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Intraspecific variation in characters such as arm sucker and gill lamellae counts in octopodids is yet to be thoroughly investigated, potentially hampering our ability to recognize species. In this study, data from 13 specimens of Muusoctopus hydrothermalis collected at four hydrothermal vents on the East Pacific Rise between 8°38′N and 12°48′N are considered. Although the northern and southern octopuses differ minimally in size, mean sucker count by arm in the northern group is 11.7 to 22.8% higher than it is in the southern group; in addition these octopuses typically have an additional gill lamella and bulkier funnel organs. The arms of each individual carry a different number of suckers. The difference is significant on nonadjacent arms, a pattern that merits examination in a broader taxonomic context. Why these differences exist among conspecifics remains unknown, the incidence of parasitic copepods is not different between the groups and the between-group variation in arm suckers seen here compares well with a previous report of variation among 18 specimens from the type locality. Increases in meristic characters (counts) in fish are attributed to lower temperatures during embryonic development following Jordan's rule. Northern and southern vents offer the octopuses a wide temperature range, but vent fluid chemistry differs. Northern vent fluids may be more toxic; if so, developing octopus embryos may survive only minimal vent fluid exposure and therefore develop at low temperatures. At the less toxic southern vents, eggs may survive greater exposure to vent fluids and thus develop at higher temperatures.
Deep-sea organisms are generally considered to be restricted to the great ocean depths, unable to enter shallower habitats except perhaps at the isothermic polar latitudes, due to physiological constraints, or more intense biotic interactions. Wood-boring bivalves of Xylophaga, however, are here shown to be exceptional. A clade, formally recognized by Turner in 2002, united by the derived characters of a complex mesoplax and a truncated excurrent siphon, papillose incurrent siphon with a longitudinal dorsal trough bordered by lappets, and often carrying white or glass-like granules, occurs at significantly shallower and warmer depths than do other species of the genus. Here, description of the new species X. multichela from 106 m depth off the Pacific coast of Guatemala illustrates the characters that unite the clade. Because wood-boring bivalves face intense competition and high levels of predation at depths of 2200 to 3250 m, they may not perceive biotic interactions at shallower depths to be dramatically more intense. In addition, members of this group are hypothesized to tolerate reduced oxygen availability, a requisite if animals are to occur in warm waters.
A feeding frenzy of 12 octopuses of Vulcanoctopus hydrothermalis was observed from the manned submersible ‘Alvin’ at Parigo, a 2620 m deep hydrothermal vent on the East Pacific Rise. The aggregated benthic octopuses at the active vent used their arms and webs to forage on bathypelagic amphipods, apparently targeting their attacks based on contact with the swarming amphipods. Individual octopuses wrapped their arms around the mantles of smaller octopuses, apparently in competition for prey. Although members of the prey species, Halice hesmonectes, are individually small (<5 mm long), the density of their swarms may make them attractive prey for the octopus. Inactive sulphide spires encircled part of this vent site; octopuses that climbed these spires had easy access to the dense prey swarms. The presence of the spires may uniquely enable this site to support simultaneous foraging by large numbers of octopuses.
The most familiar examples of erectile tissue are the genitalia of mammals, notably the penis and clitoris. Among the soft-bodied invertebrates, erectile tissue is virtually unknown, even in sex organs. Here we report that the ligula, the copulatory organ that is a modification of an arm tip of male octopuses, is erectile in Octopus bimaculoides. The normally minute ligula was observed during an unsuccessful mating attempt to be engorged. Histological sections of the ligula reveal striking structural convergences with mammalian erectile tissue: large, well-vascularized internal cavities subdivided by networks of collagen fibres and enclosed by an array of collagen fibres. This internal structure differs markedly from the dense, three-dimensional array of muscles and connective tissues seen in the other octopodid ligulae examined. Erectile tissue may represent an evolutionary compromise between opposing selective forces. Small ligulae may be advantageous because O. bimaculoides hunts in daylight and the white-faced ligula may be conspicuous to predators. Large ligulae, however, may be advantageous if the intense sexual selection thought to occur among octopodids selects for large ligulae, which can transfer larger spermatophores that carry more sperm. An erectile ligula may minimize the impact of these opposing selective forces.
The relationship between sperm reservoir and total spermatophore length among 168 spermatophores from 44 species in 11 genera has been considered. Bivariate plots show that four Atlantic species of the genus Eledone produce spermatophores with relatively large sperm reservoirs that differ from all others. Measurements of the remaining spermatophores are so tightly correlated that a single equation explains over 96% of the variation. Functional constraints on gross spermatophore morphology may be so strong that males cannot manipulate sperm reservoir size independently of spermatophore size to maximize the sperm delivered at a single copulation. Alternate means to assure male paternity may have evolved in the group as a result. Despite the overall uniformity of the relationship among all species other than those of Eledone in the Atlantic, these measurements distinguish the overtly similar species Octopus bimaculatus–Octopus bimaculoides and separate Benthoctopus januarii from all others.
The gut contents of a female specimen of Graneledone cf. boreopacifica collected from the caldera wall of Axial Volcano, near an active hydrothermal vent in the Northeast Pacific Ocean are reported. At least 30 individual gastropods and 46 individual polychaetes are represented in the gut contents by hard parts. Shell fragments and shells removed from the gut allow ready identification of the gastropods Provanna variabilis and Lepetodrilus fucensis, both of which are known only from North Pacific hydrothermal vents. Jaws of polychaete worms are identified as those of the nereidid, Nereis piscesae, and the predatory polynoids, Levensteiniella kincaidi and an unidentified species in the subfamily Branchinotogluminae. Not only was a considerable volume of prey hard parts ingested, the gastropod shells had been crushed before being ingested. The large size of the beaks in this genus of octopus and the increased area they offer for insertion of the superior mandible muscle, the prime mover in beak closure, support the hypothesis that these beaks exert sufficient force to crush the gastropod shells. Although cephalopods had been reported to be absent from hydrothermal vents, the data presented here demonstrate that not only do they occur in vent habitats, they actively prey on vent fauna.
A synthesis of current research suggests several trends in deep sea taphonomy. First, epibiosis (the condition of having bionts on exoskeletons that is not obligate) appears to decrease with increasing depth. Epibionts, such as serpulid polychaetes and barnacles that take advantage of ephemeral hard substrates offered by crab exoskeletons, sea urchin spines or gastropod shells, appear to be more diverse and common at depths of less than 500 m. At abyssal depths, microbial communities on living shells are common and may contribute to the breakdown of molluscan periostracum. These communities may also have patchy distributions.
Second, shell-inhabiting symbionts, such as sea anemones uniquely associated with shells of pagurid crustaceans, are most common from 300 to 700 m. These unusual sea anemones cover hermit crab-inhabited gastropod shells with a chitinous material (Dunn et al. 1980) that may affect shell longevity. While gastropod shells without a periostracum may degrade quickly in calcium-poor waters, the chitinous covering secreted by the anemone may increase the postmortem longevity of the shell.
Third, deep sea gastropods usually have eroded apical whorls but buttress this weakness with internal calcium plugs (J. McLean, pers. comm., 1992). In addition, not all deep sea gastropods, like buccinids, have thick periostracal layers. How differing periostracal thicknesses affect taphonomy is undetermined.
Finally, deep sea gastropod fossils occur throughout the Cenozoic from many geographic localities: Eocene (Oregon, United States), Upper Oligocene (France), Miocene (New Zealand, Japan), Miocene/Pliocene (Australia), and Pleistocene (Italy). Sedimentary deposits containing these gastropods vary from thin interbedded sediments within oceanic basalt (e.g., Eocene, Oregon), volcanic breccia with bathyal sediments (e.g. Miocene, Japan), turbiditic sequences (Neogene, Japan) and bathyal clays (Pleistocene, Italy). Fossil quality varies from poorly preserved internal molds with altered shell material, to excellently preserved specimens with calcareous layers intact. We will discuss how this record bears on the geographic and preservational differences among the various deep sea gastropod fauna.
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