title: Reproductive phenology of 233 species from four herbaceous-shrubby communities in the Gran Sabana Plateau of Venezuela.
abstract: OBJECTIVE
  Herbaceous-shrubby communities in the Gran Sabana (Great Savanna) Plateau of Venezuela grow under non-zonal conditions. We speculated that this would produce specific patterns of reproductive phenology within these different soil-climate-vegetation associations. Specifically, we tested the hypothesis that the reproductive phenology patterns of four herbaceous-shrubby communities are determined by climate, plant life-forms and soil properties.
  
  
  METHODS
  The reproductive phenology of 233 plant species of the Gran Sabana Plateau of the Venezuelan Guayana Highlands was studied taking into account their life-forms (i.e. trees, shrubs, climbers, annual herbs, perennial herbs, epiphytes and parasites/hemiparasites) in four herbaceous-shrubby communities: (i) shrubland, (ii) secondary bush, (iii) savanna and (iv) broad-leaved meadow. Patterns of flowering, and occurrence of unripe fruit and ripe fruit were studied at two levels of intensity for 24 months within a 5-year span. Two phenological records for each month of the year and between two and four replicates for each community type were made. Randomly selected 2-3 ha plots were used. General phenological patterns were established using <25% of the plants of each species in each plot to give the total duration of each phenological phase. High-intensity phenological patterns were established using >25% of individuals in each plot to establish times of high abundance of flowers, and presence of unripe fruit and/or ripe fruit on individual plants. This generated phenological peaks for each species.
  
  
  RESULTS
  Non-seasonality of general flowering and unripe fruiting in each of the four communities was related to non-seasonal flowering and unripe fruiting patterns in the plant life-forms studied and to low variation in precipitation throughout the year. Flowering activity in the shrubland and broad-leaved meadow peaked twice. The bush community had only one flowering peak while the savanna gave a non-seasonal flowering peak. The peak unripe fruiting pattern was not clearly related to unripe fruit phenological patterns of the most abundant life-forms. Unripe fruit patterns and precipitation were only correlated for shrubs, climbers and trees in the shrubland. Ripe fruiting patterns peaked during the short-dry season in the bush and shrubland, and were negatively correlated with precipitation in the shrubland. General and peak ripe fruiting patterns were non-seasonal in the savanna and broad-leaved meadow and related to the dominance of herbaceous species with prolonged ripe fruiting times, low climate seasonality, high plant species richness and diversity, and dispersal syndromes.
  
  
  CONCLUSIONS
  The reproductive phenology of the herbaceous-shrubby communities is mainly influenced by the composition of the life-forms, the precipitation regime and soil type.
authors.affiliation: Facultad de Ciencias ,  Instituto de Biología Experimental ,  Centro Botánica Tropical ,  Universidad Central de Venezuela ,  Aptdo. 48312, Caracas 1041A ,  Venezuela.
authors.forename: Nelson
authors.initials: N
authors.lastname: Ramírez
authors.forename: Herbert
authors.initials: H
authors.lastname: Briceño