PROCEEDINGS OF THE UNITED STATES NATIONAL MUSEUM

issued il^i'\A.Sl'^M>il ^H '^*

SMITHSONIAN INSTITUTION
U. S. NATIONAL MTJSEUM

Vol. 106 Washington : 1956 No. 3374

FURTHER DATA ON AFRICAN PARASITIC CUCKOOS

By Herbert Friedmann

The data assembled in this paper extend our knowledge of African

parasitic cuckoosfrom the basic facts presented in my book entitled
"The Parasitic Cuckoos of Africa" published in 1949 and the little
addendum (Friedmann, 1949b) published later in the same year.
to it
In the present account only those data pertinent to the problems of
parasitic reproduction are included. For this reason, some species
of cuckoos are not discussed at all as there were no new facts to
describe. Even in those that are treated in this paper almost no
space is given to such items as call-notes, migration, food of the adult,
plumages, etc.
For sending me unpublished information, for answering many
queries, and for other acts of kind cooperation, I am indebted to the
following naturalists: C. F. Belcher, J. P. Chapin, F. Haverschmidt,
R. Liversidge, P. Millstein, E. Pike, O. P. M. Prozesky, R. A. Reed,
and W. Stanford. The photographs illustrating this paper were
supplied by Mr. Liversidge in addition to his information. His
observations on the red-chested cuckoo (Liversidge, 1955), unpub-
lished when sent to me, fortunately appeared in print just as the
present manuscript was about to leave my hands. I have therefore
merely summarized his data insofar as integrating all the information
has permitted.
386753—56- 377
378 PROCEEDINGS OF THE NATIONAL MUSEUM vol. loe

Clamator glandarius (Linnaeus)

Great-spotted cuckoo

Three new hosts may be added to those listed m

my book, one of
whicli a race of a species of which another form was already
is onl}^

recorded as a victim of the great-spotted cuckoo. The first of these

is the dwarf raven (Corvus corax edilhae), a bird of which Belcher

(1949a, p. 37) records a nest found near Gabredarre, Ogaden, Italian

Somaliland, containing tliree or four eggs of the great-spotted cuckoo
and three or four eggs as well as one young of the host. Previously
only the Spanish race of this raven (C. c. hispanus) had been reported
as a host of the nominate race of the cuckoo.
The second is the piapiac (Ptilostomus afer). I have no data on
this other than that it is mentioned as a victim of the great-spotted
cuckoo by Mackworth-Praed and Grant (1952, p. 505).
The third new host is the long-tailed glossy starling {Lamprotornis
caudatus). Madden (1934, p. 94) saw a fledgling great-spotted cuckoo
apparently being fed by a pair of these starlings at El Obeid, Kordo-
fan, Sudan, in November 1932. Aside from adding an additional
species to the known hosts of this parasite, this record seems to be
the first for any hole-nesting bird north of South Africa. As pointed
out in my book (Friedmann, 1949a, p. 10), the data then available
showed a puzzling divergence in the choice of victims by this cuckoo
in South Africa on the one hand, and ever3''where else in its range on
the other. In the former area most of the records were of hole-nesting
birds (three species of starlings), while all the records from Nyasaland
and Southern Khodesia north to the Mediterranean lands were of birds
making open nests in trees or on ledges (crows and magpies of several
species). It is true that one open-nest builder, the Cape rook, was
known to be parasitized in South Africa; now we may further reduce
the apparent geographical disparity in host selection wTth this indica-
tion of the choice of a hole-nesting victim as far north as the Sudan.
In my earlier report (Friedmann, 1949a) I considered all the pop-
ulations of the great-spotted cuckoo as one taxonomic unit. Since
then, Clancey (1951, p. 141) has separated the birds south of the
Sahara under the name choragium, restricting the name glandarius
to the birds that breed in Spain, Portugal, Mediterranean Africa,
Cyprus, Greece, Asia Minor, and Iran and that migrate south in
winter to tropical Africa. It follows from this that there again seems
to be a regional difference in host selection. The nominate race is
known to utilize only open nests, chiefly of corvids of several kinds,
whereas the African choragium lays in such nests and in those of
hole-nesting starlings as well.
 AFRICAN PARASITIC CUCKOOS —FRIEDMANN 379
Clamator jacobinus (Boddaert)

Jacobin cuckoo

New information gathered on the jacobin cuckoo does more than

merely yield additional instances of its parasitism on previously
recorded hosts. Many additional records of its eggs in nests of
Layard's bulbuls (Pycnonotus tricolor layardi), sombre bulbuls
(xindropadus imjmrlunus im/portunus) and fiscal shrikes (Lanius
,

collaris collaris) corroborate the predominance these hosts play in

the economy of the parasite, but, merely as records, they add no
reveahng new data. Two additional instances of the bakbakiri
(Telephonus zeylonus) added to the three listed in my book suggest
that this shrike is more frequentlj^ victimized than was formerly
suspected. These instances are: (1) a nest found at Butterworth,
Cape Province, by Pike on February 4 with one egg of the host and
one of the parasite, and (2) a case reported by A. W. Vincent (1949,
p. 138) from Kichmond, Natal.
Two new host species may be recorded. Skead (1954, p. 46)
reports that an egg of this cuckoo was found in a nest of the paradise
flycatcher (Terpsiphone viridis perspicillata) at Fleet Ditch Kloof,
near King Wilham's Town, eastern Cape Province, on Dec. 18, 1954.
In the Victoria Memorial Museum at Salisbury in 1951 I found a set
of two eggs of the grassbu'd {Sphenoeacus ojer transvaalensis) with
one egg of the jacobin cuckoo, taken at Inyanga, Southern Rhodesia
(no date), by Flight Lt. E. F. Allen. This is not only a record of a
new host but also is the only instance kno\\ai to me of the parasite
using a nest built close to the ground. The one record of the jacobin
parasitizing a kingfisher, described in m}^ book (Friedmann, 1949a,
p. 31), may well be questioned; it may have been a honey-guide's agg.
In my earher account (Friedmann, 1949a, p. 36) of the hosts of this
parasite I raised some doubts as to two cases recorded by de Klerk
of the jacobin cuckoo laying its eggs in nests of the yellow- throated
sparrow (Petronia superciliaris) and suggested that the records might
refer to honey-guides. However, de Klerk's measurements are too
large for any hone^^-guide's eggs and agree very closely v/ith other,
authentic eggs of the jacobin cuckoo. The records must therefore
be accepted, but it remains that the host is a very unusual one,
nesting, as it does, in holes in trees, a t^^pe of nesting site not otherwise
known to be utilized by the jacobin cuckoo.

Incubation Period

The incubation period is still to be determined, but Skead's (1951,

pp. 171-172) incomplete evidence suggests a shorter period thantdid
my own similarly fractional data (Friedmann, 1949a,\p. 37)._^It may
380 PROCEEDINGS OF THE NATIONAL MUSEUM vol. loe

be recalled that the nest I was watching was destroyed 14 days after
the last host's egg was laid and the smashed cuckoo's egg in the nest
seemed then about in hatching condition. One of Skead's cases
suggested a possible incubation period of 12)^ days, while his second
one seemed three days less, and caused him to ask if the cuckoo's
egg might begin its development some time prior to ovulation. It
would seem unlikely that the incubation period is under 12^ days.
Evicting Habit

Our knowledge of the evicting habit in the jacobin cuckoo stiU

requires clarification. In 1949 (Friedmann, 1949a, p. 37) I was aware
of no real evidence and could only conclude that while the young
cuckoo was almost always the sole survivor of a nest, the absence of
any of the rightful young might be the result of their failure to compete
for food with their parasitic nest-mate rather than that they were
actually evicted alive by it. Skead (1951, pp. 172-173) has described
an instance in which the nestling jacobin tolerated eggs and young
in the nest for up to four days, and another in which it made no
attempt to evict eggs for four days but in which a chick of the host
was found to disappear within less than a da}^ after hatching (but
not necessarily evicted by the cuckoo). In the first case, Skead
conmients that, considering how the young cuckoo covered and almost
suffocated its nest-mate (a Layard's bulbul), the "possibility of death
of nestlings b}^ this means must not be ruled out. The chick was so
weak . . . when retrieved . . . that had it remained there much
longer, it would have been smothered. Therefore, I wish to draw
attention to the suffocation of cuckoo's nest-mates and the possibility
of their subsequent removal by the foster-parents during nest-sani-
tation . . .
."

In this connection, an observation by Pike recorded by Godfrey

(1939, p. 23) and inadvertently overlooked in my earlier report is of
interest. At Butterworth, Transkei, Cape Province, he found a nest
of a fiscal shrike on January 15 containing only one egg— a jacobin
cuckoo's. On February 27 the nest was revisited and was found to
hold a young jacobin and three j^oung shrikes. Still later he saw
the four young birds perched on twigs near the nest, and still later
he noted that the young shrikes had left but the cuckoo was still
being fed by an adult shrike. This is the only definite instance
known to me of the nest-mates surviving together with a jacobin
cuckoo. From this, and from the two instances described b}' Skead,
in which eviction by the cuckoo, if any took place, did not occur for
some days after hatching, it follows that evicting is certainly not
invariable or immediate, and, for that matter, it is still not possible
to say that the young parasite was responsible for any evictions.
 AFRICAN PARASITIC CUCKOOS —FRIEDMANN 381

Actually, thisis not surprising when we consider that in the related

great-spotted cuckoo, the rightful 3"ouug often, and perhaps even

regularly, grow up together with the parasite. Furthermore, two
j^oung jacobin cuckoos have been noted as growing up together with
one of the young of a babbler host (Argya rnalcomli) in India, a further
indication of the lack of evicting behavior (Bates, 1938, p. 125).

Egg Laying
Pike (again cited in Godfrey, 1939) found, on December 4, another
nest of the fiscal slmke containing one egg of the jacobin cuckoo; two
days later contained two shrike's eggs in addition to the cuckoo's,
it

and three days later still it held five eggs of the slniko and the one
egg of the jacobin. Pike was unable to visit the nest again. It would
appear from this case and the one referred to previously that the
cuckoo may ia}'^ occasionally into nests before the builder has begun
to lay. It is, of course, not impossible that in each instance there
may have been a single shrike's egg present and that the cuckoo
removed it Vv^hen laying its own. It is, however, not very likely that
this was the case, as the usual number of eggs m a clutch of the shrilve
is four or five, and was no reduction as would
in these cases there
have been the case had the cuckoo removed an egg. In my earlier
account of this parasite (Friedmann, 1949a, p. 30) I summarized the
then available information on this point, and I have seen no evidence
to cause me to change it since: "No one has witnessed the actual
deposition of the egg, but judging by the fact that the number of
eggs of the host is usually less than the fuU complement by the number
of cuckoos' eggs in the nest, it seems that the cuckoo usually removes

an egg before laying into the nest ." Of course, if

. . . as seems to —
have been the case in the two instances described above the cuckoo's —
egg is laid first there can be no such removal of a host's egg.
An observation suggesting that the jacobin cuckoo may at times
be an egg-eating nest robber is the following sent me by Pike, who
once saw some red bishop birds (Euplectes orix) chasing one of these
cuckoos away from their nests. Two of the nests each had a hole of
about an inch and a half m
diameter torn in the lower part; one of
the nests contained two eggs of the builder, one being badly pecked
and empty and the other also broken; the bottom of the nest was
moist with spilled yolk.
In my earlier account of this cuckoo (Friedmann, 1949a, p. 30) it
M^as mentioned that only a single instance was known in Africa of
more than two eggs of this bird in one nest, although in India Stuart
Baker had recorded six nests with three eggs of the parasite apiece,
two nests with four, and one nest with six eggs attributable to the
jacobin cuckoo. The lone African instance was a nest of the sombre
382 PROCEEDINGS OF THE NATIONAL IVIUSEUM vol. loe

bulbul (Andropadus imjportunus) with five eggs of the cuckoo and

two of the host, reported by Ivy (1901, pp. 23-24). It was to be
expected that the situation in the African population of this cuckoo
would prove to be similar to that in the Asiatic part, and this has
recently been found to be the case. Air. H. M. Miles of Salisbury,
Southern Rhodesia, informs me (in litt.) that nearby, at Plumtree
School, on Mar. 2, 1954, Mr. Ian Canncll found a nest of Layard's
bulbul containing seven eggs of the cuckoo and one broken egg of
the bulbul. He also found another nest of the same host with four
eggs of the cuckoo and one egg of the bulbul.

Clamator levaillantii (Swainson)

Stripe-breasted cuckoo

Recently m South Africa, Milstein (1954, pp. 4-5) produced evi-

dence indicating that this cuckoo may \?iy pm-e white eggs as well as
the greenish blue and pinkish ones described in my book. The case
is as follows. He saw two stripe-breasted cuckoos perching in a
wild fig tree in which there was a nest of a yellow-vented bulbul.
The cuckoos repeatedly hopped towards the nest and the bulbuls
kept them at baj^, diving onto their backs, gripping, and literally
riding them to the ground. The intruders, fluttering wildly, never
retaliated even when one of the bulbuls yanked out a tuft of whitish
breast feathers from one of the cuckoos. Milstein watched this
repeated attack and counterattack for over an hour and a half. He
then left, but returned several hours later when he examined the
bulbul's nest for the first time. It contained tvro eggs of the bulbul
and four large white eggs, presumably of a cuckoo. The two bulbul
eggs and three of the parasitic eggs were snugly settled in the nest,
but the fourth v/hite egg, slightly pinkish (freshest?) was on the rim
of the nest, almost falling out. Milstein was inclined to assume that
the fresh egg had been laid during his absence by one of the stripe-
breasted cuckoos that had shown so much interest in the nest a
few hours earlier. The four white eggs measured, in millimeters, as
follows: 27 X 22.25, 26.6 X
21, 26.25 X
21.5, and 25.25 20.75. X
In this respect they agree with Ivnown eggs of both the stripe-breasted
cuckoo and the pied cuckoo; in rolor they agree with authentic eggs
of the latter. It is to be hoped that fm-ther observations, including
allo\ving such eggs to hatch and develop into diagnostically feathered
chicks, may be forthcoming.
To the stiU meager data on the breeding season of this cuckoo in
various parts of its range maj^ be added the fact that Verheyen
(1953, pp. 305-306) records that it breeds during the rainy season in
the Upemba Park, Belgian Congo, where males with swollen gonads
were collected on October 7 and March 15.
 AFRICAN PARASITIC CUCKOOS —FRIEDMANN 383
Cuculus canorus gularis Stephens

African yellow-billed cuckoo

Information about this bird accumulates very slowly; however, it
is possible to add a new species to the small list of its known hosts.
Mr. H. ]M. Miles, of Salisbury, writes me that a correspondent of his,
J. R. Peters, found a nest of Layard's bulbui {Pycnonotus tricolor
layardi) containing a young yellow-billed cuckoo on Oct. 9, 1955, at a
spot called Guinea Fowl, 12 miles southeast of Gwelo, Southern
Rhodesia. At that time the nest contained two newly hatched
bulbuls as well as the 3^oung cuckoo. Ten days later the nest was
revisited and was found to hold only the .young parasite. It was at
this visit that the young cuckoo, then well feathered, was identified
as Cuculus canorus gularis.
Neuby-Varty (1950, p. 38) found the gray-headed sparrow (Passer
griseus diffusus) to be victimized in
Southern Rhodesia. At his farm
'Torre," near Marandellas (no date given), he found a nest of this
sparrow in a natural hole in a sugarbush containing two eggs of the
sparrow and one of the African yellow-billed cuckoo. The egg of the
latter was very light bluish gray with blotches of slate and dark
greenish olive brown, mostly at the thick end but also scattered over
the rest of the egg, and measured 23.5 X 16 mm. This agrees very
well with the fully authenticated oviduct egg described by the same
observer (Neuby-Varty, 1948, p. 158) two years earher. This record
is the first indication that this cuckoo lays in nests in holes in trees,

the few previous records involving open nests in exposed situations.

In the Victoria Memorial Aluseum, Salisbury, is an egg, supposedly
of this cuckoo, collected by D. Townley at Rumani, Southern Rho-
desia,Dec. 20, 1929, from a nest of an unidentified babbler. The label
on the egg bears the words "identity sure," and the egg is not unlike
authentic ones of this cuckoo. No babbler has yet been recorded as a
victim of this bird, but this case must remain unidentified.

Cuculus solitarius Stephens

Red-chested cuckoo
Plates 1-3
Our knowledge of several phases of the life history of this cuckoo
has increased greatl}' in the past few years. For easier reference and
discussion these data maj- be treated under various subheadings.

Eggs and Egg Laying

Whencompiling the data for my book I was unable to find an
instance ofmore than one egg of this cuckoo in any single nest. Tait
(1952, p. 135) has since recorded such a case —
a nest of Cossypha
384 PROCEEDINGS OF THE NATIONAL MUSEUM vol. iM

caffra drakenshergi with one egg of the host and two of the cuckoo.
The^cuckoo eggs were sufficiently dissimilar in color to suggest that
they were laid by different hens. There is, as yet, no evidence to
indicate that the same cuckoo may lay more than one egg in a nest.
While it is true, as stated in my book (Friedmann, 1949a, pp.
68-69), that the great majority of the known eggs of the red-chested
cuckoo are plain pale chocolate brown or olive brown without any
markings, a sufficient number of divergent types since have been
recorded from southern Africa to make it seem that there is more
variation than was formerly apparent. Thus, in the Bryanston dis-
trict, near Johannesburg, Transvaal, Reed (in litt.) found a red-
chested cuckoo's egg in a nest of a Cape robin-chat together with
one egg of the owner, which it closely resembled, being olive green in
color and heavily blotched with reddish brown, the blotches forming
an almost solid mass at the obtuse end of the egg. In another nest
of the same host species he found another egg of this cuckoo (identity
certain because the egg was allowed to hatch and the development
of the chick foUov/ed in detail). This egg has a fawn colored ground
and was heavil}- blotched with dark reddish brown. Still another
parasitized nest of the Cape robin-chat was found containing an egg
that presumably was of this cuckoo. It vv'as "off white" in color and
very heavily speckled with large, dark brown spots.^ That three
such divergent, blotched or spotted eggs were found in one locality
seems to eliminate, or to render doubtful, the possibility of their being
unusual or pathological in any sense. Still another color variant has
recently been described in Northern Rhodesia by Haydock (1950,
pp. 149-150) as deep cream in color and with a very rough shell
textm'e.
So few observations are available on the question of whether or
not the hen cuckoo removes an egg of the host when depositing one
of its own that the following case, incomplete as it is, is of some
interest. Liversidge (in litt.) writes me of a parasitized nest of a
Cape robin-chat, a bird whose name occurs under several headings in
our present discussion. The nest was found by the observer's land-
lady, who was certain that one day the nest was empty and the next
day it contained an egg of the cuckoo and one of the host. This
would suggest that the cuckoo laid the same day as the robin-chat
and did not remove an egg at that time. It is, of com-se, not impos-
sible that the cuckoo may have laid its agg earher in the day than did
the host, and that there was no egg for it to remove. As I have
recorded previously (Friedmann, 1949a, pp. 69-70), the red-chested
cuckoo seems to lay not infrequently in nests before the host has

' This epg Is not dissimilar to known eggs of Cucidus cafer; the possibility cannot be ruled out that the
egg may have been of the black cuckoo.
 AFRICAN PARASITIC CUCKOOS —FRIEDMANN 385

begun laying. In at least one instance where the cuckoo laid in a

nest already containing eggs of the host,it did remove one of them

when depositing its own.

Incubation Period

The nest of a Cape robin-chat studied by Liversidge, mentioned

above, yields the following information on this topic. The person
who first showed him the nest was quite definite that the eggs (one
each of the cuckoo and the host) were first present on September 21
(about a week before Liversidge first saw the nest). The cuckoo's egg
hatched on October 7, which implies that at least 16 days elapsed
between the time the egg was laid and the date of hatching. It is not
known, however, if the host began to incubate on or immediately
after September 21; but at least it is clear that no eggs were laid in
the nest subsequent to the two seen on September 21, and it would
seem that incubation may well have started on or just after the date
of laying. This account leaves much to be desired, but it suggests an
incubation period somewhat longer than the one of 12}^ days reported
for the related European cuckoo, Cuculus canorus. More accurate
data are needed to clarify this point.

Host Records

Accumulation of many additional host records indicates even more

strongly than before that the robin-chat Cossypha cafra is the chief
victim of the red-chested cuckoo wherever the two occur together.
At least three subspecies of the Cossypha are now knovfn to be para-
sitized: thenominate one, for which I had numerous records previously
and which I could add at least as many new ones now; the tropical
for
race iolaema, earlier listed on the basis of a single instance to which
several more may now be added (Wiley, 1948, p. 2) and the subspecies
;

drakensbergi, recently found to be victimized at Underberg, Natal,

by Tait (1952, p. 135).
The Natal robin-chat was previously listed as a host of the red-
chested cuckoo solely on the strength of Roberts (1939, p. 8), who lists
it in this capacity. No specific records or other data were given by
him. Recently Mr. H. M. Miles has informed me that at Melsetter,
Southern Rhodesia, a correspondent of his saw a fledgling of this cuckoo
attended by a Natal robin-chat. There is nothing unexpected about
this record, but it is a welcome additional indication to the not too
informative statement given by Roberts.
One species has been added to the list of known hosts by Haydock
(1950, pp. 149-150) at Luanshya, Northern Rhodesia. This is
another species of robin-chat, Cossypha heuglini, in a nest of which he
found, on October 22, two eggs of the owner. An hour later a red-
386753—56 2
386 PROCEEDINGS OF THE NATIONAL MUSEUM vol. los

chested cuckoo was heard caHmg near the nest. It was perched on a
small branch nearb}'. One of the chats was standing on the edge of
the nest in a threatening attitude. The next morning the nest con-
tained one of the original eggs and another much larger one that
measured 25.7X18.2 mm. It was ovate, deep cream in color, and had
a very rough shell texture. A week later the nest was found to be
empty when revisited. Benson (1951, p. 98) found another parasitized
nest of this robin-chat at Ketete, Nyasaland, on September 22. It
contained one egg of the host and one of the cuckoo. Near IMar-
andellas. Southern Rhodesia, Neuby-Varty has reported seeing a
young fledgling red-chested cuckoo with a Heuglin's robin-chat,
according to information received from Mr. Miles.
Haydock (1950, pp. 149-150) records the following observation
suggestive of the possible utilization of a hole-nesting host, a starling,
by the red-chested cuckoo. He writes that on Dec. 17, 1948, a pair of
Lamprocolius were seen feeding a 3'oung cuckoo recently out of
sp.
the nest. On close inspection (the bird was caught) the latter appeared
to be a red-chested cuckoo chick, having "the tj^pical yellow legs and
feet, and differing from C. gularis in being practically a uniform black,
though there were some traces of rufous on the thighs and marginal
tail feathers, and odd feathers of the breast were tipped with a dirty
white . .
."
. The identification of the starling fosterer was im-
possible as the birds were not collected, but Haydock was inclined to
feel that the species probably was Lamprocolius chloropterus elisa-
beth, a form previously known to be nesting in the area.
Aside from the uncertainty as to the species of starhng involved,
the evidence is not suflficiently conclusive to enable us to add it (even
as Lamprocolius sp.) to the list of birds definitely recorded as hosts of
this parasite. Merely seeing a bird feeding a fledged cuckoo is not
necessarily proof that it raised the latter. In tliis connection, it may
be pointed out that Harding (1948, p. 2) saw a young red-chested
cuckoo, out of the nest, being fed by a Fischer's slaty flycatcher
{Dioptrornis fischeri) The next day he observed the same cuckoo
.

being fed by a pair of collared sunbirds (Anthreptes collaris).

In their recent book, Mackworth-Praed and Grant (1955, pp. 318-
319) write that the white-winged scrub-robin (Erythropygia leucophrys
leucophrys) is a common
victim of the red-chested cuckoo. Just
what data or how many actual instances of parasitism were known to
these authors is, unfortunately, not stated. In my earlier account I
listed but a single record under the name Erythropygia leucophrys
brunneiceps. This refers to the same bird, although it may be pointed
out that Mackworth-Praed and Grant use the name brunneiceps now
for a race of Erythropygia zambesiana, a species not yet found to be
imposed upon b}^ the red-chested cuckoo.
 AFRICAN PARASITIC CUCKOOS —FRIEDMANN 387

The Cape wagtail (Motacilla capensis) was previously listed as a

host of the red-chested cuckoo on the basis of two records. To these
may be added two more, a nest found in the Transkei, Cape Province,
containing one egg" of the cuckoo (E. Pike, in litt.), and a nest of the
race Motacillo capensis wellsi with three eggs of the wagtail and one of
the cuckoo found at Kabete, Kenya Colony, Apr. 26, 1944, by Wiley
(1948, p. 2).
A second record of the dusky flycatcher (Alseonax adustus) as a
victim of this cuckoo comes to me from Walter Stanford (in litt.).
On Dec, 22, 1952, at Deep Gat, Somerset West, Cape Province, he
found a nest of this flycatcher containing one egg of its own and one
egg of the red-chested cuckoo. The nest was built in a little recessed
hollow in a tree trunk. On revisiting the nest two weeks later, Stan-
ford found that the cuckoo's eggs had not hatched, while the fl}''-
had done so and the young Alseonax was well feathered.
catcher's egg
A host overlooked when compiling my earlier account is the Cape
sparrow (Passer melanurus melanurus). C. J. Bergh (1942, p. 99)
writes from Belfast, Transvaal, that he has known the red-chested
cuckoo to lay its eggs in a "mossie's nest." The name "mossie" is
loosely used for several species of sparrows but is usually intended to
refer to the Cape sparrow.
Evicting Instinct

Considering the complete absence of any information on this topic

up to the time of the completion of my 1949 book, and, indeed, the
amount of evidence suggestive of the lack of such behavior, it is
gratifying that a number of detailed pertinent observations have since
been made. There can no longer be aiy doubt that the evicting of
nest-mates is commonly, perhaps regulrrly, done by newly hatched
red-chested cuckoos, but it still seems n-^cessary to quahfy this with
the statement that it seems to be less universal in this species than
in the European cuckoo. In the latter bird it is extremely rare to find
the young cuckoo tolerating any nest-mates.
Aside from Pringie's account, which I have briefly described
(Friedmann, 1949b, p. 515), I now have data on three more instances,
all in nests of the Cape robin-chat (Cossypha cafra). Reed (1952,
pp. 14-15) has described one such instance, and has kindly supplied
me with fuller details, which are given here as they constitute a
better record of events than his condensed printed one.
On Dec. 25, 1951, near Johannesburg, Reed discovered a nest of the
robin-chat with three eggs, one of which was a red-chested cuckoo's.
The nest close observation, and on the morning of
was kept under
January he found that the cuckoo's egg had hatched but that the
1

robin's eggs were still intact in the nest. By 7 a. m. of January 3

388 PROCEEDINGS OF THE NATIONAL MUSEUM vol. loe

both robin eggs had been ejected from the nest. One lay on the ground
at the base of the tree and the other lay in a fork of the tree beside
the nest. Fortunately, neither egg was visibly damaged. Reed
replaced one egg in the nest with the cuckoo chick and watched the
bird try immodiatel}^ to eject it. However, when the chick had gone
through the performance of carrying the egg on its back to the edge of
the nest and the egg had rolled back into the nest, Reed removed it.
He repeated the experiment on January 4 with the same result, but
when he tried again on January 5 he was unable to induce the chick
to attempt to eject the egg. To make quite certain that the bird
had lost all desire to eject it, ho placed the egg on its back, but the
chick lay passiveh^ and made no attempt even to remove it. Reed
then broke both robin-chat eggs and found no sign of incubation in
them.
Another nest was found by the same observer on November 29.
It contained a newly hatched cuckoo and a robin-chat's egg. On
December 2 the host's egg was found lying on the ground just below
the nest. Reed replaced this egg in the nest two da3'-s later, but not
even putting it directly on the back of the chick would induce the
latter to attempt to evict it. The young cuckoo was at least 6 days
old at that time; therefore, this case agrees with the more fully de-
scribed one in that the instinct to eject the nest-mates was found to
disappear before that age.
Near Cape Town, Liversidge (in litt.) watched a nest of a Cape
robin-chat containing one egg of the cuckoo and one of the host and
made detailed notes. (Also, he kindly supphed the photographs for
use in this paper.) The cuckoo egg hatched, and within 24 hours the
host's egg was out of the nest. Liversidge put it back, but a few houis
later, when Dr. Broekhuysen was out again. The
visited the nest, it

lady in whose garden the nest was located was asked to keep up
observations, to replace the egg in the nest each time, and to see how
often the young cuckoo would evict the egg. After three more
evictions in 10 minutes, the observer lost some of her attentiveness
but reported that the chick threw out the egg many more times that
day, the total of such evictions being in excess of 25 by the time the
observations were called off. The chick was later given eggs of various
sizes and shapes, and even small stones, all of which it evicted, or, at
least, attempted to eject. This went on until its fourth day. On
the fifth day an egg, when placed in the nest, was allowed to remain
there, but the ejection movements of the j'^oung cuckoo could still be
induced by prodding its back. Liversidge writes that the chick did
not have any hollow space on its back, which was quite broad and flat.
Wlien ejecting an object, however, the back becomes slightly concave,
as in the case of the well known European species, Cuculus canorus.
PROC. U. S. NAT. MUS.. VOL. 106 FRIEDMANN, PLATE 1

Cape robiii-chai bringing food to a \"oung red-chcstcd cuckoo in its nest.

(Photo b\" Livcrsidge.)
PROC. U. S. NAT. MUS.. VOL. 105 FRIEDMANN, PLATE 2

()i)c-cla\-old rcd-clu'Slcd cuckoo cjccling host's cgi; from ncsi. ScquL-iicf; Ictl lo right
top to bottom. (Photos b\- Liversidge.)
PROC. U. S. NAT. MUS.. VOL. 106 FRIEDMANN. PLATE 3

tirowlh of young red-chested cuckoo. Tup: 3 daxs old; first feathers breaking out of
tlieir sheaths. Bottom: 19 da\ s old; one da\- before leaving the nest. (Photos by
Liversidge.)
 AFRICAN PARASITIC CUCKOOS —FRIEDMANN 389

The Nestling and Its Development

When I wrote my earlier account of this cuckoo there were ahnost

no data on the giowth and development of the young bird in the nest.
Within the next year Hughes (1949, pp. 2-4) filled in part of this gap
in our knowledge, while Liversidge (1955) has since filled a good deal
more. In addition to these data, there are available some observa-
tions kindly supplied me by R. A. Reed. It is from these three com-
bined sources that the following description is compiled.
To begin with the more important items: The duration of the
nestling stage (i. e., from the time of hatchmg to leaving the nest) of
this cuckoo in nests of the Cape robin-chat was found by Reed to be
from 17K to 19 days, by Liversidge to be 20 days ±6 hours, while in
the nest watched by Hughes the young cuckoo was found to have been
killed by a predator in the nest on its 20th day; one leg, severed from
the body, was all that remained. The nestling period for the related
European cuckoo, Cuculus canorus, is 20 days, agreeing with the
data for the present species.
On hatching, the young cuckoo is devoid of down and is very dark
all over. According to Hughes it is shining black; Reed describes it
as dark bluish black; and Liversidge records it as brown, changing
within the next two days to purplish brown. The gape at first is
yellowish orange with yellowish edges, but becomes wholly bright
orange by the end of the second day. The bill is dark horn color
with a whitish egg-tooth; the feet are dark flesh color. Liversidge
found a day-old chick to weigh 4.9 grams.
On the fourth day the quill-like, stiff feather sheaths appear on the
upper surface of the wings. The next day these sheaths are 3 mm.
long and show some black pigment basally; on this day sheaths first
appear on the cervical tract. On the sixth day the skin over the eyes
begins to open, but only slightly, hardly enough to enable the bird to
see. On this day the sheaths begin to show up on the chin, forehead,
and sides of the breast, flanks, and abdomen, the longest ones are
twice as long as they were on the fifth day, 6 to 7 mm. By the seventh
day the cuckoo, now more than four times its natal weight, shows the
first sprouting of feathers out of the sheaths on the wings and tail

according to Hughes; Liversidge reports this as taking place on the

eighth day, and notes that the feathers are grayish with a white
terminal bar. On this day sprouting also begins on the sides of the
breast, and by the end of the ninth day this condition extends along
the entire ventral tract.
On the 10th day the eyes fully open, a most important change in
the development and behavior of the chick. The rest of the plumage
breaks from the enclosing sheaths on this day and the next and the
bird suddenly becomes quite fluffy, a marked change in appearance.
390 PROCEEDINGS OF THE NATIONAL MUSEUM vol. loa

On the nth day the bird begms to ruffle its plumage, a type of move-
ment not indulged in hitherto. The feathers, now all out of the
sheaths, become darker, more pronouncedly blackish and less grayish
due to the increased area of their newh^ freed portions. The ventral
edge of the bend of the wing is white, the alula is black, and the under
primary coverts are black with white bars; the feathers of the breast
and sides of the abdomen are hght buff with a black subterminal and
a broad buff terminal bar; the feathers of the midabdomen and thighs
are paler. On the 12th day the rectrices show the contrasting white
tips very markedly because of the growth of the dark proximal portion
of the feathers. By this day, or, in one case, even two days earUer,
the young cuckoo has grown so that it completely fills the nest.
On the loth day the remiges and rectrices are noticeably longer than
the rest of the plumage. From this stage on until the chick leaves
the nest, the changes are only matters of rapid growth. The feet are
now yellow, whereas originally they were flesh colored.
By the time it leaves the nest the cuckoo has grown in weight more
than 11 times its hatching size. Liversidge's bu'd weighed 4.9 grams
at bu'th and nearly 60 grams when it left the nest.
Aside from the purely physical development of the bird, as outlined
above, Liversidge has put on record some highly interesting observa-
tions of the growth and change of its behavior. For the first five
days, as noted in our discussion of the evicting instinct, the chick
reacts to nest-mates (eggs or other chicks) by getting them on its
back and pushing them out of the nest. This reaction is very weak
on the first day, very strong on the second, and normall}^ ceases on
the fourth day, but it can still be evoked by artificial stimulation on
the fifth day. During this period the gaping response of the chick
is usually straight up in direction.
A
second stage begins with the parting of the skin over the eyes.
Gaping is now directed toward the entrance of the nest, i. e., toward
the direction of the hght, rather than merely upwards, and may be
induced by either touching and sUghtly jarring the nest or by a sharp
whistled sound near the nest. Liversidge, however, considers that
this directional gaping is more the result of an originally conditioned
stimulus than of a visually oriented one. The location of the nest
necessitated the same approach by the foster parents when visiting
it, and gaping became directed toward the entrance. It is true,
however, that the gaping did become more directional at the time
the eyes began to open, even though the first day or so they were
not open enough to give the chick really effective or efficient vision.
Both Liversidge and Reed observed the first threatening reactions on
the eighth or ninth days. Reed noted that the chick erected the
feathers of the head and neck and opened the bill, revealing the
 AFRICAN PARASITIC CUCKOOS —FRIEDMANN 391

bright orange gape. On a closer approach, the bird reacted by

darting forward to strike the hand that was extended toward
its bill

it, an action much more aggressive than any feeding behavior. Liver-
sidge found that on the eighth day the eyes were open shghtlj" early
in the morning but were closed again in the evening. It was not
until the 10th day that the bird seemed able to make full and con-
tinuous use of its ej^es; on this daj^ the chick first reacted to move-
mejits of his hand. ma}^ be noted, is a little later than in
This, it

Reed's experience, and ma}' reflect the range of individual variation

in the rate of this stage of the development.
The first noise uttered by the chick, according to Liversidge, was
heard on the 10th da}^, a quiet, chattering note. The next day this
note, now mach louder, was used apparent!}^ only in connection with
begging for food. The original gaping reaction given in response to
touchmg the nest continued up to the 11th da}', but by the evening
of the 12th day such stimulation ojily elicited threatening reactions.
The period from the 15th day of nestling life onward to the time
the chick is fledged and leaves the nest is characterized by a greater
variety of activities. The bird first begins to preen on the 15th day,
at first in a haphazard, apparently nonfunctional way, but rapidly
becoming fuUy functional. On this day the nestling was first seen
tr^-ing to stand up, unsuccessfully at first, but soon becoming better
at it.On the 18th day Liversidge's bird first practiced stretching its
wings, and from then on it quivered its wings whenever begging for
food or being fed. On this da,y, also, it first seemed to distinguish
the call of the foster parents, but showed no ability to differentiate
between the various notes given by tiie robin-chats while at the nest.
Reed observed that when the young cuckoo had left the nest it appar-
ently was able to distinguish the alarm note and to react accordingly.
We still know very httle about the postnestling stage of the young
cuckoo's development, the stage when it becomes separated from the
care of its host and becomes self-sufficient. Reed banded a young
red-chested cuckoo before it left the nest of its foster parents and
found it being fed by them as much as 13 days after it left the nest.
It continued to remain withm a radius of 50 yards of the nest for at
least a month, after which it was seen no more.
Stanford (in htt.) watched a pair of Cape robin-chats feeding a
young red-chested cuckoo some days out of the nest. He first found
the latter by hearing its low-pitched but incessant "tsip-tsip-tsip"
notes. When came close to it, the bird's caUing
the robin-chats
increased in intensity; the young cuckoo bent foi*ward, opened its
bill, partly opened and drooped its ^^dngs and ruffied its plumage.

The call became much louder and was repeated more frequently, the
bill opening and closing spasmodically all the while. Once the bird
392 PROCEEDINGS OF THE NATIONAL MUSEUM vol.106

had been fed it calmed do'uai promptly. He saw it fed eight times in
30 minutes, mostly on what seemed to bo grubs, spiders, and
grasshoppers.
Breeding Range

The first indication that the red-chested cuckoo may breed in

West Africa is a record of a female with a well yolked egg in the
ovary collected on August 6 at Lake Bambalue, near Bamenda,
British Cameroons (Bannerman, 1951, p. 274).

Cuculus cafer Lichtenstcin

Black cuckoo

Since the publication of my earlier account of this rather shy and

secretive cuckoo, additional data, especially on the development of
the nestling and, to a lesser extent, on the evicting habit, have been
recorded. This is principally due to Skead's (1951, pp. 163-168) ob-
servations. In his area in the eastern Cape Province he found the
boubou shrike (Laniarius Jerrugineus) to be the regular, and, as far as
hisown experience went, the only host of the black cuckoo.
The new data, filling what was formerly a complete blank m the
record concerning the growth and development of the young bird, may
be summarized briefly as follows. On the first day after hatching the
bird naked, dark purplish in color, the gape pink, the back noticeably
is

flat (not concave), and the eyes closed. On the fourth day the sheaths
of the rectrices, remiges, and flank feathers begin to emerge; on the
fifth day those of the feathers on the nape, throat, and forehead begin
to appear; by the sixth day the bird has sheaths all over except on the
back which is still nude. The eyes begin to open on the seventh day,
and the bird begins to squeak softly. The feather sheaths begin to
emerge on either side of the bare spinal area on the eighth day. By
the nuith day the eyes are fully open; the back is still largely bare.
The feathers start bursting from their sheaths on the 11th day, but
the primary remiges are still encased in their sheaths. On the 16th
day the bird is well feathered all over, and shows fear reactions by
gaping with the head up and back, not forward as when expecting food;
on the 21st day the bird was gone from the nest and was never seen
again. The nestling period is therefore not more than 21 days. In
the Em-opean cuckoo it is said to vary from 20 to 23 days.
In another nest of the boubou shrike Skead found a newly hatched
black cuckoo and two eggs of the host. The next da}^ one of the shrike's
eggs hatched. The following day the young cuckoo was the sole oc-
cupant of the nest and no sign could be found of the shrike's egg or
chick. The probabihty is that they were ejected by the cuckoo, but
the ejection was not witnessed. The nest, being a rather shallow cup,
 AFRICAN PARASITIC CUCKOOS —FRIEDMANN 393

was such would present no obstacle to ejection by the cuckoo. The

as
inference that the young parasite (definitely identified as a black
is

cuckoo by its plumage when the feathers had grown out) tolerated, or
at least did not eject, its nest-mates for the first 48 hours. It is, of
course, unsafe to draw any hard and fast comparisons between a single
instance in this species with what is Imown in its relatives, the red-
chested cuckoo and the European cuckoo, but it may be pointed out
that the eviction of its nest-mates was delayed in the case of the black
cuckoo for at least 48 hours, as compared with less than 24 hours in a
similar situation involving the red-chested cuckoo, and about the
same in cases of the European cuckoo.

Cuciilus poliocephalus Latham

Lesser cuckoo

Although this cuckoo does not remain in Africa during its breeding
season as far as known, some courtship feeding behavior of interest has
recently been recorded in Northern Rhodesia. Grimwood (in Smith-
ers, 1952, p. 107) watched five of these cuckoos at Danger Hill, 25
miles from Mpika. Of these birds, two "pairs were indulging in what
appeared to be courtship flights, chasing one another from perch to
perch, and on alighting going through a feeding behaviour, though no
food was seen to be transferred." While not surprising, it is of inter-
est to find still another parasitic cuckoo exhibiting this atavistically
revealing behavior pattern.

dialcites cupreus (Shaw)

Emerald cuckoo
Additional data on this bu'd have to do with its host species and with
itscourtship behavior.
Host Records

In 1949 Iquoted a peculiar account (Friedmann, 1949b, pp. 516-

517) of the emerald cuckoo based on observations made on Principe
Island that were attributed to Keuiemans and published by BuUer
(1873, p. 76). Mr. F. Haverschmidt has since sent me a translation
from the Dutch of Keuiemans' (1866, pp. 363-401) observations
on the birds of Principe Island. There is no mention of the queer
story related by Buller, but merely Keuiemans' statement to the
eft'ect that he was told b}'' the inhabitants that this cuckoo deposits

its eggs in the nests of Cuphopterus Dohrni {^Horizorhinus dohrni)

and Parinia leucophaea (=Speirops lucophaea), which breed in the

summer, and that it probably deposits them in the nests of other
species as well.
 —
394 PROCEEDINGS OF THE NATIONAL MUSEUM vol. loe

A much later, and Kttle known, paper by Keulemans (1907, pp.

245-247) gives further welcome details. He writes that the female
emerald cuckoo lays her eggs in the nests of smaller bkds that build
open, cuplike nests, and mentions Zosterops jicedulinus and Speriops
leucophaea as the usual foster parents. Of the latter species of host
he relates a remarkable observation he made in July 1863 on his planta-
tion on Principe Island, "Behind my small dwelling house grew a
large Acacia tree, the branches of which nearly touched the balcony.
A pair of Parinias {=Speirops) had built their nest so near the
verandah that I could hear the 3'oung ones while they were being
fed. One morning the pleasant sound of this young family calling for
food had suddenly ceased. A week later I noticed a young Cuckoo
lifting its head above the nest. Stranger still, two or three days later
a second j'oung cuckoo was to be seen in the same nest, and both these
intruders were fed, not onl}^ by their original foster parents but also
by at least a dozen other small birds." The nest and its contents
came to grief a few days later.
These three hosts in Principe Island Horizorhinus dohmi, Zos-
terops Jicedulinus, —
and Speirops leucophaea are all additional to our
previous list. It is unfortunate that no details of actual instances are
available for either of the first two. While there is nothing improbable
in their being parasitized by the emerald cuckoo, Keulemans' unsup-
ported statement does not constitute evidence in itself, and it cannot
be claimed that bh'ds of the first two species are known definitely
as fosterers of the emerald cuckoo.
It may be pointed out that the Speirops case described above gives
us the first evidence that more than one emerald cuckoo egg may be
laid in one nest. We have no way of knowing if the two eggs were
the product of a single bird or of two different individuals,
Winterbottom (1951, p, 27) hsts the puff-backed shi-ike (Dryoscopus
cuhla) as a host of the emerald cuckoo but he gives no fm-ther data,
Mackworth-Praed and Grant (1952, p, 509) also list puff-backed
shrikes, together with bulbuls, as the usual hosts of this cuckoo.
However, I am aware of but a single specific instance of the puff-back
in this capacity (J. Vincent, 1934, p. 761; a record overlooked before).
The puff-back shrike is known better as a victim of the black cuckoo.
However, its ecological habitat is much sylvan than that
less strictly
of the emerald cuckoo, being more like that of the
black cuckoo.
On the other hand, hosts additional to those previously hsted are
indicated in the following observations, Guichard (1950, p. 168)
found the Abyssinian bulbul (Pycnonotus harbatus schoanus) to be
victimized at Addis Ababa, Ethiopia, while in Nyasaland Benson
(1953, p, 35) records eggs of the emerald cuckoo in nests of Chalomitra
senegalensis gutturalis and Cyanomitra olivacea alfredi. Benson also
 AFRICAN PARASITIC CUCKOOS —FRIEDMANN 395

lists (p. 113) a nest of the paradise llycatclicr (Terpsiphone viridis)

with two eggs of its own and one egg of the emerald cuckoo found at
Blantyre, Nyasaland, on November 26. This flycatcher was previ-
ously known as a host from a single instance in the Belgian Congo.
The subspecies in Nyasaland is T. v. granti, while the Congo bird is

T. V. viridis. The crimson-crested black weaver (Malimbus

forest
malimbicus crassirostris) is added to the list of the known victims on
the following basis. The van Somerens (1949, p. 95) record an egg
(identified as an emerald cuckoo's by Pitman) from a nest of this forest
weaver in the Bwamba area, Toro, western Uganda, An egg of the
weaver, punctured and empty, lay on the ground below the nest.
One other bird not definitely known to be parasitized hj the emerald
cuckoo has been reported in a way suggestive of such a relationship.
Holliday and Tait (1953, p. IIG) watched a nesting pair of olive
barbets {Buccanodon olivaceum woodwardi) at Ngoye Forest, Zulidand.
Once, when the barbets had gone foraging for food for then- nesthngs,
an emerald cuckoo "approached the nest, made certain that the adult
barbets were nowhere in the vicinity, and disappeared from view into
the nest hole. All the time this was going on the young birds kept
up their continuous clamour. Whilst the intruder was still within
the nest, one of the parents arrived on the scene, and finding the un-
wanted visitor, attacked it fiercely and drove it away from the nest
site. Owing to the fact that the nest was inaccessible, an examination
of the nesthngs was not possible, but one might assume that one of
the chicks was probably a young cuckoo, and the parent was taking
an active interest in its welfare, for it is not likely that it would be
looking for a suitable nest in which to lay its eggs at this stage. As
little is known about the breeding habits of Emerald Cuckoos, it is

felt that this record is of some value, for some observers have seen it
frequenting the nests of other species of barbets during the breeding
season." The assumption that the adult cuckoo came to the nest
because of a presumed interest in the welfare of an equally hypothetical
nestling of its own kind has little enough to support it, but the fact
that the cuckoo showed some interest in the nest suggests that such a
site might be within the range of its potential choice of a receptacle
for its eggs. Hitherto no hole-nesting bird as been found to be para-
sitizedby the emerald cuckoo, or, for that matter, by any of the
African metallic cuckoos of the genus Chalcites.

CoTJRTsniP Behavior

When I wi'ote my
account of this cuckoo nothing was known
earlier
of its com-tship behavior.Since then this gap has been partly fiUed
by the interesting and valuable observations recorded by Haydock
(1950, p. 150). He saw a female emerald cuckoo perched on a bare
396 PROCEEDINGS OF THE NATIONAL MUSEUM vol. loe

branch of a defoliated A male was on a higher branch, calling

tree.
loudly; it flew down hen
to theand, with wings drooping and tail
erect, bobbed in front of her, and presented her with a large hairy
caterpillar. This offering was accepted and devoured, the cock bird
calling loudly with its head thrown v^^ell back during this time. A
second similar display was then gone through, after which coition was
attempted unsuccessfully. Shortly after this both birds flew away.
Haydock's observations were made in central Northern Rhodesia in
the third week in January.
In my account of this cuckoo, mention was made of the fact
earlier
that male birds seemed to be "territorial" and to establish singing
posts to which they adhered day after day. These observations I
made in western Kenya Colony. Welcome corrobation by virtue of
similar observations now is available from the Upemba Park, Belgian
Congo, where Verheyen (1953, pp. 319-320) found the same situation.
He goes further and writes that the males reserve a w^ell defined area
which they defend against the intrusion of other males. This is,
however, an unsupported statement as he does not describe any
actual instances of territorial defense, and all I knew previously was
merely the fact that males sang persistently from their singing posts.
In Principe and San Thome Islands, Keulemans (1907) found this
cuckoo to be migratory, being present on the islands only from Feb-
ruary until November.

Chalcites klaas (Stephens)

Klaas's cuckoo

Host Species

All the new information on cuckoo refers merely to additional

this
host records, the total of which may now
be raised from the 33 species
listed in my earlier accounts to 42 species, or, if we count species and
subspecies, to 50 forms. Of the total of all instances of this bird's
parasitism known to me, 30 percent are with sunbirds as hosts, 23
percent with flycatchers, 16 percent with weavers, and the rest single
or, at most, two records for wagtails, bulbuls, babblers, starlings,
white-ej^es, and finches. While 30 percent of the parasitized nests
are those of sunbirds, the 10 species of this family recorded as hosts
of the cuckoo amount to not quite 25 percent of the total known
species of victims. Similarly, seven species of flycatchers, or about
16 percent of the total of 42 kinds of hosts, account for almost 25
percent of the recorded instances.
The arrow-marked babbler {Turdoides jardinei) is listed as a host
of Klaas's cuckoo by Winterbottom (1951, p. 15), but no specific
instances are given. I am not aware of any published pertinent data.
 ArEICA>: PARASITIC CUCKOOS —FRIEDMANN 397

Another new host is the dusky flycatcher {Alseonax adusius), a pan-

of which were seen feeding a young Klaas's cuckoo on December 6 in
Hottentot's Holland, Cape Pro%'ince, by MacLeod and Murray (1952.
p. 22). Still another species of Alseonax, the Uganda pygmy dusky
flycatcher {A. minimum jpumilus), may be added to the list, this one
on the basis of three records. Williams (1946, p. 138) found a nest
of this bhd at Kampala, Uganda, on Apr. 27, 1946, that contamed
two fresh eggs of the builder and one very slightly incubated egg of
the cuckoo. At Kitale, Kenya Colony, Stoneham (1952, p. 7) found
two nestling Klaas's cuckoos, one in each of two nests of this fly-
catcher, near the garden of the museum eai'ly in 1952.
According to Benson (1953, p. 35), the gray tit-babbler (Parisoma
plumbeum orientale) has been found to be parasitized in Nyasaland; he
gives no further data on it, and Hsts it as an '"apparent" host of Klaas's
cuckoo.
In my book (Friedmann, 1949a, p. 140) I listed the Cape flycatcher
{Batis capensis) as a host of this cuckoo solely on Joubert's statement
that it was victimized in the Cape Province. Since then MacLeod
and Murray (1952, p. 17) have recorded two instances in the Hotten-
tot's Holland area, and Stanford (in htt.) sends me the following
additional cases. On December
6 at Picnic Bush, Somerset West,
Cape Province, he and G. ^MacLeod found a young Klaas's cuckoo,
J.
not quite ready to fly, completely fiihng a nest of a Cape flvcatcher.
Within 30 yards of this nest was another nest of the same species that
had had a young Klaas's cuckoo in it a day or two earlier. In this
area, in the years 1951-1952, Stanford and MacLeod found a total of
five instances of the Cape flycatcher serving as host for Klaas's
cuckoo.
To the previous meager data on the Cape paradise fl^^catcher
(Terpsiphone perspiciUata) may be added another instance, recorded
at Hottentot's Hofland by 2\IacLeod and Murray (1952). The related
Terpsiphone viridis is a bird of which two races, the nominate one and
suahehca, have been previously listed as victims of Klaas's cuckoo. It
seems that the race spcciosa may serve in this capacity as weU. Chapin
(1953, p. 725) mentions a nest at Avakubi that contained one egg of the
builder and one egg that probably was of this cuckoo.
The rufescent swamp warbler (Calamocichia refcsccns) is a bird not
hitherto recorded as a victim of Klaas's cuckoo. Chapin writes me
that near Tshibati, Belgian Congo, on Feb. 27, 1954, he found a nest
of this bird high up on a fork of a stalk of elephant grass that con-
tained a weU-feathered nesthng Klaas's cuckoo. The foster pai-ents
were scolding^nearb}' as he examined the nest. The race of the host
in the eastern Congo is probablj'/oxi.
398 PROCEEDINGS OF THE NATIONAL MUSEUM vol. loe

At the time of my first report (Friedmann, 1949a, p. 141) I knew of

only two instances of the bar-throated warbler (Apalis thoracicus
thomcicus) being parasitized. This numl^cr may now be increased as
a result of observations by Skead (1952, p. 12), who found one such
case in December 1946 and two more in January 1947.
The crombec {Sylmetta rujescens transvaalensis) was
long-billed
previously known on the basis of a single
as a victim of this cuckoo
case. To this may be added another quite unusual one. Mr. O. P. M.
Prozesky informed me when I was in the Transvaal that on Dec. 27,
1950, at Buffelspoort, 50 miles northwest of Pretoria, he found a pair of
crombecs feeding two fledgling Klaas's cuckoos.
Another host previously known from one record and for which a
second has now come to my attention is the gray-backed bush warbler
(Camaropfera brevicaudata abessinica). Sir Charles F, Belcher Avrites
me that he found a parasitized nest of this bird in Kenya Colony.
The warbler had deserted it.
The tawn3'--flanked longtail, Prinia mistacea (subsp. melanor-
hyncha), was knovv^ to be victimized in the Gold Coast. Its eastern
race, immutabilis, has since been found to be similarly affected in
Kenya Colony, where Stoneham (1952, p. 7) found a nest at Kitale
with three eggs of the longtail and one egg of Klaas's cuckoo.
Skead's (1954, p. 87) recent study of the greater double-collared
sunbird has added six more instances of its usage as a victim by
Klaas's cuckoo. It must be regarded as one of the chief hosts of the
parasite in theCape Province.
The East African mouse-colored sunbird (Chaleomitra veroxii
jischeri) has been found to act as a host for Klaas's cuckoo in Kenj^a
Colony by Belcher (in litt.). I had previously listed it as such in
northern Zululand, but it now seems that the latter record refers to
the nominate race of the sunbird.
Cinnyris venustus jalkensteini may be added to the known hosts of
Klaas's cuckoo, as Mackworth-Praed and Grant (1955, p. 779) record
itas being occasionally parasitized in East Africa. Previously I had
known of only a single instance, involving the Abyssinian race Cinnyris
venustus fazoqlensis (Friedmann, 1949a, p. 143).
The Kenya rufous sparrow (Passer iagoensis rufocinctus) is an

addition to the recorded victims of Klaas's cuckoo. Belcher (1949b,

p. 19)mentions, without further details, that he has in his collection
eggs of Klaas's cuckoo taken from a nest of this sparrow. I under-
stand from correspondence that there were two eggs attributed to the
cuckoo in this nest and that the identification of the parasite is not
really definite.
Chapin (1954, p. 339) records the masked weaver (P/oce^(5 inter-
medius intermedius) as a host of Klaas's cuckoo on the strength of two
 AFRICAN PARASITIC CUCKOOS —FRIEDMANN 399

parasitized nests found in Uganda by Pitman. Each contained one

egg of the weaver and one egg of the cuckoo.
The southern race of this weaver, Ploceus intermedius cabanisi, has
also been found to be parasitized by Klaas's cuckoo in Southern
Rhodesia, according to Mr. H. M. Miles. It had been recorded
previously only as a host of the didric.
Layard's spot-backed weaver (Ploceus nigriceps) was described as
a victim of Klaas's cuckoo on the basis of two such instances at Taveta,
Kenya Colony (Friedmann, 1949a, p. 146). Farther south, at Mikin-
dani in extreme southern Tanganyika Territory, it also appears to
be a frequent victim, Grote (1924, p. 34) considers it a bird often
imposed upon there by both this cuckoo and the related didric.
Masterson (1953, p. 51) recentl}^ added the red-headed weaver
(Anaplectes rubriceps) to the known victims of this cuckoo. On
Nov. 6, 1952, at Murambinda, Buhera District, Southern Rhodesia,
he found a nest of this weaver that contained three eggs of its own
and one egg that was presumed to be of a Klaas's cuckoo. The latter
egg was freckled and was paler and slightly narrower than the host's
eggs. A Klaas's cuckoo was calling excitedly in the vicinity and
three times it tried to enter the nest in spite of the nearness of Master-
son's natives who had climbed to the nest. Later, in January 1953,
Masterson saw a pair of these weavers feeding a young lOaas's cuckoo
in his garden.
The red bishop bird (Eupledes orix), a frequent host of the didric,
also has been listed as an apparent host of lOaas's cuckoo in Nyasa-
land (Benson 1953, p. 35), but without further supporting details.

Eggs and Egg Laying

Recently MacLeod and Hallack (1956, pp. 2-5) described four eggs
of this cuckoo — all laid in nests of the Cape flycatcher (Batis capen-
sis) —as pinkish white and heavily spotted with reddish blotches
forming a fairly definite ring around the blunt ends. No pinkish
white eggs of this cuckoo were previously recorded. These authors
think (but do not know) that all four eggs were laid by the sam.e hen
cuckoo. This is based on the similarity of the eggs, on the fact that
all were found in nests within a limited area, and that all were found

in nests of the same species of host. The evidence, if such it may be

called, is merely suggestive, but, aside from adding another color type
to the known eggs of Klaas's cuckoo, it is the first indication of the
number of eggs laid and of host specificity in this species.
Eviction of Nest-Mates

In my earlier account I stated that while the young Klaas's cuckoo

is usually the sole survivor in a nest, no information was available as
to whether it evicts the other young or eggs in the nest or starves them
400 PROCEEDINGS OF THE NATIONAL JMUSEUM vol. loe

out in competition for the food brought to the nest by the hosts.
While the present paper was awaiting publication, MacLeod and
Hallack (1956) supplied our first definite observations on this point.
They watched a parasitized nest of a Cape flycatcher. The Klaas's
cuckoo hatched on November 9, and one of the host's eggs hatched
on November 13. On November 14 the J^oung cuckoo evicted the
3^oung flycaU'her, At 9 a, m. on that day the cuckoo was noted
underneath the young flycatcher, trjang to heave it out of the nest;
this attempt was unsuccessful, and was repeated at 9:25, again un-
successfully. At 11:42 another attempt succeeded. The young Batis
was left dangling by one foot, head down, from the outside of the
nest. The adult Batis returned with food, took no notice of its right-
ful young, and fed the young parasite. A quarter of a hour later the
young flj^catcher fell to the rocks below.

Chalcites flaviguUtris (Shelley)

Yellow-throated cuckoo

This rare cuckoo remains still virtually unobserved. Its range

may now be extended a short distance eastward to the Bwamba area,
Toro, western Uganda, where the van Somerens (1949, p. 29) collected
a breeding female at Ntandi in July and where Ridley, Percy, and
Percy (1953, p. 163) obtained another female in full breeding con-
dition on September 27 near Hakitengya. The stomacli contents of
both birds consisted of hairy cateipillars, plus scale insects in the
July bird and beetles in the September one.

Chalcites caprius (Boddaert)

Didric cuckoo

The new information on this cuckoo may be treated under several

subdivisions.
Host Records

A considerable number of species of birds not listed in my earlier

report have now been found to be parasitized by the didric cuckoo,
bringing the total of its known hosts to 46 species, or55 species and
subspecies of birds. In addition, our knowledge of some that were
included in the 1949 list has been considerably augmented. It be-
comes increasingly clear that the Ploceidae are the chief victims of the
didric cuckoo; approximately three-fourths of all the nests recorded
with eggs or j^oung of the cuckoo belong to birds of this family.
The Cape wagtail (Motacilla capensis capensis) was included in my
(1949a) book on the basis of one, not wholly satisfactory, record.
However, an overlooked statement by Nicholson (1897, pp. 142-143)
 AFRICAN PARASITIC CUCKOOS —FRIEDMANN 401

reveals that in his garden at Pretoria a didric cuckoo layed an egg in a

nest of the Cape wagtail and tlie egg was hatched and the young
parasite reared by the host. Recently, Pike (in litt.) saw one of
these wagtails feeding a fledghng didric on January 8 in the Transkei
District, Cape Province. Skcad (1952, p. 4) lists three more instances
in the Capo Province. The Cape wagtail must be considered as a
regular host of this cuckoo.
Winterbottom (1951, p. 15) wiites that the didric is parasitic on the
arrow-marked babbler {Turdoides jardinei). 1 know of no definite
instance, although it may be recalled that Swynnerton (1911, p. 19)
once shot a didric as it was leaving a nest of this babbler. There was
no cuckoo's egg in the nest, however.
The small-billed brown flycatcher (Bradornis imcrorhynchus) is a
new host of the didric. Belcher (in litt.) informs me that he collected
an egg of this cuckoo from a nest of this bird near Ngong, Kenya
Colony. He felt satisfied as to the identity of the species of cuckoo
involved.
The Shoa olive sunbird (Cyanomitra olivacea ragazzii) is a possible
addition to our previous list of hosts. Reichenow (1881, p. 16)
mentions a didric's egg found by G. A. Fischer in a nest of this sunbird.
This egg is said to have been given to the Nehrkom Collection, but
the catalog of that collection makes no mention of it. It is not
stated how the identification of the Qgg was decided; it may have been
a Klaas's cuckoo.
Skead (1952, p. 5) has tentatively listed the yellow-throated
sparrow (Petronia superciliaris) as a victim of the didric cuckoo, but
he states that the parasitic egg seen in the one nest involved may have
been that of a greater honey-guide. I think the latter is probably
correct; the record should be deleted from the list of didric hosts.
Vieiliot's black weaver (Ploceus nigerrimus) was known previously
to be parasitized in Uganda and in the Belgian Congo. To this may
be added that Serle (1954, p. 55) found a fledghng didric cuckoo,
unable as yet to fly, beneath a nesting colony of these black weavers
on January 24 at Kumba, British Cameroons.
The black-headed weaver {Ploceus 7nelanocephalus) is a species new
to our list of victims, but it was actually recorded in this capacity as
long ago as 1899 by Louis Petit (1899, pp. 66-67) in the French Congo.
The race of the weaver in that area is P. m. cajntalis (Latham).
Petit writes that the didric is parasitic on this weaver, and that the
latter rears the young parasites.
Two other races of the black-headed weaver may also be mentioned
in the present connection. Indefinite observations by Verheyen
(1953, p. 315) suggest that Dubois' black-headed weaver {Ploceus
402 PROCEEDINGS OF THE NATIONAL MUSEUM vol. loe

melanocephalus duboisi) may be parasitized by the didric in the

Bcls^an Congo.
The Uganda yclIow-coUared weaver (Ploceus melanocephalus
dimidiaius) has been recorded, -without supporting data, as a victim of
the didric cuckoo. Distant (1897, p. 143) states that Jackson found
an Qgg "considered to be of this cuckoo" in a nest of this weaver.
To the meager data previously available on Layard's spot-backed
weaver {Ploceus nigriceps) as a cUdric victim in southern Tanganjdka
Territorj' may be added that Benson (1953, p. 35) found it to be
parasitized in Nyasaland as well.
In ray earlier account (1949a, p. 173) it was noted that Cabanis'
masked weaver (Ploceus intermedius cahanisi) had been reported as a
fosterer of the didric in South Africa by Roberts. Recently Lamm
(1955, p. 34) wTOte that this weaver appears to be the principal host
of this cuckoo in southern Mozambique. He also suggested that the
spotted-backed weaver {Ploceus spilonotis spilonotus) is probably a
frequent victim of the didric in that region. Previous data on this
bird as a host were all from the Union of South Africa.
I have recorded (Friedmann, 1949a, p. 175) the spectacled weaver
{Ploceus ocularius ocularius) as not too certainly established as a
Icnown host of the didric cuckoo. A definite instance has smce been
recorded by Skead (1952, p. 9) at King WDliam's Town, Cape Province.
Previously (Friedmann, 1949a, p. 176), I included Smith's golden
weaver {Ploceus suhaureus subaureus) as a victim of the didric only on
the strength of its being so listed by Patterson and by Sparrow, since
no specific instances were loiow^n to me at the time. Since then,
D. C. H. Plowes has infonned me that at Umkomaas, Natal, on Jan. 2,
1949, he collected a nest of this weaver containing two eggs of the host
and one of the didric cuckoo.
New to our earher list of victims is the Zambesi brown-throated
weaver {Ploceus xanthopterus) . A young didric cuckoo w^as found in a
nest of this bird at Karonga, Nyasaland, on February 25 (Benson and
Benson, 1949, p. 165).
Also new to the roster of the didric's hosts is the red-winged ana-
plectes (Anaplectes rubiginosus) listed in this capacity in Nyasaland by
Benson (1953, p. 35).
The red bishop bird {Euplectes orix) was recorded earlier as a
frequent victim, but how frequently it suffers from the attention of
the parasite has recently been made clear by Reed (1953, pp. 138-140),
who found no fewer than 23 parasitized nests in the Transvaal.
The Zanzibar red bishop bird {Euplectes nigroventris) may be a host
that previously was overlooked. Fischer (1880, p, 190) found a nest
of this weaver with a live female didric caught entangled on it. The
 AFRICAN PARASITIC CUCKOOS —FRIEDMANN 403

nest contained three eggs, of which one was smashed and the others
seemed to be Euplectes eggs. It must be cautioned that there is no
proof of parasitism in this case; the cuckoo may have come to lay
there or to eat one of the eggs in the nest.
The lire-crowned bishop bu*d (Euplectes hordeacea sylvatica) does
not appear to have been recorded as a victim of the didric cuckoo
until Benson (1953, p. 35) listed it as a host of this parasite in Nyasa-
land, but ^\^thout further details. In the Upemba Park, Belgian
Congo, Verheyen (1953, p. 315) saw a hen didric entering a colony of
the nominate race of this bishop bhd but he did not obtain evidence
of any actual parasitism.
The white-winged whydah {ColUispasser albonotatus albonotatus) is

suspected of being a host of the didric in Southern Rhodesia by

Edwards, according to Miles (1951, p. 4), but no definite evidence
seems to be known.
The golden-breasted bunting {Emheriza jiaviventris) has been
definitel}" added to the known victims of this cuckoo by Haj^dock

(1950, p. 150; 1951, p. 3) in Northern Rhodesia. He observed

nesthng didrics, one in each of tlu'ee nests of this bunting, as well as
three young didrics already out of the nest apparently attended hy
the buntings.
When discussing the Cape rock bunting {Fringillaria capensis) as a
host of the didric, I mentioned (Friedmann, 1949a, p. 177) that
although several ^vriters had listed it as such, no localities or other
particulars were available. It seems that these authors were copying
from one another, and that the original source of them all was Mrs.
Barber, who apparently made her observations in the Transvaal.
Sharpe's edition of Layard's "Birds of South Africa" (1875, p. 155)
appears to contain the first mention of ^Irs. Barber's data.

Egg Laying
In my earlier report I stated that there is good evidence to the effect
that the didric hen often, if not regularly, may remove an egg from
the nest when lading its own into it. Skead (1952, p. 9) came to a
similar conclusion as but Reed (1953, pp. 138-140) concluded
vrell,

that the opposite seemed to be the case. Reed studied the didric's
parasitism on the red bishop bird, and found that the cuckoo "does
not appear to remove an egg of the host the normal Red
. . . . . .

Bishop clutch is three eggs and nearh^ all nests containing cuckoo eggs
carried a total of four eggs." While he gives data on some 23 instances
in which the red bishop bird was parasitized by the didric, most of
these cases were of nests mth j^oung cuckoos; in only six nests does
he record four eggs each (thi'ee bishop bu-d eggs and one didric egg).
404 PROCEEDINGS OF THE NATIONAL MUSEUM vol. loe

Inasmuch as this matter of removing an egg often involves the eating

of it by the didric, it may be noted that Caldcr (1951, p. 36) watched
a hen didric enter a nest of a thick-billed weaver {Amblyospiza
albifrons), emerge with an egg in its bill, and then eat the contents.
No cuckoo's egg was in the nest when it was examined a iew minutes
later, but it would seem that this weaver may have been intended as
a fosterer. It would not be surprising if someone were to find it to
act in this capacity.
In the disputed matter of adaptive similarity between the eggs of
the cuckoo and those of its victims, the recently amassed data are just

as conflicting as was the older information. Reed noted a definite

simila^rity between didric eggs in nests of the red bishop bh'd and
those of the host, while Skead found noticeable contrast between didric
eggs and those of the builders in nests of Cape wagtails and Cape
weavers.
Evicting Habit

The evicting habit in this cuckoo still presents unsolved problems,

but the following data, subsequent to my earlier account (Friedmann,
1949a, pp. 179-181), are of interest. No one has yet seen a didric
chick in the act of ousting either eggs or young from the nest. The
evidence from which eviction is assumed to have taken place is merely
the fact that eventually the parasite is usually the sole occupant of
the nest. Assuming that the young cuckoo is responsible for the
disappearance of the other nest contents, Skead (1952, pp. 7, 9) found
that a nestling didric in a Cape wagtail's nest "tolerated" the two
eggs of the host for two days after it hatched, when one egg was found
below the nest and the other was out of the nest early the next
morning (i. e., after 2}^ days). In a Cape weaver's nest there were
two weaver's eggs when the didric hatched; the next day one of the
weaver's eggs hatched, but the following day both it and the unhatched
egg were gone, leaving the didric as sole occupant. In a spectacled
weaver's nest there was one egg of the host when a didric hatched;
both remained in the nest for two daj^s, after which the cuckoo was the
only occupant. From these three examples it appears that 2 or 2)^
days may be the usual time for the "eviction" (or disappearance from
the nest) of the cuckoo's nest-mates. However, this is by no means
universal, as Reed (cit. supra) found in the case of a parasitized nest
of the red bishop bird. When the didric hatched there were two of
the host's eggs and one of its chicks in the nest, while four days later
the nest contained three chicks of the bishop bird and the young
cuckoo. Unfortunately, Reed's observations ended at that point.
However, in another nest of the same host species he found a didric
about four days old as the sole occupant. At 1 :30 p. m. he placed
two eggs of the bishop bird in the nest; by 6 p. m. (4^^ hours later)