Antitropical distribution and evolution in the Indo-West Pacific Ocean

Antitropical distributions of continental shelf, Indo-West Pacific species are probably not due to transgression of the tropics during the glacial periods, isothermic submergence, island integration, rising Neogene temperatures, or the Mesozoic dispersal of fragments from a Pacific continental mass. Characteristics of common antitropical patterns, plus information from systematic works on a variety on a variety of animal and plant groups, indicate that the long discarded "relict theory" of Theel (1885) appears to best fit the evidence, for it provides a mechanism whereby antitropical distribution may be brought about. The relict theory is compatible with the concept that the East Indies part of the Indo-West Pacific has been functioning as a center of evolutionary origin. It suggests that antitropical and associated disjunct patterns are produced as an older species, that has spread out to occupy a broad range, loses ground and gradually becomes supplanted by a younger species that had subsequently evolved in the East Indies. As this process goes on, the older species becomes restricted to a few isolated localities on the fringe of its original range. These isolates are often found to the north and south of the equatorial region but may include relict populations at the western edge of the Indian Ocean.     

Extinction and replacement in the Indo-West Pacific Ocean

Aim To provide evidence suggesting the existence of a dynamic system of extinction and replacement. Location The Indo-West Pacific Ocean. Methods Utilization of species distribution patterns produced by detailed systematic works. Results The distribution patterns appear to suggest a sequence of events that is consistent with the centre of origin hypothesis. The East Indies centre is considered to operate according to the centrifugal speciation model. Main conclusions Disjunct patterns appear to have originated in the East Indies and spread out from there. In cases where the vacated area is occupied by a sibling species, competitive (sympatric) speciation may have occurred. Over time the extinction pattern will gradually extend outward until it is played out among small populations far from their centre of origin     

Speciation and diversity on tropical rocky shores: a global phylogeny of snails of the genus Echinolittorina

A phylogenetic approach to the origin and maintenance of species diversity ideally requires the sampling of all species within a clade, confirmation that they are evolutionarily distinct entities, and knowledge of their geographical distributions. In the marine tropics such studies have mostly been of fish and reef-associated organisms, usually with high dispersal. In contrast, snails of the genus Echinolittorina (Littorinidae) are restricted to rocky shores, have a four-week pelagic development (and recorded dispersal up to 1400 km), and show different evolutionary patterns. We present a complete molecular phylogeny of Echinolittorina, derived from Bayesian analysis of sequences from nuclear 28S rRNA and mitochondrial 12S rRNA and COI genes (nodal support indicated by posterior probabilities, maximum likelihood, and neighbor-joining bootstrap). This consists of 59 evolutionarily significant units (ESUs), including all 50 known taxonomic species. The 26 ESUs found in the Indo-West Pacific region form a single clade, whereas the eastern Pacific and Atlantic species are basal. The earliest fossil occurred in the Tethys during the middle Eocene and we suggest that the Indo-West Pacific clade has been isolated since closure of the Tethyan seaway in the early Miocene. The geographical distributions of all species (based on more than 3700 locality records) appear to be circumscribed by barriers of low temperature, unsuitable sedimentary habitat, stretches of open water exceeding about 1400 km, and differences in oceanographic conditions on the continuum between oceanic and continental. The geographical ranges of sister species show little or no overlap, indicating that the speciation mode is predominantly allopatric. Furthermore, range expansion following speciation appears to have been limited, because a high degree of allopatry is maintained through three to five branching points of the phylogeny. This may be explained by infrequent long-distance colonization, habitat specialization on the oceanic/continental gradient, and perhaps by interspecific competition. In the eastern Pacific plus Atlantic we identify five cases of divergence on either side of the Isthmus of Panama, but our estimates of their ages pre-date the emergence of the Isthmus. There are three examples of sister relationships between species in the western Atlantic and eastern Atlantic, all resulting from dispersal to the east. Within the Indo-West Pacific, we find no geographical pattern of speciation events; narrowly endemic species of recent origin are present in both peripheral and central parts of the region. Evidence from estimated divergence times of sister species, and from a plot of the number of lineages over time, suggest that there has been no acceleration of diversification during the glacio-eustatic cycles of the Plio-Pleistocene. In comparison with reefal organisms, species of Echinolittorina on rocky shores may be less susceptible to extinction or isolation during sea-level fluctuations. The species richness of Echinolittorina in the classical biogeographic provinces conforms to the common pattern of highest diversity (11 species) in the central "East Indies Triangle" of the Indo-West Pacific, with a subsidiary focus in the eastern Pacific and western Atlantic, and lowest diversity in the eastern Atlantic. The diversity focus in the East Indies Triangle is produced by a mosaic of restricted allopatric species and overlap of a few widespread ones, and is the result of habitat specialization rather than historical vicariance. This study emphasizes the plurality of biogeographic histories and speciation patterns in the marine tropics.     

中国海结螺属四新纪录(腹足纲,骨螺科)

在对中国科学院海洋研究所历年来采集的骨螺科Muricidae标本进行整理分类时,共鉴定出结螺属Morula12种,其中4个中国新纪录：1）刺猬结螺Morula（M.）echinata（Reeve,1846）;2）紫结螺Morula（M.）purpureocincta（Preston,1909）;3）石优美结螺Morula（H.）lepida（Houart,1994）;4）白优美结螺Morula（H.）ambrosia（Houart,1994）.     

The first-stage zoeas of Carpilius convexus (Forskal, 1775) and Carpilius maculatus (Linnaeus, 1758) (Crustacea: Decapoda: Brachyura: Xanthoidea: Carpiliidae): an example of heterochrony

The first-stage zoeas of Carpilius convexus (Forskål,1775) and Carpilius maculatus (Linnaeus, 1758) are describedand fully illustrated. Both these Indo-West Pacific speciesexhibited a unique xanthoidean character for the basis of thesecond maxilliped, which possesses five (arranged 1,1,1,2) setaeinstead of the expected four (arranged 1,1,1,1). A comparisonwith the zoeal stages of Carpilius corallinus (Herbst, 1783)as reported by Laughlin et al. (Laughlin et al., 1983) revealedmarked differences including the possession of carapace lateralspines (v. absent in C. convexus only), the subterminal setationof the distal maxillule endopod segment with two subterminalsetae (versus one subterminal seta in Indo-West Pacific species),the terminal setation of the distal maxillule endopod segmentwith four setae (versus three setae in C. convexus only), twolateral spines on the telson (versus three in Indo-West Pacificspecies) and the number of zoeal stages. The first-stage zoeasof the two Indo-West Pacific species appear to have hatchedin a more advanced state of development than those of C. corallinus,and the expression of a number of characters has been accelerated(early onset). In fact, the zoeal stages of both Indo-West Pacificspecies and carpiliid species appear to be abbreviated becausethe first zoeas are considered to be equivalent to the third-stagezoeas of C. corallinus.     

Diversity of Scleractinia and Octocorallia in the mesophotic zone of the Great Barrier Reef, Australia

Mesophotic coral reefs in the Indo-West Pacific, the most diverse coral reef region on earth, are among the least documented. This study provides the first detailed investigation of the diversity of Scleractinia and Octocorallia of the mesophotic Great Barrier Reef (GBR). Specimens were collected by 100-m rock dredge tows at 47–163 m depth on 23 sites in four regions (15.3°–19.7° latitude South). Twenty-nine hard coral species from 19 families were recorded, with the greatest diversity found at <60 m depth, and no specimen was found >102 m. Many of these species are also commonly observed at shallower depths, particularly in inshore areas. Twenty-seven octocoral genera were collected, 25 of which represented azooxanthellate genera. Generic richness of octocorals was highest at depths >60 m. Sixteen of the 25 azooxanthellate genera were either absent or very rare at <18 m, and only five azooxanthellate genera were common on both shallow and mesophotic reefs. Species-area models indicated that the total diversity of hard corals on the deep mesophotic reefs sampled during this study was ~84 species while octocorals were represented by ~37 genera; however, the wide 95% confidence limits indicates that more intensive sampling effort is required to improve the accuracy of these estimates. Nonetheless, these results show that the taxonomic richness, particularly of hard corals, on mesophotic reefs may be much higher than previously thought, a finding that has implications for the comprehensive and adequate protection of the full range of biodiversity of the GBR.     

Migratory routes of red‐necked phalaropes Phalaropus lobatus breeding in southern Chukotka revealed by geolocators

The migration routes of red‐necked phalaropes breeding around the Bering Sea are poorly known, despite the fact that the Bering Sea could mark the boundary between the East Palearctic populations that winter in the Pacific Ocean around the East Indies and the West Nearctic populations that winter in the Pacific Ocean off the coast of South America. Geolocator data retrieved from two male phalaropes tagged in southern Chukotka, Far Eastern Russia, confirm that birds breeding in this region belong to the East Palearctic population and winter in the East Indies, suggesting that the division line with the West Nearctic population is farther to the east. The routes taken by the two phalaropes were almost entirely pelagic, totaling around 18 000–20 000 km round‐trip, with the birds continuously on the move during migration, rather than resident in any particular stopover site, contrary to most other migratory shorebirds.     

A phylogenetic test of multiple proposals for the origins of the East Indies coral reef biota

Aim  We analysed data on Indo-Pacific coral reef taxa to test four mechanisms proposed for the origins of the biodiversity centre in the East Indies Triangle: (1) the centre of origin hypothesis, (2) the centre of accumulation hypothesis, (3) the centre of overlap hypothesis, and (4) the bioaccumulation hypothesis.
Location  The Indian and western Pacific oceans.
Methods  The data set consisted of eight clades of fishes, four of corals and one of molluscs, consisting of a total of 95 species, with 72 informative nodes, in 29 areas. We used phylogenetic analysis for comparing trees (PACT) to reconstruct reticulate area relationships.
Results  An analysis using PACT produced very little congruence between area relationships among the histories of the taxa examined. Only two sets of two taxa each showed substantial congruence; several other taxa showed partial congruence.
Main conclusions  Despite numerous 'explanatory' claims for the incredible biodiversity of the marine macrofauna of the East Indies Triangle, the patterns obtained here did not accord with any of the hypotheses proposed. It is possible that, with the addition of more taxa, additional patterns would emerge. Much more systematic work within the East Indies is required to resolve this problem.     

Oceanic barriers as indicated by scombrid fishes and their parasites

Four genera of scombrid fishes (26 species) and their copepod (32 species) and monogenean ectoparasites (25 species) were used to test the hypothesis that the East Pacific Barrier is responsible for the most pronounced break in the circum-tropical warm water fauna of the continental shelves, and not the New World Barrier. Analysis at the species level showed that there is a primary centre of diversity in the West Pacific, and a secondary centre in the West Atlantic. The former, almost entirely, shares its species of the largely coastal Scomberomorus and Grammatorcynus and their parasites with seas located to the west. Only four parasites (all copepods) are shared by the East and West Pacific, and they are circum-tropical. In contrast, the West Pacific shares species of the more pelagic Scomber and their parasites with seas both to the east and west, although at the genus level, only two circum-tropical monogenean genera are shared by the E and W Pacific. We conclude that the East Pacific Barrier has been a 100% or almost 100% effective barrier to dispersal of species of Scomberomorus, Grammatorcynus and their parasites, whereas for species of Scomber and their parasites, the East Pacific has been a less effective barrier.     

The Chromodorididae (Opisthobranchia: Mollusca) of the Indo-West Pacific: Chromodoris epicuria, C. aureopurpurea, C. annulata, C. coi and Risbecia tryoni colour groups

Twenty-five species of chromodorid nudibranchs from the Indo-West Pacific are described including nine which are new to science. There are six new species of Chromodoris , two of Nounea and a new species which forms the type of a new monotypic genus. Of the other species discussed in this paper, ten are species of Chromodoris , five Risbecia and one Digidentis. Species with the following colour patterns are discussed: white with orange or yellow spots; white with yellow (or orange) and purple spots; white with purple spots or patches and mantle border; white with a sinuous submarginal brown band. A full review of the relevant literature is included as is a discussion on Indian Ocean and West Pacific species pairs.     

The Chromodorididae (Opisthobranchia: Mollusca) of the Indo-West Pacific: Chromodoris splendida, C. aspersa and Hypselodoris placida colour groups

Eleven species of Chromodoris, two species of Hypselodoris, three species of Mexichromis and one species of Noumea are described from the Indo-West Pacific, of which four species of Chromodoris and one of Mexichromis are new species. All species have a colour pattern dominated by red, purple or black spots. All previously described species with similar colour patterns, from both the Indo-West Pacific and other regions, are discussed.     

The Chromodorididae (Opisthobranchia: Mollusca) of the Indo-West Pacific: Chromodoris aureomarginata, C. verrieri and C. fide lis colour groups

Nineteen species of chromodorid nudibranchs are described including three new species of Chromodoris , two of Glossodoris and one each of Noumea, Cadlina and Thorunna from the Indo-West Pacific. The colour of all species is white with either a single yellow or gold band at the edge of the mantle or a double band of red and yellow, or orange and red or purple. All previously described species with similar colour patterns, from both the Indo-West Pacific and other regions are discussed.     

The European Tertiary Neritiliidae (Mollusca, Gastropoda, Neritopsina): indicators of tropical submarine cave environments and freshwater faunas

The oldest freshwater neritiliid, Neritilia bisinuata , is described from the Middle Eocene of the Loire Basin. Another European species, N. neritinoides , ranging from the Lower Oligocene to Lower Miocene (Upper Burdigalian) is recognized; its habitat appears to have been freshwater, but very close to the sea. Two new marine neritiliid species from the Aquitaine Basin are described: Bourdieria favia sp. nov. from the Upper Oligocene and Pisulinella aucoini  sp. nov. from the Lower Miocene. A third undescribed species from the Lower Miocene is referred to the same family and related to Pisulinella . The Oligocene species has a strong spiral sculpture, a character completely absent in previously known neritiliid species. The genus Agapilia , founded on juvenile N. neritinoides and adult Vitta picta , appears to be a junior synonym of the genus Vitta. The associated occurrence of shells of the families Neritiliidae, Neritopsidae and Pickworthiidae (well-known inhabitants of Indo-West Pacific submarine caves) at Peyrère suggest the first occurrence of a characteristic assemblage of dark submarine caves during the Oligocene. Both factorial analysis and relative abundance show that at Peyrère these families are associated with other cryptic fossils (various gastropods, bivalves, Brachiopoda, corals, Annelida). However, there are indications of other submarine cave assemblages in various Cenozoic deposits from the Palaeocene to the middle Miocene.  © 2004 The Linnean Society of London, Zoological Journal of the Linnean Society , 2004, 140 , 447–467.     

A review of the genus Janua, including Dexiospira (Polychaeta: Spirorbinae)

Janua (Dexiospira) brasiliensis and J. (D.) foraminosa are fully described for the first time and compared by a pictorial key with other species of Dexiospira and Pillaiospira . The former is very close to J. (D.) pseudocorrugata (= "Sptrorbis corrugatus" ) and particularly to J. (D.) nipponica , whilst the latter occupies a central position in the group of five species (including J. (D.) steueri ) which are characterized by having large four-lobed juvenile talons.
Janua (Janua) pagenstecheri is the most widespread species of the genus, whereas J. (Leodora) knightjonesi seems to be confined to coral seas. In all, 14 species of Janua are recognized, whilst 19 names are here regarded as synonyms or varieties. Eleven other species sometimes placed in Dexiospira or Leodora should rather constitute the genera Circeis, Sinistrella and Romanchella . These are re-defined and their affinities are studied.
The subgenus Dexiospira seems to be centred in the Indo-West Pacific, where it reaches maximum species-diversity and extension into cooler latitudes. J. (D.) pseudocorrugata var. mannarensis is pantropical and overlaps in the Mediterranean with the west European form and in the West Pacific with vars praeceps and fenestrata , and with subspecies ainu , which are characterized by small modifications of the tube or abdominal setae.     

Mitochondrial DNA variation in Indo-Pacific populations of the giant tiger prawn,Penaeus monodon

Surveys of mitochondrial DNA (mtDNA) variation in the giant tiger prawn, Penaeus monodon, using restriction fragment length polymorphisms have provided the first clear evidence that the Indo-West Pacific region is a site of accumulation of genetic diversity rather than a site of origin of genetic diversity. No haplotyes were found in common between a group of five southeast African populations and a group of five Australian (including Western Australia) and three southeast Asian populations. The dominant haplotype was different in the Australian and southeast Asian population groups. Genetic diversity (pi) was greatest in Indonesia (pi averaged 0.05), less in the Philippines and Australia (pi averaged 0.01), and markedly less in the southeast African and the West Australian populations (pi averaged 0.003). The high diversity of the southeast Asian populations resulted from the occurrence in those populations of a set of haplotypes found only in southeast Asia but derived from the southeast African haplotypes. These genetic variants therefore evolved in the Indian Ocean and later migrated into the Indo-West Pacific region. Low genetic variation in the geographically marginal populations in southeast Africa and Western Australia is considered to be the result of bottlenecks, but mismatch distributions suggest that large population sizes have been maintained in Indonesian populations for long periods.     

Copepoda from deep-sea hydrothermal vents and cold seeps

Recent explorations of hydrothermal vents in the eastern Pacific (Juan de Fuca spreading zone, Guaymas Basin in the Gulf of California, East Pacific Rise at 21° N and 13° N, and Galapagos Rift) and on the Mid-Atlantic Ridge have revealed many copepods, mostly siphonostomatoids with few poecilostomatoids. In these habitats in depths from 1 808 to 3 650 m water temperatures may reach nearly 15 ° C. Among more than 22 000 copepods from vents examined two new families, 11 new genera, and 32 new species were represented.In addition, two new copepods were found in 3 260 m at cold seeps at the base of the West Florida Escarpment in the Gulf of Mexico, an environment not thermally active, with water temperatures about 4.39 °C.Some of these copepods were associated with host invertebrates such as a Nuculana-like protobranch bivalve, a polychaete, and two species of shrimps. Others were obtained from washings of bivalves or vestimentiferans or by means of corers or slurp guns.     

Highly restricted gene flow and deep evolutionary lineages in the giant clam <Emphasis Type="Italic">Tridacna maxima</Emphasis>

The tropical Indo-West Pacific is the biogeographic region with the highest diversity of marine shallow water species, with its centre in the Indo-Malay Archipelago. However, due to its high endemism, the Red Sea is also considered as an important centre of evolution. Currently, not much is known about exchange among the Red Sea, Indian Ocean and West Pacific, as well as connectivity within the Indo-Malay Archipelago, even though such information is important to illuminate ecological and evolutionary processes that shape marine biodiversity in these regions. In addition, the inference of connectivity among populations is important for conservation. This study aims to test the hypothesis that the Indo-Malay Archipelago and the Red Sea are important centres of evolution by studying the genetic population structure of the giant clam Tridacna maxima. This study is based on a 484-bp fragment of the cytochrome c oxidase I gene from 211 individuals collected at 14 localities in the Indo-West Pacific to infer lineage diversification and gene flow as a measure for connectivity. The analysis showed a significant genetic differentiation among sample sites in the Indo-West Pacific (Φ_st = 0.74, P < 0.001) and across the Indo-Malay Archipelago (Φ_st = 0.72, P < 0.001), indicating restricted gene flow. Hierarchical AMOVA revealed the highest fixation index (Φ_ct = 0.8, P < 0.001) when sample sites were assigned to the following regions: (1) Red Sea, (2) Indian Ocean and Java Sea, (3) Indonesian throughflow and seas in the East of Sulawesi, and (4) Western Pacific. Geological history as well as oceanography are important factors that shape the genetic structure of T. maxima in the Indo-Malay Archipelago and Red Sea. The observed deep evolutionary lineages might include cryptic species and this result supports the notion that the Indo-Malay Archipelago and the Red Sea are important centres of evolution. Communicated by Biology Editor Dr. Ruth Gates     

On the affinities of three fossil ocypodid crabs and their relevance to the time and place of origin of the genus Macrophthalmus (Crustacea: Brachyura)

All the species of Macrophthalmus described from fossil material belong to the still extant subgenus Venitus. M. vindobonensis and M. aquensis , the two earliest known species of the genus, are closely related to the extant M. latreillei , while M. guamensis is conspecific with M. leptophthalmus.
Doubt is cast on Remy's (1952) theory of the late Tertiary migration of Macrophthalmus from the Mediterranean, its suggested place of origin, to the Indo-West Pacific, its present distributional range. Instead it is proposed that Macrophthalmus arose in the Indo-West Pacific during the late Eocene-Oligocene, and extended its range into the Mediterranean during the early-middle Miocene. The Mediterranean species became isolated during the late Miocene by a land bridge, and probably became extinct following the elevation of their brackish habitat during the late Miocene-Pliocene.