Distribution of Trypetesa lampas (Cirripedia,Acrothoracica) in various gastropod shells

The distribution of Trypetesa lampas (Hanoock) and its rates of infection of different species of gastropod shells in the waters of Anglesey were studied. T. lampas was found in depths from 3 to at least 25 fathoms (5 to 46 m). It infected dead shells, mainly of Buccinum undatum L., Colus gracilis (da Costa), Neptunea antiqua (L.) and Natica alderi (Forbes); it was absent from shells occupied by hermit crabs in the intertidal zone. Significantly more B. undatum shells were infected than shells of N. antiqua and Natica alderi, and more of both C. gracilis and N. antiqua than of Natica alderi. Larger shells of B. undatum were more likely to be infected than were the smaller ones. The number of females per shell varied with the shell length raised to the power 2.39 instead of as expected with the square of the length.     

Gametogenesis of Pollicipes cornucopia (Cirripedia: Scalpellomorpha) in north-west Spain

The gametogenesis and gonad microanatomy of the stalked barnacle Pollicipes cornucopia (Leach) is described. An histological classification of gametogenic development is proposed for the firt time for a species of the Cirripedia. This species has an annual gametogenic cycle on the north-west Spanish coast. Spawning takes place mainly from March to September. Ovaries release gametes at least twice during the gametogenic cycle.     

The unusual floatation collar around nauplii of certain parasitic barnacles (Crustacea: Cirripedia: Rhizocephala)

Nauplii of the rhizocephalan families Peltogastridae and Lernaeodiscidae carry a torus-shaped collar around the body. It consists of an exceedingly thin cuticle connected to the general body cuticle along a continuous narrow ridge. In nauplii of some species, the collar is very large and its surface ornamented by a very conspicuous reticulated pattern of ridges. In other species the collar is smaller and with a smooth surface that impedes its detection when using a light microscope. The collar is absent from nauplii of all investigated species of the Sacculinidae. Transmission electron microscopy shows that the collar of the succeeding nauplius instar is formed in an unexpanded state beneath the old cuticle and it must therefore be inflated at or immediately after ecdysis. At ecdysis the collar of the old instar breaks along the attachment ridge, leaving the empty collar and the exuvium of the general body as separate objects. The collar must have a profound influence on the hydrodynamic properties of the nauplius, both when swimming and passively sinking. We therefore consider it as a floatation device, a view supported by the absence of the collar in the rapidly swimming cypris larvae. There is no obvious homology to the collar outside the Rhizocephala, and it therefore appears to have evolved only once.Communicated by L. Hagerman, Helsingør     

Size regulation in larvae of the crustacean Balanus eburneus (Cirripedia: Thoracica)

We tested the hypothesis that larval size in the acorn barnacle Balanus eburneus Gould (Cirripedia Thoracica) varies in relation to food availability. In March–November 1980, and March–July 1981, larvae were obtained from adult Balanus eburneus collected in the Newport River, North Carolina, USA. Carapace length and width of larvae reared at three different food concentrations were measured. Mean naupliar instar size was independent of food concentration. Mean size of the cypris instar increased with increasing food level. Greater cypris size could be attributed to increased food reserves in the preceding naupliar stage, and was coinciden with inmarked increase in metamorphic success. Variation in instar size remained constant or declined during naupliar development, but increased sharply at the molt to the cyprid. Naupliar size regulation involved: (1) conservation of a molt increment specific for each naupliar-naupliar molt, (2) an inverse relationship between premolt size and the molt increment during the first five naupliar instars, and (3) an increase in the precision of the molt increment at the molt to the sixth naupliar instar. Experimental evidence implies that size regulation in Balanus eburneus limits variation about a fixed final naupliar size (e.g. volume). Measurement of naupliar size, accumulated energy reserves, survival and development time, and cypris metamorphic success indicated that naupliar cuticular growth is the most conservative feature of larval development. The data suggest that maximum naupliar size is limited by escalating metabolic costs during development, while minimum naupliar size is limited by size-related feeding effectiveness.     

The functional morphology and development of the alimentary tract of larval and juvenile barnacles (Cirripedia: Thoracica)

The alimentary tract of the nauplius larva of Balanus spp. consists of cuticle-lined foregut and hindgut, with intervening endodermal midgut constricted into anterior and posterior regions. The anterior midgut cells in the region of the constriction (constriction cells) secrete proteins (probably digestive enzymes). The remaining anterior midgut cells, often containing lipid droplets, form the absorptive region of the tract. Glycoprotein globules and lipid droplets within anterior midgut cells are the remants of the yolk in a pre-hatched larva, this yolk additionally supporting the larva through the non-feeding first nauplius stage. Nauplius Stages II to VI are actively feeding planktonic stages which increase in size and build up lipid reserves. These accumulated reserves support the non-feeding cyprid, first through its planktonic life and then through settlement and subsequent metamorphosis to the juvenile barnacle. Juvenile barnacles start to feed between 2 and 5 days after metamorphosis.     

Scanning electron microscopy of metamorphosis in four species of barnacles (Cirripedia Thoracica Balanomorpha)

Metamorphosis and early juvenile development was followed in the laboratory in Balanus amphitrite and B. improvisus (family Balanidae) and in Semibalanus balanoides and Elminius modestus (family Archaeobalanidae) from cyprid settlement until 24 h after ecdysis. Stages of development were studied in vivo and with scanning electron microscopy. Events in metamorphosis and early juvenile development are very similar in all four species and can be interpreted in terms of a highly modified moult. Notably, there was no indication whatsoever of the amorphous decorticated settler previously described from metamorphosis of B. amphitrite. The shape of a juvenile barnacle with cirri and incipient shell plates is apparent immediately after the shedding of the cypris carapace, and rudimentary peduncle can be distinguished below the developing wall plates. A basal row of cuticular hairs encircles the peduncle in all species except S. balanoides. These hairs seem to serve a restraining function during early development, when the juvenile barnacle is only attached by the initially secreted cyprid cement. Similarly situated hairs are also present in very young juveniles of the lepadomorph Scalpellum scalpellum. In Semibalanus balanoides nothing indicates that the rostrum originates from the fusion of two latera as previously claimed, since this plate is single as soon as it can be distinguished.     

Development of the gooseneck barnacle Pollicipes polymerus (Cirripedia: Lepadomorpha): Fertilization through settlement

Pollicipes polymerus (Sowerby, 1833) is a common barnacle in rocky intertidal zones of the West Coast of North America whose development has been previously undescribed in detail. In the present study, certain aspects of the development of P. polymerus are recorded from fertilization through settlement. In vitro fertilization has been accomplished using already-implanted sperm plus ovulating eggs and oviducal gland fluid. The eggs are small (about 100 μ diameter at fertilization; 90 x 140 μ at first cleavage) and somewhat yolky. Embryos obtained from brooding adults and from in vitro fertilization were reared in glass dishes until hatching. Cleavage was total, asynchronous and unequal. Division of the D macromere lagged behind division of the other blastomeres, and gastrulation occurred by epiboly so that blastoderm surrounded the macromeres. The nauplii hatched an average of 25.4 days (range was 20.3 to 30.1 days) after fertilization, and molted within 1 day to Stage II nauplii. From Stages II through VI the nauplii required food. As they grew from stage to stage, the number of setae per appendage increased. Nauplii have feathery and hispid-type setae, previously described only in Chthamalus aestuarii. In contrast to other pedunculate larvae which have been described, P. polymerus nauplii are small and have specialized feathery setae. Cypris-larva settlement was stimulated only when healthy adult peduncles were available, thus showing an affinity for its own species.     

Balanus amphitrite (Cirripedia: Thoracica) — A potential indicator of fluoride,copper, lead,chromium and mercury in North Adriatic lagoons

The capacity of the barnacle Balanus amphitrite Darwin to accumulate pollutants above ambient levels was examined in two North Adriatic lagoons with respect to fluoride, copper, lead, chromium and mercury. Levels in soft tissues ranged from 138 to 312 ppm (dry weight) for fluoride, 41 to 109 ppm for copper, 7.1 to 11.7 ppm for lead, 2.10 to 3.89 ppm for chromium, and from 0.96 to 1.35 ppm for mercury. The concentration factor was of the order of 10² for fluoride, 10³ for copper, and possibly higher than 10³ for lead, chromium and mercury. The differences of about 2:1 in the fluoride and copper concentrations in the waters of the two lagoons were clearly reflected by the levels in the barnacles. B. amphitrite that had set on experimental panels had levels similar to those found in specimens collected from long-term natural populations as early as 42 days after immersion of the panels. Compared with literature data, the accumulation levels found in B. amphitrite for fluoride were among the highest known for invertebrates; the values for copper and lead were considerable, but exceeded by others published for B. balanoides; the results for chromium and mercury were one order of magnitude lower than values reported for other suspension-feeders or indicator organisms. B. amphitrite appears to possess most of the properties considered essential for a biological indicator. Eventual determination of the response time of the barnacle to changes in environmental level could profitably be carried out utilizing experimental panels.     

Functional morphology of Lithotrya dorsalis (Cirripedia: Thoracica) in relation to its burrowing habit

Burrowing mechanisms of the scalpellid barnacle Lithotrya dorsalis collected from Indian Key, Florida, USA and Fort Point, Jamaica, West Indies, in the fall and spring of 1983 and 1984 were investigated. Calcite spicules covering the peduncule of this animal mechanically abrade carbonate substratum and are described by light and scanning electron microscopy. These spicules undergo a morphological transformation during the molt cycle. Initial observations on the burrowing habit of L. dorsalis indicate that it also employs a chemical agent during bioerosion. Characteristic etching patterns on both optically pure calcite (Iceland spar) and shell material lend further support to this contention. Possible sources of this chemical agent are discussed.     

Prolonged withdrawal: A possible predator evasion behavior in Balanus glandula (Crustacea: Cirripedia)

The duration of cirral withdrawal in Balanus glandula (Darwin) varies by a factor of three depending on the type of stimulus applied. Contact with potential predators including thaidid gastropods (Thais emarginata, T. lamellosa), and forcipulate asteroids (Leptasterias hexactis, Pycnopodia helianthoides) elicits significantly longer withdrawal durations than contact with an herbivorous gastropod (Tegula pulligo), a grazing, spinulosid asteroid (Henricia leviuscula) or a neutral, brown algal stimulus (Fucus distichus). By substantially attenuating the release of metabolites, prolonged withdrawal probably increases the likelihood of being bypassed by nonvisual predators relying on chemical cues to verify that barnacles are alive. The reduced response to the non-predatory species indicates that this is not a generalized response to gastropods or asteroids, but rather that it appears to be specific to potential predatory species.     

Reproduction in Balanus amphitrite Darwin (Cirripedia: Thoracica): influence of temperature and food concentration

Balanus amphitrite, an acorn barnacle, is distinctly euryhaline, eurythermal and a dominant fouling organism found in warm and temperate waters throughout the world. In this study, the influence of temperature and food concentration on the reproductive biology of this species collected from a tropical habitat was evaluated. Adult barnacles were maintained at 20, 25 and 30°C temperatures at different concentrations of food (50, 100, 150 and 200 Artemia ind⁻¹ day⁻¹). In this previously believed obligatory cross-fertilizing hermaphrodite, self-fertilization was observed. The rise in temperature from 20 to 30°C resulted in a longer interbreeding interval (6–7 days, 200 Artemia ind⁻¹ day⁻¹; 11–13 days, 50 Artemia ind⁻¹ day⁻¹). Computed carbon gained through feeding during the interbreeding interval indicated an inverse relationship to the temperature. At 20°C, although a greater amount of carbon was gained through feeding, the numbers of larvae produced were fivefold less when compared to those raised at 30°C. At 20°C, 2.3 μg C was required to produce a single larva, whereas at 30°C it was 0.4 μg C. A rise in rearing temperature also influenced the molting rate positively. Observations on temporal variation in the gonad development of this species in a tropical coastal environment influenced by the monsoons indicated gonad development to be positively related to chlorophyll a concentration.     

Interactions between shear rates and biofilms affecting exploratory behaviour by cyprids ofElminius modestus (Cirripedia)

The behaviour ofElrninius modestur Darwin cyprids suggests interaction between surface bacterial films acclimated to contrasting shear regimes and the range of shear rates over which observations were made. Exploratory behaviour is encouraged by filmed surfaces rather than biologically clean surfaces. Increasing shear rates encourage attachment to all surfaces, but cyprids detach from high-shear-acclimated films less often than from either low-shear-acclimated films, or clean surfaces. Furthermore, numbers of attaching cyprids are consistently lower on low-shear-acclimated films than on high-shear films.     

An energy budget for the free-swimming and metamorphosing larvae of Balanus balanoides (Crustacea: Cirripedia)

Analysis of biochemical components and measurements of oxygen consumption rates of cypris larvae of Balanus balanoides (L.) maintained in the laboratory at 10°C for up to 5 weeks after capture shows that lipid is the primary energy reserve, although later protein is utilised. Initially, the cyprids swim freely with an oxygen consumption rate of ca. 37×10^-3 l O₂ h^-1 cyprid^-1, but within a few days the rate falls to ca. 21×10^-3 l O₂ h^-1 cyprid^-1 when they cease swimming and explore the substratum. The cost of metamorphosis was calculated both from the loss of biochemical components and oxygen consumption rates during metamorphosis; the values were 2.8×10^-2 and 3.2×10^-2 cal cyprid^-1, respectively. A budget was collated from the data on respiration and biochemical composition, whereby the energy per cyprid was partitioned into that required for essential structural components (6.8×10^-2 cal), that needed for metamorphosis (3.0×10^-2 cal) and an excess available for swimming and exploring, which in the batches studied was about 5.0×10^-2 cal. This excess is mainly derived from the utilisation of lipid reserves and is used up usually 2 1/2 to 4 weeks after capture. During these measurements, samples of cyprids were taken at weekly intervals to test the rate of settlement and success of metamorphosis. The results showed that they lose their competence to metamorphose successfully approximately at the same time (3 to 4 weeks) that the energy supply for swimming and exploration is used up.     

Stable carbon isotope ratios in Red Sea barnacles (Cirripedia) as an indicator of their food source

The stable isotope ratio ¹²C/¹³C was used to investigate the source of carbon in free-living barnacles and in coral-inhabiting barnacles from the Red Sea. The δ¹³C of most of the barnacles collected on the open shore ranges between −17.5 and −19.7‰, indicating relative enrichment of light carbon originating from the open-sea phytoplankton. Those collected in closed habitats showed heavier isotopic composition. The δ¹³C of the coral-inhabiting barnacles ranges from −14.1 to −16.7‰, suggesting that the carbon contribution of open-sea plankton to these barnacles is less important than it is to free-living barnacles. We hypothesize that coral organic matter and zooxanthellae expelled by the host coral contribute carbon to the barnacle, and that a mixture of this relatively heavy carbon with carbon from other sources is responsible for the high values of δ¹³C in coral barnacles. Received: 28 February 1997 / Accepted: 16 September 1997     

Settlement behavior and antennulary biomechanics in cypris larvae of Balanus amphitrite (Crustacea: Thecostraca: Cirripedia)

The settlement behavior, antennulary biomechanics and morphology were investigated in the intertidal cypris larvae Balanus amphitrite. The very maneuverable antennule of the cyprid consists of four jointed segments, and it is the prime appendage for substratum exploration and final settlement. We recorded several previously undescribed settling behaviors, such as the "tight direction change", for this intertidal species and discovered that the antennulary morphology is modified to facilitate such behaviors. Segment 2 has a modified lateral cuticle, which gives the segment an additional higher degree of maneuverability compared to non-intertidal cirripede species of Rhizocephala. Additional supporting muscle groups were also found in the antennule of our species, which were not found in the investigated rhizocephalan larvae. The results could very well be associated with the higher needs of substratum recognition in intertidal species, due to more heterogeneous habitats such as intertidal rock flats.     

Some observations of factors affecting embryonic and larval growth of Pollicipes polymerus (Cirripedia: Lepadomorpha) in vitro

Observations on the effects of several environmental conditions in embryonic and larval cultures are reported for the gooseneck barnacle Pollicipes polymerus (Sowerby, 1833). When the growth rates of control embryos kept in finger bowls at ambient-temperature sea water (ca. 12°–15°C) were compared to those of embryos grown under variously modified circumstances, it was found that growth rates were faster under conditions closely simulating the adult barnacle mantle-cavity (e.g. darkness and aeration). Addition of antimicrobial drugs and reduction of egg-mass size also promoted fast growth and development. Nauplii were fed several species of algae; only three species promoted growth to Stage V or older. Naupliar growth was fastest in larvae fed the combination of Prorocentrum micans/Platymonas sp. Larvae fed Prococentrum micans/pennate diatom grew slowest, and most larvae died at Naupliar Stage III. Interactions between algal species may have affected their nutritional value for Pollicipes polymerus nauplii. Less larval activity, slower growth rates, and higher mortality rates were observed in individuals given small amounts of food. The development of embryos and larvae in culture depends greatly on the culture conditions. These conditions should be described if comparison of timetables and envents are to be made between studies.     

Genetic structure of populations of two species of Chthamalus (Crustacea: Cirripedia) in the north-east Atlantic and Mediterranean

Protein electrophoresis on starch gels was used to investigate population genetic structure of the barnacles Chthamalus montagui Southward and C. stellatus (Poli) over their north-east Atlantic and Mediterranean ranges. In each species, a single locus exhibited marked differentiation of allele frequencies between Atlantic and Mediterranean localities; in C. stellatus, genetic differentiation between the two basins had not previously been noted. In both species, mean heterozygosity per locus appeared higher in the Mediterranean samples than in the Atlantic, and Mediterranean populations had more alleles at the loci studied. Possible explanations for the differentiation between the Atlantic and the Mediterranean populations are discussed. Received: 30 May 1996 / Accepted: 17 September 1996     

Variability of shell growth and morphology of the wall-plate junctions of the intertidal barnacle Tesseropora rosea (Cirripedia: Tetraclitidae)

Variability in shell growth and morphology of the wallplate junctions of Tesseropora rosea (Krauss) were studied in 1983 in New South Wales, Australia, at two sites with differing wave exposure. The wall-plate junctions, a combination of dado (geometrically interlocking) and lap bints, increase in complexity with age, but do not differ between sites. The number of parietal tubes is fixed at settlement and does not differ between sites, but differs among wall plates in a manner consistent with their evolutionary history (rostrum > laterals > carina). The number of secondary septa increases with age, but does not differ between sites. In epilithic T. rosea, growth of the alae contributes to orifice enlargement. Epizoic T. rosea (on limpets) have conspicuous radii and display diametric growth. At the more exposed site, the shell wall was sleeper, and the aperture length, height and weight of the shell increased at a faster rate than at the less exposed site, but the maximum height of the shell was less. These differences appear to be due to a combination of greater feeding rates and greater erosion rates at the exposed site. The greater maximum height attained at the more sheltered site may be due to greater individual longevity. Juveniles at the sheltered site had thinner septa in the laterals than those at the exposed site. Septa in adults were rhinner at the exposed site, but this is due to age differences between sites in the adult populations.