Timing and duration of primary molt in Northern

Hemisphere skuas and jaegers

Authors: van Bemmelen, Rob S. A., Clarke, Rohan H., Pyle, Peter, and
Camphuysen, Kees (C. J.)
Source: The Auk, 135(4) : 1043-1054
Published By: American Ornithological Society
URL: https://doi.org/10.1642/AUK-17-232.1

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 Volume 135, 2018, pp. 1043–1054
DOI: 10.1642/AUK-17-232.1

RESEARCH ARTICLE

Timing and duration of primary molt in Northern Hemisphere skuas and

jaegers
Rob S. A. van Bemmelen,1,2* Rohan H. Clarke,3 Peter Pyle,4 and Kees (C. J.) Camphuysen5
1
Wageningen Marine Research, Wageningen University and Research, IJmuiden, The Netherlands
2
Resource Ecology Group, Wageningen University and Research, Wageningen, The Netherlands
3
School of Biological Sciences, Monash University, Victoria, Australia
4
Institute for Bird Populations, Point Reyes Station, California, USA
5
Royal Netherlands Institute for Sea Research (NIOZ), Texel, and Utrecht University, The Netherlands
* Corresponding author: rvanbemmelen@gmail.com
Submitted December 6, 2017; Accepted June 15, 2018; Published August 29, 2018

ABSTRACT
We compared the primary molt of the 4 species of skuas and jaegers (Stercorariidae) that breed in the Northern
Hemisphere: Long-tailed Jaeger (Stercorarius longicaudus), Parasitic Jaeger (S. parasiticus), Pomarine Jaeger (S.
pomarinus), and Great Skua (S. skua). We analyzed primary molt data of 1,573 individuals of multiple age classes,
mostly collected from photographs taken at sea but also from museum specimens and beached individuals. Whereas
molt duration generally increased with species’ size, molt duration in Parasitic and Pomarine jaegers was surprisingly
similar given their size difference. Larger species started primary molt earlier and showed more overlap with
postbreeding migration, such that there was complete overlap in Great Skua but no overlap in Long-tailed Jaeger.
Within jaeger species, the first primary molt cycle took longer than later molt cycles. We suggest that, unlike birds in
their first primary molt cycle, birds in their second or subsequent primary molt cycles are time-constrained to complete
primary molt before the onset of prebreeding long-distance migration. By contrast, molt duration did not differ
between age classes of Great Skuas. Adult Great Skuas may have overcome the time constraint by completely
overlapping molt and postbreeding migration. Molt-migration overlap is generally rare in birds but may be feasible for
Great Skuas given their shorter migration distance and low migration speed.
Keywords: annual cycles, molt-migration overlap, primary molt, Stercorariidae

Tiempo y duración de la muda primaria en especies de Stercorariidae del Hemisferio Norte

RESUMEN
Comparamos la muda primaria en cuatro especies de Stercorariidae que crı́an en el Hemisferio Norte: Stercorarius
longicaudus, S. parasiticus y S. pomarinus, y Skua skua. Analizamos datos de la muda primaria de 1573 individuos de
múltiples clases de edad, colectados mayormente a partir de fotografı́as tomadas en el mar pero también de
especı́menes de museo e individuos varados. Mientras que la duración de la muda generalmente aumentó con el
tamaño de la especie, la duración de la muda de S. parasiticus y S. pomarinus fue sorprendentemente similar dadas sus
diferencias de tamaño. Se encontró que las especies más grandes comenzaron la muda primaria más temprano y
mostraron más superposición con la migración posterior a la reproducción, de tal manera que hubo una superposición
completa en Skua skua pero no hubo superposición en S. longicaudus. Dentro de las especies de Stercorarius, el primer
ciclo de muda primaria tomó más tiempo que los ciclos de muda posteriores. Sugerimos que, a diferencia de las aves
en su primer ciclo de muda primaria, las aves en su segundo ciclo o en ciclos subsecuentes de muda primaria están
limitadas por el tiempo para completar la muda primaria antes del inicio de la migración de larga distancia previa a la
reproducción. En contraste, la duración de la muda no difirió entre las clases de edad en S. skua. Los adultos de S. skua
pueden haber superado la restricción de tiempo superponiendo completamente la muda y la migración posterior a la
reproducción. La superposición entre muda y migración es por lo general rara en las aves pero puede darse en S. skua
debido a su corta distancia y a su baja velocidad de migración.
Palabras clave: ciclos anuales, muda primaria, Stercorariidae, superposición entre muda y migración

INTRODUCTION events in the annual cycle of all birds. Several strategies

have evolved to fit molt into a bird’s annual cycle, generally
Primary molt, the replacement of the large remiges avoiding temporal overlap with other demanding activities
attached to the ‘‘hand’’ of the wing, is one of the major such as breeding and migration (Hedenström 2006). Non-

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 1044 Primary molt in Stercorariidae R. S. A. van Bemmelen, R. H. Clarke, P. Pyle, and C. J. Camphuysen

overlap may be achieved by adjusting the timing of molt with age and breeding success (Furness 1987). Individ-
(e.g., postponing molt until after the most intense period of uals initially stay out at sea during the boreal summer of
chick care, or after migration) or by shortening the their second calendar year and may return to the
duration of molt (usually by growing more feathers breeding grounds in their third or fourth calendar years,
simultaneously; Rohwer and Rohwer 2013). Speeding up where they arrive later in the season and remain for a
of molt can lead to decreased flight performance shorter duration than adults (de Korte 1984, Furness
(Hedenström and Sunada 1999) or a lower quality of 1987). Recruitment to the breeding population occurs
feathers (Dawson et al. 2000, Serra 2001). after 3 yr in the Long-tailed Jaeger (de Korte 1985) and
Most variation in molt duration between species is may take even longer in the other species (Furness 1987).
explained by body size (larger species taking longer to The contrasts in body size and migration distance
molt; Rohwer et al. 2009), whereas molt duration and between species, and the contrast in the degree of
timing may vary between and within species according to migratory and breeding behavior between ages within
breeding and migration behavior (Lindström et al. 1993a, species, allow us to explore the effect of these factors on
Serra et al. 1999, Pyle 2008, Dietz et al. 2013). Despite this, the timing and duration of primary molt. Our expecta-
few studies have quantified differences in molt duration tions were that (1) larger species will require more time
and timing in detail for closely related species or for to renew all primaries than smaller species; (2) by
different groups within species. In seabirds, this is partly starting earlier, larger species will require larger overlap
due to a lack of primary molt data for many species (Bridge with migration; (3) within a species, populations
2006). Studying molt of flight feathers in seabirds is wintering farther south will have a shorter molt duration;
compromised by the fact that the birds disperse over vast and (4) molt duration will decrease with age as time
expanses of ocean where collection of adequate sample constraints imposed by migration and breeding set in.
sizes can be difficult. These challenges have been overcome
in recent years, now that digital photography has been METHODS
shown to facilitate molt scoring of flying birds (Keijl 2011,
Vieira et al. 2016) and increasing numbers of photos taken Primary molt scores were obtained from several sources.
during offshore surveys and opportunistic cruises are Most were scored from photographs, which were either
shared online. sourced from the Internet (in particular from online
Stercorariidae, the family of skuas and jaegers, sighting portals), supplied by photographers, or taken by
comprises 7 species. We compared the primary molt of the authors. A dataset obtained in the same way and
all 4 species of skuas and jaegers that breed in the published previously by Newell et al. (2013) was also
Northern Hemisphere: Long-tailed Jaeger (Stercorarius included, but all molt scores were reassessed by the first
longicaudus), Parasitic Jaeger (S. parasiticus), Pomarine author to ascertain uniformity. Only the best visible wing
Jaeger (S. pomarinus), and Great Skua (S. skua). Parasitic was scored. Photographic records were filtered for
and Long-tailed jaegers are sister species, and evidence duplicates, which could be identified by date and location,
has been accumulating that the Pomarine Jaeger is a by molt status, and by individual plumage characters such
sister group of the Great Skua (Janssen and Mundy as the length and shape of the central tail feathers, the
2017). These species range in body mass from ~300 g in amount of barring on the underparts, and the color of the
Long-tailed, ~400 g in Parasitic, and ~700 g in Pomarine head cap. Additional primary molt data were obtained
jaegers to ~1,400 g in the Great Skua (Cramp and from tideline corpses on Dutch beaches (Nederlandse
Simmons 1983). Breeding in the (sub)Arctic and Stookolieslachtoffer Onderzoek database), as well as from
spending the nonbreeding period on the oceans, all museum specimens from three sources in California, USA:
skuas are migratory, but the distances covered vary the California Academy of Sciences, San Francisco (CAS);
considerably between species and individuals. Nonbreed- Museum of Vertebrate Zoology, Berkeley (MVZ); and
ing areas vary between 508S and 208N in the smallest Western Foundation of Vertebrate Zoology, Camarillo
species, the Long-tailed Jaeger (Gilg et al. 2013, van (WFVZ). Finally, we included published molt scores of
Bemmelen et al. 2017), and between 108N and 508N in museum specimens from Stresemann and Stresemann
the largest species, the Great Skua (Furness 1987, (1966), Lambert (1980), Melville (1983), and de Korte
Magnusdottir et al. 2012). Migration of Parasitic and (1985). Individual jaegers were assigned to 4 age classes
Pomarine jaegers is less well studied. Pomarine Jaegers (first-cycle, second-cycle, third-cycle, and adult) repre-
are thought to winter mainly around 0–208N (albeit with senting primary molt cycles, based on plumage characters
regular occurrence to ~368S off Australia and else- and coloration of tarsi as outlined by Howell (2007) and
where), whereas Parasitic Jaegers mainly migrate to 10– Pyle (2008). In Great Skuas, we only distinguished between
508S (Olsen and Larsson 1997, Menkhorst et al. 2017). first-cycle birds and older birds, because no reliable ageing
The timing and extent of migration are thought to vary characters are established to distinguish second- and third-

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 R. S. A. van Bemmelen, R. H. Clarke, P. Pyle, and C. J. Camphuysen Primary molt in Stercorariidae 1045

TABLE 1. Mean relative mass per primary (thus summing to 1), sample size, and the total mass of all primaries for 4 species in the
family Stercorariidae as measured from dead specimens. Primaries are numbered distally, p1 being the innermost.
Mean
Species p1 p2 p3 p4 p5 p6 p7 p8 p9 p10 mass (g) n

Long-tailed Jaeger 0.034 0.041 0.050 0.063 0.080 0.098 0.121 0.143 0.168 0.202 2.17 1
Parasitic Jaeger 0.035 0.043 0.055 0.068 0.084 0.103 0.121 0.140 0.163 0.189 3.21 4
Pomarine Jaeger 0.039 0.046 0.056 0.070 0.085 0.103 0.119 0.141 0.159 0.181 5.08 4
Great Skua 0.045 0.053 0.063 0.077 0.091 0.106 0.117 0.133 0.151 0.163 7.91 9

cycle birds from older individuals. Note that the ‘‘adult’’ Skua sample to be representative for that species’ wing
class will include an unknown proportion of individuals shape.
that did not (yet) recruit to the breeding population, or Mean start date and duration of molt were estimated
that skipped or failed breeding in any given year. by modeling PFMG values against day since July 1 (a date
In all 4 species, primary molt starts at the innermost that falls well outside the primary molt period in most
primary and completes at the outermost primary (Pyle age classes). This was done using likelihood models,
2008). We assigned molt scores of 0 (old), 1–4 (growing referred to as UZ models (Underhill and Zucchini 1988)
and in pin or 10–33%, 33–67%, or 67–99% grown, and implemented in the package ‘‘molt’’ 2.0.0 in R 3.4.0
respectively), or 5 (new and fully grown) (Ashmole (Erni et al. 2013, R Core Team 2017). In contrast to other
1962). Feather molt scores of 1 and 2 are usually methods, UZ models have specifically been designed to
indistinguishable in photographs (for examples, see analyze molt data (e.g., dealing with the heteroscedas-
Appendix Figure 4), because these feathers are clearly ticity of molt data and biases inherent to regression
dropped but the new feathers are not yet visible beyond the methods; Summers et al. 1983, Underhill and Zucchini
primary coverts. In the case of a single invisible feather, 1988). Only individuals in active primary molt were
this was scored as 1, whereas 2 invisible feathers were considered (type 3 data: Underhill and Zucchini 1988).
scored as 1 (outermost) and 2 (innermost). Fully grown R2 values were obtained from linear models of observed
primaries collected from birds found dead were cleaned, vs. fitted values. Age was included in the models as a
dried, and weighed to the nearest 0.001 g. Using the mean covariate for molt duration, starting date, and standard
relative mass for each of the 10 primary feathers (Table 1), deviation in starting date. Given the smaller sample sizes
primary scores were converted into proportion of feather of second- and third-cycle jaegers (Table 2), we
mass grown (PFMG), a molt index that increases combined these into a single age class (hereafter
approximately linearly over time (Summers et al. 1980). ‘‘second/third-cycle’’). The adult age class was taken as
The mean relative mass of each primary reflects the shape the baseline level, because this class held the largest
of the wing, which shows little variation between sample size. Optimization procedures for UZ models are
individuals or age classes within species, but large sensitive to starting values (Erni et al. 2013). Therefore,
differences between species (Dawson 2005). Indeed, the convergence problems for the model of Parasitic Jaeger
inclusion of several juveniles in our sample (1 Parasitic were solved by providing starting values, based on
Jaeger, 2 Pomarine Jaegers, and 4 Great Skuas) did not separate models for each molt cycle, and a standard
affect our results. Moreover, because the relative mass of deviation parameter of 5 days. To estimate the mean
each primary (numbered p1 to p10) varied, on average, by feather growth rate of adults, we divided the mean total
only 2.5–4.9% within Parasitic Jaegers, Pomarine Jaegers, feather mass by the estimated molt duration.
and Great Skuas, we considered our single Long-tailed Considering the effect of nonbreeding-area latitude on
molt timing and duration shown in other species (Serra et
TABLE 2. Number of individuals in active primary molt per age al. 1999, 2006, Underhill 2003, Pyle 2008), we also explored
class for 4 species in the family Stercorariidae, as used in the UZ this by adding the covariate ‘‘latitudinal zone’’ for onset and
models. No reliable plumage characteristics are known for duration of molt and comparing their Akaike’s Information
ageing second/third-cycle Great Skuas, which are consequently Criterion (AIC) values against models without covariates.
included in the adult class.
We defined 3 latitudinal zones to investigate the potential
Long-tailed Parasitic Pomarine Great effect of (nonbreeding-area) latitude on molt parameters:
Jaeger Jaeger Jaeger Skua
north of 358N in the Atlantic (mainly from the North Sea
Adult 209 175 540 221 or U.S. east coast) or north of 408N in the Pacific (mainly
Second/third-cycle 37 53 157 – off the U.S. west coast), from these latitudes south to the
First-cycle 32 23 88 38
equator (mainly California Current and Canary Current),
Total 278 251 785 259
and south of the equator (mainly Australia and the

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 1046 Primary molt in Stercorariidae R. S. A. van Bemmelen, R. H. Clarke, P. Pyle, and C. J. Camphuysen

Benguela Current). These latitudinal zones reflect areas RESULTS

where jaegers normally do not spend winter, areas that can
be regarded as regular ‘‘northern’’ nonbreeding areas and Sample Size
regular ‘‘southern’’ nonbreeding areas, respectively (Olsen A sample of 1,573 individuals of Northern Hemisphere–
and Larsson 1997). breeding Stercorariidae was included in the UZ models,
The critical assumption of UZ models of a constant rate comprising 278 Long-tailed Jaegers, 251 Parasitic Jaegers,
of change in molt index can be violated by birds that 785 Pomarine Jaegers, and 259 Great Skuas. For all species,
suspend molt (Underhill and Zucchini 1988, Erni et al. the largest sample size was represented by the adult age
2013). In molt suspension, molt is temporarily halted, to be class (69–85%; Table 2). Most records were obtained from
resumed later. It results when a newer, full-grown feather photographs (98% of those for Long-tailed, 83% for
falls distal to a substantially older feather, with no feathers Parasitic, and 86% for Pomarine jaegers; 94% for Great
missing (Pyle 2008, Pyle and Reid 2016; Appendix Figure Skua); the remainder originated from museum specimens,
5). We expect primary molt suspension to mainly take except for some beached Great Skuas (3%).
place in birds that start molt when still well north of their Because UZ models are sensitive to outliers (Erni et al.
nonbreeding area, suspend molt during migration, and 2013), some records that showed combinations of date and
resume after arrival at the main nonbreeding area. We molt score not compatible with any molt cycle were
therefore explored the effect of exclusion of individuals removed from the dataset (prior to the above tabulation).
that (after being ‘‘sampled’’) may have suspended primary Excluded records were as follows: (1) a first-cycle Long-
molt—those that were molting when (still) in the northern tailed Jaeger on August 9, 2015, off the Scilly Islands, UK,
latitudinal zone—on model parameters. with growing p8–9 and old p10; (2) an adult Long-tailed
In order to relate timing and duration of primary molt Jaeger sitting in poor condition on the beach in Mauritania
to migration timing, we derived individual-level timing of on May 7, 2013, with p9 growing and p10 old; (3) a Long-
departure from the breeding area, arrival at and tailed Jaeger, presumably an adult, reported on September
departure from the nonbreeding area, and southbound 23, 1981, in New Zealand, with p7 growing and p8–10 old;
overall migration speed from light-based geolocation (4) Long-tailed Jaegers picked up moribund in New
data for Parasitic and Long-tailed jaegers. Parasitic Zealand during the 1982 ENSO event appeared to have
Jaegers were captured at Slettnes, Norway (71.08198N, delayed molt (Melville 1983), as has been shown in other
28.21028E; R. van Bemmelen et al. personal observation), seabirds (Guerra et al. 1988, Howell and Corben 2000); (5)
with data spanning 3 yr (2014–2017), and Long-tailed an adult Pomarine Jaeger collected on August 1, 1967, at
Jaegers at Ammarnäs, Sweden (66.00488N, 16.18428E; Barrow, Alaska, with p1–4 full-grown, p5 score 4, p6 score
van Bemmelen et al. 2017), with data spanning 4 yr 2, and p7–10 old (CAS 68455); and (6) a Pomarine Jaeger
(2011–2015). Both datasets were analyzed using the same on March 31, 2007, from New South Wales, Australia,
techniques (for details, see van Bemmelen et al. 2017). showing delayed Staffelmauser (see below).
Because both datasets span multiple years, we assume
that year effects are leveled out. However, mean timing of Staffelmauser
migration for the Swedish Long-tailed Jaegers may be up Five first- to second-cycle Pomarine Jaegers showed
to a few weeks earlier than the overall species’ average, Staffelmauser (Stresemann and Stresemann 1966; Appen-
considering that breeding and migration are timed later dix Table 4), in which the second molt cycle had
at higher latitudes (Conklin et al. 2010), and this Swedish commenced before outer-primary replacement during
population is closer to the southern border of the the first molt was finished. Four of these, all collected in
breeding distribution. The breeding distribution of Monterey Bay, California (CAS 10932, 10965, 10979, and
Parasitic Jaegers extends farther south (Olsen and 11096), were duplicated to be included as both first- and
Larsson 1997), with Slettnes taking a central position in second-cycle birds in the models because their PFMG
the distribution range, and we expect that our sample values were within the range of other first- and second-
therefore corresponds reasonably well with the species’ cycle Pomarine Jaegers. However, a fifth Pomarine Jaeger
overall mean migration timing. For Pomarine Jaeger and showing Staffelmauser, photographed off New South
Great Skua, approximate migration periods and distanc- Wales, Australia, lagged several months behind in both
es were obtained from the literature (Cramp and cycles and was excluded from the UZ models because it
Simmons 1983, Furness 1987, Higgins and Davies 1996, caused errors during model fitting. We did not find
Olsen and Larsson 1997, Magnusdottir et al. 2012) and— examples of Staffelmauser in the other species.
with the exception of geolocator data for Great Skuas
(Magnusdottir et al. 2012)—are not specific to certain Suspended Molt
years or sites. We therefore assume they are generaliza- Several individuals showed suspended molt. These were 3
tions over large areas and multiple years. first-cycle (up to p7 or p8 new) and one adult (up to p9

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 R. S. A. van Bemmelen, R. H. Clarke, P. Pyle, and C. J. Camphuysen Primary molt in Stercorariidae 1047

FIGURE 1. Progress of molt in adult (black), second/third-cycle (blue), and first-cycle (red) jaegers and Great Skuas, expressed as the
proportion of feather mass grown (PFMG). Shaded areas represent 95% confidence intervals. Box plots above plots of Long-tailed
and Parasitic jaegers show (from left to right) start of southbound migration, arrival at nonbreeding grounds, and departure from
nonbreeding grounds as inferred from geolocator data. Approximate timing of migration based on the literature is shown by
horizontal lines above the plots of Pomarine Jaeger and Great Skua.

new) Long-tailed Jaegers, one adult Parasitic Jaeger (only birds, and this timing difference increased with species’
p1 new), 3 second-cycle (up to p2, p7, or p8 new) and 2 body weight. In the 3 jaeger species, molt took
adult Pomarine Jaegers (up to p1 and p2 new), and a single considerably longer in first-cycle birds than in adults
adult Great Skua (up to p9 new). We may have missed (longer by 50, 43, and 54 days); while in Great Skuas, molt
suspended molt in other records, given that the detection duration was similar between those two age classes (4
depends on the quality of the picture and the difference in days longer in first-cycle birds). Duration of molt in
age of old and new feathers. second/third-cycle birds was more or less similar to that
of adults. The model outputs suggest that second/third-
Model Results cycle Long-tailed Jaegers start at the same time as adults
All 4 species display clear temporal patterns in molt but have a 35-day-longer molt duration, whereas second/
timing. Among adults, Great Skuas showed the longest third-cycle Parasitic and Pomarine jaegers have a molt
molt duration and Long-tailed Jaegers the shortest duration that is shorter than that of adults by 17 and 20
(Figures 1 and 2; Table 3). However, the molt durations days, respectively. All models had very high R2 values.
of Pomarine and Parasitic jaegers were virtually the same, Adults were the only age class with sufficient data from
despite the distinctly larger size of the former. Projected separate latitudinal zones to test for the effect of latitude.
end dates for adults were February 13 for Long-tailed The distribution of records of adults among the 3
Jaegers, March 18 for Parasitic Jaegers, February 23 for latitudinal zones was as follows, from north to south: 0,
Pomarine Jaegers, and February 18 for Great Skuas. The 65, and 144 Long-tailed Jaegers; 13, 43, and 119 Parasitic
molt of first-cycle birds was well separated in time from Jaegers; 22, 275, and 243 Pomarine Jaegers; and 213, 8, and
that of the adults, as well as from second/third-cycle 0 Great Skuas. For all 3 jaeger species, adding ‘‘latitudinal

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 1048 Primary molt in Stercorariidae R. S. A. van Bemmelen, R. H. Clarke, P. Pyle, and C. J. Camphuysen

and Long-tailed jaegers traveled, on average, 273 km day1

(n ¼ 29) and 293 km day1, respectively (n ¼ 60).
According to the literature, Great Skuas breeding in
Scotland complete a migration of 3,000–3,500 km to Iberia
or northwest Africa (based on light-level geolocator data in
Magnusdottir et al. 2012) in ~2 mo (Cramp and Simmons
1983, Furness 1987). This suggests they travel approxi-
mately 50–60 km day1.

DISCUSSION

Consistent with our expectations, we found that molt of

primaries in the larger Great Skua generally took longer,
started earlier, and had a larger temporal overlap with
migration than in the 3 jaeger species. Within jaeger
species, differences in molt duration between age classes
suggest time constraints to finish molt before northward
migration. While these findings are largely in line with
FIGURE 2. Parameter estimates for starting date (x-axis) and previous publications (Olsen and Larsson 1997, Wiley and
molt duration (y-axis) for adult, second/third-cycle, and first- Lee 1998, Howell 2007, 2010, Pyle 2008, Pyle and Reid
cycle jaegers and Great Skuas. Note that the upper SE of first- 2016), we provide more precise estimates of molt onset
cycle Parasitic Jaeger extends beyond the bounds of the figure.
and duration and a uniform data analysis for all 4 species
and identifiable age classes, allowing interspecific and
zone’’ did not result in large decreases of AIC values (DAIC intraspecific comparisons.
. 2) compared to UZ models without covariates. Molt durations of adult Parasitic and Pomarine jaegers
We found no apparent effects related to Parasitic and were surprisingly similar, given that they differ significantly
Pomarine jaegers that may have suspended molt after in body weight (335–470 vs. 542–917 g, respectively;
being sampled. Removal of samples from the northern Cramp and Simmons 1983). A longer molt duration in
latitudinal zone had only a small effect on molt duration Parasitic Jaegers, approximating that of Pomarine Jaegers,
(,0.4 days) and on mean starting date (,2 days) in both is difficult to explain. Molt suspension may lead UZ
species. We could not perform this analysis for Long-tailed models to overestimate the duration of active molt, but
Jaegers, given the lack of molting adults in the northern models of both species appeared to be robust to the
latitudinal zone, or for Great Skuas, which largely remain inclusion of individuals that may have suspended molt
within the latitudinal zone. In addition, we could not after being sampled. Another possible modeling issue
repeat this analysis for first-cycle or second/third-cycle would be a poor model fit due to covariance of molt
datasets, because of smaller sample size. parameters, which is a known problem for type 3 UZ
models (Underhill and Zucchini 1988, Erni et al. 2013).
Migration Although this was the case in models for both Parasitic (r ¼
Timings of departure from the breeding areas, and arrival 0.897) and Pomarine (r ¼ 0.912) jaegers, we can see no
at and departure from the nonbreeding areas, are reported reason why this would affect the models for the 2 species
for Parasitic and Long-tailed jaegers in Figure 1. Parasitic differently.

TABLE 3. Model estimates for mean start date and duration (days) of primary molt in 4 species in the family Stercorariidae. In
parentheses, SD values are indicated for the mean start date and SE values for molt duration. For sample size per species and age
class, see Table 2. Parameters are missing for second/third-cycle Great Skuas because no reliable characters are known to distinguish
these age classes from older birds.
First-cycle Second/third-cycle Adult
Species Start date Duration Start date Duration Start date Duration R2

Long-tailed Jaeger January 16 (6 22) 132 (6 41) November 27 (6 22) 117 (6 25) November 23 (6 15) 82 (6 10) 0.905
Parasitic Jaeger February 12 (6 42) 165 (6 76) December 17 (6 42) 105 (6 22) November 16 (6 18) 122 (6 8) 0.889
Pomarine Jaeger February 13 (6 35) 172 (6 27) November 23 (6 35) 98 (6 21) October 29 (6 20) 118 (6 6) 0.871
Great Skua March 21 (6 28) 151 (6 34) – – September 24 (6 27) 147 (6 16) 0.739

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 R. S. A. van Bemmelen, R. H. Clarke, P. Pyle, and C. J. Camphuysen Primary molt in Stercorariidae 1049

Pomarine Jaegers by a relatively low feather growth rate.

This would be remarkable, given that the predominant
strategy for adjusting molt duration in birds is by
modifying molt intensity rather than feather growth rate
(Rohwer and Rohwer 2013). The benefit of decreasing
feather growth rate could be higher feather quality, given
that shortening molt duration has been shown to decrease
feather quality (Dawson et al. 2000, Serra 2001). High
feather quality may be especially important in Parasitic
Jaegers to avoid feather damage when performing frantic
aerial pursuits of terns and other seabirds.
In addition to the effect of body size on molt duration
(Rohwer et al. 2009), time costs of migration may have an
additive effect leading to shorter molts (De la Hera et al.
2009). This may result from ‘‘squeezing in’’ the molt
between breeding and postbreeding migration, but when
molt is postponed to the wintering period—as in our
study species—shorter molts may result from less time
spent at the wintering grounds due to longer migration
durations. Although Great Skua indeed has the shortest
migration distance and the longest molt duration, the
pattern does not hold for jaegers; the Pomarine Jaeger has
the shortest winter period and the Long-tailed Jaeger the
longest, but molt duration is shorter for the latter.
Moreover, we expected any effect of migration distance
on molt duration also to be reflected within species, but
this was not the case; molt duration in all 3 jaegers did
not differ significantly between areas north and south of
the equator.
Previous studies indicated that shorebirds and procel-
lariiform seabirds that migrate to the tropics or the
Southern Hemisphere display longer molt durations than
FIGURE 3. (A) Molt intensity (number of primaries growing observed in related migratory species or populations of the
simultaneously) throughout the molt (upper panel) and (B) same species that remain in the Northern Hemisphere
deduced mean feather growth rate (lower panel) in adults of 4 (Serra et al. 1999, 2006, Underhill 2003, Pyle 2008). Birds
species in the family Stercorariidae. that remain in the Northern Hemisphere appear to be
under greater time constraints to complete molt before the
Alternatively, model results could reflect a true similar- onset of winter conditions, with fewer resources, less
ity in molt duration between Parasitic and Pomarine favorable weather, and reduced daylight in which to forage.
jaegers. Molt duration is determined by both the number Our results did not reveal such a latitudinal difference
of simultaneously growing feathers (molt intensity) and the between (1) subtropical areas north and (2) subtropical to
time required to grow individual feathers (growth rate). temperate areas south of the equator, despite differences
While total feather mass is clearly larger in Pomarine in, for example, day length. Jaegers that remain in northern
Jaegers (Table 1), molt intensity was not different between temperate waters during the nonbreeding period (which is
the 2 species at any time during primary molt (Figure 3), uncommon in Pomarine, rare in Parasitic, and extremely
which suggests that feather growth rate (grams per day) rare in Long-tailed jaegers; Olsen and Larsson 1997) may
could explain the similar molt durations. Given that face time constraints, but our samples from this region
feather growth rate tends to increase with body mass in were too small to robustly estimate the timing and
birds (Rohwer and Rohwer 2013), we expected the duration of primary molt in these waters. Similarly, it
Parasitic Jaeger’s feather growth rate to be intermediate would be interesting to test for a difference in molt
between those of Long-tailed and Pomarine jaegers, but in duration between Great Skuas from the present study that
fact it was surprisingly similar to that of the former and probably wintered off Africa and Iberia and those
lower than that of the latter (Figure 3). Thus, Parasitic wintering in the Grand Banks area near Newfoundland,
Jaegers may arrive at a molt duration similar to that of Canada (Magnusdottir et al. 2012).

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 1050 Primary molt in Stercorariidae R. S. A. van Bemmelen, R. H. Clarke, P. Pyle, and C. J. Camphuysen

Among our study species, larger species started molt while actively molting (Croxall et al. 2005). In albatrosses,
earlier, thereby creating a temporal overlap of primary this may be facilitated by a low molt intensity, thereby
molt and southbound migration. Migratory bird species minimizing the molt gap and negative effects on flight
usually molt before or after migration, or divide molt performance (Prince et al. 1993). Great Skuas, however,
between these periods. In the latter case, molt can be can have substantial molt gaps. Rather, molt-migration
suspended during active migration, which means that no overlap in Great Skuas may be facilitated by a short
wing surface gap is present. In the smallest species in the migration distance and low travel speeds. Indeed, Parasitic
Stercorariidae, the Long-tailed Jaeger, we found neither and Long-tailed jaegers travel ~300 km day1, whereas
adults that suspended molt in the inner primaries nor Great Skuas breeding in Scotland travel at only ~50 km
birds of any age class that started primary molt well north day1. In fact, travel speed in Great Skuas is so low that one
of the nonbreeding areas and could have suspended molt could argue they are more or less stationary most of the
later on. In the Parasitic Jaeger, starting primary molt in time. In addition to travel speeds, high food availability en
northern temperate regions seems an uncommon strat- route may facilitate molt-migration overlap in Great Skuas
egy. Only a few of the molting adults in our data were migrating through productive waters of the North Sea and
from northern temperate areas, and all were photo- Gulf of Biscay. By contrast, while migrating between
graphed in September–October and in early molt; one staging in productive areas, jaegers move fast over
adult from Monterey Bay, California, USA, had suspend- unproductive subtropical and tropical waters from the
ed molt after growing the innermost primary (MVZ central North to the South Atlantic (e.g., van Bemmelen et
101196). In line with this, Olsen and Larsson (1997) state al. 2017), where molt may need to be postponed or
that 5% of Parasitic Jaegers start primary molt in suspended not only because of food limitation. Time
temperate waters. Some of these adults may stay constraints, and the need of waterbird species that remain
throughout the nonbreeding area in northern temperate in the Northern Hemisphere for the nonbreeding season to
waters instead of the usual tropical or southern temperate complete molt more quickly than species that occupy
nonbreeding areas. Pomarine Jaegers start postbreeding more southerly nonbreeding areas, may also affect the
migration later than Parasitic Jaegers and are commonly degree to which jaegers and skuas undergo active primary
encountered in early primary molt off central California. molt during migration (Pyle 2008).
Here, they become much scarcer later in the boreal Although mean starting dates of adults differed by 2 mo,
winter (Briggs et al. 1987), which suggests that birds that Long-tailed Jaegers, Pomarine Jaegers, and Great Skuas
have begun molt move southward later on. Individuals finished within 10 days of each other, whereas Parasitic
may then suspend molt at p1 or p2 during migration, as Jaegers finished 23 days later. These mean end dates, from
indicated by 2 adult specimens from Monterey Bay, late January through early March, are just before the onset
California, USA (MVZ 17765 and 17800). Thus, both of northbound migration as inferred from geolocator data
Parasitic and Pomarine jaegers can suspend molt during and literature. We found no evidence of molt-migration
southbound migration, but whether some individuals overlap in the period January–May, in contrast to
migrate with molt gaps remains unknown. individuals in active molt during southbound migration
By contrast, our results indicate a complete overlap of at northern temperate latitudes. Our interpretation is
migration with primary molt in Great Skuas. Great Skuas therefore that individuals start northbound migrating only
start primary molt at or directly after departure from the after they have finished primary molt. Among migratory
breeding grounds (Cramp and Simmons 1983, Furness birds breeding in the Northern Hemisphere, migration
1987). Data collected at Jan Mayen and included in our speed is usually higher during northbound than during
study indicate that, in pairs with chicks close to fledging, at southbound migration, indicating a high selection pressure
least one of the partners had dropped the inner primary. for timely and fast migration (Nilsson et al. 2013).
Molt then continues during southbound migration. In Conceivably, overlap of molt with northbound (prebreed-
November–December, when adult Great Skuas are virtu- ing) migration entails larger costs than overlap with
ally absent from the North Sea (Camphuysen and Leopold southbound (postbreeding) migration. For example, the
1994, Furness et al. 2006) and have moved farther south elevated energy requirements (Lindström et al. 1993b) and
and west, molt has progressed only halfway. Such an decreased flight performance (Hedenström and Sunada
extensive overlap of molt and migration is usually 1999) of molt may constrain migration speed and lead to
considered rare among migratory birds, but less rare later arrival in the breeding area (Gorney and Yom-Tov
among short-distance migrants and species that feed on 2001), which can affect breeding performance (Harrison et
the wing, such as terns and hirundines (Yuri and Rohwer al. 2011).
1997, Zenatello et al. 2002). It is not clear to what extent The absence of molt-migration overlap during north-
other seabird species molt primaries during migration, but bound migration and the (nearly) coinciding dates of molt
at least some albatross species may cover huge distances completion and departure from the nonbreeding areas

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 R. S. A. van Bemmelen, R. H. Clarke, P. Pyle, and C. J. Camphuysen Primary molt in Stercorariidae 1051

(Figure 1) suggest that molt in adults is timed to finish just the Northern Hemisphere breeding area. However, this
prior to northbound migration. That this reflects a time should also have been reflected in Long-tailed Jaegers:
constraint is suggested by the ~50 days longer molt third-calendar-year birds (thus between the second and
duration and larger variation in starting dates of first-cycle third cycles) of this species are regularly recorded at the
jaegers compared to adults (Pyle 2008, Howell 2010). breeding grounds (de Korte 1984, van Bemmelen 2010).
Unlike adults, first-cycle jaegers are not constrained by The molt parameters presented here can inform us
breeding duties or migration back to the breeding areas. when specific primaries are molted, which is critical
Shorter primary molt duration of adults compared to first- information for study design and data interpretation when
cycle individuals has also been reported in two other sampling biomarkers from primaries. This applies not only
Arctic-breeding, long-distance migrants, the Black-bellied to future studies, but also to reinterpretation of published
Plover (Pluvialis squatarola) and the Red Knot (Calidris results. For example, Furness et al. (2006) reported a
canutus), in which second-calendar-year individuals do gradual change through p1 to p10 in both d15N and d13C
not breed and usually remain on the nonbreeding grounds stable isotopes in Great Skuas and suggested that this
during the boreal summer (Serra et al. 1999, Dietz et al. reflected a gradual change in diet to lower trophic levels.
2013). In contrast, most second-calendar-year Wood However, given that Great Skuas actively molt during
Sandpipers (Tringa glareola) migrate north directly after migration, an alternative explanation is that this merely
an incomplete primary molt on the nonbreeding grounds, reflects the environmental gradients in stable isotope ratios
and this molt takes as long as that of adults (Remisiewicz et over which Great Skuas traverse. Indeed, isoscapes
al. 2010). An alternative or additional reason for a longer published after Furness et al. (2006) show gradients in
molt duration in first-cycle jaegers may be primary molt stable isotope ratios of both elements in near-surface
suspension, which was observed in some first-cycle Long- plankton along the Great Skua’s migration route (Graham
tailed Jaegers (Pyle and Reid 2016) and also may explain et al. 2010), which is expected to be reflected in higher
first-cycle Long-tailed Jaegers being in active primary molt trophic levels, including within the growing feathers of top
in the (late) boreal summer (Wiley and Lee 1998) or even predators such as the Great Skua. Given that the smaller
Pomarine Jaegers showing Staffelmauser. Potentially, first- species also may move during primary molt (van
cycle jaegers might suspend primary molt during a (poorly Bemmelen et al. 2017), we suggest sampling several
known) northbound migration. Their rarity at the breeding primaries to obtain biomarker signals along molting
areas during the boreal summer suggests that they usually locations.
do not migrate all the way to the Arctic (de Korte 1984, The present study is one of only a few that have used
van Bemmelen 2010), and first-cycle birds are apparently digital photographs to obtain avian molt data (Keijl 2011,
absent from the Benguela nonbreeding areas during the Conklin and Battley 2012, Vieira et al. 2016). We gathered
same period (Lambert 1980), which suggests that most stay these largely from the Internet. With the increase in
in tropical or temperate waters (Howell 2010). Finally, popularity of (pelagic) birding, digital photography, and the
first-cycle jaegers may be constrained by lower foraging ever-increasing capacity to share large numbers of
success due to less experience or exclusion from high- observations and images on the Internet, we consider this
quality habitats. However, these options remain speculative approach to have great potential for the study of molt in
until the movements of younger jaegers are uncovered. In other seabird species. There is a clear need for such
contrast to the 3 jaeger species, molt duration and studies, given the increasing use of biomarkers sampled
variation in starting date were virtually equal in first-cycle from flight feathers and other feather tracts, while
and older Great Skuas, despite similar contrasts between adequate molt data remain lacking for many species
age classes in breeding and migration as in the jaegers. (Bridge 2006, 2011).
Adult Great Skuas may have compensated for the time
constraint by a greater overlap of molt with southbound
migration. ACKNOWLEDGMENTS
We also expected a longer molt duration in second/
third-cycle jaegers than in adults, but our results were Thanks to the many photographers who were willing to share
inconclusive. We found a longer duration in second/third- their photos. Special thanks to M. de Boer, J. Davies, B. Flood,
M. Gottschling, S. Howell, A. Richardson, P. Ryan, R.
cycle than in adult Long-tailed Jaegers, and shorter
Stephenson, and A. Williams, who provided large numbers
durations for Parasitic and Pomarine jaegers. Again,
of photos and/or helped with additional requests. Many
covariance between the parameters for duration and onset photographs were found on the Internet, which would not
of molt may have led to underestimated durations for have been possible without the numerous birders sharing
Parasitic and Pomarine jaegers. Alternatively, second/ their data online and the developers of these portals: www.
third-cycle Parasitic and Pomarine jaegers already face waarneming.nl, www.observado.org, www.ebird.org, www.
similar time constraints as adults if they are migrating to birdpix.nl, www.flickr.com, www.birdguides.com, and https://

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 1052 Primary molt in Stercorariidae R. S. A. van Bemmelen, R. H. Clarke, P. Pyle, and C. J. Camphuysen

macaulaylibrary.org. P.P. thanks curators of the California Part two: Arrival, site tenacity and departure. Beaufortia 34:1–
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specimens under their care. Thanks to D. Newell, S. Howell, Part three: Clutch size, laying date and incubation in relation
and D. López-Velasco for exchanging datasets of Great Skua to energy reserves. Beaufortia 35:93–127.
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APPENDIX TABLE 4. Details of Pomarine Jaegers showing Staffelmauser (see text). PFMG (proportion of feather mass grown) values
for the first cycle (the molt wave that has advanced farther toward the outer primaries) are calculated by setting new growing
feathers of the second molt cycle (inner primaries) to 5 (fully grown), whereas those grown or growing in the first cycle are set to 0
(old) to calculate PFMG values for the second cycle.
PFMG

Specimen Date Location Molt score First cycle Second cycle

CAS-10932 August 1, 1907 Monterey Bay, CA, USA 2155555554 0.977 0.011
CAS-10979 September 25, 1907 Monterey Bay, CA, USA 3155555554 0.977 0.030
CAS-11096 August 22, 1907 Monterey Bay, CA, USA 2155555542 0.867 0.011
CAS-10965 August 22, 1907 Monterey Bay, CA, USA 1555555554 0.997 0.005
March 31, 2007 New South Wales, Australia 5515555420 0.701 0.092

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 1054 Primary molt in Stercorariidae R. S. A. van Bemmelen, R. H. Clarke, P. Pyle, and C. J. Camphuysen

APPENDIX FIGURE 4. Adult Pomarine Jaeger in active primary

molt, off Mauritania on November 11, 2016. Primary molt was
scored as 5542100000, resulting in a PFMG value of 0.171. Note
that p4–5 are invisible and scores for these feathers were
inferred. Photo credit: Rob van Bemmelen

APPENDIX FIGURE 5. Second-cycle Pomarine Jaeger that

suspended molt after growing p1–2, off Wollongong, New
South Wales, Australia, October 23, 2010. Photo credit: Raja
Stephenson

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