Maintaining Biodiversity in Freshwater Ecosystems on Oceanic Islands of the Tropical Pacific

Duringthepastfiveyears,aresearchgroupfromtheHawai'iDivisionofAquaticRe-sourcesandtheLouisianaStaeUniversityMuseumofNaturalSciencehascollaboratedonaseriesofstudiesconcerningthebiologyandconservaionofstreamanimalsintheHawaiianIs-lands.Fromtheviewpointsofbothgeologyandbiology,theseislandsrepresentthenorth-ernmostextensonofPolynesia.StreamanimalsinHawai'ihaverelativesattheleveloffami-ly,genus,andoccasionallyevenspeciesnotonlyinPolynesiabutthroughoutMelanesiaandMicronesiaaswell.Forthisreason…     

Life History of the Hawaiian Fish Kuhlia sandvicensis as Inferred from Daily Growth Rings of Otoliths

Kuhlia sandvicensis, the aholehole, is a native Hawaiian fish found in both marine and freshwater habitats. In the lower reaches of streams, they are predators on stream fishes, invertebrates, and insects. Aholehole are an important food fish in the Hawaiian Islands and were often used by ancient Hawaiians in traditional ceremonies. Although aholehole are an important part of stream ecosystems and Hawaiian culture, little is known about their life history, specifically, whether a freshwater phase is obligatory. In this study, light microscopy and electron microprobe techniques were used to analyze otolith daily increments. The analysis estimated age of juveniles and provided information regarding salinity of a fish's habitat at specific points in its life. Sr/Ca profiles from otoliths of juvenile and adult fish from fresh and salt water indicated that this species' use of stream habitats is facultative. Unlike Hawaiian freshwater gobies and at least one other member of the Kuhliidae from the Western Pacific, there is no physiological requirement of fresh water at a specific point in the life cycle of K. sandvicensis. Future research will provide a greater understanding as to the importance of streams as nursery habitats for this species. The research is expected to bolster the argument for maintaining the stream-ocean corridor for access by amphidromous gobies and perhaps also for the aholehole.     

Habitat utilization by Hawaiian stream fishes with reference to community structure in oceanic island streams

Synopsis Much of the freshwater biota on high Pacific islands is derived from marine ancestors. Traces of this marine origin are seen in the amphidromous life history patterns of these species. Investigation of the habitat utilization of an assemblage of freshwater gobies in the Hawaiian Islands indicates some resource partitioning within this group. The three major stream species appear to exhibit distinct habitat preferences, which in conjunction with their longitudinal zonation in streams may have served to facilitate their colonization or co-existence.     

Stable Isotope Analysis of Amphidromous Hawaiian Gobies Suggests Their Larvae Spend a Substantial Period of Time in Freshwater River Plumes

Synopsis We employed stable isotope analysis (δ¹³C, δ¹⁵N) to evaluate the sources of nutrients used by amphidromous gobiid fishes (Lentipes concolor, Sicyopterus stimpsoni, Awaous guamensis) caught migrating into and living in Hakalau Stream, Hawaii. Although considerable variation amongst the stable isotope values of stream items was noted across all 4 years of our study, the relationships between the fishes were relatively constant. Stable isotope values of recruiting gobies were consistently closer to those of both inshore plankton and freshwater adults than those of offshore plankton, suggesting that the larvae of these species derive much of their nutrition from inshore environments influenced by fresh water. Small differences between the stable values of these species further suggested that their larvae come from different inshore locations. After entering fresh water all species appear to swim rapidly upstream without feeding. Finally, once well upstream, adult L. concolor and A. guamensis appear to assume an omnivorous diet while adult S. stimpsoni rely upon autochthonous production within streams. We propose that freshwater food webs play an integral yet complex role in the lives of both larval and adult amphidromous Hawaiian fishes.     

Amphidromy links a newly documented fish community of continental Australian streams, to oceanic islands of the west Pacific

Background

Indo-Pacific high island streams experience extreme hydrological variation, and are characterised by freshwater fish species with an amphidromous life history. Amphidromy is a likely adaptation for colonisation of island streams following stochastic events that lead to local extirpation. In the Wet Tropics of north-eastern Australia, steep coastal mountain streams share similar physical characteristics to island systems. These streams are poorly surveyed, but may provide suitable habitat for amphidromous species. However, due to their ephemeral nature, common non-diadromous freshwater species of continental Australia are unlikely to persist. Consequently, we hypothesise that coastal Wet Tropics streams are faunally more similar, to distant Pacific island communities, than to nearby faunas of large continental rivers.

Methods/Principal Findings

Surveys of coastal Wet Tropics streams recorded 26 species, 10 of which are first records for Australia, with three species undescribed. This fish community is unique in an Australian context in that it contains mostly amphidromous species, including sicydiine gobies of the genera Sicyopterus, Sicyopus, Smilosicyopus and Stiphodon. Species presence/absence data of coastal Wet Tropics streams were compared to both Wet Tropics river networks and Pacific island faunas. ANOSIM indicated the fish fauna of north-eastern Australian coastal streams were more similar to distant Pacific islands (R = 0.76), than to nearby continental rivers (R = 0.98).

Main Conclusions/Significance

Coastal Wet Tropics streams are faunally more similar to distant Pacific islands (79% of species shared), than to nearby continental fauna due to two factors. First, coastal Wet Tropics streams lack many non-diadromous freshwater fish which are common in nearby large rivers. Second, many amphidromous species found in coastal Wet Tropics streams and Indo-Pacific islands remain absent from large rivers of the Wet Tropics. The evolutionary and conservation significance of this newly discovered Australian fauna requires clarification in the context of the wider amphidromous fish community of the Pacific.     

Hawaiian biogeography and the islands' freshwater fish fauna

Aim This paper describes known patterns in the distributions and relationships of Hawaiian freshwater fishes, and compares these patterns with those exhibited by Hawaii's terrestrial biota. Location The study is based in Hawaii, and seeks patterns across the tropical and subtropical Indo‐west Pacific. Methods The study is based primarily on literature analysis. Results The Hawaiian freshwater fish fauna comprises five species of goby in five different genera (Gobiidae). Four species are Hawaiian endemics, the fifth shared with islands in the western tropical Pacific Ocean. All genera are represented widely across the Indo‐west Pacific. All five species are present on all of the major Hawaiian islands. All five species are amphidromous – their larval and early juvenile life being spent in the sea. Although there has been some local phyletic evolution to produce Hawaiian endemics, there has been no local radiation to produce single‐island endemics across the archipelago. Nor is there evidence for genetic structuring among populations in the various islands. Main conclusions In this regard, the freshwater fish fauna of Hawaii differs from the well‐known patterns of local evolution and radiation in Hawaiian Island terrestrial taxa. Amphidromy probably explains the biogeographical idiosyncrasies of the fish fauna – dispersal through the sea initially brought the fish species to Hawaii, and gene flow among populations, across the archipelago, has hitherto inhibited the evolution of local island endemics, apparently even retarding genetic structuring on individual islands.     

Insular Freshwater Fish Faunas of Micronesia: Patterns of Species Richness and Similarity

Freshwater fishes are an important but relatively little known component of the highly diverse fish fauna of Micronesia. Localities supporting communities of freshwater fishes include large high islands, with considerable habitat complexity, and smaller low islands, such as atolls and raised coral islands, with limited freshwater habitat. Both types of islands may support species with adult life history styles that are (a) amphidromous and catadromous, (b) euryhaline (often estuarine), or (c) marine species which enter freshwater from time to time. We compared patterns of species richness and similarity between Micronesian localities for amphidromous and catadromous, euryhaline and marine species (ACEM) pooled, and for amphidromous and catadromous species (ACFW). Species richness of both ACEM and ACFW fishes was greatest on larger high islands compared to smaller high and all low islands. Cluster analysis of similarity indices for ACEM species between localities revealed two faunal components: high islands and low islands. High islands were further partitioned into a Caroline Islands cluster and a separate Mariana Islands cluster. Cluster analysis of ACFW species was more complex. One cluster consisted of a low island and a small high island, both in the Carolines chain and with limited freshwater habitat. The second cluster was partitioned into high islands and low islands that reflected influences of both size and geographical location.     

Rapid acquisition of directional preferences by migratory juveniles of two amphidromous Hawaiian gobies, Awaous guamensis and Sicyopterus stimpsoni

The native freshwater fish fauna of the Hawaiian Islands is composed of four gobiid species and one eleotrid. All five species are amphidromous. They spawn in freshwater, go to sea as free embryos and, as juveniles (sometimes referred to as post-larvae), enter fresh water again. Juveniles of two of these species, Awaous guamensis and Sicyopterus stimpsoni, had been observed maintaining directed migratory activity in the absence of water flow during the early freshwater phase of their upstream migration. Our experiments demonstrate that these juveniles are strongly attracted to flowing water. But, when water flow stops the fish persist in migratory behaviour in the same compass direction even when the tank is rotated, controlling for local landmarks and chemical cues, or when the trap that was the original water source is moved. This indicates a rapid (10 min or less) learning of directional cues. Preliminary data indicates a similar ability in a third species, Lentipes concolor. Rapid acquisition of directional preferences may be critical during upstream migration, when these tiny fish traverse water falls and other barriers by climbing in intermittent water flows. We suggest that rapid directional learning may have evolved in gobies as part of their predator avoidance repertoire and been subsequently applied to migration in streams.     

Overcoming urban stream syndrome: Trophic flexibility confers resilience in a Hawaiian stream fish

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Ontogenetic change in novel functions: waterfall climbing in adult Hawaiian gobiid fishes

Juveniles from three species of Hawaiian gobiid fishes climb waterfalls as part of an amphidromous life cycle, allowing them to re-penetrate adult upstream habitats after being swept out to the ocean upon hatching. The importance of climbing for juvenile stream gobies is well established, but adult fish in upstream island habitats also face potential downstream displacement by periodic disturbances. Thus, retention of climbing ability could be advantageous for adult stream gobies. Climbing performance might be expected to decline among adults, however, due to the tendency for mass-specific muscular power production to decrease with body size, and a lack of positively allometric growth among structures like the pelvic sucker that support body weight against gravity. To evaluate changes in waterfall-climbing ability with body size in Hawaiian stream gobies, we compared climbing performance and kinematics between adults and juveniles from three species: Awaous guamensis , Sicyopterus stimpsoni and Lentipes concolor . For species in which juveniles climbed using 'powerbursts' of axial undulation, adult performance and kinematics showed marked changes: adult A. guamensis failed to climb, and adult L. concolor used multiple pectoral fin adductions to crutch up surfaces at slow speeds, rather than rapid powerbursts. Adult S. stimpsoni , like juveniles, still used oral and pelvic suckers to 'inch' up surfaces and climbed at speeds comparable to those of juveniles. However, unlike juveniles, adult S. stimpsoni also add pectoral fin crutching to every climbing cycle. Thus, although powerburst species appear to be particularly susceptible to size-related declines in waterfall-climbing performance, the addition of compensatory mechanisms prevents the loss of this novel function in some species.     

Mid-ocean isolation and the evolution of Hawaiian reef fishes

The Hawaiian fish fauna has close affinities with the fauna of the Indo-west Pacific from which it is derived, but is depauperate. It is characterized by a large number of endemic species (30% of inshore fishes), which are often the most abundant species in their families in Hawaii. Although there is evidence of local adaptation, there has been no radiation of species within the island chain, as occurs in the terrestrial biota of isolated islands. Three major factors have contributed to these trends: (1) the geographic isolation of the islands, and oceanographic features, especially current patterns; (2) the life history characteristics of the fishes, especially their dispersal capabilities; and (3) the extent of adaptive differentiation to environmental conditions after they reached Hawaii.     

Benthic Community Structure and Invertebrate Drift in a Pacific Island Stream,Kosrae, Micronesia1

Tropical Pacific island streams have poorly understood communities that deserve scientific attention. We examined benthic macroinvertebrates and fishes of the Inem River on Kosrae, Federated States of Micronesia. Larval chironomids, lepidopterans, odonates, and freshwater shrimps dominated the benthos and drift. Diel periodicity in drift was not evident. Nine fishes, two shrimps, and one snail species were identified. Kosrae's stream fauna appears even more depauperate than other Pacific high islands, possible due to its extreme isolation.     

Morphological selection and the evaluation of potential tradeoffs between escape from predators and the climbing of waterfalls in the Hawaiian stream goby Sicyopterus stimpsoni

Environmental pressures may vary over the geographic range of a species, exposing subpopulations to divergent functional demands. How does exposure to competing demands shape the morphology of species and influence the divergence of populations? We explored these questions by performing selection experiments on juveniles of the Hawaiian goby Sicyopterus stimpsoni, an amphidromous fish that exhibits morphological differences across portions of its geographic range where different environmental pressures predominate. Juvenile S. stimpsoni face two primary and potentially opposing selective pressures on body shape as they return from the ocean to freshwater streams on islands: (1) avoiding predators in the lower reaches of a stream; and (2) climbing waterfalls to reach the habitats occupied by adults. These pressures differ in importance across the Hawaiian Islands. On the youngest island, Hawai'i, waterfalls are close to shore, thereby minimizing exposure to predators and placing a premium on climbing performance. In contrast, on the oldest major island, Kaua'i, waterfalls have eroded further inland, lengthening the exposure of juveniles to predators before migrating juveniles begin climbing. Both juvenile and adult fish show differences in body shape between these islands that would be predicted to improve evasion of predators by fish from Kaua'i (e.g., taller bodies that improve thrust) and climbing performance for fish from Hawai'i (e.g., narrower bodies that reduce drag), matching the prevailing environmental demand on each island. To evaluate how competing selection pressures and functional tradeoffs contribute to the divergence in body shape observed in S. stimpsoni, we compared selection imposed on juvenile body shape by (1) predation by the native fish Eleotris sandwicensis versus (2) climbing an artificial waterfall (～100 body lengths). Some variables showed opposing patterns of selection that matched predictions: for example, survivors of predation had lower fineness ratios than did control fish (i.e., greater body depth for a given length), whereas successful climbers had higher fineness ratios (reducing drag) than did fish that failed. However, most morphological variables showed significant selection in only one treatment rather than opposing selection across both. Thus, functional tradeoffs between evasion of predators and minimizing drag during climbing might influence divergence in body shape across subpopulations, but even when selection is an important contributing mechanism, directly opposite patterns of selection across environmental demands are not required to generate morphological divergence.     

Distribution of freshwater macroinvertebrates in streams with dams and associated reservoirs on a subtropical oceanic island off southern Japan

The conservation of endemic fauna in freshwater ecosystems is a topical issue on small oceanic islands. Because these endemics have limited distributions, they are more vulnerable to extinction. This study is the first to clarify the distribution of freshwater macroinvertebrates including endemic and alien species in streams with dams and associated reservoirs on the Ogasawara Islands in the northwestern Pacific Ocean. In 2007, we conducted a field survey in streams and reservoirs of the Yatsuse River system and collected 22 taxonomic groups from 13 stations. Hierarchal cluster analysis and non-metric multi-dimensional scaling (NMDS) were performed for the presence/absence data of the macroinvertebrates, and the results indicated that (1) most of endemic species were present in inlet streams of dam reservoirs, (2) these endemic species were absent in the bottom sediments of the reservoirs because of oxygen depletion and (3) dams may be barriers to the migration of some species of amphidromous crustaceans. Because human modifications, such as dams and associated reservoirs, on a small oceanic island can rapidly result in fragmentation or loss of freshwater habitats of endemic species, the remaining habitat of these species, such as headwater streams, must be protected and preserved to avoid species extinction.     

The Use of Two Modified Breder Traps to Quantitatively Study Amphidromous Upstream Migration

Amphidromous stream fauna represent three phyla that have a tropical distribution among oceanic islands, with a few continental representatives. This lifecycle involves marine larval development, with postlarval migration, growth and reproduction as adults occurring in freshwater streams. Many tropical archipelagoes are on the brink of heavy commercial development, threatening freshwater resources in tropical regions. Because of this threat and the isolated nature of tropical islands, quantitative studies are needed to better understand this unique lifecycle. In this paper, we present the use of two modifications of the original Breder trap to study the migration dynamics of tropical amphidromous fish and shrimp postlarvae. The first modification was used in a cemented and channelized stream. The second modification was used in two streams with natural substrates of large, embedded and immovable boulders and basalt outcrops. Quantitative trapping was standardized by time and numbers of traps used, to give results presented as postlarvae trap^–1 h^–1. The number of traps used in the natural streams was based on channel width at the trapping location (i.e., equal number of traps per meter), thus providing equivalent trapping effort between streams of different sizes and flow magnitude. Both trap designs were useful for quantitative monitoring of postlarvae over several months and years, among different streams, and were equally practical for assessing diel and species-specific migration dynamics. Postlarval recruitment of Hawaiian amphidromous species showed temporal variation between months and years within the same stream, significant differences between two streams of different flow magnitude, and distinct diel patterns in diurnal and nocturnal fish and shrimp migration, respectively. A direct correlation between stream flow and total fish postlarval migration was documented.     

Reproductive biology of an endemic,amphidromous goby Lentipes concolor in Hawaiian streams

Synopsis Egg size, fecundity, nest site selection, and breeding structure of the amphidromous Hawaiian goby Lentipes concolor were studied to determine if there was any relationship between these aspects of the breeding biology of this fish and the very long larval period of Hawaiian stream gobies. To do this, these parameters were compared with those known for other gobioid fishes. While eggs were small and females were found to lay up to four nests in a season, reproductive biology and mating behavior of L. concolor were typical of many gobies in spite of its amphidromous life history pattern. Lentipes concolor nests were found from October to June with a spawning peak in February. Nests tended to be positioned closer to the stream bank, and under larger rocks than if they were situated randomly. Depth and mean water column velocity over nest sites did not differ from that at randomly selected points in the stream.     

Floristic biogeography of the Hawaiian Islands: influences of area, environment and paleogeography

Aim A detailed database of distributions and phylogenetic relationships of native Hawaiian flowering plant species is used to weigh the relative influences of environmental and historical factors on species numbers and endemism. Location The Hawaiian Islands are isolated in the North Pacific Ocean nearly 4000 km from the nearest continent and nearly as distant from the closest high islands, the Marquesas. The range of island sizes, environments, and geological histories within an extremely isolated archipelago make the Hawaiian Islands an ideal system in which to study spatial variation in species distributions and diversity. Because the biota is derived from colonization followed by extensive speciation, the role of evolution in shaping the regional species assemblage can be readily examined. Methods For whole islands and regions of each major habitat, species–area relationships were assessed. Residuals of species–area relationships were subjected to correlation analysis with measures of endemism, isolation, elevation and island age. Putative groups of descendents of each colonist from outside the Hawaiian Islands were considered phylogenetic lineages whose distributions were included in analyses. Results The species–area relationship is a prominent pattern among islands and among regions of each given habitat. Species number in each case correlates positively with number of endemics, number of lineages and number of species per lineage. For mesic and wet habitat regions, island age is more influential than area on species numbers, with older islands having more species, more single‐island endemics, and higher species : lineage ratios than their areas alone would predict. Main conclusions Because species numbers and endemism are closely tied to speciation in the Hawaiian flora, particularly in the most species‐rich phylogenetic lineages, individual islands’ histories are central in shaping their biota. The Maui Nui complex of islands (Maui, Moloka‘i, Lāna‘i and Kaho‘olawe), which formed a single large landmass during most of its history, is best viewed in terms of either the age or area of the complex as a whole, rather than the individual islands existing today.     

Progressive island colonization and ancient origin of Hawaiian Metrosideros (Myrtaceae)

Knowledge of the evolutionary history of plants that are ecologically dominant in modern ecosystems is critical to understanding the historical development of those ecosystems. Metrosideros is a plant genus found in many ecological and altitudinal zones throughout the Pacific. In the Hawaiian Islands, Metrosideros polymorpha is an ecologically dominant species and is also highly polymorphic in both growth form and ecology. Using 10 non-coding chloroplast regions, we investigated haplotype diversity in the five currently recognized Hawaiian Metrosideros species and an established out-group, Metrosideros collina, from French Polynesia. Multiple haplotype groups were found, but these did not match morphological delimitations. Alternative morphologies sharing the same haplotype, as well as similar morphologies occurring within several distinct island clades, could be the result of developmental plasticity, parallel evolution or chloroplast capture. The geographical structure of the data is consistent with a pattern of age progressive island colonizations and suggests de novo intra-island diversification. If single colonization events resulted in a similar array of morphologies on each island, this would represent parallel radiations within a single, highly polymorphic species. However, we were unable to resolve whether the pattern is instead explained by ancient introgression and incomplete lineage sorting resulting in repeated chloroplast capture. Using several calibration methods, we estimate the colonization of the Hawaiian Islands to be potentially as old as 3.9 (-6.3) Myr with an ancestral position for Kaua'i in the colonization and evolution of Metrosideros in the Hawaiian Islands. This would represent a more ancient arrival of Metrosideros to this region than previous studies have suggested.     

Coalescent and biophysical models of stepping‐stone gene flow in neritid snails

Marine species in the Indo‐Pacific have ranges that can span thousands of kilometres, yet studies increasingly suggest that mean larval dispersal distances are less than historically assumed. Gene flow across these ranges must therefore rely to some extent on larval dispersal among intermediate ‘stepping‐stone’ populations in combination with long‐distance dispersal far beyond the mean of the dispersal kernel. We evaluate the strength of stepping‐stone dynamics by employing a spatially explicit biophysical model of larval dispersal in the tropical Pacific to construct hypotheses for dispersal pathways. We evaluate these hypotheses with coalescent models of gene flow among high‐island archipelagos in four neritid gastropod species. Two of the species live in the marine intertidal, while the other two are amphidromous, living in fresh water but retaining pelagic dispersal. Dispersal pathways predicted by the biophysical model were strongly favoured in 16 of 18 tests against alternate hypotheses. In regions where connectivity among high‐island archipelagos was predicted as direct, there was no difference in gene flow between marine and amphidromous species. In regions where connectivity was predicted through stepping‐stone atolls only accessible to marine species, gene flow estimates between high‐island archipelagos were significantly higher in marine species. Moreover, one of the marine species showed a significant pattern of isolation by distance consistent with stepping‐stone dynamics. While our results support stepping‐stone dynamics in Indo‐Pacific species, we also see evidence for nonequilibrium processes such as range expansions or rare long‐distance dispersal events. This study couples population genetic and biophysical models to help to shed light on larval dispersal pathways.     

Use of fish behavior in assessing the effects of Hurricane Iniki on the Hawaiian island of Kaua'i

Synopsis Declared the most compact and powerful storm known to have reached the Hawaiian Archipelago, Hurricane Iniki destroyed terrestrial and freshwater habitats island-wide when it struck Kaua'i late on the afternoon of Friday, 11 September 1992. Five weeks after the storm, a research group began underwater studies on the north shore of Kaua'i to assess the physical and biological effects of the hurricane and to track at four-month intervals the recovery of three high quality streams that are home to Hawaii's native and mostly endemic freshwater fishes, mollusks, and crustaceans. Early in the investigation, information on the social behavior of resident fishes showed particular promise for assessing the impact of the hurricane and for tracing the recovery of stream ecosystems. In Hanakapi'ai Stream, the stream damaged most by the hurricane and a subsequent flash flood, two downstream species,Eleotris sandwicensis andStenogobius hawaiiensis, were missing, and members of a third one,Sicyopterus stimpsoni, were sufficiently crowded into the lower section of the stream that aggressive behavior, occasionally resulting in injury, sharply curtailed feeding and prevented all but the earliest phases of courtship. Although the study began in the middle of the spawning season forAwaous guamensis, the few males found were in poor condition and showed neither territorial behavior in response to other males nor courtship when gravid females were encountered. Farther upstream, males ofLentipes concolor directed an inordinate amount of effort toward maintaining territories. Similar, but less dramatic alterations in distribution and behavior were seen in fishes from the less damaged Wainiha River. In Nu'alolo Stream, protected from hurricane winds by nearly vertical valley walls, the behavior was typical for fishes throughout its length except forE. sandwicensis in the damaged section at the stream mouth. Most progress in the return to social behaviors characteristic of pre-hurricane conditions for fishes in Hanakapi'ai Stream and Wainiha River occurred during the first four months of the study, but it was not until October 1993, more than a year after Iniki, that the behavior of the fishes signaled a return to conditions that existed before the storm. The study demonstrated that Hawaii's stream fishes are remarkably capable of rebounding from the effects of hurricanes because of the hardiness of adults and the recruitment of young animals from the ocean as a natural part of each species' amphidromous life cycle.