Colonization by barnacles on fossil Clypeaster : an exceptional example of larval settlement

Abstract

The presence of c. 1450 individuals of the balanid barnacle Balanus crenatus Bruguiere encrusting the test of a clypeasteroid sea urchin from the Late Miocene of the Guadalquivir Basin (southwestern Spain) allows proposing a settlement pattern linked to the growth of the encrusting organism. The possible influence of dip angle was controlled by dividing the test into four concentric zones ranging from lowest margin to apex (0–15°, 15–30°, 30–50° and 0°). Contour diagrams were prepared to identify areas of highest barnacle density as well as size categories distribution in relationship to the pitch of the sea urchin test. The orientation of balanid tests was recorded and plotted on rose diagrams from 0° to 180°. Four size categories of barnacles were distinguished: (1) < 1 mm, (2) 1–2 mm, (3) 2–3 mm and (4) 3–4 mm; these correspond to a growth sequence ranging from post-larval forms to juveniles. Two areas of maximum settlement density are situated on the posterior margin of the test, on aboral as well as oral surfaces. The aboral surface shows the maximum number of barnacles. Two groups of individuals are defined on the basis of their location, that is, those encrusting the posterior medium part of the urchin, and those located on the anterior half. The results suggest that larval settlement was initially controlled by the availability of free space and afterwards by an intensification effect. Orientation and dip of the test may have played a secondary role in the settlement of the larvae. Substrate colonization seems to have been closely related to the biostratinomic history of the sea urchin test and although several scenarios are possible, our data are congruent with a synchronous settlement of both surfaces (aboral and oral) by one spat or several.