The vertical distribution of Chthamalus montagui and Chthamalus stellatus (Crustacea,Cirripedia) in two areas of the NW Mediterranean Sea

The Mediterranean Sea is characterised by a small tidal range (0.3–1 m). Despite this, intertidal communities are well established and their upper limits often extend above mean high water level. Organisms living in the intertidal region and in the supralittoral zone rely on both tides and wave action to perform their biological functions. Lack of food, desiccation and predation are common stresses in such a harsh environment. The present study deals with the vertical distribution of two species of intertidal barnacles, Chthamalus montagui Southward and Chthamalus stellatus (Poli), which are the main constituents of the barnacle belt along Mediterranean rocky shores. Previous work, carried out in the Atlantic, showed that where the distribution ranges of the two Chthamalus species overlap, C. montagui is more common in the upper barnacle zone while C. stellatus is dominant lower down. The main aims of our study are: (1) to establish if there is a relationship between position and extension of the barnacle belt on the shore and tidal range and/or wave exposure, (2) to test the hypothesis that in the study areas C. montagui is more abundant than C. stellatus high on the shore, and that the pattern is reversed lower down. Barnacle populations were monitored in summer 1998 in the Gulf of Genoa (Ligurian Sea) and in the Gulf of Trieste (North-Adriatic Sea). The two areas differ in tidal range and hydrodynamism, the former presenting quite strong wave action and a tidal range of 30 cm, the latter having limited wave action and 1 m tidal range. Three shores were randomly selected in each gulf and two transects on each shore. Counts of barnacles in 10 ^* 10 cm quadrats were done at different shore heights along each transect. The data was subjected to analysis of variance. Results showed that a more pronounced hydrodynamic regime corresponded to a shift of the barnacle belt towards the higher shore (Gulf of Genoa), while in more sheltered areas (Gulf of Trieste), the barnacle distribution was confined to the intertidal region. The relative spatial distribution of C. montagui and C. stellatus within the barnacle belt varied locally, even between transects on the same shore, and this obscured the distribution pattern along the vertical gradient. Nevertheless, it was still possible to conclude that at mid and high shore in Genoa, C. stellatus was more abundant than C. montagui, while in Trieste the pattern was reversed.     

Aspects of the reproductive biology ofPollicipes pollicipes (Cirripedia; Lepadomorpha) from the southwest coast of Portugal

The pedunculated cirripedePollicipes pollicipes is of considerable economic importance in Portugal, particularly along the southwest coast. Current knowledge of its biology is poor, as evidenced by the paucity of available literature. This paper reports on the reproduction of twoP. pollicipes populations, Castelejo and Zavial, on the southwest coast of the Algarve. Differences in reproductive patterns between the two sites were minor and not statistically significant, despite differences in environmental conditions between the two sites. Adult animals carried fertilised eggs from April throughout September, and the percentage of individuals with egg masses correlated positively with air but not with sea water temperature. Brooding success was greater with average air temperature in excess of 14°C during months of strong upwelling and presumed food availability. Functional ovaries and testes are available in the population all year round but fertilisation does not appear to occur during the winter months. Reproduction ofP. pollicipes on the southwest coast of the Algarve appears closely linked to the upwelling phenomenon with the rate of deposition of gonadal tissue probably being determined by environmental temperature. The cues which determine the cessation of fertilisation remain to be investigated.     

Spatial variation in the recruitment of the intertidal barnacles Chthamalus montagui Southward and Chthamalus stellatus (Poli) (Crustacea: Cirripedia) over an European scale

Spatial variation in the recruitment of the intertidal barnacles Chthamalus montagui and Chthamalus stellatus was examined over an European scale. The study was carried out using standardised protocols at a series of locations. The five locations chosen (SW Ireland, NW Spain, SW Portugal and NW and NE Italy) span a large part of the range of these species in Europe. The spatial scales were location (hundreds of kilometres) and shore (thousands of metres).Estimates of total cumulative recruitment (cyprids and metamorphs) summed over the year (April 1997 to March 1998) showed substantial variation between locations which was dependent on the species. Recruitment was highest in SW Portugal for C. montagui and in SW Ireland and NW Spain for C. stellatus. Overall recruitment of C. montagui was higher than that of C. stellatus at all locations except SW Ireland, where recruitment of the two species was not significantly different. There were significant differences among shores in each location.The recruitment period of both species varied with location, with recruitment beginning earlier further south. In general, recruitment of C. montagui and C. stellatus was recorded in 8 months in NW Spain and NE Italy, while only in 7 months in SW Ireland. Recruitment of C. montagui occurred in 10 months in SW Portugal, but no recruits of C. stellatus were found. In all locations there was at least one distinct peak of recruitment. In SW Ireland both species showed only one peak of recruitment, a month after initiation. At the more southerly Atlantic locations, as well as in the Mediterranean, two unequal peaks of recruitment were generally seen. During recruit census, the number of cyprids, in comparison to metamorphs, found at any location was very low. In SW Ireland and NW Spain cyprids of both species were found, while in SW Portugal and in the Mediterranean, only cyprids of C. montagui were found.     

Densely packed concentrations of sessile barnacles (Cirripedia: Sessilia) from the Early Pliocene of SE Spain

Sessile barnacle assemblages, dominated by Concavus concavus (Bronn) and Balanus perforatus Bruguière, are very abundant in the Lower Pliocene deposits of the Almería-Níjar and Carboneras basins (southeastern Spain). They occur in shallow-coastal siliciclastic and mixed siliciclastic-carbonate sediments, forming dense concentrations in two contexts: (1) sheltered shallow-marine depressions and (2) the mouth of distributary channels feeding a delta lobe. Extensive colonization took place during periods of quiescence with a high nutrient and food supply inducing the formation of hummocks. Crowding of high morphotypes was presumably triggered by a high larval supply and recruitment rate. The barnacles are exceptionally well preserved, often as in situ clusters, many with opercula, and include a range of ontogenetic stages with some specimens preserving original colouration. Good preservation is probably due to sudden burial without further reworking.     

Reproductive Stage of the Life Cycle in the Rhizocephalan Barnacle Polyascus polygenea (Crustacea: Cirripedia)

The specific features of the reproductive stage of the life cycle have been studied in the rhizocephalan barnacle Polyascus polygenea, a parasite of the coastal crab Hemigrapsus sanguineus. It is shown that a single crab can bear 1 to 8 externae of P. polygenea. The fecundity of the parasite depends on the size of the externae and their number on the host and may reach as much as 50000 eggs for one externa. In Peter the Great Bay, Sea of Japan, this species repeatedly reproduces during the entire spring–autumn period; externae with developing embryos in the mantle cavity occur from May to September, and planktonic larvae occur from June to October. One externa produces during the season of reproduction no less than three generations of larvae. Thus, the reproductive strategy in P. polygenea comprises a three-stage cascade of reproduction: asexual reproduction via budding of the interna; the development of several generations of one or several externae; and several reproduction cycles of each externa. This allows the parasite to produce a very great number of larvae and ensures the parasitization of a significant proportion of the host crab population. The structure of the ovaries and oogenesis in rhizocephalans and free-living cirripede barnacles have many common features, which provides evidence for integration of these two groups within one monophyletic taxon.     

Patterns of adult abundance in Chthamalus stellatus (Poli) and C. montagui Southward (Crustacea: Cirripedia) emerge during late recruitment

The aim of this study was to investigate when adult distribution patterns are established in the barnacles Chthamalus stellatus and C. montagui. Adult ‘zones’ were identified by analysing field counts of both species at mid and upper shore heights. Monthly collections of cyprids, < 1 month old metamorphs and recruits (all metamorphosed individuals older than approximately 1 month) were made for C. stellatus and C. montagui in natural barnacle beds at six shores in SW Ireland. This was carried out over one year in 1996/1997, using a hierarchical sampling design. Abundance of total recruits (0-3 months old) was compared between adult zones after the main settlement season had ended. In addition, scales of variability in 0-3 month recruitment into adult zones were compared between the species at two scales: shores (1000s of metres) and sites within shores (10s of metres). Older recruits of each species, up to 11 months of age, were also compared between adult zones.The majority of settlement (measured as attached cyprids) occurred between August and October 1996. In October, there was no effect of adult zone on the abundance of total (0-3 month) recruitment up to that point in either species. Despite this homogeneity in recruitment between adult zones, significant spatial variation was found in 0-3 month recruits of both species at both of the scales examined. In C. stellatus the amount of variation associated with the larger scale (shore) was more than twice that of sites or of the residual variation (replicates within sites). 0-3 month recruitment in C. montagui was also most variable at the scale of shores but the residual variability (between replicates within site) was of similar magnitude to that of shores. Variability in 0-3 month C. montagui recruitment was relatively low at the scale of sites.There was a small but consistent input of recruits to adult zones over 9 months of the year, complicating the assessment of when adult patterns were set-up in these species. By June 1997, characteristic patterns of adult dominance had been established at all shores. Settlement had completely ceased by this time and individual barnacles were potentially 11 months old. Neither settlement nor early recruitment are significant in determining adult zonation patterns in these species. Instead, differential mortality patterns in individuals up to the age of 11 months are implicated in determining patterns of distribution of both species.     

Distribution and abundance of marine wood borers on the west coast of Peninsular Malaysia

Twenty one species of marine wood borers were recorded in this study. There are seventeen species of bivalves of which sixteen are from the family Teredinidae and one from the family Pholadidae. The crustaceans comprised the family Sphaeromatidae with three species and the family Limnoriidae with one species. Seven of the fourteen known genera of the teredinids were recorded. The bivalve wood borers were widely distributed in the local mangroves. Martesia striata was the most abundant wood borer at the Degaussing Range jetty, Lumut. Lyrodus pedicellatus was the most abundant shipworm in wooden panels. Shipworms were abundant on the subtidal panels where the genus Bankia, was most common while M. striata was abundant on the intertidal panels. M. striata numbers were higher on the top surfaces as compared to the under surfaces of wooden panels.     

Littoral barnacles of The Snares islands,southern New Zealand (Cirripedia: Thoracica)

Abstract

Ibla idiotica, Lepas australis, Calantica villosa, Chamaesipho columna, Tetraclitella purpurascens, Notobalanus vestitus, and Notomegabalanus campbelli are reported for the first time from The Snares. The occurrence of Calantica spinosa and Epopella plicata is confirmed. Habitat information and recorded distribution in New Zealand are given for each species. The Snares barnacle fauna is restricted both in diversity and in numbers of individuals. This may be due to high wave energies, strong winds which aid desiccation, and counter-flow of currents, which might disrupt larval dispersal from the north and north-west.     

Larval development of the rhizocephalanSacculina polygenea (Crustacea: Cirripedia)

Larval development of the rhizocephalanSacculina polygenea (Crustacea: Cirripedia: Rhizocephala) parasitizing the coastal crabHemigrapsus sanguineus was studied in Vostok Bay, the Sea of Japan. At 22–23°C, the entire cycle of larval development takes 2.5 days and includes five naupliar stages and one cypris stage. Like other rhizocephalans, the larvae ofS. polygenea are lecithotrophic and only grow slightly in size in the course of development, and like all sacculinids, they have no flotation collar. The naupliar stages IV and V have a tubercle between the furcal rami; this tubercle is absent in the larvae of the genusPeltogasterella, but it has been described inS. carcini. The first seta of the antennule only disappears completely at the fourth stage, although it is markedly reduced at the third stage. No morphological differences, except differences in size, are found between male and female nauplii.     

Immunocytochemical studies on the naupliar nervous system of Balanus improvisus (Crustacea, Cirripedia, Thecostraca)

The nervous system of nauplii of the crustacean taxon Cirripedia was analysed in the species Balanus improvisus Darwin, 1854 using for the first time immunocytochemical staining against serotonin, RFamide and α-tubulin in combination with confocal laser scanning microscopy. This approach revealed a circumoesophageal neuropil ring with nerves extending to the first and second antennae and to the mandibles, all features typical for Crustacea. In addition, RFamidergic structures are present in the region of the thoraco-abdomen. A pair of posterior nerves and a pair of lateral nerves run in anterior-posterior direction and are connected by a thoracic nerve ring and a more posteriorly situated commissure. A median nerve is situated along the ventral side of the thoraco-abdomen. The innervation of frontolateral horns and the frontal filaments are α-tubulin-positive. Several pairs of large neurons in the protocerebrum, along the circumoesophageal connectives and in the mandibular ganglion stain only for serotonin. Due to the almost complete absence of comparable data on the neuroanatomy of early (naupliar) stages in other Crustacea, we include immunocytochemical data on the larvae of the branchiopod, Artemia franciscana Kellogg, 1906 in our analysis. We describe several characteristic neurons in the brains of the nauplius larvae of both species which are also found in decapod larvae and in adult brains of other crustaceans. Furthermore, our data reveal that the naupliar brain of cirripedes is more complex than the adult brain. It is concluded that this ontogenetic brain reduction is related to the sessile life style of adult Cirripedia.     

Molecules and the Body Plan: TheHoxGenes of Cirripedes (Crustacea)

Among arthropods, Cirripedia (barnacles) are remarkable in that they completely lack abdominal segments. This feature prompted us to study theHoxgenes of three cirripede species, representing a wide array of the diversity of these organisms, a segmented sessile barnacle,Elminius modestus(Thoracica), the parasite of a crab,Sacculina carcini(Rhizocephala), and the burrowing barnacleTrypetesa lampas(Acrothoracica). Using PCR amplification of genomic DNA and cDNA and library probing, we have found seven clear cirripedian homologues of the eight homeoticHoxgenes known in insects, includinglabialandproboscipediahomologues, that were not previously reported in crustaceans. In addition we have isolated a divergentAntp-like gene, namedDiva, that we homologize to theftzgene of insects. The homeotic geneabdominalA(abdA) was not retrieved from any of these three cirripede species. By contrast, we have found all eight homeotic homologue genes, includingabdA, inUlophysema oeresundense, a crustacean possessing a well-developed abdomen, belonging to the Ascothoracica, generally thought to be the sister group of Cirripedia. Since we have found in barnacles homeobox-containing genes that are more divergent from theAntennapediatype than the typicalabdA, we believe that abona fide abdAgene would not have escaped our search. Hence, theabdAgene has been lost or is profoundly derived in sequence during the evolution leading to the cirripedian lineage. If confirmed, the lack ofabdAwould represent the first case in which the loss of a homeotic gene is correlated with a change in body plan during the evolution of metazoans.     

Phylogeny and evolution of life history strategies of the parasitic barnacles (Crustacea, Cirripedia, Rhizocephala)

The barnacles (Crustacea, Cirripedia) consist of three well-defined orders: the conventional filter-feeding barnacles (Thoracica), the burrowing barnacles (Acrothoracica), and the parasitic barnacles (Rhizocephala). Thoracica and Acrothoracica feed by catching food particles from the surrounding seawater using their thoracic appendages while members of Rhizocephala are exclusively parasitic. The parasite consists of a sac-shaped, external reproductive organ situated on the abdomen of its crustacean host and a nutrient-absorbing root system embedded into the heamolymph of the host. In order to resolve the phylogenetic relationship of the order Rhizocephala and elucidate the evolution of the different life history strategies found within the Rhizocephala, we have performed the first comprehensive phylogenetic analysis of the group. Our results indicate that Rhizocephala is monophyletic with a filter-feeding barnacle-like ancestor. The host-infective stage, the kentrogon larva, inserted in the lifecycle of the rhizocephalan suborder, Kentrogonida, is shown to be ancestral and most likely a homologue of the juvenile stage of a conventional thoracican barnacle. The mode of host inoculation found in the suborder Akentrogonida, where the last pelagic larval stage directly injects the parasitic material into the heamolymph of the host is derived, and has evolved only once within the Rhizocephala. Lastly, our results show that the ancestral host for extant rhizocephalans appears to be the anomuran crustaceans (Anomura), which includes hermit crabs and squat lobsters.     

Speciation,phenotypic plasticity,or ontogeny,the case of the genus Galkinius (Pyrgomatidae,Cirripedia, Crustacea)

Barnacles of the genus Galkinius occupy a large spectrum of host corals, making it one of the least host‐specific genera within the Pyrgomatidae. Molecular analyses show that within the genus Galkinius there are highly supported clades, suggesting that the genus Galkinius is a complex of evolutionarily significant units (ESUs). The morphology of the opercular valves has been used as the basis for the separation of species of Galkinius. In this study, morphological variability was found both between specimens within ESUs extracted from different host species and between specimens extracted from the same colony. Identifications based on the opercular valves cannot therefore be assigned to different species despite being genetically distinguishable. It is proposed that in many cases the differences between valve morphology of different species of Galkinius are the outcome of ontogeny. Allometric growth of the valves has resulted in differences in the proportions of the parts of the valve. © 2015 The Linnean Society of London     

Phylogeny of coral-inhabiting barnacles (Cirripedia; Thoracica; Pyrgomatidae) based on 12S, 16S and 18S rDNA analysis

The traditional phylogeny of the coral-inhabiting barnacles, the Pyrgomatidae, is based on morphological characteristics, mainly of the hard parts. It has been difficult to establish the phylogenetic relationships among Pyrgomatidae because of the apparent convergence of morphological characteristics, and due to the use of non-cladistic systematics, which emphasize ancestor-descendant relationships rather than sister-clade relationships. We used partial sequences of two mithochondrial genes, 12S rDNA and 16S rDNA, and a nuclear gene, 18S rDNA, to infer the molecular phylogeny of the pyrgomatids. Our phylogenetic results allowed us to reject previous classifications of Pyrgomatidae based on morphological characteristics. Our results also suggested the possibility of paraphyly of the Pyrgomatidae. The hydrocoral barnacle Wanella is not found on the same clade as the other pyrgomatids, but rather, with the free-living balanids. The basal position of Megatrema and Ceratoconcha is supported. The archeaobalanid Armatobalanus is grouped with Cantellius at the base of the Indo-Pacific pyrgomatines. Fusion of the shell plate and modification of the opercular valves are homoplasious features that occurred more than three times on different clades. The monophyly of the "Savignium" group, comprising four nominal genera, is also not supported, and the different taxa are placed on different clades.     

Distribution and Abundance of Fauna on Living Tissues of Two Brazilian Hermatypic Corals (Mussismilia hispida (Verril 1902) and Siderastrea stellata Verril, 1868)

This study describes the distribution and abundance patterns of the associate fauna on the living surface of the corals Siderastrea stellata Verril, 1868 and Mussismilia hispida (Verril 1902) using a non-destructive method, on the northern coast of Rio de Janeiro State. For each coral species, infestation density and proportions of infested colonies, colonies attached and unattached to the substrate were estimated. A total of 474 colonies of S. stellata and 452 colonies of M. hispida were examined. The barnacle Ceratoconcha floridana (Pilsbry, 1931) was the dominant coral associate found, followed by gall-crabs of the family Cryptochiridae Paulson, 1875 and the bivalve Lithophaga bisulcata (d’Orbigny, 1842). Both coral species presented similar patterns of infestation dominance. S. stellata colonies were more commonly infested and showed a greater mean infestation density of 0.62 ind/cm² at Armação dos Búzios, whereas M. hispida colonies had infestation densities of only 0.20 ind/cm². Infestation density does not appear to impact negatively on corals of Armação dos Búzios. A clear negative relationship between the number of associates in the coral colony and coral size was found. Evidently abundance and frequency of occurrence of associated fauna is highly related to coral community structure and composition and the results highlight the importance of local scale studies.     

The benthic phase of the life cycle ofParapenaeus longirostris (Crustacea,Decapoda, Penaeidae) along the Mediterranean coast of Israel

The population ofP. longirostris along the Mediterranean coast of Israel spends the benthic phase of its life cycle (from body size over 15 mm) on muddy bottom deeper than 45 m. New age groups are recruited within the depth zone of 45–300 m and migrate in both inshore and offshore directions. Inshore migration is limited by unsuitable sandy ground. The limiting line for offshore migration was not found.An age group could be detected during one year within a body size range of TL = 40 mm to TL = 84.5 mm for males and to TL = 102.5 mm for females. Reproductive activity in shallow water, down to a depth of 73 m, takes place during the whole year, while in depths of 150–300 m there is an arrest of reproductive activity from June to August.     

Thoughts on the ecology and evolution of the intertidal biota of the Azores and other Atlantic islands

The special features of the intertidal ecosystems of remote islands are reviewed briefly before focusing on the littoral zone of Macaronesia (Azores, Madeira and Canaries). Distribution patterns are briefly compared with other European shores. Species missing from the Azores compared with Madeira, the Canaries and continental European shores are listed. The degree of genetic differentiation within and amongst selected important species of gastropods (Patella spp.) and barnacles (Chthamalus spp.) in Macaronesia is reviewed. The role of isolated islands in allopatric speciation of these organisms is discussed in relation to dispersal. Possible threats to the populations and communities of rocky shores in Macaronesia are considered in the context of the endemic nature of certain species. The importance of studies of basic ecology and genetics to inform resource management and conservation is highlighted. Future research directions are indicated, emphasizing the usefulness of Macaronesia as a model system to explore speciation in intertidal organisms.     

莱州湾强壮滨箭虫种群数量及个体大小季节分布

基于2011年5月至2012年4月(12月和翌年1—2月冰期除外)在渤海莱州湾逐月采集的数据资料,分析莱州湾强壮滨箭虫丰度空间分布和体长的季节变化,估算其生产力和摄食率,以掌握其种群动态特征,为评价该种在莱州湾生态系统结构功能中地位提供参数和依据。结果显示:强壮滨箭虫丰度和生物量年变化曲线为双峰型,最高峰出现于5月,次高峰出现于8—9月;4 mm和15 mm体长的个体分别是丰度和生物量的主要贡献者。生产力估值为0.25 mg C m~(-3)d~(-1),摄食率估值为0.88 mg C m~(-3)d~(-1)。种群月均体长以3—4月最高(10 mm)、5月和11月最低(5 mm);月均体长与月表层水温、盐度呈负相关。春季5月、夏季7—8月和秋季10—11月中,小型个体丰度所占的比例较高,应为种群数量增长期。各月的体长频数分布可分辨出平均体长5mm(Cohort Ⅰ)、介于5—10 mm(Cohort Ⅱ)和10 mm(Cohort Ⅲ)的3个同生群。Cohort Ⅲ和Cohort I高丰度区的表层水温和盐度分别介于15—17℃和29—32。Cohort Ⅲ基本不出现于水温大于24℃的站位,其丰度与水温呈显著负相关。Cohort Ⅰ基本不出现于水温低于11℃的站位,其丰度与水温呈显著正相关。Cohort Ⅱ高丰度区的表层水温和盐度为26—27℃和30—32。由上推测强壮滨箭虫发生明显体型更替的水温阈值应不超过15—17℃。     

Combined Effects of Seawater Temperature and Salinity on Development of the Larvae of the Rhizocephalan Peltogaster reticulatus (Crustacea: Cirripedia)

The effects of seawater temperature (12, 16, 20, 22, and 25°C) and salinity (of 8 to 34) in different combinations on the larvae of the rhizocephalan Peltogaster reticulatus (Crustacea: Cirripedia), a parasite of the hermit crab Pagurus proximus, were examined. The development of P. reticulatus is comprised of five naupliar stages and one cypris stage. Nauplii have a specific structure, the flotation collar encircling the dorsal side of the larval body. Larvae lack the pigmented nauplius eye, and they show no positive phototaxis. Successful naupliar development occurred in temperature and salinity ranges of 16–25°C and 20 to 34, respectively; but all nauplii died at 12°C and 16. The duration of each naupliar stage increased under lowering of the seawater temperature. At 22–25°C and 26–28, the entire development cycle was completed in 72–80 h; and at 16°C and 20 it lasted 153 h. The cypris larvae showed a greater resistance to decreased salinity in comparison with the nauplii. At temperatures of 16–25°C and salinities of 14 to 34, the lifespan of cyprids was 6 to 12 days, and it decreased at increasing temperature.