Brazilian Central Plateau Research Articles

Abundance of epigaeic arthropods in a Brazilian savanna under different fire frequencies

ABSTRACT. Fire is a major determinant of structure and dynamics in savannas, and the rapid increase of human activities inthis biome has changed the natural burning regime. The effects of fire on the fauna of the cerrado (Brazilian savanna) arestill poorly understood, and studies comparing sites frequently and infrequently burned are scarce. In this study, theabundance of epigaeic arthropod orders and trophic guilds was assessed in cerrado sites located in the Brazilian CentralPlateau that were subjected to three burning frequencies: frequent (HighFi), intermediary (MidFi), and infrequent (LowFi).In general, we found a positive relationship between the abundance of epigaeic arthropods and fire frequency. Whenarthropods were analyzed by orders, the abundance of Collembola and Orthoptera was lower in the LowFi site, while forHemiptera, it was higher in the MidFi site. No significant differences were found for Hymenoptera, Coleoptera, and Araneae.The abundance of detritivores and herbivores decreased from HighFi to LowFi, but did not change significantly for omni-vores and predators. These results indicate that some arthropod groups may not only be resilient to fire effects, but actuallymight benefit from fire effects in the cerrado. Characterizing arthropod responses to burning frequency at high taxonomicor functional levels is important for applied studies. Based on the results of the current study, springtails and ants seem tobe particularly appropriate focal groups for further exploratory studies on the effects of fire at the species level.KEY WORDS. Arachnida; burning; cerrado; Insecta; trophic guilds.

Composição florística e fitofisionomia de remanescentes disjuntos de Cerrado nos Campos Gerais, PR, Brasil - limite austral do bioma

O cerrado possui seu limite austral de ocorrência nos Campos Gerais do Paraná onde, na forma de pequenos fragmentos disjuntos, tem grande afinidade com as formações savânicas de São Paulo e do Planalto Central do Brasil. Encontra-se, na sua maior parte, em propriedades particulares, sob forte pressão da agropecuária, tendo sido até então pouco estudado. Foram realizadas análises florísticas e fitofisionômicas de 30 remanescentes de cerrado em seis municípios, com determinação de um total de 1.782 táxons. Asteraceae foi a família com maior riqueza de espécies (256), seguida por Fabaceae (207), Poaceae (183), Myrtaceae (72) e Melastomataceae (54). Encontrou-se um gradiente latitudinal e geomorfológico na distribuição das fisionomias: cerrado stricto sensu (50%) e cerradão (13%) distribuem-se nos planaltos areníticos ao norte da região; enquanto cerrado rupestre (17%) e campo sujo com fácies de cerrado (20%) estão mais representados nas porções meridionais, nos relevos escarpados dos vales dos rios. Os remanescentes mostraram similaridade florística com áreas core do Bioma e podem ser consideradas áreas marginais da amplitude geográfica de muitos táxons, compartilhando espécies típicas, características da Província Sulina. Portanto, estas áreas campestres seriam melhor classificadas como savanas do que como estepes. A fitocenose mais austral em Ponta Grossa, com alta riqueza de espécies, encontra-se isolada biogeograficamente dos demais elementos do Bioma, constituindo um possível centro de endemismo.

Crustacea, Cladocera, Anomopoda, Ilyocryptidae, Ilyocryptus sarsi Stingelin, 1913: a new record and geographic distribution in Brazil

Ilyocryptus sarsi Stingelin, 1913 has a wide distribution in South America, with records from Brazilian states of São Paulo, Mato Grosso do Sul, Minas Gerais, Bahia, Maranhão, Pará, and Amazonas. The present paper reports, for the first time, the occurrence of I. sarsi in the Brazilian Central Plateau. The specimens were found in a wet campo in a natural protect area Gama-Cabeça-de-Veado in Brasília, Federal District, during the rainy season.

Florística e estrutura da vegetação arbustivo-arbórea em uma área de cerrado rupestre no parque estadual da Serra de Caldas Novas, Goiás

O cerrado sentido restrito é a fitofisionomia predominante do bioma Cerrado, ocupando grandes extensões do Planalto Central brasileiro. Quando essa vegetação se desenvolve sobre Neossolos Litólicos recebe a denominação de cerrado rupestre. O conhecimento sobre as comunidades arbustivo-arbóreas deste subtipo fitofisionômico do Cerrado é muito limitado. O objetivo deste estudo foi analisar a composição florística e a estrutura da vegetação arbustivo-arbórea em uma área de cerrado rupestre localizada no Parque Estadual da Serra de Caldas Novas (PESCAN), Goiás, Brasil e comparar suas características florísticas e estruturais com aquelas de outras áreas de cerrado sentido restrito localizadas no Brasil Central. Foram amostrados todos os indivíduos com diâmetro a 30 cm do solo > 5 cm, presentes em 10 parcelas aleatórias de 20 × 50 m. A riqueza florística registrada foi de 66 espécies, distribuídas em 53 gêneros e 31 famílias botânicas. A vegetação estudada tem características florísticas e estruturais semelhantes àquelas registradas em outras áreas de cerrado sentido restrito sobre solos profundos, porém, com a presença de algumas espécies típicas de cerrado rupestre, como Schwartzia adamantium (Cambess.) Gir.-Cañas e Wunderlichia mirabilis Riedel ex Baker e algumas raras para o bioma, por exemplo, Peltogyne confertiflora (Mart. ex Hayne) Benth., Myrcia canescens O. Berg e Myrcia variabilis DC. A área basal encontrada foi relativamente elevada (12,39 m².ha-1) devido à alta densidade de indivíduos (1.357 ind.ha-1), bem como a abundância de indivíduos de grande porte, de espécies como Sclerolobium paniculatum Vogel, Pterodon pubescens (Benth.) Benth., S. adamantium e P. confertiflora. Dessa forma, o substrato raso sob o cerrado rupestre do PESCAN, não limitou o desenvolvimento da vegetação arbustivo-arbórea, nem condicionou o estabelecimento de uma flora diferenciada. No entanto, a elevada importância estrutural de poucas espécies lenhosas típicas e adaptadas aos ambientes rupestres garante certa peculiaridade florística à comunidade estudada.

Shrinkage of microaggregates in Brazilian Latosols during drying: significance of the clay content, mineralogy and hydric stress history

SummaryLatosols (Oxisols, Ferralsols) are characterized by a poor soil horizon differentiation, with a strong microgranular structure formed by microaggregates 50 to 300 μm in size. This microgranular structure is generally considered to be very stable and the shrinkage properties of microaggregates as poor or absent. The objective of our study was to analyse the shrinkage properties of microaggregates in diagnostic Bw horizons collected in Latosols with various particle size distributions and mineralogical compositions depending on their location in a regional toposequence across the Brazilian central plateau. We measured the water retention properties at −300 and −1500 kPa with a centrifugation method. The morphology and size of elementary particles were examined by transmission electron microscopy; specific surface area (SSA) was determined by nitrogen adsorption. We also determined the pore size distribution of microaggregates by the combined use of mercury intrusion and the nitrogen desorption isotherm. Our results show that microaggregates of the Latosols studied were not rigid and shrank during drying. The pore volume of the −300 and −1500 kPa‐treated samples, that of the dried microaggregates and the SSA were closely related to the clay content. The mineralogy of particles <2 μm played a secondary role in the properties studied. Our results show that microaggregate shrinkage at −300 kPa and −1500 kPa varied with the location of the Latosols in the landscape. Thus, microaggregates of Latosols located on the upper highland surface of Late Tertiary shrank less than those of the lowland surface of Late Quaternary, and this difference was attributed to their different hydric stress histories.Propriétés de retrait des microagrégats de Latosols brésiliens: importance de la teneur en argile, de la composition minéralogique et de l’histoire hydrique.RésuméLes Latosols (Oxisols, Ferralsols) sont caractérisés par une faible différenciation de leurs horizons et une structure microgranulaire représentée par des microagrégats de 50 à 300 μm de diamètre. Ces microagrégats sont généralement considérés comme étant très stables et leurs propriétés de retrait comme étant très limitées, voire absentes. L’objectif de cette étude est d’analyser les propriétés de retrait des microagrégats dans l’horizon diagnostic Bw de Latosols de composition granulométrique et minéralogique variable en fonction de leur position le long d’une toposéquence régionale sélectionnée pour l’étude dans le Plateau Central brésilien. Nous avons mesuré les quantités d’eau retenues à−300 et −1500 kPa en utilisant la méthode par centrifugation. La morphologie et la taille des particules élémentaires ont étéétudiées en microscopie électronique à transmission et à l’aide de mesure de surface BET. Nous avons aussi déterminé la distribution de taille des pores dans les agrégats déshydratés en combinant les données de la porosimétrie au mercure et celles issues de l’isotherme de désorption d’azote. Nos résultats montrent que les microagrégats des Latosols étudiés ne sont pas rigides et diminuent de volume lors de la dessiccation. Le volume poral des microagrégats à−300 et −1500 kPa ainsi que leur volume poral après déshydratation sont étroitement liés à la teneur en argile. En revanche, la composition minéralogique de la fraction <2 μm ne jouerait au plus qu’un rôle secondaire dans les variations du volume poral. Nos résultats ont montré que les propriétés de retrait des microagrégats varient en fonction de la position des Latosols dans le paysage. Ainsi, les microagrégats des Latosols situés sur les hauts plateaux de la fin du Tertiaire diminuent moins de volume lors de la dessiccation que les agrégats situés sur les surfaces du Quaternaire récent et situées en contrebas dans le paysage. Une telle différence de comportement serait liée à l’histoire hydrique des Latosols, ceux situés sur les surfaces les plus anciennes ayant vraisemblablement subi des contraintes hydriques plus élevées que ceux situés sur les surfaces les plus récentes.

Astyanax xavante, a new species of characid from middle rio Araguaia in the Cerrado region, Central Brazil (Characiformes: Characidae)

Astyanax xavante is described from a creek of the middle rio Araguaia drainage located in the Cerrado region of the Brazilian Central Plateau. The new species is distinguished from congeners by the combination of pigmentation, morphometric, meristic, morphologic, and tooth features. Ecological notes for A. xavante are included.

Development and origin of the microgranular structure in latosols of the Brazilian Central Plateau: Significance of texture, mineralogy, and biological activity

Brazilian Latosols are characterized by low activity clay, little horizon differentiation, a weak macrostructure and a strong microgranular structure. The development of the latter that was reported as being possibly related to the fine material mineralogy, location in the landscape, parental material and biological activity is still under discussion. The aim of this study is to discuss the origin of the microgranular structure of Latosols located in the Brazilian Central Plateau. Ten Latosols (L) developed in different parent materials were selected along an approximately 350-km long regional toposequence across the South American Surface (SAS) (L1 to L4) and Velhas Surface (VS) (L5 to L10). The structure of the Latosols was studied in the field and samples of the diagnostic Bw horizons were collected for laboratory analyses. Basic soil characterization was performed on the <2-mm material according to the Brazilian standard procedures. The SiO2, AL2O3, Fe2O3 and TiO2 content was determined after dissolution with 1:1 H2SO4. The microstructure was studied in optical microscopy and electron scanning electron microscopy in thin sections. The results showed a varying compound strong microgranular structure and weak to moderate medium sub-angular blocky structure independently of the landscape position as well as the parental material, except for the Latosol developed on a quartzite. The Latosols were classified as gibbsitic-sexquioxic, kaolinitic non-sesquioxid, or kaolinitic-sesquioxid Latosols but there was no relationship between their composition and the development of the microgranular structure in the Bw horizon. The analyses showed indeed several types of microgranular structure with different proportions but without any relationship with their texture and mineralogy. Actually, the types of microgranular structure appeared to be mainly related to soil bioturbation by termites and eventually secondarily by ants. Thus, the weak macrostructure and strong microgranular structure of the Latosols studied, the little vertical differentiation of the horizons, and the lack of a clear relationship between their landscape position and parent material characteristics would result from long-term biotic action with high probability.

Variation of the kaolinite and gibbsite content at regional and local scale in Latosols of the Brazilian Central Plateau

The mineralogy of the Latosols of the Brazilian Central Plateau remains under discussion in the absence of a clear relationship with their age according to their geomorphic location. The aim of this study was thus to clarify the origin of the kaolinite and gibbsite content variation by studying a regional toposequence and using data from the literature. Chemical composition and soil color were used to discuss mineralogy. The mineralogy of the clay fraction was also investigated using X-ray diagrams. Our results showed that the large variation of kaolinite and gibbsite content can be explained by taking into account both their local and regional location, the variation of the hematite and goethite content remaining limited. The model that is proposed to explain such variation combines a regional component, which is mainly associated to the age of the geomorphic surface and a local component which is mainly associated to the hydraulic conditions along the toposequence.

Taxonomic notes on Pogonieae (Orchidaceae): Cleistesiopsis, a new genus segregated from Cleistes, and description of two new South American species, Cleistes batistana and C. elongata

A new genus, Cleistesiopsis, was segregated from Cleistes based on morphological and molecular characteristics, and two new species of Cleistes: C. batistana and C. elongata, both occurring in the Brazilian Central Plateau (Central-Western Brazil), are described and illustrated. Furthermore, a key to genera currently recognised within Pogonieae is presented.

Soil chemical factors and grassland species density in Emas National Park (central Brazil)

Studies of grasslands on specific soil types suggest that different nutrients can limit biomass production and, hence, species composition and number. The Brazilian cerrado is the major savanna region in America and once covered about 2 million km(2), mainly in the Brazilian Central Plateau, under seasonal climate, with wet summer and dry winter. In view of the importance of soil chemical factors in the distribution of the vegetation forms within the Cerrado domain and which may influence the number of species, we analyzed some soil characteristics in three herbaceous vegetation forms -- hyperseasonal cerrado, seasonal cerrado, and wet grassland -- in Emas National Park, a core cerrado site, to investigate the relationship between number of species and soil characteristics. We collected vegetation and soil samples in these three vegetation forms and submitted the obtained data to multiple linear regression. We found out that aluminum and pH were the best predictors of species density, the former positively related to species density and the latter negatively related. Since the predictable variation in species density is important in determining areas of conservation, we can postulate that these two soil factors are indicators of high species density areas in tropical grasslands, which could be used in selecting priority sites for conservation.

Características produtivas e reprodutivas de vacas Holandesas e mestiças Holandês × Gir no Planalto Central

Com o objetivo de comparar o desempenho leiteiro e reprodutivo de vacas da raça Holandesa e mestiças Holandês × Gir na região do Planalto Central foram utilizados dados de 1.456 vacas de cinco composições raciais, originadas das raças Holandesa (H) e Gir (G): puras H; ¾H ¼G; ½H ½G; ³/8H 5/8G; e ¼H ¾G. Analisaram-se as características produção média diária de leite (PMD), número de dias em lactação (DL) e produção total da lactação corrigida para 305 dias (P305), idade ao primeiro parto (IPP), período de gestação (PG) e intervalo de partos (IEP). Os maiores valores de PMD e P305 foram obtidos para os animais ¾H ¼G e ½H ½G e os menores, para os animais com maior proporção da raça Gir (¼H ¾G). Os valores das características reprodutivas IPP e IEP, no entanto, foram menores para os animais ¾H ¼G.Esses resultados evidenciam a adaptação das vacas mestiças ao ambiente a que foram submetidas ou que os animais puros da raça Holandesa, em razão do estresse nutricional e/ou térmico, não expressaram todo o seu potencial genético para produção leiteira.Além disso, os dados comprovaram a importância da utilização de sistemas de cruzamento na manutenção da produção dos animais e na sua adaptação ao ambiente.

RE: o refazer do Sertão-Brasília

As histórias de vida dos sertanejos que participam do vídeo Sertão-Brasília e as dos seus filhos e netos residentes em Brasília denunciam a crise instalada no sertão do Grande Sertão: Veredas, de João Guimarães Rosa. Ela advém do pacto de modernidade firmado entre Juscelino Kubitchek e o capital financeiro internacional, no início da construção de Brasília. Fundado no paradigma da complexidade de Edgar Morin, nas metáforas do pacto e do namoro extraídas do romance rosiano e nos depoimentos que integram o vídeo, este artigo aponta a cultura sertaneja como fonte de possibilidades para saída do pacto, reorganização do sistema Sertão- Brasília e reconquista da sustentabilidade.

Characidium xanthopterum (Ostariophysi: Characiformes: Crenuchidae): a new species from the Central Brazilian Plateau

Characidium xanthopterum is described from tributaries of the upper rio Paraná and upper rio Tocantins basins, in the Central Brazilian Plateau, Goiás State, Brazil. The new species is diagnosed among congeners by the absence of dark bars on the sides of the body in adult specimens, and by the deep yellow coloration in all fins. Ontogenetic change of color pattern is recorded for the first time for Characidium species. Specimens smaller than 32 mm SL possess dark bars on body. These bars disappear with growth between 32 and 35 mm SL, and are always absent in individuals larger than 35 mm SL.

Um estudo de Arqueologia Regional no médio curso do rio Tocantins, TO, Planalto Central brasileiro

Este artigo apresenta alguns dos resultados da pesquisa realizada na bacia do córrego Água Fria, médio curso do rio Tocantins, TO, Brasil. Teve como objetivo identificar os sistemas de assentamento dos grupos sociais que ocuparam a área ao longo do tempo e a forma como esses grupos organizaram o espaço interno de suas áreas de habitação e de atividades. Isso implicou na realização de levantamentos sistemáticos em âmbito regional e intervenções nos 38 sítios arqueológicos identificados. Como resultado elaboramos um modelo interpretativo de organização e uso deste espaço desde há 12.000, quando inicia a ocupação da área por grupos de caçadores e coletores, até o advento dos grupos ceramistas agricultores. Suas conclusões contribuem para as discussões sobre as rotas de povoamento antigo e o surgimento, deslocamentos e crescente complexidade sociopolítica que teria ocorrido entre as sociedades de ceramistas agricultores do Brasil Central e das regiões vizinhas.

Vochysia palmirana (Vochysiaceae), a new species from Goiás and Tocantins, Brazil

Vochysia palmirana is described and illustrated as a new species. It is a member of sect.Vochysiella subsect.Decorticantes. It grows in sandy Cerrados of the Central Brazilian Plateau and is similar toV. sessilifolia. A key to the species ofVochysia sect.Vochysiella subsect.Decorticantes is provided.

Preliminary toxicity study of dichloromethane extract of Kielmeyera coriacea stems in mice and rats

Kielmeyera coriacea Mart. (Clusiaceae), known as “Pau Santo” or “Saco de Boi” in the central Brazilian plateau region, is used to treat several tropical diseases. The present study evaluated the toxic effects of dichloromethane (DcM) extract of Kielmeyera coriacea stems, administered to rodents. In the acute toxicity tests, mice receiving doses of this extract by the oral and intraperitoneal routes, showed reversible effects, with LD 50 values of 1503.0 and 538.8 mg/kg, respectively. In the repeated-dose oral (90 days) toxicity tests, male and female Wistar rats were treated by gavage with different doses of DcM extract (5, 25 or 125 mg/kg). In biochemical and haematological evaluations, the results varied widely in respect to dose and sex, with no linear profile, and did not show clinical correlations. In the histopathological examinations, the groups exhibited some changes, but there were no significant differences between the groups compared to the controls. In conclusion, these investigations appeared to indicate the safety of acute and repeated oral administration of the DcM extract of Kielmeyera coriacea stems, which can therefore be continuously used with safety.

Hydraulic properties of the diagnostic horizon of Latosols of a regional toposequence across the Brazilian Central Plateau

The Brazilian Central Plateau covers about 40% of the Cerrados Biome and represents 24% of the Brazilian territory. The Latosols correspond to about 40% of the Central Plateau. We analyzed the hydraulic properties of a set of Latosols varying according to clay content and mineralogy along a regional toposequence across the Brazilian Central Plateau. We studied the properties of the diagnostic horizon (Bw) of Latosols located on the South American and Velhas Surfaces. We measured the bulk and particle density, the water retention properties at − 1, − 6, − 10, − 33, − 300, and − 1500 kPa by using the centrifugation method, and determined the saturated hydraulic conductivity ( K s) in the field using the Guelph permeameter. Results showed that the total pore volume explained a proportion of the variance of the water retained that decreased with the water potential. On the other hand, the clay content explained a proportion of that variance that increases when the water potential decreased. The great proportion of variance (90.7%) explained for by the clay content alone at − 1500 kPa showed that there is little variability that can be attributed to clay mineralogy variation. The K s value was related to an effective porosity for the Bw of Latosols studied and was defined as the volume proportion of pore with an equivalent diameter of > 300 μm, since it was occupied by air at − 1 kPa. Results showed that water retention properties and K s varied mainly according to the clay content and development of large pores without any close link with the mineralogy of the clay fraction.

Reproductive and productive performance of water buffaloes in central plateau of Brazil

Data were collected from 1994 to 2006 at crossbred water buffaloes of Jafarabadi, Murrah and Mediterranean breeders, in Quirinópolis (18o33’S, 50o27’W and 470m of altitude), Goiás, Brazil. This buffalo herd was collectively maintained, with bovine, in pasture of 75% of Panicum maximum and 25% of Andropogon gayanus. The carrying capacity was of 1.2 AU/ha (about 540 kg LW/10,000 m2). The annual birth rate average was 89.4%, ranged between 75 and 100%. The calving distribution (n=104) from January to December was: 3.74, 3.08, 24.30, 34.58, 14.95, 5.61, 0.00, 1.87, 0.00, 0.00, 0.00 and 1.87%. The breeding incidence occurs, more frequently, between April and June, showing a seasonal reproduction. The amount of calving incidence in the first semester of the year was 96.3% of the total. The age to the first calving (n=30) was of 1,114 ± 171 days. The postpartum interval, for an estimate gestation period of 315 days, was 94 ± 105 days. The average calving interval (n=76) was 409 ± 105 days. The average birth weight was 33.3 ± 2.6 kg; 33.5 ± 2.6 for males and 32.9 ± 2.6 kg for females, but no sex difference (p<0.05). The calves’ weight at weaning time was 232.4 ± 30.9 kg at 265 ± 37 days of age; 237.0 ± 30.5 kg and 265 ± 38 days for males and 227.3 ± 31.0 kg and 267 ± 35 days for females. The average daily weight gain from birth to weaning was 0.874 ± 0.162 kg; 0.905 ± 0.135 kg for males and 0.842 ± 0.182 kg for females, but no sex difference (p<0.05). The pre weaning mortality of calves and among adults was 2.9 and 3.8%, respectively. These results, except for the average weaning weight, which were better, agree with available data from Brazilian literature, although some concerning exists for calves up to 2.2 months older.

Demographic history of wood stork (Mycteria americana) Brazilian Pantanal colonies revealed by mitochondrial DNA analysis

We used mitochondrial DNA (mtDNA) sequences to investigate the demographic history of the wood stork (Mycteria americana) populations in the Brazilian Pantanal. Sequences of 390/460 bp fragment of the mtDNA control region were analyzed in 62 wood stork specimens from 8 colonies using neutrality tests, phylogeographic, and coalescent analyses. Population expansion was supported by the significantly negative values of Tajima’s (D = -2.071) and Fu’s (Fs = -14.544) statistics and the unimodal pattern of mismatch distribution. Nested clade analyses indicated a historic range expansion event and recurrent gene flow that was restricted by isolation by distance as explanations for the haplotype distribution among the sampled colonies. High genetic diversity and the strictly unidirectional gene flow pattern emphasized the conservation importance of preserving the southern Pantanal colonies. Coalescence analyses suggested that northern and southern colonies diverged approximately 6,250 years before the present (YBP), and that their most recent common ancestor was approximately 18,900 YBP. Our results suggest that the contemporary wood stork Pantanal population originated from a more geographically limited founder population. Potential source populations may have occurred in the southern Pantanal or ancestry may reside in populations inhabiting the Brazilian central plateau or areas closer to the equatorial region.

Flora de um brejo de altitude na escarpa oriental do planalto da Borborema, PE, Brasil

Os estudos nas florestas montanas nordestinas (brejos de altitude) indicam que as ombrófilas apresentam elevada riqueza de espécies e são mais relacionadas com a floresta ombrófila de terras baixas. Considerando o desconhecimento sobre as florestas ombrófilas sub-montanas, foi realizado o levantamento do Brejo de Bonito, Pernambuco, com o objetivo de conhecer seu perfil florístico. São apresentadas informações sobre o coletor/número, hábito e padrão de distribuição de cada espécie. Os resultados foram comparados a 22 levantamentos da Floresta Atlântica (latu sensu) nordestina. No brejo de Bonito foram coletadas 217 espécies, distribuídas em 65 famílias. Dessas, cerca de 50% são citados em outras florestas montanas de Pernambuco e de terras baixas de Pernambuco e da Bahia. A maior parte das espécies encontradas tem distribuição na América do Sul, ocorrendo desde a porção norte, penetrando na costa leste brasileira e também na porção central do Brasil e, às vezes, no domínio do semi-árido, nos Brejos de Altitude. Outra porção representativa distribui-se desde a América Central até a porção centro-norte da América do Sul, com poucas ocorrendo também na parte sul da América do Norte (Estados Unidos e México).