Bird Study (2005) 52, 166–172

Hatching success in Avocet Recurvirostra avosetta

and Black-winged Stilt Himantopus himantopus

JOSÉ JAVIER CUERVO

Estación Experimental de Zonas Áridas, Consejo Superior de Investigaciones Científicas, Calle General
Segura 1, E-04001 Almería, Spain

Capsule Daily survival rate of nests was influenced by laying date and nesting period (laying versus
incubation) in both species and by colony size in Black-winged Stilts.
Aims To investigate the effect of clutch size, colony size, reproductive phenology and nesting period on
hatching success of Avocets and Black-winged Stilts.
Methods The study was undertaken at two localities in southwestern Spain (Doñana and Odiel Natural
Parks) during 1990 and 1991 breeding seasons. The Mayfield method was used to estimate hatching
success.
Results Both species bred colonially and some colonies included nests of both species. Neither year,
locality, species or clutch size had a significant effect on hatching success. Daily survival rate was lower
for late nests than for early and intermediate, and lower during egg-laying than during incubation in both
species. Hatching success of Black-winged Stilt nests was lower in large colonies (>50 nests) than in
medium-sized ones (11–50 nests).
Conclusion Late clutches within a colony (clutches with the least hatching success) might be replacements
or belong to late breeders, but adults were not individually marked, precluding clarification. The results
suggest a high probability of clutch predation and/or desertion in early stages of the nesting period,
especially shortly after laying the first egg. The fact that hatching success in large colonies was lower
than in medium-sized ones, but only for Black-winged Stilts (not for Avocets), might help to explain the
difference in coloniality of the two species.

Avocets Recurvirostra avosetta and Black-winged Stilts of clutch size, colony size, reproductive phenology and
Himantopus himantopus are the two species of recurvi- nesting period (laying, incubation) on hatching success
rostrids that breed in the western Palearctic (Cramp & and daily survival rate of nests are investigated in
Simmons 1983). The reproductive biology of Avocets the two species. The study of Black-winged Stilts is
has been extensively studied in Germany in recent especially important owing to the dearth of informa-
years (Hötker 1998, 2000, Hötker & Segebade 2000). tion for this species, but the study of Avocets is also
However, the reproductive biology of Black-winged important because previous studies either have been
Stilts is poorly known, although some studies have undertaken in other countries, probably under different
been carried out in Italy (Tinarelli 1990, 1992) and ecological conditions (e.g. Hötker 2000), or results
Spain (Castro 1993, Arroyo 2000, Cuervo 2003, 2004). have been inconsistent (Arroyo 2000).
Studies of hatching success in the Spanish populations
of the two species have focused mainly on the effect
METHODS
of habitat type (Arroyo 2000) and nest location or
characteristics (Arroyo 2000, Cuervo 2004). However,
Study areas
the study of other factors which might affect hatching
success or daily survival rate of nests has either been The study was carried out during the 1990 and 1991
neglected or different patterns reported. Here, the effects breeding seasons (April–June) in two wetlands in
southwestern Spain, the Doñana and Odiel marshes,
Email: [email protected] located approximately 70 km apart. Avocet and Black-

© 2005 British Trust for Ornithology

 Nesting of recurvirostrids 167

winged Stilt nests were surveyed for two years in Veta A nest was considered successful when at least one
la Palma (36°57′N, 6°14′W), Doñana Natural Park, egg hatched. However, nest fate is often difficult to
Isla Mayor, Seville. However, La Herradura (37°14′N, determine, because adult recurvirostrids remove
7°00′W), Odiel Natural Park, Huelva, was surveyed eggshells immediately after hatching (Sordahl 1994)
only in 1991, and the Black-winged Stilt was the only and chicks leave the nest permanently within one or
recurvirostrid species breeding there. The study area in two days, or even within a few hours if they are
Doñana Natural Park was a former dry marsh recently disturbed (Hamilton 1975, Adret 1984). It is also
flooded for shrimping, consisting of large (c. 100 ha) possible that predators carry away the eggs leaving no
shallow (c. 1 m) brackish ponds. Avocets and Black- evidence of predation (Green et al. 1987). A nest was
winged Stilts nested on man-made dykes, small islands, deemed successful if (i) a recently hatched chick was
and pond shores. Vegetation on emerged land was found in the nest or nearby, (ii) at least one egg showed
patches of typical dry marsh vegetation, mainly evidence of imminent hatching (cracked or drilled
scattered glasswort (Arthrocnemum spp.) up to 50 cm eggshell), (iii) date of egg disappearance matched
high. For a description of ecological and climatic expected hatching date (when egg age was known) and
characteristics of the area see García Novo et al. no sign of predation was found. A nest was considered
(1977). The study area in Odiel Natural Park was a to have failed if (i) eggs were partially or totally
brackish marsh, profoundly altered by humans, made submerged, (ii) eggshell or remains of egg content were
up of dykes, canals and shallow, formerly tidal ponds. found, (iii) the nest had been deserted (cold eggs), (iv)
Black-winged Stilts nested on small islands and dykes. egg disappearance was prior to expected hatching date
Vegetation was scarce and typically halophile (adapted (when egg age was known). The fate of all other nests
to soils with very high concentration of salt). For was considered unknown. Only nests with known fate
detailed information on climate, vegetation and were used to assess hatching success.
ecological conditions see Rubio & Figueroa (1983) and
Rubio (1985).
Nesting periods
Daily survival rate was calculated for three different
Data collection
periods: (i) between laying of first and second egg, (ii)
Breeding areas were surveyed every 1–5 days in search between laying of second and last egg, and (iii)
of recurvirostrid nests, and content of all known nests between laying of last egg and hatching of first egg.
determined. Due to the conspicuousness of adults, Time between laying of two consecutive eggs was
scarce vegetation, and limited suitable nesting assessed including only nests found when one egg had
locations, almost all the nests in the study area were been laid, revised every two days at most, and surviving
found. Every nest was identified by means of a small until clutch completion. Length of the second period
numbered peg stuck into the ground, about 50 cm from (between laying of second and last egg) was calculated
the nest. Colony disturbance always lasted for less than separately for each species, taking into account the
one hour, and extreme temperature periods (early number of nests with different clutch sizes and the time
morning, noon) were avoided. No predators were between laying of two consecutive eggs in that parti-
observed to have taken advantage of researcher- cular species. The length of the third period was
induced disturbance to depredate on eggs or chicks. assessed including only nests found during egg-laying,
Groups of nests more than 100 m apart were considered with at least one hatched egg, and revised every three
different colonies, because that was the maximum dis- days at most.
tance birds chased potential aerial predators. Thus, any
nest within 100 m of a colony might benefit from anti-
Statistical analysis
predator behaviour of members of the colony. Every
nest was classified as an early (first 25%), intermediate Hatching success was assessed by the Mayfield method
(middle 50%) or late (last 25%) nest within its colony. (Mayfield 1961, 1975, Klett & Johnson 1982).
Since colony size might have some influence on However, hatching success estimated with the standard
hatching success, all colonies were classified into three method (percentage of nests with at least one egg
groups by number of nests of the two species: small hatching) is also shown to facilitate comparison with
(2–10 nests), medium-sized (11–50 nests), and large other studies. Calculation of the variance of the
colonies (>50 nests). Mayfield estimator was necessary to compare the daily

© 2005 British Trust for Ornithology, Bird Study, 52, 166–172

168 J.J. Cuervo

mortality rate between different colonies, localities or a Early Intermediate Late

years.a Once daily survival rate is assessed, it is possible 100

to estimate mean reproductive success for Avocet and

Successful nests (%)

Black-winged Stilt nests, simply by raising daily sur- 80

vival rate to the power of the number of days. If partial

60
egg loss in successful nests (nests with at least one egg
hatching) is considered, the egg survival rate, i.e. pro-
40
portion of eggs that survived until hatching age, can be
estimated. All statistical tests were two-tailed and the 20
significance level was 0.05.
0
Doñana Doñana Total
RESULTS 1990 1991
A total of 276 Avocet and 274 Black-winged Stilt nests b
were found and surveyed, in 13 colonies and three soli- 100
tary Black-winged Stilt nests. Six colonies were formed
exclusively of Black-winged Stilts, four colonies were

Successful nests (%)

80
mixed-species with Black-winged Stilt predominance,
and three were mixed-species colonies with Avocet 60
predominance. Percentages of nests with different
clutch sizes were 3% (2 eggs), 13% (3 eggs), 83% (4 40
eggs), and 1% (5 eggs) for Avocets and 2% (2 eggs),
19% (3 eggs), and 79% (4 eggs) for Black-winged Stilts. 20

0
Nest fate and phenology Doñana Doñana Odiel Total
1990 1991 1991
Nest fate was unknown for 117 (42%) Avocet and 51 Figure 2. Percentage of successful nests (at least one egg hatched)
(19%) Black-winged Stilt nests and these nests were in Avocets (a) and Black-winged Stilts (b) considering laying date
therefore not included in subsequent analyses. All (early, intermediate, or late) of each clutch within its colony.
Hatching success and laying dates were significantly related for
other nests either succeeded or failed due to a known Black-winged Stilts in Doñana 1990, Black-winged Stilts in Odiel
cause (Fig. 1). Only 59% of Black-winged Stilt nests 1991, and Avocets in 1991 (G-test, in all three cases 31 < n < 114
with known fate succeeded in hatching at least one egg, nests, G > 12.8, df = 2, P < 0.01), and also for all Avocets (n =
138 nests, G = 30.6, df = 2, P < 0.001) and all Black-winged Stilts
while the percentage of successful nests was 64% in (n = 146 nests, G = 23.9, df = 2, P < 0.001). The relationship was
Avocets. Hatching success was lower in late nests (Fig. not significant in the two cases with the smallest samples (Avocets in
1990: n = 24 nests, G = 3.1, df = 2, ns; Black-winged Stilts in
Doñana 1991: n = 22 nests, G = 2.5, df = 2, ns).
100 Avocets Stilts

80 2), except the smallest samples, where the trend was

not significant (Avocets in 1990, and Black-winged
Stilts in Doñana 1991; see Fig. 2 caption).
Nests (%)

60

40
Nest survival and colony size
20 Daily Avocet and Black-winged Stilt nest survival rates
were estimated for the two years and both localities
0 (Table 1). No significant differences between years for
Successful Depredated Flooded Deserted Avocet or Black-winged Stilt nests were found, nor
Figure 1. Percentage of Avocet (n = 159) and Black-winged Stilt were differences between localities for Black-winged
(n = 223) nests that were successful, depredated, flooded or Stilt nests significant (in all three tests, z < 1.6, ns). In
deserted. Percentage of successful nests (G-test, Gadj = 0.8, df = 1,
ns) or causes of reproductive failure (G = 3.0, df = 2, ns) did not the later analyses daily survival rates are pooled from
differ significantly between species. different years and localities for each species. Daily

© 2005 British Trust for Ornithology, Bird Study, 52, 166–172

 Nesting of recurvirostrids 169

Table 1. Daily survival rate for Avocet and Black-winged Stilt nests in different years and localities.

n Nest-days Failed nests Daily survival rate Hatching success Hatching success
(se) (Mayfield method) (standard method)

Avocets 1990 42 542.5 18 0.967 (0.008) 0.428 0.571

1991 121 1605.0 39 0.976 (0.004) 0.541 0.678
Total 163 2162.5 57 0.974 (0.003) 0.514 0.650
Black-winged Stilts Doñana 1990 178 2376.5 74 0.969 (0.004) 0.454 0.584
Doñana 1991 19 232.0 11 0.953 (0.014) 0.299 0.421
Odiel 1991 36 440.5 9 0.980 (0.007) 0.602 0.750
Total 233 3049.0 94 0.969 (0.003) 0.454 0.597

Daily survival rate was estimated for the period between laying of the first egg and first hatching: 25.3 days for Avocets and 25.1 days for
Black-winged Stilts (see the text).

Table 2. Daily survival rate for Avocet and Black-winged Stilt nests in small (2–10 nests), medium-sized (11–50 nests) and large (>50 nests)
colonies.

Colony size n Nest-days Failed nests Daily survival rate Hatching success Hatching success
(se) (Mayfield method) (standard method)

Avocets Medium-sized 28 344.0 12 0.965 (0.010) 0.406 0.571

Large 135 1843.5 45 0.976 (0.004) 0.541 0.667
Black-winged Stilts Small 23 269.5 9 0.967 (0.011) 0.431 0.609
Medium-sized 80 1265.5 27 0.979 (0.004) 0.587 0.663
Large 130 1514.0 58 0.962 (0.005) 0.378 0.554

Three solitary Black-winged Stilt nests were pooled with nests in small colonies. Daily survival rate was estimated for the period between
laying of the first egg and first hatching: 25.3 days for Avocets and 25.1 days for Black-winged Stilts (see the text).

survival rate was not significantly different in mono- and 21.8 days for Black-winged Stilts (n = 39, se =
specific (Black-winged Stilt) colonies, mixed-species ±0.2). Between-species difference in duration of the
colonies with Black-winged Stilt predominance, or third period was not significant (Student’s t-test; t =
mixed-species colonies with Avocet predominance for –0.68, df = 47, ns).
either species (in all four tests, z < 1.6, ns). Avocets Daily survival rate in the three periods mentioned is
only nested in medium-sized and large colonies, and shown in Table 3. Survival was lower in the period with
daily survival rates did not differ between these two only one egg in the nest than in the other two periods
types of colony (z = 1.0, ns; Table 2). Daily survival rate for both species (in all four tests z > 3.45, P < 0.001).
of Black-winged Stilt nests did not differ significantly In Black-winged Stilts, daily survival rate was lower in
between small colonies or solitary nests and medium- the second period (rest of egg-laying) than in the
sized (z = 1.0, ns) or large colonies (z = 0.4, ns). period with complete clutch (z = 2.44, P = 0.015), but
However, survival was higher in medium-sized colonies survival difference between these two periods in
compared to large ones (z = 2.7, P < 0.01; Table 2). Avocets did not reach statistical significance (z = 1.93,
ns). Daily survival rate in the period with complete
clutch was compared between clutches with three and
Nest survival and nesting period
four eggs, and the difference was not significant for
Time between laying of two consecutive eggs was 1.4 either species (in the two tests, z < 1.14, ns).
days (n = 8, se = ±0.1) for Avocets, and 1.2 days (n = Comparisons were not made with two- and five-egg
25, se = ±0.1) for Black-winged Stilts, and this was clutches owing to the scarcity of such nests.
considered the duration of the period between laying of
first and second egg. Length of the second period
Egg survival
(between laying of second and last egg) was 2.4 days for
Avocets and 2.1 days for Black-winged Stilts. The third Nesting success (proportion of nests with at least one
period (between laying of last egg and hatching of first egg hatching) for Avocets was: 0.7111.4 × 0.9512.4 ×
egg) lasted 21.5 days for Avocets (n = 10, se = ±0.5) 0.98221.5 = 0.372; and for Black-winged Stilts: 0.6411.2

© 2005 British Trust for Ornithology, Bird Study, 52, 166–172

170 J.J. Cuervo

Table 3. Daily survival rate for Avocet and Black-winged Stilt nests in three different periods: one egg (between laying of first and second
egg), rest of egg-laying (between laying of second and last egg), and complete clutch (between laying of last egg and hatching of first egg).

Period n Nest-days Failed nests Daily survival rate (se)

Avocets One egg 35 45.0 13 0.711 (0.068)

Rest of egg-laying 47 183.5 9 0.951 (0.016)
Complete clutch 144 1959.0 35 0.982 (0.003)
Black-winged Stilts One egg 68 97.5 35 0.641 (0.049)
Rest of egg-laying 83 146.0 10 0.932 (0.021)
Complete clutch 184 2803.5 48 0.983 (0.002)

See text for period duration.

× 0.9322.1 × 0.98321.8 = 0.348. Daily survival rate of Stilts, but not for Avocets. Hötker (2000) has previ-
Avocet eggs was 0.998, which means 0.951 hatching ously reported for Avocets breeding in northern
success for the whole period with eggs in the nest. Daily Germany that colony size (once nest density had been
survival rate of Black-winged Stilt eggs was 0.999 and controlled) did not significantly affect hatching
hatching success 0.975. Only 336 out of 376 eggs success. Arroyo (2000) found a similar result in
(89%) in successful Avocet nests reaching hatching age southern Spain for Avocets. There is little information
eventually hatched. Only 469 out of 497 eggs (94%) in on the effect of colony size on hatching success in
successful Black-winged Stilt nests reaching hatching Black-winged Stilts. Arroyo (2000) found no signifi-
age eventually hatched. If the probability of nest sur- cant relationships between the two variables, except in
vival is multiplied by probability of egg survival in one year, when the relationship was significant and
successful nests and this value is multiplied by hatching positive, that is, the larger the colony size the better
probability of eggs reaching hatching age, hatching hatching success. This result does not agree with the
probability of just-laid eggs is obtained. This is 0.372 × findings of this study, and should be accepted with
0.951 × 0.894 = 0.316 for Avocets and 0.348 × 0.975 × caution, since it was found in only one year, sample size
0.944 = 0.320 for Black-winged Stilts. was rather small and the relationship was very weak
(Arroyo 2000, p. 195).
Coloniality has both advantages and disadvantages,
DISCUSSION
and their combination determines the degree of
Daily survival rate of nests was not significantly coloniality and characteristics of the colony (for a
different between species, years, localities, or between review see Brown & Brown 2001). Why was hatching
three-egg and four-egg clutches. The only two factors success of Black-winged Stilts lower in large than in
that seemed to affect hatching success to any degree medium-sized colonies? One important advantage of
were laying date and colony size. In general, hatching coloniality in recurvirostrids is defence against aerial
success was the highest in early nests and lowest in late predators, but this function is probably achieved in
nests (Fig. 2). However, hatching success did not differ medium-size colonies, and does not improve in larger
significantly among early, intermediate and late nests colonies (Hötker 2000). Moreover, rats Rattus spp.
in the smallest samples (Avocets in 1990, Black-winged caused almost all egg predation in this study (most
Stilts in Doñana 1991). The negative relationship nests with depredated clutches contained eggshell
between laying date and hatching success is widespread fragments with rat teeth marks), and coloniality is not
among avian species (for a review in seabirds see efficient against terrestrial predators. On the other
Moreno 1998) and different hypotheses (age and hand, breeding in large colonies might entail some
experience, food availability, predation risk, etc.) have costs. With increasing numbers of individuals in the
been proposed to explain it (Moreno 1998). colony, agonistic interactions might also increase,
Determining the specific factors responsible for poor especially owing to nest-site competition and nest
hatching success in late Avocet and Black-winged Stilt defence. Recurvirostrids are not highly territorial, but
nests was beyond the aims of this study. they defend a small area around their nests (Cramp &
The other factor affecting hatching success was Simmons 1983). Excessive aggression might lead to
colony size. Daily nest survival rate was higher in inappropriate nest attendance and even to nest
medium-sized, than in large colonies for Black-winged desertion.

© 2005 British Trust for Ornithology, Bird Study, 52, 166–172

 Nesting of recurvirostrids 171

In this study, Avocets bred only in medium-sized and constructive criticism on a previous version of the manu-
large colonies, while Black-winged Stilts bred in small script. Arrocerías Herba S. A. and Agencia de Medio
ones also, thus showing more breeding flexibility and Ambiente de Huelva kindly allowed research and provided
less coloniality. Moreover, previous studies have shown accommodation in Doñana and Odiel Natural Parks, respec-
tively. The study was supported by a grant from the Spanish
that Avocets are more gregarious than Black-winged
Ministry of Education and Science.
Stilts when breeding, with smaller inter-nest distances
within the colony (Cuervo 2004). The difference in
coloniality of the two species may be because breeding ENDNOTE
in large colonies is detrimental for Black-winged Stilts a. According to Johnson (1979): variance = (exposure –
(hatching success of this species in large colonies was losses) × losses/exposure3. Standard error (se) was the square
lower than in medium-sized ones), but not for Avocets. root of the variance. A z-value (area under the normal curve)
Daily survival rate of nests was lower during egg- shows whether differences in daily survival rate are statisti-
laying than during incubation for both species, and was cally significant (Johnson & Shaffer 1990): z = |s1 – s2|/(v1
especially low immediately after laying of the first egg. + v2). In this formula, s is daily survival rate and v is the
If a nest site is prone to flooding (e.g. located on wind- variance.
ward shores) or easily accessible to predators (e.g. close
to rat dens), egg mortality will probably occur soon
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(MS received 25 September 2003; revised MS accepted 26 May 2004)

© 2005 British Trust for Ornithology, Bird Study, 52, 166–172