ORNITOLOGIA NEOTROPICAL 8: 121-132, 1997@ The Neotropical Ornithological Society

DISTRIBUTION AND DRY-SEASON ECOLOGY OF PFRIMER'SCONURE PYRRHURA PFRIMERI, WITH A REAPPRAISAL

OF BRAZILIAN PYRRHURA "LEUCOTIS"

Fábio Olmos1,2, Paulo Martusce"i2 & Robson Silva e Silva2,3

1 Rua Antonio F. Gandza 182, sao Vicente, SP, 11390-250, Brazil.

2The Coragyps Bros, C. P. 194, Peruíbe, SP, 11750-970, Brazil.2,3Rua sao José 48/31, Santos, SP. 11040-200, Brazil.

Resumo. pyrrhura pfrimeri é urna espécie pouco conhecida pertencente ao grupo leucotis, end&mica das florestassecas decíduas que crescem em solos derivados de calcário e maci~os rochosos em urna área limitada a oeste daSerra Geral, Brasil Central. A espécie tem urna das menores distribui~oes geográficas entre os psitacídeos neotropi-cais. Durante o início da esta~ao seca os periquitos alimentam-se principalmente de flores e sementes, utilizandofigos no final da esta~ao. A reprodu~ao parece ocorrer no final da esta~ao seca. Devido a sua distribuis:ao restritae a destrui~ao rápida de seu habitat a espécie deve ser considerada amea~ada de extin~ao. As diferens:as morfol6gicase ecol6gicas, bem como as distribui~oes alopátricas dos tr&s taxa brasileiros classificados como pyrrhura leucotissao marcantes o suficiente para lhes dar status específico pleno.

Abstract. Pfrimer's Conure pyrrhura pfrimeri is a poorly known taxon belonging to the leucotis species group. Thespecies is endemic to central Brazil, where it is locally common, but restricted to dry, deciduous forests growingon limestone-derived soil patches and rocky massifs west of the Serra Geral, giving it one of the smallest distribu-tions among Neotropical parrots. During the early dry season, the conures feed heavily on flowers and seeds,switching to figs later in the year. Breeding seems to occur in the early rainy season. Owing to its limited distribu-tion and the rate of habitat destruction, P; pfrimeri should be considered threatened with extinction. The morpho-logical and ecological differences and allopatric distributions of three Brazilian taxa grouped under pyrrhura "leuco-tis" are marked enough to give them full specific status. Accepted 20 MaTCh 1997.

distribution and habitat of Pfrimer's Conure,provide the first observations on its ecology andcomment on its taxonomical status and rela-tionship with other endemic Brazilian taxagrouped with Pyrrhura leucotis.

INTRODUCTION

Pfrimer's Conure Pyrrhura pfrimeri was de-scribed by Miranda-Ribeiro (1920), based on anundesignated type from Santa Maria de Tagua-tinga, present-day Taguatinga do Tocantins(12°26'S,45°40'W), central Brazil. Later, Berla(1946) designated the lectotype and paralecto-types of the taxon and treated it as a subspeciesof pyrrhura leucotis. This treatment was chal-lenged by da Silva (1989, 1995), who consideredthe form sufficiently distinct and geographicallyisolated to warrant full specific status; he consid-ered it one of the endemic species of the drydeciduous forests growing between the sao Fran-cisco and Araguaia rivers in Brazil. Nothing hasbeen reported on the ecology of P. pfrimeri andeven its distribution is poorly understood, asmost authors (Vriends 1979, Forshaw & Cooper1989, Arndt 1993) consider it known only fromthe type locality, evidently ignoring Pinto(1938), who listed specimens from localitiesother than the typical one. Here we describe the

METHODS

Localities where R pfrimeri was known to occuror to have occurred, as documented by collectedspecimens housed in the Museu de Zoologia daU niversidade de sao Paulo (MZUSP) and by daSilva (1989), plus intervening areas of suitablehabitat, were visited in 14-23 June and 27 Octo-ber- 03 November 1995. More time was spentat two localities, Nova Roma and sao Domingos(Fig. 1) to obtain ecological data. In each locality,searches for conures were made by walking alongtransects through natural vegetation patches andadjacent anthropogenic habitats. Every conureobserved or heard was recorded, as was additionaldata including group size and habitat type (dryforest, riverine forest, cerrado, pasture, disturbed

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ments from the analyses because of the largenumber of tail-damaged specimens in the sample.

RESULTS AND DISCUSSION

Habitat and Distribution. pyrrhura pfrimeri isrestricted to semi-deciduous (riverine) or decid-uous (dry) forests growing on limestone outcropsor limestone-derived soil patches from Taguatin-ga de Goias south to Iaciara, a stretch about 300km long and, at some points, only 30 km widealong the foothills west of the Serra Geral massif(Fig. 1, 2). This limestone patch, pre-Cambrianin age, was exposed by the erosion of the morerecent sandstone and quartzite deposits thatcovered it and of which the Serra Geral and theChapada dos Veadeiros are examples (RADAM-BRASIL 1989). The climate in the region isseasonal, with a dry season extending from Mayto September and an average annual precipitationaround 1,300 mm (RADAMBRASIL 1989).During 1994 a pronounced drought affected allof Brazil; the rains began 3-4 months later thanusual, while in 1995 the rains were also late, withno substantial precipitation being recorded up toearly November. Whereas most of central Brazilis covered by cerrado (savanna) vegetation, themore fertile limestone-derived soils supportforests presenting an almost continuous canopy15-20 m high. That canopy is dominated by1abebuia impetiginosa (Bignoniaceae), Astroniumurundeuva (Anacardiacea), Pseudobombax grandi-florum., Chorisia venticosa, Cavallinesia arborea(Bombacaceae), Caesalpin~a ferrea, Enterolobium

contortisiliquum, Copaifera langsdorfii, Piptade-nia sp. (Leguminosae), with occasional emer-gents 25 m or more high (mostly Cavallinesiaarborea.). A dense undergrowth of lianas and, insome areas, bamboo (Actinocladum or Guaduaspp.) occurs in the more disturbed patches. Onrocky outcrops several species of columnar cactiand terrestrial bromeliads occur, in an associa-tion much like that found in semi-arid north-eastern Brazil (see Ratter et al. 1978, RADAM-BRASIL 1989). In fact, Ratter et al. (1978) con-sidered similar forests on the eastern side of theSerra Geral as representing an arboreal type ofcaatinga, the xeric deciduous formation en-demic to northeastern Brazil (Rizzini 1979). Thehabitat of pyrrhura pfrimeri (Fig. 3) represents acaatinga forest island amid the cerrado, a con-

areas). Habitat categories followed RADAM-BRASIL (1989). Whenever a group of parrotswas observed perched or feeding, plant foodspecies, part eaten (flower, seed, fruit pulp orwhole fruit), presence and number of juvenilesand general behavior were recorded.

Each time a group of conures was foundeating we recorded a feeding bout. Differentgroups congregating at a food source could.generally be told apart as they arrived or depart-ed, because they did not mingle while flying(despite doing so while feeding); in this case werecorded one feeding bout for each group. Weattempted to infere habitat preferences and rel-ative abundances of the conures by censusingthem along line transects. Transects in eachhabitat were walked at least twice; total distancescovered, as recorded with the aid of a pedometer,were 45.7 km in June and 57.5 km in October-November. Of thest; 29.8 km were in areaswhere R pfrimeri did not occur, and they areexcluded from analyses. In all, 50.8 km werewalked through dry forest, 19.3 km along a mo-saic of pasture and small forest fragments, and3.5 km along selectively logged riverine forest(the only area of this habitat we could find). Tocalculate relative population densities, we con-sidered transect width to be 60 m in dry forestand more open areas and 20 m at the vine-tangledriverine forest.

We assessed the species' geographical distri-bution by visiting localities where it had notbeen previously recorded but with the seeminglysuitable habitat typt; based on the localitieswhere we observed the conures. We also visitedadjacent areas harboring different plant com-munities. We examined the area from Iaciaranorth to Campos Belos along the western foot-hills of the Serra Geral (Fig. 1), plus a explorato-ry visit to dry forests between Barreiras and For-mosa do Rio Preto on the eastern side.

Specimens of pyrrhura leucotis ssp. (7 pfrime.ri, 15 leucotis, and 12 griseipectus, Appendix 1)housed in the Museu de Zoologia da Universi-dade de sao Paulo (MZUSP) were examinedand measured with calipers to the nearest mm.Because of the small sample size sexes were pool-ed. We measured wing and taillength, billlengthfrom tip to the joint of the upper maxilla, billwidth at its bast; and mandibular height at thebase. During analyses we discarded tail measure-

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ECOWGY OF PYRRHURA PFRIMERI

rax cyanopogon) (pers. obs.). These dry forests arequite distinct from the surrounding savanna (fora description of cerrado see Eiten 1972, 1982),their presence on the limestone patches probablyarising from a combination of more fertile soils

clusion supported by the presence there, but notin surrounding cerrado, of bird species endemicto the caatinga, e. g., Great Xenops (Megaxenopspamaguae), Scarlet-throated Tanager ( Composo-thraupis loricata), and White-naped Jay ( Cyanoco-

15D

FIG. 1. The dry forest region of centrnl Brnzil. The square in the inserted map shows its generallocation. 5pottedareas represent the original extent of forest, now greatly reduced, at the foothills of the 5errn Gernl. Dots representlocalities where pyrrhura pfrimeri has been observed: A) Nova Roma (13°50'5, 46°52'W); B) Riverine forests ofthe rio Parana (13°55'5, 47°OO'W); C) Fazenda 5alobro (14°055, 46°47W); D) Guarnni de Goiás (14°01'5,

46°31'W); E) Fazenda Floresta (13°27'5, 46°29'W).

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OLMOSETAL

We did not observed R pfrimeri in the cerrado,despite that representing a flying distance of lessthan one kilometer. In addition, many cultivatedfruiting mango trees atop the massif attractedhundreds of Canary-winged Parakeets (Brotogerisversicolorus) that carne from the forest during themorning and spent the whole day feeding there.pyrrhura pfrimeri 's reluctance to come to thesetrees was all the more surprising because duringthe period (November) the forest was almost

completely dry and there were few fruiting trees,and the species does feed on cultivated crops (see

below).

Group Size and Population Densities. Group sizein June ranged from 2 to 32, with a mean of 12.6(SD = 6.93, n=36 sightings). The largest groups

may have resulted from the fusion of smallergroups, as observed in a large agregation of over50 conures, not included in our analyses, thatwere feeding on cultivated rice (see below). InOctober-November group size ranged from 2 to33 (mean=12, SD = 8.09, n=26), not being sta-

tistically different from the previous period

(t=0.759, P>0.05). Fledglings (begging young-sters) were observed only in June.

In dry forest areas conure relative densities(pooled data from June and October-Novembertotaling 602 individual sightings) averaged 197.66individuals/km2 (11.9 individuals/walked km),wheres in nearby pasture-isolated forest it wasonly 23.4 individuals/km2 (1.4 individuals/walked km, or only 27 recorded individuals), andall conures were observed on the few remainingforest blocks left on the pasture. We think resultsfor the dry forest censuses are somewhat of anoverestimate, because in June one of the mainfood plants of the conures was a ruderal thatgrew mostly along the tracks used as transectline. In October-November most, if not all, ofthe conures at Nova Roma (where all the cen-suses were conducted in the period) were gather-ing to feed in a sole fig tree (Ficus gomeleira) nearthe track. Although those facts probably in-duced errors in absolute population estimatesbecause the conures congregated along the trans-ect tracks, the data is useful to compare therelative densities among habitats and show thatconures are far less common in fragmentedhabitat. In the riverine forest we found only one

and, perhaps, more efficient drainage patterns(Ratter et al. 1978).

During the two periods we visited the area,both in the dry season, most trees had shed theirleaves, except for the uncommon Ficus and somespecies of legume. A few tree species were inflower. This was especially evident in June, whenthe purple flowers of the abundant 1abebuiaimpetiginosa tinged whole stretches of forest and.could be used as a visual aid to locate dry forestp.atches. This contrast is striking in the saoDomingos region, where the forested limestonehills raise quite abruptly from the surroundingcerrado-covered plain.

pyrrhura pfrimeri was found in every patch ofdry forest we surveyed (Fig. 1), even in somequite degraded patches near Nova Roma and saoDomingos, and it is probable that its distri-bution is continuous wherever forest survives. Insurveys made in the surrounding cerrado andeast of the Serra Geral, between Barreiras (12 oS,45°W) and Formosa do Rio Preto (10O57'S,45 OO5'W), where a somewhat different kind ofdry forest occurs, we failed to find the species,despite walking 23.6 km of transects. The onlysimilar-sizedparrots found in these areas were thePeach-fronted Conure (Aratinga aurea), wide-spread all over the cerrado and also in forest nearFormosa do Rio Preto, and the Cactus Conure(A. cactorum) found only east of the Serra Geralin lower and drier vegetation. Apparently Ara-tinga aurea avoids areas of continuous dry forest,entering only fragmented and disturbed patches,thereby suggesting a degree of ecological segrega-tion between this species and P. pfrimeri.

The dry forests east of the Serra Geral fromMinas Gerais to Bahia have been ornithologi-cally explored, starting with Spix in the lastcentury, but no pyrrhura has been found there(da Silva 1989). That is puzzling considering thatthe taxon geographically, closest to P. pfrimeri isP.leucotis from eastern Brazil's coastal forests (seebelow) and some related taxa linking themwould be expected to occur in the interveningarea east of the Serra Geral, especially with thesimilarities between the forests there and to thewest.

At N ova Roma dry forest and dense cerradovegetation meet in a quite abrupt transition, withcerrado growing atop a quartzite massif and dryforest on the foothills and surrounding plain.

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ECOWGY OF PYRRHURA PFRJMERJ

FIG. 2. sao Domingos, Goiás. Remnant dry-forest patches, easily detected by flowering Tabebuia impetiginosa,grow on limestone-derived soils. The sar1dstone cliffs of the Serra Geral, covered by cerrado, are in the background.

Food Habits and Behavio1: A total of 46 feedingbouts were recorded in June, plus 25 in October-November. We noted striking differences in foodplants eaten in the two periods. The flowers of1abebuia impetiginosa were eaten in 43 % of allfeeding bouts observed. The conures spent hourspicking one flower at a time, chewing around itsbase, discarding it and repeating the process,usually ignoring the approach of an observerright to the base of the food tree. The secondmost important item (33 %) were the immatureseeds of Hyptis sp. (Labiatae), an introducedruderal herb very common along the dirt roadsand tracks. The conures carne to the ground tofeed on Hyptis. Other food items during theperiod were the catkins of Cecropia sp. (Cecro-piaceae), the fruits of Ficus gomeleira (Moraceae)and Psidium guajava (Myrtaceae), and theflowers of a Bauhinia sp. (Leguminosae), eachaccounting for 6 % of the feeding bouts. All figswe opened were heavily infested by Agaonidaewasps, and these probably contributed proteinto the diet in a season when the birds had

group with 11 conures, sampling being too smallfor comparisons.

Censusing conure groups at Nova Roma asthey flew from their feeding trees to roost on thelimestone hills during sunset produced a pro-bable total population of 120-150 individualsfor the whole area (around 2 km2 of forest)or 60-75 individuals/km2, probably a moreaccurate figure than the estimate from the trans-ects.

Overall, pyrrhura pfrimeri is fairly easilyfound wherever dry forest persists, and, in fact,it is the commonest parrot in this habitat, beingobserved even in disturbed and fragmented forestpatches. Our observations suggest that Pfrimer'sConure is able to persist and even breed in smallforest fragments, although in very reduced num-bers. But considering that widespread deforesta-tion is a recent phenomenon in its area ofoccurence it is possible such isolated groups willbe unable to persist in the long term, especiallyas such remnants are vulnerable to fire and con-tinue to be selectively logged (see Conservation).

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case of Pyrrhura pfrimeri it is clear that flowersare important in a season when there are fewalternative food sources, accounting for53 % offeeding bouts in June, much as has been shownfor Pionu5 maximiliani living in semi-deciduousforests in southeastern Brazil (Galetti 1993).Besides Tabebuia impetigino5a, some of the mostcommon trees in the dry forest, like P5eudobom-bax grandijlorum, Chori5ia ventico5a, and Caval-line5ia arborea, flower gregariously in the dry sea-son and present large and nectar-rich flowers,providing additional food sources after theflowering of Tabebuia has ended. The abundanceof nectar-rich species that flower during the dryseason and the presence of fig trees, resourcesthat are absent from the surrounding cerradovegetation, explain the existence of a conurebelonging to a rainforest group inhabiting a sea-sonal, even xeric, habitat.

AII feeding bouts were observed in forestsgrowing over flatter ground; none was observedduring transects in the limestone outcrops. AtNova Roma, where more extensive observationswere made, the conures flew in early morningfrom the higher parts with exposed rock to thebase of the hills and surrounding plain to feed,returning at dusk. This pattern was consistentand, coupled with the observation that forest onflatter ground is less xeric (Ratter et al. 1978)suggests that the combination of flatter groundand foothill forest may be critical for the survivalof the conures.

The observations of Pyrrhura pfrimeri feedingat ground level are somewhat surprising, as mostspecies in the genus are restricted to the canopy(Vriends 1979, Roth 1984, Forshaw 1989);feeding on the ground is similar to some smallAratinga conures like A. cactorum and A. aurea(pers. observ.). Feeding on the ground hasobvious advantages in the seasonal dry forest,where the diversity of resources at any one timeis limited.

Like other Pyrrhura, P. pfrimeri is a noisybird and groups were usually first detected bytheir calls. In June, when fledglings were present,their continuous begging was the best clue tolocate groups. pyrrhura pfrimeri flies at or belowcanopy level, dashing among the trees, andperching both in the crowns and at lower levels.We observed groups perched on very exposeddry small trees amid pasture, the conures flying

dependent fledglings. This also suggests that therecord of a pyrrhura leucotis with a crop full ofinsects (Schubart et al. 1965) may represent a caseof deliberate, rather than accidental, ingestion(see also Martuscelli 1994 for another instance of

insect-eating).Pyrrhura pfrimeri carne to the vicinity of

human habitations to feed on the fruit pulp ofcultivated Guava (Psidium guajava) but the seeds.were discarded. Our calculations did not includetQe observation of c. 50 conures feeding onculti-vated rice Oryza sativa because we could not besure of how many different groups gatheredthere. As when feeding on Hyptis, the conurescarne to the ground to feed on the mature seeds,while several sentinels kept watch in the nearbytrees and shrubs.

In October-November the conures were ob-served to feed mainly on the fruits of Ficus gome.leira (72 %), but also on the fruit pulp of a Poute-ria sp. (Sapotaceae, 12 %) and the inflorescencesof Maclura tinctoria (Moraceae, 8 % ). Odd itemswere young leaves of Astronium. urundeuva andshoots of a Psittacanthus sp. (Loranthaceae, 4 and8% respectively). Over 120 conures would con-gregate in the area's sole fruiting fig tree, spend-ing most of the day in its vicinity. Figs were themost important resource during this period,with all conures in the area feeding from a singletree. Like other pyrrhura (Roth 1984, Forshaw &Cooper 1989, Pizo et al. 1996), P. pfrimeri usesabundant food resources clumped both in timeand space, with different groups congregating toshare the resource, and showing a preferencefor soft items. Flowers, mainly large ones withabundant nectar, have been recorded as a fooditem of some importance for the rainforest Pyrr-hura picta, & P. perlata (Roth 1984) and P. albipec.tus (Krabbe & Sornoza 1994), and as making25% of the diet of P. frontalis (Pizo et al. 1996),but were absent from the diet of P. leucotis in thelowland humid forests of Espirito Santo, whereit was found to feed only on soft fruit and seeds,especially Cecropia spp. (Simao et al. 1998), sug-gesting an interesting ecological difference be-tween leucotis and pfrimeri.

Although there are few quantitative data onthe importance of flowers for Neotropical par-rots, this resource may be critical to species orpopulations inhabiting seasonal forests with amarked dry season (Galetti, pers. com.). In the

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ECOWGY OF PYRRHURA PFRIMERI

FIG. 3. A pair of pyrrhura pftimeri feeding on cultivat-ed guava Psidium guajava at Fazenda Floresta. Noticethe absence of the auricular patch and the extent ofblue in the head.

comparatively low (3-6 m) over such open areas.Conures feeding on the ground or at low levelsdisplayed an evident sentinel system, one ormore conures always being positioned atop anearby tree. Sentinels gave alarm calls, causingthe group to fly, as soon any possible threat wasdetected. The conures were also observed to fleeafter a group of Jandaya Conures (Aratingajandaya) distressed by a soaring Black-and-White.Hawk-eagle (Spizastur melanoleucus) gave alarmcalls. Another conure group with young fledfrom approaching White-naped Jays (Cyanocorax

cyanopogon).Mating was observed in June. A pair was

engaged in allopreening when the male embracedthe female with his left wing and leg and half-mounted her; the female lowered its body andraised and twisted her tail out of the way of themale's vent. The male did not grab the femalewith his bill. The remainder of the group keptfeeding with no overt notice of the copulation.The presence of several individuals with worntails in October-November suggests that theycould be nesting during that period. Nest siteswere not found but because the conures alwaysflew towards exposed limestone outcrops duringsunset and one male collected in July (MZUSP15765) shows a worn tail with encrusted soil, itis possible this conure nests in rock cavities,which are very common in the area (pers.

observ.).We were unable to find the exact spots the

conures spent the night and therefore wereunable to assess if the whole group sleeps in acavity, as pyrrhura picta (Roth 1984) and R leuco.tis griseipectus (pers. observ.), but from theirbehavior at sunset we believe it is likely.

That young were observed in June suggeststhat breeding occurs in the late rainy season toearly dry season (April-May). Nevertheless,1994 and 1995 were drought years in Brazil, withthe rains being late by several months andcausing most bird species either to forgo breedingor to breed later than usual (pers. observ.). Thesame probably happened with R pfrimeri. Theobservations of birds with worn tails in Novem-ber, just when the very first rains, late by twomonths, were falling, support the view that thebirds adjust their breeding cycle to the rain pat-terns.

Most pairs that bred produced two fledglings,as observed in June, but we also observed pairswith three young. Overall, fledglings made uponly 20.6 % of the individuals of the 6 groups wecould positively identify and count them, sug-gesting most pairs had not bred or were unsuc-cessful during the 1994-1995 breeding season.

Conservation. The greatest, and most pressing,threat to the survival of pyrrhura pfrimeri ishabitat destruction, as limited trade or persecu-tion has been detected. Few Brazilian avicul-turists have the species in their collections, andfew conures are in captivity. The conures re-portedly have been bought from bird dealersbased at Brasília, the country's capital (N. Ka-wall, pers. com.). If trading continues at this lowlevel it is unlikely to be a source of concern inthe short-term. That can change, nevertheless,

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OLMOSETAL

regard to the use of forest on flatter groundbelow the limestone outcrops. As these forestsare fast disappearing, so may be a critcal resourcefor the species. Also, although forest on the lime-stone outcrops are usually spared, the proximityof a major developing center at the country'scapital at Brasília makes one wonder how long itwill take for a cement company to becomeestablished in the area.

It is difficult to produce a population esti-mate for pyrrhura pfrimeri because the actualextent of remnant habitat is unknown, and thepopulation densities obtained from transectsmay be an overestimate. Supposing that 40 % ofthe forest existing in the middle 80's persists andconure densities are around the lower estimateswe made for the Nova Roma region (60-75 indi-viduals/km2), the total conure population maybe around 162,000-202,500 individuals. On theother hand, it is important to note that whereasNova Roma still had forest on flatter groundadjacent to rocky outcrops, all of these has beencleared over most of sao Domingos north toCampos Belos, so densities are probably not thesame. Considering the limited extent of its habi-tat, that habitat's quick destruction and fragmen-tation, and probable simultaneous decline inoverall numbers of conures, we feel pyrrhura pfri-meri deserves at least vulnerable status, followingthe criteria adopted by the IUCN (1995).

A proposal to create a national park at saoDomingos, mostly to protected a cave-rich karstformation, has been under consideration by Bra-zilian federal authorities for almost a decade.That would also protect a sizeable patch of dryforest and populations of pyrrhura pfrimeri,albeit of unknown size. More recently, after lob-bying by local NGO's, the Goiás state govern-ment agreed to establish a 50,000 ha state park inthe same area as an "environmental compen-sation" for the area to be flooded by a hydroelec-tric project on the headwaters of the Tocantinsriver, west of the Chapada dos Veadeiros. TerraRonca State Park had its limits decreed in No-vember 1996, and the first 15,000 ha of the parkare already scheduled to be bought. This is thefirst action ever to be taken to protect a dry-fo-rest area. The park reaches the Serra Geral,including the many rivers that drain into thekarst area, protecting significat areas of cerradobesides some of the best remnant dry forests

if the species comes into fashion among avi-culturists, thereby increasing the demand.

The total area covered by dry forest west ofthe Serra Geral was estimated to be around 6,750km2 in the middle 1980's (RADAMBRASIL1989), that representing roughly 75 % of theoriginal forested area. The dry forest patches havebeen cleared for pasture and explored by loggers,since woods like A5tronium urundeuva and 1abe..buia impetigino5a, prized for their endurance, arein great demand for the making of fence poles.c'omparing vegetation maps in RADAMBRA-SIL (1989) with what we observed in the area, wecan attest that all forest around Alvorada doNorte is gone, as well as most of the largeforested area that once existed between Iaciaraand the Rio Parana (Fig. 1). Judging from ourexperience in the region only 40 % of the forestremain, and most ofit continues to be logged.

In October-November we were impressed bythe changes observed around Nova Roma sinceJune. Two forest fragments isolated amid pastur~where we had observed PyrrhuTa pfrimeri hadvanished and a third had been badly damaged byfire. Signs of forest burning were widespread and,upon investigating, we discovered that most fireshad originated from logging camps and fromattempts to improve the local poor pasture. Inaddition to burning the forest, the loggers wereremoving all 1abebuia trees over 50 cm dbh, ef-fectively depleting the conure's food supply. Firealso killed all saplings and damaged thin-barkedspecies like fig trees.

The largest remnants of dry forest we ob-served are around Guarani de Goiás, where anextensive tract with over 50 km2 was beingquickly logged in June. Further north, towardssao Domingos and Campos Belos, forest remainsonly on limestone outcrops too steep and rockyto be of any use, and in a few properties that havenot been completely colonized, like FazendaFloresta, some of them being fairly extensive(over 50 km2). We found most, if not all, foreston the flatter areas has been destroyed and thatthe remnants are being selectively logged. Allof the riverine forests we visited were badlydamaged, and are now dominated by bamboo,second growth and liana-covered trees. Althoughpyrrhura pfrimeri seems to withstand a fairlyhigh degree of habitat disturbance, its ecologicalneeds are incompletely known, especially with

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in northeastern Brazil: Serra do Baturité and

adjácent ranges (around 04°10'S, 38°50'W),Serra Negra (38°S, 08°40'W) and Murici(09°47'S, 36°50'W) (Yamashita & Coelho 1985,Teixeira et al. 1988). These widely dispersedlocalities represent natural forest islands isolatedin regions otherwise covered by xeric vegetation(Baturit~ and Serra Negra) and the largestremaining montane forest tract remaining in theregion (Murici). It is interesting that the lowlandrainforests of this part of Brazil had been wellexplored (before being almost completely cut inthe 1970-1980's to make room for sugarcaneplantations) without the species being recorded(Pinto 1940, 1956; Pinto & Camargo 1961;Teixeira 1986, Teixeira et al. 1988).

The three Brazilian pyrrhura «leucotis» havecompletely allopatric distributions, being effec-tively isolated from each other by thousands ofkilometers. P. pfrimeri shows both the mostextreme disjunction and distinctive habitat whencompared to all other taxa of the group. It is theonly species restricted to dry forests and tooccurs outside the Atlantic forest domain (Rizzi-ni 1979). pyrrhura pfrimeri and P. leucotis aresimilar in size, with no significant differences in

wing length (t=0.937, P>0.1, df=20), billlength(t=0.173, P>0.5, df=20) and mandibular height(t=0.936, P>0.2, df=20), but differ in colorpattern, the most striking character being theabsence of the white to light buffy auricularpatch present in leucotis and griseipectus. It alsohas a pale blue forehead, crown, occiput, andnape, with this color spreading to the sides of theneck and throat before grading to green down tothe breast. In contrast, leucotis has blue only onthe forehead (sometimes also just above the eyes),crown and nape being buff or gray. Consistent,but minor, differences in color are the blackishexternal primaries (P10) of pfrimeri, with blueonly along the raquis (mostly blue feathers inleucotis and griseipectus) and the almost completelack of the dark narrow terminal band of thebreast feathers. Nevertheless, in the living bird,the most striking characteristic of pyrrhurapfrimeri is the contrast between the dark red faceand the light blue head, and the absence of theauricular patch. These characters are probablyimportant in species recognition and communi-cation.

(K. Ramos Alves, pers. com.). The effective pro-tection of the state park at sao Domingos willalso benefit a population of the near-threatenedBrazilian Black-tyrant Knipolegus franciscanusand other dry-forest endemics (see da Silva &Oren 1992), a welcome measure that could becomplemented by a second reserve at Guarani deGoiás, where the extent and condition of the dryforests make the area worth conserving.

1axonomy. The pyrrhura leucotis group com-prlses five taxa with disjunct distributions. Twoof these (P. leucotis emma and P.I. auricularis) arerestricted to rain and cloud.forests in the coastalranges of northern Venezuela (Phelps & Phelps1958, Meyer de Schauensee & Phelps 1978,Arndt 1993), and are isolated from the Brazilianpopulations by thousands of kilometers occu-pied by the several forms of the related Pyrrhurapicta complex, a group with an Amazonian dis-tribution (Arndt 1993). Whether this is anothercase of species or sibiling species with easternBrazilian and northern Venezuelan ranges iso-lated by the bulk of the Amazonian forest (Willis1992), or if the leucotis species group as currentlyknown is poliphyletic, are contrasting hypothe-sis that deserve to be tested but are beyond thescope of this paper.

Pyrrhura leucotis is a bird of the lowland andfoothill ( under 500 m asl) forests east of theBrazilian coastal range from southern Bahia(c. 14 OS), south through Espirito Santo to Sepeti-ba, just south of the city of Rio de Janeiro(Pelzeln 1871), including the adjacent lowlandforests of eastern Minas Gerais. There is norecord or specimen of the species from furthersouth (Miranda-Ribeiro 1920, Pinto 1938, 1978;Pinto & Camargo 1961); the record of it oc-curring to sao Paulo is in error (Sick 1993). Thesouthernmost present-day locality is Serra doTinguá, near Rio de Janeiro (22 o36'S, 43 o27'W;Scott & Brooke 1985). The available recordsshow Pyrrhura leucotis to be a bird of the low-land forests of Amazonian affinities restricted tothis stretch of eastern Brazil and adjacent foot-hill forests (see Rizzini 1979), being replaced bypyrrhura frontalis at higher altitudes.

Pyrrhura griseipectus (treated as pyrrhura ana-ca by Teixeira 1991) has been recorded from onlythree montane ( above 500 m asl) evergreen todeciduous (at the lower altitudes) forest isolates

OLMOS ET AL

Although pyrrhura pfrimeri is morphologi-cally close to leucotis, the combined morphologi-cal differences, lack of intermediates or inter-vening populations that would allow gene flowbetween the two taxa, the very distinctivehabitat, implying in significant ecological dif-ferences, are significant enough to warrantreturning pyrrhura pfrimeri to full specific status,as suggested by da Silva (1989, 1996).

Although both the Atlantic forest taxa maybe thought to be closely related, based on geo-graphical proximity, as suggested by Hellmayr(1929) and Pinto & Camargo (1961), Pyrrhuragriseipectus in fact seem to be more divergentfrom R leucotis than R pfrimeri. Both are distin-guished by differences of the periophthalmicring (dark blue in leucotis and pfrimeri, white ingriseipectus), auricular patch (cream to yellowishin leucotis, often with a buff tinge; pure white orcream in griseipectus, being noticiably larger inthe latter), head color (front, nape and sides ofneck suffused with blue in leucotis, with no bluein griseipectus) and breast feathers (green with ablue suffusionmore intense near the neck andwitha broad light grey or buff subterminal bandand a narrow blackish terminal one in leucotis,dusky grey with a broad cream to light buffterminal band in griseipectus). pyrrhura griseipec-tus is the same size as leucotis, as measured fromtheir wing (t=O.183, P>O.5, df=25), but animportant difference, previously noted by Hell-mayr (1929), is the significantly longer bill ofgriseipectus (t=4.75, P<O.OO1, df=25), and itshigher mandible (t=3.49, P<O.O1, df=25), itlooking proportionally larger-headed than theother Brazilian taxa, something anatomicalstudies may prove. Interestingly, there is no diffe-rence in bill width between griseipectus and leuco-tis (t=O.OOO524, P>O.9, df=25). As in the case ofR pfrimeri, the even more marked morpho-logical differences, the geographycal isolationbetween the taxa, and different habitats may beevoked to assign full species status to pyrrhuragriseipectus, as proposed by da Silva (1989).

The avifaunas of eastern Brazilian forestsfrom Espirito Santo to Ceará have a strong Ama-zonian influence (Pinto 1956, Pinto & Camargo1961, Cracraft 1985, Sick 1993, Willis 1992, Gon-zaga et al. 1995), with several species or sub-species being allopatric representatives of Ama-zonian taxa (Haffer 1974), e. g. the endemic

Baturité Toucanet (Selenidera gouldii baturitensis)(Pinto & Camargo 1961). It is conceivable thatboth leucotis and griseipectus arose from ancestorsof Amazonian origin related to pyrrhura pic:tathat colonized the forests of eastern Brazil whenthey were continuous with the Amazon forest,perhaps during one of the last climatic opti-mums 30-23,000 years BP (Ab'Saber 1990), andlater populations became isolated as the forestshrank, especially the ones in northeastern Bra-zil.

Comparing pyrrhura griseipectus with Pyrr-hura picta, the subspecies that seems closest to itboth in size and general morphology is R p. pictafrom northern Brazil, which share most charac-teristics including the well-developed red bend ofthe wing (absent from other R picta subspecies),narrow blue half-collar, and dusky to brownishgrey breast feathers with broad grey or light buffterminal bands. Some R p. picta from Amapá(but not all) differ from typical specimens in thatthey lack blue on the head, very like griseipectus,but this characteristic may be age-related. Themost evident difference between griseipectus andnominate picta is the dark central patterri ofthe breast feathers, lanceolate in the latter andsquarish in former, and the apparently evenstronger mandible of R p. picta.

Although the origin of the Atlantic foresttaxa from Amazonian ancestors (not necessarilythe same) seems plausibl¡; the history of pyrrhu-ra pfrimeri presents difficulties. Its habitat isquite different from those of all other taxa in thepicta-leucotis complex, including the similar Rleucotis, and the form of pyrrhura picta geo-graphically closest (amazonum) is too different incolor pattern to be considered a near relative.The closest morphological similarity with Rleucotis suggest a colonization of the centralBrazilian forests from the Atlantic forest, proba-bly through the "Januária bridge" that connectedthe Parana valley with the upper Doce riverduring wetter periods (Willis 1992). The exist-ence of endemic birds of the Atlantic forest inthe isolated mountains of Chapada Diamantina,central Bahia (Gonzaga et al. 1995) also point tothe fact that, in northeastern Brazil, rainforestwith Atlantic affinities was at one time muchmore extensive further inland than it is todayand this may have been the source of theancestral stock from which R pfrimeri originated.

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ECOLOGY OF PYRRHURA PFRIMERI

After the recent shrinking of the rainforestswith the drying of the climate, only the flowerand fig-bearing dry forests offered conditions forthe survival of a population of Pyrrhura of theleucotis group west of the Serra Geral. Theproblem with this hypothesis is the absence of aPyrrhura in the dry forests east of the SerraGeral, but the di:ier climate of this area (meanannual precipitation around 900-1,000 mm,ppposed to 1,200-1,300 mm on the westernsl'¡)pes, RADAMBRASIL 1989) may be theclue to answer this question. The dry season isprobably just too long for a forest pyrrhura tolive east of the Serra Geral, despite the presenceof dry forest. The more modern techniques ofmolecular biology and DNA analyses may, oneday, shed light on this subject, and on therelationships of the pyrrhura picta-Ieucotis com-plex, one of the most interesting problems in

Neotropical parrot taxonomy.

ACKNOWLEDGMENTS

This work was funded by the ZoologischeGesellschaft für Arten und Populationsschutz.Special thanks to Thomas Arndt and RolandWirth for making the project feasible. We aregreatly indebted to Antonio and Cleide Nevesfor providing alllogistical support. Marco Auré-lio Pizzo and Isaac Simao provided data on thefood ecology of Atlantic forest pyrrhura, JoséFernando Pacheco and Douglas Stotz made help-ful comments on the distribution of eastern Bra-zilian conures and on the manuscript, and Mau-ro Galetti provided some important insights onthe importance of flowers to seasonal forestparrots. Kleber Ramos Alves from Instituto Gea-brasil provided information on Terra Ronca StatePark. The existence of this park, the best hopefor the Brazilian dry forests, is owed to theperseverance and hard work of Kleber and hiscompanions at Geabrasil.

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APPENDIX

pyrrhura pfrimeri. Brazil: Cana Brava, Nova Roma,Goiás: MZUSP 15766, 15768, 15770 (lO', 29 9 );Barra do rio sao Domingos, Goiás: MZUSP 15765,15767, 15769 (2 O' , 1 9 ); Santa Maria de Taguatinga,Goiás: MZUSP 11333 (type specimen, unsexed).

pyrrhura griseipectus. Brazil: Serra do Baturité, Ceará:MZUSP 33046, 33047,41509, 41510,41511, 41512,

41513,41514,41515,41516,41517 (8O', 49).

pyrrhura leucotis. Brazil: Rio Muriaé, Rio de Janeiro:MZUSP 27269, 27270 (lO', 19 ); Rio Doce, MinasGerais: MZUSP 6734, 24808, 24810, 24812, 24813,24814, 24816, 24817 (4O', 49), Itabuna, Bahia:MZUSP 10165 (10'); Jequié, Serra do Palhao, Bahia:MZUSP 13989 (10'); rio sao José, Espírito Santo:MZUSP 28103 (10'); rio Itaúnas, Espirito Santo:MZUSP 34495 (10').

pyrrhura picta picta. Brazil: Mazagao, Amapá: MZUSP42862 (19); British Guyana: MZUSP 6496 (10').

132