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ORNITOWGIA NEOTROPICAL 7: 165-169, 19% @ The Neotropical Ornithological Society

NOTES ON THE DISTRIBUTION, BODY MASS, FOODS AND VOCAL MIMICRY OF THE GRAY ( INTERMEDIA)

Betsy Trent Thomas

Waterfield, 125 Harry Settle Road, Castleton, VA 22716, U.S.A.

Key words: Gray Seedeater, Sporophila intermedia, distribution, body mass, vocal mimicry, foods and feeding behavior.

INTRODUCTION SITE AND METHODS There are over thirty members of the My observations of Gray Seedeatersare all from Sporophila in South America (Ridgely & Tudor , primarily at two study sites. One 1989). Some of these are also found in Central (Fig. 1, location 6) in the state of Guárico in the America, and at least one Sporophila torqueola central llanos at the cattle ranch Masaguaral ranges as far north assouthern Texas (American (Thomas 1979), and the other (Fig. 1, location 7) Ornithologists' Union 1983). Many sporophi- in the state of Miranda at Los Anaucos 30 km lines in South America are widespread in tropical south of Caracas in the coastal mountains (Tho- and subtropical areasof from Colom- mas 1993). I observed the at other places bia south to northern , while others briefly, mostly north of the Orinoco River. have quite restricted ranges (Ridgely & Tudor 1989). Often they are found in medium to large RESULTS mixed species f1ocks wherever local grassesare Range and Body Mass. Gray Seedeatersare usually seeding. The sexes are dimorphic, the males listed as occurring from west through usually have distinctive plumage, while generally Venezuela and , including the island of females are drab brown and difficult to identify Trinidad (Meyer de Schauensee 1966). More t:o speciesin the field. Except for a few members recently it has been reported also from the of the genus that have particularly interested northern Brazilian state of Roraima (Silva & ornithologists such as the problematic Lined Willis 1986, Silva & Oren 1990, Sick 1993). The Seedeater Sporophila lineola and the Lesson.s Gray Seedeater is often found in flocks with . bouvronides (Schwartz 1975, Ridgely other sporophiline in Venezuela and & Tudor 1989, da Silva 1995) most have received Trinidad (Table 1), where it ranges up to 1200 m only cursory attention in the literature. Two au- in the tropical and subtropical zones (ffrench thors have described some sporophiline relation- 1973, Meyer de Schauensee & Phelps 1978). ships (Meyer de Schauensee 1952, Short 1969) Most books define the size of the bird by length, based mainly on museum specimens. Almost no because museum specimens are abundant and attention has been focused on behaviors of indi- easy to measure (Meyer de Schauensee& Phelps vidual speciesin South America in what may be 1978, Hilty & Brown 1986, Ridgely & Tudor at times a tightly packed, highly competitive ge- 1989). However, body mass is generally conside- nus at ephemeral food sources. Here I give data red to be one of the best measurements of body on range, body mass, breeding season, vocal and size (Amadon 1943). Mass of some sympatric feeding behavior of the Gray SeedeaterSporophila sporophiline finches as given in Junge & Mees intermedia. I hope that field workers in the range (1961), Snow & Snow (1963), Haverschmidt of this specieswill take an interest in it and make (1968) and Thomas (1982, 1990) are summarized a thorough study to elucidate the causes and in Table 1. Data from ffrench (1973) cannot be results of its interesting behaviors. used becausehis data are combined with those of GENERAL BIOWGY

TABLE 1. Mass of sympatric sporophilines in Trinidad and Venezuela.

schistacea intermedia plumbea lineola /x;uvronides nigricollis minuta

Range of body 0"10-12.5 0"10.9-13.5 0"9-9.7 0'7.5-12 0'8-10.9 0'8.8-10 0'6.5-9.1 mass (g) 911 910.5-13 9- 98.5-10 97.5-8.5 98.5-11.2 96.5-8.4

Site of Sporophila sympatric data 1. Trinidad, w. I. x x x x x x 2. Monagas/ Anzoategui x x x x x ,3. Rio Orinoco x x x x x x 4. Amazonas x x x x x x 5'~ Cinaruco/Capanaparo x x x x x 6. Guárico x x x x 7. Los Anaucos x x x x 8. Henri Pittier x x x x x 9. Morrocoy x x x x x

Body mass from Junge & Mees 1961; Snow & Snow 1963; Haverschmidt 1968; Thomas 1982, 1990. Note: see Fig. 1 for site locations and references.

Snow & Snow (1963). Here I omit the Dull Vocal Mimicry. O!l 12 December 1982, at Los Seedeater Sporophila obscura now placed in the Anaucos in a secondary-scrub habitat, where I genus Tiaris (Collins & Kemp 1976, Ridgely & was very familiar with the typical gee-gee-gee-gee Tudor 1989), although Peters (1970) still inclu- vocalization of the ,I was stunned ded it under Sporophila, but with a note that Paul to find a male who gave a masterful c. 15 min Schwartz believed it was a Tiaris. The Gray series of mimicked calls while perched in a small Seedeateris larger by about 10% than its sympa- tree at 6-7 m. It imitated five other species of tric congeners, but seeDunning (1992) for other , all of which were common or abundant in sporophiline mass data outside of the Gray the area (Thomas 1993). Each series, or bouts of Seedeaterrange. However, the sparse mass data calls, began with a deep warbled gee-gee.gee-gee for the uncommon S. schistacea suggests an that I had previously associatedwith the species, approach to the mass of S. intermedia. then followed a rapid-fire series of imitations of the most cornrnon vocalizations of the Rufous- Breeding Months. In the llanos of Venezuela I tailed Jacamar Galbula ruficauda, Barred Ant- found an active nest in July, and in the northern shrike Thamnophilus doliatus, White-fringed mountains I observed a copulation on 2 Septem- Antwren Formicivora grisea, Bananaquit Coereba ber 1977 (Thomas 1979, 1993). Cherrie (1916) jlaveola, and Blue-gray Thraupis epis- reported two nests in June, Friedmann & Smith copus, all with great fidelity. The following year, (1955) gave September-October as breeding in the same area on 5 September 1983, I found a months, while Schafer & Phelps (1954) listed male who made two multi-mimicry songs each May-October for Henri Pittier Park in the ending with the plaintive call of the Pale-breasted Venezuelan coastal mountains. In Trinidad Spinetail Synallaxis albescens.On 6 November ffrench ( 1973) found that it bred from June-Sep- 1983 again a male mimicked the vocalizations of tember (locations in Fig. 1). Hilty & Brown the Silver-beaked Tanager Ramphocelus carbo, (1986) reported nests or breeding condition birds White-lined Tanager 1achyphonusrufus, and Ul- from Colombia in January, April, May, June, tramarine Grosbeak Cyanocompsa brissonii. The and November. A summary of these data show preceding model birds are listed in taxonomic that it appears to breed nearly throughout the order as I did not record the order in which each year except during the months of February, gave its vocalizations. None of these Gray March and December which are at the height of Seedeater males was banded so I cannot tell if the dry seasonin this northern part of the conti- they were the same or different individuals. In nent. Trinidad ffrench (1973) found that Gray Seed-

1M eater make "some imitations of other species", Rubiaceae shrub Randia venezuelensis.I watched but no species are listed. Junge & Mees (1961) a male for 4 min while it hopped from one reported that there is much imitation in its song flower cluster to another pinching the small including the familiar cry of the Yellow-bellied blossoms at the base to extract the abundant, Elaenia (Elaenia jlavogaster). Sick (1993) reported very sweet noctar. At one of the same flowering that the Plumbeous SeedeaterSporophila plumbea bushes both male and female Ruby-topaz Hum- is also a vocal mimic in . mingbirds Chrysolampis mosquitus were feeding by taking nectar through the calyx opening. Foods and Feeding Behavior. The other unusual A number of the stem-gleaning guild of observations I have are foods eaten by Gray Seed- sporophilines are migratory, following seasonal eaters. Sporophiline finch foods are usually seed ripening (Schwartz 1975, Remsen & Hunn given, l;Inqualified, as seeds (Ridgely & Tudor 1979). I found at least a few Gray Seedeatersin all 1989). With others of the genus, Gray Seedeaters months of the year at both the llanos and the often took small dry seedssuch as those of the northern Venezuelan study sites (Thomas 1979, common grass gamelote Panicum spp. and dry 1993), this suggests permanent residence and seeds of forbs, in the manner of stem-foragers. agreeswith other Venezuelan reports (Schafer & However, on 21 May 1977 at 17:45 in Los Phelps 1954, Friedmann & Smith 1955). It may Anaucos I observed three Gray Seedeaters,along be the only member of the genus that is sedenta- with a Saffron Finch Sicalis fla'1X!ola, making ry or resident in central Venezuela. Thus, it is expert 2-4 m sallies to catch insects at a fresh not surprising that inorder to survive the end of hatch of flying alates.Junge & Mees (1961) found the dry season at those sites (March-June), that Trinidadian Gray Seedeatersalso sometimes when most grasses and forbes are not in seed, foraged in the air like flycatchers. that the Gray Seedeaterwould need to exploit In the Venezuelan llanos I have records of alternate sources of food by taking both insects both male and female Gray Seedeatersnectar and nectar. foraging. On 17 and 18 Apri11982, and again on 19 May 1984 they took nectar from the common DISCUSSION sweet smelling, small white flowers of the Mayr (1964) says that an increase in size has a definite selective advantage. As a stem-forager, the about 10 % heavier weight of the Gray Seed- eater would allow it to bend down the seeding grass to a hard substrate for more efficient seed gleaning, probably giving it an advantage over other seed-eatingspecies. Residency would have great advantagesfor rapid breeding in an environ- ment where rainfall, with its quick attendant grass seed production, is somewhat irregular (Thomas 1985). This could account for Gray Seedeaterbreeding reports covering nine months of the year. Furthermore, its apparently sedenta- ry habit should also be advantageous, becauseof familiarity with the habitat, food sources and predators, as compared with the other Venezu- elan sporophiline finches that are all reported to be wanderers, migrants, or seasonal vagrants (Schafer & Phelps 1954, ffrench 1973, Hilty & FIG. 1. Sites in central Venezuela and Trinidad with Brown 1986, Ridgely & Tudor 1989). lists of sympatric Sporophila species. 1. ffrench 1973,2. Friedmann & Smith 1955,3. Cherrie 1916,4. Schwartz It can be argued that residency, and the 1979,5. Goodwin & I.entino 1990,6. Thomas 1979,7. advantages that this confers, forces Gray Seed- Thomas 1993,8. Schafer & Phelps 1954,9. I.entino & eatersinto a more catholic choice of foods during Goodwin 1991. the season when grass and forb seeds are not

167 GENERAL BIOWGY

abundant. Alternatively, their wider diet may REFEREN CES allow them to be resident while other more obli- gate seedeatersmust make seasonalmovements to Amadon, D. 1943. Bird weights as an aid in . areas with sufficient food resources. Wilson Bull. 55: 164-177. American Ornithologists' Union. 1983. Check-list of The behavior of vocal mimicry is more North American birds, 6th ed. American Orni- puzzling. The nine other speciesthat male Gray thologists' Union, Washington, D. C. Seedeatersimitated at Los Anaucoo are all com- Cherrie, G. K. 1916. A contribution to the ornitholo- mon residents of that habitat. None, however, gy of th\" Orinoco region. Bull. Brooklyn Inst. responded to the mimic perhaps ba:ause Septem- Arts and Science, Vol. 2, No.6: 133-374. .ber, November and December are at end of, or Collins, C. T., & M. H Kemp. 1976. Natal pterylosis during, their non-breeding months (Thomas of Sporophila finches. Wilson Bull. 88: 154-157. 1993). My limited data on the Gray Seedeater Dunning, J. B. 1992. Avian body masses.Boca Raton, breeding season in Los Anaucos (Thomas 1993) Florida. suggests the mimicked vocalizations may have Faaborg, J. 1985. Ecological constraints on West Indian been given as territorial advertisement, or indi- bird distributions. Pp. 621~53 in Buckley, P. A., vidual identification. Other reports of vocal Foster, M. s., Morton, E. S., Ridgely, R. S., & F. S. mimicry have also noted that the mimic made Buckley (eds.). Ornithological Monographs No. the vocalizations during its own breeding season 36. Washington, D. C. (MarshalI1950), and at a time when the models ffrench, R. 1973. A guide to the birds of Trinidad and were not breeding. However, it is not clear what Tobago. Wynnewood, Pa. advantage this might give the mimic. ffrench, R. 1985. Changes in the avifauna of Trinidad. There is also a possible conservation problem Pp. 986-991 in Buckley, P. A., Foster, M. S., Morton, E. S., Ridgely, R. S., & F. s. Buckley (eds.). with the Gray Seedeaterin Trinidad. In his 1985 Ornithological Monographs No.36. Washington, paper ffrench reported that there is such heavy, D.C. unregulated trapping of Gray Seedeatersin Trini- Friedmann, H., & F. D. Smith,Jr. 1955. A further con- dad for the cage-bird trade, that it has gone from tribution to the ornithology of northeastern Vene- abundant to rare in recent years. Other sporo- zuela. Proc. U. S. Nat. Mus. 104 (3345): 463-524. philine finches that are trapped in Trinidad, the Goodwin, M. L, & M. Lentino. 1990. Bird list of Ci- Slate-colored Seedeater Sporophila schistacea, naruco-Ca¡unaparo National Park, Apure State, , and Yellow-bellied Seed- Venezuela. Soc. Conserv. Audubon de Venezuela. eater Sporophila nigricollis are migratory. Thus, Caracas. continental South American populations are Haverschmidt, F. 1968. Birds of Surinam. Edinburgh. potential sources of immigration. The origin of Hilty, S. L., & w: L. Brown. 1986. A guide to the birds Gray Seedeaters on Trinidad may have been of Colombia. Princeton. during the Pleistocene when Trinidad was joined Junge, G. C. A., & G. F. Mees. 1961. The avifauna of to mainland South America (Faaborg 1985, Trinidad and Tobago. Zoologische Verhandelingen. Snow 1985). Since the Gray Seedeaterappears to No.37. Leiden. be not migratory, recolonization of Trinidad Lentino, M., & M. L. Goodwin. 1991. Bird list of Morrocoy National Park, Cuare Wild1ife Refuge from nearby Venezuela where it is still abundant, and adjacent areas, Falcon State, Venezuela. Soc. may not readily occur. Much interesting work Conserv. Audubon de Venezuela. Caracas. remains to be done in studying the life history of Marshall, A. J. 1950. The function of vocal mimicry in the Gray Seedeaterand its relationship with its birds. Emu 50: 5-16. congeners. Mayr, E. 1965. Systernatics and the origin of species. New York. ACKNOWLEDGEMENTS Meyer de Schauensee,R 1952. A review of the genus Sporophila. Proc. Acad. Nat. Sci. Phila. 104. I thankJ. B. Trent for some financial support; T. 153-196. Blohm for long-time accessto Masaguaral; C. T. Meyer de Schauensee,R. 1966. The species of birds of Collins for suggestions on an early draft, and South America with their distribution. Narbeth, some references;J. Ingels for discussion of these Pa. problems; as well as additional references and Meyer de Schauensee,R., & w: H. Phelps, Jr. 1978. useful comments from an anonymous reviewer. A guide to the birds of Venezuela. Princeton.

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Peters, J. L. 1970. Check-list of the birds of the world, Silva, da, J. M. C. 1995. Seasonal distribution of the Vol. 13. Cambridge, Ma. Lined SeedeaterSporophila lineola. Bull. Brit. Orn. Ridgely, R. S., & G. Tudor. 1989. The birds of South Cl. 115: 14-21. America Vol. 1. The oscine . Oxford. Snow, D. W, & B. K. Snow; 1963. Weights and wing- Remsen, J. V., & E. S. Hunn. 1979. First records of lengths i of some Trinidad birds. Zoologica 48: Sporophila caerulescensfrorn Colombia; a probable 1-12. long distance migrant frorn southem South Ameri- Snow, D. 1985. Affinities and recent history of the avi- ca. Bu]]. Brit. Orn. Cl. 99: 24-26. fauna of Trinidad and Tobago. Pp. 238-246 in Schafei, E., & ~ H. Phelps. 1954. Las aves del Parque Buckley, P. A., Foster, M. S., Morton, E. S., Ridge- Nacional "Henri Pittier" (Rancho Grande) y sus ly, R. S., & F. S. Buckley (eds.). Ornithological funciones ecologicas. Bol. Soc. Venezolana de Cien- Monographs No.36. Washington, D. C. cias Naturales Tomo XVI No.83. Caracas. Thomas, B. T. 1979. The birds of a ranch in the Vene- Schwartz, P. 1975. Solved and unsolved problems in the zuelan llanos. Pp. 213-232 in Eisenberg, J. F. (ed.). Sporophila lineola/bouvronides complex (Aves: Vertebrate ecology of the northern Neotropics. Emberizidae). Annals of Carnegie Mus. Vol. 45. Washington, D. C. Article 14. Pittsburgh, Pa. Thomas, B. T. 1982. Weights of some Venezuelan birds. Schwartz, P. 1979. Atlas de la region sur. Codesur. Mi- Bull. Brit. Orn. Cl. 102: 48-52. nisterio del Ambiente, Caracas, Venezuela. Thomas, B. T. 1985. Coexistence and behavior diffe- Short, L. L. 1969. Relationships among some South rences among the three western hemisphere storks. American seedeaters(Sporophila), with a record of Pp. 921-931 in Buckley, P. A., Foster, M. S., S. hypochroma for Argentina. Wilson Bu]]. 81: Morton, E. S., Ridgely, R. S., & F. S. Buckley (eds.). 216-219. Ornithological Monographs No.36. Washington, Sick, H. 1993. Birds in Brazil. Princeton. D.C. Silva, da, J. M. C., & E. o. Willis. 1986. Notas sobre Thomas, B. T. 1990. Additional weights of Venezuelan distribui~ao de quatro espéciesde aves AmazOnica birds. Bull. Brit. Orn. Cl. 110: 48-51. Brasileira. Bol. Mus. Para. Emilio Goeldi. Ser. 2oo- Thomas, B. T. 1993. Birds of a northern Venezuelan se- logia 2: 151-158. condary-scrub habitat. Bull. Brit. Orn. Cl. 113: Silva, da, J. M. C., & D. Oren. 1990. Resultos de urna 9-17. excur~ao omiol6gica a Ilha de Maracá, Roraima, Brasil. Goeldiana Zoologia 5: 1-8. Accepted 15 December 1996.

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