Natural populations often experience the weakening or removal of a source of selection that had been important in the maintenance of one or more traits. Here we refer to these situations as 'relaxed selection,' and review recent studies that explore the effects of such changes on traits in their ecological contexts. In a few systems, such as the loss of armor in stickleback, the genetic, developmental and ecological bases of trait evolution are being discovered. These results yield insights into whether and how fast a trait is reduced or lost under relaxed selection. We provide a prospectus and a framework for understanding relaxed selection and trait loss in natural populations. We also examine its implications for applied issues, such as antibiotic resistance and the success of invasive species.

/pdf/relaxed-selection-in-the-wild-58btxdtkqk.pdf

Relaxed selection in the wild

Methods for the Study of Marine Benthos

Background 
Although polychaetes are one of the dominant taxa in marine communities, their distributions and taxonomic diversity are poorly understood. Recent studies have shown that many species thought to have broad distributions are actually a complex of allied species. In Canada, 12% of polychaete species are thought to occur in Atlantic, Arctic, and Pacific Oceans, but the extent of gene flow among their populations has not been tested.


Methodology/Principal Findings 
Sequence variation in a segment of the mitochondrial cytochrome c oxidase I (COI) gene was employed to compare morphological versus molecular diversity estimates, to examine gene flow among populations of widespread species, and to explore connectivity patterns among Canada's three oceans. Analysis of 1876 specimens, representing 333 provisional species, revealed 40 times more sequence divergence between than within species (16.5% versus 0.38%). Genetic data suggest that one quarter of previously recognized species actually include two or more divergent lineages, indicating that richness in this region is currently underestimated. Few species with a tri-oceanic distribution showed genetic cohesion. Instead, large genetic breaks occur between Pacific and Atlantic-Arctic lineages, suggesting their long-term separation. High connectivity among Arctic and Atlantic regions and low connectivity with the Pacific further supports the conclusion that Canadian polychaetes are partitioned into two distinct faunas.


Conclusions/Significance 
Results of this study confirm that COI sequences are an effective tool for species identification in polychaetes, and suggest that DNA barcoding will aid the recognition of species overlooked by the current taxonomic system. The consistent geographic structuring within presumed widespread species suggests that historical range fragmentation during the Pleistocene ultimately increased Canadian polychaete diversity and that the coastal British Columbia fauna played a minor role in Arctic recolonization following deglaciation. This study highlights the value of DNA barcoding for providing rapid insights into species distributions and biogeographic patterns in understudied groups.

/pdf/a-tri-oceanic-perspective-dna-barcoding-reveals-geographic-4y8dx1telk.pdf

A tri-oceanic perspective: DNA barcoding reveals geographic structure and cryptic diversity in Canadian polychaetes.

Many coral species spawn simultaneously and have compatible gametes, leading to controversy over the nature of species boundaries and the frequency with which hybridization occurs. Three western Atlantic corals, Montastraea annularis, M. faveolata, and M. franksi, typify this controversy; they all spawn sympatrically on the same evenings after the fall full moons. Here we show, in both Panama and the Bahamas for multiple years, how a variety of mechanisms may act in concert to reproductively isolate all three species. Field studies indicate that M. franksi spawns two hours earlier than the other two species, and the eggs released during this earlier period disperse an average of 500 m by the time the other species spawn. Field measures of fertilization indicate that peak fertilization occurs when spawning synchrony is high and that corals that spawn at the tails of the spawning distributions have greatly reduced fertilization success. Laboratory studies indicate that there is a gametic incompatibility between M. faveolata and the other two species. There are regional differences in the gametic compatibility of M. franksi and M. annularis. In Panama, the two species are completely compatible, whereas in the Bahamas, M. franksi sperm can fertilize M. annularis eggs but the reciprocal cross often fails. Gamete age influences patterns of fertilization, such that very young eggs seem resistant to fertilization and old sperm lose viability after two hours. In sum, the combination of temporal differences in spawning, sperm aging, gamete dispersal and dilution, and gametic incompatibility act in various combinations among the three species, making it unlikely that hybrid fertilization would occur.

/pdf/mechanisms-of-reproductive-isolation-among-sympatric-56v2b8pzxv.pdf

Mechanisms of reproductive isolation among sympatric broadcast-spawning corals of the montastraea annularis species complex

Hypervariability is a prominent feature of large gene families that mediate interactions between organisms, such as venom-derived toxins or immunoglobulins. In order to study mechanisms for evolution of hypervariability, we examined an EST-generated assemblage of 170 distinct conopeptide sequences from the venoms of five species of marine Conus snails. These sequences were assigned to eight gene families, defined by conserved elements in the signal domain and untranslated regions. Order-of-magnitude differences were observed in the expression levels of individual conopeptides, with five to seven transcripts typically comprising over 50% of the sequenced clones in a given species. The conopeptide precursor alignments revealed four striking features peculiar to the mature peptide domain: (1) an accelerated rate of nucleotide substitution, (2) a bias for transversions over transitions in nucleotide substitutions, (3) a position-specific conservation of cysteine codons within the hypervariable region, and (4) a preponderance of nonsynonymous substitutions over synonymous substitutions. We propose that the first three observations argue for a mutator mechanism targeted to mature domains in conopeptide genes, combining a protective activity specific for cysteine codons and a mutagenic polymerase that exhibits transversion bias, such as DNA polymerase V. The high D:(n)/D:(s) ratio is consistent with positive or diversifying selection, and further analyses by intraspecific/interspecific gene tree contingency tests weakly support recent diversifying selection in the evolution of conopeptides. Since only the most highly expressed transcripts segregate in gene trees according to the feeding specificity of the species, diversifying selection might be acting primarily on these sequences. The combination of a targeted mutator mechanism to generate high variability with the subsequent action of diversifying selection on highly expressed variants might explain both the hypervariability of conopeptides and the large number of unique sequences per species.

/pdf/mechanisms-for-evolving-hypervariability-the-case-of-478ko9efm6.pdf

Mechanisms for Evolving Hypervariability: The Case of Conopeptides

The extraordinary diversity of larval form and function in marine invertebrates has motivated many studies of development, ecology, and evolution. Among organisms with pelagic development via a larval stage, this diversity is often reduced to a dichotomy between two broad nutritional categories: planktotrophy and lecithotrophy. Despite the clear utility of the planktotrophy-lecithotrophy dichotomy to those interested in the history or consequences of life history patterns, it is also clear that a number of larval forms do not fit neatly into either of these general categories. Here we review studies of these intermediate larval forms, focusing on descriptions of larvae known as facultative feeders. Recent descriptions of larval development suggest that facultative feeders and other intermediate larval forms are not as rare as commonly assumed. We assess the importance of these forms for models of life-history evolution and call for a more-detailed and nuanced view of larval biology to account for their existence. Clearer knowledge of the phylogenetic distribution and frequency of occurrence of larvae that exhibit intermediate nutritional requirements is also essential for evaluating current ideas on evolutionary transitions between planktotrophy and lecithotrophy. Finally, intermediate larval types provide valuable and underutilized opportunities for testing hypotheses in the fields of larval ecology and the evolution of development.

http://wmpeople.wm.edu/asset/index/jdallen/allenandpernet2007

Intermediate modes of larval development: bridging the gap between planktotrophy and lecithotrophy.

Knowledge of the attributes of the venom-injecting radular teeth of Conus can enhance understanding of the functional biology of feeding and the systematics of this large and taxonomically difficult genus of gastropods. We define and provide a scheme for coding the states of Conus radular tooth characters, in order to facilitate their use in taxonomic and phylogenetic studies. To exemplify these characters, we describe and illustrate teeth of putatively primitive species and of species representing generalized, vermivorous, molluscivorous and piscivorous feeding groups within the genus. We define and address the intraspecific and interspecific variation of ten presenceabsence characters and 15 continuous characters, of which at least five and ten, respectively, are present in most species. Some continuous characters are bimodally distributed among the species sampled. If this distribution still obtains when additional species are examined, these characters could also be coded unambiguously as having two states. We also review the several previously proposed classification schemes of Conus radular teeth and of species according to tooth characters. As Troschel suggested in 1866, a tooth length:width ratio greater than or less than 20 separates the genus into two distinct species groups, now known to comprise the molluscivorous and piscivorous species in the former, and vermivorous and generalized species in the latter. Important subsidiary characters are the number of barbs, presence/ absence of blade, cusp, and spur, length of serrations, relative width of base, and tooth length:shell length ratio.

/pdf/snail-spears-and-scimitars-a-character-analysis-of-conus-4ag8cizjbw.pdf

Snail spears and scimitars: a character analysis of conus radular teeth

The evolution of gamete incompatibility between free-spawning marine invertebrate species has been explained by three hypotheses: (1) independent divergence at gamete recognition loci; (2) selection against hybrids; and (3) a process of sexual selection involving polymorphic gamete recognition loci (Metz and Palumbi 1996). The first two hypotheses predict that gamete incompatibility appears only after gene flow has been halted for other reasons and the third that gamete incompatibility appears simultanously with blocks to gene flow. Here I show that gametes of three sympatric polychaetes in the genus Arctonoe are compatible in all crosses, over a broad range of gamete concentrations and contact times. Although at least some hybrid crosses produce fertile adults, allozyme and mitochondrial DNA sequence data indicate that the three species do not regularly exchange genes. These data are consistent with predictions of the first two hypotheses for the evolution of gamete incompatibility, but allow rejection of the third hypothesis. Gametes of the three species are compatible despite estimated divergence times of 1-3 M.Y.B.P.; in several other marine invertebrates, divergence times of the same magnitude are associated with asymmetric or complete gamete incompatibility. It appears likely that segregation of symbiotic adults on their respective host species restricts mating opportunities, and thus gene flow, among Arctonoe species.

https://onlinelibrary.wiley.com/doi/pdf/10.1111/j.1558-5646.1999.tb03779.x

Gamete interactions and genetic differentiation among three sympatric polychaetes.

Evolutionary loss of the requirement for feeding in larvae of marine invertebrates is often followed by loss of structures involved in capturing and digesting food. Studies of echinoderms suggest that larval form evolves rapidly in response to loss of the requirement for feeding, but a lack of data from other taxa makes it difficult to assess the generality of this result. I show that many members of a large clade of annelids, the Sabellidae, retain ancestral systems for particle capture despite loss of the need and ability to feed. In at least one species, Schizobranchia insignis, an opposed-band system of prototrochal, food-groove, and metatrochal ciliary bands can concentrate suspended particles and transport them to the mouth, but captured particles are invariably rejected because larvae lack a functional gut. The persistence of particle capture systems in larvae of sabellids suggests that they have lost larval feeding very recently, that opposed bands are inexpensive to construct and operate, or that opposed bands have some alternative function. These observations also suggest a hypothesis on how the ability to feed is lost in larvae of annelids and other spiralians following increases in egg size.

Persistent Ancestral Feeding Structures in Nonfeeding Annelid Larvae

The calcareous housing tubes of fossil species of the polychaete wormsDiplochaetetes andDodecaceria from the Tertiary of Lower California (Mexico) are composed, as well as the tubes of recent species ofDodecaceria, by micritic, peloidal lamellae. The wall ofDiplochaetetes spp. consists of only a few, that ofDodecaceria spp. of numerous lamellae. They form a stromatolitic fabric with intercalated lenses of fibrous calcite/aragonite. Histological investigation of the tubes ofDodecaceria showed that the tube formation is related to two processes. The initial process is weakly controlled by the worm itself (matrix mediated). The worm produces acidic organic mucus substances which are enriched between the soft tissue and the tube wall. The entire mucus inhibits the mineralization of the mucus for a certain time. The mineralization events of the mucus are responsible for the stromatolitic microfabric. Within the spaces between the primary lamellae non-specific EPS (extracellular polymeric substances)-rich mucus is enriched, which controls the formation of fibrous aragonitic crystals and peloidal fabrics. This is a very characteristic organomineralization process which is not controlled directly by the organism. The entire process is comparable with those seen in complex rigid modern microbialites from salt and alkaline lakes in Western Australia and Nevada.

Bruno Pernet

Papers

Intermediate modes of larval development: bridging the gap between planktotrophy and lecithotrophy.

Snail spears and scimitars: a character analysis of conus radular teeth

Gamete interactions and genetic differentiation among three sympatric polychaetes.

Persistent Ancestral Feeding Structures in Nonfeeding Annelid Larvae

Organomineralization of cirratulid annelid tubes-fossil and recent examples