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in the western North Atlantic. Pages 166-17 1 in A. C. Pierrot-Balts, S. van der

Spoel, B. J. Zahuranec and R. K. Johnson, eds. Pelagic biogeography. UNESCO
Technical Papers in Marine Science 49, Paris.
S. K., H. P. 198 1. Identifying humpback whales using
their natural markings. Polar Record 20:439-444.
S. K., P. J. S. S. D. 1980. Humpback
whales: a catalogue of individuals identified in the western North Atlantic by means
of fluke photographs. College of the Atlantic, Bar Harbor, ME 04609. 169 pp.
D. K., P. J. S. K. G. S. 1989. Population
composition of humpback whales, novaeangliae, on Silver Bank, 1984.
Canadian Journal of Zoology 67:281-285.
E. W., W. L. 195 3. Preliminary report on the mammary
glands of whales. Norsk Hvalfangst-tidende 42:249-258.
H. 1982. Populations of humpback whales in the Northwest Atlantic.
Report of the International Whaling Commission 32:345-353.
H. E., R. K. A. G. 1975. Population estimates of the
humpback whale (Megaptera in the West Indies by visual and acoustic
techniques. Journal of the Fisheries Research Board of Canada 32:499-506.

Cetacean Research Program,

Center for Coastal Studies, P.O. Box
1036, Provincetown, Massachusetts T.
02657; Cetacean
Research Unit, P.O. Box 159, Gloucester, Massachusetts 01930. Received
December 17, 1992. Accepted May 28, 1993.

MARINE MAMMAL SCIENCE, 9(4):434-438 (October 1993)

0 1993 by the Society for Marine Mammalogy

DIURNAL AND SEASONAL BEHAVIOR

PATTERNS OF BOTTLENOSE DOLPHINS
(TURSIOPSTRUNCATUS)

Most animals show daily and seasonal patterns in their behavior. However,
behavior patterns of free-ranging cetaceans have rarely been described. This note
analyzes the diurnal and seasonal (summer and fall) occurrence of three common
activities, composed of suites of individual behaviors-foraging, traveling, and
socializing-in bottlenose dolphins.
From 1 June to 30 November 199 1, I recorded the behavior of bottlenose
dolphin groups during 97 trips (27 to 6 I trips per month; 3 10.8 h) in the bay
system of Galveston, Texas, and adjacent waters of the Gulf of Mexico, from
a 6.5-m research vessel. This was part of an ongoing photoidentification study
of the Texas A&M University Marine Mammal Research Program.
Bottlenose dolphins are present in Galveston waters throughout the year.
Over 1,000 individuals were identified from this area over a 7-mo period
(Henningsen 199 l), but many of these dolphins were transients. I estimate that
only about 200 individuals are apparently resident, with resightings of many at
different times of the year according to analyzed photos from two years (Brager
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NOTES 435

June to August 1991

18 43 58 56 69 46 19 24 23 9 4 14 7

80%

60%

40%

20%

0%
6 7 8 9 10 11 12 13 14 15 16 17 18 19
Time of day

Figwe 1. Observation frequencies of the main behaviors during daylight hours in

Galveston waters between June and August 1991. Sample sizes are indicated above the
columns.

1992). All age classes, from small newborn calves to large older animals, are
represented.
Figures 1 and 2 show the distribution of behaviors over the daylight hours
(0600-1900 h) in the summer (JuneAugust) and in the fall (September-
November) of 199 1. Feeding, socializing, and traveling were defined according
to Shane (1990~~). Feeding and probable feeding behavior were distinguished
by repeated (long) dives in varying directions in one location. They also indude
observations of groups foraging behind trawling shrimp boats. Socializing be-
havior was distinguished by some or all group members being in almost constant
physical contact with one another, and often displaying social, playful, aggressive,
and/or exploratory surface behaviors. Traveling was distinguished as moving
steadily in one direction. Other behaviors, such as resting or riding on the bow
of a moving ship, were observed only occasionally and are pooled for this analysis.
Each group (defined as any number of dolphins observed in apparent association,
moving in the same direction and often, but not always, engaged in the same
activity; Shane 1990~) was included only once per hour and once per behavior
(regardless of group size and duration of observation). Groups displaying different
behaviors simultaneously or within the same hour were put in all relevant
categories. Therefore, Figures 1 and 2 represent frequencies of observed activities
rather than time budgets. Summertime observations were transformed from
daylight saving time to standard central time.
From June to August a well-defined diurnal pattern was evident, with feeding
mainly in the morning and a secondary smaller peak in late afternoon. Socializing
increased as feeding decreased, with peak socializing in the afternoon. Traveling
was observed most in late afternoon, when dolphins usually returned to the bay
from Gulf waters. From September to November this pattern vanished almost
totally, with decreasing amounts of socializing and traveling, and much feeding
throughout the day.
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436 MARINE MAMMAL SCIENCE, VOL. 9, NO. 4, 1993

September to November 1991

3 12 16 34 35 39 50 52 36 44 21 11 9
100%

80%

60%

20%

0%
6 7 8 9 10 11 12 13 14 15 16 17 18 19
Time of day

Figzlre 2. Observation frequencies of the main behaviors during daylight hours in

Galveston waters between September and November 1991. Sample sizes are indicated
above the columns.

Feeding may increase in fall because of possibly higher energy requirements

due to declining water temperatures. In Florida bottlenose dolphins increased
the time spent feeding when water temperature dropped from 26.7’-3 1.7”C to
15.6”20.6”C (Shane 199O~z), as it does in Texas waters (NOAA 1974).
However, I also believe that an extended feeding regime is manifested because
of decreasing prey abundance due to emigration of schooling fish out of the bay
system during fall (Gunter 1945, Nelson 1992). One third of all stomachs of
bottlenose dolphins from the northeastern Gulf of Mexico contained exclusively
fish; the rest contained mixtures of fish and invertebrates (Barros and Odell
1990), and it is, therefore, evident that fish play an important role in the diet
of dolphins from this area.
The diurnal and interseasonal changes in the frequencies of occurrence of the
three main behaviors of feeding, traveling, and socializing show dramatic sim-
ilarities between the present and two previous studies along the Texas coast
despite different methods of data collection and analysis (Gruber 1981, Mat-
agorda Bay, southwest of Galveston by 200 km; Shane 19906, Corpus Christi
Bay, southwest of Galveston by 290 km). In all three studies in disparate
locations, feeding peaked in the morning and decreased during the day. So-
cializing (which Gruber 1981, called “mating”) was highest in the second half
of the day. Traveling peaked in the afternoon or evening in all three studies.
As well, Gruber (1981) and Shane (19906) found the same increase in feeding
and concomitant decrease in socializing and traveling from summer to fall.
Although these patterns are so similar for the Texas studies, they differ from
the results Shane (1990~) gathered for bottlenose dolphins in southern Florida,
where water temperatures do not vary as much over the year and where fish
proved to be the sole diet in 68% to 82% of all stomachs (Barros and Odell
1990). Colder water temperatures at the Texas coast in winter could cause a
stronger emigration of fish prey and subsequently a shift in dolphin diet to more
cephalopods and crustaceans.
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NOTES 437

In South Africa, Saayman et al. (1973) found the behavior of free-ranging

bottlenose dolphins also to be “significantly influenced by a diurnal rhythm.”
The dolphins came into the bay early in the morning to feed, left at noon, and
returned to the bay for feeding again in the evening. Bottlenose dolphins in
Argentina displayed a different behavior pattern of resting in the morning, feeding
in deeper water at noontime, and socializing and feeding in the afternoon (Wiirsig
and Wiirsig 1979).

ACKNOWLEDGMENTS

I thank the Marine Mammal Research Program of Texas A&M University in Galveston
for funding this study, and its staff for help on the boat and in the laboratory. Special
thanks to Bernd Wursig and two anonymous referees for reviewing drafts of the manuscript
and for improving my English. This represents Contribution No. 2 5 of the Marine Mammal
Research Program, Texas A&M University at Galveston.

LITERATURE
CITED

BARROS,N. B., APITD

D. K. ODELL. 1990. Food habits of bottlenose dolphins in the
southeastern United States. Pages 309-328 in S. Leatherwood and R. R. Reeves,
eds. The bottlenose dolphin. Academic Press, San Diego, CA.
BRAGER,S. 1992. Untersuchungen zur Ortstreue und zum Vergesellschaftungsmuster
des Grossen Tiimmlers, Tursiops trzlncatzls (Montagu, 182 1). Unpublished diploma
thesis, Christian-Albrechts-Universitat, Zentralbibliothek, DW-2300 Kiel, F. R. of
Germany.
GRuBER,J. A. 198 1. Ecology of the Atlantic bottlenosed dolphin (Tursiops truncatus)
in the Pass Cavallo Area of Matagorda Bay, Texas. Unpublished M.Sc. thesis, Texas
A&M University, Library, College Station, TX.
GUNTER, G. 1945. Studies of marine fishes of Texas. Publications of the Institute for
Marine Sciences, University of Texas l:l-190.
HENNINGSEN,T. 1991. Zur Verbreitung und zur okologie des Grossen Tfimmlers
(Tursiops truncatus) in Galveston, Texas. Unpublished diploma thesis, Christian-
Albrechts-UniversitHt, Zentralbibliothek, DW-2300 Kiel, F. R. of Germany.
NATIONAL OCEANICAND ATMOSPHERIC ADMINISTRATION. 1974. Temperature, salinity,
oxygen, and phosphate in waters off U.S. Volume II: Gulf of Mexico. U.S. De-
partment of Commerce, Washington, DC.
NELSON, D. M., ed. 1992. Distribution and abundance of fishes and invertebrates in
Gulf of Mexico estuaries, Volume I: data summaries. ELMR Rep. No. 10. NOAA/
NOS Strategic Environmental Assessments Division, Rockville, MD.
SAAYMAN,G. S., C. K. TAYLERAND D. BOWER. 1973. Diurnal activity cycles in captive
and free-ranging Indian bottlenose dolphins (Tursiops aduncus Ehrenburg). Behaviour
44:212-233.
SHANE,S. H. 199Oa. Behavior and ecology of the bottlenose dolphin at Sanibel Island,
Florida. Pages 245-265 in S. Leatherwood and R. R. Reeves, eds. The bottlenose
dolphin. Academic Press, San Diego, CA.
SHANE,S. H. 1990b. Comparison of bottlenose dolphin behavior in Texas and Florida,
with a critique of methods for studying dolphin behavior. Pages 541-558 in S.
Leatherwood and R. R. Reeves, eds. The bottlenose dolphin. Academic Press, San
Diego, CA.
WuRSIG, B., AND M. W~RSIG. 1979. Behavior and ecology of the bottlenose dolphin,
Tursiops truncatus, in the South Atlantic. Fishery Bulletin 77:399-412.
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438 MARINE MAMMAL SCIENCE. VOL. 9. NO. 4. 1993

STEFAN BRaGER,~ Marine Mammal Research Program, Texas A&M University,

4700 Ave. U, Bldg. 303, Galveston, TX 7755 1. lCurrent address: Scharstorfer
Weg 12, DW-2308 Schellhorn, F. R. of Germany. Received December 7,
1992. Accepted March 9, 1993.