Defran, Caldwell, Lang and Weller

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PHOTOGRAPHIC EVIDENCE FOR GEOGRAPHIC BOUNDARIES IN
BOTTLENOSE DOLPHINS (TURSIOPS TRUNCATUS) ALONG THE PACIFIC
COASTS OF CALIFORNIA AND BAJA CALIFORNIA NORTE

Bottlenose dolphins (Tursiops truncatus) off the Pacific coast of California and Baja

California Norte are highly nomadic, regularly ranging over extensive distances (Wells et

al. 1990, Defran et al. 1999). Recent investigations have shown that a majority of

dolphins in this coastal population utilize an area extending from at least Ensenada, Baja

California Norte in the south, to Monterey Bay, California in the north (Feinholz 1996,

Defran and Weller 1999, Defran et al. 1999). Historical records suggest this species once

occurred as far north as San Francisco Bay, California (Orr 1963); however, more

contemporary accounts have considered southern Los Angeles County, with occasional

sightings extending as far north as Point Dume, California (Dohl et al. 1981), to be the

typical northern distribution limit (Norris and Prescott 1961, Leatherwood and Reeves

1982, Hansen 1990). Following the 1982-1983 El Niño event, bottlenose dolphins

extended their northern distribution to Monterey Bay, and perhaps as far as San Francisco

(Wells et al. 1990, Feinholz 1996). This range extension appears permanent, as dolphins

continue to regularly occur off central California1 and occasional sightings further north

suggest that the distribution of this population may have returned to its known historical

limit (Ferrero and Tsunoda 1989, Feinholz 1996, Wells and Scott 1999).

While the northern range limit for bottlenose dolphins along the California coast may

be determined by cold water tolerance limits, such a temperature barrier does not

1Personal Communication from Lisa Giesick and Tom Kieckhefer, Pacific Cetacean Group, 3239 Imjin
Road #122, Marina, CA 93933, January 2000.
 Defran, Caldwell, Lang and Weller
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occur to the south until the presumed southern extent of bottlenose dolphin distribution

off central Chile (Jefferson et al. 1993, Wells and Scott 1999). Research by Curry (1997)

found no genetic differences between bottlenose dolphins off California and South

America, indicating that some level of population mixing was probably occurring along

the entire range of the species. Population distinctions within this distribution, however,

remains poorly understood.

To better determine the southern movement patterns for dolphins photographically

identified in several study sites off southern and central California (Fig. 1), a series of

photo-identification surveys were conducted off Ensenada, Baja California Norte (Defran

et al. 1999). A high percentage (88%) of 68 dolphins identified off Ensenada (Table 1)

were matched to a catalog of 516 known individuals identified during a long-term study

(1981-1998) off San Diego (Hansen 1990, Weller 1991, Defran and Weller 1999, Dudzik

1999). In addition, a high number of dolphins sighted in Ensenada were also sighted in

more northern study sites off Orange County, Santa Barbara, and Monterey Bay

(Feinholz 1996, Defran et al. 1999). These results suggested that if a southern range

boundary for coastal bottlenose dolphins off California existed, it was likely to be south

of Ensenada.

The research presented here was conducted in the coastal waters 32 km south of San

Quintín, Baja California Norte (30o 18’ N, 115o 50’ W) between April and August 1990.

Our San Quintín study area was located approximately 376 km south of San Diego and

200 km south of Ensenada. The study area consisted of a 38 km coastal region from

Punta Azufre, near the entrance to San Quintín Bay, to Cañon del Rosario, and extended

2 km offshore (Fig. 1). Survey and photo-identification procedures were identical to

 Defran, Caldwell, Lang and Weller
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those employed during similar studies off California and Baja California Norte, and are

detailed elsewhere (Defran et al. 1990, Weller 1991, Defran and Weller 1999).

A total of eight boat-based photo-identification surveys were conducted in the San

Quintín study area during three sampling periods: April 9 to 12 (n = 2); June 13 to 16 (n

= 3); and August 8 to 11 (n = 3). During the first survey on 9 April 1990, the research

boat was launched at Punta Azufre, and the survey track extended south to El Socorro

(Fig. 1). For all subsequent surveys, the boat was launched from the beach off our base

camp, and surveyed south to Cañon del Rosario. Two hundred bottlenose dolphins in 14

schools were observed during the study. The average school size was 14.3 dolphins (SD

= 7.53), and ranged from 5 - 30 individuals. A total of 105 individual dolphins were

identified, and 38% (n = 40) of these dolphins were photographed on two or more

surveys. In most cases, resightings of an individual were obtained during a single

sampling period (i.e., within days), reflecting short-term site fidelity similar to that

displayed by dolphins off more northern study sites (Defran and Weller 1999, Defran et

al. 1999).

Dorsal fin photographs collected during each survey in San Quintín were compared

to an existing catalog of 545 dolphins identified in southern and central California and off

Ensenada from 1981 to 1998 (Hansen 1990, Weller 1991, Feinholz 1996, Defran and

Weller 1999, Defran et al. 1999, Dudzik 1999). In contrast to the high levels of

photographic match-up reported for dolphins identified in study sites between Ensenada

and Monterey Bay (Table 1), only two (2%) of the 105 dolphins identified in San Quintín

were previously or subsequently identified in any northern study site. Dolphin 006, first

photographed in 1982 off San Diego by Hansen (1990), was also photographed twice in
 Defran, Caldwell, Lang and Weller
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Ensenada and five times in San Diego between 1982-1989. This dolphin was then

photographed twice in San Quintín during August 1990, twice in San Diego during

February 1991, and eight times in San Diego between March 1996 and August 1998.

Dolphin 739, first photographed in San Quintín in June 1990, was subsequently

photographed in the same school as dolphin 006 off San Diego on 14 August 1998.

Several aspects of the present study may have contributed to the low level of

photographic mixing observed between coastal locations. For example, the limited

number of surveys (n = 8) conducted in San Quintín may not have been adequate to

document photographic matches between regions. However, earlier studies in Baja

California Norte and in Southern California (Defran and Weller 1999, Defran et al. 1999)

indicated that the San Quintín sample size was sufficient to detect photographic matches

between coastal locations. For example, a relatively low number of surveys were also

conducted off Santa Barbara (n = 12) and Ensenada (n = 11). When photographs from

each of these study sites were compared to a catalog of 516 dolphins identified off San

Diego between 1981-1998 (Caldwell 1992, Defran and Weller 1999, Dudzik 1999),

resighting rates of 88% (Ensenada) and 92% (Santa Barbara) were documented (Table 1).

Further, more dolphins were identified in San Quintín (n = 105) than in either Santa

Barbara (n = 49) or Ensenada (n = 68) (Table 1), making the probability of finding an

inter-study area match more likely.

The spring-summer seasonal sampling bias off San Quintín was also unlikely to have

had a significant influence on the low photographic overlap detected between coastal

study areas, because the occurrence and distribution patterns for dolphins off San Diego,

our most intensively sampled study area (Table 1), were not seasonally-based (Defran
 Defran, Caldwell, Lang and Weller
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and Weller 1999). Therefore, we conclude that the photo-identification techniques

employed during the present study were sufficient to detect greater inter-study area

overlap had it existed.

It is also improbable that the 376 km distance between the northern San Quintín and

southern San Diego study area borders represented a significant barrier. Coastal

movements of 450 km were commonly observed for dolphins off Baja California Norte

and California (Defran et al. 1999), and travel distances in excess of 900 km were

recorded for dolphins moving between Ensenada and Monterey Bay (Feinholz 1996).

The findings presented here, in combination with results from previous longitudinal

research on Pacific coast bottlenose dolphins (Defran and Weller 1999, Defran et al.

1999), suggest that at least two Tursiops populations may occur off the coastline of

California and Baja California Norte. We hypothesize that the coastal region between

Ensenada and San Quintín functions as the southern range boundary for most members of

a “California” population of bottlenose dolphins that range between Ensenada and

Monterey Bay, and as the northern range boundary for a “Baja California Norte”

population.

Range boundaries typically occur where there are breaks in resources, where

environmental conditions become inhospitable, or where geographic features deter

movement (Mayr 1970). Our review of the literature regarding the ecology,

oceanography, and marine geology of the eastern Pacific seaboard between San Quintín

and Ensenada2 (see also Caldwell 1992) revealed no apparent explanation for the range

2Personal Communication from Eduardo Morteo, Ciencias Marinas, Universidad Autónoma de Baja
California, Km 103 Carr. Tijuana-Ensenada, 22800 Ensenada, Baja California Norte, January 2000.
 Defran, Caldwell, Lang and Weller
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boundary proposed here. The determination of two potentially discrete Tursiops

populations along the Pacific coast of California and Baja California Norte provides the

basis for better defining management and conservation efforts regarding this delphinid

species, and functions as an important reference point in continued basic science studies

of bottlenose dolphin behavioral ecology. Future research programs incorporating

genetic comparisons along the California and Baja California Norte coast, and boat-based

photo-identification surveys between Ensenada and San Quintín may help to better

clarify the apparent population distinctions reported here.

ACKNOWLEDGEMENTS

The authors extend their sincere gratitude to the many individuals who made

important contributions to the field, laboratory, and manuscript phases of this research.

A. Sandoval and D. Holmgren of Centro De Investigación Científica Y De Educación

Superior De Ensenada, Ensenada, Baja California Norte, and J. Bullis and A. French of

San Diego State University provided essential logistical support in the field. A. Bassett,

L. Barre, and numerous members of the Cetacean Behavior Laboratory maintained and

updated our photographic catalogs and associated databases. E. Morteo critically

reviewed Spanish language publications on the marine biology of the Southern California

Bight and San Quintín regions and J. Barlow made valuable suggestions about

terminology. This study was conducted under the provisions of a research permit issued

by the Secretaría de Medio Ambiente Recursos Naturales Y Pesca to Dr. G. Hamman,

Faculdad de Ciencias Marinas, Universidad Autónoma de Baja California, Km 103 Carr.

Tijuana-Ensenada, 22800 Ensenada, Baja California Norte.

 Defran, Caldwell, Lang and Weller
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LITERATURE CITED

Caldwell, M. 1992. A comparison of bottlenose dolphins identified in San Quintín and

the Southern California Bight. M.S. thesis, San Diego State University, San Diego,

CA. 59 pp.

Curry, B. 1997. Phylogenetic relationships among bottlenose dolphins (Genus Tursiops)

in a worldwide context. Ph.D. dissertation, Texas A&M University, College Station,

TX. 138 pp.

Defran, R. H., G. M. Shultz and D. W. Weller. 1990. A technique for the photographic

identification and cataloging of dorsal fins of the bottlenose dolphin (Tursiops

truncatus). Report of the International Whaling Commission (Special Issue 12):53-

55.

Defran, R. H., and D. W. Weller. 1999. The occurrence, distribution, site fidelity and

school size of bottlenose dolphins (Tursiops truncatus) off San Diego, California.

Marine Mammal Science 15:366-380.

Defran, R. H., D. W. Weller, D. L. Kelly and M. A. Espinosa. 1999. Range

characteristics of Pacific coast bottlenose dolphins in the Southern California Bight.

Marine Mammal Science 15:381-393.

Dohl, T. P., K. S. Norris, R. C. Guess, J. D. Bryant and M. W. Honig. 1981. Summary

of marine mammal and seabird surveys of the Southern California Bight area, 1975-

1978. Volume III. Investigator’s Reports. Part II. Cetacea of the Southern California

Bight. National Technological Information Service. 434 pp. Available from

National Technical Information Service, Technology Administration, U.S.

Department of Commerce, Springfield, VA 22161.

 Defran, Caldwell, Lang and Weller
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Dudzik, K. J. 1999. Population dynamics of the Pacific coast bottlenose dolphins

(Tursiops truncatus). M.S. thesis, San Diego State University, San Diego, CA. 63

pp.

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.

Ferrero, R. C., and L. M. Tsunoda. 1989. First record of a bottlenose dolphin (Tursiops

truncatus) in Washington State. Marine Mammal Science 5:302-305.

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.

Jefferson, T. A., S. Leatherwood and M. A. Webber. 1993. Marine mammals of the

world. FAO species identification guide. FAO, Rome, 320 pp.

Leatherwood, S., and R. R. Reeves. 1982. Bottlenose dolphin (Tursiops truncatus) and

other toothed cetaceans. Pages 369-414 in J. A. Chapman and G. A. Feldhammer,

eds. Wild mammals of North America: Biology, management, economics. The

John Hopkins University Press, Baltimore, MD.

Mayr, E. 1970. Population, species, and evolution: an abridgment of animal species and

evolution. Belknap Press of Harvard University Press, Cambridge, MA.

Norris, K. S., and J. H. Prescott. 1961. Observations on Pacific cetaceans of California

and Mexican waters. University of California Publications of Zoology 63:291-402.

 Defran, Caldwell, Lang and Weller
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Orr, R. T. 1963. A northern record of the Pacific bottlenose dolphin. Journal of

Mammalogy 44:424.

Weller, D. W. 1991. The social ecology of Pacific coast bottlenose dolphins. M.S.

thesis, San Diego State University, San Diego, CA. 93 pp.

Wells, R. S., and M. D. Scott. 1999. Bottlenose dolphin Tursiops truncatus (Montagu,

1821). Pages 137-182 in S. H. Ridgway and R. Harrison, eds. Handbook of marine

mammals. Volume 6. The second book of dolphins and the porpoises. Academic

Press, San Diego, CA.

Wells, R. S., L. J. Hansen, A. Baldridge, T. Dohl, D. L. Kelly and R. H. Defran. 1990.

Northward extension of the range of bottlenose dolphins along the California coast.

Pages 421-431 in S. Leatherwood and R. R. Reeves, eds. The bottlenose dolphin.

Academic Press, San Diego, CA.

 Defran, Caldwell, Lang and Weller
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R. H. Defran3, Marthajane Caldwell 4, Aimée R. Lang, and David W. Weller5, Cetacean

Behavior Laboratory, Department of Psychology, San Diego State University, San Diego,

California 92182-4611, U.S.A.; e-mail: rdefran@sunstroke.sdsu.edu.

3 Author to whom correspondence should be addressed.

4 Current Address: Department of Biology, University of Miami, P.O. Box 249118, Coral Gables, FL
33124, U.S.A
5 Current Address: Southwest Fisheries Science Center, National Marine Fisheries Service, NOAA, P.O.
Box 271, La Jolla, CA 92038-0271, U.S.A.
 Defran, Caldwell, Lang and Weller
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Table 1. Summary information on survey effort, study period, and photographic

data for San Quintín and other Pacific coast study areas.
Number of Number
Study Number of Study dolphins resighted to
area surveys period identified San Diego
San Diego 245 1981 - 1991, 1996 - 1998 516 -------
Orange County 44 1982 - 1989 133 123 (92%)
Ensenada 11 1985 - 1986 68 60 (88%)
Santa Barbara 12 1987 & 1989 49 45 (92%)
Monterey Baya 52 1990 - 1993 58 47 (81%)
San Quintín 8 1990 105 2 (2%)
a
Reflects our reanalysis of the Feinholz (1996) photographic data.
 Defran, Caldwell, Lang and Weller
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FIGURE CAPTION

Figure 1. San Quintín study area. Inset shows Pacific coast study area locations
mentioned in the text.
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