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A. Palm dominated dry forest (Átures area). (At-r = Attalea racemosa Spruce; Bu-p = Buchenavia parvifolia Ducke; Co-s = Costus spiralis (Jacq.) Roscoe; Me-m = Melocactus mazelianus; Mi-m = Mimosa microcephala; Pi-a = Pitcairnia armata Maury; Pi-p = Pitcairnia pruinosa; Sy-o = Syagrus orinocensis (Spruce) Burret; Za-l = Zamia lecointei Ducke. B. Forest margin on granitic outcrop (Sipapo area). Ac-p = Acanthella pulchra Gleason; De-o = Decagonocarpus oppositifolius Spruce ex Engl.; Ko-x = Koellensteinia sp.; Ku-r = Kunhardtia radiata Maguire & Steyerm.; Ma-g = Macrolobium gracile Spruce ex Benth.; Na-b = Brewcaria brocchinioides (L.B. Sm.) B. Holst; Pa-l = Parodiolyra luetzelburgii (Pilger) Soderstr. & Zuloaga; Sc-c = Scleria cyperina Kunth; Sp-x = Sphagnum sp.; Ut-s = Utricularia subulata L.; Ut-x = Utricularia sp).
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The Guayana Shield, located in north-eastern South America, consists of a highly complex and composite mosaic of landscape elements. Amongst these, inselbergs are very conspicuous, because of their peculiar shape and their unusual associated vegetation. Geologically, these rock outcrops are part of the underlying ancient igneous-metamorphic basemen...
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Citations
... These plants often display stress-tolerant ecological strategies (Negreiros et al., 2014;de Paula et al., 2015;Dayrell et al., 2018), clonal growth, cluster roots , desiccation tolerance, succulence (Biedinger et al., 2000;), as well as leaf traits related to water storage, drought, and frost resistance (Camerik & Werger, 1981;Vitarelli et al., 2016;de Paula et al., 2019). In addition to these attributes, plant communities on rock outcrops show high alpha and beta diversity, high levels of endemism (Berry & Riina, 2005;Gröger & Huber, 2007;Scarano, 2007;Hoorn et al., 2018), and species with reduced dispersal and low genetic flow (Barbará et al., 2007). Despite their similar physical appearance, rock formations have heterogeneous species composition, and their ecosystems are named differently around the globe, such as fynbos in South Africa (Rebelo et al., 2006), kwongan in Australia (Tsakalos et al., 2019), campos rupestres in Brazil , and tepuis in the Amazon (Prance, 1996). ...
The geodiversity of rocky ecosystems includes diverse plant communities with specific names, but their continental‐scale floristic identity and the knowledge on the role of macroclimate remain patchy. Here, we assessed the identity of plant communities in eastern Brazil across multiple types of rocky landscapes and evaluated the relative importance of climatic variables in constraining floristic differentiation. We provided lists of diagnostic species and an assessment of the conservation status of the identified floristic groups. We compiled a data set of 151 sites (4498 species) from rocky ecosystems, including campos rupestres, campos de altitude , granitic‐gneiss lowland inselbergs, and limestone outcrops. We used unsupervised clustering analysis followed by ANOSIM to assess floristic groups among sites. We performed a random forest variable selection to test whether the identified floristic groups occupy distinct climatic spaces. Six groups (lithobiomes) segregated floristically according to lithology and climate. Alongside campos de altitude and limestone outcrops, inselbergs were divided according to the biome in which they occur (Atlantic Forest or Caatinga), and campos rupestres were largely segregated according to their lithological matrix (ironstone or quartzitic). Plant communities of Caatinga inselbergs were more similar to limestone outcrops, while Atlantic Forest inselbergs communities resembled campos de altitude . The composition of plant communities on outcrops seems to be largely constrained by lithology, but climatic factors are also meaningful for sites with similar lithology. The current network of protected areas does not cover these unique ecosystems and their floristic heterogeneity, with Caatinga inselbergs and limestone outcrops being the least protected.
... Inselbergs are good model systems to study the strategies of plants living in extreme ecosystems because their high floristic diversity and the extraordinary number of endemic species rule out dispersal limitation as an important community assembly mechanism at local scales (Parmentier et al., 2005;Parmentier and Hardy, 2009;Sarthou et al., 2017;Yates et al., 2019). Inselbergs (from German Insel = island and Berg = mountain) are pre-Cambrian black monolithic rock outcrops that form terrestrial dome-like 'islands' that rise above the surrounding forest or savanna landscape ( Fig. 1; Porembski and Barthlott, 2000a, b;Gröger and Huber, 2007;Porembski, 2007;Giraldo-Cañas, 2008;Lüttge, 2008). The distinct vegetation community of tropical inselbergs is subjected to multiple environmental stressors: high and highly variable air temperatures (23-40 °C, with record low temperatures of 18 °C), variable relative humidity (20-100 %), intermittent water availability, and shallow acid soils with low nutrient content (Porembski and Barthlott, 2000a;Gröger and Huber, 2007;Giraldo-Cañas, 2008;Lüttge, 2008;Sarthou et al., 2017). ...
... Inselbergs (from German Insel = island and Berg = mountain) are pre-Cambrian black monolithic rock outcrops that form terrestrial dome-like 'islands' that rise above the surrounding forest or savanna landscape ( Fig. 1; Porembski and Barthlott, 2000a, b;Gröger and Huber, 2007;Porembski, 2007;Giraldo-Cañas, 2008;Lüttge, 2008). The distinct vegetation community of tropical inselbergs is subjected to multiple environmental stressors: high and highly variable air temperatures (23-40 °C, with record low temperatures of 18 °C), variable relative humidity (20-100 %), intermittent water availability, and shallow acid soils with low nutrient content (Porembski and Barthlott, 2000a;Gröger and Huber, 2007;Giraldo-Cañas, 2008;Lüttge, 2008;Sarthou et al., 2017). ...
Background and aims:
Trait-based frameworks assess plant survival strategies using different approaches. Some frameworks use functional traits to assign species to a priori defined ecological strategies. Others use functional traits as the central element of a species ecophysiological strategy. We compared these two approaches by asking 1) what is the primary ecological strategy of three dominant co-occurring shrub species from inselbergs based on the CSR scheme? 2) What main functional traits characterize the species ecophysiological strategy based on their carbon, water, and light use?
Methods:
We conducted our study on a Colombian inselberg. In this extreme environment with multiple stressors (high temperatures and low resource availability), we expected all species to be stress-tolerant (S in the CSR scheme) and have similar ecophysiological strategies. We measured 22 anatomical, morphological, and physiological leaf traits.
Key results:
The three species have convergent ecological strategies as measured by CSR (S: Acanthella sprucei and S/CS: Mandevilla lancifolia and Tabebuia orinocensis) yet divergent resource-use strategies as measured by their functional traits. A. sprucei has the most conservative carbon use, risky water use, and a shade-tolerant strategy. M. lancifolia has acquisitive carbon use, safe water use, and a shade-tolerant strategy. T. orinocensis has an intermediate carbon use, safe water use, and a light-demanding strategy. Additionally, easy-to-measure stomatal traits are valuable to describe resource-use strategies because they correlate highly with two hard-to-measure physiological functions: stomatal conductance and maximum photosynthesis per unit mass.
Conclusions:
The two approaches provide complementary information on species strategies. Plant species can co-occur in extreme environments like inselbergs because they exhibit convergent primary ecological strategies but divergent ecophysiological strategies, allowing them to use limiting resources differently.
... In rock outcrops, because soils are spatially structured, they determine the distribution of vegetation patches and the structure of tree communities (Parmentier 2003;Conceição et al. 2007;Scarano 2007). Droughtprone soils unable to store water for long periods of time tend to contain low-diversity communities with highly specialized species (Gröger and Huber 2007). In limestone outcrops, soil fertility drives the phylogenetic composition of tree community patches (Aguiar- Campos et al. 2020), and in lowland moist tropical forests and Amazonian palm communities, low fertility sites tend to have closely related species (Schreeg et al. 2010;Muscarella et al. 2019). ...
Granite-gneiss rock outcrop inselbergs are ancient stable ecosystems with old, climatically-buffered infertile landscapes (OCBILs). Although inselbergs provide key ecosystem services, little is done for their conservation and, so far, a lot of their unknown evolutionary history has already been lost by human activities. Using a fine-scale approach, here we tested if habitat and environmental filtering (the inselberg’s harshness) affect the evolutionary diversity of an Atlantic Forest inselberg in Brazil. We recorded all trees with a diameter at breast height ≥ 5 cm in 20 plots in four habitat types (total sampled area of 0.8 hectares), from highest to lowest: island, hillside, foothill, and semideciduous forest (matrix). We also collected soil samples for chemical, textural and physical soil characterization. We fitted linear models to test the effects of soil and habitat on plot-level metrics of phylogenetic diversity and structure, lineage diversity, phylogenetic β-diversity, and evolutionary distinctiveness. We found that the upper inselberg habitats contain a distinct set of ancient, closely related, harsh-tolerant lineages, as well as a subset of lineages that persist under harsh conditions with a certain degree of water availability. The inferior inselberg habitats harbor higher lineage diversity than expected by chance. Soil strongly predicted evolutionary diversity. We concluded that soil depth, slope, nutrients and texture (environmental filtering) and habitat types and topography (habitat filtering) shape the evolutionary history contained in fine-scale inselberg habitats, which should encourage the conservation of these ancient ecosystems.
... Several studies have shown how habitat or micro-habitat in rock outcrops determine different flora and vegetation patterns (Porembski et al. 1997;Conceição & Pirani 2005;Gröger & Huber 2007;Jacobi et al. 2007;Porembski 2007;Paula et al. 2017). According to these authors, patterns are related with soil depth, humidity and shading. ...
... & Arn., a species noted in the literature as an epiphyte (Mucunguzi 2007;Alves et al. 2008;Ceballos et al. 2016). In addition, the genus Peperomia was described by Gröger & Huber (2007) as presenting shadetolerant lithophyte species. Therefore, the typical, more xeric conditions of the top habitat, may be attenuated if the surrounding forest shadows the outcrops. ...
Abstract Rock outcrop vegetation is recognized worldwide by its singular and biodiverse flora. Campo Rupestre forms hyperdiverse mosaics in rocky environments across a wide latitudinal and altitudinal gradient, with high species turnover at macro- and micro-scales. The surrounding biomes, climate, and geological formations are the main drivers of species turnover on a macro-scale while micro-habitat seems to be the main one determining the peculiarities of the Campo Rupestre on a micro-scale. In a quartzitic Campo Rupestre area we evaluate how the outcrop micro-habitats influence floristic composition and functional traits. The study area is located in the municipality of Ouro Preto, Minas Gerais state, southeastern Brazil. Two main outcrop habitats were considered: top surfaces, with bare rock, shallow depressions and ephemeral ponds; and lateral surfaces, with clefts and crevices. We recorded the vascular species, their respective life-forms (according to Raunkiaer’s system) as well as their coverage in 18 plots. We identified 71 species in 31 families. The floristic spectra and species composition were similar between top and lateral surfaces. There was no significant difference among the vegetational spectra. However, hemicryptophytes were slightly dominant on top, while on the sides chamaephytes were the dominant life-form. Understanding plant adaptations to these environments provides insights into the mechanisms underlying geomorphological heterogeneity in plant communities.
... Se encuentran rodeando parte de la laguna El Pañuelo hasta el río Orinoco y en el extremo oriental de la Reserva en los cerros Morrocoy, Campana y el Karikari, así como los cerros aledaños como Cerro tigre, Calaballo y Ventanas. La vegetación está determinada por un complejo sistema de gradientes, principalmente por la profundidad del suelo y disponibilidad de agua, que forman un mosaico de hábitats marginales donde un gran número de especies encuentran nichos apropiados para crecer (Gröger y Huber 2007). En el caso de los afloramientos presentes en la Reserva y en los afloramientos aledaños se pueden reconocer los siguientes. ...
... Bosques. Los bosques dentro de los afloramientos rocosos son los definidos por Gröger y Huber (2007) como "bosques bajos a medianos siempreverdes", por Vincelli (1981) como "monte de Syagrus" y como "bosques asociados a cerros rocosos" por Mendoza (2007). En la Reserva se encuentran en grandes formaciones cóncavas o rodeando las laderas de la roca. ...
... Las divisiones de este hábitat se realizan de acuerdo a los estados de sucesión en que se encuentra la vegetación. Gröger y Huber (2007) destacan la presencia de un centro de endemismo (Atures Figura 7. Afinidad florística entre los departamentos que conforman la región de la Orinoquia en Colombia y la Reserva Natural Bojonawi. Centre of Endemism) en la esquina nor-occidental del estado Amazonas y en el adyacente estado Bolívar en Venezuela, el cual está en el área de influencia de la Reserva para el lado colombiano. ...
Descripción de los hábitat de la reserva natural de Bojonawi Vichada, Colombia
... (Viana et al. 2016;Mota et al. 2018;Zappi et al. 2019). There are also island-like lowland ecosystems, such as white sand campinaranas, savannas, and low elevation granitic domes or inselbergs, associated with arenitic and often waterlogged soil in the Amazon region (Gröger and Huber 2007;Adeney et al. 2016;Costa et al. 2019;Henneron et al. 2019;Devecchi et al. 2020). ...
The world's largest mineral iron province, Serra dos Carajás, is home to an open vegetation known as canga, found on top of isolated outcrops rising out of the Amazon rainforest. Over one thousand vascular plants species have been recorded in these canga sites, including 38 edaphic endemics. A new survey adds to our investigation of biogeographic relationships between sixteen canga outcrops and the effect of the distance between site pairs on the number of shared species, regional species turnover and species distribution patterns. Plant collecting expeditions to the westernmost site, the Serra de Campos of São Félix do Xingu (SFX), were carried out followed by the identification of all collected specimens and the creation of a species database, built to perform biogeographical analyses. Floristic relationships among the sites were investigated regarding their similarity, using multivariate analyses. The correlation between canga areas and species richness was tested, as well as the geographical distance between pairs of outcrops and their shared species. Vascular plants at SFX total 254 species including 17 edaphic endemics. All canga sites are grouped with 25% of minimum similarity, and the SFX falls within a large subgroup of outcrops. The total species number shared between site pairs does not change significantly with geographical distance but is positively correlated with the area of each outcrop. Meanwhile, shared endemic species numbers between site pairs decline when geographical distance increases, possibly imposed by the barrier of the rainforest. Our data suggest higher shared similarity between the largest and species-richest sites as opposed to geographically nearby sites, and provide useful insight for drafting conservation and compensation measures for canga locations. The size of the canga outcrops is associated to higher floristic diversity but connectivity among islands also plays a role in their similarity.
... Fine-scale changes in community composition and species richness have been observed in other studies in drylands, where this mosaic of patches is assumed to be the result of close feedback from resource limitations (soil water) and the dynamics of the vegetation (Rietkerk and van de Koppel 2008;Meloni et al. 2017). In some studies, it has been argued that changes in community composition among vegetation profiles of tropical inselbergs are mainly conditioned by habitat structure and water deficit (Lüttge 1997;Gröger and Huber 2007;Porembski 2007). Thus, one of the challenges for future research is to assess the relationship between climate, altitude and local factors, and how these affect diversity patterns on tropical inselbergs. ...
Plant communities are shaped by multiple factors along environmental gradients; however, studies are limited on how environmental filtering drives community composition and species richness on tropical inselbergs. We evaluate the influence of altitude and climatic variables related to temperature and precipitation on plant community composition and species richness on Brazilian inselbergs. We assume as a premise that both climate and altitude would induce changes on plant community composition and species richness at the local level. We used plant inventory data from 370 sampling units across four inselberg sites in the Atlantic Forest of Espírito Santo State, south-eastern Brazil. We tested the univariate and multivariate effects of altitude and climate variables on community composition and species richness with multiple models. Differences in species richness between inselbergs were evaluated using sample-based data to estimate rarefaction and extrapolation curves. In addition, differences in species composition and taxonomic beta diversity were examined via novel frequency-based metrics. A contrasting climate pattern was observed between the inselberg sites, with south sites being wet compared to the dry conditions found in northern sites. Species richness by rarefaction showed a similar pattern within regional sites; however, there were marked differences between regions. Species richness and beta diversity showed significant differences among sites, with higher values in southern sites than in northern sites. In a multi-model comparison between inselberg sites, altitude significantly influenced community composition and species richness and explained more variance than climate models. This finding suggested that climate could act to some extent on these tropical inselbergs; however, altitude was a better predictor of plant community composition and species richness at the local level.
... beta diversity patterns between seasons at each site. In different rocky outcrops, there are plant communities that are well adapted to extreme environmental conditions, mainly to water and nutrient deficits, as well as shallow soils (Lüttge 1997;Gröger & Huber 2007). These species have different mechanisms and adaptive strategies to tolerate scarce substrates and water, as well as the high temperatures due to the bare rock surface (i.e., Lüttge 1997). ...
... These species have different mechanisms and adaptive strategies to tolerate scarce substrates and water, as well as the high temperatures due to the bare rock surface (i.e., Lüttge 1997). Several studies in America and Africa show that the vegetation on rocky outcrops is dominated by succulents, geophytic and xerophytic species, and drought-tolerant species and drought-avoiding species such as therophytes (Lüttge 1997;Gröger & Huber 2007). Our results allow us to confirm the hypothesis that seasonality, based on changes between a dry and a wet season, determines species composition and beta diversity, but not species richness. ...
... Some plants that are not drought-resistant use drought avoidance as a mechanism to allocate more energy to survival during certain periods, to germinate and grow sufficient aerial biomass during a short period of time, and to conclude their entire life cycle before the end of wet season. For example, therophytes that germinate after the beginning of the wet season can develop in a short period of time while the humidity is relatively high (Lüttge 1997;Gröger & Huber 2007;Villa et al. 2018). Likewise, geophytes are another group of plants that exhibit this mechanism. ...
Background and aims – Seasonality exerts strong controlling forces on species diversity in herbaceous species communities, however, this control process remains poorly understood in tropical lithologically different rocky outcrops. We aim to investigate the effect of seasonality and the variability of soil properties on changes in the herbaceous species richness and species composition of two different herbaceous species communities on rocky outcrops in Brazil. We hypothesize that seasonality, determined by variation in precipitation, and soil fertility, determined by variability in nutrient-related soil properties, drives species diversity (i.e., richness and beta diversity) patterns of herbaceous communities at local scale.Methods – To investigate how the variation between dry and wet seasons affects species richness and beta diversity, we studied plots on rocky outcrops of Iron Quadrangle (40 plots, 1 × 1 m) and Carajás (20 plots, 1 × 1 m). Key results – We observed similar richness patterns between seasons, without significant differences between sites, using rarefaction and extrapolation curves. However, we observed significant differences in beta diversity between seasons. Our results indicate that seasonality determines the temporal variation of the herbaceous species composition, but not species richness. Likewise, our tested models indicated that seasonality shape beta diversity in the studied rocky outcrops. Conclusions – The predictable seasonal precipitation is closely related to the community composition on this type of rocky outcrop formation, where there typically is a marked seasonal water deficit pattern, with increased deficit during the dry season. We presume that seasonality is an important driver in determining plant community assembly at local scale on the studied rocky outcrops.
... Some Neotropical mountains are well-known for their high diversity and endemism, including parts of the Andes (Moraes and Beck 1992;Gentry 1995;Gradstein 1995;Rangel 1995), and the Tepuis of the Guyana Shield (Gröger and Huber 2007). Few comparative studies have been undertaken in those mountains habitats, however, and little is known about their ecosystem function and processes (Martinelli 2007). ...
We investigated the species richness, endemism, and similarity of the bryoflora (mosses and liverworts) on five Brazilian Amazon mountains (four Tepuis and one rocky outcrop), to determine the floristic relationships between them using presence/absence matrix. Cluster analysis based on the Jaccard index was used to determine similarities between the plant assemblages in five areas. We also performed a principal component analysis to determine which abiotic variables best explained the variations between those mountains. We surveyed the bryophyte species on each of the five mountains to identify the diversity of mosses and liverworts and to answer to the following questions: (1) How many bryophyte species are there in total area (gama diversity) and on each mountain (alpha diversity)? (2) Do mosses and liverworts share patterns of diversity? (3) Can patterns of alpha diversity be used to predict patterns of beta diversity? (4) What are the species distribution patterns? (5) How many endemic species are there in the mountains? and, (6) Is there high similarity among those bryophyte floras? We encountered 425 species, 144 genera, and 51 families of bryophytes, with Lejeuneaceae and Lepidoziaceae being the richest families. A considerable number of floristic novelties were encountered: 18 species new to Brazil; 39 species new to northern Brazil; 21 species new to Amazonas State; and 2 species new to science and only recently described. Over 43% of the bryophytes have a tropical America distribution, approximately 7% are Pantropical, 6.5% are Afro-American, and 3% are endemic to Brazil (6 liverworts and 6 mosses), and 6% are widespread globally. We present a checklist of 425 bryophyte species (144 genera and 51 families) of which 12 are endemic. Patterns of endemism of the bryophyte flora on the Tepuis differ from those exhibited by angiosperms (in the latter three times higher). Thirty-nine of these species were new for the Amazonia domain in Brazil, increasing by 7% the previous total of Amazonian recorded species. The similarity between sites was low, ranging from 3 to 25%. PCA analysis showed temperature variables explaining the greater part of the variance (76.6%) between the mountains. Our results demonstrate that patterns of alpha diversity cannot be used to predict patterns of beta diversity. This paper represents the first evaluation of bryophytes on five Brazilian Amazon mountains, giving insight into their species richness, endemism, and similarity, and providing baseline information for analyzing species’ turnover rates, migration, invasion events, etc. These results also represent an increase in our general knowledge of the Brazilian bryoflora and are relevant to the conservation of bryophyte diversity in the Amazon forest, and represent a contribution toward the targets of the Global Strategy for Plant Conservation.
... For terrestrial species, the positive size effect was rather mediated by the increase in the diversity of terrestrial habitats present on a given inselberg (H 1b , Fig. 3c-d), in line with various studies on vegetation patches (Auerbach and Shmida 1987, Honnay et al. 1999, Dufour et al. 2006). The effect of habitat diversity was here related to floristic differences between habitats typical of rock-savanna vegetation, such as continuous rock surfaces, temporary ponds, crevices with their variants according to the presence of soil, rock or woody debris (Sarthou and Villiers 1998, Gröger and Huber 2007, Parmentier and Hardy 2009. For epiphytic species, the indirect size effect was mediated by the diversity of shrubs and lianas (Fig. 5b). ...
Disentangling the multiple factors controlling species diversity is a major challenge in ecology. Island biogeography and environmental filtering are two influential theories emphasizing respectively island size and isolation, and the abiotic environment, as key drivers of species richness. However, few attempts have been made to quantify their relative importance and investigate their mechanistic basis. Here, we applied structural equation modelling, a powerful method allowing test of complex hypotheses involving multiple and indirect effects, on an island-like system of 22 French Guianan neotropical inselbergs covered with rock-savanna. We separated the effects of size (rock-savanna area), isolation (density of surrounding inselbergs), environmental filtering (rainfall, altitude) and dispersal filtering (forest-matrix openness) on the species richness of all plants and of various ecological groups (terrestrial versus epiphytic, small-scale versus large-scale dispersal species). We showed that the species richness of all plants and terrestrial species was mainly explained by the size of rock-savanna vegetation patches, with increasing richness associated with higher rock-savanna area, while inselberg isolation and forest-matrix openness had no measurable effect. This size effect was mediated by an increase in terrestrial-habitat diversity, even after accounting for increased sampling effort. The richness of epiphytic species was mainly explained by environmental filtering, with a positive effect of rainfall and altitude, but also by a positive size effect mediated by enhanced woody-plant species richness. Inselberg size and environmental filtering both explained the richness of small-scale and large-scale dispersal species, but these ecological groups responded in opposite directions to altitude and rainfall, that is positively for large-scale and negatively for small-scale dispersal species. Our study revealed both habitat diversity associated with island size and environmental filtering as major drivers of neotropical inselberg plant diversity and showed the importance of plant species growth form and dispersal ability to explain the relative importance of each driver.
