Acquisition of dwarf male "harems" by recently settled females of Osedax roseus n. sp. (Siboglinidae; Annelida)

After the deployment of several whale carcasses in Monterey Bay, California, a time-series analysis revealed the presence of a new species of Osedax, a genus of bone-eating siboglinid annelids. That species is described here as Osedax roseus n. sp. It is the fifth species described since the erection of this genus and, like its congeners, uses a ramifying network of "roots" to house symbiotic bacteria. In less than 2 months, Osedax roseus n. sp. colonized the exposed bones of a whale carcass deposited at 1018-m depth, and many of the females were fecund in about 3 months post-deployment. As with other Osedax spp., the females have dwarf males in their tube lumens. The males accrue over time until the sex ratio is markedly male-biased. This pattern of initial female settlement followed by gradual male accumulation is consistent with the hypothesis that male sex may be environmentally determined in Osedax. Of the previously described species in this genus, Osedax roseus n. sp. is most similar to O. rubiplumus, but it has several anatomical differences, as well as much smaller females, dwarf males, and eggs. Osedax roseus n. sp. is markedly divergent (minimally 16.6%) for mitochondrial cytochrome oxidase subunit I (mtCOI) sequences from any other Osedax species.     

Molecular taxonomy of Osedax (Annelida: Siboglinidae) in the Southern Ocean

We report the discovery of three new species of Osedax in the deep Southern Ocean, expanding the diversity and geographical range of this genus of bone‐eating worms. Osedax rogersi sp. n. and Osedax crouchi sp. n. were found on a whale skeleton at 1444 m in the Kemp Caldera in the East Scotia Sea during the Chemosynthetic Ecosystems of the Southern Ocean (ChEsSo) project. The recently described species, Osedax antarcticus, found on whale bones implanted at a depth of 550–650 m off Smith Island in the Bransfield Strait, was also found inhabiting the whale skeleton in the Kemp Caldera. Osedax crouchi as well as another new species Osedax nordenskjoeldi sp. n. have also been found on the implanted whale bones off Smith Island. These two localities are approximately 1800 km apart demonstrating a wide distribution of species within the genus. We describe the three new species, O. rogersi, O. crouchi and O. nordenskjoeldi and report the second record of O. antarcticus. We also present a new phylogenetic analysis for Osedax, including data examining genetic connectivity between the Scotia Arc and the Bransfield Strait.     

World-wide whale worms? A new species of Osedax from the shallow north Atlantic

We describe a new species of the remarkable whalebone-eating siboglinid worm genus, Osedax, from a whale carcass in the shallow north Atlantic, west of Sweden. Previously only recorded from deep-sea (1500-3000m) whale-falls in the northeast Pacific, this is the first species of Osedax known from a shelf-depth whale-fall, and the first from the Atlantic Ocean. The new species, Osedax mucofloris sp. n. is abundant on the bones of an experimentally implanted Minke whale carcass (Balaenoptera acutorostrata) at 125m depth in the shallow North Sea. O. mucofloris can be cultured on bones maintained in aquaria. The presence of O. mucofloris in the shallow North Sea and northeast Pacific suggests global distribution on whale-falls for the Osedax clade. Molecular evidence from mitochondrial cytochrome oxidase 1 (CO1) and 18S rRNA sequences suggests that O. mucofloris has high dispersal rates, and provides support for the idea of whale-falls acting as 'stepping-stones' for the global dispersal of siboglinid annelids over ecological and evolutionary time.     

Between a whale bone and the deep blue sea: the provenance of dwarf males in whale bone-eating tubeworms

When researchers first caught a glimpse of the lush carpet of pink tubeworms covering the scattered bones of a dead grey whale 2900 m below the surface of Monterey Bay, the excitement onboard the Western Flyer (the mother ship of the remotely operated vehicle the Tiburon) must have been electrifying. The discovery of a new genus and several species of whale bone-eating Osedax tubeworms (Annelida, Siboglinidae) a mere 6 years ago from the deep sea was itself noteworthy. But what the researchers peering into the video monitors aboard the Western Flyer could not have known at that moment was that in the gelatinous tubes of those worms clung even more peculiar forms: harems of tiny, paedomorphic males of Osedax, numbering in the hundreds at times. Whereas female tubeworms bore into the marrow of whale bones (possibly via enzymes from their endosymbiotic bacteria), the dwarf males secondarily colonize the tubes of the resident females. The number of males in a female's tube increases over time in a curvilinear fashion. Dwarf males are known from all Osedax species examined to date, yet the origin of the males was an open question. In this issue, Vrijenhoek et al. provide compelling evidence that dwarf males found in the tubes of female Osedax worms are derived from a common larval pool and are unlikely to be the sons of host females or the progeny of females in the local genetic neighbourhood. This study provides an important foundation for future work on the ecology and evolution of extreme male dwarfism in Osedax and sexual size dimorphism more generally.     

Genetic diversity and potential function of microbial symbionts associated with newly discovered species of Osedax polychaete worms

We investigated the genetic diversity of symbiotic bacteria associated with two newly discovered species of Osedax from Monterey Canyon, CA, at 1,017-m (Osedax Monterey Bay sp. 3 "rosy" [Osedax sp. MB3]) and 381-m (Osedax Monterey Bay sp. 4 "yellow collar") depths. Quantitative PCR and clone libraries of 16S rRNA gene sequences identified differences in the compositions and abundances of bacterial phylotypes associated with the newly discovered host species and permitted comparisons between adult Osedax frankpressi and juveniles that had recently colonized whalebones implanted at 2,891 m. The newly discovered Osedax species hosted Oceanospirillales symbionts that are related to Gammaproteobacteria associated with the previously described O. frankpressi and Osedax rubiplumus (S. K. Goffredi, V. J. Orphan, G. W. Rouse, L. Jahnke, T. Embaye, K. Turk, R. Lee, and R. C. Vrijenhoek, Environ. Microbiol. 7:1369-1378, 2005). In addition, Osedax sp. MB3 hosts a diverse and abundant population of additional bacteria dominated by Epsilonproteobacteria. Ultrastructural analysis of symbiont-bearing root tissues verified the enhanced microbial diversity of Osedax sp. MB3. Root tissues from the newly described host species and O. frankpressi all exhibited collagenolytic enzyme activity, which covaried positively with the abundance of symbiont DNA and negatively with mean adult size of the host species. Members of this unusual genus of bone-eating worms may form variable associations with symbiotic bacteria that allow for the observed differences in colonization and success in whale fall environments throughout the world's oceans.     

The Potent Respiratory System of Osedax mucofloris (Siboglinidae,Annelida) - A Prerequisite for the Origin of Bone-Eating Osedax?

Members of the conspicuous bone-eating genus, Osedax, are widely distributed on whale falls in the Pacific and Atlantic Oceans. These gutless annelids contain endosymbiotic heterotrophic bacteria in a branching root system embedded in the bones of vertebrates, whereas a trunk and anterior palps extend into the surrounding water. The unique life style within a bone environment is challenged by the high bacterial activity on, and within, the bone matrix possibly causing O(2) depletion, and build-up of potentially toxic sulphide. We measured the O(2) distribution around embedded Osedax and showed that the bone microenvironment is anoxic. Morphological studies showed that ventilation mechanisms in Osedax are restricted to the anterior palps, which are optimized for high O(2) uptake by possessing a large surface area, large surface to volume ratio, and short diffusion distances. The blood vascular system comprises large vessels in the trunk, which facilitate an ample supply of oxygenated blood from the anterior crown to a highly vascularised root structure. Respirometry studies of O. mucofloris showed a high O(2) consumption that exceeded the average O(2) consumption of a broad line of resting annelids without endosymbionts. We regard this combination of features of the respiratory system of O. mucofloris as an adaptation to their unique nutrition strategy with roots embedded in anoxic bones and elevated O(2) demand due to aerobic heterotrophic endosymbionts.     

Not whale-fall specialists, Osedax worms also consume fishbones

Marine annelid worms of the genus Osedax exploit sunken vertebrate bones for food. To date, the named species occur on whale or other mammalian bones, and it is argued that Osedax is a whale-fall specialist. To assess whether extant Osedax species could obtain nutrition from non-mammalian resources, we deployed teleost bones and calcified shark cartilage at approximately 1000 m depth for five months. Although the evidence from shark cartilage was inconclusive, the teleost bones hosted three species of Osedax, each of which also lives off whalebones. This suggests that rather than being a whale-fall specialist, Osedax has exploited and continues to exploit a variety of food sources. The ability of Osedax to colonize and to grow on fishbone lends credibility to a hypothesis that it might have split from its siboglinid relatives to assume the bone-eating lifestyle during the Cretaceous, well before the origin of marine mammals.     

Bone-boring worms: characterizing the morphology, rate, and method of bioerosion by Osedax mucofloris (Annelida, Siboglinidae)

Osedax worms possess unique "root" tissues that they use to bore into bones on the seafloor, but details of the boring pattern and processes are poorly understood. Here we use X-ray micro-computed tomography to investigate the borings of Osedax mucofloris in bones of the minke whale (Balaenoptera acutorostrata), quantitatively detailing their morphological characteristics for the first time. Comparative thin-sections of the borings reveal how the bone is eroded at the sub-millimeter level. On the basis of these results we hypothesize a model of boring that is dependent on the density and microstructure of the bone. We also present evidence of acidic mucopolysaccharides in the mucus of the root tissue, and hypothesize that this plays an important role in the boring mechanism. We discuss the utility of these new data in evaluating Osedax trace fossils and their relevance for O. mucofloris ecology. Measured rates of bone erosion (6% per year) and evidence of enhanced sulfide release from the borings indicate that Osedax worms are important habitat modifiers in whale-fall communities.     

High symbiont diversity in the bone-eating worm Osedax mucofloris from shallow whale-falls in the North Atlantic

Osedax worms are whale-fall specialists that infiltrate whale bones with their root tissues. These are filled with endosymbiotic bacteria hypothesized to provide their hosts with nutrition by extracting organic compounds from the whale bones. We investigated the diversity and distribution of symbiotic bacteria in Osedax mucofloris from shallow-water whale-falls in the North Atlantic using comparative 16S rRNA sequence analysis and fluorescence in situ hybridization (FISH). We observed a higher diversity of endosymbionts than previously described from other Osedax species. Endosymbiont sequences fell into eight phylogenetically distinct clusters (with 91.4-98.9% similarity between clusters), and considerable microdiversity within clusters (99.5-99.7% similarity) was observed. Statistical tests revealed a highly significant effect of the host individual on endosymbiont diversity and distribution, with 68% of the variability between clusters and 40% of the variability within clusters explained by this effect. FISH analyses showed that most host individuals were dominated by endosymbionts from a single cluster, with endosymbionts from less abundant clusters generally confined to peripheral root tissues. The observed diversity and distribution patterns indicate that the endosymbionts are transmitted horizontally from the environment with repeated infection events occurring as the host root tissues grow into the whale bones.     

The plam-feeding planthopper genus Ommatissus (Homoptera: Fulgoroidea: Tropiduchidae)

Abstract The tribe Trypetimorphini, the genus Ommatissus , and its two subgenera and eleven species are redescribed or described as new. Evidence for the monophyly of both the genus and of the tribe to which it belongs is examined; it is concluded that while the genus is monophyletic, the monophyly of the tribe is in doubt. Keys for the separation of the three genera of the tribe and for all of the known species of Ommatissus are presented. A new subgenus of Ommatissus is described: O.(Opatissus) , with the type-species O.vietnamicus . Four new species are described: Ommatissus magribus from Morocco, O.kamerunus from Cameroon, O.natalensis from South Africa and O.vietnamicus from Vietnam. The 'Dubas bug', Ommatissus lybicus Bergevin, formerly regarded as a variety of O.binotatus Fieber, is raised to full species status. Trichoduchus is synonymized with Trypetimorpha resulting in the new combinations Trypetimorpha biermani Dammerman, T.china Wu and T.japonicus Ishihara. Trichoduchus japonicus Fennah is synonymized with Trypetimorpha japonicus Ishihara.     

Evolutionary innovation: a bone-eating marine symbiosis

Symbiotic associations between microbes and invertebrates have resulted in some of the most unusual physiological and morphological adaptations that have evolved in the animal world. We document a new symbiosis between marine polychaetes of the genus Osedax and members of the bacterial group Oceanospirillales, known for heterotrophic degradation of complex organic compounds. These organisms were discovered living on the carcass of a grey whale at 2891 m depth in Monterey Canyon, off the coast of California. The mouthless and gutless worms are unique in their morphological specializations used to obtain nutrition from decomposing mammalian bones. Adult worms possess elaborate posterior root-like extensions that invade whale bone and contain bacteriocytes that house intracellular symbionts. Stable isotopes and fatty acid analyses suggest that these unusual endosymbionts are likely responsible for the nutrition of this locally abundant and reproductively prolific deep-sea worm.     

Genetic Diversity and Potential Function of Microbial Symbionts Associated with Newly Discovered Species of Osedax Polychaete Worms

We investigated the genetic diversity of symbiotic bacteria associated with two newly discovered species of Osedax from Monterey Canyon, CA, at 1,017-m (Osedax Monterey Bay sp. 3 “rosy” [Osedax sp. MB3]) and 381-m (Osedax Monterey Bay sp. 4 “yellow collar”) depths. Quantitative PCR and clone libraries of 16S rRNA gene sequences identified differences in the compositions and abundances of bacterial phylotypes associated with the newly discovered host species and permitted comparisons between adult Osedax frankpressi and juveniles that had recently colonized whalebones implanted at 2,891 m. The newly discovered Osedax species hosted Oceanospirillales symbionts that are related to Gammaproteobacteria associated with the previously described O. frankpressi and Osedax rubiplumus (S. K. Goffredi, V. J. Orphan, G. W. Rouse, L. Jahnke, T. Embaye, K. Turk, R. Lee, and R. C. Vrijenhoek, Environ. Microbiol. 7:1369-1378, 2005). In addition, Osedax sp. MB3 hosts a diverse and abundant population of additional bacteria dominated by Epsilonproteobacteria. Ultrastructural analysis of symbiont-bearing root tissues verified the enhanced microbial diversity of Osedax sp. MB3. Root tissues from the newly described host species and O. frankpressi all exhibited collagenolytic enzyme activity, which covaried positively with the abundance of symbiont DNA and negatively with mean adult size of the host species. Members of this unusual genus of bone-eating worms may form variable associations with symbiotic bacteria that allow for the observed differences in colonization and success in whale fall environments throughout the world's oceans.     

Marine worms (genus Osedax) colonize cow bones

Bone-eating worms of the genus Osedax colonized and grew on cow bones deployed at depths ranging from 385 to 2893m in Monterey Bay, California. Colonization occurred as rapidly as two months following deployment of the cow bones, similar to the time it takes to colonize exposed whalebones. Some Osedax females found on the cow bones were producing eggs and some hosted dwarf males in their tubes. Morphological and molecular examinations of these worms confirmed the presence of six Osedax species, out of the eight species presently known from Monterey Bay. The ability of Osedax species to colonize, grow and reproduce on cow bones challenges previous notions that these worms are 'whale-fall specialists.'     

Phylogeny of the genus Omphalotus based on nuclear ribosomal DNA-sequences

The evolutionary history of the genus Omphalotus was inferred from DNA sequences of the ITS1-5.8S-ITS2 rDNA region. We analyzed 32 collections from different geographical areas: O. olearius (Europe), O. illudens (Europe, North America), O. subilludens (North America), O. olivascens var. olivascens (North America) and var. indigo (Mexico), O. mexicanus (Middle America), O. nidiformis (Australia), and O. japonicus (Japan). Phylogenetic analysis was performed declaring Nothopanus eugrammus as outgroup. Our analyses show that the genus Omphalotus is split into two major clades, the first consisting of O. illudens and O. mexicanus and the second comprising O. olearius, O. olivascens, O. japonicus, O. nidiformis and O. subilludens. Moreover, the often discussed synonymy of O. illudens and O. olearius is rejected. Omphalotus japonicus, a species formerly placed in the genus Lampteromyces Sing. for morphological reasons, clustered as the sister group of O. olearius.     

Resurrection of Mesoplodon hotaula Deraniyagala 1963: A new species of beaked whale in the tropical Indo‐Pacific

We present genetic and morphological evidence supporting the recognition of a previously synonymized species of Mesoplodon beaked whale in the tropical Indo‐Pacific, Mesoplodon hotaula. Although the new species is closely‐related to the rare ginkgo‐toothed beaked whale M. ginkgodens, we show that these two lineages can be differentiated by maternally (mitochondrial DNA), biparentally (autosomal), and paternally (Y chromosome) inherited DNA sequences, as well as by morphological features. The reciprocal monophyly of the mtDNA genealogies and the largely parapatric distribution of these lineages is consistent with reproductive isolation. The new lineage is currently known from at least seven specimens: Sri Lanka (1), Gilbert Islands, Republic of Kiribati (1+), Palmyra Atoll, Northern Line Islands, U.S.A. (3), Maldives (1), and Seychelles (1). The type specimen (Sri Lanka) was described as a new species, M. hotaula, in 1963, but later synonymized with M. ginkgodens. This discovery brings the total number of Mesoplodon species to 15, making it, by far, the most speciose yet least known genus of cetaceans.     

基于16S rRNA基因扩增子测序分析日本囊对虾肠道菌群结构与功能的特征

【背景】肠道菌群在对虾的生理活动中起关键作用。日本囊对虾是我国海水养殖虾类中的主要品种之一,迄今为止有关其肠道菌群结构与功能的研究还鲜有报道。【目的】利用高通量测序技术探究日本囊对虾肠道菌群的组成结构与功能作用,揭示虾体肠道菌群与外源菌群结构间的相关性。【方法】60 d的养殖周期结束后,分别采集日本囊对虾肠道样品(归为虾肠组,n=3)、养殖水体样品(归为水体组,n=3)和对虾饲料样品(归为饲料组,n=3),提取各样品总DNA进行16SrRNA基因扩增子测序,基于生物信息学方法分析与比较样品间的菌群结构特征,并使用PICRUSt软件预测日本囊对虾肠道菌群功能。【结果】3组样品测序共获得822 713条有效序列,抽平处理后可聚类为3 416个OTU。虾肠组样品中有28.49%、59.30%的OTU可以依次在水体组、饲料组样品中检测到。门水平上,虾肠组样品中的优势菌门为变形菌门(Proteobacteria)、拟杆菌门(Bacteroidetes)、厚壁菌门(Firmicutes)和梭杆菌门(Fusobacteria)。水体组、饲料组与虾肠组样品中的优势菌门结构不尽相同,但均由变形菌门和拟杆菌门组成。属水平上,虾肠组样品中的优势菌属包括弧菌属(Vibrio)、另类弧菌属(Aliivibrio)、假交替单胞菌属(Pseudoalteromonas)、假黄棕杆菌属(Pseudofulvibacter)、科尔韦尔氏菌属(Colwellia)、小纺锤状菌属(Fusibacter)、发光杆菌属(Photobacterium)、脱硫弧菌属(Desulfovibrio)、嗜冷杆菌属(Psychrobacter)以及弓形杆菌属(Arcobacter)。水体组和饲料组中检出的核心菌属结构与虾肠组相比有明显差异,其中海命菌属(Marivita)和假单胞菌属(Pseudomonas)分别为养殖水体及对虾饲料样品中的最优势菌属。PICRUSt预测结果显示,日本囊对虾肠道菌群的基因功能主要与新陈代谢类功能有关,包含氨基酸代谢、碳水化合物代谢与能量代谢等。【结论】日本囊对虾肠道菌群与其他种类对虾肠道菌群的结构间存在共性,其形成在一定程度上受到了外源菌群的干预,并在虾体的日常代谢活动中发挥了一定的作用。     

SKULL MORPHOLOGY AND PHYLOGENETIC RELATIONSHIPS OF A NEW DIMINUTIVE BALAENID FROM THE LOWER PLIOCENE OF BELGIUM

Abstract:  A new small balaenid is described and compared to all fossil and living balaenid taxa. The specimen represents a new genus and species and is named Balaenella brachyrhynus . It was discovered in the Lower Pliocene of Kallo (north-west Antwerp, Belgium) and adds new information on the diversity and evolution of Balaenidae. Based on both comparative morphology and phylogenetic analysis, Balaenella brachyrhynus is morphologically closer to the genus Balaena , including the living Greenland Bowhead whale ( B. mysticetus ), and two Pliocene species ( B. montalionis and B. ricei ) from central Italy and the eastern USA. Balaenella brachyrhynus has very short nasals, a short rostrum relative to the total skull length and a horizontal supraoccipital. A cladistic treatment of 81 morphological character states scored for 10 balaenids and nine non-balaenid cetaceans revealed that the other small balaenids generally included in the genus Balaenula (including Balaenula astensis, B. balaenopsis and a Pliocene Balaenula sp. from Japan) are closer to the living genus Eubalaena (the Right whale). As the new skull is so different from the nominal Balaenula species, and as it is more closely related to Balaena than to Eubalaena , it is concluded that a small body size was a common condition in different Balaenidae clades.     

The Osedax trophosome: organization and ultrastructure

The polychaete family Siboglinidae, which is currently construed as comprising the Frenulata, Monilifera (composed of Sclerolinum), Vestimentifera, and Osedax, has become known for its specialized symbiont-housing organ called the trophosome. This organ replaced the digestive system of the worms and is located in the elongated trunk region in Frenulata, Sclerolinum, and Vestimentifera. Currently two types of trophosomes have been described: in the taxa Frenulata and Sclerolinum the bacteriocytes originate from endoderm, and in Vestimentifera they originate from mesoderm. In Osedax, a trophosome was described as lacking (Rouse et al., 2004), but bacteriocytes are located in Osedax's characteristic root tissue. Here, we argue for a consistent name for the symbiont-housing tissue, namely trophosome, as in other siboglinids. In this study we provide morphological evidence that in Osedax the bacteriocytes are derived from somatic mesoderm. We show that the trophosome in Osedax is an apolar tissue composed of bacteriocytes and nonsymbiotic cells. As in vestimentiferans, a specific cell cycle was identified; however, in this case it is directed from the posterior to the anterior end of the worms instead of from the center toward the periphery. Comparison of all siboglinid trophosomes and re-evaluation of their body regions allows us to discuss whether the trophosomes are homologous and to hypothesize about the organization of the last common ancestor of Siboglinidae.     

APPEARANCE, DISTRIBUTION, AND GENETIC DISTINCTIVENESS OF LONGMAN'S BEAKED WHALE, INDOPACETUS PACIFICUS

Longman's beaked whale, Indopacetus pacificus , was known previously from only two skulls. Here we describe four new specimens of this species from strandings in the western and central Indian Ocean. Two juveniles, previously misidentified from external morphology as Hyperoodon planifrons , were identified as I. pacificus through diagnostic characteristics of mitochondrial (mt) DNA sequences derived from the holotype of this species. Images of the external appearance and teeth of the species are presented for the first time. Comparison of the color pattern of these new specimens with that of "tropical bottlenose whales" sighted in the tropical Indian and Pacific oceans confirm that those unidentified whales represent I. pacificus. Moore (1968) erected a new genus, Indopacetus , for this species (described initially as Mesoplodon pacificus ) based primarily on cranial morphology. Phylogenetic analyses of short mtDNA fragments available from the specimens known to date were unable to resolve the validity of this genus. However, the diagnostic osteological features highlighted by Moore (1968) for Indopacetus were also observed in the new specimens. Rib count and number of fused cervical vertebrae may also be diagnostic. Rostrum depth at mid-length and melon shape further distinguish this species from Mesoplodon beaked whales. As such, we see no reason on morphological grounds to overturn Moore's (1968) proposal that Longman's beaked whale is sufficiently distinct to be afforded its own genus.