Chemical host detection and differential attraction in a parasitic pea crab infecting two echinoids

Mechanisms of host detection and host choice are important for mobile parasites or symbionts that frequently move among host individuals. The parasitic pea crab Dissodactylus primitivus is an ectoparasite exploiting asymmetrically two spatangoid echinoid hosts, Meoma ventricosa and Plagiobrissus grandis. The entire life cycle takes place on M. ventricosa, while only adults colonize P. grandis, the host where female's fecundity is the greatest. Dissodactylus primitivus frequently exhibits host-switching, a behaviour that raises the question of host detection. To test if adult crabs are able to detect potential hosts and are attracted to the host where their fitness is greater, crab olfactory responses were investigated using a Y-tube. We first showed that D. primitivus are attracted by olfactory cues of M. ventricosa, versus those of a non-host echinoid species, Clypeaster rosaceus sharing the same habitat. This attractiveness was not due to a yellowish water-soluble substance secreted by the host. We then showed that the chemical attraction to the two hosts was different according to the origin of the crabs. Crabs coming from M. ventricosa were more attracted to this host than to P. grandis, a pattern compatible with an imprinting phenomenon. However, crabs collected from P. grandis were equally attracted to both host species, as if imprinting was lost after host change. We propose that the pattern of D. primitivus host detection could be intermediate between the behaviour of specialist and generalist symbionts.     

Characterization of eight microsatellite loci for the sea urchin Meoma ventricosa (Spatangoida,Brissidae) through Next Generation Sequencing

Eight microsatellite loci were characterized for Meoma ventricosa (Lamarck, 1816), a burrowing sea urchin that can be afflicted by a bacterial disease causing localized mass mortality. For the analyzed population (29 individuals from St. Croix, US Virgin Islands), we observed 8.125 mean number of alleles, 0.640 mean observed heterozygosity (H_o) and 0.747 mean expected heterozygosity (H_e). Two loci showed significant deviations from Hardy–Weinberg equilibrium. Overall, the described loci were characterized by a moderately high level of polymorphism suggesting that these markers are useful for a population genetic study in the Caribbean Sea.     

Tithonia oxfordiana,a new irregular echinoid associated with Jurassic seep deposits in south‐east France

Abstract: The infaunal irregular echinoid, Tithonia oxfordiana, is described and compared to congeneric species previously described from Upper Jurassic and lowermost Cretaceous strata. This new species characterizes a monospecific echinoid assemblage, which occurs only in some places where deep‐marine middle Oxfordian deposits are exposed in south‐east France. Specimens are closely packed and clearly concentrated at the top of small carbonate chemoherms; a close connection of the echinoids with the emission of reduced chemicals, which were oxidized by chemoautotrophic bacteria, is highly probable. Based on general test shape and plate architecture, T. oxfordiana probably was a deposit feeder on chemosynthetic organic matter produced by such bacteria. In view of the fact that T. oxfordiana is the sole species of the Jurassic genus Tithonia known from Oxfordian strata, it is postulated that chemoherms possibly acted as refugia for these peculiar echinoids, which have an episodic record between the Callovian and Valanginian.     

A Dense Infestation of Round Pits in the Irregular Echinoid Hemipneustes striatoradiatus (Leske) from the Maastrichtian of the Netherlands

A test of the holasteroid echinoid Hemipneustes striatoradiatus (Leske) from the upper Meerssen Member (subunit IVf-6; Maastricht Formation) in the type area of the Maastrichtian Stage (Late Cretaceous) was infested by 170 non-penetrative pits assigned to Oichnus excavatus (Donovan and Jagt). The echinoid is assumed to have been alive at that time. The distribution of pits is approximately bilaterally symmetrical and mainly supraambital; pits on the oral surface may have been carried subambitally by echinoid plate growth. Many pits are comparatively shallow; they probably were partly filled by the echinoid after death of the pit-forming organisms. The area around the periproct is devoid of pits as is the anterior ambulacrum (amb III). The latter was probably kept free of larval pit-formers by the tube feet; other ambulacra, with tube feet adapted for respiration, lacked such defense.     

Scaling of feeding biomechanics in the horn shark Heterodontus francisci: ontogenetic constraints on durophagy

Organismal performance changes over ontogeny as the musculoskeletal systems underlying animal behavior grow in relative size and shape. As performance is a determinant of feeding ecology, ontogenetic changes in the former can influence the latter. The horn shark Heterodontus francisci consumes hard-shelled benthic invertebrates, which may be problematic for younger animals with lower performance capacities. Scaling of feeding biomechanics was investigated in H. francisci (n=16, 19–59 cm standard length (SL)) to determine the biomechanical basis of allometric changes in feeding performance and whether this performance capacity constrains hard-prey consumption over ontogeny. Positive allometry of anterior (8–163 N) and posterior (15–382 N) theoretical bite force was attributed to positive allometry of cross-sectional area in two jaw adducting muscles and mechanical advantage at the posterior bite point (0.79–1.26). Mechanical advantage for anterior biting scaled isometrically (0.52). Fracture forces for purple sea urchins Strongylocentrotus purpuratus consumed by H. francisci ranged from 24 to 430 N. Comparison of these fracture forces to the bite force of H. francisci suggests that H. francisci is unable to consume hard prey early in its life history, but can consume the majority of S. purpuratus by the time it reaches maximum size. Despite this constraint, positive allometry of biting performance appears to facilitate an earlier entry into the durophagous niche than would an isometric ontogenetic trajectory. The posterior gape of H. francisci is significantly smaller than the urchins capable of being crushed by its posterior bite force. Thus, the high posterior bite forces of H. francisci cannot be fully utilized while consuming prey of similar toughness and size to S. purpuratus, and its potential trophic niche is primarily determined by anterior biting capacity.     

Rogerella isp. Infesting the Pore Pairs of Hemipneustes striatoradiatus (Leske) (Echinoidea: Upper Cretaceous,Belgium)

An otherwise well-preserved test of the holasteroid echinoid Hemipneustes striatoradiatus (Leske) from the Emael Member, Maastricht Formation (Maastrichtian, Upper Cretaceous) of Belgium, is infested by encrusting bivalves and foraminiferans and the boring Rogerella isp. In this specimen, Rogerella preferentially infested and modified the ambulacral pore pairs of the echinoid close to the apex. This was not a commensal association. The echinoid test shows no growth deformations in response to this invasion; pore pairs are locally strongly infested; and encrusting invertebrates testify to the long post-mortem residence time of the test on the sea floor. Rather, the pore pairs of the dead echinoid were crannies attractive to settling larvae of acrothoracian barnacles, the producers of Rogerella.     

The contribution of the facultative feeding period to echinoid larval development and size at metamorphosis: a comparative approach

Planktotrophic invertebrate larvae have two dissociable stages during development, a facultative feeding period, whose length is determined by the amount of maternal provisioning in the egg, and an obligate feeding period, whose length is determined by the quantity of exogenous energy needed to reach metamorphic competence. Here we set out to experimentally test the impact of feeding during the facultative feeding period at two food concentrations (limiting and nonlimiting) on larval development time and juvenile quality. We used two closely related subtropical sand dollar species that differ in the quantity of maternal investment for these comparisons: Leodia sexiesperforata (large egg, long facultative feeding period) and Mellita tenuis (small egg, short facultative feeding period). We found that feeding during the facultative period accelerates development to metamorphosis only in M. tenuis and only at the high food ration. Feeding during the facultative feeding period had no effect on development time for M. tenuis at a food limiting concentration and for L. sexiesperforata at either food concentration. Furthermore, we found feeding during the facultative period to significantly increase quantity of carbohydrates and lipids at metamorphosis only for M. tenuis in nonlimiting food concentration. Thus, our data reveal a two-fold benefit of the facultative feeding period for a poorly provisioned species under high food conditions but little effect on a well-provisioned species. We discuss our results in reference to McEdward's [McEdward, L.R., 1997. Reproductive strategies of marine benthic invertebrates revisited: facultative feeding by planktotrophic larvae. Am. Nat. 150, 48-72] facultative feeding model.     

Chronic spearfishing may indirectly affect reef health through reductions in parrotfish bite rates

The grazing behaviour of two Caribbean parrotfish, a fished species, the stoplight parrotfish Sparisoma viride and a non-fished species, the striped parrotfish Scarus iseri, were studied in the presence (fished site) and absence (marine reserve) of chronic spearfishing activity. Diurnal feeding periodicity did not differ between the sites in either species: roving individuals had significantly higher bite rates in the afternoon, while territorial individuals foraged consistently throughout the day. Mean bite rate varied between sites in both species. Abundance, biomass and bite rates of S. viride were all significantly higher within the reserve, except for roving S. viride which had a higher mean bite rate in the afternoon outside the reserve compared with within it, attributable to maximisation of feeding in the afternoon when fishing risk was lower. Scarus iseri mean abundance and bite rate were greater outside the reserve, potentially because reduction in large territorial herbivores allowed S. iseri to feed more rapidly. By reducing the grazing potential of the remaining S. viride individuals the effect of fishing is greater than would be predicted from biomass changes alone. Less grazing by S. viride would not be compensated for by the increase in grazing by S. iseri because the latter feeds on different algae. Spearfishing of key parrotfish species reduces grazing potential directly by extraction and indirectly by changing behaviour.     

Shallow Traces (Pits) in the Test of the Irregular Echinoid Echinocorys scutata Leske from the Chalk (Upper Cretaceous) of the United Kingdom

Specimens of epifaunal irregular echinoids in the Upper Cretaceous of northern Europe have been reported with patterns of circular, nonpenetrative parabolic pits, Oichnus paraboloides (Bromley), in the apical region. Specimens of Echinocorys scutata Leske from the Chalk at two sites in southeast England were commonly penetrated by this trace, generated by an indeterminate pit-forming organism. Pits commonly surround the apical system and, less commonly, occur within it; they occur preferentially anteriorly. Pits occur within plates, not along margins or sutures. Crosscutting of pits indicates that multiple spatfalls probably occurred. The host echinoid added new test plates adjacent to the apical system; thus, plates bearing O. paraboloides were moved abapically. The reduction in number of pits away from the apex, including those with echinoid tubercles reestablished on the base, indicates that, following death of an infester, the echinoid “reclaimed” and infilled them with calcite. The pit-former was most probably an unmineralized invertebrate that used E. scutata as a domicile which provided good access to food-rich currents for suspension feeding. Although the systematic position of the pit-former is unknown, similar infestations are known from other Upper Cretaceous echinoids and Mississippian crinoids.     

Convolvulus grandiflorus Jacq. (Convolvulaceae) re-examined

Convolvulus grandiflorus Jacq. has been treated as a synonym ofOperculina ventricosa andStictocardia tiliifolia. It is here accepted as identifiable withIpomoea (sect.Calonyction)macrantha which is the correct name for Jacquin’s taxon. Full synonymies ofIpomoea macrantha andOperculina ventricosa are given, along with a comment about a variant occurring in Micronesia.     

Site Selectivity of the Pit Oichnus excavatus Donovan and Jagt Infesting Hemipneustes striatoradiatus (Leske) (Echinoidea) in the Type Maastrichtian (Upper Cretaceous,The Netherlands)

Two tests of the holasteroid echinoid Hemipneustes striatoradiatus (Leske) from the type area of the Maastrichtian Stage (Upper Cretaceous) bear a varied infestation of episkeletozoans (oysters, bryozoan colony, and serpulids), borings (probable Caulostrepsis isp., Oichnus simplex Bromley), surface abrasion (Gnathichnus? isp.), and pits (O. excavatus Donovan and Jagt). Only O. excavatus represents a premortem infestation. In one specimen, the four individual pits of this ichnospecies are each associated with a different ambulacrum and pore pairs that, in life, bore respiratory tube feet; the anterior ambulacrum, of different gross morphology, is not infested. In the second test, three out of four of the same ambulacra are infested, although there are also O. excavatus in the interambulacra. The association between O. excavatus and the ambulacra of the echinoid, and thus its tube feet, is open to several plausible explanations, but most likely provided some form of feeding or protective advantage to the pit-forming organism.     

Mate Choice by Both Sexes Maintains Reproductive Isolation in a Species Flock of Pupfish (Cyprinodon spp) in the Bahamas

Female and male mate choices can reinforce reproductive isolation after sympatric speciation. Using a binary choice design, we examine the importance of visual cues in female mate choice in all three sympatric species of pupfish on San Salvador Island. We also examine the importance of olfactory cues in female choice of the hard‐shelled invertebrate specialist (Cyprinodon brontotheroides). We examine male mate choice in two of the three species, the scale eater (C. desquamator) and the detritivore (C. variegatus). Females of all three species use visual cues and prefer conspecific males. C. brontotheroides females do not use olfactory cues to discriminate between conspecific and heterospecific males. Males of C. desquamator and C. variegatus also preferentially court conspecific females. Thus, mutual mate choice, where both females and males exhibit mate choice, acts as a strong behavioral pre‐mating isolation mechanism in these sympatrically speciated pupfish.     

Studies on maternal inheritance in polyploid wheats with cytoplasmic DNAs as genetic markers

Summary Restriction fragment patterns of DNA fragments obtained after EcoRI cleavage of chloroplastic (cp) and mitochondrial (mt) DNAs isolated from different wheat species were compared. T. aestivum, T. timopheevi, Ae. speltoides, Ae. sharonensis and T. urartu gave species specific mt DNA patterns. Consequently, the cytoplasmic genomes of wheat cannot have originated from contemporary Ae. speltoides, Ae. sharonensis and T. urartu species. It is shown that cp and mt DNAs of Ae. ventricosa, a tetraploid used to transfer eyespot resistance into T. aestivum, contains cp and mt DNAs differing from DNAs isolated from T. aestivum and other wheats. In contrast, the cytoplasmic DNAs of Ae. ventricosa and Ae. squarrosa reveal an important homology, suggesting that Ae. squarrosa was the female parent of Ae. ventricosa. Disomic addition lines (T. aestivum — Ae. ventricosa) in both Ae. ventricosa cytoplasm and T. aestivum cytoplasm contained cytoplasmic DNAs identical to those of the maternal parent. Restriction patterns of the cp and mt DNAs isolated from eight lines of Triticale differing in their cytoplasm have been compared to those of the maternal parent. A strict maternal inheritance has been observed in each case.     

A New Species of Centrohelid Heliozoan Acanthocystis amura n. sp. Isolated From Two Remote Locations in Russia

A new species of centrohelid heliozoan Acanthocystis amura from the Amur River, the Far East of Russia was studied with light‐ and electron microscopy. Acanthocystis amura has simple oval plate scales (1.2–2.6 × 1.0–1.8 μm) with an axial thickening and a thin border as well as two types of spine scales. The spine scales of type 1 were 1.3–4.8 μm long and had four small hooks on their tips. The spine scales of type 2 were shorter, 1.0–3.6 μm long and had four teeth at the distal end. The morphology of A. amura resembles that of Acanthocystis quadrifurca. They both have similar spine scales but their plate scales are completely different. Acanthocystis valdiviense differs from the new species in absence of hook‐bearing scales. Molecular phylogeny based on 18S rDNA consistently placed A. amura into Acanthocystis nichollsi/Acanthocystis costata clade, but relationships between those three species were not resolved. Morphology of another centrohelid strain from another location (the South Urals) has been compared with that of the original strain and few minor differences in size characteristics of the scales have been revealed.     

Invaders eating invaders: exploitation of novel alien prey by the alien shimofuri goby in the San Francisco Estuary,California

The shimofuri goby (Tridentiger bifasciatus), which is native to Asian estuaries, was recently introduced to the San Francisco Estuary, California, USA. We conducted gut content analyses to examine the gobys feeding ecology in this highly invaded estuary. Shimofuri gobies were generalist predators on benthic invertebrates, consuming seasonally abundant prey, especially amphipods (Corophium spp.). In addition, shimofuri goby utilized two novel prey items not exploited by other resident fishes – hydroids (Cordylophora caspia) and barnacle (Balanus improvisus) cirri, both of which are alien. The shimofuri gobys feeding ecology appears well-suited to the fluctuating environment of the San Francisco Estuary and may partially explain observed increases in shimofuri goby abundance compared with declines in populations of some native species.     

Relationships between butterflyfish (Chaetodontidae) feeding rates and coral consumption on the Great Barrier Reef

This study explored differences in the feeding rate among 20 species of coral reef butterflyfishes (Chaetodontidae) from Lizard Island, Great Barrier Reef. Feeding rate, measured as bites per minute (b.p.m.), varied between 2.98 ± 0.65 and 12.29 ± 0.27 (mean ± SE) according to species and was positively related to the proportional consumption of coral (r ² = 0.40, n = 20, P < 0.01), independent of phylogeny (standardised independent contrasts r ² = 0.29, n = 19, P < 0.05). All species fed actively throughout the day, with obligate corallivores having a higher feeding rate at all times than either facultative corallivores or non-corallivores. The feeding rate of the obligate corallivores was also highest during the middle of the day. For eight of the species for which data was available, there was a positive correlation between bite rate and competitive dominance (r = 0.71, P < 0.05). Chaetodon ephippium was the only species for which the feeding rate of pairs was higher than for solitary individuals.     

Stratigraphic analysis of a fossil Neogoniolithon-capped patch reef and associated facies,San Salvador,Bahamas

An unusual Pleistocene patch reef is exposed in a coastal cliff at Grotto Beach, San Salvador, Bahamas. The reef is a coralline framestone constructed mainly by Porites astreoides together with a few large heads of Diploria strigosa and Montastrea annularis, and is capped by a dense thicket of Neogoniolithon strictum that is interpreted as marking the subtidal/intertidal boundary. The reef is flanked to the northeast by laminated to low-angle cross-laminated intraclastic grainstones and to the southwest by skeletal rudstone of reefal and interreefal derivation. Uranium-series dating of pure aragonite from a Diploria corallum yielded an age of 123 000±9000 years. Reef growth began on an erosional surface underlain by steeply crossbedded eolian grainstone. As the reef grew upward, it also grew laterally over adjacent penecontemporaneous subtidal sediments. The reef was eventually buried by 2.3 m of shallow subtidal and beach sediments that apparently prograded seaward during a highstand, or possibly while sea level was still rising. The shallow subtidal sediments are mainly peloidal, ooidal and skeletal grainstones that are pervasively bioturbated. The overlying beach facies comprises predominantly laminated, sparsely burrowed grainstone. The beach and shallow subtidal facies contain boulders of fine-grained laminated grainstone that are interpreted as storm-tossed blocks of beachrock. Living analogs of the Grotto Beach fossil reef lie off East Beach, San Salvador. Several of these have a flourishing cap of Neogoniolithon that extends above low-tide level and we believe that the Neogoniolithon cap of Grotto Beach reef did likewise. Wherever found in the stratigraphic record this facies should serve to identify the subtidal/intertidal boundary. The uppermost Pleistocene beach sediments associated with Grotto Beach fossil reef lie 5.8 m above present-day mean sea level, which ist strong evidence that this portion of San Salvador has undergone little subsidence since the Grotto Beach section was deposited.     

Holotestoid: a computational model for testing hypotheses about echinoid skeleton form and growth

Regular echinoid skeletons, or tests, comprise plate patterns and overall shapes that have proven challenging to analyse solely on the basis of any one approach or process. Herein, we present a computational model, Holotestoid, that emulates four macrostructural ontogenic processes involved in test growth (plate growth, plate addition, plate interaction, and plate gapping). We devise a geometric representation for analysing tests and describe how we use analogies (bubble interactions and close-packing) to emulate the processes. In the computational model, the emulated processes are used to determine the plate size and plate shape and combined to simulate a growth zone. We simulated growth zones for Arbacia punctulata and for Strongylocentrotus franciscanus by changing the value for one parameter, the ambulacral column angle. We quantitatively compared morphological features for simulated forms to those for real specimens to test the computational model. Additionally, we simulated growth zones for A. punctulata, S. franciscanus, Eucidaris thouarsii, and Mellita quinquiesperforata by changing three parameters, ambulacral column angle, peristome radius to apical system radius ratio, and apical system radius to column length ratio. Holotestoid can be used to explain morphological disparity among echinoid tests.     

Diversity of hopanoids and squalene-hopene cyclases across a tropical land-sea gradient

Bacterial hopanoids are ubiquitous in Earth surface environments. They hold promise as environmental and ecological biomarkers, if the phylogeny and physiological drivers of hopanoid biosynthesis can be linked with the distribution of hopanoids observed across a breadth of samples. Here we survey the diversity of hopanoid cyclases from a land‐sea gradient across the island of San Salvador, in the easternmost part of the Bahamas. The distribution of lipids was determined for the same sites, for the first time overlaying quantification of bacteriohopanepolyols with sqhC phylogeny. The results are similar to previous reports: environmental sqhCs average < 65% translated amino acid identity to their closest named relatives, and sequences from putative Proteobacteria dominate. Additionally, a new and apparently ubiquitous group of marine hopanoid producers is identified; it has no identifiable close relatives. The greatest diversity of hopanoid lipids occurs in soil, but hopanoids represent a minor fraction of total soil‐derived lipids. Marine samples contain fewer identifiable hopanoids, but they are more abundant as a fraction of the total extractable lipids. In soil, the dominant compounds are 35‐aminobacteriohopane‐32,33,34‐triol and adenosylhopane. In an upper estuarine sample, bacteriohopanetetrol and 32,35‐anhydrobacteriohopanetetrol dominate; while in lower estuarine and open marine samples, the most abundant are bacteriohopanetetrol and bacteriohopaneribonolactone. Cyclitol ethers are trace components in the soil, absent in the estuary, and of moderate abundance in the open marine setting, suggesting a dominant marine source. Conversely, aminotriol and aminotetrol decrease in abundance or disappear completely from land to ocean, while 2‐methyldiplopterol shows the opposite trend. Small quantities of 2‐methylbacteriohopanepolyols are detectable in all samples. The overall hopanoid distributions may correlate to the major phylogenetic families of hopanoid producers or to the environments in which they are found.     

Comparative morphology of stingray lateral line canal and electrosensory systems

Elasmobranchs (sharks, skates, and rays) possess a variety of sensory systems including the mechanosensory lateral line and electrosensory systems, which are particularly complex with high levels of interspecific variation in batoids (skates and rays). Rays have dorsoventrally compressed, laterally expanded bodies that prevent them from seeing their mouths and more often than not, their prey. This study uses quantitative image analysis techniques to identify, quantify, and compare structural differences that may have functional consequences in the detection capabilities of three Eastern Pacific stingray species. The benthic round stingray, Urobatis halleri, pelagic stingray, Pteroplatytrygon (Dasyatis) violacea, and benthopelagic bat ray, Myliobatis californica, show significant differences in sensory morphology. Ventral lateral line canals correlate with feeding ecology and differ primarily in the proportion of pored and nonpored canals and the degree of branching complexity. Urobatis halleri shows a high proportion of nonpored canals, while P. violacea has an intermediate proportion of pored and nonpored canals with almost no secondary branching of pored canals. In contrast, M. californica has extensive and highly branched pored ventral lateral line canals that extended laterally toward the wing tips on the anterior edge of the pectoral fins. Electrosensory morphology correlates with feeding habitat and prey mobility; benthic feeders U. halleri and M. californica, have greater electrosensory pore numbers and densities than P. violacea. The percentage of the wing surface covered by these sensory systems appears to be inversely related to swimming style. These methods can be applied to a broader range of species to enable further discussion of the relationship of phylogeny, ecology, and morphology, while the results provide testable predictions of detection capabilities. J. Morphol., 2008. © 2008 Wiley‐Liss, Inc.