Bangarang January 2014 Backgrounder1

Fin Whales! (especially in British Columbia)

Eric Keen

Abstract This is a review of the natural history of the fin whale, with special emphasis on those occurring in British Columbian waters, with even more emphasis on those occurring within the Kitimat Fjord System (Gitga’at First Nation marine territory and the study area of CetaceaLab and the Bangarang Project). Contents

Etymology Taxonomy Measurements Description Phylogeny Range and Habitat General Proximity to Shore British Columbia Seasonal Movements Reproduction Life History Mating Calving Behavior Acoustics Food and Foraging Diet Timing of Feeding Foraging Competition Predation

Parasites Whaling Current Status Threats Protection Ongoing BC Research

                                                                                                               1 Bangarang Backgrounders are imperfect but rigorous reviews – written in haste, not peer-reviewed – in an effort to organize and memorize the key information for every aspect of the project.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.

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Fin Whales Balaenoptera physalus (Linnaeus, 1758)

Etymology Balaeno- Baleen (whale) ptera - wing or fin (referring to dorsal fin) physalus “bellows” (ref. either to its ventral pleats or its blow)2

“rorqual whale”3 “a kind of toad that puffs itself up” – inflation of ventral pouch during feeding4, 5,6 “a wind instrument”7 (Grk. physalis, perhaps for the flute-like inhalation sound.)

Other English common names:

Finback8 Common rorqual9 Razorback (after the sharp dorsal ridge on her caudal peduncle)10 Herring whale11 True fin whale12 Gibbar13 Finner14

Notable other languages:15 Norwegian: finefisk, finvhal,sildror, Japanese: Nagasu-kujira Russian: Finval, seldianoi polostaik, kiit German: Finnfisk, sillval, rorval Icelandic: Sildrek, seldreki, hunfubaks French: Baleine americaine, vraie baleine Dutch: Vinvis, gewone vinvis Eskimo: Tykyshkok, keporkarnak Spanish: Ballena de Aleta Italian: Capidolio (not Capidoglio, or sperm whale)

Taxonomy This species was originally described by Frederik Martens in 167516, and again by Paul Dudley in 1725.17 Linneaus assigned this species the name Balaena physalus in 1758.

                                                                                                               2 Reeves et al. 2002. 3 Reeves et al. 2002. 4 Reeves et al. 2002. 5 Gambell 1985. 6 Leatherwood and Reeves 1983. 7 Gambell 1985. 8 Reeves et al. 2002. 9 Reeves et al. 2002. 10 Reeves et al. 2002. 11 Gambell 1985. 12 Gambell 1985. 13 Gambell 1985. 14 Gambell 1985. 15 Gambell 1985. 16 Aguilar 2008 17 Aguilar 2008

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Subsequently, Lacepede moved the species into the newly defined Balaenoptera genus in 1804, assigning it the name Balaenoptera rorqual18. In 1903, Bacovitza synonymized B. rorqual with two other redundant descriptions – (1) B. australis from Gray 1846, described from voyage of HMS Erebus and Terror, and (2) B. patachonicus from Burmeister 186519 -- all under the single name Balaenoptera physalus20. As early as 1829, the two hemispheric populations were known to be distinct. The southern population was recognized as subspecies B.p. quoyi (Fischer 1829)21. The northern subspecies is B. physalus physalus. The fin whale’s subspecies taxonomy is an area of active research, and revisions are expected (see “Phylogeny”).

Measurements Most sources disagree. All metrics found in the literature are shown. At Birth:

Length: 19’2-21’ (6-6.5m)22 19’6-23’ (6-7m)23

Weight: 4,000 to 6,000 lbs (1,800 – 2,700 kg)24 2200 – 3300 lbs (1,000 – 1,500 kg) 25

Adult Sexually mature fin whales are dimorphic; the females are 5-10% longer26,27. Maximum Length:

Northern hemi.: 79’ (24m)28 Southern Ocean: 89’ (27.1m)29

Average Length: Northern hemi.: 70’ (21.5m)30

72’ – 78’ (22-24m)31 59’-75’ (17 – 23m) 32 74’ (22.5m) for females, 68’ (21m) avg. for males33

Southern Ocean: 82 – 89’ (25-27m)34 85’ (26m) average female, 82’ (25m) average males35

Note that body weight can vary substantially throughout the year due to winter fasting. The result is that the relative mass of body tissue varies seasonally according to nutritive condition36. Maximum Weight:

260,000 lbs (120,000 kg, 117 metric tons)37 198,000 lbs (90,000 kg, 90 metric tons)38

                                                                                                               18 Gambell 1985. 19 Gambell 1985. 20 Gambell 1985. 21 Gambell 1985. 22 Reeves et al. 2002. 23 Aguilar 2008 24 Reeves et al. 2002. 25 Aguilar 2008 26 Aguilar 2008 27 Reeves et al. 2002. 28 Reeves et al. 2002. 29 Reeves et al. 2002. 30 Eder 2001. 31 Gambell 1985. 32 Creswell et al. 2007. 33 Aguilar 2008 34 Gambell 1985. 35 Aguilar 2008 36 Lockyer and Waters, 1986 37 Reeves et al. 2002. 38 Lockyer 1976.

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Average Weight: Northern hemi: 88,184 – 110,230 lbs (40,000 – 50,000 kg, 40-50 metric tons)39

92,000 lbs (42,150 kg, 42.14 metric tons)40 Southern Ocean: 132,000 – 176,370 lbs (60,000 – 80,000 kg, 60-80 metric tons)41

176,370 lbs (80,000 kg, 80 metric tons.)42 Organ Weights The largest captured Antarctic female fin whale (total weight: 57 tons) that was weighed organ by organ yielded the following results.43

Ovaries: 2.7 ton ovaries Blubber: 13.78 tons of blubber Meat: 25.22 tons of meat Heart: 0.13 ton heart Bone: 11.42 tons of bone

Skull: 2.62 ton skull Vertebrae 4.76 tons of vertebrae Mandible: 1.25 ton jaw

Of interest: the fin whale’s right lung is known to be 10% heavier than the left.44

Description The fin whale is the second largest animal on earth45, but of the great whales she is the fastest in the ocean. BODY SHAPE The fin whale’s slender, sleek fusiform body is greatly compressed towards the caudal region46. Her maximum girth is 40-50% of her total length.47 PIGMENTATION - BODY Dark gray above and white- to cream-colored below.48 Flukes are bordered with gray underneath.49 Individuals can be identified by means of scarring, pigmentation patterns, dorsal fin shapes, and/or dorsal nicks50. PIGMENTATION - JAW The right mandible is white (mneumonic: “White on Right”), a trait diagnostic for fin whales51. This pale coloration is also found on the right front baleen plates52. PIGMENTATION – HEAD Fin whales have strikingly asymmetrical head pigmentation53.

                                                                                                               39 Aguilar 2008 40 Barlow et al. 2008. 41 Aguilar 2008 42 Eder 2001. 43 Nishiwaki 1950. 44 Aguilar 2008 45 Reeves et al. 2002. 46 Gambell 1985. 47 Aguilar 2008 48 Reeves et al. 2002. 49 Reeves et al. 2002. 50 COSEWIC 2005, Agler et al. 1990, Falcone et al. 2011. 51 Reeves et al. 2002. 52 Reeves et al. 2002. 53 Aguilar 2008

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They boast a V-shaped Chevron pattern across the back behind head (this is used for photo-identification)54 as well as swirls of pigmentation (“blazes”) on the right side of the head55. A dark stripe runs up and back from each eye.56 HEAD & ROSTRUM The fin whale’s highly pointed rostrum contributes greatly to its sleek look. The head occupies 20-25% of the body length57. When the mouth is closed the lower jaw extends 10-20cm beyond top of the snout.58 BLOW: The fin whale’s blow is tall and columnar59, 13’-19’6 (4-6m) in height60, 61. BALEEN 260 to 480 plates62 of black or olive green baleen63 (except for the cream-colored plates on the anterior right), a maximum of 70cm in length64, can be found on each side of the mouth. VENTRAL PLEATS 50-100 pleats65 of the fin whale’s ventral groove blubber extend from the mandible at least to the umbilicus66. FECES: Fin whale feces appear as bright orange, football-sized floating masses67. DORSAL This specie’s namesake dorsal fin can be up to 2’ (60cm) tall68, is 2.1-2.5% of the body length69, and is located at 75% of the total body length70. This is larger than that of blue whales, but lower than in Sei and Bryde’s whales. The fin whale’s dorsal is variably shaped and unique in every individual and is used by researchers for photo-identification71. The dorsal’s leading edge rises smoothly from the back at an angle of approximately 40 degrees72,73. This smooth curve is in contrast to the sharp onset of the dorsal found in sei and Bryde’s whales. FINS The fin whale’s acute tail flukes have a conspicuous central indentation.74 The pectoral fins are relatively small and lancet-like75 and are 7.5-9.9% of total body length. AT SURFACE Fin whales surface with their blowholes showing before their dorsal does. When a fin whale begins its dive, it “appears to wheel over”76 and raises its body further out of the water. However, it almost never flukes. SIMILAR SPECIES The fin whale is most easily confused with the Sei and Bryde’s whales.77 However, both these smaller species lack the distinctive facial coloration of the fin whale and their highly falcate dorsal fins are set further forward on

                                                                                                               54 Reeves et al. 2002. 55 Reeves et al. 2002. 56 Gambell 1985. 57 Gambell 1985. 58 Gambell 1985. 59 Reeves et al. 2002. 60 Gambell 1985. 61 Eder 2001. 62 Reeves et al. 2002. 63 Reeves et al. 2002. 64 Aguilar 2008 65 Ivashin 1972 66 Reeves et al. 2002. 67 Richard 1936, Orsi Relini & Cappello 1992 68 Gambell 1985. 69 Gambell 1985. 70 Aguilar 2008 71 Reeves et al. 2002. 72 Gambell 1985. 73 Leatherwood and Reeves 1983. 74 Gambell 1985. 75 Gambell 1985. 76 Gambell 1985. 77 Reeves et al. 2002.

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their backs.78 Unlike the wheel-like surfacing of fin whales, in which the blowholes first appear then the dorsal, Bryde’s whales show the two at once.79 Less similar species include the blue and humpback whales. The blue whale, besides being much larger, has blue-gray mottling, a much smaller dorsal fin, and a broad and blunter head.80 The humpback is darker, smaller, and stockier than the fin whale, and frequently raises its flukes when diving.81

Phylogeny Higher Order Within the mammalian taxon Cetacea, fin whales are mysticetes (baleen whales). The fin whale’s closest living relative is the humpback whale, from which it diverged 6-7 million years ago82. The humpack-fin clade is sister to the gray whale, and the humpback-fin-gray clade is sister to the remainder of the rorqual whales.83 The fin whale shares the same chromosome number (2n=44) with its congenics84. Although the blue whale is not the fin’s closest living relative, several blue-fin hybrids have been described.85 In one case, a female hybrid was pregnant.86 No humpback-fin whale has been described. From McGowen et al. 2009:

87 Subspecies Within the fin whale species, the taxonomy of geographically separate populations has come into question. There are small variations in body proportion and coloration among fin whales from various regions of the world’s oceans.88 For instance, flippers of whales from the northern hemisphere have been described as shorter and broader than those of their southern counterparts.89 The Society of Marine Mammalogy currently recognizes                                                                                                                78 Reeves et al. 2002. 79 Reeves et al. 2002. 80 Reeves et al. 2002. 81 Reeves et al. 2002. 82 Geisler et al. 2011, McGowen et al. 2009. 83 McGowen et al. 2009. 84 Aguilar 2008 85 Aguilar 2008 86 Aguilar 2008 87 McGowen et al. 2009. 88 Aguilar 2008 89 Aguilar 2008

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three subspecies90: B. p. physalus (Linnaeus 1758) in the Northern Hemisphere, B. p. quoyi (Fischer 1829) in the Southern, and B. p. patachonica (Burmeister 1865), the pygmy fin whale recently described by Clarke (2004) based on a stranding at the mouth of Rio de la Plata in Argentina. Given that the Isthmus of Panama closed 4 million years ago91 and that the species’ range does not extend into the Arctic passages, it is likely that Atlantic and Pacific fin whales have been isolated for some time and probably should not be considered the same subspecies.92 A 2013 paper on mitogenomic phylogenetics confirms this suspicion.93 Current data suggest that fin whales from all oceans share a common ancestor that lived 2 million years ago94. North Pacific fin whales diverged from the Southern Ocean/Atlantic Ocean population 1.94 mya95, probably due to the the immigration of a Southern Ocean group across the equator. The North Atlantic/Mediterranean fin whales diverged from the southern hemisphere group approx.1 mya96. Significant genetic97 and acoustic98 differences separate the Mediterranean and North Atlantic poplulations99. In the species’ phylogenetic tree, North Pacific fin whales are distributed in three clades, meaning they are polyphyletic and suggesting there were at least three colonizations of the North Pacific by Southern Hemisphere fin whales: the original immigration that established a North Pacific population, then two subsequent colonizations in the 2 million years since. It does not seem like this immigration was two-way; no North Pacific haplotypes are found in the Southern hemisphere. This strongly indicates that Northern Hemisphere fin whales should be assigned to multiple subspecies, but further study is needed. As a result of the Archer et al. (2013) study, the global taxonomy of the species may go into revision soon.100 North Pacific fins Perhaps as a result of these multiple colonizations of the North Pacific, fin whale population structure within the ocean basin is unclear101. Although it has been suggested that fin whale populations break into independent breeding units102, that structure has not been detected in genetic studies. Panmixia is not assumed103, but the actual structure remains unknown. Mizroch et al. (1984) cites evidence for subpopulations in the North Pacific104. Gambell (1985) writes, “In the North Pacific there is evidence from whale marking, blood typing, and morphological analyses for three stocks – on the eastern and western sides which intermingle and overlap to varying extents in the Aleutian area, and in the east China Sea.”105 The International Whaling Commission recognizes two stocks of fin whales in the North Pacific: East China Sea and the rest of the North Pacific106. The MMPA stock assessment reports recognize three stocks of fin whales in the North Pacific: 1) the Cal/OR/WA stock, 2) the Hawaii stock, and 3) the Alaska stock. The Archer et al. (2013) mitogenomics data hinted at a divide to the north and south of Pt. Conception in southern California. However, none of these studies are conclusive. Fujino (1964) suggested an isolated stock existed off BC, in addition to eastern and western N Pacific stocks.107 British Columbia fins The increasing presence of fin whales in BC waters over the course of the season and the presence of mature females throughout the season supports Fujinon’s (1964) hypothesis that BC fin whales comprise a discrete population.108 However, it is also possible that BC fin whales belong to populations to the north or south109.

                                                                                                               90 Committee on Taxonomy 2012. 91 Kiegwin 1978 92 Archer et al. 2013 93 Archer et al. 2013 94 Archer et al. 2013 95 Archer et al. 2013 96 Archer et al. 2013 97 Berube et al. 2002, Berube et al. 1998. 98 Castellote et al. 2011. 99 Notobartolo-di-Sciara et al. 2003. 100 Archer et al. 2013 101 Gregr et al. 2006. 102 Aguilar 2008, Berube et al .1998 103 Carretta et al. 2011. 104 Mizroch et al. 1984 105 Gambell 1985. 106 Donovan 1991 107 Fujino 1964. 108 Gregr et al. 2000 109 Gregr et al. 2006.

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According to Gregr et al. (2006), “there is no way to presently determine whether animals sighted in Pacific Canadian waters are from either of the [California-Oregon-Washington and Alaska] stocks defined by NMFS [National Marine Fisheries Service]. Indeed, there is currently no evidence to determine whether these two putative stocks are truly distinct populations or whether they represent a single, eastern North Pacific population.”110 These populations are thought to move seasonally between international, Canadian, US, and possibly Mexican territorial waters.111

Range and Habitat General The fin whale can be found in all major oceans112, 113, but it is usually confined to the temperate and polar latitudes114,115. It is almost never seen near the equator or the ice limit in extreme latitudes116. In the North Pacific, fin whales are rarely encountered south of the Baja Peninsula117. Fin whales can also be found in relatively confined seas. Fin whales are the only mysticetes found regularly in the Mediterranean Sea.118,119, where an estimated 900 individuals concentrate in the Corsican-Ligurian Basin120. There is also a resident population in the Gulf of California.121 However, there are similar habitats in which they are not found: the eastern Mediterranean, the Baltic Sea, the Persian Gulf, the Red Sea, and equatorial seas.122 British Columbia    Fin whales occur in Pacific Canadian waters year-round, with highest numbers seen in the summer months123. The range of the whales that occur in BC waters is unknown but it may extend from the Aleutians to Southern California124. During the summer, the fin whale’s distribution is concentrated between Vancouver Island and the Aleutians125. Historically, fin whales were observed in BC’s exposed coastal area (Hecate Strait and Queen Charlotte Sound) and occasionally in more protected waters of Queen Charlotte Strait, the Strait of Georgia, and the Intra-Coastal Zone (ICZ)126. Only 17% of fin whale kills (1908-1967) for which positions were recorded by BC coastal whalers were on the continental shelf127. The Kitimat Fjord System was one of the few areas within the ICZ that were used by fin whales, which may speak to the unique inland habitat it offers128. Whaling data (1943 – 1967, from Coal Harbour) suggest a seasonal trend in fin whale presence in the study area and other sectors of the ICZ. Next page, from Gregr and Trites (2001): Fin whale monthly probability predictions overlaid with the positions of whale kills between 1943 and 1967. Dark areas represent a high probability of whale occurrence129. The zoomed-in figures of the study area are screenshots taken by EM Keen.

                                                                                                               110 Gregr et al. 2006. 111 Gregr et al. 2006. 112 Reeves et al. 2002. 113 Reeves et al. 2002. 114 Reeves et al. 2002. 115 Aguilar 2008 116 Aguilar 2008 117 Hamilton et al. 2009. 118 Jahoda et al. 2003. 119 Reeves et al. 2002. 120 (Forcada et al.1995) 121 Gambell 1985. 122 Aguilar 2008 123 Gregr et al. 2006. 124 Spalding 1998. 125 Spalding 1998. 126 Pike and MacAskie 1969, Gregr et al. 2002, Gregr 2004. 127 Gregr et al. 2006. 128 Pilkington et al. 2011. 129 Gregr and Trites 2001

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a) April (44 kills, 0 in study area). b) May (404 kills, 0 in study area)

c) June (620 kills, 1 in study area) d) July (727 kills, 6 in study area)

e) August (828 kills, 3 in study area) f) September (410 kills, 10 in study area)

April: None caught in ICZ (Intra-Coastal Zone) or study area. May: One in Hecate Strait, 2 near Dixon Entrance. None caught in ICZ or study area. June: 9 caught within Hecate Strait, 8 in Dixon Entrance, and 1 in the study area: 1 right on the border of

Caamano Sound. July: Many in QC Sound, 10 In Hecate Strait, 20 in Dixon Entrance, 5 in outer Milbanke Sound, and 7 in the

study area: 4 in Caamano Sound, 1 in Campania Sound, 1 in what looks like Taylor Bight (south Gil Island), 1 in Estevan Sound,

August: 26 in Eastern Hecate Strait, 15 in Dixon Entrance, 7 in Milbanke Sound (one of which is FAR up a fjord), and 3 in the study area: 2 in Campania, 1 near Fawcett Pt in Squally Channel.

September: 8 in E Hecate, 2 in QC Sound, 1 far up fjords in Milbanke Sound, 7 in Dixon Entrance, and 9 in the study area: 8 in Caamano Sound, 1 near fin island in Squally.

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Over the course of BC whaling’s second era (1948 to 1967), the distance of capture from the nearest shore increased significantly130 until they were extirpated from coastal waters entirely. In the early 2000’s, sightings of fin whales in Pacific Canadian waters were predominantly from the west coast of Vancouver Island, Hecate Strait, and QC Sound, and occurred in summer and winter.131 Fin whales were considered to prefer offshore waters but to sometimes stray into Hecate Strait, Queen Charlotte Sound, and occasionally Queen Charlotte Strait132. The waters off Vancouver Island have been suggested to contain a summer feeding aggregation.133 The Department of Fisheries and Oceans (DFO)-is actively surveying the coast to define critical habitat areas for BC waters (see “Current Research” section at end). These DFO surveys found fin whales widely distributed over BC’s continental shelf with concentrations found in the most productive whaling grounds: west of Vancouver Island and Haida Gwaii, in Queen Charlotte Sound, in southern Hecate Strait, and in Dixon Entrance. They were also encountered in the confined waterways of Caamano Sound and Squally Channel in the Kitimat Fjord System.134 Proximity to Shore Most of the literature refers to fin whales as a shelf, slope, and/or offshore species:

“Tend to concentrate in coastal and shelf waters, but can be found in the deep ocean.”135

“Occurs throughout the world in both near-shore and deep waters beyond the edge of the continental shelf.”136

“The only recent record from sheltered waters is of a dead fin whale pushed into Tacoma in 1985 by the bow of a freight vessel.”137

“Mediterranean fin whales are sighted almost exclusively offshore138 (mean water depth of sightings: 2,248m139), though there have been instances of near-coast sightings140.”

However, fin whales have been seen in the inland/intra-coastal waters of the Gulf of St. Lawrence141 and Bay of Fundy. Seasonal Movements Southern Ocean fin whales are known to follow migration patterns typical of intermediate-sized mysticetes: regular seasonal movements between temperate waters where mating and calving occur, and more polar feeding grounds (Mackintosh 1965)142, 143. They move toward poles in spring and toward the equator in fall.144 Their migration routes seem to follow areas of low geomagnetic intensity and gradient145. Not all components of the population’s demographics move together; pregnant females are the first to initiate movement146 while lactating females and juveniles of both sexes are the last to migrate.147 It is thought that northern and southern hemisphere populations do not come into contact because of their alternate migration schedules.148 This lack of contact has led to genetic isolation.

                                                                                                               130 Gregr et al. 2000 131 Gregr et al. 2006. 132 Spalding 1998. 133 Pike and MacAskie 1969 134 Ford et al. 2010a 135 Reeves et al. 2002. 136 Creswell et al. 2007. 137 Spalding 1998. 138 Jahoda et al. 2003. 139 Zanardelli et al 1992 140 Notobartolo-di-Sciara et al. 2003. 141 Sergeant 1977, Ray et al 1978 142 Gambell 1985. 143 Aguilar 2008 144 Leatherwood and Reeves 1983. 145 Aguilar 2008, Walker et al. 1992. 146 Aguilar 2008 147 Aguilar 2008 148 Aguilar 2008

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Earlier literature (and some field guides still) report similar migration patterns in the northern hemisphere149,150. There does seem to seasonal movement of some kind, but its pattern and extent is still unclear151 , 152. Mediterranean fin whales remain in their semi-enclosed basin year-round, meaning they calve there153. In the Northeast Pacific, fin whales have been said to winter in waters from southern California down to 20 degrees South154, and summer from central California into the Chuckchi Sea.155 However, northeast Pacific (NEP) fin whales are found over a broad latitudinal range throughout the year. Some whales remain in higher latitudes during the coldest months of the year or in low latitudes year-round if food is available156. BC whaling records show that fin whales were regularly caught north of 40 degrees latitude in the winter months157. There are fin whale aggregations year-round in southern/central California158,159,160 , in the Gulf of California161, in summer in Oregon162,163, and in summer/autumn in Shelikof Strait/ the Gulf of Alaska164. Acoustic signals are detected year-round off northern California, Oregon, and Washington, with a concentration of activity between Sept. and Feb165. It is unknown where BC fin whales go in the winter. In the BC whaling years, distance from shore changed significantly from month to month for female fin whales.166 However, the proportion of pregnant females killed remained constant, suggesting reproductive state did not influence seasonal movements167. The 20 Hz call has been observed offshore of BC, suggesting that fin whale breeding may be occurring in this area168. Conversely, whaling data do suggest age-structured movements. Larger individuals arrived in British Columbian waters ahead of small ones169 and shelf catches were primarily of immature animals170,171, suggesting that BC waters were a feeding ground primarily for sub adult animals172. These loose patterns (or lack thereof) have led to several alternative hypotheses to stereotyped migration. It has been suggested that some populations may shift in winter to occupy the summer habitats of others173. But the prevailing idea is that, in the North Pacific, fin whale individuals “that concentrate near the coast in the feeding season tend to disperse into open waters during the winter, therefore being more difficult to detect.”174

                                                                                                               149 Leatherwood and Reeves 1983. 150 Spalding 1998 151 Mizroch et al. 2009 152 Aguilar 2008 153 Notobartolo-di-Sciara et al. 2003. 154 Leatherwood and Reeves 1983. 155 Leatherwood and Reeves 1983. 156 Reeves et al. 2002. 157 Mizroch et al. 2009 158 Dohl et al. 1983 159 barlow 1997 160 Forney et al 1995 161 Tershy et al 1993 162 Green et al 1992 163 McDonald 1994 164 Brueggeman et al 1990 165 Moore et al 1998 166 Gregr et al. 2000 167 Gregr et al. 2000 168 Ford et al. 2010. 169 Gregr et al. 2000 170 Nichol and Heise 1992 171 Gregr et al. 2000 172 Pike and MacAskie 1969 173 Reeves et al. 2002. 174 Aguilar 2008

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Reproduction Life History Fin whale population sex ratio is 1:1175. Sexual maturity is reached at 18.3m (females) and 17.7m (males) in the north Pacific176, and 20m and 19m, respectively, in the southern hemisphere177. These lengths correspond to approximately 6-7 years in males and 7-8 years in females.178 Age of first parturition and birth rates diminished during the height of whaling.179 Males grow faster than females but stop growing sooner.180 Physical maturity (defined by ossification of vertebral column) happens at ~25 yrs. in both sexes.181 Longevity has not been determined, but individuals of up to 80-90 years old are known.182 Mating Very little is known about the species’ mating system183; mating has never been observed. No distinct breeding or calving grounds have been identified for the fin whale.184 Considering the proportionally reduced mass of male fin whale testes, the species’ mating system probably relies on competition among males for females (with no sperm competition).185 Chasing has been observed among pairs of fin whales or groups of three in late autumn on the feeding grounds. This behavior is possibly related to courtship.186 Interspecific mating between the fin and blue whales has been known to occur.187 In the northern hemisphere, the mating period is Dec. – Feb.; in the southern hemisphere, it is May to July188. Breeding – calving and mating is believed to occur in the winter189,190. In the Mediterranean, births peak in Nov. but occur throughout the year at lower rates191. Calving Gross pregnancy rates (number of pregnant females in relation to that of adult females) are between 38-49%192. Gestation is 11-12 months.193 Females give birth every 2-3 years to a single calf194. Twins have been observed in utero, but there is no evidence that any survive195. Calves stay with their mother for 6-8 months196, when the calf is 11-13m long197. This weaning is followed by 6 months of rest and recuperation for the mother, after which mating takes place again.198

                                                                                                               175 Clapham et al 1997 176 Gambell 1985. 177 Aguilar 2008 178 Aguilar 2008 179 Gambell 1985. 180 Aguilar 2008 181 Aguilar 2008 182 Aguilar 2008 183 Reeves et al. 2002. 184 Reeves et al. 2002. 185 Aguilar 2008 186 Reeves et al. 2002. 187 Reeves et al. 2002. 188 Aguilar 2008 189 Reeves et al. 2002. 190 Leatherwood and Reeves 1983. 191 Notobartolo-di-Sciara et al. 2003. 192 Leatherwood and Reeves 1983. 193 Reeves et al. 2002. 194 Reeves et al. 2002. 195 Reeves et al. 2002. 196 Reeves et al. 2002. 197 Aguilar 2008 198 Aguilar 2008

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Behavior Surface Behavior Fin whales practice a wheel-like surfacing, in which the blowholes first appear then the dorsal199. The fin whale rarely raises its flukes when diving200,201. Aerial Behavior It seems that, in most populations of fin whales, breaching is rare.

“Breaching is very rare, except when the whale is harassed.” 202 “They do breach on occasion.” 203

This author has observed fin whales porpoising in the confined channels of BC in what looked like a particularly energetic feeding frenzy. The one population in which breaching seems more regular is in the Mediterranean Sea. Breaching was observed in 4% of sightings in the Tyrrhenian Sea204, and in 6.9% of sightings in the Ligurian-Corsican-Provencal Basin (1990-1999).205 The mean number of consecutive breaches there is 2.45 (range=1-6)206.

207

                                                                                                               199 Reeves et al. 2002. 200 Reeves et al. 2002. 201 Gambell 1985. 202 Reeves et al. 2002. 203 Leatherwood and Reeves 1983. 204 Marini et al 1996c 205 Notobartolo-di-Sciara et al. 2003. 206 Notobartolo-di-Sciara et al. 2003. 207 www.tethys.org

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Speed Bursts Known as the “greyhound of the sea”208, fin whales are the fastest of the large whales209. Their top speed bursts have been reported as 11 knots210, 25 knots211, “over 20 knots”212, “in excess of” 17.3 knots213, and 15 knots214. Sustained swimming Sustained cruising speed has been reported as 5-8 knots215, 3.5-7.5 knots in the Northeast Pacific (discerned through acoustic localization) 216,, 4.9 knots217, 2.0-7.8 knots near Iceland218, and 2.7 knots in the Mediterranean (averaged from 24 focal follows).219Tag data suggest that during sustained travel fin whales cover between 90 and 180 miles a day220. Respiratory Cycles A respiratory cycle is composed of a dive, or prolonged apnoea phase, then a near-surface period of frequent breathing. The most rigorous respiration interval study for fin whales was conducted in the Mediterranean: 2068 breaths organized in 477 respiratory cycles were recorded.221 In the Mediterranean, the near-surface period averaged 62 seconds in length with an average of 4.6 breaths222. Diving Fin whales practice very shallow dives between their breaths followed by a deep dive during their prolonged apnoea phase. The mechanics and evolution of fin whale dive behavior has been thoroughly studied (see the Lunge Feeding Backgrounder for more). The average dive time for Mediterranean fin whales was 3.75 minutes223. The longest recorded dives are as long as 16.9 minutes, which is much shorter than their Theoretical Aerobic Dive Limit (TADL) of 28.6 min224. This is due either to the dispersal behavior of prey or the high energetic cost of foraging225. The deep dive occurs during both traveling and feeding, though the depth and duration of the dive changes by behavioral context. Foraging dives are deeper and longer in duration than traveling dives and are characterized by vertical excursions where lunging occurs226. In California waters, fin whales dived to 97.9m and for 6.3 min when foraging and to 59.3m and for 4.2 min when not foraging.227 In a foraging dive, lunges into the prey field occur at depth. These lunges are vertical excursions of 21.2m or more that occur an average of 1.7 times per dive (maximum of 8) 228. Lunges seem to occur during ascent, making the rate of ascent higher than that of descent during a dive229. Dive Depth Fin whales are generally deeper divers than blue and sei whales. Median depths reported for foraging dives (deeper than traveling dives) are between 100 and 230m230. The deepest dives known for the species have been observed in the Mediterranean231. Dive depths exceeding 470m have been observed twice, dives of at                                                                                                                208 Reeves et al. 2002. 209 Reeves et al. 2002. 210 Watkins 1981 211 Reeves et al. 2002. 212 Gambell 1985. 213 Leatherwood and Reeves 1983. 214 Aguilar 2008 215 Aguilar 2008 216 McDonald et al. 1995. 217 Ray 1978. 218 Watkins et al 2006 219 Lafortuna et al. 2003. 220 Watkins et al. 2006 221 Lafortuna et al. 2003. 222 Lafortuna et al. 2003. 223 Lafortuna et al. 2003. 224 Croll et al. 2001. 225 Croll et al. 2001. 226 Croll et al. 2001. 227 Croll et al. 2001. 228 Croll et al. 2001. 229 Croll et al. 2001. 230 Leatherwood and Reeves 1983. 231 Panigada et al 1999

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least 150m were routinely performed around sunset232, and the maximum observed dive depth on record is 565m233. Social Associations Mean Group Size Fin whales are usually found travelling alone or in small groups234. Here we define a group, after Notobartolo-di-Sciara et al. (2003), as “affiliations in which two or more individuals swim side by side within 1-2 body lengths and generally coordinate at least their surfacing and diving, as well as their speed and direction of movement.”235 In the Mediterranean mean group size is 1.4, which is slightly less than reported elsewhere.236 In our study area, the confined channels of BC’s north coast, observations from 2007-2010 reveal an average group size of 1.8, ranging from 1 to 6.237 Field guides have cited common group sizes of “single or in pairs…3 to 10 to 20” 238 and “3 to 7 individuals”.239 Larger groups may coalesce into a broad concentration of 100 or more unassociated individuals240 on feeding grounds.” 241 Group Stability Stable groups appear to be rare in fin whales242. The only stable social interaction is between mom-calf pairs and this interaction vanishes at weaning243. Fin whales are sometimes associated with blue whales244 and humpback whales245.

                                                                                                               232 Notobartolo-di-Sciara et al. 2003. 233 Panigada et al. 2003. 234 Reeves et al. 2002. 235 Clapham 2000. 236 Notobartolo-di-Sciara et al. 2003. 237 Pilkington et al. 2011. 238 Gambell 1985. 239 Leatherwood and Reeves 1983. 240 Reeves et al. 2002. 241 Gambell 1985. 242 Reeves et al. 2002. 243 Aguilar 2008 244 Reeves et al. 2002. 245 this author, unpubl.

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Acoustics All fin whale populations of the world are known to vocalize. Both male and female fin whales make very loud, low-frequency vocalizations that can travel over hundreds of miles in deep water246 (reviewed in Edds-Walton 1997). The bandwidth used by fins is 14-750 Hz, with most of their energy in the 20-40 Hz range247. Propagation Fin whale moans can be heard for at least tens and probably hundreds of kilometers. Payne and Webb (1971) estimated source level of 65-100 dB re: 1 dyne/cm2 at 1 yard, probably 80 dB248. This call could be heard from 45 miles away (assuming spherical loss) to 525 miles (through the deep-sound channel). In ideal conditions (which would never occur in reality), 700 miles would be the upper limit249. In practice, Southern Ocean fin whales have been detected up to 56km from the recorder. The average source level for these calls is 189 + - 4dB re: 1 microPa -1m over 15-28 Hz.250 The implications of this propagation is that “fin whales…may be in tenuous acoustic contact throughout a relatively enormous volume of ocean and that such contact might be of use for finding each other or for joining, or keeping together in, widely dispersed herds”251. Adaptive Acoustics The reason for fin whale vocalizations may never be fully understood. The long-distance communication may allow mating to occur without the need for aggregating on breeding grounds; this may explain why winter breeding grounds have not yet been found for fin whale populations252. If male fin whale songs (the 20 Hz pulses, below) attract females from great distances to aggregations of patchily distributed prey, as has been proposed253 , there could be some functional association between food resources and male reproductive display254. Fin whale calls are probably not acting as vertical sonar; they are too low frequency and unnecessarily loud for that255. However, it is possible that the calls serve as a kind of horizontal radar256, using countercalling among the pod to measure oceanic sound speed profiles257. The amplitudes of the horizontal travel path of an incoming call and those of its reflected paths off the sea surface and seafloor could be compared. The ratios of these amplitudes may be indicative of sound speed and temperature profiles, which are results from changes in the depth of the thermocline (which control productivity in an area). If so, fin whales may be able to remotely assess the productivity of a foraging ground.258 This needs to be tested, and who the heck knows how that is going to happen? Call Types and Descriptions Regional differences in fin whale call types have been found between Gulf of California and several Atlantic and Pacific regions259,260,261. Worldwide, however, calls can be placed into 3 general groups: 1) the stereotyped, “classic” 20 Hz pulse; 2) the irregular higher frequency (30-90 Hz) downswept call; and 3) “other” 262,263. All of their calls are frequency modulated downward.

                                                                                                               246 Reeves et al. 2002. 247 Edds 1998 248 Payne and Webb 1971. 249 Payne and Webb 1971. 250 Sirovic et al. 2007. 251 Payne and Webb 1971. 252 Payne (2004) 253 Croll et al 2002 254 Clark et al 2002 255 McDonald et al. 1995. 256 Clark 1993 257 McDonald et al. 1995. 258 McDonald et al. 1995. 259 Aguilar 2008 260 Hatch and Clark 2008 261 Clark et al. 2002. 262 Gambell 1985. 263 McDonald and Fox 1999

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Regular 20 Hz Downswept Pulse The classic, stereotyped pulse contains energy between 28 and 15 Hz264 with most around 20 Hz, has a downward sweep in frequency of about 6 Hz265, and has little or no harmonic energy. In the Mediterranean, this pulse modulates from 25 to 19 Hz266. The pulse direction is 0.8s267 to 1s268,269. The interval between pulses ranges from 6 to 46 seconds270, with reported averages of 30 seconds271 and 19 seconds in the California Current272. From Ford et al. 2010b:

It is believed that the 20 Hz pulse is a male breeding call273,274,275,276. In the Eastern North Pacific, the 20 Hz pulse is seasonal with most calling occurring between October and April, coinciding with the winter breeding period, and little calling occurring during the summer months277. These calls have been heard in association with courtship displays (Watkins 1981) in the W North Atlantic278, perhaps to attract females from great distances to prey aggregations279. Irregular 30-90 Hz Downswept Pulse These “high frequency” fin whale calls are still all under 100Hz. The downsweep has been reported to occur between 75 and 40 Hz280 or between 90 and 30 Hz 281. The irregular series can last from only a few minutes to more than a day. These irregular calls generally represent only a small fraction of fin whale calls, but in the

                                                                                                               264 Castellote et al. 2012. 265 Watkins et al. 1987, Thompson et al. 1992 266 Notobartolo-di-Sciara et al. 2003. 267 Ford et al. 2010. 268 Notobartolo-di-Sciara et al. 2003. 269 Schevill et al 1964, Thompson et al 1979, Watkins et al 1987, Richardson et al 1991, Thompson et al 1992 270 Watkins et al 1987 271 Rothenberg 2008. 272 McDonald et al. 1995. 273 Watkins et al 1987 274 Croll et al 2002 275 Watkins et al 1987 276 Reeves et al. 2002. 277 Watkins et al 1987 and Watkins et al 2000 278 Gambell 1985. 279 Croll et al 2002 280 Gambell 1985. 281 McDonald and Fox 1999

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North Pacific during the summer they can comprise 90% of all calls recorded282. Irregular calls seem to be used in a variety of behavioral contexts283, perhaps for communication among nearby fin whales284 From Ford et al. 2010b:

Other “Ragged” low-frequency pulses and “rumbles” have also been reported285. In the Mediterranean, a narrow-band pulse type (1s, 22-19 Hz), referred to as a “back beat”, has been observed286. Pacific Northwest Seasonality Fin whale calls are heard year round on recorders offshore in the Gulf of Alaska and Bering Seas, with a broad peak in the fall287. DFO’s Cetacean Research Program has 9 seafloor-mounted acoustic recording packages (5 – 1000 Hz) monitoring British Columbia’s coast.

288                                                                                                                282 McDonald and Fox 1999 283 McDonald and Fox 1999 284 Gambell 1985. 285 Gambell 1985. 286 Notobartolo-di-Sciara et al. 2003. 287 Moore et al. 2006

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From Nichol and Ford (2011 Fig 2). “Passive acoustic monitoring sites. Langara Is.: winters 2005 to 2009, year-round 2009 to 2011; Bowie Seamount: deployed July 2011; Cape St. James: May 2008 to May 2011; Triangle Island: deployed May 2011; Brooks Peninsula: deployed July 2010; Union Seamount: February to June 2006; LaPérouse Bank: May to September 2007; Barkley Canyon NEPTUNE: operational December 2009; Swiftsure Bank: May 2009 to May 2011.”

The Union seamount and La Perouse Bank recorders have been analyzed thus far. Fin whales were detected frequently on both, showing seasonality in occurrence289. The stereotyped 20 Hz pulse call was detected a low number of times in February and April at Union Seamount and not at all at La Perouse Bank290. Irregular repetition interval calls were detected at fairly consistent but low rate at both sites with the exception of a notable increase in August-September at LaPerouse Bank.291 From Ford et al. 2010b:

                                                                                                                                                                                                                                                                                                                                                                                           288 Nichol and Ford (2011). 289 Ford et al. 2010. 290 Ford et al. 2010. 291 Ford et al. 2010.

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Food and Foraging Diet General The species’ diet has been described with some variation among sources: “[Fin whales eat] Krill and various small schooling fish, notably herring, capelin, and sandlance.”292

“[They feed] mainly on planktonic crustaceans but also consume some fish and cephalopods. There is considerable variation by area and season…In both the North Atlantic and North Pacific oceans fish are commonly taken: herring, cod, mackerel, Pollock, sardine, and capelin, together with squid and euphausiids and copepods. Just what is consumed is probably determined by availability as much as preference in these northern areas.”293

“Fin whales eat a wide variety of food, including krill and other invertebrates, capelin, sand lance, squid, herring, and lanternfish.”

“[They] feed on a wide variety of organisms, depending on availability294. Possibly, diet varies with season and locality.” 295

There is no difference in diet between male and female whales.296 Other Regions Mediterranean fins have never been observed feeding on anything other than the euphausiid species Meganytiphanes norvegica297 (this includes investigations of stomach contents)298. In the southern hemisphere, fin whale diet is almost exclusively krill (Euphausia vallentini) but also E. superba and sometimes the copepod Calanus tonsus299. The summer diet of fin whales in the North Atlantic is dominated by crustaceans300:

“Preferred prey… seems to be krill composed of the euphausiid Meganyctiphanes norvegica, although other species of planktonic crustaceans (Thysanoiessa inermis, Calanus finmarchicus), schooling fishes such as capelin (Mallotus villosus), herring (Clupea hargengus), mackerel (Scomber scombrus) and blue whiting (Clupea harengus), and even small squids are also consumed.”301

North Pacific It has been estimated that North Pacific fin whales consume between 490,329 and 488,367 tons of prey annually302. Another study yielded a lower estimate: 318,582 tons annually303. One study estimates that individual fin whales eat up to 1 ton of euphausiids per day304. North Pacific fin whales eat planktonic crustaceans, schooling fishes, and squids – in that order of preference305. In the California Current, large zooplankters are thought to constitute 70%306-80%307 of fin whale diet308. The remainder is small squid (5%), “small pelagic fish” (5%), mesopelagic fish (5%), and “miscellaneous fish” (5%),                                                                                                                292 Reeves et al. 2002. 293 Gambell 1985. 294 Kawamura 1980 295 Aguilar 2008 296 Flinn et al. 2002 297 Notobartolo-di-Sciara et al. 2003. 298 Notobartolo-di-Sciara et al. 2003. 299 Aguilar 2008 300 Flinn et al. 2002 301 Aguilar 2008 302 Barlow et al. 2008. 303 Trites et al. 1997. 304 Aguilar 2008 305 Gregr et al. 2006. 306 Kawamura 1980 307 Barlow et al. 2008. 308 Barlow et al. 2008.

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resulting in an effective trophic level of 3.4309. In the Aleutians the whaled fins had a higher proportion of fish, mainly sardines and pollack, in their stomach310. The most important zooplankton prey taxa are several species of euphausiids and calanoid copepods311. Kawamura (1982) hypothesized that fin whales in the nearby Gulf of Alaska prey switch from euphausiids (abundant in late spring and early summer) to copepods (most abundant in summer and fall). BC fins The majority of North Pacific diet studies pertain to regions farther offshore or north of BC312. British Columbia’s historical whaling database reveals regional prey preferences. Stomach data comes from studies between 1963 and 1967, when data collection was higher in quality and more consistent313. 578 of 650 killed fin whales were examined for stomach contents between 1963 and 1967 314 . Non-empty FW stomachs contained mainly (sometimes exclusively) euphausiids in all five years 315, but the proportion of diet components changed significantly between years. In 1964 and 65, higher percentages of copepods, fish, and cephalopods were found. The euphausiids preyed upon by fin whales included Euphausia pacifica, Thysanoessa spinifera, T. longipes, and T. inermis. Important calanoid species include (Neo)Calanus cristatus, C. plumchrus, C. finmarchicus, and Metridia lucens316. Low frequency of fish in the fin whale diet is surprising when compared to the larger role fish play in other areas317. Fish in the diet of the whales killed off the BC coast were primarily found in stomachs of whales killed in the spring318. Timing of Feeding Seasonal Both Southern and Northern Hemisphere populations are thought to feed in the summer and fast while breeding in the winter319,320. Daily Feeding activity seems to escalate in the crepuscular hours. Tagged individuals in the Mediterranean reveal diving behavior that coincides with the evening vertical migration of their prey, M. norvegica. The tagged whale began performing deep (greater than 400m) dives at 5:00pm321, then showed a marked decrease in diving depth, dropping from 400m at 7:40pm to less than 50m by 9:50pm322.

                                                                                                               309 Barlow et al. 2008. 310 Spalding 1998. 311 Kawamura 1980, Nemoto 1957, Nemoto 1959, Nemoto and Kasuya 1965 312 Kawakami 1980, Kawamura 1980, 1982; Nemoto and Kasuya 1965, Pike 1950. 313 Flinn et al. 2002 314 Flinn et al. 2002 315 Flinn et al. 2002 316 Flinn et al. 2002 317 Flinn et al. 2002 318 Flinn et al. 2002 319 Aguilar 2008 320 Aguilar 2008 321 Panigada et al. 2003. 322 Panigada et al. 2003.

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From Panigada et al. 2003:

Lunging As rorqual whales, fin whales feed by lunging323. Lunges may involve bursts of speed up to 25 knots324. Lunges can occur at depth or at the surface, though there may be some geographic structure to such behavior (which may correspond to the primary prey of a region). In the Ligurian Sea, where deep euphausiids comprise the majority of fin whale diet, feeding behavior at the surface has not been observed325. See the dedicated Lunge Feeding Backgrounder for more information. Use of Right Jaw The following information comes from field guides that do not cite their sources:

“Lunge-feeding is commonly seen, during which the whale often turns on its right side.”326

“Fin whales have been observed feeding at the surface by swimming on their right sides and making a lateral scoop with the open mouth and distended throat region.”327

“It lunges at concentrated prey as other Mysticetes do, but the fin whale turns on its right side and swings on its right flipper while lunging. In this position, the dark left side and white lips on the right side maintain the whale’s countershading, making it less obvious to its prey.”328

Foraging Fin whales are known to associate with their prey and/or the oceanographic conditions that tend to aggregate their prey, such as oceanic fronts329. In BC high, localized abundance of euphausiids around bathymetric margins allows the fin whales to migrate to the continental shelf and feed continuously throughout summer330. In the Bay of Fundy, fin whales occurred primarily in shallow areas with high topographic relief and these occurrences were correlated with herring and euphausiid concentrations331. In summer in the northeastern US and Bay of Fundy distribution is associated with low surface temperatures332. In the Mediterranean fin whale sightings are very closely correlated to high M. norvegica densities in very deep waters333. Due to this deep diving, associations between whales and marine birds are extremely scarce in the Ligurian-Corsican-Provencal Basin334. In the Gulf of St. Lawrence, distribution of blue, fin, and humpback whales was associated with the

                                                                                                               323 Reeves et al. 2002. 324 Reeves et al. 2002. 325 Notobartolo-di-Sciara et al. 2003. 326 Eder 2001. 327 Gambell 1985. 328 Spalding 1998. 329 Hain et al. 1992 330 Flinn et al. 2002 331 Woodley and Gaskin 1996 332 Woodley and Gaskin 1996 333 Relini et al 1994 334 Notobartolo-di-Sciara et al. 2003.

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formation of sea surface temperature (SST) gradients335. The temperature gradient itself may have a herding effect on krill336.Thermal fronts are created by tide or wind-induced upwelling and this upwelling may also increase productivity and attract whales. Island wake systems are known to predictably aggregate plankton and weak nekton337. In the Bay of Fundy, plankton and weak nekton concentrated in eddies behind islands where tidal currents were reduced in flow and vorticity was high338. The mesoscale eddies that occur to the south of Haida Gwaii are considered prime habitat for fin and blue whales in BC waters339. Beyond this, however, there has been limited effort to relate whale distribution to habitat features in BC waters340,341. Competition Given the fin whale’s diverse diet and large range, interspecific competition is likely to occur. It is especially likely that fin whales compete with blue whales342 and humpback whales. Trites et al. (1999) suggested that some species of fish are significant competitors of whales in the Bering Sea.343 Swordfish swords have been recovered from fin whale carcasses on multiple events344; who the hell knows why swordfish have problems with fin whales. Predation Fin whales can be victims of killer whale attack345,346. They are often seen with signs of past attack on flippers, flukes, and flanks347. A mother-calf pair was seen being pursued by transients in the study area (H. Meuter, pers. comm.). Other potential predators include false killer whales and the white shark348. Parasites Ectoparasites The fin whale’s external parasites are primarily crustaceans349. The copepod Penella balaenoptera is particularly common in the southern hemisphere350 and the Mediterranean351. Cirripeds (barnacles) and Cyamus whale lice infect fin whales in warmer waters but are lost in the colder southern waters352. In adults skin of the rear trunk flanks are covered by round scars and stripes attributed to the attachment of lampreys and remoras353. Marine lampreys (Petromyzon marinus) seem to be particularly common ectoparasites in the Mediterranean354. Diatoms are contracted as surface films in the summer months in the Antarctic355,356; this has also been observed in BC waters (this author).

                                                                                                               335 Doniol-Valcroze et al. (2007) 336 Doniol-Valcroze et al. (2007) 337 Wolanksi & Hamner 1988 338 Johnston et al. 2005. 339 Nichol and Ford 2011 340 Dalla Rosa (2010). 341 Gregr and Trites 2001 342 Aguilar 2008 343 Trites et al. 1999. 344 Gambell 1985; Jongsgard 1962; Peers and Karlsson 1976 345 Leatherwood and Reeves 1983. 346 Vidal and Pechter 1989 347 Aguilar 2008 348 Connor 2000 349 Gambell 1985. 350 Gambell 1985. 351 Richiardi 1874, Anthony & Calvet 1905 352 Gambell 1985. 353 Aguilar 2008 354 Notobartolo-di-Sciara et al. 2003. 355 Gambell 1985. 356 Aguilar 2008

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357 Endoparasites Internal parasites, found during the whaling years and in necropsies, include tapeworms and Acanthocephala (Mackintosh and Wheeler 1929)358.

                                                                                                               357 http://www.tparazitolderg.org/text.php3?id=285 358 Gambell 1985.

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Whaling Fin whales have been hunted in larger numbers than any other whale species in the 20th century359. Before the invention of steam power and the explosive harpoon, fin whales were too fast to catch360. They became targeted at the onset of modern operations in 1870361 and after the severe depletion of blue whale stocks362. In the 20th century, 725,000 fin whales were killed in the Southern Hemisphere alone363. However, due to the fin whale’s greater original abundance, it was not as severely depleted as the blue whale (roughly 300,000 blues were taken, reducing their population to 0.1% of historical abundance).364

North Pacific 46,000 fin whales were taken from the North Pacific by commercial whalers between 1947 and 1987365. From 1919 to 1965, 5,000 were taken from the US west coast366,367,368. The ecological repercussions of these whaling efforts have been a topic of exciting scientific debate369.

Aboriginal “[The] fin whale was the largest species eaten by the Makah and Quileute off the Olympic Peninsula, though aboriginal whalers may have taken this large species only when they found a dying whale floating.”370 Current Whaling In 1983, the fin whale was still hunted in Iceland (Hvalfordur Station) and Spain371. The Spain program was discontinued in 1985, and Iceland stopped hunting fins in 1989372 but reinitiated in 2006. In 2009, fin whales were still exploited in West Greenland (19 individuals per year), Antarctica (3-10 individuals per year since 2005 under the Japanese Special Permit Program), and Iceland (7 individuals were taken in 2006 when whaling was reinitiated) 373.

British Columbia From 1908 to 1967, at least 7,605 fin whales were killed374, 375. The history of these whaling operations is covered in its own Backgrounder. The catch records are reviewed in Pike and MackAnsie (1969), Nichel and Heise (1992), and more recently in Nichol et al. (2002) and Gregr et al. (2006). Fin whales, which were locally depleted by the end of commercial whaling, represent one third of total kills from BC stations376,377.  The largest fin whale catches occurred in the 1950s and 1960s378. In the first era of modern BC whaling (pre-World War II), their contribution to total catch declined from a peak in 1912 to less than 20/yr in 1942.379 In the second era all kills were from the Coal Harbour station and there was a dramatic rise and fall in the number of fin whales caught, peaking in 1958 with 573 animals380.

                                                                                                               359 Reeves et al. 2002. 360 Reeves et al. 2002. 361 Aguilar 2008 362 Leatherwood and Reeves 1983. 363 Reeves et al. 2002. 364 Aguilar 2008 365 Gregr et al. 2006. 366 Rice 1974 367 Tonnessen and Johnsen 1982 368 Clapham et al 1997 369 e.g., Trites et al. 1999. 370 Spalding 1998. 371 Gambell 1985. 372 Aguilar 2008 373 Aguilar 2008 374 Gregr et al. 2000 375 Gregr et al. 2006. 376 Gregr et al. 2000 377 Mizroch et al 1984 378 Gregr et al. 2006. 379 Gregr et al. 2000 380 Gregr et al. 2000

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In all months, male and female fin whales were caught in roughly equal numbers381,382. In both sexes, the shortest fin whale individuals were taken in June and August. Both monthly and annual fin whale lengths were below the estimated length at sexual maturity (but above legal limit), implying a geographic population structure in which immature whales were found more inshore 383. The annual proportion of pregnant females killed decreased between 1948 and 1967384.

From Nichol and Ford (2011 Figure 9): Distribution of geo-referenced Fin Whale catch. Naden Harbour (green symbols) operated 1911-1941 geo-referenced catch 1924-1928, Rose Harbour (yellow symbols) operated 1910-1943 georeferenced catch 1924-1928, Coal Harbour (red symbols) operated 1948-1967, all catch geo-referenced, Kyuquot (black symbols) operated 1907-1925 geo-referenced catch 1924-1928, Sechart (orange symbol) operated 1905-1917 no catch geo-referenced (BC Historical Whaling database DFO; Nichol et al.2002).

                                                                                                               381 Gregr et al. 2000 382 Gregr et al. 2000 383 Gregr et al. 2000 384 Gregr et al. 2000

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From Gregr et al. 2006:

385

                                                                                                               385 Gregr et al. 2000

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Current Status Generally, fin Whales are considered moderately abundant but depleted in the North Atlantic and North Pacific386. Other Regions It is unknown whether they are recovering in Antarctica – the only current population estimate is 15,000387. The central North Atlantic contains an estimated 25,800 fin whales388, the northeastern North Atlantic has about 4,100389, the Spain-Portugal-British Isles area contains around 17,400390, the western Mediterranean basin contains about 3,500391, and in northeastern North American waters (incl. Gulf of St. Lawrence) there are about 2,800392. A minimum of 149 individually identified fin whales have been found in the Gulf of California393. North Pacific Before whaling, there were 42,000-45,000 fin whales in the North Pacific394. By 1973, the North Pacific population was reduced to 13,620-18,680395, of which 8,520 –10,960 belonged to eastern Pacific stock. In 1985 there were an estimated 20,000396, though this has since been revised downward. At their most depleted, North Pacific fin whales were at less than 38% of historic carrying capacity397. In the northeastern Pacific, there are currently about 14,000398 fin whales which have an estimated total biomass of 88,473 tons399. 5,700 of these whales are in the Bering Sea, Aleutian Islands, and Gulf of Alaska400. As of 2002, the NEP fin whale status was uncertain401 and no growth rate estimates were available402. There was some indication that fin whales were increasing in California coastal waters in the 1990’s403, but NOAA line transect surveys in the early 2000s did not suggest this404. However, Bayesian analyses of these data do in fact suggest that California fin whales have been increasing405 at a rate of 4.8% per year which is similar to growth rate estimates in Alaskan waters406. The current best estimate of fin whale abundance in California, Oregon and Washington waters is 3,044-3,300 whales407. The absolute minimum estimate is 2,624. Their Potential Biological Removal (PBR) is 16408. British Columbia The present population in BC waters is inferred to be less than 50% of its level 60-90 years ago409. As of 2006, no current abundance estimates or trends were known for fin whales in western Canadian waters. Researchers recognized an “urgent need for information on their abundance and distribution, their habitat, and the threats they face”410. Williams and Thomas (2007) released the first population estimate for the coast, which was based on distance sampling transects from the Strait of Juan de Fuca to the Alaskan border: 496, with a 95%

                                                                                                               386 Reeves et al. 2002. 387 Aguilar 2008 388 Aguilar 2008 389 Aguilar 2008 390 Aguilar 2008 391 Notobartolo-di-Sciara et al. 2003. 392 Aguilar 2008 393 Tershy et al. 1990 394 Ohsumi and Wada 1974 395 Ohsumi and Wada 1974 396 Allen 1980 397 Mizroch et al 1984 398 Creswell et al. 2007. 399 Barlow et al. 2008. 400 Aguilar 2008 401 Reeves et al. 2002. 402 Best 1993 403 Barlow 1994, 1997. 404 Carretta et al. 2011 405 Moore and Barlow 2012. 406 Zerbini et al. 2006. 407 Aguilar 2008. 408 Carretta et al. 2011. 409 Gregr et al. 2006. 410 Gregr et al. 2006.

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confidence interval of 201-1220. – a very high degree of uncertainty411. The authors divided the intracoastal zone into primary sampling units (PSUs), from which they randomly selected a few for transect coverage. The Kitimat Fjord System was one of these PSUs selected, but the 2004-2005 surveys did not detect any fin whales in the inland waters412. Fin whale sightings from Williams and Thomas (2007):

Threats The most imminent threats in Pacific Canadian waters are collisions with vessels, noise from industrial and military activities, and habitat displacement resulting from shifts in the physical and biological structure of the ocean413. Acoustic: Increasing levels of anthropogenic sound is a habitat concern for many large whale species414. Whales are known to move away from airgun arrays415. Vessel noise, low-frequency active sonar416, and other anthropogenic sound sources may induce behavioral responses of unknown severity417. Researchers have detected modifications in whale acoustic communication to compensate for increased background noise and that a sensitization process may play a role in observed temporary displacement418. In high noise conditions the fin whale’s 20-Hz note duration has been observed to shorten, decrease bandwidth, decrease centre frequency, and decrease peak frequency419. Ship Strike: Fin whales continue to be at risk from ship strikes420. From 1972 to 2001, 46 of 287 fin whale carcasses found (16%) were certainly killed by boats421. In US waters before 2003, 75 out of 292 ship strikes were fin whales422, suggesting that the most-whaled whale species is now the most-struck. Ship strikes were implicated in the deaths of four fin whales and the injury of another from 2004-2008423. The average observed

                                                                                                               411 Williams and Thomas 2007 412 Williams and Thomas 2007. 413 Gregr et al. 2006. 414 Croll et al. 2002, Payne 2004. 415 Castellote et al. 2012. 416 Clark and Altman 2006. 417 Southall et al. 2012. 418 Castellote et al. 2012. 419 Castellote et al. 2012. 420 Gregr et al. 2006, Panigada et al. 2006. 421 Panigada et al. 2006. 422 Jensen and Silber 2003. 423 Carretta et al. 2011.

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mortality/injury due to ship strikes is 1 per year424 though additional mortality probably goes unreported425. Despite these risks, it is uncertain whether the current frequency of strikes could significantly affect population 426 In BC, there have been 8 accounts of fatal fin whale ship strikes between 1999 and 2006427. The highest relative risk of ship strike occurs in bottlenecks, regions where both whale and boat densities are concentrated428. Williams and O’Hara (2010) modeled ship strike risks for BC fin whales based on the only systematic abundance survey available (Williams and Thomas (2007), which found no fin whales in the mainland’s confined channels). Because those abundance data come from the years before fin whales had re-entered the study area, no ship-strike risk was noted for the proposed Northern Gateway and LNG tanker routes429. The impact assessments cited this area as low risk despite the NCCS intervener report demonstrating fin whale presence in the Confined Channel Assessment Area430. Whaling: There is no current risk of fin whaling in BC waters today unless international policies substantially change. Illegal and scientific whaling is still a concern near Iceland, Antarctica, and West Greenland431. Entanglement: Fin whales can incidentally entangle in fishing gear, but this is not thought to be common432. The offshore gillnet fishery is the only likely taker of FWs433, but fishermen report that large whales usually swim through nets without entangling and with very little damage to the nets434. One fin whale death has been observed since 1990, when NMFS began observing the fishery435. Habitat & Trophic Displacement: Degrading habitat conditions due to vessel noise and fishery intensity may displace fin whales from their historical foraging grounds. Decadal oceanographic oscillations and anthropogenic climate change may also bring about shifts in suitable fin whale habitat436. It is possible but unlikely that the remaining populations may be too small to recover437. There is the risk that large baleen whales may have been “replaced” in the ecosystem to some extent by ecologically-equivalent finfish stocks438. On the upside, due to their low trophic level and oceanic distribution, overfishing is not of great concern439. Predation: Increased abundance could lead to increased predation440. Pollutants: Overall contaminant levels in fin whales are extremely low compared to other marine mammals441. Behavioral Response Studies: Mediterranean fin whales responded to disturbance from biopsy sampling efforts from small inflatable craft by moving away and altering respiration patterns.442 The behavioral response of fin whales to navy sonar is currently being written up by Cascadia Research Collective (Ann Allen, pers. comm.)

                                                                                                               424 Carretta et al. 2011. 425 Carretta et al. 2011. 426 Aguilar 2008. 427 Williams and O’Hara 2010. 428 Williams and O’Hara 2010. 429 Williams and O’Hara 2010. 430 Pilkington et al. 2011. 431 Aguilar 2008 432 Aguilar 2008 433 Carretta et al. 2011. 434 Carretta et al. 2011. 435 Carretta et al. 2011. 436 Calambokidis et al. 2009. 437 Gregr et al. 2006. 438 Payne et al. 1990 439 Aguilar 2008 440 Gregr et al. 2006. 441 Aguilar 2008, O’Shea and Brownell 1994, Law 1996 442 Jahoda et al. 2003.

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Protection Global As a species, fin whales received protected status by the IWC in 1976443,444,445. The IUCN (International Union for Conservation of Nature) lists the species as “Endangered”446. The Convention on the International Trade in Endangered Species of Wild Fauna and Flora (CITES) lists fin whales under Appendix 1 (species threatened with extinction, in which international trade is prohibited)447. USA The fin whale was formally listed as endangered in the Endangered Species Act. As such, the west coast stock was automatically listed as “depleted” and “strategic” under the MMPA448. Recently, however, the endangered status for some Northern Hemisphere populations has recently been challenged449. Canada The fin whale is currently listed as Threatened under both the Canadian Species At Risk Act (SARA)450 and the Committee on the Status of Endangered Wildlife in Canada (COSEWIC)451. Both Atlantic and Pacific stocks were originally considered a single unit and were designated of Special Concern in April 1987. This single unit was subsequently split into two populations (Atlantic and Pacific) under SARA in May 2005. The Pacific stock was designated as Threatened in May 2005452. SARA prohibits harm (killing, harassing, capture, or take) to listed species, includes provisions to protect critical habitat, and requires the development of a recovery strategy for each listed species453. No licenses for the taking of cetaceans have been issued for Pacific Canadian waters since 1967454. Any application for a Scientific License for invasive or disturbance-based sampling would require a rigorous assessment based on Section 73 of the SARA455. BC Protected Areas Currently no marine areas are designated to protect the habitat of blue, fin, or sei whales specifically. Under the Canada National Marine Conservation Areas Act, Parks Canada is responsible for the creation of National Marine Conservation Areas (NMCAs). As of 2006, the only NMCA under consideration was south of Haida Gwaii. The Fisheries Act has provisions to protect marine mammal habitat. MPAs may also be established under the Oceans Act. Once critical habitat is identified, approaches for its protection under the provisions of the SARA will be more easily determined456. The Pacific North Coast Integrated Management Area (PNCIMA), a collaborative effort of Department of Fisheries and Oceans and nongovernmental organizations, cited the mainland’s confine channels (the Bangarang study area) as an “Important Area” for fin whales, but the federal government withdrew funding from this initiative in September 2011 after pressure from the pipeline lobby and the PNCIMA initiative was abandoned457,458. Critical Habitat Designation

                                                                                                               443 Carretta et al. 2011. 444 Carretta et al. 2011. 445 Gregr et al. 2006. 446 Gregr et al. 2006, Klinowska 1991. 447 Gregr et al. 2006. 448 Carretta et al. 2011. 449 Reeves et al. 2002. 450 Nichol and Ford 2011. 451 Gregr et al. 2006. 452 Gregr et al. 2006. 453 Gregr et al. 2006. 454 Gregr et al. 2006. 455 Gregr et al. 2006. 456 Gregr et al. 2006. 457 http://www.davidsuzuki.org/media/news/downloads/2011/PNCIMAFundingBackgrounder.pdf 458 http://www.pncima.org/

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Designation under SARA requires that critical habitat in Canada and threats to that habitat be identified for the threatened species. Critical habitat has not yet been designated for the fin whale, though DFO is actively conducting the research required to do so (Nichol and Ford 2011). The recovery plan for fin whales (and other large balaenopterids) lists the following strategic steps459:

1. Identify critical habitat a. “Basic abundance and distribution data is required in order to identify critical habitat.”460

2. Rigorously determine species abundance and distribution 3. Outline means of mitigating threats

As with all highly mobile animals, “the recovery of these populations is unlikely to be accomplished by Canadian efforts alone. The need for multi-lateral and international cooperation is therefore considered essential to the successful recovery of these species.”461

Ongoing BC Research Coast-wide In an effort to identify critical habitat for BC’s largest baleen whales, Department of Fisheries and Oceans’ (DFO) Cetacean Research Program (CRP) has been conducting ship-based surveys since 2002 which have totaled 2,000 hours of effort in all seasons and covered over 40,000km of ocean462. The fin whale was the third-most encountered whale species during these surveys463, with an encounter rate of 1.161 individuals per 100km464. An extensive acoustic monitoring network has been installed on the BC coast as well.

465 From Nichol and Ford (2011 Figure 7): Fin Whales sightings, DFO surveys, 2002 to 2010 (Ford et al. 2010a; CRP-DFO unpubl. data).

                                                                                                               459 Gregr et al. 2006. 460 Gregr et al. 2006. 461 Gregr et al. 2006. 462 Nichol and Ford (2011), Ford et al. 2010a 463 Nichol and Ford (2011). 464 Nichol and Ford (2011). 465 Nichol and Ford (2011).

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Identification photos from these surveys are being prepared for mark-recapture analysis to generate a regional BC abundance estimate466. To determine broader movement patterns and Northeast Pacific stock structures, DFO is also planning to compare fin whale photos from these surveys to photo-ID catalogs from Washington, California and Alaska.467 Nichol and Ford (2011) summarize this decade of effort. “BC waters support foraging whales of these species where habitat features contribute to concentrating and sustaining prey, although the occurrence of fin whales year-round and the detection of male mating calls underscore the need to better understand the function of BC habitat for Fin Whales.” In order to complete the critical habitat identification process for fin whales, DFO reported in 2011 that the following research efforts are urgently needed468:

1. Expanded acoustic monitoring, both offshore and inshore, with a special emphasis on identifying the behavioral context of fin whale vocalization.

2. Documenting the fine-scale habitat features with which fin whales and their prey are associated “particularly in Hecate Strait and Caamano Sound. Study approaches should include systematic visual surveys, photo-identification, satellite tagging and collection of hydroaoustic data to document potential prey.”

3. Systematic survey coverage in key inshore areas such as Caamano Sound. “The occurrence of fin whales in Caamano Sound and adjoining waterways on the northern mainland coast is intriguing as it is one of few inshore areas where fin whales are found in BC and where they were encountered historically as well.”

4. Linking whale distribution to oceanographic processes. “Concurrent data on whale distribution and oceanographic processes at appropriate spatial and temporal scales are lacking.”

Kitimat Fjord System The confined channels near Caamano Sound, to which Nichol and Ford (2011) refer so regularly above, have been monitored visually and acoustically for whales by the North Coast Cetacean Society (NCCS) for the past decade and is now the centerpiece of the Bangarang study area as well. Between 1968 and 2006 (38 years), there were only a few sightings of fin whales in the confined channels by DFO or reported on the BC Cetacean Sighting Network (BCCSN): one in Beauchemin Channel (August 1994), one in Squally Channel (July 1992), and one in Campania Sound (2005)469. Almost all fin whale sightings were from the continental shelf off Vancouver Island, southern Haida Gwaii, and western Dixon Entrance470. Then, in 2006, NCCS had 5 fin whale sightings over 3 days471. In the seasons since, the number of sightings has increased dramatically. Between 2006 and 2011, NCCS had 116 days with fin whales, with 28 individuals identified472. These developments may represent the re-occupation of a historic inland fin whale feeding ground473.

                                                                                                               466 Nichol and Ford (2011). 467 Nichol and Ford (2011). 468 Nichol and Ford (2011). 469 JKB Ford, pers. comm. 470 Nichol and Ford (2011). 471 Pilkington et al. 2011. 472 Pilkington et al. 2011. 473 Gregr et al. 2006.

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Literature Cited

Agler, A.B., Beard, J.A., Bowman, S.R., Corbett, H.D., Frohock, S.E., Hawvermale, M.P.,! Katona, S.K., Sadove, S.S., and Seipt, I.E. (1990). Fin Whale (Balaenoptera ! physalus) Photographic Identification: Methodology and Preliminary Results from ! the Western North Atlantic. Report to the International Whaling Commision: ! Special Issue 12: 349-346.

Aguilar, Alex. 2009. Fin Whale: Balaenoptera physalus. In Encyclopedia of Marine Mammals, eds. William F. Perrin, Bernd Würsig and J.G.M. Thewissen. Academic Press and Elsevier.

Allen 1980 – fin whale population estimates

Anthony, R. & Calvet, J. (1905) Recherches faites sur le cétacé capturé a Cètte, le 6 octobre 1904 – Balaenoptera Physalus (Linné). Bulletin de la Société Philomatique.

Archer, FI, PA Morin, BL Hancock-Hanser, KM Robertson, MS Leslie, M Berube, S Panigada, BL Taylor. 2013. Mitogenomic phylogenetics of fin whales (Balaenoptera physalus spp.): Genetic evidence for revision of subspecies. PLOS ONE 8(5):e63396.

Barlow, J. 1994. Abundance of large whales in California coastal waters: a comparison of ship surveys in 1979/80 and in 1991. Rept. Int. Whal. Commn. 44:399-406.

Barlow, J. 1997. Preliminary estimates of cetacean abundance off California, Oregon, and Washington based on a 1996 ship survey and comparisons of passing and closing modes. Admin. Rept. LJ-97-11 available from Southwest Fisheries Science Center, P.O. Box 271, La Jolla, CA. 25 pp.

Barlow, J., M. Kahru, B.G. Mitchell. 2008. Cetacean biomass, prey consumption, and primary production requirements in the California Current ecosystem. Marine Ecology Progress Series 371:285-295.

Berube (and 9 others). 1998a. Population genetic structure of North Atlantic, Mediterranean Sea and Sea of Cortez fin whales, Balaenoptera physalus (Linnaeus 1758): Analysis of mitochondrial and nuclear loci. Mol. Ecol. 7:585-599.

Berube M, A Aguilar, D Dendanto, F Larsen, GN DiSciara, et al. 1998b. Population genetic structure of North Atlantic, Mediterranean Sea and Sea of Cortez fin whales, Balaenoptera physalus (Linnaeus 1758): analysis of mitochondrial and nuclear loci. Mol Ecol 7:585-599.

Berube M, J Urban, AE Dizon, RL Brownell, PJ Palsboll. 2002. Genetic identification of a small and highly isolated population of fin whales (Balaenoptera physalus) in the Sea of Cortez, Mexico. Conservation Genetics 3:183-190.

Berube, M., and Aguilar, A. 1998. A new hybride between a blue whale, Balaenoptera musculus, and a fin whale, B. physalus: Frequency and implications of hybridization. Mar. Mamm. Sci. 14(1): 82-98.

Best, P. B. 1993. Increase rates in severely depleted stocks of baleen whales. ICES J. Mar. Sci. 50:169-186.

Brueggeman, J. J., G. A. Green, K. C. Balcomb, C. E. Bowlby, R. A. Grotefendt, K. T. Briggs, M. L. Bonnell, R. G. Ford, D. H. Varoujean, D. Heinemann, and D. G. Chapman. 1990. Oregon-Washington Marine Mammal and Seabird Survey: Information synthesis and hypothesis formulation. U.S. Department of the Interior, OCS Study MMS 89-0030.

Calambokidis, J., JP Barlow, JKB Ford, TE Chandler, AB Douglas. 2009. Insights into the population structure of blue whales in the Eastern North Pacific from recent sightings and photographic identification. Marine Mammal Science 25(4):816-832.

Carretta, JV, KA Forney, E Oleson, K Martien, MM Muto, MS Lowry, J Barlow, J Baker, B Hanson, D Lynch, L Carswell, RL Brownell Jr, J Robbins, DK Mattila, K Ralls, MC Hill. 2011. US Pacific Marine Mammal Stock Assessments. NOAA-TIM-NMFS-SWFSC-488, US Department of Commerce.

Castellote M, CW Clark, MO Lammers. 2011. Fin whale (Balaenoptera physalus) population identify in the western Mediterranean Sea. Marine Mammal Science 28:325-344.

35

Castellote, Manuel, C.W. Clark, M.O. Lammers. 2012. Acoustic and behavioural changes by fin whale (Balaenoptera physalus) in response to shipping and airgun noise. Biological conservation.

Clapham, P. J., S. Leatherwood, I. Szczepaniak, and R. L. Brownell, Jr. 1997. Catches of humpback and other whales from shore stations at Moss Landing and Trinidad, California, 1919-1926. Mar. Mamm. Sci. 13(3):368-394.

Clapham, P.J. (2000) The humpback whale: seasonal feeling and breeding in a baleen whale, In: Cetacean Societies: Field Studies of Dolphins and Whales (Ed. by J. Mann, R.C. Connor, P.L. Tyack & H. Whitehead), pp. 173–196. University of Chicago Press, Chicago and London.

Clark, C. W. (1993). "Bioacousticso f baleenw hales:F rom infrasonicsto complexs ongs,"J . Acoust.S oc.A m. 94, 1830,A bstract.

Clark, C.W., and Altman, N.S. 2006. Acoustic detections of blue whale (Balaenoptera musculus) and fin whale (B. physalus) sounds during SURTASS LFA exercise. IEEE J. Oceanic Eng. 31(1): 120-128.

Clark, CW, JF Borsani, G Notobartolo-di-Sciara. 2002. Vocal activity of fin whales, Balaenoptera physalus, in the Ligurian Sea. Mar. Mamm. Sci. 18:286-295.

Committee on Taxonomy. 2012. List of marine mammal species and subspecies. Society for Marine Mammalogy.

Connor, R.C. (2000) Group living in whales and dolphins, In: Cetacean Societies: Field Studies of Dolphins and Whales (Ed. by J. Mann, R.C. Connor, P.L. Tyack & H. Whitehead), pp. 199–218. University of Chicago Press, Chicago and London.

COSEWIC 2005. COSEWIC assessment and update status report on the fin whale Balaenoptera physalus in Canada. Committee on the Status of Endangered Wildlife in Canada. Ottawa. ix + 37 pp. http://publications.gc.ca/collections/Collection/CW69-14-428-2005E.pdf

Creswell, Graeme, Dylan Walker and Todd Prusser. 2007. Whales and dolphins of the North American Pacific. Harbour Publishing: China.

Croll, D., CW Clark, A Acevedo, B Tershy, S flores, J Gedamke, J urban. 2002. Only male fin whales sing loud songs. Nature 117 809

Croll, D.A., C.W. Clark, A. Acevedo, B. Tershy, S. Flores, J. Gedamke, and J. Urban. 2002. Only male fin whales sing loud songs. Nature 417:809

Croll, DA, A. Acevedo-Gutierrez, B.R. Tershy, J. Urban-Ramirez. 2001. The diving behavior of blue and fin whales: is dive duration shorter than expected based on oxygen stores? Comparative Biochemistry and Physiology Part A 129:797-809.

Dalla Rosa, L. 2010. Modeling the foraging habitat of humpback whales. Ph.D. Thesis, University of British Columbia. pp171.

Dohl, T. P., R. C. Guess, M. L. Duman, and R. C. Helm. 1983. Cetaceans of central and northern California, 1980-83: Status, abundance, and distribution. Final Report to the Minerals Management Service, Contract No. 14-12-0001-29090. 284p.

Doniol-Valcroze, T, D. Berteaux, P. Larouche, and Sears, R. 2007. Influence of thermal fronts on habitat selection by four rorqual whale species in the Gulf of St. Lawrence. Mar. Ecol. Prog. Ser. 335: 207–216

Donovan, G. P. 1991. A review of IWC stock boundaries. Rept. Int. Whal. Commn., Special Issue 13:39-68.

Edds-Walton, PL. 1997. Acoustic communication signals of mysticete whales. Bioacoustics 8(1&2):47-60.

Edds, PL. 1988. Characteristics of finback Balaenoptera physalus vocalizations in the St. Lawrence River. Journal of Mammalogy 63:346-347.

Eder, Tamara. 2001. Whales and other marine mammals of British Columbia and Alaska. Lone Pine Publishing: Canada.

36

Falcone, E., Diehl, B., Douglas, A. and J. Calambokidis. 2011. Photo-identiication of fin whales (Balaeanoptera physalus) along the US west coast, Baja California, and Canada. CAscadia Research Collective, Olympia, WA. Final report for order# JF13F09SE516,20pp.

Flinn, R.D., AW Trites, EJ Gregr,RI Perry. 2002. Diets of fin, sei, and sperm whales in British Columbia: An analysis of commercial whaling records, 1963-1967. Marine Mammal Science 18(3):663-679.

FORCADA, J., G. NOTARBARTOLO DI SCIARA AND F. FABBRI. 1995. Abundance of fin whales and striped dolphins summering in the Corso-Ligurian Basin. Mammalia 59:127–140.

Ford, J.K.B., Abernethy, R.M., Phillips, A.V., Calambokidis, J., Ellis, G.M. and. Nichol, L.M. 2010a. Distribution and relative abundance of cetaceans in western Canadian waters from ship surveys, 2002-2008. Can. Tech. Rep. Fish. Aquat. Sci. 2913: v + 51 p.

Ford, J.K.B., Koot, B., Vagle, S., Hall-Patch, N., and Kamitakahara, G. 2010b. Passive acoustic monitoring of large whales in offshore waters of British Columbia. Can. Tech. Rep. Fish. Aquat. Sci. 2898: v + 30 p.

Forney, K. A., J. Barlow, and J. V. Carretta. 1995. The abundance of cetaceans in California waters. Part II: Aerial surveys in winter and spring of 1991 and 1992. Fish. Bull. 93:15-26.

FUJINO, K. 1964. Immunogenetic and marking approaches to identifying subpopulations of the North Pacific whales. The Scientific Reports of the Whales Research Institute, Tokyo 15:85-141.

Gambell, Ray. 1985. Fin whale – Balaenoptera physalus. In Handbook of Marine Mammals, Volume 3 The Sirenians and Baleen Whales. Ed. Sam H. Ridgway and Sir Richard Harrison, F.R.S. Academic Press: London.

Geisler JH, MR McGowen, Guang Yang, J Gatesy. 2011. A supermatrix analysis of genomic, morphological, and paleontological data from crown Cetacea. BMC Evolutionary Biology 11:112.

Green, G. A., J. J. Brueggeman, R. A. Grotefendt, C. E. Bowlby, M. L. Bonnell, K. C. Balcomb, III. 1992.Cetacean distribution and abundance off Oregon and Washington, 1989-1990. Ch. 1 In: J. J.Brueggeman (ed.). Oregon and Washington Marine Mammal and Seabird Surveys. Minerals Management Service Contract Report 14-12-0001-30426.

Gregr, E. J. 2002. Whales in Northern BC: Past and Present. Pages 74-78 in T. Pitcher, M. Vasconcellos, S. Heymans, C. Brignall, and N. Haggan, editors. Information Supporting Past and Present Ecosystem Models of Northern British Columbia and the Newfoundland Shelf. Fisheries Centre, University of British Columbia,Vancouver.

Gregr, E. J. 2004. Marine mammals in the Hecate Strait ecosystem. Canadian Technical Report of the Fisheries and Aquatic Sciences 2503:56p.

Gregr, E. J., and A. W. Trites. 2001. Predictions of critical habitat for five whale species in the waters of coastal British Columbia. Canadian Journal of Fisheries and Aquatic Science 58:1265-1285.

Gregr, E. J., L. Nichol, J. K. B. Ford, G. Ellis, and A. W. Trites. 2000. Migration and population structure of northeastern Pacific whales off coastal British Columbia: An analysis of commercial whaling records from 1908-1967. Marine Mammal Science 16(4):699-727.

Gregr, E.J., J. Calambokidis, L. Convey, J.K.B. Ford, R.I. Perry, L. Spaven, M. Zacharias. 2006. Recovery Strategy for Blue, Fin, and Sei Whales (Balaenoptera musculus, B. physalus, and B. borealis) in Pacific Canadian Waters. In Species at Risk Act Recovery Strategy Series. Vancouver: Fisheries and Oceans Canada. vii + 53 pp.

Gregr, EJ, L Nichol, JKB Ford, G Ellis, AW Trites. 2000. Migration and population structure of northeastern Pacific whales off coastal British Columbia: an analysis of commercial whaling records from 1908 – 1967.

Hain, J. H. W., M. J. Ratnaswamy, R. D. Kenney, and H. E. Winn. 1992. The fin whale, Balaenoptera physalus, in waters of the northeastern United States continental shelf. Report of the International Whaling Commission 42:653-669.

Hamilton TA, JV Redfern, J Barlow, LT Ballance, T Gerrodette, et al. 2009. Atlas of Cetacean Sightings for Southwest Fisheries Science Center cetaceans and ecosystem survey, 1986-2005. La Holla, CA: US Dept

37

of Commerce, National Oceanic and Atmospheric Administration, National Marine Fisheries Service, Southwest Fisheries Science Center. 70p.

Hatch, LT, Clark CW. 2008. Acoustic differentiation between fin whales in both the North Atlantic and North Pacific Oceans, and integration with genetic estimates of divergence. International Whaling Commission Scientific Meeting.

Ivashin,MV., Popov, LA, and Tsapko, AS. 1972. Morskie Mlekopptayschie. Pischevaya Promychiennost, Moscow.

Jahoda, M., C.L. LaFortuna, N Biassoni, C Almirante, A. Azzellino, S Panigada, M Zanardelli, G. Notarbartolo-di-Sciara. 2003. Mediterranean fin whale’s (Balaenoptera physalus) response to small vessels and biopsy sampling assessed through passive tracking and timing of respiration. Marine Mammal Science 19(1):96-110.

Jensen, A.S. and G.K. Silber. (2003). Large Whale Ship Strike Database. U.S.! Department of Commerce, NOAA Technical Memorandum. NMFS-OPR- , 37 pp.

Johnston, D.W., L.H. Thorne, A.J. Read. 2005. Fin whales (Balaenoptera physalus and minke whales Balaenoptera acutorostra exploit a tidally driven island wake ecosystem in the Bay of Fundy. Marine Ecology Progress Series 305:287-295.

Jongsgard 1962

Kawamura, A. 1980. A review of food of balaenopterid whales. Sci. Rep. Whales. Res. Inst. Tokyo, 32 155-197.

Kawamura, A. 1982. Food habits and prey distributions of three rorqual species in the North Pacific Ocean. Scientific Report of the Whales Research Institute 3$:59-91.

KAWAMURA, A., 1982. Food habits and prey distributions of three rorqual species in the North Pacific Ocean. Scientific Reports of the Whales Research Institute, Tokyo 34:59-91.

Kiegwin LDJ. 1978. Pliocene closing of the Isthmus of Panama, based on biostratigraphic evidence from nearby Pacific Ocean and Caribbean Sea cores. Geology 6:630-634.

Klinowska, M. 1991. “Dolphins, Porpoises and Whales of the World.” The IUCN Red Data Book. IUCN, Gland, Switzerland, 429 pp.

Lafortuna, CL, M Jahoda, A Azzellino, F Saibene, a. Colombini. 2003. Locomotor behaviours and respiratory pattern of the Mediterranean fin whale (Balaenoptera physalus). Eur J Appl Physiol 90: 387-395.

Law, R.J. 1996. Metals in marine mammals. In “Environmental contaminants in wildlife: interpreting tissue concentrations.” (WN Beyer, GH Heinz, and AW Redmon-Norwood, eds), pp. 357-376. CRC Press In, Boca Raton, USA.

Leatherwood, Stephen, and Randall R. Reeves. 1983. The Sierra Club Handbook of Whales and Dolphins. Tien Wah Press: Singapore.

Lockyer and Waters 1986

Lockyer, C. 1976. Body weights of some species of large whales. J. Cons. Cons. Int. Explor. Mer 36: 259-273.

Mackintosh, Na, and JFG Wheeler. 1929. Southern blue and fin whales. Disc. Rep. 1:257-540.

Mackintosh, NA. 1965. The stocks of whales. Fishing News, London.

Marini, L., Consiglio, C., Catalano, B., Valentini, T. & Villetti, G. 1996. Aerial behavior of fin whales in the Mediterranean Sea. Marine Mammal Science, 12, 489–495.

McDonald, M. A., J. A. Hildebrand, and S. C. Webb. 1994. Blue and fin whales observed on a seafloor array in the Northeast Pacific. (unpubl. ms.).

McDonald, M.A., and Fox, C.G. 1999. Passive acoustic methods applied to fin whale population density estimation. J. Acoust. Soc. Am. 105(5): 2643-2651.

McDonald, M.A., J.A. Hildebrand and S.C. Webb. 1995. Blue and fin whales observed on a seafloor array in the Northeast Pacific. J. Acoust. Soc. Am. 98(2);712-721.

38

McGowen, MR, M Spaulding, J Gatesy. 2009. Divergence date estimation and a comprehensive molecular tree of extant cetaceans. Molecular phylogenetics and evolution 53:891-906.

Mizroch, S. A., D. W. Rice, and J. M. Breiwick. 1984. The fin whale, Balaenoptera physalus. Mar. Fish. Rev. 46:20-24.

Mizroch, S.A., Rice, D.W., Zwiefelhofer, D., Waite J. and Perryman, W.L. 2009. Distribution and movements of fin whales in the North Pacific Ocean. Mammal. Rev. 39(3): 193-227

Moore, JE, J Barlow. 2012. Bayesian state-space model of fin whale abundance trends from a 1991-2008 time series of line-transect surveys in the California Current. Journal of Applied Ecology 48(5)

Moore, S. E., K. M. Stafford, M. E. Dahlheim, C. G. Fox, H. W. Braham, J. J. Polovina, and D. E. Bain.1998. Seasonal variation in reception of fin whale calls at five geographic areas in the North Pacific. Mar. Mamm. Sci. 14(3):617-627.

Moore, SE, KM Stafford, DK Mellinger, JA Hildebrand. 2006. Listening for large whales in the offshore waters of Alaska. BioScience 56:1.

Nemoto, T. 1957. Foods of baleen whales in the northern Pacific. Scientific report of the Whales Research Institute 12:33-89.

Nemoto, T. 1959. Food of baleen whales with reference to whale movements. Scientific Report of the Whales Research Institute 14:149-290.

Nemoto,T. and T. Kasuya. 1965. Foods of baleen whales in the Gulf of Alaska of the North Pacific. Scientific Report of the Whales Research Institute 19:45-51.

Nichol, L. and K. Heise. 1992. The historical occurrence of large whales off the Queen Charlotte Islands. Prep. for South Moresby/Gwaii Haanas National Parks Reserve, Canadian Parks Service, Queen Charlotte City, B.C. 68 pp.

Nichol, L. M., E. J. Gregr, R. Flinn, J.K.B. Ford, R. Gurney, L. Michaluk, and A. Peacock. 2002. British Columbia Commercial Whaling Catch Data 1908 to 1967: A Detailed Description of the B.C. Historical Whaling Database. Can. Tech. Rep. Fish. Aquat. Sci. 2396: viii + 76 p.

Nichol, LM and JKB Ford. 2011. Information relevant to the assessment of critical habitat for Blue, Fin, Sei and North Pacific Right Whales in British Columbia. Canadian Science Advisory Secretariat. Research Document 2011/137.

NICHOLL, ., AND K. HEISE1. 992. The historical occurrence of large whales off the Queen Charlotte Islands. Prepared For South MoresbyiGwaii Haanas National Parks Reserve, Canadian Parks Service, Queen Charlotte City, BC. 68 pp.

Nishiwaki 1950

Notarbartolo-di-Sciara, G., M Zanardelli, M Jahoda, S Panigada, S Airoldi. 2003. The fin whale Balaenoptera physalus (L. 1758) in the Mediterranean Sea. Mammal Review 33:105-150.

O’Shea, T.J., and Brownell,R.L. 1994. Organochlorine and metal contaminants in baleen whales: a review and evaluation of conservation implications. Sci. Total Environ. 154:179-200.

Ohsumi, S. and S. Wada. 1974. Status of whale stocks in the North Pacific, 1972. Rept. Int. Whal. Commn. 25:114-126.

Orsi Relini, L., Relini, G., Cima, C., Fiorentino, F., Palandri, G., Relini, M. & Torchia, G. (1992) Una zona di tutela biologica ed un parco pelagico per i cetacei del Mar Ligure. Bollettino dei Musei e degli Istituti Biologici dell’Università di Genova, 56, 57, 247–281.

Panigada, S, G Pesante, M Zanardelli, F Capoulade, A gannier, MT Weinrich. 2006. Mediterranean fin whales at risk from fatal ship strikes. Marine Pollution Bulleting 52: 1287-1298.

Panigada, S., G. Pesante, M. Zanardelli and S. Oehen. 2003. Day and night-time behaviour of fin whales in the Western Ligurian Sea. Proceedings of the Conference Oceans 2003, September 22-26, 2003, San Diego, California. Pp 466-471.

39

Panigada, S., Zanardelli, M., Canese, S. & Jahoda, M. (1999) How deep can baleen whales dive? Marine Ecology Progress Series, 187, 309–311.

Payne, M. P., D. N. Wiley, S. B. Young, S. Pittman, P. J. Clapham, and J. W. Jossi. 1990. Recent fluctuations in the abundance of baleen whales in the sourthern Gulf of Maine in relation to changes in selected prey. Fisheries Bulletin 88(4):687-696.

Payne, R. S. 2004. Long-range communication in large whales, ocean noise and synergistic impacts. International Whaling Commission. 22-23

Payne, R., D Webb. 1971. Orientation by means of long range acoustic signaling in baleen whales. Annals New York Academy of Sciences.

Payne, RS. 2004. Long-range communication in large whales, ocean noise and synergistic impacts. IWC-SC Report: IWC56 Annex K Appendix 2.

Peers and Karlsson 1976

PIKE, G. 1950. Stomach contents of whales caught off the coast of British Columbia. Fisheries Research Board Canadian Pacific Progress Report 83:27-28

Pike, G. C., and I. B. MacAskie. 1969. Marine Mammals of British Columbia. Fisheries Research Board of Canada Bulletin 171, Ottawa, ON.

Pilkington, J, H Meuter and J Wray. 2011. Occurrence of fin whales (Balaenoptera physalus in the Confined Channel Assessment Area between Wright Sound and Caamano Sound from North Coast Cetacean Society for the Period 2004 to 2011. Written Evidence Submission to the Joint Review Panel Assessing the Northern Gateway Pipeline Application.

Ray, G.C., Mitchell, E.D., Wartzok, D., Kozicki, V.M. & Maiefski, R. (1978) Radio tracking of a fin whale (Balaenoptera physalus). Science, 202, 521–524.

Reeves, Randall R., Brent S. Stewart, Phillip J. Clapham, James A. Powell. 2002. Guide to Marine Mammals of the World. Alfred A. Knopf, Inc. Hong Kong.

Relini, G., Orsi Relini, L., Siccardi, A., Fiorentino, F., Palandri, G., Torchia, G., Relini, M., Cima, C. & Cappello, M. (1994) Distribuzione di Meganyctiphanes norvegica e Balaenoptera physalus in Mar Ligure all’inizio della primavera. Biologia Marina Mediterranea, 1, 89–94.

Rice, D. W. 1974. Whales and whale research in the eastern North Pacific. pp. 170-195 In: W. E. Schevill (ed.). The Whale Problem: A Status Report. Harvard Press, Cambridge, MA.

Richard, J. (1936) Notes sur les cétacés et les pinnipèdes. In: Documents sur les cétacés et pinnipèdes des croisières du Prince Albert I de Monaco (Ed. by J. Richard), pp. 34–61. Résultats des campagnes scientifiques accomplies sur son yacht par Albert I Prince Souverain de Monaco, 94.

Richiardi, S. (1874) Sulle variazioni individuali della Balaenoptera musculus. Atti della Società Toscana di Scienze Naturali, 1, 246–259

Rothernberg, David. 2008. Thousand mile song. Basic Books: New York.

Sergeant, D.E. (1977) Stocks of fin whales, Balaenoptera physalus L. in the North Atlantic Ocean. Report of the International Whaling Commission, 27, 460–473.

Sirovic, A, JA Hildebrand, SM Wiggins. 2007. Blue and fin whale call source levels and propagation rane in ther Southern Ocean. J Acoust Soc Am 122(2): 1208-1215.

Southall, B, D Moretti, B Abraham, J Calambokidis, S Reuter, P Tyack. 2012. Marine mammal behavioral response studies in Southern California: advances in technology and experimental methods. Marine Technology Society Journal 46(4):48-59.

Spalding, David A.E. 1998. Whales of the West Coast. Harbour Publishing, Canada.

Tershy, B. R., D. Breese, and C. S. Strong. 1990. Abundance, seasonal distribution and population composition of balaenopterid whales in the Canal de Ballenas, Gulf of California, Mexico. Rept. Int. Whal. Commn., Special Issue 12:369-375.

40

Tershy, B. R., J. Urbán-R., D. Breese, L. Rojas-B., and L. T. Findley. 1993. Are fin whales resident to the Gulf of California? Rev. Invest. Cient., Univ. Auton. De Baja California Sur. 1:69-71

Thompson, P.O., Findley, L.T., and Vidal, O. 1992. 20 Hz pulses and other vocalizations of fin whales, Balaenoptera physalus, in the Gulf of California Mexico. J. Acoust. Soc. Am. 92: 3051-3057.

Tonnessen, J. N., and A. O. Johnsen. 1982. The History of Modern Whaling. Univ. Calif. Press, Berkeley and Los Angeles. 798pp.

Trites et al. 1997. Competition between fisheries and marine mammals for prey and primary production in the Pacific Ocean. J Northwest Atl Fish Sci 22:173-187.

Trites, A. W., P. A. Livingston, M. C. Vasconcellos, S. Mackinson, A. M. Springer, and D. Pauly. 1999. Ecosystem change and the decline of marine mammals in the Eastern Bering Sea: testing the ecosystem shift and commercial whaling hypotheses. Fisheries Centre Research Reports 7(1):106.

Vidal, O. & Pechter, G. (1989) Behavioral observations on fin whale, Balaenoptera physalus, in the presence of killer whale, Orcinus orca. Fishery Bulletin, 87, 370–373.

Walker, M.M., Kirschvink, JL, Ahmed, G., and Dizon, AE. 1992. Evidence that fin whales respond to the geomagnetic field during migration. J. Exp. Biol. 171, 67-78.

Watkins, W. A. (1981). "Activities and underwater sounds of fin whales," Sci. Rep. Whales Res. Inst. 33, 83-117.

Watkins, W.A., Daher, M., Repucci, G.M., George, J.E., Martin, D.L., DiMarzio, N.A., and Gannon, D.P. 2000. Seasonality and distribution of whale calls in the North Pacific. Oceanography. 13(1): 62-67.

Watkins, W.A., Tyack, P. and Moore, K.E. 1987. The 20-hz signals of finback whales (Balaenoptera physalus). J. Acoust. Soc. Am. 82(6): 1901-12.

Williams, R, and P O’Hara. 2009. Modelling ship strike risk to fin, humpback and killer whales in British Columbia, CA. J. Cetacean Res. Manage. 11(1):1-8.

Williams, R., L Thomas. 2007. Distribution and abundance of marine mammals in the coastal waters of British Columbia, CA. J Cetacean Res. And Management 9(1):15-28.

Wolanski E, Hamner WM (1988) Topographically controlled fronts in the ocean and their biological influence. Science 241:177–181

Woodley, T. H., and D. E. Gaskin. 1996. Environmental characteristics of north Atlantic right and fin whale habitat in the lower Bay of Fundy, Canada. Canadian Journal of Zoology 74(1):75-84.

Zanardelli, M., Notarbartolo di Sciara, G. & Acquarone, M. (1992) Cetacean sightings by amateurs: a twosided coin. European Research on Cetaceans, 6, 79–82.

Zerbini, A.N., Waite, J.M., Laake, J.L. & Wade, P.R. (2006) Abundance, trends and distribution of baleen whales off western Alaska and the central Aleutian Islands. Deep-Sea Research, 53, 1772–1790.