Wells Randall S.

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Randall S.

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  • Article
    Vocal copying of individually distinctive signature whistles in bottlenose dolphins
    (The Royal Society, 2013-02-20) King, Stephanie L. ; Sayigh, Laela S. ; Wells, Randall S. ; Fellner, Wendi ; Janik, Vincent M.
    Vocal learning is relatively common in birds but less so in mammals. Sexual selection and individual or group recognition have been identified as major forces in its evolution. While important in the development of vocal displays, vocal learning also allows signal copying in social interactions. Such copying can function in addressing or labelling selected conspecifics. Most examples of addressing in non-humans come from bird song, where matching occurs in an aggressive context. However, in other animals, addressing with learned signals is very much an affiliative signal. We studied the function of vocal copying in a mammal that shows vocal learning as well as complex cognitive and social behaviour, the bottlenose dolphin (Tursiops truncatus). Copying occurred almost exclusively between close associates such as mother–calf pairs and male alliances during separation and was not followed by aggression. All copies were clearly recognizable as such because copiers consistently modified some acoustic parameters of a signal when copying it. We found no evidence for the use of copying in aggression or deception. This use of vocal copying is similar to its use in human language, where the maintenance of social bonds appears to be more important than the immediate defence of resources.
  • Article
    Field energetics and lung function in wild bottlenose dolphins, Tursiops truncatus, in Sarasota Bay Florida
    (Royal Society, 2018-01-17) Fahlman, Andreas ; Brodsky, Micah ; Wells, Randall S. ; McHugh, Katherine ; Allen, Jason ; Barleycorn, Aaron ; Sweeney, Jay C. ; Fauquier, Deborah A. ; Moore, Michael J.
    We measured respiratory flow rates, and expired O2 in 32 (2–34 years, body mass [Mb] range: 73–291 kg) common bottlenose dolphins (Tursiops truncatus) during voluntary breaths on land or in water (between 2014 and 2017). The data were used to measure the resting O2 consumption rate (V˙O2, range: 0.76–9.45ml O2min−1 kg−1) and tidal volume (VT, range: 2.2–10.4 l) during rest. For adult dolphins, the resting VT, but not V˙O2, correlated with body mass (Mb, range: 141–291 kg) with an allometric mass-exponent of 0.41. These data suggest that the mass-specific VT of larger dolphins decreases considerably more than that of terrestrial mammals (mass-exponent: 1.03). The average resting sV˙O2 was similar to previously published metabolic measurements from the same species. Our data indicate that the resting metabolic rate for a 150 kg dolphin would be 3.9 ml O2 min−1 kg−1, and the metabolic rate for active animals, assuming a multiplier of 3–6, would range from 11.7 to 23.4 ml O2 min−1 kg−1.
  • Article
    Resting metabolic rate and lung function in wild offshore common bottlenose dolphins, Tursiops truncatus, near Bermuda
    (Frontiers Media, 2018-07-17) Fahlman, Andreas ; McHugh, Katherine ; Allen, Jason ; Barleycorn, Aaron ; Allen, Austin ; Sweeney, Jay C. ; Stone, Rae ; Faulkner Trainor, Robyn ; Bedford, Guy ; Moore, Michael J. ; Jensen, Frants H. ; Wells, Randall S.
    Diving mammals have evolved a suite of physiological adaptations to manage respiratory gases during extended breath-hold dives. To test the hypothesis that offshore bottlenose dolphins have evolved physiological adaptations to improve their ability for extended deep dives and as protection for lung barotrauma, we investigated the lung function and respiratory physiology of four wild common bottlenose dolphins (Tursiops truncatus) near the island of Bermuda. We measured blood hematocrit (Hct, %), resting metabolic rate (RMR, l O2 ⋅ min-1), tidal volume (VT, l), respiratory frequency (fR, breaths ⋅ min-1), respiratory flow (l ⋅ min-1), and dynamic lung compliance (CL, l ⋅ cmH2O-1) in air and in water, and compared measurements with published results from coastal, shallow-diving dolphins. We found that offshore dolphins had greater Hct (56 ± 2%) compared to shallow-diving bottlenose dolphins (range: 30–49%), thus resulting in a greater O2 storage capacity and longer aerobic diving duration. Contrary to our hypothesis, the specific CL (sCL, 0.30 ± 0.12 cmH2O-1) was not different between populations. Neither the mass-specific RMR (3.0 ± 1.7 ml O2 ⋅ min-1 ⋅ kg-1) nor VT (23.0 ± 3.7 ml ⋅ kg-1) were different from coastal ecotype bottlenose dolphins, both in the wild and under managed care, suggesting that deep-diving dolphins do not have metabolic or respiratory adaptations that differ from the shallow-diving ecotypes. The lack of respiratory adaptations for deep diving further support the recently developed hypothesis that gas management in cetaceans is not entirely passive but governed by alteration in the ventilation-perfusion matching, which allows for selective gas exchange to protect against diving related problems such as decompression sickness.
  • Article
    Signal-specific amplitude adjustment to noise in common bottlenose dolphins (Tursiops truncatus)
    (Company of Biologists, 2019-11-08) Kragh, Ida M. ; McHugh, Katherine ; Wells, Randall S. ; Sayigh, Laela S. ; Janik, Vincent M. ; Tyack, Peter L. ; Jensen, Frants H.
    Anthropogenic underwater noise has increased over the past century, raising concern about the impact on cetaceans that rely on sound for communication, navigation and locating prey and predators. Many terrestrial animals increase the amplitude of their acoustic signals to partially compensate for the masking effect of noise (the Lombard response), but it has been suggested that cetaceans almost fully compensate with amplitude adjustments for increasing noise levels. Here, we used sound-recording DTAGs on pairs of free-ranging common bottlenose dolphins (Tursiops truncatus) to test (i) whether dolphins increase signal amplitude to compensate for increasing ambient noise and (ii) whether adjustments are identical for different signal types. We present evidence of a Lombard response in the range 0.1–0.3 dB per 1 dB increase in ambient noise, which is similar to that of terrestrial animals, but much lower than the response reported for other cetaceans. We found that signature whistles tended to be louder and with a lower degree of amplitude adjustment to noise compared with non-signature whistles, suggesting that signature whistles may be selected for higher output levels and may have a smaller scope for amplitude adjustment to noise. The consequence of the limited degree of vocal amplitude compensation is a loss of active space during periods of increased noise, with potential consequences for group cohesion, conspecific encounter rates and mate attraction.
  • Article
    What’s in a voice? Dolphins do not use voice cues for individual recognition
    (Springer, 2017-08-08) Sayigh, Laela S. ; Wells, Randall S. ; Janik, Vincent M.
    Most mammals can accomplish acoustic recognition of other individuals by means of “voice cues,” whereby characteristics of the vocal tract render vocalizations of an individual uniquely identifiable. However, sound production in dolphins takes place in gas-filled nasal sacs that are affected by pressure changes, potentially resulting in a lack of reliable voice cues. It is well known that bottlenose dolphins learn to produce individually distinctive signature whistles for individual recognition, but it is not known whether they may also use voice cues. To investigate this question, we played back non-signature whistles to wild dolphins during brief capture-release events in Sarasota Bay, Florida. We hypothesized that non-signature whistles, which have varied contours that can be shared among individuals, would be recognizable to dolphins only if they contained voice cues. Following established methodology used in two previous sets of playback experiments, we found that dolphins did not respond differentially to non-signature whistles of close relatives versus known unrelated individuals. In contrast, our previous studies showed that in an identical context, dolphins reacted strongly to hearing the signature whistle or even a synthetic version of the signature whistle of a close relative. Thus, we conclude that dolphins likely do not use voice cues to identify individuals. The low reliability of voice cues and the need for individual recognition were likely strong selective forces in the evolution of vocal learning in dolphins.
  • Article
    How do marine mammals manage and usually avoid gas emboli formation and gas embolic pathology? critical clues from studies of wild dolphins
    (Frontiers Media, 2021-02-05) Fahlman, Andreas ; Moore, Michael J. ; Wells, Randall S.
    Decompression theory has been mainly based on studies on terrestrial mammals, and may not translate well to marine mammals. However, evidence that marine mammals experience gas bubbles during diving is growing, causing concern that these bubbles may cause gas emboli pathology (GEP) under unusual circumstances. Marine mammal management, and usual avoidance, of gas emboli and GEP, or the bends, became a topic of intense scientific interest after sonar-exposed, mass-stranded deep-diving whales were observed with gas bubbles. Theoretical models, based on our current understanding of diving physiology in cetaceans, predict that the tissue and blood N2 levels in the bottlenose dolphin (Tursiops truncatus) are at levels that would result in severe DCS symptoms in similar sized terrestrial mammals. However, the dolphins appear to have physiological or behavioral mechanisms to avoid excessive blood N2 levels, or may be more resistant to circulating bubbles through immunological/biochemical adaptations. Studies on behavior, anatomy and physiology of marine mammals have enhanced our understanding of the mechanisms that are thought to prevent excessive uptake of N2. This has led to the selective gas exchange hypothesis, which provides a mechanism how stress-induced behavioral change may cause failure of the normal physiology, which results in excessive uptake of N2, and in extreme cases may cause formation of symptomatic gas emboli. Studies on cardiorespiratory function have been integral to the development of this hypothesis, with work initially being conducted on excised tissues and cadavers, followed by studies on anesthetized animals or trained animals under human care. These studies enabled research on free-ranging common bottlenose dolphins in Sarasota Bay, FL, and off Bermuda, and have included work on the metabolic and cardiorespiratory physiology of both shallow- and deep-diving dolphins and have been integral to better understand how cetaceans can dive to extreme depths, for long durations.
  • Preprint
    Identifying signature whistles from recordings of groups of unrestrained bottlenose dolphins (Tursiops truncatus)
    ( 2011-09-29) Janik, Vincent M. ; King, Stephanie L. ; Sayigh, Laela S. ; Wells, Randall S.
    Bottlenose dolphins (Tursiops truncatus) have individually-distinctive signature whistles. Each individual dolphin develops its own unique frequency modulation pattern and uses it to broadcast its identity. However, underwater sound localization is challenging, and researchers have had difficulties identifying signature whistles. The traditional method to identify them involved isolating individuals. In this context, the signature whistle is the most commonly produced whistle type of an animal. However, most studies on wild dolphins cannot isolate animals. We present a novel method, SIGID, that can identify signature whistles in recordings of groups of dolphins recorded via a single hydrophone. We found that signature whistles tend to be delivered in bouts with whistles of the same type occurring within 1-10 s of each other. Non-signature whistles occur over longer or shorter periods, and this distinction can be used to identify signature whistles in a recording. We tested this method on recordings from wild and captive bottlenose dolphins and show thresholds needed to identify signature whistles reliably. SIGID will facilitate the study of signature whistle use in the wild, signature whistle diversity between different populations, and potentially allow signature whistles to be used in mark-recapture studies.
  • Preprint
    Convergence of marine megafauna movement patterns in coastal and open oceans
    ( 2017-09) Sequeira, Ana M. M. ; Rodríguez, Jorge P. ; Eguíluz, Víctor M. ; Harcourt, Robert ; Hindell, Mark ; Sims, David W. ; Duarte, Carlos M. ; Costa, Daniel P. ; Fernández-Gracia, Juan ; Ferreira, Luciana C. ; Hays, Graeme ; Heupel, Michelle R. ; Meekan, Mark G. ; Aven, Allen ; Bailleul, Frédéric ; Baylis, Alastair M. M. ; Berumen, Michael L. ; Braun, Camrin D. ; Burns, Jennifer ; Caley, M. Julian ; Campbell, R. ; Carmichael, Ruth H. ; Clua, Eric ; Einoder, Luke D. ; Friedlaender, Ari S. ; Goebel, Michael E. ; Goldsworthy, Simon D. ; Guinet, Christophe ; Gunn, John ; Hamer, D. ; Hammerschlag, Neil ; Hammill, Mike O. ; Hückstädt, Luis A. ; Humphries, Nicolas E. ; Lea, Mary-Anne ; Lowther, Andrew D. ; Mackay, Alice ; McHuron, Elizabeth ; McKenzie, J. ; McLeay, Lachlan ; McMahon, Cathy R. ; Mengersen, Kerrie ; Muelbert, Monica M. C. ; Pagano, Anthony M. ; Page, B. ; Queiroz, N. ; Robinson, Patrick W. ; Shaffer, Scott A. ; Shivji, Mahmood ; Skomal, Gregory B. ; Thorrold, Simon R. ; Villegas-Amtmann, Stella ; Weise, Michael ; Wells, Randall S. ; Wetherbee, Bradley M. ; Wiebkin, A. ; Wienecke, Barbara ; Thums, Michele
    The extent of increasing anthropogenic impacts on large marine vertebrates partly depends on the animals’ movement patterns. Effective conservation requires identification of the key drivers of movement including intrinsic properties and extrinsic constraints associated with the dynamic nature of the environments the animals inhabit. However, the relative importance of intrinsic versus extrinsic factors remains elusive. We analyse a global dataset of 2.8 million locations from > 2,600 tracked individuals across 50 marine vertebrates evolutionarily separated by millions of years and using different locomotion modes (fly, swim, walk/paddle). Strikingly, movement patterns show a remarkable convergence, being strongly conserved across species and independent of body length and mass, despite these traits ranging over 10 orders of magnitude among the species studied. This represents a fundamental difference between marine and terrestrial vertebrates not previously identified, likely linked to the reduced costs of locomotion in water. Movement patterns were primarily explained by the interaction between species-specific traits and the habitat(s) they move through, resulting in complex movement patterns when moving close to coasts compared to more predictable patterns when moving in open oceans. This distinct difference may be associated with greater complexity within coastal micro-habitats, highlighting a critical role of preferred habitat in shaping marine vertebrate global movements. Efforts to develop understanding of the characteristics of vertebrate movement should consider the habitat(s) through which they move to identify how movement patterns will alter with forecasted severe ocean changes, such as reduced Arctic sea ice cover, sea level rise and declining oxygen content.
  • Preprint
    Rehabilitation and release of marine mammals in the United States : risks and benefits
    ( 2007-04) Moore, Michael J. ; Early, Greg A. ; Touhey, Kathleen M. ; Barco, Susan G. ; Gulland, Frances M. ; Wells, Randall S.
    Rehabilitation of stranded marine mammals elicits polarized attitudes: initially done alongside display collections, but release of rehabilitated animals has become more common. Justifications include animal welfare, management of beach use conflict, research, conservation, and public education. Rehabilitation cost and risks have been identified which vary in degree supported by data rather than perception. These include conflict with fisheries for resources, ignorance of recipient population ecology, poor understanding of long term survival, support of the genetically not-so-fit, introduction of novel or antibiotic resistant pathogens, harm to human health and cost. Thus facilities must balance their welfare appeal against public education, habitat restoration, human impact reduction, and other conservation activities. Benefits to rehabilitating marine mammals are the opportunity to support the welfare of disabled animals and to publish good science and so advance our understanding of wild populations. In specific cases, the status of a population may make conservation the main reason for rehabilitation. These three reasons for rehabilitation lead to contrasting, and sometimes conflicting, management needs. We therefore outline a decision tree for rehabilitation managers using criteria for each management decision, based on welfare, logistics, conservation, research and funding to define limits on the number of animals released to the wild.
  • Article
    Bubbles in live-stranded dolphins
    (The Royal Society, 2011-10-12) Dennison, Sophie ; Moore, Michael J. ; Fahlman, Andreas ; Moore, Kathleen M. T. ; Sharp, Sarah M. ; Harry, Charles T. ; Hoppe, Jane M. ; Niemeyer, Misty E. ; Lentell, Betty J. ; Wells, Randall S.
    Bubbles in supersaturated tissues and blood occur in beaked whales stranded near sonar exercises, and post-mortem in dolphins bycaught at depth and then hauled to the surface. To evaluate live dolphins for bubbles, liver, kidneys, eyes and blubber–muscle interface of live-stranded and capture-release dolphins were scanned with B-mode ultrasound. Gas was identified in kidneys of 21 of 22 live-stranded dolphins and in the hepatic portal vasculature of 2 of 22. Nine then died or were euthanized and bubble presence corroborated by computer tomography and necropsy, 13 were released of which all but two did not re-strand. Bubbles were not detected in 20 live wild dolphins examined during health assessments in shallow water. Off-gassing of supersaturated blood and tissues was the most probable origin for the gas bubbles. In contrast to marine mammals repeatedly diving in the wild, stranded animals are unable to recompress by diving, and thus may retain bubbles. Since the majority of beached dolphins released did not re-strand it also suggests that minor bubble formation is tolerated and will not lead to clinically significant decompression sickness.
  • Preprint
    Quantifying parameters of bottlenose dolphin signature whistles
    ( 2009-01-06) Esch, H. Carter ; Sayigh, Laela S. ; Wells, Randall S.
    Bottlenose dolphins (Tursiops truncatus) produce individually distinctive vocalizations called signature whistles, first described by Melba and David Caldwell (1965). The Caldwells observed that isolated, captive dolphins produced whistles with individually distinctive frequency contours, or patterns of frequency changes over time, and hypothesized that these whistles were used to transmit identity information (Caldwell and Caldwell 1965; Caldwell et al. 1990). Since the Caldwell’s work with isolated, captive dolphins, several studies have documented signature whistles in a variety of contexts, including free-swimming captive dolphins (e.g., Janik and Slater 1998; Tyack 1986), briefly restrained wild dolphins (e.g., Sayigh et al. 1990, 2007, Watwood et al. 2005), and free-ranging wild dolphins (e.g., Watwood 2003; Watwood et al. 2004, 2005; Buckstaff 2004; Cook et al. 2004). Janik and Slater (1998) demonstrated that signature whistles are used to maintain group cohesion, thus supporting the Caldwells’ hypothesis. Janik et al. (2006) verified experimentally that bottlenose dolphins respond to signature whistles produced by familiar conspecifics even after voice featured have been removed, reinforcing the notion that the contour of a signature whistle carries identity information.
  • Article
    The Sarasota Dolphin whistle database: a unique long-term resource for understanding dolphin communication.
    (Frontiers Media, 2022-08-04) Sayigh, Laela S. ; Janik, Vincent M. ; Jensen, Frants H. ; Scott, Michael D. ; Tyack, Peter L. ; Wells, Randall S.
    Common bottlenose dolphins (Tursiops truncatus) produce individually distinctive signature whistles that are learned early in life and that help animals recognize and maintain contact with conspecifics. Signature whistles are the predominant whistle type produced when animals are isolated from conspecifics. Health assessments of dolphins in Sarasota, Florida (USA) provide a unique opportunity to record signature whistles, as dolphins are briefly separated from conspecifics. Recordings were first made in the mid 1970’s, and then nearly annually since 1984. The Sarasota Dolphin Whistle Database (SDWD) now contains 926 recording sessions of 293 individual dolphins, most of known age, sex, and matrilineal relatedness. The longest time span over which an individual has been recorded is 43 years, and 85 individuals have been recorded over a decade or more. Here we describe insights about signature whistle structure revealed by this unique and expansive dataset. Signature whistles of different dolphins show great variety in their fundamental frequency contours. Signature whistle types (with ‘whistle type’ defined as all whistles visually categorized as sharing a particular frequency modulation pattern) can consist of a single stereotyped element, or loop (single-loop whistles), or of multiple stereotyped loops with or without gaps (multi-loop whistles). Multi-loop signature whistle types can also show extensive variation in both number and contour of loops. In addition, fundamental frequency contours of all signature whistle types can be truncated (deletions) or embellished (additions), and other features are also occasionally incorporated. However, even with these variable features, signature whistle types tend to be highly stereotyped and easily distinguishable due to the extensive variability in contours among individuals. In an effort to quantify this individual distinctiveness, and to compare it to other species, we calculated Beecher’s Information Statistic and found it to be higher than for any other animal signal studied so far. Thus, signature whistles have an unusually high capacity to convey information on individual identity. We briefly review the large range of research projects that the SDWD has enabled thus far, and look ahead to its potential to answer a broad suite of questions about dolphin communication.
  • Dataset
    Bottlenose dolphin mothers modify signature whistles in the presence of their own calves
    (Woods Hole Oceanographic Institution, 2023-05-22) Sayigh, Laela S. ; El Haddad, Nicole ; Tyack, Peter L. ; Janik, Vincent M. ; Jensen, Frants H. ; Wells, Randall S.
    Human caregivers interacting with children typically modify their speech in ways that promote attention, bonding and language acquisition. Although this “motherese,” or child-directed communication (CDC), occurs in a variety of human cultures, evidence among non-human species is very rare. We looked for its occurrence in a non-human mammalian species with long-term mother-offspring bonds that is capable of vocal production learning, the bottlenose dolphin (Tursiops truncatus). Dolphin signature whistles provide a unique opportunity to test for CDC in non-human animals, because we are able to quantify changes in the same vocalizations produced in the presence or absence of calves. We analyzed recordings made during brief catch-and-release events of wild bottlenose dolphins in waters near Sarasota Bay, Florida, USA, and found that females produced signature whistles with significantly higher maximum frequencies and wider frequency ranges when they were recorded with their own dependent calves vs. not with them. These differences align with the higher fundamental frequencies and wider pitch ranges seen in human CDC. Our results provide the first evidence in a non-human mammal for changes in the same vocalizations when produced in the presence vs. absence of offspring, and thus strongly support convergent evolution of motherese, or child-directed communication, in bottlenose dolphins. CDC may function to enhance attention, bonding and vocal learning in dolphin calves, as it does in human children. Our data add to the growing body of evidence that dolphins provide a powerful animal model for studying the evolution of vocal learning and language.
  • Article
    Surface and diving metabolic rates, and dynamic aerobic dive limits (dADL) in near‐ and off‐shore bottlenose dolphins, Tursiops spp., indicate that deep diving is energetically cheap
    (Wiley, 2023-04-05) Fahlman, Andreas ; Allen, Austin S. ; Blawas, Ashley ; Sweeney, Jay ; Stone, Rae ; Trainor, Robyn Faulkner ; Jensen, Frants H. ; McHugh, Katherine ; Allen, Jason B. ; Barleycorn, Aaron A. ; Wells, Randall S.
    High‐resolution dive depth and acceleration recordings from nearshore (Sarasota Bay, dive depth < 30 m), and offshore (Bermuda) bottlenose dolphins (Tursiops spp.) were used to estimate the diving metabolic rate (DMR) and the locomotor metabolic rate (LMR, L O2/min) during three phases of diving (descent, bottom, and ascent). For shallow dives (depth ≤ 30 m), we found no differences between the two ecotypes in the LMR during diving, nor during the postdive shallow interval between dives. For intermediate (30 m < depth ≤ 100 m) and deep dives (depth > 100 m), the LMR was significantly higher during ascent than during descent and the bottom phase by 59% and 9%, respectively. In addition, the rate of change in depth during descent and ascent (meters/second) increased with maximal dive depth. The dynamic aerobic dive limit (dADL) was calculated from the estimated DMR and the estimated predive O2 stores. For the Bermuda dolphins, the dADL decreased with dive depth, and was 18.7, 15.4, and 11.1 min for shallow, intermediate, and deep dives, respectively. These results provide a useful approach to understand the complex nature of physiological interactions between aerobic metabolism, energy use, and diving capacity.
  • Article
    Modeling tissue and blood gas kinetics in coastal and offshore common bottlenose dolphins, Tursiops truncatus
    (Frontiers Media, 2018-07-17) Fahlman, Andreas ; Jensen, Frants H. ; Tyack, Peter L. ; Wells, Randall S.
    Bottlenose dolphins (Tursiops truncatus) are highly versatile breath-holding predators that have adapted to a wide range of foraging niches from rivers and coastal ecosystems to deep-water oceanic habitats. Considerable research has been done to understand how bottlenose dolphins manage O2 during diving, but little information exists on other gases or how pressure affects gas exchange. Here we used a dynamic multi-compartment gas exchange model to estimate blood and tissue O2, CO2, and N2 from high-resolution dive records of two different common bottlenose dolphin ecotypes inhabiting shallow (Sarasota Bay) and deep (Bermuda) habitats. The objective was to compare potential physiological strategies used by the two populations to manage shallow and deep diving life styles. We informed the model using species-specific parameters for blood hematocrit, resting metabolic rate, and lung compliance. The model suggested that the known O2 stores were sufficient for Sarasota Bay dolphins to remain within the calculated aerobic dive limit (cADL), but insufficient for Bermuda dolphins that regularly exceeded their cADL. By adjusting the model to reflect the body composition of deep diving Bermuda dolphins, with elevated muscle mass, muscle myoglobin concentration and blood volume, the cADL increased beyond the longest dive duration, thus reflecting the necessary physiological and morphological changes to maintain their deep-diving life-style. The results indicate that cardiac output had to remain elevated during surface intervals for both ecotypes, and suggests that cardiac output has to remain elevated during shallow dives in-between deep dives to allow sufficient restoration of O2 stores for Bermuda dolphins. Our integrated modeling approach contradicts predictions from simple models, emphasizing the complex nature of physiological interactions between circulation, lung compression, and gas exchange.