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Abstract

The superfamily Stromboidea is a clade of morphologically distinctive gastropods which include the iconic Strombidae, or ‘true conchs’. In this study, we present the most taxonomically extensive phylogeny of the superfamily to date, using fossil calibrations to produce a chronogram and extant geographical distributions to reconstruct ancestral ranges. From these results, we confirm the monophyly of all stromboidean families; however, six genera are not monophyletic using current generic assignments (Strombidae: Lentigo, Canarium , Dolomena , Doxander ; Xenophoridae: Onustus, Xenophora ). Within Strombidae, analyses resolve an Indo-West Pacific (IWP) clade sister to an East Pacific/Atlantic clade, together sister to a second, larger IWP clade. Our results also indicate two pulses of strombid diversification within the Miocene, and a Tethyan/IWP origin for Strombidae – both supported by the fossil record. However, conflicts between divergence time estimates and the fossil record warrant further exploration. Species delimitation analyses using the COI barcoding gene support several taxonomic changes. We synonymise Euprotomus aurora with Euprotomus bulla , Strombus alatus with Strombus pugilis , Dolomena abbotti with Dolomena labiosa , and Dolomena operosa with Dolomena vittata . We identified cryptic species complexes within Terebellum terebellum , Lambis lambis , “Canarium” wilsonorum, Dolomena turturella and Maculastrombus mutabilis . We reinstate Rimellopsis laurenti as a species (previously synonymised with R. laurenti ) and recognise Harpago chiragra rugosus and Lambis truncata sowerbyi valid at the rank of species. Finally, we establish several new combinations, rendering Lentigo , Dolomena , and Canarium monophyletic: Lentigo thersites , Dolomena robusta , Dolomena epidromis , Dolomena turturella , Dolomena taeniata, Dolomena vanikorensis , D. vittata , “Canarium” wilsonorum , Hawaiistrombus scalariformis , Maculastrombus mutabilis , Maculastrombus microurceus .

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We present the first systematic description of a Tortonian (late Miocene) gastropod assemblage from the Ambug Hill section in the Tutong District in Brunei Darussalam. The low-diversity assemblage comprises 62 species of which 37 are unknown from other Neogene faunas of the Indo-West Pacific Region (IWP), 23 species are formally described as new. Carnivorous, scavenging and detritus feeding species predominate and the ecological requirements of extant relatives suggest a mud-bottom environment of the inner shelf in several tens of meters water depth with reduced vegetation. The small size of most specimens, the frequent occurrence of subadult shells and small size of several species relative to congeneric species might point to suboptimal environmental conditions during the deposition of this part of the Seria Formation. The very low relations with Neogene gastropod faunas from Indonesia at species level might be explained by the biogeographic isolation between the faunas of the Java and Celebes seas and that from the South China Sea. A severe undersampling and rather spotty taxonomic descriptions of Neogene IWP-faunas and a lack of assemblages from identical depositional environments from Indonesia may also contribute to the seemingly endemic character of the Ambug Hill fauna. Rhinoclavis pulcherrima n. sp., Rissoina tutongensis n. sp., Ficus parvissima n. sp., Dolomena bruneiensis n. sp., Scalptia verheckeni n. sp., Phos bruneiensis n. sp., Nassarius pseudoovum n. sp., Indomitrella acuticonica n. sp., Hemifusus charlieleei n. sp., Prunum seriaense n. sp., Cymbiola ambugensis n. sp., Ziba waltercernohorskyi n. sp., Amalda bruneiana n. sp., Tomopleura furcata n. sp., Unedogemmula nuttallin. sp., Gemmula sculpturata n. sp., Crassispira strangulata n. sp., Paradrillia pachyspira n. sp., Conasprella paupera n. sp., Conasprella trianginodus n. sp., Triplostephanus wilfordi n. sp., Duplicaria aequalis n. sp. and Architectonica beetsi n. sp. are established as new species.
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The TethysOcean existed between the continents of Gondwana and Laurasia from the Triassic to the Pliocene. Analyses of multiple biogeographic and phylogenetic histories reveal that the subsequent breakup of the Tethys greatly influenced the distributions of many species. The ancestral Tethyan realm broke into five biogeographic provinces, including the present-day East Pacific, West Atlantic, East Atlantic, Mediterranean Sea, and Indo-West Pacific. Palaeogeographic maps illustrate the Mesozoic Atlantic opening, the Cenozoic closure of the Tethys, the Messinian Salinity Crisis, the mid-Miocene closure of the Central American Seaway, and Quaternary geological changes. Further, we consider Cenozoic sea-level changes and the formation of freshwater habitats. These reconstructions allow assessment of patterns of aquatic diversification for marine and freshwater animals, and comparison of vicariance and dispersal processes. Estimated divergence times indicate that fragmentation of the Tethys was responsible for the vicariant speciation of aquatic animals because these dates are consistent with associated tectonic events. The opening of the Atlantic Ocean during the Cretaceous is responsible for the earliest isolation between the West and East Atlantic. The mid-Miocene closure of the Tethys, which blocked global equatorial currents, appears to have isolated the Atlantic/Mediterranean Sea and Indo-West Pacific. Finally, formation of the Isthmus of Panama isolated East Pacific and West Atlantic marine organisms. Dispersals related to the Messinian Salinity Crisis and Quaternary sea-level changes influenced population structuring. Tethyan changes affected marine habitats, created new freshwater habitats, inland caves and ancient lakes along the Alps and Himalayas, and influenced anchialine caves at the edge of the ancient sea. The extensive new habitats provided opportunities for colonisation and rapid diversification. Future work should focus on testing the biological impact of the series of Tethyan changes.
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Phylogenies that are reconstructed without fossil material often contain approximate dates for lineage splitting. For example, particular nodes on molecular phylogenies may be dated by known geographic events that caused lineages to split, thereby calibrating a molecular clock that is used to date other nodes. On the one hand, such phylogenies contain no information about lineages that have become extinct. On the other hand, they do provide a potentially useful testing ground for ideas about evolutionary processes. Here we first ask what such reconstructed phylogenies should be expected to look like under a birth-death process in which the birth and death parameters of lineages remain constant through time. We show that it is possible to estimate both the birth and death rates of lineages from the reconstructed phylogenies, even though they contain no explicit information about extinct lineages. We also show how such phylogenies can reveal mass extinctions and how their characteristic footprint can be distinguished from similar ones produced by density-dependent cladogenesis.
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In this study, we first determined the complete mitochondrial genome of the endangered marine strombid gastropod Strombus luhuanus, an important shellfish traditionally distributed around Hainan island of China and parts of Papua New Guinea. The complete mitogenome sequence was 15,799 bp in length containing 13 protein-coding genes, two rRNA genes, 22 tRNA genes, and one non-coding control region (D-loop). We further compared the mtDNA sequences of S. luhuanus and S. gigas, and calculated the moderate sequence variations in their protein coding regions, which would further provide valuable genome variation information for conservative studies of S. luhuanus.