The influence of feeding on behavior and nematocyst discharge of the sea anemone Calliactis tricolor

A decrease in basitrichous isorhiza and spirocyst nematocysts is observed in a fully fed sea anemone, Calliactis tricolor, as compared to unfed animals. Discharge of tentacle nematocysts of an intact living animal is inhibited by anesthetics and a decrease in temperature. The chemical (food origin) threshold for discharge of basitrichs is higher than for spirocysts and the implications of these results are discussed. The most probable cause for the observed decrease in nematocyst discharge is a combination of food present in the gastrovascular cavity and a physical stretching of the cavity.     

The gnathobasic spine microstructure of recent and Silurian chelicerates and the Cambrian artiopodan Sidneyia: Functional and evolutionary implications

Gnathobasic spines are located on the protopodal segments of the appendages of various euarthropod taxa, notably chelicerates. Although they are used to crush shells and masticate soft food items, the microstructure of these spines are relatively poorly known in both extant and extinct forms. Here we compare the gnathobasic spine microstructures of the Silurian eurypterid Eurypterus tetragonophthalmus from Estonia and the Cambrian artiopodan Sidneyiainexpectans from Canada with those of the Recent xiphosuran chelicerate Limulus polyphemus to infer potential variations in functional morphology through time. The thickened fibrous exocuticle in L. polyphemus spine tips enables effective prey mastication and shell crushing, while also reducing pressure on nerve endings that fill the spine cavities. The spine cuticle of E. tetragonophthalmus has a laminate structure and lacks the fibrous layers seen in L. polyphemus spines, suggesting that E. tetragonophthalmus may not have been capable of crushing thick shells, but a durophagous habit cannot be precluded. Conversely, the cuticle of S. inexpectans spines has a similar fibrous microstructure to L. polyphemus, suggesting that S. inexpectans was a competent shell crusher. This conclusion is consistent with specimens showing preserved gut contents containing various shelly fragments. The shape and arrangement of the gnathobasic spines is similar for both L. polyphemus and S. inexpectans, with stouter spines in the posterior cephalothoracic or trunk appendages, respectively. This differentiation indicates that crushing occurs posteriorly, while the gnathobases on anterior appendages continue mastication and push food towards and into the mouth. The results of recent phylogenetic analyses that considered both modern and fossil euarthropod clades show that xiphosurans and eurypterids are united as crown-group euchelicerates, with S. inexpectans placed within more basal artiopodan clades. These relationships suggest that gnathobases with thickened fibrous exocuticle, if not homoplasious, may be plesiomorphic for chelicerates and deeper relatives within Arachnomorpha. This study shows that the gnathobasic spine microstructure best adapted for durophagy has remained remarkably constant since the Cambrian.     

Spines and epibionts as antipredator defenses in the thorny oyster Spondylus americanus Hermann

Observations and experiments on the cemented pelecypod Spondylus americanus Hermann at Discovery Bay, Jamaica were made to identify the predators of this animal and learn whether their spines, or their epibionts, are important in deterring predation. Spine growth was studied also and found to be very rapid (≈ 1mm/day), and broken spines broken were repaired at this rate.The predators of Spondylus include the gastropod Fasciolaria tulipa Linnaeus (responsible for half of the observed predation), and various crushing predators, including the spiny lobster [Panulirus argus (Latreille)], rays [Aetobatis narinari (Euphrasen) and Dasyatus americanus Hildebrand], the porcupine fish (Diodon hystrix L.), and an unidentified stomatopod. Predators show an increased frequency of attacks in shallow water versus deep, and probably account for the scarcity of Spondylus americanus above −15 m.The concealment effect of epifauna was found to be more important than spines as an antipredator defense. Removal of spines and epibionts significantly increased levels of mortality only when both features were manipulated. There was no statistically significant increase in predation following the removal of epibionts because the shells were rapidly grown over by sponges. Spines alone do not appear to deter predators, but probably serve to attract epibionts and extend them beyond the more vulnerable shell margin. These results do not support the view that spondylid spines evolved as structural deterrents to predators. I argue that they most likely evolved to attract other organisms to settle on the shell and conceal it from predators.     

The morphology and ecology of Dipolydora armata (Polychaeta, Spionidae) from the western Mediterranean Sea

Bick. A. 2001. The morphology and ecology of Dipolydora armata (Polychaeta, Spionidae) from the western Mediterranean Sea. — Acta Zoologica (Stockholm) 82 : 177–187
The cosmopolitan spionid Dipolydora armata is a dominant species of epifauna on Thais haemastoma shells in the western Mediterranean Sea. Some characters, particularly number, thickness and arrangement of notopodial spines, were extremely variable among specimens. Three morphological forms were distinguished based on the general appearance of the bundles of spines; one form possesses at least one bundle of stout spines, whereas bundles of medium spines or thinner acicular spines characterise the other forms. Differences in prostomial shape, as well as in the extension and form of the caruncle, were artefacts (preservation or angle of observation). The present study supports D. rogeri as a junior synonym of D. armata. D. armata was regularly found on shells inhabited by hermit crabs ( Calcinus tubularis , Clibanarius erythropus ) and on damaged, empty shells. Abundance, dominance, size frequency, occurrence of larvae and occurrence of regenerated specimens varied, possibly as a result of fouling strategy and recurrent settlements.     

Prey preferences in captivity of the freshwater crab Potamonautes lirrangensis from Lake Malawi with special emphasis on molluscivory

Freshwater crabs play an important role for the diversification of shell morphologies in freshwater gastropods. For example, the radiation of the freshwater crab genus Platythelphusa in Lake Tanganyika is thought to have driven shell diversification of the lake’s snail fauna, promoting the evolution of thalassoid shells. No comparable thalassoid snails are known from Lake Malawi. Accordingly, it was hypothesized that the lake’s only freshwater crab, Potamonautes lirrangensis, is not a snail predator. We tested this hypothesis using feeding experiments with specimens caught in the southern part of Lake Malawi. Individual crabs were held in experimental containers offshore and were presented with various food items overnight, after which ingestion frequency was recorded. Potamonautes lirrangensis can be characterized as a scavenger that is opportunistically carnivorous. A preference for fish and snail flesh could be observed, indicating a bias toward carnivory. We observed occasional cracking of the shell in different snail species, with frequent ingestion of artificially crushed specimens, suggesting that crabs do attempt to feed on snails. However, the investigated Lake Malawi gastropods appear to be partly protected against crab predation through thick-walled and low-spired shells (especially Lanistes and Bellamya), obviating the evolution of thalassoid shells carrying rims, ridges, or spines.     

Nematocysts and Taxonomy in Laomedea, Gonothyraea and Obelia (Hydrozoa, Campanulariidae)

The cnidoms of Laomedea flexuosa, Gonothyraea loveni, Obelia geniculata, O. longissima and O. dichotoma were studied by light and scanning electron microscopy to find out whether the nematocysts could be used as taxonomic characters. Three b-rhabdoid heteroneme nematocysts (microbasic b-mastigophore) and three isorhizous haploneme nematocysts (atrichous isorhiza) were distinguished. A small b-rhabdoid nematocyst with spindle-shaped capsule occurred in all the species examined. In the polyp and planula of G. loveni , and the planula of L. flexuosa it was the only nematocyst present. Specific for L. flexuosa was a b-rhabdoid with curved capsule. In the polyp and newly-liberated medusa of O. longissima another b-rhabdoid appeared with bean-shaped capsule and markedly long spines at the distal tube. The polyp and newly-liberated medusa of O. dichotoma were characterized by two different isorhizas. A special type of isorhiza occurred in the polyp of O. geniculata . The curved capsules of the different isorhizas varied somewhat in shape and size. Differences in nematocyst structure and occurrence are presumed to provide characters for taxonomy. Thus, O. dichotoma and O. Iongissima are regarded as separate species due to their distinctly different nematocysts, in conjunction with other morphological and ecological differences.     

Novel cnidocysts of narcomedusae and a medusivorous ctenophore, and confirmation of kleptocnidism

Cnidocysts have been examined from the tentacles of the ctenophore Haeckelia rubra (Euchlora rubra) and five species of hydrozoan narcomedusae (Solmundella bitentaculata, Aegina citrea, Solmissus marshalli, Solmissus albescens, and Cunina sp.) using TEM, both in sections and by firing whole cnidocysts onto EM grids. The study revealed that these apotrichous isorhiza cnidocysts have a novel morphology in which the intracapsular inverted tubule has five circumferential pleats when viewed in transverse section, rather than the usual three pleats. Accordingly, the definition of helicoptychoneme cnidocysts has been broadened to include both the usual three-pleated cnidocysts and these new five-pleated cnidocysts. In general, apotrichous isorhizas have subspherical capsules with a thick, bilayered wall, whose interior is nearly filled with the regularly coiled, helically folded, five-pleated inverted tubule. Upon discharge, the everted tubule is several mm long and the five circumferential pleats become manifested as five helical rows of spines running up the tubule, which has three morphologically different segments. The very short basal segment is devoid of ornamentation; the remaining proximal portion is characterized by five spirals of uniform, closely packed short spines; the long distal portion is characterized by a single spiral of regularly spaced large spines that derive from all five spirals-the five spirals are otherwise demarcated in the distal portion by 'scales' that are visible only with the electron microscope.     

Effects of shell stress on the growth of hermit crabs

Growth rates of the hermit crabs Pagurus longicarpus Say and P. pollicaris Say maintained in preferred shells and in shells smaller than the preferred size have been compared. Changes in wet weight, shield length, and duration of the molt cycle were determined. Crabs in preferred shells grew significantly faster than those in small shells. P. pollicaris molted at approximately the same rate in both cases but grew more each molt in preferred shells. Von Bertalanffy growth curves were fitted to changes in shield length among crabs in preferred shells. These curves indicate that P. longicarpus may mature four months after settling from the plankton and reach its asymptotic size within the next eight months while P. pollicaris also matures four months after leaving the plankton but does not reach its asymptotic size for approximately three years. The rapid growth of P. longicarpus may enable it to preempt shells which are required for the successful brooding of a large clutch before these shells are required by more aggressive competitors, such as P. pollicaris and Clibanarius vittatus.     

Nematocysts of Rhopilema nomadica (Scyphozoa: Rhizostomeae), an immigrant jellyfish in the eastern mediterranean

Rhopilema nomadica—a recently discovered scyphomedusa in the eastern Mediterranean—is considered a lessepsian migrant. Its nematocysts were extracted from the scapular and mouth-arm tentacles and examined using light and electron microscopy techniques. The morphometric parameters of the nematocysts were measured before and after complete discharge. Three categories of nematocysts were identified: heterotrichous isorhiza haploneme, holotrichous isorhiza haploneme, and heterotrichous microbasic eurytele. The relative abundance of the nematocysts and their occurrence in tissues of the jellyfish were noted. A brief discussion concerning the classification of certain types of nematocysts is given. A comparison with the available data on other Rhopilema species revealed that the nematocyst categories of R. nomadica are more similar to those of the Atlantic R. verrilli than to those of the Western Pacific R. esculentum. A brief comparison of the injuries caused by these species is given. © 1995 Wiley-Liss, Inc.     

Organelle survival in a foreign organism: Hydra nematocysts in the flatworm Microstomum lineare

Nematocysts are characteristic organelles of the phylum cnidaria. They are designated kleptocnidae when sequestered in animals that feed on cnidaria. Kleptocnidae are known for more than a century. Nevertheless it is still enigmatic how selected nematocyst types survive in the predator and how they reach their final destination in the foreign body. In the free-living Platyhelminth Microstomum lineare the fate of nematocysts of the prey Hydra oligactis was analyzed at the ultrastructural level and by fluorescence microscopy using hydra polyps that had been stained in vivo with the fluorescent dyes TROMI and TRITC. M. lineare digested hydra tissue in its intestine within 30?min and all nematocyst types were phagocytosed without adherent cytoplasm by intestinal cnidophagocytes. Desmoneme and isorhiza nematocysts were digested whereas cnidophagocytes containing the venom-loaded stenotele nematocysts started to migrate out of the intestinal epithelia through the parenchyma to the epidermis thereby traversing the subintestinal and subepidermal muscle layer. Within one to two days, M. lineare began to form a muscle layer basolateral around epidermal cnidophagocytes. Epidermal stenoteles survived in M. lineare for at least four weeks. The ability of epidermal stenotele nematocysts to discharge suggest that this hydra organelle preserved its physiological properties in the new host.     

A complete styginid trilobite from the Ordovician of Sweden

Styginidae are a small group of trilobite species, which are usually rare and most of which are incompletely known. Here we describe a complete specimen of Raymondaspis grandigena n.sp. from the Middle Ordovician (Darriwilian, upper Arenig) of Sweden. Among the group it has an unusual combination of exceptionally large genal spines, a thorax with notably short pleural spines in the anterior tergites, and a wide concave pygidial margin. The hypostome is documented for the first time in situ in a styginid, and its attachment can be best described as semi-impendent.     

Age, growth rate and season of recruitment of Pinna nobilis (L) in the Croatian Adriatic determined from Mg:Ca and Sr:Ca shell profiles

Stable oxygen isotope analyses of U-shaped spines removed at intervals along profiles of the outer shell surface of Pinna nobilis (L) were used to reconstruct sea surface temperature (SST) and validate the periodicity of adductor muscle scar rings on the inner shell surface. Elemental ratios (Mg:Ca and Sr:Ca) of spines, determined using Inductively Coupled Plasma-Atomic Emission Spectrometry (ICP-AES), were compared with the SST estimated from the stable oxygen isotopic composition of the shell deposited at the same time. The elemental ratios and the stable oxygen isotopic composition recorded in the shell were significantly correlated: Mg:Ca ratio=0.0002 (seawater temperature)+0.0095 (r²=0.445), Sr:Ca ratio=0.00005 (seawater temperature)+0.0014 (r²=0.887). The ratios in the spines were highest when the SST was warmest during July and August and were lowest between January and February when the SST was minimal.The positions of the first and second adductor muscle scar rings, unlike the later rings, are often difficult to discern, and in large shells they are obscured by nacre. Seasonal patterns in the elemental ratios were used to characterise the age in regions of the outer shell surface corresponding to the first two years of shell growth. A combination of both elemental ratios and muscle scar rings were used to estimate the age and hence the growth rate of P. nobilis from three locations in the Croatian Adriatic. Annual cycles of shell growth, inferred from the seasonal pattern in the element ratios, were used to determine the season of recruitment of fan mussels at several localities along the Croatian coastline. Pinnids generally settled during late autumn and winter although one shell from Mali Ston Bay settled during the summer. P. nobilis from Mali Ston Bay exhibited the fastest growth reaching a length of ∼60 cm and an age of 9 years, whereas those from Malo jezero grew the slowest attaining a length of ∼50 cm at 12 years of age.     

Sclerotised spines in the female bursa associated with male’s spermatophore production in cantharidin-producing false blister beetles

Cantharidin is a defence chemical synthesised in only two beetle families Meloidae and Oedemeridae. In Meloidae, cantharidin is used as a defence chemical in eggs. However, in Oedemeridae the function of cantharidin remains unclear. Based on morphological comparison of female internal reproductive organs in 39 species of Oedemeridae, we found that some species have sclerotised spines in the bursa copulatrix (bursal spines), while others have no such spines. Molecular phylogenetic trees inferred from mitochondrial 16S and nuclear 28S rRNA gene sequences suggested multiple evolutionary origins of bursal spines from an ancestor without spines. In the species which lacked spines, males transferred small amounts of ejaculates to females; however, in species with spines, males transferred large spermatophores. Deposited spermatophores gradually disappeared in the bursa, probably owing to absorption. To compare the amounts of cantharidin in eggs laid by species with and without bursal spines, we constructed a new bioassay system using the small beetle Mecynotarsus tenuipes from the family Anthicidae. M. tenuipes individuals were attracted to droplets of cantharidin/acetone solution, and the level of attraction increased with cantharidin concentration. This bioassay demonstrated that the eggs of Nacerdes caudata and N. katoi, both of which species have conspicuous bursal spines, contain more cantharidin than the eggs of N. waterhousei, which lacks spines. In the former species, males transfer large spermatophores to the female, and spermatophores are eventually broken down and digested within the female’s spiny bursa. Thus, females with bursal spines may be able to provide more cantharidin to their eggs.     

Sex-specific differences and the role of predation in the interaction between the hermit crab, Pagurus longicarpus, and its epibiont, Hydractinia symbiolongicarpus

The interaction between the hermit crab, Pagurus longicarpus, and the shell epibiont, Hydractinia symbiolongicarpus, varies from mutualism to parasitism based on the environmental context. We tested the hypothesis that this interaction also varies as a function of hermit crab sex. Given that recent work showed a negative effect of Hydractinia on female reproduction, we predicted a greater frequency of males in Hydractinia shells in the field and a stronger preference by males than females for shells with Hydractinia. Field collections documented a significantly greater proportion of males than females occupying shells with Hydractinia, and a significantly greater proportion of ovigerous females than non-ovigerous females in shells with Hydractinia. In laboratory shell-switching experiments, a greater proportion of males than females chose to enter shells with Hydractinia, but there was no difference in the proportions of males and females that vacated shells with Hydractinia.We examined whether the presence of Hydractinia influenced predation rates. Blue crabs fed on more than twice as many hermit crabs in shells with Hydractinia as compared to bare shells, but there was no significant difference for stone crabs. Laboratory experiments showed that the force required to crush shells was significantly greater for shells without Hydractinia. Thus, the lower occupancy and preference exhibited by females than males for shells with Hydractinia appears to result both from the decreased reproduction shown in past studies and an increase in predation risk.     

Mechanical defensive adaptations of three Mediterranean sea urchin species

In the Mediterranean, Paracentrotus lividus and Sphaerechinus granularis are important drivers of benthic ecosystems, often coexisting in sublittoral communities. However, the introduction of the invasive diadematoid Diadema setosum, which utilizes venomous spines, may affect these communities. To describe the mechanical properties of the test and spines of these three species, specimens were collected in winter of 2019 from the sublittoral zone of the Dodecanese island complex, southeastern Aegean Sea. This region serves as a gateway for invasive species to the Mediterranean Sea. Crushing test was conducted on live individuals, while 3‐point bending test was used to estimate spine stiffness. Porosity and mineralogy of the test and spine, thickness of the test, and breaking length of the spine were measured and compared, while the microstructural architecture was also determined. The test of S. granularis was the most robust (194.35 ± 59.59 N), while the spines of D. setosum (4.76 ± 2.13 GPa) exhibited highest flexibility. Increased porosity and thickness of the test were related to increased robustness, whereas increased flexibility of the spine was attributed to high porosity, indicating that porosity in the skeleton plays a key role in preventing fracture. The spines of S. granularis exhibited highest length after fracture % (71.54 ± 5.5%). D. setosum exhibited higher values of Mg concentration in the test (10%) compared with the spines (4%). For the first time, the mineralogy of an invasive species is compared with its native counterpart, while a comparison of the mechanical properties of different species of the same ecosystem also takes place. This study highlights different ways, in which sea urchins utilize their skeleton and showcases the ecological significance of these adaptations, one of which is the different ways of utilization of the skeleton for defensive purposes, while the other is the ability of D. setosum to decrease the Mg % of its skeleton degrading its mechanical properties, without compromising its defense, by depending on venomous bearing spines. This enables this species to occupy not only tropical habitats, where it is indigenous, but also temperate like the eastern Mediterranean, which it has recently invaded.     

Effect of pH on the morphology and viability of Scrippsiella trochoidea cysts in the hypoxic zone of a eutrophied area

To investigate the cause of morphological change of Scrippsiella trochoidea cysts and its ecological significance in the hypoxic zone of a eutrophied area, the effect of pH on the morphology and viability of S. trochoidea cysts was studied. In the acidification experiment, the dissolution of calcareous spines of S. trochoidea cysts was observed at less than pH 7.39, indicating that the morphological change of S. trochoidea cysts is caused by the low pH levels in acidic sediments of hypoxic zone. After being exposed to intense acidic environments, cysts of S. trochoidea without calcareous spines were able to germinate; however, they seem to be easily linked to degradation in the sediments. These results suggest that the survival of S. trochoidea cysts is being threatened by environmental conditions in the hypoxic zone of eutrophied area.     

Structural features of neural spines of the caudal vertebrae of protoceratopoids (Ornithischia: Neoceratopsia)

The structure of caudal neural spines of protoceratopoids displays adaptation for aquatic and terrestrial mode of life. The increasing height of caudal neural spines in the series Leptoceratops, Udanoceratops, Protoceratops, Bagaceratops is connected with the extent of adaptation for swimming and changes in inclination of neural spines are connected with the mechanical balance of the lever. Thus, the anterior caudal vertebrae (1cd–15cd) of Protoceratops and Bagaceratops show an anticliny, which promotes extension (rise) of a heavy tail in terrestrial conditions. In the middle part of the tail (16cd–23cd), with the greatest height of neural spines, a decrease in width and increase in thickness counteract transverse loads accompanying movements on land. At the same time, the supraspinal ligament prevents divergence of neural spines caused by curvature of the tail as it is raised above the ground.     

Geographical variation in shell shape of the pod razor shell Ensis siliqua (Bivalvia: Pharidae)

The present study assessed the existence of variation in the shell shape of the pod razor shell (Ensis siliqua) throughout its distributional range in the north-eastern Atlantic. Shells of E. siliqua caught at seven collecting sites (three in Portugal, three in Spain and one in Ireland) were studied by geometric morphometric methods, using both landmark- and contour-based methods. Both approaches (landmarks inside the valves and shell outline) discriminated the shells from Aveiro (centre of Portugal) and Strangford Lough (Ireland) from those caught in the nearby localities (remaining Portuguese and Spanish sites, maximum distance of 550 km by sea). Landmark analysis revealed that shells from Aveiro were more similar to shells from Ireland (~1,500 km far away). Contour analysis revealed that shells from Aveiro had a shape with a comparatively larger height-to-width ratio, whereas shells from Ireland showed a slightly more curved outline than in the remaining sites. Landmark- and contour-based methods provided coherent complementary information, confirming the usefulness of geometric morphometric analyses for discerning differences in shell shape among populations of E. siliqua. A brief review of previous applications of geometric morphometric methods to modern bivalve species is also provided.     

Reproductive costs of the symbiotic hydroid Hydractinia symbiolongicarpus (Buss and Yund) to its host hermit crab Pagurus longicarpus (Say)

Studies on the interaction between the hermit crab Pagurus longicarpus and its symbiotic hydroid Hydractinia symbiolongicarpus have focused on positive effects of hydroids on their host hermit crabs (e.g., protection from predators). Yet, these benefits may be balanced with reproductive costs, which are rarely studied. Results from field observations, laboratory trials, and a mesocosm experiment indicate that female hermit crabs in hydroid-colonized shells exhibit depressed ovigery, smaller clutch sizes, and increased clutch failure relative to females in bare shells. Frequent switching between bare and hydroid-colonized shells may alleviate negative effects when the density of hydroids in the environment is low, but at high densities Hydractinia may significantly impact hermit crab reproduction.     

Cuticular structures of the stomodeum in Paulinia acuminata (De Geer) and Marellia remipes Uvarov (Orthoptera : Pauliniidae)

The cuticular lining of the stomodeum in orthopteroid insects usually bears cuticular structures, consisting of ridges, teeth, and spines, which are sometimes fairly complex. In some of these insects, particularly cockroaches and Orthoptera s. str., some of these structures have systematic value, mainly at the higher systematic levels. The cuticular lining of the stomodeum in Paulinia acuminata (De Geer) and Marellia remipes (Uvarov) (Orthoptera : Pauliniidae) (the only 2 genera included within the family) have been studied. Wide differences in the pattern of the stomodeal armature, and also in the shape, size and distribution of the spines in its different parts, are recorded. The proportions and limits of the crop and proventriculus, and the form and spatial distribution of spines and other characters, relative to the ridges, appear to be of taxonomic significance. They indicate that the 2 genera may not be closely related, as has been suggested, based on other evidence. While Marelliaappears to be related to the Acrididae, Pauliniaseems to be isolated from other acridoids, except for certain features in common with Proscopiidae and Eumastacidae.