Isthmus Of Panama Research Articles

Integrative taxonomy unravels the species diversity of Parachristianella (Cestoda : Trypanorhyncha) from both sides of the Panamanian isthmus

The uplift of the Panamanian isthmus in the Pliocene caused the termination of gene flow among members of many ancestral populations of marine lineages leading the diversification of geminate species confined to opposite sides of Central America. This palaeogeographical event has been evoked to explain the diversification of few lineages of batoids with trans-isthmian distribution. As such, there is the expectation that this vicariance event also affected lineages of parasites associated with them. Our study suggests that this event can explain the diversification of Parachristianella Dollfus, 1946 (Trypanorhyncha:Eutetrarhynchidae) in the Caribbean Sea and tropical eastern Pacific Ocean. We provide molecular, morphological and biogeographical evidence to recognise five lineages within this genus inhabiting the coastal waters of Panama, including P. parva Campbell & Beveridge, 2007 and four new species: P. mendozai, sp. nov., P. kuchtai, sp. nov., P. campbelli, sp. nov. and P. soldanovae, sp. nov. These species can be diagnosed by unambiguous sets of molecular characters. The morphological cohesiveness of sister species, which most likely diverged from around 3million years ago through the uplift of the Panamanian isthmus, suggests that the traditional emphasis on the tentacular armature to circumscribe species within trypanorhynchs might underestimate the diversity of recently diverged lineages.

Parallel diversifications of Cremastosperma and Mosannona (Annonaceae), tropical rainforest trees tracking Neogene upheaval of South America.

Much of the immense present day biological diversity of Neotropical rainforests originated from the Miocene onwards, a period of geological and ecological upheaval in South America. We assess the impact of the Andean orogeny, drainage of Lake Pebas and closure of the Panama isthmus on two clades of tropical trees (Cremastosperma, ca 31 spp.; and Mosannona, ca 14 spp.; both Annonaceae). Phylogenetic inference revealed similar patterns of geographically restricted clades and molecular dating showed diversifications in the different areas occurred in parallel, with timing consistent with Andean vicariance and Central American geodispersal. Ecological niche modelling approaches show phylogenetically conserved niche differentiation, particularly within Cremastosperma. Niche similarity and recent common ancestry of Amazon and Guianan Mosannona species contrast with dissimilar niches and more distant ancestry of Amazon, Venezuelan and Guianan species of Cremastosperma, suggesting that this element of the similar patterns of disjunct distributions in the two genera is instead a biogeographic parallelism, with differing origins. The results provide further independent evidence for the importance of the Andean orogeny, the drainage of Lake Pebas, and the formation of links between South and Central America in the evolutionary history of Neotropical lowland rainforest trees.

Phylogenetics and geography of speciation in New World Halichoeres wrasses

The New World Halichoeres comprises about 30 small to medium sized wrasse species that are prominent members of reef communities throughout the tropical Western Atlantic and Eastern Pacific. We conducted a phylogenetic analysis of this group and related lineages using new and previously published sequence data. We estimated divergence times, evaluated the monophyly of this group, their relationship to other labrids, as well as the time-course and geography of speciation. These analyses show that all members of New World Halichoeres form a monophyletic group that includes Oxyjulis and Sagittalarva. New World Halichoeres is one of numerous labrid groups that appear to have radiated rapidly about 32 Ma and form a large polytomy within the julidine wrasses. We reconstruct the tropical Western Atlantic to be the ancestral area of New World Halichoeres, with four invasions of the Eastern Pacific and one reversal from East Pacific to Western Atlantic. These five speciation events were spread across the history of the group, with none corresponding closely to the time of the closure of the Isthmus of Panama. Three speciation events within the Atlantic occurred across the Orinoco-Amazon outflow and within the Pacific, five involve splits between lineages that occupy coastal reef systems and offshore islands. Of eight sister species pairs, seven show complete allopatry and one is fully sympatric.

Ecomorphological trajectories of reef fish sister species (Pomacentridae) from both sides of the Isthmus of Panama

The vicariance model of biogeography focuses on allopatric speciation through fragmentation from an ancestral biota via a barrier that interrupts gene flow between populations. The evolutionary processes that occur over time in sister species on each side of the vicariance event influence their traits by a compromise between divergence and conservatism. The eastern Pacific and the Caribbean Sea were separated by the Isthmus of Panama ~ 3–6 Mya and allopatric speciation occurred on either side of the isthmus. Differences in ecological conditions on each side of the Isthmus of Panama separating the sister species may have shaped their niches and morphologies over evolutionary time. The objectives of this study were to: (1) analyze the variation in niche, morphology, and size in each pair of sister damselfish species on both sides of the Isthmus of Panama, (2) determine whether these variables show specific patterns on each side of the isthmus, and (3) determine whether these variables are correlated through evolution. The results showed no relationship between morphology and niche, however, size was related to both niche and morphology. Sister damselfish species on either side of the Isthmus of Panama differ in terms of niche, morphology, and size. Nevertheless, they also show similarities, indicating environmental constraints and conservatism. This study describes a model in which adaptation or divergence and conservatism shaped the evolution of sister damselfish species on both sides of the Isthmus of Panama. These mechanisms are fundamental to population biology, and they act in opposite directions.

Phylogeography of the Mantled Howler Monkey (Alouatta palliata; Atelidae, Primates) across Its Geographical Range by Means of Mitochondrial Genetic Analyses and New Insights about the Phylogeny of Alouatta

We analyzed 156 specimens of diverse howler monkey taxa (Alouatta; Atelidae, Primates) for different mitochondrial genes (5,567 base pairs), with special emphasis on A. palliata and related taxa. Our results showed no relevant differences among individuals of different putative taxa, A. p. palliata, A. p. aequatorialis, A. coibensis coibensis, and A. c. trabeata. We found no spatial differences in genetic structure of A. p. palliata throughout Costa Rica, Nicaragua, and Honduras. A. p. mexicana (genetic distance: 1.6-2.1%) was the most differentiated taxon within A. palliata. Therefore, we postulate the existence of only 2 clearly defined subspecies within A. palliata (A. p. palliata and A. p. mexicana). A. palliata and A. pigra (traditionally considered a subspecies of A. palliata) are 2 clearly differentiated species as was demonstrated by Cortés-Ortiz and colleagues in 2003, with a temporal split between the 2 species around 3.6-3.7 million years ago (MYA). Our results with the Median Joining Network procedure showed that the ancestors of the cis-Andean Alouatta gave rise to the ancestors of the trans-Andean Alouatta around 6.0-6.9 MYA. As Cortés-Ortiz et al. showed, A. sara and A. macconnelli are differentiable species from A. seniculus, although the first 2 taxa were traditionally considered subspecies of A. seniculus. Our findings agree with the possibility that the ancestor of A. sara gave rise to the ancestor of A. pigra in northern South America. In turn, the ancestor of A. pigra originated the ancestor of A. palliata. Two of our results strongly support the hypothesis that the South American A. palliata (the putative A. p. aequatorialis) was the original population of this species; it has high genetic diversity and no evidence of population expansion. The Central America A. palliata is the derived population. It has low genetic diversity and there is clear evidence of population expansion. However, A. palliata and A. pigra probably migrated into Central America by 2 different routes: the Isthmus of Panama (A. palliata) and Caribbean island arch (A. pigra). Finally, the red howler monkeys from the island of Trinidad in the Caribbean Sea were not A. macconnelli (= A. s. stramineus) as Groves maintained in his influential 2001 publication on primate taxonomy. This taxon is more related to A. s. seniculus, although it formed a monophyletic clade. Future molecular and karyotypic studies will show if the Trinidad red howler monkeys should be considered as an extension of the Venezuelan taxon, A. arctoidea, as a subspecies of A. seniculus(A. s. seniculus), or, in the case of extensive chromosomal rearrangements, even a new species.

Mitogenomic divergence between three pairs of putative geminate fishes from Panama

We sequenced the complete mitochondrial genomes of three pairs of congeneric peripheral fishes distributed on either side of the Isthmus of Panama in order to test their status as geminate species pairs. Our phylogenetic analysis did not support a sister relationship between Gobiomorus dormitor and G. maculatus and therefore they cannot be considered geminates. The average genetic distance of protein-coding genes between Sicydium altum and S. salvini was more than two times larger than between Atlantic and Pacific Awaous banana, suggesting different timings for their divergence across the Isthmus of Panama.

Phylogeography of Nasutitermes corniger (Isoptera: Termitidae) in the Neotropical Region

BackgroundThe Neotropical Region is known for its biodiversity and ranks third in number of known termite species. However, biogeographic and phylogeographic information of termites of this region is limited compared to other world geographic regions. Nasutitermes corniger is widely distributed in the region and is of considerable economic importance. The goal of this study was to describe the phylogeography of N. corniger in the Neotropical Region, to better understand its evolutionary processes.ResultsThe sampled populations of N. corniger showed high genetic variation. Results indicated strong geographic structure among N. corniger populations, with most haplotypes not broadly shared among separated locations. Phylogeographic analyses showed a dispersal route for N. corniger from Central America into South America via the Isthmus of Panama, with subsequent dispersal through the highlands east of the Andes and into eastern regions of the continent. The majority of haplotypes were limited in distribution to proximal regions, corresponding to particular biomes (Atlantic Forest, Amazonia, Chaco, Cerrado and Caatinga).ConclusionsNasutitermes corniger is suggested to be a good model for biogeographic and phylogeographic studies in the Neotropical Region. This study clarified the phylogeographic history of N. corniger and can contribute to the understanding of biogeographic dispersion processes in the Neotropical Region.

Why Are There No Sea Snakes in the Atlantic?

Roughly 70 species of sea snakes inhabit the Indo-Pacific but are absent from the Atlantic Ocean. Paleoclimatic conditions in the Coral Triangle were favorable for evolutionary transitions to the sea, while those in the Caribbean region and coastlines bordering the Atlantic Ocean were less favorable. The dispersal of sea snakes from the Indian to Atlantic Oceans around the Cape of Good Hope has been prevented by low water temperatures and a lack of precipitation related to the presence of the Benguela Current along the Atlantic coast of southern Africa. The Isthmus of Panama fully separated the Pacific and Atlantic Oceans before the speciation and dispersal of the sole pelagic species of sea snake to reach the Central American Pacific shoreline. Future climatic changes could bring declines or extinctions of sea snakes within their present ranges, but they appear unlikely to favor the dispersal and establishment of sea snakes in the Atlantic Ocean.

Phylogeography and connectivity of molluscan parasites: Perkinsus spp. in Panama and beyond

Panama is a major hub for commercial shipping between two oceans, making it an ideal location to examine parasite biogeography, potential invasions, and the spread of infectious agents. Our goals were to (i) characterise the diversity and genetic connectivity of Perkinsus spp. haplotypes across the Panamanian Isthmus and (ii) combine these data with sequences from around the world to evaluate the current phylogeography and genetic connectivity of these widespread molluscan parasites. We collected 752 bivalves from 12 locations along the coast of Panama including locations around the Bocas del Toro archipelago and the Caribbean and Pacific entrances to the Panama Canal, from December 2012 to February 2013. We used molecular genetic methods to screen for Perkinsus spp. and obtained internal transcribed spacer region (ITS) ribosomal DNA (rDNA) sequences for all positive samples. Our sequence data were used to evaluate regional haplotype diversity and distribution across both coasts of Panama, and were then combined with publicly available sequences to create global haplotype networks. We found 26 ITS haplotypes from four Perkinsus spp. (1–12 haplotypes per species) in Panama. Perkinsus beihaiensis haplotypes had the highest genetic diversity, were the most regionally widespread, and were associated with the greatest number of hosts. On a global scale, network analyses demonstrated that some haplotypes found in Panama were cosmopolitan (Perkinsus chesapeaki, Perkinsus marinus), while others were more geographically restricted (Perkinsus olseni, P. beihaiensis), indicating different levels of genetic connectivity and dispersal. We found some Perkinsus haplotypes were shared across the Isthmus of Panama and several regions around the world, including across ocean basins. We also found that haplotype diversity is currently underestimated and directly related to the number of sequences. Nevertheless, our results demonstrate long-range dispersal and global connectivity for many haplotypes, suggesting that dispersal through shipping probably contributes to these biogeographical patterns.

The Monkeying of the Americas: Primate Biogeography in the Neotropics

Monkeys first arrived in the Neotropics about 36 Ma, and the ancestry of all living Neotropical primates (Platyrrhini) traces to a single common ancestral population from 24 to 19 Ma. The availability of lush Amazonian habitat, the rise of the Andes, the transition from the lacustrine to riverine system in the Amazon Basin, and the intermittent connection between the Amazon and the Atlantic tropical forests have all shaped how primates spread and diversified. Primates outcompeted native South American mammals but faced an influx of North American fauna with the closing of the Isthmus of Panama. Humans, extreme newcomers in the Neotropics, have influenced primate habitat and ecology over the last 13,000 years, with radical transformations in the last 500 years as a result of European colonization and land use change. Neotropical primate biogeography informs taxonomic work and conservation efforts, with a mind toward mitigating effects of direct human impact and human-mediated climate change.

Tethyan changes shaped aquatic diversification.

The Tethys Ocean 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.

Unraveling the diversification history of grasshoppers belonging to the "Trimerotropis pallidipennis" (Oedipodinae: Acrididae) species group: a hotspot of biodiversity in the Central Andes.

The Andean Mountain range has been recognized as one of the biodiversity hotspots of the world. The proposed mechanisms for such species diversification, among others, are due to the elevation processes occurring during the Miocene and the intensive glacial action during the Pleistocene. In this study we investigated the diversification history of the grasshopper Trimerotropis pallidipennis species complex which shows a particularly wide latitudinal and altitudinal distribution range across the northern, central and southern Andes in South America. Many genetic lineages of this complex have been so far discovered, making it an excellent model to investigate the role of the central Andes Mountains together with climatic fluctuations as drivers of speciation. Phylogenetics, biogeographic and molecular clock analyses using a multi-locus dataset revealed that in Peru there are at least two, and possibly four genetic lineages. Two different stocks originated from a common ancestor from North/Central America—would have dispersed toward southern latitudes favored by the closure of the Panama Isthmus giving rise to two lineages, the coastal and mountain lineages, which still coexist in Peru (i.e., T. pallidipennis and T. andeana). Subsequent vicariant and dispersal events continued the differentiation process, giving rise to three to six genetic lineages (i.e., clades) detected in this study, which were geographically restricted to locations dispersed over the central Andes Mountains in South America. Our results provide another interesting example of “island diversification” motored by the topography plus unstable climatic conditions during the Pleistocene, pointing out the presence of a hotspot of diversification in the Andean region of Peru.

Phylogeography of the sand dollar genus Encope: implications regarding the Central American Isthmus and rates of molecular evolution

Vicariant events have been widely used to calibrate rates of molecular evolution, the completion of the Central American Isthmus more extensively than any other. Recent studies have claimed that rather than the generally accepted date of ~3 million years ago (Ma), the Isthmus was effectively complete by the middle Miocene, 13 Ma. We present a fossil calibrated phylogeny of the new world sand dollar genus Encope, based on one nuclear and four mitochondrial genes, calibrated with fossils at multiple nodes. Present day distributions of Encope are likely the result of multiple range contractions and extinction events. Most species are now endemic to a single region, but one widely distributed species in each ocean is composed of morphotypes previously described as separate species. The most recent separation between eastern Pacific and Caribbean extant clades occurred at 4.90 Ma, indicating that the Isthmus of Panama allowed genetic exchange until the Pliocene. The rate of evolution of mitochondrial genes in Encope has been ten times slower than in the closely related genera Mellita and Lanthonia. This large difference in rates suggests that splits between eastern Pacific and Caribbean biota, dated on the assumption of a “universal” mitochondrial DNA clock are not valid.

Origins of the Greenland shark (Somniosus microcephalus): Impacts of ice-olation and introgression.

Herein, we use genetic data from 277 sleeper sharks to perform coalescent‐based modeling to test the hypothesis of early Quaternary emergence of the Greenland shark (Somniosus microcephalus) from ancestral sleeper sharks in the Canadian Arctic‐Subarctic region. Our results show that morphologically cryptic somniosids S. microcephalus and Somniosus pacificus can be genetically distinguished using combined mitochondrial and nuclear DNA markers. Our data confirm the presence of genetically admixed individuals in the Canadian Arctic and sub‐Arctic, and temperate Eastern Atlantic regions, suggesting introgressive hybridization upon secondary contact following the initial species divergence. Conservative substitution rates fitted to an Isolation with Migration (IM) model indicate a likely species divergence time of 2.34 Ma, using the mitochondrial sequence DNA, which in conjunction with the geographic distribution of admixtures and Pacific signatures likely indicates speciation associated with processes other than the closing of the Isthmus of Panama. This time span coincides with further planetary cooling in the early Quaternary period followed by the onset of oscillating glacial‐interglacial cycles. We propose that the initial S. microcephalus–S. pacificus split, and subsequent hybridization events, were likely associated with the onset of Pleistocene glacial oscillations, whereby fluctuating sea levels constrained connectivity among Arctic oceanic basins, Arctic marginal seas, and the North Atlantic Ocean. Our data demonstrates support for the evolutionary consequences of oscillatory vicariance via transient oceanic isolation with subsequent secondary contact associated with fluctuating sea levels throughout the Quaternary period—which may serve as a model for the origins of Arctic marine fauna on a broad taxonomic scale.

Data-driven Discovery: A New Era of Exploiting the Literature and Data

Abstract In the current data-intensive era, the traditional hands-on method of conducting scientific research by exploring related publications to generate a testable hypothesis is well on its way of becoming obsolete within just a year or two. Analyzing the literature and data to automatically generate a hypothesis might become the de facto approach to inform the core research efforts of those trying to master the exponentially rapid expansion of publications and datasets. Here, viewpoints are provided and discussed to help the understanding of challenges of data-driven discovery. The Panama Canal, the 77-kilometer waterway connecting the Atlantic and Pacific oceans, has played a crucial role in international trade for more than a century. However, digging the Panama Canal was an exceedingly challenging process. A French effort in the late 19th century was abandoned because of equipment issues and a significant loss of labor due to tropical diseases transmitted by mosquitoes. The United States officially took control of the project in 1902. The United States replaced the unusable French equipment with new construction equipment that was designed for a much larger and faster scale of work. Colonel William C. Gorgas was appointed as the chief sanitation officer and charged with eliminating mosquito-spread illnesses. After overcoming these and additional trials and tribulations, the Canal successfully opened on August 15, 1914. The triumphant completion of the Panama Canal demonstrates that using the right tools and eliminating significant threats are critical steps in any project. More than 100 years later, a paradigm shift is occurring, as we move into a data-centered era. Today, data are extremely rich but overwhelming, and extracting information out of data requires not only the right tools and methods but also awareness of major threats. In this data-intensive era, the traditional method of exploring the related publications and available datasets from previous experiments to arrive at a testable hypothesis is becoming obsolete. Consider the fact that a new article is published every 30 seconds (Jinha, 2010). In fact, for the common disease of diabetes, there have been roughly 500,000 articles published to date; even if a scientist reads 20 papers per day, he will need 68 years to wade through all the material. The standard method simply cannot sufficiently deal with the large volume of documents or the exponential growth of datasets. A major threat is that the canon of domain knowledge cannot be consumed and held in human memory. Without efficient methods to process information and without a way to eliminate the fundamental threat of limited memory and time to handle the data deluge, we may find ourselves facing failure as the French did on the Isthmus of Panama more than a century ago. Scouring the literature and data to generate a hypothesis might become the de facto approach to inform the core research efforts of those trying to master the exponentially rapid expansion of publications and datasets (Evans & Foster, 2011). In reality, most scholars have never been able to keep completely up-to-date with publications and datasets considering the unending increase in quantity and diversity of research within their own areas of focus, let alone in related conceptual areas in which knowledge may be segregated by syntactically impenetrable keyword barriers or an entirely different research corpus. Research communities in many disciplines are finally recognizing that with advances in information technology there needs to be new ways to extract entities from increasingly data-intensive publications and to integrate and analyze large-scale datasets. This provides a compelling opportunity to improve the process of knowledge discovery from the literature and datasets through use of knowledge graphs and an associated framework that integrates scholars, domain knowledge, datasets, workflows, and machines on a scale previously beyond our reach (Ding et al., 2013).

Phylogeny and biogeography of a subclade of mangoes (Aves, Trochilidae)

In this study we explore the phylogenetic relationships within the hummingbird genera Doryfera, Schistes and Colibri (Family Trochilidae), distributed in the Andes, the Pantepui, the southern Brazilian uplands and the lowlands of South America including the Choco, the Amazon Basin, the Chaco, the Cerrado, and the southeastern Brazilian coast, as well as Central and Mesoamerica. To do this, we included a comprehensive sampling of the 16 traditionally recognized subspecies within this group. We found that Doryfera, Schistes and Colibri form a well-supported monophyletic group, and that most of the traditionally recognized subspecies are indeed evolutionary lineages. As there is a high likelihood that the ancestors of this clade of hummingbirds were distributed in the lowlands, we ask: what events might account for the diversification of this subclade of mangoes into the Andes with a later potential dispersal episode from the Andes to the lowlands? We found that several phenomena such as the uplift of the Andes, the marine transgressions of the Plio–Pleistocene, the final closure of the Isthmus of Panama and the climatic oscillations of the Pleistocene might be at least, in part, responsible for the diversification of this group in both the lowlands and the highlands of this region of South America.

Neogene Proto-Caribbean porcupinefishes (Diodontidae).

Fossil Diodontidae in Tropical America consist mostly of isolated and fused beak-like jawbones, and tooth plate batteries. These durophagous fishes are powerful shell-crushing predators on shallow water invertebrate faunas from Neogene tropical carbonate bottom, rocky reefs and surrounding flats. We use an ontogenetic series of high-resolution micro CT of fossil and extant species to recognize external and internal morphologic characters of jaws and tooth plate batteries. We compare similar sizes of jaws and/or tooth-plates from both extant and extinct species. Here, we describe three new fossil species including †Chilomycterus exspectatus n. sp. and †Chilomycterus tyleri n. sp. from the late Miocene Gatun Formation in Panama, and †Diodon serratus n. sp. from the middle Miocene Socorro Formation in Venezuela. Fossil Diodontidae review included specimens from the Neogene Basins of the Proto-Caribbean (Brazil: Pirabas Formation; Colombia: Jimol Formation, Panama: Gatun and Tuira formations; Venezuela: Socorro and Cantaure formations). Diodon is present in both the Atlantic and Pacific oceans, whereas the distribution of Chilomycterus is highly asymmetrical with only one species in the Pacific. It seems that Diodon was as abundant in the Caribbean/Western Atlantic during the Miocene as it is there today. We analyze the paleogeographic distribution of the porcupinefishes group in Tropical America, after the complete exhumation of the Panamanian isthmus during the Pliocene.

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.

Comment (2) on "Formation of the Isthmus of Panama" by O'Dea et al.

O'Dea et al. challenged the inference that the Isthmus of Panama has been in place for the last 10 million years or more and from "an exhaustive review and reanalysis of geological, paleontological, and molecular records," they argued for a "formation of the Isthmus of Panama sensu stricto around 2.8 Ma." I review environmental changes since ~5 Ma throughout Earth, and I argue that environmental changes in the Central American-Caribbean region have been part of a concurrent, worldwide phenomenon that requires a global, not local, explanation. Accordingly, evidence of environmental change from the Central American-Caribbean region does not implicate the emergence of the Isthmus of Panama.

Comment (1) on "Formation of the Isthmus of Panama" by O'Dea et al.

A review and reanalysis of geological, molecular, and paleontological data led O'Dea et al. (1) to propose (i) that reports by Montes et al. (2) and Bacon et al. (3) regarding a middle Miocene closure of the Central American Seaway (CAS) are unsupported, and (ii) a new age of the formation of the Isthmus at 2.8 million years ago (Ma). Here, we reject both of these conclusions.