General
description:
By late September, the primary moult is in its
last stage, with only the outermost primary P10 still not completely fully
grown in some birds. But many birds show fully grown P10 and therefore
have finished the complete moult, which started in April.
As can be seen in the August section, the partial autumn moult in
2cy michahellis has started and by September this partial moult is
well in progress in Etaples and
Boulogne-sur-Mer,
NW France. The partial moult overlaps with the last stage of the
complete moult.
Most 2cy birds include the inner greater coverts,
central and outer median coverts and most lower lesser
coverts in the partial moult. The upper tertials and more rarely some lesser
coverts may be replaced again as well. The final extend of the partial autumn
moult in 2cy michahellis can be seen in the October
section.
Moult stage and
pace of remiges and rectrices:
The table below represents two primary
moult scores for September 21 and 22 2002 at Etaples, Calais region, NW
France. It's clear that most 2cy michahellis have completed or nearly completed
the moult of the primaries, with only P10 falling slightly short. Throughout the summer months, from June to September, the primary
moult scores in 2cy michahellis remained very predictable, with
very little variation (compared to e.g. 2cy Lesser Black-backed Gulls in
the Netherlands), as can be read from the low standard deviation
scores. By mid September, slightly less than 50% of the birds have the
primary moult completed.
By the end of July, the average primary
moult score is P6 fully grown by 28th July. The most
advanced birds by late July show an obvious short wing, with P10 dropped
and P7 already fully grown. One bird has P8 fully grown. The low SD values
(standard deviation) in the July
tables imply a small range of variation.
By the end of August, 2cy michahellis
show an average increase of one primary in NW France: the average PMS is
7.9 (n = 29) on August 22 and the average PMS is 8.1 (n = 53) on August
25.
By September, this average has increased by about 1 point: an average PMS
of 9.3 on September 22 and an average score of 9.6 (n = 39) on September
29.
These michahellis figures can be
compared to 2cy argenteus. The differences in average PMS between
the two species are limited but apparent. 2cy Larus argentatus l.s.
from the Netherlands are about 1.2 points behind, compared to 2cy michahellis
from NW France. Note however that from September onwards, Scandinavian
argentatus join the local population of argenteus, resulting
in an obvious increase in the range of individual primary scores, as can
be read from the increase in standard deviations from the end of the month
onwards. The majority of 2cy michahellis
have finished moult in the rectrices by the end of August and have
finished moult in the secondaries by the second week of September. By
the end of September, moult in the secondaries may be seen again in just
less than 20% of 2cy michahellis. Birds grow in plain white
tail-feathers or white feathers with limited black vermiculation in the partial
moult. Table three shows an
overview of rectrices moult scores of 22 birds from mid-September. Since
the
tail-feathers are moulted centrifugally, the replaced tail-feathers are normally the inner rectrices R1 and
R2.
Timing and strategy of partial moult
In the complete moult during the summer, 2cy michahellis moult all the
scapulars, wing-coverts, tertials and rectrices. In the July and
August
sections the progress of this complete moult is described. By August, the
complete moult in the wing-coverts is finished with replacement of the last
central greater coverts and finally the feathers in the carpal edge (outer lesser
coverts). As those last lesser coverts are replaced for second
generation feathers, the partial autumn moult starts in 2cy michahellis,
starting again
with the upper tertials and outer median coverts. The
box below enlarges upon the partial moult in autumn in 2cy michahellis.
Partial moult in
2cy Yellow-legged Gull. What exactly does it mean?
Feathers
wear, bleach, break and deteriorate by use and climatologically
exposure. Birds have to replace the feathers to renew their complete
plumage, normally in cycles and in most northern hemisphere species
at least once a year. Second calendar year Yellow-legged Gull is no
exception, and the complete plumage is replaced in summer. The
complete moult starts with shedding the innermost primary P1 and
finishes with a fully grown outer primary P10. In autumn, a second
moult wave is started, including body-feathers for most: some of the
wing-coverts and tertials are moulted to a next generation feathers.
Some individuals include tail-feathers in this moult.
These two images clearly show what is meant by the partial moult and
illustrates the notations used in our surveys. Wing-coverts are
labeled from the body towards the outer wing and the tertials are
numbered from
upper feather downwards. Best clue for recognition of fvresh
feathers are the grey
bases and neat white fringes, compared to the bleached
adjacent feathers.
In these images, the following feathers are included in the partial moult: tertials
#1-#2 (tt1, tt2) are new, the lower tertials are still old. In the greater coverts, #2 (gc2) has been replaced and
#5 is missing. In the median coverts #1-#2 (mc1, mc2) have been
replaced, #4 is missing, #6 has been replaced and all feathers from
#8 outwards are new. The inner lower lesser coverts (illc) #1, #4-#7
are new, #8-9 are old, #10 is
missing. In the upper lesser coverts (lc) only one innermost feather
has been renewed. The exact generation of the new wing-coverts and
especially the scapulars is very difficult to ascertain. 2cy michahellis
has an extensive partial post-juvenile moult, where many
wing-coverts are replaced. In complete moult in summer (2cy), these
wing-coverts are replaced for third generation feathers. In the
partial autumn moult in 2cy, as described in this box, some
wing-coverts are renewed again, to fourth generation.
Scapulars are moulted 'continuous' as it seems,
although at a lower pace in winter. By May, 2cy michahellis
may show already 95% third generation scapulars and a vast majority
is moulting again in June and (still?) in August. Some scapulars may
thus apply to third generation feathers (especially in the lowest
row), but likewise some apply to fourth
(maybe fifth) generation scapulars. In the picture, the abraded
lower scapulars with brown centres (ls) are probably old third
generation feathers, the fresh plain grey feathers in the lower
upper scapulars are probably fourth generation. In the mantle (m),
similar grey feathers are growing in. |
At the bottom of this page, a table is
included to show the stage of partial moult in late September in 2cy michahellis.
The exact sequence of the partial moult varies from one individual to
another, but most 2cy michahellis replace some of the following
feathers in autumn: upper tertials, wing-coverts, scapulars and in a few cases
(about 20% of the birds) tail-feathers.
In most individuals, the moult strategy follows a common strategy,
although it may vary between individuals. 2cy michahellis start
the partial moult with shedding the outermost median coverts, followed by
the upper tertials. This moult stage can be found commonly in late August
(see e.g. image 05 in the
August section) but may be found in late September birds as well (image
4733). Most extensive moult takes place in the median and
lower lesser coverts. Moult in the greater coverts is often confined to
the inner feathers and moult in the tertials is often confined to the
upper three feathers. 2cy michahellis hardly replace lesser
coverts, and if they do, there is no distinguishable sequence.
The September moult stage: the innermost greater coverts
and the innermost median coverts are shed. By mid-September, the upper two
tertails are growing and the gap in the outer median covert row is closed
in descendant order (the replacement of new feathers starts in the
outer-wing). In advanced birds, the replacement of the outer median
coverts takes place by late August (see image
09 in the August section), but it can be found commonly in early
September (see this, this
and this individual).
By late September, the partial moult normally includes the upper two
to three tertials (often with the upper tertial growing and the other two
missing), the innermost 1-4 greater coverts, the outer median coverts
(which are already completely grown in many birds) and the inner median
coverts (which may still be growing). The lower lesser coverts are dropped
after the outer median coverts are replaced. By late September, most
individuals have missing both the inner and outer lower lesser coverts, with a few remaining central feathers, and seemingly
random, lesser coverts are dropped as well.
So, by late September, 2cy michahellis
can be found in active moult in almost all wing-covert tracts, while about
20% of the birds also start dropping one or a few tail-feathers (see table
below).
The pattern of the new feathers in the upper-parts vary from plain adult-like grey feathers
to feathers hardly distinguishable from the old second generation
feathers. Those feathers show a anchor pattern or barred pattern (dark
brown to blackish) with a pronounced dark broad shaft-streak on a warm
brown base (but the brown tone of these feathers will bleach to almost
white with time). Some birds show intermediate patterns: a plain grey
base of the feather with a brownish upper half, an anchor pattern and a
clear white fringe.
Some birds may look very advanced, similar to 3cy birds,
after completion of the partial moult, with plain grey feathers in all the
scapulars, the upper four tertials, the inner 10 greater coverts, all
median and lower lesser coverts and all visible lesser coverts. In
general: grey upper-parts and white under-parts in these birds.
It's hard to label the feathers to exact
generation. As can be seen in the 1cy michahellis October section, the post-juvenile moult in michahellis can be very
extensive. Most michahellis moult inner wing-coverts to second
generation in 1cy October. They replace these wing-coverts again in the
complete moult in July-August. Then again, inner wing-coverts are replaced
by the partial moult in 2cy michahellis in September, but not
necessarily the same wing-coverts as in the post-juvenile moult.
This also holds for the tail-feathers.
And the scapulars may have been moulted to fourth or fifth generation feathers, although the pattern is still not completely adult-like. Due to the
considerable moult in the scapulars in 1cy birds in August, in 2cy birds in
April-May, in July and again in September, we exclude the scapulars from
further analysis.
Movements
The majority of michahellis in NW
France arrive from SW origins, the colonies from Marseille and the
Mediterranean coast, Italy, Switzerland and the German Rhine colony. By
September, the majority has arrived in NW France and in the Netherlands,
largest groups can be found in this month. From mid-October, numbers drop
again, as birds migrate south along the Atlantic coast.
Tables
2cy
michahellis scores of new second generation primaries:
Etaples & Le Portel, NW France. |
new
prims |
Sept
21 2002 |
Sept
22 2002 |
Sept
29 2001* |
Sept
29 2001* |
p8 |
1 |
2 |
0 |
0 |
p9 |
51 |
34 |
15 |
5 |
p10 |
23 |
20 |
24 |
12 |
n: |
75 |
56 |
39 |
17 |
m: |
9.3 |
9.3 |
9.6 |
9.7 |
SD: |
0.49 |
0.54 |
0.49 |
0.47 |
Notes:
- * survey at Etaples.
- ** survey at Boulogne-sur-Mer.
- Longest fully grown primaries are scored on two days. Not necessarily
exactly fully grown, but in the folded wing exceeding the previous
primary.
- n: total number; m: average; SD: standard deviation. |
2cy
argenteus scores of new second generation primaries:
September 2000 & 2001 at Scheveningen, the Netherlands. |
new
prims |
05.2001 |
07.2000 |
08.2001 |
09.2000 |
14.
2000 |
21.2000 |
30.2000 |
p6 |
5 |
2 |
1 |
1 |
3 |
1 |
0 |
p7 |
42 |
16 |
18 |
15 |
10 |
18 |
2 |
p8 |
37 |
21 |
17 |
24 |
34 |
56 |
37 |
p9 |
1 |
1 |
1 |
3 |
8 |
25 |
16 |
p10 |
0 |
0 |
0 |
0 |
0 |
5 |
9 |
n: |
85 |
40 |
37 |
43 |
55 |
105 |
64 |
m: |
7.4 |
7.5 |
7.5 |
7.7 |
7.9 |
8.1 |
8.5 |
SD: |
0.62 |
0.64 |
0.61 |
0.64 |
0.73 |
0.79 |
0.78 |
Notes:
- 05.2001 is sample from September 05 2001.
- Longest fully grown primaries are scored. Not necessarily
exactly fully grown, but in the folded wing exceeding the previous
primary.
- n: total number; m: average; SD: standard deviation. |
Moult
score of twenty-two 2cy michahellis, September 20-22 2002, Boulogne-sur-Mer
/ Etaples, NW France
.
|
ind: |
tertials |
gr
cov |
med
cov |
l
l cov |
les
cov |
prims |
recs |
1. |
old |
2
new |
1
new |
2
new |
2
new |
p9 |
old |
2. |
old |
2
new |
2
new |
7
new, 4 miss |
5
new |
p10 |
R1
right miss |
3. |
1
new, 1 miss |
1
new, 1 miss |
10
new, 4 miss |
2
new, 1 miss |
2
miss |
p10 |
old |
4. |
1
new, 1 miss |
1
new, 2 miss |
16
new |
5
new, 13 miss |
25%
miss, 75% new |
p9 |
R1
miss |
5. |
1
new, 1 miss |
2
new |
16
new |
2
new, 2 miss |
2
miss |
p9 |
old |
6. |
1
new, 1 miss |
1
new, 1 miss |
1
new |
old |
4
miss |
p9 |
old |
7. |
2
miss |
2
miss |
1
new, 13 miss |
2
new, 10 miss |
5
miss |
p9 |
old |
8. |
2
miss |
1
new, 1 miss |
13
new |
4
new, 8 miss |
7
miss |
p9 |
- |
9. |
2
new |
2
new |
2
new, 2 miss |
2
miss |
7
miss |
p9 |
- |
10. |
2
new |
2
new |
6
new |
1
new, 1 miss |
old |
p9 |
old |
11. |
2
new |
2
new |
all
new |
3
new |
3
new |
p10 |
R5
new |
12. |
2
new |
2
new, 1 miss |
15
new |
5
new, 3 miss |
2
miss |
p10 |
R1-R6
new |
13. |
2
new |
3
new |
14
new |
5
new |
25%
new |
p10 |
old |
14. |
3
miss |
2
new |
9
new, 2 miss |
1
new |
2
miss |
p9 |
old |
15. |
3
miss |
1
new, 2 miss |
16
new |
13
new, 1 miss |
11
new, 5 miss |
p9 |
old |
16. |
2
new, 1 miss |
1
new, 1 miss |
11
new |
3
new, 15 miss |
2
new, 2 miss |
p10 |
old |
17. |
2
new, 1 miss |
2
new |
14
new |
16
new |
3
miss |
p9 |
old |
18. |
2
new, 1 miss |
3
new |
all
new |
16
miss |
3
miss |
p9 |
old |
19. |
3
new |
4
new |
15
new, 1 miss |
3
new, 1 miss |
50%
miss |
p10 |
old |
20. |
3
new |
4
new |
1
new, 14 miss |
all
new |
40%
new, 20% miss |
p9 |
old |
21. |
3
new |
2
new |
12
new |
4
new, 3 miss |
2
new, 2 miss |
p10 |
old |
22. |
2
new, 2 miss |
1
new, 1 grow |
1
miss |
old |
old |
p9 |
old |
Notes:
Number (amount) of replaced tertials, wing-coverts and scapulars in
1cy michahellis in NW France.
Feather tracts mentioned:
gr cov = greater coverts; med cov = median coverts; l l cov = lower
lesser coverts; les cov = lesser coverts; prims = primaries; recs =
tail-feathers. For details on feather tracts, see the topography
page.
About the scores:
Only recently moulted feathers, included in the partial autumn moult
are mentioned. Moult in the lesser coverts is interpreted by judging
the stage of moult per individual and the pattern of adjacent
feathers. Moulting lesser coverts may still be categorized as
complete moult by September, hence are left out of the scores.
new: feather fully grown in the partial autumn moult;
miss: feather missing or growing, not fully grown;
prim scores are longest fully grown primary, by
September always P9 or P10;
recs are moulting tail-feathers included in the
partial autumn moult, where third generation rectrices replace
second generation feathers. '-' means the tail-feathers were not
seen well enough to fully establish the moult stage.
|
|