Marine Biology

, 165:35 | Cite as

Hidden in plain sight: species richness and habitat characterisation of sublittoral pebble beds

  • Julian Evans
  • Martin J. Attrill
  • Joseph A. Borg
  • Peter A. Cotton
  • Patrick J. Schembri
Original paper


Sublittoral pebble beds are generally considered to be impoverished, but the physical and biological characteristics of these habitats are poorly known. We characterised nineteen pebble bed sites in the Maltese Islands, providing detailed habitat data for Mediterranean sublittoral pebble beds for the first time. Nearly 40,000 individuals belonging to 332 taxa were recorded in all, with total richness estimated to reach 440 taxa; molluscs, crustaceans and polychaetes were the most diverse faunal groups. This high diversity is likely due to the structural complexity of the pebble beds, which had a vertically stratified arrangement of sediment particles that is likely maintained through periodic physical disturbance. Variation in the biotic assemblage from site to site was correlated with changes in the quantity of sand and silt, with the area of the pebble bed, with water depth, and with the thickness of the pebble layer. This indicates that pebble-bed macrofaunal assemblages are sensitive to changes in hydrodynamic conditions and sediment loading, to alterations to the stratification of the pebble beds, and to fragmentation of the habitat patches. These results contradict assertions that sublittoral pebble beds are impoverished, instead showing that they can be highly diverse habitats supporting biotic assemblages that respond to a complex set of environmental variables. The present findings enable better understanding of the ecological importance of pebble beds and of the potential impacts of anthropogenic disturbance, enabling more informed decisions for habitat conservation and management.



We are grateful to K. Liversage and an anonymous referee for their comments on a previous version of this paper.

Compliance with ethical standards


The present work was partly funded by the European Union-European Social Fund through a Strategic Educational Pathways Scholarship (Malta) Grant awarded to JE.

Conflict of interest

The authors have no conflict of interest.

Ethical approval

This article does not contain any studies with animals performed by any of the authors.


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Copyright information

© Springer-Verlag GmbH Germany, part of Springer Nature 2018

Authors and Affiliations

  1. 1.Department of BiologyUniversity of MaltaMsidaMalta
  2. 2.Marine Biology and Ecology Research CentrePlymouth UniversityPlymouthUK

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