Metazoan/microbial biostalactites from present‐day submarine caves in the Mediterranean Sea

Biostalactites formed by metazoan–microbialite associations from three submerged marine caves in the Plemmirio Peninsula (south of Syracuse, Ionian Sea) are randomly distributed and show different sizes and morphologies, as well as variations in surface roughness/smoothness. The biostalactites consist of crusts a few centimeters thick of small serpulids and other metazoans, associated with fine‐grained carbonate; the larger ones often include a nucleus of serpulid tubes (Protula). The metazoans include mainly serpuloideans, sponges, bryozoans and foraminifers but microbial carbonates are also significant components. The composition of both the living communities and thanatocoenoses on the outer surfaces, as well as the composition and fabric of the internal framework, were analysed and used to reconstruct the history of the caves. All of the identified sessile faunas mainly consist of cryptic and sciaphilic dwellers that reflect cave conditions and their variations through time. The distribution pattern, composition and abundance of the present‐day dwellers largely depend on the degree of roughness of the biostalactite surfaces and their positions within the caves. It has been suggested that the Protula specimens in the nuclei represent pioneer populations that formed aggregates during the early cave colonization phase, in response to relatively high food supply from seawater inflow and intruding continental waters. By contrast, the outer metazoan–microbialite carbonates reflect more confined conditions in the caves caused by Holocene sea‐level rises. Hypotheses are proposed for biostalactite growth, taking into account information about the growth rates of some constituents, and evidence of dissolution effects. Similarities and differences between these biostalactites and other Holocene deposits previously described from submarine caves in the Mediterranean Sea and in tropical reefs are discussed.     

Horizontal Transfer of Matter by a Cave-Dwelling Mysid

Abstract. The role of a cave-dwelling mysid population as a matter carrier was evaluated in a cave of the Medes Is. (NW Mediterranean) during 1988–89. Hemimysis spehmcola (Ledoyer , 1963) is a gregarious mysid whose swarms migrate daily from the inner end of the cave - where they remain during the day–to the exterior where they feed during the night. Fecal pellet composition, pellet egestion and pellet decomposition were measured in order to evaluate the transfer of matter by mysids. Based on the strong daily behavioural rhythms of these mysids. special importance was attached to the sampling frequency (every 2 h). In order to assess seasonal variability, four daily cycles were evaluated within a year. In situ incubations were carried out to determine changes on the egestion rates, morphology, weight and composition (AA. C: N ratios) of pellets in the course of a day. The number of pellets deposited inside the cave was estimated using containers distributed along the cave bottom by SCUBA divers. Fecal pellets showed an amorphous composition, whereby diatom frustules, dinoflagellate loricae and coccolithoporids were very scarce. About 25% of the body weight were estimated to be egested daily as fecal pellets, suggesting a detritivorous feeding habit. The population oscillated seasonally between 1 and 12 millions. Individuals egested between 1.6 and 3.5 pellets a day into the cave, each averaging from 9.4 to 11.9 μg DW, 0.5 to 1.0 μg C and 0.046 to 0.27 μg N. Therefore, the population carried daily about 20 407 g DW POM. 2–21 g C and 0.5 2.7 g N from outside to inside the cave. Pellets decomposed very quickly; between 20 to 50% of both C and N were released from pellets in less than 2 h after egestion. Oxidation of pellets theoretically consumes an amount of oxygen which agrees with the high BOD values previously reported for this cave. Marine caves are generally viewed as strictly oligotrophic systems; dense mysid populations, however, could strongly modify the trophic relationships in marine caves.     

Colonisers of the dark: biostalactite‐associated metazoans from “lu Lampiùne” submarine cave (Apulia,Mediterranean Sea)

Metazoan/microbial bioconstructions, or biostalactites (BSTs), discovered in submarine caves of Apulia c. 20 years ago—and later found in several shallow‐water Mediterranean caves—are receiving increasing attention in the last years. Examination of a single BST from the “lu Lampiùne” cave (Apulia), at the limit between the Adriatic and the Ionian seas, has been addressed for the first time in this paper. The BST started growing at c. 6,000 years ago with a rapid accretion of large‐sized serpulids (Protula), slowing down since about 3,000–4,000 years ago with a shift in main bioconstructors, probably caused by environmental changes. The present‐day community on the outer BST surface is dominated by skeletonised epibionts, mostly small‐sized serpulids, bryozoans and foraminifers, which contribute carbonate to the BST growth, by encrusting sponges, and by a few endobionts, including boring bivalves and insinuating sponges. New data remarkably increase biodiversity known for the “lu Lampiùne” cave and the cave habitat in the region. Thirty‐five taxa (16 bryozoans, 10 serpulids, 3 brachiopods, 2 foraminifers, 2 sponges, 1 bivalve and 1 cirriped) are recorded for the first time from Apulian marine caves, highlighting the need for further research in the area. In addition, the BST‐associated community seems to differ from those of individual BSTs from other Mediterranean caves, revealing the individuality of these communities.     

Water Flow, Trophic Depletion, and Benthic Macrofauna Impoverishment in a Submarine Cave from the Western Mediterranean

Abstract. Biotic zonation and severe impoverishment of benthic macrofauna are two of the most conspicuous features of submarine caves. The prevailing explanation assumes trophic depletion caused by reduction of water flow. However, no isolation of water was found in the studied Catalan submarine cave, even in the wall microlayer. Fluorescein diffusion was so fast that it was detectable everywhere in the cave already only a few minutes after the injection. The rate of dissolution of plaster spheres was even greater in the cave than in a nearby tunnel – without benthic macrofauna impoverishment – showing a considerable water flow. The oxygen concentration of water in dialysis bags placed at varying distances from the cave walls showed that no wall microlayer gradients were present.
Biochemical gradients did not indicate any decrease in food supply. Although photosynthetic pigment concentration generally decreased inside the cave, and although the gradients were not linear but formed different patterns throughout the cave, the BOD₅, the POM, the C:N ratio, the [³H]-thymidine incorporation rate, and the ETS activity were higher in the inner part of the cave. There was an accumulation of detritic material and bacteria in the inner parts that constitute a plausible food supply for benthic macrofauna.
If neither water motion nor food supply can be invoked, research into the causes of zonation and disappearence of benthic macrofauna is proposed to be carried out on biotic interactions and behavioural processes.     

Unexpected records of ‘deep‐sea’ carnivorous sponges Asbestopluma hypogea in the shallow NE Atlantic shed light on new conservation issues

Marine cave communities have been a continued source of ecological surprises, among other things because of their close ecological and evolutionary ties with the deep sea. The discovery of cladorhizid sponges, the deepest occurring poriferan family, in shallow Mediterranean caves in the 1990s was one such surprise, leading to the generally accepted hypothesis that the whole family was carnivorous, an unprecedented feeding mode for sponges. The recent observation of the cave species Asbestopluma hypogea in the Mediterranean bathyal, confirmed the view that some shallow caves can occasionally shelter otherwise deep‐dwelling species. Here we present new distribution data of A. hypogea, from deep Mediterranean locations, and for the first time from Atlantic locations. Among the new Atlantic records, the most surprising ones are located in three different geographic areas (Ria de Arousa, Groix Island and Cherbourg) of the NW European coasts, from the Iberian Peninsula to the English Channel, where A. hypogea reaches SCUBA depths (5–50 m), while not sheltered in marine caves. The carnivorous sponge however reaches its shallowest occurrence (5 m), in a small cave at Groix Island. The ecological significance of these discoveries, particularly the very patchy distribution and peculiar dynamics, are noteworthy, and the shallow occurrence of A. hypogea, together with other deep‐water or uncommon species, constitute unique assemblages that must be considered in conservation plans.     

Do Mediterranean fish assemblages associated with marine caves and rocky cliffs differ?

Fish assemblages associated with marine caves and rocky cliffs were investigated in the Salento Peninsula (SE Italy, Mediterranean Sea) by using visual census methods. Sampling was done at three sites, each of which included 4 habitat types: the external and the internal portions of caves, and shallow and deep rocky cliffs. 10 and 13 species were found exclusively inside cave habitats (e.g. Corcyrogobius lichtensteini, Thorogobius ephippiatus and Grammonus ater) or in rocky cliffs (e.g. Diplodus annularis, Sarpa salpa, Sparisoma cretense, Spondyliosoma cantharus), respectively. The four habitat types shared 10 species, and the external portions of the caves shared the most species (both with the internal cave portions and the external rocky cliffs). As a general rule, dissimilarity in the fish assemblage structure between habitats was far greater than dissimilarity between sites. Apogon imberbis (mostly associated with caves) and Chromis chromis (typifying rocky cliffs, mainly the deep ones) mostly contributed to dissimilarities between caves and rocky cliffs. Apogon imberbis (mostly associated with internal caves) and Coris julis (mainly associated to external cave portions) contributed strongly to dissimilarities between internal and external cave portions, while C. chromis, Symphodus mediterraneus and C. julis (associated with the deeper cliffs) and Thalassoma pavo (mostly present in shallow cliffs) differentiated deep and shallow cliffs. Diplodus vulgaris, Oblada melanura and Mullus surmuletus showed a marked increase in density during the cold season in the caves. These results show that fish assemblages associated with rocky reefs rich in marine caves (in terms of relative densities, species composition, species richness, exclusive species and presence of juveniles of some valuable species) may be affected by the peculiar ecological conditions within caves, which could provide additional resources for fishes (e.g. food availability, refuge against predators, sand patches within a rocky matrix) compared to rocky reefs without caves. These results suggest that stretches of rocky coasts rich in marine caves should be considered within management/conservation programs (e.g. when establishing Marine Protected Areas).     

Seasonal abundance, distribution and recruitment of mud crabs (Scylla spp.) in replanted mangroves

The abundance and distribution of mud crabs were studied in a replanted mangrove forest in Buswang, Aklan, Philippines. Two fishing gears, lift nets and bamboo traps, were used to monitor relative abundance of Scylla spp. populations from March 2002 to December 2003 inside the mangrove forest. A third gear, a stakenet set across a creek, was used to monitor crabs migrating out of the mangroves during the ebb tide. Scylla olivacea formed 99.3% and 70.3% of the catch in the mangrove and the stakenet, respectively. The percentage of Scylla tranquebarica increased from <1% in the mangrove catches to 29% in the stakenet. Scylla serrata was present at very low levels in both catches. The lack of modal progression in the size–frequency plots and the year-round catch rate of gravid females suggested that recruitment was constant throughout the year. Even though relative abundance decreased over the study period indicating that the stock is being over-exploited, mud crab production is more than equivalent to that of most natural mangroves.     

Alien biodiversity in Mediterranean marine caves

The number of alien species in the Mediterranean Sea is increasing rapidly, but few attempts have been made to evaluate impacts on specific habitat types. The present study investigated alien biodiversity in Mediterranean marine caves, both by contributing new records of Eastern Mediterranean cave aliens, and by reviewing the scattered existing literature; the main goals were to highlight potential impacts and investigate the importance of cave environments for the expansion of alien species. Seven new alien species were found in marine caves of the Aegean and Ionian seas, raising the total number of aliens reported from Mediterranean marine caves to 56 species, classified as molluscs, cnidarians, bryozoans, polychaetes, crustaceans, macroalgae, fishes and tunicates. Most cave aliens (66%) were recorded from the Southeastern Levantine coasts, specifically from Lebanese caves. Shipping and Lessepsian migration have been suggested as the main pathways of alien introduction into caves of the Mediterranean Sea. The comparison of alien cave biodiversity with the updated Mediterranean alien inventory (32 species added to the latest inventory) showed similar patterns and trends for species richness, biogeographical origin and major introduction pathways, corroborating previous findings on the high local representativeness of Mediterranean cave biodiversity. Alien species seem to have invaded mostly the entrance and semi‐dark zones of shallow and/or semi‐submerged caves and tunnels, whereas only a few have reached the dark inner sectors or caves of the anchialine type; thus, the unfavourable cave environment seems to be naturally protected from impacts related to opportunistic invasive species, at least to a certain point. Currently there is no research confirming any direct impacts of alien biota on the native cavernicolous one. However, some issues have emerged, implying potential threats that need to be further explored: (i) the presence of a considerable proportion of aliens in most studied marine caves of the Southeastern Mediterranean basin, (ii) the recently observed population explosion of alien cave‐dwelling fishes in the same area, (iii) several indications that alien diversity in marine caves is much higher than we know today. Quantitative surveys and monitoring schemes are needed in order to evaluate potential effects of alien diversity on cave community structure and the role of marine caves as stepping stones for its expansion in the Mediterranean.     

The distribution of iodide and iodate in anchialine cave waters — Evidence for sustained localised oxidation of iodide to iodate in marine water

A study of inorganic iodine speciation in the waters of seven Croatian coastal caves is described. These are anchialine caves as they are connected hydraulically with Adriatic Sea surface water, with the tide inside the cave rising and falling with that outside, but replenishment of the water is restricted by the karst rock. In effect, the water in the cave probably acts more like a piston, and although moving slightly vertically, has a long residence time compared to a fully-flushing cave. Anchialine environments display a number of unusual features, e.g., a well-developed pycnocline, hypoxia and endemic fauna. Iodate and iodide were determined by differential pulse voltammetry and cathodic stripping square wave voltammetry, respectively. Low iodide concentrations (< 10 nM) have been consistently identified in the bottom water of the caves where concentrations of 90–100 nM would ordinarily be expected from intrusion of Eastern Adriatic surface seawater. Where total inorganic iodine concentrations behave conservatively with salinity the loss of the iodide implies oxidation to iodate. As iodide oxidation remains one of the enduring academic problems of the marine iodine system the study of iodine in anchialine caves may help unravel it. Iodate reduction was observed in mid-water, at the halocline, and mechanisms for the reduction involving either respiration or chemolitho-autotrophic bacteria are considered. The respiration mechanism is favoured because of enhanced alkalinity found in the near surface waters of the caves.     

Organic pollution of an inland marine cave from Bermuda

Inland marine caves have served as biological refugia and, as such, frequently contain unique assemblages of taxonomically significant relict populations known only from a single cave or cave system. However, the absence of photosynthesis in this totally lightless environment and the presence of a highly stratified water column and long residence times for cave waters, characteristically result in depleted dissolved oxygen levels. Under such circumstances, organic pollution, even at low levels, can produce anoxic conditions and may consequently result in the extinction of entire species of cavernicolous fauna. We here present data on the normal environment of marine caves in Bermuda and cite as an example a cave which has become grossly polluted as a result of indiscriminate dumping.     

Integrative taxonomic description of Plakina kanaky,a new polychromatic sponge species from New Caledonia (Porifera: Homoscleromorpha)

Four similar sponges of different colors, all unknown to science, were collected in submarine caves of New Caledonia. We aimed at determining whether the four chromotypes represented different species or phenotypic variations of a unique new species. We used an integrative taxonomic approach combining morphologic, molecular and metabolomic analyses. The main traits that define these specimens are a skeleton made of monolophose, trilophose and tetralophose calthrops only, high chemical diversity and a high abundance and diversity of prokaryotic symbionts. The symbiotic community includes two unique prokaryote morphotypes, which are described for the first time in Homoscleromorpha, and appeared to be vertically transmitted. Although several features slightly differ among chromotypes, the most parsimonious conclusion was to propose a single new species Plakina kanaky sp. nov. Our phylogenetic analysis indicated the paraphyly of the Plakina genus, with P. kanaky sp. nov. belonging to a clade that includes Plakina jani and Plakina trilopha. The present work demonstrates that integrative taxonomy should be used in order to revise the entire Plakinidae family and especially the non‐monophyletic genus Plakina.     

Distribution and movement of scalloped hammerhead Sphryna lewini and smooth hammerhead Sphyrna zygaena sharks along the east coast of southern Africa

Knowledge of population distribution and movement is crucial for the conservation and management of shark species occurring in coastal waters. From 1984 to 2009, 641 scalloped hammerheads Sphyrna lewini, 1 342 smooth hammerheads Sphyrna zygaena and 1 352 unspecified hammerheads Sphyrna spp. were tagged and released along the east coast of South Africa, with recapture rates of 1.9%, 1.5%, and 0.7% respectively. Maximum and average distance moved was 629 km and 147.8 km (95% CI = 33.0–262.7 km) for S. lewini and 384 km and 141.8 km (95% CI = 99.1–184.5 km) for S. Zygaena respectively. The majority of sharks (68% S. lewini, 74.1% S. zygaena and 33.5% Spyrna spp.) were tagged in the Transkei region, with the largest number tagged in Port St Johns. Across regions, most tagged sharks were >50–100 cm precaudal length (PCL), except in Transkei where more sharks >100–150 cm PCL were tagged. In the Western Cape, Southern Cape and Eastern Cape, few sharks were tagged during the autumn/winter months, whereas in KwaZulu-Natal and Transkei sharks were tagged throughout the year. Large-scale directional movements observed may have been migrations in response to seasonal sea surface temperature changes. We identify coastal locations in Transkei that are of importance to juvenile and subadult hammerhead populations year-round.     

Seasonal succession and vertical distribution of phytoplankton in Lake Hayes and Lake Johnson,South Island,New Zealand

In two small monomictic lakes near Queenstown, South Island, New Zealand, algal associations characteristic of eutrophic waters were present throughout most of the period from December 1969 to February 1972.

In Lake Hayes, five of the six algal genera which were recorded in the plankton in 1948–50 were still present, but Anabaena appeared to have increased. Blooms of A. flos‐aquae which were not recorded 20 y ago (Jolly 1952) occurred in the lake in the three summers of this study and in winter of the second year. Melosira granulata dominated the plankton in winter; at other times of the year Closterium aciculare, Cyclotella kuetzingiana, and Staurastrum spp. were dominant.

In Lake Johnson the major algae differed from one year to another. Blooms of Anabaena flos‐aquae formed during the first two summers but were absent in the third, when Closterium aculum var. variabile was dominant. Peridinium cinctum was abundant throughout the first year, especially in December 1969 and October 1970, when concentrations of more than 1000 cells per ml occurred at the surface. During autumn and winter of the second year, Staurastrum bibrachiatum dominated the plankton.

In both lakes micro‐algae were abundant in late spring and early summer.

In winter when the lakes were isothermal, algae were distributed fairly uniformly with depth. During the period of thermal stratification, algae were mainly confined to the epilimnion. Although green algae and diatoms were usually dispersed fairly uniformly throughout this zone, Cryptomonas and Peridinium were often concentrated at a certain depth. In Lake Johnson in late summer 1971, a layer of purple photosynthetic bacteria, of which concentrations of 0.5–1.3 × 10° cells per ml were recorded, was present at a depth of 7–8 m.

Algae were more abundant in summer than in winter in both lakes. The mean standing crop of algae for the trophogenic zone of Lake Hayes was 6000 cells per ml, and a maximum crop of 65 000 cells per ml was recorded in December 1971 during an Anabaena bloom: the mean volume of phytoplankton was 1.93 mm³ per litre for the trophogenic zone with a maximum of 7.49 mm³per litre in January 1970 when Anabaena and Peridinium were abundant. In Lake Johnson the mean number of algae in the trophogenic zone was 11000 cells per ml and the mean volume of phytoplankton was 9.37 mm³ per litre: a maximum volume of 33.12mm³ per litre was recorded in October 1970 when Peridinium was abundant.     

Seasonal growth in brown trout in two New Zealand streams

Brown trout Salmo trutta Linn, were marked with individual tags and recaptured throughout a year in two small Canterbury streams. The growth recorded showed a marked seasonal variation which differed in pattern in the two streams. There was a considerable difference in the growth rate between the two populations and fish transferred from the slow‐growth stream to the faster‐growth stream showed an immediate response to the change.     

Protozoa in the diets of Neocalanus spp. in the oceanic subarctic Pacific Ocean

Copepod species of the genus Neocalanus dominate the zooplankton biomass of the oceanic subarctic Pacific Ocean. Neocalanus spp. populations in the subarctic Pacific environment are successful: they feed, accumulate lipid, and persist from year to year. Prior experimental observations derived from a variety of methods indicated that, although their functional morphology is such that they clear the small phytoplankton cells characteristic of the oceanic subarctic Pacific environment efficiently, Neocalanus spp. do not consume sufficient phytoplankton to meet even basic metabolic requirements in that environment. Hence, their success in the subarctic Pacific must depend on their ability to obtain nutrition from other sources. As part of the SUPER (SUbarctic Pacific Ecosystem Research) program, experiments were performed to test the hypothesis that N. plumchrus and N. cristatus obtain a significant portion of their nutrition from planktonic Protozoa. The experiments demonstrate that Protozoa alone do not provide sufficient nutrition for N. cristatus to meet its basic metabolic needs. Protozoa constitute the major dietary component of N. plumchrus however, in agreement with the predictions of Frost's (1987) model of the subarctic Pacific ecosystem. At a minimum this diet permits N. plumchrus to meet basic metabolic requirements. Copepod grazing activities appear to be sufficient to control protozoan stocks in the oceanic subarctic Pacific during late spring and early summer when Neocalanus spp. inhabit the upper water column.     

Attachment and feeding of pelagic copepods on larvacean houses

The attachment and feeding of pelagic copepods on discarded and occupied oikopleurid larvacean houses were investigated from May 1989 to August 1991 at an offshore station in the Nansei Islands, which is strongly influenced by the Kuroshio Current, and an inshore station in the central Seto Inland Sea.Oncaea spp.,Microsetella norvegica, and other calanoid and cyclopoid copepods were found attached onto discarded and occupied houses throughout the year at the offshore station, whereas neither discarded nor occupied houses, carried copepods at the inshore station at any time. The feeding behavior ofOncaea sp. in a discarded house was observed with a video tape recorder system.     

Oscarella balibaloi,a new sponge species (Homoscleromorpha: Plakinidae) from the Western Mediterranean Sea: cytological description,reproductive cycle and ecology

A new homoscleromorph sponge species was observed for the first time in 2002 along the coast of Provence (France) and since then, several investigations have been implemented. The new homoscleromorph species is devoid of skeleton and it is thus related to the Oscarella genus. Oscarella balibaloi sp. nov. has been found in several littoral submarine caves of the Western Mediterranean Sea (France and Spain). After several years of monitoring, we show that this sponge is becoming abundant in several locations. The species has a peculiar habit, often overgrowing massive sponges, gorgonians or some erected bryozoans. Oscarella balibaloi sp. nov. is thinly encrusting, with an irregular morphology, lobate surface, and soft and very slimy consistency. Its color is white to orange. At the cytological level, this new species is characterized by three distinct types of cells with inclusions in its mesohyl: one with vacuoles and the others with paracrystalline inclusions. The reproduction pattern is very similar to other Oscarella species. We also present a specific metabolic fingerprint and compare it to that of other Mediterranean Oscarella species.     

中国龙虾栖息习性的观察

于1999年5月在集美大学水产学院海水养殖试验场进行中国龙虾栖息习性的试验观察。结果表明 ,正常温度、盐度条件下 ,中国龙虾有隐匿穴居、昼伏夜出的栖息习性 ,喜欢栖息于紧身或身体易于找到依靠的水平洞穴中 ,并有同穴共居现象 ,低层洞穴的栖息率高于高层洞穴 ;温度低于12℃时 ,中国龙虾夜间也很少离穴活动 ,温度低于8.6℃或高于34℃ ,盐度低于14.1或高于44.6时 ,中国龙虾都会逃离洞穴     

Feeding of copepods on natural suspended particles in Tokyo Bay

Community grazing rates of copepods were estimated from data taken during three cruises in Tokyo Bay, based on bottle incubations and a temporal variation of gut fluorescence. Special attention was paid to the feeding selectivity in the estimations. Differential grazing was observed in the copepod communities:Acartia omorii, abundant in February, selectively fed on the particles of dominant size classes, whileOithona davisae, dominant throughout the year, andCentropages abdominalis selected large particles (>20µm). The maximum filtering rates on certain size classes were several times the average. In addition, a 34-hr investigation of the gut fluorescence of copepods revealed nocturnal feeding inParacalanus spp.,Pseudodiaptomus marinus andOithona davisae.Copepod communities collected with a net (95-µm mesh opening) were estimated to graze, in February 3.0%, in August 3.1–4.5% and in November 4.2–11.9% of the standing crops of phytoplankton or suspended particles per day.