Colonization and demographic expansion of freshwater fauna across the Hawaiian archipelago.

It is widely accepted that insular terrestrial biodiversity progresses with island age because colonization and diversification proceed over time. Here, we assessed whether this principle extends to oceanic island streams. We examined rangewide mtDNA sequence variation in four stream-dwelling species across the Hawaiian archipelago to characterize the relationship between colonization and demographic expansion, and to determine whether either factor reflects island age. We found that colonization and demographic expansion are not related and that neither corresponds to island age. The snail Neritina granosa exhibited the oldest colonization time (~2.713 mya) and time since demographic expansion (~282 kya), likely reflecting a preference for lotic habitats most prevalent on young islands. Conversely, gobioid fishes (Awaous stamineus, Eleotris sandwicensis and Sicyopterus stimpsoni) colonized the archipelago only ~0.411-0.935 mya, suggesting ecological opportunities for colonization in this group were temporally constrained. These findings indicate that stream communities form across colonization windows, underscoring the importance of ecological opportunities in shaping island freshwater diversity.

Local adaptation despite high gene flow in the waterfall-climbing Hawaiian goby, Sicyopterus stimpsoni.

Environmental heterogeneity can promote the emergence of locally adapted phenotypes among subpopulations of a species, whereas gene flow can result in phenotypic and genotypic homogenization. For organisms like amphidromous fishes that change habitats during their life history, the balance between selection and migration can shift through ontogeny, making the likelihood of local adaptation difficult to predict. In Hawaiian waterfall-climbing gobies, it has been hypothesized that larval mixing during oceanic dispersal counters local adaptation to contrasting topographic features of streams, like slope gradient, that can select for predator avoidance or climbing ability in juvenile recruits. To test this hypothesis, we used morphological traits and neutral genetic markers to compare phenotypic and genotypic distributions in recruiting juveniles and adult subpopulations of the waterfall-climbing amphidromous goby, Sicyopterus stimpsoni, from the islands of Hawai'i and Kaua'i. We found that body shape is significantly different between adult subpopulations from streams with contrasting slopes and that trait divergence in recruiting juveniles tracked stream topography more so than morphological measures of adult subpopulation differentiation. Although no evidence of population genetic differentiation was observed among adult subpopulations, we observed low but significant levels of spatially and temporally variable genetic differentiation among juvenile cohorts, which correlated with morphological divergence. Such a pattern of genetic differentiation is consistent with chaotic genetic patchiness arising from variable sources of recruits to different streams. Thus, at least in S. stimpsoni, the combination of variation in settlement cohorts in space and time coupled with strong postsettlement selection on juveniles as they migrate upstream to adult habitats provides the opportunity for morphological adaptation to local stream environments despite high gene flow.

Finding paradise: cues directing the migration of the waterfall climbing Hawaiian gobioid Sicyopterus stimpsoni.

A series of waterfall-climbing trials were conducted to identify cues that direct the climbing of juvenile Sicyopterus stimpsoni. In the first experiment, whether climbing juveniles preferentially ascend water sources with conspecifics or whether the presence of just stream water is sufficient to attract fish to ascend a climbing path were assessed. In the second experiment, whether climbing juveniles create a trail of mucus that facilitates the ability of conspecifics to follow their lead was determined. The results indicate that juvenile S. stimpsoni are less likely to climb in waters devoid of organic cues but are strongly attracted to stream water with or without the odour of conspecifics. Once climbing, performance did not differ for juveniles climbing in differing water choices, suggesting an all-or-nothing commitment once climbing commences. Climbing S. stimpsoni did produce a mucous trail while climbing that was associated with a mucous gland that dramatically increases in size just prior to juveniles gaining the ability to climb. The trail was not followed closely by subsequent juveniles traversing the same channel, however, suggesting only weak trail-following in waterfall climbing S. stimpsoni. Previous genetic studies suggest that juvenile S. stimpsoni do not home to natal streams in the face of strong near-shore oceanic currents. Instead, these fish appear primarily to rely on cues that suggest the presence of organic growth in streams, a factor that may indicate suitable habitat in an ever-changing stream environment but which may also be vulnerable to interference through human activity.