Observations on Pacific Cetaceans of Californian and Mexican Waters
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OBSERVATIONS ON PACIFIC CETACEANS OF CALIFORNIAN AND MEXICAN WATERS i BY j KENNETH S. NORRIS and JOHN H. PRESCOTT J UNIVERSITY OF CALIFORNIA PRESS BERKELEY AND LOS ANGELES 1961 1 UNIVERSITY OF CALIFORNIA PUBLICATIONS IN ZOOLOGY EDITORS (Los ANGELES); W. H. FURGASON, R. B. COWLES, T. L. JAHN Volume 63, No.4, pp. 291--402, plates 27-41, 12 figures in text Submitted by editors September 16, 1960 Issued November 21, 1961 Price. $2.25 UNIVERSITY OF CALIFORNIA PRESS BERKELEY AND Los ANGELES CALIFORNIA CAMBRIDGE UNIVERSITY PRESS LoNDON, ENGLAND PRINTED IN THE UNITED STATES OF AMERICA CONTENTS Introduction 291 Acknowledgments . 291 Cooperative Behavior 292 Assisted Locomotion. 295 Accounts of Species . 303 Cuvier's beaked whale 303 Distribution. 303 Capture observations and sightings . 304 Morphology. 304 Behavior . 305 Mortality and disease. 305 Gray's long-snouted porpoise . 306 Nomenclature . 306 Distribution. 306 Capture observations and sightings 306 Morphology. 306 • Pacific eommon dolphin. 309 Distribution . 309 Morphology . 309 Reproduction 309 Behavior. 311 Atlantic bottlenose porpoise 311 Behavior . 311 __ Pacific bottlenose porpoise . 313 Nomenclature . 313 Distribution. 313 Capture observations and sightings . 314 Morphology . 321 Reproduction . 321 Behavior . 321 Northern right whale porpoise 323 Distribution. 323 Capture observations and sightings . 324 Morphology 326 Behavior . 326 Parasitism . 326 • Pacific striped porpoise . 326 Distribution. 326 Capture observations and sightings . 327 Morphology. 329 Reproduction 329 Behavior . 330 Parasitism 330 [ iii] iv Contents Killer whale . 330 Distribution 330 Capture observations and sightings . 330 Morphology . 331 Behavior . 332 False killer whale . 334 Distribution. 334 Capture observations and sightings . 335 Morphology . 335 Behavior . 336 Pacific pilot whale 336 Distribution. 336 Capture observations and sightings . 337 Morphology . 338 Reproduction 342 Behavior . 342 Mortality and disease. 347 Parasitism. 348 Harbor porpoise . 348 Distribution. 348 Capture observations and sightings . 349 Morphology . 349 Gulf of California harbor porpoise. 349 Distribution. 349 Capture observations and sightings . 349 Dall porpoise . 351 Distribution. 351 Capture observations and sightings . 351 Morphology . 355 Reproduction 356 Behavior . 356 Parasitism. 358 California gray whale 358 Capture observations and sightings . 358 Behavior . 359 Little piked whale. 361 Nomenclature 361 Distribution. 361 Capture observations and sightings . 362 Morphology . 364 Reproduction 364 Literature Cited 367 Plates. 372 I OBSERVATIONS ON PACIFIC CETACEANS OF CALIFORNIAN AND MEXICAN WATERS BY KENNETH B. NORRIS AND .TOHN H. PRESCOTT INTRODUCTION THE CETACEANS are possibly the world's least-knmvn major group of mammals, at least with respect to ecology and behavior. This dearth of knowledge stems directly from the difficulties of observing these fleet aquatic mammals. In the past, knowl I edge has been derived largely from the fragmentary observations of whalers and J of a few biologists who spent long hours at sea waiting for the rare moments when these animals could be seen. Recently, several species of cetaceans have been observed in large salt-water viewing tanks, or oceanaria. The result has been a very significant increase in knowledge of some of the smaller cetaceans (see, e.g., McBride, 1940; McBride and Rebb, 1948; Kritzler, 1952; Essapian, 1953; Schevill and Lawence, 1953; Brown and Norris, 1956; Tavolga and Essapian, 1957). Most of this new literature deals with a single species, the Atlantic bottlenose porpoise, Tursiops truncatus (Montagu), which must now be classed as the best-known cetacean species. At Marineland of the Pacific the normal operations of capture and maintenance of cetaceans for exhibit have afforded remarkable opportunities for observation of several of the smaller Pacific coast species. For five years we have systemat ically gathered information of every available kind related to these animals. Fif teen species have been observed in sufficient detail to be considered here. These are Ziphius cavirostris Cuvier, SteneUa euphrosyne (Gray), Delphinus bairdi Dall, Tursiops truncatus (Montagu), Tursiops gilli Dall, Lissodelphis borealis (Peale), Lagenorhynchus obUquidens Gill, Orcinus orca (Linnaeus), Pseudorca crassidens (Owen), Globicephala scammoni Cope, Phocoena phocoena (Linnaeus), Phocoena sinus Norris and McFarland, Phocoenoides dalli (True), Eschrichtius glaucus (Cope), and Balaenoptera acutorostrata Lacepede. Even with our unusual opportunities for cetacean study, the bulk of our infor mation has come from chance observations of short duration. As a result, our observations are mostly fragmentary, and are of many different kinds. Probably, understanding of cetacean ecology will continue to be built up in this unsystematic piecemeal fashion, particularly for species of little commercial or exhibition value. We have arranged these diverse data by species, utilizing the phylogenetic scheme and nomenclature of Miller and Kellogg (1955), except where otherwise noted. A uniform series of subheadings has been used under each species. Information on two classes of behavior, cooperative behavior and assisted locomotion, so often crossed species lines that they seemed to require separate consideration and are presented first. ACKNOWLEDGMENTS Many people have helped us in gathering the observations involved in this report. Most important has been the help of the crew of the Marineland collecting vessel [291 ] 292 University of California Publications in Zoology Geronimo. Captain Frank Brocato and his able assistant Frank Calandrino are both keen and accurate observers, and have gathered much of the information presented here. We also wish to thank David H. Brown, Donald Hackett, Arthur Kelly, the late Conrad Limbaugh, William N. McFarland, and Wheeler North for observations reported here. John Evich, captain of the purse seiner Clermont, and Homer Moore, of the vessel GM, supplied us with specimens taken during fishing operations. We wish to thank John Fitch, senior research biologist of the California Department of Fish and Game, for identification of porpoise stomach contents listed in tables 6 and 13. Ira Cornwall provided identification of parasitic barnacles found on Lagenorhynchus obliquidens and GZobicephaZa scammoni, and Donald Heyneman has identified parasitic roundworms taken from Phocoenoides dalli. Thomas Lang, Howard Kelly, Moe William lli>sen, and Andrew Fabula of the Naval Ordinance Test Station, China Lake, California, have given advice on certain of the hydrodynamic aspects of the paper. Carl L. Hubbs, William E. Schevill, G. A. Bartholomew, and James Kubeck have given valuable editorial advice during preparation of the manuscript. Finally, our thanks go to our faithful and skillful secretary, Mrs. Muriel Johnson. COOPERATIVE BEHAVIOR It has become increasingly obvious in recent years that many species of cetaceans will offer assistance when a "schoolmate" becomes injured or otherwise unable to maintain normal locomotion. Usually this assistance consists of a normal animal or animals swimming with the incapacitated member, helping it stay at the surface, or actually taking it away from the apparent source of danger. The first recorded observation of this sort is found in the writings of Aristotle, set down twenty-four centuries ago (see Aristotle, p. 156). He noted a Mediterranean dolphin (possibly Delphinus delphis Linnaeus) supporting the body of a dead baby dolphin. Even very recently the existence of such behavior has been denied (Sanderson, 1956, p. 59), but evidence has begun to accumulate which shows that, in this particular, Aristotle was correct. McBride (1940) reported the case of a stillborn baby Tursiops truncatus that was carried to the surface and held there for an extended period by the mother. Hubbs (1953) reported an adult of the species Tursiops gilZi carrying a decomposing baby porpoise on its dorsal fin, off Imperial Beach, San Diego County, California. Moore (1953) saw an adult of Tursiops truncatus carrying a disemboweled and partially decayed newborn young on its head in Whipray Basin of southern Florida. The observations of Hubbs and Moore suggest that the dead young had been carried, presumably by the mother, for a matter of days, and perhaps as long as a week or more. Hubbs (1953) also reports the cooperative behavior of Lagenorhynchus obZiquidens, in which school mates attempted to force a harpooned individual away from its captors by swim ming against it and by standing between it and the ship. Siebenaler and Caldwell (1956) give two examples of cooperative behavior in Tursiops truncatus. An adult of this species began to swim erratically after being stunned by an under water explosion. Two other members of its school came up under the dazed animal and supported it. The cooperating animals took station below the stricken animal, one on each side, below each pectoral flipper. In the other instance, an adult of Norris-Prescott: Observations on Pacific Cetaceans 293 T. truncatus was stunned while being lowered into a holding pen. The animal sank in 7 feet of water and was then raised to the surface by two other captives, who supported it in a manner similar to that described above until it recovered. Brown and Norris (1956) reported three instances in which stricken adults of the species Lagenorhynchus obliquidens or Delphinus bairdi were assisted by other porpoises who swam beneath them, forcing them toward the surface. The