Annotated type catalogue of the Bothriembryontidae and Odontostomidae (Mollusca, Gastropoda, Orthalicoidea) in the Natural History Museum, London

The type status is described for specimens of 84 taxa classified within the families Bothriembryontidae and Odontostomidae (superfamily Orthalicoidea) and kept in the Natural History Museum, London. Lectotypes are designated for Bulimus (Liparus) brazieri Angas, 1871; Bulimus broderipii Sowerby I, 1832; Bulimus fuligineus Pfeiffer, 1853; Helix guarani d'Orbigny, 1835; Bulimus (Tomigerus) ramagei E.A. Smith, 1890; Helix rhodinostoma d'Orbigny, 1835; Bulimus (Bulimulus) ridleyi E.A. Smith, 1890. The type status of the following taxa is changed to lectotype in accordance with Art. 74.6 ICZN: Placostylus (Euplacostylus) cylindricus Fulton, 1907; Bulimus pyrostomus Pfeiffer, 1860; Bulimus turneri Pfeiffer, 1860. The following taxon is synonymised: Bulimus oblitus Reeve, 1848 = Bahiensis neglectus (Pfeiffer, 1847).     

Annotated type catalogue of the Orthalicoidea (Mollusca, Gastropoda) in the Royal Belgian Institute of Sciences, Brussels, with descriptions of two new species

The type status is described of 57 taxa from the superfamily Orthalicoidea in the collection of the Brussels museum. Two new species are described: Stenostylus perturbatussp. n., and Suniellus adrianisp. n. New lectotypes are designated for Bulimulus (Naesiotus) amastroides Ancey, 1887; Bulimulus blanfordianus Ancey, 1903; Bulimulus montivagus chacoensis Ancey, 1897; Bulimus coloratus Nyst, 1845; Plecochilus dalmasi Dautzenberg, 1900; Placostylus porphyrostomus elata Dautzenberg, 1923; Bulimulus ephippium Ancey, 1904; Bulimus fulminans Nyst, 1843; Bulimus funckii Nyst, 1843; Orphnus thompsoni lutea Cousin, 1887; Bulimus melanocheilus Nyst, 1845; Orphnus thompsoni nigricans Cousin, 1887; Orphnus thompsoni olivacea Cousin, 1887; Bulimulus pollonerae Ancey, 1897; Orphnus thompsoni zebra Cousin, 1887. New combinations are: Bostryx borellii (Ancey, 1897); Bostryx carandaitiensis (Preston, 1907); Protoglyptus mazei (Crosse, 1874); Kuschelenia (Vermiculatus) sanborni (Haas, 1947). New synonymies are established for the following nominal taxa: Orphnus thompsoni var. lutea Cousin, 1887 = Kara thompsonii (Pfeiffer, 1845); Orphnus thompsoni var. nigricans Cousin, 1887 = Kara thompsonii (Pfeiffer, 1845); Thaumastus nystianus var. nigricans Cousin, 1887 = Drymaeus (Drymaeus) nystianus (Pfeiffer, 1853); Orphnus thompsoni var. olivacea Cousin, 1887 = Kara thompsonii (Pfeiffer, 1845); Orphnus thompsoni var. zebra Cousin, 1887 = Kara thompsonii (Pfeiffer, 1845).     

Annotated type catalogue of the Amphibulimidae (Mollusca, Gastropoda, Orthalicoidea) in the Natural History Museum, London

The type status is described of 39 taxa classified within the family Amphibulimidae (superfamily Orthalicoidea) and kept in the London museum. One taxon, Bulimus elaeodes Pfeiffer, 1853, is removed to the Strophocheilidae. Lectotypes are designated for Bulimus adoptus Reeve, 1849; Bulimus (Eurytus) eros Angas, 1878; Helix onca d'Orbigny, 1835; Amphibulima pardalina Guppy, 1868. The type status of the following taxon is changed to lectotype in accordance with Art. 74.6 ICZN: Strophocheilus (Dryptus) jubeus Fulton, 1908.As general introduction to this and following papers on Orthalicoid types in the Natural History Museum, a brief history of the London collection is given and several examples of handwriting from different authors are presented.     

Annotated type catalogue of the Megaspiridae,Orthalicidae, and Simpulopsidae (Mollusca,Gastropoda, Orthalicoidea) in the Natural History Museum,London

The type status is described for 65 taxa of the Orthalicoidea, classified within the families Megaspiridae (14), Orthalicidae (30), and Simpulopsidae (20); one taxon is considered a nomen inquirendum. Lectotypes are designated for the following taxa: Helix brephoides d’Orbigny, 1835; Simpulopsis cumingi Pfeiffer, 1861; Bulimulus (Protoglyptus) dejectus Fulton, 1907; Bulimus iris Pfeiffer, 1853. The type status of Bulimus salteri Sowerby III, 1890, and Strophocheilus (Eurytus) subirroratus da Costa, 1898 is now changed to lectotype according Art. 74.6 ICZN. The taxa Bulimus loxostomus Pfeiffer, 1853, Bulimus marmatensis Pfeiffer, 1855, Bulimus meobambensis Pfeiffer, 1855, and Orthalicus powissianus var. niveus Preston 1909 are now figured for the first time. The following taxa are now considered junior subjective synonyms: Bulimus marmatensis Pfeiffer, 1855 = Helix (Cochlogena) citrinovitrea Moricand, 1836; Vermiculatus Breure, 1978 = Bocourtia Rochebrune, 1882. New combinations are: Kuschelenia (Bocourtia) Rochebrune, 1882; Kuschelenia (Bocourtia) aequatoria (Pfeiffer, 1853); Kuschelenia (Bocourtia) anthisanensis (Pfeiffer, 1853); Kuschelenia (Bocourtia) aquila (Reeve, 1848); Kuschelenia (Bocourtia) badia (Sowerby I, 1835); Kuschelenia (Bocourtia) bicolor (Sowerby I, 1835); Kuschelenia (Bocourtia) caliginosa (Reeve, 1849); Kuschelenia (Bocourtia) coagulata (Reeve, 1849); Kuschelenia (Bocourtia) cotopaxiensis (Pfeiffer, 1853); Kuschelenia (Bocourtia) filaris (Pfeiffer, 1853); Kara indentata (da Costa, 1901); Clathrorthalicus magnificus (Pfeiffer, 1848); Simpulopsis (Eudioptus) marmartensis (Pfeiffer, 1855); Kuschelenia (Bocourtia) nucina (Reeve, 1850); Kuschelenia (Bocourtia) ochracea (Morelet, 1863); Kuschelenia (Bocourtia) peaki (Breure, 1978); Kuschelenia (Bocourtia) petiti (Pfeiffer, 1846); Clathrorthalicus phoebus (Pfeiffer, 1863); Kuschelenia (Bocourtia) polymorpha (d’Orbigny, 1835); Scholvienia porphyria (Pfeiffer, 1847); Kuschelenia (Bocourtia) purpurata (Reeve, 1849); Kuschelenia (Bocourtia) quechuarum Crawford, 1939; Quechua salteri (Sowerby III, 1890); Kuschelenia (Bocourtia) subfasciata Pfeiffer, 1853; Clathrorthalicus victor (Pfeiffer, 1854). In an addedum a lectotype is being designated for Bulimulus (Drymaeus) interruptus var. pallidus Preston, 1909. An index is included to all taxa mentioned in this paper and the preceding ones in this series (Breure and Ablett 2011, 2012, 2014).     

Annotated catalogue of the types of Indo-Burmese non-marine Gastropoda deposited in the Faculty of Tropical Medicine,Mahidol University,Thailand

ABSTRACT

Four decades after its publication, Rolf A.M. Brandt’s 1974 monograph on the non-marine molluscs of Thailand remains the main authority on freshwater and estuarine species for Southeast Asia and includes up to 165 new species of snails and bivalves described by Brandt and colleagues in the same book and preceding publications. All the holotypes are lodged at the Forschungsinstitut und Naturmuseum Senckenberg in Germany, and are largely inaccessible to Thai and other Southeast Asian researchers, who rely heavily on the Brandt collection as a key reference. Paratypes were, however, donated to various other collections, including some in Thailand. We present the first catalogue of 45 paratypes of gastropods of the Brandt collection, described from Thailand, Laos and Cambodia, which are lodged at the Faculty of Tropical Medicine, Mahidol University, Thailand.     

The anatomy of Addisonia (Mollusca,Gastropoda)

Summary The organization of Addisonia lateralis (Requien, 1848) and A. brophyi McLean, 1985 is described. Addisonia species have a thin, asymmetrical, cup-like shell and a very simple shell muscle. Eyes, oral lappets and epipodial tentacles are absent and the right cephalic tentacle is also used as a copulatory organ. Most characteristic is the enormously developed gill which is enlarged into the right subpallial cavity. It is composed of about 30 leaflets with skeletal rods and its epithelia are uniquely arranged. The heart is large and the single auricle is situated anteriorly left. There are two kidneys: the left is small, while the right forms large coelomic cavities and has no connection with the pericardium or the hermaphroditic genital system. Testis and ovary are separate: both have a simple duct proper (vas deferens, oviduct). They are connected to the copulatory organ by an open seminal groove; a small receptaculum is present. The mouth opening is typically triangular, with no jaws or subradular sense organs. Addisonia possesses tuft-like salivary glands, a radula diverticulum and distinct, tubular oesophageal glands. The oesophagus itself is simple. The radula and the posterior alimentary tract are unique; the stomach is completely reduced and the intestine forms a pseudostomach. The streptoneurous, hyoathroid nervous system has pedal cords with three commissures. The visceral loop is also cord-like. A single (left) osphradium is present and the small statocysts have several statocones.The peculiarities and unique combination of primitive and advanced characters in Addisonia reflect a highly enigmatic organization among the Archaeogastropoda. Possible relationships to other archaeogastropod groups are discussed.     

Dart formation in Helix aspersa (Mollusca,Gastropoda)

Summary Dart formation in Helix aspersa has been investigated by SEM of isolated darts at progressive stages in their development, and by histology of dart sacs at the same times. Dart formation begins at the tip of a tubercle where a small group of epithelial cells secrete an organic material filling a small CaCO₃ cone that is the first mineralized part of the shaft. Subsequent secretory activity by an increasing area of the tubercle epithelium results in an increase in the diameter and anterior lengthening of the shaft. Continued secretion by the tubercle and dart sac epithelium produces the flare and finally the corona. A pattern of deposition is also evident in the fine structure of the mineral. In the shaft and vanes there is an inner layer of spherulitic prismatic structure which is covered by a layer of irregular patches of simple prismatic structure. The outermost layer of the shaft and vanes has a continuous simple prismatic structure. Two layers are present in the flare, an inner granular amorphous layer and an outer spherulitic prismatic layer. The corona consists of a single rarefied prismatic layer. A mechanism of dart formation is suggested that involves two types of organic matrix, calcifying and non-calcifying. Measurements of the calcium content of darts, dart sacs, and collars indicate that the hemolymph is the probable source of calcium for the dart.     

Basal chromodorid sperm ultrastructure (Nudibranchia, Gastropoda, Mollusca)

The relationship between three genera considered basal in the Chromodorididae (Cadlina, Tyrinna, Cadlinella) has not yet been resolved by traditional morphological means. Here we examined the sperm ultrastructure of Tyrinna nobilis, Tyrinna evelinae, Cadlina flavomaculata and Cadlina cf. nigrobranchiata, with the expectation of finding phylogenetically informative characters. No Tyrinna or Cadlina species showed sperm similarities to Cadlinella. Both Cadlina species and Tyrinna nobilis (but not T. evelinae) exhibited coarse striations in the acrosomal pedestal. The putative fibers that occurred between the coarse striations of the pedestal are condensed into a layer in Cadlina and Tyrinna, but not in other species that also have coarse striations (Gymnodoris), and may constitute evidence for a close relationship. Tyrinna evelinae possessed fine acrosomal striations, which was shared with other Chromodorididae, Actinocyclidae and the cryptobranchs Rostanga and Aphelodoris. We also examined the sperm ultrastructure of ‘Chromodoris’ ambiguus, an animal which has shown molecular affinities to species of Cadlina, and not Chromodoris. The sperm of ‘C.’ ambiguus did not exhibit the typical Cadlina characteristics, but also showed important differences to other investigated Chromodoris species.     

An Ultrastructural Study of Basommatophoran Spermatozoa (Mollusca, Gastropoda)

Spermatozoa of the basommatophoran pulmonate families Ellobiidae ( Ophicardelus ornatus Ferussac), Amphibolidae ( Salinator fragilis Lamarck, Salinator solida von Martens), Siphonariidae ( Siphonaria funiculata Reeve) and Lymnaeidae ( Lymnaea lessoni (Deshayes)) were studied using transmission electron microscopy. Spermatozoa of all species, like most euthyneuran spermatozoa, possess ( 1 ) an acrosome composed of an apical vesicle and acrosomal pedestal (many differences between families), (2) a helically keeled, posteriorly invaginated nucleus, ( 3 ) a midpiece composed of paracrystalline and matrix materials and a variable number of incorporated glycogen-filled helices (one in Salinator and Siphonaria , two or three in Lymnaea , three in Ophicardelus ) and (4) an axoneme associated with coarse fibres (periodically banded in neck region) and rows of intra-axonemal granules. The wide diversity of spermatozoon structure in the species studied, in particular the midpiece structure of Siphonaria (which resembles closely that of certain opisthobranchs) indicates that the Basommatophora may not represent a valid taxonomic unit. A comparison of basommatophoran sperm with other euthyneurans and with euspermatozoa of prosobranchs is given.     

Annotated type catalogue of land snails collected from Taiwan (Formosa) in the Natural History Museum,London

The present catalogue lists the type specimens of land snail species, collected from Taiwan and deposited in the Natural History Museum, London. Thirty-seven nominal species described by Pfeiffer, Adams, Nevill, Moellendorff, Godwin-Austen and Gude were traced. I present here information on type status, collection data obtained from the registers and labels of each collection, and annotations on the current taxonomic affiliation. Lectotypes of 28 nominal (sub)species were newly designated. One holotype was fixed originally and two holotypes newly fixed by monotypy. Syntypes of two species and paralectotypes of three species were also discovered in the Museum. No specimen of the species Pupina adamsi Sowerby, 1878, which was supposed to be deposited in the NHM, was found. Pictures of the name-bearing types are provided for further research on biodiversity of the island.     

Molecular phylogenetics of Caenogastropoda (Gastropoda: Mollusca)

Caenogastropoda is the dominant group of marine gastropods in terms of species numbers, diversity of habit and habitat and ecological importance. This paper reports the first comprehensive multi-gene phylogenetic study of the group. Data were collected from up to six genes comprising parts of 18S rRNA, 28S rRNA (five segments), 12S rRNA, cytochrome c oxidase subunit I, histone H3 and elongation factor 1alpha. The alignment has a combined length of 3995 base positions for 36 taxa, comprising 29 Caenogastropoda representing all of its major lineages and seven outgroups. Maximum parsimony, maximum likelihood and Bayesian analyses were conducted. The results generally support monophyly of Caenogastropoda and Hypsogastropoda (Caenogastropoda excepting Architaenioglossa, Cerithioidea and Campanilioidea). Within Hypsogastropoda, maximum likelihood and Bayesian analyses identified a near basal clade of nine or 10 families lacking an anterior inhalant siphon, and Cerithiopsidae s.l. (representing Triphoroidea), where the siphon is probably derived independently from other Hypsogastropoda. The asiphonate family Eatoniellidae was usually included in the clade but was removed in one Bayesian analysis. Of the two other studied families lacking a siphon, the limpet-shaped Calyptraeidae was associated with this group in some analyses, but the tent-shaped Xenophoridae was generally associated with the siphonate Strombidae. The other studied hypsogastropods with an anterior inhalant siphon include nine families, six of which are Neogastropoda, the only traditional caenogastropod group above the superfamily-level with strong morphological support. The hypotheses that Neogastropoda are monophyletic and that the group occupies a derived position within Hypsogastropoda are both contradicted, but weakly, by the molecular analyses. Despite the addition of large amounts of new molecular data, many caenogastropod lineages remain poorly resolved or unresolved in the present analyses, possibly due to a rapid radiation of the Hypsogastropoda following the Permian-Triassic extinction during the early Mesozoic.     

Crossing type variability associated with cytoplasmic incompatibility in Australian populations of the mosquito Culex quinquefasciatus Say

An analysis of cytoplasmic crossing type variation in Australian populations of Culex quinquefasciatus, a member of the Culex pipiens complex of mosquitoes, revealed high levels of variability causing partial incompatibility between natural populations. Segregating crossing types were commonly found together within sampled sites. No correlation was evident between similarity of crossing type and environmental parameters of the sites, nor distance between sites. The nature of the observed variation did not support the hypothesis of paternally expressed nuclear 'restorer' genes. Such high levels of crossing type variation would be likely to impede attempts to control populations of the Culex pipiens complex using cytoplasmic incompatibility.     

Potential key characters in Opisthobranchia (Gastropoda, Mollusca) enhancing adaptive radiation

Five potential key characters which might have enhanced species radiation in the Opisthobranchia (Gastropoda) are discussed. These are: 3–4 cuticular plates in the gizzard of Cephalaspidea s.str., kleptoplasty in Sacoglossa, kleptocnides in Aeolidoidea, a symbiotic relationship with unicellular algae in Phyllodesmium Ehrenberg, 1831, and mantle dermal formations in Chromodorididae. Interpretation of the characters as key innovations is based on phylogeny and/or comparison of species numbers in subgroups. Possible adaptive zones are discussed, and alternative interpretations indicated.     

Global diversity of gastropods (Gastropoda; Mollusca) in freshwater

The world’s gastropod fauna from continental waters comprises ∼4,000 valid described species and a minimum of 33–38 independent lineages of Recent Neritimorpha, Caenogastropoda and Heterobranchia (including the Pulmonata). The caenogastropod component dominates in terms of species richness and diversity of morphology, physiology, life and reproductive modes and has produced several highly speciose endemic radiations. Ancient oligotrophic lakes (e.g., Baikal, Ohrid, Tanganyika) are key hotspots of gastropod diversity; also noteworthy are a number of lower river basins (e.g., Congo, Mekong, Mobile Bay). But unlike many other invertebrates, small streams, springs and groundwater systems have produced the most speciose associations of freshwater gastropods. Despite their ecological importance in many aquatic ecosystems, understanding of even their systematics is discouragingly incomplete. The world’s freshwater gastropod fauna faces unprecedented threats from habitat loss and degradation and introduced fishes and other pests. Unsustainable use of ground water, landscape modification and stock damage are destroying many streams and springs in rural/pastoral areas, and pose the most significant threats to the large diversity of narrow range endemics in springs and ground water. Despite comprising only ∼5% of the world’s gastropod fauna, freshwater gastropods account for ∼20% of recorded mollusc extinctions. However, the status of the great majority of taxa is unknown, a situation that is exacerbated by a lack of experts and critical baseline data relating to distribution, abundance, basic life history, physiology, morphology and diet. Thus, the already considerable magnitude of extinction and high levels of threat indicated by the IUCN Red List of Threatened Species is certainly a significant underestimate. Guest editors: E. V. Balian, C. Lévêque, H. Segers and K. Martens Freshwater Animal Diversity Assessment     

Experimental and comparative morphology of radula renewal in pulmonates (Mollusca,Gastropoda)

Summary The continuous renewal of the pulmonate radula and the histology and regeneration of its concomitant epithelia were studied by light and electron microscopy, autoradiography and electron microprobe analysis. The two species investigated show histological differences and the results were compared with those of a preceding study on a prosobranch radula. The radula is an intricate cuticular structure of the foregut. Only the fully grown part, which is active during feeding, lies in the buccal cavity while it is constantly renewed by the coordinated cooperation of specialized cells forming the radular sheath. The end of the sheath is occupied by cells which produce the organic matrix of the radula. In taeniogloss prosobranchs, seven multicellular cushions of small odontoblasts lie at the end of the sheath and produce the seven teeth of each cross-row. In pulmonates, the multidenticular radula is generated by numerous groups of a few voluminuous cells. Despite these histological differences, prosobranchs and pulmonates generate the radula matrix by microvilli, cytoplasmatic protrusions and apocrine secretions. The epithelia of the radular sheath contribute to the transport, tanning and mineralization of the radula. The concomitant epithelia are replaced in limited proliferation zones at the end of the radular sheath and their cells migrate anteriorly to the buccal cavity. The ultrastructure of the sheath cells and the alterations which they undergo in connection with their functions are discussed. The proliferation zone of the superior epithelium is strictly confined and the cells move together with the radula forward. In prosobranchs, the cells of the superior epithelium begin to degenerate in the middle of the radular sheath and the entire epithelium is simply extruded into the buccal cavity. In pulmonates, the opening of the radular sheath is closed by the cuticular collostylar hood which is generated by a distinct epithelium which is proved to be stationary. When leaving the proliferation zone, the superior epithelium differentiates into supporting cells and mineralizing cells; the latter cause the hardening of the radular teeth and already degenerate in the middle of the sheath. The whole superior epithelium degenerates at the border to the collostylar hood-epithelium. In Lymnaea the degeneration zone is strictly confined whereas in Cepaea the collostylar hood and its generating epithelium extend into the radular sheath and the degeneration zone ranges over a distance of 3–5 rows of teeth. The proliferation zone of the inferior epithelium extends over the posterior half of the radular sheath, but the replacement rate is much lower than in the superior epithelium. Although the inferior epithelium carries the radula, it migrates slower than the radula. Obviously the radula has to be transported actively by apical protrusions of the cells, which penetrate into the radular membrane. At the opening of the radular sheath the inferior epithelium generates the adhesive layer and degenerates. During feeding, the adhesive layer has to maintain the firm mechanical connection between radula and distal radular epithelium. Autoradiographic experiments demonstrate that the distal radular epithelium is stationary. Nevertheless, the radula is known to advance to its degeneration zone. Special attention is paid to this problem. We strongly suspect that the transport of the adhesive layer and the radula is based on pseudopodial movements of apical protrusions characteristic for the distal radular epithelium. These protrusions interdigitate with the lower face of the adhesive layer. The mechanical connection has to be maintained and so the respective structures (tonofilaments and hemi-desmosomes) have to be continually renewed. This needs a high amount of energy and obviously results in the conspicuous concentration of mitochondria near the apical surface.Abbreviations al adhesive layer - ax axon - bc buccal cavity - bce buccal cavity epithelium - bl basal layer - bla basal labyrinth - bm basal membrane - bp basal plate - bpc basal plate cell - c cilia - ch collostylar hood - che collostylar hood-epithelium - cl cuticular layer - col collostyle - cr cell remnant - cts connective tissue sheath - d desmosome - dl upper layer - dre distal radular epithelium - dz degeneration zone - fe front edge - g granula - gol dictyosome - hd hemidesmosome - hl haemolymph - ie inferior epithelium - j jaw - ma tooth matrix - mc mineralizing cell - mem membranoblast - mfb microfibrills - mfl microfilaments - mgb multigranular body - mi mitochondria - mit mitosis - ml middle layer - mt microtubuli - mv microvilli - mw membrane whirl - n nucleus - nc necrotic cluster - nf nerve fibres - nsg neurosecretory granula - o odontophor - od odontoblast - odg odontoblast group - pod pre-odontoblast - rb residual body - rer rough endoplasmatic reticulum - rm radular membrane - rt radula teeth - sc supporting cell - se superior epithelium - sj septate junction - sro subradular organ - ss secretion substance - tf tonofilaments - tsm supramedian tensor muscle - tw terminal web - v vacuole - ves vesicle     

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.     

Ultrastructure and Phylogenetic Significance of Notaspidean Spermatozoa (Mollusca, Gastropoda, Opisthobranchia)

Spermatozoa of five notaspidean opisthobranchs [Berthellina citrina, Berthella ornata, Pleuro-branchus peroni, Pleurobranchaea maculata, Umbruculum sinicum] were examined using TEM. In all five species, the acrosome (sensu lato) consists of an apical vesicle (the acrosomal vesicle) and acrosomal pedestal. The acrosomal pedestal overlaps the nuclear apex, and in P. peroni (and possibly B. ornata) is periodically banded—-the first reported incidence of this type of substructure in any euthyneuran acrosome. Although sperm nuclei of P. peroni, B. ornata and B. citrina differ in length and also the number of keels present (nucleus 7 μm long with four/five keels present in Pleurobranchus; 17 μm long with one keel in Berthella; 15 μm long with a very weak keel in Berthellina), the basal invagination to which the centriolar derivative, axoneme and coarse fibres are attached is always poorly developed, and very little overlap between nucleus and midpiece occurs. In P. maculata and U. sinicum, the nucleus forms a helical cord around the axoneme and mitochondrial derivative such that it is not possible to recognize exclusively ‘nuclear’ and ‘midpiece’ regions of the spermatozoon. In all notaspideans investigated, (1) the axoneme, coarse fibres and glycogen helix are enclosed by the paracrystalline and matrix components of the mitochondrial derivative and (2) a dense ring structure (attached to the plasma membrane) and glycogen piece are observed. While the glycogen piece is very short (0.85–1.43 μm) with a very degenerate axoneme in B. citrina, B. ornata and P. peroni, this region of the spermatozoan is well developed (30–35 μm long) in U. sinicum and exhibits a fully intact 9 + 2 axoneme. The ‘glycogen piece’(or its presumed homologue) in P. maculata spermatozoa is very short (0.65 μm), devoid of any axonemal remnant and constructed of a hollow, internal cylinder attached to an outer (incomplete) shell, and contains scattered (glycogen) granules. Spermatozoal structure supports a close relationship between the genera Berthellina, Berthella and Pleurobranchus. These three genera have more distant links with Pleurobranchaea, while Umbraculum maintains an isolated, specialized position within the Notaspidea.     

On the taxonomy of Littorina africana (Mollusca: Gastropoda)

L. africana and L. knysnaensis are regarded as two morphs of a single species which exhibits a genetic cline along the south-eastern coast of southern Africa. The dark brown morph knysnaensis dominates the western, cooler end of the cline and is replaced by the pale blue morph africana at the warmer end of the cline. These conclusions are based on evidence from the latitudinal distributions, the complete range of intermediate forms regarding shell colour and shell morphology and the lack of differences in redular morphology, penial morphology or habitat.     

Ultrastructure of spermatozoa in Orthalicidae (Mollusca,Gastropoda, Stylommatophora) and its systematic implications

Sperm morphology of orthalicid gastropods Clessinia pagoda, Spixia tucumanensis, Plagiodontes daedaleus (Odontostominae) and Drymaeus hygrohylaeus, D. poecilus, Bostryx stelzneri (Bulimulinae) are examined and described for the first time using transmission electron microscopy. Spermatozoa show the general characteristic of Pulmonata: an acrosomal vesicle, sperm nucleus helical, mitochondrial derivative forming a continuous sheath with paracrystalline material and coarse fibers associated with axonemal doublets. Features in the acrosomal complex and shape of the nucleus distinguish orthalicid sperms from other stylommatophoran. The acrosomal pedestal is traversed by fine striations in all species examined except in S. tucumanensis. The structure and thickness of the perinuclear sheath with a single or double layer of electron-dense material ensheathing the nuclear apex is characteristic of the group. The presence of a subnuclear ring in Drymaeus, Bostryx and Clessinia species is also reported. A data matrix of eleven species per 34 characters (16 sperm plus 18 anatomical and shell characters) from orthalicids plus other stylommatophoran and systellommatophoran representative species was constructed. Three cladistic analyses (sperm-based, anatomical-based and a combined sperm + anatomical-based) were performed to test the phylogenetic potential of sperm ultrastructure in orthalicid systematics and understand how sperm characters affect the topology and resolution of the obtained trees. Stylommatophora resulted in a monophyletic clade in the sperm-based and in the combined-character analysis. Orthalicidae is monophyletic only in the combined-character cladogram. Within Orthalicidae, Odontostominae is recovered as a monophyletic clade in all analyses, while Bulimulinae is paraphyletic in all trees except in the combined phylogeny. The present study and cladistic analyses performed support the hypothesis that characters on sperm ultrastructure are informative for stylommatophoran systematic and phylogenetic approaches, providing synapomorphies at familiar, subfamiliar and generic level.