PROCEEDINGS OF THE UNITED STATES NATIONAL MUSEUM
issued il^i'\A.Sl'^M>il ^H '^*
SMITHSONIAN INSTITUTIONU. 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 Africanparasitic cuckoos from the basic facts presented in my book entitled"The Parasitic Cuckoos of Africa" published in 1949 and the littleaddendum to it (Friedmann, 1949b) published later in the same year.In the present account only those data pertinent to the problems ofparasitic reproduction are included. For this reason, some speciesof cuckoos are not discussed at all as there were no new facts todescribe. Even in those that are treated in this paper almost nospace is given to such items as call-notes, migration, food of the adult,plumages, etc.For sending me unpublished information, for answering manyqueries, and for other acts of kind cooperation, I am indebted to thefollowing naturalists: C. F. Belcher, J. P. Chapin, F.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 weresupplied by Mr. Liversidge in addition to his information. Hisobservations on the red-chested cuckoo (Liversidge, 1955), unpub-lished when sent to me, fortunately appeared in print just as thepresent manuscript was about to leave my hands. I have thereforemerely summarized his data insofar as integrating all the informationhas permitted.386753?56- 377
378 PROCEEDINGS OF THE NATIONAL MUSEUM vol. loeClamator glandarius (Linnaeus)Great-spotted cuckooThree new hosts may be added to those listed m my book, one ofwhicli is onl}^ a race of a species of which another form was alreadyrecorded as a victim of the great-spotted cuckoo. The first of theseis the dwarf raven (Corvus corax edilhae), a bird of which Belcher(1949a, p. 37) records a nest found near Gabredarre, Ogaden, ItalianSomaliland, containing tliree or four eggs of the great-spotted cuckooand three or four eggs as well as one young of the host. Previouslyonly the Spanish race of this raven (C. c. hispanus) had been reportedas a host of the nominate race of the cuckoo.The second is the piapiac (Ptilostomus afer). I have no data onthis other than that it is mentioned as a victim of the great-spottedcuckoo by Mackworth-Praed and Grant (1952, p. 505).The third new host is the long-tailed glossy starling {Lamprotorniscaudatus). Madden (1934, p. 94) saw a fledgling great-spotted cuckooapparently being fed by a pair of these starlings at El Obeid, Kordo-fan, Sudan, in November 1932. Aside from adding an additionalspecies to the known hosts of this parasite, this record seems to bethe first for any hole-nesting bird north of South Africa. As pointedout in my book (Friedmann, 1949a, p. 10), the data then availableshowed a puzzling divergence in the choice of victims by this cuckooin South Africa on the one hand, and ever3''where else in its range onthe other. In the former area most of the records were of hole-nestingbirds (three species of starlings), while all the records from Nyasalandand Southern Khodesia north to the Mediterranean lands were of birdsmaking open nests in trees or on ledges (crows and magpies of severalspecies). It is true that one open-nest builder, the Cape rook, wasknown to be parasitized in South Africa; now we may further reducethe 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. Sincethen, Clancey (1951, p. 141) has separated the birds south of theSahara under the name choragium, restricting the name glandariusto the birds that breed in Spain, Portugal, Mediterranean Africa,Cyprus, Greece, Asia Minor, and Iran and that migrate south inwinter to tropical Africa. It follows from this that there again seemsto be a regional difference in host selection. The nominate race isknown to utilize only open nests, chiefly of corvids of several kinds,whereas the African choragium lays in such nests and in those ofhole-nesting starlings as well.
AFRICAN PARASITIC CUCKOOS?FRIEDMANN 379Clamator jacobinus (Boddaert)Jacobin cuckooNew information gathered on the jacobin cuckoo does more thanmerely yield additional instances of its parasitism on previouslyrecorded hosts. Many additional records of its eggs in nests ofLayard's bulbuls (Pycnonotus tricolor layardi), sombre bulbuls(xindropadus imjmrlunus im/portunus) , and fiscal shrikes (Laniuscollaris collaris) corroborate the predominance these hosts play inthe economy of the parasite, but, merely as records, they add noreveahng new data. Two additional instances of the bakbakiri(Telephonus zeylonus) added to the three listed in my book suggestthat this shrike is more frequentlj^ victimized than was formerlysuspected. These instances are: (1) a nest found at Butterworth,Cape Province, by Pike on February 4 with one egg of the host andone 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 paradiseflycatcher (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 setof two eggs of the grassbu'd {Sphenoeacus ojer transvaalensis) withone 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 anew host but also is the only instance kno\\ai to me of the parasiteusing a nest built close to the ground. The one record of the jacobinparasitizing 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 thisparasite I raised some doubts as to two cases recorded by de Klerkof the jacobin cuckoo laying its eggs in nests of the yellow-throatedsparrow (Petronia superciliaris) and suggested that the records mightrefer to honey-guides. However, de Klerk's measurements are toolarge for any hone^^-guide's eggs and agree very closely v/ith other,authentic eggs of the jacobin cuckoo. The records must thereforebe 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 otherwiseknown to be utilized by the jacobin cuckoo.Incubation PeriodThe incubation period is still to be determined, but Skead's (1951,pp. 171-172) incomplete evidence suggests a shorter period thantdidmy own similarly fractional data (Friedmann, 1949a,\p. 37)._^It may
380 PROCEEDINGS OF THE NATIONAL MUSEUM vol. loebe recalled that the nest I was watching was destroyed 14 days afterthe last host's egg was laid and the smashed cuckoo's egg in the nestseemed then about in hatching condition. One of Skead's casessuggested a possible incubation period of 12)^ days, while his secondone seemed three days less, and caused him to ask if the cuckoo'segg might begin its development some time prior to ovulation. Itwould seem unlikely that the incubation period is under 12^ days.Evicting HabitOur knowledge of the evicting habit in the jacobin cuckoo stiUrequires clarification. In 1949 (Friedmann, 1949a, p. 37) I was awareof no real evidence and could only conclude that while the youngcuckoo was almost always the sole survivor of a nest, the absence ofany of the rightful young might be the result of their failure to competefor food with their parasitic nest-mate rather than that they wereactually evicted alive by it. Skead (1951, pp. 172-173) has describedan instance in which the nestling jacobin tolerated eggs and youngin the nest for up to four days, and another in which it made noattempt to evict eggs for four days but in which a chick of the hostwas found to disappear within less than a da}^ after hatching (butnot necessarily evicted by the cuckoo). In the first case, Skeadconmients that, considering how the young cuckoo covered and almostsuffocated its nest-mate (a Layard's bulbul), the "possibility of deathof nestlings b}^ this means must not be ruled out. The chick was soweak . . . when retrieved . . . that had it remained there muchlonger, it would have been smothered. Therefore, I wish to drawattention to the suffocation of cuckoo's nest-mates and the possibilityof 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 ofinterest. At Butterworth, Transkei, Cape Province, he found a nestof a fiscal shrike on January 15 containing only one egg?a jacobincuckoo's. On February 27 the nest was revisited and was found tohold a young jacobin and three j^oung shrikes. Still later he sawthe four young birds perched on twigs near the nest, and still laterhe noted that the young shrikes had left but the cuckoo was stillbeing fed by an adult shrike. This is the only definite instanceknown to me of the nest-mates surviving together with a jacobincuckoo. From this, and from the two instances described b}' Skead,in which eviction by the cuckoo, if any took place, did not occur forsome days after hatching, it follows that evicting is certainly notinvariable or immediate, and, for that matter, it is still not possibleto say that the young parasite was responsible for any evictions.
AFRICAN PARASITIC CUCKOOS?FRIEDMANN 381Actually, this is not surprising when we consider that in the relatedgreat-spotted cuckoo, the rightful 3"ouug often, and perhaps evenregularly, grow up together with the parasite. Furthermore, twoj^oung jacobin cuckoos have been noted as growing up together withone of the young of a babbler host (Argya rnalcomli) in India, a furtherindication of the lack of evicting behavior (Bates, 1938, p. 125).Egg LayingPike (again cited in Godfrey, 1939) found, on December 4, anothernest of the fiscal slmke containing one egg of the jacobin cuckoo; twodays later it contained two shrike's eggs in addition to the cuckoo's,and three days later still it held five eggs of the slniko and the oneegg of the jacobin. Pike was unable to visit the nest again. It wouldappear from this case and the one referred to previously that thecuckoo may ia}'^ occasionally into nests before the builder has begunto lay. It is, of course, not impossible that in each instance theremay have been a single shrike's egg present and that the cuckooremoved it Vv^hen laying its own. It is, however, not very likely thatthis was the case, as the usual number of eggs m a clutch of the shrilveis four or five, and in these cases there was no reduction as wouldhave been the case had the cuckoo removed an egg. In my earlieraccount of this parasite (Friedmann, 1949a, p. 30) I summarized thethen available information on this point, and I have seen no evidenceto cause me to change it since: "No one has witnessed the actualdeposition of the egg, but judging by the fact that the number ofeggs of the host is usually less than the fuU complement by the numberof cuckoos' eggs in the nest, it seems that the cuckoo usually removesan egg before laying into the nest . . . ." Of course, if?as seems tohave been the case in the two instances described above?the cuckoo'segg is laid first there can be no such removal of a host's egg.An observation suggesting that the jacobin cuckoo may at timesbe an egg-eating nest robber is the following sent me by Pike, whoonce saw some red bishop birds (Euplectes orix) chasing one of thesecuckoos away from their nests. Two of the nests each had a hole ofabout an inch and a half m diameter torn in the lower part; one ofthe nests contained two eggs of the builder, one being badly peckedand empty and the other also broken; the bottom of the nest wasmoist with spilled yolk.In my earlier account of this cuckoo (Friedmann, 1949a, p. 30) itM^as mentioned that only a single instance was known in Africa ofmore than two eggs of this bird in one nest, although in India StuartBaker had recorded six nests with three eggs of the parasite apiece,two nests with four, and one nest with six eggs attributable to thejacobin cuckoo. The lone African instance was a nest of the sombre
382 PROCEEDINGS OF THE NATIONAL IVIUSEUM vol. loebulbul (Andropadus imjportunus) with five eggs of the cuckoo andtwo of the host, reported by Ivy (1901, pp. 23-24). It was to beexpected that the situation in the African population of this cuckoowould prove to be similar to that in the Asiatic part, and this hasrecently been found to be the case. Air. H. M. Miles of Salisbury,Southern Rhodesia, informs me (in litt.) that nearby, at PlumtreeSchool, on Mar. 2, 1954, Mr. Ian Canncll found a nest of Layard'sbulbul containing seven eggs of the cuckoo and one broken egg ofthe bulbul. He also found another nest of the same host with foureggs of the cuckoo and one egg of the bulbul.Clamator levaillantii (Swainson)Stripe-breasted cuckooRecently m South Africa, Milstein (1954, pp. 4-5) produced evi-dence indicating that this cuckoo may \?iy pm-e white eggs as well asthe greenish blue and pinkish ones described in my book. The caseis as follows. He saw two stripe-breasted cuckoos perching in awild fig tree in which there was a nest of a yellow-vented bulbul.The cuckoos repeatedly hopped towards the nest and the bulbulskept them at baj^, diving onto their backs, gripping, and literallyriding them to the ground. The intruders, fluttering wildly, neverretaliated even when one of the bulbuls yanked out a tuft of whitishbreast feathers from one of the cuckoos. Milstein watched thisrepeated attack and counterattack for over an hour and a half. Hethen left, but returned several hours later when he examined thebulbul's nest for the first time. It contained tvro eggs of the bulbuland four large white eggs, presumably of a cuckoo. The two bulbuleggs and three of the parasitic eggs were snugly settled in the nest,but the fourth v/hite egg, slightly pinkish (freshest?) was on the rimof the nest, almost falling out. Milstein was inclined to assume thatthe fresh egg had been laid during his absence by one of the stripe-breasted cuckoos that had shown so much interest in the nest afew hours earlier. The four white eggs measured, in millimeters, asfollows: 27 X 22.25, 26.6 X 21, 26.25 X 21.5, and 25.25 X 20.75.In this respect they agree with Ivnown eggs of both the stripe-breastedcuckoo and the pied cuckoo; in rolor they agree with authentic eggsof the latter. It is to be hoped that fm-ther observations, includingallo\ving such eggs to hatch and develop into diagnostically featheredchicks, may be forthcoming.To the stiU meager data on the breeding season of this cuckoo invarious parts of its range maj^ be added the fact that Verheyen(1953, pp. 305-306) records that it breeds during the rainy season inthe Upemba Park, Belgian Congo, where males with swollen gonadswere collected on October 7 and March 15.
AFRICAN PARASITIC CUCKOOS?FRIEDMANN 383Cuculus canorus gularis StephensAfrican yellow-billed cuckooInformation about this bird accumulates very slowly; however, itis 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 tricolorlayardi) containing a young yellow-billed cuckoo on Oct. 9, 1955, at aspot called Guinea Fowl, 12 miles southeast of Gwelo, SouthernRhodesia. At that time the nest contained two newly hatchedbulbuls as well as the 3^oung cuckoo. Ten days later the nest wasrevisited and was found to hold only the .young parasite. It was atthis visit that the young cuckoo, then well feathered, was identifiedas Cuculus canorus gularis.Neuby-Varty (1950, p. 38) found the gray-headed sparrow (Passergriseus diffusus) to be victimized in Southern Rhodesia. At his farm
'Torre," near Marandellas (no date given), he found a nest of thissparrow in a natural hole in a sugarbush containing two eggs of thesparrow and one of the African yellow-billed cuckoo. The egg of thelatter was very light bluish gray with blotches of slate and darkgreenish olive brown, mostly at the thick end but also scattered overthe rest of the egg, and measured 23.5 X 16 mm. This agrees verywell with the fully authenticated oviduct egg described by the sameobserver (Neuby-Varty, 1948, p. 158) two years earher. This recordis 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, supposedlyof this cuckoo, collected by D. Townley at Rumani, Southern Rho-desia, Dec. 20, 1929, from a nest of an unidentified babbler. The labelon the egg bears the words "identity sure," and the egg is not unlikeauthentic ones of this cuckoo. No babbler has yet been recorded as avictim of this bird, but this case must remain unidentified.Cuculus solitarius StephensRed-chested cuckooPlates 1-3Our knowledge of several phases of the life history of this cuckoohas increased greatl}' in the past few years. For easier reference anddiscussion these data maj- be treated under various subheadings.Eggs and Egg LayingWhen compiling the data for my book I was unable to find aninstance of more 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 thatthey were laid by different hens. There is, as yet, no evidence toindicate 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-chestedcuckoo are plain pale chocolate brown or olive brown without anymarkings, a sufficient number of divergent types since have beenrecorded from southern Africa to make it seem that there is morevariation 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 withone egg of the owner, which it closely resembled, being olive green incolor and heavily blotched with reddish brown, the blotches formingan almost solid mass at the obtuse end of the egg. In another nestof the same host species he found another egg of this cuckoo (identitycertain because the egg was allowed to hatch and the developmentof the chick foUov/ed in detail). This egg has a fawn colored groundand was heavil}- blotched with dark reddish brown. Still anotherparasitized nest of the Cape robin-chat was found containing an eggthat presumably was of this cuckoo. It vv'as "off white" in color andvery heavily speckled with large, dark brown spots.^ That threesuch divergent, blotched or spotted eggs were found in one localityseems to eliminate, or to render doubtful, the possibility of their beingunusual or pathological in any sense. Still another color variant hasrecently been described in Northern Rhodesia by Haydock (1950,pp. 149-150) as deep cream in color and with a very rough shelltextm'e.So few observations are available on the question of whether ornot the hen cuckoo removes an egg of the host when depositing oneof its own that the following case, incomplete as it is, is of someinterest. Liversidge (in litt.) writes me of a parasitized nest of aCape robin-chat, a bird whose name occurs under several headings inour present discussion. The nest was found by the observer's land-lady, who was certain that one day the nest was empty and the nextday it contained an egg of the cuckoo and one of the host. Thiswould suggest that the cuckoo laid the same day as the robin-chatand 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 didthe host, and that there was no egg for it to remove. As I haverecorded previously (Friedmann, 1949a, pp. 69-70), the red-chestedcuckoo 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 theegg may have been of the black cuckoo.
AFRICAN PARASITIC CUCKOOS?FRIEDMANN 385begun laying. In at least one instance where the cuckoo laid in anest already containing eggs of the host, it did remove one of themwhen depositing its own. Incubation PeriodThe nest of a Cape robin-chat studied by Liversidge, mentionedabove, yields the following information on this topic. The personwho first showed him the nest was quite definite that the eggs (oneeach of the cuckoo and the host) were first present on September 21(about a week before Liversidge first saw the nest). The cuckoo's egghatched on October 7, which implies that at least 16 days elapsedbetween the time the egg was laid and the date of hatching. It is notknown, however, if the host began to incubate on or immediatelyafter September 21; but at least it is clear that no eggs were laid inthe nest subsequent to the two seen on September 21, and it wouldseem that incubation may well have started on or just after the dateof laying. This account leaves much to be desired, but it suggests anincubation period somewhat longer than the one of 12}^ days reportedfor the related European cuckoo, Cuculus canorus. More accuratedata are needed to clarify this point.Host RecordsAccumulation of many additional host records indicates even morestrongly than before that the robin-chat Cossypha cafra is the chiefvictim of the red-chested cuckoo wherever the two occur together.At least three subspecies of the Cossypha are now knovfn to be para-sitized: the nominate one, for which I had numerous records previouslyand for which I could add at least as many new ones now; the tropicalrace iolaema, earlier listed on the basis of a single instance to whichseveral more may now be added (Wiley, 1948, p. 2) ; and the subspeciesdrakensbergi, 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 listsit in this capacity. No specific records or other data were given byhim. Recently Mr. H. M. Miles has informed me that at Melsetter,Southern Rhodesia, a correspondent of his saw a fledgling of this cuckooattended by a Natal robin-chat. There is nothing unexpected aboutthis record, but it is a welcome additional indication to the not tooinformative 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 isanother species of robin-chat, Cossypha heuglini, in a nest of which hefound, on October 22, two eggs of the owner. An hour later a red-386753?56 2
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chested cuckoo was heard caHmg near the nest. It was perched on asmall branch nearb}'. One of the chats was standing on the edge ofthe nest in a threatening attitude. The next morning the nest con-tained one of the original eggs and another much larger one thatmeasured 25.7X18.2 mm. It was ovate, deep cream in color, and hada very rough shell texture. A week later the nest was found to beempty when revisited. Benson (1951, p. 98) found another parasitizednest of this robin-chat at Ketete, Nyasaland, on September 22. Itcontained one egg of the host and one of the cuckoo. Near IMar-andellas. Southern Rhodesia, Neuby-Varty has reported seeing ayoung 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 observationsuggestive 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 ofLamprocolius sp. were seen feeding a 3'oung cuckoo recently out ofthe nest. On close inspection (the bird was caught) the latter appearedto be a red-chested cuckoo chick, having "the tj^pical yellow legs andfeet, and differing from C. gularis in being practically a uniform black,though there were some traces of rufous on the thighs and marginaltail feathers, and odd feathers of the breast were tipped with a dirtywhite . . . ." The identification of the starling fosterer was im-possible as the birds were not collected, but Haydock was inclined tofeel 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 (evenas Lamprocolius sp.) to the list of birds definitely recorded as hosts ofthis parasite. Merely seeing a bird feeding a fledged cuckoo is notnecessarily proof that it raised the latter. In tliis connection, it maybe pointed out that Harding (1948, p. 2) saw a young red-chestedcuckoo, out of the nest, being fed by a Fischer's slaty flycatcher{Dioptrornis fischeri) . The next day he observed the same cuckoobeing 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 leucophrysleucophrys) is a common victim of the red-chested cuckoo. Justwhat data or how many actual instances of parasitism were known tothese authors is, unfortunately, not stated. In my earlier account Ilisted but a single record under the name Erythropygia leucophrysbrunneiceps. This refers to the same bird, although it may be pointedout that Mackworth-Praed and Grant use the name brunneiceps nowfor a race of Erythropygia zambesiana, a species not yet found to beimposed upon b}^ the red-chested cuckoo.
AFRICAN PARASITIC CUCKOOS?FRIEDMANN 387The Cape wagtail (Motacilla capensis) was previously listed as ahost of the red-chested cuckoo on the basis of two records. To thesemay 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 therace Motacillo capensis wellsi with three eggs of the wagtail and one ofthe cuckoo found at Kabete, Kenya Colony, Apr. 26, 1944, by Wiley(1948, p. 2).A second record of the dusky flycatcher (Alseonax adustus) as avictim of this cuckoo comes to me from Walter Stanford (in litt.).On Dec, 22, 1952, at Deep Gat, Somerset West, Cape Province, hefound a nest of this flycatcher containing one egg of its own and oneegg of the red-chested cuckoo. The nest was built in a little recessedhollow 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}''-catcher's egg had done so and the young Alseonax was well feathered.A host overlooked when compiling my earlier account is the Capesparrow (Passer melanurus melanurus). C. J. Bergh (1942, p. 99)writes from Belfast, Transvaal, that he has known the red-chestedcuckoo to lay its eggs in a "mossie's nest." The name "mossie" isloosely used for several species of sparrows but is usually intended torefer to the Cape sparrow. Evicting InstinctConsidering the complete absence of any information on this topicup to the time of the completion of my 1949 book, and, indeed, theamount of evidence suggestive of the lack of such behavior, it isgratifying that a number of detailed pertinent observations have sincebeen made. There can no longer be aiy doubt that the evicting ofnest-mates is commonly, perhaps regulrrly, done by newly hatchedred-chested cuckoos, but it still seems n-^cessary to quahfy this withthe statement that it seems to be less universal in this species thanin the European cuckoo. In the latter bird it is extremely rare to findthe 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 suppliedme with fuller details, which are given here as they constitute abetter record of events than his condensed printed one.On Dec. 25, 1951, near Johannesburg, Reed discovered a nest of therobin-chat with three eggs, one of which was a red-chested cuckoo's.The nest was kept under close observation, and on the morning ofJanuary 1 he found that the cuckoo's egg had hatched but that therobin's eggs were still intact in the nest. By 7 a. m. of January 3
388 PROCEEDINGS OF THE NATIONAL MUSEUM vol. loeboth robin eggs had been ejected from the nest. One lay on the groundat the base of the tree and the other lay in a fork of the tree besidethe nest. Fortunately, neither egg was visibly damaged. Reedreplaced one egg in the nest with the cuckoo chick and watched thebird try immodiatel}^ to eject it. However, when the chick had gonethrough the performance of carrying the egg on its back to the edge ofthe nest and the egg had rolled back into the nest, Reed removed it.He repeated the experiment on January 4 with the same result, butwhen he tried again on January 5 he was unable to induce the chickto attempt to eject the egg. To make quite certain that the birdhad lost all desire to eject it, ho placed the egg on its back, but thechick lay passiveh^ and made no attempt even to remove it. Reedthen broke both robin-chat eggs and found no sign of incubation inthem.Another nest was found by the same observer on November 29.It contained a newly hatched cuckoo and a robin-chat's egg. OnDecember 2 the host's egg was found lying on the ground just belowthe nest. Reed replaced this egg in the nest two da3'-s later, but noteven putting it directly on the back of the chick would induce thelatter to attempt to evict it. The young cuckoo was at least 6 daysold 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 todisappear before that age.Near Cape Town, Liversidge (in litt.) watched a nest of a Caperobin-chat containing one egg of the cuckoo and one of the host andmade detailed notes. (Also, he kindly supphed the photographs foruse in this paper.) The cuckoo egg hatched, and within 24 hours thehost's egg was out of the nest. Liversidge put it back, but a few houislater, when Dr. Broekhuysen visited the nest, it was out again. Thelady in whose garden the nest was located was asked to keep upobservations, to replace the egg in the nest each time, and to see howoften the young cuckoo would evict the egg. After three moreevictions in 10 minutes, the observer lost some of her attentivenessbut reported that the chick threw out the egg many more times thatday, the total of such evictions being in excess of 25 by the time theobservations were called off. The chick was later given eggs of varioussizes and shapes, and even small stones, all of which it evicted, or, atleast, attempted to eject. This went on until its fourth day. Onthe fifth day an egg, when placed in the nest, was allowed to remainthere, but the ejection movements of the j'^oung cuckoo could still beinduced by prodding its back. Liversidge writes that the chick didnot 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 righttop 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 oftlieir sheaths. Bottom: 19 da\ s old; one da\- before leaving the nest. (Photos byLiversidge.)

AFRICAN PARASITIC CUCKOOS?FRIEDMANN 389The Nestling and Its DevelopmentWhen I wrote my earlier account of this cuckoo there were ahnostno 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 gapin our knowledge, while Liversidge (1955) has since filled a good dealmore. 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 thenestling stage (i. e., from the time of hatchmg to leaving the nest) ofthis cuckoo in nests of the Cape robin-chat was found by Reed to befrom 17K to 19 days, by Liversidge to be 20 days ?6 hours, while inthe nest watched by Hughes the young cuckoo was found to have beenkilled by a predator in the nest on its 20th day; one leg, severed fromthe body, was all that remained. The nestling period for the relatedEuropean cuckoo, Cuculus canorus, is 20 days, agreeing with thedata for the present species.On hatching, the young cuckoo is devoid of down and is very darkall over. According to Hughes it is shining black; Reed describes itas dark bluish black; and Liversidge records it as brown, changingwithin the next two days to purplish brown. The gape at first isyellowish orange with yellowish edges, but becomes wholly brightorange by the end of the second day. The bill is dark horn colorwith a whitish egg-tooth; the feet are dark flesh color. Liversidgefound a day-old chick to weigh 4.9 grams.On the fourth day the quill-like, stiff feather sheaths appear on theupper surface of the wings. The next day these sheaths are 3 mm.long and show some black pigment basally; on this day sheaths firstappear on the cervical tract. On the sixth day the skin over the eyesbegins to open, but only slightly, hardly enough to enable the bird tosee. On this day the sheaths begin to show up on the chin, forehead,and sides of the breast, flanks, and abdomen, the longest ones aretwice as long as they were on the fifth day, 6 to 7 mm. By the seventhday the cuckoo, now more than four times its natal weight, shows thefirst sprouting of feathers out of the sheaths on the wings and tailaccording to Hughes; Liversidge reports this as taking place on theeighth day, and notes that the feathers are grayish with a whiteterminal bar. On this day sprouting also begins on the sides of thebreast, and by the end of the ninth day this condition extends alongthe entire ventral tract.On the 10th day the eyes fully open, a most important change inthe development and behavior of the chick. The rest of the plumagebreaks from the enclosing sheaths on this day and the next and thebird suddenly becomes quite fluffy, a marked change in appearance.
390 PROCEEDINGS OF THE NATIONAL MUSEUM vol. loaOn 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 thesheaths, become darker, more pronouncedly blackish and less grayishdue to the increased area of their newh^ freed portions. The ventraledge of the bend of the wing is white, the alula is black, and the underprimary coverts are black with white bars; the feathers of the breastand sides of the abdomen are hght buff with a black subterminal anda broad buff terminal bar; the feathers of the midabdomen and thighsare paler. On the 12th day the rectrices show the contrasting whitetips very markedly because of the growth of the dark proximal portionof 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 thanthe rest of the plumage. From this stage on until the chick leavesthe nest, the changes are only matters of rapid growth. The feet arenow yellow, whereas originally they were flesh colored.By the time it leaves the nest the cuckoo has grown in weight morethan 11 times its hatching size. Liversidge's bu'd weighed 4.9 gramsat bu'th and nearly 60 grams when it left the nest.Aside from the purely physical development of the bird, as outlinedabove, Liversidge has put on record some highly interesting observa-tions of the growth and change of its behavior. For the first fivedays, as noted in our discussion of the evicting instinct, the chickreacts to nest-mates (eggs or other chicks) by getting them on itsback and pushing them out of the nest. This reaction is very weakon the first day, very strong on the second, and normall}^ ceases onthe fourth day, but it can still be evoked by artificial stimulation onthe fifth day. During this period the gaping response of the chickis 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., towardthe direction of the hght, rather than merely upwards, and may beinduced by either touching and sUghtly jarring the nest or by a sharpwhistled sound near the nest. Liversidge, however, considers thatthis directional gaping is more the result of an originally conditionedstimulus than of a visually oriented one. The location of the nestnecessitated the same approach by the foster parents when visitingit, and gaping became directed toward the entrance. It is true,however, that the gaping did become more directional at the timethe eyes began to open, even though the first day or so they werenot open enough to give the chick really effective or efficient vision.Both Liversidge and Reed observed the first threatening reactions onthe eighth or ninth days. Reed noted that the chick erected thefeathers of the head and neck and opened the bill, revealing the
AFRICAN PARASITIC CUCKOOS?FRIEDMANN 391bright orange gape. On a closer approach, the bird reacted bydarting its bill forward to strike the hand that was extended towardit, an action much more aggressive than any feeding behavior. Liver-sidge found that on the eighth day the eyes were open shghtlj" earlyin the morning but were closed again in the evening. It was notuntil 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. This, it ma}^ be noted, is a little later than inReed's experience, and ma}' reflect the range of individual variationin the rate of this stage of the development.The first noise uttered by the chick, according to Liversidge, washeard on the 10th da}^, a quiet, chattering note. The next day thisnote, now mach louder, was used apparent!}^ only in connection withbegging for food. The original gaping reaction given in response totouchmg the nest continued up to the 11th da}', but by the eveningof the 12th day such stimulation ojily elicited threatening reactions.The period from the 15th day of nestling life onward to the timethe chick is fledged and leaves the nest is characterized by a greatervariety of activities. The bird first begins to preen on the 15th day,at first in a haphazard, apparently nonfunctional way, but rapidlybecoming fuUy functional. On this day the nestling was first seentr^-ing to stand up, unsuccessfully at first, but soon becoming betterat it. On the 18th day Liversidge's bird first practiced stretching itswings, and from then on it quivered its wings whenever begging forfood or being fed. On this da,y, also, it first seemed to distinguishthe call of the foster parents, but showed no ability to differentiatebetween 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 youngcuckoo's development, the stage when it becomes separated from thecare of its host and becomes self-sufficient. Reed banded a youngred-chested cuckoo before it left the nest of its foster parents andfound 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 atleast a month, after which it was seen no more.Stanford (in htt.) watched a pair of Cape robin-chats feeding ayoung red-chested cuckoo some days out of the nest. He first foundthe latter by hearing its low-pitched but incessant "tsip-tsip-tsip"notes. When the robin-chats came close to it, the bird's caUingincreased in intensity; the young cuckoo bent foi*ward, opened itsbill, partly opened and drooped its ^^dngs and ruffied its plumage.The call became much louder and was repeated more frequently, thebill opening and closing spasmodically all the while. Once the bird
392 PROCEEDINGS OF THE NATIONAL MUSEUM vol.106had been fed it calmed do'uai promptly. He saw it fed eight times in30 minutes, mostly on what seemed to bo grubs, spiders, andgrasshoppers. Breeding RangeThe first indication that the red-chested cuckoo may breed inWest Africa is a record of a female with a well yolked egg in theovary collected on August 6 at Lake Bambalue, near Bamenda,British Cameroons (Bannerman, 1951, p. 274).Cuculus cafer LichtenstcinBlack cuckooSince the publication of my earlier account of this rather shy andsecretive cuckoo, additional data, especially on the development ofthe nestling and, to a lesser extent, on the evicting habit, have beenrecorded. This is principally due to Skead's (1951, pp. 163-168) ob-servations. In his area in the eastern Cape Province he found theboubou shrike (Laniarius Jerrugineus) to be the regular, and, as far ashis own experience went, the only host of the black cuckoo.The new data, filling what was formerly a complete blank m therecord concerning the growth and development of the young bird, maybe summarized briefly as follows. On the first day after hatching thebird is naked, dark purplish in color, the gape pink, the back noticeablyflat (not concave), and the eyes closed. On the fourth day the sheathsof the rectrices, remiges, and flank feathers begin to emerge; on thefifth day those of the feathers on the nape, throat, and forehead beginto appear; by the sixth day the bird has sheaths all over except on theback which is still nude. The eyes begin to open on the seventh day,and the bird begins to squeak softly. The feather sheaths begin toemerge on either side of the bare spinal area on the eighth day. Bythe nuith day the eyes are fully open; the back is still largely bare.The feathers start bursting from their sheaths on the 11th day, butthe primary remiges are still encased in their sheaths. On the 16thday the bird is well feathered all over, and shows fear reactions bygaping 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 seenagain. The nestling period is therefore not more than 21 days. Inthe Em-opean cuckoo it is said to vary from 20 to 23 days.In another nest of the boubou shrike Skead found a newly hatchedblack cuckoo and two eggs of the host. The next da}^ one of the shrike'seggs 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 orchick. The probabihty is that they were ejected by the cuckoo, butthe ejection was not witnessed. The nest, being a rather shallow cup,
AFRICAN PARASITIC CUCKOOS?FRIEDMANN 393was such as would present no obstacle to ejection by the cuckoo. Theinference is that the young parasite (definitely identified as a blackcuckoo by its plumage when the feathers had grown out) tolerated, orat least did not eject, its nest-mates for the first 48 hours. It is, ofcourse, unsafe to draw any hard and fast comparisons between a singleinstance in this species with what is Imown in its relatives, the red-chested cuckoo and the European cuckoo, but it may be pointed outthat the eviction of its nest-mates was delayed in the case of the blackcuckoo for at least 48 hours, as compared with less than 24 hours in asimilar situation involving the red-chested cuckoo, and about thesame in cases of the European cuckoo.Cuciilus poliocephalus LathamLesser cuckooAlthough this cuckoo does not remain in Africa during its breedingseason as far as known, some courtship feeding behavior of interest hasrecently been recorded in Northern Rhodesia. Grimwood (in Smith-ers, 1952, p. 107) watched five of these cuckoos at Danger Hill, 25miles from Mpika. Of these birds, two "pairs were indulging in whatappeared to be courtship flights, chasing one another from perch toperch, and on alighting going through a feeding behaviour, though nofood was seen to be transferred." While not surprising, it is of inter-est to find still another parasitic cuckoo exhibiting this atavisticallyrevealing behavior pattern.dialcites cupreus (Shaw)Emerald cuckooAdditional data on this bu'd have to do with its host species and withits courtship behavior. Host RecordsIn 1949 I quoted a peculiar account (Friedmann, 1949b, pp. 516-517) of the emerald cuckoo based on observations made on PrincipeIsland that were attributed to Keuiemans and published by BuUer(1873, p. 76). Mr. F. Haverschmidt has since sent me a translationfrom the Dutch of Keuiemans' (1866, pp. 363-401) observationson the birds of Principe Island. There is no mention of the queerstory related by Buller, but merely Keuiemans' statement to theeft'ect that he was told b}'' the inhabitants that this cuckoo depositsits eggs in the nests of Cuphopterus Dohrni {^Horizorhinus dohrni)and Parinia leucophaea (=Speirops lucophaea), which breed in thesummer, and that it probably deposits them in the nests of otherspecies as well.
394 PROCEEDINGS OF THE NATIONAL MUSEUM vol. loeA much later, and Kttle known, paper by Keulemans (1907, pp.245-247) gives further welcome details. He writes that the femaleemerald cuckoo lays her eggs in the nests of smaller bkds that buildopen, cuplike nests, and mentions Zosterops jicedulinus and Speriopsleucophaea as the usual foster parents. Of the latter species of hosthe relates a remarkable observation he made in July 1863 on his planta-tion on Principe Island, "Behind my small dwelling house grew alarge Acacia tree, the branches of which nearly touched the balcony.A pair of Parinias {=Speirops) had built their nest so near theverandah that I could hear the 3'oung ones while they were beingfed. One morning the pleasant sound of this young family calling forfood had suddenly ceased. A week later I noticed a young Cuckoolifting its head above the nest. Stranger still, two or three days latera second j'oung cuckoo was to be seen in the same nest, and both theseintruders were fed, not onl}^ by their original foster parents but alsoby at least a dozen other small birds." The nest and its contentscame to grief a few days later.These three hosts in Principe Island
?
Horizorhinus dohmi, Zos-terops Jicedulinus, and Speirops leucophaea?are all additional to ourprevious list. It is unfortunate that no details of actual instances areavailable for either of the first two. While there is nothing improbablein their being parasitized by the emerald cuckoo, Keulemans' unsup-ported statement does not constitute evidence in itself, and it cannotbe claimed that bh'ds of the first two species are known definitelyas fosterers of the emerald cuckoo.It may be pointed out that the Speirops case described above givesus the first evidence that more than one emerald cuckoo egg may belaid in one nest. We have no way of knowing if the two eggs werethe product of a single bird or of two different individuals,Winterbottom (1951, p, 27) hsts the puff-backed shi-ike (Dryoscopuscuhla) as a host of the emerald cuckoo but he gives no fm-ther data,Mackworth-Praed and Grant (1952, p, 509) also list puff-backedshrikes, together with bulbuls, as the usual hosts of this cuckoo.However, I am aware of but a single specific instance of the puff-backin 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 less strictly sylvan than thatof the emerald cuckoo, being more like that of the black cuckoo.On the other hand, hosts additional to those previously hsted areindicated in the following observations, Guichard (1950, p. 168)found the Abyssinian bulbul (Pycnonotus harbatus schoanus) to bevictimized at Addis Ababa, Ethiopia, while in Nyasaland Benson(1953, p, 35) records eggs of the emerald cuckoo in nests of Chalomitrasenegalensis 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 atBlantyre, 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 isT. V. viridis. The crimson-crested black forest weaver (Malimbusmalimbicus crassirostris) is added to the list of the known victims onthe following basis. The van Somerens (1949, p. 95) record an egg(identified as an emerald cuckoo's by Pitman) from a nest of this forestweaver in the Bwamba area, Toro, western Uganda, An egg of theweaver, punctured and empty, lay on the ground below the nest.One other bird not definitely known to be parasitized hj the emeraldcuckoo has been reported in a way suggestive of such a relationship.Holliday and Tait (1953, p. IIG) watched a nesting pair of olivebarbets {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 adultbarbets were nowhere in the vicinity, and disappeared from view intothe nest hole. All the time this was going on the young birds keptup their continuous clamour. Whilst the intruder was still withinthe nest, one of the parents arrived on the scene, and finding the un-wanted visitor, attacked it fiercely and drove it away from the nestsite. Owing to the fact that the nest was inaccessible, an examinationof the nesthngs was not possible, but one might assume that one ofthe chicks was probably a young cuckoo, and the parent was takingan active interest in its welfare, for it is not likely that it would belooking for a suitable nest in which to lay its eggs at this stage. Aslittle is known about the breeding habits of Emerald Cuckoos, it isfelt that this record is of some value, for some observers have seen itfrequenting the nests of other species of barbets during the breedingseason." The assumption that the adult cuckoo came to the nestbecause of a presumed interest in the welfare of an equally hypotheticalnestling of its own kind has little enough to support it, but the factthat the cuckoo showed some interest in the nest suggests that such asite might be within the range of its potential choice of a receptaclefor its eggs. Hitherto no hole-nesting bird as been found to be para-sitized by the emerald cuckoo, or, for that matter, by any of theAfrican metallic cuckoos of the genus Chalcites.CoTJRTsniP BehaviorWhen I wi'ote my earlier account of this cuckoo nothing was knownof its com-tship behavior. Since then this gap has been partly fiUedby 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. loebranch of a defoliated tree. A male was on a higher branch, callingloudly; it flew down to the hen and, with wings drooping and tailerect, bobbed in front of her, and presented her with a large hairycaterpillar. This offering was accepted and devoured, the cock birdcalling loudly with its head thrown v^^ell back during this time. Asecond similar display was then gone through, after which coition wasattempted unsuccessfully. Shortly after this both birds flew away.Haydock's observations were made in central Northern Rhodesia inthe third week in January.In my earlier account of this cuckoo, mention was made of the factthat male birds seemed to be "territorial" and to establish singingposts to which they adhered day after day. These observations Imade in western Kenya Colony. Welcome corrobation by virtue ofsimilar observations now is available from the Upemba Park, BelgianCongo, where Verheyen (1953, pp. 319-320) found the same situation.He goes further and writes that the males reserve a w^ell defined areawhich they defend against the intrusion of other males. This is,however, an unsupported statement as he does not describe anyactual instances of territorial defense, and all I knew previously wasmerely the fact that males sang persistently from their singing posts.In Principe and San Thome Islands, Keulemans (1907) found thiscuckoo to be migratory, being present on the islands only from Feb-ruary until November.Chalcites klaas (Stephens)Klaas's cuckooHost SpeciesAll the new information on this cuckoo refers merely to additionalhost records, the total of which may now be raised from the 33 specieslisted in my earlier accounts to 42 species, or, if we count species andsubspecies, to 50 forms. Of the total of all instances of this bird'sparasitism known to me, 30 percent are with sunbirds as hosts, 23percent with flycatchers, 16 percent with weavers, and the rest singleor, at most, two records for wagtails, bulbuls, babblers, starlings,white-ej^es, and finches. While 30 percent of the parasitized nestsare those of sunbirds, the 10 species of this family recorded as hostsof the cuckoo amount to not quite 25 percent of the total knownspecies of victims. Similarly, seven species of flycatchers, or about16 percent of the total of 42 kinds of hosts, account for almost 25percent of the recorded instances.The arrow-marked babbler {Turdoides jardinei) is listed as a hostof Klaas's cuckoo by Winterbottom (1951, p. 15), but no specificinstances are given. I am not aware of any published pertinent data.
ArEICA>: PARASITIC CUCKOOS?FRIEDMANN 397Another new host is the dusky flycatcher {Alseonax adusius), a pan-of which were seen feeding a young Klaas's cuckoo on December 6 inHottentot's Holland, Cape Pro%'ince, by MacLeod and Murray (1952.p. 22). Still another species of Alseonax, the Uganda pygmy duskyflycatcher {A. minimum jpumilus), may be added to the list, this oneon the basis of three records. Williams (1946, p. 138) found a nestof this bhd at Kampala, Uganda, on Apr. 27, 1946, that contamedtwo fresh eggs of the builder and one very slightly incubated egg ofthe cuckoo. At Kitale, Kenya Colony, Stoneham (1952, p. 7) foundtwo 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 (Parisomaplumbeum orientale) has been found to be parasitized in Nyasaland; hegives no further data on it, and Hsts it as an '"apparent" host of Klaas'scuckoo.In my book (Friedmann, 1949a, p. 140) I listed the Cape flycatcher{Batis capensis) as a host of this cuckoo solely on Joubert's statementthat it was victimized in the Cape Province. Since then MacLeodand Murray (1952, p. 17) have recorded two instances in the Hotten-tot's Holland area, and Stanford (in htt.) sends me the followingadditional cases. On December 6 at Picnic Bush, Somerset West,Cape Province, he and J. G. ^MacLeod found a young Klaas's cuckoo,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 thathad had a young Klaas's cuckoo in it a day or two earlier. In thisarea, in the years 1951-1952, Stanford and MacLeod found a total offive instances of the Cape flycatcher serving as host for Klaas'scuckoo.To the previous meager data on the Cape paradise fl^^catcher(Terpsiphone perspiciUata) may be added another instance, recordedat Hottentot's Hofland by 2\IacLeod and Murray (1952). The relatedTerpsiphone viridis is a bird of which two races, the nominate one andsuahehca, have been previously listed as victims of Klaas's cuckoo. Itseems 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 thebuilder and one egg that probably was of this cuckoo.The rufescent swamp warbler (Calamocichia refcsccns) is a bird nothitherto recorded as a victim of Klaas's cuckoo. Chapin writes methat near Tshibati, Belgian Congo, on Feb. 27, 1954, he found a nestof 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-entswere scolding^nearb}' as he examined the nest. The race of the hostin the eastern Congo is probablj'/oxi.
398 PROCEEDINGS OF THE NATIONAL MUSEUM vol. loeAt the time of my first report (Friedmann, 1949a, p. 141) I knew ofonly two instances of the bar-throated warbler (Apalis thoracicusthomcicus) being parasitized. This numl^cr may now be increased asa result of observations by Skead (1952, p. 12), who found one suchcase in December 1946 and two more in January 1947.The long-billed crombec {Sylmetta rujescens transvaalensis) waspreviously known as a victim of this cuckoo on the basis of a singlecase. 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 ofcrombecs feeding two fledgling Klaas's cuckoos.Another host previously known from one record and for which asecond has now come to my attention is the gray-backed bush warbler(Camaropfera brevicaudata abessinica). Sir Charles F, Belcher Avritesme 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 easternrace, immutabilis, has since been found to be similarly affected inKenya Colony, where Stoneham (1952, p. 7) found a nest at Kitalewith three eggs of the longtail and one egg of Klaas's cuckoo.Skead's (1954, p. 87) recent study of the greater double-collaredsunbird has added six more instances of its usage as a victim byKlaas's cuckoo. It must be regarded as one of the chief hosts of theparasite in the Cape Province.The East African mouse-colored sunbird (Chaleomitra veroxiijischeri) has been found to act as a host for Klaas's cuckoo in Kenj^aColony by Belcher (in litt.). I had previously listed it as such innorthern Zululand, but it now seems that the latter record refers tothe nominate race of the sunbird.Cinnyris venustus jalkensteini may be added to the known hosts ofKlaas's cuckoo, as Mackworth-Praed and Grant (1955, p. 779) recordit as being occasionally parasitized in East Africa. Previously I hadknown of only a single instance, involving the Abyssinian race Cinnyrisvenustus fazoqlensis (Friedmann, 1949a, p. 143).The Kenya rufous sparrow (Passer iagoensis rufocinctus) is anaddition to the recorded victims of Klaas's cuckoo. Belcher (1949b,p. 19) mentions, without further details, that he has in his collectioneggs of Klaas's cuckoo taken from a nest of this sparrow. I under-stand from correspondence that there were two eggs attributed to thecuckoo in this nest and that the identification of the parasite is notreally 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 399parasitized nests found in Uganda by Pitman. Each contained oneegg of the weaver and one egg of the cuckoo.The southern race of this weaver, Ploceus intermedius cabanisi, hasalso been found to be parasitized by Klaas's cuckoo in SouthernRhodesia, according to Mr. H. M. Miles. It had been recordedpreviously only as a host of the didric.Layard's spot-backed weaver (Ploceus nigriceps) was described asa 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 tobe a frequent victim, Grote (1924, p. 34) considers it a bird oftenimposed 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. OnNov. 6, 1952, at Murambinda, Buhera District, Southern Rhodesia,he found a nest of this weaver that contained three eggs of its ownand one egg that was presumed to be of a Klaas's cuckoo. The latteregg was freckled and was paler and slightly narrower than the host'seggs. A Klaas's cuckoo was calling excitedly in the vicinity andthree 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 cuckooin 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 LayingRecently MacLeod and Hallack (1956, pp. 2-5) described four eggsof this cuckoo?all laid in nests of the Cape flycatcher (Batis capen-sis)?as pinkish white and heavily spotted with reddish blotchesforming a fairly definite ring around the blunt ends. No pinkishwhite eggs of this cuckoo were previously recorded. These authorsthink (but do not know) that all four eggs were laid by the sam.e hencuckoo. This is based on the similarity of the eggs, on the fact thatall were found in nests within a limited area, and that all were foundin nests of the same species of host. The evidence, if such it may becalled, is merely suggestive, but, aside from adding another color typeto the known eggs of Klaas's cuckoo, it is the first indication of thenumber of eggs laid and of host specificity in this species.Eviction of Nest-MatesIn my earlier account I stated that while the young Klaas's cuckoois usually the sole survivor in a nest, no information was available asto 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 andHallack (1956) supplied our first definite observations on this point.They watched a parasitized nest of a Cape flycatcher. The Klaas'scuckoo hatched on November 9, and one of the host's eggs hatchedon November 13. On November 14 the J^oung cuckoo evicted the3^oung flycaU'her, At 9 a, m. on that day the cuckoo was notedunderneath 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 Batiswas left dangling by one foot, head down, from the outside of thenest. 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 theyoung flj^catcher fell to the rocks below.Chalcites flaviguUtris (Shelley)Yellow-throated cuckooThis rare cuckoo remains still virtually unobserved. Its rangemay now be extended a short distance eastward to the Bwamba area,Toro, western Uganda, where the van Somerens (1949, p. 29) collecteda breeding female at Ntandi in July and where Ridley, Percy, andPercy (1953, p. 163) obtained another female in full breeding con-dition on September 27 near Hakitengya. The stomacli contents ofboth birds consisted of hairy cateipillars, plus scale insects in theJuly bird and beetles in the September one.Chalcites caprius (Boddaert)Didric cuckooThe new information on this cuckoo may be treated under severalsubdivisions. Host RecordsA considerable number of species of birds not listed in my earlierreport have now been found to be parasitized by the didric cuckoo,bringing the total of its known hosts to 46 species, or 55 species andsubspecies of birds. In addition, our knowledge of some that wereincluded in the 1949 list has been considerably augmented. It be-comes increasingly clear that the Ploceidae are the chief victims of thedidric cuckoo; approximately three-fourths of all the nests recordedwith 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 anest of the Cape wagtail and tlie egg was hatched and the youngparasite reared by the host. Recently, Pike (in litt.) saw one ofthese wagtails feeding a fledghng didric on January 8 in the TranskeiDistrict, Cape Province. Skcad (1952, p. 4) lists three more instancesin the Capo Province. The Cape wagtail must be considered as aregular host of this cuckoo.Winterbottom (1951, p. 15) wiites that the didric is parasitic on thearrow-marked babbler {Turdoides jardinei). 1 know of no definiteinstance, although it may be recalled that Swynnerton (1911, p. 19)once shot a didric as it was leaving a nest of this babbler. There wasno cuckoo's egg in the nest, however.The small-billed brown flycatcher (Bradornis imcrorhynchus) is anew host of the didric. Belcher (in litt.) informs me that he collectedan egg of this cuckoo from a nest of this bird near Ngong, KenyaColony. He felt satisfied as to the identity of the species of cuckooinvolved.The Shoa olive sunbird (Cyanomitra olivacea ragazzii) is a possibleaddition 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, butthe catalog of that collection makes no mention of it. It is notstated how the identification of the Qgg was decided; it may have beena Klaas's cuckoo.Skead (1952, p. 5) has tentatively listed the yellow-throatedsparrow (Petronia superciliaris) as a victim of the didric cuckoo, buthe states that the parasitic egg seen in the one nest involved may havebeen that of a greater honey-guide. I think the latter is probablycorrect; the record should be deleted from the list of didric hosts.Vieiliot's black weaver (Ploceus nigerrimus) was known previouslyto be parasitized in Uganda and in the Belgian Congo. To this maybe added that Serle (1954, p. 55) found a fledghng didric cuckoo,unable as yet to fly, beneath a nesting colony of these black weaverson January 24 at Kumba, British Cameroons.The black-headed weaver {Ploceus 7nelanocephalus) is a species newto our list of victims, but it was actually recorded in this capacity aslong 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 thelatter rears the young parasites.Two other races of the black-headed weaver may also be mentionedin 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 theBcls^an Congo.The Uganda yclIow-coUared weaver (Ploceus melanocephalusdimidiaius) has been recorded, -without supporting data, as a victim ofthe didric cuckoo. Distant (1897, p. 143) states that Jackson foundan Qgg "considered to be of this cuckoo" in a nest of this weaver.To the meager data previously available on Layard's spot-backedweaver {Ploceus nigriceps) as a cUdric victim in southern TanganjdkaTerritorj' may be added that Benson (1953, p. 35) found it to beparasitized 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 afosterer of the didric in South Africa by Roberts. Recently Lamm(1955, p. 34) wTOte that this weaver appears to be the principal hostof this cuckoo in southern Mozambique. He also suggested that thespotted-backed weaver {Ploceus spilonotis spilonotus) is probably afrequent victim of the didric in that region. Previous data on thisbird 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 aIcnown host of the didric cuckoo. A definite instance has smce beenrecorded by Skead (1952, p. 9) at King WDliam's Town, Cape Province.Previously (Friedmann, 1949a, p. 176), I included Smith's goldenweaver {Ploceus suhaureus subaureus) as a victim of the didric only onthe strength of its being so listed by Patterson and by Sparrow, sinceno 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 hostand one of the didric cuckoo.New to our earher list of victims is the Zambesi brown-throatedweaver {Ploceus xanthopterus) . A young didric cuckoo w^as found in anest of this bird at Karonga, Nyasaland, on February 25 (Benson andBenson, 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 byBenson (1953, p. 35).The red bishop bird {Euplectes orix) was recorded earlier as afrequent victim, but how frequently it suffers from the attention ofthe 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 hostthat previously was overlooked. Fischer (1880, p, 190) found a nestof 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 othersseemed to be Euplectes eggs. It must be cautioned that there is noproof of parasitism in this case; the cuckoo may have come to laythere or to eat one of the eggs in the nest.The lire-crowned bishop bu*d (Euplectes hordeacea sylvatica) doesnot appear to have been recorded as a victim of the didric cuckoountil Benson (1953, p. 35) listed it as a host of this parasite in Nyasa-land, but ^\^thout further details. In the Upemba Park, BelgianCongo, Verheyen (1953, p. 315) saw a hen didric entering a colony ofthe nominate race of this bishop bhd but he did not obtain evidenceof any actual parasitism.The white-winged whydah {ColUispasser albonotatus albonotatus) issuspected of being a host of the didric in Southern Rhodesia byEdwards, according to Miles (1951, p. 4), but no definite evidenceseems to be known.The golden-breasted bunting {Emheriza jiaviventris) has beendefinitel}" added to the known victims of this cuckoo by Haj^dock(1950, p. 150; 1951, p. 3) in Northern Rhodesia. He observednesthng didrics, one in each of tlu'ee nests of this bunting, as well asthree young didrics already out of the nest apparently attended hythe buntings.When discussing the Cape rock bunting {Fringillaria capensis) as ahost of the didric, I mentioned (Friedmann, 1949a, p. 177) thatalthough several ^vriters had listed it as such, no localities or otherparticulars were available. It seems that these authors were copyingfrom 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 LayingIn my earlier report I stated that there is good evidence to the effectthat the didric hen often, if not regularly, may remove an egg fromthe nest when lading its own into it. Skead (1952, p. 9) came to asimilar conclusion as vrell, but Reed (1953, pp. 138-140) concludedthat the opposite seemed to be the case. Reed studied the didric'sparasitism on the red bishop bird, and found that the cuckoo "doesnot appear to remove an egg of the . . . host . . . the normal RedBishop clutch is three eggs and nearh^ all nests containing cuckoo eggscarried a total of four eggs." While he gives data on some 23 instancesin which the red bishop bird was parasitized by the didric, most ofthese cases were of nests mth j^oung cuckoos; in only six nests doeshe record four eggs each (thi'ee bishop bu-d eggs and one didric egg).
404 PROCEEDINGS OF THE NATIONAL MUSEUM vol. loeInasmuch as this matter of removing an egg often involves the eatingof it by the didric, it may be noted that Caldcr (1951, p. 36) watcheda hen didric enter a nest of a thick-billed weaver {Amblyospizaalbifrons), 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 minuteslater, but it would seem that this weaver may have been intended asa fosterer. It would not be surprising if someone were to find it toact in this capacity.In the disputed matter of adaptive similarity between the eggs ofthe cuckoo and those of its victims, the recently amassed data are justas conflicting as was the older information. Reed noted a definitesimila^rity between didric eggs in nests of the red bishop bh'd andthose of the host, while Skead found noticeable contrast between didriceggs and those of the builders in nests of Cape wagtails and Capeweavers. Evicting HabitThe 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 didricchick in the act of ousting either eggs or young from the nest. Theevidence from which eviction is assumed to have taken place is merelythe fact that eventually the parasite is usually the sole occupant ofthe nest. Assuming that the young cuckoo is responsible for thedisappearance of the other nest contents, Skead (1952, pp. 7, 9) foundthat a nestling didric in a Cape wagtail's nest "tolerated" the twoeggs of the host for two days after it hatched, when one egg was foundbelow the nest and the other was out of the nest early the nextmorning (i. e., after 2}^ days). In a Cape weaver's nest there weretwo weaver's eggs when the didric hatched; the next day one of theweaver's eggs hatched, but the following day both it and the unhatchedegg were gone, leaving the didric as sole occupant. In a spectacledweaver'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 theonly occupant. From these three examples it appears that 2 or 2)^days may be the usual time for the "eviction" (or disappearance fromthe nest) of the cuckoo's nest-mates. However, this is by no meansuniversal, as Reed (cit. supra) found in the case of a parasitized nestof the red bishop bird. When the didric hatched there were two ofthe host's eggs and one of its chicks in the nest, while four days laterthe nest contained three chicks of the bishop bird and the youngcuckoo. Unfortunately, Reed's observations ended at that point.However, in another nest of the same host species he found a didricabout four days old as the sole occupant. At 1 :30 p. m. he placedtwo eggs of the bishop bird in the nest; by 6 p. m. (4^^ hours later)
AFRICAN PARASITIC CUCKOOS?FRIEDMANN 405both of these were out of the nest, the inference being that they wereejected by the cuckoo. Keed's general observations, based not onl}^on those two cases but on many other less detailed instances, are thatthe didric nestling "appears to eject the eggs or young of its host inall cases because in no single instance were large Didric Cuckoo chicksfound -with Red Bishop .young in the same nest."Contrary to all the above are the cases observed by Pitman and byChapin (Friedmann, 1949a, pp. 180-181) in which the cuckoo and itsnest-mate grew up together until leaving the nest. The type of ex-periment tried by Reed should be repeated, with constant observationto see if, and how, ejection is performed.Fledgling FeedingAnother observation of fledgling feeding by a didric cuckoo comesto me from Pike, who saw, in the Transkei area, Cape Province, onDecember 11, a well grown young didric being fed by an adult maleof the same species. This further supports tlie general conclusionthat fledgling feeding is done by the male, and, to this extent, bolstersthe thought that fledgling feeding may be only courtship feeding witha grown young being mistaken for a hen by the cock bu-d.Additional cases of a male didric feeding another of its kind havebeen pubUshed by Maclaren (1953, p. 167; 1952, pp. 684-685), butthe light was poor on all occasions and he was unable to determineif the birds being fed were adult females or fidly fledged young.Literature citedBannerman, David A.1951. Birds of tropical West Africa, vol. 8, 552 pp.Bates, R. S. P.1938. On the parasitic habits of the pied crested cuckoo [Clamator jacobinus(Bodd)]. Journ. Bombay Nat. Hist. Soc, vol. 40, p. 125.Belcher. Charles F.1949a. Eggs from Somalia. Oolog. Rec, vol. 23, pp. 35-41.1949b. Review of "Parasitic Cuckoos of Africa." Nature in East Africa,ser. 2, No. 2, pp. 14-20.Benson, C. \V.1951. Breeding and other notes fromNyasaland and the Sundazi District ofNorthern Rhodesia. Bull. Mus. Comp. Zool., vol. 106, pp. 69-114.1953. A check list of the birds of Nyasaland. vi+ 118pp.Benson, C. W., and Benson, F. M.1949. Notes on birds from northern Nyasaland and adjacent TanganyikaTerritory. Ann. Transvaal Mus., vol. 21, pp. 155-177.Bergh, C. J.1942. Some birds of Belfast, Transvaal. Ostrich, vol. 13, pp. 98-99.BuLLER, Walter L.1873. History of the birds of New Zealand, xxiii+ 384 pp.
406 PROCEEDINGS OF THE NATIONAL MUSEUM vol. loeCalder, D. R.1951. A note on the didric cuckoo, Chrysococcyx caprius (Boddaert).Ostrich, vol. 22, p. 36.Chapin, James P.1953. Birds of the Belgian Congo, vol. 3. Araer. Mus. Nat. Hist. Bull.75A, 821 pp., 14 pis.1954. Birds of the Belgian Congo, vol. 4. Araer. Mus. Nat. Hist. Bull.75B, 846 pp., 27 pis.Clancey, p. a.1951. Notes on birds of the South African subcontinent. Ann. Natal Mus.,vol. 12, pt. 1, pp. 137-151.Distant, W. L.1897. (Editorial comment.) Zoologist, ?er. 4, vol. 1, p. 143.Fischer, G. A.1880. BrJefliche Berichte aus Ost-Afrika. Journ. Ornith., vol. 28, pp.187-193.Friedman N, Herbert1949a. The parasitic cuckoos of Africa. Washington Acad. Sci. Mono-graph No. 1 (1948), 204 pp., 10 pis.1949b. Additional data on African parasitic cuckoos. Ibis, vol. 91, pp.514-519.Godfrey, Robert1939. The black-cre.sted cuckoo. Ostrich, vol. 10, pp. 21-27.Grote. Hermann1924. Zum Brutparasitismus von Vidua serena (L.). Bcitr. Fortpflang-zungsbiol. Vogel., vol. 1, p. 34.Guichard, K. M.1950. A summary of the birds of the Addis Ababa region, Ethiopa. Journ.East Africa Nat. Hist. Soc, vol. 19, pp. 154-178.Harding, A. K.1948. Bird notes. Nature in East Africa, No. 7, p. 2.Haydock, Edward L.1950. Supplementary notes on African cuckoos. Ibis, vol. 92, pp. 149-150.1951. Extracts from a Northern Rhodesian diary. Babbler, No. 6, pp. 1-3.Holliday, C. S., and Tait, I. C.1953. Notes on the nidification of Buccanodon olivacea woodwardi (ShellejO-Ostrich, vol. 24, pp. 115-117.Hughes, Jean1949. Notes on a young red-chested cuckoo. Nature in East Africa, ser. 2,No. 2, pp. 2-4.Ivy, R. H.1901. Notes on the nesting and other habits of some South-African birds.Ibis, ser. 8, vol. 1, pp. 18-28.Keulemans, John G.1866. Opmerkingen over de Vogels van de Kaapverdische eilande en vanPrins-eiland (Ilha do Principe) in de Bocht van Guinea gelegen.Nederlandsch Tijdsschr. Dierk., vol. 3, pp. 363-401. '? ?' ,1907. The golden cuckoo. Bird Notes, vol. 5, pp. 245-247.Lamm, Donald W. i1955. Local migratory movements in southern Mozambique. Ostrich,vol. 26, pp. 32-37.
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