MARINE MAMMAL SCIENCE, 15(2):366-380 (April 1999)

0 1999 by the Society for Marine Mammalogy

OCCURRENCE, DISTRIBUTION, SITE

FIDELITY, AND SCHOOL SIZE OF
BOTTLENOSE DOLPHINS
(TURSIOPS T R U N C A T U S ) OFF
SAN DIEGO, CALIFORNIA
R. H. DEFRAN
DAVIDW. WELLER~
Cetacean Behavior Laboratory,
Department of Psychology,
San Diego State University,
San Diego, California 92182-4611, U S A .
E-mail: [email protected]

ABSTRACT
The occurrence, distribution, site fidelity, and school size of bottlenose
dolphins (Ttlrsiops trancattls) in the coastal waters of north San Diego County,
California were assessed during a six-year boat-based photoidentification
study. A total of 146 photographic surveys were conducted between January
1984 and December 1989. Dolphin schools were encountered on 79% of
all surveys, and 2,869 individuals were observed in 145 separate schools.
Three-hundred seventy-three dolphins were individually identified. All
schools were sighted within 1 km of shore, and more than two thirds of
the schools were encountered in the southern half of the 32-km long study
area. School size (mean = 19.8, SD = 18.40) and the number of dolphins
encountered per survey (mean = 26.8, SD = 22.30) were highly variable.
Low resighting rates of known individuals provided little evidence for long-
term site fidelity. When our six-year photoidentification database was com-
bined with previous data, 404 dolphins were identified in the study area
from September 1981 to December 1989. Jolly-Seber population estimates
during the 1984-1989 study period varied between 234 and 285. The
combination of regular dolphin occurrence, low site fidelity by known in-
dividuals, and the continuous increase in the rate at which new dolphins
were identified indicates that numerous different individuals were visiting
the study area across and within years. The open California coastline differs
in habitat structure and prey distribution from more protected study areas
where bottlenose dolphins display site fidelity. These habitat differences may

Current Address: Marine Mammal Research Program, Texas A&M University, 4700 Avenue
U, Building 303, Galveston, Texas 77551-5923, U.S.A.

366
 DEFRAN A N D WELLER: BOTTLENOSE DOLPHINS 367

help to explain the observed intraspecific behavioral variability of this spe-

cies.
Key words: bottlenose dolphins, Tursiops truncatus, California, photoidentifi-
cation, occurrence patterns, distribution, site fidelity, school size.

Investigations of geographically distinct populations of bottlenose dolphins

(Tursiops truncatus) have clarified striking and subtle intraspecific behavioral
differences commonly interpreted as specific adaptations to local ecological
conditions (e.g., Wiirsig and Wursig 1977, 1979; Wiirsig 1978; Wells 1986;
Ballance 1990, 1992; Shane 1990; Bearzi e t al. 1997). The reported variation
in patterns of occurrence, distribution, school size, and site fidelity of bottle-
nose dolphins has been suggested to reflect differing habitat characteristics
(Ballance 1992). This apparent interplay between environment and behavior
is particularly evident when bottlenose dolphins occurring in protected marine
environments are compared to communities living in more open habitats
(Shane et a f . 1986, Wells et a f . 1987, Ballance 1992). Protected habitats have
been found to foster relatively small school sizes, some degree of regional site
fidelity, and limited movement patterns (Wells et af. 1987, Shane 1980). In
contrast, semi-open habitats often sustain larger school sizes, diminished levels
of site fidelity, and more expansive home ranges (Wursig 1978, Ballance 1992,
Bearzi et al. 1997).
The research presented here provides new information on the biology of
bottlenose dolphins occurring in the open and fully exposed nearshore habitat
of southern California. This coastline is quite dissimilar from regions where
most other long-term studies on this species have been conducted and provides
the opportunity to further evaluate the influence of habitat on bottlenose dol-
phin behavior.

METHODS
Study Area

The San Diego study area extended along 32 km of coastline from Scripps
Pier, La Jolla (32"52'N) north to South Carlsbad State Beach (33'08") (Fig.
1). This coastline is located near the center of the Southern California Bight,
a 732-km stretch of coastline extending from Point Conception in the north
to Punta Colnett in the south. The sharp eastern break in the California
coastline south of Point Conception marks a conspicuous change in the coastal
climate and marine fauna due to the interruption of the California Current
system (Southern California Coastal Water Research Project (SCCWRP) 1973,
Dailey et al. 1993, Hickey 1993). The departure of the California Current
from the coast results in a warming of the water within the Bight and creates
the Southern California Countercurrent, The most significant effect of this
nearshore countercurrent is the creation of a region in which northern, south-
ern, western, and upwelling bottom waters converge to form a variable ocean-
368 MARINE MAMMAL SCIENCE, VOL. 15, NO. 2 , 1999

Figure 1. Map of San Diego study area. Inset shows relative location along Cali-
fornia coastline.
 DEFRAN AND WELLER: BOTTLENOSE DOLPHINS 369

ographic and biological coastal ecosystem (SCCWRP 1973, Dailey et al. 1993,
Hickey 1993).
Beaches in the San Diego study area offer little natural protection from
oceanic conditions and are variable in composition, including gently sloping
sand, steeply inclined cobblestone, estuary mouths, and rocky outcrops. Near-
shore underwater topography ranges from submerged reefs, sea grass flats, and
dense kelp canopies to relatively barren, sandy expanses. Vessel traffic within
the study area is minimal and consists primarily of lobster and gillnet fish-
ermen and occasional recreational anglers.

Photoident$cation Surveys
Photographic surveys involved travel in a 4.3- or 5.2-m boat moving par-
allel to the coast, 90-180 m offshore of the surf line. All surveys were con-
ducted in Beaufort sea state 1 3 and under visibility conditions adequate for
finding and photographing dolphins. Two to four observers searched the area
from the shore to 2 km offshore until a school of dolphins was sighted. To
ensure that all members of a spatially separated school were located, the re-
search vessel progressed approximately 1 km past these dolphins and then
motored to a vantage point approximately 60-90 m from the school. Initial
estimates of school size, as well as information on time, location, environmen-
tal conditions, and behavior were then noted.
During most surveys, dolphins alternated between coalescing into a larger
school and separating into smaller subgroups. These smaller subgroups usually
remained within 500 m of each other and were considered to be part of the
larger school from which they originated. Thus, a school was defined as any
dolphins observed in close proximity to one another and usually moving in
the same direction and engaged in similar behavior. School-size estimates were
based on a consensus among all trained observers on board the research vessel
and represented the maximum estimate during an observation period.
Following initial estimates of school size, the vessel maneuvered to within
3-12 m of the school, and individual dorsal-fin notch patterns were photo-
graphed with 35-mm motor driven cameras, a 400-mm f4.0 telephoto lens
and Kodak Tri-X film. Attempts were made to photograph every dolphin
within a school. Initial estimates of school size were revised as necessary, and
contact with the school was maintained until photographic effort was com-
pleted. The research vessel then motored offshore, where all film was labeled
with the date, location, and school number. Identical procedures were repeated
as the research vessel resumed travel on the predetermined survey route and
as additional dolphin schools were encountered. Upon reaching the northern
terminus of the study area, the research vessel motored offshore back to Scripps
Pier. These offshore return trips varied from 1 to 4 km from shore and per-
mitted us to opportunistically sample for bottlenose dolphins occurring out-
side the nearshore coastal study area.
Two survey types were conducted during the study: complete and partial.
Complete surveys covered the entire 32-km study area. Partial surveys, short-
370 MARINE MAMMAL SCIENCE, VOL. 15, NO. 2, 1999

ened by inclement weather or equipment failure, covered only a portion of the

study area and always encountered at least one dolphin school.

Photoidentijication
Our method for analyzing dorsal-fin photographs has been detailed else-
where (Defran et al. 1990). Briefly summarized, only clear photographs of
distinctive dorsal fins were used to establish a “type specimen” to which all
other photographs were compared. Subsequently, only unambiguous matches
with the “type specimen” were accepted as resightings. Two modifications were
made in our application of this procedure: (1) the criteria used to categorize
a fin as distinctive became more conservative as the size of the photoidentifi-
cation catalog grew; (2) all photographs were examined by two, and more
commonly, three people. As a consequence of these methodological refine-
ments, minor changes in the summary statistics from earlier stages of our
work (Defran et al. 1986, Hansen and Defran 1990) have resulted. Important
population parameters and the site-fidelity characteristics described in these
earlier summaries, however, remained unchanged.
Our initial photographic catalog consisted of dorsal-fin photographs of 114
individuals which met our criteria for distinctiveness. One hundred and three
of these dolphins were selected from those photographed by Hansen (1990)
between September 1981 and January 1983, and 11 were selected from those
photographed by Hansen and his associates between August 1983 and No-
vember 1983 at the National Marine Fisheries Service Southwest Fisheries
Science Center (NMFS). Unless otherwise noted, all photographs taken be-
tween September 1981 and November 1983 are referred to as the NMFS
sample, while photographs taken between January 1984 and December 1989
are referred to as the Cetacean Behavior Laboratory (CBL) sample. Results from
the NMFS sample have been summarized by Hansen (1990). Thus, results
presented here are mostly from the six-year CBL sample. For several analyses,
however, the NMFS and CBL data sets were combined to form a nine-year
sample.

RESULTS
Survey Eflort
One hundred and forty-six boat-based photoidentification surveys were con-
ducted in the San Diego study area from January 1984 to December 1989.
The mean interval between surveys was 15 d and the mean interval between
surveys on which at least one dolphin was photographed was 22 d. Approxi-
mately 200 h were spent in direct observation and photography of 2,869
dolphins in 145 separate schools. Observations on individual schools averaged
83 min.
Complete surveys accounted for 75% (n = 109) of all surveys conducted.
Sixteen of these extended to Oceanside (33’12’ N), 7.4 km beyond the north-
 DEFRAN AND WELLER: BOTTLENOSE DOLPHINS 371

60 -
r
gg 50-
E
s'L1 40-
8'9
-U
0 2 30-
ZP,
E a 20-
If
w 10-
I
0

ern terminus of our study area (Fig. 1). Partial surveys constituted 25% (n =
37) of all surveys. Twenty-five partial surveys were terminated in the southern
half of the study area (<33"00'N), while 12 surveys extended by varying
distances into the northern portion of the study area. Most partial surveys
(84%) were carried out between 1984-1986. Only one of the 29 complete
surveys between 1984-1986 encountered more than one dolphin school, sug-
gesting that the single schools encountered on partial surveys were likely to
have been the only school in the study area.

Occurrence
Dolphin schools were encountered on 79% ( n = 116) of all surveys. En-
counter percentages varied annually and ranged from a low of 60% in 1987
to a high of 95% in 1989. Of the 7 9 complete surveys on which dolphin
schools were sighted, 75% ( n = 59) encountered only one school. Multiple
schools (range = 2-5) were sighted on 25% ( n = 20) of these 7 9 surveys.
Seventy percent ( n = 14) encountered two schools, 20% ( n = 4) encountered
three schools, and 10% (n = 2 ) included either four or five schools. The mean
number of dolphins encountered on a complete survey varied considerably both
within and across years (Fig. 2).

SeusonaZity
Survey effort, expressed as the number of surveys, was similar in all four
seasons: winter (December-February; n = 32), spring (March-May; n = 38),
summer (June-August; n = 41), fall (September-November; n = 35). Dol-
phins photographed 10 or more times between January 1984 and December
1989 ( n = 38) were used to test for evidence of seasonal occurrence. We
restricted our analysis to this subset of frequently sighted individuals to reduce
the possibility of creating spuriously high seasonal trends by inclusion of dol-
372 MARINE MAMMAL SCIENCE, VOL. 15, NO. 2 , 1999

15
10

1984 1985 1986 1987 1988 1989

Year
Figure 3. Mean school size by year. Error bars represent standard deviations.

phins sighted infrequently and over short periods of time. No evidence of

seasonal occurrence patterns was apparent from this sample. Eighty-seven per-
cent ( n = 33) of these 38 frequently sighted dolphins were photographed in
all four seasons, while the remaining 13% ( n = 5) were photographed in at
least three different seasons.

Distribution
Sightings of bottlenose dolphins were distributed throughout the study
area. These dolphins were the only odontocetes encountered during coastal
surveys. All dolphin schools were sighted within 1 km of shore and most often
were within 250 m of the shoreline. No bottlenose dolphins were encountered
on our offshore return route.
The majority of dolphins (70%) and dolphin schools (67%) were sighted
in the southern portion of the study area between Torrey Pines State Park
(32'52' N) and Solana Beach (33"OO' N) (Fig. 1). Similarly, when we analyzed
only complete surveys, 60% of sightings occurred in the southern portion of
the study area, indicating that this trend was not an artifact of our south-to-
north survey strategy.

School Size, Site Fidelity, and Population Size

Mean school size ranged from 12.7 in 1986 to 28.8 in 1988 with an overall
mean of 19.8 individuals (range = 2-90, SD = 18.40) (Fig. 3). School sizes
were highly variable. Many schools (22%) contained between two and five
dolphins (n = 32), and 75% ( n = 109) were composed of 25 or fewer dolphins
(Fig. 4). On no occasion was a solitary dolphin observed.
A total of 373 individuals were photographically identified in the study
area from 1984 to 1989. When the study period was extended to include the
27 mo of NMFS data, the number of identified individuals increased to 404
(Fig. 5). The rate at which individual dolphins were first identified (rate of
 DEFRAN AND WELLER: BOTTLENOSE DOLPHINS 373

35

School Size
Figwe 4. Frequency distribution of school size.

discovery) was analyzed by partitioning the combined NMFS and CBL pho-
tographic data sets (1981-1989) into blocks of 10 consecutive surveys in
which at least one dolphin was identified (n = 125 surveys, Fig. 5). Rate of
discovery was plotted as both the percent of new sightings per 10 surveys,
and the cumulative number of new individuals identified. The cumulative rate
of discovery curve showed a continued increase in the number of dolphins
identified from survey blocks 1-12 (Fig. 5). One of the greatest increases in
the number of new dolphins identified occurred late in the study period be-
tween blocks 9 and 10. By the end of 1989 (block 12), however, this function
appeared to be asymptotic and only 4% of the dolphins photographed were
"new."

Sighting frequencies for the 373 dolphins identified during the study

"
I 2 3 4 5 6 7 8 9 1011 12
Blocks of Ten Surveys

Figure 5. Percentage of new sightings and rate of discovery curve for dolphins
photographed in San Diego from 1981 to 1989. National Marine Fisheries Service
(NMFS) (1981-1983) and.CBL (1984-1989) data sets plotted in blocks of 10 surveys.
Only surveys on which at least one identifiable dolphin was photographed were used.
Blocks 1 and 2 contain data from 26 Hansen and NMFS surveys. Block 12 contains
data from only nine surveys.
374 MARINE MAMMAL SCIENCE, VOL. 1 5 , NO. 2 , 1999

100, t

90 -
2 80
-
B
E
70
P 60
50
3
c
40
30
ki
0 20
5 10
-
0
1 2 3 4 5 6 7 8 9 10 11 12 13 15 16 17 18 24
Number of Times Photographed

Figure 6. Sighting frequencies for dolphins identified from 1984-1989.

ranged between 1 and 24 (mean = 4.6) (Fig. 6). The mean interval between
resightings was 203 d, and many dolphins (24%) were sighted only once.
Sixty-six percent (n = 245) of dolphins identified were resighted fewer than
six times, an average of less than one sighting per year.
Sightings-per-opportunityratios, which reflect sighting frequency as a func-
tion of effort, were calculated for dolphins “frequently” photographed (I six
times). Ratios were derived by dividing the number of sightings for an in-
dividual dolphin by the number of surveys (opportunities) on which at least
one dolphin was photographed (n = 99) over the six-year study period. The
mean number of sightings per opportunity for these “frequently” sighted dol-
phins was 0.09 (SD = 0.030). This low mean value would have been further
reduced had the calculation included surveys on which no dolphin schools
were sighted.
Hansen (1990) reported that 25% of the dolphins he identified were pho-
tographed only once. We examined the CBL photographic data set for the
presence of the 103 dolphins we retained from the Hansen (1990) data set.
Thirty (29%) of the dolphins first identified by Hansen in 1981-1983 were
not photographed during the 1984-1989 study period. Among the subset of
dolphins Hansen (1990) most frequently sighted (5-9 sightings), four were
never photographed by us and six were sighted less than once per year. One
of Hansen’s most frequently sighted dolphins, however, was also our most
frequently sighted dolphin.
Multiple mark-recapture estimates were calculated using the Jolly-Seber
population estimator golly 1965; Caughley 1977; Seber 1982; Krebs 1985,
1989).This estimator, which is appropriate for open populations, was selected
because of the low resighting frequencies (Fig. 6) and low sightings-per-op-
portunity ratios for dolphins in this study. Additional findings on coastal
movement patterns (Defran e t al. 1999) indicate that dolphins photographed
in San Diego are part of a larger open population. Dolphins identified in San
Diego in 1984 (n = 98) were used as the marked individuals, and those
identified each year between 1985 and 1988 were used as the recaptures.
 DEFRAN AND WELLER: BOTTLENOSE DOLPHINS 375

Results from these calculations yielded the following annual population esti-
mates and associated 95% confidence intervals: 1985 = 237 (CI 214-262),
1986 = 234 (CI 205-263), 1987 = 285 (CI 265-306), 1988 = 284 (CI
274-294).

DISCUSSION
The behavioral ecology of most animals is thought to reflect specific adap-
tive strategies shaped, in part, by the unique ecological conditions under which
a species or population exists. Bottlenose dolphins have now been studied in
a variety of marine environments, with the most detailed and long-term in-
formation resulting from the work of Wells (1986) and his colleagues in Sar-
asota Bay, Florida (Wells et a/. 1987). The West Florida barrier-island habitat
differs significantly from the open coastal environment of San Diego. Differ-
ences in patterns of site fidelity, school size, and population size between
Florida and San Diego may be related to these habitat differences. While
information on bottlenose dolphins in other habitats similar to San Diego has
been collected (Wursig and Wursig 1977, 1979; Wursig 1978; Ballance
1990, 1992), the relatively short-term nature of these studies limits longitu-
dinal comparison with the present research.

School Size
School sizes for coastal bottlenose dolphins range from one to over 100
individuals but are most commonly 2-15 dolphins (Shane et al. 1986). The
mean school size for bottlenose dolphins in San Diego (19.8) was larger than
for most other coastal bottlenose dolphin populations. School sizes reported
for this species are variable, due in part to differing definitions of a “school”
(Shane et al. 1986). Coastal bottlenose dolphins in Argentina (Wursig 1978),
the Gulf of California (Ballance 1990), and Portugal (dos Santos and Lacerda
1987) assemble in schools similar in size to those observed in San Diego. In
contrast, considerably smaller mean school sizes (range = 3-7) have been
reported for study areas in Texas (Shane 1980, Gruber 1981, Henningsen
1991, Brager 1992, Fertl 1994), the northern Adriatic Sea (Bearzi et a/. 1997),
and Florida (Wells 1986, Shane 1990).
Weller (1991) examined the social ecology of bottlenose dolphins in San
Diego and concluded that the relatively large school sizes for dolphins in this
area may be a response to a generally patchy distribution of primary prey
species within the Southern California Bight (Dailey et a/. 1993). Variations
in school size were further hypothesized to serve as a sociobiological adaptation
to permit exploitation of variable and unstable food resources.

Rate of Discovery
The rate at which previously unidentified dolphins were discovered re-
mained relatively constant across the duration of the study, only approaching
376 MARINE MAMMAL SCIENCE, VOL. 1 5 , NO. 2 , 1999

a possible asymptote by 1989 (Fig. 5, blocks 10-12). Without further pho-

tographic data it is impossible to determine if this trend would have persisted.
The continuous increase in the slope of the discovery curve may indicate that
dolphins in the San Diego study area belong to a population that is larger
than previously estimated (Hansen 1983). Hansen (1983) calculated a closed
population estimate of 173-240 bottlenose dolphins in San Diego. The growth
in the number of recognizable dolphins over time, however, along with con-
siderations raised in the following sections, suggests that this population is
more likely to be open (size of population may change during study period)
than closed (see Jolly 1965; Seber 1982; Krebs 1985, 1989 for a review of
distinctions between open and closed populations).
Results of studies on bottlenose dolphins from other regions suggest that
rate of discovery curves for new individuals reach an asymptote over shorter
periods of time. The shape of such acquisition curves is, of course, related to
the size and distribution of the population under study, the geographic scope
of surveys, and the number of days in the field. Wells (1986) reported that
after the initial 27 d of field work in Sarasota, Florida, a majority of the
dolphins identified during the entire study had been photographed. Ballance
(1990) reported similar findings for bottlenose dolphins in the Gulf of Cali-
fornia, with most individuals identified approximately one month into the
study. Finally, Shane (1987) reported that the rate of discovery for bottlenose
dolphins at Sanibel Island, Florida, began to reach an asymptote by the end
of her one-year study. It is important to consider, however, that the studies of
Ballance (1990) and Shane (1987) were of limited duration and may not be
directly comparable to the long-term work of Wells (1986) or the research
presented here.

Population Size
Bottlenose dolphins in San Diego appear to belong to an open population
estimated to range in size from 234 to 284 animals. Our population estimates
closely match bottlenose dolphin abundance estimates recently derived from
tandem aerial surveys in southern California (Carretta et al. 1998). Both of
these estimates, however, may be slightly conservative. Aerial survey work
(Carretta et al. 1998) covered only a portion of the known range for California
coastal bottlenose dolphins (Defran et al. 1999). Photoidentification studies
determined that approximately 65% of all dolphins observed had distinctively
marked fins (Hansen and Defran 1990). Therefore, if the occurrence and dis-
tribution patterns of unmarked dolphins are similar to those of marked dol-
phins, the total number of identified dolphins may underestimate the popu-
lation size by at least 35%.

Site Fidelity and Distribution

Bottlenose dolphins occur throughout the year in the nearshore waters of
San Diego but are not year-round or seasonal residents. Numerous lines of
 DEFRAN AND WELLER: BOTTLENOSE DOLPHINS 377

evidence suggest that dolphins are essentially transient within the San Diego
study area. Individual sighting frequencies (Fig. 6) are low across the six-year
study period, as are sightings-per-opportunityratios. Also, the high variability
in the number of dolphins encountered during a survey, both within and across
years (Fig. 2), indicates that many dolphins present on some occasions were
not present on others. Similarly, on numerous surveys there were no dolphins
observed in the study area, especially during those years with low daily en-
counter probabilities (e.g,, 1984, 1985, 1987).
Hansen (1990) concluded that the San Diego study area served as part of a
permanent year-round home range for these dolphins and as part of a seasonal
home range for others. Most of the dolphins Hansen identified, however, were
sighted only once or twice during our study. Furthermore, the 21 dolphins
Hansen identified as showing site fidelity to the San Diego study area were
sighted on less than half of his surveys. In fact, many of these dolphins ac-
cumulated a high number of resightings over short periods of time (<1 mo)
suggesting a pattern of short rather than long-term fidelity to the area.
All schools encountered during photoidentification surveys were within 1
km of shore, and no dolphin schools were ever encountered 1-4 km offshore
during our return surveys. Similarly, Hansen (1990) always found dolphin
schools within 1 km of shore during boat-based and aerial surveys. Carretta
et al. (1998) reported that 87% of all dolphin groups sighted during aerial
surveys were within 300 m of shore, and Hanson and Defran (1993) reported
that 99% of all dolphin sightings during a 12-mo cliff-based behavioral study
were within 1 km of shore. This nearshore distribution pattern coincides with
water depths ranging between 10 and 30 m. Wiirsig and Wiirsig (1979) found
that bottlenose dolphins off Argentina showed a preference for water less than
10 m deep, and Ross and Cockcroft (1990) reported similar preferences for
coastal bottlenose dolphins off southern Africa, where most sightings occurred
in water depths of 15-30 m.
When our distributional data collected nearshore are combined with find-
ings of Barlow (1993) and Shane (1994) the probable dimensions of the pre-
ferred habitat for California coastal bottlenose dolphins are further refined.
Barlow (1993) conducted systematic boat-based surveys of California waters
up to 555 km offshore and found bottlenose dolphin schools to be concentrated
near and among the offshore Channel Islands. Shane (1994) reported no match-
es when photographs of recognizable bottlenose dolphins from Catalina Island
(-42 km offshore) were compared to identified individuals from San Diego.
We propose, therefore, that a potential distributional gap exists between in-
shore bottlenose dolphins and those found near the offshore islands within the
Southern California Bight. Kenney (1990) reported similar evidence for a dis-
tributional gap between inshore and offshore bottlenose dolphins off the north-
eastern United States. The preferred habitat of bottlenose dolphins off San
Diego is a narrow “coastal corridor” which extends at least the 32-km length
of our study area, with an offshore boundary no farther than 1 km from shore.
Movement pattern data presented in Defran et al. (1999) suggest that this
corridor also extends well to the north and south of San Diego.
378 MARINE MAMMAL SCIENCE. VOL. 1 5 , NO. 2 . 1999

Bottlenose dolphins in the nearshore waters of San Diego differ in their

distribution, site fidelity, and school size from other coastal populations of this
species. Differences in habitat structure and prey distribution between the
open California coastline and the more protected study areas where bottlenose
dolphins display some site fidelity and smaller school sizes undoubtedly con-
tribute to these reported intraspecific differences.
A recent extension of the research presented here has been the incorporation
of photoidentification studies of coastal bottlenose dolphins at other locations
within the Southern California Bight (Defran et al. 1999). Most of the bot-
tlenose dolphins identified in these study areas have also been photographed
in San Diego, suggesting an extensive coastal home range for this population.

ACKNOWLEDGMENTS

This research was carried out under the authorization of National Marine Fisheries
Service Permit 387 and 619. The authors wish to acknowledge the important field
work and laboratory contributions of a number of individuals. During the early years
of this work, A. Weaver was a valued partner in our field work, as was G. Shultz who
also refined and managed our photoidentification activities. In more recent times, J.
Scott, M. Caldwell, and A. Kesaris assisted in the photoidentification process, L. Quig-
ley and E. Tepper verified many of our calculations, and A. Acevedo, L. Ballance, and
one anonymous reviewer gave us thoughtful and thorough comments on earlier drafts
of the manuscript. Our gratitude is also extended to J. Barlow who advised us on
population estimates, to B. Wiirsig for laboratory support, and the National Marine
Fisheries Service, Southwest Fisheries Science Center, La Jolla, for camera equipment
indispensable to our research. Special appreciation is extended to L. Hansen who pa-
tiently introduced us to this work, gave us access to his photographic data set, and
provided important feedback on our evolving interpretation of the data. Finally, we
acknowledge the loving contributions of Barbara J. Weller (1938-1992) to the authors,
and we dedicate this article to her.

LITERATURE
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Received: 12 June 1995
Accepted: 27 July 1998