Exploring the Potential Use of Seismic Waves As a Communication Channel by Elephants and Other Large Mammals1
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AMER.ZOOL., 41:1157±1170 (2001) Exploring the Potential Use of Seismic Waves as a Communication Channel by Elephants and Other Large Mammals1 C. E. O'CONNELL-RODWELL,2*L.A.HART,² AND B. T. ARNASON³ *Center for Conservation Biology, Department of Biological Sciences 371 Serra Mall, Stanford University, Stanford, California 94305-5020 ²Department of Population Health and Reproduction, University of California, Davis, California 95616 ³Tezar Inc., P.O. Box 26235, Austin, Texas 78755-0235 SYNOPSIS. Bioseismic studies have previously documented the use of seismic stim- uli as a method of communication in arthropods and small mammals. Seismic signals are used to communicate intraspeci®cally in many capacities such as mate Downloaded from https://academic.oup.com/icb/article/41/5/1157/343557 by guest on 30 September 2021 ®nding, spacing, warning, resource assessing, and in group cohesion. Seismic sig- nals are also used in interspeci®c mutualism and as a deterrent to predators. Al- though bioseismics is a signi®cant mode of communication that is well documented for relatively small vertebrates, the potential for seismic communication has been all but ignored in large mammals. In this paper, we describe two modes of pro- ducing seismic waves with the potential for long distance transmission: 1) loco- motion by animals causing percussion on the ground and 2) acoustic, seismic- evoking sounds that couple with the ground. We present recordings of several mammals, including lions, rhinoceroses, and elephants, showing that they generate similar acoustic and seismic vibrations. These large animals that produce high amplitude vocalizations are the most likely to produce seismic vibrations that prop- agate long distances. The elephant seems to be the most likely candidate to engage in long distance seismic communication due to its size and its high amplitude, low frequency, relatively monotonic vocalizations that propagate in the ground and have the potential to travel long distances. We review particular anatomical fea- tures of the elephant that would facilitate the detection of seismic waves. We also assess low frequency sounds in the environment such as thunder and the likelihood of seismic transmission. In addition, we present the potential role of seismic stimuli in human communication as well as the impact of modern anthropogenic effects on the seismic environment. INTRODUCTION brations of the earth substrate (Ewing, Bioseismic cues are known to be impor- 1989). tant for many arthropods (Cocroft et al., Vibration signal energy depends mostly 2000), ®sh, reptiles, amphibians and small on the mass and available muscular power of the signal producer (Markl, 1983). The mammals in intraspeci®c and heterospeci®c source signal intensity and attenuation dur- communication, prey detection and predator ing transmission, together with the sensitiv- avoidance and navigation (see O'Connell- ity and depth of receptors in the receiver, Rodwell et al., 2000 for review). Two pri- and the threshold at which the receptor will mary methods of initiating bioseismic cues be stimulated relative to the frequency and are: 1) percussion that causes an impact strength of the stimulus de®ne the spatial with the earth and produces waves in re- extent of vibration signals. The Weber- sponse to direct contact and 2) vocaliza- Fechner law states that the magnitude of an tions which produce wave movements that observer's psychological response is direct- are then coupled with the earth to cause vi- ly related to the logarithm of the intensity of the stimulus (Landing et al., 1998). Sig- nal detection theory (SDT) further stipu- 1 From the Symposium Vibration as a Communi- lates that detection also depends on the ex- cation Channel presented at the Annual Meeting of the Society for Integrative and Comparative Biology, 3±7 pectation, motivation, in situ conditions, January 2001, at Chicago, Illinois. sensitivity, decision making and ®nally, 2 E-mail: [email protected] noise level (Tanner and Swets, 1954). 1157 1158 C. E. O'CONNELL-RODWELL ET AL. It appears there is a ``sweet zone'' for humans may also have used seismic cues at seismic signal transmission ranging from 10 one time as a means of long distance com- Hz to 40 Hz, where there is a maximum munication and prey detection. ef®ciency of transmission of seismic energy (O'Connell-Rodwell et al., 2000). Ambient Seismic vibrations produced by percussion seismic noise on land from ocean waves Foot-drumming of banner-tailed kanga- creates peaks at about 0.14 Hz and about roo rats (Randall, 1989) and the chela 0.07 Hz (White, 1965). With increasing fre- drumming of the male ®ddler crab (Aicher quency, these low frequency and storm mi- and Tautz, 1990) are percussion-induced croseisms sharply decline to negligible lev- seismic signals. Markl (1983) suggests that els by 10 Hz. Although noise due to micro- drumming-induced communication is a seisms decreases to trivial levels above 10 close-ranged communication system. More Downloaded from https://academic.oup.com/icb/article/41/5/1157/343557 by guest on 30 September 2021 Hz, the attenuation of seismic pulses in- recent studies of the Cape mole rat suggest creases with frequency (Frantii et al., that seismic signals produced by drumming 1962). Pre-historically, the range around 20 propagate at least an order of magnitude be- Hz was a quiet seismic region, carrying yond acoustical signals (Narins et al., 1992, only vibrations associated with thunder and 1997), providing evidence that seismic sig- earth tremors making it available to ele- nals might also be used in long distance phants and other large mammals. communication. Both acoustic and seismic waves are sub- Large terrestrial mammals inevitably ject to interference and alteration due to en- cause far greater impact on the seismic en- vironmental factors. Wind shear and tem- vironment than the invertebrates and small perature gradients in¯uence the acoustic mammals. Trunk banging displays of the propagation of sound, whereas the soil type Asian elephant produce a booming sound and heterogeneity are among the factors in- heard for a great distance (Tennent, 1867; ¯uencing the propagation of a seismic sig- Sanderson, 1878). The sound seems to be nal (O'Connell-Rodwell et al., 2000). Air- produced by the sudden percussion of the borne sound waves spread spherically rath- column of air in the trunk as it is expelled er than cylindrically, attenuating more rap- (Krishnan, 1972). A female may drum her idly than ground surface waves such as trunk following the birth of a calf (Vincent, Rayleigh waves, losing 6 dB for every dou- 1946), when a musth bull joins the herd bling of distance as opposed to 3 dB. (LAH, unpublished observations), as a There is also an outer limit to airborne threat to an intruder (O'Connell-Rodwell, transmission (Uman, 1984) which is not the unpublished observations), or even when case for surface seismic waves. In this pa- testing the soundness of a bridge (Baker, per, we address the production of both 1890/1988). acoustic and seismic waves by large terres- An elephant seal lying on the ground re- trial mammals, especially Asian and Afri- sponds to a seismic wave caused by drop- can elephants, that are known to produce ping an object at a distance of 20 m (Shi- high amplitude acoustic vocalizations. pley et al., 1992). Although the psycho- The primary aim of this paper is to pre- physical parameters have not yet been sent a conceptual framework for examining worked out, perhaps these animals obtain bioseismic cues produced by large verte- seismic information about the size and brates through percussion and the coupling strength of an opponent in an episode of of low frequency vocalizations, especially con¯ict. where such signals might be used for long The locomotion of large mammals pro- distance communication. We review what is duces ground-borne vibrations. An elephant known about biological sense organs that mock charge ends in a foot stomping be- could potentially be used to detect seismic havior that produces a substantial seismic signals and discuss the data that support the signal, modeled to be capable of traveling possibility of the elephant being capable of up to 32 km (O'Connell-Rodwell et al., detecting the seismic signals produced by 2000). The seismic energy generated by a conspeci®cs. We address the possibility that stampede of bison was apparently detect- SEISMIC WAVES AS A COMMUNICATION CHANNEL 1159 able by Native Americans and used as a phone, with a ¯at response of 20±20,000 form of prey detection. A herd of zebra or Hz. Seismic signals were recorded using a giraffe running may propagate a series of Mandrel 10 Hz MD-79 vertical polarized seismic waves with characteristics unique geophone with a transduction coef®cient of to that species. The characteristic gaits of 0.230 V/cm/sec. Geophones were buried 10 four-legged animals relate to the size and cm into the ground. structure of the body and differ with species The lion roars were recorded at approx- (Hildebrand, 1995). The natural period for imately 300 m. Rhino vocalizations were a walking elephant is 1.6 to 2.2 sec, for the recorded at approximately 100 m and the horse, 1.2 to 1.8 sec, and for the deer, 0.8 African elephant rumble was recorded at 20 to 1.0 sec (Hildebrand, 1985), each setting m. The Asian elephant rumble was recorded in motion a characteristic seismic wave. In from a female at 5 m. The thunder was re- Downloaded from https://academic.oup.com/icb/article/41/5/1157/343557 by guest on 30 September 2021 theory, a seismic eavesdropper, such as a corded at varying distances approximately lion on the African plain, could assess 1 km away. whether vibrations are from antelope or ze- The seismic components of vocalizations bra, making more selective hunting forays were ®ltered and ampli®ed separately from for preferred food.