OCCURRENCE PATTERNS, SITE FIDELITY, AND MOVEMENTS

OF PACIFIC COAST BOTTLENOSE DOLPHINS

(TURSIOPS TRUNCATUS) IN THE

SOUTHERN CALIFORNIA BIGHT

_______________

A Thesis

Presented to the

Faculty of

San Diego State University

_______________

In Partial Fulfillment

of the Requirements for the Degree

Master of Science

in

Interdisciplinary Studies: Animal Behavior

_______________

by

Aimée R. Lang

Spring 2002
THE UNDERSIGNED FACULTY COMMITTEE APPROVES

THE THESIS OF AIMÉE R. LANG:

___________________________________________ ____________
R. H. Defran, Chair Date
Department of Psychology

___________________________________________
Vanessa Malcarne
Department of Psychology

___________________________________________
David W. Weller
Department of Biology

SAN DIEGO STATE UNIVERSITY

Spring 2002
 iii

ACKNOWLEDGEMENTS
 iv

TABLE OF CONTENTS

PAGE
ACKNOWLEDGEMENTS..................................................................................................... iii
LIST OF TABLES.................................................................................................................. vii
LIST OF FIGURES ............................................................................................................... viii
INTRODUCTION .....................................................................................................................1
Space-Use Patterns………………………………………………………………………...1
Ecological Factors..........................................................................................................2
Internal Constraints........................................................................................................5
Pacific Coast Bottlenose Dolphin Research ........................................................................6
Objectives ..........................................................................................................................10
METHODS…. .........................................................................................................................12
Photographic Survey Procedure.........................................................................................16
Photo-Identification Procedures.........................................................................................17
Computer Analysis.............................................................................................................19
Occurrence Patterns .....................................................................................................20
Distribution ..................................................................................................................21
Site Fidelity..................................................................................................................21
Movements...................................................................................................................22
Rate of Discovery Curves ............................................................................................22
RESULTS ……........................................................................................................................24
Occurrence .........................................................................................................................24
Distribution ........................................................................................................................25
Site Fidelity........................................................................................................................25
Movements.........................................................................................................................30
Rate of Discovery ..............................................................................................................32
Comparisons with Past Studies..........................................................................................32
Occurrence Patterns .....................................................................................................32
El Niño .........................................................................................................................37
 v

PAGE
RESULTS (continued)
Distribution ..................................................................................................................40
Site Fidelity..................................................................................................................40
Movements...................................................................................................................43
Rate of Discovery ........................................................................................................43
DISCUSSION ..........................................................................................................................46
Occurrence Patterns ...........................................................................................................47
Distribution ........................................................................................................................51
Site Fidelity........................................................................................................................54
Movements.........................................................................................................................57
Rate of Discovery Curve....................................................................................................59
Comparisons with Other Study Areas................................................................................61
CONCLUSIONS......................................................................................................................66
REFERENCES ........................................................................................................................69
APPENDIX
PHOTOGRAPHIC QUALITY CRITERIA........................................................................79
ABSTRACT ..........................................................................................................................82
 vi

LIST OF TABLES

TABLE PAGE
1. Habitat Features of Study Area Zones in San Diego and Santa Barbara........................29
2. Summary Information on Survey Effort, Study Period, and Photographic
Results from San Diego Between 1984 and 1989...........................................................34
3. Summary Information on Survey Effort, Study Period, and Photographic
Data for Southern California Bight Study Areas 1982-1999 ..........................................44
 vii

LIST OF FIGURES

FIGURE PAGE
1. Map of the Southern California Bight .............................................................................13
2. Map of the San Diego Study Area...................................................................................14
3. Map of the Santa Barbara Study Area.............................................................................15
4. Computation of the Dorsal Ratio.....................................................................................18
5. Mean Number of Dolphins Observed per Complete Survey as a Function of
Oceanographic Season in San Diego and Santa Barbara between April 1998
and August 1999..............................................................................................................26
6. Mean School Size as a Function of Oceanographic Season for San Diego and Santa
Barbara between April 1998 and August 1999 ...............................................................27
7. Number of Dolphins Observed in 1 Km Zones of the San Diego and Santa Barbara
Study Areas Divided by the Number of Surveys Covering Each Zone ..........................28
8. Number of Dolphins Identified in San Diego and Santa Barbara between April 1998
and August 1999 with Sighting per Kilometer Surveyed Ratios in Specified
Categories ........................................................................................................................31
9. Cumulative Number of Dolphins Identified in San Diego and Santa Barbara between
April 1998 and August 1999 ...........................................................................................33
10. Mean School Size across Years for All Schools Sighted in San Diego
between 1984-1999 .........................................................................................................35
11. Mean Number of Dolphins Sighted per Complete Survey in San Diego
between 1984 and 1999 ...................................................................................................36
12. Mean Number of Dolphins per Complete Survey as a Function of Oceanographic
Season in San Diego between 1984 and 1999.................................................................38
13. Mean School Size as a Function of Oceanographic Season for Surveys
Conducted in San Diego between 1984-1999 .................................................................39
14. Number of Dolphins Observed per Unit Survey Effort over 1 Km Segments
of the San Diego Study Area between 1984-1999 ..........................................................41
15. Sighting Frequencies of Dolphins Identified in San Diego between 1984-1999 ............42
16. Rate of Discovery Curve Showing the Cumulative Number of Dolphins
Identified over Blocks of Five Surveys ...........................................................................45
 1

INTRODUCTION

The objective of this research was to study space-use patterns of Pacific bottlenose
dolphins (Tursiops truncatus) in two coastal areas of the Southern California Bight: San
Diego and Santa Barbara, California. Results from this study expanded on findings obtained
from long-term photo-identification studies of bottlenose dolphins along the Pacific coast,
which began in 1981 and continued through 1999 (Hansen 1990, Wells et al. 1990, Weller
1991, Caldwell 1992, Feinholz 1996, Defran and Weller 1999, Defran et al. 1999, Dudzik
1999, Marsh 2000, Defran et al. In prep). Bottlenose dolphins occurring along the west coast
of North America generally remain within 1 km of shore (Hanson and Defran 1993, Caretta
et al. 1998, Defran and Weller 1999) and are known to have a coastal range of at least 830
km, from Ensenada, Baja California, Mexico to Monterey Bay, California (Wells et al. 1990,
Defran et al. 1999). Members of this population demonstrate little site fidelity to any one
coastal area within their range (Defran and Weller 1999, Defran et al. 1999) and live in a
highly dynamic social system characterized by fluid social affiliations between dolphins
(Weller 1991, Marsh 2000). Past research on Pacific coast bottlenose dolphins has been
largely centered in San Diego, with more limited survey effort conducted off three coastal
California areas: Orange County, Santa Barbara, and Monterey Bay; and two coastal regions
along the Pacific coast off Baja California Norte, Mexico: Ensenada and San Quintín. This
study used both Santa Barbara and San Diego as primary study areas, allowing direct inter-
study area comparisons to be made. Primary objectives were to document bottlenose dolphin
site fidelity and occurrence patterns, as well as to describe the distribution and movement
patterns within and between these coastal areas. Information gathered during the study offers
a broader understanding of the ecological and biological factors affecting space-use patterns
of coastal bottlenose dolphins inhabiting the Southern California Bight.

Space-use Patterns

Information regarding the movements of animals within their environment plays an

important role in understanding the behavioral ecology of a species or population. For most
mammalian species, individuals are not distributed randomly but instead form characteristic
 2

patterns of distribution, grouping, ranging, and association (Crook et al. 1976). These
behavioral patterns are shaped not only by spatial and temporal changes in the environment
but also by internal biological constraints of individuals, such as age and sex. In the context
of ecological and internal influences, behavioral strategies evolve to optimize the conditions
under which individuals can perform vital functions to maximize fitness (Crook et al. 1976).
Vital functions include resource exploitation, predator avoidance, and reproduction.
Movement is an important and necessary component of these vital functions; without
movement it would be difficult to procure food, avoid predators, or find mates with which to
reproduce.
Traditionally, studies of animal space-use patterns have focused on documenting the
home range of individuals. Home range refers to the area an animal normally uses for
routine activities, such as feeding, mating, and care of young (Burt 1943, Jewell 1966). More
recently, studies have focused on how animals use space within their home range. Most
animals do not use all areas of their home range with the same intensity; instead, it is more
common for certain regions to be occupied with greater frequency than others (Dixon and
Chapman 1980). Areas of concentrated use are termed “core areas” (Burt 1943, Kaufmann
1962, Samuel et al. 1985). Studying core areas and movements within the range is an
important part of understanding factors determining space-use and in helping to understand
interactions with other individuals and the environment (Samuel et al. 1985).

Ecological Factors
On an ecological time scale (i.e., the lifespan of an individual), fitness is defined by
survival and reproduction. Survival necessitates extracting life-sustaining energy from
resources within the environment, while simultaneously avoiding being killed by predators.
Reproduction requires finding receptive individuals with which to mate. Since resource
availability, predation pressure, and access to mates vary across environments, they represent
ecological constraints that influence the behavioral strategies animals adopt to maximize
fitness given the specific structure of the habitat in which they live (Crook et al. 1976).
Resource availability has a fundamental ecological influence on the behavior of
animals, as an individual must first obtain sufficient food resources to satisfy metabolic
requirements before it is able to avoid predation or find mates. On the most basic level, the
home range of an animal must be large enough to provide adequate food resources to satisfy
 3

energetic requirements (McNab 1963). Abundance, distribution, and predictability of food

resources influence both the size of the area needed, and the way space within that area is
used (Davies and Houston 1984). Optimal foraging theory, which states that an individual
will maximize food intake while minimizing the amount of energy used to obtain food
(Krebs and McCleery 1984), has been used to draw several generalizations about the
interaction between resource availability and space-use patterns. Where food resources are
abundant, animals tend to use smaller ranges than in areas where resources are limited and
individuals must travel farther to obtain food (Schoener 1968, Clutton-Brock 1975).
Abundant resources dispersed into patches, however, may also result in increased range
sizes, as animals must travel between patches (Orians 1961, Schoener 1968, Geist 1974,
Jarman 1974, Clutton-Brock 1975, Hladik 1975). Resource predictability may further affect
the distribution of individuals (Brown 1964). Resources which are dispersed into patches of
consistent abundance and distribution often favor the development of territorial strategies, as
an individual can survive on one patch and will expend less energy in defending that patch
than in traveling between patches. Unpredictable resources, however, are difficult to defend
and tend to favor more wide-ranging space-use strategies.
Competitive interactions between individuals of the same species or different species
occupying the same ecological niche may further modify the abundance and spatial and
temporal distribution of resources and thereby influence space-use strategies. As more
individuals exploit an area of initially abundant resources, the resource value of the area
declines until it becomes more beneficial for an individual to move to another area (Milinski
and Parker 1991).

Space-use patterns, which maximize resource exploitation, may vary on both a

temporal and spatial scale in response to changes in resource availability. Bottlenose
dolphins off the west coast of California shifted their range northward during the 1982-1983
El Niño. Prior to 1983, the northern range boundary for Pacific coast bottlenose dolphins
was considered to be San Pedro, California (Hansen 1990). Following the El Niño event,
dolphins identified off San Diego were subsequently sighted as far north as Monterey Bay
(Wells et al. 1990); this range extension appears to be long-term, as dolphins have continued
to use the Monterey Bay area through 2001 (Fienholz 1996; Pacific Cetacean Group,
personal communication). The spatial shift in the range of Pacific coast bottlenose dolphins
 4

was thought to be in response to shifts in prey distribution due to changing water

temperatures and oceanic currents brought by the El Niño. Similar shifts in territory use have
also been documented for red foxes (Vulpes vulpes) living in Oxford, England (Doncaster
and MacDonald 1991). Red foxes live in social groups, with each group inhabiting an
exclusive territory. In neighboring suburban areas of Oxford, group territories were
primarily stable. However, within the city of Oxford red fox groups demonstrated a pattern
of “drifting territoriality”, where territories shifted in a directional, fixed spatial arrangement
across the landscape. Territory shifts were considered to be a behavioral response to the
instability of the urban environment, where food resources fluctuated rapidly and
unpredictably (Doncaster and MacDonald 1991).
Predation pressure is another environmental constraint influencing space-use patterns.
Predation has an obvious and direct correlation with fitness; there are few failures more
detrimental to fitness than being killed. Predation may affect not only where an animal
feeds, but also how it feeds. Studies of foraging behavior in gray squirrels (Sciurus
carolinensis) indicated that individuals fed preferentially in patches closest to cover
(Newman and Caraco 1987) and increased travel speed between and time spent in patches
offering little cover (Newman et al. 1988).
Access to mates, and thereby the opportunity to reproduce, acts as an additional
environmental parameter thought to influence behavioral patterns. While space-use
strategies used to satisfy metabolic demands or avoid being killed by predators may increase
chances of survival, maximizing fitness also entails passing genes on to the next generation.
Strategies to increase access to mates are often integrated with strategies to exploit food
resources. For example, in some species males may attempt to control access to valuable
food resources, as females will be drawn to areas of high resource quality. Male territorial
systems evolve in areas where resources are easily defended, such as environments with
stable, abundant, and dispersed food resources. In areas where the costs of territoriality are
high (i.e., areas of unstable and/or limited resources), males may attempt to control access to
females by preventing other males from mating with females of his group, often using
aggression (Orians 1961, Brown 1964, Emlen and Oring 1977). Space-use patterns
developed to maximize opportunities to mate may also be expressed in sex-based differences
among space-use patterns, which are discussed in the following section.
 5

Internal Constraints
Within the context of ecological variation, internal constraints reflect individual
requirements for energy and may be influenced by a range of biological factors, including
size, sex, age, and reproductive condition. Body size represents one of the most obvious
correlations with movements, as an individual of larger size requires more energy to survive.
As such, large animals may require a greater area to provide enough energy for metabolic
requirements (McNab 1963).
Space-use patterns may also be influenced by demographic factors such as age and
sex of individuals. Sex-specific differences in space-use patterns of mammals often result
because some resources may be more important to one sex than to the other. Since most
male mammals are not involved in rearing young, access to mates and increased opportunity
to reproduce are in some respects more important determinants of reproductive success than
survival of offspring. Reproductive success of females, however, is dependent on the
number of young produced and raised to maturity. Thus female space-use patterns may be
more directly correlated with access to resources necessary for producing and caring for
young, such as food, shelter, and water; while male space-use patterns may be more closely
associated with distribution of potential mates (Brown 1966, Eisenberg 1966, Trivers 1972,
Bradbury and Vehrencamp 1977, Emlen and Oring 1977).
Reproductive condition, particularly of females, may further shape movement
patterns. A female that is pregnant or nursing a calf will have increased energetic demands.
A study of coyotes (Canis latrans) in Nebraska found that females had smaller home ranges
during gestation and nursing periods than during pre-breeding and breeding seasons (Andelt
and Gipson 1979). Similar results were found in a study of roe deer (Capreolus capreolus),
where females with fawns had smaller ranges than females without fawns, presumably
because of increased energetic demands (Tufto et al. 1996).
Space-use patterns may also vary with age of individuals. In a radio-tracking study of
opossums (Didelphis marsupialis) in Kansas, immature opossums covered a smaller area on
a daily basis than adult opossums did (Fitch and Shirer 1970). Immature opossums,
however, did occasionally make dispersive movements, presumably to explore their
environment, and progressively extended their range of movements with age. These range
extensions suggested that young animals simply needed time to discover the limitations of
 6

their environment. A similar trend was found in white-crowned sparrows (Zonotrichia

leucophrys nutalli) where first year male sparrows had significantly smaller territories than
mature males (Ralph and Pearson 1971). Differences in territory sizes were correlated with
breeding success, however, and suggested that younger sparrows lacked the experience to
defend more optimal territories.

Pacific Coast Bottlenose Dolphin Research

Boat-based photo-identification studies of Pacific coast bottlenose dolphins were

started by Hansen (1990) in 1981 and continued through 1999. While Hansen’s research was
concentrated in the San Diego area, photo-identification studies have since incorporated
several other areas of the Pacific coast, ranging as far south as San Quintín, Baja California
Norte, Mexico (30°15’N, 121°48’W) north to Monterey Bay, California (36°48’N,
121°48’W) (Wells et al. 1990, Weller 1991, Caldwell 1992, Feinholz 1996, Defran and
Weller 1999, Defran et al. 1999, Dudzik 1999, Marsh 2000, Defran et al. In prep). Studies of
Pacific coast bottlenose dolphins have spanned almost two decades, and the longitudinal
nature of the research has provided a relatively detailed understanding of the interaction
between the highly dynamic environment found along the Pacific coast and the behavior of
bottlenose dolphins inhabiting the area.
Hansen’s (1990) study included 22 boat-based photo-identification surveys conducted
between September 1981 and January 1983. Surveys covered the area extending from
Scripps Pier in La Jolla, California north to Oceanside Harbor (See Figure 2). Dolphins were
sighted on 86% (n = 12) of all surveys encompassing the entire study area, with most
sightings occurring between Torrey Pines State Beach and south Carlsbad. All sightings
were made within 1 km of shore. One hundred twenty-three dolphins were photographically
identified, with 42% (n = 52) sighted only one time and the remainder sighted between two
and nine times during the study. A small subset of identified individuals (n = 21) were
sighted five or more times throughout the study. Closed population models estimated that
between 173 – 240 dolphins used the area. Seven aerial surveys, covering all (n = 5) or part
(n = 2) of the area between the Mexican border and Long Beach Harbor, were also conducted
as part of Hansen’s (1990) study. Dolphins were encountered on all aerial surveys, with the
number of dolphins counted per survey ranging from 55 to 128.
 7

Based on the results from his study, Hansen (1990) considered the range of Pacific
coast bottlenose dolphins to include an area of at least 155 km between La Jolla and San
Pedro. While dolphins were present in the San Diego study area year-round, sighting
frequencies of individuals varied considerably, and Hansen concluded that the San Diego
area was part of a seasonal range for some individuals and a permanent range for the small
subset of the population (n = 21) sighted most often.
Defran and colleagues (Defran and Weller 1999, Defran et al. 1999) continued
research on Pacific coast bottlenose dolphins between 1984 and 1989. One hundred forty-six
boat-based photo-identification surveys were conducted in San Diego between January 1984
and December 1989. The study area used was slightly smaller than Hansen’s study area, and
extended 32 km from Scripps Pier in La Jolla to south Carlsbad. Dolphins were sighted on
72% (n = 79) of all surveys covering the entire area, but encounter percentages varied
annually, ranging from a low of 60% in 1987 to a high of 95% in 1989. Both number of
dolphins encountered per survey (mean = 26.8, SD = 22.30) and school size (mean = 19.8,
SD = 18.40) were also variable. While sightings were made within 1 km of the coastline and
were distributed throughout the study area, the majority (70% of dolphins, 67% of schools)
of sightings were concentrated in the southern portion of the study area, between Torrey
Pines State Park and Solana Beach.
Three hundred seventy-three individual dolphins were identified between 1984 and
1989. New dolphins were continually being identified throughout the study, although the
rate at which previously unidentified dolphins were photographed had begun to decrease by
1989. Identified individuals were sighted between one and twenty-four times, with an
average of 4.6 sightings. The majority (66%) of identified dolphins were resighted six times
or less, for an average of less than one time per year. Sighting per opportunity ratios were
calculated to reflect sightings per unit effort, and were derived by dividing the number of
sightings for each individual by the number of surveys on which dolphins were encountered.
The overall mean sighting per opportunity ratio was 0.09.
Association patterns of identified dolphins were analyzed by Weller (1991) using the
half-weight index, which yields values ranging from 0.00 for animals never sighted together,
to 1.00 for animals always sighted together. Ninety-five percent of the calculated
coefficients of association were between 0.00 and 0.39, and mean levels of association for
 8

individual dolphins ranged from 0.14 to 0.29. The mean number of affiliates ranged from 1-
24, with the number of affiliates increasing with the number of sightings of each individual.
The low coefficients of association and high number of affiliates indicated that Pacific coast
bottlenose dolphins are members of a fluid and dynamic social system.
Twenty-nine percent (n = 30) of the dolphins sighted by Hansen were not
photographed by Defran and Weller (1999) between 1984 and 1989. Of Hansen’s subset of
frequently (>5 times) sighted dolphins (n = 21), four were not photographed during Defran
and Weller’s study, and six were seen less than once per year. One of Hansen’s frequently
sighted dolphins, however, was the most frequently sighted dolphin between 1984 and 1989.
Photographic survey efforts in San Diego ceased in 1989 but were resumed by
Dudzik (1999) between March 1996 and August 1998. She conducted 66 photographic
surveys, during which 233 dolphins were identified. Fifty-six percent (n = 131) of these
dolphins had been previously identified during the 1984-1989 study by Defran and Weller
(1999). Comparison of occurrence patterns, as measured by the percentage of surveys on
which dolphins were encountered, the number of dolphins per complete survey, and average
school size, between the 1984-1989 surveys and the 1996-1998 surveys showed no
significant differences, suggesting that occurrence of dolphins was stable over time.
Population size estimates were also compared across study periods, using Chao’s Mth model.
Given the low sighting frequencies of most identified dolphins in the population, this model
was considered the most appropriate because it allowed capture probabilities to vary by both
time and individual. Estimates were derived for three different study periods (1984-1986,
1987-1989, 1996-1998) and ranged from 289 to 356. Similar estimates across study periods
suggested that population size also remained stable across the eleven year time span.
Regional studies have also been carried out in other areas of the range of Pacific coast
bottlenose dolphins. Between 1981 and 1989, boat-based photo-identification surveys were
conducted in three secondary study areas within the Southern California Bight: Orange
County and Santa Barbara, California; and Ensenada, Baja California Norte, Mexico (Defran
et al. 1999). The majority of dolphins identified in Santa Barbara (88%), Orange County
(92%), and Ensenada (88%) were also photographed in San Diego. One hundred twenty
(58%) of the 207 dolphins identified in the three secondary study areas were documented to
 9

move between areas. Within each study area, most individuals were sighted only one time,
and the majority of resightings were made within days or weeks.
During the 1982–1983 El Niño, Pacific coast bottlenose dolphins were documented to
shift the northward limit of their range (Wells et al. 1990, Feinholz 1996). Bottlenose
dolphins were first sighted north of Point Conception in May 1983 (Wells et al. 1990).
Opportunistic sightings indicated bottlenose dolphins traveled as far north as Monterey Bay
between 1983 and 1988, and several of these sightings included dolphins previously
identified within the Southern California Bight (Wells et al. 1990).
Systematic photo-identification surveys of Monterey Bay began in 1990. Feinholz
(1996) conducted 84 boat surveys between October 1990 and November 1993. Bottlenose
dolphins were sighted on 79% (n = 66) of all surveys. Sixty-eight individual dolphins were
identified; 63% (n = 43) of the dolphins identified had previously been sighted within the
Southern California Bight. Two dolphins sighted within Monterey Bay had previously been
sighted as far south as Ensenada, a distance approximately 830 km south of Monterey Bay.
Twenty-six percent (n = 18) of the dolphins identified were sighted only one time. Feinholz
(1996) considered 13 individuals (19% of identified dolphins) to demonstrate some level of
site fidelity to the Monterey Bay area, based on sighting frequencies and presence in the area
at least once during each year of the study.
Eight boat-based photo-identification surveys were conducted in San Quintín, Baja
California Norte, Mexico between April and August 1990 (Caldwell 1992, Defran et al. In
prep) to determine a southern range boundary for Pacific coast bottlenose dolphins. One
hundred five dolphins were identified. The majority (62%) of dolphins identified were
sighted only one time, with most resightings occurring within days of the first sighting. Only
one dolphin had previously been sighted within the Southern California Bight. Dolphin #006
was sighted 5 times in San Diego and twice in Ensenada prior to being photographed on two
consecutive days in San Quintín. Subsequent to its sighting in San Quintín, dolphin #006
was also photographed several times in San Diego. While survey effort in San Quintín was
relatively limited, comparable effort in Santa Barbara and Ensenada was sufficient to detect
movements of known individuals between areas. Thus, results from the San Quintín study
indicated that a probable southern range boundary for Pacific coast bottlenose dolphins exists
between Ensenada and San Quintín. The boundary was unlikely to represent physical
 10

limitations of distance dolphins were able to cover, since Santa Barbara and San Quintín are
roughly the same distance from San Diego. The exact nature of the boundary, and any
causal mechanisms, have yet to be determined (Caldwell 1992, Defran et al. In prep).
Results from studies spanning many years and several different study areas have
illustrated the complexity of interactions between the dynamic environment found along the
Pacific coast and bottlenose dolphin behavior. Coastal California waters experience
significant daily, monthly, and yearly variability, creating patchy and unpredictable patterns
of resource availability (Cross and Allen 1993, Dailey et al. 1993). The high mobility,
extensive coastal distances traveled, variable school sizes, and apparent lack of site fidelity
demonstrated by Pacific coast bottlenose dolphins may be a reflection of patchy and
unpredictable abundance and distribution of prey fish along the Pacific coast (Defran and
Weller 1999, Defran et al. 1999). Changes in prey distribution and abundance are also
presumed to have affected range boundaries, demonstrated both by the northward range
extension of Pacific coast bottlenose dolphins following the 1982-1983 El Niño (Wells et al.
1990), and by the existence of a probable southern range boundary north of San Quintín
(Caldwell 1992, Defran et al. In prep). The high degree of behavioral variation over both
time and space that has been demonstrated by Pacific coast bottlenose dolphins would have
been difficult to document in a study of more limited scope, and emphasizes the importance
of conducting studies covering broad temporal and geographic scales.

Objectives

While research on Pacific coast bottlenose dolphins was conducted for almost two
decades prior to this study, important questions remained unanswered. The extension of
research efforts between 1998 and 1999 offered further opportunity to examine the stability
of behavioral patterns over time, allowing more detailed information on the ecological and
biological factors influencing individual behavior to be obtained. Research presented here
involved the continuation of boat-based photo-identification studies of Pacific coast
bottlenose dolphins in San Diego, California, as well as the inclusion of an additional study
area in Santa Barbara, California, 271 km to the north of San Diego and near the northern
terminus of the Southern California Bight. While Pacific coast bottlenose dolphins had been
studied in areas other than San Diego before, such studies had never been conducted over the
 11

same time span and with comparable effort. The addition of regular survey effort in Santa
Barbara in combination with continued survey effort in San Diego allowed movements of
individuals, occurrence patterns, and individual sighting frequencies to be compared between
areas, and permitted more detailed examination of the effect that ecological variables may
have on space-use patterns of Pacific coast bottlenose dolphins. Further study in both areas
also allowed determination of sex and reproductive state of more individuals. Comparison of
behavioral patterns of individuals of known sex and reproductive state to the rest of the
identified population permitted preliminary analyses of how internal differences factored into
behavioral variation between individuals. Furthermore, comparison of results from this study
to results from prior datasets allowed documentation of any changes in occurrence patterns or
individual sighting frequencies of Pacific coast bottlenose dolphins that may have occurred
over time in response to ecological variation.
The strength of the study was derived from the many comparisons that could be
made. As such, the objectives of the study were classified into three categories: (1) within
study area comparisons, (2) between study area comparisons, and (3) past-to-present
comparisons. Within each study area, documentation of the occurrence of bottlenose
dolphins throughout the year-long study allowed comparison of space-use patterns over
seasons. Comparisons of the results from surveys in San Diego and Santa Barbara permitted
documentation of movements between areas and analysis of space-use patterns of each area
by individuals. The distribution of sightings within each area was also compared to
determine if regions of heavy use within study areas shared similar ecological features.
Patterns of occurrence, distribution, and individual sighting frequencies between 1998 and
1999 were further compared to past research conducted on Pacific coast bottlenose dolphins
to determine the stability of usage patterns over time.
 12

METHODS

The research presented here utilized two coastal areas within the Southern California
Bight: San Diego and Santa Barbara, California. The Southern California Bight (Figure 1)
extends 732 km from Point Conception (34º33’N, 120º28’W) in the north to Punta Colnett
(30º57’N, 116º20’W) in the south. The coastal environment within the Southern California
Bight can be characterized as open and dynamic. Point Conception marks a sharp eastward
break in the coastline which interrupts the flow of the California current system, resulting in
the creation of the Southern California Countercurrent and producing a region where
northern, southern, western, and upwelling bottom waters converge. Converging waters
create a marked change in coastal climate and marine fauna within the Bight. Waters of the
Southern California Bight are subject to both short- and long-term temperature fluctuations
dependent on oceanic currents, with average maximum surface temperatures peaking at 19°C
between July and September, and falling to an average minimum temperature of 14.5°C in
late winter. Nearshore salinities vary slightly, with a high of 33.6 ppt in July and a low of
33.4 ppt in late winter (Dailey et al. 1993).
The San Diego study area (Figure 2) consisted of a narrow strip of coastline
extending from Scripps Pier in La Jolla (32º52’N, 117º15’W) 32 km north to Tamarack in
south Carlsbad (32º08’N, 117º20’W). Nearshore underwater topography was variable,
ranging from submerged reefs, dense kelp beds, and sea grass flats to barren sandy bottoms.
Beaches also varied in composition, with gently sloping sand, steeply inclined cobblestone,
estuary mouths, and rocky outcrops. Low levels of commercial and recreational vessel
traffic were present. The study area was similar to the area used by Hansen (1990), and
identical to that used by Defran and Weller (1999) and Dudzik (1999).
The Santa Barbara study area (Figure 3) extended from the Santa Barbara Harbor (34º
25’N, 119º 42’W) in the north 38 km southeast to Emma Wood State Beach (34º18’N,
119º21’W) in Ventura County. While Santa Barbara and San Diego were similar in
nearshore underwater topography and beach composition, the Santa Barbara coastline is
oriented northwest to southeast, as opposed to the more direct north-south orientation of the
San Diego coastline. Much of the Santa Barbara area is also sheltered by the Channel Islands
 13

Point Conception
Santa Barbara

34

Orange County

33
San Diego

Pacific
Ocean
N
32 Ensenada

50 km

Punta
31 Colnett

120 119 118 117

Figure 1. Map of the Southern California Bight.

 14

117o 31' 117o 23' 117o 15'

33o14 '

33o12 '
O c ea n s id e

33o10 '
P a ci fi c C arl s b ad
O ce an
33o08 '
N

33o06 '
4 km

Leu ca d i a
33o04 '

E n c in i tas
33o02 '
Ca r d if f

33o00 ' S o la n a B e a ch

32o58 '
Del M ar
C
al

32o56 '
ifo
rn
ia

To rre y P i n es
S tat e P ark
32o54 '

32o52 ' S a n D ie g o S cri p p s P ie r

S t u d y A re a M e xic o

32o50 ' La J o l la

Figure
Figure 2 2.
Figure San
.2.San Diego
Diego
San study
Study
Diego Area.area
study area
 15

119o40' 119o30'

Goleta Santa Montecito

Barbara Summerland

Carpinteria
Santa Barbara
Harbor
34o20'

Pacific Ventura
Ocean
N Ventura
Harbor

7 km Oxnard

Anacapa
Santa Cruz
34o00'

Figure 3. Santa Barbara study area.

 16

(San Miguel, Santa Rosa, and Santa Cruz islands), which provided relief from wind and sea
conditions and resulted in generally calmer waters than in the San Diego study area.

Photographic Survey Procedure

Photo-identification surveys were conducted twice a month in San Diego and four
times a month in Santa Barbara. San Diego surveys used a 5.8 m powerboat equipped with a
115 hp engine, while Santa Barbara surveys utilized a 4.6 m inflatable boat with a 40 hp
engine. All surveys were conducted in Beaufort sea state of ≤3, to ensure conditions
adequate for sighting, observing, and photographing dolphins.
Boat surveys consisted of travel 90-180 m outside of the surf line while systematic
observations from the beach to 2 km offshore were conducted. When a school of dolphins
was detected, time and location (using Global Positioning System) where the school was
initially sighted were recorded. Attempts were then made to photograph all dolphins present
in the school. After all dolphins in the school were thought to have been photographed,
location and time where observations were terminated were recorded, and school
composition (i.e., number of calves present) and school size were noted. The survey then
proceeded until the next sighting was made or the end of the study area was reached. All
surveys attempted to cover the entire study area. Surveys covering the entire study area were
labeled as complete surveys and were used in all analyses. Surveys that failed to cover the
entire study area due to weather, sea state, or equipment failure were labeled as partial
surveys and were used in selected analyses.
Definitions of schools, calves, and probable mothers followed those used by Weller
(1991), Defran and Weller (1999), Defran et al. (1999), and Dudzik (1999) so that data could
be compared. Schools were defined as any dolphins observed in close proximity to one
another and usually moving in the same direction and engaged in similar behavior. Calves
were defined using three criteria: (1) an animal consistently demonstrating close affiliation
with a larger dolphin; (2) an animal displaying an awkward and immature surfacing pattern;
and (3) an animal small in size, possessing fetal folds, and/or distinct neonatal coloration.
Probable mothers were defined as in Weller (1991): a larger dolphin exhibiting prolonged
and exclusive or near-exclusive association with a dolphin identified as a calf. In addition,
those dolphins defined as probable mothers (n = 12) by Weller (1991) between 1984 and
 17

1989 were combined with those dolphins defined as probable mothers during the present
study to represent a subset of females in the population.

Photo-identification Procedures

A Canon A-1 35 mm camera equipped with a 400-mm lens and high-speed motor
drive was used to photograph dolphin dorsal fins. Kodak Tri-X 400 ISO black and white
film was used for all photographs. Methods used to sort, match, identify, and catalog dorsal
fin photographs are described in detail elsewhere (Defran et al. 1990) but are briefly
summarized as follows.
After developing film from each survey, the resulting negatives were assessed for
photographic quality and notch pattern distinctiveness. Photographic quality was evaluated
on the basis of focus, clarity, lighting, and parallax of the image; any negative not meeting
the pre-set criteria was excluded from analyses. Only dorsal fins containing a notch pattern
allowing unequivocal identification from other fins in the database were used. Once usable
negatives were sorted out, negatives from each school were grouped according to
recognizable individuals. The best negative (based on photographic criteria presented in
Appendix) of each dorsal fin was considered the type specimen and was placed in a slide
mount labeled with date, school number, area of sighting, and a temporary identification
number. Type specimens were projected and enlarged to fit a 10x17 cm frame and
subsequently traced by hand onto white paper. The tracing process allowed all traced fins to
be standardized for size and orientation. After tracing, dorsal ratios were calculated for all
dorsal fins with two or more notches on the trailing edge. Dorsal ratios were computed by
dividing the distance between the two largest notches on the trailing edge of the fin by the
distance between the top of the fin and the notch nearest the bottom of the fin (Figure 4). If
the fin contained notches of similar size, the notches furthest apart were used for the dorsal
ratio.
Tracings of previously identified dolphins were placed into one of four catalogs: (1)
dorsal fins with a single notch; (2) dorsal fins with a notch in the top of the fin; (3) dorsal fins
with two to three notches; and (4) dorsal fins with four or more notches. Within catalogs,
tracings were organized numerically according to dorsal ratios. Organization of identified
individuals by number of notches and dorsal ratios facilitated the cataloging process. The
 18

TOP Dorsal
Ratio =
A to B
B to Top

A 19 mm
44 mm
= .439

Computation of the dorsal ratio (from Defran et al. 1990).

Figure 4. Computation of the dorsal ratio
 19

analyst first searched for matches of the unidentified dolphins under the appropriate catalog
and dorsal ratio. Upon finding a match, slides of the new dolphin and all slides of the
previously identified dolphin were compared. If all slides matched, the image was then
labeled and filed with the identification number of the previously identified dolphin. If no
matches were made, all catalogs were searched twice before the dolphin was considered a
new sighting and given a permanent identification number. For all steps of the cataloging
process, three people must either have agreed to the match or have searched all catalogs
twice before the process was considered complete.
All negatives were given a photographic quality rating based on four factors (see
Appendix): focus of fin image, contrast between fin image and background, proportion of fin
visible in image, and size of fin image. Negatives given the lowest rating on any one of these
factors were not used for any analysis.

Computer Analysis

Data collected in San Diego and Santa Barbara between April 1998 and August 1999
were stored in two Excel databases: a photographic database consisting of date and area of
sightings of all individuals; and a sighting database, in which the date, time, and location of
sightings, as well as estimated school size and number of calves present, were recorded.
These databases are referred to as the short-term photographic and sighting databases.
Within the short-term databases, data collected in San Diego by Dudzik (1999) between
April 1998 and August 1998 were consolidated with data collected in San Diego for this
study
Data collected in San Diego between 1998 and 1999 were then combined with data
collected in San Diego between January 1984 and December 1989 (Defran and Weller 1999)
and between March 1996 and March 1998 (Dudzik 1999) to form a long-term dataset
consisting of all bottlenose dolphin sightings and identifications in San Diego between 1984
and 1999. This dataset was also divided into a photographic database, containing the dates of
sighting for all dolphins, and a sighting database with the date, time, location, and
composition of all schools. These databases are referred to as the long-term photographic
and sighting databases, respectively.
 20

The sighting databases (both long and short-term) were used in analysis of occurrence
patterns and distribution of sightings. The photographic databases were used to examine site
fidelity and movement patterns, as well as to construct rate of discovery curves.

Occurrence Patterns
Occurrence patterns were measured in several ways: total number of dolphins
observed in the study area per survey, percentage of surveys encountering dolphins, and size
of schools encountered. The first three parameters were analyzed using data from the survey
database. Only complete surveys were used to analyze the number of dolphins observed per
survey and the number of surveys encountering dolphins, while all surveys were used for
school size analyses.
Occurrence pattern parameters were analyzed using analysis of variance methods
(ANOVAs) generated by SPSS 7.5. Although the Santa Barbara study area is larger than the
San Diego study area, the difference is slight (~ 6 km), and occurrence patterns were
compared directly between the two areas. Within each study area, occurrence patterns were
also compared between oceanographic seasons, which were defined by Hickey et al. (1993)
to include the Upwelling period (March -July), Oceanic period (August-October), and
Davidson current period (November-February). The oceanographic seasons are defined by
water temperatures and as such may more directly influence dolphin and prey distribution
than would traditional solar seasons.
The final comparisons of occurrence patterns were made by combining data collected
in San Diego between 1998 and 1999 with data collected in San Diego between 1984 and
1989 (Defran and Weller 1999) and between 1996 and 1998 (Dudzik 1999). Occurrence
patterns were then compared across years, as well as across oceanographic seasons, to
determine the stability of occurrence patterns over time. The long-term survey database was
also used to compare occurrence patterns across El Niño, La Niña, and normal months.
Months were classified into categories using the Bivariate Enso time series index (Smith and
Sardeshmukh 2000), which is based on both sea surface temperature anomalies and
atmospheric pressure differences.
 21

Distribution
Distribution of sightings within each study area was examined by dividing each area
into zones of 1 km² and calculating a ratio of the number of dolphins observed in each zone
divided by the number of surveys covering that zone. Location where dolphins were first
sighted was used in order to avoid potential influences on dolphin behavior by boat
interaction. Zones were then classified as follows: (1) low if the usage ratio was below the
mean minus the standard deviation (for both areas this was a negative number, so zones with
no usage were classified as low), (2) high if the usage ratio was above the mean plus the
standard deviation, and (3) moderate for all other ratios. Although this analysis is recognized
as being preliminary, ratios were employed to provide a rough comparison of usage between
zones. Within San Diego, ratios were calculated using both 1998-1999 sightings and 1984-
1999 sightings to examine consistency of patterns over time.

Site Fidelity
For the 1998-1999 study, the degree of site fidelity to each area was examined. This
analysis calculated a ratio of sightings per kilometer surveyed for all identified individuals in
each area as a way to account for unequal survey effort between areas. The number of
kilometers covered in each survey was determined by plotting the GPS location where
surveys were terminated onto the 1 km zones used for distribution analyses. The last 1 km
zone completely covered by the survey was considered the number of kilometers covered.
Kilometers covered in surveys of each area were then summed, and the number of sightings
of identified dolphins in each area was divided by the corresponding sum of kilometers
covered for each area. An ANOVA was then used to compare sighting per kilometer ratios
between areas to determine if mean levels of site fidelity differed. As previously mentioned,
since the majority of consecutive day surveys occurred in Santa Barbara, sightings on
consecutive days were excluded from the analysis (and effort calculations were adjusted) to
eliminate any potential bias introduced if dolphins were more likely to be resighted on
consecutive day surveys rather than surveys separated by a week or more.
A preliminary analysis of biological constraints contributing to individual variation
in site fidelity characteristics was also conducted, by subdividing the population by sex and
lactational state. Dolphins identified as probable mothers (i.e., lactating dolphins) between
1998 and 1999 represented one subset, and their sightings per kilometer ratios in each area
 22

were compared to those of the rest of the identified population. Probable mothers identified
between 1998 and 1999 were combined with dolphins identified by Weller (1991) as
probable mothers between 1984 and 1989 to represent a subset of females. The sightings per
kilometer ratios of all females were then compared with the ratios of the remainder of the
population. These analyses were considered preliminary, as they represent only one subset
of each factor (i.e., only females or only lactating dolphins) and are made up of a small
fraction of those subsets. However, these analyses may provide a basis for generating ideas
for future study.

Movements
The movements of identified dolphins between San Diego and Santa Barbara between
1998 and 1999 were documented. The mean, range, and standard deviation of interval of
days between sightings in different areas were analyzed by calculating the maximum travel
speed between areas. The number of dolphins sighted in both areas over the long-term study
period between 1984 and 1999 was also calculated.

Rate of Discovery Curves

Rate of discovery curves, which plot the number of dolphins identified in an area over
time or effort, were generated for both the short- and long-term photographic datasets. For
the short-term study, separate rate of discovery curves were generated for the San Diego and
Santa Barbara study areas by plotting the cumulative number of dolphins identified over each
month of the study. Curves were then compared between the two areas in order to determine
if similar rates of acquiring previously unidentified dolphins existed in each area. The slope
of each curve was also examined to provide further information on occurrence patterns.
While a continuously rising slope would indicate that previously unidentified dolphins are
moving into an area at a steady rate, an irregular slope, with periods of rapid increase
interspersed with periods of little or no increase, would be indicative of pulses of new
dolphins moving into the area over certain months or seasons.
A collective rate of discovery curve for the San Diego study area was also generated
using dolphins identified in San Diego between 1984 and 1999. This curve was plotted using
the cumulative number of dolphins identified over blocks of five surveys on which at least
one dolphin was identified. The curve was then examined to determine the point at which it
 23

began to reach an asymptote, which was used as an indication that the majority of dolphins in
the population had been identified.
 24

RESULTS

Between April 1998 and August 1999, 43 surveys were conducted in San Diego coastal
waters, and 61 surveys were conducted in Santa Barbara waters. Of these surveys, 65.1% (n
= 28) in San Diego and 70.5% (n = 43) in Santa Barbara covered the entire study area and are
termed “complete” surveys.

Occurrence

A total of 912 dolphins in 58 schools were observed in San Diego, while 915 dolphins
in 72 schools were observed in Santa Barbara. Calves represented 8.3% (n = 76) of all
dolphins observed in San Diego and 7.8% (n = 71) of all dolphins observed in Santa Barbara.
In both study areas between one and four schools were observed per survey, with
42.9% (n = 12) of complete surveys in San Diego and 27.9% (n = 12) of complete surveys in
Santa Barbara encountering only one school. Both complete and partial surveys were used to
compare school sizes between areas. Mean school size was slightly lower in Santa Barbara
(mean = 12.7, SD = 8.27, range 2-35) than in San Diego (mean = 15.7, SD = 8.26, range 1-
96), but these differences were not significant (F(1, 128) = 1.49, P>0.05).
The number of dolphins sighted during complete surveys in San Diego ranged from 1
to 135 (mean = 20.6, SD = 28.50), and dolphins were encountered on 82.1% (n = 23) of
complete surveys. In Santa Barbara, 67.4% (n = 29) of complete surveys encountered
dolphins, with the number of dolphins sighted per survey ranging from 2 to 54 (mean = 15.7,
SD = 15.02). No significant differences were observed in the number of dolphins observed
per survey (F(1, 69) = 0.89, P>0.05) or the percentage of surveys encountering dolphins ( ²
= 1.84, df = 1, P>0.05).
Seasonal patterns of occurrence were examined using the oceanographic seasons
defined by Hickey (1993), which include: Upwelling (March through July), Oceanic (August
through October), and Davidson (November through February). Using only complete
surveys of each study area, no significant difference across oceanographic seasons was
observed in the number of dolphins per survey in Santa Barbara (F(2,40) = 0.41, P>0.05)
(Figure 5). In San Diego, however, the number of dolphins observed per complete survey
 25

was significantly greater (F(2,25) = 10.712, P<0.001) during the Davidson period than in
either the Oceanic or Upwelling period (Figure 5). The number of complete surveys on
which dolphins were encountered did not show significant variation across oceanographic
seasons in Santa Barbara (X² = 0.35, df = 2, P>0.05) but was significantly higher during the
Upwelling season than during the Oceanic season in San Diego (X² = 6.26, df = 2, P =
0.044).
Both complete and partial survey data were combined to compare school size (Figure
6). While school size did not differ across oceanographic seasons in Santa Barbara (F(2, 69)
= 0.17, P>0.05), school sizes in San Diego were significantly larger during the Davidson
period than during either the Oceanic or Upwelling periods (F(2, 55) = 5.48, P = 0.007).

Distribution

Ratios of total dolphins observed per unit effort were graphed for 1 km² zones in each
study area (Figure 7). In San Diego, the mean number of dolphins observed in each zone
per unit survey effort was 0.7 (SD = 0.95, range 0-4.3), while in Santa Barbara the mean
number of dolphins per unit survey effort was 0.5 (SD = 0.41, range 0-1.4). Zones of low
usage in San Diego included zones 5, 11, 13, 17, 20, and 24; while zones of high usage
included zones 1, 2, and 6. In Santa Barbara, zones of low usage included zones 10-12, 14,
20, 30, and 36-38; and zones of high usage included 1, 4, 5, 17, 28, and 31. Table 1 details
some of the zones containing topographical or environmental features that might affect
dolphin distribution.

Site Fidelity

Between April 1998 and August 1999, 204 dolphins were identified in San Diego,
and 178 dolphins were identified in Santa Barbara. In San Diego, dolphins were sighted
between one and eight times, and the mean number of sightings per dolphin was 2.0 (SD =
1.49). One hundred five dolphins (51.5%) were sighted only one time. In Santa Barbara,
dolphins were identified between one and seven times, and the mean number of sightings
was 2.1 (SD = 1.30). Eighty-two dolphins (46.1%) were sighted only one time. The mean
interval between sightings of the same individual was 97.8 days (SD = 95.11) in San Diego
 26

100
90
San Diego
Mean Number of Dolphins
80
Santa Barbara
70
60
50
40
30
20
10
0
Upwelling Oceanic Davidson
Oceanographic Season

Figure 5. Mean number of dolphins observed per complete survey as a function of

oceanographic season in San Diego and Santa Barbara between April 1998 and August 1999.
Error bars represent standard deviations.
 27

45
40 San Diego
35 Santa Barbara
Mean School Size

30
25
20
15
10
5
0
Upwelling Oceanic Davidson
Oceanographic Season

Figure 6. Mean school size as a function of oceanographic season for San Diego and Santa
Barbara between April 1998 and August 1999. Error bars represent standard deviations.
 28

4.5

Dolphins Per Unit Survey

4
3.5
3
Effort
2.5
2
1.5
1
0.5
0
1
3
5
7
9
11
13
15
17
19
21
23
25
27
29
31
San Diego 1 km Zones

1.6
Dolphins Per Unit Survey

1.4
1.2
1
Effort

0.8
0.6
0.4
0.2
0
1

10

13

16

19

22

25

28

31

34

37
Santa Barbara 1 km Zones

Figure 7. Number of dolphins observed in 1 km zones of the San Diego and Santa Barbara
study areas divided by the number of surveys covering each zone. Zones run from south to
north in San Diego and from southeast to northwest in Santa Barbara.
 29

Table 1. Habitat features of study area zones in San Diego and Santa Barbara.

Study Area Zone Habitat Feature

San Diego 1-2 Scripps Canyon
5-6 Rocky reef (Ward 1999)
8-9 Los Penasquitos lagoon
13-14 San Dieguito river mouth
18-19 San Elijo lagoon
26-27 Batiquitos lagoon
Santa Barbara 1 ~ 1 km northwest of the Ventura Estuary
9-11 Oil piers
15-16 Rincon Creek mouth
21-23 El Estero lagoon
38 <1 km southeast of the Santa Barbara Harbor
 30

and 96.5 days (SD = 88.27) in Santa Barbara, and was not significantly different between
areas (F(1,397) = .02, P>0.05).
To compare sighting frequencies across areas with unequal survey effort, a ratio of
number of sightings per km surveyed was calculated for individual dolphins identified in
each area (Figure 8). Sighting per km ratios were significantly higher for dolphins identified
in San Diego than for those identified in Santa Barbara (F(1,380) = 59.77, P<0.001).
The greatest number of dolphins identified in any one month of the study occurred in
January 1999 in the San Diego study area, when 79 dolphins (38.7% of all dolphins
identified in San Diego) were identified. Forty percent (n = 164) of all identifications made
in San Diego accrued between the months of November 1998 and January 1999 (no surveys
were conducted in December in San Diego). In Santa Barbara, the highest number of
dolphins identified during any one month occurred in June 1999 and included 40 dolphins
(23% of the total number of dolphins identified in Santa Barbara).
Analyses of the effect of sex and lactational state on sighting frequencies were also
run for each area. No significant differences were found in sighting ratios for probable
mothers (F(1, 202) = 0.20, P>0.05) or females (F(1, 202) = 2.99, P>0.05) and the remainder
of identified dolphins within the San Diego study area between 1998-99. In Santa Barbara,
females (F(1, 176) = 12.80, P<0.001) and probable mothers (F(1,176) = 10.42, P = 0.001)
showed significantly higher sightings per kilometer surveyed than did other dolphins in the
population. Due to small sample sizes in the number of known females and probable
mothers, these results should be considered preliminary.

Movements

A total of 293 dolphins were identified during the study, with 30.4% (n = 89) of
identified dolphins sighted in both study areas between April 1998 and August 1999. One
hundred thirty-one inter-study area movements were documented, with individual dolphins
moving between study areas between one and four times. Fifty-five (61.8%) of the dolphins
sighted in both areas were documented to move between study areas only one time. Sixty-
nine (52.7%) of the observed movements were from San Diego north to Santa Barbara. Of
those dolphins sighted in only one area, 115 were identified in San Diego and 89 in Santa
Barbara.
 31

120
51%
San Diego
100
Santa Barbara
Number of Dolphins

44%
80

60 24%
22%
40
13% 11%
11%
20 6%
4% 5% 2%
1% 1% 2%
0
0.00- 0.50- 1.00- 1.50- 2.00- 2.50- 3.00- 3.50- 4.00- 4.50- 5.00- 5.50- 6.00- 6.50- 7.00-
0.49 0.99 1.49 1.99 2.49 2.99 3.49 3.99 4.49 4.99 5.49 5.99 6.49 6.99 7.49

Sightings Per Km Surveyed (x10-3)

Figure 8. Number of dolphins identified in San Diego and Santa Barbara between April
1998 and August 1999 with sighting per kilometer surveyed ratios in specified categories.
Numbers above bars represent percent of the total number of dolphins identified in each area
with ratios in each category. Consecutive day sightings of the same individual have been
removed.
 32

The mean number of days between sightings in different areas was 96.0 days (SD =
73.04) and ranged from five to 383 days. The fastest travel speed recorded averaged 61.6
km/day and was displayed by one dolphin first identified in San Diego on 28 February 1999
and subsequently resighted in Santa Barbara on 5 March 1999.

Rate of Discovery

Rate of discovery curves were generated for both areas using the cumulative number
of dolphins identified per month (Figure 9). By the last month of the study, only two new
dolphins (15% of all dolphins identified that month) were identified in San Diego, and one
new dolphin (8% of all dolphins identified that month) was identified in Santa Barbara.
However, the rate at which new dolphins were identified each month fluctuated considerably,
and survey effort differed between months as well as between study areas. The curves
showing the cumulative number of dolphins identified in each area shows a slight leveling
trend in both areas by the end of the study.

Comparisons with Past Studies

Sighting history and survey data collected in San Diego between April 1998 and
August 1999 were pooled with data collected in San Diego between January 1984 and
December 1989 (Defran and Weller 1999) and between March 1996 and April 1998 (Dudzik
1999) to allow comparisons to be made over time. Information on survey effort for each
study is presented in Table 2.

Occurrence patterns
Occurrence patterns, as measured by encounter rate, total number of dolphins
encountered, school size, and percentage of calves per school, were compared across years.
All observed schools were used to compare school size, which yielded no significant
difference across years (F(9, 256) = 1.641, P>0.05) (Figure 10). Both the encounter rate (X²
= 23.181, df = 9, P = 0.006) and the number of dolphins observed per survey (F(9, 255) =
4.300, P<0.001) were significantly different across years (Figure 11). With the exception of
1997, which contained both the lowest encounter rate and the lowest number of dolphins
 33

250
Cumulative Number of Dolphins Identified

SB
200
SD

150

100

50

0
Apr- May- Jun- Jul- Aug- Sep- Oct- Nov- Dec- Jan- Feb- Mar- Apr- May- Jun- Jul- Aug-
98 98 98 98 98 98 98 98 98 99 99 99 99 99 99 99 99

Month

Figure 9. Cumulative number of dolphins identified in San Diego and Santa Barbara
between April 1998 and August 1999. Thin dotted line (between December 1998 and
January 1999 in San Diego) denotes month with no survey effort.
 34

Table 2. Summary information on survey effort, study period, and photographic results from
San Diego between 1984 and 1989. Photographic results differ from those given in Defran
and Weller 1999 and Dudzik 1999 due to revision of the dataset over time and elimination of
sightings not meeting the specified photographic quality criteria.

Number Number Sighting

Study Period Number of of of School Size Dolphins Frequency ResightingsResightings
Surveys Dolphins Schools Mean±SD Identified Mean±SD from from
(Number Complete) (Range) (Range) 1984-1989 1996-1998
January 1984 - 146 (109) 2869 145 19.8 ± 18.41 323 4.6 ± 3.25
December 1989 (2 - 90) (1 - 19)

March 1996 - 52 (20) 1233 63 19.6 ± 25.19 206 2.0 ± 1.21 108
March 1998 (1 - 140) (1 - 7)

April 1998 - 43 (28) 912 58 15.7 ± 18.83 204 2.0 ± 1.49 103 124
August 1999 (1 - 96) (1 - 8)
Total 241 (157) 5014 266 18.9 ± 20.29 468 5.0 ± 4.21
(1 - 140) (1 - 24)
 35

45
40
35
Mean School Size

30
25
20
15
10
5
0
1984 1985 1986 1987 1988 1989 1996 1997 1998 1999
Year

Figure 10. Mean school size across years for all schools sighted in San Diego between 1984-
1999. Error bars represent standard deviations.
 36

60

Mean Number of Dolphins Per Survey

50

40

30

20

10

0
1984 1985 1986 1987 1988 1989 1996 1997 1998 1999
Year

Figure 11. Mean number of dolphins sighted per complete survey in San Diego between
1984 and 1999. Error bars represent standard deviations.
 37

observed per survey, both of these parameters were higher in the latter years of the study
(1988-1989, 1996-1999) than during the earlier years (1984-1987).
Occurrence patterns were also compared across oceanographic seasons for the long-
term database. When data from all complete surveys were pooled, the mean number of
dolphins observed per complete survey (F(2,154) = 5.31, P = 0.006) was lower during the
Oceanic season than during the other two seasons (Figure 12). Dolphins were also
encountered significantly less often on complete surveys during the Oceanic season than
during the Upwelling season (X² = 10.94, df = 2, P = 0.004), though no significant difference
was found between the Oceanic and Davidson period. School size was significantly greater
(F(2,263) = 12.50, P<0.001) during the Davidson season when compared with other
oceanographic seasons (Figure 13). Each year with observations across all three
oceanographic seasons was also examined individually for evidence of seasonal occurrence
patterns. Results varied, with some years demonstrating no evidence of seasonal patterns or
showing seasonal patterns inconsistent with those observed in 1998-1999. In general,
however, the seasonal pattern of increased usage of the San Diego study area during the
Davidson period was more marked during the latter years of the study (1996-1999) than
during the earlier years (1984-1989).

El Niño
The influence of El Niño events on occurrence patterns of Pacific coast bottlenose
dolphins was examined using data collected in San Diego between 1984 and 1999. This
period encompassed a relatively mild El Niño/La Niña event between 1987 and 1988 and a
stronger El Niño/La Niña event between 1997 and 1999. For the 1987-1988 El Niño/La Niña
event, no significant differences between El Niño, La Niña, and normal months were found
in school size (F(2, 207) = 0.27, P>0.05), number of dolphins observed per survey (F(2,127)
= 1.71, P>0.05), or number of surveys encountering at least one group of dolphins (X² = 5.36,
df = 2, P>0.05). Similarly, no significant differences between El Niño, La Niña, or normal
months were found in school size (F(2, 213) = 0.52, P>0.05), number of dolphins observed
per survey (F(2, 116) = 0.47, P>0.05), or number of surveys encountering at least one group
of dolphins (X² = 3.56, df = 2, P>0.05) for the 1997-1999 El Niño/La Niña event.
 38

45
40
Mean Number of Dolphins

35
30
25
20
15
10
5
0
Upwelling Oceanic Davidson
Oceanographic Seasons

Figure 12. Mean number of dolphins per complete survey as a function of oceanographic
season in San Diego between 1984 and 1999. Error bars represent standard deviations.
 39

45
40

Mean School Size

35
30
25
20
15
10
5
0
Upwelling Oceanic Davidson
Oceanic Season

Figure 13. Mean school size as a function of oceanographic season for surveys conducted in
San Diego between 1984-1999. Error bars represent standard deviations.
 40

Distribution
Distribution of sightings was examined using the 1984-1999 dataset in the San Diego
study area. The mean number of dolphins found in each 1 km segment per unit survey effort
was 0.5 (SD = 0.26) (Figure 14). This ratio of dolphins per unit effort was used to classify
each zone as one of low, moderate, or high usage according to the same criteria used for the
short-term database. Zones of low usage included zones 19 and 17, while zones of high
usage included zones 1, 6, 8, and 14. In general, zones located in the southern half of the
study area (zones 1-16) were used more heavily than those in the northern portion (zones 17-
32). Table 1 shows some of the important topographical or environmental features in each
zone.

Site Fidelity
A total of 468 dolphins were identified in the San Diego study area between 1984 and
1999. The number of sightings per dolphin (Figure 15) ranged from one to 24, and the mean
number of sightings per dolphin was 5.0 (SD = 4.21). One hundred four (22.2%) of the
dolphins identified were sighted only one time. Of the 204 dolphins identified in San Diego
between 1998-1999, 77.0% (n = 157) had been identified in previous studies. Only 15.0% (n
= 70) of the dolphins identified were sighted during all three study periods.
The sighting histories of the dolphins (n = 63) sighted ten or more times in the San
Diego study area, and therefore considered to be “frequently sighted”, were examined in
detail. Fifty-three (84.1%) of these dolphins were identified in all seasons, with the
remaining 15.9% (n = 10) being sighted in both the Upwelling and Davidson seasons but not
the Oceanic season. Forty-eight (76.2%) of these dolphins were sighted in areas other than
San Diego. The mean number of years this subset of frequently sighted dolphins was
identified was 5.8 (SD = 1.58) and ranged between two and nine years.
The number of sightings of dolphins identified as females (n = 22) during any part of
the study was compared with the number of sightings of the remainder of identified dolphins.
Females had higher sighting frequencies (F(1,466) = 19.64, P<0.001) than other identified
dolphins.
 41

1.4

Dolphins Per Unit Survey 1.2

0.8
Effort

0.6

0.4

0.2

0
1
3
5
7
9
11
13
15
17
19
21
23
25
27
29
31
1 km Zones

Figure 14. Number of dolphins observed per unit survey effort over 1 km segments of the
San Diego study area between 1984-1999. Segments run south to north.
 42

120

Number of Dolphins 100

80

60

40

20

0
1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24
Number of Sightings

Figure 15. Sighting frequencies of dolphins identified in San Diego between 1984-1999.
 43

Movements
Movements between areas were examined by comparing the San Diego 1984-1999
dataset with the Santa Barbara 1998-1999 dataset as well as with datasets from surveys
conducted in Santa Barbara, Orange County, and Ensenada between 1982 and 1989 by
Defran and colleagues (see Defran et al. 1999). Survey effort and number of dolphins
identified in each area are shown in Table 3. Two hundred fifty-four (54.3%) of the dolphins
identified in San Diego were seen in at least one other study area. Five dolphins were
identified in all four study areas, and 59 dolphins were seen in at least three study areas. Of
the 47 dolphins identified in Santa Barbara in 1987 and 1989, 34.0% (n = 16) were resighted
in Santa Barbara between 1998 and 1999.

Rate of Discovery
A rate of discovery curve plotting the cumulative number of dolphins identified in the
San Diego study area between 1984-1989 is shown in Figure 16. Survey effort was divided
into blocks of 5 surveys on which at least one dolphin was identified (n = 162 surveys), with
the exception of block 19 and block 33 which contained only four and three surveys,
respectively. A break in survey effort of over six years occurred between block 19, which
ended in December 1989, and block 20, which began in March 1996. The curve shows a
continual increase throughout most of the study but begins to reach an asymptote toward the
end of the study period, with only one new dolphin identified in San Diego during the final
months of the study.
 44

500
450
CumulativeNumber of
Dolphins Identified
400
350
300
250
200
150
100
50
0
1 3 5 7 9 11 13 15 17 19 21 23 25 27 29 31 33
Survey Blocks

Figure 16. Rate of discovery curve showing the cumulative number of dolphins identified
over blocks of five surveys on which at least one dolphin was identified. Block 19 includes
only 4 surveys, and Block 33 contains only 3 surveys. A break in survey effort (shown with
dotted line) from Dec 1989 to Mar 1996 occurs after block 19.
 45

Table 3. Summary information on survey effort, study period, and photographic data for
Southern California Bight study areas 1982-1999. Photographic data differ slightly from
information presented in Defran et al. 1999 due to revision of the dataset over time and
the elimination of sightings not meeting the specified photographic quality criteria.

Study Area Dates Number of Number of Dolphins Number of Dolphins

Surveys Identified Resighted in San Diego
San Diego 1984-1989, 1996-1999 241 468 ------------------------------------
Santa Barbara 1987, 1989, 1998-1999 73 209 75.6% (n=158)
Orange County 1982-1989 44 118 82.2% (n=97)
Ensenada 1985-1986 11 66 80.3% (n=53)
Total 1982-1999 346 533 ------------------------------------
 46

DISCUSSION

This study analyzed the occurrence patterns, site fidelity, and movements of coastal
bottlenose dolphins within two areas of the Southern California Bight. Although Pacific
coast bottlenose dolphins have been studied since 1981 (Hansen 1990, Wells et al. 1990,
Weller 1991, Caldwell 1992, Feinholz 1996, Defran and Weller 1999, Defran et al. 1999,
Dudzik 1999, Marsh 2000, Defran et al. In prep), many questions remain about the effect of
ecological and environmental variables on their behavior. The present study contributed to
our understanding of the behavioral ecology of the population by allowing comparisons to be
made over both space and time.
The extension of previous studies of Pacific coast bottlenose dolphins in San Diego
coastal waters permitted temporal changes in behavior to be examined. Bottlenose dolphins
are long-lived mammals, existing in complex societies and exhibiting a high degree of
behavioral flexibility. Long-term studies allow documentation of the persistence of
behavioral patterns in the population as a whole over time, offering further understanding of
the effect of environmental variables on behavior. Longitudinal studies also present an
opportunity to observe changes in the behavior of individuals over time, providing an
indication of the effect of internal constraints on behavior.
In addition, concentrated study of Pacific coast bottlenose dolphins in Santa Barbara
waters provided a spatial dimension for comparison, by allowing dolphin behavior from the
same population in two different parts of their range to be compared. While both areas share
similar types of habitats, some differences in topography and temperature exist between the
two areas (Dorman and Palmer 1981). Santa Barbara lies approximately 270 km to the north
of San Diego and is somewhat sheltered from prevailing wind and sea conditions by the
Channel Islands. The San Diego coastline, however, is fully exposed to direct western swells
(Littler 1980). The continental slope also drops more rapidly and steeply off the San Diego
coastline than in Santa Barbara, where the slope is more gradual (Dorman and Palmer 1980).
Furthermore, the southernmost end of the San Diego study area contains a submarine canyon,
providing an area of increased topographical relief. These differences in topography and
openness are unlikely to affect dolphin behavior directly. They may influence the
 47

distribution of prey resources, however, and thereby provide an indirect index of the effect of
subtle habitat differences on dolphin behavior. Additionally, continued study of Pacific coast
bottlenose dolphins has allowed more direct comparisons to be made with other long-term
studies of coastal bottlenose dolphins in Sarasota Bay, Florida (Wells 1986) and in Aransas
Pass, Texas (Weller 1998, Maze and Würsig 1999). Geographic comparisons of this nature
allow the effects of large-scale habitat differences on behavior of bottlenose dolphins to be
explored.

Occurrence Patterns

Examination of the occurrence patterns of Pacific coast bottlenose dolphins within the
Southern California Bight can provide one indication of the effect of environmental factors
on the behavior of the population. Occurrence patterns are primarily influenced by a
combination of direct limitations to environmental characteristics (e.g. water temperature and
salinity), predation pressure, the need to reproduce, and prey distribution and abundance.
Bottlenose dolphins within the Southern California Bight live in a temperate environment,
while populations in other regions, such as off the coast of Scotland, are known to withstand
more extreme oceanographic conditions (Wilson et al. 1997a). Thus, variables such as water
temperature and salinity should not directly limit dolphin movements within the coastal
Southern California Bight environment. Predation pressure, as well, probably provides
minimal influence on the behavior patterns of coastal California bottlenose dolphins (Weller
1991). The shark species most commonly associated with bottlenose dolphin predation,
which include tiger (Galeocerdo cuvier), dusky (Carcharhinus obscurus), bull
(Carcharhinus leucas) and great white (Carcharadon carcharias) sharks (Wood et al. 1970,
Corkeron et al. 1987), are rarely found within the coastal Southern California Bight
environment (Barnhart 1936). Killer whales, considered to be another dolphin predator
(Würsig and Würsig 1979), are also infrequently sighted in the area. Influences associated
with the need to reproduce would be reflected in behavioral differences between individuals
of different sex, age, or reproductive classes, and will be discussed in later sections.
Given these factors, occurrence patterns of the population as a whole are most likely
to be influenced by prey distribution and abundance. Little direct information, however, is
available about the diet of Pacific coast bottlenose dolphins. Walker (1981) analyzed the
 48

stomach contents of nine southern California coastal bottlenose dolphins and concluded that
the primary prey species of coastal dolphins were fish and invertebrates inhabiting the littoral
and sublittoral zones. The diet was diverse, with fish from 12 families and invertebrates
from six families represented in the stomachs, but the majority (62%) of the species ingested
came from two families of fish, Sciaenidae (croakers) and Embioticidae (perches). These
results were based primarily on stomach contents of stranded animals, however, and as such
may not have reflected the typical diet of healthy animals.
Despite the paucity of direct information on the diet of coastal bottlenose dolphins
within the Southern California Bight, indirect information on the influence of prey
distribution and abundance on dolphin behavior can be provided by examining occurrence
patterns across study areas, seasons, and years. Between 1998 and 1999, occurrence patterns
of bottlenose dolphins in San Diego and Santa Barbara coastal waters were similar,
indicating that dolphins used both of these areas with comparable intensity. When
occurrence patterns were compared across oceanographic seasons, however, only San Diego
showed evidence of seasonal patterns of occurrence. While dolphins were present in San
Diego waters year-round, the number of dolphins observed per survey as well as the size of
observed schools was greatest during the Davidson season.
Similar results were found during behavioral studies by Ward (1999) between 1996
and 1997, who postulated that the larger school sizes of coastal bottlenose dolphins in San
Diego during late fall and winter were correlated with the presence of market squid (Loligo
opalescens) off La Jolla during winter months. While market squid spend much of their life
cycle in offshore waters, they move inshore to spawn, often forming large aggregations near
marked submarine topographic features (Recksiek and Frey 1978). In southern California
spawning is most evident between December and February (Fields 1965, Recksiek and Frey
1978) and has been observed near La Jolla Canyon (McGowan 1954, Limbaugh and Shepard
1957, Fields 1965), which is adjacent to Scripps Canyon and the southern terminus of the
San Diego study area. The influx of dolphins into the San Diego study area during the
Davidson period, as reflected by both greater numbers of dolphins within the study area as
well as larger school sizes, may represent dolphins taking advantage of this concentrated prey
resource. While this short-term exploitation may entail dolphins feeding directly on squid
 49

aggregations, the spawning squid may also be attracting predatory fish that are important
prey of the dolphins.
Increased usage of the San Diego study area during the Davidson oceanographic
season was not as evident over the long-term dataset (1984-1999) as it was during the short-
term study (1998-1999). While school sizes remained significantly higher during the
Davidson period than during other oceanographic seasons, numbers of dolphins in the area
were similar during the Davidson and Upwelling seasons and decreased during the Oceanic
season. Less marked increase in the number of dolphins using the area during the Davidson
season over the long-term study may be further related to exploitation of market squid.
Because squid have a relatively short life span, generally less than two years, they are
extremely responsive to inter-annual fluctuations in biological and physical variables (Boyle
and Boletzky 1996), and their impact on the ecosystem as both predator and prey varies
considerable from year to year (Beddington et al. 1990, Rodhouse and Nigmatullin 1996).
Those years with no pattern of increased usage of the San Diego study area during the
Davidson period may represent years in which the size of market squid spawning
aggregations was reduced, resulting in more widespread distribution of dolphins.
As well, variation in the degree of upwelling across oceanographic seasons may
affect prey distribution and abundance and thereby influence seasonal occurrence patterns of
dolphins. May and June are the maximum upwelling periods in the vicinity of Scripps Pier
(Dailey et al. 1993), which are often correlated with the presence of unusually high
concentrations of demersal fish nearshore (Moyle and Cech 1988). Additionally, the
majority of coastal marine fish species in southern California, including those consumed by
bottlenose dolphins, aggregate in spring and summer to spawn (Hansen and Defran 1993).
The overall increased availability of prey in San Diego during the Upwelling season may
serve to bring high numbers of dolphins into the area, but dolphins may be dispersed into
smaller groups as a reflection of abundant and more evenly dispersed prey resources (Weller
1991). Seasonal occurrence patterns during the long-term study (1984-1999) supported this
hypothesis, as the numbers of dolphins in the area were similar during the Davidson and
Upwelling seasons, while school sizes were greater in the Davidson period than in the
Upwelling season. During the Davidson season, when overall productivity within the
Southern California Bight is reduced, prey resources may be less evenly distributed and more
 50

difficult to find. Larger schools may be beneficial in this situation, as dolphins may integrate
their sensory capabilities in order to increase the probability of finding food (Norris and Dohl
1980).
Annual variation in occurrence patterns of Pacific coast bottlenose dolphins in the
San Diego study area was also evident. While school size did not differ significantly across
years (1984-1999), both the number of dolphins observed per survey and the number of
surveys encountering at least one school varied across years. Variation in dolphin occurrence
patterns may be a reflection of the highly dynamic ecosystem of the Southern California
Bight, which undergoes significant daily, monthly, seasonal, and yearly fluctuations
(SCCWRP 1973, Dailey et al. 1993). The effect of such environmental variability can be
seen at the lowest trophic levels, with changes in both phytoplankton and zooplankton
biomass thought to be correlated with yearly changes in the flow of the California Current
system as well as with changes in the intensity and timing of local coastal upwelling events
(Tont 1976, 1981; Roesler and Chelton 1987). The influence of environmental variability is
further manifested in the recruitment and spawning success of some species of fish. For
example, rockfish recruitment within the Southern California Bight increases during warmer
years, when both transport by the California Current and upwelling are lower, and decreases
during colder years when transport and upwelling are higher (Norton 1987).
As well, during the latter years of the study (1997-1999), the Southern California
Bight was being affected by an El Niño event of similar strength to that of 1982-1983
(Wolter and Timlin 1998). During El Niño events, water temperatures within the Southern
California Bight are elevated several degrees above normal (Dailey et al. 1993). While it is
unlikely that changing water temperatures affect Pacific coast bottlenose dolphins directly,
changes in distribution and abundance patterns (Fiedler et al. 1986, Bragg 1991, Hollowed
and Wooster 1992, Ainley et al. 1995, Stull and Tang 1996) as well as species composition
(Westerhagen 1993) of marine biota are known to occur as a consequence of changes in
habitat quality (Woodbury 1999) and direction of currents (Yoklavich et al. 1996, Connolly
and Roughgarden 1999). As mentioned previously, the range of Pacific coast bottlenose
dolphins was extended following the 1982-83 El Niño, presumably in response to changing
prey distributions (Wells et al. 1990). El Niño events have also been documented to affect
foraging in California sea lions (Zalophus californianus). During the 1982-83 El Niño, the
 51

relative contributions of different food types in the diet of California sea lions was altered,
apparently in response to changing availability of prey (DeLong et al. 1991). California sea
lions were also affected by the 1997-1998 El Niño, when pup production decreased by 64%,
presumably due to nutritional stress (Forney et al. 2000).
Based on the Bivariate Enso time series index (Smith and Sardeshmukh 2000),
however, the occurrence patterns of Pacific coast bottlenose dolphins did not vary between El
Niño, La Niña, and normal months. While it is possible that Pacific coast bottlenose dolphin
occurrence patterns did not shift in response to El Niño conditions, the index used, which
was based on sea surface temperature and atmospheric pressure, may not have accurately
reflected environmental cues dolphins use to change behavior, or some lag time might have
existed before changing conditions were reflected in dolphin behavior.

Distribution

While comparison of occurrence patterns between years, seasons, and study areas has
provided information on large-scale environmental factors affecting the behavioral ecology
of Pacific coast bottlenose dolphins, examination of sightings within each study area can
offer an indication of more finely scaled variables shaping dolphin behavior. Although
characterizing all potentially influential habitat variables was beyond the scope of this study,
some general patterns emerged when the distribution of sightings was examined. As
demonstrated in previous studies of Pacific coast bottlenose dolphin distribution (Hansen
1990, Caldwell 1992, Hanson and Defran 1993, Feinholz 1996, Caretta et al. 1998, Defran
and Weller 1999, Defran et al. 1999, Ward 1999, Dudzik 1999), the majority of sightings in
both study areas were made within 0.5 km from shore, and no dolphins were sighted beyond
1 km of the coastline. Nearshore waters of the Southern California Bight are biologically
rich and diverse, supporting numerous trophic levels and a wide diversity of habitats
including submarine canyons, sandy bottoms, rocky reefs, and kelp beds. Coastal waters also
receive a greater supply of nutrients than do waters further from the coast, due to terrestrial
runoff and increased vertical mixing of nutrients caused by upwelling and tidal cycling. The
productive waters, in combination with the low levels of predation and interspecific
competition thought to exist in coastal waters (Weller 1991), make the nearshore zone a
favorable environment for Pacific coast bottlenose dolphins.
 52

Within the San Diego study area, the most prominent distributional pattern was heavy
usage of areas associated with submarine canyon habitat, both in the long-term (1984-1999)
and short-term (1998-1999) datasets. Behavioral studies by Ward (1999) found increased
feeding of Pacific coast bottlenose dolphins in canyon versus non-canyon habitat during his
behavioral study between 1996 and 1997. Submarine canyons are nutrient rich areas due to
increased levels of upwelling, which are generated as tide-driven water masses flow up and
down the canyon walls (Svedrup et al. 1942). The topography of submarine canyons also
produces a diversity of microhabitats, ranging from rocky canyon shoulders to dense mats of
surf grass and kelp detritus. This diversity hosts organisms of many different trophic levels,
beginning with amphipods and invertebrates inhabiting mats and attracting both demersal
and pelagic fish (Vetter 1994). These fish may provide a prey resource for Pacific coast
bottlenose dolphins. Increased usage of canyon habitat during this study was also consistent
with the observed occurrence patterns, which indicated that dolphins use the San Diego study
area heavily during winter months when market squid are spawning near the canyon. The
combination of both consistent prey resources, in the form of fish attracted to the area by the
diversity of micro-habitats and increased levels of nutrients due to upwelling, and temporally
abundant resources, represented by aggregations of market squid during spawning season in
some years, probably makes areas surrounding the canyon preferential habitat for Pacific
coast bottlenose dolphins.
Areas of heavy use within both the San Diego and Santa Barbara study areas were
also examined for correlations with estuaries. Estuaries are sites of high concentrations of
nutrients, which support large numbers of invertebrates and fish (Vannucci 1969, Thomson
1973, Moyle and Cech 1988). Previous studies comparing Pacific coast bottlenose dolphin
use of estuarine areas within the San Diego study area have had mixed results. Although no
formal analysis was conducted, Hansen (1990), whose study area extended north to
Oceanside, observed that the distribution of Pacific coast bottlenose dolphins was
concentrated between Torrey Pines State Reserve and south Carlsbad and attributed this in
part to the presence of lagoon mouths providing nutrient rich areas. Hanson (1990),
however, found the proportion of time spent feeding to be greater in areas containing reefs
than in those surrounding estuarine areas. Results from this study were inconclusive. While
several small lagoon estuaries exist within both the San Diego and Santa Barbara study areas,
 53

most estuarine areas showed only low to moderate levels of usage between 1998 and 1999.
When data collected in San Diego between 1984 and 1999 were combined, however, zones
associated with the San Dieguito and Los Penasquitos lagoons demonstrated moderate to
high levels of usage, while zones associated with the San Elijo and Batiquitos lagoons
exhibited low to moderate levels of usage. Many of the small lagoon estuaries along the
California coast are closed to the sea for most of the year by wave built sand spits, which are
only breached during winter when rainfall is highest (Elwaney et al. 1998, Emmett et al.
2000). Without consistent contact with the surrounding ocean, the high levels of productivity
typically associated with estuaries may be too unpredictable to attract dolphins on a regular
basis.
Two 3 km stretches containing no sightings existed within the Santa Barbara study
area and included the zones lying directly southeast of the Santa Barbara Harbor and the
zones surrounding a pier used by ships associated with oil well operations. Both areas
contain high levels of large vessel traffic. While bottlenose dolphins in this study area have
been observed bow-riding in front of large boats, they may avoid use of these areas for
activities other than traveling and playing due to the increased levels of disturbance as well
as higher levels of pollution in the water associated with boats and oil well operations.
Comparison of the distribution of dolphin sightings in the San Diego study area
between 1998 and 1999 with the distribution of sightings during the combined 1984-1999
period demonstrated that sightings were spread more evenly throughout the study area over
the long-term study. The more regularly dispersed sightings over the long-term study
indicate that most portions of the study area are valuable to Pacific coast bottlenose dolphins
in some capacity and may be related to prey distribution. The high daily, monthly, and
annual variability in the coastal ecosystem of the Southern California Bight creates patchy
and unpredictable patterns of distribution of marine organisms, including prey species of the
bottlenose dolphin (SCCWRP 1973, Mearns 1974, DeMartini and Allen 1984, Ware and
Thompson 1991, Cross and Allen 1993, Dailey et al. 1993). Over longer time scales,
however, prey patches would occur over most portions of the study area, resulting in more
uniform usage of the entire area by bottlenose dolphins.
As expected, these results indicate that habitat usage by Pacific coast bottlenose
dolphins in the Southern California Bight is determined by a combination of many different
 54

factors. More complete analysis of habitat usage should incorporate not only more extensive
documentation of the habitats within the coastal Southern California Bight, but also
behavioral observations within different habitat types, extending the work done by Ward
(1999). Currently, Geographic Information System mapping techniques are often used in
both terrestrial and marine systems to overlay animal sightings with an array of different
habitat variables. The application of such techniques to the habitat of Pacific coast
bottlenose dolphins would provide a much more thorough understanding of the complex
factors affecting dolphin distribution.

Site Fidelity

Previous studies of Pacific coast bottlenose dolphins have reported low sighting
frequencies of individuals in San Diego coastal waters (Defran and Weller 1999, Dudzik
1999) as well as in less extensively studied secondary areas within the Southern California
Bight (Defran et al. 1999). Regional studies within the Southern California Bight further
indicated that the majority of dolphins identified in Santa Barbara (76%) and Orange County
(82%), California and Ensenada (80%), Baja California Norte, Mexico had also been sighted
within the San Diego study area. Taken together, these observations suggested that Pacific
coast bottlenose dolphins range over extensive coastal areas, demonstrating little preference
for any one area (Defran and Weller 1999, Defran et al. 1999, Dudzik 1999).
Results from the present study further indicated that Pacific coast bottlenose dolphins
are essentially transient within the Southern California Bight. Dolphins identified in both
study areas had an average of only two sightings despite the high number of surveys
conducted. Lack of site fidelity among Pacific coast bottlenose dolphins has been attributed
to the patchy and unpredictable distribution of prey (Defran and Weller 1999, Defran et al.
1999). Within the coastal Southern California Bight environment, most fish species either
are found in temporary local concentrations or are widely but sparsely distributed (SCCWRP
1973, Mearns 1974, DeMartini and Allen 1984, Ware and Thomson 1991, Cross and Allen
1993, Dailey et al. 1993). The patchy or sparse distributions of dolphin prey may be a factor
in the long distances traveled by Pacific coast bottlenose dolphins (Defran et al. 1999), as
patchy and sparse resource distributions necessitate that animals range widely to locate
sufficient prey (Krebs and Davies 1981). As well, Pacific coast bottlenose dolphin school
 55

sizes are large when compared with most other coastal bottlenose dolphin populations (Wells
et al. 1987, Corkeron 1990, Shane 1990, Smolker et al. 1992, Bräger et al. 1994, Fertl 1994),
which may be a further indication of patchy resource distribution within the Southern
California Bight. Large school sizes are thought to increase sensory integration and thereby
aid in locating difficult to find resources (Norris and Dohl 1980), and they may represent an
adaptation by Pacific coast bottlenose dolphins to increase efficiency in locating patchily
distributed prey (Weller 1991, Defran and Weller 1999, Defran et al. 1999).
The combined long-term database yielded further evidence that dolphins demonstrate
only low levels of site fidelity to the San Diego study area, with individual dolphins sighted
an average of only five times over the ten-year period of study. More detailed examination
of sighting patterns also indicated that lack of site fidelity was not correlated with seasonal
movements within the range, as most of those dolphins with the highest sighting frequencies
were observed in the San Diego study area over all three oceanographic seasons.
While dolphins in both study areas were sighted infrequently, sightings as a function
of effort were significantly higher within the San Diego study area. Higher ratios may reflect
the influx of dolphins into the San Diego study area during the Davidson season rather than
increased site fidelity to the area over the entire study period. The number of dolphins
identified in San Diego during January 1999 was almost twice as high as the greatest number
of dolphins identified during any one month in Santa Barbara, despite higher survey effort in
Santa Barbara than in San Diego for the months compared. Thus the higher levels of site
fidelity observed in the San Diego study area between 1998 and 1999 may not be an
indication of consistent and long-term differences in site fidelity patterns between the two
areas but may instead reflect differences in prey resource distribution during the Davidson
oceanographic season of this particular study period.
Individual differences in site fidelity were explored by comparing dolphins identified
as probable mothers and females to the rest of the population. Between 1998 and 1999,
dolphins identified as females, as well as the subset of those females identified as probable
mothers, had higher sighting per kilometer ratios than did other dolphins identified in Santa
Barbara. Females and probable mothers in San Diego did not demonstrate increased site
fidelity between 1998 and 1999; however, females in San Diego did have higher sighting per
kilometer ratios over the long-term study (1984-1999). Higher site fidelity of females and
 56

probable mothers is likely related to differences in reproductive strategies between sexes. In

most mammals, males provide no parental care and focus on impregnating as many females
as possible to maximize their reproductive success. Female reproductive success, however,
relies on raising young to maturity. The higher energetic demands of both pregnancy and
lactation may require that females maximize energy intake while minimizing movements
within the range. As such, female distribution, when compared with that of males, is often
more closely linked with resource distribution (Bradbury and Vehrencamp 1977, Emling and
Oring 1977, Wrangham 1980).
The higher levels of site fidelity in females not identified as mothers may reflect
formation of groups of females to protect calves, as has been observed in sperm whales
(Whitehead and Weilgart 2000). Evidence of increased levels of association between
probable mothers in the Santa Barbara study area between 1998 and 1999 was documented
(Marsh 2000), indicating that at least those dolphins with calves may group together over
relatively short time spans. However, predation pressure on coastal bottlenose dolphins
within the Southern California Bight is thought to be low (Weller 1991), and no evidence of
long-term stable associations between individuals has been observed (Weller 1991, Marsh
2000). The higher levels of site fidelity of females over the long-term study in San Diego
may simply reflect that these females were pregnant or were, although not identified as such,
associated with calves during the study. Bottlenose dolphin calves generally remain with
their mothers for three to four years after birth (Wells et al. 1987, Smolker et al. 1992) and
often are not weaned until the female is in the latter stages of her next pregnancy (Mann et al.
2000). Thus, most females of reproductive age in the population would probably have been
either pregnant or lactating during the study, and as such would have been under increased
nutritional demands when compared to the remainder of the population.
The lack of increased site fidelity in probable mothers and females observed in San
Diego during the short-term study may be due to several factors. While a higher number of
total dolphins were identified in San Diego than in Santa Barbara coastal waters, fewer
probable mothers and females were documented in San Diego during the short-term study.
This difference is probably not due to greater number of females or probable mothers using
the Santa Barbara study area but rather is related to the more strenuous survey conditions
found in the San Diego study area. Calf behavior was observed closely before observers
 57

made decisions on which adult would be identified as the probable mother. The calmer and
more protected waters of the Santa Barbara study area facilitated close and lengthy
observations of dolphins, while more direct exposure to swells within San Diego coastal
waters often prevented the close and prolonged observations necessary to identify probable
mothers. While these same conditions might be responsible for true differences between
areas, with mothers with calves preferring the more sheltered waters provided by the Santa
Barbara coastline, the higher levels of site fidelity observed in females over the long-term
study in San Diego argue against any consistent differences between the two areas. In
general, however, ambiguity in results from analyses of female site fidelity between and
within areas may be due in part to the small number of dolphins identified as females and as
probable mothers relative to the total number of identified dolphins and illustrates the need
for further studies to be conducted before any conclusions are drawn.
While females and probable mothers did demonstrate increased site fidelity when
compared to other identified individuals, levels of site fidelity in both female and other
Pacific coast bottlenose dolphins are low when compared to other populations containing
more residential dolphins (e.g. Wells et al. 1987). Dolphins identified as females in San
Diego coastal waters were identified a mean of only nine times over the ten-year span of the
study, and all except two identified females (10%) had been sighted in other study areas.
Thus while female occurrence patterns may be more closely linked with prey resource
distribution than those of males, females must still range over extensive areas to meet
nutritional demands.

Movements

Between 1998 and 1999, 89 dolphins were documented to move between San Diego
and Santa Barbara, representing 50% and 44% of the number of dolphins identified in Santa
Barbara and San Diego, respectively. Since the study areas were not surveyed on a daily
basis, these percentages likely underestimate the actual number of individuals moving
between the two areas. They can, however, be considered a minimum estimate of the
number of dolphins moving between the two areas during the seventeen months of the study.
The frequency of documented movements between the two areas was even higher, with some
individuals moving between areas as many as four times. The high percentage of individuals
 58

documented to move between the two areas, combined with the large number of observed
movements, provides further evidence that Pacific coast bottlenose dolphins are highly
mobile within the Southern California Bight.
When the long-term dataset was used to examine movements of dolphins between
areas, over half of the dolphins identified in San Diego between 1984 and 1999 had been
sighted in at least one other region of the Southern California Bight. Even among the subset
of most frequently sighted dolphins, most were sighted in other study areas, indicating that
even those dolphins with relatively high sighting frequencies in San Diego waters range
widely throughout the Southern California Bight.
Results obtained by combining data collected between 1996 and 1999 in San Diego
and Santa Barbara with that obtained in San Diego, Orange County, and Santa Barbara,
California and Ensenada, Baja California Norte, Mexico between 1982 and 1989 (Defran et
al. 1999) demonstrates the effect of continued survey effort on resighting percentages. In
both Ensenada and Orange County, where survey effort was discontinued after 1989, the
percent of dolphins identified in those areas and resighted in at least one other study area
increased (from 88% to 91% in Ensenada and from 92% to 95% in Orange County) with the
addition of the 1996 through 1999 surveys. The high percentages indicate that even without
coverage of most parts of the range of Pacific coast bottlenose dolphins, continued survey
effort over time in only small parts of the range is sufficient to identify the majority of
dolphins in the population.
The fastest travel speed documented during the seventeen-month study was 61.6
km/day, reached by one dolphin moving between San Diego and Santa Barbara over five
days. This speed is higher than the maximum speed previously observed, in which three
dolphins averaged 47 km/day over 2 days when moving between San Diego and Orange
County, California (Defran et al. 1999). Bottlenose dolphins in other areas have been
observed to travel at similar or faster speeds. One dolphin off the southwest coast of Britain
traveled 1076 km over 20 days, for an average of 53.8 km/day (Wood 1998). Two stranded
and rehabilitated bottlenose dolphins were released into the wild off the west coast of Florida
and subsequently tracked using satellite tags to determine the success of their release (Wells
et al. 1999). One individual traveled at an average of 48 km/day, covering 2050 km over the
first 43 days after his release and attaining speeds of at least 75 km/day as he rounded the tip
 59

of Florida. The second dolphin traveled 4200 km over 47 days, for an average of 89 km/day.
Both dolphins were thought to have been of the offshore ecotype, which may range over
greater distances than coastal dolphins. However, they illustrate the fact that bottlenose
dolphins are capable of covering great distances in a short amount of time.

Rate of Discovery Curve

Rate of discovery curves represent the cumulative number of individual dolphins

identified within a study area, generally plotted against either survey effort or time. These
curves never flatten out entirely due to new dolphins being “born” (i.e., becoming marked
and thereby identifiable) into the population; however, when the majority of individuals have
been identified the curve begins to reach an asymptote which can be roughly correlated with
the number of marked individuals in the population. Previous studies of the Pacific coast
bottlenose dolphin population by Defran and Weller (1999) between 1984 and 1989 and by
Dudzik (1999) between 1996 and 1998 have demonstrated that the rate at which new
dolphins are first identified in the San Diego study area continues to increase with effort even
over extended periods of time. The rate of discovery curve generated by Defran and Weller
(1999) began to reach a possible asymptote only after 146 surveys across six years of study,
during which a total of 373 dolphins were identified. Dudzik (1999) identified a total of 233
dolphins over 66 surveys, with only a slight leveling of the curve after 29 months of study.
While several methods have been employed to estimate the size of the population of
Pacific coast bottlenose dolphins (Hansen 1990, Defran and Weller 1999), the most recent
calculations by Dudzik (1999) estimated that between 289 and 356 dolphins use the San
Diego study area as part of their range. These estimates used sighting data between 1984 and
1998 and indicated that the population size of Pacific coast bottlenose dolphins over the
fourteen -year period was relatively stable. Abundance estimates have also been generated
using tandem aerial surveys covering the area between the U.S. Mexican border and Point
Conception. This method, which represents not population size but rather abundance of
dolphins using the study area, yielded estimates ranging between 78 and 271 dolphins
(Caretta et al. 1998).
Rate of discovery curves can also be used to provide rough estimates of the number
of identifiable individuals within a population. The rate of discovery curve generated for San
 60

Diego between 1984 and 1999 showed a continual increase throughout most of the study,
although the curve flattened out over the last four months, during which time only one
previously unidentified dolphin was photographed. By the end of the study, a total of 468
dolphins had been photographed in the San Diego coastal waters, and the increased leveling
of the curve over the summer of 1999 indicated that the majority of dolphins in the
population had been identified. Estimates of the number of identifiable dolphins within a
population, as determined through rate of discovery curves, are not directly comparable with
population estimates for two reasons. First, rate of discovery curves do not account for
unmarked (and thus unidentifiable through photo-identification) dolphins in the population.
Along the Pacific coast, Hansen and Defran (1990) estimated that approximately 35% of
coastal bottlenose dolphins were unmarked, and thereby not represented in the curve. The
rate of discovery curve also does not account for mortality of any identified dolphins over the
sixteen-year span of the study. Wells and Scott (1990) determined the mortality rate for
bottlenose dolphins in Sarasota Bay, Florida, to be between 0.010 and 0.038 per year. If
mortality rates for the Pacific coast population are consistent with those observed in Florida,
and the population estimates of Dudzik (1999) are assumed to be accurate, then between 46
and 216 dolphins may have died between 1984 and 1999. Incorporation of both these factors
with the number of identified individuals determined by the rate of discovery curve would
produce estimates more closely corresponding to those calculated by Dudzik (1999).
Rate of discovery curves produced from data collected in San Diego and in Santa
Barbara between 1998 and 1999 were compared to determine if any differences existed in the
rate at which dolphins were identified in each area. Both rate of discovery curves continued
to increase throughout the study, with only a slight leveling trend in both curves toward the
end of the study. While the curves are not entirely equivalent due to differences in survey
effort in each area, some interesting patterns emerged when the curves were compared. The
curve produced for Santa Barbara, which demonstrated no differences in seasonal occurrence
patterns during the study, increased at a relatively consistent rate throughout the study, with
the largest increase in the number of new dolphins identified in Santa Barbara occurring
between August and September 1998. The curve for San Diego, however, showed more
dramatic fluctuations in the rate of increase, with a leveling trend observed between July and
October 1998 followed by a sharp increase from October 1998 to January 1999. This pattern
 61

is consistent with the observed seasonal occurrence patterns, in which school size and total
number of dolphins using the area were highest during the Davidson season and lowest
during the Oceanic season. Decrease in the number of previously unidentified dolphins using
the San Diego study area during the Oceanic season corresponded with an increase, although
less marked than the seasonal pulse observed in San Diego, of new dolphins into the Santa
Barbara study area.

Comparisons with Other Study Areas

On an even larger scale, comparisons between bottlenose dolphin populations

occupying different habitats can help identify the selective environmental factors shaping
behavior and ecology. For this purpose, results obtained through long-term study in the
Southern California Bight have been contrasted with those from two other studies of
bottlenose dolphins in Sarasota Bay, Florida, and Aransas Pass, Texas. Studies in these two
areas were chosen for comparison because they represent longitudinal research in
environments with habitat characteristics different from those found along the Pacific coast.
Studies in all three areas have been ongoing for many years. Research conducted in
Sarasota Bay began in 1970, and continues to date (Irvine and Wells 1972, Wells et al. 1980,
Irvine et al. 1981, Wells et al. 1987, Scott et al. 1990, Wells 1991, Barros and Wells 1998).
This study is impressive not only in its longitudinal scope but also in the diversity of work
conducted in the area. The sheltered and shallow waters of the inner bays have allowed
dolphins to be temporarily captured, and as a result age, sex, reproductive condition, and
familial relationships are known for many members of the Sarasota community. This
information, in conjunction with knowledge of individual sighting histories and occurrence
patterns obtained through boat-based photo-identification studies, has been used not only in
studies of behavior and ecology but also in research on the acoustics, genetics, and
physiology of the population (Hohn et al. 1989, Sayigh et al. 1990, Duffield and Wells 1991,
Tolley et al. 1995). The integration of knowledge gained from such a range of biological
disciplines has led to one of the most comprehensive pictures of a dolphin society available
today. Work in both the Southern California Bight and along the Texas coast has been more
limited in duration and scope, in part because of the inaccessibility of these environments
relative to that found in Sarasota. Work along the Southern California Bight began in 1981
 62

and has continued through 1999 (Hansen 1990, Wells et al. 1990, Weller 1991, Caldwell
1992, Feinholz 1996, Defran and Weller 1999, Defran et al. 1999, Dudzik 1999, Marsh 2000,
Defran et al. In prep). Initial work along the Texas coast consisted of several short-term
studies in different areas along the coastline (Shane 1980, Shane and Schmidly 1978, Gruber
1981, Jones 1988, Henningsen 1991, Bräger 1993, Bräger et al. 1994, Fertl 1994, Lynn 1995,
Maze and Würsig 1999). The first comprehensive effort, however, which covered four
different study areas along the Texas coast, was begun by Weller (1998) in 1991. Although
work from other areas is still ongoing, results from surveys of Aransas Pass, Texas, between
1991 and 1997, are presented in Weller (1998) and are used for the comparisons made here.
The environments found within the Southern California Bight, Sarasota Bay, and
Aransas Pass study areas differ in both habitat structure and oceanography. The environment
of the Southern California Bight is characterized as open, exposed, and highly dynamic, with
current-driven variation in water properties on daily, seasonal, and annual scales (Dailey et
al. 1993). Water temperatures vary from 14.5° C in late winter to 19°C in summer and early
fall, with corresponding changes in salinity from a low of 33.4 ppt to a high of 33.6 ppt
(Dailey et al. 1993). The distribution of dolphin prey species within the Bight is thought to
be patchy and unpredictable, as a response to the high variability in oceanographic variables
(Defran et al. 1999). At the other extreme, waters of Sarasota Bay, located on the western
coast of Florida, are sheltered by a series of barrier islands, containing a system of large bays
and inlets connected to the open ocean by channels and passes. Inner waters consist
primarily of shallow seagrass beds, and waters remain relatively calm due to the protection of
the barrier islands (Wells et al. 1987). Although fish distribution and abundance in most
marine systems is patchy at some spatial scale (Nybakken 1993), the small size
(approximately 125 km²) of the dolphins’ range within Sarasota Bay precludes wide dispersal
of patches, and the primary dolphin prey species are thought to be present year-round (Barros
and Wells 1998). The habitat structure of Aransas Pass, Texas, closely resembles that of
Sarasota Bay, with a series of bays and inner waters sheltered by barrier islands and
connected to the open ocean via dredged shipping lanes (Weller 1998). However, seasonal
variability in both salinity and temperature is more pronounced within Aransas Pass when
compared with the variability typical of Florida (Weller 1998). Sea surface temperatures
 63

range from a high of 30.0°C in the summer and a low of 8.3°C in the winter, while salinity
ranges from a high of 40 ppt in the summer to a low of 17 ppt in the spring (Copeland 1965).
The occurrence patterns and site fidelity characteristics of bottlenose dolphins in each
area reflect these differences in habitat structure and oceanography. Within the Southern
California Bight, between 289 and 356 dolphins (Dudzik 1999) range over an area of at least
830 km², demonstrating little site fidelity to any one area and exhibiting frequent movements
between different parts of their range (Defran and Weller 1999, Defran et al. 1999, Dudzik
1999). These frequent movements and the lack of site fidelity are thought to be an adaptation
to patchy and unpredictable distribution of prey, which necessitates that dolphins range
widely to locate prey resources (Defran et al. 1999). Relatively large group sizes, averaging
19 dolphins, are also thought to be an adaptation to patchily distributed prey, as larger groups
may be able to locate prey resources more efficiently (Norris and Dohl 1980, Weller 1991,
Marsh 2000). Although seasonal patterns were not evident in all parts of the dolphins’ range,
dolphins were more abundant during the Davidson (November through February) and
Upwelling (March through July) seasons over the long-term study in San Diego. This
seasonal flux may be due to the presence of temporally abundant prey resources, represented
by semi-annual aggregations of spawning squid within the study area, during the Davidson
season as well as increased levels of prey concentration around upwelling areas during spring
and early summer.
Dolphins within Sarasota Bay, however, exhibit markedly different behavioral
patterns. Approximately 100 dolphins have been identified as year-round residents within an
area of approximately 125 km² (Wells et al. 1980, Irvine et al. 1981, Wells 1986, Scott et al.
1990). Site fidelity of dolphins and the relatively small home-range dimensions of the
community indicates that prey species are consistently available across seasons and years.
School sizes are small, averaging 7 dolphins (Wells et al. 1987), which is considered a
reflection of feeding on solitary and non-obligate schooling prey (Wells 1991, Barros and
Wells 1998). Although dolphins remain in the area year-round, some seasonal shifts in
distribution within the study area occur, as dolphin abundance is highest in channels, passes,
and offshore areas during fall and winter and increases within the inner waters of the bays
during spring and summer (Scott et al. 1990). These seasonal shifts have been linked to
 64

changes in water temperatures, either through the thermal requirements of dolphins or as a

response to changes in prey and predator distribution (Irvine et al. 1981, Wells et al. 1980).
Given the similarity in habitat structure between the two areas, dolphin behavioral
patterns within Aransas Pass, Texas, were expected to parallel those in Sarasota Bay (Weller
1998). However, longitudinal research in the area indicated behavior was more similar to
that observed on the Pacific coast. Dolphins belonged to a large and open population, with
782 dolphins identified over a four-year period. While most dolphins demonstrated no
evidence of site fidelity to the area, a small proportion of the population appeared to be semi-
residential within the area. Short-term studies in both Aransas Pass and other areas along the
Texas coast have also identified small groups of resident or semi-resident dolphins (Shane
1977, Gruber 1981, Bräger 1992, Fertl 1994, Lynn 1995, Maze and Würsig 1999),
suggesting that the population of bottlenose dolphins off the Texas coast may be separated
into semi-residential subpopulations inhabiting bays and inner waters, and a larger group of
transient animals traveling extensive distances along the coast. Dolphins were present in the
Aransas Pass study area year-round, but abundance peaked in late fall and winter and
correspondingly decreased in spring and summer. Seasonal sighting patterns were also
evident among individual dolphins, with most identified during only one season.
Research along the Texas coast provided an interesting contrast with results generated
by work conducted in Sarasota Bay and along the Pacific coast (Weller 1998). Past studies
along the Pacific coast and in Sarasota Bay have presented what were generally considered to
be two extremes of intraspecific variation in bottlenose dolphin behavior. This variation has
been attributed in large part to differences in habitat structure between the two areas (Defran
et al. 1999). At one extreme, small groups of residential dolphins, such as those of Sarasota
Bay, are known to inhabit protected bays and inlets thought to contain relatively abundant
and spatially stable prey resources. At the other end of the continuum, larger populations of
essentially transient dolphins inhabit the more open and dynamic environment of the Pacific
coast, ranging widely in search of unpredictable and patchily distributed prey resources.
Research in Aransas Pass, however, illustrated the importance of other ecological variables in
the relationship between habitat and behavior. While habitat structure within Aransas Pass
most closely resembles that of Sarasota Bay, dolphin behavioral patterns were more similar
to those observed along the Pacific coast. Weller (1998) theorized that the more pronounced
 65

seasonal variability in water temperatures and prey availability along the Texas coast, which
resembles that of the Pacific coast, might account for similarities in dolphin movements
along the Texas and Pacific coastlines. Within the Texas coast population, however, a small
proportion of individuals do exhibit site fidelity characteristics more similar to dolphins of
Sarasota Bay. Residential or semi-residential dolphins are primarily found in protected bays
and inlets of the Texas coast, which closely resemble the bays and barrier islands of the
Sarasota study area. Thus, the variability of bottlenose dolphin behavioral patterns along the
Texas coast may reflect the effect of both large-scale (habitat structure) and small-scale
(temperature variability) ecological differences on behavior of bottlenose dolphins, as well as
illustrating the complexity of determining which environmental factors are important in
shaping behavior.
 66

CONCLUSIONS

The primary objectives of this study were to examine the occurrence patterns, site
fidelity, and movements of Pacific coast bottlenose dolphins over both space and time.
Bottlenose dolphins, which can be found in most of the world’s oceans and in a wide variety
of different habitats, have demonstrated a high degree of behavioral flexibility in response to
environmental variables (Shane et al. 1986). This study was able to examine Pacific coast
bottlenose dolphin behavior across three different spatial scales. At the finest scale,
distribution of sightings within the San Diego and Santa Barbara study areas were examined
for correlations with habitat characteristics. While some behavior patterns were stable at this
scale (e.g., dolphins always remained within 1 km of shore), other patterns, such as increased
sightings in canyon habitat of San Diego, provided evidence of differential use of habitats
within study areas. At the next level, behavioral patterns were compared between the San
Diego and Santa Barbara study areas between 1998 and 1999. While analysis of occurrence
patterns indicated that dolphins used these two areas with comparable intensities, dolphin
occurrence in the Santa Barbara study area was similar across oceanographic seasons, while
dolphins utilized the San Diego study area most heavily during the Davidson season. More
intense usage of the San Diego study area during this season suggested that dolphins were
exploiting a temporally abundant resource, in the form of aggregations of spawning squid
near Scripps Canyon. Comparison of site fidelity patterns indicated that dolphins were
sighted more often in San Diego than in Santa Barbara during the seventeen-month study,
and closer examination of sighting patterns indicated that increased levels of site fidelity in
San Diego might also have been correlated with seasonal usage patterns. Thus comparison
of occurrence and site fidelity patterns between areas indicated that dolphins do use portions
of their range differently, at least over time scales of 1-2 years, and suggested that
exploitation of temporally abundant resources may play an important role in shaping
occurrence patterns of Pacific coast bottlenose dolphins.
At the broadest spatial scale, continued and long-term study of bottlenose dolphins
along the Pacific coast allowed comparisons with longitudinal studies in other areas. Two
such areas, Sarasota Bay, Florida, and Aransas Pass, Texas, were chosen for comparison
 67

because of the large-scale differences in habitat structure and oceanography of both areas
when compared to the Southern California Bight. While Sarasota Bay and Aransas Pass are
similar in habitat structure, containing relatively protected inner waters sheltered from the
open ocean by barrier island chains, they differ oceanographically, with Aransas Pass
undergoing more intense seasonal fluctuations than those occurring in Sarasota Bay. The
habitat structure of the Southern California Bight, on the other hand, is quite different from
both Sarasota Bay and Aransas Pass, with coastal waters open and exposed to the full force
of the ocean. Although seasonal fluctuations in temperature and salinity are not as marked as
those found in Aransas Pass, waters of the Southern California Bight are highly dynamic and
undergo daily, seasonal, and annual fluctuations in response to changes in the California
Current system. In Sarasota Bay, where waters are protected and resources are considered to
be abundant and evenly distributed, dolphins are residential within a small area year-round.
In the more open, exposed, and spatially variable environment of the Southern California
Bight, however, dolphins travel extensive coastal differences and exhibit little site fidelity
within their range, presumably as an adaptation to widely distributed and unpredictable prey
resources. Dolphins in Aransas Pass seem to employ both behavioral strategies, with small
groups of semi-residential dolphins, similar to those in Sarasota Bay, inhabiting protected
bays while a larger subset of the population ranges widely and exhibits seasonal shifts in
distribution. Thus comparison of these three areas indicates that while habitat structure may
play a significant role in shaping behavioral patterns of bottlenose dolphins, variation in
oceanographic characteristics also has considerable influence on occurrence patterns, site
fidelity, and movements of dolphins.
Comparisons of behavioral patterns over time were also valuable in illuminating
environmental factors influencing dolphin behavior. Over the sixteen-year span of the study,
some patterns remained consistent over time, while others fluctuated on both a yearly and/or
seasonal basis. For example, over the course of the long-term study dolphins were most
often observed within 0.5 km of the coastline, and were always within 1 km of shore. This
pattern likely developed due to a combination of abundant nearshore prey resources and
limited predation pressure and interspecific competition. Within the range of Pacific coast
bottlenose dolphins, these factors are consistent over time and space and as such lead to the
development of stable behavioral strategies. The number of dolphins observed within the
 68

study area, however, varied with both season and year. Thus, while the environmental forces
shaping latitudinal distribution may remain constant over time, those factors influencing
alongshore distribution may be inconsistent over both short (within year) and long (between
years) time scales, making a behaviorally flexible strategy most effective in exploiting
resources within the 1 km² zone.
In summary, extension of photo-identification studies of Pacific coast bottlenose
dolphins over both time and space provided the opportunity to learn more about the forces
shaping behavior in this coastal predator. Understanding behavior and the factors
influencing it are important from a conservation and management viewpoint. While the
coastal bottlenose dolphin population in the Southern California Bight appears stable, other
populations (e.g. Moray Firth , Wilson et al. 1997b; mid-Atlantic seaboard of the United
States, Wang et al. 1994) might be considered threatened (Connor et al. 2000), and
knowledge of the factors influencing behavior in Pacific coast bottlenose dolphins may
provide valuable insight into risks facing other populations. As well, the coastal environment
of bottlenose dolphins is exposed to high levels of anthropogenic impacts, including boat
traffic, pollution, and habitat alteration. By providing information on baseline dolphin
behavior, long-term studies allow any changes in behavior, which could be a response to
such impacts, to be detected. The health of bottlenose dolphins, as top-level predators
within the coastal environment, may also provide a valuable indication of the overall health
of the habitat and ecosystem (Tyack et al. 2000).
 69

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 80

APPENDIX

PHOTOGRAPHIC QUALITY CRITERIA*

Negatives were be judged for quality based upon the following parameters:

1) Focus (Sharpness) - The focus of a photograph will be assessed using the

following 3-point scale:
3 = Blurry but general outline visible.
2 = Questionable whether small nicks can be detected. (slightly blurred
edge or “soft” edge)
1 = Adequate to detect small nicks. (sharp edge)

2) Exposure - The contrast of a photograph will be assessed using the following 3-

point scale:
3 = Fin and background (or glare) similar in shading so that notch
pattern is not easily distinguished.
2 = Fin somewhat lighter than background (or glare) but notch pattern
can still be distinguished. (For color: fin somewhat darker than
background.)
1 = Fin very light against a dark background with fin still lighter in
color than glare: notches are very easily distinguished. (For
color: fin very dark against light background.)

3) Proportion of Fin/ID area visible - The percentage of the dorsal fin region visible
in each negative will be assessed and labeled using the following 3 point
scale: Note: consult reference diagram for this measure.
3 = 1/3-2/3 of the fin is visible
2 = 2/3 or more of the fin is visible
1 = All or nearly all of the fin is visible (more than 2/3)

*
Adapted from Dudzik 1999
 81

4) The area that a dorsal fin occupies on a given negative will be assessed using
transparent grids designed specifically for this task. The following three-point
scale will be used for this measure:
4 = fin occupies box containing 4 squares
3 = fin occupies box containing 9 grid squares
2 = fin occupies box containing 16 squares
1 = fin larger than box containing 16 squares
If any portion of the fin is outside of the box, then designate the fin as
occupying the next larger box.
ABSTRACT
 ABSTRACT OF THE THESIS

Occurrence Patterns, Site Fidelity, and Movements

of Pacific Coast Bottlenose Dolphins
(Tursiops truncatus)
by
Aimée R. Lang
Master of Science in Interdisciplinary Studies: Animal Behavior
San Diego State University, Spring 2002

Boat-based photo-identification surveys conducted over a seventeen-month period off

San Diego (n=43) and Santa Barbara (n=61), California were used to study occurrence
patterns, site fidelity and movements of Pacific coast bottlenose dolphins (Tursiops
truncatus). Within the Santa Barbara study area, mean size of observed dolphin schools was
12.7, with 67.4% of surveys encountering dolphins and an average of 15.7 dolphins sighted
per survey. Although mean school size ( x =15.7), mean number of dolphins observed per
survey ( x =20.6), and percent of surveys encountering dolphins (82.1%) were higher in the
San Diego study area, these differences were not significant, indicating that dolphins used
both areas with similar intensities. When occurrence patterns were compared across
oceanographic seasons, only San Diego demonstrated evidence of seasonal patterns. Both
school sizes and number of dolphins observed per survey were significantly higher during the
Davidson oceanographic season than during all other seasons (P<0.01), which was attributed
in part to the presence of aggregations of spawning squid within the San Diego study area
between November 1998 and February 1999. Squid aggregations likely represented a
temporally abundant prey resource that was exploited either directly or indirectly by dolphins
during this time period.
Over the seventeen-month study, 204 dolphins were photographically identified in
San Diego, while 178 dolphins were identified in Santa Barbara. Dolphins demonstrated
little site fidelity in either area, with a mean of 2.1 sightings per individual in Santa Barbara
and a mean of 2.0 sightings per individual in San Diego. Eighty-nine dolphins were
photographed in both areas, with a total of 131 documented inter-study area movements. The
mean number of days between sightings in different areas was 96.0 days. One dolphin was
documented to move between areas over only five days, for a mean travel speed of 61.6
km/day.
Data collected in the San Diego study area during the seventeen-month study was
combined with that collected in San Diego waters by Defran and Weller between 1984 and
1989 and by Dudzik between 1997 and 1999 to examine the stability of occurrence patterns
and site fidelity of dolphins over time. A total of 241 surveys were conducted during the
combined study period and were used to compare dolphin occurrence patterns across years
and oceanographic seasons. While school sizes did not vary significantly across years, the
encounter rate (P =0.006) and the number of dolphins sighted per survey (P <0.001) were
significantly different across years, with a general trend of higher occurrence of schools
during the latter years of the study. Increased dolphin occurrence during the Davidson
oceanographic season was not as marked over the long-term dataset; instead, seasonality in
occurrence patterns was reflected in decreased encounter rates and number of dolphins per
survey during the Oceanic season. No effect of the 1987-88 El Niño or the stronger 1997-
1998 El Niño on dolphin occurrence patterns was detected.
Four hundred sixty-eight dolphins were photographed in San Diego between 1984
and 1999, and identified individuals were sighted a mean of five times. Of the subset of most
frequently sighted dolphins, the majority had been sighted in other study areas of the
Southern California Bight, providing further evidence that dolphins are highly mobile within
their range. As well, the majority of dolphins in this subset were photographed during all
three oceanographic seasons, indicating that the lack of observed site fidelity in Pacific coast
bottlenose dolphins was not correlated with seasonal movements within the range. The high
mobility and lack of site fidelity demonstrated by identified dolphins, in combination with
the observed variability in occurrence patterns over seasonal and annual time scales, were
considered a reflection of the highly dynamic nature of the coastal environment of the
Southern California Bight.