Matthew L. Knope

Cope’s rule in the evolution of marine animals

Getting bigger all the time In todays world, many animal species are large, with even larger species only recently extinct, but the first animals to evolve were tiny. Was this increase in size due to active selection or to some more random process? Heim et al. test the classic hypothesis known as Copes rule, which posits that there is selection for increasing body size. They analyzed a data set that spans over 500 million years and includes more than 17,000 marine animal species. In support of Copes rule, body volumes have increased by over five orders of magnitude since the first animals evolved. Furthermore, modeling suggests that such a massive increase could not have emerged from a random process. Science, this issue p. 867 Diversification produced a 150-fold increase in the mean size of marine animals over the past 542 million years. Cope’s rule proposes that animal lineages evolve toward larger body size over time. To test this hypothesis across all marine animals, we compiled a data set of body sizes for 17,208 genera of marine animals spanning the past 542 million years. Mean biovolume across genera has increased by a factor of 150 since the Cambrian, whereas minimum biovolume has decreased by less than a factor of 10, and maximum biovolume has increased by more than a factor of 100,000. Neutral drift from a small initial value cannot explain this pattern. Instead, most of the size increase reflects differential diversification across classes, indicating that the pattern does not reflect a simple scaling-up of widespread and persistent selection for larger size within populations.

Metabolic dominance of bivalves predates brachiopod diversity decline by more than 150 million years

Brachiopods and bivalves feed in similar ways and have occupied the same environments through geological time, but brachiopods were far more diverse and abundant in the Palaeozoic whereas bivalves dominate the post-Palaeozoic, suggesting a transition in ecological dominance 250 Ma. However, diversity and abundance data alone may not adequately describe key changes in ecosystem function, such as metabolic activity. Here, we use newly compiled body size data for 6066 genera of bivalves and brachiopods to calculate metabolic rates and revisit this question from the perspective of energy use, finding that bivalves already accounted for a larger share of metabolic activity in Palaeozoic oceans. We also find that the metabolic activity of bivalves has increased by more than two orders of magnitude over this interval, whereas brachiopod metabolic activity has declined by more than 50%. Consequently, the increase in bivalve energy metabolism must have occurred via the acquisition of new food resources rather than through the displacement of brachiopods. The canonical view of a mid-Phanerozoic transition from brachiopod to bivalve dominance results from a focus on taxonomic diversity and numerical abundance as measures of ecological importance. From a metabolic perspective, the oceans have always belonged to the clams.

Molecular support for marine sculpin (Cottidae; Oligocottinae) diversification during the transition from the subtidal to intertidal habitat in the Northeastern Pacific Ocean

Sculpins in the genera Ruscarius, Artedius, Clinocottus, and Oligocottus are common intertidal and subtidal benthic fishes of the Northeastern Pacific Ocean. While there has been a long history of attempts to reconstruct the evolutionary relationships within this clade, studies have largely resulted in conflicting conclusions. Current ideas regarding the limits of species and genera in this subfamily (Oligocottinae) and their branching order are based primarily on morphology [Bolin, R.L., 1944. A Review of the Marine Cottid Fishes of California. Natural History Museum of Stanford [corrected] University, Stanford [corrected] University, California; Bolin, R.L., 1947. The Evolution of the Marine Cottidae of California with a Discussion of the Genus as a Systematic Category. Stanford [corrected] University, California]. The primary objectives of this study are: (a) to determine if the phylogenetic relationships inferred from DNA characters are concordant with those inferred from morphological characters and (b) to determine if a habitat shift from the subtidal to the intertidal environment resulted in the diversification of the group. Cytochrome b and Nicotinamide Adenine Dinucleotide Dehydrogenase subunit one mitochondrial gene fragments and one nuclear intron (S7 ribosomal protein) were sequenced in order to infer the phylogenetic relationships within this subfamily. Maximum likelihood and Bayesian algorithms were employed to reconstruct phylogenetic trees. We found that several genera in this clade are not monophyletic and that there is a clear phylogenetic signal indicating that a habitat shift from the subtidal to the intertidal habitat has resulted in the diversification of the Oligocottinae.

Evolutionary history, immigration history, and the extent of diversification in community assembly

During community assembly, species may accumulate not only by immigration, but also by in situ diversification. Diversification has intrigued biologists because its extent varies even among closely related lineages under similar ecological conditions. Recent research has suggested that some of this puzzling variation may be caused by stochastic differences in the history of immigration (relative timing and order of immigration by founding populations), indicating that immigration and diversification may affect community assembly interactively. However, the conditions under which immigration history affects diversification remain unclear. Here we propose the hypothesis that whether or not immigration history influences the extent of diversification depends on the founding populations’ prior evolutionary history, using evidence from a bacterial experiment. To create genotypes with different evolutionary histories, replicate populations of Pseudomonas fluorescens were allowed to adapt to a novel environment for a short or long period of time (approximately 10 or 100 bacterial generations) with or without exploiters (viral parasites). Each evolved genotype was then introduced to a new habitat either before or after a standard competitor genotype. Most genotypes diversified to a greater extent when introduced before, rather than after, the competitor. However, introduction order did not affect the extent of diversification when the evolved genotype had previously adapted to the environment for a long period of time without exploiters. Diversification of these populations was low regardless of introduction order. These results suggest that the importance of immigration history in diversification can be predicted by the immigrants’ evolutionary past. The hypothesis proposed here may be generally applicable in both micro- and macro-organisms.

Adaptive morphological shifts to novel habitats in marine sculpin fishes.

Sculpin fishes of the North American Pacific Coast provide an ideal opportunity to examine whether adaptive morphological character shifts have facilitated occupation of novel habitat types because of their well‐described phylogeny and ecology. In this group, the basal‐rooted species primarily occupy the subtidal habitat, whereas the species in the most distal clades are found in the intertidal. We tested multiple evolutionary models to determine whether changes in body size and changes in number of scales are adaptive for habitat use in sculpins. Based on a statistically robust, highly resolved molecular phylogeny of 26 species of sculpins, in combination with morphometric and habitat affinity data, our analyses show that an adaptive model based on habitat use best explains changes in body size and number of scales. The habitat model was statistically supported over models of neutral evolution, stabilizing selection across all habitats, and three clade‐based models. We suggest that loss of scales and reduction of body size in the intertidal may facilitate cutaneous breathing in air when tidepools become hypoxic during low tides. This study demonstrates how the combined use of phylogenetic, ecological and statistical approaches helps to identify traits that are likely adaptive to novel habitats.

Phylogenetics of the marine sculpins (Teleostei: Cottidae) of the North American Pacific Coast.

With 92 species along the North American Pacific Coast, marine sculpins represent the most species-rich radiation of fishes in this region. I used the mitochondrial cytochrome b gene and the nuclear ribosomal S7 intron for 99 species (76 North American, 19 Asian, and four North Atlantic) to produce the most complete phylogenetic hypothesis yet generated for this assemblage. Maximum likelihood and Bayesian analyses produced highly similar tree topologies. While many previously proposed groupings based on morphology are recovered, the molecular data suggest that a number of genera are para- or polyphyletic. However, this analysis supports the monophyly of one large clade that is found exclusively along the North American Pacific Coast (Chitonotous-Ruscarius-Artedius-Orthonopius-Clinocottus-Leiocottus-Oligocottus). Some sibling species have disjunct ranges, suggesting allopatric speciation. However, many other sibling species have largely overlapping ranges, and repeated habitat shifts appear to have facilitated diversification.

Limited role of functional differentiation in early diversification of animals

The origin of most animal phyla and classes during the Cambrian explosion has been hypothesized to represent an ‘early burst’ of evolutionary exploration of functional ecological possibilities. However, the ecological history of marine animals has yet to be fully quantified, preventing an assessment of the early-burst model for functional ecology. Here we use ecological assignments for 18,621 marine animal genera to assess the relative timing of functional differentiation versus taxonomic diversification from the Cambrian to the present day. We find that functional diversity increased more slowly than would be expected given the history of taxonomic diversity. Contrary to previous inferences of rapid ecological differentiation from the early appearances of all well-fossilized phyla and classes, explicit coding of functional characteristics demonstrates that Cambrian genera occupied comparatively few modes of life. Functional diversity increased in the Ordovician and, especially, during the recoveries from the end-Permian and end-Cretaceous mass extinctions. Permanent shifts in the relationship between functional and taxonomic diversity following the era-bounding extinctions indicates a critical role for these biotic crises in coupling taxonomic and functional diversity.

Invasive congeners are unlikely to hybridize with native Hawaiian Bidens (Asteraceae)

UNLABELLED PREMISE OF THE STUDY Invasive plant species threaten native plants in multiple ways, one of which is genetic assimilation through hybridization. However, information regarding hybridization between related alien and native plant species is generally lacking. In Hawaii, the invasive Central American species Bidens pilosa and Bidens alba have colonized natural areas and often grow alongside the native Hawaiian Bidens species, a clade representing an adaptive radiation of 27 endemic taxa, many of which are threatened or endangered. • METHODS To assess the risk of hybridization between introduced and native Hawaiian Bidens (which will readily hybridize with one another), we undertook crosses in cultivation between the invasive species and nine native Bidens taxa. • KEY RESULTS The majority of the crosses formed no viable seed. Although seed did mature in several of the crosses, morphological screening of the resulting seedlings indicated that they were the result of self-pollination. • CONCLUSIONS This result suggests that B. alba and B. pilosa are incapable of hybridizing with these Hawaiian Bidens taxa. Further, we found that B. alba in Hawaii was self-compatible, despite self-incompatibility throughout its native range, and that the tetraploid species B. alba and the hexaploid species B. pilosa were cross-compatible, although pollen fertility was low.