The ruff (
Calidris pugnax) is a medium-sized wading bird that breeds
in marshes and wet meadows across northern Eurasia. This highly
gregarious sandpiper is migratory and sometimes forms huge flocks in
its winter grounds, which include southern and western Europe, Africa,
southern Asia and Australia.
The ruff is a long-necked, pot-bellied bird. This species shows marked
sexual dimorphism; the male is much larger than the female (the
reeve), and has a breeding plumage that includes brightly coloured
head tufts, bare orange facial skin, extensive black on the breast,
and the large collar of ornamental feathers that inspired this bird's
English name. The female and the non-breeding male have grey-brown
upperparts and mainly white underparts. Three differently plumaged
types of male, including a rare form that mimics the female, use a
variety of strategies to obtain mating opportunities at a lek, and the
colourful head and neck feathers are erected as part of the elaborate
main courting display. The female has one brood per year and lays four
eggs in a well-hidden ground nest, incubating the eggs and rearing the
chicks, which are mobile soon after hatching, on her own. Predators of
wader chicks and eggs include mammals such as foxes, feral cats and
stoats, and birds such as large gulls, corvids and skuas.
The ruff forages in wet grassland and soft mud, probing or searching
by sight for edible items. It primarily feeds on insects, especially
in the breeding season, but it will consume plant material, including
rice and maize, on migration and in winter. Classified as "least
concern" on the
IUCN Red List
IUCN Red List criteria, the global conservation
concerns are relatively low because of the large numbers that breed in
Scandinavia and the Arctic. However, the range in much of Europe is
contracting because of land drainage, increased fertiliser use, the
loss of mown or grazed breeding sites, and over-hunting. This decline
has seen it listed in the Agreement on the Conservation of
African-Eurasian Migratory Waterbirds (AEWA).
1 Taxonomy and nomenclature
3 Distribution and habitat
4.2 Nesting and survival
5 Relationship with humans
6 Conservation status
7 Biology of variation among males
10 Further reading
11 External links
Taxonomy and nomenclature
A seventeenth-century painting of a woman wearing a ruff, the
decorative collar from which the English name of the bird is derived.
The ruff is a wader in the large family Scolopacidae, the typical
shorebirds. Recent research suggests that its closest relatives are
the broad-billed sandpiper,
Calidris falcinellus, and the sharp-tailed
Calidris acuminata. It has no recognised subspecies or
This species was first described by
Carl Linnaeus in his Systema
Naturae in 1758 as Tringa pugnax. It was moved into its current
genus by German naturalist
Blasius Merrem in 1804. Both parts of
the binomial name refer to the aggressive behaviour of the bird at its
mating arenas; pugnax from the
Latin term for "combative".
The original English name for this bird, dating back to at least 1465,
is the ree, perhaps derived from a dialectical term meaning
"frenzied"; a later name reeve, which is still used for the female,
is of unknown origin, but may be derived from the shire-reeve, a
feudal officer, likening the male's flamboyant plumage to the
official's robes. The current name was first recorded in 1634, and is
derived from the ruff, an exaggerated collar fashionable from the
mid-sixteenth century to the mid-seventeenth century, since the male
bird's neck ornamental feathers resemble the neck-wear.
The ruff has a distinctive gravy boat appearance, with a small head,
medium-length bill, longish neck and pot-bellied body. It has long
legs that are variable in colour but usually yellow or orange. In
flight, it has a deeper, slower wing stroke than other waders of a
similar size, and displays a thin, indistinct white bar on the wing,
and white ovals on the sides of the tail. This species shows sexual
dimorphism. Although a small percentage of males resemble females,
the typical male is much larger than the female and has an elaborate
breeding plumage. He is 29–32 cm (11–13 in) long with a
54–60 cm (21–24 in) wingspan, and weighs about
180 g (6.3 oz). In the May-to-June breeding season, the
typical male's legs, bill and warty bare facial skin are orange, and
he has distinctive head tufts and a neck ruff. These ornaments vary on
individual birds, being black, chestnut or white, with the colouring
solid, barred or irregular. The grey-brown back has a scale-like
pattern, often with black or chestnut feathers, and the underparts are
white with extensive black on the breast. The extreme variability
of the main breeding plumage is thought to have developed to aid
individual recognition in a species that has communal breeding
displays, but is usually mute.
Outside the breeding season, the typical male's head and neck
decorations and the bare facial skin are lost and the legs and bill
become duller. The upperparts are grey-brown, and the underparts are
white with grey mottling on the breast and flanks.
Territorial male in breeding plumage
The female, or "reeve", is 22–26 cm (8.7–10.2 in) long
with a 46–49 cm (18–19 in) wingspan, and weighs about
110 g (3.9 oz). In breeding plumage, she has grey-brown
upperparts with white-fringed, dark-centred feathers. The breast and
flanks are variably blotched with black. In winter, her plumage is
similar to that of the male, but the sexes are distinguishable on
size. The plumage of the juvenile ruff resembles the non-breeding
adult, but has upperparts with a neat, scale-like pattern with dark
feather centres, and a strong buff tinge to the underparts.
Typical adult male ruffs start to moult into the main display plumage
before their return to the breeding areas, and the proportion of birds
with head and neck decorations gradually increases through the spring.
Second-year birds lag behind full adults in developing breeding
plumage. They have a lower body mass and a slower weight increase than
full adults, and perhaps the demands made on their energy reserves
during the migration flight are the main reason of the delayed
Ruffs of both sexes have an additional moult stage between the winter
and final summer plumages, a phenomenon also seen in the bar-tailed
godwit. Before developing the full display finery with coloured ruff
and tufts, the males replace part of their winter plumage with striped
feathers. Females also develop a mix of winter and striped feathers
before reaching their summer appearance. The final male breeding
plumage results from the replacement of both winter and striped
feathers, but the female retains the striped feathers and replaces
only the winter feathers to reach her summer plumage. The striped
prenuptial plumages may represent the original breeding appearance of
this species, the male's showy nuptial feathers evolving later under
strong sexual selection pressures.
Adult males and most adult females start their pre-winter moult before
returning south, but complete most feather replacement on the
wintering grounds. In Kenya, males moult 3–4 weeks ahead of the
females, finishing before December, whereas females typically complete
feather replacement during December and early January. Juveniles moult
from their first summer body plumage into winter plumage during late
September to November, and later undergo a pre-breeding moult similar
in timing and duration to that of the adults, and often producing as
brightly coloured an appearance.
Two other waders can be confused with the ruff. The juvenile
sharp-tailed sandpiper is a little smaller than a juvenile female ruff
and has a similar rich orange-buff breast, but the ruff is slimmer
with a longer neck and legs, a rounder head, and a much plainer
face. The buff-breasted sandpiper also resembles a small juvenile
ruff, but even the female ruff is noticeably larger than the
sandpiper, with a longer bill, more rotund body and scaly-patterned
Distribution and habitat
Illustration of a lek by
Johann Friedrich Naumann
Johann Friedrich Naumann (1780–1857)
The ruff is a migratory species, breeding in wetlands in colder
regions of northern Eurasia, and spends the northern winter in the
tropics, mainly in Africa. Some Siberian breeders undertake an annual
round trip of up to 30,000 km (19,000 mi) to the West
African wintering grounds. There is a limited overlap of the
summer and winter ranges in western Europe. The ruff breeds in
extensive lowland freshwater marshes and damp grasslands. It avoids
barren tundra and areas badly affected by severe weather, preferring
hummocky marshes and deltas with shallow water. The wetter areas
provide a source of food, the mounds and slopes may be used for leks,
and dry areas with sedge or low scrub offer nesting sites. A
Hungarian study showed that moderately intensive grazing of grassland,
with more than one cow per hectare (2.5 acres), was found to
attract more nesting pairs. When not breeding, the birds use a
wider range of shallow wetlands, such as irrigated fields, lake
margins, and mining subsidence and other floodlands. Dry grassland,
tidal mudflats and the seashore are less frequently used. The
density can reach 129 individuals per square kilometre (334 per square
mile), but is usually much lower.
The ruff breeds in Europe and Asia from Scandinavia and Great Britain
almost to the Pacific. In Europe it is found in cool temperate areas,
but over its Russian range it is an Arctic species, occurring mainly
north of about 65°N. The largest numbers breed in Russia (more than 1
million pairs), Sweden (61,000 pairs), Finland (39,000 pairs) and
Norway (14,000 pairs). Although it also breeds from Britain east
through the Low Countries to Poland,
Germany and Denmark, there are
fewer than 2,000 pairs in these more southerly areas.
Wintering in India
It is highly gregarious on migration, travelling in large flocks that
can contain hundreds or thousands of individuals. Huge dense groups
form on the wintering grounds; one flock in
Senegal contained a
million birds. A minority winter further east to Burma, south
China, New Guinea and scattered parts of southern Australia, or
on the Atlantic and Mediterranean coasts of Europe. In Great Britain
and parts of coastal western Europe, where the breeding and wintering
ranges overlap, birds may be present throughout the year.
Non-breeding birds may also remain year round in the tropical
wintering quarters. The
Ruff is an uncommon visitor to Alaska (where
it has occasionally bred), Canada and the contiguous states of the US,
and has wandered to Iceland, Middle America, northern South America,
Madagascar and New Zealand. It has been recorded as breeding
well south of its main range in northern Kazakhstan, a major migration
The male, which plays no part in nesting or chick care, leaves the
breeding grounds in late June or early July, followed later in July by
the female and juveniles. Males typically make shorter flights and
winter further north than females; for example, virtually all
wintering ruffs in Britain are males, whereas in
Kenya most are
females. Many migratory species use this differential wintering
strategy, since it reduces feeding competition between the sexes and
enables territorial males to reach the breeding grounds as early as
possible, improving their chances of successful mating. Male ruffs may
also be able to tolerate colder winter conditions because they are
larger than females.
Birds returning north in spring across the central Mediterranean
appear to follow a well-defined route. Large concentrations of ruffs
form every year at particular stopover sites to feed, and individuals
marked with rings or dye reappear in subsequent years. The refuelling
sites are closer together than the theoretical maximum travel distance
calculated from the mean body mass, and provide evidence of a
migration strategy using favoured intermediate sites. The ruff
stores fat as a fuel, but unlike mammals, uses lipids as the main
energy source for exercise (including migration) and, when necessary,
keeps warm by shivering; however, little research has been conducted
on the mechanisms by which they oxidise lipids.
Ruff in Nederlandsche Vogelen,
Vol. 1 (1770)
Males display during the breeding season at a lek in a traditional
open grassy arena. The ruff is one of the few lekking species in which
the display is primarily directed at other males rather than to the
females, and it is among the small percentage of birds in which the
males have well-marked and inherited variations in plumage and mating
behaviour. There are three male forms: the typical territorial
males, satellite males which have a white neck ruff, and a very rare
variant with female-like plumage. The behaviour and appearance for an
individual male remain constant through its adult life, and are
determined by its genes (see §Biology of variation among males).
The territorial males, about 84% of the total, have strongly coloured
black or chestnut ruffs and stake out and occupy small mating
territories in the lek. They actively court females and display a high
degree of aggression towards other resident males; 5–20
territorial males each hold an area of the lek about 1 m
(1.1 yd) across, usually with bare soil in the centre. They
perform an elaborate display that includes wing fluttering, jumping,
standing upright, crouching with ruff erect, or lunging at rivals.
They are typically silent even when displaying, although a soft
gue-gue-gue may occasionally be given.
Territorial males are very site-faithful; 90% return to the same
lekking sites in subsequent seasons, the most dominant males being the
most likely to reappear. Site-faithful males can acquire accurate
information about the competitive abilities of other males, leading to
well-developed dominance relationships. Such stable relationships
reduce conflict and the risk of injury, and the consequent lower
levels of male aggression are less likely to scare off females.
Lower-ranked territorial males also benefit from site fidelity since
they can remain on the leks while waiting for the top males eventually
to drop out.
Satellite males, about 16% of the total number, have white or mottled
ruffs and do not occupy territories; they enter leks and attempt to
mate with the females visiting the territories occupied by the
resident males. Resident males tolerate the satellite birds
because, although they are competitors for mating with the females,
the presence of both types of male on a territory attracts additional
females. Females also prefer larger leks, and leks
surrounded by taller plants, which give better nesting habitat.
Satellite male with white ruff
Although satellite males are on average slightly smaller and lighter
than residents, the nutrition of the chicks does not, as previously
thought, influence mating strategy; rather, the inherited mating
strategy influences body size. Resident-type chicks will, if provided
with the same amount of food, grow heavier than satellite-type chicks.
Satellite males do not have to expend energy to defend a territory,
and can spend more time foraging, so they do not need to be as bulky
as the residents; indeed, since they fly more, there would be a
physiological cost to additional weight.
A third type of male was first described in 2006; this is a permanent
female mimic, the first such reported for a bird. About 1% of males
are small, intermediate in size between males and females, and do not
grow the elaborate breeding plumage of the territorial and satellite
males, although they have much larger internal testes than the ruffed
males. This cryptic male, or "faeder" (
Old English "father") obtains
access to mating territories together with the females, and "steals"
matings when the females crouch to solicit copulation. The faeder
moults into the prenuptial male plumage with striped feathers, but
does not go on to develop the ornamental feathers of the normal male.
As described above, this stage is thought to show the original male
breeding plumage, before other male types evolved. A faeder can be
distinguished in the hand by its wing length, which is intermediate
between those of displaying males and females. Despite their
feminine appearance, the faeders migrate with the larger 'normal'
lekking males and spend the winter with them.
The faeders are sometimes mounted by independent or satellite males,
but are as often "on top" in homosexual mountings as the ruffed males,
suggesting that their true identity is known by the other males.
Females never mount males. Females often seem to prefer mating
with faeders to copulation with normal males, and normal males also
copulate with faeders (and vice versa) relatively more often than with
females. The homosexual copulations may attract females to the lek,
like the presence of satellite males.
Not all mating takes place at the lek, since only a minority of the
males attend an active lek. As alternative strategies, males can also
directly pursue females ("following") or wait for them as they
approached good feeding sites ("intercepting"). Males switched between
the three tactics, being more likely to attend a lek when the
copulation rate the previous day was high or when fewer females were
available after nesting had started. Lekking rates were low in cold
weather early in the season when off-lek males spent most of their
The level of polyandry in the ruff is the highest known for any avian
lekking species and for any shorebird. More than half of female ruffs
mate with, and have clutches fertilised by, more than one male, and
individual females mate with males of both main behavioural morphs
more often than expected by chance. In lekking species, females can
choose mates without risking the loss of support from males in nesting
and rearing chicks, since the males take no part in raising the brood
anyway. In the absence of this cost, if polyandry is advantageous, it
would be expected to occur at a higher rate in lekking than among
Nesting and survival
Eggs, Collection Museum Wiesbaden
The carrion crow will raid the nests of wetland waders for eggs and
The nest is a shallow ground scrape lined with grass leaves and stems,
and concealed in marsh plants or tall grass up to 400 m
(440 yd) from the lek. Nesting is solitary, although several
females may lay in the general vicinity of a lek. The eggs are
slightly glossy, green or olive, and marked with dark blotches; they
are laid from mid-March to early June depending on latitude.
The typical clutch is four eggs, each egg measuring 44 mm
× 31 mm (1.7 in × 1.2 in) in size and
weighing 21.0 g (0.74 oz) of which 5% is shell. Incubation
is by the female alone, and the time to hatching is 20–23 days, with
a further 25–28 days to fledging. The precocial chicks have buff
and chestnut down, streaked and barred with black, and frosted with
white; they feed themselves on a variety of small invertebrates,
but are brooded by the female. One brood is raised each year.
Predators of waders breeding in wet grasslands include birds such as
large gulls, common raven, carrion and hooded crows, and great and
Arctic skuas; foxes occasionally take waders, and the impact of feral
cats and stoats is unknown. Overgrazing can increase
predation by making nests easier to find. In captivity, the main
causes of chick mortality were stress-related sudden death and twisted
neck syndrome. Adults seem to show little evidence of external
parasites, but may have significant levels of disease on their
tropical wintering grounds, including avian malaria in their
inland freshwater habitats, and so they might be expected to
invest strongly in their immune systems; however, a 2006 study
that analysed the blood of migrating ruffs intercepted in Friesland
showed that this bird actually has unexplained low levels of immune
responses on at least one measure of resistance. The ruff can
breed from its second year, and the average lifespan for birds that
have passed the chick stage is about 4.4 years, although a Finnish
bird lived to a record 13 years and 11 months.
The ruff normally feeds using a steady walk and pecking action,
selecting food items by sight, but it will also wade deeply and
submerge its head. On saline lakes in East Africa it often swims like
a phalarope, picking items off the surface. It will feed at night
as well as during the day. It is thought that
Ruff use both visual
and auditory cues to find prey. When feeding, the ruff frequently
raises its back feathers, producing a loose pointed peak on the back;
this habit is shared only by the black-tailed godwit.
Rice paddies are a favoured winter feeding ground
During the breeding season, the ruff’s diet consists almost
exclusively of the adults and larva of terrestrial and aquatic insects
such as beetles and flies. On migration and during the winter, the
ruff eats insects (including caddis flies, water-beetles, mayflies and
grasshoppers), crustaceans, spiders, molluscs, worms, frogs, small
fish, and also the seeds of rice and other cereals, sedges, grasses
and aquatic plants. Migrating birds in Italy varied their diet
according to what was available at each stopover site. Green aquatic
plant material, spilt rice and maize, flies and beetles were found,
along with varying amounts of grit. On the main wintering grounds
in West Africa, rice is a favoured food during the later part of the
season as the ricefields dry out.
Just before migration, the ruff increases its body mass at a rate of
about 1% a day, much slower than the bar-tailed godwits breeding in
Alaska, which fatten at four times that rate. This is thought to be
because the godwit cannot use refuelling areas to feed on its
trans-Pacific flight, whereas the ruff is able to make regular stops
and take in food during overland migration. For the same reason, the
ruff does not physiologically shrink its digestive organs to reduce
bodyweight before migrating, unlike the godwit.
Relationship with humans
1897 illustration of ruffs being trapped for food with a net
Ruffs were formerly trapped for food in England in large numbers; on
one occasion, 2,400 were served at Archbishop Neville's enthronement
banquet in 1465. The birds were netted while lekking, sometimes being
fattened with bread, milk and seed in holding pens before preparation
for the table.
...if expedition is required, sugar is added, which will make them in
a fortnight's time a lump of fat: they then sell for two Shillings or
half-a-crown a piece… The method of killing them is by cutting off
their head with a pair of scissars [sic], the quantity of blood that
issues is very great, considering the size of the bird. They are
dressed like the Woodcock, with their intestines; and, when killed at
the critical time, say the Epicures, are reckoned the most delicious
of all morsels.
The heavy toll on breeding birds, together with loss of habitat
through drainage and collection by nineteenth-century trophy hunters
and egg collectors, meant that the species became almost extinct in
England by the 1880s, although recolonisation in small numbers has
occurred since 1963. The draining of wetlands from the
1800s onwards in southern Sweden has resulted in the ruff's
disappearance from many areas there, although it remains common in the
north of the country. The use of insecticides and draining of
wetlands has led to a decrease in the number of ruff in Denmark since
the early 1900s. There are still areas where the ruff and other
wetland birds are hunted legally or otherwise for food. A
large-scale example is the capture of more than one million waterbirds
(including ruffs) in a single year from
Lake Chilwa in Malawi.
Although this bird eats rice on the wintering grounds, where it can
make up nearly 40% of its diet, it takes mainly waste and residues
from cropping and threshing, not harvestable grain. It has sometimes
been viewed as a pest, but the deeper water and presence of
invertebrate prey in the economically important early winter period
means that the wader has little effect on crop yield.
The ruff has a large range, estimated at
1–10 million square kilometres (0.38–3.8 million
square miles) and a population of at least 2,000,000 birds. The
European population of 200,000–510,000 pairs, occupying more than
half of the total breeding range, seems to have declined by up to 30%
over ten years, but this may reflect geographical changes in breeding
populations. Numbers in Asia do not appear to be declining, and more
ruffs are wintering in Africa. The species as a whole is therefore not
believed to approach the thresholds for the population decline
criterion of the
IUCN Red List
IUCN Red List (that is, declining more than 30
percent in ten years or three generations). For these reasons, the
ruff is classified as "least concern".
Male in non-breeding plumage in India
The most important breeding populations in Europe, in Russia and
Sweden are stable, and the breeding range in Norway has expanded to
the south, but populations have more than halved in Finland, Poland,
Latvia and The Netherlands. Although the small populations in these
countries are of limited overall significance, the decline is a
continuation of trend towards range contraction that has occurred over
the last two centuries. The drop in numbers in Europe has been
attributed to drainage, increased fertiliser use, the loss of formerly
mown or grazed breeding sites and over-hunting.
Fossils from the
Pleistocene suggest that this species bred further
south in Europe in the cool periods between glaciations than it does
now. Its sensitivity to changing climate as well as to water table
levels and the speed of vegetation growth has led to suggestions that
its range is affected by global warming, and the ruff might act as an
indicator species for monitoring climate change. Potential threats
to this species may also include outbreaks of diseases to which it is
susceptible such as influenza, botulism and avian malaria.
The ruff is one of the species to which the Agreement on the
Conservation of African-Eurasian Migratory Waterbirds (AEWA) applies,
where it is allocated to category 2c; that is, the populations in need
of special attention as they are showing "significant long-term
decline" in much of its range. This commits signatories to
regulate the taking of listed species or their eggs, to establish
protected areas to conserve habitats for the listed species, to
regulate hunting and to monitor the populations of the birds
Biology of variation among males
The ruff has three male forms, which differ in mating behaviour and
in appearance: the typical territorial males which have a dark neck
ruff, satellite males which have a white neck ruff, and the very rare
cryptic males known as "faeders" which have female-like plumage. The
behaviour and appearance of each individual male remain constant
through its adult life, and are determined by a simple genetic
polymorphism. Territorial behaviour and appearance is recessive to
satellite behaviour and appearance, while preliminary research
results suggest that the faeder characteristics are genetically
controlled by a single dominant gene. It was originally thought
that the difference between territorial and satellite males was due to
a sex-linked genetic factor, but in fact the genetic locus relevant
for the mating strategy is located on an autosome, or non-sex
chromosome. That means that both sexes can carry the two different
forms of the gene, not just males. The female does not normally show
evidence of its genetic type, but when females are given testosterone
implants, they display the male behaviour corresponding to their
Skeleton of a ruff
In 2016, two studies further pinpointed the responsible region to
chromosome 11 and a 4.5-Mb covering chromosomal rearrangement.
The scientists were able to show that the first genetic change
happened 3.8 million years ago on the resident chromosome, when a part
of it broke off and was reintroduced in the wrong direction. This
inversion created the faeder allele. About 500,000 years ago another
rare recombination event of faeder and resident allele in the very
same inverted region lead to the satellite allele. The 4.5 Mb
inversion covers 90 genes, one of them is the centromere coding gene
N- CENPN-, which is located exactly at one of the inversion
breakpoints. The inactivation of the gene has severe deleterious
effects and pedigree data of a captive ruff colony suggests that the
inversion is homozygous lethal. Over the course of the past 3.8
million years, further mutations have accumulated within the inversion
i.e. three deletions ranging from 3.3 to 17.6 kb. Two of these
deletions remove evolutionary highly conserved elements close to two
HSD17B2 and SDR42E1-both holding important roles in metabolism
of steroid hormones. Hormone measurements around mating time showed
that whereas residents have a sharp increase of testosterone, faeders
and satellites only experience higher androstenedione levels, a
substance which is considered an intermediate in testosterone
biosynthesis. The authors conclude that one or more of the
deletions act as a cis-acting regulatory mutation which is altering
the expression of one or both genes and eventually contributes to the
different male phenotypes and behaviour.
^ a b
BirdLife International (2015). "
Calidris pugnax". IUCN Red List
of Threatened Species. Version 2015.4. International Union for
Conservation of Nature. Retrieved 23 March 2016.
^ Thomas, Gavin H.; Wills, Matthew A.; Székely, Tamás (2004). "A
supertree approach to shorebird phylogeny". BMC Evolutionary Biology.
4: 28. doi:10.1186/1471-2148-4-28. PMC 515296 .
PMID 15329156. Figure 7 shows relevant relationships.
Supplementary Material[permanent dead link]
^ a b c d e f g h i j Hayman, Marchant & Prater 1986,
^ Linnaeus 1758, p. 148
^ Merrem 1804, no.168 col.542
^ Simpson 1979, p. 883
^ Lockwood 1984, pp. 127–128
^ a b c Cocker & Mabey 2005, pp. 211–212
^ a b c d e f g h Mullarney et al. 1999, p. 156
^ a b c Jukema, Joop; Piersma, Theunis (2006). "Permanent female
mimics in a lekking shorebird" (PDF). Biology Letters. 2 (2):
161–164. doi:10.1098/rsbl.2005.0416. PMC 1618908 .
^ a b c d e f Robinson, R.A. (2009). "
Ruff Philomachus pugnax
[Linnaeus, 1758]". BirdFacts: profiles of birds occurring in Britain
& Ireland (BTO Research Report 407). Thetford: British Trust for
Ornithology. Retrieved 16 April 2009.
^ Lank, David B.; Dale, James (2001). "Visual signals for individual
identification: the silent "song" of Ruffs" (PDF). The Auk. 118 (3):
^ Karlionova, Natalia; Meissner, Włodzimierz; Pinchuk, Pavel (2008).
"Differential development of breeding plumage in adult and second-year
male Ruffs Philomachus pugnax". Ardea. 96 (1): 39–45.
^ Jukema, Joop; Piersma, Theunis (2000). "Contour feather moult of
Ruffs Philomachus pugnax during northward migration, with notes on
homology of nuptial plumages in scolopacid waders". Ibis. 142 (2):
^ a b Pearson, D.J. (1981). "The wintering and moult of ruffs
Philomachus pugnax in the Kenyan Rift Valley". Ibis. 123 (2):
^ Britton, David (August 1980). "Identification of Sharp-tailed
Sandpipers" (PDF). British Birds. 73 (8): 333–345.
^ Vinicombe, Keith (May 1983). "Identification pitfalls and assessment
problems: 4. Buff-breasted
Sandpiper Tryngites subruficollis" (PDF).
British Birds. 76 (5): 203–206.
^ a b Vaillancourt, Eric; Prud'Homme, Sophie; Haman, François;
Guglielmo, Christopher G.; Weber, Jean-Michel (2005). "Energetics of a
long-distance migrant shorebird (Philomachus pugnax) during cold
exposure and running" (PDF). Journal of Experimental Biology. 208 (Pt
2): 317–325. doi:10.1242/jeb.01397. PMID 15634851.
^ a b c d e f g Snow & Perrins 1998, pp. 628–632
^ a b c d "
Calidris pugnax". Species factsheet. BirdLife
International. Retrieved 23 March 2016.
^ Blokhin, Yury Yu. (1998). "Spatial and temporal dynamics of wader
numbers in the delta complexes of the northern subarctic" (PDF).
Wader Studies. 10: 214–220.
^ a b "
Ruff Philomachus pugnax" (PDF). United Kingdom Special
Protection Areas. Joint Nature Conservation Committee. Retrieved 16
April 2009. Estimates for the Russian population vary widely,
and I have used the minimum figure given in Snow & Perrins 1998,
^ Slater 1970, p. 310
^ Johnson, James A.; Lanctot, Richard B.; Andres, Brad A.; Bart,
Jonathan R.; Brown, Stephen C.; Kendall, Steven J.; Payer, David C.
(September 2007). "Distribution of breeding shorebirds on the Arctic
coastal plain of Alaska" (PDF). Arctic. 60 (3): 277–293.
^ "Saryarka – steppe and lakes of northern Kazakhstan". World
United Nations Environment Programme
United Nations Environment Programme and World
Conservation Management Centre. Retrieved 24 March 2016.
^ Khrokov, V. (1988). "Breeding record of
Ruff Philomachus pugnax in
Northern Kazakhstan". Ornitologiya (in Russian). 23: 224–225.
^ a b Baccetti, N.; Gambogi, R.; Magnani, A.; Piacentini, D.; Serra,
L. (1998). "Stop-over strategy of
Ruff Philomachus pugnax during the
spring migration" (PDF). International
Wader Studies. 10:
^ a b Berthold, Bauer & Westhead 2001, pp. 51–52
^ Roulin, Alexandre (2004). "The evolution, maintenance and adaptive
function of genetic colour polymorphism in birds" (PDF). Biological
Reviews. 79 (4): 1–34. doi:10.1017/S1464793104006487.
^ Payne 1984, pp. 13–14
^ Widemo, Fredrik (1997). "The social implications of traditional use
of lek sites in the ruff Philomachus pugnax" (PDF). Behavioral
Ecology. 8 (2): 211–217. doi:10.1093/beheco/8.2.211.
^ a b c Lank, David B.; Smith, Constance M.; Hanotte, Olivier; Burke,
Terry; Cooke, Fred (November 1995). "Genetic polymorphism for
alternative mating behaviour in lekking male ruff Philomachus pugnax"
(PDF). Nature. 378 (6552): 59–62. doi:10.1038/378059a0.
^ Hill, Wendy L. (December 1991). "Correlates of male mating success
in the ruff Philomachus pugnax, a lekking shorebird". Behavioral
Ecology and Sociobiology. 29 (5): 367–372.
^ Hugie, Don M.; Lank, David B. (1997). "The resident's dilemma: a
female choice model for the evolution of alternative mating strategies
in lekking male ruffs (Philomachus pugnax)" (PDF). Behavioral Ecology.
8 (2): 218–225. doi:10.1093/beheco/8.2.218.
^ Lank, David B.; Smith, Constance M. (1992). "Females prefer larger
leks: field experiments with Ruffs (Philomachus pugnax)". Behavioral
Ecology and Sociobiology. 30 (5): 323–329.
^ Hoglund, Jacob; Widemo, Fredrik; Sutherland, William J.; Nordenfors,
Helena (June 1998). "Ruffs, Philomachus pugnax, and distribution
models: can leks be regarded as patches?". Oikos. 82 (2): 370–376.
doi:10.2307/3546978. JSTOR 3546978.
^ Karlionova, Natalia; Pinchuk, Pavel; Meissner, Włodzimierz;
Verkuil, Yvonne (2007). "Biometrics of Ruffs Philomachus pugnax
migrating in spring through southern Belarus with special emphasis on
the occurrence of 'faeders'". Ringing & Migration. 23 (3):
^ Verkuil, Yvonne I.; Jukema, Joop; Gill, Jennifer A.; Karlionova,
Natalia; Melter, Johannes; Hooijmeijer, Jos C.E.W; Piersma, Theunis
(November 2008). "Non-breeding faeder Ruffs Philomachus pugnax
associate according to sex, not morphology: capsule faeders (males
that are female look-alikes) associate with males rather than females,
at several different spatial scales".
Bird Study. 55 (3): 241–246.
^ Hooijmeijer, Jos, ed. (2009). "Faeders are supermen" (PDF).
Ruff Research 2009: 13–14. Archived from the original
(PDF) on 2016-04-12.
^ Lank, David B.; Smith, Constance M. (March 1987). "Conditional
lekking in ruff (Philomachus pugnax)". Behavioral Ecology and
Sociobiology. 20 (2): 137–145. doi:10.1007/BF00572636.
^ a b Lank, David B.; Smith, Constance M.; Hanotte, Olivier; Ohtonen,
Arvo; Bailey, Simon; Burke, Terry (2002). "High frequency of polyandry
in a lek mating system" (PDF). Behavioral Ecology. 13 (2): 209–215.
^ a b Coward 1930, pp. 150–154
^ a b Achim, Johann. "EAZA Husbandry guidelines for the Ruff
(Philomachus pugnax)" (PDF). European Association of Zoos and Aquaria
(EAZA). Archived from the original (PDF) on 10 May 2004. Retrieved 23
^ "Management of wet grassland habitat to reduce the impact of
predation on breeding waders: Phase 2". Science and research projects.
Department for Environment, Food and Rural Affairs
Department for Environment, Food and Rural Affairs (DEFRA). Retrieved
23 April 2009.
^ Bolton, Mark; Tyler, Glen; Smith, Ken; Bamford, Roy (2007). "The
impact of predator control on lapwing Vanellus vanellus breeding
success on wet grassland nature reserves". Journal of Applied Ecology.
44 (3): 534–544. doi:10.1111/j.1365-2664.2007.01288.x.
^ a b van der Wal, R.; Palmer, Stephen C. (2008). "Is breeding of
farmland wading birds depressed by a combination of predator abundance
and grazing?" (PDF). Biology Letters. 4 (3): 256–258.
doi:10.1098/rsbl.2008.0012. PMC 2610041 .
^ Threlfall, William; Wheeler, Terry A. (1986). "Ectoparasites from
birds in Newfoundland". Journal of Wildlife Diseases. 22 (2):
273–275. doi:10.7589/0090-3558-22.2.273. PMID 3712656.
^ a b Mendes, Luisa; Piersma, Theunis; Lecoq, M.; Spaans, B.;
Ricklefs, Robert E. (2005). "Disease-limited distributions? Contrasts
in the prevalence of avian malaria in shorebird species using marine
and freshwater habitats". Oikos. 109 (2): 396–404.
^ Piersma, T. (1997). "Do global patterns of habitat use and migration
strategies co-evolve with relative investments in immunocompetence due
to spatial variation in parasite pressure?". Oikos. 80 (3): 623–631.
doi:10.2307/3546640. JSTOR 3546640.
^ Mendes, Luisa; Piersma, Theunis; Hasselquist, Dennis; Matson, Kevin
D.; Ricklefs, Robert E. (2006). "Variation in the innate and acquired
arms of the immune system among five shorebird species". Journal of
Experimental Biology. 209 (Pt 2): 284–291. doi:10.1242/jeb.02015.
^ Lozano, G.A.; Lank, D.B. (2004). "Immunocompetence and
testosterone-induced condition traits in male ruffs (Philomachus
Animal Biology. 54 (4): 315–329.
^ del Hoyo, Elliot & Sargatal 1996, pp. 530–531
^ Onrust, J.; Loonstra, A. H. J.; Schmaltz, L. E.; Verkuil, Y. I.;
Hooijmeijer, J. C. E. W.; Piersma, T. (2017). "Detection of earthworm
Ruff Philomachus pugnax". Ibis. 159 (3): 647–656.
^ Baccetti, N.; Chelazzi, L.; Colombini, I.; Serra, L. (1998).
"Preliminary data on the diet of migrating Ruffs Philomachus pugnax in
northern Italy" (PDF). International
Wader Studies. 10:
^ Trolliet, B.; Girard, O. (2001). "Numbers of
Ruff Philomachus pugnax
wintering in West Africa" (PDF).
Wader Study Group Bulletin. 96:
^ Piersma, Theunis (1998). "Phenotypic flexibility during migration:
optimization of organ size contingent on the risks and rewards of
fueling and flight?". Journal of Avian Biology. 29 (4): 511–520.
doi:10.2307/3677170. JSTOR 3677170.
^ a b Pennant 1776, p. 460
^ Dahlfors, Steve (2009). "Brushane Philomachus pugnax" (in Swedish).
SOF – Sveriges Ornitologiska Förening. Retrieved 20 April
^ "Brushane" (in Danish). Danish Forest and Nature Agency. 15 May
2009. Archived from the original on January 15, 2012. Retrieved 20
^ Lane, Brett (1993). "Waterbird hunting on the Red River Delta,
Northern Vietnam" (PDF). The Stilt. 22: 51.
^ Hayman, Marchant & Prater 1986, p. 29
^ Balmaki, Behnam; Barati, Ahmad. "Harvesting status of migratory
waterfowl in northern Iran: a case study from Gilan Province"
(PDF). in Boere, Galbraith & Stroud 2006, pp. 868–869
^ Kanstrup, Niels. "Sustainable harvest of waterbirds: a global
review" (PDF). in Boere, Galbraith & Stroud 2006,
^ Trolliet, B.; Girard, O. (1991). "On the
Ruff Philomachus pugnax
wintering in the
Senegal Delta" (PDF).
Wader Study Group Bulletin. 62:
^ Bocheński, Zygmunt (November 2002). "
Bird remains from Obłazowa
– zoogeographical and evolutionary remarks" (PDF). Acta Zoologica
Cracoviensia. 45: 239–252.
^ Zockler, Christoph (April 2002). "Declining
Ruff Philomachus pugnax
populations: a response to global warming?" (PDF).
Wader Study Group
Bulletin. 97: 19–29.
^ "Annex 2: Waterbird species to which the Agreement applies" (PDF).
Agreement on the conservation of African-Eurasian migratory Waterbirds
(AEWA). AEWA. Archived from the original (PDF) on 4 April 2016.
Retrieved 22 April 2008.
^ "Annex 3: Waterbird species to which the Agreement applies" (PDF).
Agreement on the conservation of African-Eurasian migratory Waterbirds
(AEWA). AEWA. Archived from the original (PDF) on 4 April 2016.
Retrieved 22 April 2008.
^ Lank, David. "
Ruff Project". Simon Fraser University. Retrieved 11
^ Lank, D.B.; Coupe, M.; Wynne-Edwards, K.E. (November 1999).
"Testosterone-induced male traits in female ruffs (Philomachus
pugnax): autosomal inheritance and gender differentiation" (PDF).
Proceedings of the Royal Society B. 266 (1435): 2323–2330.
^ Lamichhaney, Sangeet; Fan, Guangyi; Widemo, Fredrik; Gunnarsson,
Ulrika; Thalmann, Doreen Schwochow; Hoeppner, Marc P.; Kerje, Susanne;
Gustafson, Ulla; Shi, Chengcheng (2016-01-01). "Structural genomic
changes underlie alternative reproductive strategies in the ruff
(Philomachus pugnax)". Nature Genetics. 48 (1): 84–88.
doi:10.1038/ng.3430. ISSN 1061-4036. PMID 26569123.
^ a b Küpper, Clemens; Stocks, Michael; Risse, Judith E.; dos
Remedios, Natalie; Farrell, Lindsay L.; McRae, Susan B.; Morgan, Tawna
C.; Karlionova, Natalia; Pinchuk, Pavel (2016-01-01). "A supergene
determines highly divergent male reproductive morphs in the ruff".
Nature Genetics. 48 (1): 79–83. doi:10.1038/ng.3443.
ISSN 1061-4036. PMID 26569125.
Berthold, Peter; Bauer, Hans-Günther; Westhead, Valerie (2001). Bird
migration: a general survey. Oxford University Press.
Boere, G.C.; Galbraith, C.A.; Stroud, D.A. (2006). Waterbirds around
the world. Edinburgh: The Stationery Office.
Cocker, Mark; Mabey, Richard (2005). Birds Britannica. Chatto &
Windus. ISBN 0-7011-6907-9.
Coward, Thomas Alfred (1930). The birds of the British Isles and their
eggs. Volume 1 (4th ed.). Frederick Warne.
del Hoyo, Josep; Elliot, Andrew; Sargatal, Jordi, eds. (1996).
Handbook of the Birds of the World. Volume 3: Hoatzin to Auks.
Barcelona: Lynx Edicions. ISBN 84-87334-20-2.
Hayman, Peter; Marchant, John; Prater, Tony (1986). Shorebirds: an
identification guide to the waders of the world. Boston: Houghton
Mifflin. ISBN 0-395-60237-8.
Liddell, Henry George; Scott, Robert (1980). A Greek-English Lexicon
(Abridged ed.). United Kingdom: Oxford University Press.
Linnaeus, C. (1758). Systema naturae per regna tria naturae, secundum
classes, ordines, genera, species, cum characteribus, differentiis,
synonymis, locis. Tomus I. Editio decima, reformata (in Latin).
Holmiae [Stockholm]: Laurentii Salvii.
Lockwood, W.B., ed. (1984). The Oxford Book of British
Oxford: Oxford University Press. ISBN 0-19-214155-4.
Merrem, Blasius (1804). Allgemeine literatur-zeitung (in German).
Mullarney, Killian; Svensson, Lars; Zetterstrom, Dan; Grant, Peter
Bird Guide. Collins. ISBN 0-00-219728-6.
Payne, Robert B. (1984). Sexual selection, lek and arena behavior, and
sexual size dimorphism in birds (ornithological monographs: No. 33).
Washington, DC: American Ornithologists’ Union.
Pennant, Thomas (1776). British Zoology. Volume 2: Water Birds.
London: B. White.
Simpson, D.P. (1979). Cassell's
Latin Dictionary (5th ed.). London:
Cassell Ltd. ISBN 0-304-52257-0.
Slater, Peter (1970). A field guide to Australian birds:
non-passerines. Adelaide: Rigby. ISBN 0-85179-102-6.
Snow, David; Perrins, Christopher M., eds. (1998). The Birds of the
Western Palearctic concise edition (2 volumes). Oxford: Oxford
University Press. ISBN 0-19-854099-X.
Banks, R.C. (2012). "Classification and nomenclature of the Sandpipers
(Aves: Arenariinae)" (PDF). Zootaxa. 3513: 86–88.
Gibson, R.; Baker, A. (2012). "Multiple gene sequences resolve
phylogenetic relationships in the shorebird suborder Scolopaci (Aves:
Charadriiformes)". Molecular Phylogenetics and Evolution. 64 (1):
Wikimedia Commons has media related to Philomachus pugnax.
Wikispecies has information related to Philomachus pugnax
Wikisource has the text of the 1920
Encyclopedia Americana article
Ruff species text in The Atlas of Southern African Birds
Ruff mating strategy introduction and research
Ruffs on postage stamps: www.birdtheme.org and www.bird-stamps.org
Ageing and sexing (PDF; 2.0 MB) by Javier Blasco-Zumeta &
Calidris pugnax". Avibase.
Ruff media". Internet
Ruff photo gallery at VIREO (Drexel University)
Interactive range map of
Calidris pugnax at
IUCN Red List
IUCN Red List maps
Audio recordings of
Ruff on Xeno-canto.
Philomachus pugnax in the Flickr: Field Guide Birds of the World
Ruff media at
Fauna Europaea: 96818