THE LACS DES LOUPS MARINS HARBOUR SEAL,

Phoca vif~linu mellonae Doutt 1942:

ECOLOGY OF AN ISOLATED POPULATION

A Thesis

Presented to

The Facuity of Graduate Studies

of

The University of Guelph

by

IUCHARD JOHN SMITH

In partid fbElment of requirements

for the degree of

Doctor of Philosophy

August, 1999

O Richard John Smith, 1999

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ABSTRACT

THE LACS DES LOUPS MARINS HARBOUR SEAL, Phoca vi'hrina mellonae Doutt 1942:

ECOLOGY OF AN ISOLATED POPULATION

Richard John Smith University of Guelph, 1999

Advisor: Dr. DM. Lavigne

Investigations were undertaken to determine to what extent the Lacs des Loups MarEis

harbour seal (Phoca v i t u l i ~ mellonoe) is distinct and isolated £tom aceanic harbour

seals. The population occurs in the area of Lacs des Loups Marins (Lower Seal Laices)

(56-57W, 73-74OW), 160 km East of Hudson Bay, on the Ungava penmsula of northem

Quekc. Written referencs b the mique appearance and behaviour of this seal date

back to 18 18. The subspecies was describeci primarily on the basis of a characteristic

morphology and presumed long-time geographic isolation fkom neighbouring oceanic

harbour seals.

A craniometric d y s i s of P. v. mellome conhned that it is morphologically

distinguishable fkom oceanic harbour seal subspecies in the Atlantic and Pacific. There

was also evidence ofbehavioural differences: Pupping seerns to take place substantially

earlier (early May) than in other harbour seal populations at similar latitudes. An analysis

of DNA sequences nom the mitochondrial control region indicated that the Lacs des

Loups Marins animds exhibit some haplotypic divergence fkom other harbour seals.

Analyses of stable isotope ratios in hair and fatty acid signatures in blubber indicated that

P. v. mellonae feeds exclusively in hshwater. Monitoring of the movements of eight

fieshwater animals with satellite telemetry over portions of a two year period provided

firrther evidence that not ody are the seals resident within the Lacs des Loups Marins

area, but that individual animals exhi'bit considerabIe site fidelïty. The diffierentiation

exhibited by P. v. meIIome is similar to that demoa~frated by isolated ringed seal

popdations in Lakes Saimaa, Fiidand (Pusa LWpik suimenrii) and Ladoga, Russia (P. h

ladogemis).

The totality of the evidence collected provides support for the designation of the Lacs

des Loups Marins seal as a distinct subspecies. The population warrants c ldcat ion as

an Evolutionarily Signifiant Unit. The infiinnation gatheted on the cange of P. v.

mellonae has implications for friture efforts to conserve the population, which is now

listed as 'tulnerable" by the Cornmittee on the Status of Endangerd Wiidlife in Canada

(COSEW).

For John Petagumskum, Sr.,

who hows more about uchihnzjdi than 1 ever wili.

It was the Mauritius pïnk pigeon that inspired this thesis. Voracious reading of Geraid

Durrell as a kid got me thinking about the unique bond ttiat can develop between a

researcher and their study animal. As I absorbai the tales of Corfi and other fa.-away

places, I figured the natural and necessary complement to becoming the "'world expert"

on the arcane aspects of a littIe hown (tbough wonderfid) creahire's biology is a

responsibility for the consewation of that species. I feel very fortunate that this project

afforded me the opportunity to explore all of these sides of an incrediile animal that

exists - almost unknown - in my own country.

This thesis was a long tune in comùig, and it was only with the support and

encouragement of many people that its completion was possible. I hope that my Mom

and Dad see a lot of their early teachings in this work Xt was their love of the outdoors

and our family came trips and weekends at Lake Chateaugay that first made me redise

that 1 wanted to becorne a forest ranger (or some simila. occupation!). My sister Lori.

my grandparents John and Uajorie Braive, and my late grandmother Joan Smith (who

was responsible for our growing up very much aware of our Newfoundand, sea-centmi,

roots) have always been very supportive of my biological bent.

I'm very grateful to Dave Gaskin for giving me my start in marine mammal biology.

The amazhg diversity and number of people at Dave's mernorial service last year was

testimony to his decency and positive impact on people's lives. His lab in Zoology

1

Annex 1, and the Grand Manan Whale & Seabïrd Research Station, were great places to

leam and to discuss marine mammal issues, and were a meeting place for a dedicated

group of colleagues and Eends that 1 respect tremendousiy: Per Berggren, Dave

Johnston, Heather Koopman, Aieksija Neimanis, Andy Read, Sue Wallace, Andrew

"Sparl@' Westgate and Thom W d e y . 1 had my £kt 'harine mammal eXpenence7'

with Andrew and Thom, travelling back and forth in very straight lines across the Bay of

Fundy. Adrew and Heather aided enormously in the work of this thesis and never

doubted its completion (or not that they told me anyway).

My thesis project, and my love of (some might say obsession with) seals in general,

wodd never have corne to be without Dave Lavigne - a tnisted advisor, an admired

colleague and Enend. 1 remember the exact moment that 1 decided to work with him. Tt

was the &y &er the Sandanistas lost the election, and 1 walked into MammaIogy class to

see table after table of stuffed microtine rodents and bats. 1 observed that it seemed

pretty pointless to be studying obscure mamrnals when a great revolution had just been

defeated in Nicaragua. Dave dryly told me to sit d o m and focus on the lab, because

there were plenty of mammal-related matters to worry about. Nearly ten years, and

many animal w e k e campaigns later, his words seem quite prophetic. Seals are not, as

he once told me, "just Wre mal1 furry whales": they're much more interesting than that.

Thanks are due to the other members of my advisory cornmittee - Tom Nudds, Brad

White and BiU Leoaard - for their counsel and most especialiy their patience. Tom also

kindly stepped in as CO-advisor when needed. Thank you to the members of my

examination cornmittee for helpfûi comments and dcisms: Dm. W. Beamkh, V.

.. ll

Thomas and 1.A- McLaren. 1 also appreciate the support of my lahates, Glenn Boyle,

Carolyn Cailaghan and Isabelle Schmelzer. The denizens of the International lbfarine

Mammai Association, Inc. - especiaily, Jan Hannah, Sue Wallace, Dave Johnston, Peter

Meisenheimer and Thom Woodley, graciously shared their workspace over the course of

many years. Peter, a fiieml and confidante h m both the student movement and animal

welfare movement sides ofmy We, ais0 gave up his apartment floor as the clock ticked

d o m to the end of this project

One of the most rewarding aspects of this work was the close collaboration with the

Cree community of Whapmagoosiui. I am gratefiil to David Masty and Matthew

Mulcash for facilitating this relationship. John Petagumskum, Sr., and Sandy

Petagumskum provided invaluable advice and support in the field, and John opened a

window on to the rich history of the relationship between the fieshwater seals and his

people. Wachyia!

To my fiends (and fellow residents of 63 Koch and 134 Paisley), Ian Adams, Paul and

Irene Heaven, Mike McMullen, Merrill Stephen, Julie Stewart, and Tim Tinker, 1 owe

more thanks than 1 can say. They were my home in Guelph. 1 also owe a debt to a

number oforganisations, *ch, though they prolonged my dliliation with the

University of Guelph, taught me as much as this thesis project: the Canadian Federation

of Students, the University of Guelph Graduate Students' Association (CFS Local 62),

CUEW Local 13/CUPE Local 3913 and the International Fund for Animal Welfare

(IFAW). 1 am gmtefiil to the good fiends that 1 met through the studeut movemeut -- ..- 111

Cass Koenen, Brad Lavigne, Nona Robinson, Sarah Schmidt, Michael Temelini and

Marty Williams. Dr. Temehi - another politically involved and chronically undefendeci

Ph.D. Candidate - served as an inspiration to finish this work My IFAW colleagues and

firiends demomtmted tolenmce and good humour as I struggled to balance a fidi-time job

with the demands of this thesis.

1 thank all the individuah and institutions that provided tissue samples and access to

their zoological colIectiom: Ammcan Museum of Natlual History, New York, N. Y.; P.

Béland and the St. Lawrence Nationai ïnstitute of EcotoxicoIogy; Canadian Museum of

Nature, Ottawa, Ontario; Carnegie Museum, Pittsburgh, Pennsylvania; Department of

Fisheries and Oceans both in Montreal and in Montjoli, Quekc; Grand Manan Whale

and Seabird Research Station, North Head, NB; Marine Mammal Strauding Centre, New

Jersey; National Museum of Natural History, Washington, D.C.; Naturhistoriska

Riksmuseet, Stockholm, Sweden; New England Aquarium, Boston, Mass-; University of

Alaska Museum, Fairbanks, Alaska; University of Amsteràam Museum, Amsterdam, the

Netherlands; Zoologisk Museum and Greenland Environmental Research Institute,

Copenhagen, Denmark. 1 thank P. Wilson for aiding with the DNA analysis; T. Cox for

her Arcview expertise; K. Hobson for a i h g with the stable isotope analysis and C. van

Kessel of the Department of Soi1 Science, University of Saskatchewan, for the use of its -

facilities to analyse the stabIe-isotope samp1es; S. Iverson, Department of Biology,

Dalhousie University, for analysing the fatty acid samples; and the staff of the Centre

d'études nordiques for field support. M. McMuiIen (the originator of the Lacs des Loups

Marins seal capture technique and "Eye of the Tiger" cornpanion), D. Heaven, P. and 1

iv

Heaven (who, if you analysed their tissue d3c and S"N ratios, would Iikely stilt show

traces of Lacs des Loups Marias b m k char feeding), 1. Schmelzer, C . M . Smith, and

A. Westgate - great fiends (and in one case, f d y ) d, provided three years of sterling

field assistance. This project wouidn't have happened were it not for them.

1 owe the completion of this thesis and this chapter of my Life to Jennifer, my best

fien& Our relatiotlship began (again) as 1 was racing out the door for my nrst field

season. She asked me to move in with ber over short-wave radio. "In Iust - SprEig.. ."

AU sampling was done in cornpliance with University of Guelph Animal Care

Committee regdations and pennits fkom the Ministère de l'Environnement et de la Faune

of the Government of Qué'bec. This research was fimded by the International Marine

Mammal Association Inc., Guelph, Ont., the World Wildlife Fund-Canada Endangered

Species Recovery Fund, and a N a d Sciences and Engineering Research Council of

Canada (NSERC) operating grant to David Lavigne. 1 was provided with financial

assistauce by two Ontario Graduate Scholarships, an NSERC Post-Graduate Scholarshïp

and the Norman James Aquatic Mammalogy Scholarship.

TABLE OF CONTENTS

ACKNOWLEDGEMENTS

TABLE OF CONTENTS

LIST OF TABLES

LIST OF FIGURES

PREFACE

CWAPTERS:

1 . General Introduction ............................................................. 9

2 . Subspecïfïc variation in crania of the harbour seai. Phoca vitdina L .

Introduction Materials and Methods Redts Discussion Appendùr 1

3 . Genetic analysis of western Atlantic. Phoca vitzdina concoior Deffiy 1842. and Lacs des Loups Marins. P . v . meilonae Doutî 1942. harbour seals using mitochondrial DNA

Introduction Materials and Rfethods Redts Discussion

4 . Distinguishing between populations of fksh and saitwater harbour seals. Phoca vitulina L., using stable-isotope ratios and fatty acid profiles

Introduction MateriaIs and Methods Resuits Discussion

5 . Movements of Lacs des Loups Marins harbour seals. Phoca vitulina nrellonae Doutt 1942. monitored by satellite telernetry

Introduction Materiais and Methods R e d i s Discussion Appendix 1

6 . General Discussion

1 . Status of the Lacs des Loups Marins harbour seal, Phoca vituIiita mellome. in Canada ................................................. 186

LIST OF TABLES

Table 2.1:

Table 2.2:

Table 2.3:

Table 3.1:

Table 3.2:

Table 4.1 :

Table 4.2:

Table 4.3:

Table 4.4:

Table 5-1:

Standardid canonical discriminant function coefficients for first two fimctions. Percentage variations accounted for by the two hctiom are also given ................................................... 45

A post~ori cldcations based on discriminant scores for five P. vituiii~ subspecies, P. lmgho, and Pura hhpida ................... 47

Merins of adjusted (Iog,,-transfid and C-scored) cranial chraacta measurements arranged according to snGdy gmup (folowing Amano and Miyazaki 1992). Withinrows,meansthatareannoQtedwiththesame qxmxipt letters are signifïcady differeat (P < 00.1) ........................... 48

Specimeas of Phma w~tulina COIICO~O~ and P. v. mello~e used in the walysis of the mitocbandrial control region. ................................ 72

Kimura's DNA disbnce measures of 6 Phoca vïtuIina rnellorue and 11 P. v. concolor assuming a 2-to-1 transition-transversion substitution ratio ................................................. 73

Specimem of Phoca vitdina richaraki, P. v. concolor and P. v. rneZZome used in the siable isotope (SI) and

................ fatîy acid (FA) analyses.. ........ ,.. ..............~... .... 1 04

Stable-carbon and nitrogen isotope values (mean * SD) of hait and serum fiom seals fiom fie&, brackish and At-watex ............................................................................................. 108

Fish fauna of Lacs des Loups Marins, Que'bec: abundance and stable-carbon and nitrogen isotope values (mean * SD) fiom muscle tissue ............................................... 109

Levels of selected fatty acids in the blubber of three subspecies of harbour seai (Phoca vitulina) .......................................... 1 10

Description ofharbour seals (Phoco vihtlina mellonae) that were atlked with satellite transmitters in Lacs des

.............................................. Loups Merias, Québec in 1995 and 1996 137

Table 5.2: Mean number of uplinks per day; numkr of telemetemi positions used in analysis: average daily distance (ADD); and minimum convex polygon (MCP) and kernel home range estimates for 9 Lacs des Loups Marins seals observecl in 1995 and 1996.........-... ..... . -....-...-. .. ................................. 138

Table 6.1 : Cornparison of hshwater seal populations ui Lacs des Loups Marins and Lakes Saimaa (Finland) and Ladoga (R-a) ....... -.. ..~..... . ..-.. -.--....... .-... .... .................. . . . . 185

LIST OF FIGURES

Fig. 1.1: Known range of Phca v i td i . melLonae in relation to Hydco-Quelbec's proposed Grande Baleine hydroelectric Project .................................................................................................... 27

Fig- 2.1 : Range of the barbout seal (Phoca vïtuIiina L.) ........................................ 5 1

Fig. 2.2: Relative differencw between the group means of cranial rneasurements of 5 subspecies of harbour seal (Phoca v i f u l i ~ ~ ~ concolor,- P. v, mellonae; P. v- vitulina; P. v. richarhi and P. v. ~t tgjng~ri) , the spotted seal (Pm largha) and the ringed seal (Aua hup&dkzo) basecl on plots of tint, second and thud canonid variates ........................-................................ 53

Fig. 3.1: Consensus tree of 500 bootsirap neighbour-joining distance trees for 480 b.p. of the mitochondrial control region for 6 Phoca v i i u l i ~ m e f f o ~ e and 11 P. v. concofor. The grey seal (Halichoenu g g p s ) is the outgroup. The number 1Ocafed at the nodes indicates the percentage of bootstrap trees that contain this pattern ............................................................... 75

Fig. 3.2: Consensus tree of 500 bootstrap neighbour-joining distance trees for 385 bp. of the mitochondrid control region for 6 Phoca vituiina mellonae, Il P- v. concolor, and 34 harbour seai haplotypes reported by Stanley et al. (1996). The grey seal (Halichoerus gs>pus) is the outgroup. The number located at the nodes indicates the percentage of bootîtrap

................................................................. trees that contain this pattem 77

Fig. 4.1 : Di~ï .u î ïon of stable-carbon and nitrogen isotope ratios in the haïr of harbour seals with diets derived fiom marine (Phoca vitulina concolor; Northwest Atlantic), fieshwater and marine (P- v. concolo~ Kasegalik Lake), and hshwater (P. v. mellonae) sources. Subscripts 1 and 2 denote the 1994 and 1995 values, respectively, for the one

.................... ........ fieshwater sed that was marked and recaptured ... 1 12

Fig. 4.2: Plot of total n-3 vs. total n-6 fatty acids (as percent of total fatty acids) in the blubber of marine (Phoca Wl~lina r i c h d i

........ and P. v. concolor) and fieshwater (P. v. mellonae) harbour seais 1 14

Fig. 5.1: Study area in northem Quebec as depicted by National Topographie Data Base digitised maps ................................................ 140

............ Fig. 5.2: Sightings of Lacs des Loups Marins harbour 4 s : 1994-96 .... 142

Fig. 5.3: Movements of 8 Lacs des Loups lkhins harbour d s as monitored by satellite telemetry in 1995 and 1996. Minimum Convex Polygons enclose ail estimateci positions ................. 144

Fig. 5.4: Kemel home ranges (sith confidence areas of 25%, 50%, 75%, and 95%) of 8 Lacs des Loups Marins harbour seals

...................... dculated from sateilite-teiemetered positions: 1995-96 153

Fig. 5.5: Utilisation distri'bution (with confidence areas of 25%, SO%, 75%, and 95%) caicuiated using combined positions of 8 satellite-telemetered Lacs des Loups Marins harôour seals: 1995-96 .................................................................................. 162

PREFACE

Facts and Fables

You have to go to a windowless building in downtown Pittsburgh, Pennsylvania, in

order to see the only known museum specirnens of the mysterious Lacs des Loups Marins

harbour sezl (Ph- v i t u l i ~ rneIIonue). The story of how these specimens got there is

quite i n d i l e : the seal is probabIy the only endemic Canadian mammal to have a

popda. adventure story written about its nrSt encornter with science (Twomey 1942).

And this story isn't the only one. In fact, upon beginning this thesis project one of the

things that stnick me was the amazing numkr and muddled quality of the anecdotes

swirling around this subspecies. Separating fact fkom fiction was sometimes a challenge.

As one example, a Hydro-Quek researcher once told me with a straight f z e that they

had concluded the Lacs des Loups Marins seal was herbivorous, and spent much of its

time rooting about for vegetation at the bottom of the lake -certainiy an imprecedented

behaviour for a pinniped and one *ch the anaiyses in this thesis found to be false.

Another story that at £kt blush seemed just as improbable was relayed by a Cree elder

in the fïrst year of the project. He told me that the fkeshwater seal (or achiRicnIpiih) is

almost always solitary, and when disturbed in the middle of the lake wiil make a beeline

back to shore and try to use the contours of the shoreline to escape. Not only did this

tum out to be entirely accurate, but the propensity for hugging the shore was exactly the

behaviour that ailowed us to capture the animals and complete the project.

In addition to its sometimes strenge existence at the junction of science and science

fiction, the Lacs des Loups Marins seal has ken, and continues to be, embroiled in

events unfolding in one of the most volatile, and pol i t idy cornplex, regions of Caoada

The Carnegie Museum and Canadian Disinterest

The fieshwater seals of Ungava have been known about by Europeans since at least

1757, when Nicolas Bellin f h t mappeâ the peniasula They were the objects of interest

by Hudson's Bay Company explorers and fut traders throughout the lgm Cenniry- The

Geological Survey of Canada officially t w k note of thei. presence in 1898, but it was not

until an expedition fiom Pittsburgh's Carnegie Museum tcaveled to Lacs des Loups

Marins in 1938 that the population was describeci scientifically.

One of the more puzzlïng questions suwmding this seal is why it has been known of

for so long, with so little work having been done on its biology. Part of the reasm for

this may be the rather late transfer of the no- Ungava peninsula to fidl povincial

status (1 9 12) and, until the 1970s, the area's comparatively inaccessible and undeveloped

nature. In addition, the fieshwater d occupies a uniquely ambiguous legal position: it

is a marine mammal (a federai jurisdiction) in freshwater (mostiy a provincial

jurisdiction) (Anonymous 1990). The province is charged with the seal's management,

yet because ail govemmental marine mammal researchers are employai at the federd

level, has minimal expertise in discharging this responsibility. To hther cornpikate

matters, Lacs des Loups Marins exists just North of the 55' parallel at the northem

extreme of traditional Cree hunting territory According to the James Bay and Northern

2

Québec Agreement; however, this is huit-controlled land, placing the seais not only in a

legal grey zone between the federal and provincial govetaments, but also between the

Cree and the govemnent of Qué'bec (the Cree enjoy considerabIe autonomy in the

region), and between the Cree and Inuit.

#en h. W.E. Clyde Todd, Cunitor of Oniithology for the Carnegie Museum wrote

to the Canadian government on January 8&, 1937 to request assistance m motmthg the

expedition to Ungava, the response was astonishing. The letter back to Clyde Todd h m

Dr. RM. Anderson, Chief of the Dominion Division of Biology, exemplifies the low

priority that the Canadian govemment stül attaches to marine mammal research:

By fm the greater part of the aerial surveying in the noah

of Canada is in connection with development of

prospecthg and mining, and as there is so much pressure

brought to bear for surveys which may throw some light on

developments which have what is caiied economic

[emphasis original] value, that there is hardly any hope that

any m e y party would be sent out by aeroplme for purely

biological collecting.

Anderson went on to say that he is "sorry that me does] not have mmy constructive

ideas on Hudson Bay exploration", hopes the Carnegie wil l 'bot be disappointed if [the

seals] do not prove to be something 'new'" and suggests that Clyde Todd shouid contact

the governent of Quebec, not the govemment of Canada, regarding specimen collection

permit!%

Undeterred by Ottawa's disinterest, the Carnegie team traveled to Lacs des Loups

Marins in March, 1938, and were successful in collecting two seal specimens that were

used to describe a new subspecies. The expedition was so arduous that the accom

when published as the adventure story N e d e to the North, has has d e d the "classic

narration of the journey-que& (James 1982, p. 5). The team reached the lake by

sledding overland fiom the Hudson Bay coast, but became dangerously short of food

The photograph of the happy faces of the team members just after two se& were kiiled,

is testament to how quickly the m a t of these valuable scientific specimens was rendered

into a badly-needed stew. The pelts and s M s made it back to Pittsburgh intact.

The Last Fifty Years

Since 1938, the bulk of research occiaring in the area has been instigated by threatened

new construction by Hydro-Quekc. When Robert Bourassa was re-elected in 1986, the

architect of the James Bay hydroeleîtnc project qyickly moved ahead with "James Bay

: huge new proposed impoundments on the Great Whale, Nottaway, Broadback and

Rupert Rivers. At the National En- Board hearings in 1990, Hydro-Québec initially

tried to daim that there was no such thing as a fkeshwater seal (Posluns 1993)' while at

the same time initiating a variety of investigations of them (Consortium Gilles Shooner

& Associés 1991). The Whapmagwstui Cree spent a significant amount of t h e during

4

their presentation to the Board speakïng about their knowledge of the M w a t e r seals

(Richardson 1991; Posluns 1993). A year later, Hydra-Qué'bec issued a report stating

that the fieshwater seals were to be found in numerous lakes in the area (Consortium

Gilles Shooner & Associés 199 1)-

The genesis of this thesis also lies in the proposed Great M e hydtoelectric project -

&ch bas now been hdefinitely shelved by the Qué'bec government- In 1991, the

International Marine Mammal Association published a report entitled "Potential impacts

of Hydroelectric Development on Matine Mamrnals in Northem Q u e W ( W d e y et

al. 1992). Much of this report dealt with the potential impacts on the Lacs des Loups

Marins seal, and a later iteration of it was prrsented in 1994 to the Great Whale project

environmental assessment process (Review Bodies 1994). Mso in 1994,I authored a

status report on the Lacs des Loups Marins seal for the Cornmittee on the Statw of

Endangered Wildlife in Canada (COSEWIC) (Smith 1997). The Cornmittee was

concemed enough about the possible impacts of development on the Ungava penuwla to

List the seal as 'tulnerable" (Smith 1997). Around this same time, the International

Union for the Conservation of Nature end Naturai Resources (NCN) iisted the seais as

ccsuspected but not definitely known to be endangered, vulnerable, or rare due to a lack of

reliable information" (Reijnders et al. 1993). This thesis grew out of attempts to provide

some of the answers needed to assess the consemation status of this unique Canadia.

population.

Ironîcally, the Lacs des Loups Marins harbour seai, which has had stories told about it

for hundreâs of years, ody became the f i of susfaineci biological investigation at the

point when it seemed threatened with possible extinction by the rising waters of Hydro-

Québec dams. The animal s t i l l has no legal protection. UnWre the United States, Canada

has no Endangered Species Act or Matine Mammal Protection Act There are long-

outstanding proposais to establish both federal and provincial parks in the Lacs des

Loups Marins area, and the govenunent of Que* has listed the seal as "Wrely to be

designated as threatened or vulnerable" since 1992 (Queb 1992). R.M. Anderson's

comment nom 1937 that Y do not think that hau seals are protected any place in Canada,

particdarly in the open sea" is as tnie today as it was then.

References

Anonymous. 1990. A qui apparti-ent ce phoque? L'Actualité. 15 March.

Consortium Gilles Shooner & Associés, SOMER, and Environnement Illimité Inc, l99l.

Complexe Grande-Baleine, Avant-projet Phase 2, Büan des coanaissances sur le

phoque d'eau douce. Vice-présidence Enviromemeot, Hy&o-Que%ec

James, W.C. 1982. Introductï~n~ Pages 5-21 in Neede to the North. A. C. Twomey.

Oberon Press, Ottawa.

Posluns, M. 1993. Voices fkom the Odeyak. NC Press Limited, Toronto.

Québec (Province). 1992. Liste des espèces de la faune vertébrées suscepti'bles d'être

désignées menacées ou vulnérables. Ministère de l'Environnement et de la Faune,

QuéYbec.

Reijnders, P.J.H., S. Brasseur, JI. van der Toom, P. van der Wolf, 1. Boyd, J. Harwood,

D. Lavigne, L. Lowry, and S. Stuart w-1. 1993. Seals, fur se&, sea lions and

walruses: status of pinnipeds and conservation action plan. IUCN, Gland,

Switzerland.

M e w Bodies respom'ble for the environmental assessrnent of the proposed Great

Whaie hydroelectric project. 1994. Joint report on the conformity and quality of

the environmental impact statement for the pposed Greaî Whaie Riva

hydroelectric project.

Richardson, B. 199 1. Strangers devour the land. Douglas & McIntyre, Toronto.

Smith, RJ. 1997. Status of the Lacs des Loups Marins Harbour Seai, Phoca vituIiina

mellonae, in Canada, Can. Field-Nat. 1 1 1 : 270-276-

Twomey, A.C. 1942. Needle to the North: The story of an expedition toUngava and

the Belcher Islands, Herbert Jenkins LtL, London.

Woodey, T-H., RJ. Smith and D.M. Lavigne. 1992. Potential impacts of hydroelectric

development on marine mammals in northem Que-. IMMA Tecbnical Report

Number 92-02. International Marine Marnmal kssociation Inc., Guelph, Ontario.

THE LACS DES LOUPS MARINS HARBOUR SEAL,

Phoca vitrrlim mellonae Doutt 1942:

ECOLOGY OF A N ISOLATED POPULATION

We hold the highest respect for the fkeshwater seal; because although it was hard to hunt, it saved the Iives of entire f d e s at times past when fimine struck When all else was not avdable, by miracle a hunter killed a fieshwater seai. It was as if the Creator planned it this way.

John Petagumkum Sr., testrBng b&re the National Energy B& Ottawa, 1990 (Richardson 1991, p. 360)

CHAPTER 1 : GENERAL INTRODUCTION

Harbour seals (Phoca vitulina L.) are well hown for occasionally fiequenting lakes

and rivers throughout the world (Erlandson 1834; DeKay 1842; Ailen 1880; Browne

1909; Grenfell 1910; Prkhard 1911; Strong 1930; Dunbar 1949; Fisher 1952; Wheeler

1953; Harper 1954; Harper 1961; Beck et al. 1970; Paulbitski 1974; Roffe and Mate

1984; Wiiliamson 1988). With the possible exception of Lake niamna, Alaska, however,

the harbour seal (P. v. mellonae) population in the area of Lacs des Loups Marins (Lower

Seal Lakes) (Fig. 1.1) on the Ungava peninsula of northem Québec, is the only one

hown for which there exists evidence of year-round residency in fieshwater. Much of

this evidence is andotal and historical, and includes repeated references to the Lacs des

Loups Marins seai's restricted distribution and unique morphology and behaviour.

Historical References

The Cree people of Whapmagoostui, who have Iived and hmted in the area in question

for at least a millennium (Crowe 1991). consider the cumnt range of P. v. m e l l o ~ e to be

Lacs des Loups Marins, Petit Lac des Loups Marias, and Lac Bourdel, with some reports

of animals having once been in Lac à l'Eau Claire (Clearwater Lake) (Posluas 1993; J.

Petagumslaim Sr., Whapmagoostui, pers, comm.). This information is conoboraied by

the Cree toponyms in this area, wuhich make reference to achikw (seal) and achiRun@ih

(fieshwater seal) (Consortium Gilles Shooner & Associés 1991). The Cree contend that

the fieshwater seal bas a number of unique attri'butes, wbich are describeci quite

specifically in a recent interview with one of their elders, James Kawapit:

The fieshwater seds look différent in bat their fiir is much

darker. Their markings are more numerous and are dark.

The marine seals are much iighter in colour. They also

have different habits, The other difference is in the taste,

The Ecesbwater se& taste more like fish and their fat tastes

different îrom the marine ones (Posluns 1993, p. 90).

Kawapit's account of the morphological distinctness of P. v. meIIonae is strikingly

similar to that of George Atkinson II who. on July 22, 18 18, noted that:

At these ripp1es is a fishing place whae the indians set nets

for seals in the winter; they are quite a different species to

those on the sea coast; their skin being covered with short

silky hair; yesterday and this day we saw several in the

lakes and rivers; they are in general SIMU and shyer than

the 0th- and much darker in color (p. 61).

Though Atkinson's sighting of seals in Upper Seal Lake (Petit Lac des Loups Marins),

during one of the earliest Hudsonts Bay Company expeditions hto the Ungava interior,

is the first written description of these animals, the phrase "Lacs des Loups Mark&' is in

fact much older. The French cartogapher Nicolas Bellin's "Petite Atlas Maritime''

(1757) seems to have coined the term, establi-ching a written record of seals in this erea

dating back nearly 250 years. Sub~e~uently, other sightïngs were made in the vicinity of

Lac d'Iberville and Petit Lac des Loups Uarins (J. Clouston, July, 1820) and Lacs des

Loups Marins (W. Hendry, July, 1828; N. Fuilayson, July, 1830; Rev. E.J. Peck, July,

1884 in Lewis 1904). Near the tum of the century, during his traverse of the Ungava

peninsula for the Geological Sinvey of Canada, A.P. Low observed seals in Lacs des

Loups Marins (Juiy, 1896)' and reported that "the hdians kill annually more than thirty,

showing that the animal breeds fieely in the fie& water" &ow 1898, p. 13).

Written references to the dissimilarity between the Lacs des Loups Marins seals and

oceanic harbout seals are not restricted to those of Kawapit and Atkinson. Hendry's

(1828) journal makes mention of the '%ne quality" (p. 84) of the fieshwater seal s h

and, in 1 888-89, there are a nimiber of specinc references to fkeshwater seal pelts in trade

in the records of the Little Whale River pst (Hudson's Bay Company Archives, Little

Whale River Inventories, Country Produce, B.373W22, fo. 14d and B.373/d/23, fo. 13d).

During his travels on the Ungava peninsula, Flaherty (19 18) was told of seals in Lake

Minto (just to the North of Lacs des Loups Marins):

11

The Iake is famous among the Eskimos as the habitat of the

fiesh-water seal, hunted primarily not as food, but for the

pelt, which, much darker, softer, and more 1ustrous than

that of the sait-water variety, is used for their fher

garments (p. 119).

More recently, d e r extensive research among the Inuit of northem Qué'bec, Grabum

(1969) descri'bed the fkshwater seaï slrins as king among '?he softest and most

beautifidly marked of ail ... [that] btmg top prices when traded." (p. 20).

Subspecies Description

The Lacs des Loups Marins s d was bmught to the attention of a wider audience in

1942 by the publication of its subspecific description (Doutt 1942) and a popular book

entitled Neede to the North (Twomey 1942). Both these works were the products of a

Carnegie Museum expedition to Lacs des Loups Marins in March, 1938, which resulted

in the collection of two type specimens. Doutt was f d a r with the anecdotal accounts

of P. v. ntellonae 's u n d chatacteristics and restricted distribution, and he

endeavoured to use a more rigorous scientinc analysis to evaluate the evidence. The

subspecies was d e s m i primarily on the basis of its unusually dark pelage and an

enlarged coronoid process on the manàiible, and on the premise that the population had

been isolated foc 3000-8000 years, trapped by the Ungava peninsuia's isostatic rebound

since the retreat of the Laurentian ice sheet Wutt 1942). Interestingiy, this was an idea

that had fkst been advanceci by Low (1898):

The harbour seal is known to travel ovdand for

considerabIe distances, but its presence in this Iake nearly a

hmdred miles h m Salt-water at an elevation of nearly 800

feet above the sea, can hardy be due to its migration up

such a rough stream as the Nastapoka, Another way in

which it might have reached the lake was during the

subsidence ofthe land at the close ofthe glacial period,

when the lake was nearer sea-level than at present by more

than 600 feet, and when the deep bay extended inland up

the present valley of the Nastapoka to or near the outlet of

the lake, with such conditions it would be easy for seals to

reach the lake, and having found it full of fish they

probably lost the inclination to r e m to the sea (Low 1898,

p. 13).

Radio-carbon dating has recently been used to estimate mme precisely the deglaciation

date in the coastal hiUs of the Nastapoca River area to be 7300 years BP (Allard and

Séguin 1985).

Doutt's subspecies description was accepted by a number of authors such as Simpson

(1944), Anderson (1946), Davies (1958), Scheffer (1958), Banfield (1974). and Bigg

(198 1). George Gaylord Simpson, in his classic text Tentpo and Mode in Evolution,

acnially used the Lacs des Loups Uarins seal, and Doutt's estimate of its time of

isolation, as a case study for calculahg a rate of evolution (Simpson 1944). In Doutt's

words:

I estunated that a new subspecies was produced in about

5,500 y-, plus or minus 2,500 years. Simpson (1944, p.

19) points out that this wouîd k about 1,000 generations.

He suggests that in some rodents subspecific differentiation

might taLe place in even less than 3 0 grnerations @outt

1955).

0th- authors disputed the subspecies designation, arguing that the supposed

craniological anomalies of P. v. mehnae were inconclusive, and that the seals were

likely able to travel b I y between salt and fieshwater (Maasfield and McLaren 1958;

Mansfield 1967; Smith and Horonowitsch 1987; also see Honaclci et al. 1982; King

1983; Wüg 1989; Reeves et al. 1992). Mansfield (1967) also contended that because

Arctic harbour seals are, as a rule, much darker than those fiom the southern part of the

range, the dark colour of P. v. rnel201~1e's pelage is not diagnostic, but rather a

characteristic that the fieshwater animais share with nearby oceanic harbour d s .

Since Doutt's description, a continueà number of seal sightings fiom ail times of the

year have been made in Lacs des Loups M&hx In August, 1953 (Doutt 1954); August,

1976 (Power and Gregoire 1978); April, 1984 (i3errouard 1984); and Mach, 1986

(Smith and Horonowitsch 1987). Other recent evidence, however, nom sightings and

hydro-acoustic data compiled by Hy&o -Québec, indicate that P. v- mel lo~e may be

distriiuted over a wider area than previously thought and, consequently, have a greater

potential to overlap the range of oceanic harborn 4 s (Coosortïum Gilles Shooner &

Associés 199 1).

Impiicatiom for Conservation

Though it is rlifficuit to make precise estimates of population size due to a lack of

information, it seems clear that the Lacs des Loups h k h s seals are few in nmber

(Consortium Gilles Shooner & Associés 1991). The tendency of harbour seals to be

distriiuted in small local populations makes them sensitive to disturbance (Boulva and

McLaren 1979; Maine Seal 1994), and there are a number of examples of local harbour

sed populations beuig extirpated, or their numbers reduced, by human activity (DeKay

1842; Allen 1880; Teilmaun and Die= 1993; Reijnders et al. 1993). The likelihood that

the Lacs des Loups Marins seal population is similady vulnerable, and the potential

threat of fùture development - particularly hydroelectnc development - on the Ungava

peninsula, prornpted the Intemational Union for the Conservation of Nature and Natural

Resources (nrCN) to List P. v. me2Ionae as being "suspected but not dennitely known to

be endangered, YUlIlerable, or rare due to a lack of diable informatioa" (Reijnders et al.

1993). The Cornmittee on the Status of Endangered Wildlife in Cariada (COSEWIC) has

designated the population as "vuinerable" (Smith 1997). The government of Quékc bas

also listed the population as "likely to be designated as threatened or VUlIlerable" (Quebec

1992a). and is considering whether to give legal protection to a portion of P. v.

melIonuefs range (Dubreuil1983; Quebec 1992b). Further investigation of the

distribution of P. v. m e l l o ~ e and the degree to which it is a distinct population is,

therefore, extremeIy relevant to fuhne conservation efforts.

Thesis Description

The objective of this thesis was to test Doutt's hypothesis that the Lacs des Loups

Marins seal is an isolated group that is distinguishable fiom neighbouring oceanic

harbour seal populations. This was achieved through the mtegration of sources of data

fiom investigations ofnatural history, morphology, molecuiar genetics, feeding, and

range and distribution.

The second chapter of the thesis builds on Doun's original analysis and contains

morphometric cornparisons of crania from P. v. mellonue aod the 0 t h four putative

subspecies of harhur seal, according to the classification of Bigg (1 98 1). The third

chapter is an analysis of the genetic diversity of harbour seals in northeastem North

Amerka, including P. v. mellonae, accomplished through the cornparison of a 500 bp

region of the mitochondrial control region. The fourth chapter is an investigation of

habitat use by P. v. mellonae as revealed by stable isotope ratios and fatty acid profiles.

In the fifth chapter, satellite telernetry techniques are used to examine the distn'bution and

range of P. v. melZonae. The status report on P. v. tneilonoe that was prepared for the

Cornmittee on the Status of Endangered Wildlife in Canada (Smith 1997) is provided in

Appendix 1.

References

Allard, M., and M X SégUm. 1985. La déglaciation d'une partie du versant hudsonien

que'bécois: bassins des rivières Nastapoca, Shekdrake et A L'eau Claire.

Géographie physique et Quaternaire 39: 13 :M.

M e n , J.A. 1880. History of North American pinnipeds - A monograph of the walnises,

sea-Eons, sea-bears and d s of North AmerÏca- US. Geological and

Geographicai Swvey of the Territories. Misce11aneous Publication No.12.

Anderson, RM. 1946. Catalogue of Canadian recent mammals. National Museum of

Canada Bulletin No. 102. King's Rinter, Ottawa

Atkinson, G. 1818. J o d of George Atkinson II. Page 6 1 in Noahem Quebec and

Labrador j o d s and correspondence 18 19-35. 1963. Edited by KG. Davies

and A. M. Johnson. The Hudson's Bay Record Society, London.

Banfield, A.W.F. 1974. The mammals of Canada. University of Toronto Ress,

Toronto, Ont.

Beck, B., T.G. Smith, and A.W. Mansfield. 1970. Occurrence of the harbour seal,

Phoca vihrlina, Linnaeus in the Thlewiaza River, N. W.T. Cm. Field-Nat.

84:297-300.

Berrouard, D. 1984. Résultats &me reconnaissance aérienne dans le but de repérer des

sous-populations de phoques communs (Phoca vitulina) dans la région des lacs

des Loups Marins. Ministère de l'Environnement, Direction régionaie du

Nouveau-Que'bec,

Bigg, M.A. 1981. Hahm seai - Phoca vitdina Linnaeus, 1758 and P. lmgha Pallas,

18 1 1, Pages 1-27 nt Handbook of marine msmimals, Vol. 2: Seals. Edr'fed by

S.H. Ridgway and RJ. Harrison. Academic Ress, London.

Boulva, J., and LA. McLaren. 1979. Biology of the harbor seal, Phoca vitulina, in

eastern Canada. Bull. Fish. Res. Board Canada 200.

Browne, P.W. 1909. Where the fishers go - The story of Labrador. The Musson Book

Company Ltd., Toronto.

Clouston, J. 1820. Second journal of James CIouston. Pages 29-68 in Northern Qwbec

and Labrador joumals and correspondence 18 19-35. 1963. Edited by KG.

Davies and A. M. Johnson. The Hudsods Bay Record Society, London.

Consortium Gilles Shooner & Associés, SOMER, and Environnement Illimite Inc. 1991.

Complexe GrandedeBaleine, Avant-projet Phase 2, Büan des connaisuuices sur le

phoque d'eau douce. Vice-présidence Environnement, Hy&o-Quebec.

Crowe, kJ. 199 1. A history of the original peoples of northern Canada. McGill-

Queen's University Press.

Davies, IL. 1958. Pleistocene geography and the dktri'bution of n o f pin.peds.

Ecology 39:97-113.

DeKay, J.E. 1842. Naturai history of New York, Part 1. Zoology. D. Appleton and Co.

and Wiley and Putnam, New York, NY.

Doutt, J.K. 1942. A review of the genus Phoco. Ann. Carnegie Mus. 29:61-125.

Doutt, I.K. 1954. Observations on mammals dong the east Coast of Hudson Bay and the

interior of Ungava. Ann. Carnegie Mus. 33 :23 5-249.

Doutt, I.K. 1955. Tenninology of microgeographic races in mammals Syst. 2001.

4: 179-185.

Dubreuil, C. 1983. Un statut de conservation pour le phoque commun habitant les eaux

douces du Que*- Ministère de l'Enviromement, Direction des réserves

écologiques et des sites naturels.

Dunbar, M.J. 1949. The pinnipedia of the Arctic and Subarctic. Bull. Fish. Res. Board

Canada 85: 1-22.

19

Erlandson, E. 1834. J o d of Erland Erlandson. Pages 247-259 m Northem Quebec

and Labrador joumals and correspondence 18 19-35. 1963. E d i t d by K-G.

Davies and AM. Johnson. The Hudson's Bay Record Societyy London.

Everitt, m., and &W. Braham. 1980. Aeriaf survey of PacifIc harbor d s in the

southeastem Bering Sea Northwest Sci. 54:28 1-288.

Finlayson, N. 1830. Journal ofNicol Finlayson. Pages 101476 in Northern Quebec

and Labrador joumals and correspondence 18 19-35. 1963. Edited by KG.

Davies and AM. Johnson. The Hudson's Bay Record Societyy London-

Fisher, H.D. 1952. The statu ofthe harbour seal in British Columbia, with particuiar

reference to the Skeena River. Bull. Fish. Res. Board Canada 93.

Flahertyy R J. 19 18. Two traverses across Ungava peninsula, Labrador. Geographicai

Review 6: 116-132.

Grabum, N-EtH. 1969- Eskimos without igloos - Social and ecowmic development in

Sugluk. Littie, Brown and Company, Boston.

Grenfeil, W.T. 1910. Labrador - The country and the people. The MacMillan

Company, New York, NY.

Harper, F. 1956. The mammals of Keewatin. University of Kensas Museum of Natural

fistory Mi-scel.Ianeous Publication 12: 1-94,

Harper, F. 1961. Land and fiesh-water mamrnds of the Ungava peninsula University

of Kaasas Museum of Nahwl EIïstory Miscellaneous Publication 27: 1-178.

Hendry, W. 1828. Journal of William Hendry. Pages 69-99 in Northern Quebec and

Labrador j o d s and correspondence 18 19-35. 1963. EdM by K.G. Davies

and A.M Johnson. The Hudson's Bay Record Society, London.

Honacki, J.H., LE. Kinman, and LW. Koeppl. 1982. Mammal species of the world.

Auen Press and the Association of Systematics Coliections, Lawrence, Kansas.

James, W.C. 1982. Epüogue. Pages 203-205 in Needle to the North. A. C. Twomey.

Oberon Press, Ottawa.

King, 1. 1983. Seals of the world. 2nd Edition. British Museum ( N a d History) and

Corne11 U~versity Press, Ithaca, NY.

Lewis, A. 1904. The life and work of EJ. Peck among the Eskimos. Hodder and

S toughton, London.

Low, A.P. 1898. Report on a traverse of the northern part of the Labrador penmsula

Srom Richmond Guifto Ungava Bay. Geological Sinvey ofcanada Annual

Report (n.s.) 9A-43.

Maine SeaL 1994- Boater etiqyette around harbour seals. Maine Seal, Lincoln, MA.

Manfield, A-W., and I.A. McLaren. 1958. Status of the m u r seal in the eastem

Canadian Arctic. Pages 46-50 in Fisheries Research Board of Canada, Arctic

unit, annual report and investigators' summaries. Edired by H.D. Fisher.

Fisheries Research Board of Canada, Montreal, QuéIbec.

Mansfield, A.W. 1967. Distrriution of the harbor seal, Phoca vitulina Linnaeus, in

Canadian arctic waters. J. Mammal. 48:249-257.

Paulbitski, PA. 1974. Pinnipeds obsened in rivers of northem California, California

Fish and Game 60:48-49.

Poslrms, M. 1993. Voices firom the Odeyak. NC Ress Limiteci, Toronto.

Power, G., and I. Gregoire. 1978. Redation by fieshwater seals on the fish community

of Lower Seal Lake, Quebec. Jorn. Fish. Res. Board Canada 35:844-850.

Richard, H.H. 19 1 1. Through trackless Labrador. William Heinemann, London.

22

Que'bec (Province). 1992% Liste des espèces de la faune verte'brées suscepb'bles d'être

désignées menacées ou vuhérables. Ministère de l'Environnement et de la Faune,

Québec.

Q u e h (Province). 1992b. Réseau des réserves icologiques au Que%ec (projets et

réalisations). Ministère de l'Environnement, Direction de la consewation et du

patrimoine éoologique.

Reeves, R&, B.S. Stewart, and S. Leatherw006 1992, The Sierra Club handbook of

seals and sitenians- Sierra Club Books, San Francisco, CA.

Reijnders, P.J.H., S. Biasseur, J. van der Toom, P. van der Wolf, 1. Boyd, J. Harwood,

D. Lavigne, L. Lowry and S. Stuart [eds.], 1993. Seals, fut se&, sea lions and

wairuses: status of pinnipeds and conservation action plan. IUCN, Gland,

Switzerlaad.

Richardson, B. 1991. Strangers devour the land. Douglas & McIntyre, Toronto.

Roffe, T.J., and B.R Mate. 1984. Abundances and feeding habits of pinnipeds in the

Rogue River, Oregon. I. Wildlife Mimage. 48: 1262-1274.

Scheffêr, V.B. 1958. Seals, sea lions and w a b s - A review of the pinnipedia

Stadord University Press, Stanford, California.

23

Simpson, G.G. 1944. Tempo and mode in evolution* Columbia University Press, New

York, NY.

Smith, RJ. 1997. Status of the Lacs des Loups Marins Harbour Seal, Phoca Mtulina

rnellonae, in Canada. Cm. Field-Nat, 11 1: 270-276.

Smith, T.G., and G. Horonowitsch. 1987. Harbour seals in the Lacs des Loups Miirim

and easteni Hudson Bay drainage- Can. Tech. Rep. Fish. AQuat. Sci. 1539.

Strong, W.D. 1930. Notes on mammals of the Labrador interior. J. Mammal. 1 1 : 1- 10.

T e h m J., and R Dietz. 1993. Status of the harbour seal (Phoca Mtulina concolor

L.) in Greenlaad- Greenland Environmental Research Institute, Technical Report.

Twomey, AC. 1942. Needle to the North: The story of an expedition to Ungava and

the Beicher Islands. Herbert Jenkins Ltd-, London.

Wheeler, E.P. 1953. Notes on pinnipedia. J. Mammal. 34353-255.

Wiig, 0. 1989. A description of common seals, Phoca vitdina L. 1758, fiom Svalbard.

Mar. Mamm. Sci, 5: 149-158,

Williamson, G E 1988. Seals in Loch Ness. Sci. Rep. Whales Res. Iiist. 39: 15 1-157.

Fig. 1.1. Known range of P h m Mîulina mellonoe in relation to Hydro-Que'bec's

proposed Grande Baleine hydroelectric project.

AREA

O 20 40 60 80 lWKM -

CHAPTER 2: SUBSPECIFIC VARIATION IN CRANIA

OF TEE HARBOUR SEAL, Phoca v i~Iina L.

A mythic beast requires a mythic country to inhabit ... Now known as Nouveau-QueTbec, Ungava stretches east h m Hudson Bay to the Labrador boundary, and north fiom the Eastmain and Hamilton Rivers to Hudson Strait (James 1982, p. 9)

htrod~ction.

The harbour sed, Phoca vituIina K.), occurs in both the North Atlantic and Noah

Pacinc Oceans over a wide range of latitudes (Fig. 2.1). Five subspecies were descri'bed

by Bigg (1981) on the basis of geographical disiribution: P. v. vituZina - found fiom

northern Portugal (rare) to the Barents Sea (rare), Iceland, the Faeroe Islands, and

Spitsbergen; P. v. concolor - found fkom northern Florida (rare South of Massachusetts)

to Hudson Bay, to Admiralty Inlet on Baffin Island and to southeasteni and western

Gredand, but absent West of Melville Peninsula and northeni Baffin Island to the

MacKenzie River delta; P. v. mellonae - a land-locked popdation found in the area of

Lacs des Loups Marins (Seal Lakes), Que'bec, Canada; P. v. richardsi - found dong the

centraLwest coast of Baja California, Mexico, North to the Aleutian Islands, Bristol Bay,

and the h i i l o f Islands; P. v. stejnegeri - found on northeastern Hokkaido, Japan, the

Kuril Islands to southem and eastern Kamchatka and Commander Islands, and with an

uncIear range around the Aleutian Islands (also see Rice 1998).

The phylogenetic relationships among the different populations of P. vitulina are

poorly understood (Bigg 1981), with the one study of worldwide diversity in harbour

28

seals having omitted P. v. mellonae h m the analysis (Staniey et al. 1996). The extent of

isolation between P. v. n'char& and P. v. stejnegeri in the Aleutians has been a matter of

contention, with Burns et al. (1984) estimating that the point of demarcation between the

two groups occurs near the end of the Alaska peninsula and more tecent investigations

(O'Corry-Crowe and W d a k e 1997; Rice 1998) pointing to Ùitergmdation between the

groups. The ice-breeding spotteci seai, P. lmgho, o f f e North Pacific, previously

regarded as a subspecies of P. Mr~rii l l~ (Burns et al 1984). was eievated to the level of

species on the basis of morphological, physiological and behavioural difEêrences

(Shaugbnessy and Fay 1977; Burns et al. 1984; Delyamure et aL 1984). It is now known

that p. hghu is genetidy distinguishable fkom P. v i l u i i ~ (Mouchaty et ai. 1995;

O'Corry-Crowe and Westlake 1997).

Pacitïc and Atlantic harbour seds can be differentiated by a few skulî characteristics

(Doutt 1942; Chapskü 1969). DBerentiation among Atlantic harbour seals is, however,

less ce*. Stanley et al. (1996) were the nrSt to review the taxonomy of P. v. virulina

and P. v. concolor since Scheffei (1958). and found that they were also genetically

differentiated.

P. v. mellonae was descri'bed as a subspecies by Doutt (1942) on the basis of the

musually dark colour of its pelage and the enlarged coronoid process of its maadi'ble.

Doutt (1942) and Davies (1958) believed that this seal has been land-lucked in the

interior of northem Qué'bec for approximately 5000 years, having been trapped in Lacs

des Loups Marins since the last ice-age by the isostatic reboimd of the Ungava peninsula

This hypothesis was disputed by hlansneld (1967), who suggested that the supposeci

morphological anomalies of the P. v. mellonae specimens could k aitif- of DoutYs

small sample size (n = 2) rather than king characteristic of an entire population-

Mansfield (1967) also thought it Uely that the seals travelled k l y between Lacs des

Loups Marins and the ocean, ody entering fkeshwater to feed oppommistidiy. He

pointed out that this was a common behaviour of barbout seals (Beck et 41. 1970; Roffe

and Mate 1984; Williamson 1988). The most rrcent review ofthe subject (Smith et ol.

1994), however, which was based on data nom this thesis, was unable to reject the

hypothesis that the Lacs des Loups Marins seal is a unique subspecies. Rice (1998)

subsequently accepted P. v- mellonae's subspecific designation in his review of marine

mammal systematics.

Though there remain considerable disagreements over some aspects of pinniped

phylogeny, there is an emerging consensus in the literature regardhg the biogeographic

history of the phocids (Hoberg and Adams 1992). It is suggested that the earliest phocid

seals appeared in the North Atlantic approximately 15 million years ago, subsequently

dispersing into the Pacific through the Arctic basin during the nrst opening of the Bering

Strait, approximately 3 million years ago (Repenning et al. 1979; Wyss 1987). Recent

work on the organisation and structure of heIminthic parasite fauaas among phocid hosts

supports these conclusions (Hoberg and Adams 1992).

The fossil record, and parasitological and molecular genetic data indicate, therefore,

that P. v. concolor is most closely related to P. v. viîulina, P. v. richarhi is most closely

related to P. v. stejngd and, following the reasoning of Doutî (1942). P. v. mellonae is

most closely related to P. v. concolor. In die first global examination ofharbour seal

morphology, the present shidy investigated these hypothesised relatioaships using a

multivariate analysis of craniomeûic traits. Such analyses have been extensively used

with other marine mammai species to elucidate patterns in infita-specinc variation (e.g

Hyvarinen and Nieminen 1990; Ross and Cockcroft 1990; Amano and M i y d 1992).

Materials and Methods

Collections of P. vitdina, Pm lmgha and ringed seal (Pusu hispida) mania were

examined at the following institutions: Arctic Biological Station, Fisbaies and Oceans

Canada, Montréal, Qué'bec; Carnegie Museum, Pittsburgh, Pennsylvania; National

Museum of Natrrral History, Washington, D. C.; Canadian Museum of Nature, Ottawa,

Ontario; Grand Manan Whde and Seabid Research Station, Grand Manan Is., N.B.

Canada; University of Alaska Museum, Fairbanks, Alaska; American Museum of Nahirai

History, New York, N. Y.; Naturhistoriska RüEsmuseet, St~kholm, Swedeu; Zwlogisk

Museum, Copenhagen, Denmark; University of Amsterdam hTseum, Amsterdam, the

Netherlands (Appeadix 1).

A total of 303 crania were examine& 5 P. v. meflonae, 61 P. v. vituIina, 60 P- v.

concolor, 45 P. v. richmdpi, 41 P. v. stejnegmi, 42 spotteci seals (P. largha) and 49

ringed seals (P. hiâpida) (Appendix 1). hie to the tiagmentary nature of some

specimens, only 25 metric and 4 non-metric characters describeci by an existùig

experimental protocol (Burns et ai. 1984) were examined on each skull (Table 2.1).

Measurements were log,,-transformed and the data analysed using the Statktical

Package for the Srniai Sciences (SPSS-X).

A MANOVA on the sexed specimens indicated that there was no interaction between

taxonomie group and sex (wilk's X = 0.53, P = 0.3 1); thus, data fiom male, f d e and

unsexed specimeas were pooled in subsequent analyses. The effects of individual size

ciiffierences were minimiseci using the "C-score" method (HoweMs L986), which ùivolved

calcdating the difference between the 2-score of a single measmement for a given

individual, and the mean 2-score of that individual for all the measurements used in the

analysis. The means of each study gmup's C-scored character measurements were

calcuiated, and signiscant ciiffierences between these means analysed using ANOVAs and

Tukey-Kramer HSD tests (P c 0.01). The data were then submitted to a discriminant

analysis (Zar 1984). F-statistics were calcuiated nom the Mahalanobis D~ distance

measures computed between the 7 groups of seals and a postet+ori classifications made

based on the discriminant scores.

Results

The first two bct ions of the discriminant analysis together accounted for about 83.6%

of the variation between the 7 groups of seals (T'able 2.1). A plot of the nrst three

canonical variates showed a separation between the 7 groups consistent with the

hypothesised relationships: the Pacifie harbour seal subspecies were closest in the

discrifnitlsiflt space; the Atlantic harbour seai subspecies and P. v. meIZonue grouped

closest together; P. loghn and P. hhpiirh were at some distance fkom the harbour seai

subspecies (Fig. 22). The F-statistics calculated nom the Mahalonobis Df measurements

indicated that a l I group meam were significantly dïffierent nom one another (P <

0.000 1). A postenori classifications, based on the discriminant scores, were 83% for P.

v. rnellonue as compared to 98% for P. hispida, 87% for P. v. concolor, and 78% for P-

v. richardlri (Table 2.2).

AU 29 characters demonstrated signifiant subspecinc variaîïon (ANOVA; P < 0.01,

(Table 2.3). When submiaed to Tulcey-Kramer HSD tests, many more d a l characters

(n = 20) exhibited signifïcant differences between Atlantic and Pacinc Ocean harbour

sed subspecies than did characters which exhiaiteci signifiant variation between

subspecies within the same ocean (n = 5) (Table 2.3).

In general, the Pacinc rrania were larger than the Atlantic cranïa (character CL in

Table 2.3). The extent of contact between the premaxillaries and nasais (character PTN

in Table 2.3) was found to be valuable in distinguishing Atlantic (iittle contact) from

Pacific (extensive contact) animals. It was also found that the media1 length of the nasals

which were in contact with the f?ontals (character NFO) was greater in the Atlantic

subspecies than in the Pacinc subspecies. The Least interorbitai width was si@cantly

different between P. v. vitdina (larger) and P. v. stejnegefi (smaller).

Within the Atlantic ocean, P. v. vitdina showed significant differences nom P. v.

concolor in only 4 cbaracters: total length of the nasais (LN), the length of the maxillo-

fiontal suture to the anterior end of the nasals (MAN), the greatest width of the extemal

nares and the degree of extension of the premaxillaries towards the d s 0

(Table 2.3).

Within the Pacinc ocean, P. v. richar& showed significant differences fiom P. W.

stejnegeri in ody 1 c h t e r : the width of the buiia (WEl).

P. v. m e l l m e also exhi'bited signiticant differences fiom the Pacfic Ocean subspecies

in the degree of the extension of the prrmaullary towards the n a d (PTN) and was

simila. to P. v. concolor with respect to this character. When compared to P. v.

concolor, the population to which it is most closely relateci, P. v. mellonue was found to

have upper premolars which are set paraUeI, rather than obliqyely, to the angle of the jaw

(SPT), a longer distance between the end of the snout and the anterior edge of the d s

(SAN) and a smaller interorbital width (Lw. Interestingly, the mandile height at the

coronoid, one of the primary criteria with whkh Doutt (1942) descriid the subspecies,

was not significantly larger in P. v. mellonae than in P. v. concolor.

The crania of P. Iurgha and P. hispidu were weil-dinérentiated (Table 2.3). The height

of the foramen magnum (HFM) in P. logho was si@cantly smaller than that of the

Atlantic Ocean harbour seal subspecies but not si@cantly diffetent than that of the

Pacific Ocean harbour seai subspecies. The teeth of P. larghu were generally set paralle1

rather than obliqudy to the angîe of the jaw (character SPT).

Discussion

The results of the craniometnc analyses support the classification of Bigg (198 1): the

relatively large amornt of variabüity between the Atlantic and Pacinc harbour seai

subspecies and reiatively d amount ofvarïability between the subspecies within each

ocean are consistent with the known biogeopphic history of the phocid seals.

According to Davies (1958), "... if vitdina differentiation dates h m the last glacial, then

the separaton between Pacifc and Atlautic groups wodd be sïightly older than that

between eastern and western groups ui each ocean." Davies (1958) went on to speculate

that the evolution of the P. v i t u l i ~ subspecies was dependent on sequentiai episodes of

dispersai (fiom the Atlantic via the Arctic) and subsequent vicariance. This view has

been supported by other studies of biogeography and pinniped host-parasite associations

(Hoberg and Adams 1992) and cladistic analyses (DeMuizon 1982). The molecular

genetic analysis of Stanley et al. (1996) indicated that harbour seal populations in the

Atlantic and Pacinc Oceans and East and West coast populations of these oceans were

significantly genetically diA'erentiated, and that Pacific and Atlantic harbour seals have

been isolated for 1.7-2.2 million years. Fuaher, in the Atlantic, the pattern of genetic

divergence suggests that the colonisation proceeded fiom the West Atlantic coast of

North America to the North and then East to Europe, and the degree of divergence

between European and West Atlantic populations suggests this colonisation may have

begun fiom 0.9 to 1.3 mülion years ago (Stanley et al. 1996). In the P d c Ocean,

genotypes in the Northwest are basal to those in the East, suggesting a West to East

colonisation pattern similar to that found in the Atlantic (Stanley et al. 1996). The

morphological, behaviourai and genetic evidence that P. v. richardri and P - v- stqhegeeri

have doser affinities with North Atlantic popdations of f . vihrzim than with P. h @ a

suggests the importance of mulbiple events of dispersal and vi-ance maing the Mid to

Late Pleistocene (Hoberg and Adams 1992).

The r e d t s of the present study confirm the coaclusioos of many extant cr;miometric

analyses. The extent of contact between the premaxillaries and nasals was nrsC descri'bed

by True (1899) to be vaiuabie in distiaguishing Atlantic nom Pacific animais. The

medial length of the nasals in contact with the fkontals was used by Chapskii (1969) for

the same purpose. In agreement with the observation of Doutt (1942), the degree of

extension of the premaxillary towards the nasal (PTN) in P. v. meIlorne was similar to

that of P. v. concolor. The signincant difference between the Least interorbital width of

Atlantic and Pacific populations is interesting, considering that it is a rneasurement which

Wiig (1989) noticed to be remarkably s m d in his P. v. vituIina crania nom Svalbard.

The fact that tk =& western Atlantic populations showed signincant differences

in only 4 characters and, when mis-classified by the apteriori analysis, wcre rnost

ofien mistaken for each other (87% correct classification for the western Atlantic animals

and 92% for the eastern Atlantic animals) (Smith et al. 1994). is reflective of Doutt's

(1 942) difficdty in stating "how specimens (of hatbour seals ] fiom the American side of

the Atlantic can be distinguished fiom specirnens taken on the European side." That the

eastern and western Pacinc populations showed a sigdicant difference in ody 1

character, and that, in the a posteriori classificatioa, these two groups were more ofien

mis-classified as each other than any other two groups (78% correct classification for

each group) (Smith et al. 1994). is consistent with the confusion which nurounds the

1Ocation of the demarcati*on betweeu these groups (Burns et al 1984). The observation

of Shaughnessy and Fay (1977). that "richardsi dîffets fiom Zargha in the same

characters and in the same ways as does stejnegdt, was also supporteci (see characters

MW, CWI, DMM, LW, PWM and PMI). The small height of the foramen magnum

and the fact that the teeth are set paraUeL to the angle of the jaw in P. I q h are bot.

characters that have been previously descriibed (Shaughnessy and Fay 1977; Guzylack

and Robineau 1993)-

The results indicated that P. v- mellonae is differmtiated fkom other harbour seal

subspecies and that, consistent with the biogeographic evidence, is most closely related to

P. v. concolor (Dom 1942; Davies 1958). AU P. W. melZonae crani-a were correctly

classified by the craniometric analysis, and ody one P. v. siejmgerï crania was

incomectly classifieci into this g r o p (Table 2.2). These r d t s are similar to those of a

smaiier craniometric analysis that was undertaken by Hydro-Québec researchers

(Consortium Giiles Shooner & Associes et al. 199 1).

Using a similar analysis, Temte (1 99 la) was able to differentiate between crania of

local populations of harbour seals on the Pacinc coast of North America He found a

significant relationship between diffei.eaces in cranid morphometry and population

differences in budi timing, and suggested that these "dochronic", as well as aiiopatric

factors, may be acting to promote population differemtiation. With respect to P. v.

mellonae there is evidence that, in addition to the population's apparent geographic

isolation fiom other harbour seai populations, similar dochronic mechanisms may be at

play. From his conversations with the indigenous peoples of northem Qué'bec, Doutt

(1942) reported that the mean pupping date of the population o c c d in early May.

This date has also been corrobotated by more recent work (Coosottium Gilles Shooner &

Associés et al. 1991). The f d e P. v. mellonae specimen coliected by h u i t (1942) on

March 22,1938 was carrying a 60-cm f m . By applyiag this information to a prenatal

growth cuve that was fonnulated for the P. v. c011color population fiom Sable Island,

Nova Scotia (which included a iinear inçrease in length of 3 -6 mm pet day, and a mean

lengih at birth of between 76 and 85 cm), the mean birth &te for P. W. mellonae can be

estimated as king between 6 and 3 1 May (Boulva and McLaren 1979). Using a prenatal

growth rate of 4.4 mm pet &y and a mean length at birth of 8 1.6 cm, such as were

observed in P. v. ficharhi in British Columbia (Bigg 1969), the estimated mean birth

date of P. v. mellonae is 10 May. Both of these estimates for the mean bïrth date of P. v.

mellonae are substantiaily earlier than for other harbour sed populations at a similar

latitude (for example, June 10 in northem Labrador, Iune 27 at Schleswig/Holstein,

Gennany, and Jime 29 at Port Molk, AK) (Temte et al. 1991). If Temte (19914 199 1b;

1994) was correct in his belief that biah timing in harbour seals has a heritable

component and may, îherefore, hction to create reproductive barriers between adjacent

populations, my calculations would fiirtber suggest that P. v. mellonae is reproductively

isolated fkom P- W. concolor.

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Condylohasai Length (CL)

Length of Upper 'ïoothw (LUT)

Mastoid Widîh

Length of Mandiile (LM)

MandiMe Height at Coraooid @tlHC)

Length of Lower Tooümw (LLT)

Depth of MandiMe Behind Ml @MM)

Total h g t h of Nasals 0 Length of Maxillo.Ftantd Suitne to Anterior End of Nasal (MAN)

Greatest Width of Extemal Nares CNEN)

Width of Snouî at Canines (WSC)

Least Interorbital Width 0 Greatest h g t h of PM2 (GLP)

Palaiai Widîh at Ml (PWM)

Wichh of Bulla (WB)

Length of W a (LB)

Condylar Width (CW2)

Height of Foramen Mignum 0

Width of Ehh fimm TÈp of Audaory 0.265 Proces to Anteria Edge of Carat-d Foramen (AAC)

Length of Nasal m Caitia With Frontai, -0.848 Relative to Their Overail Length (NF01

Set of Premolar Teeth (Sm 0,394

Variance 59.4% 24.2%

Table 22. A postenon' classZEications based on disaiminant scores for five P.

vitdina subspecies, P. I q h , and P ura hikpida.

% correctly

Groups PVC PVM PVR PVS PW PL PH classified

%

Corredy 88 83 80 78 92 91 98 100

classified

Table 2.3. Meaas o f a d i (logf,,-transfmed and C-scored) aaniai charade

rneasurements arranged acconi8ig to shidy group (following Amam a d Miyaraki 1992).

Wxthin rows, means that are lrmotated wial the same slipaYaipC leüers are signincantly

different (P < 0.01).

Char- PVC PVM PVR PVS P W PL PH acter

CL

LUT

MW

CWI

ZW

HC

LM

MHC

LLT

DMM

LN

MAN

WEN

WSC

LIW

GLP

PWM

WB

LB

cw2 WFM

HFM

SAN

GLJ

AAC -0,118- 4-244 O.O1lC 0262AD -0.309- 0,123E O22Off

NF0 0257- 0.245 -0318"" 0 . 6 7 F O,17SEF -0.6lp

PTN 0.223- 1 . 0 2 y - 0 . ~ 6 2 ~ -0-937- 1 . 1 6 1 ~ ~ - 0 . 9 3 ~ ~ 027gGH1

PUJ -0.302- -0 .931~ -0.38F -1-042GH -0,189G 1. lmçM 0.98780CG

SPT 0.343- -0.982 -O.SSOAC -0,473- 0.492- -0.292E 0219

Fig. 2.1. Range of the mur seai (Ph- vituIina L.).

Fig. 2.2. Relative di f fe~e~~ces between the grwp merms of craaiai measurements of 5

subspecies ofbatbour seai (Phoca viiulina concolor; P. W. mellonae; P P. v. vitulim; i? v.

richardsi and P. v. stejnegen3, the spotted seai (P. largha) and the ringed seal (Puno

hispida) based on plots of first, second and third caaonical variates.

c 0

O v

F z 0 O O hispida

Appendk 1

Sources and acquisition nwnbers of harbour, spotted and ringed sed -a used in these

analyses.

American Museum of Naîural History

Phoca vitulina concoIor= 100,80201,144651, 164600,232379,232390,232406,

232414,232415,232420,232424,232425,232427,232431,232438,232440,232441,

232443,232449

P. Zmgha: 18169

P u a hispida: 19308,193 10

Arctic Biologicai Station

P. W. meIIorne: LM-89-0 1, LM-89-02, LM-90-0 1, LM-90-02

Canadian Museum of Nature

P. v. concolor: A20685,198429-SIV1,10358,12502,52622,52630,52631,52639,

52640,52641,52642,52643,52644,52645,52650,52651,52652,52653,52654,52658,

52659,52660,52661

P. v. richarAi: 2597,52662,52663

Carnegie Museum

P. v. concoloc 6671,17848,17849,18748

P. v. mellonue: 15215

Grand Manan Whaie and Seabird Research Station

P, v. concolor= GM-9 1-06, PV-92-02, PV-92-03

National Museum of Natural History

P. v, concolo~or, 180285,188823,188826,504298,504299

P. v. richarhi: 49550,81516,81518,81520, 146429,14643 1, 146435,146436,146437,

154015,154016,219866,219868,219873,219874,219876,2 19879,245914,2459l5,

253041,253042,253235,250712,250713,273532,504842

P. v. stejnegeri: 213 10,213 12,290653,290654,290655

P, v. virulina: 238153,238154

Zoologisk Museum

P. v, concolor= CN79, CN82, CN83, CN84, CN85, a 1 9 6

P. v- vitulinu: CN159, CN253, CN321, CN75 1, CN847, CNlO85, CNl 11 1, CNll42,

CNllM, CN1149, CNl155

University of AlasLa Museum, Fairbanks

P. v. n'chardi: 3409,3704,11475, 11627,11727,11740,11742, 11746,11750, 11751,

11752, 11758

P. v. stejngen*: 11718, 11720, 11721,11723,L1724, 11728, 11729, 19127, 19128,

19130,19134, 19135, 19139,19141,19143,19146,19149,19150,19151,19152,19155,

19156,19183, 19184,19185,19186,19187,19190, 19191,19192,19199,19202,19203,

19205, 19206,19207

P. Iargha: Il458.ll459, 11593,11594,11599,11602,11605, 11613,11634, 11797,

11799, 11801,11804,11805,11891,16606,16607, 18602,18604, 18605, 18610, 18612,

186 13.18617, 18619,18620,1862 1,18622,18623, 18624,18625,18641,18642,18643,

18659,18666, 18675,18680,18683,18686

Pusa hispida: 2088,7183,L1565,11566,11568,11575,11576, 11578,L1579, 11580,

1 158 1,11584, 1 tS85,11586,11587,ll~88, llS9O,ll892,Ll895, 1 l896,ll897,ll898.

11899,16599,16600,19055,19058,19063,19068, 19080,19084, 19085, 19095, 190%,

19098,19100, 19102,19108,19118

University of Amsterdam Museum

P. v, vitulim: 11.116, 11.1 19, 11.742, 12.647, 12.919, 15.146, 15.987, 18,454, 18.455,

19.397, 19.398,23.713,23.716,23.726, 23.735.23.737, 2O90, 2593.4290, 8276, 8723

P. largha: 24.142

Pusu hispida: 17.184,21.065,24.348

CHAPTER 3: GENETIC ANALYSIS OF WESTERN ATLANTIC,

Phoco vitrrlha concolor DeKay 1842, AND LACS DES LOUPS MARINS,

P, v, mel lo~e Doutt 1942, HARBOUR SEALS USING MITOCHONDNAL DNA

Does the layman understand what the sight of a piece of stlange sealski. means? 1t means the clue to possible discovery o f a new large mammal, and the discovery of any new large mammal is now so rare that even the possibiiity gïves a scientist the jitkrs (Twomey 1942, p. 4).

Introduction

The original subspecies description of the Lacs des Loups Marins harbour seai (P. v.

mellonue) was made on the b i s of morphological evidence and a presumed

geographical isolation of the population since the retreat of the Laureutide ice sheet

(Doutt 1942). The most prominent critique of the subspecies designation was that of

Mansfield (1967), who noted:

Several seals in Company could well have been the

fomding membem of the population in Upper and Lower

Seal Lakes, their establishment in these two lakes probably

depending on the presence ofrapid sections with open

water in the wiuter and au adeqyate food supply. There is

no evidence of migration ofthese seals, the lakes appearing

to be their permanent homes. In spite of thk, there appears

to be no need to postdate a long isolation by slow

emergence of the lakes fiom the level of the pst-glacial

seas, since access to them at the present t h e is quite

possible @. 256).

Smith and Horonowitsch (1987, p. 8) thought that Mansfiefd's interpretation "hit the

area contaios a resident population occasionaily augmeoted by wanderuig seals entering

from the sea and undertaking d o m ovaland migrationsy' was a reasonable hypothesis.

Mansfield (1967) pointed to Inuit toponyms that seemed to iadicate that seais were once

more wi&ly distnbued throughout the Rzvière aux FeuIlles drainage system (also

corroborateci by Consortium mes Shooner & Associés 1991), and suggested that:

It may well have been possible that before the tum of the

century the waîershed lakes were a favoted summeer resort

of young wsf~ldering harbor seds a few reaching Upper

and Lower Seal Lakes fkom time to time. When the rifle

came the coastal harbor d s were heaviiy exploited,

resulting in virtual isolation of the lake seals (p. 256).

Analyses of mitochondrial DNA, in partidar the control region, have beeu

increasuigiy used to elucidate population sub-structure in a varïety of marine mammal

species (see Hoelzel et al. 1991; Maldonado et al. 1995; Mouchaîy et al. 1995; Rose1 et

al. 1995; Malik et al. 1997). The mitochoudrial DNA of harbour d s (Phoca vil~fim)

was nrst analysed by h a s o n and Johasson (1992). In the* subsequent study of

worldwide mitochondrial DNA diversity in the harbour seal, Stanley et al. (1996) used a

limited sample of harbour seals fkom the western Atlantic (P. v. concolor) and no

sampies fÏom Lacs des Loups Marins (P. v. mellonae). Other analyses of hatbour seal

58

genetic diversity have also excludcd these two subspecies (Burg 1996; Lamont et al.

1996; O'Corry-Crowe and Westlake 1997). The purpose of the preseat study was to

undertake a preliminary investigation ofthe extent to which P. v. aeIIonue is genetically

distinct fiom oceanic harbour seals, partïdarly those h m easfern North Amerka (P. v.

concolor), through the ~e~uencing of a portion of the mitochondrial control region and

the comparhon of these SeQuences to the d t s of Stanley et ai. (1996).

Materials and Methods

Sample Collection

Seventeen samples were sequenaxi for the purposes of the aoalysis (Table 3.1).

Muscle samples were fiozen until use, and blood was taken nom the hind flippers of

captured se& (Chapter 5) in senun separatim hibes and centrifuBed as soon as possible

after collection before king stored in liquid nitrogen.

DNA Analysis

Blood and muscle samples (approximately 0.5 to 1.0 g of muscle tissue) were prepared

in 1 x lysis buffet (4 M urea, 0.2 M NaC1,0.5% n-lauroyl sarcosiw, 10 mM CDTA (1,2-

cyclohexanediamine), 0.1 M Tris-HCI pH 8.0) and ground using a mortar and pestle

cooled in liquid nitrogen. DNA extractions were performed according to Guglich et ai.

(1994).

DNA served as a template for amplification using the polymerase chah reaction

(PCR). Two PCR primers -Tl 15 5'-ATGACCCTGAAGAA(G/A)GAACCAG-3' and

59

WKT283 5'-TACACTGGTmGTMCC-3') were used to a m p e a 520 b.p, product

containing a portion of the tRNA threoniae and probe and part of the control region

(Lamont et al. 1996). The DNA was amplified mder the following reaction conditions:

1 O m M Tris-HCI (pH 8.4), 50 mM potassÏum chloride (KCI), 0.00 1% Triton X-100.2.0

mM magnesium chloride (MgCI2). 0.2 mM dNTP's, 0 2 mM of each primer, 1.5 U of

Taq DNA polymerase (Perkùi-Elmer-Cetus), aad 1 0 ng of template DNA carried out in

a 25 pl volume- AmpIifidoa was pediormed under a temperature regime of: 9S°C for

90 S. 52O C for 60 s, 72°C for 60 s for 1 cycle; 94OC for 30 s, 52" C for 30 s, 72°C for 30

s for 25 cycles; 94OC for 30 s, 52O C for 30 s, 72°C for 7 minutes for 1 cycle. PCR

products were isolated through a L.5% low-melting point agarose gel aud excised and

purifieci using a phenol-chloroform and chlorofonn extraction.

Amplineci product (2 - 300 ng) was used for cycle sequencing according to the

P R I S M ~ ~ Ready Reaction Dye Deoxy Terminator Rotocol (Applied Biosystems hc.,

Foster City, CA). DNA sequences were analyzed by the cornputer program Image

Quantification (Molecuiar Dynamics). The proâuct was sequenced using both primers to

codkm the sequence using both strands.

Analysis of Sequence Variation

Two cornparisons were undden: the £ïrst among the 17 saquenced samples ; the

second berneen the q e n c e d samples and the 34 haplotypes descri'bed by Stanley et al.

(1996) (GenBank accession numbers U363424J36375). For both cornparisons a

published DNA sequence of the -y seal (Haïichoems grypus) was used as the outgroup

60

(sequeme firom h a s o n et al. 1993). DNA h m the 17 sequenced samples were

reduced to 480 b.p. to aügn the sequences in the computer program Genetic Data

Environment Next. 385 b.p. of these sequences were aligned with those of Stanley et al.

(1996). For each of the two commsons, sequence variation was estimatecl using the

computer program Phylip 3 . 5 ~ (Felseosiein 1993). DNA distances ofpiUnvise seq~eace

cornparisons were obtained using Kimura's (1980) 'Wo-parameter" mode1 assumhg a 2-

to-1 transition to transversion substitution in the Phylip program DNAdist, A statistical

analysis using the bootstrap method was performed to gerierate coaf ihce intervals on

the phylogenies. The Phylip prognun Seqboot resampled the original data by replacing

cbaracters, in this case nucleotides, and estimateci the variation among the repliCates

using the DNAdist program. Neighbor-joining trees were caiculated for the distance

measures of each replicate using the pro- Neighbor. A consensus tree - a majorîty

d e tree that is wnstnicted acwrding to the groups or clusters that are gewnited most

often in the replicates - was then created fkom bootstrap data in the program Consense to

estimate the confidence interval at each node. A total of 500 bootstrap data bases were

generated to test the phylogenies (Felsemtein 1985). In addition, a pamimony analysis

was undertaken using the DNApam program. A bootstrap of 1ûûû replicates, using the

programs listed above, was applied to the e o n y analysis.

Results

A total of 14 different mitochondrial DNA haplotypes were found amoog the 17

sequenced samples (Table 3.2). Thne of the individuals fkom Lacs des Loups Marins

FM-89-01, LM-94-03, and LM-94-04) SM a haploîype, and two of the hdividuals

l?om Massachusetts shared a haplotype.

Within the 480 bop. examineci, the observed number of painvise différences among

haplotypes mged nom 1 to 19. Conected DNA distances for cornparisons between the

17 sequenced samples ranged hm 0.008 to 0.05 (T'able 3.2). Lacs des Loups hilarins

had four unique haplotypes, three ofwhich were Smilar and grouped together 647% of

the time (Fig. 3.1). The fourth Lacs des Loups Uarins haplotype (LM-90-02) was more

differentiated than the other harbour se& that were sequenced, with a mean of 16 0.6

substitutions as wmpared to a mean of 6.6 * 0.4 substitutions for all 17 samples (Table

3.2).

When compered with the sequences nom Stanley et al. (1996), all P. v. meIIorne and

the majority of the P. v. concolor samples grouped with other harbour seals firom the

western Atlantic (Fig. 3.2). The exceptions were the sample nom New Jersey, which

was identical to a sample h m Churchiii, Manitoba froxn the Stanley et al. (19%) study

and wnich - as it did in the Stanley study - grouped with eastem Atlantic Ocean

haplotypes (Fig. 3.2). In addition, one of the haplotypes fiom the St. Lawrence River

was similar to one ofthe haplotypes described in Stanley et al. (1996) and they grouped

together close to the eastem Atlantic samples.

The most parsimouious tree generated grouped all of the Lacs des Loups Marins

samples together 47.2% of the time.

62

Discussion

Doutt (1942) summa"sed his hypothesis concerning P. v. meiIonae in this way:

it would seem that we have a large species, with few

generatiom per century; but it has been thrust into a new

environment which would cause it to change more rapidly

than wodd otherwZse be the case. T'us, these two factors

tend to cance1 one snother ...Thus it seems probable that

about 5,500 years, phis or minus 2,500 years, has been the

time raquirrd to produce a new subspecies under the

conditions set forth above (p. 78).

In response, Mansfield (1967) indicaîed that there is evidence of harbour seals having

once been more widely distriiuted in the Arctic in localiseci populations, but that they

were eliminated by over-hunting. He noted that:

The localized distribution of the barbot sed In the arctic

makes its fiiture somewhat precafious. Already in Ungava

Bay, swthern BatFn Island, and southem Southampton

Island it has been elimiuated fkom some local territories,

for the Eskimos often know exactly where to h d this

species in m e r @. 252).

Though there are known to still be d 1 harbow seal populations in estuaries in Ungava

Bay, western Hudson Bay and the Iower Churchill River, Manitoba (S. h e s ,

Department of Fisheries and Oceans, pers. comm.), the over-hunting that Mansfield

(1967) referred to has obviously contributed to the M e r isolation of P. v. meilonae

63

fiom aeighbouring oceanic harbour seals. hnsfïeld's argument was that the Lacs des

Loups Marins popdation is sùnply a remnant of harbow sed diversity that was once

widespread on the Ungava peninsuIa

Given that the mitochondrial control region is a neutral marker (Hoeh1 et al. 199 1)

and would exhibit little, if any. change over the 7,300 year perïod since the retreat of the

Laurentide ice sheet (AlIard and Séguin 1985), it is not possi'ble to f d a t e any

conclusions regarding the t h e of isolation of P. v. mello~e fioul the present genetic

data. Those selectable morphological and behaviod characteristics that seem to have

changed in the population (Chapter 2) do, however, lend weight to Doutt's (1942)

contention that the popdation has been isolated for some tirne. It is unlürely that a shift

in reproductive seasonality, such as P. v. mellonae seems to exhi'bit (Chapter 2). would

be possible in the centmy or so that Mansfield (1967) postulated as the t h e perîod over

which other harbour seals in nodem Québec were eliminated,

Consistent with the arguments ofboth Doutt (1942) and UansfieId (1967), the Lacs des

Loups Marins seals appeared to be most closely related to harbour seals f?om the western

Atlantic Ocean (Figs. 3.1 and 3.2). Though the four uaique haplotypes and particular

difference exhiited by one individual fkom Lacs des Loups Marins sample are

intrïguing, and potentiaiiy indicative of a founder effect in the population, it is difficdt to

formulate any conclusions regarding the genetic relationship of P. v. mellonae to harbour

seals that used to exist in Arctic Canada without M e r andysis of historic samples. The

samples that were included in the analysis fiom the closest geographic locations to Lacs

64

des Loups Marins were the one animal h m Churchill, Uanitoba and two fkom

Greenland The Churchill animal was more closely related to animals fiom the New

Jersey coast and the eastem Atlantic (Table 32; Figs 3.1 and 3.2). One haplotype fkom

Greenland was most closely related to haplotypes h m Sable Island, N.S., Miquelon and

the St Lawrence. The second Greeniand haplotype, though it grouped closely with Lacs

des Loups Marins 2, was not supported by a very large bootstrap value (38.8%) @ig.

3 - 1)-

The number ofpainvise substitutions in the combineci sample of 17 Seqllences was very

similar to that observecl by Stanley et ai. (1996) (1 to 23 substitutions in 435 b.p.).

LnterestingIy, Burg (1996) and Lamont et al. (1996) fotmd higher numbers of pairwise

substitutions in theu sampling of Pacific animals ( average of 2.6% * 0.29%. and 1 to 16

among 320 b-p. respectively). Lamont et al. (1996) concluded thaî this relatively large

amount of genetic variation in Pacific harbour seals was Wrely due to the lack of a severe

population bottleneck - as has been experienced by other pinniped populations due to

over-exploitation - at some tirne in the past. In addition, Lamont et al. (1996) concluded

that there is limited gene fîow among Pacinc harbour seais dong the coasts of

Washington, Oregon, and California and limiteci, if any. gene flow between Puget Sound

and the Pacinc coast of Washington. In reference to the work of Temte et al. (1991).

Lamont et al. (1996) concluded that the timing of harbour seal birüis in Puget Sound is

most iikely genetically regulated, and possibly follows the schedule of harbour seals off

British Columbia Burg (1996) aiso reporteci significant stnrcturing of the harbour seal

population in B.C. and Alaska. From her analysis, she discemed three significantly

65

separate groups, suggesting that the Pacifie Ocean was colonised by harbour seais twice.

One m a t d heage hvaded the Pacifie approxiniately 670,000 years ago and is oow

restricted to southem Vancouver Island, Puget Sound and Iapaa A second invasion

occurred approximately 380,000 years ago. This group of animals colonwd Japan and

Aiaska and a small group of f d e s moved south firom northem B.C. and Alaska to

wloniw southem B-C.

The conclusion of both Burg (1996) and Lamont a al. (1996) is that there are

reproductively isolated groups ofharbour seais within P. W. richar&i- This conclusion

based on genetic evidence coafirms previous o h a t i o n s of harbour seals' extreme site

fidelity (Boulva and McLaren 1979; Bigg 198 1; Keliy 198 1). These conclusions are

consistent with those of O'Corry-Crowe et al. (1997), who found - through an analysis

of the mitochondriai DNA control region - that P. v. stejnegeri and P. v. richarthi do not

occur as genetidy distinct clades. It codd very weli be that, with fÙrther analysis of P.

v. concolor genotypes, similai sub-structuring cornes to light.

References

Ailarà, M., and M.K. Séguin. 1985. La dglaciation d'me partie du versant hudsoaien

québécois: bassins des rivières Nastapoca, SheIdrake et A L'eau Claire.

Géographie physique et Quatemaire 39: 13 :M.

h a s o n , U'., and E. lohnsson. 1992. The oomplete mitochoadna1 DNA sequeme of the

harbor seal, Phoca v i t u l i ~ ~ ~ ~ J, Mol. Evol. 34493-505.

hx~ason, u., A- Guliberg, E. Jobnsson, and C. Ledje. 1993. The nucleotide Sequence of

the mitochondrial DNA molde of the grey seai, Holchoerics grypus, and a

cornparison with mitochondrial sequences of other true seals. J. Mol. Evol.

37:323-330.

Bigg, U A . 198 1. Harbour seal - Phoco vitdina Linnaeus, 1758 and P. lmgha Pallas,

18 1 1. Pages 1-27 in Handbook of marine mammals Vol. 2: Seals. Edited by

S.H. Ridgway and RJ. Harrison. Acaâemic Press, London.

Boulva, J., and I.A. McLaren. 1979. Biology of the harbor seal, Phoca vihrIi~, in

eastern Canada. Bull. Fish. Res, Board Canada 2 0 .

Bmg, TM- 1996. Geuetic analysis of Eastern Pacific harbour seals (Phoca v ï ~ I i n a

richar&i) fnnn British Columbia and parts of Alaska using mitochondriai DNA

and microsateliites. M.Sc. Thesis, Department of Zwlogy. University of British

Columbia, Vancouver, BC-

Consortium Gilles Shwner & Associés3 SOMER, anri E n h ~ e r n e n t nlimité Inc. 1991.

Complexe GrandeBaIeine, Avant-projet Phase 2, Bilan des connaissances sur le

phoque d'eau douce. Vice-présidence Envkomement, Hydro-Québec.

DeKay, LE. 1842. Naîural history of New YorL, Part 1. Zoology. D. Appleton and Co.

and Wdey and Putnam, New York W.

Doutt, J. K. 1942. A review o f the genus Phoca. AM. Carnegie Mus. 29:6 1- 125.

Felsenstein, 1.1985. Confidence limits on phylogenetics: an approach ushg the

bootstrap. Evolution 39:783-791.

Felsenstein, 1. 1993. PHYLIP Manual Version 3%. University Herbarium, Uaiversity of

California, Berkeley CA.

Guglich, E.A., P.J. Wilson and B.N. White. 1994. Foremic application of repetitive

DNA markers to the species identification of animal tissues. Joumai of Forensic

Sciences 39:353-36 1.

Hoelzel, AIR, I M Hançock, and GA. Dover. 1991. Evolution of the cetacean

mitochoadrial D-Ioop region. Mol. Biol. Evol. 8:475-493.

Kelly, B.P. 198 1. Pelage polymorphism in Pacific harbor seals. Cm. J. Zod. 59: 1212-

1219-

Kimura, M. 1980. A simple mode1 for estimating evolutioaafy rates of base substitutions

through comparative studies ofnucleotide aquences. Journal o f Molecular

Evoluîion 16: 1 1 1- 120-

Lamont, MM., J.T. Vida, J.T. Harvey, S. Jemes, R Brown, H.H. Hubet, R DeLong,

and W K Thomas- 1996. Genetic substnrcture of the Pacific harbor seal (Phocu

virulina n'char&) off Washington, Oregon, and California Mar. Mamm. Sci.

12:402413.

Maldonado, LE., F. O. Dada, B.S. Stewart, E. Geffen, and RK. Wayne. 1995.

Intraspeciiïc genetic differentiation in California sea lions (Zolophus

c u I z ~ o m i ~ ) h m southern California and the Gulf of California. Mar. Mamm.

Sci. 1 1 :46-58.

Malik, S., P.J. Wilson, RJ. Smith, D . U Lavigne, and B.N. White. 1997. Pianiped

penises in trade: a molecular-genetic investigation. Conserv. Biol. 1 1 : 1365-

1374.

Mansfield, A.W. 1967. Distrifiution of the harbor seal, Phoca vituiim Linnaeus, in

Canadianadian arctk waters. J. M d . 48:249-257,

Mouchaty. S., I.A. Cook and G.F. Shields. 1995. Phylogenetic d y s i s of noahem hair

seah based on nucleotide sequences of the mit~chond~al cytochrome b gene. J.

Mammal- 76: 1 178- 1 185-

O'Corry-Crowe, GM., and EL. Wdake. 1997. Molecular investigations of spotted

seais (Phoca Iargha) aud harbor seals (P. vi tuI i~)3 and theu relationship in areas

of sympatq~ Pages 29 1-304 in Molecular genetics of marine mamrnais. Special

Publication Number 3. Edited by A.E D h n , S.J. Chivers and W.F. Perrin. The

Society for Marine Mammdogy, Lawrence, KS.

Rosel, P.E., AB. D b n , and M.G. H a y g d 1995. Variability of the mitochondrial

wntrol region in populations of the harbout prpoise, Phocoena phocoena, on

interoceanic and regional scaies. Can. J. Fish. Aquat. Sci. 52: 12 10-1 2 19.

Smith, T.G., and G. Horonowitsch. 1987. Harbour seals in the Lacs des Loups Marins

and eastem Hudson Bay drainage. Can. Tech. Rep. Fish. Aquat. Sci. 1539.

Stanley, H.F.. S. Casey, LM. Cunahan, S. Goodman, J. Harwood, and RK. Wayne.

1996. Worldwide patterns of mitochondrial DNA differentiation in the harbor

seal (Phoca vituIina). Mol. Biol. EvoL 13:368-3 82.

70

Temte, J.L., MA. Bigg, and 0 Wiig. 199 1. Cliws revisited: The timing of pupping in

the harbour seal (Phoca vitdina). I. Zool., Lond 224:617-632.

Twomey, A C 1942. Needle to the North: The story of an expedition to Ungava and

the Belchet Islands. Herbert JenkIns Ltd., London.

-

Sample Number Tissue Collection Location CoUection Date

LM-89-01 muscle Lacs des Loups Marins 89-08

muscle

blwd

blood

blood

b l d

muscle

muscle

muscle

muscle

muscle

muscle

musde

muscle

blood

muscle

muscle

Lacs des Loups Marins

Lacs des Loups Marios

LacsdesLoupshilarias

Lacs des Loups Marllis

LacsdesLoupsUarins

S. Aappilattoq, G r d m d

Anorluitsoq, Greeniand

Massachusetts

Massachusetts

Massachusetts

Massachusetts

Grand Manan, N.B.

Nova Scotia

New Jersey

Gaspé, Qué.

S t Lawrence River, Qué.

Fig. 3.1. Consensus tree of500 bootstrap neighbour-joining distance trees for 480 b.p. of

the mitochondrial control region for 6 Phoca vituIina mellonae and 11 P. v. concolor.

The grey seal (Halehoma g r j p ) is the outgroup. The number located at the nodes

indicates the percentage of bootstrap trees that contain this pattern.

I Lacs des Loups hbrins (LM) 3

64.7% -t Lacs des Loups Matins (LM) 4

Lacs des Loups Marins (LM) 1 P

I Massachusetts (MA) 1

I r Grand Manan (GM) 1

- St. Lawrence (SL) 2

Greenland (GR) 2

Nova Scotia (NS) 1

-

1

- St. Lawrence (SL) 1

LI

Lacs des Loups Marins (LM) 2

- Massachuseüs (MA) 2

Greenland (GR) 1

Massachusetts (MA) 3

t- New Jersey (NJ) 1

Fig. 3.2. Consensus tree of 500 bootstrap neighbom-joinmg distance trees for 385 b-p. of

the mitochondrial control region for 6 Phoca vi îzdi~ meflon<e, 1 1 P. v. concolor, and 34

harbour seal haplotypes reporteci by Stanley et al. (1996). The grey seal (Hoichhwrur

gr)(pilr) is the outgroup. The number located at the nodes indicates the percentage of

bootstrap trees that contain this pattein.

1 Gc ChurcW Man, New Jersey 1

W Baiîic W Baitic

ScotlaaàiN hlaud

- I œ b d Sable k

-

C MiqueIon/Sable Is. Nova Scoiia 1 - Grand Mauan 1

L

- Iceland/N Sea/W Baltk E Baltic

Commander 1s. Commander Is,

a

I rœ~d/Scotiilll~ Mtic ' - W Baltic

r J w Bait-c E Baltic

A Gwtïc

CHAPTER 4: DISTINGWSHING BETWEEN POPULATIONS OF FRESH

AND SALWATER HARBOUR SEALS, Phoca vituZi~ L., USING

STABLE-ISOTOPE RATIOS AND FATTY ACID PROFTLES

No one knows how many years rnight have passed . .. had not the search for bagea, the unknown seal of Seal Lakes of Ungava, received a sharp and sudden impetus. During his 1935 summer work at Great Whaie River, [Doutt] ... noticed the s e a l s h haady-bag which hung h m an hdian's shoulder. The bag seemed to be made fiom the skin of a hair seal, but not the skin of nefchek, comrnon hair seal of salt water, or any hair seai known to science (Twomey 1942, p. 10)

Introduction

W e it is common for harbour seals (Phoca vituIina) to feed occasionaüy in lakes and

rivers throughout their world-wide distribution (Reeves et al. 1992), the apparent

residency of harbour seals in Lacs des Loups Marins is unique (Smith et al. 1994; Smith

1997). One way to investigate the degree of the Lacs des Loups Marias seal's (P. v.

rnellonae) use of fieshwater, and the extent to which this behaviour distinguishes the

population fiom marine harbour seals, is through a study of feeding habits.

The diet of P. v. mellonue was examined using biochemical techniques that have

proven effective in other, somewhat similar, investigations: stable-carbon and aitrogen

isotope ratios in hair (reviewed by Peterson and Fry 1987; see also Lajthe and Michener

1994), and chemometry of fatty acid profiles in the tissue Lipids (Grahl-Nielsen et ai.

1993; Kakela et al. 1993).

The stable-isotope approach is based on the fact that stable isotope ratios of carbon

( i 3 ~ / ' 2 ~ ) and nitrogen (1%/14N) in animai tissue are ultimately related to diet (DeNiro

and Epstein 1978, 1981). Stable-carbon isotope ratios ofprimary producers in

fieshwater food webs are typically l e s enriched in "C than those of marine food webs

and this diîférence is passed on to higher-order consumers (Chisholm et al. 1982; Fry and

Sherr 1988; Hobson 1990; Hobson and Sealy 1991; Ramsay and Hobsoa 1991; Walker

and Macko 1999). Thus, seals from marine food webs should show relatively more "C

in their tissues than those fiom freshwater food webs. Any enrichment of 13c in P. v.

melZonue tissue fiom expected values for fkeshwater endpoints would provide evidence

of marine input to the diet. When used in conjunction with "C measurements, stable-

nitrogen isotope analysis can provide additional information about the diet of seals as

well as their use of marine and fkeshwater food webs (Hobson and Welch 1992, 1995).

Stable-nitrogen isotope ratios in consumer tissues tend to be enriched by about 3%0

compared to dietary values and this step-wïse enrichment in '% in food webs has been

used to estimate the trophic levels of animals, including marine mammals (Rau 1982;

Minagawa and Wada 198+ Wada et al. 1987; Fry 1988; Hobson and Welch 1992, 1995).

Tissue fatty acid composition is also related to an animal's diet (Hilditch and Williams

1964). Freshwater and marine food webs are characterïsed by different levels of specific

fatty acids, particularly long-chah polyunsaturated fatty acids (PUFAs) @ilditch and

Williams 1964). Pnmary producers in fkeshwater envkonments have significantly Iowa

n-3/n-6 ratios than their marine cornterparts (Sargent 1976) rdting in freshwater fishes

having lower n-3h-6 ratios, lower levels o fCo and C, umatwated fatty acids, and

higher levels of C,, PUFA than marine fishes (Henderson and Tocher 1987). In

p~hnipeds, blubber Ïs the most important site of lipid storage (Pond 1978) and the

chemical composition of this tissue is thought to reflect that of the animal's diet (e.g.

Iverson et ai. 1995, 1997). Thus, the f q acid composition of the blubber of marine

seals which presumably prey exclusively on saltwater fisb, shouid mirror the chemical

profile detailed above. Similady, a fkeshwater signature in the fatty acid composition of

P. v. mellonae's bluôber would indicate that it feeds primarily on fieshwater fi& species.

1 analysed the tissues of P. v. m e I Z o ~ e and two marine populations of harbour seals (P.

v. richarhi, eastem Pacifîc Ocean; P. v. concolor, western Atlantic Ocean and eastem

Canadian Arctic) to test the hypothesis that P. v. mellonne feeds primarily in a fkeshwater

habitat, and is disthguished fiom its oceanic cornterparts by the fieshwater signature in

its tissue stable-carbon and nitrogen isotope ratios and fatty acid profiles (Smith et al.

1996). 1 also tested whether stable isotope ratios in P. v. mellonae were distinguishable

fiom those in a group of lacustrine P. W. concolor which, befause of their proximity to

the ocean, probably feed in both freshwater and marine habitats (Smith et al. 1996).

Materials and Methods

Sample Collection

Stable-carbon and nitrogen isotope values were examinai in three groups of seals: P.

v. concolor fiom Kasegalik Lake, Belcher Islands, N.W.T., P. v. concolor fiom the

80

Northwest Atlantic Ocean, and P. v. meIIome- Fatty acid values were examined for the

latter two of these groups, in addition to P. v. rr'charciIri nom the eastern Pacinc Ocean.

Most of the haïr samples used for stable isotope analyses were gamered h m &ed

pelts in the folIowïng collections: the Carnegie Museum, Pittsburgh, PA (Table 4.1; CM

accession numbers); the Canadian Museum of Nature, Ottawa, Ont. (CMN accession

numbers); and the Greenland Environmental Research Insiitute, Copenhagen, Denmark

(PV-Gl). The remainder of the samples that were used in both d y s e s was fiesWfiozen

tissue collected fiom animals that were shot (Department of Fisheries and ûceans

Canada, Nanaimo Research Station, sarnples PV-11 through -26; University of Guelph,

samples LM-94-02 and LM-95-04; Greenland Environmental Research White, sampfes

PV-1527, -1612 and -1618), found dead (Grand Manan Whale and Seabird Research

Station, North Head, N.B., samples PV-92-01 tlirough -05 and PV-93-01), or drowned

incidentally in gillnets (Department of Fisheries and Oceans Canada, Institut Maurice-

Lamontagne, samples LM-89-0 1 and -02).

Hair, blood and blubber biopsy samples fiom LM-94-0 1, LM-94-03 through -05, LM-

95-01 through -03, LM-95-05 and 4, and LM-96-01 through -05 were collected fiom

live animals that were captured and released in Lacs des Loups Marins in August and

September of 1994, 1995 and 1996 (Chapter 5). Blood was taken fkm the hind flippers

in s ~ u m separatioa tubes and cenfrifùged as soon as possible after collection before

being stored Biopsies were taken nom the animals' side near the pelvis; the açea was

first fiozen with a small injection of lidocaine hydrochloride O(ylocaine, Astra Pharma

81

Inc., Mississauga, Ont) before the application of a 6 mm diameter biopsy punch (Acu-

Punch, Acuderm hc., Ft- Lauderdale, FL).

The fish fauna of Lacs des Loups Marins was sampled in 1995 and 1996 using a

scientific sampling mononlament gillnet (Iength of approximately 42.5m; divided into 7

panels of different sized mesh, ranging nom 2.5 cm to approximately 10 cm in 1.25 cm

increments). The fish were sampled both to gather a rough idea of the composition of

the lake's fish community, and to obtain samples of potential prey or foodweb

components for stable isotope analysis.

S table-isotope Analyses

The collected hair samples were washed in detergent using an ultrasonic cleaner and

then rinsed repeatedly in distilled water before being dried in an oven at 60°c. Samples

were then soaked in ether, rinsed, and a i . dried before king ground to a powder in a

dental amalgam baii mill (Wig-L-Bug). Fish muscle tissue was freeze-dried and lipids

extracted using repeated rimes with 2: 1 chloroform:methenol. Samples were then air

dried and powdered, Samples for 1 3 ~ / 1 2 ~ analysis were loaded into tin cups and

combusted in a CN elemental analyser that was part of the Europa Robo-Rep coatinuous

flow mass spectrometer used for 13c analysis of CO2. Samples for '% analysis were

loaded into quartz tubes with CuO, elemental Cu, Ag wire, and Cao, evacuated and then

sealed using a propane torch. These samples were then combusted at 850°C for 2 hours

before king allowed to cool slowly. The resultant N2 gas was introduced directly into a

SIRA-12 isotope ratio mass spectmmeter. Stable-isotope ratios wexe expresed ushg the

following notation:

6x = (%mpIe/Rsrandard - 1) X 1

where X = or %, R = '3~/'2~ or '%/'% and the conventioaal standards were

PeeDee Belemnite (PDB) carbonate and atrnospheric (AIR) nitrogen for carbon and

nitrogen samples, respectively. Analytical error for both 613c and 6% analyses was

estimated at I0.3%0. ANOVAs (a = 0 .01 ) were used to compare S"C and 6'%Ivalues

among the groups of seals.

Fatty Acid Analyses

hed i a t e ly after biopsy, blubber samples (approximately 0.5 g each) fkom LM-94-01

to -05 were placed in 2 mL of 2: 1 chloroform:methanoI (0 with 0.005% butylated

hydroxytoluene, in Kimax test tubes with teflon caps. Samples were kept at -20°C until

M e r analysis, at which tirne an additional 7 ml of the C:M mixture was added to each

tube. Blubber samples fiom the other seals (Table 4.1) were taken fkom îÏesh carcasses

and fiozen until fkther analysis (also at -20°C). Approximately 0.5 g of each of these

samples was placed in 9 mL of the C:M solution in Kimax tubes with Teflon caps. Al1

sarnples were mashed manually with a glas homogeniser until thin and transparent, then

vortexed for 20 seconds and ailowed to soak for 24 hours. Samples (total iipid) were

extracted according to a modifieci Folch procedure (Folch et al. 1957; see Iverson 1988).

Fatty acids of total lipïds were converted to methyl esters by placùig 50 mg of üpid

with 1 mi, hexane and 1 mL 10% BFj in methanol (samples LM-94-01 to -OS), or by

piacing 50 mg ofüpid with 1 mL hexane and 1 mL 1PA BF3 in butanol (all other

çamples) in Kimax tubes, which were then incubated at 100°C for 1 hour and ailowed to

cool to room temperature. Deioaised H z 0 (3 mL) was added, and the sample was

vortexed and centrifugeci at 480 x g for 5 min. The hexane layer was removed, washed

with 5 mL deionised H20, and dried with sodium sulphate. Samples were flushed with

nitrogen and capped tightly.

Fatty acid composition was deteRnined using temperature-programmed capillary gas-

liquid chromatography as desmid in Iverson et al. (1995) (methyl esters) and in

Koopman (1994) (butyl esters). Fatty acids were identifïed ushg chromatograms of

hown standard rcixtiaes (Nu Check Rep, Elysian, MN) and fiom samples earlïer

identified using silver nitrate chromatography (Iverson 1988). Ideutified fatty acid

components were converted to a mass percentage of the total array of fatty acids plus

h o w n compounds. The small sample size restrïcted the number of statistical tests that

could be made; thus only individual fatty acids and fatty acid f d e s that have revealed

habitat ciifferences in other species (e-g. see Henderson and Tocher 1987; m e l a et al.

1993) were compared between groups of seals: 18:2n-6, 18:3nw3,20:ln-9, 20:4n-6,

22: ln- 1 1, total C,, PUFA, total saturates, total monounsaturates, total PUFA and the n-

31x1-6 ratio. The three subpopuiations of seals were wmpared using ANOVA on tanked

data (equivalent to a Kniskal-Wallis non-parametric test) with a = 0.05.

Results

S table-isotope Analyses

The three sampled populations exhiibited diffefent 13c content (ANOVA, FlW =

142.6, p4 M O O 1): P. v. mellonue showed the most negative mean 613c value; the

Kasegalik Lake sample of P. v. concoIor showed an intermediate value; and the marine

population of P. v. concolor showed the most positive value (Table 4.2, Fig. 4.1).

Applying the diet-hair 6I3c htionation value for seds of +2.8%0 (Hobson et al. 1996),

mean dietary 613c values for these three populations of seals were cdculated as -25.9,

-2 1 -9, and -1 8 . s for the P. v. melIonae, Kasegaiik Lake, and marine P. v- concolor

samples, respectively. Senmi 613c values for P. v. inelIonue were very similar to the

values derived fiom the analysis ofhair samples (mean of-27.2% after applyhg the

fkactionation value).

Stable-nitrogen isotope values of seds fiom the two fieshwater locations were les

variable than those fiom the broader marine sample (Table 4.2, Fig. 4.1). Using the seai

diet-hair 6% fractionation value of +3.% (Hobson et al. 1996), mean dietary 6%

values of the P. v. mellonae, Kasegalik Lake, and marine P. v. concolor samples were

9.9, 10.6, and 13.2%0, respectively. The mean senmi 6% value for P. v. meMonue was

identical to the value derived fkom hair (9.9%).

One fieshwater seal that was marked in 1994 (LM-94-01) was subsequently recaphned

in 1995 (LM-95-01). To avoid analyshg the same animal twice, the means of this

animal's 1994 and 1995 613c and 6% values (see Fig. 4.1) were used for the purposes

of the ANOVA and the calculation of mean 613c and 6 '% values for P. v. melIonaee

The fish species collecteci included- longnose sucker (Catostomus caîostomur), cisco

(Coregonus artedii), lake whitefish (C. clupe~omik), d p i n (Conus sp.), lake chub

(Couesius plumbeus). threespine stickleback (Gasterosteus aculea~cs), round whitefish

(Prosopium cylindraceum), nlnespine stickleback (Pmgitius pimgirus), brook char

(Sabelinus fonti~&), and lake char (S. rtamuycllsh) (Table 4.3)- Stable-carbon and

nïtrogen isotope ratios for the sampled Lacs des Loups Marins fish are also presented in

Table 4.3,

Fatty acid Analyses

The blubber of P. W. nteIIonae coatained sipnincantly higher levels of l8:2nd, 18:3n-

3, 20:4n-6, C,, PUFA and total PUFA than did the blubber of either P. v. concolor and P-

v. richardri. Phoca v. mellonae also had sisnificantly lower levels of 20: 10-9 and 22: ln-

1 1 and a lower n-3/n-6 ratio than the two groups of marine seafs (Table 4.4). The most

obvious clifference between the fieshwater and marine seals examineci here was the large

proportion of n-6 faîty acids in P. v. nzellonae's blubber. Levels of total n-3 fatty acids

in these animals were also higher than both marine seal groups, but were closer to the

range of vaiues observed for marine seals than were the levels of n-6 f e acids Pig.

4.2).

There were some signincant dinerences between the fatty acid composition of the

blubber of the two groups of marine d s : P. v. concolor (Northwest Atlantic) had

higher levels of 18:2nd, 20: ln-9,22:ln-11, C,, PUFA and a larger II-3/n-6 ratio in their

blubber than did P. v. richardi (eastern Pacifïc).

Discussion

Results nom the stable-isotope and fatty acid anaIyses both indicated that the diet of

Lacs des Loups M&im seais is of fkhwater orïgin. Stable-carbon isotope malysis of

seal hair allowed the successful segregation of the three populations of seals fkom

fkeshwater and marine habitats (Fig. 4.1). As predicted, P. v. meIIonae showed the most

depleted 6% values, wtiich is consistent with a diet derived from fkeshwater prey

during the period when the hair was grown - likely at least haifa year. The results fiom

the analysis of P. v. meIIo~e se-, which are similar to the results fiom the haïr

analysis are consistent with a diet derived fkom fhshwater since the last major feeduig.

The fact that there was no discemie difference between the isotopic signatures of the P.

v. mellonae samples fiom 1936-38, 1989, and 1994-96, and that the marked and

recaptured animal (LM-94-01, LM-95-01) had a 6I3c vaîue that was vimially identical in

both years (-22.9 in 1994, -22.5 in 1995), is fkîher evidence of prolonged fieshwater

feeding.

The Cree of Whapmagoostui have long contended that the Lacs des Loups Marins se

feed in fieshwater and that these animals taste differently compareci to oceanic harbour

87

seals, making them a f a v o d target for hunting (John Petagumshim Sr., p. comm.;

Twomey 1942; Posluns 1993). The Cree also consider that this fieshwater diet is one of

the reasous why the Lacs des Loups Marins seals' pelage is darker and more lustrous

than its dtwater counterpart (John Petagumskum Sr., pers. comm.; Posluns 1993). This

belief that the pelage of the fieshwter animais is ~uanfiably différent has been

remarked upon by a variety of authors over the course of nearly two centuries (Atlonson

18 18; Flaherty 19 18; Doutt 1942; ûraburn 1969), and formed one of the bases for P .v.

melZonue's subspecific description @outt 1942).

Aside nom these anecdotai accormts, few previously published &ta are available on

the diet of the Lacs des Loups Marins seal (Smith et al. 1996; Smith 1997). Power and

Gregoire (1978) conducted a study in which fish fiom Lacs des Loups Marins were

compared with samples caught in nine nearby Ungava peninsula lakes. M e r

determining that brook char (S. fontimIis) was the dominant fish species of Lacs des

Loups Marins, and that lake char (x nanrqmrh) and lake whitefish (C clupeafomik)

populations in the lake were depressed compared to the other lakes, they concluded that

sed predation on the latter two species was responsible for the observed alterations in

Lacs des Loups Marins' fish community. Even though brook char showed evidence of

high mortality in Lacs des Loups Marins, Power and Gregoire hypothesised that their

musual dominance of the lake's fish fauna was attributable to the fact that their

spawnhg occurs in sheltered tri'butary streams, making them inaccessible to potential

seai predation during this vulnerable pe r id Despite this interesting hypothesis, these

researchers provided no direct evidence of the prey preference or feeding habits of the

fkeshwater seals.

The lake whitefish and char populations in Lacs des Loups Marins are cesident

populations (Consortium GilIes Shooner & Associés et al. 199 lb), and limited evidence

fkom the stomach contents of the 4 available seai specimens indicates that P. v.

mellonae's diet consisî~~ in large part, of îhese fish species: CM-152 13, 'îweil-digesîed

pieces of fish" (Doutt 1942, p.71); LM-89-01, "2 otoliths similar to SaIveZirntLF sp.,

numerous mal1 pebbles, and the dorsal spine of Gasterosteus acuIealtls" (Coosortium

Gilles Shooner & Associés et al. 1991a); LM-89-02, "60 otoliths appearing to be those of

the genus Salvelintls, 3 otoliths of Si fontinalis and the pelvic and dorsal spuies of G.

uculeatus" (Consortium Gilles Shooner & Associés et al. 1991a); and LM-94-02,7

otoliths of genus Salvelinus (Smith 1997). However, given that the sL3c values of the

char and whitefish sampled nom Lacs des Loups Marins were even less than those

calculated for the mean seal diet, it is clear that a more extensive carbon isotopic survey

of potential prey of P. v. mellonae is required before any fbrther inferences can be made

about the composition of seal diets within Lacs des Loups Marins.

While few isotopic studies have been conducted on fish in Arctic drainages, Hessiein et

al. (1989, 1991) demonstrated that variation in isotope values among species can indicate

différent sources of feeding both within and outside of local lakes. The 6 " ~ valws for

char in Lacs des Loups Marins (Table 4.3) overlap those found for char (S. alpinus) at

Char Lake, N.W.T. (Hobson and Welch 1995). Whitefish and stickleback values are

similar to those found by Hesslein et al. (199 1) in Travaillant Lake, N.W.T.

Phoco v. concolor h m Northwest Atlantic marine populations showed 6 I 3 c values

that are typical ofmarine mammals measured elsewhere in north-temperate regions

(Ramsay and Hobson 199 1; Hobson and Welch 1992; Hobson et ai. 1997). The

b g a l i k Lake sample was intermediate in 613c signature between the marine and Lacs

des Loups Marins values. Kasegalik Lake is, at some points, l e s than 2 Imi fiom the

ocean, and the Arctic char that fiequent the lake are anadromous (SaLLikiluaq Harnlet

Council 1978). Thus, seals collected nom Kasegalik Lake had access to both fieshwater

and marine-derived carbon, either because of their use, or because of their prey's use, of

both kinds of aquatic habitat.

The stable-nitrogen isotope values for the marine P. v. concolor sample indicated that it

occupied at lest two trophic levels. This is consistent with published stomach content

and scat analyses that have indicated a broad diet consisting of invertebrates as well as

planktivorous and omnivorous fish (Boulva and McLaren 1979; Payne and Selzer 1989).

The P. v. rnellonae and Kasegalik Lake P. v. concolor were similar in 6% values,

suggesting that they occupy similar trophic positions. The estirnateci mean 6% values

of 9.9 to lO.6%0 for the prey of fieshwater seals is in close agreement with those values

found for fish in Lacs des Loups Marins (Table 4.3). Freshwater seals also showed leu

variation in trophic position; probably due to fewer dietary alternatives in high-latitude

lakes compared with Arctic marine systems (e.g. Kling et al. 1992; Hobson and Welch

1992, 1995).

The fatty acid composition of the blubber of P. v. mellonae also indicated that the lipid

in these seals is derived fiom a fieshwata source. The plot of n-3 vs. n-6 fatty acids

(Fig. 4.2) shows that levels of n-6 alone can distinguish the fkeshwater harbour seals

fiom the two marine populations. The pattern of high levels of n-6, Cl, PUFA and low

levels of monounsatiaiexi C, and C, in the Lacs des Loups Marins seals is typical of

the fatty acid composition of organisms in fieshwater environments (Henderson and

Tocher 1 987). Henderson and Tocher (1987) noted that the n-3/n-6 ratio in fieshwater

fish ranges fkom 0.5 to 3.8, while in marine fish the value is typically beîween 4.7 and

14.4. Assuming similar ratios for higher level consumers like d s , this m e r indicates

that P. v. mellonae (ratio = 2.05) is feeding in fkesh water, whereas P. v. concolor (ratio

= 7.65) and P. v. richardri (ratio = 6.3 1) are consuming marine prey.

There are few comparative stuclies of the fatty acid composition of other mammals in

fieshwater environments; however, those tbat do exist have arrived at similar

conclusions. Kakela et al. (1993) examinecl the fatty acid composition of the blubber of

S&aa ringed seals (Phoca hispida suimen&) fiom a fieshwater lake and found that it

contained higher levels of 2OAn-6 (3.02 %) than did the blubber of ringed seah (P. h-

botnicu) fiom the Baltic (0.63 %). However, these authors found the levels of 18:în-6

and the n-3/n-6 ratio in the blubber of Sairnaa seais (4.50 % and 4.58, respectively) to be

the same as those of the Baltic seals (4.43 % and 4.53, respectively), and suggested the

91

presence of a gradient of fatty acids h m kshwater to brackish to marine food webs-

Kâkelâ et aL(1995) found the levels of&n-6 fatty acids in the blubber of fieshwater

ringed seals fkom lakes Saimaa and Ladoga to be twice as high as those in marine ~ g e d

seals fiom the Arctic Ocean. In the present study, the separation behreen marine and

fkeshwater seals on the basis of these fatty acids was even clearer; the blubk of the

marine seals contaiwd less than one-tenth the amount of QI-6 fatty acids observed in

the fieshwater seals (P. v. richardi, 0.18%; P. v- concolor, 0,13%; P- v. meIZonae,

1.95%). The Amazon river dolphin (InUI geoflenrik) is completely riverine and its

blubber is characterised by an extremely low n-3h-6 ratio (0.27). which is primarily the

resdt of high levels of 18:2n-6 (6.61 %) (Ackman et al. 197 1). Inia geoflensik blubber

also contains low levels of 20: 1 (0.52 %) (Achan et al. 1971), similar to those observed

here in P. v. mellonae (0.6 1 %).

The two marine harbour seal subspecies examined exhibited clear marine signatures in

the fatty acid patterns of theV blubber. Surveys of the composition of the blubber of five

phocid seal speîies revealed n-3/n-6 ratios ranging nom 9 to 24 , and levels of 20:l and

22: 1 ranging nom 5.0 to 16.6 and 0.7 to 6.5, respectively (Engelhardt and Waiker 1974;

West et al. 1979). While the fatty acid compositions of the blubber of P. v. concolor

and P. v. richmdri are similar to those of 0 t h marine seals, they can ais0 be

distinguished fiom one another on the basis of levels of monounsatrnited C, and G.

These fatty acids originate in marine copepods (Ackman 1980), and the significantly

higher levels of these compounds in the P. v. concolor sample (Table 4.4) are probably a

reflection of variation in the diet of the primary prey species in each area Levels of

92

different isomers of monoumatumted C, fatty acids can also be used to provide fiirther

evidence that the Lacs des Loups Uarins seals are feeding in fkeshwaîer; 22: l n 4 1

originates in marine copepods, while 22: ln-9 is found in fieshwater invertebrates

(Ratnayake and Ackrnau 1979; Bell et al. 1994)- The ratio of 22:ln-1 l to 22:ln-9 in the

P. v. mellonae seals (0.56) was much lower than in the P. v. richar& and P. v. concolor

seals (7.52 and 8.21, respectively), and 22: ln4 1 was virtuaily absent in the fkeshwater

seals, suggesting M e r that the Lacs des Loups Marins animals are not feedùig in a

marine food web,

This study has demonstrated the utility of two relatively new analytical techniques for

the study of marine mammals with access to marine or fieshwater ecosystems. While the

present analyses were performed on samples that did not allow year-round inférences of

seal diet, they do provide additional evidence that P. v. mellonoe is resident in fieshwater

and point to interesting fùture research possibilities in this unique aquatic ecosystern.

F u r t h isotopic adys is of bone collagen would, for example, provide dietary inferences

based on the lifetime of individuals (Tieszen and Boutton 1988). In addition, isotopic

and fatty acid analysis based on samples collected throughout the year would reveal

seasonal dietary patterns.

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Lajtha, &, and RH. Michener- 1994. Stable isotopes in ecology and envUonmentd

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101

Smith, RI., DM. Lavigne, and W K Leonard 1994. Subspacific status of the

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betweea populations of hsh- and Salt-water harbour seais (Phoca vitulina) using

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Smith, R J . 1997. Status of the Lacs des Loups Marins Harbou. Seal, Phoca vi~Iina

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the Belcher Islands. Herbert Jenkins Ltd., London,

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Waker, I.L., and S.A. Macko. 1999. Di- studies of marine mammals Usmg stable

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West, G.C., LI. Burns, and M. Modafferi. 1979. Fatty acid composition of blubber fiom

the four species of Bering Sea phocid seais. Can, J. 2001. 57: 189-195.

Table 4.1. Specimens of Phoca vzMtuIina n'char&i, P. v. concoior and P. v. meZIonae used

in the stable isotope (SI) and fatty acid (FA) d y s e s .

Specimen Types of Location Collected Date Sex YS= Collected

P. v. richarhi

PV-11

PV-12

PV-13

PV-14

PV- 15

PV-16

PV-17

PV-18

PV-19

PV-20

PV-21

PV-22

PV-23

PV-24

PV-25

PV-26

Danger Reefs, B.C.

Danger Reefs, B.C.

Snake Island, B.C.

Nanaimo, B.C.

Snake Island, B.C.

Snake Island, B.C.

Crofton, B.C.

Miami Island, B.C.

North Stuart Channel, B.C.

North Stuart Channel, B.C.

Miami Island, B-C.

North Reef (Crofton), B.C.

Escape Reef, B.C.

Thetis Island, B.C.

Miami Island, B.C.

Danger Reefs, B.C.

Grand Manan Island, N.B.

Grand Manan Island, N.B.

Grand Mmm Island, N.B.

Grand Manan Island, N.B.

Grand Manan Island, N.B.

PV-93-01

PV- 1 527

PV-1612

PV-1618

PV-G1

CMN-6138

CMN-93 1 1

CMN-10358

CMN-20289

CMN-20290

CMN-2029 1

CMN-20600

CMN-20685

CMN-20687

CM-13161

CM-152 L7

CM-15218

CM-15219

CM915220

CM015221

CM017679

CM-18698

CM- 18745

CM-18746

CM- 18747

CM- 18748

CM- 19445

Grand Manas Island, NB.

Anorluitso~, Greenland

S. Aappilattoq, Gmdand

Anorluitsoq, Greenland

Qaqortoq, ~ ~ e ~ a n d

Nettilling Fiord, Banin 1s.

Saguenay Co., Qué,

Cape Dorset, Bafnn 1s.

Ha Ha Bay, Qué-

Ha Ha Bay, Qué.

Ha Ha Bay, Qué.

Anticosti Is., Qué.

Grand Grève, Qué.

Grand Grève, Qué.

Freshwater Lake, Belcher 1s.

Kasegahk Lake, Belcher 1s.

Kasegdik Lake, Belcher Is.

Kasegalik Lake, Belcher 1s.

Kasegalik Lake, Belcher 1s.

Kasegalik Lake, Belcher 1s.

La Tabatière, Qué*

La Tabatière, Qué.

Lake, Belcher is., N. W.T.

Lake, Belcher Is., N.W.T.

Lake, Belcher Is., N.W.T.

Lake, Belcher Is., N. W.T.

Lake, BeIcher Is., N.W.T.

93-09-09

86-05-0 1

86-05-04

86-05- 1 1

1993

26-03-13

28-06-04

28-10-16

50-06-16

50-06-15

50-06-16

52-06-25

5 1-07-23

5 1-07-23

37-10-1 1

3 7-07. 15

38-06-30

38-06-30

38-06-30

38-07-19

39-07-0 1

39- 12-23

Spring 1940

Spring 1940

Spring 1940

Spring 1940

40-08-1 1

P. v. mellonae

CM01521 1

CM-15212

CM-15213

CM-15214

CM-15215

CM-17744

LM-89-0 1

LM-89-02

LM-94-01 (Ml)

LM-94-02

LM-94-03 (AF2)

LM-94-04 (M3)

LM-94-05 (AF4)

LM-95-01 (AFS)

(recaptured AF 1)

LM-95-02 (AF6)

LM-95-03 (AF7)

LM-95-04

SI

SI

SI

SI

SI

sr

Both

Both

Both

Both

Both

Both

Both

SI

SI

SI

SI

Lacs des Loups Marins, w- Lacs des Loups Marins, W. Lacs des Loups Marins, Qué- Lacs des Loup Uarins, Qué. Lacs des Loups Marins, Qué. Lacs des Loups Marins, Qué.

Lacs des Loups Marins, W. Lacs des Loups Marins, Qué.

Lacs des Loups Marins, Qué.

Lacs des Loups Marins, Qué.

Lacs des Loups Marins, Qué.

Lacs des Loups Marins, Qué.

Lacs des Loups Marins, Qué.

Lacs des Loups Marins, Qué.

Lacs des Loups Marins, Qué.

Lacs des Loups Marins, W. Lacs des Loups Marins, Qué.

Summer 1936

Summer 1937

3 8-03-22

38-03-22

38-03-23

1938-39

89-08

89-08

94-09-04

94-09-05

94-09-07

94-09-10

94-09-1 1

95-09-04

95-09-07

95-09-09

95-09-09

LM-95-05 (AFS) SI Lacs des Loups Marins, 95-09-1 1 M w* Lacs de^ LOUPS Marins, 95-09-14 M Qué- L~CS de^ LOUPS MannS, 96-08-25 M Qué*

Lacs des Loups Marins, 96-08-27 F Qué* Lacs des LOUPS Marins, 96-08-27 F Qué-

Lacs des Loups Marins, 96-08-30 F w* Lacs des Loups Marins, 96-08-3 1 F W.

Table 4.2. Stable-cdma and nitrogen isotope values (meaa SD)

of hait and serum h m s d s fiom h h , brackish and sait-wter,

Popdation n 8I3c (%O) o%(%o)

- - -- --

Phocta vitdina mellonae (hair) 23 -23.l*l.4 12.%1.1

P. v- donae (serum) 10 -24-4*1.0 12.9M.8

P.v.concolor(KasegaWrLake) 11 -19.1*1.0 13.ét1.2

P. v- concolor (marine) 16 -6.112 1623.4

Table 4.3. Fi& fauna of Lacs des Loups Marins, QuéTbec: abundance and stable-carbon

and nitrogen isotope values (mean * SD) nom muscle tissue.

Species Numeric Relative Stable Isotope Analyses

Abundance Abundance

Coregonus artedii 41 11.3 23 -25.5M.8 10.M.6

Prosopium cyZid-aceunt 23 6.4 15 -2Oht1.8 8SM.4

* Specimens were opporRniistically obtained using dip-nets and therefore not included

in abundance calculations.

Table 4.4. Levels of selected fatty acïds in the blubber of thtee subspecies of tiarbour

seal (Phoca vituiina).

P. v, concolor P, v. richardri P. v. mellonae (marine) (marine) (fFeshwata)

Fatty acids

18:2n-6

18:3n-3

20: ln-9

20:4n-6

22:ln-11

Total saturates

Total monounsaturates

Total PUFA

Total C,, PUFA

n-3/n-6 ratio

Note: Levels of fatty acids are given as percent wet weight (* SD.) of total fatty acids

and mknowns present in the sample. Levels of each fatty acid in a subspecies foilowed

by different letters were significantly different (p < 0.05). Sample s k s were 5, 16,

and 7 for P. v. concolor, P. v. n'char&, and P. W. mellonae, rrespeîively.

Fig. 4.1. Distri'buîion of stable-carbon and nitmgen isotope ratios in the hair of hsnbour

seals with die& derived 60m marine (Phoca vitdina concolor; Northwest Atlantic),

fieshwater and marine (P. v. concolo~ Kase- Lake), and 6eshwater (P. v.

mellonae) sources. Subsaipts 1 and 2 denote the 1994 and 1995 values, respectively,

for the one fieshwater seai that was marked and recaptuted.

A P.v.mellonae

0 P x concolor (Kasegalik Lake)

1 a P. v. concoior (NW Atlantic)

Fig. 4.2. Plot of total n-3 vs. total n-6 fâtty acids (as percent of total fatty acids) in the

blubber of marine (Phoca v i~I ina nkharaki and P- v. concob) and fieshwater (P- v.

d o n a e ) harbour seals.

0 P. v. richardsi

P . v-coricdor I

A P. v. rnelbriae

O 2 4 6 8 I O 12 14

Total m6 fatty acids (%)

the Ungava pmïmda's isostatic rebound since the retreat of the Lamentian ice sheet

(Doutt 1942). Other authors have disputed this interpretation, arguing that the seals are

likely able to travel fieely between salt and fieshwater (Mansfield and McLaren 1958,

Mansfield 1967, Smith and Horonowitsch 1987, also see Honacki et al. 1982, King 1983,

Wiig 1989, Reeves et al. 1992). Smith and Horonowitsch (1987) noted that the large

waterfklls on the Rivière Nastapoca wodd seem to make this avenue of exchange with

Hudson Bay (a distance of approximately 160 km) &eIy, but that the more placid

Rivière aux Feuilles, Rivière aux Mélèzes and Rivière DuGué, that drain toward Ungava

Bay (a distance of approximately 300 km), might provide a more likely access to the

ocean.

The question of seasonal changes in distribution has also been the subject of reguiar

speculation (e.g. Doutt 1942, Mansfield 1967, Smith and Horonowitsch 1987,

Consortium Gilles Shooner & Associés et al. 199 1)- Consortium Gilles Shooner &

Associés et al. (1991) hypothesised that the seals congregate in larger bodies of water,

such as Lacs des Loups Marins, Lac Bourde1 and Petit Lac des Loups Marins in the

winter, but then disperse into the swromding watersheds in the spring (Fig. 5.1). Unlike

ringed seals (Pusa hispida), harbour seals do not possess elongated nails on theù anterior

flippers that cm be used to mate breathing holes in the ice. As a consequence, any

harbour seals that over-winter in Ungava lakes are dependent on naturally occurring ice-

fkee areas. Doutt (1942, p. 7 1) quoted information relayed to him by his Cree guides:

Open water is the place to get seals when the lakes are

fiozen. Such places are found in the rivers wtiere steep

116

hills narrow the channel and make swift water. The holes

change rapidly with changes in the weather - wann

weather opens new holes and makes old ones larger - cold

weather closes up the smalier ones entirely and narrows the

larger ones to small dimensions.

This observation of seal behaviour was the same as that provided to me by Cree hunters

in Whapmagwstuï (J. Petagumshim. Sr., pers. corniil). There have also been

suggestions that the Lacs des Loups Marins harbour seals may take advantage of

shoreline deformations of ice caused by droppiag water for protection - a mturaliy-

o c c h g analogy to the ice caverns that are excavated by ~ g e d seals (Twomey 1942;

Smith and Horonowitsch 1987; Dean Consulting & Research Associates Inc. 1991).

The use of satellite-linked transmitters is an increasingiy common method of

monitoring the movements and behaviour of marine mammals (Stewart et al. 1989;

Lesage et ai. 1995; Stewart et ai. 1996; Westgaie and Read 1998). The objective of the

present study was to investigate the seasonal movements of Lacs des Loups Marins

harbour seais with satellite telemetry in order to test Doutt's hypothesis that they have a

rehcted range and are resident in Lacs des Loups Marins throughout the year.

Materials and Methods

Seai Capture and Transmitter Configurations

Harbour seals (P. v. meIIorme) were observed during field studies at Lacs des Loups

Marins in August and September of 1994.95 and 96. Satellite-linked transmitters were

117

placed on nine harbour seais that were captured in 1995 and 1996 (Table 5.1). Seals

were captured in braided monofilament tangIe nets (dimensions approxjmately 60 m x 3

m, mesh size of 10 cm) that were anchored on land and set perpendicular to the shore.

Seals were removed nom the net and strepped to a modi£Ied stretcher for restraint. Body

mass was measiired using a spring balance, and a variety of samples were taken

(Chapters 3 and 4).

After the area of attachent was thoroughiy cleaned and dried usùig acetone and a

towel, satellite-linked transmitters were placed on each seal using five-minute epoxy

(Devcou, Danvers, Massachusetts, USA) that was vigorously worked into the pelage.

Two different transmitter configurations were used. In 1995, the transmitters were

epoxy-cast satellite-Iinked thne depth recorders (TDRs) (Wildlife Cornputers,

Woodinville, Washington, USA), and were placed between the animals' shouider blades.

These TDRs had a 17 cm whip antenna, measured 6 cm x 14 cm x 6.5 cm and weighed

approximately 400 g. Each instrument was powered by five 213 A Lithium cells. To

conserve battery Me, the TDRs were duty-cycled 8 hourslday, with the exception of the

TDR on seal AF5 (5132-951, which was set to transmit every other &y. Unfortunately,

the conductivity of Lacs des Loups Marins was so low that it was bene& the level of

sensitivity of the TDRs' salt-water switch (which prevents TDR transmission when the

seal is submerged), so it became necessary to tum the transmitters on permanently,

rendering impossible the use of the TDRs' environmental sensors.

In 1996, the configuration of instnunent that was used was a flat-board ST-10

transmitter (Telonics, Mesa, Arizona, USA) mounted in a low profle, rectangular, lexan

box. These transmitters had a 17 cm whip antenna, measrned 13.4 cm x 4.7 cm x 1.9 cm

and weighed 200 g. The ST-10s were set to &t 6 hours out of 48 with a 60 second

repetition rate. Because they were substantiaily smaller than the TDRs, it was possible to

attach them nearer to the backs of the animais' heads.

Data Analysis

Location data fiom each seal were obtained fiom Service ARGOS, Inc. (Landover,

Maryland, USA). Service ARGOS provided information on the @ty of the estimated

location, and divided the location classes into seven categories: Class 3 (at least six

uplinks received in a single satellite pas, position accuracy better than 150 m); Class 2

(five uplinks received in a single satellite pass, position accuracy within 350 m); Class 1

(four uplinks received in a single satellite pass, position accuracy within 1 km); Class O

(less than four uplinks received in a single satellite pas, position accinacy greater thau 1

km); Class A (three uplinks received; no estimate of position accuracy); Class B (two

uplinks received; no estimate of position accuracy); and Class Z (a single uplink

received; no latitude and longitude provided) (Service ARGOS 1996). Clam Z upünks

produced no useable positions for the purpose of the analysis. Ail estimated locations

were filtered using plausibility checks on speed and geographic position. Consecutive

positions resulting in an average travel speed of greater than 7.2 kmhour were excluded;

for an example of the use of such a nIter see Read and Westgate (1997). This filter value

was selected based on published travel speeds of phocid seals m e s 1984). Each

119

estimated location was also Iïitered by comparing the geographic location ofco~lsecutive

positions and excluding those positions that were improbable or impossible based on

known pinnïped behaviour and the shapes of the animals' satellite tracks. For example,

positions that were many km fiom any body ofwater, and "spikes" in the anhals' tracks

(an extremely large position movement between two other, very close, positions) were

validated using this Nter. If there was any doubt a s to the plausibility of a position, it

was left in the andysis.

Analysis of movement &ta was performed using Arcview Geograpbic Information

System (GIS) (ESRI 1994). Only the best position obtained per &y nom each seal was

included to avoid bias associated with multiple daily positions. Positions (filtered, best

per &y) were plotted on digitised maps of the study area provided by the National

Topographie Data Base (NTDB), and consecutive positions comected with tcack he s .

Average daily disiance (ADD) traveled was calculated by d g the distance (km)

between the best position received each day for ail days of the deployment and then

dividing this value by the number of days of the deployment.

Two different calculations of home range were performed with the filtered, best per

day positions, and plotted on the NTDB maps. Minimum Convex Polygons (MCP) were

calculated fkom each animal's positions (ESEU 1994)' and the amount of land within each

polygon subtracted fiom the total polygon area to yield esthates of the amount of the

lakes' d a c e area potentiaüy utilised by each animal. Total area used by ali seais was

calculated by constnicting a MCP around locations of all seals. Though MCP is stiil

120

ofim used by researchers to describe home range, the greatest disadvantage of this

method is that the sïze of the home range estimate can increase indefinitely as the number

of locations increases (White and Garrott 1990). Thus, calculations of kernel home range

(Worton 1989) were also undertaken for each animal and for the total of all anhals

capture& The kemel home range method does not d e r fiom a relationship between

sample size and home range size (White and Garrott 1990). It is an estimate of the

harmonic mean home range, and weights the points based on the ''utilisation

distn'bution." A point gets a higher weight if it is in a cluster ofother points (i.e. a high

utilisation area) (Worton 1989; White and Garrott 1990). Kernel home range (ceil sïze:

250 m) was calculated with least-squares cross-validation of h (the smoothing parameter)

(see Worton 1989). Arcview GIS allows the caiculation of various probabilities of use,

and confidence areas of 2S%, 50%, 75% and 95% were chosen. The surface area of

water withui each of these probability areas was then calculated

Resulîs

A total of 39 seal sighthgs were made in Lacs des Loups Marins during August and

September of 1994,9S and 96 @ig. 5.2).

Satellite transmitters placed on the seals provided &ta fiom 29 to 177 days (Table 5.1).

A total of 1067 locations were received on 438 tracking days. The mean number of

positions per day per animal for all location classes ranged from 1.6 1 O. 1 to 3 .9 * 0.2

(Table 5.2). Positions with location classes A, B, O, 1,2, and 3 accounted for 50.7% of

all position estimates. Positions that were incorporated into the analyses (non-2, best per

day, filtered) accounted for 20.4% of al1 position estimates (Appendix 1).

The transmîtter on seal 5 132-96 ody produced 3 position estimates (non-Z, best per

&y, fltered) likely due to a malfiinction soon after deployment (Appendix 1). These

positions were therefore excluded fiom Mer analysis. Average daily distances

traveled ranged fiom 1.5 to 9.8 km and Minimum Convex Polygon home ranges varied

fiom 100.2 to 625.7 km2 (Table 5.2; Fig. 5.3). The size of these home ranges were very

much hcreased by outlying points: estunated locations at a distance fiom the animals'

primary areas of utilisation. The majority of these outlying points were location classes

A and B, which are the least precise estimates of location (Service ARGOS 1996). These

points often had an IQ value of O, indicating that the location estimate was of poor

quality (IQ is an indicator that estimates residual error on the signal freqyency

calculation and transmitter oscillator fiequency drift between two sateliite passes)

(Service ARGOS 1996). Examples of such locations are italicised and underlined in

Appendix 1. Despite the likely poor quality of some of these locations they were left in

the anaiysis because of the conservative nature of the filter employed

From an examination of Figures 5.3 and 5.4, the ciifferences in the movements and

ranges of the 8 se& becorne clear. Seals 5130-95 and 5132-95 had ranges in simila.

locations, and spent the majority of their t h e slightly South of the geographic centre of

Lacs des Loups Marins. Seal 5 13 1-95 spent the majority of its time in a branch of Lacs

des Loups Marins slightly North and West of the centre of the laice, near Lac Boutdel.

122

Seai 5 133-95 was tagged in Lacs des Loups Marins and then spent the next two and a

half months in a s m d river that comects Lacs des Loups Marins and Petit Lac des

Loups Marins. Seals 513 1-96 and 5 133-96 spent the majority of theu t h e slightly North

of the centre of Lacs des Loups Marins. Seal 9941-96 spent its time in the southern

portion of Lacs des Loups Mirim. The two most southeni position estimates for this

animal. which are weil outside Lacs des Loups Miuïns, and which greatly increased the

size of its MCP-calculated home range, are location class £3, with an IQ of0 (Appendix

1). Similarly, seai 9942-96 was tagged near the outfîow of Lacs des Loups Marins into

the Nastapoca River. This animal spent the majority of its time in this area, with the two

location estimates in Lac à l'Eau Claire being location class B, IQ = 0.

There was no evidence of monthly shifts in movement or distribution patterns by the

seals (Figs. 5.3 and 5.4). The number of satellite uplinks was dramatically decreased

with the onset of ice cover on Lacs des Loups Marins. *ch usuaily occurs around the

first week of November (Consortium Gilles Shooner & Associés et al. 1991). The

predictability with which the traosminers ceased o p t i o n - in both years of the study,

despite very diffêrent transmitta coafiguratiom - around this tùne is strikùig. The dates

of the 1st upünks with nu-c location classes for each animal (except 5 132-96, which

malfûnctioned soon after deployment) are as follows: 5 1 30-95 (Nov. 7); 5 1 3 1-95 (Nov.

i 1); 5132-95 (Nov. 23); 5 133-95 Wov. 23); 513 1-96 (Nov. 03); 5 133-96 (Nov. 23);

9941-96 (Jan. 13); 9942-96 (Jan. 18). Given the paucity of location estimates d e r

November 15, it was not possible to test whether there was any movement of seals

toward naturaIly-occ-g ice-fk areas in the winter. With the exception of the last

123

position with a numeric location class for 9942-96, wtùch was out of Lacs des Loups

Marins on the boundary of the Lac à l'Eau Claire watershed, al1 positions recorded after

Nov. 7 were in simila- geographic locations as those recorded prior to the onset of ice

cover (Appendix 1).

By combining the locations of al i 8 snnnaln (Fig. 5.5) and calcdating the utilisation

distriiution, it was possible to visualise the fkct that the seals had two distinct areas of

utilisation: seven d a l s utiliseci Lacs des Loups Manns/Lac Bourde1 proper, and 5 133-

95 utilised the river between Lacs des Loups Marins and Petit Lac des Loups Marins.

Discussion

These results support the hypothesis that P. v- rnellonae is resident in the Lacs des

Loups Marins area throughout the year. In addition, the results provide evideme that

individual seais preferentially utilise specinc sites within the lake (Figs. 5.3 and 5.4).

Though there were few location estimates available subsequent to the onset of ice cover

on the lake, what few satellite uplinks that did occur indicated that the seals remained in

the area. Given the site fidelity exhibited by P. v. mel lo~e , and the fact that a large

number of animals were captwed in the vicinity of the base camp that was utilised over

the three years of the shidy (located at the geographic centre of Lacs des Loups Marins,

56.524' N, 73.78 O W), the utilisation distribution of the telemetered harbour seals is

clearly partly a fhction of capture effort. The numeric location-class positions obtained

fiom seals 5 133-95 and 9942-96 do, however, provide evidence of some habitat useage

outside of Lacs des Loups MirbLac Bourde1 proper.

124

Considering that the battery power of the -tters should have been ample to

provide uplinks into January- and that 6 out of 8 transmitters failed concurrent with the

omet of ice cover in early November (and the two that did keep transmi-tting began to do

so with much reduced reliability amund this tirne) (Appendix l), it is reasonable to

conclude that it was winter conditions that impaïred or premaîurely ended transmissions.

There are two possible explanations for this: 1) Both configurations of transmïtters were

more sensitive to malfiuictionkg in the cold than was expected; or 2) The seals behaved

in such a way that they destroyed the t d i t e r s - such as shearing the antennae as they

exited or enterai the water through the ice - or rendered tnuismissîons impossible -

such as spending the majority of their tirne undemeath the îce as suggested by Twomey

(1942) and Smith and Horonowisch (1987). The first possibility is d ike ly given the

extent to which other shuiies of pagophilic phocids have successfbiiy used the same

configurations of transmïttets (e-g. Lydersen and Hammill 1993; Lydersen 1995). The

second possibility, therefofo seems the most lürely explanation and should S o m the

experimental design of any M e r field -es in this area

The results of the present investigation are similar to those that have been undertaken

of harbour seal populations in Europe and on the East and West coasts of North Amerka

Harbour seals are weli-known for king non-migratory and exhibithg a high degree of

site fidelity (Yoçhem et al. 1987; Thompson 1993). Olesiuk et al. (1995) found that 13

of 15 translocated seals found their way back to their original haul-out sites, some

swimming 272 km at swimming speeds of up to 62 km/dayy Thompson (1993)

125

recognised two broad categories ofharbour seal movements: 1) those between W-out

sites and the sea, which are primarily for forsging and occur within 50 lan of haul-out

sites; and 2) those that occur between different haul-out sites, which may involve

dispersal and cm occur seasonally *en seais switch to sites that are more SUitabIe for

pupping or are closer to foraging areas. Consistent with this summary, Lesage et al.

(1995) reported that one satellite-telemeterd barbour d in the S t Lamence River

remained within 80 km of its capture site thrwghout the winter, and in November a

second animal lef€ the area where it had been captureci and moved 520 km to the Baie des

Chaleurs where it remained for most of the winter before retmûng in late March to the

capture location. In southern California, Stewart et al. (1989) used satellite telemetry to

monitor the movements of a single seal and reported that the animal ranged up to 48 km

fkom its haul-out site, although most trips were within 5 km. Thompson and Miller

(1990) and Thompson et al. (1994) observed seals foraging up to 45 km nom haul-out

sites, but females with pups restricted their movements to within 2 km of haul-out sites

during the early part of the lactation p e r i d Other authors have observed foraging to be

limited to 20 km of haul-out sites (Bjerge et al. 1995); l e s than 5.6 lan (Surym and

Harvey 1998); and within 30 km (Tollit et al. 1998). Evidence h m seals that are

resident in fieshwater (Lake Saimaa ringed seal, Pusa hispida saimmik) indicates that

home range size in these animals may be even smailer than oceanic Purcr subspecies.

Maximum distances traveled by individual Saimaa seais ranged fiom 3.4 to about 18 km,

much shorter distances than ringed seals in the Arctic Ocean, which sometimes swim up

to hundreds of km (Sipila and Hyvarinen 1998).

The range size of harbour seals has been observed to Vary accordhg to sex and

reproductive condition. Using Minimum Convex Polygon calculaîions, Thompson et al.

(1994) and Vau Parijs et al. (1997) showed that female harbour seals restnct their range

dining the early part of the lactation @od (to approximately 5-35 km2) but that they

make foraging trips in late lactation and are more widely dispersed when in oestrus

(ranges of 50- 1 15 w). Male home range sizes are generally larger at ail times of the

year, but also Vary seasonally: males continue to travel widely during the early puppîng

period (ranges of65480 km2), but then restnct their ranges to 4-70 Lm2 at around the

tirne that females start to make foraging trips in late lactation. These MCP home range

sizes are similar to those observed in Lacs des Loups Marins seais (Table 5.2).

UnfortunateIy, due to a paucity of data points, I was unable to test sex-related or seasoual

variations in home range size in P. v. d o n a e .

Implications for conservation

The redts derived fiom satellite telemetry provide information on the habitat utilïsed

by the Lacs des Loups Marins harbour seal that will be useful to the Govemment of

Québec should it proceed with plans to protect the range of P. v. melloMe (Dubreuil

1983; Québec, Province 1992). Such protection should, at a minimum, encompass the

watersheds of Lacs des Loups Marins, Lac Bourde1 and Petit Lac des Loups Marins (Fig.

5.5).

In light of the evidence that harbour seals rarely venture out of this relatively small

area, it is dinicult to hiow how to interptet Hydro-Québec's claim to have recorded

127

vocalisations ofharbour seals in 49 Ungava peninsula water bodies, ranging as far South

as the reservoir at LG4, part of the original James Bay 1 hydroelectnc project

(Consortium GiUes Shooner & Associés et ai. 1991). Hydro-Que- reported the

recording of 8 distînctiy diffecent kinds of vocalisations: 1) resembling the "click"

recorded firom harbour seais by Renoufand Davis (1982); 2) vaguely resembling the

'-serine caU" prohiced by harp d s (Pagophiüus groeniandicz~s) (hbhL et al. 1975);

3) resembling the 6C~hirp" of the harp seal @&hl et al. 1975); 4) resembhg the "tap" of

the walrus (Odobemrs rosmmils) (Sîirling et al. 1983); 5) resembliag a bearded seal

(Erignathw bmbattrs) vocalisation (Cleator et aL 1989); 6) resembling a Weddeii seal

(Leptonychotes weddellli) vocalisation (Thomas and Stirling 1983); 7) resembbg another

sort of harbour seal vocalisation (Renouf and Davis 1982); and 8) a new sound not

recorded in the Literature (Consortium Gilles Shooner & Associés et al. 199 1). During

the same observations, Hydro-Québec biologists reported that the male and f d e

harbour seals exhibited externdy visible sexual dimorphism, with males having a

rnarkedly more buibous forehead than females (Consortium Gilles Shwner & Associés et

al. 199 1). Hy&~Québec9s fhdings regarding sexuai dimorphism and vocalisations, and

the widespread distriauton of Lacs des Loups Marins harbour seals are not consistent

with any other data, either historical or contemporary (Boulva and McLaren 1979; Bigg

198 1; Smith 1997).

References

Bellin, N. 1757. Petite Atlas Mantirne.

Bigg, M.A. 198 1. Harbour seai - Phoca viîzdina Linnaeus, 1758 and P. Imglra Pallas,

181 1. Pages L-27 in Handbook of marine mammals, Vol. 2: Seals. Edited by

S.H. Ridgway and RI. Harrison. Academic Press, London.

Bjmge, A., D. Thompson, P. Hammond. M. Fedak, E. Bryant, H. Aarefjord, R Roen,

and M. OIsen. 1995. Habitat use and diving behaviour of harbour seals in a

coastal archipelago in Norway. Pages 2 1 1-223 in Whales, seals, fish and man.

Edited by A. S. Biix, L. W a l k and 0. Uiltang. Elzevier Science, Norway.

Boulva, J., and I.A. McLaren. 1979. Biology of the harbour seal, Phoca vitulina, in

eastern Canada. Fi&. Res. Board Can, Bull. 200.

Cleator, H.J., 1. Stirling, and T.G. Smith. 1989. Underwater vocalizations of the

bearded seal (Erignathus barbatus). Can. L Zool.67: l9oO-l9lO.

Consortium Gilles Shwner & Associés, SOMER, and Enviro~ernent alimité hc. 199 1.

Complexe Grande-Baleine, Avant-projet Phase 2, Bilan des connaissances sur le

phoque d'eau douce. Vice-présidence Environnement, Hydro-Que*.

Crowe, KJ. 1991. A history of the original pegles of northern Canada McGiU-

Queen's University Press,

Dean Consdting & Research Associates hc. 199 1. Investigations of the uuder-ice

habitats of fksh-water seah in the Lacs des Loups Mirbu region. A Report for

Hydro-Quebec, Service HyQauIique, Division Hydrometrie. Dean Comdting &

Research Associates, Inc. No-ch, Vermont-

Doutt, J.K 1942. A review of the genus Phoca. Am. Carnegie Mus. 29:61-125.

Dubreuil, C. 1983. Un statut de conservation p u r le phoque commun habitant les eaux

douces du Québec. Ministère de l'Environnement, Direction des réserves

écologiques et des sites naturels.

ESRI. 1994. Using Arcview GIS. Envitonmental Systems Research Institute, RedIands,

CA.

Honaclci, J.H., EE. Kinman, and J. W. Koeppl. 1982. Mammal species of the world.

Allen Press and the Association of Systexnatics Collections, Lawrence, Kansas.

Innes, H. S. 1984. Swimming energetics, metabolic rates and huid limb muscle anatomy

of some phocid seals. Ph.D. Thesis. University of Guelph, Guelph, Ont.

King, J. 1983. Seals of the world 2nd Edition. British Museum (Natuml History) and

Cornell University Press, 1- W.

Lesage, V., M.O. Hammill, and ICU Kovacs. 1995. Cornparison of dive behaviour and

movements of harbour and gcey s d s h m the St Lawrence estuary, Quebec,

Canada. Abstract at the Eleventh Bienniai Conference on the Biology of Marine

Mammals, Orlando, FL.

Lydersen, C. 1995. Energetics of pregnancy, lactation and neonatal development in

ringed seals (Phoca IUFpda)). Pages 3 19-327 in Whales, seals, fish and man.

Edited by A. S. Blix, L. W a l k and 0. ULltang. EIzevier Science, Norway.

Lydersen, C., and M.0. H m . 1993. Diving in ~ g e d seal (Phoca hispida) pups

during the nursing period. Cm. J. Zool. 71 :99l-996.

Mansfield, A, W., and I.A. McLaren. 1958. Status of the harb0u.r seal in the eastern

Canadian Arctic. Pages 46-50 in Fisheries Research Board of Canada, Arctic

unit, annual repott and investigators' summaries. Edied by H.D. Fisher.

Fishexies Research Board of Canada, Montreal, Que*.

Mansfield, A.W. 1967. Distri'bution of the harbor seal, Phoca vituïina Linnaeus, in

Canadian arctic waters. J. Mammal. 48:249-257.

W, B., J.M. Terhune and K. Ronald 1975. Undmter calls of the harp seai,

Pugophilus groenIandiw. Rapp. P.-v. Rem. ht. Explo. Mer 169533-543.

Olesiuk, P.F., T.G. Smith, G. Horonowitsch, and GM. Ellis. 1995. Translocation of

harbour seals (Phoca vitulina): a demonstration of homuig abilïty and site

fidelity. Abstract at the Eleventh Biennial Conférence on the Biology ofUarine

Mammais, Orlando, FL-

Quebec (Provioce). 1992, Réseau des résewes écologiques au Quebec (projets et

réalisations). Ministère de L'Enviromement, Direction de la conservation et du

paîrimoine écologique.

Read, A.J., and A.I. Westgate. 1997. Monitoring the movements of harbour porpoises

(Phocoeno phocoeno) with satellite telemeîry. Mar. Biol. l3O:3 1 5-322.

Reeves, R.R, B.S- Stewart, and S. Leatherwd 1992. The Sierra Club handbook of

seals and sirenians, Sierra Club Books, San Francisco, CA-

Renouf, D., and M.B. Davis. 1982. Evidence that seals may use echolocation. Nature

300:635-637.

Service ARGOS. 1996. User Manual, 1.0. Sexvice ARGOS, hc., Landover, Maryland,

USA.

132

Sipila, T., and H. Hyvhen. 1998. Status and biology of Saimaa (Phoca hispida

saimertsb) and Ladoga (Phoca hhppida ladogemik) ringed seals. Pages 83-99 i n

Ringed seais in the North Atlantic. Edited by MJ. Heide-Jmgensen and C.

Lydersen. The North Atlantic Marine Uammal Commission, Tromse, Norway.

Smith, R.J. 1997. Status of the Lacs des Loups Marins Harhur Seai, Phoca vitulina

rnellonae, in Canada. Can. Field-Nat. 1 1 1:270-276.

Smith, T.G., and G. Horonowitsch. 1987. Harbour seals in the Lacs des Loups Marins

and eastern Hudson Bay ürainage. Cm. Tech. Rep. Fish. Aquat. Sci. 1539.

Stewart, B.S., S. Leatherwwd, PX. Yochem, and M.P. Heide-Jmgensen. 1989.

Harbor seal tracking and telemetry by satellite. Mar. Mamm. Sci. 5361-375.

Stewart, B.S., E.A. Petrov, E.A. Baranov, A. Timonin, and M. fvanov. 1996. Seasonal

movements and dive patterns of juvenile Baïkal seals, Phoca sibirica. Mar.

Mamm. Sci. 12:528-542.

Stirling, L, W. Calvert and H. J. Cleator. 1983. Underwater vocaiiations as a tool for

studying the distri'bution and relative abundance of wintering pianipeds in the

High Arctic. Arcîic 36:262-274.

Suryan, KM., and KT. Harvey. 1998. Tracking harbor seaIs (Phoca vihrlina richarthi)

to determine dive behavior, foraging activity, and haul-out site use. Mar. U.

Sci, 14361-372.

Thomas, I.A., and 1. Stirling. 1983. Geographic variation in underwater vocalizations of

Weddell seds (Lqtonychotes wddeZZi) fiom Palmer Peniasula and McMurdo

So~nd, Antarctica, Can. J, Zool, 612203-2212.

Thompson, P.M. 1993. Harbour seal movement patteros. Symp. ml. Soc. Lond

66:225-239-

Thompson, P.M., and D. Miller. 1990. Summer foraging activity and movements of

radio-tagged common seals (Phoca vitdinu, L.) in the Moray Firth, Scotland 5.

Appl. Ecol. 27:492-50 1.

Thompson, P.M., D. Miller, R Cooper, and P.S. Hammond 1994. Changes in the

distribution and activity of female harbour seals during the breeding season:

implications for theù lactation strategy and mating patterns. 5. Anim. Ecol.

63 :24-30.

Toilit, D-J., A.D. Black, PM- Thompsm, k MacKay, HM- Corpe, B. Wilson, SM. Van

Parijs, K, Greilïer, and S. Parlane. 1998- Variations in harbour seal PIroca

vituIiltcl diet and dive-depths in relation to fotagiog habitat, J. Zml., Lond

244:209-222.

Twomey, A.C. 1942. Needle to the North: The story of a . expedition to Ungava and

the Belcher Islands. Herbert Jenkins Ltd., London.

Van Parijs, SM., PM. Thompson, D.J- TOU& and A. MacKay. 1997. Distri'bution and

activity of male harbour seals d h g the mating season. Anim. Behav. 54:35-43.

Westgate, AL, and A.J. R e d 1998. Applications of new technology to the

consenation of porpoises. MTS Journal 32:7û-8 1.

White, G.C., and RA. Garrott. 1990. Analysis of wildlife tracking data. Academic

Press. San Diego, USA.

Wiig, 0. 1989. A description of cornmon se&, Phoca vitdina L, 175 8, fkom Svalbard.

Mar. Mamm. Sci. 5:149-158.

Worton, B.J. 1989. Kemel methods for estimating the utilization distribution in home-

range snidies. Ecology 70: 164-168.

Yochem, P.K., B.S. Stewart, RL. DeLong, and DP. Demer- 1987. Die1 haul-out

patterns and site fidelity of harbor seals (Phocu vitulina rich&) on San Miguel

Island, California, in autumn, Mar, Mamm. Sci. 3:323-332.

Table 5.1. Description of harbour seals (Phoca vitulirra meIZonae) b t were

affixed with satellite tnmsmitters in Lacs des Loups Marios, Que* in 1995

and 1996.

Seal Sex Standard Body Capture Satellite Tracking

ID. Length Mass Date Package Period

(a) (kg) Cd)

1995 AFS F 119 55 4 Sep SLTDR 81

7 Sep SLTDR

I l Sep SLTDR

14s- SLTDR

25Aug ST-10

27 Aug ST-IO

27 Aug ST-10

30Aug ST-10

31Aug ST-10

Table 5.2. Mean number of uplinks per day; number of telemeterd positions used in

anaiysis; average daily distance (ADD), and minimum convex polygon (MCP) and

kernel home range estimates for 9 Lacs des Loups Marins se& observed in 1995 and

1996.

Seal ID Mean No, ADD MCP Kemel Kerne1 Kernel Kernel

number of of (km) Water (25) (50) (75) (95)

uplinks per psi- (km2) Water Water Water Water

&y tions 0an2) (km2) (kmL) (lan2)

used

5130-95 3.3 *O.l 52 9.8 502.5 14.8 29.9 51.0 291-4

Fig. 5.1. Saidy area in northem Quebec as depicted by National Topograpbic Data Base

digitised maps.

Fig. 5.2. Sightings of Lacs des Loups Marins harbour seais: 1994-96.

Sightings

Fig. 5.3. Movements of 8 Lacs des Loups Marins harbour seals as monitored by satellite

telemetry in 1995 and 1996. Minimum Convex Polygoas enclose aii estimated positions.

51 32-95 (4 Sep 1995 - 23 N ov 1 9951

9942-96 [31 Aug 1996- 18 Jan 19971

Fig. 5.4. Kernel home ranges (with confidence areas of 25%, 50%, 75%, and 95%) of 8

Lacs des Loups Marins harbour seals caiculated fkom satellite-telemetered positions:

1995-96.

Fig. 5.5. Utilisation distribution (with confidence areas of 2S%, 5û%,75%, and 95%)

calculated using combined positions of 8 sateIlite-telemeterd Lacs des Loups Marùis

harbour seals: 1995-96.

Position estimates for 9 Lacs des Loups Marins harbour seals used in this analysis.

Positions are frltered, best per day, and do not include locations fkom class Z (Service

A R G o S 1996).

Date Time (GMT) Latitude Longitude Location IQ

CIass

CHAPTER 6: GENERAL DISCUSSION

Mer alI, does it matter? Does it matter tbat Phoca vi.ïir'na

mellonue may not be a 3Iibspecifically distinct race? Does it matter that this supposedly land-locked race may not have been isolated in the Seal Lakes of Ungava for thousands of years? Such questions are of course important to biologists, and rightly so. But new discoveries have always had a way of consignïng old discoveries to the dustbin of history. For the reader of N e d e to the North the status of bagea, the mythic seal ofthe Ungava lakes, is secure: we know that k g e a is not netcheit. And we also lmow that ka~agea may be a-ed ody after the most arduous of joumeys, taken in mid-&ter, with the special knowledge of the nght assistants and guides, upoa completion of a period of patient waiting, and only d e r corning through the reqyisite tests and ordeals (James 1982, p. 205).

Subspecies and ESUs

The balance of evidence fkom this thesis indicates that Doutt (1942) was correct in his

subspecific designation of Phoca vi~ulina rnellonue. Subspecies were defined by Wilson

and Brown (1953) as king "genetidy distinct, geographicaliy separate populations

belonging to the same species" (p. 99). Expanding on this definition, McIvor et al.

(1995) cautioned that:

It is unlikely that any single ecological, morphological, or

gewtics-based approach will or should m e r the question

of subspecific status. Rather, an integrated approach

shodd be used involving natuml history, morphology,

range and disîri'bution, and molecular genetic data @.

Just such an "integrated approach" was used in the preceding chapters to address the

question of the Lacs des Loups Marins harbour d ' s subspecinc status (Table 6.1). As a

result of the publication of Smith et al. (1994) - which included a portion of the data

presented in Chapter 2 - the most recent and definitive summary of m a r k mammal

systematics has t e - m e d this status (Rice 1998). The extremely restricted and isolateci

range of P. v. m e l l o ~ e (Chapter 5), the evidence of year-round fieshwater feeding

(Chapter 4), si@caut differentiation in biah timing and morphoIogy (Chapter 2) al l

corroborate the historical and anecdotal evidence of the population's distuictness.

Though the mitochondriai DNA analysis was preliminary, it did reveal some unique

haplotypes in the Lacs des Loups Marins animals (Chapter 3). A larger genetic analysis

of harbour seal samples nom museums will be necessary before a complete picnüe can

be had of historical harbour sed genetic diversity in the Canadian Arctic, which was

likely greater than it is today (Mansfield 1967).

Though the concept of the subspecies is stili widely used to descnï intra-specific

levels of variation, Wilson and Brown (1953, p. 1 10) asaitely observed that "as the

analyses of geographical variation becorne more complete, the trinomial nomenclatorial

system will be revealed as inefficient a d super£luous." This dilemma was summarised

by Doutt (1955, p. 181) himself, when he s t a td

If we recognïze as a subspecies every local population

which cm be shown to be recognizable, we wïil eventuaUy

have to name as subspecies the mice fkom every valley and

every mountainside - perhaps even fiom every woodlot!

173

It was "out of a sense of fiiustration with the limitations of currmt mammalian

taxonomy in determinhg which named subspecies acnially represmt significant adaptive

variation" that Ryder (L986) advanced the concept of the "Evolutionarily Signiscant

Unit" or ESU. In identifjing ESUs, Ryder (1986) noted that the recognition of

inevitable uncertainty in the classifiCafion of potentially significa~t populations

necessitates that concordance be achieved between sets of data denved by differing

techniques (such as naturai history information, morphometrics, range and distribution

data, as weil as protein electrophoresis and molecular genetic techniques).

Since the time of Ryder's writing, the concept of ESUs has been widely adopted,

paaicularly by conservation biologists, and become an "operational term for a group of

organisms that should be the minimai unit for conservation management (Vogler and

DeSalle 1994). ESUs have been elucidated for organisms that had previously been

classified as species (Moritz 1994; V o g k and DeSalle 1994); subspecies (McIvor et al.

1995); stocks of marine rnammals (Duon et al. 1992); and individual populations of

Pacinc salmon (Penwck and Dimmick 1997; Waples 1998). Moritz (1994) attempted to

take the ESU concept £ùrther by distinguishing:

between two types of conservation units, both important for

management: E.S.U.s, concerned with historical

population structure, mtDNA phylogeny and long-term

conservation needs; and M.U.s Management Units],

addressing cumnt population structure, ailele fkquencies

and short-tenn management issues (p.374).

In contrast, Mallet (1995) advocated uncoupling taxonomic and conservation biology

concerns completely, making a strong case for his particdar "species definition for the

Modem Synthesis" M e at the same tirne noting:

We are much more interesteci in collserying a d

morph010gicaI~ ecological and genetic diversity than in

stnicturing conservation around a nebulws taxonomic level

about which, in the past, there has been so rnuch

disagreement (p. 298).

With the exception of Moritz (1994), P. W. mellonae fullills the criteria for ESU

designation as laid out by a variety of authors (Ryder 1986; Dizon et al. 1992; Vogler

and DeSalle 1994; Waples 1998). The Lacs des Loups Marins seal exhibits the

'~orphological, ecological and genetic diversity" that pieces of legislation which use the

ESU concept, such as the United States Endangered Species Act, are designed to protect

(Mailet 1995)-

DiEerentiation and Freshwater Seals

The harbour seal has the widest breeding distribution of any pinniped, with colonies

spread over 16,000 km fiom the East Baltic to Japan (Bigg 198 1; Stanley et ai. 1996;

Rice 1998). Boulva and McLaren (1979) noted that variation in numbers of pst-canine

teeth suggested thet muring between local harbour seal populations was limiteci, and

Stutz (1967) discussed this lack of mixing with respect to characteristic pelage patterns.

175

Further research has continued to reveal more reproductively isolated groups withui

subspecies (see Yochem et al. 1988; Temte 199 1; Lehman et al. 1992; Storr-Hansen and

Splüd 1992). in the words of Ray (1976), "Phma vitulina seems to be actively

differentiating if not in the proces of developing a species swarm."

Given that Temte (1991) for example, was able to distinguish significant

morphological variation between adjacent P. v. richmdri populations - wtiere

reproductive isolation seems solely the product of differences in annual birth timing

(which he labeled "allochronic" as opposed to aiIopatric diffaentiation) - it is perhaps

not surjsing that P. v. olellome is measiaably Merentiateci fiom P. v. concolor.

It is instructive in this regard to compare P. v. melZonue to the isolated ringed seal

populations in Lake Saimaa, Finland, Pusa hbpido sairnensIS, and in Lake Ladoga,

Russia, Puîa hrSpido ladogensii (Table 6.1). These two subspecies, the Baikal seal

(Pusa sibin'ca), the Caspian seal (Pwa caspïca) - which iives in brackish water - , and

P. v. mellonae comprise the sum total of the world's seal populations that are known to

be resident in f ieh or brackish water (Rice 1998). Both of the ringed seal subspecies

seem to have been isolated in fieshwater for a similar period of time as P. v. mellonae,

and though Lakes Saimaa and Ladoga are substaatially larger than Lacs des Loups

Marins, they are closer to each 0 t h and to the Baltic Sea (approximately 50 km) than

Lacs des Loups Marins is to Hudson Bay. Both of these ringed seal populations are few

in number, and both exhibit morphology and behaviour that are distinct from each other

and fkom ringed seals in the Baltic (SipilH and Hyvafulen 1998). It seems clear fkom

176

Table 6.1 that a small population of pinnipeds, when isolateci for approximately 7,000-

9,000 years, can becorne measurably diffiientiateed

Implications for Conservation

A number ofhatbour seai populations throughout the world have been extirpated (e.g.

in Lake Ontano), or are thteatened and in deciine (e-g. in Greenland; Sable Island, Nova

Scotia; Hokkaido, litpan; and Tugidak Island, Alaska) @ehy 1842; AUen 1880; Bodva

and McLaren 1979; Boveng 1988; Rougerie 1990; Pitcher 1990; Wada et ai- 1991;

Reijnders et al. 1993; TeiImann and Dietz 1993; Ellis 1998). Ironically, one of these

populations was ükely extirpated by the collecting that redted in sorne of the samples

used in this thesis. Between 1937 and 1940, the Carnegie Museum, Pittsburgh, PA,

collected a nll~llber of specimens of harbour seals fiom fkeshwater lakes on the Belcher

Islands, NWT: there does not w w appear to be any seals remaining in this area (Smith

et al. 1996).

Udess the govecnments of Canada and Quebec act soon to provide adequate legal

protection for P. v. meilorne, whkh bas recently been Listed as %ulnerable" by the

Committee on the Status of Endangered Wildlife in Canada (Smith 1997), the mail

group of seals in Lacs des Loups Marins may yet join the Iist of extirpated harbour seal

populations (Alfonso and McAllister 1994).

Referaces

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Table 6.1. Cornparison of fkshwater seal populations in Lacs des Loups Marins and

Lakes Saimaa (Finland) and Ladoga (Russia).

Attributes Lacs des Loups Lake Saimaa Lake Ladoga

seal (Phoca ( P m hispida (Pusa hispida

S d a c e area of lake (km2) 535 ' 4,400 19,890

Lake shoreline length (km) 1,830 ' 15,000 N/A

Distance fkom next nearest 160 ' 50 50

harbour seaI population (lm)

Likely time of isolation 7,300 8,000 9,000 2

(Years before present)

Number of individuals in Most recent 200 Minimum of

population estimate 5,000

Population density Approximately 0.05 seals/kxn2 0.25 seals/km2

0.1 seals/lan2 lw4

Morphologically distinct Yes * Yes 2,6*7 Yes L6*7

Behaviourally distinct Yes Yes Yes

Haplotypic Merences Yes ' No data No daîa

Exclusively fkeshwater Yes Yes 'O Yes 'O

feeding

Site fidelity in fieshwater Yes " Yes Yes

Footnotes: (1) Consortium Gilles Shootler & Associés et al. 1991; (2) Sipila and Hyvikiuen 1998; (3) A i i d and Séguin 1985; (4) Smith 1997; (5) Chapter 2 and Smith et al, 1994; (6) MiiUer-WiUe 1969; (7) Hyvarinen and Nieminen L990; (8) Ghapter 3; (9) Chaptet 4 and Smith et al, 1996; (10) Kakelit et al. 1995; (1 1) Chapter 5

STATUS OF THE LACS DES LOUPS MAIUNS HARBOUR SEAL, PHOCA ViVUUNA MZLONAE, IN CANADA

Richard J, Smith Department of Zoology University of Guelph

Guelph, Ontario, NIG 2 W 1

Status Recommended: Vulnerable

Status of the Lacs des Loups M&hs hatbour seai, Phoca wCfuIimz mellonae, in Canada

RJ. Smith Department of Zoology, University of Guelph, Guelph, Ontario, NLG 2W1

Smith, RI. 1994. Statu of the Lacs des Loups Marins harbour seal, Phoca vitdina meIIonae, in Canada. Report to the Cornmittee on the Staîus of Endangered Wildlife in Canada (COSEWIC). Canadian Wildlife Service, Ottawa, Ontario K1A OH1

The Lacs des Loups Marins harbour seal (phmu viiufim melhme) is a subspecies that occurs in the area of Lacs des Loups Marùis (Lower Seal Lakes) (56-57W. 73-74OW), 160 km east of Hudson Bay, on the Ungava peninsula of northern Quebec. W1th the possible exception of Lake Iliamna, Alaska, it is the only known harbour seal population that is resident in fkeshwater year-round. Wntten references to the unique appeamnce and behaviour of this seai date back to 18 18. The subspecies was descri'bed primarily on the bask of its Mique morphology and presumed long-tirne geographic isolation fkom neighboiiring oceanic harbour seals. Estimates of the population's size are imprecise, and range fiom 100 to 600 animals. Little is known of the seais' habitat requirements other than that they seem to feed exclusively in fkeshwater, and are likely reliant on some specific environmental features such as under-ice air pockets to sustain them through the winter. Pupping seems to take place substantially earlier (mid-April to mid-May) than in 0th- harbour seai populations at sunilar latitudes. The only known cause of human-induced mortality is occasional hunting by aboriginal peoples. Both the WCN and the Que- govemment have recognized that Phoca vitdina meIfonare is potentiaiiy VUltlerable or threatened because of its srnail population size, resbricted range, and suscept'bility to disturbance. A COSEWIC &tus of "vdnerablet' is recommended.

INTRODUCTTON

Subspecies description

The Lacs des Loups Marins harbour seal, Phoca v i . ~ mellome, is confineci to the an?

of Lacs des Loups Mkïm (Lower Seal Lakes), approximately 160 km east of Hudson Bay, on

the Ungava penllisula of northern Q u e k (Figure 1) [Doutt 1942; Anderson 1946; Scheffer

1958; Bigg 19811.

While there are mimaous references to batbout seaIs occUmng in kshwater worldwide

(Erlandson 1834; DeKay 1842; Auen 1880; Browne 1909; Grediell1910; Richard 1911; Strong

1930; Dunbar 1949; Fisher 1952; Wheeler 1953; Harper 1956; Harper 1961; Beck et al. 1970;

Paulbitski 1974; Roffe and Mate 1984; Williamson 1988), with the possible exception of Lake

Iliamna, Aiaska (Everitt and Braham 1980), Phoca vihrIina mellonae is the only kuown harbour

seal population resident in fkeshwater year-round (Atkinson 18 18; Clouston 1820; Hendry 1828;

Finlayson 1830; Low 1898; Lewis 1904; Flaherty 1918; Twomey 1938; Doua 1942; Manning

1946; Doutt 1954; Grabum 1969; Power and Gregoire 1978; Smith and Horonowitsch 1987;

Consortium Gilles Shwner & Associés et al. 1991).

Written references to the unique appearance and behaviour of Phoca vituiina mellonae

date back to Atkinson (1818). The subspecies was descriid primariiy on the basis of an

unusuaiiy dark pelage and an enlarged coronoid process on the mmdi'ble @outt 1942). with the

presumption that the population had been isolated for 3000-8000 years, trapped by the Ungava

peninsula's isostatic rebound since the retreat of the Laurentian ice sheet. ûther authors disputed

this interpretation, however, arguing that supposed morphologicai anomalies of Phoca vitdina

mellonae are merely artifacts of a small sample size, and that the seals are Wrely able to travel

fkeeiy between sait and Wwater (Mkfield and McLaren 1958; W e l d 1967; Srnit& and

Horonowitsch 1987; o h see Honaçki et al. 1982; King 1983; Wïig 1989; Reeves et al. 1992).

Other work, the majority of it recent, strongly supports the validity of Phoca vituiina meIZonae's

subspecifïc desiguation (Davies 1958; Consortium Gilles Shoona & Associés et al. 1991; Smith

et al. 1994, 1996; Smith 1996).

Taxonomy

Order: Carnivora

Family: Phocidae

Scientific name: Phoca vituiiltcl m e l l o ~ e

Common names: Lacs des Loups Marins seal, Seal Lakes sed, Ungava seal, phoque

d'eau douce, qasigiaq (Iiiuktitut), achiguanipe (Cree)

DISTRIBUTION

This is clearly a population with restncted distriiution (Figure 1). There are histoncai

references to the presence of this seal in Lac Minto, at the head of Rivière aux Feuiiles (Flaherty

19 18; Manning 1947), Lac Beneta, situated in the basin of Rivière aux Mélèzes (Manning 1947).

Petit Lac des Loups Marins (Atkinson 18 18; Clouston 1820; Doutt 1942). and Lacs des Loups

Marins (Hendry 1828; Finlayson 1830; Low 1898; Lewis 1904; Doutt 1942; Doutt 1954; Power

and Gregoire 1978; Berrouard 1984; Smith and Horonowitsch 1987). Several sightings have

been made by Hydro-Quelbec employees and contractors in the Rivière aux Feuilles, Lac Melvin

and Rivière Delay (Consortium Gilles Shooner & Associés et al. 199 1). Inuit hunters,

i n t e ~ e w e d by Hydro-Québec contractors, nported seeing or küling fieshwater seals in Lac

Guillaume-Delisle, Rivièce Nestapoca, Rivière Boniface, RïRive Niagumaq, Rivière Kuunga,

Rivière Longland, Lac Tasialuk, and Lacs des Loups Marins (Archéotec inc. 1990). The Cree

nation of Whapmagooshii considers the range of Phoca vifufina mefZonue to be Lacs des Loups

Marins, Petit Lac des Loups Marins, and Lac Bourdei, with some reports of animals having once

been in Lac à l'Eau Claire (Clearwater Lake) (J. Petagumskwn, Whapmagoostui, personal

communicati~n)~

Hydro-Quek has recently compileci observations made of these fieshwater seals

between 1970 and 1990 (Consortium Gilles Shooner & Associés et al. 1991). Though the

preponderance of their survey efforts have been concentrated in Lacs des Loups Marins, Hydto-

Québec's &ta nevertheless indicate the presence of seals in Rivière Nastapoca, Lacs des Loups

Marins, Petit Lac des Loups Marins, Lac Bourdel, Lac à 1Eau Claire, and Petite Rivière de la

Baleine. In addition, evidence nom recordmgs of undemater vocdizations suggests the

presence of seals in Rivière aux Feuilles, Rivière aux MéIèzes, Rivière du Gué, Grande Rivière

de la Baleine and La Grande Rivière (Consortium Giues Shooner & Associés et al. 199 1).

hiring the autumn of 1995,4 seais were capturd in Lacs des Loups Marins and affixed

with satellite-linked time-depth recorders (Wildlife Cornputers, WA). Ali four tags transmitted

nom early September to mid-Novernber, and during that t h e al1 four seals remained within

Lacs des Loups Marins or in the immediate vicinity (RI. Smith, unpublished data).

PROTECTION

Because Canada possesses no specific marine mammal or endangered specïes legislation,

and because it is u n c h whether Phoca vz'tulim mello~e, a marine miunmal in fieshwater, falls

within a provincial or federal jurisdiction, the population currently has minimal legal protection.

Freshwater seals north of the 55th pardel are üsted as a protected species under the James Bay

and northem Quebec Agreement (Québec 1976); however, this protection does not have the

force of law (J. Gimn, Ministère de l'Environnement et & la Faune, M~adisson, Qué. personal

communication). Phoca vitdina mellonae was recently listed by the Intemational Union for the

Conservation of Nature and Nahiral Resources (TUCN) as king "insufficiently known", meaning

that it is "suspected but not defhitely known to be endangered, vulnerable, or rare due to a lack

of reliable information" (Reijnders et al. 1993). The govemment of Qw'bec has listed the

population as "likely to be designated as threatened or VuInetable" (Que- 1992a), and is

considering whether to give legal protection to a portion of Phoca Mtulina mellonae's habitat

(Ihbreuil 1983; Québec 1992b). This protection should be a priority given that the proposed

Grande Baleine hydcoelectric development could have an adverse impact on a large portion of

this population's range (Romthal and Beyea 1989; RougerÏe 1990; Woodley et al. 1992; Smith

et al. 1994).

POPULATION S m AND TREND

Estimates of the size of this mail population are imprecise. A maximum of 500 animals

was the "guess" of Dom (1957), cited in Scheffer (1958). Power and Gregoire (1978) estimated

200 and 600 animals by two différent summations. The most recent estimate by Consortium

Gilles Shooner & Associés et al. (1991) was approximately 100 animal% or 0.1 seals/lmi2, in

Lacs des Loups Marins and Lac Bourdel. Population trends over tune obviously caunot be

calculated.

HABITAT

Little is known of the habitat and ecological requirernents of this subspecies.

The few dead animais that have been examineci were found to have h o n i d (SolveZmtrs

sp.) otoliths in their stomachs (Consortium Gilles Shooner & Associés et al. 1991; Smith et al.

1996). Cornparisons of the stable-isotope ratios and fatty acid profiles of the tissues of P. v.

mellonae and harbour seals collected fkom oceanic locations indicate thaf over a two year

period, P. v. mellonae seemed to be feeding exclusively in fieshwater (Smith et al. 1996).

Recent investigations found no permanent hadout sites on Lacs des Loups Mwns and

Lac Bourdel (Consortium Gilles Shooner & Associés et al. 1991). In winter, when the vast

majority of the lakes and rivers are covered in ice, the seals may rely on several physical feahaes

for their sources of air: areas that remain ice-fiee because of strong currents, fissures in the ice,

and air pockets created by the shoreline's complicated geometry or by the unddations in the

bottom of the sheet ice on the lake's d a c e (Smith and Horonowitsch 1987; Consortium Gilles

Shooner & Associés et al. 199 1; Dean Consulting & Research Asfociates Inc. 199 1).

None of the habitat of this population is protected. It is entirely on Crown land that

could be adversely affected by Hydro-Québec's constniction of the proposed Grande Baleine

hydroelectric project ( W d e y et ai. 1992) which, though indefinitely postponed by the current

provincial govemment, has not been cancelied altogether. One of the results of the Grande

Baleine environmental assessrnent pocess has been that Hydro-Quek is now r e q W to

evaluate properly the Wrely impacts ofthe pro* on the population, prim to comtmciion

(Review Bodies 1994). Some of these potential impacts inclde the disappearauce of ice-fkee

areas and under-ice shoreline shelters, upon which the seals may rely in the winter, ïrï inter

courses with altered flows arising fkom hydroelectric development The Grande Baieine project

may also affect the disîrïbution and abundance of the seals' prey, and contaminate the animais

with methyl mercury released h m the flooded, decomposing vegetation ( W d e y et al. 1992)-

The negative effects of this habitat destruction couid lead to a decline in the seal population and

an impoverishment of its genetic diversity (Alfonso and McAUister 1994).

GENERAL BIOLOGY

Reproduction probably occurs between mid-April and mid-May in the Lacs des Loups

Marins area; substantially earlier than 0th- harbour seal populations at a similar latitude (Doutt

1942; Archéotec inc. 1990; Consortium Gilles Shooner & Associés et al. 1991; Temte et al.

1 99 1 ; Smith et al. 1994). Since the lakes are still iced over at the time of pupping, and no birihs

have been observed on the ice, several authors have postulated that pupping takes place in under-

ice shelters (Consortium Gilles Shooner & Associés et al. 1991). like those of ringed seals

(Phoca hispida) (Smith and Stirling 1975).

The only known cause of human-induced mortality is occasional huntïng of the seals by

aboriginal peoples (Clouston 1820; Low 1898; Flaherty 19 18; Doutt 1942; Doutt 1954;

Consortium Gilies Shooner & Associés et al. 1991; J. Petagumskum personal cornmimication).

Seasonal movements of the population are poorly known, though the sporadic

observations of Gilles Shooner & Associ6s et al. (199 1) hùit at seals spending the wlnter mmths

in larger bodies of water iike Lacs des Loups hilarins, Lac Boutdel, and Petit Lac des Loups

Marins, with some dispersal into outlyïng, d e r bodies ofwater upon the melting of the ice.

These investigators report Einding many wom trails between of water fieqynted by the

seals, some as long as 0.15 km, and on inclines as steep as 25". There is no evidence that

animals move between the area of Lacs des Loups Marins and Hudson or Ungava Bays.

Howeva, though there are a number of impassable waterfds on the Rivière Nastapoca, some

authors believe that if the seals could move into the more placid nvers that flow north into

Ungava Bay, îhis would be a faible avenue of exchange between the M and salîwater

populations (Mansfield 1967; SOGEAM 1985; Smith and Horonowitsch 1987).

Preliminary evidence h m DNA sequencing of region 1 of the mitochondrial D-loop

indicate that P. v. meIZonae has haplotypes that are unique when compared to harbour seals in

the eastern Canadian arctic and Northwest Atlantic (Smith 1996).

A-als hauled out in the spring months are usually in s m d groups, whereas at the end

of the Sunmer, they are usually hauled out shgly or in pairs: This behaviour is probably related

to the rnoulting process (Consortium Gilles Shooner & Associés et ai. 199 1).

LIMITTNG FACTORS

The tendency of harbour seals to be distriibuted in small local popuiations makes them

vulnerable to disturbance (Maine Seal 1994). There are a aumber of examples of local harbour

seal populations king extirpated, or their numbers drastically reduced, by human activity. For

example, a s m d population that seemed to fiequent Lake Ontario was eliminated by the early

1800s (DeKay 1842; Auen 1880); the population in Greenland is practicaily extirpated

(Teilmm and Dietz 1993; R Dietz, personal communication), an important reason behg the

intensity with which it has been hunted and entangled in fishing gear; the population in

Hokkaido, lapan, is very small, with removals h m incidental catches in fishing gear exceedbg

recruitment (Reijnders et al. 1993). Given such evidence, the Lacs des Loups Marins seal

population is likely sensitive even to limiteci disturbance by humans.

SPECIAL SIGNIFICANCE OF THE SUBSPECIES

This population of harbour seais is unique, in that it is the object of reverence by the

aboriginal peoples of northem Quebec (Posluns 1993; Archéotec inc. 1990; M. George,

Whapmagoosnii, personal communication); it is the object of a wealth of historical references,

and seems to be unusual in a number of ways, including aspects of its biology (Consortium

Gilles Shooner & Associés et al. 199 1; Smith et al. 1994, 1996; Smith 1996). The population

has also ac-ed something of a public profile over the last few years (e.g. Dubreuil 1987).

RECOMIblENDATIONS

Future research should address how geneticaliy distinct this population is nom nearby

ocean populations, estimate cuwnt abundance and trends in the sue of the population, expand

our howledge of the population's range and seasonal movements, and collect basic biological

data such as the timing of reproduction, and critical habitat requirements.

This subspecies should be classified as Milnerable by both COSEWIC and Q u e k

provincial legislation, and cruciai areas of its habitat protected, possibly through Quek's

ecological reserve programme, or the establishment of a national pack

The recommendations expressed here are those of the author, and not necesdy those of

COSEWC or its member agencies.

EVALUATXON

It is essential for the fimne viability of this subspecies that the potential impacts on the

popdation the Grande BaIeme hydroeIectrïc project, and any other fuhire deveIopment in

Québec's north, be eliminated or serioudy mitigated.

Because of the inaccessibilïty of the population, there is no known trade in the

subspecies, legal or otherwise.

ACKNOWLEDGEMENTS

1 thank DM. Lavigne for his support. The coments and criticisms of RR Campbell

and two other members of COSEWr's Fish and Marine Mammal Subcommittee were dso

helpful. G. Luste provided some of the more obscure references. Funding for the production of

this manuscript was provided by the Natural Sciences and Engineering Research Council of

Canada and the International Minine M h m a l Association Inc., Guelph, Ontario.

AUTHOR'S RECOMMENDATION OF STATUS

Because of the population's uniqueaess, smail size, restricted range, and the possibility of

its being adversely impacted by future development on the Ungava paillwla, Phoca vitzdina

mellonae should be classified as vulnerable.

LITERA- CITED

Alfonso, N., and D.E. McAllistec. 1994. Biodivefsity and the Great Whale hydroelectnc

project. Great Whale Environmental Assessment: Background Paper No. 11, Great

Whaie Public Review Support Office. 75 p.

Allen, I.A. 1880. History of North American pinaipeds - A mortograph of the walnises, sea-

lions, sea-bears and seals ofNorth Arnerica U.S. Geologîcal and Geogqhical Slwey

of the Territones. Miscellaneous Publication No. 12. 785 p.

Anderson, RM. 1946. Catalogue of CanRdian recent mammals. National Museum of Canada

Bulletin No. 102. Kin@ Prhter, Ottawa. 238 p.

Archéotec inc. 1990. Complexe Grande-Baleine, Avant-projet, Phase 2, Le phoque d'eau

douce: Eléments pour une compre%ension de son utilisation par les Autochtones du

Nouveau-Québec, vice-présidence Environ~lement, Hydto-Québec. 145 p.

Atkinson, G. 181 8. Journal of George Atlàoson II. Page 61 in Northem Quebec and Labrador

journals and correspondence 18 19-35. 1963. Edired by KG. Davies and A.M. Johnson.

The Hudson's Bay Record Society, London. 415 p.

Beclq B., T.G. Smith, and A.W- MansfieId. 1970. Occurrence of the harbour seal, Phoca

vitdina, Linnaeus in the Thlewiaza River, N.W.T. The Canadian Field-Naturalist

84:297-300.

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Figure 1. Known range of Photo vir~lilt4 meZIonae in relation to Hydro-Que'bec's proposeci

Grande Baleine hydroelectric project.

AREA

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