Lowland Tropical South America Research Articles

Dipole Response of Millennial Variability in Tropical South American Precipitation and δ18Op during the Last Deglaciation. Part I: Rainfall Response

Abstract Oxygen isotope speleothems have been widely used to infer past climate changes over tropical South America (TSA). However, the spatial patterns of the millennial precipitation and precipitation δ18O (δ18Op) response have remained controversial, and their response mechanisms are unclear. In particular, it is not clear whether the regional precipitation represents the intensity of the millennial South American summer monsoon (SASM). Here, we study the TSA hydroclimate variability during the last deglaciation (20–11 ka ago) by combining transient simulations of an isotope-enabled Community Earth System Model (iCESM) and the speleothem records over the lowland TSA. Our model reasonably simulates the deglacial evolution of hydroclimate variables and water isotopes over the TSA, albeit underestimating the amplitude of variability. North Atlantic meltwater discharge is the leading factor driving the TSA’s millennial hydroclimate variability. The spatial pattern of both precipitation and δ18Op show a northwest–southeast dipole associated with the meridional migration of the intertropical convergence zone, instead of a continental-wide coherent change as inferred in many previous works on speleothem records. The dipole response is supported by multisource paleoclimate proxies. In response to increased meltwater forcing, the SASM weakened (characterized by a decreased low-level easterly wind) and consequently reduced rainfall in the western Amazon and increased rainfall in eastern Brazil. A similar dipole response is also generated by insolation, ice sheets, and greenhouse gases, suggesting an inherent stability of the spatial characteristics of the SASM regardless of the external forcing and time scales. Finally, we discuss the potential reasons for the model–proxy discrepancy and pose the necessity to build more paleoclimate proxy data in central-western Amazon. Significance Statement We want to reconcile the controversy on whether there is a coherent or heterogeneous response in millennial hydroclimate over tropical South America and to clearly understand the forcing mechanisms behind it. Our isotope-enabled transient simulations fill the gap in speleothem reconstructions to capture a complete picture of millennial precipitation/δ18Op and monsoon intensity change. We highlight a heterogeneous dipole response in precipitation and δ18Op on millennial and orbital time scales. Increased meltwater discharge shifts ITCZ southward and favors a wet condition in coastal Brazil. Meanwhile, the low-level easterly and the summer monsoon intensity reduced, causing a dry condition in the central-western Amazon. However, the millennial variability of hydroclimate response is underestimated in our model, together with the lack of direct paleoclimate proxies in the central-west Amazon, complicating the interpretation of changes in specific paleoclimate events and posing a challenge to constraining the spatial range of the dipole. Therefore, we emphasize the necessity to increase the source of proxies, enhance proxy interpretations, and improve climate model performance in the future.

Neotropical Clavulina: Two new species from Mexico and a re-evaluation of Clavulina floridana

ABSTRACT Clavulina comprises ca. 90 described species distributed worldwide in both tropical and temperate regions. However, only one species (C. floridana) has been described so far from tropical North America. We used morphological and molecular data from three DNA loci (nuc rDNA internal transcribed spacer region ITS1-5.8S-ITS2 [ITS], a portion of nuc 28S rDNA [28S], and a fragment of DNA-directed RNA polymerase II second largest subunit [RPB2]) from basidiomata and ectomycorrhizas collected in tropical ecosystems from three biogeographic provinces of Mexico and one tropical province in the USA to investigate the phylogenetic and taxonomic diversity of Clavulina in the region. Nine new species-level clades were discovered, two of which are proposed as new species (C. arboreiparva and C. tuxtlasana). Specimens of C. floridana recently collected in Florida were included in our analyses, for which a modern description is provided. In addition, C. floridana is a new record for Mexico. The diversity of Clavulina in tropical North America is comparable to that found in lowland tropical South America. However, some of the species found in tropical deciduous forests produce small, rare, and inconspicuous basidiomata, which easily go unnoticed, and therefore are poorly represented in collections. Many species remain undescribed in tropical regions of North America.

Evolutionary diversity in tropical tree communities peaks at intermediate precipitation

Global patterns of species and evolutionary diversity in plants are primarily determined by a temperature gradient, but precipitation gradients may be more important within the tropics, where plant species richness is positively associated with the amount of rainfall. The impact of precipitation on the distribution of evolutionary diversity, however, is largely unexplored. Here we detail how evolutionary diversity varies along precipitation gradients by bringing together a comprehensive database on the composition of angiosperm tree communities across lowland tropical South America (2,025 inventories from wet to arid biomes), and a new, large-scale phylogenetic hypothesis for the genera that occur in these ecosystems. We find a marked reduction in the evolutionary diversity of communities at low precipitation. However, unlike species richness, evolutionary diversity does not continually increase with rainfall. Rather, our results show that the greatest evolutionary diversity is found in intermediate precipitation regimes, and that there is a decline in evolutionary diversity above 1,490 mm of mean annual rainfall. If conservation is to prioritise evolutionary diversity, areas of intermediate precipitation that are found in the South American ‘arc of deforestation’, but which have been neglected in the design of protected area networks in the tropics, merit increased conservation attention.

Using tree species inventories to map biomes and assess their climatic overlaps in lowland tropical South America

AbstractAimTo define and map the main biomes of lowland tropical South America (LTSA) using data from tree species inventories and to test the ability of climatic and edaphic variables to distinguish amongst them.LocationLowland Tropical South America (LTSA), including Argentina, Bolivia, Brazil, Ecuador, Paraguay, Peru and Uruguay.Time periodPresent.Major taxa studiedTrees.MethodsWe compiled a database of 4,103 geo‐referenced tree species inventories distributed across LTSA. We used a priori vegetation classifications and cluster analyses of floristic composition to assign sites to biomes. We mapped these biomes geographically and assessed climatic overlaps amongst them. We implemented classification tree approaches to quantify how well climatic and edaphic data can assign inventories to biomes.ResultsOur analyses distinguish savanna and seasonally dry tropical forest (SDTF) as distinct biomes, with the Chaco woodlands potentially representing a third dry biome in LTSA. Amongst the wet forests, we find that the Amazon and Atlantic Forests might represent different biomes, because they are distinct in both climate and species composition. Our results show substantial environmental overlap amongst biomes, with error rates for classifying sites into biomes of 19–21 and 16–18% using only climatic data and with the inclusion of edaphic data, respectively.Main conclusionsTree species composition can be used to determine biome identity at continental scales. We find high biome heterogeneity at small spatial scales, probably attributable to variation in edaphic conditions and disturbance history. This points to the challenges of using climatic and/or interpolation‐based edaphic data or coarse‐resolution, remotely sensed imagery to map tropical biomes. From this perspective, we suggest that using floristic information in biome delimitation will allow for greater synergy between conservation efforts centred on species diversity and management efforts centred on ecosystem function.

First Record of Leucocytozoon (Haemosporida: Leucocytozoidae) in Amazonia: Evidence for Rarity in Neotropical Lowlands or Lack of Sampling for This Parasite Genus?

Birds harbor an astonishing diversity of haemosporidian parasites belonging to the genera Haemoproteus, Leucocytozoon, and Plasmodium. Currently there are more than 250 morphologically described avian haemosporidian species and 2,828 unique lineages found in virtually all avian clades and zoogeographic regions, except for Antarctica. Our report is based on PCR and microscopic screening of 1,302 individual avian samples from Brazil to detect the underrepresented genus Leucocytozoon. This survey primarily focuses on passerine birds collected from Amazonia, the Atlantic Rain Forest, and Pantanal. We also summarize studies conducted in Brazil that report haemosporidian prevalence using both microscopy and molecular tools and present for the first time a record of Leucocytozoon infecting an avian host population in Amazonia. Based on our findings, we suggest that high average temperatures may be constraining both the distribution and diversity of Leucocytozoon in lowland tropical South America.

Impact of mid- to late Holocene precipitation changes on vegetation across lowland tropical South America: a paleo-data synthesis

AbstractA multi-proxy paleo-data synthesis of 110 sites is presented, exploring the impact of mid- to late Holocene precipitation changes upon vegetation across Southern Hemisphere tropical South America. We show that the most significant vegetation changes occurred in southwest Amazonia and southeast Brazil, regions reliant on precipitation derived from the South American summer monsoon (SASM). A drier mid-Holocene in these regions, linked to a weaker SASM, favored more open vegetation (savannah/grasslands) than present, while increased late-Holocene precipitation drove expansion of humid forests (e.g., evergreen tropical forest in southwest Amazonia,Araucariaforests in southeast Brazil). The tropical forests of central, western and eastern Amazonia remained largely intact throughout this 6000-year period. Northeastern Brazil’s climate is “antiphased” with the rest of tropical South America, but a lack of paleo-data limits our understanding of how vegetation responded to a wetter (drier) mid-(late) Holocene. From this paleo-data perspective, we conclude that ecotonal forests already close to their climatic thresholds are most vulnerable to predicted future drought, but the forest biome in the core of Amazonia is likely to be more resilient. Of greater concern are widespread deforestation and uncontrolled anthropogenic burning, which will decrease ecosystem resilience, making them more vulnerable than they might be without current anthropogenic pressures.

On the importance of geographic and taxonomic sampling in phylogeography: A reevaluation of diversification and species limits in a Neotropical thrush (Aves, Turdidae)

Phylogeographic studies seeking to describe biogeographic patterns, infer evolutionary processes, and revise species-level classification should properly characterize the distribution ranges of study species, and thoroughly sample genetic variation across taxa and geography. This is particularly necessary for widely distributed organisms occurring in complex landscapes, such as the Neotropical region. Here, we clarify the geographic range and revisit the phylogeography of the Black-billed Thrush (Turdus ignobilis), a common passerine bird from lowland tropical South America, whose evolutionary relationships and species limits were recently evaluated employing phylogeographic analyses based on partial knowledge of its distribution and incomplete sampling of populations. Our work employing mitochondrial and nuclear DNA sequences sampled all named subspecies and multiple populations across northern South America, and uncovered patterns not apparent in earlier work, including a biogeographic interplay between the Amazon and Orinoco basins and the occurrence of distinct lineages with seemingly different habitat affinities in regional sympatry in the Colombian Amazon. In addition, we found that previous inferences about the affinities and taxonomic status of Andean populations assumed to be allied to populations from the Pantepui region were incorrect, implying that inferred biogeographic and taxonomic scenarios need re-evaluation. We propose a new taxonomic treatment, which recognizes two distinct biological species in the group. Our findings illustrate the importance of sufficient taxon and geographic sampling to reconstruct evolutionary history and to evaluate species limits among Neotropical organisms. Considering the scope of the questions asked, advances in Neotropical phylogeography will often require substantial cross-country scientific collaboration.

New species of Elaphomyces (Elaphomycetaceae, Eurotiales, Ascomycota) from tropical rainforests of Cameroon and Guyana.

The sequestrate false truffles Elaphomyces favosus, E. iuppitercellus, and E. labyrinthinus spp. nov. are described as new to science from the Dja Biosphere Reserve, Cameroon. Elaphomyces adamizans sp. nov. is described as new from the Pakaraima Mountains of Guyana. The Cameroonian species are the first Elaphomyces taxa to be formally described from Africa, occurring in lowland Guineo-Congolian tropical rainforests dominated by the ectomycorrhizal (ECM) canopy tree Gilbertiodendron dewevrei (Fabaceae subfam. Caesalpinioideae). The Guyanese species is the third to be discovered in lowland tropical South America, occurring in forests dominated by the ECM trees Pakaraimaea dipterocarpacea (Dipterocarpaceae) and Dicymbe jenmanii (Fabaceae subfam. Caesalpinioideae). Macromorphological, micromorphological, habitat, and DNA sequence data are provided for each new species. Molecular and morphological data place these fungi in Elaphomycetaceae (Eurotiales, Ascomycota). Unique morphological features are congruent with molecular delimitation of each of the new species based on a phylogenetic analysis of the rDNA ITS and 28S loci across the Elaphomycetaceae. The phylogenetic analysis also suggests that a common ancestor is shared between some Elaphomyces species from Africa and South America, and that species of the stalked, volvate genus Pseudotulostoma may be nested in Elaphomyces.

Tropical niche conservatism as a historical narrative hypothesis for the Neotropics: a case study using the fly family Muscidae

Aim We evaluate the extent to which the tropical conservatism hypothesis can explain the evolutionary development of the Muscidae. Furthermore, we compare the geographical patterns of muscid phylogenetic structure with biogeographical regions that have been identified for Neotropical insects. Location Central and South America. Methods We modelled the geographic distributions of 658 species using Maxent and 19 environmental variables. A generic-level supertree of the Muscidae was assembled using matrix representation with parsimony and used to map the geographic pattern of mean root distance (MRD), a metric of the relative evolutionary development of assemblages. Regression models (ordinary least squares and regression trees) were used to examine temperature and other environmental correlates of MRD to explore potential environmental drivers of muscid diversification. We used the regression tree results to recognize variable intervals that best explained MRD, and these intervals were mapped to recognize and compare with biogeographical regions of Neotropical insects. Results The geographic pattern of MRD was consistent with the tropical conservatism hypothesis: species in genera that diversified relatively early, as measured by their distance from the tree root, dominate lowland tropical South America, whereas species in genera that diversified more recently occupy extra-tropical areas, sub-Antarctic areas and the Andean highlands. Temperature was the strongest correlate of MRD. Three biogeographical regions were recognized and they coincided with two regions known for insects. Main conclusions Evolutionary responses of muscid flies to post-Eocene climate change taking the form of an expansion of a tropical group into regions with colder climates may be fundamental to explaining their distribution in the Neotropics. Our biogeographical regions delimited by temperature and the phylogenetic metric, surrogates of the tropical conservatism hypothesis, were very similar to general insect patterns, supporting the ‘tropical origin and evolutionary response to climate cooling’ as a broadly based historical narrative for the Neotropics.

Review of plant biogeographic studies in Brazil

Abstract Molecular phylogenetic studies have become a major area of interest in plant systematics, and their impacts on historical biogeographic hypotheses are not to be disregarded. In Brazil, most historical biogeographic studies have relied on animal phylogenies, whereas plant biogeographic studies have largely lacked a phylogenetic component, having a limited utility for historical biogeography. That country, however, is of great importance for most biogeographic studies of lowland tropical South America, and it includes areas from a number of biogeographic regions of the continent. Important biogeographic reports have been published as part of phylogenetic studies, taxonomic monographs, and regional accounts for small areas or phytogeographic domains, but the available information is subsequently scattered and sometimes hard to find. In this paper we review some relevant angiosperm biogeographic studies in Brazil. Initially we briefly discuss the importance of other continents as source areas for the South American flora. Then we present a subdivision of Brazil into phytogeographic domains, and we cite studies that have explored the detection of biogeographic units (areas of endemism) and how they are historically related among those domains. Examples of plant taxa that could be used to test some biogeographic hypotheses are provided throughout, as well as taxa that exemplify several patterns of endemism and disjunction in the Brazilian angiosperm flora.

Floral trimorphism and monomorphism in continental and island populations of Eichhornia paniculata (Spreng.) Solms. (Pontederiaceae)

Eichhornia paniculata (Spreng.) Solms. (Pontederiaceae) is a short-lived perennial or annual of marshes, seasonal pools and ditches of lowland tropical South America, primarily NE Brazil, and the Caribbean islands of Cuba and Jamaica. Comparisons, made under uniform glasshouse conditions, of populations originating from seed collected in the two regions revealed striking differences in their floral biology and breeding systems. The majority of populations sampled in NE Brazil are tristylous. Floral trimorphism is associated with pollen trimorphism and minor differences in anther size and pollen production among the three stamen levels. Unlike the majority of heterostylous plants the floral morphs of E. paniculata are highly self-fertile. Populations sampled on the island of Jamaica are composed exclusively of self-pollinating, semi-homostylous, mid-styled forms. Flowers from these populations are smaller and less showy, with reduced pollen heteromorphism and significantly fewer pollen grains and ovules per flower, in comparison with trimorphic populations from Brazil. Individual genotypes from Jamaican populations display considerable developmental instability in floral expression, particularly with respect to filament elongation of the lower stamen level. It is proposed that self-pollinating populations of E. paniculata are evolutionarily derived from tristylous ancestors and that the shift in breeding system is favoured at low density, following population bottlenecks, where pollinator service is unreliable.

Crinipellis perniciosa (Agaricales)

Crinipellis perniciosa (Stahel) Singer is one of the more serious plant pathogens, being the causal agent of witches' broom disease of cocoa. Distribution is confined to the cocoa areas of lowland tropical South America, with the disease often reaching epidemic levels in Ecuador, Guyana, Surinam, Trinidad and Tobago. It is also recorded from Bolivia, Brazil, Colombia, Grenada and Peru (C.M.I. Distribution Map No. 37, ed. 3, I975). The fungus attacks species of Herrania Goudst and Theobroma L. (Sterculiaceae), although it is not confined to those genera nor is it always parasitic. Desrosiers & Buchwald (1949) reported basidiocarps resembling C. perniciosa on dead lianas, bamboo and unidentified wood in Ecuador, whilst Evans (I977) found typical basidiocarps on both hanging and climbing lianas, fallen debris caught in understorey canopy and on unidentified fallen tree branches on the forest floor. The host is attacked by infection of the breaking vegetative buds, flower-cushions and young fruit. Hyperplasia results and the proliferated growths form the 'witches brooms' which characterize the disease. The disease has been thoroughly reviewed by Baker & Holliday (1957) and Thorold (1975: 11-26). Published descriptions of the agaric basidiocarps are largely confined to the original account given by Stahel (1915: 16), based on material from Surinam, together with additional information on the microcharacters by Singer (1942: 503), following a study of the authentic material. Dennis (1951: 429, pl. 20/4) gave a brief, illustrated account of specimens collected from Theobroma in Trinidad. These accounts were subsequently summarised by Baker & Holliday (1957) and Holliday (1970). Following an extended field investigation of the disease in Ecuador, during 1973-1976, by Dr. H. C. Evans, it has become apparent that the basidiocarps are far more variable than hitherto realized. Constant differences may be recognized in size and pigmentation of the basidiocarp, host range and spore size range. It is decided for these reasons to provide a revised detailed description, recognizing three distinct varieties.

Crinipellis perniciosa . [Descriptions of Fungi and Bacteria

Abstract A description is provided for Crinipellis perniciosa . Information is included on the disease caused by the organism, its transmission, geographical distribution, and hosts. HOSTS: Restricted to Theobroma spp. and Herrania spp. Recorded on: T. bicolor, T. cacao, T. calodesmis, T. glauca, T. grandiflora, T. microcarpa, T. obovata and T. subincana (37: 151); H. albiflora, H. nitida and H. purpurea (39: 684). DISEASE: Witches' broom disease of cacao. Infection of the breaking vegetative buds, flower cushions and young fruit leads to hypertrophy, reduction in leaf size, abnormal proliferation of axillary buds, growth of vegetative shoots from cushions and distortion of the fruit. These pathogenic effects may be due to the production of enzymes which inactivate the host auxins (48, 9g, 2864). The pathogen is generally confined to the hypertrophied areas which eventually become necrotic and later develop sporophores. At the epidemic level (reached in Ecuador, Surinam, Trinidad and Tobago) nearly all the vegetative growing points are infected and killed, up to 75% of the crop being directly destroyed. GEOGRAPHICAL DISTRIBUTION: Confined to parts of lowland tropical South America, Trinidad, Tobago and Grenada (CMI Map 37 ed. 2, 1965). Endemic on wild species of Theobroma in the Amazon and Orinoco river systems (17: 801; 23: 56; 42: 78). TRANSMISSION: Air dispersed basidiospores, released mostly between 1800 hr and 0600 hr, probably do not survive the day following the night on which they are shed. No conidial state is known.