<Emphasis Type="Italic">Hypericum</Emphasis> species in the Páramos of Central and South America: a special focus upon <Emphasis Type="Italic">H. irazuense</Emphasis> Kuntze ex N. Robson

Knowledge about members of the flowering plant family Clusiaceae occurring in the tropical mountain regions of the world is limited, in part due to endemism and restricted distributions. High altitude vegetation habitats (Páramos) in Central and South America are home to numerous native Hypericum species. Information related to the phytochemistry of páramo Hypericum, as well as ecological factors with the potential to influence chemical defenses in these plants, is briefly reviewed. Results of the phytochemical analysis of Hypericum irazuense, a species collected in the páramo of the Cordillera de Talamanca in Costa Rica, are presented. Lastly, guidelines for the viable and sustainable collections of plant material, to facilitate future investigations of these interesting plants, are given.     

Cryptogamic epiphytes in primary and recovering upper montane oak forests of Costa Rica – species richness,community composition and ecology

Species richness, community composition and ecology of cryptogamic epiphytes (bryophytes, macrolichens) were studied in upper montane primary, early secondary and late secondary oak forests of the Cordillera de Talamanca, Costa Rica. Canopy trees of Quercus copeyensis were sampled with the aim of getting insight in patterns and processes of epiphyte succession and recovery of diversity in secondary forest following forest clearing. Species richness of cryptogamic epiphytes in secondary and primary forests were nearly the same, showing that primary forests are not necessarily more diverse than secondary forests. High species richness of secondary forests was presumed due to the closed canopy, resulting in permanently high atmospheric humidity in these forests. Similarity in species composition of secondary and primary forests increases with forest age, but after 40 years of succession one third (46 species) of primary forest species had not re-established in the secondary forest. Community composition in primary and secondary forests differed markedly and indicates that a long time is needed for the re-establishment of microhabitats and re-invasion of species and communities adapted to differentiated niches. Genera and species exclusive to primary forests are relevant as indicator taxa and conservation targets. Forty percent (68 species) of all species recorded are restricted to secondary forests, indicating the important contribution of secondary forest diversity to total species richness of the oak forests of Costa Rica.     

Tropical and Temperate: Evolutionary History of Páramo Flora

Biogeography of the tropical alpine flora of South and Central America, the páramo flora, has been studied by dividing genera into tropical, temperate, and cosmopolitan chorological flora elements. Published molecular phylogenies of páramo genera are reviewed to summarize knowledge about evolutionary history of the páramo flora and to assess congruence between chorological and phylogenetic approaches. Molecular phylogenies suggest that both the tropical and temperate regions have been important source areas for evolution of the páramo flora. Conclusions derived from chorological patterns regarding origin of genera in páramo are mostly supported by phylogenetic data. Nevertheless, in Chuquiraga, Halenia, Huperzia, and Perezia the chorological scenario is rejected, and in Chusquea-Neurolepis, Elaphoglossum, Gunnera, Halenia, Jamesonia-Eriosorus, and Lasiocephalus independent colonization events from one or several source areas are suggested. Tropical and temperate genera contributed equally to modern species richness of the páramo flora. Among temperate genera, the northern hemisphere genera gave rise to more species in páramo than did genera from the southern hemisphere. So far, no unequivocal evidence has been provided for migration of páramo genera to the temperate zones.     

Altitudinal distribution,diversity and endemicity of Carabidae (Coleoptera) in the páramos of Ecuadorian Andes

Species richness and diversity of Carabidae (Coleoptera), as well as rates of endemicity, are studied along altitudinal transects in the páramo of Ecuadorian Andes, from 3500 to 5000 m. Whereas a global tendency to reduction of species richness is evident from 4200 m upwards, two zones of high diversity and high proportion of endemic species occur at 3800–4000 m and at 4200–4400 m. Species turnover between grass páramo and superpáramo is significantly higher in drier mountains, especially in the Western Cordillera, than in humid mountains of the Eastern Cordillera. The altitudinal range of Carabid species tends globally to decrease along the vertical gradient, but with important local variations due to microenvironmental factors, especially humidity rate. When compared with recent phytogeographical studies, these results tend to support the idea that the majority of tussockgrass páramo is a secondary anthropogenic ecosystem. On the contrary, it is argued that the xeric landscape of the Chimborazo “arenal” is primordial, based on the presence of a stenotopic and possibly relict species, Pelmatellus andium Bates 1891.     

Polystichum lilianae sp. nov. (Dryopteridaceae) and its relationships to P. fournieri and P. turrialbae

Morphological, isozyme, and cytological analyses of the small, pale-scaled polystichums from oakdominated montane rain forests in Costa Rica and Mexico reveal the presence of a separable undescribed species endemic to the Cordillera de Talamanca of Costa Rica. The new taxon,Polystichum lilianae, is an allotetraploid hypothesized to have the sympatricP. turrialbae as one diploid progenitor based on isozyme characters. The isozyme and morphometric data also support the inclusion ofP. smithii, described from southern Mexico, inP. turrialbae, described from Costa Rica. The namePolystichum fournieri, formerly used for all of these plants, applies to species endemic to Oaxaca and Chiapas, Mexico. It is not a progenitor ofP. lilianae.     

Bird species diversity in forest understory: Analysis of mist-net samples

Summary Mist-netted samples of forest understory birds in neotropical (Costa Rica) and temperate (Illinois) forests (during spring migration) supported certain earlier observations: the neotropical understory sample contained more species, more rare species, and more species that use resources not sufficiently available in temperate forest.During spring migration in Illinois, the number of bird species in a mist-netted sample of the community approaches (albeit temporarily) that of neotropical forests and is sometimes even equivalent. But the constitution of these samples seems to be quite different in different regions. Costa Rican samples showed greater within-habitat variability and individual recapture distances were shorter, implying a greater local patchiness of bird distributions than in Illinois in spring.     

Growth form evolution and hybridization in Senecio (Asteraceae) from the high equatorial Andes

Changes in growth forms frequently accompany plant adaptive radiations, including páramo–a high‐elevation treeless habitat type of the northern Andes. We tested whether diverse group of Senecio inhabiting montane forests and páramo represented such growth form changes. We also investigated the role of Andean geography and environment in structuring genetic variation of this group. We sampled 108 populations and 28 species of Senecio (focusing on species from former genera Lasiocephalus and Culcitium) and analyzed their genetic relationships and patterns of intraspecific variation using DNA fingerprinting (AFLPs) and nuclear DNA sequences (ITS). We partitioned genetic variation into environmental and geographical components. ITS‐based phylogeny supported monophyly of a Lasiocephalus‐Culcitium clade. A grade of herbaceous alpine Senecio species subtended the Lasiocephalus‐Culcitium clade suggesting a change from the herbaceous to the woody growth form. Both ITS sequences and the AFLPs separated a group composed of the majority of páramo subshrubs from other group(s) comprising both forest and páramo species of various growth forms. These morphologically variable group(s) further split into clades encompassing both the páramo subshrubs and forest lianas, indicating independent switches among the growth forms and habitats. The finest AFLP genetic structure corresponded to morphologically delimited species except in two independent cases in which patterns of genetic variation instead reflected geography. Several morphologically variable species were genetically admixed, which suggests possible hybrid origins. Latitude and longitude accounted for 5%–8% of genetic variation in each of three AFLP groups, while the proportion of variation attributed to environment varied between 8% and 31% among them. A change from the herbaceous to the woody growth form is suggested for species of high‐elevation Andean Senecio. Independent switches between habitats and growth forms likely occurred within the group. Hybridization likely played an important role in species diversification.     

A new species of Columnea and range extension in the Gesneriaceae from Costa Rica

Columnea lariensis is described and illustrated as a new species from the Talamanca Mountains, Costa Rica. It is similar to C. canarina from Panama. Alloplectus weirii is here reported for the first time from Mesoamerica and an illustration is provided.

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Resumen   Columnea lariensis de la Cordillera de Talamanca, Costa Rica, es descrita e ilustrada. Es similar a C. canarina de Panamá. Alloplectus weirii se reporta para Mesoamérica por primera vez y se incluye una ilustración de la misma.

     

High litterfall in old-growth and secondary upper montane forest of Costa Rica

Tropical upper montane forests usually comprise trees of small stature with a relatively low aboveground productivity. In contrast to this rule, in the Cordillera de Talamanca (Costa Rica), tall trees (>35 m in height and more than 60 cm in diameter) are characteristic for the upper montane old-growth oak forests which are growing at an altitude of almost 3,000 m close to the alpine timberline. For these exceptional forests, productivity data are not yet available. In this study, we analyzed litterfall and its components (tree leaves, litter of epiphytic vascular and non-vascular plants, mistletoes, twigs and other canopy debris) in three forest stands belonging to different successional stages and related seasonal changes in litterfall to micrometeorological variables. The studied stands were early-successional forest (10–15-year-old), mid-successional forest (40-year-old), and old-growth forest. The stands are dominated by Quercus copeyensis and are located at 2,900-m altitude. Total litterfall was highest in the mid-successional forest (1,720 g m⁻² y⁻¹), and reached 1,288 g m⁻² y⁻¹ in the old-growth forest and 934 g m⁻² y⁻¹ in the early-successional forest. Litter mass was dominated by leaves in all stages (56–84% of total litterfall). In the old-growth forest, however, twigs and small canopy debris particles (33%), epiphytes (6%), and mistletoes (5%) also contributed substantially to litter mass. Leaf litterfall showed a clear seasonal pattern with a negative correlation to monthly precipitation and highest values in the dry season (January–April). However, the strongest correlation existed with minimum air temperature (negative), probably because temperatures already dropped at the end of the rainy season, when precipitation had not yet declined and leaf shedding already increased. In contrast, litterfall of epiphyte mass, and twigs and other debris was mostly dependent on occasional strong winds. We conclude that the upper montane oak forests of the Cordillera de Talamanca are exceptional with respect to the large tree size and the relatively high productivity as indicated by litterfall. Litter mass was especially high in the mid-successional and old-growth forests, where the observed annual totals are among the highest recorded for tropical forests so far.     

Ecophysiology in Relation to Exposure of Pendant Epiphytic Bryophytes in the Canopy of a Tropical Montane Oak Forest1

We investigated ecophysiological and morphological traits affecting light and water use of four commercially important pendant epiphytic bryophyte species in a montane oak‐bamboo forest in Costa Rica. Predictions based mostly on ecophysiological studies of temperate bryophytes and vascular plants were experimentally tested on tropical montane bryophyte species ranked on the basis of their distributions in the canopy from the most protected (1) to the most exposed sites (4): (1) Phyllogonium viscosum; (2) Pilotrichella flexilis; (3) Dendropogonella rufescens; and (4) Frullania convoluta. As predicted, highly exposed species tended to have higher light saturation and compensation points, higher dark respiration rates, more chlorophyll, higher chlorophyll a:b ratios, and higher nitrogen concentrations. Contrary to predicted trends, exposed‐site species had lower water contents at full saturation; there were no detectable differences among species in the rate of water loss. Rates of carbon gain in all species reached asymptotes with increasing water content, but the moisture compensation point for carbon uptake of the moss D. rufescens was substantially higher than in the other species. Observed patterns result from interactions among processes related to water storage and transport operating at different scales and are determined by various morphological traits including the density, size, and disposition of phylloids, as well as by whole‐clump architecture.     

WOOD SPECIFIC GRAVITY GRADIENTS IN TROPICAL DRY AND MONTANE RAIN FOREST TREES

Pith to bark specific gravity (SG) trends were investigated in 18 tropical dry forest and six montane rain forest tree species of Costa Rica. Eleven dry forest species showed statistically significant increases in SG with distance from pith. The increases ranged from 20–80%; the greatest changes were exhibited by species which are known to occur in tropical wet as well as tropical dry forests. The other seven species showed no change in SG with distance from pith. Of the montane forest species, one showed a significant decrease of 20%, and three showed significant increases ranging from 20–40%. Two species exhibited no change in SG. Comparison of these changes with trends found in tropical wet forest and temperate forest suggests that the increase in SG with size is most common in tropical wet forest, least common in temperate forest, and intermediate in tropical dry and montane forests.     

Angiosperm flora and biogeography of the páramo region of Colombia,Northern Andes

Páramo is the neotropical high elevation ecosystem in the northern Andes and Central America consisting of multiple dissected open areas above 3000 m a.s.l. Complex evolutionary processes that occurred within these ecosystems gave rise to a unique tropical Andean flora. Previous phytogeographical classifications for Colombian páramos have been based on subjective assessments of species distributions. However, a detailed floristic analysis highlighting affinities between páramo regions in Colombia has not yet been proposed. The aim of this study is to provide an analytical scenario for the patterns of regional plant diversity in a hierarchical framework based on the biogeographical history of the páramos, based on 30 localities of the Colombian páramos. Parsimony analysis of endemicity (PAE) and Jaccard similarity indices were applied to a presence/absence data matrix of páramo angiosperm species. Altitudinal distribution of species was used to determine the percentage of the páramo flora endemic to this ecosystem. Genera are shared among páramos, whereas species composition is highly endemic. 86% of the páramo angiosperm plant species is endemic to this ecosystem. The Colombian Páramo areas can be grouped using PAE into five biogeographical units, based on angiosperm species composition. These are: (1) Páramos de la Cordillera Oriental; (2) Páramos de la Cordillera Central and Macizo Colombiano; (3) Páramos de Antioquia; (4) Páramos del Norte; (5) Páramos de la Cordillera Occidental.     

Tylopilus alkalixanthus, a new species of Boletaceae from Costa Rica and Japan

Tylopilus alkalixanthus is described from the Cordillera Talamanca of Costa Rica. At the present time it is known only from Costa Rica and Japan.     

Size and Structure of Fine Root Systems in Old-growth and Secondary Tropical Montane Forests (Costa Rica)

The fine root systems of three tropical montane forests differing in age and history were investigated in the Cordillera Talamanca, Costa Rica. We analyzed abundance, vertical distribution, and morphology of fine roots in an early successional forest (10–15 years old, ESF), a mid‐successional forest (40 years old, MSP), and a nearby undisturbed old‐growth forest (OGF), and related the root data to soil morphological and chemical parameters. The OGF stand contained a 19 cm deep organic layer on the forest floor (i.e., 530 mol C/m²), which was two and five times thicker than that of the MSF (10 cm) and ESF stands (4 cm), respectively. There was a corresponding decrease in fine root biomass in this horizon from 1128 g dry matter/m² in the old‐growth forest to 337 (MSF) and 31 g/m² (ESF) in the secondary forests, although the stands had similar leaf areas. The organic layer was a preferred substrate for fine root growth in the old‐growth forest as indicated by more than four times higher fine root densities (root mass per soil volume) than in the mineral topsoil (0–10 cm); in the two secondary forests, root densities in the organic layer were equal to or lower than in the mineral soil. Specific fine root surface areas and specific root tip abundance (tips per unit root dry mass) were significantly greater in the roots of the ESF than the MSF and OGF stands. Most roots of the ESF trees (8 abundant species) were infected by VA mycorrhizal fungi; ectomycorrhizal species (Quercus copeyemis and Q. costaricensis) were dominant in the MSF and OGF stands. Replacement of tropical montane oak forest by secondary forest in Costa Rica has resulted in (1) a large reduction of tree fine root biomass; (2) a substantial decrease in depth of the organic layer (and thus in preferred rooting space); and (3) a great loss of soil carbon and nutrients. Whether old–growth Quercus forests maintain a very high fine root biomass because their ectomycorrhizal rootlets are less effective in nutrient absorption than those of VA mycorrhizal secondary forests, or if their nutrient demand is much higher than that of secondary forests (despite a similar leaf area and leaf mass production), remains unclear.     

The effect of conspecific density,herbivory, and bamboo on seedling dynamics of a dominant oak in a Neotropical highland forest

Conspecific negative density dependence (CNDD) is one of the main mechanisms influencing diversity maintenance in tropical forests. Tropical highland forests, in contrast to most lowland forests, are commonly dominated by a few tree species, and testing the importance of density dependence effects on seedling establishment of dominant trees may provide insights on the mechanisms regulating population dynamics and forest composition of tropical highlands. We tested the effect of CNDD regulation on seedling survival and recruitment of Quercus costaricensis, a monodominant oak in the Talamanca highland forests of Costa Rica. We used Ripley's K and generalized linear mixed models to test the effects of conspecific density, distance to the nearest adult, density of Chusquea bamboo shoots, and herbivory on the annual survival probability of 3579 seedlings between 2014 and 2017. We did not find a significant effect of CNDD on seedling survival. However, bamboo density and herbivory both significantly decreased oak seedling survival. All seedlings had signs of herbivory and predator satiation may explain the lack of density dependent regulation in seedlings of this species. We argue that the lack of intraspecific density regulation at the seedling stage may contribute to explain the dominance of Q. costaricensis in the highland forests of Costa Rica. Local seedling dynamics of this endemic oak are instead regulated by herbivory and the density of Chusquea. Abstract in Spanish is available with online material.     

Low temperature resistance in saplings and ramets of Polylepis sericea in the Venezuelan Andes

The frequent occurrence of all year-round below zero temperatures in tropical high mountains constitutes a most stressful climatic factor that plants have to confront. Polylepis forests are found well above the continuous forest line and are distributed throughout the Andean range. These trees require particular traits to overcome functional limitations imposed on them at such altitudes. Considering seedling and sapling stages as filter phases in stressful environments, some functional aspects of the regeneration of Polylepis sericea, a species associated to rock outcrops in the Venezuelan Andes, were studied. We characterized microclimatic conditions within a forest, in a forest gap and surrounding open páramo and determined low temperature resistance mechanisms in seedlings, saplings and ramets. Conditions in the forest understory were more stable compared to the forest gaps and open surrounding páramo. Minimum temperatures close to the ground were 3.6 °C lower in the open páramo compared to the forest understory. Maximum temperatures were 9.0 °C higher in the open páramo. Ice nucleation and injury temperatures occurred between ?6 and ?8 °C for both ramets and saplings, an evidence of frost avoidance to low nighttime temperatures. In this particular forest, this resistance ability is determinant in their island-like distribution in very specific less severe temperature habitats.     

Vegetation Zonation in a Neotropical Montane Forest: Environment,Disturbance and Ecotones

Gradual changes in vegetation structure and composition are expected to result from continuous environmental change with increasing elevation on mountains. Hence, the occurrence of abrupt or discrete ecotones in vegetation patterns is intriguing and may suggest key controls on community assembly in montane forests. We review tropical montane forest (TMF) zonation patterns focusing on a case study from the Cordillera Central, Hispaniola where a striking discontinuity in forest composition occurs consistently at ～2000 m elevation, with cloud forest below and monodominant pine forest above. We propose that a discontinuity in climatic factors (temperature, humidity) associated with the trade‐wind inversion (TWI) is the primary cause of this and other ecotones in TMFs that occur at a generally consistent elevation. Low humidity, fires and occasional frost above the TWI favor pine over cloud forest species. Fires in the high‐elevation pine forest have repeatedly burned down to the ecotone boundary and extinguished in the cloud forest owing to its low flammability, reinforced by high humidity, cloud immersion and epiphytic bryophyte cover. Small‐scale fire patterns along the ecotone are influenced by topography and where forest structure is impacted by hurricanes and landslides. Analogous patterns are observed worldwide in other TMFs where the TWI is important, high‐elevation fires are frequent, and the flora contains frost‐tolerant species (often of temperate lineage). The response of this and other TMFs to anthropogenic climate change is highly uncertain owing to potentially countervailing effects of different climatic phenomena, including warming temperatures and decreased frost; changes in the TWI, high‐elevation drought or cloudiness; and increased frequency or intensity of hurricanes and El Niño‐Southern Oscillation events.     

Spatial relations and population structure of a dominant tree along a treeline ecotone in the Tropical Andes: interactions at gradient and plant-neighbourhood scales

Background: Studies in temperate mountains suggest that plant–plant interactions modulate tree establishment above the forest line. In tropical mountains worldwide this issue remains largely unexplored.

Aims: To analyse the population structure and local spatial relationships of a dominant tree at a species-rich tropical Andean forest line.

Methods: We determined changes in the population structure of Diplostephium venezuelense along an elevation gradient between continuous forest and open páramo and analysed plant community structure and superficial rock cover in the neighbourhood of saplings and adults at the upper forest line.

Results: Sapling and adult densities were highest in low-altitude páramos adjacent to the forest line and at the borders of small forest islands. Saplings showed local spatial aggregation, were positively associated with small boulders and low shrubs, and negatively associated with mosses and lichens. However, no spatial association was found between scattered adults in the páramo and saplings of other forest trees.

Conclusions: Complex species-specific local spatial interactions, suggesting both facilitative and antagonistic effects, seem to modulate the establishment of the dominant tree D. venezuelense at and above the upper forest line. Nevertheless, the establishment of other tree species above continuous forests does not appear to be facilitated by the canopy cover offered by the isolated D. venezuelense individuals established in open páramo environments.     

A new Costa Rican species of centropogon (Campanulaceae-Lobelioideae)

A newCentropogon (sect.Wimmeriopsis subsect.Colombiani) from the Cordillera de Talamanca, Costa Rica, is described and compared toC. gutierrezii (Planch. & Oerst.) F. E. Wimmer, a species restricted to the Cordillera Central with which it previously has been confused.     

Notes on the bryophyte communities oe lowland Tropical Rain Forest,with special reference to Moraballi Creek,British Guiana

Discussion and conclusions The four main bryophyte synusiae in the Moraballi Creek rain forest (high epiphytes, shade epiphytes, epiphyllae, synusia on dead wood) are remarkably distinct both in structure and floristic composition, though there are a few very frequent species, e.g. Calymperes lonchophyllum, Octoblepharum albidum, which occur in all the synusiae except the epiphyllous, which consists almost entirely of highly specialised species exclusive to itself. The differences in the structure of the synusiae depend on the different growth forms of the more abundant species. Data are not available for a satisfactory analysis of these growth forms (like that, for instance of Gimingham & Robertson, 1950, for some British bryophyte communities), but from the general observations given above it is obvious, for instance, that there are wide differences between the growth forms of the strongly xeromorphic high epiphyte synusia and those of the shade epiphyte synusia, characterised by markedly hygromorphic features such as freely projecting or dangling shoots and large thin-walled cells. Again, there are conspicuous differences between the growth forms prevailing in the shade epiphyte and dead wood communities. The differences in floristic composition and structure between the various bryophyte synusiae reflect the marked differences in microclimatic and other ecological factors existing between the various microhabitats available for bryophytes within the rain forest community.When the bryophyte synusiae of the Moraballi Creek rain forest are compared with those of temperate deciduous or coniferous forests the most striking differences seen are the absence in the former of any bryophyte synusia on the ground, the presence of an epiphyllous community and the pronounced dominance of hepaticae, especially of the family Lejeuneaceae, in all microhabitats except the dead trunks and stumps. In conclusion it may be noted that the bryophyte flora of the lowland rain forest is not excessively rich in species, indeed area for area it is probably poorer than many temperate forests. The comparatively restricted bryophyte flora, compared with the overwhelmingly rich angiosperm flora is a surprising fact. These comparisons, though based in the first place on the Moraballi Creek forest alone, would apply to lowland rain forests in Africa or the Indo-Malayan region. In montane forests conditions differ greatly from those in the lowlands and some of the above conclusions no longer apply.Received for publication 11.XI.1953.