2012 COURSE DATES: AUGUST 4 – 17, 2012 - Sirenian International

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Anim Cogn (2009) 12:43–53 51 the prey to escape or allowing the calf to eat the prey, as well as the energy to play with the prey rather than just eating it quickly and efficiently herself. It would be more efficient and less costly for the mothers to have simply caught and consumed the fish, or perhaps to have left the calf in the care of an alloparent, as opposed to this altered foraging behavior observed in these events. Furthermore, the chase latencies and foraging behaviors were drastically different with their calves present, not just with the longer latencies but also with the presumed referential behaviors toward the prey objects and allowing the still nursing calf to participate, which would be more indicative of social learning. If the calves were merely a distraction, the mothers could have either immediately consumed the prey or disciplined the calf, as previously observed by mothers in the population for undesirable behaviors (Miles and Herzing 2003), rather than allowing the calf to participate. Therefore, because the calves appeared attentive, interacted with the prey and the mother, and appeared to need little care during the event, they were not likely a distraction to the mother. Reciprocally, the mothers may have been altering their foraging behavior because their calves were attentive, so that a calf’s attention may have stimulated the altered maternal foraging behavior. Further analysis into the calves’ behavior is warranted and may also elucidate the exact learning mechanism at play on behalf of the calf. For the majority of the mothers, there was a difference in maternal age between the events observed foraging without their calves compared to the events foraging with their calves. It is not likely that the age differences, or resulting level of experience would have resulted in the observed differences in chase latencies and number of body-orienting movements. Rather, an older or more experienced dolphin should be expected to have quicker chase latencies and fewer body-orienting movements due to increased efficiency, not the longer latencies and increased number of body-orienting movements as observed here. Additionally, of the nine mothers observed in this study, three of the mothers, Little Gash, Mugsy, and PR2, were observed foraging both with and without their calves present during the same year, and each of these three mothers were observed foraging both with and without calves during the same field encounter. For these three mothers, mean chase latencies were much longer foraging with calves than foraging alone at the same age and comparable to the mean chase latencies of foraging alone at an earlier age (Table 2). This suggests that there was not a difference in foraging behavior due to difference in age or experience, but rather due to the presence of their calves. There were also no significant differences for chase latencies or number of body-orienting movements for the prey species observed in this study: thus the altered foraging behaviors were also not likely due to the difficulty of catching a specific type of prey. The three episodes in which a target dolphin was observed foraging with a non-calf individual, juvenile in age or older, produced chase latencies and number of bodyorienting movements comparable to when these dolphins foraged alone, and both substantially different from the levels of behavior observed when these dolphins foraged with their calves present. This suggests that the change in behavior was not merely a social phenomenon seen in the presence of other individuals, but instead reflects teaching behavior targeted at a naïve observer. However, the naïve observers in this study were presumably the calves of the observed mothers. Future research is needed to explore if these same mothers or other alloparents also exhibit similar altered foraging behavior when in the presence of other naïve observers that are not their offspring. This altered foraging behavior on behalf of the mothers may be a valuable social learning mechanism for Atlantic spotted dolphins. The mothers in the study clearly altered their typical foraging behavior in the presence of their calves at a potential cost to themselves due to the exaggerated behavior, as well as sometimes foregoing ingestion of the prey. However, as per Caro and Hauser’s definition of teaching, the observer must benefit from the model’s altered behavior, in this case through more rapid or skillful acquisition of foraging behavior. In dolphin society this skill would be essential, as mothers give birth approximately every 3 years, at which time the older calf would become weaned and rely more on independently caught fish. It is advantageous for the mother to ensure the success of her offspring, presumably by investing her time and energy into the calf’s foraging capabilities before the calf is weaned. Weaning is a gradual process in spotted dolphins, and although the calves were not dependent upon fish for food, consuming the fish may have been an important part of the development of the calves’ foraging behavior, perhaps because it reinforced the social-learning process. Future research is warranted to explore the development and skillfulness of the young calf’s pre-weaning foraging capabilities in order to examine the full effect of the mother’s presumed teaching efforts. If this is true teaching behavior, the calf will derive some benefit from observing the mother’s altered foraging behavior. Additionally, a comparison is needed to compare the calves of teaching and non-teaching or less attentive mothers. However, because all of the mothers observed in this study demonstrated the altered foraging behavior, it may be difficult to assess the consequences of naturally occurring individual differences in the mothers’ foraging behavior on the calves. Although the full benefit for the calf is unknown, it seems 123
52 Anim Cogn (2009) 12:43–53 that the calf is indeed the target of this altered behavior, which was observed only in the presence of the calf. Additionally, further research is needed into the calf’s behavior and attention to the mother. Future research can hopefully clarify if the calf is indeed attentive to the mother’s potential teaching behavior to gain something from the experience, as well as clarify what mechanism the calves may be using to learn from the mothers, such as imitation, stimulus enhancement, or local enhancement. Data from this study, such as the calves being predominantly in the observation position relative to the mother during the events, some of the calves chasing and ingesting the fish, and the calves’ interactions with both the mothers and prey objects, may be evidence that the calves were attentive to the mothers’ altered foraging behavior and support the argument for teaching. Future data from the calf behavior will also hopefully strengthen our argument that the altered maternal foraging behavior may be an example of teaching. The data from the calf behavior are currently being collected and analyzed as part of a separate ongoing project and will be available for future publication. Despite this drawback, we believe that this study detailing the altered foraging behavior of the mothers is a significant finding in the area of animal cognition, even without data on the calf behavior. Despite previous research that has shown that dolphins pass mirror self-recognition tests (Reiss and Marino 2001), understand referential pointing (Herman et al. 1999), and spontaneously use referential gesturing in captive situations (Xitco et al. 2001), further evidence is needed before attributing theory of mind to dolphins, which some researchers argue is required for true teaching (Tomasello et al. 1993). Regardless of the lack of conclusive evidence supporting theory of mind in dolphins, the perspectivetaking abilities by dolphins supported by previous research (Herman et al. 1999; Xitco et al. 2001) might be sufficient for the presumed teaching behavior shown here. Although the cognitive abilities behind the clear alteration of foraging behavior of the mother dolphins in the presence of their calves are yet to be determined, the observed teaching behaviors in dolphins are nonetheless remarkable. Mother dolphins provide opportunities for calves to observe foraging behaviors, and sometimes even provide opportunities for calves to practice foraging. Teaching, then, may be an important way in which aspects of cetacean social learning and possibly culture are transmitted from one generation to the next. Acknowledgments We would like to thank Stan Kuczaj and Jesse Bering for their helpful comments on earlier drafts of this manuscript, and John Bender, Miley Fishero, Megan Rothrock, Melissa Ingui, Wisline Shepherd, Sheryl Spencer, and the Wild Dolphin Project for their assistance on the project. 123 References Bering JM (2001) Theistic percepts in other species: can chimpanzees represent the minds of non-natural agents? J Cogn Cult 1:107– 137 Bering JM, Povinelli DJ (2003) Comparing cognitive development. In: Maestripieri D (ed) Primate psychology. Harvard University Press, Cambridge, pp 205–233 Bjorklund DF, Bering JM (2003) Big brains, slow development, and social complexity: the developmental and evolutionary origins of social cognition. In: Brüne M, Ribbert H, Schiefenhövel W (eds) The social brain: evolutionary aspects of development and pathology. Wiley, New York, pp 133–151 Bjorklund DF, Yunger JL, Bering JM, Ragan P (2002) The generalization of deferred imitation in enculturated chimpanzees (Pan troglodytes). Anim Cogn 5:49–58 Boesch C (1991) Teaching in wild chimpanzees. Anim Behav 41:530–532 Boesch C (1993) Toward a new image of culture in chimpanzees. Behav Brain Sci 16:514–515 Boesch C, Tomasello M (1998) Chimpanzee and human cultures. Curr Anthropol 39:591–614 Caro TM, Hauser MD (1992) Is there teaching in nonhuman animals? Q Rev Biol 67:151–174 Greenfield P, Maynard A, Boehm C, Schmidtling EY (2000) Cultural apprenticeship and cultural change: tool learning and imitation in chimpanzees and humans. In: Parker ST, Langer J, McKinney ML (eds) Biology, brains, and behavior: the evolution of human development. School of American Research Press, Santa Fe, pp 237–277 Guinet C, Bouvier J (1995) Development of intentional stranding hunting techniques in killer whale (Orcinus orca) calves at Crozet Archipelago. Canadian Journal of Zoology 73:27–33 Herman LM (2002) Vocal, social, and self-imitation by bottlenosed dolphins. In: Nehaniv C, Dautenhahn K (eds) Imitation in animals and artifacts. MIT Press, Cambridge, pp 63–108 Herman LM, Abichandani SL, Elhajj AN, Herman EYK, Sanchez JL, Pack AA (1999) Dolphins (Tursiops truncatus) comprehend the referential character of the human pointing gesture. J Comp Psychol 113:347–364 Herzing DL (1996) Vocalizations and associated underwater behavior of free-ranging Atlantic spotted dolphins, Stenella frontalis, and bottlenose dolphins, Tursiops truncatus. Aquat Mamm 22:61–79 Herzing DL (1997) The life history of free-ranging Atlantic spotted dolphins (Stenella frontalis): age classes, color phases an female reproduction. Mar Mamm Sci 13:576–595 Herzing DL (2005) Transmission mechanisms of social learning in dolphins: underwater observations of free-ranging dolphins in the Bahamas. In: Delfour F (ed) Autour de L’ethologie et de la cognition animale. Presses Universitaires de Lyon, Lyon, pp 185–194 Herzing DL, Brunnick BJ (1997) Coefficients of association of reproductively active female Atlantic spotted dolphins, (Stenella frontalis). Aquat Mamm 23:155–162 Kruetzen M, Mann J, Heithaus M, Connor R, Bejder L, Sherwin WB (2005) Cultural transmission of tool use in bottlenose dolphins. Proc Natl Acad Sci USA 105:8939–8943 Kuczaj SAII, Paulos RD, Ramos JA (2005) Imitation in apes, children and dolphins: implications for the ontogeny and phylogeny of symbolic representation. In: Namy L (ed) Symbol use and symbolic development. MIT Press, Cambridge, pp 221–243 Mann J, Smuts BB (1999) Behavioral development in wild bottlenose dolphin newborns (Tursiops sp.). Behaviour 136:529–566 Marino L, Connor RC, Fordyce RE, Herman LM, Hof PR, Lefebvre L, Lusseau D, McCowan B, Nimchinsky EA, Pack AA, Rendell
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ECOLOGY, BEHAVIOR & CONSERVATION OF
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discomfort without proper protectio
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ABOUT BELIZE Research Site: The pro
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FIELD TRAINING AND ASSIGNMENTS Duri
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ENVIRONMENTAL CONDITIONS SBCRC is s
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HEALTH INFORMATION Routine Immuniza
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• Consult the CDC for immunizatio
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Oil or oil-based repellant (e.g. ol
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Ecology, Behavior, and Conservation
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MINIMUM / MAXIMUM CLASS SIZE: 6-24
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COURSE FEE & ADDITIONAL INFORMATION
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Ecology, Behavior & Conservation of
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2012 Field Course Policy & Liabilit
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2012 Field Course Policy & Liabilit
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FIELD JOURNALS You are required to
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OUR INVESTIGATION Figure 2. G. crut
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OUR INVESTIGATION Introduction The
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Introduction Our Investigation Befo
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2010 VT: Scholar, BANNER, Safe Zone
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presentation to Horn Point Environm
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LaCommare, Katherine S. November 20
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2006. Holy Redeemer Day Camp. Conta
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during the summer months while othe
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polyandrous - often mating with sev
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in the summer. Chessie, a male Flor
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ecords indicate that manatees have
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Florida manatee. National Biologica
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parturition: the action or process
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(a) (b) Manatees on the Belize Barr
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direction, and (e) when ‘travelli
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male travelled at least as far as B
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NEWS OF THE WEEK MEETINGBRIEFS>> 10
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Functional Ecology 2008, 22, 284-28
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288 S. R. Noren Kelly, H.R. (1959)
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172 NOTES Caribbean Journal of Scie
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174 FIG. 1. Drowned Cayes study are
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176 severe event occurred in 1998 (
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Bradbury RH, Seymour RM, Antonelli
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Social Interactions in Captive Fema
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The Journal of Experimental Biology
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Fig. 1. Rescue locations of manatee
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The present study is the first to c
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a. b. c. Power Power Power 100 80 6
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