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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.

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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

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Biology and Impacts of Pacific Island Invasive Species. 2. Boiga irregularis, the Brown Tree Snake (Reptilia: Colubridae)
Gordon H. Rodda and Julie A. Savidge, 307

The Brown Tree Snake, Boiga irregularis (Merrem, 1802), was accidentally transported to the island of Guam shortly after World War II. Over the following two decades it spread throughout the island with little public or professional recognition of its extent or impacts. This secretive nocturnal arboreal snake occurs in all habitats on Guam, from grasslands to forests. Under the right conditions, it is capable of high rates of reproduction and population growth. The Brown Tree Snake caused the extirpation of 13 of Guam’s 22 native breeding birds and contributed to the extirpation of several species of native bats and lizards. Guam’s 12 forest birds were especially impacted, with 10 species eliminated and the other two severely reduced. In addition, the snake continues to substantially impact domestic poultry, pets, the island’s electrical power infrastructure, and human health. To protect other vulnerable Pacific islands, the U.S. government annually spends several million dollars inspecting cargo outbound from Guam to exclude Brown Tree Snakes. Cargo destinations most at risk are in Micronesia, especially the Northern Mariana Islands, but Guam also has direct air transportation links to Hawai‘i that will soon be supplemented with direct ship traffic. Ultimately, all Pacific islands are at risk but especially those obtaining cargo through Guam.

Human Impacts on the Nearshore Environment: An Archaeological Case Study from Kaua‘i, Hawaiian Islands
Alex E. Morrison and Terry L. Hunt, 325

Archaeology provides a long-term framework to document prehistoric resource use and habitat modification. Excavation at Nu‘alolo Kai, Kaua‘i, yielded a large, well-preserved shellfish assemblage. Analysis determined the susceptibility of mollusk communities to human foraging pressures in the past. Some coral reef and intertidal species, such as Turbo sandwicensis and Strombus maculatus, declined in abundance as a result of heavy exploitation. In contrast, shoreline mollusk communities remained fairly stable through time. Archaeological research provides baselines for modern conservation efforts and fisheries management.

Soil Phosphorus and Agricultural Development in the Leeward Kohala Field System, Island of Hawai‘i
Molly Meyer, Thegn N. Ladefoged, and P. M. Vitousek, 347

The leeward Kohala Field System on the island of Hawai‘i was one of the most intensive pre–European contact dryland agricultural systems. Archaeological and soil analysis has documented changes in soil nutrients over time. Soils were collected under agricultural field walls of different relative ages within the Kohala Field System. These field walls preserved soil from the time of their construction (between ca. A.D. 1400 and 1800), so soil samples from underneath older field walls have been exposed to a shorter period of cultivation than the soils under more recent field walls. Total P and P : Nb ratios of these buried soils were greater under walls than in once-cultivated surface soils, and greater under older walls than under younger walls. These results suggest that precontact cultivation decreased soil P reserves in this intensive agricultural landscape.

Coral (Anthozoa: Scleractinia) Recruitment at Bahías de Huatulco, Western México: Implications for Coral Community Structure and Dynamics
R. A. López-Pérez, M. G. Mora-Pérez, and G. E. Leyte-Morales, 355

Over the past decades there has been an increasing awareness of community structure and dynamics in eastern Pacific coral reef systems, yet the processes producing these patterns are poorly known. We conducted a quantitative analysis of patterns of sexual and asexual recruitment through fragmentation at six localities in Huatulco, México. Between January 2001 and January 2002, sexual recruitment was evaluated by using terracotta tiles. Fragmentation was addressed twice using quadrats. Two hundred ninety-two corals (291 Porites panamensis Verrill, 1 Pocillopora sp.) were recruited to the settlement tiles. Changes in abundance of recruits among sites were determined by coral cover of P. panamensis at each area. Fragmentation was restricted to Pocillopora spp., and processes producing fragments had no connection with those promoting their reattachment and survival. Sexual and asexual recruitment patterns and potential survival asymmetries displayed by P. panamensis and Pocillopora spp. in the area are of capital importance in the occurrence of local communities and potentially of those of the entire eastern Pacific region. Sexual and asexual recruitment patterns suggest that recovery of frame-building corals following disturbance is highly species-specific. Recovery of P. panamensis following coral removal can be relatively fast, but greatly prolonged for Pocillopora; however, in communities with low to moderate disturbance where patches of Pocillopora were preserved reef recovery can proceed at a moderate to relatively fast pace following disturbance. Coordinated multidisciplinary and interinstitutional efforts, including genetic, histological, and ecological approaches, are necessary to determine unequivocally the processes controlling community structure and dynamics in the area.

Linckia multifora (Echinodermata: Asteroidea) in Rarotonga, Cook Islands: Reproductive Mechanisms and Ecophenotypes
Terry J. Crawford and Bruce J. Crawford, 371

In Rarotonga, Linckia multifora (Lamarck) exists in two forms: a blue gray type that is found on the reef intertidally and a red form that is found subtidally. Both types reproduce asexually by regeneration of autotomized arms, as well as sexually, but the relative potential for sexual reproduction varies greatly between these different sites. In the laboratory, reciprocal crosses of the blue gray intertidal form and the red subtidal form developed as successfully as the controls and were indistinguishable in morphology. In addition, both the blue gray intertidal form and the red subtidal form contain two different classes of haplotypes of the mitochondrial gene cytochrome oxidase subunit I (COI), which exhibit 12 fixed differences. These results suggest that L. multifora of Rarotonga has a dual origin and that the two different forms seen in the two environments belong to a single interbreeding population and may represent ecophenotypes.

Estimating Abundance of Reef-Dwelling Sharks: A Case Study of the Epaulette Shark, Hemiscyllium ocellatum (Elasmobranchii: Hemiscyllidae)
M. R. Heupel and M. B. Bennett, 383

Benthic reef sharks play an important role in reef ecosystems, but little is known about their abundance or population dynamics. Abundance of the epaulette shark, Hemiscyllium ocellatum (Bonnaterre), on Heron Island Reef, Great Barrier Reef, Australia, was examined via a mark-recapture study. A total of 496 sharks was tagged between July 1994 and August 1997 in a 0.25-km² area of reef flat, with 80 tagged sharks recaptured for a total of 102 recapture events. Captured individuals ranged in size from juveniles to adults (285–750 mm total length). Recaptured sharks were collected after 1–725 days at liberty and at distances of 0–329 m from their original capture point. The overall recapture rate was 20.6% with an estimated 17.5% tag loss. Population size was estimated using both closed and open population models. Closed population models produced various abundance estimates, with the Chao M(th) ranked best in model performance with an estimate of 2,224 sharks and 95% confidence intervals ranging from 1,730 to 2,916. Open population models produced lower estimates, with the Jolly D model producing an estimate of 559 individuals within the study site and confidence intervals ranging from 26 to 1,092. All models produced density estimations of 0.3 to 1.2 sharks per 100 m². Based on thorough examination of model assumptions and results, open population models appear to provide the best population estimate within the study area.

A Visual Sighting and Acoustic Detections of Minke Whales, Balaenoptera acutorostrata (Cetacea: Balaenopteridae), in Nearshore Hawaiian Waters
Shannon Rankin, Tom F. Norris, Mari A. Smultea, Cornelia Oedekoven, Ann M. Zoidis, Ethan Silva, and Julie Rivers, 395

Minke whales, Balaenoptera acutorostrata (Lacépède), have been considered a rare species in Hawaiian waters due to limited sightings during visual and aerial surveys. However, our research suggests that they are more common than previously considered. In spring 2005, a combined visual-acoustic survey of cetaceans in Hawaiian waters resulted in the sighting of a minke whale within 22 km of Kaua‘i. Minke whale vocalizations were also detected at several other locations near Kaua‘i and O‘ahu. These 2005 reports are the first from nearshore (<50 km) Hawaiian waters despite years of previous shipboard and aerial surveys. The lack of historical sightings is likely due to misidentification or the inability to detect these animals during poor sighting conditions. We recommend that future cetacean surveys in Hawaiian waters include a passive acoustic component to increase the likelihood of detecting minke whales.

Neosabellaria vitiensis, n. sp. (Annelida: Polychaeta: Sabellariidae), from Shallow Water of Suva Harbor, Fiji
Julie H. Bailey-Brock, D. W. Kirtley, Eijiroh Nishi, and Susanne M. J. Pohler, 399

A new species of the genus Neosabellaria Kirtley, 1994, is described from shallow-water locations of Suva Harbor, Fiji. Neosabellaria vitiensis Bailey-Brock, Kirtley, Nishi, & Pohler, n. sp., is a gregarious sabellariid; its tubes are constructed of sand and shell debris and form small “reefs” exposed during low tides. The new species is distinguished by the structure of opercular paleae in the middle row, which are shoe-shaped with circular tips, and paleae in the outer row, which have distal lateral teeth and denticulate median plume. Detailed morphological features of the new species are described and compared with other Pacific sabellariid species, particularly with most closely related N. clandestina (Menon & Sareen, 1966). Neosabellaria vitiensis is endemic to the Fiji Islands.

A New Species of the Genus Lepidodactylus Fitzinger (Squamata: Gekkonidae) from the Mortlock Islands, Chuuk State, Federated States of Micronesia
Donald W. Buden, 407

A new species in the Group II complex of the gekkonid lizard genus Lepidodactylus Fitzinger is described based on recently collected material from Namoluk Atoll, Mortlock Islands, Chuuk State, Federated States of Micronesia. Lepidodactylus oligoporus Buden, n. sp., is distinguished from other members of Group II by differences in the numbers of midbody scale rows (130–134), fourth-toe scansors (15–19), interorbital scales (34–35), and precloacal/femoral pore-bearing scales (12–15) and by the lack of cloacal spurs and the presence of a moderate amount of webbing between the toes. In body size and scutellation, it most resembles L. novaeguineae Brown & Parker from New Guinea and L. pulcher Boulenger from the Admiralty Islands.

Reptiles of Satawan Atoll and the Mortlock Islands, Chuuk State, Federated States of Micronesia
Donald W. Buden, 415

Twenty species of reptiles are recorded from the Mortlock Islands, Chuuk, Federated States of Micronesia. The eight geckos and eight skinks together comprise 80% of the herpetofauna; amphibians are absent. Most of the species are widespread in the west-central Pacific, but the recently described gecko Lepidodactylus oligoporus is known only from the type locality on Namoluk Atoll. Hemidactylus frenatus appears to be displacing Gehyra mutilata, which is common only on Namoluk Atoll, where H. frenatus is unrecorded. Five species of skinks of the genus Emoia are sympatric on Satawan Atoll. Partial habitat segregation was observed in three morphologically very similar species of Emoia, with E. cyanura being more frequently encountered in beach strand and other open, sun-exposed areas; E. caeruleocauda in shady forest; and E. impar in sun-dappled forest patches.

First Report of Gastrointestinal Helminths from the Wokan Cannibal Frog, Lechirodus melanopyga (Amphibia: Limnodynastidae) from Papua New Guinea
Stephen R. Goldberg, Charles R. Bursey, and Fred Kraus, 429

The initial gastrointestinal helminth list is established for Lechriodus melanopyga (Doria) from Papua New Guinea. Examination of the digestive tracts of 16 L. melanopyga from April–May (n = 14) and October (n = 2) revealed six helminth species: Digenea: Mesocoelium monas; Nematoda: Aplectana macintoshii, Cosmocerca novaeguineae, Oswaldocruzia bakeri, Abbreviata sp. (larvae in cysts); Acanthocephala: Acanthocephalus bufonis. Cosmocerca novaeguineae was present in the greatest numbers (171) and shared the highest prevalence (88%) with Acanthocephalus bufonis. Lechriodus melanopyga represents a new host record for each of these helminths. New Guinea is a new locality record for Mesocoelium monas and Acanthocephalus bufonis.

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Conservation Status of the Endemic Bees of Hawai‘i, Hylaeus (Nesoprosopis) (Hymenoptera: Colletidae)
Karl N. Magnacca, 173

The 60 species of native Hylaeus bees in the Hawaiian Islands are important pollinators in native ecosystems, but they have been almost completely ignored in conservation studies. The only previous assessment of the conservation status of the individual species was not based on recent collections. Here I report on conservation status of all known species, based on collections made from 1999 to 2002. Species are arranged into six categories according to degree of threat, and species considered to be threatened are discussed individually. Five species have not been collected recently from one or more islands from which they are historically known, seven are restricted to endangered habitat, 10 are considered to be very rare and potentially endangered, and 10 have not been collected recently and could be extinct. With such a high proportion of rare species and the importance of Hylaeus species as pollinators, further work on their ecology is needed.

Reproductive Biology of the Brown Tree Snake, Boiga irregularis (Reptilia: Colubridae), during Colonization of Guam and Comparison with That in Their Native Range
Julie A. Savidge, Fiona J. Qualls, and Gordon H. Rodda, 191

Since their introduction to Guam shortly after World War II, brown tree snakes, Boiga irregularis (Merrem), have seriously impacted the biota and human population of the island. Understanding the biology of this exotic species will likely be important to the success of control programs. We compared the reproductive biology of 782 B. irregularis caught on Guam during the 1980s with results from published studies of native-range populations. Average and maximum sizes of mature snakes on Guam were larger than those from Australian populations. The majority of female brown tree snakes matured at snout-vent lengths (SVLs) of 910–1,025 mm, and most males matured at SVLs of 940–1,030 mm on Guam. Based on growth rates from the early 1990s on Guam, sexual maturity is estimated to occur during a snake’s third or fourth year. Only one female (0.3%) in our data set had oviductal eggs. Clutch size was estimated at 4.3 (SD = 2.2), based on large vitellogenic ovarian follicle (≥30 mm in length) and oviductal egg counts. Unlike their Australian counterparts, the Guam population reproduced year-round. Our data offer insights into the likely reproductive patterns of brown tree snakes should they infest other islands in the Pacific region.

Consistent Frequency of Color Morphs in the Sea Star Pisaster ochraceus (Echinodermata: Asteriidae) across Open-Coast Habitats in the Northeastern Pacific
Peter Raimondi, Raphael D. Sagarin, Richard Ambrose, Christy Bell, Maya George, Steven Lee, David Lohse, C. Melissa Miner, and Steven Murray, 201

The sea star Pisaster ochraceus (Brandt, 1835) is among the most conspicuous members of northeastern Pacific rocky-shore fauna due to its dramatic color variation, ranging from bright yellowish orange to brown to deep purple. Despite a large body of ecological and developmental biology information on P. ochraceus, few studies have rigorously examined color patterns or their causes across its geographic range. We used thousands of observations of sea star color and size taken from southern California to northern Oregon to show that the frequency of orange sea stars is approximately 20% with little variation across a broad latitudinal band. However, the frequency of orange sea stars in a population increases with the size of the animals in most populations. We consider several alternative hypotheses for these color patterns but find that the most parsimonious explanation is that adult color is a selectively neutral genetic trait that expresses itself ontogenetically. These novel findings point to the need for renewed study of the basic biology of this key ecological species.

Interspecific Spawning between a Recent Immigrant and an Endemic Damselfish (Pisces: Pomacentridae) in the Hawaiian Islands
Karen P. Maruska and Kimberly A. Peyton, 211

The Indo-Pacific damselfish Abudefduf vaigiensis (Quoy & Gaimard, 1825) was first observed in the Hawaiian Islands in the early 1990s and is now clearly established as a breeding population in the Islands. Sightings of fish with color patterns intermediate between those of A. vaigiensis and the very similar endemic Hawaiian sergeant, Abudefduf abdominalis (Quoy & Gaimard, 1825), suggest that hybridization of the two has occurred naturally. This study provides direct evidence of crossbreeding from observations and video footage of two separate spawnings in nearshore waters of O‘ahu and a third spawning in a public aquarium display tank. Reproductive behaviors were similar in intra- and interspecific spawning. However, one important difference was the absence of courtship by the male A. abdominalis toward the female A. vaigiensis in the interspecific spawnings. Instead, the female A. vaigiensis initiated spawning and the male A. abdominalis remained to fertilize, guard, fan, and clean the hybrid clutch along with a previous clutch until the embryos hatched. Embryos collected from one hybrid clutch showed normal embryonic development and subsequently hatched to produce viable swimming larvae. These observations represent a rare example of interspecific spawning in the damselfish family (Pomacentridae) and an exceptional opportunity to study hybridization and introgression in a wild population of coral reef fishes.

Genetic Population Structure of the Hawaiian Alien Invasive Seaweed Acanthophora spicifera (Rhodophyta) as Revealed by DNA Sequencing and ISSR Analyses
Daniel C. O’Doherty and Alison R. Sherwood, 223

Acanthophora spicifera (Vahl) Børgesen is the most widespread and invasive alien macroalga on coral reefs throughout the main Hawaiian Islands. This alga disperses from harbors and ports to coral reefs throughout the state, producing high quantities of biomass that affect a wide range of reef flora and fauna. Population samples of A. spicifera from across the main Hawaiian Islands were collected and compared through two kinds of analyses: DNA sequencing (based on a variable region of the nuclear large subunit ribosomal RNA gene, and the mitochondrial cox 2-3 spacer region) and fragment techniques (Inter-Simple Sequence Repeats [ISSRs]). DNA sequencing revealed no variation for the two markers, even when collections from other areas of the Pacific and Australia were included. In contrast, ISSR analyses revealed highly structured Hawaiian populations of A. spicifera with a substantial range of both within- and among-population variation, with individual plants forming discrete clusters corresponding to geographic locality.

Modeling Streamflows and Flood Delineation of the 2004 Flood Disaster, Mānoa, O‘ahu, Hawai‘i
Aly I. El-Kadi and Eric Yamashita, 235

In October 2004 a flood caused extensive damage to the University of Hawai‘i (UH) campus and neighboring residential areas in Mānoa Valley, O‘ahu, Hawai‘i. This modeling study was aimed at streamflow evaluation and flood delineation for the area impacted by the flood. The study concluded that the HEC-1 model of the U.S. Army Corps of Engineers is suitable for simulating storm runoff response for the study area, considering the nature of small Hawai‘i watersheds, which generate hydrographs with steep rising and falling limbs. The curve-number method of the U.S. Soil Conservation Service is also suitable because it predicts reasonably well the main features of streamflow hydrographs, including runoff duration and time of peaks. To improve on accuracy, however, there is a need for better characterization of spatial rainfall distribution through measurements. A flood delineation model, which treats the flood as a hypothetical dam break, was used to predict the floodwater pathway, flood zone extent, maximum flood depth, and the time to reach that depth. The model predicted an upper value for storm total flow volume that would not cause flooding on the UH campus. Although not fully validated, the developed models can guide data-collection and decision-making processes. For example, the models demonstrated that it is possible to mitigate the flood through streamflow diversion and stream dredging, realignment, and lining. For efficient management, we recommend defining a new subwatershed of the Ala Wai basin (to be called the West Mānoa Watershed) that contains the university campus.

Endemic Land Snail Fauna (Mollusca) on a Remote Peninsula in the Ogasawara Archipelago, Northwestern Pacific
Satoshi Chiba, Angus Davison, and Hideaki Mori, 257

Historically, the Ogasawara Archipelago harbored more than 90 native land snail species, 90% of which were endemic. Unfortunately, about 40% of the species have already gone extinct across the entire archipelago. On Hahajima, the second-largest island and the one on which the greatest number of species was recorded, more than 50% of species are thought to have been lost. We report here the results of a recent survey of the snails of a remote peninsula, Higashizaki, on the eastern coast of Hahajima. Although the peninsula is small (∼0.3 km²) and only part is covered by forest (<0.1 km²), we found 12 land snail species, all of which are endemic to Ogasawara. Among these species, five had been thought to already be extinct on Hahajima, including Ogasawarana yoshiwarana and Hirasea acutissima. Of the former, there has been no record since its original description in 1902. Except for the much larger island of Anijima and the main part of Hahajima, no single region on the Ogasawara Archipelago maintains as great a number of native land snail species. It is probable that the land snail fauna of the Higashizaki Peninsula is exceptionally well preserved because of a lack of anthropogenic disturbance and introduced species. In some circumstances, even an extremely small area can be an important and effective refuge for threatened land snail faunas.

Odonata of Yap, Western Caroline Islands, Micronesia
Donald W. Buden and Dennis R. Paulson, 267

Fifteen species of Odonata are recorded from Yap, Micronesia—two Zygoptera and 13 Anisoptera. None is endemic to Yap. Hemicordulia lulico occurs elsewhere only in Palau, whereas most of the other species are widespread in the western Pacific and Indo-Australian regions. Macrodiplax cora and Tramea loewi, both recorded by Lieftinck in 1962, were the only species not encountered during this study; Tramea loewi remains known in Micronesia only from a single male collected in Yap by R. J. Goss in 1950. Six of the breeding species in Yap that are widespread in Indo-Australia occur no farther east in the Caroline Islands except possibly as unusual extralimital records in the cases of Agriocnemis femina and Neurothemis terminata; the four other species reaching only as far east as Yap are Anaciaeschna jaspidea, Agrionoptera insignis, Orthetrum serapia, and Rhyothemis phyllis. Orthetrum serapia is reported from Micronesia for the first time, although a very old single specimen record of O. sabina from Tobi Island may possibly pertain to O. serapia. The odonate fauna of the outer islands of Yap State is poorly known; only six species have been recorded from among four of the 15 island groups. In addition, Tramea transmarina euryale rather than T. t. propinqua was found to be the subspecies occurring in the Chuuk Islands, contrary to earlier publications.

A New Species of Cophixalus (Anura: Microhylidae) from Misima Island, Papua New Guinea
Stephen J. Richards and Paul M. Oliver, 279

Cophixalus misimae Richards & Oliver, n. sp., is described from low-altitude rain forest on Misima Island, Louisiade Archipelago, Papua New Guinea. It is a small (males 15.5–16.1 mm, females 19.3–19.6 mm) terrestrial species with a visible tympanum, a snout that is distinctly truncate in dorsal view, unwebbed toes, a dark brown lateral stripe, and a call consisting of a train of high-pitched pulses. It is the third species of Cophixalus known from the Louisiade Archipelago and is currently known only from Misima Island.

First Record of Vegetative Cells of Pyrodinium bahamense (Gonyalucales: Goniodomataceae) in the Gulf of California
Aída Martínez-López, Ana E. Ulloa-Pérez, and Diana C. Escobedo-Urías, 289

As part of an ongoing monitoring study of phytoplankton in coastal lagoons on the east side of the Gulf of California, Pyrodinium bahamense Plate, 1906 var. bahamense was collected in the Topolobampo–Santa Maria–Ohuira coastal lagoon system in the Gulf of California in May 2005. Average concentrations of P. bahamense were 100 cells liter^-1. This finding is the first observation of vegetative cells of this tropical species in the Gulf of California and represents its northernmost occurrence to date.

Metopograpsus oceanicus (Crustacea: Brachyura) in Hawai‘i and Guam: Another Recent Invasive?
Gustav Paulay, 295

The grapsid crab Metopograpsus oceanicus (Jacquinot, 1852) is recorded from the Hawaiian Islands for the first time; it appears to be established at least in Kāne‘ohe Bay on O‘ahu. I review the ecology of the species in Oceania and argue that it was introduced both to the Hawaiian Islands and Guam, likely by shipping traffic. A brief review of Metopograpsus in the Hawaiian Islands is also presented.

Association Affairs, 301

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Introducing a New Series: Biology and Impacts of Pacific Island Invasive Species
David R. Clements and Curtis C. Daehler, 1

Biology and Impacts of Pacific Island Invasive Species. 1. A Worldwide Review of Effects of the Small Indian Mongoose, Herpestes javanicus (Carnivora: Herpestidae)
Warren S. T. Hays and Sheila Conant, 3

The small Indian mongoose, Herpestes javanicus (E. Geoffroy Saint-Hilaire, 1818), was intentionally introduced to at least 45 islands (including 8 in the Pacific) and one continental mainland between 1872 and 1979. This small carnivore is now found on the mainland or islands of Asia, Africa, Europe, North America, South America, and Oceania. In this review we document the impact of this species on native birds, mammals, and herpetofauna in these areas of introduction.

Vegetation History of Laysan Island, Northwestern Hawaiian Islands
J. S. Athens, J. V. Ward, and D. W. Blinn, 17

Paleoenvironmental investigations were undertaken on Laysan Island in the remote Northwestern Hawaiian Islands to investigate its flora before historical observations. Substantial impacts occurred to the island as a result of late nineteenth- and early twentieth-century guano mining, commercial feather collecting, and denudation of vegetation by feral rabbits. An account of Laysan’s historically known vegetation is presented, followed by discussion of results from the investigation of a 16.41-m sediment core from Laysan’s central hypersaline lake. The 7,000-year pollen and seed record, besides indicating the former importance of Pritchardia palms on Laysan, showed the former presence of seven previously unknown taxa, only four of which could be identified. Diatom analysis indicated fresh to brackish lake water during the early Holocene, a finding supported by the mollusk assemblage. Diatom diversity gradually decreased over time until there is a near monoculture, with types indicating a gradual increase of salinity. Hypersaline conditions were first recognizable near the top of the sequence with the appearance of Artemia zooplankton. Generally wetter conditions seem to have characterized the island before about 5,150 yr B.P., with drier condi