Environmental Correlates of Fish Distribution Pattern

Abstract  The underlying mechanisms responsible for ecological plasticity and consequent invasive character of non-native freshwater fish species, variations in growth and life history traits in gibel carp Carassius gibelio (Bloch) were compared in natural and artificial water bodies of Turkey. Females significantly outnumbered males in all natural and most artificial waters. Discriminate function analysis differentiated gibel populations into three separate groups (natural lakes, artificial water bodies and running waters), with significant differences among separated groups in growth index, standard length and age at maturity, relative fecundity and gonado-somatic index, but not in egg diameter and both generalised and relative condition. Growth features (e.g. growth index and relative condition) and reproductive features (e.g. relative and absolute fecundity) positively correlated with water body area. No correlations were found for any growth or life history trait with depth, latitude and altitude. With the exception of smaller size at maturity, all traits were higher in populations from artificial water bodies than those inhabiting running waters, suggesting gibel carp is required to exert more reproductive effort to invade natural ecosystems than artificial waters.

Background
Many postglacial lakes contain fish species with distinct ecomorphs. Similar evolutionary scenarios might be acting on evolutionarily young fish communities in lakes of remote islands. One process that drives diversification in island freshwater fish species is the colonization of depauperate freshwater environments by diadromous (migratory) taxa, which secondarily lose their migratory behaviour. The loss of migration limits dispersal and gene flow between distant populations, and, therefore, is expected to facilitate local morphological and genetic differentiation. To date, most studies have focused on interspecific relationships among migratory species and their non-migratory sister taxa. We hypothesize that the loss of migration facilitates intraspecific morphological, behavioural, and genetic differentiation between migratory and non-migratory populations of facultatively diadromous taxa, and, hence, incipient speciation of island freshwater fish species.

Results
Microchemical analyses of otolith isotopes (88Sr, 137Ba and 43Ca) differentiated migratory and non-migratory stocks of the New Zealand endemic Gobiomorphus cotidianus McDowall (Eleotridae). Samples were taken from two rivers, one lake and two geographically-separated outgroup locations. Meristic analyses of oculoscapular lateral line canals documented a gradual reduction of these structures in the non-migratory populations. Amplified fragment length polymorphism (AFLP) fingerprints revealed considerable genetic isolation between migratory and non-migratory populations. Temporal differences in reproductive timing (migratory = winter spawners, non-migratory = summer spawners; as inferred from gonadosomatic indices) provide a prezygotic reproductive isolation mechanism between the two ecotypes.

Conclusion
This study provides a holistic look at the role of diadromy in incipient speciation of island freshwater fish species. All four analytical approaches (otolith microchemistry, morphology, spawning timing, population genetics) yield congruent results, and provide clear and independent evidence for the existence of distinct migratory and non-migratory ecotypes within a river in a geographically confined range. The morphological changes within the non-migratory populations parallel interspecific patterns observed in all non-migratory New Zealand endemic Gobiomorphus species and other derived gobiid taxa, a pattern suggesting parallel evolution. This study indicates, for the first time, that distinct ecotypes of island freshwater fish species may be formed as a consequence of loss of migration and subsequent diversification. Therefore, if reproductive isolation persists, these processes may provide a mechanism to facilitate speciation.