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Bangarang February 2014 Backgrounder1 Marine Tetrapods (of the Kitimat Fjord System) Eric Keen Abstract Marine tetrapods are vertebrates secondarily adapted for marine environment who obtain most or all of their nourishment from the sea. This includes marine reptiles, marine mammals (cetaceans, pinnipeds, sirenians, sea otters, sea bats and polar bears) and seabirds. This Backgrounder reviews their general natural history and compiles information relevant to the status, ecology and distribution of those marine tetrapods expected in the Kitimat Fjord System. Of marine mammals, the Kitimat Fjord System is commonly host to two mysticetes, four odontocetes, two phocids, one otariid, and one mustelid. Depending on how one deals with the seasonal use of marine habitats, 35-55 seabirds are expected in the area (excluding shorebirds). Contents Natural History Taxonomy Marine tetrapods Marine mammals Seabirds Evolution Water: The subtle difference Marine mammals Seabirds Biology Anatomy, Morphology Energetics Diving Life History – Marine Mammals Life History – Seabirds Foraging Marine Mammals of the Kitimat Fjord System Toothed cetaceans Mustached cetaceans Pinnipeds Mustelids Seabirds of the Kitimat Fjord System Taxon by Taxon Important Bird Areas (IBAs) 1 Bangarang Backgrounders are imperfect but rigorouss reviews – written in haste, not peer-reviewed – in an effort to organize and memorize the key information for every aspect of the project. They will be updated regularly as new learnin’ is incorporated. 1 Natural History Taxonomy For our purposes, tetrapods (amphibians, reptiles, birds and mammals) are considered marine if they obtain most or all of their sea from the marine environment. Marine Mammals The term “marine mammal” is not a natural biological grouping; it encompasses 130 species of cetaceans, pinnipeds (these are the two most common and well known marine mammal groups), sirenians, and fissipeds (Carnivora members with separate digits, including the otters and polar bears), all of whom retrieve most of their food from the sea. As mammals, all of these groups are endothermic, nurse live young, and have diagnostically mammalian skulls. As marine mammals, they obtain all their food from the sea2. These disparate groups represent 5 or 6 recolonizations of aquatic habitat. Marine mammal taxonomy and systematics is highly controversial and currently in a state of flux. These marine mammal groups are united not by close relation, but by a similar story that has transformed their life histories and ecologies in shared respects3. Marine mammals are extraordinarily derived, rivaling the adaptive ingenuity of bats. To varying degrees and with exceptions, all are streamlined and have reduced appendages, modified physiology and anatomy for thermo- and osmo-regulation, unique strategies and “equipment” for foraging, enhancement or loss of certain senses, and myriad internal adaptations to marine living. Their secondarily marine life has also fundamentally restructured their life history, reproductive strategies, intelligence, and sociality. All marine mammals are directly or indirectly impacted by human activities, some to the point of extinction4. The zoogeography, distribution, and migratory behaviors of marine mammal species are highly variable and prohibit summary. One available generalization is that little is known of the particular factors that limit the occurrence of species. The physical and bathymetric features that drive megascale patterns in ocean circulation and productivity are ultimately responsible for many marine mammal distributions5. Cetaceans are a monophyletic group that diverged from other artiodactyls (even-toed ungulates) >50mya and underwent punctuated periods of radiation, survived by ~86 extant species. ). As the most derived secondarily marine mammal, they are completely independent of land, practically hairless, well-blubbered, and extremely hydrodynamic (fusiform bodies, no hind appendages, telescoped skull, etc.) 6. The Cetacea can be split into two monophyletic groups: Suborder Mysteceti (baleen whales): 4 families, 14 species. Mysticetes have no functional teeth beyond the fetus stage, and both nostrils are present as blowholes; their skulls are symmetrical, and they are universally large. The most speciose family is the Balaenopteridae, the lunge-feeding, dorsal-finned rorquals, which includes the largest animals to have ever lived (i.e. the blue whales). Other families are the Balaenidae (right and bowhead whales), Neobalaenidae (pygmy right whales, resurrected cetotheres), and Eschrichtiidae (gray whales) 7. Suborder Odontoceti (toothed whales): 10 families, 65 species. Odontocetes bear homodont teeth, have asymmetrical skulls, and all are known or assumed to echolocate. They are generally smaller than mysticetes, but male sperm whales (the lone species of the family Physteridae) can be over 60ft long. Many odontocetes are specialists at foraging at great depths, and include some of the deepest divers on earth. In addition to sperm whales, families include the Kogiidae (2 species; pygmy and dward sperm whales), Ziphiidae (21+ species; beaked whales), four families of river dolphins, all of whom returned independently of each other to freshwater 2 Ballance, L. 2013. Marine mammal taxonomy. Marine Tetrapods. Scripps Institution of Oceanography. Lecture 2. 3 Jefferson, T.A., M.A. Webber, and R.L. Pitman. 2008. Introduction. Marine Mammals of the World. Academic Press. 4 Jefferson, T.A., M.A. Webber, and R.L. Pitman. 2008. Introduction. Marine Mammals of the World. Academic Press. 5 Jefferson, T.A., M.A. Webber, and R.L. Pitman. 2008. Introduction. Marine Mammals of the World. Academic Press. 6 Ballance, L. 2013. Marine mammal taxonomy. Marine Tetrapods. Scripps Institution of Oceanography. Lecture 2. 7 Ballance, L. 2013. Marine mammal taxonomy. Marine Tetrapods. Scripps Institution of Oceanography. Lecture 2. 2 habitats (4 families, 4 genera, 4 species), the Monodontidae (beluga and narwhal), the Phocoenidae (porpoises; 3 genera, 7 species), and finally, the Delphinidae (17 genera, 36 species, the largest being the killer whale) 8. The Sirenia, the only marine mammal herbivores, boast a fossil record that extends back >50mya; they evolved from proboscideans (shared ancestors of elephants and hyraxes). There are two families (Trichechidae, 3 manatee species; Dugongidae, 1 dugong species)9. Marine mammals in the order Carnivora include all members of the Suborder Pinnipedia (fins modified as non- digited flippers; 34 species). Pinnipeds are a monophyletic group of amphibious carnivores that diverged from a (likely) ursid ancestor 30-35mya. There are five major lineages of pinnipeds, three of which are extant (tusked odobenid walruses, 1 species; eared otariid seals, 16 species; and earless phoecid “true” seals, 19 species)10. All species are tied in some way to land, all have skin with hair underlain with a blubber layer, some exhibit extensive sexual dimorphism, and all have small litters (~1 pup). The marine fissipeds (Carnivora), otters and bears, are relative newcomers. Otters (Mustelidae: Lutrinae, six of 13 species are marine-living, 1 obligately marine) are only a few million years old. Polar bears, the only marine ursid, diverged from brown bears ~1.3 mya. It is the least aquatic and least derived of the marine mammals11. Another marine representatives of the Carnivora one species of the Ursidae (polar bear, require sea ice and land; 1 genus, 1 species) 12. Within the Order Chiroptera, there is one bat species in the Noctilionidae that preys upon aquatic and marine fishes13. Identifying the representatives of these taxa can be difficult, but good identification guides are essential for wildlife watching, research, and education programs. Virtually every marine mammal species exhibits variability among its geographic populations. Significant dimorphism among sexes, life stages, seasonal appearances (though less important in marine mammals), uncommon color morphs, individuals (perhaps due to scarring or injuries), and sighting conditions – as well as the possibility of hybrids and intergrades between species -- may also confound efforts to identify and describe a species14. Seabirds The term “seabird” is a loosely taxonomic grouping that encompasses ~350 species in many evolutionarily disparate clades, including penguins, “tubenose” seabirds, tropicbirds, pelicans, frigatebirds, boobies and gannets, cormorants, and gulls and terns and their close relatives. As birds, all of these groups possess feathers, forelimbs modified into wings, no teeth, highly modified skeletons for flight, and an extensive airsac system throughout their body. Birds are also homeothermic and oviparous. As seabirds, they obtain all or most of their food from the sea. Notably, this group does not include sea ducks, osprey or sea eagles, and it does include species like gulls and terns who often obtain their food from land15. The penguins (O. Sphenisciformes, f. Spheniscidae, 6 genera and 18 species) are strictly marine, currently restricted to the southern hemisphere, and are primarily temperate and subpolar16. The “tubenoses” (O. 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