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MARINE MAMMAL SCIENCE, 10(3):283-298 (July 1994).

0 1994 by the Society for Marine Mammalogy

CETACEANS OF THE NORTHERN GULF OF

CALIFORNIA: DISTRIBUTION,
OCCURRENCE, AND RELATIVE ABUNDANCE
GREGORY K. SILBER~
Institute of Marine Sciences, University of California, Santa Cruz, California 95064

MICHAEL W. NEWCOMER
22371 Hartman Drive, Los Altos, California 94024

PATRICIAC. SILBER
Catalyst Consulting Team, 2284 Dunster Lane, Potomac, Maryland 20854

HECTOR PEREZ-CORTeS M.*

A.P. 787, La Paz, Baja California Sur, 23000 Mexico

GRAEME M. ELLIS
Pacific Biological Station, Nanaimo, British Columbia, Canada V9R 5K6

A total of 1,7 15 km of boat-based surveys and 1,52 1 km of aircraft-based

surveys was conducted from 1986-1989 to assess the distribution, relative
abundance, and ecological relationships of cetaceans in the northern Gulf of
California. Seven cetacean species were seen; in decreasing frequency of groups
encountered they were: bottlenose dolphins, Tursiops truncatus; vaquitas, Pho-
coena sinus; common dolphins, Delphinus delphis; fin whales, Balaenoptera phy-
salus; Bryde’s whales, B. edeni; gray whales, Eschrichtius robustus, and killer
whales, Orcinus orca. Common dolphins were numerically dominant and bot-
tlenose dolphins were seen most often. Bryde’s whales and vaquitas had the
smallest group sizes. In general, the odontocete cetaceans were separated spatially,
whereas the distribution of Bryde’s and fin whales overlapped considerably.
Key words: cetaceans, Gulf of California, distribution, relative abundance, oc-
currence, ecology, group size.

The Gulf of California, Mexico, is an oceanographically unique basin pos-

sessing ecologically diverse provinces and great productivity (Alvarez-Borrego

* Present address: Marine Mammal Commission, 1825 Connecticut Avenue, N.W., Room 5 12,
Washington, DC 20009.
2 Present address: CRIP La Paz, Km 1 Carr a Pichilingue, La Paz, Baja California Sur, 23000
Mexico.

283
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284 MARINE MAMMAL SCIENCE,VOL. lO,NO.3, 1994

1983). The extreme upper Gulf is characterized by shallow depths (generally

less than 200 m), large seasonal temperature fluctuations (15”26’ C), great
tidal amplitudes (up to 9.6 m), and turbid, well-mixed water (Maluf 1.983).
Much has been written on the abundant and varied marine mammal fauna
occurring throughout the Gulf of California (for reviews, see Tershy et al. 1990,
Vidal et al. 1993). However, few studies have been conducted on cetaceans in
the upper Gulf. Moreover, there are no published reports involving extensive
surveys and no attempts to describe cetacean habitat use and distribution. in the
region. Several surveys were made but were limited to a few days or weeks (Van
Gelder 1960, Balcomb et al. 1979, Wells et al. 1981, Vidal et al. 119876,
Silber and Norris 1991, Barlow et al. 1993). Aside from these studies, all
published records of cetaceans in the upper Gulf consist of beach strandings or
opportunistic sightings from shore.
The primary objective of this study was to characterize habitat use and
distribution of the vaquita, Phocoena sinus. Secondary objectives were tcl char-
acterize the distribution, relative abundance, and ecological relationships of
cetaceans in the upper Gulf. To accomplish this, we conducted boat-based
surveys, augmented by aerial surveys, in the northern Gulf from 1986 to 1989
(Silber 1988, 1990a, b).

MATERIALS AND METHODS

In this paper the “northern” and “upper” Gulf of California refer to the
area north of the Midriff Islands. Although some surveys were conducted as far
south as 29”34’N, the study area was confined to the area north of 30”OO’N.
Logistical constraints, primarily the size and range of survey platforms used,
limited the area of more intensive survey.
Surveys were conducted in 1986 through 1989 in the northern Gulf over a
total of 3,236 km: 1,715 km from boats (Fig. 1) and 1,521 km from aircraft
(Fig. 2). Most surveys were conducted during spring when the best weath.er for
SUNey work was likely to be encountered. We also conducted a small number
of surveys in December 1988 and September 1989.
An 8-m Boston Whaler was used to survey for cetaceans north of 30”30’N
on a total of 7 1 d from 1986 to 1988 (Fig. 3). Two to four observers positioned
3.5 m above the water surface used the unaided eye and binoculars (7 X: and
10 x ) to search the area in front, and 200-300 m on either side, of the vessel’s
track, although observers also regularly scanned to greater distances. During all
surveys the vessel traveled at about 10-l 1 km/h. For each cetacean sighting,
species, group size, and location were determined.
Transects conducted in 1986 were nonrandom and restricted primarily to
nearshore areas. In subsequent years inshore surveys were augmented by randomly
allocated transect lines farther from shore. Searches for cetaceans were also
conducted when making passage to or from an anchorage and between transect
lines.
For each sighting, locations were determined by triangulation from landmarks,
and group sizes were estimated by agreement among all observers. Water depths
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SILBER ET AL.: GULF OF CALIFORNIA CETACEANS 285

Golfo
de
Californ

Figwe 1. Boat surveys (-) in 1986, 1987, and 1988.

were obtained with a JRC color video depth sounder or from a nautical chart.
Water clarity (as determined by using a secchi disc) and sea surface temperature
were measured at most sighting locations. Sea states were estimated according
to the Beaufort scale. No surveys were conducted when sea state exceeded
Beaufort 4, and the majority were conducted in sea states of one.
On four occasions surveys were conducted 4- 12 km into the Colorado River.
At these times we utilized strong flood tides to enter the narrow tidal channels
that meandered throughout the delta and then retreated on the ebb tides.
Surveys were also conducted from aircraft (Cessna 205) in the areas surveyed
by boat, as well as south to 2Y34’N (Fig. 2). A total of 1,521 km was flown
3-5 May 1988 and 2-6 September 1989. All surveys were flown at 152 m,
except one that was conducted at an altitude of 2 13 m. Three observers, each
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286 MARINE MAMMAL SCIENCE, VOL. 10, NO. 3, 1994

l,,
\
\\ il
L.-.--.-.,--.- 1
I
BAJA
- Aerial Survey
3-S my 1988

UNITED \
h.,, STATES 1 CALIFORNIA

Figwe 2. Aircraft surveys in May 1988 (-) and September 1989 (. . . . .).

familiar with both aerial survey techniques and P. sinus, were used on each
survey. Surveys were made with unaided eye, observers concentrating their efforts
within 200-500 m of the flight path, but they also scanned to greater distances.
Two observers were positioned on the port side of the aircraft and one on the
starboard. Untrained observers (i.e., pilots and other passengers) also searched
for cetaceans. For each sighting, species, group size (by observer consensus), and
location were determined; water depths were acquired later from a chart.
For boat- and aircraft-based surveys, we have presented sighting rates (number
of sightings/100 km surveyed, number of individuals/100 km surveyed), as
well as the estimated total number of individuals observed for each cetacean
species as indices of relative abundance.
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SILBER ET AL.: GULF OF CALIFORNIA CETACEANS 287

n 1986 n=26

sIn-
6
0 1987 n=23
q 1988 n=22
%
0)4
3
2
g,

0
1234123412341234
Feb. Mar. Apr. May
Weeks
Figure 3. Effort: time spent conducting boat-based surveys in the northern Gulf of
California (1986, 1987, and 1988).

RESULTS

Seven cetacean species were seen in the northern Gulf; in order of decreasing
frequency of groups seen they were: bottlenose dolphins, Tursiops truncatus;
vaquitas, P. sinus; common dolphins, Delphinus delphis; fm whales, Balaenoptera
physalus; Bryde’s whales, B. edeni; killer whales, Orcinus orca and gray whales,
Eschrichtius robustas (Table 1). Our analysis and discussion focuses primarily
on the three most commonly seen species-the common dolphin, bottlenose
dolphin, and vaquita.
The sighting rate was highest for the bottlenose dolphin, followed by the
vaquita and the common dolphin (Table 2). Common dolphins were numerically
dominant and were found in large schools containing up to 1,100 individuals.
The number of common dolphins was one order of magnitude greater than the
number of bottlenose dolphins, which was one order of magnitude greater than
the vaquita. Fin whales were the most commonly seen mysticete whale. Gray,
Bryde’s, and killer whales were least common, being seen on less than seven
occasions each. Unidentified balaenopterid whales were seen on three occasions.
In general, the odontocete cetaceans were separated spatially (Fig. 4). Bot-
tlenose dolphins showed little spatial overlap with common dolphins or with
vaquitas, but the distribution of vaquitas overlapped that of common dolphins;
the latter two species were occasionally seen in the same general vicinity (< 1.5
km apart). Habitats utilized by odontocetes in the upper Gulf of California can
be characterized roughly as follows: bottlenose dolphins were observed in highly
turbid, inshore waters, generally < 15 m deep, and within 10 km of shore, with
a distribution that favored the eastern perimeter (Fig. 4). Common dolphins
inhabited relatively clear, offshore waters, generally in excess of 30 m deep, and
> 15 km from shore, and most sightings occurred in the central portions of the
upper Gulf (Fig. 4). Vaquitas were seen in narrowly defined water depths,
generally between 10 and 30 m, around 15-20 km from shore, in waters that
varied in clarity (Table 1). Vaquitas also exhibited an areal distribution inter-
mediate between those of bottlenose and common dolphins.
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288 MARINE MAMMAL SCIENCE, VOL. 10, NO. 3, 1994

Table 1. Distance to shore, water depth, clarity, and temperature of cetacean sighting
locations in the northern Gulf of California.

Water
Distance Water temper- Water
to shore depth atnte clarity
Species (km) (4 (0 b-4
Tursiops truncatus N 140 140 38 23
f 5.4 12.4 19.0 2.6
SD 5.79 9.60 2.08 1.90
Range 0.125-34 2.5-46 15-21 0.5-6.5
Phocoena sinus N 58 58 32 32
32 18.4 26.6 21.5
SD 5.84 7.67 2.23 3%
Range 2.4-24 13.5-56.0 17-26 0.9--12.0
Delphinus delphis N 56 56 19 1.6
19.7 41.0 19.4 6.5
:D 10.98 24.39 2.00 2.10
Range 1.6-63.6 5-105 15-23 3.0-10.5
Balaenoptera physalus N 38 38 7
17.9 44.7 19.7 674
:D 11.30 22.92 3.25 3.34
Range 1.6-35.4 9-110 16-23 2.5-10.0
Balaenoptera edeni N 6 6 2
30.0 54.3 18.8
:D 11.05 33.16
Range 1 l-40 26-105 18.5-19.0
Orcinus orca N 2
3525 57.5
:D 2.;2 7.78
Range 34-37 52-63
Eschrichtius robustus N 2
3 2.5 45.5
SD 2.00 13.44
Range 15-18 36-55

Mean water depth and distance to shore were greatest for common dolphins,
least for bottlenose dolphins, and intermediate for vaquitas (Fig. 5, 6; Table
1). The range of water depths utilized by common dolphins was greatest (5-
105 m), while that of the vaquita was the most narrowly defined (14-56 m).
Mean water temperatures differed little at sighting locations for the three odon-
tocete species. Water temperatures were lowest at bottlenose dolphin sighting
locations and greatest at locations where vaquitas were seen (Table 1). In general,
water clarity was least where bottlenose dolphins were encountered and greatest
where common dolphins occurred, although the ranges of water clarity values
for each species were substantial (Table 1).
Although cetacean sightings during the September 1989 aerial surveys were
relatively few, the distribution and relative abundance of bottlenose dolphins,
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SILBER ET AL.: GULF OF CALIFORNIA CETACEANS 289

T&e 2. Group size and sighting rate of cetaceans in the northern Gulf of California.

Sighting Indi-
rate viduals
Group (#/ (#/
Species size 100 km) 100 km)
Tursiops truncatus N 140
10.1 boat 7.6 75.3
ZD 11.04 aircraft 0.6 8.2
Range l-60 combined 4.3 43.8
Total individuals 1,416
Phocoena sinus N 58
1.9 boat 2.7 5.0
:D 1.20 aircraft 0.8 1.6
Range l-7 combined 1.8 3.4
Total individuals 110
Delphinus delphis N 56
2 254.3 boat 2.6 596.0
SD 309.37 aircraft 0.8 264.2
Range 4-1,100 combined 1.7 440.0
Total individuals 14,239
Balaenoptera physalus N 38
f 5.7 boat 2.2 12.5
SD 10.48 aircraft 0.0 0.0
Range l-50 combined 1.2 6.6
Total individuals 215
Balaenoptera edeni N 6
4 1.2 boat 0.4 0.4
SD 0.41 aircraft 0.0 0.0
Range l-2 combined 0.2 0.2
Total individuals 7
Orcinus orca N 2
3 boat 0.6 0.1
SD $9 aircraft 0.7 1.0
Range 2-15 combined 0.1 0.5
Total individuals 17
Eschrichtius robustus N 2
f 1.5 boat 0.2
SD 0.71 aircraft :-A 0.0
Range l-2 combined 0:1 0.1
Total individuals 3

vaquitas, and fin whales appeared to be consistent with spring surveys. This
suggests that these species may be year-round residents, although bottlenose
dolphin distribution may have shifted slightly offshore. Bottlenose dolphins were
observed during every month surveyed. Common dolphins were seen over the
same periods, but were absent during September 1989. Bryde’s whales also
were not seen in September. This is the period of highest mean water temperature
in the upper Gulf (Robinson 1973), and the absence of common dolphins in
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290 MARINE MAMMAL SCIENCE. VOL. 10. NO. 3. 1994

ONORA

Golfo
de
California

Figwe 4. Sightings of bottlenose dolphins, Tursiops truncatus, (0); common dol-

phins, Delphinus delphis, (0); vaquitas, Phocoena sinus, (H); and killer whales, Orcinus
orca, (A) in the northern Gulf of California.

this period suggests that the species leaves the region to utilize areas of greater
productivity elsewhere.
Among the odontocete species, mean group size was greatest for common
dolphins (2 54.3 + 309.37 [SD)), although group size varied considerably for
this species, followed by that of bottlenose dolphins and vaquitas (10.1 f I! 1.04
{SD) and 1.9 + 1.20 [SD], respectively, Table 2). Vaquita group size was
always small; the largest group consisted of seven individuals.
Bottlenose dolphins were the only marine mammals seen in the Colorado
River. In the channels of the river, the dolphins were surrounded on virtually
all sides by land or mud flats and were never more than 200 m from shore.
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SILBER ET AL.: GULF OF CALIFORNIA CETACEANS 291

60

Depth (m)
Figwe 5. Frequency of occurrence of water depths at sighting locations for bottlenose
dolphins, vaquitas, and common dolphins.

The habitat consisted of shallow (2-m maximum at low tide), exceedingly silted
waters which had high tidal current velocities. The density of dolphins was high
within the river mouth, and density appeared higher there than most other areas
of the upper Gulf. Even at an estimated 12 km beyond the river mouth, the
density of bottlenose dolphins did not appear to diminish.

Delphinus

40 11-20 21-30 31-40 >40

Distance from shore (km)

Figwe 6. Frequency of occurrence of distance from nearest shore at sighting locations
for bottlenose dolphins, vaquitas, and common dolphins.
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292 MARINE MAMMAL SCIENCE, VOL. IO, NO. 3, 1994

‘\ i ,__..
‘\ I;.- ,_._.- ?- . 8. physalus
‘1. !
7 I 0 B. edeni
0 Eschrichtius
UNITED 1
‘-, STATES \ CAL

Figwe 7. Sightings of fin, Balaenoptera physalus, (U); Bryde’s, Balaenoptera edeni,

(0); and gray, Eschrichtius robustus, (17) whales in the northern Gulf of California.

The number of sightings of Bryde’s whales was small, therefore our statements
with respect to that species should be regarded as preliminary. Nonetheless, in
contrast to the odontocete cetaceans, the spatial distribution of Bryde’s and fin
whales overlapped considerably (Fig. 7), as did the range of water depths (Fig.
8) and distances from shore where these whales occurred (Fig. 9). However,
mean water depth and mean distance to the nearest shore (54.3 -C 33.16 {SD]
and 30.0 + 11.5 [SD), respectively) were greater at locations in which Bryde’s
whale sightings occurred than were mean water depth and distance from nearest
shore (44.7 rt 22.92 [SD] and 17.9 f 11.30 {SD}, respectively) at fin whale
sighting locations. The range of water depths where fin whale sightings occurred
(9- 110 m) was greater than the range of water depths for Bryde’s whale sightings
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SILBER ET AL.: GULF OF CALIFORNIA CETACEANS 297

Depth (ml
Figwe 8. Frequency of occurrence of water depths at sighting locations for Bryde’s
and fin whales.

locations (26-105 m), although for Bryde’s whales this may be an artifact of
small sample size. Mean group size for fin whales was 5.7 ? 10.48 (SD), but
loose aggregations of up to 50 individuals were seen. Bryde’s whales were always
seen singly or in pairs.
Gray whales and killer whales were seen only a few times and appear to be
infrequent visitors to the northern Gulf. These species were seen in moderate
water depths (36-63 m) and more than 15 km from shore. Gray whales were
observed in small groups and, judging by their body size, were probably sub-
adults.

DISCUSSION

Our cetacean surveys have yielded results similar to those of earlier surveys
within the upper Gulf. Cetacean species commonly sighted included bottlenose
dolphins, common dolphins, and fin whales (Balcomb et al. 1979, Wells et al.
1981, Vidal et al. 1987a). Van Gelder (1960) reported only fin whales, and
a total of four sightings of vaquitas were reported by Wells et al. (198 l), Vidal
et al. (1987a), and Barlow et al. (1993).
The following cetaceans reported north of Angel de La Guarda Island (central
Gulf) probably occur only infrequently in the northernmost Gulf. Gilmore
(1957) reported beach strandings of sperm whales, Physeter macrocephalus, in
the northern Gulf. A beach-cast pygmy sperm whale, Kogia breviceps, was found
north of San Felipe by Brownell (1969), and a live stranding of this species
occurred at Puerto Penasco in 1984 (Vidal et al. 19876). Sightings of humpback
whales, Megaptera novaeangliae, near El Golfo de Santa Clara and Bahia San
Luis Gonzaga were described by Villa-R. (1986), and Heyning (1986) reported
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294 MARINE MAMMAL SCIENCE. VOL. 10. NO. 3. 1994

g -
50

59
n B.edeni
40
0 B. physhfs

80 30

L
8 t

11-20 21-30 31-40 >40

Distance from shore (km)

Figwe 9. Frequency of occurrence of distance from nearest shore at sighting locations
for Bryde’s and fin whales.

a single beach-cast specimen of a rough-toothed dolphin, Steno bredanensis, near

El Golfo de Santa Clara. Two sightings of pilot whales, Globicephalus macro-
rhynchus, were made just north of Angel de La Guarda Island (Balcomb et al.
1979).
Tershy and Breese (199 1) reported a total of 3 1 gray whale sightings between
1979 and 1989 in the Gulf, north of 28”00’N, with some whales observed
feeding. They suggested that some gray whales visit the northern Gulf to exploit
its rich patches of prey. These sightings as well as others reported for the Gulf
(Wells et al. 1981, Vidal et al. 1987a, Silber 19906) consisted of small groups,
generally one or two individuals.
Killer whales have been reported in various locations throughout the Gulf,
including the southern (Tarpy 1979), and mid-Gulf regions (Vidal and Pechter
1989, Breese and Tershy 1993, Vidal et al. 1993). One of our killer whale
sightings has been described previously; it involved an attack by killer whales
on a Bryde’s whale (Silber et al. 1990). We believe these to be the only published
sightings of killer whales in this region.
A significant amount of work has been conducted on the relative abundance,
seasonal occupancy, habitat utilization, and social behavior of balaenopterid
whales in the central Gulf (Tershy et al. 1990, 1991; Tershy 1992; Breese and
Tershy 1993). With minor exceptions, there is general agreement between our
findings on Bryde’s and fin whales and work conducted on these species in the
central Gulf. Some individual fin whales were common to both regions (as
confirmed by photoidentification), as well as other areas in the Gulf (Tershy et
al. 1990>, suggesting that portions of the fin whale population utilize the entire
Gulf.
Our findings on Bryde’s and fin whale group sizes are similar to those reported
by other researchers for the upper Gulf (Balcomb et al. 1979, Wells et al.
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SILBER ET AL.: GULF OF CALIFORNIA CETACEANS 295

1981, Vidal et al. 1987a) and elsewhere in the Gulf (Tershy 1992, Breese and
Tershy 1993, Vidal et al. 1993). Tershy (1992) and Breese and Tershy (1993)
indicated that Bryde’s whales frequently were seen feeding alone or gathered in
small feeding aggregations (median group size = 3.0), whereas fin whales were
seen feeding in larger groups (median = 5.0); the latter species often formed
feeding aggregations consisting of coordinated groups of two to four individuals.
Delphinid group sizes in the upper Gulf were similar to those reported for
other locations. In general, dolphin species which utilize inshore habitats occur
in smaller group sizes than those species in pelagic habitats (for review see Wells
et al. 1980, Wiirsig 1986), and we observed this pattern among bottlenose
and common dolphins. Common dolphin groups in the upper Gulf were typically
large, and the mean group size was comparable to sizes reported in other regions
of the North Pacific (Evans 1975, Hui 1979, Smith et al. 1986) and for the
Gulf of California (Wells et aZ. 198 1, Breese and Tershy 1993). Very large
schools (up to several thousand individuals) of common dolphins have been
noted occasionally elsewhere, and group sizes varied seasonally in this species
(Hui 1979, Smith etal. 1986). Schools exceeding an estimated 1,100 individuals
were not seen in the northern Gulf. Smaller group sizes in this region, relative
to those found in open ocean habitats, may be related to differences in spatial/
temporal distribution of prey between the semienclosed upper Gulf and more
pelagic environments. Mean group size observed in bottlenose dolphins in the
upper Gulf of California was within the range of group sizes reported for this
species in other locations in the Atlantic and Pacific Oceans (for review, see
Shane et al. 1986). Vaquita group sizes were always small and were similar to
those previously reported for this species (Wells et al. 198 1, Vidal et al. 1987a).
The closely related harbor porpoise, P. phocoena, occurs in similarly small group
sizes in the North Pacific and North Atlantic and is commonly seen in pairs or
small groups of up to 10 individuals (Gaskin et al. 1974, Leatherwood et al.
1982).
Odontocete species exhibited little spatial overlap, whereas the distribution
of Bryde’s and fin whales overlapped. The use of different habitats (i.e., water
depths and distances to shore) by odontocete species in the upper Gulf may
contribute to their ecological separation. Two studies, conducted in different
oceans, described spatial separation among odontocete species and provide note-
worthy similarities to our study. Saayman et al. (1972) and Saayman and Tayler
(1973) described the distribution of Sousa sp., Stenella coeruleoalba, and Tursiops
sp. off the coast of South Africa, where these species occupied inshore, pelagic,
and intermediate habitats, respectively. An analogous situation, involving T.
truncatus (inshore), Lagenorhynchus obscurus (pelagic), and Phocoena spinipinnis
(intermediate), was described for the coastal waters of Argentina (Wiirsig 1979;
Wiirsig and Wiirsig 1979, 1980).

ACKNOWLEDGMENTS

This study would not have been possible without important field contributions by D.
Breese, T. Jefferson, A. Robles, G. Barros, L. Torrez-M., and A. Valezquez-R. We extend
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296 MARINE MAMMAL SCIENCE,VOL. 10, NO. 3, 1994

special thanks for support to the Center for the Study of Deserts and Oceans, Project
Lighthawk, and the West Coast Whale Research Foundation. Drs. K. Norris, El. Villa-
R., and R. Wells provided invaluable advice. The project received financial support from
the Nature Conservancy, the Center for Marine Conservation, the Homeland Foundation,
and the American Cetacean Society (Los Angeles Chapter). The study was conducted
under Scientific Research Permit numbers 301856, 300422, 400036 issued by the
Secretaria de Pesca, Mexico. The paper was improved by comments from B. Tershy.

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SILBER ET AL.: GULF OF CALIFORNIA CETACEANS 297

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Received: 26 February 1993
Accepted: 11 March 1994