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Soaring Styles of Extinct Giant Birds and Pterosaurs bioRxiv preprint doi: https://doi.org/10.1101/2020.10.31.354605; this version posted November 1, 2020. The copyright holder for this preprint (which was not certified by peer review) is the author/funder. All rights reserved. No reuse allowed without permission. 1 2 Soaring styles of extinct giant birds and pterosaurs 3 4 5 Authors: 6 Yusuke Goto1*, Ken Yoda2, Henri Weimerskirch1, Katsufumi Sato3 7 8 Author Affiliation: 9 1Centre d’Etudes Biologiques Chizé, CNRS – Université de la Rochelle, 79360 Villiers 10 En Bois, France. 11 2Graduate School of Environmental Studies, Nagoya University, Furo, Chikusa, 12 Nagoya, Japan. 13 2Atmosphere and Ocean Research Institute, The University of Tokyo, Chiba, Japan. 14 15 16 Corresponding Author: 17 Yusuke Goto 18 Centre d’Etudes Biologiques Chizé, CNRS – Université de la Rochelle, 79360 Villiers 19 En Bois, France. 20 e-mail: [email protected] 21 TEL: +814-7136-6220 22 23 1 bioRxiv preprint doi: https://doi.org/10.1101/2020.10.31.354605; this version posted November 1, 2020. The copyright holder for this preprint (which was not certified by peer review) is the author/funder. All rights reserved. No reuse allowed without permission. 24 Abstract 25 Extinct large volant birds and pterosaurs are thought to have used wind-dependent soaring 26 flight, similar to modern large birds. There are two types of soaring: thermal soaring, used by 27 condors and frigatebirds, which involves the use of updrafts to ascend and then glide 28 horizontally; and dynamic soaring, used by albatrosses, which involves the use of differences in 29 wind speed with height above the sea surface. However, it is controversial which soaring styles 30 were used by extinct species. In this study, we used aerodynamic models to comprehensively 31 quantify and compare the soaring performances and wind conditions required for soaring in two 32 of the largest extinct bird species, Pelagornis sandersi and Argentavis magnificens (6–7 m 33 wingspans), two pterosaur species, Pteranodon (6 m wingspan) and Quetzalcoatlus (10 m 34 wingspans), and extant soaring birds. For dynamic soaring, we quantified how fast the animal 35 could fly and how fast it could fly upwind. For thermal soaring, we quantified the animal’s sinking 36 speed circling in a thermal at a given radius and how far it could glide losing a given height. 37 Consistently with previous studies, our results suggest that Pteranodon and Argentavis used 38 thermal soaring. Conversely, the results suggest that Quetzalcoatlus, previously thought to have 39 used thermal soaring, was less able to ascend in updrafts than extant birds, and Pelagornis 40 sandersi, previously thought to have used dynamic soaring, actually used thermal soaring. Our 41 results demonstrate the need for a comprehensive assessment of performance and required 42 wind conditions when estimating soaring styles of extinct flying species. 43 44 Introduction 45 Flying animals have evolved a wide range of body sizes. Among them, there have been 46 incredibly large species of birds and pterosaurs (Fig. 1). Pelagornis sandersi and Argentavis 47 magnificens are the largest extinct volant birds. Their estimated wingspans reached 6–7 m (1– 48 4), twice as large as that of the wandering albatross, the extant bird with the longest wingspan 49 (Table 1). Several large species of pterosaurs appeared in the Cretaceous period. Pteranodon, 50 presumably the most famous pterosaur, is estimated to have had a wingspan of 6 m (Table 1) 51 (5). The azhdarchids are one of the most successful Cretaceous pterosaur groups and include 52 many large species with wingspans of approximately 10 m (Table 1) (6–9). Quetzalcoatlus 53 northorpi, one of the azhdarchid species, is regarded as one of the largest flying animals in 54 history. 55 Discoveries of these giant winged animals have fascinated paleontologists and 56 biologists for over a century. Their flight ability has been of great interest because their huge 2 bioRxiv preprint doi: https://doi.org/10.1101/2020.10.31.354605; this version posted November 1, 2020. The copyright holder for this preprint (which was not certified by peer review) is the author/funder. All rights reserved. No reuse allowed without permission. 57 size must significantly affect their flight. With increasing size, flapping becomes more costly; the 58 power required to fly increases faster than the power muscles can produce. Two main 59 arguments are often debated about the flight of extinct giants, both stemming from this physical 60 constraint: the first is about whether they were able to take off (4, 10–12); the second is about 61 what kind of soaring flight style they employed (1, 4, 13–15). This study focuses on the latter 62 debate. Due to the high cost of flapping, large extant birds prefer to fly utilizing wind energy, that 63 is, they prefer to soar. It is presumed that extinct large animals also employed soaring flight as 64 their primary mode of transportation (1, 4, 14). For example, in Quetzalcoatlus, flapping flight 65 might have required anaerobic exercises that were difficult to sustain for long periods of time 66 (14). 67 There are two main soaring flight styles among extant birds: dynamic soaring and 68 thermal soaring (16). In dynamic soaring, birds extract flight energy from wind shear—the 69 vertical gradient in horizontal wind speed over the ocean (Fig. 2A). In extant birds, seabirds 70 (e.g., albatrosses, shearwaters, and petrels) employ this soaring style and can routinely travel 71 hundreds of kilometers per day over the sea. In thermal soaring, birds first fly circling in warm 72 rising-air columns (thermals). They climb to a substantial height and then glide off in the desired 73 direction while losing their height (Fig. 2C-E). By repeating this up-down process, birds travel 74 over vast distances. Various terrestrial bird species (e.g., vultures, eagles, and storks) and 75 seabirds (e.g., frigatebirds and pelicans) employ thermal soaring. 76 This study investigated which soaring styles were employed by four large extinct 77 species, that is, Pelagornis sandersi, Argentavis magnificens, Pteranodon, and Quetzalcoatlus. 78 Their soaring capabilities were quantified using two factors. The first is performance, for 79 example, the available speed and efficiency of soaring. The second is the minimum wind speed 80 required for sustainable soaring flight. 81 For thermal soaring, performance was evaluated in the gliding and soaring up 82 phases. In the gliding phase, a bird’s performance is its glide ratio, which is the ratio of distance 83 a bird traverses to the height the bird loses to cover that distance, i.e., (glide ratio) = (horizontal 84 speed)/(sinking speed) (Fig. 2E). A bird with a higher glide ratio can traverse a longer distance 85 for the same amount of height lost as a bird with a lower glide ratio. Plotting a curve known as a 86 ‘glide polar’ is the conventional method to quantify glide ratio and the associated speed in 87 artificial gliders and birds (16). This curve is a plot of the sinking speed against horizontal speed 88 when a bird glides in a straight line. We can determine the flyer’s highest glide ratio and the 89 associated travel speed by finding the line that passes the origin and tangents of the glide polar. 3 bioRxiv preprint doi: https://doi.org/10.1101/2020.10.31.354605; this version posted November 1, 2020. The copyright holder for this preprint (which was not certified by peer review) is the author/funder. All rights reserved. No reuse allowed without permission. 90 91 92 Fig. 1. A size comparison and soaring styles of extinct giant birds (Pelagornis sandersi 93 and Argentavis magnificens), pterosaurs (Pteranodon and Quetzalcoatlus), the largest 94 extant dynamic soaring bird (wandering albatross), the largest extant thermal soaring 95 terrestrial bird (California condor), the largest extant thermal soaring seabird 96 (magnificent frigatebird), and the heaviest extant volant bird (kori bustard). The icons 97 indicate dynamic soarer, thermal soarer, and poor soarer and summarize the main 98 results of this study. The red arrows indicate the transition from a previous expectation 99 or hypothesis to the knowledge updated in this study. 4 bioRxiv preprint doi: https://doi.org/10.1101/2020.10.31.354605; this version posted November 1, 2020. The copyright holder for this preprint (which was not certified by peer review) is the author/funder. All rights reserved. No reuse allowed without permission. Species Mass Wingspan Wing Aspect Wing Ref (kg) (m) area (m2) ratio loading Pelagornis sandersi 21.8, 6.06, 6.13, 2.45 13.0, 51.0 (1) and 6.40 and ~ 14.0, ~ 40.1 7.38 4.19 and 15.0 161 Argentavis magnificens 70.0 7.00 8.11 6.04 84.7 (4) Extinct Pteranodon 36.7 5.96 1.99 17.85 181 (5) Quetzalcoatlus 259 9.64 11.36 8.18 224 (5) wandering albatross 8.64 3.05 0.606 15.4 140 (48)* (Diomedea exulans) black-browed albatross 3.55 2.25 0.376 13.4 87.5 (49)* (Thalassarche melanophris) white-chinned petrel 1.37 1.4 0.169 11.6 79.5 (50) Dynamicsoaring (Procellaria aequinoctialis) magnificent frigatebird 1.52 2.29 0.408 12.8 36.5 (17) (Fregata magnificens) California condor 9.5 2.74 1.32 5.7 70.6 (4) (Gymnogyps californianus) brown pelican 2.65 2.10 0.45 9.80 57.8 (17) (Pelecanus occidentalis) Thermalsoaring black vulture 1.82 1.38 0.327 5.82 54.6 (17) (Coragyps atratus) white stork 3.4 2.18 0.540 7.42 61.8 (4) (Ciconia ciconia) kori bustard 11.9 2.47 1.06 5.76 110 (16) - (Ardeotis kori) Non Soaring 100 101 Table 1.
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