Nest Insulation and Incubation Constancy of Arctic Geese
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Nest insulation and incubation constancy of arctic geese STEVEN C. THOMPSON and DENNIS G. RAVELING Introduction as better insulated nests may be more conspicuous (M 0 ller 1984). Proper and timely development of avian In geese (Anserini), only females incu embryos depends upon maintenance of bate. Nests are constructed in shallow incubation temperatures within relatively depressions on the ground which are usually narrow limits (e.g. Lundy 1969; Drent lined with dry vegetation collected from the 1975). For many birds, especially species area immediately around the nest (Owen nesting in cold environments, insulation of 1980). Finally, the nest is lined with down nests conserves heat transferred to eggs and and a few contour feathers from the breast minimises the cost of incubation to parent of the female. When leaving the nest, geese birds. Various species of birds nesting at cover their eggs with a blanket of material high elevations or latitudes have bulkier or pulled from the lining of the nest. better insulated nests than close relatives Female geese rely heavily on endogenous nesting at lower elevations or latitudes reserves to sustain incubation (e.g. Ankney (Pearson 1953; Corley Smith 1969; Collias and Maclnnes 1978; Raveling 1979a, b) and and Collias 1971; M0ller 1984). Furth are extremely attentive to their nests ermore, the amount of nest insulation may (Cooper 1978; Aldrich and Raveling 1983; be negatively correlated with nest attentive Thompson and Raveling 1987). There is, ness (White and Kinney 1974). however, considerable variation in patterns In many species, the incubating bird of incubation. For example. Emperor leaves the nest for a period (a recess) to G eese Anser canagicus, Cackling Canada obtain food and/or engage in other main G eese Branta canadensis minima and Black tenance activities. Nest insulation can B rant B. bernicla nigricans all nest on the reduce the rate of heat loss from eggs and Yukon-Kuskokwim Delta of Alaska, but thereby increase the length of recess with exhibit a two-fold difference in recess out causing harm to embryos, and minimise lengths and a thirteen-fold difference in the time and energy expended to rewarm recess frequencies, resulting in a twenty the eggs. Species with single-sex incubation fold difference in daily recess time between tend to have large, well insulated nests the least and most attentive species (Table compared with species in which both sexes 1). These differences are thought to be incubate on a rotational basis (O’Conner primarily related to the abilities of the 1978). It may therefore be expected that different-sized goose species to repel dif variation in the insulation of nests among ferent types of predators (Thompson and similar species in a given environment may Raveling 1987). be related to differences in incubation There are evident differences in nest rhythms. Conversely, predation may limit structure among Emperor Geese, Cackling nest insulation by selecting for cryptic nests G eese and B rant (Figure 1). B rant nests Tabic I. Nest attentiveness of Emperor Geese, Cackling Canada Geese and Black Brant, from, respectively, Thompson and Raveling (1978), Aldrich (1983) and Thompson (unpubl.). Total % time Modal Mean Mean Mean incubating recess recess recesses recess Species N (range) length (min) length (min) per day time/d (min) Emperor Goose 11 99.5 8 13 0.5 7 (99.1 - 99.7) Cacking Canada 12 93.6 15 26* 3.5* 92 Goose (89.1 - 96.6) Black Brant 4 89.6 22 22 6.7 148 (87.3 - 94.9) ‘Derived from published regression. 124 W ildfow l 39 ( 19NK): 124-132 Nest Insulation in arctic geese 125 Figure 1. Typical natural nests illustrating eggs exposed and covered of Emperor Goose (a,b). Cackling Goose (c,d) and Brant (e,f). 126 Steven C. Thompson and Denis G. Raveling contain by far the greatest quantities of Thermistor probes (YS1 model 400) were down and feathers, whereas Emperor implanted in 3 eggs of each clutch by drilling Goose nests contain the least amounts; the a hole down the long axis to the centre of intermediate amounts in Cackling Geese each egg, inserting the probe, and sealing nests differ more from Brant than from it in place with melted paraffin. Emperor Geese. Experiments were conducted in a The purpose of this study was to compare coldroom at approximately 6°C, providing differences in nest insulation quality of the ambient thermal conditions typical of those three species by measuring the cooling rates at Kokechik Bay (Eisenhauer and Kirk of eggs in their nests, and to relate those patrick 1977). Each nest was arranged in a differences to the incubation constancy of 5 cm deep depression in moist sand of the each species. The possible role of predation approximate natural dimensions reported as an agent selecting for differences in by Mickelson (1975). Although the sand conspicuousness and vulnerability of the was different from the material available to nests was also considered. geese on the Delta, it provided a standardi sed substrate. Clutches were heated in a drying oven at Methods approximately 40°C until internal egg tem peratures reached approximately 38°C, Six nests each of Emperor Geese, Cackling close to natural incubation temperatures Geese and Black Brant were collected, at (Drent 1975). The clutch was then immedi Kokechik Bay on the Yukon-Kuskokwim ately placed in a nest and covered with the Delta, Alaska, after depredation late in nest lining from the edges of the nest bowl. incubation or after hatch, to be certain that Cooling of eggs was then monitored for 30 little or no more material would have been minutes using strip chart recorders con added. Only nests judged to be relatively nected to the thermistors. Air temperature undisturbed by wind or predators were in the coldroom was simultaneously mea taken, and returned to the University of sured by a probe placed 10 cm from the California, Davis (UCD), where they were edge of the nest. Individual eggs of each weighed to the nearest gram on an electric species were placed in approximately the balance. same positions (Figure 2) in all tests to Eggs from the three goose species were minimise the effect of differences among emptied and then refilled with paraffin. This eggs. These positions simulated those in gave eggs with heating and cooling prop natural nests. Each of the 6 nests of each erties similar to those of natural eggs species was tested to discern the effects of (Schw artz et al. 1977), but approximately nest insulation relative to the combined 15% less in mass. The paraffin tended to effect on cooling of different clutch and egg expand upon heating and crack the shells sizes. Thus, each Emperor Goose nest was of eggs. Despite this problem, eggs retained tested containing, in random order, the their original mass and shape and provided clutches of Emperor Goose eggs, Cackling valid results for the comparative purposes Goose eggs, and Brant eggs. Each nest-egg of this study. Clutches were assembled of combination was tested in “calm” air (no 6 eggs of Emperor Geese, 5 of Cackling wind speed detectable with a Dwyer ane Geese, and 4 of Brant. These were typical mometer) and with a 10 km/h air flow over of natural clutches and eggs (Table 2). the nest (“windy”). This was provided by Table 2. Sizes and masses of praffin-filled eggs in experimental clutches. All eggs Thermistor-probed eggs Mean Mean Mean Mean Mean Mean egg egg egg egg egg egg Species n mass (g) length (mm) width (mm) n mass (g)* length (mm) width (mm) Emperor Goose 6 103 81 52 3 102 81 52 Cackling Goose 5 86 75 50 3 86 76 50 Black Brant 4 75 71 48 3 75 70 48 * Before insertion of thermistor probes. Nest Insulation in arctic geese 127 O BLACK BRANT o o EMPEROR CACKLING GOOSE GOOSE Figure 2. Arrangements of paraffin-filled eggs of Emperor Geese, Cackling Geese and Brant in experimental nests. Numbered eggs are those with thermistors. Arrows indicate direction of air flow over nests under windy conditions. a standard household box fan placed 50 cm (243.2±24.3 g) or Cackling Goose nests from the nest edge. Each nest was left (193.2+42.0 g) (PcO.OOl), but differences empty of eggs for at least an hour between in mass between the nests of the latter two tests, adequate for nest and substrate to species were not significant (P>0.05). The return to the ambient temperature. differences in mass were primarily due to Differences among values of initial egg the varying amounts of vegetation present temperatures, egg cooling, coldroom tem (Figure 1). peratures and nest masses were investigated Mean coldroom temperature during tests by Analysis of Variation (ANOVA), and was 6.41±0.04°C and did not vary signifi significance of differences between means cantly during tests of nest, egg or wind type. tested by the T-method. Average initial temperature of eggs was 38.7±0.1°C, and slight variations during tests for nest, egg or wind type or egg Results positions were not significant. ANOVA showed that relative differ The mean mass (x±s.e.) of Black Brant ences among nest types were the same nests (38.3±2.7 g) was significantly less regardless of the species of eggs they con than that of either Emperor Goose tained (Table 3). Variation in cooling of 128 Steven C. Thompson and Denis G. Raveling 6 o o W INDY LÜ OC 3 I- < CC LJ Ol S ÜJ CD e> LÜ CO < LÜ CC CALM O £ LJ CU LT EMPEROR CACKLING BLACK GOOSE GOOSE BRANT NEST TYPE Figure 3.