First record of an epibiosis between the sand crab Emerita analoga (Stimpson, 1857) (Decapoda: Hippidae) and the mussel Semimytilus algosus (Gould, 1850) (Bivalvia, Mytilidae) in southern Peru

Coastal zones of the Humboldt Current Upwelling System (HCUS) are composed both of rocky and sandy beaches inhabited by macrozoobenthic communities. These show oscillating changes in the dominance of species; the abundance of the sand crab Emerita analoga is linked to phases of the El Niño Southern Oscillation (ENSO). The biogenic surfaces of these crabs serve as substrate for opportunistic colonizers. This study is the first record of an epibiosis between E. analoga and the rock mussel Semimytilus algosus, detected at a southern Peruvian sandy beach. Mussels fouled a wide size-range of adult E. analoga (7.3%) but they themselves belonged to small-size classes. The largest S. algosus was 17.4 mm in length. Highest permanence of epibionts was found on larger sand crabs (maximum between 24 and 27 mm). Significantly more mussels were found on the ventral surface (39.4%) compared to 10 other surface areas of the sand crab. Possible benefits and disadvantages of the observed epibiosis for both the basibiont and the epibiont are discussed.     

Hydrozoan and protozoan epibionts on two decapod species, Liocarcinus depurator (Linnaeus, 1758) and Pilumnus hirtellus (Linnaeus, 1761), from Scotland

Crabs of the species Liocarcinus depurator and Pilumnus hirtellus, collected on the west coast of Scotland, showed hydrozoan and protozoan epibionts. The epibionts of Liocarcinus depurator were the protozoan Ephelota plana and the hydrozoans Clytia gracilis and Leuckartiara sp. The epibionts of Pilumnus hirtellus were the protozoans E. plana and Zoothamnium sp. For Liocarcinus depurator the number of Ephelota per crab fluctuated between 0 and 47 and the greatest number of suctorians were located on the chelipeds, carapace and anterior pereiopods. The hydrozoans, for Liocarcinus depurator, showed densities of 0-20 (Clytia gracilis on the second pereiopod) and 0-507 individuals per crab (Leuckartiara sp., principally on chelipeds, carapace and the fourth right pereiopod). For Pilumnus hirtellus, Ephelota plana showed densities between 3 and 56 individuals per crab, the greatest number of suctorians being located on the same areas as on Liocarcinus depurator. There was a density of 10-69 individuals per crab of Zoothamnium sp. on Pilumnus hirtellus (located on the carapace). Ephelota plana has not been observed previously as an epibiont on crustacea, nor had Zoothamnium sp. as an epibiont on Pilumnus hirtellus. Both hydrozoans, Leukartiara and Clytia, have not been previously described as epibionts on Liocarcinus depurator. Data about the morphological characteristics and distribution of these epibionts are included.     

Evidence of dietary feedback in a facultative association between deep-sea gastropods and sea anemones

While epibiotic associations between macrobenthic invertebrates are common, the role they play in the feeding ecology of intervening species is often incompletely understood. The diets of epibiotic sea anemones Allantactis parasitica and their gastropod hosts were analyzed using digestive tract contents, lipid biomarkers and observations of live specimens in an attempt to detect dietary feedback from the facultative association. Comparisons were made using symbiotic individuals and asymbiotic counterparts collected at depths of 191-627 m from three neighbouring areas in the northwest Atlantic. Gastropods carrying one or two epibionts had higher stomach indices than those harbouring three epibionts or no epibiont. The diet of symbiotic gastropods was also more diversified based on stomach contents and lipid analysis. Among other things, symbiotic gastropods contained four times more lipids and a greater proportion of Σn−3 fatty acids. Gastrovascular cavity content indices of asymbiotic sea anemones were generally lower than those of symbiotic counterparts. Their cavities were more often empty, and their diet less diversified with fewer benthic items, suggesting that foraging of gastropods through the sediments makes more food available to sea anemones living as epibionts. Lipid analysis showed some disparities between symbiotic and asymbiotic sea anemones at the regional scale, including in percent phospholipids and in the proportion of certain fatty acids. Together these findings indicate that mutual protection against predators leads to prolonged and more efficient foraging for gastropods and increased time spent deployed (feeding) in food-rich areas for sea anemones, thus enabling both partners to fully exploit food resources that may be limited at bathyal depths.     

The shell of <Emphasis Type="Italic">Hexaplex (Trunculariopsis) trunculus</Emphasis> (Gastropoda: Muricidae) as a mobile hard substratum for epibiotic polychaetes (Annelida: Polychaeta) in the Ria Formosa (Algarve coast—southern Portugal)

This work reports and illustrates secondary colonisation of Hexaplex (Trunculariopsis) trunculus shells by polychaetes in the Ria Formosa lagoon (Algarve coast—southern Portugal). Information is presented in terms of qualitative and quantitative data on epibiotic polychaetes, fouling frequency, degree of epibiosis, preferential areas for colonisation and incidence of shell damages. A total of 2,880 polychaetes belonging to 10 families were identified. Besides Spirorbidae (not quantified), the most abundant families were Serpulidae (67.5%), Sabellidae (10.2%), Sabellariidae (9.6%) and Cirratulidae (8.5%). From a total of 2,264 T. trunculus individuals analysed for epibiosis, 28.4% were non-fouled, 66.8% were intermediately fouled and 4.8% were heavily fouled. The fouling frequency, degree of epibiosis and incidence of damages in the shell spire increased markedly with T. trunculus shell length. Some epibiotic polychaetes were located preferentially on/or nearby particular features of T. trunculus shells, which probably facilitate settlement and tube construction, in addition to provide some protection against tube damage. The main consequences of epibiosis (benefits and disadvantages) for the gastropod basibiont (T. trunculus) are discussed. The present study apparently indicates that besides burrowing into soft bottom substrates, sporadic inter-tidal exposure and mutual predation on epibiosis, T. trunculus lacks other typical antifouling defence adaptations. Handling editor: K. Martens     

Invasive zebra mussel colonisation of invasive crayfish: a case study

We investigated the interaction between two invasive invertebrate species in a shallow Central European flooded sandpit: the epibiosis of Ponto-Caspian zebra mussels Dreissena polymorpha on the American crayfish Orconectes limosus. Between 2004 and 2005, we followed the seasonal variation in number and size of the mussels attached to crayfish bodies, and microhabitats preferred by mussels. The proportion of crayfish colonised by mussels varied seasonally: in spring and early summer it was consistently over 75%, afterwards it dropped temporarily due to loss of bivalves during the crayfish moult, and later increased again due to re-colonisation by often relatively large juvenile mussels. Three different pathways of mussel settlement on crayfish hosts are likely: (1) primary settlement of free-swimming pediveliger larvae; (2) secondary settlement of plantigrade mussels and juveniles; (3) active re-attachment of grown mussels from the substrate to crayfish. This epibiosis was promoted by lack of suitable substrates at the studied locality. Electronic supplementary material The online version of this article (doi: ) contains supplementary material, which is available to authorized users.     

Taxonomy and distribution of the hydrozoan and protozoan epibionts on Pagurus bernhardus (Linnaeus, 1758) (Crustacea, Decapoda) from Scotland

Protozoan and hydrozoan epibionts on the hermit crab Pagurus bernhardus and its shell, collected near Cumbrae Island (Scotland) were studied. The epibionts found were the following: (1) protozoans: the suctorian ciliates Ephelota plana, Acineta compressa, Conchacineta constricta, Corynophrya anisostyla; the peritrich ciliates Cothurnia mobiusi and Zoothamnium plumula; the chonotrich ciliate Chilodochona quennerstedti; and (2) hydrozoans: the species Leuckartiara sp. and Clytia sp. The morphological characteristics of the epibionts were analysed, as well as their taxonomic position. The distribution of epibionts on the crab surface and its shell was studied, and the density and biomass of epibionts were calculated on each anatomic unit. There was a differential distribution according to the type of epibiont: hydrozoans dominated in biovolume and were present mainly on the shell, meanwhile protozoans represented the highest fraction of density and they were found exclusively on the crab (principally on eyes, antennulae, antennae, maxillipeds, pereopods and uropods). The anterior area of the cephalothorax was the most colonized. On this area, the maxillipeds and second pereopods showed the highest densities. The location of each epibiont species was described. There was a correlation between the length of the crab and the total number of hydrozoans. There was a significant correlation between the right and the left units of the crab, taking into consideration the mean densities of epibionts on each anatomical unit. The shell was colonized by the same species of hydrozoa that appeared on the crab, although in a much higher density (mean 3024.38 per shell; 6.9 per crab). There was a significant difference between both species of hydrozoan epibionts with respect to the mean densities on the different areas of the shell. The zone of the shell more occupied by Clytia sp. was the apical zone of the shell, while the highest densities of Leuckartiara sp. were registered near the aperture of the shell. The hydrozoan and protozoan epibiont species found on P. bernhardus in this study represent the first mention of their presence on this hermit crab.     

Associational resistance of fouled blue mussels (<Emphasis Type="Italic">Mytilus edulis</Emphasis>) against starfish (<Emphasis Type="Italic">Asterias rubens</Emphasis>) predation: relative importance of structural and chemical properties of the epibionts

Several epibiotic species reduce starfish (Asterias rubens) preference for the blue mussel Mytilus edulis in the Baltic. The aim of this study was to reveal whether this associational resistance was caused by structural or chemical aspects of the different epibionts. To assess structural epibiont effects, an in situ experiment was conducted with unfouled mussels and mussels equipped with artificial epibionts (dummies) exposed to natural predation by A. rubens. The chemically inert dummies closely matched the structural properties of the locally common epibionts Balanus improvisus (barnacle), Ceramium strictum (red alga), Halichondria panicea (sponge), and Laomedea flexuosa (hydrozoan). Starfish fed indiscriminately in all treatments. Chemical effects of epibionts on the attractiveness of mussels for A. rubens were investigated by incorporating freeze-dried epibionts or mussel tissue into Phytagel pellets at natural concentrations. Starfish were allowed to choose among these structurally similar but chemically different prey items in an in vitro experiment. The predators exhibited significant preferences among the food pellets, which closely matched their preferences for corresponding natural mussel–epibiont associations. Thus, chemical aspects of epibionts appear to play a larger role in this associational resistance than do structural aspects. Implications of these indirect interactions for benthic communities are discussed.Communicated by H.-D. Franke     

Effects of barnacle epibionts on the periwinkle Littorina littorea (L.)

In a sandy bay with mussel beds in the Wadden Sea (Island of Sylt, eastern North Sea), periwinkles Littorina littorea (L.) were often strongly overgrown with the barnacle Balanus crenatus Bruguière in the lower intertidal zone. Consequences of this epibiosis on mobility, reproduction and mortalityof the snail were examined. B. crenatus growing on L. littorea increased snail volume up to 4-fold and weight up to 3.5-fold and crawling speed of fouled L. littorea was significantly slowed down. The epibiotic structure also caused a decrease in reproductive output. In laboratory experiments, egg production of fouled L. littorea was significantly lower than in snails free of barnacles. Presumably, copulation of the periwinkles is hampered by the voluminous and prickly cover of barnacles. Field studies demonstrated an increased mortality of overgrown L. littorea. A decrease in reproductive output and a lower survival of snails with a cover of barnacles suggest that B. crenatus epibionts may have a significant impact on the population of L. littorea. Received: 20 June 1998 / Accepted: 2 October 1998     

Epibiotic demosponges on the Antarctic scallop <Emphasis Type="Italic">Adamussium colbecki</Emphasis> (Smith, 1902) and the cidaroid urchins <Emphasis Type="Italic">Ctenocidaris perrieri</Emphasis> Koehler, 1912 in the nearshore habitats of the Victoria Land,Ross Sea,Antarctica

The importance of epibiosis in Antarctic benthic communities is highlighted here considering the specific diversity of sponges living on shells of the scallop Adamussium colbecki and on spines of the cidaroid urchin Ctenocidaris perrieri. Scallops are from three different areas along the Victoria Land [Tethys Bay (TB), New Harbour (NH), Dunlop Island (DI)], while cidaroid urchins are from NH but not present in the two other stations. Homaxinella balfourensis is the commonest species both on the scallops and cidaroid urchins. Other common species on scallops are Myxilla (Myxilla) asigmata, Lissodendoryx (Ectyodoryx) nobilis and Iophon unicorne at NH, Iophon unicorne at DI, and Iophon radiatum, Haliclona sp. 1, Iophon unicorne and Lissodendoryx (Ectyodoryx) nobilis at TB. The highest number of sponge species we found on a single scallop was ten and the sample was collected at NH. On the spines of C. perrieri, Isodictya erinacea, Iophon unicorne and Haliclona (Rhizoniera) dancoi are present too. A. colbecki and C. perrieri, generally living on soft bottoms, represent important substrata for several sponge species. In this way, sponges may increase their dispersal exploiting valves and spines as stepping stones.     

Marine epibiosis

Summary Polysyncraton lacazei is a colonial tunicate (family didemnidae) living in the NW-mediterranean rocky sublitoral. A thorough scanning of numerous colonies revealed that in spite of an apparently heavy local fouling pressure only one fouling species — a kamptozoan — is encountered with some regularity on Polysyncraton. We try to define the epibiotic situation of sessile marine organisms as composed of four epibiotic parameters: longevity or exposure time (A), epibiont load (E), colonizer pool (CP) and fouling-period (FP). Subsequently, these factors are combined to propose an Antifouling Potential index: AFP=(1–E/CP)×A/(FP+A). This index is intended to permit evaluating the relative antifouling defense potency to be expected in a given organism in a given epibiotic situation and to compare different cases of epibiosis and fouling.