Densidad Poblacional y Disponibilidad de Híbitat de la Danta de Montaña (Tapirus pinchaque) en los Andes Centrales de Colombia1

Population density of Tapirus pinchaque was estimated in 7000 ha Parque Nacional Natural Los Nevados in the central Andes of Colombia. Eight variables were measured in 110 left rear foot prints of the mountain tapir: total length and width, and length and width of the three toes. This information was analyzed with Cluster Analysis by means of UPGMA and Euclidean distance using the scores obtained by means of a Principal Component Analysis (PCA) using the eight variables. Habitat availability was measured in 147,000 ha using aerial photographs taken during the 1950s and 1980s. The results of the cluster analysis showed that there are between 11 and 15 individuals, for a mean density of one individual for each 551 ± 85 ha. The measurements of forest cover in the 147000 ha showed that habitat availability for the mountain tapir (montane forests and forestry plantations) increased 18l4lha from the 1950s to the 1980s. This increment in forest cover seems to have a positive effect on the local population size of T. pinchaque, since in areas previously covered by pastures, there are now secondary forests where tapirs tracks and feces can be found.     

DNA Barcoding Survey of Anurans across the Eastern Cordillera of Colombia and the Impact of the Andes on Cryptic Diversity

Colombia hosts the second highest amphibian species diversity on Earth, yet its fauna remains poorly studied, especially using molecular genetic techniques. We present the results of the first wide-scale DNA barcoding survey of anurans of Colombia, focusing on a transect across the Eastern Cordillera. We surveyed 10 sites between the Magdalena Valley to the west and the eastern foothills of the Eastern Cordillera, sequencing portions of the mitochondrial 16S ribosomal RNA and cytochrome oxidase subunit 1 (CO1) genes for 235 individuals from 52 nominal species. We applied two barcode algorithms, Automatic Barcode Gap Discovery and Refined Single Linkage Analysis, to estimate the number of clusters or “unconfirmed candidate species” supported by DNA barcode data. Our survey included ~7% of the anuran species known from Colombia. While barcoding algorithms differed slightly in the number of clusters identified, between three and ten nominal species may be obscuring candidate species (in some cases, more than one cryptic species per nominal species). Our data suggest that the high elevations of the Eastern Cordillera and the low elevations of the Chicamocha canyon acted as geographic barriers in at least seven nominal species, promoting strong genetic divergences between populations associated with the Eastern Cordillera.     

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.     

Facilitation by pioneer shrubs for the ecological restoration of riparian forests in the Central Andes of Colombia

Riparian vegetation has great functional importance in agroscapes because it establishes physical connections between natural ecosystems embedded in an agricultural matrix. Throughout the tropics, the historical demand for cropland and pastures in the lands adjacent to streams has led to the replacement of forests by exotic grasses. In order to find new methods for initiating the restoration of riparian forests, we evaluated the use of the pioneer shrubs Tithonia diversifolia and Piper auritum as nurse species for their effects on (1) competition with dominant grasses; (2) natural regeneration and the ecological facilitation of six native tree species of different successional stages; and (3) influence on microclimate. Over a period of 15 months, 4.4% coverage of P. auritum was insufficient to inhibit grasses. In contrast, 81% coverage of T. diversifolia limited the growth of dominant grasses such as Cenchrus purpureus, Paspalum paniculatum, and Cynodon plectostachyus likely by intercepting more than 90% of photosynthetically active radiation (PAR), even though other factors cannot be ruled out. T. diversifolia showed simultaneous effects of facilitation and competition by promoting higher survival of the planted native trees while slowing the growth of pioneer species and inhibiting the regeneration of native woody plants probably as a result of high light interception. This study suggests that planting T. diversifolia as a nurse shrub may facilitate early stages of restoration by inhibiting the growth of pastures, but requires pruning, both to increase light availability and promote the growth of planted trees and the recruitment of woody plants.     

Late Quaternary paleosols in the western and central Cordillera of Colombia

More or less well preserved paleosols were studied in the submontane and montane zone of the Western and Central Cordillera, Departamento del Valle, Colombia. Their Pleniglacial age was ascertained by radiocarbon analysis of charcoal which was, in several sections, found associated with the basis of volcanic ash deposits. The two ash periods which provided the bulk of the volcanic parent material seem to have occurred about 30 000 and 10 000 yr. B.P. The paleosols developed before and immediately after the older period (Middle Pleniglacial). The characteristic morphological traits — a uniform grey horizon of low humus content and with a basal iron pan — indicates a cold humid environment of low evapotranspiration and low organic turnover. Such soils do not form today but were characteristic for the Middle Pleniglacial soil cover from altitudes of 1500 to 3500 m. Increasing stable humus content in paleosols after 30 000 yr. B.P. apparently indicate a turn towards drier conditions, i.e. a growing influence of Gramineae after a predominance of wooded vegetation, quite in agreement with the pollen-based paleoclimatic concept of Van der Hammen. This paper demonstrates the distribution of truncated and transported paleosols relicts as a result of the great degradational phase which reshaped the soil mantle probably between 20 000 and 10 000 yr. B.P. The type of paleopedological soil formation process is discussed in some detail.     

Studies in Neotropical Paleobotany. XV. A Mio-Pliocene palynoflora from the Eastern Cordillera, Bolivia: implications for the uplift history of the Central Andes

An assemblage of 33 fossil pollen and spores, recovered from the 3600-m high Pislepampa locality of E. W. Berry, Eastern Cordillera, Bolivia, adds considerably to our knowledge of three aspects of the region in late Neogene time: (1) the paleovegetation, (2) the paleoclimate, and (3) the paleoelevation of the Central Andes. The plant microfossils recognized are Isoetes, Lycopodium (three types), Cnemidaria, Cyathea (three types), Grammitis, Hymenophyllum, Pteris, trilete fern spores (two types), Danaea, monolete fern spores (four types), Podocarpus, Gramineae, Palmae, Ilex, cf. Oreopanax, Cavanillesia, cf. Pereskia, Compositae (three types), Ericaceae, Tetrorchidium, and unknowns (three types). The diversity of the Compositae suggest that this flora has a maximum age around the Miocene-Pliocene boundary, that is, 6-7 million years. All members of the paleocommunity presently grow in the bosque montano húmedo (cloud forest) along the eastern slope of the Central Andes of Bolivia, which occurs between MATs (mean annual temperatures) of ～10° and 20°C. The Pislepampa flora probably represents the lower limits of this forest because the fossil leaves collected by Berry from the same locality all have entire margins, suggesting that the flora grew near the cloud forest-tropical forest transition. Presently, the lower limit of the cloud forest forest has MATs of ～20°C, a mean annual precipitation between 1000 and 1500 mm, and that part containing most of the identified genera of fossil pollen is found at elevations ～1200-1400 m. These conditions are thus inferred for the Pislepampa flora; however, because of the uncertainty of the magnitude of global climate change and of possible changes in the ecological range of plant genera, we estimate an error of at least ±1000 m for the paleoelevation estimate. When the total uplift is corrected for probable amounts of erosionally driven isostatic rebound, the paleoelevation estimate suggests that from one-third to one-half of the uplift of the Eastern Cordillera had occurred by the beginning of the Pliocene. This estimate is consistent with other paleoelevation estimates from the Central Andes.     

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.     

Habitat edge context and the distribution of phyllostomid bats in the Andean forest and anthropogenic matrix in the Central Andes of Colombia

Bats play an important role in the ecosystem as seed dispersers, pollinators and pest controllers. In particular, frugivorous bats are important for regeneration processes in open and degraded areas, because they disperse the seeds of pioneer plant species, which are essential for succession. Depending on the type of habitat that is established at the fragment edge, resources and bat movement patterns toward open areas can be affected. The structure and composition of bats was compared between two ‘interior-edge-pasture’ gradients, in an Andean forest fragment located at the Reserva Natural la Montaña del Ocaso (Quindío, Colombia). The two edge-types considered were forest-edge and bamboo-edge (Guadua angustifolia, Poaceae), both located in the same fragment. Bat abundance was significantly different in the two edge habitats. The forest-edge is a soft edge, in that it allows bat species to move from the interior of the forest to the pasture in front of it. In contrast, the bamboo-edge can be defined as semi-permeable, because it allows less movement of species and individuals from the interior to the pasture. Here we evaluate the possible effects of habitat edge type on bat movement in degraded areas in the main coffee producing region of Colombia.     

Variation in freshwater fish assemblages along a regional elevation gradient in the northern Andes,Colombia

Studies on elevation diversity gradients have covered a large number of taxa and regions throughout the world; however, studies of freshwater fish are scarce and restricted to examining their changes along a specific gradient. These studies have reported a monotonic decrease in species richness with increasing elevation, but ignore the high taxonomic differentiation of each headwater assemblage that may generate high β‐diversity among them. Here, we analyzed how fish assemblages vary with elevation among regional elevation bands, and how these changes are related to four environmental clines and to changes in the distribution, habitat use, and the morphology of fish species. Using a standardized field sampling technique, we assessed three different diversity and two structural assemblage measures across six regional elevation bands located in the northern Andes (Colombia). Each species was assigned to a functional group based on its body shape, habitat use, morphological, and/or behavioral adaptations. Additionally, at each sampling site, we measured four environmental variables. Our analyses showed: (1) After a monotonic decrease in species richness, we detected an increase in richness in the upper part of the gradient; (2) diversity patterns vary depending on the diversity measure used; (3) diversity patterns can be attributed to changes in species distribution and in the richness and proportions of functional groups along the regional elevation gradient; and (4) diversity patterns and changes in functional groups are highly correlated with variations in environmental variables, which also vary with elevation. These results suggest a novel pattern of variation in species richness with elevation: Species richness increases at the headwaters of the northern Andes owing to the cumulative number of endemic species there. This highlights the need for large‐scale studies and has important implications for the aquatic conservation of the region.     

Upper quaternary vegetational and climatic sequence of the fuquene area (Eastern Cordillera,Colombia)

In this paper the results of a pollen-analytical study of three bore holes of lake deposits in the Fuquene-Valle de Ubaté area, Colombian Cordillera Oriental (5° N; elev. 2,580 m) are given. By means of ¹⁴C dates and comparison with other pollen diagrams and dates from the Eastern Cordillera, it was possible to correlate the local pollen zones and chronostratigraphical units with the European chronostratigraphical sequence. The longest diagram (Fuquene II) represents some 30, 000 years, including part of the Middle Pleniglacial, the Upper Pleniglacial, the Late Glacial and the Holocene.During the Middle Pleniglacial, Polylepis wood was an important constituent of the vegetation. During climatic extremes of the Upper Pleniglacial an open páramo vegetation existed.During the main part of the Upper Pleniglacial (from ca. 21,000 to ca. 13,000 B.P.) the lake level was low and the climate dry. The lake level rose again in the beginning of the Late Glacial, and the area around the lake became forested. At the beginning of the El Abra Stadial (Late Dryas Stadial) there was a marked cooling and the lake became lower once more; this had a great impact on the vegetation which became partly open again. The beginning of the Holocene is marked by a gradual increase of forest elements, especially of Quercus (oak.) Oak forests dominated the area during the greater part of the Holocene. During the “hypithermal”, elements growing today at a lower level, such as Cecropia, Acalypha (and possibly Alchornea), must have been growing in the area, intermixed with the oak forest. At the same time there was a considerable increase of Urticaceae in the undergrowth of these forests. The vegetation zones were probably situated several hundreds of metres higher than they are at the present time.The influence of man on vegetation had started already before the beginning of our era, but there is a sudden increase of this influence (e.g., increase of Dodonaea, decrease of Quercus and Urticaceae) somewhere around 2,000 B.P. Later on, the diagram shows the rapid ultimate decline of the forest elements and an increase of Gramineae. Today nothing is left of the original oak forests and their place is taken by arable land or by secondary plant communities with low brushwood.In the lake sediments, thin layers of volcanic ash were found dated approximately 9,000 B.P., 11,000 B.P., and 21,000 B.P., respectively; a series of three ash-layers was deposited between approximately 22,000 B.P. and 26,000 B.P. and one possibly at approximately 30,000 B.P. These layers have been found in several other sections in the Eastern Cordillera and are the basis for a tephrochronology.     

Vegetation patterns,regeneration rates and divergence in an old-field succession of the high tropical Andes

Vegetation restoration during old-field succession was studied in an alpine Andean ecosystem (paramo). 123 plots with different fallow times (1 to 12 years) and 8 plots under natural vegetation were sampled. The results indicate that secondary succession in the paramo, like in other extreme environments, can be interpreted as an autosuccession: there are mainly changes in species relative abundance and little floristic relay (i.e. species turnover). Only a few herbaceous species, mostly introduced (e.g. Rumex acetosella), act as strict pioneers and strongly dominate the early stages. Then, they undergo a progressive decline, while native forbs (e.g. Lupinus meridanus) and grasses (e.g. Vulpia myuros) have their peak abundance in intermediate stages. The characteristic paramo life forms, sclerophilous shrubs (e.g. Baccharis prunifolia, Hypericum laricifolium) and giant rosettes (e.g. Espeletia schultzii), appear very early and gradually increase in abundance during succession, becoming dominant in the late stages and showing a dual behaviour, both as ruderal and stress tolerant species. The 1st axis of a Detrended Correspondence Analysis arranges the sites according to their fallow time. The 2nd and 3rd axes, associated with diverging pathways of regeneration, are correlated with topographic factors and physio-chemical soil characteristics. Hence, structural divergence between plots increases along succession as community composition starts to reflect the conditions of each site. We found evidence of a constant rate of succession during the first 12 years, contradicting the generally accepted hypothesis in the succession literature of a continous slow down up to the climax. Regeneration of vegetation physiognomy is relatively fast, questioning the prevailing idea of slow restoration in alpine ecosystems. However, 12 years of fallow are insufficient to attain the species richness of the natural paramo. Under the current trend of fallow length reduction observed in traditional potato cultivation in the Andes, our results raise doubts about the conservationist value of this management strategy.     

Holocene space-time succession of the Middle Atrato wetlands, Chocó biogeographic region, Colombia

Successional sequences from forested wetlands in the Middle Atrato River Basin were reconstructed using characterisation of present vegetation communities and palynological analysis. A 4.8 km transect, drawn across a river meander, and two 6 and 8 m deep sediment cores (San Martín and Villanueva) were collected in the floodplain within two different vegetation assemblages. Based on the floristic and environmental characteristics of the local vegetation communities, ecological changes spanning the last 4 ka (cal years BP) were analysed in San Martín and Villanueva cores. Present vegetation is dominated by four communities determined by flood tolerance and drainage conditions. We found Euterpe oleraceae, Mauritiella macroclada-Campnosperma panamensis and Oenocarpus bataua forests, and mixed forest and open vegetation in a gradient from poor to improved drainage conditions. Vegetation changes in the palynological record suggest that sedimentation and erosion processes on flood basins are due to changes in drainage conditions and to variable flooding levels.A wet period in the 4 to 2.7 ka interval is postulated, which might be related to sea level rise or local subsidence. Lower flooding levels and improved drainage conditions dominated the 2.7 to ∼ 1.6 ka interval, whereas a flooding event (and a hiatus) occurred between 1.5 and 0.5 ka. This flooding event might be synchronous with analogous events as recorded in the Colombian Amazonia between 1.6 and 1.45 ka. Forest disturbance, probably of anthropogenic origin, is recorded in both sites since 0.5 ka.     

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.     

Elevational Ranges of Montane Birds and Deforestation in the Western Andes of Colombia

Deforestation causes habitat loss, fragmentation, degradation, and can ultimately cause extinction of the remnant species. Tropical montane birds face these threats with the added natural vulnerability of narrower elevational ranges and higher specialization than lowland species. Recent studies assess the impact of present and future global climate change on species’ ranges, but only a few of these evaluate the potentially confounding effect of lowland deforestation on species elevational distributions. In the Western Andes of Colombia, an important biodiversity hotspot, we evaluated the effects of deforestation on the elevational ranges of montane birds along altitudinal transects. Using point counts and mist-nets, we surveyed six altitudinal transects spanning 2200 to 2800m. Three transects were forested from 2200 to 2800m, and three were partially deforested with forest cover only above 2400m. We compared abundance-weighted mean elevation, minimum elevation, and elevational range width. In addition to analysing the effect of deforestation on 134 species, we tested its impact within trophic guilds and habitat preference groups. Abundance-weighted mean and minimum elevations were not significantly different between forested and partially deforested transects. Range width was marginally different: as expected, ranges were larger in forested transects. Species in different trophic guilds and habitat preference categories showed different trends. These results suggest that deforestation may affect species’ elevational ranges, even within the forest that remains. Climate change will likely exacerbate harmful impacts of deforestation on species’ elevational distributions. Future conservation strategies need to account for this by protecting connected forest tracts across a wide range of elevations.     

Altitudinal variation in fish assemblage diversity in streams of the central Andes of Colombia

This study documents differences in fish assemblages for 32 freshwater streams located between 258 and 2242 m a.s.l. on the eastern slopes of the central range of the Colombian Andes. A total of 2049 fishes belonging to 62 species, 34 genera and 16 families were collected. Species richness declined rapidly with altitude; nearly 90% of the species were recorded between 250 and 1250 m a.s.l. Three of the four physico‐chemical variables, of the water, temperature, dissolved oxygen and pH, explained 53·5% of the variation in species richness along the altitudinal gradient, with temperature the most important (37·6%). An analysis of species composition showed that the distinctiveness of the fish fauna increased with elevation, with the greatest turnover observed between 1000 and 1750 m a.s.l. On this altitudinal gradient, turnover was dominated by the loss of species rather than gain, and dominance by just a few species was greater at higher elevations. Turnover was also observed along the altitudinal gradient in the structure of the three functional groups (torrential, pool and pelagic species). The study focused on understanding the pattern of diversity of fish communities inhabiting the Andes in Colombia. Anthropogenic effects on the altitudinal distribution of fish species in the region, however, are largely unknown and would require further investigations.     

Los pumas (Puma concolor) como agentes tafonómicos. Análisis actualístico de un sitio de matanza en los Andes de Mendoza, Argentina

A taphonomic study of the carcass of an adult guanaco (Lama guanicoe) recently killed and consumed by one or more pumas (Puma concolor) in the Laguna del Diamante reserve (Mendoza province, Argentina) is exposed, and the case is discussed in the light of the available taphonomic information on this agent in America. The carcass was dispersed along more than 30 m and basically devoid of soft tissues. Thirty-four percent of the bones presented tooth modifications, removal of bone tissue and edge damage, mostly attributable to pumas. Some of these modifications are considerable, as in the case of the skull. The results of the study suggest that bone modifications are intense as compared to other taphonomic studies on this felid. The implications of this are discussed, and the conclusion is reached that more knowledge is needed on the range of variation in puma taphonomic action and the conditions under which such variation occurs.     

Biomass allocation in herbaceous plants under grazing impact in the high semi-arid Andes

Among the plant traits that affect performance, vitality and herbivore resistance in rangeland vegetation, biomass partitioning ranks top, commonly outweighing processes at single leaves (e.g., photosynthesis, respiration). We explored the allometry of a broad sample of herbaceous species from two high elevation sites in the Andes in order to explain biomass partitioning under harsh climatic conditions and risk of biomass losses under strong camelid grazing pressure. We combined data from NW-Argentina and W-Bolivia from elevations between 4200 and 4250 m in a landscape dominated by tall Festuca orthophylla tussocks and a drought driven seasonality (rainfall only between November and March). Across 10–20 taxa per region we found less investment in leaves with a mean leaf mass fraction of only 11% and instead a massive storage in below-ground compartments (rhizomes, tap roots), particularly at the colder Argentinean site. Though grazing pressure was much greater in Bolivia, the foliage mass fraction was there larger than in Argentina. The inter-tussock space in these open, dry plains (‘pajonal’) was dominated by rosette forming species with a below-ground shoot apex and massive tap roots (70% of all species), rendering these species less sensitive to grazing and trampling. The storage organs of these species represented more than 50% of total biomass. Llamas, which represented the main vertebrate herbivore in these open plains at the Bolivian site, preferred non tap-root herbs and species with low leaf nitrogen concentration. Palatable forbs for llamas (22% of all species at the Bolivian site) only survived when nested (facilitated) in the rigid, tall Festuca orthophylla tussocks or thorny shrubs. In conclusion, these extremely high elevation rangeland herbs invest preferentially in structures for persistence (K-strategy) rather than maximizing carbon gain.     

Endemic Melastomataceae of the Sierra Nevada de Santa Marta,Colombia

The 16 species of Melastomataceae restricted to the middle and upper slopes of the Sierra Nevada de Santa Marta, Colombia, are listed. Included are seven previously undescribed species:Graffenrieda santamartensis, Miconia insueta, Miconia oreogena, Miconia tricaudata, Huilaea kirkbridei, Kirkbridea tetramera, Kirkbridea pentamera. The newly described genusKirkbridea is placed in the tribe Miconieae.     

Pronophiline butterflies (Satyridae) of the Cordillera de Mérida, Venezuela

The 36 species of the tribe Pronophilini known from the Cordillera de Merida, the main Andean mountain range in Venezuela, are listed, together with three others probably erroneously described or labelled as from the range. The general ecological characteristics of the tribe are described, using the Merida species as examples. These include its restriction to high altitudes, endemism in mountain 'islands', species partitioning up the altitude gradient, mimicry within the tribe, and seasonality. The evolutionary relationships of the 16 endemic species are analyzed. Two new genera, six new species, two new subspecies and one new form are described. Three new synonymies are established, three new combinations made, new status is given to two taxa and the status of one taxon is revised.