MEMOIRS OF THE ~ QUEENSLAND MUSEUM BRISBANE VOLUME 30 1 JULY, 1991 PART 2 Preface Papers contained in this number of the Memoirs of the Queensland Museum were among those presented at the Humpback Whale Conference hosted by the Queensland Museum in September 1990. Delegates from Australia, Colombia, Japan, New Zealand, South Africa and the United States of America honoured Drs R.G. Chittleborough and W.H. Dawbin, the pioneers of Australasian Humpback Whale research during the last, and probably final, period of exploita- tion of the species in the 1950s and early 1960s. Research initiatives in the post-whaling period have reflected increasing concern not only for Humpback Whales but also for the ecosystem in which they live, It was particularly appropriate that the official ceremony to acknowledge the display of life size models of Humpback Whales over the front entrance to the Museum was held during the conference and conference delegates were in attendance (see photograph over page). With the increasing numbers of Humback Whales appearing off the Australian coast, greater understanding ,through the sorts of research reported herein, will be necessary to develop harmony between whales and humans. The growing tourist interest in Humpback Whale activities in Hervey Bay must be accommodated but the interests of the whales must also be considered especially in view of the past decimation of the species wrought by commercial whalers. The Humpback Whale Conference was most notable for the many different aspects considered by the speakers (photo-id, anatomy, strandings, songs, management, modelling, to name but a few) and this diversity is reflected in the volume that follows despite its containing only a percentage of papers delivered at the conference. The Queensland Museum is proud to have been associated with this meeting of the Humpback fraternity, both as host and as publisher of the proceedings. Peter A. Jell and Robert A. Paterson Brisbane 10 May 1991. MEMOIRS OF THE (QUEENSLAND MUSEUM BRISBANE © Queensland Museum PO Box 3300, South Brisbane 4101, Australia Phone 06 7 3840 7555 Fax 06 7 3846 1226 Email qmlib@qm.qld.gov.au Website www.qm.qld.gov.au National Library of Australia card number ISSN 0079-8835 NOTE Papers published in this volume and in all previous volumes of the Afemoirs of the Queensland Museum maybe teproduced for scientific research, individual study or other educational purposes. Properly acknowledged quotations may be made but queries regarding the republication of any papers should be addressed to the Editor in Chief. Copies of the journal can be purchased from the Queensland Museum Shop. A Guide to Authors is displayed at the Queensland Museum web site A Queensland Government Project Typeset at the Queensland Museum POTENTIAL IMPACTS OF CLIMATIC CHANGE ON THE SOUTHERN OCEAN ECOSYSTEM R.G. CHITTLEBOROUGH Chittleborough, R.G. 1991 07 Q1: Potential impacts of climatic change on the Southern ocean ecosystem, Mehoirs of the Queensland Museum 30(2):245-247. Brisbane. ISSN 0079-8835. Global climate change has the potential to disrupt the delicately poised thermal balance in surface waters of the Southern Ocean around Antarctica, threatening this highly productive ecosystem with severe and permanent collapse, Pivotal in this process is a diminution of the important (bul little studied) CO2 sink in the Southern Ocean, resulting in a series of feedback loops accelerating global warming and intensifying impacts upon the Southern Ocean ecosystem. As well as outlining the processes involved and stressing the urgent need for further research, this paper underlines our wider social responsibilities to press for fresh policies essential Lo arrest the global changes before irreversible harm is done to the Antarctic environment and ecosystems, with {heir global consequences to our life support system. R.G. Chitleborough, 24 Wau Sc, Swanbourne, Western Australia 6010; 4 January 1991. The Convention for Conservation of Antarctic Marine Living Resources, (CCAMLR), while “RECOGNISING the importance of safeguard- ing the environment and protecting the integrity af the ecosystem of the seas surrounding An- tarctica’: places its main emphasis on the im- pacts which harvesting may have, not only upon target species but also upon ecalogical relation- ships. Article UI of the Convention also commits us to the conservation principle of “prevention of changes or minimisation of the risk of changes in the marine ecosystem which are not potential- ly reversible over two or three decades...” However, even if there was no exploitation of living (or mineral) resources in the Antarctic, the environment and ecosystems there are now threatened by accelerating climatic changes being triggered globally by mankind. Not surprisingly, the main focus of attention presently being given to global climatic change is how the changes will impact on ourselves. We have begun to consider how quickly we may have to adapt in terms of water supply, agricul- ture, forestry, transport. coastal developments, ele, Far less attention is being given to potential impacts upon natural ecosystems, particularly the more remote Antarctic ecosystem. PROCESSES MAINTAINING THE SOUTHERN OCEAN ECOSYSTEM The main driving force of this system is the annual pulse of winter sea ice extending northwards from Antarctica to cover same 20 million square kilometres of the Southern Ocean, retreating each summer almost to the mainland coast. This annual pulse drives the vertical circulation of these waters (Fig. 1), cold brine released by the formation of sea ice sinking along the continental shelf, with compensatory upwelling of nutrient rich water farther offshore at the Antarctic Divergence (Sverdrup et al.,1942). Microalgae growing from the base of the sea ice, and phytoplankton blooms each spring and summer within the nutrient rich An- tarctic Surface Water, are the main basis of the high productivity of the Southern Ocean ecosys- tem (Chittleborough, 1984). Estimates of gross annual production of phytoplankton are 6.1-38 billion tonnes. This represents an annual uptake of 1.5-10 billion tonnes of CO:. Part of that COz is released again during metabolic activity of consumers, but a proportion sinks as detrital organic carbon into Antarctic Bottom Water. Some dissolved CO: from the atmosphere ts also carried down in both the Antarctic Bottom Water and the Antarctic Intermediate Water sinking at the Antarctic Covergence, While it is widely accepted that the Southern Ocean is a major sink for CO measurements of the sink are not available. Takahashi (1987) es- timated that the Southern Ocean removes 6,67 billion tonnes of COz per year from the atmos- phere or 70% of total uptake flux of all oceans. t = HC ba! YS UBZANTARCTIC a — _&- MEMOIRS OF THE QUEENSLAND MUSEUM Fig, 1, Representation of currents and waler masses of the Antarclic regions (after Sverdrup et al,, 1942). THERMAL STABILITY The advent of satellite sensing affords a prac- tical means of assessing long term variations in thermal stability of the Southern Ocean, Jacka (1983) stressed the importance of monitoring the extent of winter sea ice, as this parameter is likely to be one of the earliest indicators of any significant climatic change. As satellite images of the winter sea ice around Antarctica became available, extensive open waters within the ice (polynyas) were dis- covered. The largest of these, the “Weddell Polynya” (actually situated to the east of the Weddell Sea) measured 1 L00 x 650 km in Sep- tember 1975. The Weddell Polynya appeared in three consecutive winters, 1974-1976, then dis- appeared until 1980 (Comiso and Gordon, 1987), Farther east at c, 45°E, the Cosmonaut Polynya appeared in 1973,1975, 1979,1982 and 1986, While it is not suggested that these polynyas weré caused by the greenhouse effect, they do serve to illustrate the delicate balance in the formation and maintenance of sea ice. Rather more information is available on the extent and distribution of residual sea ice during summer, as this is the time of greatest shipping activity in the Southern Ocean. While much of this information is still to be collated, an analysis by Bentley (1984) indicates that the extent of the Antarctic sea ice in summer decreased by 2.5 million square kilometres. between 1973 and 1980, and perhaps by more since the 1930's. There is. evidence of a recent increase in air temperature over the Southern Ocean. Budd (1980) recorded an increase in mean annual air temperature at subantarctic islands of 0.4° C during 1958-1978 and by 0.6° C at stations on the edge of the Antarctic continent. At Ker- guelen Island, situated on the northern edge of Antarctic Surface Water, the mean annual air temperature has risen during 1964-1982 by 2.1° C (Jacka, Christou & Cook, 1986), the increase being more marked in summer than in winter. Similar rises in mean annual air temperature are recorded for Amsterdam Island and Marion Is- land, On glaciated subantarctic islands such as Heard Island, glacial retreat has accelerated dramatically in recent decades (Allison and Keage, 1986). Heard Island, located towards the outer margin of Antarctic Surface Water, and its CLIMATIC CHANGE IN THE SOUTHERN OCEAN ECOSYSTEM glaciers moving i ell on steep slopes (short residence time of the ice), affords sensitive in- dicators of changes in climate. While there is a paucity of hard data on variability within the Southern Ocean region, it is evident from the great changes occurring seasonally and in the longer term, that the ther- mal balance of this region ts delicately poised. POTENTIAL FOR CHANGE Global climate models used {o predict surface air temperature changes due to increasing atmos- pheric COz, generally indicate greater increases in temperature at higher latitudes, For example, Rind (1984) indicated that a doubling of atmos- pheric CO: would raise mean annual air temperatures over most of Australia by 44°C, while over the Southern Ocean around An- tarctica increases of 6-8°C could be anticipated. Sea surface temperatures in the Southern Ocean can therefore be expected to rise by a greater amount than in lower latitudes, The most immediute effect of Souther Occan surface isotherms contracting southwards would be to further restrict the distribution of cold lolerant species living within Antarctic Surface Water. For example, most of the slock of Euphausia superba is confined 10 waters less than 2°C (Marr, 1962), while E. crystallorophias is restricted to waters of even lower temperature. The shrinking range of such key food species will compress dependent consumer species into a narrower band around Antarctica. increasing competition between predators, Around some sub-Antaretic islands, vital food resources may then be beyond the foraging range of adult seals and birds during the critical period of rearing the young. Croxal et al. (1987) showed that this already occurs sporadically al South Georgia. One of the most far reaching effects of rising sca surface temperatures in the Southern Ocean would be a progressive reduction in the extent of sea ice, both in winter and summer, In an initial modelling of the potential impact upon Southern Ocean sea ice, Parkinson and Bindschadler (1984) concluded that a rise of S°C in air lemperature could result in the winter extent and volume of sea ice to be halved. As the increase in wir temperature at high latitudes is anticipated to continue to rise well beyond that level, the winter extent of sea ice could be reduced even further. Direct ecological impacts of a reduction in sea ice include loss of substrate for ice ulgae (an important component of primary production during winter and early spring), and less ice floes suilable for pupping and mating of ice seals (particularly crabeater and leopard seals) at the critical period during late October and carly November each year. One of the physical effects of a reduction in the extent of sea ice would be a loss of albede and further absorption of solar energy into the surface of the Southern Ocean, accelerating the change in the energy balance. An even more important feedback loop from a much reduced extent of winter sea ice would be a severe reduction in the pulse driving the vertical circulation of the waters of the Southern Ocean. With Jess sea ice formed, there would be less brine released to sink as Antarctic Bollom Water and hence diminution of the passage of dissolved CQz to be held in the deep ocean sink- With the weakening of the vertical circulation there would also be a decline in compensatory upwelling of nutrient rich water upon which phytoplankton and all higher consumers are to- tally dependent. A failure of phytoplankton blooms would represent a massive reduction im the fixation of COs in Antarctic Surface Water, again feeding back to accelerate global warming. A severe reduction in primary production within Antarctic Surface Water would have a disastrous impaci on the Southern Ocean ecosys- tem as a whole. including the harvested specics which CCAMLR is allempling to manage and conserve. Furthermore, a diminution of krill stocks through man-induced climate changes would severely retard (or reverse) the recovery of previously depleted populations of blue. fin and humpback whales, negating much of the hard-won ground by the lWC, Further potential for impact upon the Southern Ocean ecosystem derives from the ultraviolet wavelengths penetrating the ozonc hole now evi- dent in the stratosphere over Antarctica each spring and early summer. Increasing penctration of UY band tnto the sea surface has potential to depress photosynthesis or even to be lethal to the more sensitive species in the phytoplankton. again depressing the productivity of the ecosys~ tem as well as reducing the uptake of aimos- pheric COz Precise field measurements are lacking, but Pittock ct al. (1981) suggested that phytoplankton is surface waters would suffer Sppicctebhe mortality by a reduction of the ozone shield in the range of 16-30%, In October 1985 ozone levels over Antarctica declined by 50% (Ember ¢} a)., 1986), Concencrations of 246 MEMOIRS OF THE QUE ENSLAND MUSEUM Excessive resource consumption Warmer southern ocean albedo Greenhouse gases| accumulate in troposphere Warmer carbon troposphere Less phytoplankton production Less Faster rise in atmospheric C02 Less iil less whales Less detrital upwelling of nutrients Ozone hole in stratosphere over Antarctica Less penguins less seals Eye damage to new-born seals & birds “UV(B) penetrates surface water Less Antarctic bottom water Much reduced ocean sink of CO2 Fig. 2. Potential feedback loops within the Southern Ocean ecosystem resulting from climatic changes. phytoplankton some tens of metres below the surface may be afforded some protection from UV radiation, but turbulent wind mixing bring- ing these organisms close to the sea surface may well make them more vulnerable. Of more immediate impact could be the eye damage that the UV band might bring to new- born seals and birds. Particularly vulnerable here would be the pups of crabeater seals, born on ice floes in high latitudes late in October, close to the peak of the ozone hole. Again, direct meas- urements of sensitivity are lacking. There are many potential feedback loops im- pacting on the Southern Ocean ecosystem from global warming and weakening of the ozone shield (Fig.2). Even if subsequent research finds that one or two of these processes are of relative- ly minor extent, the overall prognosis for the Southern Ocean ecosystem is extremely poor. Furthermore, the collapse of the important Southern Ocean COz sink has global implica- tions, greatly accelerating the rates of climate change that each of us will have to face. It should be noted that other feedback systems in the Antarctic, not directly relating to the Southern Ocean ecosystem, but having potential to affect global climate change, are not discussed here. These include acceleration of break-up from the fringes of the Antarctic glacial ice sheet, and a weakening circulation of the Southern Ocean triggering a permanent ENSO phen- omenon. As stressed by Thomas (1984) “we cannot rule out the possibility that a climate change of magnitude predicted for CO2 doubling could radically alter ocean circulation”. In par- ticular, he points out that if the relatively warmer Circumpolar Deep Water is able to reach the major ice shelves without considerable cooling, ice shelves could thin enough for massively en- hanced calving. “Clearly”, he concluded, “we need to learn more about ocean behaviour neat Antarctica.” CONCLUSION There can be little doubt that the Southern Ocean ecosystem is threatened with severe and permanent collapse as a result of impending climatic changes. Such a collapse would halt the present recovery of humpback whale popula- tions. As scientists, we can design fascinating CLIMATIC CHANGE IN THE SOUTHERN OCEAN ECOSYSTEM studies of the physical and biological impacts upon the Antarctic environment as global climatic changes progress. But do we not also have a wider responsibility to press for policy changes aimed at arresting the global processes before irreversible harm is done to the Antarctic environment and ecosystems? Alihough it is quite evident that we need far more research in this area, we can hardly afford to regard the Southern Ocean as a giant experimental unit if we are likely to lose control of the experiment. Our role should be far more than the gathering of information. We would be failing in our social responsibilities if we do not make a clear state- ment on the urgent need for action to circumvent the setting up in the Southern Ocean of irre- versible processes having high potential to cause massive environmental and ecological changes around Antarctica, as well as greatly accelerat- ing changes in climate (and sea level) throughout the world, Unless we act quickly and decisively, the conservation strategies presently being pursued by Australia within the WC, CCAMLR and the Antarctic Treaty itself, become meaningless ges- (ures. As stated recently by Dr Noel Brown of UNEP, the next decade is our last window of opportunity to make effective changes. Let’s use that time to the full. LITERATURE CITED ALLISON, I.F. AND KEAGE, P.L. 1986. Recent changes in the glaciers of Heard Island. Polar Record 23 (144); 255-271. BENTLEY, C.R. 1984. Some aspects of the cryo- sphere and its role in climatic change. Geophysi- cal Monographs 29: 207-220. BUDD, W.-F. 1980. The importance of the polar regions for the atmospheric carbon dioxide con- centrations, 115-128. In G.l. Pearman (ed.), ‘Carbon Dioxide and climate,’ (Australian Academy of Science; Canberra), CHITTLEBOROUGH, R.G. 1984, Nature, extent and management of Antaractic living resources, 135-161. In §. Harris, ed.. ‘Australia’s An- tarctic Policy Options’, Centre for Resource and Environmental Studies, ANU Mon. 11; 135- 161. COMISQ, I.C. AND GORDON, A.L. 1987. Recur- ting polynyas over the Cosmonaut Sea and Maud Rise. J. Geophys. Res, 92; 2819-2833, CROXALL, J.P. et al. 1987. Reproductive perfor- mance of seabirds and seals at South Georgia and Signy Island, South Orkney Islands, 1976-1987: implications for Southem Qcean monitoring studies. SC-CAMLR. Selected Scientific Papers 1987: 445-447, EMBER, L.R. et al. 1986, Tending global commons. Chemical & Engineering News 64 (47): 14-64. JACKA, T.H, 1983, A computer data base for An- larclic sea ice extent, ANARE Research Notes 13: 1-54. JACKA, T.H., CHRISTOQU, L. AND COOK, B.F. 1984. A data bank of mean monthly and annual surface temperatures for Antaretica, the Southern Ocean and South Pacific Ocean, ANARE Research Notes 22: 1-97. MARR, J.W,S. 1962. The natural history and geog- raphy of the Antarctic krill (Euphausia superba Dana). Discovery Reports 32; 37-463. PARKINSON, C.L. AND BINDSCHADLER, R.A. 1984, Response of Antarctic sea ice to uniform atmospheric increases, Geophysical Mono- graphs 29; 254-264. PITTOCK, A.B. et al. 1981. Human impact on the global atmosphere: impacts for Australia, Search 12; 260-272, RIND, D_ 1984. Global climate in the 21st. Century. Ambio 13: 148-151. SVERDRUP, H-V_etal. 1942. ‘The oceans’ .(Prentice hall: N.Y). TAKAHASHI, T, 1987, Assessment of seasonal and geographic variability in CO2 sinks and sources in the ocean, In Reichle, D.E. et al., ‘Environ- mental Sciences Div. Ann. Progr, Rep. for period ending Sept, 30 1986", (Oak Ridge Na- lional Lab.: Tennessee). THOMAS, R.H. 1984. Responses of the polar ice sheets to climatic warming. 301-316. In ‘Glaciers, ice sheets and sea level: effects of a C0o-induced climatic change. Report to U.S. Dept. of Energy. DoB/ER/60235-1. MEMOIRS OF THE (QUEENSLAND MUSEUM BRISBANE © Queensland Museum PO Box 3300, South Brisbane 4101, Australia Phone 06 7 3840 7555 Fax 06 7 3846 1226 Email qmlib@qm.qld.gov.au Website www.qm.qld.gov.au National Library of Australia card number ISSN 0079-8835 NOTE Papers published in this volume and in all previous volumes of the Afemoirs of the Queensland Museum maybe teproduced for scientific research, individual study or other educational purposes. Properly acknowledged quotations may be made but queries regarding the republication of any papers should be addressed to the Editor in Chief. Copies of the journal can be purchased from the Queensland Museum Shop. A Guide to Authors is displayed at the Queensland Museum web site A Queensland Government Project Typeset at the Queensland Museum HUMPBACK WHALE SONGS ALONG THE COAST OF WESTERN AUSTRALIA AND SOME COMPARISON WITH EAST COAST SONGS WILLIAM H. DAWBIN AND ELIZABETH J_EYRE Dawbin, W.H. and Eyre, E.J, 1991 07 01; Humpback Whale songs along the coasi of Western Australia and some comparison wilh east coast songs, Memoirs of the Queenslana Museum 30(2): 249-254, Brisbane. ISSN 0079-8835. Humpback Whale songs have been recorded since 1986 off Western Australia. These songs share nu themes with those recorded off eastern Australia, supporting other evidence that the two breeding stocks are separate despite Some overlap in Antarctic feeding areas. William H, Dawbin, Australian Museum, 6-8 College Street, Sydney, New South Wales, 2000, Elizabeth]. Eyre, 16/83-85 Alfred Street, Ramsgate Beach, New South Wales 2217; 27 March 1997, Humpback whales regularly migrate along continental coastlines in the Southern Hemi- sphere (Dawbin, 1966), and produce complex songs (Payne and McVay, 1971) which change over time (Payne et al,, 1983). To date most detailed studies refer to Northern Hemisphere stocks. This report describes the results of studies from the west coast of Australia which began in 1986, together with comparison with data collected from the east coast in 1989 Studies of populations of humpbacks in the Northern Hemisphere have shawn that songs are similar within oceans, but differ significantly where oceans are isolated by a land mass (Payne and Guinee, 1983 ; Winn et al., 1981). The Australian populations of Areas TV and V also exhibit major differences from each other in song length and content. Evolution of songs can he a gradual or rapid process, with songs changing not only from year to year, but also within the year, as the whales migrate to and from the breeding grounds, This paper will examine some of the song changes that have occurred amongst the Area IV Feet during 1986-1989, with recent data rom 1990 included. Recordings from the cast coast will be examined briefly for comparison between populations. METHODS Equipment used was a Clevite Ordnance Oyster hydrophone (CH-15) with 30 m of cable, a preamplifier and a Sony WMD6 cassette re- corder. Recordings used for analysis were made during the migration period from Dampier (20° 39°S, 116° 45°E) to Rottnest Island (32° 00°S, 115° 30°B), Other recordings have been made along the west coast from its northern most to southern most extremities (Dawbin and Gill, this memoir). Spectrograms of the songs and shart hand use of descriptive words for each sound were incorporated for song analysis. The spectrograms were traced for the purposes of this report to eliminate background noise and enable clearer definition of song units, Themes. were numbered by allocating the last theme as that which included sounds which were nearest in character to the “surface ratchet” described by Winn et al. (1971). As songs may differ between singers, comparisons here are made on what are regarded as representative song samples. Table 1, demonstrating song change, uses methods as described by Helweg et al. (1990). The Western Australian 1956 song is used as a reference, with each theme: occurring after the surface ratchet labelled successively from A to E. Themes from following years which cor- respond to any of those in 1986 fall into A to E. but new themes are given subsequent new letters. RESULTS The song changed in the first theme from 1986 to 1988 and 1990 (Fig. 1), Between the 1986 and 1987 songs there was substantial change, with little similarity between themes. In 1988 there was substantial change from the 1987 song rep- ertoire. From 1988 onwards, however, the arran- gements of sounds within themes became more similar, with only slight progressive change be- tween units of sound from 1988 to 1990, The change in song from 1986 to 1987 was not as great as it Was from 1987 (o 1988 (Table 1). The number of themes remained the same over MEMOIRS OF THE QUEENSLAND MUSEUM 250 “po1ind90 aARy yey) Sodueyo ay} oJRASN]]I O} swuPIsONIAdS Jo SSulors] BuIsn ‘YG6] PUP SQ6T O1 OQHT WO eI[eIISNY JO ISROD JSOM dy} BuopR OW} ISI OY) JO uOSRdWIOD "| “O]4 spuosas 9T 8 0 | 0661 ‘T asnsny qutod eqqong 8soL *gT TaqweaoNn jyaoy uea7Q L861 ‘ET tequeadas ilatdueg gget ‘Atnr z pueTs—T 3sauq joy 2uy 251 HUMPBACK WHALE SONGS, W.A. ‘aypunxosdde si ayeos awit) ay,f, “ISEOD YRA UdaMIJaq AWAY) JSP] 94) JO JUOJUOS ay) Ul aouaIAJJIP B SayesISUOWAP PUP ISBOD JSP AY) WO) S} Sure) psy) ayy, ‘aseayd so8uo] & ul SuNjnses Yssy ay) ULL) SUN punos Jo suoadas BOW SMOYs d[dwIRs pUOdIS DY “PILRAISMY WIAISOM Suoye ‘Ajaanaadsas “Suos uonessiw wisyynos pur usayou ayessnyy! sojdwes Om) ySJJ OU] “6861 SulNp SAulpsooal JUdIIyJIP WOIY UDY) ISB] OY) JO aspayd aanejuasaidos & Suedwoo ssuroes wesdonoads "7 ‘O14 spuovas OT 8 0 Ks ding ‘ 3¢ et + , PORE TPLEUT ES yt wider Hiatt ' 4 Wana y| saa ‘emi QZ 3snsny keg Aasaray Roe apes teh OZ 14284909790 PUBTSI 4sauq joy Ww na sa a ‘pat ZI asngsny FA UOAIPUIP) 2 yy MEMOIRS OF THE QUEENSLAND MUSEUM fer ‘suoHeinp oseiyd jusJajJIP SNYy) puke ‘s}1uN puNos Jo suOI Nadal Jo siaquinu juasa}JIp MOYs Sa]dures jseOD ISOM OY) ‘UIPdYy °Z “BLY SB RIEP OWS OY) JO} HRT SULINP SBuIpsodai JUsIIJJIP Woy BWIY) IS11J 9) JO oSeIYd dANeyUasoIdas B SuLeduiod sBuldey wWeIdoNads “¢ “OL spuovas gf 8 0 — ES Se 9% Isnsny feq Avaray 0Z 13q0399 pueTST 4sauqqoy * te ' = — a POR \ aed i } ; * Why) a 4 vet | ‘es 2 Le pole aan ; I \ u Ps | \ s4 } cs T ‘ \ : ale ' es My ae ae a bia rg tee Hee re YO Zp TeneRy ! : | { Zz uoareuie) zuy HUMPBACK WHALE SONGS, WA, 1986 and 1987 and then decreased from 5 to 4 in 1988 (Table 1). The song remained stable during 1988 and 1989, and then in 1990 two themes were incorporated into one, dropping the theme number from 4 to 3. For further comparison, Fig. 2 shows the last theme from the year 1989, and Fig. 3 the first theme from the same year. Both northern and southern migration songs are represented from Western Australia and these resemble each other, These first and last themes of 1989 resemble the first and last themes of 1988 and 1990 (Fee 1). TABLE |, Repeated and new themes over five years on the west coast and wcomparison with one year from the east coast of Australia. The 1990 song consists of three themes, but the first theme is actually an incor- poration of the first and second theme from 1958 and 1989. ‘The east and west coast show no similarity in song content. The 1989 Hervey Bay sample has no shared themes with the west from any years {Table 1). There is no similarity between cousts in song elements in the first and last themes during 1989 (Figs 2,3). This contrasts with the sharing of at leastsome themes between Hawaii, Mexico and Bonin Islands (Helweg etal., 1990), Evidence from “Discovery” marks indicates that two animals marked off eastern Australia were killed off Western Australia (Chitleborough, 1965) but acoustic data from the present study to wa ye} suggest a high degree of independence between east and west coast breeding stocks, DISCUSSION Song change can occur on many levels. Recordings from both coasts of Australia have shown that the content and structure of a song may change between years or within years during the northern and southern migrations. There can also be differences between individual singers and each song rendition of an individual. Change can occur either suddenly or progres- sively, and even within the songs themselves there can be different rates of evolution. Recordings from east and west show that little or progressive change tends to occur in the last and first theme, and that the majority of change occurs in the ‘body’ of the song. Therefore gradual change is more likely to happen at the end or beginning, with more rapid change taking place in the middle. This rapid change was seen in the middle themes from 1986 and 1987, with the first and last themes evolving at a slower rate. In contrast, a slower change from 1988 to 1990) occurred during the whole song, with some themes not changing at all. Recordings were made during migration as opposed to on the breeding grounds. This is a difficult environment to record in, as the whales are in transit on active migration, and weather conditions are more frequently unfavourable. This enabled only a small sample of song to be collected each year, and the identification and subsequent resighting of individual singers was virtually impossible. Our studies have shown, however, that Australian Humpback Whale song differs be- tween the west and east coast and that these songs change over time. The variability in song and the different rates it can occur at have been presented here, but what determines the rate of change remains unknown. ACKNOWLEDGEMENTS Thanks go to Chris Burton, Robert Mannell, Hec Goodall and the siaff at The Pet Porpoisc Pool, Ross Isaacs and Peter Gill. This project has received financial support from a Marine Sciences and Technology Research Grant to Dawbin and Cato. Discussions with Dr. Doug Cato have been helpful and much appreciated, The N.S.W, National Parks and Wildlife Service 254 have provided valuable vessel support for sound recording. LITERATURE CITED CHITTLEBOROUGH, R.G. 1965. Dynamics of two populations of the Humpback Whale Megaptera novaeangliae (Borowski). Aust. J. Mar. Freshw. Res. 16: 33-128. DAWBIN, W.H. 1966. The seasonal migratory cycle of Humpback Whales. 145-170. In K.S. Norris (ed.), ‘Whales, dolphins and porpoises’, (Univ. Calif. Press: Berkeley). DAWBIN, W.H. AND GILL, P.C. this memoir. Humpback Whale survey along the west coast of Australia: a comparison of visual and acoustic observations. Mem. Qd Mus. 30(2): 255-257. HELWEG, D.A., HERMAN, L.M., YAMAMOTO, S. AND FORRESTALL, P.H. 1990, Com- parison of songs of Humpback Whales (Megap- tera novaeangliae) recorded in Japan, Hawaii and Mexico during the winter of 1989. Sci. ‘Repts. Cetacean Res. 1: 1-20. PAYNE, K., TYACK, P. AND PAYNE, R. 1983. Progressive changes in the songs of the MEMOIRS OF THE QUEENSLAND MUSEUM Humpback Whales (Megaptera novaeangliae): A detailed analysis of two seasons in Hawaii, 9-57. In R. Payne (ed.), ‘Communciation and behaviour of whales’, (Westview Press: Boulder), F PAYNE, R.S. AND McVAY, S. 1971. Songs of Humpback Whales. Science 173: 585-597. PAYNE, R. AND GUINEE, L. 1983, Humpback Whale (Megaptera novaeangliae) songs as an indicator of “stocks”, 333-359. In R. Payne (ed.), ‘Communication and behaviour of whales’. (Westview Press: Boulder). THOMPSON, T.J., WINN, H.E. AND PERKINS, P.J. 1979. Mysticete Sounds, 403-431. In H.E. Winn and B. Olla (eds), ‘Behaviour of marine mam- mals, vol. 3: Cetaceans. (Plenum Press: New York). WINN, H.E., THOMPSON, T.J., CUMMINGS, W.C., HAIN, J.. HUDNALL, J.. HAYS, H. AND STEINER, W.W. 1981. Song of the Humpback Whale - population comparisons. Behavioural Ecology and Sociobiology 8: 41- 46, MEMOIRS OF THE (QUEENSLAND MUSEUM BRISBANE © Queensland Museum PO Box 3300, South Brisbane 4101, Australia Phone 06 7 3840 7555 Fax 06 7 3846 1226 Email qmlib@qm.qld.gov.au Website www.qm.qld.gov.au National Library of Australia card number ISSN 0079-8835 NOTE Papers published in this volume and in all previous volumes of the Afemoirs of the Queensland Museum maybe teproduced for scientific research, individual study or other educational purposes. Properly acknowledged quotations may be made but queries regarding the republication of any papers should be addressed to the Editor in Chief. Copies of the journal can be purchased from the Queensland Museum Shop. A Guide to Authors is displayed at the Queensland Museum web site A Queensland Government Project Typeset at the Queensland Museum HUMPBACK WHALE SURVEY ALONG THE WEST COAST OF AUSTRALIA: A COMPARISON OF VISUAL AND ACOUSTIC OBSERVATIONS WILLIAM H. DAWBIN AND PETER C GILL Dawbin, W.H. and Gill, PC. 1991:07:01; Humpback whale survey along the wes! coast of Australia: a comparison of visual and acoustic observations. Memoirs of the Queensland Museum 30(2): 255-257. Brisbane. ISSN 0079 8835. The results of asurvey of humpback whales, using visual and acoustic delection techniques, along the Western Australian coast during the winter and spring of 1989 are reparted- William H. Dawbin, Australian Museum, 6-8 College Sireet, Sydney, N.S.W, 2000; Peter C. Gill, Oceanic Research Foundation, PO Box 247, Windsor, New South Wales 2756; 10) January, 199), The migratory cycle of humpback whales (Dawbin,1966), and the population charac- teristics of Area [TV humpbacks (Chittle- borough, 1965) and their increase in abundance in Area IV since exploitation ceased in 1962 (Bannister,1990) are well known. Less well studied is distribution of Area IV animals off Western Australia, when they congregate in Warm inshore waters to breed. This study was conducted from a 20m schooner, the R/V ~Thistlethwayte”, while on a circumnavigation of Australia during the austral winter and spring of 1989, The voyage allowed a survey of inshore waters along the Western Australian coast during the humpback breeding season, Acoustic and visual detection of whales has been used to determine migration routes, Folkow. and Blix (1990) detected several species during a sound survey of the mid-Atlantic, while Clapham and Mattila (1990) detected humpback song at sca in the western Atlantic, distant {rom land, Studies of humpback song on breeding grounds elsewhere are well known and too numerous to list here, but humpback sounds are known to be indicators of stock identity (Payne and Guince, 1983), and songs from Western Australia have been shown to differ from those on the east coast (Dawbin and Eyre, this mem- oir), WHD and Chris Burton have recorded humpback sounds in various latitudes off the Western Australian coast, as far south as Cape Leeuwin (34° 20'S). It was therefore decided to use acoustic and visual observations to monitor humpbacks during the southward passage. The vessel left Darwin in mid July 1989, and sound samples were taken at intervals of no more \han four hours, navigation and weather permit- ting, while at sea. The vessel stayed as clase ta {he coast as was navigationally prudent. On July 24, humpbacks were first encountered visually and acoustically at 15° 17'S, 123° 52'E. This was considerably further north than the generally accepted limits of Area TV breeding grounds (Harrison and Bryden, 1988), though fisherfolk familiar with these walers laler reported thal humpbacks are commonly seen in the area (P.Canney, pers.comm.), We regarded our timing as favourable, because the majorily of whales are still moving north al this time. Sound was monitored over 33 days al more than 100 stations, with 92 stations between the first and last recordings; stations spanned 15° of latitude and 1600 nautical miles. Generally the weather was calmer north of Broome, with fre- quent short spells of bad weather to the south. Sightings and sound recordings were made as faras 30° 24’S, with animals being detected three limes as often by sound as by sight, A total of 40 humpbacks were sighted on 21 occasions, and a conservalive total af 127 heard on 64 occasions. Conservative, because when more than one animal was singing faintly, it was often difficult to distinguish how many were singing. In these cases, the minimum definite number was recorded, Animals were heard singing af all times of the day and night. Humpback sounds typically consisted of a “chorus” of several animals singing the same song, apparently inde- pendently of cach other. Occasionally “social” (non-song) sounds were heard, often during ab- served vigorous displays of social behaviour, as described by Silber re). Only at 23 (25%) slalions were no Whales detected at all. In 13 of these, detection may have been hampered by surface waves (sound), or by darkness or whitecaps (sight). Dawbin (1956) discussed factors affecting the coastal migration routes of humpbacks. To these 256 MEMOIRS OF THE QUEENSLAND MUSEUM 14S Os 500 Km a 200 miles 314 PERTH 114E ALBANY FIG. 1. Map of Western Australia, showing numbers and locations of humpback whales located during the survey. Animals detected by sound alone are denoted by a plain number; those detected visually are denoted by acircled number. This does not mean that the sightings and soundings are mutually exclusive; sound was often detected during sightings. Not all monitoring stations are shown. we must add consideration of the possible effects | and which may even have resulted in a localised of large-scale offshore oil and gas development. modification of the humpback migration path. Gas rigs in the vicinity of Barrow Island created Bowhead whales in the Beaufort Sea have shown mechanical underwater noise levels which avoidance behaviours in reponse to oil explora- would certainly hinder cetaceancommunication, _ tion and drilling (Richardson et al.,1985,1986). ACOUSTIC AND VISUAL HUMPBACK SURVEY, W.A. The extent of Area IV humpback breeding grounds is still undetermined. Townsend (1935) showed that humpbacks were taken off the northwest coast, west of Broome. In this study, almost all of the animals observed south of Broome were travelling in a northward direction, while animals to the north of Broome were generally static. Two cows with young calves were seen north of Broome, a town previously tegarded as the extreme northern limit of the breeding area: these were the only calves seen during the survey. This indicates a wide latitudinal range in which humpbacks may calve: parturition has been reported as far south as Albany, at 35°S (Chittleborough, 1965). Water temperatures and depth profiles north of Broome are comparable to those of humpback breeding areas in the West Indies (Whitehead and Moore,1982). Animals apparently engaged in courtship were also observed north of Broome. Visual monitoring would have indicated only 1/3 the whales detected by hydrophone. This is despite the fact that Humpback songs are re- garded as being almost exclusively produced by males. Past catch records indicate that females occur in almost equal abundance off W.A. Acoustic monitoring has the advantage of being able to operate in darkness or reduced visibility; even in reasonable conditions whales may evade visual detection where they are not expected. In 1986 WHD recorded humpback sounds off Rottnest Island near Perth, when there had been no visual reports for many years. The combined results of this survey, and earlier sur- veys by WHD and Chris Burton show that humpbacks vocalise, and can therefore be monitored acoustically, along the Western Australian coast between 15-34°S. ACKNOWLEDGEMENTS Special thanks to the crew of the “Thistleth- wayte’ for their enthusiastic help during the sur- vey. Thanks also to Pam Canney and Ian Lew of F/V “Rachel” in Broome, for their hospitality and valuable information about humpback sight- ings. Financial assistance was providedby a Marine Sciences and Technology Research Grant, and by a grant from the Australian Whale Conservation Society LITERATURE CITED BANNISTER,J.L., KIRKWOOD,G.P. AND WAYTE,S.E. 1990. Population increase in 257 “Group IV” humpback whales, Western Australia. SC-[WC/42/PSI, CHITTLEBOROUGH,R.G. 1965. Dynamics of two populations of the humpback whale, Megaptera novaeangliae (Borowski). Aust. J. Mar. Freshw. Res.16: 33-128. CLAPHAM,P.J. AND MATTILA,D.K. 1990. Humpback whale songs as indicators of migra- tion routes. Marine Mammal Science 6(2): 155— 160. DAWBIN, W.H. 1956. The migrations of humpback whales which pass the New Zealand coast. Trans. R. Soc. N.Z. 84:147-196. DAWBIN, W.H. 1966. The seasonal migratory cycle of Humpback Whales. 145-170. In K.S. Norris, (ed.), ‘Whales, dolphins and porpoises’, (Univ. of California Press: Berkeley). DAWBIN, W.H. AND EYRE, E.J. this memoir. Humpback Whale songs along the coast of Western Australia and some comparison with east coast songs.Mem, Qd Mus.30(2):249-254, FOLKOW,L.P. AND BLIX,A.S. 1990. Norwegian whale sighting and acoustic surveys in the At- lantic Ocean during the winter of 1989-90. IWCSC/42/06. HARRISON,R. AND BRYDEN,M.M.(eds) 1988. ‘Whales, dolphins and porpoises’. (Golden Press: Silverwater). PAYNE,R. AND GUINEE,L.N. 1983. Humpback Whale (Megaptera novaeangliae) songs as an indicator of “stocks”. 333-359. In R.Payne, ed., ‘Communication and Behaviour of Whales’. (Westview Press: Boulder). RICHARDSON,W.J., FRAKER,M.A., WURSIG,B., AND WELLS,R.S. 1985. Behaviour of bowhead whales, Balaena mysticetus, summer- ing in the Beaufort Sea: reactions to industrial activities. Biol.Conserv.32, 195-230. RICHARDSON,W.J., WURSIG,B. AND GREENE, C.R. 1986. Reactions of bowhead whales, Balaena mysticetus, to seismic exploration in the Beaufort Sea. J. Acoust. Soc. Am. 79(4), 1117-1128. SILBER, G.K. 1986. The relationship of social vocalisations to surface behaviour and aggres- sion in the Hawaiian humpback whale (Megap- tera novaeangliae). Can. J. Zool. 64, 2075-2080. TOWNSEND, C.H. 1935. The distribution of certain whales as shown by log book records of American whaleships. Zoologica 19: 1-50. WHITEHEAD, H. AND MOORE, M.J. 1982. Dis- tribution and movements of West Indian humpback whales in winter. Can. J. Zool. 60, 2203-2211. MEMOIRS OF THE (QUEENSLAND MUSEUM BRISBANE © Queensland Museum PO Box 3300, South Brisbane 4101, Australia Phone 06 7 3840 7555 Fax 06 7 3846 1226 Email qmlib@qm.qld.gov.au Website www.qm.qld.gov.au National Library of Australia card number ISSN 0079-8835 NOTE Papers published in this volume and in all previous volumes of the Afemoirs of the Queensland Museum maybe teproduced for scientific research, individual study or other educational purposes. Properly acknowledged quotations may be made but queries regarding the republication of any papers should be addressed to the Editor in Chief. Copies of the journal can be purchased from the Queensland Museum Shop. A Guide to Authors is displayed at the Queensland Museum web site A Queensland Government Project Typeset at the Queensland Museum WESTERN AUSTRALIAN HUMPBACKS SINCE 1963 J.L.BANNISTER Bannister, J.L. 1991 07 01: Western Australian humpbacks since 1963. Memoirs of the Queensland Museum 30(2): 258. Brisbane. ISSN 0079-8835. Aerial surveys off Shark Bay, Western Australia, since 1976 have demonstrated a sig- nificant increase in Southern Hemisphere “Group IV” Humpback Whales since whaling ceased off Western Australia in 1963. “Group IV” animals are humpbacks which summer in Antarctic Area IV (70°E —-130°E) and winter north of that area, off Western Australia. Flights have been timed for comparison with data available from whaling company spotter aircraft operations in 1963. Since 1977 they have been undertaken over a ten day period in mid-July, when most sighted whales are still travelling northwards. Results demonstrate that the number had at least doubled between 1963 and 1982. The best estimate of the increase rate is 8.8% per annum. It is possible that recovery may have been somewhat delayed. To detect any further increase, it is planned to repeat the surveys, using the standardised techniques already adopted, for the same period and at the same time of the year, in 1991 and 1994. Given an estimate of some 500 animals remaining at the end of 1963, of which at least half would have been immature, and allowing for a possible delay in recovery, the present number off Western Australia in winter/spring seems to be around 2,000-3,000. That result is broadly consistent with the sighting rate of 17 per day, mostly moving north, in the Shark Bay aerial survey in 1988 and an encounter rate of 1.5 per hour, mostly moving south, in field operations for photographic identification off the Dampier Archipelago in 1990. John L. Bannister, Western Australian Museum, Francis Street, Perth, Western Australia 6000; 8 February, 1991. MEMOIRS OF THE (QUEENSLAND MUSEUM BRISBANE © Queensland Museum PO Box 3300, South Brisbane 4101, Australia Phone 06 7 3840 7555 Fax 06 7 3846 1226 Email qmlib@qm.qld.gov.au Website www.qm.qld.gov.au National Library of Australia card number ISSN 0079-8835 NOTE Papers published in this volume and in all previous volumes of the Afemoirs of the Queensland Museum maybe teproduced for scientific research, individual study or other educational purposes. Properly acknowledged quotations may be made but queries regarding the republication of any papers should be addressed to the Editor in Chief. Copies of the journal can be purchased from the Queensland Museum Shop. A Guide to Authors is displayed at the Queensland Museum web site A Queensland Government Project Typeset at the Queensland Museum SIGHTING ANALYSIS AND PHOTO-IDENTIFICATION OF HUMPBACK WHALES. OFF WESTERN AUSTRALIA, 1989 CHRISTOPHER L.K. BURTON Burton, C.L.K. 1991 07 O01; Sighting analysis and photo-identification of humpback whales, off Western Australia, 1989, Memoirs of the Queensland Museum 30(2): 259-270. Brisbane, ISSN 0079-8835. During 1989, 492 humpback whales Megaptera novaeangliae af the Southern Hemisphere Group IV stock were observed off the Western Australian coast, and 118 of these were photographically identified using tail fluke and lateral body pigmentation patterns. Most observations were made duting whale-watching cruises conducted off Perth, with the highest sighting rates (whales/sighting hour) being recorded in September and October. Sightings of mother-calf pairs increased in November. To improve efficiency of identifica- tion, animals were classified into 4 Types according to the proportions of white and black pigment on the lateral body and the ventral side of the tail flukes. The degree of coverage of yellow colouration on the tail flukes, presumably caused by the diatom Cocconeis, was classified using the same proportional method. [JHumpback Whale, Megaplera novaeangliae, pigmentation, photo-identification, sighting, Western Australia. Christopher L.K. Burton, c/- W.A. Marine Research Laboratories, P.O. Box 20, North Beach, Western Australia 6020; 20 December 1990 Humpback whales (Megaptera novaeangliae) which migrate along the Western Australian coast, comprise the Southern Hemisphere Group TV stock that was severely depleted by commer- cial whaling from an estimated 12,000—-17,000 animals to c. 800 by 1962 (Chittleborough, 1965). Recent aerial surveys show that the stock has been increasing, having at least doubled be- tween 1963 and 1982, and having continued to increase since then (Bannister, this memoir), On that basis the stock may now be more than 2000 animals. Tail fluke photography is a valid means for humpback whale identification (Katona, 1979; Glockner & Venus, 1983; Baker et al., 1986; Stone, Katona & Tucker, 1987). Photographing the lateral body pigmentation patterns in addi- tion to flukes increases the number of individual- ly identified whales (Kaufman, Smultea & Forestell, 1987). Photographic identification of individuals will assist in analysis of migration patterns, distribu- tion and stock identity of this population of humpback whales. Whale watching tours pro- vide an opportunity to photograph many in- dividuals and collect abundance and pod composition data over a short time span within a specific area, Watching humpback whales has become a viable industry on the east coast of Australia, where much research has been condiicted into identification and monitoring of the Area V population (Bryden & Slade, 1987; Bryden, Corkeron & Slade, 1988; Kaufman, Smultea & Forestell, 1987; Paterson & Paterson, 1989). In Hawaii, observations from consecutive years of commercial whale watching have been used to investigate changes in humpback pods and calf encounters (Salden,1988). In Perth, whalewatching tours by Underwater World fram Hillary’s Boat Harbour provideda valuable plat- form for research on humpback whales. This Paper presents in two parts, results from analysis of photographs obtained during 1989 and from data collected during the whale watching cruises. PART 1 - PHOTO-IDENTIFICATION METHODS During July and August 1989, humpback whales were photographed in Shark Bay from a 5.3m runabout, and off North West Cape from a professional fishing boat. From September to November 1989, photographs were taken during whale watching and froma 5.4m runabout. Loc- ations of all sampling areas are shown in Fig. 1, A 35mm SLR camera with a 35-200mm zoom lens was used for all photographs. The larger tour operators’ vessels operators came no 260 VISE 125E ; ra y, f 20s om ’ DAMPIER ARCHIPELAGO Pan NORTH-WEST CAPE WESTERN 14/5 \ SHARK BAY AUSTRALIA ‘\ aTa/ 106 | } aes -) PERTH FIG, 1. Humpback whale observations and photographs, 1989. (observations/photo-ids). closer to the whales than 50m. Interference to the pods was minimised, and often good photographs were obtained when whales them- selves became the curious observers. Data collected included details of pod struc- ture, behaviour, time and photographs. Adults were presumed to be female only if in close association with a calf. Differentiation between adults and subadults was not made. The number of whales observed daily was recorded, as not all whales could be photographed. Copies of the photographs were sent to the Western Australian Museum and to the Pacific Whale Foundation for use in compiling catalogues. Classification of the lateral colouration of individuals follows that described by Kaufmann et al. (1987) (Fig. 2) using the proportion of white pigmentation on the lateral body, with Type 1 having the greatest and Type 4 the least (Fig.3). Whales which did not present sufficient lateral body when surfacing to accurately class- ify were classified as undetermined. Colouration of the ventral sides of the tail flukes was also assessed, using a similar propor- tional method to the lateral colouration, with Type 1 mostly white and Type 4 mostly black. The flukes were divided by a straight line from MEMOIRS OF THE QUEENSLAND MUSEUM TYPE 4 FIG. 2, Lateral body pigmentation types displayed by Australian Humpback Whales. Illustrations by Lili Hagen. courtesy of G. Kaufman, Pacific Whale Foun- dation, TYPE 1, While coloration reaches above the horizontal mid-line of the body. generally extending anterior to dorsal fin, TYPE 2, White coloration ex- tending to body midline or above, with coloration generally observed near the caudal peduncle. TYPE 3, An obvious but less distinct whitish-grey coloration patch along the dorsal surface of the caudal peduncle posterior to the dorsal fin. TYPE 4, Lack of obvious white pigmentation patterns. HUMPBACK WHALE SURVEY, W.A., 1989 261 SS — si = —— FIG. 3. Identified Humpback Whales from W.A., showing 4 lateral pigmentation types. FIG. 4. Tail fluke pigmentation types of humpbacks off WA. 262 MEMOIRS OF THE QUEENSLAND MUSEUM ee r U FIG. 5. Identified humpback whales from WA showing 4 tail fluke pigmentation types. Type 1Y Babel FIG. 6. Photographs of identified whales off WA, showing 3 of the 4 tail fluke diatom colouration categories (yellow). HUMPBACK WHALE SURVEY, W.A., 1989 263 fo me ie FIG. 7. Tail fluke pigmentation and d 12 July, 1989. B, Perth, 27 and 30 September, 1989. 264 the fluke tip to the centre of the median strip between each fluke to assist in determining Type 2 and Type 3 animals (Fig. 4). A Type 2 fluke is categorised as having <50% black pigmentation and a Type 3 fluke having >50% black (Fig.5). Many identified animals had some yellow colouration on the underside of the flukes, presumably due to a film of the Antarctic diatom Cocconeis ceticola. The typically yellowish colour of this species on the skin of whales in warmer waters of the Southern Hemisphere has been taken to indicate movement of the whale from colder waters (Bannister, 1977).The degree of diatom coverage on the underside of the tail flukes was assessed using the proportional method described above for the white/black pig- mentation patterns, with Type 1 having little yellow and Type 4 being mostly yellow. Types 2 and 3 animals have < and >50% yellow colouration, respectively (Fig.6). Tail flukes of animals which were too distant and had uneven lighting, or which were at too low an angle to accurately see the black and white or yellow colouration, were classified as undetermined. RESULTS LOCATION NO. OF WHALES Northwest Cape Dampier Archipelago TABLE 1. Humpback whales off WA during 1989. Observations were made by the author except for the one off the Dampier Archipelago. 417 observations and 69 IDs were made during whale watching tours. ‘Class [Lateral body |Lateral body TFluke ‘orty 17 Total op fluke a Adult Se Calves awe Sarees [Toral_| 18. (15.2%) | 48 40.7%) | 52 (44.1%) [118 | TABLE 2. Humpback whales identifed on lateral or fluke pigmentation patterns. Calves identified by lateral observation as none were observed fluking up. Female MEMOIRS OF THE QUEENSLAND MUSEUM LATERAL PIGMENTATION No. of whales % of types mostly white TABLE 3.Classification of whales on degree of lateral pigmentation. Undetermined whales were identified by body marks, scars or dorsal fin characteristics. FLUKE PIGMENTATION No. of whales pe 3 (>50% black Type 4 (mostly black) Undetermined TABLE 4. Classification on degree of white and black pigmentation on the underside of the tail flukes. DIATOM COLOURATION No. of whales Undetermined 29 a ee ee TABLE S. Classification on degree of yellow coloura- tion (diatoms) on the underside of tail flukes. ee ee es TOTAL 15 24 2 29 TABLE 6. The degree of diatom coverage according to fluke categorisation type. One animal, identified in Shark Bay, was categorised as Type 2 yellow, and was still in that HUMPBACK WHALE SURVEY, W.A,, |989 category when resight ed off Perth 77 days later with very little change in diatom distribution on its tail flukes (Fig. 7). INITIAL SIGHTING No. 2.7.89 9.11.89 | 100 | Perth | 15.11.89] 9.1180] tor] Perth | TABLE 7, Resightings during 1989, Shark B. | Perth Perth DISCUSSION During the present study which focussed predominantly on the southern migration, 41% of animals were identified using only lateral and 44% using only fluke pigmentation patterns, compared to 25% and 17% of the animals of the Area V stock on the east coast (Kaufman er al., 1987), Futhermore, a higher number of iden- tified animals (58%) was obtained using cither lateral or fluke patterns on the east coast com- pared to 15% on the west coast. This large dif- ference may reflect the grealer ease of man- oeuvrability of smaller craft and the persistence of the researchers in obtaining both lateral and fluke photographs ie. a difference in methods. 1989 was the first year in which a concerted effort was made to identify many animals on the west coast. LATERAL PIGMENTATION The 28% Type | and 40% Type 4 of my oDservations compare with 7% Type 1 and 37% Type 4 in eastern Australia (Kaufman e al, 1987). From these initial data, it appears that the Group IV stock has.a larger proportion of Type | animals, FLUKE PIGMENTATION The 70% Type 1 and 95% Types 1+2 of my observations compare with 87% of flukes from 1984,85 animals of the Group V stock being 75% white (Kaufman et al., 1987). In the Northern Hemisphere nearly 62% of animals, including calves, had mostly black flukes (Glockner and Venus, 1983); their Types 4 and 5 equate to my Type 4. Although | identified no mostly black flukes, al least 2 Were photographed by other people (G. Pobar, pers comm). 265 Categorising fluke patterns may help simplify tesight analysis, as cach ID can be placed in a type category rather than being checked against all other animals. DIATOM COVERAGE Assessinent of yellow colouration on specitic whales which may be resighted after their north- em migration may enable quantitative com- parison over time of the change in diatom coverage, Colour photographs of flukes are bet- ler for assessment of diatom coverage than are black and white photographs, Of 41 individuals categorised, 95% had <50% diatom coverage whereas on the east coast there was little evidence of diatom coverage at all (M.Osmond, pers. comm). Difference in diatom coverage be- tween Groups IV and V stocks may reflect geographical differences in the Southem Ocean feeding areas, but further comparisons are neees- sury before making any conclusions. lt is unclear how differences in diatom coverage relate to different fluke colourations. The extent to which diatom coverage may have been reduced by time spent in warm waters is unknown, REstaHTs A mother-calf pair was sighted twice in 7 days in the same area off Perth, while another in- dividual was resighted after 4 days. Many more whales are scen off Perth and south coastal areas from September to December than at other times, Certain population classes, for example, mother- calf pairs, may use the large bays and embay- ments off Western Australia (Shark Bay. Geographe Bay, Perth-Rotinest) for ‘resting’ areas during their southern migration. Photo-ID in Hawaii indicates residence times of up to 11 weeks (Darling,Gibson & Silber, 1983), and in Hervey Bay, Queensland, times of mainly 1 to 2 days, and up to 19 days (Bryden, Corkeron & Slade, 1988), With the development of whale-watching as an industry off the coast of Perth, information on the distribution, behaviour, identification and ecology of whales will be essential, to properly understand and manage the interactions between people and whales. PART 2 - SIGHTING ANALYSIS METHODS Whale-watching tours are conducted from September to November during the southern 266 ——— Tnoviles Sen FIG. 8. The area of Indian Ocean between Perth and Rottnest Island where the whale watching tours were conducted during 1989. migration of humpback whales which occurs between August and December. AREA The Indian Ocean, adjacent to Perth and north of Rottnest Island, was divided into grids of approximately 2 x 1.8 nautical miles to define sightings distribution (Fig 8). Tours were con- ducted west of the shallow limestone reefs which run parallel to the coast, and north of Rottnest Island, generally when wind speeds were below 15-20 knots. MEMOIRS OF THE QUEENSLAND MUSEUM 2nmiles Pein FIG, 9. Numbers of humpback whales (adults + cal- ves)observed in each grid square during the southern migration. VESSEL Two vessels were used for the tours; a 15m aluminium monohull fishing vessel and a 1Om fibreglass shark cat. The larger vessel had radar, and was used on most occasions. The vessels departed from the boat harbour at Hillary's and headed southwest towards Rottnest Island around the southern most Centaur Reef. At least 2 or 3 observers were continually looking for whales, and on most occasions the blow was the initial sighting cue. Data COLLECTION Cruise description data gave starting and en- ding times and weather conditions during obser- vation outside the fringing reefs. Locations of pods were specified by radar or compass bear- ings. A cruise path description was made on a grid as it was not possible to continuously plot each cruise using more accurate equipment. Sighting data consisted of individual whale counts, pod sizes, reproductive status, behaviour and photographs. Searching time included that spent with each pod and assumes that searching continues while with one pod. Searching time with the vessel moving would have been much HUMPBACK WHALE SURVEY, W.A., 1989 ADULTS CI CALVES RN NUMBER OF WHALES CRUISE DAYS FIG. 10. Adults + calves sighted per day during the cruises, from September to December (southern migration), Perth, 1989. less if time spent with each pod had not been included. PHOTOGRAPHIC DOCUMENTATION Efforts were concentrated in obtaining photos of the ventral side of the tail flukes and lateral views of each whale. Whales were inferred to be female if accompanied by a calf; mother—calf NUMBER OF WHALES 9 10 1 12 MONTH FIG. 11. Humpback whales (adults + calves) per daily sighting hour; southern migration, Perth, September- December, 1989 (mean + SD), 267 pairs were obvious, even when other adults were nearby. RESULTS CRUISES ADULTS CALVES “| | REPEAT PODS os TABLE 8. Monthly data collected during whale watching cruises. MONTH DAYS CRUISES WHALES MEAN HR/CRUISE WHALES/HR TABLE 9. Summary of cruise searchtime, number of whales and daily sighting rate. An ANOVA was performed on the trans- formed data (log10) to test differences between months for whales/cruise hour. A significant dif- ference was noted (F=5.32, df=43,P=.0035), and tests (Tukey,Scheffe,SNK) between monthly means indicate that December is different from the remaining months. Removing December also resulted in a significant difference for November (P=.0092) from Sept—Oct. DISCUSSION No comparative data on humpback whale numbers in the near shore waters off Perth are available; estimated population size is more than 2000. It may be that a significant proportion of the population migrate south close to the coast, using the large bays as resting areas. To what extent and duration these bays are used remains 268 FIG.12. Pods of humpback whales observed per day during the cruises; southern migration, Perth, 1989. to be determined. The photo-id work in 1989 indicates residence times of 4—7 days off Perth. The distribution of whales over the gridded area suggests that they limit their approach to the coast to depths greater than 20m. Concentration of sightings in the southern region of the study area reflects cruise paths. A series of aerial sur- vey flights over the area would provide a more accurate estimation of their distribution and give some indication of the proportion of whales which pass al a greater distance from the coast and are not seen by the cruise vessels. The effect of the warm, southward flowing Leeuwin cur- rent along the Western Australian coast on the spatial distribution of these animals may have to be considered. The number of adults decrease fram October. Since the cruises did not com- mence until late September the initial rise in numbers was missed, and may follow a normal disribution curve, similar to that observed in Queensland from shore (Bryden et al., 1987). The number of calves increased in November, as previously noted (Chittleborough, 1965). This earlier work by Chittleborough also indi- MEMOIRS OF THE QUEENSLAND MUSEUM ADULT POD SiZE FIG, 13. Frequency of pod sizes observed during the southern migration, Perth, 1989. N=37 FIG. 14. Frequency of calves with different adull pod sizes observed during the southern migration, Perth, 1989. HUMPBACK WHALE SURVEY, W.A., 1989 cated that the southern migration had passed the Australian mainland by the end of October with a few stragglers remaining in November and December. The numbers of whales secn off Perth in 1989 during November and December were fewer than in October, but still accounted for 17% of observations, Some were also reported in Geographe Bay, south of Perth in November and December; they may be part of the migration which pass close to Perth, Further observations will be necessary to determine if there has been a real change in migratory behaviour compared with that observed by Chittleborough (1965). It is likely that the apparent change in migration over 30 years reflects differences in data collec- tion, On the east Australian coast Paterson and Paterson. (1989) recorded a similar migration pattern in 1987 to that recorded by Chit- tleborough in 1961] but they received reports of some late southward migration during the sum- mer months. However, Stone et al. (1987) have shown differences in the timing of the migration past Bermuda in the North Atlantic Ocean in recent years compared with historical records from the 17—19th centuries. ACKNOWLEDGEMENTS 1 thank Mr Bill Mills and Mr Gavin Barr who skippered both vessels with expertise and regard for the whales during the Underwater World whale-watch cruises, [ sincerely thank Mr Henry Hall, who has given considerable time and effort while taking photographs and recording songs off Perth. Special thanks to Mr Trevor Cook of Carnarvon, who has helped me in a similar man- ner in Shark Bay. Mr Danny Cassidy from Ex- mouth co-operated in obtaining photographs from North West Cape and Mr Ken Maley of Karratha supplied the IB from Dampier. LITERATURE CITED BAKER, C.S.. HERMAN, L.M., PERRY. A, LAW- TON. W.S..STRALEY,.M.WGLMAN,A,A.. KAUFMAN, G.D., WINN, H.E., HALL, J.D.. REINKE, J.M. AND OSTMAN, J, 1986. Migralury movement and population structure of humpback whales (Megaprera novaeangliac) in the central and eastern North Pacific. Murine Ecology 31:105-119, BANNISTER, J\L. 1977, Sperm whales off Albany, Western Australia. Repl. int Whal. Commn 27: 274-276, BANNISTER, J.L. 1988. Report on aerial survey for 269 Humpback Whales in Wesiern Australia 1988. W.A,. Museum report (u the Australian Nationul Parks and Wildlife Service. BANNISTER, J.1.. this memoir, Western Australian hompbacks since 1963, Mem. Qd Mus. 30(2): 258. BRYDEN, M.M. AND SLADE, R.W, 1987, Survey: of Humpback whales, Megaplera novaeangliae, off eastern Australia, 1987, Report, Department of Veterinary Anatomy, University of Sydney, (unpubl. ). BRYDEN, M.M., CORKERON, P.J. AND SLADE, R.W, 1988. Humpback whales in Hervey Bay, Queensland, 1988. Report, Department of Veterinary Anatomy, University of Sydney. (ua- publ). CHITTLEBOROUGH, R.G. 1965. Dynamics of wa populations of the Humpback whale, Megaptera novaeangliae (Borowski). Aust. J. Mar. Freshw. Res, 16: 33-128, DARLING, J.D,, GIBSON, K.M. AND SILBER, G.K. 1983, Observations on the abundance and behavior of Humpback Whales (Meguptere Novaeangliae) off West Maui, Hawaii, 1977-79, 201-222. In Payne, R. (ed.), ‘Communication and behavior of whales’, (Westview Press; Boulder). GLOCKNER, D.A. AND VENUS, S.C, 1983. Iden- tification growth rate and behavior of Humpback Whale (Megaptera novavangliac) cows and cal- ves in the waters off Maui, Hawaii, 1977-79. 223-258, In Payne, R., (ed,), ‘Communication and behavior of whales’. (Westview Press: Boulder). KATONA, S.. BAXTER, B., BRAZIER, 0, KRAUS, S., PERKINS, J. AND WHITEHEAD, H, 1979, Identification of Humpback Whales by fluke photographs. 33-44. In Winn, H.E. and Olla, B.L. (eds), “Behavior of marine animals, vol, 3°, (Plenum Press: New York), KAUFMAN, G.D., SMULTBA, M.A, AND FORE- STALL, P. 1987. The use of lateral body pig- Mentation patterns for photographic identification of east Australian (Area V) hump- back whales. Celus 7: 5-13. PATERSON, R. AND PATERSON, P. 1989, The status of the recavering stock of humpback whales, Megaprera novaeangliae in east Australian waters. Biological Conservation 47: 33-48. SALDEN, D.R. 1988. Humpback whale encounter rates off shore of Maui, Hawaii, J, Wildlife Management 52: 301-304, STONE, G.S., KATONA, S.K. ANB TUCKER. E.B. 1987, History, migration and present status of 270 MEMOIRS OF THE QUEENSLAND MUSEUM humpback whales, Megaptera novaeangliae at Bermuda. Biological Conservation 42: 133- 145.. 262 MEMOIRS OF THE QUEENSLAND MUSEUM 7 4 » = _— - — a a = 2 FIG. 5. Identified humpback whales from WA showing 4 tail fluke pigmentation types. Type 1Y a i —- OEE FIG. 6. Photographs of identified whales off WA, showing 3 of the 4 tail fluke diatom colouration categories (yellow). HUMPBACK WHALE SURVEY, W.A., 1989 a al SS = FIG. 7. Tail fluke pigmentation and diatom colouration photographs of a tesighted humpback; A, 12 July, 1989. B, Perth, 27 and 30 September, 1989. 263 Shark Bay, MEMOIRS OF THE (QUEENSLAND MUSEUM BRISBANE © Queensland Museum PO Box 3300, South Brisbane 4101, Australia Phone 06 7 3840 7555 Fax 06 7 3846 1226 Email qmlib@qm.qld.gov.au Website www.qm.qld.gov.au National Library of Australia card number ISSN 0079-8835 NOTE Papers published in this volume and in all previous volumes of the Afemoirs of the Queensland Museum maybe teproduced for scientific research, individual study or other educational purposes. Properly acknowledged quotations may be made but queries regarding the republication of any papers should be addressed to the Editor in Chief. Copies of the journal can be purchased from the Queensland Museum Shop. A Guide to Authors is displayed at the Queensland Museum web site A Queensland Government Project Typeset at the Queensland Museum A HUMPBACK WHALE, MEGAPTERA NOVAEANGLIAE, SKELETON DISCOVERED AT AN UNUSUAL LOCATION ON MORETON ISLAND, QUEENSLAND V.C. BUSHING Bushing, V.C. 1991 07 01: A humpback whale, Megaptera novaeangliae, skeleton discovered at an unusual location on Moreton Island, Queensland. Memoirs of the Oueensland Museum 30(2): 271—275.Brisbane. ISSN 0079-8835, Chance discovery of a humpback whale, Megaptera novaeangliae, skeleton on Moreton Island is described. Its distance above and behind present high water mark poses conjecture as to the manner and timing of its deposition. Radiocarbon dating indicates that its age is less than 1500 years BP thus excluding the possibility of deposition during the last interglacial high sea level period. It is most likely that the whale was washed ashore during a cyclonic surge or other extreme weather conditions. [JHumpback whale, Megaptera novaeangliae, skeleton, Moreton Island, Queensland, radiocarbon dating. V.C. Bushing, National Parks and Wildlife Service, Department of Environment and Heritage, Queensland 4001. Present address:- Queensland Forest Service, Department of Primary Industries, GPO Box 944, Brisbane, Queensland 4001;20 December, 1990, During a routine patrol on sand dunes behind Spitfire Creek (27°04’S, 153°27’E) on the east- ern or oceanic shore of Moreton Island, on 29 October 1984, large animal bones were seen in a sandblow (Figs 1-4). At first they were con- sidered to be those of a horse but closer inspec- tion and hand-digging revealed large cetacean vertebrae. The site was re-examined on 4 November 1984 and skeletal elements, including a maxi- llary tip, cervical, thoracic and caudal vertebrae as well as the sternum and ribs, were recovered. The skeleton was identified as that of a humpback whale when a radius, characteristic of the species, was located (Fig.5). The skeletal elements were deposited and registered (JM 5252) in the Queensland Museum and com- parison with other Humpback Whale material in that collection (Paterson,1986) indicated that they were those of an adult. The skeleton lay in a SE/NW line in the sandblow which sloped to the northeast (Fig.6). The maxillary fragment and caudal vertebrae lay at the extreme northwest and southeast aspects of the site, respectively. The highest point of the sand dune in which the site was incorporated is 12.6 m and the midpoint of the skeletal deposit, from where the thoracic elements were recovered, is 4.8m above and 190m behind present high water mark. During the last interglacial period, c. 120,000 yr BP (Flint,1971) sea levels were approximate- ly 5m higher than at present. If the whale had stranded at that time it may have been located in this position in normal weather conditions. Radiocarbon dating with correction for isotopic fractionation dates the skeleton at 1410 + 70 yr BP. However, its age may be substantially less because of depletion of C'* in the Antarctic oceanic water mass. This phenomenon is termed the reservoir effect and a correction is applicable to material of Antarctic origin (Omoto,1983). Although Southern Hemisphere humpback whales spend considerable periods in temperate waters (Dawbin,1966) their periods of rapid skeletal growth occur at the Antarctic feeding grounds (Matthews,1937). Accordingly, reser- voir effect correction of JM5252 is required. Freshly killed seals and penguins in the Ross Sea had radiocarbon dates ranging between 1750 + 70 and 1300 + 50 yr BP (Omoto, 1983). It is clear that JM 5252 does not date from the interglacial period and other reasons for its location must be considered. Anatomical distribution of skeletal elements in the sandblow excludes the remote possibility that they were relocated by humans from a site nearer to the sea. Forman et al., (1987) observed that whale bones are gradually scattered along the littoral zone by tide and weather action. The relatively undisturbed skeleton of JM 5252 sug- gests that it had not been subjected to further significant tidal action. The Spitfire Creek depression is the last sub- stantial breach in the present frontal dune system proximal to the Cape Moreton headland (Fig. 1) and the sea enters its easterly aspect in rough, southeast weather. In the winter of 1957 water 272 MEMOIRS OF THE QUEENSLAND MUSEUM FIG.I. Spitfire Creek, Moreton Istand. Cape Moreton is seen in the far right of the photograph. The sand-blow in which JM 5252 was found is in the centre of the photograph, FIG.2. The sand-blow which contained JM 5252. A view seawards from its northwest aspect. The thoracic elements were recovered from the region of the marking stick. UNUSUALLY LOCATED HUMPBACK ON MORETON IS. 273 FIG.3. The sand-blow which contained JM 5252. A view from the present low frontal dunes to demonstrate the extent of the Spitfire Creek depression. FIG.4. Skeletal elements of JM 5252 at the time of discovery. was trapped to a depth of approximately I.5m behind a barrier of logs and dune grass at the entrance to Spitfire Creek (R.A. Paterson, pers. comm.). This debris had accumulated during a prolonged period of rough weather associated with strong southeast winds. Decomposing whales are extremely buoyant, particularly if their integument is intact (Tonnessen and Johnsen, 1982). It is suggested that the most likely reason for the unusual loca- tion of JM 5252 is that its carcass was washed into the Spitfire Creek depression during cyclonic or other extreme weather conditions associated with abnormally high tides. 274 MEMOIRS OF THE QUEENSLAND MUSEUM FIG.6. The sand-blow which contained JM 5252. Its northern aspect is to the right of the photograph. UNUSUALLY LOCATED HUMPBACK ON MORETON IS. ACKNOWLEDGEMENTS Peter Jell of the Queensland Museum kindly atranged for radiocarbon dating of JM5252 which was performed by Beta Analytic Inc. Ms Gillian Taylor of the N.W.G. MacIntosh Centre for Quaternary Dating kindly provided informa- tion concerning the Antarctic reservoir effect. Mr Neville Hawkins transported the skeletal material from Moreton Island and Dr Robert Paterson assisted with the excavation and iden- tified the specimen. LITERATURE CITED DAWBIN, W.H. 1966. The seasonal migratory cycle of humpback whales. 145-170. In K.S. Norris (ed.), ‘Whales, dolphins, and porpoises’. (University of California Press: Berkeley and Los Angeles). 275 FLINT, R.F. 1971.'Glacial and Quaternary geology’. (John Wiley and Sons: London). 339p. FORMAN, S.L., MANN, D.H. AND MILLER, G.H. 1987. Late Weichselian and Holocene relative sea-level history of Bréggerhalvdya, Spitsber- gen. Quaternary Research 27: 41-50. MATTHEWS, L.H. 1937. The Humpback Whale - Megaptera nodosa. Discovery Reports 17: 7— 92. OMOTO, K. 1983. The problem and significance of radiocarbon geochronology in Antarctica, 450— 452. In R.L. Oliver, P.R. James and J.B. Jago (eds),‘Antarctic earth science’. (Australian Academy of Science: Canberra). PATERSON, R.A. 1986, A list of specimens of the Order Cetacea in the Queensland Museum. Mem. Qd Mus. 22: 309-312. T@NNESSEN, J.N. AND JOHNSEN, A.D. 1982. ‘The history of modern whaling’. (Hurst: Lon- don). MEMOIRS OF THE (QUEENSLAND MUSEUM BRISBANE © Queensland Museum PO Box 3300, South Brisbane 4101, Australia Phone 06 7 3840 7555 Fax 06 7 3846 1226 Email qmlib@qm.qld.gov.au Website www.qm.qld.gov.au National Library of Australia card number ISSN 0079-8835 NOTE Papers published in this volume and in all previous volumes of the Afemoirs of the Queensland Museum maybe teproduced for scientific research, individual study or other educational purposes. Properly acknowledged quotations may be made but queries regarding the republication of any papers should be addressed to the Editor in Chief. Copies of the journal can be purchased from the Queensland Museum Shop. A Guide to Authors is displayed at the Queensland Museum web site A Queensland Government Project Typeset at the Queensland Museum SONGS OF HUMPBACK WHALES: THE AUSTRALIAN PERSPECTIVE DOUGLAS H. CATO Cato, D. 1991 07 01: Songs of Humpback Whales: the Australian perspective. Memoirs of the Queensland Museum 30(2): 277-290. Brisbane. ISSN 0079-8835. Humpback whale songs have been recorded in Australian waters since 1979, mainly off the east coast and generally these have shown the complex and stereotyped structure associated with Humpback Whale songs observed in other parts of the world. Detailed study of the 1982/83 song off the east coast demonstrated a stereotyped, repeating song and established “rules” that governed the song for that period. These rules are broadly similar to those established for the Northern Hemisphere with differences in detail. The sounds that make up the song are also different. The song has changed progressively with time. In some years this has been gradual, but in 1984 the change was so substantial that the 1985 song bore little resemblance to that for 1982-84, and was relatively unstructured. Studies in Australian waters provide a somewhat different perspective to those of the Northern Hemisphere because of differences in the environment and the access to whales. Whereas most Northern Hemisphere recordings have been obtained on the breeding grounds (with the notable exception of the Bermuda area) we have access to the whales along the migration paths which extend for thousands of kilometres along the east and west coastlines. From our observations, the integrity of the song appears to be maintained over large distances (1500 km) of the migration paths, although the sample size is small. However, songs off the east coast are distinctively different to those off the west coast.) Humpback Whales, songs, marine acoustics, animal behaviour. Douglas H. Cato, Materials Research Laboratory (MRL), DSTO PO Box 50, Melbourne, Victoria 3032; 3 January, 1991, Humpback Whale songs have been studied extensively for stocks in the Northern Hemi- sphere. Payne and McVay (1971) first demon- strated the existence of a structured, stereotyped and repeating song from recordings of migrating whales off Bermuda, and Winn, Perkins and Poulter (1971) observed similar songs in the West Indies. The song is repeated for long periods and appears to be stereotyped between singers of the same geographical breeding stock, but different oceans have different songs (Payne, 1978; Winn and Winn, 1978; Winn et al., 1981). In addition, the songs of any stock gradually change with time (Winn and Winn, 1978; Payne and Payne, 1985). Sounds at- tributed to Southern Hemisphere Humpback Whales were first recorded in the late 1950's (Kibblewhite et al., 1967), but song analysis has been reported for only one recording (Winn et al., 1981), apart for some preliminary work on the east Australian song (Cato, 1984). With the notable exception of the Bermuda recordings, much of the work in the Northern Hemisphere has been in tropical breeding grounds such as Hawaii and the West Indies, and discussion has often focussed on the significance of the song to breeding activities. In the North Pacific and North Atlantic Oceans, Humpback Whales are more readily accessible on breeding grounds than on the migration paths. The situa- tion is reversed in the Australian region, where migration paths follow coastlines for thousands of kilometres, and breeding grounds are diverse (Chittleborough, 1965; Dawbin, 1966). Recent sightings of humpbacks extend along the entire coastline of eastern Australia (Paterson and Paterson, 1984, 1989; Simmons and Marsh, 1986). Known concentrations of whales are along the migration paths rather than on breeding grounds. This paper discusses songs recorded between 17°40’S and 35°20’S on the east coast and between 19°40’S and 32°S on the west coast. While some results are from breeding grounds, most were obtained on migration routes. There is some interest in the songs of Southern Hemisphere Humpback Whales. It seems that stocks in the two hemispheres are well segre- gated because most stocks do not usually cross the equator and because migrations are seasonal and thus six months out of phase. Songs in the two hemispheres may have evolved inde- pendently over a considerable period of time. MONTE BELLO,.---1 198 6 ‘ ABROLHOS —% . 1990 i: MEMOIRS OF THE QUEENSLAND MUSEUM JERVIS BAY 1979, 80, 84 FIG, 1. Map showing the location and year of the song recordings discussed in this paper. The first Southern Hemisphere recordings of Humpback Whale songs were obtained off New Zealand during 1958-1963 by Kibblewhite, Denham and Barnes (1967). They observed in- tense activity during April-September, the time of year when most Humpback Whales might be expected in these latitudes (Dawbin, 1966). During 1960-1963 sound activity decreased markedly, coincident with the similarly marked decline in the Humpback Whale population. No analysis of song patterns has been published. This paper reviews some results of studies of Humpback Whale songs in Australian waters with particular reference to differences in perspective that the Australian region provides, and how these may lead to a better understanding of the song and its purpose. Samples of songs discussed in this paper are available on compact disc entitled “Songs of the Humpback and Frog Calls of Brisbane and En- virons” issued by the Queensland Museum. METHODS Data RECORDING Sound recordings have been made at a number of locations on migration paths and in breeding grounds along Australian coastlines including those discussed in this paper (Fig. 1). These were single session recordings of a few hours dura- tion, except off Point Lookout, Stradbroke Is. (27°25’S), where data have been recorded on a number of days every year since 1981. Since 1982, recordings have been made off Coffs Har- bour (30°18’S) and single session recordings have been made at other locations by an Australian Museum team (W.H. Dawbin, pers. comm.; Dawbin and Eyre, this memoir). Recordings off Stradbroke Is. were coor- dinated with the observations of Paterson and Paterson (1984, 1989), and were timed to coin- cide with the peak of the migration past this point. Recordings have been made during the southern migration in each year (late September, HUMPBACK WHALE SONGS early October) and during the northern migration (late June, early July) in 1984, 1989, and 1990. Stradbroke Is. is at the most easterly part of the east. coast, and this is where the migration paths converge. On the basis of experience during whaling activities in the area, most humpbacks could be expected to pass within 15 km of the shore (Chittleborough, 1965). When this study commenced, this was the largest known con- centration of Humpback Whales on the east coast. Even so, long periods of monitoring were required to obtain even a small sample of the song because of the small population at the time (<1/2 the present number;see Paterson and Pater- son,1989), As a consequence, sample sizes during the early years of recording were small. Recordings off Stradbroke Is. were made from a small boat which was allowed to drift in the vicinity of the whale paths off Point Lookout. Whales are in transit at this point, and although they show some meandering, there are only oc- casional departures from general north or south passage through the area (Cato, 1984). This is an exposed oceanic coast and weather conditions have been unsuitable for recording on about half the days allocated to monitoring. The recording system consisted of a hydrophone connected via a high pass filter to a low noise preamplifier, and from there to a tape recorder. The preamplifier provided a sufficient- ly high input impedance to give adequate low frequency response. During recordings, it is desirable to minimise any water flow past the hydrophone, because the pressure fluctuations of the flow, although not acoustic, are detected by the pressure sensitive hydrophone and appear as low frequency noise on the recording. Under some conditions they may be of sufficient level to overload the preamplifier and cause distor- tion. There is little problem if, as in the absence of wind, the boat and hydrophone drift with the water mass. Usually, however, there is sufficient wind drift to cause some motion of the hydrophone relative to the water. In most cases it was necessary to use the high pass filter to attenuate the low frequencies to avoid overload- ing the amplifier. Since this filter has a gradual toll off with decreasing frequency, it was pos- sible to correct for its effect in the measurements from tape replay. By experimenting with fairings on the cable, it was found that string wound spirally along the cable substantially reduced the noise from waterflow, presumably by disrupting the shedding of vortices. This was effectively from 1987. 279 During 1981-1983 the system used was a General Instrument Corporation Z3B_ hydro- phone on 30m of cable, RAN Research Lab- oratory (RANRL) designed low noise pre- amplifiers and a Kudelski Nagra III tape re- corder, System response was +3dB from 20Hz to 17 kHz, but it was often necessary to use a high pass filter (-6 dB at 55 Hz, —20 dB at 20 Hz) to attenuate low frequency noise from turbulence. From 1984, Clevite CH17 hydrophones and Sony WMD6 or TCDSM cassette recorders were used, giving a system response of 30 Hz—15 kHz, modified by the above filter response when used. Data ANALYSIS Sonagrams (spectrograms) were produced using a Kay Elemetrics Corp. Digital Sonagraph and a Spectral Dynamics SD-311 analyser. Spectral measurements were made using a Hewlett-Packard 3582A analyser and Bruel and Kjaer 2112 analyser and 2305 level recorder. Payne and McVay (1971) defined a “sound unit” as any sound that is continuous to our ears when heard in “real time” (i.e., when replayed at the same tape speed as recorded). These are the individual sounds, or elements of the song. Winn and Winn (1978) used the term “syllable” for a similar purpose. A Humpback Whale song cycle comprises hundreds of sound units but it can be shown that these can be categorised as belonging to one or other of a small number (typically 12-15) different sound types. The term “sound type” is defined here to mean the particular type of sound that a unit belongs to. One sound type is distinguished from another by having different acoustical characteristics. These characteristics can be measured by various methods of analysis, to demonstrate that the sound types are distinctly different. The characteristics of Humpback Whale sounds are such that they are well suited to human aural perception. Thus, with a little experience we can easily distinguish between different sound types by ear. To simplify discus- sion, descriptive names are used to identify sound types, e.g. “growl”, “moan”, “whistle”. The choice of a name is, of course, subjective but once chosen becomes the definitive name for that particular sound type . It is convenient, in this definition, to allow sound units of a par- ticular sound type to show some variation in character, so long as this variation is small com- pared with the variation between different sound types. The characteristics of the sound units of a sound type may change with time. If the change exceeds the acceptable variation in the sound 280 MEMOIRS OF THE QUEENSLAND MUSEUM FREQUENCY (kHz) 1 TIME (seconds) (b) VIOLIN FREQUENCY (kHz) TIME (seconds) 2 (c) GROWL FREQUENCY (kHz) TIME (seconds) FIG. 2. Sonagrams of two samples of the following sound types: (a) “moan”, (b) “violin” and (c) “growl”, from theme 5 of the 1983 song off the east coast. The filter bandwidth was 11.3 Hz. type characteristics, a new sound type must be Because sound types are so readily distin- defined. Sounds type names can be identified guished by the human ear, a major part of the with particular years by affixing the year, sothat analysis consists of listening to the tapes and “moan(82)” would apply to 1982 only. noting the sequence of sounds using some shor- GROWL GROWL GROWL GROWL VIOLIN VIOLIN MOAN MOAN (ZH4) AINANDAYS TIME (s) HUMPBACK WHALE SONGS 281 FIG. 3. Sonagram showing how the sound types of Fig. 2 go together to form one complete phrase of theme 5 of the 1983 song off cast Australia’ The filter bandwidth Wasl 1.3 Hz. thand notation. From this, the song structure or pattern can be determined, The sounds are also displayed visually as sonagrams which show as a grey scale plot the sound intensity as a function of frequency and time (Figs 2,3), The distinction between sound types and the way they are ar- ranged to form the sang pattern can also be determined from sonagrams. The two methods complement each other. Some subtleties detec- table by ear are difficult to display or distinguish on spectro-graphic analysis. On the other hand, quantitative comparisons in frequency and time are possible on sonagrams. Sound types can be more readily detectable by aural analysis when the signal to noise ratio is poor (as is the case of distant whale sounds), and a significant propor- tion of the data can only be analysed effectively by ear. Generally, aural and visual analysis produce consistent results. In the one case we use our aural perception for acoustic analysis and interpretation, in the other we use an instrument to do the analysis and our visual perception for the interpretation. The ear is an effective acoustic analyser, but our aural perception is not equally sensitive to all measurable characteristics of a sound. On the other hand, our instruments have limitations, and their qutputs require some as- sessment, such as the visual interpretation of the sonagram. The sonagram, for example, does not contain all the information in the original sound If we are interested in the importance of the sounds to the whales, we would like to know what characteristics. of a sound are detected by the aural perception of a whale. This, of course, is not known. It may be, however, that our aural perception of a sound rather than our visual perception of a sonagram, is closer to the whales’ aural perception of the same sound. If we are interested in other aspects of the sounds, such as the mechanism of production, and how they propagate through water, the measurable physi- cal characteristics of the sounds may be more useful than our aural perception. PROPAGATION OF HUMPBACK WHALE SOUNDS AND THEIR DETEC- TION AND RECOGNITION The question of how far Humpback Whale sounds are detectable is of some importance ina study of the song, because it determines the tange over which the song can have an effect, whatever its purpose. Unfortunately, because of the nature of sound propagation in water, there is no simple answer to this, Sound is a fairly 282 localised phenomenon in air because the high absorption attenuation causes the level to fall rapidly with distance from the source. Such at- tenuation is very much lower in water (at least at the frequencies of Humpback Whale sounds) so that sound levels fall far more gradually with distance. A doubling of distance in typical ocean conditions results in a decrease in level of about 6 dB, whether the doubling is from 100-200m, or from 5—10 km. The detectability depends on the signal to noise ratio at the receiver, that is, the ratio of the whale sound level (the signal) to that of the background noise level. A variation in the signal to noise ratio of 10 db would not be unusual, either through variation in propagation conditions or in the level of the background noise. This corresponds to a variation of a factor of three in detection range. Thus on this basis alone, the uncertainty in estimating the distance to which the sounds are detectable is a factor of three. The other factors required in this calculation are the source level and the minimum signal to noise ratio for which detection is possible ( the detection threshold). Winn et al. (1971) measured broad band source levels of different sounds from the song in the West Indies to range 175-188 dB re 1Pa~ at lm. Thompson, Cum- mings and Ha (1986) measured source levels of non-song sounds of comparable level. Because of the difficulties of identifying singers and measuring their distances during our recordings, we have no estimates of source levels of sounds in Australian waters. We can, however, estimate a lower limit by using the closest possible es- timates of the distances of whales during record- ings, where this is known. The results support the estimates of Winn et al. Variation in the received level between sound types of the 1982 song recorded from one singer off Stradbroke Is. was about 10 dB. This result and the measurements of Winn et al. indicate that variation in the source level of different sound types from one singer would cause the detection distance to vary by a factor of 3-5. Greater variation would occur if source levels varied between individuals. Under typical ocean conditions, the distance the sounds would propagate for the levels to fall to that of the average background noise, i.e to give a broadband signal to noise ratio of O dB is about 5.6km for the lowest source levels measured by Winn et al., and 25km for the highest source levels. However, the sounds would actually be detectable at significantly lower signal to noise ratios and thus much greater MEMOIRS OF THE QUEENSLAND MUSEUM distances because the acoustical characteristics of the sounds are distinctively different from the background noise. Just how much further the sounds can be detected depends not only on these relative acoustical characteristics, but also on the auditory perception if the receiver is an animal, or on the signal processing gain if the instrumen- tation is used. A simple experiment in the detec- tion of some Humpback Whale sounds with varying signal to noise ratios by a human subject showed that the threshold of detection occurred at a signal to noise ratio of about -14 dB, cor- responding to an increase in detection range of a factor of about five. This was not a rigorous, controlled experiment, and the result should be used only as an indication that detection by a human listener would be at substantially greater distances than the values at which the signal to noise ratio falls to 0 dB. It seems likely that whales are also able to detect the sounds when signal level is less than noise level. It seems likely, therefore, that Humpback Whales should be able to detect and recognise the song from other individuals for large distan- ces, probably some tens of kilometres, but detec- tion distance will vary significantly under varying conditions, and the actual values cannot be determined without knowledge of the capabilities of the auditory perception of the whales. This may be an important factor in the significance of the song. If the song is detectable for distances of tens of kilometers, behavioural reactions in response to the song may also occur over such distances. The possibility needs to be investigated and the Australian coastlines pro- vide the opportunity for such studies along migration paths. EARLY RECORDINGS The first recordings of Humpback Whale sounds in Australian waters were made by Mil- lington and Sneesby off Jervis Bay in October 1979 and in the same area in October 1980. These recordings were obtained incidental to other work and were passed to the author for analysis. In 1979 two whales were observed by aircraft at the time of the recordings (the actual species was not identified). They were estimated to be about 10 km from the hydrophones. There is no direct evidence to indicate that the singer was one of these. However, the recorded sound levels were consistent with a source at that dis- tance, based on the source levels of Winn et al. (1971). Also, given the size of the population in HUMPBACK WHALE SONGS 1979, and the evidence that southward migration paths tend to broaden outwards from the coast well before the latitude of Jervis Bay is reached, itis unlikely (although of course not impossible) that there would have been another whale in audible range. A sample from the 1979 record- ings is given on track 1 of the compact disc. The 1979 recordings are characteristic of an intense, distant source. There is some waxing and waning of the signal consistent with varia- tion in propagation loss and some echoing char- acteristic of deep water. The background noise was characteristic of the deep ocean, being predominantly from sea surface motion and dis- tant shipping (Cato, 1978). As such, it is charac- teristic of broadband noise — rather featureless, so allowing the distinctive Humpback Whale sounds to be more readily detected than in an environment where a variety of other sounds are present. The 1979 recordings comprised about four hours of song. Several different sound types were evident and there was evidence of repeti- tion or cycling over periods of minutes, consis- tent with the Humpback Whale song structure. At the time, identification of the sounds, as due to Humpback Whales, relied on comparison with the extensive knowledge of sounds from other marine animals. The acoustical characteristics of the sounds are such that they are beyond the capability of fish and are indicative of marine mammals. Of the known whale sounds, only those of the right whale have similar acoustical characteristics. However, recordings by the author of sounds of a right whale five weeks earlier (September, 1979) off Wollongong, were different, and showed no evidence of the cyclic and structured characteristics of the sounds recorded off Jervis Bay and typical of the Humpback Whale song. The 1980 recording was about 15 minutes duration, and sound types were generally similar to those of the 1979 recordings. Given the extensive data now available of Humpback Whale sounds off east Australia, there can now be little doubt that the 1979 and 1980 sounds are from humpbacks, Even though the actual sound types are different to any recorded later, their characteristics fall within the same range of variability observed for later recordings. There are significant and regular gaps in the 1979 and 1980 recordings. This is to be expected for sounds from a distant Humpback Whale. Measurements of source levels of Humpback Whale sounds by Winn et al. (1971) showed a 283 range of 13 dB from the sounds of the highest source levels to those of the lowest source levels. Given a typical propagation loss of 6 dB per distance doubled for deep water, 13 dB propaga- tion loss corresponds to a ratio of propagation distances of four to five. Thus, the higher source level sounds of Winn’s data would be detectable at four to five times the distance of the sounds of lower source levels, other things being equal (detectability will also depend on the actual char- acteristics of the sounds and their relationship to those of the background noise). Although the sounds recorded by Winn et al. (1971) are dif- ferent to those recorded in Australian waters, it is evident that high source level sound types will be audible at significantly greater distances than those of lower source levels. From song record- ings in later years it is apparent that as the signal levels fall relative to the background noise, the lower source level sound types cease to be detected. Gaps in the data appear in a manner similar to those of the 1979 and 1980 recordings. Since some of the sound types are missing in the 1979 and 1980 data, it has not been possible to make a complete analysis of the song struc- ture. It is clear, however, that the sound types in these songs are unlike those in later recordings off the east coast of Australia, the first of these being off Stradbroke Is. in September, October 1981. This poses the intriguing question: why are the 1981 recordings so different to those of 1979 and 1980? Was there a substantial change in the east Australian song in the 11 months between the 1980 and 1981 recordings, or were the 1979 and 1980 songs alien to this stock? Since we have never observed alien songs in later recordings, and the 1979 and 1980 songs are very similar, it seems unlikely that they were alien songs, and more likely that the song did undergo a substantial change. We cannot answer this question without further data from that period, but that, of course, is not possible. These songs are now extinct. The next recordings were obtained off Stradbroke Is. in September, October 1981. Al- though song data totalled about 65 minutes not all sound types were audible. In July 1982, Dr Dawbin recorded about 20 minutes of song off Cape Byron (pers. comm.). ESTABLISHMENT OF THE SONG PATTERN AND CHARACTERISTICS To establish the existence of a repeating stereotyped song, and to determine the rules 284 governing the pattern for comparison with the songs of the Northern Hemisphere, requires the analysis of many song cycles recorded from a number of individuals. Although there have been recordings of Humpback Whale sounds in Australian waters since 1979, the first data that could be considered adequate for this purpose were not obtained until September, October 1982 off Stradbroke Is. during the south bound migration. This comprised 27 song cycles on 26 September, 7 on 28 September, and | on 1 Oc- tober. Earlier recordings were either too short in duration or not all sound types were detectable. THEME SEQUENCE OF SOUND UNITS | TYPE OF IN PHRASE PHRASE MOAN | 6-11 WHISTLES REPEATING SIGH MOAN 2 3-6 VIOLINS BER ASE SIGH MOAN VIOLIN 3 2 SQUEALS REPEATING) SHORT SQUEAL SQUEAL 1,2 MOANS 4 1,2 VIOLINS REPEATING 4-8 GULPS 2 MOANS 5 1,2 VIOLINS BREESE 3-6 GROWLS 2 MOANS SIGH Several CHUGS,2 SIGHS 6 (repeats) EVOLVING CHAINSAW (repeats) Several YAPS,CHIRPS (repeats) TABLE 1. Basic structure of the Humpback Whale song off east Australia in 1982 and 1983. The se- quence proceeds down the table. The phrases of themes | to 5 repeat a number of times to form the theme. There is only one phrase of about 20 steps in theme 5. The song pattern was initially determined from the 4hrs 15mins of data recorded between 0740 and 1240 on 26 September 1982 off Stradbroke Island. Sounds were audible from the time listen- MEMOIRS OF THE QUEENSLAND MUSEUM ing commenced until it ceased (because of deteriorating weather conditions) so the period of continuous calling may have extended well beyond Shrs. In all, 27 song cycles were recog- nisable, of which 18 were recorded in full. Seven cycles were incomplete because of gaps in recording either due to repositioning the boat or by loss of signal as another boat passed by. The gaps were short enough (equivalent to one to three themes in length) for the continuity of the song to be followed, i.e. the duration of the gap equalled the usual duration of the missing themes. The remaining two cycles were incom- plete because of an aberration in the singing. The song jumped from theme 3 of one cycle to theme 2 of the next, with a fragment of theme 6 separat- ing the two, and with no apparent pause in sing- ing. There was an additional gap of 30 min possibly representing 3 cycles of the song, sug- gesting that there might have been a total of 30 cycles over Shrs. Song cycle durations (com- plete) varied from 7 1/2—11 1/2 mins, the average being 9 1/4 mins. During the period of recording, two groups, each of three whales were observed at distances varying from a few hundred metres to a few kilometres. One group included a calf, however, a comparison of the received signal levels, with the estimates of source level of Winn, Perkins and Poulter (1971), indicated that the other group of three was responsible for the song. For a short period a more distant song was audible. The general pattern of the song established from these recordings is shown in Table 1. The choice of theme 1 as the start of the song was based on the following criteria, using data from this and later years. On 10 out of the 11 occasions when we heard a whale stop singing at the end of a song session, it has been at theme 6 or the equivalent theme in later years. The eleventh occasion was a rare example in our experience of aberrant singing where themes had been omitted in the two cycles preceding the cycle in which the session stopped (at theme 2). On the one occasion when we observed the start of a song session, it began with theme 1. Usually the song in 1982 progressed from theme to theme without pause, with the excep- tion of the transition from theme 6 to theme 1 where a pause was sometimes, though not al- ways, observed (data from later years showed a more pronounced pause at this point). Often the sound intensity diminished towards the end of theme 6. On the basis of Northern Hemisphere work, this is indicative of the point in the song HUMPBACK WHALE SONGS where the whale approaches the surface to breath, and is thus a further criterion of the end of the song. A reduction in intensity 1s to be expected as a Singer approaches the surface be- cause of the interference between the sound waves incident on (stnking) the surface and those reflected from it, since they are out of phase, Winn el al, (1971) and Winn and Winn (1978) were able to localise on singers in the West Indies using a directional shipboard sonar, and found that singers came to the surface to breathe at a particular part of the song, Tyack (1981) noted the reduction in sound intensity that occurréd at this point of |he song and used this a5 a means of identifying the singer. lt may, however, be misleading to talk of the start and end of a song cycle. Theme 6 could alternatively be viewed as a link between cycles and it is interesting to note that a song session usually stops part way through theme 6 rather than al the end. The structure of theme 6 also differs from that of the other themes. In structure, the 1982 song broadly follows the rules established for the songs of the Northern Hemisphere. The themes are emilted in the order shown (with rate exceptions) and cach com- prises a variable number of repetitions of a phrase specific to that theme, except that theme 6 has only one phrase. Phrases comprise a num- ber of sound units in a fixed order separated by gaps of silence which vary from <1/2 sec. to 3 secs. The duration of the Sound units varies from O.lsecs for the “whistles” to 4.3 secs for the “chainsaw”. There is some variation in the num- ber of repetitions of certain sound types in a theme (Table 1), Theme 6 differs from the others in that it contains only ane long phrase of about 2 mins duration with about 20 steps. Early steps com- prise several “chuggs ” and terminate wilh one or two “sighs”. Following steps consist of one long sound unit —“chain saw” — rather like the merging of several “chuggs"” to farm one con- tinuous sound. The final steps consist of several discrete units — the “yaps” — and terminate in “chirps”, Themes 1 to 5, on the other hand, contain a number of repetitions (up ta 12) of short phrases of 15-25 secs duration cach com- prising a number ( 5-13 ) of sound units, There are more subtle patterns within the basic pattern (Table 1), and some minor variations, These will be discussed in a later paper, This basic song pattern was generally adhered to in the 40 song cycles from five individuals recorded in September and October 1982. The only exception was the omission of themes in two consecutive cycles out of the 27 recorded on 26 September. Themes 4 and 5 wert missing from one cycle, theme 1 from the next. Other- wise, theme order and phrase structure was n- changed, Songs recorded off Stradbroke Is. in Septem- ber and October 1983, and off Heron fs. in Oc- tober 1983 were similar to those recorded a year before with only minor changes. There was su little change in the song pattern. that the structure in Table 1 can be considered to apply equally to both the 1982 and 1983 songs. Some of the sound \ypes showed changes in character but were still clearly recognisable as variants on the ]982 ver- sions, Sonagrams of samples of three of the sound types from the 1983 song (Fig. 2) have two conseculive samples of each sound type, all from a phrase of theme 5. Note that acoustical charac teristics ot the sound types are different, but each sound type shows some variation between the two samples, though much less than the dif- ference between sound types. Both “moan” and “violin” are harmonic sounds and would be per- ccived as having characteristics similar to musi- cal sounds, The fundamental frequency is equal to the interval between the harmonics and this is substantially higher in the case of the “violin”. so that it is perceived as having a higher piich. The frequency (and thus the pitch) is relatively steady in the case of the “violin” but varies significantly over the duration of the “moun”. initially falling, then rising. The “growl” has different acoustical characteristics consisting of a rapid succession of broad band pulses. Figure 3 shows the sound types of Fig. 2 as they occur together to form a phrase of theme 5 (1983). A sample of the 1983 song commencing with theme 2, is given on track 2 of the compact disc. There are two phrase renditions of theme 2, f of theme 3, 1 of theme 4, and 3 of theme 5 in this particular sample. Theme 6, as discussed above, has only one long phrase. The song ends al the “chainsaw” of theme 6 where the whale stopped singing. The sonagrams of Figs 2 and 3 were taken from this particular sample, The recording was made off Point Lookout, Stradbroke Is, on 29 September 1983 in the presence of seven whales (Cato, 1984). CHANGE IN THE SONG WITH TIME Over the 12 months from September 1982, the 286 east coast song changed in the character of some of the sound types and in the song pattern. Not all sound types are audible in the recordings of September 1981 so the pattern for only about 2/3 of the song can be determined. This part of the song is, however, similar in pattern, to the song of 1982, and the audible sound types are similar in character to their 1982 versions. The song recorded in June, July 1984 (four singers, on separate days) was generally similar to that of 1982/83 in pattern, and changes in the character of the sound types were small. From July to September 1984, however, the song changed substantially. Recordings in early Oc- tober and November (two singers, on separate days) showed that themes 3 and 5 were much diminished in duration, and the character of the sound types in themes 1, 2 and 6 had changed significantly. By 1985 the changes were so pronounced that it was difficult to recognise similarities to the 1982/83 song (eight singers, over three days). Themes 3 and 5 were extinct, and many of the sound types of the other themes had changed almost beyond recognition - they had to be considered to be new sound types, even though they may have evolved from the old ones. Themes | and 2 became relatively unstructured. Theme 6 now occupied 1/2 the song, partly be- cause of reduction in the earlier themes and expansion of theme 6. The result was a shorter, significantly more poorly structured song than in 1982/83, Thus, while the song showed a relatively gradual change from late in the singing season of 1981 to early in the season of 1984, the change during the season in 1984 was substantial. By late 1985 song was quite different to that of 1982/83, not only in song pattern and character of the sound types, but also in form of song, being relatively poorly structured. Mednis (this memoir) shows from an analysis of songs recorded off Stradbroke Is. in 1988, that the song later returned to the well structured form of 1982/83. As discussed above, the sound types observed in the 1979 and 1980 songs off the east coast were different from those of 1981 and later years. The possibility that this was the result of a sub- stantial change in the song in the 12 months from October 1980, thus seems more plausible, in view of the substantial change observed in 1984, A sample of the 1985 song is given on track 3 of the compact disc. Other singers are audible in the background. This sample is one of the few examples where the last theme (the one that MEMOIRS OF THE QUEENSLAND MUSEUM evolved from theme 6 of 1982/83) is repeated, so it is rather longer than most song cycles of 1985. GEOGRAPHICAL FACTORS IN SONG CHARACTERISTICS AND SINGING BEHAVIOUR Studies of Humpback Whale songs in the Northern Hemisphere have tended to con- centrate on the tropical breeding grounds, with the notable exception of Bermuda (32°20’N). In the north Pacific and North Atlantic, Humpback Whales are most accessible on breeding grounds, because of the congregations in near shore shallow waters such as Hawaii and the islands of the West Indies. Migration paths, on the other hand, tend to be well off shore, and it is only where they approach islands such as Bermuda that there have been significant record- ings on the migration paths. Studies to relate song production and behaviour have con- centrated on breeding grounds, notably Hawaii (Tyack, 1981, 1983). The situation is reversed in Australian waters. We have most ready access to the whales along the migration paths which follow the thousands of kilometres of the east and west coastlines. Breeding grounds on the east coast appear to be between the Great Barrier Reef and the coast (Paterson and Paterson, 1984, 1989; Simmons and Marsh, 1986). On the west coast they appear to lie in the shelf area off northwest Australia. No concentration of whales on either breeding ground has been discovered. As a consequence, most recordings of the song have been from the migration paths, with very little from what could be considered to be breeding grounds (only the two most northerly points on the east coast (Fig. 1) could be considered to be on the breeding grounds). Stradbroke Is. was chosen as the place to record the song when this study commenced in 1981, because this was where a significant concentration of whales were known. More recently, a concentration of humpbacks in Her- vey Bay during the southern migration has been established (Bryden et al., 1988). Humpback whales have been observed along the entire east coast of Australia (10-38°S). The shortest distance by sea between these points while keeping within, say, 50 km of shore is in excess of 3500 km. Humpback whales have also been observed over a large range of latitudes off the west coast (Bannister, 1985; Dawbin and Gill, this memoir). In this study, songs have HUMPBACK WHALE SONGS been recorded off the east coast at a number of locations between Jervis Bay (35°20'S) and Cowley Beach, near Innisfail (17°40°S). The shortest distance by sea between these twa loca- lions is about 2400 km. Off the west coast, recordings have been made between Perth (32°S) and the Monte Bello Islands (19°40°S), Beeause the sang changes with time, com- parisons between recordings from different loca- tions must be confined to recordings that could be considered to be contemporaneous, i.e. separated in time by no more than a few weeks, if any observed differences are to be associated with the differences in position. Recordings off the east coast (Fig.!) were contemporaneous with recordings off Stradbroke ts., with the ex- ception of the 1979 and 198U recordings off Jervis Bay. In these comparisons, no greater differences could be detected between contem- porancous songs. at different locations than be- tween different songs at the same location. The largest scparations in these comparisons were Cowley Beach to Stradbroke Is, (1400 km) and Stradbroke Is. to Jervis Bay (970 km), The first recordings of Humpback Whale songs off the west coast were obtained by a naval vessel in October 1983 off Perth and the Monte Bello Islands, a separation of 1600 km by shortest path. The recordings were separated by only 13 days and comprised one song cycle al the northern Incation and little more than half the cycle in the southern recording. The part of the song that is common ta both recordings is similar, On the other hand these recordings are distinctively different from any recordings off ihe east coast, including those recorded within a few days off Stradbroke Is, The difference is fundamental: the sound types recorded off the west coast were different in their acoustical char- acteristics to those off the east coast. Recordings in Jater years have also shown the difference between the two coasts, for example, the songs recorded near the Monte Bello Islands in 1988 (several song cycles, two singers), and those discussed by Dawbin and Eyre (this memoir). Although the sample size is small, these results suggest Ihat the Humpback Whale song retains its integrity without geographical variation over a wide range of latitudes along the migration paths of the east coast of Australia. Humphack whale populations off the cast and west coasts of Australia are considered to be separate breeding stocks, with some intermin- gling in the Antarctic feeding grounds (Chil- tleborough, 1965; Dawhin, 1966). Evidence 287 from marks recaptured from whales during whaling activities from 1950 to 1962 (Dawbin, 1966) indicates that outside the feeding grounds, the two. stocks are well segregated but that a small amount of interchange does sometimes ocevr, Of whales marked off the east coast, 46 were later captured off the east coast, two off the west coast, Evidence from sightings, catches and catch compositions also indicates that stocks were well segregated, but Chittleborough takes the absence of any statistical differences in mor- phology as an indication of a small interchange between stocks. Humpback whale songs in the North Atlantic Ocean differ subsianuially from those of the North Pacific Ocean (Payne, 1978 Winn and Winn, 1978; Winn et al., 1981), Geographical barriers da not allow the opportunity for inter- change between stocks in these oceans as there is south of Australia, Qn the other hand, similar songs have been observed between breeding grounds which are widely separated by open water at similar latitudes in the same ocean, even (hough the separation 1s greater than between the cast and west coasts of Australia, The similarities of songs between Hawani and the Revillagigedo Is., (west of Mexico) in the North Pacific Ocean and between the West Indies and Cape Verde in the North Atlantic Ocean haye been reported by Payne and Guinee (1983) and Winn etal, (1981), the separation distances being 5000 km and 4300 km respectively, Using tail fluke photographs to identify individuals, Darling and Jurasz (1983) found one example of interchange between Hawaii and Revillagigedo Is. These results sug- gest, therefore, thal 4 geographical barrier be- tween breeding grounds or between breeding grounds and a substantial part of the migration route is more important than the separation dis- tance in determining different songs. It might be expected that the geographical separation would need to extend sufficiently far to separate that part of the migration paths over whith sig- nificant singing occurs. However, & more definite conclusion requires comparisons from other areas, ¢.g. between east Australia and New Zealand. TENTATIVE SONG RULES 1982-1985 To favilitate comparisons between hemi- spheres it is useful to formulate some tentative “rules” which govern the song pattern and sound character off Australia, These “rules” are designed to match as closely zs possible the 288 “rules” of song siructure already determined for the Northern Hemisphere and given in the papers. cited above. They are thus not necessarily new findings but rather an expression in a way designed to simplify comparison. Song pattern” is defined as the order in which sound units of the different sound types occur. ‘Sound charac- ter” is defined as the spectral and temporal char- acteristics of a particular sound ee The definition of “sound type” is given in (he section on “Methods” From the analysis the following features of the song have been established :- (a) The song consists of several themes (six in 1982/83) in a hxed order, each comprising a variable number af renditions of a phrase which is Specific to that theme (b) Each phrase consists of several discrete sounds (syllables or units) in @ fixed order separated by short periods of silence (typically 0).5-4secs). (c) Each sound unit is a sample of one or other of a small number of basic sound types (12 in 1982/83) appropriate to thal particular song. These are distinguished by their temporal and spectral characteristics (referred to as “character” in this paper). (a) The song paliern and the sound character at uny time appear to be maintained by in- dividuals of a stock for great distances along the migration paths and breeding grounds, although the sample size is small. (e) The songs are distinctively different be- tween stocks on the east and west coasts. (f) Both song pattem and character of the sound types change progressively with time, (h) Songs heard simultaneously appear to be independent and unsynchronised. DISCUSSION AND CONCLUSIONS Humpback whale songs recorded in Australian waters show a structure that is basically similar to that of the Northern Hemisphere songs. Al- though a song may contain some hundreds of sound units, all belong to one or other of a small number of sound types (12 in the 1982/83 song off east Australia). The song changes with time at anon uniform rate, Off east Australia the rate of change was gradual in 1982 and 1983, but a substantial change during the singing season in 1984 led to a substantially differentsong in 1985. This song was unusual in being poorly structured compared to the earlier songs. There are more subtle patterns in the song than have been addressed in this paper. There are also MEMOIRS OF THE QUEENSLAND MUSEUM small variations in pattern and in the charac- teristics of the sound types that warrant more detailed analysis. Information theory indicates that a signal can carry information only to the extent that itis unpredictable. Thus ina generally stereotyped song, il ts the variation from Lhe stereotype that has the potential to carry infor- mation, Songs are distinctively different between the east and west coasts of Australia. It seems [hat a geographical barrier between stocks where most of the singing occurs, i.e. on the migration paths and breeding grounds may be enough to ensure. that songs ate unrelated, even though the stocks may intermingle in the feeding grounds. In the samples available, the song appears to maintain its integrity over the very wide range of latitudes over which singing occurs off the cast coast. This leads ta a number of interesting ques- tions. How are changes in the song communi- cated along the coast, and at what rate? Do the changes originate at any point where singing occlifs or are there preferential source regions? At what distance from a singer does the seng affect the behaviour of other whales? The Australian coastlines provide access to Humpback Whales for thousands of kilometres along the migration paths and into the breeding grounds. The breeding grounds appear to be diverse and there may be no clear demarcation between these and the migration paths. Although whales appear in Hervey Bay as they migrate south, their behaviour shows greater proportion of activity of the type associated with Lhe breed- ing grounds than is observed further south off Stradbroke [s. These conditions provide the op- portunity to study the relative significance of the song in breeding and migration. Does the propor- tion of whales singing and related behavioural reactions vary along the coastline? The popula- tion of Humpback Whales off the cast coast has more than doubled since the early recordings were made (Paterson and Paterson,!1989) and there are now more peaple making recordings, so that we now have prospects for obtaining much larger data samples than were available far the analysis on which this paper was based. There are good prospects for progress in under- standing the significance of the song in migta- lion and breeding. ACKNOWLEDGEMENTS tam particularly indebted to Dr Robert and Patricia Paterson for their continued support and HUMPBACK WHALE SONGS encouragement throughout the study, They us- sisted in many ways, in particular in providing information on whale movements during the recording sessions. This was part of their general study of Humpback Whale migration. I um also indebted to Dr. John Quayle for providing the boat and his participation in the recordings off Stradbroke Island in 198] and 1982. to Les Nash for providing the boat and technical assistance from }983. 1 gratefully acknowledge the many discussions. with Dr W.H, Dawbin, Australian Museum, who has also been recording Humpback Whales in Australian waters, | am indebted to the following people for providing recordings of Humpback Whale sounds in Australian waters:- Mick Millington and Dick Sneesby (Department of Defence), near Jervis Bay 1979, 1980, 1984; Alastair Trail, near Heron Is, 1983; Royal Australian Navy, off West Australia, 1983; Frank Marian, Cowley Beach, 1987;Rob McCauley (James Cook University), near Monte Bello Is. (W.A.), 1988,1990; Ross Issacs, Hervey Bay, 1989, John Hayes, Hervey Bay, 1989, [am also grateful to Professor Michael Bryden and Dr Peter Corkeron for inviting me to work with them in Hervey Bay. 1 would also like to thank Geof Plant of the National Acoustics Laboratory for_allowing me to use their sonagraph, and Brian Jones of DSTO for the many times that he repaired, modified or built euuipment for the sound recordings. LITERATURE CITED BANNISTER, JL. 1985: Southern Right (Eubaleena australis) and Humpback (Megaptera navacangliae) whales off Western Australia: Some recent work. 105-114. In J.R- Ling and M.M. Bryden (eds) ‘Studies of sea mammals in southern latitudes. (South Australian Museum: Adelaide) BRYDEN, M.M., CORKERON, PJ. AND SLADE, R.W, 1988, Humpback whules in Hervey Bay.. Queensland, 1988, Aust. Nat. Parks Wildl. Serv., Canberra, Res. Rep. vi + 42 pp. CATO, D.H. 1978. Review of ambient noise in the ocean: nun biological saurces, Bull. Aust. Acoust, Sog, 6: 31-36. CATO, D.H. 1984, Recording Humpback While sounds off Stradbroke Island. 285-290. In RJ, Coleman, J. Covacevich and P. Davie, (eds) ‘Focus on Stradbroke’. (Boolarong Publica- lions; Brisbane). CHITTLEBOROUGH, RG, 1965. Dynamics of two populations of the Humpback Whale, Megay- tera novaeangliae (Borowski), Aust. J, Mar. Freshw. Res. 16: 33-128. DARLING, J.D. AND JURASZ, CM, 1983 Migralory destinations of North Pacilic Humpback Whales (Megaptera navaeangliae} 59-80. In R. Payne (ed.), ‘Communication and behaviour of whales’. (Westview:Colorado), DAWBIN, W.H. 1966, The seasonal migratory cycle of the Humpback Whale. 145-170. In K.S, Nor> ris (ed.) ‘Whales, dolphins und porpoises’. (University of California Press:Berkeley). TDAWBIN, W.H. & EYRE, E-J. this memoir. Humpback Whale songs along the ‘coast of Western Australia and same comparison wilh cast coast songs. Mem, Od Mus, 30(2);249- 254. KIBBLEWHITE, A.C., DENHAM, R.N. AND BAR- NES, D.J. 1967, Unusual low-frequency signals observed in New Zealand waters, J. Acoust. Soc, Am. 41: 644-655. MEDNIS, A, this memoir. An acoustic analysis of the 1988 song of the Humpback Whale, Meguplera novaeangliae, off eas! Australia, Mem, Od Mus. 30(2): 323-332. PATERSON, R. AND PATERSON, P. 1984. A study of the past and present status of Humpback Whales in east Australian waters, Biol, Consery. 29: 321-343. PATERSON. R. AND PATERSON, P. 1989. The status of the recovering stock of Humpback Whales Megaptera novaeangliae in east Austrahan walers. Biol. Conserv, 47: 33-48. PAYNE, R.S. AND McVAY, S. 1971. Songs of Humpback Whales. Science 173: 585-597, PAYNE, R.s, 1978, Behaviour and vocalisations of Humpback Whales (Megaptera sp.). 56-77. lo K.S. Norris and R, Reeves (eds), Report on a workshop on problems related to Humpback Whales (Megaptera novaeangliae) in Hawaii. (U.S. Marine Mammal Commission: Washington D.C.). PAYNE, R.S. AND GUINEE, L.N, 1983, Humpback whale (Megaptera novaeangliae) songs as an indicator of “stocks”. 333-358, In R.S, Payne (ed.) ‘Communication and behaviourof whales’, (Westview: Colorada). PAYNE, K. AND PAYNE, R.S. 1985. Large scale changes over 19 years in songs of Humpback Whales in Bermuda. Z. Tierpsychol, (J. Carn- parative Ethology) 68: 89-114. SIMMONS, M.L. AND MARSH, H, 1986. Sightings of Humpback Whales in Great Barrier Reel waters. Sci. Rep, Whales Res. Inst. 37; 31-48, THOMPSON, P.O., CUMMINGS, W.C. AND HA_ 290 S.J. 1986. Sounds, source levels and associated behaviour of Humpback Whales, southeast Alaska. J. Acoust. Soc. Am. 80: 735-740. TYACK, P. 1981. Interactions between singing Hawaiian Humpback Whales and conspecifics nearby. Behav. Ecol. Sociobiol. 8: 105-116. TYACK, P. 1983. Differential response of Humpback Whales, Megaptera novaeangliae, to playback of song or social sounds. Behav. Ecol. Sociobiol. 13: 49-55. WINN, H.E., PERKINS, P.J. AND POULTER, T.C. 1971. Sounds of the Humpback Whale. Proc. 7th MEMOIRS OF THE QUEENSLAND MUSEUM Annual Conf. on Biol. Sonar. 7: 39-42. (Stan- ford Research Institute:Menlo Park). WINN, H.E. AND WINN, L.K. 1978. The song of the Humpback Whale Megaptera novaeangliae in the West Indies. Marine Biology. 47: 97-114. WINN, H.E., THOMPSON, T.J.. CUMMINGS, W.C., HAIN, J.. HUDNALL, J., HAYS, H. AND STEINER, W.W., 1981. Song of the Humpback Whale - population comparisons. Behav. Ecol. Sociobiol. 8: 41-46. MEMOIRS OF THE (QUEENSLAND MUSEUM BRISBANE © Queensland Museum PO Box 3300, South Brisbane 4101, Australia Phone 06 7 3840 7555 Fax 06 7 3846 1226 Email qmlib@qm.qld.gov.au Website www.qm.qld.gov.au National Library of Australia card number ISSN 0079-8835 NOTE Papers published in this volume and in all previous volumes of the Afemoirs of the Queensland Museum maybe teproduced for scientific research, individual study or other educational purposes. Properly acknowledged quotations may be made but queries regarding the republication of any papers should be addressed to the Editor in Chief. Copies of the journal can be purchased from the Queensland Museum Shop. A Guide to Authors is displayed at the Queensland Museum web site A Queensland Government Project Typeset at the Queensland Museum HUMPBACK WHALES MEGAPTERA NOVAEANGLIAE IN THE GORGONA ISLAND, COLOMBIAN PACIFIC BREEDING WATERS: POPULATION AND POD CHARACTERISTICS L. FLOREZ-GONZALEZ Florez-Gonzdlez, L. 1991 07 01; Humpback Whales Megaptera novaeangliae in the Gorgona Island, Colombian Pacific breeding waters: population and pod chatacteristics. Memoirs of the Queensland Museum 3((2):291-295. Brisbane. ISSN 0079-8835, Photu-identification was used to study the Humpback Whale population which arrived at the Parque Nacional Natural Isla Gorgona between 1986 and 1988, Fifty animals were identified in 1986, 40 in 1987 and 35 in 1988 for a total of 108. Eleven individuals were common to 1986/1987 and 6 to 1987/1988. Three individuals were common for the three years. Using Petersen's formula, the estimates of the population for the 3 years ranged Irom 170 to 450, Gorgona has suitable environmental conditions required for humpback reproduction, The importance of Gorgona as a calving area is shown by the fact that 26.5% of animalsseen were calves, The groups that staved the longest time in the area were females accompanied by their calves but in general the frequencies were low. Humpbacks visit the region from June to November, the breeding season for Southern Hemisphere populations. During the years of study, behaviour of individuals and groups was analysed. L. Flérez-Gonzdlez, Universidud Nacional de Colombia = Fundacion Yubarta A.A, 33141 Cali (Valle), Colombia; 10 January, 1997, This is the first biological contribution on humpbacks in Colombia. Due to its geographic position, Colombia may potentially be a meeting place for Northern and Southern Hemisphere populations; therefore, this study may help to identify part of the migratory routes of both populations and may contribute to the protection and management of the species. Photo-identification (Katona et al.,1979) per- mits identification of individuals based on pig- mentation patterns, natural markings and scars; it has been used by various authors in different areas, and represents a useful tool for the estima- tion of populations, distribution, migration pat- terns and social dynamics (Herman et al., 1979; Whitehead, 1981; Whitehead, 1983; Balcomb and Nichols, 1982; Darling and MacSweeney, 1985; Mayo et al,, 1985; Baker et al., 1987; Clapham and Mayo, 1987; Kaufman et al,, 1987; Chu and Nieukirk, 1988). Humpback whale research in the Southern Hemisphere in America is limited. There are some studies that discuss the presence of this species (Alberica, 1986)as well as the lack of conservation and management policies (Ag- uayo-Lobo and Torres,1967; Paiva and Gran- geiro, 1970; Pinedo and Castelo, 1980; Cardenas etal., 1986; Guerra-Correaetal,, 1987),Only one sludy similar to that reported hercin has pre- viously been carried out in South America; that study, in 1988, at Abrolhos Bank, an important breeding site off northeastern Brazil employed photo-identification and identified 64 in- dividuals (Siciliano et al,, 1990), In Area | (feeding site) of the Antarctic, Stone and Hamner (1988) identified 32 individuals in 1985-1986. With the aid of a photographic catalogue, a Humpback Whale has been iden- tified at both the Antarctic Peninsula and Gor- gona Island areas, This is the first time that a Humpback Whale has been shown to cross the equator, and the first time an Antarctic Hump- back Whale has been documented in South American waters (Stone, Flérez-Gonzalez and Katona, 1990). STUDY AREA The research was carried out in the Parque Nacional Natural Isla Gorgona, an area of ap- proximately 49200 hectares, including Gorgona and Gorgonilla Islands (Fig.1) and established in June 1985. It is on the Pacific continental shelf of Colombja (2°47'-3°06’N and 78°06'— 78°18°W), 56 km from Guapi the nearest main- land town. Gorgona is approximately 10x3 km; its maximum height is 330m, The study area is located within the Intertropical Convergence Zone and thus endures variable winds and high pluviosity (7000-8000 mm annually). Salinity 292 COLOMBIA FIG. 1. Location map of study area. The broken line designates the Parque Nacional Natural Isla Gorgona limits. The dotted lines indicate isobaths in metres. The dot on the inset map indicates Gorgona Island in the Colombian Pacific. around Gorgona is 31-33 parts per thousand. Water temperature is 26-28°C (INDERENA, 1986; Llinas et al.,1990). METHODS Monitoring was carried out daily between 10 am and 6 pm. The surveys were conducted from a fibreglass taxi boat with a 25 horsepower out- board motor. On some occasions, the animals were followed 15-20 km from the island, but normally no more than 5 km. When conditions prevented use of a boat observations were made from the high bay coast guard tower. Song recordings were made with a Sony, TC- DSM, Tape recorder and a hydrophone with a frequency response 50—10000 Hz. Analysis of the song recordings have not been included in the present paper, however this field information permitted location and identification of in- dividual singers. RESULTS AND DISCUSSION MEMOIRS OF THE QUEENSLAND MUSEUM Three years of observations in the area per- mitted the photo-identification of 108 in- dividuals. Fifty animals were identified in 1986, 40 in 1987 and 35 in 1988; 17 individuals were sighted in more than one year: 11 were common to 1986-1987 and 6 were common to 1987-1988, 3 individuals were common for the three years . The 17 common individuals were: 3 females, 2 males, 1 calf, 4 adults, 1 immature and 6 non discriminated. Using Petersen’s formula (Begon, 1979), the population for the three years was estimated at 170-450. Of the 108 individuals, 42 (39%) were iden- tified by their back and the ventral surface of the flukes, 44 (41%) were identified exclusively by the ventral surface of the flukes and 22 (20%) using only the back. In 30 individuals, other characters (pectorals, dorsal region of the flukes, ventral region of the body and the head) were employed in identification. Five (5) fluke and six (6) back photographs show a thin layer of diatom growth. Five (5) underside tail region photographs show killer whale teeth scars. As previously established, Humpback Whales, during breeding activities, prefer meeting in groups near continents and around islands, in shallow waters, with protected beaches and temperatures 24—28°C. The eastern and extreme southern regions of Gorgona present these fac- tors and this makes the island an ideal breeding site. Here 26.5% of observed animals were cal- ves. INDIVIDUAL ACTIVITIES Apparently, irrespective of age or sex, whales were observed in the following activities: spin- ning breach, full breach, chin breach, tail breach, lobtail, flippering, rolling, spyhop, and drifting. These activities have been described in hump- back populations in different regions (White- head, 1981,1985a,b; Mayo et al., 1985; Pittman and Danton, 1985; Kaufman and Forestell, 1986; Silber, 1986). Group CHARACTERISATION The Humpback Whale population in Gorgona Island was observed in 5 classes of groupings: 1, solitary animal; 2, female and her calf; 3, two animals without calf; 4, three individuals usually a female with her calf and an escort; 5, group of more than three individuals. Of 35 solitary animals the 14 found in 1986 included 7 male singers; the 17 seen in 1987 included 13 male singers, in 1988 4 were seen HUMPBACK WHALES, GORGONA ISLAND, COLOMBIAN PACIFIC but it was impossible to determine if they were male singers, Of the 35 solitary individuals, 7 were heard from the boat without any electronic aid. The solitary animals generally presented an clusive behaviour making immersions of 15-20 sahnateh and showing little activity on the sur- ace, Groups including a female and calf were fre- quently found near the island, especially on the east, or protected side. For all observed females, epimeletic behaviour was evident, Three females and their calves were found on two occasions in a period of 8 days. One female and her calf were observed in two consecutive years (1987-1988). In the three years, 13 groups of two individuals, none calves, Were observed, Most were adults but it was not possible to determine the sex; they showed little activity on the surface. Mobley and Herman (1985) found that (he most stable social unit for groups without a calf was thal of adult pairs. However, as in this study, it was not possible to determine the sex of the pairs. Tyack and Whitchead (1983), suggested that the pairs are formed by an adult female and an adult male, Six trios had a female, her calf and an escort. Only two escorts were certainly males; for the remaining il was impossible to determine sex, Two triples were observed twice in the same year in the space of 2 and 8 days. respectively. Similar results have been described by Baker and Herman (1984a) who cited observation of a trio on two consecutive days. However, this does nol necessarily imply thai the trio stayed (together during these days, All groups with escorts were wivelling, and courtship behaviour was never seen. On all ocassions the small animal was swimming on top or beneath the female and the escort was generally protecting the group, lacat- ing itself between the ship and the couple, and sometimes using distractive tactics. Herman and Antinoja (1977) suggested that the escort whale could have a protective func- tion, However, Tyack and Whitehead (1983), proposed that the escorts are probably males waiting for the opportunity to mate if the female is receptive. This hypothesis is now accepted and jt has been supported by other researchers (Baker and Herman, 1984a; Mobley and Herman, 1985). On the other hand, Clapham and Mayo (1987) determined the sex of 91 escorts oul of 138 observed, and found that 55 (60.5%) were females and 36 (39.5%) males. This grouping pattern needs further interpretation and to that end determination of the sex of the individuals will probably be useful. 293 On September 6, 1986 a trio of a female and two males were observed in aggressive be- haviour for nearly 15 minutes; one individual left and the other two showed excitement on the surface for more than 10 minutes; they even jumped with their bodies ventrally joined for several seconds, Because it was mostly under- water, the specific function of this activity was not clear, Sixteen groups of more than 3 individuals were seen in the three years. The maximum number was nine. Most, showed agonistic behaviour (collisions, persecutions, excitement, loud noises, bubble expulsion and other aggressive activities) and constant coming together and splitting off of individuals. Agonistic and union-disunion behaviour, like those described here, have been reported by in- vestigators in other breeding sites (Herman and Antinoja, 1977; Tyack and Whitehead, 1983; Baker and Herman, 1984a, b; Mobley and Her- man, 1985; Silber, 1986). It seems that a familiar bond may exist among some individuals because they were secn together on more (han one uccasion in the same year and even in two different years. Unfor- junately, it was not possible jo determine sex and age. Traditionally, if has been assumed that Humpback Whales formed stable associations: family groups or reproductive couples (Whitehead, 1983). However, recent investiga- tions show that this is not completely truc. In the Antilles and in Hawaii, where humpbacks reproduce in the winter, groups stay together for a short period of time, excluding those formed by a calf (up to 1 year old) and her mother (Whitehead, 1981; Tyack and Whitehead, 1983; Baker and Herman, 19844; Mobley and Herman, 1985). Further observations are needed to con- firm whether the species forms stable family groups and whether it is monogamous or polygamous RECOMMENDATIONS A major obstacle to understanding the soctal behaviour of humpbacks has been the difficulty of determining the age and sex of the individuals. With long term studies, based on photo-iden- lfication and underwater observations to deter- mine s¢x, the knowledge of this species can be expanded making management and protection policies optimal. Therefore, investigations in the area of jhe park must be continued. 29d With (he objective of elucidating the migratory routes of the species in the Colombian Pacific. investigations in other areas north of Gorgona must be carried out (Negritos Island, Palma Is- land, Malaga Bay, Utria Inlet, Cupica Gulf), It has been recommended to the governmental management institute (INDERENA), to strictly control access of ships, boats and tourism to the Island, to avoid a threat to the Humpback Whale population. Some sites in the Colombian Pacifie where Humpback Whales have been reported are proposed to be managed as whale sanctuaries to ensure preservation through better management of this species which is seriously threatened. ACKNOWLEDGEMENTS This paper forms part of my muster thesis tn Inarine biology at the Universidad Nacional de Colombia. This research would not have been possible without funds from World Wildlife Fund in 1986, from (he Long Term Research Institute in 1987 and from Lylia Gonzalez (my mother) in 1988, Especially, | want to thank Dr Roger Payne, to whom the initiation of this re- search is attributed; for his advice in the study of the humpbacks and his help in finding funds. 1 wish to thank the Instituto Nacional de los Recursos Naturales Renovables (INDERENA), The Museo del Mar of the Universidad Jorge Tadeo Lozano, the Instituto de Investigaciones Marinas de Punta de Betin (INVEMAR) and the Universidad Nacional, for their logistic support. [ thank Wilfredo Henao, Luis Fernando Con- stain, Gustavo Bravo, Thomas Lyrholm for their help in the field work. I give special thanks to Carlos Patifo, Kathleen O'Connell, Roberto Tovar, Elvira Alvarado, Walter Mankel, Sven Zea, Arturo Acero, Fernando Patifio and Orlando Rodriguez who constantly encouraged and sup- ported my research. | want lo express my gratitude to my family for their support in the realization of all my prajecis and lo marine biologists Elvira Alvarado, Sven Zea and Adriana Beltran, for translation and corrections of this manuscript. REFERENCES ALBERICO, M. L986. Los Mamiferao. 191-210. In Prahl, HV. and Alberico, M,, (eds), ‘Isla de Gorgona’, (Biblioteca Banco Popular: Bogota) 2352p. AGUAYO-LOBO, A. AND TORRES, D. 1967, Ob- MEMOIRS OF THE QUEENSLAND MUSEUM servaciones sobre mamiferos marinos durante la Vigésima comision Antarctica chilena. Rev, Biol, Mar, |3(1): 1-58, BAKER,.C.S. AND HERMAN, L.M. 1984. Aggres- sive behaviour between Humpback Whales (Megaptera novaeangliae) wintering in Hawaiian walers. Can.J.Zool. 62(10);1922— 1937, BAKER, C,S. AND HERMAN, L.M. J984b. Seasonal contrast in the social hehaviour of the Humpback Whale. Cetus 5(2): [4-16. BAKER, C.S.. PERRY, A. AND HERMAN, L.M. 1987. Reproductive histories of female humpback whales Megaptera novaeaneliae iy the north Pacific. Mar, Ecol,Prog,Ser, 41; 103- 114, BALCOMB, K,C, If AND NICHOLS, G, 1982, Humpback whale censuses in the West Indies. Rep, Int, Whal. Commn 32: 401-406, BEGON, M. 1979. ‘Investigating unimal abundance: capture-recapture for biologists’ (University _ Park Press: Baltimore.) CARDENAS, J.C,, OPORTO, J, AND STUTZIN, M, 1986, Problemas de manejo que afectan a las pobluciones de celiceos en Chile. Proposiciones pura una politica de conservacion y manejo, Amb.y,Des, 2(3); 107-116. CHU, K.C. AND NIEUKIRK, S. 1988, Dorsal fin scars us indicator of age, sex, and social status in Humpback Whales (Megaprera novaeangliae), Can,J. Zool, 66; 416-420, CLAPHAM, P.J. AND MAYO, C.A. 1987. Reproduction and recruitment of individually identified Humpback Whales, Megaptera novaeangliae observed in Massachusetts Bay, 1979-1985. Can.J.Zool. 65(11): 2853-2863. DARLING, J.D. AND McSWEENEY, D, 1985, Ob- servalions om the migrations of north Pacific Humpback Whales (Megaptera novacangliae), Can.J,Zool, 63(2); 308-314, GUERRA-CORREA, C., WAEREBEEK, K. van, PORTFLITT-KANDORA, G. AND LUNA- JORQUEVA, G. 1987. Presencia de cetaceos frente a la segunda region de Chile, Estud. Oceanal, 6: 87-96. HERMAN, L.M. AND ANTINOJA, R.C. 1977. Humpback whales in the Hawaiian breeding water: Population and pods characteristics. Sci,Rep, Whales Res.Inst. 29: 59-85. HERMAN, L.M., FORESTELL, PH. AND AN- TINOJA, R.C., 1979. The 1976/1977 migration of Humpback Whales into Hawaiian waters: composile description, Mar,Mam.Com, Rep, MMC-77/19. Washington, D.C. 55p. HUMPBACK WHALES, GORGONA ISLAND, COLOMBIAN PACIFIC INDERENA. 1986. Colombia. Parques Nacionales, 2a ed. FEN edils. Bogota. 263p. KATONA, S.K., BAXTER, B., BRAZIER, O., KRAUS, S., PERKINGS, J. AND WHITEHEAD, H. 1979. Identifications of Humpback Whales by fluke photographs. 33- 44. 1n Winn, H.E. & Olla, B.L. (eds), “Behaviour of marine animals - current perspectives in resarch. v.3, Cetaceans,” (Plenum Press, New York). KAUFMAN, G.D. AND FORESTELL, P.H. 1986. ‘Hawaii's Humpback Whales’. (Pacific Founda- tion Press; Maui), 176p, KAUFMAN, G.D., SMULTEA, M.A. AND FORES- TELL, P.A. 1987. Use of lateral body pigmen- tation patlerns for photographic identification of east Australian (Area V) humpback whales. Cetus 7(1): 5-13, - LLINAS, R.R., PINTO, J.N., PENA, F. AND CARO, F. 1990. Geologia. 55-64. In Aguirre, J, and Rangel, O. (eds), ‘Biota y ecosistemas de Gor- gona. (Féndo PEN—Colombia, Editorial Presen- cia Ltda; Bogota). 303p. MAYO, C., CARLSON, C., CLAPHAM, P.J. AND MATTILA, D.K. 1985. ‘Humpback whales of the southern Gulf of Maine’. (Center for Coastal Studies; Provincetown), 62p. MOBLEY, J.R. Jr. AND HERMAN, L,M, 1985, Transcience of social affiliations among Humpback Whales (Megaptera novaeangliae) on the Hawaiian wintering grounds, Can,J,Zool. 63; 762-72. PAIVA, M.P, AND GRANGEIRO, B.F, 1970. Inves- tigations on the whaling seasons 1964-1967 off northeastern coast of Brazil. Arq, Cien. Mar 10(2): 111-126. PINEDO, M.C. AND CASTELO, H.P. 1980. Estudos de celéceos no oceano Atlantico sul occidental e a falta de uma politica de conseryacao e manejo no Brasil, Bolm Inst. Oceanogr., Sao Paulo 29(2): 319-21. PITTMAN, 8. AND DANTON, C. 1985. Humpback Whale behaviour. Apendix II. 48-53. In Mayo, C. et al, SICILIANO, S., LODI, L, SALES, G. AND GONCHOROSKY, L. 1990, Observacoes de baleias jubarte Megaptera novaeangliae, no Banco dos Abrolhos, Costa Nordeste do Brasil. 40. In 4a Reunidn de Trabajo de Especialistas en Mamiferos Acuaticos de América del Sur. 12- 15 de Nov. 1990. (CIMMA-Universidad Austral de Chile: Valdivia), 75p. SILBER, G. 1986, The relationship of social vocaliza- tions to surface behaviour and aggression in the Hawaiian Humpback Whale (Megaptera novaeangliae). Can.J.Zool. 64(1): 2075-2080. STONE, G.S. AND HAMNER, W.M. 1988. Humpback whales Megaprera novaeangliae and southern right whales Eubalaena australis in Gerlache Strait, Antarctica. Polar Recard 24(148): 15-20. STONE, G.S., FLOREZ-GONZALEZ, L. AND KATONA, 5. 1990, Whale migration record, Nature 346: 705. TYACK, P. AND WHITEHEAD, H. 1983. Male competition in large groups of wintering Humpback Whales. Behaviour 83; 132-154. WHITEHEAD, H.P. 1981. The behaviour and ecol- ogy of the Humpback Whale in the northwest Atlantic. PhD thesis, Univ. Cambridge.(un- publ.). WHITEHEAD, H.P, 1983. Structure and stability of Humpback Whale groups off Newfoundland. Can.J.Zool. 61(6): 1391-1401, WHITEHEAD, H. 1985a. Why whales leap. Scien- tific American 251(3): 84-93. WHITEHEAD, H. 1985b. Humpback whale breach- ing. 117-155, In Pilleri, G. (ed.), ‘Investigations on Cetacea, vol. 17. MEMOIRS OF THE (QUEENSLAND MUSEUM BRISBANE © Queensland Museum PO Box 3300, South Brisbane 4101, Australia Phone 06 7 3840 7555 Fax 06 7 3846 1226 Email qmlib@qm.qld.gov.au Website www.qm.qld.gov.au National Library of Australia card number ISSN 0079-8835 NOTE Papers published in this volume and in all previous volumes of the Afemoirs of the Queensland Museum maybe teproduced for scientific research, individual study or other educational purposes. Properly acknowledged quotations may be made but queries regarding the republication of any papers should be addressed to the Editor in Chief. Copies of the journal can be purchased from the Queensland Museum Shop. A Guide to Authors is displayed at the Queensland Museum web site A Queensland Government Project Typeset at the Queensland Museum LARGE-SCALE PLANNING FOR ASSESSMENT AND RECOVERY OF HUMPBACK WHALE POPULATIONS STEVEN K. KATONA Katona, 8,K, 1991] 07 01; Large-scale planning for assessment and recovery of humpback whale populations. Wemairy of the Queertsland Museum 3Q(2):297-305. Brisbane. ISSN OU79-8835, Planning to help humpback whale populations recover from past commercial hunting must acknowledge their seasonal diversity of habitats, frequently governed by different nations, where they are affected by human activities such as shipping, fishing. dredging, waste disposal. and others. Human-induced changes in ocean ecology or resource abundance make it unlikely that populations could regain aboriginal sizes. Comprehensive research programs to assess present distribution and population sizes will usually require coordinat- ed efforts by scientists [rom many nations, Estimation of present population size is facilitated by seasonal aggregation of whales at feeding or breeding locations, but poor esumation of prehunting abundance make it hard to choose a desired population size or to evaluate present vs past populations. Some populations may require many decades to reach desired population size, wwing lo the whales” tendency to return to traditional locations and their slow tate of reproduction, Monitoring programs must be long term. The U.S. Humpback Whale Recovery Plan is discussed as an example of a large scale plan covering 3 stocks of humpback whales. in two oceans. A research proposal, entitled Years of the North Atlantic Humpback (YONAH), to assess population sizes and movements by an intensive international synoptic study throughout the summer and winter range of the species is summarized. Steven K. Katona, College of the Atlantic, Bar Harbor, Maine, USA 04609; 20 December, 1990, The humpback whale, Megaptera novaeangliae, was the second species to be protected by the Intemational Whaling Commis- sion, beginning in 1955 with prohibition of commercial (non-subsistence) hunting in the North Atlantic Ocean. Protection was extended to the North Pacific and Southern Hemisphere populations after the 1963 hunting season. The species has not been hunted subsequently except for small subsistence. hunts in Greenland (up to {0 per year until 1980 [Kapel, 1979], last whale taken in 1987); Bequia Island, Lesser Antilles (several per year, none caught since 1987 [ Ward, 1987; Price. 1985; Adams, 1971, 1975]); and Tonga (up to 10 per year until hunting was stopped in 1978 [Winn and Reichley, 1985]. Winnand Reichley (1985) also noted that at least one humpback was taken by pirate whalers. Forty-one (41) humpbacks were taken in eastern Canada from 1969-1971 by scientific permit (Mitchell, 1973). Hunting depleted population sizes so greatly throughout their range that Humpback Whales was listed as. “Endangered” in waters of the U_S- (Braham, 1984) and “threatened” in Canada (Whitehead, 1987) and are included in Appen- dix I of the 1973 Convention on International Trade in Endangered Species (CITES). Classification of an animal as “endangered” entails substantial governmental and scientific responsibilities. In most such cases, active management is required by law, when possible, to facilitate recovery of the depleted popula- lion(s). Scientific studies may be required to define more exactly the species’ status, potential for recovery, and any actions to be taken, Sub- stantial costs in personne! time and money may be required. Since the main historical threat to humpback whales, hunting, effectively ended 25 years ago, it might be sufficient to do nothing and simply wait for the protected populations to increase of their own accord, This would be a tempting alternative for governments. beset with other acute problems. In the end. time alone might be the most important factor necessary for recovery. However, a suite of problems that did not exist in the days of hunting now affects some popula- tions of humpbacks and other whales. Many of these threats are much harder to manage than was hunting, and their combined effects could be equally detrimental. These threats are not inten- 298 tionally directed at humpback whales or other species. Some are unintentional side effects either of Jungstanding human uses of the sea, such us fishing or shipping, or sloppiness (e.g,, pollution by sanitary, chemical or solid wastes). Others involve potential competition between humans and whales for food resources, such as capelin, herring and lately krill, or space, for example where coastal, recreational or industrial development may impact locations used for feeding or breeding (e.g. Nitta and Naughton, (989). Some of these threats are subtle and difficult to quantify, Even where effects are dramatic, such as for entrapment or entanglement of whales in fishery gear (Lien etal., 1990) mitiga- thon may be very difficult if no satisfactory alter- native methods are available for humans. For example, prohibiting the use of shore-fast nets along the coast of Newfoundland would save the lives of several dozen humpback whales every year, but it would also ¢liminate the only source of income available to most coastal residents. The curnulative or synergistic effects of different threats, such as food limitation or contamination and challenge by pathogens (Geraci, 1989), are difficult to analyze. Successful management of the humphack is complicated by their seasonal migrations. During their annual cycle, the whales range through a large geographic urea, frequently in- cluding waters under the jurisdiction of several nations, Studies sufficiently broad ranging to describe topics such as migration, population structure, or abundance and trends benefit from or require the collaboration by many scientists, often in dilleremt countries, Since the whales have relatively long lifetinves, some studies (e.g. investigations on age-specific natalily, compurit- live reproductive fitness, or heritability of be- havioral or morphological features} may require decades, sometimes longer than the span of an individual scientific career. Successful incar- poration of research results into plans for effec- tive management of the species also laces geographic, temporal and political obstacles, For example, effective management of summer habitats to slow adequate opportunities for feed- ing is not sufficient jo preserve the species if Winter habitats for reproduction are not also maintained, Coordination between governments is necessary for management of most hampback whale stocks. Confounding the issue still further, the terms “management”. “conservation” or “pres- MEMOIRS OF THE QUEENSLAND MUSEUM ervation” are temporilly unbounded. They imply periodic assessment, evaluation and ac- tions to keep a species within certain numeric limits forever, Since natality and mortality vary over time in relation to prevailing ecological forces, management-relaled research must be carried out at appropriate intervals to determine population sizes and trends. This may become tricky if human-induced mortality causes total mortality to approach natality, In such a situa- tion, and especially in the presence of stochastic factors, it may not be possible at any one time to evaluate whether a population is headed for sur- vival or extinction, Gaskill and Lien (in prep,) emphasized that very long periods may be re- quired to assess the true “health” of a population of whales. Their computer models produced in- stances in which apparently “healthy” popula- tions went extinct over periods of hundreds of years, even while showing several 50-year periods of steady increase during whict scien- (isis or managers would have concluded that the species was progressing wonderfully. Cautioned by such awareness, management of the humpback whale or of any endangered species cannot be regarded as a task to complete, While there may be some actions that need to be accomplished only once, for example designat- ing certain locations as sanctuaries for the species, other actions may need to be overseen or repeated forever, or until a decision is laken that the specics should not be protected. The following accounts present highlights of two recent approaches to long term, broad scale managemen! or study of humpback whales, namely the U.S. Humpback Whale Recovery Plan and a new research proposal entitled YONAH (Years of the North Atlantic Humpback Whale), Communication of some of the goals, problems and progress of these projects may help them to succeed and may help athers to replicate and improve them. U.S. HUMPBACK WHALE RECOVERY PLAN The U.S Endangered Species Act (ESA) of 1973, (If USC 1531 et seg., amended 1978, 1982, 198h) requires Federal agencies 10 use all reasonable methods available to conserve species of plants and animals that are endangered or threalened with extinclion. The U.S. Marine Mammal Protection Act (MMPA) of 1972 (16 usc 1361] et seq., amended [990) assigned to the National Marine Fisheries Service (NMFS) PLANNING RECOVERY GF HUMPBACK WHALE POPULATIONS responsibility for managing most marine mam- mal populations, including endangered species, Section 4 (g) of the ESA requires the Ad- munistrator of an agency with oversight respon- sibility for an endangered species to determine whether populations of that species would benefit from a Recovery Plan, and if so, to con- struct and implementone. A Recovery Plan is “a guide that delineates, justifies and schedules those actions required for restoring und securing an endangered/threatened species as a viable, self-sustaining member of its ecosystem.” Guidelines for constructing Recovery Plans specify thal a numerical goal is usually required, along with means for evaluating progress toward achievement of that goal. In 1987 the Administrator of NMFS con- cluded that humpback whales would benefit from a Recovery Plan and assigned the NMFS Office of Protected Resources to construct it. An advisory group, the Humpback Whale Recovery Teain, Was sclected by NMFS to assist with the Plan (NMFS, 1990), The 10-person Team in- cluded experts on marine mammals from the private sector, academia, and government. Con- struction of the Plan, now in final stages of approval to become policy for NMFS, required nearly 2 years, 2 team mectings, 7 dralis, and 2 periods for public review and comment, The Plan concentrates primarily on popula- tions of humpback whales occurring seasonally or permanently in waters of the North Atlantic and North Pacific Oceans under U.S, jurisdic- tion, but it also includes recommendations for populations using waters around U.S. Trust Ter- nitories jn (he South Pacific. The Plan reviews the natural history of humpback whales; sum- murizes current knowledge about populations in the North Atlantic Ocean and (he North Pacific Ocean; discusses known and potential threats to the species; then details a series of actions recommended to achieve recovery goals and a prioritized schedule estimating the time and moncy required for (hase actions. Defining an overall goal for the Plan was nol simple. The objective contained in the recovery planning guidelines, namely restoring the species to be a functional member of its ecosys- tem could not easily be defined or evaluated, so it would not suffice. The goal of encouraging populations (o increase to the abundance prevail- ing before commercial hunting sounded appeal- ing, but reviewers correctly noted that it might not be possible or desirable. since many other vonditions had also changed since bunting 299 began. Some new equilibrium would have to be sought that reflected expanded human use and abuse of the sea, since loday’'s oceans could not support aboriginal numbers of whales as well as increased human populations The Team then tried (a detine a suitable per- centage of aboriginal numbers for which to strive, but this also proved to be premature. How could one choose such a percentage, or evaluate progress toward that goal, without confiden| knowledge of prehunting abundance. Despite meticulous historical research (e.g. Miichell and Reeves, 1983) the few existing estimates of early abundance for the population in the western North Atlantic Ocean Were ot precise enough ta tely upon, and even less mformaltion was available for other oceans. Consequently, the Plan now identifies several longterm goals: (1) the “BIQLOGICAL GOAL” of building and maintaining populations large enough to be resilient to chance events such as epizootics, episodic oceanographic changes, in- breeding or anthropogenic environmental catastrophes; (2) the “NUMERICAL GOAL” of achieving population sizes equal to al least 60% of jhe historical carrying rapacy for popula- tions in the North Atlantic and North Pacific Oceans, since large mammal populations are thought to achieve maximal productivity begin- ning at approximately that abundance (Fowler and Smith, 1981); and (3) the “POLITICAL GOAL" of being able to teclassify populations of this species fram “endangered” to “threatened” or even “unprotected”. While tasks identified in the Plan define those longterm goals more precisely, the Plan suggests adoption of an INTERIM GOAL to double the abundance of populations during the next 20 years, Periodic assessments of population abundance are called for in the Plan, Data showing statistically sig- nificant trends of population increase overall and reoccupation of portions of the range knowit jo have been occupied during historical times are specified as acceptable evidence of progress (owatd recovery goals, Recovery Planning Guidelines require that each action recommended be assigned a priority. Priority 1 denotes an action thal must be taken fn prevent extinction or to prevent the species from declining irreversibly in the foreseeable future. Priority 2 is an action that must be taken to prevent a significant decline in the species population or habitat quality, or some other sig- nificant negative impact short of extinction, Priority 3 is all other actions necessary to provide 400 for full recovery of the species. According to these guidelines, there is nothing that should (or could) now be done for humpback whales that could be classified as Priority 1. However, many things should de classified as Priority 2. Actions (lasks) were recommended in four general categories: 1) Maintain and enhance habitats used by humpback whales currently or historically; 2) Identify and reduce direct, human-related mortality, injury and disturbance; 3) Measure and monitor key population parameters; and 4) Improve administration and coordination of recovery efforts. Foremost among the recommendations for habitat maintainance ts entifying and designat- ing critical habitat, particularly in the Hawaiian Islands winter range, bul also if needed in other locations. Official designation as critical habital adds # layer of Federal protection to any area within the jurisdiction of the U.S, This jask will require better definition of the habitats and habitat features utilized by humpback whales. Other nations are encouraged to tuke similar actions to protect humpback whale habitats of special importance, as the Dominican Republic had already done by proclaiming Silver Bank a sanctuary for that species. Studies are recom- mended to ascertain historical abundance or the potential for reoccupation of habitals that may have been more important to the species in the past und that could be used if populations in- crease. The development of protocol for munitoring physical and chemical factors that could deerease habitat quality, as well as parasite load. biotoxins. and anthropogenic contaminant levels in tissucs of whales and their prey is called for. The Plan recommends reducing disturbance from human- produced underwater aoise in Hawaiian waters and in other important habitats whenever humpback whales are present. Finally, recognizing the necessity for providing adequate nutrition for whale populations, the Plan recom- mends monitoring levels oi prey abundance, identifying and evaluating the {ype and amount of fisheries competition, and preventing the in- itiation of new large scale fisheries for primary prey of humpback whales. Foremost among the recommendations designed to identify and reduce direct human-re- lated injury and mortality and perhaps casiest to achieve is continuing the ban on commercial hunting of humpback whales. With luck, this alone might allow recovery of the species throughout its range. However luck cannot be trusted in the face of {he various known and MEMOIRS OF THE QUEENSLAND MUSEUM potential impacts that have been identified, which include subsistence hunting, entrapment and entanglement in fishing gear, collisions with ships; disturbance from vessels and aircraft; un- derwater noise from ships, boats and aircraft, commercial whale-watching boats (Atkins and Swartz, 1988: Beach and Weinrich, 1989) and research boats; nvise fronr industrial activities; habitat degradation from chemical pollution, in- cluding petroleum: activities associated with in- creased coastal development; and competition for resources with humans. While some of thase impacts might become more severe in the future, the most currently pressing task appears to he working with commercial fishermen and tegional fisheries managemen| councils to modify fishing gear and/or fishing regulations, where possible, ta prevent entrapment or ene tanglement of humpbuck whales. Further evaluation of the effect of collisions with ships on humpback populations is also called for. Im- proved protocol for reporting and investigating carcasses, Stranded specimens, and for photographing living whales will all help to pro- vide information necded for these and other tasks. Important recommendations for measuring and monitoring key population parameters in- clude re-analyzing existing data with improved techniques; systemalizing sampling methods for eslimating population size; and developing bet- tet facilities for obtaining, archiving and analyz- ing data on humpback whales, Rescarch vessels need to be dedicated to studying humpback whales and other endangered species, by build- ing a new vessel for use in the North Pacifie and another for the North Atlantic Ocean, and/or by chartering existing vessels. New field studies on population dynamics are recommended to ex- amine age-specific rates. of birth, survivorship and mortality. Better definition of geographic subdivisions of populations are called for, both by analyzing and evaluating existing informa- tion and by implementing immediately snitial surveys of selected regions, Migrationroutes and transil (imes need to be defined more precisely by using long-term radio tags, underwater Jisten- ing stations and genetic techniques- Finally, bet- ter assessments of abundance, status and trends of humpback whale populations are called for (hrough new census surveys, participation in in- ternational sightings surveys and implementa- tion of improved sampling programs for caplure- recapture studies using individual identification photographs. PLANNING RECOVERY OF HUMPBACK WHALE POPULATIONS Of foremost importance to the objective of improving administration and coordination of the humpback whale recovery program will be selecting a Director for the overall recovery el- fort. Expansion or reconstitution of a Reenvery Implementation Team to include wider repre- sentation of agencies such as the Mineral Management Service and National Park Service. and preparation of detailed regional work plans will improve the Plan’s chance of success. An important recommendation is to en- courage multinational cooperation to protect humpback whale habitats. Discussing the Plan at international mectings; distributing it to other countries. and providing apprapriate follow-up communication; integrating recommendations with goals of the International Whaling Com- missions and encouraging other nations to develop recovery plans for conservation and management of humpback whales are important long term goals that will benefit the species. This sounds like a tall order, especially since NMPS, like most government agencies. is al- ready overburdened und underfunded. Some of ihe recommendations in this Plan might conflict with those in other plans, Forexample, maintain- ing large populations of sand lance. herring and capelin lor humpback whales could conflict with goals. of the Right Whale Recovery Plan if the fish competed with right whales by consuming too many copepods. NMFS ts also preparing a plan for managing shark populations, some of which are under heavy fishing pressure. Some of the sharks thus saved will probably eat young humpback or right whales. How all of this will work out remains to be seen. YONAH (YEARS OF THE NORTH ATLANTIC HUMPBACK) As demonstrated above, two simple questions need to be answered as a prelude to successful munagement of a species. Where are they? ancl How many are there’? Answers remain elusive for most species of cetaceans, Fortunately those questions are somewhat casicr to answer for humpback whales than for other balaenopterids, because the whales are easier to sec and because they gather at traditional locations during winter. Nevertheless, it has not yet been possible to estimate population abundances sufficiently precisely to allow trends to be detected with statistical conlidence, YONAH (Years of the North Atlantic Humpback Whale) isa research project designed 4A lu answer those questions by sampling all populations in the North Athentec Ocean with standardized methods during a two-year period using two techniques relatively new to whale science, photographic jdentification (Hammond et al., 1890) and DNA-biapsy sampling (Baker et al., 1990) of individual whales. Other unique features of YONAH include simultaneous sam pling of al] known habitats used during summer by humpback whales in the entire North Atlantic Ocean fortwo consecutive summers; and inten- sive sampling on. the winter range during the preceding and [ollawing winter seasons. Large sample sizes will provide substantially improved description of populatiun abundance, distribu- tion and interchange. for an accurate snapshot of the species ocean wide status, YONAH requires international cooperation, because most Humpback whales in the North Atlantic Ocean feed, migtate, overwinter and/or breed in waters under the jurisdiction of two or more countries. Furthermore, it would be dif- ficult for one governmenl to provide all the fund- ing necessary for such 2 comprehensive project, and it would be nevrly impassible for one re- search team to. accomplish the synoptic sampling program within the lime penod necessary to achieve the program's scicntific goals. YONAH will benetit from large scale models of collaboration that have already been estab- lished. For example, in 1987, Denmark, the Faroes, lceland, Norway and Spain collaborated in an extensive shipboard and acrial census of whales in the eastern North Atlantic thal used scientists from & nations (Oritsland ef al., 1989). A significant problem confronts workers using photo-identification or DNA-biopsy tech- niques to study long range migrations, popula- lion Subdivision and oceanic abundance. Hammond (1986,1990) described how oppor- tunistic methods used in many such studies may not sample all portions of a population equally owing lo geographic. temporal and biological biases. A brief examination of shortcomings in samples currenily available for regions of the North Atlantic Ocean és instructive. Geographic biases include the following ex- amples. Newfoundland’s Jong coastline is seasonally home to most of the whales. in the western North Atlantic, but no comprehensive photographic sampling has been carried out since Whitehead’s pioneering studies (c.g. Whitehead et al,, 1982; Whitehead and Glass, 1985). Even those studies visited only a small portion of the coast and spread the work 302 over several years. The situation in Iceland is even more difficult. The conclusions that whales from Iceland’s Denmark Strait form a separate feeding aggregation and visit the West Indies to breed (Katona and Beard, 1990) were drawn from a sample of only 20 whales. Luckily, 4 of them were subsequently sighted on the winter range and one returned to Iceland during a sub- sequent summer. At the other extreme, the Gulf of Maine, a relatively small region seasonally inhabited by somewhat fewer than 500 whales, each year produces thousands of photographs. However, most of those photographs are taken by naturalists or scientists working mutualisti- cally with commercial whale-watch boats, whose working range is usually constrained by the 4 to 5 hour length of scheduled trips. Conse- quently, relatively accessible locations are sampled disproportionately. Mitchell and Reeves’ (1983) early concern that photo-iden- tification studies have not fully determined the boundaries of the populations sampled is still true. Temporal biases also exist. For example, most photographs in the Gulf of Maine are obtained from April 15 to October 15, the season when commercial whale-watch boats operate. Whales that appear early, perhaps on the way to other feeding aggregations, or whales that stay late, perhaps remaining throughout winter, do not have an equal opportunity to be photographed, particularly if they occur offshore. Dispropor- tionate sampling during summer months is a common feature of studies throughout the North Atlantic. Larger scale temporal inequalities in sampling intensity also occur. For example, en- tire regions may be sampled during some years, but not others. A suite of interesting biological biases also exist, such that different age classes, reproduc- tive classes or genders may be over- or under- represented in samples. Resulting problems can be particularly vexing if sampling methods used in different regions cause different types of biases. For example, sampling on the winter range has focused on surface active courtship groups, which present more opportunities for photographing flukes than do mothers with cal- ves or single whales. Since such groups are predominantly male, photographs of that gender may be over-represented. On the other hand, singing whales, thought to be only males, remain submerged for up to 30 minutes, and are likely to be sampled less than other classes. Similarly, a female accompanied by a calf flukes up less MEMOIRS OF THE QUEENSLAND MUSEUM often than do other animals, particularly on the winter range. Calves photographed on the winter range are normally excluded from calculations of abundance, because they rarely fluke up and their fluke pigmentation patterns are not yet clearly formed. Awareness of these biases and recognition that international cooperation would be needed to eliminate them led to the idea for a carefully designed, intensive synoptic investigation of all humpback whale habitats in the North Atlantic Ocean. A research proposal was prepared by scientists from 6 nations (see Acknowledge- ments) and submitted to the International Whal- ing Commission (Document SC/40/025) and the Marine Mammal Committee of the International Council for Exploration of the Sea (ICES) (C.M 1990/N:18). Major goals of YONAH are:- 1, Photographs for individual identification will be obtained from approximately 1500 whales on the summer range during each of two successive summer seasons and 2000 whales on the winter range during the previous and inter- vening winters. Biopsy samples are anticipated from about half of those animals. Standardized protocols will be used to select areas in the summer range for searching, locate whales or groups of whales, select individual whales from groups for photographic or DNA sampling, and to terminate activity with a whale if samples have not been obtained within a reasonable amount of time. The overall sampling objectives will be to equalize the opportunity for every individual whale to be sampled, compensate for any inequalities in resulting analyses, and achieve precise confidence intervals by obtain- ing sufficiently large sample sizes. 2. Individual-identification photographs will be centrally archived and analyzed as described by Katona and Beard (1990), in order to be comparable with previous results. Procedures will be developed for providing appropriate ac- cess to photographic samples for YONAH col- laborators or other interested scientists. 3. Protocol for processing and archiving biop- sy samples will insure that at least half of each biopsy sample collected will be transferred to YONAH and used to determine gender of in- dividuals; the number of matrilineal lines (haplotypes) present in mitochondrial DNA in the North Atlantic Ocean and their distribution in different subregions; and to identify nuclear gene sequences that would permit Mendelian analyses for delineating panmictic population PLANNING RECOVERY OF HUMPBACK WHALE POPULATIONS unils, Additional studies for genetic fingerprint- ing and identification of paternal markers may be attempted after primary analyses are com- pleted, Tissue remaining after primary analyses. along with appropriate descriptive dala, will be preserved in a central archive at -S0° C 10 insure its long term viability and availability. The YONAH organizing committee will welcome proposals for further analyses of tissue samples and grani permission for their use us appropriate, in consultation with the Project Coordinators for YONAH subregions involved, 4. A comprehensive centralized database will be constructed containing all information on jn- dividual photo-identifications, individual genetic data, sightings from which photographic and/or biopsy samples were allempled, and sear- ching effort when locating groups of whales to sample. Data auditing checks will track and coordinate linkage of field and laboratory results so that no data are lost, YONAH collaborators will receive updated copies of the entire database periodically. 5, The YONAH database for the entire North Atlantic Ocean will be analyzed to estimate total population size and rates of exchange of whales belween sub regions of the summer range. The representativeness Of data collected at the regional level will be investigated and results used to modify methods for estimating popula- tion abundance, Likelihood models will be developed for estimating abundance and also for defining DNA haplotypes, Sensitivity analyses will also be conducted to determine the sen- sitivity of estimates of abundance 4nd jnter- change to various assumptions. 6. Population abundance Will be estimated by applying the Petersen capture-recapture method ta individual-identification photographs. The planned sampling design, two successive seasons on the summer range (S) and $2) along with the previous and intervening scasons on the winter range (W) and W2) will permit several independent population estimates using cither Wi/Su Si/W2; W2 /S2, Wi/W2 or Si/Sz as the capture-recapture samiples, Estimates will be prepared for population subregions, including the Gull of Maine, Gulf of St. Lawrence, New- foundland, southwestern Greenland, [celand, and Norway; and also for the whole North Atlan- tic Ocean . This is an ambitious project. However, scien- tific interest is high and. it dovetails nicely with some national and international goals of several countries. NMFS appears to be willing to support further planning for organization and develop- ment of scientific protocols. Proposals for aduli- tional funding are in preparation. Current hopes are that Work al sea might begin by January. 1992, ACKNOWLEDGEMENTS The following people served on the Humpback Whale Recovery Team: C. Scott Baker (University of Wellington, New Zealand); Howard W. Braham (National Marine Manimal Laboratory. NMFS, Seattle. Washington); Jofin J. Burs (Living Resources, Inc., Fairbanks, Alaska); Douglans G, Chapman (University of Washington, Seattle); Deborah Glockner-Ferrari {Center far Whale Stidies, Walnut Creek. Calitorma); Steven K. Katona (College of the Aduntie, Bar Harbor, Maine); James H. Lecky (NMFS, Terminal) Island, California); John H. Prescolt (New England Aquarium, Boston, Mas sachusctts); Gerald P. Scott (NMFS, Miami, Florida) and William A, Watkins (Woods Hole Oceanographic Institution, Woods Hole, Mas- sachusetts}, Charles A, Mayo, Ir, (Center for Coustal Studies, Provincetown, Massachusetts), Roger Payne (Long Term Rescarch Institute, Lincoln, Massachusetts) and Gloria Thompson {Office of Protected Resources, NMFS, Silver Spring, Maryland) served as Technical Advisors io the Team. Thanks are also due to Nancy Fostor, Director of the Office of Protected Resources, NMFS, and Charles Karnella, Assis- lant Director of that office, for assistance during all phases of Recovery Plan construction. | thank College of the Atlantic fur sabbatical leave and the New England Aquarium for office space, support services and housing subsidy during 6 months in Boston spent working on the Recavery Plan, The discussion of the U.S, Humpback Recovery Plan summarizes my own experience. My comments do not necessarily represent views of the Recovery Team, NMFS or any agency or party other than myself. The Plan is still being discussed and evaluated by NMFS and some of the recommendations mentioned could be modified ot eliminated before the Plan he- comes NMFS policy. Copies of the Humpback Whale Recovery Plan and the Recavery Plan for Northern Right Whales can be obtained by wril- ing to NMFS, Office of Protected Resources, 1335 East West Highway, Silver Spring, Maryland, USA 20910, The idea thal grew into YONAH was generated by David K. Mattila, Philip J. Clapham and Charles “Stormy” Mayo, Jr. of the Center for Coastal Studies, Provincetown, Mas- sachusetts, Thanks are duc to Tim Smith (North- east Fisheries Laboratory, NMFS, Woods Hole, Massachusetts) for taking on the task of coor- dinating the YONAH planning effort and help- ing to bring that important research project from idea to reality, Subsequent planning and prepara- tion of research proposals involved Nils Oien, (Norwegian Institute for Marine Research), Johann Sigurjonsson, (Iceland Fisheries Re- search Institute), Finn Larsen (Greenland Fisheries Research Institute), Per Polsboll (University of Copenhagen), Philip Hammond (Sea Mammals Research Unit, Cambridge University), Jon Lien (Memorial University of Newfoundland), David Mattila, Philip Clapham, Stormy Mayo (Provincetown Center for Coustal Studies), Tim Smith and Tom Polacheck (North- east Fisheries. Laboratory, NMFS), and Steven K. Katona and Judith A. Beard (College of the Atlantic). LITERATURE CITED ADAMS, J.E. 1971. Historical geography of whaling in Bequia Island, Wesi Indies. Caribbean Studies 11(3%55-74. ADAMS, LE. 1975, Primitive whaling in the West Indies. Sea Frontlers 21(5):303-313, ATKINS, N, ANDSWARTZ,S.L, 1988, Proceedings of the workshop to review and revaluate whale watching programs and management needs, November 14-16, 1988, Monterey, California. Center for Marine Conservation, Washington, D.C., and NMFS Office of Protected Resources. 1335 East West Hwy., Silver Spring, MD, 20910. 53p, BAKER, C,.S,, PALUMBI, S.R., LAMBERTSEN, R.H., WEINRICH, M.T., CALAMBOKIDIS. J. AND O'BRIEN, S.J. 1990. Influence of Sesonal migration on the geographic distribu- Hion of mitochondrial DNA haplotypes in whales. Nature 344: 238-240. BEACH, D.W. AND WEINRICH, M.T. 1989, Watching the whales; is an educational adven- jure for humans turning oul to be another threat for endungered species? Oceanus 32(1):84-88. BRAHAM, H.W. 1984. The status of endangered whales; an overview, Mar, Fish, Rev, 46(4):2-6. FOWLER, C.W. AND SMITH, T.D. 1981. ‘Dynamics of Large Mammal Populations’. (lohn Wiley and Sons, Inc.: New York) 477p. GASKILL, HLS. AND LIEN, J. in prep. Theoretical MEMOIRS OF THE QUERBNSLAND MUSEUM contribution of prenancy rates which dynamieal- ly adjust ta population density to growth in populations of Blue Whales (Balaenoptera mus- coulis). GERACI, J.R. 1989. Clinical investigation of the 1987-1988 mass mortality of bottlenose dol- phins along the U.S. Central and South Auantic Coast, Final report to National Marine Fisheries Service, U.S. Navy, Office of Naval Research, and Marine Mammal Commission. 63p-(un- publ.). HAMMOND, PS, 1986, Estimating the size of naturally marked whale populations using cap- \ure-réecapture techniques, Rep, int, Whal, Commn (special issue 5); 253-282, HAMMOND, P.S. 1990. Heterogengity in the Gulf of Maine? Estimating population size fram in- dividual recapture data when capture prob- abilities are not equal, Rep, int. Whal, Commn (special issue 12): 135-140, HAMMOND, P.S., MIZROCH. S.A. AND DONOVAN, G.P. (eds.). 1990 Individual recog- nition of cetaceans: use of photo-identificanon and other techniques to estimate population parameters. Rep. int. Whal. Commn (special jssuc 12), 440p, KAPEL. F.O, 1979. Exploitation of large whales in west Greenland in the 20th century, Rep. int. Whal. Commn 29: 197-214. KATONA,S,K, AND BEARD, J.A, 1990, Population size, migrations anid teeding ayerepations of the humpback whale (Megaprera novaeangliae) in the western North Atlantic Ocean. Rep. Int. Whal, Commn, (special issue 12); 295-306. LIEN, J., STENSON, G.B. AND NI, H. 1990, A review of incidental entrapment of seabirds, seals und whales in inshore fishing gear in New- foundjand and Labrador! a problem for fisher- men and fishing gear designers. 67-71. In ‘Proceedings of the World Symposium on Fish- ing Gear and Fishing Vessel Design’. (New- foundland-Labrador Institute of Fisheries and Marine Technology: St. Johns, Newfoundland), MITCHELL, E.D. 1973, Draft report on humpback Whales taken under special scientific permit by Canadian land stations, Rep. Int, Whal. Commn 23;138-154. MITCHELL, E.D. AND REEVES. RR, 1983. Catch history, abundance, and present status of northwest Atlantic humpbuck whales. Rep. Int. Whal. Commn (special issue 5); |53-212. NITTA, E.T, AND NAUGHTON, J.J, 1989, Species profiles: Life histories and environmental re- quirements of coastal vertebrates and inver- tebrates, Pacific Ocean Region; Report 2, PLANNING RECOVERY OF HUMPBACK WHALE POPULATIONS Humpback whale, Megaptera novaeangliae. Technical Report EL-89-10, prepared by Na- tional Marine Fisheries Service, National Oceanic and Atmospheric Administration, Honolulu, Hawaii, for the US Army Engineer Waterways Experiment Station, Vicksburg, Mississippi. Available from NTIS, 5285 Port Royal Rd., Springfield, VA 22161. NMFS 1990, Humpback Whale Recovery Plan. Of- fice of Protected Resources. 1335 East West Highway, Silver Spring, Maryland. 126p. ORITSLAND, T., OIEN, N., CALAMBOKIDIS, J., CHRISTENSEN, I., CUBBAGE, J.C., HARTVEDT, S., JENSEN, P.M.. JOYCE, G.G., TELLNES, K. AND TROUTMAN, B.L. 1989. Norwegian whale sightings surveys in the North Atlantic, 1987. Rep. int. Whal. Commn 39:411-415. PRICE, W.S. 1985. Whaling in the Caribbean: histori- cal perspective and update. Rep. Int. Whal. Commn 35:413-420. WARD, N. 1987. The whalers of Bequia. Oceanus 30(4);89-93. WHITEHEAD, H. 1987. Updated status of the humpback whale, Megaptera novaeangliae, in Canada, Canadian Field—Naturalist 101(2):284- 294. WHITEHEAD, H., SILVER, R. AND HARCOURT, P. 1982. The migration of humpback whales along the northeast coast of Newfoundland, Can. J. Zool., 60:21732179. WHITEHEAD, H. AND GLASS, C, 1985, The sig- nificance of the Southeast Shoal of the Grand Bank to humpback whales and other cetacean species. Can, J, Zool. 63:2617-2685. WINN, H.E. AND REICHLEY, N. 1985, Humpback whale Megaptera novaeangliae, 241-274. In Ridgway, S.H. and Harrison, R. (eds) *Hand- book of marine mammals, Vol. 3:The Sirenians and baleen whales.’ (Academic Press:London). MEMOIRS OF THE (QUEENSLAND MUSEUM BRISBANE © Queensland Museum PO Box 3300, South Brisbane 4101, Australia Phone 06 7 3840 7555 Fax 06 7 3846 1226 Email qmlib@qm.qld.gov.au Website www.qm.qld.gov.au National Library of Australia card number ISSN 0079-8835 NOTE Papers published in this volume and in all previous volumes of the Afemoirs of the Queensland Museum maybe teproduced for scientific research, individual study or other educational purposes. Properly acknowledged quotations may be made but queries regarding the republication of any papers should be addressed to the Editor in Chief. Copies of the journal can be purchased from the Queensland Museum Shop. A Guide to Authors is displayed at the Queensland Museum web site A Queensland Government Project Typeset at the Queensland Museum HUMPBACK WHALES {MEGAPTERA NOVAEANGLIAE) IN THE WESTERN NORTH ATLANTIC OCEAN STEVEN K. KATONA AND JUDITH A, BEARD Katona, §.K. and Beard, J.A, 1991 07 01; Humpback whales (Megaptera novaeangliae) in the western North Atlantic Ocean. Memoirs of the Queensland Museum 30(2): 307-321. Brisbane. ISSN 0079-8835. individual humpback whales can be recognized by natural markings on their bodies, The North Atlantic Humpback Whale Catalogue contains photographs of the ventral flukes of over 4000 humpback whales. More than a thousand resightings over the past 15 years have yielded information on seasonal distribution, population substructure and abundance. This paper reviews the biology of humpback whales in the North Atlantic Ocean, highlighting contributions made by phata-identification studies, and indicates avenues for future re- search. During boreal summer, North Atlantic humpbacks form at least 5 geographically distinct feeding aggregations al Jalitudes from aboul 42°—78° N. Known feeding aggregations occur in the Gult of Maine (c. 400 individuals), Gulf of St. Lawrence (c.200); Newfoundland and Labrador (c.2500); western Greenland (c.350); lceland-Denmark Strait (up to 2,000); and Norwegian Sea (1000). Photo-identified whales from all feeding aggregations including and to the west of Iceland spend boreal winter near the Virgin Islands, Puerto Rico and the Dominican Republic (c.17--22°N), where courtship, interbreeding and. calving occur. Some of these whales pass close to Bermuda during the northward migration. The winter ground(s) for European humpbacks is nul yet known. A variance-weighted estimate for lhe total population of humpback whales in the North Atlantic Ocean west of Iceland during 1978-1988 is 506623266) (95% Cl). Steven K, Katona and Judith A, Beard, College of the Atlantic, 105 Eden Si,, Bar Harbor, Maine, USA 04609; 20 December, 1990, Humpback whales (Megaptera novacangliae) play important roles im the ecology and economy of the North Aulantic Ocean. Here, as in other oceans, they were economically significant to aboriginal people and commercial hunters as a source of meat and oil. Now, the accessibility of their coastal habitats, their willingness. to ap- proach boats, and the frequent opportunities they provide for people ta watch feeding, breaching or other interesting behaviors have made them the mainstay of whale watching industries along the New England coast and in eastern Canada. Commercial whale watching tours in New England, for example. now take out nearly 1,500,000 people each year, representing over $23,000,000 in ticket sales alone, On the other hand, fishermen along the coast of Newfound- land regard humpbacks as pests, because their frequent entrapment or entanglement in shorefast nets set for salmon, cod or capelin, cause gear damage, loss of fishing time and economic hardship (Lien et al., 1990), Although the difficulty of estimating prehunt- ing humpback abundance (NMFS, 1990) con- founds attempts to evaluate its ecological role in the past, it is certainly important at some loca- tions today. During summer, the species. is the major cetacean predator in the southwestern Gulf of Maine and at some locations in the Canadian Maritimes. During winter, humpbacks migrate to warm, low latitude waters where they feed only rarely (Baraff et al., 1991), but where they may be the dominant cetacean in biomass, and their sounds, excretions, or other factors may be of ecological significance (Katona and Whitehead, 1988), Early descriptions of humpbacks in the North Atlantic came from opportunistic observations at sea or dead animals killed during hunting or found dead along the shore (e.g., True, 1904; Allen, 1916; Ingebritsen, 1929), The com- prehensive studies of large samples in the Southern Hemisphere (e.g. Matthews, 1937; Chittleborough, [958,1965) or the North Pacific (e.g. Nishiwaki, 1959) were not carried out in the North Atlantic, because humpbacks were al- 308 NORTH AMERICA 40 GREENLAND NORTH ATLANTIC OCEAN MEMOIRS OF THE QUEENSLAND MUSEUM BEAR T. + BARENTS SEA AFRICA 28 4) 28 FIG, 1. Map of North Atlantic Ocean. Modified from Jonsgard (1966). teady scarce before such studies became routine. Schevill and Backus’s (1960) observation over a 10 day period of a humpback whale swim- ming near Portland, Maine, was one of the first attempts to study a free-living humpback for purely scientific purposes. Their use of distinc- tive dorsal fin shape and fluke pigmentation to conclude that the same individual was present on different days may have been the first modern use of the technique that has become a cornerstone of humpback whale research. Ex- amples of the development and use of fluke photographs to identify humpback whales in the North Atlantic and other oceans are contained in Hammond et al. (1990). METHODS FOR PHOTO IDENTIFICA- TION OF HUMPBACK WHALES Pigmentation patterns on the ventral surface of the flukes of humpback whales were HUMPBACK WHALES IN THE WESTERN NORTH ATLANTIC photographed, printed in standard format and analyzed as discussed in Katona and Whitehead (1981), Katona and Beard (1990)and Lien and Katona (1990). Approximately 10.000 photographs of the flukes of humpback whales collected by research workers and amuteurs throughout the Atlantic region from 1967 to the present are maintained at College of the Atlantic (COA) in Bar Harbor, Maine, as the North At- lantic Humpback Whale Catalogue. Con- tributors are asked fo submit at least one fluke photograph of every individual photographed each season. Each photograph obtained is com- pared to the entire collection to determine whether it represents a previously knawn whale or a new whale. Photographs of the same in- dividuals laken al different times and places are used to study long distance migrations. pupula- tion subdivisions, and to estimate population abundance using capture-recapture techniques (Hammond, 1986). Updated sightings data from the COA catalogue are provided periodically to contributors. of photographs, some of whom maintain regional photographic collections con- taining detailed sighting records for individual whales in localized study areas that are used, for example, to chart reproductive histories of individual females (Clapham and Mayo, 1987a,b; 1990), or investigate social behayiar (Weinrich and Kuhlberg, in press). HUMPBACK WHALES IN THE NORTH ATLANTIC OCEAN A. DISTRIBUTION AND FEEDING ON THE SUMMER RANGE During summer, humpback whales feed over the continental shelf and along coastlines from the British Isles north to Bear Island (75°N) and GULF OF MAINE GULF OF ST. LAWRENCE NEWFOUND- LAND LABRADOR n=1477 R=213 (14%) n=594 R=307 (62%) 309 Spitsbergen (78°N), and around Iceland, south- western Greenland, Newfoundland and Lab- tador, the Gulf of St. Lawrence, and the Gulf of Maine (Tomilin, 1967; Leatherwood eral., 1976; Whitehead et al., 1982; Mitchell and Reeves, (983; Katona et al., 1983; Perkins et al., 1984, 1985; Payne et al., 1986; Whitehead, 1987). This distribution was documented relatively early from locations of Whaling catches, but pre- vious investigators (e.g. Kellogg, 1929: Mit- chell, 1974) had little or no information about movements between those regions and no effec- tive method for obtaining more. About 15 years ago, the questions raised by those authors began to be answered by charting the movements of photographically-identified humpbacks. Our conclusions regarding migrations, population subdivisions and abundance of humpback whales in the North Atlantic Ocean are drawn from the study of 10,566 photographs, represesenting 4,021 individual whales, By December. 1988, 1,428 individual whales had been seen on more than one day (a total of 4,012 sightings) and 1,083 individuals had been seen in more than one season or in different gea- graphic regions (a total of 3,173 sightings). Photographs from field seasons subsequent to 1988 are still being analyzed. Geographic patterns of resightings on the summer range (Fig. 2) suggest that during sum- mer the population of humpback whales from Iceland westward is divided into several relative- ly distinct units. Individually-identified whales from Iceland, southwestern Greenland, New- foundland and Labrador, the Gulf of St. Lawrence. ar the Gulf of Maine returned Tepeatedly to those same waters. We use the term “feeding aggregations” to describe the groups of whales using those separate parts of the feeding GREENLAND ICELAND EASTERN NORTH ATLANTIC n=155 R=a2 2 “12 (27%) FIG. 2. Movements of photographically identified Humpback Whales on the North Atlantic summer range. Boxes represent regional feeding aggregations. Number of individuals photographed (n); number resighted in a region in different years (R); and percentage resighled in the region in different years are shown. Numbers af individuals photographed in more than one region are shown between boxes. LOCATION Iceland Greenland Newfoundland capelin (Mallorms yillosus), sand lance (Arunodyies spp. ), Gull of St. Lawrence Nova Scotia lance (Avynedyte's sp.), cuphausiids east cous! USA mackerel (Scamber scombrus), krill norvewica) ismall fish, krill, sund Janee (Avmmnodyles sp. herring (Clupeu harengus), krill (Meganveniphanes narvegica) sand Jance (Ammadyres sp.), herring (Clupea harengus), MEMOIRS OF THE QUEENSLAND MUSEUM PREY SPECIES SOURCES Kapel,1979; Perkins etal. 1982 cuphausiids, haddock,| Mitchell,1974; Bredin,1983; White- (Melanogrammus aeglefinus), mackerel (Scomtber scambrus),| head and Glass, 1985; Whitehead and uid (lex llecebrasus Carseaddén, 1085 herring (Clupea harenyus), capelin (Mallors villosus), sand|R. Sears pers. comm, Brodie et al..1978; C. Haycock pers, comm... 5. Katona unpubl, data Meyer el al.,1979; Overholts and Nic- olas, 1979: Watkins and Schevill,1979; Hain et al.,1982; Katona et al.,1983; Kenney, 1984; Hays et al.,1985; Kenney et al.,l985, Kenney and Winn.1987: Winn et al.,1987; Mayo et al., 1988; Geract etal, N89 (Meganyctiphanes TABLE 1, Prey species utilized by humpback whales in the North Atlantic Ocean range, Little interchange was seen between feed- ing aggregations (Fig. 2). In same cases, inter- change shown between regions may be animals that happened to be seen while enroute to their feeding destination, For example, the three in- dividuals photographed in both the Gulf of Maine and Newfoundland may have used the Gulf of Maine only for migration, since they were observed there at the very beginning and very end of the feeding season, Interchange be- tween the feeding aggregation in the Gulf of St. Lawrence and those in Newfoundland (includ- ing Labrador) or the Gulf of Maine is relatively large because of their proximity. Individuals bound for the Gulf of St. Lawrence probably swim through waters used by other feeding ag- gregalions, Photo-identification studies from the eastern North Atlantic Ocean are planned or in progress, but not cnough data are yet available for com- parable analysis. No resightings were found of 12 individuals from the eastern Atlantic repre- sented in the COA collection. Distribution on the summer range is direetly dependent on the distribution and abundance of prey. However, since humpbacks in the North Atlantic utilize several types of prey (Table 1), the pattern is somewhat more complex than in the Antarctic, where krill, primarily Luphausia superba, dominates the diet (Matthews, 1937; Laws, 1985). In the western North Atlantic, humpbacks arrive by mid-April at the Mas- sachussetts coast, the portion of the feeding tange closest to the winter range. In more nor- therly ateas they usually appear by May or early June, with peak numbers in July or August, Typical annual movements within a feeding region are keyed to annual cycles of prey abun- dance. For example, the intensive Whalewatch- ing industry operating from mid-April to mid-October along the Massachusetts coust ex- ists because many ‘whales remain in thal region throughout summer, feeding mainly ona popula- tion of sand lance resident at Stellwagen Bank (Mayo et al., 1958). In contrast, humpbacks migrate steadily along the coast of Newfound- land following the northward progression of capclin spawning from June through October (Whitehead et al., 1982). Feeding areas have changed substantially be- tween Weeks or years depending on local abun- dance and distribution of prey (see Table 1). No strong evidence of age or sex class segregation has been found on the summer range. For example, in continental shelf waters of the U.S., the geographic distributjon of mothers with calves and of juveniles is similar to that of other humpbacks (Goodale, 1982). By late auiumn, most humpbacks begin migrating to lower latitudes. However, winter records from Newfoundland (Williamson, 1961; Jon Lien, pers. comm.), the Gulf of Maine (CETAP,, 1982; Mayo et al., 1988; C. Haycock, pers. comm.) and northern Norway (Rister, 1912, cited in Kellogg, 1929) indicate that some whales may stay at high latitudes all winter. HUMPBACK WHALES IN THE WESTERN NORTH ATLANTIC DOMINICAN REPUBLIC VIRGIN ISLANDS n=1305 RBZ (6%) n=131 R=3 (2%) FIG.3 31) CAPE VERDE ISLANDS n=0 . Movements of photographically identified humpback whales on the West Indies winter range. Number of whales photographed (n) is shown lor each breeding location, along with number (R) and percentage of whales photographed at that location during dillerent winters. The number of whales photographed in more than one localion is also shown. Events observed during late November in the southwestern Gulf of Maine, preceeding migra- tion to the West Indies winter range, include increased numbers of whales, energetic swim- ming behavior and song production (Mattila et al., 1987). Migration routes of different feeding aggrega- tions between the summer range and the winter Tange are not known. Humpback whales probab- ly migrate well offshore to their principal winter range around the Greater and Lesser Antilles, since no winter sightings have been recorded along the U.S. coast or at Bermuda. Songs of humpback whales enroute to the winter range were heard on underwater sound recordings made east and south of Bermuda and also south- west of there (Clapham and Mattila, 1990). B, DisTRIBUTION ON THE WINTER RANGE From late December through carly April, most of the western North Ailantic humpback popula- tion is found at Silver Bank and Navidad Bank, reefs approximately 607 nm and 157 nm, respectively, located about 120 km north and northwest of Puerto Plata, Dominican Republic. At least 2000 humpbacks vecur there from December to early March (Balcomb and Nichols, 1978,1982; Whitehead and Moore, 1982). Winn et al. (1975) estimated that 85% of the entire western North Atlantic breeding population used Silver and Navidad Banks. Sig- nificant numbers of humpback whales also winter along the Dominican coast (Mattila et al., 1988), the northwest coast of Puerto Rico (Winn et al., 1975; Mattila, 1983) and on the Virgin Bank (Winn and Winn, 1978; Mattila and Clapham, 1989). Movement of whales between those portions of the winter range occurs within a season and between years (Fig. 3). The remainder of the population may be scattered throughout the Lesser Antilles as far as Venezuela (Winn and Winn, 1978). The wintering grounds of eastern Atlantic humpbacks. have not yet been documented by either recovery of artificial tags or pholo-iden- tification studies. Humpbacks were killed by hunters during winter around the Cape Verde Islands (Kellogg. 1928; Townsend, 1935; Mitchell and Reeves, 1983) and along the coust of northwestern Africa (Kellogg, 1928). Kellogg’s (1928) map of humpback migrations showed all eastern North Atlantic humpbacks using those winter grounds, but the possibility that some easiern Atlantic humpbacks winter around the Antilles can not be ruled out, The Cape Verdes are now being suryeyed using photo-identification techniques (F, Wenzel, pers. comm.). Courtship (Tyack and Whitehead, 1983), sing- ing (Winn et al., 1981; Payne and Guinee, 1983: Payne and Payne, 1985), newborn calves and nursing (Mattila and Clapham, 1989) have all been observed on the winter range. Copulation, which has never been documented in this species, is also presumed to take place there. Differential habitat use by reproductive class occurs on the West Indies winter range. Calm waters around coral heads and reefs providing lee from the trade winds are used preferentially by females with calves (Whitehead and Moore, 1982; Goodale, 1982). Singing males may prefer locations with flat bottoms (Whitehead and Moore, 1982). Migrations to or from the West Indies winter range have been documented for 339 photographically-identified humpbacks from all feeding aggregations from Iceland west (Fig. 4}. Photo-identification has also shown that surface- active courtship groups contain whales from dif- ferent feeding aggregations (Mattila et al., td — tm GULF OF MAINE GULF OF ST. LAWRENCE NEWFOUND- LAND MEMOIRS OF THE QUEENSLAND MUSEUM GREENLAND ICELAND LABRADOR n=594 n=132 \\ n=1477 232 n=155 Va WEST INDIES WINTER RANGE n=1826 FIG. 4. Migrations between eastern North Atlantic summer range feeding aggregations and West Indies winter range. Number of individuals (n) photographed at different locations is shown along with the number photographed in different locations (on interconnecting lines). 1989), indicating that all humpbacks from the western North Atlantic, at least, probably inter- breed. Some humpbacks remain in the West Indies through March, but most depart earlier. Peak numbers at Silver Bank occured in February (Balcomb and Nichols, 1982). Photo-identified individuals (n=26) from all five western Atlantic feeding aggregations have been observed during northward migration near Bermuda in April and early May (Stone et al., 1987) (Fig. 5), but some whales probably take a a mid-ocean route to the summer range. Minimum mean swimming speeds of 3.29 km/hr (2,684 km in 34 days) and 2.28 km/hr (2,351 km in 43 days) were computed for two photographically-identified whales migrating from the West Indies to the Gulf of Maine feeding aggregation (Clapham and Mat- tila, 1988). The slower rate was close to mean speeds calculated by Dawbin (1966). If all western North Atlantic humpbacks inter- breed, as suggested by Mattila et al. (1989), and since a calf must nurse during its first migration to the summer range, persistent subdivision of the population on the summer range must be maternally directed. Such divisions probably are caused by the tendency of a calf to return to the portion of the summer range used by its mother and learned during its first summer (Martin etal., 1984; Baker et al., 1990), rather than by a geneti- cally fixed behavior pattern. Several studies indicate that maternally directed behavioral fidelity to feeding grounds has been maintained for many generations. In the North Atlantic, analysis of the amount of pig- mentation on the ventral surface of flukes revealed significant differences between most feeding aggregations (Beard et al., in prep.). In the North Pacific, Baker et al. (1990) found significant differences in mitochondrial DNA haplotypes between humpback whales sampled in southeastern Alaskan waters and those sampled offshore from central California. C. REPRODUCTION Reproductive parameters for humpbacks from the New England coast obtained from photo- identified individuals in the Gulf of Maine gave a crude birth rate ranging from 0.45 in 1981 to 0.103 in 1983; the same data yielded reproduc- HUMPBACK WHALES IN THE WESTERN NORTH ATLANTIC GULF OF MAINE GULF OF ST, LAWRENCE NEWFOLIND- LAND 313 GREENLAND ICELAND LABRADOR, n=594 bars n=132 n=1477 SIZ n=ZO il n=155 BERMUDA nellS WEST INDIES WINTER RANGE n=1826 FIG. 5, Distribution of resightings of individually identified humpback whales photographed tive rates of 0.30-0.43 calves per mature female per year (Clapham and Mayo, 1987a). Those authors and their colleagues at the Center for Coastal Studies (CCS), Provincetown, Mas- sachusetts, now have a sample population of 97 mothers and 207 calves. The numbers of calves produced by individually identified mothers as of September, 1990, are 7 (n=l), 6 (n=3), 5 (n=5), 4 (n=6), 3 (n=14), 2 (n=22) and | (n=46). Nine (9) females are now grandmothers, two of them for the second time (L. Baraff, pers. comm.). As detailed by Clapham and Mayo (1987b), calves were usually born at intervals of 2 or 3 years. Two whales known from their birth year and seen annually bore calves of their own, at ages of no more than 4 years and 6 years, respectively; others with incomplete sighting histories calved at minimum ages of § (n=2), 6 (n=3), 7 (n=2) and 9 (n=l) years. D. HistoricAL ABUNDANCE Mitchell and Reeves (1983) summarized the history of humpback whale hunting in the western North Atlantic Ocean, starting at Ber- muda in 1611, Maine in 1675, Georges Bank in 1725, the Gulf of St. Lawrence in 1819, the West Indies wintering grounds in 1822, and the Canadian maritimes in the late 1800’s, They accounted for at least 9.125 humpbacks killed during 1850-1971, and used their assembled catch estimates to calculate that the population size in 1865 was greater than 4,700. Breiwick et al, (1983) used the same data, but incorporated estimates for annual natural mortality (4%) and net recruitment (3%), and revised that estimate to 6,300 whales. However, many more humpback whales may have been present originally, because humpbacks had been hunted for several centuries before 1865, although catches were poorly documented (Reeves and Mitchell, 1982). Winn and Reichley (1985) listed 10,000+ as their estimate for the original population in the western North Atlantic. Commercial hunting could have reduced the North Atlantic humpback population to as few as 700 animals by 1932 (Breiwick et al.,1983). Hunting of females with calves at locations where humpbacks returned annually for breed- ing probably contributed to their rapid depletion on some winter grounds Winn and Scott (1981). No estimate of historical abundance is avail- able for humpbacks in eastern North Atlantic waters. As reported in Reeves and Mitchell (1983), Ingebritsen (1929) calculated that 3,300 314 whales were killed (here during 1885-1927, in- cluding 1,500 off Finnmark (central Norway) and 1,500 off Iceland, Since we now know that humpbacks from at least as far cast as. Iccland migrate to the West Indies und probably inter- breed with the western North. Atlantic stock (Mattila et al, 1989), it is unclear how Inge- britsen’s data could be used to evaluate historical abundance for eastern Atlantic humpbacks, E, ESTIMATE OF CURRENT ABUNDANCE Population estimates for the western North Atlantic have shown an upward trend since the 1960's, however the increase is believed to be principally due to improvements in sampling effort and methodology (Whitehead, 1987). We report here (Fig. 6) a new estimate of S066 whales (95% confidence interval, 1800-820), based on whales photo- identified during 1978- L988 and computed by methods detailed in Katona and Beard (1990). This represents about 80% of Breiwick et al.’s (1983) estimate for the population in 1865. oestimate is available for current abundance of humpbacks in European waters from Iecland eastward. Luckily, Ingebritsen’s ( 1929) state- ment that hunting appeared “to have entircly exterminated” the species in thuse waters was not correct, and studies using phote-jdentifica- tion and census surveys are tn progress. Gunn- laugsson and Sigurjonnson (1990) used sight- ings from ships to cstimate a population of less than 2000 humpbacks around Iceland. This was considered a great increase since protection began in 1955 anda considerable increase since the 1960°s (Sigurjonsson and Gunnlaugsson, 1990), However, as shown in Sigunjonsson and Gunolaugsson (1990), that estimate includes many sightings of humpbacks in the Denmark Strait that are probably affiliated with the western North Atlantic population, plus others sighted east of [cejand that could be from the eastern Atlantic feeding aggregation(s). Visual surveys [rom ships, by the Norwegian Institute for Marine Research indicated abour |,000 humpbacks in the Norwegian See and wlong the Norwegian coast in summer(Oicn, 1991). PHOTO-IDENTIFICATION AND FUTURE DIRECTIONS FOR INVESTIGATIONS ON HUMPBACK WILALES IN THE NORTH ATLANTIC OCEAN For studying migration and population sub- MEMOIRS OF THE QUEENSLAND MUSEUM divisions, identification of individuals using natural markings provides the same kind of in- formation that was previously obtained by ap- plying artifical tags. For a species like the humpback, which possesses great individual variation in naturdl markings and an equally salient variely of acquired scars, each individual is marked so distinctively that artifical tags are nol needed, Changes in pigmentation that occur in some individuals during carly years da not compromise the accuracy of results if photo- comparisons are done carefully (Carlson et al., 1990). Photo-identification studies do not harm target animals, are rclatively inexpensive, and produce many more recaptures than did artificial tagging. For example, out of approximately 3000 humpback whales tagged with “Discovery” tags during commercial whaling or scientific studies in the Antarctic areas IV and V, 123 tags (4%) were recovered (Dawhbin, 1966), The recovery rate is even lower in the western North Atlantic, where, according to Mitchell and Reeves (1983), over [OU successful implantations of Discovery tags in humpback whales produced only one significant recapture, an animal that retumed to western Greenland waters in successive years (Mitchell, 1974, 1977). By comparison, the per- centage of whales resighted in the same feeding ugeregalion during different years ranged from 5% for Iceland, which has. the smallest sample size (n=20), to 62% for the Gulf of Maine, the most heavily sampled region (n=594). The number of identified humpbacks in te- search collections is probably now nearly 8,000, including collections from Antarctica (50, Cal- lege of the Atlantic, Stone et al., 1990, and unpublished data); Australia (600, Kaufman, 1990); Brazil (60, S_ Siciliano, pers. comm.), Colombia (SQ, L. Flérez-Gonzalez, pers. comm.), and the North Pacific Ocean (perhaps 2,500, Sally Mizroch, National Marine Mammal Laboratory, NMFS, pers, comm.,, including col- lections from Mexico, Hawaii, Japan, California, and Alaska), The amount of informatton on whale movements and life history will inercase steadily os the number of photographs increases in future field seasons and as comparisons be- tween regional catalogues are donc. Sophisticated cameras and film necessary for high speed, high resolution distance photag- raphy of whales did not exist during most of the period of artificial tagging, nor did the high speed hoats that now bring photographers clase to whales. Consequently, photo-identification AUMPBACK WHALES IN THE WESTERN NORTH ATLANTIC studics now carried out on many cetacean species (Hammond et al., 1990) probably would not have supplanted artificial tagging much ear- lier than they did. Despite jis current utility, studies of hump- buck migration using natural markings and those using artificial marking share the Weakness that unly the beginning and end points of the move- ment are documented. Tracing migration roules will require improvements in tracking with radio tags using satellite technology (Matec, 1989), Estimating population abundance using photo-identification is a useful, and sometimes preferable (Hammond, 1986) alternative to visual census surveys from ships or airplanes (e.g. Scott and Winn, 1980; CETAP 1982) and acoustic census surveys (c.g, Winn et al., 1975; Levenson and Leapley, 1978). Abundance es- timates presented here for humpback popula- lions of the North Atlantic and similar estimates for the North Pacific Ocean (Perry ct al., 1990) are probably the most accurate available for any ectacean population over a Whole ocean basin. Nevertheless, this method can still be im- proved. Temporal and regional biases in sam- pling effort exist, and the opportunistic sampling methods usually used may bias for or against certain portions of the population (Hammond, 1986.1990). Use of a standardized samplin protocol to equalize the opportunities for a Whales within @ population’s range to be photographed would improve the accuracy and prectsion of population estimates. Data from photo-identification studies pro- vide interesting contrasts to information from hunted specimens. Inspection of tens of thousands of carcasses allowed scientists such as Chiteborough (1958,1965) ta compute mean Values for life history parameters such as age, age of sexual maturity, fecundity, or growth rate. Precise us they are, such results contain lwo grades of uncertainty. First, they are necessarily indirect. For example, corpora albicantia, embryos or fetuses show thal ovulation or preg- naney occured, but do not indicate whether a calf was born, whether it survived, or exactly how frequently reproduction occurred. Second, mean results do not describe what any real live, in- dividual whale ever did. Data from long term photo-identification of individuals creates the opposite problem. It can yield precise information on the reproductive rate of individuals (Glockner-Ferrari and Ferrari, 1984: Baker et al., 1Y86; Clapham ani Mayo, 19&7b, 1990) and, poiemially, the survival and 315 reproduction of their calves. However, since the large sample sizes provided by commercial whale huntiig can not be obtained in a shor time, stalisticully precise means for the popula. tion are more difficult ta achieve. Nevertheless, the ability to describe the activily of a small number of real individuals offers a new perspec- tive on the biology of the species. A technical problem confronting large scale photo-identification studies is the increased time required for sorting large collections of photographs. It takes only a moment to read the numbers an a “Discovery” tag, and anybody can do it. Comparing photographs requires special skills; patience, allentiveness, facility at pattern- recognition and good visual memory, Only people possessing those skills should match photographs for scientific purposes. Computers gel high marks tor patience, attentiveness and information retention, but instructing them to find and retmember a pattern is laborious. A program developed by Mizroch etal. (1990) uses videodise storage of photographs and combines the strengths of people and computers to speed photo-comparisons. [t is now used routinely tn our laboratory and several others. Use of computers will increase the rate at which photographs can be matched, reducing the number of photographs that must be checked visually, and minimizing tedium. Because it al- lows photographic collections to be sorted in many ways, Mizroch et al.'s (1990) system has also helped researchers to find duplicates in photographic collections. In collaboration with Sally Mizroch and the National Marine Mammal Laboratory (NMFS), we are incorporating digi- tal analysis of images for comparison of photographs, to enhance speed and accuracy, One side benefit will be to enable researchers ta compare large collections of photographs from differcnl oceans, a project loo expensive and time-consuming to be undertaken manually. While contemplating such technological im- provements, it ts worth emphasizing that our cyes and brains have undergone millions of years of evolution for improved pattern recognition, These organs routinely perform complex visual operations such as translation, rolatton, compen- sation, and decision-making that are cumber- some for even the best of today’s computers. Machines will never entirely replace skilled people in analyzing individual identification photographs. Continuing to loak toward the filuns, chul- lenging scientific puzzles remain to be solved for Alé this species in the North AUuantic and other oceans, (1) During this deeade, studies of migra- tions in the eastern North Atlantic using photo- identification will probably define the location of winter ranges and reveul the degree of separa- tion between those animals and the western At- lantic population. (2) Precise description of migration routes for individual humpbacks is needed to provide clues about navigational mechanisms, evaluale polential energetic benefits from using wave energy to assist locomotion (Bose and Lien, 1990), facilitate detailed studies of migration and ascertain potential environmental risks to whales as they migrate through different regions. (3) Descrip- tion of where, when and how copulation occurs is needed for comparison with other mating sys- ters (Brownell and Ralls, 1986; Clutton-Brack, 1989), (4) Continued intensive regional sam- pling of individuals, particularly with improved sampling procedures, is needed to provide im- proved estimates of natality and mortalily, allow construction of a life table, and reveal long term patterns of social behavior. Enough mothers need to be photographed on the winter ground with calves and then Jateron the summer ground to provide an estimate of calf mortality during the first six months of life. A particularly dif- ficult, but important goal is te learn more about causes of natural mortality. The only causes specifically identified to date are killer whales (Orcinus orca) (Dolphin. 1987; Katona et al., 1988), sharks (Winn and Reichley, (985; Pater- son and Van Dyck, this memoir) and dinoflagel- late toxins (Geract et al,, 1989). (5) Expanded use of DNA-sampling is needed to facilitate gender identification (Lambertsen ct al., L988: Baker et al., unpublished manuscript), elucidate gene flew produced by migrations (Baker et al., 1990), and contribute to interpretation of Jongierm matrilineal, patrilineal and social relationships. Some of the solutions to those puzzles will not only sate our curiusily, but will allow us to respond more effectively to the overall challenge of managing humpback whales and their habitats successfully enough so that the populations recover from depletion by commercial whaling and swim the wide aceans for countless genera- tions ta come, Until Star Trek TV comes true, (here is probably nothing more beneficial that we can do for this species than gather information like this and use it wisely. [Lis humbling to reflect on how much of the data and how many of the future objectives MEMOIRS OF THE QUEENSLAND MUSEUM presented here are merely fleshing out of the skeleton erected by previous workers, including ern Drs. R.G. Chittleborough and W.H. awhin, in whose honor this conference has been organized. Everyone working on popula- tion biology, migration and reproduction is in- debted to their pathbreaking research, as every lest of references testifies. ACKNOWLEDGEMENTS Muintainance of the North Atlantic HJumpback Whale Cafalogue is funded by the Northeast Fisheries Center, NMFS, Woods Hole, Massachusetts. Substantial collections of photographs were contributed by workers as- sociated with the Atlantic Cetacean Research Center (Gloucester, Massachusetts); Center for Coastal Studies (Provincetown, Massachusetts); Long Term Research Institute (Lincoln, Massa- chusetts); Cetacean Research Unit (Gloucester, Massachusetts): Ocean Research Education Society (Gloucester, Massachusetts); Bricr Is- land Ocean Study (Westport, Nova Scotia); Dal- housie University (Halifax, Nova Scotia); Mingan Islands Cetacean Study (Sept Iles, Quebec, Canada); Memorial University (St. John’s, Newfoundland); Fisheries Research tn- stitute (Reykyavik, [celand); University of Ber- gen (Bergen, Norway); Sea Mammal Research Unit (British Antarctic Survey, Cambridge University, Cambridge, England), A more com- plete list of contributors may be found in Ac- knowledgements to Katona and Beard(1990), We are particularly prateful to the follawing peuple for photographs, logistic assistance, financial support or technical advice contributed lo the North Atlantic Humpback Whale Catalogue: Kenneth C. Balcomb Ill, Lisa Baral, Edward McC. Blair, Robert Bowman, Carole Carlson, Philip Clapham, Tim Cole, Harriet Cor- bett, Kevin Chu, Steve Frohock, Harold Graham, Mr. and Mrs. H.8. Guthrie, Carl Haycock, Philip A, Hammond, Finn Larsen, Jon Lien, Anthony R. Martin, David K. Mattila, Charles A. Mayo, Jr, Scoll Mercer, Nancy Miller, Sally A, Miz- roch, Karen Moore, George Nichols, Jr., Mr. and Mrs, Francis W. Peabody, Judy Perkins, Margo Rice, Kim Robertson, Richard Sears, Johann Sigurjonsson, Mr. and Mrs. Donald B- Straus, Edward B. Tucker, Gordon Waring, William A. Watkins, Mason Weinrich, Frederick P. Wenzel, Hal Whitehead, J, Michael Williamson. Thanks are also duc to College of the Atlantic for provid- ing office space and computer facilitics for this HUMPBACK WHALES IN THE WESTERN NORTH ATLANTIC project; and to the U.S. Coast Guard, Southwest Harbor Group, for permission to use facilities at Mt. Desert Rock Lightstation, Maine. LITERATURE CITED ALLEN, G.M. 1916. 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Song of the Humpback Whale — population comparisons. Behav. Ecol. Sociobiol. 8:41—-46. WINN, H.E. AND SCOTT, G.P. 1981. The humpback whale: present knowledge and future trends in 321 research with special reference to the western North Atlantic. 171-180. In ‘Mammals in the Seas. Vol. 3. General Papers and Large Cetaceans’. (United Nations Food and Agricul- ture Organization:Rome). WINN, H.E. AND REICHLEY, N. 1985. Humpback Whale-Megaptera novaeangliae.. 241-274. In Ridgway, S.H. and Harrison, R. (eds) ‘Hand- book of Marine Mammals. Vol. 3: The Sirenians and Baleen Whales’. (Academic Press:London). 362p. WINN, H.E., HAIN, J.H.W., HYMAN, M.A.M. AND SCOTT, G.P. 1987, Whales, dolphins, and por- poises. 375-382. In Backus, R. and Bourne, D.W. (eds) ‘Georges Bank.’ (MIT Press: Cambridge, Mass.). MEMOIRS OF THE (QUEENSLAND MUSEUM BRISBANE © Queensland Museum PO Box 3300, South Brisbane 4101, Australia Phone 06 7 3840 7555 Fax 06 7 3846 1226 Email qmlib@qm.qld.gov.au Website www.qm.qld.gov.au National Library of Australia card number ISSN 0079-8835 NOTE Papers published in this volume and in all previous volumes of the Afemoirs of the Queensland Museum maybe teproduced for scientific research, individual study or other educational purposes. Properly acknowledged quotations may be made but queries regarding the republication of any papers should be addressed to the Editor in Chief. Copies of the journal can be purchased from the Queensland Museum Shop. A Guide to Authors is displayed at the Queensland Museum web site A Queensland Government Project Typeset at the Queensland Museum AN ACOUSTIC ANALYSIS OF THE 1988 SONG OF THE HUMPBACK WHALE, MEGAPTERA NOVAEANGLIAE, OFF EASTERN AUSTRALIA ASTRIDA MEDNIS Mednis, A. 1991 07 01: An acoustic analysis of the 1988 song of the Humpback Whale, Megaptera novaeangliae, off east coast Australia. Memoirs of the Queensland Museum 30(2): 323-332. Brisbane. ISSN 0079-8835. This paper presents an acoustical analysis of the 1988 Humpback Whale song off east coast Australia, particularly of the sound types which comprise the song. The analysis shows that the song represented generally a well structured form, broadly similar to that of 1982/83, even though there was a change to a less structured song in 1984 (Cato, this memoir). A relatively detailed analysis of the structural components of the song, specifically the sound types, has been undertaken. Although the song is relatively stereotyped, variation in the acoustical characteristics of the sound types is observed which may be signiflcant in terms of structural components of the song. The full range structural complexity in the Humpback Whale song may not be fully known. [|] Humpback Whale, song, marine acoustics, animal behaviour, marine mammals. Astrida Mednis, Centre for Marine Science, University of New South Wales, Present address: Australian Heritage Commission, G.P.O. Box 1567, Canberra 2601; 18 February, 1991, Payne and McVay (1971) recognised that Humpback Whale sounds recorded off Bermuda occurred in fixed patterns to form songs. They found that individual sound units were grouped to form phrases which were repeated to form themes. This ordered sequence comprised the song. Based on studies of Humpback Whale songs in eastern Australian waters since 1979, Cato (1984, this memoir) established the existence of song structures that were broadly similar to those of the Northern Hemisphere. The song of 1982/83 was well structured with six themes, but a rapid change in 1984 lead toa relatively poorly structured song. Cato introduced the concept of “sound type” as a means of categorising the sound units according to acoustical charac- teristics. He found that sound units of the 1982/83 song could be classified into 12 types. I present here an analysis of the 1988 song off eastern Australia with particular emphasis on the characteristics of sound types. These are given descriptive names which are definitive to this paper. Sound types are classified according to acoustical characteristics and some of the sub- tleties in variation of characteristics are ad- dressed. This may allow interpretation in greater detail and contribute to functional understanding of structural components of the song. Cato (1984) noted that, because the song is ste- reotyped, its potential to carry information is limited. Information transmission depends on variations in the song. METHODS Song recording Song data for 1988 were collected by Dr D. Cato, during the southward migration, 2-4 Oc- tober, 1988, off Stradbroke Island latitude 27°S. This was close to the peak time of the southward migration as identified by Chittleborough (1965) and confirmed by Paterson and Paterson (1989). Recordings were made from a 3m motor boat, using a hydrophone at about 20m depth, a pre- amplifier, aSony Walkman Professional WMD6 tape recorder. The frequency responses are uniform within +3dB over the frequency range 50 Hz-15 kHz. Song analysis Song analysis involved aural and spectro- graphic analysis. Most of the energy of Hump- back Whale sounds lies in the range 100-4000 Hz and the sounds are distinctive to our ears (human hearing is within the range of c. 20Hz— 20kHz). A considerable amount of analysis of the song structure was therefore done aurally by noting down sound units sequentially using graphic symbols, where each symbol represents a sound type, thus identifying the order and timing of sound units, phrases, themes and song 324 TABLE 1. Phrase structures of the 1988 song. THEME| SOUND UNITS IN ORDER OF OCCURRENCE RENCES n-chug 19-28 1 zp 1 oink 1 n-chug evolve into whistles 11-12 cry 2 n—whistles cry 5-8 n-whistles OCCUR- tN 3 n—violin 1 coarse roar 2 down moan i] up moan 1 down moan 1 up moan 1 down moan 1 up moan l flat roar I moan staccato 3-4 screal i] up moan down moan up moan down moan up moan flat roar hmm 5-7 screal 1 a oo up moan 1 down moan | up moan 1 down moan 1 ] 1 up moan flat roar whoomp 5-8 screal 7 whoomp 2 chain-saw growl 1 cycles. The aural analysis was repeated until the song structure was fully established. This in- volved a detailed analysis of the data, looking for patterns in the occurrence of the sound units. Sound units were grouped to form phrases and similar phrases were then grouped into themes. As patterns were established, they were checked for consistency between the different song MEMOIRS OF THE QUEENSLAND MUSEUM cycles of an individual singer and between the songs of presumably different singers. Acousti- cal characteristics for each sound type and phrase structure were determined from sonagrams produced on a Kay Elemetrics sonagraph. Seventeen hours of recorded data were analysed, which included 64 song cycles able to be analysed. Intensity levels for different sound units were determined with a Bruel and Kjaer level recorder type 2305, played through a Rock- land low pass filter, to remove unwanted low frequency noise. RESULTS SONG STRUCTURE The 1988 song structure (Table 1) comprises 7 themes in a fixed order, where each theme com- prises a particular phrase with a variable number of renditions. This is a well structured song, similar to that of 1982/83 (Cato,1984,this memoir). It has 7 themes and 15 sound types compared with 6 themes and 12 sound types in 1982/83. This shows that the song off east Australia has returned to a well structured form after the change to a relatively poorly structured song in 1985. On average, the song consists of about 160 sound units, depending on the number of phrase and sound unit renditions in different themes. The Humpback Whale song is complex and relatively stereotyped; considerable analysis may be done in the description and interpretation of the song characteristics. Even though the song is relatively stereotyped, there are certain sub- tleties in the complex song which are not stereotyped. DESCRIPTION OF THE SOUND TYPES. The frequencies of all sound types is in the range 50-8000Hz (more than 7 octaves), the lowest in frequency being the “whoomps” and “down moan” and the highest in frequency being the “n-whistles” and “screal” (Table 2). In the 1988 song the range in the fundamental frequen- cy was 50-4000Hz while Payne and Payne (1985) observed a range of 30-4000Hz in songs in Bermudan waters over 19 years. The sound types vary from acoustically simple to complex and are classified into four categories (Table 2). These categories are determined from sonagrams and are audibly quite distinctive. Each category can be subdivided according to the frequency range, fundamental frequency and ACOUSTIC ANALYSIS, 1988 HUMPBACK WHALE SONG, E. AUST TABLE 2. General sound type spectral and temporal characteristics. SOUND TYPE FREQUENCY FREQUENCY OF NO, OF TIME | RANGE FUNDAMENTAL HARMONICS (s) (Hz) OR LOWEST S.L.* /S.L.* Hz) HARMONIC — rising frequency n-whistles 600-8000 600-3000 2-5 0.1-0.3 cry S00-7500 500-700 10-11 0.3=1,35 up moan 200-4000 200-350 2-<16 0.8-1.7 n-violin LOO—6000 Loo z 0.6 oink 150-2500 150 7 0.4 down moan 50-5500 50-200 2-19 0.8—2.2 — Steady frequenc hmm 120-4000 120-250 5-16 0,45-0.7 ~ collective 0.9-1.7 BROAD BAND WITH SPECTRAL LINES coarse roar flat roar 250-4500 400-3750 400-600 iz 2-8 1.24.45 250-350 —20 0.8-2.4 chain saw growl 100-4000 100-1000 6 3.0-5.7 screal 900-8000 901-1450 3-8 2.0-3.5 BROAD BAND IMPULSIVE zp 2000 80 7 O38 n-chugs 80-6000 impulsive - 0.2-0.35 COMPLEX iz — moan staccato 1.1-3.0 moan 100-340 3-15 0.8-1.8 slaccatu 300-6800 300-450 3-12 a3 — whoomp harmonic 50-1300 50-100 impulsive 600—4000 50000 impulsive O.1-0.5 — pulse 600-4000 600-1100 pulsation 0.10.2 *S.L. = spectral lines duration. The harmonic sound types “cry” and “up moan”, and the complex sound type “moan staccato” (Table 2) (which is actually a combina- tion of two harmonic sounds) have similarities: however, differences in fundamental frequen- cies, frequency contour on the sanagram and duration separate them (Figs 1-3). The “coarse roar” and “flat roar” (Table 2) are similar, how- ever, the former has a higher fundamental fre- quency range. The “down moan” is distinctive relative to aj! other sound types (Fig. 4). Harmonic sound types with a rising frequency The “cry” sound type is representative of this subcategory (Fig. 1). It lasts 0.3-1.35 secs depending on its position in the phrase in theme 2, the only theme in which it occurs. Silences between cries range from 1,1—2.8 secs, The “up moan” occurs in 4 different themes with some variation in the sound character be- tween different themes and within the same phrase of a theme depending on the position of the sound unit in the phrase, i.e. inter theme and intra phrase variation (Figs 2,4). This variation 326 FREQUENCY (kHz) MEMOIRS OF THE QUEENSLAND MUSEUM TIME (s) FIG. 1. Sonagram of an harmonic with a rising frequency sound type “cry”. is in the fundamental frequency and duration. The duration of the sound type ranges from 0.6—-2.1 secs. Silences of following units in themes 4 and 5 are 1.5-1.9 secs. Harmonic sound types with a falling frequency The “down moan” (Table 2) occurs in four different themes with some variation in fun- damental frequency and duration between themes (Fig. 4). Broad band with spectral lines not harmonic- allv related These sound types are not harmonic but have energy spread over some bandwidth, and spectral peaks evident as spectral lines. The fre- TABLE 3. Variation of “screal” between themes. quencies of these lines are not harmonically re- lated. The “screal” (Table 2) has 3-10 spectral lines (Fig. 5) and occurs once in 3 different themes, consistently as the last sound unit in a phrase. Sound character variation in the different themes is shown in Table 3, The “chain saw growl” (Table 2) generally occurs at a number of frequencies between 100 and 1600 Hz (Fig. 6). It generally increases in duration with each phrase rendition in the theme. The sound type occurs singularly in the phrase of theme 7. Harmonic sound types with a steady frequency The “hmm” (Table 2) occurs in a group of units, like the “‘n-chugs”, where 24 sound units FREQUENCY ee FUNDAMENTAL piace Ls NO. OF SPECTRAL LINES ——o a 900-8000 500-1400 | oa0-ss00 500 900-1400 1000-6800 1000-1450 | SS 3.0-3.3 2.0-3.0 ACOUSTIC ANALYSIS, 1988 HUMPBACK WHALE SONG, E. AUST 327 FREQUENCY (kHz) TIME (s) FIG, 2. Sonagram of an harmonic with a rising frequency sound type “up moan”. (a) (a) (b) (b) (a) FREQUENCY (kHz) TIME (s) FIG. 3. Sonagram of a complex sound type “moan staccato” including a, moan and b, staccato endings. MEMOIRS OF THE QUEENSLAND MUSEUM FREQUENCY (kHz) TIME (s) FIG. 4. Sonagram of an harmonic with a falling frequency sound type a, “up moan” and b, “down moan”. may occur in one group. For example, (hmm hmm hmm) (hmm hmm hmm hmm) (hmm hmm hmm), where each hmm represents one rendition of the sound unit. Each sound unit is 0.45-0.7 secs in duration while a collective group may extend 0.9-1.7 secs. The period of silence be- tween units of the same group is c. 0.05-0.1 secs and between groups there are longer silences of 0.40.9 secs. It occurs in theme 5 only. 4 Broad band impulsive sound types The impulsive sound types are the “n-chugs”, occurring in themes 1 and 2 and “zp” occurring in theme 1. The “zp” occurs only once in the phrase of theme 1. The “n-chug” sound type occurs as a collective unit. For example a collection of “‘n-chugs” may comprise 28 sound unit repetitions in a particular pattern i.e. (111) (1111) (1111) (11111) FREQUENCY (kHz) bo TIME (s) FIG. 5. Sonagram of a broad band with spectral lines sound type “screal”. ACOUSTIC ANALYSIS, 1988 HUMPBACK WHALE SONG, E. AUST 329 FREQUENCY (kHz) TIME (s) FIG, 6. Sonagram of a broad band with spectral lines sound type “chain-saw growl”, (111111) where 1=1 chug. So that the whole collective consists of subgroups each with a vari- able number of chug repetitions (Fig. 7). These units are impulsive in structure with most energy between 80—2000Hz and extending to 6000Hz. Silences of 0.20-).55 secs between subgroups and 0.15 secs between renditions of units within one group were observed. Complex sound types Complex sound types are combinations of components which may be harmonic or impul- sive. The “whoomp” sound type starts as a har- monic and evolves to an impulse with a pulsating, “possum” ending (Fig. 8). This sound type occurs in themes 6 and 7 with sound char- acter variation between these themes being of a lower frequency, less harmonic in structure and without the terminal impulse in theme 7. While in theme 6 the impulse ending may or may not occur. It is evident from the above discussion that FREQUENCY (kHz) Oo — ~ TIME (s} FIG. 7. Sonagram of a broad band impulsive sound type “n-chugs”. (a) (b) (c) FREQUENCY (kHz) MEMOIRS OF THE QUEENSLAND MUSEUM (a) (b) (c) (ce) TIME (s) FIG, 8 Sonagram of a complex sound type - “whoomp”, including (a) harmonic, (b) impulse, and (c) pulsating ending. there are variations in the characteristics of the sound lypes in the song. DuRATION OF THE SOUND TYPES Sound types vary in duration from 0.1-5.7 secs, the shortest being the “n-whistles” (theme 2) and the longest being the “chain saw growl” (theme 7). Cato observed sound type durations of 0.1-4.3 secs in the 1982 song, where the “whistles” were the shortest and the “chain saw” the longest. However, the “chain saws” of the 1982 song are not the same sound type as the “chain saw growls” in the 1988 song. In Ber- mudan waters the longest sound type was 8 secs and the shortest <2% of the longest (Payne and Payne 1985). VARIATION IN SOUND TyPE ABUNDANCE AND Dis- TRIBUTION THROUGHOUT THE SONG The sound types are variable in terms of dis- tribution through the different phrases of the song (Table 1), Some sound types are particular to certain themes i.e. *zp" and “oink” to theme 1 and “n-whistle” ta theme 2, whilst other sound types are relatively ubiquitous, I.c. “up moan”, “down moan”, “screal” and “flat roar”, occurring in three different themes and the “whoomp” occurring in two different themes, Nine of the sound types occur more than once in a phrase. The remaining occur only once ina phrase, for example “n-violin” in theme 3 and, “screal” and “flat roar” in themes 4, 5 and 6. VARIATION IN SOUND TyPE CHARACTER- INTER- THEME AND INTRAPHRASE VARIATION As sound types are repeated throughout a phrase or as they occur in different themes, the sound character (e.g. a slight change in fun- damental frequency, frequency range or dura- lion) may vary depending on the sound type’s position in the song. For example, in theme 4 the “up moan” is emitted 3 times, each rendition becoming shorter. However, the same sound type in theme 5 is generally shorter in duration. Additionally, the “moan staccato” may vary in sound character with repetition in a phrase of theme 4, and the number of staccato units may vary with each repetition. The staccato endings are mostly present but sometimes absent. Similarly, the “whoomp” in theme 6, is repre- sented with “possum-like” endings in some phrases but not in others, The character of a sound type may therefore change with respect to its position in the phrase or in the song. Although there are subtle varia- tions in the character of a particular sound type, the sound types are gencrally similar from one rendition of the song to the next. Even though there is some variation in the acoustical character of 4 sound type, this variation is small compared to the differences in acoustical characteristics between different sound types, Perhaps these observed variations can be re- lated to accentuation of meaning i.e. possibly these variations have specific functions in terms ACOUSTIC ANALYSIS, 1988 HUMPBACK WHALE SONG, E. AUST of communication and general behaviour of the singer. A slight modification in the sound type may change the signification of the signifier i.e. the meaning of the sound type. SOUND INTENSITY From a limited sample of 6 song cycles over 2 days, intensity varied between different sound types. The “moan staccato” has the highest level of all sound types while the “flat roar”, “hmm” and “screal” are the next highest. These are mean values and not always consistent with cach phrase rendition. It is important to consider this feature relative to overall rhythm (Guinee and Payne, 1988) and possible “accentuation” of cer- tain sound units. Additionally, sound level diminishes at theme 7, which may be due to attenuation of sound when the whale approaches the surface to breath (Tyack,1981). SILENCES BETWEEN SOUND UNITS In the 1988 song, silences between sound units of up to 3.2 secs were observed where the longest silences generally occurred between sound units of themes | and 7. Shortest silences were c. 0.15 secs between “n-whistles” in theme 2 and 0.2 secs between “n-chugs” in theme 1. Payne and Payne (1985) observed silences as very short or lasting up to 6 secs. The range in duration of silences between sound units may be important structural features of the song. CONCLUSIONS The 1988 song of east Australia shows a well- developed structure similar to that in the 1982- early 1984, compared to the “unstructured” song of 1985. The 1988 song consisted of 15 sound types. Sound types representative of particular acousti- cal groups are described with regard to spectral and temporal characteristics, showing the dif- ferences between sound types and between ren- ditions of the same sound type. The analysis indicates that while the sound types do not vary to a great extent with different song renditions, there are subtle variations with renditions of the same sound type which may contribute to the overall function of the song, for example the information content. It is possible that these sub- tleties function as components of the song struc- ture. A more detailed analysis of the song structure and the sound units may reveal the function of these structures and thereby con- tribute to the understanding of the song. 331 This paper presents an analysis of the song at a certain level of magnification, however, it is not fully understood whether this level of detail is at a scale appropriate to the Humpback Whale physiological sensory apparatus. It is yet to be determined how representative these identified song components are for communication. Analysis of these structural details may lead to a greater understanding of Humpback Whale song evolution, learning capacity and behaviour, and the role of song in the reproductive success and possible social structure of the species. ACKNOWLEDGEMENTS I would like to thank Dr D. Cato (Defence Science and Technology Organization ) for shar- ing his data and for his enthusiastic encourage- ment and guidance and Dr M. Augee (University of New South Wales ) for his general guidance. Additionally, I would like to thank Mr G. Plant, National Acoustics Laboratory, for the use of the Kay Elemetrics Sonagraph. Thanks also go to Dr R. Paterson and Mr L. Nash for their support in collecting the song data at Stradbroke Island. Thanks are also due to the DSTO for providing financial support for this study. LITERATURE CITED CATO, D.H, 1984. Recording Humpback Whale sounds off Stradbroke Island. 285—290. In R.J. Coleman, J. Covacevich and P. Davie, (eds), ‘Focus on Stradbroke’, (Boolarong,: Brisbane). CATO, D.H. this memoir Songs of Humpback Whales: the Australian perspective. Mem. Qd Mus. 30(2): 277-290. CHITTLEBOROUGH, R.G. 1965. Dynamics of two populations of the Humpback Whale, Megap- tera novaeangliae (Borowski). Aust. J. Mar. Freshw. Res. 16: 33-128. GUINEE, L.N. AND PAYNE, K.B. 1988. Rhyme- like repetitions in songs of Humpback Whales. Ethology 79; 295-306. PATERSON, R. AND PATERSON, P. 1989. The Status of the recovering stock of Humpback Whales, Megaptera novaeangliae, in east Australian waters. Biological Conservation 47: 33-48. PAYNE, R.S. AND McVAY, 1971. Songs of Humpback Whales. Science 173: 585-597. PAYNE, K. AND PAYNE, R. 1985. Large scale changes over 19 years in songs of Humpback Whales in Bermuda. Z. Tierpsychol. J. Com- parative Ethology 68:89-114. 332 MEMOIRS OF THE QUEENSLAND MUSEUM TYACK, P. 1981. Interactions between singing population comparisons. Behav. Ecol. Humpback Whales and conspecifics nearby. Sociobiol. 8: 41-46. Behav. Ecol. Sociobiol. 8: 105-116. WINN, H.E. AND WINN, L.K. 1978. The song of the WINN, H.E., THOMPSON, T.J., CUMMINGS, Humpback Whale, Megaptera novaeangliae, in W.C., HAIN, J., HAYS, H. AND STEINER, the West Indies. Marine Biology 47: 97-114. W.W. 1981. Song of the Humpback Whale — MEMOIRS OF THE (QUEENSLAND MUSEUM BRISBANE © Queensland Museum PO Box 3300, South Brisbane 4101, Australia Phone 06 7 3840 7555 Fax 06 7 3846 1226 Email qmlib@qm.qld.gov.au Website www.qm.qld.gov.au National Library of Australia card number ISSN 0079-8835 NOTE Papers published in this volume and in all previous volumes of the Afemoirs of the Queensland Museum maybe teproduced for scientific research, individual study or other educational purposes. Properly acknowledged quotations may be made but queries regarding the republication of any papers should be addressed to the Editor in Chief. Copies of the journal can be purchased from the Queensland Museum Shop. A Guide to Authors is displayed at the Queensland Museum web site A Queensland Government Project Typeset at the Queensland Museum THE MIGRATION OF HUMPBACK WHALES MEGAPTERA NOVAEANGLIAE IN EAST AUSTRALIAN WATERS R, A. PATERSON Paterson, R.A. 1991 0701: The migration of Humpback Whales. Weeaplera novavangliae, in east Australian waters. Mentors of the Queensland Museum 32); 333-341. Brisbane. ISSN 0079-8835. The east Australian Humpback Whale, Megeplera novaeangliae, stock, severely depleted by 1962. is now recovering atapproximately 9.7% perannum. Available evidence indicates that Shotebased whaling did not affect migration patterns in east Australia, The stack disperses in the sheltered waters of the Great Barrier Reet to breed although some calving occurs al higher latitudes, Fulure studies may determine if the breeding area for this stock includes the Coral Sea east of the Great Barrier Reef. QWumpback Whale, Megapiera muvaoeangliae, migration, cast Australta, Robert A. Paterson, Queensland Museum, PO Box 300, South Brisbane, Queensland 4101; 8 February, 1991, Humpback whales were exploited from five locations in east Australia during this century (Table 1). After the Second World War, when the re-establishment of Australian shore stations was under consideration, fishermen and other interested persons were requested to report whale sightings (Anon., 1948). Reports from east Australia indicated large numbers of Humpback Whales at 23-27°S during July, August and September. Specific site reports in- cluded Cape Capricorn, Lady Elhott Island, Double Island Point and Stradbroke Island (Col- well, 1969). During the final phase (1952-62) of Humpback Whale exploitation in east Australia, extensive biological data, including timing of northern and southern migrations in the vicinily of 27-28°S, were obtained (Chittleborough. 1965). However, the routes and timings of migrations at lower latitudes were not deter- mined, Recoveries of “Discovery” tags indicated that Humpback Whales marked in cast Australia tended to return after feeding in Antarctic waters although small rates of interchange within Area V (130°E-170°W) occurred between east Australia and New Zealand (Dawbin, 1964) in addition to an isolated exchange from Fijian waters (17°40°S, 178°55"E), Information con- cerning movement during the northern migration in a single season was also obtained. A Humpback Whale marked at St George Head, N.S.W. (35°11°S, 150°45°E) was captured al Tangalooma (27°11'S, 153°23°E) six days later (Robins, 1955) and another marked in the Cook Strait. New Zealand (41°21°S, 174°25°E) was captured at Tangalooma twenty days later (Daw- bin, 1964), In addition to movements within Area V a small number of interchanges with Area IV (70°E—130°E) were demonstrated (Chit- tleborough, 1965). When the Southern Hemisphere humpback whaling industry collapsed in 1962-63 the Area V population was estimated to be <5% of the original population of J0,000 (Chittle- borough, 1965; Chapman, 1974). During a whale sighting cruise through the known (Omura, 1953) Area V feeding grounds between January and March 1973 concern was expressed because only seven Humpback Whales were sighted (Machida, 1974). It was not until the early 1980s thal the first tentative accounts of recovery in the east Australian Humpback Whale population were reported (Paterson, 1980, 1981). In the post-whaling era Humpback Whale migrations in east Australia have been assessed by direct observation (Bryden,1985:; Paterson and Paterson,1984,1989; Simmons and Marsh,1986) and photo-identification of in- dividuals (Kaufman, Smultea and Fores- tell, 1987). The results of shore-based surveys at Point Lookout (27°26'S, 153°33°E) are dis- cussed in this paper together with reports. from other locations in east Australia. POINT LOOKOUT SURVEY Point Lookout is the northeastern headland of North Stradbroke Island and was within the operational range of Tangalooma whalechasers os TABLE 1. East Australian Humpback Whale exploita LOCALITY PERIOD Twofold Bay until 1929 37°05'S; 149°54°E Jervis Bay 1912-1913 35°03'S; 150°45°E Norfolk Island 29°01°S; 167°58°E Byron Bay 28°37'S; 153°38"E Tangalooma 2771S. 153929 E until 1927 1956-14962 1954-1962 1952-1962 (Fig.1). Sightings of Humpback Whales made on a weekly basis from a 67m high position during 1987-90 are shown in Figs 2,3 and 4. Observation methods were as described by Paterson and Paterson (1984, 1989). The peak of the northern migration occurred between late June and late July respectively in those years. In the same vicinity in 1961 the peak occurred between the last two weeks of June and the first week of July (Chittleborough, 1965). Dawbin (1956) suggested that undetermined Southern Ocean climatic factors may affect the onset of the northern migration with resultant variations in the timing of the annual peaks. The sighting tates of northbound Humpback Whales passing the east Australian shore-stations between June TABLE 2: Northbound Humpback Whales seen per 100 hours at 27°S, on the east Australian coast in 1961-1962 and 1987-1990. a nn PERIOD RELATION TO SIGHTINGS WHALING (100 HRS OPERATIONS penullimate whaling year 12/6/1961- 6/8/1961 18/6/1962- final whaling year 5/8/1962 15/6/1987- | 25 years post-whaling 65.3 5/8/1987 11/6/1988- | 26 years post-whaling 70.7 3/8/1988 11/6/1989- 3/8/1989 11/6/90- 5/8/90 27 years post-whaling 28 years post-whaling MEMOIRS OF THE QUEENSLAND MUSEUM tion since 1900. CATCHES Numbers not recorded REFERENCE Davidson, 1988 small relic industr 7400-600 12,500 barrels of oil Numbers not recorded 884 1,146 Dakin, 1934 Dakin, 1934 Chittleborough, 1965 Chittleborough, 1965 6,277 Chittleborough, 1965 Cape Morston Moreton Tangalooma Island South Pacific Ocean Point Lookout North Stradbroke Island QUEENSLAND NEW SOUTH WALES Byron Bay FIG. 1. Location map of former east Australian whal- ing stations (Table 1) nearest to Point Lookout, 335 HUMPBACK WHALE MIGRATION, EAST AUSTRALIA UY 5 AUGUST WH QW... J . Qg KW. OCTOBER OO Yy gs: S 4 | - 32 5 2229 5 1219 26 3 10 17 24 31 SEPTEMBER hern migration in 1990 (Fig.4). Although tantial numbers of northbound Humpback n index of Whales may have passed Point Lookout after the week of August in 1990, observations were K ice) fo oD & ion =) no} ¢ 2 : r= 3 2 = Ps 6 a Y i= i ~n - OO ®@ & © H tT ON F& a Z, Fr} S SHNOH OL Yad a = £ a2 k = BEs NaaS S31VHM MOVadWNH JO YaSWNN i 5 o Sate > = ' u S WH; 2 % sgoeeees 3 N 3S 4B eRe ROIS 8F g N é > |] 8S3-S8 45 z = a & U5 = we cr 3 ra ZL 2s wo * 5) Can bal i ABR aAY E pe F207" 2 + Sen OS > % 20% wiS8z 3 NS 2 agbearee? i \ = VYeEszereso a 2 wD Br wy MES 5% SBD BOYES 9% i) cS) Aglto4tce a “eee beso y © 2 SSenagens ph! 4 Zs > pa} 2 gSz2SSE3 oo OF OH TM N fF A £ Pees” se au E <8 ERs. oo ON OHM TOA a SYNOH OL HAd > 5 ae ZCAGS fe} 4 Ww = = >a 2 GALHDIS SS1VHM MOVEdANH 4O YASANN j BOT RS SRR SHNOH OL YAd Fa 4 N SeQSed Br NaS SS1VHM MOVEdWNH JO HaSWNN x WW oO wa8o 8S tose Ww Ww = EEA SO BYU > = he Seo eB oron 1988 FIG. 3. Humpback Whale sightings from Point Lookout, per 10 hrs of observation, during 1988 and 1989 336 14 Pea) eo ot2 no a 1 4 z,'° 9 22 ee ge” zg 6 xmra 5 uw o 4 i uw 6 = + z { Kg WEEK ENDED 16 23 30 7 14 2128 4 JUNE JULY AUGUST 1990 FIG. 4. Humpback Whale sightings from Point Lookout, per 10 hrs of observations, during 1990, terminated then to conform with the data collee- tions in 1988-89 (Fig.3). The extended observations. in 1987 confirm the findings of Chittleborough (1965) that the northern migration continues at Jeast until late October, albeit at a reduced rate. In 1987, 15.5% of northbound Humpback Whales passed Point Lookout between late August and October. The Antarctic pack ice does not teach its northern limit until November (Slijper, 1962) and presumably “late” northbound Humpback Whales obtain adequate nutrition because those observed at close quarters off Point Lookout in October 1982 were in prime condition (Pater- son, 1983). The Jack of a distinct peak during the southern migration in late August, September and Oc- tober was evident in 1987 and is consistent with the findings of Chittleborough (1965) and Pater- son (1984). Small numbers of southbound Humpback Whales are sighted from Point Lookout during mid-summer including two scen on 27 January 1985. During annual surveys at Point Lookout during 1978-89 almost 1,500 Humpback Whale groups were observed, Whenever possible, their route in relation to Boat and Flat Rocks was noted (Fig.5). During the northern migration 87.5% passed cast of both racks while 3.5% passed cast of Boat Rock and west of Flat Rock, During the MEMOIRS OF THE QUEENSLAND MUSEUM 153°33'E 153°33'E FIG, 5. Maps of the courses taken by Humpback Whales when migrating northwards (upper) and southwards (lower) past Point Lookout (1978-1989), HUMPBACK WHALE MIGRATION, EAST AUSTRALIA southern migration 25% passed cast of both rocks while 47.5% passed cast of Boat Rock and west of Flat Rock, No appreciable alteration in those patterns was noled in any year during 1978-89, The different routes may result from the geographic relationship of North Stradbroke and Moreton Islands (Fig.l). Northbound Humpback Whales generally puss Point Lookout ona northnortheast course until thev reach Flat Rock when they move northwest towards Moreton [sland. Southbound whales which pass close to Moreton Island may upproach Point Lookout from the westnorthwest. They are !re- quently seen near Shap Rock (Fig.5) as they move ast towards Boat Rock before resuming a southbound course, REPORTS FROM OTHER EAST COAST LOCATIONS During 1979-86, annual requesis for informa- lion concerning Humpback Whale migratrons were published in Australian Fisheries a journal circulated inter alia to holders of Australian commercial fishing licences, The response was generous and included reports from amateur and commercial fishermen, lighthouse keepers, sur- veillance and tuna-spotting pilots and the general public. A comprehensive sighting network developed and many data resulted (Figs 6,7). A survey of this type, which relies on public cooperation, may reflect bias towards areas of high human density (Simmons and Marsh, L986) and will not result in reports of all sightings (Paterson and Paterson, 1984). Nevertheless, it established the broad patlern of Humpback Whale migration in cast Australia. The peak of the northern migration [ron the Antarclic feeding grounds occurs during June/July and includes small numbers which pass castwards through Bass Strail, Although northbound Humpback Whales generally adapt a directcourse there are records of some entering bays andestuaries between 27°S and 42°S during June/July, including the Derwen) Estuary, Tas- mania; Corio Bay, Victoria; Broken Bay, N.S.W. and Moreton Bay, Queensland, Dawbin (1964) noted when Humpback Whales reach a coastal shore they tend to remain close fo it and those findings were confirmed by Bryden (1985) during aerial surveys off the southern Queensland and northern N.S.W. coasts. However, after (he migration stream pas- ses Breaksea Spit (2436°S, 153°20°E) a the northern end of Fraser Island at enters the shel- tered waters of the Great Barrier Reef and dis- perses widely between the outer Barrier and the coast, which inclines to the northwest. Northward movement slows and sightings peak in August with only small numbers reaching latitudes lower than 15°S. The fatitudinal and monthly concentrations are similar to those recorded in other Southern Hemisphere breeding locations (Townsend, 1935). Calving has been witnessed at 18-19°S inside the Great Barrier Reef but also occurs at higher latitudes as Humpback Whales accompanied by new born calves have been scen migrating northwards along the southern Queensland coast (Paterson and Paterson, 1989). A dedicated survey. using photo-identification (Kaufman et al.,1987), may demonstrate the movements of individual Humpback Whales in the Great Barrier Recf region. Small numbers of Humpback Whales cam- mence their southern migration in late July (Figs 3,4) but the majority leave the Great Barrier Reef region between mid-August and mid-October. The migration pattern in the Fraser Island region differs from the northern migration in that large numbers of Humpback Whales pass west of Breaksca Spit and enter Hervey Bay, This phenamenon has resulted in the development of a large whale-watching industry in recent years as the public has the opportunity to observe Humpback Whales at close quarters in sheltered wulers af! a predictable time of the year. The geographic relationship of Breaksea Spit and Fraser Island to the mainland may explain in part why Humpback Whales enter Hervey Bay. A similar, although less dramatic, course variation during the southern migration compared with the northern migration in the Point Lookout region (Fig. 5) has already been discussed. By November/December most Humpback Whales have left the east Australian coast for the Antarctic feeding grounds although small num- bers have been reported at relatively low latitudes in January/February (Paterson, 1987), Dawbin (1966) noted that southbound Humpback Whales captured at Great Barrier Island, New Zealand (36°S) invariably showed evidence of recent feeding and there is evidence to suggest occasional Humpback Whale feeding activity at similar latitudes on the cast Australian coast (Paterson,] 487; Ellis, 1989), DISCUSSION The combination of legal and illegal catches of 338 MEMOIRS OF THE QUEENSLAND MUSEUM 10°S 15°s 20°s 25°s 30°s 35°s 40°s 20°s 25°s 30°s 35°S 40°s FIG. 6. Spatial distribution of Humpback Whales in east Australian waters, May-August, 1978-1989. Each marking represents one or more Humpback Whale Groups seen during a particular month in one or more years. HUMPBACK WHALE MIGRATION, EAST AUSTRALIA 339 10°s 15°s 20°s 25°S October September 30°5 35°s 40°s 20°s 25°S November December 30°s 35°S 40°s 155°E FIG. 7. Spatial distribution of Humpback Whales in east Australian waters, September-December, 1978-1989. Each marking represents one or more Humpback Whale Groups seen during a particular month in one or more years. 340 Area V Humpback Whales during 1952-63 resulted in a population decline extreme even by the standards of modern whaling (Tonnessenand Johnsen, 1982). The paucity of incidental sight- ings in the post-whaling decade prompted con- jecture that Humpback Whales deliberately avoided the regions of the former east Australian shore-stations (Anon., 1976), However, similarity of the migration patterns for 1961 (Chittleborough,1965) and 1987 (Fig.2) sug- gests that exploitation of Humpback Whales. in east Australia did not affect their migratory be- haviour. Although the sheltered waters of the Great Barrier Reef are 4 major destination for those Humpback Whales which migrate northwards along the east Australian coast, it has not been established that those waters are the sole destina- tion. Dawbin and Falla (1949) suggested that the Chesterfield Reefs, northwest of New Culedonia in the eastern Coral Sea, were the destination of the cast Australian stock as Humpback Whales had been captured there in the nineteenth century (Townsend, 1935), However, the “Discovery” tag program failed to establish evidence of such a pathway and the present situation with regurd to Humpback Whale migration in the Coral Sea east of the Great Barrier Reef remains un- resolved. Paterson and Paterson (1989) considered that the east Australian Humpback Whale stock was increasing at approximately 9.7% per annum on the basis of annual surveys at Point Lookout during 1981-87. The 1987/88 and 1988/89 data were consistent with that estimate but the low increase demonstrated in 1989/90 was almost certainly due to the “late” northern migration in 1990 (Fig.4). Such a variation notwithstanding, the evidence from Table 2 indicates that the east Australian portion of the Area V stock has doubled since the cessation of whaling in 1962 and now exceeds 10% of the pre~ exploitation estimate of 10,000. ACKNOWLEDGEMENTS Particular thanks are due to Martin Bowerman, formerly editor of Ausfralian Fisheries, who in- stigated the annual whale sighting appeal in 1979 and to the officers of the Great Barrier Reef Marine Park Authority who collated data ob- tained by Coastwatch surveillance aircrews. Numerous persons from many locations on the east Australian coast generously forwarded reports which enhanced this study. MEMOIRS OF THE QUEENSLAND MUSEUM LITERATURE CITED ANONYMOUS 1948, Whaling is big business, Fisheries Newsletter. 7(2): 2-3. ANONYMOUS 1976. Moreton Island environment impact study and strategic plan. Co-ordinator General's Department, Brisbane, Queensland, 104p. BRYDEN, M.M. 1985. Studies of Humpback Whales (Megaptera novaeangliae), Area V. 115-123. 1n J.K. Ling and M.M, Bryden, (eds), ‘Studies of sea mammals in south latitudes’. (South Australian Museum: Adelaide). CHAPMAN, D.G, 1974, Status of Antarctic rorqual stocks. 218-238, In WE, Schevill, (ed,), "The Whale problem, a Status report’. (Harvard University Press: Cambridge, Mass. ). CHITTLEBOROUGH. R.G, 1965. Dynamics of two populations. of the Humpback Whale, Megap- tera nuvueaneliae (Borowski), Aust. J. Mar. Freshw, Res, |G: 33-128. COLWELL, M. 1969. ‘Whaling around Australia’, (Rigby: Adelaide) |Oxp, DARIN, W.J. 1934.‘ Whalemen adventurers’, (Angus & Robertson: Sydney), 263p. DAVIDSON, R. 1988. “Whalemen of Twofold Bay’, (René Davidson; Eden), 1 72p, DAWBIN, W.H, 1956, The migrations of Humpback Whales which pass the New Zealand coast, Trans. R. Soc, N.Z. 84: 147-196. DAWBIN, W.H, 1964. Movements. of Humpback Whales marked in the South West Pacific Ocean 1952 to 1962, Norsk Hvalfangst-Tidende 3: 64— 78. DAWBIN, W.H. 1966. The seasonal migratory cycle of Humpback Whales. 145-170. [n K.S. Norris, (ed.), “Whales, dolphins and porpoises”, (Univ, California Press: Berkeley and Los Angeles). DAWBIN, W.H. AND FALLA, R.A. 1949, A con- tribution to the study of the Humpback Whale based on observations at New Zealand shore stations. Proc. 7th Pacific Sci. Congr. 4: 373— 382. ELLIS, R. 1989. Whales down under, Australian Geographic 16; 59-79, KAUFMAN, G.D., SMULTEA, M.A. AND FORES- TELL, P. 1987. Use of lateral body pigmentation patterns for photographic identification of East Australian (Area V) Humpback Whales, Cetus 7 (1): 5-13, MACHIDA, S. 1974. The voyage ol the Konan Maru No 16 to the Antarctic whaling grounds. Sei. Repts Whales Res. Inst. 26: 289-302, HUMPBACK WHALE MIGRATION, EAST AUSTRALIA OMURA, H. 1953. Biological study on the Humpback Whales in the Antarctic whaling Areas IV and V. Sci. Repts Whales Res. Inst. 8: 81-102. PATERSON, R. 1980. Encouraging sightings of Humpback Whales off east coast. Australian Fisheries 39(4): 8-10, PATERSON, R. 1981. ‘Whale watch’ brings en- couraging reports. Australian Fisheries 40(4): 30-33. PATERSON, R. 1983, ‘Whale watch’ brings more encouraging reports. Australian Fisheries 42(4): 32-33, PATERSON, R. 1984. Migration patterns of Humpback Whales (Megaptera novaeangliae) in the waters adjacent to Moreton and North Stradbroke Islands. 342-347. In R.J. Coleman, J. Covacevich and P. Davie, (eds), “Focus on Stradbroke’. (Boolarong, Brisbane). PATERSON, R. 1987. Recovery in the East Australian Humpback Whale stock. Australian Fisheries 46(8): 32-36. PATERSON, R. AND PATERSON, P. 1984. A study 341 of the past and present status of Humpback Whales in East Australian waters. Biological Conservation 29: 321-343. PATERSON, R. AND PATERSON, P. 1989. The Status of the recovering stock of Humpback Whales Megaptera novaeangliae in East Australian waters. Biological Conservation 47: 33-48. ROBINS, J.P. 1955. Whale marking and its uses . Fisheries Newsletter 14(4): 15,19. SIMMONS, M.L. AND MARSH, H. 1986, Sightings of Humpback Whales in Great Barrier Reef waters. Sci. Repts Whales Res. Inst. 37: 31-46. SLIJPER, E.J. 1962. ‘Whales’. (Hutchinson:London). 475p. T@NNESSEN, J.N. AND JOHNSEN, A.O. 1982. ‘The history of modern whaling’. (Hurst and ANU Press: London and Canberra), 798p. TOWNSEND, C.H. 1935. The distribution of certain whales as shown by log book records of American whaleships. Zoologica 19: 1-50. MEMOIRS OF THE (QUEENSLAND MUSEUM BRISBANE © Queensland Museum PO Box 3300, South Brisbane 4101, Australia Phone 06 7 3840 7555 Fax 06 7 3846 1226 Email qmlib@qm.qld.gov.au Website www.qm.qld.gov.au National Library of Australia card number ISSN 0079-8835 NOTE Papers published in this volume and in all previous volumes of the Afemoirs of the Queensland Museum maybe teproduced for scientific research, individual study or other educational purposes. Properly acknowledged quotations may be made but queries regarding the republication of any papers should be addressed to the Editor in Chief. Copies of the journal can be purchased from the Queensland Museum Shop. A Guide to Authors is displayed at the Queensland Museum web site A Queensland Government Project Typeset at the Queensland Museum STUDIES OF TWO HUMPBACK WHALES. MEGAPTERA NOVAEANGLIAE, STRANDED AT FRASER ISLAND, QUEENSLAND R.A. PATERSON AND S. VAN DYCK Paterson, R.A. and Van Dyck, S., 1991 07 O1: Studies of wo Humpback Whales, Meguplera novaeangltae, stranded at Praser tsland, Queensland, Memoirs of the Queensland Museum 32); 343-350, Brisbane, ISSN 0079-8835, In 1989 two Humpback Whales, Megapicra novaeangliae, stranded at Fraser Island, Queensland, One specimen wis an &.lm female yearling and the other a 4.2m male new-born calf. Skeletal material. soft tissue, baleen, blubber and externul parasites were recovered, Histological examinations were made of tissues from the heart, lungs, liver, kidneys and frontal bone (call only). The cause of death in the yearling was not determined whereas the call probably died from shark attack. [>] Humpback Whale, Megaplera novacangliae, srandings, Fraser Island, Queensland, Robert A. Paterson and Stephen Van Dyck, Queensland Museum, PO Box 300, Sauth Brisbane, Queensland 4/71; & February, 1991. Although Humpback Whales migrate close to Southern Hemisphere continental shores (Daw- bin, 1966), records of strandings are relatively uncommon (Bannister, 1989). Prior to 1989 the Queensland Museum (QM) had records of five strandings (Paterson, 1986) but the deposited nvaterial was fragmentary apart from a skull, the collection of which was described in detail by Welsby (1931). Material recovered from two Humpback Whales stranded at Fraser Island (Fig. 1) in 1989 Was a significant addition to the QM collection, The specimens are registered JM7302 and IM7303 in the Queensland Museum Mammal Department. SPECIMEN DESCRIPTIONS FEMALE YEARLING, JM7302 was 8.1m long, indicating a yearling, and stranded during good weather conditions near Brown's Rocks (24°46'S, 153°16°E) on 3 July 1989. Eye-witness accounts indicated that the animal was alive in the surf-zone but dead when washed ashore. When examined on the day following stranding, the right dorsolateral aspect of the carcass was buried due to tidal action (Fig. 2). Apart from superficial shark bites on the flukes and caudal peduncle there was no cxternal evidence of injury. Barnacles, Coronula diadema and Conchoderma auritum, were noted particularly on the protuberance beneath the chin and on the pectoral fins, flukes and genital aper- ture (Figs 3,4). The majority of C, aurtium were attached to C- diadema. Cyamids were not iden- tified. The head and baleen were removed and blubber samples taken. The gastro-intestinal tract was empty. When the site was revisited on 2 September 1989 it was discovered that the carcass had been washed 7km further north during stormy weather and the majority of the cervical and lumbar ver- tebrae were lost, MALE CALF, JM7303 was 4.2m long and stranded at Moan Point (25°14'S, 153°00°E) on 17 October 1989, Eyc-witness accounts indicated that a larger (adult) Humpback Whale was present in deep water adjacent to Moon Point earlier on the same day while a calf, presumably the subsequently stranded animal, was seen swimming steadily in shallow watcr clase to the shore. The actual stranding was not witnessed, When discovered the animal was bleeding from a large right axil- lary wound and died soon after, The carcass was recovered some hours later, takcn to Urangan on the mainland where it was frozen, and sub- sequently transported to the OM, It had suffered numerous shark bites including a large, although healing, ventral wound, The humerus and large severed vessels were evident in the base of the previously noted axillary wound (Fig. 5). There were 24 ventral grooves and no evidence of external parasites. The specimen was cast and moulded for permanent display in the Queensland Museum. The larynx was removed for separate examina- tion (Quayle, this memoir), The pleural and peritoneal cavities appeared normal. The prin- 344 MEMOIRS OF THE QUEENSLAND MUSEUM \ + 25°8 \ | / Locality Map \ / wmnelanit i } ie Bay FIG. 1. Location of stranding sites on Fraser Island . cipal gastric content was sand, possibly ingested during stranding. Faecal material was present in the intestines. Histological examination of the heart, lungs, liver and kidneys was normal apart from evidence of terminal pulmonary oedema. There were 15 pairs of ribs and the vertebral formula was C7; T15; L11; Cd 18 =51. Healing fractures involving the anterior aspects of the larger ribs were noted and the radiological find- ings (Fig. 6) suggest that the injuries had been sustained approximately four weeks previously. in addition a histologically confirmed post- traumatic pericranial reaction of similar duration to the rib fractures was noted on the supero- lateral aspect of the supraorbital process of the right frontal bone (Fig. 7). Similar pericranial injuries may occur in humans during birth and are termed cephalhaematomata (Caffey, 1973). There is no evidence that they are associated with mortality or persistent morbidity in humans and similarly the skeletal injuries suffered by JM7303 are considered to be unassociated with its death. While post-natal skeletal trauma is not entirely excluded there was no evidence of over- lying cutaneous or subcutaneous injury. FIG. 2. Partly buried carcass of JM7302 near Brown's Rocks. STRANDED HUMPBACK WHALES, FRASER ISLAND 345 Ae ™~), we aap tae, ot a 7. FIG. 4. Barnacles, Coronula diadema and Conchoderma auritum adjacent to the genital aperture of JM7302. 346 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 5, Humpback Whale calf, JM7303, showing large right axillary wound. FIG. 6. Radiograph of anterior aspects of the left 8th and 9th ribs of JM7303 showing healing fractures. STRANDED HUMPBACK WHALES, FRASER ISLAND 347 FIG. 7. Skull of JM7303 showing post-traumatic reaction to the supero-lateral aspect of the supra-orbital process of the right frontal bone. 348 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 8. Baleen of JM7302 showing band of white plates anteriorly. BLUBBER The mid-dorsal blubber thicknesses of JM7302 and JM7303 were 9.7cm and 4.5cm, respectively and are consistent with normal nutrition for Humpback Whales of those lengths (Matthews, 1937). It is assumed that JM7302 was northbound from the Antarctic and had been recently weaned. Accordingly its blubber would be thicker than that of JM7303 which was new- born and in the early stages of the southern migration. BALEEN The baleen bristles were greyish-white in both specimens. The plates were black with the ex- ception of a few that were pale, almost white, anteriorly in JM7302 (Fig. 8). They numbered 342 and 310+5 on the left in JM7302 and JM7303 respectively (Figs 9,10). Doubt con- cerning the exact count in JM7303 resulted from the extremely small size of the anterior plates. The largest plates were 46cm and 9.5cm long in JM7302 and JM7303, respectively. DISCUSSION Given the large size of some cetaceans and the unpredictable nature of their stranding it is not surprising that recovery of useful biological material is often limited. The study of JM7302 typified those problems. Although first ex- amined on the day following stranding, its loca- tion and available resources dictated the amount of material that could be recovered. The Queensland Museum was fortunate in the case of JM7303 in that prompt retrieval and freezing of the latter by a local fisherman provided a rare opportunity to examine and prepare an excellent- ly preserved specimen. Sheltered waters of the Great Barrier Reef are an important calving area for the east Australian Humpback Whale stock (Paterson and Paterson, 1984,1989; Simmons and Marsh,1986). Most Humpback Whales pass northwards along the southern Queensland coast between June and early August and pass southwards between late August and October. In the vicinity of Fraser Island they pass the eastern (oceanic) shore during both migrations but enter Hervey Bay on STRANDED HUMPBACK WHALES, FRASER ISLAND 349 FIG. 9, Lingual and buccal views of left baleen row of JM7303. the western shore in large numbers only during the southern migration (Paterson and Paterson, 1989). The location and timing of the strandings of JM7302 and JM7303 are consistent with those migration patterns, ACKNOWLEDGEMENTS We are indebted to many people who assisted with the retrieval and preparation of the specimens. JM7302 was examined and collected in accordance with a permit issued by the Mini- ster for Primary Industries. Shamus Conway and Merv Toms of QNPWS, Wayne Kelly of Sandy Cape Lightstation, Ron Marshall, Ben Byth and Tom Paterson assisted with the retrieval of JM7302 from Fraser Island. Ken Thomas and the staff at the Brisbane City Council's Luggage Point Sewerage Station allowed us to prepare skeletal material at the Station. We are most grateful to Vic Hislop who retrieved JM7303 and transported it to Brisbane after arranging its freezing. Bruce and Carolyn Cowell, OM, patiently ap- plied their respective photographic and prepara- tion skills. X-rays were taken by Sophie Kupis, Jindalee Medical Centre and John Musgrave of Sullivan, Nicolaides and Partners, kindly ar- ranged the preparation and histological ex- FIG, 10, Lingual and buccal views of right baleen row of IM7302. amination of numerous tissue samples. Peter Davie, QM, identified the barnacles. LITERATURE CITED BANNISTER, J.L. 1989. Balaenopteridae. 982-987, In D.W. Wallon and B.J. Richardson, (eds), ‘Fauna of Australia. Vol. 1B*. (Aust. Govt Publ. Service; Canberra). CAFFEY, J. 1973_ ‘Paediatric X-ray diagnosis. Vol. 1’. 6th ed. (Year Book Medical Publs: New York), 714p. DAWBIN, W.H. 1966. The seasonal migratory cycle of Humpback Whales. 145-170. In K.S. Norris, (ed.), ‘Whales, dolphins and porpoises’. (Univ, Calif. Press: Berkeley and Los Angeles). MATTHEWS, L.H. 1937. The Humpback Whale - Megaptera nodosa, Discovery Reports 17: 7— 92. PATERSON, R.A. 1986. A list of specimens of the Order Cetacea in the Queensland Museum. Mem. Qd Mus. 22(2): 309-311. PATERSON, R. and PATERSON, P. 1984. A study of the past and present status of Humpback Whales in east Australian waters. Biological Conservation 29: 321-343. PATERSON, R. and PATERSON, P. 1989, The status of the recovering stock of Humpback Whales 350 MEMOIRS OF THE QUEENSLAND MUSEUM Megaptera novaeangliae in east Australian humpback whales in Great Barrier Reef waters. waters. Biological Conservation 47: 33-48. Sci. Repts Whales Res. Inst. 37: 31-46. SIMMONS, M.L. and MARSH, H. 1986. Sightingsof WELSBY, T. 1931. ‘Sport and pastime in Moreton Bay’. (Simpson Halligan & Co.: Brisbane). MEMOIRS OF THE (QUEENSLAND MUSEUM BRISBANE © Queensland Museum PO Box 3300, South Brisbane 4101, Australia Phone 06 7 3840 7555 Fax 06 7 3846 1226 Email qmlib@qm.qld.gov.au Website www.qm.qld.gov.au National Library of Australia card number ISSN 0079-8835 NOTE Papers published in this volume and in all previous volumes of the Afemoirs of the Queensland Museum maybe teproduced for scientific research, individual study or other educational purposes. Properly acknowledged quotations may be made but queries regarding the republication of any papers should be addressed to the Editor in Chief. Copies of the journal can be purchased from the Queensland Museum Shop. A Guide to Authors is displayed at the Queensland Museum web site A Queensland Government Project Typeset at the Queensland Museum A DISSECTION OF THE LARYNX OF A HUMPBACK WHALE CALF WITH A REVIEW OF ITS FUNCTIONAL MORPHOLOGY CJ, QUAYLE Quayle, C.J. 199) 07 0}; A dissection of the larynx of a humpback whale calf with review of its. functional morphology, Memoirs of the Queensland Museum 30(2): 351-354. Brisbane. ISSN 0079-8835. The larynx of a humpback whale calf was dissected. The anatomy conformed with the general mammalian pallern with the addition of a ventral diverticulum. Some thoughts as to how the organ might function ure considered. [7] Humpback whale, larynx, anatomy, funetion, C.J, Quayle, 79 Wickhum Terrace, Brisbane 4000; 20 December 1990, A humpback whale calf (QMJM7303) stranded at Moon Point, Fraser Island, in Qe- wber, 1989 was frozen soon after death and delivered to the Queensland Museum, Brisbane. I presented an almost unique opportunity to examine a larynx of manageable size with min- imal postmortem changes. The calf was 4.2 m long and estimated to be about four weeks old. The respiratory tract began with two blowholes, slits about 5em long on cach side of the midline and inclined so that their rostral ends Were closest together, The blowholes were lined with black skin which continued to the base of the skull where it became continuous with the mucous membrane. The blowholes were surrounded by fibro-fatty tissue without obvious musculature, From their external entrance the blowholes ran almost horizontally caudally in bony grooves ending as the bony posterior choanae, The vomerine bone of the skull here separated the airway into two channels, The air passage then became a single thin-walled tube which continued, inclined ventrally, to meet the thick-walled muscular oropharynx at an acute angle. This arrangement suggests a valve. This length of tube appeared to be the equivalent of the human nasopharynx. The larynx itself was horizontally disposed. It measured 20cm from the tip of the epiglattis to the first tracheal ring. Compared with its human counterpart it was in close proximity to the skull base with the tip of the epiglottis extending al- most to the posterior choana within the conduit, ! have called, the nasopharynx. In this respect clongation of the human pharynyes is said to be the result of a vertical stance. The food passage passed on each side of the laryngeal opening as two pyriform fossae ta become a single tube, the oesophagus, dorsal to the trachea. The larynx was a tubular organ with a car- tilaginous skeleton. There were three unpaired cartilages, the epiglottic, thyroid and cricosd, and one pair of arytenoid cartilages. Caudally the larynx became continuous with the trachea, The tube was lined with what appeared to be squamous epithelium, This was tough and close- ly applied particularly over the arytenoid car- tilages (compare human vocal cords). The cartilages were joined by muscles most of which could be identified as following the general mammalian pattern. The thyroid cartilage consisted of two alae or plates continuous in the ventral midline and widely deficient dorsally, There was a small rostral cornua and a long caudal one equivalent to the greater cornua of the human thyroid car- tilage. The latter articulated with the cricoid car- tilage on cach side. There was a flat ovoid facet contiguous on each cartilage and the joint had a thick fibrous capsular membrane. The cricoid' cartilage was complete dorsally where it formed a large rectangular plate 18x12em. It was deficient ventrally (Fig.1), The dorsal plate merged with the upper tracheal rings forming a substantial crico-tracheal plate. Caudally and laterally was the facet for articula- tion with the greater cornu of the thyroid car- tilage. On cach rostral shoulder was a’ large synovial joint articulating with its respective arytenoid cartilage. The lateral plates of the cricoid were narrow rostrally and broad caudal- ly. Within the ventral deficiency lay the ventral diverticulum, A very thick band of muscle, lar- gely transversally disposed (the thyroarytenoid muscle) filled the space between the lateral alac and covered the diverticulum, Each arytenoid was complex in shape ar- ticulating with the cricoid cartilage as described, we ws te MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 1. Longitudinal section of Humpback Whale larynx (JM7303) in medial aspect (thyroid cartilage removed). (A) epiglotiic cartilage: (B) fibrous allachment to thyroid cartilage; (C) arytenoid cartilage - corpus; (D) arytenoid cartilage - rostral end; (E) cricoid cartilage; (F) tracheal Jumen; (G) fundus of the ventral diverticulum; (H) neck of the ventral diverticulum: (1) Lhyroarylenoid muscle; (J) interarytenoid fibro elastic connection. The body of the arytenoid lay largely within the arch of the cricoid. A short thick process cx- tended from the cricoarytenoid articulation to join the body of the cartilage in its middle third, The body was a banana shaped mass 15cm long with the rostral and the caudal extensions. Caudal processes were connected in the midline by a thick fibro-elastic band. The beak shaped rostral processes were concave ventrally and together were almost embraced by the epiglottis. The epiglottis together with the rostral processes of the arytenoids and the aryepiglottic folds formed the arytenoepiglottideal tube. The medial borders of the corpus were almost flat and covered with tough skin. They approximated in the midline. Some accounts refer to purt of this body as the cunciform cartilage (Hosokawa, 1950:28). A partial cleft in this specimen sug- gests that the cuneiform and arytenoid cartilages may be fused, the rostral process representing the cuneiform cartilage. The epiglottis was 15 cm long and roughly spoon-shaped concave dorsally. It was relatively elastic and attached by a thick fibrous band to the thyroid cartilage near its cranial notch. The in- tegument on the luminal side had thick lon- gitudinal grooves which seemed almost to interdigitate with those on the adjacent surface of the arytenoid. The trachea was comparatively short being approximately 4cm long from the cricoid plate to the bifurcation. Dorsally the proximal car- tilages were not fused but “jumbled”. The car- tilages. were deficient ventrally where they had rounded ends. The ventral diverticulum was recessed into the defect between these ends, the tracheal Jumen being effectively divided into twa passages, A pouch (ventral diverticulum) opened into the laryngeal lumen by a longitudinal slit on its floor between the medial borders of the arytenoid cartilages. There is no human counter- part. It was 10cm Jong and was 3 cm in diameter al the fundus. The wall was fibrous and appeared nondistensible. There were some shallow pock- ets in the fundus and the whale was covered by a thick muscular mass. The fundus invaginated into the tracheal lumen on its ventral side and appeared to all but obliterate that lumen. INTERPRETATION 1, the aryteno epiglottideal tube appeared to form a conduil to carry the air stream from the posterior choanae to the laryngeal inlet. 2, the airstream appeared to be directed into the ventral diverticulum (Fig. 2), The opening into the diverticulum may well be closed during normal inspiration. 3, air passing into the trachea must pass be- tween the flattened margins of the bodies of the adjacent arytenoid cartilages. 4, the ventral diverticulum seemed unlikely to HUMPBACK WHALE LARYNX 353 FIG. 2. Longitudinal section of Humpback Whale larynx (JM7303) in medial aspect to show course of air stream into trachea. be distensible and as a reservoir would have been of relatively small size. 5, air expelled from the diverticulum would apparently pass over the free margins of the arytenoids and through the arytenoepiglottideal tube into the nasopharynx. 6, the ventral diverticulum could apparently function as a valve able to isolate the supraglottic air space from that in the thorax. 7, among cetaceans ventral diverticula have been described only in baleen whales (Slij- per, 1962:147). 8, baleen whales produce sounds organised into a complex form and distinct from the repeti- FIG. 3. Schematic drawing laryngeal inlet (i) humpback whale; (ii) human. (A) epiglottis; (B) arytenoid; (C) glottic chink. 354 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 4. Human larynx (displayed horizontally). Longitudinal section in medial aspect. (A) epiglottic cartilage; (B)aryteno-epiglottideal “tube”; (C) cricoid cartilage; (D) tracheal rings; (E) tracheal lumen. tive clicking sounds produced by toothed whales (Schevill,1964:308). FUNCTION As in the human the larynx is almost certainly a valve to stop food and water going down into the trachea. In this respect it may have an added burden as the whale forces enormous amounts of water from its mouth through the baleen plates during feeding. The larynx seems too complicated a structure to be a simple valve and the remarkable similarity between the human and whale glottic inlets (Figs 3,4) suggests that it is morpholog- ically capable of phonation. There are no vocal cords as such, but phonation could occur be- tween the apposing arytenoids. In physical terms an air cavity with fluctuating volume is an efficient source of sound in water. Perhaps forcing air from the ventral diverticulum between the adducted arytenoids would make the air column both within the diverticulum and the nasopharynx vibrate. The concept of a whale singing with sound bubbles coming out of its mouth is almost cer- tainly incorrect. The similar physical charac- teristics (i.e.acoustic impedances) of the soft tissue of the whale and the surrounding sea water would mean that sound generated would pass through the body of the whale into the surround- ing water in all directions with little attenuation. I postulate that air forced from the ventral diverticulum between the arytenoids causes the air column in the diverticulum and perhaps the nasopharynx to vibrate. The resulting pressure fluctuations are transmitted through the soft tis- sues of the whale into the surrounding water. The air stream does not transmit the sound waves but generates pressure fluctuations in the column of air which bear on the enclosing tissues and pass through the body into the water. The same muscle action would compress the diverticulum and adduct the arytenoids. Al- though the volume of air in the diverticulum is small in this specimen (which may not have been vocal) it is known that the diverticulum may be large in an adult whale. This would provide sufficient air for the relatively short bursts of phonation characteristic of the humpback whale song. While the diverticulum was compressed the trachea would be occluded and gas exchange could presumably continue uninterrupted in the lungs. Conceivably the diverticulum could be refilled from the thoracic air while the whale remained submerged. ACKNOWLEDGEMENTS I thank Ms Sally Elmer of the Queensland Museum for preparing the drawings, and Dr Douglas H. Cato of the Materials Research Laboratory, Department of Defence, for advice on the physics of sound transmission. LITERATURE CITED HOSOKAWA, H., 1950. On the cetacean larynx, with special remarks on the laryngeal sack of the sei whale and the aryteno-epiglottideal of the sperm whale.Sci. Repts Whale Res. Inst. 3:23- 62. SCHEVILL, W.E., 1964. Marine bio-acoustics (Per- gamon: New York). SLIJPER, E.J., 1962. Whales (Hutchinson:London). CONTENTS CHITTLEBOROUGH, R.G. Potential inpacts of climatic change on the Southern Ocean ecosystem ................- 243 DAWBIN, W.H. AND EYRE,E.J. Humpback Whale songs along the coast of Western Australia and some comparison with CBS COASU SOMES 56 ic seat eer Be eae) Pam paren 4-4 wa velo Rain GES oh bubs we Pees 249 DAWBIN, W.H. AND GILL, P.C. Humpback Whale survey along the west coast of Australia: a comparison of visual and BCOUStiC OlmeIV ALIGNS. (acy oy te Sone es nes Sevan er epee ed ee ee ae ee 255 BANNISTER, J.L. 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An acoustic analysis of the 1988 song of the Humpback Whale, Megaptera novaeangliae, OPP SANE Use Ae TT Ope gt eee te ar, se eR cee teen ae cee ne: A eae 323 PATERSON, R.A. The migration of Humpback Whales Megaptera novaeangliae in east Australian waters ... 333 PATERSON, R.A. AND VAN DYCK S. Studies of two Humpback Whales, Megaptera novaeangliae, stranded at Fraser Island , Cueenslad ihe pv eeee ine op yee me Ts te be KERR Yea ed 343 QUAYLE, C.J. A dissection of the larynx of a Humpback Whale calf with a review of its functional (OU (21 0178s A ia a te ei Ay PO a AOL Are Pee Sie ce ae nate Br PRT ba 351