Sandpipers
(Scolopacidae)
Class: Aves
Order: Charadriiformes
Suborder: Charadrii
Family: Scolopacidae
Thumbnail description
Small to medium-sized waders with short to long legs; bill short to long, may be straight, decurved, or slightly recurved.
Size
4.7–26 in (12–66 cm);.03–3.0 lb (14.5–1,360g)
Number of genera, species
23 genera; 86 species
Habitat
Coastal and inland wetlands; breed in tundra, marshland, grassland, steppe, scrub, and forest
Conservation status
Extinct: 2 species; Critical: 2 species; Endangered: 2 species; Vulnerable: 6 species; Near Threatened: 10 species
Distribution
Worldwide, except Antarctic
Evolution and systematics
The fossil record suggests that the family Scolopacidae arose about 40 million years ago in the early Tertiary, during explosive evolution after a wave of extinctions at the end of the Cretaceous. Some fossil material, possibly representing the genera Limosa (godwits) and Tringa (shanks), is available from the late Eocene, the period when this radiation probably occurred. Although there is great diversity within the Scolopacidae, morphological and biochemical evidence suggest all groups currently included in the family have a common ancestor. Recent osteological studies suggest this ancestor may also have given rise to the jacanas (Jacanidae) and the painted snipes (Rostratulidae), while DNA-DNA hybridization studies suggest these two groups, with the seedsnipes (Thinocoridae) are the most closely related outgroups of a monophyletic scolopacid family. The relationship of the Scolopacidae with plovers (Charadriidae) is disputed, and most recent authors suggest that the two are not sister groups as formerly assumed.
Although phalaropes (Phalaropus and Steganopus) have often been separated into their own family, molecular genetic techniques confirm they belong within Scolopacidae, as do other morphologically and behaviorally diverse taxa such as woodcocks (Scolopax) and turnstones (Arenaria). All three groups are given subfamily status (Phalaropodinae, Scolopacinae, and Arenariinae respectively) within Scolopacidae, which also has three other subfamilies: the Gallinagininae (snipes), Tringinae (shanks and allies), and Calidrinae (sandpipers). The Tringinae are usually subdivided into three tribes, the Numeniini (godwits and curlews), Tringini (shanks), and Prosoboniini (Polynesian sandpipers). Four subfamilies, the Scolopacinae, Arenariinae, Calidrinae, and Phalaropodinae, are probably monophyletic.
Within the family, 86 species (64 monotypic) are currently recognized in 23 genera (12 monotypic). Most species breed in Arctic or boreal environments, but when the family first appeared the climate was warmer. About 1.5 million years ago, at the end of the Pliocene, the earth's climate became similar to that of today and since then, during the Pleistocene, alternating ice ages and interglacial periods have had great impacts on the geographical distribution of scolopacid waders and extent of their habitats. Such climatic changes may have played a major role in evolution of species and subspecies. Recent DNA studies suggest that extant subspecies of the dunlin (Calidris alpina) originate from climatic events that caused repeated fragmentation of populations towards the late Pleistocene, from about 200,000 years ago. On the other hand, the five subspecies of the red knot (Calidris canutus) appear to have arisen only about 10,000 years ago and may all be derived from a small population that survived during retreat of northern ice-caps.
Physical characteristics
Scolopacid waders vary in length from about 5–26 in (12–66 cm) and in many species the bill makes up a large proportion of total length. Great variation in bill morphology reflects great diversity of food resources exploited and a wide range of habitats used. The bill is often long and decurved, this feature being especially prominent in the curlews (Numenius), while godwits and the Terek sandpiper (Xenus cinereus) have an upcurved bill. Turnstones have a short, relatively strong, wedge-shaped bill, while the spoon-billed sandpiper (Eurynorhynchus pygmaeus) has a spatulate bill tip. In most species the bill tip contains many tactile receptors for locating buried prey, while the tip of the upper mandible is flexible, allowing it to be opened and closed while probing, to grasp prey. Compared with plovers, most species have relatively narrow skull and small eyes, correlated with use of more tactile foraging methods. In woodcock and snipe, the eyes are placed high on the side of the head, to give 360° vision that aids predator detection.
Body shape varies from slender to stocky; the tail is short and the neck and legs are often fairly long. Feet usually have three long front toes and a short hind toe, and in some species are partially webbed. Most species can run rapidly and, although they can swim, many species habitually wade in shallow water. In most species, wings are long and pointed, well-adapted to long-distance migration.
Sexes are usually indistinguishable in plumage features. Females are often slightly larger and longer-billed, especially in the Tringinae, but males are larger in some Calidrinae. Nonbreeding plumages are typically dull and cryptic, upper-parts being brown, gray, or blackish with pale streaks or spots, and the underparts pale with variable dark markings. Breeding plumage is often much brighter, with extensive rufous or black areas. Breeding-plumage male godwits are more brightly colored than females, but the ruff (Philomachus pugnax) takes sexual dimorphism to the extreme in the family: the male is almost 25% larger than the female and develops a ruff and head tufts of erectile feathers during the breeding season. In contrast, female phalaropes, which court males and defend territories, are more brightly colored than males. Many species have white on the tail, rump, or lower back, and often a white bar on the upperwings. The bill, legs, and feet are sometimes brightly colored and in some species may change color with age or season, becoming brightest prior to breeding.
Three eco-morphological groups are distinguishable: the snipes, sandpipers, and phalaropes. The snipes and woodcocks have very long bill and very cryptic plumage. Phalaropes are small, plumpish but elegant birds with straight, narrow bill and short legs. The remaining genera are lumped into the sandpiper group: wading birds with short to long legs and short to very long, narrow bill.
Downy hatchlings of scolopacid waders are cryptically colored and patterned, and have well-developed legs. Most have a small bill that gives little indication of length and shape of the adult bill.
Distribution
The family occurs virtually worldwide, with the exception of the Antarctic. Most species inhabit the Northern Hemisphere during the breeding season, occurring in boreal, sub-arctic, and Arctic regions, and only a few breed in temperate or tropical zones. Northern-breeding species are highly migratory and many winter widely in the tropics and the Southern Hemisphere, from South America and Africa through southern Asia to Australasia.
Some scolopacid waders breed farther north than members of any other bird family. The sanderling (Calidris alba) nests north to Franz Joseph Land, the Zemlya Islands, and the northernmost tip of Greenland, areas not reached by the northernmost-breeding Pluvialis and Charadrius plover species.
All polytypic genera are represented in both Eurasia and North America, generally having different species in each region, but a few species have a circumpolar breeding distribution. Numenius has four North American and three Eurasian breeding species, and one circumpolar species, the whimbrel (Numenius phaeopus). Calidris has seven representatives in Eurasia, four in North America, and seven that breed in both regions. The snipe genus Gallinago has one northern-breeding species, the common snipe (Gallinago gallinago), that breeds across virtually the entire Holarctic region.
Migrations are usually approximately north-south, so that Nearctic breeding species and subspecies typically winter in the Americas and Palearctic breeding taxa in Europe, Africa, Asia, and Australasia. Although many migrant species have very extensive wintering areas, some have very restricted breeding ranges. The western sandpiper (Calidris mauri) that winters coastally in the United States and south through Central America to Peru breeds only around the Bering Strait, while the sharp-tailed sandpiper (Calidris acuminata) that winters throughout Australasia and Melanesia east to Tonga breeds only in a restricted area of north-central and northeast Siberia.
Only 11 species breed in the tropics. All are primarily resident, and most have restricted ranges. They include three woodcock species, the rufous woodcock (Scolopax saturata) of Sumatra, Java, and New Guinea, the Sulawesi woodcock (Scolopax celebensis), and the Moluccan woodcock (Scolopax rochussenii). Seven other species are snipes: the widely distributed African snipe (Gallinago nigripennis), the Madagascar snipe (Gallinago macrodactyla), and five South American species, including the noble snipe (Gallinago nobilis) that is restricted to the northern Andes from Venezuela to Ecuador, and the imperial snipe (Gallinago imperialis) that has a patchy distribution in Peru. The remaining species, the Tuamotu sandpiper (Prosobonia cancellata), occurs only in French Polynesia.
Habitat
Although scolopacid waders occupy a wide range of habitat types in the nonbreeding season, breeding habitats are less diverse and almost all species nest in inland habitats, predominantly in freshwater wetland systems, the only species sometimes breeding at coastal salt marshes being the common redshank (Tringa totanus), Nordmann's greenshank (Tringa guttifer), and the willet (Catoptrophorus semipalmatus).
Species such as the jack snipe (Lymnocryptes minimus) and Gallinago snipes breed in marshes, swamps, floodplains, bogs, and moist grasslands, although some also use more wooded habitats. The wood snipe (Gallinago nemoricola) of central Asia breeds mainly in woodland and scrub. The dowitchers (Limnodromus) and most godwits also breed in marsh, bog, or wet grassy habitats, but the bar-tailed godwit (Limosa lapponica) nests in tundra and woodland. Curlews exploit many breeding habitat types, including taiga, woodland, tundra, moorland, bogs, wet grassland, farmland, prairies, and lakeshores. Most shanks, and the two tattler (Heteroscelus) species, breed in wet wooded and forested areas such as muskeg and taiga. Most Calidris sandpipers breed north of the taiga in the low to middle Arctic tundra zone, while species such as red knot, purple sandpiper (Calidris maritima), and sanderling breed still farther north, in tundra that may be gravelly, stony, or rocky.
Woodcocks inhabit forests and other wooded habitats. The two migratory species, the American woodcock (Scolopax minor) and the Eurasian woodcock (Scolopax rusticola) often breed in deciduous woodland. Both occupy similar habitat in wintering areas, but most other migrant scolopacid species occupy different habitats, and take different food, in breeding and wintering areas. Many species winter at tropical, non-tidal wetlands, including river deltas, floodplains, lakeshores, pans, lagoons, and marshes. Species commonly using such habitats include the curlew sandpiper (Calidris ferruginea), little stint (Calidris minuta), wood sandpiper (Tringa glareola), marsh sandpiper (Tringa stagnatilis), spotted redshank (Tringa erythropus), bar-tailed godwit, and the snipes. In Africa, ruff and black-tailed godwit (Limosa limosa) rely extensively on manmade habitats such as rice fields. Buff-breasted sandpipers (Tryngites subruficollis) winter on pampas and grasslands in South America.
Coastal intertidal sand and mud attract different species, such as curlews, common redshank, terek sandpiper, willet, turnstones, and many Calidris sandpipers. Species such as the greenshank (Tringa nebularia) and other large Tringa species are equally at home in freshwater and marine wetlands. The two phalaropes are pelagic in winter.
The common sandpiper (Actitis hypoleucos) and the spotted sandpiper (Actitis macularia) show little seasonal variation in habitat preferences, both inhabiting margins of waterbodies such as rivers, ponds, and lakes all the year. A few species, notably the rock sandpiper (Calidris ptilocnemis), the purple sandpiper, and surfbird (Aphriza virgata), winter on rocky shores that offer high densities of marine invertebrate prey.
Behavior
Many species are territorial during the breeding season, but others, such as the Asian dowitcher (Limnodromus semi-palmatus), the common redshank, and some godwits and curlews are colonial or semi-colonial. Outside the breeding season most species feed and roost mainly in flocks. Only those that forage visually tend to scatter widely. All species are strong fliers and tightly packed flocks often perform spectacular and complex aerial movements with great precision.
Outside the breeding season, activity patterns often relate to tides, birds feeding by day or night at low tides and roosting at high tides. On the northernmost breeding grounds, where daylight is continuous, activity levels are high and little rest is taken during the six to eight weeks when breeding takes place.
Reasons for this diversity of mating systems may be related to the short breeding season and ability of a single parent to incubate eggs and raise young unaided in situations where food is abundant and predators are few.
Song is an important component of display flights in the family. Snipes, however, accompany display flights by a variety of bleating or winnowing sounds produced by modified tail feathers during high-speed descents. The least vocal species are the phalaropes, which have no elaborate flight display or song, and species such as the ruff, which display in leks on the ground. Most species also have loud alarm calls given when an intruder approaches the breeding territory, while characteristic short alarm calls are uttered to warn of danger and to instigate and coordinate flocking.
Most species that breed at moderate to high latitudes in the Northern Hemisphere undertake extensive southward migrations after breeding. Generally, more northern breeding populations undertake the longest migrations and the temperate breeders the shortest migrations. Many species migrate along well-defined north-south "flyways." Migrants fly in flocks, usually of one species, and may fly at altitudes of over 16,000 ft (5,000 m). These birds can navigate with great precision and, by accumulating very large fat deposits, can make sustained flights of several thousand miles.
Feeding ecology and diet
Scolopacid waders eat predominantly invertebrates, including lumbricid, polychaete, and oligochaete worms; mollusks; crustaceans; arachnids; insects; small fish; and amphibians. On northern breeding grounds most migratory species appear to eat predominantly insects and arachnids, which are abundant. On migration and on wintering grounds, very different animal prey may be taken and some species become dietary specialists. Few species take predominantly plant material, although ruffs and black-tailed godwits periodically eat only rice. When newly returned to the breeding grounds, many species may find animal food scarce and may temporarily supplement their diet with seeds and shoots, while great knots (Calidris tenuirostris) feed predominantly on berries throughout the breeding season. Berries may also have been the staple food of the Eskimo curlew (Numenius borealis) when breeding.
Four principal feeding techniques may be distinguished in the family. Pecking with occasional probing is characteristic of short-billed species of shorelines and relatively dry-land habitats, such as the common sandpiper. Another technique, probing soft substrates, is practiced by snipe, woodcock, and most other species in the family. The third method, running with the bill submerged to chase fish, is used by shank species such as the greenshank and the greater yellowlegs (Tringa melanoleuca). In the nonbreeding season large Tringa species operate in flocks to drive schools of small fish into shallow water.
The fourth technique, rapidly pecking tiny prey from the water, is practiced by species with a short, relatively fine bill— phalaropes and some Calidris species. Phalaropes forage while swimming, when they take shrimps, copepods, and other planktonic animals from the water. They often spin around rapidly, probably to bring deeper prey closer to the surface. Small sandpipers such as the little stint and the western sandpiper often feed by stitching—making rapid vertical pecks into the substrate.
Several species have specialized feeding habits on the nonbreeding grounds. Turnstones turn over stones, shells, and seaweed to locate hidden prey. Mollusks dominate the diet of the surfbird, which hammers or tugs at its prey with its strong, short bill. Other species, such as the red knot, also take many mollusks. The spoon-billed sandpiper feeds like the much larger spoonbills (Platalea, family Threskiornithidae), sweeping its spatulate bill from side to side in the water to catch small prey. The Terek sandpiper makes short sprints to catch small burrowing crabs when they surface to feed or court. Several curlew species take larger burrowing crabs.
Some species hold winter feeding territories and scolopacid waders have been the subjects of classic studies on feeding dispersion and territoriality in relation to prey availability. Results suggest that defense of resources is not tenable at very low or very high food densities, but occurs at intermediate prey densities and may enhance short-term food availability, prevent long-term depletion, and reduce the disturbance of prey.
Reproductive biology
Mating systems in the family are remarkably varied, although most species are monogamous. Polygyny, in which males pair with more than one female, is characteristic of species that display in leks and also occurs in the Eurasian woodcock, the white-rumped sandpiper (Calidris fuscicollis), and the sharp-tailed sandpiper. Polyandrous species (female mates with more than one male) include the spotted sandpiper (which also shows sex-role reversal) and possibly the spotted redshank and the upland sandpiper (Bartramia longicauda). Phalaropes are usually monogamous, though polyandry is recorded in all three species. They also show sex-role reversal: females defend territories and court males that undertake incubation duties and raise the chicks alone.
Many species pair rapidly after arrival on the breeding grounds and begin nesting within a few days. Attraction of mates is often achieved by persistent flight singing, followed by ground displays associated with nest-site selection. Some species, such as Temminck's stint (Calidris temminckii) sing from perches. The ruff, buff-breasted sandpiper and great snipe (Gallinago media) have ground leks at which males display to visiting females and mating occurs.
The nest of most scolopacid waders is a shallow scrape or depression on open ground or in herbage, usually sparsely lined with soft vegetation. A few species make more substantial nests. Baird's sandpiper (Calidris bairdii) builds a cup nest in herbage and Nordmann's greenshanks build substantial twig and lichen nests in trees. Green sandpipers (Tringa ochropus) and solitary sandpipers (Tringa solitaria) use old tree nests of other birds.
The most common clutch size is four eggs, but two to three eggs are laid by dowitchers and by woodcocks and snipes of temperate and tropical latitudes. Scolopacid eggs are pyriform, with a cryptic pattern of brown to black markings on a paler ground. Incubation begins when the clutch is complete and takes about three weeks. Chicks hatch almost simultaneously, are precocial and can walk and feed when a few hours old. They leave the nest within a day of hatching and are brooded and guarded by one or both parents. Only woodcocks and snipes feed their chicks. Parental distraction displays include repetitive display flights, running away like rodents with feathers fluffed like fur and making mammal-like squeaks, and injury-feigning.
Parental duties are shared roughly equally by both sexes in many species, although the female tends to leave chicks earlier. In some species, the female leaves before brood-rearing, or even incubation, is complete. Males of polygynous species undertake no parental care, while in successively polyandrous species, the male raises the first brood while the female may lay and raise another clutch.
When chicks are old enough not to need brooding, the broods of different parents and even different species may form crèches, allowing some adults to desert young early. The remaining attendants desert the young at fledging or immediately afterwards, and most adults migrate south before the young.
In most species, fewer than half the fledglings survive the first year. Although some species breed when one year old, first-year birds of many species remain on wintering grounds and return to breed only when two or three years old. In many Northern Hemisphere breeding areas, breeding success is cyclical and thought to be linked to the population cycles of lemmings (Cricetidae), which in turn influence levels of predation on breeding birds. As a result, curlew sandpipers suffer increased predation of eggs and young by arctic foxes (Alopex lagopus) in years of decreased populations of lemmings, the foxes' normal prey.
Conservation status
Twenty species are of conservation concern, including 10 that are considered Near Threatened. Two species have become extinct since 1600, both from French Polynesia: the white-winged sandpiper (Prosobonia leucoptera) from Tahiti, and Ellis's sandpiper (Prosobonia ellisi) from Moorea. Both probably occurred along highland streams and were exterminated during the eighteenth century, probably by introduced rats. The only surviving member of the genus, the Tuamotu sandpiper, is endangered, having a very small range and population.
Of the two Critically Endangered species, the Eskimo curlew may be extinct as there have been no sightings since the mid-1980s. This small curlew, which bred in Canada and wintered in South America, was hunted close to extinction in the late nineteenth century. Its fate was sealed by the almost complete loss to agriculture of the prairie habitat on which it relied during spring passage. The slender-billed curlew (Numenius tenuirostris), which breeds in Siberia and winters in northwest Africa, has declined dramatically in numbers, the key factor probably being hunting in the nineteenth century, possibly aggravated by habitat loss.
The endangered Nordmann's greenshank breeds in the Russian Far East and winters coastally in south Asia. It is threatened by habitat loss throughout its range, and also by hunting and disturbance. Of the six vulnerable species, the spoon-billed sandpiper is threatened by habitat loss. The bristle-thighed curlew (Numenius tahitiensis) that breeds in western Alaska and winters on oceanic islands in the Pacific (including the Hawaiian islands) suffers heavy losses to introduced predators while flightless during its autumn molt on the wintering grounds. The wood snipe, which breeds in the Himalayas, suffers habitat loss and hunting on its southern Asian wintering grounds. The Chatham snipe (Coenocorypha pusilla) is confined to four small islands off New Zealand and is vulnerable to local extinction by introduced species. Two Asian woodcocks, the Amami woodcock (Scolopax mira), of the Nansei Shoto islands in southern Japan, and the Moluccan woodcock, which is known from only three localities, are threatened by loss of forest habitats.
All ten Near Threatened species, including one woodcock, four snipe, two curlews, the Asian dowitcher, and the buffbreasted sandpiper, are threatened by habitat loss and by hunting. Habitat loss and disturbance are potential threats to many wader species, especially in tropical regions and at temperate staging areas close to large human populations. In northern breeding areas habitat loss may not have been so significant, but has still adversely affected some species. Pressures on habitats are likely to increase, while hunting is still a significant factor and rapid climate change may have significant impacts on habitats and migratory flight routes.
Significance to humans
The common snipe has been closely associated with folklore in Europe and America, and its arrival in spring has been related to events of significance. For example, in Newfoundland it is associated with the arrival of lobsters inshore, while in the United States it is associated with appearance of some commercial fish species. In many parts of Europe the snipe is associated with rain, while some Eskimos of Alaska believe that killing a snipe would bring bad weather.
In Australia, where the return of waders coincides with the first rains of the wet season and is thus associated with good health, one Aborigine tribe carries out "sandpiper dances" as part of its initiation ceremonies. Other species may be associated with sexual appetite, and in the Russian Far East, the lekking behavior of ruffs has impressed the local people of the Chukchi Peninsula so much that they celebrate it with an imitative dance.
The principal significance of scolopacid waders to humans has always been for food and sport. In Europe in the Middle Ages, ruffs were captured on southward passage and were fattened for consumption during winter. Red knots were also highly regarded as a food item. Snipe and woodcock have long been a major quarry of hunters worldwide, and are shot annually in large numbers in Europe and North America. An estimated 1.5 million common snipe are killed by hunters annually in Europe and half a million in North America, while annually up to 3.7 million Eurasian woodcock are shot in Europe and about 2 million American woodcock in North America.
In North America, European immigrants slaughtered an immense number of scolopacid waders in the nineteenth century—godwits, dowitchers, knots, and many other species, even small sandpipers. They were not only trapped and shot, but also blinded by lights at night so that they could be captured by hand. The precipitous decline of the once abundant Eskimo curlew can be attributed to such market hunting, and by the beginning of the twentieth century, this species had become a rarity.
Waders are rarely considered nuisance species, but in some Sahel countries, ruffs and black-tailed godwits are often considered pests because they eat rice. Many species are potentially beneficial by virtue of the insect pests they eat. Upland sandpipers, which breed on North American plains, serve farmers in this way. This role was so unappreciated that in the nineteenth century they were poisoned in the belief that they ate grain.
Species accounts
Eurasian woodcockAfrican snipe
Long-billed curlew
Spotted redshank
Tuamotu sandpiper
Ruddy turnstone
Great knot
Spoon-billed sandpiper
Ruff
Wilson's phalarope
Resources
Books:Del Hoyo, J., A. Elliott, and J. Sargatal, eds. Handbook of the Birds of the World. Vol. 3. Barcelona: Lynx Edicions, 1996.
Higgins, P. J., and S. J. J. F. Davies, eds. Handbook of Australian, New Zealand and Antarctic Birds. Vol. 3. Melbourne: Oxford University Press, 1996.
Matthiessen, P. The Wind Birds. Shorebirds of North America. Shelburne, Vermont: Chapters Publishing Ltd., 1994.
Rosair, D., and D. Cottridge. Hamlyn Photographic Guide To The Waders Of The World. London: Hamlyn, 1995.
Stattersfield, A. J., and D. R. Capper, eds. Threatened Birds Of The World: The Official Source For Birds on the IUCN Red List. Cambridge: BirdLife International, 2000.
Periodicals:Boland, J. M. "An Overview Of The Seasonal Distribution Of The North American Shorebirds." Wader Study Group Bulletin. 62 (1991): 39–43.
Jonsson, P. E., and T. Alerstam. "The Adaptive Significance Of Parental Role Division And Sexual Size Dimorphism In Breeding Shorebirds." Biological Journal of the Linnean Society 41 (1991): 301–314.
Underhill, L. G., R. P. Prys-Jones, E. E. Syroechkovski, et al. "Breeding of Waders (Charadrii) and Brent Geese (Branta bernicla bernicla) at Pronchishcheva Lake, Northeastern Taimyr, Russia, in a Peak and a Decreasing Lemming Year." Ibis 135 (1993): 277–292.
Organizations:African-Eurasian Migratory Waterbird Agreement (AEWA). UN Premises in Bonn, Martin Luther-King Str, Bonn, D-53175 Germany. E-mail: aewa@unep.de Web site:
International Waterbird Census. Web site:
Wader Specialist Group, Mr. David Stroud. Monkstone House House, City Road, Peterborough, PE1 1JY United Kingdom. Phone: +44 1733 866/810. Fax: +44 1733 555/448. E-mail: stroudd@jncc.gov.uk
Wader Study Group, The National Centre for Ornithology. The Nunnery, Thetford, Norfolk JP24 2PU United Kingdom.
Western Hemisphere Shorebird Reserve Network (WHSRN). Manomet Center for Conservation Science, P O Box 1770, Manomet, MA 02345 USA. Phone: (508) 224-6521. Fax: (508) 224-9220. E-mail: info@manomet.org Web site:
Woodcock and Snipe Specialist Group. Director, European Wildlife Research Institute, Bonndorf, Glashuette D-79848 Germany. Phone: 949 7653 1891. Fax: 949 7653 9269. E-mail: wildlife@ewi.t-online.de
Working group on International Wader and Waterfowl Research (WIWO). Stichting WIWO, c/o P O Box 925, Zeist, 3700 AX The Netherlands. Web site:
[Article by: Barry Taylor, PhD]
