(199) THE HATCHING AND FLEDGING OF SOME COOT

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1 (199) THE HATCHING AND FLEDGING OF SOME COOT BY RONALD ALLEY AND HUGH BOYD. SUCCESS INTRODUCTION. THE following data were obtained during the summer of 196, from observations carried out at Blagdon Reservoir, North Somerset, by kind permission of the Bristol Waterworks Company. During the most critical periods we were able between us to make almost day-to-day and day-long visits to the reservoir. We concentrated upon the Coot (Fulica a. atra) population of two adjacent bays. Choice of these bays was due to their convenience for observation. The conditions there prevailing, with regard to growth of vegetation, depth of water, disturbance by man (the lake is intensively fished during the summer), and in other ways, were not considered atypical of the reservoir as a whole. Of the fourteen pairs of Coot that attempted to nest within the study-area, seven were in their territories when we first watched the bays in March, one arrived late in that month, and the remainder came in a rush within ten days in May. Because of inclement weather, and especially a rise in the waterlevel, which delayed the emergence of growth suitable for nesting purposes and swamped two early layings, nesting was somewhat late. In the study-area the first egg was laid, by IV pair, on April 23rd. This was taken by crows and no further egg was laid until May 12th. Most pairs lost their first clutches in some manner, and no pair in the two bays (and very few elsewhere) reared more than one brood. VI pair alone laid after hatching young, and here the circumstances were of considerable interest: June 27th. Four eggs (marked by us) and two newly-hatched chicks. June 28th. One fresh egg laid alongside the other four. June 29th. Three young and five eggs one addition. June 30th. Three young and six eggs. July 1st. Nest found wrecked, eggs gone. We are satisfied that the fresh eggs were laid by VI female. Many cases of compressed cycles have been recorded in the literature (though not for the Coot), but we know of none where the overlap has been so marked. A certain amount of promiscuous laying took place within the study-area, and one pair (O) actually took over the incubation of I pair's second clutch of nine eggs, adding three of their own. As all eggs were, however, laid by the fourteen pairs, this remarkable, though perhaps characteristic, behaviour does not invalidate a quantitative treatment of hatching and fledging success as such. It is hoped to treat of nesting-behaviour in a subsequent paper.

2 200 BRITISH BIRDS. [VOL. XL. HATCHING. In all, 121 eggs were laid in the study-area, and of them 2, or 3.7%, were hatched. Egg-loss was therefore considerable. The following table summarizes our knowledge of the factors responsible : Cause of loss. TABLE I. CAUSES OF EGG-LOSS. No. of pairs affected. Swamping 2 Infertility... Abandonment 2 Knocking-out 3 Burying 3 Predators : (a) Man... 3 (b) Crows... 7 (c) Unknown 2 Undetermined Factors... Total No. of eggs lost % of eggs lost Infertile eggs. Three of the infertile eggs were found floating a few feet from the nests in which they had been laid and are thought to have been deliberately ejected: one had been sucked. The fourth was left for several days in the nest of L pair, after the young had been moved (see below). Subsequently it was found submerged near the nest. We have no evidence as to the agency responsible, for this. Burying. Loss by burying was due to the over-layering of eggs by the addition of material to the nest after they had been laid. The nest may be far from complete when laying begins, and materials may continue to be added so long as the nest is in use. Eggs may be wholly built-in within 8 hours. We have retrieved such eggs from abandoned nests four or more inches below the surface. Effective incubation ceases, of course, when the eggs are thus covered. The large number (10) of eggs lost in this way is due to the aberrant behaviour of O pair (see above), who buried no less than six. Crows. As may be seen, crows destroyed many eggs : their toll may well have exceeded 19. The open-cup nest of the Coot is very vulnerable to such depredation. Carrion Crows (Corvus c. corone) made frequent patrols along the shoreline, alert to pitch on any unguarded nest. VIII and IX pairs, nesting along a

3 VOL. XL.] HATCHING AND FLEDGING. 201 particularly exposed and crow-haunted stretch, lost nine of their ten eggs to crows. (The other was knocked out by one of VIII pair). Magpies (Pica p. pica) hunted the shoreline less often, but one was seen to be driven from III pair's nest by the owner, suggesting that they probably took some toll. Unknown predators. It is likely that some of the eggs placed in this category were removed by man. This class we use to cover the loss (not simultaneous) of two entire clutches, including that of VI pair, noted above. Knocking-out. Not infrequently, the Coot knocks out eggs from the shallow cup of the nest when leaving it in a hurry. Eggs falling into the water are lost, but those remaining on the side of the nest, or on the runway (where present) may be hooked back into the cup. Abandonment. Within a few hours of being hatched, chicks are led to a brooding platform some yards from the nest. Very often one adult will attend the young on their new platform while the other incubates such eggs as remain. As hatching may be spread over three or four days, the pull of the young proved sometimes too great, and the last eggs were abandoned. These were subsequently taken by crows. FLEDGING SUCCESS. In all, 28 young are thought to have fledged, 23.1% of the total of eggs laid, or 66.7% of eggs hatched; 2 of the young (the product of six pairs) remained with their parents in the latter's territory until seen to fly. The other four disappeared before they were seen to fly, but may fairly confidently be counted as fledged. In one case the parent birds and their three young abandoned the territory when the young were 7! weeks old, and are believed to have joined the flocks present in the area at this time. Ill pair's single surviving young disappeared when aged about 9 weeks, and a grey-bellied newcomer to the flock in the neighbouring bay is believed to have been it.* (Although we colour-ringed all the young produced in the two bays, we could seldom glimpse the combination on a bird whose identity puzzled us). Mortality of young was severe in the first few days of life, decreasing up to the end of the third week, and slight thereafter. Generally young would disappear without trace; only in the few cases recounted below was the cause of death ascertained. One young (10 days old) was killed in our sight by the adults into whose territory it had wandered. Attacks by territoryholding Coot upon straying young are very frequent until the young, by painful experience, learn (at about three weeks) to keep within the boundaries of their parents' territories. An adult will advance upon a trespassing chick, grab it around the back of the head, lift it clear of the water, and viciously shake and peck *This young returned to breed in the study-area in 197. Authors. N

4 202 BRITISH BIRDS. [VOL. XL. it. When released, the victim floats prostrate', so still the spectator thinks it dead, and then slowly raises its head and makes off; sometimes it is so dazed and lost that it goes in the wrong direction straight in to another attack. If the young Coot was not amazingly tenacious of life, very few could survive to be fledged. The number actually lost as the result of such assaults cannot be assessed; but the recovery of the body of another chick well within the territory of a neighbouring pair suggests that it may be considerable. The form of assault practised by adults on intruding young is the same as that used at times by parents on their own offspring, though in general more vigorous and persistent. Consideration of the significance of these parental assaults is outside the scope of this paper, but it may be remarked that they might well be a contributory cause, if no more, to the death of weak or sickly young. In the first week or so of life young Coot may be endangered by their lack of strength. An incident observed outside the study area illustrates the hazards to which they are exposed. One of us (R.A.) approached a chick aged about seven days, which dived in alarm. Surprised by its failure to reappear, he looked into the water, and " saw it motionless about six inches below the surface. For some seconds I watched to see what it intended, then perceived it was held across the nape by a thick strand of rush, and could not emerge" (extract from field notes). When released the chick was stiff; later it was found dead. One of the young in the study-area, three days old, was found floating alongside the nest in which it was hatched. It proved to be unusually light (we had previously noted its undersized appearance), and on examination a Cestode worm* was found constricting the intestine. It seems very likely that the parasite was responsible for the death of the bird, though the case presents difficulties, which it is not proposed to discuss here. No other instance of death resulting from disease or parasites was established. Danger to young from crows must be limited to the first few days of life, when the chicks pass most of the time on the nest or the " brooding platform." No evidence was obtained to show whether in fact crows did attack any young in the area. There are no hawks resident in the neighbourhood likely to prey on young Coot. Attacks on young by Great Crested Grebes (Podiceps c. cristatus) and Little Grebes (P. r. rttficollis) were seen, but did not compare in ferocity with the assaults of adult Coot; deaths resulting from attacks by grebes seem improbable. No evidence that ducks killed any young was obtained, though " hostile " manifestations by them were observed. Pike are rare in the reservoir, but eels and trout are numerous. The latter attain a large size. If fish took any young on the studyarea, it may be presumed that these two species were responsible. However, we failed to establish any record of such an occurrence. Dr. H. D. Crofton, of Bristol University, has kindly identified this worm as Dibothriocephalus sp. Authors.

5 VOL. XL] HATCHING AND FLEDGING. 203 SUMMARY. I. The nesting success of 1 pairs of Coot was closely studied during the summer of eggs were laid by these pairs, and of them 2, or 3.7% hatched. Factors responsible for egg-loss are tabulated and discussed. Nearly one-quarter of the egg-loss was caused by activities of the parents young are thought to have fledged, 23.1% of the total of eggs laid, or 66.7% of eggs hatched. Causes of mortality are detailed for a few cases : the evidence suggesting that adult Coot are a considerable factor here also.

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