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.     

Above-ground biomass structure of a Chusquea tessellata bamboo páramo,Chingaza National Park,Cordillera Oriental,Colombia

The present study was carried out in the bamboo (Chusquea tessellata) páramo of Parque Natural Nacional de Chingaza, Eastern Cordillera, Colombia from December 1987 to April 1988. Above-ground biomass structure of bamboo páramo was quantified in 16 plots. These data are compared with previous results on above-ground biomass structure of bunch-grass (Calamagrostis spp.) páramos.The total (non-living and living) above-ground biomass of a Chusquea tessellata bamboo páramo was low (2,625 g DW · m^–2) compared to bunch-grass páramo. Nevertheless, higher values of standing living biomass and litter are found in the bamboo páramo due to the leaf shed of the bamboo. The thick litter layer may inhibit germination and growth of nearby plants.Maximum biomass is found near the ground surface. Cumulative LAI (In transformed) and height in the bamboo vegetation are related parabolically for Chusquea tessellata and linearly for bunch-grass due to differences in leaf distribution. The mean bifacial LAI of living Chusquea tessellata leaves is 2.2 m² · m^–2, whereas it is 2.5 m² · m^-2 for all Poaceae.     

<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.     

Ordination and classification of vegetation along an altitudinal gradient in the Venezuelan páramos

By means of ordination and classification techniques, the relationships between climate, soils, human activities and vegetation along an altitudinal gradient of the Venezuelan páramos are analyzed and interpreted. The altitudinal gradient chosen is characterized by decrease of temperature, precipitation, soil fertility, soil water-holding capacity, and plant cover as altitude increases. The ordination results suggest vegetation changes to be primarily related to environmental changes occurring with altitude, and secondly to disturbances caused mainly by grazing. Some results point toward a disjunction in the vegetational gradient occurring at ca. 3 500 m.a.s.l. and separating low and high páramo. This disjunction might have been caused by the glacial history of the páramos and the occurrence of frequent night-frosts.The soil samples were kindly analyzed by the Laboratorio de Edafologia, Centro Nacional de Investigaciones Agropecuarias. Help in plant identification was generously obtained from the specialists of Instituto Botánico, Instituto Nacional de Parques, Caracas.Nomenclature follows Vareschi (1970).Acknowledgements: This work was supported by the Decanato de Investigaciones, Universidad Simón Bolivar. I wish to thank A. Pacheco for help in the field sampling. Dr O. Arenas help was invaluable in the mathematical treatment of the data. Drs A. Vivas and J. M. B. Smith provided useful criticism to an earlier version of this work.     

The growth form composition of plant communities in the ecuadorian páramos

A growth form classification for the plants of the Ecuadorian páramos is proposed, consisting of ten forms: stem rosettes, basal rosettes, tussocks, acaulescent rosettes, cushions and mats, upright shrubs, prostrate shrubs, erect herbs, prostrate herbs, and trailing herbs.The growth form composition of 192 samples of páramo vegetation from twelve different regions in Ecuador was analysed using multivariate techniques to determine physiognomic types. The distribution of growth forms was significantly related to altitude, rock cover, bare ground and to measures of disturbance and exposure.The dominant growth forms in all the samples belonged to one of Hedberg's (1964) five types, but other forms were also significant components of the vegetation. The majority of Ecuadorian páramo vegetation showed a relatively consistent growth form composition, dominated by tussocks. The accompanying growth forms were mostly acaulescent rosettes, cushions, upright shrubs, prostrate shrubs, erect herbs and prostrate herbs, sometimes with stem rosettes, basal rosettes or trailing herbs.At higher altitudes, the dominance of tussocks was reduced. At first, acaulescent rosettes became dominant, but higher still their dominance was shared with cushions. At the highest altitudes of all, where plant cover was thin, no single growth form was dominant. In other locations where plant cover was sparse, once again no single growth form was dominant. In humid páramos, stem rosettes were co-dominant with tussocks or erect herbs. Basal rosettes, erect herbs and prostrate herbs were locally co-dominant at higher altitudes.The growth form composition of the Ecuadorian páramos showed similarities with other tropical alpine regions, though no comparable quantitative data for these regions are available yet. The quantitative determination of growth form composition may also lead to a better understanding of community structure and the mechanisms which govern it.     

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.     

Patterns of energy content in plants from the venezuelan paramos

Summary Determinations of energy and ash content were made on the organs of some common plant species from an elevational gradient in the Venezuelan páramos. The results indicate differences in energy content among the life forms sampled. Giant rosette plants show the highest values in the roots while woody plants have the highest energy content in the leaves. The reproductive organs show a higher energy content than the vegetative organs. There is a tendency to increase energy content as elevation and environmental stress increase. The adaptive significance of these patterns is discussed. The high average energy value for páramo plants, 20.18 KJ/g ash free dry weight, is similar to values reported for alpine tundra plants and conifer forests.     

Pubescence,floral temperature and fecundity in species of Puya (Bromeliaceae) in the Ecuadorian Andes

Summary The effects of inflorescence pubescence on floral temperatures and fecundity were examined in four species of Puya in the Ecuadorian Andes. These species span an elevational range from 1,980–4,000 m and comparisons among these species showed that pubescence production increases significantly with elevation. Flower temperatures of glabrous or slightly pubescent species of Puya from lower elevations closely tracked air temperature, while those of the pubescent páramo species did not. Pubescence removal experiments on Puya hamata, a páramo species, demonstrated that pubescence is an effective insulator, maintaining elevated flower temperatures. In Puya clava-herculis (also a páramo species) elevated flower temperatures were associated with higher seed set. Thus, the greater production of inflorescence pubescence in páramo species of Puya may be an important factor contributing to reproductive success in these higher elevation species.     

Superpáramo plant species diversity and phytogeography in Ecuador

In 18 superpáramo sites in Ecuador we found 388 species of vascular plants belonging to 146 genera and 52 families, making the Ecuadorian superpáramo flora richer in species than that of Venezuela and that of Colombia which appears to have fewer species although the number remains uncertain. The most species rich families were Asteraceae (83 species) and Poaceae (49) which also dominate the grasspáramo that surrounds the superpáramo. Otherwise the superpáramo is dominated by families that are mostly herbaceous such as Cyperaceae, Brassicaceae, Caryophyllaceae and Valerianaceae, whereas shrubby families that dominate the subpáramo and grasspáramo, such as Melastomataceae, Ericaceae, and Solanaceae, have only few species or are absent in the superpáramo. The generic spectrum was dominated by a suite of species-rich genera (Lachemilla, Gentianella, Valeriana, Calamagrostis, Draba) with many páramo-endemic species reflecting a high level of autochthonous speciation in the (super)páramo. Species richness varied from 71 to 149 in the individual superpáramos surveyed, but species richness was only weakly correlated to their area and log-transformed area, and negatively correlated to the vertical range of each superpáramo. β-diversity was significantly correlated to the vertical range (i.e., the number of surveyed altitudinal levels) but it was not correlated to the area or log-transformed area of each superpáramo. Most of the species were narrowly distributed and 112 (29%) of them were found in a single superpáramo, while eight (2%) occurred in all 18 superpáramo sites. Floristic similarity was not correlated to the geographical distance between the sites. Redundance Analysis suggested that geological origin of the substrate (metamorphic versus volcanic bedrock) was important for the floristic composition. Occurrence of mountains built from metamorphic rocks, is however, correlated to areas with high rainfall and the amount of rainfall may be a stronger determinant for species distribution than the presence of volcanic versus metamorphic bedrocks. TWINSPAN cluster analysis divided the 18 sites into three groups, which corresponded to dry, humid and very humid superpáramos. The groups were also separated along the first ordination axis in Correspondence Analysis, while the second axis may correlate to the volcanic history of the area.     

A new species of piha (Cotingidae: Lipaugus) from the Cordillera Central of Colombia

The Chestnut-capped Piha Lipaugus weberi sp. nov., is described from sub-Andean forest on the northern slope of the Central Cordillera of the Colombia Andes. The new species appears most closely related to Lipaugus fuscocinereus of the Northern Andes including the Central Cordillera, but is much smaller, with a distinctive chestnut-brown crown, yellow orbital ring, two modified primaries in the male, an overall darker grey coloration and unique vocalizations. It appears to be restricted to a narrow belt of premontane very humid forests (1500–1820 m asl) where it is fairly common. The restricted range and specific ecological requirements of Lipaugus weberi make the species of great conservation concern as the Central Cordillera has been severely deforested and remaining forests are highly fragmented. We present notes on the behaviour, ecology and conservation of this new species.
Se describe Lipaugus weberi sp. nov, de bosque subandino de la vertiente norte de la Cordillera Central de los Andes de Colombia. La nueva especie parece estar bastante relacionada con Lipaugus fuscocinereus de los Andes septentrionales incluyendo la Cordillera Central, pero es mucho más pequeña, presenta una distintiva gorra castaño opaco, anillo ocular amarillo, dos remeras primarias modificadas, la coloración gris más oscura y vocalizaciones ünicas. Esta especie se restringe a una franja angosta de bosque muy hümedo premontano (1500–1820 msnm), donde es relativamente comün. El rango restringido y los requerimientos ecológicos especificos de Lipaugus weberi hacen que esta especie sea de gran relevancia para la conservación, más aün debido al estado de deforestación severa de la Cordillera Central, en la cual los bosques remanentes son muy fragmentados. Presentamos anotaciones sobre la ecologia, el comportamiento y la conservación de esta nueva especie.     

The calanoid fauna (Crustacea,Copepoda) of the Cordillera Oriental of the Colombian Andes

Calanoid copepods belonging to four species and one subspecies from lakes and ponds mainly from the Cordillera Oriental (22) and Cordillera Central (1) of the Colombian Andes are recorded. Most of the water bodies studied are located in the páramo region, between 2996 and 4085 m altitude. The family Boeckellidae is recorded for the first time from Colombia. Taxonomic characteristics of Boeckella occidentalis Marsh, 1906, Boeckella gracilis (Daday,1902) and the diaptomid Prionodiaptomus colombiensis (Thiébaud, 1914) are discussed. A new diaptomid genus, Colombodiaptomus, with one species and one subspecies are established.     

Seed Dispersal in Páramo Plants: Epizoochorous and Hydrochorous Taxa

Abstract: On the basis of a recent checklist of plant diversity in páramos, diaspores collected from herbaria were studied for adaptations to dispersal on animals and by water. This study shows that the páramo flora has a relatively high percentage of genera with morphological adaptations to epizoochorous and to hydrochorous diaspore dispersal. Genera with hooked and straight appendages are present throughout the páramo belt, while their number decreases in the higher páramo zones. About half of the hydrochorous genera and one-third of the epizoochorous ones can be found throughout all páramo zones. The contribution of holarctic epizoochorous genera to the páramo flora seems to be greater than that of austral-antarctic genera, whereas in hydrochorous genera it is the reverse.     

Areas of endemism and distribution patterns for Neotropical Piper species (Piperaceae)

Aim The study aimed to establish areas of endemism and distribution patterns for Neotropical species of the genus Piper in the Neotropical and Andean regions by means of parsimony analysis of endemicity (PAE) and track‐compatibility analysis. Location The study area includes the Neotropical region and the Northern Andean region (Páramo‐Punan subregion). Methods We used distribution information from herbarium specimens and recent monographic revisions for 1152 species of Piper from the Neotropics. First, a PAE was attempted in order to delimit the areas of endemism. Second, we performed a track‐compatibility analysis to establish distribution patterns for Neotropical species of Piper. Terminology for grouping Piper is based on recent phylogenetic analyses. Results The PAE yielded 104 small endemic areas for the genus Piper, 80 of which are in the Caribbean, Amazonian and Paranensis subregions of the Neotropical region, and 24 in the Páramo‐Punan subregion of the Andean region. Track‐compatibility analysis revealed 26 generalized tracks, one in the Páramo‐Punan subregion (Andean region), 19 in the Neotropical region, and six connecting the Andean and Neotropical regions. Both the generalized tracks and endemic areas indicate that distribution of Piper species is restricted to forest areas in the Andes, Amazonia, Chocó, Central America, the Guayana Shield and the Brazilian Atlantic coast. Main conclusions Piper should not be considered an Andean‐centred group as it represents two large species components with distributions centred in the Amazonian and Andean regions. Furthermore, areas of greater species richness and/or endemism are restricted to lowland habitats belonging to the Neotropical region. The distribution patterns of Neotropical species of Piper could be explained by recent events in the Neotropical region, as is the case for the track connecting Chocó and Central America, where most of the species rich groups of the genus are found. Two kinds of event could explain the biogeography of a large part of the Piper taxa with Andean–Amazonian distribution: pre‐Andean and post‐Andean events.     

Taxonomic revision of hispaniola tiger beetles in the genus brasiella rivalier 1954 (coleoptera, carabidae, cicindelinae)

The Brasiella tiger beetle fauna on Hispaniola, the second largest island of the Greater Antilles, has more species diversity than currently recognized as all populations previously have been assigned to the insular endemic Brasiella dominicana (Mandl). A comparative study of adult morphology, particularly male genitalic and female abdominal characters, for available Brasiella specimens from populations on Hispaniola, proposes eight additional new species also endemic to this island. Except for three sympatric species in the Sierra de Baoruco in southern Dominican Republic occurring in different habitats, all the Brasiella on Hispaniola appear to be allopatric. Most species occur in the major mountainous regions of Hispaniola. Two species, however, are known only from river floodplains in the southern coastal plain of the Dominican Republic. Brasiella dominicana (Mandl) and Brasiella ocoa, new species, occur along river floodplains emanating from the eastern end of the Cordillera Central in the Dominican Republic. Two new Brasiella species, Brasiella bellorum, and Brasiella philipi, occur in the Cordillera Central, Dominican Republic, the former species from central portions, and the latter species from north slopes of this mountain range, respectively. Three new Brasiella species, Brasiella rawlinsi, Brasiella iviei, and Brasiella youngi, are isolated in the Sierra de Baoruco, Dominican Republic, where each occupies a different habitat along an altitudinal gradient. The two new Brasiella species in Haiti are Brasiella darlingtoniana, in the Massif de la Selle, and Brasiella davidsoni, in the Massif de la Hotte. All nine Brasiella species on Hispaniola, along with Brasiella viridicollis (Dejean) and its two subspecies on Cuba, belong to the viridicollis species group of the genus Brasiella based on criteria presented in earlier published phylogenetic studies of Brazilian and West Indian tiger beetles. The subspecies Brasiella viridicollis fernandozayasi (Kippenhan, Ivie and Hopp) may represent a distinct species within this species group, whereas removal of Brasiella wickhami (W. Horn) from this species group seems warranted based on evidence presented. A general overview of species relationships for the Brasiella on Hispaniola are discussed, along with the current and ancestral geographic distributions of the Brasiella viridicollis species group in the West Indies.     

Spatio‐temporal history of the endemic genera of Madagascar

Madagascar is renowned for its unparalleled species richness and levels of endemism, which have led, in combination with species extinction caused by an unprecedented rate of anthropogenic deforestation, to its designation as one of the most important biodiversity hotspots. It is home to 10 650 species (84% endemic) of angiosperms in 1621 genera (19% endemic). During the last two centuries, botanists have focused their efforts on the provision of a taxonomic framework for the flora of the island, but much remains to be investigated regarding the evolutionary processes that have shaped Madagascan botanical diversity. In this article, we review the current state of phylogenetic and biogeographical knowledge of the endemic angiosperm genera. We also propose a new stratified biogeographical model, based on palaeogeographical evidence, allowing the inference of the spatio‐temporal history of Madagascan taxa. The implications of past climate change and extinction events on the evolutionary history of the endemic genera are also discussed in depth. Phylogenetic information was available for 184 of the 310 endemic genera (59.3%) and divergence time estimates were available for 67 (21.6%). Based on this evidence, we show the importance of phylogenetic clustering in the assemblage of the current Madagascan diversity (26% of the genera have a sister lineage from Madagascar) and confirm the strong floristic affinities with Africa, South‐East Asia and India (22%, 9.1% and 6.2% of the genera, respectively). The close links with the Comoros, Mascarenes and Seychelles are also discussed. These results also support an Eocene/Oligocene onset for the origin of the Madagascan generic endemic flora, with the majority arising in the Miocene or more recently. These results therefore de‐emphasize the importance of the Gondwanan break‐up on the evolution of the flora. There is, however, some fossil evidence suggesting that recent extinctions (e.g. Sarcolaenaceae, a current Madagascan endemic, in southern Africa) might blur vicariance patterns and favour dispersal explanations for current biodiversity patterns. © 2013 The Linnean Society of London     

Aulonemia bogotensis (Poaceae: Bambusoideae), a new species from the Cordillera Oriental of Colombia

A new species of Aulonemia from the páramos of the Cordillera Oriental, Cundinamarca, Colombia, is described and illustrated. Aulonemia bogotensis is distinguished by its diminutive size, abaxially tomentose foliage leaf blades, and spikelets with muticous lemmas. Two similar Colombian endemics, A. pumila and A. trianae, are compared and contrasted with the new species.     

Adaptation and Convergent Evolution within the Jamesonia-Eriosorus Complex in High-Elevation Biodiverse Andean Hotspots

The recent uplift of the tropical Andes (since the late Pliocene or early Pleistocene) provided extensive ecological opportunity for evolutionary radiations. We test for phylogenetic and morphological evidence of adaptive radiation and convergent evolution to novel habitats (exposed, high-altitude páramo habitats) in the Andean fern genera Jamesonia and Eriosorus. We construct time-calibrated phylogenies for the Jamesonia-Eriosorus clade. We then use recent phylogenetic comparative methods to test for evolutionary transitions among habitats, associations between habitat and leaf morphology, and ecologically driven variation in the rate of morphological evolution. Páramo species (Jamesonia) display morphological adaptations consistent with convergent evolution in response to the demands of a highly exposed environment but these adaptations are associated with microhabitat use rather than the páramo per se. Species that are associated with exposed microhabitats (including Jamesonia and Eriorsorus) are characterized by many but short pinnae per frond whereas species occupying sheltered microhabitats (primarily Eriosorus) have few but long pinnae per frond. Pinnae length declines more rapidly with altitude in sheltered species. Rates of speciation are significantly higher among páramo than non-páramo lineages supporting the hypothesis of adaptation and divergence in the unique Páramo biodiversity hotspot.     

The Taxonomic Status of Mazama bricenii and the Significance of the Táchira Depression for Mammalian Endemism in the Cordillera de Mérida,Venezuela

We studied the taxonomy and biogeography of Mazama bricenii, a brocket deer classified as Vulnerable by the IUCN, drawing on qualitative and quantitative morphology and sequences of the mitochondrial cytochrome-b gene. We used Ecological Niche Modeling (ENM) to evaluate the hypothesis that M. bricenii of the Venezuelan Cordillera de Mérida (CM) might have become isolated from populations of its putative sister species, Mazama rufina, in the Colombian Cordillera Oriental (CO). This hypothesis assumes that warm, dry climatic conditions in the Táchira Depression were unsuitable for the species. Our analyses did not reveal morphological differences between specimens geographically attributable to M. bricenii and M. rufina, and phylogenetic analyses of molecular data recovered M. bricenii nested within the diversity of M. rufina. These results indicate that M. bricenii should be regarded as a junior synonym of M. rufina. ENM analyses revealed the existence of suitable climatic conditions for M. rufina in the Táchira Depression during the last glacial maximum and even at present, suggesting that gene flow between populations in the CO and CM may have occurred until at least the beginning of the current interglacial period and may continue today. Because this pattern might characterize other mammals currently considered endemic to the CM, we examined which of these species match two criteria that we propose herein to estimate if they can be regarded as endemic to the CM with confidence: (1) that morphological or molecular evidence exists indicating that the putative endemic taxon is distinctive from congeneric populations in the CO; and (2) that the putative endemic taxon is restricted to either cloud forest or páramo, or both. Only Aepeomys reigi, Cryptotis meridensis, and Nasuella meridensis matched both criteria; hence, additional research is necessary to assess the true taxonomic status and distribution of the remaining species thought to be CM endemics.     

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.