404 Short Communicationsand Commentaries [Auk, Vol. 110

CRAMP, S., AND K. E. L. SIMMONS (Eds). 1977. Birds LOOMIS,L. M. 1918. Expedition of the California of the western Palearctic, vol. I. Oxford Univ. Academyof Sciencesto the GalapagosIslands, Press, Oxford. 1905-1906. XII. A review of the albatrosses,pe- DEVILLERS,P. 1977. The skuas of the North American trels, and diving petrels.Proc. Calif. Acad. Sci. Pacific coast. Auk 94:417-429. (4th set.) 2(2):1-187. FORBUSH,E. H. 1929. Birds of Massachusetts and MURPHY,R.C. 1936. Oceanic birds of South America. other New Englandstates, part I. Massachusetts AmericanMuseum of Natural History,New York. Departmentof Agriculture,Norwood, Massachu- NORTON, A. I-I. 1922. The Pintado Petrel (Daption setts. capense)in Maine. Auk 39:101-102. JONES,E., AND I. J.S•cIRA. 1979. Breedingdistribution PIECE,N. 1873. Sub-tropicalrambles in the land of of the Great Skua at Macquarie Island in relation the Aphanapteryx.Personal experiences, adven- to numbers of rabbits. Emu 79:19-23. tures,and wanderingsin and around the island LAWRENCE,G.N. 1853a. Descriptionsof new species of Mauritius. Harper and Brothers,New York. of birdsof the generaOrtyx Stephens, Sterna Linn., WARHAM,J. 1969. Notes on some Macquarie Island and IcteriaVieillot. Ann. Lyceum Nat. Hist. (New birds. Notornis 16:190-197. York) 6:1-4. WATSON, G.E. 1975. Birds of the Antarctic and sub- LAWRENCE,G.N. 1853b. Additions to North Amer- Antarctic.American Geophysical Union, Wash- ican ornithology no. 3. Ann. LyceumNat. Hist. ington, D.C. (New York) 6:4-7. LEE,D. S. 1986. Secondrecord of the Cape Petrel in Received18 September1992, accepted 4 December1992. the western North Atlantic. Chat 50:118-119.

The Auk 110(2):404-406, 1993

Experimentally Induced Infanticide: The Removal of Birds and Its Ramifications

MARC BEKOFF Departmentof Environmental, Population, and Organismic Biology, University of Colorado, Boulder,Colorado 80309, USA

Therecurrently is greatinterest in the ethicalissues detail many of the ethicalproblems involved when surroundingresearch on nonhumananimals (here- performingmanipulations that includethe removal after animals),including field studiesof behaviorand of animals from, or the addition of animals to, already behavioralecology (American Society of Mammalo- existinggroups (the former of which is a common gists1987, Oring et al. 1988,Michener 1989, Animal manipulationin studiesof territorialityand parental Behavior Society 1991, Bekoff and Jamieson1991, care;Cuthill 1991).However, the guidelinesdo ad- Cuthill 1991, Bekoff et al. 1992). Thus, a study (Emlen dress some of the adverse effects of field research that et al. 1989)of experimentallyinduced infanticide in can stem from trapping, banding, and visiting nests WattledJacanas (]acana jacana) deserves scrutiny, for (seealso Henson and Grant 1991,Wilson et al. 1991), there are some matters of concern centering on: (i) all of which may be usedin studiesinvolving social the review processfor publication;(ii) the methods manipulation of animal groups. used to remove two adult female jacanas;(iii) the The paperby Emlenet al. (1989)was submittedto maimingand killing of sevenof their infantsby re- the Auk and reviewed after the AOU guidelines (Or- placementfemales as a directresult of the deliberate ing et al. 1988)on the useof wild birds in research removalof the mothersof the youngbirds (for further were published.Why was this study publishedin a discussionof someethical implications of infanticide journal sponsoredby a societythat had alreadyes- studies, see Elwood 1991; for recent review of studies tablishedguidelines concerning the ethicsof various of cognition,pain, and stress in birds,see Elzanowski types of research,some of which appear to be vio- and Abs 1991, Gentle 1992); and (iv) questions con- lated?Thus, while it is probably true that the two cerningtrade-offs between the importanceof acquir- adult females collected by shooting (a fact not in- ing differenttypes of knowledgeand the typesof cluded in the original paper) were "humanely col- animalresearch that are permissiblein the pursuitof lected" in accordancewith the AOU guidelinesthat this knowledge.The AOU guidelines(Oring et al. "Humane use of firearms necessitates that birds be 1988)also do not specificallyoutline proceduresfor killed outright" (Oring et al. 1988:10a),it is not clear studyinginfanticide, and do not directly addressin why alternativemethods of removing and replacing April 1993] ShortCommunications andCommentaries 405 the adult females were not used. Furthermore, the fering and the acquisitionof knowledge do not sub- experimental design that allowed subsequentand mit to simple answers (Driscoll and Bateson 1988, continuous harm to come to seven chicks as a direct Bekoff 1991, Bekoff and Jamieson 1991, Bekoff et al. resultof the killing of their mothersappears to violate 1992),especially when humanshave played a signif- at least two AOU guidelines (Oring et al. 1988:4a), icant role in bringing about the pain and suffering. namely, the requirements that: (a) Procedureswith With respectto the study by Emlen et al. (1989), be- animals must avoid or minimize distressand pain to causeWattled Jacanasare polyandrousand sexroles the animals,consistent with sound researchdesign. are reversed,the data may be interestingfrom a com- (b) Proceduresthat may causemore than momentary parative and evolutionary perspective,especially be- or slight pain or distress to the animals should be causethey are consistentwith data from reports on performedwith appropriatesedation or analgesia,ex- naturally occurring infanticide in nonpolyandrous cept when justified for scientificreasons in writing mammals.Yet, one can still questionfrom an ethical by the investigatorin advance. perspectivethe experimental methodsemployed by Terminologyis alsoimportant to consider,for there Emlen et al. (1989) that involved deliberately killing are some potential problems concerning the word adult females, the result of which included attacks on "collected."While it may be that useof the word "col- chicksby replacementfemales, aggressionthat was lected" is a nonissuefor some ornithologistsin that allowed to continueuntil chickswere alsokilled. Why it is an obvious synonym for "killed" (Oring et al. was it impossible to remove and then either replace 1988:10a-lla), I and one reviewer, for example, did or relocate the adult females?Why were the chicks not know that collectedstipulatively meant killed. given no opportunityto escapefrom the attacksthat Furthermore,professional journals are not only read were directedat them (e.g. Huntingford 1984)?Why by professionalsin the field. Thus, all readers, re- did the researchers choose not to interfere with on- viewers,and editors are entitled to know exactlywhat going aggressiondirected toward the chicks,attacks methods were used for collection, for different forms for which the researcherswere responsible,or oth- of collectionwill exposeanimals to varying degrees erwise protect the chicks from harm (e.g. Perrigo et of suffering (Oring et al. 1988:10a-lla). Researchers al. 1989)?It is inarguable that future studiesof infan- often use a variety of words either intentionally or ticide and other interestingsocial and nonsocialphe- unintentionally to refer to the act of killing other nomena must give seriousconcern to all individuals animals (e.g. sacrificing,euthanizing, culling). These who are being studied (Bekoff 1992), including those terms may serve both to distance or to detach re- who are affecteddirectly and indirectly by the ex- searchersfrom what they have done to the animals perimental methods that are used. they studyand to objectifymethods and the animals I thank Innes Cuthill, Susan E. Townsend, Dale they have chosento use (Lynch 1988,Lacy 1991,Ver- Jamieson, and two reviewers for comments on an hoog 1991).By talking or writing about the details of earlier draft of this paper. the typesof manipulationsthat have been performed in their studies,researchers may become more sen- LITERATURE CITED sitive to what they have actuallydone. This realiza- tion may force them to seek alternative and more ANIMAL BEHAVIOR SOCIETY. 1991. Guidelines for the humane methodsof study. use of animals in research. Anim. Behar. 41:183- One also needsto ask questionsabout the ethicsof 186. performing experimental studies to test hypotheses AMERICAN SOCIETY OF MAMMALOGISTS. 1987. Ac- that predictpossible killing or maimingas a resultof ceptablefield methodsin mammalogy:Prelimi- the manipulationsof animal groups (Huntingford nary guidelines approved by the American So- 1984). Emlen et al. (1989:5) indicated that there were ciety of Mammalogists.J. Mammal. 68(4, Supp1.): numerous intensive attacks on some chicks and that 1-18. seven of nine chicks had been "eliminated" after their BEKOFF,M. 1991. The animal'spoint of view, animal mothers had been removed. Indeed, some chicks were welfare, and some other related matters. Behar. brutalizedby replacementfemales to the point that Brain Sci. 14:753-755. they limped aboutor were unable to maintain their BEKOFF,M. 1992. What is a "scale of life"? Environ. balancewhile walking, prior to their disappearance. Values 1:253-256. The investigatorspresumed (with reason)that injured BEKOFF•M., AND D. JAMIESON. 1991. Reflective ethol- chicks who could not be followed died from their ogy, applied philosophy,and the moral statusof injuries. animals. Perspect.Ethol. 9:1-47. Were the lives of the killed females and chicks BEKOFF,M., L. GRUFmN,S. E. TOWNSEND,AND B. E. ROLLIN. worth a studyof experimentallyinduced infanticide? 1992. Animals in science: Some areas reconsid- Of course,questions concerning the importance of ered. Anim. Behav. 44:473-484. acquiringdifferent types of knowledgeare extremely CUTHILL,I. 1991. Field experiments in animal be- difficult to answer(Midgley, 1989).Also, discussions havior: Methods and ethics. Anim. Behar. 42: concerningtrade-offs between animal pain and suf- 1007-1014. 406 ShortCommunications and Commentaries [Auk, Vol. 110

DRISCOLL,J. W., AND P. BATESON. 1988. Animals in MICHENER, G. D. 1989. Ethical issues in the use of behavioral research. Anim. Behav. 36:1569-1574. wild animals in behavioral and ecologicalstud- ELWOOD,R.W. 1991. Ethical implicationsof studies ies. Natl. Agric. Libr., U.S. Department of Agri- on infanticide and maternal aggression in ro- culture. September, pages 37-43. dents. Anim. Behav. 42:841-849. MIDGLEY, M. 1989. Wisdom, information, and EEZANOWSKI,A., AND M. ABS(conveners). 1991. Pain knowledge: What is knowledge for? Routledge, and stressin birds. Pages1901-1940 in Acta XX New York. Congressus Internationalis Ornithologici. ORING, L. W., K. P. ABLE, D. W. ANDERSON, L. F. BAP- Christchurch, New Zealand, 1990. Ornithol. TISTA,J. C. BARLOW,A. S. GAUNT, F. B. GILL, AND Congr. Trust Board,Wellington. J. C. WINGFIELD. 1988. Guidelines for the use of EMLEN,S. T., N.J. DEMONG,AND D. J. EMLEN. 1989. wild birds in ornithologicalresearch. Auk 105(1, Experimental induction of infanticide in female Suppl.):la-41a. Wattled Jacanas.Auk 106:1-7. PERRIGO,G., W. C. BRYANT,L. BELVIN, AND F. $. VOM GENTLE,M. J. 1992. Pain in birds. Animal Welfare SAAL. 1989. The use of live pups in humane, 1:235-247. injury-free test for infanticidal behavior in male HENSON, P., AND T. A. GRANT. 1991. The effects of mice. Anim. Behav. 37:897-898. human disturbanceon Trumpeter Swan breeding VEmtOOG,H. 1991. The scientific perception of an- behavior. Wildl. Soc. Bull. 19:248-257. imals as object.Between the Species7:208-212. HUNTINGFORD,F.A. 1984. Some ethical issues raised WILSON, R. P., B. CULIK, R. DANFIELD, AND D. ADELUNG. by studies of predation and aggression.Anim. 1991. Peoplein Antarctica.How much do Ad61ie Behav. 32:210-215. Penguins Pygoscelisadeliae care? Polar Biol. 11: LACY,R.C. 1991. Zoos and the surplus problem: An 363-370. alternative solution. Zoo Biol. 10:293-297. LYNCH, M.E. 1988. Sacrifice and the transformation Received18 February1992, accepted17 August1992. of the animal body into a scientific object:Lab- oratory culture and ritual in the neurosciences. Soc. Stud. Sci. 18:265-289.

The Auk 110(2):406-409, 1993

Ethics and Experimentation:Hard Choicesfor the Field Ornithologist

STEPHEN T. EMLEN Sectionof Neurobiologyand Behavior,Cornell University, Ithaca, New York 14853,USA; and SmithsonianTropical Research Institute, Box 2072, Balboa, Republic of Panama

Every scientistmust make difficult ethical decisions that I welcome the opportunity to reply to the com- when designing experiments,whether such experi- mentary of Bekoff (1993). mentsare conductedin the laboratoryor in the field. Bekoff(1993) criticized our study (Emlen et al. 1989) Typically, thesedecisions require weighing the likely of experimentally induced infanticide in jacanason scientific gain (in terms of new information to be ethical grounds and chastisedthe American Orni- learned) againstthe animal cost(in terms of suffering thologists' Union for publishing our article in the of the individuals involved). The question of when Auk. As the senior author of the challenged paper, I the pursuit of knowledge justifiesthe imposition of wish to justify our specificexperiments, as well as suffering on animal subjectsis one that should be address the broader issue of ethical trade-offs in ex- honestly confronted and constantly reassessed.Most perimental science. scientific societieshave published guidelines to help If asked, everyone would agree that unnecessary individual scientistsformulate their answers(e.g. Or- and unnatural pain and suffering in animals should ing et al. 1988, Dawkins and Gosling 1992, Anony- be minimized wherever possible, but there exists a mous 1987, 1992). However, even with such guide- spectrum of opinions on when and whether inter- lines, there is no magic "threshold" of agreement. vention and experimentation are appropriate. At one Rather, there is a broad grey area within which dif- end, few would disagreethat many birds are kept in ferent opinions are vehemently expressed.Peer feed- captivity under sufficiently inhumane conditions that back is useful in defining these grey areas and in no degree of scientific justification can excusetheir stimulatingdiscussion about them. It is in this light poor care.At the other, field ornithologistsroutinely