Penguin (Eudyptes Chrysocome Moseleyi)
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Polar Biol 02000) 23: 812±816 Ó Springer-Verlag 2000 ORIGINAL PAPER Charles M. Drabek á Yann Tremblay Morphological aspects of the heart of the northern rockhopper penguin Eudyptes chrysocome moseleyi ): possible implication in diving behavior and ecology? Accepted: 20 May 2000 Abstract We compared the heart morphology of the Kooyman and Ponganis 1997; Schreer and Kovacs 1997; small, deep-diving northern rockhopper penguin to the Kooyman and Ponganis 1998). Drabek 01989) published hearts of small, shallow-diving and large, deep-diving the ®rst suggestion that the heart morphology of pen- penguin species. The rockhopper penguin had a heart guins might play a role in their diving capabilities. He larger than expected for its body mass, and its heart noted that the penguin heart size relative to body mass weight/body weight was signi®cantly greater than in the 0heart ratio) was larger than expected for birds and that larger Ade lie penguin. We found the rockhopper's right the right ventricle of the emperor penguin 0Aptenodytes ventricle weight/heart weight to be signi®cantly greater forsteri) was more muscular in proportion to both the than this relationship in both the larger chinstrap and left ventricle and the heart than that of the chinstrap Ade lie penguins. The relationship of the right to left 0Pygoscelis antarctica) and Ade lie penguins 0P. adeliae). ventricular weights in the rockhopper heart is not dif- The long, deep diving capabilities of the emperor pen- ferent to that of the large, deepest-diving emperor pen- guin have long been documented 0Kooyman et al. 1971), guin. A larger heart in the rockhopper penguin might be and Drabek 01989) suggested the heart morphology related to its diving behavior and ecology if it contrib- might contribute to diving eciency during feeding by utes to diving eciency during foraging by increasing increasing lung perfusion to decrease surface recovery lung perfusion during surface recovery. This would lead times between dives. In a subsequent study, Drabek to decreased surface time. 01997) compared and contrasted the heart morphology of these deep-diving penguin species with that of the little penguin 0Eudyptula minor). The little penguin rep- resents the smallest of all the penguin species and is known to be a brief, shallow diver 0Bethge et al. 1997). Introduction Drabek reported that the little penguin heart is not larger than expected for its body mass and that the right Although it has been three decades since Kooyman et al. ventricle relative to the heart and left ventricle is sig- 01971) reported on a path®nding investigation of deter- ni®cantly less muscularized than right ventricles in the mining the behavior of freely diving penguins, it has only larger, deeper-diving penguin species. Tremblay et al. been in the last 10 years that biologists have used similar 01997) reported the ®rst data on maximum diving depths and even more sophisticated techniques 0e.g., Kooyman attained by the northern rockhopper penguin, 0Eudyptes and Kooyman 1995) to elucidate the diving capabilities chrysocome moseleyi), at Amsterdam Island. They found of most of the 18 penguin species. Excellent reviews of that although it is the third smallest 0to the little penguin these studies are numerous 0Burger 1991; Wilson 1995; and Galapagos penguin) 02.3 kg), it is capable of diving deeply 0168 m). This is nearly the depth 0175 m) attained by the Ade lie penguin 0Whitehead 1989), a penguin C. M. Drabek 0&) twice the body mass of the rockhopper. More recently, Biology Department, Whitman College, Cherel et al. 01999) showed that the most remarkable Walla Walla, Washington 99362, USA feature of diving behavior in northern rockhopper pen- e-mail: [email protected] guins was the high percentage of the time spent diving Y. Tremblay 069% of their time at sea), which was mainly due to a Centre d'Etudes Biologiques de Chize , high dive frequency 044 dives per hour at sea). U.P.R. 1934 du Centre National de la Recherche Scienti®que, The purpose of this study was to describe the heart 79360 Villiers-en-Bois, France and ventricle sizes of the northern rockhopper and to 813 make comparisons with the hearts of the smaller, shal- greater 0P < 0.001) than in both the Ade lie and little low-diving little penguin and with the large, deep-diving penguins 0Fig. 1). In the rockhopper, the weight of the Antarctic penguins. right ventricle relative to the weight of the left ventricle is no dierent from this relationship in the emperor but signi®cantly greater 0P < 0.001) than in the little, chinstrap, and Ade lie penguins. The weight of the right Materials and methods ventricle relative to the heart weight in the rockhopper penguin is signi®cantly less 0P < 0.001) than this rela- The hearts of seven male and four female adult northern rock- tionship in the emperor and signi®cantly greater hopper penguins 0E. chrysocome moseleyi) from Amsterdam Island 0P < 0.001) than in the little, chinstrap, and Ade lie 037°50¢S, 77°31¢E) in the southern Indian Ocean were weighed and measured. The hearts were removed from autopsied penguins that penguins. had died from natural causes 0mainly predation) near the colonies. The hearts were initially ®xed in 10% formalin and stored in 30% isopropyl alcohol until measurements were taken. Since the hearts were not weighed upon removal, there was no check made of any in¯uence of the preservatives. This was the same protocol followed Discussion by Drabek 01989, 1997). The brachiocephalic artery and pericar- dium were removed and excess alcohol was wicked away before The general anatomical aspects of the cardiovascular the weights were recorded on an Ohaus triple-beam balance to the morphology of penguins have been described in several nearest 0.1 g. Helios needle calipers were used to measure the length of the heart from the apex to the ori®ce of the pulmonary investigations 0Watson 1883; Pycraft 1907; Glenny 1944, valve. The circumference and width were measured at the greatest 1947; Andrews and Andrews 1976). Drabek 01989) was breadth of the ventricles. Measurements were made to the nearest the ®rst to publish heart and ventricular weight data for 0.1 mm. The left ventricle weight included the interventricular any penguin species and the ®rst to suggest that the heart septum. The dierences between the means were compared using morphology might in¯uence diving capabilities. Drabek's Student's t-test and ANOVA. The 5% level of probability was accepted as indicating statistical signi®cance. 01997) heart study of the little penguin elucidated the dierences between the heart anatomy of a shallow- diving penguin and the hearts of penguins known to be deeper divers. He felt the proportionately larger right Results ventricle contributes to diving eciency by increasing lung perfusion during surface recovery. Table 1 shows the heart measurements for the rock- Tremblay et al. 01997) recorded a dive depth of hopper penguin in comparison to those of the little, 168 m for the northern rockhopper and compared their emperor, chinstrap, and Ade lie penguins. The length/ data to the allometric equation 0y 47.6 + 18.0x) used width relationship of the rockhopper's heart was sig- by Wilson 01995) to relate maximum diving depth 0x)to ni®cantly dierent from this relationship in the emper- a penguin's body mass 0y). The 2.66-kg rockhoppers or's heart 0F4,57 5.36, P < 0.001). The rockhopper's Tremblay et al. studied would be predicted to dive to a heart is proportionately wider. The heart weight relative maximum of only 95 m, and Wilson's equation indicates to the body weight of the rockhopper penguin is no that only a penguin 3 times heavier 06.69 kg) would have dierent from that of the chinstrap but signi®cantly the ability to dive to 168 m. Even by considering usual Table 1 The heart parameter data for the rockhopper Parameter Little penguin Emperor Chinstrap Ade lie Rockhopper penguin are presented as 0N 27) 0N 8) 0N 8) 0N 8) 0N 1) means standard error, with ranges in parentheses. The little Body weight 0g) 1188 56 ± 3900 160 4400 170 2340 156 penguin data are from Drabek 0780±1940) 03200±4500) 03700±4900) 01400±2950) 01997) and the emperor, chin- Heart mass 0g) 9.9 0.4 174.8 5.4 42.4 1.1 37.5 1.4 24.7 1.19 strap, and Ade lie data are from 06.6±14.1) 0154.9±198.0) 038.6±48.0) 029.0±42.6) 017.8±31.3) Drabek 01989) 0H heart mass; Length 0mm) 28.9 0.5 84.9 2.4 50.6 1.1 50.7 0.8 38.8 0.8 B body weight; L heart length; 023.8±36.9) 074.0±91.9) 047.2±56.0) 047.8±55.7) 032.5±41.8) W heart width; RV right ven- Width 0mm) 23.9 0.4 64.7 1.1 45.5 0.9 46.2 0.9 33.6 1.11 tricle weight; LV left ventricle 020.2±30.7) 061.9±71.1) 041.0±49.0) 042.5±49.8) 027.8±38.2) weight) Circumference 72.3 1.3 180.3 3.5 116.9 1.8 115.6 1.8 97.3 1.65 0mm) 060.0±89.0) 0163.0±190.0) 0110.0±123.0) 0107.0±120.0) 086±104) H/B 0%) 0.86 0.04 ± 1.07 0.03 0.85 0.03 1.07 0.05 00.50±1.16) 00.90±1.20) 00.73±1.02) 00.86±1.37) L/W 1.22 0.02 1.31 0.03 1.12 0.03 1.10 0.03 1.17 0.04 01.06±1.44) 01.18±1.43) 00.96±1.25) 01.00±1.24) 00.87±1.3) RV/H 0%) 6.7 0.46 21.26 1.21 9.3 0.48 10.9 0.93 15.97 0.688 02.9±12.2) 015.9±46.1) 07.7±11.8) 07.8±15.5) 012.78±19.58) RV/LV 0%) 17.9 1.14 37.36 2.41 15.8 0.88 18.9 2.15 33.52 1.1 06.1±28.1) 026.9±46.1) 012.4±19.2) 011.2±27.6) 027.58±38.54) 814 would be able to dive to 168 m 0i.e., a 6.69-kg bird, using the allometric equation from Wilson 1995) would be predicted to have a 46.0-g heart.