Tethyan vicariance, relictualism and speciation: evidence from a global molecular phylogeny of the opisthobranch genus Bulla

Aim Our aims were: (1) to reconstruct a molecular phylogeny of the cephalaspidean opisthobranch genus Bulla, an inhabitant of shallow sedimentary environments; (2) to test if divergence times are consistent with Miocene and later vicariance among the four tropical marine biogeographical provinces; (3) to examine the phylogenetic status of possible Tethyan relict species; and (4) to infer the timing and causes of speciation events. Location Tropical and warm‐temperate regions of the Atlantic, Indo‐West Pacific, Australasia and eastern Pacific. Methods Ten of the 12 nominal species of Bulla were sampled, in a total sample of 65 individuals, together with cephalaspidean outgroups. Phylogenetic relationships were inferred by Bayesian analysis of partial sequences of the mitochondrial cytochrome c oxidase I (COI) and 16S rRNA and nuclear 28S rRNA genes. Divergence times and rates of evolution were estimated using uncorrelated relaxed‐clock Bayesian methods with fossil calibrations (based on literature review and examination of fossil specimens), implemented in beast . The geographical pattern of speciation was assessed by estimating the degree of overlap between sister lineages. Results Four clades were supported: Indo‐West Pacific (four species), Australasia (one species), Atlantic plus eastern Pacific (three species) and Atlantic (two species), with estimated mean ages of 35–46 Ma. Nominal species were monophyletic, but deep divergences were found within one Indo‐West Pacific and one West Atlantic species. Species‐level divergences occurred in the Miocene or earlier. The age of a sister relationship across the Isthmus of Panama was estimated at 7.9–32.1 Ma, and the divergence of a pair of sister species on either side of the Atlantic Ocean occurred 20.4–27.2 Ma. Main conclusions Fossils suggest that Bulla originated in the Tethys realm during the Middle Eocene. Average ages of the four main clades fall in the Eocene, and far pre‐date the 18–19 Ma closure of the Tethys Seaway. This discrepancy could indicate earlier vicariant events, selective extinction or errors of calibration. Similarly, the transisthmian divergence estimate far pre‐dates the uplift of the Panamanian Isthmus at about 3 Ma. Speciation events occurred in the Miocene, consistent with tectonic events in the central Indo‐West Pacific, isolation of the Arabian Sea by upwelling and westward trans‐Atlantic dispersal. Differences in habitat between sister species suggest that ecological speciation may also have played a role. The basal position of the Australasian species supports its interpretation as a Tethyan relict.     

Seasonal changes in environmental variables,biomass, production and nutrient contents in two contrasting tropical intertidal seagrass beds in South Sulawesi,Indonesia

Seasonal dynamics were studied by monthly monitoring of biological and environmental variables in permanent quadrats in two contrasting intertidal seagrass beds in South Sulawesi, Indonesia, from February 1991 to January 1992. Datasets were analysed with canonical correlation analysis for correlations between environmental and biological variables. Considerable variation in biomass, production and plant tissue nutrient contents in a monospecific seagrass bed of Enhalus acoroides, growing on a coastal terrigenous mudbank (Gusung Tallang), was assumed to be related to riverine influences of the nearby Tallo River. The variation in seagrass variables at this site could, however, not be significantly correlated to seasonal patterns in rainfall, salinity, tides, nutrient availability, water motion or turbidity. A seasonal cycle in biomass, production and nutrient contents in a mixed seagrass bed of Thalassia hemprichii and E. acoroides, growing on carbonate sand on the reef flat of an offshore coral island (Barang Lompo), was found to be largely determined by tidal exposure and water motion. Exposure of the intertidal seagrass bed during hours of low water during spring tides showed a gradual shift from exposure during the night (January-June) to exposure during daylight (July-December). Daylight exposure resulted in a significant loss of above-ground plant biomass through desiccation and burning of leaves. The observed seasonal dynamics of the seagrass bed on reef sediment contrast with reports from the Caribbean, where the effect of tidal exposure on comparable shallow-water seagrass communities is relatively insignificant due to a small tidal amplitude.     

A new genus of rodent from Wallacea (Rodentia: Muridae: Murinae: Rattini), and its implication for biogeography and Indo‐Pacific Rattini systematics

We describe Halmaheramys bokimekot Fabre, Pagès, Musser, Fitriana, Semiadi & Helgen gen. et sp. nov. , a new genus and species of murine rodent from the North Moluccas, and study its phylogenetic placement using both molecular and morphological data. We generated a densely sampled mitochondrial and nuclear DNA data set that included most genera of Indo‐Pacific Murinae, and used probabilistic methodologies to infer their phylogenetic relationships. To reconstruct their biogeographical history, we first dated the topology and then used a Lagrange analysis to infer ancestral geographic areas. Finally, we combined the ancestral area reconstructions with temporal information to compare patterns of murine colonization among Indo‐Pacific archipelagos. We provide a new and comprehensive molecular phylogenetic reconstruction for Indo‐Pacific Murinae, with a focus on the Rattus division. Using previous results and those presented in this study, we define a new Indo‐Pacific group within the Rattus division, composed of Bullimus, Bunomys, Paruromys, Halmaheramys, Sundamys, and Taeromys. Our phylogenetic reconstructions revealed a relatively recent diversification from the Middle Miocene to Plio‐Pleistocene associated with several major dispersal events. We identified two independent Indo‐Pacific dispersal events from both western and eastern Indo‐Pacific archipelagos to the isolated island of Halmahera, which led to the speciations of H. bokimekot gen. et sp. nov. and Rattus morotaiensis Kellogg, 1945. We propose that a Middle Miocene collision between the Halmahera and Sangihe arcs may have been responsible for the arrival of the ancestor of Halmaheramys to eastern Wallacea. Halmaheramys bokimekot gen. et sp. nov. is described in detail, and its systematics and biogeography are documented and illustrated. © 2013 The Linnean Society of London     

Integrated sedimentological,ichnological and sequence stratigraphical studies of the Koti Dhaman Formation (Tal Group), Nigali Dhar Syncline,Lesser Himalaya,India: paleoenvironmental,paleoecological, paleogeographic significance

Abstract

Integrated ichnology, sedimentology and sequence stratigraphy of the Lower Quartzite Member to the Arkosic Sandstone Member of the Koti Dhaman Formation (Cambrian Series 2, Stage 4), Tal Group, Nigali Dhar Syncline, Lesser Himalayan lithotectonic zone are presented. Trilobite traces of Gondwanan affinity i.e., Cruziana salomonis, Cruziana fasciculata, Rusophycus dispar and Rusophycus burjensis are recorded along with Arenicolites isp. and Skolithos isp. from the Lower Quartzite Member. A rich and diverse ichnoassemblage attributed to the Cruziana ichnofacies is described for the first time from the Arkosic Sandstone Member of the same formation. Seven ichnofossil assemblages, i.e., Cruziana-Rusophycus, Planolites-Palaeophycus, Cruziana problematica, Diplichnites, Cochlichnus anguineus, Bergaueria perata and Psammichnites gigas have been recognized in the Lower Quartzite to Arkosic Sandstone members of the Koti Dhaman Formation. Seven sedimentary facies i.e., sandstone–shale facies (FT1), cross-bedded (trough and planar) sandstone (FT2), bedded sandstone facies (FT3), shale facies (FT4), shale–sandstone facies (FT5), shale-rippled sandstone facies (FT6) and planar and trough cross-laminated sandstone (FT7) and four facies associations FA1-FA4 are identified in the Koti Dhaman Formation. The formation contains shallowing upward parasequences of a tidal flat complex. Overall, two major events are recognized: i) the break in sedimentation between the Lower Quartzite Member and the overlying Shale Member probably related to forced-regressive event and ii) the facies shift from FT6 to FT7 of the Arkosic Sandstone Member represents an erosive transgressive event; the surface is interpreted as wave ravinement surface, which also serves as a sequence boundary. Integrated ichnology, sedimentology and sequence stratigraphic studies indicate that the Lower Quartzite Member was deposited in a shallow subtidal sand sheet complex and tidal flat complex; the Shale Member was deposited in a mud flat setting of a tidal flat complex, and the Arkosic Sandstone Member in a mixed-flat (tidal flat complex) to sand sheet complex front and margin (subtidal sand sheet complex). Overall, the lower to middle part of the Koti Dhaman Formation represents a tide-dominated shallow subtidal–intertidal to mud-flat subenvironments of the tidal flat complex. A palaeogeographic reconstruction of lower Cambrian (516–514?Ma) is presented based on the distribution of trilobite traces from the Lesser Himalaya and the Bikaner–Nagaur area of Peninsular India (eastern Gondwana), Egypt, Jordan, Turkey (western Gondwana) and Canada (Avalonia).     

Memorial

Taxonomy and palaeoecology of flysch trace fossils: The marnoso‐arenacea formation and associated facies (Miocene, northern Apennines, Italy)

Taxonomy and palaeoecology of flysch trace fossils: The Marnoso‐arenacea Formation and associated facies (Miocene, Northern Apennines, Italy), Alfred Uchman, 1995. Beringeria, Würzburger geo‐wissenschaftliche Mitteilungen, 15, 115 p., 16 pl., 58 DM [Orders and other information available from Fruende der Würtzburg Geo‐wissenschaften e. V., Pleicherwall 1, 97070 Würtzburg, Germany].

Track attack

Dinossaurios e a batalha de Cantique by Antonio M. Galopim de Carvalho, 1994. Editorial Noticias, Lisboa, 291 p.

Annotated Atlas of South America tetrapod footprints (Devonian to Holocene) by Giuseppe Leonardi, 1994. Publication of the Com‐panhia de Pesquisa de Recursos Minerais, Brasilia, 248 p., 35 pls.     

Phylogenetic relationships of tropical western Atlantic snappers in subfamily Lutjaninae (Lutjanidae: Perciformes) inferred from mitochondrial DNA sequences

Phylogenetic relationships among 20 nominal species of tropical lutjanine snappers (Lutjanidae) (12 from the western Atlantic, one from the eastern Pacific, and seven from the Indo‐Pacific) were inferred based on 2206 bp (712 variable, 614 parsimony informative) from three protein‐coding mitochondrial genes. Also included in the analysis were DNA sequences from two individuals, identified initially as Lutjanus apodus, which were sampled off the coast of Bahia State in Brazil (western Atlantic), and from three individuals labelled as ‘red snapper’ in the fish market in Puerto Armuelles, Panama (eastern Pacific). Bayesian posterior probabilities and maximum‐likelihood bootstrap percentages strongly supported monophyly of all lutjanines sampled and the hypothesis that western Atlantic lutjanines are derived from an Indo‐Pacific lutjanine lineage. The phylogenetic hypothesis also indicated that oceans where lutjanines are distributed (western Atlantic, eastern Pacific, and Indo‐Pacific) are not reciprocally monophyletic for the species distributed within them. There were three strongly supported clades that included all western Atlantic lutjanines: one included six species of Lutjanus from the western Atlantic, two species of Lutjanus from the eastern Pacific, and the monotypic genera Rhomboplites and Ocyurus (western Atlantic); one that included three, probably four, species of Lutjanus in the western Atlantic; and one that included Lutjanus cyanopterus (western Atlantic), an unknown species of Lutjanus from the eastern Pacific, and three species of Lutjanus from the Indo‐Pacific. Molecular‐clock calibrations supported an early Miocene diversification of an Indo‐Pacific lutjanine lineage that dispersed into the western Atlantic via the Panamanian Gateway. Divergent evolution among these lutjanines appears to have occurred both by vicariant and ecological speciation: the former following significant geographic or geological events, including both shoaling and closure of the Panamanian Gateway and tectonic upheavals, whereas the latter occurred via phenotypic diversification inferred to indicate adaptation to life in different habitats. Taxonomic revision of western Atlantic lutjanines appears warranted in that monotypic Ocyurus and Rhomboplites should be subsumed within the genus Lutjanus. Finally, it appears that retail mislabelling of ‘red snapper’ in commercial markets extends beyond the USA. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 102 , 915–929.     

Positive range–abundance relationships in Indo‐Pacific bird communities

Reeve et al. (2016, Ecography, 39 , 990–997) recently reported negative range–abundance relationships in Indo‐Pacific bird communities and speculated that geographical isolation facilitates the evolution of broad‐niched, small‐ranged and abundant species. We tested this relationship using a large independent data set on range and abundance of birds across New Caledonia (over 4,000 bird census points for 17,300 km²). In contradiction to Reeve et al. (2016, Ecography, 39 , 990–997), we found clear evidence that range–abundance relationships are positive and endemic species have narrower habitat niches than wide‐range species. Our findings are likely valid also for other islands in the Indo‐Pacific.     

Facies,paleoenvironment, carbonate platform and facies changes across Paleocene Eocene of the Taleh Zang Formation in the Zagros Basin,SW-Iran

The Paleocene–Eocene Taleh Zang Formation of the Zagros Basin is a sequence of shallow-water carbonates. We have studied carbonate platform, sedimentary environments and its changes based on the facies analysis with particular emphasis on the biogenic assemblages of the Late Paleocene Sarkan and Early Eocene Maleh kuh sections. In the Late Paleocene, nine microfacies types were distinguished, dominated by algal taxa and corals at the lower part and larger foraminifera at the upper part. The Lower Eocene section is characterised by 10 microfacies types, which are dominated by diverse larger foraminifera such as alveolinids, orbitolitids and nummulitids. The Taleh Zang Formation at the Sarkan and Maleh kuh sections represents sedimentation on a carbonate ramp.

The deepening trends show a gradual increase in perforate foraminifera, the deepest environment is marked by the maximum occurrence of perforate foraminifers (Nummulites), while the shallowing trends are composed mainly of imperforate foraminifera and also characterised by lack of fossils in tidal flat facies.

Based on the facies changes and platform evolution, three stages are assumed in platform development: I; algal and coralgal colonies (coralgal platform), II; coralgal reefs giving way to larger foraminifera, III; dominance of diverse and newly developing larger foraminifera lineages in oligotrophic conditions.     

Thanetian–Ilerdian coralline algae and benthic foraminifera from northeast India: microfacies analysis and new insights into the Tethyan perspective

Thanetian–Ilerdian carbonate deposits from the Lakadong Limestone in Assam Shelf, Meghalaya, northeast India, are studied with respect to microfacies distributions and controlling ecological factors on dominant biogenic components. Palaeoenvironmental implications are inferred following the detailed analysis of microfacies characterized by rich assemblages of coralline red algae and benthic foraminifera. The carbonate sediments have been interpreted as lagoonal to outer shelf facies. It is envisaged that the analysed benthic communities thrived in a meso‐oligotrophic regime above the fair‐weather wave base. The Lakadong Limestone constitutes a well‐preserved record of Late Palaeocene–earliest Eocene shallow marine carbonate ecosystem and has high potential to decipher its response to an interval of distinct changes in climate and tectonic settings. The abundance of oligotrophic larger benthic foraminifera in the Lakadong Limestone is comparable to the foraminiferal assemblages of west Tethys. The phylogenetic changes (‘Larger Foraminiferal Turnover’, LFT) and subsequent rapid radiation of typical Eocene larger benthic foraminifera (Alveolina, Nummulites) usually observed in the west Tethys have also been observed in the upper part of the Lakadong Limestone. The eastward migration of Eocene foraminifera from the west coincided with the India‐Asia collision and global warming events at the Palaeocene–Eocene boundary that may have produced a wide array of modifications in biogeography, seasonal run‐offs and ocean circulation pathways. The data indicate that rapid rate of migration from west before the onset of geographic barriers and/or timely restoration of pan‐Tethyan environmental conditions ensured the incidence of these forms in the earliest Eocene sediments.     

Hydraulic sorting of microbiota in calciturbidites— A dinantian case study from the rheinische schiefergebirge,Germany

Summary A calciturbidite bed from the lower part of the Kieselkalk Formation (late cd II) at Wallau, eastern Rheinisches Schiefergebirge, displays ideal grading of reworked calcareous shallow-water microbiota, ranging from plurimillimetric agglutinated foraminifers and fragments of calcareous algae (Koninckopora sp.) to plurimicronic calcispheres, radiolarians and sponge spicules. Microbiota derived from all levels of the platform. Correspondingly, several carbonate microfacies types could be discerned. The early diagenetic micrite base of the turbidite preserved the anoxic basinal facies. The turbidite bed belongs to Foraminiferal Zone 15 (V3bα), theAlbaillella cartalla Zone (radiolarian chronology), and is the Lower and part of the Uppertexanus Zone (standard conodont zonation). From the few published data on foraminifers, the Kieselkalk is thought to range from Mid Viséan V2b to Late Viséan V3b gamma.     

Isolation and characterization of novel microsatellite markers for Cymodocea serrulata (Cymodoceaceae), a seagrass distributed widely in the Indo‐Pacific region

Cymodocea serrulata is a tropical seagrass species distributed widely in the Indo‐Pacific region. We developed 16 novel microsatellite (simple sequence repeat) markers for C. serrulata using next‐generation sequencing for use in genetic studies. The applicability of these markers was attested by genotyping of 40 individuals collected from a natural population in the Philippines. Of the 16 loci, 15 showed polymorphism. For the 15 polymorphic markers, the number of alleles per locus ranged from two to seven, and the observed and expected heterozygosities ranged from 0.131–1.000 and 0.124–0.788, respectively. These markers are useful tools for elucidating genetic diversity, connectivity, and structure in this foundational coastal species.     

Latest Oligocene through early middle Miocene diatom biostratigraphy of the eastern tropical Pacific

Study of DSDP Sites 71, 77, and 495 has allowed the development of a refined diatom biostratigraphy for the latest Oligocene through early middle Miocene of the eastern tropical Pacific which is well correlated to the low-latitude zonations for planktonic foraminifers, coccoliths, and radiolarians. Six zones and 7 subzones are proposed, and correlation with high-latitude diatoms zonations for the North Pacific, the Norwegian Sea, and the Southern Ocean is suggested by the discovery of selected diatoms in these tropical sediments which were previously thought to be restricted to high latitudes. Six new species and one new variety of diatoms which are stratigraphically useful are proposed: Actinocyclus hajosiae, n. sp., A. radionovae, n. sp., Coscinodiscus blysmos, n. sp., C. praenodulifer, n. sp., Craspedodiscus rydei, n. sp., Thalassiosira bukryi, n. sp., and Coscinodiscus lewisianus var. robustus n. var.     

Oceanographic significance of Pacific late miocene calcareous nannoplankton

An analysis of the variability in the composition and distribution of Pacific Late Miocene calcareous nannoplankton about their average biogeography shows that there are primarily two environmental factors causing that variability, climate and dissolution. Climate produces a latitudinal, biogeographic differentiation of the Late Miocene nannoflora, while selective dissolution superimposes a bathymetric differentiation of the nannoflora on that due to climate. Together, these two factors produce three distinct Late Miocene nannofloral assemblages, a high-latitude, temperate assemblage characterized by Reticulofenestra pseudoumbilica and Coccolithus pelagicus, and two tropical assemblages, their differences in composition depending on water depth and surface-water productivity: (1) in shallower water and beneath areas of higher organic production and sedimentation of calcite there is an undissolved assemblage characterized by sphenoliths, small elliptical placoliths and Coccolithus pataecus; (2) in deeper water and areas of lower productivity there is a dissolved assemblage dominated by discoasters.Selective dissolution produces most of the apparent biogeographic variation in Pacific Late Miocene nannoplankton compositions, the variation in compositions observed between the seventeen sites studied. Dissolution preferentially removes the more soluble constituents of the tropical nannoflora so that increasing dissolution tends to give tropical nannoflora a cooler, more temperate aspect. At the same time, selective dissolution shifts the composition of the warmer, tropical component towards its more resistant taxa.Nannoplankton records show a period of greatly decreased calcite dissolution in deep tropical and temperate South Pacific sites between about 8 and 10 m.y. ago. This decrease is strongly correlated with a temporary increase in the ¹³C composition of Pacific deep waters. Calcite dissolution increased during this same period in the deep North Pacific.Nannoplankton records of Late Miocene climate in the tropics are distinctly different from those at higher, south temperate latitudes. Tropical records show a sharp warming in the earliest Late Miocene after a generally cool late Middle Miocene. This was followed by a temporary cooling, nearly to Middle Miocene levels, about 7 m.y. ago. Toward the end of the Late Miocene, the tropical Pacific warmed again and remained warm into the Pliocene. Warming of temperate climates occurred much later. Not until latest Miocene did the southern the Pliocene. Warming of temperate climates occurred much later. Not until latest Miocene did the southern temperate latitudes warm appreciably. Southern subpolar climate cooled continuously through the Late Miocene. We attribute the resulting increases in the latitudinal climatic contrast across the southern Pacific Ocean to the development and migration of a strong subtropical convergence.On the basis of the nannoplankton oceanographic records we postulate that beginning about 10.5 m.y. ago Pacific surface circulation became primarily zonal and the production of deep and bottom waters in the Southern Ocean increased sharply. This produced a northward decrease in calcite preservation, an increase in benthic ¹³C, and a strong climatic gradient across southern latitudes. The period of most vigorous deep Pacific circulation ended 7 m.y. ago in response, we speculate, to the reduced ocean salinities during the Messinian.     

Longtail tuna Thunnus tonggol (Bleeker, 1851) shows genetic partitioning across,but not within,basins of the Indo‐Pacific based on mitochondrial DNA

Genetic stock structure is atypical in tuna species, with most species demonstrating geographically‐broad, panmictic populations. Here, genetic data suggest a distinct pattern for Thunnus tonggol across the Indo‐Pacific region. The genetic variation in the coastal tuna T. tonggol sampled from across the South China Sea was examined using the highly variable mitochondrial DNA displacement loop (D‐loop) gene region. One hundred and thirty‐nine specimens were sampled from four locations in Indonesia, Vietnam and the Philippines. Phylogenetic reconstruction of genetic relationships revealed no significant ?_ST statistics and hence no population structure within the South China Sea. However, subsequent analysis with sequence data from coastal northwest India infers discrete genetic stocks between the Indian Ocean and the South China Sea. Consistent with previous genetic analyses of tuna species in the Indo‐Pacific, the findings in this study infer no population structure within each basin, but rather show a significant partitioning across the wider region. Furthermore, these results have implications for the management of the commercially valuable Thunnus tonggol across national boundaries, and thus requiring collaboration among countries to ensure its sustainable use.     

Phylogeography of the humbug damselfish,Dascyllus aruanus (Linnaeus, 1758): evidence of Indo‐Pacific vicariance and genetic differentiation of peripheral populations

The phylogeographical structure of coral‐associated reef fishes may have been severely affected, more than species from deeper habitats, by habitat loss during periods of low sea level. The humbug damselfish, Dascyllus aruanus, is widely distributed across the Indo‐West Pacific, and exclusively inhabits branching corals. We used mitochondrial cytochrome b sequence and seven microsatellite loci on D. aruanus samples (260 individuals) from 13 locations across the Indo‐West Pacific to investigate its phylogeographical structure distribution‐wide. A major genetic partition was found between the Indian and Pacific Ocean populations, which we interpret as the result of geographical isolation on either side of the Indo‐Pacific barrier during glacial periods. The peripheral populations of the Red Sea and the Society Islands exhibited lower genetic diversity, and substantial genetic differences with the other populations, suggesting relative isolation. Thus, vicariance on either side of the Indo‐Pacific barrier and peripheral differentiation are considered to be the main drivers that have shaped the phylogeographical patterns presently observed in D. aruanus. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 113 , 931–942.     

The systematics of Lobophora (Dictyotales,Phaeophyceae) in the western Atlantic and eastern Pacific oceans: eight new species

Lobophora is a common tropical to temperate genus of brown algae found in a plethora of habitats including shallow and deep‐water coral reefs, rocky shores, mangroves, seagrass beds, and rhodoliths beds. Recent molecular studies have revealed that Lobophora species diversity has been severely underestimated. Current estimates of the species numbers range from 100 to 140 species with a suggested center of diversity in the Central Indo‐Pacific. This study used three molecular markers (cox3, rbcL, psbA), different single‐marker species delimitation methods (GMYC, ABGD, PTP), and morphological evidence to evaluate Lobophora species diversity in the Western Atlantic and the Eastern Pacific oceans. Cox3 provided the greatest number of primary species hypotheses(PSH), followed by rbcL and then psbA. GMYC species delimitation analysis was the most conservative across all three markers, followed by PTP, and then ABGD. The most informative diagnostic morphological characters were thallus thickness and number of cell layers in both the medulla and the dorsal/ventral cortices. Following a consensus approach, 14 distinct Lobophora species were identified in the Western Atlantic and five in the Eastern Pacific. Eight new species from these two oceans were herein described: L. adpressa sp. nov., L. cocoensis sp. nov., L. colombiana sp. nov., L. crispata sp. nov., L. delicata sp. nov., L. dispersa sp. nov., L. panamensis sp. nov., and L. tortugensis sp. nov. This study showed that the best approach to confidently identify Lobophora species is to analyze DNA sequences (preferably cox3 and rbcL) followed by comparative morphological and geographical assessment.     

Molecular data illuminate cryptic nudibranch species: the evolution of the Scyllaeidae (Nudibranchia: Dendronotina) with a revision of Notobryon

Scyllaeidae represents a small clade of dendronotoid nudibranchs. Notobryon wardi Odhner, 1936, has been reported to occur in tropical oceans from the Indo‐Pacific and eastern Pacific to temperate South Africa. The systematics of Notobryon has not been reviewed using modern systematic tools. Here, specimens of Notobryon were examined from the eastern Pacific, the Indo‐Pacific, and from temperate South Africa. Additionally, representatives of Scyllaea and Crosslandia were studied. Scyllaeidae was found to be monophyletic. Notobryon was also found to be monophyletic and is the sister group to Crosslandia plus Scyllaea. The molecular data also clearly indicate that within Notobryon, at least three distinct species are present, two of which are here described. Genetic distance data indicate that eastern Pacific and South African exemplars are 10–23% divergent from Indo‐Pacific exemplars of Notobryon wardi. Scyllaea pelagica has been regarded as a single, circumtropical species. Our molecular studies clearly indicate that the Atlantic and Indo‐Pacific populations are distinct and we resurrect Scyllaea fulva Quoy & Gaimard, 1824 for the Indo‐Pacific species. Our morphological studies clearly corroborate our molecular findings and differences in morphology distinguish closely related species. Different species clearly have distinct penial morphology. These studies clearly reinforce the view that eastern Pacific, Indo‐Pacific, and temperate biotas consist largely of distinct faunas, with only a minor degree of faunal overlap. © 2012 The Linnean Society of London, Zoological Journal of the Linnean Society, 2012, 165 , 311–336.     

SYMBIODINIUM (DINOPHYTA) DIVERSITY AND STABILITY IN AQUARIUM CORALS1

Indo‐Pacific reef corals growing for years in closed‐system aquaria provide an alternate means to investigate host–symbiont specificity and stability. The diversity of dinoflagellate endosymbionts (Symbiodinium spp.) from coral communities in private and public aquaria was investigated using molecular‐genetic analyses. Of the 29 symbiont types (i.e., species) identified, 90% belonged to the most prevalent group of Symbiodinium harbored by Indo‐Pacific reef corals, Clade C, while the rest belonged to Clade D. Sixty‐five percent of all types were known from field surveys conducted throughout the Pacific and Indian oceans. Because specific coral–dinoflagellate partnerships appear to have defined geographic distributions, correspondence of the same symbionts in aquarium and field‐collected specimens identifies regions where particular colonies must have been collected in the wild. Symbiodinium spp. in clade D, believed to be “stress‐tolerant” and/or “opportunistic,” occurred in a limited number of individual colonies. The absence of a prevalent, or “weedy,” symbiont suggests that conditions under which aquarium corals are grown do not favor competitive replacements of their native symbiont populations. The finding of typical and diverse assemblages of Symbiodinium spp. among aquarium corals living many years under variable chemical/physical conditions, artificial and natural light, while undergoing fragmentation periodically, indicates that individual colonies maintain stable, long‐term symbiotic associations.