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This document summarizes a 1997 study on populations, ecology, and threats to the Corncrake (Crex crex) in Europe. It finds that Corncrake populations have declined across most of its European breeding range. Eastern European countries with less intensive agriculture generally have larger populations and higher densities than western countries. Corncrakes prefer tall vegetation for nesting and feeding. Drainage of marshes and conversion of grasslands to other land uses are major threats, as is agricultural intensification. Early mowing of hay and silage fields destroys nests and chicks. Earlier harvesting due to agricultural changes is a key threat driving declines across Europe.

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European Action Plan

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Bogdan Zagavei
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This document summarizes a 1997 study on populations, ecology, and threats to the Corncrake (Crex crex) in Europe. It finds that Corncrake populations have declined across most of its European breeding range. Eastern European countries with less intensive agriculture generally have larger populations and higher densities than western countries. Corncrakes prefer tall vegetation for nesting and feeding. Drainage of marshes and conversion of grasslands to other land uses are major threats, as is agricultural intensification. Early mowing of hay and silage fields destroys nests and chicks. Earlier harvesting due to agricultural changes is a key threat driving declines across Europe.

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This document summarizes a 1997 study on populations, ecology, and threats to the Corncrake (Crex crex) in Europe. It finds that Corncrake populations have declined across most of its European breeding range. Eastern European countries with less intensive agriculture generally have larger populations and higher densities than western countries. Corncrakes prefer tall vegetation for nesting and feeding. Drainage of marshes and conversion of grasslands to other land uses are major threats, as is agricultural intensification. Early mowing of hay and silage fields destroys nests and chicks. Earlier harvesting due to agricultural changes is a key threat driving declines across Europe.

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European Action Plan

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Bogdan Zagavei
This document summarizes a 1997 study on populations, ecology, and threats to the Corncrake (Crex crex) in Europe. It finds that Corncrake populations have declined across most of its European breeding range. Eastern European countries with less intensive agriculture generally have larger populations and higher densities than western countries. Corncrakes prefer tall vegetation for nesting and feeding. Drainage of marshes and conversion of grasslands to other land uses are major threats, as is agricultural intensification. Early mowing of hay and silage fields destroys nests and chicks. Earlier harvesting due to agricultural changes is a key threat driving declines across Europe.

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VOGELWELT 118: 117 – 134 (1997) 117

Populations, ecology and threats to the Corncrake Crex crex in


Europe

Rhys E. Green, Gerard Rocamora & Norbert Schäffer

Green, R. E., G. Rocamora & N. Schäffer 1997: Populations, ecology and threats to
the Corncrake Crex crex in Europe. Vogelwelt 118: 117 – 134.
Declines in the population of the Corncrake have been reported from most states in the
breeding range in Europe. States in eastern Europe with lower intensity of agriculture tend
to have larger populations and higher population densities than states in western Europe.
Information on the ecology of the Corncrake is reviewed. Corncrakes select tall vegetation
that is not so dense that it is difficult for them to penetrate. Marshes and grasslands are
important habitats, so drainage, conversion to other uses and agricultural intensification
are threats to Corncrake populations. In most states many Corncrakes occupy grassland
managed for the production of hay or silage. The nest is placed on the ground and the
chicks are reared by the W in tall vegetation. W are capable of producing two broods of
young in one summer, so the potential breeding season is long (May - August). Mowing
hay and silage during the breeding season causes destruction of nests and chicks. The ear-
lier that mowing occurs, the larger the negative impact on Corncrake breeding success.
Agricultural development leading to earlier and more rapid harvesting of hay and silage is
therefore another major threat to Corncrake populations in Europe.
Keywords: Crex crex, population trends, grassland management, breeding biology, Europe.

1. Introduction
The Corncrake Crex crex (L.) is a rail (Gruiformes: trends, ecology, threats and future prospects for the
Rallidae) that is considered to be declining in num- species. Questionnaires were received from 34 states,
bers in most of its world range (COLLAR et al. 1994; though some questions were not answered for some
TUCKER & HEATH 1994). As a result, the species is states because of lack of the required information.
listed as being threatened with global extinction
(COLLAR et al. 1994; GROOMBRIDGE 1994) and is thus 2. Geographical range
of high priority for conservation action. Although
there is a substantial literature on the ecology of the Corncrakes formerly bred over much of northern and
Corncrake and the likely reasons for its decline are central Europe between about 41° N and 62° N,
known, there has been no review of its conservation extending into Asia in western Siberia as far as 120°
biology. In 1994 BirdLife International co-ordinated E. The distribution is now much restricted within the
such a review as part of preparations for a Species former range, and is fragmented in western and cen-
Action Plan to assist the recovery of European Corn- tral Europe. Corncrakes are found from sea level up
crake populations (CROCKFORD et al. 1996). This paper to 1,400 m a.s.l. in the Alps and to 3,000 m a.s.l. in
summarizes the results of the review. We avoid, as Russia. Corncrakes winter mainly in southern Africa.
much as possible, the duplication of general accounts Autumn migration appears to be concentrated through
of the biology of Corncrakes given in standard texts Egypt (STOWE & BECKER 1992).
(GLUTZ VON BLOTZHEIM et al. 1973; CRAMP & SIMMONS
1980). Instead we concentrate on recent findings 3. Population size
about the ecology of the species of direct relevance
3.1. Source of data
to conservation management.
This paper is based on reviews of published Most questionnaires gave estimates of the Corncrake
papers, unpublished reports and theses, on discussions population of the range state as a whole and two thirds
with ecologists who have studied Corncrakes and a of them also gave details of estimates and trends for
questionnaire which was sent to ornithologists in 35 certain regions within the state. Reports from Mace-
states in the Corncrake’s European breeding range. donia and Turkey stated that Corncrakes are no longer
Questions were asked about population size and thought to be breeding there. No information was
118 R. E. GREEN et al.: Populations, ecology and threats to the Corncrake in Europe

obtained from Albania. For each population estimate, M in Russia (over 400,000), Belarus (26,000-30,000),
a reliability code was given ranging from 0 (a guess) and probably Ukraine (25,000-55,000) and several
to 3 (probably accurate to 10 %). thousand each in Poland, Latvia, Lithuania, Estonia,
The main method used to count Corncrakes is Bulgaria and Romania, although several of these
night-time counts of singing M. Full counts through- countries have data of poor reliability (Table 1). The
out the national breeding range were used in Belgium, uncertainty concerning the estimates for Russia, the
France, Hungary, Moldova, Netherlands, Slovenia, Baltic States and the Ukraine must be emphasised.
Spain, Sweden and United Kingdom, all of which Additional nationwide surveys are in progress in Bul-
have relatively small populations. In some countries garia, Poland and Romania. All surveys have been
full counts were used in some regions while in Au- funded by the Royal Society for the Protection of
stria, Belarus, Bulgaria, Czech Republic, Estonia, Birds (RSPB). Finland, Moldova, Slovakia,and Slo-
Finland, Germany, Ireland, Italy, Poland, Slovakia venia have over 500 singing M and Croatia and Bos-
and Russia numbers were estimated from counts in nia-Herzegovina probably hold similar numbers.
sample areas. Only in a few cases (e. g. Ireland) were Hundreds of singing M are found in the Czech Repub-
these estimates based on strictly random sample lic and Hungary.
areas. Daytime counts have also been used in some In western Europe estimates are more precise.
areas in Lithuania, Austria, Norway, France, Ireland France holds the largest population with over 1,000
and Slovenia as well as in Belarus as the main method M, Germany has the second largest with about 800
before 1994. Many countries also used reports from singing M. Populations from the Great Britain, Italy,
local people: Lithuania, Ukraine, Austria, Sweden, Sweden, Austria and Ireland are significant in size
Belarus, Norway, Switzerland, Ukraine, Spain, Den- (>100), but those from other countries can only be
mark, Estonia, Slovenia and Italy. In Latvia use was considered as relicts (Belgium, Denmark, the Nether-
made of point counts and transects. lands, Luxembourg, Norway, Spain and Switzerland.)
3.2. Population estimates 3.3. Population size in relation to the size of the
country
Table 1 gives the breeding population estimate availa-
ble in 1995 for each European state, together with its Large countries would be expected to hold more
accuracy as assessed by respondents. For Latvia the Corncrakes than small countries, so numbers were
more recent 1996 estimate (KEISS 1997) was used. As compared with the land area of the state. Most coun-
Luxembourg and Romania provided no population tries provided a range rather than a single figure for
estimates in the questionnaire, numbers from TUCKER the number of singing Corncrakes present (Table 1).
& HEATH (1994) were used. However, assuming that these ranges include the true
The European population excluding Russia was number of birds, the Corncrake population of each
estimated at 105,000 - 138,000 singing M from the state could be placed in a category according to its
questionnaires based on 1985-1995 data (Table 1). order of magnitude, that is, 1 = 1 - 10 birds, 2 = 11 -
However, the biggest part of the European population 100 birds etc. These scores were plotted against the
is concentrated in Russia west of the Ural mountains. logarithm of the surface area of the country (Figure
For this large area reasonable estimates have only 1). In Norway, Sweden, Finland, European Russia,
now become available (MISCHENKO et al. 1997), indi- Spain and Italy the range is thought to be constrained
cating 260,000 - 400,000 M in nine regions surveyed by climate and only the land surface area within the
and yielding an estimate of the total population as historical latitudinal limits of the Corncrake was used.
high as 1 Mio. M. Obviously, these data are not com- The variation among states in Corncrake numbers
parable to earlier estimates which were not based on is considerably larger than would be expected from
survey data. variation in the size of the states alone (Fig. 1). For
For Europe excluding Russia, our new estimates example, states 10,000 - 100,000 km2 in extent have
are somewhat higher than the 82,000 - 100,000 M Corncrake populations spanning four orders of mag-
given by TUCKER & HEATH (1994). The questionnaires nitude. The deviations from the general relationship
yielded more precise data than that available to between Corncrake population size and surface area
TUCKER & HEATH for nine states. There were large differ according to the recent political history of the
differences between the two sets of estimates for state. Eastern European states that, until recently, had
Belarus (26,000-30,000 instead of 55,000-60,000), planned economies tended to have higher Corncrake
Ukraine (25,000-55,000 instead of 2,000-2,500), Bul- populations, after allowing for surface area, than we-
garia (1,000-2,500 instead of 100-1,000), Lithuania stern states with capitalist or mixed economies. An
(3,000-4,000 instead of 2,000-3,000) and Germany index of Corncrake density was obtained by subtrac-
(800 instead of 260). ting the logarithm (base 10) of land surface area from
The largest Corncrake populations are in central the Corncrake population order of magnitude. This
and eastern Europe, with more than 10,000 singing index was significantly higher for eastern states than
VOGELWELT 118: 117 – 134 (1997) 119

for western states (MANN- Tab. 1: Recent estimates (mostly from 1994) of Corncrake populations (number of
WHITNEY U-test; P < singing males) in 33 European range states. The accuracy of estimates was assessed by
0.001), regardless of whe- contributors on a scale from 0 (a guess) to 3 (a census accurate to 10 % of the true
ther the unified Germany number). The population trend over the 10 years prior to the most recent estimate is
given as -2 (decrease of >50 %), -1 (decrease of 20 - 49 %), 0 (change of <20 %), F
is treated as eastern or
(fluctuating, with changes of >20 % but no clear trend). - Schätzungen (Stand 1994,
western. This difference außer für Lettland und Rußland) der Wachtelkönigbestände (rufende M) in 33 europäi-
may be attributable to dif- schen Ländern. Die Genauigkeit der Angaben wurde von den Bearbeitern mit 0 (Vermu-
ferences in farming sys- tung) bis 3 (Zählung mit einem Fehler von weniger als 10 % der tatsächlichen Anzahl)
tems, but climate and other angegeben. Weiterhin ist die Bestandsentwicklung der vergangenen 10 Jahre in ver-
factors may also be invol- schiedene Kategorien eingeteilt: -2 (Rückgang >50 %), -1 (Rückgang 20 - 49 %), 0
ved. Using the Corncrake (Änderung <20 %), F (schwankend, >20 %, aber kein klarer Trend).
population data published
by TUCKER & HEATH range state number of accuracy year of trend
(1994), GREEN & RAYMENT singing M code estimate
rufende M Genauigkeit Jahr
(1996) found that variation
among 17 European states Austria 140 - 180 2 1989 - 91 (-1)
in the average population
Belarus 26,000 - 30,000 2 1990 -1
density of Corncrakes was
Belgium 17 - 21 2-3 1992 - 94 F
negatively correlated with Bosnia 300 - 1,000 1 1987 ?
three indices of the inten-
Bulgaria 1,000 - 2,500 *2 2 1980 - 94 -1
sity of agriculture (the ave-
Croatia 400 - 1,000 1 1990 ?
rage yield of milk from Czech Republic 200 - 400 1 1985 - 89 -1
dairy cattle and indices of
Denmark 6 3 1991 -2
the prevalence of tractors
Estonia 5,000 2 1993 -1
and of fertilizer use). In Finland 500 - 1,000 2 1994 0
that study the relationship
France 1,100 - 1,200 3 1991 - 92 -1
between Corncrake den-
Germany 800 2 1994 0
sity and agricultural inten- Great Britain 480 3 1993 -1
sity was significant even if
Hungary 350 - 450 2 1993 - 94 F
the west-east trend was
Ireland *1 174 3 1993 -2
allowed for using partial Italy 250 - 300 0 1994 ?
correlation.
Latvia 26,000 - 38,000 3 1996 -1
3.4. Main Corncrake Liechtenstein 8 3 1994 0
populations within Lithuania 3,000 - 4,000 2 1994 ?
countries Luxembourg <10 0 - -1
Moldova 450 2 1985 -1
The regions with important Netherlands 50 - 150 2 1989 - 91 -2
concentrations of Corn- Norway 15 - 35 1-2 1994 -2
crakes were identified for Poland 6,600 - 7,800 *2 1 1993 -2
most countries as follows. Romania 3,000 - 6,000 *2 0-1 - ?
Lower Austria, Styria and Russia 400,000- 1 Mio.*3 2 1995 -1
Vorarlberg (Austria); Serbia-Montenegro >100 0 1991 ?
Fagne and Famenne (Bel- Slovakia 600 - 900 1 1992 ?
gium); Livansko Polje Slovenia >500 3 1992 - 93 -1
(Bosnia-Herzegovina); So- Spain 24 - 31 2 1993 - 94 -1
fia and Burgas regions, Sweden 250 - 1,000 0 - ?
along the Black Sea coast, Switzerland 4 1 1993 0
Smoljan region, the Bal- Ukraine 25,000 - 55,000 1 1993 -1
kans range (mainly bet-
ween 800 and 1400 m asl), total 505,000 – 1,1 Mio.
Dobrudja and Trakia plain
*1
(Bulgaria); alluvial wet- includes 9 M in Northern Ireland (part of United Kingdom) – einschließlich 9
lands of the Sava River (in M in Nordirland (Teil des U.K.).
*2
cluding Turopolje, Lonjs- nation-wide surveys in progress – landesweite Kartierung in Bearbeitung.
ko, Mokro, Jelas Polje), *3
new estimate: 1 million (Central European Russia); 2-2.5 million (Russia)
Drava and Danube flood- (MISCHENKO et al. 1997). – Neue Schätzung: 1 Million (Central European
plains and NE of Karlovac Russia); 2-2.5 Millionen (Rußland) (MISCHENKO et al. 1997).
120 R. E. GREEN et al.: Populations, ecology and threats to the Corncrake in Europe

Ebro depression in Catalonia (Spain); Oland, Got-


land and Uppland (Sweden); Neuchatel and Vaud
cantons of the Jura Mountains (Switzerland); NW
and central Ukraine (Ukraine); Outer Hebrides, Ork-
ney, Skye, Coll and Tiree in Scotland (United King-
dom). Main concentrations were not identified for
Russia, Belarus, Estonia, Latvia, Romania and Slo-
vakia where the species is quite widespread and there
was relatively little survey information.
3.5. Long-term population trends
Declines in European Corncrake populations were
reported as long ago as the latter half of the 19th
century in Denmark, Germany, Britain and Ireland
(NORRIS 1947; VON HAARTMAN 1958; GREEN 1995). In
France the decline of the Corncrake was reported in
several regions early this century (BROYER 1985). HOL-
Fig. 1: Recent estimate of numbers of singing M Corncrakes GERSEN (1958) noted a decline in numbers in Norway.
in relation to the land surface area of European range states. In central European Russia declines were first repor-
Former communist states are indicated by triangles and ted around 1930 (SPANGENBERG & OLIGER 1949; PTU-
other states by circles. The unified state of Germany is SHENKO & INOZEMTSEV 1968) and became widespread
plotted as a circle with dot. Land north of 62° N and south and obvious in the 1950s and 1960s (PTUSHENKO &
of 41° N was excluded from the area estimates. - Schät- INOZEMTSEV 1968; BALDAEV 1973; MALCHEVSKIY & PU-
zung des Wachtelkönigbestandes (rufende Männchen) in KINSKY 1983). Long term declines were reported in
Abhängigkeit von der Flächengröße europäischer Länder. the questionnaire survey from numerous countries:
Ehemals kommunistische Länder sind als Dreiecke, andere
als Kreise dargestellt. Der Sonderfall von Deutschland ist
in Austria over the past 30-40 years, in Flanders (Bel-
als Kreis mit Punkt angegeben. Gebiete nördlich 62° N und gium) between 1950 and 1960, in the Czech Repub-
südlich 41° N wurden nicht berücksichtigt. lic, mainly between the 1950s and 1970s, in Germany
and Macedonia over the last 15 - 20 years and in
Switzerland between 1960 and 1980.
among the River Kupa (Croatia); Sumava, Krkonose, The countries which show stable or fluctuating
Jeseniky, Beskydy (Czech Republic); Northern trends during the past 5 - 10 years either experienced
Jutland: Bornholm, Sealand (Denmark); SE Finland, large declines previously (Finland, Germany, Bel-
especially S Karelia, and the southern coast (Finland); gium and Switzerland), or have only been properly
Basses Vallées Angevines, Loire valley, Charente surveyed recently (Hungary and Sweden) and hence
valley, Saone valley and Carentan marshes in reliable data on trends are not available. However,
Normandy (France); Unteres Odertal in Brandenburg, Hungary was classified as a country with strong decli-
Murnauer Moos and vicinity in Bayern, and NW Nie- nes by TUCKER & HEATH (1994). Thus virtually all
dersachsen (Germany); Bodrogkoz, Hansag, Bodva- European range states appear to have experienced
volgy (Hungary); Shannon Callows, North Donegal long term declines in Corncrake populations.
and Moy valley (Ireland); Udine province, Friuli-Ve- 3.6. Recent population trends
nezia Giulia region, Veneto region- especially
Vicenza and Treviso provinces (Italy); Ruggeller Riet Questionnaire respondents were asked to describe the
(Liechtenstein); Nemunas valley and delta, Jura overall trend in their national Corncrake population,
Valley, Central Lithuanian Plain, Katra Valley by selecting from several statements the one best des-
(Lithuania); near Fentage, Erpeldange/Bous and Pis- cribing their situation and giving comments. Answers
sange (Luxembourg); NW, E and SW Moldova, espe- could be based on information on numbers or, where
cially the middle Prut valley (Moldova); Rhine-Waal- that was not possible, on geographical range. Infor-
Yssel area and NE Groningen province (Netherlands); mation was provided in questionnaires for 24 coun-
Co Rogaland, Møre and Romsdal, Akershus and Bus- tries on the five year trend (5YT) and/or ten year
kerud (Norway); Narew valley, Biebrza valley, trend (10YT) of their Corncrake populations. Trends
Przemysl and Krosno areas, foothills of Bieszczady were classified according to the scale of TUCKER &
mountains (Poland); Vojvodina, especially the Sava HEATH (1994) as -2 (decrease of >50 %), -1 (decrease
and Danube valleys, Lake Plavsko jezero at the head of 20 - 49 %), 0 (change of <20 %), or F (fluctuating,
of the Lim River (Serbia-Montenegro); Lake Cer- with changes of >20 % but no clear trend). For the
knisko, Ljubljansko barje, West Julian Alps, Pla- 22 countries providing information on the 10YT, the
ninsko polje, Reka-valley (Slovenia); Cinca basin and average score was -0.95, which is equivalent to a de-
VOGELWELT 118: 117 – 134 (1997) 121

cline of 20-50 %. Of the 16 countries which supplied (Moldova, Romania, Italy). The regions where the
information on the 5YT, the average score was only largest declines are thought to have occurred in the
-0.44, with certain countries (Czech Republic, Den- last five, 10 or 20 years are: Rhine delta and Lake
mark, Estonia, Russia) which had reported a decline Constance (Austria); Carentan marshes and Loire
over the past ten years reporting population stability valleys (France); N Donegal and Mayo (Ireland);
(code 0) over the last five years. Groninjer/Friesland (Netherlands); Rogaland, Møre
Table 1 summarises the data gathered through the and Romsdal (Norway); Malopolska (Poland); Cadca,
questionnaire on overall trends of Corncrake popu- Vranov (Slovakia); Gotland (Sweden); Orkney, Skye,
lations within the last ten years. For the five countries Islay, Northern Ireland (UK).
with insufficient data from the questionnaire (Austria, Regions where more moderate declines have been
Croatia, Lithuania, Romania and Switzerland) data reported are: Saone and Charente valleys (France);
from TUCKER & HEATH (1994) for trends in the period Eastern Rodopi, Sakar (Bulgaria); Shannon Callows
1970-1990 have been used. A decline has occurred (Ireland); Akershus and Buskerud coastal areas (Nor-
over most of Europe within the last ten years. Only way); Biebrza (Poland); Moscow region (Russia);
Liechtenstein reported an increase in its very small Zahorie (Slovakia); Lubljansko barje (Slovenia); and
population. Several countries such as Ireland, Nether- central Ukraine.
lands, Denmark, Norway and Poland have experienced Regions where populations appear to be fluctua-
recent declines of more than 50 %; whereas Italy, Swit- ting, stable or increasing are: Sofia region (Bulgaria);
zerland, Hungary, Belgium, Germany, Sweden and Wallonia and Flanders (Belgium); Sumava, Krkonose
Finland appear to have stable or fluctuating populations in Czech Republic; Basses Vallées Angevines
in this period. All other countries appear to have expe- (France); Untere Oder, Murnauer Moos (Germany);
rienced declines of less than 50 %. Kombinat Wizna, Narew (Poland); Dolny, Kubin, Hu-
Detailed surveys have demonstrated the alarming menné (Slovakia); Cerknica and Planinsko polje (Slo-
extent of declines in several countries. The best docu- venia); Olland, Uppland, Delarm (Sweden); Lipetsk
mented examples are presented in Figure 2. In Great (Russia); Lewis, Coll and Tiree (UK).
Britain there has been a decline of 40% between 1978
and 1993, including a 17% decline during 1988-93
(GREEN 1995). In Ireland the declines were even more 4. Ecology
severe over the same period with a decline of 81%
4.1. Habitat selection
between 1988 and 1993 (SHEPPARD & GREEN 1994). In
France a decline of about 40% in numbers and range Corncrakes are found in drier habitats than most other
has been observed between 1984 and 1992. In western rails. Breeding occurs in open or semi-open
Ukraine probably more than 60%
of the population has disappeared
since 1976, including a 10 %
decline during the period 1988-93.
In the Netherlands the population
has been reduced by more than
75 % since the early 1980s. In
Belarus, the decline has probably
been greater than 10 - 15 % in the
last 10 years.
3.7. Variations in population
trends within countries
The majority of countries reported
obvious variations in population
trends between parts of the country.
Except Estonia, the ones where Fig. 2: Examples of changes in numbers of singing M Corncrakes in states or
such variations were not mentioned large regions. (Reliability of the count: France = 3; Great Britain = 3; Hungary
are either countries with relict or = 2; Ireland = 3; Sweden = 0; western Ukraine = 1; see Table 1). Fr = France,
small populations (Switzerland, GB = Great Britain, Hu = Hungary, Ir = Ireland (Republic and Northern
Ireland), Sw = mainland Sweden, WU = Westukraine. - Beispiele für die
Luxembourg, Macedonia, Spain, Bestandsänderungen von rufenden Wachtelkönigen in Ländern oder großen
Netherlands, Norway) or those Gebieten (Zuverlässigkeit der Zahlen: Frankreich = 1; Ungarn = 2; Irland,
where the total population appea- Großbritannien = 3; Schweden = 0; Westl. Ukraine = 1; siehe Tabelle 1). Fr
red stable (Hungary, Finland) or = Frankreich, GB = Großbritannien, Hu = Ungarn, Ir = Irland (Republik
where very few data are available und Nordirland), Sw = Festland Schweden, WU = Westukraine.
122 R. E. GREEN et al.: Populations, ecology and threats to the Corncrake in Europe

Tab. 2: Principal habitats of Corncrakes in 30 range states as indicated by hay (CRAMP & SIMMONS 1980).
responses to a questionnaire. The importance of each habitat is denoted by a Suitable habitats include moist, un-
score from 0 (not used) to 3 (most birds in this habitat). The table has been fertilized grassland and regularly
summarised by counting, for each habitat, the number of states in which it cut meadows in areas with agricul-
was given the highest or equal highest score. Some other habitats were recor-
tural methods of low intensity.
ded in a few states and these are discussed in the text. – Hauptsächlich ge-
nutzte Habitate des Wachtelkönigs in 30 europäischen Staaten (Ergebnisse Such areas include wetland areas
nach Fragebögen). Die Wichtigkeit jedes Habitats wird mit einem Wert zwi- along rivers and marshes, the frin-
schen 0 (nicht genutzt) und 3 (die meisten Vögel halten sich in diesem Habi- ges of moorland (peatland, mires),
tat auf) angegeben. Die letzte Zeile gibt für jedes Habitat die Anzahl der meadows on the margin of standing
Länder wieder, in denen der Skalenwert für eine Zeile am höchsten oder gleich water, cleared grassy areas in fo-
den benachbarten Spalten/Habitaten ist. Einige Habitate kommen nur ver- rests and mountain pastures. The
einzelt vor und werden im Text diskutiert. species is found on abandoned
agricultural land, but changes in
Range dry floodplain alpine marshes crops vegetation may make such habitats
state meadows meadows meadows
unsuitable after a few years (see
Austria 1 1 3 1 1 below). Singing M can regularly be
Belarus 2 2 0 2 1 heard in fertilized meadows or
Belgium 2 2 0 1 1 fields sown with cereals, but suc-
Bulgaria 1 3 3 2 1 cessful reproduction is believed to
Czech Republic 0 0 2 0 1 be infrequent. In these habitats
Denmark 0 3 0 0 2-3
Estonia 2 2 0 1 1
herbaceous vegetation on field
Finland 2 0 0 1 2 margins or fallow land may be used
France 0 3 1 1 0 as alternative habitats during the
Germany 1 2 0 1 1 harvest.
Great Britain 2 2 0 2 0 The importance of various habi-
Hungary 1 2 0 1 1 tats for Corncrakes, as reported in
Ireland 2 2 0 0 0 the questionnaire survey, is shown
Italy 1 0 3 0 0 in Table 2. Hay or silage fields in
Latvia 2 3 0 0 2 valleys liable to flooding seem to
Liechtenstein 0 1 0 3 0 be of highest importance, but the
Lithuania 3 2 0 1 1
Luxembourg 0 1 0 1 1
birds also breed in hay/silage
Moldova 2 1 0 1 1 fields in dry lowlands and in mar-
Netherlands 1 3 0 1 1 shes, crops or in hay/silage fields
Norway 2 1 0 1 2 in (sub-)alpine areas (listed in
Poland 2 2 1 1-2 0-1 order of decreasing importance).
Romania 3 0 0 0 1 M are regularly found singing in
Russia 1 3 0 1 1 clear cuts in forests in Russia,
Slovakia 0 2 3 0 1 Lithuania, Belarus, Poland and
Slovenia 1 3 2 1-2 0-1 Belgium, in pastures in Lithuania
Spain 0 1 0 2 3 and Latvia, and on abandoned
Sweden 2 0 0 2 0-1
Switzerland 1 0 2 2 0
land in Slovenia, Czech Republic,
Ukraine 2 3 0 2 0 Denmark, Austria and Germany.
Detailed information on habitat
number with selection and breeding biology has
top score 12 17 6 7 4 come from studies which used
radio-tracking (STOWE & HUDSON
landscapes, mainly in meadows of tall grass. The 1991; GRABOVSKY 1993; SCHÄFFER & MÜNCH 1993;
species is almost always concealed in tall vegetation. TYLER 1996; TYLER & GREEN 1996). Two important
Tall vegetation around single bushes and hedgerows, features of selected habitats are that they are tall
stands of tall herbs, reeds or other tall marsh plants enough to provide cover (>20cm) and that they are
are used as singing places. The original breeding not so dense that they are difficult for the birds to walk
habitats of the Corncrake would almost certainly have through (TYLER 1996; SCHÄFFER 1997). In most habitats
been riverine meadows of Carex-Iris-Typhoides and selected by Corncrakes much of the annual production
alpine, coastal and fire-created grasslands with few of plant material is removed each year by mowing,
trees or bushes present. grazing or winter floods. Stands of dead grasses from
Today Corncrakes are strongly associated with the previous year’s growth tend to be avoided by Corn-
agricultural grassland managed for the production of crakes. It is believed that, though sufficiently tall, the
VOGELWELT 118: 117 – 134 (1997) 123

dense mat and tussock structure renders them un- 4.2. Diet and foraging
suitable. Other vegetation with closely spaced and
In the breeding season the food of the Corncrake in-
robust stems or leaves, such as tussock-forming species
cludes a wide range of invertebrates, including taxa
of Juncus may also be unsuitable for this reason.
living on plants, on the soil surface and in the soil.
Closely spaced grasses in fertilised meadows may also
Small vertebrates, such as fish and amphibians, are
be too dense, and are certainly so if they are pushed to
also taken occasionally. In Scotland and Ireland earth-
the ground by wind and rain. On nutrient-rich soils,
worms and molluscs are an important part of the diet
the vegetation on abandoned meadows and other farm-
(TYLER 1996) whereas in Poland insects are the most
land tends to become too dense for Corncrakes within
common prey (SCHÄFFER 1997). The availability of
a few years. However, the species may remain for deca-
these prey in moist habitats may be another reason
des after abandonment if the soil is poor, as in the
that moist grassland is good habitat for Corncrakes in
Bohemian Forest, Czech Republic.
addition to the suitability of the vegetation structure
In some parts of the breeding range the distri-
of some marsh vegetation. The principal prey (beetles,
bution of singing Corncrakes, and probably also their
other large insects, earthworms, snails, slugs) are wide-
abundance, is strongly affected by the availability of
spread in habitats other than those used by Corncrakes,
suitable vegetation in spring. In western Scotland in
so it appears that the species has specialised more in
May and early June too little cover is provided by
the structure of the vegetation that it occupies than in
grass in hay and silage meadows. At this time stands
the food it takes. Studies of foraging methods have
of tall herbs, such as Urtica dioica and Anthriscus
not been made, but birds are active and apparently
sylvestris and marsh vegetation (Iris pseudacorus,
foraging only by day and mostly in cover. It is presu-
Phragmites australis, Phalaris arundinacea) are
med, from the structure of the bill, that prey is pecked
selected (CADBURY 1980). Only when the height of
from vegetation and the soil surface.
grass in the meadows exceeds about 20 cm (in mid
Some seeds are taken in the breeding season
June) are they much used. In Scotland and Ireland
(CRAMP & SIMMONS 1980), but it is uncertain whether
GREEN (1996) found that Corncrakes occurred where
they are ingested directly, or in the guts of invertebrate
a sufficient area of vegetation of a suitable height
prey, or in place of grit. Corncrakes in captivity were
and structure in spring occurred close to areas of hay
observed to change their diet from invertebrates to
and silage meadows which did not become suitable
seeds in autumn, even when there was no change in
until later in the summer. However, the species was
the availability of food (SCHÄFFER unpublished). In
absent if the average date of mowing of hay and silage
Germany and Poland Corncrakes prey mainly on
was earlier than late July. In the Kostroma region of
insects in the length range 5 - 12 mm (SCHÄFFER 1997).
Russia radio-tagged M Corncrakes were shown by
These are large insects relative to the average size
GRABOVSKY (1993) to inhabit hay meadows and also
available. The density of large insects decreases when
tall grass and herbage at the edges of forest.
mowing of grassland is abandoned (SCHÄFFER 1997).
In Germany singing M select tall ground vege-
tation with some robust or woody stems present. The
4.3. Mating system
distribution of singing M is strongly related to Salix
bushes and the edges of reed beds (Phragmites aus- The mating system of most rails is monogamy with
tralis) where the stems of the previous year’s growth biparental care. Corncrakes are sequentially polyga-
provide cover (SCHÄFFER & MÜNCH 1993). Neighbou- mous. M advertise for mates and probably defend
ring areas with shorter vegetation are used relatively territory by a loud disyllabic song, which is given
early in the season as feeding habitats during daylight occasionally by day and almost continuously at night
(SCHÄFFER & MÜNCH 1993). from tall vegetation (SCHÄFFER 1995a; 1997). W arrive
Singing M Corncrakes have been recorded in on the breeding grounds only slightly later than M
arable crops, especially cereals, in several parts of and are attracted to singing M. A W associates closely
the range. Although arable farmland is not usually with a M for several days (SCHÄFFER & MÜNCH 1993;
considered an important habitat, cereal crops can pro- TYLER & GREEN 1996; SCHÄFFER 1997). During this
vide valuable cover for Corncrakes in regions where period the M sings only infrequently (SCHÄFFER 1995a,
they are not harvested until after the hay crop is cut 1997; TYLER & GREEN 1996). The pair-bond breaks
(CADBURY 1980). In England in the 19th Century, during egg-laying and the M then resumes singing.
when Corncrakes were widespread, they were fre- In some cases M move a considerable distance to a
quently recorded in cereal crops after hay meadows new singing area (TYLER & GREEN 1996; SCHÄFFER
had been mowed (JEFFRIES 1875). In the north-east 1997). W may lay a second clutch later in the summer
Netherlands up to about 200 singing M Corncrakes and may also move before doing so. Hence both M
have been recorded in arable crops in May and June and W are likely to change partners between breeding
(VOSLAMBER 1989), though it is not known whether attempts. Incubation and care of the chicks is done
this is a self-sustaining population. only by the W (TYLER 1996; SCHÄFFER 1997).
124 R. E. GREEN et al.: Populations, ecology and threats to the Corncrake in Europe

4.4. Arrival on the breeding grounds rent years, clumps of singing Corncrakes are found
at different locations even if habitat characteristics
Several days may pass between the arrival of the birds
appear identical, (c) radio-tagged M often associate
in the breeding area and the beginning of singing
together (within 200 m) and displace one another
activity (observations in Russia). During this period,
from singing places. The number of singing M within
the M can be stimulated to call by tape recorded
one clump may vary from about three to ten, though
songs. W arrive soon after the M and pairing occurs
isolated M and regular spacing patterns are also found.
as described above.
Social attraction of singing M may enhance their
Except where detailed studies are made it would
ability to attract W, as in lek-forming species.
usually be assumed that the beginning of singing re-
presents the arrival of M. This assumption would 4.7. Breeding biology
usually lead to errors of only a few days in the esti-
The nest is on the ground in dense vegetation and is
mation of the time of arrival. The time of the main
constructed from dead stems and leaves collected
arrival of Corncrakes was reported on questionnaires
from nearby. Studies of egg-laying by radio-tagged
for 28 states. Where a range was given, the date of
W show that they usually lay one egg per day, but that
peak arrival was taken to lie in the middle of the
eggs can also be laid at shorter intervals (e. g. 12
range. The mean date of the main arrival was 21 May
eggs in 8 days: TYLER 1996; SCHÄFFER 1997). In Scot-
(range 30 April - 15 June). There were only weak
land the first nests of the season of most W were usual-
and non-significant correlations between the date of
ly placed in stands of tall herbs or marsh vegetation
the main arrival and latitude (rS = 0.183) and lon-
(STOWE & HUDSON 1991), but in Ireland, where grass
gitude (rS = 0.183).
growth begins earlier, many first nests were in mea-
4.5. Song dows (TYLER 1996). The average clutch size is about
10 eggs. In Scotland nests of radio-tagged W that had
The song of the M Corncrakes is very loud and cha-
mated with a known radio-tagged M were within 200
racteristic. M usually call almost continuously for
m of the male’s singing place (TYLER & GREEN 1996).
about 4 hours during the night and less frequently at
Incubation is by the W only and begins with the
other times (HUDSON et al. 1990; GRABOVSKY 1993).
completion of the clutch (TYLER & GREEN 1996; SCHÄF-
In Scotland radio-tagged M without W called for about
FER 1997). In Scotland incubation began on first nests
90 % of the time during the night, but those accom-
from about 20 May to 12 June (TYLER 1996). Incu-
panied by W called for 10 - 15 % of the time (TYLER
bating W only left the nest briefly by day and walked
& GREEN 1996). In Germany and Poland paired M
around foraging within a few metres of the nest before
reduce their continuous calling performance at night
returning. Observations in Scotland indicated that,
during egg-laying and sing occasionally by day
apart from the destruction of nests by mowing, nest
(SCHÄFFER 1995a; 1997). DELOV (1995) also suggested
success was remarkably high - 93 % of nests survived
that in Bulgaria M switched from singing at night to
from laying to hatching (TYLER 1996). In Scotland
singing in the early morning during egg-laying. The
incubation of the first nest took about 18 days.
average number of songs per night from 6 birds recor-
The chicks leave the nest soon after hatching and
ded over 9 nights between 23.15 and 03.15 hours in
are fed by the W bill-to-bill. Foraging of broods is by
Germany was 21,207 (range 17,206 - 24,275) (FANG-
day within 100 - 200 m of the nest. In Scotland some
RATH 1994, FANGRATH & SCHÄFFER 1994; SCHÄFFER
whole broods were lost, amounting to 10 - 20 % bet-
1995a). The time of year at which most Corncrakes
ween hatching and independence (TYLER 1996). Par-
stop singing was reported on the questionnaires for
tial losses of chicks also occurred, almost all within
24 states. Where a range was given, the date was
the first 5 - 6 days after hatching. In Scotland an
taken to lie in the middle of the range. The mean
average of 41 % of first brood chicks remained alive
date of ceasing to sing was 8 July (range 12 June -
at independence in broods in which at least one chick
19 July).
survived (TYLER 1996). At two weeks of age the dis-
4.6. Population density and dispersion tinctive calls of young birds and the W can be heard
so giving reliable evidence of breeding (SCHÄFFER
Singing M often are not dispersed evenly within an
1994).
area but, at least in the beginning of the breeding
In Scotland first brood chicks were abandoned
season, are concentrated at a few points (GLUTZ VON
by radio-tagged W at about 12 days old (TYLER 1996).
BLOTZHEIM et al. 1973; SCHÄFFER 1995a). There are
They then lived independently and became capable
two potential reasons for this: suitable habitats are
of flight at about 35 days old. All radio-tagged W
found only in a few locations or social attraction bet-
which reared their first brood before mid July then
ween singing M. The evidence that social attraction
associated immediately with a singing M and pro-
is important is as follows. (a) Singing M often form
duced a second clutch of eggs (TYLER 1996). In north-
clumps within a homogeneous habitat, (b) in diffe-
east Poland the production of two clutches by the
VOGELWELT 118: 117 – 134 (1997) 125

same W was also observed when mowing


of hay meadows was delayed so that mea-
dow habitat which was normally remo-
ved by mowing remained available in late
summer (SCHÄFFER 1995b; SCHÄFFER
1997). In Poland M have been observed
to associate with W accompanying broods
of chicks near to independence (SCHÄF-
FER 1995b). It has been shown by radio-
tracking and capture/ringing that these M
are not related to the chicks of the
female’s first brood (SCHÄFFER 1997).
In Scotland 80 % of second nests were
placed in tall grass in meadows (TYLER
1996). On average incubation began on the
second nest 12 days after the W left the first
brood. The average interval between the
start of incubation on the first and second
nests was 42 days (TYLER 1996).
In Scotland the incubation period of
second nests was slightly shorter (16 - 17
days) than for first nests (TYLER 1996).
Nest success was as high as for first nests,
provided that mowing did not occur. The
loss rate of whole broods was similar for
first and second broods, but partial losses
of chicks were lower for second broods,
67 % of chicks surviving to independence.
W stayed with their second broods longer
(15 - 20 days). Overall, second breeding
attempts were more productive than first
attempts, mainly because of better chick
survival (TYLER 1996).
4.8. Timing of breeding and number
of clutches per female
A typical schedule of egg-laying, incu- Fig. 3: Timing of breeding of Corncrakes. (a) Breeding schedule of a
bation and chick-rearing, based on the typical W Corncrake in the Hebridean Islands of Scotland, based on
studies of radio-tagged W in Scotland, is observations of radio-tagged birds (see text). Each horizontal bar
presented in Figure 3(a) using results from shows, for each of two successive clutches of the same W, the periods
TYLER (1996). The distribution of hat- of laying (solid), incubation (hatched), care of chicks by the W (large
cross-hatch) and flightless independent development of chicks (small
ching dates of the broods of radio-tagged
cross-hatch). The asterisks show the mean date of the main arrival of
W in the Scottish study areas showed two Corncrakes on the breeding grounds throughout Europe and the mean
peaks caused by the hatching of the first date on which most M cease to sing, taken from the questionnaires.
and second broods (Figure 3(b)). In (b) - (d) Hatching dates of Corncrake broods in the Hebridean Islands
Poland SCHÄFFER (1995b) studied timing of Scotland, north-east Poland and western France (see text). - Brut-
of breeding by finding the nests of radio- zeit von Wachtelkönigen. (a) Typischer Brutverlauf eines Weibchens
tagged W, by estimating the age of cap- auf den Hebriden, Schottland, basierend auf Beobachtungen teleme-
tured chicks and by assessing the develop- trierter Vögel (siehe Text). Die horizontalen Balken zeigen für jede
ment of embryos in nests destroyed during der beiden aufeinanderfolgenden Bruten desselben Weibchens die
mowing of hay. This study took place in Legeperiode (schwarz), die Bebrütung (schraffiert), Führen von Jung-
vögeln durch das Weibchen (kreuzschraffiert) sowie Entwicklung flug-
areas where some mowing was delayed
unfähiger Jungvögel (eng kreuzschraffiert). Die Sternchen zeigen den
so that nesting habitat was available durchschnittlichen Ankunftszeitpunkt von Wachtelkönigen in den Brut-
throughout the summer. In France SALA- gebieten in ganz Europa und das mittlere Ende der Rufperiode der
MOLARD et al. (1995) and BROYER (1995) Männchen (Auswertung der Fragebögen). (b) - (d) Schlupfzeitpunkte
observed chicks during hay mowing and von Wachtelkönigbruten auf den Hebriden (Schottland), in Nordost-
estimated their age and hence their hat- polen und Westfrankreich (siehe Text).
126 R. E. GREEN et al.: Populations, ecology and threats to the Corncrake in Europe

ching dates. All of these studies in Scotland, France This suggests that the song period of M Corncrakes
and Poland concur in finding that broods hatch over is adapted to encompass the long period during which
a long period and that there are two peaks in the distri- W produce clutches.
bution of hatching dates (Fig. 3(b) - (d)). In all three In the questionnaire survey, respondents were
areas the two peaks were 40 - 45 days apart, which asked to indicate the half-months in which nests and
agrees with the time between hatching of clutches of flightless chicks were found in their country. Data
individual females. The observations in France sug- were available for nests from 24 states and for chicks
gest that breeding there may be about 15 days earlier from 21 states. Nests were recorded in more than
than in Scotland and Poland. These results suggest half the states from the second half of May to the
that female Corncrakes are able to rear two broods first half of July inclusive (Table 3).
of chicks per year in these three widely separated In 12 states the period in which nests were found
areas. However, in the study area in Poland it was spanned two months or more. Chicks were recorded
only possible to produce two broods when hay mo- in more than half of the states between the second
wing was delayed beyond the normal date. half of June and the second half of July, with many
The average times of year at which M begin and states also reporting flightless chicks in August. The
cease singing, derived from the questionnaire respon- long duration of the period in which nests and chicks
ses, are also shown in Figure 3a. It can be seen that were reported could be partly attributable to variation
the main period of singing coincides with the time between states in the time at which breeding started.
between the beginning of egg-laying on the first However, the length of the period with nests and
clutch and the end of egg-laying of the second clutch. chicks within many of the states suggests that the

Tab. 3: Timing of breeding of Corncrakes in 26 range states as indicated by responses to a questionnaire. Occurrence
of nests with eggs and flightless chicks in each half-month is denoted by x, or by + where respondents indicated that
only a few examples were known. The table has been summarised by counting, for each half-month, the number of
states in which nests or chicks were recorded. – Brutzeitraum von Wachtelkönigen in 26 europäischen Staaten (Ergeb-
nisse nach Fragebögen). Vorkommen eines Geleges oder flugunfähiger Jungvögel in einer entsprechenden Monatshälf-
te werden mit x wiedergegeben, oder mit +, wenn nur wenige Beispiele bekannt waren. Die letzte Zeile gibt für jede
Monatshälfte die Anzahl der Staaten an, in denen Nester oder Jungvögel gefunden/gesehen wurden.

nest with eggs flightless chicks


May June July August May June July August
range state 1 2 1 2 1 2 1 2 1 2 1 2 1 2 1 2
Belarus x x x x x x x x x x
Belgium no data x
Bulgaria x x x x x x x x x
Czech Republic x x x x no data
Denmark x x x x x x
Estonia x x x x x x
France x x x x x x x x x
Germany x x x
Great Britain x x x x x x + + x x x x x +
Ireland x x x x x x + + x x x x x +
Italy x x x x x
Latvia x x x no data
Liechtenstein no data x
Lithuania x x x x x no data
Luxembourg x x
Moldova x x
Netherlands x x x x x x x
Poland x x x x x x x x x
Romania x x x x x no data
Russia x x x x x
Slovakia x x x x x x
Slovenia x x x
Spain x x x x
Sweden x x x x x x +
Switzerland x x x x x x x no data
Ukraine + +
total 5 14 19 16 17 9 3 0 0 3 7 15 14 12 10 5
VOGELWELT 118: 117 – 134 (1997) 127

production of two clutches by individual W within In parts of the Netherlands, Germany and Poland
one breeding season is a widespread phenomenon. mowing of hay in May or June results in the removal
In many countries hay mowing removes much of the of such a large part of the Corncrake’s habitat that
tall vegetation suitable for Corncrakes before a se- birds which settle in these areas must be forced to
cond clutch could be produced, but by delaying move considerable distances after mowing (VAN DEN
mowing experimentally in Poland, SCHÄFFER (1995b) BERGH 1991).
showed that W could produce a second clutch in July
in areas where nests are usually only observed in May 5. Threats
and June.
5.1. Principal threats on the breeding grounds
4.9. Moult
Questionnaire respondents were asked to give a rating
Both M and W begin a simultaneous moult of remiges from 0 to 3 to each of seven named threats and to
and rectrices in July and August after breeding. add any others that they thought were important. All
Within a few days they loose all their primaries and of the scores were added together and the value for
secondaries (SCHÄFFER 1997, TYLER unpublished). As each threat was then expressed as a percentage of
a result they are unable to fly. Radio-tagging in Po- the total. Within each state the threats which received
land, Scotland and Ireland indicates that the birds the highest score (usually 3) were noted and the num-
walk only short distances during this period and re- ber of states giving a threat its highest score was cal-
main within the habitats used for breeding. culated.
Averaging scores over the 30 states with usable
4.10. Migration and wintering
data, mechanization of mowing was regarded as the
Migration to the wintering habitats starts in August greatest threat to Corncrake populations, followed
and continues until October. Several weeks before by earlier mowing dates, loss of hay meadows, loss
leaving the M stop singing and evidence of the pre- of wetlands, abandonment of agriculture, increased
sence of Corncrakes is then difficult to obtain. Hence numbers of predators, hunting and changes to grass-
there is very little information about the precise lands caused by fertilizers (Table 4). Assessing the
timing of the departure from the breeding areas. Pre- threats in terms of the number of states giving it the
sent knowledge about migration and wintering is maximum score gave similar results. Rarely reported
reviewed by STOWE & GREEN (1997, this issue). threats not shown in Table 4 were fire (1 state) and
afforestation (1 state). Afforestation can presumably
4.11. Dispersal and return rates
regarded as a form of loss of hay meadows and/or
Knowledge of dispersal and annual survival of Corn- wetlands.
crakes is rudimentary. There are no reliable estimates
of first-year or adult sur-
vival. This is because the Tab. 4: Rating of threats to Corncrake populations by questionnaire respondents in 30
extent of dispersal has European states. Each threat was scored from 0 (no importance) to 3 (very important).
only been partly quan- Threat scores for each state were summed and each score then expressed as a percentage
tified. The evidence of the total. The table shows the mean and range of the percentages and the number of
available suggests that states giving the highest or equal highest rating to each threat. – Einteilung der Gefähr-
movements of more than dungsursachen von Wachtelkönigen in 30 europäischen Ländern in eine Skala von 0
(unwichtig) bis 3 (sehr wichtig) (Ergebnisse nach Fragebögen). Für jede angeführte
10 km and up to about 50 Gefährdungsursache wurden die Werte der einzelnen Länder aufsummiert und anteilig
km between natal and als Prozentwerte der Gesamtheit angegeben. Die Tabelle gibt die Mittelwerte und den
breeding site or succes- minimalen und maximalen Prozentwert für eine Gefährdungsursache bzw. die Anzahl
sive breeding sites are not der Staaten mit dem höchsten oder gleich hohem Skalenwert an.
infrequent (ALNAS 1974;
VAN DEN BERGH 1991). Re- threat % score number of
turn rates of ringed birds mean minimum maximum states giving
to small study areas were top rank
low, but in the absence of loss of wetlands 16 0 43 9
detailed information on loss of hay-meadows 17 0 40 10
movements it is difficult mechanisation of mowing 26 0 66 21
to assess whether this was earlier mowing 22 0 42 16
because of low survival or reseeding of meadows 1 0 14 0
many movements (ALNAS abandonment of farming 9 0 30 2
1974; SWANN 1986; VAN increase of predators 5 0 33 0
excessive hunting 2 0 18 0
DEN BERGH 1991; FOX
collisions with fences and cables 1 0 9 0
1993; GRABOVSKY 1993).
128 R. E. GREEN et al.: Populations, ecology and threats to the Corncrake in Europe

5.2. Effects of mowing on Corncrake breeding rarely reported killed by mowing. In Russia in the
success and survival 1970s an average of 2 - 3 nests per km2 and 7 - 30
birds per km2 (mainly chicks) were reported as being
The questionnaire survey identified the timing and
destroyed during mechanized mowing of silage from
method of mowing of hay and silage as threatening
the outside of the field inwards (MANUSH 1974).
Corncrake populations in many range states. Effects
The mobility of Corncrake chicks increases with
of mowing on nest success and the survival of chicks
age. The distance moved between successive loca-
and adults has been recognised previously as a pro-
tions less than two hours apart of radio-tagged chicks
bable cause of Corncrake population declines (NORRIS
was less than 50 m for chicks less than 1 week old,
1947; VON HAARTMANN 1958). In 1938 NORRIS (1945)
up to about 100 m for chicks 1 week to 3 weeks old
obtained information on the timing of the Corncrake
and up to 400 m for older chicks (SCHÄFFER & WEI-
decline in different regions of Britain and Ireland
ßER 1996). When disturbed by mowing, only recently
and (NORRIS 1947) on the proportion of hay which
hatched chicks were observed to move so slowly that
was cut using mowing machines, which were mainly
there was a risk of their being overtaken by the side-
horse-drawn, rather than by scythe. It was found that
ways progress of the mower (TYLER 1996). Where
rapid declines in Corncrake numbers were never
mowing was proceeding from the periphery of the
known to precede mechanisation of mowing and that
meadow towards the centre, chicks could sometimes
differences between regions in the timing of the Corn-
be seen at the edge of the uncut block, at the opposite
crake decline were correlated with differences in the
side to that being mowed at the time. They sometimes
time at which most mowing became mechanised.
broke cover and ran to concealment in standing vege-
Mowing machines pose a direct threat to Corn-
tation or under a cut swathe, but more often turned
crakes for two reasons.
back into the standing grass. Where a small patch or
(1) Mechanised mowing can result in large areas of
narrow strip of hay was being mowed, or a meadow
hay or silage being mowed in one session. Mowing
was being mowed from the centre outwards, these
usually proceeds from the outside of the field inwards.
movements away from the mower would be expected
Fully-winged adults and juveniles may fly out of the
to result in most individuals leaving the danger area
grass when close to the mower, but flightless chicks
without breaking cover.
which do not break cover are often killed. Chicks
In Scotland and Ireland about 40 % of the chicks
which attempt to escape by crossing the open, already
observed during mowing from the periphery of the
mowed area of the field may be taken by predators
meadow inwards were killed, or would have been
such as gulls Larus spp. and White Storks Ciconia
killed if they had not been captured by the observers
ciconia.
(TYLER 1996). Observations of mowing in France in-
(2) Because it is more rapid, mechanised mowing
dicated that 86 % of chicks were killed by mowing
allows the whole hay or silage harvest to be comple-
from the outside of the field inwards (BROYER 1996).
ted earlier in the summer. Hence, fields which would
The overall risk of mowing to chicks would be expec-
have been mowed in late summer if mowing was done
ted to be higher, because some individuals would be
by hand are harvested earlier, within the breeding
exposed to several mowing episodes during their 35
period of the Corncrake. Even improvements in the
day flightless period and because the presence of ob-
power and speed of machinery (not the change from
servers probably prevented some predation by gulls.
horse to tractor) in northern England between the
1950s and 1970s led to a 50 % decline in the duration
5.3. Effects of mowing on Corncrake population
of the time required to harvest the hay crop (SMITH &
trends
JONES 1991). In some areas the production of silage
rather than hay is associated with earlier mowing, In addition to studies of the effects of hay and silage
usually as part of a process of intensification to mowing on nest success and chick survival there have
produce higher yields. Earlier mowing results in a also been investigations of the relationship between
larger proportion of Corncrake nests and chicks being the timing of mowing and trends in the population of
at risk (TYLER 1996). adult Corncrakes. GREEN & WILLIAMS (1994) found a
Observations during mowing and tracking of radio- correlation between the mean date of mowing in dif-
tagged adults and chicks during real or simulated hay ferent regions of Britain and Ireland and the trend in
mowing have been used to assess the impact of mecha- numbers of singing M Corncrakes counted between
nised mowing on the survival of nests and chicks 1988 and 1991. Corncrake populations in regions
(TYLER 1996; SCHÄFFER & WEIßER 1996). Virtually all where the average date of mowing was earlier than
nests in mowed meadows were destroyed, but incu- late July were undergoing rapid declines, whereas
bating W were only rarely found killed. Adults and those in regions with later mowing were stable or
chicks tended to run away from the tractor and mower declining more slowly. It can be estimated from Figure
into uncut grass. Although adults become flightless 2 of GREEN & WILLIAMS (1994) that an average mowing
during the moult in July and August they are very date in mid August would be required for a stable
VOGELWELT 118: 117 – 134 (1997) 129

Corncrake population. BROYER (1995) adop- Tab. 5: Rating of mowing methods used in areas with Corncrakes by
ted a similar approach to an analysis of questionnaire respondents in 30 European states. Each method was
Corncrake population trends in France. He scored from 0 (no importance) to 3 (very important). Scores for each
found that Corncrake populations had de- state were summed and each score then expressed as a percentage of
the total. The table shows the mean and range of the percentages. –
clined markedly between 1982 and 1992
Einteilung der Mahdmethoden in Gebieten mit Wachtelkönigvorkom-
in regions where more than 50 % of hay men in 30 europäischen Ländern (Ergebnisse nach Fragebögen).
had been mowed by 15 July, but populations Jeder Methode wurde ein Wert zwischen 0 (unwichtig) und 3 (sehr
had declined more slowly or had increased wichtig) zugeordnet. Die Skalenwerte der Länder wurden aufsum-
in regions with later mowing. Given that miert und als anteilige Prozentzahl der Gesamtheit für jede Mahd-
the timing of the Corncrake breeding season methode angegeben. Die Tabelle gibt den Mittelwert der Prozent-
is earlier in France than in Britain and Ire- zahlen und deren Minima bzw. Maxima wieder.
land (Figure 3), these two studies are in
close agreement that late mowing of hay is % score
associated with stable or increasing Corn- mowing method mean minimum maximum
crake populations. scythe and other hand mowing 12 0 60
horse-drawn mowing machine 3 0 18
5.4. Proportion of Corncrakes in hay tractor-drawn mowing machine 50 20 75
and silage meadows tractor-drawn forage harvester 36 0 75
Given that factors which relate to hay and
silage meadows ranked as the top three among per- significantly correlated with the percentage score for
ceived threats, it was considered relevant to examine mowing by hand and horse-drawn machines (rS =
data of the importance of mowed grassland to Corn- 0.677, P < 0.01). This correlation may indicate a ten-
crakes. The scores given to Corncrake habitats by dency for high density Corncrake populations to have
questionnaire respondents were summed for each persisted longer in states with incompletely mecha-
state and expressed as a percentage of the total (Table nised agriculture. These states are predominantly
2). On average, the percentage scores given to the those which recently had communist political sys-
three categories of hay and silage meadows were 62 tems. This correlation might illustrate cause and
% of the total of all scores (range 17 - 100 %) indi- effect, but this is uncertain because of the confoun-
cating that the management of hay and silage mea- ding effect of geography.
dows could affect an important part of the Corncrake As noted in the section on Population, states in the
population in most states. east of Europe have higher Corncrake density indices
than those in the west. The proportion of mowing by
5.5. Mowing methods used in areas with
hand or horse-power is also significantly correlated
Corncrakes
with longitude (rS = 0.647, P < 0.01), so the observed
Mechanisation of mowing has occurred in most parts correlation between Corncrake density and mowing
of the Corncrake’s European breeding range. Respon- methods might be spurious, resulting from an effect
dents were asked to give a score from 0 to 3 to the of climate or some other factor which varies geographi-
importance of four methods used to cut hay or silage cally. However, GREEN & RAYMENT (1996) found that
(hand scythe, horse-drawn mower, tractor-drawn the population density of Corncrakes was correlated
mower and forage/silage harvester). They could also with agricultural intensity even when longitude was
identify and give a score to any other method. One taken into account by partial correlation.
respondent identified a cutting method other than the
5.6. Proportion of mowing occurring during the
four listed above: mowing steep slopes with a moto-
Corncrake breeding season
rized mower pushed by hand. This was included with
hand scythe mowing. All of the scores for a state Questionnaire respondents were asked to give a score
were added together and the value for each method from 0 to 3 according to the amount of cutting of
was then expressed as a percentage of the total. hay or silage within 8 half-months from May to Au-
In most states mowing with tractor-drawn mowers gust and for cutting after the end of August. All of
and forage/silage harvesters was the most common the scores for a state were added together and the
form of harvesting in areas where Corncrakes occur value for each time period was then expressed as a
(Table 5). However, mowing or cutting of hay by percentage of the total. A cumulative percentage was
hand or with hand operated or horse-drawn machines then calculated which was regarded as a crude index
was an important method in some states. of the percentage of the total area of hay and silage
There was a tendency for mowing by hand or with that had been cut by the end of each half-month. In
horses to be more prevalent in areas with large Corn- some countries the same fields may be cut more than
crake populations relative to their surface area (Figure once in one year, but this was disregarded in the ana-
4). The index of Corncrake density (see above) was lysis.
130 R. E. GREEN et al.: Populations, ecology and threats to the Corncrake in Europe

Fig. 4: The most recent estimate of numbers of singing M


Corncrakes in relation to the land surface area of European
range states. The size of circles is related to the proportion
of mowing recorded as being carried out by hand or horse-
drawn mower in areas with Corncrakes recorded. The
smallest dots represent states with no mowing by hand or
horse-power and the largest states with 60 % mowing by
these methods in areas with Corncrakes. – Neuste Schät-
zungen von Wachtelkönigzahlen in Abhängigkeit von der
Flächengröße europäischer Länder. Die Größe der Krei-
se gibt den Prozentsatz von Flächen an, die mit Hand oder Fig. 5: Comparison of the timing of breeding by Corn-
pferdegezogenen Mähwerken gemäht werden. Die klein- crakes and that of mowing of hay and silage. (a) The num-
ber of states in which Corncrake nests were reported in
sten Kreise zeigen Länder, in denen nicht mit Hand oder
the questionnaire in each half-month. (b) The number of
Pferden gemäht wird. Länder, in denen über 60 % der
Flächen mit den genannten Methoden bewirtschaftet wer- states in which flightless Corncrake chicks were reported
in the questionnaire in each half-month. (c) A score (curve)
den, sind als die größten Kreise dargestellt.
representing the mean cumulative percentage of hay and
silage mowed by the end of successive half-months in areas
Most mowing of hay and silage in areas where with Corncrakes in 30 range states. Vertical lines show
Corncrakes occur was reported to take place in June the range of values of the cumulative percentage. - Ver-
and July, but there was considerable variation among gleich des Brutzeitpunktes von Wachtelkönigen und des
states (Figure 5). In some states most mowing was in Mahdzeitpunktes von Heu und Silage (basierend auf Um-
May and June, while in some others most was in late frageergebnissen). (a) Anzahl von Ländern, in denen Wach-
July and August. Figure 5 also shows the timing of telkönignester in einzelnen Monatshälften nachgewiesen
Corncrake reproduction from the questionnaire results wurden. (b) Anzahl von Ländern, in denen flugunfähige
on the timing of breeding shown in Table 3. Com- Jungvögel in den einzelnen Monatshälften nachgewiesen
parison of Figure 5 with Figure 3 shows that there is wurden. (c) Verlauf der Mahd von Heu und Silage am Ende
der jeweiligen Monatshälften (aufsummiert) in 30 Län-
good agreement between the breeding schedule of a
dern des Verbreitungsgebietes. Vertikale Linien zeigen die
typical W Corncrake in Scotland and the timing of ob- Spannweite der Werte des aufsummierten Prozentwertes.
servations of nests and chicks from the questionnaires.
There is a striking degree of overlap between the
mowing season and the Corncrake breeding season.
In many states much mowing of hay and silage occurs mowing occurs sufficiently early that there is little
before the first clutch has hatched or while the chicks possibility of a second brood being successfully fled-
of the first brood are small. Mowing earlier than late ged in hay or silage and in some even the production
June will destroy the first nests. Early mowing also of the first brood is severely threatened.
removes tall vegetation habitat. Mowed fields are un- Early mowing would also be expected to affect
likely to grow tall enough after mowing to allow the the survival of adults and chicks and the ability of W
production of replacement nests in the same field. In to produce replacement nests by its effect on the area
any case, mowed fields are often either grazed after of suitable tall vegetation. However, direct estimates
mowing or mowed again. In most range states of these effects are not available.
VOGELWELT 118: 117 – 134 (1997) 131

Wachtelköniglebensraum im
Murnauer Moos, Bayern.
Aus Naturschutzgründen
oder bedingt durch hohen
Wasserstand werden Wiesen
(meist Streuwiesen) erst spät
im Jahr gemäht. In Verbin-
dung mit Altschilfstreifen
(hohe Vegetationsdeckung
an den Rufplätzen bei An-
kunft der Vögel im Frühjahr)
finden sich hierdurch ideale
Voraussetzungen für erfolg-
reiche Wachtelkönigbruten.
Foto: N. SCHÄFFER

Wachtelköniglebensraum
im Biebrzatal in Nordost-
polen. Nach Aufgabe der
Bewirtschaftung (früher
Streuwiesenmahd per Hand)
"verfilzen" die Flächen zu-
nächst durch abgestorbenes
Pflanzenmaterial. Mittel-
und langfristig verbuschen
und bewalden die meisten
Flächen. Der Lebensraum
des Wachtelkönigs geht
hierdurch verloren.
Foto: N. SCHÄFFER.

Wachtelkönigweibchen mit
Jungvogel.
Foto: A. LIMBRUNNER.
132 R. E. GREEN et al.: Populations, ecology and threats to the Corncrake in Europe

5.7. Loss of suitable habitat Scotland and Ireland which were occupied by Corn-
crakes in 1978 but not in 1988 had less tall marsh
Loss of wetland and hay meadow habitats were also
vegetation (Iris pseudacorus, Phalaris arundinacea
rated as important threats to Corncrake populations
and Phragmites australis) and hay meadows than sites
in many range states. Wetlands, in particular the edges
which were occupied by Corncrakes in both 1978 and
of marshes, provide important habitats for Corncrakes
1988. GREEN & STOWE (1993) also found that the rate
and are thought to have good food supplies and vege-
of change of the Corncrake population in different re-
tation with an open structure which allows Corncra-
gions of Scotland and Ireland was correlated with chan-
kes to pass through it easily. These habitats provide
ges in the quantity of tall vegetation suitable for Corn-
refuges for Corncrakes when drier grasslands are un-
crakes. Corncrake populations tended to decline in
suitable, for example, because the grass is too short
regions where tall marsh vegetation and hay meadows
during the spring or after mowing in late summer.
decreased and short pasture grazed by sheep increased.
Wetland margins are also likely to be grazed or
mowed late in the summer, if they are managed at 5.8. Predators
all, so the risk of damage to nests and chicks is low.
Mortality of adult birds in Scotland and Ireland has
Wetlands are threatened by drainage and by flood
been estimated by studies of radio-tagged birds. The
prevention schemes on rivers.
main proximate cause of death was predation by
Hay and silage meadows are also important habi-
mammals (otter Lutra lutra, mink Mustela vison and
tats for Corncrakes. These habitats are threatened by
domestic/feral cat Felis catus). Deaths and injuries
conversion to arable cropland, forestry plantations
to untagged Corncrakes caused by collisions with fen-
and other uses. An example is the conversion of hay-
ces, overhead wires and road traffic were observed,
meadows in France to maize Zea mais fields and
but no deaths of tagged birds occurred. Total mortality
poplar Populus ssp. plantations. Intensification of
rates varied substantially between areas and this
management of mowed grasslands is also a threat,
variation did not appear to explain variations in popu-
both because of the increased risk of damage to Corn-
lation trends. In fact adult mortality was higher in an
crakes by mowing (which is discussed above), and
increasing population than in one which was decrea-
because of other changes that are less well under-
sing (TYLER, unpublished data). There were indica-
stood. Some questionnaire respondents believed that
tions that the killing of adult birds by predators was
increased fertilizer application and reseeding cause
most likely to occur when they were located in small
changes in the plant species composition and physical
patches of tall cover which were easy for predators
structure of grasslands used for mowing which reduce
to search, or where the Corncrake was observed and
their suitability for Corncrakes. The mechanisms
became vulnerable when moving between patches.
underlying these effects are believed to be reductions
Hence increasing the area and continuity of suitable
in the abundance of important invertebrate prey and
tall cover at times of year when it is scarce might
increases in the stem density of the grassland which
reduce losses of adults to predators.
make it difficult for Corncrakes to penetrate.
In some states agricultural abandonment is a threat
to habitats which are at present suitable for Corncrakes.
6. Conclusion
Changes in socioeconomic factors may also be impor- Although it remains a widespread species with a large
tant where traditional management of vegetation is world population, concern for the future persistence
necessary to maintain the habitat in good condition of the Corncrake is justified because of population
for Corncrakes. Low-intensity grazing and hay-making declines in many parts of its range. Hay meadows,
may create favourable conditions for Corncrakes at wet pastures and the fringes of marshes are important
wetland margins so reductions in numbers of farmers habitats which are likely to be subject to more drai-
may be harmful in some areas. Reduction in cattle nage and agricultural intensification in future. Loss
farming has led in some areas to the abandonment of of these habitats is likely to cause further declines in
agriculture on dry grasslands which were previously Corncrake numbers. Changes in the management of
grazed or mowed. Such changes may produce a tem- agricultural grasslands, such as mechanisation of hay
porary increase in tall vegetation suitable for Corn- and silage mowing and increases in the power and
crakes, but the eventual development of a dense grass speed of mowing machines are also likely to have a
ground layer and scrub is likely to cause the disappea- negative effect on Corncrake populations by causing
rance of Corncrakes. In other areas, especially in Scot- increased destruction of nests and chicks. Conser-
land and parts of Ireland, cattle farming and hay vation measures should aim to maintain the extent
production has been replaced by sheep grazing which of suitable habitat and enhance breeding success by
has reduced the area of tall vegetation habitats suitable influencing the timing and method of mowing of hay
for Corncrakes. STOWE et al. (1993) found that sites in and silage.
VOGELWELT 118: 117 – 134 (1997) 133

Acknowledgements: We are extremely grateful to the ques- KURESOO, A. KÜRTHY, P. LINA, V. MALETIC, A. MIKITYUK, T.
tionnaire respondents and participants in the European Corn- MIKUSKA, I. MIROWSKI, A. MISCHENKO, T. MOKWA, A. MOSA-
crake Species Action Plan Workshop in Gdansk, Poland, LOV, W. MÜLLER, A. MUNTEANU, D. MUNTEANU, I. ØIEN, E.
October 1994 and especially to J. BROYER, R. BUDRYS, P. OSIECK, R. OTTVALL, F. PERCO, A. PETRYSHYN, J. PETTERSSON,
BURGER, G. C. CABAS, C. CASEY, P. CEMPULIK, T. CONZEMIUS, J. PRIEDNIEKS, J. PYKAL, J. RADOVIC, M. REMISIEWICZ, P. RYE-
J. COVENEY, N. CROCKFORD, M. CZYZAK, V. DELOV, M. DEMKO, LANDT, P. SACKL, E. SÉRUSIAUX, A. SIKORA, I. SKILSKY, K.
J. DEWYSPELAERE, J. DIXON, M. DVORAK, J. ELTS, I. FARRONATO, STANDRING, M. STRAZDS, A. TISHECHKIN, P. TOUT, P. TRON-
M. FLADE, A. FOLVIK, C. GACHE, B. GODERT, I. GORBAN, M. TELJ, L. VAN DEN BERGH, F. VASSEN, Y. VERGELES, Z. WALICZ-
GRELL, B. HEREDIA, M. HEATH, J. HORA, R. HORVATH, J. HUY- KY, G. WILLI, J. WILSON, J. WINKELMAN, M. ZIELINSKI, K.
SECOM, P. IANKOV, W. KANIA, P. KANUCH, D. KARASKA, K.-H. ZUB and M. ZURBA. M. AVERY, T. STOWE and G. WILLIAMS
KOLB, O. KEISS, J. KORNAN, P. KOSKIMIES, T. KULAKOWSKI, A. made helpful criticisms of a draft.

7. Zusammenfassung
Green, R. E., G. Rocamora & N. Schäffer 1997: Bestand, Ökologie und Gefährdung des Wachtelkönigs Crex
crex in Europa. Vogelwelt 118: 117 – 134.
In den letzten Jahren und Jahrzehnten war in fast allen können zwei Gelege je Brutsaison hervorbringen, wobei
Ländern seines Verbreitungsgebietes ein Bestandsrück- sich die Brutzeit von Mai bis August erstreckt. Durch
gang des Wachtelkönigs festzustellen. Dabei wird ein ein- Mahd während der Brutsaison werden alljährlich ein gro-
facher Zusammenhang augenfällig: In den Ländern Mittel- ßer Teil der Nester zerstört und viele junge Wachtelkönige
ost- und Osteuropas, deren Landwirtschaft gekennzeich- getötet. Früher Mahdzeitpunkt, schnelle Mähmaschinen,
net ist durch geringe Intensität oder geringe Effektivität, einheitliche Mahdtermine, größere Bewirtschaftungsein-
sind die Gesamtzahlen von Wachtelkönigen wie auch die heiten sowie die Mahd von außen nach innen brachten
Bestandsdichten wesentlich höher als in Westeuropa. Die den Wachtelkönig auf die Liste global gefährdeter Arten.
vorliegende Arbeit gibt einen Überblick über den derzei-
tigen Wissensstand zu seiner Ökologie. Wichtige Lebens-
raumkriterien sind hohe Vegetationsdichte und geringer Rhys E. Green, Royal Society for the Protection of
Raumwiderstand. Hierdurch können Wachtelkönige ver- Birds, 17 Regent Terrace, Edinburgh EH7 5BN, Scot-
steckt vor Prädatoren lange Strecken durch die Vegetati- land, UK.
on laufen. Dauergrünland und Feuchtgebiete sind die wich-
tigsten Lebensraumtypen des Wachtelkönigs. In den mei-
Gerard Rocamora, Ligue pour la Protection des Oise-
sten Ländern besiedeln Wachtelkönige Dauergrünland, das aux/BirdLife International, BP 263, La Corderie Roya-
zur Heu- oder Silagegewinnung genutzt wird. Das Nest le 17305 Rochefort, France.
wird am Boden angelegt und die Jungen in Flächen mit Norbert Schäffer, Landesbund für Vogelschutz in Bay-
hoher Vegetationsdichte geführt. Wachtelkönigweibchen ern e. V., Postfach, D-91161 Hilpoltstein, Germany.

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