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FINAL REPORT

DISTRIBUTION AND ABUNDANCE OF HARBOR AND GRAY SEALS ACADIA NATIONAL PARK AREA

Contract between the National Park Service and Maine Department of Sea and Shore Fisheries Contract No. 14-10-9-900-380

Period Covered - July 1, 1971 to July 1, 1973

Report Prepared by: DavidT. Richardson., Principal Investigator TABLE OF CONTENTS

I .. Introduction 3

II - Present Abundance, Distribution ------6 Abstract ------6 A. Study Area Description ------8 B. Methods ------'------11 C. Results: Aerial and Boat Census Methods ------13 D. Discussion ------23 E. Population Analysis ------...:---- 27

III - Past Abundance, Distribution ------30

Abstract ------30 A. Methods ------31 B. Results 31

IV - Behavior, Reproduction ------34

Abstract ------34 A. Behavior ------35 B. Reproduction ------37

V - Seals and Man ------41 Abstract ------41 A. Hunting Habits: Relation to Conunercial.Fishing --- 41 B. Food Preferences ------43 C. Population Limiting Factors ------44 D. Protection of Marine Mammals ------49 E. Recommendations ------50

Acknowledgements ------54 Literature Cited ------55

References ------57 Appendices: ' .. 1. Aerial Surveys (raw data) 2. P. L. 92-522 (Marine Mammal Protection Act of 1972) ... , «~i'2i'2 &1! I"~

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I - INTRODUCTION

A two-year study of Harbor seals (Phoca vitulina) and Gray seals (Halichoerus grypus) on the Maine coast in the vicinity of Acadia National Park was initiated "in July 1971. The study was funded by the National

Park Service, U. S. Department of Interior, Contract Number 14-10-9-900-380, I . and was contracted through the Maine Department of Sea and Shore Fisheries under the guidance of Marine Research Director, Robert Dow. David Richard- son was hired by the Department as a research biologist to carry out the study. The principal goals were: 1. To determine past distribution and abundance of the two seal species. 2. To determine present distribution and abundance, including a determination of feeding areas, resting or sleeping areas, whelping sites, and other significant seal habitat. 3. T·o find reasons for changes which have occurred in distribu­ tion and abundance, if any. 4. To determine if legislation is needed in order to protect present and future populations of harbor and gray seals and, if so, draft a bill which would adequately protect the two species. 5. To determine what opportunities now exist for park visitors to observe seals. 6. To provide a detailed explanatory· listing of research needs concerning the ecology of seals in the vicinity of Acadia National Park. In January, 1972, the first semi-annual report was submitted detailing the results of the first fifteen week summer field study. This report described in detail the study area, seal habits and ecology and factors thought most important in limiting the populations of seals. This first report also summarized the findings of two aerial surveys and boat surveil- ,. I -4- I

lance techniques used in estimating the population and distribution of seals in the waters about Acadia National Park.

I In July~ 1972, the second semi-annual report was submitted covering t ,f studies completed during the period January 1 to June 30~ 1972. This report detailed observations made of seals during the whelping period of mid-May ! to mid-June and summarized the findings of three completed aerial surveys j of the study area. The July report also summarized a proposed research r study of the hunting and feeding behavior of seals and its relation to commercial fishing and an expanded effort to census the seal populations of the Maine coast. The author is currently conducting these studies with support from the National Geographic Society and the Maine Department of

Sea and Shore Fisheries. The third semi-annual progress report gave results of a thorough _aerial census completed in July, 1972 and also summarized census and dis- ). tribution data for the two summer field periods. Estimates of pup production

were revised using fecundity~ pup~ and year-class mortality data from popula- tion studies of harbor seals carried out in the Canadian Maritime Provinces and in British Columbia. Updated information related to the past abundance of seals in the Acadia waters gained through personal interviews, a formal questionnaire, and extractions from a naturalist's field notes were dis- cussed. Finally, protection and management of Maine's seals afforded by the recent Marine Mammal Protection Act of 1972 were interpreted. This final report restates and discusses data pertinent to the afore-

mentioned study goals regarding seal habits~ ecology and populations. In addition to presenting salient findings of the two summer field periods, this report recommends future research and encourages further use of wild­ life interpr-etive programs to heighten public awareness of marine mammals

occurring in the area of the Park. rI~ -5-

This report presents new data regarding the seasonal distribution of

~ray seals on the Maine coast and results of an aerial census of Blue Hill Bay on June 8, 1973. Given recent census flights of the coastal embayments and shoreline from Kittery to pemaquid Neck, some preliminary remarks are given regarding the relative importance of the waters about Acadia National Park as desirable breeding and resident habitat for harbor and gray seal populations. Each section of this report will begin with an abstract of pertinent findings and will then present the methodology, findings and analysis supporting that summary. -6-

II - PRESENT ABUNDANCE, DISTRIBUTION

Aerial and boat censuses of the waters about Acadia National Park, Mount Desert Island, reveal that 500 square nautical miles of water in Blue Hill, Frenchman and Jericho Bays and their approaches support an esti­ mated resident population of 1600 harbor seals and an estimated seasonal population of 40 gray seals.· An average density of 3.2 seals per square nautical mile and 26.3 seals per haulout area is estimated for the study area shown in Figures 1 and 2. Stated another way, there are 39.5 seals per linear nautical mile. The estimated numbers of seals and percent of the total population by regions are as follows: 484 or 29.7% in Blue Hill Bay north of Pond Island and Bass Harbor; 279 or 17.1% in the waters of Frenchman, Flanders and Sullivan Bays and the Skillings River; 127 or

7.8% in the waters and bays east and north of Schoodic Peninsula to Petit Manan; 720 or 44.1% in the Swans Island and Isle Au Haut archipeligo and Jericho Bay; 22 orl.3% in the waters of Somes Sound. The seasonal dis­ tribution of seals derived from four aerial censuses (Table 3 and Figure 3) indicates an apparent decline of approximately 9.~~ in both Blue Hill and Frenchman Bays during and following the May-June pupping season, and a con­ current increase of 16.4% in the numbers of seals in more offshore waters. Applying findings of Canadian maritime and west coast studies of harbor seals, it is estimated that from a postwhelping population of 1600 seals in 1972,20.47% or 328 pups (164 male and·164 female) were born to the study area that y~ar and that mortality rates differ for the sexes by age classes as predicted by the life table given (Table 6). -8-

A. Study Area Description The study area includes all those open waters, bays, and rivers within the boundaries described as follows: (Refer to Figure 2, Study Area Map) Beginning at Petit Manan Point and running to Petit Manan f t Island to a point approximately N 440 22', W 67 0 52', thence

0 southwesterly (on a course 257 ) to a point southwest of Isle i! I Au Haut approximately N 43 0 56', W 68 0 42', thence northeasterly

0 (on a course 74 ) to a point northeast of Isle Au Haut approxi­ mately N 440 08', W 68 0 34', thence north-northeasterly (on a

0 0 course 39 0 ) to Naskeag Point approximately N 44 14', W 68 31'. The study area cited includes all the waters of Blue Hill and Frenchman Bays, the northeast waters of Isle Au Haut Bay, the eastern waters of Jericho Bay, and the smaller Gouldsboro and Dyer Bays east of Schoodic Point.

The study area is approximately 41.5 linear nautical miles at its southern boundary, and the total water area, including upper bays and rivers, is estimated to be approximately 506 square nautical miles. For purposes of analysis, the total study area is divided into five major regions (refer to Figure 2). The hydrography of each is as follows: The Blue Hill Bay region (I) includes all the waters north of the 440 14' parallel (a line drawn from Naskeag Point to a point just north of Bass Harbor Light) and includes the more sheltered waters of the central bay, Blue Hill Harbor, Morgan Bay, Union River Bay, and the Western Bay east to Mt. Desert Narrows. These waters are protected from open ocean winds and swells and receive warm summer land breezes and greater seasonal fluctuations ~\), ""As 2 . ! ~ 64f'.,,.V~r !" b 0 I C\}fl 4 I ,l) I • J I ~. 9! .1

?~.)"L_I.. ~ /' '" $.1 ./ j,/ til • '/ ('t • ~ • Po "o:J • /' «< .... • • ~ • > ~ • /' • '1 '1 • (D (D • /' • N ":::: )/ PI ." 'd /' c::s ./ /' Arabicnumberals designate all haulouts used by seals during 1971 and 1972 field studies (see Appendix 1)

Underlined numerals designategray;seal . " haulouts. ~ Roman numerals designate regions.

/ " . ./ ./ 108 ./ • fl\T. /' »£~E~T ~ ./ -, /' CoOrt """m d " III' Ill] I 111 t lIZg fill Wi j':;U !,c"Mf'IT-illf=m:I!;'; ;; ==f!::'U=-'""~""'r?;:;::=:::::;::: "!}gi5bF! -10-

in water temperature. The shoreline is moderately populated and there is light boating activity. The Frenchman Bay region (II) includes all waters north of the 440 20'parallel (a line drawn from Great Head, Mt. Desert I Island, to Schoodic Point) including Eastern Bay, Skillings f River, Taunton River, Flanders Bay, and the Winter Harbor area. I This region is influenced by weather factors similar to Region I except that lower Frenchman Bay is somewhat less protected from onshore winds and swells. The Northeast region (III) includes all the open waters and the finger-like bays having feeder rivers and estuaries east of Schoodic peninsula to Petit Manan Island. This area contains 5 seal haulouts expqsed to open ocean conditions and 3 seal ledges within embayments which afford protection and estuarine conditions. Region IV, or the Offshore Islands Region, includes all the waters south and southwest of Mt. Desert Island, including the Cranberry Isles, the Swans Island archipeligo, and Isle Au Haut archipeligo. This region, excepting those waters immediately north of Swans Island (Mackerel Cove and Casco Passage area), is characterized by unprotected and highly changeable weather condi­ tions but more stable colder water temperatures characteristic of open ocean conditions. Seal haulout areas are more remote and thus experience less pleasure boat traffic, land breezes and possi- ble pollution effects. Region V or the Fjord Region, is a small, specialized area including the waters of Southwest and Northeast Harbors west of Sutton Island and the fjord waters of Somes Sound. This area is -11-

densely populated, has much boat traffic, and the waters of the upper sound are not exchanged completely with the tidal cycle. B. Methods Boat Surveillance - Numerical counts of harbor and gray seal adults and harbor seal pups were taken throughout both summers by the investigator, Acadia National Park naturalists and staff, and by Coast Guardsmen on duty at offshore light stations. Observations were usually noted on standard field data cards and then logged on a master chart giving all haul out locations and dates of observations. Separate counts were kept of gray seals. In late August and September of both summers several days were spent covering the more remote offshore ledges for a census of gray seals in the study area. Also, during the 1972-1973 winter several excursions were made to lower Penobscot, Jericho and Blue Hill Bays to determine whether gray seals remained in Maine waters to breed. Aerial Census - All aerial surveys employed the use of a light single engine hi-wing plane flying at approximately 400 feet over coastal shoreline, small coastal islands and halftide ledges. All photographs were taken with a 35 mm hand-held camera using either 52 mm or 135 mm lenses and appropriate polarizing or haze filters. The first census in June of 1971 utilized black and white 8 x 10 enlargements for counting purposes with 2 or 3 frames usually taken of. each haulout location (see photocopy, page~. Aerial surveys taken in September 1971, May 1972, July 1972, June 1973 used high speed (ASA 160) Kodak Ektachrome film. All aerial shots were taken at 1/500th second shutter speed. Developed transparencies were illuminated ;. ~ :. ;:' . .,~

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P.erial Photor:ranh of Colony of Bnsking Harbor Seals Crm>1 Island '. Lcd,,;e, ~·1c;c~ernl Cove, S~.j'ans Island. June 15, 1971 :·;ot.:: ;p~s i:1.(1ic2tCC! )'v arr01;·!S. -13- on a lenticular screen for counting with theaid~ at times, of fiX binocu- lars.

As nearly as possible, time of tide~ sea state and illumination con- ditions considered most attractive to induce seals to haul out to sun and rest were sought in choosing days for aerial census and photography. Low- drain tides occuring in the late morning to early afternoon on days with maximum sunlight and calm wind and sea state were chosen

C. Results: Aerial and Boat Census Methods Table 1 shows the data gained from both aerial and boat census methods including all haulout sites from all regions (regardless of whether some sites may not have been included in one census method or the other). Thus, in Table 1 the values in colurrm "g Max. No. ff refer to the total seals in each region found by each sampling method and calculated by summing the maximum number of seals ever photographed and counted at each haulout site. The values in colunm ,,& Ave. No." refer to the total seals in each region found by each sampling method and calculated by summing the average number I of seals also photographed and counted at each haulout site. I Table 1 also summarizes sampling frequency for haulouts known in each ~ ~ region, indicating the total haulouts where seals were ever censused by each method. The "total census opportunitiesff colurrm refers to the number I of times in the entire two-year period counts of seals were made within I each region by each census method. i Comparing values for total haulouts with values for total haulouts visited by each method, it is seen that only about 50 percent of all haul- outs were visited by boat (except Blue Hill Bay where 74 percent were accessible and were visited). Blue Hill Bay also had the closest fit of maximum numbers of adult harbor seals seen by each census method. -14- Table 1 Summary of Census Methods and Population Estimates by Region Using All Haulouts Sampled for Two Years

1 2 ion Total Census Total Total t Max. No. t Ave. No. Haulouts Method Haulouts Census A=Aeria1 Each Opportunities B=-Boat Method

23 A 19 40 611 432 B 11 53 580 397 --, I 10 A 9 22 490 263 B 5 45 282 111 II 8 A 1 11 211 193 B 4 9 151 104 V 51 A 49 90 1212 923 B 21 89 891'. 689 2 A 2 3 23 18 B 1 4 8 4

1 94 A 86 166 2547 1829 B 54 200 19.18 1311

1. Values are calculated by summing for each region the maximum number of seals ever counted visually or by photograph at each . haulout location. 2. Values are calculated by summing for each region the average number of seals counted by photograph at each hau10ut location. -15-

Comparing the total haulouts for all regions to the total haulouts reached by each method, it is seen that boat surveillance reaches fewer (54 of 95, or 57%) of the total number of haulouts in the study area than does aerial census (which reaches 86 of 94 locations, or 92%). Table 2 compares the abundance and distribution of seals by regions using only those haulout sites which were sampled by both aerial and boat census methods. The decimal values beside population counts are percent of the total population for each calculation method and each census method. Figure 3 compares by histogram the distribution of seals by regions (sampled by both methods) using values for the average number of seals per haulout summed for each region. Except for the Somes Sound region with only two haulout sites, all regions vary by twenty percent or less with respect to sampling method. Table 3 summarizes the raw data (given in Appendix 1) for all aerial surveys conducted to date and gives by region the number of haulouts, adults and pups counted. Comparing the July survey with the three prior surveys, the number of seals rose substantially to 1395 (see discussion to follow). Figure 4 plots the changes in distribution of seals by region from May to September, combining data from aerial surveys of both summers. Values are expressed as percent of the total population for each region on each aerial census date. An overall trend appears to be a net increase of seals of about sixteen percent in the waters of the offshore islands (Region IV) from late May to September. Blue Hill Bay and Frenchman Bay fluctuate but show a net loss of about nine percent over the same four- . month period. It appears that the bays may lose as much as ten percent of their population to the offshore waters during and following the f Table 2 -16-

Comparison of Seal Distribution by Regions Using Two Sampling Methods. Hau10uts ·are those Sampled for the Two Year Period.by Both Methods.

Lon Total Census t Max. No. t Ave. No •. t Max. No. Hau10uts Method Gray Seals A=Aeria1 B=Boat· * * 13 A 484 .29.7 308 27.2 --- B 535 30.0 352 27.6 0

r 4 A 279 17.1 182 16.1 B 267 14.9 173 13.6 4

II 3 A 127 7.8 112 9.9 B 148 8.3· 98 7.7 2

V 23 A 720 44.1 513 45.3 B 824 46.2 647 50.8 33

1 A 22 1.3 17 1.5 B 8 0.45 4 0.31 0 .-.---- ;al 44 A 1632 1132 --- B 1782 1274 39

* Values in columns beneath asterisks are percent of the total population. -17-

Comparison of Seal Distribution by Regions Using Two Sampling Methods.

~ Aerial 0 Boat s:: 0 •.-4 ~res 50 r-I ::s Cl. 0 Il,. r-I ..,res 40 0 .. E-4 i! . )'. ~ 0 .j.)s:: 30 Q) () ~ Q) Il,. 20

10

Region I II III IV ·v'

Figure 3. Values are percent seals for each region of the total population . calculated by summing the average number of seals per haulout sampled by both methods. .-t'able 3 Changes in Distribution of Seals by Region

Aerial Surveys - Isle au Haut to Petit Manan July 5,6 Average Region Census Date: June 15,16 Sept. 30 May 23 1971 1971 1972 1972

Percent total population * * * * * *

I Blue Hill Bay 9 No. Haulouts 11 6 9 243 27.5 199 21.4 293 29.8 263 18.9 250 23.8 Adults (36) Pups (52)

II Frenchman Bay 2 7 7 No. Haulouts 7 17.8 94 10.6 123 13.3 218 22.2 312 22.4 187 Adults (19) (11) I Pups I-' co I III Northeast 4 2 4 No. Haulouts 6 100 9.5 136 15.4 84 9.5 80 8.1 83 5.9 Adults (2) Pups (0)

IV Offshore Islands 21 18 32 No. Haulouts 22 504 47.9 377 42.7 522 56.3 392 39.9 726 52.0 Adults (11) (44) Pups

V Fjord 0 0 1 . No. Haulouts 2 0.8 11 l.0 33 3.7 0 0 0 0 11 Adults 0 0 0 (0) Pups 43 Total 33 36 53 No. Haulouts 48 1395 1047 883 928 983 Adults (74) (101) 88 Pups Parentheses indicate values known to be conservative counts

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• Changes in Distribution of Seals , . v i I By Region, May to September, :1972 . i 1

60 -----­je16.4 . ----- 50 ,,--- , . I " IV .".,,1 40 -,'

s:: 0 I ..-I 30 4.l Cd r-I ;:J 0. 0 ~I .... .-8.7 Cl... 20 , -.. .. , c .. ,.,., ,, , ...... Cd \ I ..... 4.l , .' I ..- . -- 8 0 ,. '. I ---_ - .9 E-; .~ ... • .",, I. 4.l .. " e. s:: 10 III···· · -•.

Figure 4.

Values are percent of the total population for each region on each aerial census date. Refer to 'Table 3 for data. 1

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~r ~ I, whelping period. On the other hand, it is possible that sampling error , ' I, 11 '; 'I I' between regions could account for some or all variance shown between Iif ~ Ii , . I sampling (census) dates. ~r 'i J It Table 4 relates the density distribution of seals per square nautical ,: L• ;1 mile and per haulout by region using the July 1972 census data. The I! !Ii 1 ~ , t: I approximate water area and percent study area values for each region were I u I gained by summing graph squares within each region and the total study area. :~ The density of seals in Frenchman Bay is shown to be highest both in terms I of water area and haulout sites chosen. Region III, east of Schoodic peninsula, is shown to have a low density of seals in terms of water area but when viewed in numbers of chosen haulout sites, the density appears comparable to other regions. Densities in Area V, Somes Sound, being based on only one colony, should not be compared with other regions. It appears from Table 4 that seals are likely to be found in comparable densities in all regions during July; colonies being most likely to number twenty seals or more.

The two seal species differ in their distribution in the study area. The harbor seal is found on all locations cited, whereas the gray seal chooses only the remote, offshore ledges on which to sun and rest. No gray seals were seen on the ledges in central or upper reaches of any bays.

The study area map (Figure 2) indicates gray seal haulouts by underlined , " i arabic numerals. Over the two-summer period, 40 gray seals (summing the I 1 J maximum seen at each haulout for both summers) were viewed at 12 locations. i .~ ] Gray seals haul out in close proximity to harbor seals. Several gray seals at one site can be grouped or scattered among many harbor seals. No gray seals were found in the study area during the winter boat surveys. Both species prefer half tide ledges or small islands of low profile and good visibility to surrounding waters. Such sites are usually well away DENSITY DISTRIBUTION OF SEALS BY REGION

Densities Number of Seals 'Per Sq. Per Haul- Approx. Water Area percent of July 1972 Aerial Number of Nautical out Region Sq. Nautical Miles Study Area Census Haulouts Mile

I 80 15.8 263 9 3.3 29.2

II 61 12.1 312 7 5 .. 1 44.6

III 97 19.1 83 4 0.9 20.8

265 52.3 726 ,32 2.8 22.7 IV I N t-' I V 4 0.7 11 1 2.8 11.0

• Total 506 1395 53 Average 2.76 26.3 ! 11

-22- from populated sections of the mainland or islands. Preferred ledges are of gentle slope, seaweed covered, and remote from intense boat traffic. During a visit to the study area in February 1972, fishermen and residents of the Mt. Desert Island area confirmed that seals had been seen in the upper bays until mid to late November. ,However, during the winter months seals are not often seen by fishermen and usually those spotted are swimming. It is likely that seals slmply avoid the cold air temperatures and stay almost exclusively in the water during the months of December through March. It is likely, also, that offshore movement and a more pelagic winter existence is advantageous to seals for hunting fish stocks which are known to leave inshore waters in the late fall and winter. In certain waters of the study area, however, seals were viewed throughout the winter by a Sorrento resident. At the mouth of Sullivan

Harbor and in Flanders Bay seals were seen all winter. Following a bay freeze-over in late February, seals were noted to haul out on the edge of the ice and would often drift out into the open parts of Sullivan Harbor and Frenchman Bay. Only one or two seals were seen occup'ving an ice floe at anyone time and movement off or on the floe was not observed. Table 5 shows the most recent results of an abbreviated aerial census of the study area compared with the ~ ~ coverage of the past two years. On June 8, 1973, a census of Blue Hill Bay and Frenchman Bay covered all of the ledges in Regions I and II of the study area. l/

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d,:~ Table 5

Three-Year Comparison* of Blue Hill and Frenchman Bay Seal Populations

June 15 Sept. 30 May 23 July 5 June 8 1971 1971 1972 1972 1973 Blue Hill adults 243 199 293 263 305 Bay pups 52 94

Frenchman adults 50 123 169 229 170 Bay pups 9 40

*Comparison based on partial coverage of the study area

D. Discussion Seal colonies tolerated well one or two passes by the plane over the ledge at approximately 400 feet. During each census, photographs from perhaps one or two of fifty total locations would show seals leaving the ledge. Yet these seals were usually easily counted in the water, particularly in shallow cove or shoal areas. Although Vaughn (1971) reports that count- ing is facilitated during times other than near high noon when shadow casting by the subject provides contrast, his background is one of light colored sand on the beaches of the Wash (Lincolnshire), England. Counting of seals on the irregular and dark seaweed-covered ledges of our coast did not pro- vide this advantage. It was found during morning and afternoon flight times that approaching the ledge facing the sun was poorest for taking photo- graphs due to light reflectance from the water. However, approaching the ledge facing the sun did prevent the plane from casting a shadow over the ledge (a possible frightening stimulus). Subsequently, the plane could circle at right angles to the sunTs oblique rays followed by a short inter- val with the rays behind the plane affording more time over the ledge and best lighting conditions for photographing. , Ii

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Boat and aerial surveillance experience during the two summers indi- ," I I' , t cated that intense sunlight and warm temperatures are first-order induce-

ments for seals to haul out to rest. Under these conditions they also seem most reluctant to leave the ledge for cold waters. Census activity

during these favorable weather conditions proved to be more successful.

Seals are also most reluctant to leave the ledges just prior to and

I' 'I during the molting season in late July and August. At this time seals are ,1 quite lethargic, are in a state of fasting and seek resting ledges during the high tide hours. During this period of hormonal stress, captive seals

have been shown to have lowered resistance to infection, to act in peculiar and unfamiliar ways and in general to be Tfpathologically sickTf (Geraci, J., DVM" personal commun..ication). It is possible that during late July and

August when molting occurs seals are most reliably seen sunning and resting

and remain visible longer for aerial census purposes.

Table 1, comparing aerial and boat census o~ sampling methods, reveals

that either by taking maximum counts, or by averaging counts over the two- year period, aerial and boat census methods compare closely. Maximum number counts taken by boat for the whole study area are 9.2% greater than those taken by aerial census. Average number counts taken by boat are

12.3% greater than those taken by air. These variances would seem to be

within probable sampling error for this study. Thus, in the case of censusing gray seals where viewing the profile

of the head is crucial to identification, reasonable confidence might be placed on boat census values if sufficient surveillance opportunities or , sampling frequency is pursued. Table 1 presents a summary of data collected from ,all haul out sites

for the entire two-year period. Thus, the maximum count of 2547 seals is, -25-

as expected, high. Such a total does not account for distributional changes but rather disregards the sampling date and shows only the sum of the maxi­

mum count taken at each haul out. The average number counts for aerial

census shown in Table 1 are, likewise, expected to be rather high since on any given day seals would not be expected to be utilizing all 86 haul outs. However, the total average value for boat census shown in Table 1 repre­

sents fifty-four haul outs (comparable to both the June 1971 and July 1972 total haul out counts during aerial sampling) and gives a total population of 1371 seals. This compares closely with the July 1972 aerial census giving a total of 1395 seals.

Several aspects of the July 1972 census make it the most thorough and reliable of any conducted. The flat calm sea condition (see December 1972 progress report, Appendix 1, Photo 2) provided opportunity to test the hypothesis that the entire seal population hauls out to rest and sun on the low tide. During the entire two-day census no seals were sighted in the water. Visual observations and photographs at oblique angles and directly over open water and close to ledges having large seal colonies showed no seals present in the water, submerged, or at the surface. This was true even though many vertical shots show clearly the subtidal topography and buoys, birds, and other floating objects. In two instances where seals were

ITspooked lT from ledges by the repeated passing of the aircraft over the ledge, aerial shots clearly show the seals diving, submerged, and swimming at the surface.

Another factor making this aerial census more thorough than previous ones was the use of a small two-seat seaplane with a slower air speed, allowing more maneuverability and time for vertical shots over the seal ledges. as

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It is with some confidence, then, that the investigator adopts the

important assumption that seals strongly select low tide hours to rest and : i sun, and that for census purposes the entire population is visible during favorable weather conditions. This assumption is also made by Van Bemmel

(1956). This judgment is further reinforced by boat surveillance where uniformly large seal colonies are observed hauled out even during foggy or overcast low tide hours with attendant colder temperatures as well as j . during sunny, warm, low tide hours. In addition, persons living near seal

ledges often report snarls, growls, and boisterous social communication among large herds of seals hauled out at night.

In summary, seals appear to be opportunistic, utilizing the low tide

hours to rest under most weather conditions, day or night. This assumption does not hold for winter months with cold air temperatures. Due to thorough area coverage and photo counts during the July 1972

aerial census, a weighted average seems more realistic than the straight

average of 1047 seals given in Table 3. As shown in Table 5, the numbers of seals in both bays have not changed

greatly since the census of May 1972. The apparent increase compared with the two 1971 surveys is thought to reflect more thorough and experienced flight coverage and photography in later censuses. An actual increase in seals in the two-year period may, however, be a component of the 25.6%

increase in Blue Hill Bay. The very low number of 50 seals seen during the first census of Frenchman Bay cannot be adequately explained. Given some shortcomings in both census or sampling methods, one is still prompted to choose a lTworkinglT figure for the study area population. The July 1972 postwhelping census of 1395 seals is probably a close approxi­ mation of the population. Yet, it may be somewhat conservative since only -28-

Bigg speculates that increased mortality for males corresponds with attainment of maturity and that reproductive activity such as males fighting during the breeding season (males have been noted by this investi- gator to be scarred about the neck and chest area) might contribute to increased mortality.

This differential mortality by sex may be in error, however, as Bigg derives it from his shot sample of 245 seals and assumes that it represents the age structure of animals dying from natural death. This may be so or it may be that mature males, by virtue of different hunting habits, curiosity for man's activities, or general gregarious behavior could be favored more strongly in a shot sample than in an age-frequency sample dying from natural causes. Bigg does attempt to lessen sources of known or suspected bias by eliminating from his shot sample the young, less wary seals and instances where collection methods involved suspected bias by age or sex.

Using data from Bigg's (1969) studies requires an assumption regarding similar habitat, whelping sites and rates of pup and adult mortality. The Maine rocky coast habitat may have some hazards that the sand beach Sable Island whelping area does not have. Tidal reefs and sand bars, east of

Vancouver, B. C., are common haul outs for harbor seals in Bigg's study. These haul outs seem to compare closely to whelping sites in the bays of the Acadia study area.

Using Bigg's and Boulva's data for recruitment rate and Bigg's data for mortality rates by age and sex class, Table 6 was constructed as the most likely approximation of the Acadia National Park seal population structure. This life table assumes that from a post whelping population of 1600 seals, 20.4% or 328 pups (164 female and 164 male) are born each year and that mortality rates are markedly higher for mature males than for females (29% versus 15% after age 5). --e 6 -29- Life Table for Harbor Seals in Acadia Area

Females Males Total 5 *Mortality Total Females *Mortality Total ~ales Individuals Per Year Class Per Year Per Year Class (% ) (%) Class

20 164 21 164 328

20 131 2l 130 26l 20 105 21 103 208 20 84 21 81 165 -20 67 .-ll 64 131 15 57 29 45 102 15 48 29 32 80 15 41 29 23 64 15 35 29 16 51 15 30 29 11 41 15 25 29 8 33 15 21 29 6 27 15 18 29 4 22 15 15 29 3 18 15 13 29 2 15 15 11 29 1 12

15 9 29 9 15 8 29 8

15 7 29 7 15 6 29 6 Total 895 693 1588 'om Bigg (1969) ;sumes a total postwhe1ping population of 1600 and a recruitment of 20.47%. -30-

The life table' (Table 6) gives an approximation of the age structure which nearly yields a balanced or stable population. The author's conclu- sion from surveys of the population covering three whelping periods is

that this prediction of a stable population should be a short-term working hypothesis. Small yearly increments or decrements to the population may be disguised by such changes in census technique. Likewise, impressions

of changing numbers of seals given by residents, fishermen, or seal col- , !: I i I lectors (see Section III, Past Abundance, Distribution) may reflect distri- ,I Ii :1 ! I! butional changes in the population occurring along the cruising route, near , I' : i' the, familiar fishing grounds or in the TThunting area" of these observers.

III - PAST ABUNDANCE, DISTRIBUTION Evidence for the historical abundance and distribution of seals on

the Maine coast is sketchy, difficult to assess objectively, and conflict- ing. Interviews were conducted with fishermen, permanent and summer residents, two veteran seal collectors, and Sea and Shore Fisheries wardens

in the study area. Questionnaires attempted to assess past and present abundance, distribution, habits and ecology of seals. Attitudes regarding the above people's interactions with seals were gained. Accounts of seal observations were sought from local and university libraries. A natural-

ist's field notes were extracted for seal counts dating from 1941 to 1950. A quantitative estimate of past seal abundance and distribution in the Acadia area is not possible to make at this time. Field notes of the late

Dr. Alfred O. Gross taken in the area of Boothbay Harbor 30 years ago indicate that seal abundance and distribution remains similar to that

observed in recent years. -31-

A. Methods During bath summers, investigatian af the past abundance of seals has been conducted by interviews with fishermen, village and lang-time summer residents, two veteran seal callectars and area Sea and Share Fisheries wardens. A questiannaire was distributed to fishermen during the summer of 1971 and a mare comprehensive questiannaire (see December 1972 pragress repart, Appendix 2) was given to. fishermen, residents, seal callectars I i and wardens during the summer of 1972. Dr. William Drury kindly made ·1, i: I available extracted counts of seals which were made an field excursio.ns by II ;1 the late Dr. Alfred O. Gross af Bawdain College. Grass's abservations were made during the decade 1941 to. 1950 at numeraus offshore ledges along the caast. They represent the only quantitative observatians faund by this investigator. Local and university libraries were searched far any refer- ence to Maine's seals. A shart news accaunt of the study was placed with the Bar Harbor Times and the Ellswarth American requesting information fram local histarians and residents regarding personal knawledge of seal abundance in the waters abaut Acadia National Park.

B. Results Person to. person interviews yielded an interesting canflict of apinion amang respondents regarding past abundance af seals. Mast fishermen feel that seals are as abundant now as in past years and even decades. All fishermen interviewed were quite accurate in citing ledges where seals cauld be regularly found; however, in same cases numbers af seals at spe- cific locatians prov.ed to be averestimated. Many village residents of Mt. Desert Island stated that seals have declined in numbers at certain haul auts where in the past they were -32- easily viewed from land. Similarly, long-time summer residents often cite locations near their homes where they no longer see seals sunning. Since seals are of considerable interest to both full-time and seasonal residents, one is inclined to find such observations credible. It is not known, how- ever, whether such instances of seals having deserted previous sunning ledges reflects a decline in total population or simply an avoidance of waters which have become too populous with summer visitors, boat traffic, and harassment. Such local changes may be temporary or may reflect changes in numbers of schooling fish now coming to inshore and harbor waters.

Two Bar Harbor residents interviewed gave interesting accounts of past seal,sitings but no comparisons to present abundance. One former steamboat captain, now 96 years old, remembers the "bay seal" as never very abundant nor varying much in numbers back in the 1920's.

Another Bar Harbor resident recalls the winter of 1917-18 when French- man Bay froze over and "hundred of seals - possibly a thousand" hauled out on ice floes. She recalls that the steamer had to continually cross the

Bay to keep a passage open and some floes were "black with seals. TT Two interviews of importance concerning past abundance of seals were conducted with two seal collectors who for many years in the late spring have combed the ledges of the Boothbay Harbor region and the Blue Hill Bay region for seal pups. One collector lives in West Boothbay Harbor and for 25 years has taken seals from the ledges of Southport Island, the Sheepscot and Damariscotta rivers. In more recent years he has had difficulty ,meet- ing his usual quota of approximately 40 seals (mostly pups) and has had to set nets near ledges in Casco Bay off Harpswell Sound. He cannot give definite reasons for the decline in seals in the Sheepscot River and is unsure whether it is an actual population decrease or a movement of once -33-

larger herds out of ·the area to other locations. He does allege that seal

collecting for many years in Casco Bay has not affected that local popula­

tion. A healthy seal pup could be sold last year for fifty dollars, includ-

ing crating and shipment, to destinations as far as Cleveland, Ohio.

Another seal collector of Tremont, Mt. Dese~t Island, has collected

seals in Blue Hill Bay for over ten years. His most effective method has been to approach a ledge quickly in a motorized canoe and capture by hand

or throw-net the pups and drowzy adults before they can leave the ledge.

He estimated the present seal population in Blue Hill Bay and offshore

islands (Regions I and II) to be about one thousand seals. His estimate was remarkably astute; correlating respectably with the 684 seals spotted in this same area in the June 8, 1973 aerial survey. However, he is con- vinced that the number of seals has declined 50% in the last ten years.

Like most fishermen interviewed, this collector feels that although occa- sional shooting of seals does occur, it is not as significant a factor as in past years. He feels that more seals have been taken from the area in recent years by out~of-state zoological societies, amusement and animal park collectors, and scientists seeking research specimens. He also cites an increased incidence in the past several years of abandoned pups; many having skin lesions or sores which may be bacterial" or viral induced (Wilson and

Sweeny, 1970, and Wilson et.al., 1972). Field notes from the late Dr. Alfred o. Gross, formerly of Bowdoin College, give the only quantitative field data of seals in Maine waters found to date by the investigator. Dr. William Drury of the Massachusetts

Audubon Society extracted counts of seals which were made by Gross while censusing shore birds in the decade 1941 to 1950 and kindly made these available to the investigator. GrossTs data is most useful in the approaches

,! •

-34- to penobscot Bay and in the Boothbay area where comparisons of past and present seal counts are possible. Seal counts in the Mount Desert area were too few to make comparisons with counts from this study. Counts made by Gross in the Boothbay Harbor area such as Pumpkin Ledges (200 seals) and Hypocrites Ledges (25 seals) slightly exceed t~e total of approximately 175 seals censusedat those locations by the investigator in November, 1972. Local residents and amateur historians responded to newspaper requests for information giving accounts of where they now were able to see seals. No definitive historical records or field notes of past abundance or sea- sonal changes in distribution were received. Local and university libraries yielded no quantitative data regarding seals in the Mount Desert Island area. Some accounts of seal sitings on the Maine coast during whaling and commercial trading were read in Allen's (1880) History of North American Pinnipeds. However, no information use­ ful in deducing the abundance of seals was included. In summary, memory accounts and limited field counts by one naturalist cannot substantiate an impression of past abundance of the Acadia seal population. The author speculates that the harbor seal may have occurred in greater numbers at near-shore haul-out sites in past decades of this century. It is not known whether the total numbers of seals in the study area has increased, decreased, or behaved cyclically during this century.

IV .~ BEHAVIOR, REPRODUCTION The harbor seal is a resident species to Acadia waters. The gray seal is migratory and arrives in Maine waters during spring months following a late winter whelping period in maritime waters. The two species share more remote offshore ledges; inshore and bay ledges are occupied only by harbor ....

I 'i i!1 -35- J

seals. Harbor seals whelp on ledges during May and June, the cows choosing to haul out at some distance from males and juveniles of the herd. Reaggre- gat ion of sexes and age classes appears to occur during the physiologically

stressful period of molting in late July and August. Mating occurs in the water following termination of lactation in late June-July. A delayed implantation of the embryo until September is followed by a seven and one-

half month gestation period. New-born harbor seal pups are capable of

swimming immediately. The mother-pup relationship is characterized by frequent affectionate nuzzling and biting actions in water, frequent nursing on land, and persistent teaching of the pup to swim and to dive with the mother. Pups are weaned within several weeks, having doubled their birth weight of

11 kg. (25lbs). Behavior is closely tied to the tide cycle, with resting, sleeping, grooming, and most social interaction occurring during low tide

hours, and with feeding and some resting occurring on the high tide hours.

Feeding appears to be opportunistic, favoring readily available finfish species in waters near the haul out ledge to which individuals regularly return.

A. Behavior The habits of the harbor and gray seals are closely tied to the tide

cycle with resting, sunning, sleeping, and presumably most social inter- action occurring on the falling, low and early rising stages of the tide. II Hunting activity and some resting behavior occurs on the high tide. Regard­ i•'I iii, less of time of day, approximately at midfalling tide seals appear, usually

alone but sometimes with one or several others, begin to undulate their way onto the most prominent points of a chosen ledge and are often heard making grunting sounds (of annoyance?) as another seal attempts to crowd onto the ...... ------

-36- limited ledge space. Sometimes the larger adult seals will flay their heads against the neck or back of a seal attempting to climb on a crowded ledge, presumably to discourage him from doing so. Large adult seals are often perched in higher, more vulnerable locations on the ledge, possibly acting as sentries or exercising their status in a "pecking order." These large bulls were usually the last to leave a ledge when the investigator slowly drifted toward the ledge. Upon "spooking" into the water, however, harbor s.eals demonstrated remarkable curiosity for the boat and intruder. As if now assured of their mobility and safety in the water, they would circle the boat and surface as close as 20 feet away, stare, and often playfully dive and splash near the surface. Such behavior reveals the sealTs apparent understanding of manTs long-standing predatory role, yet substantiates their curiosity for manTs activities.

During the low tide hours when weather is sunny with warm temperatures and calm sea sta~e, seals are rarely in the water. Occasionally a seal will leave, swim about the ledge and soon crawl back out to rest. On hot days seals can be seen waving rear limbs while arching their bodies in a crescent form. This presumably aids in heat dissipation. A return to the cold waters may be necessary in preventing heat prostration caused by the superior heat retention of blubber and hair pelage. Occasionally seals will leave the ledge to playfully chase one another and "porpoiseTT or jump clear out of the water. Two seals may chase and nip at each otherTs rear limbs with tumultuous splashing and snarling. Rather uneventful hours of sunning and resting, interrupted by frequent scratching of lice and mites and the watching of passing boats is followed by the flooding tide forcing arched seal bodies at successively higher points r

-37-

on the ledge to leave and hunt for food until the ebbing tide again reveals the preferred ledge.

The high tide hours are occupied predominantly by hunting for food. However, during July and August when molting occurs seals are very lethargic,

are thought to fast, and often seek small islands !or high tide haulouts.

During August of 1972 several gray seals were observed and timed as they dived and surfaced near a ledge on the high tide. Consistent submerged times of 13 to 14 minutes and surface times of 15 to 30 seconds indicate

that they probably sleep and rest by effortlessly descending and surfacing in one spot. Harbor seals are known to behave this way in captivity and

no doubt spend time during high tide hours resting on the bottom. Harbor

seals, when surfacing from resting or foraging activity, usually quickly look , about to orient and then raise their nose skyward for several seconds. This is probably reflexive behavior to check for danger nearby and then to more efficiently hyperventilate with straightened trachae and reduced

chances of water or spray entering the nostrils.

B, Reproduction

Seasonal changes in distribution of seals from more offshore ledges to upper bays during the whelping season probably reflects a preference for the more protected bay ledges as whelping sites. Section II of this report

presents evidence for such a seasonal distribution change (see Figure 4).

Aerial census has shown whelping to occur throughout the study area, but greater numbers of pups are born among large groups of seals in Morgan Bay near Jed Island, Western Bay, near Green Island, Flanders Bay, on Halftide

Ledge, Skillings River, on Great Ledge, and on several ledges near York Narrows, Swans Island, and near Johns, Opechee and Eagle Islands in lower , Blue Hill Bay. These active whelping areas might be called "nursery" areas -38- and deserv"e protective surveillance during the pupping season. These areas are indicated on the Study Area Map C, Figure 5. Harbor seals whelp on the ledges during May and June with the peak of pups born in late May and early June. The utilization of a greater number of ledges in a given haulout area was noted during both summers during the pupping season. The pregnant females and cows with pups choose to haul out together at some distance from males and juveniles of the herd. Consolida- tion of the herd appears to take place as pups are weaned. Following termination of lactation in late June-July mating occurs, presumably in the water as copulation has not been observed on land. A delayed implantation of the embryo during the stressful period of molting in late July and August until late September is followed by a seven and one-half month gestation period (Fisher, 1954). Detailed field notes were taken during May, 1972, of behavior shown by approximately ten mothers and pups just following birth, during nursing and resting hours on the ledges, and while swimming. Although too lengthy to detail here, the following points summarize these observations: 1. Mothers appear confused at the sight of the newborn pup but

quickly adopt possessive behavior and ensure that the pup interacts solely with herself. The mother ignores the placenta she expels minutes after birth on the ledge. It is devoured by gulls and ravens or is washed away by the tide. It is possible

that only following weaning do pups interact with one another. Mothers remain aloof from other nursing mothers but not from pregnant females. 2. Pups are large, relative to other mammalian offspring; approxi- mately 25 pounds or 10 to 15 percent of the mother's weight. ~'LL~Q\J crJrl~~ l~C'4S ' on ~ 64!..fI7- C ...... S~~· l" ~ fS(J l>

""- 7t~ ][ , .to',, CJ'J I ~ ~.e.. AS c:: /fS(. '!""V"\ · ./ ~ ~., · · ,/ .C) '" · ~' I r;;;:;,.C ·• M .fI:~ fl\ ' · ./ !:P (~""" iIi,..' /' ~- '-)~ ~ .' ·:/' '"tI i .-/ n . ~o !"'. IS[ q~]A \ \a /" . ~~ ./ \ ...... 1 ~ qt ./ \ •.1 /. ,""..,' • A': frN-- :: .. ./' ,0 11 ~/' _.-----._.'- Naturalist-Guided Cruise Routes o'.. , ~ /' I / Significant whelping areas. ./ Top number indicates pups counted on ./ July 1972 aerial census; bottom number d'~ on June 1973 census. /" ~ /' /' ./ D MT. ./ Pi:~E(2.T ./ !Zoc.":'

,- ""''''''',,:;:v..;.:."~.~.-...-~,_~ ~. """"".-;':",..c,.; ••• ;.,., .... : .-" .•.. :.~' ,. ,. ··,~·:~~~s-:~·:~;~~?-~??~~~~~~~~~~~;~~~-;:~~:-~~~·~~-·:~·~~=-.. -~~:~;::~ ::==.:~?~:~.~~.~.:~~~;:~;~~~-C~=::==-==~.~ >~~"""""''''''''t<:'} ...... ''''q=)''9' •.;.;.....;..:.,~

" ..-_._. --.~----.--:-----~ ~-. - :"'::':'::':'::':'=:::-.'::::.-'"":. ,,- --

-40-

Pups have loose skin of steel gray color with mixed light spots. 3. Pups have advanced motor coordination at birth; can adeptly swim immediately after birth without aid. However, the mother

usually keeps the pup just behind her head and often buoys the pup slightly in her back and neck while swimming. 4. Mothers and pups display an intimate and affectionate relationship by mouth and forelimb contact while in the

water and on the ledges. A ritual of the pup biting or Ii \ sucking at the neck of the motherTs uplifted head, followed by the mother mouthing the pupTs head is repeated frequently while in the water. This ritual may be necessary for pup recognition when the mother leaves the pup and returns to a crowded nursery. It may also be simp'ly erotic and

affectionate behavior by the mother. 5. The hungry pup gives human-like cries, actively seeks out

the mother and in~tiates the nursing activity by stroking the motherTs abdomen lengthwise in search of the hidden

nipples. Mothers willingly nurse until the pup moves away. Newborn pups appear to nurse at least hourly. 6. Fear of man is learned early from the mother. Newborn seals taken by seal collectors befriend man immediately. Many accounts have been written of pet seals raised and weaned by their captors. When the investigator has approached ledges having newborn pups, the mother leaves quickly and beats the water with her forelimbs and barks to induce the pup to leave the ledge and follow. -41-

I'UPS nurse for se-,':S:'al wee..l..cs during which time they double their

I, II III weight of appro~::-.:.tely 11 Kg. (25 lb.) (Boulva 1971). The pups are

111"ili',ht to fast for SCIT!l:::: period of days following weaning until feeding ''''IIIIVlor is learned fr·::YiT. older pups or juveniles.

" SEALS AND MAN.

A reputation for ~~iding entrained fish and an alleged predation of

':cl/ t lobsters by seal.== :"as maintained some covert hunting pressure by

I i :~hermen. Other pOJ;:"::'ation limiting factors have included capture of

I'lIpS and juveniles fc~ zoos, public display, and zoological and medical

\'\'t-;earch, killing of seals for hides and meat, and possible pollution "I'l'ects. Natural mor-::ality rates are determined by stillbirth, accidents, l including storms ane abandonment), disease, infestation by parasites, aud predation by ha\>.·~s, eagles, sharks, and possibly whales. Opportunities to view seals are most rewarding in the wild. Recommendations are made I'or the Park Service to aid in implementing the Marine Mammal Protection

~ct of 1972 by explaining the Act's protection measures in its interpretive program for the public and continuing seasonal assessment of seal stocks in the study area. It is recommended that the Park continue and expand, dS needed, its excellent support of naturalist-guided boat cruises among the islands and ledges about Acadia National Park. The educational and functional merits of this guided tour of coastal island habitat ensures protection of wildlife and appreciative close viewing of seals by park visitors.

A. Hunting Habits: Relation to Commercial Fishing The investigator is currently studying the hunting and feeding behavior of harbor and gray seals and the impact of seal predation on commercial -42- fishing in Maine. Funds for the study are given by the National Geographic Society and salary support is given by the Maine Department of Sea and Shore

Fisheries. The extent to which the feeding activity of seals competes with manTs interests is a practical research problem and should be investigated for the intelligent implementation of long-term Rrotection and management policies for seals. Knowledge of the feeding habits of seals has been gained during the two summers by literature review and through interviews conducted with fishermen, residents, fisheries wardens, and naturalists and marine mammal biologists. No effort will be made here to review the literature on the feeding ecology of seals. Rather, results of the investigatorTs current research in this area, including such a review, will be later forwarded to the National Park Service. The knowledge gained of seals' interactions with fishing activity in the Acadia area will be reported here. The predominant activity of seals during high tide hours is assumed to be hunting for food. Fishermen consistently relate having seen seals surface with large fish such as sea bass (Centropristus striata), striped bass (Morone saxitalis), cod (Gadus morhua), sculpin (Myoxocephalus sp.), and flounder (probably winter flounder (Pseudopleuronectes americanus).

In the case of flatfish, it is alleged that the fish is worked in the sealTs mouth until it assumes a tubular shape before it is swallowed whole. Large fish are brought to the surface so they may be more easily torn apart by violent shaking of the head, whereas small fish can easily be swallowed whole underwater. Some fishermen temper their enmity toward seals by watch­ ing where large groups of seals congregate. It is here, they say, that productive fishing of herring may be found. They allege that seal herds will move during the spring and summer seasons in response to changing dis- •

-43-

tributions of schooling fish. Thus, popular seal haulouts may act as "markers" for such fishing grounds. Worthington (1964) concurs, stating

.•. "seals, like fishermen, are opportunists, so when fisheries are operat- ing at a maximum, seal activity is also greatest in that area. IT Seals used to raid fishing weirs which were once numerous in the Mount

Desert Island area. One fisherman related that a clever bull would dive several hundred feet outside the leader net, enter the weir and thrash about among the entrapped herring. Following this chaotic feeding, the harbor seal would leave the weir and surface several hundred feet distant from the frustrated weir tenders. Lawrence Jordan, formerly fishing a weir in Gouldsboro Bay, remembers seals climbing up his nets five or six feet out of water to enter and feed on the schooling fish within. It is also told

that seals would chew an entrance hole in lighter nets and feed unseen.

B. Food Preferences Although many fishermen interviewed and returning questionnaires

alleged that seals actively preyed on soft or shedding lobsters, such feed- ing is not borne out by research conducted by Montreuil (1958, and personal communication) nor by attempted feeding to captive seals of whole lobsters

or picked lobster meat.* Montreuil analyzed close to 1,000 harbor and gray seal stomachs and intestinal tracts for intestinal parasites and for evi- dence that seals consumed lobster. He found no instance of pigment or

carapace remains or any hint that lobsters had been eaten during this 10-year analysis. Moreover, he states, ITIn 1956 or 1957, the construction of a causeway joining Grindstone and Amherst Islands closed off a large lagoon, entrapping a small population of gray seals which apparently stayed for at least two years. Samples of these animals obtained as much as 2 years after

*Harry Goodridge, Rockport, Maine, personal communication. r

-44-

their isolation, revealed that although lobster was abundant in the area,

the seals showed no evidence of having consumed any; in fact, the gray

seais in the bay seemed to have subsisted almost exclusively on Fundulus

~. and Gasterosteus ~., which we found to be virtually the only fish

species present in the area." (Personal corronunic~tion).

Fishermen, no doubt accurately, recount seeing seals struggling at the surface with a lobster in its mouth. Montreuil states that fishermen in the Magdalen Islands " ..• readily admitted that the seals seemed to be trying to get rid of the lobster rather than trying to swallow them. I believe it likely that seals may occasionally tangle with lobster while rooting around the bottom and may get their nose or lips caught by one of

these, surface in a panic, and put up quite a fuss while trying to scrape

off the attacking lobster with their flippers. 1T (Personal communication). Such an explanation correlates well with the description given by Bass

Harbor fishermen that seals flay their head as if shaking the lobster and

Tf use their forelimbs to "bend the 10bster and ITcrack it in two. IT Seals have been observed to haul out in upper Bay areas near Boothbay Harbor, apparently following the spring run of alewives. A captive seal

in Rockport (Argosy Magazine, May 1971, Down East Magazine, Sept. 1967) is known to prefer alewives to thawed mackerel. It is possible that the relatively fresh alewives are more palatable. Most studies of harbor seal

feeding habits rank herring high in dietary preference but conclude that seals are opportunists and probably eat a variety of items available in the haul out area rather than forage at great distances for specific species.

C. Population Limiting Factors Natural causes of mortality for both seal species include stillbirth,

accidents (including storms and abandonment), disease and infestation by pi

-45-

parasites. Pups may suffer predation by eagles and hawks. Seals may be attacked by sharks and possibly some whales which infreqently come inshore. But seals, like man, are at the top of the food chain and have few natural

predators.

Boulva (1971) found that 12% of the harbor seal pups on Sable Island died in the first year of life. Van Bemmel (1956) assumed that the harbor

seal in Holland had a 20% mortality in its first year, and Bigg (1969a)

estimated first yea~ mortality to be 21%. As expressed by the life table (Table 6) in Section II, approximately 41% of female harbor seals reach potential breeding maturity but somewhat fewer females survive during years

of peak fecundity in year classes beyond five. For example, only about 13% of the females reach age 12. During the field study, mortality caused by infectious disease or lesions, abandonment by the mother, and attack of a pup by osprey were noted by the investigator. Concern for an increased incidence of bacterial or viral sores or lesions was noted by two seal collectors in the Blue Hill Bay region during the past several years.

Whether naturally occurring or induced by pollutants (see next section),

the collectors state that during the summer of 1971 "several pups would be inspected carefully before finding one free of sores, whereas in past years most pups were free of the lesions with only an occasional one affected."

It is not known whether greater incidence of such lesions is naturally occurring and possibly cyclical or if it may be induced by sewage effluents

(J. R. Geraci, DVM, personal communication). It is likely that ascarid worms, including Porrocaecum decipiens,

Anisakis ~. and Contracaecum sp. keep a strong check on the populations of .harbor and gray seals. Scott and Fisher (1958) reported that in 318 -46- harbor seals and 127 gray seals analyzed, both species were always infected with P. decipiens. An average of 50 worms per harbor seal stomach with no geographic variation occurred in maritime waters. Mansfield (1968) similarly reports a mean of 73 worms per harbor seal in the Maritime population of approximately 5,000 to 6,000 seals (Boulva, personal communication). Scott and Fisher (1958) state that male harbor seals have more worms than female seals and that young harbor seals become hosts of P. deci;Eiens in their first summer. P. decipiens is the familiar codworm. Control of the seal populations in an effort to reduce incidence of the worm in commercial stocks of cod has been suggested in fisheries management papers. However, it is not known that seals are the only intermediate host of this parasite. Control of the seal population for an uncertain decline in codworm infesta- tion is a questionable policy.

Man-induced causes of mortality include: 1. The indiscriminate shooting of seals by fishermen and others.* 2. Commercial fishing activity limiting food supply for seals. 3. The possible, but undocumented, taking of seals for their skins

(for skiis as climbing skins and the pup pelage for coats, etc.)* 4. The taking of seals for mink or other domesticated animal foods.* 5. The subtle effects of pollution which might include organic

sewage as well as industrial inorganic chemicals and chemical

pesticides and herbicides. 6. The taking of live seals (usually young pups) for economic exploitation in zoos, animal TTamusement parks, TT and the use I-\ f; of seals for marine, biological, and biomedical research f ! organizations.* ! *Action now illegal under Marine Mammal Protection Act of 1972. -47-

The shooting of seals has been mentioned and is thought to be less significant now than in past years. To quantify such activity would per- haps be impossible. During both field study periods, however, no cases of seal·shooting were documented by the investigator in the Mount Desert area.

One shot seal was found in the fall of 1972 at Mount Desert Rock. Some , summer residents have cited cases of seals having washed ashore in previous i '~ summers. Some had been shot or in one case had died of deep cuts in the forelimbs. It would appear logical that the reduced numbers of schooling fish such as herring, haddock, halibut and cod found in the study area might have affected the present seal population. However, the extent that increased commercial catches of these species of fish affected the seal population would depend on the willingness or capability of seals to sub- stitute other dietary items. An unwillingness to make such substitutions would at least have distributional effects; that is, seals may have moved within or away from the study area to sites where the remaining stocks would be more readily hunted. As mentioned, it is more likely that seals readily substitute dietary finfish species which are conveniently caught near chosen haul out areas. The harvesting of either seal species for skins or meat is known to occur in the Maritime Provinces, but is thus far undocumented on the coast of Maine. Pollution effects on the existing North Atlantic seal population are not well known. Harbor seals are known to be distributed in areas such as Portsmouth Harbor and the Piscataqua River, New Hampshire, where 81 to 100 percent of the clam flats are closed due to pollution. Likewise, seals seem well distributed in central and lower penobscot Bay, having 61 to 80 'j'

-48-

percent of its clam flats closed. The study area lies within a zone declared to have less than 20 percent of its clam flats closed due to pollution.*

Thus seals would appear to be exposed to less sewage and bacterial contamina-

tion than along most of the rest of the Maine coast. Gaskin (1971) reported alarmingly high residues of organochlorine pesti- cide in the tissues of harbor porpoises (Phoecoena phocoena) taken in the

Bay of Fundy. The average DDT level in the blubber of males was 306.74 ppm, exceeding the highest published values for any wild mammal. A recent report

by Gaskin, et.al. (1973) analyzing mercury, DDT, and PCB levels in harbor

seals taken in the Boothbay Harbor area were measured. DDT and PCB levels

were of the same magnitude as reported for harbor porpoises and were found

to be lowest in a lactating female. Virtually no o,pT-DDT and relatively little dieldrin were found in seal fat, in contrast to porpoises. Mercury

values in liver varied from 0.5 to 7.9 ppm in Maine seals; somewhat less than the 1.7 to 50.9 ppm found in Bay of Fundy harbor seals.

Dr. A. V. Olden has indicated (personal communication) that in compar-

ing deposition of heavy metals and pesticides in consumer tissues, there

is not much alteration in the ratios of zinc, copper, lead and cadmium found in fish species and the seals feeding on fish. In the case of mercury

and organochlorine residues, however, there is a considerable accumulation

shown in this last step of the food chain. It is likely that seals and other marine mammals will prove to be

valuable TTindicator TT organisms in monitoring the levels and rates of change

of contaminants in water masses resulting from manTs activities. Information

*Values taken from: Maine Coastal Resources Renewal, 1971. State Planning Office, Augusta, Maine -49- regarding levels of heavy metals, pesticides and other organic residues of industrial use which have potential human toxicity may be provided by tissue analysis of seals and other marine manunals.

, :.

D. Protection of Marine Mammals

On October 28, 1972, the Marine Manunal Protection Act of 1972 was signed and 60 days later it became law. Information and support for passage of this Act given to the Merchant Marine and Fisheries Conunit~has been detailed in earlier reports.

A copy of the Act is included in Appendix 2 of this report, as well as in Appendix 4 of the last progress report. It clearly addresses a funda- mental goal pursued by the initiators of this study of Acadia's seal popula- tions -- that of protection of this wildlife species in waters integral to Park land holdings. Important protective measures afforded by the Act are:

1. A complete moratorium on the taking or importing of all marine manunals or products of marine manunals for any pur-

pose unless approved by either the Department of Interior which has jurisdiction over walrus, polar bears, manatees

and sea otters, or the Department of Conunerce (National

Marine Fisheries Service) in the case of whales, porpoises,

seals and sea lions.

"\ 2. A Marine Manunal Commission and a Conunittee of Scientific II n Advisors are established and are to oversee the assessment II II ,. I!H of stocks, changes in numbers of stocks over periods of \ time, and are to interpret the carrying capacities for

species and their environments. 3. Marine manunal species are to be allowed to attain their

Tfoptimum sustainable populationTT consistent with their m ~ :I , iiiII. ::! Ii ,I: :; !I -50- I, ,I ;i ,]:I' i habitat and its carrying capacity. Such levels and defini- j' ,

tions will be interpreted by the Commission and Advisors. 4. The term TTmoratoriumTT means a complete cessation of the

taking of marine mammals; where TTtaking TT is defined as

n ••• to harass, hunt, capture, or kill or attempt to harass,

hunt, capture, or kill any marine mammaL. n 5. Enforcement of the measures of this Act will be by Federal

agents "and through the voluntary or contractual arrange-

ments between the above Departments and individual states. The Park Service may wish to establish some formal understanding with the Marine Mammal Commission offering aid in implementing measures of the Act in the vicinity of Acadia National Park. It is the opinion of this investigator that the protective measures of this Act very adequately pro- tect the seal populations in the waters about Acadia National Park provided that fishermen and residents comply with the laws set forth. Established attitudes of fishermen should be challenged by an effective publicity cam- paign based on facts assessing the role of seals to inshore fisheries. Fear of penalty will not be effective, given the impractical nature of constant patrol efforts and the covert nature of seal shooting near isolated seal ledges. The sympathies of fishermen and the public at large must be gained. Any tools available for public education regarding the intrinsic and practi- cal values of marine mammal species should be utilized.

E. Recommendations

At the present time the Acadia National Park staff is coordinating an effective interpretive program for Park visitors and residents. Inclusion of information on seals and other marine mammals occurring in waters about the Park and information, in brief, explaining the new laws protecting these mammals is recommended. The naturalist-guided boat cruises are by far r

-51-

the most rewarding means of viewing and photographing seals at close range

without danger of harassment. This author has witnessed the delight of passengers viewing seals at 50 feet or less as the boat slowly backs toward

a ledge; the seals acting almost unconcerned. Seals are quickly conditioned

to the harmless nature of these boats and allow viewing at closer range than possible with small pleasure boats. The naturalist can explain inter-

esting aspects of seal biology and habitat (and now protective measures)

during viewing when the learning experience is most meaningful. The investigator has witnessed numerous times when visitors could not

be accommodated on these cruises. If possible, the Park should encourage

more cruises to meet the heavy demand. During 1972, six daily cruises

(four routes, two running twice daily) were sponsored by the naturalist

program.

It is recommended that the Park make information on seal whelping

sites and popular haulouts known to local town conservation commissions

and other groups active in coastal shoreline zoning and wildlife and resource planning activities. It is not recommended that the Park encourage seal

viewing from private land. Such locations are very limited and could not

accommodate public access. At present, the College of the Atlantic has a TTWhale WorkshopTT composed

of students under the guidance of Ecology instructor, Dr. Steven Katona.

Plans for whale watching activities at Mount Desert Rock and other light stations and research projects by students on whale and seal ecology might be a source of cooperative and imaginative exchange with the naturalist

staff at Acadia. Participation by these students.in Park interpretive

programs could be pursued by the Park staff. .'..... ~.:.: ...... :~.~ ;!::...... y.. : ... ,~ .... ':.: .... ""\\'\I\\\\\\~ \":'.

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, •• ,.\\':\\\.)Jo.\\\\,'.~.• :.; ... '.' ., ., .•..;,. .. "\W"'~ .,,"{ .)'~ ...: .. : ... ::. ,.. ' . :.\,'W.'\·.,.".::.:~.~;;>;-\:::·::::~~.:. """\" ,\ " ,~~~\\,,\\\ ~ "':~ ---'-'-----._. __ .__ .__ . -----_._-.. _--_ .. ----- Christopher Packard Photo Top Photo Basking Mul t Harbor Seal' Ht. 'Desert Rock, Note lesions on neck cnd abdomen

1· Bottom Photo Colony of Harbor Seals i North Rock, Gr~nd Hanan Channel -53-

By initiating a census of Maine's seals, the Park Service has produced in this study a valuable baseline of data for a significant segment of the

Maine coast. During aerial surveys subsequent to this study, sections from

Kittery to Cape Small and Cape Small to pemaquid Neck have been censused during June, 1973. Continuing through the current summer, the seal popula- tions in the penobscot Bay area and the coast east of Petit Manan will be censused. From these surveys it should be possible to make estimates for numbers of seals in all the coastal embayments.

At present, the Mount Desert Island area with its well protected bays and numerous ledges within two major archipelagos, emerges as a distinct

"center" of seal concentration. Other centers will likely prove to be the lower penobscot Bay area, the lower Sheepscot Bay including islands off Boothbay Harbor and possibly the Englishman-Machias Bay area.

It is likely that the seals found near Acadia National Park are close to one-fourth of the total population of the entire coast; although this estimate is speculation, based on coverage of only about one-half the linear coastline. Nevertheless, the Mount Desert area contributes very significantly to Maine's total seal population. The unique placement of a National Park with growing concern for island habitat near seal haulouts provides oppor- tunity to further support monitoring of the two species. This investigator hopes that the National Park Service can encourage, through the Department of Interior's influence and participation in implementing the Marine Mammal Protection Act, continued monitoring of the Maine seal population. It is hoped that funds will be forthcoming, as authorized in the Act, for research and assessment of population stocks during fiscal year 1974. It is recom­ mended that efforts by this investigator with the Maine Department of Sea and Shore Fisheries complement similar efforts by the Park Service to obtain necessary grants for continuing population studies. -54-

Efficient and accurate estimates of Mainers seal populat1on should be possible through use of a probability sampling design using aerial photography and knowledge of haulout locations gained during the 1973 coastal census. Progress in developing such a sampling program will be submitted to Acadia National Park Superintendent,Keith Miller as soon as possible.

Records, documents, photographs, including aerial census transparencies and other pertinent data too lengthy or in clumsy format for this report will be submitted to naturalist William Dengler at Acadia National Park for safekeeping and reference. In the interests of brevity and reduced , ·1 xerography, several appendices referred to in this final report but appear- i, ing in one of the three semiannual progress reports will not be included here. References to the appropriate report are given.

Acknowledgements:

The author wishes to thank John Good, Robert Binnewies, William Dengler,

Robert Dow, and Keith Miller for valuable assistance and support throughout the study. Special appreciation is given to the Maine fishermen and resi- dents who provided information for this report. Sea and Shore Fisheries area wardens and Don McIntosh are thanked for aid in gaining essential data for the study. I

I j -55- LITERATURE CITED Allen, J. A. 1880. History of North American Pinnipeds, Washington Government Printing Office, 785 pp. Bigg, M. A. 1968. Clines in the Pupping Season of the Harbor Seal, Phoca vitulina, Fish. Res. Bd. Can. 26:449-455.

Bigg, M. A. 1969. The Harbour Seal in Dritish Columbia. Fish. Res. Bd. Can. Bull. 172:33 p. Boulva, J. 1972. Temporal Variations in Birth Period and Characteristics of Newborn Harbor Seals. Presented at the Symposium on the Biology of the Seal, held at Guelph, Ontario, Canada, 14-17 of August, 1972.

Boulva, J. 1971. Observations on a Colony of Whelping Harbor Seals, Phoca vitulina concolor, on Sable Island, Nova Scotia, J. Fish. Res. Bd. Canada, 28:755-759. Davies, J. L. 1957. The Geography of the Grey Seal, J. of Mammalogy, Vol. 38, No.3. Fisher, H. D. 1952. The Status of the Harbor Seal in British Columbia with Particular Reference to the Skeena River, Bull. Fish. Res. Bd. Can., No. 93:1-58. Fisher, H. D. 1954. Delayed Implantation in the Harbour Seal, Phoca vitulina L. Nature, 173:879. Gaskin, D. E. 1971. Organochiorine Pesticide Residues in Harbour porpoises from the Bay of Fundy Region. Nature, Vol. 233. Gaskin, D. E., et.al., 1972. Mercury, DDT and PCB in Harbour Seals (Phoca vitUlina) from the Bay of Fundy and Gulf of Maine, J. Fish. Res. Bd. Can., 30:471-475. Hunt, J. H. 1948. The Atlantic Harbour Seal in the Coastal Waters of Maine. Maine Department of Sea and Shore Fisheries reprint. Mansfield, A. W. 1967. Distribution of the .Harbor Seal, Phoca vitulina, Linnaeus, in Canadian Arctic Waters. Journal of Mammalogy, Vol. 48, No.2. Mansfield, A. W., ms, 1968. Seals as Vectors of Porrocaecum in the Mari­ time Provinces, Ann. Rep. 1967-68, Arctic BioI. Sta., Fish. Res. Bd. Can., p. 31-33. Montreuil, P. A. 1958. Corynosoma Magdaleni Sp. Nov. (Acanthocephala), A Parasite of the Gray Seal in Eastern Canada. Canadian Journal of Zoology 36, 205. Scott, D. M. and H. D. Fisher. 1958. Incidence of the Ascarid Porrocaecum dieipiens in the Stomachs of the Three Species of Seals along the Southern Canadian Atlantic Mainland. J. Fish. Res. Bd. Can., 15(4), pp. 495-516. -56-

Sergeant, D. E. 1973. Feeding, Growth, and Productivity of Northwest Atlantic Harp Seals (Pagophilus groenlandicus). J. Fish. Res. Bd. Can. 30:17-29. Van Bemmel, A. C. V. 1956. Planning a Census of the Harbour Seal (Phoc~ vitulina) on the Coasts of the Netherlands. Beaufortia, Vol. 5, 54:121-132.

Vaughan, R. W. 1971. Aerial Survey of Seals in the Wash, reprint, The National Environment Research Council, occasional publications: No.2, 1971. Wilson, T. M. and Sweeny, P. R. 1970. Morphological Studies of Seal Poxvirus. J. of Wildlife Diseases, Vol. 6. Wilson, T. M. et. a1. 1972. Pox in Young, Captive Harbor Seals. J.A.V.M.A., Vol. 161,~o.-g, pp. 611-617. Worthington, E. B. 1964. Gray Seals and Fisheries, Nature 103:116-118.

,l ,! I I i Ii

(' ii'i ~:i .j :i -57-

REFERENCES

Andrews, J. C. and P. R. Mott. 1967. Gray Seals at Nantucket, Mass. J. of Mammalogy, Vol. 48(4), p. 657-658. Beck, B.; T. G. Smith; A. W. Mansfield. 1970. Occurrence of the Harbour Seal, Phoca vitulina, Linnaeus, in the Thlewiaza River, N.W.T., The Canadian Field Naturalist, Vol. 84.

Bishop, R. H., ms., 1967. Reproduction, Age Deter~ination and Behavior of the Harbour Seal, Phoca vitulina L., in the Gulf of Alaska. M. Sc. thesis, Univ. of Alaska, College, Alaska, 121 p. Burns, J. J. and S. J. Harbo (no date available). An Aerial Census of Ringed Seals, Northern Coast of Alaska. Alaska Dept. of Fish and Game, Fairbanks, Alaska, U.S.A. University of Alaska, College, Alaska, U. S.A. Cameron, A. W. 1966. Breeding Behavior in a Colony of Western Atlantic Gray Seals. Canadian J. of Zoology, Vol. 45. Caughley, G. and J. Goddard. 1972. Improving the Estimates from Inaccu­ rate Censuses, J. of Wildlife Management, 36:1.

Drury, W. H. 1965. Lets Save Nantuckets Horseheads. Yankee Magazine, Vol. 29(5), 62:101-105. Geraci, J. R. and M. C. Keys (no date given on report). Veterinary Medicine in the Conservation and Management of Marine Mammal Resources, J.A.V.M.A., Vol. 157(2). Johnston, D. G. and S. Ridgway, 1969. Parasitism in Some Marine Mammals, J.A.V.M.A., Vol. 155(7). Joseph, D. C. 1970. Dormant Embryos of Mammals. BioScience, Vol. 27(7). Mansfield, A. W. Seals and Walrus. Science, History, and Hudson Bay. (no further reference available). Mansfield, A. W. and H. D. Fisher, 1970. Age Determination of the Harbour Seal, Phoca vitulina L., Nature, Vol. 186(4718), p. 92-93. Mansfield, A. W. 1966. The Gray Seal in Eastern Canadian Waters. Canadian Audubon, Vol. 28(4), p. 161-166. McGinnis, S. M. 1968. Biotelemetry in Pinnipeds, The Behavior and Physiology of Pinnipeds, ed. by R. H. Harrison et.al., Appleton­ Century-Crofts, New York, N. Y. McLaren, I. A. 1958. The Biology of the Ringed Seal (Phoca hispida Schreber) in the Eastern Canadian Arctic, Bull. No. 118, Fish. Res. Bd. Can. -58-

McLaren, I. A. 1960. On the Origin of the Caspian and Baikal Seals and the Paleoclimatological Implication. American J. of Science, Vol. 258, p. 47-65.

McLaren, I. A. 1960. Are the Pinnipedia Biphylectic? Systematic Zoology, Vol. 9(1).

McLaren, I. A. Methods of Determining the Numbers and Availability of Ringed Seals in the Eastern Canadian Arctic. J. of Arctic Research, (reprint, date and volume unknown). McLaren, I. A. 1966. Taxonomy of Harbour Seals of the Western North Pacific and Evolution of Certain other Hair Seals. J. of Mammalogy, Vol. 47(3). ! . McLaren, I. A. 1966. Analysis of an Aerial Census of Ringed Seals. J. Fish. Res. Bd. Can., 23(5). McLaren, I. A. 1967. Seals and Group Selection. Ecology, Vol. 48(1).

Montreuil, Paul. May 2,1972. Personal communication (letter). Pike, G. C. and I. B. MacAskie. 1969. Marine Mammals of British Columbia, Fish. Res. Bd. Can., Bull. 171. Rae, B. B. 1960. Seals and Scottish Fisheries, Edinburgh, Her Majesty's Stationery Office, Marine Research No.2. Rae, B. B. 1968. The Food of Seals in Scottish Waters, Edinburgh, Her Majesty's Stationery Office, Marine Research No.2. Rae, B. B. 1973. Further Observations of the Food of Seals, J. of Zoology, London, 169, p. 287-297. Rough, V. 1968. Gray Seals in New England. Massachusetts Audubon, Vol. LII(3). Scheffer, V. B. 1958. Seals, Sea Lions and Walruses. Stanford Univ. Press. Slater, L. 1963. BIOTELEMETRY: The Use of Telemetry in Animal Behavior and Physiology in Relation to Ecological Problems. Proceedings of Interdisciplinary Conference, N. Y., MacMillan Co., N. Y. Smith, T. G. 1970. Computer Programs for Analysis of Ringed Seal popula­ tion Data. Fish. Res. Bd. of Can., Technical Report No. 224.

Stutz, S. S. 1966. Foetal and Postpartum Whitecoat Pelage in Phoca vitulina, J. Fish. Res. Bd. Can., 23(4). Vaughan, R. W. 1971. Aerial Survey of Seals in the Wash. Seals Research Unit NERC, occasional publications: No.2. -59-

Vaughan, R. W. 1971. Aerial Photography in Seals Research. The Nature Conservancy, Edinburg. Vedros, N. A. et.al. 1971. Leptospirosis Epizootic among California Sea Lions. SCIence, Vol. 172, pp. 1250-1251. Wilke, F. and K. W. Kenyon. 1952. Notes on the Food of Fur Seals, Sea Lions and Harbor Porpoises. J. Wildlife Management 16(3), p. 396. Wilke, F. 1953. Seals of Northern Hokkaido; J. of Mammalogy, Vol. 35(2). Young, P. C. 1971. The Relationship Between the Presence of Larval Anisakine Nematodes in Cod and Marine Mammals in British Home Waters. J. of Applied Ecology (reprint - volume unknown).

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