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Cooperative Breeding by Rufous Hornbills on Mindanao Island, Philippines

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Cooperative Breeding by Rufous Hornbills on Mindanao Island, Philippines October1993] ShortCommunications andCommentaries 933 for granting us the use of the study area,the Campos MARTELLA, M.B. 1985. Observaciones sobre el com- family for their hospitality, and J. Navarro and H. portamientode la cotorraMyiopsitta monachus con Merlini for their help with the fieldwork. We appre- especialenfasis en la comunicacionsonora. Ph.D. ciatedthe helpful commentsof M. Notes, L. Baptista, thesis, C6rdoba Univ., C6rdoba, Argentina. E. Klaas,and E. Morton on the manuscript.This work MARTELLA,g. B., J. L. NAVARRO,AND E. H. BUCHER. was funded by the Consejo Nacional de Investiga- 1987. M6todo para la captufade cotorras(Myiop- clones Cientificas y T•cnicas of Argentina (PID sitta monachus)en sus nidos. Vida Silvestre Neo- 3-908101/85). tropical 1:52-53. NAVARRO,J. L., L. F. MARTIN,AND E. H. BUCHER.1992. LITERATURE CITED The use of remige molting for age-classidenti- fication in Monk Parakeets. Hornero 13:261-262. BROWN,J. L. 1978. Avian communal breeding sys- ROWLEY,I. 1983. Mortality and dispersalof juvenile tem. Annu. Rev. Ecol. Syst. 9:123-244. GalahsCacatua roseicapilla in the western Austra- BROWN,J.L. 1987. Helping and communalbreeding lian wheatbelt. Aust. Wildl. Res. 10:329-342. in birds: Ecologyand evolution. Princeton Univ. SAUNDERS,D. A. 1982. The breeding behavior and Press, Princeton, New Jersey. biology of the short-billedform of the White- EMLEN,S. T. Observations on a captive colony of tailedBlack Cockatoo Calyptorhynchus funereus. Ibis Quaker Parakeets. Watchbird. In press. 124:422-455. FORSHAW,J. M. 1978. Parrots of the world, 2nd ed. Landsdowne Editions, Melbourne. Received9 April 1992,accepted 22 November1992. The Auk 110(4):933-936, 1993 Cooperative Breeding by Rufous Hornbills on Mindanao Island, Philippines MARK C. WITMER Sectionof Ecologyand Systematics,Cornell University, Ithaca, New York 14853,USA Hornbills (Bucerotidae)are unique in their nesting a distinct casquemorphology and bill coloration (il- habits;females seal themselves within the nest cavity, lustrated in Hachisuka 1934). In all subspeciesthe where they lay the clutch and remain with the grow- sexes are similar and all show the same pattern of ing young for most or all of the nesting period. In plumage maturation (McGregor 1909). The juvenal somespecies the male helps with the sealingprocess plumage is distinct with white featherson the head, from outsidethe nestcavity. The nestlingsand female neck, chest,and belly. Subsequently,the pale juvenal are fed by the male through a narrow vertical slit in plumage is replacedby the dark black and chestnut the sealednest opening (Kemp 1979 and references body plumage of adults.The casqueand bill develop therein). Thus far, four speciesof hornbills have been more slowly; many birds have adult plumage color- found to have cooperative breeding systemswith ation but incomplete development of the casqueand helpersat the nest;nonbreeding birds assistthe male bill coloration. Maturation is probably complete in in feeding the nest mates(Stacey and Koenig 1990). about four years (Kemp 1979). Mindanao Rufous Theseare the Bushy-crestedHornbill (Anorrhinusgal- Hornbills develop from a black bill with a low, eritus;Madge 1969), Southern Ground Hornbill (Bu- humped casque in juvenals to an ivory bill with a corvuscafer [=leadbeateri; see Browning 1992];Kemp prominent, red, anvil-shaped casquein mature adults and Kemp 1980), White-crowned Hornbill (Aceros (see Hachisuka 1934). ! use "immature" to designate comatus;Leighton 1982), and Brown Hornbill (Ptilo- birds that have adult plumage coloration, but which laemustickelli; Poonswad et al. 1983).I report helpers have not yet developed the definitive adult mor- at the nest in another hornbill species,the Rufous phologyand colorationof the bill and casque("adult" Hornbill (Buceroshydrocorax mindanensis) on Minda- refers exclusivelyto thesebirds). nao Island in the Philippines. I observedbreeding activity at two RufousHornbill Each of the three subspeciesof Rufous Hornbill (B. nestson the island of Mindanao, Philippines: one at h. hydrocorax,Luzon and Marinduque;B. h. semigalea- Mt. Apo National Park near Davao City, Davao del tus, Bohol, Leyte, Panaon, and Samar; and B. h. min- Sur Province (7øl'N, 125ø22'E)from 31 July to 25 Au- danensis,Mindanao and Basilan[McGregor 1909])has gust 1984;and the other at Lake Sebu, South Cotabato 934 ShortCommunications and Commentaries [Auk, Vol. 110 Province (6ø12'N, 124ø41'E)from 4 to 14 August 1985. I beganobservations on 31 July and watchedthe nest Both sitesare approximately 1,000 m in elevation. The from a ground blind located20 m from the nest tree primary forest vegetation is tropical lowland ever- for a total of 62 h on 9 days (5-8 h per day) from 4- green rain forest (Whitmore 1984) dominated by trees 25 August.On 5 Septembera singlenestling fledged. in the family Dipterocarpaceae(Lewis 1988). I also The nest was a cavity about 14-m high in a 20-m-high recorded Rufous Hornbill group sizes at the Paper rotten tree trunk (102-cm dbh). I could see no seal at Industries Corporation of the Philippines (PICOP) this nest; the female had apparentlybroken out of logging concession (7ø50'N, 126ø14'E) during two the nest by the time I began observations,and the weeks in November and December 1986. The PICOP cavity was either left unsealedor I was unable to site is 600 m in elevation with vegetationof approx- discern remaining sealing material. Thus, I observed imately 20-year-oldselectively logged forest. thesehornbills during the final 45 days(nest discov- The Lake Sebu nest was found on 4 August 1985. ery to fledging)of a presumedfour-month nesting I opportunistically observedhornbill activities at and cycle(see Kemp 1979),during which time the female around the nest daily through 14 August; total ob- was out of the cavity and feeding the nestling. servation time was 12 h. The nest cavity was 30-m Three Rufous Hornbills tended this nest. Both high in the trunk of a huge emergent dipterocarp adults,presumably the matedpair, had blue-grayeyes. (40-m high, 220-cm dbh). This hornbill group was in These birds could be individually identified by dif- the initial phasesof breeding activity and included ferences in tail molt and bill coloration; one had a four adults and one immature (black bill and casque; short central rectrix and a dirty-white bill tip (adult the undevelopedcasque of this hornbill suggeststhat A), while the other had a complete tail with three it was an immediate postjuvenalbird). I was not able new feathersthat were lighter than the rest of the to individually identify the adults. tail and an ivory-white bill tip (adult B). The other An adult hornbill, probably the female, spent 49% hornbill seen at the nest was an immature bird, rec- of the total observation time (5.88 h) in the nest cham- ognizableby a dark-graybill tip and marooncasque. ber. This bird excavatedthe chamber by chiselling at This bird's casquewas a raised anvil-shaped protu- the cavitywalls and dumping out beak-fullsof wood berance,but it was not as prominent as in the adults; chips. Other group members, including the imma- its eyes were yellow-brown. ture, often perched outside of the cavity entrance, During the nine daysof observationI was able to facing the nest hole, and hammered at the outer rim recognizethe individual feedingthe nestlingfor 66 of the cavity opening. of 72 feeding visits:22 (33%)by adult A; 29 (44%)by The presumedfemale was fed by at leasttwo adults adult B; and 15 (23%)by the immature.The feeders seven times during the two days (10 and 11 August) tendedto arrive and departtogether when delivering that shespent the mosttime (104and 127min) within food to the nest;group feeding visits were followed the cavity. Foodstransferred included fruits (9 of 10 by long periodsof inactivityat the nest.The feeders visible items) and arthropods(1 item). visited the nest singly, in rapid succession,perching On 4 and 8 August I observedcopulation. The first on the bottomlip of the cavity entranceto regurgitate copulationtook placein the nesttree and the second food to the nestling.Individual feedersusually stayed in an adjacenttree. With the pair perchedon the same at the nest cavity for lessthan one minute. The mean branch facing the same direction, the male sidled up number of hornbills tending the nest during a group next to the female and lowered his head forward low visit was 1.8 ñ SD of 0.76 (n = 35). Ratesof visitation under the female's chin, gently shaking his head. to the nestby thesethree birds were notablyconstant. After 1 to 2 s, he hopped laterally over her to land Intervals between feedingsby the group (defined as on the samebranch, where he repeated the behavior a visit by any or all of the feeders)averaged 1.35 ñ from the oppositeside. With eachsuccessive hop the 0.75 h and ranged from 0.25 to 3.00 h (n = 32). Known male appearedcloser to mounting the female. After visit intervals were not significantly different be- five or six of thesehops, the male mountedthe female tween individual birds (ANOVA, Fz35 = 0.04, P = for 1 to 2 s. In the first observation the male then 0.96). Of 83 individual items that I observed being performedthree or four more of the precopulatory transferredto the nestling, 78 were fruits and 5 were hopping displaysand mountedthe femaleagain (with animal prey (2 cicadasidentified). his tail to the right side of hers) for about 5 s. Just Repeatedsightings of territorial RufousHornbill prior to the 8 August copulation,the female had al- groupsat the two nest sitesand at the PICOP site lopreenedwith anotherof the adult hornbills. enabled me to determine the number of individuals On 4 August three adults moved about the tree in in thesegroups. Family group sizes ranged from three a closegroup, while the immature bird lagged be- to seven birds (œ= 4.3, n = 7).
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