A remarkable sand-dwelling fish assemblage from central Amazonia, with comments on the evolution of psammophily in South American freshwater fishes

Abstract

We studied a specialised assemblage of sand-dwelling fish species from a streamlet in central Amazonia. The five sand-dwelling species comprised 15.2 % of the total number in the streamlet (33 species). Two of the sand-dwellers are daytime foragers, Characidium cf. pteroides (Crenuchidae) and Stauroglanis gouldingi (Trichomycteridae), whereas three ones are night-time foragers, Gymnorhamphichthys rondoni (Rhamphichthyidae), "Imparfinis" pristos and Mastiglanis asopos (Heptapteridae). With the exception of C. cf. pteroides, the remainder species bury in the sand during their resting period. All five species bear a cryptic, sand-colour pattern, and some are translucent, traits which we regard as advantageous both for evasion from predators and for approaching prey (for the daytime foragers). All five species are microphagous carnivores and their foraging tactics range from generalised sit-and-wait (C. cf. pteroides) to active searching for interstitial prey (G. rondoni). A unique "drift-trap" system made up by the extended barbels and filamentous first pectoral-fin rays is employed by M. asopos. We regard the small size and low number of vertebrae (which is functionally expressed by fast displacements in the sand) as additional features advantageous to explore the sand habitat, allowing diving quickly in the substrate to evade predators and to forage for small prey in sand interstices or plant debris. The relationship between morphological and behavioural characters suited to life in sandy substrates is examined under the perspective of the current phylogenies for each group of psammophilous fishes here studied. The mapping of these characters demonstrates that not all of them evolved specifically for strict psammophily. Instead, some of them may represent the final step of a transformation series or synapomorphies of higher hierarchical levels. Several characters linked to psammophily, such as small body, large eyes, and scarce pigmentation are probably paedomorphic traits. We consider that the little variable specific composition of fish sand-dwelling assemblages and the wide (although patchy) geographical distribution of their species in the Amazon are due to the widespread occurrence and low complexity of this specialised habitat. The pre-existence of benthic habits, small size, and the broad distribution of rivers with sandy beds, mainly in the Amazon and Orinoco basins, possibly played a determinant role in the origin and diversification of psammophilous assemblages among freshwater fishes in South America. Copyright © 2006 Sociedade Brasileira de Ictiologia.