Temporal accumulation of phenylpropanoids in male fruit flies, <Emphasis Type="Italic">Bactrocera dorsalis</Emphasis> and <Emphasis Type="Italic">B. carambolae</Emphasis> (Diptera: Tephritidae) following methyl eugenol consumption

Summary. Males of dacine tephritids, Bactrocera carambolae and B. dorsalis are strongly attracted to, and compulsively feed on methyl eugenol (ME), a potent attractant for many Bactrocera species. While ME was shown to be biotransformed into phenylpropanoids, 2-allyl-4,5-dimethoxyphenol and (E)-coniferyl alcohol, and temporarily stored in the rectal gland of male B. dorsalis prior to release during courtship at dusk, B. carambolae male produces only the latter compound along with its de novo synthesized pheromone components. Both species were also shown to have different age-related response, sensitivity and consumption levels of ME. Here, we monitored and compared temporal changes in the accumulation profiles of these phenylpropanoids by the two sibling species, with male rectal glands being individually excised at different time intervals from 15 minutes to 20 days after initial ME feeding and analysed quantitatively. Results are discussed in light of plant-fruit fly co-evolution relationship.     

Penetration and storage of sponge spicules in tissues and coelom of spongivorous echinoids

When echinoids feed on sponges, silicate spicules of the sponge were found to enter their body either by penetration through the wall of the food canal into coelomic cavities or by penetration into skeletal plates and spines. The spicules, which have penetrated into the coelom, obviously evoke a kind of protective answer. They were found to be entangled by clusters of cell remnants, the so-called brown bodies. The brown bodies contain melanin and gather at special sites of the echinoid body; these are the Stewart Organs, the gills and the inner side of the ambulacral plates. Sometimes the silicate spicule becomes surrounded by a calcareous sheath. The length of the sponge spicules makes their removal impossible, so that they are stored. The spicules penetrating into the plates are partly incorporated into the stereom. The four species examined in this study were Asthenosoma ijimai, Araeosoma owstoni, Diadema setosum (collected in Sagami Bay, Japan in 1991) and Hapalosoma gemmiferum (collected in Suruga Bay, Japan in 1991).     

Formation des spicules de Didemnidae (ascidies composées). I. L'apparition des spicules chez l'oozoïde après la métamorphose

The formation of the calcareous spicules of the Didemnidae has been studied in the oozooid of Trididemnum cereum (Giard, 1872). The first spicules become visible within the paired, goblet-shaped lateral organs. From there, they appear gradually throughout the entire tunic. A histological study confirmed this in vivo observation. Moreover, in the region of the lateral organs, a certain relationship between the growing spicule and one single cell is visible, both in the light and electron microscopes. In addition, electron micrographs show that the spicule is surrounded by a double-layered membrane. Therefore, it seems unlikely that the spicules of the Didemnidae owe their existence exclusively to physico-chemical processes. Thus, the systematic value of the spicules cannot be doubted.

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Formation des spicules de Didemnidae (ascidies composées). I. L'apparition des spicules chez l'oozoïde après la métamorphose

Conclusion L'étude de la calcification chez l'oozoïde de Trididemnum cereum (Didemnidae) a permis d'observer l'apparition des premiers spicules au niveau des organes thoraciques latéraux et la distribution progressive des spicules dans la tunique. Cette observation confirme le rôle important des OTL dans la calcification, rôle déjà signalé par Michaelsen (1919a, b), Hartmeyer (1923–1924) et Van Name (1945). L'existence d'un contact étroit entre le spicule et une cellule, visible en microscopie électronique, confirme la possibilité d'une intervention cellulaire directe dans la formation du calcaire. Cette observation concorde partiellement avec les travaux de Woodland (1907).La valeur systématique des spicules ne semble plus devoir être mise en doute. Elle ne devrait plus être rejetée à l'heure actuelle, contrairement aux opinions d'Elredge (1967) et Monniot (1970).

     

Biochemical genetic divergence and systematics in sponges of the genera Corticium and Oscarella (Demospongiae: Homoscleromorpha) in the Mediterranean Sea

The sponge sub-class Homoscleromorpha is generally considered to include just two families, the Oscarellidae (without spicules) and the Plakinidae (with simple spicules). In May 1990, an unusual sponge was found deep inside a submarine cave in the western Mediterranean Sea. On the basis of externally visible characters this sponge appeared indistinguishable from the common plakinid species Corticium candelabrum Schmidt, 1862. However, on closer examination in the laboratory the new sponge proved to be devoid of spicules. Therefore, despite great morphological similarities to C. candelabrum, the new sponge should, by taxonomic convention, have been placed in the Oscarellidae. On the basis of other criteria, the similarities to C. candelabrum were great and the new sponge was at first considered to be conspecific. Thus, the taxonomic position of the new sponge and its relationship to C. candelabrum are highly confusing. It could be an aspiculate morph of C. candelabrum, or a new and undescribed related species or, lacking spicules, it could justifiably be placed in a different family (Oscarellidae). The relationship of the new sponge to C. candelabrum and also to two species of Oscarella (Oscarellidae) was assessed by the use of enzyme electrophoresis to estimate genetic divergence between species. It was found that the new sponge was reproductively isolated from sympatric C. candelabrum, with 6 of 16 loci proving diagnostic. Thus it is clear that the new sponge belongs to a different biological species. Surprisingly it was also found that, although this new species was fairly closely related to C. candelabrum (level of genetic identity, I0.47), the two Oscarella species were similarly closely related to C. candelabrum (I0.31 to 0.41) and rather less closely to the new species (I0.17 to 0.28). Indeed from genetic identity estimates, O. tuberculata is more closely related to C. candelabrum than it is to O. lobularis. It is concluded that all homoscleromorph sponges should be placed in the single family Plakinidae.     

Comparative sensitivity to and consumption of methyl eugenol in three <Emphasis Type="Italic">Bactrocera dorsalis</Emphasis> (Diptera: Tephritidae) complex sibling species

Summary. Sensitivities to methyl eugenol of three sibling species in the Bactrocera dorsalis complex were compared. The degree of species sensitivity to methyl eugenol, i.e. B. dorsalis > B. papayae > B. carambolae (in decreasing order), was concomitant with the species age-related response to methyl eugenol as previously reported. The ability to consume methyl eugenol by the three sibling species showed similar trend - the average ME consumption per male was 0.70 ml for B. dorsalis, 0.58 ml B. papayae and 0.18 μl B. carambolae. Results obtained were discussed in relation to area-wide control of fruit fly. Received 21 April 2002; accepted 9 July 2002     

Cytotoxic haplosclerid sponges preferred: a field study on the diet of the dotted sea slug<Emphasis Type="Italic"> Peltodoris atromaculata</Emphasis> (Doridoidea: Nudibranchia)

Trophic specializations are widespread among opisthobranch molluscs. One purported example from the Mediterranean Sea is the dotted sea slug Peltodoris atromaculata. It has been hypothesized that this species is strongly monophagous on the sponge Petrosia ficiformis. However, the small amount of evidence that has been found for this hypothesis is based just on laboratory tests. Here we study the feeding habits and the diet of Peltodoris atromaculata in its natural habitat. We observed and videotaped 161 individuals together with the organisms on which they were found (their living substrata). Feeding scars were identified and videotaped as well. Individuals and their living substrata were sampled for further analysis in the laboratory. The composition of faeces of Peltodoris, especially undigested sponge spicules, was analyzed by light and scanning electron microscopy and compared to the composition of the living substrata. Most of the faecal samples consisted of undigested sponge spicules. Although Peltodoris was found on 11 species of sponges, only 2 of them, Petrosia spp. and Haliclona fulva, form its diet (76% out of n=121 samples). In accordance with this, feeding scars in the habitats were exclusively observed on these two sponges. Estimation of electivity indices suggests that Haliclona is preferred over Petrosia. One remarkable feature of the exclusive feeding of Peltodoris on Petrosia and Haliclona is that both sponges share specific fulvinol-like polyacetylenes that show cytotoxic activity in bioassays. Potential benefits and evolutionary aspects of this trophic specialization are discussed. Besides sponge-containing faeces, we found spicule-free faeces (24%, n= 29). These were very small in volume compared to sponge-containing faeces, and only few distinct structures were present. However, the use of food other than sponges is not necessarily indicated by this, because the spicule-free faeces might also represent left-overs from the stomach and digestive gland after sponge spicules have been released.Communicated by O. Kinne, Oldendorf/Luhe     

Comparison of phenylpropanoid volatiles in male rectal pheromone gland after methyl eugenol consumption, and molecular phylogenetic relationship of four global pest fruit fly species: Bactrocera invadens, B. dorsalis, B. correcta and B. zonata

Males of many tephritid fruit fly species of the genus Bactrocera show a very strong affinity to methyl eugenol (ME). An attracted male compulsively ingests ME, which is then biotransformed before its metabolites are accumulated into the rectal gland. The glandular organ is known to serve as a reservoir for sex pheromone in some species. Upon ME-feeding, males of the oriental fruit fly, Bactrocera dorsalis, selectively accumulated two metabolites, 2-allyl-4,5-dimethoxyphenol (DMP) and (E)-coniferyl alcohol (E-CF), in the rectal pheromone gland. We compared the profiles of phenylpropanoid metabolites accumulated by three other species of very high economic and quarantine importance—Bactrocera invadens, Bactrocera zonata and Bactrocera correcta, with that of B. dorsalis. Males of each species were fed artificially on ME and the metabolites stored in the rectal glands were examined by means of chromatography and spectroscopy. Similar to B. dorsalis, males of laboratory-raised B. invadens accumulated DMP and E-CF, in almost equal quantities, in the rectal sac. The sum of DMP and E-CF increased gradually with time after ME consumption and reached as high as 150 μg/male 2 days post ME-feeding. Wild males of B. invadens captured in Kenya also possessed both the compounds in varying quantities. In contrast, males of B. zonata accumulated DMP and (Z)-coniferyl alcohol (Z-CF) in an approximate ratio of 1:1; whereas B. correcta is known to convert ME to (Z)-3,4-dimethoxycinnamyl alcohol (Z-DMC) and Z-CF also in an approximately 1:1 ratio. Thus, there are three types of binary combinations of rectal phenylpropanoid volatiles (i.e. DMP + E-CF; DMP + Z-CF; Z-CF + Z-DMC) utilized among the four Bactrocera species. Such differences in phenylpropanoid ingredients may play a critical role in differentiating these species if encountered in the natural habitat. In this context, the two putative sibling species—B. invadens and B. dorsalis, possess the identical subset of rectal volatiles (DMP and E-CF) in a similar proportion. Furthermore, the phylogenetic analyses of the four Bactrocera species by comparing nucleotide sequences in the mitochondrial genes showed that B. invadens clearly belonged to the same clade as B. dorsalis species. Therefore, we consider the two as the same biological species, and certainly not distinct.     

Calcium uptake and net calcification rates in the octocoral Eunicella papillosa

This paper presents the results of a preliminary study on the uptake of ⁴⁵Ca in the mesoskeleton of the gorgonian Eunicella papillosa (Octocorallia). Hourly seawater aliquots over 12 h periods displayed considerable fluctuations of activity with time, reflecting a disappearance and reappearance of tracer during the experiment. Analysis of total coenenchyme mass showed an initial rapid but fluctuating uptake of tracer and a progressive decrease with time. Distinct differences in the calcification rate of specific growth regions of the coral were detected, the rate being higher in branch tips than in lower branch regions. Chase experiments on colonies previously incubated for 24 h revealed that up to 45% of the tracer taken up during labelling were returned to the seawater. A comparison of the tracer content of the coenenchyme and the calcite spicules revealed that up to 70% of the calcium taken up remained in the coral tissue. Quantification of isotopic exchange phenomena with dead corals and isolated spicules emphasized the importance of the live coral tissue as a barrier and regulator of Ca uptake. The results are discussed in the light of the limitations of radioisotope techniques for the determination of calcification rates, and an attempt to compare rates with data in the literature is made.     

Gonad development and the planulae of the temperate Australian soft coral Capnella gaboensis

Capnella gaboensis Verseveldt, 1977 was sampled at four sites in Sydney Harbour, during 1981–1984. This soft coral has an annual cycle of gonad development, with gonad number reaching a peak in May several weeks prior to spawning, and gonad size reaching a peak in May–June at spawning. The gonads develop during the warm months, and colonies spawn their gametes in late autumn and early winter. Gonad development is neither synchronous within colonies nor within populations, possibly reflecting the protracted nature of spawning. The histology of the developing oocytes and spermaries is described in detail C. gaboensis is a surface-brooder. The planulae are similar in structure to the larvae of other octocorals. The larvae are benthic, settling quickly upon suitable substratum, metamorphosing into polyps with mouths, tentacles and spicules, approximately one week after settling.     

Determination of canthaxanthin in the red coral (<Emphasis Type="Italic">Corallium rubrum</Emphasis>) from Marseille by HPLC combined with UV and MS detection

Corallium rubrum is the most famous and precious coral due to the intense characteristic red colour of its skeleton. We have determined the presence of carotenoids in natural samples of C. rubrum from Marseille, Riou, and investigated their chemical composition. Analysis was performed both on soft tissues and hard tissues including spicules and skeleton. Since hard tissues are made of a mineral fraction and an organic fraction obtained after demineralization, extraction was performed both with and without demineralization by EDTA. The extracts were analyzed by complementary methods of TLC, HPLC/DAD and HPLC/MS. The components were separated by RP-18 chromatography column using acetonitrile/methanol for HPLC/DAD and acetonitrile/water gradient for HPLC/MS analysis. Our results give the first evidence of the presence of canthaxanthin, 4,4′-diketo-β-carotene, as the major carotenoid in all samples. Spicules exhibited higher content in carotenoids than skeleton. Demineralization treatment improved the efficiency of carotenoid extraction by an average factor of 5 and showed that organic matrix contains canthaxanthin.     

Secondary metabolite and inorganic contents in<Emphasis Type="Italic"> Cystodytes</Emphasis> sp. (Ascidiacea): temporal patterns and association with reproduction and growth

Temporal variation is one of the least known components of defence production in marine organisms. Here we examined whether there is a predictable temporal pattern in the production of chemical and structural defences by a marine invertebrate. To assess the seasonal variation in chemical defence, we measured ascididemin, the main pyridoacridine alkaloid in the blue Mediterranean morph of the ascidian Cystodytes sp. Structural defence variability was assessed on the basis of colony ash content, as it contains mainly spicules. Ascididemin concentration and the colony ash content displayed an annual cycle, reaching a minimum in spring and peaking in summer. Cross-correlation analyses with existing data on growth and reproduction suggested a significant trade-off between investment in reproduction and the other biological parameters considered (growth, inorganic content and ascididemin concentration). Our results suggest that optimization of resource allocation, probably influenced by biotic interactions and physical factors, shaped the temporal trends observed in secondary metabolite concentration and inorganic content.     

Inorganic skeleton of the demosponge Halichondria panicea. Seasonality in spicule production in the Baltic Sea

Seasonality in spicule production of the demosponge Halichondria panicea (Pallas, 1766) was studied in the Western Baltic from July 1992 to July 1993. Spicule production increased in parallel with increasing temperatures in spring, being most pronounced during summer after release of larvae. Large proportions of up to 30% immature spicules occurred in the population during skeleton production peaks, and this resulted in reduction of mean spicule dimensions by ≃20%. Effects on spicule width were more obvious than on spicule length. In winter, when wave forces were largest, higher percentages of mineral skeleton were observed: 50% of dry weight compared to 40% in summer. Enhanced spicule densities in the tissues were not attained by marked spicule production but through a reduction in organic content by shrinking. In spring, growth of body tissue was faster than the build-up of skeleton, so that the relative amount of the latter was reduced. The indirect, growth-related developments of the skeleton were inversely proportional to temperature and directly to salinity, silicate concentrations and wave energy. Spicule size and the number of immature spicules were correlated with temperature and only slightly with salinity. No correlation with oxygen concentration or pH could be detected. Received: 3 April 1997 / Accepted: 8 August 1997     

Sponge-feeding fishes of the West Indies

In an analysis of the stomach contents of 212 species of West Indian reef and inshore fishes, sponge remains were found in 21 species. In eleven of these, sponges comprised 6% or more of the stomach contents; it is assumed that these fishes feed intentionally on sponges. Sponges comprise over 95% of the food of angelfishes of the genus Holacanthus, over 70% of the food of species of the related genus Pomacanthus, and more than 85% of the food of the filefish, Cantherhines macrocerus. Lesser quantities of sponges are ingested by the remaining fish species. Fishes that feed on sponges belong to highly specialized teleost families, suggesting that this habit has evolved in geologically late time. The small number of fish species that concentrate on sponges as food suggests that the defensive characters of sponges—mineralized sclerites, noxious chemical substances, and tough fibrous components—are highly effective in discouraging predation. The two sponges most frequently eaten by fishes have a low percentage of siliceous spicules relative to organic matter, but among the 20 next most frequently consumed species no striking correlation occurs with respect to spicule content. Color and form of the sponge show no special correlation with frequency of occurrence in fish stomachs. Three species of fishes appear to concentrate on one species of sponge, but in these cases over 60% of the food taken consists of a variety of other organisms. Those fishes, more than half of whose diet consists of sponges, tend to sample a wide variety of species. No strong evidence is provided by our data that fish predation is a significant factor in limiting sponge distribution in the West Indian region.     

Effect of dye auxiliaries on chemical oxygen demand and colour competitive removal from textile effluents using Posidonia oceanica

The use of an abundant and widely distributed seagrass species, Posidonia oceanica, as a biosorbent for the direct dye Yellow 44 was successfully shown. The studies were performed on the single dyestuff dissolved in water and in a dyebath containing agents commonly used in the textile industry, i.e. a surfactant (Lavotan TBU), a sequestring agent (Meropan DPE), a softening agent (Eurosoft CI10) and a salt (NaCl). The colour reduction results showed that P. oceanica was found to be more efficient for removal of Yellow 44 from an aqueous solution (162 mg g^?1) than from a dyebath solution (135 mg g^?1), according to the Langmuir isotherm model. For the single dyestuff sorption, Fourier transform infrared and X-ray photoelectron spectroscopy studies highlighted chemical sorption between the dye alcohol function and the sorbent acid function. The chemical oxygen demand removal percentages were found to be 54.9 and 76.6% for Yellow 44 dissolved in aqueous solution and in dyebath solution, respectively. This confirmed the both sorption of the chemical auxiliaries and the dye on P. oceanica.     

Effect of temperature on life cycles of nematodes associated with the mangrove (Rhizophora mangle) detrital system

The effect of temperature on the life history of various representatives of the meiofauna associated with decaying mangrove (Rhizophora mangle) leaves was investigated. Life cycles, recorded in days, for 6 species of marine nematodes cultured at a temperature of 24°C are as follows: Rhabditis marina 2 1/4; Diplolaimelloides sp., 7; Diplolaimella ocellata, 11 1/2; Enoplus paralittoralis, 22; Oncholaimus sp., 29; Haliplectus dorsalis, 34. In general, life cycles become shorter with increased temperatures; however, as temperatures approach the upper limits which support reproduction, life cycles become slightly lengthened. For most species, the ability to complete a life cycle was inconsistent within the temperature range of 33° to 35°C. Studies with two harpacticoid copepod and two foraminifera species tend to support the 33° to 35°C range as being the thermal stress zone.Contribution No. 1697 from the University of Miami, Rosentiel School of Marine and Atmospheric Science, 10 Rickenbacker Causeway, Miami, Florida 33149, USA.     

Regional differences in duration of the planktonic larval stage of reef fishes in the eastern Pacific Ocean

Regional variation in the duration of the planktonic larval phase of three species of reef fishes, Thalassoma lucasanum (Labridae), Stegastes flavilatus, and Microspathodon dorsalis (Pomacentridae) was investigated between 1982 and 1991 at several sites in the tropical eastern Pacific over a distance of 3500 km, encompassing virtually their entire range of distribution. Durations of the larval phase, determined from counts of daily otolith increments, were significantly different (1.3 to 1.6 x) between sites. Populations of all three species had a consistently shorter larval life at the most northern site, Cabo San Lucas (Mexico) compared to Panamá and the offshore islands of Galápagos and Cocos. Analyses of otolith increment width over the precompetent period revealed that this disparity in larval duration primarily reflected differences in larval growth rates: faster growing fish spent less time in the plankton. In T. lucasanum, some of the variation in larval duration between Panamá and offshore sites (Galápagos Islands and Cocos Island) may be accounted for by a higher frequency of individuals delaying metamorphosis at the offshore sites. These data indicate that conditions in the planktonic environment are not homogeneous throughout the tropical eastern Pacific and may have a profound effect on aspects of the larval ecology of reef fishes in this region.     

Photo- and electro-catalysis for solar disinfection of Escherichia coli-contaminated water using Ag–TiO2/graphite

A titanium dioxide film on a graphite substrate was synthesized by chemical bath deposition from TiCl₄ as precursor and with the surfactant cetyl trimethyl ammonium bromide as a linking and assembling agent. Silver was loaded on the TiO₂ film by electrodeposition at 0.025?A. Water contaminated with Escherichia coli was disinfected under sunlight irradiation by photolysis (Lys), photocatalysis (PC), photoelectrocatalysis (PEC), and electrocatalysis (EC). The highest rate constant, k, was achieved with EC; k was 5.1?×?10^?2 colony forming units (CFU) mL^?1?min^?1. However, auto-oxidation of Ag occurred during EC and PEC. Meanwhile, the rate constant of disinfection by means of PC was lower than EC and PEC, and k was 3.82?×?10^?2 CFU?mL^?1?min^?1. Nevertheless, the auto-oxidation of Ag in the Ag–TiO₂/graphite tablet did not occur during the disinfection process.     

Silica uptake of the marine sponge Halichondria panicea in Kiel Bight

The silica uptake of Halichondria panicea Pallas, 1766 was measured in laboratory experiments from February until August 1995. Uptake rates were determined by measuring the decrease of dissolved silica in the surrounding seawater. All specimens were collected in Kiel Bight and maintained in aquaria up to several weeks prior to the experiments. Up to 5.27 μmol Si g⁻¹ AFDW h⁻¹ were incorporated by the sponges, with a strong, size-independent individual variation. A positive correlation between the content of dissolved silica in the seawater and the silica uptake was found. Temperature had no decisive effect on uptake rates. The nutritional condition of the sponges turned out to be of great importance. After 1 wk of starvation, Halichondria panicea uptake rates were only 15% of the previous amount, which indicates that silica transport is an energy-consuming process. During the most intense phase of reproduction activity in spring, female specimens showed a significant drop in their silica uptake. Obviously they did not produce spicules during this time. An attempt to estimate the influence of H. panicea on its habitat as a consumer of dissolved silica led to the conclusion that in summer the sponges might affect the phytoplankton species composition by competing with diatoms for dissolved silica. Received: 4 August 1996 / Accepted: 14 October 1996     

Chemical defense and evolutionary trends in biosynthetic capacity among dorid nudibranchs (Mollusca: Gastropoda: Opisthobranchia)

Summary. An evolutionary scenario incorporating recent advances in phylogenetic research begins with an opisthobranch-pulmonate common ancestor that was herbivorous and had some diet-derived chemical defense. The Nudibranchia and their closest relatives, the Notaspidea, form a lineage the ancestors of which had switched to feeding upon sponges and deriving protection from metabolites contained in them. Subsequently there have been repeated shifts in food and defensive metabolites, and trends are evident in the ability to detoxify, sequester and utilize metabolites from food, as well as to synthesize defensive compounds de novo. The Notaspidea display a minor adaptive radiation that foreshadows a more extensive one in the various lineages of nudibranchs. This review emphasizes changes that have occurred within the Holohepatica, or dorid nudibranchs (order Doridacea). Their sister-group, the Cladohepatica, consists of three other orders, Dendronotacea, Arminacea, and Aeolidiacea, in which there has been a shift from sponges to Cnidaria as food. The Dendronotacea often feed upon Octocorallia, which combine spicules, chemical defense, and stinging capsules and thereby suggest a transition from feeding on sponges. A previous diet of Octocorallia is suggested by the defensive use of prostaglandins in the dendronotacean Tethys fimbria, which eats crustaceans. A shift to bryozoans in some Arminacea is accompanied by use of different metabolites. Dorid nudibranchs evidently began as sponge-feeders, but some lineages have shifted to a variety of other food organisms, and others have specialized in the kind of sponges they feed on and how they do it. There have been shifts to bryozoans (Ectoprocta) and ascidians (Chordata: Urochordata) that track metabolites rather than the taxonomy of the food. There is a crude correlation between the genealogy and the defensive metabolites of the sponge-feeding dorids. De novo synthesis is well documented in this order and the metabolites are appropriately positioned so as to have an adaptive effect. The hypothesis that the capacity for de novo synthesis was acquired by gene transfer across lineages is rejected, partly on the basis of different chirality of metabolites in the nudibranchs and their food organisms. Instead it is proposed that there has been a preadaptive phase followed by evolution in a retrosynthetic mode, with selection favoring enzymes that enhance the yield of end products that are already present in the food. Received 5 February 1999; accepted 26 July 1999     

Fight or flight? A geographic mosaic in host reaction and potency of a chemical weapon in the social parasite <Emphasis Type="Italic">Harpagoxenus sublaevis</Emphasis>

Ant social parasites use chemical warfare to facilitate host colony takeover, which is a critical but recurring step in their life cycle. Many slave-making ants use the secretion of the Dufour gland to manipulate host behaviour during parasitic nest foundation and slave raids. Harpagoxenus sublaevis applies this chemical weapon onto defending Leptothorax host workers, which elicits deadly fights amongst them. Host species are expected to evolve counter-adaptations against this behavioural manipulation and in this study we investigated the geographic structure of this co-evolving trait. We compared the effectiveness of the parasitic gland secretion from different H. sublaevis populations in host colonies from various sites and analysed the occurrence of local adaptation. The two host species L. muscorum and L. acervorum generally showed different responses to the parasites’ chemical weapon: L. acervorum attacked nestmates treated with Dufour gland secretion, while L. muscorum workers fled. Flight, instead of intraspecific fights, is an adaptive host reaction as it results in fewer host fatalities during raids. Beside interspecific host differences, we found a geographic mosaic of host resistance: parasites from a German population strongly manipulated the behaviour of both sympatric Leptothorax populations. Russian or Italian hosts instead did not react with intracolonial aggression, but fled when confronted with the gland secretion of their sympatric parasite. Not only variation in host resistance explains differences in the effectiveness of the parasitic gland secretion but also interpopulational differences in its chemical composition, which were revealed by gas chromatography and mass spectrometry.