J. CETACEAN RES. MANAGE.

1(1):3-24

Population estimates of Pacific Coast bottlenose dolphins (Tursiops

truncatus) photographed off San Diego, California

Kimberly J. Dudzik, R. H. Defran

Contact e-mail: dudziktaz@att.net

Cetacean Behavior Laboratory

Department of Psychology
San Diego State University
San Diego, CA 92182-4611

ABSTRACT

A 29 month photo-identification study was conducted from 1996 to 1998 in the San Diego study

area to obtain new mark- recapture based population estimates for the Southern California Bight

population of coastal bottlenose dolphins (Tursiops truncatus). Chao’s closed model Mth was

applied to the current and earlier photographic data collected in this area from 1984 to 1989. These

earlier data were partitioned into 2.5 year sample periods (84/86, 87/89) comparable in duration to

the 96/98 sample period. A photo quality rating system was applied to photographic data from all

three sample periods to minimize heterogeneous capture probabilities. Abundance estimates for the

three sample periods were: 28984/86 (95% CI = 230 - 398), 35487/89 (95% CI = 330 - 390) and

35696/98 (95% CI = 306 - 437). The 1984-1986 sample period encountered fewer schools and

dolphins and had fewer resights and thus may have been negatively biased relative to other sample

periods. Abundance estimates derived for all sampling periods are probably lower than the true

population size because they exclude individuals among the 37% of this population that lack

identifying marks, and because the sighting frequencies vary among members of this population.

When considering the expected negative bias of the 84/86 estimate and the similarity of the 87/89

and 96/98 estimates, the size of population seems to have remained relatively stable across a 15 yr

time span. Occurrence patterns and school size characteristics from the 1996-1998 and the earlier

1984-1989 data sets were similar, providing further evidence of stability over time. No obvious

impacts from the 1997/1998 El Niño were detected in occurrence, school size, or population

estimates.

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Keywords: ABUNDANCE ESTIMATE; PACIFIC OCEAN; INDEX OF ABUNDANCE;

COMMON BOTTLENOSE DOLPHIN; MARK-RECAPTURE; PHOTO-ID

Submitted to Journal of Cetacean Research and Management – Please do not cite or circulate without authors’
permission.

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INTRODUCTION

Photo-identification research on Pacific coast bottlenose dolphins (Tursiops truncatus) since

1981 has shown that members of this population inhabit a narrow coastal corridor of over 900 km

from Ensenada, Baja California Norte, Mexico to Monterey Bay, California (Feinholz, 1996;

Defran et al., 1999). Within this range, members of this population are highly nomadic and make

frequent movements between coastal areas. A number of mark-recapture estimates of this

population have been derived using both photo-identification (Hansen, 1990; Defran and Weller,

1999) and aerial survey (Carretta et al. 1998) sampling techniques.

Hansen (1990) photographed coastal bottlenose dolphins occurring off northern San Diego

County from September 1981 to January 1983 and used Tanaka’s closed model to derive an

estimate of 240 individuals (95% CI = 120- 477). The broad confidence intervals associated with

this estimate diminishes its utility and are probably the result of low survey effort, the short interval

between a number of the surveys and the small size of the photographic data set (Cf. Hansen, 1990;

Defran and Weller, 1999).

Defran and Weller (1999) carried out photo-identification surveys in north San Diego County

from 1984 to 1989 and estimated population size using the Jolly-Seber technique. Population size

was estimated for each year possible between 1984 and 1989, and yielded estimates ranging from

234 (95% CI = 205-263) to 285 (95% CI = 265-306). Population size estimated by the Jolly-Seber

model, however, is likely to be biased downward due to violations of equal catchability that

assumes that every animal in the population has the same capture probability (Pollock et al., 1990).

When capture probabilities are not equal among all individuals, this estimate will be negatively

biased (Hammond, 1986; Pollock et al., 1990). Defran and Weller (1999) found that sighting

frequencies among individuals during the 84/89 study varied greatly from a single sighting to 24

sightings. Such heterogeneous sighting frequencies could be due to individual movement patterns,

individual distinctiveness, sampling effort and photographic quality or some combination of these

factors.

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In the current study we carried out new photo-identification surveys in the San Diego study area

in order to provide a more contemporary estimate for this population. Further, we attempted to

diminish negative bias associated with equal catchability violations by correcting for the effects of

lower photographic quality and by selecting a closed model estimator less affected by variations in

catchability. Closed population estimators operate under three assumptions that must be met in

order to obtain precise population estimates: 1) the population is closed to births, deaths,

immigration, and emigration; 2) each individual has the same probability of being captured; 3)

marks remain stable over time (Pollock et al., 1990). Each of these assumptions is examined in

detail below.

Closure
The first assumption requires that no births, deaths, immigration or emigration occur during the

study period. Bottlenose dolphins tend to be long lived animals with low mortality rates and low

birth rates (Wells and Scott, 1990). Choosing shorter sample periods, as in this study, minimizes

violations of this assumption since births and deaths occurring during the shorter periods would

have little effect on the estimate (Calambokidis et al., 1989; Calambokidis et al., 1993; Wilson et

al., 1999). Accordingly prior data collected in San Diego from 1984 to 1989, and in the current

study from 1996 to 1998 were partitioned into 2.5 y sample periods.

The second closed model assumption is that each individual in the population has the same

probability of being captured. When capture probabilities are heterogeneous among individuals in a

population, the resulting abundance estimates are negatively biased (Hammond 1986). Two sources

of heterogeneous capture probabilities in the current study were photographic quality and the

extensive geographic range of dolphins in this population.

Photographic Quality
Differences in photographic quality such as focus and exposure may have an effect on whether

some dolphins can be resighted and lead to heterogeneous capture probabilities (Hammond, 1986).

We minimized heterogeneity due to photographic quality by developing and applying a rating

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system that allowed us to eliminate low quality (difficult to match) photographs. (Arnbom, 1987;

Cerchio, 1998; Friday et al., 2000; Urban et al., 1999; Whitehead et al., 1997; Wilson et al., 1999).

Geographic range
Variability in the probability that a dolphin will visit some portion of the population range, such

as a more geographically limited study area, may diminish equal catchability and thus induce a

negative bias on the population estimate. Defran et al. (1999) and Feinholz (1996) reported that

bottlenose dolphins photographed along the central California and Southern California Bight

coastline utilize 905km range from Ensenada, Baja California Norte, Mexico to Monterey Bay, CA.

However, photographic sampling in Defran and Weller (1999) and in the current research only

occurred within a very small portion (32km) of the entire range. Between 88% and 94% of the

dolphins photographed off Santa Barbara, Orange County, and Ensenada from 1981 to 1989 were

also photographically captured in San Diego (Defran et al., 1999; Fig. 1). Similarly, 81% of

dolphins photographed in Monterey Bay between 1992 and 1994 were previously identified in San

Diego1. Furthermore, low resighting rates and regular movement patterns between these areas

showed limited site-fidelity to any particular area (Defran et al., 1999). Thus, the San Diego study

area, centrally located within this population’s range and visited by many members of this

population, offered an opportunity to sample individuals from across this region. However, the

high sighting variability of San Diego dolphins combined with their less than 100% photographic

overlap with dolphins from other Southern California Bight (SCB) study areas must be considered.

It seems likely, therefore, that some individuals may utilize the San Diego area less frequently than

others, or not at all, and that geographic bias may contribute to lower population estimates.

Mark recognition
The last closed model assumption requires that marks remain stable over time and individuals are

not mismatched. Bottlenose dolphins obtain notches along the trailing edge of the dorsal fin that

tend to remain stable over time, and a photographic record of unique notch patterns over time

1
Based on our reanalysis of the Feinholz (1996) photographic data.

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 J. CETACEAN RES. MANAGE. 1(1):3-24

allows the investigator to see subtle changes (Wursig and Jefferson, 1990). The San Diego

photographic catalog extends from 1981 to 1998 and has numerous examples of bottlenose dolphins

showing only subtle fin changes across 10 or more years.

Both open and closed population models share the equal catchability assumption, which is

difficult to meet in most population studies (Hammond, 1986; Pollock et al., 1990). Several closed

models have been developed, however, in order to allow for certain varying capture probabilities to

occur (Pollock et al., 1990). For the current estimates we selected Chao’s model Mth which

assumes that capture probabilities can change from one sampling period to the next (time variation),

and that each member of the population has a unique capture probability due to sex, age, home

range, etc., that is independent of all other members of the population (heterogeneity) (Chao and

Jeng, 1992; Otis et al., 1978).

METHODS

Study area
The San Diego study area is a 32km strip of coastline that extends from Scripps Pier, La Jolla

(320 52’N) to South Carlsbad State Beach (330 08’N; Fig. 1). The shoreline is characterized by

steep cliffs with intermittent sandy beaches. Dolphins in the area utilize a variable habitat that

consists of submerged reefs, rock outcrop, sloping sand and estuary mouths. Although much of the

study area is exposed to the full force of the open ocean, there are also areas bordered by dense

patches of kelp located about .5km offshore.

Photographic surveys
Survey procedures closely followed those of Defran and Weller (1999) and are briefly

summarized. Surveys were conducted aboard a 5.7m Chaparral boat equipped with a 115hp

outboard motor. The survey vessel was motored along the coastline, 90- 180m offshore of the surf

zone. At least three observers scanned the water’s surface from the beach to about 2km offshore.

Once dolphins were sighted, the boat proceeded approximately 1km past the school to ensure that

all members of the school were located. Information on location, time, sighting number, dolphin

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behavior, and field estimates of the number of adults and calves present were recorded before dorsal

fin photography began. School size and calf definitions were the same as in Defran and Weller

(1999).

The research vessel was positioned to within 3- 12m of the school in order to photograph

individual dorsal fins. All dorsal fin photographs were taken with a Canon AE-1, 35 mm camera

equipped with a Canon 400mm lens and high-speed motor drive (5 exposures/sec) and loaded with

Kodak Tri-X, 400 ASA black and white film. An attempt was made to photograph all dolphins

without regard to the apparent distinctiveness of their dorsal fin notch patterns. Once photographic

effort was considered complete, the boat was motored offshore where all film shot was labelled

with date, location, school number, and roll number. The research vessel then resumed travel along

the coastline to locate additional dolphin schools.

Photo-identification
Procedures for identifying and matching individual dorsal fins closely followed the methodology

of Defran et al. (1990) and are briefly summarized. A clear photograph of each distinctive dorsal

fin was chosen as a “type specimen” to which all other photographs were compared. Only

unambiguous matches to the “type specimen” were considered resightings. Further, all photographs

were rated by three experienced photo analysts on four criteria of photographic quality: focus (three

levels), exposure (three levels), proportion of the fin above the waterline (three levels), and size of

the fin within the frame (four levels). Photographs that had the lowest possible rating on any of

these four criteria were labelled ‘poor quality’ and were excluded from population analyses, while

the remaining photographs were labelled as ‘good quality’ (See Dudzik, 1999 for additional details

of the photo quality analysis).

Data analysis
Student t-tests were used to evaluate occurrence and school size differences. The program

CAPTURE (Rexstad and Burnham, 1991) was used for closed model analyses. CAPTURE features

several closed model estimators and Chao’s model Mth was determined to be the most appropriate

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model for the current study. CAPTURE includes estimates of abundance, point estimate averages,

standard errors and confidence intervals.

Power analysis was used to determine if the population estimates had enough power to detect a

trend over the 15 year time span. Power was calculated using Gerrodette’s (1987) general model:

r2n3>12cv2(z α/2 + zβ)2

where r is the rate of change over time, n is the number of estimates of abundance, cv is the

coefficient of variation of estimated total population size, z α/2 is the probability of making a Type I

error and zβ is the probability of making a Type II error.

Terminology
‘Photographic catalog’ refers to all individuals identified in coastal areas of the Southern

California Bight from 1981 to 1989 (cf. Defran and Weller, 1999), central California from 1990 to

1993 (Feinholz, 1996) and San Diego from 1996 to 1998 (current study). Two San Diego study

area photographic datasets are distinguished: ‘84/89’ – dolphins photographed by Defran and

Weller (1999) from 1984 to 1989; ‘96/98’ – dolphins photographed in the current study from 1996

to 1998. Finally, photographic data from the San Diego study area were also partitioned into three

sample periods: ‘84/86’ and ‘87/89 from Defran and Weller (1999); ‘96/98’ from the current study.

RESULTS

Survey effort, occurrence, school size

Sixty-six photographic surveys were conducted between March 1996 and August 1998 in the

San Diego study area. Dolphins were encountered on 79% (n = 52) of all surveys, and 1,497

dolphins (field estimate) were observed in 84 schools. Eleven percent of all dolphins encountered

were classified as calves. Although some surveys were cancelled due to unfavorable weather

conditions and equipment failures, survey effort was partitioned more or less equally across each

year with a small increase in survey effort in 1997 (1996 surveys = 19; 1997 surveys = 27; 1998

surveys = 20). Even though more surveys were conducted in 1997, considerably fewer dolphins

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were encountered (1996: n = 736; 1997: n = 239; 1998: n = 522). Comparisons of encounter rates,

number of schools and resighting rates between current and past research are summarized in Table

1.

In the current study overall mean school size was 17.8 (SD = 23.42) with yearly means of 25.4 in

1996 (SD = 33.31), 11.4 in 1997 (SD = 13.15) and 15.4 in 1998 (SD = 15.77). School sizes were

variable and ranged from a single dolphin to 140 dolphins. Comparisons of occurrence patterns

were made between the 96/98 data and the 84/89 data (Defran and Weller, 1999). No significant

difference was found between the percentage of surveys on which dolphins were encountered,

average school size, number of dolphins encountered per complete survey, and percentage of calves

encountered (Table 2).

Photographic dataset
A total of 233 dolphins were photographically identified in the current study. Sighting

frequencies were variable and ranged from 1 to 14 ( x = 2.6). Thirty-seven percent of identified

individuals were sighted only once and 56% (n = 131) of the individuals identified from 1996 to

1998 were resightings from 1984 to 1989.

In order to analyze the rate at which individuals were first identified, rate of discovery was

plotted as the cumulative number of individuals identified over blocks of five consecutive surveys

in which at least one dolphin was identified (Fig. 2). The rate of discovery curve shows a

continuous increase throughout the study period with one large increase between blocks two and

three. By the end of the study period, between 11% and 17% of all individuals were newly

identified.

Photographic quality and Population estimates

For each dataset, only those photographs meeting “good quality” rating criteria and those having

distinctive fins were included in the analysis of population size. After applying the photo quality

rating criteria to the 84/89 data, 73 poor quality photographs were removed from the catalog. This

removal resulted in a reduction of the number of individuals in the 84/89 data from 373 to 366

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individuals. The number of individuals that qualified as ‘good quality’ from the total in each

sample period were: 84/86: n = 160 out of 165 total individuals; 87/89: n = 284 out of 309; 96/98: n

= 225 out of 233. Abundance estimates derived by Chao’s closed model Mth were as follows:

84/86 - 289 (95% CI = 230 - 398), 87/89 - 354 (95% CI = 330-390), 96/98 - 356 (95% CI = 306 -

437). Power analysis indicated that the estimates had only moderate power to detect a trend (power

= .61 at α = .05).

DISCUSSION

Closed model population estimates based on the 96/98 photo-identification data collected in this

study, as well as those based on our selected subset of the 84/89 photographic data, were higher

than earlier estimates reported by Defran and Weller (1999) or Hansen (1990). These new

estimates, along with the recently collected occurrence and school size data, provided an

opportunity to examine trends in these parameters spanning a 15 y period. In addition, possible El

Niño effects on these population estimates were evaluated.

We reduced negative bias in our estimates by using Chao’s model Mth which allows for

variations in heterogeneity and time (see Introduction) and through our application of a

photographic quality rating system. The resulting estimates for our most recent sample periods

were the highest obtained and were quite similar (87/89est = 354, 96/98est = 356), while the estimate

for the earliest sample period (84/86est = 289) was somewhat lower. In our interpretation of these

values and in comparisons between them we reviewed possible sources of bias associated with the

assumption of equal catchability. Some of the bias we identified applies to all sample periods while

other sources relate to a specific sample period.

Although long-term photographic effort reveals that a high proportion of dolphins utilize the San

Diego study area as a part of their range (81% - 94% over 9yrs) choosing the short sampling periods

necessary for closed models probably introduced a degree of geographic bias. Some dolphins in the

population utilize the San Diego study area more often than others as evidenced by variable sighting

frequencies reported by Defran and Weller (1999) during their 84/89 study and reported in the

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current research. An additional negative bias in our population estimates occurs because not all

dolphins have distinctively marked dorsal fins. Approximately 37% of the dolphins encountered in

the San Diego study area, including the 11% calves, are not distinctly marked and are thus excluded

from the population estimate (Defran and Weller, 1999).

Although negatively biased, the point estimates we derived reveal a possible stable trend in the

population size (est. = 354-356) between the 87/89 and 96/98 sample periods. The 84/86 sampling

period was further biased due to fewer resightings as well as fewer schools and dolphins (Table 1).

When capture probabilities are low (low resighting rate), the estimate is biased downward (Chao

and Jeng, 1992; Hammond, 1986). Considering the low resighting rate for the 84/86 sampling

period, the estimate may actually have been more similar to the 87/89 and 96/98 estimates. If so,

this population of coastal bottlenose dolphins may have remained relatively stable over a 15 y

period. Although power analysis indicates only moderate power to detect a trend, this decrease in

power is most likely due to a small sample size of only two estimates (Gerrodette, 1987). Stability

in the population is also evidenced in the occurrence patterns compared over time (Table 2). One

would expect that if a major shift in population size had occurred, then a similar shift might be

reflected in school size, encounter rates, percentage of calves, or other parameters.

El Niño effects
The 1997/1998 El Niño event occurred during our 96/98 sample period. Negative impacts

associated with El Niño have affected several populations of marine organisms. For example,

California sea lion (Zalophus californianus) pup production decreased by 64% in 1998 due to the

97/98 El Niño (Forney, et al., 2000). Pinniped pups generally succumb to nutritional stress due to

strong storms separating mothers from pups, and reduced food availability that forces mothers to

remain at sea for longer periods instead of returning to feed their pups (Riedman, 1990).

Pacific coast bottlenose dolphins within the SCB have also been affected by El Niño in the past.

Following the major 1982/1983 El Niño event there was a northward range extension of SCB

bottlenose dolphins from Los Angeles to Monterey Bay (Wells et al., 1990). The 1997/98 El Niño

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event offered an opportunity to examine possible effects on population size as well as occurrence

patterns and school size. Statistical comparisons of these parameters (Table 2) between the 84/89

and 96/98 datasets showed comparable values and no significant differences. It appears, therefore,

that coastal bottlenose dolphins in the SCB escaped obvious negative effects of the 1997 El Niño.

We are cautious about these interpretations, however, as it is possible that some 1997/1998 El Niño

effects may have occurred too late to appear in the 96/98 sample period, or that they affected

parameters we did not evaluate.

In conclusion, the mark-recapture techniques and model used in this research helped to decrease

bias in population estimates in order to better understand the population characteristics of SCB

coastal dolphins. Variable sighting frequencies of individuals photographed in the San Diego study

area suggest that significant distributional shifts occur within this population and contribute to

geographic bias still evident in the estimates. A continued photographic effort in the secondary

study areas would help to further eliminate bias due to geographic distribution and, therefore,

strengthen population estimates. The population characteristics reported in this research, however,

provide a long-term view of trends within the SCB coastal population and the most current

abundance estimate.

ACKNOWLEDGMENTS

The authors would like to thank A. Lang, J. Marsh, T. Dedecker, E. Howarth, and J. Oswald

for their valuable assistance in the field. We are grateful to A. Lang, J. Marsh, and the many

Cetacean Behavior Laboratory interns who provided their time and assistance in the laboratory.

Appreciation goes out to K. Forney, J. Carretta, and J. Barlow of the National Marine Fisheries

Service, Southwest Fisheries Science Center, and to J. Calambokidis of the Cascadia Research

Collective who provided initial insights and comments about our research design and analysis.

Appreciation also goes to A. Acevedo-Gutierrez of the San Francisco California Academy of

Sciences for his assistance with the power analysis. We acknowledge the valuable and constructive

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comments offered by J. Barlow, D. Weller, and two anonymous reviewers on earlier drafts of the

manuscript.

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coast bottlenose dolphins (Tursiops truncatus) in the Southern California Bight. Mar.
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truncates). M.S. thesis, San Diego State University, San Diego, CA. 63pp.
Feinholz, D.M. 1996. Pacific coast bottlenose dolphins (Tursiops truncatus gilli) in Monterey Bay,
California. M.S. thesis, San Jose State University, San Jose, CA. 78 pp.
Forney, K.A., Barlow, J., Muto, M.M., Lowry, M., Baker, J., Cameron, G., Mobley, J., Stinchcomb,
C., and Carretta, J.V. 2000. U.S. Pacific Marine Mammal Stock Assessments: 2000. U.S.
Department of Commerce. 276 pp.
Friday, N., Smith, T.D., Stevick, P.T. and Allen, A. 2000. Measurement of photographic quality
and distinctiveness for the photographic identification of humpback whales, (Megaptera
noveangliae). Mar. Mamm. Sci. 16:355-374.
Gerrodette, T. 1987. A power analysis for detecting trends. Ecology 68:1364-1372.
Hammond, P.S. 1986. Estimating the size of naturally marked whale populations using capture-
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Hansen, L.J. 1990. California coastal bottlenose dolphins. Pages 403-420 in S. Leatherwood and
R. R. Reeves, eds. The Bottlenose Dolphin. Academic Press, San Diego, CA.
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Otis, D.L., Burnham, K.P., White, G.C., and Anderson, D.R. 1978. Statistical inference from
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Pollock, K.H., Nichols, J.D., Brownie, C. and Hines, J.E. 1990. Statistical inference for capture-
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Berkeley, CA.
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Table 1.
Summary of encounter, school, and resight statistics for each sample
period. “Number resighted” reflects the number of individuals resighted
within each study period.

Dolphins Number Number

Sample Period Encountered Schools Resighted

84/86 975 61 68

87/89 1894 85 250

96/98 1497 84 146

Table 2.
Summary of encounter, calf and school size statistics for 84/89 and 96/98 datasets.

Mean # of
Dolphins
% Surveys Encountered Mean
Dolphins Per Complete School
Dataset Encountered Survey % Calves Size

1984/1989 79% 26.8 11% 19.8

1996/1998 79% 21.1 11% 17.8

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Figure 1. San Diego study area. Inset shows location of other photo-
identification study areas along the California and Baja California
Norte coastline mentioned in the text.

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250
225
Cumulative Number of
Identified Dolphins

200
175
150
125
100
75
50
25
0
1 2 3 4 5 6 7 8 9 10
Blocks of Five Surveys

Figure 2. Rate of discovery for dolphins photographed in the San

Diego study area from 1996 to 1998. Only surveys on which at least
one identifiable dolphin was photographed were used. Block 10
represents the last four surveys.

18