nkx3.2 mutant zebrafish accommodate jaw joint loss through a phenocopy of the head shapes of Paleozoic jawless fish.

The vertebrate jaw is a versatile feeding apparatus. To function, it requires a joint between the upper and lower jaws, so jaw joint defects are often highly disruptive and difficult to study. To describe the consequences of jaw joint dysfunction, we engineered two independent null alleles of a single jaw joint marker gene, nkx3.2, in zebrafish. These mutations caused zebrafish to become functionally jawless via fusion of the upper and lower jaw cartilages (ankylosis). Despite lacking jaw joints, nkx3.2 mutants survived to adulthood and accommodated this defect by: (a) having a remodeled skull with a fixed open gape, reduced snout and enlarged branchial region; and (b) performing ram feeding in the absence of jaw-generated suction. The late onset and broad extent of phenotypic changes in the mutants suggest that modifications to the skull are induced by functional agnathia, secondarily to nkx3.2 loss of function. Interestingly, nkx3.2 mutants superficially resemble ancient jawless vertebrates (anaspids and furcacaudiid thelodonts) in overall head shape. Because no homology exists in individual skull elements between these taxa, the adult nkx3.2 phenotype is not a reversal but rather a convergence due to similar functional requirements of feeding without moveable jaws. This remarkable analogy strongly suggests that jaw movements themselves dramatically influence the development of jawed vertebrate skulls. Thus, these mutants provide a unique model with which to: (a) investigate adaptive responses to perturbation in skeletal development; (b) re-evaluate evolutionarily inspired interpretations of phenocopies generated by gene knockdowns and knockouts; and (c) gain insight into feeding mechanics of the extinct agnathans.

Evolutionary transformation of mouthparts from particle-feeding to piercing carnivory in Viper copepods: Review and 3D analyses of a key innovation using advanced imaging techniques.

BACKGROUND: Novel feeding adaptations often facilitate adaptive radiation and diversification. But the evolutionary origins of such feeding adaptations can be puzzling if they require concordant change in multiple component parts. Pelagic, heterorhabdid copepods (Calanoida) exhibit diverse feeding behaviors that range from simple particle feeding to a highly specialized form of carnivory involving piercing mouthparts that likely inject venom. We review the evolutionary history of heterorhabdid copepods and add new high-resolution, 3D anatomical analyses of the muscular system, glands and gland openings associated with this remarkable evolutionary transformation. RESULTS: We examined four heterorhabdid copepods with different feeding modes: one primitive particle-feeder (Disseta palumbii), one derived and specialized carnivore (Heterorhabdus subspinifrons), and two intermediate taxa (Mesorhabdus gracilis and Heterostylites longicornis). We used two advanced, high-resolution microscopic techniques - serial block-face scanning electron microscopy and two-photon excitation microscopy - to visualize mouthpart form and internal anatomy at unprecedented nanometer resolution. Interactive 3D graphical visualizations allowed putative homologues of muscles and gland cells to be identified with confidence and traced across the evolutionary transformation from particle feeding to piercing carnivory. Notable changes included: a) addition of new gland cells, b) enlargement of some (venom producing?) glands, c) repositioning of gland openings associated with hollow piercing fangs on the mandibles, d) repurposing of some mandibular-muscle function to include gland-squeezing, and e) addition of new muscles that may aid venom injection exclusively in the most specialized piercing species. In addition, live video recording of all four species revealed mandibular blade movements coupled to cyclic contraction of some muscles connected to the esophagus. These behavioral and 3D morphological observations revealed a novel injection system in H. subspinifrons associated with piercing (envenomating?) carnivory. CONCLUSIONS: Collectively, these results suggest that subtle changes in mandibular tooth form, and muscle and gland form and location, facilitated the evolution of a novel, piercing mode of feeding that accelerated diversification of the genus Heterorhabdus. They also highlight the value of interactive 3D animations for understanding evolutionary transformations of complex, multicomponent morphological systems.

Remarkably loud snaps during mouth-fighting by a sponge-dwelling worm.

Many aquatic animals, including mammals, fishes, crustaceans and insects, produce loud sounds underwater [1-6]. Soft-bodied worms would seem unlikely to produce a loud snap or pop because such brief, intense sounds normally require extreme movements and sophisticated energy storage and release mechanisms [5]. Surprisingly, we discovered a segmented marine worm that makes loud popping sounds during a highly stereotyped intraspecific agonistic behavior we call 'mouth fighting'. These sounds - sound pressures up to 157 dB re 1 µPa at 1 m, with frequencies in the 1-100 kHz range and a strong signal at ∼6.9 kHz - are comparable to those made by snapping shrimps, which are among the most intense biological sounds that have been measured in the sea [6]. We suggest a novel mechanism for generating ultrafast movements and loud sounds in a soft-bodied animal: thick, muscular pharyngeal walls appear to allow energy storage and cocking; this permits extremely rapid expansion of the pharynx within the worm's body during the strike, which yields an intense popping sound (likely via cavitation) and a rapid influx of water. Clearly, even soft-bodied marine invertebrates can produce remarkably loud sounds underwater. How they do so remains an intriguing biomechanical puzzle that hints at a new type of extreme biology.

Hagfish from the Cretaceous Tethys Sea and a reconciliation of the morphological-molecular conflict in early vertebrate phylogeny.

Hagfish depart so much from other fishes anatomically that they were sometimes considered not fully vertebrate. They may represent: (i) an anatomically primitive outgroup of vertebrates (the morphology-based craniate hypothesis); or (ii) an anatomically degenerate vertebrate lineage sister to lampreys (the molecular-based cyclostome hypothesis). This systematic conundrum has become a prominent case of conflict between morphology- and molecular-based phylogenies. To date, the fossil record has offered few insights to this long-branch problem or the evolutionary history of hagfish in general, because unequivocal fossil members of the group are unknown. Here, we report an unequivocal fossil hagfish from the early Late Cretaceous of Lebanon. The soft tissue anatomy includes key attributes of living hagfish: cartilages of barbels, postcranial position of branchial apparatus, and chemical traces of slime glands. This indicates that the suite of characters unique to living hagfish appeared well before Cretaceous times. This new hagfish prompted a reevaluation of morphological characters for interrelationships among jawless vertebrates. By addressing nonindependence of characters, our phylogenetic analyses recovered hagfish and lampreys in a clade of cyclostomes (congruent with the cyclostome hypothesis) using only morphological data. This new phylogeny places the fossil taxon within the hagfish crown group, and resolved other putative fossil cyclostomes to the stem of either hagfish or lamprey crown groups. These results potentially resolve the morphological-molecular conflict at the base of the Vertebrata. Thus, assessment of character nonindependence may help reconcile morphological and molecular inferences for other major discords in animal phylogeny.

Connecting pattern to process: Growth of spiral shell sculpture in the gastropod Nucella ostrina (Muricidae: Ocenebrinae).

Shell morphology is a well-suited and underused system to examine the development of novel forms. The three-dimensional structure produced (the shell) is separate from the largely two-dimensional tissue that secretes it (the mantle), allowing us to disentangle the pattern from the process. Despite knowing a great deal about the mechanics of shell secretion (process), and the variety of shell shapes that exist (pattern), no effort has been made to understand how the mantle changes to produce different shell shapes. We investigated this question in the dimorphic snail Nucella ostrina, which exhibits both smooth and ribbed shells to determine how ribs are formed by the mantle. Rib thickenings are produced only in the outer calcitic shell layer and secreted by the distal Outer Mantle Epithelium (OME) with increased acid phosphatase activity. The evenly thick inner aragonitic layers are secreted by the proximal OME which expresses acid phosphatase. Here we show that locally thicker ribs in N. ostrina are produced by changing the dimensions of the distal OME: elongation in the direction of growth and increased cell height. This should increase the amount of shell material secreted, producing locally thicker shell (ribs). Preliminary evidence suggests this mechanism may be widespread in gastropods.

Lamellose Axial Shell Sculpture Reduces Gastropod Vulnerability to Sea Star Predation.

Marine gastropods exhibit a stunning diversity of shell sculpture, but the functional significance of many sculpture types remains unknown. Unfortunately, experimental tests of the functional significance of differences between species are complicated by other morphological differences, such as shell microstructure, aperture shape, and shell thickness, that may confound interpretation. The most robust experimental tests are therefore performed using different shell forms within a species. We took advantage of the extensive intraspecific shell variation in the common intertidal gastropod Nucella lamellosa to test the adaptive significance of axial lamellae, a type of shell sculpture found in numerous marine gastropod subfamilies. We offered three forms of N. lamellosa (lamellose, artificially smooth, and naturally smooth) to the predatory sea star Pisaster ochraceus under controlled laboratory conditions. Pisaster ochraceus consumed significantly fewer lamellose snails than either artificially or naturally smooth snails. We suggest that shell lamellae deter sea star predation by impairing their ability to capture or manipulate snail prey or by increasing prey effective size. These results suggest a credible hypothesis for the adaptive significance of lamellar sculpture in marine gastropods and provide a valuable missing piece to the story about adaptive phenotypic plasticity in N. lamellosa shell form.

Parallel Saltational Evolution of Ultrafast Movements in Snapping Shrimp Claws.

How do stunning functional innovations evolve from unspecialized progenitors? This puzzle is particularly acute for ultrafast movements of appendages in arthropods as diverse as shrimps [1], stomatopods [2], insects [3-6], and spiders [7]. For example, the spectacular snapping claws of alpheid shrimps close so fast (∼0.5 ms) that jetted water creates a cavitation bubble and an immensely powerful snap upon bubble collapse [1]. Such extreme movements depend on (1) an energy-storage mechanism (e.g., some kind of spring) and (2) a latching mechanism to release stored energy quickly [8]. Clearly, rapid claw closure must have evolved before the ability to snap, but its evolutionary origins are unknown. Unearthing the functional mechanics of transitional stages is therefore essential to understand how such radical novel abilities arise [9-11]. We reconstructed the evolutionary history of shrimp claw form and function by sampling 114 species from 19 families, including two unrelated families within which snapping evolved independently (Alpheidae and Palaemonidae) [12, 13]. Our comparative analyses, using micro-computed tomography (microCT) and confocal imaging, high-speed video, and kinematic experiments with select 3D-printed scale models, revealed a previously unrecognized "slip joint" in non-snapping shrimp claws. This slip joint facilitated the parallel evolution of a novel energy-storage and cocking mechanism-a torque-reversal joint-an apparent precondition for snapping. Remarkably, these key functional transitions between ancestral (simple pinching) and derived (snapping) claws were achieved by minute differences in joint structure. Therefore, subtle changes in form appear to have facilitated wholly novel functional change in a saltational manner. VIDEO ABSTRACT.

How reversible is development? Contrast between developmentally plastic gain and loss of segments in barnacle feeding legs.

Segmented organisms and structures have fascinated biologists since William Bateson first described homeotic transformation and recognized the fundamental evolutionary significance of segmental organization. On evolutionary time scales, segments may be lost or gained during major morphological transitions. But how segment loss compares to gain on developmental time scales remains mysterious. Here, we examine the ease of reverse development (opposite to normal growth) by comparing developmentally plastic leg segment loss versus gain in individual barnacles transplanted between different water flow conditions. Plastic segment addition occurred rapidly (one to two molts) and exclusively near the limb base. In contrast, developmentally plastic segment loss-the first observation in any arthropod-took much longer (>10 molts) and, remarkably, occurred throughout the leg (23% of losses occurred mid-limb). Segment loss was not a simple reversal of segment addition. Intersegmental membranes fused first, followed by elimination of duplicate tendons and gradual shortening (but not loss) of duplicate setae. Setal loss, in particular, may impose a severe developmental constraint on arthropod segment fusion. This asymmetric developmental potential (time lag of phenotypic response)-plastic segment addition (amplified normal development) is faster and more orderly than segment loss (reverse development)-adds a new dimension to models of developmental plasticity because the cost of making a developmental mistake in one direction will be greater than in the other.

What determines direction of asymmetry: genes, environment or chance?

Conspicuous asymmetries seen in many animals and plants offer diverse opportunities to test how the development of a similar morphological feature has evolved in wildly different types of organisms. One key question is: do common rules govern how direction of asymmetry is determined (symmetry is broken) during ontogeny to yield an asymmetrical individual? Examples from numerous organisms illustrate how diverse this process is. These examples also provide some surprising answers to related questions. Is direction of asymmetry in an individual determined by genes, environment or chance? Is direction of asymmetry determined locally (structure by structure) or globally (at the level of the whole body)? Does direction of asymmetry persist when an asymmetrical structure regenerates following autotomy? The answers vary greatly for asymmetries as diverse as gastropod coiling direction, flatfish eye side, crossbill finch bill crossing, asymmetrical claws in shrimp, lobsters and crabs, katydid sound-producing structures, earwig penises and various plant asymmetries. Several examples also reveal how stochastic asymmetry in mollusc and crustacean early cleavage, in Drosophila oogenesis, and in Caenorhabditis elegans epidermal blast cell movement, is a normal component of deterministic development. Collectively, these examples shed light on the role of genes as leaders or followers in evolution.This article is part of the themed issue 'Provocative questions in left-right asymmetry'.

Mesoscale morphology at nanoscale resolution: serial block-face scanning electron microscopy reveals fine 3D detail of a novel silk spinneret system in a tube-building tanaid crustacean.

BACKGROUND: The study of morphology is experiencing a renaissance due to rapid improvements in technologies for 3D visualization of complex internal and external structures. But 3D visualization of the internal structure of mesoscale objects - those in the 10-1000 µm range - remains problematic. They are too small for microCT, many lack suitable specific fluorescent markers for confocal microscopy, or they require labor-intensive stacking and smoothing of individual TEM images. Here we illustrate the first comprehensive morphological description of a complete mesoscale biological system at nanoscopic resolution using ultra-modern technology for 3D visualization - serial block-face scanning electron microscopy (SBF-SEM). The SBF-SEM machine combines an in-chamber ultramicrotome, which creates a serial array of exposed surfaces, with an SEM that images each surface as it is exposed. The serial images are then stacked automatically by 3D reconstruction software. We used SBF-SEM to study the spinneret (thread-producing) system of a small, tube-dwelling crustacean that weaves tubes of silk. Thread-producing ability is critical for the survival of many small-bodied animals but the basic morphology of these systems remains mysterious due to the limits of traditional microscopy. RESULTS: SBF-SEM allowed us to describe - in full 3D - well-resolved components (glands, ducts, pores, and associated nerves and muscles) of the spinneret system in the thoracic legs and body segments of Sinelobus sp. (Crustacea, Peracarida, Tanaidacea), a tube-building tanaid only 2 mm in body length. The 3D reconstruction by SBF-SEM revealed at nanoscale resolution a unique structure to the gland and duct systems: In each of three thread-producing thoracic segments, two separate ducts, derived from two separate glands located in the body, run through the entire leg and merge at the leg tip just before the spinneret pore opening. We also resolved nerves connecting to individual setae, spines and pores on the walking legs, and individual muscles within each leg segment. CONCLUSIONS: Our results significantly expand our understanding of the diversity of spinneret systems in the Crustacea by providing the first well-resolved view of spinneret components in the peracarid crustacean order, Tanaidacea. More significantly, our results reveal the great power of SBF-SEM technology for comprehensive studies of the morphology of microscopic animals.

Shaving a Shell: Effect of Manipulated Sculpture and Feeding on Shell Growth and Sculpture Development in Nucella lamellosa (Muricidae: Ocenebrinae).

Gastropod shell sculpture offers a novel tool for studying morphological patterning. Existing shell features may be manipulated experimentally to test how alteration affects subsequent shell growth and form. Axial sculpture occurs in many gastropod groups, and spacing of sculpture may be regular or irregular. But how gastropods control sculpture placement during shell growth is unknown. We studied the growth and positioning of axial lamellae in the muricid Nucella lamellosa, and compared these to the superficially similar axial varices seen in other muricids. First, we tested whether the feeding rate had any effect on the rate of addition or positioning of new lamellae. Second, we tested what effect previous shell sculpture had on lamellar placement, and shell growth in general, by removing all shell sculpture and allowing snails to grow over the "shaved" shell surface. Lamellar growth appeared to be relatively plastic; spacing was highly variable both within and among individual snails, and 1-2 weeks were required to complete the addition of a new lamella. Body growth rate was the primary determinant of lamellar growth; past lamellae had no effect on placement of new lamellae or rate of shell length increase. Feeding rate and body size affected only growth in shell length, and had no direct effect on spacing or on the rate of addition of new lamellae. The growth of axial lamellae in N. lamellosa differed from that of varices by exhibiting neither a) regular spacing nor b) a growth hiatus after completion of a lamella. Significantly, despite the obvious impediment of previous sculpture to future shell growth, removal of this sculpture had no observable effect on the rate of body growth or on any aspect of subsequent lamellar growth.

Handed behavior in hagfish--an ancient vertebrate lineage--and a survey of lateralized behaviors in other invertebrate chordates and elongate vertebrates.

Hagfish represent an ancient lineage of boneless and jawless vertebrates. Among several curious behaviors they exhibit, solitary individuals in one dominant genus of hagfish (Eptatretus spp.) regularly rest in a tightly coiled posture. We present the first systematic treatment of this distinctive behavior. Individual northeastern Pacific hagfish (E. stoutii) exhibited significant handedness (preferred orientation of coiling). However, right-coiling and left-coiling individuals were equally common in the population. Individual hagfish likely develop a preference for one direction by repeating the preceding coiling direction. We also revisit classical accounts of chordate natural history and compare the coiling behavior of Eptatretus with other handed or lateralized behaviors in non-vertebrate chordates, lampreys, and derived vertebrates with elongate bodies. Handed behaviors occur in many of these groups, but they likely evolved independently. In contrast to vertebrates, morphological asymmetries may bias lateralized larval behaviors toward one side in cephalochordates and tunicates. As a consequence, no known handed behavior can be inferred to have existed in the common ancestor of vertebrates.

Intertidal sea stars (Pisaster ochraceus) alter body shape in response to wave action.

Sea stars are some of the largest mobile animals able to live in the harsh flow environment of wave-exposed, rocky intertidal shores. In addition, some species, such as the northeastern Pacific Pisaster ochraceus, are ecologically significant predators in a broad range of environments, from sheltered lagoons to the most wave-exposed shorelines. How they function and survive under such an extreme range of wave exposures remains a puzzle. Here we examine the ability of P. ochraceus to alter body form in response to variation in flow conditions. We found that sea stars in wave-exposed sites had narrower arms and were lighter per unit arm length than those from sheltered sites. Body form was tightly correlated with maximum velocity of breaking waves across four sites and also varied over time. In addition, field transplant experiments showed that these differences in shape were due primarily to phenotypic plasticity. Sea stars transplanted from a sheltered site to a more wave-exposed site became lighter per unit arm length, and developed narrower arms, after 3 months. The tight correlation between water flow and morphology suggests that wave force must be a significant selective factor acting on body shape. On exposed shores, narrower arms probably reduce both lift and drag in breaking waves. On protected shores, fatter arms may provide more thermal inertia to resist overheating, or more body volume for gametes. Such plastic changes in body shape represent a unique method by which sea stars adapt to spatial, seasonal and possibly short-term variation in flow conditions.

Something Darwin didn't know about barnacles: spermcast mating in a common stalked species.

Most free-living barnacles are hermaphroditic, and eggs are presumed to be fertilized either by pseudo-copulation or self-fertilization. Although the common northeast Pacific intertidal gooseneck barnacle, Pollicipes polymerus, is believed only to cross-fertilize, some isolated individuals well outside penis range nonetheless bear fertilized eggs. They must therefore either self-fertilize or-contrary to all prior expectations about barnacle mating-obtain sperm from the water. To test these alternative hypotheses, we collected isolated individuals bearing egg masses, as well as isolated pairs where at least one parent carried egg masses. Using 16 single nucleotide polymorphism markers, we confirmed that a high percentage of eggs were fertilized with sperm captured from the water. Sperm capture occurred in 100 per cent of isolated individuals and, remarkably, even in 24 per cent of individuals that had an adjacent partner. Replicate subsamples of individual egg masses confirmed that eggs fertilized by captured sperm occurred throughout the egg mass. Sperm capture may therefore be a common supplement to pseudo-copulation in this species. These observations (i) overturn over a century of beliefs about what barnacles can (or cannot) do in terms of sperm transfer, (ii) raise doubts about prior claims of self-fertilization in barnacles, (iii) raise interesting questions about the capacity for sperm capture in other species (particularly those with short penises), and (iv) show, we believe for the first time, that spermcast mating can occur in an aquatic arthropod.

Effects of body size and shape on locomotion in the bat star (Patiria miniata).

Among taxa ranging from cnidarians to vertebrates, absolute speed of locomotion generally increases with increasing body size. Despite the unique mode of locomotion in echinoderms, crawling speed also appears to increase with increasing body size, at least in some species of asteroids and echinoids. We used an escape-response assay to assess how maximum crawling speed varied with body size in the bat star Patiria miniata. We also tested the effect of arm number on maximum crawling speed by comparing speeds of five- and six-armed individuals. Contrary to prior reports for a single sea urchin and sea star species, both absolute crawling speed and crawling speed relative to body size actually declined with increasing body mass, increasing arm length, and increasing oral surface area, in both five- and six-armed individuals. Arm number did not appear to have a significant effect on crawling speed. The reasons for this negative relationship between crawling speed and body size in P. miniata remain unclear, but we suspect that the disproportionate increase in body mass relative to total tube-foot cross-sectional area may make locomotion proportionally more difficult in larger-bodied sea stars.

Developmental plasticity and the origin of novel forms: unveiling cryptic genetic variation via "use and disuse".

Natural selection eliminates phenotypic variation from populations, generation after generation-an observation that haunted Darwin. So, how does new phenotypic variation arise, and is it always random with respect to fitness? Repeated behavioral responses to a novel environment-particularly those that are learned-are typically advantageous. If those behaviors yield more extreme or novel morphological variants via developmental plasticity, then previously cryptic genetic variation may be exposed to natural selection. Significantly, because the mean phenotypic effect of "use and disuse" is also typically favorable, previously cryptic genetic variation can be transformed into phenotypic variation that is both visible to selection and biased in an adaptive direction. Therefore, use-induced developmental plasticity in a very real sense "creates" new phenotypic variation that is nonrandom with respect to fitness, in contrast to the random phenotypic effects of mutation, recombination, and "direct effects" of environment (stress, nutrition). I offer here (a) a brief review of the immense literature on the effects of "use and disuse" on morphology, (b) a simple yet general model illustrating how cryptic genetic variation may be exposed to selection by developmentally plastic responses that alter trait performance in response to "use and disuse," and (c) a more detailed model of a positive feedback loop between learning (handed behavior) and morphological plasticity (use-induced morphological asymmetry) that may rapidly generate novel phenotypic variation and facilitate the evolution of conspicuous morphological asymmetries. Evidence from several sources suggests that handed behaviors played an important role both in the origin of novel forms (asymmetries) and in their subsequent evolution.

Juvenile colour polymorphism in the red rock crab, Cancer productus: patterns, causes, and possible adaptive significance.

Juveniles of the common red rock crab of the Northeastern Pacific, Cancer productus, display a stunning diversity of colours and patterns, while adults all have the same drab colouration. Although this is widely known, key questions remain: (1) Does the frequency of different juvenile colours or patterns vary among collection sites or seasonally? (2) Does juvenile colour polymorphism reflect genetic heterogeneity or phenotypic plasticity in response to variable environmental conditions? (3) Do juveniles of different colours or patterns prefer substrata of different heterogeneity or brightness? We therefore: (i) described the variation in colour and pattern of juvenile C. productus; (ii) tested for associations between colour/pattern morphs and crab size, collection site, and season, in the field; (iii) conducted preliminary tests for habitat preferences (background colour, substratum type, light level) of different colour/pattern morphs in laboratory experiments, and (iv) tested the effect of diet (mussels versus shrimp) and feeding rate (high versus low) on juvenile colour/pattern. We describe 30 phenotypes that embrace a wide range of colour and pattern variants. The proportions of these phenotypes did not vary significantly among four collection sites, but they did vary significantly with season: over the summer and fall, juvenile colour and pattern variation was gradually replaced by the uniform adult colouration. The number of crabs displaying adult colouration also increased with crab size. Laboratory experiments suggest no significant preferences of different juvenile morphs for different backgrounds, substrata, or light levels. Diet (mussels versus shrimp) and feeding frequency had no effect on colour/pattern. Collectively, these results, although limited in scope, are not consistent with two likely hypotheses that could explain the extensive colour and pattern variation in juvenile C. productus: (i) selection for background matching by different cryptic forms and (ii) direct effects of diet or feeding rate on colour or pattern. Most probably, the large variety of different juvenile morphs is a result of frequency-dependent selection in which abundant variants are attacked disproportionately often and rarer forms are favoured. Juvenile colour polymorphism in C. productus may reduce the vulnerability to visual predators, impede the formation of a search image, and consequently decrease the risk of predation during the juvenile stages.

Elevated CO2 affects shell dissolution rate but not calcification rate in a marine snail.

As CO(2) levels increase in the atmosphere, so too do they in the sea. Although direct effects of moderately elevated CO(2) in sea water may be of little consequence, indirect effects may be profound. For example, lowered pH and calcium carbonate saturation states may influence both deposition and dissolution rates of mineralized skeletons in many marine organisms. The relative impact of elevated CO(2) on deposition and dissolution rates are not known for many large-bodied organisms. We therefore tested the effects of increased CO(2) levels--those forecast to occur in roughly 100 and 200 years--on both shell deposition rate and shell dissolution rate in a rocky intertidal snail, Nucella lamellosa. Shell weight gain per day in live snails decreased linearly with increasing CO(2) levels. However, this trend was paralleled by shell weight loss per day in empty shells, suggesting that these declines in shell weight gain observed in live snails were due to increased dissolution of existing shell material, rather than reduced production of new shell material. Ocean acidification may therefore have a greater effect on shell dissolution than on shell deposition, at least in temperate marine molluscs.

Scale-eating cichlids: from hand(ed) to mouth.

Two recent studies in BMC Biology and Evolution raise important questions about a textbook case of frequency-dependent selection in scale-eating cichlid fishes. They also suggest a fascinating new line of research testing the effects of handed behavior on morphological asymmetry.

Experimental evidence for the rapid evolution of behavioral canalization in natural populations.

Canalization-the evolutionary loss of the capacity of organisms to develop different phenotypes in different environments-is an evolutionary phenomenon suspected to occur widely, although examples in natural populations are elusive. Because behavior is typically a highly flexible component of an individual's phenotype, it provides fertile ground for studying the evolution of canalization. Here we report how snail populations exposed for different lengths of time to a predatory crab introduced from Europe to America exhibit different degrees of canalization of an adaptive antipredator behavior: soft tissue withdrawal, measured as angular retraction depth. Where crab-snail contact is shortest (60 years), snails showed the highest behavioral flexibility. Where crabs invaded 110 years ago, snails showed significantly less behavioral flexibility, and where the interaction is ancient (Europe), snails exhibited highly canalized behavior. Selection therefore appears to have acted rapidly to increase canalization in wild snail populations, leading ultimately to the hard-wired behavior seen in European conspecifics.