Distributional relationships among subtidal algae, sea urchins and reef fish in northeastern New Zealand

Abstract

Interactions among large brown macroalgae, sea urchins, and fishes were investigated in northeastern New Zealand during the period 1988 - 1993. The Cape Rodney to Okakari Point Marine Reserve was the site of many of these investigations. The patterns of abundance of large brown macroalgae and urchins down depth gradients over a wide geographic range were compared with those reported from earlier studies, and 3 major trends were identified. First, the fucoid alga Carpophyllum flexuosum now occurs at many sites which are exposed to wave action, in contrast to earlier studies. This alga occurred most abundantly on urchin-grazed coralline flat areas. Second, at four sites in the Marine Reserve, the densities of the echinometrid urchin Evechinus chloroticus decreased with increasing depth, rather than reaching maximal densities at mid-depths, as had previously been described. Finally, at sites of decreased exposure to wave action, the coralline flats habitat did not occur at all, and dense stands of. C. flexuosum occurred, in conjunction with the ubiquitous laminarian alga, Ecklonia radiata. Following the discovery of this new algal component to exposed rocky reefs, a more detailed study of its population characteristics was initiated. The population size structure of C. flexuosum on coralline flat areas was markedly different from that of the same species in sites sheltered from wave action. These differences occurred at both offshore islands and sites near Leigh, suggesting that it was a general pattern. C. flexuosum plants on coralline flats were smaller than those from sheltered sites, and had a greater number of smaller laminae, heavier stipes, and a greater degree of branching. There was some evidence of temporal change in the morphology of C. flexuosum from coralline flats. Comparisons between a site with C. flexuosum and a site with coralline flats suggested that the activity of fish was 75% lower in the vegetated than in the unvegetated site, and the feeding rate in the vegetated site was less than 50% that in the unvegetated site. I speculate that future effects on fish activity of the invasion of C. flexuosum into a habitat which previously lacked macroalgal vegetation may depend on changes in the morphology of plants. An investigation of many aspects of the biology of E. chloroticus in different habitats was undertaken. Analysis of the body dimensions of E. chloroticus suggested that this species was relatively tall compared to other echinometrids (average ratio of test height: test diameter = 0.54), Comparisons among habitats with differing amounts of vegetation revealed only small differences in the relationship between test diameter and test height. Small E. chloroticus (<40 mm test diameter) lived in crevices, while larger individuals grazed freely over the substratum. In vegetated habitats, the crevice-dwelling habit was maintained at test diameters about l0 mm greater than in unvegetated habitats. Very small (<20 mm test diameter) E. chloroticus frequently covered themselves with shell. Population size structures of E. chloroticus within the Cape Rodney to Okakari Point Marine Reserve were bimodal; other localities had unimodal populations. Modal sizes varied among localities, with smallest modes (50-60 mm TD) being found at Inner Hauraki Gulf sites, and largest modes at the offshore Mokohinau Islands (70-80 mm TD). Habitat did not predictably affect population size structure. A bimodal population structure was maintained at Waterfall Reef rock flats throughout the 5-year study period. Gonad size showed seasonal fluctuations at several sites, being greatest in summer. There were few consistent differences in gonad size between biological habitats. Gonad colour varied among sites and habitats, with orange gonads generally being more prevalent in vegetated habitats, and black gonads being represented more in unvegetated habitats. Smaller urchins had greater proportions of orange gonads, while larger urchins had greater proportions of brown and black gonads. Although highly variable among individual urchins, movement of E. chloroticus was greater at unvegetated sites (0.7 m per 5 days) than at vegetated sites (0.4 m per 5 days), in the Marine Reserve. Feeding of E. chloroticus was studied at a number of sites in the Marine Reserve. Urchins frequently consumed drift algae, particularly E. radiata. C. flexuosum was consumed at less than half the rate of other macroalgae in several laboratory feeding experiments, and was chosen least frequently in a field assay of feeding preferences among 8 species of macroalgae. Boosting densities of E. chloroticus in stands of E. radiata to 60 m-2 led to destructive grazing of plants over a 2 month period - at lower densities, the urchins dispersed. Densities of C. flexuosum were effectively unchanged when urchin densities were increased to these elevated levels. As a result of these observations I speculate that feeding preferences of E. chloroticus may have a role in allowing C. flexuosum to survive on coralline flats. In a laboratory experiment, urchins from a feeding aggregation did not graze algae at higher rates than individuals from outside aggregations. Diets of both E. radiata and C. flexuosum consistently produced similar gonad volumes in urchins held in the laboratory, although gonad volumes produced were low. A preliminary experiment suggested that C. flexuosum from exposed sites was consumed at lower rates than C. flexuosum from sites which were sheltered from wave action. These differences in palatability are mirrored in the formation of stable borders between coralline flats and C. flexuosum of the sheltered morphology, and the ability of C. flexuosum of the exposed morphology to survive in the coralline flats habitat. The fish fauna of the Cape Rodney to Okakari Point Marine Reserve was shown to be different from that of a nearby area. A number of species were more abundant within the Marine Reserve. Subsequent surveys showed that there were differences in abundances of 3 large carnivorous fishes among sites within the Marine Reserve, and that population size structure and the distance within which divers could approach one species, (Pagrus auratus), clearly varied between areas within the Marine Reserve. Mean standard length of P. auratus in the central marine reserve was 40% larger than that of P. auratus outside the central marine reserve, and the average minimum approach distance was 70% less in the central marine reserve. Feeding of fish by humans in the central part of the Marine Reserve was suggested to be the main cause of the differences in responses to divers. Population size structure of, and crevice occupancy by, E. chloroticus, clearly differed between the Marine Reserve and an adjacent area, with bimodal population size structures and a 10 mm greater size of crevice occupancy occurring in the Marine Reserve. The implications of these findings for extrapolating from experiments done in one area to other areas are discussed. The major biological components of rocky reef habitats identified in this study were broadly similar to those identified in previous studies in northeastern New Zealand, and have parallels in overseas studies. Long term changes to the flora of rocky reefs in northeastern New Zealand have occurred, and appear to persist by a mechanism which had previously been discounted. Similar processes to those observed in overseas studies appear to maintain habitats (consistent recruitment of algae or urchins to habitats which they dominate), or cause them to change from one habitat state to another (e.g. grazing outbreaks by urchins). However, the predictability of the persistence of these habitats at a particular site appears to be low. Further, the precise mechanisms whereby habitats may change from one to another may also be unpredictable. I argue that there is little scope for general statements concerning the spatial and temporal occurrence, or mode, of habitat transitions on temperate subtidal reefs. This study emphasises the value of repeated descriptions of patterns of abundance, and highlights problems of extrapolation and generalisation in marine ecology. Insufficient information exists at present to comment adequately on the persistence of subtidal habitat types. This may in part stem from the types of information which have been collected in the past. Methodological problems with the use of quadrats to sample densities of organisms in areas of differing topography are therefore addressed. In conclusion, it is suggested that sampling protocols which incorporate a variety of information, gathered over as wide an area, and as intensively as possible, should be used in future research of this type.