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The Composition of Petrel Eggs

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The Composition of Petrel Eggs Condor 85:194-199 0 The Cooper Ornithological Soaety 1983 THE COMPOSITION OF PETREL EGGS JOHN WARHAM ABSTRACT. -Examination of fresh eggsfrom 23 speciesof procellariiform birds showsthat the relative amounts of albumen and yolk in the eggdepend upon egg size, large eggshaving proportionately more albumen and less yolk than smaller ones. Similar allometric relationships also exist within two species, Diomedea immutabilis and Pufinus griseus.The proportions of these components in petrel eggsdiffer from those in the eggsof either precocial or altricial birds in general. The yolk in petrel eggslighter than 25 g weighs more than that in eggsof similar size produced by typical precocial birds. Shell weight is a relatively constant percentage of egg weight, except for fulmars, which nest among rocks and on ledges:their eggshave heavier and thicker shellsthan other petrel eggsof similar size. Heinroth (1922) pointed out that the eggsof these advantages, little information is avail- altricial birds contain less yolk than those of able about the constituents of the eggsof al- precocial birds. The yolk reserves carried by batrosses,shearwaters, and other petrels. the hatchlings in their yolk sacsare also gen- In this paper, I report the yolk, albumen, erally larger in precocial birds than in altricial and shell weights for eggs of 23 members of ones of similar body weights. Ar and Yom- this order, including the southern fulmarine Tov (1978) calculated that, on average, pre- speciesMacronectes giganteus, Fulmarus gla- cocial hatchlings have twice as much yolk as cialoides, Daption capense, and Pagodroma altricial ones. nivea previously studied by Etchecopar and Romanoff and Romanoff (1949) compiled Prevost (1954). data on the weights of egg components with The study was undertaken (1) to test the respectto the weightsof intact eggs.These may hypothesis that the proportions of the con- be summarized: stituents of the eggsvary in a regular manner Altricial species-yolk 12-27%; albumen 68- among procellariiform speciesaccording to the 79%; shell 5-10%. size of their eggs, (2) to seek anomalies re- Precocial species-yolk 32-4 1O/o; albumen quiring further examination, and (3) to deter- 51-56%; shell 9-14%. They pointed out that mine if the intraspecific trends noted by the the proportions of yolk, albumen, and shell Romanoffs in the fowl also exist among petrels. vary with the size of egg in both altricial and (In this paper, “petrel” is used as a general precocial species,larger eggshaving relatively term for any procellariiform bird.) smaller yolks, larger whites, and heavier shells than small eggs.They also noted, as had Curtis METHODS (19 12) that similar size-related differences in Freshly laid eggswere hard-boiled, cooled, and the egg’s constituentsexist intraspecifically in after separation, the shell with its underlying the domestic fowl (Callus gallus, var. domes- membranes, the albumen, and the yolk were ticus). weighed. Some whole eggswere weighed under The study of the egg’s contents has been field conditions before and after boiling, using stimulated recently becauseof its importance spring balances; others were weighed indoors to the energetics of parental care and devel- on Mettler balances,as were all the individual opmental processes(see, for example, Lack constituents. Boiling reduced egg weight by 1968a, b, Ricklefs 1977, Ar and Yom-Tov about l-3%, probably because of water loss 1978, Carey et al. 1980). In uncovering basic from the albumen. I therefore increased al- relationshipsbetween the egg,its producer, and bumen weight by the difference between the its product, the order Procellariiformes has fresh and the boiled egg weight, as recom- special characteristicsthat may make it par- mended by Curtis (19 12). ticularly useful. First, the group is monophy- Because all the birds were protected, only letic; second,its members exhibit a great range small numbers of eggswere usually available. of body weight from about 20 g in the Least They came from breeding placesin the Pacific Storm-Petrel (Halocyptena microsoma) to and Indian Oceans,from Antarctica, and from about 8,700 g in the Royal Albatross (Di- Scotland. Larger samples,e.g., from the Sooty omedea epomophora);and third, the clutch of Shearwater (Pujkus griseus)and the Laysan all the speciesconsists of only one egg.Despite Albatross (Diomedea immutabilis), consisted [I941 PETREL EGGS 19.5 whereas yolk or shell weight and egg weight were not (P < 0.0 1 to 0.1). Logarithmic plots of the data for the species listed in Table 1 (Fig. 3) indicate that the egg’s constituents were highly correlated with egg weight. The correlations between the log,, of the component weightsand the log,, of the egg weightswere very high (r values between 0.983 and 0.999), but again, they were highest for albumen weight, as they were for Sooty Shear- waters and Laysan Albatrosses(Fig. 2). Comparing incubation periods with albu- men and yolk weights disclosed similar rela- tionships. I have satisfactory figures on the :,..;‘ :‘ “true” incubation periods (i.e., the lengths of time elapsingbetween the laying and hatching of eggsthat receive uninterrupted incubation) of 16 of the speciesin Table 1. (This definition Y 20. of the incubation period is important because petrel eggsare resistant to chilling [Matthews 19541 and temporary desertion is quite com- mon in some species,giving the impression of long “apparent” incubation periods.) For these 16 species,the correlation coefficient Ybetween incubation period and albumen weight was 0.957 (P < O.OOl), and that between incuba- 0 100 200 300 400 tion period and yolk weight was 0.942 (P < Egg Wtcg) 0.00 1). FIGURE 1. The relative proportions of the components making up the eggsof 23 procellariiform species.Circles DISCUSSION are values for shell (s), closed squaresvalues for yolk (y), Considering the small number of eggsthat I and open squaresvalues for albumen (a). % = (weight of part/weight of egg) X 100. examined, the data show surprisingly few anomalies, the differences in the proportions of the parts being regular and consistentamong eggs of different size. For example, fulmars of surface and/or abandoned eggs.(Burrowing (Macronectes, Fulmarus, Daption, and Pago- petrels often lay eggs on the surface of the droma), which have the fastest development ground: theseare not incubated, and some eggs ratesamong petrelsboth as embryos and chicks are abandoned even by birds of speciesthat (Warham 1968) do not have eggs with ab- nest on the ground itself. Such eggsare prob- normally small yolks. Thus, their relatively ably laid by the younger members ofthe colony short incubation periods are not a consequence and tend to be smaller than those of estab- of eggs with small yolks, which might favor lished breeders (Serventy 1967, Warham, un- earlier hatching. (The quality or energy content publ.). Hence, they may differ slightly in com- of fulmar yolks may nevertheless differ from position from the eggsof older birds. those of other petrels.) The ratios of yolk to albumen in the eggsof RESULTS procellariiforms range from 0.36 to 0.70, the The trends noted by Romanoff and Romanoff percentagesof yolk from 24.3 to 39.0, those (1949) for birds in general also apply to petrels of albumen from 54.5 to 68.5 (Table 1). These (Fig. 1). Larger eggscontained more albumen, values lie between the ranges listed by Ro- yolk, and shell than smaller ones, but the pro- manoff and Romanoff (1949) for altricial and portion of albumen was higher, whereas the precocial species.Nice (1962:26), commenting proportion of yolk was smaller. Shell weight on gtchecopar and Prevost’s data for penguins was a relatively constantpercentage of the egg’s and petrels, wrote, “We do not know whether weight, irrespective of the latter’s size. the large amount of yolk is an adaptation to Similar relationships appear to exist within extreme cold or is a characteristicof thesebirds speciesof petrels too, as indicated by my data throughout their range. It may indicate that for the Laysan Albatrossand Sooty Shearwater penguins and albatrossesand perhaps petrels (Fig. 2). In both species,albumen weight and are really more semi-precocial than semi-al- eggweight were highly correlated (P < 0.00 l), tricial.” 196 JOHN WARHAM 80 60 I I 0 I 80 100 120 Egg Wt cg) FIGURE 2. Components in the eggsof 15 Sooty Shearwaters(A) and 16 LaysanAlbatrosses (B). Symbols as in Figure 1. Equations for the regressionlines in A: Albumen Weight = 0.79 Egg Weight - 20.39 (I = 0.964; P < 0.001) Yolk Weight = 0.15 Egg Weight + 17.83 (Y = 0.553; P < 0.05) Shell Weight = 0.06 Egg Weight + 2.56 (r = 0.628; P < 0.01) and in B: Albumen Weight = 0.78 Egg Weight - 45.49 (r = 0.918; P < 0.001) Yolk Weight = 0.18 Egg Weight + 35.55 (r = 0.462; P < 0.1) Shell Weight = 0.05 Egg Weight + 9.49 (r = 0.485; P < 0.1). My data, however, suggestthat the com- Fairy Prion (Pachyptila turtur) precocial be- ponents of petrel eggsdo not vary according cause they attain homeothermy within a day to climate or latitude, and that their rather high or so after hatching. Clearly, it is hard to define proportions of yolk are characteristic of these these terms precisely becausethe statesof ma- birds throughout their breeding range-be it turity of newly hatched chicks form a contin- tropical, temperate, or polar. uum between the extreme precocity of a young Nice (1962) recognized intermediates be- megapode (which can fly within a few hours tween the “true” altricial and “true” precocial posthatching, is homeothermic, and receives groups. Consequently, she developed a de- no parental care) and the extreme altricial state tailed systemof classifyingneonatal birds based of a neonatal passerine (which is wholly de- on eggcomposition and other criteria, such as pendent on its parents for food and most of the chick’s ability to move about within its its warmth, and does not become homeo- first day or two after hatching.
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