Different antipredator behaviour in two anuran tadpoles: effects of predator diet

Recent investigations have indicated that animals are able to use chemical cues of predators to assess the magnitude of predation risk. One possible source of such cues is predator diet. Chemical cues may also be important in the development of antipredator behaviour, especially in animals that possess chemical alarm substances. Tadpoles of the common toad (Bufo bufo) are unpalatable to most vertebrate predators and have an alarm substance. Tadpoles of the common frog (Rana temporaria) lack both these characters. We experimentally studied how predator diet, previous experience of predators and body size affect antipredator behaviour in these two tadpole species. Late-instar larvae of the dragonfly Aeshna juncea were used as predators. The dragonfly larvae were fed a diet exclusively of insects, R. temporaria tadpoles or B. bufo tadpoles. R. temporaria tadpoles modified their behaviour according to the perceived predation risk. Depending on predator diet, the tadpoles responded with weak antipredatory behaviour (triggered by insect-fed predators) or strong behaviour (triggered by tadpole-fed predators) with distinct spatial avoidance and lowered activity level. The behaviour of B. bufo in predator diet treatments was indistinguishable from that in the control treatment. This lack of antipredator behaviour is probably related to the effective post-encounter defenses and more intense competitive regime experienced by B. bufo. The behaviour of both tadpole species was dependent on body size, but this was not related to predator treatments. Our results also indicate that antipredator behaviour is largely innate in tadpoles of both species and is not modified by a brief exposure to predators. Received: 22 August 1996 / Accepted after revision: 31 January 1997     

The effect of mirrors on African clawed frog (Xenopus laevis) larval growth, development, and behavior

We examined the behavioral and developmental responses of Xenopus laevis larvae to their mirror images in three experiments. The mirrors allowed us to visually simulate increased density, without the tadpoles’ behavior being confounded by chemical cues from additional tadpoles. In the first experiment, we demonstrated that Xenopus tadpoles have a right eye preference for mirrors, contrary to the left eye preference of all other anuran species studied to date. This lateralized eye use disappeared, however, as tadpoles approached metamorphosis. Next, we examined how mirrored aquaria walls affected tadpole growth and development. We found that tadpoles raised in aquaria with partially mirrored walls showed depressed growth compared to tadpoles raised without mirrors, despite the fact that Xenopus larvae normally thrive when raised in visual contact with conspecifics. The tadpoles raised with mirrors had, though not significantly, proportionally larger bodies relative to their tail length (d = 0.51). This suggests that a phenotypically plastic response in body proportions was induced in these tadpoles solely by the sight of other tadpoles. The third experiment established that X. laevis tadpoles are more active in front of a mirror; i.e., they turn more often and spend more time in front of mirrored surfaces. We consider this increased activity to be an aberrant behavior of the tadpoles, which were attempting to school with their own images. We suggest that this extra activity reduced the amount of energy available for growth, accounting for the depressed growth seen in our second experiment.     

Diet and prey selectivity of scyphomedusae from Port Phillip Bay,Australia

Scyphomedusae collected from Port Phillip Bay, Victoria, Australia, between 1984 and 1986, consumed a variety of zooplankton. The percentage composition of gut contents of Cyanea capillata (Linné) in order of decreasing importance was larvaceans 31%, cladocerans 29%, fish eggs 14%, copepods 11%, hydromedusae 9%, and ascidian tadpoles 3%. The percentage composition of gut contents of Pseudorhiza haeckeli Haacke was fish eggs 41%, copepods 33%, larvaceans 8%, cladocerans 4%, crab zoea 4%, and decapod larvae 1%. Both species of scyphomedusae showed strong positive selection for fish eggs and yolk-sac larvae, and negative selection for other prey items. When fish eggs were omitted from the selectivity analyses, C. capillata showed positive selection for amphipods, decapods, crab zoea, Podon spp., larvaceans and ascidian tadpoles, and negative selection for Evadne spp. and all copepod taxa. Pseudorhiza haeckeli showed positive selection for amphipods, decapod larvae, crab zoea and cladocerans, and negative selection for cirripede larvae, larvaceans and hydromedusae. Amongst copepods, P. haeckeli showed positive selection for calanoid and harpacticoid copepods and negative selection for cyclopoid copepods.     

Lack of sequestration of host plant glucosinolates in <Emphasis Type="Italic">Pieris rapae</Emphasis> and <Emphasis Type="Italic">P. grarricae</Emphasis>

Summary. Sequestration of plant toxins in herbivores is often correlated with aposematic coloration and gregarious behaviour. Larvae of Pieris brassicae show these conspicuous morphological and behavioural characteristics and were thus suggested to sequester glucosinolates that are characteristic secondary metabolites of their host plants. P. rapaeare camouflaged and solitary, and are thus not expected to sequester. To test this hypothesis and to check the repeatabi-lity of a study that did report the presence of the glucosinolate sinigrin in P. brassicae, larvae were reared on three species of Brassicaceae (Sinapis alba, Brassica nigra and Barbarea stricta), and different leaf and insect samples were taken for glucosinolate analysis. The major host plant glucosinolates could only be found in traces or not at all in larval haemolymph, bled or starved larvae, faeces or pupae of both species or P. brassicae regurgitant. Haemolymph of both Pieris spp. was not rejected by the ant Myrmica rubra in dual-choice assays; the regurgitant of P. brassicae was rejected. This suggests the presence of compounds other than glucosinolates that might be sequestered in or produced by P. brassicae only. In faeces of both Pieris spp. a compound which yielded 4-hydroxybenzylcyanide (HBC) upon incubation with sulfatase was detected in high concentrations when larvae had been reared on S. alba. This compound may be derived from hydrolysis of sinalbin, the main glucosinolate of that plant. The unidentified HBC progenitor was apparently not sequestered in the two Pieris spp., and was not detected in faeces of larvae reared on B. nigra or B. stricta. Received 18 July 2002; accepted 11 September 2002.     

全氟辛烷磺酸盐(PFOS)及替代品对两栖类蝌蚪的急性毒性

PFOS及其4种替代品对两栖动物非洲爪蟾和黑斑蛙蝌蚪的急性毒性结果为:用调聚法合成的织物三防整理剂对非洲爪蟾蝌蚪和黑斑蛙蝌蚪的96h-LC50分别为8和21mg·L-1,而PFOS对两种蝌蚪的96h-LC50分别为92和81mg·L-1。此实验结果说明织物三防整理剂的急性毒性高于PFOS。用电解氟化法合成的C4、C6织物三防整理剂和50%的全氟丁基有机铵盐阳离子表面活性剂浓度在100mg·L-1时,对两种蝌蚪都没有毒性。这说明,从急性毒性的角度,C4、C6织物三防整理剂和表面活性剂可作为PFOS的替代品使用,但是织物三防整理剂的急性毒性比PFOS大,作为替代品使用应慎重考虑。另外,PFOS和织物三防整理剂对黑斑蛙蝌蚪的急性毒性与对非洲爪蟾蝌蚪的急性毒性存在差异。出于保护我国本土两栖动物的目的,使用黑斑蛙开展毒性评价比使用非洲爪蟾更有现实意义。     

Aposematism in a soft-bodied insect: a case for kin selection

Summary This paper describes the influence on predator behaviour, and the survival of an aposematic aphid, Aphis nerii, in comparison with a palatable, cryptic aphid, Acyrthosiphon pisum, when offered to two predators with different foraging tactics. The experiments were designed to test Fisher's (1930) suggestion that aposematism could evolve by kin selection, since aposematic animals often occur in aggregations of relatives. Initially, spiders (Zygiella x-notata) and birds (Parus major) killed high proportions of distasteful A. nerii (60% and 54% respectively). With experience, the predators killed and ate fewer A. nerii. The decreasing mortality of A. nerii after initial encounters with predators, coupled with its apparently obligate parthenogenesis, indicate that the evolution of aposematism in this soft-bodied insect is consistent with kin selection.     

Benefit of suites of defensive behavior induced by predator chemical cues on anuran tadpoles, Hyla japonica

When predator chemical cues are present, low activity of prey is a commonly seen defensive behavior. However, few studies have explored the functional implications of the defensive behaviors and, thus, elucidated the possible linkages between behavioral responses and its consequences. In this study, we experimentally investigated how behavioral responses of Hyla japonica tadpoles to predator chemical cues affect vulnerability to a dragonfly nymph Anax parthenope julius. The frequency of tadpoles attacked by dragonfly nymphs was lower with chemical cues of predator was present than without chemical cues, and most of attacks occurred when tadpoles were mobile. When tadpoles were exposed to chemical cues, on the other hand, their swimming speed was quicker and swimming distance was longer, respectively, and the rates of being approached of the swimming tadpoles by dragonfly nymph was lower than those not exposed to chemical cues. We found that the tadpoles are induced by predator chemical cues not only to generally lower activity but also to swim in bursts as additional behavior and that the suite of their behavioral responses reduce the vulnerability against dragonfly nymph. Tadpoles can receive information about the predation risks by chemical cues and adjust their defensive behavior accordingly.     

Sequestration of phorbol esters by aposematic larvae of Hyles euphorbiae (Lepidoptera: Sphingidae)?

Summary. The larvae of the hawkmoth species Hyles euphorbiae have a conspicuous aposematic colouration and show gregarious behaviour. It has thus been suggested that they sequester phorbol esters from their food plants which include different species of the genus Euphorbia (Euphorbiaceae) for chemical protection against predators. To test this hypothesis in more detail, we fed larvae an artificial diet with three doses of 12-tetradecanoyl-phorbol-13-acetate (TPA), then examined the faeces and the larval tissues, such as integument, haemolymph and gut of the caterpillars for the presence of TPA. In order to determine the ability of the larvae to detoxify phorbol esters, other larvae were directly injected with a TPA solution and analysed in the same manner. Our study indicates that the larvae of Hyles euphorbiae do not sequester phorbol esters. Upon oral application TPA was not found in the larval integument or the haemolymph. Instead, it was mostly metabolised (about 70–90%). Nevertheless, about 10-30% were retained and recovered in the faeces. The larvae were also able to metabolise and thus detoxify the phorbol ester when TPA was injected directly into the body. These hawkmoth caterpillars are relatively large and have a gut full of plant material, which they regurgitate into the direction of the predator when attacked in nature. Since phorbol esters are very potent toxins and irritants, we postulate that the gut content (and especially the plant slurry disgorged as regurgitant from the anterior gut) alone could be aversive for a potential predator, even if some metabolism has taken place. Thus, although H. euphorbiae caterpillars do not actively sequester phorbol esters, their aposematic colouration appears to be based on chemical defence through phorbol esters retained in the gut.     

Intracellular bacteria associated with the ascidian<Emphasis Type="Italic"> Ecteinascidia turbinata</Emphasis>: phylogenetic and in situ hybridisation analysis

The ascidian Ecteinascidia turbinata (Herdman) is a colonial sea squirt found in the Caribbean and Mediterranean Seas. In the present study, the bacterial complement of E. turbinata has been assessed by 16S rRNA gene analysis and the most commonly occurring strains identified by restriction fragment length polymorphism. Three strains were found to predominate using this approach, with one representing >50% of clones from both larval and adult material. The 16S rRNA gene sequence of the most commonly occurring strain did not match with any known bacterial sequences and could only be assigned to the γ-proteobacteria subdivision. The two other frequently occurring strains were assigned to the Mollicutes. In situ hybridisation analysis with eubacterial probes to 16S rRNA revealed the presence of apparently endosymbiotic bacteria in adult and larval tissue, and electron microscopy confirmed the presence of putative bacteriocytes in the larval tissue. The presence of the same bacteria in the brooded larvae suggested that they were vertically transmitted from parent to offspring. Further hybridisation using a novel probe designed to be specific to the 16S rRNA sequence of the dominant strain, highlighted the same cell types as that revealed by the eubacterial probe. The results suggest that the bacteria represent a novel strain, denoted "Candidatus Endoecteinascidia frumentensis", and that they may have an important role in the biology of E. turbinata.     

Chemically mediated avoidance of a predatory odonate (Anaxjunius) by American toad (Bufoamericanus) and wood frog (Ranasylvatica) tadpoles

Predators can have significant nonlethal effects on prey by modifying prey behaviour through chemically mediated interactions. We examined behavioural responses of wood frog (Ranasylvatica) and American toad (Bufoamericanus) tadpoles to both direct and indirect chemical signals associated with a predatory odonate (Anaxjunius). In laboratory trials, tadpoles of both species responded strongly to water conditioned with Anax nymphs by decreasing foraging rates, becoming immobile, and moving away from the stimulus. The responses to water conditioned with starved Anax versus Anax that fed on conspecific tadpoles did not differ significantly; these results suggest that tadpoles rely primarily on direct signals to detect odonates. Rana did not respond to water conditioned with conspecific tissue extracts, while Bufo responded with behaviours that were indistinguishable from those of tadpoles exposed to Anax chemicals. In a field experiment, the responses of R. sylvatica tadpoles to Anax chemicals were similar to those of tadpoles observed in the laboratory. Collectively, our data indicate that tadpoles of both species use chemical cues to assess predation risk from other community members. Tadpoles can selectively distinguish members who pose a threat, and only evacuate food patches or reduce foraging rates when in danger. These behaviours appear to be adaptive and are consistent with the predictions of optimality theory. Received: 1 August 1997 / Accepted after revision: 22 November 1997     

Strange parental decisions: fathers of the dyeing poison frog deposit their tadpoles in pools occupied by large cannibals

Parents may increase the probability of offspring survival by choosing suitable rearing sites where risks are as low as possible. Predation and competition are major selective pressures influencing the evolution of rearing site selection. Poison frogs look after their clutches and deposit the newly hatched tadpoles in bodies of water where they remain until metamorphosis. In some species, cannibalism occurs, so parents deposit their tadpoles singly in very small pools. However, cannibalism also occurs in species that deposit tadpoles in larger pools already occupied by heterospecific or conspecific larvae that could be either potential predators or competitors. Here, I test the hypothesis that, given the choice, males of Dendrobates tinctorius would deposit their newly hatched tadpoles in low-risk sites for their offspring. I characterised the pools used by D. tinctorius for tadpole deposition, conducted experiments to determine the larval traits that predict the occurrence of and latency to cannibalism, and tested whether parents deposit their tadpoles in low-risk pools. I found that (1) neither pool capacity nor the presence of other larvae predict the presence/absence or number of tadpoles; (2) cannibalism occurs often, and how quickly it occurs depends on the difference in size between the tadpoles involved; and (3) the likelihood of males depositing their tadpoles in occupied pools increases with the size of the resident tadpole. I suggest that predation/cannibalism is not the only factor that parents assess when choosing deposition sites, and that the presence of larger conspecifics may instead provide information about pool quality and stability.     

Vertical distribution of larvae of Euphausia nana and E. similis (Crustacea: Euphausiacea) in Sagami Bay and Suruga Bay,Central Japan

Vertical distributions of the larval stages of Euphausia nana Brinton and E. similis G. O. Sars in Sagami Bay and Suruga Bay, Central Japan were studied. Most of the metanauplius larvae of E. nana occurred between 25 and 80 m depth, and they were found at greater depths than the eggs and calyptopis larvae. The nauplii and metanauplii of e. similis were mainly found between 50 and 100 m depth, and they also occurred deeper than the eggs and calyptopes. The larvae of the two species from calyptopis I demonstrate diurnal vertical migration. However, this phenomenon was not clear in the season (March) when a seasonal thermocline was absent. Calyptopes and fruciliae of E. similis occurred deeper and migrated over greater vertical distances than those of E. nana. The distance of migration of furcilia I larvae of E. similis was estimated to be about 200 m.     

Kin preference behavior in Bufo boreas tadpoles

Summary The ontogeny of sibling recognition behavior was studied in the laboratory in tadpoles of the western toad (Bufo boreas boreas) to test the hypothesis that Bufo tadpoles associate with siblings and to compare this behavior with two species of anurans previously studied. Tadpoles reared exclusively with sibs demonstrated a preference to associate with sibs over non-sibs both early and late in development but tadpoles reared with sibs and non-sibs (mixed rearing groups) exhibited no preference. Larvae that developed a preference for sibs after being reared with them for 75 days lost this preference following exposure to a mixed group for 2 to 6 days. Additionally, larvae reared in a mixed group did not develop a preference for the familiar mixed group nor was a preference exhibited by individual larvae that were reared exclusively with 5 non-sibs. These results suggest that larvae learn by prior association to discriminate sibs from non-sibs and that preferences are rapidly modifiable following exposure to non-sibs. However, the latter two experiments suggest that social preferences are not totally labile and are not based entirely on familiarity. Field observations of larval activity, aggregation behavior, and dispersal patterns in B. boreas indicate that tadpoles mix with non-sibs from early larval stages throughout development. Because even short term exposure to nonsibs resulted in a loss of sib preference in our experiments, we suggest that a kin selection interpretation of B. boreas aggregation behavior is not parsimonious. Results differ with those of other studies of sib recognition in larval anurans.     

Bright plumage in the magpie: does it increase or reduce the risk of predation?

Conspicuous color patterns in birds may lead to increased risk of predation. Alternatively, bright birds may be aposematic or unprofitable prey, which leads to decreased predation. During four autumns, I examined whether the black-and-white plumage of (stuffed) magpies Pica pica increases or reduces the risk of attack by migrating goshawks Accipiter gentilis. Attack risk was higher for normal magpies than for cryptic, brown magpies whether the mounts were placed near one another or far apart. However, the brown magpie may have been avoided because of its novelty. For magpies and cryptic jays Garrulus glandarius exposed together, the attack risk was similar. In 2 years, magpie and jay mounts were also exposed far apart. In 1994, with an invasion of migrating jays, attack risk was much higher for jays than for magpies. In 1995, with a normal density of jays, hawks attacked the magpies more often. The results demonstrate frequency-dependent prey selection by goshawks, which would influence any predation cost of bright plumage. The attacks on normally colored magpie mounts suggest that magpies are not aposematic. Trials with photographs and human observers indicated that normal magpies were somewhat easier to detect than jays. The plumage of the magpie possibly increases the risk of predation, but may be favoured by sexual or social selection. Received: 18 January 1996 / Accepted after revision: 6 October 1996     

Evidence for a water-borne cue inducing metamorphosis in the dorid nudibranch mollusc Adalaria proxima (Gastropoda: Nudibranchia)

The dorid nudibranch Adalaria proxima (Alder & Hancock) is a specialist predator of the cheilostome bryozoan Electra pilosa (L.). Natural induction of metamorphosis of the pelagic lecithotrophic larva of A. proxima was assessed in response to solutions from sonicated prey tissue and (live) E. pilosa-conditioned seawater (Electra-CSW). We exploited the tendency of larvae to become entrapped (rafted) at the air-water interface in cultures to examine whether larvae require direct contact with the live prey for metamorphosis to proceed. Larvae metamorphosed when rafted above colonies of live E. pilosa, above plankton mesh bags isolating live E. pilosa, and in choline chloride controls; there was no metamorphosis of larvae that were rafted in filtered seawater controls. Entrapped veliger shells remained rafted throughout the experimental period in all cases. No metamorphosis occurred in treatments containing either the supernatants or pelleted particulates obtained from sonicated colonies of E. pilosa. Both one-colony and three-colony Electra-CSW induced metamorphosis of larvae. These data are at variance with previous results in showing that direct contact with the live prey is not necessary for metamorphosis to proceed. Furthermore, the fact that competent larvae metamorphosed in response to Electra-CSW in the absence of any other cue strongly suggests that the inductive cue is water-borne.     

Larval distribution and reproductive ecology of Thysanoessa macrura (Crustacea: Euphausiacea) in the Scotia Sea

In the Scotia Sea, the larvae of Thysanoessa macrura G.O. Sars, 1885 were scarce in spring (September 27 to October 6, 1971) and mainly present at the Calyptopis I stage, while in summer (December 17, 1974 to January 3, 1975) they were much more abundant and much further developed (up to Furcilia VI). The spawning season commences as early as September and, as indicated by the presence of numerous nauplii and metanauplii in summer, is rather long. T. macrura begins to spawn much earlier in West-Wind-Drift waters than in waters of the Weddell Sea. It starts to spawn simultaneously with Euphausia frigida and E. triacantha, but significantly earlier than E. superba. These timings of the breeding process are related to differential biogeographical characteristics of these species. Plankton collections made at depths of 500 or 1000 m have, for T. macrura, revealed a developmental ascent analogous to that of E. superba. Ontogenetical deep-water migrations are, consequently, characteristic not only for E. superba, but also for other Antarctic euphausiids (in particular E. frigida, but possibly not for inshore E. crystallorophias) and, as correctly postulated by several other investigators, for all oceanic euphausiid species.     

Effects of predation risk and hunger on the behaviour of two species of tadpoles

Predation and hunger are threats for most organisms, and appropriate behavioural responses to both factors should be shaped by natural selection. In combination, however, the behavioural demands of predation avoidance and effective foraging often cannot be satisfied at the same time and lead to a conflict within organisms. We examined the behavioural responses of two closely-related species of tadpoles, Rana lessonae and R. esculenta, to simulated predation by fish and hunger. Tadpoles, hatched and reared in the laboratory, were tested in a three-way factorial (predation risk × hunger × species) experiment with four predation levels and four hunger levels. Both species decreased their swimming activity with increasing predation risk. Predation risk did not influence the amount of activity time invested in feeding but caused the tadpoles to spend less time in patches with food. Refuges were not used to avoid predation. R. esculenta was more sensitive to predation risk than R. lessonae. Hunger increased both the activity of tadpoles and the amount of activity time invested in feeding, thus indicating an increased energy intake. No interactions were observed between predation risk and hunger. These results show that tadpoles possess genetically-based behavioural mechanisms that allow them to respond in a graded manner to predation and hunger. However, they did not balance the two conflicting demands of predation avoidance and effective foraging; the two mechanisms appeared to act independently. Correspondence to: R.D. Semlitsch     

Feeding,growth, and survival of Engraulis mordax larvae reared in the laboratory

Methods are described for the successful rearing of northern anchovy larvae (Engraulis mordax Girard) on cultured foods. Larvae were fed successively on the unarmored dinoflagellate Gymnodinium splendens, the veliger of the gastropod Bulla gouldiana, and nauplii of the brine shrimp Artemia salina. Rearing containers ranging in capacity from 4.5 to 510 l were tested; the smaller ones were found to be most useful for laboratory experimentation. Irreversible starvation occurred when E. mordax were denied food for more than 1.5 days after yolk absorption. Growth rates of larval anchovies fed different diets were compared. Larvae fed G. splendens grew for 1 week at the same rate as animals fed wild plankton, but did not maintain this rate. Laboratory survival of E. mordax larvae on a diet of G. splendens alone, did not differ significantly when veligers supplemented the diet. However, when G. splendens and veligers were fed simultaneously to E. mordax larvae, growth rate was greatly improved, although still not matching the growth attained on a diet of wild plankton. Length (L) versus weight (W) analyses were made for all larvae at all diets. The results showed that weight could be calculated most accurately from length by the relationship log W=3.3237 log L-3.8205, regardless of diet.     

Induction of micronuclei in tadpoles of Odontophrynus americanus (Amphibia: Leptodactylidae) by the pyrethroid insecticide cypermethrin

Cypermethrin (CY) is an active cyano pyrethroid effective against a wide range of pests encountered in agriculture and forestry. Although CY is not mutagenic in in vitro assays for gene mutation, in vivo assays showed conflicting results. In vivo genotoxicity of the synthetic pyrethroid CY in erythrocytes of Odontophrynus americanus tadpoles was examined. The frequency of micronuclei (MN) was recorded in blood smears obtained from tadpoles exposed in vivo to four different nominal concentrations 5, 10, 20 or 40?µg?L^?1 of the compound and fixed at two sampling times 48 and 96?h. As a positive control larvae were exposed to 40?mg?L^?1 of cyclophosphamide (CP). Tadpoles exposed to all CY treatments showed a significant increase in single small MN compared to the negative control group after 48?h and at 5 and 10?µg?L^?1 of CY at 96?h. Results obtained here demonstrated the genotoxic effects of the commercial formulation CY in the anuran larvae analyzed. Thus, data suggest that measurements of MN and other erythrocytes morphological aberrations performed in circulating blood samples of O. americanus tadpoles is a method for detecting cytogenetic damage in other native species.     

Inhibition and facilitation of bryozoan and ascidian settlement by natural multi-species biofilms: effects of film age and the roles of active and passive larval attachment

Sessile marine invertebrate larvae can recognize suitable settlement substrata by using various environmental cues, including organic/microbial biofilms. In laboratory choice assays, the effect of biofilms of varying ages on the settlement behaviour of two fouling organisms was assessed. The species included the arborescent cheilostome bryozoan Bugula flabellata (Thompson) and the solitary ascidian Ciona intestinalis (L.), both of which are characteristic of temperate sublittoral hard substratum assemblages in northwest Europe. Experiments were carried out using polystyrene petri dish substrata preconditioned with multispecies biofilms from natural laboratory-aquarium seawater for 1, 3, 6 or 12 d. Unfilmed (new, initially sterile) dishes were used as control substrata. Whereas the coronate larvae of B. flabellata generally were inhibited by biofilming, irrespective of film age, the settlement of tadpole larvae of C. intestinalis was facilitated on biofilmed substrata, and numbers of settled larvae generally increased with biofilm age: the highest mean numbers were counted on 12 d-old biofilms. In C. intestinalis, settlement and metamorphosis are processes which can be temporally separate and are possibly induced by different environmental cues. This study therefore distinguished between C. intestinalis larvae which were attached to the biofilm surface by the anterior, and those larvae entrapped by the biofilm but not settled in the conventional meaning of the term. As reported in previous studies, we did observe that such entrapped larvae could subsequently attach and develop successfully into sessile juveniles. Both the numbers of “attached” and “trapped” tadpoles increased with biofilm age. Assuming that “settlement” is essentially a process involving the active behavioural response of larvae to environmental cues, it seems that the facilitated attachment of C. intestinalis onto biofilmed substrata is due to the combined effect of active habitat selection and passive deposition/“entrapment” of larvae onto the “sticky” substratum. Received: 21 August 1996 / Accepted: 21 November 1996