DOI: https://doi.org/10.3354/meps14937
copiedAge-specific body length, mass, and energetic cost of growth in humpback whales
- M. van Aswegen
- A. Szabo
- J. J. Currie
- S. H. Stack
- J. Straley
- J. Neilson
- C. Gabriele
- K. Cates
- C. S. Baker
- D. Steel
- A. A. Pack
- K. L. West
- N. J. Hofmann
- R. J. Toonen
- C. Loockerman
- F. A. Sullivan
- T. Cheeseman
- F. Sharpe
- L. Bejder
ABSTRACT:
Body size is a fundamental parameter in modeling the energetic trade-offs related to growth, activity, maintenance, and reproduction. This study integrated photogrammetry from an unoccupied aerial system (UAS; drone) with individual life histories, historical whaling records, and post-mortem tissue samples to estimate body mass and compare somatic growth costs in humpback whales Megaptera novaeangliae. Using 2651 UAS-derived length measurements and demographic information of 1503 individuals, sex-specific von Bertalanffy growth curves were developed from birth to 50 yr. Asymptotic lengths were estimated at 13 m for females and 12.34 m for males, with calves reaching 71-75% of their asymptotic length by weaning (~1 yr). Volume-at-age curves of whales in average body condition were converted to mass using UAS-derived body volume estimates, post-mortem tissue density samples, and published estimates of tissue composition. The cost of growth in calves and adults was modeled using tissue-specific lipid and protein concentrations derived from post-mortem tissue samples and published estimates, respectively. We found that postnatal offspring growth requires ~38 times more energy than prenatal growth, with calves needing 6-8 times more energy per day than mature adults. The cumulative cost of growth averaged 713 GJ for females and 607 GJ for males. We highlight a critical period of energetic demand for lactating females, who must support one-third of their offspring’s lifetime growth costs in just 1 yr. Estimating body mass and the cost of growth in humpback whales offers valuable insights for bioenergetic models assessing the effects of disturbance and a rapidly changing climate.
‘Ōlelo Hawai‘i: ‘O ka nui kino kekahi mea ko‘iko‘i i mea e ana ai i ka ho‘okaulike i ka mea ‘ai no ka ulu ‘ana, ka hana ‘ana, ka mālama ‘ana i ke kino, a me ka hānau a hānai ‘ana i ke keiki. Ho‘ohana kēia ‘imi noi‘i i ka pa‘i ki‘i ‘ana o ka helekopa uila li‘i a me ka mo‘olelo ola o kekahi mau koholā Megaptera novaeangliae, ‘ike kahiko ‘ō koholā, a me ka mea kino hō‘ike‘ike ‘u‘uku o nā kupapa‘u koholā no ke koho ‘ana i ke kaupaona a me ka ho‘ohālikelike ‘ana i nā koina ulu o ke kino ma ke koholā. Me 2651 mau ana lō‘ihi mai ka helekopa uila li‘i a me ka ‘ike keka o 1503 mau koholā, ua ho‘okumu ‘ia nā pi‘o ulu von Bertalanffy no nā kāne a me nā wāhine mai ka hānau ‘ia ‘ana a hiki i ka makahiki 50. ‘O 13 m ka lō‘ihi pālahalaha i koho ‘ia mai ka pi‘o no nā wāhine, a 12.34 m no nā kāne. Ulu nō nā keiki a hiki i 71-75% o ko lākou lō‘ihi pālahalaha ma mua o ka ukuhi ‘ana (ma kahi o 1 makahiki). Ua ho‘olilo ‘ia nā pi‘o nui-makahiki i ke kaupaona no nā koholā i ka momona kino ma‘amau ma o ka ho‘ohana ‘ana i nā koho o ka nui o ke kino mai ka helekopa uila li‘i, ka pa‘apū o ka mea kino hō‘ike‘ike ‘u‘uku o nā kupapa‘u koholā, a me nā koho o ke ‘ano o nā ‘i‘o like ‘ole i ho‘olaha ‘ia. Ana ‘ia ke koina ulu ma nā keiki a me nā mākua me ka nui o nā lipid a me nā protein ma nā mahele ‘i‘o ‘oko‘a mai ka mea kino hō‘ike‘ike ‘u‘uku o nā kupapa‘u koholā a me nā koho i ho‘olaha ‘ia. Hō‘ike nō mākou, ‘oi pākanakolukūmāwalu aku ke koina ulu keiki ma hope o ka hānau ‘ia ‘ana ma mua o ka hāpai ‘ia ‘ana, a ‘oi pāono-pāwalu aku ke koina mea ‘ai o nā keiki ma mua o nā mākua. ‘O 713 GJ no nā wāhine a 607 GJ no nā kāne ke koina ulu waena no 50 mau makahiki. Hō‘ike nō mākou i kēia wā nui loa o ke koina mea ‘ai no nā makuahine e hānai ana, nāna e mālama pono i ka ulu ‘ana o ka hapakolu o ke ola holo‘oko‘a o kā lākou keiki i ho‘okahi wale nō makahiki. ‘Ō‘ili mai ka ‘ike waiwai mai ke koho ‘ana i ke kaupaona kino a me ke koina ulu no nā ana koina mea ‘ai e kilo i nā hopena o nā pilikia a me nā loli ‘āwīwī o ke ‘ano o ka honua.
KEYWORDS
Adamczak SK, McHuron EA, Christiansen F, Dunkin R and others (2023) Growth in marine mammals: a review of growth patterns, composition and energy investment. Conserv Physiol 11:coad035 PubMed Crossref PubMed Crossref
Agbayani S, Fortune SME, Trites AW (2020) Growth and development of North Pacific gray whales (Eschrichtius robustus). J Mammal 101:742-754
Akaike H (1974) A new look at the statistical model identification. IEEE Trans Automat Contr 19:716-723 Crossref
Alaska Whale Foundation (2024) Annual research report, 2023. Alaska Whale Foundation, Petersburg, AK
Aoki K, Isojunno S, Bellot C, Iwata T and others (2021) Aerial photogrammetry and tag-derived tissue density reveal patterns of lipid-store body condition of humpback whales on their feeding grounds. Proc R Soc B 288:20202307 PubMed Crossref
Ash CE (1952) The body weight of whales. Nor Hvalfangst-Tid 41:364-374
Baker CS, Herman LM (1984) Aggressive behavior between humpback whales (Megaptera novaeangliae) wintering in Hawaiian waters. Can J Zool 62:1922-1937 Crossref
Baker CS, Steel D, Calambokidis J, Falcone E and others (2013) Strong maternal fidelity and natal philopatry shape genetic structure in North Pacific humpback whales. Mar Ecol Prog Ser 494:291-306 Crossref
Barratclough A, Jepson PD, Hamilton PK, Miller CA, Wilson K, Moore MJ (2014) How much does a swimming, underweight, entangled right whale (Eubalaena glacialis) weigh? Calculating the weight at sea, to facilitate accurate dosing of sedatives to enable disentanglement. Mar Mamm Sci 30:1589-1599 Crossref
Baty F, Ritz C, Charles S, Brutsche M, Flandrois JP, Delignette-Muller ML (2015) A toolbox for nonlinear regression in R: the package nlstools. J Stat Softw 66:1-21 Crossref
Bierlich KC, Hewitt J, Schick RS, Pallin L and others (2022) Seasonal gain in body condition of foraging humpback whales along the Western Antarctic Peninsula. Front Mar Sci 9:1036860 Crossref
Bierlich KC, Kane A, Hildebrand L, Bird CN and others (2023) Downsized: gray whales using an alternative foraging ground have smaller morphology. Biol Lett 19:20230043 PubMed Crossref
Bosco L, Otterbeck A, Fransson T, Lindén A, Piha M, Lehikoinen A (2023) Increasing winter temperatures explain body size decrease in wintering bird populations of Northern Europe—but response patterns vary along the spatioclimatic gradient. Glob Ecol Biogeogr 32:2100-2110 Crossref
Bowen WD, den Heyer CE, McMillan JI, Iverson SJ (2015) Offspring size at weaning affects survival to recruitment and reproductive performance of primiparous gray seals. Ecol Evol 5:1412-1424 PubMed Crossref
Braithwaite JE, Meeuwig JJ, Hipsey MR (2015) Optimal migration energetics of humpback whales and the implications of disturbance. Conserv Physiol 3:cov001 PubMed Crossref PubMed Crossref
Brose U, Jonsson T, Berlow EL, Warren P and others (2006) Consumer-resource body-size relationships in natural food webs. Ecology 87:2411-2417
Burton T, Rollinson N, McKelvey S, Stewart DC, Armstrong JD, Metcalfe NB (2020) Adaptive maternal investment in the wild? Links between maternal growth trajectory and offspring size, growth, and survival in contrasting environments. Am Nat 195:678-690 PubMed Crossref
Calambokidis J, Steiger GH, Straley JM, Herman LM and others (2001) Movements and population structure of humpback whales in the North Pacific. Mar Mamm Sci 17:769-794 Crossref
Calambokidis J, Falcone EA, Quinn T, Burdin A and others (2008) SPLASH: structure of populations, levels of abundance and status of humpback whales in the North Pacific. US Department of Commerce, Seattle, WA
Calder WA (1982) The pace of growth: an allometric approach to comparative embryonic and post embryonic growth. J Zool 198:215-225 Crossref
Calder WA (1984) Size, function, and life history. Harvard University Press, Cambridge, MA
Cartwright R, Venema A, Hernandez V, Wyels C, Cesere J, Cesere D (2019) Fluctuating reproductive rates in Hawaii’s humpback whales, Megaptera novaeangliae, reflect recent climate anomalies in the North Pacific. R Soc Open Sci 6:181463 PubMed Crossref
Caruso NM, Sears MW, Adams DC, Lips KR (2014) Widespread rapid reductions in body size of adult salamanders in response to climate change. Glob Change Biol 20:1751-1759 PubMed Crossref
Cates KA, Atkinson S, Pack AA, Straley JM, Gabriele CM, Yin S (2020) Corticosterone in central North Pacific male humpback whales (Megaptera novaeangliae): pairing sighting histories with endocrine markers to assess stress. Gen Comp Endocrinol 296:113540 PubMed Crossref
Cheeseman T, Southerland K, Park J, Olio M and others (2022) Advanced image recognition: a fully automated, high-accuracy photo-identification matching system for humpback whales. Mamm Biol 102:915-929 Crossref PubMed Crossref
Cheeseman T, Southerland K, Acebes JM, Audley K and others (2023) A collaborative and near-comprehensive North Pacific humpback whale photo-ID dataset. Sci Rep 13:10237
Cheeseman T, Barlow J, Acebes JM, Audley K and others (2024) Bellwethers of change: population modelling of North Pacific humpback whales from 2002 through 2021 reveals shift from recovery to climate response. R Soc Open Sci 11:231462 PubMed Crossref
Chittleborough RG (1955) Aspects of reproduction in the male humpback whale, Megaptera nodosa (Bonnaterre). Mar Freshw Res 6:1-29 Crossref
Chittleborough RG (1955) Puberty, physical maturity, and relative growth of the female humpback whale, Megaptera nodosa (Bonnaterre), on the Western Australian coast. Mar Freshw Res 6:315-327 Crossref
Chittleborough RG (1959) Determination of age in the humpback whale, Megaptera nodosa (Bonnaterre). Mar Freshw Res 10:125-143 Crossref
Chittleborough RG (1965) Dynamics of two populations of the humpback whale, Megaptera novaeangliae (Borowski). Mar Freshw Res 16:33-128 Crossref
Christiansen F, Dujon AM, Sprogis KR, Arnould JPY, Bejder L (2016) Noninvasive unmanned aerial vehicle provides estimates of the energetic cost of reproduction in humpback whales. Ecosphere 7:e01468 Crossref
Christiansen F, Vivier F, Charlton C, Ward R, Amerson A, Burnell S, Bejder L (2018) Maternal body size and condition determine calf growth rates in southern right whales. Mar Ecol Prog Ser 592:267-281 Crossref
Christiansen F, Sironi M, Moore MJ, Di Martino M and others (2019) Estimating body mass of free-living whales using aerial photogrammetry and 3D volumetrics. Methods Ecol Evol 10:2034-2044 Crossref
Christiansen F, Sprogis KR, Gross J, Castrillon J, Warick HA, Leunissen E, Bengtson Nash S (2020) Variation in outer blubber lipid concentration does not reflect morphological body condition in humpback whales. J Exp Biol 223:jeb213769 PubMed Crossref
Christiansen F, Rodríguez-González F, Martínez-Aguilar S, Urbán J and others (2021) Poor body condition associated with an unusual mortality event in gray whales. Mar Ecol Prog Ser 658:237-252 Crossref
Christiansen F, Bejder L, Burnell S, Ward R, Charlton C (2022) Estimating the cost of growth in southern right whales from drone photogrammetry data and long-term sighting histories. Mar Ecol Prog Ser 687:173-194 Crossref
Christiansen F, Madsen PT, Andrews-Goff V, Double MC and others (2025) Extreme capital breeding for giants: effects of body size on humpback whale energy expenditure and fasting endurance. Ecol Modell 501:110994 Crossref
Clapham PJ (1992) Age at attainment of sexual maturity in humpback whales, Megaptera novaeangliae. Can J Zool 70:1470-1472 Crossref
Clapham PJ (1996) The social and reproductive biology of humpback whales: an ecological perspective. Mammal Rev 26:27-49 Crossref
Clapham PJ, Wetmore SE, Smith TD, Mead JG (1999) Length at birth and at independence in humpback whales. J Cetacean Res Manage 1:141-146
Clutton-Brock TH, Albon SD, Guinness FE (1989) Fitness costs of gestation and lactation in wild mammals. Nature 337:260-262 PubMed Crossref
Currie JJ, van Aswegen M, Stack SH, West KL, Vivier F, Bejder L (2021) Rapid weight loss in free ranging pygmy killer whales (Feresa attenuata) and the implications for anthropogenic disturbance of odontocetes. Sci Rep 11:8181 PubMed Crossref
Damuth J (1981) Population density and body size in mammals. Nature 290:699-700 Crossref
Dawson SM, Bowman MH, Leunissen E, Sirguey P (2017) Inexpensive aerial photogrammetry for studies of whales and large marine animals. Front Mar Sci 4:366 Crossref
de Araujo Lira AF, do Nascimento Andrade de Almeida Rego F, Salomão RP, de Albuquerque CMR (2020) Effects of habitat quality on body size of the litter dwelling scorpion Tityus pusillus in fragmented rainforests of Brazil. J Arachnol 48:295-299 Crossref
Doroshenko NV (2000) Soviet catches of humpback whales (Megaptera novaeangliae) in the North Pacific. In: Yablokov AV, Zemsky VA (eds) Soviet whaling data (1949-1979). Center for Russian Environmental Policy, Marine Mammal Council, Moscow, p 48-95
Elzhov TV, Mullen KM, Spiess AN, Bolker B (2023) Minpack.lm: R interface to the Levenberg-Marquardt nonlinear least-squares algorithm found in MINPACK, plus support for bounds. Link
Fortune SME, Trites AW, Mayo CA, Rosen DAS, Hamilton PK (2013) Energetic requirements of North Atlantic right whales and the implications for species recovery. Mar Ecol Prog Ser 478:253-272 Crossref
Fortune SME, Moore MJ, Perryman WL, Trites AW (2021) Body growth of North Atlantic right whales (Eubalaena glacialis) revisited. Mar Mamm Sci 37:433-447 Crossref
Frankel AS, Gabriele CM, Yin S, Rickards SH (2022) Humpback whale abundance in Hawai’i: temporal trends and response to climatic drivers. Mar Mamm Sci 38:118-138 Crossref
Frölicher TL, Fischer EM, Gruber N (2018) Marine heatwaves under global warming. Nature 560:360-364 PubMed Crossref
Gabriele CM, Straley JM, Neilson JL (2007) Age at first calving of female humpback whales in southeastern Alaska. Mar Mamm Sci 23:226-239 Crossref
Gabriele CM, Lockyer C, Straley JM, Jurasz CM, Kato H (2010) Sighting history of a naturally marked humpback whale (Megaptera novaeangliae) suggests ear plug growth layer groups are deposited annually. Mar Mamm Sci 26:443-450 Crossref
Gabriele CM, Neilson JL, Straley JM, Baker CS, Cedarleaf JA, Saracco JF (2017) Natural history, population dynamics, and habitat use of humpback whales over 30 years on an Alaska feeding ground. Ecosphere 8:e01641 Crossref
Gabriele CM, Amundson CL, Neilson JL, Straley JM, Baker CS, Danielson SL (2022) Sharp decline in humpback whale (Megaptera novaeangliae) survival and reproductive success in southeastern Alaska during and after the 2014-2016 Northeast Pacific marine heatwave. Mamm Biol 102:1113-1131 Crossref
Gardner JL, Peters A, Kearney MR, Joseph L, Heinsohn R (2011) Declining body size: A third universal response to warming? Trends Ecol Evol 26:285-291 PubMed Crossref
Gillooly JF, Brown JH, West GB, Savage VM, Charnov EL (2001) Effects of size and temperature on metabolic rate. Science 293:2248-2251 PubMed Crossref
Ginther SC, Cameron H, White CR, Marshall DJ (2024) Metabolic loads and the costs of metazoan reproduction. Science 384:763-767 PubMed Crossref
Gittleman JL, Thompson SD (1988) Energy allocation in mammalian reproduction. Am Zool 28:863-875 Crossref
Glockner DA (1983) Determining the sex of humpback whales (Megaptera novaeangliae) in their natural environment. In: Payne R (ed) Communication and behavior of whales. Westview Press, Boulder, CO, p 447-464
Goldbogen JA, Pyenson ND, Madsen PT (2023) How whales dive, feast, and fast: the ecophysiological drivers and limits of foraging in the evolution of cetaceans. Annu Rev Ecol Evol Syst 54:307-325 Crossref
Gompertz B (1825) On the nature of the function expressive of the law of human mortality, and on a new mode of determining the value of life contingencies. Philos Trans R Soc Lond 115:513-583
Gough WT, Segre PS, Bierlich KC, Cade DE and others (2019) Scaling of swimming performance in baleen whales. J Exp Biol 222:jeb204172 PubMed Crossref
Gough WT, Cade DE, Czapanskiy MF, Potvin J and others (2022) Fast and furious: energetic tradeoffs and scaling of high-speed foraging in rorqual whales. Integr Org Biol 4:obac038 PubMed Crossref
Herman LM, Antinoja RC (1977) Humpback whales in the Hawaiian breeding waters: population and pod characteristics. Sci Rep Whales Res Inst 29:59-85
Herman LM, Pack AA, Rose K, Craig A, Herman EYK, Hakala S, Milette A (2011) Resightings of humpback whales in Hawaiian waters over spans of 10-32 years: site fidelity, sex ratios, calving rates, female demographics, and the dynamics of social and behavioral roles of individuals. Mar Mamm Sci 27:736-768 Crossref
Herrera E, Amusquivar E (2000) Lipid metabolism in the fetus and the newborn. Diabetes Metab Res Rev 16:202-210 PubMed Crossref
Holyoake C, Stephens N, Coughran D (2012) Collection of baseline data on humpback whale (Megaptera novaeangliae) health and causes of mortality for long-term monitoring in Western Australia. Murdoch University, Perth PubMed Crossref
IJsseldijk LL, Hessing S, Mairo A, 10 Doeschate MTI and others (2021) Nutritional status and prey energy density govern reproductive success in a small cetacean. Sci Rep 11:19201
Irvine LG (2018) Humpback whale migration: new insights into the energetics of a long-distance migrant. PhD thesis, University of Tasmania, Hobart
IT’IS Foundation (2022) Tissue properties database, v4:1. Link
Johnson CN (1999) Relationships between body size and population density of animals: the problem of the scaling of study area in relation to body size. Oikos 85:565-569 Crossref
Jørgensen C, Enberg K, Mangel M (2016) Modelling and interpreting fish bioenergetics: a role for behaviour, life-history traits and survival trade-offs. J Fish Biol 88:389-402 PubMed Crossref
Jurasz CM, Palmer VP (1981) Censusing and establishing age composition of humpback whales (Megaptera novaeangliae), employing photodocumentation in Glacier Bay National Monument, Alaska. Report to the National Park Service, Anchorage, AK
Katona S, Baxter B, Brazier O, Kraus S, Perkins J, Whitehead H (1979) Identification of humpback whales by fluke photographs. In: Winn HE, Olla BL (eds) Behavior of marine animals: current perspectives in research. Springer, Boston, MA, p 33-44 Crossref
Laird AK (1966) Postnatal growth of birds and mammals. Growth 30:349-363 PubMed
Lockyer C (1976) Body weights of some species of large whales. ICES J Mar Sci 36:259-273 Crossref
Lockyer C (1981) Estimation of the energy costs of growth, maintenance and reproduction of the female minke whale, (Balaenoptera acutorostrata), from the Southern Hemisphere. Rep Int Whal Commn 31:337-343
Lockyer C (1981) Growth and energy budgets of large baleen whales from the Southern Hemisphere. In: FAO Fish Series 5: mammals in the seas, Vol 3: general papers and large cetaceans. FAO, Rome, p 379-487
Lockyer C (1984) Review of baleen whale (Mysticeti) reproduction and implications for management. Rep Int Whal Commn Spec Issue 6:27-50
Lockyer C (1987) Evaluation of the role of fat reserves in relation to the ecology of North Atlantic fin and sei whales. In: Huntley AC, Costa DP, Worthy GAJ, Castellini MA (eds) Approaches to marine mammal energetics. Society for Marine Mammalogy, Lawrence, KS, p 183-203
Lockyer C (2007) All creatures great and smaller: a study in cetacean life history energetics. J Mar Biol Assoc UK 87:1035-1045 Crossref
Lockyer C, McConnell LC, Waters TD (1985) Body condition in terms of anatomical and biochemical assessment of body fat in North Atlantic fin and sei whales. Can J Zool 63:2328-2338 Crossref
Mazerolle MJ (2015) AICcmodavg: model selection and multimodel inference based on (Q)AIC(c). R version 2.3-3. Link
McHuron EA, Adamczak S, Costa DP, Booth C (2023) Estimating reproductive costs in marine mammal bioenergetic models: a review of current knowledge and data availability. Conserv Physiol 11:coac080 PubMed Crossref
McMahon C, Burton H, Bester M (2000) Weaning mass and the future survival of juvenile southern elephant seals, Mirounga leonina, at Macquarie Island. Antarct Sci 12:149-153 Crossref
Mikhalev Y (2019) Whales of the Southern Ocean: biology, whaling and perspectives of population recovery. Springer International Publishing, Cham
Miller CA, Best PB, Perryman WL, Baumgartner MF, Moore MJ (2012) Body shape changes associated with reproductive status, nutritive condition and growth in right whales Eubalaena glacialis and E. australis. Mar Ecol Prog Ser 459:135-156 Crossref
Millward DJ, Garlick PJ, Reeds PJ (1976) The energy cost of growth. Proc Nutr Soc 35:339-349 PubMed Crossref
Napoli C, Hirtle N, Stepanuk J, Christiansen F and others (2024) Drone-based photogrammetry reveals differences in humpback whale body condition and mass across North Atlantic foraging grounds. Front Mar Sci 11:3389 Crossref
Nishiwaki M (1959) Humpback whales in Ryukyuan waters. Sci Rep Whales Res Inst 14:49-87
Oftedal OT (2000) Use of maternal reserves as a lactation strategy in large mammals. Proc Nutr Soc 59:99-106 PubMed Crossref
Ogle DH, Doll JC, Wheeler P, Dinno A (2023) FSA: simple fisheries stock assessment methods. R version: 0.9.3. Link
Oke KB, Cunningham CJ, Westley PAH, Baskett ML and others (2020) Recent declines in salmon body size impact ecosystems and fisheries. Nat Commun 11:4155 PubMed Crossref
Oliver ECJ, Burrows MT, Donat MG, Sen Gupta A and others (2019) Projected marine heatwaves in the 21st century and the potential for ecological impact. Front Mar Sci 6:734 Crossref
Pack A, Herman L, Craig A, Spitz S, Deakos M (2002) Penis extrusions by humpback whales (Megaptera novaeangliae). Aquat Mamm 28:131-146 Link
Pack AA, Herman LM, Spitz SS, Hakala S, Deakos MH, Herman EYK (2009) Male humpback whales in the Hawaiian breeding grounds preferentially associate with larger females. Anim Behav 77:653-662 Crossref
Pack AA, Herman LM, Craig AS, Spitz SS and others (2017) Habitat preferences by individual humpback whale mothers in the Hawaiian breeding grounds vary with the age and size of their calves. Anim Behav 133:131-144 Crossref
Pettersen AK, White CR, Marshall DJ (2015) Why does offspring size affect performance? Integrating metabolic scaling with life-history theory. Proc R Soc B 282:20151946 PubMed Crossref
Pirotta E, Mangel M, Costa DP, Goldbogen J and others (2019) Anthropogenic disturbance in a changing environment: modelling lifetime reproductive success to predict the consequences of multiple stressors on a migratory population. Oikos 128:1340-1357 Crossref
Pirotta E, Bierlich KC, New L, Hildebrand L, Bird CN, Fernandez Ajó A, Torres LG (2024) Modeling individual growth reveals decreasing gray whale body length and correlations with ocean climate indices at multiple scales. Glob Change Biol 30:e17366 PubMed Crossref
Plön S, Andra K, Auditore L, Gegout C and others (2024) Marine mammals as indicators of Anthropocene ocean health. npj Biodiversity 3:24 Crossref
Polidori C, Gutiérrez-Cánovas C, Sánchez E, Tormos J, Castro L, Sánchez-Fernández D (2020) Climate change-driven body size shrinking in a social wasp. Ecol Entomol 45:130-141 Crossref
Potvin J, Cade DE, Werth AJ, Shadwick RE, Goldbogen JA (2021) Rorqual lunge-feeding energetics near and away from the kinematic threshold of optimal efficiency. Integr Org Biol 3:obab005 PubMed Crossref
Queiros Q, McKenzie DJ, Dutto G, Killen S, Saraux C, Schull Q (2024) Fish shrinking, energy balance and climate change. Sci Total Environ 906:167310 PubMed Crossref
R Core Team (2023) R: a language and environment for statistical computing. R Foundation for Statistical Computing, Vienna
Reading CJ (2007) Linking global warming to amphibian declines through its effects on female body condition and survivorship. Oecologia 151:125-131 PubMed Crossref
Ricker WE (1979) Growth rates and models. In: Hoar WS, Randall DJ, Bret JR (eds) Fish physiology: bioenergetics and growth. Academic Press, Orlando, FL, p 677-743 Crossref
Roberts SB, Young VR (1988) Energy costs of fat and protein deposition in the human infant. Am J Clin Nutr 48:951-955 PubMed Crossref
Rode KD, Amstrup SC, Regehr EV (2010) Reduced body size and cub recruitment in polar bears associated with sea ice decline. Ecol Appl 20:768-782 PubMed Crossref
Ronget V, Gaillard JM, Coulson T, Garratt M, Gueyffier F, Lega JC, Lemaître JF (2018) Causes and consequences of variation in offspring body mass: meta-analyses in birds and mammals. Biol Rev Camb Philos Soc 93:1-27 PubMed Crossref
Russell G, Cagnazzi D, Colefax A, Sprogis KR, Christiansen F (2023) Cost of migration and migratory timing in Western Australian humpback whales. Mar Mamm Sci 40:e13074
Sargent RC, Gross MR, Van Den Berghe EP (1986) Male mate choice in fishes. Anim Behav 34:545-550 Crossref
Schmidt-Nielsen K (1984) Scaling. Why is animal size so important? Cambridge University Press, Cambridge Crossref
Searing KB, Lomolino MV, Rozzi R (2023) Melting climates shrink North American small mammals. Proc Natl Acad Sci USA 120:e2310855120 PubMed Crossref
Sharpe DMT, Hendry AP (2009) Life history change in commercially exploited fish stocks: an analysis of trends across studies. Evol Appl 2:260-275 PubMed Crossref
Speakman JR (2005) Body size, energy metabolism and lifespan. J Exp Biol 208:1717-1730 PubMed Crossref
Speakman JR, Selman C, McLaren JS, Harper EJ (2002) Living fast, dying when? The link between aging and energetics. J Nutr 132:1583S-1597S PubMed Crossref
Spitz SS (1999) Size measurement of humpback whales using underwater videogrammetry. PhD Thesis, University of Hawai‘i at Mānoa, Honolulu, HI
Spitz SS, Herman LM, Pack AA, Deakos MH (2002) The relation of body size of male humpback whales to their social roles on the Hawaiian winter grounds. Can J Zool 80:1938-1947 Crossref
Stewart JD, Durban JW, Knowlton AR, Lynn MS and others (2021) Decreasing body lengths in North Atlantic right whales. Curr Biol 31:3174-3179.e3 PubMed Crossref
Stewart JD, Durban JW, Fearnbach H, Barrett-Lennard LG, Casler PK, Ward EJ, Dapp DR (2021) Survival of the fattest: linking body condition to prey availability and survivorship of killer whales. Ecosphere 12:e03660 Crossref
Stewart JD, Joyce TW, Durban JW, Calambokidis J and others (2023) Boom-bust cycles in gray whales associated with dynamic and changing Arctic conditions. Science 382:207-211 PubMed Crossref
Straley JM, Gabriele CM, Baker CS (1994) Annual reproduction by individually identified humpback whales (Megaptera novaeangliae)in Alaskan Waters. Mar Mamm Sci 10:87-92 Crossref
van Aswegen M, Szabo A, Currie JJ, Stack SH and others (2025) Maternal investment, body condition and calf growth in humpback whales. J Physiol 603:551-578 PubMed Crossref
van Aswegen M, Szabo A, Currie JJ, Stack SH and others (2025) Energetic cost of gestation and prenatal growth in humpback whales. J Physiol 603:529-550 PubMed Crossref
van der Hoop J, Corkeron P, Moore M (2017) Entanglement is a costly life-history stage in large whales. Ecol Evol 7:92-106 PubMed Crossref
Víkingsson GA (1990) Energetic studies on fin and sei whales caught off Iceland. Rep Int Whal Commn 40:365-373
Víkingsson GA, Auðunsson G, Elvarsson B, Gunnlaugsson T (2013) Energy storage in common minke whales (Balaenoptera acutorostrata) in Icelandic waters 2003-2007—chemical composition of tissues and organs. Paper SC/F13/SP10 presented to the IWC Scientific Committee
Vivier F, Wells RS, Hill MC, Yano KM and others (2023) Quantifying the age structure of free-ranging delphinid populations: testing the accuracy of unoccupied aerial system photogrammetry. Ecol Evol 13:e10082 PubMed Crossref
von Bertalanffy L (1938) A quantitative theory of organic growth (inquiries on growth laws. II). Hum Biol 10:181-213
Wilson LN, Gardner JD, Wilson JP, Farnsworth A and others (2024) Global latitudinal gradients and the evolution of body size in dinosaurs and mammals. Nat Commun 15:2864 PubMed Crossref
Yuan R, Hascup E, Hascup K, Bartke A (2023) Relationships among development, growth, body size, reproduction, aging, and longevity—trade-offs and pace-of-life. Biochemistry (Mosc) 88:1692-1703 PubMed Crossref
Zepelin H, Siegel JM, Tobler I (2005) Mammalian sleep. In: Kryger MH, Roth T, Dement WC (eds) Principles and practice of sleep medicine, 4th edn. WB Saunders, Philadelphia, PA, p 91-100
Zimova M, Weeks BC, Willard DE, Giery ST, Jirinec V, Burner RC, Winger BM (2023) Body size predicts the rate of contemporary morphological change in birds. Proc Natl Acad Sci USA 120:e2206971120 PubMed Crossref
M. van Aswegen (Corresponding Author)
- Marine Mammal Research Program, Hawai‘i Institute of Marine Biology, University of Hawai‘i at Mānoa, Hawai‘i 96744, USA
- Alaska Whale Foundation, PO Box 1927, Petersburg, Alaska 99833, USA
A. Szabo (Co-author)
- Alaska Whale Foundation, PO Box 1927, Petersburg, Alaska 99833, USA
- Hawai‘i Institute of Marine Biology, University of Hawai‘i at Mānoa, Hawai‘i 96744, USA
J. Currie (Co-author)
- Marine Mammal Research Program, Hawai‘i Institute of Marine Biology, University of Hawai‘i at Mānoa, Hawai‘i 96744, USA
- Pacific Whale Foundation, Maui, Hawai‘i 96793, USA
S. Stack (Co-author)
- Pacific Whale Foundation, Maui, Hawai‘i 96793, USA
- Southern Ocean Persistent Organic Pollutants Program, School of Environment and Science, Griffith University, Queensland 4111, Australia
J. Straley (Co-author)
- University of Alaska Southeast, Alaska 99835, USA
J. Neilson (Co-author)
- Glacier Bay National Park and Preserve, Gustavus, Alaska 99826, USA
C. Gabriele (Co-author)
- Glacier Bay National Park and Preserve, Gustavus, Alaska 99826, USA
K. Cates (Co-author)
- University of Alaska Fairbanks, College of Fisheries and Ocean Sciences, Alaska 99775, USA
C. Baker (Co-author)
- Marine Mammal Institute, Oregon State University, Oregon 97365, USA
D. Steel (Co-author)
- Marine Mammal Institute, Oregon State University, Oregon 97365, USA
A. Pack (Co-author)
- Psychology and Biology Departments, University of Hawai‘i at Hilo, Hawai‘i, USA and The Dolphin Institute, Hilo, Hawai‘i 96720, USA
K. West (Co-author)
- Health and Stranding Lab, College of Tropical Agriculture and Human Resources, University of Hawai‘i at Mānoa, Hawai‘i 96822, USA
N. Hofmann (Co-author)
- Health and Stranding Lab, College of Tropical Agriculture and Human Resources, University of Hawai‘i at Mānoa, Hawai‘i 96822, USA
R. Toonen (Co-author)
- Hawai‘i Institute of Marine Biology, University of Hawai‘i at Mānoa, Hawai‘i 96744, USA
C. Loockerman (Co-author)
- Hawai‘i Institute of Marine Biology, University of Hawai‘i at Mānoa, Hawai‘i 96744, USA
F. Sullivan (Co-author)
- Pacific Whale Foundation, Maui, Hawai‘i 96793, USA
T. Cheeseman (Co-author)
- Marine Ecology Research Centre, Southern Cross University, Lismore, New South Wales 2480, Australia
- Happywhale, Santa Cruz, California 95060, USA
F. Sharpe (Co-author)
- Alaska Whale Foundation, PO Box 1927, Petersburg, Alaska 99833, USA
L. Bejder (Co-author)
- Marine Mammal Research Program, Hawai‘i Institute of Marine Biology, University of Hawai‘i at Mānoa, Hawai‘i 96744, USA
- Zoophysiology, Department of Bioscience, Aarhus University, Aarhus 8000, Denmark
Handling Editor:
Peter Corkeron, Nathan, Queensland, Australia
Reviewers:
T. Horton and 2 anonymous referees
Acknowledgements:
The present work was part of M.v.A.’s PhD thesis. Field research was conducted under National Marine Fisheries Service (NMFS) permit numbers 21476, 21321, and 19703, with all activities approved by the University of Hawaiʻi Institutional Animal Care and Use Committee. Stranding response, necropsy, and sample collection of the humpback calves were conducted under NMFS permit numbers 18786 and 24359. We thank Cameron Nemeth and Kalikoaloha Martin for translating the abstract into Hawaiian. We thank the numerous research assistants who helped with fieldwork and data processing. We also thank PacWhale Eco-Adventures, Chrissy Lovitt and Emma Nelson (Maui Ocean Adventures), Lee James, Jason Jones, and the Guth family for their support. We thank Charles Jurasz, Susan Jurasz, Peter Jurasz, and Ginny Palmer (Sea Search) for providing valuable long-term sighting data. We are also grateful to Marc Lammers (Oceanwide Science Institute) for providing sighting date information. We thank Elmer Ka‘ai for his support relating to humpback whale strandings and sample collection. We thank the following public Happywhale data contributors: Dennis Rogers, Jim Nahmens, and Andy Wolcott (Alaska Sea Adventures), Atlantis Cruises, Kenneth O’Brien (Fantasy Cruises), Jonathon Whitney, Capt. Steves Rafting Adventures, Bruce Whittington (Bluewater Adventures), Paul Pudwell (Sooke Whale Watching), Kenneth Held (Safari Boat Excursions Maui, Maui Adventure Cruises), Brooke McKinley (Hawai‘i Ocean Sports), Eric Austin Yee (PacWhale Eco-Adventures), Tracy Desiderio (Harv and Marv’s Outback Alaska), John Forde (The Whale Centre Tofino), Dive Maui, Mindy Huston, Heather Baskey, Emily Cramer, and K. C. Dermody (Jayleen’s Alaska). We are grateful to Zoltan Nemeth (cetek-art.com) for providing illustrations. This paper represents Hawai‘i Institute of Marine Biology and School of Ocean and Earth Science and Technology contribution nos. 1998 and 11963, respectively. Hawai‘i fieldwork was funded through the University of Hawai‘i at Mānoa, the US Department of Defense’s Defense University Research Instrumentation Program (N00014-19-2612), ‘Our Oceans’, Netflix, Wildspace Productions and Freeborne Media, Office of Naval Research (N000142012624), Omidyar Ohana Foundation, the National Marine Sanctuary Foundation (21-10-B-334), and PacWhale Eco-Adventures, as well as members and donors of Pacific Whale Foundation. Southeast Alaska research was funded through awards from the National Geographic Society, the Lindblad Expeditions-National Geographic (LEX-NG) Funds, and the North Pacific Research Board (award no. 2114). Graduate assistantships for M.v.A. were funded by a Denise B. Evans Oceanography Fellowship, North Pacific Research Board grant (2114), and the Dolphin Quest General Science and Conservation Fund. Long-term humpback whale population monitoring in Glacier Bay and Icy Strait was funded by the National Park Service. Stranding response, necropsy, and tissue processing of the humpback whale calves were supported by the NOAA John H. Prescott Marine Mammal Rescue Assistance Grant Program.
