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Alarm Calling in Yellow-Bellied Marmots: I

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Alarm Calling in Yellow-Bellied Marmots: I Anim. Behav., 1997, 53, 143–171 Alarm calling in yellow-bellied marmots: I. The meaning of situationally variable alarm calls DANIEL T. BLUMSTEIN & KENNETH B. ARMITAGE Department of Systematics and Ecology, University of Kansas (Received 27 November 1995; initial acceptance 27 January 1996; final acceptance 13 May 1996; MS. number: 7461) Abstract. Yellow-bellied marmots, Marmota flaviventris, were reported to produce qualitatively different alarm calls in response to different predators. To test this claim rigorously, yellow-bellied marmot alarm communication was studied at two study sites in Colorado and at one site in Utah. Natural alarm calls were observed and alarm calls were artificially elicited with trained dogs, a model badger, a radiocontrolled glider and by walking towards marmots. Marmots ‘whistled’, ‘chucked’ and ‘trilled’ in response to alarming stimuli. There was no evidence that either of the three call types, or the acoustic structure of whistles, the most common alarm call, uniquely covaried with predator type. Marmots primarily varied the rate, and potentially a few frequency characteristics, as a function of the risk the caller experienced. Playback experiments were conducted to determine the effects of call type (chucks versus whistles), whistle rate and whistle volume on marmot responsiveness. Playback results suggested that variation in whistle number/rate could communicate variation in risk. No evidence was found of intraspecific variation in the mechanism used to communicate risk: marmots at all study sites produced the same vocalizations and appeared to vary call rate as a function of risk. There was significant individual variation in call structure, but acoustic parameters that were individually variable were not used to communicate variation in risk. ? 1997 The Association for the Study of Animal Behaviour When alarmed by predators, many species pro- specific’, in that call structure in some way varies duce specific vocalizations (Klump & Shalter according to situation. 1984). Some species vary calls according to the Situationally specific calls can be produced type of predator detected (Seyfarth et al. 1980; several ways (Blumstein 1995a). Animals could (1) Davis 1984; Sherman 1985; Cheney & Seyfarth produce acoustically distinctive call types (an 1990; Marler et al. 1992; Macedonia & Evans apparent precursor to externally referential com- 1993), and others vary calls according to the munication), (2) vary the rate or number of times degree of risk the caller experiences, perhaps a single call type is produced, and/or (3) vary the according to the ‘response urgency’, or imminence overall intensity (i.e. volume) of a single call. Each of predation, that the caller faces (Robinson 1980; of these ‘mechanisms’ could be used singularly or Owings & Hennessy 1984; Blumstein 1995a). The in combination. A general assumption is that each distinction is important, because it was gener- species uses a single mechanism or a single com- ally assumed that only humans could com- bination of mechanisms to communicate variation municate about events and stimuli external to in situation. themselves; non-humans supposedly only com- To study the degree of situational specificity municated about their internal states (reviewed and to determine the degree of external referenti- in Marler 1985). Regardless of whether they are ality in vocalizations, it is necessary to study both externally referential or not, both types of variable ‘production specificity’ and ‘perception specificity’ alarm calls are referred to as ‘situationally (Marler et al. 1992; Macedonia & Evans 1993; Blumstein & Arnold 1995). If stimulus type Correspondence: D. T. Blumstein, Department of System- uniquely covaries with the vocal response, there is atics and Ecology, University of Kansas, Lawrence, KS a high degree of production specificity. Thus, if 66045, U.S.A. (email: [email protected]). yellow-bellied marmots have highly referential 0003–3472/97/010143+29 $25.00/0/ar960285 ? 1997 The Association for the Study of Animal Behaviour 143 144 Animal Behaviour, 53, 1 communication, they should, for instance, have neither Waring nor Davis conducted playback uniquely different ‘raptor’ and ‘canid’ calls that experiments to experimentally document marmot should only be produced in response to raptors responses to call variants, the question of how and canids, respectively. Perception specificity yellow-bellied marmots produce and perceive situ- means that acoustic variants (e.g. raptor calls ationally specific vocalizations remains unclear. versus canid calls) should elicit the appropriate In this paper, we describe the means whereby response in a conspecific who hears the call in yellow-bellied marmots produce and perceive situ- absence of the stimulus that normally elicits the ationally specific alarm calls. We used a combi- call and without other contextual cues associated nation of natural observations and simple field with alarm calling (reviewed in: Cheney & experiments to study factors that influence alarm- Seyfarth 1990; Marler et al. 1992; Evans et al. call production (part 1) and marmots’ responses 1993; Macedonia & Evans 1993). to alarm-call variants (part 2). Throughout, we There is some question about the degree to address the fundamental assumption of whether which yellow-bellied marmots produce externally intraspecific variation exists in the way a species referential alarm calls. Waring (1966) described encodes situational information. the vocal repertoire of yellow-bellied marmots and published spectrograms of several of their vocaliz- ations. He suggested that marmots varied the rate PART 1: ALARM-CALL PRODUCTION of their most common alarm call, their ‘primary whistle motif’, as a function of risk, not predator Methods type. More recently, Davis (1991) performed a more detailed micro-structural analysis of yellow- Study sites and subjects bellied marmot alarm calls. Davis concluded, based on a multivariate discriminant function Marmots (genus Marmota, ca 14 species) are analysis, that yellow-bellied marmots had predator- large (3–5 kg), obligately hibernating, moderately specific calls. Of 23 variables he examined, the to highly social, ground-dwelling sciurid rodents maximum frequency at peak amplitude was the (Barash 1989; Bibikow 1996). We studied alarm variable best able to discriminate predator type (at communication by yellow-bellied marmots at best, discriminant functions were able to correctly three locations during most or parts of two sum- classify only 37–65% of the calls to one of four mer active seasons for 745 h at three study lo- eliciting stimuli). Davis did not point out that cations. We studied marmots for 147 h in 1994 (29 recorded peak amplitudes might covary with June–20 July) and 415 h in 1995 (7 June–29 July) sound intensity (given that higher frequencies are in and around the Rocky Mountain Biological more likely to attenuate) and may also vary with Laboratory, Colorado, U.S.A. (RMBL; site de- body size (larger animals produce lower frequency scription and marking and census methods in alarm calls). Intensity variation alone is not an Armitage 1991); in 1995 we also studied marmots ideal mechanism to encode information about for 154 h (1–29 May 1995) in Capitol Reef predator type because perceived intensity, and National Park (Torrey, Utah, U.S.A.), and for 29 h therefore perhaps peak frequencies, would be in- (14–20 August 1995) in the City of Boulder Open fluenced by the distance and relative orientation in Space Parks (Boulder, Colorado, U.S.A.). Unless space between the signaller and perceiver. Thus, otherwise indicated, quantitative results presented perceivers at different distances from the caller below excluded the Boulder observations. might potentially draw different inferences about At RMBL, we studied 60 different individually the type of predator present. Similarly, if fre- identified non-pup marmots (38 in 1994, 31 in quency was a function of body size, marmots 1995) from 11 different social groups in the sub- would have to know the identity of the signaller to alpine East River Valley (7 groups in 1994, 6 properly assess the meaning of a signal. Recently, groups in 1995), and about 17 unidentified indi- Blumstein & Daniel, in press noted that yellow- viduals about 10 km away living in a higher alpine bellied marmots live in environments with particu- meadow (North Pole Basin). Yellow-bellied mar- larly poor acoustics, and that micro-structural mot habitat patches are typically described as alarm-call variants might not be transmitted well ‘colonial’ or ‘satellite’ (Armitage 1991). Colonies enough to be differentiated by perceivers. Because consist of ‘. one or more males, resident Blumstein & Armitage: Marmot alarm calls 145 Table I. Known predator kills of yellow-bellied marmots in the East River valley (Gothic, Colorado, U.S.A.): 1962–1994 Predator Young Yearling Adult Total Coyote, Canis latrans 1182140 Badger, Taxidea taxus 34 4 11 49 Eagle, Aquila chrysaetos 10 3 13 Bear, Ursus americanus 224 Marten, Martes americana 213 Long-tailed weasel, Mustela frenata 11 2 Red fox, Vulpes vulpes 22 Domestic dog, Canis familiaris 11 females, usually yearlings (animals one year old), frequency of occurrences varied from 11 to 35% and young (animals <4 months old)’ but satellite (Van Vuren 1991; R. A. Powell, unpublished sites contain a single female with her offspring data). For a cohort of yearling marmots (Armitage 1991, page 381). Colonies contained implanted with radiotransmitters, coyotes, Canis one or more ‘social
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