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

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AMERICAN MUSEUM No Iltates PUBLISHED by the AMERICAN MUSEUM of NATURAL HISTORY CENTRAL PARK WEST at 79TH STREET, NEW YORK, N.Y 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 hon- eyguide (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 honey- guides 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.
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