BREEDING BIOLOGY AND BEHAVIOUR OF THE GREY PHALAROPE PHALAROPUS FULCARIUS IN EAST SIBERIA

1975 GREY PHALAROPE IN SIBERIA 285

BREEDING BIOLOGY AND BEHAVIOUR OF THE GREY PHALAROPE PHALAROPUS PULICARIUS IN EAST SIBERIA

A. A. KISTCHINSKI

Received 12 July 1973

The biology and behaviour of phalaropes are of particular interest because of the reversed role of the sexes in events linked with reproduction. Direct knowledge in this field seems to be especially poor for the Grey Phalarope Phalaropus fulicarius, in spite of the fact that this bird is common in arctic tundras. Observations in east Siberian tundras, scattered in faunistic papers (Birula 1907, Gladkov 1958, Sdobnikov 1959, Kapitonov 1962, Uspenski el al. 1962, Vorobyev 1963, Gladkov & Zaletayev 1965, Uspenski 1965, Portenko 1972, Haviland 1915), are fragmentary, and deal mainly with timing of the annual cycle and descriptions of nests. More detailed observations have been made in the North American arctic, Greenland and Svalbard (Manniche 1910, Bent 1927, Sutton 1932, Brandt 1943, Bailey 1948, Lsvenskiold 1954, 1964, Manning & Macpherson 1961). Recent accounts of the behaviour of Grey Phalaropes (Bengtson 1968, Hohn 1971a) have described many peculiar displays, including ritualized ones.

METHODS

Observations reported below were made at the localities shown in Figure 1. In the Chukotski Peninsula, 31 May-27 July 1970, we surveyed the northern coast between Cape Schmidt and Wankarem. In the delta of the Indigirka river, 3-28 June and 31 July- 4 August 1971, observations were made near the mouth of the Keremesit river flowing into the southernmost channel (Kolymskaya) of this delta: on 29 June-30 July we crossed the whole southeastern part of the delta. In the eastern part of the delta of the Yana river, data were gathered during 19 June-21 July 1972.

0 100 200 300 .c--l

km

e l 9 7 1

Study areas and years

FIGURE 1. Sketch map of east Siberia, showing localities mentioned in the text.

286 A. A. KISTCHINSKI IBIS 117

Dates of egg-laying were determined by finding incomplete clutches, by killing females in laying condition, or by deduction from the incubation stage of complete clutches, based on an incubation period of 19-24 days (Pedersen, cited in Salomonsen 1950; Conover 1926). Altogether, 93 nests were found and 126 birds were dissected. Specimens were also examined in the collection of the Zoological Museum, Moscow State University.

Simultaneous observations on Red-necked Phalarope Phalaropus lobatus provided comparisons, supplementing published reports (Tinbergen 1935, Hohn 1968).

HABITATS AND NUMBERS Both in the Indigirka delta, as noted by Mikhel (1935) and Uspenski et al. (1962), and

on the northern Chukotski coast, the Grey Phalarope and the Lapland Longspur Calcariw lapponicus are the two most numerous bird species. Here Grey Phalaropes inhabit low, flat, polygonal tundra. At the annual thaw, this tundra is flooded by shallow ponds, separated from one another by low dry ridges. From the end of June, these ponds progressively dry out, until water remains only in the centres of polygons and in the numerous permanent lakes. At the middle of June, Grey Phalaropes were evenly distributed throughout the tundra in the delta (especially near the shores of lakes) at a density of 1-2 pairs ha-l (cf. 0.3-0.5 pairs ha-l for the Red-necked Phalarope). South of the Kolymskaya Channel, where waterways are bordered by wide belts of willow thickets, polygonal tundra forms only about half the total area. The population density of Grey Phalaropes is correspondingly lower: on 12-13 June 1971, 19 pairs were counted in a sample area of 21 ha, (i.e., 0.9 pairs ha-1 cf. 0.5 pairs ha-l for the Red-necked Phalarope, which normally frequents permanent water bodies). In the maritime zone of the delta, 10-15 km wide, polygonal tundra is replaced by wet meadows dominated by Dupontia psilosantha and Eriophmum spp. These meadows are almost completely flooded in spring, and phalaropes nest there only sporadically.

In the eastern part of the Yana delta, Grey Phalaropes inhabit flat moss-sedge tundra with tussocks and numerous swamps and ponds, outside the willow-alder riparian belt. Counts during the incubation period (28 June-7 July) within a census strip 50-80m wide (51 ha) gave an average density of 0.57 nesting males per hectare, with a maximum of 1.5 males ha-l. Males were also found in the floodplains, among sedge marshes with low dwarf birch ridges and on old channels overgrown by Arctophila fulva, but in these habitats were less abundant. Females were observed up to the first days of July but subsequently disappeared. Further to the southwest, in the shrub tundra, Grey Phalaropes were absent while Red-necked remained common.

On the northern coast of the Chukotski Peninsula, Grey Phalaropes bred in the flat moss-sedge arctic tundra with innumerable lakes, ponds and (in June) temporary water- bodies. It was impossible to count birds, because they did not keep to any permanent home ranges (see below); but they appeared to be as common during the breeding time as in the Indigirka delta, and the density was probably similar. Red-necked Phalaropes also bred there, but at about one twentieth the density of Greys. On the island of the Amguema delta, tundra is of the more southern type, and banks of channels are bordered by willow thickets. Grey Phalaropes nested there in the central parts of the islands among wet polygonal tundra, outside willow growth, but were much less abundant than in the arctic tundra of the coastal flats. Numbers of both phalaropes were approximately equal.

A chain of shallow lagoons runs along the coast of the Chukotsk Sea. They are bordered by wet meadows dominated by Dupontia psilosantha, Carex subspathacea, Stellaria humifusa, Cochlearia arctica (the moss-sedge cover is absent) with innumerable brackish ponds. Phalaropes do not nest there, but many were found foraging.

Some time after completion of egg-laying, flocks consisting of females, non-breeding and unsuccessful males occurred near the breeding grounds, on the lakes among flat or


rolling coastal tundra. In 1970, in the Chukotsk tundra we did not see them during brief trips inland, 15 km and more from the sea. In 1971, in the Indigirka delta, such flocks were abundant until the middle of July on lakes in the maritime zone of the delta, but not more than 25 km from the sea. Later, these flocks moved on to the brackish coastal lagoons and to the sea.

The Grey Phalarope is known to be abundant in low coastal tundras in the other parts of eastern Siberia; near Anahar Bay (Uspenski 1965), in the delta of the Kolyma river and westwards (Buturlin & Dementiev 1934, Vorobyev 1963), around Tchaun Bay (Lebedev & Filin 1959), etc. In northern Alaska numbers fluctuate from year to year (Holmes 1966), and comparable fluctuations have been reported from the mouth of the Yenissei river (Haviland 1915) and western Taimir Peninsula (Vinokurov 1971).

BREEDING CHRONOLOGY Grey Phalaropes require sufficient areas of open water in the form of temporary ponds

of melted water or open strips along lakeshores. Accordingly, the times of their arrival depend on spring phenology. Thus, arrival of Grey Phalaropes has been reported in the Anadyr drainage during 26 M a y 4 June (Portenko 1939), near the Kolyma mouth at the end of May (Vorobyev 1963), in the Lena delta and Tiksi Bay from the end of May to 12-14 June in different years (Gladkov 1958, Kapitonov 1962). In the extreme north, in Greenland, Svalbard, and northern Taimir Peninsula, they do not arrive until middle of June or even later.

TABLE 1 Breeding chronology of the Grey Phalarope

Events

Northern The delta coast of the of the Chukotski Indigirka Peninsula river

(1970) (1971)

Arrival of the first birds First 'circle flights', courtship

displays, and attempts to copulate

Peak of arrival Mass pairing Peak of courtship behaviour,

First complete clutches Mass egg-laying Lessening of courtship displays Flocks of non-breeders,

mass copulations

unsuccessful males, and females having finished laying

Intensive moulting of adults (except incubatikg males) into winter plumage

Hatching Males leave nesting grounds First appearance of full-grown

flying- young Flocks of migrating birds on the

coastal lagoons and on the sea

1 June

1-3 June 6-10 June

13-18 June 15 June

17-25 June 19-30 June

-

21 June-10 July

Since 11 July 10-1 5 July'

?

25 July

8-27 July

4 June

4-7 June 8-10 June 11 June

11-14 June 13 June

15 June-1 July 15 June-2 July

4-16 July'

Since 10 July 7-22 July'

26 July-1 August

28 July

?

Notes: Birds observed only in the maritime zone of the delta where we arrived on 4 July. In 1960, intensive migration of flocks containing predominantly females took place on the seashore northwards of the delta on 20 June-6 July (Priklonski et al. 1963).

* Calculated on the base of laying dates (see text). Downy chicks were seen in the maritime parts of the Indigirka delta on 12 and 23 July.


In 1971 in the Indigirka delta the first Grey Phalaropes appeared on 4 June, and a peak was reached on 8-10 June (Tables 1 and 2); previous reports have noted arrival on 9-12 June 1930 (Mikhel 1935), and 5-13 June 1960 (Uspenski et d. 1962). By this date, large belts of open water had appeared along lakeshores, and many temporary ponds had emerged. Time and again, pairs and flocks flew in, low above the ground, and always from the east. Arriving birds dropped to the surface of a pond and immediately began to feed. The preceding non-stop flight was probably very long; many of the birds taken had empty stomachs. Westwards of the Indigirka delta, near the mouth of Bogdashkina river, it has been reported that Grey Phalaropes arrived in spring 1960 from the west; post- breeding flocks of females and non-breeders migrated both to the east and to the west (Priklonski et al. 1963).

TABLE 2

Sex ratio in Grey Phalaropes in the Indigirka delta (1971)

4-7 June 8-11 June 12-19 June (beginning (peak of (courtship

of the the time) arrival) arrival) Total

Total birds observed 90 273 177 540

Number sexed 71 273 177 521 Percent in pairs 56 92 99

Sex ratio (males per 100 females) 73 104 99 97

On arrival,courtship displays, chasing of males by females and attempts to copulate were seen at once. The peak of courtship activity, however, occurred in the middle of June and continued not more than a week, although residual displays were observed up to 2 July.

Dates of laying in 1970 and 1971 are presented in Figure 2. Females were not fully synchronized in reproductive condition. In 1970, for instance, of five females killed on 19 June three had shelled eggs in their oviducts but the two others had only ovarian follicles not exceeding 9-10 mm in diameter; the first full clutch was observed on 15 June. In 1971, in the southern part of the Indigirka delta, first incomplete clutches were found on 12 June, and in the central delta some females continued egg-laying till 8-9 July (Fig. 2). In 1972, incubated clutches were found on 30 June and 4 July; in the latter nest, the chicks hatched on 12 July.

Differences in the breeding chronology of Grey Phalaropes can be related to local conditions and spring weather (see Gladkov 1951, Kozlova 1961, Hohn 1965). Egg- laying seems to be earliest in the Yukon delta and near the Kolyma mouth, and latest on the Novosibirski Islands, in Greenland and in Svalbard (where the nesting schedule varies from year to year, L~venskiold 1964). On most arctic tundras, breeding schedules in general coincide with our observations. Data presented in Figure 2 imply that in 1971 egg-laying in the arctic tundras of the central part of the Indigirka delta occurred 7-10 days later (or was still more delayed) than in the tundra of more southern, subarctic types situated only 40 km to the southwest.

After laying, pairs break up (Table 1). Females gather in flocks, leave the nesting habitat and resort to the lakes and coastal lagoons; they are joined by non-breeding and unsuccessful males. Some flocks move to the sea before the end of June, and in the middle of July practically all Grey Phalaropes except incubating males are found on the seashore outside the breeding grounds. However, in some places (probably depending on food supply) non-breeders may remain until the end of July (Hohn 1971a; our observations).

1975 GREY PHALAROPE I N SIBERIA 289

At the end of July, males abandon the young, before they can fly, and leave the tundra altogether. The young complete their growth and also leave the nesting grounds in the first half of August. We found young in 1971 up to 6 August (cf. S. G. Priklonski, in litt., in 1960 up to 9 August).

c 11-15 16-20 21-25 26-30 1-5. 6-10

June July Dates of completion

FIGURE 2. Time of egg-laying in Grey Phalaropes: 1970-northern coast of the Chukotsk Peninsula; 1971-southern part of the Indigirka delta, mouth of the Keremessit river (W), and central parts of the Indigirka delta (U).

BEHAVIOUR AND DISPLAYS

In this report, we only deal with forms of behaviour linked with reproduction; special attention is paid to ritualized displays. Alertposture (Bengtson 1968) (= apprehensive posture of Hohn, 1971a). Any bird, either on the ground or on the water, can assume this posture when approached by another phalarope or by an unknown opponent. The neck is stretched vertically and the bill held horizontally. During the alert posture (and the flight which often follows it) the Grey Phalarope utters the common note muit-cruit or, when strongly excited, the quick tchirr- Zyk. This posture is also characteristic of both other phalaropes, the Red-necked and Wilson’s Phalarope Phaluropus tricolor (Hohn 1967).

AGGRESSIVE BEHAVIOUR

When swimming, the bird ruffles the feathers on its head, neck and back, lowers the head, pulling it between the shoulders, with the bill pointing forward, attacks another


bird on the water, and chases it. The same display is performed by Red-necked Phalaropes (Tinbergen 1935, pers. obs.). A similar display is often performed on the ground: as when swimming, the active bird ruffles its feathers, lowers its head, stretches its bill ahead and lunges towards the opponent, sometimes running several steps.

These displays occur when birds of the same sex approach one another closer than 20-30 cm; they are also frequent during the performance of courting swims (see below) by the pair towards an alien female. In the majority of cases, strong aggression is exhibited by female to female. Sometimes, however, a male displays towards another male, a female towards a male, or even a male towards a female which is attempting to chase or court him. Usually this aggressive display is restricted to one attack, after which the sub- mitting bird flies or swims away; sometimes, a short chase terminates in flight and pursuit in the air.

In the air, the active phalarope flies up, with legs lowered and with much flapping of the wings, and chases another bird, following it close to the ground and attacking it periodically. This is a common display between two females pursuing the same male.

Agressive displays are frequent during the courtship period - from arrival to egg-laying. Almost always the female is the aggressor, and aggression is mainly directed against females; reactions between males are much rarer and weaker. This aggression is evidently aimed to protect the proximity of a mate or to keep an individual distance, and not to defend any territory. As a rule, aggressive behaviour is only manifested in these ritualized displays; actual fights and overt aggression can almost never be seen. Similar postures are known in Wilson’s (Hohn 1967) and Red-necked Phalaropes. In addition, Grey Phalaropes may sometimes perform aggressive displays towards Red-necked and, according to Bengtson (1968), towards Purple Sandpiper Calidris maritima and Dunlin Calidris alpinu; Red-necked Phalaropes may act similarly towards Sanderlings Crocethiu a h and Turnstones Armaria interpres (Hohn 1971a). Driving flight. A peculiar type of aggressive behaviour is performed by a brooding male towards females which approach a recently completed clutch, for which I propose the term ‘Driving flight’. When approached by a female, the male flies up and ‘shivers’ 10- 50 cm above the ground, with tail sunk almost vertically and legs dangling in the air. With wings flapping very quickly (as when rattling, see below) and white underwing exposed, the male attacks the female, and beats her repeatedly with his wings until she leaves. This behaviour seems to serve as a mechanism for breaking up pairs and removing females from the nesting grounds. We saw many driving flights during 21 June-3 July 1971 and one on 7 July 1972, but this behaviour was never observed before the first clutches had been completed. Similar driving flights were seen, although more rarely, in Red-necked Phalaropes. These flights closely resemble behaviour of male Wilson’s Phalarope reported by Hohn (1967).

COURTSHIP BEHAVIOUR

Circle-flights (Bengtson 1968). The female flies round in circles of 100-200 m diameter, with slow wing-strokes, tilting from one side to the other to expose the white undersides of the wings, uttering the common cry w i t . Only females perform circle flights; searching for and attracting a male seem to be the functions of this display. Sometimes a female circles while chasing a male in aerial pursuit. Not unfrequentiy one male is chased by two or three females, and aggressive displays (see above) can occur between them. Circle flights seem to be a first phase of the precopulatory behaviour. They usually terminate when a male lands, and then follows the female (seldom the reverse); after this, various ground and/or on-water displays occur. Circle flights are most common in the first half of June (before egg-laying) although they can also be observed less frequently, up to the departure of females. The ‘ceremonial flights’ of Red-necked Phalarope (Tinbergen 1935)

1975 GREY PHALAROPE IN SIBERIA 29 1

are probably homologous with circle flights. Female Wilson’s Phalaropes often chase males in the air, but do not perform ritualized flights of a comparable type (Hohn 1967).

Kristoffersen (1926) stated that pairing Grey Phalaropes remained a long time on the wing, sometimes rising to a great height, and then dropping like a stone until they were close to the ground. Such behaviour was not seen by us. Courting Swim. Joint swimming and joint feeding of the pair are necessary elements of further precopulatory ritual. The courting swim posture (Bengtson 1968) is very common. In this, a female follows a male on the water, pressing her head down between the shoulders with bill pointing forward; this posture resembles to some extent the aggressive display on water. The male can either adopt the same posture, or alert posture, or be inactive. During this, the male and the female exchange cooing contact calls, very quiet but rather high pitched kryu Kryu kryu kryu. This call is uttered to solicit a flying partner and in various ritualized precopulatory displays either on the water or the ground. Courtship touchings. During joint swimming and joint feeding a male and a female swim very close to one another, either face to face or parallel and pressed flank to flank; similar postures were mentioned by Andrew (1955). After this, a bird touches with its bill the partner’s bill or feathers on the neck or breast. When doing this the female usually holds her head lower than the male. During Courtship touchings contact calls are constantly uttered, similar to those uttered during courting swims but quicker and speeding up as (presumably) the excitement grows.

Sometimes both mates draw their heads between their shoulders, push their bills forward (as in the courting-swim) and swim together, touching one another by the bills. In other cases, mates approach one another on land and perform similar courtship touchings. ‘Pushing’, described by Bengtson (1968), also seems to be another variation of courtship touching. A female lowers her head and presses the bill into her breast feathers. We rarely observed this posture. Hohn (1971a) believed it to be homologous with the ‘head down’ display of Wilson’s Phalarope described by him earlier (Hohn 1967). The ‘head-up’ posture of Wilson’s Phalarope (Hohn 1967) has been observed in Grey Phalaropes in the Canadian Arctic (Hohn 1971a), but we did not see it.

During joint swimming and courting, both on land and on water, the pair frequently perform aggressive reactions towards other Grey Phalaropes which approach too closely. Rattling. The bird suddenly flies up a few centimetres above ground or water and hovers for a moment in the air, with wings beating very quickly to produce a whirring sound. Neck and head are stretched forward, and the legs dangle. This display is performed by both sexes in the course of precopulatory ritual, but females do so much more often, sometimes in the absence of males. Rattling was described by Tinbergen (1935) and seen by us in Red-necked Phalaropes, but there is no similar display in Wilson’s Phalarope (Hohn 1967). Copulation. The male stands on the back of the female and hovers to maintain equili- brium while mating, a posture common to many waders. When copulating, the female always stands on the ground or in water 1-3 cm deep, as do Wilson’s Phalarope (Hohn 1967), whereas Red-necked always copulate in water, the female swimming and never standing on the bottom.

Precopulatory behaviour, on the whole, follows a definite sequence of ritualized displays. Circle flights are followed by aerial pursuits of males by females which terminate in both birds landing. After this, rather prolonged joint swimming and joint feeding of the newly formed pair are accompanied by various types of courtship touchings, rattlings and frequent contact calls, which serve to keep the partners together. Aggressive encounters with strange birds are common. All the precopulatory displays seem to contribute to the sexual stimulation of both sexes. Contact calls, courtship touchings and rattlings become more and more frequent; birds move onto the land or shallows, and the whole sequence U VOL. 117


ends in the attempt to copulate. The initiative in the precopulatory events is usually taken by the female.

Attempts to copulate are not always successful. Sometimes, the female moves or flies away, or the pair assumes the initial position but copulation remains uncompleted. After failure, the male jumps down and, after a new series of precopulatory displays, may repeat his attempt. After a copulation or after an unsuccessful attempt, both birds begin to feed or preen. The pair may remain together, or separate at once.

The sequence described above is not always maintained. It can be interrupted at any stage, when the pair may part or continue to feed together. Some steps of the display sequence can be missed, or the male remain inactive. In rare cases, the male may initiate courting and copulation, as on 17 June 1970: twice a male flew up to a female, performed rattling and alighted on the water; in response, the female began the courting swim. Portenko (1972) saw active courting by a male towards a female resulting in a successful copulation, and on several occasions we observed successful copulation, without preceding display, of a male with a female which arrived from elsewhere a moment before. Com- parable observations have been made by Andrew (1955) and for Wilson’s Phalarope by Hohn (1967) who, however, did not report whether the copulation was successful. According to Tinbergen (1935), in the Red-necked Phalarope, males initiate copulations rather frequently, but all successful acts observed were initiated by females.

At present we do not have sufficient knowledge to discuss the evolution of breeding behaviour in Phalaropinae. In general, aggressive displays are similar in all the three species. Courtship displays (circle flight, rattling, courtship touchings, courting swim) can be homologized in the northern phalaropes but those of the Wilson’s Phalarope are rather more different. Calls uttered during courtship displays differ between all species. In the most terrestrial, Wilson’s Phalarope, Hohn (1967) described head-up and head- down precopulatory postures which are performed on the ground. These may be related to some postures in Grey Phalarope but are not common in the latter species. On the other hand, in Wilson’s Phalarope we do not know any postures corresponding to those seen during courting on the water, so typical of the northern phalaropes. On the whole, breeding behaviour of phalaropes, especially of the northern species, seems to be more diversified than that of many other waders, due primarily to the development of various ritualized forms of courtship on water.

TERRITORIALITY As already stated by Hohn (1971a), with whom I agree, territories, i.e., individual or

family areas defended from conspecific birds (Howard 1920, Hinde 1956), do not exist in Grey Phalaropes. At courtship time, the attachment to some sort of home range may exist but, lacking data from marked birds, it appears that ranges are rather large and include a set of feeding ponds. Within the home range, hundreds of other birds (both breeders and non-breeders) can forage. In the Chukotski tundras we observed movements of females and of pairs (which demonstrated various courting displays) for hundreds of metres, and one bird might itself feed over an area of the same diameter. In the tundra adjacent to the Indigirka river, where in the middle of June 1971 numerous ponds were found, pairs kept to smaller areas of 50-100 m in diameter. Even here, however, other pairs fed on the same ponds, sometimes 2-3 pairs on one pond 15-20 m wide. We have no evidence indicating long attachment of a bird or a pair to an area. On the contrary, during 13-18 June 1970 we saw hundreds of Grey Phalaropes of both sexes, feeding on several groups of ponds; on 19-23 June they disappeared from this site but large numbers appeared on other lakes.

In regions where the species is rarer, stronger attachment to the home range has been reported (Manniche 1910, Bengtson 1968), but no forms of true territorial behaviour have been observed by us, Bengtson (1968) or Hbhn (1971a) in the Grey Phalarope. The


aggressive displays are not aimed to defend any particular area, but to maintain an individual distance of 20-50 cm or to keep a space of the same size around the mate (already available or to be expected). Sometimes these displays are performed in response to unsuccessful courting. Aggressive acts can happen on any body of water and not only on that habitually frequented by the bird.

No territorial behaviour was discovered in Wilson’s Phalarope (Hohn 1967). In Red- necked Phalaropes, although Tinbergen (1935) used the term ‘territory’, his detailed observations showed that he did not refer to a defended territory in the classical sense. Observations by Hohn (1968) and ourselves have confirmed that true territories are not maintained by Red-necked Phalaropes.

SEXUAL DIMORPHISM As in other phalaropes, female Grey Phalaropes are larger than males. In our samples,

69 males weighed 41-60 g (mean 50-8&0.53 g, s.d. 4.37), and 51 females, excluding cock-plumaged (see below), 49-73 g (mean 61.0*0.75 g, s.d. 5-36). No significant differences were discovered between the three areas studied.

Females have also bright nuptial plumage. The hormonal mechanisms involved have been studied by Johns (1964), Hohn (1967, 1970, 1971b) and Fevold & Pfeiffer (1968). The nuptial plumage and behavioural peculiarities of females can be related to an unusually high secretion of testosterone in their ovaries in spring, while the lack of brood patches and absence of brooding behaviour are linked with reduced secretion of prolactin, which is however produced copiously in males.

On 19 June 1970, we took from one flock two cock-feathered females in dull, male-like plumage; there were many white feathers of winter plumage on the abdomen, which had not moulted. These females weighed only 55 and 51 g. On 21 June, we killed a ‘half- cock-feathered’ female (weight 52 g) intermediate between the male and female breeding plumage. None of these females had ovarian follicles exceeding 3.5 mm in diameter. They kept in mixed flocks together with other non-breeders and some breeding birds. Pre- sumably disturbances in hormonal secretion (in particular, testosterone production) were responsible for their condition.

Females in full nuptial plumage are monochromatic. But among males two types are distinguishable: (a) those with grey cheeks and all the abdomen rusty-coloured, and (b) those with yellowish buff-grey cheeks and whitish feathers in the middle of the abdomen. Both variations were commonly observed, including paired and incubating males; intermediates were also seen. In the Yana delta in 1972, of 43 males, 24 (56%) were of type (a), 18 (42%) of type (b) and 1 (2%) intermediate. In the collection of the Zoo- logical Museum of the Moscow State University, of 39 males, 20 (51%) are of type (a), 16 (41%) of type (b), and 3 (8%) intermediate. No relation was discovered between the locality and percentage of colour types. It is also unknown whether these plumage types are age-dependent.

INTERRELATIONS BETWEEN SEXES IN THE BREEDING TIME Pair-formation can occur during migration, immediately after arrival or even later.

Reports postulating arrival in pairs mainly relate to the extreme north of the species’ range. In the Indigirka delta, where Grey Phalaropes appear rather late, most arriving birds in 1971 were paired; in the first few days females predominated, but later the sex ratio equalized (Table 2). On the northern Chukotsk coast, in 1970, most phalaropes arrived at the first days of June in flocks, not apparently paired. Near Tiksi Bay at the end of May and the beginning of June, Grey Phalaropes appeared mainly in pairs, rarely in flocks (Kapitonov 1962). Courtship displays, circle flights, chasing of males, various forms of courting on water, rattling, and copulatory attempts can be seen from the day of arrival.


T 18- s 8 16- 32 i 14 .- U

u- c 6 12-

- I O -

Males have large testes from arrival until the end of egg-laying (Figure 3). Gonads of arriving females, by contrast, are still poorly developed: of 12 females taken on 4-9 June, the diameter of the largest follicle in 9 was 3 mm, and only three had follicles in the range 5-9 mm.

0

-

o o + o

8

f I I I I I I I I I I 1 k k df arrival

f

U 4 iii :I 2

Mass laying - B 8 0

0 X

x* *::

0 6 .

0 0

I I I I 1 I I I I I I I ] 5 10 15 20 25 30 5 10 15 20 25 30

June July

FIGURE 3. Seasonal cycle in testicular length of Grey Phalaropes, 1970-72. (0) 1970; (0) 1971; ( X ) 1972.

In 1971, in the south part of the Indigirka delta, all Grey Phalaropes were paired by 12 June, i.e., the sex ratio was effectively parity (Table 2). All the males and females were in full nuptial plumage. Intensive courtship behaviour continued merely for a few days (Table 1): after 14 June, circle flights, aerial pursuits, and rattling could be seen only rarely. In the central part of the delta, where nesting was delayed (Fig. 2), residual courtship displays and copulations occurred until 2 July. Only three of 23 females and none of 34 males taken in 1971 during the laying season and later (17 June-23 July) proved to be non-breeders.

A different situation was discovered on the Chukotsk coast in 1970. The sex ratio was 86 (n = 164), which is not significantly different from parity (x2 = 0.88). Among both males and females, there were birds in full nuptial plumage as well as birds that had not finished their prenuptial moult; the latter had some feathers or areas of winter plumage and sometimes winter feathers even predominated. On 19 June we took a non-breeding female in full winter plumage changing into a new plumage, also of the winter type. All the other birds collected on 19-21 June, which had not completed their prenuptial moult, were not changing plumage, i.e., the moult had been suspended for the breeding period.

Only females in full nuptial plumage courted males; females collected with unfinished moult proved to be non-breeders. By contrast, males which participated in courtship and brooded clutches were in various stages of moulting. All the males collected with unfinished moult were taken from courting pairs. In the sample of 14 females taken during egg-laying period or later (19 June-1 1 July), seven were non-breeders (including two without any anomalies in moult or plumage); among males collected at the same time,


non-breeders made up 2 of 21. Thus, the sex ratio among the actively breeding part of the population showed a predominance of males, and a proportion of non-breeding females seemed to be essential.

In 1970, no permanent pairs were seen. Throughout June, Grey Phalaropes fed mostly in flocks of up to 50 birds, but sometimes in pairs. However, the composition of the groups, including pairs, was continuously changing, with members suddenly flying to other ponds up to 500m distant. This activity was spontaneous, and very common; it demonstrated free use of range. Occasionally a female began to court a male of the same or of another flock; the pair kept together and the sequence of precopulatory rituals might successfully occur, terminating in copulation. On other occasions, such a sequence was broken off, and the birds separated. In this case, the female flew up and/or sought another mate. Sometimes a female of the pair in any phase of courting began to chase another female, and the pursuit took it so far that (as far as we could see) it never returned, while the male continued to feed on the spot.

On 21 June 1970, a male and a female, both in full nuptial plumages, stood side by side on a tussock amidst a swamp and were preening, then began to feed together. Suddenly, another male in incomplete breeding plumage landed and started to feed beside the female. Shortly afterwards, he successfully copulated with her without any preceding display. After this, the new male and the female flew off together while the first male continued his foraging.

After copulation, some pairs remained together for a time, while others broke up at once. In the latter case, females joined any group, often flying hundreds of metres. Females in laying condition were twice taken from pairs, and once from a mixed flock dominated by non-breeders.

The observations reported above are inadequate for firm conclusions, but suggest the following possible pattern. In 1970 pairs formed only for the short time of egg-laying (3-4 days). Only during 16-23 June did we see pairs with any frequency. It seemed that the only biological function of pair-formation was to find a male for incubation. Copu- lation seemed to be promiscuous within the local population, and in some cases polyandry was probable. The clutch of a female was not necessarily incubated by the male which had copulated with her.

More permanent pairs might have occurred, but in such large promiscuous population we could not be certain in finding them. Thus, in one case (1971) all Grey Phalaropes in a population split up into distinct pairs that existed from arrival to the end of egg-laying (8-20 days), while in another case (1970) promiscuous mating seemed to occur, involving the whole local population. In spite of the lack of information for any region through a succession of years, it can be assumed that the social structure of a population in any area can vary from year to year.

In most faunistic papers, ‘pairs’ of Grey Phalaropes are mentioned; I believe this terminology is inferred by tradition, rather than based firmly on proved facts. A situation such as that observed by us in 1970 (i.e., birds staying in flocks throughout the whole summer; pairs ephemeral) may not be too rare but has been inadequately studied. When numbers are low, the Grey Phalaropes associate in pairs from arrival until the beginning of incubation, and the sex ratio approximates parity (Manniche 1910, Birula 1907, Llavenskiold 1964, Bengtson 1968). In June 1958, on Prince of Wales Island, there were 5-16 Grey Phalaropes per km2, mainly in pairs (Manning & Macpherson 1961). On one of the Hudson Bay Islands, six pairs nested in 1967; they spent most of the time in flocks, and attempts by members of established pairs to copulate with outsiders were noted (Hohn 1971a). Along the shores of Anabar Bay in 1959 (Uspenski 1965) and in the tundras adjacent to the Indigirka river in 1960 (Uspenski et al. 1962) Grey Phalaropes were numerous and kept in pairs from arrival, as they did in 1971. Around Tiksi Bay in 1957, phalaropes were abundant and occurred both in pairs and in flocks (Kapitonov


1962); this author reported, without details, that birds probably do not form permanent pairs, and females repeatedly copulate with different males within one season. Lastly, in south Iceland, according to S.-A. Bengtson (in litt.), Grey Phalaropes are rare and males predominate in the population. The following sex ratios are recorded: in 1969 146 (n = 32), in 1970 127 (n = 25), in 1971 125 (n = 18), in 1972 S.-A. Bengtson wrote: ‘It was difficult to recognize any definite pairs; the birds often courted and foraged in mixed groups’.

It is possible that promiscuous mating occurs when the sex ratio is unbalanced. The problem needs further study by means of marking, but one important conclusion can be already drawn: the social system of the Grey Phalarope is not invariable, but can alter in response to local and seasonal situations.

It is possible that social systems are equally variable in other phalarope species. In Wilson’s Phalarope, in Alberta in 1962-4, females predominated in the population; phalaropes were paired from arrival until the end of egg-laying (Hohn 1967). Dawson (1940) supposed that polyandry occurred in this species in a situation (similar to ours in 1970) when a considerable proportion of females did not mate, and males predominated in the breeding stock. Hohn (1967), however, has rejected this possibility; he considered the laying of two clutches by one phalarope female to be hardly possible. However, at least occasionally, a female seems to be able to lay additional eggs. On June 19 1970, we took two Grey Phalarope eggs out of three in an incomplete set; on June 21 there were four eggs in this nest. In the Red-necked Phalarope in Sweden, in a small colony of four females and six males, two females laid two clutches each during 13 days; each of these two mated with two different males (Raner 1972).

The latter observation definitely proves the possibility of ‘successive biandry’, when males prevail in a small population. Even if such a situation is uncommon, promiscuous mating by one female with more than one male, and vice versa, seem to be not uncommon. For the Grey Phalarope our observations of 1970 provided evidence, as did Hohn (1971a)’ and for the Red-necked Phalarope Tinbergen (1935). A male Red-necked Phalarope which mated with two females in succession was reported by Congreve & Freme (1930); females predominated in this population.

NEST LOCATION AND STRUCTURE As a rule, Grey Phalaropes nest among sedges in wet places in moss-sedge tundra,

usually on the edge of a temporary pond or on a tussock amidst flooded swamp. In such places we found 79 nests (25 in 1970, 52 in 1971, and 2 in 1972). More rarely, nests are built near the shore of a permanent lake or on islets in small lakes (3 found in 1970 and 4 in 1971), or on dry tussocks or ridges in the polygonal tundra (7 nests, all in 1971). In Svalbard and in the Canadian Arctic, nests have sometimes been found on open dry spots covered with lichens, or even among sand and gravel; such nests were merely shallow holes almost without lining (Eifrig 1905, Lavenskiold 1964).

The nest is lined with materials gathered nearby: in the polygonal swamps by dry sedges Carex stuns, etc., and cotton-grass; on dry ridges by small dead willow leaves and lichens ThamnoZka, as well as sedges. In wetter sites the lining is thicker; nonetheless, at the beginning of incubation the floor of the nest under eggs is often damp. As incubation proceeds, temporary ponds gradually dry up, and by the time of hatching many nests are on dry ground, sometimes 100-200 m from the nearest water, forcing the male to move rather far to feed. Nests are nearly always well hidden by overhanging blades of sedges and grasses of the previous year, which form a sort of canopy over the incubating bird.

Groups of 5-10 nests have been found by Salomonsen (1950), Kapitonov (1962) and Hohn (1971a). In Svalbard, Grey Phalaropes sometimes nest in clusters within colonies of Arctic Terns Sternu paradisma (Larson 1960, Lsvenskiold 1954), and nests have been


found seven paces from each other (Riippell 1968, cited by Hohn 1971a). During our work, most nests were distributed rather evenly, 40-80 m apart or farther, but in 1970, I discovered two with brooding males 6 m apart, and in 1971 Dr V. E. Flint found two clutches 3 m apart. I n June 1971, five pairs of Grey Phalaropes kept to the periphery of a colony of Ross’ Gulls (one nest found), and on 22 June a clutch was observed close to an Arctic Tern’s nest.

Nest location and structure were similar in all the areas studied in 1970-72; the same pattern has been reported in other parts of east Siberia. The nests of the Red-necked and Wilson’s Phalaropes look like those of Grey Phalaropes; Red-necked breed more often near lakeshores, Wilson’s on wet meadows. Clumped nesting has been discovered among all phalaropes (Hohn 1967, Raner 1972; our observations).

Building of the nest by the male only was observed by Manniche (1910) and Brandt (1943); Hohn (1971a) reported building by both the sexes. On 24 June 1971 I saw a male making a nest cup, and on 16 June 1971 V. E. Flint saw a female doing the same. As do other phalarope species (Tinbergen 1935, Hohn 1967 ; our observations), Grey Phalaropes typically make several nest hollows; only one is ultimately used for laying, while the others are not lined and remain empty. We saw such empty cups many times, especially in 1971, as has been reported by Larson (1960) and Hohn (1971a).

INCUBATION AND PARENTAL CARE

Incubation begins after the laying of the second or third egg. Males were flushed from two of five sets of 2 eggs, and from all sets of 3 eggs. On one set of 3 eggs a female was sitting, but females probably replace males only in order to lay. In the vicinity of nests with incomplete clutches, both birds were usually present, but near full clutches we saw females only twice (17 and 23 June 1971).

After laying, males begin to drive females away from the nests, and pairs break up. Fluttering flights of males (see above) chasing females are common at the end of June. After the departure of females, only incubating males remain in the breeding habitat.

Only males have brood patches. Some authors have reported that they flushed female Grey Phalaropes from nests, that they found females attending broods, or showing concern when an observer approached the young (Haviland 1915, Sutton 1932, Brandt 1943, Larson 1960, Dhondt et al. 1969). Examination of each of the cases reported shows that when females were seen on the nests, the possibility could not be ruled out that it was a final stage of egg-laying (except, probably, in the case reported by Brandt, 1943). Observations dealing with attendance and care of young are evidently reliable. Never- theless, such a behaviour in females is a rare exception and may well be due to endocrine disturbances.

In the Red-necked Phalarope, females have only twice been flushed from nests (Glad- stone 1907) - once from an incomplete set, another time from one probably just finished. No reliable evidence of females incubating eggs or accompanying broods has been offered for Wikon’s Phalarope (Hohn 1967), although females can stay near the nests, sometimes in groups, and can appear alarmed when one approaches the nest (Nelson 1877).

Incubating males feed either on their own or in groups of 4-5 birds on small lakes and ponds. At the end of June and in July, most temporary ponds dry up, and feeding males gather on remaining bodies of water, sometimes far from their nests. Not infrequently, they join mixed phalarope flocks which consist of non-breeders, unsuccessful males and females that have finished laying. No aggressive reactions between males exist at this time; they are indifferent to each other, even if their nests are located 3-6m apart. Groups of feeding males form and disband freely. On the ponds around any nest other Grey Phalaropes can freely feed, and the incubating male show no hostility to them. The tendency of males to drive away females is probably strong only in the first days of incubation. However, it has been reported (Levenskiold 1964, Bengtson 1968) that if a


strange female comes close to the young, the attending male drives her off without delay.

Grey Phalarope males are close sitters. They usually fly only if approached closer than 1-3 m. However, some males left their nests when the observer was still 10 m away and walked several metres through the high grass before flying. After the disappearance of danger, the male, having swam and fed nearby or having sat 5-20m from the nest, reapproached the nest on foot, quietly making his way through the grass. From fresh eggs males flew rather far, and silently; when flushed from advanced clutches, they sometimes fluttered around, from time to time sitting down 5-10 m away. Distraction display was seen on only one occasion. On July 4 1972, we approached to 1.5 m of a clutch, 8 days before hatching; the male flew up, sat down and then ran several metres through the grass with feathers ruffled and wings half-spread; later he flew. Other descriptions of ‘active’ distractions can only be found in the reports by Manniche (1910) and Sutton (1932).

In the middle of June, the young hatch and the behaviour of males changes. By this time, temporary ponds have dried up, and the birds concentrate near permanent lakes. When one approaches the brood, the male flies out from a distance of 20-50m and, uttering the alarm call tchirr-Zik, circles in the air or flutters over the intruder, time and again alighting at a distance of 10-12 m. He is usually joined by other males feeding nearby. We did not see distraction displays similar to those described for Wilson’s Phalarope (Hohn 1967). With downy chicks (12 and 23 July 1971, 12 July 1972), a male kept very close. Later, according to several authors, males often feed rather far away from their broods, but keep permanent contact with the young and, if necessary, guard them, attacking and driving off Arctic Terns, skuas and Grey Phalarope females if they appear in the vicinity (Bertram & Lack 1933, L~venskiold 1964, Bengtson 1968). By the end of July males aggregate in flocks, leave their young, and abandon nesting grounds. The young stay behind on various ponds, including very small ones, until the beginning of August.

BREEDING SUCCESS

Complete clutches may contain 2-4 eggs, but mean clutch-size is usually close to 4; only in 1971, in the central parts of the Indigirka delta where nesting was delayed, was mean clutch-size lowered (Table 3). Females can probably lay more than 4 eggs (see above) but due to their early departure from the nesting grounds, repeat clutches seem to be rare. Repeat clutches after the loss of the first set were reported by Nelson (1877) in Wilson’s Phalarope and by Gladstone (1907) in Red-necked. Sometimes six or seven eggs have been found in the nests of Grey Phalaropes (Koenig 1911, Bent 1927, Bengtson, in litt.), and even eight eggs in two nests of Red-necked Phalarope (Congreve & Freme 1930), but it was believed that these instances each involved two laying females.

Nests of Grey Phalaropes are easy to find; so, one can suppose that their loss may be high. Indeed, in the first half of July, both in 1970 and 1971, we saw and took many males that had lost their clutches. Of the total number of males taken after 17 June (i.e., at the laying time and later), unsuccessful males in 1970 amounted to 11 of 21, and in 1971, 10 of 34. These data are not fully comparable, but on the whole, they show that the percentage of unsuccessful males was high. In the middle of July, they were moulting intensively, and their brood patches were rapidly being covered by feathers. These features allowed us to distinguish these males both from non-breeders (which had no brood patches at all) and from incubating males which had well-developed brood patches and did not moult until the end of July.

Nest losses seemed to be especially high in 1971, when we found many destroyed nests in the Indigirka delta, and witnessed a heavy migration of unsuccessful males in the middle of July. On 20-21 July 1971 we counted, in the typical habitats of the central delta, 74 males evidently attending broods per 150 ha, i.e. 0.5 male ha-1. Thus, the density


TABLE 3

Clutch size

Number of nests with clutch

Locality and year 2 3 4 Meanis.e. s.d.

Northern coast of the Chukotski Peninsula, 1970 - 1 25 3.96-10.039 0.20

the Keremessit river (southern parts of the Indigirka delta), 1971 - 1 21 3.9510.045 0.21

The central parts of the Indigirka delta, 1971 4 5 24 3.6110.123 0.70

The delta of the Yana river, 1972 - -

The lower reaches of

2 4.00

Note: Clutch-sizes were determined on the basis both of clutches found (when we were sure that they were completed) and examination of several collected females having just finished egg-laying.

of males was two to four times lower than in June. The number of full-grown young counted during 1-3 August 1971, on an area of 110 ha, was 0.2 ha-l; this indicates a population gain of only 10-20%. This poor breeding success, in spite of a low percentage of non-breeders (see above) can probably be attributed to predation by Arctic Foxes Alopex lagopus, gulls and skuas, under the conditions of an unusually dry summer, when nests were easily accessible and when numbers of lemmings were low. I n the Red- necked Phalarope too, breeding success was low in 1971. Large losses have been reported in Wilson’s Phalaropes as well (Hohn 1967).

DEPARTURES OF FEMALES AND NON-BREEDERS

After laying, females join non-breeders of both sexes to form mixed flocks which move to the maritime parts of the tundra, These flocks can consist of several birds to several hundreds. Both females in egg-laying condition and incubating males temporarily join such flocks. For example, on 19 June-23 July of 1970 and 1971, there were taken from mixed flocks: 8 non-breeding females, 2 non-breeding males, 4 females in laying condition, 10 females having finished laying, 6 brooding males and 21 unsuccessful males. Grey Phalaropes have a strong tendency to aggregate in flocks. For instance, Hohn (1971a) has observed that all the females of his study area left together, among them one which had not laid at all and another which had only laid two eggs. In our experience, in 1970, phalaropes stayed in flocks throughout the summer, and the transformation of ‘courting flocks to post-breeding’ ones was hardly noticeable; we could register it only by changes in the condition of gonads in the birds collected.

The composition of post-breeding flocks is not constant. They continuously break up and form again, even in a span of hours. As a rule, Red-necked and Grey Phalaropes form mixed flocks. In July, more and more unsuccessful males join these flocks. At the beginning of July, birds in these flocks begin to moult into winter plumage; males and females which had interrupted moulting for the courtship season (1970) now resumed their moult. In the middle of July moulting was very intensive, and by the time of departure to the sea, some birds are halfway to winter plumage.


Post-breeding flocks perform regular circling flights over the lakes, all members turning together so concordantly that the whole flock changes colour, from the simultaneous exposure of white undersides of the wings. In 1970, the flocks in the Chukotsk tundra gradually moved in July to the brackish lakes, the coastal lagoons and the sea. In 1971 we observed in the Indigirka delta an eastward migration of mixed flocks which was intensive from 14-17 July and then ceased. In the most northern parts of the breeding range females sometimes stay longer, until the end of July (Bengtson 1968). In the east Siberian coastal waters Grey Phalaropes can remain till September or even early October (Portenko 1972: 412).

ACKNOWLEDGMENTS I would like to thank Dr V. E. Flint who took an active part in the study in 1971, and who kindly

put his important personal observations at my disposal; also Dr S. G. Priklonski for his unpublished notes made in northeast Yakutia. Dr S.-A. Bengtson kindly sent me his unpublished observations on the Grey Phalarope in Iceland, and read the whole manuscript and made valuable comments; a first submission was critically reviewed by Prof. N. Tinbergen. Collections of the Zoological Museum of the Moscow State University were studied by courtesy of the Museum staff.

SUMMARY Studies were made in 1970 in the Chukotski Peninsula, in 1971 in the delta of the Indigirka river

and in 1972 in the delta of the Yana river. Grey Phalaropes inhabit polygonal and tussocky moss-sedge tundra rich in swamps, lakes and (in June) temporary ponds. Population density in favourable habitats may reach 1-2 pairs ha-’. Data on breeding chronology are presented, and various aggressive and courtship displays described. Most phalaropes seem to keep within a home range, sometimes large, during courtship time, but no defended territories and no forms of territorial behaviour exist. Many birds, both local nesters and wanderers, can feed on any pond. Sexual dimorphism is described. In 1970, three non-breeding cock-plumaged females were taken. Pairs are formed both before arrival and on the nesting grounds. AU courtship displays are wholly or mostly initiated by females. In 1971, in the Indigirka delta, all the Grey Phalaropes were paired by 12 June, and stayed in pairs until the end of egg-laying. In 1970, on the northern Chukotsk, phalaropes seemed to form no (or very few) permanent pairs. Throughout June, most birds occurred in mixed flocks constantly moving between lakes or ponds. Copulation seemed to be promiscuous within the local population; polyandry cannot be excluded in some cases. Pairs appeared to be created only for the time of egg- laying; probably, the only biological role of pair-formation is to find a male for incubating. Thus, a definite social system is not a species-specific feature; it can vary depending on local and yearly situations, including probably sex ratio. Nests are usually situated in very wet places, sometimes on the water edge. They can be found as little as 3 m apart, but are usually 40-80 m apart, or further. Incubation begins after the second or third egg. After the end of egg-laying, males drive females away from the nests, and pairs break up. Females and non-breeders gather in flocks and move onto the lakes of maritime tundra, and later on to the sea. The composition of the flocks is not constant: they often join together or part. Brooding males feed near their nests, sometimes in groups; not unfrequently they join flocks of females and non-breeders for some time. The normal average clutch- size is c. 4 eggs; when nesting was delayed (in the central Indigirka delta in 1971) the average was 3.61. The loss of nests was great in 1971 ; numbers of young on 1-3 August was 10 times lower than adult numbers in June.

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Documents

BREEDING BIOLOGY AND BEHAVIOUR OF THE GREY PHALAROPE PHALAROPUS FULCARIUS IN EAST SIBERIA