Aboral Surface Research Articles

Covering behaviour in the sea urchin Pseudechinus magellanicus (Temnopleuridae) in central Patagonia, Argentina

ABSTRACT The sea urchin Pseudechinus magellancus exhibits a covering behaviour through which it captures different items from the environment and arranges them on its aboral surface with the assistance of podia. The items mainly used include detached algae and inorganic items available in the substrate. Spatial (between sites) and vertical (along a coastal depth gradient from intertidal tidepools to upper circalittoral zones) variability was found in the composition of covering items, indicating that coverage is strongly influenced by the availability of the utilised items. Inorganic biogenic items (eg shell remnants) were prevalent in the lower intertidal, while detached algae predominate in subtidal environments. A positive selection of larger items from those available in the environment was found at least in the intertidal tidepools. The species exhibited an equal degree of coverage between juveniles and adults, and also a higher proportion of individuals with high coverage in the intertidal zone compared to the subtidal zone. The degree of coverage between sites with dissimilar sediment input was similar. The covering behaviour in P. magellanicus may not be a response to a single factor but rather an advantageous trait with multiple functions that could have been selected simultaneously.

Effect of flow structures on the transport of gametes in broadcast-spawning sea urchin

Broadcasted fertilization for the reproduction of invertebrates is accomplished through a complicated interaction between spawned gametes and the surrounding flows. Most gametes encounter each other in the vicinity of the sea urchin body where unique flow structures develop, and so analysis of local flow characteristics allows us to better understand the effect of flow on the fertilization process. This study applied a Lagrangian framework based on computational fluid dynamics to estimate the fertilization rate of eggs in a range of flow velocities (0.025–0.2 m/s) and the fertilization rate was the highest at U = 0.1 m/s, which is an intermediate flow speed. Among the four classified sub-zones, such as the aboral surface, wake, substrate, and water column, fertilization occurred most frequently in the wake and substrate regions. The relationship between fertilization rate and flow structures was investigated using three parameters: 1) standardized Morisita index to quantify the pattern of gamete distribution, 2) length of the recirculation zone to specify the areas where the eggs are most frequently fertilized, and 3) integral scale to estimate the dimension of vortex structures downstream. The results of this study show that the fertilization rate is higher inside the recirculation zone, especially when strong vortex structures are observed because the flow provides a favorable condition for the gametes to aggregate and collide with each other.

Description of two species of the genus Astrodia Verrill, 1899 (Ophiuroidea, Euryalida, Asteronychidae), including a new species from seamounts in the West Pacific.

Five specimens of Ophiuroidea from deep-sea seamounts in the West Pacific were collected and identified as two species, Astrodiaduospina sp. nov. and Astrodiaabyssicola. The new species, Astrodiaduospina sp. nov., can be distinguished from its congeners by having indistinct or underdeveloped oral papillae, relatively short genital slits, crescent-shaped lateral arm plates, and plate-shaped external ossicles on the aboral surface of the disc. One specimen was identified as Astrodiaabyssicola, which has been reported in the north-western Pacific and the north-eastern coast of Japan. The most recent tabular key of Astrodia was revised with two more key characteristics added, the shape and presence of oral papillae and the number of arm spines. The phylogenetic relationship of Astrodia and Asteronyx was analyzed based on 16S and COI sequences. The discovery of the two species further expanded the geographical distribution of the genus Astrodia.

Born With Bristles: New Insights on the Kölliker’s Organs of Octopus Skin

The entire skin surface of octopus embryos, hatchlings and juveniles bears scattered tufts of tiny chitinous setae within small pockets, from which they can be everted and retracted. Known as Kölliker’s organs (KO), they disappear before the subadult stage. The function of these structures during the early life of the octopus is unknown, despite having been first described nearly two centuries ago. To investigate these organs further, general trends in size of KO distribution and density were analyzed in hatchlings and juveniles of 17 octopod species from all oceans, representing holobenthic, holopelagic and meropelagic modes of life. The size of the KO is fairly constant across species, unrelated to mode of life or hatchling size. The density of KO is similar on ventral and dorsal body surfaces, but hatchlings of smaller size tend to have a higher density of KO on the aboral surface of the arms. Analysis of a series of post-hatchingOctopus vulgarisshows KO size to be constant throughout ontogeny; it is therefore a consistent structure during the octopus early life from planktonic hatchling to benthic juvenile. New KO are generated on the skin of the arm tips during the planktonic period and initial benthic lives of juveniles. Their density, on both the mantle and arms, gradually decreases as the octopus grows. In older benthic juveniles, the KO degrades, losing its setae and the base of its follicle becomes exposed as a nearly circular stump of muscle. It is estimated that fully everted KO increase the body surface area by around two-thirds compared to when the KO are retracted. This modular mechanism of body surface extension and roughness probably influences flow-related forces such as drag and propulsion of the moving surface of the young octopus while it is of small size with a relatively large surface area. In addition, the distribution of these organs on the aboral surface of the arms of the octopus and their birefringent properties suggest a role in camouflage. Further research is needed to test these hypotheses of KO function in live animals.

Morphology of the lips, cheeks, and the hard palate of the Egyptian water buffalo (Bubalus bubalis): a focus on histological, histochemical, and ultrastructural aspects.

The available data is scanty about Egyptian water buffalo lips, cheeks, and palate. The current investigation was focused on describing the morphology of the lip, cheek, and palate. Our study included the gross, light, and electron microscopic examinations of ten heads of the Egyptian water buffalos. The nasolabial plate surface carried numerous scales of keratinized epithelium. Internal labial surface and labial mucocutaneous junctions were covered with stratified squamous keratinized epithelium. Two types of hair follicles in the dermis included ordinary and cavernous types characterized by cavernous space. The conical papillae on the internal aspect of the oral commissure were projected from the mucous membrane. Seromucous glands were occasionally observed under the oral mucous membrane of the commissure and gave positive PAS and AB. Conical papillae density on the inner cheek surface had some variations: the rostral part had large papillae, while the dorsal part had numerous papillae than the ventral part, the caudal part had a smaller number of papillae, while the middle part was devoid of papillae. Parotid duct opening in the buccal vestibule was without papillae. Conical papillae had two surfaces; the rostral surface was highly keratinized than the caudal one. The buccal gland was a compound tubuloacinar mixed (mucoserous) gland and mucus acini only reacted to PAS and AB. The oral surface of palatine rugae was covered with highly keratinized epithelium than the aboral surface. Palatine glands showed PAS and AB positive. The result describes the relationship between the available food particles, environmental conditions and the lip, cheek, and palate appearance, and structure.

Growth and mineralogy in dental plates of the holocephalan Harriotta raleighana (Chondrichthyes): novel dentine and conserved patterning combine to create a unique chondrichthyan dentition

The dentition in extant holocephalans (Chondrichthyes) comprises three pairs of continuously growing dental plates, rather than the separate teeth characterizing elasmobranchs. We investigated how different types of dentine in these plates, including hypermineralized dentine, are arranged, and how this is renewed aborally, in adult and juvenile dentitions of Harriotta raleighana (Rhinochimeridae). Individual plates were analysed using x-ray computed tomography (μCT), scanning electron microscopy (SEM) in back scattered mode with energy dispersive X-ray (EDX) analysis, and optical microscopy on hard tissue sections.ResultsHarriotta dental plates are made entirely of dentine tissue, mostly as trabecular dentine, bone itself being absent. Hypermineralized dentine forms in restricted ovoid and tritor spaces within trabecular dentine, inside a shell of outer and inner dentine layers. Trabecular dentine is ubiquitous but changes to sclerotic osteodentine near the oral surface by increasing density, remaining less mineralized than the hypermineralized dentine. All structures are renewed aborally, within a vascular dental pulp, a tissue suggested to be a source of stem cells for tissue renewal. Ca density profiles and concentrations of Mg, P, and Ca ions reveal extreme differences in the level and type of mineralization. Early mineralization in ovoids and tritors has very high levels of Mg, then a sudden increase in mineralization to a high total mineral content, whereas there is gradual change in trabecular dentine, remaining at a low level.Hypermineralized dentine fills the prepatterned ovoid, rod and tritor spaces, early at the aboral surface within the trabecular dentine. Deposition of the hypermineralized dentine (HD, proposed as new specific name, whitlockin replacing pleromin) is from surfaces that are lined with large specialized odontoblasts, (whitloblasts, instead of pleromoblasts) within cell body spaces connecting with extensive, ramifying tubules. Early mineralization occurs amongst this maze of tubules that penetrate far into the dentine, expanding into a mass of saccules and membranous bodies, dominating in the absence of other organic matrix. This early stage has hydroxyapatite, also significantly rich in Mg, initiated as a poorly crystalline phase. In the hypermineralized dentine, formation occurs as clusters of variably shaped crystals, arising from a sudden phase transition.ConclusionsIn the hypermineralized dentine, high MgO + CaO + P2O5 suggests that almost pure Mg containing tricalciumphosphate (MgTCP: (ß-Ca3(PO4)2) (whitlockite) is present, with little or no hydroxyapatite. Serial replacement of tritors and ovoids is suggested to occur within the dental plate, probably representing a relic of patterning, as classically found in elasmobranch dentitions.

A new species of Fibularia from Japanese waters with a redescription of F. japonica and F. ovulum (Echinodermata: Echinoidea: Clypeasteroida).

A new species, Fibularia coffea sp. nov., occurs from shallow waters in Japan. This new species is distinguished from the other species of Fibularia by the following characters: test height is low, oral surface is slightly depressed toward the peristome, number of pores of petal III continues to increase with the test growth, reaching over 30 at TL > 7.5 mm, and black pigments form symmetric pentaradial on aboral surface in living animals. Two further Japanese species, Fibularia japonica and F. ovulum, are redescribed based on the type specimens (F. japonica) and additional specimens (F. ovulum), respectively. A tabular key to the extant species of Fibularia is also provided. A partial fragment of the mitochondrial gene cytochrome oxidase subunit I (COI) of the type specimens of F. coffea sp. nov. and the additional specimen of F. japonica was sequenced for barcoding in future works.

Cephalotrichella echinicola, sp. nov. (Palaeonemertea : Cephalotrichellidae), a new nemertean associated with sea urchins from Nha Trang Bay (South China Sea)

About 50 nemertean species have been reported to live in symbiotic relationships with other invertebrates, but only two hoplonemertean species are associated with echinoderms (starfish). The palaeonemertean Cephalotrichella echinicola, sp. nov. is described from samples collected in Nha Trang Bay, Vietnam, South China Sea. The species is the first known nemertean associated with sea urchins (Metalia sternalis and M. spatagus), living on both the oral and the aboral surfaces of the host and freely moving among its spines. The internal morphology of the new species is described based on histological sections and confocal laser scanning microscopy with phalloidin and antibody labelling. Sequences of three nuclear gene markers (18S rRNA, 28S rRNA, and histone H3) and two mitochondrial gene markers (16S rRNA and COI) were compared with those of other palaeonemertean species, and a phylogenetic analysis suggested that C. echinicola is closest to the free-living Cephalotrichella alba Gibson & Sundberg, 1992. Both the morphological data and the phylogenetic analysis provide additional evidence for distinguishing the families Cephalotrichidae and Cephalotrichellidae and support the rejection of Cephalotrichella as a junior synonym of Cephalothrix. A new diagnosis of the genus Cephalotrichella is given. http://zoobank.org/urn:lsid:zoobank.org:pub:091B5D56-71B2-4F4C-9AD8-F666B4610DE2

Whole-body photoreceptor networks are independent of 'lenses' in brittle stars.

Photoreception and vision are fundamental aspects of animal sensory biology and ecology, but important gaps remain in our understanding of these processes in many species. The colour-changing brittle star Ophiocoma wendtii is iconic in vision research, speculatively possessing a unique whole-body visual system that incorporates information from nerve bundles underlying thousands of crystalline 'microlenses'. The hypothesis that these might form a sophisticated compound eye-like system regulated by chromatophores has been extensively reiterated, with investigations into biomimetic optics and similar supposedly 'visual' structures in living and fossil taxa. However, no photoreceptors or visual behaviours have ever been identified. We present the first evidence of photoreceptor networks in three Ophiocoma species, both with and without microlenses and colour-changing behaviour. High-resolution microscopy, immunohistochemistry and synchrotron tomography demonstrate that putative photoreceptors cover the animals' oral, lateral and aboral surfaces, but are absent at the hypothesized focal points of the microlenses. The structural optics of these crystal 'lenses' are an exaptation and do not fulfil any apparent visual role. This contradicts previous studies, yet the photoreceptor network in Ophiocoma appears even more widespread than previously anticipated, both taxonomically and anatomically.

The first Western Gondwanan species of Triodus Jordan 1849: A new Xenacanthiformes (Chondrichthyes) from the late Paleozoic of Southern Brazil

Triodus is a well-known genus of Xenacanthiformes, previously recorded from the late Bashkirian (Lower Pennsylvanian, Carboniferous) to the middle Artinskian (Cisuralian, Permian), mainly from Laurasian deposits (Europe and USA). For the first time, this genus is recorded from the Western Gondwana, based on isolated teeth that are referred to Triodus richterae sp. nov. The new species were found associated with other shark teeth (another xenacanthiforms and a possible euselachian), palaeoniscoid teeth and scales, labyrinthodont teeth, tetrapod bony remains, macroscopic charcoal and leaf fragments. This fossil assemblage was collected in a conglomerate layer from the Barro Alto site (São Gabriel municipality, Rio Grande do Sul State, Brazil), with an estimated Capitanian age, from the Morro Pelado Member, Rio do Rasto Formation, Paraná Basin. The new species has teeth with an almost oval base and the aboral surface has a smooth concavity and a rounded to horseshoe-shaped basal tubercle. The coronal surface of these teeth has a tricuspid crown, a rhomboid-shaped coronal button with rounded edges with a lingually directed shaft and some oral foramina predominantly situated at the lingual margin of the base and flanking the lingual shaft. The lateral cusps bear a variable number of non-branching vertical cristae, distributed from the apex to their proximal portion, making the transversal section of these cusps asterisk-shaped. Microstructurally, these teeth have both base and cusps composed of orthodentine with an opened pulp cavity. T. richterae sp. nov. represents the youngest species of Triodus, considering it comes from Capitanian (late Guadalupian) beds, which are at least 15 Ma younger than T. kraetschmeri, the previously youngest species of this genus. The depositional interpretation, as well as the fossil assemblage in the type locality of the new species and of the Rio do Rasto Formation as a whole, indicates another freshwater record for xenacanthid sharks.

Covering behavior of deep-water echinoids in Antarctica: possible response to predatory king crabs

Covering behavior refers to the propensity of echinoids (Echinoidea) to lift materials from the surrounding environment onto their aboral surfaces using their tube feet and spines. This behavior has been widely documented in regular echinoids from a variety of well-lit, shallow- marine habitats. Covering behavior in the deep sea, however, is rarely observed, and the functional significance of covering when it does occur remains speculative. During a photographic survey of the seafloor off Anvers Island and Marguerite Bay along the western Antarctic Pen - insula, we imaged 11 benthic transects at depths ranging from 390 to 2100 m. We recorded the number of echinoid species, incidence of covering behavior, types of materials used for covering, potential predators of echinoids, and potential prey items for predators. The echinoid Sterechinus spp. was found at all depths, and the percentage of individuals exhibiting covering behavior increased with depth between 390 and 1500 m. There was a significant positive correlation between the incidence of covering behavior in Sterechinus spp. and the density of king crabs (Anomura: Lithodidae), crushing predators that may be expanding their bathymetric range up the Antarctic continental slope as a consequence of ongoing climatic warming. In contrast, covering behavior was not positively correlated with the densities of non-crab predators, the total densities of predators, or the availability of prey. Our results document rarely observed covering behavior in echinoids living in the deep sea and suggest that covering could be a behavioral response to predation pressure by king crabs.

Holistic morphometric analysis of growth of the sand dollar Echinarachnius parma (Echinodermata:Echinoidea:Clypeasteroida).

Holistic morphometrics is a term implying complete shape characterization of all of the structural parts of an organism. The skeleton of an echinoid is comprised of hundreds of individual plates arranged in a closed 3-dimensional mosaic forming the test. GIS software and techniques were used to generate topologically correct digital models of an ontogenetic series of specimens of the sand dollar echinoid Echinarachnius parma. Plate growth can be considered in proportion to overall skeleton growth, resulting in a linear model of relative growth. Alternatively, separate logistic equations can be fit to the ontogenetic series of homologous plate areas using nonlinear least squares regression to result in a model for instantaneous growth. The linear and logistic parameters of the models describe the allometric growth of plates from different viewpoints. Growth is shown to fall into characteristic patterns defining distinct plate growth domains associated with development of the imago (larval) skeleton just prior to metamorphosis, early growth associated with expansion of the corona and fold-over (forming the flattened body form), juvenile growth and formation of petals, and adult growth. Functions of growth, plate translocation, plate juxtaposition between aboral and oral surfaces, and relationships with internal buttressing are quantified. Results offer explanations for general skeletal symmetry, distinction between ambulacral and interambulacral growth, the relationship of growth to internal buttressing, existence of a distinct petalodium, and anterior-posterior asymmetry during development. The parametric values of growth functions derived from the results are a basis for computational modeling of growth and development in sand dollars.

DRILL HOLES IN THE IRREGULAR ECHINOID,FIBULARIA, FROM THE OLIGOCENE OF NEW ZEALAND

Abstract A detailed analysis of 813 tests of the irregular echinoid Fibularia sp. collected from multiple depositional horizons at a single Oligocene locality on South Island, New Zealand revealed 57 specimens with drill holes, representing a 7% drilling frequency (57 drilled individuals). Among those drilled, 38% (22 individuals) had multiple drill holes with up to four holes present. Most holes were complete, circular to sub-circular in outline, averaging 0.5 mm in diameter, and belong to the ichnospecies Sedilichnus simplex. The distribution of holes on the test was non-random: 1) the aboral surface was drilled preferentially, and 2) the central region of the aboral surface had a significantly higher number of holes (p 0.1). The distribution of drill holes as well as their shape and size make it likely that they were produced by a predatory g...

The Cambrian edrioasteroid Stromatocystites (Echinodermata): Systematics, palaeogeography, and palaeoecology

The Cambrian edrioasteroid Stromatocystites is reported and described from Spain, Sweden and Turkey. All previously known occurrences of the genus are critically reviewed, and S. flexibilis is reinterpreted as a junior synonym of S. pentangularis. Stromatocystites was biogeographically widespread and colonized different areas of Baltica, Gondwana (Arabian, eastern and western margins) and Laurentia (western Newfoundland). Stratigraphically, it ranges from Cambrian Series 2, Stage 4 to Cambrian Series 3, Drumian. Stromatocystites lived in quiet water environments with stabilized substrates. It was attached directly to the substrate by its aboral surface. As these environments were widespread throughout Baltica, Gondwana and Laurentia, availability of suitable substrates for larval settlement and oceanic palaeocurrents led to the successful development of Stromatocystites colonies.

Siphuncular structure in the Jurassic belemnitid Megateuthis (Cephalopoda: Coleoidea)

Each septum of the phragmocone in the Jurassic belemnitid Magateuthis is composed of a thick, lamello-fibrillar, nacreous layer and a thin, chitinous layer, here termed the additional chitinous layer, which covers the aboral surface of the septum. The nacreous and additional chitinous layers continue from the septum into the septal neck. The inner surface of the septal neck is covered by a thick, tubular, conchiolin layer that originates on the septal surface around the siphuncle. The connecting ring is composed of three conchiolin layers with high elasticity: the additional chitinous layer that fuses with the tubular chitinous layer and continues into the connecting ring, and an outer chitinous layer that is a non-calcified continuation of the nacreous layer of the septal neck and is often destroyed by diagenesis. As seen in higher magnification, the tubular layer consists of tightly packed tubes with chitinous walls that are oriented at right angles to the long axis of the siphuncle. At about half of its length, each tube divides into three branches that are oriented towards the siphuncular cavity. The siphuncular cavity contains diaphragms composed of flat calcareous sheets. The siphuncular structure in belemnitid Megateuthis is similar to that in aulacocerid Mojsisovicsteuthis? In extant Spirula, the inner layer of the septal neck is composed of calcareous pillars embedded in a chitinous substance, whereas, in Megateuthis, it consists of chitinous tubes. The chitinous tubular layer in Megateuthis continues into the connecting ring.

Relative effects of gamete compatibility and hydrodynamics on fertilization in the green sea urchin Strongylocentrotus droebachiensis.

Intraspecific variation in gamete compatibility among male/female pairs causes variation in the concentration of sperm required to achieve equivalent fertilization levels. Gamete compatibility is therefore potentially an important factor controlling mating success. Many broadcast-spawning marine invertebrates, however, also live in a dynamic environment where hydrodynamic conditions can affect the concentration of sperm reaching eggs during spawning. Thus flow conditions may moderate the effects of gamete compatibility on fertilization. Using the green sea urchin Strongylocentrotus droebachiensis as a model system, we assessed the relative effects of gamete compatibility (the concentration of sperm required to fertilize 50% of the eggs in specific male/female pairs; F50) and the root-mean-square of total velocity (urms; 0.01-0.11 m s(-1)) on fertilization in four locations near a spawning female (water column, wake eddy, substratum, and aboral surface) in both unidirectional and oscillatory flows. Percent fertilization decreased significantly with increasing urms at all locations and both flow regimes. However, although gamete compatibility varied by almost 1.5 orders of magnitude, it was not a significant predictor of fertilization for most combinations of position and flow. The notable exception was a significant effect of gamete compatibility on fertilization on the aboral surface under unidirectional flow. Our results suggest that selection on variation in gamete compatibility may be strongest in eggs fertilized on the aboral surface of sea urchins and that hydrodynamic conditions may add environmental noise to selection outcomes.

A Tremadocian asterozoan from Tasmania and a late Llandovery edrioasteroid from Victoria

Jell, P.A., 2014. A Tremadocian asterozoan from Tasmania and a late Llandovery edrioasteroid from Victoria. Alcheringa 38, 528–540. ISSN 0311-5518.An asterozoan, Maydena roadsidensis gen. et sp. nov., is described from the mid-Tremadocian (La1b Zone) Florentine Valley Formation in southwestern Tasmania and is the oldest known asterozoan in the world. Although only a single, largely dissociated, specimen is available, enough is preserved to recognize distinctive ambulacral plates similar to those of Archegonaster Jaekel from the Llanvirn of the Czech Republic. Reciprocodiscus transambus n. gen., n. sp. is an isorophid edrioasteroid from the uppermost late Llandovery Springfield Formation exposed in the bed of Deep Creek, near Springfield, 65 km north northwest of Melbourne. It occurs with a low-diversity trilobite fauna indicating a deepwater, subphotic environment. This edrioasteroid has in each ambulacrum a single series of floor plates that are not visible on the oral surface, indicating its isorophid affinity and retains certain apparently primitive features that are not seen in post-Cambrian edrioasteroids, such as the very large plates of the marginal circlet, plated aboral surface and ambulacral tips extending onto the marginal circlet plates.Peter A. Jell [p.jell@uq.edu.au] School of Earth Sciences, The University of Queensland, St Lucia Queensland 4072, Australia.

A new Xenacanthiformes shark (Chondrichthyes, Elasmobranchii) from the Late Paleozoic Rio do Rasto Formation (Paraná Basin), Southern Brazil

The Brazilian records on Xenacanthiformes include teeth and cephalic spines from the Parnaíba, Amazonas and Paraná basins. This work describes a new species of Xenacanthidae, collected in an outcrop of Serrinha Member of Rio do Rasto Formation (Wordian to Wuchiapingian), Paraná Basin, municipality of Jacarezinho, State of Paraná. The teeth of the new species are two or three-cuspidated and the aboral surface show a smooth concavity and one rounded basal tubercle. The coronal surface presents one semi-spherical and subcircular coronal button, and also two lateral main cusps and one central (when present) with less than one fifth of the size of the lateral cusps in the labial portion. The lateral cusps are asymmetric or symmetric, rounded in transversal section, lanceolate in longitudinal section, devoid of lateral carinae and lateral serrations, and with few smooth cristae of enameloid. In optical microscope the teeth show a trabecular dentine (osteodentine) base, while the cusps are composed by orthodentine, and the pulp cavities are non-obliterated by trabecular dentine. The fossil assemblage in the same stratigraphical level and in the whole Rio do Rasto Formation indicates another freshwater record for xenacanthid sharks.

Fertilization in a sea urchin is not only a water column process: effects of water flow on fertilization near a spawing female

Fertilization efficiency in free-spawning invertebrates in the marine environment depends on the complex interaction between biological and physical factors. Experimental evi- dence indicates that in some taxa, a considerable amount of fertilization may take place on the substrate and within flow structures in close proximity to a spawning female. Gametes can be retained on a spawning animal, resulting in their slow release over relatively long periods of time, and retained eggs can be fertilized before they are released into the water column. Hydrodynamic conditions are likely to influence both the retention of gametes and their subsequent mixing in the water column as well as the relative importance of fertilization in the water column and in other locations near spawning animals. Here, fertilization in the broadcast-spawning sea urchin Strongy - locentrotus droebachiensis was explored over a range of flow velocities (u- = 2 to 15 cm s �1 ) to determine the effect of velocity on fertilization and the relative contribution of different locations (aboral surface of the female, water column, wake, and the substrate behind the female) to overall fertilization. As velocity increased, the percentage of eggs fertilized declined in all locations. At all velocities, more eggs were fertilized on the aboral surface than in the water column. Further, as velocity increased, the relative contribution of the aboral surface to overall fertilization increased. These results highlight the importance of considering the interaction of hydrodynamics, organism morphology, and gamete properties in studies of fertilization in broadcast-spawning invertebrates.

Effects of Oscillatory Flow on Fertilization in the Green Sea Urchin Strongylocentrotus droebachiensis

Broadcast spawning invertebrates that live in shallow, high-energy coastal habitats are subjected to oscillatory water motion that creates unsteady flow fields above the surface of animals. The frequency of the oscillatory fluctuations is driven by the wave period, which will influence the stability of local flow structures and may affect fertilization processes. Using an oscillatory water tunnel, we quantified the percentage of eggs fertilized on or near spawning green sea urchins, Strongylocentrotus droebachiensis. Eggs were sampled in the water column, wake eddy, substratum and aboral surface under a range of different periods (T = 4.5 – 12.7 s) and velocities of oscillatory flow. The root-mean-square wave velocity (rms(u w)) was a good predictor of fertilization in oscillatory flow, although the root-mean-square of total velocity (rms(u)), which incorporates all the components of flow (current, wave and turbulence), also provided significant predictions. The percentage of eggs fertilized varied between 50 – 85% at low flows (rms(u w) <0.02 m s−1), depending on the location sampled, but declined to below 10% for most locations at higher rms(u w). The water column was an important location for fertilization with a relative contribution greater than that of the aboral surface, especially at medium and high rms(u w) categories. We conclude that gametes can be successfully fertilized on or near the parent under a range of oscillatory flow conditions.