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Population structure in harbour porpoises (Phocoena phocoena) of British Columbia and widespread hybridization… Crossman, Carla Anne 2012

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Population structure in harbour porpoises (Phocoena phocoena) of British Columbia and widespread hybridization in cetaceans by Carla Anne Crossman BSc, Queen’s University, 2010  A THESIS SUBMITTED IN PARTIAL FUFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF  MASTER OF SCIENCE in The Faculty of Graduate Studies (Zoology)  THE UNIVERSITY OF BRITISH COLUMBIA (Vancouver) December 2012 © Carla Anne Crossman, 2012 i  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.  ii  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.  iii  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  2  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  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  iv  2.3.3 2.4  3  Discussion ..................................................................................................................... 34  2.4.1  Hybridization ....................................................................................................... 39  2.4.2  Contributions ........................................................................................................ 40  An Analysis of Cetacean Hybridization .......................................................................... 42 3.1  Introduction ................................................................................................................. 42  3.1.1 3.2  Research objectives .............................................................................................. 46  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  4  Microsatellites ....................................................................................................... 30  Discussion ..................................................................................................................... 60  3.4.1  Species barriers ..................................................................................................... 61  3.4.2  Potential benefits of interspecific mating ......................................................... 62  3.4.3  Conclusions ........................................................................................................... 63  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  v  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  vi  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  vii  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  viii  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 ix  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.  x  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.  1  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  2  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 3  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  4  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).  a  b  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.  7  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.  8  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).  9  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 fullterm 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, 10  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. 11  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. 12  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 13  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).  14  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  15  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 16  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  17  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 MgCl 2 (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  18  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  19  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. 20  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.  21  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.  22  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).  P  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.  23  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.  Q  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.  24  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  25  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.  26  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).  27  3 475  90  JX475289 JX47532 9 JX4754 05 JX475 3 89 JX47 5386 JX4 75 JX4 322 753 44 JX 4 7 JX4 5304 JX 75310 4 JX 7534 47 JX 53 5 JX 4753 28 JX 475 28 JX 475 364 JX 47 34 JX 47 538 6 47 52 2 52 93 98  Porpoise Dall's Finless Porpoise  1 3 7 32 7 5 5 3 91 7 4 7 2 JX X4 475 533 60 J X 47 53 5 J X 7 37 8 J X4 75 31 J X4 75 42 J X4 753 58 J X4 53 5 J X47 541 J X47 317 J X 47 5 J  03 53 09 47 53 9 JX 47 533 1 JX X47 531 3 J 47 41 JX 475 314 JX 475 87 3 5 JX 7 4 JX 02 754 JX4 75409 JX4 5412 7 JX4 5391 JX47 397 5 JX47 03 4 JX475 6 JX47541 JX475411 JX475349 JX475410 JX475320 JX475401 JX4753 JX475 48 JX4 306 JX4 75348 JX4 75401 JX 75351 JX 47538 JX 4753 3 JX 475 08 JX 475 347 JX 47 335 J 47 530 J X47 536 0 J X4 53 3 J X4 753 00 JXX47 753 00 9 4 75 529 4 35 0 6  JX  06 54 38 47 53 68 9 3 7 JX X4 475 537 4 J X 47 29 8 5 2 J JXX47 753 333 J X4 75 53 J X4 753 8 2 J 4 JXX47553294 J X47 340 J 75 JXX447536797 J 4753 JX 323 JX47755302 JX4 5361 JX47 JX475343 JX475378 JX475362 JX475373 JX475376 JX47529 4 JX 5 4 7 380 JX47 5 94 2 JX4 7 5 3 81 JX 4 754 19 J X 4 7 JX4 5341 7 JX 5359 4 JX 7 54 1 4 8 JX 7532 JX 4753 7 JX 4753 65 JX 475 72 JX 475 396 JX 47 30 JX 47 539 3 5 4 75 40 5 40 8 4  16 53 366 7 4 5 13 JX X47 53 93 J 47 53 2 JX 47 539 JX 47 367 JX 475 331 JX 475 21 JX 4753 5 JX 7532 JX4 75336 JX4 75326 JX4 5296 JX47 5323 JX47 23 3 JX475 4 7 JX4753 JX475385 JX475324 JX475414 JX475305 JX475299 JX475357 JX475 JX47 292 JX47 5417 JX4 5400 JX4 75400 JX 75398 JX4475354 JX 753 JX 475 99 JX 475 301 JX 47 307 J 47 531 JXX47 531 2 J 4 53 5 J X4 75 34 JX X47 753 294 47 52 52 53 95 70  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 timereversible 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 phocoenoides) and Dall’s porpoise were used as outgroups.  28  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 -0.70 100.70 -0.00705 0.68±0.016 Two Sampling 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 -  29  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 30  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  Mean LnP(K)  -3000  -3200  -3400  -3600  2  4  6  8  10  K  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).  31  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.  32  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 0.25 99.75 0.0025 0.25 ± 0.015 Two Sampling 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).  33  0.7 0.6 0.5 0.4 0.3 0.2 0.0  0.1  Proportion of alleles  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 F ST < 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  34  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 -  35  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; BarrettLennard 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 36  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 (EscorzaTreviño 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  37  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.  38  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  39  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  40  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).  41  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 42  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  43  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 44  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.  45  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 postzygotic, 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 46  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 Max1 or Max2 ) (Larger of Min1 or Min2 ) Smaller of (Max1 Min1 or Max2 Min2 )  47  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: 10,000,000100,000,000 km2; large: >100,000,000 km2), water depth (shallow: 0-200 m; medium: 2001000 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 (  )  48  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  49  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.  50  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 rvalues 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.  51  Non-hybridizing species pairs  Hybridizing 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.  52  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  Natural Range Overlap  0.021  -0.387  -0.577  -0.461  Vocalization Frequency  -0.239  -0.283  0.383  0.413  Species  Proportion of Variation  24.32% 19.76% 14.63% 10.09%  Accounted For  53  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 species pairs  Hybridizing 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. 54  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  Natural Range Overlap  -0.034  -0.251  -0.630  0.360  Vocalization Frequency  -0.352  -0.310  0.342  -0.495  Species  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  55  (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)  56  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  57  Non-hybridizing species pairs  Hybridizing 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.  58  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  26.03%  21.71%  16.64%  10.66%  Proportion of Variation Accounted For  59  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 60  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  61  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  62  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.  63  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. 64  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 subpopulations 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  65  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 66  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.  67  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. 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Canadian Journal of Zoology 81:1755–1762.  84  Appendices A.1 Appendix S1 – Chapter 2 Table S1.1 Case numbers and additional organizations associated with each sample Species  DFO  AHC  Dall's Porpoise  AHC 00-3075  Dall's Porpoise  AHC 02-3166  Dall's Porpoise  AHC 04-1780  Dall's Porpoise  AHC 11-3920  Dall's Porpoise  DFO 1075  AHC 03-2768  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  Dall's Porpoise  DFO 4631  Dall's Porpoise  DFO 4838  Dall's Porpoise  DFO 5563  AHC 02-2110  LBL  Other Case Number  Other Org.  04-SJ004 LBL 03-14  AH 03OBDM06  LBL 02-18 LBL 02-19  AHC 09-01909  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, 97-05  From AHC  Dall's Porpoise  LBL 00-09  Ron Lewis 98-405, 97-10  From AHC  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 97-11  From AHC  LBL 00-13  Ron Lewis 98-1468  From AHC  LBL 00-14  Ron Lewis 98-1473  From AHC  Dall's Porpoise  DFO 3763  Dall's Porpoise Dall's Porpoise  AHC 02-02110  LBL 02-18  Dall's Porpoise  DFO 4287  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  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  Dall's Porpoise  DFO 2910  AHC 07-2879  LBL 07-10  030WCPPF10  FOS 3056 CR  85  Species  DFO  AHC  LBL  Other Case Number  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  Dall's Porpoise  LBL 98-17A  Dall's Porpoise  LBL 99-02  Other Org.  G. Ellis 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  Harbour Porpoise  AHC 03-2356  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  Harbour Porpoise  AHC 08-1592  Harbour Porpoise  AHC 09-01252  Harbour Porpoise  DFO 5026  LBL 02-28 2002SJ-006  06Pp21NovW1-06  Vancouver Aquarium  09-0SJ001  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  Harbour Porpoise  AHC 10-4951  DFO 5733  Harbour Porpoise  AHC 10-4952  DFO 5737  LBL 09-07  86  Species  DFO  AHC  Harbour Porpoise  AHC 10-4953  Harbour Porpoise  AHC 10-514  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  LBL  DFO 6627  AHC 11-3143  Harbour Porpoise  DFO 6779  AHC 11-3921  Harbour Porpoise  Other Org.  09Pp07AugSK-01  AHC 11-2487  Harbour Porpoise  Other Case Number  11Pp01JanWI-01  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  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  Harbour Porpoise  DFO 3340  AHC 08-02977  Harbour Porpoise  DFO 3399  AHC 08-3529  Harbour Porpoise  DFO 3406  AHC 08-03815  Harbour Porpoise  DFO 3410  AHC 08-03531  B. Hanson  LBL 08-25 LBL 08-22 AH 08PpEsqM06 LBL 08-18  87  Species  DFO  AHC  LBL  Other Case Number  Harbour Porpoise  DFO 3519  AHC 02-02112  LBL 02-16  AH 02PpM02  Harbour Porpoise  DFO 3539  Harbour Porpoise Harbour Porpoise  DFO 3599A? DFO 3599B  Harbour Porpoise  DFO 3639  Harbour Porpoise  DFO 3691  Harbour Porpoise  SWDP 93-46, HP93-15 AHC 02-02115  LBL 02-12  AH 02PhF08  AHC 02-02115  LBL 02-33  AH 02PpF08  Other Org. SWDP  SWDP 94-08, HP94-01A  SWDP  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  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  Harbour Porpoise  DFO 3974  Harbour Porpoise  DFO 3993  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  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  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, Pp92-1  DFO 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  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  AHC 01-03253  LBL 98-08  HP98-02B  M753-2 SWDP 94-09, HP94-02A  AHC 01-04520  SWDP  AH 01JdFM20  AHC 98-00445  379, AH 01HSPP002  AHC 01-03741  LBL 03-21  88  Species  DFO  AHC  Harbour Porpoise  DFO 4661  AHC 01-3813  Harbour Porpoise  LBL  Other Case Number  Other Org.  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  Harbour Porpoise  DFO 5556  Harbour Porpoise  DFO 5566  AHC 04-01950  Harbour Porpoise  DFO 5649  AHC 10-01948  LBL 10-12  Harbour Porpoise  DFO 5651  AHC 10-01946  LBL 10-10  Harbour Porpoise  DFO 5656  AHC 10-01952  LBL 10-16  Harbour Porpoise  DFO 5723  Harbour Porpoise  DFO 5733  Harbour Porpoise  DFO 5738  Harbour Porpoise  DFO 6292  Harbour Porpoise  DFO 6443  Harbour Porpoise  DFO 6445  AHC 09-03814  RR 09-0254  AH 16PpRRF10, RR 100084 AH 20PpVicF10  AHC 10-4591 AHC 02-01779  SJ057 SWDP94-?, HP94-10, 172  SWDP B. Hanson  Harbour Porpoise Harbour Porpoise  LBL 00-02  SIRS  Vancouver Aquarium SIRS  Harbour Porpoise  LBL 00-16  Ron Lewis 98-406  T. Guenther  Harbour Porpoise  LBL 00-17  T. Guenther  Harbour Porpoise  LBL 00-18  Ron Lewis 98-406, SWDP 97-08 Ron Lewis 98-408  LBL 00-19  Ron Lewis 98-409, SWDP 94-06  From AHC  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 97-13 Ron Lewis 98-417 SWDP 95-27  From AHC  Harbour Porpoise  Harbour Porpoise  JackVA  DFO 4656  DFO 4565  LBL 00-33  T. Guenther  From AHC  89  Species  DFO  AHC  Harbour Porpoise  LBL  Other Case Number  LBL 02-09  Other Org. 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  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  Vancouver Aquarium  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  90  Species  DFO  AHC  LBL  Harbour Porpoise  LBL 98-14  Harbour Porpoise  LBL 98-15  Harbour Porpoise  LBL 98-16  Harbour Porpoise  LBL 98-17B  Harbour Porpoise  LBL 98-19  Other Case Number  Other Org. Nitnat Fishery Area 12-1 test fishery  Ron Lewis #97/003539 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  Porpoise  LBL 07-11  WDFW 1207-04  LBL 00-39  Ron Lewis 98-444, SWDP 97-16  Porpoise  AHC 02-02114  LBL 02-11  AH 02PpM06  Porpoise  AHC 02-02113  LBL 02-15  AH 02PpF05  AHC 07-3624  LBL 07-09  Porpoise  DFO 3027(8)  T. Guenther  Vancouver 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  91  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 101.82 -0.01818 0.82±0.011 Three sampling -1.82 groups -0.99 100.99 -0.00991 0.75±0.013 Two sampling 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  -  92  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 99.75 0.00247 0.25±0.015 Two sampling groups 0.25  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 0.00078  Outside/North 0.35±0.016 -  93  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  Mean LnP(K)  -2700  -2800  -2900  -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.  94  Table S1.10 Coordinate data used in Geneland Sample  Latitude  Longitude  Sample  Latitude  Longitude  Sample  Latitude  Longitude  AHC 03-2356 AHC 06-02388 AHC 06-1962 AHC 06-3276 AHC 07-2921 AHC 09-01252 AHC 09-02921 AHC 09-3238 AHC 10-01596 AHC 10-0514 AHC 10-0515 AHC 10-0516 AHC 10-1618 AHC 10-1907 AHC 10-4951 AHC 11-150 AHC 11-1945 AHC 11-1946 AHC 11-1987 AHC 11-2196 AHC 11-2197 AHC 11-2198 AHC 11-2199 AHC 11-2200 AHC 11-2201 AHC 11-2487 AHC 11-3143 AHC 11-3921 AHC 11-930 AHC 12-1018 AHC 12-1022 AHC 12-1023 AHC 2011-1641 AHC 2011-1642 AHC 2011-1643 Birch Bay 2006 DFO 1071 DFO 1081 DFO 1833 DFO 2227 DFO 2246 DFO 2250 DFO 2400 DFO 2787 DFO 2788 DFO 2820 DFO 3340 DFO 3399 DFO 3406 DFO 3410 DFO 3519  48.454 48.071 48.418 46.928 47.972 48.454 49.848 47.241 47.244 48.318 48.198 48.744 50.127 47.607 48.751 49.580 48.412 48.406 48.756 48.418 48.431 48.432 48.407 48.411 48.425 47.966 48.306 49.360 49.021 49.169 48.422 49.008 49.357 48.378 48.408 48.915 49.286 49.273 49.247 48.414 48.419 48.826 48.417 50.683 50.684 49.152 49.229 49.293 48.413 48.409 48.424  -122.937 -123.027 -123.415 -124.175 -122.478 -123.012 -124.534 -124.220 -124.219 -122.375 -122.354 -122.720 -125.360 -122.350 -123.413 -124.797 -123.381 -123.362 -123.445 -123.468 -123.447 -123.429 -123.339 -123.313 -123.430 -122.476 -123.543 -124.442 -123.100 -123.935 -123.421 -123.119 -124.437 -123.520 -123.367 -122.758 -123.216 -123.269 -123.262 -123.388 -123.389 -123.441 -123.392 -126.695 -126.691 -125.914 -123.243 -126.037 -123.384 -123.378 -123.383  DFO 3539 DFO 3599B DFO 3639 DFO 3742 DFO 3819 DFO 3897 DFO 3919 DFO 3964 DFO 3974 DFO 3993 DFO 4007 DFO 4024 DFO 4064 DFO 4065 DFO 4098 DFO 4104 DFO 4126 DFO 4239 DFO 4240 DFO 4242 DFO 4273 DFO 4298 DFO 4327 DFO 4345 DFO 4346 DFO 4448 DFO 4494 DFO 4497 DFO 4635 DFO 4697 DFO 4723 DFO 4788 DFO 4819 DFO 4838 DFO 5206 DFO 5566 DFO 5649 DFO 5651 DFO 5738 DFO 6445 JackVanAqua LBL 00-02 LBL 00-16 LBL 00-17 LBL 00-18 LBL 00-20 LBL 00-22 LBL 00-23 LBL 00-25 LBL 00-26 LBL 00-27  49.008 48.426 48.410 48.798 48.417 53.654 48.405 48.421 49.920 54.138 48.645 49.019 48.406 49.804 48.417 48.407 48.417 49.262 49.198 49.671 49.179 48.330 48.369 49.048 53.313 48.351 48.350 50.027 48.408 48.428 49.662 48.417 50.779 48.408 49.276 48.299 48.410 48.811 50.029 48.235 49.376 49.069 48.410 48.410 48.457 48.370 48.419 49.069 48.437 48.384 49.509  -123.093 -123.382 -123.326 -123.202 -123.370 -131.905 -123.355 -123.396 -125.184 -130.279 -123.803 -122.808 -123.353 -124.527 -123.317 -123.345 -123.400 -126.230 -123.963 -124.934 -122.554 -123.617 -123.712 -125.726 -132.789 -123.531 -123.530 -125.239 -123.336 -123.473 -124.928 -123.300 -126.700 -123.341 -123.170 -123.533 -123.381 -123.602 -125.243 -123.567 -123.274 -125.755 -123.341 -123.340 -123.293 -123.717 -123.419 -125.755 -123.291 -123.515 -124.245  LBL 00-30 LBL 00-31 LBL 00-33 LBL 02-11 LBL 02-15 LBL 03-15 LBL 05-30 LBL 07-07 LBL 07-08 LBL 07-09 LBL 07-12 LBL 08-19 LBL 08-23 LBL 09-11 LBL 10-04 LBL 10-05 LBL 10-08 LBL 10-09 LBL 10-13 LBL 10-14 LBL 10-15 LBL 10-17 LBL 10-18 LBL 10-19 LBL 10-20 LBL 10-21 LBL 10-22 LBL 10-23 LBL 10-24 LBL 10-25 LBL 10-29 LBL 98-08 LBL 98-09 LBL 98-14 LBL 98-19 SWDP 96-11 SWDP 97-09  49.647 48.450 48.419 48.373 48.416 48.403 48.420 50.286 49.254 50.590 49.021 48.333 48.254 48.746 48.798 48.428 48.407 48.405 48.300 48.415 48.416 48.373 49.129 49.155 49.155 49.020 48.554 48.406 47.651 48.408 49.017 53.696 48.668 48.666 49.268 48.357 48.352  -126.848 -122.963 -123.387 -123.715 -123.398 -123.348 -123.421 -125.350 -124.072 -127.101 -122.806 -123.547 -123.393 -123.264 -123.243 -123.394 -123.368 -123.359 -123.534 -123.409 -123.406 -123.715 -125.904 -125.909 -125.905 -122.807 -124.420 -123.365 -122.295 -123.372 -122.794 -131.871 -124.862 -124.854 -126.142 -123.822 -123.544  95  Table S1.9 Locations of 112 mtDNA haplotypes resolved by sequencing 545 base pairs of d-loop from 134 harbour porpoises. The localities “inside”, “outside” and “northern” water are shown in Fig. 2.1. Haplotype (Genbank Accession)  Northern Waters (5)  Outside Waters (14)  Inside Waters (109)  JX475334 JX475335 JX475353 JX475380 JX475407 JX475304 JX475306 JX475340 JX475355 JX475328 JX475362  1 1 1 1 1 0 0 0 0 0 0  0 0 0 0 0 1 1 1 1 1 1  0 0 0 0 0 0 0 0 0 3 0  JX475373 JX475294  0 0  1 1  0 4  JX475376 JX475383 JX475387 JX475406 JX475411 JX475413 JX475290 JX475295 JX475305 JX475313 JX475317 JX475323 JX475319 JX475332  0 0 0 0 0 0 0 0 0 0 0 0 0 0  1 1 1 1 1 1 0 0 0 0 0 0 0 0  0 0 0 0 0 0 1 1 1 1 1 3 1 1  JX475338 JX475343 JX475357 JX475384 JX475388 JX475392 JX475405 JX475401 JX475417  0 0 0 0 0 0 0 0 0  0 0 0 0 0 0 0 0 0  1 1 1 1 1 1 1 2 1  Haplotype (Genbank Accession)  Northern Waters (5)  Outside Waters (14)  Inside Waters (109)  JX475419  0  0  1  JX475296 JX475301 JX475302 JX475307 JX475310 JX475312 JX475314 JX475315 JX475322  0 0 0 0 0 0 0 0 0  0 0 0 0 0 0 0 0 0  1 1 1 1 1 1 2 1 1  JX475325 JX475326 JX475327 JX475333 JX475336 JX475344 JX475361 JX475402 JX475410 JX475414 JX475348 JX475298 JX475299 JX475300  0 0 0 0 0 0 0 0 0 0 0 0 0 0  0 0 0 0 0 0 0 0 0 0 0 0 0 0  1 1 1 1 1 1 1 1 1 1 2 1 1 3  JX475303 JX475308 JX475309  0 0 0  0 0 0  2 1 1  JX475311  0  0  1  JX475316 JX475321 JX475331 JX475339 JX475346 JX475350 JX475352 JX475354  0 0 0 0 0 0 0 0  0 0 0 0 0 0 0 0  1 1 1 1 1 1 1 1  JX475356 JX475359 JX475369 JX475370 JX475372 JX475374 JX475377  0 0 0 0 0 0 0  0 0 0 0 0 0 0  1 1 1 1 1 1 1  96  Haplotype (Genbank Accession)  Northern Waters (5)  Outside Waters (14)  Inside Waters (109)  JX475378 JX475379 JX475395 JX475396 JX475404 JX475408 JX475409 JX475412 JX475418 JX475291 JX475318 JX475347 JX475349 JX475351 JX475360 JX475363 JX475371 JX475375 JX475391 JX475403 JX475415 JX475416 JX475337 JX475342 JX475394 JX475400 JX475397 JX475398 JX475399 JX475324 JX475292 JX475293  0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0  0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0  1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 2 1 1 1 1 1 1  JX475320  0  0  1  97  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  Water Depth  Average Group Size  No  Species' Range Size Medium  NA  NA  NA  9.33  No  Medium  Medium  Medium  NA  10  12.8  Yes  Medium  Deep  Medium  Moridae, Gonatidae, Macrouridae, Cradchiidae  4.15  3.89  3.98  Yes  Large  Deep  Solitary  Cepolidae, Melamphaidae, Myctophidae  5.5  6.93  5.27  7.54  Yes  Large  Deep  Solitary  Histioteuthidae, Gonatidae, Cranchiidae, Ommastrephidae, Vampyroteuthidae, Bolitaenidae, Stauroteuthidae  Gervais' Beaked Whale (Mesoplodon europaeus) Ginkgo-Toothed Beaked Whale (Mesoplodon ginkgodens) Gray's Beaked Whale (Mesoplodon grayi) Hector's Beaked Whale (Mesoplodon hectori) Hubb's Beaked Whale (Mesoplodon carlhubbsi)  3.54  4.56  NA  NA  Yes  Medium  Deep  Solitary  Stomiidae, Octopoteuthidae, Lophogastridae  NA  NA  NA  NA  Yes  Large  NA  NA  NA  NA  NA  NA  Yes  Medium  Deep  Solitary  Merluccidae, Phosichthyidae, Myctophidae  3.65  4.34  NA  NA  Yes  Medium  NA  Solitary  Octopoteuthidae  4.96  5.3  4.9  5.32  Yes  Medium  NA  NA  Pygmy Beaked whale (Mesoplodon peruvianus) Longman's Beaked Whale (Mesoplodon pacificus) Northern Bottlenose Whale (Hyperoodon ampullatus) Southern Bottlenose Whale (Hyperoodon planifrons) Sowerby's Beaked Whale (Mesoplodon bidens)  3.26  3.72  NA  NA  Yes  Medium  NA  Solitary  NA  NA  NA  NA  Yes  Large  Deep  Solitary  7.3  9.8  6  8.7  Yes  Medium  Deep  Solitary  NA  6.94  5.7  7.45  Yes  Medium  Deep  Solitary  4.09  5.5  4.1  5.1  No  Medium  Shallow  Solitary  Andrew's Beaked Whale (Mesoplodon bowdoini) Arnoux's Beaked Whale (Berardius arnuxii) Baird's Beaked Whale (Berardius bairdii) Blainville's Beaked Whale (Mesoplodon densirostris) Cuvier's Beaked Whale (Ziphius cavirostris)  Length MaleMin  Length MalMax  Length FemMin  Length FemMax  Sexually Dimorphic  3.9  4.4  3.94  4.87  8  9.34  8.84  9.5  11.9  3.49  Prey  NA  Gonatidae, Mastigoteuthidae, Melamphaidae, Onychoteuthidae, Histioteuthidae, Myctophidae, Stomiidae, Octopoteuthidae Myctophidae, Nemipteridae Cranchiidae, Onychoteuthidae, Chiroteuthidae, Histioteuthidae Gonatidae, Myopsidae, Oegposidae, Clupeidae, Gadidae Cranchiidae, Onychoteuthidae, Enoploteuthidae, Neoteuidae, Psychroteuthidae, Gonatidae Gadidae, Merluccidae  98  Species  Length MaleMin  Length MalMax  Length FemMin  Length FemMax  Sexually Dimorphic  Water Depth  Average Group Size  Yes  Species' Range Size Medium  Straptoothed Whale (Mesoplodon layardii)  4.87  5.5  5  6.25  Stejneger's Beaked Whale (Mesoplodon stejnegeri) Tasman Beaked Whale (Tasmacetus shepherdi) True's Beaked Whale (Mesoplodon mirus) Atlantic Humpbacked Dolphin (Sousa teuszii) Atlantic Spotted Dolphin (Stenella plagiodon/frontalis) Atlantic White-Sided Dolphin (Lagenorhynchus acutus) Black Dolphin (Cephalorhynchus eutropia) Bottlenose Dolphin (Tursiops truncatus)  3.89  5.3  4.34  5.96  7.35  4.6  Prey  NA  Solitary  5.24  No  Medium  Deep  Solitary  Vampyroteuthidae, Chiroteuthidae, Cranchiidae, Cycloteuthidae, Gonatidae, Histioteuthidae, Mastigoteuthidae, Octopoteuthidae, Ommastrephidae, Onchoteuthidae Gonatidae, Cranchiidae  NA  NA  No  Medium  Deep  Solitary  Merlucciidae, Serranidae, Bythitidae  5.3  4.87  5.18  Yes  Medium  NA  NA  2  2.48  NA  2.35  Yes  Small  Shallow  Medium  Haemulidae, Clupeidae, Mugilidae  1.66  2.26  1.67  2.29  Yes  Medium  Shallow  Medium  2.44  2.75  1.94  2.43  Yes  Medium  Shallow  Medium  Gadidae, Clupeidae, Carangidae, Sciaenidae, Congridae, Trichiyridae, Triglidae Loliginidae, Ammodytidae, Osmeridae, Scombridae  1.24  1.65  1.23  1.61  Yes  Small  Shallow  Medium  Munididae, Loliginidae  2.02  3.81  1.9  3.67  Yes  Large  Medium  Medium  Clymene Dolphin (Stenella clymene) Commerson's Dolphin (Cephalorhynchus commersonii)  1.76  1.97  1.71  1.83  Yes  Medium  Deep  Social  Engraulidae, Apongidae, Trichiuridae, Synodontidae, 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 Myctophidae  1.3  1.67  1.39  1.74  Yes  Small  Shallow  Solitary  Short Beaked Common Dolphin (Delphinus delphis)  1.71  2.6  1.67  2.44  Yes  Medium  Medium  Medium  Dusky Dolphin (Lagenorhynchus obscurus)  1.67  2.11  1.67  2.05  No  Medium  Shallow  Medium  Cranchiidae, Loliginidae  Merlucciidae, Atherinopsidae, Lithodidae, Loliginidae, Rhodomelaceae, Laminariaceae, Sphacelariaceae , Ceramiaceae, Sertulariidae, Nereidae, Mysidae, Euphausiidae, Sphaeromatidae, Diastylidae, Ophiomixidae, Styelidae, Clupeidae, Gnathiidae, Sphaeromatidae, Cirolanidae, Pyuridae Argentinidae, Bathylagidae, Batrachoididae, Melamphaidae, Scomberesocidae, Myctophidae, Sciaenidae, Engraulidae, Merlucciidae, Ophidiidae, Scombridae, Stromateidae, Loliginidae, Onychoteuthidae Engraulidae  99  Species  Length MaleMin  Length MalMax  Length FemMin  Length FemMax  Sexually Dimorphic  False Killer Whale (Pseudorca crassidens)  3.96  6.1  3.4  5.06  Fraser's Dolphin (Lagenodelphis hosei)  2.31  2.7  2.06  2  3.2  1.51  Indo-Pacific Humpbacked Dolphin (Sousa chinensis) Heaviside's Dolphin (Cephalorhynchus heavisidii) Hector's Dolphin (Cephalorhynchus hectori) Hourglass Dolphin (Lagenorhynchus cruciger) Irrawaddy Dolphin (Orcaella brevirostris)  Killer Whale (Orcinus orca)  Long Beaked Common Dolphin (Delphinus capensis)  Water Depth  Average Group Size  Yes  Species' Range Size Large  Prey  Deep  Medium  2.64  Yes  Large  Deep  Social  2.16  2.44  Yes  Medium  Shallow  Medium  1.74  1.59  1.66  No  Small  Shallow  Solitary  1.17  1.38  1.28  1.53  Yes  Small  Shallow  Solitary  Mugilidae, Uranoscopidae, Moridae, Ommastrephidae, Arripidae  1.63  1.87  1.66  1.83  No  Small  Deep  Medium  Mycophidae  2.2  2.35  2  2.32  Yes  Small  Shallow  Solitary  5.2  9.75  4.57  8.53  Yes  Large  Medium  Medium  2  2.4  1.9  2.2  Yes  Small  Shallow  Social  Cyprinidae, Teraponidae, Apogonidae, Chirocentridae, Pangasiidae, Cuttlefish, Engraulidae, Clupeidae, Synodontidae, Hemirhamphidae, Psettodidae, Leigignathidae, Nemipteridae, Pomadasyidae, Sillaginidae, Platycephalidae Eschrichtiidae, Ohyseteridae, Balaenopteridae, 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 Engraulidae, Myctophidae, Phosichthyidae, Atherinopsidae, Meriuccidae, Clupeidae, Centrolophidae, Carangidae, Scombridae, Triglidae, Scomberesocidae, Sphyraenidae, Normanichthyidae, Batrachoididae, Ophichthidae, Galatheidae  Lycoteuthidae, Ommastrephidae, Scombridae, Coryphaenidae, Sciaenidae, Lateolabracidae, Salmonidae, Ariidae, Gadidae, Gonatidae, Cranchiidae Onychoteuthidae, Ophichthyidae, Scomberoscocidae, 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 Mugilidae, Engraulidae, Pristigasteridae, Haemulidae, Sparidae, Callichthyidae, Sciaenidae, Trichiuridae Merlucciidae, Ophidiidae, Gobiinae, Loliginidae  100  Species  Length MaleMin  Length MalMax  Length FemMin  Length FemMax  Sexually Dimorphic  Water Depth  Average Group Size  Prey  Yes  Species' Range Size Large  5  6.1  4.05  4.72  Medium  Medium  2.57  Yes  Medium  Deep  Social  1.95  2.3  Yes  Medium  Deep  Social  2.5  1.7  2.36  Yes  Medium  Medium  Medium  Octopodidae, Gonatidae, Ommastreohidae, Mastigoteuthidae, Loligoinidae, Argentinidae, Gadidae, Pleuronectidae, Macrouridae, Zoarcidae, Ammoditidae, Trichiuridae, Pandalidae, Galatheidae, Chiroteuthidae, Brachioteuthidae, Sepiolidae, Cranchiidaae, Histioteuthidae Ommastrephidae, Histioteuthidae, Loliginidae, Onycoteuthidae, Chiroteuthidae, Mastigoteuthidae, Cranchiidae, Enoploteruthidae, Myctophidae, Paralepididae, Scopelarchidae Gonatidae, Onychoteuthidae, Loligindae, Enoploteuthidae, Histioteuthidae, Centrolophidae, Melamphaidae, Scomberesocidae, Merlucciidae, Myctophidae, Bathylagidae, Paralepididae Engraulidae, Merlucciidae, Loliginidae  Melon-Headed Whale (Peponocephala electra)  2.05  2.64  2.11  Northern Right Whale Dolphin (Lissodelphis borealis)  2.11  3.1  Pacific White-Sided Dolphin (Lagenorhynchus obliquidens) Peale's Dolphin (Lagenorhynchus australis) Pygmy Killer Whale (Feresa attenuata) Risso's Dolphin (Grampus griseus)  1.7 1.6  2.18  1.63  2.1  No  Small  Shallow  Solitary  2.07  2.59  2.07  2.45  No  Large  Medium  Medium  2.53  3.6  2.4  3  No  Small  Medium  Medium  Rough-Toothed Dolphin (Steno bredanensis)  2.09  2.65  2.12  2.55  Yes  Large  Medium  Medium  Short-Finned Pilot Whale (Globicephala macrorhynchus)  4.24  4.91  3.34  3.92  Yes  Medium  Medium  Social  Southern Right Whale Dolphin (Lissodelphis peronii)  NA  NA  NA  NA  Yes  Medium  Medium  Social  Long-Finned Pilot Whale (Globicephala melaena)  Myxinidae, Ommastrephidae, Clupeidae, Moridae, Merlucciidae, Octopodidae, Loliginidae Delphinidae Sepiidae, Brachioteuthidae, Cranchiidae, Mastigoteuthidae, Onychoteuthidae, Histioteuthidae, Ommastrephidae, Argonautidae, octopodidae, Ascidiacea, Pyrosomidae, Salpidae, Enoploteuthidae, Chiroteuthidae, Loliginidae, Sepiolidae Tremoctopodidae, Onychoteuthidae, Coryphaenidae, Trichiuridae, Atherinopsidae, Scomberesocidae, Belonidae, Atherinopsidae, Centriscidae, Coryphaenidae, Ommastrephidae, Atherinidae, Loliginidae Octopodidae, Enoploteuthidae, Histioteuthidae, Loliginidae, Mastigoteuthidae, Chiroteuthidae, Cranciidae, Brachioteuthidae, Lepidoteuthidae, Ommastrephidae, Melamphaidae Ommastrephidae, Mastigoteuthidae, Cranchiidae, Gonatidae, Bathylagidae, Photichthyidae, Myctophidae, Merlucciidae, Engraulididae  101  Species  Length MaleMin  Length MalMax  Length FemMin  Length FemMax  Sexually Dimorphic  Water Depth  Average Group Size  Prey  Yes  Species' Range Size Large  Spinner Dolphin (Stenella longirostris)  1.36  2.35  1.29  2.04  Deep  Social  1.63  Yes  Large  Deep  Social  Mastigotruthidae, Congridae, Stomiidae, Paralepididae, 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 NA  Pan tropical Spotted Dolphin (Stenella attenuata) Striped Dolphin (Stenella coeruleoalba)  1.66  2.57  1.63  2.15  2.56  1.85  2.36  Yes  Large  Deep  Medium  Guiana dolphin (Sotalia guianensis) White-Beaked Dolphin (Lagenorhynchus albirostris) Beluga (Delphinapterus leucas) Narwhal (Monodon monoceros) Franciscana (Pontoporia blainvillei)  1.31  1.87  1.38  2.06  No  Small  Shallow  Solitary  2.51  3.1  1.74  2.78  Yes  Medium  Shallow  Medium  3.5  4.7  3.1  3.9  Yes  Medium  Medium  Solitary  4.1  4.7  3.4  4.15  Yes  Small  Deep  Solitary  1.21  1.58  1.37  1.74  Yes  Small  Shallow  Medium  Burmeister's Porpoise (Phocoena spinipinnis)  1.51  1.75  1.53  1.85  Yes  Small  Shallow  Solitary  Ateleopodidae, Myctophidae, Microstomatidae, Melamphaeidae, Bathylagidae, Gempylidae, Brachioteuthidae, Cranchiidae, Cycloteuthidae, Enoploteuthidae, Grimalditeuthidae, Histioteuthidae, Mastigoteuthidae, Octopoteuthidae, Ommastrephidae, Onychoteuthidae, Pholidoteuthidae, Tremoctopodidae, Sternoptychidae, Nomeidae, Paralepididae, Phosichthyidae, Scopelarchidae Clupeidae, Scianidae, Batrachoididae, Trichiuridae, Loliginidae Gadidae, Clupeidae, Osmeridae Osmeridae, Clupeidae, Ammodytidae, Cyclopteridae, Characidae, Osmeridae, Salmonidae, Gadidae, Illicinae Gadidae, Pleuronectidae, Salmonidae, Clupeidae Gobiidae, Penaeidae, Ophidiiae, Cynoglossidae, Trichiuridae, Carangidae, Phycidae, Antherinidae, Poatmidae, Engraulidae, Sciaenidae, Loliginidae, Batrachoidiae, Gadidae, Stromatidae, Congridae, Stromateidae, Nomeidae Merlucciidae, Engraulidae, Loliginidae, Euphausiidae, Centrolophidae, Carangidae, Clupeidae, Congridae, Centropomidae, Atherinopsidae, Sciaenidae, Gadidae, Sparidae  102  Species  Cochito [Vaquita] (Phocoena sinus)  Length MaleMin  Length MalMax  Length FemMin  Length FemMax  Sexually Dimorphic  1.27  1.44  1.35  1.48  Yes  Species' Range Size Small  Water Depth  Average Group Size  Shallow  Solitary  Prey  Sciaenidae, Haemulidae  Dall's Porpoise (Phocoenoides dalli)  1.75  1.8  1.74  1.77  Yes  Medium  Medium  Medium  Bolitaenidae, Enoploteuthidae, Gonatidae, 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 Loliginidae, Apogonidae, Leiognathidae, Sepiidae, Engraulidae Octopodidae, Clupeidae, Gadidae, Gobiidae, Anguillidae, Gonostomatidae, Marlucciidae, Sparidae, Zoarcidae, Ammodytidae, Pleuronectidae, Sebastidae, Illicinae, Sternoptychidae, Merlucciidae, Stromateidae, Scombridae, Sepiolidae, Myxinidae, Loliginidae Engrauliidae  Finless Porpoise (Neophocaena phocaenoides) Harbour Porpoise (Phocoena phocoena)  1.32  2.27  1.32  2.06  Yes  Medium  Shallow  Solitary  1.23  1.6  1.38  1.7  Yes  Medium  Shallow  Solitary  Spectacled Porpoise (Phocoena dioptrica) Gray whale (Eschrichtius robustus) Blue Whale (Balaenoptera musculus) Bryde's Whale (Balaenoptera edeni) Fin Whale (Balaenoptera physalus) Antarctic Minke Whale (Balaenoptera bonaerensis) Common Minke Whale (Balaenoptera acutorostrata)  1.89  2.24  1.74  2.04  Yes  Small  Deep  Solitary  11.1  14.6  11.7  15  Yes  Small  Shallow  Solitary  20  25  21  33.6  Yes  Large  Medium  Solitary  Pachychilidae, Pinnotheridae, Galatheidae, Nephropidae, Atylidae, Ampeliscidae Euphausiidae, Temoridae  12  14.2  13.7  15.5  Yes  Medium  Medium  Solitary  Clupeidae, Engraulidae, Euphausiidae, Carangidae  17.7  25  18.3  27  Yes  Large  Medium  Medium  7.3  8.6  7.9  9  Yes  Medium  Deep  Solitary  6.7  8.2  7.2  8.8  Yes  Medium  Shallow  Solitary  Sei Whale (Balaenoptera borealis)  12.8  15.9  13.3  16.1  Yes  Large  Deep  Solitary  Ommastrephidae, Euphausiidae, Osmeridae, Calanidae, Clupeidae, Gadidae Channichthyidae, Myctophidae, Paralepididae, Nototheniidae, Euphausiidae, Hyperiidae Euphausiidae, Gadidae, Salmonidae, Ammodytidae, Clupeidae, Osmeridae, Scombridae, Anarhichadidae, Squalidae, Merlucciidae Clausocalanidae, Hyperiidae, Temoridae, Euphausiidae, Eucalanidae, Metridinidae, Calanidae, Engraulidae, Clupeidae, Scomberesocidae, Myctophidae  103  Species  Length MaleMin  Length MalMax  Length FemMin  Length FemMax  Sexually Dimorphic  Water Depth  Average Group Size  Yes  Species' Range Size Large  Humpback Whale (Megeptera novaeangliae) Bowhead Whale (Balaena mysticetus) North Pacific Right Whale (Eubalaena japonica) North Atlantic Right Whale (Eubalaena glacialis) Southern Right Whale (Eubalaena australis) Pygmy Right Whale (Caperea marginata) Indo-pacific bottlenose dolphin (Tursiops aduncus)  12  14.8  13.9  15.5  11.6  15.5  14  15  17.1  11  Prey  Deep  Solitary  18  Yes  Small  Medium  Solitary  Mysidae, Euphausiidae, Pandalidae, Clupeidae, Osmeridae, Gadidae Euphausiidae, Calanidae  15.5  18.3  Yes  Medium  Shallow  Solitary  Euphausiidae, Calanidae  12.9  11  18  Yes  Small  Shallow  Solitary  Euphausiidae, Calanidae  11.3  15.2  12.3  16.5  Yes  Small  Shallow  Solitary  Euphausiidae, Calanidae  5.47  6.09  6  6.45  Yes  Small  NA  Solitary  2.09  2.43  2.01  2.38  No  Medium  Shallow  Medium  Sperm Whale (Physeter macrocephalus)  15.2  183  10.4  12.5  Yes  Large  Deep  Solitary  Pygmy Sperm Whale (Kogia breviceps)  2.7  3.3  2.66  3.3  No  Medium  Deep  Solitary  Dwarf Sperm Whale (Kogia sima)  2.19  2.34  2.1  2.34  No  Medium  Deep  Solitary  Paracalanidae, Centropagidae, Calanidae, Acartiidae, Clausocalanidae, Onceidae, Oithonidae, Hyperiidae Apogonidae, Leiognathidae, Lethrinidae, 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 Ceratiidae, Gadidae, Macrouridae, Trachipteridae, Icosteidae, Scorpaenidae, Anoplomatidae, Hexagrammida, Nototheniidae, Gnathophausia, Majidae, Cancridae, Architeuthis, Ommastrephidae, Onychoteuthidae, Gonatidae, Pholidoteuthidae, Octopoteuthidae, Histipteuthidae, Cranchiidae, Vampyroteuthidae, Octopodidae Cranchiidae, Enoploteuthidae, Histioteuthidae, Lycoteuthidae, Ommastrephidae, Myctophidae, Gadidae, Gempylidae Ommastrephidae, Cranchiidae, Onychoteuthidae, Lycoteuthidae, Enoploteuthidae, Octopoteuthidae, Chiroteuthidae, Vampyroteuthidae, Gonatidae, Gonostomatidae, Macrouridae, Sternoptychidae, Congridae, Argentinidae, Loligiginidae, Sepiidae, Histioteuthidae, Octopodidae, Moridae, Myctophidae, Penaeidae, Acanthephyridae, Aristeidae, Argentinidae, Microstomatidae  104  Species  Andrew's Beaked Whale (Mesoplodon bowdoini) Arnoux's Beaked Whale (Berardius arnuxii)  Baird's Beaked Whale (Berardius bairdii)  Blainville's Beaked Whale (Mesoplodon densirostris) Cuvier's Beaked Whale (Ziphius cavirostris)  Gervais' Beaked Whale (Mesoplodon europaeus) Ginkgo-Toothed Beaked Whale (Mesoplodon ginkgodens) Gray's Beaked Whale (Mesoplodon grayi) Hector's Beaked Whale (Mesoplodon hectori) Hubb's Beaked Whale (Mesoplodon carlhubbsi) Pygmy Beaked whale (Mesoplodon peruvianus) Longman's Beaked Whale (Mesoplodon pacificus)  Northern Bottlenose Whale (Hyperoodon ampullatus)  Predators  Max Temp (°C) 19  Vocalizatio n Frequency  Dalatiidae  Min Temp (°C) 13.0  Parasites  Species Known to Associate with  Medium  NA  NA  Dalatiidae  NA  NA  Medium  NA  Dalatiidae, Delphinidae  15.0  29  Medium  Dalatiidae  25.0  29  Medium  Cyamidae, Lepadidae, Anisakidae, Tetrameridae, Monocotylidae, Phyllobothriidae, Diphyllobothriidae, Brachycladiidae, Cestodes, Trematodes, Nematodes Anisakidae, Tetrabothriidae, Lepadidae  Killer Whale (Orcinus orca), Hourglass Dolphin (Lagenorhynchus cruciger), Peale's Dolphin (Lagenorhynchus australis) Northern Right Whale Dolphin (Lissodelphis borealis)  Dalatiidae, Lamnidae, Delphinidae  25.0  29  High  Dalatiidae  NA  NA  Medium  Dalatiidae  NA  NA  Dalatiidae  NA  Dalatiidae  NA  Phyllobothriidae, Tetrameridae  NA  NA  Medium  Lepadidae, Phyllobothriidae, Cyamidae NA  NA  Medium  NA  NA  NA  NA  Medium  Lepadidae, Phyllobothriidae  NA  Dalatiidae  NA  NA  Medium  Pennellidae  NA  NA  18.2  19.3  Medium  Anisakidae, Campulidae  NA  Dalatiidae  27.0  30  Medium  NA  Delphinidae  -1.3  -0.9  Medium  Tetrabothriidae, Anisakidae, Polymorphidae  Short-Finned Pilot Whale (Globicephala macrorhynchus), Bottlenose Dolphin (Tursiops truncatus), Spinner Dolphin (Stenella longirostris) Killer Whale (Orcinus orca)  NA  References (and sources cited within) 1, 3, 12, 202, 339 2, 9, 12, 46, 101, 200, 204, 258, 340 7, 8, 9, 12, 101, 200, 340  4, 9, 11, 12, 69, 207, 258, 319, 339 9, 10, 11, 12, 79, 103, 131, 201, 207, 258, 304, 319, 338 9, 12, 69, 71, 258, 314, 339 9, 12, 69, 258, 339 9, 12, 46, 69, 71, 258, 339 6, 9, 12, 46, 69, 258, 339 1, 5, 12, 69, 339 12, 68, 69, 71, 312, 313, 339 12, 34, 69, 80, 309, 339  9, 12, 46, 49, 103, 126, 300, 304, 336, 339  105  Species  Southern Bottlenose Whale (Hyperoodon planifrons)  Dalatiidae  Min Temp (°C) NA  Sowerby's Beaked Whale (Mesoplodon bidens)  Dalatiidae  NA  NA  Medium  Straptoothed Whale (Mesoplodon layardii)  Dalatiidae  10.0  16  Medium  Phyllobothriidae, Tetrabothriidae, Lepadidae, Tetrameridae, Anisakidae, Polymorphidae Lepadidae  Dalatiidae, Lamnidae  NA  NA  Medium  Tetrameridae, Tetrabothriidae  NA  NA  13.0  19  NA  NA  NA  Dalatiidae  13.0  19  Medium  Coronulidae, Pennellidae, Anisakidae  NA  Atlantic Humpbacked Dolphin (Sousa teuszii)  NA  18.8  24.5  High  Cyamidae  Bottlenose Dolphin (Tursiops truncatus)  Atlantic Spotted Dolphin (Stenella plagiodon/frontalis)  NA  19.0  27  High  Atlantic White-Sided Dolphin (Lagenorhynchus acutus)  NA  4.0  10  High  Lepadidae, Coronulidae, Cyamidae, Echeneidae, Campulidae, Heterophyidae, Brauninidae, Pseudaliidae, Anisakidae Heterophyidae, Brachycladiidae, Tetrabothriidae, Phyllobothriidae, Tetrabothriidae, Pseudaliidae , Tetrameridae, Pseudaliidae, Polymorphidae, Pseudaliidae, Heterophyidae  Black Dolphin (Cephalorhynchus eutropia)  NA  11.0  14.6  Low  Rough-Toothed Dolphin (Steno bredanensis), Bottlenose Dolphin (Tursiops truncatus), Risso's Dolphin (Grampus griseus), Pan tropical Spotted Dolphin (Stenella attenuata) Killer Whale (Orcinus orca), Bottlenose Dolphin (Tursiops truncatus), Short Beaked Common Dolphin (Delphinus delphis), Long-Finned Pilot Whale (Globicephala melas), White-Beaked Dolphin (Lagenorhynchus albirostris), Fin Whale (Balaenoptera physalus), Humpback Whale (Megeptera novaeangliae) Peale's Dolphin (Lagenorhynchus australis), Commerson's Dolphin (Cephalorhynchus commersonii)  Stejneger's Beaked Whale (Mesoplodon stejnegeri) Tasman Beaked Whale (Tasmacetus shepherdi) True's Beaked Whale (Mesoplodon mirus)  Predators  Max Temp (°C) NA  Vocalizatio n Frequency  Parasites  Medium  Tetrameridae  Anisakidae, Brauninidae, Campulidae, Polymorphidae  Species Known to Associate with  Killer Whale (Orcinus orca), Peale's Dolphin (Lagenorhynchus australis), Hourglass Dolphin (Lagenorhynchus cruciger) NA  NA  References (and sources cited within) 9, 12, 46, 70, 126, 258, 336, 339 9, 12, 46, 69, 71, 103, 173, 258, 339 9, 12, 46, 67, 69, 81, 258, 339 9, 12, 69, 71, 201, 258, 310, 319, 339 9, 12, 102, 123, 202, 308 9, 12, 69, 71, 174, 202, 258, 315, 339 25, 37, 137, 187, 262, 285, 306, 307, 340 13, 16, 38, 46, 94, 212, 217, 340, 341, 345  17, 21, 26, 27, 49, 51, 50, 51, 55, 340, 344, 346  17, 46, 60, 111, 170, 188, 348  106  Species  Bottlenose Dolphin (Tursiops truncatus)  Clymene Dolphin (Stenella clymene)  Predators  Dalatiidae, Carcharhinidae, Lamnidae, Hexanchidae  Dalatiidae  Min Temp (°C) 15.0  Max Temp (°C) 29  Vocalizatio n Frequency  Parasites  High  Brauninidae, Nasitrematidae, Heterophyidae, Anisakidae, Brachycladiidae, Phyllobothriidae, Campulidae, Tetrameridae, Diphyllobothriidae, Pseudaliidae, Polymorphidae  20.2  28.5  High  Coronulidae, Cyamidae, Pseudaliidae, Phyllobothriidae, Nasitrematidae  Species Known to Associate with  Guiana dolphin (Sotalia guianensis), Atlantic Spotted Dolphin (Stenella plagiodon/frontalis), Melon-Headed Whale (Peponocephala electra), ShortFinned Pilot Whale (Globicephala macrorhynchus), Atlantic Humpbacked 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), RoughToothed 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) Pan tropical Spotted Dolphin (Stenella attenuata), Short Beaked Common Dolphin (Delphinus delphis), MelonHeaded Whale (Peponocephala electra)  References (and sources cited within) 13, 15, 17, 154, 158, 159, 160, 186, 187, 207, 246, 265, 266, 319, 322, 326, 340, 341  13, 16, 61, 62, 130, 169, 172, 212, 217, 340, 341, 345  107  Species  Predators  Commerson's Dolphin (Cephalorhynchus commersonii)  NA  Min Temp (°C) 1.0  Max Temp (°C) 16  Vocalizatio n Frequency  Parasites  Low  Brachycladiidae, Anisakidae, Brauninidae, Heterophyidae, Tetrabothriidae  Short Beaked Common Dolphin (Delphinus delphis)  Dalatiidae, Carcharhinidae, Lamnidae, Delphinidae, Sphyrnidae  7.0  23  High  Anisakidae, Pseudaliidae, Tetrameridae, Phyllobothriidae, Campulidae, Heterophyidae  Dusky Dolphin (Lagenorhynchus obscurus)  Hexanchidae, Delphinidae, Lamnidae  10.4  19  High  Tetrabothriidae, Phyllobothriidae, Anisakidae, Trematodes, Pseudaliidae, Brauninidae  False Killer Whale (Pseudorca crassidens)  Dalatiidae, Delphinidae  9.0  31  Medium  Anisakidae, Pseudaliidae, Polymorphidae, Brachycladiidae, Nasitrematidae, Cyamidae, Coronulidae  Species Known to Associate with  Peale's Dolphin (Lagenorhynchus australis), Black Dolphin (Cephalorhynchus eutropia), Burmeister's Porpoise (Phocoena spinipinnis) Striped Dolphin (Stenella coeruleoalba), Risso's Dolphin (Grampus griseus), Clymene Dolphin (Stenella clymene), Atlantic White-Sided Dolphin (Lagenorhynchus acutus), Northern Right Whale Dolphin (Lissodelphis 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), LongFinned Pilot Whale (Globicephala melas), Spinner Dolphin (Stenella longirostris) Southern Right Whale Dolphin (Lissodelphis peronii), Short Beaked Common Dolphin (Delphinus delphis), Long-Finned Pilot Whale (Globicephala melas), Killer Whale (Orcinus orca), Risso's Dolphin (Grampus griseus), Burmeister's Porpoise (Phocoena spinipinnis), Heaviside's Dolphin (Cephalorhynchus heavisidii) Killer Whale (Orcinus orca), Indo-pacific bottlenose dolphin(Tursiops aduncus), Rough-Toothed Dolphin (Steno bredanensis), Fraser's Dolphin (Lagenodelphis hosei), Risso's Dolphin (Grampus griseus), Bottlenose Dolphin (Tursiops truncatus), Melon-Headed Whale (Peponocephala electra), ShortFinned Pilot Whale (Globicephala macrorhynchus)  References (and sources cited within) 17, 46, 108, 109, 110, 185, 200, 348  15, 17, 23, 24, 27, 94, 95, 124, 143, 149, 201, 202, 205, 246, 258, 318, 319, 340, 341  17, 46, 49, 50, 51, 55, 82, 201, 202, 205, 304, 319, 340, 344, 346  16, 41, 42, 84, 207, 225, 258, 319, 340  108  Species  Max Temp (°C) 29  Vocalizatio n Frequency  Dalatiidae  Min Temp (°C) 25.0  High  Tetrabothriidae, Phyllobothriidae, Tetrabothriidae, Brachycladiidae, Anisakidae, Pseudaliidae, Polymorphidae  Indo-Pacific Humpbacked Dolphin (Sousa chinensis)  Lamnidae, Delphinidae  23.9  29.6  High  Anisakidae, Pseudaliidae, Cyamidae  Heaviside's Dolphin (Cephalorhynchus heavisidii) Hector's Dolphin (Cephalorhynchus hectori)  Lamnidae, Hexanchidae  9.0  19  Low  Cyamidae, Coronulidae  Hexanchidae, Carcharhinidae  6.3  22  Low  NA  -0.3  13.4  High  Cyamidae, Strigeidae, Brauninidae, Halocercinae, Pseudaliidae , Anisakidae, Acariidae, Polymorphidae, Campulidae, Phyllobothriidae Anisakidae  Carcharhinidae  20.0  35  Low  Fraser's Dolphin (Lagenodelphis hosei)  Hourglass Dolphin (Lagenorhynchus cruciger)  Irrawaddy Dolphin (Orcaella brevirostris)  Predators  Parasites  Trematode , Schistosomatidae, Nematode  Species Known to Associate with  Sperm Whale (Physeter macrocephalus), False Killer Whale (Pseudorca crassidens), Striped Dolphin (Stenella coeruleoalba), Pan tropical Spotted Dolphin (Stenella attenuata), Spinner Dolphin (Stenella longirostris), ShortFinned Pilot Whale (Globicephala macrorhynchus), Melon-Headed Whale (Peponocephala electra), Risso's Dolphin (Grampus griseus) Bottlenose Dolphin (Tursiops truncatus), Killer Whale (Orcinus orca), Southern Right Whale (Eubalaena australis), Long Beaked common Dolphin (Delphinus capensis) Dusky Dolphin (Lagenorhynchus obscurus) NA  Fin Whale (Balaenoptera physalus), Antarctic Minke Whale (Balaenoptera bonaerensis), Common Minke Whale (Balaenoptera acutorostrata), Sei Whale (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) Finless Porpoise (Neophocaena phocaenoides), Spinner Dolphin (Stenella longirostris)  References (and sources cited within) 16, 17, 96, 122, 132, 133, 134, 207, 340, 344  37, 187, 189, 262, 273, 306, 319, 340  17, 31, 200, 203 17, 97, 112, 129, 179, 200, 201, 319, 203  46, 49, 50, 51, 55, 58, 136, 205, 275, 340, 344, 346  17, 40, 47, 139, 180, 253, 321, 340  109  Species  Predators  Killer Whale (Orcinus orca)  Dalatiidae  Min Temp (°C) 1.7  Long Beaked Common Dolphin (Delphinus capensis)  Dalatiidae  NA  Max Temp (°C) 26  Vocalizatio n Frequency  NA  High  Medium  Parasites  Fasciolidae, Tetrabothriidae, Phyllobothriidae, Anisakidae  Anisakidae  Species Known to Associate with  Southern Right Whale (Eubalaena australis), Sei Whale (Balaenoptera borealis), Fin Whale (Balaenoptera physalus), Bryde's Whale (Balaenoptera edeni), Blue Whale (Balaenoptera musculus), Gray whale (Eschrichtius 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) Pacific White-Sided Dolphin (Lagenorhynchus obliquidens), IndoPacific Humpbacked Dolphin (Sousa chinensis), Bryde's Whale (Balaenoptera edeni)  References (and sources cited within) 16, 17, 103, 149, 204, 205, 207, 225, 258, 292, 293, 294, 303, 304, 305, 306, 340  94, 95, 124, 156, 173, 184, 197, 258, 340, 341  110  Species  Long-Finned Pilot Whale (Globicephala melaena)  Melon-Headed Whale (Peponocephala electra)  Predators  Max Temp (°C) 22  Vocalizatio n Frequency  Delphinidae  Min Temp (°C) 0.6  Parasites  High  Brachycladiidae, Phyllobothriidae, Diphyllobothriidae, Tetrabothriidae, Anisakidae, Pseudaliidae, Polymorphidae  Dalatiidae  25.0  29  High  Nasitrematidae, Phyllobothriidae, Pseudaliidae, Anisakidae  Species Known to Associate with  Southern Right Whale Dolphin (Lissodelphis peronii), Humpback Whale (Megeptera novaeangliae), Killer Whale (Orcinus orca), Atlantic WhiteSided 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) Bottlenose Dolphin (Tursiops truncatus), Spinner Dolphin (Stenella longirostris), Pygmy Killer Whale (Feresa attenuata), Fraser's Dolphin (Lagenodelphis hosei), Humpback Whale (Megeptera novaeangliae), Short-Finned Pilot Whale (Globicephala macrorhynchus), RoughToothed Dolphin (Steno bredanensis), Clymene Dolphin (Stenella clymene), Pan tropical Spotted Dolphin (Stenella attenuata), False Killer Whale (Pseudorca crassidens), Sperm Whale (Physeter macrocephalus)  References (and sources cited within) 15, 17, 24, 46, 85, 103, 115, 202, 205, 304, 343  16, 49, 63, 64, 65, 66, 103, 207, 258, 346  111  Species  Predators  Northern Right Whale Dolphin (Lissodelphis borealis)  Pacific White-Sided Dolphin (Lagenorhynchus obliquidens)  Max Temp (°C) 19  Vocalizatio n Frequency  NA  Min Temp (°C) 8.0  Parasites  High  Nasitrematidae, Tetrameridae, Anisakidae, Phyllobothriidae, Coronulidae, Pennellidae  Delphinidae, Lamnidae  12.0  13  High  Nasitrematidae, Brachycladiidae, Phyllobothriidae, Tetrabothriidae, Anisakidae, Tetrameridae  Species Known to Associate with  Pacific White-Sided Dolphin (Lagenorhynchus obliquidens), Short 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) Northern Right Whale Dolphin (Lissodelphis borealis), Killer Whale (Orcinus orca), Risso's Dolphin (Grampus griseus), Striped Dolphin (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)  References (and sources cited within) 36, 46, 178, 311, 347  17, 46, 50, 51, 52, 53, 54, 55, 210, 317, 319, 340, 344, 346  112  Species  Peale's Dolphin (Lagenorhynchus australis)  Pygmy Killer Whale (Feresa attenuata)  Predators  Max Temp (°C) 9  Vocalizatio n Frequency  NA  Min Temp (°C) 6.0  Dalatiidae  25.0  29  NA  High  Parasites  Tetrabothriidae  Lepadidae, Anisakidae, Pseudaliidae, Tetrabothriidae, Nasitrematidae  Species Known to Associate with  Long-Finned Pilot Whale (Globicephala melas), Arnoux's Beaked Whale (Berardius arnuxii), Killer Whale (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) Rough-Toothed Dolphin (Steno bredanensis), Melon-Headed Whale (Peponocephala electra), Risso's Dolphin (Grampus griseus)  References (and sources cited within) 50, 51, 55, 56, 57, 182, 183, 340, 344, 346  16, 48, 207, 210, 258, 316, 319  113  Species  Risso's Dolphin (Grampus griseus)  Rough-Toothed Dolphin (Steno bredanensis)  Predators  Max Temp (°C) 35  Vocalizatio n Frequency  Dalatiidae, Lamnidae  Min Temp (°C) 7.5  Parasites  Medium  Phyllobothriidae, Tetrabothriidae, Tetrameridae, Pseudaliidae, Anisakidae, Nasitrematidae, Cyamidae, Coronulidae  Dalatiidae, Delphinidae  25.0  29  Medium  Tetrabothriidae, Polymorphidae, Anisakidae, Cyamidae  Species Known to Associate with  Bottlenose Dolphin (Tursiops truncatus), Fin Whale (Balaenoptera physalus), Sperm Whale (Physeter macrocephalus), Short-Finned Pilot Whale (Globicephala 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) False Killer Whale (Pseudorca crassidens), Bottlenose Dolphin (Tursiops truncatus), Pygmy Killer 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)  References (and sources cited within) 13, 16, 44, 59, 83, 200, 205, 210, 258, 316, 319  13, 16, 17, 90, 122, 201, 207, 262, 340  114  Species  Short-Finned Pilot Whale (Globicephala macrorhynchus)  Southern Right Whale Dolphin (Lissodelphis peronii)  Spinner Dolphin (Stenella longirostris)  Predators  Min Temp (°C) 25.0  Max Temp (°C) 29  Vocalizatio n Frequency  Parasites  Species Known to Associate with  High  Brachycladiidae, Campulidae, Nasitrematidae, Tetrabothriidae, Phyllobothriidae, Anisakidae, Pseudaliidae  Somniosidae, Nototheniidae  1.0  20  High  Nasitrematidae, Pseudaliidae, Anisakidae, Tetrabothriidae, Opisthorchiidae, Phyllobothriidae  Dalatiidae, Lamnidae, Carcharhinidae, Tetraodontidae, Delphinidae  22.0  27.5  High  Echeneidae, Lepadidae, Anisakidae, Pseudaliidae, Spiruridae, Brachycladiidae, Nasitrematidae, Tetrabothriidae, Phyllobothriidae, Polymorphidae  Bottlenose Dolphin (Tursiops truncatus), Fraser's Dolphin (Lagenodelphis hosei), Melon-Headed Whale (Peponocephala electra), Northern Right Whale Dolphin (Lissodelphis borealis), Pacific WhiteSided 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) Dusky Dolphin (Lagenorhynchus obscurus), Bottlenose Dolphin (Tursiops truncatus), Long-Finned Pilot Whale (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) Bottlenose Dolphin (Tursiops truncatus), Pan tropical Spotted Dolphin (Stenella attenuata), Rough-Toothed Dolphin (Steno bredanensis), Longman's Beaked Whale (Mesoplodon pacificus), Fraser's Dolphin (Lagenodelphis hosei), MelonHeaded 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)  Dalatiidae  References (and sources cited within) 13, 17, 85, 103, 116, 117, 118, 195, 207, 258, 343  35, 36, 46, 181, 201, 202, 205, 319, 347  13, 16, 17, 92, 120, 121, 149, 207, 212, 213, 317, 319, 323, 340, 341, 345  115  Species  Pan tropical Spotted Dolphin (Stenella attenuata)  Striped Dolphin (Stenella coeruleoalba)  Guiana dolphin (Sotalia guianensis) White-Beaked Dolphin (Lagenorhynchus albirostris)  Beluga (Delphinapterus leucas)  Predators  Min Temp (°C) 25.3  Max Temp (°C) 28  Vocalizatio n Frequency High  Phyllobothriidae, Tetrabothriidae, Brauninidae, Nasitrematidae, Anisakidae, Tetrameridae  Dalatiidae  20.0  30  High  Dalatiidae, Delphinidae, Carcharhinidae  15.0  31  High  Cyamidae , Pennellidae, Lepadidae, Coronulidae, Tetrabothriidae, Phyllobothriidae, Nasitrematidae, Brachycladiidae, Heterophyidae, Campulidae, Brachycladiidae, Anisakidae, Anguillicolidae, Tetrameridae, Pseudaliidae, Polymorphidae Nasitrematidae, Pseudaliidae, Anisakidae  Delphinidae  8.1  17.2  High  Delphinidae, Ursidae, Somniosidae  0.0  16  Medium  Dalatiidae, Delphinidae, Caleocerdo cuvier  Parasites  Anisakidae, Pseudaliidae, Cyamidae  Brachycladiidae, Campulidae, Diphyllobothriidae, Anisakidae, Tetrameridae, Pseudaliidae, Ascarididae, Polymorphidae  Species Known to Associate with  Atlantic Spotted Dolphin (Stenella plagiodon/frontalis), Spinner Dolphin (Stenella longirostris), Clymene Dolphin (Stenella clymene), Bottlenose Dolphin (Tursiops truncatus), Fraser's Dolphin (Lagenodelphis hosei), Risso's Dolphin (Grampus griseus), Rough-Toothed Dolphin (Steno bredanensis), MelonHeaded Whale (Peponocephala electra), Short-Finned Pilot Whale (Globicephala macrorhynchus), Bryde's Whale (Balaenoptera edeni) Short Beaked Common Dolphin (Delphinus delphis), Fraser's Dolphin (Lagenodelphis hosei), Pacific WhiteSided Dolphin (Lagenorhynchus obliquidens), Risso's Dolphin (Grampus griseus), Gray whale (Eschrichtius robustus), Fin Whale (Balaenoptera physalus)  Bottlenose Dolphin (Tursiops truncatus)  Harbour Porpoise (Phocoena phocoena), Killer Whale (Orcinus orca), LongFinned Pilot Whale (Globicephala melas), Atlantic White-Sided Dolphin (Lagenorhynchus acutus), Fin Whale (Balaenoptera physalus), Humpback Whale (Megeptera novaeangliae), Sei Whale (Balaenoptera borealis), Short Beaked Common Dolphin (Delphinus delphis), Bottlenose Dolphin (Tursiops truncatus) Narwhal (Monodon monoceros), Killer Whale (Orcinus orca)  References (and sources cited within) 13, 16, 17, 38, 39, 46, 103, 207, 209, 212, 217, 258, 319, 323, 340, 341, 345  13, 16, 17, 89, 93, 150, 151, 205, 217, 258, 340, 341, 345  17, 22, 46, 91, 201, 214, 252, 258, 340, 341 21, 23, 46, 50, 51, 55, 152, 194, 340, 344, 346  14, 17, 49, 99, 103, 193, 201, 218, 219, 221, 222, 319, 325, 340  116  Species  Narwhal (Monodon monoceros)  Franciscana (Pontoporia blainvillei) Burmeister's Porpoise (Phocoena spinipinnis)  Cochito [Vaquita] (Phocoena sinus) Dall's Porpoise (Phocoenoides dalli)  Finless Porpoise (Neophocaena phocaenoides)  Harbour Porpoise (Phocoena phocoena)  Predators  Min Temp (°C) -2.0  Max Temp (°C) 5  Vocalizatio n Frequency  Delphinidae, Carcharhinidae, Hexanchidae NA  16.0  27  NA  4.0  19.5  Low  Lamnidae, Carcharhinidae, Alopiidae, Hexanchidae, Delphinidae, Lamnidae  17.0  32  Low  12.4  24  Dalatiidae, Lamnidae  5.7  Dalatiidae, Delphinidae, Lamnidae, Somniosidae,  6.0  Odobenidae, Delphinidae, Ursidae, Somniosidae,  High  Parasites  Species Known to Associate with  Cyamidae, Anisakidae, Ascarididae, Pseudaliidae  Beluga (Delphinapterus leucas), Killer Whale (Orcinus orca)  Coronulidae, Cirolanidae, Cymothoidae, Cocconeidaceae, Anisakidae, Polymorphidae Campulidae, Heterophyidae, Nasitrematidae, Brauninidae, Anisakidae, Pseudaliidae, Polymorphidae, Cyamidae, Coronulidae Tetrameridae, Campulidae, Coronulidae  NA  NA  Pseudaliidae, Nasitrematidae, Brachycladiidae, Anisakidae, Tetrameridae, Phyllobothriidae, Polymorphidae, Cyamidae  25.6  NA  17  Low  Brachycladiidae, Nasitrematidae, Tetrameridae, Pseudaliidae, Diphyllobothriidae Brachycladiidae, Campulidae, Opisthorchiidae, Heterophyidae, Diphyllobothriidae, Anisakidae, Pseudaliidae, Ascarididae, Polymorphidae  Harbour Porpoise (Phocoena phocoena), Killer Whale (Orcinus orca), Fin Whale (Balaenoptera physalus), Blue Whale (Balaenoptera musculus), Gray whale (Eschrichtius robustus), Humpback Whale (Megeptera novaeangliae), Northern Right Whale Dolphin (Lissodelphis borealis), Pacific WhiteSided Dolphin (Lagenorhynchus obliquidens), Risso's Dolphin (Grampus griseus), Sei Whale (Balaenoptera borealis) Irrawaddy Dolphin (Orcaella brevirostris)  Bottlenose Dolphin (Tursiops truncatus), Dusky Dolphin (Lagenorhynchus obscurus), Commerson's Dolphin (Cephalorhynchus commersonii) NA  Dall's Porpoise (Phocoenoides dalli), Bottlenose Dolphin (Tursiops truncatus), Killer Whale (Orcinus orca), Pacific White-Sided Dolphin (Lagenorhynchus obliquidens), White-Beaked Dolphin (Lagenorhynchus albirostris), Fin Whale (Balaenoptera physalus), Common Minke Whale (Balaenoptera acutorostrata), Humpback Whale (Megeptera novaeangliae)  References (and sources cited within) 17, 46, 100, 145, 201, 221, 254, 253, 319, 320, 323, 340 17, 33, 98, 248, 249, 250, 251, 319, 323 17, 46, 86, 114, 284, 289  17, 46, 113, 175, 244, 245, 246, 289 17, 46, 87, 88, 210, 232, 235, 236, 319  17, 28, 29, 30, 87, 240, 241, 242, 258 17, 21, 75, 76, 77, 103, 176, 177, 224, 227, 232, 233, 234, 258, 289, 319  117  Species  Max Temp (°C) 9.5  Vocalizatio n Frequency  NA  Min Temp (°C) 5.5  Low  NA  NA  Delphinidae  0.0  25  Low  Notocotylidae, Tetrabothriidae, Pseudophyllidae, Polymorphidae  Blue Whale (Balaenoptera musculus)  Dalatiidae, Delphinidae  14.1  21.6  Low  Notocotylidae, Tetrabothriidae, Anisakidae, Tetrameridae, Ascarididae, Polymorphidae  Bryde's Whale (Balaenoptera edeni)  Dalatiidae, Delphinidae  18.5  19  Low  Polymorphidae  Killer Whale (Orcinus orca), Bottlenose Dolphin (Tursiops truncatus), Striped Dolphin (Stenella coeruleoalba), Short Beaked Common Dolphin (Delphinus delphis), Dall's Porpoise (Phocoenoides dalli), Short-Finned Pilot Whale (Globicephala macrorhynchus), Northern Right Whale Dolphin (Lissodelphis borealis), Pacific WhiteSided Dolphin (Lagenorhynchus obliquidens), Risso's Dolphin (Grampus griseus) Killer Whale (Orcinus orca), Fin Whale (Balaenoptera physalus), Pacific WhiteSided Dolphin (Lagenorhynchus obliquidens), Dall's Porpoise (Phocoenoides dalli), Bottlenose Dolphin (Tursiops truncatus), Rough-Toothed Dolphin (Steno bredanensis) Killer Whale (Orcinus orca), Long Beaked common Dolphin (Delphinus capensis), Rough-Toothed Dolphin (Steno bredanensis), Indo-pacific bottlenose dolphin (Tursiops aduncus), Spinner Dolphin (Stenella longirostris), Pan tropical Spotted Dolphin (Stenella attenuata)  Spectacled Porpoise (Phocoena dioptrica) Gray whale (Eschrichtius robustus)  Predators  Parasites  Species Known to Associate with  References (and sources cited within) 17, 32, 238, 289 14, 19, 49, 103, 135, 222, 253, 340  16, 18, 49, 125, 141, 161, 164, 166, 258, 300, 304, 340  16, 18, 49, 128, 141, 164, 166, 174, 196, 246, 247, 258, 291, 292, 302, 340  118  Species  Predators  Fin Whale (Balaenoptera physalus)  Dalatiidae, Delphinidae  Min Temp (°C) 16.0  Max Temp (°C) 23  Vocalizatio n Frequency  Parasites  Species Known to Associate with  Low  Brachycladiidae, Notocotylidae, Diphyllobothriidae, Phyllobothriidae, Tetrabothriidae, Anisakidae, Tetrameridae, Polymorphidae  Killer Whale (Orcinus orca), Blue Whale (Balaenoptera musculus), Atlantic White-Sided Dolphin (Lagenorhynchus acutus), Hourglass Dolphin (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) Hourglass Dolphin (Lagenorhynchus cruciger), Peale's Dolphin (Lagenorhynchus australis), Pygmy Right Whale (Caperea marginata)  Antarctic Minke Whale (Balaenoptera bonaerensis)  Dalatiidae, Delphinidae  -1.9  21.8  Low  Pennellidae, Cyamidae, Anisakidae  Common Minke Whale (Balaenoptera acutorostrata)  Dalatiidae, Delphinidae  6.1  20.1  Low  Pennellidae, Brachycladiidae, Diphyllobothriidae, Tetrabothriidae, Anisakidae, Cyamidae  Sei Whale (Balaenoptera borealis)  Dalatiidae, Delphinidae  5.0  18.8  Low  Notocotylidae, Diphyllobothriidae, Tetrabothriidae, Polymorphidae, Tetrameridae, Anisakidae  Humpback Whale (Megeptera novaeangliae), Rough-Toothed Dolphin (Steno bredanensis), Bottlenose Dolphin (Tursiops truncatus), Hourglass Dolphin (Lagenorhynchus cruciger), Harbour Porpoise (Phocoena phocoena) Killer Whale (Orcinus orca), Hourglass Dolphin (Lagenorhynchus cruciger), Northern Right Whale Dolphin (Lissodelphis borealis), Pacific WhiteSided Dolphin (Lagenorhynchus obliquidens), Peale's Dolphin (Lagenorhynchus australis), WhiteBeaked Dolphin (Lagenorhynchus albirostris), Pygmy Right Whale (Caperea marginata), North Atlantic Right whale (Eubalaena glacialis), Humpback Whale (Megeptera novaeangliae), Dall's Porpoise (Phocoenoides dalli)  References (and sources cited within) 18, 21, 49, 103, 141, 164, 166, 167, 246, 258, 300, 304, 340  46, 78, 140, 141, 142, 164, 166, 189, 204, 205, 206, 258, 282, 301, 340 21, 46, 140, 141, 144, 164, 166, 258, 300, 340  18, 49, 103, 140, 141, 164, 166, 206, 258, 282, 283, 301, 304, 340  119  Species  Max Temp (°C) 19  Vocalizatio n Frequency  Parasites  Species Known to Associate with  Dalatiidae, Delphinidae  Min Temp (°C) 13.0  Low  Diphyllobothriidae, Anisakidae, Brachycladiidae, Tetrameridae, Polymorphidae  Delphinidae  -1.6  2  Low  North Pacific Right Whale (Eubalaena japonica)  NA  3.0  17  Medium  Brachycladiidae, Notocotylidae, Phyllobothriidae, Polymorphidae, Tetrameridae Cyamidae, Tetrabothriidae, Polymorphidae  Bottlenose Dolphin (Tursiops truncatus), Killer Whale (Orcinus orca), Common Minke Whale (Balaenoptera acutorostrata), Rough-Toothed Dolphin (Steno bredanensis), Dall's Porpoise (Phocoenoides dalli), Atlantic WhiteSided Dolphin (Lagenorhynchus acutus), Long-Finned Pilot Whale (Globicephala melas), Melon-Headed Whale (Peponocephala electra), Northern Right Whale Dolphin (Lissodelphis borealis), Pacific WhiteSided 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) NA  North Atlantic Right Whale (Eubalaena glacialis)  NA  2.2  21.8  Medium  Cyamidae  Delphinidae  13.0  19  Medium  Cyamidae, Hydrophilidae, Tetrabothriidae, Polymorphidae  Pygmy Right Whale (Caperea marginata)  Dalatiidae  5.0  20  Low  NA  Indo-pacific bottlenose dolphin (Tursiops aduncus)  Dalatiidae  13.4  24.1  High  Anisakidae, Cyamidae, Pseudaliidae, Tetrameridae, Phyllobothriidae, Coronulidae  Humpback Whale (Megeptera novaeangliae)  Bowhead Whale (Balaena mysticetus)  Southern Right Whale (Eubalaena australis)  Predators  Humpback Whale (Megeptera novaeangliae) Humpback Whale (Megeptera novaeangliae), Sei Whale (Balaenoptera borealis) Killer Whale (Orcinus orca), Peale's Dolphin (Lagenorhynchus australis), Indo-Pacific Humpbacked Dolphin (Sousa chinensis) Long-Finned Pilot Whale (Globicephala melas), Sei Whale (Balaenoptera borealis), Antarctic Minke Whale (Balaenoptera bonaerensis) False Killer Whale (Pseudorca crassidens), Bryde's Whale (Balaenoptera edeni), Spinner Dolphin (Stenella longirostris)  References (and sources cited within) 17, 18, 21, 49, 103, 138, 162, 202, 204, 205, 258, 261, 300, 340  14, 18, 46, 49, 103, 163, 222, 280, 281, 340 18, 46, 49, 104, 146, 147, 278, 279, 340 18, 46, 49, 105, 146, 192, 198, 340 46, 49, 104, 106, 107, 146, 147, 190, 191, 202, 304, 340 46, 72, 73, 258, 340  17, 74, 153, 155, 180, 190, 258, 276, 277, 287, 340, 341  120  Species  Predators  Min Temp (°C) 12.0  Max Temp (°C) 30.3  Vocalizatio n Frequency  Parasites  Species Known to Associate with  Medium  Brachycladiidae, Tentaculariidae, Diphyllobothriidae, Tetrabothriidae, Anisakiidae, Phyllobothriidae, Tetrameridae, Polymorphidae, Pennellidae, Coronulidae, Lepadidae, Cyamidae, Echeneidae  Pacific White-Sided Dolphin (Lagenorhynchus obliquidens), Northern Right Whale Dolphin (Lissodelphis borealis), Risso's Dolphin (Grampus griseus), Melon-Headed Whale (Peponocephala electra), Fraser's Dolphin (Lagenodelphis hosei), ShortFinned Pilot Whale (Globicephala macrorhynchus), Southern Right Whale Dolphin (Lissodelphis peronii) NA  Sperm Whale (Physeter macrocephalus)  Delphinidae, Dalatiidae, Somniosidae  Pygmy Sperm Whale (Kogia breviceps) Dwarf Sperm Whale (Kogia sima)  Delphinidae, Lamnidae  26.9  30.9  Medium  Delphinidae, Lamnidae  26.0  26.4  Medium  References for Species Associations (and sources cited within)  Phyllobothriidae, Tetraphyllidea incertae sedis, Anisakidae, Tetrameridae, Pseudaliidae, Polymorphidae, Pennellidae  NA  References (and sources cited within) 16, 81, 327, 332, 333, 334, 339  289, 304, 319, 328, 330, 331 16, 120, 289, 328, 329, 330, 331, 332, 335  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, 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, 304, 312, 327, 330  121  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. 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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  143  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  144  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) Male body length Female body length Sexual Dimorphism Range Size Water Depth Water Temperature Prey Species Predator Species Parasite Species Average Group Size Known Associate Species Natural Range Overlap Vocalization Frequency  PC1 0.0160 0.0179 -0.0414 -0.0289 -0.0486 -0.0151 -0.0019 -0.0093 -0.0044 -0.0057 -0.0028 -0.0151 0.0025  PC2 PC3 -0.0025 -0.0032 -0.0019 -0.0019 0.0488 0.0297 0.0306 0.0308 0.0134 0.0019 -0.0116 -0.0078 -0.0004 0.0008 -0.0004 0.0049 0.0006 0.0031 -0.0058 -0.0008 -0.0007 0.00003 -0.0089 -0.0026 -0.0074 -0.0055  PC4 -0.0020 -0.0044 -0.0045 -0.0021 0.0070 0.0010 0.0034 0.0020 -0.0002 0.0694 0.0022 0.0032 0.0112  145  

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