NON-SONG PHONATIONS AND ASSOCIATED SURFACE BEHAVIOR OF THE HAWAIIAN HUMPBACK , MEGAPTERA NOVAEANGLIAE

A Thesis

Presented to

The Faculty of the Department of Biological Sciences

San Jose State University

In Partial Fulfillment

of the Requirements for the Degree

Master of Arts

By

Gregory K. Silber

May, 1986 ABSTRACT

Humpback whale (Megaptera novaeangliae) non-song phonations were recorded off West Maui, Hawaii, in the winters of 1981, 1982 and 1984. Non-song phonations or social sounds, differed from humpback song in acoustic structure and behavioral context. Social phonations were produced by large socializing whale groups which exhibited a great deal of surface activity and dramatic aerial displays. It was theorized that social phonations were associated with competition among humpback males because: 1) intraspecies aggression was observed only in large active groups, and was absent in other groups, 2) social phonations were produced at high rates in large active groups, and very rarely among non-aggressive groups, and 3) aggression in large active groups has been attributed to male competition for a sexually mature female. A catalog of the sound types was prepared. A descriptive analysis of the sounds, the social groups in which they occur and associated surface behavior was presented. The relative use of individual sound types, sound patterns and individual behavior types was quantified. Surface activity rates were not significantly different in active groups of varying size. However, phonation rates differed with group size, and rates increased dramatically when a new member joined a competing group. Although the sounds may serve a variety of functions, it is likely that the majority of sounds acted in conjunction with visual displays to communicate threats and to aid in the establishment and maintenance of a social dominance among males competing for proximity to a female.

iii TABLE OF CONTENTS

Page

Abstract iii

List of Figures ...... V List of Tables ...... vl List of Appendices vii Acknowledgements ...... viii

Introduction 1

Materials and Methods ...... 4

Results ...... 7 Discussion 13

Literature Cited 20

Appendix 48

iv List of Figures

Figure 1. Example of technique used to measure sound characteristics with digitizing tablet.

Figure 2. Sample of song structure.

Figure 3. Social phonation tracings illustrating sound spacing and association with surface activity and group dives.

Figure 4. Frequency of occurrence of each call type in the humpback whale social phonation repertory.

Figure 5. Frequency of occurrence of various sound patterns.

Figure 6. Mean phonation rate versus group size (#/hr).

Figure 7. Mean phonation rate per individual (#/whale/hr).

Figure 8. Frequency of occurrence of surface behavior types in active humpback groups.

Figure 9. Mean surface activity rate versus group size (#/hr).

Figure 10. Mean surface activity per individual (#/whale/hr).

Figure 11. The number of phonations versus changes in group size, Group B, 3/1/82.

Figure 12. The number of phonations versus changes in group size, Group I, 3/20/82.

v List of Tables

Table 1. Social sound catalog.

Table 2. Humpback whale groups monitored for sound.

Table 3. Group phonation rate versus activity level.

Table 4. The difference in phonation rate before and after a joiner enters a surface active group.

vi Appendices

Appendix 1. Surface active groups from which underwater recordings were made 1980, 1981.

Appendix 2. Surface active groups from which recordings were made, 1982.

Appendix 3. Surface active groups monitored for sound from which no sounds ere heard, 1981, 1982.

Appendix 4. Cow/calf and cow/calf/escort groups monitored for sound, 1981, 1982.

Appendix 5. Solitary non-singing adults monitored for sound, 1981, 1982.

Appendix 6. Groups containing two adult which were monitored for sound, 1981, 1982.

Appendix 7. Cow/calf and cow/calf/escort groups observed but not monitored for sound, 1981, 1982.

Appendix 8. Characteristics of social sound types in the acoustic repertory of the humpback whale.

Appendix 9. Phonation rate versus surface display rate.

vii ACKNOWLEDGEMENTS

I would not have had the opportunity to study the humpback whale if it had not be~n for the encouragement and constant support of Jim Darling, Roger Payne and Katy Payne. Trish Cutler and Beth Mathews contributed immeasurably to the field work. Drs. John Ford, Chris Clark and Peter Tyack contributed equipment and advice during analysis. My committee members Bernd Wursig, Ken Norris, John Oliver and Randy Wells provided advice and critical reading of various drafts of the thesis. My advisor Bernd Wlirsig has given me invaluable support throughout all phases of the project and he deserves special thanks. The Moss Landing Marine Laboratories created an outstanding learning environment for which I will always be grateful. Financial support was provided by the American Cetacean Society (Los Angeles and Monterey Bay Chapters), the Packard Foundation, Sigma XI, the Maui Whale Watchers, the New York Zoological Society, the West Coast Whale Research Foundation and Ms. Sylvia Shugrue. The 1981 season was supported in part by the National Geographic Society, the Vancouver Aquarium, the World Wildlife Fund, the Clifford E. Lee Foundation and the New York Zoological Society. The thesis is dedicated to Robert L. and Wanda M. Silber who always encouraged excellence, but never demanded it.

viii 1.

Introduction

In winter months, humpback whales (Megaptera novaeangliae) aggregate in waters surrounding the

Hawaiian islands. During this period the humpback produces a complex series of sounds defined as "song"

(Payne and McVay, 1971). The humpback emits another very different series of sounds which are called "social sounds" (Payne, 1978; Tyack, 1982). These phonations differ from song in several ways. Unlike song, social calls are produced in both summering and wintering areas

(Thompson and Kennison, 1977; Payne, 1978). Songs are rhythmic and continuous (Payne and McVay, 1971) and singers are lone, stationary adult whales (Payne, 1982;

Tyack, 1982). In contrast, social phonations are erratic and sound series possess no known pattern.

Although some authors briefly described humpback social vocalizations (Payne, 1978; Tyack, 1982; Tyack,

1983), no study has quantified the occurrence of the sounds, described them in detail or given an account of the behavioral contexts in which they occur.

Individual humpbacks frequently interrupted activities such as singing to travel considerable distances to join groups that were producing social sounds (Tyack, 1983). Likewise, humpbacks were strongly 2. drawn toward social sound stimuli during playback experiments (Tyack, 1983).

Several authors characterized the structure, composition, and behavior of humpback surface active groups (Glockner-Ferrari and Ferrari, 1981 and 1983;

Darling et. al., 1983; Tyack, 1983; Tyack and Whitehead,

1983; Baker and Herman, 1984). Darling et.al. (1983),

Tyack and Whitehead (1983) and Glockner-Ferrari and

Ferrari (1983) described aggression between competing individuals. Tyack and Whitehead (1983) and Baker and

Herman (1984) concluded that agonistic encounters, manifested as dramatic surface displays and collisions between individuals in large active groups, resulted from male-male competition for a sexually mature female.

The data presented here were derived from three field seasons of study on the underwater acoustics and associated behavior of humpbacks wintering in Hawaiian waters. My objective was to document and quantify the use of non-song vocalizations by wintering humpbacks. I have: 1) presented a catalog of humpback social phonations, 2) provided a descriptive analysis of the sounds and an account of the frequency of occurrence of each sound type, 3) established the relative usage of the sounds by various social group types, and 4) presented an analysis of the relative occurrence of 3. individual surface activities that were associated with social sounds. In addition, I have advanced a hypothesis concerning the function of the vocalizations. 4. Materials and Methods Social sounds were recorded off West Maui, Hawaii in January - April, 1981, February - April, 1982, and March, 1984. Behavioral observations and underwater recordings were made from small craft. A Nakamichi 550 portable

cassette tape recorder was used in conjunction with a

Barcus Berry preamplifier, and a single omni-directional hydrophone of the following makes: Gould CH-17U, Aquadyne

AQ-17 or U.S. Navy Sonobuoy. The hydrophones were deployed at depths ranging from 10-16 meters, depending

upon the individual hydrophone in use. The entire system had a frequency response of 50-10,000 Hz ~ 5 dB.

Underwater recordings were made by "leapfrogging" in advance of moving whale groups. We positioned our boat

in front of a group and recorded as the whales approached

our location. All observations and recordings were made

when the whales were within an estimated 250 m of the

craft. The whale groups studied were: 1) lone non-singer = single adult, not singing, 2) two adults, 3) cow and calf = single adult with calf of the year, 4) cow, calf and escort = two adults and a calf of the year, 5) surface active group = three or more adults moving rapidly and exhibiting a great deal of surface activity.

Classes 1 through 4 were generally not active at the

surface and behavioral data from those groups were not

used in determining activity rates. 5 •

Each surface activity other than normal respiration was tabulated as a single event. Activities included: breach, tail slap, flipper slap, underwater blow, etc.

All data were divided into five minute intervals.

Phonation rates were determined after dividing all five minute intervals into three levels of activity using the

following criteria: LOW = 0-12 surface events/whale/hour; MID = 13-32 events/whale/hour; HIGH = 33-54 events/whale/hour.

A social sound was defined as any phonation that did

not possess the rhythmic and continuous pattern of song.

A phrase was defined as one or more sounds surrounded by

at least ten seconds of silence. A sample of 5,433

clearly audible sounds was used to determine phonation

rates. Sounds were divided into 25 categories, by spectral analysis and judgement of aural quality. A

selection of 147 sounds that represented all 25

categories was derived from the entire sample, and

spectral analysis was performed on a Kay Spectrograph

Model 7029A. Depending upon the frequency range of the

individual sound, spectrograms were made with 22.5, 45.0,

90.0 and 180.0 Hz filter-band widths, which permitted

analysis from 20-2000, 40-4000, 80-8000 and 160-16,000 Hz

respectively. Sound structure was measured by a hand digitizing tablet coupled with an Apple II microcomputer. 6 .

A standardized acetate overlay was positioned over a spectrogram by placing the horizontal axis on the zero frequency line. The vertical axis was placed at the point where the first sound energy occurred. The digitizing wand was touched to predetermined points on the overlay, where each point represented a specific parameter (FIG. 1) (Ford, 1984). The following measurements were made: duration; maximum sound energy; maximum frequency of the concentrated (e.g. fundamental) energy; and side band (or harmonic) measurements, including a value for the highest and lowest frequency at the beginning, middle and end of a selected side-band.

A Model I analysis of variance (ANOVA) (Zar, 1974) was used to determine the significance of differences in phonation and surface activity rates relative to group

size. The same test (ANOVA) was used to determine the

significance of differences in phonation rate versus activity levels. A Student Newman-Kuels test (Zar, 1974) was used to determine if significant differences existed in the vocalization rate between groups of differing

size. A Spearman rank correlation (Zar, 1974) was used to determine the relationship between vocalization rates and group size. 7.

Results

I conducted behavioral observations and monitored underwater acoustics for 153 whale groups (Appendicies 1-7). During this period 18.3 ohours of underwater recordings were obtained which were analyzed for phonation and surface activity rates.

There were fundamental differences between humpback social sounds and that which is known about song. Unlike the predictable recurring cadence of song (FIG. 2), social phonations occurred singularly or in sudden bursts of sound surrounded by periods of silence, and they possessed no apparent consistent sequencing or pattern

(FIG. 3). Social calls were heard in conjunction with dramatic surface displays in large groups. Song is generally attributed to stationary and relatively inactive single adults (Payne, 1982; Tyack, 1982; and others). Furthermore, while song undergoes gradual change (Payne et al., 1983), social sounds did not appear to change in any consistant pattern throughout several seasons.

Qualitative observation indicated that some social calls were also used as part of the song of that year, although a detailed analysis was not conducted. The humpback social sound repertory was diverse in frequency 8. range, duration and number of call types (TABLE 1,

Appendix 8). Many calls were stereotypic while others exhibited considerable variabiTity. Some of the sound types that are represented here as being variable (TABLE

1), may be viewed as a part of a continuum rather than discrete units. The sounds ranged in duration from less than 0.25 seconds to over 5.00 seconds. They exhibited a frequency range from 50.0 Hz to 10.0 kHz. The majority of sound energy was below 3.0 kHz and most of the

"fundamental" or concentrated energy was below 2.0 kHz.

Sounds with simple structure, particularly frequency-modulated upsweeps, occurred most often (FIG.

4) •

There was considerable variation in the number and type of sounds used in sequences or phrases (FIG. 5).

The most common format was a single sound, preceded and followed by at least 10 seconds of silence. In addition, sequences of similar or unlike sounds were strung together in rapid succession (FIG. 5). A series of 20 or more sounds were heard, although strings containing from two to ten sounds were more common. Phonations in a series generally possessed brief intercall periods and were short in duration. In contrast, more complex and longer duration sounds were most common in series which contained several sound types. Some sound types were 9.

always heard together. Sound types 1,2,3,14,16,17,21 and

23 (Table 1) were often produced in rapid succession without interspersion of dissimilar sounds. The remainder of the calls were heard singularly or combined

with other unlike sounds. It was common to hear whales

phonating simultaneously (FIG. 5). In these instances multi-unit phrases were employed.

Social phonations were heard almost exclusively in

large surface active groups (TABLE 2) at an overall mean rate of 43.1 + sd 55.52/whale/hour. Likewise, surface

behavior that was interpreted as intraspecies aggression was observed only in surface active groups. Social calls were rarely heard in groups that contained less than

three adult whales or in socializing but non-aggressive

groups. Although sounds were not heard in some groups,

we cannot conclude that those groups were always silent.

While samples were too small to obtain phonation rates

for all group sizes, these data indicate that larger

groups phonate at much higher rates than do smaller

groups.

The phonation rate differed significantly in active

groups of different size (F=5.27, p<0.005), and there was

a significant positive correlation between vocalization

rate and increasing group size (r=0.829, 0.05

6). There was a significant difference in vocalization 10. rate per individual relative to group size (F=2.52, p<0.05), and there was a slight (not significant) correlation between individual vocalization rate and increasing group size (r=0,029, p<0.05) (FIG. 7).

Phonation rates appeared greatest in groups that were most active at the surface, but the differences were not significant (F=0.994, O.SO

Qualitative observations indicate that social sounds were often heard just prior to and during group surfacing.

Whales in large groups performed a wide variety of surface actions (FIG. 8) that were readily observed.

Surface displays were produced at a rate of 11.7 ± s.d. 11.50/whale/hr. There was no significant difference in the mean surface display rate relative to group size

(F=O.l25, p

Likewise no significant difference existed between displays per individual with group size (F=0.0052, p

(r=-1.00, p=O.Ol) with increasing group size (FIG. 10).

A large percentage of the surface activities, particularly those involving the tail flukes, were clearly directed at other group members and some resulted in actual physical contact. We observed whales striking 11. each other and intentionally butting and colliding with other group members. Many of these activities resulted in severely chafed skin and open wounds along a whale's dorsal fin, tail flukes and head. These actions and various other displays were interpreted as intraspecific aggression. Incidents which involved conspecific body contact and striking blows with tail flukes were clearly audible underwater (sound #25) and this type of sound was commonly heard (FIG. 4). These sounds were high amplitude, rapid onset and broadband, and possessed a rasping quality.

All social groups were highly fluid and group sizes, particularly large groups, changed frequently. The mean size of active groups was 5.5 ~ s.d. 1.51 whales (range = 3- 20; n = 47) (Appendices 1-3). It was common to observe whales joining an active group. Phonation rates often increased dramatically after joining occurred

(TABLE 4). Two incidents serve as good examples of this pattern. They illustrate a dramatic increase in phonation rate with the addition of another whale, followed by a decrease in the number of sounds even though group size remained constant (FIGS. 11 and 12).

Whales that departed from active groups often began to sing. On two occassions singing was heard very near an 12. active group although it was not determined if the singing whale was actually in the group.

Whales in active groups g~nerally surfaced synchronously. Active groups spent 51.2% of the time at the surface, which is much higher than qualitative observations of other group types, such as cow/calf and cow/calf/escort groups. Nine (21.4%) surface active groups contained calves. 13.

Discussion

Social sounds played an important role in complex social interactions involving whales in large groups.

The sounds were closely linked to aggression, both of which occurred almost exclusively in large active groups.

Visual displays were used with acoustic signals as males threatened each other. The diversity and complexity of the acoustic repertory suggests that the sounds may have been graded to indicate levels of excitement or agitation between interacting humpbacks. Social vocalizations occurred in behavioral contexts that differed dramatically from that of the song, indicating that the two types of phonations served very different communicative functions.

Glockner-Ferrari and Ferrari (1981) and Darling

(1983) showed that humpback active groups consisted of multiple males and one or more females. Baker et al.

(1981), Tyack (1982), Tyack and Whitehead (1983), and

Baker and Herman (1984) concluded that activities in large active groups resulted from the persistent and aggressive attempts of numerous males to gain access to a position near a female. Baker and Herman (1984)

concluded that aggression in large groups resulted from male-male competition for a sexually mature female and

documented the occurrence of butting and body contact by 14. males. The high-amplitude underwater sound of whales striking or colliding with a conspecific (TABLE 1) support the contention that agonistic encounters occured in active groups, as males jostled and dueled for position. Darling et al. (1983) and Glockner-Ferrari and

Ferrari (1983) described several separate incidents in which a peripheral male successfully displaced another male in its role as the principal escort to the female.

Social phonations were heard almost exclusively in large active groups and only very rarely in other groups

(TABLE 2). Likewise, whales in large active groups were highly aggressive, which was absent in other social groups. It is likely therefore, that many of the sounds acted as critical elements in male agonistic behavior.

Social phonations probably served to threaten or warn another group member about the likelihood of attack, or

conversely they circumvented an attack by signifying low

status. Many sounds were low frequency, harsh and

guttural in character and they may have been produced by dominant animals to intimidate or frighten an opponent.

Similarly there were high frequency chirps and twitters that subordinate individuals emitted to appease a dominant. This generalization about acoustic

communication has been described in ungulates (Kiley,

1972), and a wide variety of birds and mammals (Collias, 15.

1960; Morton, 1977}. In addition, the sounds may have announced the location or identity of a competitor.

Frequency-modulated sounds are easily located (Marler,

0 1957}, and many of the humpback calls posessed this quality. I believe that multi-call phrases (which exhibited variations in the number of repetitions} (FIG.

5} were used to convey information about the relative level of aggression or agitation. Calls that were highly variable in duration or frequency, may also have been graded for relative level of excitement. Many calls were highly complex in harmonic structure and frequency range.

Clark (1983} found that (Eubalaena australis} sound types exhibited greater complexity as group structure became more complicated. A similar pattern may be occurring in humpback active groups. The occurrence of two or more group members phonating simultaneously

(FIG. 5}, particularly in multi-sound sequences, suggests that two whales were producing acoustic threats concurrently.

The vocalization rate was positively correlated to group size (FIGS. 6 and 7}, indicating that the number of acoustic threats increased relative to the number of males in the group. A greater number of individuals phonating simultaneously was heard following the

occurrence of whales joining a large group than prior to 16. joining, indicating that threats were produced concurrently. Likewise, the relative number of sounds produced by each individual was correlated to increasing group size, suggesting that each group member contributed to overall sound production. In some groups the vocalization rate increased dramatically with the addition of new whales (FIGS. 11 and 12}, supporting this contention. However, the techniques used in this study

(e.g. a single omni-directional hydrophone} precluded the determination of which individual in the group was producing the sounds. Therefore it was not possible to assess if a single whale (e.g. principal escort} increased its phonation rate or if more sounds were heard as a result of the increased group size.

Visual displays were important to male-male

competition (Baker and Herman, 1984} which accounted for

the occurrence of observed surface activites in large

groups (FIG. 8}. It is likely that social sounds were

used in association with visual displays, to heighten the

effect of a threat. The simultaneous use of two or more

signalling pathways is common behavior in a wide variety

of social species (Smith, 1977}, and it has been

described in canids (Lehner, 1968}, felids (Marler, 1961}

and ungulates (Kiley, 1972}. 17.

The surface activity rate {FIG. 9) and activity rate per individual {FIG. 10) were negatively correlated to group size, although the latter exhibited a stronger correlation. These data differ f,rom previous reports by

Herman {1977), Tyack {1982) and Tyack and Whitehead

{1983) which indicated that activity increased proportionally with group size. The negative trend

corresponding to increasing group size suggests that one

individual produced the majority of the displays at a

relatively constant rate that was independent of group

size. It is likely that the principal escort was most active and engaged in threats directed at periphery

escorts, to maintain its location near the female.

Glockner-Ferrari and Ferrari {1983), Tyack and Whitehead

{1983) and Baker and Herman {1984) concluded that most of

the surface activity could be attributed to the principal male escort. The data presented here and my qualitative

observations are in agreement with those studies.

The fluidity of large groups may be attributed to

intraspecific competition and/or dominace hierarchies

within the group. Single whales and pairs traveled

considerable distances to join large active groups

{Tyack, 1983; Tyack and Whitehead, 1983; pers. obs.). We

observed whales that interrupted activities including

singing, to swim rapidly toward active groups. The 18.

social phonations drew whales toward the group and localization of the group was almost certainly mediated acoustically. Sound playback experiments indicated that social vocalizations provided a very strong attracting

stimuli (Tyack, 1983). Males in the surrounding area were presumably drawn to the sounds associated with

competition so they might enter the group and endeavor to

secure a position near the female. Qualitative

observations reveal that whale densities, particularly singing whales, were lower in areas where a surface

active group had recently passed. This suggests that the

group "collected" whales as it progressed through a given

area. In addition, it is probable that some of the whales that departed from active groups were those that

were displaced during competition. Singing whales are

generally believed to be males (Winn et. al., 1973;

Darling, 1983). Therefore it is interesting that

departing whales often began to sing. It is likely that

departing whales resorted to a different strategy

(singing) to attract females as suggested by Tyack (1982)

or to communicate to other males as proposed by Darling

(1983). The addition of joining whales may have

disrupted an established dominance structure within the

group and therefore resulted in an increased number of phonations and acoustic threats (FIGS. 11 and 12, TABLE

4). Presumably, the phonation rate decreased as the 19. hierarchical structure of the group was re-established, and the role of the principal escort clearly defined.

Humpbacks produce social sounds in the northern

latitudes (Thompson and Kennison, 1977), where the whales are engaged primarily in feeding during the summer months

(Jurasz and Jurasz, 1979; and others). Aggression is

rare during that period (Baker and Herman, 1984) and the

sounds undoubtedly serve functions that differ from the

Hawaiian contexts. In many other species the meaning of

a given signal is dependent upon the context in which it

is produced (Smith, 1968, 1977; Kiley, 1972). Both the

Hawaiian and Alaskan repertories may include sounds which

aid in detecting, localizing and maintaining contact with

distant individuals. However, additional data are needed

to more fully explore these hypotheses. 20.

LITERATURE CITED

Baker, SC, LM Herman and W Stifel, 1981. Agonistic behavior among humpback whales: evidence of male-male competition. Fourth Conf. on the of Marine Mammals, Abstracts, p. 7.

Baker, SC and LM Herman, 1984. Aggressive behavior between humpback whales (Megaptera novaeangliae) wintering in Hawaiian waters. Can. J. Zool. 62: 1922-38.

Beamish, P. 1979. Behavior and significance of entrapped baleen whales. In: HE Winn and BL Olla (eds.), Behavior of Marine Mammals - current perspectives in research - vol. 3: Cetaceans. New York, Plenum Press, p. 291-309.

Collias, NE, 1960. An ecological and functional classification of animal sounds. In: Lanyon, WE and Tavolga, WN (eds.) Animal sounds and communication. Amer. Inst. Biol. Sci., Pub No. 7, Washington, Pages 368-391.

Darling, JD, 1983. Migrations, abundance and behavior of Hawaiian humpback whales, Megaptera novaeangliae (Borowski). Unpubl. Doctoral dissertation, University of California, Santa Cruz, CA.

Darling, JD, KM Gibson and GK Silber, 1983. Observations on the abundance and behavior of humpback whales off West Maui, Hawaii, 1977-1979. In: RS Payne (ed.), Communication and behavior of whales. AAAS Selected Symposia Series, Westview Press, Boulder, CO., p. 201-222.

Ford, JKB, 1984. Call tradition and dialects in killer whales (Orcinus orca) in British Columbia. Unpubl. Doctoral dissertation, University of British Columbia, Canada.

Glockner-Ferrari, DA and M Ferrari, 1981. Correlation of the sex and behavior of individual humpback whales Megaptera novaeangliae to their role in the breeding population. Fourth Biennial Conf. on the Biology of Marine Mammals, Abstracts, p. 34. Glockner-Ferrari, DA and M Ferrari, 1983. Reproduction, aggression and sexual activites of the humpback whale. Fifth Biennial Conf. on the Biology of Marine Mammals, Abstracts, p.37.

Herman, LM, 1977. Humpback whales in the Hawaiian breeding waters: behaviors. KS Norris and RR Reeves (eds.), Report on a workshop on problems related to humpback whales (Megaptera 21. novaeangliae) in Hawaii. Marine Mammal Commission Report #77/03. Jurasz, CM and VP Juarasz, 1979. Feeding modes of humpback whales, Megaptera novaengliae, in Southeastern Alaska, USA. Sci. Rep. Whales Inst. Tokyo ~(31):69-84.

Kiley, M, 1972. The vocalizations of ungulates, their causation and function. z. Tierpsychol. 31:171-222.

Lehner, PN, 1978. Coyote vocalizations: a lexicon and comparisons to other canids. Anim. Beh. 26:712-722.

Marler, PJ, 1961. The logical analysis of animal communication, J. Theor. Bio., 1:295-297.

Marler, PJ, 1967. Animal communication signals. Science 157:769-774. Morton, ES, 1977. On the occurrence and significance of motivation-structural rules in some bird and mammal sounds. American Naturalist 111(981):855-869.

Payne, K, P Tyack and RS Payne, 1983. Progressive changes in the songs of humpback whales (Megaptera novaeangliae). In: RS Payne (ed.), Communication and behavior of whales, AAAS Selected Symposia Series, Westview Press, Boulder, co., p. 295-332.

Payne, RS, 1977. Behavior and vocalizations of humpback whales (Megaptera ~). In: KS Norris and RR Reeves (eds.), Report on a workshop on problems related to humpback whales (Megaptera novaeangliae) in Hawaii. Marine Mammal Commission Report #77/03.

Payne, RS, 1982. New light on the singing whales. National Geographic 161(4):463-477. Payne, RS and S McVay, 1971. Songs of humpback whales. Science 173:585-597.

Smith, WJ, 1968. Message-meaning analysis. In: Sebeok, TA (ed.) Animal Communication. Univ. Indiana Press, Bloomington, 44-60.

Smith, WJ, 1977. The behavior of communicating, an ethological approach. Harvard Univ. Press, Cambridge, MA.

Thompson, PO and SJ Kennison, 1977. Sound production of humpback whales, Megaptera novaeangliae, in Alaskan waters. J. Acoust. Soc. Am. 62(l):S89. 22.

Tyack, P, 1982. Humpback whales respond to the sounds of their neighbors. Unpubl. Doctoral dissertation, , NY.

Tyack, P, 1983. Differential response of humpback whales Megaptera novaeangliae, to playback of song or social sounds. Behav, Ecol. and Sociobiol. 13(49):49-55.

Tyack, P and H Whitehead, 1983. Male competition in large groups of wintering humpback whales. Behaviour 83: 132-154.

Watkins, WA, 1967. Air-borne sounds of the humpback whale, Megaptera novaeangliae. J. Mamm. 48(4):573-578.

Winn, HE, WL Bischoff and AG Taruski, 1973. Cytological sexing of cetacea. Marine Biology 23(4):343-346.

Zar, JH, 1874. Biostatistical Analysis. Prentice-Hall, Inc. Englewood Cliffs, N.J. 23.

FIGURE 1. Example of technique used to ~ake acoustic measurements with a digitizing tablet.

2.0

·~

@ ® r1.0

J-;Pasurements definina t¥.'0 sid~_::_c;!~ 1. Duration 2. Maximum eneray 5. 3eginninq - higher frequency 3. Minimum enerqy =· Beginning - lo~er frequency 4. Maximum of concentrated 7. Middle - higher frequency enerrry (e.q. fundamental e. Middle - lower frequency sideband max. freque~cy) 9. End - higl1er frequ11Cy 10. End- lower fr~guency NOT S'-lm-IN 11. Duration to next call in multiple unit sound 24.

'igure 2. Sample of humpback whale song structure. Song :racings illustrate phrases and continuous, rhythmic >atterns. From Payne, Tyack and Payne, 1983. 25.

MARCH 19, 1977 ..... -::-! ••• - -- .... !1.. - -1-.-- - _,_ ....,. - ~t--- -· I I I - ..r ••••• 1111- ~, .•.••••••• -- -~·······- -~.- •.:.:·.:,,- -, •• ~ ...... ------r=·------1-__ ,_, ------1- _____ ,:: __ -- ---

~------~r~_.._._;~·-·-· ___- __.4-l·_·_·~·r·-'-'-'-'--'"---'-k'_,_Jr'-'-,_'_,_,.~~----·-kl··,·~·*··''-'-' __ ..___ ·_·,-~:-_._._~_-_-_-_. ____ .._·r--·J_·_-, -1------1------1------... r~: .. - --- -·-,_~1·- '--' :_1.~~. 1•••. · L.. ~--·~ . L _ I___ I -1 --r -~r ~=I ~~~- --r ~~

jc:.:·=,. : '"='== i

MARCH 24,1976

~;-· . . l . ·;.•. ,. ... ~ .. ... ;_~1-!"•:('f\ .... ., ..... __ --- :1.-l"";.u .. .i~...:J:i:.ii:ll~- -I··.··~... - ...-.. - -- ~- .. - .. -~------::i ~ :lJ :Jj:.ij:J -- -~ j ~ J :.i z::: :-:J -::: -~ .. .:.::;.:.!;t!: --- - :.:. .. ~:.~":.u~ -- -

~ .., .. :. .. !.1 • ..:. ; ~-~!) -· ~ .., ...... ;..~J::. ..i:.~~J:. -~ ...... ------" :" "' -,- -I~ - .... -"1.-'lo-..-"""'- ..... , , , J.J.J-- .1 • ./ J - j ·-' """' -"' """" -"\ -"\ """' l- • ' r-· .J.J.Jl- - ·-' ·- .J --·,,-· - - - _, ----1------. ---1 - - J ----.1--- - -' -----1- --- _, ..... 1- --1

~ _, .... 1- --- ...,. 0 ••• - --- -J' -00 .1--- . - ~: _j:__:- - - _, ...... -- - -- ,__ r~-:: ...... j • . j ; :-.>' J J .,,. J --"'· I _.-;:. ~ i I J I I J J ..:..:.:::.; ~=;J7'"' • -·~ ....--,..,.,..·--:1· ~~ • ' ~-~-+ ._.J.. ____

- __;.,... - --: ·-·;[- ,,- - -~--~1- -. - ·- ==~!- ---·- I

! I • • .. " .. .. 26.

'igure 3. Social phonation tracings illustrating sound ;pacing and association with surface activity and group lives. Letter symbols denote individual surface activities !Xhibited by the group, 4 April, 1982. BS = bubblestream, :s = chin slap, HU = head-up, LB = loud blow, OM = open 1outh, PV = pivot, RO = roll, TL = tail lob, TOR = torpedo, JB = underwater blow, VF = vertical flukes, WW = white mter. 27 ,·

-04 APRIL 1982

:t;;~am ;~~g;; 't·•·· •.•.••••• :: :::r: ))})} JJ)J)J ~Jill J· ~

:;:;:;:::;:::::::;:;:::::::::::::::~::~:­ :;:;:;;:;::;:;:;:;:;:;:;:;:;:;:::::;:;:;:;. zo:so

15 30 45 60 75 90 105 120 TIME (SECONDS) 28.

~gure 4. Frequency of occurrence of each call type in the ~pback whale social phonation repertory. 29.

F R E Q U E N C Y 0 F O.C C U R R E N C E (%) 5 10 15 20 25 30

2

3 -:j J r---~~------~ 4 -:1 ~ 1--~ 5 -:l•· lf--,~...1 6 -:I c S1eond1 7 -:ldil 0 2.0 4.0 8 -:1= 9 -:luuwmJ

10

II UJ 0.. ,_>- 12

Q 3 z 14 ..0" 15

16

17

18

19

20.

21

22

23 .: I ~ 24 ' -- 25 ~I I 1------l 30.

igure 5. Frequency of occurrence of various sound atterns. 0 5 I I kHz Seconds

2+Anlmals Simultaneously :~ JJJ$JJ~ 1-

One Sound Type ~ J

0 10 20 30 40

FREQUENCY OF OCCURRENCE(%) 500 ( 12) t(39) 400 f (221 300 Mean Number or Sounds (# sounds/hr) 200 w f (73) t (68) .N t(6) 100

0•------+------r------~----~----~----__, 2 3 4 5 6 7 a+ Group Size(# whales)

Figure 6. The number or sounds relative to group size. Mean, one standard error and sample sizes are Indicated. 80.0

70.0 t(39l ( 12)

60.0

50.0 Mean number of (6) sounds 40.0 t (73) (#sounds/ (22) t w whale/hr) fc6a> 30.0 .w

20.0

10.0

0.0 3 4 5 6 7 a+ Group Size ("'whales)

Figure 7. The number of sounds produced per whale versus group size. Vocalization data have been standardized by the number of whales In each group. Mean, one standard deviation and sample sizes (N) are Indicated. 34.

'igure 8. Frequency of occurrence of surface behavior types n active humpback groups. 35.

>­

LL. 0 w a_ >- 1-

0 ,...,0 10 0 I!') 0 I!') 0 "' N N

25.0 (6)

20.0

15.0 Neon number or surface displays ("displays/whale /hr) · ~(60) 10.0 f(JB) W· -..!

5.0 f<21)

0.0 ·-----+-----+-----+------t------1 4 5 6 7 at Group Size (" whales) Figure 10. The meen number of surface displays per whale relative to group size. These data have been standardized by the number of whales In the group. Nean, one slendard error and sample size are Indicated. 38.

'igure 11. The number of phonations versus changes in group ;ize, Group B, 3/1/82. 39.

NUMBER OF WHALES

"-1 :::E 1-

0 0 ID 10

1'o'A~:3.LNI ::J.LnNIVIJ :3AI.::l ~3d SNOI.l'o'NOHd .::!0 ~38WnN 4 0.

Lgure 12. The number of phonations versus changes in group ize, Group I 3/20/82. 41.

NUMBER OF WHALES 0

It) .. '

It) \~======~==~ l'l'l

0 l'l'l -en 1.1.1 1- It) ::::> C\.1 :z :E 0 - C\.1 1.1.1 ::!: 1- It)

0

1'1i'A~3.LNI 3.LnNIII'.I 3AI~ ~3d SNOI.L'Ii'NOHd ~0 ~38WnN TABLE 1. Sound types in tho acoustic repertory o! the humpback whale. Tho sounds are grouped into pulsive and tonal subsections. Sound types which vary in duration are B. Between 0.5 - 3,0 seconds duration, described as variable and duration ranges are given. Sound types vhich possess stetootyped duration are ! SOHE CHARACTERISTICS described as discrete. FH ~ !tequency modulation. t. Pulslve variable duration (1,0-J.O secon~sl1 highly puls~d. but 6 A. Less than 0.5 seconds duration. tonal qualityr rich har~nicsr gradual onset, always heard singularly, unifor~ frequency ! EXAHPL£ SOHE CHARACTERISTICS 3 discrete, simple structure, kHz .. little or no FH, generally ~.;:::. heard in aeries of 2-J unital 3 :;::-:; - moat energy belov 1000 Hz 1<11: poppiny soundar variable dur- -~ ation up to 5,0 secondsl1 1 l DUltlple broadband units1 I~ variable (but generally lov} amplitude and variable rep- discrete, segmented, two-part etition rate sound, second segment lover Wlq in frequency; bath component& variable duration (l.D-5.0 ' ,.. as a aingle unit are often 8 seconds),broadbandl noisy, ~ used in series o! repeated sounds! very harsh, -breathy• sound- -"' most energy below 1000 Hz kH:l [3$0 ingl amplitude modulation! N always heard singularly

variable duration (0,5-2.0 discrete, simple structure• 3 seconds), like a7, series of gradual onset, rapid FH up­ k!lz broadband popping sounds which aweepJ broadband, most ener­ • increase in repetition rate, kH:1 gy below 500 H~1 heard bo~h 1111111111111111111) ending in FH upsweep grunt I J in series and singularly gradual, low amplltud~ onset, often followed by sertes of upsweep grunts variable duration {1.0-J.O • disc rete, ai111ple structure, 3 seconds), pulsed w th some 4 ?!:, rich har~nics, FH dovnsveep' kllz tonal qualttyJ broadband, kH%:1 most energy below 1000 Hz; 10 noisy1 •elephant screamNr ~ always heard singularly rich harmontcsl gradual onsetJ all energy above 500 H~1 heard 1 singularly or series of 2-3 units

loud blow1 discrete, gradual onset, wheeze sound associated 5 with exhalation at the sur!ace1 0 1 .o 2.0 surface co~ponent has character­ istics similar to those depicted seconds here (Watkins, 1977)J also termed •trumpetingN (Thompson et, al., 1977)

d 1.0 2.0 seconds Ilo!2!!.!! C. Greater than 1.0 seconds duration'. •• La•• than 0, 5 seconds duration, ! SOME CHARACTERISTICS ! EXAMPLE SOH£ CHARACTERISTICS variable duration (0,5-'i.O multiple unit, each unit is seconds)a highly pulsive, discrete1 overall sound i11 11 J vvvvvvvv variable duration (0,5-2,0 strident, noisyJ co~plex kH:z kHz structural FH and amplitude 14 1 seconds)a high frequency, modulationJ gradual onset1 rapid down-up FH, •chirp• alvaya heard singularly o.s 1.0 sec, like• sound1 all energy between 3000-6000 Hz

variable duration (1,0-5,0 'I discrete, short, simple struc­ seconds) low frequency •tog• ture• rapid upsweep FH, vide horn• sounda uniform frequency, IS kH:l side band1 alvays singular! rich harmonics, gradual onset1 ~ all energy above 1000 H:e amplitude modulation) most ener­ o!s t!o sec, gy below 1500 HzJ always heard singularly d\screte, always in series or J ~,., 16 3 units1 little or no FHa kHz ~~~ ~st energy between 500- variable duration ( 1.0- 5.0 ,, 3000 Hz 3 seconds) 1 broadband I exz:eeding- 1 l j . , ly noisi, stridentr amplitude --· J ITtT.-.~=-~~~. e .. w·y *.... modulat on a •l.lreathy• quality, kl!:t: ·~~f!!t,i'i;f{· 111ay be associated with under­ discrete, si~ple structure, .-...:._"'· .7- , •• :J::,;e,~ .... ~~~~ vater air ·release I BOme de- gradual ons~t, rapid upsweep l scending !requency qualitYI 17 FHa heard in series or singu­ t______all energy above 1000 Hz larly! ·most 'energy belov 1000 Hz

variable duration (0,5-2,0 J seconds), multiple unit, high­ IB kHz ly complex structure, rapid up and down FH1 rich harmonics, 0 1.0 2.0 I seconds number or units is variable! '------all energy between 500-4000 ll:e

discrete, complex structurea J upsweep FH1 ·~heeze-like• 19 ldfz sounda rich harmonicsagenerally low amplitude. always heard I singularly

0 1.0 2.0 seconds 44 .

• S9i11 iJWM"'jt!\I:!TICS :Uacrn•, rldnq ~•n ~••c•ndl.nr; f"''l fl~h 1\IMOCUUt:ll J"'ldi~UI 1!~1 llan

~iK'r"ltlo fl•llll lti'"ACtUrll l.lVIfl , c:ontdn11 1 "niu 'lhli:h Htlr.'lltl ... rhlnq r~. !oUt..-.q, ddr.r;t rh:" h1r:~~~ntcu 010at -•r~~r aoo .... " lOCO HI .. ... :

.UiaGl• ~>!Utl.on \lol•'),,Q I.C• ond1l1 ca•pl•,. tc:uer;ur•, .,ltlllll ...... -'-" u.nh.11 -..ltll'l• !rl't'JIC'ICT an,ttor ;-n•r•llf lu;h tr~•nc:-r, .,..rr .. ..:1 vld• dd• aondu Ill •n•r1r lt•t_..n 1000-11,000 1'11 0 =:uw:~.-nul aco..-. ---- .woo Ha ••• '·' '·' -·

'IU'hale d>U"It!On \Z,Q-0"5,3 I.CCindl)l :::::: ll"'Pl• lt;-.U:f:l!'U ~1nh f"''l aU .. •n•r;r 11:>1:1v• ~coo a:, vuia .. t• ..,u­ -- ~...._., ¥id• lld• ti&MI

t:Uo llnflgdf '";tnJ!St/'"l"l'l§lpn, 11&1 or o:ovhadonl o::an It• •nt•utv• or flllin~l nouy, bra1d. aand1 o~u;» 11n1•t• 'l'flr't hi;h 1511lit'-'d•r ;•n•t11:,7 " !'!•ud in Qrlu vltn !\!vntr vuullt• npn1 ~111n ~au 45.

'ABLE 2. Humpback whale groups monitored for sound.

Percent # of groups Observation Phonation ;roup groups producing time rate ;tructure monitored sounds (min.) (#/whale/hr)

Jone adult 34 0% 180.0 o.o :+c 7 0% 78.0 0.0 :+C+E 14 7.1% 171.0

~wo adults 49 2.0% 372 0 0

!+ adults 49 87.8% 4462.0 43.12"_55.52 :active group) 46.

rable 3. Group phonation rate versus activity rate. Rates •ere not significantly different (ANOVA: F=.994; error 1f=208; o.50

LOW MID HIGH

K+SD 38.8+56.81 41. 8+44. 08 60.3+52.20 Range o-12 o-146.4 o-194.0 )l 149 38 19 Table 4. The change in vocalization rate as new whales enter large active groups. The rates are significantly different (Chi-square: X=125.32,df=4,p<0.001).

EEUlE AFTER

Observation Vocalization Rate Vocalization Rate Example# Period( min) Group Size (#/whalelhr) Group Size (#/whale/h r)

1 11.9 4 62.5 5 35.8* 2 19.1 5 6.3 6 143.8 3 35.4 4 75.0 7 257.8 4 70.9 5 10.6 6 72.5 5 27.0 8 8.5 9 26.1 ./>. -.J

•vocalization rate decreased after joining occurred 48.

Appendix 1. Surface active groups from which underwater recordings were made, 1980, 1981. "

Observation #of 5-min. #of Whales Calf? Date Group Time (min.) Intervals X±SD(N) (No=O; Yes=1)

2/20/80 • ? ? 2/1/81 B 67 4 3.0±0.00 0 2/3/81 c 105 11 2.7±0.50 0 2/15/81 R 52 7 3.0±0.00 0 3/5/81 N 140 12 2.5±0.50 0 3/8/81 C+A 60 6 ? 3/12/81 A 60 7 5.5±0.50 0 3/1 8/81 J 29 3 7.3±0.60 0 3/18/81 z 125 7 4.1±0.40 0 3/19/81 E 22 3 4.3±0.60 0 3/20/81 z 85 10 5.3±0.50 0 3/24/81 J 60 7 5.4±3.20 0 3/24/81 z 111 10 3.8±0.60 0 4/4/81 A+C 107 7 7.6±1.80 1 4/1 0/81 •• ?

TOTAL 17hr. 3min. 94 X 78.0min./gr. 7 .2/gr. 4.6whales SD ±36.00 ±2.90 ±1.65

*recorded by P. Tyack **recorded by J. Ford 49.

Appendix 2. Recordings from surface active groups in 1982.

# 5-min. #of Whales Calf? Date Group lime (min.) Intervals (i<±SD) (No=O; Yes=1) 2/14 B 158 9 4.7±0.40 0 2/14 c 25 2 4.0t0.00 0 2/15 c 40 5 6.0±0.00 1 2/15 J 60 11 4.5±0.50 0 2/16 I 54 6 4.0±0.00 0 2/18 A 307 17 4.5±1.00 1 2/22 E 89 4 4.5±0.60 0 2/23 A 70 4 7.0±0.00 0 3/1 D 220 20 5.0±0.00 1 3/2 B 37 2 4.6±0.60 0 3/3 A 81 5 3.6±0.50 0 3/6 A 15 2 5.0±0.00 1 3/6 c 30 4 6.0+0.00 0 3/7 c 115 9 7.7±:2.20 0 3/11 A 47 3 6.0±0.00 0 3/11 B 32 2 4.0±0.00 1 3/16 c 144 12 5.6±0.50 0 3/18 B 190 11 5.3±0.50 1/0 3/20 I 153 14 5.9±1.20 0 3/21 D 109 5 5.2±1.10 0 3/21 E 71 5 4.4±0.90 0 3/22 A 235 16 4.9±,,00 0 3/22 B 36 8 10.0±0.00 0 3/26 F 100 2 6.0±1.40 0 3/26 G 28 1 5.0±0.00 1 3/29 B 127 11 6.5±1.60 0 3/31 B 244 6 4.8±0.40 0/1 4/2 A 282 4 5.0±0.00 0 4/4 c 112 15 6.1t1.10 0

TOTAL (1982): 56hr.41 min. 220int. x 113.0 min. 7.3 int./gr. 5.7/gr. 27.6% SD ±82.85 ±5.18 ±2.99 calves

TOTALS (1980-82) 73hr.44min. 314int.

X 102 min. 7.3int./gr. 5.5/gr. 22.0% SD ±73.1 ±4.56 ± 1 .51 calves 50.

Appendix 3. Surface active groups monitored for sound · from which no sounds were heard. 1981, 1982. c =calf.

Time Under Date Group #of Whales Observation (min.)

1/24/81 c 4 4.0 2/3/81 D 3 15.0 2/19/81 H 3 6.0 3/3/81 3 4.0 3/9/81 J 4+C 5.0 3/19/81 L 4-5 4.0

TOTAL 38.0 min. 51.

Appendix 4. Cow and calf groups monitored for sound, 1981 and 1982. Sounds were heard from only two groups. C+C=cow and calf, E=escort.

Time of Date Group Composition Observation (min.)

1981 2/2 A C+C 4.0 2/2 B C+C+E 3.0 2/7 B C+C 9.0 2/12 A C+C+E 3.0 2/17 A C+C 3.0 3/1 A C+C+E 7.0 3/18 A C+C+E 17.0 3/19 D C+C+E 21.0 3/23 c C+C+E 10.0 3/23 J C+C 5.0 3/24 H* C+C+E 4.0 3/26 D C+C 15.0 4/7 F C+C+E 18.0 4/4 c· C+C+E 6.0 4/14 c C+C 16.0

1982 3/15 A C+C+E 5.0 3/16 A C+C+E 13.0 3/20 A C+C+E 5.0 3/24 B C+C => C+C+E => C+C+2E 38.0 3/26 B C+C+E 8.0 3/29 B C+C+E 6.0 4/1 A C+C+E 13.0 4/4 A C+C 26.0 4/4 B C+C+E 32.0

TOTAL 4 hrs.47 min.

•social sounds heard 52.

Appendix 5. Solitary non-singing adults monitored for sound, 1981 and 1982. No sounds were heard.

Date Group Time of Observation (min.)

1/23/81 I 5.0 1/23/81 G 4.0 1/24/81 B 6.0 1/24/81 A 18.0 2/1/81 3.0 2/1/81 c 9.0 2/3/81 A 6.0 2/4/81 c 5.0 2/4/81 D 4.0 2/7/81 c 12.0 (former singer) 2/1 0/81 4.0 2/14/81 F 6.0 2/1 6/81 D 5.0 2/18/81 H 5.0 3/2/81 B 4.0 3/3/81 7.0 3/5/81 5.0 3/6/81 3.0 3/14/81 F 4.0 3/21/81 I 4.0 3/23/81 c 5.0 3/23/81 F 3.0 3/23/81 L 3.0 3/24/81 B 5.0 3/24/81 F 3.0 3/26/81 H 3.0 3/26/81 z 6.0 4/4/81 4.0 4/7/81 2.0 4/7/81 7.0 2/22/82 C+D 5.0 (former singer) 3/17/82 B 4.0 3/20/82 E 4.0 3/26/82 c 3.0 TOTAL 3.0 hrs. o min. 53.

Appendix 6. Groups containing two adult whales which were monitored for sound, 1981 and 1982. No sounds were heard. Sta~onary= "breathholders". Moving implies that a recent joining took place. E=escort.

Time of Date Group Observation(min.) Com mens

1981 1/17 A 11.0 Moving 1/21 c 10.0 . Moving; surface displays 1/21 D 5.0 Moving 1/21 4.0 Stationary 1/22 A 14.0 Moving; surface displays 1/23 A 8.0 Stationary 1/23 D 15.0 Moving 1/23 E 6.0 Moving 1/23 F 7.0 Moving 1/23 G 4.0 Former singer joined by E 1/25 A 6.0 Moving 1/28 A 9.0 Stationary 1/29 B 10.0 Moving 1/31 A 12.0 Stationary 2/1 G 4.0 Moving 2/1 H 4.0 Stationary 2/2 D 5.0 Stationary 2/4 A 8.0 Moving 2/7 A 4.0 Moving 2/7 D 5.0 Moving 2/14 2.0 Stationary 2/14 3.0 Moving 2/14 G 8.0 Moving 2/16 B 9.0 Stationary 2/17 A 5.0 Stationary 2/17 H 3.0 Stationary 2/17 p 4.0 Moving 3/2 A 9.0 Stationary 54.

Appendix 6 continued.

Time of Date Group Observation(ll)in.) Comments

1981 3/3 c 5.0 Moving, former singer+ E 3/3 0 3.0 Moving, former singer+ E 3/5 K 8.0 Stationary 3/9 H 18.0 Stationary 3/10 B 5.0 Stationary 3/19 I 3.0 Surface displays; individuals split after surface activity 3/20 H 12.0 Stationary 3/23 G 5.0 Social sounds heard 3/26 N 4.0 Stationary 3/26 28.0 Moving 4/10 A 6.0 Moving

1982 2/15 I 4.0 Stationary 2/21 c 8.0 Moving 2/21 c 25.0 Moving 2/22 B 5.0 Stationary 2/22 B 5.0 Moving; surface displays 3/1 B+C 3.0 Moving; former singer+ E; joined then split 3/2 E 2.0 Stationary 3/16 B 4.0 Surface displays; Adult (cow?) + subadult 3/29 E 17.0 Adult (cow?) + subadult 4/4 B 5.0 Former singer+ E

TOTAL TIME: 6hrs. 12 min. 55.

Appendix 7. Cow and calf groups observed, but not monitored for sound, 1981 and 1982. C+C=cow and calf, E=Single escort.

Date Group Composition

1981 2/16 H C+C 2/16 Q C+C+E 2/21 C+C 3/3 C+C 3/6 c C+C 3/11 z C+C+E 3/13 M C+C+E 4/11 A C+C+E 4/11 c C+C+E 4/19 C+C+E

1982 2/15 E C+C+E 2/16 F C+C+E 2/16 G C+C+E 2/16 H C+C 2/20 B C+C+E 2/21 D C+C 3/17 A C+C+E 3/20 B C+C 3/24 A C+C+E 3/29 F C+C+E 3/30 A C+C+E 3/31 A C+C

TOTAL GROUPS= 23

69.6% contained escorts 56.

APPENDIX 8. Characteristics of social sound types in the acoustic repertoire of the humpback whale. Maximum and minimum energy values define the entire frequency range of the sound. Maximum concentrated energy values characterize the fundamental. Mean and standard deviation of side bands (harmonics) were computed from measurements at the beginning, middle and end of a selected side band. Segmented = the sound is composed of two or more parts possessing different structure. Y = yes. N = No. Series - Y = the sound is commonly produced in a sequence of like sounds. Series - N = the sound is most often heard singularly. MAXIMUM MEAN SOUND DURATION MAXIMUM MINIMUM CONC, SIDE BAND SEG- MENTED SERIES NUMBER {SEC. ) ENERGY{Hz) ENERGY(Hz) ENERGY( Hz) WIDTH(Hz) (X:tSD)

l .23 2712 121 520 71. 7:t6.10 y y

2 .31 2996 160 652 119 . 0:!4 7 • 00 y y

3 .32 4014 50 352 73.7+31.86 N y

4 .66 3222 50 210 48. 3;t31. 21 N N

5 .91 1855 78 1680 N N U1 --J

6 1.26 7225 853 1834 42.0+4.08 N N

7 1.35 3933. 414 2345 y N

8 2.02 2035 889 1319 N N

9 2.25 3654 50 1925 65.3+18.15 y N

10 2.89 3348 718 2332 82.7:t66.07 N N

11 5.00 2260 80 1512 33.0:t1.06 N N

12 5.00 3207 50 1226 59.3:t24.24 N N

13 5.00 3680 1475 3336 N N ~lliXIHUN NEAN CONC. SEG- SOUND DURATION NAXINUN l'!ININUN SIDE BAND HENTED NUNBER (SEC. ) ENERGY( Hz) ENERGY(Hz) ENERGY(Hz) WIDTH(Hz) SERIES (X±_SD) 14 .10 7235 1520 4250 1451.3+20.12 y N

15 .ll 5850 4000 5254 y y

16 .ll 2930 150 770 103. 7±100 .4 y y

17 .24 3160 147 1448 89.3+57.8 N y

Ul 18 .31 7630 994 1500 924.3±250.55 y y CD

19 .46 4160 695 2410 167.3+22.53 N N

20 .95 3994 221 515 266.3+163.48 N N

21 1.19 6332 695 980 806.9±107.06 y N

22 1.25 10,000 782 3662 1084.1+537.35 y N

23 1.20 10,000 2615 4488 3553.7±167.04 y N

24 5.00 4458 656 824 700.0±52.90 N N

25 .23 4500 152 2864 N y