APPENDIX 
GENERAL BEACH ECOLOGY 
The pioneer work on sandy beaches is Btologiske Studier Over Sand-strands- 
faunen, Saerlig Ved De Danske Kyster by Mortensen in 1923,as cited by Dahl 
(1953). Mortensen is credited by Dahl with first posing the main problems in 
sandy beach ecology. The earliest study in this country of species assemblages 
on sandy beaches was conducted at Beaufort, North Carolina (Pearse, Humm, and 
Wharton, 1942). This Beaufort study also constituted the first description of 
the fauna at Fort Macon. Although it was not a quantitative study, it did 
note the intertidal zonation and general abundance of £. Talpotda and Donax 
spp. 
There are few other studies of high-energy sandy beaches. Leber (1977) 
studied the community ecology of the beach at Emerald Isle. This same beach 
was used as a comparison beach in the present study. He monitored the inter- 
tidal macrofauna for 14 months and established a qualitative energy flow model 
for the macrofaunal community. Matta (1977) performed a quantitative survey 
of the organisms located on transects from the swash zone to a distance of 
60 meters offshore at Duck, North Carolina. 
Other studies of sandy beach macrofaunal communities have been of beaches 
exposed to lower wave energies. Dexter (1967) reported the distribution and 
niche diversity of haustoriid amphipods; Dexter (1969) reported the faunistics 
of the sandy beach community of Radio Island at Beaufort, North Carolina. In 
addition she compared the fauna of Pacific and Atlantic sandy beaches of 
Panama (Dexter, 1972), Costa Rica and Columbia (Dexter, 1974), and described 
the sandy beach fauna of Mexico (Dexter, 1976). Croaker (1967) studied the 
amphipods of sandy beaches in Georgia. Pamatmat (1968) carried out a quanti- 
tative faunistic study of the beaches in Washington. Grant (1965) studied the 
fauna of protected beaches in Massachusetts. Whithers (1977) compared the 
faunal communities of a series of sand and sand-mud beaches in Wales, and 
McLachlan (1977a, 1977b) described both macrofaunal and meiofaunal communities 
on a series of protected beaches in South Africa. Saloman (1976) reported the 
effects of a hurricane on the beach and the nearshore beach fauna of Panama 
City Beach, Florida. Holland and Polgar (1976) studied the ecology of a high- 
energy intertidal sandflat in Georgia. All these investigators concluded that 
sandy beach macrofaunal communities generally are: (1) low in measures of 
community structure such as species diversity (Shannon and Weaver, 1963), 
species richness (Pielou, 1969), and species evenness (Pielou, 1966); and (2) 
dominated by one or two species which are particularly well adapted to the 
environmental stresses of the habitat in which they live (Sameoto, 1969). This 
type of low diversity system is thought by some (Holland and Polgar, 1976; 
Leber, 1977) to constitute a classic example of a physically controlled com- 
munity as defined by Sanders (1968). The physically controlled sandy beach is 
subjected to large fluctuations in various abiotic ecofactors including temper- 
ature, salinity, oxygen availability, redox potential, water movement, and pH 
(Hartwig, Gluth, and Wieser, 1977). 
In general, ecological communities are composed of primary producers, first- 
order consumers (sometimes called secondary producers), predators (sometimes 
called second-order consumers), and decomposers. The high-energy sandy beach 
is an exception, however, in that it lacks any important primary production. 
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