LEAF DEMOGRAPHY OF A NEOTROPICAL RAIN-FOREST CYCAD, ZAMIA-SKINNERI (ZAMIACEAE)

Tropical rain forest understories are extremely light-limited environments. Little is known about the demography of leaves in these habitats. To determine leaf longevity and causes of mortality for a species in the understory of a neotropical rain forest, we studied 634 leaves on 173 individuals of the cycad shrub Zamia skinneri for 10 yr at the La Selva Biological Station, Costa Rica. Five different seasonal cohorts of leaves were followed. No significant differences among cohorts were found in median leaf longevity, but the cohorts did differ in the proportion of leaves dying in annual intervals from < 1 to greater-than-or-equal-to 6 yr. Median minimum time alive for all leaves was 4.6 yr. Epiphyll coverage increased from 0% in the first year of life to 75%-100% by year 4. Leaf mortality was well described by two separate exponential functions, approximately 10% mortality/yr until year 4, then abruptly increasing to ca. 51%/yr. Of 625 leaf deaths, 77.0% were due to unexplained causes, 16.5% to physical damage by falling litter, 4.8% to attacks by larvae of the lycaenid butterfly Eumaeus minyas, and 1.8% to other causes. Leaf longevity was positively correlated with leaf size. Individuals of Z. skinneri frequently survive more than a century in very dark understory conditions. We suggest that leaf longevity in these circumstances is determined by the inevitable deterioration in leaf carbon balance due to leaf aging and increasing epiphyll coverage, and by loss due to physical damage and attacks by the specialist herbivore.