Neotropical flowering epiphyte diversity: local composition and geographic affinities

Worldwide, the highest diversity of vascular epiphytic plants resides in the Neotropics. The general pattern of taxonomic composition of the neotropical epiphytic flora has been described, but information regarding the magnitude and geographic distribution of species richness is lacking. In this paper, we carried out a regional scale analysis in order to provide an overview of the richness, composition and geographic affinities among several neotropical epiphyte floras. Our database comprised 7,524 flowering epiphyte species (48 % of the estimated total of neotropical epiphytes) from eight representative localities with politically-defined boundaries. The epiphyte quotient (e.g., percentage of epiphyte species in the total flora) per locality ranged from 5.2–27.7 % (mean: 17.5 %) of the flowering plants. Ecuador represented the most species diverse locality (4,247 spp.), followed by Costa Rica (2,611 spp.). At the family level, the epiphyte composition among localities was roughly homogeneous and dominated by emblematic epiphytic groups: Orchidaceae, Bromeliaceae, and Araceae. However, the generic composition of the epiphytic floras was more heterogeneous. Ecuador and Cuba showed the highest epiphyte endemism (37 and 26 %, respectively), and in some cases vascular epiphytes represented 30 % or more of the total endemic flowering plants at a particular locality. From the available information, four main regional epiphyte floras were identified: the northwestern Andean region, the northern and southern Mesoamerican regions; Cuba as a representative of the Caribbean region remained as a separated group, though weakly related to the Mesoamerican groups. This study identified important geographic localities as targets for the conservation of neotropical epiphyte diversity.     

The geographic range size and vulnerability to extinction of angiosperm epiphytes in the Atlantic Forest of Brazil

Aim

Angiosperm epiphytes have long been reported to have larger geographic ranges than terrestrial species, despite evidence of their outstanding diversity and endemism. This apparent contradiction calls for further investigation of epiphytes' poorly understood range size patterns. Here, we address the question of whether epiphytes have larger geographic ranges and different vulnerability to extinction than terrestrial species.

Location

The Atlantic Forest of Brazil, a global centre of tropical epiphyte diversity with relatively well-known flora, where we can estimate the geographic ranges of a large number of species with reasonable confidence.

Time period

Occurrence records from the 17th century to the year 2021.

Major taxa studied

Flowering plants (angiosperms).

Methods

We downloaded, processed and cleaned all occurrence records for the angiosperm species native to the Atlantic Forest of Brazil available in the speciesLink network and the Global Biodiversity Information Facility. We estimated the extent of occurrence and area of occupancy of 12,679 native flowering plants, including 1251 epiphytic species. We compared the geographic ranges of epiphytes and other life forms at broad (e.g. Angiosperms, Monocots) and more restricted taxonomic scales (e.g. individual families), assuming species are independent entities and also when accounting for species phylogenetic dependence.

Results

We found that epiphytes have among the smallest geographic ranges of flowering plants. We found no consistent evidence that epiphytism leads to differences in geographic ranges between close relatives. However, both epiphytes and non-epiphytes in epiphyte-rich lineages have small ranges and likely a high vulnerability to extinction.

Main Conclusions

Our findings contrast with the long-held hypothesis that epiphytes have larger geographic ranges than terrestrial species. Epiphytes and their close relatives share many diversification mechanisms and ecological adaptations (‘epiphyte-like traits’), which probably explain why both sets of species have small range sizes and high vulnerability to extinction.     

Chromosome numbers and DNA content in Bromeliaceae: additional data and critical review

For the large Neotropical plant family Bromeliaceae, we provide new data on chromosome numbers, cytological features and genome size estimations, and combine them with data available in the literature. Root‐tip chromosome counts for 46 species representing four subfamilies and a literature review of previously published data were carried out. Propidium iodide staining and flow cytometry were used to estimate absolute genome sizes in five subfamilies of Bromeliaceae, sampling 28 species. Most species were diploid with 2n = 50 in Bromelioideae, Puyoideae and Pitcairnioideae, followed by 2n = 48 observed mainly in Tillandsioideae. Individual chromosome sizes varied more than tenfold, with the largest chromosomes observed in Tillandsioideae and the smallest in Bromelioideae. Genome sizes (2C‐values) varied from 0.85 to 2.23 pg, with the largest genomes in Tillandsioideae. Genome evolution in Bromeliaceae relies on two main mechanisms: polyploidy and dysploidy. With the exception of Tillandsioideae, polyploidy is positively correlated with genome size. Dysploidy is suggested as the mechanism responsible for the generation of the derived chromosome numbers, such as 2n = 32/34 or 2n = 48. The occurrence of B chromosomes in the dysploid genus Cryptanthus suggests ongoing speciation processes closely associated with chromosome rearrangements. © 2014 The Linnean Society of London, Botanical Journal of the Linnean Society, 2014, 176 , 349–368.     

Floristic patterns of epiphytes in the Brazilian Atlantic Forest,a biodiversity hotspot

To assess the main factors driving epiphytic angiosperm distribution throughout the Brazilian Atlantic Forest, we compiled 57 floristic surveys and analysed species composition under the influence of environmental variables, space and vegetation type using canonical correspondence analysis (CCA), similarity (Sorensen) and Mantel's tests. The indicator value index (IndVal) was used to find indicator species of each Brazilian Atlantic Forest vegetation type. Group sharpness analysis was performed in order to determine the appropriate group partition level. CCA showed a separation of the epiphytic flora reflecting temperature and rainfall gradients. Mantel's test showed that environment and space were highly correlated with floristic similarity. Cluster analysis, indicating floristic similarity, resolved five groups, mainly grouped by region. Clear differentiation of the Brazilian Atlantic Forest epiphytic flora on a north–south axis with a strong correlation with temperature and rainfall gradients was found. The role of space and environment on species composition varied according to distinct epiphytic species groups. In particular, for Bromeliaceae and Orchidaceae, the main factor associated with floristic similarity was space. Indicator species were found for all vegetation types apart from the Seasonal Semideciduous Forest that seemed to represent a subset of a more humid forest type. © 2015 The Linnean Society of London, Botanical Journal of the Linnean Society, 2015, 179 , 587–601.     

Arbuscular mycorrhizas and dark septate endophytes in bromeliads from South American arid environment

Most plant roots are associated with glomalean fungi forming arbuscular mycorrhizas (AM) and a wide range are also colonized by ascomycetous dark septate endophytes (DSE). Bromeliaceae species can be epiphytic, rupicolous or terrestrial but their mycorrhizal status is poorly studied. We examined the AM and DSE status of 5 epiphytic and 4 terrestrial Bromeliaceae from an arid area of Central Argentina. The terrestrial species were either dually associated (AM and DSE) or non-associated whereas the epiphytes were only DSE colonized. Terrestrial Bromeliaceae that formed AM-DSE associations were likely responding to the arid conditions of the area and the availability of AM fungal (AMF) spores in the soil. The terrestrialBromelia ubaniana was not colonized either by AMF or DSE. This could reflect its root morphology and high number of root hairs. DSE are endosymbiotic in the stressful ecosystems experienced by canopy epiphytes in the studied environment. The different fungal associations are discussed in relation to the three Bromeliaceae subfamiles and we suggest that environmental features determinethe type of association formed by species in this plant family.     

Exotic tree monocultures play a limited role in the conservation of Atlantic Forest epiphytes

The Brazilian Atlantic Forest suffered a severe geographic contraction along the last five centuries that reduced drastically most vascular epiphyte populations. Among the range of man-made matrixes, tree monocultures have the potential to contribute positively to the maintenance of the regional epiphyte diversity. Here, we test the similarity in abundance, richness, and species composition between vascular epiphytic communities established in managed monocultures of exotic and native species with natural communities occurring in neighboring native Araucaria Forest patches. In the São Francisco de Paula National Forest (Rio Grande do Sul state, Brazil), we recorded 62 epiphyte species from 300 phorophytes occurring in 12, one-hectare plots of Araucaria Forest and managed plantations of Pinus, Eucalyptus and Araucaria. Species richness, rarefied richness and abundance were significantly higher in Araucaria Forest in comparison to the exotic stands. Species composition was also substantially differentiated as Araucaria Forest patches harbored a greater number of zoochorous species than those of the exotic stands. Additionally, plantations of Araucaria angustifolia, a native species, sustained more individuals and more species than the exotic plantations. Neither tree height nor DBH explained epiphyte richness; however, both phorophyte diversity and stand age together accounted for 92% of the among-site variation in epiphytic species richness. We conclude that substrate heterogeneity in combination with time available for colonization contribute significantly to beta-diversity of epiphytes in Araucaria forests. However, demographic experimental studies are necessary in order to disentangle the role of substrate quality from metapopulation processes, such as dispersal limitation, at both temporal and spatial scales.     

Genetic structure of sigmodontine rodents (Cricetidae) along an altitudinal gradient of the Atlantic Rain Forest in southern Brazil

The population genetic structure of two sympatric species of sigmodontine rodents (Oligoryzomys nigripes and Euryoryzomys russatus) was examined for mitochondrial DNA (mtDNA) sequence haplotypes of the control region. Samples were taken from three localities in the Atlantic Rain Forest in southern Brazil, along an altitudinal gradient with different types of habitat. In both species there was no genetic structure throughout their distribution, although levels of genetic variability and gene flow were high.     

First records of Synoeca septentrionalis Richards, 1978 (Hymenoptera, Vespidae, Epiponini) in the Brazilian Atlantic Rain Forest

Nests of Synoeca septentrionalis were collected in two Brazilian Atlantic Rain Forest localities (Itabuna and Santa Terezinha, in the state of Bahia and Alfredo Chaves in the state of Espírito Santo). Synoeca septentrionalis was previously recorded only from Central America and northwestern South America. This findingextends its geographical distribution to Northeast and Southeast regions of Brazil, and represents the first record for Synoeca septentrionalis in the Brazilian Atlantic Rain forest, raising to three the number of Synoeca species known from Bahia State.     

Breeding systems of Bromeliaceae species: evolution of selfing in the context of sympatric occurrence

Co-occurring congener plant species in the highly diverse Atlantic forests of southeastern Brazil may act as natural laboratories for evaluating evolution of reproductive shifts. We assessed the breeding systems in a sympatric assemblage of bromeliad species and compiled literature available for the family to compare our experiments with available information. We performed controlled experiments of autonomous selfing, self- and cross-pollination in 40 species of 11 genera of two subfamilies that, in general, overlap their blooming period and share the same pollination vectors. We also tested for differences between self-compatible (SC) and self-incompatible (SI) species with regard to ecological factors such as abundance, co-flowering and co-occurrence. Most species experimentally tested and surveyed in the literature (75%) were SC. Species from the subfamily Tillandsioideae were predominantly SC, while Bromelioideae showed greater variation in breeding systems. About 43% of the species studied set fruits spontaneously in the absence of pollinator vectors. We found that SC species were the more abundant, were more frequently associated with other relatives and overlapped their blooming period with other species more than SI species. Thus, our results suggest that self-compatibility is common for Bromeliaceae and do not support the traditional hypothesis of reproductive assurance used to explain the evolution of selfing. We discuss self-compatibility as a reproductive isolating mechanism in the presence of pollen flow among sympatric congener species.     

Genome size and base composition of Bromeliaceae species assessed by flow cytometry

Flow cytometry (FCM) has been used to estimate the nuclear DNA content of Bromeliaceae species, which constitutes relevant information for studies of taxonomy, evolution, genetic diversity, and reproductive biology in bromeliads. Nevertheless, C values have only been estimated for 58 out of the 3,140 existing Bromeliaceae species. Aiming to contribute to the genome database of Bromeliaceae, the current study was carried out to measure the nuclear DNA content and base composition of Bromelioideae and Tillandsioideae species occurring in the Atlantic Rainforest. The most adequate FCM procedure provided histograms exhibiting G₀/G₁ peaks with coefficients of variation below 5%, so that these histograms were used to measure the mean 2C and AT% values for all collected Bromelioideae and Tillandsioideae species. These values were statistically compared, and dendrograms were plotted. A second comparison was performed among all mean 2C values reported for Pitcairnioideae, Tillandsioideae, and Bromelioideae species. In accordance with previous statistical comparisons, two groups were formed: cluster 1, composed by Tillandsia loliacea, Tillandsia usneoides, and Tillandsia cyanea, and cluster 2, gathering other 69 species. Based on these results, we concluded that FCM was a rapid, accurate, and reliable technique to assess genome size and base composition. Furthermore, the FCM data reported here will contribute to the Monocot and Bromeliaceae database, which still displays several ongoing gaps, especially for endemic species.     

Synopsis of the tribe Hureae (Euphorbioideae, Euphorbiaceae)

A synopsis of the tribe Hureae is presented with nomenclatural updates, a discussion of diagnostic features, and summaries of geographical distributions. This study is based on the analysis of approximately 300 voucher specimens, including collections and photographs of types, in addition to bibliographic documentation. Seventeen species distributed in three genera were recognized: Algernonia (11 species), Hura (2), and Ophthalmoblapton (4). All species are American with the majority distributed within the Atlantic Forest, particularly in southeastern Brazil. A key for the identification of genera and species is provided along with illustrations, information on the geographic distributions and conservation status. Lectotypification for Algernonia leandrii is proposed.     

Spatial and temporal patterns of land flatworm assemblages in Brazilian Araucaria forests

Abstract. Araucaria forests are among the most threatened biomes of one of the world's 25 biodiversity hotspots, the south Brazilian Atlantic Rain Forest. This study was focused on flatworm community structure in three protected areas located on the eastern border of the Araucaria Plateau in south Brazil. We addressed three main questions: (1) How species‐rich are Araucaria forests? (2) Are there any differences in the community structure within the three areas? (3) What is the distribution pattern of land flatworms in this type of forest? A total of 51 flatworm species in ten genera and two subfamilies were found. Non‐metric multidimensional scaling analysis showed that the assemblage structure differed within the three areas. The abundance and richness varied in time, being affected by an interaction between season and areas. Araucaria forests on the eastern border of the Araucaria Plateau should be considered a hotspot of land planarian diversity, harboring an estimated 64 species. The high combined species richness in the protected areas studied emphasizes the importance of their preservation. Notwithstanding their close proximity, they are not redundant in maintaining regional biodiversity.     

Climate change threatens protected areas of the Atlantic Forest

Only 7 % of the Atlantic Forest Biodiversity Hotspot is currently protected, though it holds 18 % of all amphibian species in South America. How effective would the Atlantic Forest network of protected areas (PAs) be in a changing climate? Are there some intrinsic features of PAs that drive species loss or gain inside them? We addressed these questions by modeling the ecological niches of 430 amphibian species in the Atlantic Forest and projecting their distributions into three future climate change simulations. We then assessed changes in species richness inside PAs for different time frames and tested their significance via null model. The number of species should decline within Atlantic Forest network of PAs under changing climate conditions. Only altitude was a good predictor of species gains or lost inside PAs. Therefore, we suggest that new PAs established in highlands would be more effective to alleviate the effects of climate change on this imperiled fauna.     

Comparative ecophysiology of CAM and C3 bromeliads. I. The ecology of the Bromeliaceae in Trinidad

Abstract This article deals with the physiological ecology of the Bromeliaceae, a large neotropical family containing both terrestrial and epiphytic forms, as well as many species with crassulacean acid metabolism (CAM). The article is in two parts. In the first, we review what is known of the occurrence of CAM and C₃ species in the Bromeliaceae. The photosynthetic pathways are discussed in the context of the major taxonomic divisions within the family and the great diversity of bromeliad life-forms. Of the three subfamilies, the Pitcairnioideae contain both C₃ and CAM species and are essentially all terrestrial. In contrast, the Tillandsioideae are entirely epiphytic or saxicolous, with CAM species being restricted to the genus Tillandsia, And in the Bromelioideae all species show CAM, but terrestrial and epiphytic forms are found in about equal numbers. The evidence suggests that both CAM and the epiphytic habit arose more than once in the family's evolutionary history. In the second part we consider the photosynthetic ecology of the various bromeliad life-forms in more detail using the specific example of Trinidad (West Indies). CAM bromeliads tend to be centred on the drier regions of the island and C₃ forms on the wetter areas. However, at any one site there is a marked vertical stratification of species within the forest profile. Based on the known habitat preferences of the bromeliads, six contrasting sites were selected for field studies in Trinidad. These ranged from arid coastal scrub to montane rain forest, the vegetational and climatic characteristics of which are described here. The constancy of δ¹³C values (carbon-isotope ratios) for individual CAM species in these markedly different habitats emphasized the need for ecophysiological studies to characterize environmental effects on CO₂ assimilation and transpiration. The following papers in this series present the results of a comparative investigation of gas exchange and leaf water relations of CAM and C₃ bromeliads in situ at the various sites.     

The systematic implications of the complexity of leaf flavonoids in the bromeliaceae

In a leaf survey of 61 species of the Bromeliaceae, an unexpectedly wide spectrum of flavonoid constituents was encountered. The family is unique amongst the monocotyledons in the frequency and variety of flavonoids with extra hydroxylation or methoxylation at the 6-position. More common flavonols (in 43% of species) and flavones (in 13%) are distributed throughout the family whereas the rarer flavonoid classes are restricted to one or two of the three subfamilies. Thus 6-hydroxyflavones were found in both the Pitcairnioideae (in 50%) and the Tillandsioideae (in 14%) but patuletin (in 19%), gossypetin (in 1 species) and methylated 6-hydroxymyricetin derivatives (in 24%) were detected only in the Tillandsioideae. A new flavonol, 6,3′,5′-trimethoxy-3,5,7-4′-tetrahydroxyflavone, was identified as the 3-glucoside in Tillandsia usneoides and a new glycoside, patuletin 3-rhamnoside, in Vriesea regina. Myricetin glycosides were found only in the Bromelioideae and their presence here and the concomitant absence of 6-hydroxyflavonoids could indicate the primitive condition of this subfamily. The flavonoid results, in toto, confirm the view based on morphology, that the Bromeliaceae occupies an isolated position in relation to other monocot families.     

Diversity of dry forest epiphytes along a gradient of human disturbance in the tropical Andes

Question: Disturbance effects on dry forest epiphytes are poorly known. How are epiphytic assemblages affected by different degrees of human disturbance, and what are the driving forces? Location: An inter‐Andean dry forest landscape at 2300 m elevation in northern Ecuador. Methods: We sampled epiphytic bryophytes and vascular plants on 100 trees of Acacia macracantha in five habitats: closed‐canopy mixed and pure acacia forest (old secondary), forest edge, young semi‐closed secondary woodland, and isolated trees in grassland. Results: Total species richness in forest edge habitats and on isolated trees was significantly lower than in closed forest types. Species density of vascular epiphytes (species per tree) did not differ significantly between habitat types. Species density of bryophytes, in contrast, was significantly lower in edge habitat and on isolated trees than in closed forest. Forest edge showed greater impoverishment than semi‐closed woodland and similar floristic affinity to isolated trees and to closed forest types. Assemblages were significantly nested; habitat types with major disturbance held only subsets of the closed forest assemblages, indicating a gradual reduction in niche availability. Distance to forest had no effect on species density of epiphytes on isolated trees, but species density was closely correlated with crown closure, a measure of canopy integrity. Main conclusions: Microclimatic changes but not dispersal constraints were key determinants of epiphyte assemblages following disturbance. Epiphytic cryptogams are sensitive indicators of microclimate and human disturbance in montane dry forests. The substantial impoverishment of edge habitat underlines the need for fragmentation studies on epiphytes elsewhere in the Tropics.     

THE FOLIAR EPIDERMIS IN TILLANDSIOIDEAE (BROMELIACEAE) AND ITS ROLE IN HABITAT SELECTION

Structural features of the foliar epidermal surfaces of selected epiphytic members of subfamily Tillandsioideae of Bromeliaceae were examined to determine why xeromorphic nonimpounding (atmospheric) species with a dense layer of foliar trichomes are seldom found in deeply shaded humid portions of the forest profile, whereas tank-forming members of the same subfamily often abound in such sites. Two features of the peltate trichome which appear to play a major role in enforcing habitat preferences among members of Tillandsioideae are (1) anatomical features of the walls of certain trichome cells which affect the mobility of the shield and. (2) variations in juxtaposition of trichomes and stomates on the leaf surface. Both features are involved in inhibition of gas exchange in wet leaves; trichome shield flexibility is also associated with high leaf reflectivity when the leaf surface is dry.     

Tree dominance and diversity in Minas Gerais,Brazil

Quantifying diversity is an old challenge for ecologists and is also a social demand given the increasing threats to natural areas. We sought to work on these issues by using data from 158 vegetation remnants (over 350,000 trees) in southeastern Brazil (Minas Gerais State—MG, nearly 600,000 km²). Specifically, we sought to answer the following questions. (1) How many trees and tree species currently exist in MG vegetation remnants? (2) How much of such biodiversity is present in each vegetation domain (Atlantic Forest, Cerrado and Caatinga) and vegetation type (Seasonally Dry tropical forest “semideciduous”, Seasonally Dry tropical forest “deciduous”, Rain Forest, Swamp, Cerrado and Cerradão) in MG? (3) How much has been lost in regards to tree amount and tree species? We built a 0.1-degree cell grid to estimate the number of trees via spatial regression and used Fisher’s alpha and Fisher’s log series to provide estimates on how many tree species there are and were in MG. We found the number of trees in Minas Gerais to be approximately 24.5 × 10⁹, and the number of tree species to be between 3592 and 3743. The most abundant species distribution among the vegetation domains and vegetation types followed inversely the environmental heterogeneity of the classes. Consequently, the most abundant species in MG belonged to the Cerrado domain, where there was less environmental heterogeneity. The numbers of trees and tree species lost were estimated at 68.54 and 4.03–8.42% of the original values, respectively. We discuss that due to the consequences of human impacts that go back in the region for over a thousand years, other processes not considered in this study, such as habitat degradation by isolation, alterations of food webs, unsustainable use, and climate change, might have caused local extinctions and potentially increased the number of species lost. We believe our results may guide conservation initiatives by providing a base for future environmental laws, parks planning, and the development of more appropriate vegetation management techniques in MG. In addition, our results may inspire future quantitative ecological studies in the tropics.