Eastern Pacific Barrier Research Articles

Geographic genetic variation in the Coral Hawkfish, Cirrhitichthys oxycephalus (Cirrhitidae), in relation to biogeographic barriers across the Tropical Indo-Pacific

The Tropical Indo-Pacific (TIP) includes about two thirds of the world’s tropical oceans and harbors an enormous number of marine species. The distributions of those species within the region is affected by habitat discontinuities and oceanographic features. As well as many smaller ones, the TIP contains seven large recognized biogeographic barriers that separate the Red Sea and Indian Ocean, the Indian from the Pacific Ocean, the central and eastern Pacific, the Hawaiian archipelago, the Marquesas and Easter Islands. We examined the genetic structuring of populations of Cirrhitichthys oxycephalus, a small cryptic species of reef fish, across its geographic range, which spans the longitudinal limits of the TIP. We assessed geographic variation in the mitochondrial cytb gene and the nuclear RAG1 gene, using 166 samples collected in 46 localities from the western to eastern edges of the TIP. Sequences from cytb show three well-structured groups that are separated by large genetic distances (1.58–2.96%): two in the Tropical Eastern Pacific (TEP), one at Clipperton Atoll another occupying the rest of that region and the third that ranges across the remainder of the TIP, from the central Pacific to the Red Sea and South Africa. These results indicate that the ~4,000 km wide Eastern Pacific Barrier between the central and eastern Pacific is an efficient barrier separating the two main groups. Further, the ~950 km of open ocean that isolates Clipperton Atoll from the rest of the TEP is also an effective barrier. Contrary to many other cases, various major and minor barriers from the Central Indo-Pacific to the Red Sea are not effective against dispersal by C. oxycephalus, although this species has not colonized the Hawiian islands and Easter Island. The nuclear gene partially supports the genetic structure evident in cytb, although all haplotypes are geographically mixed.

First records of two rays and three bony fishes for the Galapagos Islands

Abstract The Galapagos Islands lie within the oceanic ecoregion of the Tropical Eastern Pacific, which has a unique fish assemblage composition due to the influence of several ocean currents and El Niño Southern Oscillation (ENSO) events. In the El Niño phase of these events, water temperature changes facilitate the movement of fish species between oceanic ecoregions, as well as across the Eastern Pacific Barrier. Here, we present five new fish records for the Galapagos Marine Reserve based on underwater imagery. These include two rays (Mobula thurstoni and Myliobatis longirostris) and three bony fishes (Lobotes pacifica, Lutjanus colorado and Sphyraena stellata). Of these, the first species is proposed as potentially resident to the Galapagos, and the latter four as vagrant species in the Galapagos until further sightings can conclusively determine their status. The effects of ENSO, the use of underwater video technology, and the importance of up-to-date and accurate species listings to understand the impact of the climate crisis are discussed.

Ancient Tethyan Vicariance and Long-Distance Dispersal Drive Global Diversification and Cryptic Speciation in the Red Seaweed Pterocladiella.

We investigated the globally distributed red algal genus Pterocladiella, comprising 24 described species, many of which are economically important sources of agar and agarose. We used DNA-based species delimitation approaches, phylogenetic, and historical biogeographical analyses to uncover cryptic diversity and infer the drivers of biogeographic patterns. We delimited 43 species in Pterocladiella, of which 19 are undescribed. Our multigene time-calibrated phylogeny and ancestral area reconstruction indicated that Pterocladiella most likely originated during the Early Cretaceous in the Tethys Sea. Ancient Tethyan vicariance and long-distance dispersal have shaped current distribution patterns. The ancestor of Eastern Pacific species likely arose before the formation of the formidable Eastern Pacific Barrier—a first confirmation using molecular data in red algae. Divergences of Northeast and Southeast Pacific species have been driven by the Central American Seaway barrier, which, paradoxically, served as a dispersal pathway for Atlantic species. Both long- and short-distance dispersal scenarios are supported by genetic relationships within cosmopolitan species based on haplotype analysis. Asymmetrical distributions and the predominance of peripatry and sympatry between sister species suggest the importance of budding speciation in Pterocladiella. Our study highlights the underestimation of global diversity in these crucial components of coastal ecosystems and provides evidence for the complex evolution of current species distributions.

The Eastern Tropical Pacific coral population connectivity and the role of the Eastern Pacific Barrier

Long-distance dispersal is believed to strongly influence coral reef population dynamics across the Tropical Pacific. However, the spatial scale and strength at which populations are potentially connected by dispersal remains uncertain. To determine the patterns in connectivity between the Eastern (ETP) and Central Tropical Pacific (CTP) ecoregions, we used a biophysical model incorporating ocean currents and larval biology to quantify the seascape-wide dispersal potential among all population. We quantified the likelihood and determined the oceanographic conditions that enable the dispersal of coral larvae across the Eastern Pacific Barrier (EP-Barrier) and identified the main connectivity pathways and their conservation value for dominant reef-building corals. Overall, we found that coral assemblages within the CTP and ETP are weakly connected through dispersal. Although the EP-Barrier isolates the ETP from the CTP ecoregion, we found evidence that the EP-Barrier may be breached, in both directions, by rare dispersal events. These rare events could explain the evolutionary genetic similarity among populations of pocilloporids in the ecoregions. Moreover, the ETP may function as a stronger source rather than a destination, providing potential recruits to CTP populations. We also show evidence for a connectivity loop in the ETP, which may positively influence long-term population persistence in the region. Coral conservation and management communities should consider eight-key stepping stone ecoregions when developing strategies to preserve the long-distance connectivity potential across the ETP and CTP.

Strong trans-Pacific break and local conservation units in the Galapagos shark (Carcharhinus galapagensis) revealed by genome-wide cytonuclear markers.

The application of genome-wide cytonuclear molecular data to identify management and adaptive units at various spatio-temporal levels is particularly important for overharvested large predatory organisms, often characterized by smaller, localized populations. Despite being "near threatened", current understanding of habitat use and population structure of Carcharhinus galapagensis is limited to specific areas within its distribution. We evaluated population structure and connectivity across the Pacific Ocean using genome-wide single-nucleotide polymorphisms (~7200 SNPs) and mitochondrial control region sequences (945 bp) for 229 individuals. Neutral SNPs defined at least two genetically discrete geographic groups: an East Tropical Pacific (Mexico, east and west Galapagos Islands), and another central-west Pacific (Lord Howe Island, Middleton Reef, Norfolk Island, Elizabeth Reef, Kermadec, Hawaii and Southern Africa). More fine-grade population structure was suggested using outlier SNPs: west Pacific, Hawaii, Mexico, and Galapagos. Consistently, mtDNA pairwise ΦST defined three regional stocks: east, central and west Pacific. Compared to neutral SNPs (FST = 0.023-0.035), mtDNA exhibited more divergence (ΦST = 0.258-0.539) and high overall genetic diversity (h = 0.794 ± 0.014; π = 0.004 ± 0.000), consistent with the longstanding eastern Pacific barrier between the east and central-west Pacific. Hawaiian and Southern African populations group within the west Pacific cluster. Effective population sizes were moderate/high for east/west populations (738 and 3421, respectively). Insights into the biology, connectivity, genetic diversity, and population demographics informs for improved conservation of this species, by delineating three to four conservation units across their Pacific distribution. Implementing such conservation management may be challenging, but is necessary to achieve long-term population resilience at basin and regional scales.

Historical Biogeography of Delphininae Dolphins and Related Taxa (Artiodactyla: Delphinidae)

Delphinine dolphins arose via a recent, rapid radiation, probably within the last four million years. Although molecular phylogenies are increasingly well resolved, patterns of morphology-ecology-geography are hard to link to phylogeny or to translate into taxonomy. Such problems might be tackled through understanding the drivers of the delphinine radiation. Here, we examine delphinine historical biogeography using the phylogeny of McGowen et al. (Mol Phylogenet Evol 53:891–906, 2009) as our working hypothesis. We used the “Spatial Analysis of Vicariance” method to delimit modern distribution patterns, including disjunctions involving sister nodes in the Delphininae. The analysis identified disjunct sister nodes, allowing some interpretation of Delphininae biogeography. The Central American Seaway was probably an important gateway for early delphinids, but the succeeding “hard” barrier of the Panama Isthmus had little influence. Southern African waters form the Atlantic-Indo-Pacific gateway, which is sometimes considered a “soft” barrier because of the variation in the Benguela and Agulhas currents, in turn driven by tectonic changes and/or Pleistocene glacial and interglacial cycles. The latter cycles probably fragmented coastal habitats, allowing allopatric speciation. Geological patterns of turnover in Southern Ocean diatoms, which link to physical oceanic change, closely match the main cluster of delphinine divergences. The Eastern Pacific Barrier, and perhaps the associated Humboldt Current and equatorial “cold tongue,” affect modern distributions, but cause and effect are poorly understood. Future research should involve molecular-morphological phylogenetics for all species, subspecies, and ecomorphs. Complete distributions must be known for all taxa to understand how vicariance and dispersal shaped the distribution of delphinines.

Adaptive radiation of damselfishes (Perciformes, Pomacentridae) in the eastern Pacific

Pomacentridae is one of the most abundant families in tropical and temperate rocky and coral reefs. They present an extraordinary diversity of habitat preferences, feeding, morphologies and behavior. The eastern Pacific is biogeographically isolated by the Isthmus of Panama and the eastern Pacific barrier. There is an agreement about the origin of the fauna of the Tropical Eastern Pacific, suggesting three main factors, mixture, dispersal and vicariance. In this study, by cluster analyses and parsimony analysis of endemism, the distribution of damselfishes within the eastern Pacific was analyzed to elucidate the provinciality and the history of their radiation. Six main provinces were found: (1) Easter Pacific equatorial islands, (2) North, (3) Center, (4) South, (5) California Province and (6) Clipperton. The Gulf of California and Galapagos islands are the two main centers of species richness. Three main factors limited the radiation of damselfishes: in islands, the distance of the island to the mainland and the size of the island; while in the mainland, the temperature appears to be the main barrier. Our results show the radiation process of Pomacentridae in the eastern Pacific as a dynamic dispersion system, which can be categorized in three main steps: (1) mixture and speciation of species with close affinity to west Atlantic ancestral stocks in the Central Province, (2) dispersion due to favorable conditions to Galapagos islands and Gulf of California and (3) the more complex and perhaps long, gradual dispersal and radiation to temperate areas and isolated or marginal environments.

A sea water barrier to coral gene flow

Land is not the only barrier to dispersal encountered by marine organisms. For sedentary shallow water species, there is an additional, marine barrier, 5000 km of uninterrupted deep-water stretch between the central and the eastern Pacific. This expanse of water, known as the ‘Eastern Pacific Barrier’, has been separating faunas of the two oceanic regions since the beginning of the Cenozoic. Species with larvae that cannot stay in the plankton for the time it takes to cross between the two sides have been evolving independently. That the eastern Pacific does not share species with the rest of the Pacific was obvious to naturalists two centuries ago (Darwin 1860). Yet, this rule has exceptions. A small minority of species are known to straddle the Eastern Pacific Barrier. One such exception is the scleractinian coral Porites lobata (Fig. 1). This species is spread widely throughout the Indo-Pacific, where it is one of the major reef-builders, but it is also encountered in the eastern Pacific. Are eastern and central Pacific populations of this coral connected by gene flow? In this issue of Molecular Ecology, Baums et al. (2012) use microsatellite data to answer this question. They show that P. lobata populations in the eastern Pacific are cut off from genetic influx from the rest of the Pacific. Populations within each of the two oceanic regions are genetically connected (though those in the Hawaiian islands are also isolated). Significantly, the population in the Clipperton Atoll, the westernmost island in the eastern Pacific, genetically groups with populations from the central Pacific, suggesting that crossing the Eastern Pacific Barrier by P. lobata propagules does occasionally occur.

No gene flow across the Eastern Pacific Barrier in the reef‐building coral Porites lobata

The expanse of deep water between the central Pacific islands and the continental shelf of the Eastern Tropical Pacific is regarded as the world's most potent marine biogeographic barrier. During recurrent climatic fluctuations (ENSO, El Niño Southern Oscillation), however, changes in water temperature and the speed and direction of currents become favourable for trans-oceanic dispersal of larvae from central Pacific to marginal eastern Pacific reefs. Here, we investigate the population connectivity of the reef-building coral Porites lobata across the Eastern Pacific Barrier (EPB). Patterns of recent gene flow in samples (n = 1173) from the central Pacific and the Eastern Tropical Pacific (ETP) were analysed with 12 microsatellite loci. Results indicated that P. lobata from the ETP are strongly isolated from those in the central Pacific and Hawaii (F(ct) ' = 0.509; P < 0.001). However, samples from Clipperton Atoll, an oceanic island on the eastern side of the EPB, grouped with the central Pacific. Within the central Pacific, Hawaiian populations were strongly isolated from three co-occurring clusters found throughout the remainder of the central Pacific. No further substructure was evident in the ETP. Changes in oceanographic conditions during ENSO over the past several thousand years thus appear insufficient to support larval deliveries from the central Pacific to the ETP or strong postsettlement selection acts on ETP settlers from the central Pacific. Recovery of P. lobata populations in the frequently disturbed ETP thus must depend on local larval sources.

Global diversification of mangrove fauna: a molecular phylogeny of Littoraria (Gastropoda: Littorinidae)

The genus Littoraria is one of very few molluscan groups that are closely associated with mangroves. We document its global evolutionary radiation and compare biogeographic patterns with those of mangrove plants, based on phylogenetic and fossil evidence. Using sequences from three genes (nuclear 28S rRNA, mitochondrial 12S rRNA and COI) we reconstruct a phylogeny of 37 of the 39 living morphospecies. Six monophyletic subgenera are defined (Bulimilittorina, Lamellilitorina, Littoraria, Palustorina, Protolittoraria, Littorinopsis) and we synonymize L. coccinea and L. glabrata. A deep division between Palustorina from the Indo-West Pacific and Littoraria from the Atlantic and Eastern Pacific is estimated by a Bayesian relaxed-clock method to be of Middle Eocene to Palaeocene age (43.2–62.7Ma), which far predates the Early Miocene (18Ma) closure of the Tethyan Seaway; this, as in mangrove plants, may reflect vicariance by climatic cooling, rather than tectonic processes. The age of Littoraria angulifera in the Atlantic is, however, consistent with Early Miocene vicariance of a Tethyan ancestor. We infer that speciation events are mainly of Miocene or older age, and that diversification has not been driven by depletion of mangrove habitats during recent glacial intervals. Parsimonious reconstruction of ancestral habitats suggests that the genus has inhabited mangrove or wood substrates since its origin, while the rock-dwelling habit of the four members of Protolittoraria is derived. Three species span the Eastern Pacific Barrier, and one is amphi-Atlantic, consistent with a long larval phase of up to 10weeks. Allopatric speciation is inferred, but usually with subsequent range overlap. Ovoviviparity (interpreted as an adaptation to life in mangroves) has arisen twice.

Crossing the impassable: genetic connections in 20 reef fishes across the eastern Pacific barrier

The 'impassable' Eastern Pacific Barrier (EPB), ca 5000 km of deep water separating the eastern from the central Pacific, is the World's widest marine biogeographic barrier. Sequencing of mitochondrial DNA in 20 reef fish morphospecies encountered on both sides of the barrier revealed cryptic speciation in two. Among the other 18 species only two showed significant differentiation (as revealed by haplotype networks and FST statistics) between the eastern and the central Pacific. Coalescence analyses indicated that genetic similarity in the 18 truly transpacific species resulted from different combinations of ages of most recent invasion and of levels of recurrent gene flow, with estimated times of initial separation ranging from approximately 30000 to 1 Myr (ago). There is no suggestion of simultaneous interruptions of gene flow among the species. Migration across the EPB was previously thought to be exclusively eastward, but our evidence showed two invasions from east to west and eight cases in which subsequent gene flow possibly proceeded in the same direction. Thus, the EPB is sporadically permeable to propagules originating on either side.

Tropical Transpacific Shore Fishes

Tropical Transpacific Shore Fishes

PHYLOGEOGRAPHY OF THE PANTROPICAL SEA URCHIN TRIPNEUSTES: CONTRASTING PATTERNS OF POPULATION STRUCTURE BETWEEN OCEANS

To understand how allopatric speciation proceeds, we need information on barriers to gene flow, their antiquity, and their efficacy. For marine organisms with planktonic larvae, much of this information can only be obtained through the determination of divergence between populations. We evaluated the importance of ocean barriers by studying the mitochondrial DNA phylogeography of Tripneustes, a pantropical genus of shallow water sea urchin. A region of cytochrome oxidase I (COI) was sequenced in 187 individuals from locations around the globe. The COI phylogeny agreed with a previously published phylogeny of bindin that barriers important to the evolution of Tripneustes are: (1) the cold water upwelling close to the tip of South Africa, (2) the Isthmus of Panama, (3) the long stretch of deep water separating the eastern from the western Atlantic, and (4) the freshwater plume of the Orinoco and the Amazon rivers between the Caribbean and the coast of Brazil. These barriers have previously been shown to be important in at least a subset of the shallow water marine organisms in which phylogeography has been studied. In contrast, the Eastern Pacific Barrier, 5000 km of deep water between the central and the eastern Pacific that has caused the deepest splits in other genera of sea urchins, is remarkably unimportant as a cause of genetic subdivision in Tripneustes. There is also no discernible subdivision between the Pacific and Indian Ocean populations of this genus. The most common COI haplotype is found in the eastern, central, and western Pacific as well as the Indian Ocean. Morphology, COI, and bindin data agree that T. depressus from the eastern Pacific and T. gratilla from the western Pacific are, in fact, the same species. The distribution of haplotype differences in the Indo-Pacific exhibits characteristics expected from a sea urchin genus with ephemeral local populations, but with high fecundity, dispersal, and growth: there is little phylogenetic structure, and mismatch distributions conform to models of recent population expansion on a nearly global scale. Yet, comparisons between local populations produce large and significant FST values, indicating nonrandom haplotype distribution. This apparent local differentiation is only weakly reflected in regional divergence, and there is no evidence of isolation by distance in correlations between FST values and either geographical or current distance. Thus, Tripneustes in the Indo-Pacific (but not in the Atlantic) seems to be one large metapopulation spanning two oceans and containing chaotic, nonequilibrium local variation, produced by the haphazard arrival of larvae or by unpredictable local extinction.

Dispersal barriers in tropical oceans and speciation in Atlantic and eastern Pacific sea urchins of the genus Echinometra.

Echinometra is a pantropical sea urchin made famous through studies of phylogeny, speciation, and genetic structure of the Indo-West Pacific (IWP) species. We sequenced 630 bp of the cytochrome oxidase I (COI) mitochondrial gene to provide comparable information on the eastern Pacific and Atlantic species, using divergence between those separated by closure of the Isthmus of Panama 3.1 million years ago (Ma) to estimate dates for cladogenic events. Most recently (1.27-1. 62 Ma), the Atlantic species E. lucunter and E. viridis diverged from each other, at a time in the Pleistocene that sea levels fell and Caribbean coral speciation and extinction rates were high. An earlier split, assumed to have been coincident with the completion of the Isthmus of Panama, separated the eastern Pacific E. vanbrunti from the Atlantic common ancestor. Transisthmian COI divergence similar to that in the sea urchin genus Eucidaris supports this assumption. The most ancient split in Echinometra occurred between the IWP and the neotropical clades, due to cessation of larval exchange around South Africa or across the Eastern Pacific Barrier. Gene flow within species is generally high; however, there are restrictions to genetic exchange between E. lucunter populations from the Caribbean and those from the rest of the Atlantic. Correlation between cladogenic and vicariant events supports E. Mayr's contention that marine species, despite their high dispersal potential, form by means of geographical separation. That sympatric, nonhybridizing E. lucunter and E. viridis were split so recently suggests, however, that perfection of reproductive barriers between marine species with large populations can occur in less than 1.6 million years (Myr).

PHYLOGEOGRAPHY OF THE PANTROPICAL SEA URCHIN EUCIDARIS IN RELATION TO LAND BARRIERS AND OCEAN CURRENTS.

The pantropical sea urchin genus Eucidaris contains four currently recognized species, all of them allopatric: E. metularia in the Indo-West Pacific, E. thouarsi in the eastern Pacific, E. tribuloides in both the western and eastern Atlantic, and E. clavata at the central Atlantic islands of Ascension and St. Helena. We sequenced a 640-bp region of the cytochrome oxidase I (COI) gene of mitochondrial DNA to determine whether this division of the genus into species was confirmed by molecular markers, to ascertain their phylogenetic relations, and to reconstruct the history of possible dispersal and vicariance events that led to present-day patterns of species distribution. We found that E. metularia split first from the rest of the extant species of the genus. If COI divergence is calibrated by the emergence of the Isthmus of Panama, the estimated date of the separation of the Indo-West Pacific species is 4.7-6.4 million years ago. This date suggests that the last available route of genetic contact between the Indo-Pacific and the rest of the tropics was from west to east through the Eastern Pacific Barrier, rather than through the Tethyan Sea or around the southern tip of Africa. The second cladogenic event was the separation of eastern Pacific and Atlantic populations by the Isthmus of Panama. Eucidaris at the outer eastern Pacific islands (Galapagos, Isla del Coco, Clipperton Atoll) belong to a separate clade, so distinct from mainland E. thouarsi as to suggest that this is a different species, for which the name E. galapagensis is revived from the older taxonomic literature. Complete lack of shared alleles in three allozyme loci between island and mainland populations support their separate specific status. Eucidaris galapagensis and E. thouarsi are estimated from their COI divergence to have split at about the same time that E. thouarsi and E. tribuloides were being separated by the Isthmus of Panama. Even though currents could easily convey larvae between the eastern Pacific islands and the American mainland, the two species do not appear to have invaded each other's ranges. Conversely, the central Atlantic E. clavata at St. Helena and Ascension is genetically similar to E. tribuloides from the American and African coasts. Populations on these islands are either genetically connected to the coasts of the Atlantic or have been colonized by extant mitochondrial DNA lineages of Eucidaris within the last 200,000 years. Although it is hard to explain how larvae can cross the entire width of the Atlantic within their competent lifetimes, COI sequences of Eucidaris from the west coast of Africa are very similar to those of E. tribuloides from the Caribbean. FST statistics indicate that gene flow between E. metularia from the Indian Ocean and from the western and central Pacific is restricted. Low gene flow is also evident between populations of E. clavata from Ascension and St. Helena. Rates of intraspecific exchange of genes in E. thouarsi, E. galapagensis, and E. tribuloides, on the other hand, are high. The phylogeny of Eucidaris confirms Ernst Mayr's conclusions that major barriers to the dispersal of tropical echinoids have been the wide stretch of deep water between central and eastern Pacific, the cold water off the southwest coast of Africa, and the Isthmus of Panama. It also suggests that a colonization event in the eastern Pacific has led to speciation between mainland and island populations.

Massive gene flow across the world's most potent marine biogeographic barrier

The Eastern Pacific Barrier (EPB), 5400 km of uninterrupted deep water between the central and eastern Pacific, constitutes the greatest marine obstacle to the dispersal of shallowwater organisms. ...

Clipperton Atoll (eastern Pacific): oceanography, geomorphology, reef-building coral ecology and biogeography

Coral reef geomorphology and community composition were investigated in the tropical northeastern Pacific during April 1994. Three areas were surveyed in the Revillagigedo Islands (Mexico), and an intensive study was conducted on Clipperton Atoll (1,300 km SW of Acapulco), including macro-scale surface circulation, sea surface temperature (SST) climatology, geomorphology, coral community structure, zonation, and biogeography. Satellite-tracked drifter buoys from 1979–1993 demonstrated complex patterns of surface circulation with dominantly easterly flow (North Equatorial Counter Current, NECC), but also westerly currents (South Equatorial Current, SEC) that could transport propagules to Clipperton from both central and eastern Pacific regions. The northernmost latitude reached by the NECC is not influenced by El Nino-Southern Oscillation (ENSO) events, but easterly flow velocity evidently is accelerated at such times. Maximum NECC flow rates indicate that the eastern Pacific barrier can be bridged in 60 to 120 days. SST anomalies at Clipperton occur during ENSO events and were greater at Clipperton in 1987 than during 1982–1983. Shallow (15–18 m)and deep (50–58 m) terraces are present around most of Clipperton, probably representing Modern and late Pleistocene sea level stands. Although Clipperton is a well developed atoll with high coral cover, the reef-building fauna is depauperate, consisting of only 7 species of scleractinian corals belonging to the generaPocillopora, Porites, Pavona andLeptoseris, and 1 species of hydrocoral in the genusMillepora. The identities of the one Pocilpopora species and one of the twoPorites species are still unknown. Two of the remaining scleractinians (Pavona minuta, Leptoseris scabra) and the hydrocoral (Millepora exaesa), all formerly known from central and western Pacific localities, represent new eastern Pacific records. Scleractinian corals predominate (10–100% cover) over insular shelf depths of 8 to 60m, and crustose coralline algae are dominant (5–40% cover) from 0.5 to 7m. Spur and groove features, constructed of alternating frameworks ofPocillopora andPorites, and veneered with crustose coralline algae, are generally well developed around most atoll exposures. Although crustose coralline algae predominate in the breaker zone (with up to 100% cover), a prominent algal ridge is absent with only a slight buildup (ca. 10 cm) to seaward. Frequent grazing by the pufferfishArothron meleagris results in the removal of large amounts of live tissue and skeleton fromPorites lobata. Acanthaster planci is present, but rare. The grazing of large diadematid sea urchins, (2 species each ofDiadema andEchinothrix) on dead corals cause extensive erosion in some areas. Large numbers of corals on the 15–18 m terrace had recently suffered partial (P. lobata, 60–70% maximum of all colonies sampled) or total (Pocillopora sp., 80% maximum) mortality. The lengths of regenerating knobs and the rates of linear skeletal growth inP. lobata, determined by sclerochronologic analysis, indicated a period of stress during 1987. Massive skeletal growth is significantly higher at intermediate (16–17 m) than shallow (6–8 m) depths with mean extension rates of 1.5 mm yr−1 inP. lobata and 1.4 mm yr−1 inP. minuta at intermediate depths. Skeletal growth inP. lobata was depressed during the 1987 El Nifio event at Clipperton. The branching coralPocillopora sp. demonstrated high and similar skeletal growth rates at both shallow (25.4 mm yr−1) and intermediate (26.5 mm yr−1) depths. The presence of widely distributed Indo-Pacific zooxanthellate corals at Clipperton and the Revillagigedo Islands indicates that these NE Pacific Islands probably serve as a stepping stone for dispersal into the far eastern Pacific region.

The Genus Antemetula Rehder in the Indo-Westpacific Area, with the Description of Two New Fossil Species

Rehder (1943: 199) pointed out, that Buccinum clathratum Adams & Reeve, 1848, is the type of the genus Metula H.& A. Adams, 1858. For the species congeneric with Buccinum metula Hinds, 1844, he proposed the generic name Antemetula, with that species as the type. Bucchium metula was described by Hinds from a specimen dredged by the Sulphur off the West coast of Veragua, at a depth of a few fathoms. There are no other records of the species from the American West coast. Smith (1904: 465) called attention to the fact that young specimens of Buccinum mitrella Adams & Reeve, 1848, match Hinds’s figure of B. metula. B. mitrella was described from one or more specimens dredged by the Samarang in the China Sea, at a depth of 10 fathoms, and has been recorded since from some more localities in the Indo-Westpacific area. Smith thought B. mitrella to be a synonym of B. metula, and the type locality of the latter possibly erroneous. It is, indeed, unlikely that a benthonic prosobranch inhabiting the West coast of America between the tropics will also occur in the Indo-Westpacific area, because the eastern Pacific barrier is practically insurmountable for littoral species, as Ekman (1935: 105—107) has shown. If therefore Smith’s opinion, that B. mitrella is a synonym of B. metula be right, the type locality mentioned by Hinds must be wrong. The Sulphur dredged in the China Sea also, and therefore it seems possible that material from that sea was confounded with shells from the American West coast.