Bewuchsuntersuchungen auf Natursteinsubstraten im Gezeitenbereich des Nordsylter Wattenmeeres: Algen

Zusammenfassung 1. Zur Analyse der Besiedlungsdynamik und der Substratabhängigkeit makroskopischer benthischer Pflanzen wurden 6 verschiedene Substrate unter In-situ-Bedingungen untersucht: Solnhofener Plattenkalk, Muschelkalk, Buntsandstein, Granit, Säulenbasalt und Basaltlava. Ausgewählt nach Farbe, Korngröße und Relief wurden die Substrate in 3 Wasserbereichen (Hochwasser-, Niedrigwasser- und Unterwasserbereich) vertikal an einem Gerüst am Hafen von List/Sylt angebracht und während zweier aufeinander folgender Jahre regelmäßig kontrolliert.2. Physikalische Substrateigenschaften beeinflußten die Besiedlung stärker als chemische. Erstere wirkten sich in den drei Wasserbereichen — unabhängig von anderen Umweltfaktoren — jeweils verschieden auf den Bewuchs aus. Auf dem Hochwasserniveau (Trockenfall-Perioden von durchschnittlich 8 Std. während eines Gezeitenganges) waren vor allem folgende Faktoren siedlungsbegrenzend: a) Temperatur des Substrates (abhängig von Farbe, Wärmeverlust durch Verdunstung), b) Wassergehalt (abhängig von Gesteinsporosität), c) Stärke des Reliefs (Schutz vor Abrasion der Organismen) und Oberflächenrauhigkeit (Stabilität des Anheftens), d) Widerstandsfähigkeit des Substrats selbst gegen Erosion und Abrasion. Auf dem Niedrigwasserniveau (Trockenfall-Perioden von durchschnittlich 1 Std. während eines Gezeitenganges) wirkten besonders in der Brandungshohlkehle mechanische Kräfte auf die Algenentwicklung ein, während sich der Einfluß von Oberflächentemperatur und Wassergehalt der Gesteine verringerte. Auf dem Unterwasserniveau (keine Trockenfall-Perioden) verloren die physikalischen Eigenschaften der Substrate an Bedeutung, vermutlich infolge der geringen Temperaturschwankungen und der meist abgeschwächten Turbulenz. Biologische Faktoren bestimmten in zunehmendem Maß das Bewuchsmuster.3. Die Korngröße war — unabhängig vom Härtegrad des Materials — entscheidend für Ausdehnung und Bewuchsdichte einiger Algen.Enteromorpha sp. entwickelte sich optimal auf fein- bis mittelkörnigen Substraten (Solnhofener Plattenkalk und Buntsandstein), weniger dicht auf dem vorwiegend feinkörnig auskristallisierten Gesteinsfluß des Säulenbasalts und den feinkörnigen Bestandteilen des Muschelkalks. DasCodiolum-Stadium einerUrospora-Art überzog ausschließlich die glatte und feinkörnige Fläche von Säulenbasalt, Solnhofener Plattenkalk und entsprechend feinkörnige Stellen im Muschelkalk.Porphyra sp., meistPorphyra purpurea, bevorzugte die grob strukturierten Substrate Granit, Muschelkalk und die durch Poren zerrissene Oberfläche der Basaltlava. Säulenbasalt wurde an den Stellen besiedelt, wo größere Einsprenglinge den gleichartigen, feinkörnigen Gesteinsfluß durchsetzten.Ceramium rubrum reagierte in gleicher Weise wiePorphyra sp. Mit Hilfe von Parallelbeobachtungen an aus Quarzsand 5 abgestufter Korngrößen hergestellten Substraten wurde eine Korngrößenabhängigkeit der Algenbesiedlung ermittelt. Die ausgewählten Fraktionsbereiche (< 0,25; 0,25–0,5; 0,5–1,0; 1,0 bis 2,0; 2,0–4,0 mm) konnten in etwa denen der Natursteine gegenübergestellt werden. Unterhalb von 0,5 mm Korndurchmesser entwickelten sich ausschließlichEnteromorpha-Arten, oberhalb davonPorphyra-Arten undCeramium rubrum. Die Zahl der siedelnden Rotalgen erhöhte sich mit zunehmender Korngröße. Die Versuchsergebnisse der Sandsubstrate entsprachen denen der gegenübergestellten Natursteine ähnlicher Fraktionsbereiche und wiederholten sich in den 3 Horizonten.4. Dichte und Porosität der Substrate bestimmten im Hochwasserbereich nicht nur das Bewuchsmuster, sondern auch die obere Grenze der Algenvegetation. Die obere Grenze befand sich bei Muschelkalk (Zurückhalten des Wassers in der zerklüfteten Oberfläche) und Basaltlava (Wasserspeicherung im Porenraum) über der Mitteltiden-Hochwasserlinie. Bei dem dichten Gestein Säulenbasalt wurde die Mitteltiden-Hochwasserlinie vom Bewuchs nicht mehr erreicht. Im Frühling und Herbst dagegen entwickelte er sich auf Säulenbasalt bei günstigen Lufttemperaturen und Wasserständen bis fast zur gleichen Höhe wie auf Basaltlava. Die obere Grenze derEnteromorpha-Besiedlung befand sich auf dem wasserspeichernden Buntsandstein höher als auf Solnhofener Plattenkalk mit seiner geringen Porosität. Auf Granit erreichte der Algenbewuchs die niedrigste vertikale Ausdehnung.5. Abhängig von der Farbe bestimmte der Erwärmungsgrad eines Substrates die vertikale Ausdehnung der Vegetation. Im Bereich der Mitteltiden-Hochwasserlinie verschwanden die Algen von dunklen Substraten (Säulenbasalt, Granit), so wie die Wärmestrahlung im Juli/August zunahm und entwickelten sich von neuem im Herbst bei günstigeren Umweltbedingungen.6. Das Zusammenwirken von Erosion und Abrasion wurde — ohne Frostsprengung — am Solnhofener Plattenkalk deutlich. Dünne Substratschichten hoben sich plattig zusammen mit Bewuchs von der Oberfläche ab. Im Hochwasser- und Niedrigwasserbereich machte sich eine Abtragung durch Wellenbewegung stärker als auf dem Unterwasserniveau bemerkbar.

---

Fouling studies on natural-stone substrates in the tidal zone of the north sylt wadden sea: Algae

Six different types of test substrates were exposed in the tidal zone of the wadden sea near the harbour of List (Island of Sylt, North Sea): Solnhofen limestone, Middle Triassic limestone, Bunter sandstone, granite, basalt, and basaltic lava. The test substrates were fixed to a panel at the midtide to high-water level, the midtide to low-water level, and 75 cm below the latter (sublittoral level); they were arranged in such a way, that substrates with fine granulation alternated with those of rough granulation. A defined boundary between the growth zones ofEnteromorpha sp. (green algae) andPorphyra purpurea (red algae) was observed at the midtide to low-water level.Enteromorpha sp. was found on fine-granulated substrates (chalk of Solnhofen, mottled sandstone).Porphyra purpurea settled preferably on rough granulated substrates (limestone, granite, and basaltic lava). With increasing irregularity of the substrate-surface structure, the abundance of red algae increased, especially on basaltic lava. In order to examine granulation effects more closely, chemically equivalent artificial solid substrates were installed under identical environmental conditions. The material chosen consisted of quartz. Grain size fractions of 0.25 mm, 0.25–0.5 mm, 0.5–1.0 mm, 1.0–2.0 mm, 2.0–4.0 mm were fixed to panels, providing five artificial substrates of increasing surface roughness. The boundary between the growth of green algae and red algae was found at 0.5 mm.Enteromorpha sp. settled on substrates with granule size < 0.5 mm;Porphyra purpurea on substrates > 0.5 mm. The population density of the red algae increased with increasing granule size. These observations led to the assumption that the surface structure of the test substrate exerts more influence on algal settlement than substrate hardness or chemical composition.

     

Vicious circle in the intertidal: Facilitation between barnacle epibionts, a shell boring polychaete and trematode parasites in the periwinkle Littorina littorea

We studied interactions between three organisms associated with a common gastropod of northern Atlantic shores, the periwinkle Littorina littorea: barnacle epibionts Balanus crenatus, a shell boring polychaete Polydora ciliata, and tissue invading trematodes which use the periwinkles as first intermediate host. Snails collected shortly after barnacle settlement with > 50% cover of barnacles had significantly higher infestation of shell boring worms compared to unfouled snails, while trematode infestation was similar. The result was the same at two sites, and we conclude that the worm P. ciliata facilitates barnacle fouling on snails. The reverse was also the case. In an experiment with 14 weeks of exposure, snail treatments with barnacle epibionts had a significantly higher P. ciliata load than unfouled and cleaned snails. Again, trematode infestations were similar. The reciprocal positive interactions between barnacle epibionts and shell boring worms on snail houses is regarded as a case of facultative mutualism. On the other hand, for the snail basibiont, both barnacles and shell boring worms exert strongly negative effects by reducing fecundity, growth, and survival, resulting in a vicious circle for the snails. The combined effects of these associated organisms may rival in importance any competitive or predacious effects on the host L. littorea.     

The effect of surface colour on the adhesion strength of Elminius modestus Darwin on a commercial non-biocidal antifouling coating at two locations in the UK

A number of factors affect the adhesion strength of organisms to fouling-release coatings, and except for a few studies focussing on black or white surfaces none have dealt specifically with the effect of coating colour. The aim was to test the effect of colour on the adhesion strength of the barnacle Elminius modestus. Panels coated in six commercial colours of Intersleek 700® were submerged at two field sites and barnacles were pushed-off using a standard assay procedure. The strength of adhesion (SOA) varied between and within sites for colour and by barnacle basal area, SOA per unit area being higher for smaller barnacles. Higher SOA with a small basal area may be because of size-specific predation, differential hydrodynamic effects or adhesive failure with age. The complex effect of colour on barnacle adhesion may be because of physico-chemical surface characteristics varying with pigments, and their interactions with local environmental conditions, as well as interactions with the settling barnacle larvae.     

In situ ATR–FTIR characterization of primary cement interfaces of the barnacle Balanus amphitrite

A method is presented for characterizing primary cement interfaces of barnacles using in situ attenuated total reflection–Fourier transform infrared spectroscopy. Primary cement of the barnacle, Balanus amphitrite (Amphibalanus amphitrite), was characterized without any disruption to the original cement interface, after settling and growing barnacles directly on double sided polished germanium wafers. High-quality IR spectra were acquired of live barnacle cement interfaces, providing a spectroscopic fingerprint of cured primary cement in vivo with the barnacle adhered to the substratum. Additional spectra were also acquired of intact cement interfaces for which the upper portion of the barnacle had been removed leaving only the base plate and cement layer attached to the substratum. This allowed further characterization of primary cement interfaces that were dried or placed in D₂O. The resulting spectra were consistent with the cement being proteinaceous, and allowed analysis of the protein secondary structure and water content in the cement layer. The estimated secondary structure composition was primarily β-sheet, with additional α-helix, turn and unordered components. The cement of live barnacles, freshly removed from seawater, was estimated to have a water content of 20–50% by weight. These results provide new insights into the chemical properties of the undisturbed barnacle adhesive interface.     

Differential growth of the barnacle Notobalanus flosculus (Archaeobalanidae) onto artificial and live substrates in the Beagle Channel, Argentina

In the Beagle Channel, southern South America (ca. 55°S 67°W), about 20% of false king crabs (Paralomis granulosa) >80 mm carapace length are fouled with the barnacle Notobalanus flosculus. To evaluate differences in growth rates of barnacles attached to artificial and live substrates, clay tiles were anchored as collectors to the bottom at two different sites in the Beagle Channel in September 1996: in Ushuaia harbour (low currents and high levels of suspended matter) and around the Bridges Islands (strong currents and low level of suspended matter). Another set of collectors was deployed at the same sites in October 1998 to collect barnacles for histological studies. Tiles were removed from each place, approximately, on a monthly basis. Carapaces of P. granulosa with the epizoic N. flosculus were sampled between November 1996 and 1997, and between March 1998 and September 1999, to study sexual maturation of barnacles. Growth of barnacles was compared between the collectors and P. granulosa carapaces following a qualitative approach. A sexual maturity scale was defined, based on the stage of development of the female reproductive apparatus of N. flosculus. Growth rate of barnacles was highest in the harbour, intermediate on P. granulosa, and lowest around the Bridges Islands. Presence of oocytes was registered only in epizoic barnacles, suggesting that at least a proportion of these individuals is able to spawn on the carapaces. The potential advantages of settling on a living substrate, namely increased availability of food particles and decreased predation risks are discussed.     

Direct and indirect effects of Littorina littorea (L.) on barnacles growing on mussel beds in the Wadden Sea

On the extensive sedimentary tidal flats of the Wadden Sea, beds of the blue mussel Mytilus edulis represent the only major hard substratum and attachment surface for sessile organisms. On this substratum, the barnacle Semibalanus balanoides is the most frequent epibiont. In summer 1998, it occurred on over 90% of the large mussels (>45 mm shell length) and the dry weight of barnacles reached 65% of mussel dry weight. However, the extent of barnacle overgrowth is not constant and differs widely between years. Periwinkles (Littorina littorea) may reach densities >2000 m^–2 on intertidal mussel beds. Field experiments were conducted to test the effect of periwinkle grazing on barnacle densities. An experimental reduction of grazing and bulldozing pressure by periwinkles resulted in increased recruitment of barnacles, while barnacle numbers decreased with increasing snail density. The highest numbers of barnacles survived in the absence of L. littorea. However, a lack of periwinkle grazing activity also facilitated settlement of ephemeral algae which settled later in the year. Field experiments showed that the growth rate of barnacles decreased in the presence of these ephemeral algae. Thus, L. littorea may reduce initial barnacle settlement, but later may indirectly increase barnacle growth rate by reducing ephemeral algae. It is suggested that periwinkle density may be a key factor in the population dynamics of S. balanoides on intertidal mussel beds in the Wadden Sea.     

Desiccation,predation, and mussel-barnacle interactions in the northern Gulf of California

Summary Field experiments were conducted in order to determine the potential for desiccation and predation to mediate the effect of mussels (Brachidontes semilaevis) on barnacles (Chthamalus anisopoma) in the highly seasonal northern Gulf of California. We did this by removing both mussels and a common mussel predator (Morula ferruginosa: Gastropoda) and by spraying selected sites with sea water during summertime spring low tides. We also determined the effect of crowding on resistance to desiccation in barnacles, and the effect of barnacles on colonization by mussels. The mussel-barnacle community was not affected by keeping experimental quadrats damp during daytime low tides throughout the summer. Exposure to summertime low tides, however, did affect the survivorship of isolated, but not crowded, barnacles; and barnacle clumps enhanced the recruitment of mussels. Hence crowding in barnacles had a positive effect on both barnacle survivorship and mussel recruitment. Morula had a negative effect on mussel density, and mussels had a negative effect on barnacle density. The effect of Morula on barnacle density was positive, presumably due to its selective removal of mussels. These results suggest an indirect mutualism between barnacles and the gastropod predator, because barnacles attract settlement or enhance the survival of mussels, and the predator reduces the competitive effect of mussels on barnacles.     

Settlement Dynamics of the Encrusting Epibenthic Macrofauna in Two Creeks of the Caeté Mangrove Estuary (North Brazil)

The epibenthic encrusting fauna of 2 creeks of the Caeté mangrove estuary, northern Brazil, was studied over a 13 month period using collectors fixed at 2.5 and 3.5 m above the creek bottom and in which upper and lower sides of ceramic and wooden panels were used as settlement substrates. The number of individuals of the most abundant organisms (barnacles, oysters and mussels) settling per panel was determined each month, for each substrate type, panel orientation and height above creek bottom. The barnacle, Fistulobalanus citerosum has a peak settlement period during the wet season whereas both peaks in the numbers of settlers of the oyster Crassostrea rhizophorae were recorded during the dry season and such discrete temporal patterns in settlement have also been observed for barnacles and oysters in other mangroves and estuaries. In contrast to other studies, settlement of the mussel Mytella falcata was generally low during the study period and may be related to over-exploitation of stocks in the region. Overall, settler density was usually greater on the underside of ceramic panels close to the creek bottom, similar to results of other studies of epibenthic settlement in diverse habitats.     

New Genus and Species of Chirotheroid Tracks in the Detfurth-Formation (Middle Bunter,Lower Triassic) of Central Germany

Archosaur tracks are described that have been found in 1999 in a single track bed in the Dethfurth-Formation of the Middle Bunter (Lower Triassic) near Wolfhagen (Central Germany). The taxonomic evaluation of these tracks requires the erection of the new genus and species Protochirotherium wolfhagense ichnogen. n. ichosp. n. Apart from this a rearrangement of groups of chirotheroid tracks is proposed due to the digital proportions.     

Host‐specific acrothoracid barnacles on Middle Devonian platyceratid gastropods

Borings, attributed to acrothoracic barnacles, occur on the platyceratid gastropod Naticonema lineatum (Conrad) from the Middle Devonian Hamilton Group of western New York and rarely in specimens as old as the Early Devonian. These latter are the oldest known acrothoracid borings are in the fossil record. The borings are consistently developed as laterally compressed, inequilateral pouches exclusively on these gastropods, commonly as dense infestations.

Naticonema shells yielding borings typically occur associated with partially articulated qrinoid remains, and they are sometimes found attached to crinoids in a manner similar to coprophagous Platyceras. In addition to barnacle borings, Naticonema shells often bear thin encrustations of bryozoans which are usually perforated by these borings but sometimes also overgrow them. Barnacles bored live hosts; gastropods prevented shell penetration by producing cyst‐like secondary secretions of calcite beneath acrothoracid boreholes.

The relative antiquity of these borings and their association with coprophagous platyceratids makes their discovery particularly significant in revealing aspects of the early ecology of barnacles. Attachment to the host commensal gastropods was one of the first successful life modes of these crustaceans prior to their later diversification to other habitats. Mississippian and Pennsylvanian occurrences of similarly bored gastropods demonstrate continuity of the barnacle‐gastropod‐crinoid ecological association from the Middle to Late Paleozoic.     

Lepadiform and scalpelliform barnacles from the Oligocene and Miocene of the Paratethys sea

Abstract: A review on the Oligocene and Miocene stalked barnacles of the Paratethys Sea is presented. The fauna comprises two lepadiform and eight scalpelliform species. Only two species are known from Oligocene deposits, four are documented from Lower Miocene formations, and another four species occur in the Middle Miocene. Only one Oligocene and one Miocene species occur in two different Paratethyan basins in roughly coeval environments, and only a single Middle Miocene species is also known from the adjacent Mediterranean Sea. This low degree of biogeographical similarity is probably a result from the still low number of publications dealing with this group. Stalked barnacles are often indicating bathyal or at least deep sublittoral environments. Especially, the abundance of the benthic deep‐water lepadiform Poecilasma in evaporitic deposits of the Middle Miocene Badenian Salinity Crisis may serve as new evidence for a deeper marine depositional environment. Scalpellum paratethyianum sp. nov. and Lepas sattmanni sp. nov. are described as new species; Arcoscalpellum renevieri (Mayer and Gümbel in Gümbel, 1861) is introduced as new combination.     

‘Flying barnacles’: implications for the spread of non-indigenous species

The presence of adult barnacles of Fistulobalanus pallidus (Darwin) and Fistulobalanus albicostatus (Pilsbry) attached to field-readable plastic leg rings on the Lesser Black-backed Gull Larus fuscus in Northern Europe is reported. L. fuscus is a long-distance palaearctic migrant, breeding in temperate areas spreading widely over inland and marine habitats outside the breeding season. The species is known to perform long-distance migration to Africa and the Middle East. Combining present knowledge on the birds' migratory pattern and the home range of the barnacle species, it is concluded that the cypris larvae of F. pallidus must have settled in African waters, whereas the area where F. albicostatus settled on the bird leg rings is less certain. The barnacles were of adult size and must thus have been attached for a period of no less than 2 months. More than 30 individual barnacles could occur together on a single field-readable plastic leg ring. The barnacles could therefore, if ported alive to a new area, reproduce successfully and thus either introduce the species or genetically affect other native populations. This may pose a new and wholly unexpected transportation pathway for barnacles as invasive species.     

The symbiotic relationship between the hydrocoral Millepora dichotoma and the barnacle Savignium milleporum

Translocation of radioactive ¹⁴C and ³²P between the pyrgomatine barnacle Savignium milleporum and the hydrocoral Millepora dichotoma in the Red Sea was investigated in order to discover any mutual nutritional benefits. Translocation of photosynthetic products from endosymbiotic zooxanthellae to the hydrocoral was demonstrated. There was no evidence that carbon was further translocated to the barnacle. However, hydrocorals bearing barnacles accumulated significantly more ¹⁴C and ³²P than those with no barnacles. The possibility that the hydrocorals recycle substances excreted by the barnacles is discussed in the context of the oligotrophic environment of the Red Sea.     

The Impact of Desiccation on the Adhesion of Barnacles Attached to Non-stick Coatings

Fouling-release coatings prevent fouling of ships' hulls through hydrodynamic forces generated as the ship moves through the water. The effectiveness of such coatings may be evaluated by measuring the adhesion strength of settled organisms, e.g. barnacles. The influence of desiccation of the barnacle adhesive on such measurements was investigated. Shear forces required to remove barnacles of the genus Balanus increased during the course of desiccation up to the point when the barnacles suddenly self-detached. The increase was thought to be due to the rising cohesive strength of the adhesive. Growing tensile forces within the weakly cross-linked adhesive, however, are suggested to have led to self-detachment. The shear forces required to remove barnacles of the genus Elminius were generally low and did not differ significantly during the course of desiccation. The different results may be attributed to specific base morphologies. It was concluded that measuring the adhesion strength of members of the Balanidae on non-stick surfaces in air could produce flawed results due to the influence of desiccation of the barnacle adhesive. The investigations have also provided new insights into the characteristics of barnacle adhesive.     

Distribution and abundance of the acmaeid limpet,Patelloida latistrigata,and its interaction with barnacles

Summary At Cape Banks, New South Wales, adults of the small intertidal limpet, Patelloida latistrigata occur exclusively in the barnacle zone, and are primarily associated with the barnacle, Tesseropora rosea. Limpet density increases with barnacle density. Juvenile limpets can be found throughout the barnacle zone, and on patches of bare rock that may be temporarily available at lower levels on the shore. The failure of juveniles to survive and grow in places other than among Tesseropora is due to a combination of factors. These include desication at high levels on the shore, smothering by rapidly growing algae low on the shore, and the grazing activities of the larger limpet, Cellana tramoserica. These latter two factors also reduce the survival of experimentally transplanted adult Patelloida: algae by covering the substratum and smothering the limpets, and Cellana by outcompeting them for food. The density of Cellana is greater on patches of bare rock than among barnacles, and these large limpets may be unable to move and feed effectively over the irregular surface created by Tesseropora. Patelloida, however, is small enough to feed over and among these barnacles, and hence has a refuge from competition with Cellana. Barnacles may also provide shelter from the effects of desiccation and strong wave action, and thus increase the survival of juvenile Patelloida. By being associated with barnacles, however, Patelloida becomes vulnerable to intermittent predation by the whelk, Morula marginalba. This effect may be serious enough to eliminate small local populations of limpets, either by direct predation or by removing the refuge-providing barnacles. The association with barnacles may also limit the maximum size to which Patelloida can grow. Patelloida is not always found with Tesseropora, and adults in different localities can be found in association with other sessile organisms. It may be argued that small species of limpets require a spatial refuge from physical and/or biological pressures. To examine this hypothesis, the relationship between Patelloida and Tesseropora is compared to other published accounts of limpets with specialised modes of life.     

Increasing precision in randomised field experiments: barnacle microtopography as a predictor of Littorina abundance

We investigated whether measurements of barnacle microtopography could be used as covariates to increase precision in randomised field experiments with intertidal Littorina populations. We used image analysis to quantify the microtopography of the barnacles within 100 cm2 quadrats in July 1994 using as variables: number of barnacles, number of dead empty barnacles, mean distance to nearest neighbouring barnacle, total area covered by barnacles, and number of pockets of different size-classes [extra-small (< 0.15 cm2), small (0.15 and < 0.30 cm2), medium ( 0.30 and < 0.45 cm2), large ( 0.45 and < 0.60 cm2), and extra-large (0.60 cm2)] at least 75% surrounded by barnacles. We then used these variables to predict the abundance of Littorina spp. in the same quadrat. Two variables: the number of extra-small pockets and the number of small pockets accounted for 50% of the variation in total littorinid snail abundance among quadrats for January 1995 and for 47% of the variation for August 1994 but for only 16% of the variation for June 1994. However the single variable, the number of barnacles, accounted for 29–36% of the variation in snail abundance on these dates and can be measured without special equipment. The best combination of covariates was the number of extra-small pockets alone. The number of extra-small pockets between barnacles may be influencing the snail abundance by providing a refuge from wave shock and heat stress/desiccation. We suggest that the availability of these refuges among the barnacles could act as a density-dependent regulator of the total population size of a particular species of Littorina.     

Speciation Versus Phenotypic Plasticity in Coral Inhabiting Barnacles: Darwin's Observations in an Ecological Context

Speciation and phenotypic plasticity are two extreme strategic modes enabling a given taxon to populate a broad ecological niche. One of the organismal models which stimulated Darwin's ideas on speciation was the Cirripedia (barnacles), to which he dedicated a large monograph. In several cases, including the coral-inhabiting barnacle genera Savignium and Cantellius (formerly Pyrgoma and Creusia, respectively), Darwin assigned barnacle specimens to morphological ``varieties' (as opposed to species) within a genus. Despite having been the subject of taxonomic investigations and revisions ever since, the significance of these varieties has never been examined with respect to host-associated speciation processes. Here we provide evidence from molecular (12S mt rDNA sequences) and micromorphological (SEM) studies, suggesting that these closely related barnacle genera utilize opposite strategies for populating a suite of live-coral substrates. Cantellius demonstrates a relatively low genetic variability, despite inhabiting a wide range of corals. The species C. pallidus alone was found on three coral families, belonging to distinct higher-order classification units. In contrast, Savignium barnacles exhibit large between- and within-species variations with respect to both micromorphology and DNA sequences, with S. dentatum ``varieties' clustering phylogenetically according to their coral host species (all of which are members of a single family). Thus, whereas Savignium seems to have undergone intense host-associated speciation over a relatively narrow taxonomic range of hosts, Cantellius shows phenotypic plasticity over a much larger range. This dichotomy correlates with differences in life-history parameters between these barnacle taxa, including host-infestation characteristics, reproductive strategies, and larval trophic type. Received: 18 January 1999 / Accepted: 9 May 1999     

Symbiotic association of a pyrgomatid barnacle with a coral from a volcanic middle Miocene shoreline (Porto Santo,Madeira Archipelago,Portugal)

Abstract: The bioclaustration of the pyrgomatid barnacle Ceratoconcha aff. costata within the carbonate skeleton of a colonial hermatypical coral (Tarbellastrea reussiana) is described from a middle Miocene basaltic rocky palaeoshore on a small north‐eastern Atlantic islet of Porto Santo (Madeira Archipelago, Portugal). The resulting structure is named as a new trace fossil Imbutichnus igen. nov., characterized by a small, funnel‐like cavity, a circular to oval cross‐section, with a nearly cylindrical shape in the upper part and a conical shape towards the base. Imbutichnus costatum isp. nov. is defined as a bioclaustration structure produced by the overgrowth of a pyrgomatid barnacle by a coral. From an ethologic point of view, Imbutichnus is attributed to the Impedichnia category. In terms of palaeoecology, it is interpreted as the result of a parasitic relationship. This is also the oldest record of pyrgomatid barnacles in the eastern Atlantic and clearly demonstrates that the Miocene palaeogeographic range of Ceratoconcha was much wider than previously assumed.     

Influence of environmental variation on the pollination of the ambophilous sedge <Emphasis Type="Italic">Rhynchospora ciliata</Emphasis> (Cyperaceae)

In typically anemophilous families, there are species, such as Rhynchospora ciliata (Cyperaceae), in which both wind and insects have been registered as pollen vectors (ambophily). However, the contribution of each vector to pollination of ambophilous species can change under different environmental conditions. Therefore, this study aimed to determine the primary pollination system, abiotic or biotic, of R. ciliata, comparing wet and dry seasons. We tested the following hypotheses: (1) in dry months, wind is the main pollen vector of R. ciliata; (2) in wet months, insects will contribute more to the reproductive success of R. ciliata and compensate for low fruit formation by the abiotic vector. Our hypothesis was not confirmed. We found that higher wind speed can favor pollination by wind in R. ciliata even in wetter months. Because insects were less susceptible to environmental variation, they contributed more in dry months when wind speed was low, thus compensating for lower fruit set by wind. High rainfall and humidity can result in faster settling of pollen grains suspended in the air; however, rainfall does not prevent pollination by wind. Since individuals of R. ciliata are located near the edge of the forest, grouped quite close together, and as the inflorescence is located at the apex of the floral scape, wind can act as pollen vector and the inhibiting effects of rainfall can be minimized. The contribution of each pollen vector is important to the reproductive success of R. ciliata and the effectiveness of wind varies with local environmental conditions.     

Evaluation of a fully automated method to measure the critical removal stress of adult barnacles

A computer-controlled force gauge designed to measure the adhesive strength of barnacles on test substrata is described. The instrument was evaluated with adult barnacles grown in situ on Silastic T2®-coated microscope slides and epoxy replicas adhered to the same substratum with synthetic adhesive. The force per unit area required to detach the barnacles (critical removal stress) using the new automated system was comparable to that obtained with ASTM D5618 (1994 ASTM D5618. 1994. Standard test method for measurement of barnacle adhesion strength in shear American Standard for Testing and Materials. Paint-Tests for formulated products and applied coatings 06.01 [Google Scholar]) (0.19 and 0.28 MPa compared with 0.18 and 0.27 MPa for two batches of barnacles). The automated method showed a faster rate of force development compared with the manual spring force gauge used for ASTM D5618 (1994 ASTM D5618. 1994. Standard test method for measurement of barnacle adhesion strength in shear American Standard for Testing and Materials. Paint-Tests for formulated products and applied coatings 06.01 [Google Scholar]). The new instrument was as accurate and precise at determining surface area as manual delineation used with ASTM D5618 (1994 ASTM D5618. 1994. Standard test method for measurement of barnacle adhesion strength in shear American Standard for Testing and Materials. Paint-Tests for formulated products and applied coatings 06.01 [Google Scholar]). The method provided significant advantages such as higher throughput speed, the ability to test smaller barnacles (which took less time to grow) and to control the force application angle and speed. The variability in measurements was lower than previously reported, suggesting an improved ability to compare the results obtained by different researchers.