Algal growth and species composition under experimental control of herbivory, phosphorus and coral abundance in Glovers Reef, Belize

Abstract

The proliferation of algae on disturbed coral reefs has often been attributed to (1) a loss of large-bodied herbivorous fishes, (2) increases in sea water nutrient concentrations, particularly phosphorus, and (3) a loss of hard coral cover or a combination of these and other factors. We performed replicated small-scale caging experiments in the offshore lagoon of Glovers Reef atoll, Belize where three treatments had closed-top (no large-bodied herbivores) and one treatment had open-top cages (grazing by large-bodied herbivores). Closed-top treatments simulated a reduced-herbivory situation, excluding large fishes but including small herbivorous fishes such as damselfishes and small parrotfishes. Treatments in the closed-top cages included the addition of high phosphorus fertilizer, live branches of Acropora cervicornis and a third unmanipulated control treatment. Colonization, algal biomass and species composition on dead A. palmata "plates" were studied weekly for 50 days in each of the four treatments. Fertilization doubled the concentration of phosphorus from 0.35 to 0.77 microM. Closed-top cages, particularly the fertilizer and A. cervicornis additions, attracted more small-bodied parrotfish and damselfish than the open-top cages such that there was moderate levels of herbivory in closed-top cages. The open-top cages did, however, have a higher abundance of the chemically and morphologically defended erect algal species including Caulerpa cupressoides, Laurencia obtusa, Dictyota menstrualis and Lobophora variegata. The most herbivore-resistant calcareous green algae (i.e. Halimeda) were, however, uncommon in all treatments. Algal biomass increased and fluctuated simultaneously in all treatments over time, but algal biomass, as measured by wet, dry and decalcified weight, did not differ greatly between the treatments with only marginally higher biomass (p < 0.06) in the fertilized compared to open-top cages. Algal species composition was influenced by all treatments with a maximum between-treatment Bray-Curtis similarity of only 29%. The fertilized cages showed rapid colonization by a mixed turf community largely composed of the filamentous brown (Hincksia mitchelliae) and green (Enteromorpha prolifera) species. Algal cover in the fertilized cages leveled at 80% after 20 days compared to less than 50% in the other treatments. There was no evidence that A. cervicornis suppressed algal colonization compared to the unmanipulated controls. Instead, the herbivore susceptible Padina sanctae-crucis was the most abundant algae followed by Jania capillacea in this treatment in contrast to the more chemically defended Dictyota menstrualis that dominated the unmanipulated controls. We conclude that A. cervicornis was not suppressing algae as a group and its loss cannot account for the observed changes in algal abundance in most reefs except for creating space. In contrast, A. cervicornis appears to attract aggressive damselfish that may reduce herbivory by larger herbivores. Phosphorus enrichment can lead to rapid colonization of space by filamentous turf communities but not high biomass and dominance of erect frondose algae within 50 days. Moderate levels of herbivory by large-bodied herbivores promoted moderately herbivore-resistant erect brown and green algae that are commonly reported on disturbed reefs. Consequently, all the studied factors influenced algal communities but seldom as commonly predicted.