NOAA Technical Memorandum NMFS-NE-117

This series represents a secondary level of scientific publishing. All issues employ thorough internal scientific

review; some issues employ external scientific review. Reviews are -- by design -- transparent collegial

reviews, not anonymous peer reviews. All issues may be cited in formal scientific communications.

Review of Distribution of the

Long-finned Pilot Whale (Globicephala melas)

in the North Atlantic and Mediterranean

Alan G. Abend1 and Tim D. Smith2

1Univ. of Massachusetts, Amherst, MA (Present address: Zoo New England, Boston, MA 02121)

2National Marine Fisheries Serv., Woods Hole, MA 02543

U.S. DEPARTMENT OF COMMERCE

William Daley, Secretary

National Oceanic and Atmospheric Administration

D. James Baker, Administrator

National Marine Fisheries Service

Penelope D. Dalton, Assistant Administrator for Fisheries

Northeast Region

Northeast Fisheries Science Center

Woods Hole, Massachusetts

April 1999
Note on Species Names

The NMFS Northeast Region's policy on the use of species names in all technical communications is generally to follow the

American Fisheries Society's lists of scientific and common names for fishes (i.e., Robins et al. 1991a), mollusks (i.e., Turgeon

et al. 1998b), a decapod crustaceans (i.e., Williams et al. 1989c), and to follow the Society for Marine Mammalogy's guidance

on scientific and common names for marine mammals (i.e., Rice 1998d). Exceptions to this policy occur when there are

subsequent compelling revisions in the classifications of species, resulting in changes in the names of species (e.g., Cooper

and Chapleau 1998e).

___________________

a

Robins, C.R. (chair); Bailey, R.M.; Bond, C.E.; Brooker, J.R.; Lachner, E.A.; Lea, R.N.; Scott, W.B. 1991. Common and scientific names

of fishes from the United States and Canada. 5th ed. Amer. Fish. Soc. Spec. Publ.20; 183 p.

b

Turgeon, D.D. (chair); Quinn, J.F.; Bogan, A.E.; Coan, E.V.; Hochberg, F.G.; Lyons, W.G.; Mikkelsen, P.M.; Neves, R.J.; Roper, C.F.E.;

Rosenberg, G.; Roth, B.; Scheltema, A.; Thompson, F.G.; Vecchione, M.; Williams, J.D. 1998. Common and scientific names of aquatic

invertebrates from the United States and Canada: mollusks. 2nd ed. Amer. Fish. Soc. Spec. Publ. 26; 526 p.

c

Williams, A.B. (chair); Abele, L.G.; Felder, D.L.; Hobbs, H.H., Jr.; Manning, R.B.; McLaughlin, P.A.; Pérez Farfante, I. 1989. Common

and scientific names of aquatic invertebrates from the United States and Canada: decapod crustraceans. Amer. Fish. Soc. Spec. Publ. 17;

77 p.

d

Rice, D.W. 1998. Marine mammals of the world: systematics and distribution. Soc. Mar. Mammal Spec. Publ. 4; 231 p.

e

Cooper, J.A.; Chapleau, F. 1998. Monophyly and interrelationships of the family Pleuronectidae (Pleuronectiformes), with a revised

classification. Fish. Bull. (U.S.) 96:686-726.
Page iii

Contents

Abstract ................................................................................................................................................................................................... v

Introduction ............................................................................................................................................................................................. 1

United States ........................................................................................................................................................................................... 1

Canada ..................................................................................................................................................................................................... 6

Greenland ................................................................................................................................................................................................. 9

Iceland ...................................................................................................................................................................................................... 9

Norway .................................................................................................................................................................................................. 11

Faeroe Islands ........................................................................................................................................................................................ 13

British Isles ............................................................................................................................................................................................ 15

The Netherlands .................................................................................................................................................................................... 16

France .................................................................................................................................................................................................... 16

Spain and Portugal ................................................................................................................................................................................. 16

Italy ....................................................................................................................................................................................................... 18

Discussion ............................................................................................................................................................................................. 18

Endnotes ................................................................................................................................................................................................ 19

References Cited .................................................................................................................................................................................... 19

Tables

Table 1. Total number of pilot whales taken in targeted fisheries by month and by region in Greenland from 1828 to 1991 .......... 10

Table 2. Pilot whale sightings in Icelandic and adjacent waters between 1981 and 1989 .................................................................. 10

Table 3. Pilot whale sightings off Norway from 1987 to 1990 ......................................................................................................... 13

Figures

Figure 1. Distribution of long-finned pilot whales in the North Atlantic and Mediterranean Sea based on

sighting data from 1952 to 1992 .......................................................................................................................................... vi

Figure 2. Coverage of CeTAP sighting surveys of the continental shelf and shelf edge from 1978 to 1981 ....................................... 2

Figure 3. Late winter - early spring distribution of pilot whales off the eastern United States based on sighting

data from 1978 to 1992 ......................................................................................................................................................... 4

Figure 4. Spring distribution of pilot whales off the eastern United States based on sighting data from 1978 to 1992 ...................... 4

Figure 5. Summer distribution of pilot whales off the eastern United States based on sighting data from 1978 to 1992 ................... 5

Figure 6. Autumn distribution of pilot whales off the eastern United States based on sighting data from 1978 to 1992 ................... 5

Figure 7. Distribution of long-finned pilot whales off Newfoundland based on sighting data from 1979 to 1982 ............................. 7

Figure 8. Distribution of long-finned pilot whales off Greenland based on sighting data from 1981 to 1991 ..................................... 7

Figure 9. Distribution of long-finned pilot whales off Iceland based on sighting data from 1981 to 1989 ........................................ 12

Figure 10. Distribution of long-finned pilot whales off Norway based on incidental sightings from 1971 to 1989

and sighting data from 1989 ................................................................................................................................................ 12

Figure 11. Distribution of long-finned pilot whales off Faeroe Islands based on sighting data from 1986, 1987, and 1989 ............... 14

Figure 12. Distribution of long-finned pilot whales off the British Isles based on sighting data for 1987 and 1989 .......................... 14

Figure 13. Distribution of long-finned pilot whales off Spain based on sighting data from 1981 to 1989 .......................................... 17

Figure 14. Distribution of long-finned pilot whales off Italy based on sighting data from 1981 to 1988 ........................................... 17
Page iv

Acronyms

CeTAP = (University of Rhode Island) Cetacean and Turtle Assessment Program

GFRI = Greenland Fisheries Research Institute

MBO = Manomet Bird Observatory

NASS = North Atlantic Sighting Survey

NEFSC = (NMFS) Northeast Fisheries Science Center

NMFS = NOAA National Marine Fisheries Service

SCANS = Small Cetacean Abundance in the North Sea Survey
Page v

ABSTRACT

 The spatial and seasonal distribution of the long-finned pilot whale (Globicephala melas) in the North Atlantic is

summarized based on published and some unpublished data. The data are presented separately for 12 countries, and within

each country by type of data: 1) evidential (i.e., sightings, strandings, harvests, bycatches); and 2) inferential (i.e., oceanographic processes, prey occurrences). The locations of sightings are presented in distribution maps for each country. The

locations of the several sighting surveys and other field activities are summarized, but the distribution of sighting effort is not

displayed. For each country, the general distribution pattern is summarized, seasonal differences are shown, and possible

seasonal movement patterns are suggested. The Atlantic-wide distribution is then summarized, based on these individual

country summaries.
Page vi

Figure 1. Distribution of long-finned pilot whales in the North Atlantic and Mediterranean Sea based on sighting data from 1952 to 1992.
Page 1

INTRODUCTION

 The long-finned pilot whale (Globicephala melas Traill

1809) is also referred to as the blackfish, pothead, northern

pilot whale, Atlantic pilot whale, caa'ing whale, and

grindehval. Most literature references to the long-finned

pilot whale use the short common name of "pilot whale" in

conjunction with the scientific name to distinguish it from

the short-finned pilot whale (Globicephala macrorhyncus

Gray 1846). The long-finned pilot whale is a robust whale

with a large bulbous head and a flattened forehead. Its

coloration is black with an anchor-shaped patch of grayish

white on the throat extending into a gray stripe along the

underside surrounding the navel and genital areas. Pectoral flippers, long and sharply pointed, reach one-fifth the

length of the body (Caldwell and Caldwell 1983). Males can

be up to 6 m long and weigh 2,400 kg; females are roughly

25% smaller (Sergeant 1962; Martin et al. 1987; Kasuya et

al. 1988).

 The distribution of pilot whales in the North Atlantic

(Figure 1) extends from Cape Hatteras to Greenland in the

west, to Iceland and possibly the Barents Sea in the north,

and to northwestern Africa and the Mediterranean in the

east (Sergeant 1962; Leatherwood et al. 1976; Evans 1980;

Nores and Pirez 1988). For hundreds of years, strandings

of pilot whales have occurred regularly along the coasts of

the North Atlantic (Brown 1975; Geraci and St. Aubin 1977;

Martin et al. 1987; McFee 1990).

 The principal sources of data on distribution patterns

fall into two categories: 1) evidential (i.e., dedicated and

incidental sightings, strandings, directed harvests,

bycatches in commercial fisheries); and 2) inferential (i.e.,

oceanographic processes, prey occurrences). Dedicated

sighting surveys have been conducted by several nations/

territories, including the United States, Canada, Iceland,

Faeroe Islands, Denmark, Norway, France, Spain, Great Britain, and Italy. Prior to 1987, sighting surveys for cetaceans

in the Northeast Atlantic were limited in both area and duration (Buckland et al. 1993). During 1987 and 1989, largescale cetacean sighting surveys were conducted in the

Northeast Atlantic. In the Northwest Atlantic, a linetransect survey was conducted in 1980 (Hay 1982), aerial

surveys in 1978-82, and shipboard surveys in 1980-88 (Cetacean and Turtle Assessment Program 1982; Payne and

Heinemann 1993). Data have regularly been collected on

natural strandings in many countries. Catch data have been

collected from drive fisheries in the Faeroe Islands and

United States, with records from the former dating back

several centuries. Biological samples have been collected

from Canadian and Faeroese fisheries, and from natural

strandings and bycatches in commercial fisheries in the

United States.

 Most of these data have been analyzed to determine

local or regional distribution patterns. Lacking, however, is

an integration of the large-scale distribution patterns of

pilot whales. In this paper, we describe the data available

from each of 12 countries in the North Atlantic, and through

the use of summary maps attempt to depict overall distribution and movement patterns.

UNITED STATES

 Recorded data on pilot whales began a few days after

the Pilgrims arrived from England, as pilot whales stranded

near the Pilgrims' first landing site. Recordings from the

drive fishery of Cape Cod, Massachusetts, lasted until the

1950s. Since then, dedicated shipboard and aerial surveys

have been conducted to map the distribution of marine

mammals in the eastern U.S. coastal and off-shore waters.

Additional research has involved life history data collection from strandings and from incidental takes during commercial fishing operations. These data sets provide the basis for assessing pilot whale distribution patterns along the

Mid- and North Atlantic regions of the United States.

Dedicated Sighting Surveys

 Beginning in 1978, the United States initiated dedicated

cetacean surveys of the northwestern shelf of the North

Atlantic Ocean. This effort, the Cetacean and Turtle Assessment Program (CeTAP), which ran from November 1978

to January 1982, was sponsored by the U.S. Department of

the Interior's Bureau of Land Management for making decisions on oil and gas exploration and development. The goal

of the surveys was to characterize marine mammals and

turtles in the Mid- and North Atlantic regions of the U.S.

outer continental shelf. The surveys were initially conducted

by air and sea, but shipboard surveys were suspended in

1980, with the aerial surveys continuing for the remainder

of the program.

 A second program of surveys was conducted from January 1980 to December 1988 through a contract with the

Manomet Bird Observatory (MBO) of Manomet, Massachusetts, under the auspices of the National Marine Fisheries Service's (NMFS's) Northeast Fisheries Science Center

(NEFSC) in Woods Hole, Massachusetts. Shipboard surveys were performed aboard NMFS fishery research vessels as the ships traveled along designated cruise tracks.

The objective of the surveys was to provide long-term monitoring of the distribution and abundance of cetacean and

seabird populations, which could be directly compared to

the distribution and abundance of fish stocks in the same

waters (Payne et al. 1984; Smith et al. 1988).

 The two survey programs covered all of the continental

shelf and shelf-edge waters between Nova Scotia, Canada,

and Cape Hatteras, North Carolina. The CeTAP surveys

extended from the coastline seaward to include outer continental shelf waters. The seaward projection included waters 9.3 km (5 nautical miles) beyond the 1,830-m depth contour, which included the associated deepwater canyons
Page 2

Figure 2. Coverage of CeTAP sighting surveys of the continental shelf and shelf edge from 1978 to 1981 (Source: Cetacean and Turtle

Assessment Program 1982).
Page 3

(Cetacean and Turtle Assessment Program 1982; Figure 2).

The MBO surveys were in waters less than 910 m, with

most surveys in waters less than 183 m (Payne and

Heinemann 1993). This coverage by the two survey programs encompassed a large portion of the long-finned pilot

whale's range in the western North Atlantic and a part of

the northern distribution of the short-finned pilot whale.

 The short-finned pilot whale ranges from Cape Hatteras

south to the Caribbean (Caldwell and Caldwell 1983; Leatherwood et al. 1976). The two species appear to overlap

seasonally between approximately 35°N latitude (Cape

Hatteras) and 38°-39°N latitude (Payne et al. 1991). Identification to the species level is difficult at sea. Accordingly,

the dedicated survey sightings refer only to the genus level

(Globicephala).

 A total of 619 sightings of pilot whales were recorded

in the two survey series. Depths associated with sightings

ranged from 15 to 5,121 m. This range reflects both survey

effort and the movements of the whales between inshore

and offshore waters. Pilot whales were often sighted with

other cetacean species, especially bottlenose dolphins

(Tursiops truncatus). Cow-and-calf pairs and juveniles were

primarily seen along the shelf edge from Cape Hatteras to

the Gulf of Maine, mainly in the spring and summer, with

fewer observed in the autumn and winter (Cetacean and

Turtle Assessment Program 1982). The sightings concur

with Sergeant's (1962) suggestion that pilot whale calves

are born from May to November, peaking in August.

 There have been only limited surveys in waters greater

than 2,000 m in depth. This limitation has created a gap in

our understanding of the utilization of these waters by cetaceans. During the summers of 1990 and 1991, the NEFSC

conducted marine mammal surveys in the shelf edge and

offshelf waters to the Gulf Stream between Cape Hatteras

and Georges Bank (Waring et al. 1992). The surveys ran

from August 5 to August 18, 1990, and from June 8 to July

10, 1991. Nearly two-thirds of the 1990 survey effort was

along the shelf edge off Georges Bank and within the Gulf

Strewn. The remaining one-third concentrated around a

warm-core ring near Georges Bank and in a 9.3-km buffer

zone of the Gulf Stream northern wall. The greater portion

of the 1991 survey effort primarily concentrated in shelfedge waters between the 200- and 2000-m contours, with a

smaller portion conducted in off-shelf waters (Waring et al.

1992).

 In 1990, a total of 68 pilot whales were sighted. The

sightings occurred primarily along the northern wall of the

Gulf Stream, along the shelf break, and near Oceanographer

Canyon, with a couple of sightings recorded in the midportion of the Gulf Stream near Cape Hatteras. During the

1991 survey, a total of 724 pilot whales were sighted. Most

of the 1991 sightings occurred along the shelf break, in

Oceanographer, Hydrographer, and Corsair Canyons, along

the southern edge of Georges Bank, and along the northern

wall of the Gulf Stream (Waring et al. 1992). In both years,

the whales were principally found in groups of 2-5 individuals.

 The combined CeTAP, MBO, and NEFSC sighting data

indicate that pilot whales occur primarily along the continental shelf edge between the 100- and 2000-m contours

from Cape Hatteras northerly to Georges Bank. A few

sightings also occurred inshore of the 100-m contour, as

well as seaward of the 2,000-m contour. These latter sightings

are consistent with observations from deep ocean weather

stations indicating that pilot whale distribution includes

some or all of the deep ocean (Brown 1961).

 During late winter to early spring, pilot whale concentrations along the shelf edge at approximately 35°N join

additional pilot whales (presumed from deeper off-shore

waters) and move northward along the shelf edge between

the 100- and 1000-m contours onto the continental shelf

(Payne et al. 1991; Figure 3). Between early March and

May, pilot whales substantially increase in number along

the shelf edge between 38°N and 42°N latitude (Figure 4).

 During summer, pilot whales are broadly distributed over

the shelf (Cetacean and Turtle Assessment Program 1982;

Payne et al. 1984, 1991; Figure 5). The greater percentage of

sighted pilot whales occur along the shelf edges of the

northeastern portion of Georges Bank and onto the Scotian

Shelf. During May through October, pilot whales predominantly occur on central Georges Bank, primarily along its

northern edge (Hain et al. 1981; Cetacean and Turtle Assessment Program 1982; Payne et al. 1984, 1991). This summer movement continues northward into the Gulf of Maine

and into Canadian waters (Sergeant and Fisher 1957; Sergeant 1962; Mercer 1975).

 During autumn, the largest concentrations of pilot

whales occur along the southwestern edge of Georges Bank

(Figure 6). By late autumn to early winter, many pilot whales

have already moved offshore and southward (Cetacean and

Turtle Assessment Program 1982; Payne et al. 1991). The

survey sightings are thus consistent with the Katona et al.

(1978) and Sergeant (1962) hypothesis that some pilot whales

move southward into deep offshore waters during the winter. Although, a component of the population appears to

be seasonably migratory, the sighting data indicate that

pilot whales are common year-round residents of particular

continental shelf areas, notably the southern margin of

Georges Bank (Cetacean and Turtle Assessment Program

1982).

Strandings

 Pilot whale strandings on Cape Cod, Massachusetts,

have been known to occur since 1620, and were common in

Cape Cod Bay during the l7th century (Weeden 1894; Tower

1907; Hart 1924; Thompson 1928). Since 1819, when newspapers began recording strandings, there have been 92

stranding events on Cape Cod. In addition to natural

strandings, drive fisheries caused 70 strandings between

1828 and 1950.

 Strandings follow a seasonal cyclical pattern. Strandings

between 1832 and 1899 occurred primarily during Septem-
Page 4

ber through November, while strandings between 1900 and

1959 occurred during July and August. From 1960 to present

strandings have returned to later in the year, occurring during September through December (McFee 1990).

 The stranding pattern may be due to cyclical ocean

temperatures that may regulate the movement of prey species and, therefore, the movement of pilot whales. During

summer and autumn 1987, cooler ocean temperature and

lower salinity may have influenced the migration of Atlantic mackerel (Scomber scombrus) and longfin inshore squid

(Loligo pealeii) -- both prey of long-finned pilot whales --

during the following winter. Mackerel routinely migrate to

Cape Hatteras during February and March (Anderson and

Paciorkowski 1980). However, during winter 1988, commercial fisheries data suggest that mackerel did not migrate,

remaining instead near Hudson Canyon (Fairfield et al.

1989).

 Residents of Cape Cod have reported fish congregating

within Cape Cod Bay prior to pilot whale strandings. Although the identity of these fish was unknown, it is posFigure 3. Late winter - early spring distribution of pilot whales off the eastern United Sates based on sighting data from 1978 to 1992

(Source: Cetacean and Turtle Assessment Program 1982; Payne et al. 1991; Waring et al. 1992).

Figure 4. Spring distribution of pilot whales off the eastern United States based on sighting data from 1978 to 1992 (Source: Cetacean and

Turtle Assessment Program 1982; Payne et al. 1991; Waring et al. 1992).
Page 5

sible that they were Atlantic mackerel and/or longfin in shore

squid. When squid do migrate inshore, they may have a

short occupancy period within Cape Cod Bay (Arnold et al.

1974; Macy 1982). The pilot whales entering Cape Cod Bay

could have been in pursuit of mackerel and/ or squid. Unfortunately, stranded pilot whales reveal very little about

their recent foraging as their stomachs are almost always

empty.

 Cape Cod Bay has varying water depths that could trap

the whales when the tide goes out. In fact, almost 70% of

the strandings have occurred in the varying depths of the

inner bay (McFee 1990).

Incidental Takes in Commercial Fisheries

 Beginning in 1977, fisheries compliance inspectors (i.e.,

observers) were required aboard all foreign fishing vessels

operating within the U.S. Exclusive Economic Zone (i.e.,

200-mile zone), in compliance with the Fishery ConservaFigure 5. Summer distribution of pilot whales off the eastern United States based on sighting data from 1978 to 1992 (Source: Cetacean

and Turtle Assessment Program 1982; Payne et al. 1991; Waring et al. 1992).

Figure 6. Autumn distribution of pilot whales off the eastern United States based on sighting data from 1978 to 1992 (Source: Cetacean

and Turtle Assessment Program 1982; Payne et al. 1991; Waring et al. 1992).
Page 6

tion and Management Act of 1976. The observer coverage

resulted in increased reporting of marine mammal sightings

and incidental takes (i.e., bycatch) by the fishing fleets.

 Between 1984 and 1988, 93% of all pilot whale incidental kills occurred in the foreign fishery for Atlantic mackerel

within the Mid-Atlantic Bight. This fishery followed mackerel concentrations which occur along the shelf edge during winter and spring (Anderson and Paciorkowski 1980).

Pilot whales concentrate along the shelf edge from December through May, and are mainly taken in the spring along

the shelf edge. Sixty-two percent of the pilot whale bycatch

during 1977-88 occurred between March and July, with the

remainder between December and February. The bycatch

occurred primarily near the 100-m contour (Waring et al.

1990; Fairfield et al. 1989).

 Since 1987, a drift gillnet fishery for swordfish has developed in which incidental catches of both short-finned

and long-finned pilot whales are taken (unpubl. data1

). The

fishery has operated from Cape Hatteras in winter to Georges

Bank in summer, and is usually associated with the continental shelfbreak, especially in association with Gulf Strewn

features. Bycatches of pilot whales in the Cape Hatteras

region in winter are most likely all short-finned pilot whales,

and in the Georges Bank region in summer are most likely all

long-finned pilot whales, based on those animals identified

to species.

Oceanography

 Kenney and Winn (1986) indicate that after the western inshore Gulf of Maine, the continental shelf edge is the

second-most-used cetacean habitat off the northeastern

United States. In turn, pilot whales are the second-mostobserved cetacean -- behind the bottlenose dolphin -- in

that habitat (Hain et al. 1985). Kenney and Winn (1987)

suggest that this shelf-edge habitat use is due to complex

oceanographic processes elevating productivity and providing an important habitat for the squid and fish prey of

cetaceans (Coelho 1985; Maurer and Bowman 1985; Rowell

et al. 1985).

 Pilot whales may also use submarine canyons, in addition to the shelf edge, as feeding areas. Church et al. (1984)

suggest that circulation patterns induced by undersea canyon topography promote oceanographic mixing. Squid may

be more numerous around canyons and canyon heads due

to high availability of food caused by nutrient upwelling.

According to Kenney and Winn (1987), Atlantic spotted

dolphins (Stenella frontalis), which feed on squid (Perrin

et al. 1973), showed higher densities within the submarine

canyon areas.

 In addition, pilot whale sightings were associated with

warm-core rings (Waring et al. 1992).

U.S. Summary

 Pilot whales are distributed along the eastern United

States, migrating seasonally along the edge of the continental shelf. Some animals appear to be year-round residents on the southern edge of Georges Bank, suggesting

continuous abundance and availability of prey items in that

region. Squid appear to be an important prey item but pilot

whales are known to eat mackerel, and to inhabit areas with

no squid. The nutrient-rich continental shelf edge and slope

support an extensive food web important to pilot whales

and other toothed cetaceans. Various oceanographic processes on the shelf edge and slope, including local upwelling and direct interaction with the Gulf Stream through

warm-core rings, generate high levels of productivity.

CANADA

 The majority of information on pilot whale biology in

Canada has been derived from commercial whaling operations. Sergeant (1962) based his research on samples from

harvested whales, and the apparently high rate of exploitation has spurred subsequent studies of population densities. Additional data come from sightings aboard fishing

and coast guard vessels, whale watching operations, fishery research operations, and from some land-based operations. Strandings have also contributed to the life history

of pilot whales in this region. Since strandings are few, they

are considered together with the fishery.

Ocean and Land-Based Sightings

 Historically, pilot whales have been common seasonal

migrants to Canadian waters (Sergeant and Fisher 1957;

Sergeant 1962; Mercer 1967, 1975). They range from the

offshore waters of Labrador, to the coastal and inshore

waters of Newfoundland, to the waters around Nova Scotia

and Prince Edward Island. Lynch (1987) noted that most

sightings of pilot whales occurred during autumn, and were

concentrated along the eastern coast of Newfoundland and

off the Avalon Peninsula (Figure 7). Pilot whales have been

sighted in the Labrador Sea from late May to late July, as

well as in more southerly locations in the North Atlantic

Drift (Sergeant 1968). Sightings have been irregular on the

southwestern and western coasts of Newfoundland (Sergeant and Fisher 1957).

 Sergeant (1962) suggested that the main wintering area

of the Newfoundland stock was in the waters of the North

Atlantic Current, east of the Grand Bank. Winter sightings

of pilot whales by Sergeant and Fisher (1957) and by Schevill
Page 7

(pers. comm. to Mercer 1967) are consistent with this suggestion.

 Pilot whale sightings off Canada have fluctuated, with a

recent peak in 1981; however, sightings in the late 1980s

and early 1990s have been infrequent (Lynch 1987; Jon Lien,

pers. comm.2). During 1979-82, Lynch (1987) reported few

sightings of pilot whales off Labrador or the Grand Bank.

Sightings and strandings (less than five animals per stranding) of pilot whales have most recently occurred off Nova

Scotia and Prince Edward Island in the early 1990s (Jon

Lien, pers. comm.3

).

Oceanography and Prey Distribution

 Seasonal migrations of pilot whales in Canadian waters

may relate to distribution of their prey. The concentration

of pilot whales along the eastern coast of the Canadian

Figure 7. Distribution of long-finned pilot whales off Newfoundland based on sighting data from 1979 to 1982 (Source: Cetacean and

Turtle Assessment Program 1982; Payne et al. 1991; Waring et al. 1992).

Figure 8. Distribution of long-finned pilot whales off Greenland based on sighting data from 1981 to 1991 (Source: Cetacean and Turtle

Assessment Program 1982; Payne et al. 1991; Waring et al. 1992).
Page 8

Atlantic Maritimes coincides with large aggregations of

northern shortfin squid (Illex illecebrosus) -- their prime

prey in these waters (Sergeant 1962). The arrival of northern shortfin squid in Canadian waters depends on

hydrographical conditions, feeding, and reproductive cycles

(Sergeant and Fisher 1957; Roper et al. 1984). Movement of

these squid onto the Scotian Shelf takes place during the

spring and summer when water temperatures are increasing. In addition, there is evidence that bottom water temperature influences squid distribution (Scott 1978). The preferred temperature range of the shortfin squid is 5°-15°C

(Frost and Thompson 1933).

 Adult northern shortfin squid occur over the continental shelf from the southeastern United States to Newfoundland and Labrador during summer and autumn. Black et al.

(1987) proposed that in late autumn adults move southward

to the grounds off Cape Hatteras to spawn.

 The inshore movement of pilot whales into bays normally begins in mid-July, and closely follows the movement

of northern shortfin squid. The whales arrive shortly after

the squid, and leave slightly before the last squid departs in

the autumn; no whale sightings have been recorded in Winter (Mercer 1975; Lynch 1987). Lynch (1987) reports pilot

whales were sighted off the northeastern and/or northwestern coasts of Newfoundland before the shortfin squid arrived. During the late summer and autumn, vast numbers of

shortfin squid inhabit the coastal waters and inshore bays

(Squire 1957; Black et al. 1987), but most recently, the squid

have been absent from Newfoundland waters. The relationship between inshore movement of pilot whales and shortfin

squid appears to occur only in the Newfoundland area (Mercer 1975). Shortfin squid have shown almost or complete

absence from inshore waters at times, with no detectable

regularity (Templeman and Fleming 1953). The failure of

squid to enter the bays has resulted in low or no catches of

pilot whales (Sergeant and Fisher 1957).

 In offshore waters, pilot whales also follow the movements of northern shortfin squid. During the spring and

early summer, squid concentrate at the edge of the continental shelf in waters warmer than 5°C. This shelf-edge

concentration of squid occurs when overlying cold arctic

water impedes the vertical dispersal and horizontal movement of the squid to shallow areas on the shelf (Mercer

1975; Black et al. 1987). Within the same time period, pilot

whales are widespread off Nova Scotia, Labrador Sea, Gulf

of St. Lawrence, and along the edge of the Grand Bank.

Throughout the summer, pilot whales feed on shortfin squid

that concentrate on the southwestern edge of the Grand

Bank (Brown 1961; Sergeant and Fisher 1957; Black et al.

1987). Pilot whales are reported to pass through the bank

during the autumn, moving to offshore waters that are presumably their wintering grounds (Sergeant and Fisher 1957;

Sergeant 1962).

 Although northern shortfin squid are the primary prey

of pilot whales in Newfoundland, fish are also occasionally

taken as prey (Mercer 1975). From 1951 to 1956, Sergeant's

(1962) examination of pilot whale stomach contents revealed

only shortfin squid. In late 1957, the rapid disappearance of

shortfin squid in inshore waters was followed by the disappearance of most pilot whales. The remaining whales, however, preyed on Atlantic cod (Gadus morhua). In 1959,

when the squid had begun to return, many whales were still

feeding on cod.

Pilot Whale Drive Fishery and Strandings

 For several centuries prior to 1900, small harvests of pilot whales were taken in the bays of Newfoundland by whalers and fishermen who drove herds ashore or harvested

pilot whales that had stranded (Templeman 1966). Modern

commercial whaling began in 1947 and lasted until December 1972 when Canada implemented a ban on commercial

whaling. Trinity Bay was the major whale fishery station in

Canada. Whales were also taken in Bonavista Bay and in

Notre Dame Bay (Sergeant 1962). During the 26 yr of the

fishery, approximately 54,000 pilot whales were taken, primarily by driving whales into the shallow bays and killing

them by lancing. In 1956, pilot whale kills peaked at 9,799.

Subsequently, the numbers killed declined, with only a few

hundred taken annually during the late 1960s to early 1970s

(Mitchell 1974; Mercer 1975).

 Sergeant (1962) concluded that the commercial fishery

had no effect on the population, and proposed that the

short-term decline of pilot whale landings subsequent to

record catches in 1956 was due to hydrographic conditions.

Sergeant's conclusion was based on density-dependent

factors between shortfin squid and pilot whales. However,

Mercer (1975) concluded that the 4-yr period upon which

Sergeant based his conclusion was not long enough to

influence the generation time of pilot whales which is 6 yr.

On the other hand, Mercer (1975) and Mitchell (1974), using

the cumulative catch of pilot whales between 1951 to 1961,

concluded that the initial pre-exploited population was between 50,000 and 60,000 animals, and that overexploitation

was the main cause of the decline of the pilot whale population.

 Recent estimates of pilot whale abundance in the Newfoundland/Labrador and Grand Bank region are highly variable, An aerial survey in 1980 of the coastal and off-shore

(approximately 100 nautical miles) waters of eastern Newfoundland and southeastern Labrador estimated pilot whale

abundance to be between 6,731 and 19,603 animals (Hay

1982). However, an analysis of land and sea-based sighting

records from 1976 to 1983 concluded that the abundance of

pilot whales off Newfoundland was unknown (Lynch 1987).

 Strandings have occurred primarily at Sable Island, Nova

Scotia (Sergeant et al. 1970; Geraci and St. Aubin 1977).

Between 1957 and 1976, approximately 12 strandings occurred on Sable Island ranging from 1 to 130 animals. These

whales were probably foraging for prey, and were trapped

as the tides receded.
Page 9

Canadian Summary

 Pilot whales have historically been seasonally abundant

in Canadian waters, with highest abundances in summer

and autumn. Local movements of pilot whales have reflected movements of prey species such as northern shortfin

squid. Winter distribution patterns are not well known, but

may include areas east of the Grand Bank. The abundance

of pilot whales declined in Canadian waters in recent decades due to overfishing, and possibly due to changes in

the distribution of preferred prey species. Recent distribution patterns are substantially different than historical ones.

GREENLAND

 Information on pilot whales in Greenland waters is primarily from catch statistics and, more recently, from dedicated observations compiled by the Greenland Fisheries

Research Institute (GFRI) in Nuuk, Greenland, and

Copenhagen, Denmark. Pilot whaling has always been minimal by the Greenlanders, but the catch data provide historical yearly and seasonal information.

Dedicated Sighting Surveys and Catch

Statistics

 Sightings of pilot whales in Greenland occur annually

with the majority of sightings off the western coast south

of Disko Island (Figure 8). The sightings have varied from

individuals to groups of thousands, but the average group

size ranges between 8 and 50 individuals. Only a few observations have been made on the eastern coast of Greenland.

This discrepancy could be due to two factors: 1) East

Greenland has a substantially smaller human population

than West Greenland, and therefore less potential for sighting of pilot whales; and 2) pack ice along East Greenland

lasts longer, limiting the access of pilot whales into inshore

waters.

 The movement of pilot whales appears to be influenced

by oceanographic factors. In summer, pilot whales move

through Greenland waters using deepwater corridors of 200

m or more (Heide-Jørgensen and Bunch 1991). This northwestern movement coincides with the distribution of pilot

whales in northern waters of the eastern North Atlantic

around 62°N in August, September, and October, and around

59°N from May through November (Brown 1961).

 Because dedicated sighting surveys for cetaceans were

not conducted off Greenland prior to the late 1970s, data

from catch statistics provide the only historical information

on abundance and distribution. The GFRI has compiled

whale catch records since 1828 (Table 1). The majority of

pilot whale catches have been taken between Nuuk and

Disko Island. A few whales have been caught south of

Nuuk but only about 1% have been taken in East Greenland.

Most catches occur during June-September (HeideJørgensen and Bunch 1991).

 Dedicated sighting cruises began in 1979, and since 1987

have been supplemented with aerial surveys (Larsen et al.

1989; Larsen and Nielsen 1990). The distribution of pilot

whales from these surveys is consistent with that derived

from historical catch data. Over 97% of the sightings have

been recorded south of Disko Island.

Oceanographic- and Biological-Related

Distribution

 There are indications that the abundance of pilot whales

off West Greenland is linked to prey abundance and fluctuating ocean temperatures. During periods of low water temperatures in the Davis Strait, no pilot whales were caught

(Heide-Jørgensen and Bunch 1991). Sergeant and Fisher

(1957) reported that both northern shortfin squid and pilot

whales move into the Davis Strait from Canada during warm

temperature periods. Historically, shortfin squid have not

inhabited West Greenland waters, but have occurred off

southern Greenland (Roper et al. 1984).

 The same factors that influence the occurrence of pilot

whales also influence the occurrence of other marine species (Heide-Jørgensen and Bunch 1991). A warm period in

the North Atlantic has led to a northward shift in the geographical distribution of many boreal marine animals. These

rising ocean temperatures in West Greenland have caused

the Atlantic cod population to increase, creating a topodeme

that appears to have migration circuits entirely within the

local fjords (Baker 1978). Inversely, when the temperatures

have decreased, the cod population has declined, as have

pilot whale catches and sightings.

Greenland Summary

 Historically, pilot whales have never been very abundant in Greenland. A small percentage of whales in Canada

may venture northward into Greenland waters. That

Greenlanders have never targeted pilot whales as heavily

as other cetaceans provides some evidence that the abundance of pilot whales in Greenland has never been large.

 As in Canada, the presence or absence of preferred prey

may alter the sightings of pilot whales. Water temperature

may also influence the occurrence of prey and pilot whales

in Greenland waters.

ICELAND

 Iceland has been an active whaling nation for many

years, and initial sighting information on pilot whales has

come from observers aboard whaling vessels. More re-
Page 10

Table 1. Total number of pilot whales taken in targeted fisheries by month and by region in Greenland from 1828 to 19911

Month

Region Apr. May Jun. Jul. Aug. Sep. Oct. Total

Nuuk to 0 0 720 1,735 543 561 0 3,559

Disko Island

Nuuk to 0 3 325 90 568 0 60 1,046

Cape Farvel

East Greenland 36 0 52 173 27 275 0 563

Total 36 3 1,097 1,998 1,138 836 60 5,168

______

1

From Heide-Jørgensen and Bunch, 1991.

Table 2. Pilot whale sightings in Icelandic and adjacent waters between 1981 and 1989

Survey Number of

 Date Survey Area Type Sightings Source

Jun. - Jul. 1981 Denmark Strait; Irminger Sea surface 16 Sigurjónsson 1983

Jun. 20 - Jul. 20, 1982 W, SE, & E Iceland aerial 0 Hiby et al. 1984

Denmark Strait; SE-N Iceland surface 5 Martin et al. 1984

Aug. 1983 Denmark Strait surface 2 Sigurjónsson 1985

Jun. 27 - Jul. 20, 1986 All waters surrounding Iceland aerial 7 Gunnlaugsson and

Sigurjónsson 1988

Jun. 24 - Jul. 28, 19871 SE Iceland - Scotland; SW Iceland; surface 71 Sigurjónsson et al. 1989;

Irminger Sea; Denmark Strait Buckland et al. 1993

Jul. 1-20, 19871 Adjacent waters surrounding Iceland aerial 3 Donovan and

Gunnlaugsson 1989

Jul. 10 - Aug. 14, 19891 Icelandic and adjacent waters surface 124 Sigurjónsson et al. 1991

_____

1

Conducted as part of the North Atlantic Sighting Survey. See text for more information on the survey.
Page 11

cently, researchers at the Marine Research Institute in

Reykjavik have collected sighting data from dedicated sighting surveys (Figure 9). Throughout the 1980s, aerial and

shipboard sighting surveys covered the mid-North Atlantic area around Iceland and East Greenland.

Sighting Surveys

 Sigurjónsson and Gunnlaugsson (1990) provide a detailed account of whale sightings made during 1979-88 from

whaling vessels, but these data are biased as sightings of

smaller cetaceans were reported with less accuracy than

sightings of baleen whales. The sighting data are summarized into two time periods: before and after July 15. This

split reflects seasonal whaling times based on the movement of the large baleen whales migrating into Icelandic

waters. The frequency of pilot whale sightings increases

after July 15 when concentrations appear west-northwest

of Iceland. Some whales have been sighted in shallow waters (less than 400 m), but most sightings occur in water

between 400 and 1000 m. As in the shelf waters off the

United States, there appears to be a relationship between

the high concentration of pilot whales found on the Icelandic edge, and hydrographic conditions such as ocean temperature and upwelling (Foerster and Thompson 1985). The

North Atlantic Current flows south of Iceland along the

shelf edge towards Norway, and likely influences pilot whale

movements.

 In June and July 1981, a marking cruise for baleen whales

took place in the Denmark Strait. Pilot whale sightings occurred off the southwestern and western coasts of Iceland

in water depths between 1,000 and 2,000 m. Approximately

500-800 pilot whales were encountered during the 16

sightings of this survey (Sigurjónsson 1983).

 Between 1981 and 1989, Iceland participated in joint

international surveys of the North Atlantic, including the

North Atlantic Sighting Survey (NASS) (Table 2). The NASS

began in 1987 when several nations/territories organized

joint shipboard surveys that were conducted from June

through August that year, and in summer 1989. The area

surveyed was bound by Spitsbergen and the Barents Sea

in the north, the Spanish coast to the south, the West

Greenland coast to the west, and the Norwegian coast to

the east. In addition, an aerial sighting survey occurred in

1987 covering the coastal waters of Norway, West Greenland,

and Iceland in conjunction with NASS-87 (Hiby et al. 1989).

Strandings

 Strandings of pilot whales are rare in Iceland. Only two

mass strandings are known; one in western Iceland at

Rifshofn, and the other in southwestern Iceland at

Thorlakshofn. The Rifshofn stranding occurred in August

1982, when a pilot whale school of approximately 280 individuals congregated near the harbor entrance. The

Thorlakshofn stranding occurred in October 1986, when

143 whales stranded along a gradually sloping beach. Only

a few stomachs were examined, but all were empty

(Sigurjónsson et al. 1993).

Icelandic Summary

 Pilot whales do not occur on the northern side of Iceland, but regularly appear off the southern coast. Pilot

whales prefer the edge of the shelf, with only occasional

movements into shallower waters. Pilot whales have a broad

distribution pattern in the northern Central Atlantic, and

appear to be most abundant in mid-summer.

NORWAY

 In Norway, distribution data on pilot whales have come

from historical whaling records, incidental sightings, and

sighting surveys. These data have been compiled by the

Institute of Marine Research in Bergen since 1967. Norway

was also a participant in the NASS sighting cruises that

covered the northern extreme of pilot whale distribution

(Figure 10).

Sighting Surveys and Incidental

Observations

 Incidental observations of whales within Norwegian

waters took place from 1971 to 1975 aboard whaling vessels. Actual sighting effort depended upon the workload of

the vessel and the observer. Observations in the Barents

Sea occurred during May-June 1973, July-August 1974, and

May-August 1975. The only four sightings of pilot whales

made during these cruises occurred in the Barents Sea in

May 1975 (Christensen 1977). Dedicated sighting surveys

were conducted off Norway from 1987 to 1990 in conjunction with the NASS (Table 3).

 During summer 1991, several groups of pilot whales

were sighted off Lofoten. Animals had been seen in that

region for 3-4 yr in a row (Øien 1991b). Strandings of pilot

whales are relatively rare, and those reported may have

drifted onto Norwegian shores after dying at sea in more

southern waters (Christensen 1990).

Whaling Statistics

 Whaling for small cetaceans began in the 1920s and

spread throughout Norway by the 1930s. Between 1938 and

1974, the number of participating vessels was regulated,

and a compulsory catch report system was in place

(Christensen 1975; Øien 1991b). In 1950, a closed season
Page 12

was implemented during the first 3 wk of July, with additional time limits administered in subsequent years. Though

the catch data reflect these restrictions, the commercial statistics depict in broad terms the patterns of pilot whale distribution. Catch records indicate that pilot whales concentrate in two locations: off Lofoten in northern Norway, and

off Møre in southern Norway, primarily in July and August

(Øien 1991b).

Norwegian Summary

 Pilot whales occur in Norwegian waters, but not in high

abundances. Two areas of concentration are indicated by

catch statistics: along the western coastline at Lofoten and

at More, with most catches reported in July and August.

Dedicated sighting surveys have revealed relatively few

sightings of pilot whales.

Figure 9. Distribution of long-finned pilot whales off Iceland based on sighting data from 1981 to 1989 (Source: Cetacean and Turtle

Assessment Program 1982; Payne et al. 1991; Waring et al. 1992).

Figure 10. Distribution of long-finned pilot whales off Norway based on incidental sightings from 1971 to 1989 and sighting data from 1989

(Source: Cetacean and Turtle Assessment Program 1982; Payne et al. 1991; Waring et al. 1992).
Page 13

Table 3. Pilot whale sightings off Norway from 1987 to 1990

Number of

 Date Survey Area Survey Type Sightings Source

1971-75 Barents Sea Surface 4 Christensen 1977

(whaling vessels)

July 19871 Barents, Greenland, Surface & aerial 0 Øritsland et al. 1989

& Norwegian Seas

July 1988 Barents, Greenland, Surface 0 Øien 1990

& Norwegian Seas

July 19891 Barents, Greenland, Surface 3 Øien 1991a

& Norwegian Seas

July 1990 North Sea Surface 0 Øien 1991b

______

1

Sightings from North Atlantic Sighting Survey. See text for more information on the survey.

FAEROE ISLANDS

 Long-finned pilot whales are commonly sighted around

the Faeroe Islands, and have become an integral part of the

culture of its inhabitants. A drive fishery in the Faeroes has

existed for centuries, with pilot whales being herded into

the many bays of the archipelago. An international research effort conducted from 1986 to 1988 to sample whales

killed in the drives has provided a substantial amount of

information on the life history of this species (Bloch,

Desportes, et al. 1989; Desportes 1993).

 Faeroese involvement in the NASS sighting efforts has

provided additional information on pilot whale distribution.

Since 1986, whale observers have been placed on commercial fishing vessels to record cetacean sightings. These

sightings are being compiled by the Office of the Director

of Fisheries and by the Natural History Museum in

Torshavn.

Sighting Surveys

 Since 1986, observers aboard Faeroese research and inspection vessels have recorded pilot whale sightings (Figure 11) in the Faeroe Islands - Iceland area (Bloch and Hoydal

1988, 1990). During 1987 and 1989, the Faeroese participated in the NASS sighting series with two dedicated sighting cruises. The 1987 survey was in June and July, and the

1989 survey was in July and August. The Faeroese vessel

surveyed an area between southeastern Iceland and western Ireland, bounded by 6°W and 18°W longitude, and 64°N

and 52°N latitude. However, no survey activity was conducted inside the 183-m depth contour in the area of the

British Isles, Ireland, and Iceland.

 There were only 19 sightings recorded during the 1987

survey. Sightings concentrated in three areas: 79% of the

sightings were within the Faeroe Islands archipelago, 5%

were just northeast of the islands, and 16% were just south

of the islands. The 1989 survey had only 11 pilot whale

sightings. These sightings occurred west to southwest of

the islands (45%) and northwest between the islands and

Iceland (55%) (Buckland et al. 1993; Joyce et al. 1991). The

seasonal distribution of pilot whale sightings during these

surveys was consistent with the temporal pattern observed

during the past 280 yr in the Faeroese drive fishery (Joensen

and Zachariassen 1982).

Drive Fishery and Oceanographic- and

Biological-Related Distribution

 The Faeroese drive fishery is the longest continuously

documented harvest of any wild animal, with accounts dating back to 1584 (Hoydal 1987). Although these records

contain minor biases, the historical record provides an account of the presence of pilot whales in and around the

Faeroe Islands for the past four centuries. The "grinds" or

pilot whale drives occur throughout the year, and are most

intensive during July through September when pilot whale

abundance is highest (Hoydal 1987).
Page 14

 A positive correlation exists between the occurrence

of pilot whales and surface water temperatures (Joensen

and Zachariassen 1982; Bloch, Hoydal, et al. 1989). As temperatures increase, the available food for whales increases,

particularly the European flying squid (Todarodes

saggitatus) which is the preferred prey of pilot whales in

the Faeroese region (Desportes and Mouritsen 1993). This

relation is so strong that pilot whales move to deepwater

feeding grounds north of the islands in pursuit of other

squid species if flying squid is not available (Desportes

and Mouritsen 1993). Hoydal (1987) found a high correlation between the abundance of flying squid and the catches

of pilot whales (r = 0.75), implying that pilot whales follow

their prey. This conclusion is similar to that presented by

Sergeant (1962), Mercer (1975), and Heide-Jørgensen and

Bunch (1991).

 In a comparison of Faeroe Islands and Greenland catch

statistics, the peak numbers of pilot whales landed in

Figure 11. Distribution of long-finned pilot whales off Faeroe Islands based on sighting data from 1986, 1987, and 1989 (Source: Cetacean

and Turtle Assessment Program 1982; Payne et al. 1991; Waring et al. 1992).

Figure 12. Distribution of long-finned pilot whales off the British Isles based on sighting data for 1987 and 1989 (Source: Cetacean and

Turtle Assessment Program 1982; Payne et al. 1991; Waring et al. 1992).
Page 15

Greenland occurred in the 1930s, 1960, and between 1974

and 1980 when sea temperatures had increased. The first

and last of those peaks of Greenland landings preceded

periods of good catches in the Faeroe Islands, whereas the

middle of those peaks in Greenland was at the end of a good

catch period in the Faeroe Islands (Hoydal 1987).

Faeroese Summary

 Pilot whales occur regularly in the Faeroese archipelago.

Movement patterns appear to be related to prey distribution and ocean temperature. If their favorite squid prey is

not available, pilot whales leave to feed elsewhere. Most

sightings have occurred on the western side of the islands,

specifically southwest and northwest of the islands towards

Iceland.

BRITISH ISLES

 Information on long-finned pilot whales in waters surrounding the United Kingdom and Ireland has historically

been derived from incidental observations and from

strandings. Spanish and Faeroese vessels surveyed these

waters during the 1987 and 1989 NASS sighting series (Figure 12). Recent sighting data have been obtained through

the 1994 Small Cetacean Abundance in the North Sea Survey (SCANS), with two sightings south of Ireland and one

between the Shetlands and Norway. The 1994 SCANS

sought to estimate abundance and identify summer concentrations of small cetaceans in the North Sea. The survey

was conducted by the Research Unit for Wildlife Population Assessment at Scotland's St. Andrews University.

Weather ships have also reported numerous sightings west

of Ireland and Scotland (Anonymous 1990; Evans 1976; G.

Desportes, pers. comm4

).

Land-Based Observations

 Sightings of long-finned pilot whales are most common

along the northern and southern coasts of Great Britain and

the western coast of Ireland, with the highest numbers concentrated off the northern coast of Scotland and the southwestern coast of England (Evans 1976, 1980). During April

through June, pilot whales congregate in the Irish Sea, and

along the northern and western coasts of England. In November and December, the whales are found off southwestern England and northern Scotland. Pilot whales are present

throughout the year off northern Scotland, with peak abundance during March through May. In southwestern England and in Ireland, numbers are highest in April, June,

and October. Eastern Great Britain has virtually no sighting

records through most of the year, but pilot whales may occur in small numbers in the summer and autumn.

 During fall and winter, pilot whales generally disperse

into smaller schools. In spring, the whales generally form

into larger groups which maintain themselves through September. Pilot whales have usually been sighted individually

or in schools of 11-20 individuals. The largest numbers of

pilot whales in sighted schools were 75 in April 1977 off

northern Wales, and 65 in November 1976 off Cornwall in

southern England (Evans 1980). Such high numbers are

normally only seen in the drive fishery in the Faeroe Islands

during summer. These high numbers suggest that the whales

may form large schools at particular times of the year, such

as during long migrations or at food concentrations (Evans

1976).

 Fraser (1974) suggested that pilot whales occurring in

British waters are part of a population occupying the North

Atlantic Current. This population has a movement eastward in the late months of the year, bringing the schools to

the more southerly parts of the British Isles in winter, but

not to the north until spring and summer. During November

to December, definite peaks in numbers have regularly occurred in northern Scotland and southwestern England, at

least between 1975 and 1980 (Evans 1980).

 Since these peaks occur at the opposite geographical

ends of the British Isles, the possibility of two separate

populations is implied. The northern peak may be the southern extent of the Faeroe Islands population as catches in

the Faeroe Islands are concentrated in July and August,

with few in winter. Pilot whales appear to be aggregated in

waters between the Shetland and Faeroe Islands during

much of the summer, but probably move south toward the

northern coast of Scotland later in the year (Evans 1980). A

relationship may exist between pilot whales reported off

France by Duguy (1977) and those observed off the Cornish coasts in November and December, and occasionally

in the Irish Sea in April.

Strandings

 The majority of observations of pilot whales have occurred from strandings. Strandings are most common on

the southwestern coast of England and Wales, and along

the Atlantic coast of Ireland (Evans 1980; O'Riordan 1972,

1975; Brown 1975). The number of strandings peaked in

1935, then began a decline along with the catches of seafood fish species. The dual declines may have been caused

by a sea temperature rise and water movements (Southward

1963). Strandings along all coasts of Britain have increased

since 1941 (Sheldrick 1976). The frequency of strandings

appeared to be correlated with abundance of seafood fish

species, although it may also have reflected a high mortality due to lack of prey for the whales (Sheldrick 1976).

 Strandings occurred on eastern coasts of Scotland and

England prior to 1947. Since then, there has been an increasing number of strandings on the southern and western coasts of the Britain Isles (Evans 1980). The highest
Page 16

numbers of these strandings have occurred in winter (Fraser

1974). In Ireland, strandings or captures of pilot whales

have occurred in every month except September (O'Riordan

1975).

 Most occurrences of stranded pilot whales have been

single animals; only two large mass strandings have been

reported, one with 100 whales and the other with 148 whales.

Both of these large mass strandings occurred in spring 1950.

Between November 1982 and January 1985, five mass

strandings occurred, involving 2340 whales per stranding.

Three such strandings occurred in northern Scotland and

two in eastern England. The strandings in eastern England

indicate that pilot whales will enter shallow areas in pursuit

of fish, and may strand while following fish into shoal waters. This behavior appears to be similar to that found in the

strandings in Canada and on Cape Cod.

 Fraser (1974) suggested that pilot whales move to more

southerly parts of the British Isles in winter, but this is not

consistent with more recent information by Martin et al.

(1987) indicating strandings in the north in winter.

British Isles Summary

 The waters of the British Isles and Ireland may be a crossroads for the distribution of pilot whales in the eastern North

Atlantic. On both ends of the islands, population peaks

occur at similar times of the year, suggesting that two separate stocks may frequent these waters. Strandings of single

animals are most common, but mass strandings have occurred.

THE NETHERLANDS

 Long-finned pilot whales rarely occur off the coast of

The Netherlands. Since 1594, there have only been 16 recorded events of pilot whales stranding (Husson and Van

Bree 1972, 1976; Smeenk 1986, 1989). All but two strandings

involved individual whales; there was one stranding of 61

in 1856 and another of 31 in 1825. Although there may be no

direct link between pilot whale abundance and the occurrence of stranded whales, the lack of strandings and

sightings suggests that pilot whales do not regularly inhabit Dutch waters.

FRANCE

 There have been occasional sightings and strandings,

and a few incidental kills in commercial fishery operations,

along the coasts of France (Duguy 1977; Evans 1976; Duguy

and Hussenot 1982; Anonymous 1988). Pilot whales occur

primarily along the Atlantic coast, but are also seen along

the eastern sector of the Mediterranean coastline (Duguy

1990). The highest concentration of pilot whales along the

Mediterranean coast occurs in July (Vallon et al. 1976;

Duguy 1990). During winter, pilot whales may also reside in

the Bay of Gascony (Duguy and Aloncle 1975; Duguy 1977).

During 1971-76, a total of 16 strandings occurred on French

coasts. One pilot whale stranded on the English Channel

coast, 12 on the Atlantic coast, and 3 on the Mediterranean

coast (Duguy 1977).

SPAIN AND PORTUGAL

 Distribution data come mainly from dedicated sighting

surveys, as strandings are rare in Iberian waters. Spain initiated the Ballena sighting surveys which were four sighting cruises in the offshore waters of Spain and Portugal

during July-September of 1981-82, and was a participant in

the 1987 and 1989 NASS multinational survey cruises.

Dedicated Sighting Surveys

 Because of coastlines located along the cold waters of

the Atlantic and the warm waters of the Mediterranean,

Spain provides the varied oceanographic conditions for

many species of cetaceans (Casinos and Vericad 1976). Pilot whales are common in Spanish waters, with both

shortfinned pilot whales and long-finned pilot whales occurring. Nores and Pérez (1988) suggest the northern limit

of the short-finned pilot whale is at 45°N, which is within

the Bay of Biscay where long-finned pilot whales have also

been sighted.

 Two series of sighting cruises were conducted during

the 1980s (Figure 13). The four Ballena sighting cruises in

1981-82 lasted upwards of 2 wk per cruise. Sightings of pilot

whales from these cruises were all offshore (Aguilar et al.

1983; Sanpera et al. 1984, 1985; Sanpera and Jover 1986).

However, coastal sightings would have been unlikely because none of the survey tracks were inshore.

 In addition to the Ballena sighting cruises, additional

cruises were undertaken to survey Spanish waters in 1987

and 1989. The NASS-87 cruise, during July 6 - August 3,

1987, covered the areas of western Ireland south to the

Iberian coasts. Only 11 pelagic pilot whales were sighted

(Lens et al. 1989). The NASS-89 cruise, during July 5 - August 9, 1989, covered an area limited by 25°W, the European

continental shelf, and between 42°N and 52°N. For survey

purposes, the area was divided into two parts consisting of

an offshore area and the coastal Bay of Biscay. Pilot whales

were widely distributed throughout both areas, but were

more frequent in the offshore area. The sightings of whales

on the continental slope and in pelagic waters were consistent with sightings made in other surveys. Mixed schools

of predominately pilot whales and shortbeaked common

dolphins (Delphinus delphis) occurred on two occasions

(Lens 1991).

 Strandings of pilot whales in Spain are a rare event;

only four pilot whale strandings occurred during 1973-81

(Grau et al. 1986), and these were all on the Mediterranean
Page 17

coast. There have been no reported strandings of pilot

whales in Portugal.

Spanish and Portuguese Summary

 The distributions of long-finned and short-finned pilot

whales overlap in Spanish waters, with the former species

occurring at least seasonally in the Bay of Biscay. Longfinned pilot whales occur in coastal habitats in the Spanish

Mediterranean and the offshore pelagic zones of Spanish

Atlantic waters. The pelagic distribution in the eastern

Atlantic is consistent with that found in other regions of

the North Atlantic, relating strongly to edges and slopes of

the continental shelf.

Figure 13. Distribution of long-finned pilot whales off Spain based on sighting data from 1981 to 1989 (Source: Cetacean and Turtle

Assessment Program 1982; Payne et al. 1991; Waring et al. 1992).

Figure 14. Distribution of long-finned pilot whales off Italy based on sighting data from 1981 to 1988 (Source: Cetacean and Turtle

Assessment Program 1982; Payne et al. 1991; Waring et al. 1992).
Page 18

ITALY

 The scarcity of literature on cetacean distributions in the

Mediterranean has led to conclusions about long-finned

pilot whaledistribution being founded on occasional

strandings of pilot whales and obscure sightings. Although

systematic studies of cetacean populations in the Mediterranean Sea are lacking, the most common cetaceans are the

common and bottlenose dolphins (Pilleri and Pilleri 1982).

Although pilot whale abundance is unknown, it appears

that the eastward extent of the distribution of pilot whales

in the Mediterranean is in Italian waters; there have been

no regularly recorded sightings or strandings in the eastern

Mediterranean, Adriatic, or Black Seas (Marchessaux 1980

as cited in Evans 1987; Pilleri and Pilleri 1982; Podestà and

Magnaghi 1988).

 Within the waters surrounding Italy (Figure 14), pilot

whales are considered common, but appear to be locally

concentrated in the Ligurian Sea, with undocumented

sightings in the southern seas around Sicily (Cagnolaro et

al. 1983 as cited in Cagnolaro et al. 1986; DiNatale 1983).

Sightings of pilot whales typically are in waters of 1000 in or

more, although they are occasionally very close to the coast

(Podestà and Magnaghi 1988). Virtually all of the recorded

cetacean sightings have been from crews aboard personal

yachts (Pilleri and Pilleri 1982).

 There are no records of mass strandings of pilot whales

in Italian waters. The stranding events that have occurred

have been of individual pilot whales. Many of the whales

that have washed up have been decomposed, indicating

that they died at sea (Cagnolaro et al. 1986; Centro Studi

Cetacei 1989, 1990).

 Recently, there has been a decline in the number of

sightings in areas that were previously regarded as the most

populated. This decline sparked a proposal for pilot whale

research by the Committee of Marine Vertebrates and Cephalopods of the International Commission for the Scientific

Exploration of the Mediterranean Sea (Podestà and

Magnaghi 1988).

 Bycatches of pilot whales have occurred in the pelagic

driftnet fisheries in the Ligurian Sea. In 1988, 10 pilot whales

were incidentally caught in the swordfish (Xiphias gladius)

driftnet fishery. There are possibly more being taken, as

many go unreported, die, or sink, or are purposely sunk by

fishermen to avoid charges, as Italian law protects cetaceans (Notarbartolo di Sciara 1990).

Italian Summary

 Distribution of pilot whales is concentrated within the

Ligurian Sea, based on the limited number of sightings and

driftnet bycatch. Unrecorded sightings have occurred in

and around Sicily. Individual pilot whales have washed

ashore in a decomposed state, indicating death at sea.

DISCUSSION

 The long-finned pilot whale is widely distributed in the

North Atlantic, with sightings reported in a broad band

from roughly 35°N to 65°N in the western Atlantic, and from

40°N to 75°N in the eastern Atlantic (Figure 1). There are no

major gaps within this overall distribution where pilot whales

have never been reported. Where sightings have been infrequently reported, for example in the central North Atlantic along the Mid-Atlantic Ridge, there has been little

searching effort.

 The long-finned pilot whale occurs regularly along the

continental shelf break, usually in deeper more pelagic waters, but also in the open ocean far removed from continental land masses. It occurs around several islands, especially

Greenland, Iceland, and the Faeroe Islands, again at the

island shelf breaks in deeper waters.

 The distribution of long-finned pilot whales overlaps

with that of the short-finned pilot whales along the U.S.

coast in the region from 35°N to 40°N, and along the French

and Spanish Atlantic coasts from roughly 40°N to 45°N.

The distributional overlap in the Central Atlantic is unknown. The relationship of the long-finned pilot whales

reported from Spanish and Italian Mediterranean waters to

those in the Atlantic is not clear.

 A north-south seasonal movement pattern has been suggested for the long-finned pilot whale in nearly all areas,

although the Faeroe Islands drive fishery occurs year-round,

and in the eastern United States where seasonal sighting

surveys have been conducted, animals are present in all

seasons (Figures 3-6).

 Squid are the preferred prey in all locations where information is available, although Atlantic mackerel and Atlantic cod may also be significant prey items on occasion. Pilot

whales may change their distribution patterns as preferred

prey species change in abundance.

 Directed harvesting of pilot whales has occurred in three

main areas -- on Cape Cod, in Newfoundland, and in the

Faeroe Islands -- illustrating persistent distribution patterns. At present, a consistent harvest occurs in the Faeroe

Islands; aboriginal takes occur occasionally in other areas.

Bycatch of this species has occurred in recent years in

Atlantic mackerel and squid trawl fisheries of the eastern

coast of the United States, and in swordfish drift gillnet

fisheries in both U.S. waters and in the Mediterranean Sea.

 The species is known to strand naturally, often in large

numbers, in several areas, especially along the northern

shore of Cape Cod.

 One major information gap in our understanding of distribution patterns is the low level of sighting effort in the

central North Atlantic, especially seaward of the continental shelf break in U.S. waters, and between 30°W and 50°W

between 40°N and 60°N. Another information gap is longterm diet history of pilot whales. Studies by Evans (1980),

Desportes (1985), and Desportes and Mouritsen (1993) have
Page 19

focused on stomach contents, reflecting short-term diet history. Information on long-term diet history of pilot whales

of the eastern United States has been reported by Abend

and Smith (1995, 1997).

 Given the lack of clear breaks in the distribution of pilot

whales across the North Atlantic, there is no basis from

these data to suggest that there are separate populations in

the Atlantic. The situation is less clear between the Mediterranean and the Atlantic. On the other hand, given the

occurrence of the species year-round in several distant

portions of the Atlantic, there is also no evidence of a single

population.

ENDNOTES

1. Available from: Tim D. Smith, National Marine Fisheries

Service, Woods Hole, MA.

2. Memorial University of Newfoundland, St. Johns, NF.

3. ibid.

4. Museum of Natural History, Torshavn, Faeroe Islands.

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