Published Papers - Reimer James Davis

Distribution of palytoxin in coral reef organisms living in close proximity to an aggregation of Palythoa tuberculosa

Aratake S.

Toxicon ( Toxicon )  111   86 - 90   2016.03 [ Peer Review Accepted ]

Type of publication: Research paper (scientific journal)

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Palytoxin is a strong marine toxin that was first isolated from the zoantharian Palythoa toxica and later from other species of the genus Palythoa. How the toxin gets into the animal remains an unsolved question. To study the specificity of palytoxin distribution, the toxin content of Palythoa tuberculosa and other organisms living in close association on a coral reef in Okinawa were analysed by mass spectrometry. In contrast to earlier reports, palytoxin was only detected in P. tuberculosa colonies. (C) 2016 Elsevier Ltd. All rights reserved.

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Green fluorescence of Cytaeis hydroids living in association with Nassarius gastropods in the red sea

Prudkovsky A.A.

PLoS ONE ( PLoS ONE )  11 ( 2 )   2016.02 [ Peer Review Accepted ]

Type of publication: Research paper (scientific journal)

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Green Fluorescent Proteins (GFPs) have been reported from a wide diversity of medusae, but only a few observations of green fluorescence have been reported for hydroid colonies. In this study, we report on fluorescence displayed by hydroid polyps of the genus Cytaeis Eschscholtz, 1829 (Hydrozoa: Anthoathecata: Filifera) found at night time in the southern Red Sea (Saudi Arabia) living on shells of the gastropod Nassarius margaritifer (Dunker, 1847) (Neogastropoda: Buccinoidea: Nassariidae). We examined the fluorescence of these polyps and compare with previously reported data. Intensive green fluorescence with a spectral peak at 518 nm was detected in the hypostome of the Cytaeis polyps, unlike in previous reports that reported fluorescence either in the basal parts of polyps or in other locations on hydroid colonies. These results suggest that fluorescence may be widespread not only in medusae, but also in polyps, and also suggests that the patterns of fluorescence localization can vary in closely related species. The fluorescence of polyps may be potentially useful for field identification of cryptic species and study of geographical distributions of such hydroids and their hosts.

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The resurrection of the genus Bergia (Anthozoa, Zoantharia, Parazoanthidae)

Montenegro J.

Systematics and Biodiversity ( Systematics and Biodiversity )  14 ( 1 ) 63 - 73   2016.01 [ Peer Review Accepted ]

Type of publication: Research paper (scientific journal)

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The genus Bergia was established by Duchassaing de Fonbressin and Michelotti in 1860, for two species, Bergia catelunaris and B. vialactea. Subsequently, in 1903 Duerden recognized these two species as conspecific, and used B. catelunaris in favour of B. vialactea, and transferred B. catelunaris to the genus Parazoanthus. However, over the last decade, it has been found that the genus Parazoanthus is actually polyphyletic and therefore it has gradually been divided and redefined. Based on phylogenetic analyses, Parazoanthus sensu stricto was recently limited to species which form associations with sponges, but it still comprised of three distinctive and monophyletic subclades. Of these clades, one Parazoanthus clade contains the type species for Parazoanthus, P. axinellae, while another clade was recently described as the genus Umimayanthus based on mitochondrial 16S-rDNA sequences. However, the other remaining Parazoanthus clade contains P. catenularis, the original type species of the genus Bergia. Based on a concatenated set of published data for five molecular markers (18S-rDNA, 28S-rDNA, ITS1/5.8S/ITS2-rDNA, 16S-rDNA and COI-DNA), this study confirms the monophyly of Bergia in the context of Parazoanthidae. The phylogenetic analyses strongly support the resurrection of the genus Bergia as a valid taxonomic unit.

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Preliminary analyses reveal strong genetic structure in populations of Leucothoe vulgaris (Crustacea: Amphipoda: Leucothoidae) from Okinawa, Japan

White K.

Systematics and Biodiversity ( Systematics and Biodiversity )  14 ( 1 ) 55 - 62   2016.01 [ Peer Review Accepted ]

Type of publication: Research paper (scientific journal)

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Preliminary phylogenetic analyses of specimens of a leucothoid amphipod, Leucothoe vulgaris, collected from sponges, ascidians, and coral rubble from Okinawa, Japan, were completed using mitochondrial COI and nuclear 18S ribosomal DNA sequences. Analyses of sequences from 83 specimens demonstrate that populations of L. vulgaris likely entail at least two cryptic species, which are reciprocally monophyletic and reproductively isolated. These two potentially cryptic species live in sympatry but are apparently morphologically identical, suggesting a geographically driven divergence process and secondary contact. Within each clade, two major subclades corresponding to the east and west coast of Okinawa Island were present, with divergence times of approximately 1.61-1.83 mya. This last result suggests a role of Pleistocene sea level changes in the current patterns of intra-specific genetic structure and highlights the need for a more comprehensive sampling of L. vulgaris throughout the Indo-west Pacific.

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First record of the genus Sphenopus (Anthozoa: Hexacorallia: Zoantharia) from Japan

JD Reimer, T Higashiji, M Nonaka

Fauna Ryukyuana   29   37 - 40   2016.01 [ Peer Review Accepted ]

Type of publication: Research paper (other science council materials etc.)

Unexpected diversity and a new species of Epizoanthus (Anthozoa, Hexacorallia) attached to eunicid worm tubes from the Pacific Ocean.

Kise H, Reimer JD

ZooKeys ( ZooKeys )  2016 ( 562 ) 49 - 71   2016 [ Peer Review Accepted ]

Type of publication: Research paper (scientific journal)

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Epizoanthus species are generally found in association with other marine invertebrates such as hermit crabs and gastropods. Although Epizoanthus spp. are relatively common, there is limited information about their diversity and ecology due to their habitats or hosts, often being below the depths of SCUBA diving (>similar to 50 m). In particular, the Epizoanthus fauna of the Indo-Pacific Ocean remains poorly understood. In this study, the diversity of Epizoanthus species associated with eunicid worm tubes from shallow waters in the Pacific Ocean we investigated using molecular analyses (mitochondrial cytochrome oxidase subunit 1 = COI, mitochondrial 16S ribosomal DNA = mt 16S-rDNA, nuclear internal transcribed spacer region of ribosomal DNA = ITS-rDNA) combined with morphological and ecological data. The combined data set leads us to describe two new species; Epizoanthus inazuma sp. n. and Epizoanthus beriber sp. n. Both new species are found in low-light environments: E. inazuma sp. n. on mesophotic coral reef slopes and reef floors, or on the sides of overhangs; E. beriber sp. n. has only been found in caves. Morphological characteristics of these two new species are very similar to E. illoricatus Tischbierek, 1930 but the two new species are genetically distinct. Mesentery numbers and coloration of polyps may be useful diagnostic characteristics among eunicid-associated Epizoanthus species. These results demonstrate that there is high potential for other potentially undescribed zoantharian species, particularly in underwater cave habitats.

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A new solitary free-living species of the genus Sphenopus (Cnidaria, Anthozoa, Zoantharia, Sphenopidae) from Okinawa-jima Island, Japan.

Fujii T, Reimer JD

ZooKeys ( ZooKeys )  2016 ( 606 ) 11 - 24   2016 [ Peer Review Accepted ]

Type of publication: Research paper (scientific journal)

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A new species of free-living solitary zoantharian is described from Okinawa, Japan. Sphenopus exilis sp. n. occurs on silty seafloors in Kin Bay and Oura Bay on the east coast of Okinawa-jima Island. Sphenopus exilis sp. n. is easily distinguished from other Sphenopus species by its small polyp size and slender shape, although there were relatively few differences between Sphenopus exilis sp. n. and S. marsupialis in the molecular phylogenetic analyses. Currently, very little is known about the ecology and diversity of Sphenopus species. Thus, reviewing each species carefully via combined morphological and molecular analyses by using newly obtained specimens from type localities is required to clearly understand and distinguish the species within the genus Sphenopus.

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A citizen science approach to monitoring bleaching in the zoantharian Palythoa tuberculosa.

Parkinson JE, Yang SY, Kawamura I, Byron G, Todd PA, Reimer JD

PeerJ ( PeerJ )  4 ( 3 ) e1815   2016 [ Peer Review Accepted ]

Type of publication: Research paper (scientific journal)

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Coral reef bleaching events are expected to become more frequent and severe in the near future as climate changes. The zoantharian Palythoa tuberculosa bleaches earlier than many scleractinian corals and may serve as an indicator species. Basic monitoring of such species could help to detect and even anticipate bleaching events, especially in areas where more sophisticated approaches that rely on buoy or satellite measurements of sea surface temperature are unavailable or too coarse. One simple and inexpensive monitoring method involves training volunteers to record observations of host color as a proxy for symbiosis quality. Here, we trained university students to take the 'color fingerprint' of a reef by assessing the color of multiple randomly selected colonies of P. tuberculosa at one time point in Okinawa Island, Japan. We tested the reliability of the students' color scores and whether they matched expectations based on previous monthly monitoring of tagged colonies at the same locations. We also measured three traditional metrics of symbiosis quality for comparison: symbiont morphological condition, cell density, and chlorophyll a content. We found that P. tuberculosa color score, although highly correlated among observers, provided little predictive power for the other variables. This was likely due to inherent variation in colony color among generally healthy zoantharians in midwinter, as well as low sample size and brief training owing to the course structure. Despite certain limitations of P. tuberculosa as a focal organism, the citizen science approach to color monitoring has promise, and we outline steps that could improve similar efforts in the future.

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The order Zoantharia Rafinesque, 1815 (Cnidaria, Anthozoa: Hexacorallia): supraspecific classification and nomenclature.

Low ME, Sinniger F, Reimer JD

ZooKeys ( ZooKeys )  2016 ( 641 ) 1 - 80   2016 [ Peer Review Accepted ]

Type of publication: Research paper (scientific journal)

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Many supraspecific zoantharian names have long and complicated histories. The present list is provided to advise researchers on the current state of supraspecific nomenclature of the zoantharians, particularly given the recent attention paid to the taxonomy, phylogeny, and biodiversity of this order. At the same time, several taxonomic issues brought to light by recent research are resolved. Details on the taxonomic and nomenclatural history of most groups are provided, along with appendices of invalid supraspecific names.

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沖縄島西海岸における中深度サンゴ礁生態系に関する研究

水山克, 大原拓, 大原拓, WEINSTEIN David, 藤井琢磨, 藤井琢磨, MCCLELLAND Ty, MONTENEGRO Javier, 式場はるか, 浅海竜司, YANG Sung‐yin, 河村伊織, REIMER James Davis

沖縄生物学会誌   ( 54 ) 54 - 55   2016

Type of publication: Research paper (other science council materials etc.)

Phylogenetic characterization of two echinoid species of the southeastern Mediterranean, off Egypt

Soliman T.

Egyptian Journal of Aquatic Research ( Egyptian Journal of Aquatic Research )  41 ( 4 ) 359 - 365   2015.12 [ Peer Review Accepted ]

Type of publication: Research paper (scientific journal)

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In this study we investigated the phylogenetics of two sea urchin species, Arbacia lixula and Paracentrotus lividus from the Mediterranean Sea. Specimens were collected from the east coast of Alexandria City, Egypt. Pigmentation examination showed four sympatric color morphotypes (black, purple, reddish brown, and olive green). Mitochondrial DNA was extracted from specimens and mitochondrial cytochrome oxidase subunit I (COI) and 16S ribosomal RNA (16S) were sequenced. The results showed that all black specimens constituted the species A. lixula. All other colors belonged to P. lividus, with no apparent differentiation between color morphotypes. Moreover, P. lividus showed high haplotype diversity (COI H = 0.9500 and 16S H = 0.8580) and low values of nucleotide diversity (COI π = 0.0075 and 16S π = 0.0049), indicating a high degree of polymorphism within this species. This study represents the first attempt at DNA barcoding of echinoid species in the southeast Mediterranean off the Egyptian coast, and will provide a base for future phylogenetic analyses.

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Yellow tails in the Red Sea: Phylogeography of the Indo-Pacific goatfish Mulloidichthys flavolineatus reveals isolation in peripheral provinces and cryptic evolutionary lineages

Fernandez-Silva I.

Journal of Biogeography ( Journal of Biogeography )  42 ( 12 ) 2402 - 2413   2015.12 [ Peer Review Accepted ]

Type of publication: Research paper (scientific journal)

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Aim Broadly distributed reef fishes tend to have high gene flow mediated by a pelagic larval phase. Here, we survey a reef-associated fish distributed across half the tropical oceans, from the Red Sea to the central Pacific. Our goal is to determine whether genetic structure of the broadly distributed Yellowstripe Goatfish (Mulloidichthys flavolineatus) is defined by biogeographical barriers, or facilitated via larval dispersal. Location Red Sea, Indian Ocean, Pacific Ocean Methods Specimens were obtained at 19 locations from the Red Sea to Hawai'i. Genetic data include mtDNA cytochrome b (n = 217) and 12 microsatellite loci (n = 185). Analysis of molecular variance (AMOVA), structure, a parsimony network and coalescence analyses were used to resolve recent population history and connectivity. Results Population structure was significant (mtDNA Phi(ST) = 0.68, P < 0.001; microsatellite F-ST = 0.08, P < 0.001), but mostly driven by samples from the North-western (NW) Indian Ocean (including the Red Sea) and Hawai'i. There was little population structure across the Indian Ocean to the central Pacific. Hawai'i was distinguished as an isolated population (mtDNA Phi(ST) = 0.03-0.08, P = n.s.; microsatellites F-ST = 0.05-0.10, P < 0.001). Specimens from the NW Indian Ocean clustered as a distinct phylogenetic lineage that diverged approximately 493 ka (d = 1.7%), which indicates that these fish persisted in isolation through several Pleistocene glacial cycles. Main conclusions These data reinforce the emerging themes that: (1) phylogeographical breaks within species often coincide with biogeographical breaks based on species distributions, and (2) populations on the periphery of the range (NW Indian Ocean and Hawai'i) are isolated and may be evolutionary incubators producing new species.

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Zoantharia (Cnidaria: Anthozoa: Hexacorallia) of the south china sea and gulf of thailand: A species list based on past reports and new photographic records

Reimer J.

Raffles Bulletin of Zoology ( Raffles Bulletin of Zoology )  63   334 - 356   2015.09

Type of publication: Research paper (scientific journal)

Zoantharia (Cnidaria: Anthozoa: Hexacorallia) of the south china sea and gulf of thailand: A species list based on past reports and new photographic records

Reimer JD, HB Wee, A Put Jr, BW Hoeksema

Raffles Bulletin of Zoology ( Raffles Bulletin of Zoology )  63   334 - 356   2015.09 [ Peer Review Accepted ]

Type of publication: Research paper (scientific journal)

Unexpected diversity and new species in the sponge-Parazoanthidae association in southern Japan.

Montenegro J, Sinniger F, Reimer JD

Molecular phylogenetics and evolution ( Molecular Phylogenetics and Evolution )  89   73 - 90   2015.08 [ Peer Review Accepted ]

Type of publication: Research paper (scientific journal)

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Currently the genera Parazoanthus (family Parazoanthidae) and Epizoanthus (family Epizoanthidae) are the only sponge-associated zoantharians (Cnidaria, Anthozoa). The Parazoanthidae-sponge associations are widely distributed in tropical and subtropical waters from the intertidal to the deep sea in the Atlantic and Indo-Pacific Oceans. However, the taxonomic identification of both parties is often confused due to variable morphology and wide ecological ranges. In particular, Parazoanthidae species diversity remains poorly understood in the Indo-Pacific. In the present study, the diversity of the sponge-zoanthid association in the Indo-Pacific was investigated with 71 Parazoanthidae specimens collected from 29 different locations in Japan (n = 22), Australia (n = 6) and Florida, USA (n = 1). For all specimens morphological analyses were performed and total DNA was extracted and amplified for four DNA markers (COI-mtDNA, mt 16S-rDNA, ITS-rDNA and ALG11-nuDNA). The combined data demonstrate that the specimens of this study are clearly different from those of all described Parazoanthus species, and lead us to erect Umimayanthus gen. n., within family Parazoanthidae, containing the three newly described species U. chanpuru sp. n., U. miyabi sp. n., U. nakama sp. n. The new genus also includes the previously described species U. parasiticus (Duchassaing and Michelotti, 1860; comb. nov.), previously belonging to the genus Parazoanthus. Neighbor joining, maximum likelihood and Bayesian posterior probability phylogenetic trees clearly demonstrate the monophyly of Umimayanthus gen. n. to the exclusion of all outgroup sequences. The phylogenetic results were also compared to morphological features, and polyp sizes, amount of sand content in tissues, types of connections between polyps, and cnidae data, in particular holotrichs-1, were useful in distinguishing the different species within this new genus. This new genus can be distinguished from all other Zoantharia by a unique and conserved 9 bp insertion and a 14 bp deletion in the mt 16S-rDNA region. Additionally, compared to Parazoanthus sensu stricto (i.e. P. axinellae [Schmidt, 1862]), Umimayanthus spp. are only found associated to sponges, and have a coenenchyme much less developed than Parazoanthus sensu strict. Each new species can be distinguished from other congeners by a unique DNA sequence, numbers of tentacle, maximum sizes of holotrichs, associated sponge morphology, and colony morphology. The identification of the host sponge species is the next logical step in this research as this may also aid in the distinction of Umimayanthus species. (C) 2015 Elsevier Inc. All rights reserved.

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Colony-specific investigations reveal highly variable responses among individual corals to ocean acidification and warming.

Kavousi J, Reimer JD, Tanaka Y, Nakamura T

Marine environmental research ( Marine Environmental Research )  109   9 - 20   2015.08 [ Peer Review Accepted ]

Type of publication: Research paper (scientific journal)

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As anthropogenic climate change is an ongoing concern, scientific investigations on its impacts on coral reefs are increasing. Although impacts of combined ocean acidification (OA) and temperature stress (T) on reef-building scleractinian corals have been studied at the genus, species and population levels, there are little data available on how individual corals respond to combined OA and anomalous temperatures. In this study, we exposed individual colonies of Acropora digitifera, Montipora digitata and Porites cylindrica to four pCO(2)-temperature treatments including 400 mu atm-28 degrees C, 400 mu atm-31 degrees C, 1000 mu atm-28 degrees C and 1000 mu atm-31 degrees C for 26 days. Physiological parameters including calcification, protein content, maximum photosynthetic efficiency, Symbiodinium density, and chlorophyll content along with Symbiodinium type of each colony were examined. Along with intercolonial responses, responses of individual colonies versus pooled data to the treatments were investigated. The main results were: 1) responses to either OA or T or their combination were different between individual colonies when considering physiological functions; 2) tolerance to either OA or T was not synonymous with tolerance to the other parameter; 3) tolerance to both OA and T did not necessarily lead to tolerance of OA and T combined (OAT) at the same time; 4) OAT had negative, positive or no impacts on physiological functions of coral colonies; and 5) pooled data were not representative of responses of all individual colonies. Indeed, the pooled data obscured actual responses of individual colonies or presented a response that was not observed in any individual. From the results of this study we recommend improving experimental designs of studies investigating physiological responses of corals to climate change by complementing them with colony-specific examinations. (C) 2015 Elsevier Ltd. All rights reserved.

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Effects of causeway construction on environment and biota of subtropical tidal flats in Okinawa, Japan.

Reimer JD, Yang SY, White KN, Asami R, Fujita K, Hongo C, Ito S, Kawamura I, Maeda I, Mizuyama M, Obuchi M, Sakamaki T, Tachihara K, Tamura M, Tanahara A, Yamaguchi A, Jenke-Kodama H

Marine pollution bulletin ( Marine Pollution Bulletin )  94 ( 1-2 ) 153 - 67   2015.05 [ Peer Review Accepted ]

Type of publication: Research paper (scientific journal)

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Okinawa, Japan is known for its high marine biodiversity, yet little work has been performed on examining impacts of numerous large-scale coastal development projects on its marine ecosystems. Here, we examine apparent impacts of the construction of the Kaichu-Doro causeway, which was built over 40 years ago. The causeway is a 4.75 km long embankment that divides a large tidal flat and has only two points of water exchange along its entire length. We employed quadrats, transects, sampling, visual surveys, and microbial community analyses combined with environmental, water quality data, and 1 m cores, at five stations of two paired sites each (one on each side of Kaichu-Doro) to investigate how the environment and biota have changed since the Kaichu-Doro was built. Results indicate reduction in water flow, and site S1 was particularly heavily impacted by poor water quality, with low diversity and disturbed biotic communities. (C) 2015 The Authors. Published by Elsevier Ltd.

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First record of Nanozoanthidae from the Red Sea

Reimer J.D.

Marine Biodiversity Records ( Marine Biodiversity Records )  8   2015.04 [ Peer Review Accepted ]

Type of publication: Research paper (scientific journal)

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Here we report on the first finding of Nanozoanthidae (Anthozoa: Hexacorallia: Zoantharia) in the Red Sea and the first record west of Western Australia. A single specimen of Nanozoanthus sp. was found at a depth of 13 m off Dumsuq Island, the Farasan Islands, Saudi Arabia (16�33.846′N 42�03.510′E) during SCUBA surveys. Previous research had hypothesized that the genus could potentially be widespread in the Indo-Pacific and was simply undetected due to its small and cryptic nature, and the current specimen provides support for this idea. Such findings demonstrate the importance of biodiversity surveys by taxonomic specialists in understudied marine regions.

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First record of the genus Umimayanthus from Palau and Micronesia

Montenegro J.

Marine Biodiversity Records ( Marine Biodiversity Records )  8   2015.04 [ Peer Review Accepted ]

Type of publication: Research paper (scientific journal)

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Until recently, the only sponge-associated genera in the order Zoantharia were Parazoanthus (family Parazoanthidae), Epizoanthus and Thoracactis (family Epizoanthidae), both within the suborder Macrocnemina. The taxonomy of the genus Parazoanthus, as originally described, has been undergoing revision since 2010, with several species, genera and even families described. In 2015, multiple molecular markers were used in combination with morphological characteristics to erect the genus Umimayanthus inside the family Parazoanthidae. It included three species described from southern Japan, with other records for some of the species from the Great Barrier Reef, New Caledonia and the Red Sea. However, little is known of its distribution in the Pacific Islands. Here we report on the finding of Umimayanthus specimens in Palau, Micronesia, representing the first records for this region. A total of 32 specimens of Umimayanthus were collected from seven different locations eight of the specimens were identified as Umimayanthus chanpuru, while the remaining 24 colonies were only identified to genus level.

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Effect of phase shift from corals to zoantharia on reef fish assemblages

Cruz I.C.S.

PLoS ONE ( PLoS ONE )  10 ( 1 )   2015.01 [ Peer Review Accepted ]

Type of publication: Research paper (scientific journal)

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Consequences of reef phase shifts on fish communities remain poorly understood. Studies on the causes, effects and consequences of phase shifts on reef fish communities have only been considered for coral-to-macroalgae shifts. Therefore, there is a large information gap regarding the consequences of novel phase shifts and how these kinds of phase shifts impact on fish assemblages. This study aimed to compare the fish assemblages on reefs under normal conditions (relatively high cover of corals) to those which have shifted to a dominance of the zoantharian Palythoa cf. variabilis on coral reefs in Todos os Santos Bay (TSB), Brazilian eastern coast. We examined eight reefs, where we estimated cover of corals and P. cf. variabilis and coral reef fish richness, abundance and body size. Fish richness differed significantly between normal reefs (48 species) and phase-shift reefs (38 species), a 20% reduction in species. However there was no difference in fish abundance between normal and phase shift reefs. One fish species, Chaetodon striatus, was significantly less abundant on normal reefs. The differences in fish assemblages between different reef phases was due to differences in trophic groups of fish; on normal reefs carnivorous fishes were more abundant, while on phase shift reefs mobile invertivores dominated.

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