Surface sequestration of chemical feeding deterrents in the Antarctic sponge<Emphasis Type="Italic"> Latrunculia apicalis</Emphasis> as an optimal defense against sea star spongivory

Latrunculia apicalis is a spherically shaped demosponge that previous investigations have shown is rarely preyed upon by sea stars which are the dominant spongivores in antarctic benthic communities. Prior studies have also demonstrated that L. apicalis produces organic compounds that elicit a tube foot retraction response in the keystone spongivorous sea star Perknaster fuscus that can be used as a reliable assay for feeding deterrence. L. apicalis is known to contain discorhabdin alkaloids which serve, among other roles, as the source of its green coloration. To assess the defensive nature of the discorhabdin alkaloids toward P. fuscus, we have determined discorhabdin G concentrations in discrete sponge layers and evaluated those concentrations in the P. fuscus bioassay. In discorhabdin G-bearing sponges, we found a gradient of discorhabdin G that falls off rapidly toward the center of the sponge. On average, 52% of total discorhabdin G in a given sponge was found within 2 mm of the sponge surface. Tube foot retraction responses to extracts from the surface tissues (0–2 mm depth) of L. apicalis were compared to those of an inner layer (8–10 mm depth) and to a sample comprised of the same inner layer spiked with discorhabdin G at a concentration equivalent to that of the surface tissues. Tube foot retraction response times to extracts of the surface layers and the spiked inner layers were not statistically different, but were significantly greater than responses to the unaltered inner layer and controls. These results support the predictions of the optimal defense theory as L. apicalis sequesters its defensive chemistry (discorhabdin G) in its most vulnerable surface tissues, where the likelihood of predation from sea stars is highest. As antarctic sponges are generally preyed upon by extraoral feeding sea stars rather than deeper biting predators such as fish, surface sequestration may be uniquely adaptive in sessile macroinvertebrates occurring in antarctic marine benthic environments.Communicated by P.W. Sammarco, Chauvin     

Intraguild predation by <Emphasis Type="Italic">Harmonia axyridis</Emphasis> on coccinellids revealed by exogenous alkaloid sequestration

Summary.  Under laboratory conditions, the multicolored Asian lady beetle, Harmonia axyridis is well known as an intraguild predator of other ladybirds. However the real impact of this exotic species on native species was poorly investigated in the field. Because many ladybird species produce alkaloids as defensive compounds, we propose here a new method of intraguild predation monitoring in coccinellids based on alkaloid quantification by GC-MS. In laboratory experiments, adaline was unambiguously detected in fourth instar larvae of H. axyridis having ingested one egg or one first instar larva of Adalia bipunctata. Although prey alkaloids in the predator decreased with time, traces were still detected in pupae, exuviae and imagines of H. axyridis having ingested one prey when they were fourth instar larvae. Analysis of H. axyridis larvae collected in two potato fields shows for the first time in Europe the presence of exogenous alkaloids in 9 out of 28 individuals tested. This new method of intraguild predation detection could be used more widely to follow the interactions between predators and potential chemically defended insect preys.     

Induction of host specificity in larvae of Manduca sexta: chemical dependence controlling host recognition and developmental rate

Summary. The tobacco hornworm, Manduca sexta L. (Lepidoptera: Sphingidae) is a specialist on Solanaceae. This host specificity is induced as the larva feeds on solanaceous foliage, so that solanaceous-reared larvae will refuse to feed on other plants. Experiments were designed to determine the role of dietary constituents on the induction of host specificity and the effects of these on development of M. sexta. Choice assays using leaf discs of cowpea, Vigna sinensis (Fabaceae), were used to monitor the isolation of relevant chemical cues from foliage of potato. An aqueous extract of potato foliage strongly stimulated feeding. This extract was partitioned with n-butanol under alkaline conditions to obtain a highly active butanol extract. Reversed phase flash chromatography with a water-methanol gradient gave an active fraction that was used as a supplement for wheat germ-based artificial diet. Larvae reared on this s-diet became dependent on potato allelochemicals for initiation and continuation of feeding activity. These larvae also developed faster than larvae reared on the control p-diet, but no effect on adult mass was detected. Further flash chromatography of the active fraction under alkaline conditions provided a highly active sub-fraction, and semi-preparative HPLC using gradients of water and acetonitrile resulted in the isolation of a single compound. Bioassays indicated that this compound alone can account for host recognition by solanaceous-reared larvae. The results suggest that the mechanism of induced host specificity in M. sexta involves development of dependence on this compound. Received 21 December 1999; accepted 14 March 2000     

Localization of granulatimide alkaloids in the tissues of the ascidian <Emphasis Type="Italic">Didemnum</Emphasis><Emphasis Type="Italic">granulatum</Emphasis>

Ascidians are a notable source of nitrogen-bearing secondary metabolites with a wide range of biological activities. Although many biologically active compounds have been isolated from ascidians, it is often unclear whether the animal or associated microbial symbionts such as bacteria or fungi are the true biosynthetic source of the metabolites. We have addressed the question of the biosynthetic source of the alkaloids granulatimide and isogranulatimides by localizing these compounds within the ascidian. In this work, we demonstrate that granulatimide is stored in Didemnum granulatum tunic bladder cells. Analysis by confocal fluorescence microscopy at the granulatimide emission range indicated the presence of fluorescent cells as highly vacuolated cells found dispersed in the ascidian tunic. Since this is the most exposed ascidian tissue, it is possible that these alkaloids may have a protective role, either as sunscreens and/or as feeding deterrents.     

Larval settlement in the serpulid polychaete Hydroides elegans in response to bacterial films: an investigation of the nature of putative larval settlement cue

Larval settlement in the marine polychaete Hydroides elegans (Haswell) is induced by certain bacteria in marine biofilms. The exact nature of the settlement cue that larvae of H. elegans receive from these bacteria remains unknown. In this study, we revealed some properties of the bacterially derived larval settlement cue by investigating the larval settlement inductive activity of two bacterial strains after various treatments. These two bacterial strains, Roseobacter sp. and an α-subclass Proteobacteria, are highly inductive to larval settlement of H. elegans. The larvae responded similarly to Roseobacter and Proteobacteria in all the larval settlement bioassays, suggesting that the larval settlement-inducing substances produced by these bacteria may share common characteristics. First of all, the larvae did not settle in the seawater conditioned by the bacteria attached as a film or by the bacteria that were freely suspended in seawater. The results suggest that the putative larval settlement cue is not released into seawater and, therefore, should be associated with the surface of the bacteria. Secondly, formaldehyde treatment entirely eliminated the larval settlement induction activity of the bacterial films, and streptomycin treatment reduced the percentage of larval settlement on the bacterial films in a concentration-dependent manner. Since both treatments can kill bacteria with little damage to the surface chemistry of bacterial cells, the decline in larval settlement is suggested be due to a reduction of the viable bacterial population in the bacterial films. In fact, the reduction of larval settlement in the streptomycin treatments coincided with the decrease in viable bacterial populations in broth cultures containing respective concentrations of streptomycin. These results suggest that the viability of Roseobacter and Proteobacteria is important to their settlement induction effect. Since the larval settlement induction activity of the bacterial strains appears to correlate with their viability, we suggest that the putative larval settlement cue is derived from a metabolic pathway in the bacteria and that the cue is exported to and concentrated at the extracellular polymer matrix of the bacterial cell, at which the larvae establish contact with the bacteria. The larval settlement cue may be highly susceptible to degradation so that a metabolically active bacterial film is needed to maintain the putative cue at a concentration that surpasses the threshold for induction of larval settlement. Received: 14 October 1998 / Accepted: 5 September 2000     

Dissolved histamine: a potential habitat marker promoting settlement and metamorphosis in sea urchin larvae

Many species of marine invertebrate larvae settle and metamorphose in response to chemicals produced by organisms associated with the adult habitat, and histamine is a cue for larvae of the sea urchin Holopneustes purpurascens. This study investigated the effect of histamine on larval metamorphosis of six sea urchin species. Histamine induced metamorphosis in larvae of three lecithotrophic species (H. purpurascens, Holopneustes inflatus and Heliocidaris erythrogramma) and in one planktotrophic species (Centrostephanus rodgersii). Direct comparisons of metamorphic rates of lecithotrophic and planktotrophic larvae in assays cannot be made due to different proportions of larvae being competent. Histamine (10 μM) induced metamorphosis in 95% of larvae of H. purpurascens and H. inflatus after 1 h, while the coralline alga Amphiroa anceps induced metamorphosis in 40–50% of these larvae. Histamine (10 μM) and A. anceps induced 40 and 80% metamorphosis, respectively, in the larvae of H. erythrogramma after 24 h. Histamine (10 μM) and the coralline alga Corallina sp. induced 30 and 70% metamorphosis, respectively, in the larvae of C. rodgersii after 24 h. No metamorphosis of any larval species occurred in seawater controls. Larvae of two planktotrophic species (Tripneustes gratilla and Heliocidaris tuberculata) did not metamorphose in response to histamine. Seagrasses, the host plants of H. inflatus, induced rapid metamorphosis in larvae of the two Holopneustes species, and several algae induced metamorphosis in C. rodgersii larvae. Histamine leaching from algae and seagrasses may act as a habitat marker and metamorphic cue for larvae of several ecologically important sea urchin 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     

Settlement of larval blacklip abalone, Haliotis rubra, in response to green and red macroalgae

Surfaces from the habitat of adult Haliotis rubra were tested as inducers of larval settlement to determine the cues that larvae may respond to in the field. Settlement was high on the green algal species Ulva australis and Ulva compressa (Chlorophyta), the articulated coralline algae Amphiroa anceps and Corallina officinalis, and encrusting coralline algae (Rhodophyta). Biofilmed abiotic surfaces such as rocks, sand and shells did not induce settlement. Ulvella lens was also included as a control. Treatment of U. australis, A. anceps and C. officinalis with antibiotics to reduce bacterial films on the surface did not reduce the settlement response of H. rubra larvae. Similarly, treatment of these species and encrusting coralline algae with germanium dioxide to reduce diatom growth did not significantly reduce larval settlement. These results suggest that macroalgae, particularly green algal species, may play an important role in the recruitment of H. rubra larvae in the field and can be used to induce larval settlement in hatchery culture.     

The interplay between foraging mode, habitat structure, and predator presence in antlions

Antlion larvae are sand-dwelling insect predators, which ambush small arthropod prey while buried in the sand. In some species, the larvae construct conical pits and are considered as sit-and-wait predators which seldom relocate while in other species, they ambush prey without a pit but change their ambush site much more frequently (i.e., sit-and-pursue predators). The ability of antlion larvae to evade some of their predators which hunt them on the sand surface is strongly constrained by the degree of sand stabilization or by sand depth. We studied the effect of predator presence, predator type (active predatory beetle vs. sit-and-pursue wolf spider), and sand depth (shallow vs. deep sand) on the behavioral response of the pit building Myrmeleon hyalinus larvae and the sit-and-pursue Lopezus fedtschenkoi larvae. Predator presence had a negative effect on both antlion species activity. The sit-and-wait M. hyalinus larvae showed reduced pit-building activity, whereas the sit-and-pursue L. fedtschenkoi larvae decreased relocation activity. The proportion of relocating M. hyalinus was negatively affected by sand depth, whereas L. fedtschenkoi was negatively affected also by the predator type. Specifically, the proportion of individual L. fedtschenkoi that relocated in deeper sand was lower when facing the active predator rather than the sit-and-pursue predator. The proportion of M. hyalinus which constructed pits decreased in the presence of a predator, but this pattern was stronger when exposed to the active predator. We suggest that these differences between the two antlion species are strongly linked to their distinct foraging modes and to the foraging mode of their predators. Reut Loria and Inon Scharf contributed equally to the paper.     

Resistance of the generalist moth Trichoplusia ni (Noctuidae) to a novel chemical defense in the invasive plant Conium maculatum

Summary. Conium maculatum is an apiaceous species native to Eurasia that is highly toxic to vertebrates due to the presence of piperidine alkaloids, including coniine and γ-coniceine. More than 200 years after invading the United States this species remains mostly free from generalist insect herbivores. The presence of novel chemical defenses in the introduced range could provide invasive species with a competitive advantage relative to native plants. The cabbage looper (Trichoplusia ni) is a generalist lepidopteran found throughout the US that occasionally feeds on C. maculatum. We evaluated the toxicity of piperidine alkaloids to T. ni and determined putative resistance mechanisms, both behavioral and physiological, that allows this insect to develop successfully on C. maculatum foliage. T. ni larvae raised on diets enriched with coniine and γ-coniceine showed a decrease in consumption and longer development time, but no effects on growth were found at any alkaloid concentration. In a diet choice experiment T. ni larvae showed no avoidance of alkaloid-enriched diets, suggesting that the deterrence produced by alkaloids was related to a post-ingestive metabolic response. The ability of T. ni to consume diets high in alkaloid content could be due to at least three different mechanisms: 1) a decreased consumption rate, 2) efficient excretion of at least 1/3 of ingested alkaloids unmetabolized in frass, and 3) partial detoxification of alkaloids by cytochrome P450 s, as shown by the decreased larval growth in the presence of piperonyl butoxide, a P450 inhibitor. Even though T. ni tolerates C. maculatum alkaloids, the use of this species as a host plant could be ecologically disadvantageous due to prolonged larval growth and thus increased exposure to predators. Novel plant secondary compounds do not guarantee increased resistance to generalist herbivores.     

Allelochemical effects of ergoline alkaloids fromIpomoea parasitica onHeliothis virescens

Lysergol, elymoclavine and three other van Urk-positive alkaloids were identified by 2D chromatography in the seed extract ofIpomoea parasitica. The presence of the same ergoline alkaloids was also demonstrated in vegetative tissue ofI. parasitica. Heliothis virescens larvae reared on a diet containing an alkaloid extract ofI. parasitica showed an increase in the consumption index and a reduction in the effiency of conversion of food. No difference was observed in the approximated digestibility. The percentages of pupation and emergence were reduced and the malefemale ratio was altered in insects fed on a diet containing the alkaloid extract ofI. parasitica or lysergol. The increases in the concentration of ergoline alkaloids inI. parasitica at the seedling stage and at flowering support the theory that these compounds play a defensive role against herbivory in the plant.     

Substratum preferences in planula larvae of two species of scleractinian corals, Goniastrea retiformis and Stylaraea punctata

To test whether coral planulae recruit randomly to different coral reef habitats or have specific substratum preferences, the settling behavior of planulae from two shallow water coral species from Pago Bay, Guam (13°25.02N, 144°47.30E) were examined in the laboratory in June and July of 1995. Goniastrea retiformis is generally restricted to the shallow reef front (<10 m depth) in areas dominated by crustose coralline algae (CCA), while Stylaraea punctata is abundant on inner reef flats were CCA coverage is low and sand and carbonate rubble covered by biofilms is common. When presented with four substrata (1) carbonate rock scrubbed free of biofilm and dried as a control, (2) the CCA Hydrolithon reinboldii, (3) the CCA Peyssonelia sp., and (4) naturally conditioned carbonate rubble covered by a biofilm, G. retiformis larvae showed a significant preference for H. reinboldii, and S. punctata larvae for the carbonate biofilm treatment. The preference shown by S. punctata larvae for biofilmed surfaces did not diminish with increasing larval age up to 11 days. These results suggest that the larvae of both species are capable of habitat selection, and that the preferred substrata among those tested bears a relationship to the habitats in which adult colonies were found. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Effects of furoquinoline alkaloids on the growth and feeding of two polyphagous lepidopterans

Summary. The furoquinoline alkaloids skimmianine and dictamnine were tested for effects on the feeding and growth of larvae of the generalist lepidopteran, Spodoptera litura. Skimmianine was further investigated for toxic and phototoxic effects on the larvae of Trichoplusia ni, also a generalist species. In feeding experiments with S. litura, growth and consumption of larvae decreased with increasing concentration of furoquinolines; skimmianine caused a greater reduction in growth than dictamnine. In T. ni, dietary skimmianine reduced growth and consumption; when administered topically, it significantly reduced consumption but without a concomitant reduction in growth. Phototoxicity of skimmianine was not apparent in T. ni because UV light failed to increase the negative effects of the alkaloid on larvae.     

How do slugs cope with toxic alkaloids?

Summary. Arion lusitanicus and other slugs are able to feed on a variety of plants, even those containing toxic secondary metabolites, such as alkaloids. Alkaloids, like sparteine, lupanine, quinidine and atropine are avoided in no-choice feeding experiments only when they are present in concentrations above 1 mg/g food pellet. Alkaloids (sparteine, lupanine, cytisine, quinidine, atropine, senecionine, eserine, and harmaline) are toxic to Arion lusitanicus when injected. LD₅₀ values are 10 to 20 fold higher than in vertebrates indicating that slugs have a high tolerance towards food toxins. The tolerance is higher in young animals than in adult slugs. Injected alkaloids are rapidly detoxified within 72 h. Tolerance and detoxification can be induced by feeding slugs on non-lethal doses of lupin alkaloids. Using isolated microsomal preparations from the digestive gland, active detoxification was observed in vitro. Evidence is presented that cytochrome p450 plays an important role in detoxification of the applied alkaloids. A powerful and inducible detoxification systems appears to be the main mechanism that allows slugs to feed on plants rich in secondary metabolites (when no other food is available), that are usually avoided by other herbivores.     

Systemically-induced response of cabbage plants against a specialist herbivore, Pieris brassicae

Summary. Plant responses to herbivory might directly affect the herbivore (“direct” defences) or might benefit the plant by promoting the effectiveness of natural antagonists of the herbivores (“indirect” defences). Brussels sprouts attacked by Pieris brassicae larvae release volatiles that attract a natural antagonist of the herbivores, the parasitoid Cotesia glomerata, to the damaged plant. In a previous study, we observed that feeding by caterpillars on the lower leaves of the plant triggers the systemic release of volatiles detectable by the parasitoids from upper leaves of the same plant.?The role of these systemically induced volatiles as indirect defence and the dynamics of their emission were investigated in wind-tunnel dual choice tests with C. glomerata. The systemically induced emission of volatiles varied depending on leaf age and on plant age. Systemic induction affected parasitoid effectiveness, as induced plants could be more easily located by parasitoids than non-induced ones.?The role of the systemic induction as a direct defence was investigated through behavioural and feeding tests with P. brassicae. In dual choice assays, 1st instar larvae preferred to feed and fed more on systemically induced than on non-induced leaves. In single choice assays, the leaf area consumed by caterpillars was larger on systemically induced leaves than on non-induced control leaves. However, caterpillars fed on systemically induced leaves attained the same weight as those feeding on non-induced controls. In addition, P. brassicae pupae whose larvae were fed on systemically induced leaves had longer developmental times than those of larvae fed on non-induced leaves. Adult oviposition behavior was not influenced by systemic induction.?We conclude that systemically induced responses in cabbage might reduce P. brassicae fitness both directly, by affecting their development and feeding behavior and indirectly by making caterpillars and pupae more vulnerable to attack by carnivores. The occurrence of a possible relationship between direct and indirect defence is discussed. Received 24 January 2001; accepted 3 May 2001.     

Phenological fate of plant-acquired pyrrolizidine alkaloids in the polyphagous arctiid Estigmene acrea

Summary. The alkaloid profiles of the life history stages of the highly polyphagous arctiid Estigmene acrea were established. As larvae individuals had free choice between a plain diet (alkaloid-free) and a diet that was supplemented with Crotalaria-pumila powder with a known content and composition of pyrrolizidine alkaloids (PAs). Idiosyncratic retronecine esters (insect PAs) accounted for approximately half of the PAs recovered from the larvae. These alkaloids were synthesized by the larvae through esterification of dietary supinidine yielding the estigmines, and esterification of retronecine yielding the creatonotines. The retronecine is derived from insect-mediated degradation of the sequestered pumilines (macrocyclic PAs of the monocrotaline type). With one exception, the PA profiles established for larvae were found almost unaltered in all life-stages as well as larval exuviae and pupal cocoons. The exception is the males, which in comparison to pupae and adult females, showed a significantly decreased quantity of the creatonotines and pumilines. These data support the idea that the creatonotines are direct precursors of the PA-derived male courtship pheromone, hydroxydanaidal. Crosses of PA-free males with PA-containing females and vice versa confirmed an efficient trans-mission of PAs from males to females and then from females to eggs. In single cases a male bestowed almost his total PA load to the female, and a female her total load to the eggs. The results are discussed with respect to pheromone formation, PA transmission between life-stages, and the defensive role of PAs against predators and parasitoids     

Role of secondary metabolites and pigments in the epidermal tissues, ripe ovaries, viscera, gut contents and diet of the sea cucumber Holothuria atra

The epidermal tissues, ovaries, viscera, gut contents, and the diet of Holothuria atra (Jaeger), collected from the Great Barrier Reef (GBR), Australia, just prior to spawning, contained carotenoid pigments, mycosporine-like amino acids (MAAs) and three other metabolites in varying proportions. Mycosporine-glycine (mycosporine-gly), palythine, asterina-330, shinorine, mycosporine-2-gly, porphyra-334, and palythinol were present in the epidermal tissues. Mycosporine-gly was the major MAA in the epidermal tissues, gut contents and the diet, and the only MAA present in the ripe ovaries and the viscera. Gadusol, a metabolite with antioxidant properties, which co-occurs with MAAs, and in certain instances with trace amounts of 6-deoxygadusol in unfertilised and fertilised eggs and developing larvae of some marine invertebrates and vertebrates, was absent from H. atra. However, 6-deoxygadusol, with similar physical and chemical properties to gadusol, and the proposed biosynthetic precursor of MAAs, was present, but only in the ovaries. This is first report of the presence of this proposed biogenetic precursor of MAAs in significant amounts in a marine invertebrate. Approximately 90% of the total carotenoids of the epidermal tissues, ovaries and viscera of H. atra are highly oxidised, the main component being astaxanthin followed by canthaxanthin. These were the major carotenoids present in the ovaries and the total carotenoid content was highest in the ovaries. β-carotene, a common egg carotenoid, was present in trace quantities in the ovaries but, along with the xanthophylls lutein and zeaxanthin, it occurred in significant amounts in the gut contents and the diet of H. atra. Carotenoid patterns in the epidermal tissues and viscera were strikingly similar, containing β-carotene, canthaxanthin, astaxanthin, zeaxanthin, lutein, phoenicoxanthin and echinenone. The alkaloids homarine and isomeric trigonelline, (structurally unrelated to MAAs) and 6-deoxygadusol were present only in the ovaries and the viscera. It is suggested that these metabolites and pigments are either involved with photoprotection or reproduction, or associated with both processes. The origin, biogenetic relationships and the roles of these metabolites and pigments in H. atra are discussed. Received: 21 November 1997 / Accepted: 14 September 1998     

Food selection by laboratory-reared larvae of the scaled sardine Harengula pensacolae (Pisces,Clupeidae) and the bay anchovy Anchoa mitchilli (Pisces,Engraulidae)

Food selection by laboratory-reared larvae of scaled sardines Harengula pensacolae, and bay anchovies Anchoa mitchilli, was compared. Natural plankton was fed to the larvae during the 22 days following hatching. Food levels in the rearing tanks were maintained at an average of 1,600 to 1,800 potential food organisms per liter. Larvae of both species selected as food copepod nauplii, copepodites, and copepods; initial feeding was on organisms of 50 to 75 body width. Larvae of H. pensacolae averaged 4.2 mm in total length at hatching and those of A. mitchilli about 2.0 mm. H. pensacolae larvae grew about 1.0 mm per day and A. mitchilli 0.70 mm per day. The mean number of food organisms in each digestive tract was greater in H. pensacolae than in A. mitchilli, and the difference in number increased as the larvae grew. Average size of food organisms eaten increased for both species with growth, because of selection by the larvae; the average size of copepodites and copepods in digestive tracts increased at a faste rate in H. pensacolae than A. mitchilli. A. mitchilli longer than 8 mm did not eat copepod nauplii.Contribution No. 170, Bureau of Commercial Fisheries Tropical Atlantic Biological laboratory, Miami, Florida 33149, USA.     

Implications of life history for genetic structure and migration rates of southern African coastal invertebrates: planktonic,abbreviated and direct development

The amount of genetic structure in marine invertebrates is often thought to be negatively correlated with larval duration. However, larval retention may increase genetic structure in species with long-lived planktonic larvae, and rafting provides a means of dispersal for species that lack a larval dispersal phase. We compared genetic structure, demographic histories and levels of gene flow of regional lineages (in most cases defined by biogeographic region) of five southern African coastal invertebrates with three main types of larval development: (1) dispersal by long-lived planktonic larvae (mudprawn Upogebia africana and brown mussel Perna perna), (2) abbreviated larval development (crown crab Hymenosoma orbiculare) and (3) direct development (estuarine isopod Exosphaeroma hylecoetes and estuarine cumacean Iphinoe truncata). We hypothesized that H. orbiculare, having abbreviated larval development, would employ a strategy of larval retention, resulting in genetic structure comparable to that of the direct developers rather than the planktonic dispersers. However, regional population structure was significantly lower in all species with planktonic larvae, including H. orbiculare, than in the direct developers. Moreover, nested clade analysis identified demographic histories resulting from low levels of gene flow (isolation by distance and allopatric fragmentation) in the direct developers only, and migration rates were significantly higher in all three species having planktonic larvae than in the direct developers. We conclude that the amount of genetic structure within marine biogeographic regions strongly depends on the presence or absence of free-swimming larvae. Whether such larvae are primarily exported or retained, whether they have long or short larval duration, and whether or not they are capable of active dispersal seems to have little effect on connectivity among populations.