318, 319, Right Whale Dolphin (Lissodelphis 340, 341 borealis), Pacific White-Sided Dolphin (Lagenorhynchus obliquidens), Southern Right Whale Dolphin (Lissodelphis peronii), Gray whale (Eschrichtius robustus), Dusky Dolphin (Lagenorhynchus obscurus), Risso's Dolphin (Grampus griseus), Long- Finned Pilot Whale (Globicephala melas), Spinner Dolphin (Stenella longirostris) Dusky Dolphin Hexanchidae, 10.4 19 High Tetrabothriidae, Southern Right Whale Dolphin 17, 46, 49, 50, (Lagenorhynchus obscurus) Delphinidae, Lamnidae Phyllobothriidae, Anisakidae, (Lissodelphis peronii), Short Beaked 51, 55, 82, 201,

Trematodes, Pseudaliidae, Common Dolphin (Delphinus delphis), 202, 205, 304, Brauninidae Long-Finned Pilot Whale (Globicephala 319, 340, 344, melas), Killer Whale (Orcinus orca), 346 Risso's Dolphin (Grampus griseus), Burmeister's Porpoise (Phocoena spinipinnis), Heaviside's Dolphin (Cephalorhynchus heavisidii) False Killer Whale Dalatiidae, Delphinidae 9.0 31 Medium Anisakidae, Pseudaliidae, Killer Whale (Orcinus orca), Indo-pacific 16, 41, 42, 84, (Pseudorca crassidens) Polymorphidae, bottlenose dolphin(Tursiops aduncus), 207, 225, 258,

Brachycladiidae, Rough-Toothed Dolphin (Steno 319, 340 Nasitrematidae, Cyamidae, bredanensis), Fraser's Dolphin Coronulidae (Lagenodelphis hosei), Risso's Dolphin (Grampus griseus), Bottlenose Dolphin (Tursiops truncatus), Melon-Headed Whale (Peponocephala electra), Short- Finned Pilot Whale (Globicephala macrorhynchus)

108

Species Predators Min Max Vocalizatio Parasites Species Known to Associate with References Temp Temp n Frequency (and sources (°C) (°C) cited within) Fraser's Dolphin Dalatiidae 25.0 29 High Tetrabothriidae, Sperm Whale (Physeter macrocephalus), 16, 17, 96, 122, (Lagenodelphis hosei) Phyllobothriidae, False Killer Whale (Pseudorca 132, 133, 134,

Tetrabothriidae, crassidens), Striped Dolphin (Stenella 207, 340, 344 Brachycladiidae, Anisakidae, coeruleoalba), Pan tropical Spotted Pseudaliidae, Polymorphidae Dolphin (Stenella attenuata), Spinner Dolphin (Stenella longirostris), Short- Finned Pilot Whale (Globicephala macrorhynchus), Melon-Headed Whale (Peponocephala electra), Risso's Dolphin (Grampus griseus) Indo-Pacific Humpbacked Lamnidae, Delphinidae 23.9 29.6 High Anisakidae, Pseudaliidae, Bottlenose Dolphin (Tursiops truncatus), 37, 187, 189, Dolphin (Sousa chinensis) Cyamidae Killer Whale (Orcinus orca), Southern 262, 273, 306,

Right Whale (Eubalaena australis), Long 319, 340 Beaked common Dolphin (Delphinus capensis) Heaviside's Dolphin Lamnidae, Hexanchidae 9.0 19 Low Cyamidae, Coronulidae Dusky Dolphin (Lagenorhynchus 17, 31, 200, (Cephalorhynchus heavisidii) obscurus) 203

Hector's Dolphin Hexanchidae, 6.3 22 Low Cyamidae, Strigeidae, NA 17, 97, 112, (Cephalorhynchus hectori) Carcharhinidae Brauninidae, Halocercinae, 129, 179, 200,

Pseudaliidae , Anisakidae, 201, 319, 203 Acariidae, Polymorphidae, Campulidae, Phyllobothriidae Hourglass Dolphin NA -0.3 13.4 High Anisakidae Fin Whale (Balaenoptera physalus), 46, 49, 50, 51, (Lagenorhynchus cruciger) Antarctic Minke Whale (Balaenoptera 55, 58, 136,

bonaerensis), Common Minke Whale 205, 275, 340, (Balaenoptera acutorostrata), Sei Whale 344, 346 (Balaenoptera borealis), Bottlenose Dolphin (Tursiops truncatus), Southern Right Whale Dolphin (Lissodelphis peronii), Long-Finned Pilot Whale (Globicephala melas), Arnoux's Beaked Whale (Berardius arnuxii), Southern Bottlenose Whale (Hyperoodon planifrons), Killer Whale (Orcinus orca) Irrawaddy Dolphin Carcharhinidae 20.0 35 Low Trematode , Schistosomatidae, Finless Porpoise (Neophocaena 17, 40, 47, 139, (Orcaella brevirostris) Nematode phocaenoides), Spinner Dolphin 180, 253, 321,

(Stenella longirostris) 340

109

Species Predators Min Max Vocalizatio Parasites Species Known to Associate with References Temp Temp n Frequency (and sources (°C) (°C) cited within) Killer Whale Dalatiidae 1.7 26 Medium Fasciolidae, Tetrabothriidae, Southern Right Whale (Eubalaena 16, 17, 103, (Orcinus orca) Phyllobothriidae, Anisakidae australis), Sei Whale (Balaenoptera 149, 204, 205,

borealis), Fin Whale (Balaenoptera 207, 225, 258, physalus), Bryde's Whale (Balaenoptera 292, 293, 294, edeni), Blue Whale (Balaenoptera 303, 304, 305, musculus), Gray whale (Eschrichtius 306, 340 robustus), Harbour Porpoise (Phocoena phocoena), Dall's Porpoise (Phocoenoides dalli), Narwhal (Monodon monoceros), Beluga (Delphinapterus leucas), White-Beaked Dolphin (Lagenorhynchus albirostris), Risso's Dolphin (Grampus griseus), Peale's Dolphin (Lagenorhynchus australis), Pacific White-Sided Dolphin (Lagenorhynchus obliquidens), Arnoux's Beaked Whale (Berardius arnuxii), Northern Bottlenose Whale (Hyperoodon ampullatus), Southern Bottlenose Whale (Hyperoodon planifrons), Atlantic White-Sided Dolphin (Lagenorhynchus acutus), Bottlenose Dolphin (Tursiops truncatus), Dusky Dolphin (Lagenorhynchus obscurus), False Killer Whale (Pseudorca crassidens), Indo-Pacific Humpbacked Dolphin, Long-Finned Pilot Whale (Globicephala melas), Humpback Whale (Megeptera novaeangliae), Hourglass Dolphin (Lagenorhynchus cruciger), Short-Finned Pilot Whale (Globicephala macrorhynchus) Long Beaked Dalatiidae NA NA High Anisakidae Pacific White-Sided Dolphin 94, 95, 124, Common Dolphin (Delphinus (Lagenorhynchus obliquidens), Indo- 156, 173, 184, capensis) Pacific Humpbacked Dolphin (Sousa 197, 258, 340, chinensis), Bryde's Whale (Balaenoptera 341 edeni)

110

Species Predators Min Max Vocalizatio Parasites Species Known to Associate with References Temp Temp n Frequency (and sources (°C) (°C) cited within) Long-Finned Pilot Whale Delphinidae 0.6 22 High Brachycladiidae, Southern Right Whale Dolphin 15, 17, 24, 46, (Globicephala melaena) Phyllobothriidae, (Lissodelphis peronii), Humpback 85, 103, 115, Diphyllobothriidae, Whale (Megeptera novaeangliae), Killer 202, 205, 304, Tetrabothriidae, Anisakidae, Whale (Orcinus orca), Atlantic White- 343 Pseudaliidae, Polymorphidae Sided Dolphin (Lagenorhynchus acutus), Hourglass Dolphin (Lagenorhynchus cruciger), Peale's Dolphin (Lagenorhynchus australis), White-Beaked Dolphin (Lagenorhynchus albirostris), Pygmy Right Whale (Caperea marginata), Dusky Dolphin (Lagenorhynchus obscurus), Short Beaked Common Dolphin (Delphinus delphis) Melon-Headed Whale Dalatiidae 25.0 29 High Nasitrematidae, Bottlenose Dolphin (Tursiops truncatus), 16, 49, 63, 64, (Peponocephala electra) Phyllobothriidae, Pseudaliidae, Spinner Dolphin (Stenella longirostris), 65, 66, 103,

Anisakidae Pygmy Killer Whale (Feresa attenuata), 207, 258, 346 Fraser's Dolphin (Lagenodelphis hosei), Humpback Whale (Megeptera novaeangliae), Short-Finned Pilot Whale (Globicephala macrorhynchus), Rough- Toothed Dolphin (Steno bredanensis), Clymene Dolphin (Stenella clymene), Pan tropical Spotted Dolphin (Stenella attenuata), False Killer Whale (Pseudorca crassidens), Sperm Whale (Physeter macrocephalus)

111

Species Predators Min Max Vocalizatio Parasites Species Known to Associate with References Temp Temp n Frequency (and sources (°C) (°C) cited within) Northern Right Whale NA 8.0 19 High Nasitrematidae, Tetrameridae, Pacific White-Sided Dolphin 36, 46, 178, Dolphin (Lissodelphis borealis) Anisakidae, Phyllobothriidae, (Lagenorhynchus obliquidens), Short 311, 347

Coronulidae, Pennellidae Beaked Common Dolphin (Delphinus delphis), Bottlenose Dolphin (Tursiops truncatus), Risso's Dolphin (Grampus griseus), Short-Finned Pilot Whale (Globicephala macrorhynchus), Dall's Porpoise (Phocoenoides dalli), Baird's Beaked Whale (Berardius bairdii), Sperm Whale (Physeter macrocephalus), Fin Whale (Balaenoptera physalus), Gray whale (Eschrichtius robustus), Humpback Whale (Megeptera novaeangliae), Sei Whale (Balaenoptera borealis) Pacific White-Sided Dolphin Delphinidae, Lamnidae 12.0 13 High Nasitrematidae, Northern Right Whale Dolphin 17, 46, 50, 51, (Lagenorhynchus obliquidens) Brachycladiidae, (Lissodelphis borealis), Killer Whale 52, 53, 54, 55,

Phyllobothriidae, (Orcinus orca), Risso's Dolphin 210, 317, 319, Tetrabothriidae, Anisakidae, (Grampus griseus), Striped Dolphin 340, 344, 346 Tetrameridae (Stenella coeruleoalba), Short Beaked Common Dolphin (Delphinus delphis), Long Beaked common Dolphin (Delphinus capensis), Short-Finned Pilot Whale (Globicephala macrorhynchus), Bottlenose Dolphin (Tursiops truncatus), Sperm Whale (Physeter macrocephalus), Gray whale (Eschrichtius robustus), Blue Whale (Balaenoptera musculus), Fin Whale (Balaenoptera physalus), Sei Whale (Balaenoptera borealis), Humpback Whale (Megeptera novaeangliae), Harbour Porpoise (Phocoena phocoena), Dall's Porpoise (Phocoenoides dalli), Southern Right Whale Dolphin (Lissodelphis peronii)

112

Species Predators Min Max Vocalizatio Parasites Species Known to Associate with References Temp Temp n Frequency (and sources (°C) (°C) cited within) Peale's Dolphin NA 6.0 9 High Tetrabothriidae Long-Finned Pilot Whale (Globicephala 50, 51, 55, 56, (Lagenorhynchus australis) melas), Arnoux's Beaked Whale 57, 182, 183,

(Berardius arnuxii), Killer Whale 340, 344, 346 (Orcinus orca), Southern Right Whale Dolphin (Lissodelphis peronii), Southern Bottlenose Whale (Hyperoodon planifrons), Black Dolphin (Cephalorhynchus eutropia), Antarctic Minke Whale (Balaenoptera bonaerensis), Fin Whale (Balaenoptera physalus), Sei Whale (Balaenoptera borealis), Risso's Dolphin (Grampus griseus), Commerson's Dolphin (Cephalorhynchus commersonii), Southern Right Whale (Eubalaena australis) Pygmy Killer Whale Dalatiidae 25.0 29 NA Lepadidae, Anisakidae, Rough-Toothed Dolphin (Steno 16, 48, 207, (Feresa attenuata) Pseudaliidae, Tetrabothriidae, bredanensis), Melon-Headed Whale 210, 258, 316,

Nasitrematidae (Peponocephala electra), Risso's Dolphin 319 (Grampus griseus)

113

Species Predators Min Max Vocalizatio Parasites Species Known to Associate with References Temp Temp n Frequency (and sources (°C) (°C) cited within) Risso's Dolphin Dalatiidae, Lamnidae 7.5 35 Medium Phyllobothriidae, Bottlenose Dolphin (Tursiops truncatus), 13, 16, 44, 59, (Grampus griseus) Tetrabothriidae, Tetrameridae, Fin Whale (Balaenoptera physalus), 83, 200, 205,

Pseudaliidae, Anisakidae, Sperm Whale (Physeter macrocephalus), 210, 258, 316, Nasitrematidae, Cyamidae, Short-Finned Pilot Whale (Globicephala 319 Coronulidae macrorhynchus), Short Beaked Common Dolphin (Delphinus delphis), Northern Right Whale Dolphin (Lissodelphis borealis), Dall's Porpoise (Phocoenoides dalli), Striped Dolphin (Stenella coeruleoalba), Killer Whale (Orcinus orca), Pan tropical Spotted Dolphin (Stenella attenuata), False Killer Whale (Pseudorca crassidens), Pygmy Killer Whale (Feresa attenuata), Short Beaked Common Dolphin (Delphinus delphis), Fraser's Dolphin (Lagenodelphis hosei), Pacific White-Sided Dolphin (Lagenorhynchus obliquidens), Dusky Dolphin (Lagenorhynchus obscurus), Atlantic Spotted Dolphin (Stenella plagiodon/frontalis), Peale's Dolphin (Lagenorhynchus australis), Gray whale (Eschrichtius robustus), Rough-Toothed Dolphin (Steno bredanensis) Rough-Toothed Dolphin Dalatiidae, Delphinidae 25.0 29 Medium Tetrabothriidae, Polymorphidae, False Killer Whale (Pseudorca 13, 16, 17, 90, (Steno bredanensis) Anisakidae, Cyamidae crassidens), Bottlenose Dolphin 122, 201, 207,

(Tursiops truncatus), Pygmy Killer 262, 340 Whale (Feresa attenuata), Atlantic Spotted Dolphin (Stenella plagiodon/frontalis), Melon-Headed Whale (Peponocephala electra), Humpback Whale (Megeptera novaeangliae), Spinner Dolphin (Stenella longirostris), Pan tropical Spotted Dolphin (Stenella attenuata), Common Minke Whale (Balaenoptera acutorostrata), Short-Finned Pilot Whale (Globicephala macrorhynchus), Bottlenose Dolphin (Tursiops truncatus), Bryde's Whale (Balaenoptera edeni), Blue Whale (Balaenoptera musculus)

114

Species Predators Min Max Vocalizatio Parasites Species Known to Associate with References Temp Temp n Frequency (and sources (°C) (°C) cited within) Short-Finned Pilot Whale Dalatiidae 25.0 29 High Brachycladiidae, Campulidae, Bottlenose Dolphin (Tursiops truncatus), 13, 17, 85, 103, (Globicephala macrorhynchus) Nasitrematidae, Tetrabothriidae, Fraser's Dolphin (Lagenodelphis hosei), 116, 117, 118,

Phyllobothriidae, Anisakidae, Melon-Headed Whale (Peponocephala 195, 207, 258, Pseudaliidae electra), Northern Right Whale Dolphin 343 (Lissodelphis borealis), Pacific White- Sided Dolphin (Lagenorhynchus obliquidens), Risso's Dolphin (Grampus griseus), Gray whale (Eschrichtius robustus), Longman's Beaked Whale (Mesoplodon pacificus), Pan tropical Spotted Dolphin (Stenella attenuata), False Killer Whale (Pseudorca crassidens), Rough-Toothed Dolphin (Steno bredanensis), Killer Whale (Orcinus orca), Sperm Whale (Physeter macrocephalus) Southern Right Whale Dolphin Somniosidae, 1.0 20 High Nasitrematidae, Pseudaliidae, Dusky Dolphin (Lagenorhynchus 35, 36, 46, 181, (Lissodelphis peronii) Nototheniidae Anisakidae, Tetrabothriidae, obscurus), Bottlenose Dolphin (Tursiops 201, 202, 205,

Opisthorchiidae, truncatus), Long-Finned Pilot Whale 319, 347 Phyllobothriidae (Globicephala melas), Fin Whale (Balaenoptera physalus), Hourglass Dolphin (Lagenorhynchus cruciger), Peale's Dolphin (Lagenorhynchus australis), Pacific White-Sided Dolphin (Lagenorhynchus obliquidens), Short Beaked Common Dolphin (Delphinus delphis) Spinner Dolphin Dalatiidae, Lamnidae, 22.0 27.5 High Echeneidae, Lepadidae, Bottlenose Dolphin (Tursiops truncatus), 13, 16, 17, 92, (Stenella longirostris) Carcharhinidae, Anisakidae, Pseudaliidae, Pan tropical Spotted Dolphin (Stenella 120, 121, 149,

Tetraodontidae, Spiruridae, Brachycladiidae, attenuata), Rough-Toothed Dolphin 207, 212, 213, Delphinidae Nasitrematidae, Tetrabothriidae, (Steno bredanensis), Longman's Beaked 317, 319, 323, Phyllobothriidae, Whale (Mesoplodon pacificus), Fraser's 340, 341, 345 Polymorphidae Dolphin (Lagenodelphis hosei), Melon- Headed Whale (Peponocephala electra), Fraser's Dolphin (Lagenodelphis hosei), Irrawaddy Dolphin (Orcaella brevirostris), Indo-pacific bottlenose dolphin(Tursiops aduncus), Short Beaked Common Dolphin (Delphinus delphis), Bryde's Whale (Balaenoptera edeni)

115

Species Predators Min Max Vocalizatio Parasites Species Known to Associate with References Temp Temp n Frequency (and sources (°C) (°C) cited within) Pan tropical Spotted Dolphin Dalatiidae, Delphinidae, 25.3 28 High Phyllobothriidae, Atlantic Spotted Dolphin (Stenella 13, 16, 17, 38, (Stenella attenuata) Caleocerdo cuvier Tetrabothriidae, Brauninidae, plagiodon/frontalis), Spinner Dolphin 39, 46, 103, Nasitrematidae, Anisakidae, (Stenella longirostris), Clymene Dolphin 207, 209, 212, Tetrameridae (Stenella clymene), Bottlenose Dolphin 217, 258, 319, (Tursiops truncatus), Fraser's Dolphin 323, 340, 341, (Lagenodelphis hosei), Risso's Dolphin 345 (Grampus griseus), Rough-Toothed Dolphin (Steno bredanensis), Melon- Headed Whale (Peponocephala electra), Short-Finned Pilot Whale (Globicephala macrorhynchus), Bryde's Whale (Balaenoptera edeni) Striped Dolphin Dalatiidae 20.0 30 High Cyamidae , Pennellidae, Short Beaked Common Dolphin 13, 16, 17, 89, (Stenella coeruleoalba) Lepadidae, Coronulidae, (Delphinus delphis), Fraser's Dolphin 93, 150, 151,

Tetrabothriidae, (Lagenodelphis hosei), Pacific White- 205, 217, 258, Phyllobothriidae, Sided Dolphin (Lagenorhynchus 340, 341, 345 Nasitrematidae, obliquidens), Risso's Dolphin (Grampus Brachycladiidae, Heterophyidae, griseus), Gray whale (Eschrichtius Campulidae, Brachycladiidae, robustus), Fin Whale (Balaenoptera Anisakidae, Anguillicolidae, physalus) Tetrameridae, Pseudaliidae, Polymorphidae Guiana dolphin Dalatiidae, Delphinidae, 15.0 31 High Nasitrematidae, Pseudaliidae, Bottlenose Dolphin (Tursiops truncatus) 17, 22, 46, 91, (Sotalia guianensis) Carcharhinidae Anisakidae 201, 214, 252,

258, 340, 341 White-Beaked Dolphin Delphinidae 8.1 17.2 High Anisakidae, Pseudaliidae, Harbour Porpoise (Phocoena phocoena), 21, 23, 46, 50, (Lagenorhynchus albirostris) Cyamidae Killer Whale (Orcinus orca), Long- 51, 55, 152,

Finned Pilot Whale (Globicephala 194, 340, 344, melas), Atlantic White-Sided Dolphin 346 (Lagenorhynchus acutus), Fin Whale (Balaenoptera physalus), Humpback Whale (Megeptera novaeangliae), Sei Whale (Balaenoptera borealis), Short Beaked Common Dolphin (Delphinus delphis), Bottlenose Dolphin (Tursiops truncatus) Beluga Delphinidae, Ursidae, 0.0 16 Medium Brachycladiidae, Campulidae, Narwhal (Monodon monoceros), Killer 14, 17, 49, 99, (Delphinapterus leucas) Somniosidae Diphyllobothriidae, Anisakidae, Whale (Orcinus orca) 103, 193, 201,

Tetrameridae, Pseudaliidae, 218, 219, 221, Ascarididae, Polymorphidae 222, 319, 325, 340

116

Species Predators Min Max Vocalizatio Parasites Species Known to Associate with References Temp Temp n Frequency (and sources (°C) (°C) cited within) Narwhal Odobenidae, -2.0 5 High Cyamidae, Anisakidae, Beluga (Delphinapterus leucas), Killer 17, 46, 100, (Monodon monoceros) Delphinidae, Ursidae, Ascarididae, Pseudaliidae Whale (Orcinus orca) 145, 201, 221,

Somniosidae, 254, 253, 319, 320, 323, 340 Franciscana Delphinidae, 16.0 27 NA Coronulidae, Cirolanidae, NA 17, 33, 98, 248, (Pontoporia blainvillei) Carcharhinidae, Cymothoidae, Cocconeidaceae, 249, 250, 251,

Hexanchidae Anisakidae, Polymorphidae 319, 323 Burmeister's Porpoise NA 4.0 19.5 Low Campulidae, Heterophyidae, Bottlenose Dolphin (Tursiops truncatus), 17, 46, 86, 114, (Phocoena spinipinnis) Nasitrematidae, Brauninidae, Dusky Dolphin (Lagenorhynchus 284, 289

Anisakidae, Pseudaliidae, obscurus), Commerson's Dolphin Polymorphidae, Cyamidae, (Cephalorhynchus commersonii) Coronulidae Cochito [Vaquita] Lamnidae, 17.0 32 Low Tetrameridae, Campulidae, NA 17, 46, 113, (Phocoena sinus) Carcharhinidae, Coronulidae 175, 244, 245, Alopiidae, Hexanchidae, 246, 289 Dall's Porpoise Delphinidae, Lamnidae 12.4 24 NA Pseudaliidae, Nasitrematidae, Harbour Porpoise (Phocoena phocoena), 17, 46, 87, 88, (Phocoenoides dalli) Brachycladiidae, Anisakidae, Killer Whale (Orcinus orca), Fin Whale 210, 232, 235,

Tetrameridae, Phyllobothriidae, (Balaenoptera physalus), Blue Whale 236, 319 Polymorphidae, Cyamidae (Balaenoptera musculus), Gray whale (Eschrichtius robustus), Humpback Whale (Megeptera novaeangliae), Northern Right Whale Dolphin (Lissodelphis borealis), Pacific White- Sided Dolphin (Lagenorhynchus obliquidens), Risso's Dolphin (Grampus griseus), Sei Whale (Balaenoptera borealis) Finless Porpoise Dalatiidae, Lamnidae 5.7 25.6 NA Brachycladiidae, Irrawaddy Dolphin (Orcaella 17, 28, 29, 30, (Neophocaena phocaenoides) Nasitrematidae, Tetrameridae, brevirostris) 87, 240, 241,

Pseudaliidae, 242, 258 Diphyllobothriidae Harbour Porpoise Dalatiidae, Delphinidae, 6.0 17 Low Brachycladiidae, Campulidae, Dall's Porpoise (Phocoenoides dalli), 17, 21, 75, 76, (Phocoena phocoena) Lamnidae, Somniosidae, Opisthorchiidae, Heterophyidae, Bottlenose Dolphin (Tursiops truncatus), 77, 103, 176,

Diphyllobothriidae, Anisakidae, Killer Whale (Orcinus orca), Pacific 177, 224, 227, Pseudaliidae, Ascarididae, White-Sided Dolphin (Lagenorhynchus 232, 233, 234, Polymorphidae obliquidens), White-Beaked Dolphin 258, 289, 319 (Lagenorhynchus albirostris), Fin Whale (Balaenoptera physalus), Common Minke Whale (Balaenoptera acutorostrata), Humpback Whale (Megeptera novaeangliae)

117

Species Predators Min Max Vocalizatio Parasites Species Known to Associate with References Temp Temp n Frequency (and sources (°C) (°C) cited within) Spectacled Porpoise NA 5.5 9.5 Low NA NA 17, 32, 238, (Phocoena dioptrica) 289

Gray whale Delphinidae 0.0 25 Low Notocotylidae, Tetrabothriidae, Killer Whale (Orcinus orca), Bottlenose 14, 19, 49, 103, (Eschrichtius robustus) Pseudophyllidae, Dolphin (Tursiops truncatus), Striped 135, 222, 253,

Polymorphidae Dolphin (Stenella coeruleoalba), Short 340 Beaked Common Dolphin (Delphinus delphis), Dall's Porpoise (Phocoenoides dalli), Short-Finned Pilot Whale (Globicephala macrorhynchus), Northern Right Whale Dolphin (Lissodelphis borealis), Pacific White- Sided Dolphin (Lagenorhynchus obliquidens), Risso's Dolphin (Grampus griseus) Blue Whale Dalatiidae, Delphinidae 14.1 21.6 Low Notocotylidae, Tetrabothriidae, Killer Whale (Orcinus orca), Fin Whale 16, 18, 49, 125, (Balaenoptera musculus) Anisakidae, Tetrameridae, (Balaenoptera physalus), Pacific White- 141, 161, 164,

Ascarididae, Polymorphidae Sided Dolphin (Lagenorhynchus 166, 258, 300, obliquidens), Dall's Porpoise 304, 340 (Phocoenoides dalli), Bottlenose Dolphin (Tursiops truncatus), Rough-Toothed Dolphin (Steno bredanensis) Bryde's Whale Dalatiidae, Delphinidae 18.5 19 Low Polymorphidae Killer Whale (Orcinus orca), Long 16, 18, 49, 128, (Balaenoptera edeni) Beaked common Dolphin (Delphinus 141, 164, 166,

capensis), Rough-Toothed Dolphin 174, 196, 246, (Steno bredanensis), Indo-pacific 247, 258, 291, bottlenose dolphin (Tursiops aduncus), 292, 302, 340 Spinner Dolphin (Stenella longirostris), Pan tropical Spotted Dolphin (Stenella attenuata)

118

Species Predators Min Max Vocalizatio Parasites Species Known to Associate with References Temp Temp n Frequency (and sources (°C) (°C) cited within) Fin Whale Dalatiidae, Delphinidae 16.0 23 Low Brachycladiidae, Notocotylidae, Killer Whale (Orcinus orca), Blue Whale 18, 21, 49, 103, (Balaenoptera physalus) Diphyllobothriidae, (Balaenoptera musculus), Atlantic 141, 164, 166,

Phyllobothriidae, White-Sided Dolphin (Lagenorhynchus 167, 246, 258, Tetrabothriidae, Anisakidae, acutus), Hourglass Dolphin 300, 304, 340 Tetrameridae, Polymorphidae (Lagenorhynchus cruciger), Northern Right Whale Dolphin (Lissodelphis borealis), Pacific White-Sided Dolphin (Lagenorhynchus obliquidens), Peale's Dolphin (Lagenorhynchus australis), Risso's Dolphin (Grampus griseus), Southern Right Whale Dolphin (Lissodelphis peronii), White-Beaked Dolphin (Lagenorhynchus albirostris), Dall's Porpoise (Phocoenoides dalli), Harbour Porpoise (Phocoena phocoena), Striped Dolphin (Stenella coeruleoalba) Antarctic Minke Whale Dalatiidae, Delphinidae -1.9 21.8 Low Pennellidae, Cyamidae, Hourglass Dolphin (Lagenorhynchus 46, 78, 140, (Balaenoptera bonaerensis) Anisakidae cruciger), Peale's Dolphin 141, 142, 164, (Lagenorhynchus australis), Pygmy 166, 189, 204, Right Whale (Caperea marginata) 205, 206, 258, 282, 301, 340 Common Minke Whale Dalatiidae, Delphinidae 6.1 20.1 Low Pennellidae, Brachycladiidae, Humpback Whale (Megeptera 21, 46, 140, (Balaenoptera acutorostrata) Diphyllobothriidae, novaeangliae), Rough-Toothed Dolphin 141, 144, 164, Tetrabothriidae, Anisakidae, (Steno bredanensis), Bottlenose Dolphin 166, 258, 300, Cyamidae (Tursiops truncatus), Hourglass Dolphin 340 (Lagenorhynchus cruciger), Harbour Porpoise (Phocoena phocoena) Sei Whale Dalatiidae, Delphinidae 5.0 18.8 Low Notocotylidae, Killer Whale (Orcinus orca), Hourglass 18, 49, 103, (Balaenoptera borealis) Diphyllobothriidae, Dolphin (Lagenorhynchus cruciger), 140, 141, 164,

Tetrabothriidae, Polymorphidae, Northern Right Whale Dolphin 166, 206, 258, Tetrameridae, Anisakidae (Lissodelphis borealis), Pacific White- 282, 283, 301, Sided Dolphin (Lagenorhynchus 304, 340 obliquidens), Peale's Dolphin (Lagenorhynchus australis), White- Beaked Dolphin (Lagenorhynchus albirostris), Pygmy Right Whale (Caperea marginata), North Atlantic Right whale (Eubalaena glacialis), Humpback Whale (Megeptera novaeangliae), Dall's Porpoise (Phocoenoides dalli)

119

Species Predators Min Max Vocalizatio Parasites Species Known to Associate with References Temp Temp n Frequency (and sources (°C) (°C) cited within) Humpback Whale Dalatiidae, Delphinidae 13.0 19 Low Diphyllobothriidae, Anisakidae, Bottlenose Dolphin (Tursiops truncatus), 17, 18, 21, 49, (Megeptera novaeangliae) Brachycladiidae, Tetrameridae, Killer Whale (Orcinus orca), Common 103, 138, 162,

Polymorphidae Minke Whale (Balaenoptera 202, 204, 205, acutorostrata), Rough-Toothed Dolphin 258, 261, 300, (Steno bredanensis), Dall's Porpoise 340 (Phocoenoides dalli), Atlantic White- Sided Dolphin (Lagenorhynchus acutus), Long-Finned Pilot Whale (Globicephala melas), Melon-Headed Whale (Peponocephala electra), Northern Right Whale Dolphin (Lissodelphis borealis), Pacific White- Sided Dolphin (Lagenorhynchus obliquidens), White-Beaked Dolphin (Lagenorhynchus albirostris), Harbour Porpoise (Phocoena phocoena), North Atlantic Right whale (Eubalaena glacialis), Sei Whale (Balaenoptera borealis), North Pacific Right Whale (Eubalaena japonica) Bowhead Whale Delphinidae -1.6 2 Low Brachycladiidae, Notocotylidae, NA 14, 18, 46, 49, (Balaena mysticetus) Phyllobothriidae, 103, 163, 222,

Polymorphidae, Tetrameridae 280, 281, 340 North Pacific Right Whale NA 3.0 17 Medium Cyamidae, Tetrabothriidae, Humpback Whale (Megeptera 18, 46, 49, 104, (Eubalaena japonica) Polymorphidae novaeangliae) 146, 147, 278, 279, 340 North Atlantic Right Whale NA 2.2 21.8 Medium Cyamidae Humpback Whale (Megeptera 18, 46, 49, 105, (Eubalaena glacialis) novaeangliae), Sei Whale (Balaenoptera 146, 192, 198, borealis) 340 Southern Right Whale Delphinidae 13.0 19 Medium Cyamidae, Hydrophilidae, Killer Whale (Orcinus orca), Peale's 46, 49, 104, (Eubalaena australis) Tetrabothriidae, Polymorphidae Dolphin (Lagenorhynchus australis), 106, 107, 146,

Indo-Pacific Humpbacked Dolphin 147, 190, 191, (Sousa chinensis) 202, 304, 340 Pygmy Right Whale Dalatiidae 5.0 20 Low NA Long-Finned Pilot Whale (Globicephala 46, 72, 73, 258, (Caperea marginata) melas), Sei Whale (Balaenoptera 340

borealis), Antarctic Minke Whale (Balaenoptera bonaerensis) Indo-pacific bottlenose dolphin Dalatiidae 13.4 24.1 High Anisakidae, Cyamidae, False Killer Whale (Pseudorca 17, 74, 153, (Tursiops aduncus) Pseudaliidae, Tetrameridae, crassidens), Bryde's Whale 155, 180, 190, Phyllobothriidae, Coronulidae (Balaenoptera edeni), Spinner Dolphin 258, 276, 277, (Stenella longirostris) 287, 340, 341

120

Species Predators Min Max Vocalizatio Parasites Species Known to Associate with References Temp Temp n Frequency (and sources (°C) (°C) cited within) Sperm Whale Delphinidae, Dalatiidae, 12.0 30.3 Medium Brachycladiidae, Pacific White-Sided Dolphin 16, 81, 327, (Physeter macrocephalus) Somniosidae Tentaculariidae, (Lagenorhynchus obliquidens), 332, 333, 334, Diphyllobothriidae, Northern Right Whale Dolphin 339 Tetrabothriidae, Anisakiidae, (Lissodelphis borealis), Risso's Dolphin Phyllobothriidae, Tetrameridae, (Grampus griseus), Melon-Headed Polymorphidae, Pennellidae, Whale (Peponocephala electra), Fraser's Coronulidae, Lepadidae, Dolphin (Lagenodelphis hosei), Short- Cyamidae, Echeneidae Finned Pilot Whale (Globicephala macrorhynchus), Southern Right Whale Dolphin (Lissodelphis peronii) Pygmy Sperm Whale Delphinidae, Lamnidae 26.9 30.9 Medium NA 289, 304, 319, (Kogia breviceps) 328, 330, 331 Dwarf Sperm Whale Delphinidae, Lamnidae 26.0 26.4 Medium Phyllobothriidae, Tetraphyllidea NA 16, 120, 289, (Kogia sima) incertae sedis, Anisakidae, 328, 329, 330, Tetrameridae, Pseudaliidae, 331, 332, 335 Polymorphidae, Pennellidae

References for Species 19, 25, 32, 36, 37, 42, 43, 45, 46, 48, 50, 51, 52, 55, 58, 59, 65, 73, 80, 82, 86, 87, 90, 92, 104, 122, 132, 161, 162, 168, 169, 170, 186, 205, 208, 210, Associations (and 211, 213, 215, 216, 220, 225, 226, 228, 229, 230, 231, 255, 256, 260, 263, 264, 267, 268, 269, 270, 272, 274, 286, 288, 291, 295, 296, 297, 298, 299, sources cited within) 304, 312, 327, 330

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S2.2 References for S2.1 1. Baker, A. N. 2001. Status, relationships, and distribution of Mesoplodon Bowdoini Andrews, 1908 (Cetacea: Ziphidae). Marine Mammal Science 17:473–493. 2. Hobson, R. P. and A. R. Martin. 1996. Behaviour and dive times of Arnoux’s beaked whales, Berardius arnuxii, at narrow leads in fast ice. Canadian Journal of Zoology 74:388–393. 3. Laporta, P., R. Praderi, V. Little and A. Le Bas. 2005. An Andrew’s beaked whale Mesoplodon bowdoini (Cetacea, Ziphiidae) stranded on the Atlantic Coast of Uruguay. Latin American Journal of Aquatic Mammals 4:101–111. 4. Besharse, J. C. 1971. Maturity and sexual dimorphism in the skull, mandible, and teeth of the Beaked Whale, Mesoplodon densirostris. Journal of Mammalogy 52:297–315. 5. Mead, J. G., W. A. Walker and W. J. Houck. 1982. Biological observations on Mesoplodon carlhubbsi (Ceatacea: Ziphiidae). Smithsonian Contributions to Zoology 344:1–25. 6. Mead, J. G. and A. N. Baker. 1987. Notes on the rare beaked whale, Mesoplodon hectori. Journal of the Royal Society of New Zealand 17:303–312. 7. Walker, W. A., J. G. Mead and R. L. Brownell Jr. 2002. Diets of Baird’s Beaked Whales, Berardius bairdii, in the Southern Sea of Okhotsk and off the Pacific coast of Honshu, Japan. Marine Mammal Science 18:902–919. 8. Reeves, R. R. and E. Mitchell. 1993. Status of Baird’s Beaked Whales, Berardius bairdii. Canadian field-naturalist 107:509–523. 9. Mead, J. G. 1984. Survey of reproductive data for the Beaked Whales (Ziphiidae). Report of the International Whaling Commission 91–96. 10. Santos, M. B., G. J. Pierce, J. Herman, A. López, A. Guerra, E. Mente and M. R. Clarke. 2001. Feeding ecology of Cuvier’s beaked whale (Ziphius cavirostris): a review with new information on the diet of this species. Journal of the Marine Biological Association of the United Kingdom 81:687–694. 11. McSweeney, D. J., R. W. Baird and S. D. Mahaffy. 2007. Site fidelity, associations, and movements of Cuvier’s (Ziphius cavirostris) and Blainville's (Mesoplodon densirostris) Beaked Whales off the Island of Hawai'i. Marine Mammal Science 23:666–687. 12. Macleod, C. D., W. F. Perrin, R. Pitman, J. Barlow, L. Ballance, A. D’Amico, T. Gerrodette, G. Joyce, K. D. Mullin, D. L. Palka and G. T. Waring. 2006. Known and inferred distributions of beaked whale species (Cetacea: Ziphiidae). Journal of Cetacean Research and Management 7:271–286. 13. Davis, R. W., G. S. Fargion, N. May, T. D. Leming, M. Baumgartner, W. E. Evans, L. J. Hansen and K. Mullin. 1998. Physical habitat of cetaceans along the continental slope in the north-central and western Gulf of Mexico. Marine Mammal Science 14:490–507. 14. Moore, S. E., D. P. Demaster and P. K. Dayton. 2000. Cetacean habitat selection in the Alaskan arctic during summer and autumn. Arctic 53:432–447. 15. López, A., G. J. Pierce, X. Valeiras, M. B. Santos and A. Guerra. 2004. Distribution patterns of small cetaceans in Galician waters. Journal of the Marine Biological Association of the United Kingdom 84:283–294. 16. Ballance, L. T. and R. L. Pitman. 1998. Cetaceans of the western tropical Indian Ocean: distribution, relative abundance, and comparisons with cetacean communities of two other tropical ecosystems. Marine Mammal Science 14:429–459. 17. Hoelzel, A. R. (ed). 2002. Marine mammal biology: an evolutionary approach. Blackwell Publishing, Malden, MA. 18. Jefferson, T. A., M. A. Webber and R. L. Pitman. 2008. Marine Mammals of the World: a comprehensive guide to their identification. Elsevier, San Diego, CA.

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19. Wolman, A. A. 1985. Gray Whale Eschrichtius robustus (Lilljeborg, 1861). Pp. 67–90 in S. H. Ridgway and R. Harrison, eds. Handbook of Marine Mammals: Volume 3 The sirenians and baleen whales. Academic Press Inc., Toronto, ON. 20. Mullin, K. D. and G. L. Fulling. 2004. Abundance of cetaceans in the oceanic northern Gulf of Mexico, 1996- 2001. 20:787–807. 21. Simard, P., J. L. Lawlor and S. Gowans. 2006. Temporal variability of cetaceans near Halifax, Nova Scotia. Canadian field-naturalist 120:93–99. 22. Azevedo, A. F., S. C. Viana, A. M. Oliveira and M. V. Sluys. 2005. Group characteristics of marine tucuxis (Sotalia fluviatilis) (Cetacea: Delphinidae) in Guanabara Bay, south-eastern Brazil. Journal of the Marine Biological Association of the United Kingdom 85:209–212. 23. MacLeod, C. D., C. R. Weir, M. B. Santos and T. E. Dunn. 2008. Temperature-based summer habitat partitioning between white-beaked and common dolphins around the United Kingdom and Republic of Ireland. Journal of the Marine Biological Association of the United Kingdom 88:1193 – 1198. 24. Cañadas, A., R. Sagarminaga and S. García-Tiscar. 2003. Cetacean distribution related with depth and slope in the Mediterranean waters off southern Spain. Deep-Sea Research I 49:2053–2073. 25. Van Waerebeek, K., L. Barnett, A. Camara, A. Cham, M. Diallo, A. Djiba, A. O. Jallow, E. Ndiaye, A. S. O. O. Bilal and I. L. Bamy. 2004. Distribution, status, and biology of the Atlantic Humpback Dolphin, Sousa teuszii (Kükenthal, 1892). Aquatic Mammals 30:56–83. 26. Weinrich, M. T., C. R. Belt and D. Morin. 2001. Behavior and ecology of the Atlantic White-sided dolphin (Lagenorhynchus acutus) in coastal New England waters. Marine Mammal Science 17:231–248. 27. Selzer, L. A. and P. M. Payne. 1988. The distribution of White-sided (Lagenorhychus acutus) and Common Dolphins (Delphinus delphis) vs. environmental features of the continental shelf of the Northeastern United States. Marine Mammal Science 4:141–153. 28. Jefferson, T. A., K. M. Robertson and J. Y. Wang. 2002. Growth and reproduction of the finless porpoise in southern China. The Raffles Bulletin of Zoology 10:105–113. 29. Barros, N. B., T. A. Jefferson and E. C. M. Parsons. 2002. Food habits of Finless Porpoises (Neophocaena phocaenoides) in Hong Kong waters. The Raffles Bulletin of Zoology 10:115–123. 30. Shirakihara, K., M. Shirakihara and Y. Yamamoto. 2007. Distribution and abundance of finless porpoise in the Inland Sea of Japan. Marine Biology 150:1025–1032. 31. Best, P. B. and R. B. Abernethy. 1994. Heaviside’s Dolphin Cephalorhynchus heavisidii (Gray, 1828). Pp. 289–310 in S. H. Ridgway and R. Harrison, eds. Handbook of Marine Mammals: Volume 5 The first book of dolphins. Academic Press Inc., San Diego, CA. 32. Brownell Jr., R. L. and P. J. Clapham. 1999. Spectacled Porpoise Phocoena dioptrica Lahille, 1912. Pp. 379–391 in S. H. Ridgway and R. Harrison, eds. Handbook of Marine Mammals: Volume 6 The second book of dolphins and the porpoises. Academic Press Inc., Toronto, ON. 33. Crespo, E. A. and R. González. 1998. Group size and distributional range of the Franciscana, Pontoporia blainvillei. Marine Mammal Science 14:845–849. 34. Dalebout, M. L., G. J. B. Ross, C. S. Baker, R. C. Anderson, P. B. Best, V. G. Cockcroft, H. L. Hinsz, V. Peddemors and R. L. Pitman. 2003. Appearance, distribution and genetic distinctiveness of Longman’s Beaked Whale, Indopacetus pacificus. Marine Mammal Science 19:421–461. 35. Newcomer, M. W., T. A. Jefferson and R. L. Brownell Jr. 1996. Lissodelphis peroni. Mammalian Species 531:1–5.

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36. Jefferson, T. A., M. W. Newcomer, S. Leatherwood and K. Van Waerebeek. 1994. Right Whale Dolphins Lissodelphis borealis (Peale, 1848) and Lissodelphis peronii (Lacépède, 1804). Pp. 335–362 in S. H. Ridgway and R. Harrison, eds. Handbook of Marine Mammals: Volume 5 The first book of dolphins. Academic Press Inc., Toronto, ON. 37. Jefferson, T. A. and L. Karczmarski. 2001. Sousa chinensis. Mammalian Species 1–9. 38. Perrin, W. F., E. D. Mitchell, J. G. Mead, D. K. Caldwell, M. C. Caldwell, P. J. H. van Bree and W. H. Dawbin. 1987. Revision of the Spotted Dolphins, Stenella spp. Marine Mammal Science 3:99–170. 39. Perrin, W. F. and A. A. Hohn. 1994. Pantropical Spotted Dolphin Stenella attenuata. Pp. 71–98 in S. H. Ridgway and R. Harrison, eds. Handbook of Marine Mammals: Volume 5 The first book of dolphins. Academic Press Inc., San Diego, CA. 40. Stacey, P. J. and S. Leatherwood. 1997. The Irrawaddy Dolphin, Orcaella brevirostris: A summary of current knowledge and recommendations for conservation action. Asian Marine Biology 14:195–214. 41. Baird, R. W. 2002. False Killer Whale, Pseudorca crassidens. Pp. 405–406 in W. F. Perrin, B. Würsig, and J. G. M. Thewissen, eds. Encyclopedia of Marine Mammals. Academic Press Inc., San Francisco, CA. 42. Stacey, P. J., S. Leatherwood and R. W. Baird. 1994. Pseudorca crassidens. Mammalian Species 1–6. 43. Pereira, J. N. D. S. G. 2008. Field notes on Risso’s Dolphin (Grampus griseus) distribution, social ecology, behaviour, and occurrence in the Azores. Aquatic Mammals 34:426–435. 44. Chen, I., A. Watson and L.-S. Chou. 2011. Insights from life history traits of Risso’s dolphins (Grampus griseus) in Taiwanese waters: Shorter body length characterizes northwest Pacific population. Marine Mammal Science 27:43–64. 45. Baird, R. W. and P. J. Stacey. 1991. Status of Risso’s Dolphin, Grampus griseus, in Canada. Canadian field- naturalist 105:233–242. 46. Klinowska, M. 1991. Dolphins, Porpoises and Whales of the World: The IUCN Red Data Book. IUCN, Gland, Switzerland and Cambridge, U.K. 47. Marsh, H., R. Lloze, G. E. Heinsohn and T. Kasuya. 1989. Irrawaddy Dolphin Orcaella brevirostris (Gray, 1866). Pp. 101–118 in S. H. Ridgway and R. Harrison, eds. Handbook of Marine Mammals: Volume 4 River dolphins and the larger toothed whales. Academic Press Inc., San Diego, CA. 48. Ross, G. J. B. and S. Leatherwood. 1994. Pygmy Killer Whale Feresa Attenuata Gray 1874. Pp. 387–404 in S. H. Ridgway and R. Harrison, eds. Handbook of Marine Mammals: Volume 5 The first book of dolphins. Academic Press Inc., San Diego, CA. 49. Gaskin, D. E. 1982. The Ecology of Whales and Dolphins. Heinemann, Exeter, NH. 50. Reeves, R. R., C. Smeenk, C. C. Kinze, R. L. Brownell Jr. and J. Lien. 1999. White-beaked Dolphin Lagenorhynchus albirostris Gray, 1846. Pp. 1–30 in S. H. Ridgway and R. Harrison, eds. Handbook of Marine Mammals: Volume 6 The second book of dolphins and the porpoises. Academic Press Inc., Toronto, ON. 51. Reeves, R. R., C. Smeenk, R. L. Brownell Jr. and C. C. Kinze. 1999. Atlantic White-sided Dolphin. Pp. 31–56 in S. H. Ridgway and R. Harrison, eds. Handbook of Marine Mammals: Volume 6 The second book of dolphins and the porpoises. Academic Press Inc., Toronto, ON. 52. Brownell Jr., R. L., W. A. Walker and K. A. Forney. 1999. Pacific White-sided Dolphin Lagenorhynchus obliquidens Gill, 1865. Pp. 57–84 in S. H. Ridgway and R. Harrison, eds. Handbook of Marine Mammals: Volume 6 The second book of dolphins and the porpoises. Academic Press Inc., Toronto, ON.

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53. Dahlheim, M. E. and R. G. Towell. 1994. Occurrence and distribution of Pacific White-sided Dolphins (Lagenorhynchus obliquidens) in southeastern Alaska, with notes on an attack by Killer Whales (Orcinus orca). Marine Mammal Science 10:458–464. 54. Morton, A. B. 2000. Occurence, photo-identification and prey of Pacific White-sided Dolphins (Lagenorhynchus obliquidens) in the Broughton Archipelago, Canada 1984-1998. Marine Mammal Science 16:80–93. 55. Brownell Jr., R. L., E. A. Crespo and M. A. Donahue. 1999. Peale’s Dolphin Lagenorhynchus australis (Peale, 1848). Pp. 105–120 in S. H. Ridgway and R. Harrison, eds. Handbook of Marine Mammals: Volume 6 The second book of dolphins and the porpoises. Academic Press Inc., Toronto, ON. 56. Viddi, F. A. and A.-K. Lescrauwaet. 2005. Insights on habitat selection and behavioural patterns of Peale’s Dolphins (Lagenorhynchus australis) in the Strait of Magellan, Southern Chile. Aquatic Mammals 31:176–183. 57. Schiavini, A. C. M., R. N. P. Goodall, A.-K. Lescrauwaet and M. Koen Alonso. 1997. Food habits of the Peale’s dolphin, Lagenorhynchus australis: review and new information. Report of the International Whaling Commission 0:827–834. 58. Brownell Jr., R. L. and M. A. Donahue. 1999. Hourglass Dolphin Lagenorhynchus cruciger (Quoy and Gaimard, 1824). Pp. 121–136 in S. H. Ridgway and R. Harrison, eds. Handbook of Marine Mammals: Volume 6 The second book of dolphins and the porpoises. Academic Press Inc., Toronto, ON. 59. Kruse, S., D. K. Caldwell and M. C. Caldwell. 1999. Risso’s Dolphin Grampus griseus (G. Cuvier, 1812). Pp. 183– 212 in S. H. Ridgway and R. Harrison, eds. Handbook of Marine Mammals: Volume 6 The second book of dolphins and the porpoises. Academic Press Inc., Toronto, ON. 60. Ribeiro, S., F. A. Viddi, J. L. Cordeiro and T. R. O. Freitas. 2007. Fine-scale habitat selection of Chilean dolphins (Cephalorhynchus eutropia): interactions with aquaculture activities in southern Chiloé Island, Chile. Journal of the Marine Biological Association of the UK 87:119–128. 61. Jefferson, T. A., D. K. Odell and K. T. Prunier. 1995. Notes on the biology of the Clymene dolphin (Stenella clymene) in the Northern Gulf of Mexico. Marine Mammal Science 11:564–573. 62. Fertl, D., T. A. Jefferson, I. B. Moreno, A. N. Zerbini and K. D. Mullin. 2003. Distribution of the Clymene dolphin Stenella clymene. Mammal Review 33:253–271. 63. Bryden, M. M., R. J. Harrison and R. J. Lear. 1977. Some aspects of the biology of Peponocephala electra (Cetacea: Delphinidae) I. General and reproductive biology. Australian Journal of Marine and Freshwater Resources 28:703–715. 64. Cannon, L. R. G. 1977. Some aspects of the biology of Peponocephala electra (Cetacea: Delphinidae) II.* Parasites. Australian Journal of Marine and Freshwater Resources 28:717–722. 65. Watkins, W. A., M. A. Daher, A. Samuels and D. P. Gannon. 1997. Observations of Peponocephala electra, the Melon-headed Whale, in the Southeastern Caribbean. Caribbean Journal of Science 33:34–40. 66. Brownell Jr., R. L., K. Ralls, S. Baumann-Pickering and M. M. Poole. 2009. Behavior of melon-headed whales, Peponocephala electra, near oceanic islands. Marine Mammal Science 25:639–658. 67. Sekiguchi, K., N. T. W. Klages and P. B. Best. 1996. The diet of strap-toothed whales (Mesoplodon layardii). Journal of Zoology of London 239:453–463. 68. Baker, A. N. and A. L. van Helden. 1999. New records of beaked whales, genus Mesoplodon, from New Zealand (Cetacea: Ziphiidae). Journal of the Royal Society of New Zealand 29:235–244. 69. Mead, J. G. 1989. Beaked Whales of the Genus Mesoplodon. Pp. 349–430 in S. H. Ridgway and R. Harrison, eds. Handbook of Marine Mammals: Volume 4 River dolphins and the larger toothed whales. Academic Press Inc., San Diego, CA.

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70. Slip, D. J., G. J. Moore and K. Green. 1995. Stomach contents of a Southern Bottlenose Whale, Hyperoodon planifrons, stranded at Heard Island. Marine Mammal Science 11:575–584. 71. Macleod, C. D., M. B. Santos and G. J. Pierce. 2003. Review of data on diets of beaked whales: evidence of niche separation and geographic segregation. Journal of the Marine Biological Association of the United Kingdom 83:651–665. 72. Sekiguchi, K. and B. Z. Kaczmaruk. 1992. On the feeding habits and baleen morphology of the Pygmy Right Whale Caperea marginata. Marine Mammal Science 8:288–293. 73. Baker, A. N. 1985. Pygmy Right Whale Caperea marginata (Gray, 1846). Pp. 345–354 in S. H. Ridgway and R. Harrison, eds. Handbook of Marine Mammals: Volume 3 The sirenians and baleen whales. Academic Press Inc., Toronto, ON. 74. Möller, L. M., S. J. Allen and R. G. Harcourt. 2002. Group characteristics, site fidelity and seasonal abundance of Bottlenose Dolphins Tursiops aduncus in Jervis Bay and Port Stephens, South-Eastern Australia. Australian Mammalogy 24:11–21. 75. Börjesson, P., P. Berggren and B. Ganning. 2003. Diet of Harbor Porpoises in the Kattegat and Skagerrak Seas: accounting for individual variation and sample size. Marine Mammal Science 19:38–58. 76. Gannon, D. P., J. E. Craddock and A. J. Read. 1998. Autumn food habits of Harbor Porpoises, Phocoena phocoena, in the Gulf of Maine. Fisheries Bulletin 96:428–437. 77. Raum-Suryan, K. L. and J. T. Harvey. 1998. Distribution and abundance of habitat use by Harbour Porpoise, Phocoena phocoena, off the northern San Juan Islands, Washington. Fisheries Science 96:808–822. 78. Tamura, T. and K. Konishi. 2009. Feeding habits and prey consumption of Antarctic Minke Whale (Balaenoptera bonaerensis) in the southern ocean. Journal of Northwest Atlantic Fishery Science 42:13–25. 79. Heyning, J. E. 1989. Cuvier’s Beaked Whale Ziphius cavirostris G. Cuvier, 1823. Pp. 289–308 in S. H. Ridgway and R. Harrison, eds. Handbook of Marine Mammals: Volume 4 River dolphins and the larger toothed whales. Academic Press Inc., Toronto, ON. 80. Anderson, R. C., R. Clark, P. T. Madsen, C. Johnson, J. Kiszka and O. Breysse. 2006. Observations of Longman’s Beaked Whale (Indopacetus pacificus) in the Western Indian Ocean. Aquatic Mammals 32:223–231. 81. Gaskin, D. E. 1972. Whales, Dolphins and Seals. The MacMillan Company of Canada Limited, Toronto, ON. 82. Brownell Jr., R. L. and F. Cipriano. 1999. Dusky Dolphin Lagenorhynchus obscurus (Gray, 1828). Pp. 85–104 in S. H. Ridgway and R. Harrison, eds. Handbook of Marine Mammals: Volume 6 The second book of dolphins and the porpoises. Academic Press Inc., Toronto, ON. 83. Blanco, C., M. Á. Raduán and J. A. Raga. 2006. Diet of Risso’s dolphin (Grampus griseus) in the western Mediterranean Sea. Scientia Marina 70:407–411. 84. Odell, D. K. and K. M. McClune. 1999. False Killer Whale Pseudorca crassidens (Owen, 1846). Pp. 213–243 in S. H. Ridgway and R. Harrison, eds. Handbook of Marine Mammals: Volume 6 The second book of dolphins and the porpoises. Academic Press Inc., Toronto, ON. 85. Bernard, H. J. and S. B. Reilly. 1999. Pilot Whales Globicephala Lesson, 1828. Pp. 245–279 in S. H. Ridgway and R. Harrison, eds. Handbook of Marine Mammals: Volume 6 The second book of dolphins and the porpoises. Academic Press Inc., Toronto, ON. 86. Brownell Jr., R. L. and P. J. Clapham. 1999. Burmeister’s Porpoise Phocoena spinipinnis Burmeister, 1865. Pp. 393–410 in S. H. Ridgway and R. Harrison, eds. Handbook of Marine Mammals: Volume 6 The second book of dolphins and the porpoises. Academic Press Inc., Toronto, ON.

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87. Kasuya, T. 1999. Finless Porpoise Neophocaena phocaenoides (G. Cuvier, 1829). Pp. 411–442 in S. H. Ridgway and R. Harrison, eds. Handbook of Marine Mammals: Volume 6 The second book of dolphins and the porpoises. Academic Press Inc., Toronto, ON. 88. Jefferson, T. A. 1991. Observations on the distribution and behaviour of Dall’s porpoise (Phocoenoides dalli) in Monterey Bay, California. Aquatic Mammals 17:12–19. 89. Perrin, W. F., K. M. Robertson and W. A. Walker. 2008. Diet of the Striped Dolphin, Stenella coeruleoalba, in the Eastern tropical Pacific Ocean. 90. Miyazaki, N. and W. F. Perrin. 1994. Rough-toothed dolphin Steno bradanensis (Lesson, 1828). Pp. 1–21 in S. H. Ridgway and R. Harrison, eds. Handbook of Marine Mammals: Volume 5 The first book of dolphins. Academic Press Inc., San Diego, CA. 91. da Silva, V. M. F. and R. C. Best. 1994. Tucuxi Sotalia fluviatilis (Gervais, 1853). Pp. 43–69 in S. H. Ridgway and R. Harrison, eds. Handbook of Marine Mammals: Volume 5 The first book of dolphins. Academic Press Inc., San Diego, CA. 92. Perrin, W. F. and J. W. Gilpatrick Jr. 1994. Spinner Dolphin Stenella longirostris (Gray, 1828). Pp. 99–128 in S. H. Ridgway and R. Harrison, eds. Handbook of Marine Mammals: Volume 5 The first book of dolphins. San Diego, CA. 93. Perrin, W. F., C. E. Wilson and F. I. Archer II. 1994. Striped Dolphin Stenella coeruleoalba (Meyen, 1833). Pp. 129–159 in S. H. Ridgway and R. Harrison, eds. Handbook of Marine Mammals: Volume 5 The first book of dolphins. Academic Press Inc., San Diego, CA. 94. Perrin, W. F., D. K. Caldwell and M. C. Caldwell. 1994. Atlantic Spotted Dolphin Stenella frontalis (G. Cuvier, 1829). Pp. 173–190 in S. H. Ridgway and R. Harrison, eds. Handbook of Marine Mammals: Volume 5 The first book of dolphins. Academic Press Inc., San Diego, CA. 95. Evans, W. E. 1994. Common Dolphin, White-bellied Porpoise Delphinus delphis Linnaeus, 1758. Pp. 191–224 in S. H. Ridgway and R. Harrison, eds. Handbook of Marine Mammals: Volume 5 The first book of dolphins. Academic Press Inc., San Diego, CA. 96. Perrin, W. F., S. Leatherwood and A. Collet. 1994. Fraser’s Dolphin Lagenodelphis hosei Fraser, 1956. Pp. 225–240 in S. H. Ridgway and R. Harrison, eds. Handbook of Marine Mammals: Volume 5 The first book of dolphins. Academic Press Inc., San Diego, CA. 97. Slooten, E. and S. M. Dawson. 1994. Hector’s Dolphin. Cephalorhynchus hectori Pp. 311–333 in S. H. Ridgway and R. Harrison, eds. Handbook of Marine Mammals: Volume 5 The first book of dolphins. Academic Press Inc., San Diego, CA. 98. Brownell Jr., R. L. 1989. Franciscana Pontoporia blainvillei (Gervais and d’Orbigny, 1844). Pp. 45–67 in S. H. Ridgway and R. Harrison, eds. Handbook of Marine Mammals: Volume 4 River dolphins and the larger toothed whales. Academic Press Inc., San Diego, CA. 99. Brodie, P. F. 1989. The White Whale Delphinapterus leucas (Pallas, 1776). Pp. 119–144 in S. H. Ridgway and R. Harrison, eds. Handbook of Marine Mammals: Volume 4 River dolphins and the larger toothed whales. Academic Press Inc., San Diego, CA. 100. Hay, K. A. and A. W. Mansfield. 1989. Narwhal Monodon monoceros Linnaeus, 1758. Pp. 145–176 in S. H. Ridgway and R. Harrison, eds. Handbook of Marine Mammals: Volume 4 River dolphins and the larger toothed whales. Academic Press Inc., San Diego, CA.

127

101. Balcomb III, K. C. 1989. Baird’s Beaked Whale Berardius bairdii Stejneger, 1883: Arnoux's Beaked Whales Berardius arnuxii Duvernoy 1851. Pp. 261–288 in S. H. Ridgway and R. Harrison, eds. Handbook of Marine Mammals: Volume 4 River dolphins and the larger toothed whales. Academic Press Inc., San Diego, CA. 102. Mead, J. G. 1989. Shepherd’s Beaked Whale Tasmacetus sheperdi Oliver, 1937. Pp. 309–320 in S. H. Ridgway and R. Harrison, eds. Handbook of Marine Mammals: Volume 4 River dolphins and the larger toothed whales. Academic Press Inc., San Diego, CA. 103. Dailey, M. D. and R. L. Brownell Jr. 1972. A checklist of marine mammal parasites. Pp. 528–589 in S. H. Ridgway, ed. Mammals of the Sea. Charles C. Thomas, Springfield, IL. 104. Scarff, J. E. 1986. Historic and present distribution of the right whale (Eubalaena glacialis) in the Eastern North Pacific south of 50ON and east of 180OW. Reports of the International Whaling Commission 43–63. 105. Winn, H. E., C. A. Price and P. W. Sorensen. 1986. The distributional biology of the Right Whale (Eubalaena glacialis) in the Western North Atlantic. Reports of the International Whaling Commission 129–138. 106. Aguayo L., A. and D. Torres N. 1986. Records of the Southern Right Whale, Eubalaena australis (Desmoulins, 1822) from Chile between 1976 and 1982. Reports of the International Whaling Commission 159–167. 107. Ohsumi, S. and F. Kasamatsu. 1986. Recent off-shore distribution of the Southern Right Whale in summer. Reports of the International Whaling Commission 177–185. 108. Goodall, R. N. P., A. R. Galeazzi, S. Leatherwood, K. W. Miller, I. S. Cameron, R. K. Kastelein and A. P. Sobral. 1988. Studies of Commerson’s Dolphins, Cepahlorhynchus commersonii, off Tiera del Fuego, 1979-1984, with a review of information on the species in the South Atlantic. Reports of the International Whaling Commission 3–70. 109. Leatherwood, S., R. A. Kastelein and K. W. Miller. 1988. Observations of Commerson’s Dolphin and other cetaceans in Southern Chile, January-February 1984. Reports of the International Whaling Commission 71– 83. 110. Bastida, R., V. Lichtschein and R. N. P. Goodall. 1988. Food habits of Cephalorhynchus commersonii off Tiera del Fuego. Reports of the International Whaling Commission 143–160. 111. Goodall, R. N. P., K. S. Norris, A. R. Galeazzi, J. A. Oporto and I. S. Cameron. 1988. On the Chilean Dolphin, Cephalorhynchus eutropia (Gray, 1846). Reports of the International Whaling Commission 197–257. 112. Dawson, S. M. and E. Slooten. 1988. Hector’s Dolphin, Cephalorhynchus hectori: distribution and abundance. Reports of the International Whaling Commission 315–338. 113. Vidal, O. 1995. Population biology and incidental mortality of the Vaquita, Phocoena sinus. Reports of the International Whaling Commission 247–273. 114. Goodall, R. N. P., K. S. Norris, G. Harris, J. A. Oporto and H. P. Castello. 1995. Notes on the biology of the Burmeister’s Porpoise, Phocoena spinipinnis, off Southern South America. Reports of the International Whaling Commission 317–347. 115. Desportes, G. and R. Mouritsen. 1993. Preliminary results on the diet of Long-finned Pilot Whales off the Faroe Islands. Reports of the International Whaling Commission 305–324. 116. Kasuya, T. and S. Tai. 1993. Life history of Short-finned Pilot Whale stocks off Japan and a description of the fishery. Reports of the International Whaling Commission 439–473. 117. Mintzer, V. J., D. P. Gannon, N. B. Barros and A. J. Read. 2008. Stomach contents of mass-stranded Short- finned Pilot Whales (Globicephala macrorhynchus) from North Carolina. Marine Mammal Science 24:290–302.

128

118. Taylor, B. L., R. Baird, S. M. Dawson, J. Ford, J. G. Mead, G. Notarbartolo do Sciara, P. Wade and R. L. Pitman. 2011. Globicephala macrorhynchus. IUCN 2011. IUCN Red List of Threatened Speces. Version 2011.2. [Date accessed: February 28, 2012] 119. Kleinenberg, S. E., A. V. Yablokov, B. M. Bel’kovich and M. N. Tarasevich. 1969. Investigation of the species Beluga. Translated. Smithsonian Institute. 120. Fitch, J. E. and R. L. Brownell Jr. 1968. Fish otoliths in cetacean stomachs and their importance in interpreting feeding habits. Journal of the Fisheries Research Board of Canada 25:2561–2574. 121. Dolar, M. L. L., W. A. Walker, G. L. Kooyman and W. F. Perrin. 2003. Comparative feeding ecology of Spinner Dolphins (Stenella longirostris) and Fraser’s Dolphins (Lagenodelphis hosei) in the Sulu Sea. Marine Mammal Science 19:1–19. 122. West, K. L., J. G. Mead and W. White. 2011. Steno bredanensis (Cetacea: Delphinidae). Mammalian Species 43:177–189. 123. Mead, J. G. 2002. Shepherd’s Beaked Whale, Tasmacetus shepherdi. Pp. 1078–1081 in W. F. Perrin, B. Würsig, and J. G. M. Thewissen, eds. Encyclopedia of Marine Mammals. Academic Press Inc., San Francisco, CA. 124. Perrin, W. F. 2002. Common Dolphins, Delphinus delphis, D. capensis, and D. tropicalis. Pp. 245–248 in W. F. Perrin, B. Würsig, and J. G. M. Thewissen, eds. Encyclopedia of Marine Mammals. Academic Press Inc., San Francisco, CA. 125. Sears, R. 2002. Blue Whale, Balaenoptera musculus. Pp. 112–116 in W. F. Perrin, B. Würsig, and J. G. M. Thewissen, eds. Encyclopedia of Marine Mammals. Academic Press Inc., San Francisco, CA. 126. Gowans, S. 2002. Bottlenose Whales, Hyperoodon ampullatus and H. planifrons. Pp. 128–129 in W. F. Perrin, B. Würsig, and J. G. M. Thewissen, eds. Encyclopedia of Marine Mammals. Academic Press Inc., San Francisco, CA. 127. Rugh, D. J. and K. E. W. Shelden. 2002. Bowhead Whale, Balaena mysticetus. Pp. 129–131 in W. F. Perrin, B. Würsig, and J. G. M. Thewissen, eds. Encyclopedia of Marine Mammals. Academic Press Inc., San Francisco, CA. 128. Kato, H. 2002. Bryde’s Whales, Balaenoptera edeni and B. brydei. Pp. 171–177 in W. F. Perrin, B. Würsig, and J. G. M. Thewissen, eds. Encyclopedia of Marine Mammals. Academic Press Inc., San Francisco, CA. 129. Dawson, S. M. 2002. Cephalorhynchus Dolphins, C. heavisidii, C. eutropia, C. hectori, and C. commersonii. Pp. 200– 203 in W. F. Perrin, B. Würsig, and J. G. M. Thewissen, eds. Encyclopedia of Marine Mammals. Academic Press Inc., San Francisco, CA. 130. Jefferson, T. A. 2002. Clymene Dolphin, Stenella clymene. Pp. 234–236 in W. F. Perrin, B. Würsig, and J. G. M. Thewissen, eds. Encyclopedia of Marine Mammals. Academic Press Inc., San Francisco, CA. 131. Heyning, J. E. 2002. Cuvier’s Beaked Whale, Ziphius cavirostris. Pp. 305–307 in W. F. Perrin, B. Würsig, and J. G. M. Thewissen, eds. Encyclopedia of Marine Mammals. Academic Press Inc., San Francisco, CA. 132. Jefferson, T. A. and S. Leatherwood. 1994. Lagenodelphis hosei. Mammalian Species 470:1–5. 133. Hammond, P. S., G. Bearzi, A. Bjøge, K. Forney, L. Karczmarski, T. Kasuya, W. F. Perrin, M. D. Scott, J. Y. Wang, R. S. Wells and B. Wilson. 2008. Lagenodelphis hosei. IUCN 2011. IUCN Red List of Threatened Speces. Version 2011.2. [Date accessed: February 28, 2012] 134. Dolar, M. L. L. 2002. Fraser’s Dolphin, Lagenodelphis hosei. Pp. 485–487 in W. F. Perrin, B. Würsig, and J. G. M. Thewissen, eds. Encyclopedia of Marine Mammals. Academic Press Inc., San Francisco, CA. 135. Jones, M. L. and S. L. Swartz. 2002. Gray Whale, Eschrichtius robustus. Pp. 524–536 in W. F. Perrin, B. Würsig, and J. G. M. Thewissen, eds. Encyclopedia of Marine Mammals. Academic Press Inc., San Francisco, CA.

129

136. Goodall, R. N. P. 2002. Hourglass Dolphin, Lagenorhynchus cruciger. Pp. 583–585 in W. F. Perrin, B. Würsig, and J. G. M. Thewissen, eds. Encyclopedia of Marine Mammals. Academic Press Inc., San Francisco, CA. 137. Ross, G. J. B. 2002. Humpback Dolphins, Sousa chinensis, S. plumbea, and S. teuszi. Pp. 585–589 in W. F. Perrin, B. Würsig, and J. G. M. Thewissen, eds. Encyclopedia of Marine Mammals. Academic Press Inc., San Francisco, CA. 138. Clapham, P. J. 2002. Humpback Whale, Megaptera novaeangliae. Pp. 589–592 in W. F. Perrin, B. Würsig, and J. G. M. Thewissen, eds. Encyclopedia of Marine Mammals. Academic Press Inc., San Francisco, CA. 139. Arnold, P. W. 2002. Irrawaddy Dolphin, Orcaella brevirostris. Pp. 652–654 in W. F. Perrin, B. Würsig, and J. G. M. Thewissen, eds. Encyclopedia of Marine Mammals. Academic Press Inc., San Francisco, CA. 140. Perrin, W. F. and R. L. Brownell Jr. 2002. Minke Whales, Balaenoptera acutorostrata and B. bonaerensis. Pp. 750– 754 in W. F. Perrin, B. Würsig, and J. G. M. Thewissen, eds. Encyclopedia of Marine Mammals. Academic Press Inc., San Francisco, CA. 141. Stewart, B. S. and S. Leatherwood. 1985. Minke Whale Balaenoptera acutorostrata Lacépède, 1804. Pp. 91–136 in S. H. Ridgway and R. Harrison, eds. Handbook of Marine Mammals: Volume 3 The sirenians and baleen whales. Academic Press Inc., Toronto, ON. 142. Dailey, M. D. and W. K. Vogelbein. 1991. Parasite fauna of three species of Antarctic whales with reference to their use as potential stock indicators. Fishery Bulletin 89:355–365. 143. Gibson, D. I., E. A. Harris, R. A. Bray, P. D. Jepson, T. Kuiken, J. R. Baker and V. R. Simpson. 1998. A survey of the helminth parasites of cetaceans stranded on the coast of England and Wales during the period 1990 ± 1994. Journal of Zoology of London 244:563–574. 144. Akihiko, U. and A. Jun. 2000. The ectoparasties and endoparasites in the Minke Whale, Balaenoptera acutorostrata from the Western North Pacific Ocean. Journal of the Japan Veterinary Medical Association 53:85–88. 145. Heide-Jørgensen, M. P. 2002. Narwhal, Monodon monoceros. Pp. 783–787 in W. F. Perrin, B. Würsig, and J. G. M. Thewissen, eds. Encyclopedia of Marine Mammals. Academic Press Inc., San Francisco, CA. 146. Cummings, W. C. 1985. Right Whales Eubalaena glacialis (Müller, 1776) and Eubalaena australis (Desmoulins, 1822). Pp. 275–304 in S. H. Ridgway and R. Harrison, eds. Handbook of Marine Mammals: Volume 3 The sirenians and baleen whales. Academic Press Inc., Toronto, ON. 147. Klumov, S. K. 1962. The Right Whales in the pacific ocean. Proceedings of the Institute of Oceanology 58:202– 297. 148. Baird, R. W. 2002. Risso’s Dolphin, Grampus griseus. Pp. 1037–1039 in W. F. Perrin, B. Würsig, and J. G. M. Thewissen, eds. Encyclopedia of Marine Mammals. Academic Press Inc., San Francisco, CA. 149. Ralls, K. and S. L. Mesnick. 2002. Sexual Dimorphism. Pp. 1071–1078 in W. F. Perrin, B. Würsig, and J. G. M. Thewissen, eds. Encyclopedia of Marine Mammals. Academic Press Inc., San Francisco, CA. 150. Archer II, F. I. 2002. Striped Dolphin, Stenella coeruleoalba. Pp. 1201–1203 in W. F. Perrin, B. Würsig, and J. G. M. Thewissen, eds. Encyclopedia of Marine Mammals. Academic Press Inc., San Francisco, CA. 151. Hammond, P. S., G. Bearzi, A. Bjøge, K. Forney, L. Karczmarski, T. Kasuya, W. F. Perrin, M. D. Scott, J. Y. Wang, R. S. Wells and B. Wilson. 2008. Stenella coeruleoalba. IUCN 2011. IUCN Red List of Threatened Speces. Version 2011.2. [Date accessed: February 28, 2012] 152. Kinze, C. C. 2002. White-beaked Dolphin, Lagenorhynchus albirostris. Pp. 1332–1334 in W. F. Perrin, B. Würsig, and J. G. M. Thewissen, eds. Encyclopedia of Marine Mammals. Academic Press Inc., San Francisco, CA.

130

153. Amir, O. A., P. Berggren, S. G. M. Ndaro and N. S. Jiddawi. 2005. Feeding ecology of the Indo-Pacific bottlenose dolphin (Tursiops aduncus) incidentally caught in the gillnet fisheries off Zanzibar, Tanzania. Estuarine, Coastal and Shelf Science 63:429–437. 154. Hammond, P. S., G. Bearzi, A. Bjøge, K. Forney, L. Karczmarski, T. Kasuya, W. F. Perrin, M. D. Scott, J. Y. Wang, R. S. Wells and B. Wilson. 2008. Tursiops truncatus. IUCN 2011. IUCN Red List of Threatened Speces. Version 2011.2. [Date accessed: February 28, 2012] 155. Hammond, P. S., G. Bearzi, A. Bjøge, K. Forney, L. Karczmarski, T. Kasuya, W. F. Perrin, M. D. Scott, J. Y. Wang, R. S. Wells and B. Wilson. 2008. Tursiops aduncus. IUCN 2011. IUCN Red List of Threatened Speces. Version 2011.2. [Date accessed: February 28, 2012] 156. García-Godos, I., K. Van Waerebeek, J. C. Reyes, J. Alfaro-Shigueto and M. Arias-Schreiber. 2007. Prey occurrence in the stomach contents of four small cetacean species in Peru. Latin American Journal of Aquatic Mammals 6:171–183. 157. Ballance, L. T. 1992. Habitat use patterns and ranges of the Bottlenose Dolphin in the Gulf of California, Mexico. Marine Mammal Science 8:262–274. 158. Blanco, C., O. Salomón and J. A. Raga. 2001. Diet of the bottlenose dolphin (Tursiops truncatus) in the western Mediterranean Sea. Journal of the Marine Biological Association of the United Kingdom 81:1053–1058. 159. Barros, N. B. and R. S. Wells. 1998. Prey and feeding patterns of resident Bottlenose Dolphins (Tursiops truncatus) in Sarasota, Florida. Journal of Mammalogy 79:1045–1059. 160. Barros, N. B., E. C. M. Parsons and T. A. Jefferson. 2000. Prey of offshore bottlenose dolphins from the South China Sea. Aquatic Mammals 26:2–6. 161. Yochem, P. K. and S. Leatherwood. 1985. Blue Whale Balaenoptera musculus (Linnaeus, 1758). Pp. 193–240 in S. H. Ridgway and R. Harrison, eds. Handbook of Marine Mammals: Volume 3 The sirenians and baleen whales. Academic Press Inc., Toronto, ON. 162. Winn, H. E. and N. E. Reichley. 1985. Humpback Whale Megaptera novaeangliae (Borowski, 1781). Pp. 241–273 in S. H. Ridgway and R. Harrison, eds. Handbook of Marine Mammals: Volume 3 The sirenians and baleen whales. Academic Press Inc., Toronto, ON. 163. Reeves, R. R. and S. Leatherwood. 1985. Bowhead Whale Balaena mysticetus Linnaeus, 1758. Pp. 305–344 in S. H. Ridgway and R. Harrison, eds. Handbook of Marine Mammals: Volume 3 The sirenians and baleen whales. Academic Press Inc., Toronto, ON. 164. Cummings, W. C. 1985. Bryde’s Whale Balaenoptera edeni Anderson, 1878. Pp. 137–154 in S. H. Ridgway and R. Harrison, eds. Handbook of Marine Mammals: Volume 3 The sirenians and baleen whales. Academic Press Inc., Toronto, ON. 165. Gambell, R. 1985. Sei Whale Balanenoptera borealis Lesson, 1828. Pp. 155–170 in S. H. Ridgway and R. Harrison, eds. Handbook of Marine Mammals: Volume 3 The sirenians and baleen whales. Academic Press Inc., Toronto, ON. 166. Gambell, R. 1985. Fin Whale Balaenoptera physalus (Linnaeus, 1758). Pp. 171–192 in S. H. Ridgway and R. Harrison, eds. Handbook of Marine Mammals: Volume 3 The sirenians and baleen whales. Academic Press Inc., Toronto, ON. 167. Aguilar, A. 2002. Fin Whale, Balaenoptera physalus. Pp. 435–438 in W. F. Perrin, B. Würsig, and J. G. M. Thewissen, eds. Encyclopedia of Marine Mammals. Academic Press Inc., San Francisco, CA.

131

168. Lipsky, J. D. 2002. Right Whale Dolphins Lissodelphis borealis and L. peronii. Pp. 1030–1033 in W. F. Perrin, B. Würsig, and J. G. M. Thewissen, eds. Encyclopedia of Marine Mammals. Academic Press Inc., San Francisco, CA. 169. Perrin, W. F. and J. G. Mead. 1994. Clymene Dolphin Stenella clymene (Gray, 1846). Pp. 161–171 in S. H. Ridgway and R. Harrison, eds. Handbook of Marine Mammals: Volume 5 The first book of dolphins. Academic Press Inc., Toronto, ON. 170. Goodall, R. N. P. 1994. Chilean Dolphin Cephalorhynchus eutropia (Gray 1846). Pp. 269–287 in S. H. Ridgway and R. Harrison, eds. Handbook of Marine Mammals: Volume 5 The first book of dolphins. Academic Press Inc., Toronto, ON. 171. Barros, N. B., T. A. Jefferson and E. C. M. Parsons. 2004. Feeding habits of Indo-Pacific Humpback Dolphins (Sousa chinensis) stranded in Hong Kong. Aquatic Mammals 30:179–188. 172. Jefferson, T. A. 1996. Morphology of the clymene dolphin (Stenella clymene) in the northern Gulf of Mexico. Aquatic Mammals 22:35–43. 173. Mignucci-Giannoni, A. A., E. P. Hoberg, D. Siegel-Causey and E. H. Williams. 1998. Metazoan parasites and other symbionts of cetaceans in the Caribbean. The Journal of Parasitology 84:939–946. 174. Santos, M. C. de O., S. Siciliano, A. F. de Castro Vincente, F. S. Alvarenga, E. Zampirolli, S. P. de Souza and A. Maranho. 2010. Cetacean records along São Paulo state coast, southeastern Brazil. Brazilian Journal of Oceanography 58:123–142. 175. Hohn, A. A., A. J. Read, S. Fernández, O. Vidal and L. T. Findley. 1996. Life history of the vaquita, Phocoena sinus (Phocoenidae, Cetacea). Journal of Zoology of London 239:235–251. 176. Read, A. J. 1990. Estimation of body condition in harbour porpoises, Phocoena phocoena. Canadian Journal of Zoology 68:1962–1966. 177. Lockyer, C., M. P. Heide-Jørgensen, J. Jensen, C. C. Kinze and T. Buus Sørensen. 2001. Age, length and reproductive parameters of harbour porpoises Phocoena phocoena (L.) from West Greenland. ICES Journal of Marine Science 58:154–162. 178. Ferrero, R. C. and W. A. Walker. 1993. Growth and reproduction of the northern right whale dolphin, Lissodelphis borealis, in the offshore waters of the North Pacific Ocean. Canadian Journal of Zoology 71:2335– 2344. 179. Slooten, E. 1992. Age, growth, and reproduction in Hector’s dolphins. Canadian Journal of Zoology 69:1689– 1700. 180. Chantrapornsyl, S., K. Adulyanukosol and K. Kittiwathanawong. 1996. Records of cetaceans in Thailand. Phuket Marine Biological Center Research Bulletin 61:39–63. 181. Rose, B. and A. I. L. Payne. 1991. Occurrence and behavior of the Southern Right whale Dolphin Lissodelphis peronii off Namibia. Marine Mammal Science 7:25–34. 182. Claver, J. A., M. A. Iniguez, D. M. Lombardo and I. von Lawzewitsch. 1992. Preliminary observations on the ovarian activity and sexual maturity in female Peale’s dolphin (Lagenorhynchus australis). Aquatic Mammals 18:85–88. 183. Boy, C. C., N. Dellabianca, R. N. P. Goodall and A. C. M. Schiavini. 2011. Age and growth in Peale’s dolphin (Lagenorhynchus australis) in subantarctic waters off southern South America. Mammalian Biology 76:634– 639. 184. Jefferson, T. A., D. Fertl, J. Bolaños-Jiménez and A. N. Zerbini. 2009. Distribution of common dolphins (Delphinus spp.) in the western Atlantic Ocean: a critical re-examination. Marine Biology 156:1109–1124.

132

185. Berón-Vera, B., S. N. Pedraza, J. A. Raga, A. Gil de Pertierra, E. A. Crespo, M. K. Alonso and R. N. P. Goodall. 2001. Gastrointestinal helminths of Commerson’s dolphins Cephalorhynchus commersonii from central Patagonia and Tierra del Fuego. Diseases of Aquatic Organisms 47:201–208. 186. Wells, R. S. and M. D. Scott. 1999. Bottlenose Dolphin Tursiops truncatus (Montagu, 1821). Pp. 137–182 in S. H. Ridgway and R. Harrison, eds. Handbook of Marine Mammals: Volume 6 The second book of dolphins and the porpoises. Academic Press Inc., Toronto, ON. 187. Perrin, W. F. and S. B. Reilly. 1982. Reproductive parameters of dolphins and small whales of the family Deiphinidae. Report of the International Whaling Commission Special Is:97–133. 188. Torres, P., J. A. Oporto, L. M. Brieva and L. Escare. 1992. Gastrointestinal Helminths of the cetaceans Phocoena spinipinnis (Burneister, 1865) and Cephalorhynchus eutropis (Gray, 1846) from the Southern coast of Chile. Journal of Wildlife Diseases 28:313–315. 189. Williams, R., S. L. Hedley and P. S. Hammond. 2006. Modeling distribution and abundance of Antarctic baleen whales using ships of opportunity. Ecology and Society 11:1. [online] URL: http://www.ecologyandsociety.org/. 190. Tormosov, D. D., Y. A. Mikhaliev, P. B. Best, V. A. Zemsky, K. Sekiguchi and R. L. Brownell Jr. 1998. Soviet catches of southern right whales Eubalaena australis, 1951 ± 1971. Biological data and conservation implications. Biological Conservation 86:185–197. 191. Braham, H. W. and D. W. Rice. 1984. The Right whale, Balaena glacialis. Marine Fisheries Review 46:38–44. 192. Mead, J. G. 1986. Twentieth-century records of Right Whales (Eubalaena glacialis) in the Northwestern Atlantic Ocean. Report of the International Whaling Commission Special Is:109–119. 193. Cosens, S. E. and L. P. Dueck. 1991. Group size and activity patterns of belugas (Dephinaptera leucas) and narwhals (Monodon monoceros) during spring migration in Lancaster Sound. Canadian Journal of Zoology 69:1630–1635. 194. Canning, S. J., M. B. Santos, R. J. Reid, P. G. H. Evans, R. C. Sabin, N. Bailey and G. J. Pierce. 2008. Seasonal distribution of white-beaked dolphins (Lagenorhynchus albirostris) in UK waters with new information on diet and habitat use. Journal of the Marine Biological Association of the United Kingdom 88:1159–1166. 195. Kanaji, Y., H. Okamura and T. Miyashita. 2011. Long-term abundance trends of the northern form of the short-finned pilot whale (Globicephala macrorhynchus) along the Pacific coast of Japan. Marine Mammal Science 27:477–492. 196. Pinto, R. M., L. C. Muniz-Pereira, V. C. Alves and S. Siciliano. 2004. First report of a Helminth infection for Bryde’s Whale Balaenoptera edeni Andreson, 1878 (Cetacea, Balaenopteridae). Latin American Journal of Aquatic Mammals 3:167–170. 197. Ferguson, M. C., J. Barlow, P. Fiedler, S. B. Reilly and T. Gerrodette. 2006. Spatial models of delphinid (family Delphinidae) encounter rate and group size in the eastern tropical Pacific Ocean. Ecological Modelling 193:645–662. 198. Ridgway, S. H. 1996. Final report from the Right Whale necropsy team: results, analysis, and recommendations. Naval Command and Ocean Surveillance Center. Technical Document 2934 San Diego, CA. 199. Reeves, R. R. and R. L. Brownell. 2008. Report of the assessment workshop on Indo-Pacific Bottlenose Dolphins (Tursiops aduncus) with Solomon Islands as a case study. Apia, Spain. 200. Culik, B. M. 2004. Review of small cetaceans; distribution, behaviour, migration and threats. Bonn, Germany.

133

201. Di Beneditto, A. P. M. and S. Siciliano. 2007. Stomach contents of the marine tucuxi dolphin (Sotalia guianensis) from Rio de Janeiro, south-eastern Brazil. Journal of the Marine Biological Association of the United Kingdom 87:253–254. 202. Best, P. B., J. P. Glass, P. G. Ryan and M. L. Dalebout. 2009. Cetacean records from Tristan da Cunha, South Atlantic. Journal of the Marine Biological Association of the United Kingdom 89:1023–1032. 203. A, D. and V. DeBuffrenil. 1989. Acoustic signals of the Commerson’s Dolphin, Cephalorhynchus commersonii, in the Kerguelen Islands. Journal of Mammalogy 70:449–452. 204. Lauriano, G., C. M. Fortuna and M. Vacchi. 2010. Occurrence of killer whales (Orcinus orca) and other cetaceans in Terra Nova Bay, Ross Sea, Antarctica. Antarctic Science 23:139–143. 205. Miyazaki, N. and K. Hidehira. 1988. Sitting records of small cetaceans in the southern hemisphere. Bulletin of the National Science Museum, Tokyo, Series A. 14:47–65. 206. Mizroch, S. A. and D. W. Rice. 2006. Have North Pacific killer whales switched prey species in response to depletion of the great whale populations? Marine Ecology Progress Series 310:235–246. 207. Gannier, A. 2009. Comparison of odontocete populations of the Marquesas and Society Islands (French Polynesia). Journal of the Marine Biological Association of the United Kingdom 89:931–941. 208. Herzing, D. L., K. Moewe and B. J. Brunnick. 2003. Interspecies interactions between Atlantic spotted dolphins, Stenella frontalis and bottlenose dolphins, Tursiops truncatus, on Great Bahama Bank, Bahamas. Aquatic Mammals 29:335–341. 209. Pitman, R. L., S. O’Sullivan and B. Mase. 2003. Killer whales (Orcinus orca) attack a school of pantropical spotted dolphins (Stenella attenuata) in the Gulf of Mexico. Aquatic Mammals 29:321–324. 210. Psarakos, S., D. L. Herzing and K. Marten. 2003. Mixed-species associations between Pantropical Spotted dolphins (Stenella attenuata) and Hawaiian Spinner dolphins (Stenella longirostris) off Oahu, Hawaii. Aquatic Mammals 29:390–395. 211. Herzing, D. L. and C. M. Johnson. 1997. Interspecific interactions between Atlantic Spotted dolphins (Stenella frontalis) and Bottlenose dolphins (Tursiops truncatus) in the Bahamas, 1985–1995. Aquatic Mammals 23:85–99. 212. Moreno, I. B., A. N. Zerbini, D. Danilewicz, M. C. D. O. Santos, P. C. Simões-Lopes, J. Lailson-Brito Jr. and A. F. Azevedo. 2005. Distribution and habitat characteristics of dolphins of the genus Stenella (Cetacea: Delphinidae) in the southwest Atlantic Ocean. Marine Ecology Progress Series 300:229–240. 213. Silva Jr, J. M., F. J. De Lima Silva, C. Sazima and I. Sazima. 2007. Trophic relationships of the spinner dolphin at Fernando de Noronha Archipelago, SW Atlantic. Scientia Marina 71:505–511. 214. Flores, P. A. and V. M. Da Silva. 2002. Tucuxi and Guiana Dolphin Sotalia fluviatilis and S. guianensis. Pp. 1188–1192 in W. F. Perrin, B. Würsig, and J. G. M. Thewissen, eds. Encyclopedia of Marine Mammals. Academic Press Inc., San Francisco, CA. 215. Wedekin, L. L., F. G. Daura-Jorge and P. C. Simões-Lopes. 2004. An aggressive interaction between Bottlenose Dolphins (Tursiops truncatus) and estuarine Dolphins (Sotalia guianensis) in southern Brazil. Aquatic Mammals 30:391–397. 216. Acevedo-Gutiérrez, A., A. DiBerardinis, S. Larkin, K. Larkin and P. Forestell. 2005. Social interactions between Tucuxis and Bottlenose dolphins in Gandoca-Manzanillo, Costa Rica. Latin American Journal of Aquatic Mammals 4:49–54. 217. Jefferson, T. A. and B. E. Curry. 2003. Stenella clymene. Mammalian Species 726:1–5. 218. Frost, K. J., R. B. Russell and L. F. Lowery. 1992. Killer Whales, Orcinus orca, in the southeastern Berring Sea: recent sightings and predation on other marine mammals. Marine Mammal Science 8:110–119.

134

219. Thiemann, G. W., S. J. Iverson and I. Stirling. 2008. Polar Bear diets and arctic marine food webs: insights from fatty acid analysis. Ecological Monographs 78:591–613. 220. Orr, J. R. and L. A. Harwood. 1998. Possible aggressive behavior between a Narwhal (Monodon monoceros) and a Beluga (Delphinapterus leucas). Marine Mammal Science 14:182–185. 221. Smith, T. G. and B. Sjare. 1990. Prdation of Belugas and Narwhals by Polar Bears in nearshore areas of the Canadian High Arctic. Arctic 43:99–102. 222. Melinikov, V. V. and I. A. Zagrebin. 2005. Killer whale predation in coastal waters of the Chukotka peninsula. Marine Mammal Science 21:550–556. 223. Shelden, K. E., D. J. Rugh, B. A. Mahoney and M. E. Dahlheim. 2003. Killer Whale predation on Belugas in Cook Inlet, Alaska: implications for a depleted population. Marine Mammal Science 19:529–544. 224. Taguchi, M., H. Ishikawa and T. Matsuishi. 2010. Seasonal distribution of Harbour Porpoise (Phocoena phocoena) in Japanese waters inferred from stranding and bycatch records. Mammal Study 35:133–138. 225. Visser, I. N., J. Zaeschmar, J. Halliday, A. Abraham, P. Ball, R. Bradley, S. Daly, T. Hatwell, T. Johnson, W. Johnson, L. Kay, T. Maessen, V. McKay, T. Peters, N. Turner, B. Umuroa and D. S. Pace. 2010. First record of predation on False Killer Whales (Pseudorca crassidens) by Killer Whales (Orcinus orca). Aquatic Mammals 36:195–204. 226. Read, A. J. 1999. Harbour Porpoise Phocoena phocoena (Linnaeus, 1758). Pp. 323–378 in S. H. Ridgway and R. G. Harrison, eds. Handbook of Marine Mammals: Volume 6 The second book of dolphins and the porpoises. Academic Press Inc., San Diego, CA. 227. Watts, P. and D. E. Gaskin. 1985. Habitat index analysis of the Harbor Porpoise (Phocoena phocoena) in the southern Bay of Fundy, Canada. Journal of Mammalogy 66:733–744. 228. Baird, R. W. 1998. An interaction between Pacific White-Sided Dolphins and a neonatal harbor porpoise. Mammalia 62:129–134. 229. Willis, P. M., B. J. Crespi, L. M. Dill, R. W. Baird and M. B. Hanson. 2004. Natural hybridization between Dall’s porpoises (Phocoenoides dalli) and harbour porpoises (Phocoena phocoena). Canadian Journal of Zoology 82:828–834. 230. Ross, H. M. and B. Wilson. 1996. Violent interactions between bottlenose dolphins and harbour porpoises. Proceedings: Biological Sciences 263:283–286. 231. Haelters, J. and E. Everaarts. 2011. Two cases of physical interaction between White-Beaked Dolphins (Lagenorhynchus albirostris) and juvenile Harbour Porpoises (Phocoena phocoena) in the southern North Sea. Aquatic Mammals 37:198–201. 232. Dahlheim, M. E. and P. A. White. 2010. Ecological aspects of transient killer whales Orcinus orca as predators in southeastern Alaska. Wildlife Biology 16:308–322. 233. Arnold, P. W. 1972. Predation on Harbour Porpoise, Phocoena phocoena, by a White Shark, Carcharodon carcharias. Journal of the Fisheries Research Board of Canada 29:1213–1214. 234. Williamson, G. 1963. Common Porpoise from the Stomach of a Greenland Shark. Journal of the Fisheries Research Board of Canada 20:1085–1087. 235. Jefferson, T. A. 1991. Observations on the distribution and behaviour of Dall’s porpoise (Phocoenoides dalli) in Monteray Bay, California. Aquatic Mammals 17:12–19. 236. Walker, W. A. 2001. Geographical variation of the parasite, Phyllobothrium delphini (Cestoda), in Dall’s Porpoise, Phocoenoides dalli, in the northern North Pacific, Berring Sea, and Sea of Okhotsk. Marine Mammal Science 17:264–275.

135

237. Conlogue, G. J., J. A. Ogden and W. J. Foreyt. 1985. Parasites of the Dall’s Porpoise (Phocoenoides dalli True). Journal of Wildlife Diseases 21:160–166. 238. Brownell Jr., R. L. 1975. Phocoena dioprica. Mammalian Species 66:1–3. 239. Pinedo, M. C., A. S. Barreto, M. P. Lammardo, A. L. V. Andrade and L. Geracitano. 2002. Northernmost records of the Spectacled Porpoise, Layard’s Beaked Whale, and Peale's Dolphin in the southwestern Atlantic Ocean. Aquatic Mammals 28:32–37. 240. Gao, A. and K. Zhou. 1993. Growth and reproduction of three populations of finless porpoise, Neophocaena phocaenoides, in Chinese waters. Aquatic Mammals 19:3–12. 241. Yoshida, H., N. Higashi, H. Ono and S. Uchida. 2010. Finless Porpoise (Neophocaena phocaenoides) discovered at Okinawa Island, Japan, with the source population inferred from mitochondrial DNA. Aquatic Mammals 36:278–283. 242. Parsons, E. and T. A. Jefferson. 2000. Post-mortem investigations on stranded dolphins and porpoises from Hong Kong waters. Journal of Wildlife Diseases 36:342–356. 243. Silber, G. K. and K. S. Norris. 1991. Geographic and seasonal distribution of the Vaquita, Phocoena sinus. Anales del Instituto de Biología, Universidad Autónoma del Estado de México. Serie Zoología 62:263–268. 244. Silber, G. K. 1990. Occurrence and distribution of the Vaquita Phocoena sinus in the Northern Gulf of California. Fishery Bulletin 88:339–346. 245. Silber, G. K. 1988. Recent sightings of the Gulf of California Harbour Porpoise, Phocoena sinus. Journal of Mammalogy 69:430–433. 246. Silber, G. K., M. W. Newcomer, P. C. Silber, H. Pérez-Cortés M. and G. M. Ellis. 1994. Cetaceans of the Northern Gulf of California: distribution, occurrence and relative abundance. Marine Mammal Science 10:283–298. 247. Silber, G. K., M. W. Newcomer and H. Pérez-Cortés M. 1990. Killer whales (Orcinus orca) attack and kill a Bryde’s whale (Balaenoptera edeni). Canadian Journal of Zoology 68:1603–1606. 248. de Oliveira Santos, M. C., J. E. de Faria Oshima and E. da Silva. 2009. Sightings of Franciscana dolphins (Pontporia blainvillei): the discovery of a population in the Paranaguá Estuarine Complex, southern Brazil. Brazilian Journal of Oceanography 57:57–63. 249. Ott, P. H. and D. Danilewicz. 1998. Presence of franciscana dolphins (Pontporia blainvillei) in the stomach of a killer whale (Orcinus orca) stranded in southern Brazil. Mammalia 62:605–609. 250. Di Beneditto, A. P. M. 2004. Presence of Franciscana Dolphin (Pontoporia blainvillei) remains in the stomach of a Tiger Shark (Galeocerdo cuvieri) captured in southeastern Brazil. Aquatic Mammals 30:311–314. 251. Lucifora, L. O., R. C. Menni and A. H. Escalante. 2005. Reproduction, abundance and feeding habits of the broadnose sevengill shark Notorynchus cepedianus in north Patagonia, Argentina. Marine Ecology Progress Series 289:237–244. 252. Santos, M., J. Oshima, E. Pacífico and E. Silva. 2010. Group size and composition of Guiana dolphins (Sotalia guianensis) (Van Bénèden, 1864) in the Paranaguá Estuarine Complex, Brazil. Brazilian Journal of Biology 70:111–20. 253. Kaschner, K., R. Watson, A. Trites and D. Pauly. 2006. Mapping world-wide distributions of marine mammal species using a relative environmental suitability (RES) model. Marine Ecology Progress Series 316:285–310. 254. Dietz, R. and M. P. Heide-Jørgensen. 1995. Movements and swimming speed of Narwhals, Monodon monoceros, equipped with satellite transmitters in Melville Bay, northwest Greenland. Canadian Journal of Zoology 73:2106–2119.

136

255. Rako, N., H. Draško and C. M. Fortuna. 2009. Long-term inshore observation of a solitary Striped Dolphin, Stenella coeruleoalba, in the Vinodol Channel, Northern Adriatic Sea (Croatia). Natura Croatica 18:427–436. 256. Frantzis, A. and D. L. Herzing. 2002. Mixed-species associations of Striped dolphins (Stenella coeruleoalba), Short-beaked Common dolphins (Delphinus delphis), and Risso’s dolphins (Grampus griseus) in the Gulf of Corinth (Greece, Mediterranean Sea). Aquatic Mammals 28:188–197. 257. Rosas, F. C. W., E. L. A. Monteiro-Filho, J. Marigo, R. A. Santos, A. L. V. Andrade, M. Rautenberg, M. R. Oliveira and M. O. Bordignon. 2002. The striped dolphin, Stenella coeruleoalba (Cetacea: Delphinidae), on the coast of São Paulo State, southeastern Brazil. Aquatic Mammals 28:60–66. 258. Dwyer, S. L. and I. N. Visser. 2011. Cookie Cutter Shark (Isistius sp.) bites on cetaceans, with particular reference to Killer Whales (Orca) (Orcinus orca). Aquatic Mammals 37:111–138. 259. Crovetto, A., J. Lamilla and G. Pequeño. 1992. Lissodelphis peronii, Lacépède 1804 (Delphinidae, Cetacea) within the stomach contents of a sleeping shark, Somniosus CF. pacificus, Bigelow and Schroeder 1944, in Chilean Waters. Marine Mammal Science 8:312–314. 260. Migura, K. A. and D. W. Meadows. 2002. Short-finned Pilot whales (Globicephala macrorhynchus) interact with Melon-Headed whales (Peponocephala electra) in Hawaii. Aquatic Mammals 28:294–297. 261. Steiger, G. H., J. Calambokidis, J. M. Straley, L. M. Herman, S. Cerchio, D. R. Salden, J. Urbán-R, J. K. Jacobsen, O. von Ziegesar, K. C. Balcomb, C. M. Gabriele, M. E. Dahlheim, S. Uchida, J. K. B. Ford, P. Ladrón de Guevara-P, M. Yamaguchi and J. Barlow. 2008. Geographic variation in killer whale attacks on humpback whales in the North Pacific: implications for predation pressure. Endangered Species Research 4:247–256. 262. Weir, C. R. 2010. First description of Atlantic Humpback Dolphin Sousa teuszii whistles, recorded off Angola. Bioacoustics 19:211–224. 263. Perrin, W. F. and W. A. Walker. 1975. The Rough-Toothed Porpoise, Steno bredanensis, in the eastern tropical Pacific. Journal of Mammalogy 56:905–907. 264. Ritter, F. 2002. Behavioural observation of rough-toothed dolphins (Steno bredanensis) off La Gomera, Canery Islands (1995-2000), with special reference to their interactions with humans. Aquatic Mammals 28:46–59. 265. Celona, A., A. De Maddalena and G. Comparetto. 2006. Evidence of predatory attack on a Bottlenose Dolphin Tursiops truncatus by a Great White shark Carcharodon carcharias in the Mediterranean Sea. Annales Ser. hist. nat. 16:159–164. 266. Bruce, B. D. 1992. Preliminary observations on the biology of the White Shark, Carcharodon carcharias, in South Australian waters. Australian Journal of Marine and Freshwater Resources 43:1–11. 267. Rossi-Santos, M. R., E. Santos-Neto and C. G. Baracho. 2009. Interspecific cetacean interactions during the breeding season of humpback whale (Megaptera novaeangliae) on the north coast of Bahia State, Brazil. Journal of the Marine Biological Association of the United Kingdom 89:961–966. 268. Gannier, A. 2002. Cetaceans of the Marquesas Islands (French Polynesia): distribution and relative abundance as obtained from a small boat dedicated survey. Aquatic Mammals 28:198–210. 269. Ciano, J. and R. Jørgensen. 2000. Observations on an interaction between a Humpback Whale (Megaptera novaeangliae) and Pilot Whales (Globicephala melas). Marine Mammal Science 16:245–248. 270. Williams, A. D., R. Williams and T. Brereton. 2002. The sighting of pygmy killer whales (Feresa attenuata) in the southern Bay of Biscay and their association with cetacean calves. Journal of the Marine Biological Association of the UK 82:509–511. 271. Reyes, L. M. and P. García-Borboroglu. 2004. Killer Whale (Orcinus orca) predation on sharks in Patagonia, Argentina: a first report. Aquatic Mammals 30:376–379.

137

272. Perryman, W. L., D. W. K. Au, S. Leatherwood and T. A. Jefferson. 1994. Melon-headed Whale Peponocephala electra Gray, 1846. Pp. 363–386 in R. G. Harrison and S. H. Ridgway, eds. Handbook of Marine Mammals: Volume 5 The first book of dolphins. Academic Press Inc., San Diego, CA. 273. Wang, J. Y., S. C. Yang, S. Hung and T. A. Jefferson. 2007. Distribution, abundance and conservation status of the eastern Taiwan Strait population of Indo-Pacific humpback dolphins, Sousa chinensis. Mammalia 157–165. 274. Goodall, R. N. P. 1994. Commerson’s Dolphin Cephalorhynchus commersonii (Lacépède 1804). Pp. 241–267 in S. H. Ridgway and R. G. Harrison, eds. Handbook of Marine Mammals: Volume 5 The first book of dolphins. Academic Press Inc., San Diego, CA. 275. Santora, J. A. 2012. Habitat use of hourglass dolphins near the South Shetland Islands, Antarctica. Polar Biology 35:801–806. 276. Hale, P. T., A. S. Barreto and G. J. B. Ross. 2000. Comparative morphology and distribution of the aduncus and truncatus forms of bottlenose dolphin Tursiops in the Indian and Western Pacific Oceans. Aquatic Mammals 26:101–110. 277. Dulau-Drouot, V., V. Boucaud and B. Rota. 2008. Cetacean diversity off La Réunion Island (France). Journal of the Marine Biological Association of the United Kingdom 88:1263–1272. 278. Matsuoka, K., H. Kiwada, Y. Fujise and T. Miyashita. 2007. Distribution of blue (Balaenoptera musculus), fin (B. physalus), humpback (Megaptera novaeangliae) and north pacific right (Eubalaena japonica) whales in the western North Pacific based on JARPN and JARPN II sighting surveys (1994 to 2007). Institute of Cetacean Research. Scientific papers submitted to the JARPNII REVIEW meeting called by the International Whaling Commission (2009) SC/J09/JR35 279. Tynan, C. T., D. P. DeMaster and W. T. Peterson. 2001. Endangered right whales on the southeastern Bering Sea shelf. Science 294:1894. 280. Borstad, G. A. 1985. Water colour and temperature in the Southern Beaufort Sea: Remote sensing in support of ecological studies of the Bowhead Whale. Department of Fisheries and Oceans Canada. Canadian Technical Report of Fisheries and Aquatic Sciences No. 1350 281. Laidre, K. L., M. P. Heide-Jørgensen, M. L. Logsdon, L. Delwiche and T. G. Nielsen. 2010. A whale of an opportunity: Examining the vertical structure of chlorophyll-a in high Arctic waters using instrumented marine predators. Marine Biology Research 6:519–529. 282. Jayasankar, P., A. A. Krishnan, M. Rajagopalan and P. K. Krishnakumar. 2007. A note on observations on cetaceans in the western Indian sector of the Southern Ocean (20-56°S and 45-57°30’E ), January to March 2004. Journal of Biogeography 9:263–267. 283. Watanabe, H., M. Okazaki, T. Tamura, K. Konishi, D. Inagake, T. Bando, H. Kiwada and T. Miyashita. 2012. Habitat and prey selection of common minke, sei, and Bryde’s whales in mesoscale during summer in the subarctic and transition regions of the western North Pacific. Fisheries Science 78:557–567. 284. Molina-Schiller, D., S. A. Rosales and T. R. O. de Freitas. 2005. Oceanographic conditions off coastal South America in relation to the distribution of Burmeister’s porpoise, Phocoena spinipinnis. Latin American Journal of Aquatic Mammals 4:141–156. 285. Weir, C. R. 2009. Distribution, behaviour and photo-identification of Atlantic humpback dolphins Sousa teuszii off Flamingos, Angola. African Journal of Marine Science 31:319–331. 286. Baldwin, R. M., M. Collins, K. Van Waerebeek and G. Minton. 2004. The Indo-Pacific Humpback Dolphin of the Arabian Region: A status review. Aquatic Mammals 30:111–124.

138

287. Cribb, N., C. Miller and L. Seuront. 2008. Assessment of bottlenose dolphin (Tursiops aduncus) habitat characteristics in the estuarine waters of the Adelaide Dolphin Sanctuary, South Australia. Journal of Marine Animals and Their Ecology 1:6–8. 288. Baumgartner, M. F., S. M. Van Parijs, F. W. Wenzel, C. J. Tremblay, H. C. Esch and A. M. Warde. 2008. Low frequency vocalizations attributed to sei whales (Balaenoptera borealis). Journal of the Acoustical Society of America 124:1339–1349. 289. Erbe, C. 2004. The acoustic repertoire of odontecetes as a basis for developing automatic detectors and classifiers. Defense R & D Canada – Atlantic. DRDC Atlantic CR 2004-071 290. Goodall, R. N. P. 2002. Peale’s Dolphin Lagenorhynchus australis. Pp. 844–847 in W. F. Perrin, B. Würsig, and J. Thewissen, eds. Encyclopedia of Marine Mammals. Academic Press Inc., San Francisco, CA. 291. Penry, G. S., V. G. Cockcroft and P. S. Hammond. 2011. Seasonal fluctuations in occurrence of inshore Bryde’s whales in Plettenberg Bay, South Africa, with notes on feeding and multispecies associations. African Journal of Marine Science 33:403–414. 292. Smultea, M. A., A. B. Douglas, C. E. Bacon, T. A. Jefferson and L. Mazzuca. 2012. Bryde’s Whale (Balaenoptera brydei/edeni) sightings in the southern California bight. Aquatic Mammals 38:92–97. 293. de Stephanis, R., T. Cornulier, P. Verborgh, J. Salazar Sierra, N. P. Gimeno and C. Guinet. 2008. Summer spatial distribution of cetaceans in the Strait of Gibraltar in relation to the oceanographic context. Marine Ecology Progress Series 353:275–288. 294. Lachmuth, C. L., J. J. Alava, B. E. Hickie, S. C. Johannessen, R. W. Macdonald, J. K. B. Ford, G. M. Ellis, F. A. P. C. Gobas and P. S. Ross. 2010. Ocean disposal in resident killer whale (Orcinus orca) Critical Habitat: Science in support if risk management. Department of Fisheries and Oceans Canada. Canadian Science Advisory Secretariat Research Document 2010/116 295. Kreb, D. and Budiono. 2005. Cetacean diversity and habitat preferences in tropical waters of east Kalimantan, Indonesia. The Raffles Bulletin of Zoology 53:149–155. 296. Di Sciara, G. N., M. C. Venturino, M. Zanardelli, G. Bearzi, F. J. Borsani and B. Cavalloni. 2009. Cetaceans in the central Mediterranean Sea: Distribution and sighting frequencies. Bolletino di zoologia 60:37–41. 297. Elwen, S. H., M. Thornton, D. Reeb and P. B. Best. 2010. Near-Shore Distribution of Heaviside’s (Cephalorhynchus heavisidii) and Dusky Dolphins (Lagenorhynchus obscurus) at the Southern Limit of their Range in South Africa. African Zoology 45:78–91. 298. Frantzis, A., P. Alexiadou, G. Paximadis, E. Politi, A. Gannier and M. Corsini-Foka. 2003. Current knowledge of the cetacean fauna of the Greek Seas. Journal of Cetacean Research and Management 5:219–232. 299. Frazer, J. F. D. 1976. Herd structure and behaviour in cetaceans. Mammal Review 6:55–59. 300. Hooker, S. K., H. Whitehead and S. Gowans. 1999. Marine protected area design and the spatial and temporal distribution of cetaceans in a submarine canyon. Conservation Biology 13:592–602. 301. Moore, M. J., S. D. Berrow, B. A. Jensen, P. Carr, R. Sears, V. J. Rowntree and R. Payne. 1999. Relative abundance of large whales around South Georgia (1979-1998). Marine Mammal Science 15:1287–1302. 302. Best, P. B. 1974. Two allopatric forms of Bryde’s whales off South Africa. Reports of the International Whaling Commission Special Is:10–38. 303. Dahlheim, M. E. and J. E. Heyning. 1999. Killer Whale Orcinus orca (Linnaeus, 1758). Pp. 281–322 in S. H. Ridgway and R. Harrison, eds. Handbook of Marine Mammals: Volume 6 The second book of dolphins and the porpoises. Academic Press Inc., Toronto, ON.

139

304. Jefferson, T. A., P. J. Stacey and R. W. Baird. 1991. A review of Killer Whale interactions with other marine mammals: predation to co-existence. Mammal Review 21:151–180. 305. Visser, I. N., J. Berghan, R. V. Meurs and D. Fertl. 2000. Killer whale (Orcinus orca) predation on a shortfin mako shark (Isurus oxyrinchus) in New Zealand waters. Aquatic Mammals 26:229–231. 306. Ford, J. J. B. and G. M. Ellis. 2006. Selective foraging by fish-eating killer whales Orcinus orca in British Columbia. Marine Ecology Progress Series 316:185–199. 307. Ross, G. J. B., G. E. Heinsohn and V. G. Cockcroft. 1994. Humpback Dolphins Sousa chinensis (Osbeck,1765), Sousa plumbea (G. Cuvier, 1829) and Sousa teuszii (Kukenthal, 1892). Pp. 23–42 in S. H. Ridgway and R. Harrison, eds. Handbook of Marine Mammals: Volume 5 The first book of dolphins. Academic Press Inc., Toronto, ON. 308. Pitman, R. L., A. L. van Helden, P. B. Best and A. T. Pym. 2006. Shepherd’s Beaked Whale (Tasmacetus shepherdi): information on appearance and biology based on strandings and at-sea observations. Marine Mammal Science 22:744–755. 309. Yatabe, A., N. Kubo, M. Otsuka, S. Shima, T. Kubodera and T. K. Yamada. 2010. Stomach contents and structure of a Longman’s Beaked Whale (Indopacetus pacificus) stranded in Kyushu, Japan. Aquatic Mammals 36:172–177. 310. Loughlin, T. and M. A. Perez. 1985. Mesoplodon stejnegeri. Mammalian Species 250:1–6. 311. Jefferson, T. A. and M. W. Newcomer. 1993. Lissodelphis borealis. Mammalian Species 425:1–6. 312. Jackson, A., T. Gerrodette, S. Chivers, M. Lynn, S. Rankin and S. Mesnick. 2006. Marine mammal data collected during a survey in the eastern tropical Pacific Ocean aboard NOAA ships David Starr Jordan and McArthur II, July 28-December 7, 2006. 313. Reyes, J. C., J. G. Mead and K. Van Waerebeek. 1991. A new species of beaked whale Mesoplodon peruvianus sp. N. (Cetacea: Ziphiidae) from Peru. Marine Mammal Science 7:1–24. 314. Carrillo, M. 2003. Presence and distribution of the Ziphiidae family in the southwest coast of Tenerife. Canary Islands. Pp. 1–5 in 17th Conference of the European Cetacean Society. 315. Moore, J. C. and F. Wood Jr. 1957. Differences between the beaked whales Mesoplodon mirus and Mesoplodon gervaisi. American Museum Novitates 1831:1–25. 316. Würsig, B., T. A. Jefferson and D. J. Schmidly. 2000. The Marine Mammals of the Gulf of Mexico. First Edition. Texas A&M University Press, College Station. 317. Stacey, P. J. and R. W. Baird. 1991. Status of the Pacific White-sided dolphin, Lagenorhynchus obliquidens, in Canada. Canadian Field-Naturalist 105:219–232. 318. Stacey, P. J. and R. W. Baird. 1991. Status of the False Killer Whale, Pseudorca crassidens, in Canada. Canadian Field-Naturalist 105:189–197. 319. Heithaus, M. R. 2001. Predator-prey and competitive interactions between sharks (order Selachii) and dolphins (suborder Odontoceti): a review. Journal of Zoology of London 253:53–68. 320. Campbell, R. R., D. B. Yurik and N. B. Snow. 1988. Predation on Narwhals, Monodon monoceros, by Killer Whales, Orcinus orca, in the eastern Canadian Arctic. Canadian Field-Naturalist 102:689–696. 321. Khan, M., S. Panda, A. K. Pattnaik, B. C. Guru, C. Kar, M. Subudhi and R. Samal. 2011. Shark attacks on Irrawaddy dolphin in Chilika lagoon, India. Journal of the Marine Biological Association of India 53:27–34. 322. Abrantes, K. and A. Barnett. 2011. Intrapopulation variations in diet and habitat use in a marine apex predator, the broadnose sevengill shark Notorynchus cepedianus. Marine Ecology Progress Series 442:133–148.

140

323. Weller, D. W. 2002. Predation on Marine Mammals. Pp. 985–994 in W. F. Perrin, B. Würsig, and J. G. M. Thewissen, eds. Encyclopedia of Marine Mammals. Academic Press Inc., San Francisco, CA. 324. Kemper, C. M. 2002. Pygmy Right Whale, Caperea marginata. Pp. 1010–1012 in W. F. Perrin, B. Würsig, and J. G. M. Thewissen, eds. Encyclopedia of Marine Mammals. Academic Press Inc., San Francisco, CA. 325. Watts, P. D., B. A. Draper and J. Henrico. 1991. Preferential use of warm water habitat by adult Beluga Whales. Journal of Thermal Biology 16:57–60. 326. Lopez, S., R. Meléndez and P. Barría. 2009. Alimentación del tiburón marrajo Isurus oxyrinchus Rafinesque, 1810 (Lamniformes : Lamnidae) en el Pacífico suroriental. Revista be Biología Marina y Oceanografía 44:439– 451. 327. Rice, D. W. 1989. Sperm Whale Physeter macrocephalus Linnaeus, 1758. Pp. 177–233 in S. H. Ridgway and R. Harrison, eds. Handbook of Marine Mammals: Volume 4 River dolphins and the larger toothed whales. Academic Press Inc., Toronto, ON. 328. Caldwell, D. K. and M. C. Caldwell. 1989. Pygmy Sperm Whale Kogia breviceps (de Blainville, 1838): Dwarf Sperm Whale Kogia simus Owen, 1866. Pp. 235–260 in S. H. Ridgway and R. Harrison, eds. Handbook of Marine Mammals: Volume 4 River dolphins and the larger toothed whales. Academic Press Inc., Toronto, ON. 329. Nagorsen, D. 1985. Kogia simus. Mammalian Species 239:1–6. 330. Bloodworth, B. E. and D. K. Odell. 2008. Kogia breviceps (Cetacea: Kogiidae). Mammalian Species 819:1–12. 331. Hohn, A. A., D. S. Rotstein, C. A. Harms and B. L. Southall. 2006. Report on Marine Mammal Unusual Mortality Event UMESE0501Sp: Multispecies Mass Stranding of Pilot Whales (Globicephala macrorhynchus), Minke Whale (Balaenoptera acutorostrata), and Dwarf Sperm Whales (Kogia sima) in North Carolina on 15-16 January 2005. National Marine Fisheries Service. NOAA Technical Memorandum NMFS-SEFSC-537 332. Jaquet, N. and D. Gendron. 2002. Distribution and relative abundance of sperm whales in relation to key environmental features, squid landings and the distribution of other cetacean species in the Gulf of California, Mexico. Marine Biology 141:591–601. 333. Cañadas, A., R. Sagarminaga, R. De Stephanis, E. Urquiola and P. S. Hammond. 2005. Habitat preference modelling as a conservation tool: proposals for marine protected areas for cetaceans in southern Spanish waters. Aquatic Conservation: Marine and Freshwater Ecosystems 15:495–521. 334. Whitehead, H. 2002. Sperm Whale Physeter macrocephalus. Pp. 1165–1172 in W. F. Perrin, B. Würsig, and J. G. M. Thewissen, eds. Encyclopedia of Marine Mammals. Academic Press Inc., San Francisco, CA. 335. Taylor, B. L., R. W. Baird, J. Barlow, S. M. Dawson, J. Ford, J. G. Mead, G. Notarbartolo di Sciara, P. Wade and R. L. Pitman. 2008. Kogia sima. IUCN 2012. IUCN Red List of Threatened Speces. Version 2012.1. [Date accessed: June 21, 2012] 336. Mead, J. G. 1989. Bottlenose Whales Hyperoodon ampullatus (Forster, 1770) and Hyperoodon planifrons Flower, 1882. Pp. 321–349 in S. Ridgway and R. G. Harrison, eds. Handbook of Marine Mammals: Volume 4 River dolphins and the larger toothed whales. Academic Press Inc., Toronto, ON. 337. Vidal, O., R. L. Brownell Jr. and L. T. Findley. 1999. Vaquita Phocoena sinus Norris and McFarland, 1958. Pp. 357–378 in S. H. Ridgway and R. G. Harrison, eds. Handbook of Marine Mammals: Volume 6 The second book of dolphins and the porpoises. Academic Press Inc., Toronto, ON. 338. Frantzis, A., J. C. Goold, E. K. Skarsoulis, M. I. Taroudakis and V. Kandia. 2002. Clicks from Cuvier’s beaked whales, Ziphius cavirostris (L). Journal of the Acoustical Society of America 112:34–37.

141

339. Dawson, S. M., J. Barlow and D. Ljungblad. 1998. Sounds recorded from Baird’s beaked whale, Berardius bardii. Marine Mammal Science 14:335–344. 340. May-Collado, L. J., I. Agnarsson and D. Wartzok. 2007. Reexamining the relationship between body size and tonal signals frequency in whales: a comparative approach using a novel phylogeny. Marine Mammal Science 23:524–552. 341. Oswald, J. N., J. Barlow and T. F. Norris. 2003. Acoustic identification of nine delphinid species in the eastern tropical Pacific Ocean. Marine Mammal Science 19:20–37. 342. Watkins, W. A., W. E. Schevill and P. B. Best. 1977. Underwater sounds of Cephalorhynchus heavisidii (Mammalia: Cetacea). Journal of Mammalogy 58:316–320. 343. Rendell, L. E., J. N. Matthews, A. Gill, J. C. D. Gordon and D. W. Macdonald. 1999. Quantitative analysis of tonal calls from five odontocete species, examining interspecific and intraspecific variation. Journal of Zoology of London 249:403–410. 344. Leatherwood, S., T. A. Jefferson, J. C. Norris, W. E. Stevens, L. J. Hansen and K. D. Mullin. 1993. Occurrence and sounds of Fraser’s dolphins (Lagenodelphis hosei) in the Gulf of Mexico. The Texas Journal of Science 45:349–354. 345. Herzing, D. L. 1996. Vocalizations and associated underwater behavior of free-ranging Atlantic Spotted dolphins, Stenella frontalis and Bottlenose dolphins, Tursiops truncatus. Aquatic Mammals 22:61–79. 346. May-Collado, L. J., I. Agnarsson and D. Wartzok. 2007. Phylogenetic review of tonal sound production in whales in relation to sociality. BMC Evolutionary Biology 7:136. 347. Rankin, S., J. Oswald, J. Barlow and M. Lammers. 2007. Patterned burst-pulse vocalizations of the northern right whale dolphin, Lissodelphis borealis. Journal of the Acoustical Society of America 121:1213.

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Appendix S2.3

Thanks for Helping! As part of my Master’s I am investigating whether cetacean species that share particular evolutionary or ecological traits might be more likely to hybridize with one another. Part of doing this requires weighting different traits according to their likelihood of influencing interspecific mating. In order to come up with such a weighting, I am looking for the support of fellow biologists to provide their own opinion on the relative importance of these traits in promoting hybridization. I'm asking you to rate on a scale from 0-10 how important each trait might be in influencing hybridization. Species Traits:

Less 2 3 4 5 6 7 8 9 Very No Important Important Influence 1 10 0 Male Body Length (at physical maturity) Female Body Length (at physical maturity) Sexual Dimorphism (in colour or size) Preferred Water Depth Preferred Water Temperature Prey Species Predator Species Parasite Species Mean Group Size Species Range Size Cetacean Species Known to Interact With Shared Range Overlap Vocalization Frequency

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Table S2.4 Eigenvectors of the first four principal components of variation in similarity of traits for all cetacean species comparisons by taking the average of 10,000 subsampled principal component analyses where each species was only represented once. Variables that are more important for each principal component have larger values (+ or -).

Trait (all species) PC1 PC2 PC3 PC4 Male body length -0.0449 0.0065 -0.0066 0.0064 Female body length -0.0187 0.0341 0.0225 0.0032 Sexual Dimorphism -0.0161 0.0203 0.0203 0.0031 Range Size -0.0037 -0.0011 0.0014 0.0527 Water Depth 0.0213 -0.0049 0.0019 -0.0088 Water Temperature 0.0201 -0.0035 0.0015 -0.0057 Prey Species -0.0114 0.0033 -0.0058 0.0064 Predator Species -0.0009 0.0009 0.0021 0.0007 Parasite Species -0.0007 0.0006 0.0001 0.0029 Average Group Size -0.0067 0.0029 -0.0022 0.0029 Known Associate Species -0.0019 0.0011 0.0011 0.0008 Natural Range Overlap -0.0207 -0.0036 -0.0078 0.0036 Vocalization Frequency 0.0077 -0.0003 0.0056 0.0110

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Table S2.5 Eigenvectors of the first four principal components of variation in similarity of traits for cetacean species with 44 chromosomes by taking the average of 10,000 subsampled principal component analyses where each species was only represented once. Variables that are more important for each principal component have larger values (+ or -).

Trait (2n=44) PC1 PC2 PC3 PC4 Male body length 0.0160 -0.0025 -0.0032 -0.0020 Female body length 0.0179 -0.0019 -0.0019 -0.0044 Sexual Dimorphism -0.0414 0.0488 0.0297 -0.0045 Range Size -0.0289 0.0306 0.0308 -0.0021 Water Depth -0.0486 0.0134 0.0019 0.0070 Water Temperature -0.0151 -0.0116 -0.0078 0.0010 Prey Species -0.0019 -0.0004 0.0008 0.0034 Predator Species -0.0093 -0.0004 0.0049 0.0020 Parasite Species -0.0044 0.0006 0.0031 -0.0002 Average Group Size -0.0057 -0.0058 -0.0008 0.0694 Known Associate Species -0.0028 -0.0007 0.00003 0.0022 Natural Range Overlap -0.0151 -0.0089 -0.0026 0.0032 Vocalization Frequency 0.0025 -0.0074 -0.0055 0.0112

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