SETTLEMENT AND EARLY POST SETTLEMENT SURVIVAL OF SESSILE MARINE-INVERTEBRATES ON TOPOGRAPHICALLY COMPLEX-SURFACES - THE IMPORTANCE OF REFUGE DIMENSIONS AND ADULT MORPHOLOGY
Lj. Walters et Ds. Wethey, SETTLEMENT AND EARLY POST SETTLEMENT SURVIVAL OF SESSILE MARINE-INVERTEBRATES ON TOPOGRAPHICALLY COMPLEX-SURFACES - THE IMPORTANCE OF REFUGE DIMENSIONS AND ADULT MORPHOLOGY, Marine ecology. Progress series, 137(1-3), 1996, pp. 161-171
We predicted that both refuge dimension and growth form would influenc
e settlement and short-term post-settlement success (less than or equa
l to 7 d) of sessile marine invertebrates that live attached to hard s
ubstrata in low energy environments. Individuals with unlimited attach
ment to the substrata should rapidly be protected by their growth form
, thus decreasing their need to settle in refuges and limiting the len
gth of time any locations on heterogeneous substrata act as refuges. A
lternatively, organisms with limited attachment to the substrata shoul
d remain susceptible to the causes of mortality for a longer time, and
as a result should settle in high quality refuges(sites that protect
individuals from competitors, predators or physical disturbance events
until either a size refuge or reproductive maturity is obtained). Res
ults agreed with these predictions for 4 species of invertebrates exam
ined on both the topographically complex surface of the solitary ascid
ian Styela plicata (hereafter Styela) and on settlement plates with un
iformly spaced roughness elements that mimicked the heights of roughne
ss elements (2.0 and 5.0 mm) found on Styela in Beaufort, North Caroli
na, USA. On all surfaces, the 2 species with limited attachment to the
substrata, Balanus sp. (aclonal, solitary) and Bugula neritina (clona
l, arborescent), settled almost exclusively in the location that provi
ded individuals with the best refuge: the crevices formed where the ba
ses of roughness elements intersect with the flat surfaces. Additional
ly, when roughness elements of various heights were present (Styela, r
ange: 0.6 to 8.8 mm), intermediate size roughness elements (2.0 < x le
ss than or equal to 5.0 mm) were picked over 72% of the time. Settleme
nt locations and locations where survival were enhanced were less cons
istent for the 2 species with unlimited attachment to the substrata: a
clonal, encrusting form (Schizoporella errata) and a clonal stolen-ma
t form (Tubularia crocea). Fewer individuals of these 2 species settle
d on roughness elements on Styela and when they did, they were not res
tricted to the bases of the roughness elements. On the plate surfaces,
most settlement did occur in crevices, but both species grew away fro
m this location within days and short-term survival was not consistent
ly greater in this location. Additional trials were run on plates with
pits of the same maximum dimensions as the tested roughness elements
(2.0 and 5.0 mm depth) to see if crevices and pits provide refuges of
equal quality for newly settled individuals. Only survival of Balanus
sp. recruits was greatest in both crevices and pits. Evidence for acti
ve choice of settlement location comes from consistent results in tria
ls in which some larvae settled in greater numbers on specific size ro
ughness elements on Styela and in areas of high erosion. Overall, thes
e results show that one must be very cautious when generalizing about
refuge quality on heterogeneous surfaces, and to determine if a locati
on is a spatial refuge, it is critical to consider: (1) the dimensions
oi the larva, (2) the relative dimensions of the individual and poten
tial refuge:location at any point in time from the moment settlement o
ccurred, and (3) the growth form of the individual which is related to
its need for protection from biotic and abiotic sources of mortality.