Larval development of the pedunculate barnacles Octolasmis angulata Aurivillius 1894 and Octolasmis cor Aurivillius 1892 (Cirripedia: Thoracica: Poecilasmatidae) from the gills of the mud crab,Scylla tranquebarica Fabricius, 1798

Detailed studies of larval development of Octolasmis angulata and Octolasmis cor are pivotal in understanding the larval morphological evolution as well as enhancing the functional ecology. Six planktotrophic naupliar stages and one non-feeding cyprid stage are documented in details for the first time for the two species of Octolasmis. Morphologically, the larvae of O. angulata and O. cor are similar in body size, setation patterns on the naupliar appendages, labrum, dorsal setae-pores, frontal horns, cyprid carapace, fronto-lateral gland pores, and lattice organs. Numbers of peculiarities were observed on the gnathobases of the antennae and mandible throughout the naupliar life-cycle. The setation pattern on the naupliar appendages are classified based on the segmentation on the naupliar appendages. The nauplius VI of both species undergoes a conspicuous change before metamorphosis into cyprid stage. The cyprid structures begin to form and modify beneath the naupliar body towards the end of stage VI. This study emphasises the importance of the pedunculate barnacle larval developmental studies not only to comprehend the larval morphological evolution but also to fill in the gaps in understanding the modification of the naupliar structures to adapt into the cyprid life-style.     

An early Cambrian chelicerate from the Emu Bay Shale,South Australia

The Emu Bay Shale Lagerstätte (Cambrian Series 2, Stage 4) occurs on the north coast of Kangaroo Island, South Australia. Over 50 species are known from here, including trilobites and non‐biomineralized arthropods, palaeoscolecids, a lobopodian, a polychaete, vetulicolians, nectocaridids, hyoliths, brachiopods, sponges and chancelloriids. A new chelicerate, Wisangocaris barbarahardyae gen. et sp. nov., is described herein, based on a collection of some 270 specimens. It is up to 60 mm long, with the length of the cephalic shield comprising about 30% that of the exoskeleton. The cephalic margin has three pairs of bilaterally‐symmetrical small triangular spines. A pair of small eyes is placed well forwards on the ventral margin of the cephalic shield. The trunk comprises 11 segments that increase in length while narrowing posteriorly, each possibly bearing a pair of biramous appendages; the most posterior segment is almost square whereas the others are transversely elongated. The spatulate telson is proportionately longer than in taxa such as Sanctacaris, Utahcaris and Leanchoilia. Up to eight (?four pairs) of 3 mm‐long elements bearing alternating inward‐curving short and long spines beneath the cephalic shield are interpreted as basipodal gnathobases, part of a complex feeding apparatus. A well‐developed gut includes a stomach within the cephalic shield; it extends to the base of the telson. In a few specimens there are shell fragments within the gut, including those of the trilobite Estaingia bilobata (the most common species in the biota); these fragments have sharp margins and extend across the gut lumen. The species may have been a predator or a scavenger, ingesting material already broken up by a larger predator/scavenger. The morphology of this taxon shares many overall body features with Sanctacaris, and some with Sidneyia, particularly its gnathobasic complex. These chelicerate affinities are corroborated by phylogenetic analyses.     

Larval development of Japanese 'conchostracans': part 2, larval development of Caenestheriella gifuensis (Crustacea, Branchiopoda, Spinicaudata, Cyzicidae), with notes on homologies and evolution of certain naupliar appendages within the Branchiopoda

As part of a larger project examining and comparing the ontogeny of all major taxa of the Branchiopoda in a phylogenetic context, the larval development of Caenestheriella gifuensis (Ishikawa, 1895), a Japanese spinicaudatan ‘conchostracan’, is described by scanning electron microscopy. Seven different larval stages are recognised, in most cases based on significant morphological differences. They range in length from about 200 to 850 μm. Nauplius 1 has a plumb and lecithotrophic appearance with a rounded hind body and a labrum with an incipient medial spine. Limb segmentation is mostly unclear but the second antennae have more putative segments delineated than are expressed in the later stages. Feeding structures such as the mandibular coxal process and antennal coxal spine are only weakly developed. Nauplius 2 is very different from nauplius 1 and has three large spines on the labral margin and two long caudal spines. Feeding structures such as the mandibular coxal process and various spines and setae are developed, but whether feeding begins at this stage was not determined. The mandible has developed an ‘extra’ seta on endopod segment 1, absent in Nauplius 1. The segmentation of the second antenna has changed significantly due to fusions of various early segments. Nauplius 3 is like nauplius 2 in morphological detail, but larger and more elongate. Nauplius 4 has developed a pair of small anlagen of the carapace and rudiments of the first five pairs of trunk limbs, and the coxal spine of the antenna has become distally bifid. Nauplius 5 has a larger carapace anlage, externally visible enditic portions of the elongate trunk limbs, and a pair of primordial dorsal telson setae. Nauplius 6 has a larger and partly free carapace and better-developed, partly free trunk limbs with incipient enditic, endopodal, and exopodal setation. A pair of caudal spines, dorsal to the large caudal spines, has appeared. Nauplius 7 is quite similar to nauplius 6 but is larger and has slightly longer caudal and labral spines; also, the setation of the most anterior trunks limbs is better developed. The larval development is largely similar to that of other spinicaudatans. The larval mandible, which is evolutionarily conservative within the Branchiopoda, reveals a setation pattern similar to that of the Anostraca and Notostraca (two setae on mandibular endopod segment 1). Most other spinicaudatans and all examined laevicaudatans share another setal pattern (one seta on mandibular endopod segment 1), which could indicate a close relationship among these taxa. The second antenna undergoes a special development, which provides an insight into the evolution of this limb within the Branchiopoda. In nauplius 1 the basipod, endopod, and exopod are all superficially divided into a relatively high number of segments. In later nauplii some of these have fused, forming fewer but larger segments. We suggest that this ontogeny reflects the evolution of antennae in the conchostracans. Various aspects of the morphology of the antennae are discussed as possible synapormorphies for either the Diplostraca or subgroups of the Conchostraca.     

Larval development of Octomeris sulcata Nilsson-Cantell (Cirripedia: Thoracica: Chthamalidae) from Japan and Korea

Embryos obtained from gravid adults of the chthamalid barnacle Octomeris sulcata Nilsson-Cantell from Japan and Korea were cultured through six naupliar stages to the cyprid and juvenile barnacle stage in laboratory conditions, fed either the diatom Skeletonema costatum (Grev.) Cleve or the dinoflagellate Prorocentrum minimum (Pavillard) Schiller. The nauplii were planktotrophic and, depending on diet, reached the cyprid stage 9 or 17 days after hatching in individual cultures at 22 °C with 24 h illumination. The survival rate was higher and the duration of the naupliar stages was shorter when fed P. minimum rather than S. costatum. This is probably due to the presence of feathered setae on the antennae. Feathered or plumose setae in nauplii of different cirripede taxa are apparently linked to the type of phytoplankton in the seas when these taxa first evolved.The larval stages of O. sulcata are described, and morphological differences between larvae reared from Japanese andKorean adults are compared. The polygonal cephalic shield and unilobed labrum, a pair of posterior shield spines after naupliar stage IV, feathered setae and a hispid seta on the coxa of the antenna, a cuspidate seta on the mandible, and the gnathobase of the antenna are important in distinguishing the nauplii of this species from other species, including Chthamalidae.     

Three-dimensional reconstruction of the naupliar musculature and a scanning electron microscopy atlas of nauplius development of Balanus improvisus (Crustacea: Cirripedia: Thoracica)

An atlas of the naupliar development of the cirripede Balanus improvisus Darwin, 1854 using scanning electron microscopy (SEM) is provided. Existing spikes on the hindbody increase in number with each moult and are an applicable character for identification of the different nauplius stages, as is the setation pattern of the first antennae. The naupliar musculature of B. improvisus was stained with phalloidin to visualise F-actin, followed by analysis using confocal laser scanning microscopy (CLSM) with subsequent application of 3D imaging software. The larval musculature is already fully established in the first nauplius stage and remains largely unchanged during all the six nauplius stages. The musculature associated with the feeding apparatus is highly elaborated and the labrum possesses lateral muscles and distal F-actin-positive structures. The alimentary tract is entirely surrounded by circular muscles. The extrinsic limb musculature comprises muscles originating from the dorsal and the ventral sides of the head shield, respectively. The hindbody shows very prominent postero-lateral muscles that insert on the dorso-lateral side of the head shield and bend towards ventro-posterior. We conclude that the key features of the naupliar gross anatomy and muscular architecture of B. improvisus are important characters for phylogenetic inferences if analysed in a comparative evolutionary framework.     

Chemical Defence in Cephalaspidean Gastropods: Origin, Anatomical Location and Ecological Roles

Within the opisthobranchs, the cephalaspideans are traditionallyconsidered a transitional group between typical testacean prosobranchsand shell-less opisthobranchs. The cephalaspidean anatomy, includingthe presence of a cephalic shield, is related to burrowing throughsoft sediment. Recent studies have shown that some herbivorousand carnivorous cephalaspideans contain secondary metabolites.The micro-herbivorous bubble snails of the Bullidae and Haminoeidaefamilies are known to have secondary metabolites which have differentecological roles. The polypropionates isolated from Bulla gouldianaand B. striata were deterrent to fishes while the secondarymetabolites of Haminoea callidegenita, H. fusari, H. hydatis,H. navicula, H. orbignyana and H. orteai were alarm pheromonesemployed during cross copulation. In Bulla gouldiana and B.striata, the defensive secretion was located mainly in a whitegland along the margin of the mantle. In Haminoea species, alarmpheromones were located in external parts (cephalic shield,parapodial lobes and posterior pallial lobe). The carnivorous cephalaspideans Navanax inermis and Philinopsisdepicta employ chemotaxis to follow the slime trail of theirprey, which include other cephalaspideans or even congenericindividuals. N. inermis and P. depicta sequester alarm pheromonesand allomones from their cephalaspidean prey, which are ejectedwhen N. inermis and P. depicta are disturbed. The specific metabolic patterns of Mediterranean cephalaspideanssuggest that these patterns can be used as chemotaxonomic markers.We propose the use of a single Thin Layer Chromatography todifferentiate among Mediterranean Haminoea species. (Received 3 February 1997; accepted 22 May 1998)     

Larval development of Lynceus brachyurus (Crustacea, Branchiopoda, Laevicaudata): redescription of unusual crustacean nauplii, with special attention to the molt between last nauplius and first juvenile

The larval development of "conchostracans" has received only scattered attention. Here I present the results of a study on the larval (naupliar) development and the metamorphosis of Lynceus brachyurus, a member of the bivalved branchiopod order the Laevicaudata. Lynceus brachyurus is the only species of the "Conchostraca" in Denmark. The phylogenetic position of the Laevicaudata has traditionally been a source of controversy, and this study does not solve the question completely. This work focuses on features potentially important for phylogeny. The general appearance of the larvae of L. brachyurus has been known for more than a century and a half, and some of its unique features include a large, larval dorsal shield; a huge, plate-like labrum; and a pair of immovable, horn-like antennules. However, many details relating to limb morphology, potentially important for phylogeny, have not been studied previously. Based on size categories, five or six larval stages can be recognized. The larvae approximately double their length and width during development (length: 230-520 microm). Most morphological features stay largely unchanged during development, but the antennal coxal masticatory spines are significant exceptions: they become bifid after one of the first molts. In all larval stages only the antennae and the mandibles actively move. In late naupliar stages the trunk limbs become visible as rows of laterally placed, undeveloped, and still immovable lobes. Swimming is performed by the antennae, whereas the mandibles appear to be involved mainly in feeding, as in other branchiopod larvae. The last naupliar stage undergoes a small metamorphosis to the first juvenile stage, the details of which in part were studied by following the premolt juvenile condition through the cuticle of the last stage nauplius. Among other changes there is a characteristic change in the shape and morphology of the univalved dorsal naupliar shield to a bivalved juvenile carapace. The general morphologies of the antennae and the mandibles are very similar to those of other branchiopod larvae and fall well within the "branchiopod naupliar feeding apparatus" recognized as a branchiopod synapomorphy by Olesen (2003), but some specific features shared with the larvae of other "conchostracans" are also identified. These special "conchostracan" features include: 1) a similar antennular setation; 2) a similar comb-like setulation of the bifid antennal coxal processes; and 3) mandibular palpsetae with setules condensed. In light of recent suggestions concerning branchiopod phylogeny (Cyclestheria as a sister group to the Cladocera), these similarities probably do not support a monophyletic "Conchostraca" but rather are symplesiomorphies of this taxon. A final decision must await a phylogenetic analysis of a more complete set of characters.     

The nauplius stages of the cirripede Tetraclita squamosa rufotincta Pilsbry

The nauplius stages of the cirripede Tetraclita squamosa rufotincta Pilsbry from Elat have been cultured and described. There are the usual six larval stages followed by the cypris but the increase in size during development is small compared with many other species. This small increase compares favourably with other species having very large embryos containing an excess of yolk and which do not take external food during larval development. The setation of the larval appendages is less than in other species and on the antenna and mandible does not increase after stage III. It is suggested that the lack of setation, coupled with a reduced development of the labrum, may be a consequence of the lack of a necessity for this species to feed externally during its planktonic life.     

Scanning Electron Microscopy of Mouth Appendages in Six Species of Barnacles (Crustacea Cirripedia Thoracica)

The morphology and setation of mouth appendages (trophi) are investigated with scanning electron microscopy in the balanomorph barnacles Semibalanus balanoides, Balanus balanusand B. nubilusand in the pedunculate barnacles Pollicipes polymerus, P. cornucopiaand Lepas anatifera.It is difficult to uphold a clearcut distinction between denticles and setae and several types of setae also intergrade with each other. The trophi of Pollicipes polymerusand P. cornucopiahave the most ‘generalized’ morphology. The palps of Pollicipesand L. anatiferaare simpler than those in balanomorphs and carry a single type of seta. In L. anatiferathe mandibles and maxillules are adapted to a semi-predatory feeding habit by carrying large, pointed teeth, but this species lacks the foliate-serrate setae which populate the palps in the other species studied. Compared with Pollicipesand Lepas, the three balanomorphs have palps with a complex setation. The trophi in S. balanoidesdiffer from the two species of Balanusin numerous features of setation and denticulation, notably in having the palps populated by plumodenticulate setae homologous to purely denticulated types in Balanus.It is suggested that SEM studies of barnacle trophi will provide characters of use in estimating phylogeny.     

A key for the identification of the known larval stages of the Mediterranean Brachyura

Summary

Based on the accounts by various authors, an identification key has been constructed for the planktonic zoeae of the 70 species of Mediterranean Brachyura whose larvae are known. The key is based on primary (spines of the carapace, exopodite of the antenna, spines of telson forks) as well as secondary (setation of mouthparts, processes and spines on abdominal somites) morphological characteristics. It allows the identification of the zoeae of 64 species plus those of Liocarcinus genus (six species).     

The reticulated species of Chydorus (Cladocera,Chydoridae): two new species with suggestions of convergence

Of the two small, compact, dusky, reticulated species of Chydorus from Sri Lanka, one arbitrarily has been selected as the taxon to which Daday's name reticulatus henceforward will be attached, the other herein being described as new. A third species from eastern North America, also described herein as new, resembles the two Asian species in certain gross features but otherwise is very distinct. Particularly noteworthy among these differences are the structure of the labrum and of the male postabdomen and copulatory hook. The question is raised whether the gross morphological features that seem to unite the taxa or the features of the labrum and of the male that separate them are the more conservative. No unequivocal answer is provided. Because of their obvious close similarity, the two Asian species are established as a species group, the one from North America being divergent and therefore not included. The possibility of convergence is considered.The North American taxon in north Florida occurs in waterbodies with a pH less than 5 (down to 4.2), conductivity less than 40 µS, and usually with no more than a trace of dissolved color. They are naturally acidic, non-bog lakes. The number of species of chydorids and of total littoral Cladocera in them are considerably greater than found by Fryer in waterbodies of comparable acidity in England. The occurrence also of a diversity of macrophytes and fishes in these lakes indicates that pH per se is not the factor forcing the decreasing diversity associated with changes resulting from acid precipitation.     

Juvenile morphology of the xanthid crab Leptodius exaratus (H. Milne-Edwards, 1834) (Decapoda: Brachyura), with notes on the appearance of sexual dimorphism

Summary

The morphometrical and meristic features of the carapace, cephalic appendages (antenna, antennule), mouthparts (maxillule, maxilla, first-third maxillipeds), sternum, pereiopods, abdomen, and pleopods of juveniles and the onset of morphological sexual dimorphism were described for the xanthid crab Leptodius exaratus (H. Milne Edwards, 1834), based on laboratory-reared and wild adult specimens collected from Tateyama Bay, Japan. First instar juveniles shared some of the features of adults (e.g. gross appearance of the carapace and cheliped propodus proportions), but differed from adults on almost all other morphological parameter examined. Morphological development was still not complete at the ninth instar; extrapolation from the rate of morphological changes between instars 1–9 suggests that L. exaratus requires about 13 ecdyses to transform into adults, including development of reproductive structures. Differences in the number and morphology of pleopods and abdomen width allowed early distinction of the sexes. Thus, males formed gonopods in the first abdominal somite and lost the paired vestigial pleopods in somites 3–5 from the fourth instar; females retained the pleopods in somites 2–5, but these became biramous and had increased setation. The abdomen grew wider in females than in males from the fifth instar. Several morphological features of juveniles have phylogenetic and taxonomic implications: carapace motifs clearly place L. exaratus in the superfamily Xanthoidea, whereas the patterns of setation in the scaphognathite and first maxilliped epipod allow separation of this (xanthid) species from crabs of other Xanthoidean families.     

The first-stage zoeas of Carpilius convexus (Forskal, 1775) and Carpilius maculatus (Linnaeus, 1758) (Crustacea: Decapoda: Brachyura: Xanthoidea: Carpiliidae): an example of heterochrony

The first-stage zoeas of Carpilius convexus (Forskål,1775) and Carpilius maculatus (Linnaeus, 1758) are describedand fully illustrated. Both these Indo-West Pacific speciesexhibited a unique xanthoidean character for the basis of thesecond maxilliped, which possesses five (arranged 1,1,1,2) setaeinstead of the expected four (arranged 1,1,1,1). A comparisonwith the zoeal stages of Carpilius corallinus (Herbst, 1783)as reported by Laughlin et al. (Laughlin et al., 1983) revealedmarked differences including the possession of carapace lateralspines (v. absent in C. convexus only), the subterminal setationof the distal maxillule endopod segment with two subterminalsetae (versus one subterminal seta in Indo-West Pacific species),the terminal setation of the distal maxillule endopod segmentwith four setae (versus three setae in C. convexus only), twolateral spines on the telson (versus three in Indo-West Pacificspecies) and the number of zoeal stages. The first-stage zoeasof the two Indo-West Pacific species appear to have hatchedin a more advanced state of development than those of C. corallinus,and the expression of a number of characters has been accelerated(early onset). In fact, the zoeal stages of both Indo-West Pacificspecies and carpiliid species appear to be abbreviated becausethe first zoeas are considered to be equivalent to the third-stagezoeas of C. corallinus.     

Larval stages of the crab Mithrax tortugae (Brachyura: Mithracidae) with comparisons between all species of Mithrax

The postembryonic larval stages of Mithrax tortugae are described, illustrated and compared with the described zoeae of other Mithrax species. Mithrax tortugae showed morphological features in all the stages of larval development that differed from those observed in other species of Mithrax, especially M. hispidus. In the Zoea I stage, M. tortugae and M. pleuracanthus lacked the minute spine on the dorsal spine observed in M. hispidus; M. tortugae exhibited a terminal spine on the inner lobe of the coxal endite of the maxilla, which was not observed in M. hispidus or M. pleuracanthus. Also, M. tortugae exhibited furcae with spines that are not spinulated, whereas in M. hispidus and M. pleuracanthus these spines are spinulated. In the Zoea II stage, M. tortugae showed a terminal spine on the coxal endite of the maxillula, whereas in M. hispidus and M. pleuracanthus this spine is absent. In the Megalopa stage, we also observed differences in the sternal plate setation between M. tortugae and M. hispidus, where M. tortugae had eight simple setae and M. hispidus showed two simple and four plumodenticulate setae. Partial sequences of the 16S rRNA and COI genes of the parental female were analysed, providing additional evidence for species identification. Together, our analyses of larval morphology and the results of the molecular analyses reinforced recognition of the relationships among M. tortugae, M. hispidus and M. pleuracanthus.     

Functional morphology of mouthparts and digestive system during larval development of the cleaner shrimp Lysmata amboinensis (de Man, 1888)

Mouthpart and alimentary canal development was examined in Lysmata amboinensis larvae using scanning electron microscopy and histology. The gross morphological features of external mouthparts and internal digestive tract structures of larvae at different developmental stages indicate that ingestive and digestive capabilities are well developed from early on. With increasing age of the larvae the mouthpart appendages increased in size, the hepatopancreas in tubular density and the midgut in length. The density of setae and robustness of teeth and spines of individual structures increased. The most pronounced changes from early to late stage larvae involved formation of pores on the paragnaths and labrum, transformation of the mandibular spine‐like teeth to molar cusps, development of the filter press in the proventriculus and of infoldings in the previously straight hindgut. The results suggest that early stage L. amboinensis larvae may benefit from soft, perhaps gelatinous prey, whereas later stages are better equipped to handle larger, muscular or more fibrous foods. J. Morphol. 2011. © 2011 Wiley‐Liss, Inc.     

Seasonal species composition of barnacle larvae (Cirripedia: Thoracica) in Rhode Island waters, 1977-1978

Cirripede larvae can occur year-round in temperate and tropicalwaters, often in significant numbers, yet the species compositionof the ‘Balanus sp.’ component is rardy studied.Weekly plankton samples were analyzed qualitatively for larvalcirripede species and stage over a year (1977–1978) attwo Rhode Island stations. Six species of larvae were foundin Lower Narragansett Bay (30° salinity). Semibalanus balanoidesand Balanus balanus have a single winter brood. S. balanoidesis the predominant winter breeder with a minor release of naupliiin early December and major release in March followed by cypridsin mid-April.B. balanus populations release all larvae in Marchwith cyprids in mid-April. Balanus crenatus is mainly a winterbreeder, but has multiple broods; it does not breed in July-Septemberwhen the water temperature is above 18°C. Balanus venustusis the predominant summer breeder, and larvae were observedfrom May through December (water >8°C). Larvae of Chthamalusfragilis and Balanus eburneus occur in low numbers from May-October.At the Pettaquamscutt River site (12 salinity), Balanus improvisuslarvae predominate and early stage nauplii (I-II) occur in samplesyear round (0–27°C). Two peaks of later stage naupliiand cyprids occur in late spring (May) and early winter (Nov.-Jan.).Continued temperatures bdow 5°C or above 20°C appearto inhibit larval development. Comparison of results with existing literature reveals severalsignificant findings. The bimodal rdease of S. balanoides larvaeis unusual and may be in response to the phytoplankton dynamicsof the year; however, the existence of distinct races of S.balanoides may also be a factor. Larvae of B. venustus predominatein the lower bay during the summer, yet this species is unreportedin past studies. B. improvisus nauplii are more cold tolerantthan previously reported. Comparison of findings with reportedbreeding patterns in Florida indicate significant differencesin temperature responses between northern and southern populations. ¹Contribution No. 188 from EPA Environmental Research Laboratory,South Ferry Rd., Narragansett, RI 02882