The life cycle of Eutima commensalis sp. nov. (Eutimidae,Hydromedusae)

Developmental stages of the medusa Eutima commensalis sp. nov. were obtained from plankton collections made at 2 fixed stations, one at Fareway Buoy and the other at Aroor in the Cochin backwater system. Six stages were arbitrarily assigned and described. The hydroid and the medusa of this form were found to occur in these waters throughout the year, except during the monsoon months from June to September. The hydroid is endocommensalic on wood-boring molluscs such as Nausitora hedleyi, Teredo furcifera and Martesia striata.     

The life cycle ofClytia attenuata (calyptoblastea: Campanulariidae)

The life cycle of the hydroidClytia attenuata (Calkins) (Calyptoblastea: Campanulariidae) has been completed in the laboratory including development of the medusa, previously described asPhialidium lomae Torrey (Leptomedusae: Campanulariidae). Under laboratory conditions, the hydroid exhibits some morphological variation. Characteristic branching of the hydroid occurs at temperatures between 17° to 19°C. At 13° to 15°C the colonies are unbranched and cannot be distinguished fromClytia cylindrica L. Agassiz. Young medusae are similar to other young species ofPhialidium. Development to the adult form requires 25 to 30 days at 17° to 24°C. The adult medusae are 6 to 10 mm in diameter, watch-glass shaped, and have 20 to 28 tentacles. Based on the adult medusa,Clytia attenuata is maintained as a valid species.     

Latitudinal trends in size-dependence of bioluminescence in the midshipman fish Porichthys notatus

The bioluminescent fish Porichthys notatus (plainfin midshipman), has a discontinuous distribution along the Pacific coast of North America. The fish is present from Cape Mendocino southward to Baja California, Mexico, absent off the coast of Oregon, USA, and abundant, northward, in Puget Sound, Washington. Interestingly, the population in Puget Sound lacks the substrate (luciferin) necessary for the luminescence reaction and, despite possessing an otherwise fully functional photophore system, is nonluminescent. The California population of P. notatus is uniformly luminescent south of Monterey Bay, but 15% of the speciments tested from San Francisco Bay and the Gulf of the Farallons have been reported to be nonluminescent. Explanations for nonluminescent midshipman in both Puget Sound and the San Francisco Bay area have focussed on a dietary requirement for luciferin. To gain further insight into reasons for nonluminescence in the San Francisco Bay region, the distribution of bioluminescence in P. notatus was studied from Monterey Bay to Cape Mendocino during 1985. A complex pattern of bioluminescence was found, in which nonluminescent individuals reflected neither a local anomaly in the San Francisco Bay region nor a simple gradient of decreasing luminescence towards the northern end of the range of the California population. Instead, a distinct size-dependent component in luminescence capability of the fish was observed. Aspects of the life history of P. notatus and related factors which might influence the bioluminescence characteristics of this population are discussed.Please address all correspondence to Dr. F.I. Tsuji at the Osaka Bioscience Institute     

Invertebrate introductions in marine habitats: two species of hydromedusae (Cnidaria) native to the Black Sea,Maeotias inexspectata and Blackfordia virginica,invade San Francisco Bay

The hydrozoans Maeotias inexspectata Ostroumoff, 1896 and Blackfordia virginica Mayer, 1910, believed to be native to the Black Sea (i.e. Sarmatic) and resident in a variety of estuarine habitats worldwide, were found as introduced species in the Petaluma River and Napa River, California, in 1992 and 1993. These rivers are mostly-estuarine tributaries that flow into north San Francisco Bay. Both species appeared to be well-established in this brackishwater habitat. Salinities at the collection sites were about 11 during the summer, rising to nearly 20 in the early autumn and falling to near 0 in the winter. Large numbers of all sizes of both species of medusae were observed and collected, indicating that the hydroid stages of the life cycles of the two are also well-established in these rivers. In the Petaluma River, populations of both species were at maximum in late July, with numbers of individuals declining through August and into September; the Napa River was sampled only in October, and at that time only B. virginica was found. Examination of full guts of M. inexspectata and B. virginica medusae revealed that both species had fed nearly exclusively on small crustaceans, principally barnacle nauplii, copepods and their eggs and nauplii, and crab zoea larvae (M. inexspectata only). All the M. inexspectata medusae were males, indicating that the population has probably developed from the introduction of perhaps only a single male polyp or polyp bud. In spite of its inability to reproduce sexually, this population appears to be maintained by the prodigious ability of the polyp to bud and reproduce asexually, and is fully capable of invading additional low-salinity habitats from its present Petaluma River site. Male and female B. virginica medusae were collected in both the Petaluma River and the Napa River, indicating that B. virginica may have been introduced by either the polyp or medusa stage (or both), but that multiple individuals (of both sexes) must have arrived from another port in one or more invasions. As indicated for M. inexspectata, the B. virginica population will also probably seed new populations in San Francisco Bay and elsewhere. Based on its cnidome as well as the morphology of both medusa and polyp, M. inexspectata has been reclassified by moving it from the family Olindiidae, Limnomedusae, to the family Moerisiidae, Anthomedusae.     

Distribution of male and female mtDNA lineages in populations of blue mussels,Mytilus trossulus and M. galloprovincialis,along the Pacific coast of North America

Geographic variation in mitochondrial large subunit (16S) ribosomal RNA haplotypes was examined for blue mussels, Mytilus trossulus Gould, 1850 and M. galloprovincialis Lamarck, 1819, sampled from ten sites along the Pacific coast of the USA in January of 1993. Using polymerase chain reaction/restriction fragment length polymorphism (PCR/RFLP) assays we determined haplotype frequencies for both the male and female mussel mitochondrial DNA (mtDNA) lineages. Populations from Morro Bay south to San Diego, California, contained only M. galloprovincialis male and female haplotypes, while those from Arcata Bay, California, north to Port Orford, Oregon, were fixed for M. trossulus haplotypes. Populations from Monterey Bay to Bodega Bay, California, contained a mixture of M. trossulus and M. galloprovincialis haplotypes. Overall only 2 of 97 heteroplasmic individuals had a mixed M. trossulus/M. galloprovincialis mitochondrial genotype indicating that hybridization is uncommon in the populations sampled. Further, there was no evidence of extensive introgression between these mussel taxa at the level of mtDNA. This is in contrast to previously published results which suggested the significant introgression of M. trossulus haplotypes into southern populations containing primarily M. galloprovincialis nuclear genotypes. We feel the discrepancy lies in the ability of our assays to detect haplotypes corresponding to both the male and female mtDNA lineages. Potential explanations for the lack of mtDNA introgression include, low levels of backcrossing between hybrids and parental taxa, epistatic interaction between nuclear and mitochondrial genes and the breakdown of a sex-specific inheritance pattern for mtDNA in hybrids.     

A commensal relationship between the scyphozoan medusae<Emphasis Type="Italic"> Catostylus mosaicus</Emphasis> and the copepod<Emphasis Type="Italic"> Paramacrochiron maximum</Emphasis>

The copepod Paramacrochiron maximum was found in high numbers (up to 5,675 copepods/medusa) on the oral arms of the scyphozoan Catostylus mosaicus. This association was considered to be commensalism for the following reasons: P. maximum (Lichomolgidae) was abundant on the medusae (approximately 805 copepods/kg of medusae) and very rare in the water column (approximately 5.99×10^-4 copepods/kg of water); copepodites and adults of the symbiont were present on the host; the copepods were on the medusae both day and night, at different times (nine occasions between March 1999 and May 2000) and different locations (Botany Bay and Lake Illawarra, NSW, Australia). Over 40 taxa of plankton were found on the oral arms of C. mosaicus (including protists, cnidarians, polychaetes, molluscs, a wide range of holoplanktonic and meroplanktonic crustaceans, chaetognaths and fish eggs). These taxa were abundant in the water column and we concluded that they were prey. Symbiotic amphipods and carangid fishes were found with medusae. We conclude that there is a symbiotic association between P. maximum and C. mosaicus and care should be taken not to confound these copepods with the prey of C. mosaicus. Poecilostomid copepods are well known for consuming mucus and feeding is likely to be a major reason for the association.Communicated by G.F. Humphrey, Sydney     

Allozyme and morphological evidence for a newly introduced species of Aurelia in San Francisco Bay,California

The jellyfish Aurelia aurita (Linnaeus, 1758) is usually considered to be a cosmopolitan species. Aurelia sp. medusae observed at Foster City, San Francisco Bay, California, USA, since 1988 are morphologically distinct from Aurelia sp. collected 200 km away in Monterey Bay, but are morphologically similar to aquarium-cultured Aurelia sp. from Japan. They differ consistently in radial canal morphology. In allozyme electrophoresis, significant differences at 12 of 14 polymorphic loci strongly suggest that Aurelia sp. from Foster City and Tokyo Bay belong to one species, while Aurelia sp. from Monterey Bay and Vancouver Island belong to a second species. We propose that Aurelia sp. at Foster City is a recent introduction, possibly from Japan via ships' ballast water. The identities and taxonomic affinities of the two Aurelia defined in this study, and their relationships with the Linnaen A. aurita described from the North Atlantic, will require genetic and morphological study of the currently recognized species A. aurita and A. limbata (Brandt, 1838) from several zoogeographical provinces.     

Genetic evidence for the cryptic species pair, <Emphasis Type="Italic">Lottia digitalis </Emphasis>and <Emphasis Type="Italic">Lottia austrodigitalis</Emphasis> and microhabitat partitioning in sympatry

It has been proposed that the common West Coast limpet, Lottia digitalis, is actually the northern counterpart of a cryptic species duo including, Lottia austrodigitalis. Allele frequency differences between southern and northern populations at two polymorphic enzyme loci provided the basis for this claim. Due to lack of further evidence, L. austrodigitalis is still largely unrecognized in the literature. Seven additional enzyme loci were examined from populations in proposed zones of allopatry and sympatry to determine the existence of L. austrodigitalis as a sibling species to L. digitalis. Significant allele frequency differences were found at five enzyme loci between populations in Laguna Beach, southern California, and Bodega Bay, northern California; strongly supporting the existence of separate species. Both species exhibit two microhabitat morphotypes, a gooseneck barnacle morph in the mid-intertidal zone and a rock morph in the high-intertidal zone. In sympatry, L. austrodigitalis was more abundant higher in the intertidal on rocks, whereas L. digitalis was more abundant lower in the intertidal on barnacles. This finding supports earlier claims of microhabitat partitioning in this sibling species pair. In addition to this finding, the transition zone between the species was found to have shifted substantially northward in only two decades, from Monterey Peninsula, CA to near Pigeon Point, CA, where L. digitalis previously dominated.     

Feeding behavior of the ctenophore <Emphasis Type="Italic">Thalassocalyce inconstans</Emphasis>: revision of anatomy of the order Thalassocalycida

Behavioral observations using a remotely operated vehicle (ROV) in the Gulf of California in March, 2003, provided insights into the vertical distribution, feeding and anatomy of the rare and delicate ctenophore Thalassocalyce inconstans. Additional archived ROV video records from the Monterey Bay Aquarium Research Institute of 288 sightings of T. inconstans and 2,437 individual observations of euphausiids in the Gulf of California and Monterey Canyon between 1989 and 2005 were examined to determine ctenophore and euphausiid prey depth distributions with respect to temperature and dissolved oxygen concentration [dO]. In the Gulf of California most ctenophores (96.9%) were above 350 m, the top of the oxygen minimum layer. In Monterey Canyon the ctenophores were more widely distributed throughout the water column, including the hypoxic zone, to depths as great as 3,500 m. Computer-aided behavioral analysis of two video records of the capture of euphausiids by T. inconstans showed that the ctenophore contracted its bell almost instantly (0.5 s), transforming its flattened, hemispherical resting shape into a closed bi-lobed globe in which seawater and prey were engulfed. Euphausiids entrapped within the globe displayed a previously undescribed escape response for krill (‘probing behavior’), in which they hovered and gently probed the inner surfaces of the globe with antennae without stimulating further contraction by the ctenophore. Such rapid bell contraction could be effected only by a peripheral sphincter muscle even though the presence of circumferential ring musculature was unknown for the Phylum Ctenophora. Thereafter, several live T. inconstans were collected by hand off Barbados and microscopic observations confirmed that assumption.     

Ecological gradients and relative abundance of native (Mytilus trossulus) and invasive (Mytilus galloprovincialis) blue mussels in the California hybrid zone

Marine communities are experiencing unprecedented rates of species homogenization due to the increasing success of invasive species, but little is known about the mechanisms that allow a species to invade and persist in a new habitat. In central California, native (Mytilus trossulus Gould 1850) and invasive (Mytilus galloprovincialis Lamarck 1819) blue mussels and their hybrids co-exist, providing an opportunity to analyze the mechanisms that determine the distributions of these taxa. Spatial and temporal variation in temperature and salinity and the relative frequencies of these mussel taxa were examined between 2000 and 2004 at four sites in San Francisco Bay and four in Monterey Bay, which were chosen for their different positions along inferred estuarine/oceanic gradients in the hybrid zone. Mussels were genetically identified as the parent species or hybrids by amplifying regions of two species-specific loci: the adhesive byssal thread protein (Glu-5′) and the internal transcribed spacer region of ribosomal DNA (ITS 1). The proportion of M. trossulus at the eight hybrid zone sites correlated negatively with average salinity (R ²=0.60) and positively with maximal temperature (R ²≥0.72), a somewhat unexpected result given what is known about the phylogeography of this species. The proportion of M. galloprovincialis showed the opposite pattern. The proportion of hybrids was correlated neither with habitat temperature nor salinity. Genotypes of mussel populations at an additional 13 sites from Coos Bay, Oregon (latitude 43.35°N) to Long Beach, California (latitude 33.72°N), sampled at various intervals between 2000 and 2004, were also determined. This survey confirmed previous reports that the hybrid zone lies between Monterey and the Cape Mendocino region (latitudes 36.63°N–40.5°N). Within Monterey and San Francisco Bays, however, the temporal comparisons (1990s vs. 2000s) revealed abrupt changes in the proportions of the two parent species and their hybrids on annual and decadal time scales. These changes indicate that the blue mussel populations are in a highly dynamic state. The survey also showed that, regardless of habitat, M. trossulus is consistently of smaller average size than either M. galloprovincialis or hybrids.

---

A comparison of absorption and assimilation efficiencies between four species of shallow- and deep-living fishes

We captured two species of deep-sea zoarcids, Melanostigma pammelas and Lycodapus mandibularis, from Monterey Bay California and maintained them in the laboratory. One shallow-water zoarcid, Eucryphycus californicus, and an ecologically and morphologically similar stichaeid fish Xiphister atropurpureus were collected from intertidal and subtidal habitats in Monterey Bay. We investigated the absorption and assimilation efficiencies of these fishes to determine whether deep-sea species have evolved mechanisms to increase their efficiency of food use. Fishes were placed in experimental chambers and fed a known quantity of food. Ammonia excretion was measured and feces were collected daily. Both food and feces were analyzed for water, protein, lipid and ash to determine specific absorption efficiencies. Absorption ranged from 87.7 to 92.4% and assimilation efficiencies from 84.0 to 86.5%. Meal sizes from 0.5 to 4.0% of body weight did not affect the results. No significant differences were found between deep-sea and shallow-water species fed single meals or fed ad libitum for 10 days. This suggests that the selective pressure to maximize absorption and assimilation is universal and is not affected by the productivity of the habitat occupied. However, the relative size of the intestine in the deep-sea species was significantly smaller suggesting that they had a lower metabolic cost to maintain their digestive apparatus. It could not be concluded whether this was the result of pressure to conserve energy or rather the tendency of the shallow-living species to ingest more refractory material (i.e. sediment, algae).     

Occurrence and detrimental effects of the bivalve-inhabiting hydroid <Emphasis Type="Italic">Eutima japonica</Emphasis> on juveniles of the Japanese scallop <Emphasis Type="Italic">Mizuhopecten yessoensis </Emphasis>in Funka Bay,Japan: relationship to juvenile massive mortality in 2003

In November 2003, we first observed prevalent occurrence of a hydroid, Eutima japonica, on soft body tissues of age zero Japanese scallop (Mizuhopecten yessoensis) juveniles cultured in large areas of Funka Bay, Hokkaido. The occurrence coincided with massive death of juvenile scallops. A major objective was to clarify ecological relationships between the symbionts, and to infer the relationship between symbiosis and the massive mortality. To do this, we investigated distributions of association rates of hydroids with juvenile scallops at 15–34 sites over 3 years (2003–2005), with age one adult scallops at 24 sites in 2003, and with mussels at 13 sites in 2004. We studied seasonal changes in association rates with juvenile scallops, and numbers of polyps per juvenile scallop at three sites from November 2003 to June 2004. We also quantified the hydroid impacts on juvenile scallop shell length growth and triglyceride accumulation in the digestive gland. The association rate of E. japonica polyps with juvenile scallops was high in large areas of Funka Bay in 2003, and overlapped the distribution of mussels bearing polyps. Association rates with age one adult scallops were very low in November 2003, even at the sites where polyps were very common on juvenile scallops. Levels of hydroid occurrence in juvenile scallops varies by year. We found that hydroids presence in juvenile scallops declined drastically in 2004 and 2005. The association rates with juvenile scallops, and numbers of polyps per juvenile scallop declined during winter, until they disappeared completely in the following June. Since polyps were rare in adult scallops, we believe that infection of juvenile scallops was probably initiated from the planulae produced by medusae released from polyps growing on Mytilus spp., especially M. galloprovincialis. Subsequently, the inhabitation spread intraspecifically and interspecifically. In juvenile scallops, inhabitation of polyps reduced shell length growth by 43%, and triglyceride accumulation in digestive glands by 24–47%. Inhabitation of E. japonica on juvenile scallop is best regarded as parasitism, rather than inquilinism or commensalism. Occurrence of polyps was probably not a direct lethal factor for juvenile scallops, because there were some sites where association rates were high, but mortalities were low. Massive mortalities in 2003 may have resulted from simultaneous impacts of heavy polyp load and stresses caused by the way in which the animals were handled (transferred from cages for pre-intermediate culture to cages for intermediate culture), because the massive mortality occurred within a month of the transfer. The presence of polyps in juvenile scallops does not affect the quality of the product in Funka Bay, because market size scallops are hydroid-free.     

Sexual reproduction in the corallimorpharian sea anemone Corynactis californica in a central California kelp forest

The seasonal cycle of sexual reproduction in the corallimorpharian sea anemone Corynactis californica (Calgren, 1936) was studied for 18 consecutive months (July 1987 to December 1988) at a subtidal area in the Hopkins Marine Refuge (HMLR), Pacific Grove, Monterey Bay, California. Samples were collected, histological sections were prepared, and gametocytes were examined and measured. C. californica grows by multiple or longitudianl fission to form single-sexed clones of various color and size. In female clones, oocytes appeared in late August and early September; they increased in size steadily in fall, and peaked in early December in both years. In male clones, spermatogenesis was synchronous with the female oogenic cycle, and motile sperm were observed in most testes in early December. Spawned gonads were found in both sexes from late November to early December. Spawning was induced in the laboratory in early December, and external fertilization was followed by development of free-swimming larvae. Gametogenesis and spawning are correlated with seasonal increases in seawater temperature and phytoplankton abundance, and we discuss and compare timing of sexual reproduction in this corallimorpharian to those in several actiniarian and scleractinian species.     

Reproductive ecology of the brooding,hermaphroditic clam Lasaea subviridis

The reproductive ecology of a small intertidal clam was studied for two years (September 1980-November 1982) at two mid-intertidal sites on the northern shore of Monterey Bay, California. Lasaea subviridis is hermaphroditic and incubatory; the young are released as juveniles. Brooding adults are found throughout the year. All embryos in a clam are at the same stage of development, but the number and stage of embryos varies among clams at any one time. Individuals are iteroparous and produce small numbers of oocytes and sperms simultaneously. All stages of gonad development are found in different individuals at any one time. There was no evidence of seasonality. Young reared in the laboratory after being dissected from the parent reached brooding size (shell length of 1.7 mm) within 6 mo. Laboratory experiments indicated that juveniles crawl away soon after release from the parent. These reproductive characteristics may contribute to this taxon's ability to colonize new environments rapidly.     

Hybridization between the blue mussels Mytilus galloprovincialis and M. trossulus along the Pacific coast of North America: evidence for limited introgression

Two species of blue mussel, Mytilus galloprovincialis and M. trossulus, co-occur and hybridize along the Pacific coast of North America. Using a set of polymerase chain-reaction (PCR)-based genetic markers which diagnostically identify these species, we show that they are sympatric from the Cape Mendocino region to the Monterey Peninsula in northern and central California, USA. Mussels with hybrid genotypes were detected in all populations sampled in the region of sympatry, and the frequency of hybrid genotypes in individual hybrid populations ranged from 13 to 44%. Significant frequencies of first-generation backcross genotypes were detected in two individual hybrid zone populations (Berkeley and Monterey Marina) and in the hybrid zone as a whole, indicating that the potential exists for introgression between M. galloprovincialis and M. trossulus. Despite this potential, we found no evidence of advanced introgression beyond first-generation backcrosses, suggesting that gene flow between M. galloprovincialis and M. trossulus has been quite limited. The frequency of mussels with M. trossulus and hybrid genotypes declined abruptly south of Monterey Peninsula, while the frequency of mussels with M. galloprovincialis and hybrid genotypes declined precipitously north of Cape Mendocino. These abrupt genetic discontinuities indicate that this blue mussel hybrid zone is presently positioned between two prominent coastal features and there is little, if any, export of alleles from the hybrid zone into bordering parental populations. Received: 20 August 1997 / Accepted: 26 October 1998     

Life cycle ofRhopilema nomadica: a new immigrant scyphomedusan in the Mediterranean

Rhopilema nomadica is an Indopacific scyphomedusan, which has migrated into the eastern Mediterranean in recent years. Large aggregations of the medusae were recorded in Haifa Bay, Israel, reaching 5.5×10⁵ medusae per square nautical mile during summer 1989. The life cycle ofR. nomadica from planula to young medusa is described. Fertilization is external and planulae are formed within a few hours at 20°C. After settlement, polyps were fed withArtemia sp. nauplii and developed into polydisc strobilae within 45 d. The strobilation process was completed within 7 d, and the liberated ephyrae developed into young medusae within 2 mo. Asexual reproduction occurred mainly via podocyst formation. The population explosion ofR. nomadica could be attributed to its high reproductive potential.     

Hepatic mixed-function oxidases in California flatfishes are increased in contaminated environments and by oil and PCB ingestion

Hepatic mixed-function oxidases (MFOs) were measured in the bothid flatfishes Citharicthys sordidus and C. stigmaeus from relatively uncontaminated and from polluted coastal populations of California, USA, at various times of the year during 1979–1980 and in individuals fed crude oil and polychlorinated biphenyl-augmented food in the laboratory. For C. sordidus, aryl hydrocarbon hydroxylase (AHH) specific activity was generally highest around the Los Angeles County sewage outfall on the Palos Verdes Shelf, intermediate near the 7-mile Hyperion sewage outfall in Santa Monica Bay and around a petroleum seep in the Santa Barbara Channel, and lowest in relatively unpolluted Monterey Bay. For C. stigmaeus, which had about ten times less specific activity than the foregoing species, specimens from the Santa Barbara petroleum seep had significantly greater AHH specific activity than those from Monterey Bay. Fishes from contaminated environments also showed increases of microsomal proteins with molecular weights of 56, 54, 57 and 46×10³; moreover, the content of cytochrome P-450 was elevated in specimens of C. sordidus from such environments. Augmentation of food with seep oil or polychlorinated biphenyls (PCBs) induced significant increases in the specific activity of AHH and amounts of microsomal proteins in C. stigmaeus. Thus, these two species of flatfishes are good candidates for monitoring biologically meaningful levels of petroleum and polychlorinated biphenyls in contaminated environments. Moreover, the mixed function oxidase pattern in fish populations from the Santa Barbara petroleum seep is evidently a functional adaptation to chronic intake of petroleum hydrocarbons.