AMERICAN MUSEUM No Iltates PUBLISHED by the AMERICAN MUSEUM of NATURAL HISTORY CENTRAL PARK WEST at 79TH STREET, NEW YORK, N.Y

Home , Anteater chat, Campethera, Cardinal woodpecker, Crested barbet, Cuckoo paper wasp, Dendropicos

AMERICAN MUSEUM No iltates PUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORY CENTRAL PARK WEST AT 79TH STREET, NEW YORK, N.Y. 10024 Number 2825, pp. 1-46, figs. 1-21 August 9, 1985

Behavioral Notes on the Nest-Parasitic Afrotropical Honeyguides (Aves: Indicatoridae)

LESTER L. SHORT1 AND JENNIFER F. M. HORNE2

ABSTRACT New data from field studies ofAfrotropical hon- bonds, assisting female honeyguides to enter well- eyguides, examination of label data from speci- defended nests of their hosts. Some Lesser Hon- mens in most ofthe major collections having honeyguide (L minor) males seek out duetting pairs eyguides, and review ofthe literature are bases for of their barbet hosts, monitor them, and defend updating the biology ofAfrotropical honeyguides, them against conspecific male honeyguides. Hon- last treated by Friedmann (1955). Three species eyguides parasitizing barbets monitor barbet ac- (Prodotiscus zambesiae, Indicator meliphilus, and tivities about the barbets' roosting or nesting holes L narokensis) have been elevated from subspecific even in the nonbreeding periods. A nestling hon- status, and two new species (Melignomon eisen- eyguide (L minor) was seen making its initial de- trauti, and Indicatorpumilio) have been described parture from its host's (Stactolaema anchietae) since 1955. Emphasizing habitat, foraging habits, nest; a host barbet arriving to feed it shifted rec- foods, interspecific behavior, acoustic and visual ognition from that of a (foster) nestling to that of displays, hosts, and territoriality and breeding a "honeyguide," and immediately and violently habits, new insights are provided into honeyguide attacked the young honeyguide, which was driven biology, although much remains to be accom- out of the barbets' territory. Young honeyguides plished, and the biology of some species is vir- essentially must be able to fend for themselves tually unknown. Honeyguiding habits seem re- when they exit from the nest. The destruction of stricted to Indicator indicator. At least some the host's eggs or young, by the laying female hon- honeyguides (L indicator, L variegatus, and L mi- eyguide or later by the young honeyguide, is im- nor) are aggressive about sources ofbeeswax, with portant to insure that the nestling honeyguide se- a dominance hierarchy inter- and intraspecifically. cures maximum feeding. The same Lesser Exact pair relations largely remain to be elucidat- Honeyguide situation involving that nestling pro- ed, but evidence is presented that there is coop- vided evidence ofcontinual monitoring ofthe nest eration between individuals, hence extended pair by one and sometimes two Lesser Honeyguides ' Chairman and Curator, Department of Ornithology, American Museum of Natural History; Adjunct Professor, City University of New York. 2 Research Associate, National Museums of Kenya, Nairobi, Kenya. Copyright © American Museum of Natural History 1985 ISSN 0003-0082 / Price $4. 10 2 AMERICAN MUSEUM NOVITATES NO. 2825

right up until the fledging ofthe young honeyguide. guides. For the sake of completeness, full lists of The sight and sound of adult honeyguides thus honeyguide hosts are provided for each species, may be familiar to a young honeyguide even before and the information provided effectively sum- it leaves the hosts' nest. New vocal and visual marizes what is known of Afrotropical honey- displays are described for a number of honeyguides in 1984.

INTRODUCTION The honeyguides (Indicatoridae) include over short periods of time. Honeyguides are approximately 17 species of four genera dis- drably colored, generally inconspicuous birds. tributed in tropical Africa (the Afrotropics, a Their nest parasitism often involves hole- now widely accepted term for sub-Saharan nesting species, and the presence of honey- Africa) and tropical Asia. Only two species guides in the nests offoster parents often goes of one genus (Indicator) are Asian, the other undetected by human observers. No one, for species ofthat genus and the other three gen- example, knows what terminates the young era being endemic to Africa. These piciform honeyguides' relations with the foster parents birds are known especially for their honey- (but see below, under Indicator minor). guiding habit, and for their nest parasitism of other birds. In fact, these are little known birds, with most published information deal- LOCATION OF FIELDWORK ing with honey-guiding, which is actually Field studies have taken place during di- proven for only one species (Indicator indi- verse times of the year in Kenya in all years cator), and with details, some speculative or from 1976 to the present. Study sites have circumstantially derived, on hosts of these regularly included Kakamega Forest, Che- parasitic birds. The most recent treatment of sumot Farm near Kericho, 01 Ari Nyiro (Lai- the family is that ofFriedmann (1955). Since kipia Ranching Company, on the Laikipia that publication, scattered information has Plateau), the Kedong Valley, Karen, Olor- been published by Friedmann (1958, 1958a, gesailie National Prehistoric Site, the Loita 1963, 1968, 1969, 1970, 1978a, and 1978b), Plains and Narok region, and the coastal, and there have been notes in diverse journals threatened Arabuko-Sokoke Forest on nu- but few (e.g., Chapin, 1958, 1962; Colston, merous occasions. Other sites have included: 1981; Louette, 1981; Prigogine, 1978; Rob- the Endebess region and slopes of Mount El- erts, 1956; and Short and Home, 1979, 1983) gon in December 1982; the Lake Magadi re- that contain many details about honeyguides. gion and southern Ewaso Nyiro River in Jan- Our purpose here is to provide information uary 1983; the north (Kenyan) slope ofMount on the biology of Afrotropical honeyguides, Kilimanjaro in November 1982; Narosura, summarizing the scattered published details, the Lenkutoto Plateau, and the upper south- and adding new information on species that ern Ewaso Nyiro River in July 1979; the we have studied since 1976. Some ofthis new northwestern Mau Forest in October and No- information raises questions about the feed- vember 1976; the Naro Moru region in March ing habits, aggressiveness, territoriality, and- 1976; the area around Ngobit and Nyeri in other aspects of honeyguide behavior as pre- October 1976; the Loldaika region, Maralal, sented earlier by Friedmann (1955), and as and the area about Barsaloi in October 1976; generally accepted in the ornithological lit- Lewa Downs in July and August 1977; Lake erature about this interesting group of birds. Baringo and Tangul Bei in August 1977; the This report supplements and updates Fried- Narok-Mosiro region in July 1978; the Shim- mann's treatise, without duplicating its de- ba Hills in August 1978; Diani Beach, the scriptions and other information. remnant Jadini Forest, and Shimoni in No- We stress that the honeyguides are incom- vember 1981; the area about Mount Suswa pletely known, generally because individual in November 1980; the Kapsabet-Muhuroni birds cover large areas, fly rapidly, and con- region in September 1980; the area about centrate their feeding at beeswax or other sites Koru, the Nandi Scarp, and Chemelil in De- 1985 SHORT AND HORNE: HONEYGUIDES 3

: . FIG. 1. Study site of beeswax experiments, 01 Ari Nyiro Ranch, central Kenya, December 1983. Open riverine acacia woods (Acacia xanthophloea), with surrounding slopes in mixed dense acacias and other trees, elevation about 1770 m. METHODS AND cember 198 1; the Chyulu Hills in November MATERIAL 1982; and the Ngulia region of Tsavo West Observations were made with 8 to 10 pow- National Park in July 1977. We also have er binoculars, and notes were taken by writ- made study visits to Amboseli, Masai Mara, ing, and at times on tape recorders, as the and the Aberdare national parks, as well as observations occurred. Vocalizations were studying sporadically along roads throughout recorded by Home on a Stellavox SP-7 re- western, central, and southern Kenya, and corder, generally with a 72 cm parabolic re- coastal Kenya north to the mouth ofthe Gal- flector, and a Schoeps CMC-45 microphone. ana River. Home has studied birds in Africa Playback was by Short using a Sony cassette for three decades, and has made observations recorder. Audiospectrographic analysis was on honeyguides throughout Kenya (e.g., the with use ofthe Ornithology Department's Kay Lake Turkana region, the Tana River, Mar- Elemetrics Sonagraph 6061B, using wide- sabit, Lake Jipe, Kisii, the Ngurumans, Ku- band filter. lal, and elsewhere, and on frequent occasions Experiments were conducted on 01 Ari in Uganda, Tanzania (where she lived for ten Nyiro in December 1983, using commercial years), Zaire, and Ethiopia. Short has ob- "brown honeycomb" (a mixture of broken served honeyguides in Cameroun and South beeswax from honeycombs, dried particles of Africa, and both authors studied them in Ak- honey, and pieces of larvae and other stages agera Park, Rwanda, in early January 1982; of bees) and pure, transparent, white to in northern, central, and western Malawi dur- brownish honeycomb obtained from an ing August and September 1980; and in the abandoned bees' nest in an elephant-felled Ndola and Choma regions of Zambia in Oc- tree by Stephen Njugai. These two materials tober and November 1983. were placed at opposite sides of an emptied 4 AMERICAN MUSEUM NOVITATES NO. 2825

FIG. 2. Insertion ofbeeswax into abandoned, cut-out (by honey-gathering people) hive, below central large acacia in fig. 1, by field assistant Stephen Njugai. This and all following photographs (except fig. 4) were taken at 01 An Nyiro, Kenya, in December 1983. beehive in an Acacia xanthophloea tree; the Paolo Gallman and Mr. and Mrs. Colin Fran- abandoned hive had been cut open and the combe greatly assisted our studies on 01 Ari honey removed at least several months pre- Nyiro. Ann and Pat Donnelly helped us, and viously by human honey-gatherers (figs. 1- hosted us in coastal Kenya. Iris and Sandy 3). On ten days we made the two materials Wilson in Malawi, Jean-Pierre Vande weghe available to local honeyguides ad libidum for in Rwanda, and Margaret and Paddy Bruce- from 3 to 6 hours a day within that opened Miller, Loretta and Clide Carter, Dylan As- hive, the two sides being 25 cm apart. The pinwall, and Robert Stjernstedt in Zambia results of the experiments are described be- gave us a great deal of assistance. We also low. thank Cecilia Muringo, Claude Chappuis, Jo- seph Mwaki, Lorio Lukiro, Michael Webley, Anna and Karl Merz, John Wilson, Isobel ACKOWLEDGMENTS Bennett, Sybil Sassoon, Mulji Modha, Anna We are grateful to the Office of the Presi- and Chris Obura, J. Colebrook-Robjent, and dent of Kenya for authorizations to conduct many others who helped us in the field, in- research in Kenya, and to Richard Leakey cluding many game wardens and forestry of- and G. R. Cunningham-van Someren for their ficials. help in planning and facilitating our Kenyan Herbert Friedmann kindly commented on research. The Ministry ofTourism and Wild- a version of this manuscript, benefiting it life provided appropriate permits and facil- greatly. Anthony Diamond contributed un- itated research on reserves and in parks. Mrs. published observations on Indicator indica- 1985 SHORT AND HORNE: HONEYGUIDES 5

10 lk .OF

FIG. 3. The abandoned hive offigure 2. Pure honeycomb was placed at the far (left) end, and "brown" comb (broken comb with honey and bee fragments) was placed at the near (right) end, just inside the "lip" of the hive entrance. tor, for which we are greatly obliged. G. R. to species or described as new within the past Cunningham-van Someren made helpful quarter century, the latest being Melignomon suggestions regarding the manuscript, and eisentrauti (Louette, 1981). We have field ex- identified some stomach contents, for which perience with all the Afrotropical species ex- we thank him. cept Prodotiscus insignis, Melignomon zen- Financial assistance has come from the keri, M. eisentrauti, and Indicator willcocksi. Leonard C. Sanford and Ritter-Eisenmann Nonetheless, we treat all species -for the sake funds of the American Museum of Natural of completeness and because data on each History, and from Mrs. Marianna Collins. species affect the interpretation of data Without this assistance, which is gratefully available for all other species. acknowledged, we could not have conducted Detailed descriptions and discussions of our studies. systematics are to be covered elsewhere (Short and Home, in prep.), and we assign subspe- cies based on geographical reasons unless we state our other reasons for so designating SPECIES ACCOUNTS them. For each species we treat: range sum- The accounts that follow are in the system- mary, habitat, foraging habits, foods-, inter- atic framework provided by Snow (1978), specific behavior (unrelated to nest-parasit- which we follow very tentatively, for we feel ism), acoustic displays, visual displays, hosts, that taxonomic problems are numerous in and territoriality and breeding habits. Our this family. Indeed, almost a quarter of the concentration is on recently acquired infor- species now recognized have been elevated mation. Older information is either covered 6 AMERICAN MUSEUM NOVITATES NO. 2825 by citations or mentioned in detail only for branches, creeping over the bark, but it also the sake of clarity. flycatches from a perch and hovers before flowers to obtain insects. Sometimes it is found low in grass (Benson and Stuart Irwin, GENUS PRODOTISCUS SUNDEVALL 1967) or flowers in gardens. It forages with These birds are best called "honeybirds," flocks of white-eyes (Zosterops spp.) and tits as they certainly do not "guide" to honey (Parus). There are reports that it occasionally sources. Their hosts are diverse, including enters cavities, but whether to forage or to many that are not hole-nesting species. Thus, seek hosts' nests is uncertain (see Friedmann, honeybirds are the only piciform birds not 1955, p. 260). closely associated with cavities or holes in FoODS: This is primarily an insectivorous trees. They are inconspicuous, their voices species, eating aphids, lepidopterous and co- are little known, and few observers have not- leopterous larvae, and especially scale insects ed them more than casually. They have a fine, (Coccidae), but it also takes flying insects of sharp bill, which differs from the stubbier, unknown species. Some feeding occurs on heavier bill of most other honeyguides; but Loranthus, possibly for scale insects. Stom- they share with the others white outer tips of achs often contain the waxy exudate of scale the tail. White flank patches of feathers that insects, and beeswax itself has been reported are erected in certain displays are character- (Friedmann, 1955), although the bird is not istic. known to visit beehives. INTERSPECIFIC BEHAVIOR: Except for flock- Prodotiscus regulus Sundevall ing behavior noted above, and behavior re- Brown-backed Honeybird lating to possible hosts, there are no data. RANGE SUMMARY: It occurs disjunctly in ACOUSTIC DISPLAYS: Known calls include: West Africa about Mt. Nimba in Guinea, in a scream uttered in distress by a wounded Togo, and from eastern Nigeria to western individual; a chattering call; a single "zeep" Central African Republic; farther east in east- or a double "zeep-zeep" that is the main call central Sudan to central and southern Ethio- given in flight; a buzzy, strident note; a sib- pia; and from southern Uganda and central ilant "zzzz" reportedly uttered in courtship; Kenya southward circling the Congo forest and a supposed song, a "tseeeu tseeeu" in Rwanda, Burundi, and Tanzania, and from (Friedmann, 1955, p. 261) by a subadult fe- Angola and southern Zaire south to northern male that flipped its tail and spread it, show- Namibia, Botswana, Transvaal, eastern Cape ing the white outer tips. Province, and Natal. Southern African pop- VISUAL DISPLAYS: The tail movements of ulations may be partly migratory. a singing subadult female are mentioned HABITAT: This honeybird frequents very above. An apparent aggressive display in- diverse woodlands, even entering exotic volves the erecting ofthe white flank feathers (Australian) acacia plantations, the edges of to form conspicuous patches, used against a riparian forest, grasslands with scattered trees host species, Petronia superciliaris. When or thickets, dense thorn scrub, and orchards perched and foraging it makes distinctive side- and gardens. It avoids deserts and dense, wet to-side movements of the head, rather as if forests, but flies across open country without focusing visually on something, but the be- trees, perching occasionally on grass stalks. havior is conspicuous and species-specific (see Within woodlands it seems to occur in the P. zambesiae). These movements likely are understory and in smaller trees more often a display. The tail frequently is spread when than in the canopy, which the less frequently the birds are perched or flycatching; the spread observed P. zambesiae seems to favor. It is tail is an aggressive display generally in hon- found from sea level to approximately 2000 eyguides. In apparent courtship (or possibly m elevation (Kenya, Cameroun), perhaps an aggressive encounter), two birds pursued generally not reaching such high elevations one another, calling "zeet-zeet," then perched in southern Africa. 25 cm apart; one then drooped its wings, FORAGING HABITS: The usual feeding mode fluffed its body (especially flank feathers), and, is by gleaning on twigs amid leaves and on spreading its tail, leaned forward and called 1985 SHORT AND HORNE: HONEYGUIDES 7

"zzzz"-the second bird called "zeep" and south to northern Angola in the west, and flew, seemingly pursued by the first. northeastern Zaire and southern Uganda in HOSTS: This honeybird is known to para- the east (Snow, 1978). There are no records sitize the finch Petronia superciliaris, and the from coastal West Africa west of southern warblers Camaroptera brachyura and Cisti- Nigeria, and it is missing from large areas of colafulvicapilla. It has been noted near other central Zaire. cisticolas, and these may prove to be the ma- HABITAT: From the gaps in its range in West jor hosts. Brown-backed Honeybirds have Africa and in central Zaire, we suggest that, been seen emerging from nests occupied by although a forest species, this honeybird fa- the swift, Apus horus, and from nesting cav- vors clearings and forest edges rather than ities of the Pied Starling (Spreo bicolor). dense forest (see also Friedmann and Wil- Whether these species are parasitized, or the liams, 1971). It is found in second-growth, honeybirds were seeking insect food in those and in tall trees scattered above coffee plants, nests, is uncertain. This honeybird has been as well as in small trees at the edges offorest. seen being chased by pairs of Petronia su- It has been found from sea level to 1200 m perciliaris, Cisticola rufilata, and Nectarinia in Angola and Zaire, and up to 2200 m in senegalensis; these are known (Petronia) or western Kenya and Uganda (Britton, 1980). prospective hosts. FORAGING HABITS: This honeybird fly- TERRITORIALITY AND BREEDING HABITS: catches for insects from a perch, or flutters There is no knowledge of territoriality and or climbs from branch to branch, gleaning pairing. So sparse are data that we have sing- insects in the manner of species of Parus. ing only from a young female,3 and "court- Occasionally it joins mixed species foraging ship" noted that may in fact refer to an ag- flocks, in numbers up to three (usually only onistic encounter. Two birds, presumed to be one, or two). male and female, have been observed to- FOODS: It eats dipteran and hymenopteran gether for periods oftime while feeding, sug- insects, and probably numerous others. Some gesting that pair relations may involve more unidentified fruits have been reported (on la- than a briefmeeting for copulation. Data from bels) in stomach contents. Wax also has been specimens indicate that a female may lay five noted on labels, but whether of insects' exu- eggs or more. Breeding generally is in October dates or bees is uncertain. to March in southern Africa, and at diverse INTERSPECIFIC BEHAVIOR: Unknown, ex- times (e.g., March, May, October in Zaire) cept as related to flocking (above) or hosts not forming any pattern farther north. Ob- (below). servations suggest that the hosts raise only ACOUSTIC DISPLAYS: Various chattering one honeybird in a parasitized nest, hence notes and weak "whi-kiki" calls have been that the adult female honeybird, or the young reported (Friedmann, 1955, p. 251). bird punctures eggs or kills the young of the VISUAL DISPLAYS: Except for tail spreading host. displays as it flies in foraging, no displays are known. Prodotiscus insignis (Cassin) HOSTS: There are no absolutely certain rec- Cassin's Honeybird ords of hosts of this honeybird, although it RANGE SUMMARY: This honeybird, form- is reported or assumed to parasitize white- ing a superspecies with the more eastern and eyes (Zosterops spp.), flycatchers (Muscicap- southern P. zambesiae, inhabits West and idae), and warblers of the genus Apalis (see Central Africa from Liberia and Sierra Leone Friedmann, 1955, pp. 251-252). TERRITORIALITY AND BREEDING HABITS: east through southern Nigeria and southern Data are lacking on territoriality and hosts. Cameroun to northern Zaire, southeastern Individuals have not been followed in any Sudan, and Uganda to western Kenya, and reproductive setting. Peculiarly, Brosset (1981) has reported feeding of a juvenile of 3We heard L. Gillard's recording of2-3 second trilling this species by an adult Cassin's Honeybird, song in 1985, and used it to elicit and tape-record such the only such instance of possible parental songs in Kenya, from likely males. care known in the honeyguides. Breeding 8 AMERICAN MUSEUM NOVITATES NO. 2825 dates are based upon adult female specimens ing for two hours at 2 to 15 m, mainly at 5 with well-developed eggs in the ovary or ovi- to 10 m above the ground in ajacaranda tree, duct. These suggest November breeding in a fever tree, and in rose bushes in our Karen, Cameroun, January in Sudan, July in Ugan- Nairobi, garden, appearing to feed on two da, March in Kenya (Brown and Britton, consecutive days at the same sites and times 1980), July in Zaire, and September in An- (13:00 to 15:00 hours). Occasionally this bird gola. preened, drooped its wings in sunning itself, and scratched its head (under the wing) with its feet. It often bobbed its head when not Prodotiscus zambesiae Shelley gleaning with some movement to the side as Green-backed Honeybird well (hence its difference from P. regulus in RANGE SUMMARY: This uncommon hon- head movements is only in the up-down eybird replaces Prodotiscus insignis, its close component), as if peering at something, al- relative, in eastern and southern Africa. It though we noted no obvious object of such was considered conspecific with P. insignis scrutiny. This honeybird sometimes joins by Friedmann (1955), but now generally is mixed foraging flocks of sylviid warblers, treated as a separate species (e.g., Snow, 1978; Campephagidae (cuckoo-shrikes), Pycno- Britton, 1980). The Green-backed Honey- notidae (bulbuls), Zosteropidae (white-eyes), bird occurs from west-central Angola, north- and ploceid finches. These flocks may contain ern Zambia, southeastern Zaire, and south- two or rarely three Green-backed Honey- ern Tanzania south to northeastern Namibia, birds, and sometimes Brown-backed Honey- Zimbabwe, and southern Mozambique. Lo- birds occur in the same foraging flocks. cal, apparently somewhat disjunct popula- FOODS: It feeds chiefly on spiders and in- tions also occur in west-central Ethiopia, in sects, including scale insects and their waxy central and southern Kenya, and in north- exudate, termites, beetles, bugs (Homoptera), eastern Tanzania (Snow, 1978). moths, and flies. Stomach contents also men- HABITAT: It frequents woodlands, forest tion undetermined seeds and Loranthus ber- edges and clearings, riparian woods, and gar- ries. dens and other cultivated areas. In central INTERSPECIFIC BEHAVIOR: This honeybird Africa it reaches 2130 m and it usually is occasionally attacks tits (Parus spp.) and some found above 900 m in East Africa (Britton, other small birds, with which it may occur 1980); farther south it extends from sea level in mixed foraging flocks. No interactions have to 1400 m in Angola, 1200 m in Zimbabwe, been noted with Prodotiscus regulus, al- and 1670 m in Malawi. Its erratic presence though the latter has been seen in foraging in places in Kenya (e.g., Karen, personal ob- flocks with it. servation) and elsewhere suggests local ACOUSTIC DISPLAYS: Calls reported from movements, although this has not been es- this species are a chattering series, a note "fit," tablished. a flight call "skeee-aaa" apparently uttered FORAGING HABITS: The Green-backed during courtship (Friedmann, 1955, p. 251), Honeybird forages like a slow-moving sylviid and a likely song, "pee-eee-it" (Short and warbler, except that it frequently perches qui- Horne, 1979, pp. 3-4). The song was given etly and then sallies from the perch to seize repeatedly from a perch in the woodland of insects in the air. It often bobs its head when coastal Kenya. Playback of the call, which perched (see P. regulus above). We have seen consists ofa low "pee," followed by a whistly it creep about the bark and leafclusters, prob- rising "eee," and then a sharp clicking "it," ing into mosses and lichens, and taking scale caused the caller to move about excitedly, insects off petioles of leaves. When it shifts spreading its tail, and answering with the same to flycatching its flight often is showy, with call. In Karen, Kenya, playback ofthis "song" a flair caused by the spread, fully white outer to a nonvocal Green-backed Honeyguide tail tips; rather than directly taking the insect caused the latter to cease feeding, approach and returning to a perch, it may circle, zig- the source of the sound, and hop about with zag, or flutter to one side, then go on to another its tail repeatedly spread and shut (but it did perch. Although it often frequents the canopy not call). oftrees when foraging, we have seen one feed- VISUAL DISPLAYS: The head bobbing ofthis 1985 SHORT AND HORNE: HONEYGUIDES 9 distinctly small-headed species, with occa- GENUS MELIGNOMON REICHENOW sional side-to-side movements, may be a These small honeyguides are very little species recognition display enabling a mate known, one being described only recently or other conspecifics to maintain contact. (In (Louette, 1981). They combine a thin bill, shade the Green-backed Honeybird looks and feeding habits, and morphology, including a acts much like the flycatcher Muscicapa thick gizzard, and deep sternal notches of adusta.) One honeybird was reported to ap- Prodotiscus with the thick skin, greater num- proach another, bobbing its head up and down ber oftail feathers (12), coarser plumage, and and spreading its conspicuous white flank slit nostrils of Indicator. The sternal keel is patches, before both flew (Friedmann, 1955, intermediate between that of these two gen- p. 250, reporting J. G. Williams' observa- era. tions). Benson and Benson were cited by Friedmann (loc. cit.) as seeing an individual in flight over the trees while calling ("skeee- aaa") at intervals ofonce per second, possibly Melignomon zenkeri Reichenow a courtship flight display. Begging young Zenker's Honeyguide droop their wings and show their orangish RANGE SUMMARY: A rare forest bird, Zen- gape as they follow their foster parents about. ker's Honeyguide occurs in southern Cam- HosTs: The major hosts appear to be species eroun and adjacent Rio Muni, and farther ofwhite-eyes (Zosterops abyssinica, Z. polio- east from the southeastern Central African gastra, and Z. senegalensis), with which it Republic to northern and northeastern Zaire often can be seen foraging. It also is known and the southwestern corner of Uganda to use the Amethyst Sunbird (Nectarinia (Friedmann, 1968). amethystina) and the Black-throated Wattle- HABITAT: It is known -io occur in lowland eye (Platysteira peltata), a flycatcher (Fried- forest, reaching into the highlands of eastern mann, 1958a). G. R. Cunningham-van Zaire to an elevation of 1530 m about Lake Someren also reports having seen Dusky Fly- Kivu. catchers (Muscicapa adusta) and Paradise FORAGING HABITS: These are likely to be Flycatchers (Terpsiphone viridis) used as hosts like those of species of Prodotiscus to some by this honeybird. One seen at a hole in a extent (see foods, below) but observations are tree occupied by a pair ofSouthern Black Tits lacking. (Parus niger) may have been foraging, and FoODS: Contents ofstomachs include scale literature reports of tinkerbirds (Pogoniulus insect (Coccidae) pupae, emerging adults and spp.) and rock-sparrows (Petronia spp.) as cocoons, and gray waxy material mixed with hosts are unsubstantiated (Friedmann, 1955). insect fragments (Friedmann, 1968), proba- All ofthe known hosts build cup-shaped nests. bly from scale insects rather than wax from We have noted one Green-backed Honeybird beehives. flycatching about a pair of Montane White- INTERSPECIFIC BEHAVIOR, ACOUSTIC Dis- eyes (Zosterops poliogastra) that were gath- PLAYS, AND VISUAL DISPLAYS: Unknown. ering nesting material. HOSTS: Its hosts are unknown. Barbets have TERRITORIALITY AND BREEDING HABITS: been suggested but the similarity ofthis hon- Other than some indications ofaggression in eyguide's foods to those of species of Pro- displays, there are no data on territoriality or dotiscus equally suggests that it may use war- pair formation. Definite egg dates are from blers, flycatchers, or other hosts of groups April to July, December, and February, all used by other honeybirds of that genus. during or after rains in Kenya and northern TERRITORIALITY AND BREEDING HABITS: Tanzania, and August to October in Zim- The sparse data on breeding are gleaned from babwe (Stuart Irwin, 1981). Thejuvenal hon- specimens. A juvenile taken in March in eybirds fed by foster parents are unaccom- Cameroun suggests January and February panied by young of the hosts (inferring breeding. A Bwamba, Uganda, female had destruction ofthe host's eggs or young by the two ova well developed and a third enlarging honeybirds), although van Someren (1956) in July. A Central African Republic female found two young ofP. zambesiae in one nest taken in June had ova somewhat enlarged. of Zosterops. Prigogine (1971) and others have suggested 10 AMERICAN MUSEUM NOVITATES NO. 2825 breeding during the January to March rainy Melichneutes robustus (Bates) period in eastern Zaire, but it also appears to Lyre-tailed Honeyguide breed about Lake Kivu in (dry) August. RANGE SUMMARY: Because it is conspicu- Melignomon eisentrauti Louette ous mainly when in flight, and thus difficult Eisentraut's (or Coe's) Honeyguide to collect, this honeyguide is very little known, RANGE SUMMARY: This honeyguide appar- but its distinctive display sounds give us area knowledge of its occurrence in areas from ently is rare, being known from the which we lack specimens. It occupies forests around Mt. Nimba, Liberia, and in south- in the Mt. Nimba region of Guinea and Li- western Cameroun (Louette, 198 1; Colston, beria, and from southeastern Nigeria east to 1981). It also may occur in Ghana (Macdon- the southern Central African Republic and ald, 1980). This taxon appears to represent a formerly southwestern Uganda, thence south valid species that forms a superspecies with and M. zenkeri, from which it differs in its more to northwestern Angola and south-central olive-yellow dorsum, grayer ventral surfaces, eastern Zaire. and whitish posterior underparts. HABITAT: It is reported from forests and HABITAT: Specimens have been obtained forest edges, and also secondary forest, cleared and in secondary forest, areas in forest, and coffee plantations (Fried- in wet primary forest mann, 1978b). Although a lowland species, and it occurs likely in lowland forests gen- it reaches 2000 m elevation in eastern Zaire. erally in parts ofWest Africa where such for- to flycatch est remains. FORAGING HABITS: It is known FORAGING HABITS: It forages somewhat like for insects in the air (Chapin, 1939), and it upper parts forages about beehives. Most information sylviid warblers in the middle and comes from native collectors who find them of forest trees, according to Colston (in litt.). about bees' hives. It usually frequents the See related M. zenkeri. canopy but several have been collected in FoODS: Stomachs examined contained "in- mist nets within a meter or two ofthe ground sects," yellow "wax" or possibly pollen, seeds, (Friedmann, 1978b). There are no reports of and small fruits (Colston, 1981). its guiding animals to honey sources. INTERSPECIFIC BEHAVIOR, ACOUSTIC DIS- FOODS: From stomach contents, it is known PLAYS, AND VISUAL DISPLAYS: None reported. to eat beeswax, spiders, and undetermined HOSTS: Not known, suggested as possibly insects. woodpeckers (Campethera caroli, C. nivosa, INTERSPECIFIC BEHAVIOR: Unknown. breeding condition at same time as M. eisen- ACOUSTIC DISPLAYS: The vocalizations and trauti, but, if like Prodotiscus in foraging, mechanical sounds are surprisingly well could be parasitic on other than picids). known, and indeed are the main factor con- TERRITORIALITY AND BREEDING HABITS: A tributing to detection of the presence of this March female from Mt. Nimba was in breed- honeyguide. A purring note uttered in flight, ing condition, and molting adults suggest that a harsh call, a chattery "tarara-tarara-tar- breeding occurs there between November and ara-," and a buzzy low "ny, ny" are known June, and a December female from Mt. Cam- calls, without ascribed function. The display eroun, with an enlarged ovary may have been flight sounds for which the species is re- preparing to breed (Louette, 1981). There is markable were rendered "nyete-nyete-" by no positive indication of hosts. Chapin (1939), and usually called nyete sounds or calls. Recordings of this by C. GENUS MELICHNEUTES REICHENOW Chappuis that we have heard (see Chappuis, This monotypic genus, doubtfully distinct 198 1) are ofhonking or tooting series starting from Indicator, is characterized by a sternum as single, then giving way to double notes that deeper than that ofIndicator and highly mod- may be rendered "eh, eh, eh, eh-eh, kua, kua, ified (derived) central two pairs of tail feath- kua, kua-but, kua-but, kua-bu, kua-bu, kua- ers (Chapin, 1939, p. 554). The tail features bu, eee-eh." The end of the "call" almost is in a remarkable flight display, with sound that ofa braying, although it sometimes ends likely produced by the air moving through it. in a loud "wow-wow-wow!" There is an in- 1985 SHORT AND HORNE: HONEYGUIDES I1I crease in tempo and volume through the se- available regarding occasional guiding by L ries. Observers indicate that the sound is pro- variegatus. All other species that are reason- duced by the honeyguide when plummeting ably well known, as well as those that guide, downward in a spiral or zig-zagging flight. feed to some extent upon beeswax. Consid- VISUAL DISPLAYS: The display flight, erable uncertainty obtains concerning the sta- acoustic aspects ofwhich are described above, tus of taxa of the small- and medium-size consists of a honeyguide flying to a consid- members ofthe genus. We are not as yet pre- erable height over a forest during one or two pared to comment in depth upon the tax- minutes, then dropping rapidly over about onomy of these honeyguides (a preliminary 30 seconds, giving forth the sounds described manuscript is in preparation). However, we above. The sounds likely are produced by the do arrange the species in a unique classifi- tail feathers that are modified in both sexes. cation, based upon the clearly derived honey- Friedmann (1978b) feels that the reports of guiding habits, and sexual dimorphism ofthe this display essentially indicate that it occurs indicator species group (L maculatus, L var- year-round. Its function is unclear, but the iegatus, L indicator) of large-size honey- flights occur in proximity to feeding areas, guides. and they could be territorial, or be involved in pair formation or pair maintenance. Two birds do not usually display at the same time, Indicator willcocksi Alexander nor does a flight display elicit responses from Willcock's Honeyguide other individuals. RANGE SUMMARY: This species is known HOSTS: These are at present unknown, al- from Guinea-Bissau, Liberia, Ghana, and though Friedmann (1978b) and others (e.g., western and central Nigeria eastward through Rougeot, 1950) suspect that it parasitizes Cameroun and southern Central African Re- species of the forest barbets of the genus public to southwestern Sudan, and south- Gymnobucco, one or more of which occur ward to northern and eastern Zaire, and west- throughout the range of M. robustus. ern Uganda. It probably occurs in Ivory Coast TERRITORIALITY AND BREEDING HABITS: and other West African countries, but is often Nothing is known of its territoriality, if in- mistaken for other small honeyguides, es- deed it is territorial. Right displays appear pecially L exilis. to be not closely tied with territoriality. HABITAT: It is known as a forest bird ofthe Breeding data come chiefly from the condi- lowlands but its distribution and some ob- tion ofgonads in adult specimens. One nest- servations suggest that it actually prefers for- ling is known (Friedmann, 1978b, p. 651), est clearings and edges, riverine forest in open but its host's nest was not identified! It was country, and dense wooded patches in grass- taken during April in northeastern Zaire. lands. It occurs up to 1800 m in Zaire and Specimens of adults with enlarged gonads Uganda, usually below 1500 m (and below represent April and August in eastern Zaire, sympatric L pumilio) in eastern Zaire, but and June in the Central African Republic between 1200 and 1800 m in western Ugan- (Friedmann, 1978a). da. FORAGING HABITS: This little known honeyguide flycatches for insects from a perch, GENUS INDICATOR STEPHENS and gleans in the foliage of dense trees. The largest genus of the family, Indicator FoODS: These include flying ants, lepidop- includes the largest and smallest species, and terous (moth) larvae, other "insects," bees- is the only genus represented outside Africa wax, and wax of unknown origin, possibly (in southern Asia). Slitlike nostrils, a gener- exudates of insects. ally sturdy bill, thick skin, 12 rectrices, a tuft- INTERSPECIFIC BEHAVIOR: Unknown. ed oil gland, a thin-walled ventriculus, and a ACOUSTIC DISPLAYS: The only tape record- narrow keel ofthe sternum mark these species. ings of its calls are of its song (Chappuis, Guiding of mammals to beehives is known 1981). This is characterized by a sharp, me- for one species (I. indicator) conclusively, and chanical snapping sound at the end of each suggestive evidence requiring confirmation is three-parted note, and the lack of a distinc- 12 AMERICAN MUSEUM NOVITATES NO. 2825

tive initial note (found in related species L FOODS: Stomach contents are reported only exilis and L minor). From Chappuis' record- as beeswax and "insects." ings, which we have heard, the song can be INTERSPECIFIC BEHAVIOR: Although col- described as a "p-will (snap), p-will (snap), lected with L exilis at honeycomb sites pe-will (snap), pe-weel (snap), -." The notes (Chapin, 1958, 1962), its behavior with re- peak at about 2.5 kilohertz on their second gard to that species has not been described. element and end in a snapping sound at 2-5 ACOUSTIC DISPLAYS: Its calls are essentially kilohertz. We have played back copied songs unknown. A "tuutwi" call was reported by ofthis species in central Kenya, eliciting iden- Prigogine (1971) to be different from calls of tical replies, but so far have been unable to L exilis, but with no discussion or indication determine the identity of the singer (unpub- of function. lished notes). Prigogine (1971, p. 105) re- VISUAL DISPLAYS: Unknown. ported its vocalizations the same as those of HOSTS: Unknown. L pumilio; although it is uncertain which TERRITORIALITY AND BREEDING HABITS: vocalizations he meant, it is unlikely that two The only available data on its breeding come sympatric honeyguides would be vocally alike, from eastern Zaire specimens, namely ofju- and nothing like the above song of L will- veniles in June, August, and October, im- cocksi has been noted for I. pumilio. matures from December and January, and VISUAL DISPLAYS: Unknown. adults in breeding condition from May to HOSTS: Unknown. October. Prigogine (1971) reported breeding TERRITORIALITY AND BREEDING HABITS: It about Lake Kivu in April. is known to sing, presumably territorial songs, but data are almost nonexistent. Breeding as determined from the size ofgonads of adults Indicator narokensis Jackson is in: August and September about Mt. Nim- Kilimanjaro Honeyguide ba; February in Ghana; January, February, RANGE SUMMARY: We caution that this and May in Nigeria (Elgood, 1982); January honeyguide is unlikely to be distinguished in and February in Cameroun; and February, the field from I. meliphilus. Contrary to de- April, June, and September in Zaire (also scriptions by Forbes-Watson (1977) both Zaire juveniles in February and April). species have a loral mark, and seasonal variation in plumage coloration is so great that one cannot rely upon "greener" versus "grayer" appearance of, respectively, L meliphilus Indicator pumilio Chapin and L narokensis. From specimens, the Kil- Dwarf Honeyguide imanjaro Honeyguide occurs from the east- RANGE SUMMARY: This honeyguide occu- ern edge of Uganda across central Kenya to pies the highlands of northeastern Zaire, ad- southern Kenya (reaching woodlands near the jacent Rwanda and Burundi (Vande weghe, coast), and probably adjacent northern Tan- in litt.), and the Impenetrable Forest ofwest- zania. ern Uganda. Occurrences elsewhere are sus- HABITAT: It frequents dry woodlands, scrub pect because of confusion with other small woodlands, and wooded grasslands, includ- honeyguides, especially L willcocksi and L ing streamside vegetation in dry bushland, at exilis (Prigogine, 1978). elevations from near sea level to 1800 m. HABITAT: It occurs in montane forests at FORAGING HABITS: This honeyguide is elevations from 1500 to 2400 m, usually known to flycatch for insects from low to high above 1500 m, and at an altitude usually perches in the canopy. It also forages for bees- greater than that of I. willcocksi. wax at bees' nests. Insect foraging is con- FORAGING HABITS: This inconspicuous ducted at all heights in trees and in bushes. honeyguide was discovered only in the 1950s FOODS: Details from specimen labels in- by Chapin (1958), and its habits are little dicate beeswax, scales (likely scale insects), known. It mainly has been observed feeding other unspecified insects, and undetermined on pieces of honeycomb put out to attract fruits. honeyguides, in the company of I. exilis. INTERSPECIFIC BEHAVIOR: Suspected hon- 1985 SHORT AND HORNE: HONEYGUIDES 13 eyguides of this species (Short and Home, 1979) interacted with individuals of Indica- tor minor and were chased away by, or fled from, calling I. minor in Arabuko-Sokoke Forest, Kenya. It seems attracted by the activities, including calling, ofIndicator minor, although it is submissive to that species and usually moves away after quietly approaching the position of calling Lesser---Honey- guides. ACOUSTIC DISPLAYs: These are known only from uncollected small honeyguides believed to be ofthis species (because ofcolor pattern; specimens of it have been collected in the Arabuko-Sokoke Forest) in coastal Kenya during August. Squeaks and squeaky trills, a "yi-yi-yi-yi-eek" were uttered by such small honeyguides interacting with Lesser Honey- guides (I. minor) and with Black-collared Barbets (Lybius torquatus). The trills are at a tempo of 11-12 per second, and notes have a terminal peak sonagraphically at about 2 kilohertz. Songlike "whee-ip" calls were heard from these small honeyguides in the dry forest canopy; the "whee-ip" notes were repeated at 1.3 per second. Structurally the notes have a double peak at 4-5 kilohertz, and they FIG. 4. First known photograph of Indicator closely resemble such notes of Indicator mi- meliphilus in front ofexperimental hive, Novem- nor, I. conirostrus, and L exilis (Chappuis, ber 1984. 1981), but they tend to be more clearly double peaked. Obviously the identity ofthe calling birds has to be corroborated to insure that their songs during August in coastal Kenya, these calls are indeed those of L narokensis and thus appeared to be territorial. (see Short and Home, 1979, sonograms of figs. 1 and 3). VISUAL DISPLAYS: The crown feathers of Indicator meliphilus (Oberholser) birds suspected of representing this species Pallid Honeyguide (fig. 4) were erected during interaction with barbets RANGE SUMMARY: This small honeyguide and with Lesser Honeyguides, and the black- occurs disjunctly in East Africa (eastern tipped white outer tail feathers were spread Uganda to central and southern Kenya and in flight when these birds were chased by and northern Tanzania), and from central Angola otherwise interacted with barbets or honey- and southern Zaire through northern Zambia guides. to Malawi, central Mozambique, and east- HOSTS: These are unknown. Individuals central Zimbabwe. Since it is easily confused that called and were chased by Black-collared with Indicator narokensis in East Africa, Barbets appeared to monitor pairs ofthe lat- much care and collecting of specimens are ter, and were driven from them by Lesser required for identification, and field identi- Honeyguides. The suggestion is that Black- fication without the bird in hand is not fea- collared Barbets may be one of its hosts. sible. TERRITORIALITY AND BREEDING BEHAVIOR: HABITAT: Occupies woodlands of diverse Information is lacking, except as noted above kinds including brachystegia woods, riverine under Acoustic Displays and Hosts. Singing woods in dry country, acacia woodland, frag- birds responded by approach to playback of mented forest patches, the edges of forests, 14 AMERICAN MUSEUM NOVITATES NO. 2825 and, in Malawi, evergreen forest (Benson and Pallid Honeyguide in flying attacks upon, and Benson, 1977). The Pallid Honeyguide sup- in chases involving, Black Tits (Parus leu- posedly occurs in wetter habitats generally comelas), warblers ofthe genus Phylloscopus, than does I. narokensis, but confusion ofthese and Scimitarbills (Phoeniculus cyanomelas). two honeyguides makes it difficult to assess During these pursuits the honeyguide exhib- habitat preferences. Localities from which it ited great agility, speed, and maneuverability is known range from sea level to about 2000 in evading and in circling its slower antago- m (Britton, 1980). nists. FORAGING HABITS: These are mainly based ACOUSTIC DISPLAYS: Its voice has not been upon reports of the southern population, recorded on tape. We have heard a suspected which has fewer sympatric species with which Pallid Honeyguide utter a short "dzzzz" trill it may be confused. Birds feed on wax from during a chase involving a Scimitarbill, the bees' nests, both occupied and unoccupied trill resembling aggressive trills of L minor by the bees. Pallid Honeyguides creep about and other honeyguides. We also heard a sharp the bark oftrees, going into cracks and holes. "zit-zit" call from another (or the same) such The birds are usually very active, in almost bird a day later. Friedmann (1955, p. 226) constant motion. Individuals glean in flowers noted a soft twittering sound of an apparent and in the foliage, and flycatch for insects male near a suspected female, possibly a from a perch, often returning to the same courtship call. Other notes ascribed to it are perch (Carter, 1978). When seeking prey for a rasping "zeet-zeet" in flight, a "zit-zit" as flycatching, a honeyguide moves its head from reported above (possibly the same call as the side to side, peering, before darting after the "zeet-zeet"), a chattering note, and a sibilant insect (Carter, 1978). According to our Ke- noise like whistling through one's teeth (see nyan observations of a honeyguide likely to Friedmann, 1955). All ofthese and other calls be ofthis species, the flycatching occurs both require study and verification as to the species in low bushes and high in the canopy, and uttering them. may take place over an extended period (15 VISUAL DISPLAYS: The most conspicuous minutes, one sortie per minute). visual display is the frequent spreading or FoOD: It is known to eat beeswax, beetles, flicking of the tail, showing the white tips of and otherwise undetermined insects. It feeds the outer rectrices. This often is evident in at flowers, most likely on insects taken from Pallid Honeyguides giving chase to conspe- them. The regularity of its flycatching sug- cific or other individuals, as well as in perched gests that it feeds on various kinds of flying honeyguides. It seems to denote aggressive- insects, and that insects form a significant ness or assertiveness of the displaying hon- portion ofthe diet. It does not guide to bees' eyguide. A side-to-side movement ofthe head, nests, but feeds on beeswax obtained from as if to get a better view, is seen in perched active and abandoned hives that it finds. Pallid Honeyguides, especially between bouts INTERSPECIFIC BEHAVIOR: An individual of flycatching-this could be a display, and apparently ofthis species, observed in central certainly is a characteristic movement. Sup- Kenya, was subordinate to Indicator vari- posed courtship flight has been reported egatus and I. minor at a source of beeswax, (Friedmann, 1955), one bird fanning the tail and its small size suggests that it regularly and flitting about another, uttering a twitter- must give way to these and other larger hon- ing note (see Acoustic Displays), but, since eyguides (among sympatric honeyguides only the sexes cannot be distinguished, possibly L narokensis is smaller). In flycatching it fre- this could represent an aggressive encounter quently comes into conflict with other fly- between birds of the same sex. catching birds, and its aggressive manner of HOSTS. These are unknown; hosts ascribed spreading its tail and swerving about brings or reported are of doubtful validity. These it into encounters with other species. The are the Yellow-rumped Tinkerbird (Pogoni- Black-throated Wattle-Eye (Platysteira pel- ulus bilineatus, Friedmann, 1955, p. 227), tata), a flycatcher, has been noted attacking Bennett's Woodpecker (Campethera bennet- a Pallid Honeyguide (Carter, 1978). We have tii, Benson and Benson, 1977, p. 114), and noted an individual likely to represent the the Yellow-throated Petronia (Petronia su- 1985 SHORT AND HORNE: HONEYGUIDES 15 perciliarus, Mackworth-Praed and Grant It also has been seen investigating tree cav- 1970, p. 562). ities of diverse kinds, moving frequently at TERRITORIALITY AND BREEDING HABITS: various levels in trees. As many as three birds The only possible instance of territorial be- have been observed near one another in the havior has been mentioned above (see Visual tops of trees. We have seen it flitting about Displays). Individuals are aggressive and the mid-level of a Kenyan forest canopy, likely to be territorial, for rarely are two in- gleaning in leaf clusters, flycatching sporad- dividuals seen near one another, and they ically, and moving frequently. Another bird actively attack many small and even large moved along limbs and branches of forest birds of other species. Because of the uncer- trees, probing into mosses, lichens, and leaf tainty ofthe hosts, noted above, the breeding petioles, and flycatching occasionally. Others period is not well known, but it is likely to have been noted flycatching and feeding by breed when small, hole-nesting hosts, per- gleaning along the stem of a vertically hang- haps tinkerbirds (Pogoniulus spp.), nest. ing liana. Least Honeyguides are not known Adults in breeding condition are reported to guide humans or other mammals to bee- from Tanzania in February (Brown and Brit- hives, and are presumed to find beeswax at ton, 1980), and Malawi in September. Im- occupied or unoccupied hives on their own. mature specimens date from June and August FoODS: This honeyguide is known to eat in Kenya, from December in Tanzania, and beeswax, adult bees, bee grubs, hymenopter- from December and January in Zambia. Some ans of other kinds, aphids, ants, scale insects of these data could conceivably represent L (possibly also their wax exudates), spiders, narokensis rather than L meliphilus. some fibers, possibly yellow pollen, seeds, and fruits (spp. undetermined). INTERSPECIFIC BEHAVIOR: In Kakamega Indicator exilis (Cassin) Forest, western Kenya we had Indicator exilis Least Honeyguide respond four times by close approach to the RANGE SUMMARY: The range of this forest playback ofthe voices ofthe barbets Tricho- species extends, perhaps disjunctly, from laema hirsuta and (once) Pogoniulus bilinea- southern Senegal, Sierra Leone, and north- tus. In each case it perched nearby, flitted eastern Liberia (Colston, personal commun.) about spreading its tail, and looked at the east to Ghana, and then from southwestern recordist, over whom it circled (twice). Once Nigeria east through southern Cameroun, it kept up its circling, with spread tail, flut- southern Central African Republic, southern tering. The honeyguide did not remain for Sudan, Uganda, and western Kenya south to more than a minute, after looking us over. It Bioko (Fernando Po), Gabon, Cabinda, has been known to visit honeycomb sources northern Angola, northern Zambia, southern used by Indicator pumilio (Chapin, 1958, and eastern Zaire, and Burundi (Vande weghe, 1962), and by L variegatus (Archer and Glen, personal commun.). Adults are readily dis- 1969), but relations between species at the tinguished from other small honeyguides by hives or other sources have not been dis- the sharply contrasting blackish moustachial cussed. stripe and anterior forehead that set off the ACOUSTIC DISPLAYS: Chappuis (1 98 1) re- clear white loral marks. ported on the "song" of this honeyguide, a HABITAT: The Least Honeyguide basically series very like that of the song of L minor, is a forest-dwelling species, occurring as well but higher pitched and apparently lacking the in forest fringes, isolated forest patches, riv- initial, distinct note that characterizes the song erine forest, secondary forest, and trees in ofboth L minor and L conirostris. This may cultivated areas adjacent to forests. It extends be rendered "pew, pew, whew, whew, whew-" from lowland forests near sea level to mon- or a faster "wheer, wheer, wheer -" (from tane forest at elevations up to 2400 m in Chappuis' recordings, which we have heard Kenya, Uganda, and eastern Zaire. and analyzed). The circumstances of the FORAGING HABITS: Most observations of "song" utterances have not been described. this bird have been at bees' nests and honey- In western Kenya we have heard a songlike comb bait put out for collecting ofspecimens. "pee-pee-wheet, pee-pee-wheet" series, also 16 AMERICAN MUSEUM NOVITATES NO. 2825 rendered "sweet-bee-bee." These "songs" resented as follows: Bioko, June (also a June were uttered by a lone bird perched at 10 m subadult, and May-June adults with enlarged above a forest path in degraded forest during gonads); Cameroun, April and June (plus May December 1983. Playback ofthe song or call breeding noted); Congo, February; eastern resulted in close approach by the bird, that Zaire, February, June (two), August, and Oc- peered at us and looked from side to side, tober (three); and Sudan, February. Sub- but did not call. We got no response from adults and adults in breeding condition, tak- these Kenyan birds to playback ofChappuis' en with the juveniles just noted, suggest (1981) "songs" of L exilis recorded in Cam- breeding from January to September in east- eroun. ern Zaire, November to February in western The only other vocalizations reported are Uganda, July to November in western Ke- low siffling notes, and "tsa-tsa-tsa" calls, re- nya, and September (perhaps earlier) to Oc- peated in conflicts with one another, or in tober in Zambia (data are from specimens response to calls from other birds (Prigogine, and such sources as J. P. Chapin's carded 1971). notes, and Benson et al., 1971). Chapin's VISUAL DISPLAYS: Only a side-to-side Tshibati, Zaire, specimens (Chapin 1958, movement of the head, by lone birds, and 1962, specimens mainly in the AMNH) in- some erecting of crown feathers during play- clude fully juvenal-plumaged males with back response (presumably aggressive) are testes as large as 2 mm in diameter, and a known visual displays. subadult female having some juvenal feath- HOSTS: The hosts invariably are stated to ers with an ovary as large as 4.5 by 3 mm. be tinkerbirds of diverse species (see, e.g., These data suggest that Little Honeyguides Friedmann, 1955). Bates (1930) reported can breed before attaining a fully adult plum- young L exilis found in the nests of unspec- age (iuvenal features include an essentially ified barbets, and adults of this honeyguide white tip on rectrices 3, 4, and 5, lack of a entering holes excavated by the Yellow- white loral mark, the absence of a mousta- rumped Tinkerbird (Pogoniulus bilineatus) chial stripe, the lack ofblack on the forehead, and by the Gray-throated Barbet (Gymno- and deeper gray-green underparts than in bucco bonapartei)-the barbets were excited adults). by the presence of this honeyguide. None- theless, there are no certain hosts as yet de- Indicator conirostris (Cassin) termined, and eggs reported are from the ovi- Thick-billed Honeyguide duct of adult females, not from hosts' nests. Reaction of adult Least Honeyguides to ac- RANGE SUMMARY: This is a forest-inhab- tual and played back vocalizations ofthe bar- iting honeyguide that replaces its allospecies bets Pogoniulus bilineatus and Tricholaema L minor in the main forest region from Li- hirsuta suggests that these may be hosts. beria, southern Ivory Coast, southern Ghana, TERRITORIALITY AND BREEDING HABITS: and southern Nigeria east through central and Aggressiveness ofthis honeyguide (Prigogine, southern Cameroun to southwestern Central 1971) suggests territoriality, but there are no African Republic, northern Zaire, western details available on this aspect ofits biology. Uganda, and, disjunctly, western Kenya, There are no data on courtship. Breeding is south to northern Angola, and southern and indicated by a female held overnight on No- eastern Zaire. It generally is considered un- vember 20, which laid an egg at Ilaro, Nigeria common. The status ofthis honeyguide with (Elgood, 1983), and by oviduct eggs ready to respect to L minor needs clarification, as both lay in June in north-central Zaire and in Jan- occur together, or meet in some places in uary in eastern Zaire. At Mt. Nimba, Liberia, Uganda and western Kenya, but specimens an August adult female had laid, two females intermediate between them are known for in November had large ovarian eggs, and a Nigeria, Cameroun, the Ruwenzori region of juvenile was taken in December (Colston, Zaire, and northern Angola. Both appear to personal commun.). The duration of time have identical voices (Chappuis, 1981). over which the juvenal plumage is retained HABITAT: This forest honeyguide is re- is unknown, but juvenal specimens are rep- placed all about its range by L minor in ad- 1985 SHORT AND HORNE: HONEYGUIDES 17 jacent woodland and semiopen country, although it does occur in dense second-growth in Liberia (specimen data), and in gallery forests of Angola and Nigeria (Elgood, 1983). It generally is a lowland forest species, but extends up to 2300 m in western Kenyan montane forests. FORAGING HABITS: These are little known. Like all ofits genus it frequents beehives, and it often is seen about the regularly active colonies of the barbets of the genus Gymno- bucco. It flycatches for some insects, and comes to honeycomb put out as bait. This honeyguide is not known to guide humans or other mammals to honey sources. FoODS: It eats beeswax, bees, beetles, flying termites, and other insects. The remains of figs, other fruit, black seeds, and plant fibers have been noted in stomachs of adults. That some fruit is taken is not surprising, for barbets and other hosts regularly feed fruit to FIG. 5. Lesser Honeyguide (Indicator minor) nestling honeyguides, which obviously fledge just below experimental hive. successfully on a diet including much fruit. INTERSPECIFIC BEHAVIOR: No information most likely the other two species of Gym- is available about this honeyguide and its re- nobucco as well) as a major host (Friedmann, lations with host species, or, especially, with 1955). The only other species suggested as a the broadly sympatric and morphologically host is the tinkerbird, Pogoniulus bilineatus, very similar, though smaller, Indicator exilis. for which the single record is doubtful (Fried- ACOUSTIC DISPLAYS: Its song is so like that mann, 1955). of I. minor as to be indistinguishable. Tape TERRITORIALITY AND BREEDING HABITS: recordings of its song lent us for analysis by There are no data concerning territorial be- C. Chappuis suggest that it possibly sings havior, or breeding habits other than records slightly faster (but this may be insignificant, from collected specimens (in addition to ob- in L in servations of Thick-billed Honeyguides reg- given variation songs of minor), but ularly seen in the vicinity ofnests ofthe social pitch, quality, and structure its notes are like barbets ofthe genus Gymnobucco). Adult fe- those of L minor (see Chappuis, 1981), that males with large eggs are known from Liberia is, an introductory, dropping "wheew" fol- in September (egg in oviduct), and from east- lowed at 0.8 to 1.0 second intervals by four ern Zaire in December, and an egg was re- to seven, double-peaked, sharp "wheet" notes. ported from Cameroun in October. Large go- Most field workers consider this a quiet nads mark adults from: Liberia in November; species, and it is noteworthy that such a care- Ghana in February; Nigeria in May, August, ful observer as Chapin never heard its voice October, and December; Uganda and Kenya in all his time spent in the Congo region. in in October to Literature references give only a chirping call December; Zaire December; for it, but it is likely to utter squeaking and and Angola in February. Juveniles were found trilling calls similar to those known for L in April from Cameroun and June (nestling) minor. from western Kenya, and an April Cameroun VISUAL DISPLAYS: Unknown. specimen is subadult. HOSTS: The only definitely ascribed host is the Gray-throated Barbet (Gymnobucco bo- Indicator minor Stephens napartel), but it frequents colonies ofNaked- Lesser Honeyguide (figs. 5, 7, 8) faced Barbets (G. calvus) throughout the year RANGE SUMMARY: Generally this common and hence it very likely uses that barbet (and honeyguide occurs around the Afrotropical 18 AMERICAN MUSEUM NOVITATES NO. 2825

8 A f. I 6 'I 4 ' A 4 . #hd144 eo. e, '40. 0114 - MAA c 1.0 1.5 2.0 Ii F ~05 1.0 G 20 B 8 tI 1. :1 .v 6 :. 4 A 4 .. i. 2

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FIG. 6. Vocalizations ofLesser, Scaly-throated, and Greater honeyguides, all in wide-band sonagrams. A. Three variant songs of a single male Lesser Honeyguide (collected specimen), rendered, from left to right "pwee-weet-wee," "pwa-weet-wee-waa," and "wee-tya-wee-wit," Nov. 19, 1981, Arabuko-Sokoke Forest, coastal Kenya. B. Notes 8 to 20 of a 32-note squeaky trill ("tyeep" notes) of Lesser Honeyguide directed at White-headed Barbet (Lybius leucocephalus), September 1979, Karen, Kenya. C. Slightly wavering aggressive trill during interactions oftwo Lesser Honeyguides, Nov. 21, 1981, Arabuko-Sokoke Forest, coastal Kenya. D. Wavering aggressive trill of Lesser Honeyguide, Jan. 6, 1982, Akagera Park, Rwanda (disregard barlike elements at start and finish at about 5.5 kilohertz). E. Calls of two Scaly- throated Honeyguides in aggressive encounter over experimental honeycomb site, one giving chattery trill, other uttering screechy, long notes, Dec. 19, 1983, 01 Ari Nyiro, central Kenya. F. Two noisy chattery bursts ofinteracting Scaly-throated Honeyguides over honeycomb, Dec. 17, 1983, 01 Ari Nyiro, Kenya. G. Long screechy burst, interacting Scaly-throated Honeyguides, Dec. 17, 1983, 01 Ari Nyiro, Kenya. H. Hissing chattery birds, notes longer than in chattery trills but shorter than screechy notes (see E, F, G), of Scaly-throated Honeyguide in an encounter with another conspecific bird, Dec. 19, 1983, 01 Ari Nyiro, Kenya. I. "Tya-dah," double-noted call of interacting Scaly-throated tlIoneyguides, Dec. 17,1983, 01 Ari Nyiro, Kenya. J. Long series of squeaky calls by male Greater Honeyguide in close pursuit ofan African Drongo (Dicrurus adsimilis), Nov. 22, 1981, Arabuko-Sokoke Forest, coastal Kenya. belt from Senegal, Mali, Chad, southern and southern Central African Republic, northeastern Sudan, northern Ethiopia and northeastern and eastern Zaire, Burundi, and Tan- ern Somalia south in unforested areas to zania; and, south of the forest from northern southern Guinea, northern Ivory Coast, cen- Angola, southern Zaire, Burundi, and Tan- tral Ghana, Nigeria, central Cameroun, zania south to southern Angola, northern Na- 1985 SHORT AND HORNE: HONEYGUIDES 19

FIG. 8. Nestling Lesser Honeyguide offigure 7 just about to leave foster parent Anchieta's Barbet nest, same date and locality.

doned beehives, without guiding mammals FIG. 7. Nestling Lesser Honeyguide being fed to them, regularly investigating crevices, holes (fruit) by adult Anchieta's Barbet (Stactolaema in trees, and recently opened (by humans) anchietae) for the last time before it departed the hives. Usually solitary, two may occur near nest, not five minutes later. Near Ndola, Zambia, November 1983. a beehive, and it sometimes is seen in the company of Indicator indicator or L variegatus at hives. At times it flies to isolated mibia, northern and eastern Botswana, Or- trees in cultivated areas where it may sing or ange Free State, southwestern Cape Province, forage. We have observed Lesser Honey- Lesotho, Natal, Swaziland, and southern Mo- guides probing and gleaning on the bark of zambique, except in treeless areas. It appar- rough-barked trees in a woodpecker-like ently interbreeds with L conirostris in Nigeria manner. When perched, it often crouches, as and Cameroun (personal observ.), and per- if seeking prey, at which times the small- haps in Angola, but is usually ecologically headedness of the species is most evident. separated from that forest-dwelling species, Between foraging sites it often flies in a dip- which it meets parapatrically in Sudan, Ken- ping pattern and flicks its white-tipped tail ya, and Uganda. periodically; it may fly directly, as well, with- HABITAT: It occupies woodlands, wooded out undulations, or pause to chase individ- grassland, brushland, and the edges offorests, uals of other species that it may encounter. as well as cultivated areas with trees, avoiding Water sources, such as small streams or bird the interior offorests, and treeless grasslands watering baths put out by humans, are vis- and deserts. A broad range ofelevations from ited, and the bird drinks water occasionally. sea level to 3000 m (East Africa) is occupied FoODS: The Lesser Honeyguide regularly (Britton, 1980), although it usually occurs be- seeks out and eats beeswax, apparently di- low 2000 m. gesting it by the action of aerobic lipolytic FORAGING HABITS: This honeyguide is bacteria (Friedmann and Kern, 1956), al- adroit at flycatching for insects, and it also though A. W. Diamond (in litt.) and others gleans on the bark and in the foliage of trees. question the efficacy of aerobic bacterial It seeks beeswax from occupied and aban- digestion in the relatively anaerobic gut. When 20 AMERICAN MUSEUM NOVITATES NO. 2825 offered a choice of plain beeswax or brown, that function sufficiently as begging calls to honeyed wax containing bits ofbees and hon- cause feeding by the foster parents. Adults ey, they consistently choose the plain wax. make ruffling, mechanical sounds with the They also eat adult bees at hives, and "hawk" wings as they interact with one another or or flycatch for mayffies, winged termites, ants, with other species. Other mechanical sounds hymenopterous insects other than bees (e.g., include a so-called drumming flight display larvae and pupae of aculeate wasps), cater- made as one or two birds fly in bounding pillars, and spiders (Friedmann, 1955; per- flights up and down, with circling, fluttering, sonal observ.). Undetermined plant remains, and gliding movements. This sound is made seeds, and hairlike threads are known from by the tail (Friedmann, 1955), which often is stomachs of collected honeyguides. This spread during the flight display. The "drum- species may kill and eat small birds on rare ming" has been compared with the sound of occasions (Friedmann, 1955). Young are fed an irregularly firing motorcycle motor by many kinds of insects, and berries and other Brown (1948). Near beehives a clicklike note fruits by their foster parents (often barbets has been heard, and a strident alarm call, a and woodpeckers). "kish-kish-kish-," is known (Friedmann, INTERSPECIFIC BEHAVIOR: Other than many 1955). Squeaks, squeak-trills (fig. 6B), and honeyguide-host interactions, reported be- trill-squeaks are uttered singly or in series, low, this fast and aggressive honeyguide fre- speeding to a rattling "tyeek-tyik-tyik-tyik- quently engages in encounters with various, ik-ik-ikikikik" (Short and Horne, 1979), usually small, birds such as bulbuls (Pycno- marking interactions of Lesser Honeyguides notus spp.), Fiscal Shrikes (Lanius collaris), intraspecifically and interspecifically. These flycatchers (e.g., Batis molitor), and many may range from very squeaky and slow, at others (Friedmann, 1955; Ranger, 1955; per- five notes per second, to a fast, sharp sonal observ.). The honeyguide may initiate "dddddt" at up to 30 notes per second con- these without provocation, but its flitting and tinued for 2 to 3 seconds. The call may waver tail-spreading while making sallies for insects in pitch between 1.3 and 2.7 kilohertz, and sometimes trigger an attack by another often slows in tempo toward the end. More species -the honeyguide quickly becomes the squeaklike trills are softer, and probably aggressor unless the interactive species is very function at close range in encounters, whereas much larger. This honeyguide regularly at- louder trills (fig. 6C, D) are characteristic of tacks and drives away smaller honeyguides, chases, and of long-range interactions be- e.g., Indicator narokensis and L meliphilus tween honeyguides, or between honeyguides that it chances to find, and is dominant to and barbets (or woodpeckers). Actual contact them at beeswax sources. Its activities bring and battles often are marked by fast, high- it into conflict with the considerably larger pitched trills (personal observ.). These trills Indicator indicator and L variegatus, espe- are commonly heard during the breeding sea- cially near wax sources. These larger hon- son of this honeyguide's hosts. eyguides are dominant over L minor, and In a letter ofJuly 25, 1948, to J. P. Chapin, may chase the latter. However, L minor is Captain C. R. S. Pitman (unpublished files faster and more agile than are variegatus and ofDr. Chapin) reported a "pee-euw, pee-euw, indicator, and it may briefly return the attack, pee-euw" given during the dipping portions or, more frequently, it evades the slower at- of the flight of two possibly paired birds as tacker and circles back to the wax source. In they moved from tree to tree. This call closely this manner it is able to dart in, and secure resembles the so-called song (Friedmann, a few pieces of wax before the larger honey- 1955), a series of ten to 30 "weet," "pew," guide returns and supplants it (personal ob- or "wit" notes, introduced by a distinctive, serv.). We have seen it attack Tree Pipits (An- single "pee-yew" or "whee-ew" that may not thus trivialis), which have white tips of the be audible to an observer at a distance from outer tail like honeyguides, as the latter birds the honeyguide (Short and Horne, unpub- fly up to a tree after feeding on the ground. lished analyses). The main series of "pow" ACOUSTIC DISPLAYS: Nestling Lesser Hon- notes carries farther, to 365 m (Ranger, 1955). eyguides utter squeaky notes in long series Heard so far only from males, this repetitive 1985 SHORT AND HORNE: HONEYGUIDES 21 series of notes at about two per second is special song perches than are the typical songs. delivered for periods ofa few minutes to two These short variant songs often are uttered hours or more at 0.5 to 3.0 series per minute. in the vicinity of singing (duetting) pairs or It characteristically is given from a regularly groups ofthe barbet hosts ofthis honeyguide, used site, usually beneath the canopy of a e.g., Lybius leucocephalus and L. torquatus tree. In our experience the sites are not so (Short and Horne, 1979) in Kenya, and Stac- specific as indicated by Friedmann (1955). tolaema anchietae and Lybius chaplini in Rather, they are areas within a tree or even Zambia (personal observ.). Lesser Honey- a tree itself, and not a specific branch (one guides regularly come to our playback of the bird in western Kenya sang from at least six songs and duets of these and other barbets; points on all sides of a single tree). So-called the barbets likewise approach and act ag- song perches, which may be taken to mean a gressively (calls, displays) when we play back general site (e.g., a particular tree), appear to Lesser Honeyguide trill calls. be traditional, and are used for long periods VISUAL DISPLAYS: Tail spreading is an ag- ofthe year in southern Africa (see under Ter- gressive and assertive display given in flight, ritoriality and Breeding Habits). In eastern when one bird pursues another or sometimes Africa the song occurs over a shorter period, when no antagonist is apparent, and also while and indeed may be replaced by other calls a bird is perched and interacting with other that we have heard as far south as Zambia birds. When aggressively trilling, as in re- (personal observ., see below). sponse to playback and approach to the re- The introductory "pee-yew" note of the cordist, the bird often flicks the wings and typical song drops from an early peak ofabout moves the head from side-to-side as if peer- 4, to as low as 2 kilohertz, often with a step, ing to detect another bird. At such times the and it lasts about 0.2 second. "Pew" notes of bird often hunches its back, head lowered. the song are shorter, at 0.1 second, and they Since the spread tail is so conspicuous, the vary from being double-peaked to single- Lesser Honeyguide seems to control its spread peaked at about 4 kilohertz. These notes are very finely; when sneaking about near a host's extremely similar to, ifnot identical with, the nest, or when moving to a beeswax source in song notes of L conirostris (Chappuis, 1981). the presence oflarger species ofhoneyguides, Overtones are marked in both the "pee-yew" the white of the outer tail is completely hid- and "pew" notes. den from view dorsally, being covered by the Loud, clear songlike notes in series are more greenish central tail feathers. frequent in Kenya than are the songs de- During encounters with hosts such as bar- scribed above. These repeated series vary in bets, the Lesser Honeyguide may spread its form and length, from a short "pwee-wit," tail and half-glide into a perch, or give a pe- to phrases such as "pwee-wit-weet" and culiar fluttering, then sail to a perch. These "pwee-weet-tya-ya-ya-ya." Some variants are call attention to the often-calling honeyguide, shown in figure 6A. Sharper notes among and could serve to attract barbets from the these tend to resemble the double, introduc- nest area to attack the honeyguide. Since a tory note of the song, described above, but second honeyguide sometimes is present, they terminate, rather than begin with, a peak perched unobtrusively (and often passing un- at 2.5 to 3 kilohertz. Strong overtones occur detected by human observers), these flight in this song form, which varies in duration displays may be intraspecifically aggressive, from 0.5 second for a three-noted version to or even sexual in connotation, but such func- 1.2 second for those of seven or eight notes. tions require further study for elucidation. Such "songs" also have been recorded by us Other flight displays that involve "drum- on tape in Zambia, as for example, a "wa- ming" sounds made with the tail possibly are weet-weep-weet" from a Lesser Honeyguide territorial (Ranger, 1955). Two Lesser Hon- in November 1983 (near Ndola, northern eyguides may fly together in dipping flight Zambia). The relation of this song form to that has been taken to reflect courtship the typical song described above remains to (Friedmann, 1 955). be elucidated. We note that the variant Gaping is performed by Lesser Honey- "songs" are even less often delivered from guides that are attacked by other conspecific 22 AMERICAN MUSEUM NOVITATES NO. 2825 birds. During encounters, the bird attacked dence is as yet unavailable: the barbets Stac- seems to maintain a sleeked plumage, where- tolaema leucotis (personal observ.) and Ly- as the attacker often has the feathers of the bius vieilloti, and the Cardinal Woodpecker head, back, and thighs erected. (Dendropicos fuscescens). Possible hosts for Special mention should be made of the which data so far are inconclusive, or reports flight of this honeyguide, as it probably con- which require confirmation are: Red-fronted veys certain meaning to the birds beyond the Tinkerbird (Pogoniulus pusillus), Yellow- display flights noted above. Lesser Honey- fronted Tinkerbird (P. chrysoconus), Gray- guides can fly directly, or dip and undulate throated Barbet (Lesser Honeyguides about in flight. They are extremely fast and ma- active nests, personal observ.), Olive Wood- neuverable, completely outclassing their host pecker -(Dendropicos griseocephalus), Gray woodpeckers, and generally barbets as well Woodpecker (D. goertae), Greater Striped (only Stactolaema whytii and S. anchietae, of Swallow (Hirundo cucullata), Gray Tit (Parus all the barbets we have observed interacting afer), and Southern Black Tit (P. niger, the with Lesser Honeyguides, can catch them in species ofIndicator is uncertain, Friedmann, flight with any regularity). When interacting 1955). closely in chases, Lesser Honeyguides make Many ofthe species known to be hosts have rustling sounds with the wings, flailing with close relatives elsewhere within the range of them. Many times we have seen a barbet chase L minor, and at least some of these (e.g., a Lesser Honeyguide from one tree, only to Lybius guifsobalito, L. rubrifacies, Tricholae- have the honeyguide chasing it, pecking it, mafrontata) should prove to be hosts ofthis and beating it with its wings before the two honeyguide. reach an adjacent tree, where the barbet usu- Lists of hosts in themselves are apt to be ally lands. It is clear to us that Lesser Hon- deceptive. It is clear that barbets generally eyguides indeed entice barbets to chase them, are the preferred hosts of the Lesser Hon- usually resulting in rapid tiring ofthe barbets, eyguide, and that locally, one barbet species doubtless to the honeyguide's advantage (see may be preferred. Of 33 cases of hosts from below). Zimbabwe (Stuart Irwin, 1981), 23 are ofbar- HOSTS: The hosts ofthis honeyguide all are bets and 20 of these are of Lybius torquatus hole-nesters, chiefly barbets and woodpeck- (but nine are of the bee-eater Merops pusil- ers, but with two each ofbee-eaters and star- lus). Likewise, most records from Zaire (35 lings, and several of other groups. We con- of 50) represent barbet hosts. sider the following to be sufficiently TERRITORIALITY AND BREEDING HABITS: documented to be accepted (Friedmann, Pair relations and the occurrence of territo- 1955). The bee-eaters are Meropspusillus and riality are under study. It is clear that the M. oreobates (Friedmann, 1958). The barbets Lesser Honeyguide's breeding is more com- are: Stactolaema whytii, S. anchietae (Zam- plicated than it appeared to be earlier (Fried- bia, 1983, personal observ. offledging Lesser mann, 1955). It is the usual conception that Honeyguide), S. olivacea, Tricholaema leu- males in the breeding season sing from spe- comelaina, T. diademata, Lybius chaplini cific calling sites that are not defended, but (Colebrook-Robjent and Stjernstedt, 1976), may be used alternately by two or more males L. leucocephalus (Friedmann, 1970; Short and (rarely two sing simultaneously from the same Home, 1979), L. torquatus and Trachyphon- site, Ranger, 1955), and that females come us vaillantii. Woodpeckers parasitized in- to these sites to select a mate. In fact we can clude: Jynx ruficollis, Campethera bennettii, find no report of copulation at the calling and C. abingoni. The starlings are Spreo bi- sites, or "stud posts" (Friedmann, 1955) as color and Cinnyricinclus leucogaster. The they are designated, and we have noted (per- other hosts are: Striped Kingfisher (Halcyon sonal observ.) signs of aggression between chelicutO), White-throated Swallow (Hirundo honeyguides at diverse times of the year. albigularis), and Yellow-throated Petronia Calling sites that we have observed in Kenya (Petronia superciliaris). are no more specific than the top of a partic- The information available to us suggests ular tree, and the presumed males that sing these likely hosts, for which conclusive evi- at them are highly aggressive to other singing 1985 SHORT AND HORNE: HONEYGUIDES 23

Lesser Honeyguides that intrude, and to (hu- in the barbets' behavior.) There followed re- man) playback of these songs. peated visits to the empty nest by at least two In southern Africa north to southern Zaire, adult barbets carrying food (whether these songs are uttered from the calling sites for six included those that chased the honeyguide is to eight months of the year, generally in Au- uncertain), apparently bewildered by the sud- gust to March. In central and eastern Kenya den disappearance of "their" young-they we have rarely heard singing birds despite carried fruit to the nest, about it, into it, and field studies covering diverse seasons in the out, several times. Within 15 minutes an adult years 1976 to 1984. Secondary songs (see Lesser Honeyguide appeared, or reappeared, above) were heard more frequently in Kenya. approaching the nest and looking inside, de- Lesser Honeyguides regularly investigate spite attacks and pursuits by one or another used and vacant holes in trees (personal ob- of the Anchieta's Barbets. The honeyguide serv.), and they monitor and defend (prob- also peered into an older excavated hole lo- ably only the males) duetting barbet pairs cated above the actual nest (all from field- from other conspecific individuals, even when notes and personal observations near Ndola, the barbets are not nesting (e.g., Lybius tor- Zambia, in November 1983). quatus, Short and Horne, 1983; see also Short During nesting ofthe barbets, Lesser Hon- and Horne, 1979). As many as four males eyguides frequently are in attendance, al- replaced one another when collected in the though they may be inconspicuous except company of, and defending, a duetting barbet when trying to look into, or enter, the nest. pair (Short and Horne, 1983). Lesser Hon- On more than 40 occasions (of more than eyguides have been seen by us at the roosting 500) in which we observed interactions of a holes of nonbreeding groups of Stactolaema Lesser Honeyguide with individuals of var- whytii, Lybius melanopterus, L. torquatus, ious species ofbarbet near a nest, we have at and Gymnobucco bonapartei, in most cases least glimpsed a second, silent, apparently se- resulting in attacks and supplanting by the cretive Lesser Honeyguide nearby. These fig- barbets (personal observ.). ures are conservative, and are inexact be- In one Zambian case a Lesser Honeyguide cause we generally assumed that when one or (there may have been more than one) con- two barbets left chasing a honeyguide, and sistently approached the nest of a group of later returned with a honeyguide accompa- Anchieta's Barbets (Stactolaema) that con- nying them, the same Lesser Honeyguide was tained only a young Lesser Honeyguide, with involved. Since the birds at times were away daily visits involving calls and encounters be- for one-half hour or more, and we often saw tween the adult honeyguide and the barbets. a honeyguide while hearing a continuing chase A Lesser Honeyguide visited the nest, calling, of barbets and honeyguides in the distance, and was seen by (and saw) the young hon- more cases are likely in both categories. In eyguide moments before the latter fledged. addition, there are 55 cases in which a second The baby honeyguide had been fed regularly Lesser Honeyguide appeared and then called, by its foster parents (figs. 7, 8) and at least and became involved aggressively with the two other adult members of the group, with honeyguide already present (these likely rep- all of which it roosted nightly in the nest. resent male-male encounters). Since Lesser Immediately on its flying from the nest for Honeyguides perch frequently in the foliage, the first and only time, an approaching An- and when quiet are very difficult to locate, it chieta's Barbet carrying food to feed the young is likely that a second honeyguide was present honeyguide shifted from a "parental" to a more often than is indicated by our obser- defensive role, attacking the youngster re- vations. peatedly. Two adult barbets pursued the hon- Post-breeding Nubian Woodpeckers eyguide, hitting it in flight and chasing it be- (Campethera nubica) and Bennett's Wood- yond the limits of the barbets' territory. (It peckers (C. bennettii) also have been ob- is noteworthy that the characteristic honey- served being followed, approached closely, guide tail pattern could not be seen inside the and attacked by attendant Lesser Honey- nest, but became apparent instantly as the guides. The woodpeckers are less able than bird flew, this seeming to trigger the change barbets to chase the honeyguides away, and 24 AMERICAN MUSEUM NOVITATES NO. 2825 to pursue them with any ease, because they by the host species. The facts of the close are too slow and heavy. They engaged in long monitoring of hosts' holes and nests, even (to 40 minutes) engagements with Lesser when the latter already have a young Lesser Honeyguides, undoubtedly to the detriment Honeyguide inside, frequent conflicts be- of the woodpeckers' own accompanying tween male honeyguides (Short and Home, young (in the case ofthe Nubian Woodpeck- 1983) in proximity to the hosts, and the hon- ers). eyguide-related destruction of barbet eggs by Another incident shedding light upon the barbets suggest that hosts' holes and activities relation ofthe Lesser Honeyguide's behavior are related to territoriality, of some form, in to activities of its hosts occurred in Novem- the Lesser Honeyguide, and that the honey- ber 1983, north of Choma, Zambia. A group guide can influence hosts' nesting whether al- of Chaplin's Barbets (Lybius chaplini) at- ready parasitized or not. It is clear that hosts' tended a nest. There were occasional duets activities attract honeyguides, and the duets oftwo and even three ofthe four or five adults and other actions of hosts can serve as clues in the group. One and at times two Lesser to direct the attention of male and female Honeyguides were active about the nest, Lesser Honeyguides not only to the host, but chasing individual barbets, enticing them to to the resident Lesser Honeyguide as well. attack, and breaking up duets. Following one Limited data even suggest that young hon- ofthese broken duets and a subsequent chase eyguides in the nests of foster parents may of a honeyguide by one barbet, the barbet in be visited by adult honeyguides, and thus ex- the nest suddenly appeared at the entrance posed to the appearance and calls of conspe- with an egg in its bill. It left the nest, perched cific adults. Finally, our data suggest frequent nearby, and crushed and dropped the egg. To action in concert at prospective host nests by our surprise, the same barbet returned to the two Lesser Honeyguides, presumably paired nest, entered, came out with a second egg that birds, for male Lesser Honeyguides instantly it then broke, and methodically repeated this cease activities with hosts to attack some in- twice more, breaking four eggs in all. We later coming honeyguides (males, as shown in Short recovered the completely smashed, fresh eggs and Home, 1983), but move freely about with (hardly, ifat all, incubated), and studied them the hosts while ignoring, or even perching with Mr. Colebrook-Robjent, a veteran egg with, some other conspecific birds (likely fe- collector who had first described the eggs of males, see Short and Home, 1979, pp. 9, 1 1). this barbet (Colebrook-Robjent and Stjem- Associations ofmale and female Lesser Hon- stedt, 1976). He stated that each was clearly eyguides were reported by Roberts (1956). a barbet egg, and not a honeyguide egg. We Hoesch (1957) noted two cases of apparent have no reason to doubt this. pairs, or in any case two, Lesser Honeyguides What can we conclude from these obser- supposedly assisting pairs of the barbet vations? Obviously the matters of Lesser Tricholaema leucomelaina (a host) to drive Honeyguide territoriality and relations with away the barbet's nest-hole competitors, their hosts require additional study. It seems Riippell's Parrots (Poicephalus reuppellii). evident that the situation is complicated (it Female Lesser Honeyguides are reported may be that there is geographic variation in (Friedmann, 1955) to occasionally remove these details). The available data show that the eggs of the hosts in whose nest they lay, the Lesser Honeyguides, at least in some cases, but it is likely that the hatchling Lesser Hon- carefully and frequently monitor the activi- eyguide, with its sharply hooked bill, is main- ties oftheir hosts. They also react to the duets ly responsible for killing the hosts' young, or or playbacks of the duets of their hosts even destroying their eggs. The dead young are re- when the latter are not breeding, inciting moved from the nest by the foster parents aggression, fighting among themselves, and (Friedmann, 1955). We have found no rec- closely following the hosts about (Short and ords of two Lesser Honeyguides being raised Home, 1983). They seem particularly inter- simultaneously by any of its hosts. Attempts ested in holes used by the hosts for roosting by adult Lesser Honeyguides to enter occu- and nesting, and frequently fly to these to pied barbet nests are resisted vigorously, and examine them when not under observation Moyer's report (1980) of a Lesser Honey- 1985 SHORT AND HORNE: HONEYGUIDES 25 guide caught in the nest of, and apparently eyguide must be able to fend for itself, mainly killed by Black-collared Barbets (Lybius tor- or entirely, on fledging, as Ranger, 1955, p. quatus) indicates the risks that are taken by 86, also suggested.) The young are fed on the the honeyguides. foods ofthe hosts, which often include fruits, There are no data on laying schedules of and, of course, do not include wax. females, or on whether a single female utilizes The young are born naked, with a bill hook several host species, or specializes on one that is shed in 12 to 15 days (Ranger, 1955). alone. Sparse data suggest that individual They kill their host's young within the first Lesser Honeyguides, probably males, at least week, generally, and attempt to bite humans are attracted by calls and activities oftwo (or when handled for two weeks after hatching, more) host species, e.g., Anchieta's Barbet but not thereafter (Ranger, 1955). No host's and Black-collared Barbet in Zambia (per- young are known to have survived with a sonal observ.). young Lesser Honeyguide, which, when bar- The breeding period of this honeyguide is bets are the host, thus has to itself all of the diverse, as its wide distribution and many food brought by its foster parents and the hosts suggest. Brown and Britton (1980, p. helpers usually found in various species of 75) give about ten East African breeding rec- barbets. The feathers of the naked hatchling ords in the period April to August, during emerge in 7 to 12 days, and the eyes open in andjust after the main rains. Many host bar- 11 to 20 days (Ranger, 1955). Most Lesser bets in that region nest in the dry season (per- Honeyguides hosted by barbets probably sonal observ.), so we consider the few records leave the foster parents, at the latters' urging, ofBrown and Britton insufficient to establish when they fledge. The young honeyguide in a pattern. Specimen records include two fe- any case does not return to the nesting cavity males with a shelled egg in the oviduct from (as do the host's young), thus resulting in a Kenya in January, and one from northeastern situation unusual for the foster parents of Tanzania in February, outside ofthe months whatever host species. Reports of foster par- they report; juveniles from Kenya date April ents feeding the young Lesser Honeyguide are to December. Other laying dates are Septem- based upon circumstantial and unconvincing ber to November in Zambia (Benson et al., evidence (see Friedmann, 1955, pp. 210, 212), 1971; person. observ.) and Malawi (Benson and we frankly doubt that the hosts continue and Benson, 1977), September to January in feeding the newly fledged Lesser Honeyguide. Zimbabwe (Stuart Irwin, 1981; 30 of 35 records date October and November), and, from various sources, September to February Indicator maculatus G. R. Gray (mainly October to December) in South Af- Spotted Honeyguide rica. Juveniles from Uganda date November RANGE SUMMARY: This honeyguide occu- and December, and from eastern Zaire in Oc- pies a range complementing that of its allo- tober. Breeding is reported, with bases not species, L variegatus, in forests from Gambia always certain, or adults with enlarged go- and Guinea-Bissau across West Africa nads are noted (see Friedmann, 1955) from through all ofSierra Leone, and through Ivo- Senegal and Gambia in May and June, Sudan ry coast and Ghana to southern Nigeria, then in March, southern Somalia in March, Ugan- eastward through southern Cameroun, da in October to December, southern Zaire northern Zaire, the Ouossi River region of in September to December, Angola in No- southeastern Central African Republic, vember, and northern Botswana in Novem- southern Sudan, and southwestern Uganda, ber. and southward to Gabon, Cabinda, and cen- The Lesser Honeyguide's incubation pe- tral and eastern Zaire. It is narrowly sym- riod of 12 to an estimated 16 and one-half patric with L variegatus in southern Sudan days (three records, Friedmann, 1955) is less (Nikolaus, 1982) and western Uganda (Fried- than that ofhosts, whereas the nestling period mann and Williams, 1971). in two cases was 38 days, longer than that of HABITAT: It occupies forests, forest clear- the host Black-collared Barbets. (The long ings and edges, gallery forests, and dense nestling period suggests that the young hon- thickets that are in proximity to forests. It 26 AMERICAN MUSEUM NOVITATES NO. 2825 ranges from sea level to 1650 m in Cameroun which the birds had been trapped. These eggs (specimens) mountains, to 1500 m in eastern date January in Rio Muni (Friedmann, 1955), Zaire, and reaches a maximum elevation of representing Indicator maculatus stictotho- 2100 m in southern Sudan (Nikolaus, 1982). rax, and June from Ivory Coast (Friedmann, FORAGING HABITS: It is suggested (Fried- 1970), representing L m. maculatus. Juve- mann, 1955) that the Spotted Honeyguide niles, which have a distinctive breast pattern forages in a manner similar to that ofits close and whiter tail tips than do adults, are known relative, I. variegatus. It seeks beeswax, which from the Ivory Coast in June, Cameroun and it eats, and it readily comes to beeswax placed Gabon in August, and Sudan in March. Adults out for it by humans (Friedmann, 1978a). with gonads that are enlarged, and thus may The diet from stomach contents suggests that be in or near breeding condition, represent it flycatches for insects and takes these and Sierra Leone in March, Central African Re- spiders from the bark and foliage of trees. public in May and June (Friedmann, 1978a), FoODS: Stomach contents include spiders Sudan in March (Traylor and Archer, 1982), ofdiverse sizes, as well as caterpillars, beetles, and Uganda in July. There are reports of ants, and termites. Beeswax frequently is breeding from Cameroun in November, found, as well as small seeds, probably from northern Zaire in November to March, and fruits, pollen, and sand grains. eastern Zaire in February to March (Fried- INTERSPECIFIC BEHAVIOR: There are no data. mann, 1955). Molting birds represent the AcouSTIc DisPLAYs: Its voice has not been areas and months of the year indicated by tape recorded. Nikolaus (1982, p. 3) de- laying dates and juveniles, and, in general, scribed a loud call, probably that noted in honeyguides seem to molt from the main part most of the literature (see Friedmann, 1955) of the breeding season until just thereafter as "woe-woe-woe-," as follows: "like that of (personal observ.). a small falcon or similar to the call of the Long-crested Eagle Lophoaeetus occipitalis, Indicator variegatus Lesson but a little higher pitched." Many authors report that it has a trill or purring call very Scaly-throated (or Variegated) like the main trilling call or song (delivered Honeyguide (figs. 9-18) from "specific" song perches) of the Scaly- RANGE SUMMARY: This honeyguide occu- throated Honeyguide, whose song bears re- pies a parapatric range east and south ofthat semblance to the songs ofthe barbets Lybius ofits close relative, L maculatus, with which bidentatus and Trachyphonus vaillantii (per- it forms a superspecies (the two meet at least sonal observ.). in southern Sudan, and Bwamba Forest, VISUAL DISPLAYS: These probably are like southwestern Uganda, and probably in those of L variegatus. Its trilling song is de- northeastern Zaire, see L maculatus). The livered with throat and body feathers erected. range of variegatus extends from southern HOSTS: These are as yet unknown. The pre- Sudan, most of Uganda, central Ethiopia, and sumption is that the Gray-throated Barbet southern Somalia south (with gaps in arid (Gymnobucco bonapartei) and others of the areas lacking dense brush) around the eastern genus Gymnobucco are hosts (Friedmann, fringe of Zaire and through Zambia west to 1955). The very striking resemblance in de- north-central Angola, then south to south- tail of the head, throat, and breast patterns western Angola, northeastern Zimbabwe, and colors of L maculatus to those of the Swaziland, eastern Transvaal, Natal, and to Buff-spotted Woodpecker (Campethera ni- coastal eastern Cape Province in South Af- vosa) suggest that this may be a major host, rica, and to Mozambique in the east. but supportive data are lacking. Its range gen- HABITAT: The Scaly-throated Honeyguide erally correlates with that ofthis woodpecker. is a generally inconspicuous and even secre- TERRITORIALITY AND BREEDING HABITS: As tive resident ofthickets, woodlands, riverine its hosts are uncertain, there is little breeding forests, and forest edges and clearings. In dry information and there are no data on terri- bushland areas it is restricted to thickets and toriality. Eggs are known from two females streamside denser brush. Where its range ap- that dropped them, one each, in mist nets in proaches that of allospecies L maculatus in 1985 SHORT AND HORNE: HONEYGUIDES 27

FIG. 9. Adult Scaly-throated Honeyguide (In- dicator variegatus) below the-experimental hive of figure 3. Posture typical. FIG. 10. Adult Scaly-throated Honeyguide in somewhat submissive posture, showing cleft breast easternmost Zaire it avoids forest and is found plumage, near experimental hive. in wooded grasslands and riverine woods (Chapin, 1939). It exists in forests as well as open some nine months before by Kenyan woodlands in coastal Kenya, and in bamboo honey gatherers, and pure, clean chunks of forest on some Kenyan mountains (e.g., Mt. whitish beeswax (obtained by S. Njugai from Kenya, Britton, 1980). Highland evergreen a hive that had been cleared of honey and forest and juniper woods are occupied in abandoned by bees after elephants had pushed Ethiopia. Its altitudinal range varies as well, the tree over, onto the ground) some 25 cm for it occurs near sea level and in mountains away, at the other end ofthe opened hive. At to 3350 m (Mt. Elgon; Chapin, 1939). least seven different Scaly-throated Honey- FORAGING HABITS: A major food item of guides, and possibly as many as 15 came from adults is beeswax obtained from beehives that all directions over 30 hours of observations. the birds find in various ways. They seem In 104 clearly observed feedings, these hon- attracted by the presence of bees, they regu- eyguides selected the "honeyed" comb pieces larly inspect holes and crevices in trees, and four times, and in the other 100 instances fed they respond by approaching humans who on the bare honeycomb (figs. 13-16). The climb or bang on trees (personal observ.). Ex- feedings were marked by intense, regular, al- periments (figs. 1-3) in Kenya over a ten-day most continuous aggressive interactions (figs. period involved the placing of brown, hon- 16-17) and cautious, defensive approaches to eyed comb pieces (i.e., containing comb, hon- the honeycomb (figs. 9-12), and many rapid ey traces, remains of bee larvae, and other retreating flights from the hive by honey- debris of beehives, from an apiarist) at one guides carrying offa large piece ofbare comb. end of an abandoned hive that had been cut Honeyguides at the hive in the experimen- 28 AMERICAN MUSEUM NOVITATES NO. 2825 x%

FIG. 11. Adult Scaly-throated Honeyguide near FIG. 13. Adult Scaly-throated Honeyguide at experimental hive in full aggressive stance (crown experimental hive. The bird landed at the lower and thigh feathers extended, wings slightly out). (brown comb) end ofthe hive, but, tail closed and crown feathers appressed, it prepares to move to the upper, bare comb end of the hive, which a tal situation appeared to defend the food second honeyguide has just left. source until so disrupted from feeding (in order to feed, a honeyguide had to face into the hive and thus away from would-be assaults i.e., defending a prospective feeding site. Two by others) that they flew out, with or without complex cases at the experimental site illus- food, or were driven out by another Scaly- trate the aggressive nature ofencounters there. throated Honeyguide. We also saw, in three In one instance two honeyguides were at the cases elsewhere, a Scaly-throated Honey- hive very briefly together, one at the bare guide investigating a tree cavity turn and ward comb on one side, bird A, the other at the off an incoming, attacking conspecific bird,

FIG. 12. Stubby-tailed adult Scaly-throated FIG. 14. AdultScaly-throatedHoneyguideeat- Honeyguide, switching tail as it approaches hive. ing bare wax at the hive. 1985 SHORT AND HORNE: HONEYGUIDES 29

FIG. 15. Adult Scaly-throated Honeyguide pulling off large piece of comb (from bare comb), before flying off with it. honeyed comb on the other side, bird B. A FIG. 16. Scaly-throated Honeyguide (above) at B the hive. bare wax in hive faces off against crouching (ap- had been there when approached pressed plumage) Scaly-throated Honeyguide be- Bird B then called, gaped, approached bird low. A, and supplanted it at the bare comb, and A flew off to the canopy of the hive tree. As B began to feed on the bare comb, C, a third the adult at the bare comb, before the adult individual appeared, and it flew at B, which ceased tolerating its presence and displayed it supplanted. Bird B flew away, leaving C at at, or attacked it. On three other occasions the bare comb and A onlooking from the the subadult perched at the edge of the hive canopy. In another instance a subadult hon- without eliciting attack by a feeding adult, eyguide was allowed to perch and feed at the but it did not dare to enter to feed. honeyed comb by an adult at the bare comb. Thus, Scaly-throated Honeyguides do act After some 30 seconds, with a few aggressive, aggressively at or near food sources, both in- low calls uttered, the adult flew at the sub- traspecifically and (see below) interspecifi- adult, supplanted it, and began to feed on the cally. In the intraspecific encounters there honeyed comb. The displaced subadult seems to be a dominance hierarchy of some perched momentarily on a nearby branch, sort; at least adults are dominant to, but may then flew to the bare comb. As the subadult tolerate somewhat, subadults. Actual domi- commenced the adult left the hon- nance in feeding at honeycombs may partly feeding, depend on hunger, and on the natural ad- eyed comb and attacked the subadult, sup- vantage of an inflying bird over an already planting it. perched individual, but some birds, e.g., the We observed that subadults (or at least one stub-tailed adult, seemed to supplant and be subadult) of this species still uttered begging successful in attacks or defenses more often calls, different from the harsher aggressive than not, so a real hierarchy probably exists.4 notes, and tended to crouch, wave their wings, As for location of hives, we cannot com- and beg. In this way a subadult Scaly-throat- ment on Stager's (1967) suggestion that ol- ed Honeyguide was able to approach the hive faction may play a role in the honeyguide very closely, more so than any adult, while finding the wax, except to state that inquis- an adult fed at the bare comb. In three instances the subadult actually was able to feed 4 There exists a definite hierarchy, proven by 1985 briefly at the honeyed comb in proximity to studies of 16 color-banded birds at one hive. 30 AMERICAN MUSEUM NOVITATES NO. 2825

.4 ..i li* ,.l.,E FIG. 17. Scaly-throated Honeyguide (above) calling at retreating conspecific bird near hive below. itive, keen-eyed honeyguides that regularly trees and bushes, or in the understory and investigate holes, and are aware of human middle levels of woodlands. activities associated with honey-gathering are FOODS: Known items of food, in addition not apt to need the use of olfactory cues. We to beeswax, are bees, bee larvae and pupae, have no evidence of honey-guiding by this aphids, ants, flies, beetles, termites, caterpil- honeyguide, and we are impressed that the lars, other insects and arthropods, and some van Somerens, Skead, Chapin, Friedmann, honey, figs, seeds (most likely of fruits), and and other long time careful and interested remains of other unidentified plants. African observers have never observed this INTERSPECIFIC BEHAVIOR: Although usually species to guide humans or other animals to unobtrusive and keeping to cover, this hon- bees' nests (Friedmann, 1955). Hence, we eyguide is aggressive to other birds that may doubt that this secretive bird regularly, or approach it too closely, and to other honey- even occasionally guides humans to honey guides in feeding situations. We have seen it sources -the few reports cited by Friedmann attack Common Bulbuls (Pycnonotus bar- are unconvincing and require corroboration. batus) that were watering, and on eight oc- Ofcourse beeswax is not the only, nor even casions, Tree Pipits (Anthus trivialis), which the major, source of food for Scaly-throated have a white-edged tail, when the pipits were Honeyguides. These birds flycatch for insects in flight. from perches. They almost always forage in- In our experimental situation in central dividually. Occasionally, a Scaly-throated Kenya, described above (see Foraging Hab- Honeyguide is seen feeding with other birds its), Scaly-throated Honeyguides consistently in mixed-bird foraging flocks. It gleans from dominated Greater Honeyguides (Indicator the bark and leaves of trees, moving quietly, indicator), Lesser Honeyguides (L minor, fig. usually hidden from the sight ofthe observer. 18), and an apparent Pallid Honeyguide (L Much foraging is accomplished low in dense meliphilus), keeping them from beeswax by 1985 SHORT AND HORNE: HONEYGUIDES 31

FIG. 18. Scaly-throated Honeyguide (upper left) causing Lesser Honeyguide (lower right) to flee from near hive. actively attacking and supplanting them. Al- Greater Honeyguides approached the ex- though Lesser Honeyguides frequently were perimental site more often (figs. 20-21) than noted about the beeswax site, they were kept did Lesser Honeyguides, probably because from it for over ten days by Scaly-throated their habit ofperching conspicuously on dead Honeyguides; one Lesser Honeyguide man- branches or the tops of trees generally kept aged to dart in and secure wax only twice. them from conflict with the Scaly-throated On one occasion three Scaly-throated Hon- Honeyguides. The latter, more numerous, eyguides were actively interacting in the can- stayed inside the trees and at lower levels, opy over the hive, and a Lesser Honeyguide only occasionally venturing into the open. At perched and fed for 30 seconds on the bare least three Greater Honeyguides, an adult comb before being driven away by one ofthe male, an adult female, and a subadult male Scaly-throateds. On the other occasion a were seen in trees near the experimental hive Lesser Honeyguide darted to the honeyed, over a ten-day period. The subadult never brown comb bits and managed to grab sev- ventured into the hive tree, where Scaly- eral pieces before a Scaly-throated Honey- throateds usually were present, and it was guide present at the (bare) wax threatened it, chased from a nearby tree by one Scaly- causing the Lesser to fly off. Several chases throated. The adult Greater Honeyguides did of Lesser Honeyguides by Scaly-throated go to the hive tree, but frequently were at- Honeyguides near this site also were noted; tacked and driven from the vicinity. On only when being pursued, the faster Lesser Hon- four occasions did the (or an) adult male reach eyguide at least twice returned the attack, and the hive, and manage to eat some ofthe bare the two birds flew off calling, engaged in a wax comb. The (or a) female entered the hive flying encounter. twice, once going briefly to the bare comb, 32 AMERICAN MUSEUM NOVITATES NO. 2825 and once, harried by a Scaly-throated, to the to 3 notes at variable intervals. The second, brown, honeyed comb. In the hive tree we chattering call (fig. 6F, H) of these two calls observed ten supplanting attacks by several is somewhat like the guiding chatter of the Scaly-throated Honeyguides on the male Greater Honeyguide (L indicator), but does Greater Honeyguide, all of them successful. not appear related to honey-guiding activity. Likewise, the six attacks on the (or a) female A third, less frequent aggressive note is a more Greater Honeyguide by a Scaly-throated discrete, formed double "tza-dat" call (fig. Honeyguide were successful. On a few oc- 6I), with a buzzy beginning, dropping from casions the Greater Honeyguide returned to an initial peak, breaking and rising to 1 to a nearby tree after being driven off, but only 1.3 kilohertz, followed by a clearer second to perch and watch. On only one occasion element ending in a sharp peak at 1.7 kilo- did we see an attack by a Greater Honeyguide hertz. These show overtones, and are 0.18 to on a Scaly-throated Honeyguide-in this case, 0.2 second in duration, uttered in series of a female Greater flew at a subadult Scaly- about 2 per second. throated perched beside the hive, but the lat- A whistling call of 1-6 notes at about 2 per ter held its position, smashed at the female second, and easily mimicked, is a "foyt-foyt- Greater with bill and wings, and caused the foyt-." Its function is unclear, although hu- female to fly away. These data strongly sug- mans whistling it can attract Scaly-throated gest that I. variegatus is dominant to L in- Honeyguides. The song, or song trill (fig. 1 9A, dicator, females of which may be more ag- B), is a churring series that varies even in the gressive than are males (see account of L same bird from a froglike croaking sound to indicator). a loud, vibrant police-whistle-like series ACOUSTIC DISPLAYS: Young utter a low, (Friedmann, 1955). The song lasts 2 to 4 sec- husky "chess-chess-chess-" call with notes at onds, and commences with longer notes ut- four or five per second (Dowsett-Lemaire, in tered at a rate of 18 to 20 per second, then litt.; 1983), and when near fledging, a trill, changes over, becoming louder, giving way "foy-foy-foy-" (Dowsett-Lemaire, 1983, p. to a rapid trill at 40 to 42 notes per second. 72). Birds held in the hand after capture in Songs of 3.28 to 3.38 seconds in duration nets utter a sharp squawk. Various interactive contained 1 5 to 121 notes (central Kenyan calls have been recorded on tape, as when birds). This song is uttered from beneath the two birds (ofunknown sex) have an encoun- canopy oftrees or bushes, and usually is con- ter at a honeycomb source (fig. 6E). One in- sidered to be delivered from a very specific teractive aggressive note is a buzzy "zh-zh- song perch (Friedmann, 1955; Ranger, 1955). zh" or "tzha-tzha-" with sound concentrated We have observed a singing Scaly-throated at 5 to 8 kilohertz. Sonagraphically the notes Honeyguide more or less following several are unformed, with noise over a wide spec- Nubian Woodpeckers (Campethera nubica) trum of pitch. The notes usually last 0.5 to about, and singing from seven perches, in- 0.1 second with variable intervals of0.02 sec- cluding one on a herbaceous plant a meter ond and more between notes, and are uttered from the ground at the edge of a woodland. at up to 10 per second. Occasionally the sound Playback also tends to draw singing birds is continuous, a "zzzzh" over 0.25 to 0.35 closer to the playback recordist, the honey- second, then breaking into shorter notes at guide singing at one site, moving, singing at irregular intervals (fig. 6G). Even shorter "dz" another site, and so on, until singing over- notes in series are but 0.01 to 0.02 second in head. It is inconceivable that these sites all duration, given in bursts at 13 to 20 notes represent distinct song perches regularly used per second at the same range of pitch as the by territorial honeyguides. As in the case of "zh-zh" notes. These may speed up and con- the Lesser Honeyguide, the situation may vary tinue for 2 or more seconds, as a "dz-dz-dz- geographically in regard to song perches. As dz-dzzzzzzip." Intermediate "zh" and "dz" many as 76 songs may be given in 90 min- notes also occur. Interacting birds may utter utes, with singing particularly early in the either ofthese calls, or one may give "dz-dz- morning (even before daybreak), but also at dz-" calls continuously as the second bird intervals during the day (Ranger, 1955). repeatedly calls "zzha, zzha" in groups of 1 VISUAL DISPLAYS: These are poorly known. 1985 SHORT AND HORNE: HONEYGUIDES 33 8 A 81IF I 6 6 !II t It \1 '!I , 4 4 t 2 2 t i

0;5 1.0 1.5 2.0 G 1.0 1.5 2.0 8- B8 015 6- 6 , N Cr4- w 4 5 2- it 2- 2.0 Iv).0-0. -1- 1- 0.5 1.0 1.5 2.0 0.5 1.0 1.5 2.0 Z8- C j D 1!! &. H D 'III!! ir 6- b4 I tD}4ti$4F 4 li .3*t1t1 i *t i,; 4 w ~~~~~~~~~~~liii~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~ 2- '

0.5 1.0 1.5 2.0 0- 0.5 1.0 1.5 2.0 8] E 8-

6 X i1 6 I 4- 4-

; 1 Pi Pt 2- 41, 2- r) ..I .U. 0.5 1.0 1.5 210 V 05 I.0 1.5 2.0 TIME IN SECONDS FIG. 19. Vocalizations of Scaly-throated and Greater honeyguides in wide-band sonagrams. A, B. Full song (trill) of Indicator variegatus, Dec. 12, 1983 at 01 Ari Nyiro, Kenya. C. Peep call of Indicator indicator, August 1968, 01 Ari Nyiro, Kenya. D. Mixed piping and squeaking notes of L indicator in interactive trill, Dec. 17, 1983, 01 Ari Nyiro, Kenya. E. Three piping notes, two faint chatters, a noisy chattering note, and six squeaky piping notes during interaction at experimental site by male L indicator at L variegatus, Dec. 17, 1983, 01 Ari Nyiro, Kenya. F. Chattery trill form of guiding call of male I. indicator, showing only slight tendency to paired notes, Dec. 17, 1983, 01 Ari Nyiro, Kenya. G. Double- noted guiding call of male L indicator, August 1968, 01 Ari Nyiro, Kenya. H. Double-noted guiding call of male L indicator, Jan. 3, 1982, Akagera Park, Rwanda. I. Terminal two sets of "burr-wit" or "tor-vik" song of 10 sets by male I. indicator Aug. 26, 1980, Dzalanyama Mountains, Malawi.

Interacting Scaly-throated Honeyguides gape ters but at intervals as a honeyguide moves in a horizontal posture with head extended about, foraging or watching host species. at an antagonist; the wings and tail are flicked These honeyguides outfly most of their host frequently and there are side-to-side body and species, and often turn upon them when head movements as antagonists approach one chased, ending up perching or supplanting another. The throat feathers may be ruffled the hosts. and could serve to highlight the scaly throat HOSTS: The known hosts ofthis widespread in adults showing this feature. Flight displays honeyguide are surprisingly few. Verified are also occur, with fluttering-whether by ag- the Black-collared (Lybius torquatus) and gressive or courting adults is unknown. The Whyte's (Stactolaema whytil) barbets, and the tail is spread often, not only during encoun- Yellow-rumped Tinkerbird (Pogoniulus bi- 34 AMERICAN MUSEUM NOVITATES NO. 2825 lineatus) among capitonids, and the picid period of singing mainly is from August to Nubian (Campethera nubica), Golden-tailed December, with a declining amount of song (C. abingoni), Olive (Dendropicos griseo- thereafter, to April. Outside of southern Af- cephalus), Gray (D. goertae), and Cardinal rica the singing period is much shorter, and (D. fuscescens) woodpeckers (Friedmann, more variable. Furthermore, even within 1955, 1958a; Dowsett-Lemaire, 1983). The southern Africa songs sometimes are uttered record of the tinkerbird, which excavates a from perches other than the traditional ones, cavity so tiny as to make entry of a female and in Kenya moving birds may sing from Scaly-throated Honeyguide very difficult at diverse perches, even in bushes in grassy areas best, is accepted from van Someren's notes out ofwoodlands (personal observ.). Singing (in Friedmann, 1955, and especially Fried- may come from two birds as close to one mann, 1958a, p. 311, contra Dowsett-Le- another as 230 to 250 m. maire, 1983, p. 74). Excited, repeated re- It is presumed that females come to the sponses of a Scaly-throated Honeyguide to males' calling or stud sites (Friedmann, 1955). playback ofthe voice ofthe Green Tinkerbird There are no observations of courtship, nor (Pogoniulus simplex, Short and Home, 1979, has copulation been observed. pp. 3-5) reinforce the van Someren obser- Scaly-throated Honeyguides that are at- vations, and indicate that this honeyguide tracted to sources ofbeeswax obviously cross somehow manages to get its egg into tink- any existing territorial borders, since one can erbird nests (see below). observe up to seven about such a site (per- We consider as unsubstantiated the reports sonal observ.). The fact that there appears to ofthese hosts (Friedmann, 1955): Horus Swift be a dominance hierarchy at feeding sites, (Apus horus); White-cheeked Barbet (Stac- and that immature birds achieve some suc- tolaema leucotis); Crested Barbet (Trachy- cess in staying close to these sites without phonus vaillantii); and Yellow-throated Pe- being driven away suggest that the hierarchy tronia (Petronia superciliaris). The two may be sex-related and related to "territory," barbets among these are likely to prove hosts that is, local, perhaps territorial, individuals ofthis honeyguide. Our observations ofplay- may be dominant, although not so dominant back responses of L variegatus to voices of as to maintain exclusive control over the food the Green Tinkerbird, Green Barbet (Stac- source. tolaema olivacea, see Short and Home, 1979, Singing males (determined by collecting) p. 5), Brown-breasted Barbet (Lybius mela- actively follow vocal, interactive (e.g., males nopterus), Green-backed Woodpecker (Cam- interacting with females, and even adult-ju- pethera cailliautii), and Bearded Woodpeck- venile interactions) individuals of host er (Dendropicos namaquus) suggest these species. We watched one male singing in spo- species as possible hosts. radic bursts following a pair of Nubian It is clear that, as with the Lesser Honey- Woodpeckers that were accompanied by a guide (Indicator minor) and unlike the Great- noisy, begging young woodpecker. The hon- er Honeyguide (L indicator), the preferred eyguide sang in view of the woodpeckers, hosts of L variegatus are barbets and wood- moved with them, and came to our playback peckers. There are no data on host special- oftheir interactive calls. We obtained such a ization by individual female Scaly-throated male Scaly-throated Honeyguide that proved Honeyguides. to have large testes, after it followed Nubian TERRITORIALITY AND BREEDING HABITS: Woodpeckers, and just after it had sung over Details concerning territoriality and its mat- us in response to our playback of Nubian ing system are known only from some South Woodpecker calls. A Lesser Honeyguide (I. African studies (Friedmann, 1955; Ranger, minor) had followed the same woodpecker 1955). Presumed males maintain traditional pair, even more closely than had the Scaly- (long-term, occupied for as long as 26 years, throated Honeyguide (in fact, minor flew with, although not likely by the same bird) song was attacked by, and attacked the wood- perches on branches beneath the canopy of peckers several times as it placed itselfbeside trees, from which they sing at intervals over them). The Scaly-throated Honeyguide once the breeding season. In southern Africa the attacked the Lesser Honeyguide and chased 1985 SHORT AND HORNE: HONEYGUIDES 35 it, and was chased by it in return. We suggest (and probably variable) how long the juvenal that at least some breeding male Scaly- plumage is retained, but this is not apt to be throated Honeyguides may have a territorial longer than three to four months after hatch- system that is seasonal, and based upon ing. Southern Africanjuveniles date from Oc- breeding pairs of its hosts that are defended tober to February. Such birds are known as from other Scaly-throated Honeyguides, as follows: December, March, and June in in the case of Lesser Honeyguides, for which Ethiopia; February and March in Sudan; Feb- we have more evidence (Short and Home, ruary and May (nestlings) and April, June, 1983). July, August, and November in Uganda; June There are no data on the pair relations and to July nestlings, May to June fledglings, and activities of the female Scaly-throated Hon- others from all months except August in di- eyguide in entering the cavity nests ofits hosts. verse parts of Kenya; July, August, Novem- Dowsett-Lemaire (1983) found that scattered ber, and March in northern and central and apparently female Scaly-throated Honey- February in southern Tanzania; and January, guides occurred high on Malawi's Nyika Pla- April, October, and December for Zaire. The teau, laying in nests of Olive Woodpeckers molt occurs late in the breeding season and in areas where no songs were heard or singing immediately thereafter, and all data that are posts of male honeyguides were located. Fe- available are in accord with the seasonality males are not known to remove or destroy indicated by the laying, breeding, and young eggs of the hosts, although Dowsett-Lemaire of this honeyguide that have been discussed. (1983, p. 74) noted one possible such case. The incubation period is calculated to be Occasionally this honeyguide (same individ- approximately 18 days by Dowsett-Lemaire ual?) may lay more than one egg in a host's (1983). The young honeyguides have the typ- nest, and there even is a report (Friedmann, ical Indicator bill hook, and they usually, if 1955) of a Whyte's Barbet nest bearing three not always, destroy the eggs or occupants of eggs ofthat barbet and, supposedly, three eggs the host's nest (the one supposed record of a ofthe Scaly-throated and one egg ofthe Less- young ofthe host, an Olive Woodpecker, with er honeyguides. At least two eggs and prob- a young Scaly-throated Honeyguide is subject ably more are laid by a female. A continu- to doubt, see Sclater and Moreau, 1932; ously chattering Scaly-throated Honeyguide Friedmann, 1955; and Dowsett-Lemaire, perched with tail clinging to the nest entrance 1983). of a pair of Olive Woodpeckers may have Young are fed insects and (by barbets) fruit, been attempting to lay (Dowsett-Lemaire, the foods normally fed to their young by the 1983). foster parents. Feeding occurs up to 45 times Laying occurs in May in Kenya, in Sep- in eight hours (Dowsett-Lemaire, 1983). The tember to October in Malawi, in October in young honeyguides, according to that author, Zambia, in September to January in Zim- fledged in 27 to 35 days, and once leaving babwe, in December in Transvaal, and in Oc- the nest, the young honeyguide does not reen- tober in Natal (Benson et al., 1971; Benson ter it. Rather, it perches and begs from a site and Benson, 1977; Stuart Irwin, 1981; Brown in a small area about the nest, in the case of and Britton, 1980). Other indications of Olive Woodpeckers as the host. Dowsett-Le- breeding we have gleaned from specimens maire (1983) noted the efforts of the wood- and the literature suggest breeding in Novem- pecker foster parents to get the young hon- ber and January in Ethiopia; February in Su- eyguide back into the nest to roost for the dan; September and February in Uganda; night, without success. That author failed to May, June, and January in Kenya; August to locate young Scaly-throated Honeyguides October in Mozambique; May and July in being fed beyond the day of fledging, which, eastern Zaire and October to December in with our observations of the Lesser Honey- southern Zaire; April and July to October in guide, suggests that newly fledged honey- Malawi; February and June in Angola; and guides very rapidly, if not instantly, become October to December in South Africa. Re- independent upon fledging. Nevertheless, flecting earlier breeding are the many records fledged Scaly-throated Honeyguides have ofjuveniles; we emphasize that it is uncertain been observed being fed by Whyte's and 36 AMERICAN MUSEUM NOVITATES NO. 2825 I

FIG. 20. Adult male Greater Honeyguide (In- dicator indicator) above hive, looking up at Scaly- throated Honeyguide.

Black-collared Barbets; Yellow-rumped FiG. 21. Adult male Greater Honeyguide Tinkerbirds; and Gray, Olive, Bearded (Tay- trying, without success, to reach experimental hive lor, 1983), and Cardinal woodpeckers. Van that was occupied by Scaly-throated Honeyguide. Someren (1956) reported the enlargement of the nesting cavity by a pair ofYellow-rumped Tinkerbirds hosting a young Scaly-throated attacking and pursuing one another, often to Honeyguide, apparently in order to enable the advantage of the juvenile that was able the fat young honeyguide to leave the nest to get to the hive, and make off with some (its mother obviously must somehow have wax in the absence of adults. entered the tiny hole leading into the cavity). The only information available on behav- Indicator indicator (Sparrman) ior of independent juvenile Scaly-throated Greater Honeyguide (figs. 20-21) Honeyguides comes from our Kenyan studies ofhoneyguides at beeswax sources. We found RANGE SUMMARY: This honeyguide occurs that the single Scaly-throated Honeyguide in from the trans-African Sahel belt (southern juvenal plumage among the dozen or so that Mauritania to northern Ethiopia) south to frequented the beeswax was much more suc- northeastern Namibia, northern and eastern cessful than most of the adults in remaining Botswana, and the Cape Province and Natal, close to the beeswax, if not in feeding on it. South Africa, avoiding true deserts, extensive That is, its sporadic begging behavior, grasslands, and forests such as the main mass crouching in submission, and probably its ofthe Congolese forest (it does not reach the feather markings enabled it to bypass the forested coast of West Africa). Its behavior dominance hierarchy established among the and use of conspicuous perches make it the adults. It was allowed to approach adults more best known of the honeyguides. closely than were other adults, and even to HABITAT: Greater Honeyguides frequent perch and feed at one end ofthe experimental bushlands, wooded grasslands, and open hive containing beeswax while another adult woodland, and to a lesser extent, forest edges, was present, much less often eliciting an at- dense woodlands, riverine woods, and scrub tack and pursuit by the adults than occurred desert. They are more common in open, dry when one adult tried to approach the hive situations with scattered trees than in forest- while another adult was present. Also, when ed or densely wooded areas. Where the hab- three or four birds, including the juvenile, itat is open the species occurs up to 3000 m were present and interacting near the hive, in elevation (East Africa, Britton, 1980), but the juvenile largely was ignored, the adults it is more common below 2000 m. It does 1985 SHORT AND HORNE: HONEYGUIDES 37 not shun habitation, but occurs in cultivated guides pose special problems ofentry for the regions and even in eucalyptus groves. Be- bird is unknown. Clearly the Greater Hon- cause beeswax is a favorite food, it is apt to eyguide, like all its congeners, must locate be found where bees are abundant. active and inactive hives on its own, to obtain FORAGING HABITS: The Greater Honey- wax. It may be that the large size ofthe Great- guide is the only regularly guiding member er Honeyguide makes it difficult to gain entry of its family, attracting attention of people to some hives (however, the much more se- and certain other mammals, mainly the Ratel cretive and usually nonguiding L variegatus (Mellivora capensis), and guiding them to-ac- is virtually the same size). Or, it may feed to tive beehives. This well-known behavior is a greater degree than other congeners on the at least partly influenced by learning, for in larvae and eggs ofbees, requiring active hives. suburban and village areas where people no In addition to beeswax, and the bees, their longer seek honey from the hives of wild eggs, and larvae that are taken accidentally bees, the Greater Honeyguides infrequently or by choice when it feeds on the wax comb, or never guide, becoming less conspicuous the Greater Honeyguide flycatches from thereby (Friedmann, 1955). The guiding has perches to secure insects, probably its pri- been described in great detail by Friedmann, mary food. Honeyguides may participate in and we only summarize it here. One first no- interspecific foraging flocks that are a com- tices a persistent chattering call and notes the mon feature in African woods. Several Great- bird, which may be of either sex (immature- er Honeyguides may associate in these flocks, plumaged birds guide as well). Its tail flits and in flycatching, as well as gathering at wax apart and the bird bobs, looking at the pro- sources; immature birds especially are liable spective honey-seeker. It then moves in un- to form loose associations. At wax sources dulating flight, fluttering as it flies a variable we have noted aggression, adult females being distance in a certain direction (toward a hive), dominant to adult males. The honeyguides perches again, and incessantly calls. If fol- may forage about large grazing and browsing lowed the bird repeats the guiding a number mammals, flycatching for insects that are dis- of times until, perhaps a kilometer or more turbed by the mammals. We also have seen from the starting point, the hive is at hand. them follow woodpeckers foraging in trees, The honeyguide's final flight to the imme- flying down to pick offthe insects flushed by diate vicinity of the beehive is followed by the woodpeckers' movements. Their occa- its silent scrutiny-it depends upon the per- sional, reported feeding on eggs ofother birds son (or mammal) being guided to observe the (Friedmann, 1955) may be the reason why bees and to attempt to get the honey. While they sometimes are mobbed by small birds the activity goes on the bird perches quietly, that are not their hosts. or may move about to alternate perches. Tra- FoODS: The main foods are insects includ- ditionally tribal peoples leave a piece ofhon- ing diverse larvae, beetles, ants, termites, the eycomb conspicuously placed out for the pupae, larvae, eggs and adults of bees; and honeyguide, which moves to the hive and beeswax. A. W. Diamond (in litt.) main- commences feeding when the honey-hunters tained captive Greater Honeyguides for up have gone. Ifno one follows the bird initially, to six months on a diet ofchopped meat and the honeyguide may call at frequent intervals, hard-boiled eggs. In feeding trials with hon- for up to 5 minutes constantly at one burst. eycomb, Dr. Diamond found the captive birds Its attempts may last an hour or more, as it first sought out larvae of the bees, then ate moves from tree to tree around one (personal the wax. The honeyguides on occasion eat observ.). eggs of other birds. The young honeyguides Obviously the honeyguide must have are fed on insects, fruits, and the other foods knowledge of the location of one or several normally fed to their own nestlings by their active beehives. No one has described how foster parents. It has been assumed that the the birds find a hive, but visual clues (bees digestion of wax by the Greater Honeyguide moving in numbers, or a mass ofbees in one is through the means ofaerobic lipolytic bac- place) must play a major role. Whether or not teria (Friedmann and Kern, 1956), although the hives to which the Greater Honeyguide the action of aerobic bacteria in the confines 38 AMERICAN MUSEUM NOVITATESNNO. 2825 of the digestive organs poses some problems indicator over male indicator over subadult according to Dr. Diamond. Enzyme action indicator). Such behavioral differences may or the action ofcerolytic yeasts are other pos- be seasonal, or geographical, as Friedmann's sibilities for digestion ofbeeswax by the hon- studies largely were in southern Africa. eyguide (Friedmann, 1955). It is certain that ACOUSTIC DISPLAYS: The young Greater the honeyguide cannot exist for long on a diet Honeyguides utter a series of chattery notes solely of beeswax, as in the case of Lesser that are loud and irregular, bearing resem- Honeyguides that died in about one month blance to the guiding call. They pull the head offeeding only on beeswax (Friedmann, 1955, back and tilt the bill upward as they call at p. 75). 5 to 10 notes per second. Fry (1974) consid- INTERSPECIFIC BEHAVIOR: Because of their ered these calls to be vocally convergent upon conspicuous perching and flycatching, as well those of bee-eaters (specifically Merops bul- as their aggressiveness, Greater Honeyguides ocki), but this honeyguide has many, diverse frequently have aggressive encounters with hosts, and the voice of its young is not apt nonhost species. The African Drongo (Dicru- to resemble convergently any one more than rus adsimilis) may attack a perched Greater others. A. W. Diamond noted (in litt.) that Honeyguide, as may rollers (Coracias spp.), the begging call of one honeyguide sounds and almost inevitably the Greater Honey- like two or more calling individuals, and the guide returns the attack (see fig. 6J). It does loud calls may elicit feeding at greater rates not readily relinquish a perch unless a bird than a lone nestling otherwise might obtain. is much larger (e.g., a hornbill, Tockus spp.), Mechanical sounds include wing noises or indeed is a predator. The Greater Hon- made during presumed courtship flights, so- eyguide sometimes approaches a recordist called drumming, and a "brrr" sound made playing back the voice of the Lesser Hon- by a male near a female. The drumming eyguide (personal observ.). It may briefly sound, likely made with the wings, occurs as chase a Lesser Honeyguide that it encounters, a male circles back and forth, dipping as it the latter being faster and more maneuver- circles, giving a "Bvooommm" or "whoosh" able. At sources of beeswax we have noted sound, followed after a pause by another, and that it consistently is dominant to smaller so on (see under Visual Displays below). The honeyguides, such as I. minor, but invariably tail is closed when this flight display occurs, is driven away, supplanted, or kept from wax and the display itself has not been observed by Scaly-throated Honeyguides. We often saw frequently (see Friedmann, 1955). Courting Greater Honeyguides simply avoid proxim- individuals also snap the bill occasionally. ity to I. variegatis and the wax when the latter A squeaky, short note (Short and Home, was present. At our Kenyan study site (see 1979, fig. 2) forming a peak sonagraphically above discussion of I. variegatus) we ob- and having overtones is an interactive, ag- served ten supplanting attacks ofI. variegatus gressive call similar to squeek calls of con- on males of I. indicator (figs. 20-21), six of geners. This call, rendered "tych," "tyek," or variegatus on females ofindicator, one ofvar- "yeh" is uttered in conflicts with other species iegatus on a subadult male of indicator, and (e.g., Green Barbet, Stactolaema olivacea), one attack ofa female indicator on a subadult and may be heard between a male and female variegatus-this last failed, the variegatus Greater Honeyguide. In figure 6J are shown holding its position, then chasing the indi- a series of squeaky notes uttered by a male cator away (personal observ.). It should be I. indicator chasing an African Drongo. A noted that Friedmann (1955, p. 35) found no single piping, peep note with a peak at 4 kilo- possessiveness in honeyguides he studied, hertz, given in 0.04 second also is heard dur- mentioning the coming together of several ing interactions (fig. 1 9C). This resembles the honeyguides, even of I. minor with I. indi- "vik" or "wit" note of the song (see below). cator, without noting aggression. Our obser- Irregularly given in series of five or so per vations, in contrast, show a strict dominance second, these piping notes mark encounters hierarchy ofhoneyguide species (I. variegatus of some intensity, such as one with Green over I. indicator over I. minor), and of the Barbets (see Short and Home, 1979, fig. 1). sexes and age groups intraspecifically (female A more aggressive, trilling series ofnotes (fig. 1985 SHORT AND HORNE: HONEYGUIDES 39

1 9D) combine the form ofsqueak and piping Ranger's (1955, p. 72) observations suggest notes, given at 12 to 14 notes per second. that the song regularly starts with a soft "burr" These somewhat resemble the interactive and ends in a "wit." The "wit" is sharp and chattering calls and the guiding call (see be- fast, at 3 to 3.5 kilohertz, and often rises over low), but the notes are more piping with a its 0.02 to 0.04 second duration. The slightly peak. Possibly this is the "screaming-chat- lower pitched, buzzy "burr," 0.1 to 0.22 sec- tering" that Ranger (1955, p. 78) heard from ond in duration, consists of a long initial and a copulatory pair. Other interactive series two to six following elements, lending em- combine features of these various notes (fig. phasis to the sharper "wit," which follows 19E). Hissing notes that we have not heard the "burr" in 0.25 to 0.38 second (the next are reported from intensely aggressive birds "burr" is about 0.4 second after the "wit," by A. W. Diamond (in litt.). and this timing varies more, so the "burr- The guiding call is a long, irruptive series wit" version more accurately describes the ofchattery notes, varying from a regular trill song). The calls are audible for up to 825 m (fig. 19F) to a less double-noted series (fig. (Ranger, 1955). A call takes six to 11 seconds, 19G, H). The more regular, noisy chatter about a second per pair of "burr-wit" notes, seems to us to occur in the presence of a with six to 11 pairs per song. During the peak second honeyguide, or when the calling bird period of singing the song is uttered at about is disturbed (as near our experimental bees- one minute intervals almost continuously for wax site in Kenya). Notes are short (0.01 to several hours, especially in the morning, but 0.015 second) with a dropping tendency but also in the afternoon and sporadically through much noisy sound at varying frequencies, de- the day. There may be long pauses when the livered at 12 to 14 per second. The double- male shifts calling sites. Late in the season noted chatter is more typical ofguiding calls, the calling intervals diminish and ever greater but these may vary with the intensity of the intervals pass without songs, until they cease honeyguide's reaction to prospective follow- altogether (Ranger, 1955; see below). ers over time. At 14 to 16 notes per second VISUAL DISPLAYS: These need thorough the pairs of notes consist of a longer, usually study, especially in view ofthe marked sexual louder, first note, and a shorter, softer second difference in plumage that is not found in note, imparting the insistent quality to the other honeyguides. Aggressive birds hunch guiding call. There may be short bursts of the back with (golden) leading edges of the notes for five or six seconds, followed by a wings raised, the neck arched and the bill pause, or a bird may call continuously for open. In more intense aggression the hon- several minutes. The call may be rendered eyguide stretches its open bill at an antagonist "churra-churra-churra-churra-" for the dou- and hisses (A. W. Diamond, in litt.). A female ble-noted version, or "chur-chur-chur-chur- soliciting copulation near a male's song perch " for the simpler chattery trill. Males, fe- flicks its wings about, and its tail as well males, and juveniles may give the guiding (Friedmann, 1955, p. 128; Ranger, 1955, p. call, and guide. 78). It then stretches forward, crouches, and The song, variously termed the "vik-tor," the approaching male alights on her, stretch- "wit-burr," or "wit-cheer" call, is uttered so ing his chin and neck onto her head and cop- far as is known by males, often from tradi- ulating with wings fluttering and her tail tional song perches (see territoriality below). turned to the right (Skead, 1951; Friedmann, Most songs we have heard end in a "wit" or 1955). The male fluffs its crown feathers at "vik" note (fig. 191), as Friedmann (1955, p. times, puffs its body, arching it, and moves 121) suggested, but we have not analyzed a about the female with beating wings and sufficient number offully complete songs (the spread tail, as in turkeys (Meleagris gallo- initial note, probably usually a "burr" or pavo). "tor" note, is soft and the song is uttered at Ranger (1955, p. 78) described a copula- 40- to 80-second intervals, hence it is difficult tion differing in that the male seemed to to record its start) to state that the full song grasp the upper neck of the female, one or is indeed a "tor-vik" or "cheer-wit" rather both then gave a screaming-chattering call than the usually ascribed reverse. Indeed, throughout the end of copulation, they then 40 AMERICAN MUSEUM NOVITATES NO. 2825 flew in a chasing flight, and there followed form species, and 13 passerines (especially two "drumming" flight displays (see discus- swallows and starlings, which nest in holes, sion above under Vocal Displays) by the male or in covered nests, or in mud nests in dark over the area. places). The Hoopoe (Upupa epops) is a com- The notes of A. W. Diamond also are of mon host. Bee-eaters include: the Cinnamon- great interest regarding breeding and display, chested (Merops oreobates), Swallow-tailed and we paraphrase from his unpublished notes (M. hirundinaceus), White-fronted (M. bul- on observations of October 30, 1977, in Ke- lockoides), Carmine (M. nubicus), Little (M. nya: Male honeyguide began singing about pusillus), Little Green (M. orientalis), Mad- 10:15. At 12:05 I went to watch him, and agascar (M. superciliosus) and Bohm's (M. first saw a female on a branch of an Acacia boehmi). The Abyssinian Roller (Coracias xanthophloea. She kept up a constant wing- abyssinica) is the only coraciid (Parker, 1966). flicking, also raising and slightly spreading There are three wood-hoopoes (Scimitarbill, the tail, but not showing much of its white Phoeniculus cyanomelas, Abyssinian Scimi- edges. At this point the male was about 4 m tarbill, P. minor, and Green Wood-Hoopoe, from her, perhaps 2 m above her. The singing P. purpureus), and four kingfishers (Pygmy, male raised its head, stretched and inflated Ispidina picta, Chestnut-bellied, Halcyon its neck, with bill at a 60 degree angle to the leucocephala, Striped, H. chelucuti, and horizontal, and open. In about 5 minutes the Brown-hooded, H. albiventris). Definite bar- male flew to the female, tried landing on her bet hosts are the Pied (Tricholaema leuco- back, but sat upon her head and she flew to melaina), Black-collared (Lybius torquatus), about 10 meters away. Another male singing, and Crested (Trachyphonus vaillantii). female wing-flicking bout ensued, ending with Woodpeckers are the Rufous-necked Wry- another attempt at copulation, the male peck- neck (Jynx ruficollis), Nubian Woodpecker ing at the female's nape; the copulation ap- (Campethera nubica), Golden-tailed Wood- parently was unsuccessful. Again, after 5 pecker (C. abingoni), Knysna Woodpecker (C. minutes, this was repeated, the male finally notata), Tullberg's Woodpecker (C. tullber- landing atop the female, gripping her back gi), and Gray Woodpecker (Dendropicos with his bill, and staying on her for 10 to 15 goertae). Other groups represented by more seconds until she apparently lost her balance; than one species are: swallows, White-throat- with some squawking notes both fell, locked ed (Hirundo albigularis), Rufous-chested (H. together, into long grass below the tree. After semirufa), Larger Striped (H. cucullata), and 30 seconds the male flew up from the grass Banded Martin (Riparia cincta); starlings, into the air, then flew in a circle about 50 Pied (Spreo bicolor), Red-shouldered Glossy meters in radius, very deliberately, making (Lamprotornis nitens), and Blue-eared Glossy loud "whoosh, whoosh" noises with his wings. (L. chalybeus); thrushes, Cape Anteater Chat He did this three times before flying into (Myrmecocichla formicivora) and Anteater another tree, giving me a good view from Chat (M. aethiops); and sparrows, Grey- below-the noise definitely came from the headed (Passer griseus), and Yellow-throated feathers, and was not vocal. A few seconds Petronia (Petronia superciliaris). Other hosts later the female flew up and silently went off are the Black Tit (Parus niger) and Scarlet- in a direction away from the male, this at chested Sunbird (Nectarinia senegalensis). about 12:30. The Cardinal Woodpecker (Dendropicosfus- HoSTS: Definite hosts number 39 species, cescens) is a likely host. Other reported hosts, while evidence is unsatisfactory to verify re- records for which need corroboration (non- ports for 1 1 others. Most are hole-nesting birds, or build deep nests (the white eggs of hole-nesters are unlikely) are: Yellow-rumped the honeyguide would be conspicuous in Tinkerbird (Pogoniulus bilineatus), Red- shallow cup nests). Friedmann (1970) noted fronted Tinkerbird (P. pusillus), Cape Bulbul that 47 of 161 cases of which he knew in- (Pycnonotus capensis), Zanzibar Sombre volved bee-eaters, fewer than one-third ofthe Greenbul (Andropadus importunus), African total number. The hosts include 17 coracii- Drongo (Dicrurus adsimilis), Tropical Bou- form birds (eight are bee-eaters), nine pici- bou (Laniarius ferrugineus), Black-backed 1985 SHORT AND HORNE: HONEYGUIDES 41

Puffback (Dryoscopus cubla), Black-headed erwise, during laying. Rarely (two cases) are Oriole (Oriolus larvatus), Cape Rock Thrush two eggs laid in the same host's nest (Fried- (Monticola rupestris), and Violet-backed mann, 1955). Breeding occurs in the breeding Starling (Cinnyricinclus leucogaster). period of the host species. TERRITORIALITY AND BREEDING HABITS: Eggs are known: in July from Gambia; April Individual birds probably cover wide areas, to June and August to December (encom- and territoriality has generally not been es- passing one or both wet periods) in East Af- tablished. Even in southern Africa, where song rica (Brown and Britton, 1980); July and Sep- perches are traditional for 22 years or more, tember to October in Zambia (Benson et al., more than one male may use the same song 1971); September to November and January perch alternately and may sing simultaneous- for Zimbabwe (Stuart Irwin, 1981); Septem- ly from adjacent sites that both have used ber to October for Malawi (Benson and Ben- previously (Ranger, 1955). Little or no son, 1977); and September to January (main- aggression is evident from the literature, sug- ly November-December) for Natal, South gesting that territorial defense, ifthere is any Africa. Reported breeding, based upon adults territory, is minimal. However, at our Ke- in breeding condition, or other indications nyan experimental site we never observed are from: Gambia in July; Nigeria in January more than one adult male, one adult female, to July (Elgood, 1983); Cameroun in April; and a subadult male in the area at any given Sudan in March to April and October; Ethio- time. Furthermore, the female supplanted the pia in January to April; northern Zaire in male, and both kept the subadult male from January to May; southeastern Zaire in June, the vicinity ofthe beeswax. Although our ob- August, and October; Kenya in July and No- servations may not bear directly on territo- vember; Tanzania in July to January; Angola riality, there is good evidence for aggression in October; Zambia in September to October that is associated with such, and these hon- and March; Malawi in August to September eyguides are aggressive. A. W. Diamond (in and April; Zimbabwe from September to De- litt.) reports that caged and neighboring wild cember; northern Mozambique in January; males are regularly aggressive to one another. and South Africa, mainly from October to Each male has several calling sites in the December, but also in September and Jan- South African birds that Ranger (1955) and uary. Friedmann (1955) observed. Calling at these The very distinctive juveniles come from sites was regular from September to Decem- July and October to November (subadults) ber, with earlier singing in August, and ta- in Senegal; December in Ghana; April, June, pering off in January and February. Ranger and July (plus December subadult) in Nige- noted that, although there are primary and ria; December and March in Cameroun; Feb- secondary sites, the males may call from oth- ruary (plus January to April subadults) in Su- er than the main song perches. As discussed dan; February to April, June, and July, and under Visual Displays (above), it is to these August and September (plus December to song perches that females come to mate. March subadults) in Ethiopia; August to Sep- Ranger's evidence (1955) indicates that males tember, with a November subadult from So- mate with more than one female, and that malia; December and March, and June and females may go from male to male for breed- August (plus subadults for March, July, and ing. However, Roberts (1956) reported ob- September) in Uganda; all months ofthe year servations from Transvaal of males and fe- in Kenya, with a concentration in December males, apparently pairs, of I. minor and of I. to February, May to July, and September to indicator (in which the sexes are clearly October; December, and February to July in marked) acting in concert to enable the fe- Tanzania, with June and December sub- male to enter a host's nest. Thus, there may adults (in the dry periods); May to October be a longer pair-bond than indicated by in northern Zaire; January to March, May to Friedmann (1955) and Ranger (1955). July and December in southeastern Zaire The clutch size is four to as many as eight (Upemba specimens); December to April and glossy, white eggs. The laying female usually August, plus subadults in June and December breaks the hosts' eggs with her bill or oth- (mainly in dry periods) in southern Zaire; 42 AMERICAN MUSEUM NOVITATES NO. 2825

November, with November to January sub- The actual hosts are unknown for some hon- adults in Angola; December and April in eyguides. In particular, the behavior and Zambia; December from Malawi; February, ecology of species of Melichneutes and Me- April and September from Mozambique; De- lignomon, somewhat intermediate morpho- cember from Zimbabwe; July and November logically between Prodotiscus and Indicator, from Botswana; and August to January and are so little known that the evolution of ex- March, with subadults from November to tant honeyguides remains problematical. In May in Swaziland and South Africa. addition, the biology of the smaller species The hatchling honeyguide (the incubation (willcocksi, pumilio, meliphilus, and naro- period is uncertain) is equipped with trans- kensis) of Indicator is very poorly known. parent bill hooks that overlap, and although The feeding of honeyguides is more open the pinkish young are blind and featherless than ever for investigation, particularly the they bite any surviving young of the host, habits of locating and digesting wax. It is killing them. These hooks are lost at about abundantly clear that honeyguides and hon- 14 days of age (Friedmann, 1955; Ranger, eybirds seek wax from one source (beeswax) 1955). The young honeyguide begs and re- or another (insects' exudates). How do they ceives food from the foster parents, remain- find wax and what part does it play in their ing about 30 days in the nest. After leaving diet? Our knowledge ofhoneyguide digestion the nest the honeyguide calls loudly for food can hardly be said to be solidly based, for the but does not follow the foster parents, nor data are meager, and for but a few individuals does it return to the nest (Friedmann, 1955). of one species. Modern methods surely can There are few observations of feeding of lead quickly to breakthroughs in understand- fledged young, although they reportedly ing the digestion of wax, and the nature of (Friedmann, 1955) are fed for up to 10 days their seeming requirement for it. Honey- after leaving the nest. guiding itself, while intriguing, is far from The independent young retain the yellow- universal, and indeed only one species (L in- breasted and yellow-throated plumage for dicator) regularly guides-the honeyguides some time, perhaps to six months ofage. Sub- might be said to be misnamed, for "wax- adults, that is birds in partly adult plumage, birds" would appear to better reflect a trait but retaining the traces of yellow below or common to many, if not all, of them. (males) the blotchy black and white, or yel- A great deal has been accomplished in set- low, throat and blackish bill, may attempt to ting forth the hosts of the various honey- breed in this plumage (the golden wing mark, guides (although hosts are unknown for sev- the fully black throat in males, and the pink eral species). We are at a point where we bill in males, are the last colored areas to shift require more specific data. Does an individ- to the adult condition). Such subadults readi- ual female (or for that matter, males too) spe- ly guide to honey sources. In their juvenal to cialize in one host, or several? We have a list subadult plumages young birds may gather of hosts for some species, but only some in- in very loose feeding flocks about herds of dications of preferred hosts. There has been cattle or game animals, the main instance of an emphasis on observers finding new hosts sociality exhibited in the species. to add to the list for various honeyguides. It is time to concentrate more on the known, regularly used hosts, to ascertain their precise SUMMARY AND CONCLUSIONS ecological interrelations with the parasitic Data accumulated in the three decades since honeyguides. Do preferred hosts vary from Friedmann's (1955) monograph still leave place to place, or even from season to season many unanswered questions about the Af- in the same locality? These and many other rotropical honeyguides. Indeed, one gets the questions ought now to be posed, and an- impression that we are on the verge of learn- swers sought, especially since some host ing how these birds become independent, seek species, and thus perhaps indirectly the hon- food, recognize conspecifics, spatially organ- eyguides, are being affected adversely by hu- ize themselves, court, mate, select hosts, and man alteration of the African landscape on lay in hosts' nests at the appropriate time. an ever grander scale. 1985 SHORT AND HORNE: HONEYGUIDES 43

One new thrust has been in the wealth of in Scaly-throated and Greater Honeyguides, new information relating to honeyguides' be- is a new development bearing on aggression havioral relations with their hosts. These ap- generally in honeyguides, and on territorial- pear to be complex, at least in central and ity. We are at present expanding our exper- East Africa. The feeding of a young honey- iments to work with marked individuals. We guide by a conspecific adult (Brosset, 1981), hope to gain more information on how wide- and our observations of honeyguide activity spread the aggression is, and on the exact about recently fledged young white-eyes ac- nature of the hierarchy under various con- companying their parents suggest that even ditions. It is possible that the behavior of in the honeyguides, the forming ofpairs may conspecific honeyguides representing popu- be less briefand simple than has been thought. lations near the equator differs markedly from In the case ofseveral species ofIndicator there that of those from southern African popu- is considerable evidence that the honeyguides lations. The latter are subjected to very strict- carefully monitor their hosts, their hosts' ly defined seasons limiting breeding, and, in- nesting and roosting sites, the nesting of the sofar as hosts vary, there are fewer choices hosts, and the progress of the nesting even of host species for southern honeyguides. up to and beyond the point of fledging of Only with our studies (1979) and the re- the hosts' young, or of a young honeyguide. search ofChappuis (1981) has the analysis of Young Lesser Honeyguides may see and cer- honeyguide vocalizations really commenced. tainly often hear adults, perhaps their par- It appears that there are certain aggressive ents, calling near them. Whether or not this squeaky notes, trills, and chatters that have plays a role in facilitating learning of vocal- counterparts in several honeyguides. Of izations, or any imprinting effects remains to course the voices of some species are still be established. unrecorded on tape, and indeed there is no There is suggestive evidence from Lesser vocal (or visual) display information avail- Honeyguides that part of the interplay be- able for a few honeyguides, so we are only tween honeyguides and their barbet hosts be- beginning to evaluate such data for some yond the period of egg-laying by the hosts species. One problem that emerges is that of may relate to disruption ofthe nesting effort, the function and nature of song in the group. increasing chances ofrenesting, and thus pro- It appears that there are similarities in songs viding another opportunity for the honey- of many (e.g., Indicator narokensis, L will- guide to parasitize them. The number oftimes cocksi, L exilis, L conirostris, I. minor, and that two honeyguides have been seen about even L indicator) of the honeyguides, with a host's nest strongly suggests cooperative ac- those of Indicator variegatus and I. macu- tions that would enable the female to enter latus being divergent from others. The Lesser the host's nest. The seeking out ofthe barbets Honeyguide has an alternate song used more by a Lesser Honeyguide, eliciting chase after frequently in the tropics than is the primary chase, may be a ploy to get the barbets from song, which is so prevalent for much of the the nest to allow a female to enter. Males year in southern Africa. Correlated with this might even "play" at this to increase aggres- difference seems to be a difference in use and sive and chasing tendencies by the hosts at importance ofsinging perches in northern and times other than while actually laying, to al- southern regions. These differences may be low the drawing of the birds away from the related to the greater range of the breeding nest at a later time (the host's habituation period and the monitoring of host species to the honeyguide, resulting in its failure to that seem notable in more northern birds, but chase it, is the negative side of such activity has hardly been noted for the better known to the honeyguide). southern Lesser Honeyguides. Studies of the Until now there has been little appreciation Thick-billed Honeyguide, a tropical forest of the aggressiveness of honeyguides intra- species that sings virtually identically to the specifically, or with other species of honey- Lesser Honeyguide (to which it is very closely guides. The establishment of a dominance related, Friedmann, 1955), might aid in in- hierarchy based upon age and sex, at least at terpretation and understanding of the vari- certain times and under certain conditions, ation that seems to occur in the Lesser Hon- 44 AMERICAN MUSEUM NOVITATES NO. 2825 eyguide. The problems ofthe precise relations pp., 4 col. plates, 22 photographs, 4 among the taxa of Prodotiscus, and of the maps. smaller species ofIndicator, would be brought Britton, P. L., and D. A. Zimmerman much closer to solution if we had access to 1979. The avifauna of Sokoke Forest, Kenya. data on their acoustic displays. In any case, Jour. East Afr. Nat. Hist. Soc. and Natl. greater knowledge of the form and function Mus., no. 169, 15 pp. of honeyguide vocalizations is likely to help Brosset, A. 1981. Observation de l'indicateur parasite in understanding their apparently very com- Prodotiscus insignis nourissant unjeune plex biology. de son espece. L'Oiseau, vol. 51, pp. 59- In conclusion, we have made a number of 61. advances in our knowledge of honeyguides. Brown, L. H. Their population dynamics, ecology, repro- 1948. Notes on birds ofthe Kabaa, Ilorin and ductive behavior, and foraging habits remain N. Benin provinces ofNigeria. Ibis, vol. largely unknown. Even in areas of greatest 90, pp. 525-538. progress (in establishing the identity of host Brown, L. H., and P. L. Britton species, in aspects of the activities of young 1980. The breeding seasons of East African honeyguides with their foster parents and the birds. Nairobi, Kenya, East Afr. Nat. Hist. Soc., vi + 164 pp., 2 maps. young of the latter, and in honeyguiding by Carter, C. one species), we can only be said to have 1978. Eastern Least Honeyguide (Indicator arrived at the doorstep ofa mist-enshrouded meliphilus) at Ndola. Bull. Zambian Or- house whose form remains obscured. It is nith. Soc., vol. 10, pp. 30-31. hoped that our studies, the updating ofFried- Chapin, J. P. mann's (1955) landmark work, and the sum- 1939. The birds ofthe Belgian Congo. Part IV. marizing we have attempted will stimulate Bull. Amer. Mus. Nat. Hist., vol. 75B, further advances. pp. 1-846, figs. 1-46, plates 1-27. 1958. A new honeyguide from the Kivu Dis- trict, Belgian Congo. Bull. Brit. Ornith. Club, vol. 78, pp. 46-48. LITERATURE CITED 1962. Sibling species of small African honeyguides. Ibis, vol. 104, pp. 40-44. Archer, A. L., and R. M. Glen Chappuis, C. 1969. Observations on the behavior of two 1981. Supplement Sonore. Illustration sonore species of honey-guides Indicator var- de problemes bioacoustiques poses par iegatus (Lesson) and Indicator exilis les de la Zone (Cassin). Los Angeles Co. Mus., Con- Oiseaux Ethiopienne. trib. Sci., no. 160, pp. 1-6. Alauda, vol. 49, pp. 35-58. Bates, G. L. Clancey, P. A. 1930. Handbook of the birds of West Africa, 1975. Miscellaneous taxonomic notes on Af- London, J. Bale, Sons, and Danielson, rican birds. No. XLIII. Durban Mus. Ltd., xxi + 572 pp. Novitates, vol. 1 1, pp. 1-24. Benson, C. W., and F. M. Benson 1977. Miscellaneous taxonomic notes on Af- 1977. The birds of Malawi. Limbe, Malawi, rican birds. No. XLVII. Ibid., vol. 11, Montfort Press, 263 pp., 1 col. plate, 1 pp. 181-187. map. Colebrook-Robjent, J. F. R., and R. Stjernstedt Benson, C. W., R. K. Brooke, R. J. Dowsett, and 1976. Chaplin's Barbet Lybius chaplini: first M. P. Stuart Irwin description of eggs, a new host record 1971. The birds of Zambia. London, W. Col- for the Lesser Honeyguide Indicator mi- lins Sons and Co., Ltd., 414 pp., 8 figs., nor. Bull. Brit. Ornith. Club, vol. 96, 12 col. plates. pp. 109-111. Benson, C. W., and M. P. Stuart Irwin Colston, P. R. 1967. A contribution to the ornithology of 1981. A newly described species of Meligno- Zambia. Zambian Mus. Papers, no. 1, mon (Indicatoridae) from Liberia, West pp. 1-139. Africa. Ibid., vol. 101, pp. 289-291. Britton, P. L. (ed.) Dowsett-Lemaire, F. 1980. The birds of East Africa. Nairobi, Ke- 1983. Scaly-throated Honeyguide Indicator nya, East Afr. Nat. Hist. Soc., xiv + 271 variegatus parasitizing Olive Wood- 1985 SHORT AND HORNE: HONEYGUIDES 45

peckers Dendropicos griseocephalus in Louette, M. Malawi. Ibid., vol. 103, pp. 71-75. 1977. Die avifauna van Kameroen En Haar Elgood, J. H. Zoogeografische Interpratatie (partim 1983. The birds ofNigeria. Brit. Omith. Union Non-Passeriformes). Unpublished Check-list No. 4, 246 pp., 7 figs. Ph.D. thesis, Univ. Antwerp, vol. III, Forbes-Watson, A. 155 pp., 119 maps. 1977. Notes on the field identification of East 1981. A new species ofhoney-guide from West African honeyguides (Indicatoridae). Africa (Aves, Indicatoridae). Rev. Zool. Scopus, vol. 1, pp. 17-20. Africaine, vol. 95, pp. 131-135. Friedmann, H. Macdonald, M. A. 1955. The honey-guides. Bull. U.S. Natl. Mus., 1980. Further notes on uncommon forest birds no. 208, vii + 292 pp., 25 plates. in Ghana. Bull. Brit. Ornith. Club, vol. 1958a. The status of the gray-breasted least 100, pp. 170-172. honey-guides. Proc. Biol. Soc. Wash- Macworth-Praed, C. W., and C. H. B. Grant ington, vol. 71, pp. 65-68. 1970. Birds ofwest central and western Africa, 1958b. Advances in our knowledge ofthe hon- vol. 1. London, Longman Group Ltd., ey-guides. Proc. U.S. Natl. Mus., vol. xxvii + 671 pp. 108, pp. 309-320. Moyer, D. C. 1963. Morphological data on two sibling 1980. (On Lesser Honeyguide and Black-col- species of small honey-guides. Los An- lared Barbet.) Zambian Ornith. Soc. geles Co. Mus., Contrib. Science, no. 79, Newsletter, vol. 10, p. 159. 5 pp. 1968. Zenker's Honey-guide. Jour. fur Or- Nikolaus, G. nith., vol. 109, pp. 276-283. 1982. Further notes on some birds new to 1969. Phenotypic potential and speciation in South Sudan. Scopus, vol. 6, pp. 1-4. Indicator and Prodotiscus. Ostrich, Parker, S. A. Suppl. 8, pp. 21-26. 1966. Coracias abyssinica: an unrecorded host 1970. Further information on the breeding bi- of Indicator indicator. Bull. Brit. Or- ology of the honey-guides. Los Angeles nith. Club, vol. 86, p. 81. Co. Mus., Contrib. Science, no. 205, pp. Prigogine, A. 1-5. 1971. Les Oiseaux de l'Itombwe et de son 1978a. Results of the Lathrop Central African Hinterland. Mus. Royal de l'Afrique Republic Expedition 1976, ornithology. Centrale, Annales, Science Zool., ser. Ibid., no. 287, pp. 1-22. IN-80, no. 185, 298 pp., 16 figs. 1978b. Current knowledge of the Lyre-tailed 1978. Note sur les petits Indicateurs de la Fo- Honey-guide, Melichneutes robustus ret de Kakamega. Le Gerfaut, vol. 68, Bates, and its implications (Aves Indi- pp. 87-89. catoridae). Rev. Zool. Africaine, vol. 92, Ranger, G. A. pp. 644-656. 1955. On three species of honey-guide; the Friedmann, H., and J. Kern Greater (Indicator indicator), the Lesser 1956. Micrococcus cerolyticus, an aerobic li- (Indicator minor) and Scaly-throated polytic organism isolated from the Af- (Indicator variegatus). Ostrich, vol. 26, rican honey-guide. Can. Jour. Micro- pp. 70-87. biol., vol. 2, pp. 515-517. Ripley, S. D., and G. Heinrich Friedmann, H., and J. G. Williams 1966. Additions to the avifauna of northern 1971. The birds of the lowlands of Bwamba, Part II. Postilla, no. 95, 29 pp. Toro Province, Uganda. Los Angeles Co. Angola. Mus., Contrib. Science, no. 211, pp. 1- Roberts, H. A. 70. 1956. Notes biologiques sur l'Indicateur a Fry, C. H. queue-en-lyre: Melichneutes robustus 1974. Vocal mimesis in nestling Greater Hon- Bates. Ostrich, Suppl. 3, pp. 271-273. ey-guide. Bull. Brit. Ornith. Club, vol. Rougeot, P.-C. 94, pp. 58-59. 1950. Contribution a l'Etude des Indicato- Hoesch, W rides de la foret Gabonaise. L'Oiseau, 1957. Uber die Auswirkungen der Besiedlung vol. 20, pp. 51-63. auf den Vogelbestand in Suidwest-Afri- Sclater, W. L., and R. E. Moreau ka. Jour. fur Ornith., vol. 98, pp. 279- 1932. Taxonomic and field notes on some 281. birds of north-eastern Tanganyika Ter- 46 AMERICAN MUSEUM NOVITATES NO. 2825

ritory. Part II. Ibis, vol. 74, pp. 656- Stager, K. E. 683. 1967. Avian olfaction. Amer. Zoologist, vol. Short, L. L. 7, pp. 415-419. 1985. Neotropical-Afrotropical barbet and Stuart Irwin, M. P. woodpecker radiations: a comparison. 1981. The birds ofZimbabwe. Salisbury, Zim- In Neotropical ornithology, Ornith. babwe, Quest Publishing, xvi + 464 pp., Monographs, no. 36, pp. 559-574. 20 col. plates, 22 photos. Short, L. L., and J. F. M. Home Taylor, P. B. 1979. Vocal displays and some interactions of 1983. E. A. N. H. S. nest records scheme: 1982. Kenyan honeyguides (Indicatoridae) Scopus, vol. 6, pp. 129-142. with barbets (Capitonidae). Amer. Mus. Traylor, M. A., Jr., and A. L. Archer Novitates, no. 2684, pp. 1-19, figs. 1982. Some results of the Field Museum's 1-2. 1977 Expedition to South Sudan. Ibid., 1983. The relationship of male Lesser Hon- vol. 6, pp. 5-12. eyguides Indicator minor with duetting van Someren, V. G. L. barbet pairs. Bull. Brit. Ornith. Club, 1956. Days with birds. Fieldiana: Zool., vol. vol. 103, pp. 25-32. 38, 520 pp., 126 figs. Skead, C. J. Winterbottom, J. M. 1951. Notes on honeyguides in southeastern 1969. Lesser Honeyguide Indicator minor. Cape Province, South Africa. Auk, vol. Ostrich, vol. 40, p. 131. 68, pp. 52-62. Zimmerman, D. A. Snow, D. W. (ed.) 1972. The avifauna of the Kakamega Forest, 1978. An atlas of speciation in African non- western Kenya, including a bird popu- passerine birds. Brit. Mus. (Nat. Hist.), lation study. Bull. Amer. Mus. Nat. vii + 390 pp., 373 maps. Hist., vol. 149, pp. 255-340.

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Recent issues of the Novitates may be purchased from the Museum. Lists of back issues of the Novitates, Bulletin, and Anthropological Papers published during the last five years are available free of charge. Address orders to: American Museum of Natural History Library, Department D, Central Park West at 79th St., New York, N.Y. 10024.