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Escape Behaviour of Steamer Ducks BRADLEY C

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Escape Behaviour of Steamer Ducks BRADLEY C Escape behaviour of Steamer Ducks BRADLEY C. LIVEZEY and PHILIP S. HUMPHREY Introduction ing, partially submerged swimming, ‘rest­ ing’ with erect posture, and ‘floating’ most­ Steamer ducks Tachyeres comprise four ly submerged; steaming turbulent, spray- species of benthic divers limited in distribu­ generating surface locomotion powered by tion to southern South America (Murphy feet and wings; diving subaquatic locomo­ 1936; Hum phrey & Thompson 1981): tion accomplished by foot paddling, often Flying Steamer Duck T. patachonicus of aided by strokes of half-folded wings; hid­ marine coastlines and freshwater lakes of ing squatting motionless on land, typically Argentina, Chile, and the Falkland with head and neck extended anteriorly; Islands; Magellanic Flightless Steamer flight limited to but not universal in T. Duck T. pteneres of coastal Chile and the patachonicus (Hum phrey & Livezey 1982). Magellanic Straits; Falkland Flightless Death feigning, possibly peculiar to T. Steamer Duck T. brachypterus of marine pteneres (Humphrey et al. 1970), was not coasts of the Falkland Islands; and the observed. recently described White-headed Flightless Contingency tables were designed in Steamer Duck T. leucocephalus of coastal accord with Cochran (1954). Analyses of Chubut, Argentina. They are named for sequences of escape behaviour assumed their habit of ‘steaming’, a rapid, spray- each behaviour was not immediately re­ generating surface locomotion involving peatable, i.e., termination of one escape wings and feet. The escape behaviours of mode was defined by the beginning of a steamer ducks comprise steaming, diving, different behaviour. Statistical independ­ typical swimming, concealed swimming or ence of such behavioural transitions was ‘sneaking’, running and hiding on land, not tested rigorously because of large death-feigning, and in one species, flight differences in observed frequencies of for escape. different escape behaviours (Lemon & The probable relevance of these diverse Chatfield 1971). escape behaviours to the taxonomy, ecolo­ gy, and field identification of steamer ducks prompted us to study these be­ Results haviours during recent field work designed to clarify the Systematics of the genus. Frequencies of escape behaviours T. pteneres, T. patachonicus, and T. Study sites and methods leucocephalus were encountered with diffe­ rent frequencies (X 2 = 20-1, /■’<0-005) on We studied three species of steamer ducks land and water. Pair-wise comparisons of in Argentina during December 1980 to species indicated that T. patachonicus and February 1981: T. leucocephalus at Puerto T. pteneres were similar (X 2 = 0-4), and in Melo, Chubut; T. patachonicus at Puerto both species à of the initial encounters were Deseado, Santz Cruz; and T. patachonicus of birds on land and | on water (Table 1). and T. pteneres at Ushuaia, Tierra del T. leucocephalus differed markedly Fuego. Data recorded for pursued birds (P < 0-005) from the other two species were: species and sex; location of bird(s) studied and was found on land and water when encountered; sequence of behaviour with equal frequency. used; flock size; minimum distance of Apart from flight, the three species dif­ bird(s) from boat; wind conditions; and fered (X 2 = 129-6, P<0-001) in frequen­ occurrence of gunfire. Unless otherwise cies of other escape behaviours used (Fi­ noted, all observations of escape behaviour gure 1). Closer examination revealed dif­ were made from boats. ferences in behaviour between T. patacho­ Behaviours used during escape were nicus and T. leucocephalus (X2= 103-2, classified for statistical analyses as: swim ­ P < 0-001) and between T. leucocephalus ming surface locomotion powered by feet and T. pteneres (X 2 = 106-3, P<0-001). T. alone, includes typical ‘swimming’ with pteneres and T. patachonicus were similar head and neck erect, ‘sneaks’ or conceal- in observed frequencies of steaming, swim­ 12 Wildfowl 33 (1982): 12—16 Escape behaviour of Steamer Ducks 13 ming, diving, and running. T. leucocepha­ loadings and the resultant difficulty of lus differed from the other two species take-off (Humphrey & Livezey 1982). In primarily in its pronounced tendency to addition, 10 specimens of T. patachonicus swim and steam and lower frequencies of were flightless because of moult of re­ running and diving (Figure 1). miges. Birds in wing moult were usually Only 11% of T. patachonicus flew when encountered in flocks and, like the flight­ pursued. This reflects, in part, heavy wing less species, resorted to swimming, steam- A T. pteneres 173 acts ; n = 79 O CO LU 40 cn B T. leucocephalus c w 9 6 9 acts ; n = 4 8 6 ZD 30 w 03 13 20 - > TD C 10 C CD O 03 CL 30 - T. patachonicus 246 acts, n =107 20 10 SWIM STEAM DIVE RUN HIDE FLY Figure 1. Relative frequencies of escape modes observed at three localities in Argentina during December 1980-February 1981. 14 Bradley C. Livezey and Philip S. Humphrey ing, diving, running and hiding for escape. swimming. In T. leucocephalus swimming Proportions of swims classed as ‘sneaks’ was also generally followed by steaming, differed (X 2 = 27-9, F <0-005) among the but steaming, if terminated, most often three species (Table 1). T. pteneres and T. gave rise to more swimming. Also, the patachonicus both sneaked relatively fre­ relatively land-loving T. leucocephalus quently (21% and 14% of all swims, escaped as frequently by running followed respectively) ; T. leucocephalus rarely by swimming. sneaked (4%). Sequences of escape behaviour initiated Table 1. Mean numbers of acts, behavioural diversities (H'), percentages of swims classed as ‘sneaks’, and percentages of birds initially encountered on land and on water. Statistic T. pteneres T. patachonicus T. leucocephalus Mean number of acts 2-4 3-1 2-2 H ' (acts) 1-45 1-44 1-13 Swims classed as ‘sneaks’ (%) 21 14 4 Initially on land (%) 33 29 52 Initially on water (%) 67 71 48 T. pteneres and T. patachonicus were on land differed between the otherwise also more similar to each other than to T. similar T. pteneres and T. patachonicus, leucocephalus in diversity of escape be­ although neither species was typically first haviour. Both numbers of acts and the encountered ashore. Flightless T. pteneres Shannon-Weaver index (H') suggested low always ran to water and immediately behavioural diversity in T. leucocephalus steamed. Beached T. patachonicus most compared to the other two species studied frequently ran to almost immediate flight (Table 1). (54%); running was less commonly fol­ lowed by swimming (25%), steaming Behavioural sequences (11%) or diving (11%). Tabulation of frequencies of changes from one escape mode to another, provides further evidence that T. leucocephalus dif­ Effects o f flock size fered from T. patachonicus and T. pteneres (Table 2). In the latter two species, swim­ D ata for T. pteneres showed no significant ming gave rise more frequently to steam­ differences in escape behaviour among ing, steaming to diving, and diving to more single birds, birds in pairs, or flocks of Table 2. Two-act sequential analyses of escape behaviour. Cells contain number (%) of various escape modes that were followed by other escape modes. Second First act Species act Swim Steam Dive Run Fly Total T. pteneres Swim — 6(27) 3(60) 0(0) — 9 (64) Steam 37 (69) — 2(40) 15 (100) — 54 Dive 12 (22) 15 (68) — 0 (0 ) — 27 Run 5(9) 1(5) 0(0) —— 6 Total 54 22 5 15 — 96 T. leucocephalus Swim — 13 (72) 25 (96) 126 (68) — 164 (367) Steam 276 (87) — 1(4) 58 (31) — 335 Dive 34(11) 2(11) — 2 (1 ) — 38 Run 6 (2 ) 3(17) 0 (0 ) —— 9 Total 316 18 26 186 — 546 T. patachonicus Swim — 4(10) 23 (92) 7(25) 3 (33) 37 (98) Steam 53 (52) — 0 (0 ) 3(11) 2 (22) 58 Dive 22 (22) 27 (67) — 3(11) 4 (44) 56 Run 9(9) 6(15) 1 (4) — 0 (0 ) 16 Fly 18 (18) 2(5) 1(4) 15 (54) — 36 Total 102 39 25 28 9 203 Escape behaviour of Steamer Ducks 15 three or more. However, both T. patacho­ as a rapid, target-obscuring method for nicus (X l = 33-3, P<0-005) and T. escaping underwater predators, especially leucocephalus (X - = 2 5 -l, P < 0-005) in flightless species. showed different escape behaviour de­ Diving is probably employed as an pending on flock size. Lone birds and pairs escape behaviour by steamer ducks mostly of T. patachonicus hid, dived, and flew to avoid aerial predators. The threat of more frequently than conspecifics in larger aerial attack by Kelp Gulls Larus domini- flocks, but larger flocks were somewhat canus. Skuas Catharacta skua, and Giant more prone to steam. Steaming was also Petrels Macronectes giganteus is substantial more frequent in larger flocks of T. for downy young of Tachyeres (Pettingill leucocephalus. 1965). Diving ability is acquired early, and we observed short feeding dives in class-I ducklings of T. pteneres and T. patachoni­ Effects o f nearness o f boat cus. Foxes Dus icy on spp. are predators of Frequency data revealed no differences in adult T. patachonicus and T. pteneres on escape behaviour related to distance of mainland South America (Reynolds in approach. However, when we surprised Lowe 1934) and occurred formerly on the feeding steamer-ducks at close range, the Falklands (Clayton 1776). The utility of birds typically dived to escape us. When running to avoid canids is obvious, espe­ birds detected us at a distance they gener­ cially for flightless species during nesting. ally steamed, swam, or flew away. This The usefulness of death-feigning is not disagreement between our impression in clear in that most terrestrial predators are the field and the statistical results may be also scavengers. because escape behaviours were affected Flightlessness in waterfowl is generally by when the ducks detected us rather than associated with island faunas, presumably simply the closeness of our approach.
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