The Chromodorididae (Opisthobranchia: Mollusca) of the Indo-West Pacific: Noumea purpurea and Chromodoris decora colour groups

Nine species of chromodorid nudibranchs are described including a new species of Durvilledoris and a new species of Noumea. The Noumea purpurea colour group consists of five species with a pink, purple or reddish-purple background and a median white line, sometimes broken. The Chromodoris decora colour group also has white longitudinal lines with a pinkish background colour and usually an orange border. A non-chromodorid juvenile, probably a species of Hexabranchus , is compared with the species of the C. decora colour group.     

The Chromodorididae (Opisthobranchia: Mollusca) of the Indo-West Pacific: further species of Glossodoris, Thorunna and the Chromodoris aureomarginata colour group

Nineteen species of chromodorid nudibranchs from the Indo-West Pacific are described including 12 new to science. There are four new species of Glossodoris and one new species of Thorunna . The anatomy of two other species of Thorunna, T. daniellae and T. florens , is described and the problematical 'Hypselodoris' kulonba Burn, 1966 is shown to belong to the genus Digidentis. New species of chromodorids which are white with a yellow or red margin. or markings. from the genera Durvilledoris. Noumea and Cadlina are also described. Notes on colour variation in Glossodoris pallida and Chromodoris splendida are included.     

The Chromodorididae (Opisthobranchia: Mollusca) of the Indo-West Pacific: Chromodoris aureomarginata, C. verrieri and C. fide lis colour groups

Nineteen species of chromodorid nudibranchs are described including three new species of Chromodoris , two of Glossodoris and one each of Noumea, Cadlina and Thorunna from the Indo-West Pacific. The colour of all species is white with either a single yellow or gold band at the edge of the mantle or a double band of red and yellow, or orange and red or purple. All previously described species with similar colour patterns, from both the Indo-West Pacific and other regions are discussed.     

The Chromodorididae (Opisthobranchia: Mollusca) of the Indo-West Pacific: the genus Ceratosoma J. E. Gray

The genus Ceratosoma (Nudibranchia: Chromodorididae) is reviewed and seven species, including two new species, are recognized. Species of the genus are known only from the Indo-West Pacific and it is suggested that the external body shape is the most useful character in distinguishing species.     

Opisthobranchiate Mollusca from Ghana: Chromodorididae

Nine species of the opisthobranch family Chromodorididae are described from Ghana: Cadlina evelinae Marcus; C. rumia Marcus; C. dubia sp. nov.; Chromodoris ghanensis (Edmunds); C. kpone sp. nov.; C. luleorosea (Rapp); Hypselodoris tana sp. nov.; H. bilineata (Pruvot-Fol) and Mexichromis tricolor (Cantraine). The small number of species found belonging to this warm-water family is attributed to the cold water of the Benguela current.     

The Chromodorididae (Opisthobranchia: Mollusca) of the Indo-West Pacific: Noumea flava colour group

Six species of the genus Noumea are described including four new species. All species are yellow, and four have orange markings as well; one has a white mantle border and the last has red gills and rhinophores. Noumea flava has a widespread distribution in the Indo-West Pacific and two other species are known from the tropical Pacific. The other three species, including a yellow colour form of Noumea haliclona (Burn, 1957), are reported from southern Australia. Aspects of the reproductive system of some species of Noumea strongly indicate that the genus Verconia Pruvot-Fol, 1931 is closely related to that genus.     

The Chromodorididae (Opisthobranchia: Mollusca) of the Indo-West Pacific: Chromodoris epicuria, C. aureopurpurea, C. annulata, C. coi and Risbecia tryoni colour groups

Twenty-five species of chromodorid nudibranchs from the Indo-West Pacific are described including nine which are new to science. There are six new species of Chromodoris , two of Nounea and a new species which forms the type of a new monotypic genus. Of the other species discussed in this paper, ten are species of Chromodoris , five Risbecia and one Digidentis. Species with the following colour patterns are discussed: white with orange or yellow spots; white with yellow (or orange) and purple spots; white with purple spots or patches and mantle border; white with a sinuous submarginal brown band. A full review of the relevant literature is included as is a discussion on Indian Ocean and West Pacific species pairs.     

The Chromodorididae (Opisthobranchia: Mollusca) of the Indo-West Pacific: the genus Glossodoris Ehrenbergh (= Casella, H. & A. Adams)

The genus Glossodoris Ehrenberg is reviewed and 28 species are placed in the genus. Seventeen species are fully described and of these, seven are named as new species. Four others are considered new to science, but insufficient material is available to name them. Published information on a further 11 species is included as is a discussion on phylogenetic trends within the genus.     

The Chromodorididae (Opisthobranchia: Mollusca) of the Indo-West Pacific: a review of the genera

The full anatomy of over fifty species of the Chromodorididae is described including the type species of all but one of the nominal genera. The holotype material of the type species of Ehrenberg's problematic genera Glossodoris, Actinodoris and Pterodoris is described. Sixteen genera are recognized within the family Chromodorididae including four new monotypic genera. Glossodoris is shown to be a senior synonym of Casella , not Chromodoris , and Ceratosoma and Verconia are shown to be members of the Chromodorididae. The separation of Cadlina, Tyrinna and Cadinella into a separate sub-family Cadlininae is considered unnecessary.
A large suite of characters including features of the external shape, mantle glands, gills, foregut, jaw-plates, radula and reproductive system are used to define the genera. A tentative phylogeny of the family is proposed.     

Phylogeny of Hypselodoris (Nudibranchia: Chromodorididae) with a review of the monophyletic clade of Indo-Pacific species, including descriptions of twelve new species

This work provides an account of the systematics and phylogeny of Hypselodoris . Aspects of the morphology of 42 species are described and the systematic status of an additional 11 species is discussed. Twelve new species are described: Hypselodoris alboterminata, H. bertschi, H. bollandi, H. fucata, H. iacula, H. insulana, H. krakatoa, H. paulinae, H. reidi, H. rudmani, H. violabranchia and H. zephyra. A phylogenetic analysis supports the monophyly of Hypselodoris and Risbecia . Two distinct clades of Hypselodoris are present. One contains species from the Atlantic and eastern Pacific while the other contains species limited to the Indo-Pacific tropics and adjacent temperate regions. Species from the Atlantic and eastern Pacific are bluish in body colour and have a plesiomorphically large receptaculum seminis while Indo-Pacific taxa are variably coloured and all have a minute receptaculum seminis. The distribution and size of mantle glands provides a wealth of morphological characters. With few exceptions, mantle glands vary in closely related species and are important for distinguishing members of smaller clades. Mantle gland distribution is therefore useful in identifying preserved material that is difficult to identify to species in the absence of the pigment of living specimens. Similar colour patterns found in sympatric species of Hypselodoris appear to be a result of both common descent and convergence between less closely related lineages. Biogeographic distributions of sister taxa provide several examples of vicariance. Examination of these cases shows that no single vicariant pattern is present, but vicariance appears to occur at the margins of the Indo-Pacific rather than centrally. Some vicariance occurs even within archipelagos such as the Hawaiian Islands. These cases largely refute the generality of the hypothesis of Springer (1982), that Pacific Plate and Australasian Plate endemic sister taxa should predominate.     

Sperm ultrastructure in the nudibranch genus Halgerda with reference to other Discodorididae and to Chromodorididae (Mollusca: Opisthobranchia)

Comparative sperm ultrastructure within the molluscan nudibranch genus Halgerda (Discodorididae) was examined for the first time using transmission electron microscopy (TEM), based on 17 of the 35 known species. In addition, observations on two other discodorids are made to facilitate outgroup comparison with Halgerda, including one species of Discodoris (D. boholiensis) and Asteronotus cespitosus (currently accepted as the closest sister taxon to Halgerda). Comparison was also made with some genera of the Chromodorididae in view of sperm similarities. Spermatozoa of all species examined were of the complex, helical, elongate ( approximately 300-400 micro m) type characteristic of most heterobranch gastropods. These cells exhibit the following discrete regions (in anteroposterior sequence) : an acrosomal complex (composed of a rounded, membrane-bound vesicle and a column-like pedestal); a solid, helical nucleus; an elongate, helical midpiece (composed of an axoneme and associated nine coarse fibers, an enveloping mitochondrial derivative of matrix, and paracrystalline materials and glycogen helix); an annular complex; and a short glycogen piece. Of these regions, the midpiece is by far the longest, occupying over 90% of the total sperm length. Comparison with other members of the radula-bearing cryptobranch dorids reveals several sperm similarities to other genera in the clade, particularly those of other Discodorididae and also with the Chromodorididae. Comparison with previously studied genera reveals noteworthy sperm differences within the Discodorididae. The most notable differences are the internal structure of the acrosomal pedestal (long and homogeneous in Halgerda, Discodoris; short and homogeneous in Asteronotus; long and finely striated in Rostanga; oblong with angular electron-lucent striations in Jorunna) and the internal structure of the glycogen piece. The pronounced helical keels of most Halgerda and Discodoris nuclei contrast with the weakly helical nucleus of Asteronotus. Sperm features alone do not provide a means of defining the genus Halgerda or the family Discodorididae nor do they support the monophyletic status of the caryophyllidia-bearing dorids. Important sperm characters such as the acrosome, nucleus, and midpiece can often still be determined from specimens that have been initially fixed in formalin, then stored in ethanol for extended periods of time (i.e., museum material). Of all sperm features, the mitochondrial derivative of the midpiece is the most resistant to long-term fixation : the survival of acrosomal, nuclear, and axonemal components is variable, presumably a factor of prefixation autolysis, varied primary fixation times and temperatures, formalin quality, and duration of alcohol storage.     

Comparative immunohistochemistry of the cephalic sensory organs in Opisthobranchia (Mollusca,Gastropoda)

The structure and function of the central nervous systems of opisthobranch gastropods have been studied extensively. However, the organisation and function of the peripheral nervous system are poorly understood. The cephalic sensory organs (CSOs) are known to be chemosensory structures in the head region of opisthobranchs. In the present study, we used immunohistochemical methods and confocal laserscanning microscopy to comparatively examine the CSOs of different opisthobranchs, namely Acteon tornatilis, Aplysia punctata, Archidoris pseudoargus and Haminoea hydatis. We wanted to characterise sensory epithelia in order to infer the function of sensory structures and the organs they constitute. Immunoreactivity against the three antigens tyrosine hydroxylase, FMRFamide and serotonin was very similar in the CSOs of all investigated species. Tyrosine hydroxylase-like immunoreactivity was detected primarily in subepidermal sensory cell bodies, which were much more abundant in the anteriorly situated CSOs. This observation indicates that these cells and the respective organs may be involved in contact chemoreception and mechanoreception. The dominant features of FMRFamide-like immunoreactivity, especially in the posterior CSOs, were tightly knotted fibres which reveal glomerulus-like structures. This suggests an olfactory role for these organs. Serotonin-like immunoreactivity was detected in an extensive network of efferent fibres, but was not found within any peripheral cell bodies. Serotonin-like immunoreactivity was found in the same glomerulus-like structures as FMRFamide-like immunoreactivity, indicating a function of serotonin in the efferent control of olfactory inputs. Besides this functional implication, this study could also add some knowledge on the doubtful homology of the CSOs in opisthobranch gastropods.     

Ultrastructural and histochemical study of the salivary glands of Aplysia depilans (Mollusca, Opisthobranchia)

The digestive system of the sea hare, Aplysia depilans , includes a pair of ribbon-shaped salivary glands. A central duct and a large blood vessel run close to each other along the length of these glands and both are surrounded by a layer of muscle cells. Three cell types form the glandular epithelium: granular cells, vacuolated cells and mucocytes. The granular cells possess cilia and spherical secretion granules, located primarily in the apical region. The granules of immature cells have a low electron density and are mainly formed by neutral polysaccharides with small amounts of proteins. The granules of mature cells are larger, have a high electron density and are mainly formed by proteins with lower amounts of neutral polysaccharides. Transition stages between immature and mature granular cells are observed. The vacuolated cells are large and frequently pyramidal in shape, but after the application of histochemical techniques almost all vacuoles remain uncoloured. The numerous vacuoles contain flocculent material in a clear background and the mitochondria possess large crystalline structures in the matrix. A pyramidal shape is also typical of the mucocytes, which are filled with vesicles containing granular masses surrounded by a network of secretion material. These large cells are strongly stained by Alcian blue, revealing the presence of acidic mucopolysaccharides. This is the first ultrastructural study of the salivary glands in opisthobranch gastropods.     

A comparative ultrastructural investigation of the cephalic sensory organs in Opisthobranchia (Mollusca, Gastropoda)

Cephalic sensory organs (CSOs) are specialised structures in the head region of adult Opisthobranchia involved in perception of different stimuli. The gross morphology of these organs differs considerably among taxa. The current study aims at describing the cellular morphology of the CSOs in order to reveal cellular patterns, especially of sensory epithelia, common for opisthobranchs. Transmission electron microscopy was used to characterise the fine structure of the organs and to compare the CSOs of four different opisthobranch species. The cellular composition of the sensory system is conserved among taxa. The epidermal cells in sensory regions are always columnar and ciliated cells are frequently apparent. The sensory cells are primary receptors arranged in subepidermal cell clusters. They extend dendrites which penetrate the epithelium and reach the surface. Some of the dendrites bear cilia, whereas others only build a small protuberance. Processing of sensory information takes place in the peripheral glomeruli of all species. Moreover, few taxa possess additional peripheral ganglia at the base of their CSOs. The results of the present study might support other investigations indicating that the posterior CSOs are primarily involved in distance chemoreception, whereas the anterior CSOs might be used for contact chemoreception and mechanoreception.     

Evolution of the Sacoglossa (Mollusca, Opisthobranchia) and the ecological associations with their food plants

Evolution in the opisthobranch order Sacoglossa has been closely linked to their specialized suctorial herbivorous habits. All shelled Sacoglossa (about 20% of the species) feed on one algal genus, Caulerpa. The non-shelled Sacoglossa have 'radiated' to other diets, mainly siphonalean or septate green algae (Class Ulvophyceae). Comparing the phylogeny of sacoglossan genera with the phylogeny of the Ulvophyceae indicates that co-speciation may have taken place at the basal node of the Sacoglossa, and that host switching has taken place several times in the two non-shelled clades. It is suggested that the most important evolutionary process has been speciation by 'resource-tracking'; the resource tracked is most probably cell wall composition of the algal prey. The fossil record of extant sacoglossan genera dates back to the Eocene and, based on the fossil record of siphonalean green algae, the Sacoglossa most likely appeared in the Cretaceous. It is hypothesized that the ancestral sacoglossan was epifaunal, suctorial and herbivorous, and the 'ancestral' food plant was not Caulerpa, but filamentous, calcified, now extinct, Udoteaceae.     

Systematic position of the pelagic Thecosomata and Gymnosomata within Opisthobranchia (Mollusca, Gastropoda) – revival of the Pteropoda

The complete 18S (SSU) rRNA as well partial 28S (LSU) rRNA and partial mitochondrial COI sequences have been used to reconstruct the phylogenetic relationships within Opisthobranchia with special focus on the pelagic orders Thecosomata and Gymnosomata. Maximum parsimony, maximum likelihood, distance as well as Bayesian analysis of a combined dataset of the three genes reveals that Thecosomata and Gymnosomata are sister groups and together are closely related to Anaspidea. Possible sister taxon to Thecosomata, Gymnosomata and Anaspidea is Cephalaspidea s. str . Analysis of a taxon-extended dataset of partial 28S sequences supported a basal position of Limacina within Euthecosomata. Within Cavolinidae, Creseis is basal to the other taxa. Other phylogenetic implications from the present results are also discussed. Investigation of the morphology and histology of Thecosomata and Gymnosomata as well as several other opisthobranch taxa helped to identify autapomorphies for Thecosomata and Gymnosomata as well as apomorphies for the clades including these taxa.     

On the phylogeny of the Arminidae (Gastropoda, Opisthobranchia, Nudibranchia) with considerations of biogeography

In the past, the different genera of the family Arminidae have been diagnosed (mostly) on the basis of plesiomorphic characters and therefore their monophyly is questionable. The Arminidae are characterized by the autapomorphies 'possession of marginal sacs' and 'rhinophores with longitudinal lamellae'. The genus Heterodoris is excluded from the family. Two possible hypotheses about the phylogenetic relationships within the Arminidae are presented.
The Arminidae probably originated in the western Pacific, near Japan and have distributed in a westerly direction. The most primitive species ( Dermatobranchus ) are restricted mainly to the western Pacific region while the more derived species ( Armina ) have a world-wide distribution.     

Systematics and phylogeny of the Aglajidae (Opisthobranchia: Mollusca)

The generic limits within the Aglajidae are reviewed and modified, based on the study of 21 species. Navanax Pilsbry, 1895 is reinstated as a genus distinct from Aglaja. The generic placement of several species is altered from previous works. An evaluation of morphological characters suggests that each of six genera has distinctive external morphology. The systematic value of twelve characters is critically reviewed. The evolutionary relationships of the Aglajidae and its genera to other philinacean familes are discussed.