The complex nature of progenetic species — examples from Mesozoic ammonites

Several examples of progenetic species among Mesozoic ammonites are investigated. The ammonites are Middle Jurassic kosmoceratids, Upper Jurassic oppeliids, and Upper Cretaceous scaphites. As assessed through outgroup comparison, the progenetic species in each of these examples is characterized by small size at maturity and the presence of morphological features typical of those of the juveniles of more primitive species. However, in none of these examples is the progenetic species a duplicate form of the juvenile stage of more primitive species. There are at least three kinds of features of progenetic species that differ from those of the juveniles of more primitive species: (1) symplesiomorphous features associated with maturation, (2) unique features whose presence is not clearly related to progenetic processes, and (3) unique features that may represent epigenetic outgrowths of juvenile morphology. This novel combination of juvenile. adult. and unique features may endow progenetic species with the evolutionary potential to play a role in the origin of higher taxa. □ Progenesis, heterochrony, Ammon-oidea. Jurassic, Cretaceous.     

Progenesis in dicyemids

Dicyemids (Phylum Dicyemida) are the most common and characteristic endosymbiont living in the renal sac of benthic cephalopod molluscs. Precocious development of a hermaphroditic gonad occurs in the larvae and smaller juveniles of 40 dicyemid species from 17 cephalopod species so far and is the usual phenomenon in dicyemids. Based on the developmental and morphological features of precocious individuals, progenesis (a form of heterochrony) is the appropriate term for such precocious development. In general, progenetic individuals have much lower fecundity than normal ones because of their smaller body size, and therefore, it appears to be a disadvantageous reproductive trait. Nonetheless, the number of progenetic individuals consists of 30%–50% of the population, a relatively large proportion suggesting that the presence of progenetic individuals probably plays an important role in life history strategy. Precocious development significantly reduces growth time and enables early maturation. Progenetic individuals are common in short-living cephalopod species, in which precocious development seems appropriate for dicyemids, enabling fast larval release before the end of the host's life span.     

Ultrastructure of the ventral pharynx in the interstitial annelid <Emphasis Type="Italic">Questa paucibranchiata</Emphasis> (Orbiniidae) and its phylogenetic significance

The small taxon Questa comprises only meiofaunal species and shows oligochaete-like adaptations in its reproductive biology. Formerly, it was classified as a separate polychaete group, Questidae, of uncertain affinities, but due to these oligochaete-like similarities, a relationship with Clitellata was discussed. However, other morphological phylogenetic analyses questioned such a relationship, providing evidence for common ancestry with Paraonidae and Orbiniidae. Molecular studies revealed that questids even form an in-group of Orbiniidae, which refutes a clitellate relationship without doubt. On the other hand, morphological evidence supporting this grouping is still considered weak. Among the external characters, the chaetation, comprising crenulated capillaries, hooks and furcate chaetae, might strengthen this hypothesis. One of the internal characters stated to support this grouping is the ventral pharynx, which as such, however, is widespread among polychaetes. This study is a first attempt to look for specific structures in this organ, which might be regarded as synapomorphic of Questa and Orbiniidae. The ultrastructural analysis revealed a ventral buccal organ composed of large interstitial cells and transverse muscle fibres and surrounded by an investing muscle system similar to that found in juvenile Orbiniidae. Thus, further morphological evidence for the current position is provided. Moreover, since this structure is known as a larval or juvenile character in Orbiniidae, this finding is also indicative for a progenetic origin of Questa, which might have occurred upon invading the interstitial environment.     

Reproductive fitness consequences of progenesis: Sex‐specific pay‐offs in safe and risky environments

Progenesis is considered to have an important role in evolution because it allows the retention of both a larval body size and shape in an adult morphology. However, the cost caused by the adoption of a progenetic process in both males and females remains to be explored to explain the success of progenesis and particularly its biased prevalence across the sexes and environments. Here, through an experimental approach, we used a facultative progenetic species, the palmate newt (Lissotriton helveticus) that can either mature at a small size and retain gills or mature after metamorphosis, to test three hypotheses for sex‐specific pay‐offs of progenesis in safe versus risky habitats. Goldfish were used because they caused a higher decline in progenetic than metamorphic newts. We determined that progenetic newts have a lower reproductive fitness than metamorphic newts. We also found that, when compared to metamorphs, progenetic males have lower reproductive activity than progenetic females and that predatory risk affects more progenetic than metamorphic newts. By identifying ultimate causes of the female‐biased sex ratios found in nature, these results support the male escape hypothesis, that is the higher metamorphosis rate of progenetic males. They also highlight that although progenesis is advantageous in advancing the age at first reproduction, it also brings an immediate fitness cost and this, particularly, in hostile predatory environments. This means that whereas some environmental constraints could favour facultative progenesis, some others, such as predation, can ultimately counter‐select progenesis. Altogether, these results improve our understanding of how developmental processes can affect the sexes differently and how species invasions can impair the success of alternative developmental phenotypes.     

Unifying Concepts Learned from Methyl Farnesoate for Invertebrate Reproduction and Post-Embryonic Development

Since the discovery that methyl farnesoate (MF), the unepoxidatedform of the insect juvenile hormone (JHIII), is produced bymandibular organs of numerous crustaceans, extensive evidencehas accumulated that this compound appears to perform similarfunctions in the Crustacea as JH performs in insects. A majorfunction of MF appears to be in enhancing reproductive maturation.This was first shown by indirect experimentation with eyestalkablation, which augmented MF production. Subsequently, directtreatments of several species of crustacea with MF showed thatreproductive maturation was enhanced. A second function of MF, similar to that of the JH of insects,is in the maintenance of juvenile morphology. This is especiallytrue in the late larval transformations into juveniles, whereMF plays an inhibitory role, as well as during the transformationof juveniles into adults. These results were inferred from eyestalkremoval experiments. In the case of the larval-juvenile transition,inhibitory results were also obtained with MF by direct hormonetreatments. However, the transition from very early larval stages,such as one nauplius stage proceeding to the next, which inmany cases also involves morphogenetic changes, may be occurringin the presence of MF. Indeed, MF appears to be stimulatoryto early postembryonic larval stages of Crustacea. Again, thisfunction of MF in Crustacea appears to be similar to functionsof JH in early postembryonic insects. However, it should bepointed out that there are many more "early" stages in Crustaceathan there are in insects, and very few of these cases havebeen investigated. When considering the animal kingdom and larval metamorphosis,the question may be raised whether there are other members ofthe JH family regulating metamorphosis and reproduction. Oneplausible example appears to be among certain annelids. Thetrochophores of Capitella respond to various juvenoids, butare most responsive, within one hour, to MF and eicosatrienoicacid. This latter compound is present also in adult annelids,where it has been named "Sperm Maturation Factor," since itseems to function in the maturation of sperm in Arenicola. Therefore,eicosanoids perform in annelids two functions performed in insectsby JHs. In conclusion, it seems that there are morphogenesis promotingresponses to JHs in early larval development in crustaceans,annelids, and possibly other forms, which differ from thoseMF effects in later larvae of Crustacea where MF retards morphogenesis.Such early responses as noted here have recently also been describedfor insects. Furthermore, it is clear that the polyunsaturated8,11,14-eicosatrienoic and aracidonic acids seem to be juvenoids,and appear to function as such in annelids, and may also befunctionally active in insects and crustaceans. It seems reasonableto conclude therefore that new and novel juvenoids exist, whileothers still await discovery.     

Male morphological variation and the determinants of body size in two Otiteselline fig wasps

The chalcid wasps (Hymenoptera) that develop in fig tree inflorescences(Ficus: Moraceae) have often been used to study alternativereproductive behaviors. However, recent work suggests that suchbehaviors are more complex than previously thought. We investigatedthis in Otitesella longicauda and O. rotunda. In addition toknown dimorphisms in the two species (each have "religiosa"males that use their mandibles to fight for mates in the fig,and "digitata" males that disperse from the fig to mate), wefound that religiosa males below species-specific body sizeswitch points have relatively larger mandibles and are lesssclerotized than those above. Thus, they are actually trimorphic.We suggest that the religiosa morph variation is linked to fighter/nonfighteralternative mating behaviors, with small (nonfighting) maleshaving relatively larger mandibles because they also use themto pull females out of their galls before mating. Also, we investigatedthe determinants of wasp body size, and whether females (foundresses)adjust their offspring allocation strategies according to expectedoffspring size. We found that wasp size is larger in ovariesnear the center of the fig, and more females and fewer religiosamales are laid in such ovaries than in those further away. Thisprobably indicates that foundresses lay females when they areexpected to be large because their fitness is more body size–dependentthan that of religiosa males. We then discuss the implicationsof our findings for the study of alternative reproductive behaviorsand foundress offspring allocation strategies.     

The dynamics of sponge larvae assemblages from northwestern Mediterranean nearshore bottoms

In this study, we show for the first time the dynamics of spongelarvae assemblages from nearshore meroplankton. Plankton wascollected by SCUBA diving once or twice a week during a 2-yearperiod over a rocky artificial reef in the NW Mediterranean.Data on larval abundance were cross correlated with the valuesof environmental parameters (i.e. seawater temperature, solarradiation, wind speed and atmospheric pressure). In the laboratory,we recorded external features and main behaviors of larvae.We collected larvae belonging to 20 different taxa of sponges,which are among the most common in the sublittoral hard bottomcommunities of the NW Mediterranean and other temperate areas.There was a positive correlation between maximum abundance oflarvae and highest water temperatures. Maximum solar radiationpreceded the maximum of larval abundance. Wind speed showedno clear seasonal patterns and atmospheric pressure was overallthe lowest when larvae were most abundant. Two main patternsin the larval release periods were observed. One was shown byspecies releasing larvae in summer, right before the maximumwater temperatures (orders Dictyoceratida and Dendroceratida)and another by the species whose larvae release from the endof summer till autumn, when temperatures decrease (order Poecilosclerida).The larvae of Phorbas tenacior, Raphidoflus jolicoueri, Mycalerotalis, Tedania anhelans, Pleraplysilla spinifera, Aplysillasulfurea var. rosea and Chelonaplysilla noevus are describedfor the first time. The larvae collected mainly belonged tothe parenchymella type (except for the species Oscarella sp.and probably Cliona viridis) and showed different features andbehaviors: from the elongated parenchymellae of Scopalina lophyropoda(order Halichondrida), which show simple swimming behavior andno response to light, to the parenchymellae of Poeciloscleridaand Dictyoceratida orders with variable morphologies as adaptationsto complex swimming behaviors. Our database will hopefully contributeto the present knowledge of larval types in sponges and definitivelyhighlight the importance of this group in the dynamics of meroplanktonfrom nearshore bottoms.     

Progenesis in Eunicida ("Polychaeta," Annelida)--separate evolutionary events? Evidence from molecular data

The Eunicida are a well-defined annelid taxon currently comprising Dorvilleidae, Lumbrineridae, Eunicidae, Onuphidae, and Hartmanniellidae. Especially the Dorvilleidae include several species of small body size and apparently simple organisation, resembling larvae and juveniles of larger species. One hypothesis to explain the evolution of such species is progenesis. By this criterion the Dinophilidae were included in this group as well. Although their overall similarity suggests monophyly, parallel evolution of such progenetic taxa cannot be excluded. To infer the phylogeny of Eunicida and their presumed relationship to Dinophilidae and progenetic dorvilleids, 18S rDNA sequences from various Dinophilidae, Dorvilleidae, and members of other Eunicida were determined. The data matrix were extended by adding sequences from several annelid and molluscan species and analysed with maximum-parsimony, distance, and maximum-likelihood algorithms. Highly significant bootstrap-values support monophyly of the Dinophilidae, of a clade enclosing the Eunicidae and the Onuphidae, and of a dorvilleid group comprising six species including the presumed progenetic Parapodrilus psammophilus. The position of the Dinophilidae remains unresolved, so that their inclusion within the Eunicida can neither be corroborated nor rejected. However, a constraint monophyly of Dinophilidae and P. psammophilus is rejected by several tests, substantiating the hypothesis of separate progenetic evolutionary events. Furthermore, the molecular data indicate polyphyly of the Dorvilleidae.     

Ultrastructure of pigmented eyes in Dorvilleidae (Annelida,Errantia, Eunicida) and their importance for understanding the evolution of eyes in polychaetes

Among polychaetes, the errant forms are the only group known so far possessing true multicellular eyes in adults which are preceded by bicellular larval eyes in many species. Most likely, two pairs of such eyes showing a specific structure belong to the ground pattern of Errantia = Aciculata. However, these eyes have primarily been investigated in only two subgroups of Errantia, but data on the third main taxon, Eunicida, are available for only two taxa. In the present investigation, the eyes in two additional species of Eunicida, the dorvilleids Protodorvillea kefersteini and Schistomeringos neglecta, were studied. In P. kefersteini, usually described as possessing one pair of small eyes, two pairs could be detected, whereas in S. neglecta only one pair was found. Each eye is made up of rhabdomeric photoreceptor cells, pigment cells and unpigmented supportive cells. Lenses or vitreous bodies are absent. From their structure most likely all eyes represent adult eyes and even the small anterior eyes in P. kefersteini structurally resemble miniaturized adult eyes. Neither persisting larval eyes nor unpigmented rhabdomeric ocelli were found in the two species. The observations in Dorvilleidae confirm the hypothesis of a common origin of adult eyes in Errantia.     

Ultrastructure of pigmented adult eyes in errant polychaetes (Annelida): implications for annelid evolution

The evolution of photoreceptor cells and eyes in Metazoa is far from being resolved, although recent developmental and structural studies have provided strong evidence for a common origin of photoreceptor cells and existence of sister cell types already in early metazoans. These sister cell types are ciliary and rhabdomeric photoreceptor cells, depending on which part of each cell is involved in photoreception proper. However, a crucial point in eye evolution is how the enormous structural diversity of photoreceptor cells and visual systems developed, given the general molecular conservation of the photoreceptor cells. One example of this diversity can be observed in Annelida. Within the polychaetes the errant forms, taxon Aciculata, constitute the only group possessing true multicellular eyes in the adult stage. Thus far these organs have been investigated only in taxa of Phyllodocida, a subgroup of Aciculata. Data on Eunicida and Amphinomida as well as certain phyllodocidan taxa had been lacking. The ultrastructure of these adult eyes was investigated in various species of errant polychaetes, belonging to Amphinomidae, Eunicidae and Hesionidae, to elucidate whether they provide any phylogenetic clues regarding either the evolution of visual systems in Annelida or lophotrochozoan phylogeny in general. These eyes are composed of numerous supportive pigment cells and rhabdomeric photoreceptor cells and sometimes additional cell types. As a rule the pigment and rhabdomeric cell types form a continuous epithelium in which the two types intermingle. Presence of granules with shading pigment in sensory cells is a common feature but is apparently restricted to a taxon comprising Phyllodocida and Eunicida s. str. Very likely a lens-like structure does not belong to the ground pattern of annelid eyes, despite its presence in Phyllodocida. These lens-like structures are formed by secretions or cellular processes of the pigment cells. In many species the eye cup communicates with the exterior via a small cuticularized canal. This canal is interpreted as a rudiment due to the mode of formation in the epidermis. With respect to current phylogenetic hypotheses, these multicellular eyes have either been developed in the stem species of a taxon Aciculata nested within the polychaetes or have been evolved in the stem lineage of Annelida. Similarities to gastropod eyes are interpreted as convergent and not as indication of common origin. Except for the photoreceptor cells proper, the structure of the adult eyes in polychaetes most likely does not help to resolve lophotrochozoan phylogeny.     

A new poecilogonous species of sea slug (Opisthobranchia: Sacoglossa) from California: comparison with the planktotrophic congener Alderia modesta (Loven, 1844)

Cryptic species are increasingly recognized as commonplace amongmarine gastropods, especially in taxa such as shell-less opisthobranchsthat lack many discrete taxonomic characters. Most cases ofpoecilogony, the presence of variable larval development withina single species, have historically turned out to representcryptic species, with each possessing a single canalized typeof development. One well-characterized example of poecilogonywas attributed to the sacoglossan opisthobranch Alderia modesta;in southern California, slugs resembling this member of a monotypicgenus produce both long-lived, planktotrophic and short-lived,lecithotrophic larvae. Paradoxically, however, A. modesta isexclusively planktotrophic everywhere else in the northern Pacificand Atlantic Oceans. A recently completed molecular study foundthat slugs from poecilogonous populations south of Bodega Harbor,California, comprise an evolutionarily distinct lineage separatefrom northern, strictly planktotrophic slugs. We now describethe southern species as A. willowi n. sp., based on differencesin morphology of the dorsum and radula, characteristics of theegg mass, larval development mode and nuclear and mitochondrialgenetic markers. A DNA barcode is provided, based on 27 fixeddifferences in the cytochrome c oxidase subunit I gene thatcan reliably differentiate Pacific specimens of Alderia species.Genetic and morphological data are concordant with developmentalevidence, confirming that A. willowi is a true case of poecilogony.An improved understanding of the ecological differences betweenthese sister taxa may shed light on the selective pressuresthat drove the evolution of lecithotrophy in the southern species. (Received 1 November 2005; accepted 20 September 2006)     

Descriptions of some planktonic larvae of the Calappidae (Crustacea: Decapoda; Brachyura)

Some unidentified planktonic larval stages have been assignedto the Calappidae using Rice's key of Brachyura families. Zoeastage I from the eastern region of the Algerian coast (westernMediterranean Sea) has been attributed, according to Manningand Holthuis (1980), to Calappa granulata (L.), the only recordedrepresentative of the family from the Mediterranean Sea. Zoealstages 1-lV from the Jordanian coast of the Gulf of Aqaba, NorthernRed Sea, have been reported to the subfamily Matutinae. Comparisonsof larvae of Calappa and of Hepatus and then, of larvae of Hepatusand of Matuta, show that Hepatus larvae are more closely relatedto those of Calappa, while those of Matuta are clearly distinct.It s suggested to place the genus Hepatus with the Calappinae.Guinot (1966. 1978) placed some genera, including Hepatus. ina 'unit parthenoxystomienne, lea Aethrinae'. Larval developmentis unknown for Aethra, Actaeomorpha. Hepatella andOsachila.However, comparison of Hepatus larvae with those of Parthenopidaeshows that larval characters, particularly the setation of theendopod of the first maxilliped, are not conclusive for supportingsuch a regrouping. Moreover, the Calappidae have a combinationof primitive and advanced features which suggest that theirgrouping in the Oxystomata seems more likely to be an adaptationto a similar way of life. The morphotypal taxon Oxystornatahas been definitively rejected by Guinot (1978).     

Life history constraints on the evolution of abbreviated life cycles in parasitic trematodes

Abbreviations of the complex life cycle of trematodes, from three to two hosts, have occurred repeatedly and independently among trematode lineages. This is usually facultative and achieved via progenesis: following encystment in the second intermediate host, the metacercaria develops precociously into an egg-producing adult, bypassing the need to reach a definitive host. Given that it provides relatively cheap insurance against a shortage of definitive hosts, it is not clear why facultative progenesis has only evolved in a few taxa. Here a comparative approach is used to test whether progenetic trematodes are characterized by larger body size and egg volumes, two traits that correlate with other key life history features, than other trematodes. These traits may constrain the evolution of progenesis, because precocious maturation might be impossible when the size difference between the metacercaria and a reproductive adult is too large. First, trematode species belonging to genera in which progenesis has been documented were found not to differ significantly from other trematode species. Second, using within-genus paired comparisons across 19 genera in which progenesis has been reported, progenetic species did not differ, with respect to body size or egg size, from their non-progenetic congeners. Third, using intraspecific paired comparisons in species where progenesis is facultative, no difference was observed in the sizes of eggs produced by worms in both the intermediate and definitive host, suggesting that opting for progenesis does not influence the size of a worm's eggs. Overall, the lack of obvious differences in body or egg size between trematodes with truncated life cycles and those with the normal three-host cycle indicates that basic life history characteristics are not acting as constraints on the evolution of progenesis; trematodes of all sizes can do it. Why facultative progenesis is not more widespread remains a mystery.     

Adaptations of Host and Symbionts in the Intertidal Zone

SYNOPSIS. Different species of larval trematodes that utilizethe same intermediate host characteristically exhibit distinctivepatterns of response to environmental fluctuations. Not onlyare the responses distinctive for each species, but also theycannot be correlated with that of the host. The physiologicalresponses of the molluscan host to physical changes in the environment,however, are modified significantly by the presence of a larvaltrematode infection. The thermal metabolic acclimation patternsof infected Nassarius obsoleta are quite distinct from thoseof non-infected N. obsoleta. Thex activity patterns during thermalacclimation of cytochrome c oxidase from digestive glands ofsnails infected with different species of larval trematodesindicate that each species of larval trematodes differentiallyinfluences the hosttissue, so that the cytochrome oxidase systemdoes not respond to temperature in the same way as does uninfectedhost tissue.     

Comparative analysis of the nervous systems in presumptive progenetic dinophilid and dorvilleid polychaetes (Annelida) by immunohistochemistry and cLSM

Entire nervous systems of the dinophilids Dinophilus (two species) and Trilobodrilus (three species) and the dorvilleids Parapodrilus psammophilus and Ophryotrocha gracilis (larva) were stained with antisera directed against serotonin, Phe‐Met‐Arg‐Phe‐NH₂ (FMRFamide) and acetylated α‐tubulin and analysed by confocal laser scanning microscopy (cLSM). Adult dinophilids and the dorvilleid larva exhibit the same structure of the ventral nerve cord, with two main nerves spaced far apart, one median and two paramedian nerves. A serotonergic plexus is situated between the paramedian nerve pair, above the ventral locomotory ciliary band. These similarities between adults and larva corroborate the presumed progenetic origin of dinophilids. However, since larval nervous systems of other polychaete taxa also seem to be organized in this way, this result cannot support the view that dinophilids originate from dorvilleids. In P.psammophilus the main nerve cords are widely separated only in the last segment, indicating that this pattern may be correlated with the absence of parapodia. The unpaired median nerve of dinophilids, P. psammophilus and many other polychaetes, is considered to be part of the basic annelid body plan. The ground pattern of the ventral paired dinophilid ganglia is represented by three (anterior, main, posterior) commissures, conserved in most of the ganglia in the Dinophilus species and mostly reduced to a main commissure in the Trilobodrilus species. The dinophilid species and P. psammophilus possess six pairs of ganglia indicating six trunk segments – in contrast to former views. The two rings behind the prostomium in both the dinophilid and the dorvilleid species contain one pair of ganglia only, corroborating the presumed homology of this peristomial region in the two taxa. The Dinophilus nervous system with 12 longitudinal nerves and three perpendicular nerve rings per segment resembles orthogonal nervous structures characteristic of platyhelminths.     

The Jurassic ammonite Coelocerus: an atypical example of dimorphic progenesis elucidated by cladistics

Instances of convergence or parallelism among unrelated Lower Carixian ammonites are not uncommon. The case of polymorphitid-like ammonites is considered here, and the example of Coeloceras is investigated in detail. The small sexual dimorph (microconch) of Coeloceras pettos (Quensitedt) (Coeloceratidae), a progenetic species, provides a remarkable example of complex convergence with the microconch of Uptonia (Polymorphitidae). Homeomorphic morphology is here an amalgam of juvenile traits, of maturation-related transformations, and of completely new features bearing no obvious relation with either progenesis or maturation. A stratophenetic approach is hardly adequate for unravelling such an involved pattern of convergence. However, a cladistic analysis of ammonoid morphology suggests that Coeloceras was derived from the Tethyan Miltoceras. Indeed, greater biostratigraphic knowledge and the independent use of comparative shell morphology are both needed to improve ammonite phylogeny. □ Convergence, parallelism, progenesis, dimorphism, ammonites, Jurassic.     

污损性管栖多毛类生态特点及研究展望

污损性管栖多毛类属环节动物门,主要由龙介虫科(Serpulidae)、螺旋虫科(Spirorbidae)、缨鳃虫科(Sabellidae)和蛰龙介科(Terebllidae)4科55种组成,以华美盘管虫(Hydroides elegans)、内刺盘管虫(H.ezoensis)、龙介虫(Serpula vermicularis)和克氏无襟毛虫(Pomatoleios kraussii)等种类为优势种,通常成批的附着在物体表面,严重影响人工设施的安全和性能。污损性管栖多毛类的分布具有明显的地域性和季节性,并与深度有关。今后工作应进一步开展管栖多毛类基础生物学研究,探讨外来种的入侵及影响,丰富和发展生态和分类等方面的研究内容。     

Larval development of Hexapanopeus caribbaeus (Stimpson, 1871) (Crustacea, Decapoda, Xanthoidea, Panopeidae) reared under laboratory conditions

The genus Hexapanopeus Rathbun, 1898 contains eight species,of which three are found in the Brazilian coast. The larvaldevelopment of two of these species has already been studiedin the laboratory. This article describes in detail the morphologyof the complete larval stages of Hexapanopeus caribbaeus rearedunder laboratory conditions, and the morphology is comparedwith that of the larvae of the other species of the same genus.     

Effects of temporary food limitation on survival and development of brachyuran crab larvae

As a consequence of the combined effects of prey patchinessand diel or tidal vertical migrations in the water column, decapodcrustacean larvae may experience temporal or spatial variabilityin the availability of planktonic food. In a laboratory study,we evaluated effects of temporarily limited access to prey onthe larvae of three species of brachyuran crabs, Chasmagnathusgranulata, Cancer pagurus and Carcinus maenas. Stage-I zoeaewere fed ad libitum for 4 or 6 h per day (20 or 25% treatments;6 h tested in C. pagurus only), and rates of larval survivaland development were compared with those observed in continuouslyfed control groups (24 h, 100%). In C. granulata, we also testedif intraspecific variability in initial biomass of freshly hatchedlarvae originating from different broods has an influence onearly larval tolerance of food limitation. Moreover, we exposedembryos and larvae of this estuarine species to moderately decreasedsalinities to identify possible interactions of osmotic andnutritional stress. Finally, we evaluated in this species theeffect of food limitation on survival from hatching throughall larval instars to metamorphosis. In all three species, limitedaccess to prey had only weak or insignificant negative effectson survival through the Zoea-I stage. The strength of the effectsof temporary food limitation varied in C. granulata significantlyamong broods. However, no significant relationships were foundbetween initial larval biomass (C content) and either survivalor development duration. Strongly decreased survival to metamorphosiswas found when food limitation continued throughout larval development.Thus, early brachyuran crab larvae are well adapted to transitorylack of planktonic food. The capability of the Zoea-I stageof C. granulata to withstand nutritional stress also under conditionsof concomitant salinity stress allows them to exploit variousbrackish environments within estuarine gradients. However, continuedexposure to limited access to planktonic prey may exceed thenutritional flexibility of C. granulata larvae.