David R. Lindberg

Insights into bilaterian evolution from three spiralian genomes

Current genomic perspectives on animal diversity neglect two prominent phyla, the molluscs and annelids, that together account for nearly one-third of known marine species and are important both ecologically and as experimental systems in classical embryology. Here we describe the draft genomes of the owl limpet (Lottia gigantea), a marine polychaete (Capitella teleta) and a freshwater leech (Helobdella robusta), and compare them with other animal genomes to investigate the origin and diversification of bilaterians from a genomic perspective. We find that the genome organization, gene structure and functional content of these species are more similar to those of some invertebrate deuterostome genomes (for example, amphioxus and sea urchin) than those of other protostomes that have been sequenced to date (flies, nematodes and flatworms). The conservation of these genomic features enables us to expand the inventory of genes present in the last common bilaterian ancestor, establish the tripartite diversification of bilaterians using multiple genomic characteristics and identify ancient conserved long- and short-range genetic linkages across metazoans. Superimposed on this broadly conserved pan-bilaterian background we find examples of lineage-specific genome evolution, including varying rates of rearrangement, intron gain and loss, expansions and contractions of gene families, and the evolution of clade-specific genes that produce the unique content of each genome.

UNDERSTANDING AND PREDICTING ECOLOGICAL DYNAMICS: ARE MAJOR SURPRISES INEVITABLE

Ecological surprises, substantial and unanticipated changes in the abundance of one or more species that result from previously unsuspected processes, are a common outcome of both experiments and observations in community and population ecology. Here, we give examples of such surprises along with the results of a survey of well-established field ecologists, most of whom have encountered one or more surprises over the course of their careers. Truly surprising results are common enough to require their consideration in any reasonable effort to characterize nature and manage natural resources. We classify surprises as dynamic-, pattern-, or intervention-based, and we speculate on the common processes that cause ecological systems to so often surprise us. A long-standing and still growing concern in the ecological literature is how best to make predictions of future population and community dynamics. Although most work on this subject involves statistical aspects of data analysis and modeling, the frequency and nature of ecological surprises imply that uncertainty cannot be easily tamed through improved analytical procedures, and that prudent management of both exploited and conserved communities will require precautionary and adaptive management approaches.

Extinctions in ancient and modern seas

In the coming century, life in the ocean will be confronted with a suite of environmental conditions that have no analog in human history. Thus, there is an urgent need to determine which marine species will adapt and which will go extinct. Here, we review the growing literature on marine extinctions and extinction risk in the fossil, historical, and modern records to compare the patterns, drivers, and biological correlates of marine extinctions at different times in the past. Characterized by markedly different environmental states, some past periods share common features with predicted future scenarios. We highlight how the different records can be integrated to better understand and predict the impact of current and projected future environmental changes on extinction risk in the ocean.

Marine biotic interchange between the northern and southern hemispheres

Patterns of bipolar or antitropical distributions occur in a diverse array of marine invertebrate, vertebrate, and plant groups in the eastern Pacific Ocean. Available geologic and paleontological evidence does not support vicariance as a process in the creation of these distributions but instead favors biotic interchange between hemispheres. Moreover, the timing of these events suggests several breaches (both northward and southward) of the tropics rather than a single event. The fossil record is extremely important in delimiting potential hypotheses and allowing correlation with vicariance events. The congruence of some interchanges with major regional tectonic activity and others with Pleistocene glaciations is not surprising and argues for a plurality of mechanisms. Extinction of endemic taxa following interchange among marine invertebrates is rare, and none of the antitropical distributions reviewed here suggests that the arrival of a taxon in the adjoining hemisphere resulted in the extinction of an endemic taxon. Instead, interchange and endemic taxa coexist. In contrast to the extinction patterns, the patterns of radiations are extremely diverse with some immigrant taxa undergoing remarkable radiations, whereas other taxa are represented by single species. Temperate nearshore rocky communities in both the northern and southern hemispheres appear to be mosaics of species that share common ancestry (because of interchange), are cosmopolitan, and have independent origins within the region. Although some communities appear to be organized around products of interchange (e.g., kelp forests of California and Chile), only the taxa have immigrated; linkages and interactions between species are independent and locally derived.

HUMAN INFLUENCES ON TROPHIC CASCADES ALONG ROCKY SHORES

A three-trophic-level interaction among American Black Oystercatchers (Haematopus bachmani), limpets (Lottia spp.), and erect fleshy algae in rocky intertidal communities of central and southern California was documented via manipulative and “natural” experiments. Removal of the territorial limpet (Lottia gigantea) initially caused large increases in the percent cover of erect fleshy algae, followed by a more gradual increase in density of small limpets (Lottia spp.) and a decline in algal cover. Algal cover increased following the removal of small limpets at the sites from which L. gigantea had been removed earlier, thus demonstrating that the large and small limpets had similar inhibitory effects on plant populations. A comparison of sites with and without oystercatchers showed that L. gigantea occupied substrate inclinations in proportion to their availability at sites where oystercatchers were rare, whereas the distribution of L. gigantea was skewed toward vertically inclined substrates where oystercatchers were common. Survival rates of limpets translocated to horizontal and vertical substrates were similar in sites lacking oystercatcher predation, but were much lower on horizontal substrates where oystercatchers were common. Our results are consistent with those from several prior studies in demonstrating that shorelines frequented by humans typically lack oystercatchers. Humans also exploit L. gigantea and reduce populations to low densities of small individuals. These findings may explain why the midlittoral zone of rocky intertidal communities in western North America are so often dominated by high population densities of small limpets.

Molluscan engrailed expression, serial organization, and shell evolution

SUMMARY Whether the serial features found in some molluscs are ancestral or derived is considered controversial. Here, in situ hybridization and antibody studies show iterated engrailed‐gene expression in transverse rows of ectodermal cells bounding plate field development and spicule formation in the chiton, Lepidochitona caverna, as well as in cells surrounding the valves and in the early development of the shell hinge in the clam, Transennella tantilla. Ectodermal expression of engrailed is associated with skeletogenesis across a range of bilaterian phyla, suggesting a single evolutionary origin of invertebrate skeletons. The shared ancestry of bilaterian‐invertebrate skeletons may help explain the sudden appearance of shelly fossils in the Cambrian. Our interpretation departs from the consideration of canonical metameres or segments as units of evolutionary analysis. In this interpretation, the shared ancestry of engrailed‐gene function in the terminal/posterior addition of serially repeated elements during development explains the iterative expression of engrailed genes in a range of metazoan body plans.

Delayed herbivory and the assembly of marine benthic ecosystems

Abstract Phylogenetic analysis of the metazoan evolutionary tree as a whole, and of trees of component major clades, indicates that marine herbivores, defined here as macrophagous consumers of living multicellular attached marine plants, always occupy terminal positions at several scales of analysis. Nearly all living benthic marine herbivores are derived from microphages, detritivores, or predators, and most have post-Paleozoic origins. The derived nature of herbivory in the sea parallels the evolutionary situation among land animals. Pre-Mesozoic marine benthic ecosystems, characterized by relatively low rates of flow of energy and nutrients, may have relied even more heavily on decomposers for the transfer of carbon from primary producers to animals than do living marine ecosystems in the photic zone.

The taxonomist - an endangered race. A practical proposal for its survival.

BackgroundTaxonomy or biological systematics is the basic scientific discipline of biology, postulating hypotheses of identity and relationships, on which all other natural sciences dealing with organisms relies. However, the scientific contributions of taxonomists have been largely neglected when using species names in scientific publications by not citing the authority on which they are based.DiscussionConsequences of this neglect is reduced recognition of the importance of taxonomy, which in turn results in diminished funding, lower interest from journals in publishing taxonomic research, and a reduced number of young scientists entering the field. This has lead to the so-called taxonomic impediment at a time when biodiversity studies are of critical importance.Here we emphasize a practical and obvious solution to this dilemma. We propose that whenever a species name is used, the author(s) of the species hypothesis be included and the original literature source cited, including taxonomic revisions and identification literature - nothing more than what is done for every other hypothesis or assumption included in a scientific publication. In addition, we postulate that journals primarily publishing taxonomic studies should be indexed in ISISM.SummaryThe proposal outlined above would make visible the true contribution of taxonomists within the scientific community, and would provide a more accurate assessment for funding agencies impact and importance of taxonomy, and help in the recruitment of young scientists into the field, thus helping to alleviate the taxonomic impediment. In addition, it would also make much of the biological literature more robust by reducing or alleviating taxonomic uncertainty.

What Happened to Gray Whales during the Pleistocene? The Ecological Impact of Sea-Level Change on Benthic Feeding Areas in the North Pacific Ocean

Background Gray whales (Eschrichtius robustus) undertake long migrations, from Baja California to Alaska, to feed on seasonally productive benthos of the Bering and Chukchi seas. The invertebrates that form their primary prey are restricted to shallow water environments, but global sea-level changes during the Pleistocene eliminated or reduced this critical habitat multiple times. Because the fossil record of gray whales is coincident with the onset of Northern Hemisphere glaciation, gray whales survived these massive changes to their feeding habitat, but it is unclear how. Methodology/Principal Findings We reconstructed gray whale carrying capacity fluctuations during the past 120,000 years by quantifying gray whale feeding habitat availability using bathymetric data for the North Pacific Ocean, constrained by their maximum diving depth. We calculated carrying capacity based on modern estimates of metabolic demand, prey availability, and feeding duration; we also constrained our estimates to reflect current population size and account for glaciated and non-glaciated areas in the North Pacific. Our results show that key feeding areas eliminated by sea-level lowstands were not replaced by commensurate areas. Our reconstructions show that such reductions affected carrying capacity, and harmonic means of these fluctuations do not differ dramatically from genetic estimates of carrying capacity. Conclusions/Significance Assuming current carrying capacity estimates, Pleistocene glacial maxima may have created multiple, weak genetic bottlenecks, although the current temporal resolution of genetic datasets does not test for such signals. Our results do not, however, falsify molecular estimates of pre-whaling population size because those abundances would have been sufficient to survive the loss of major benthic feeding areas (i.e., the majority of the Bering Shelf) during glacial maxima. We propose that gray whales survived the disappearance of their primary feeding ground by employing generalist filter-feeding modes, similar to the resident gray whales found between northern Washington State and Vancouver Island.

Evolution of large body size in abalones (Haliotis): patterns and implications

Kelps and other fleshy macroalgae—dominant reef-inhabiting organisms in cool seas— may have radiated extensively following late Cenozoic polar cooling, thus triggering a chain of evolutionary change in the trophic ecology of nearshore temperate ecosystems. We explore this hypothesis through an analysis of body size in the abalones (Gastropoda; Haliotidae), a widely distributed group in modern oceans that displays a broad range of body sizes and contains fossil representatives from the late Cretaceous (60-75 Ma). Geographic analysis of maximum shell length in living abalones showed that small-bodied species, while most common in the Tropics, have a cosmopolitan distribution, whereas large-bodied species occur exclusively in cold-water ecosys- tems dominated by kelps and other macroalgae. The phylogeography of body size evolution in extant abalones was assessed by constructing a molecular phylogeny in a mix of large and small species obtained from different regions of the world. This analysis demonstrates that small body size is the plesiomorphic state and largeness has likely arisen at least twice. Finally, we compiled data on shell length from the fossil record to determine how (slowly or suddenly) and when large body size arose in the abalones. These data indicate that large body size appears suddenly at the Miocene/Pliocene boundary. Our findings support the view that fleshy-algal dominated ecosys- tems radiated rapidly in the coastal oceans with the onset of the most recent glacial age. We con- clude with a discussion of the broader implications of this change.