A single panmictic population of endemic red crabs,Gecarcoidea natalis,on Christmas Island with high levels of genetic diversity

The red crab, Gecarcoidea natalis, is endemic to Christmas Island in the Indian Ocean and largely responsible for shaping the unique ecosystem found throughout the island’s rainforests. However, the introduction and establishment of supercolonies of the highly invasive yellow crazy ant, Anoplolepis gracilipes, has decimated red crab numbers over the last several decades. This poses a significant risk to the future conservation of G. natalis and consequently threatens the integrity of the unique island ecosystem. Here we undertook a population genetic analysis of G. natalis using a combination of 11 microsatellite markers and sequencing of the mitochondrial cytochrome oxidase subunit I gene from samples collected on Christmas Island as well as a single location from North Keeling Island (located approximately 900 km west of Christmas Island). The genetic results indicate that G. natalis is a single panmictic population on Christmas Island, with no spatial genetic structure or restricted gene flow apparent between sampled locations. Further, G. natalis from North Keeling Island are not genetically distinct and are recent immigrants from Christmas Island. The effective population size of G. natalis has likely remained large and stable on Christmas Island throughout its evolutionary history with relatively moderate to high levels of genetic diversity in microsatellite loci and mitochondrial haplotypes assessed in this study. For management purposes G. natalis can be considered a single panmictic population, which should simplify conservation efforts for the genetic management of this iconic island species.     

SEA-BIRD BREEDING SEASONS ON CHRISTMAS ISLAND,PACIFIC OCEAN*

Christmas Island, at 2° N, 157° W in the Central Pacific Ocean, supports 18 breeding species of sea-birds. Data on the breeding seasons collected by the authors and other visitors to the island are analyzed in individual species accounts. Population estimates are given, the distribution of each species on the island is outlined, and for some species information on the sexual and moult cycles of marked individuals is presented. The Discussion concerns the ultimate and proximate factors controlling breeding seasons and the role of social factors in maintaining breeding synchrony in sea-birds, the diversity of the breeding regimes exhibited on Christmas Island (summary and analysis in Fig. 2 and Table 5), and comparison of these regimes with those found in the birds of Ascension Island and the Galapagos Archipelago.     

Inter-island movements and population differentiation in a pelagic seabird

We used mark-resight data and amplified fragment length polymorphism (AFLP) markers to assess movements and gene flow between Central Pacific breeding colonies of the great frigatebird, Fregata minor. Of 715 adult frigatebirds marked on Tern Island and Johnston Atoll, 21.3% were resighted at other frigatebird colonies at least 582 km away. Mark-resight data indicated regular movement of males and females between Tern Island and Johnston Atoll (873 km apart), and less frequent movements to other islands; no birds marked on Tern or Johnston were seen on Christmas Island, but one was seen in the Philippines, 7627 km from where it was marked. Despite the regular occurrence of interisland movements, Bayesian analyses of AFLP data showed significant genetic differentiation between Tern Island and Johnston Atoll, and more pronounced differentiation between these two islands and the more distant Christmas Island. The AFLP profiles of three birds breeding on Tern Island fell within the profile-cluster typical for Christmas Island birds, both in a nonmetric multidimensional scaling analysis and in a population assignment test, suggesting dispersal events from Christmas Island to Tern Island. Several factors could explain the persistence of genetic structure despite frequent movements between colonies: many movements occurred during the nonbreeding season, many breeding-season movements did not involve mate-acquisition behaviours and individuals that do disperse may be selected against, as suggested by morphometric differences between colonies. The persistence of genetic structure among breeding colonies despite significant interisland movements suggests limits to the effectiveness of migration as a homogenizing force in this broadly distributed, extremely mobile species.     

Supercolony mosaics: two different invasions by the yellow crazy ant, Anoplolepis gracilipes, on Christmas Island, Indian Ocean

Invasive species are one of the main reasons for the ongoing global loss of biodiversity. Anoplolepis gracilipes is an invasive ant that has recently received significant attention due to its negative effect on the native fauna and flora of Christmas Island, Indian Ocean. This species has contributed to a drastic change in the structure of the Christmas Island rainforest through its negative impact on the island’s endemic red land crab, the dominant consumer on the islands forest floor. In this study, we investigate the population structure of A. gracilipes on Christmas Island in order to determine whether multiple introductions occurred on the island and how they correspond to known infestations. We genotyped 578 individuals collected from 50 nests across the Island. We identify two distinct subgroups in the population that represent two different supercolonies. These supercolonies are interspersed across the island, however both nuclear (microsatellites) and mitochondrial markers strongly suggest that there is no gene flow between the two colonies. Significant heterozygote excess within the entire sampling area, with all but one worker examined being heterozygous for all seven microsatellite loci, suggests an unusual reproductive system in these ants. Our results are consistent with recent sociogenetic findings in a population of A. gracilipes in Northern Borneo.     

Pleistocene morphology and Holocene emergence of Christmas (Kiritimati) Island, Pacific Ocean

 Christmas (Kiritimati) Island is an unusually large coral atoll, of which a large proportion of the surface is presently subaerial. Extensive outcrops of in situ branching Acropora corals, together with Porites microatolls, Tridacna, and other shallow marine biota, indicate that the present low-lying area of interconnecting lakes in the island interior formed as a reticulate lagoon. Radiocarbon dating indicates that these lagoonal reefs flourished between 4500 and 1500 radiocarbon years BP, and surveying confirms that sea level was 0.5–1.0 m above present at that time, with subaerial exposure resulting from Late Holocene emergence. Boreholes undertaken for a water resources survey of the island penetrated near-surface Pleistocene limestones on the northern, southern, and eastern sides of the island. These are highly weathered and fractured, and although aragonitic clasts are preserved, U-series dating indicates a Middle Pleistocene or older age. At one location flanking the Bay of Wrecks, an outcrop of limestone, with an erosional notch, 1–2 m above present sea level, yielded a U-series age of 130 ka, and is interpreted as Last Interglacial in age. In contrast to previous interpretations which have suggested that Christmas Island comprised an atoll superstructure that is entirely Holocene, or the layer-cake interpretation appropriate for many mid-ocean atolls, Christmas Island appears to have had a form similar to its present in the Middle Pleistocene or earlier. It has undergone karstification during lowstands. Interglacials, particularly the Last Interglacial and the Holocene, appear to have resulted in only a minor veneer of coral over older limestone surfaces. Christmas Island is considered characteristic of an atoll that has not experienced significant subsidence through the Late Quaternary. Accepted: 15 May 1998     

Ecology and behavior of Gecarcoidea natalis, the Christmas Island red crab, during the annual breeding migration.

The terrestrial crab Gecarcoidea natalis is endemic to the forests of Christmas Island but must migrate each year to the coast to breed. During 1993 and 1995, radio-tracking, mark and recapture, and counting methods were used to establish the routes, walking speeds, direction of travel, and destinations of migrating crabs, as well as crab numbers and distribution. The density of crabs ranged from 0.09 to 0.57 crabs per square meter, which gave a population estimate of 43.7 million adult crabs on the island. During the dry season the crabs were relatively inactive but on arrival of the wet season immediately began their migration. The crabs generally walked in straight lines, and most crabs from around the Island traveled toward the northwest shore instead of simply walking toward the nearest shore. The maximum recorded distance walked by a red crab in one day was 1460 m, but the mean was 680 m per day in 1993 and 330 m in 1995. Comparing the 1993 and 1995 study seasons, there was a 3-week difference in the timing of the start of the migration, but the spawning date was fixed by the lunar phase and took place 17 to 18 days after mating. In 1993, late rain prompted a "rushed" migration and crabs walked directly to their shore destinations; in contrast, in 1995 most crabs made stops of 1 to 7 days during the downward migration. By giving the crabs a chance to feed along the way and minimizing the time that the population was concentrated near the shore, these stops may be important in ensuring that the animals have enough food after the long dry season. Furthermore, this behavior implies that the crabs are able to judge how far away they are from the shore during the migration.     

NOTES ON THE BIRDS OF EASTER ISLAND

Information on the occurrence and breeding status of sea-birds at Easter Island has long been scanty and incomplete. In order to obtain accurate information and in particular to investigate reports of a supposed breeding site of a "brown-plumaged petrel", we spent three weeks on the island, including 12 days camping out on the uninhabited, waterless islet of Motu-Nui.
The petrel proved to be the Christmas Island Shearwater Puffinus nativitatis , whose breeding range is thus extended by nearly 2,000 miles. Ten other species of sea-birds were definitely identified and specimens, photographs and breeding records were obtained for most of them. It is shown that the bird species frequenting Easter Island are, like the human inhabitants, predominantly of Polynesian origin—in fact the American continents are not represented at all.
The paper opens with a brief outline of present-day conditions on the main island, and concludes with an account of the "bird-cult" rituals which are known to have been a feature of life in Easter Island for centuries.     

Impacts of an invasive ant species on roosting behavior of an island endemic flying‐fox

Introduced species can cause major disruptions to ecosystems, particularly on islands. On Christmas Island, the invasive yellow crazy ant (Anoplolepis gracilipes) has detrimental impacts on many animals ranging from the iconic red crabs (Gecarcoidea natalis) to the Christmas Island Thrush (Turdus poliocephalus erythropleurus). However, the full extent of its effects on the island's fauna is not yet known. In this study, we investigated the impact of the yellow crazy ants on the island's last native mammal: the Christmas Island flying‐fox (Pteropus natalis). This species has been described as a keystone species, but has recently experienced substantial population decline to the extent that it is now listed as Critically Endangered. We examined the impacts of the yellow crazy ants on the roosting behavior of the Christmas Island flying‐fox, and on its local and island‐wide distribution patterns. We showed that the crazy ants increased behaviors in the flying‐foxes that were associated with avoidance of noxious stimuli and decreased behaviors associated with resting. Roost tree selection and roost site location were not related to variation in the abundance of crazy ants on the island. Our results indicate that the crazy ants interfere with the activity budgets of the flying‐foxes. However, the flying‐foxes failed to relocate to ant‐free roost trees or roost sites when confronted with the noxious ant, suggesting that the flying‐foxes are either not sufficiently disturbed to override strong cultural attachment to roosts, or, are behaving maladaptively due to ecological naïveté.     

Red crabs in rainforest on Christmas Island,Indian Ocean: No effect on the soil seed bank

Red land crabs (Gecarcoidea natalis) are both abundant and widespread in intact rainforest on Christmas Island, Indian Ocean, occurring at densities of ~ 1 crab m^–2. We used an on-going exclusion experiment to test the hypothesis that through their activities as seed predators and soil disturbers, these omnivores are important determinants of the density and relative species abundance of seeds in the soil seed bank of undisturbed primary rainforest. After 6.5 y of continuous crab exclusion, there was no significant difference in the density of seeds in the top 3 cm of soil between the control (3671 ± 517 seeds/0.19 m²) and exclusion plots (3285 ± 801 seeds/0.19 m²), nor was there any significant treatment difference in relative species composition. There were also no significant treatment differences when Muntingia calabura, which accounted for 91.9% of all seeds in the seed bank, was excluded from the analyses. We attribute the lack of a significant impact of red crabs to their probable inability to handle the very small seeds which occurred in the seed bank, and the low level of soil disturbance at the study site. We conclude that the observed impact of red crabs on the abundance and diversity of successional species in natural light gaps on Christmas Island is due to their impact on germinating seedlings alone, rather than any additional impact on the soil seed bank.     

Recruitment dynamics in a rainforest seedling community: context-independent impact of a keystone consumer

The influence of keystone consumers on community structure is frequently context-dependent; the same species plays a central organising role in some situations, but not others. On Christmas Island, in the Indian Ocean, a single species of omnivorous land crab, Gecarcoidea natalis, dominates the forest floor across intact rainforest. We hypothesised that this consumer plays a key role in regulating seedling recruitment and in controlling litter dynamics on the island, independent of the type of vegetation in which it occurred. To test this hypothesis, we conducted crab exclusion experiments in two forest types on the island and followed the dynamics of seedling recruitment and litter processing for six years. To determine if these effects were likely to be general across the island, we compared land crab densities and seedling abundance and diversity at ten sites across island rainforest. Surveys across island rainforest showed that seedlings of species susceptible to predation by land crabs are consistently rare. Abundance and diversity of these species were negatively correlated to red crab abundance. Although red land crabs may be important determinants of seedling recruitment to the overstorey, differences in overstorey and seedling composition at the sites suggests that recruitment of vulnerable trees still occurs at a temporal scale exceeding that of this study. These “windows” of recruitment may be related to infrequent events that reduce the effects of land crabs. Our results suggest that unlike the context dependence of most keystone consumers in continental systems, a single consumer, the red land crab, consistently controls the dynamics of seedling recruitment across this island rainforest.     

Greta's Garbo: stranded seeds and fruits from Greta Beach, Christmas Island, Indian Ocean

Aim To describe the species composition of stranded seeds and fruits drifted by ocean currents to Christmas Island, Indian Ocean. Location Christmas Island, Indian Ocean. Methods Frequent visual searches along the strand line of the island's few accessible beaches over a 4-year period 1988–92, with most effort concentrated on Greta Beach, on the east coast. Results The collection contained not fewer than sixty-three species in forty-nine genera and twenty-nine families. Leguminous seeds were by far the most common (especially Caesalpinia bonduc (L.) Roxb., Dioclea spp., Entada spp., Erythrina spp. and Mucunagigantea (Willd.) DC.), but Calophyllum inophyllum L., Guettarda speciosa L., Hernandia ovigera L., Heritiera littoralis Aiton and Terminalia catappa L. were also common. Main conclusions Only about one-third of species recorded in the drift flora are native to the island, and most disseminules stranded on the island are probably not locally derived. The most likely distant sources of drift disseminules are probably the southern Indonesian islands and Sumatra, with most disseminules probably arriving via the Timor and Arafura Seas between Indonesia and Australia. However, some disseminules may originate from as far east as the Moluccas and the east coast of Kalimantan. The majority of species recorded in the drift flora are not native to the island, and yet some of these were encountered frequently and displayed a high degree of viability on arrival (e.g. Dioclea hexandra (Ralph) Mabb., Erythrina fusca Loureiro and Mucuna gigantea (Willd.) DC.). Several possible reasons for the failure of many drift species to establish on the island are discussed.     

Spatial dynamics of supercolonies of the invasive yellow crazy ant, Anoplolepis gracilipes, on Christmas Island, Indian Ocean

Key to the management of invasive species is an understanding of the scope of an invasion, the rate of proliferation and the rate at which invaded habitats become degraded. This study examines the spatial dynamics of high-density supercolonies of the invasive yellow crazy ant, Anoplolepis gracilipes , on Christmas Island, Indian Ocean, and the associated impacts at their boundaries. Since the early 1990s, A. gracilipes supercolonies have occupied over 30% of the 10,000 ha of rainforest on Christmas Island. Thirty-four discrete high-density supercolonies formed between 1989 and 2003, ranging in size across nearly three orders of magnitude from 0.9 to 787 ha. Supercolonies boundaries are diffuse, and ants were observed in low densities in some cases up to 200 m from the main high-density supercolony. The 13 boundaries examined were all dynamic over a 10–20 observation month period: nine boundaries expanded, and the maximum rate of spread was 0.5 m day⁻¹. Across boundary transition zones, between high-density supercolonies and intact rainforest, yellow crazy ants reduced other ant species richness, occupied red crab burrows and killed resident red crabs, which was the trigger for 'invasional meltdown' on Christmas Island. The highly variable and unpredictable nature of A. gracilipes boundaries poses a challenge for incorporation into a predictive framework, as well as for their management.     

Y-chromosome and mitochondrial DNA studies on the population structure of the Christmas Island community

Christmas Island is a remote Australian territory located close to the main Indonesian island of Java. Y-chromosome and mitochondrial DNA (mtDNA) markers were used to investigate the genetic structure of the population, which comprises communities of mixed ethnic origin. Analysis of 12 Y-chromosome biallelic polymorphisms revealed a high level of gene diversity and haplotype frequencies that were consistent with source populations in southern China and Southeast Asia. mtDNA hypervariable segment I (HVS-I) sequences displayed high levels of haplotype diversity and nucleotide diversity that were comparable to various Asian populations. Genetic distances revealed extremely low mtDNA differentiation among Christmas Islanders and Asian populations. This was supported by the relatively high proportion of sequence types shared among these populations. The most common mtDNA haplogroups were M* and B, followed by D and F, which are prevalent in East/Southeast Asia. Christmas Islanders of European descent were characterized by the Eurasian haplogroup R*, and a limited degree of admixture was observed. In general, analysis of the genetic data indicated population affinities to southern Chinese (in particular from the Yunnan Province) and Southeast Asia (Thailand, Malaysia, and Cambodia), which was consistent with historical records of settlement. The combined use of these different marker systems provides a useful and appropriate model for the study of contemporary populations derived from different ethnic origins.     

Linking El Niño,local rainfall,and migration timing in a tropical migratory species

Current climate models project changes in both temperature and precipitation patterns across the globe in the coming years. Migratory species, which move to take advantage of seasonal climate patterns, are likely to be affected by these changes, and indeed, a number of studies have shown a relationship between changing climate and the migration timing of various species. However, these studies have almost exclusively focused on the effects of temperature change on species that inhabit temperate zones. Here, we explore the relationship between rainfall and migration timing in a tropical species, Gecarcoidea natalis (Christmas Island red crab). We find that the timing of the annual crab breeding migration is closely related to the amount of rain that falls during a ‘migration window’ period prior to potential egg release dates, which is in turn related to the Southern Oscillation Index, an atmospheric El Niño‐ Southern Oscillation Index. As reproduction in this species is conditional on successful migration, major changes in migration patterns could have detrimental consequences for the survival of the species. This study serves to broaden our understanding of the effects of climate change on migratory species and will hopefully inspire future work on rainfall and tropical migrations.     

Invasion,establishment, and range expansion of two parasitic nematodes in the Canadian Arctic

Climate warming is occurring at an unprecedented rate in the Arctic and is having profound effects on host‐parasite interactions, including range expansion. Recently, two species of protostrongylid nematodes have emerged for the first time in muskoxen and caribou on Victoria Island in the western Canadian Arctic Archipelago. Umingmakstrongylus pallikuukensis, the muskox lungworm, was detected for the first time in 2008 in muskoxen at a community hunt on the southwest corner of the island and by 2012, it was found several hundred kilometers east in commercially harvested muskoxen near the town of Ikaluktutiak. In 2010, Varestrongylus sp., a recently discovered lungworm of caribou and muskoxen was found in muskoxen near Ikaluktutiak and has been found annually in this area since then. Whereas invasion of the island by U. pallikuukensis appears to have been mediated by stochastic movement of muskoxen from the mainland to the southwest corner of the island, Varestrongylus has likely been introduced at several times and locations by the seasonal migration of caribou between the island and the mainland. A newly permissive climate, now suitable for completion of the parasite life cycles in a single summer, likely facilitated the initial establishment and now drives range expansion for both parasites.     

Natural immigration of brushtail possums,Trichosurus vulpecula,onto Aroha Island,Kerikeri Inlet,Bay of Islands,New Zealand

Abstract

Aroha is a 5 ha island in Kerikeri Inlet, Bay of Islands, which is joined to the North Island mainland by a 200 m long causeway. Introduced Australian brushtail possums (Trichosurus vulpecula) spread into the general area only in the mid 1970s, and were first trapped on Aroha Island in June 1979. Since then possums have regularly crossed the causeway, and have been systematically trapped to keep the island free of them. Two hundred and sixty two possums have been trapped on Aroha Island to December 1990, and the annual catch has increased steadily since 1981. About equal numbers of males and females have been trapped. Most possums were caught in March-April and fewest in July-August. Captures on the island were clustered, particularly during the breeding seasons. Average body weight was 2.2 kg; 97% were greys and 3% blacks; the young of females trapped on the island had been born in all months except December to February. In colour, body weight, and breeding pattern the possums trapped on Aroha Island were similar to other possum populations from the northern North Island. Most of the possums invading Aroha Island were 1-2 years old: in contrast to findings from previous studies of possum dispersal, Arona Island was invaded year-round by young male and female possums equally. Some older animals may also have been attracted to the island by fruits available in an orchard there. Maintenance of the possum-free state of Aroha Island will require continuous management.     

Invasive ants disrupt frugivory by endemic island birds

Biological invasions can alter direct and indirect interactions between species, generating far-reaching changes in ecological networks that affect key ecological functions. We used model and real fruit assays to show that the invasion and formation of high-density supercolonies by the yellow crazy ant (YCA), Anoplolepis gracilipes, disrupt frugivory by endemic birds on Christmas Island, Indian Ocean. The overall handling rates of model fruits by birds were 2.2–2.4-fold lower in ant-invaded than in uninvaded rainforest, and pecking rates by two bird species declined by 2.6- and 4.5-fold, respectively. YCAs directly interfered with frugivory; their experimental exclusion from fruiting displays increased fruit handling threefold to sixfold, compounding indirect effects of ant invasion on resources and habitat structure that influence bird abundances and behaviours. This invasive ant, whose high densities are sustained through mutualism with introduced scale insects, rapidly decreases fruit handling by endemic island birds and may erode a key ecological function, seed dispersal. Because most other invasive ant species form expansive, high-density supercolonies that depend in part on association with hemipteran mutualists, the effects that we report here on avian frugivore–plant associations may emerge across their introduced ranges.     

广东省川山群岛开发利用生态风险评价

随着海洋经济的迅猛发展,海岛的开发利用程度不断加大,海岛生态环境遭到极大地冲击和破坏,导致海岛生态系统出现环境恶化、灾害加剧的趋势。广东省川山群岛,主岛为上川岛和下川岛,因岛上旅游娱乐和养殖用海等开发利用活动频繁,使其生态系统面临严峻的挑战。根据EPA提出的生态风险评价框架,分析了川山群岛的生态终点、生境类型和压力源状况,构建了一个用以描述川山群岛压力源、生境和评估终点间相互关系的概念模型,采用RRM模型思想进行了川山群岛开发利用风险评价。结果表明:上川岛属于开发利用的高风险区,主要风险区域是潮间带和近海两个生态子系统,其中以岩滩的开发利用压力最大,旅游娱乐和养殖用海是风险压力的主要来源,珍稀水禽是主要受威胁的物种;下川岛较上川岛的开发利用风险低,其岛陆、潮间带和近海3个生态子系统的开发利用压力都不大,养殖用海、旅游娱乐和城镇用地是风险压力的主要来源,也是威胁珍稀水禽和猛禽的主要压力源。基于此,从旅游管理、养殖用海管理等方面给出了上川岛和下川岛的风险管理对策,以期为海岛环境保护和可持续利用提供科学依据。     

Genetic Divergence of an Avian Endemic on the Californian Channel Islands

The Californian Channel Islands are near–shore islands with high levels of endemism, but extensive habitat loss has contributed to the decline or extinction of several endemic taxa. A key parameter for understanding patterns of endemism and demography in island populations is the magnitude of inter–island dispersal. This paper estimates the extent of migration and genetic differentiation in three extant and two extinct populations of Channel Island song sparrows (Melospiza melodia graminea). Inter–island differentiation was substantial (G''''_ST: 0.14–0.37), with San Miguel Island having the highest genetic divergence and lowest migration rates. Santa Rosa and Santa Cruz Island populations were less diverged with higher migration rates. Genetic signals of past population declines were detected in all of the extant populations. The Channel Island populations were significantly diverged from mainland populations of M. m. heermanni (G''''_ST: 0.30–0.64). Ten mtDNA haplotypes were recovered across the extant and extinct Channel Island population samples. Two of the ten haplotypes were shared between the Northern and Southern Channel Islands, with one of these haplotypes being detected on the Californian mainland. Our results suggest that there is little contemporary migration between islands, consistent with early explanations of avian biogeography in the Channel Islands, and that song sparrow populations on the northern Channel Islands are demographically independent.     

Factors Affecting Restoration of Halodule wrightii to Galveston Bay, Texas

Experimental restoration of Halodule wrightii (shoalgrass) to its former range on Galveston Island, Texas, began in April 1994. We tested the effects of site, planting density, water depth, and fertilizer addition on survival and growth through June 1996. Temperature, salinity, light transmittance, turbidity, and sediment properties at two restoration sites, Redfish Cove and Snake Island Cove, were similar to those in naturally occurring grassbeds in nearby Christmas Bay. Halodule survival, coverage, and new shoot densities were affected by site (significantly higher at Redfish Cove than at Snake Island Cove, which eventually failed), by planting density (significantly higher when planted on 0.25-m or 0.5-m centers rather than on 1.0-m centers), and by water depth (significantly higher when planted in relatively shallow water). Propagation (spreading from transplant units) was significantly greater from 0.25-m or 0.5-m center plantings but was not consistently affected by site or water depth. Fertilizer enhanced propagation but not survival. After two years, Redfish Cove produced belowground biomass similar to that observed in Christmas Bay, but aboveground biomass remained significantly less. Snake Island Cove plant mortality in September 1995 may have been presaged by low root-rhizome carbohydrate levels observed in October 1994, but causes remain unknown. Further restoration of Halodule to Galveston Bay is possible at selected sites, but structural equivalency will take longer than two growing seasons to achieve.