SSC San Diego TD 627 Revision D,
Annotated Bibliography of Publications from the U.S. Navy's Marine Mammal Program, May 1998


Beach, F. A., III, and R. L. Pepper. 1971. Marine Mammal Training Procedures: The Effects of Scheduled Reinforcement in the Dolphin (Tursiops truncatus). NUC TP 214, 72 pp.

The strength of a behavior conditioned on a one-reward-for-one-response basis was compared with that conditioned on several types of schedules in which more than one response was required for a reward. All schedules provided good control, but a variable-ratio schedule produced a larger amount of work over a longer period and at a greater rate for the same amount of reward.

Beach, F. A., III, and L. M. Herman. 1972. Preliminary Studies of Auditory Problem Solving and Intertask Transfer by the Bottlenosed Dolphin. Psych. Rec. 22: 49-62.

Describes successive reversal training and discrimination learning set experiments with two bottlenosed dolphins.

Beach, F. A., III, and R. L. Pepper. 1972. Operant Responding in the Bottlenosed Dolphin (Tursiops truncatus). Jour. Exper. Anal. Behavior 17 (2): 159-160.

Describes experiments to determine the relative efficacies of various food-reinforcement schedules in a paddle-press task.

Beach, F. A., III, R. L. Pepper, J. V. Simmons, Jr., P. F. Nachtigall, and P. A. Siri. 1974. Spatial Habit Reversal in Two Species of Marine Mammals. Psych. Rec. 24: 385-391.

Two California sea lions and one Atlantic bottlenosed dolphin were tested over 19 reversals of a spatial problem. All performed well.

Chaplin, M., T. Kamolnick, M. Todd, and W. Van Bonn. 1996. Conditioning Tursiops Truncatus for Nasal Passage Endoscopy. (Abs.) 24th Annual IMATA Conf., Gold Coast, Australia, Nov. 3-8.

Details training of two Tursiops to accept a flexible fiberoptic endoscope into the blowhole to view the internal upper respiratory anatomy (epiglottal, nasal cavity and peri-blowhole structure) during echolocation to examine the role this structure may play in sound production.

Chun, N. K. W. 1978. Aerial Visual Shape Discrimination and Matching-to-Sample ProblemSolving Ability of an Atlantic Bottlenosed Dolphin. NOSC TR 236, 24 pp.

Two-dimensional geometric shapes of various configurations were presented. Large differences in perimeter lengths between any two shapes generally resulted in better performance. Other form parameters may be involved in the discriminative process. Evidence indicated problem-specific rather than conceptual learning.

Cramer, A. and A. Pitts. 1996. The Use of Commercially Available Software Programs in Marine Mammal Management. (Abs.) 24th Annual IMATA Conf., Gold Coast, Australia, Nov. 3-8.

Discusses the use of commercial data management applications for behavioral and dietary data collection in management of marine mammals, which allowed decisions regarding the animals’ health and training.

Cummings, W. C., P. O. Thompson, and J. F. Fish. 1974. Behavior of Southern Right Whales: R/V Hero Cruise 72-3. Antarctic Jour. U.S. 9 (2):33-38.

Describes behaviors and underwater sounds of right whales in Golfo San Jose, Argentina. No decided change in behavior was elicited by playback of southern right whale sounds or the sounds of northeast Pacific killer whales.

Flanigan, W. F., Jr. 1974. Nocturnal Behavior of Captive Small Cetaceans. 1. The Bottlenosed Porpoise (Tursiops truncatus). (Abs.) Sleep Research. 3: 84.

Observed behavior consisted of periods of unambiguous waking, stereotypic circular swimming with brief (20-30 seconds) eye closure and other indications of sleep, and quiescent "hanging" behavior with similar indications of sleep.

Flanigan, W. F., Jr. 1974. Nocturnal Behavior of Captive Small Cetaceans. 2. The Beluga Whale (Delphinapterus leucas). (Abs.) Sleep Research 3: 85.

Observed behavior consisted of active waking, quiet waking, and stereotypic circular swimming. The last does not meet all the criteria used to define sleep in terrestrial animals but probably reflects adaptations to an aquatic environment.

Gisiner, R. C., and R. J. Schusterman. 1991. California Sea Lion Pups Play an Active Role in Reunions With Their Mothers. Animal Behavior 41 (2): 364-366.

Presents and discusses data showing that pups less than one week old respond only to their mothers’ call and as they grow older, play an increasingly active role in reunions with others which have spent time away from the pup.

Gisiner, R. C. and R. J. Schusterman. 1991. Complex Conditional Relationships Learned by a Language-Trained Sea Lion (Abs.) 32nd Annual Meeting of the Psychonomic Society, San Francisco, November 1991. Bulletin of the Psychonomic Society 29 (6):486.

A sea lion was trained to respond appropriately when 2 to 7 signs were presented. If missing, added, or disoriented signs were presented the animal demonstrated that she had learned more than the specifically trained paired-associate relationships between signs and referents.

Gonzales, B.A., R.E. Cartee and S.H. Ridgway. 1993. Conditioning a California Sea Lion (Zalophus californianus) for Voluntary Ultrasound as a Husbandry Tool. 21st Annual IMATA Conf., Kailua-Kona, HI, Nov. 7-12.

Sea lions were trained to present different body parts for ultrasound evaluation. Methods of training were discussed.

Hall, R.W. and R. Stitt, 1996. Conditioning Dolphins for Temporary Holding Pools. 24th Annual IMATA Conf., Gold Coast, Australia, Nov. 3-8.

Navy dolphins had been temporarily maintained in "above ground" holding pools for medical procedures, obersvation, environmental emergencies and at-sea shipboard deployments. Explains the behavioral conditioning process for various pool configurations, emphasizing acclimation, medical and beaching behaviors.

Haun, J. E. 1977. Trainer and Trainer Transfer of Marine Mammals Utilizing Collateral Behaviors. In: Proceedings, IMATA Conference, D. I. McSheehy and G. B. Peiterson (eds.). New England Aquarium, Boston, pp. 65-79.

Discusses the classification and quantification of innovative or animal-initiated behaviors that occur in performance of a conditioned chain of behaviors and describes how such collateral behaviors pertain to training and can be used to facilitate transfer of an animal from one trainer to another.

Hui, C. A. 1989. Surfacing Behavior and Ventilation in Free-ranging Dolphins. Jour. Mammal. 70 (4):833-835.

Ventilation intervals are estimated from video images of Delphinus and Stenella swimming near boats. Results used to support speculations on behavioral and bioenergetic consequences of dolphins surfacing to breathe at different swimming speeds.

Irvine, B. 1971 (1972). Behavior Changes in Dolphins in a Strange Environment. Quart. Jour. Florida Acad. Sci. 34 (3):206-212.

Sluggish and unresponsive behavior was observed in dolphins when they were first moved from tanks to lagoon pens. Similar behavior was noted in animals that escaped from their pens or wandered away from the trainer during early training. It is suggested that this was a response to a strange environment.

Kamolnick, T., M. L. Reddy, D. Miller, C. Curry and S. Ridgway. 1992. Conditioning a Bottlenose Dolphin (Tursiops truncatus) for Milk Collection. 20th Annual IMATA Conf., Grand Bahamas, Nov. 1-6, p. 24.

Two lactating Tursiops truncatus was conditioned to allow for the collection of milk samples. Training methods are outlined.

Kamolnick, T., M. Reddy, D. Miller, C. Curry and S. Ridgway. 1994. Conditioning a Bottlenose Dolphin (Tursiops truncatus) for Milk Collection. Marine Mammals: Public Display and Research. 1 (1): 22-25.

Two lactating Tursiops truncatus were conditioned to allow for the collection of milk samples. Training methods are outlined and collection devices are reviewed.

Kamolnick, T., M. Todd, M. Beeler and J. Ross. 1996. Conditioning Strategy for Open Water Research. (Abs.) 24th Annual IMATA Conf., Gold Coast, Australia, Nov. 3-8.

Reports on follow-on to an earlier study in which white whales (Delphinapterus leucas) were conditioned for hearing studies at 100 meters. This extended that testing to 200 and 300 meters. Discusses the development of conditioning strategies employed in the study in the unpredictable environment of the open sea.

Leatherwood, J. S. 1974. A Note on Gray Whale Behavioral Interactions with Other Marine Mammals. Mar. Fish. Rev. 36 (4):50-51.

Porpoises of a number of species were observed riding the pressure waves of the whales. Whales were also observed riding large swells in a manner similar to that seen in smaller cetaceans.

Leatherwood, J. S. 1975. Some Observations of Feeding Behavior of Bottlenosed Dolphins (Tursiops truncatus) in the Northern Gulf of Mexico and (Tursiops cf T. gilli) off Southern California, Baja California, and Nayarit, Mexico. Mar. Fish. Rev. 37 (9): 10-16.

Seven distinct feeding behaviors, in which a variety of prey species are taken by various means, are identified and discussed.

Leatherwood, J. S. 1977. Some Preliminary Impressions on the Numbers and Social Behavior of Free-swimming Bottlenosed Dolphin Calves (Tursiops truncatus) in the Northern Gulf of Mexico. In: Breeding Dolphins: Present Status. Suggestions for the Future, S. H. Ridgway and K. Benirschke (eds.). A report to the Marine Mammal Commission. Nat’l. Tech. Info. Serv. PB-273 673, pp. 143-167.

Data from aerial observations on numbers of calves observed with respect to (1) positions within herds of subgroups containing calves, (2) positions of calves within subgroups, (3) interactions between calves and other animals in the herd, and (4) changes in the behavior of calves with age. Apparent patterns were seen. Relevant observations from other field and captive studies are discussed.

Leatherwood, J. S., and D. K. Ljungblad. 1979. Nighttime Swimming and Diving Behavior of a Radio-tagged Spotted Dolphin (Stenella attenuata). Cetology, No. 34, 6 pp.

The dolphin, radio-tracked from shipboard for 13 consecutive hours, covered 100.5 km at estimated speeds of 2.3 to 10.7 knots, with burst speeds exceeding 12 knots. The animal dove for from 1 to 204 seconds, exhibiting three diving modes tentatively identified as running, traveling, and feeding.

Ljungblad, D. K., and S. E. Moore. 1983. Killer Whales (Orcinus orca) Chasing Gray Whales (Eschrichtius robustus) in the Northern Bering Sea. Arctic 36:361-364.

Behaviors observed when 16 killer whales approached and chased feeding gray whales. No whale sounds were picked up by a sonobuoy although widely spaced killer whales exhibited apparently coordinated movements.

Ljungblad, D. K., B. Wursig, S. L. Swartz, and J. M. Keene. 1988. Observations on the Behavioral Responses of Bowhead Whales (Balaena mysticetus) to Active Geophysical Vessels in the Alaskan Beaufort Sea. Arctic. 41 (3):183-194.

Results from four field experiments support the conclusion that short-term behavioral changes occur when bowhead whales are exposed to airgun blasts from vessels within 10 km. The effects of airgun disturbance wane within an hour.

Murchison, A. E., and R. L. Pepper. 1972. Escape Conditioning in the Bottlenosed Dolphin (Tursiops truncatus). Cetology No. 8, 5 pp.

To evaluate the effectiveness of procedures other than food reward in establishing behavioral control, the use of an aversive stimulus was investigated. The animal was successfully conditioned to approach an emitting hydrophone in order to terminate the presentation of a moderately intense sound delivered underwater.

Murchison, A. E., and S. A. Patterson. 1980. The Effect of Extended Reinforcement Schedules on the Receiver Operating Characteristics (ROC) of an Echolocating Atlantic Bottlenosed Dolphin (Tursiops truncatus). (Abs.) Jour. Acoust. Soc. Am. 68 (Suppl. 1): 597.

After a dolphin was conditioned to report (by paddle press) presence or absence of a target, its performance was tested using different variable and fixed-ratio reinforcement schedules. The dolphin’s ROC remained essentially unchanged for all schedules, but when it was kept on the more extended schedules for more than eight consecutive 100-trial sessions, all responses became "target absent."

Nachtigall, P. E. 1971. Spatial Discrimination and Reversal Based on Differential Magnitude of Reward in the Dolphin (Tursiops truncatus). Eighth Annual Conf. on Biological Sonar of Diving Mammals, Menlo Park, CA, pp. 67-72.

Tursiops responds to differential reward magnitudes (four smelt versus one smelt) in a manner characteristic of other animals similarly studied.

Nachtigall, P.E. 1989. Echolocation Sameness-Difference and Matching-to-Sample: Demonstration of Dolphin Cognitive Processes. (Abs.) Animal Language Workshop, April 6-10, Honolulu, HI

A presentation summarizing echolocation sameness-difference and delayed matching-to-sample experiments that demonstrated concept formation by echolocating dolphins.

Nachtigall, P.E., J. Lien, W.W.L. Au, and A.J. Read. 1995. Harbour Porpoises: Laboratory Studies to Reduce Bycatch. DeSpil Publishers, Woerden, The Netherlands, 167 pp.

This volume was organized in response to the growing international problem of harbor porpoises becoming entrapped and dying in fishing nets. It presents a summary of laboratory experiments specifically designed and carried out to examine net entrapment and ways to prevent animals from becoming entangled. Studies included an examination of acoustic signals, echolocation signal characteristics, behavior in response to ropes, entanglement and acoustic alarms to warn porpoises about nets.

Pawloski, D. A., and P. W. B. Moore. 1987. Combined Stimulus Control of Peak Frequency and Source Level in the Echolocating Dolphin (Tursiops truncatus). 15th Annual IMATA Conf., New Orleans, LA, Oct. 26, 1987, pp. 3-9.

The training methods by which an echolocating dolphin was trained to control its emitted source level and the frequency content of the echolocation click are presented.

Pawloski, J. L. 1990. Training a False Killer Whale (Pseudorca crassidens) for an Underwater Audiogram. (Abs.) 18th Annual IMATA Conf., Chicago, Nov. 4-9, 1990.

Details the training and testing for an underwater masked hearings threshold study using a false killer whale. Also discusses training problems related to this type of testing.

Pepper, R. L., and F. A. Beach, III. 1972. Deprivation and Other Aspects of Food Reinforcement in the Dolphin. Ninth Conf. on Biological Sonar and Diving Animals, 10 pp.

Dolphin behavior in a simple automated task was found to be responsive to controlled variations in food reinforcement.

Pepper, R. L., and F. A. Beach, III. 1972. Preliminary Investigations of Tactile Reinforcement in the Dolphin. Cetology. No. 7, 8 pp.

Tactile reinforcement, gradually substituted for fish in a paddle-press task, at first maintained good response. After extensive testing, behavioral breakdown occurred. Aggressive behavior directed toward the trainer was interpreted as sexual frustration.

Pepper, R. L., and R. H. Defran. 1975. Dolphin Trainers Handbook, Part 1. Basic Training. NUC TP 432, 52 pp.

A handbook of information and guidance for dolphin trainers.

Ridgway, S. H. 1966. Studies on Diving Depth and Duration in Tursiops truncatus. 1966 Conf. on Biological Sonar and Diving Mammals, Menlo Park, CA, pp. 151-158.

Describes technique by which a bottlenosed porpoise was trained to dive to depths down to 550 feet and perform other tasks in preparation for participation in Sealab II. Total dive time to 550 feet and back averaged 163 seconds.

Ridgway, S. H., D. A. Carder, and M. M. Jeffries. 1985. Another Male "Talking" White Whale. (Abs.) Sixth Biennial Conf. on the Biology of Marine Mammals, Society of Marine Mammalogy, Vancouver, B.C., Canada, Nov. 22-26, p.67.

A male white whale nearing the attainment of sexual maturity spontaneously began to vocalize in a way that sounded like human speech heard at a distance. This phenomenon of white whale vocalization had been noted twice previously by others, but the physical characteristics of the sounds had not been presented. The spectral characteristics of distinctive phonations were measured and are described.

Ridgway, S., T. Kamolnick, M. Reddy, C. Curry, and R. Tarpley. 1993. Re-lactation and Induced Lactation in Tursiops and Analysis of Milk Collected with a Dolphin Milking Device. Tenth Biennial Conf. on the Biology of Marine Mammals, Galveston, Texas, Nov. 11-15, 1993, p. 91.

Milk was collected from two Tursiops that were induced to lactate by the presence of orphaned calves and analyzed for fat content. This was the first time milk was collected serially from dolphins conditioned to volunteer for the procedure.

Ridgway, S.H., T. Kamolnick, M. Reddy, C. Curry and R.J. Tarpley. 1995. Orphan-Induced Lactation in Tursiops and Analysis of Collected Milk. Marine Mammal Science. 11 (2): 172-182.

Reports on composition of milk voluntarily collected from two adult females induced to lactate by the presence of orphaned calves. This is the first report of cetacean females re-lactating to nurse calves not their own. It is also the first analysis of milk produced by induced lactation.

Ross, J., S. Klappenback and M. Xitco. 1996. The U.S. Navy’s Newest Recruits: Exploring Novel Directions for the Marine Mammal Program. (Abs.) 24th Annual IMATA Conf., Gold Coast, Australia, Nov. 3-8.

Reports on Navy’s PROGENY project involving five 4- and 5-year-old bottlenose dolphins (Tursiops truncatus) being studied and trained as a group. Reviews conceptual techniques such as co-operative target detection, "mimicry" or behavioral modeling, and "match-to-sample" abilities.

Chun, N. K. W. 1978. Aerial Visual Shape Discrimination and Matching-to-Sample Problem-Solving Ability of an Atlantic Bottlenosed Dolphin. NOSC TR 236, 24 pp.


Schusterman, R. J. 1981. Behavioral Capabilities of Seals and Sea Lions: A Review of Their Hearing, Visual, Learning, and Diving Skills. Psych. Rec. 31:125-143.

Compares behavioral/sensory capabilities of otariids (fur seals and sea lions) and phocids (earless seals).

Schusterman, R. J., B. K. Grimm, R. C. Gisiner, and F. B. Hangii. 1991. Retroactive Interference of Delayed "Symbolic" Matching-to-Sample in California Sea Lions (Abs.) Bulletin of the Psychonomic Society. 32nd Annual Meeting, San Francisco, CA., November 1991. 29(6):486.

Experiments showed that two sea lions demonstrated nearly complete forgetting when irrelevant comparison stimuli were shown during the delay intervals in a two-choice delayed conditional discrimination task. Simple delays of 1 second to 2 minutes in one animal did not affect performance and the other animal showed some forgetting when the delay was from 1 to 45 seconds.

Thomas, J. A., L. M. Ferm and V. B. Kuechle. 1987. Silence as an Antipredation Strategy by Weddell Seals. Antarctic Jour. of the U.S. 22 (5):232-234.

The hourly rate of underwater vocalizations over the day was collected near McMurdo Sound, Antarctica, from October through January for three seasons. In mid-December, for three years, the number of Weddell seal sounds decreased dramatically at the same time that killer whale and leopard seal vocalizations increased. The study proposes that as the two predatory species move near breeding colonies of Weddell seals, they shift from a highly vocal behavior to silence to avoid attracting attention to newly weaned seal pups.

Thomas, J. A., L. M. Ferm, and V. B. Kuechle. 1988. Patterns of Underwater Calls from Weddell Seals (Leptonychotes weddelli) During the Breeding Season at McMurdo Sound, Antarctica. Antarctic Jour. of the U.S. 23 (5): 146-148.

Seasonal changes in the hourly rate of vocalizations by Weddell seals was documented by automated cassette recorders. The rates changed in a way that predicted the onset of reproductive activities such as pupping, weaning, mating, and dispersal.

Wood, F. G., and S. H. Ridgway. 1967. Utilization of Porpoises in the Man-In-The-Sea Program. In: An Experimental 45-Day Undersea Saturation Dive at 205 Feet. ONR Report ACR-124, p. 407-411.

At Sealab II a bottlenosed dolphin named Tuffy was trained to carry objects between the surface and aquanauts working on the ocean floor. He also demonstrated his ability to carry a line from the habitat to a "lost" aquanaut. The conditioning of Tuffy and details of his transports to and from the Sealab site are described.

Wood, F. G., D. K. Caldwell, and M. C. Caldwell. 1970. Behavioral Interactions Between Porpoises and Sharks. In: Investigations on Cetacea, Vol. II, G. Pilleri (ed.). Institute of Brain Anatomy, Berne, Switzerland pp. 264-279.

Sometimes porpoises attack sharks, sometimes sharks attack (and eat) porpoises, and sometimes mutual tolerance is exhibited. The relationship of porpoises and sharks is still inadequately understood.

Wood, F. G. 1986. Social Behavior and Foraging Strategies of Dolphins. (Section introduction) In: Dolphin Cognition and Behavior, R. J. Schusterman, J. A. Thomas, and F. G. Wood (eds.). Lawrence Erlbaum Associates, Hillsdale, NJ, pp. 331-333.

Dolphin social and feeding behaviors, first observed in oceanariums, have now been studied in free-ranging animals. While conditions of captivity may distort natural patterns, observations of behavior of captive animals can be useful in interpreting that of free-living dolphins. Captive conditions appear to stimulate dolphin propensities for play and the invention of games.