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.     

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.     

The impact of roads on the avifauna of páramo grasslands in Cajas National Park,Ecuador

National parks are an important tool for conserving biodiversity, particularly in areas of high biodiversity and endemism such as the tropical Andes. However, national parks often face a variety of stressors related to recreation, road construction and illegal extraction of natural resources. Unfortunately, the influence of these stressors for biodiversity is rarely well documented. Cajas National Park in Ecuador is no exception. Despite being traversed by the Cuenca-Molleturo-Naranjal road, effects of the road construction on biodiversity have not been determined. We therefore assessed the influence of road proximity on bird species richness and abundance as well as composition of bird habitat groups in Cajas National Park using transect walks at 25 m and 250 m distance to the road (overall 18 transects, each 1 km length). In total, we recorded 1110 individuals of 28 páramo bird species. Overall species richness did not differ between transects near and far from the road. Nevertheless, the average abundance of shrubby páramo species was significantly higher far from the road than near the road (Far = 36, Near = 25). Moreover, we found a tendency towards differences in the composition of bird habitat groups between transects near and far from the road. One aspect potentially driving the observed patterns was the increasing proportion of planted non-native woody tree species within páramo grassland near the road, which may have caused reduced abundances of shrubby páramo bird species there. While roads represented a clear impact on the composition of bird species in the páramo, the major effect seems to be driven by the introduction of non-native plant species along the roadside. In order to reduce the impact of roads to a minimum, we suggest that park managers should control the introduction of such plant species.     

Ecophysiology and genetic diversity in species of the bamboo Chusquea in the high Andes,Venezuela

ABSTRACT

Background: Woody bamboos of the genus Chusquea grow along a broad range of elevations in the Venezuelan Andes. Their growth-form and density vary along the cloud forest – páramo gradient. In this article, we related ecophysiological traits and population genetic diversity information to explain the distribution of growth-form patterns of Chusquea in the Merida Andes, Venezuela.

Aims: We quantified differences in the ecophysiological response and genetic diversity of scandent cloud forest and shrub-like páramo bamboos of the genus Chusquea, taking in account the differences in their flowering patterns, growth-form and habitat.

Methods: We related low temperature resistance, water relations and leaf gas exchange variables to the growth-form, habitat, flowering patterns and genetic diversity in species of Chusquea. The genetic diversity study was based on Inter Sequence Simple Repeats and Random Amplified Polymorphic DNA markers analysis of cloud forest and páramo populations.

Results: Scandent cloud forest and shrub-like páramo species of Chusquea had a very similar ecophysiological response for all the variables analysed during wet and dry seasons and were capable of enduring freezing temperatures through moderate supercooling. Species associated with the cloud forest – páramo gradient maintained low stomatal conductance and transpiration rates that favoured high leaf water potentials, without limiting photosynthetic rates. Shrub-like bamboos growing above the continuous forest line had a small decline in net photosynthesis rates, associated with an increase in water use efficiency. Both scandent and shrub-like bamboos had a remarkably high genetic diversity, comparable to non-clonal species.

Conclusions: Species of Chusquea in the Venezuelan Andes are a physiologically relatively homogeneous group across a broad elevation gradient. Population genetic diversity appears to be more related with their flowering pattern and habitat conditions than with their growth form.     

Influence of habitat modification by livestock on páramo bird abundance in southern Andes of Ecuador

Livestock grazing on natural grasslands is widespread with negative consequences to biodiversity. In the High Andes, páramo grassland is a distinctive ecosystem where the influence of livestock grazing on páramo birds is poorly documented. We assessed the influence of habitat modification of páramo grassland related to livestock grazing on bird habitat guilds in the southern Andes of Ecuador. We recorded birds occurring along transects located in areas which showed a gradient (low to high) of grazing pressure. We found a decrease in abundance of páramo specialists in transects with more grazing pressure. We interpret this habitat modification as loss of key habitat necessary for habitat-specialized birds.     

<Emphasis Type="Italic">Espeletia</Emphasis> giant rosette plants are reliable biological indicators of time since fire in Andean grasslands

Páramo grasslands in the tropical Andes are fire-prone ecosystems and an understanding of their fire ecology is fundamental to biodiversity conservation and ecosystem management. Fire registers are normally impractical in these remote, cloud-covered landscapes, but Espeletia giant rosette plants have been proposed as biological indicators of time since fire in páramos. Espeletia giant stem rosettes tolerate fire well, protecting apical buds in at the heart of their leaf rosettes, and for some species, germination is known to be enhanced by fire. As the plant grows, its dead leaves remain attached to the stem, but fire removes these and resets the “leaf clock”. This study uses a unique register of fires in one Ecuadorian páramo to assess the robustness of this biological indicator. Dead leaf cover on Espeletia pycnophylla giant rosette plants was measured in fifteen different sites with known fire dates from 2000 to 2014. The growth rates of plants at four different elevations were measured over a 2-year period and used to estimate time since fire based on dead leaf cover in the known sites. Estimates were accurate to ± 2 year. Thus, where fire records are missing, relatively easy measurements of growth rates and dead leaf cover of Espeletia giant rosette plants can provide reliable estimates across a wide range of times since fire. This approach has value for direct investigations into fire ecology but also for studies in which controlling for fire dynamics is necessary to reveal underlying patterns. Therefore, this approach also offers a means to obtain better information on other landscape-scale processes such as the impact of climate change on biodiversity or the provision of ecosystem services.     

Ecophysiology of frailejones (<Emphasis Type="Italic">Espeletia</Emphasis> spp.), and its contribution to the hydrological functioning of páramo ecosystems

Páramos are high elevation tropical ecosystems in northern Andes, with large water yield and water regulation. One of the main and representative species growing in these páramos is the genus Espeletia, known as frailejones. There is a lack of knowledge of Espeletia ecophysiology, maybe due to its unusual anatomical modifications and the specific climatic conditions of these ecosystems. Therefore, it is important to determine the relationships between the anatomical modifications of Espeletia, its physiological functioning, and its contribution to the ecohydrologic functioning of páramos. Consequently, we studied the physiology of frailejones in two Colombian páramos, focused on the identification of conductive tissues inside the stems, calculated the age, and measured sapflow, using the heat ratio method. Results show that Espeletia spp. have a central pith that increases with height, as the size of secondary xylem decreases. Frailejones respond quickly to the changing conditions of weather factors controlling transpiration such as solar radiation, temperature, and fog presence. However, although environmental factors favor transpiration, the sapflow tends to decrease—a particular behavior of the Espeletia transpiration processes—since this occurs chaotically over time, including sapflow at night. The transformation of sapflow velocity to depth of water in a basin shows that the water lost through their transpiration is very low, which contributes to the high runoff ratio of páramo ecosystems. For the first time, we determine by radiocarbon the real ages of three E. hartwegiana, and their mean growth rates to range between 3.8 and 6.9 cm year^?1.     

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.     

Una nueva especie de Chusquea (Poaceae: Bambusoideae) de la Cordillera de Mérida, Venezuela

A new species of woody bamboo (Poaceae: Bambusoideae),Chusquea guirigayensis, is described and illustrated. This species is placed inChusquea sect.Swallenochloa (McClure) L. G. Clark because of its shrubby habit, short waxy internodes, intravaginal branching, and linearly arranged subsidiary buds/branches. In addition, we discuss related species, and describe the ecology and distribution of the new species. Separate keys based on sterile and fertile material of the Venezuelan species ofC. sect.Swallenochloa are provided.     

Ultrastructural changes and dynamic expressions of FAD7, Cu/Zn-SOD,and Mn-SOD in Neosinocalamus affinis under cold stress

As one of the fast-growing species, bamboo plays an important role in ecological stability and wood processing industry. However, low temperature limitation is the basic problem for the cultivation and introduction of bamboo. In this study, the symptoms of cold stress influence on the native bamboo (Neosinocalamus affinis (Rendle) Keng f.) and hybrid bamboo (Bambusa pervariabilis × Dendrocalamopsis grandis) were observed under transmission electron microscope, and the dynamic responses of FAD7, Cu/Zn-SOD, and Mn-SOD genes to cold stress were identified in bamboo by real-time quantitative RT-PCR. Observation by electron microscopy indicated that bamboo is one of the most chilling-sensitive species with severe ultrastructural injury induced by chilling, but the native bamboo (N. affinis) is more cold-tolerant compared with the hybrid bamboo. Results obtained by real-time quantitative RT-PCR analysis revealed that FAD7, Cu/Zn-SOD, and Mn-SOD were all cold-inducible genes in N. affinis. In addition, dynamic response patterns of N. affinis Cu/Zn-SOD and Mn-SOD under cold stress were similar. This work is a fundamental research of hardiness physiology of bamboo and may contribute to the breeding program on obtaining transgenic bamboo species.     

Diversity of zonal páramo plant communities in Ecuador

Patterns of vascular plant species diversity in high‐altitude Ecuadorian ecosystems (‘páramos’) are examined. Data from two independent surveys were used: the first from 12 different locations and 192 samples, the other from 18 locations and 243 samples. These surveys included 348 and 284 species, respectively. The data confirmed the occurrence of two main zones in terms of vascular plant species diversity. The grass páramo and superpáramo were distinguished by differences in plant cover, species richness, α‐diversity and β‐diversity. The transition between these two zones begins at around 4000 m. Grass páramo samples comprised more species but the strong dominance of tussock grasses resulted in low equitability compared with the superpáramo, where safe sites for plant survival are limited and the environment does not permit continuous grass cover. Turnover of species across the altitudinal gradient is higher in the grass páramo than in the superpáramo. This is due largely to agricultural fires at lower altitudes, which create a fine‐scale mosaic of burned patches that enhances variability at this scale. Despite the loss of equitability, intermediate levels of fire disturbance appear to promote species richness within the samples. It is suggested that the complex patterns of páramo diversity in the Ecuadorian Andes are largely the outcome of three interrelated factors: altitude, disturbance and the availability of safe sites at the highest altitudes.     

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.     

Phytogeography of the bryophyte floras of oak forests and páramo of the Cordillera de Talamanca, Costa Rica

Aim Central America is a biogeographically interesting area because of its location between the rich and very different biota of North and South America. We aim to assess phytogeographical patterns in the bryophyte floras of oak forests and páramo of the Cordillera de Talamanca, Costa Rica. Location Tropical America, in particular the montane area of Cordillera de Talamanca, Costa Rica. Methods The analysis is based on a new critical inventory of the montane bryophyte flora of Cordillera de Talamanca. All species were assigned to phytogeographical elements on the basis of their currently known distribution. Absolute and percentage similarities were employed to evaluate floristic affinities. Results A total of 401 species [191 hepatics (liverworts), one hornwort, 209 mosses] are recorded; of these, 251 species (128 hepatics, one hornwort, 122 mosses) occur in oak forests. Ninety‐three per cent of all oak forest species are tropical in distribution, the remaining 7% are temperate (4%) and cosmopolitan (3%) species. The neotropical element includes almost 74% of the species, the wide tropical element (pantropical, amphi‐atlantic, amphi‐pacific) only 19%. A significant part of the neotropical species from oak forests are species with tropical Andean‐centred ranges (27%). As compared with bryophyte species, vascular plant genera in the study region are represented by fewer neotropical, more temperate and more amphi‐pacific taxa. Bryophyte floras of different microhabitats within the oak forest and epiphytic bryophyte floras on Quercus copeyensis in primary, early secondary and late secondary oak forest show a similar phytogeographical make‐up to the total oak forest bryophyte flora. Comparison of oak forest and páramo reveals a greater affinity of the páramo bryophyte flora to temperate regions and the great importance of the páramo element in páramo. Surprisingly, oak forests have more Central American endemics than páramo. Main conclusions (1) Providing first insights into the phytogeographical patterns of the bryophyte flora of oak forests and páramo, we are able to confirm general phytogeographical trends recorded from vascular plant genera of the study area although the latter were more rich in temperate taxa. (2) Andean‐centred species are a conspicuous element in the bryophyte flora of Cordillera de Talamanca, reflecting the close historical connection between the montane bryophyte floras of Costa Rica and South America. (3) High percentages of Central American endemics in the bryophyte flora of the oak forests suggest the importance of climatic changes associated with Pleistocene glaciations for allopatric speciation.     

Some Umbelliferae of the Andean paramos of South America

Two Colombian species described earlier by the authors underPrionosciadium, and a hitherto unnamed collection from the same country are systematically compared with North AmericanPrionosciadium and the monotypic generaCotopaxia (Ecuador) andPerissocoeleum (Colombia). Habit, foliage, inflorescence, fruit, and distribution all suggest that the páramo species exhibit considerable unity ifCotopaxia is excluded. It is indicated also that the genusPrionosciadium has been attributed to South America incorrectly. Despite the somewhat discordant evidence obtained from pollen morphology, it is concluded that the most feasible disposition of the four Colombian plants is to unite them underPerissocoeleum.     

Notes on Collybia V. Gymnopus section Levipedes in tropical South America,with comments on Collybia

Five species of Gymnopus section Levipedes are reported for tropical South America. Two of these, Gymnopus macropus and G. nubicola, are newly described. Collybia laccata is considered synonymous with G. mucubajiensis comb. nov. now known from Andean Ecuador; both were originally described from the Venezuelan páramo. Gymnopus alkalivirens and G. semihirtipes are compared to G. mucubajiensis. Gymnopus spongiosus, previously known from the eastern United States, is newly reported from Colombian Quercus forests and is proposed as a new combination. Gymnopus dryophilus is, at present, not known from tropical South America, but extralimital material from nearby Costa Rica is described for comparison. No known species of Collybia s.str. have been identified from tropical South America.     

<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.     

Chusquea sect. Swallenochloa (Poaceae: Bambusoideae) and allies in Brazil

The 13 high altitude/latitude, dwarf species ofChusquea in Brazil are described, illustrated, and mapped, and their morphology, habitats, distributions, and taxonomic affinities are discussed. Two keys to species are provided, one based solely on vegetative characters, and the other on vegetative and flowering characters.Chusquea erecta, C. nutans, C. riosaltensis,C. windischii, C. caparaoensis, andC. nudiramea are described as new, andC. microphylla is elevated to specific status. Two subspecies are recognized within the variableC. mimosa: C. mimosa subsp.australis and subsp.mimosa. Seven species are formally classified withinChusquea sect.Swallenochloa; the remaining six species are classified into two informal categories, theNudiramea andHeterophylla groups. A list of all the species currently included withinChusquea sect.Swallenochloa is provided.     

Plutarchia ecuadorensis (Ericaceae,Vaccinieae): a new species and a new generic record for Ecuador

Plutarchia, a shrubby páramo genus, is here reported for the first time in Ecuador where two species are recognized: P. angulata A. C. Smith and the new P. ecuadorensis Luteyn. These two species are keyed, described, and the new species illustrated.     

A revision ofTaraxacum sect.Piesis (Compositae)

On the basis of allozyme and cultivation data, and of additional herbarium material, a taxonomic and nomenclatural revision ofTaraxacum sect.Piesis A.J. Richards exKirschner et?těpánek is provided. The section is made up of halophilous, sexually reproducing taxa. InT. stenocephalum Boiss. etKotschy,T. pindicum Kirschner et?těpánek, sp. nov., andT. perenne Kirschner et?těpánek, sp. nov., a tetraploid chromosome number has been recorded, representing the only known case of sexuality at the tetraploid level in the genus. The complex ofT. stenocephalum, includes some geographically and morphologically extreme populations treated as subspecies: subsp.gumusanicum (Soest)Kirschner et?těpánek, comb. nov., subsp.magnum Kirschner et?těpánek, subsp. nov., and subsp.daralagesicum (Schischk.)Kirschner et?těpánek, comb. nov. In addition toT. bessarabicum (Hornem.)Hand.-Mazz., a widely distributed Eurasian species,T. stenocephalum, a complex centred in Transcaucasia and Anatolia, andT. pachypodum H. Lindb., a North African endemic, four new species are described:T. salsum Kirschner et?těpánek, sp. nov., a diploid endemic confined to E Crimea,T. perenne Kirschner et?těpánek, sp. nov., a tetraploid sexual species known only from SW Crimea,T. pindicum Kirschner et?těpánek, sp. nov., a remarkable tetraploid endemic to the Pindos Mts., Greece, andT. salsitatis Kirschner, ?těpänek etYirdirimli, sp. nov., an Anatolian diploid species. Furthermore, a hybrid betweenT. salsum andT. bessarabicum from Crimea (documented on the basis of allozyme data elsewhere) is given a binomial,T. xmesohalobium Kirschner et?těpánek, nothosp. nov.