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Although the reproductive biology and early life-history stages of deep-sea corals are poorly understood, such data are crucial for their conservation and management. Here, we describe the timing of larval release, planula behavior, metamorphosis, settlement, and early juvenile growth of two species of deep-sea soft corals from the northwest Atlantic. Live colonies of Gersemia fruticosa maintained under flow-through laboratory conditions released 79 planulae (1.5–2.5 mm long) between April and early June 2007. Peak planulation in G. fruticosa coincided with peaks in the chlorophyll concentration and deposition rates of planktic matter. Metamorphosis and settlement occurred 3–70 d post-release. The eight primary mesenteries typically appeared within 24 h, and primary polyps grew to a height of ∼6–10 mm and a stalk diameter of ∼1 mm within 2–3 months. Planulae of Duva florida (1.5–2.5 mm long) were extracted surgically from several colonies and were successfully reared in culture. Primary polyps reached a height of ∼3–4 mm within 2–3 months. No budding of primary polyps was observed in either species over 11–13 months of monitoring, suggesting a very slow growth rate.
The endemic Hawaiian gastropod Smaragdia bryanae is a specialized marine herbivore that uses the endemic seagrass Halophila hawaiiana as both food and habitat. These small neritids, their grazing scars, and their egg capsules are found year-round on seagrass leaves, where they feed on protoplast contents released as the sharp outer-lateral teeth of the snail's radula puncture leaf epidermal cells; the contents of these cells are likely swept into the mouth by the long, wispy cusps of the marginal teeth. Structural differences from the typical neritid radula include elongated outer-lateral teeth with two sharply pointed cusps, delicate marginal teeth reduced in both size and number, and a compressed central section. Snails grazed on leaves of H. hawaiiana steadily in laboratory culture, and grew and reproduced on this diet. In laboratory choice experiments, snails did not graze the thalli of any of six macroalgal species growing near seagrass where snails were collected, and strongly preferred occupying seagrass. Seagrass samples from five field sites on Oahu and one on Maui showed from 30% to 94% of leaves damaged, with 11% of the total leaf standing area grazed. Snails are smaller (mean length 2.74±0.32 mm, mean width 2.15±0.17 mm, n = 217) than the width of the leaves of H. hawaiiana (mean 3.24±1.26 mm, n = 790). The snails associate constantly with their host, despite the scattered distribution, small patch size, and variability of the seagrass resource, demonstrated by a sevenfold range in the leaf area index (mean 1.11±0.61 cm2 blade surface cm– 2, n = 31) among samples. Damage on grazed leaves (mean 8.21±7.05 mm2 per leaf, or 16.5% of leaf surface, n = 511) is concentrated in the apical and central epithelia between the midrib and the marginal veins, where snails may access cells with thinner walls and few fibers. Details of the grazing interaction between these extant species in Hawai'i shed light on the ecological specialization of members of the genus Smaragdia to seagrasses over geological time.
A retractable head region somewhat resembling the introvert of sipunculans is a characteristic feature of members of the annelid taxon Fauveliopsidae. The morphology of fauvelopsids is not well known, and additional data might help to resolve their relationships with other annelids and sipunculans. Ultrastructural investigations of the anterior end of adults of Fauveliopsis cf. adriatica revealed peculiar brain and sensory structures. From the neuropil of the brain, two pairs of lobes mainly composed of neuronal somata extend posteriorly into the peristomium and the following segment. The nuchal organs are embedded in the median pair of lobes, as are additional photoreceptor-like sensory structures, the ocellar tubes, which are found at the bases of epidermal follicles that extend deeply into the brain. The retractor muscles of the prostomium are attached to the apices of these follicles, which are lined by tendon and supportive cells. The lumen of each follicle is completely filled with cuticular material that forms a rod. Monociliary sensory cells are present all along the length of each follicle; their cilia extend into the cuticle, and are oriented parallel to the longitudinal axis of the tube. Basally, each follicle forms an ovoid extension that is devoid of cuticular material and densely filled with numerous sensory processes—microvilli and cilia—of bipolar sensory cells. The terminal end of the 40-µm-deep follicle is formed by two conspicuous cells that contain numerous densely packed vesicles that resemble pigment granules. The ocellar tubes of fauveliopsids are strikingly similar to the ocular tubes of sipunculids. These similarities may reflect common ancestry or may represent convergent evolution; both alternatives are partially supported by previous morphological and molecular studies.
Autotomy of the elytra (scales) in the annelid Alentia gelatinosa occurs at a breakage plane near the junction between the elytron and its elytrophore (stalk), and requires fracture of the external epidermal cuticle. The mechanism of cuticular fracture was investigated by light and electron microscopy, glycoconjugate histochemistry, direct observation of autotomy in isolated preparations, and mechanical tests. The breakage plane crosses the elytrophoral wall at a cuticular thickening and passes through the subelytral cavity between the elytron and the terminal septum of the elytrophore. At the cuticular breakage zone (CBZ), the collagenous framework of the normal cuticle is replaced with non-collagenous microfibrils. The CBZ has a complex glycoconjugate composition and includes a strongly sulfated, uronic acid-containing glycosaminoglycan and a high proportion of disulfide or sulfydryl linkages. Tonofilament-rich epidermal cells (tendon cells) are attached to the thick cuticle on the dorsal and ventral sides of the CBZ. Dorsal tendon cells have long processes that extend into the elytron near the roof of the subelytral cavity. Ventral tendon cells are linked by connective tissue to the longitudinal and terminal sphincter muscles of the elytrophore. Mechanical tests showed that the elytrophoral wall is not inherently weaker at the autotomy plane than elsewhere. It is hypothesized that at autotomy (i) contractile force generated by the sphincter muscle is transmitted through elytrophoral tendon cells to the ventral side of the CBZ and (ii) contraction of the longitudinal and main circular muscles of the elytrophore increases hydrostatic pressure in its lumen, everts the terminal septum, and generates tension that is transmitted through elytral tendon cells to the dorsal side of the CBZ. This results in stress concentration at the basal edge of the CBZ and initiates fracture. The distinctive microstructure and macromolecular composition of the CBZ may reduce its fracture toughness and make it more susceptible to brittle failure.
Because of their reproductive biology, spiders are extremely promising subjects for testing hypotheses on sexual selection. Further, their genital morphology provides useful characteristics for taxonomy. However, the structure and functional morphology of the genitalia of members of most spider groups are poorly known. This is especially true for members of the Haplogynae. In this article, the female genitalia of three oonopid species are described, using light and scanning electron microscopy. The male palps are also briefly described. The female genitalia of all these three species do not correspond to the description of haplogyne genital systems given in the literature. Receptacula are lacking in the genitalia of Opopaea deserticola and Zyngoonops sp. Sperm are present in the uterus internus of members of these two species, indicating that fertilization occurs there or in the ovary. Females of Zyngoonops sp. have a pouch that possibly holds appendages on the male endites during copulation. Modifications on the endites might allow males to exert copulatory courtship. A secretory sac was found inside the receptaculum of Gamasomorpha lutzi. Previous studies on oonopids with the same type of genitalia showed that the sac contains sperm and that it can be discarded during copulation. Spermatozoa were also found inside the uterus internus of members of G. lutzi. A sclerite in the uterus wall of females of all three species might serve to lock the uterus during copulation in order to prevent sperm from getting into it, as suggested for a variety of other oonopids. The male palps of O. deserticola and Zyngoonops sp. are simple. Furrows on the emboli of G. lutzi suggest that males use the palps as copulatory courtship devices. The present study reveals the complex genital morphology of three species belonging to the little known spider family Oonopidae, and provides new insights into the function of their genitalia in the context of sexual selection.
Here we report on the first quantitative survey of morphological variation in the sea urchin Heliocidaris erythrogramma within Western Australia and distinguish between two subspecies found to co-occur in this region. We surveyed urchins at multiple spatial scales along the Western Australian coastline to assess variation in dermis and spine color and, using landmark-based geometric morphometrics, spine morphology. Both color and morphology proved to be useful for separating subspecies within Western Australia. There were four major color morphs: red dermis/violet spines (56%), red/violet-green (23%), red/ green (7%), and white/green (10%). Members of the first two color morphs had bulbous spines with wide, flattened tips, a morphology that is unique to Western Australia and characteristic of H. e. armigera, and members of the latter two consistently exhibited the narrow, pointed spines typical of specimens of H. e. erythrogramma, which has a broader distribution. In Western Australia, H. e. armigera was relatively abundant both within and among sites, but H. e. erythrogramma was found only in a few localized patches. Shifts in the relative abundance of these two subspecies occurred at fine spatial scales (<5 km), although environmental correlates of these transitions were unclear. Contrary to expectations, neither dermis color nor spine morphology varied with relative wave exposure: individuals with a red dermis or thickened spine morphology occurred at most sites regardless of exposure, and while white dermis and thinner spines only occurred at high-exposure sites, these features were not common across the majority of exposed sites. Both color morph frequencies and spine morphology remained stable within sites over the 3-year duration of this study. While the ecological significance of this morphological variation remains unclear, the consistency of the association between color and spine morphology, occurring across fine spatial scales, suggests that strong environmental or genetic factors are involved in maintaining morphological differentiation between these two subspecies.
Pachechinus bajulus is an endemic Australian sea urchin with an unusual mode of brooded larval development. We used mitochondrial and nuclear gene sequences to estimate the phylogenetic relationships among Pachechinus and other Echinometridae, including well-studied species of Heliocidaris with planktonic development. We found strong evidence for the planktotrophic species Heliocidaris tuberculata as the sister group to a clade of three closely related species in which development is known (Heliocidaris erythrogramma, P. bajulus) or suspected (Pachechinus australiae) to be lecithotrophic. Clade support values and likelihood ratio tests rejected monophyly of Heliocidaris species. The sister group to H. erythrogramma is most likely the two Pachechinus species. We resolve the paraphyly problem by reassigning the Pachechinus species to the genus Heliocidaris (the senior synonym), which has six extant species including Heliocidaris australiae and Heliocidaris bajulus. The phylogeny has potentially important implications for comparative studies of developmental morphology and genetics that have assumed a close sister-group relationship between H. erythrogramma and H. tuberculata, and highlights the need for such data from H. bajulus and other Heliocidaris species.
The effects of spatial competition among colonial marine organisms are often evident in the contact zones between colonies. These effects are especially pronounced when the interaction results in overgrowth or necrosis of one of the competitors. Ascidians, one of the dominant taxonomic groups in subtidal sessile communities, have specialized morula cells that provide a defense against microbial infections. Injuries resulting from interspecific competitive interactions might also act as a stimulus for this defensive mechanism. Therefore, we expected to see the recruitment of morula cells in tissues near competitor contact zones. To test the hypothesis that spatial competition elicits this immune response, we placed colonies of the ascidian Didemnum perlucidum from southeastern Brazil in four different types of competitive situations: (1) overgrowth of the competitor, (2) stand-off interactions, (3) overgrowth by the competitor, and (4) free of competitors. Our results indicate that competitive interactions increase the population of morula cells in contact zones, as more cells were observed in interactions that resulted in the overgrowth of individuals of D. perlucidum, and fewer cells were observed in colonies that were free of competitors. We identified the defensive function of the morula cells by showing the presence of the enzyme phenoloxidase within its vacuoles. Phenoloxidase is a widespread enzyme among animals and plants, and is frequently used in defense by synthesizing toxic quinones from polyphenol substrates. This is the first study to document the presence of morula cells in didemnid ascidians and the mobilization of these cells by spatial competition by heterospecifics, and one of the first studies to identify phenoloxidase activity in morula cells.
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