Paleogene and Late Cretaceous Pteropoda (Mollusca, Gastropoda, Heterobranchia) from North America.

A comprehensive discussion and survey is made of all North American Paleogene and Late Cretaceous pteropods, and their systematics reviewed. From the West Coast of North America pteropod fossils have been collected from 23 localities in Washington State, and from the Gulf Coastal Plain they have been found in 40 localities. We also review earlier published specimens from boreholes in the Atlantic Coastal Plain. As a result, six new species are introduced from the Gulf Coast (Currylimacina asperita Garvie sp. nov., Heliconoides hodgkinsoni Garvie sp. nov., Limacina texanopsis Garvie sp. nov., Limacina parvabrazensis Garvie Janssen sp. nov., Limacina pseudopygmaea Garvie Janssen sp. nov., Cheilospicata cedrus Garvie sp. nov.) and one from Washington State (Clio gailae Goedert Janssen sp. nov.). The geographical distribution and/or stratigraphic ranges of several species has been extended, for example, Limacina canadaensis Hodgkinson, 1992, is reported for the first time from western North America. Some species originally described from elsewhere, Altaspiratella multispira (Curry, 1982), Heliconoides bartonensis (Curry, 1965), L. karasawai Ando, 2011, L. aff. valvatina (Reuss, 1867), Creseis spina (Reuss, 1867) and Clio chadumica Korobkov, 1966, are reported for the first time from the Americas. New specimens of several poorly known species yielded additional information on morphology or biostratigraphy. It is shown that some species have longer temporal ranges in America, occurring in, or continuing into younger deposits than those in Europe. In addition to the only Cretaceous pteropod from Washington State known so far, several enigmatic species possibly representing Pteropoda are described from the Late Cretaceous (Maastrichtian) and Danian of Texas, which allows a salient discussion on the early history of Pteropoda. An enigmatic specimen from the Eocene of Texas might represent a very early form of Gymnosomata. This analysis shows that there may be more pteropod species represented during the Late Cretaceous and Paleocene than previously suspected.

The origin and diversification of pteropods precede past perturbations in the Earth's carbon cycle.

Pteropods are a group of planktonic gastropods that are widely regarded as biological indicators for assessing the impacts of ocean acidification. Their aragonitic shells are highly sensitive to acute changes in ocean chemistry. However, to gain insight into their potential to adapt to current climate change, we need to accurately reconstruct their evolutionary history and assess their responses to past changes in the Earth's carbon cycle. Here, we resolve the phylogeny and timing of pteropod evolution with a phylogenomic dataset (2,654 genes) incorporating new data for 21 pteropod species and revised fossil evidence. In agreement with traditional taxonomy, we recovered molecular support for a division between "sea butterflies" (Thecosomata; mucus-web feeders) and "sea angels" (Gymnosomata; active predators). Molecular dating demonstrated that these two lineages diverged in the early Cretaceous, and that all main pteropod clades, including shelled, partially-shelled, and unshelled groups, diverged in the mid- to late Cretaceous. Hence, these clades originated prior to and subsequently survived major global change events, including the Paleocene-Eocene Thermal Maximum (PETM), the closest analog to modern-day ocean acidification and warming. Our findings indicate that planktonic aragonitic calcifiers have shown resilience to perturbations in the Earth's carbon cycle over evolutionary timescales.

Fossil-calibrated molecular phylogeny of atlantid heteropods (Gastropoda, Pterotracheoidea).

BACKGROUND: The aragonite shelled, planktonic gastropod family Atlantidae (shelled heteropods) is likely to be one of the first groups to be impacted by imminent ocean changes, including ocean warming and ocean acidification. With a fossil record spanning at least 100 Ma, atlantids have experienced and survived global-scale ocean changes and extinction events in the past. However, the diversification patterns and tempo of evolution in this family are largely unknown. RESULTS: Based on a concatenated maximum likelihood phylogeny of three genes (cytochrome c oxidase subunit 1 mitochondrial DNA, 28S and 18S ribosomal rRNA) we show that the three extant genera of the family Atlantidae, Atlanta, Protatlanta and Oxygyrus, form monophyletic groups. The genus Atlanta is split into two groups, one exhibiting smaller, well ornamented shells, and the other having larger, less ornamented shells. The fossil record, in combination with a fossil-calibrated phylogeny, suggests that large scale atlantid extinction was accompanied by considerable and rapid diversification over the last 25 Ma, potentially driven by vicariance events. CONCLUSIONS: Now confronted with a rapidly changing modern ocean, the ability of atlantids to survive past global change crises gives some optimism that they may be able to persist through the Anthropocene.

The status of Cuvierina spoeli Rampal, 2002 and Cuvieria oryza Benson, 1835 (Gastropoda, Pteropoda, Cuvierinidae).

The name Cuvierina spoeli Rampal, 2002 cannot be understood as a new replacement name (nom. nov.) for Cuvierina atlantica, as published in the unavailable name Cuvierina columnella (Rang, 1827) f. atlantica van der Spoel, 1970. The name atlantica was made available as a name of the species group in the combination Cuvierina columnella atlantica Bé, MacClintock Currie, 1972, currently interpreted to represent the species Cuvierina atlantica. Cuvierina spoeli type material does not belong to C. atlantica, but to C. columnella of which it therefore is a junior subjective synonym.        The name Cuvieria oryza Benson, 1835 was recognised to represent a senior synonym of Cuvierina urceolaris (Mörch, 1850), not a junior synonym or form of C. columnella Rang, 1827, as it was frequently interpreted by authors. The name C. oryza is here declared nomen oblitum and the name C. urceolaris, because in prevailing usage, is declared nomen protectum. The conditions of ICZN (1999, art. 23.9.1.1 and .2) are met.

Biogeography and genetic diversity of the atlantid heteropods.

The atlantid heteropods are regularly encountered, but rarely studied marine planktonic gastropods. Relying on a small (<14 mm), delicate aragonite shell and living in the upper ocean means that, in common with pteropods, atlantids are likely to be affected by imminent ocean changes. Variable shell morphology and widespread distributions indicate that the family is more diverse than the 23 currently known species. Uncovering this diversity is fundamental to determining the distribution of atlantids and to understanding their environmental tolerances. Here we present phylogenetic analyses of all described species of the family Atlantidae using 437 new and 52 previously published cytochrome c oxidase subunit 1 mitochondrial DNA (mtCO1) sequences. Specimens and published sequences were gathered from 32 Atlantic Ocean stations, 14 Indian Ocean stations and 21 Pacific Ocean stations between 35°N and 43°S. DNA barcoding and Automatic Barcode Gap Discovery (ABGD) proved to be valuable tools for the identification of described atlantid species, and also revealed ten additional distinct clades, suggesting that the diversity within this family has been underestimated. Only two of these clades displayed obvious morphological characteristics, demonstrating that much of the newly discovered diversity is hidden from morphology-based identification techniques. Investigation of six large atlantid collections demonstrated that 61% of previously described (morpho) species have a circumglobal distribution. Of the remaining 39%, two species were restricted to the Atlantic Ocean, five occurred in the Indian and Pacific oceans, one species was only found in the northeast Pacific Ocean, and one occurred only in the Southern Subtropical Convergence Zone. Molecular analysis showed that seven of the species with wide distributions were comprised of two or more clades that occupied distinct oceanographic regions. These distributions may suggest narrower environmental tolerances than the described morphospecies. Results provide an updated biogeography and mtCO1 reference dataset of the Atlantidae that may be used to identify atlantid species and provide a first step in understanding their evolutionary history and accurate distribution, encouraging the inclusion of this family in future plankton research.

Time-calibrated molecular phylogeny of pteropods.

Pteropods are a widespread group of holoplanktonic gastropod molluscs and are uniquely suitable for study of long-term evolutionary processes in the open ocean because they are the only living metazoan plankton with a good fossil record. Pteropods have been proposed as bioindicators to monitor the impacts of ocean acidification and in consequence have attracted considerable research interest, however, a robust evolutionary framework for the group is still lacking. Here we reconstruct their phylogenetic relationships and examine the evolutionary history of pteropods based on combined analyses of Cytochrome Oxidase I, 28S, and 18S ribosomal rRNA sequences and a molecular clock calibrated using fossils and the estimated timing of the formation of the Isthmus of Panama. Euthecosomes with uncoiled shells were monophyletic with Creseis as the earliest diverging lineage, estimated at 41-38 million years ago (mya). The coiled euthecosomes (Limacina, Heliconoides, Thielea) were not monophyletic contrary to the accepted morphology-based taxonomy; however, due to their high rate heterogeneity no firm conclusions can be drawn. We found strong support for monophyly of most euthecosome genera, but Clio appeared as a polyphyletic group, and Diacavolinia grouped within Cavolinia, making the latter genus paraphyletic. The highest evolutionary rates were observed in Heliconoides inflatus and Limacina bulimoides for both 28S and 18S partitions. Using a fossil-calibrated phylogeny that sets the first occurrence of coiled euthecosomes at 79-66 mya, we estimate that uncoiled euthecosomes evolved 51-42 mya and that most extant uncoiled genera originated 40-15 mya. These findings are congruent with a molecular clock analysis using the Isthmus of Panama formation as an independent calibration. Although not all phylogenetic relationships could be resolved based on three molecular markers, this study provides a useful resource to study pteropod diversity and provides general insight into the processes that generate and maintain their diversity in the open ocean.

Revision of the genus Cuvierina Boas, 1886 based on integrative taxonomic data, including the description of a new species from the Pacific Ocean (Gastropoda, Thecosomata).

Shelled pteropods (Gastropoda, Thecosomata, Euthecosomata) are a group of holoplanktonic gastropods that occur predominantly in the surface layers of the world's oceans. Accurate species identifications are essential for tracking changes in species assemblages of planktonic gastropods, because different species are expected to have different sensitivities to ocean changes. The genus Cuvierina has a worldwide warm water distribution pattern between ~36°N and ~39°S. Based on an integrative taxonomic approach combining morphometric, genetic, and biogeographic information, the two subgenera of Cuvierina, Cuvierinas. str. and Urceolarica, are rejected. A new species is introduced: Cuvierina tsudaisp. n., which has to date been considered the same species as Cuvierina pacifica. Cuvierina tsudaisp. n. is endemic to the Pacific Ocean and is characterised by a shell height of 7.2-8.0 mm, a moderately cylindrical shell shape, the absence of micro-ornamentation and a triangular aperture. Cuvierina pacifica is restricted to the centre of the oligotrophic southern Pacific gyre, has a shell height of 6.6-8.5 mm, a more cylindrical shell shape, no micro-ornamentation and a less triangular aperture than Cuvierina tsudaisp. n.