Gull notes, two letters from Professor Niko Tinbergen, 1954-1956

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Seattle, 5, 1956
Dept, of Zoology-

Dear Martin,

Thanks for your letter of M a y 12. You sure saw something of gull
behavior! Tell me: do you want your report back, or shall I sendit
on to Mike and Esther?

You must by now get a fairly good idea about the taxonomic rela-
tionships in the group. One thing seems to me obvious: the large
gulls form a very closely knit group, and the ridibundus-type is
another one, with ptptoMon, rldibundus and novae-hollandiae very
close together Indeed. \

It seems to me that w4 are approaching a stage in which one can
see "primitive", original forms of display, one in this species,
one in another. Thus you say that the alarm call of Belcher’s dull
may be primitive. Your criterium is, I believe, that it is less
different from the other calls than it is in most species. I got
a similar idea as to the trumpeting call in the Glauvous-winged
Gull: this is the only thing in which it differs from the Herring
Gull; the single calls are all very much alike, and each is more
drawn-out, and resembles closely the single keew-call as used
for "Stlmmfuehlung" , just social contact. In the European Herring
Gull, there are three phases, one a raw kaw, with in the oblique,
then the throw-down or bend, with one or two smothered, hish-pitched
calls, then the throw-back to the oblique with a series of keews
beginning with a loud and high pithced one, and then the rest with
diminishing loudness and descending pitch, thehead gradually doming
forward and down. The Glaucous-winged rarely does phase 1, it does
the bend with the first call, then throws the head back slowly, and
gives a very equal, level series of Just rather long-drawn, no
staccato keews. Because each is so similar to the general keew call
this gives the impression of being closee to the original form.

In all other displays and calls I can(t see nor hear any difference.
Ecologically the species is more of an open sea fisher than our
H.G., it does much more plunge-diving off shore. It has a dark eye,
the wing tips are very light, a little darker than the blue mantle,
and further the bird is a little stouter than our H.G. But after
having se^n this I begin to wonder how the various HGs behave. I
think a close study of those very closely related species of the
HG superspecies may become very interesting indeed.

I quite agree with your ideas about the redirected attack being
primary in the grass pulling. I would be careful with the coining
of the phrase "pseudo-displacement activities"; it Is quite possible
that all d.a. are in this sense pseudo, the name d.a. ought, I think
to be kept as a provisional, descriptive term, and its causation
should be worked out, but as long as we have not done this in more
cases, I think the coining of a new term would suggest a dichotomy
which need not be there; the term wouod imply that you know that
there are real d.a. as distinct from the paeudo, and that we don't
know yet. But it is extremely interesting that the redirected attack
which in rldibundus is just that, should develop into either feeding
drinking or nest building in Belcher's. In our H.G. it certainly
develops only in grass pulling. It is another example of the posslbi
lity of finding the origin of a movement by comparative study, and
then its radiation. One species may have the primitive form of one

movement and another that of the other. Thus I believe that the
Kittiwake shows a rather primitive form of choking, in which you
can recognise the origin as throwing-down of nest material.

Once one would, In this way, movement by movement, have found the
origins, it becomes of great importance to list exactly in what way
the movement has changed in the courseof radiation, because that
will give us ideas as to what can change in speciation, in the way
of behavior mechanisms.

And all this seems to me to lead to so many intersting aspects,
of evolution, taxonomy, behavior, that it will be a major Job to
work all this out. Publication of all the descriptive stuff may be
difficult, and it may be necessary to keep the full reports as mimoe
graphed reports, andperhaps publish short descriptive accounts in
various journals (if you could do this in papers isazai of a size
acc p table to various ornithological Journals, you might get them
published with at least the drawings in such journals as Wilson
Bulletin, Conddr, Ardea eta (Ardea might accept a paper on a species
that occurs in the West Indies)), and then general conclusions in
Behaviour journals.

Another point: what cb you think of wing tip patterns? I got the
idea that they must be rather adaptive, inclusive perhaps mantle-
color and color of under wing. It struck me that H a rtlaub's gull
has got the white front bar on the upper wide of the wing Just as
ridibundus and Bonaparte* s, but that H a rtlaub's has developed a
rather canus-like wing tip pattern. Now it may be significant that
at least in South Africa there is no other species that has the argen-
tatus or canus-type; it is as if, for species-recognition, the species
in each general region diverge. Usually, when the field ornitholo-
gist uses such a thing as a field character for diagnosis, ( as in
the wing and tail patterns of waders), it seems as if the birds
themselves use it too. If so, Dwight would have taken a rather aw-
kward character to base his classification on. But I can't see any
practicable way of deciding whether these things are social signals
or not, except perhaps: a. zoo observations in which species from

all over the world would respond to flying species with similar
wing patterns (as with the Geese in the Berlin Zoo), b. statistics
about the wing patterns of species within one geographical area.

With al5- these things, I may be talking wet, Just knowing so much
less than you do. I wish we could have another gull conference with
you attending and showing your stuff. Whenever you would come to
Europe, you need only tell us and we would all converge on whatever
roost you would choose to alight l

I guess you are well informed about new developments in Oxford?
Desmond got a job in the London Zoo, Aubrey a lectureship in Edinburgh,
IJli same in Birkbeck college, Fae a fellowship in California, D a vid
Blest a lectureship in University College. That looks like spreading
the gospel indeed.

Our stay here is very pleasant and s ilating. I feel quite at home
as usual, and there vis a lot to see the surroundings. We are ex-
tremely lucky with the weather, and marvel at the Cascades, the
Olympics, the bird islands in the sound, and the giant volcanoes.

Just last weekend I slept In a large glaucous-winged colony and
could, start watching from mytent at daybreak; saw all the displays
many times, and could even take some pictures.

All the best, yrs ever

f

Bonaparte * s , June 17, 1955, I*

Watching birds on shore Dog Lake. All, apparently without ex-
ception, are first— year *birds. Some are in almost complete nuptial
plumage, some in somplete winter plumage. On the whole inactive;
but they do react to my approach, occasionally chase each other or
the Common Terns, and squabble over fence posts.

Have noticed the following hostile patterns:

A

Aerial Lons Call Note. The buzz lest and twang! est note of all.

Long drawn. Could be transcribed as something like "Kaa-aa-aa-aah or
* Kaa- aa~ aanh* .

Series of Lons Call N otes, in the air . Much shorter notes than
the single Long Call Notes. Could be transcribed as n Kaaa-kaaa-kaaa-
kaaa-kaaa. . « Also very buz zy and twangy. Uniform pitch. Done
by swiftly flying bird, with beating wings apparently. (Rather different
from the complete Long Call on the ground, which has the usual

form, and stretches through the 0 and F postures).

Alarm Call ? Very short, buzzy note, impossible to transcribe,
given by "*f lying birds flying over me.

•a, * '

Gakkering. Possibly given by flying bird, certainly by standing
bird, during disputes in which one is trying to force the other off its
fence cost. The standing bird gives it from a posture comme ca:

H

The sound itself can probably be transcribed as "KakakakakaKakakaka, *.» -
with*the N UBual buzzy quality, but in a definitely less extreme degree.
Is thi s the same sound I heard at Buffalo?

Charge Ca ll . Like a single Long Call Note; but not at all P^ly~
syllabicV harsher, very little twang, almost a scream. Apparently
accompanies the Swoop,

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Bonaparte* s, June 17, 1955, II.

Circling . Essentially like Ring-bill in physical form. Perhaps
somewhat more erratic.

Erratic Flight . Reaction to human disturbance. Just like that
of the Black-headed Gull, but even more extreme. May possibly occur
in intra-specific encounters; but a little difficult to tell exactly
what are the releasing and orientating stimuli in some of the doubtful
cases.

Pursuit Flight Pe rformance . I have only seen the Pursuer's
patterns. “Mostly chasing Common Terns. Have had a glimpse or two
of intra- specific chases.

Swoop . With Pursuit Flights. Just like the diagonal Swoops of
the Blaek-headed Gull and Franklin's Gull.

" Flying" Soar . Follows Swoops. Like ordinary Soar; but accomplished
by beating wings.

Soar . Possibly occurs during some Pursuit or Erratic Flights; but
must be rare (at best) and relatively inconspicuous.

The following incident may be significants

One bird, disturbed by me, flies up. Into loose flock of Black and
Common Terns*. Flies round and about, up and down. Occasional Swoops
and "Flying" Soars. Occasional sudden, extreme, downward plunges; with
wings half- folded, motionless. Followed by flying up again. None of
these Scoops or plunges were obviously directed at a particular object
or animal; but they may actually have been so in reality. The whole
performance was perhaps slightly reminiscent of some Erratic Flights
or hawking for insects; but the bird apparently gave Long Call Notes
in flight, and "looked” rather aggressive.

mm mm mm «s «g» r.s» ’’.an gJ sa-sr? -.-2 • s* r-j * «** <d» ssm ss»

Additions, as a result of further observations in the evening.

A large flock was assembled, including perhaps 60 birds, and also includ-
ing a few adults in complete nuptial plumage.

There is definitely a complete Long Call in the air. Sounds Just
like the complete Long Call on the ground or water. Given by rapidly
flying birds, beating their wings very rapidly, (this flight, surprisingly
enough, is not particularly buoyant or tern-like in fact, I haven t
yet seen any tern- like flight under any circumstances in this species;.
Accompanied by a very .marked Oblique Posture, much more marked than any-
thing in Franklin's G, 11 or anything I remember in the Bxack-headed Gull.
The posture is almost reminiscent of the Little Gull s aerial Vertical
posture. No sign of any aerial Forward posture. (All the observed
aerial L 0 ng Calls were given by immatures).

An adult bird was heard to give definite Alarm Calls, obvio 118 -*-^

the same call as the one I cited earlier as "Alarm Call ?* , but Just
a little different. Not as .buzzy. Sometimes one note, sometimes two,
sometimes three. "Keh* or "Keh keh" or "Keh keh keh" .

All sorts of Swoops. Interspecific, toward me. Intra-specific,
during Pursuit Flights after other gulls. Much as described above.
Shallower than those of the Black-headed Cull and (perhaps) those of
Franklin’s Cull. Very rapid, with very rapidly beating wings. Definitely
accompanied by one or two Charge Call Notes. These can be transcribed
as * Keeeeeeeeeer* or " Kreeeeeeeeeee" , (not really 11 Kw arr" ) , but this
gives a very poor impression of them. Buzzy, but definitely scream— like.
These notes stop as soon as the bird starts to fly up.

"Flying* Soar is the usual pattern after Swoops. Also relatively

shallow*

This Swoop-* Flying* Soar routine, incidentally, is not unlike
the behavior of Common Terns in similar situations. But the Swoop-

flying* Soar of the tern is much steeper and more vertical.

.

"Typical* Soars also occur. Still relatively rare and Inconspicuous.
Occur after the most shallow Swoops.

Have seen quite a number of mutual Swoop-* Flying*' Soar performances.
Like those of the Black-headed Cull; like pendulums crossing.

Erratic Flights, or what I have been calling Erratic Flights, are
remarkably common in these species. Whenever, for instance, I put up
a flock of birds from the ground, a large proportion of them will
start Erratic Flights, while it is only a much smaller proportion of
them that will start Pursuit Flights, Swoops, etc. These Erratic
Flights are very erratic indeed, with a terrific amount of twisting
and turning, flying up and down, vertical plunges (sometimes for huge

cli S 1b£lTlO00 ^

They do not appear, very often, to be orientated. It is certainly
my impression, however, that they are rather more than alarm reactions.

I would not be surprised to find that they contained a strong aggressive

element.

In general, it is obvious that all flights of this species tend
to be more erratic than those of related species. This cannot be
ascribed to the small size of the birds, (as Little Culls and xerns
are much less erratic); but it may, conceivably, be due to a combination

of small size, plus short wings.

Bonaparte's, June 17 , 1955 * III*

*

Bonaparte* s, June 18, 1955, I.

Had fewer opportunities to watch the birds today.

Must add a few words about the single adult bird, in complete
nuptial plumage, mentioned yesterday.

Almost certain this was pairing behavior. Not only did the bird do
”G-.C”s toward all nearby flying birds, but it also made little circular
flights in the interim (usually not very aggressive, no more than Long
Call Notes) , and occasionally attacked nearby birds on the ground, (I
couldn’t watch this very well, but the whole procedure was very reminiscent
of territorial defense). A post, I suppose, is exactly what one would
expect as a station in a tree-nesting gull. The displaying bird, unfort-
unately, got very little response from the flying birds. So, the adult
would occasionally fly over to a nearby first-year bird on the ground,
and would then apparently attempt an ”M. C. " • I definitely saw the adult
do Oblique - Forward - Upright and Head-flagging - Head-tossing in these
circumstances. Again, these didn’t seem to get much in the way of a
response.

Today, however, the adult may have succeeded partially. At least,
one of the nearby first-year birds goes into a pronounced Upright, facing
away from the adult, whenever the adult flies by.

I S i I I .

I am beginning to believe that this adult bird must be a female.

It is not very large, and it is not nearly as aggressive as some of the
first-year birds. In addition, the fact that the adult did Head-tossing
during its ”M.C.” is perhaps suggestive.

All the aerial hostile patterns I have observed in this adult seem
to be essentially the same, perhaps identical, with those of the first-year
birds. I have heard first-year birds give Alarm Calls exactly like those
of the adult cited yesterday. I have heard the adult give a long series
of Long Call Notes exactly like those of the first-year birds. I have
also observed a Swoop-” Flying” Soar performance by the adult exactly like
the corresponding performances by first-year birds. I can’t detect any
difference between the voice of the adult and those of the first-year
birds, (Unfortunately, however, I have not yet observed a complete aerial
Long Call and aerial Oblique Posture by the adult.)

More about glides, in both first-year birds and the adult. Both
glide frequently in ordinary flight. Obviously quite unritualized.

The first-year birds also glide very frequently during their Erratic
Flights. (I have never, incidentally, seen a really good Erratic Flight
by the adult) • These glides during Erratic Flights may be either upward
or downward. Sometimes quite prolonged — relatively. They do not
appear to be ritualized either; but they can be easily interpreted as
ambivalent hostile activities. They are thus, in all probability, strictly
comparable to those of the Ring-bill; but they seem to be confined to a
rather different situation.

Bonaparte's, June 19, 1955, I.

Correction about the adult bird previously mentioned. Apparently
a male. Attempted a forced copulation with one of the first-year birds.
Just like Black-headed Gull. Copulation Call like that of the Black-headed
Gull, except for the characteristic buz>z.y twang.

This adult continued its usual pairing behavior. Tonight, however,
it did several diagonal Swoops and Soars (at Black Tern) during its little
circular flights. These Swoops and Soars were exactly like those of the
Black-heade d Gull . in physical form. Couldn't hear accompanying call
very well, but I think it was a "Plaintive Charge Call" (i.e. long-drawn
screaming Long Call Notes) during the Swoop.

Still haven't seen adult give aerial Long Call in Oblique.

When I frighten a group of birds, they fly up, circle, silently.

This can certainly be classed as a Silent Panic.

-

Watched an Erratic Flight performance this morning, involving two

first— year Bonaparte's Gulls and one Common Tern, It was noticeable that

these "Erratic Flights" of the gulls included definite Swoops and "Flying

Soars , and that the Swoops were definitely directed, either toward the

other gull or toward the tern. This was definitely a relatively aggressive
performance.

A few additional remarks about "Gakkering" . Often given by bird
on post (first-year bird) when a flying bird tries to dislodge it.
Characteristic sequence is "Gakkering" (in semi-U or semi— 0 posture, very
well orientated, neck sometimes twisted, wings sometimes raised anchor
spread) - Oblique - Forward. The "Gakkering Call" itself is harsh,
rapid, and rhythmic. Very much like The Forward Call part of the Long
Call. Possibly louder??? The rhythm of the whole usual performance can
be indicated by the following diagram*

There seems to be a Landing Call. Given by both the adult and the
first-year birds. Relatively rare, (as one might expect in the circum-
stances, as the birds either land far apart or fight over a small post).
Can't say much about it. A series of Long Call Notes, getting successively
shorter and deeper in pitch, I think. Probably has fewer notes than the
Landing Calls of some other species.

Bonaparte* s, June 20, 1955* X

The unmated adult is still here, and so are a few first-year birds,
but the great bulk of the birds seems to have gone. Got very little
today.

The unmated adult is still behaving as before; but much duller, as
there are fewer birds around to encounter or to display to. He seems
to have formed or re-formed his connection with a first-year bird. The
latter stays around in his neighborhood, calls to him when he flies over-
head, and goes into a U when he flies very near.

Notice that the Alarm Call of flying birds is often uttered with
apparently closed bill. The adult gave continuous monosyllabic Alarm
Notes as he circled around me, and repeatedly made shallow Swoops toward
me and then moved away in shallow w Flying** Soars. Thus the Alarm Call
must contain an appreciable element of attack drive.

Saw both adult and first-year bird give repeated series of Long
Call Notes from completely relaxed, almost hunched, postures. This was
definitely not a display posture.

The voice of the adult, at moderate intensities at least, is defin-
itely less buz2.y, more of a scream, than that of his first-year ^mate” «
The difference is slight, how ever .

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