Development in an estuarine fouling community: The influence of early colonists on later arrivals.

Experiments were performed to determine if earlier colonists inhibited, enhanced, or were necessary for establishment of later colonists during development of an estuarine fouling community at Lewes, Delaware. We determined the significance of earlier stages on the successional process by functionally removing early colonizing species. Since settlement of sessile invertebrates onto our experimental test plates was seasonal, we were able to accomplish functional removal of early colonists by putting out clean test panels after these species had ceased settling. Comparisons between panels initially submerged at three different times in 1974 and 1975, and between panels put out at one-month intervals throughout the study (to describe seasonal settlement patterns) allowed us to determine interactions between adult populations of earlier colonists and colonizing individuals of later arriving species.The dominant sessile species in our system and their times of settlement were: a barnacle (Balanus improvisus) - April through June, a polychaete (Hydroides dianthus) - July and August, a tunicate (Molgula manhatensis) - June through October, a hydroid (Tubularia crocea) - July through October, and a mussel (Mytilus edulis) - November through April. All successional series eventually came to be dominated by M. edulis, and it persisted as the dominant for over a year.A variety of species interactions were observed. M. edulis inhibited colonization by all other dominants and B. improvisus partially inhibited settlement of M. manhattensis. The presence of adult M. manhattensis had no influence on summer settlement of T. crocea, but the hydroids enhanced settlement of tunicates in the fall. During both years of our study, larger settlements of mussels were noted on panels harboring tunicates and hydroids than on bare surfaces. H. dianthus, on the other hand, became established only on bare substrates, and colonization was almost totally inhibited by other dominants.Development in our fouling community did not conform to any single model of community development presented to date. Instead, components of several models were observed within our relatively simple (in terms of number of species) system. For example, facilitation (enhancement of later colonists by earlier ones) and inhibition (resistance of earlier colonists to invasion by later colonists) were both observed. However, we found no evidence earlier colonists were essential for establishment of the next developmental stage. In fact, inhibitory interactions appeared to be much more prevalent than facilitative interactions. The former may also have more profound effects on community development since they more often determine eventual species compositions.