Biology and Impacts of Pacific Island Invasive Species. 5. Eleutherodactylus coqui, the Coqui Frog (Anura: Leptodactylidae)
Karen H. Beard, Emily A. Price, and William C. Pitt, 297-316

The nocturnal, terrestrial frog Eleutherodactylus coqui, known as the Coqui, is endemic to Puerto Rico and was accidentally introduced to Hawai‘i via nursery plants in the late 1980s. Over the past two decades E. coqui has spread to the four main Hawaiian Islands, and a major campaign was launched to eliminate and control it. One of the primary reasons this frog has received attention is its loud mating call (85–90 dB at 0.5 m). Many homeowners do not want the frogs on their property, and their presence has influenced housing prices. In addition, E. coqui has indirectly impacted the floriculture industry because customers are reticent to purchase products potentially infested with frogs. Eleutherodactylus coqui attains extremely high densities in Hawai‘i, up to 91,000 frogs ha⁻¹, and can reproduce year-round, once every 1–2 months, and become reproductive around 8–9 months. Although the Coqui has been hypothesized to potentially compete with native insectivores, the most obvious potential ecological impact of the invasion is predation on invertebrate populations and disruption of associated ecosystem processes. Multiple forms of control have been attempted in Hawai‘i with varying success. The most successful control available at this time is citric acid. Currently, the frog is established throughout the island of Hawai‘i but may soon be eliminated on the other Hawaiian Islands via control efforts. Eradication is deemed no longer possible on the island of Hawai‘i.

Estimation of the Origin of Polypedates leucomystax (Amphibia: Anura: Rhacophoridae) Introduced to the Ryukyu Archipelago, Japan
Norihiro Kuraishi, Masafumi Matsui, and Hidetoshi Ota, 317-325

We attempted to estimate the origin of the exotic frog Polypedates leucomystax in the Ryukyu Archipelago. This species was first found in 1964 just in front of the U.S. military base at Kadena on Okinawajima Island and currently has established feral populations on more than 20 islands. We conducted phylogenetic analyses using mitochondrial DNA sequences of the cytochrome b gene. Samples of P. leucomystax from five islands of the Ryukyus had a single haplotype, which was identical to that of a Philippine sample but quite different from haplotypes of Vietnamese samples. Samples of P. megacephalus from Taiwan formed a clade different from the P. leucomystax clade. From these results, P. leucomystax in the Ryukyus seems to have originated through accidental transportation of very few individuals with military cargo from somewhere around the Philippines.

Endoparasites of Eleven Species of Ranid Frogs (Anura: Ranidae) from Papua New Guinea
Stephen R. Goldberg, Charles R. Bursey, and Fred Kraus, 327-337

Two hundred eighty-eight ranid frogs from Papua New Guinea collected from 2002 to 2005 were examined for endoparasites: Platymantis adiastolus, P. boulengeri, P. browni, P. gilliardi, P. papuensis, P. schmidti, Rana daemeli, R. garritor, R. jimiensis, R. milneana, and R. papua. Found were one species of Cestoda (as cysticerci), three species of Digenea (Opisthioglyphe cophixali, Diplodiscus amphichrus, and Mesocoelium monas), 18 species of Nematoda (adults of Abbreviata oligopapillata, Aplectana krausi, Aplectana macintoshii, Aplectana zweifeli, Cosmocerca novaeguineae, C. tyleri, Desmognathinema papuensis, Falcaustra papuensis, Icosiella papuensis, Meteterakis crombiei, Ochoterenella papuensis, Paracapillaria spratti, Pseudorictularia dipsarilis, Rhabdias australiensis, Seuratascaris numidica, larvae of Abbreviata sp., and Ascaridae gen. sp.), two species of Acanthocephala (Acanthocephalus bufonis and cystacanths of a second species), and one species of Pentastomida (nymphs of Kiricephalus sp.). Sixty-seven new host records, one new country record, and several new island records are reported. Nematodes composed 18/24 (75%) of the species present. Thirteen of the 24 endoparasite species found currently appear to be endemic to Papua New Guinea.

Black Rat (Rattus rattus) Predation on Nonindigenous Snails in Hawai‘i: Complex Management Implications
Wallace M. Meyer III and Aaron B. Shiels, 339-347

Understanding interactions among nonindigenous species that pose a threat to native species is crucial to effectively preserve native biodiversity. Captive feeding trials demonstrated that the black rat, Rattus rattus, will readily consume two of the most destructive nonindigenous snails, the giant African snail, Achatina fulica (100% predation), and the predatory snail Euglandina rosea (80% predation). Rats consumed snails from the entire size range offered (11.5 to 59.0 mm shell length), suggesting that there is no size refuge above which snails can escape rat predation. Damaged E. rosea shells from the captive feeding trials were compared with shells collected in the Wai‘anae Mountains, O‘ahu. This revealed evidence that R. rattus is responsible for at least 7%–20% of E. rosea mortality. However, this is likely a substantial underestimate because 67% of E. rosea shells in the captive feeding trials were damaged in such a way that they would not have been collected in the field. Therefore, we hypothesize that reduction or eradication of R. rattus populations may cause an ecological release of some nonindigenous snail species where these groups coexist. As such, effective restoration for native snails and plants may not be realized after R. rattus removal in forest ecosystems as a consequence of the complex interactions that currently exist among rats, nonindigenous snails, and the remaining food web.

A Recent Outbreak of the Hawaiian Koa Moth, Scotorythra paludicola (Lepidoptera: Geometridae), and a Review of Outbreaks between 1892 and 2003
William P. Haines, Mandy L. Heddle, Patricia Welton, and Daniel Rubinoff, 349-369

The koa moth, Scotorythra paludicola, is an endemic Hawaiian moth that undergoes sporadic outbreaks in koa forests in Hawai‘i, causing vast defoliations of its host plant, Acacia koa. We studied one such outbreak that occurred on East Maui in 2003, in which approximately 16 km² of forest were defoliated. We collected adult moths and larvae, and recorded size-class distribution of larvae in defoliated regions. Larvae at a given site tended to be of a similar size class, suggesting that outbreaks were synchronous, and mean development time from first instar to adulthood was 42 days under laboratory conditions. Mortality of field-collected, laboratory-reared larvae due to disease was high (80%), making it impossible to quantify meaningful parasitism rates, but three nonnative hymenopteran primary parasitoids were reared (the braconids Meteorus laphygmae and Cotesia marginiventris, and the ichneumonid Hyposoter exiguae). One ichneumonid hyperparasitoid, Gelis sp., was also reared. No native parasitoids were reared. We found no relationship between occurrence of five koa moth outbreaks on East Maui between 1920 and 2006 and annual or monthly precipitation or temperature during that period.

Short-Range Movements of Hawksbill Turtles (Eretmochelys imbricata) from Nesting to Foraging Areas within the Hawaiian Islands
Denise M. Parker, George H. Balazs, Cheryl King, Larry Katahira, and William Gilmartin, 371-382

Hawksbill sea turtles, Eretmochelys imbricata, reside around the main Hawaiian Islands but are not common. Flipper-tag recoveries and satellite tracking of hawksbills worldwide have shown variable distances in post-nesting travel, with migrations between nesting beaches and foraging areas ranging from 35 to 2,425 km. Nine hawksbill turtles were tracked within the Hawaiian Islands using satellite telemetry. Turtles traveled distances ranging from 90 to 345 km and took between 5 to 18 days to complete the transit from nesting to foraging areas. Results of this study suggest that movements of Hawaiian hawksbills are relatively short-ranged, and surveys of their foraging areas should be conducted to assess status of the habitat to enhance conservation and management of these areas.

Growth and Distribution of the Macroalgae Gracilaria salicornia and G. parvispora (Rhodophyta) Established from Aquaculture Introductions at Moloka‘i, Hawai‘i
Stephen G. Nelson, Edward P. Glenn, David Moore, and Brendan Ambrose, 383-396

Gracilaria salicornia and G. parvispora were introduced to the south reef of Moloka‘i, Hawai‘i, in the past 15–20 yr for aquaculture development. Both species have naturalized on the reef. Gracilaria salicornia is now considered an invasive species on O‘ahu due to its tendency to grow in dense beds that produce undesirable windrows of thalli on the beach. There is also concern that it reduces biodiversity and degrades habitats of reefs. We surveyed the south coast of Moloka‘i, where both species were introduced, and measured biomass density, growth rates, and thallus nutrient contents of G. salicornia in established beds. Both species are found in the silt-laden, nearshore zone of the reef within 50 m of shore. Gracilaria salicornia grows in dense beds containing 475 g dry weight m⁻² of biomass, but growth rates are low, 0.03%–1.28% day⁻¹. Tissue nitrogen levels are low, suggesting that these populations are nitrogen limited. Nevertheless, populations of G. salicornia persist and grow slowly on the reef, whereas those of G. parvsipora are only found in areas of local nitrogen enrichment from anthropogenic sources. Currently, G. salicornia does not appear to be negatively affecting the reef ecology on Moloka‘i, because it is confined to the disturbed, nearshore zone. However, its ability to grow slowly and persist under low-nitrogen conditions allows it to form dense beds and suggests that it will eventually spread farther along the coast.

The Soils of Kiritimati (Christmas) Island, Kiribati, Central Pacific: New Information and Comparison with Previous Studies
R. J. Morrison and C. D. Woodroffe, 397-411

Kiritimati, the largest land area atoll in the world, is undergoing rapid population increase, and, given the isolation of the island, local food production will have to be expanded to support the residents. Two soils investigations were completed in the 1960s, but no additional information on the soil resources of the island has been produced since that time. In this study, 15 soil profiles were described and analyzed. Where possible, comparison has been made with previous work, and discussion of the soil-forming factors is presented. Results confirm that soils are weakly developed (Entisols) with relatively low organic matter contents and low water-retention capacity. These properties are expected from the age of the parent materials and the relatively dry climate of the island. Total elemental analyses show that the soils contain very low concentrations of potassium and important trace elements (iron, manganese, copper, and zinc), which will limit any plant production. Classification of the soils identified eight soil families, mainly separated on the basis of content of larger coarse fragments and soil moisture regime, including the influence of groundwater. Comparison with previous studies showed that although different nomenclature and classification systems were used, similar soil patterns were observed, and the soils of Kiritimati are relatively unique in the Pacific islands.

Review of Octocorallia (Cnidaria: Anthozoa) from Hawai‘i and Adjacent Seamounts. Part 2: Genera Paracalyptrophora Kinoshita, 1908; Candidella Bayer, 1954; and Calyptrophora Gray, 1866
Stephen D. Cairns, 413-448

Nine deep-water primnoid octocoral species are described from Hawaiian waters, four of them as new species, bringing the total number of octocoral species known from Hawai‘i to 94. Candidella gigantea is reported for the first time subsequent to its original description from Fiji in 1889. To place the two new species of Calyptrophora in context, all 16 species in the genus are keyed and analyzed in a morphology-based phylogenetic analysis. Although the analysis did not support the species complexes and species groups established by Bayer, it did suggest two distinct clades based on characters such as the opercular cowl, inclination of the polyps, and cross section and sculpture of the basal scale spines.

Currently Known and Reported Discomycetes (Ascomycota) of Hawai‘i
George J. Wong and Richard P. Korf, 449-456

A species list of Discomycetes that occur in Hawai‘i has been compiled that includes all previously reported species in the literature. Comments are provided for reports if there are changes in nomenclature, author citation, or for taxonomic revisions based on reexamination of collections. Fifteen taxa, new to Hawai‘i, are reported. The list of accepted taxa includes a total of 47 species, one including two subspecies. Three previously reported species were misidentified and apparently do not occur in Hawai‘i. Three species formerly reported as Discomycetes are now excluded as Dothideomycetes. The relatively small number of species of Discomycetes recorded from Hawai‘i is probably due to lack of an exhaustive effort to survey this group of Fungi. Although some species are recorded as growing on endemic or indigenous host plants, species of Discomycetes were not designated as endemic or indigenous due to insufficient knowledge of species distribution and the wide range of variations in host preferences.

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Losing the Bounty? Investigating Species Richness in Isolated Freshwater Ecosystems of Oceania
Robert Schabetsberger, Gabriele Drozdowski, Eugen Rott, Rupert Lenzenweger, Christian D. Jersabek, Frank Fiers, Walter Traunspurger, Nicola Reiff, Fabio Stoch, Alexey A. Kotov, Koen Martens, Heinrich Schatz, and Roland Kaiser, 153-179

The South Pacific freshwater ecosystems have never been investigated systematically. Although their ecological value has long been recognized and recommended for protection, little action has been taken so far. Here, we present results of 39 lentic water bodies on 18 islands belonging to seven countries. Temperature, conductivity, and pH were measured and samples of aquatic organisms were collected. Freshwater algae, nematodes, rotifers, ostracods, copepods, cladocerans, and aquatic oribatid mites were identified to genus or species level. Sixty-six percent of all taxa recorded have a cosmopolitan distribution, 14% are circumtropical/tropicopolitan species, and for 20% a restricted distribution predominantly in Australasia has previously been reported. Eleven new copepod and three new ostracod taxa were discovered. Out of 39 water bodies we found at least 17 stocked with nonindigenous fish species. Salinization and uncontrolled introduction of alien fish species may lead to reduced species richness in these remote freshwater ecosystems. The highest species richness was recorded in old, shallow, fish-free softwater lakes at high altitude.

Dietary Shifts by Green Turtles (Chelonia mydas) in the Kāne‘ohe Bay Region of the Hawaiian Islands: A 28-Year Study
Dennis J. Russell and George H. Balazs, 181-192

The green turtle, Chelonia mydas, has modified its feeding behavior to include the increasing abundance of nonnative algae growing in the greater Kāne‘ohe Bay area of O‘ahu in the Hawaiian Islands. Changes in diet of the green turtle are correlated with an increase in abundance of seven species of nonnative algae between 1977 and 2005. Turtles were found to be eating 130 species of marine vegetation, and the three most common were the nonnative species Acanthophora spicifera, Hypnea musciformis, and Gracilaria salicornia. These three abundant and nutritious food sources are now an important part of the turtle diet in addition to native species found in and near Kāne‘ohe Bay. Chelonia mydas behavior has shifted to include these new seaweeds within 10 years of their introduction to the region. The turtles have also gradually included an additional four less-prolific slow-growing nonnative algal species (Eucheuma denticulatum, Gracilaria tikvahiae, Kappaphycus striatum, and Kappaphycus alvarezii), but the time it has taken turtles to include these species has been longer, 20–30 years, after the seaweeds were introduced. During this same 28-year time period numbers of C. mydas have increased throughout the Hawaiian Islands.

Ciguatera in the Introduced Fish Cephalopholis argus (Serranidae) in Hawai‘i and Implications for Fishery Management
Jan Dierking and Cara E. Campora, 193-204

The Peacock gouper (Cephalopholis argus) was introduced to Hawai‘i in 1956 to establish a new fishery. It has become abundant, but the fishery failed due to concerns about ciguatera fish poisoning, a neurological disease in humans caused by ingestion of fish containing ciguatoxin. The aim of this study was to provide better understanding of geographic patterns of ciguatoxicity in C. argus and of the correlation of toxicity with morphometric characters of this species, with the goal to assess the possibility of a safe fishery. Overall, 18.2% of C. argus specimens from sites around O‘ahu and Hawai‘i Island contained ciguatoxin in concentrations potentially harmful to humans. This was higher than the rate of occurrence in Hawaiian reef fishes in general, and on the scale of ciguatoxicity in species banned from sale in fish markets. Toxicity was high around both analyzed islands. However, toxic individuals were significantly less common around O‘ahu than around Hawai‘i Island (8% versus 24%). Regular geographic patterns in toxicity within islands (e.g., gradients along coastlines) were not present, and variability in toxicity within each sample site was high. Toxicity was significantly but weakly positively correlated with C. argus length but not with fish condition (measured by length at weight). In conclusion, high prevalence of toxic individuals, variability in toxicity on all analyzed spatial scales, and low explanatory power of morphometric characters make the avoidance of ciguatoxic C. argus individuals difficult. A safe fishery for this species in Hawai‘i therefore does not appear feasible at present.

Distribution, Density, and Biomass of Introduced Small Mammals in the Southern Mariana Islands
Andrew S. Wiewel, Amy A. Yackel Adams, and Gordon H. Rodda, 205-222

Although it is generally accepted that introduced small mammals have detrimental effects on island ecology, our understanding of these effects is frequently limited by incomplete knowledge of small mammal distribution, density, and biomass. Such information is especially critical in the Mariana Islands, where small mammal density is inversely related to effectiveness of Brown Tree Snake (Boiga irregularis) control tools, such as mouse-attractant traps. We used mark-recapture sampling to determine introduced small mammal distribution, density, and biomass in the major habitats of Guam, Rota, Saipan, and Tinian, including grassland, Leucaena forest, and native limestone forest. Of the five species captured, Rattus diardii (sensu Robins et al. 2007) was most common across habitats and islands. In contrast, Mus musculus was rarely captured at forested sites, Suncus murinus was not captured on Rota, and R. exulans and R. norvegicus captures were uncommon. Modeling indicated that neophobia, island, sex, reproductive status, and rain amount influenced R. diardii capture probability, whereas time, island, and capture heterogeneity influenced S. murinus and M. musculus capture probability. Density and biomass were much greater on Rota, Saipan, and Tinian than on Guam, most likely a result of Brown Tree Snake predation pressure on the latter island. Rattus diardii and M. musculus density and biomass were greatest in grassland, whereas S. murinus density and biomass were greatest in Leucaena forest. The high densities documented during this research suggest that introduced small mammals (especially R. diardii) are impacting abundance and diversity of the native fauna and flora of the Mariana Islands. Further, Brown Tree Snake control and management tools that rely on mouse attractants will be less effective on Rota, Saipan, and Tinian than on Guam. If the Brown Tree Snake becomes established on these islands, high-density introduced small mammal populations will likely facilitate and support a high-density Brown Tree Snake population, even as native species are reduced or extirpated.

Critically Endangered Fijian Crested Iguana (Brachylophus vitiensis) Shows Habitat Preference for Globally Threatened Tropical Dry Forest
Clare Morrison, Gunnar Keppel, Nunia Thomas, Isaac Rounds, and Peter S. Harlow, 223-251

Tropical dry forests are a unique and threatened ecosystem in the Pacific and globally. In Fiji, the endangered Fijian crested iguana (Brachylophus vitiensis) is endemic to tropical dry forests. Yadua Taba Island contains one of the best remaining stands of tropical dry forest in the Pacific along with the largest (and only secure) population of B. vitiensis in Fiji and has been proposed as a translocation source for iguana conservation. In this study we determined the major vegetation types on Yadua Taba and identified forest habitat preferences of B. vitiensis to (1) characterize the island’s habitats for tropical dry forest regeneration monitoring and (2) understand which forest types are preferred by iguanas for future translocation projects. Vegetation data were collected using reconnaissance, entitation, line transects, and aerial photos. Iguana abundance data were collected by nocturnal surveys of permanent transects. Six major vegetation types were identified of which tropical dry forest was the largest (46% of the island), followed by a combination of rocky cliff–shrubland/grassland vegetation (26%). Our conservative estimate of B. vitiensis population size on Yadua Taba is 12,000 iguanas, the majority of which occur in tropical dry forest. Superabundance of the dry forest understory tree Vavaea amicorum, the favorite fruit species of iguanas, may help account for the high density of iguanas observed. These results highlight the ecological link between tropical dry forest and B. vitiensis and emphasize the importance of rehabilitation or conservation of tropical dry forest habitat in potential iguana translocation sites as part of the management plan for B. vitiensis throughout the Fiji Islands.

Carlia ailanpalai (Reptilia: Scincidae): An Invasive Species of Lizard in the Federated States of Micronesia
Donald W. Buden, 243-251

Distribution of the introduced scincid lizard Carlia ailanpalai Zug in the Federated States of Micronesia (FSM) is reviewed. It is common in open grassy areas but seldom occurs in mature forest. Preliminary surveys indicate that it is well established in Yap, though less frequently encountered at increasing distance from Colonia, the main settlement, and it is unrecorded in the extreme northern and southern parts of Yap. It is the most common species of lizard in open, grassy, ruderal habitats throughout Weno Island, Chuuk, being nearly the only species encountered in the commercial district, but it is unknown elsewhere in Chuuk State. The only record for Kosrae is a single specimen collected in 1988 (first record for the FSM), but there is no evidence of an established population. There are no records for Pohnpei State. Guam is likely the primary source for the Yap and Chuuk populations (and Kosrae specimen), but the time of initial introduction is unknown. Carlia ailanpalai appears to have spread rapidly, at least on Weno, Chuuk, where it has become the predominant lizard in open habitats islandwide, possibly since the late 1960s. How C. ailanpalai interacts with other species in the FSM requires further study, but preliminary surveys of distribution and relative abundance suggest that it has a negative impact on populations of Emoia jakati and, to a lesser extent, on other Emoia species as well. Populations of C. ailanpalai in the FSM meet the criteria for invasive species status as it is defined by numerous U.S. government agencies and international conservation groups.

Evidence of a Possible Decline since 1989 in False Killer Whales (Pseudorca crassidens) around the Main Hawaiian Islands
Randall R. Reeves, Stephen Leatherwood, and Robin W. Baird, 253-261

Recent evidence indicates that there is a small, demographically isolated, island-associated population of false killer whales (Pseudorca crassidens) around the main Hawaiian Islands. Although it is known that false killer whales in Hawai‘i are sometimes killed or seriously injured in the Hawai‘i-based long-line fishery, it is not known whether such interactions have resulted in a reduction in population size or whether other factors have been negatively influencing population size. We report the results of an aerial survey in June and July 1989, the purpose of which was to obtain a minimum count of the number of false killer whales around the main Hawaiian Islands. The false killer whale was the third most commonly seen species of odontocete off the island of Hawai‘i during the survey, representing 17% of sightings. Groups of more than 300 individuals were seen on three different days, with minimum counts of 380, 460, and 470 individuals in these groups. The encounter rate, relative species ranking, and average group size from the 1989 survey were all substantially greater than those from more recent aerial and ship-based surveys. The largest group observed in 1989 (470) contained almost four times as many whales as estimated for the entire main Hawaiian Islands from recent aerial surveys (121 individuals, CV = 0.47) or mark-recapture analyses (123 individuals, CV = 0.72). Therefore, the population of false killer whales around the main Hawaiian Islands may have declined substantially since 1989. The cause or causes of such a decline are uncertain.

Andvakia discipulorum, A New Species of Burrowing Sea Anemone from Hawai‘i, with a Revision of Andvakia Danielssen, 1890
Marymegan Daly and Roger H. Goodwill, 263-275

We describe Andvakia discipulorum Daly & Goodwill, n. sp., from an intertidal mudflat of Kāne‘ohe Bay, O‘ahu, Hawai‘i. Members of this species are inconspicuous, being small and having a column covered with sand. In comparison with other species of the genus, Andvakia discipulorum, n. sp., presents distinct arrangement of mesenteries, sizes of nematocysts, and musculature. We also provide a redescription of Andvakia boninensis based on specimens collected from Saipan, Mariana Islands. These descriptions provide an opportunity to revise and update the taxonomy of Andvakia and to address the systematics of family Andvakiidae. We determine that Andvakia is the senior synonym of Decaphellia and reject earlier hypotheses of synonymy between Andvakia and Capneopsis, Ilyactis, and Octophellia. A tabular key to the species of Andvakia is provided.

A New Name for the Hawaiian Antipatharian Coral Formerly Known as Antipathes dichotoma (Cnidaria: Anthozoa: Antipatharia)
Dennis M. Opresko, 277-291

A Hawaiian species of antipatharian coral previously identified as Antipathes dichotoma Pallas, 1766, is described as Antipathes griggi Opresko, n. sp. The species forms tall, bushy colonies with elongate, upright terminal branches, often arranged uniserially. Spines are conical, mostly 0.20 to 0.26 mm tall, apically bifurcated, multilobed to jagged in appearance, and covered over most of their surface with small roundish to elongate papillae. Minute secondary spines may occur on some of the thicker branches. Polyps are 1 to 1.6 mm in transverse diameter. The species resembles A. fruticosa Gray in branching pattern, size of spines, and presence of secondary spines but differs in morphology and density of the spines (thicker, more crowded primary spines and fewer secondary spines in A. griggi). Other related species differ from A. griggi in having more widely spreading and irregularly arranged branches, no secondary spines, and either smaller spines with fewer apical lobes (A. curvata van Pesch, A. arborea Dana, and A. galapagensis Deichmann) or larger spines with the apical lobes arranged in a somewhat coronate pattern [A. spinulosa (Schultze) and A. lentipinna Brook].

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Pacific Science, the UH Press’s original journal and only journal in the natural sciences, now has a new Index to Online Archives, which contains an increasingly comprehensive listing of published tables of contents (ToCs) from each volume, with links from each article title to its online edition. From vol. 47 (1993), the ToCs also contain abstracts. All volumes from 1 (1947) through 54 (2000) will be available by open access in UH Hamilton Library’s ScholarSpace digital repository. So far, the repository contains complete volumes going back as far as the early 1970s, but we plan to have all the volumes from the 1960s, 1950s, and 1940s available online by June 2009.

A short history of the journal’s migration from print to electronic availability follows.

In 2001, Pacific Science made its first online debut in Project Muse, which still contains archives of all the issues from volume 55 (2001) through volume 61 (2007).

In 2007, Pacific Science made its debut in BioOne.2, starting with issues going back to volume 59 (2005).

In 2008, Hamilton Library at the University of Hawai‘i began scanning and adding back issues of Pacific Science to its ScholarSpace digital repository, which will eventually include all issues starting from vol. 1 (1947).

Habitat-Mediated Use of Space by Juvenile and Mating Adult Port Jackson Sharks, Heterodontus portusjacksoni, in Eastern Australia
David Mark Powter and William Gladstone, 1

Studies of spatial ecology of demersal sharks are critical to understanding the significance of habitat variation; however, limited information exists. Spatial ecology of adult Heterodontus portusjacksoni was studied at three locations on the central and southern coast of New South Wales, Australia, from January 2002 to December 2005. Juveniles within a nursery area were studied from December 2002 to December 2005. Tag-recapture, day and night underwater visual census, and acoustic tagging were used. Adults returned annually to the same coastal breeding reefs for up to four consecutive years. Individual juveniles resided within a sea-grass nursery area for at least 2 yr and were not uniformly distributed throughout the nursery. Adult females often sheltered in aggregations in gutters as a male avoidance strategy, and both sexes utilized the sand/reef interface in the absence of gutters. Juveniles aggregated infrequently due to absence of habitat features that mediated aggregation. Acoustic tracks of adults revealed periods of inactivity up to 27 hr. Juveniles spent significant amounts of time inactive, punctuated with short bouts of swimming. Juveniles utilized moderate activity spaces (3,510–583,990 m²) centered over a core area of the sea-grass bed but also ranged over much larger areas of the bay. Use of space by H. portusjacksoni is strongly influenced by habitat characteristics throughout its life history.

Characteristics of Scylla spp. (Decapoda: Portunidae) and their Mangrove Forest Habitat in Ngaremeduu Bay, Republic of Palau
Katherine C. Ewel, Stacy Rowe, Blake NcNaughton, and Kimberly M. Bonine, 15

Three species of mangrove crabs (Scylla spp.) were captured in live traps in Ngaremeduu Bay on the island of Babeldaob, Republic of Palau. Most were S. serrata, but one individual each of S. olivacea and S. paramamosain was also trapped, establishing existence of a biogeographic gradient in mangrove crab species diversity across the Micronesian archipelago. Species composition of mangrove trees along transects around the bay and along the three major tributaries was similar to that of other Micronesian islands, although trees are smaller in Palau. For 17 months in 1999–2000, crabs were trapped in the bay and captured by hand along the transects; they were trapped again for 1 month in 2004. Characteristics of the crabs and of burrows encountered along the transects suggested that only S. serrata was captured in 1999–2000 and that population density of this species was 40 crabs ha­-¹. Carapace widths for the 159 crabs captured during the entire study did not differ significantly over the 4-yr span, and averaged 153 mm for males and 137 mm for females. However, average carapace widths for the largest quartile of crabs declined significantly from 174 mm to 171 mm across the study period. Catch per unit effort was 0.28 crab per trap night in 1999–2000 and 0.45 in 2004. Although large crabs are still available in Ngaremeduu Bay, current regulations may not be sufficient to keep populations from decreasing gradually in size, especially in the face of increasing harvest pressure on the island of Babeldaob.

Behavioral Responses of the Endemic Shrimp Halocaridina rubra (Malacostraca: Atyidae) to an Introduced Fish, Gambusia affinis (Actinopterygii: Poeciliidae) and Implications for the Trophic Structure of Hawaiian Anchialine Ponds
Krista A. Capps, Caroline B. Turner, Michael T. Booth, Danica L. Lombardozzi, Scott H. McArt, David Chai, and Nelson G. Hairston Jr., 27

In the Hawaiian Islands, intentionally introduced exotic fishes have been linked to changes in native biodiversity and community composition. In 1905, the mosquito fish Gambusia affinis was introduced to control mosquitoes. Subsequently, G. affinis spread throughout the Islands and into coastal anchialine ponds. Previous studies suggest that presence of invasive fishes in anchialine ponds may eliminate native species, including the endemic shrimp Halocaridina rubra. We examined effects of G. affinis on H. rubra populations in anchialine ponds on the Kona-Kohala coast of the island of Hawai‘i. In the presence of G. affinis, H. rubra exhibited a diel activity pattern that was not seen in fishless ponds. Shrimp in ponds with fish were active only at night. This pattern was evident in anchialine ponds and in laboratory experiments. In laboratory predation experiments, G. affinis preferentially consumed smaller H. rubra, and in the field the H. rubra collected from invaded sites were larger than those from fishless ponds. Analysis of trophic position using stable isotope analyses showed that feeding of H. rubra was not significantly distinct from that of snails, assumed to feed at trophic level 2.0 on epilithic algae, but G. affinis was slightly omnivorous, feeding at tropic level 2.2. The mosquito fish diet was apparently composed primarily of algae when the defensive behavior of H. rubra made them substantially unavailable as prey. The effect of successful establishment of G. affinis on shrimp behavior has the potential to alter abundance of benthic algae and processing and recycling of nutrients in anchialine pond ecosystems.

Expressions of 1976–1977 and 1988–1989 Regime Shifts in Sea-Surface Temperature off Southern California and Hawai‘i
Laurence C. Breaker and Stephanie J. Flora, 39

Sea-surface temperatures off southern California from Scripps Pier and from Koko Head, Hawai‘i, were examined to determine what impact regime shifts that occurred in 1976–1977 and 1988–1989 had on environmental conditions at each location. Cumulative sums were employed to enhance the detection process. The cumulative sum time histories revealed major turning points at both locations at the time of the 1976–1977 event. At both locations, increases in temperature were indicated, consistent with the phase change in the Pacific Decadal Oscillation that took place at that time. The cumulative sums also indicated major turning points at both locations during the 1988–1989 event. A new procedure called the method of expanding means was employed to determine the long-term impact of these events. By comparing means before and after a given event it is possible to observe the magnitude of the change and to what extent it is sustained. For the 1976–1977 regime shift, temperatures increased rapidly and remained consistently higher, by ~1°C for 2–3 yr at Scripps Pier. This increase occurred over a period of approximately 7 months and accounts for more than half of the total warming that has occurred at that location since 1920. At Koko Head, a similar response was observed with a sustained increase of approximately +0.5°C. The oceanic response to the 1988–1989 event was quite different. At Scripps Pier, temperatures before and after this event did not show any tendency to converge to significantly different values out to periods of 2–3 yr. At Koko Head, mean temperatures did converge to slightly different values after 1 yr, with mean values being consistently lower after this event (~–0.4°C). It was shown that in some cases changes associated with these events could be identified in the original data, but without the help of cumulative sums, it is usually not possible to make a clear distinction between changes of interest and other sources of variability. Finally, decorrelation time scales for the records at both locations were estimated and found to be on the order of a year, implying spatial scales that are at least synoptic (tens to hundreds of kilometers).

Reef-Top Sediment Bodies: Windward O‘ahu, Hawai‘i
Christopher Bochicchio, Charles Fletcher, Matthew Dyer, and Thomas Smith, 61

Hawaiian fringing reefs display sand bodies on their surfaces that are potentially important components of littoral sediment budgets. This work provides a regional survey of modern reef-top sediment storage and investigated geologic controls on sediment storage potential. Sand bodies are formed when sediment accumulates in topographic depressions that are the result of meteoric water eroding the emerged carbonate reef platform during periods of lower sea level. The relief of some depressions may be modified by Holocene reef accretion. Depression morphology exerts a strong control on volume and internal distribution of sediment. In this study a total of 205 jet probe thickness measurements was collected from 54 major sand bodies on the fringing reef (0–20 m depth) adjacent to 22 km of Southeast O‘ahu coastline (Kailua, Lanikai, and Waimānalo). Volumes were determined and synthesized with previous volume estimates of coastal subaerial and deeper submarine sediment bodies (20–200 m depth), giving the total sediment storage within the coastal system. Sand bodies range from 50 to 2,800 m from shore. Measured thickness varied from 0 to greater than 3.0 m with a mean of 0.95 m. For this study sand bodies were classified into three dominate morphologies: channel, field, and karst depression. The volume of sediment stored in channels was 58,253±618×10³ m³, fields contained 171±6×10³ m³, and karst depressions contained 1,332±248×10³ m³. Correlation of sediment body distribution with reef and coastal plain morphology revealed potential geologic controls on sand body formation in this region. Meteoric runoff and reef slope are important controls on spatial distribution of sand bodies.

Hyperiid Amphipods (Crustacea: Peracarida) in Mexican Waters of the Pacific Ocean
Rebeca Gasca, 83

Information on regional diversity of hyperiid amphipods of the eastern Pacific Ocean is still largely incomplete. Recent surveys of hyperiid fauna from the Mexican Pacific motivated a revision of extant faunistic accounts. This revised list includes all records from Mexican waters of the eastern Pacific (MP) from Baja California to the southern border with Central America. A total of 150 species belonging to 19 families and 48 genera of the Hyperiidea was included in this account; seven are new records in the MP. Up to 31 nominal species were excluded from previous listings. Overall, the epipelagic infraorder Physocephalata is highly diverse in the MP (119 species); Physosomata, containing deep-living forms, are less diverse (31 species). The northern part of the MP (including the gulf and off the Baja California peninsula) harbors the highest number of species/records, whereas nearly half of the species are known from the central areas and six from the southernmost sector of the MP. This pattern reflects current knowledge of the group in these areas and also geographic differences in the sampling/research efforts, but it is not a diversity gradient. Species richness of the MP is comparable with that known from other Pacific subregions. The epipelagic hyperiid fauna of the tropical MP remains relatively unknown and should be studied further to reveal regional patterns of diversity. The deep-living hyperiid community of the tropical eastern Pacific harbors a diversity that is deserving of further study.

Evidence for a Correlation between Systematics and Bioactivity in New Caledonian Cunoniaceae and Its Implications for Screening and Conservation
Yohan Pillon and Bruno Fogliani, 97

It is generally assumed that there is a good correlation between systematics and the secondary compounds found in plants. However because of the frequent homoplasy of chemical characters this has been difficult to test using statistical methods. Here we applied two nonparametric tests on a published data set, where 50 species of New Caledonian Cunoniaceae were screened for bioactivity against several pathogenic strains. Using Moran’s I index we showed that in two of nine tests against pathogenic strains there was a significantly higher similarity than expected in bioactivities between species belonging to the same genus and a significantly higher than expected dissimilarity in bioactivity between species belonging to different tribes. When considering the bioactivities against all pathogenic strains with Mantel tests, we also found significant correlation between bioactivity and phylogenetic distance in two of four tests. This has implications in screening and conservation. Searches for new molecules and bioactivity should preferentially be made on species spread across the tree of life. There is also a need to preserve as much phylogenetic diversity as possible to make sure that the widest reservoir of natural compounds remains available for future generations.

Subfossil Land Snails from Easter Island, Including Hotumatua anakenana, New Genus and Species (Pulmonata: Achatinellidae)
Patrick V. Kirch, Carl C. Christensen, and David W. Steadman, 105

The depauperate modern terrestrial biota of Easter Island contrasts with that of most other southeastern Polynesian high islands, which characteristically support a number of endemic species of insects, land snails, birds, and plants. We investigated cultural and noncultural late Holocene deposits at Anakena, Easter Island, establishing the former presence of endemic land snails on the island. These include an unidentified helicinid, a Nesopupa species, and a previously undescribed extinct achatinellid land snail, Hotumatua anakenana Kirch, Christensen & Steadman, n. genus and n. sp. A human-introduced achatinellid, Pacificella variabilis, occurs in later stratigraphic contexts of the same site. Prehistoric deforestation may have been the primary cause of the extinction of Hotumatua, although predation by rats or other alien species may have been involved as well. Along with recently discovered extirpated species of angiosperms, seabirds, and land birds, the extinction of Hotumatua reflects the nearly complete loss of the native biota of Easter Island after Polynesian colonization about 1,000 yr ago.

A New Live-Bearing Species of Scincid Lizard (Reptilia: Scincidae) from New Caledonia, Southwest Pacific
Ross A. Sadlier, Sarah A. Smith, Anthony Whitaker, and Aaron M. Bauer, 123

A new species of skink, Kanakysaurus zebratus, is described from the ultramafic Massif de Kopéto and Massif de Koniambo on the northwestern coast of Grande Terre, New Caledonia. Although this new species is similar in overall appearance to its congener K. viviparus from the far northwest of Grande Terre and the Îles Belep, it can be distinguished by features of scalation and coloration. It is also identified as being genetically distinct from and reciprocally monophyletic with respect to populations of K. viviparus from Rivière Néhoué (type population), the Îles Belep, and a recently discovered population from Sommet Poum (reported here for the first time). The population of Kanakysaurus on Dôme de Tiébaghi (5 km southeast of Rivière Néhoué) is problematic: in morphology it is closest to K. viviparus, but DNA sequence data group part of the population with K. viviparus and part with K. zebratus, n. sp. On Kopéto the new species was found only in maquis shrubland at 500–1,000 m in elevation and on Koniambo in Gymnostoma-dominated closed forest at 700 m. Adult females collected on the Massif de Kopéto in February during the height of the wet season had well-developed embryos, confirming a live-bearing mode of reproduction for the new species, and for the genus as a whole. The summit area of Kopéto is the site of a large nickel mine and substantial portions of the known range of the new species are projected to be cleared to extract nickel-bearing ore in the future; extensive development for nickel mining is also forecast in the immediate future for Koniambo. Because of the apparently restricted range and projected degradation of habitat of this new species, it is here regarded as assignable to IUCN Red List Category Endangered and considered a high priority for conservation management.

Grammonus nagaredai, a New Viviparous Marine Fish (Ophidiiformes: Bythitidae) from the Hawaiian Islands
John E. Randall and Marc James Hughes, 137

Grammonus nagaredai is described as a new species of viviparous bythitid fish from two specimens collected from caves off O‘ahu and Hawai‘i at depths of 6–9 m. Grammonus yunokawai Nielsen, known from one specimen from a cave in 20 m in the Ryukyu Islands, is the most similar species, differing in having a deeper body, more convex nape, broader maxilla, longer predorsal length, and shorter pectoral fins.

Association Affairs, 147

Historical Tropical Cyclone Activity and Impacts in the Cook Islands
Fes A. de Scally, 443

Analysis of a recently completed database of 143 tropical cyclones in the Cook Islands revealed a minimum average frequency of 0.8 cyclones per cyclone season between 1820 and 2006, with a more-precise frequency of 1.8 cyclones per season with the beginning of satellite monitoring of cyclones in 1970. Since 1970, 31% of cyclones have reached hurricane intensity. The Southern Cooks have been more than twice as frequently affected by cyclones as the Northern Cooks, with the island of Palmerston having the greatest number of encounters. Since 1820, 96% of cyclones have occurred during the official November–April cyclone season, with February alone accounting for 29%. Since 1970, 46% of cyclones achieving hurricane status have occurred in February. Nevertheless, Cyclone Martin in October–November 1997 demonstrated the dangers of a cyclone occurring outside the official season. An increase in cyclone occurrences since the mid-1970s is probably attributable to satellite monitoring, but it is noteworthy that all six cyclones known for certain to have achieved major hurricane status have occurred since 2002. Since 1970, 56% of cyclones have occurred during El Niño events, an increase of 15% from the 1870–1969 period. Since 1891, cyclones with moderate and major human impacts have occurred on average at least every 3.8 and 8.8 yr, respectively, with the Southern Cooks more than twice as frequently affected as the Northern Cooks. However, past cyclone disasters in the latter group suggest that risk to human life is greater there due to the potential for inundation of the atolls by storm surges. Half of cyclones with human impacts have occurred during El Niño events, with weak to moderate El Niños almost as important in this respect as strong El Niños. Only 13% of cyclone impacts have occurred during La Niña events.

Was Tropical Cyclone Heta or Hunting by People Responsible for Decline of the Lupe (Ducula pacifica) (Aves: Columbidae) Population on Niue during 1994–2004?
R. G. Powlesland, D. J. Butler, and I. M. Westbrooke, 461

On 6 January 2004, Tropical Cyclone Heta devastated much of the South Pacific island nation of Niue. The forest suffered extensive damage, particularly to the north-western sector, with many trees uprooted and others stripped of branches and foliage. Even though some patches of forest in the southeast sustained little damage, many lupe (Pacific pigeon, Ducula pacifica) and kulukulu (purple-crowned fruit dove, Ptilinopus porphyraceus) entered eastern villages in search of food and water after the cyclone, a very unusual behavior. This paper details our findings from a survey of some of Niue’s forest birds carried out during September 2004 and compares these with results from a similar survey in September 1994. Five-minute point count data, an index of conspicuousness, from three transects showed that heahea (Polynesian triller, Lalage maculosa) were more abundant in 2004 than in 1994, that the results were variable from transect to transect for miti (Polynesian starling, Aplonis tabuensis) and kulukulu, but that significantly fewer lupe were detected along all three transects in 2004 than previously. We tentatively suggest that the decline in the lupe population was caused mainly by unsustainable human hunting during 1994–2004, rather than mortality caused by the cyclone.

Ecological Partitioning and Invasive Ants (Hymenoptera: Formicidae) in a Tropical Rain Forest Ant Community from Fiji
Darren Ward, 473

Determining composition and structure of ant communities may help understand how niche opportunities become available for invasive ant species and ultimately how communities are invaded. This study examined composition and structure of an ant community from a tropical rain forest in Fiji, specifically looking at spatial partitioning and presence of invasive ant species. A total of 27 species was collected, including five invasive species. Spatial partitioning between arboreal (foliage beating) and litter (quadrat) samples was evident with a relatively low species overlap and a different composition of ant genera. Composition and abundance of ants was also significantly different between litter and arboreal microhabitats at baits, but not at different bait types (oil, sugar, tuna). In terms of invasive ant species, there was no difference in number of invasive species between canopy and litter. However, the most common species, Paratrechina vaga, was significantly less abundant and less frequently collected in the canopy. In arboreal samples, invasive species were significantly smaller than endemic species, which may have provided an opportunity for invasive species to become established. However, taxonomic disharmony (missing elements in the fauna) could also play an important role in success of invasive ant species across the Pacific region. Invasive ants represent a serious threat to biodiversity in Fiji and on many other Pacific islands. A greater understanding of habitat susceptibility and mechanisms for invasion may help mitigate their impacts.

Ecology of the Endemic Land Crab Johngarthia malpilensis (Decapoda: Brachyura: Gecarcinidae), a Poorly Known Species from the Tropical Eastern Pacific
Mateo López-Victoria and Bernd Werding, 483

Johngarthia malpilensis (Faxon, 1893) is the least studied of the eight American species of Gecarcinidae. This land crab is considered endemic to Malpelo, an oceanic island of the Colombian Pacific. Several aspects of its ecology were investigated between 2003 and 2006. We estimated its population density, distribution, daily activity, reproduction, interactions, and diet by marking and monitoring 909 individuals. During our visits we recorded crabs of sizes from 5 to 82 mm carapace width. Johngarthia malpilensis shelters mainly in fissures and hollows between rocks. It is distributed all over the main island except in very steep sectors. An average density of 0.41 adults m-2 and 0.55 juveniles m-2 produced an estimated total population of 833,000. Johngarthia malpilensis showed high mobility, with crabs covering distances over 450 m in a few days on highly irregular surfaces. Activity was higher from dusk till dawn and lowest around noon. Release of larvae took place during the high tides associated with the new moon, at least during the rainy season. It is omnivorous and opportunistic, consuming practically every available resource. The crab is occasionally preyed upon by an endemic lizard and migratory birds. Its general ecology is very similar to that of J. planatus, a closely related species. As a voracious omnivore J. malpilensis is one of the most important components of Malpelo’s food web.

First Records of Butterflies (Lepidoptera) from the Republic of Nauru
Donald W. Buden and W. John Tennent, 495

Four species of butterflies are reported from Nauru for the first time and as first records of butterflies from the island republic. None is endemic. Three of the four species are widespread in Oceania: Badamia exclamationis (Fabricius), Danaeus plexippus (Linnaeus), and Hypolimnas bolina (Linnaeus). The other, Petrelaea tombugensis (Röber), belongs to a genus that also is widespread in the Pacific. The small number of widespread species found on Nauru is comparable with the situation encountered on other small, remote, low-lying Pacific islands.

The Reptiles of Nauru
Donald W. Buden, 499

Eleven species of reptiles are reported from Nauru in the first systematic treatment of the herpetofauna. Four of the species are marine; the seven others include six lizards (four geckos, two skinks) and one snake. Gehyra mutilata (Wiegman), G. oceanica (Lesson), Pelamis platura (Linnaeus), and Ramphotyphlops braminus (Daudin) are recorded on Nauru for the first time. With the exception of Emoia arnoensis Brown & Marshall, which is endemic to eastern Micronesia, the herpetofauna consists of species that range widely among the west-central Pacific Ocean islands. The only known record of E. arnoensis from Chuuk possibly is based on a misassigned locality, in which case the range of the species would be limited to the Marshall Islands, Nauru, and Kosrae. There is no evidence to suggest that habitat modification on Nauru stemming largely from more than a century of phosphate mining has reduced the number of reptile species.

Inventory of Thysanoptera Collected from French Polynesia
Mark S. Hoddle, Christina D. Hoddle, and Laurence A. Mound, 509

A survey for Thysanoptera was conducted in the Society (Tahiti, Moorea, and Raiatea), Marquesas (Hiva Oa, Nuku Hiva, Ua Huka, and Ua Pou), and Austral islands (Rurutu and Tubuai) archipelagos in French Polynesia from September 2003 to November 2005. At least 55 thrips species in 36 genera and three families were identified from 823 slide-mounted specimens that were collected from 61 host plants in 33 families. Twelve species are considered to be important pests. The greatest diversity of species, 43 (77%), was collected from the Society Islands, with 60% being recorded from Tahiti alone. Species diversity was intermediate in the Marquesas Islands at 43% (24 species collected), with 35% or 19 species being recorded from Nuku Hiva. Lowest diversity was recorded for the Austral Islands, with 38% or 21 species being found in that archipelago. Less than 10% of collected species are likely to be native, with the majority of identified thrips (>90%) in French Polynesia representing a high diversity of exotic species (leaf, flower, and fungus feeders, and four predatory species) that have successfully infiltrated other island groups in the South Pacific. Survey results and subsequent estimates of thrips species diversity in French Polynesia should be interpreted with caution due to uncontrolled variation in sampling intensity that was affected by survey duration, time of year, and visitation frequency to islands.

Attempt to control the Invasive Red Alga Acanthophora spicifera (Rhodophyta: Ceramiales), in a Hawaiian Fishpond: Assessment of Removal Techniques and Management Options
Mariska Weijerman, Rebecca Most, Kristy Wong, and Sallie Beavers, 517

Acanthophora spicifera (Vahl) Børgesen was unintentionally introduced to Hawai‘i in 1950 and has since become the most common nonindigenous algal species in the main Hawaiian Islands. On the west coast of Hawai‘i Island it has been documented at three sites, including Kaloko Fishpond in Kaloko-Honokōhau National Historical Park. The fishpond has an open connection to the sea, increasing the risk that A. spicifera will establish itself on neighboring shallow coral reefs and rocky intertidal habitats. To diminish that risk and to develop an efficient management strategy, a range of approaches was assessed to control this invasive alga in Kaloko Fishpond. Removal techniques were labor intensive and had limited effect. All experiments showed a substantial initial decrease in algal density, but the long-term effect was minimal because of rapid regrowth. The most promising removal method was the use of submerged shelters to raise local densities of herbivorous fishes. Fishes grazed the alga and quickly reduced the biomass. However, the large number of predators and absence of topographical structure will make it challenging to provide sufficient shelters to increase the herbivorous fish population in the entire fishpond. A management strategy to substantially reduce the algal biomass in the fishpond includes a combination of biological control and periodic manual removal of the alga.

Relationships Between Otolith Size and Body Size for Hawaiian Reef Fishes
Ken Longenecker, 533

Estimating body size of fishes from remains recovered from piscivores, archaeological sites, and sedimentary deposits is desirable but rarely accomplished because the relationships between the size of a fish and its durable anatomical structures are largely unknown. Regression equations to predict the size or weight of 41 common Hawaiian reef fishes from sagittae (saccular otoliths) are presented. Data are also grouped into higher taxa to permit size predictions when otoliths cannot be assigned to species.

Maximum Annually Recurring Wave Heights in Hawai‘i
Sean Vitousek and Charles H. Fletcher, 541

The goal of this study was to determine the maximum annually recurring wave height approaching Hawai‘i. The motivation was scientific as well as administrative: to enhance understanding of the recurring nature of dominant swell events, as well as to inform the Hawai‘i administrative process of determining the “upper reaches of the wash of the waves” (Hawai‘i Revised Statutes [H.R.S.] § 205-A), which delineates the shoreline. We tested three approaches to determine the maximum annually recurring wave, including log-normal and extremal exceedance probability models and Generalized Extreme Value (GEV) analysis using 25 yr of buoy data and long-term wave hindcasts. The annual recurring significant wave height was found to be 7.7 ± 0.28 m (25 ft ± 0.9 ft), and the top 10% and 1% wave heights during this annual swell was 9.8 ± 0.35 m (32.1 ft ± 1.15 ft) and 12.9 ± 0.47 m (42.3 ft ± 1.5 ft), respectively, for open North and Northwest Pacific swell. Directional annual wave heights were also determined by applying hindcasted swell direction to observed buoy data lacking directional information.

Demographic Parameters of Yellowfin Croaker, Umbrina roncador (Perciformes: Sciaenidae), from the Southern California Bight
Daniel J. Pondella II, John T. Froeschke, Lynne S. Wetmore, Eric Miller, Charles F. Valle, and Lea Medeiros, 555

The yellowfin croaker, Umbrina roncador Jordan & Gilbert, 1882, is a common nearshore and surf-zone species in the southern California bight. Age was determined for individuals (n = 1,209) using annual increments in otoliths, and size at age was modeled using the von Bertalanffy growth curve (L∞ = 307.754 mm, k = 0.278 yr-1, t0 = -0.995 yr; maximum age = 15 yr). Females (L∞ = 313.173 mm, k = 0.307 yr-1, t0 = -0.771 yr) grew significantly faster and larger than males (L∞ = 298.886, k = 0.269 yr-1, t_0 = -1.072 yr). Age and growth modeling based upon otolith length (OL) and width (OW) measurements were assessed and were consistent with body measurements. Males and females were found in all size classes and in an overall 51:49 ratio that was not significantly different from a 50% sex ratio, suggesting that these fish are gonochores. Fish were reproductive during summer months, with gonadosomatic indices (females, 5.65%; males, 5.51%) consistent with group-spawning fishes. Data from two separate monitoring programs indicated that yellowfin croaker catch-per-unit-effort (CPUE) fluctuated appreciably from 1992 to 2006 on both spatial and temporal scales. CPUE also declined significantly in the latter years of these programs. Based on samples collected between 2003 and 2004, an estimate of overall annual total mortality was A = 0.4492, and instantaneous coefficient of total mortality was estimated at Z = 0.5964. Recruitment year classes were back calculated using annual survivorship. Year class strength was variable and declined significantly by the end of this study. Considering the high temporal and spatial variation in estimates of abundance and recruitment, coupled with the likelihood that these fish employ a probable group-spawning reproductive behavior, we recommend a cautious approach for the future management of this species.

Acropora (Anthozoa: Scleractinia) Reproductive Synchrony and Spawning Phenology in the Northern Line Islands, Central Pacific, as Inferred from Size Classes of Developing Oocytes
Jean C. Kenyon, 569

Little is known of the timing of reproduction in central Pacific coral populations near the equator. Oocyte pigmentation and size comparison with sizes of mature eggs reported in published literature were used to infer intra- and interspecific synchrony and probable spawning phenology in 15 species of Acropora from Palmyra and Kingman atolls in the northern Line Islands. Sampling at both atolls took place in March–April 2002 and 2004. Oocyte sizes were determined from microdissections of fixed, decalcified samples. The majority (91.2%) of samples (n = 209) were gravid, with high levels of fertility in most (84.3%) samples. Statistically discrete oocyte size classes could be distinguished in most taxa at each atoll in each year. These discrete oocyte size classes suggest that several episodes of spawning, involving multiple species, take place over 2 or 3 months beginning in early spring. These data, which are the first observations of coral reproductive synchrony in the Line Islands, support the results of other recent studies, suggesting that reproductive synchrony can be a feature of equatorial reef assemblages where the annual ranges of sea-surface temperature and tidal amplitude are small.

Seasonal Occurrence and Aggregation Behavior of the Sea Urchin Astropyga pulvinata (Echinodermata: Echinoidea) in Bahía Culebra, Costa Rica
Juan José Alvarado, 579

Between October 2003 and July 2005, aggregation behavior of the sea urchin Astropyga pulvinta Lamarck was studied in Bahía Culebra, Costa Rica. This sea urchin forms aggregations during part of the year and then disappears. I quantified the number of individuals present in a defined area each month, their aggregation behavior between day and night, and their size. Also, temperature and nutrient concentrations of the water were sampled. There were significantly more individuals in aggregations during the colder, upwelling season (December to April). Aggregations consisted of adult individuals that exploit food during the upwelling season. Moreover, these aggregations were used as a refuge by several fish species of high commercial value for the aquarium trade. These sea urchin populations could suffer as extraction of ornamental fishes and urchins increases. Their abundance and behavior should continue to be monitored as an indication of the ecological health of the community.

First Record of a Pearlfish, Carapus mourlani, Inhabiting the Aplysiid Opisthobranch Mollusc Dolabella auricularia
Peter W. Glynn, Ian C. Enochs, John E. McCosker, and Abigail N. Graefe, 593

Adult individuals of the pearlfish Carapus mourlani (Petit, 1934) occur commonly in the mantle cavity of the opisthobranch mollusc Dolabella auricularia (Lightfoot, 1786) in shallow marine waters of the Gulf of Chiriquí, Pacific Panamá. Nearly 30% of the molluscan hosts collected during the day on a coral reef contained one or two fish. Feeding observations of a captive fish as well as the intact condition of the host’s ctenidium and other internal organs suggest that C. mourlani is an inquiline commensal and not parasitic. Fish curl around the ctenidium during the day and capture microcrustaceans when the fish emerge from their host at night to feed. From low-light infrared video recordings, Carapus was observed to accurately grasp rapidly swimming amphipods in nearly total darkness and ingest them. This symbiotic relationship appears to benefit Carapus by allowing the fish to avoid predators during the day and to forage at night.

Two New Indo-Pacific Sand Lances of the Genus Ammodytoides (Perciformes: Ammodytidae)
John E. Randall and John L. Earle, 603

One new sand lance, Ammodytoides idai Randall & Earle, n. sp., is described from 10 specimens, 67.2–121.3 mm standard length, collected on sand substratum in the depth range of 8–25 m from the D’Entrecasteaux Islands, Papua New Guinea. It is characterized by 44–46 dorsal rays, 21–22 anal rays, 14–16 pectoral rays, 103–107 pored lateral-line scales, two or three small scales dorsally on the opercle, 5–6 + 21–23 gill rakers, 55–58 vertebrae; a series of black spots distally in the dorsal fin, and a blackish posterior border on the caudal fin (at least on adult males), broadening toward lobe tips. A second similar species, A. praematura Randall & Earle, n. sp., is described from a single 61 mm specimen from the Chagos Archipelago, differing in having 48 dorsal rays, 24 anal rays, black dots in the dorsal and anal fins, no submarginal black spots in the dorsal fin, and a curved blackish bar across each lobe of the caudal fin. A key is provided for the eight known species of Ammodytoides.

Association Affairs, 613

Index to Volume 62, 617

E. Alison Kay, the longest-serving editor of Pacific Science, was remembered yesterday at a service in the chapel of Punahou School, her alma mater. Two of her books published by UH Press are still in print: Shells of Hawaii (1991, with Olive Schoenberg-Dole) and A Natural History of the Hawaiian Islands: Selected Readings II (1994).

E. Alison Kay (1928–2008) was a malacologist, environmentalist, and professor at the University of Hawaiʻi. She was born in Eleele and grew up on the island of Kauai in the Territory of Hawaii, graduated from Punahou School in 1946, and obtained her first B.A. from Mills College in 1950. She then went on to earn another B.A. in 1952 and an M.A. in 1956 from Cambridge University as a Fulbright scholar before returning to the University of Hawaiʻi, where she completed her dissertation in 1957.

Her research focused on marine mollusks in the Indo-Pacific region, and she regularly offered a graduate course in taxonomy and systematics, and another in biogeography, emphasizing in particular the ecology and distribution of island mollusks. Her dissertation was on cowrie shells and Cypraea alisonae was named for her.

However, she believed strongly in general education, was assigned to the Dept. of General Science until 1982 (when she moved to Zoology), and was one of the most engaging lecturers in the large survey courses taught at the Varsity Theater off-campus. In later years, she taught a popular course in the natural history of the Hawaiian Islands, for which she edited a textbook in 1994. She also served at various times as dean of graduate students, as department head, and as a member of the Faculty Senate. She was the longest-serving editor in chief of the journal Pacific Science, overseeing almost every issue between 1972 and 2000.

Active in many environmental projects, she helped found the Save Diamond Head Association, conducted research on the effects of the atomic bomb in the Marshall Islands, and did pioneering research on micromollusks for biomonitoring. Her research on the ecology of opihi (limpets) helped shape state regulations limiting opihi collection.

This issue is available in BioOne.2

Tropical Island Ecosystems and Sustainable Development Symposium

Tropical Island Ecosystems and Sustainable Development

Biodiversity Research on Coral Reef and Island Ecosystems: Scientific Cooperation in the Pacific Region
Makoto Tsuchiya, René Galzin, and Neil Davies, 299

Pacific Island Forests: Successionally Impoverished and Now Threatened to Be Overgrown by Aliens?
Dieter Mueller-Dombois, 303

Indigenous forests in remote islands are generally impoverished of secondary successional tree species. After canopy disturbances, the same indigenous tree species seem to resume dominance by a process known as “autosuccession” or “direct succession.” Primary forest tree species are mostly colonizer species. Mature island forests are difficult to categorize as either pioneer, successional, or climax forests by their canopy species composition. Climax forests, which characterize mature forests in less-isolated areas, are typically of distinctly different canopy species composition than the pioneer forests. In central Canada, for example, pioneer pine forests are replaced in succession by mixed hardwood/softwood forests under exclusion of fire. This process is known as “normal replacement succession” or “obligatory succession.” Another well-known ecological concept distinguishes between “primary” and “secondary” forests in the continental tropics. Secondary forests are formed by fast-growing relatively short-lived second-growth species, which quickly assemble after major disturbances. It usually takes a long time for primary tropical rain forest trees to reappear in secondary forests. In contrast, primary island forests rarely include fast-growing indigenous canopy species that form such secondary forests in the continental tropics. Instead, secondary forests in islands are now made up mostly of introduced species. In this paper I attempt to evaluate alien plant invasion in remote islands in view of these concepts of ecological succession.

Differences in Associated Crustacean Fauna and Seasonality of Sexual Reproduction between Two Color Morphs of the Photosymbiotic Ascidian Didemnum molle (Ascidiacea: Didemnidae)
Takumi Fukuda and Euichi Hirose, 309

Photosymbiotic ascidians inhabiting subtropical waters tend to have gonads in spring and summer, whereas those in tropical waters are usually sexually mature year-round. We studied the seasonality of sexual reproduction in two populations of the photosymbiotic ascidian Didemnum molle (Herdman, 1886), sampling monthly for 12 months. Although the two populations were located only about 20 km apart, their color morphs were exclusively distributed: colonies of one population were always dark gray; those of the other population were mostly brown. The seasonality of sexual reproduction differed greatly between the populations (and thus between the color morphs). Sexual reproduction was limited to summer in the population with dark gray colonies, whereas the population with brown colonies possessed embryos with tails almost year-round. Moreover, the resident crustacean fauna in the colonies also differed between the populations. The microenvironment in each habitat may have caused these differences, but there may also be some physiological differences between the color morphs that affect the seasonality of sexual reproduction and the resident crustacean fauna.

Distribution and Possible Impacts of Toxic Organic Pollutants on Coral Reef Ecosystems around Okinawa Island, Japan
S. T. Imo, M. A. Sheikh, K. Sawano, H. Fujimura, and T. Oomori, 317

Organic pollutants have detrimental effects on the environment. In this study we evaluated the current status of contamination with organochlorine pesticides (OCPs), organo-tin compounds (OTCs), and polychlorinated biphenyls (PCBs) in the waters in and around Okinawa Island, Japan. Possible toxicological effects of these pollutants on marine life including corals are also discussed. Concentrations of total OCPs in river water were in the range of 1.02–56.4 ng liter-1. Among the OCPs, α-BHC, β-BHC, and aldrin were common in river water. OTCs detected in 30 samples of seawater were (mean ± SD) monobutyl tin (MBT), 0.44 ± 0.75 ng (Sn) liter-1; dibutyl tin (DBT), 1.32 ± 2.70 ng (Sn) liter-1; tributyl tin (TBT), 0.72 ± 2.90 ng (Sn) liter-1; monophenyl tin (MPhT), 0.04 ± 0.42 ng (Sn) liter-1; diphenyl tin (DPhT), 0.007 ng (Sn) liter-1; and triphenyl tin (TPhT), 0.013 ng (Sn) liter-1. Highest concentrations of TBT, 28.5 ng (Sn) liter-1 for water and 172 ng (Sn) g-1 dry weight for sediment, were detected in samples from Itoman Port. Concentrations of total PCBs were 0.05–0.28 ng liter-1 in open ocean and from 1.59 to 2.48 ng liter-1 in coastal waters. Overall, this study shows that the coral reef ecosystems and their adjacent environments around Okinawa Island are contaminated by toxic organic contaminants (OCPs, OTCs, and PCBs). Levels of these contaminants detected in some sites have exceeded the Environmental Quality Target (EQT), which may pose a risk to health of marine life.

Reproductive Biology and Early Development of Two Species of Sleeper, Eleotris acanthopoma and Eleotris fusca (Teleostei: Eleotridae)
Ken Maeda, Nozomi Yamasaki, Masashi Kondo, and Katsunori Tachihara, 327

Reproductive biology and early development of two species of sleepers, Eleotris acanthopoma Bleeker, 1853, and E. fusca (Forster, 1801), were investigated in streams on Okinawa Island in southern Japan. Gonadal examination and morphology of the genital papillae indicated that E. acanthopoma matured at a smaller body size (ca. 28 mm in standard length) than E. fusca (ca. 50 mm). Mature ovaries were composed of oocytes that could be categorized into two size classes. Larger females of both species had several hundred thousand developed oocytes in the larger size class and may spawn them at one or several consecutive spawning events. Egg masses of both species were found in habitats typically occupied by adults and were deposited, often sparsely, on the underside of objects. Form of the egg masses and morphology of eggs and newly hatched larvae of both species were almost identical. Eggs were a nearly spherical pyriform in shape, with the widest diameters measuring approximately 0.4 mm. Newly hatched larvae were very small (1.0–1.4 mm in notochord length) and undeveloped. The mouth opened and the eyes became pigmented 3 days after hatching, and all of their yolk was consumed 4 days after hatching. Reproductive strategies of both species were characterized by high fecundity through production of small eggs and small newly hatched larvae, with high fecundity likely to mitigate the presumed increased risk associated with widespread larval dispersal.

Flicker Light Effects on Photosynthesis of Symbiotic Algae in the Reef-Building Coral Acropora digitifera (Cnidaria: Anthozoa: Scleractinia)
Takashi Nakamura and Hideo Yamasaki, 341

Reef-building corals inhabit a variety of aquatic habitats with a range of light conditions. Because the coral host depends on photosynthetic products assimilated from endosymbiotic algae, reef-building corals have to cope with irradiance fluctuations on instantaneous to seasonal time scales. Underwater high-frequency light fluctuations resulting from the lens effect on the water surface are prominent in oligotrophic coral reef environments, a phenomenon known as flicker light. Effects of flicker light on endosymbiont photosynthesis of the reef-building coral Acropora digitifera (Dana, 1846) were evaluated with pulse amplitude modulation chlorophyll fluorometry. At supersaturating light intensities, photosynthesis was less inhibited by flicker light than by constant light. Reduction in photoinhibition by flicker light was pronounced at high water temperatures. Flicker light may strongly influence endosymbiont photosynthesis of corals inhabiting shallow reef habitats, especially during periods of strong solar irradiance and high water temperature.

Molecular Phylogeography of the Endemic Five-lined Skink (Plestiodon marginatus) (Reptilia: Scincidae) of the Ryukyu Archipelago, Japan, with Special Reference to the Relationship of a Northern Tokara Population
Masanao Honda, Taku Okamoto, Tsutomu Hikida, and Hidetoshi Ota, 351

Phylogenetic relationships were inferred for populations of the Ryukyu five-lined skink Plestiodon marginatus, a species showing an extraordinary distribution across the Tokara Tectonic Strait. Phylogenetic analyses of 809 base positions of the mitochondrial 12S and 16S rRNA genes supported collective divergence of the southern Tokara and northern Amami populations, which have been classified as P. m. oshimensis. A population from Nakanoshima, an island of the Tokara Group north of the Tokara Tectonic Strait, has the closest affinity with the Okinawajima population of P. m. marginatus rather than with the geographically closer southern Tokara and northern Amami populations. This result is concordant with that of a recent allozyme study and suggests an origin of the Nakanoshima population through long-distance dispersal from the Okinawa Island Group. Also, our results strongly suggest a closer relationship of a population of P. m. oshimensis from Okinoerabujima, a southern island of the Amami Group, with P. m. marginatus from Okinawajima than with the “consubspecific” southern Tokara and northern Amami populations. Both Nakanoshima and Okinoerabujima populations are usually referred to as P. m. oshimensis, and therefore our results indicate nonmonophyly of P. m. oshimensis in the current taxonomic arrangement.

Canopy Multilayering and Woody Species Diversity of a Subtropical Evergreen Broadleaf Forest, Okinawa Island
Akio Hagihara, S. M. Feroz, and Masatsugu Yokota, 363

Woody species diversity and the spatial distribution of trees in a subtropical evergreen broadleaf forest on a silicate substrate, Okinawa Island, were investigated to determine the forest’s architectural stratification. The forest stand consisted of four architectural layers. The values of Shannon’s index Hʹ and Pielou’s index Jʹ tended to increase from the top layer downward, except for the bottom layer. The lower layers contained many species relative to their smaller height ranges. High woody species diversity of the forest depended on small trees. This trend of species diversity was different from that of forest on a limestone substrate on Okinawa Island, where high woody species diversity depended on large trees. Conservation of small trees in the lower layers, especially the bottom layer, is indispensable to maintain diversity in Okinawan evergreen broadleaf forests. Castanopsis sieboldii (Mak.) Hatusima had the highest importance value in all layers, indicating that it is typically a facultative shade species as well as a climax species. The spatial distribution patterns of trees were found to be random in the lower three layers, but in the top layer clumping seemed to occur at three spatial scales. A high degree of overlapping in spatial distributions of trees among the layers suggested that light cannot penetrate easily into the lower layers. As a result, most species in the lower layers must be shade-tolerant. Mean weight index decreased from the top toward the bottom layer, and tree density increased from the top downward. This trend resembled the mean weight-density trajectory of self-thinning plant populations.

Biogeography of the Decapod and Stomatopod Crustacea of the Tropical Pacific: Issues and Prospects
Joseph Poupin, 377

Biogeographic patterns of the Crustacea (Decapoda and Stomatopoda) are given for the tropical Pacific, based on recent taxonomic studies combined with emergence of regional databases. Conclusive results are still difficult to obtain due to incomplete regional inventories and existence of complexes of sibling species with unclear taxonomic status. A time-series graph of the number of new records plotted against time is computed for several central Pacific islands (French Polynesia, Pitcairn, Easter Island, and Clipperton). It demonstrates that the fauna is still insufficiently known in those places. A biodiversity gradient is calculated for several taxa between West and East Pacific. The traditional decrease between Australia and French Polynesia is confirmed for higher taxa (Brachyura, Anomura), but at lower taxonomic levels it is not always verified (e.g., hermit crabs, Calcinus; crabs, Trapezia). A map is presented illustrating the following provisional biogeographic results: (1) cryptic endemic species recognized in the Marquesas Islands; (2) presence of a distinct faunistic province in the South Pacific, along the 25° S parallel, including Rapa and Easter Islands; (3) theoretical position of the border between the Indo-West Pacific (IWP) and East Pacific (EP) faunistic provinces (84 W on the seamounts of Sala y Gómez/Nazca and 110° W on Clipperton); (4) differences between Clipperton, with a mixed IWP-EP fauna (43% IWP versus 57% EP species), and the Galápagos, with obvious EP affinities (10% IWP versus 90% EP species).

Genetic Relationships among Species of Meretrix (Mollusca: Veneridae) in the Western Pacific Ocean
Ayako Yashiki Yamakawa, Masashi Yamaguchi, and Hideyuki Imai, 385

We compared allozymes at 12 loci in 12 populations of six species of Meretrix: M. lusoria (Japan, Korea, and Taiwan), M. petechialis (China and Korea), M. ovum (Thailand and Mozambique), M. lyrata (China), M. lamarckii (Japan), and Meretrix sp. A (Okinawa, Japan). Our allozyme results were generally consistent with the major groupings currently recognized within the genus based on morphological characters. However, we found two cryptic or undescribed species: Meretrix sp. A from Okinawa and M. cf. lusoria from Taiwan. The shell characters of Meretrix sp. A were similar to those of M. lamarckii, but the species was genetically distinct (Nei’s genetic distance D > 0.845) from all other species examined. The Taiwanese Meretrix population was morphologically indistinguishable from Japanese M. lusoria, although the genetic distance between the Taiwanese and Japanese populations showed a high degree of genetic differentiation (D > 0.386). Meretrix lusoria seedlings were introduced into Taiwan from Japan in the 1920s, and Japanese M. lusoria was previously thought to be established as a cultured stock. However, our results suggest that the Taiwanese population may represent a sibling or cryptic species of M. lusoria.

Systematic Review of Late Pleistocene Turtles (Reptilia: Chelonii) from the Ryukyu Archipelago, Japan, with Special Reference to Paleogeographical Implications
Akio Takahashi, Hiroyuki Otsuka, and Hidetoshi Ota, 395

The Quaternary terrestrial turtle fauna of the Ryukyu Archipelago was reviewed on the basis of recently excavated fossils, as well as literature information. As a result, five extinct species (four geoemydids [Cuora sp., Geoemyda amamiensis, Mauremys sp., and another species with undetermined generic and specific status] and one testudinid [Manouria oyamai]) were recognized from Late Pleistocene cave and fissure deposits. Two of the three turtles currently occurring in this archipelago (C. flavomarginata and G. japonica) were also recognized from comparable deposits on islands, including those where they do not occur at present. These records indicate that the terrestrial turtles of the Ryukyus were much more diverse during the Late Pleistocene than at present, and that extinction has occurred during the last few tens of thousands of years not only for those five fossil species but also for some island populations of the extant species. Distributions of three of the extinct species (G. amamiensis, Cuora sp., and the geoemydid [genus and species undetermined]), confined to the central Ryukyus, are concordant with the currently prevailing hypothesis of Ryukyu paleogeography, which assumes a relatively long isolation of this region and much more recent insularization of the southern Ryukyus. In contrast, distributions of the remaining two extinct species (Man. oyamai and Mau. sp.) must be explained by some ad hoc scenario or, otherwise, drastic modification of the current hypothesis.

Low Genetic Diversity of Oval Squid, Sepioteuthis cf. lessoniana (Cephalopoda: Loliginidae), in Japanese Waters Inferred from a Mitochondrial DNA Non-coding Region
Misuzu Aoki, Hideyuki Imai, Tohru Naruse, and Yuzuru Ikeda, 403

Genetic diversity and population structure of Japanese populations of the oval squid, Sepioteuthis cf. lessoniana, were compared with populations from Taiwan and Vietnam using nucleotide sequences of the mitochondrial DNA non-coding region 2. In total, 402 nucleotide sequences representing 242 individuals from Japanese waters (Ishikawa, Japan Sea coast of Honshu; Tokushima, eastern Shikoku; Nagasaki, western Kyushu; and Okinawajima and Ishigakijima Island, in the Ryukyu Archipelago) and the East and South China Seas (Keelung, northern Taiwan; Vietnam, Gulf of Tonkin) were examined. Among the 29 haplotypes recognized, haplotype no. 1 was shared by more than 75% of individuals from Japanese localities, whereas it was found in less than 13% of specimens from the East and South China Seas populations. Conversely, the East and South China Seas populations included more than 30% individuals with haplotype no. 2, whereas less than 10% of haplotype no. 2 individuals were from Japanese localities. The differences of haplotype and nucleotide diversities between pooled Japanese populations (0.2639, 0.23%) and the East and South China Seas populations (0.7900, 1.01%) indicate that S. cf. lessoniana from Japanese waters exhibits lower genetic diversity. An analysis of molecular variance between the Japanese populations and the East and South China Seas populations was highly significant. A minimum spanning tree of 29 haplotypes and an Unweighted Pair Group Method with Arithmetic mean (UPGMA) tree based on pairwise FST comparisons also supported the separation between Japanese and the East and South China Seas populations. We suggest that the Kuroshio Current physically limits gene flow and has thus caused the differences in genetic diversity among the populations examined.

Degree and Pattern of Gene Flow in Several Scleractinian Corals in the Ryukyu Archipelago, Southern Japan
A. Nishikawa, 413

Dispersal distance of planktonic larvae of coral reef organisms is influenced by their ecological characteristics and environmental factors such as current flow and physical structure of reefs. This study reviews the degree and pattern of genetic differentiation in scleractinian corals in the Ryukyu Archipelago, compared with other regions. Small-scale genetic heterogeneity, but broad-scale homogeneity, was detected in some species, including brooders and spawners in the Ryukyus. Comparison with other regions indicated that limited gene flow on a small spatial scale (i.e., self-recruitment) seemed to occur in many regions. However, the degree of gene flow over larger distances was complex and species-dependent. With an implication for conservation in the Ryukyus, the larval source hypothesis, which states that coral larvae were recruited from the Kerama Islands to the Okinawa Islands, was consistent with results illustrating high gene flow in some species. Thus, conservation of corals in the Kerama Islands is high priority. Detection of genetic breaks between the southern and central Ryukyus was not common among species. The genetic structure observed in corals is highly variable and depends on both species and spatial scale in the Ryukyus. In addition, the complex genetic structures of corals may be related to coral-specific destructive events, such as bleaching, outbreaks of crown-of-thorns starfish, and disease. Further studies will provide new insights and a more detailed view of the genetic structure of corals by using different markers (e.g., microsatellites) and approaches (assignment tests and clustering analysis), which will provide useful information for coral reef conservation.

Dietary Habits of the Introduced Cane Toad, Bufo marinus (Amphibia: Bufonidae), on Ishigakijima, Southern Ryukyus, Japan
Noriko Kidera, Nontivich Tandavanitj, Daehyun Oh, Nozomi Nakanishi, Aya Satoh, Tetsuo Denda, Masako Izawa, and Hidetoshi Ota, 423

We examined dietary habits of the introduced cane toad Bufo marinus at three sites representing different types of habitats (pond, forest, and rice paddy) on Ishigakijima Island, southern Ryukyus, Japan. Stomach contents analysis revealed that the toad mostly utilizes terrestrial arthropods, of which hymenopterans (mostly ants), adult coleopterans, hemipterans, and araneans dominated in the frequency of occurrence, hymenopterans in the numerical proportion, and larval lepidopterans, adult coleopterans, and larval dipterans in the volumetric proportion. Comparisons in taxonomic composition of the toad’s stomach contents and pitfall and sweeping net samples suggested ignorance or avoidance of Amphipoda by the toad. Our results suggest the possibility of considerable predation pressure of B. marinus upon the native arthropods, and ants in particular, on Ishigakijima Island.

A Framework for Assessing Impacts of Marine Protected Areas in Moorea (French Polynesia)
Thierry Lison de Loma, Craig W. Osenberg, Jeffrey S. Shima, Yannick Chancerelle, Neil Davies, Andrew J. Brooks, and René Galzin, 431

Marine Protected Areas (MPAs) have been promoted as effective management tools to protect biodiversity at local and global scales, but there remains considerable scientific uncertainty about effects of MPAs on species abundances and biodiversity. Commonly used assessment designs typically fail to provide irrefutable evidence of positive effects. In contrast, Before-After-Control-Impact (BACI) designs potentially remedy many of these problems by explicitly dealing with both spatial and temporal variation. Here, we document the historical context of implementation and the scientific assessment of MPAs recently established at eight sites around the island of Moorea, French Polynesia. In 2004, we designed and implemented a monitoring plan that uses a BACI-Paired Series (BACIPS) design to quantify the effect of the MPAs. Twice per year, we monitor fish, corals, and other benthic invertebrates at 13 sites (eight within MPAs and five outside MPAs) around Moorea, in three distinct reef habitats (fringing, barrier reef, and outer slope). We present statistical analyses of data collected during five surveys (July 2004 to July 2006), before the initiation of enforcement. We also assessed the potential of our program to detect future responses to the established MPA network. Our estimates of biomass for five categories of fishes (Acanthuridae, Chaetodontidae, Serranidae, Scaridae, and fisheries target species) within MPA sites generally track estimates in paired Control sites through time. Estimated statistical power to detect MPA effects (a 192% biomass increase within the MPA) was high at the MPA network scale but varied among taxonomic categories and reef habitats: power was high on the reefouter slope and lower in the lagoon, and generally high for acanthurids and chaetodontids. It did not vary significantly between sites. We discuss limitations of our approach (shared by all MPA assessments to date) and describe solutions and unique opportunities to redress these limitations in French Polynesia.

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Biology and Impacts of Pacific Island Invasive Species. 4. Verbesina encelioides, Golden Crownbeard (Magnoliopsida: Asteraceae)
Kathleen R. Feenstra and David R. Clements, 161

Verbesina encelioides (Cav.) Benth. & Hook. f. ex A. Gray, golden crownbeard, is a sunflower-like herbaceous annual plant ranging in height from 0.3 to 1.7 m with showy yellow flowers. It is native to the southwestern United States, the Mexican Plateau, and other parts of tropical America. Its invasive characteristics include high seed production (as many as 300–350 seeds per flower and multiple flowers per plant), seed dormancy, ability to tolerate dry conditions, and possible allelopathic effects. Disturbed areas with a relatively sandy substrate within warm, arid climate zones are vulnerable to invasion by V. encelioides. Verbesina encelioides is found on all of the main Hawaiian islands except Ni‘ihau but is particularly problematic on Midway and Kure Atoll, where it may threaten the habitat of nesting birds such as Laysan and black-footed albatrosses and Christmas and wedge-tailed shearwaters. Many other Pacific islands with similar habitats could be invaded by V. encelioides. The plant has become naturalized in many other U.S. states, parts of South America, the Bahamas, Cuba, the Dominican Republic, Puerto Rico, parts of Europe, Saudi Arabia, India, Ethiopia, Morocco, Botswana, Namibia, Israel, and Australia. It is a pest of various crops in the southern United States and India and is poisonous to sheep and cattle. Verbesina encelioides can be controlled via herbicides or mechanical means, but measures must be repeated due to the presence of persistent seed banks. Further research on V. encelioides is needed to understand its population dynamics, allelopathic properties, and impacts on natural ecosystems.

Exploiting Macrofauna Diadromy for Assessing Anthropogenic Impact in American Samoa Streams
L. M. Wade, F. S. Fanolua, A. M. Vargo, K. van Houte-Howes, E. Bardi, and D. L. Vargo, 177

Stream biomonitoring is increasingly used to identify and monitor changes in water quality, stream habitat, and even the surrounding watershed. An effective biomonitoring protocol must comprise attributes able to discriminate human-caused changes from natural variation. We attempted to identify such attributes for streams of American Samoa, which, in turn, might also have widespread applicability to other oceanic islands. Owing to the diadromous nature of the macrofauna, we assessed species richness, diversity, composition, dominance, and biomass of freshwater fishes, crustaceans, and mollusks in 50 m sections in midreaches of five streams with and five streams without anthropogenic influences at the estuarine reach. We electrofished for fishes and crustaceans, and we picked mollusks from stream substrates. We discovered that two species of neritid snails of the pan-Pacific genus Clithon were significantly more abundant in the midreach of streams undisturbed by human impacts at the estuarine reach, making them potentially useful bioindicators throughout the South Pacific.

Life History of the Red Spiny Lobster, Panulirus penicillatus (Decapoda: Palinuridae), in the Galápagos Marine Reserve, Ecuador
Alex Hearn and Juan Carlos Murillo, 191

The red spiny lobster, Panulirus penicillatus (Olivier, 1791), is exploited commercially in the Galápagos Marine Reserve by the local fishing sector. Catches and catch per unit effort have declined over the past few years, leading to concerns about sustainability of the fishery. This study supports the processes regarding the fishery management of P. penicillatus by determining its distribution and growth parameters. Nearly 3,000 lobsters were tagged during surveys carried out at 13 islands between 2000 and 2004. Sex ratio did not differ significantly from 1:1, and tagging returns showed little or no movement of individuals. Mean values with 95% confidence intervals for von Bertalanffy growth parameters were estimated to be K=0.201±0.004, L∞=16.91±0.183 (cm carapace length), and Ф’=4.14±0.019 for males; and K=0.264±0.02, L∞=12.34±0.40 (cm carapace length), and Ф’=4.99±0.06 for females. Natural mortality was 0.342 for males and 0.378 for females. These results, together with comparative estimates for red spiny lobster elsewhere, illustrate the geographical variability of growth among populations of P. penicillatus, which may occur within the archipelago itself.

A Comparison of Immature Green Turtles (Chelonia mydas) Diets among Seven Sites in the Main Hawaiian Islands
Karen E. Arthur and George H. Balazs, 205

Understanding resource acquisition and feeding ecology of threatened species is integral to their conservation because diet is intimately linked with growth rate and reproductive output. We examined diets of immature green sea turtles, Chelonia mydas (L.), from seven sites on the islands of Hawai‘i, O‘ahu, Moloka‘i, and Lāna‘i in January and August 2003. Diet analysis was based on 191 samples collected from 181 live green turtles by stomach lavage. These samples were identified and quantified using dissection microscopy and the principles of microstereology. Diet of green turtles in the Main Hawaiian Islands was dominated by red algae, and diet items most commonly encountered were Acanthophora spicifera (an introduced species), Hypnea sp., Pterocladiella sp., and Cladophora sp. Sea grasses (Halophila hawaiiana and H. decipiens) were an important component of diet in turtles from Kāne‘ohe Bay. Content of green turtle diets differed among foraging grounds, and these differences may provide an insight into previously documented differences in turtle growth rates among sites.

Evolutionary Lineages in Emballonura and Mosia Bats (Mammalia: Microchiroptera) from the Southwestern Pacific
D. J. Colgan and S. Soheili, 219

The microchiropteran bat family Emballonuridae is widely distributed in archipelagos of the southwestern Pacific, with especially strong representation of genera Emballonura and Mosia. DNA sequences from three segments of the mitochondrial genome were collected from four species of Emballonura and from M. nigrescens to investigate the relationship of genetic differentiation to archipelago biogeography. Specimens of each species formed monophyletic clades in maximum parsimony and Bayesian analyses. Mosia nigrescens was genetically distant to the other four species. The other four studied species formed a monophyletic clade composed of the pairs E. beccarii, E. serii and E. raffrayana, E. semicaudata. Clades within species were strongly concordant with geography, with only two counterexamples (E. semicaudata in Fiji and E. raffrayana in the Solomon Islands) to the general finding that each island’s population of a species constitutes a monophyletic clade. Genetic results do not agree with current subspecific designations within M. nigrescens. Samples from Woodlark, Alcester, and Manus Islands are phylogenetically closer to Papuan mainland samples than to Solomon Islands and New Ireland samples supposedly belonging to the same subspecies. Results suggest that Emballonura can establish populations across wide water barriers but does so infrequently. The isolating effect of water barriers is exemplified by the substantial genetic distinctiveness of Solomon Islands and New Ireland populations of both E. raffrayana and M. nigrescens. Absence from New Britain of E. beccarii, E. raffrayana, and E. serii (all known from New Ireland) may also reflect effects of water barriers if not due to collecting artifacts.

Relative Abundance and Distribution of Mariana Swiftlets (Aves: Apodidae) in the Northern Mariana Islands
Justine B. Cruz, Shelly R. Kremer, Gayle Martin, Laura L. Williams, and Vicente A. Camacho, 233

The endangered Mariana Swiftlet, Aerodramus bartschi (Mearns, 1909), occurs in its native habitat on only three islands worldwide—Guam, Saipan, and Aguiguan. It is locally extinct on the islands of Rota and Tinian, and numbers have declined on Guam. On Saipan and Aguiguan, the bird remains common. We present previously unpublished data from reports lodged with the Commonwealth of the Northern Mariana Islands Division of Fish and Wildlife combined with an analysis of arrival count data from surveys conducted regularly on Saipan (1985–2005) and opportunistically on Aguiguan (1985–2002). Direct counts of swiftlets arriving at nesting caves did not permit islandwide population estimates but provided an index useful for assessing relative abundance. On Aguiguan, swiftlets occurred in only a few of the available caves; the population was small, more densely concentrated than on the other islands, and relatively stable. On Saipan, swiftlet numbers declined for the first part of the monitoring period (1985–1992), then increased significantly (1998–2005), and now stand at their highest level (>5,000 birds) since 1985. Large between-year fluctuations, high variation in colony attendance patterns, and occasional abandonment and recolonization of some caves were evident during the 20-yr monitoring period. Of the potential constraints to the population, pesticide use, typhoons and supertyphoons, habitat alteration by feral animals, human disturbance in the nesting caves, and predation remain areas of concern. Conservation measures may have lessened some disturbance events and nest damage by cockroaches, while other measures, such as translocation, may improve the species’ chances of persistence.

A New Small-Bodied Species of Bavayia (Reptilia: Squamata: Diplodactylidae) from Southeastern New Caledonia
Aaron M. Bauer, Todd R. Jackman, Ross A. Sadlier, Glenn Shea, and Anthony H. Whitaker, 247

A new species of diplodactylid genus Bavayia, B. goroensis, is described from the Plaine des Lacs region of the Province Sud, New Caledonia. The new gecko is the smallest member of the Bavayia cyclura clade (49 mm snout-vent length) and, based on a molecular phylogeny, is basal within this group. It differs from other members of this group in its much smaller size, more gracile body, and lower number of precloacal pores and subdigital lamellae. The new species is known from only two locations, one of which is adjacent to extensive nickel mining operations. Because of its limited distribution and the direct and indirect threats posed by the proximity of mining to one of the populations, the species is here regarded as “Endangered.”

Role of Pro-Thrombolites in the Geomorphology of a Coastal Lagoon
D. A. Siqueiros-Beltrones, 257

Thrombolites are lithified biosedimentary structures generated by entrapment, precipitation, and binding of sediments promoted by growth and metabolic activity of cyanobacteria. Beaches of the coastal lagoon known as Ensenada de La Paz in Baja California Sur, México, are bordered by sedimentary formations of cyanobacterial origin identified as pro-thrombolites (incipient thrombolites) that represent a first record for the region and México. Observed thrombolithic structures show grains of varied sizes embedded within a fine-grain micritic matrix, which may be surrounded by medium-grain cementing micrite. Different degrees of consolidation occur—some crumble easily, whereas others require some manual force to break. These pro-thrombolites consist of platforms >20 cm thick and/or fragments of assorted sizes and forms. In some cases the structures have lithified, forming rocky plates (thrombolites). The extension and wide distribution of pro-thrombolites around the La Paz lagoon suggests that these structures could have determined its evolution from an original (primitive) cove into a lagoon. That is, the formation of pro-thrombolites through the entrapment and binding of sediments may have eventually altered water circulation, promoting sand sedimentation causing the formation of the El Mogote sand bar. Likewise, pro-thrombolites may have formed large extensions of headlands through accretion. Thus, several square kilometers of populated land around the La Paz lagoon may have thrombolithic origin.

Additional Marine Benthic Algae from Howland and Baker Islands, Central Pacific
Roy T. Tsuda, Peter S. Vroom, Isabella A. Abbott, Jack R. Fisher, and Kevin B. Foster, 271

Marine benthic algae from Howland Island and Baker Island were identified from collections made during earlier expeditions in 1924, 1935, and 1964, and during five separate expeditions between 1998 and 2004. Eighty-nine (six blue-green algae, 53 red algae, five brown algae, and 25 green algae) of the 99 species represent new records for the two islands. Forty-seven and 86 species are documented with voucher specimens from Howland Island and Baker Island, respectively. This study increases the total number of benthic marine algal species from the two islands to 104 species. A similar number (107 species) was previously reported from the nearby low coral islands in the Phoenix Group located 400 km to the southwest. Only 38% (39 of 104 species) of the algal species from Howland Island and Baker Island are reported from the Phoenix Group. The presence of Udotea palmetta Decaisne on Baker Island is of interest because the record is the first for this green algal genus in the central Pacific region.

Association Affairs, 291

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Population Characteristics of the Mangrove Crab Scylla serrata (Decapoda: Portunidae) in Kosrae, Federated States of Micronesia: Effects of Harvest and Implications for Management
Kimberly M. Bonine, Eric P. Bjorkstedt, Katherine C. Ewel, and Moses Palik, 1

Apparent declines in abundance of mangrove crabs Scylla serrata (Forsskål, 1755) in Kosrae, Federated States of Micronesia, have prompted concern regarding long-term persistence of this important cultural and economic resource. To support development of effective management strategies, we gathered basic biological information about mangrove crabs on this island, where S. serrata is the only mangrove crab species present. In particular, we were interested in understanding movement patterns and evaluating spatial variation in population structure. Many population characteristics, including estimated life span, ontogenetic shifts in habitat use, sex-specific allometric relationships, male-biased sex ratios, and evidence for limited (<2 km) alongshore movement, are similar to those reported elsewhere in the range of the species. Therefore, insights from S. serrata populations elsewhere might usefully inform management of the species on Kosrae. Moreover, information reported in this study, for which there is no ambiguity about species identification, has broader relevance. Spatial variation in size structure of the population appears to be driven by variable harvest pressure that reflects distribution of the human population and location of emerging commercial harvest operations. Effective management of mangrove crabs is therefore likely to benefit from application of size-based or sex-based restrictions on harvest and might usefully incorporate spatially explicit strategies, such as partial or complete reserves. Development and implementation of effective management will necessarily depend on cultural as well as scientific information.

Hawaiian Limpet Harvesting in Historical Perspective: A Review of Modern and Archaeological Data on Cellana spp. from the Kalaupapa Peninsula, Moloka‘i Island
Mark D. McCoy, 21

The isolated Kalaupapa region, Moloka‘i Island, Hawai‘i, offers archaeologists and ecologists a unique opportunity to study traditional Hawaiian limpet (‘opihi) (Cellana spp.) harvesting from the Proto-Historic Period (1650–1795), Early Historic Period (1795–1866), and the present day. In this study, archaeological collections, modern harvests, and field observations are used to describe a regular pattern of slightly larger mean limpet size in western shoreline harvests and an increase in average limpet size from the Proto-Historic Period to the present. Although further investigations are necessary to test alternative explanations, these results suggest (1) that shelter from ocean currents and trade winds may provide a microenvironment favorable to local limpet growth, and (2) a lessening of harvesting pressure concurrent with the massive depopulation of the study area after European contact. Future studies should focus on identifying possible ecological factors impacting average size, documenting changes in limpet size using specimens from archaeological deposits, and accounting for the impact of population change on marine resources in historic and prehistoric Hawai‘i.

Growth and Mortality of Coral Transplants (Pocillopora damicornis) along a Range of Sediment Influence in Maui, Hawai‘i
Gregory A. Piniak and Eric K. Brown, 39

Fragments of the lace coral Pocillopora damicornis (Linnaeus, 1758) were transplanted to four sites on the south-central coast of Maui, Hawai‘i, to examine coral growth over a range of expected sediment influence. Corals remained in situ for 11 months and were recovered seasonally for growth measurements using the buoyant weight technique. Average sediment trap accumulation rates ranged from 11 to 490 mg cm-2 day-1 and were greater at the wave-exposed reef site than at the protected harbor sites. Coral growth was highest at the donor site and was higher in the summer than in the winter. A stepwise linear regression found significant effects of sediment trap accumulation and light on growth rates, but the partial correlation coefficients suggest that these factors may be only secondary controls on growth. This study did not show a clear link between coral growth and sediment load. This result may be due, in part, to covariation of sediment load with wave exposure and the inability of trap accumulation rates to integrate all sediment effects (e.g., turbidity) that can affect coral growth.

Environmental versus Genetic Influences on Growth Rates of the Corals Pocillopora eydouxi and Porites lobata (Anthozoa: Scleractinia)
L. W. Smith, H. Wirshing, A. C. Baker, and C. Birkeland, 57

Reciprocal transplant experiments of the corals Pocillopora eydouxi Milne Edwards & Haime and Porites lobata Dana were carried out for an 18-month period from September 2004 to March 2006 between two back reef pools on Ofu Island, American Samoa, to test environmental versus genetic effects on skeletal growth rates. Skeletal growth of P. eydouxi showed environmental but not genetic effects, resulting in doubling of growth in Pool 300 compared with Pool 400. There were no environmental or genetic effects on skeletal growth of P. lobata. Pool 300 had more frequent and longer durations of elevated seawater temperatures than Pool 400, characteristics likely to decrease rather than increase skeletal growth. Pool 300 also had higher nutrient levels and flow velocities than Pool 400, characteristics that may increase skeletal growth. However, higher nutrient levels would be expected to increase skeletal growth in both species, but there was no difference between the pools in P. lobata growth. P. eydouxi is much more common in high-energy environments than P. lobata; thus the higher flow velocities in Pool 300 than in Pool 400 may have positively affected skeletal growth of P. eydouxi while not having a detectable effect on P. lobata. The greater skeletal growth of P. eydouxi in Pool 300 occurred despite the presence of clade D zooxanthellae in several source colonies in Pool 300, a genotype known to result in greater heat resistance but slower skeletal growth. Increased skeletal growth rates in higher water motion may provide P. eydouxi a competitive advantage in shallow, high-energy environments where competition for space is intense.

Life Cycle of Chrysaora fuscescens (Cnidaria: Scyphozoa) and a Key to Sympatric Ephyrae
Chad L. Widmer, 71

The life cycle of the Northeast Pacific sea nettle, Chrysaora fuscescens Brandt, 1835, is described from gametes to the juvenile medusa stage. In vitro techniques were used to fertilize eggs from field-collected medusae. Ciliated planula larvae swam, settled, and metamorphosed into scyphistomae. Scyphistomae reproduced asexually through podocysts and produced ephyrae by undergoing strobilation. The benthic life history stages of C. fuscescens are compared with benthic life stages of two sympatric species, and a key to sympatric scyphomedusa ephyrae is included. All observations were based on specimens maintained at the Monterey Bay Aquarium jelly laboratory, Monterey, California.

A Review of the Octocorallia (Cnidaria: Anthozoa) from Hawai‘i and Adjacent Seamounts: The Genus Narella Gray, 1870
Stephen D. Cairns and Frederick M. Bayer, 83

The nine Hawaiian species of Narella are revised, including the description of six new species. All species but one (N. ornata) are described and illustrated using SEM; all species are keyed and included in a detailed table of comparison. A brief history of octocoral taxonomic research in the Hawaiian Islands is presented, resulting in a total of 90 named species for this region, only five of which occur in shallow water. Specimens were collected from throughout the archipelago and adjacent seamounts, including Cross, Pensacola, Bishop, and Bushnell, from depths of 326 to 1,977 m.

Patterns of Nestedness in Remote Polynesian Ant Faunas (Hymenoptera: Formicidae)
Lloyd W. Morrison, 117

The entire ant faunas of remote Polynesian islands consist of introduced species. An important question concerning the assembly of Pacific island ant faunas is whether these species are a random assortment of the available species pool, or whether they exhibit highly ordered occurrence patterns (i.e., nested subsets of species). I evaluated nestedness for the ant faunas of two island groups in remote Polynesia: (1) the Hawaiian Islands, and (2) French Polynesia and the Cook Islands. Wilcoxon two-sample tests were used to analyze nestedness patterns for individual species and islands; the degree of nestedness for species assemblages and archipelagos was determined by combining tail probabilities of individual species and islands. Both island groups revealed highly significant nestedness at the level of the assemblage (a per-species approach) as well as the archipelago (a per-island approach). Considered individually, most species (73–95%) and most islands (89–100%) demonstrated significant nestedness. Instances of nonsignificant nestedness were frequently associated with low statistical power. These results reveal a strong deterministic element in the assemblage of remote Polynesian ant faunas. Dispersal opportunities along with presence of appropriate habitat type are likely the most important mechanisms underlying the observed patterns.

Characteristics of the Psidium cattleianum (Myrtaceae) Seed Bank in Hawaiian Lowland Wet Forests
Amanda L. Uowolo and Julie S. Denslow, 129

Psidium cattleianum Sabine (strawberry guava) is one of Hawai‘i’s most disruptive alien plants. Dense stands can suppress growth and establishment of native species, support high populations of crop-damaging fruit flies, and preclude restoration or management of native forests. Our research investigated factors affecting persistence of P. cattleianum seeds in lowland wet forest soils. We collected soil cores from four forested sites immediately after fruit fall and 6.5 months later. We found abundant germination of P. cattleianum seeds immediately after fruit drop. Soil collected under mature P. cattleianum clumps yielded 761 viable seeds/m2. We found no viable seeds 6.5 months after fruit drop. We evaluated seed longevity using seed bags buried below the litter layer that we retrieved after 28, 56, 196, and 365 days. Seeds either germinated or deteriorated rapidly after fruit drop; after 28 days, 22.3% of the buried seeds were viable and there were no viable seeds at 196 days. Predator effects were assessed using trays with a known number of seeds with and without predator exclosures. After 28 days, 37% of the seeds in the open trays were damaged by predators. The lack of a persistent seed bank likely is due to a combination of rapid, high germination rates, postdispersal seed predation, and seed mortality. We suggest that chemical or mechanical control efforts would be most efficient and effective if conducted at least 3 months after the fruiting season, when the vast majority of seeds have either germinated or died.

Subfossil Land Snail Fauna (Mollusca) of Central Chichijima, Ogasawara Islands, with Description of a New Species
Satoshi Chiba, Tetsuro Sasaki, Hajime Suzuki, and Kazuo Horikoshi, 137

The fossil record provides useful information to estimate what island communities were like before human colonization. We examined the species composition of the subfossil land snail fauna of dune deposits at the Yatsuse River, central Chichijima, Ogasawara Islands, and compared it with the species recorded in Chichijima since the nineteenth century. The 22 species in the dune deposits included 13 species that are now extinct in Chichijima. Live specimens of 11 of these extinct species were recorded in the early twentieth century, but no living Mandarna pallasiana and Ogasawarana obtusa Chiba et al., n. sp., have ever been recorded. Age of the sediment, estimated by radiocarbon (14C) dating, was 720 years B.P., and it is possible that these two land snail species became extinct as a result of the impact of human colonization of the island, which started in 1830. Specifically, Ogasawarana obtusa, n. sp., became extinct before the start of taxonomic studies of the land snails of Ogasawara. The sample included Hawaiia minuscula, which is generally now considered a cosmopolitan species introduced from North America. This finding suggests that Hawaiia minuscula is not alien in Ogasawara but indigenous.

First Record of Fossorial Behavior in Hawaiian Leafroller Moth Larvae, Omiodes continuatalis (Lepidoptera: Crambidae)
Cynthia King and Daniel Rubinoff, 147

Larvae of the endemic Hawaiian leafroller moth, Omiodes continuatalis (Wallengren), were used in controlled exposure trials on the island of Maui, Hawai‘i, in May–August 2006, to examine effects of introduced parasitoids on native Hawaiian Lepidoptera. During the trials we observed O. continuatalis larvae burrowing up to 14 cm into the soil beneath plants on which they were deployed. This discovery reflects the first record of fossorial behavior not associated with pupation in larvae of Hawaiian Omiodes and suggests how O. continuatalis, a species once listed as extinct by the U.S. Fish and Wildlife Service, may persist despite intense pressure from introduced biological control agents.

This issue is available in Project Muse and in BioOne.2

Biology and Impacts of Pacific Island Invasive Species. 3. The African Big-Headed Ant, Pheidole megacephala (Hymenoptera: Formicidae)
James K. Wetterer, 437

In the Pacific region, the African big-headed ant, Pheidole megacephala, is now widespread in tropical areas; populations are also found at higher latitudes in Australia, New Zealand, and Japan. On most inhabited tropical islands in the Pacific, P. megacephala is well known as a household and agricultural pest. Because P. megacephala does not attack humans, this species is often not recognized as an important threat. The negative ecological impact of P. megacephala, however, may be greater than that of any other invasive ant species. In areas where it occurs at high density, few native invertebrates persist. Loss of invertebrate species that serve key functions in the natural community (e.g., important prey species) may have cascading effects leading to the subsequent loss of additional species. Pheidole megacephala tends to thrive in open, disturbed habitats with weedy vegetation that can support high densities of plant-feeding Hemiptera, which these ants tend for honeydew. Before 1900, P. megacephala was known in the Pacific region only from Aru Island (Indonesia) and Hawai‘i. By the 1930s, it was found through much of Pacific Asia, Melanesia, and Polynesia, but it was not collected in Micronesia until 1950. Currently P. megacephala is known from virtually every tropical island group in the Pacific but not from many islands within the groups, particularly uninhabited islands. Quarantine efforts might be successful in keeping P. megacephala off these islands. Because P. megacephala does not commonly dominate areas with intact natural vegetation, setting aside relatively undisturbed habitat on inhabited islands may also be effective in protecting native invertebrates from attack by this ant.

Distribution of Parmarion cf. martensi (Pulmonata: Helicarionidae), a New Semi-Slug Pest on Hawai‘i Island, and Its Potential as a Vector for Human Angiostrongyliasis
Robert G. Hollingsworth, Rachel Kaneta, James J. Sullivan, Henry S. Bishop, Yvonne Qvarnstrom, Alexandre J. da Silva, and David G. Robinson, 457

The semi-slug Parmarion cf. martensi Simroth, 1893, was first discovered on O‘ahu, Hawai‘i, in 1996 and then on the island of Hawai‘i in 2004. This species, which is probably native to Southeast Asia, is abundant in eastern Hawai‘i Island, reportedly displacing the Cuban slug, Veronicella cubensis (Pfeiffer, 1840), in some areas. A survey in July–August 2005 found P. cf. martensi primarily in the lower Puna area of Hawai‘i Island, with an isolated population in Kailua-Kona (western Hawai‘i Island). It is now established in commercial papaya plantations, and survey participants reported it as a pest of lettuce and papaya in home gardens. Survey respondents considered P. cf. martensi a pest also because of its tendency to climb on structures where it deposits its feces and because of its potential to transmit disease. Individuals of this species were found to carry large numbers of infective third-stage larvae of the nematode Angiostrongylus cantonensis (Chen, 1935), the causative agent of human angiostrongyliasis and the most common cause of human eosinophilic meningoencephalitis. Using a newly developed polymerase chain reaction test, 77.5% of P. cf. martensi collected at survey sites were found infected with A. cantonensis, compared with 24.3% of V. cubensis sampled from the same areas. The transmission potential of this species may be higher than that for other slugs and snails in Hawai‘i because of the high prevalence of infection, worm burdens, and its greater association with human habitations, increasing the possibility of human-mollusk interactions.

Recent Records of Alien Anurans on the Pacific Island of Guam
Michelle T. Christy, Craig S. Clark, David E. Gee II, Diane Vice, Daniel S. Vice, Mitchell P. Warner, Claudine L. Tyrrell, Gordon H. Rodda, and Julie A. Savidge, 469

Eight anuran species were recorded for the first time in Guam in the period May 2003–December 2005, all apparently the result of arrivals to the island since 2000. Three of the eight species (Rana guentheri, Polypedates megacephalus, and Eleutherodactylus planirostris) had well-established breeding populations by 2005. A further three (Fejervarya cf. limnocharis, Fejervarya cancrivora, and Microhyla pulchra) were recorded from a number of individuals, but it is not known whether these species have established breeding opulations. Two species (Kaloula pulchra and Eleutherodactylus coqui) appear to be incidental transportations to the island that have not established. Before 2003, five anuran species, all introductions, had been recorded from Guam. Three of these, Polypedates leucomystax, Pseudacris regilla, and Kaloula picta, were detected on Guam in incoming cargo but destroyed. Two species established: Bufo marinus was deliberately introduced and the Australian hylid Litoria fallax was probably an accidental introduction. Successful establishment of anurans on Guam has increased the risk of frog introductions to nearby islands. By providing additional food sources for the brown tree snake (Boiga irregularis), anuran introductions have increased the chance that B. irregularis might substantially increase in numbers and in turn increase the risk of the snake being accidentally transported to other islands.

Arthropod Surveys on Palmyra Atoll, Line Islands, and Insights into the Decline of the Native Tree Pisonia grandis (Nyctaginaceae)
Alex T. Handler, Daniel S. Gruner, William P. Haines, Matthew W. Lange, and Kenneth Y. Kaneshiro, 485

Palmyra Atoll, in the Line Islands of the equatorial Pacific, supports one of the largest remaining native stands of Pisonia grandis forest in the tropical Pacific Ocean. In 2003, we surveyed terrestrial arthropods to document extant native and introduced species richness, compare these lists with historical records, and assess potential threats to native species and ecosystem integrity. In total, 115 arthropod taxa were collected, bringing the total number of taxa recorded since 1913 to 162. Few native species were collected; most taxa were accidental introductions also recorded from the Hawaiian Islands, the presumed main source of introductions to Palmyra. The overlap with previous historical surveys in 1913 and 1948 was low (<40%), and new species continue to establish, with one species of whitefly reaching pest status between 2003 and 2005. We observed numerous dead or dying large Pisonia grandis, and the green scale Pulvinaria urbicola (Coccidae) was particularly abundant on trees of poor health. Abundant introduced ants, particularly Pheidole megacephala, tended this and other hemipterans feeding on both native and introduced plants. We hypothesize that the Pheidole-Pulvinaria facultative mutualism is causing the decline of Pisonia grandis. Because of the unique properties of Pisonia grandis forest on oceanic atolls, its importance for nesting seabirds, and its alarming global decline, immediate conservation efforts should be directed at controlling introduced Hemiptera and disrupting their mutualisms with nonnative ants on Palmyra Atoll.

Scale and Benthic Composition Effects on Biomass and Trophic Group Distribution of Reef Fishes in American Samoa
Marlowe G. Sabater and Saolotoga P. Tofaeono, 503

We determined spatial patterns in distribution and biomass of 163 fish species in nearshore waters around Tutuila Island, American Samoa. Visual surveys of reef fishes along 30 by 5 m belt transects were conducted using a hierarchical nested design at five spatial scales from individual transects to tens of kilometers, allowing assessment of broad geographic patterns. Benthic cover data were derived from video transect surveys to test the relationship between habitat and distributions of reef fishes. We found that fish biomass, density, and numerical abundance in American Samoa are dominated by herbivores from relatively few species in the families Acanthuridae and Scaridae. Subsets of carnivore species covaried positively with live coral, algae, and coralline algae cover. Herbivores, in contrast, covaried positively with filamentous algae and coralline algae (i.e., their foods). Biomass of fishes at different trophic categories was associated with higher abundance of food material and habitat availability. Significantly higher biomass occurred along the south shore of Tutuila and at reefs with greater exposure to wave energy, such as topographic points, despite the occurrence of lower live coral cover. Significant variations in fish biomass occurred at large spatial scales, specifically at habitat and exposure levels. Variations at these scales were apparently driven by association of the most dominant trophic group with its food source and the extent but not the quality of habitat.

Life History and Courtship Behavior of Black Perch, Embiotoca jacksoni (Teleostomi: Embiotocidae), from Southern California
Bridgette Froeschke, Larry G. Allen, and Daniel J. Pondella II, 521

The black perch, Embiotoca jacksoni Agassiz, 1853, is a common reef fish associated with nearshore marine habitats of California, with the majority of the population occurring within the Southern California Bight. Black perch were collected throughout southern California from Santa Barbara to Carlsbad, including Santa Catalina Island, to determine their physical characteristics, growth, sex ratio, periodicity of reproduction, and length of gestation. Courtship observations were conducted using scuba along the King Harbor Breakwater in Redondo Beach, California, from January 2004 to December 2005 to verify periodicity of courting and associated reproductive behaviors. Specimens captured ranged from 75 to 220 mm standard length and from 18 to 487 g in total body weight. Seven age-classes were determined by otolith aging, with the growth rate tapering off after age-class one. Seventy percent of the individuals captured were from age-classes one to three. Growth rates did not differ between sexes. Mean monthly gonosomatic indexes for males peaked from July to November, with the highest mean occurring in October. Gestating females were found from December to May, with youngest gestating females being in age-class one. Courtship behaviors were observed within aggregations and in pairs from July to November, with males being the primary aggressors. Courtship postures occurred along the base of the reef, with pairs departing into caves for copulation. This study suggests that the black perch population within the Southern California Bight has different life history characteristics and reproductive timing than those in northern California.

Vertical Distribution of Fish Larvae and Its Relation to Water Column Structure in the Southwestern Gulf of California
L. Sánchez-Velasco, S. P. A. Jiménez-Rosenberg, and M. F. Lavín, 533

The seasonal evolution of vertical distribution of fish larvae and its relationship with seasonal stratification, as measured by a quantitative stability parameter, were analyzed for a region off Bahía de La Paz in the southwestern Gulf of California. Samples were obtained with an opening-closing net (505 μm) in 50-m depth strata from surface to 200-m depth in May, July, and October 2001 and February 2002. Significant differences in total larval abundance and in dominant species (mesopelagic and epipelagic) were found among strata from May to October. More larvae were found in maximum-stability strata (from 16 ± 5 to 48 ± 17 m depth) than below the pycnocline (from 100- to 150-m depth). In February, the 100-m-deep surface mixed layer had a weak pycnocline at its base, and no significant difference was found. Results show that vertical distribution of fish larvae in this area depends mainly on the seasonal evolution of the water column structure, with most fish larvae in the pycnocline, at the most stable stratum of the water column.

Shallow-Water Sea Anemones (Cnidaria: Anthozoa: Actiniaria) and Tube Anemones (Cnidaria: Anthozoa: Ceriantharia) of the Galápagos Islands
Daphne Gail Fautin, Cleveland P. Hickman Jr., Marymegan Daly, and Tina Molodtsova, 549

We provide the first inventory of members of orders Actiniaria (sea anemones sensu stricto) and Ceriantharia (tube anemones) from the Galápagos Islands. Based on observations and collections at 48 localities throughout the archipelago that span nearly a decade, we report on eight species of actiniarians (representing families Actiniidae, Actinostolidae, Aiptasiidae, Hormathiidae, and Isophelliidae) and two of cerianthids (in families Arachnactidae and Botrucnidiferidae). We include live photographs and diagnostic features of the animals, as well as a key and map of their occurrence in the Galápagos. Two actiniarians and one cerianthid are resolved only to genus level; of those identified to species, three of the actiniarians and one of the cerianthids have an eastern Pacific distribution, one actiniarian appears to be endemic to the Galápagos Islands, and two actiniarians are broadly distributed in the Indo-West Pacific.

Vegetative and Reproductive Variability of Dictyota crenulata (Phaeophyta: Dictyotales) along the Central and Southwestern Gulf of California, México
María del Carmen Altamirano-Cerecedo and Rafael Riosmena-Rodríguez, 575

Dictyota crenulata J. Agardh is widely distributed throughout the Gulf of California. Comparative analyses of morphology, anatomy, and reproductive features of this species were conducted along the central western and southwestern regions of the Gulf of California. Thalli showed geographical variations in length and apical width. No differences were observed in anatomy of vegetative thalli or relative abundance of reproductive structures. Dictyota crenulata had unilayered or multilayered medullas in the basal region and in proliferations. Most thalli presented unilayered medullas in the middle section. Our observations indicate that number of medullary layers is indeed a phenotypically plastic character, in agreement with previously published results. Variations in thallus morphology such as proliferations and length are likely the result of environmental differences, also reflected in the reproduction of D. crenulata. The southwestern region had the highest percentage of all life cycle stages (female gametophytes and sporophytes, both 22%, and vegetative thalli, 14%). Our results demonstrate morphological variability in Dictyota crenulata across its distribution in the Gulf of California.

Checklist of Pacific Operculina (Convolvulaceae), Including a New Species
G. W. Staples, 587

A new species of Operculina (Convolvulaceae), O. polynesica Staples, is described from the Pacific. This brings to five the number of species known from Micronesia, Melanesia, and Polynesia. A key for identification is provided, nomenclature and distributions are summarized, and a list of specimens examined is included to aid herbarium curators in naming Pacific material.

Index to Volume 61, 599