Condor, 80:382-390 0 The Cooper Ornithological Society 1978 ALLOPARENTAL CARE IN THE PURPLE GALLINULE CHARLES ONEIL’ KREKORIAN Purple Gallinules ( Gallinula mwtinica) oc- The objectives for this study of Purple Gal- cupy lowland fresh-water marshy habitats linules in Costa Rica were: (1) to establish throughout the Americas (Wetmore 1965). whether they have alloparental care, and to Their breeding range extends from central learn the status or condition of the helpers; Ohio in the United States to northern Argen- (2) to describe the types of assistance pro- tina and Uruguay in South America (A.O.U. vided, and quantify the relative contributions 1957). With the exception of Gross and van made by various members of the social unit, Tynes’ ( 1929) excellent description of the nest, including the chicks; and (3) to gain infor- eggs, and nesting behavior of the Purple Gal- mation about Purple Gallinule social struc- linule on Barro Colorado Island, Panama, and ture and data that might serve for evaluating a description of its nesting behavior in Ohio the effect of helpers on the reproductive suc- (Trautman and Glines 1964)) little is known cess of the breeders. about the ethology of this species. Available information characterizes the Purple Gallinule as monogamous, with the parents setting up STUDY AREA feeding territories and sharing care of the pre- This study was conducted on the shallow 7-acre pond cocial chicks (Gross and van Tyne 1929, Mean- (2.83 ha) located on the grounds of the Centro ley 1963, Trautman and Glines 1964). Allopa- Agronomic0 Tropical de Investigation y Ensenaiiza rental care (“helpers”) in this species is not (CATIE ) near Turrialba, Costa Rica. The pond, its flora and fauna, and the climate of the area were de- mentioned in the literature although some scribed in detail by Jenni and Collier ( 1972). zoologists (Gerald Collier and Paul Fromer, The distribution of vegetation on the pond varied pers. comm.) have suspected it. “Alloparental” from year to year due to CATIEs’ management prac- is a neutral term used to describe the behavior tices. During my study, three distinct zones of vege- tation existed, each characterized by one or more of members in a social group other than the conspicuous plant species (Fig. 1). The first con- parents which assist in the care of offspring sisted of papyrus (Cyperus papyrus) which attained (Wilson 1975). a height of 3.5 m and grew on the north and east Approximately 80 species of birds in 32 dif- sides of the pond. The second zone was made up of ferent families are known to exhibit coopera- semiaquatic grasses and sedges, 0.3-2 m high, which covered the west and the east ends of the pond. The tive breeding in which helpers feed nestlings grasses (Leersia hexandru) and sedges (Eleochuris geni- and occasionally fledglings (Skutch 1961, Har- culutu) also covered mud islands ( 5-20 m diam. ) at rison 1969, Fry 1972, Grimes 1976, Rowley the west and north parts of the pond and a large 1976, Woolfenden 1976, Zahavi 1976). In ad- island (ca. 35 m x 40 m) on the southeastern half of the pond (Fig. 1). The grasses occupied about dition, some helpers aid in nest and/or terri- 35% of the pond area. The third zone, dominated by tory defense (Ligon 1970, Parry 1970, Brown broad-leaved water lilies (Nymphueu sp. ) though 1972, Zahavi 1974, Woolfenden 1975), and often mixed with other low lying aquatic plants, nest sanitation (Skutch 1961, Ligon 1970, Fry covered most of the central and remaining portions 1972). Nest building, incubation, and brood- of the pond where grasses were not found. The pro- file of the third zone was low and open, with grassless ing by helpers have been recorded less fre- mud islands scattered throughout. A small area near quently but do occur in “mutual helpers” such the center of the pond was open water. as the anis (Crotophagu; Vehrencamp 1977) and in some other groups such as bee-eaters MATERIALS AND METHODS (Meropidae; Fry 1972) and swallows (Hir- undinidae; Skutch 1961). Observations were conducted for 117 days usually between 06:30-lo:30 from 1 October 1976 to 24 Alloparental care is currently of sociobiolog- February 1977 using a 15-60~ zoom telescope and ical interest because it is used as evidence of 10 x 50 mm binoculars. The six families studied kin selection and altruism (Wilson 1975). The were observed a total of 238 h. Five hours of observa- controversy over kin selection versus individ- tions were made between 13:00-17:OO when weather conditions were cool, and the birds active. ual selection (Hamilton 1964) can be resolved Twenty-nine Purple Gallinules were trapped with only when additional studies of alloparental self-tripping drop cages (50 x 50 x 24 cm) made of care document the filial relationship of helpers coarse mesh hardware cloth on a wood platform. Platforms were placed in the pond near the shore and to breeders and the effect of helpers on re- next to the grass islands frequented by each family. production. Captured gallinules were weighed, measured (shield, [3821 ALLOPARENTAL CARE IN GALLINULES 383 ~rasr island mud island I I I I bamboo island 0 10 20 30 m FIGURE 1. The pond at Turrialba, Costa Rica, showing the distribution of vegetation during the study and the approximate location of Purple Gallinule family territories. beak, and leg) and banded with a USFWS metal rather than plumage condition. Since my use of the band and a unique combination of at least two colored term “chick” is based on behavior rather than plum- plastic bands using red ( R ), blue ( B ), yellow ( Y) , age, it does not always denote young in natal down. and orange (0) which were placed on the lower I use the term “chick” here to denote any individual tibiotarsi. With the bird facing the observer the receiving food from an adult or older juvenile. Thus, bands were recorded in sequence from top to bottom, this category included individuals up to 60 days from the observers’ right to his left. In recording the old because they received help from adults even color code a dash always follows any bands on the though they wore the juvenile plumage described right leg and precedes any bands on the left leg. In by Wetmore ( 1965). all cases, observations of families with chicks began before all members, including chicks, were banded. JUVENILE PLUMAGE Three stages in the life history of the Purple Gal- linule (chick, juvenile, and adult) generally are Head, neck, sides, and tibia buffy brown; back, recognized. Each stage is associated with a charac- rump, and tail dull brown; wings greenish-blue washed teristic appearance, but the history of molts and plum- with brown; throat, breast, abdomen, and under ages is not known in detail. Nevertheless, the follow- tail coverts white. I considered individuals to be ing descriptions, modified from Wetmore ( 1965), juveniles (9-24 weeks of age) until they had full allowed stage identification of the socially interac- adult plumage and markings. tive birds. ADULT PLUMAGE DOWNY CHICKS (Sexes alike.) Head, breast, sides, and wings deep Above black, underneath brownish black; crown, blue; back, rump, tertials, and tail dull green; abdo- sides of head from bill to back of eyes, and throat men and tibia black; under tail coverts white. with filaments of very pale bluish white. Fully mature adult Purple Gallinules have a con- Traces of brown began to appear on the black spicuous light blue frontal plate, and the bill is plumage of some chicks when they were approxi- bright red at the base and yellow over the distal mately 2 weeks old. When they were 34 weeks old, 12-20 mm. the ventral half of the body was light beige-brown The time and duration of feeding and other and the dorsum still black with a greenish-olive pertinent behaviors were recorded with a stopwatch. tint. Juvenile plumage was attained sometime be- Each time a chick fed or was being fed by a juvenile tween 5 and 7 weeks of age by some chicks. In this or adult, the identity of individuals involved (based paper, chick status is based on a behavioral criterion on the band combinations) was recorded. When the 384 CHARLES ONEIL’ KREKORIAN TABLE 1. Purple Gallinule families studied and the number of times (numbers in parentheses) banded family members were observed to feed chicks. Feedings of chicks by unbanded adults [UA] are also included. Additional family growth after March 1977 is included in the text. Family 1 Adult O-O (46), B-B (23), Y-Y (39) Juvenile R-R (37), B-O (7) Chick B-R, B-Y, R-O Family 2A Adult Y-R (8), BB-0 (13), BB-Y (4), UA(s) (2) Juvenile R-Y (7), Y-O (26) Chick Y-B, BB-R Family 2B Adult Y-R (9), BB-0 (13), BB-Y (18), R-Y (27), 00-Y (O), Y-O (0), UA(s) (14) Juvenile Y-B (4), BB-R (6) Chick RR-O, 00-R Family 3 Adult O-Y (7), RR-Y (5) Juvenile None Chick RR-B, YY-B, YY-R Family 4 Adult R-B, O-B, YY-0, 0-BB Juvenile None Chick 3 unbanded Family 5 Adult 4 unbanded Juvenile 2 unbanded Chick 3 unbanded Family 6 Adult 2 unbanded Juvenile 2 unbanded Chick 3 eggs bands could not be seen or the individuals involved Family 1 captured on 14 and 15 October had no bands, the birds were designated as chick, were: BB (306 g), O-O (263 g), Y-Y (228 juvenile, or adult and the performed behavior de- scribed.