A revision of the family Zoogonidae Odhner, 1902 (Platyhelminthes: Digenea): Introduction and subfamily Zoogoninae

Summary The 28 major morphological characters of members of the family Zoogonidae are discussed. A key to the subfamilies is given and a detailed revision of the subfamily is presented. Nine genera and 27 species are treated in detail with keys and cladograms to genera and species. The genera and species covered are: Zoogonus rubellus, Z. argentopsi, Z. dextrocirrus, Z. lasius, Z. [=Zoogonoides] mazuri (Korotaeva) n. comb., Z. pagrosomi, Proparvipyrum israelense, Parvipyrum acanthuri, Brevicreadium congeri, Diphterostomum brusinae, D. albulae, D. americanum, D. betencourti, D. indicum, D. magnacetabulum, D. vividum, Pseudozoogonoides subaequiporus, P. ugui, Glaucivermis spinosus, Neozoogonus californicus, N. longicecus, N. malacanthi, Zoogonoides viviparus, Z. acanthogobii, Z. laevis, Z. pyriformis, Z. yamagutii. ac]19851127     

Studies on the morphology and anatomy of the Antarctic nudibranch genera Pseudotritonia Thiele, 1912 and Telarma Odhner, 1934 with a discussion of the family Gharcotiidae Odhner, 1926 (Nudibranchia: Opisthobranchia)

The external morphology and anatomy of the Antarctic nudibranchs Pseudotritonia quadrangularis Thiele, 1912, P. gracilidens Odhner, 1944 and Telarma antarctica Odhner, 1934 are redescribed. Both genera, which were only known by two or one specimen, do not possess a cnidosac, but there is a terminal swelling at the termination of the digestive glandular ramifications in P. gracilidens. This sac is composed of cells with a large vacuole. A glandular stripe on the right side of the body above the genital openings, nephroproct and anal papilla is considered to be a synapomorphy for both genera. The validity of the family Charcotiidae and the affinities to the aeolid genus Notaeolidia Eliot, 1905 and taxa of the Arminacea are discussed.     

On the morphology and life-history of Hemiurus luehei Odhner, 1905 (Digenea: Hemiuridae)

Previously undescribed cystophorous cercariae which develop in sporocyst germinal sacs in the tectibranch opisthobranch Philine denticulata (Adams) are shown to be cercariae of Hemiurus luehei Odhner, 1905 (Hemiuridae), a common stomach parasite of clupeid and salmonid fishes off the Atlantic coast of Europe, in the Baltic and the Mediterranean Sea. The free-swimming cercariae are seized by calanoid copepods. Temora longicornis (Müller) and Acartia tonsa Dana acted as suitable experimental intermediate hosts. Pressure by the copepod mouth appendages causes delivery tube eversion and the injection of the cercarcial body into the haemocoel. Sagitta sp. was found naturally infected with a metacercaria of H. luehei. The cystophorous cercaria and metacercaria are described using light and scanning electron microscopy. Adults from herring were examined under the stereoscan electron microscope.     

A revision of the family Zoogonidae Odhner, 1902 (Platyhelminthes: Digenea): Subfamily Lepidophyllinae and comments on some aspects of biology

Summary A detailed revision of the zoogonid subfamily Lepidophyllinae is presented, using morphological characters discussed in an earlier paper. Twelve genera and 50 species are treated in detail with keys and cladograms to genera and species. The genera and species covered are: Lepidophyllum steenstrupi, L. appyi, L. armatum, L. brachycladium, L. cameroni, L. pleuronectini, L. pyriforme, L. schantaricum, Urinatrema hispidum, U. hirudinacea, Panopula cavernossa, P. bridgeri, P. spinosa, Limnoderetrema minutum (Manter, 1954) [formerly Deretrema] n.g. (in freshwater fishes; genital pore at oral sucker or pharynx level), n. comb., Brachyenteron peristedioni, B. acropomatis, B. campbelli, B. doederleiniae, B. magnibursatum, B. parexocoeti, B. pycnorganum, Steganodermatoides kergeleni, S. agassizi, S. allocytti, S. maceri, Neosteganoderma glandulosum, N. infundibulum, Proctophantastes abyssorum, P. gillissi, Deretrema (Deretrema) fusillus, D. (D.) cholaeum, D. (D.) pacificum, D. (Spinoderetrema) plotosi, D. (S.) acutum, D. (S.) fellis, D. (S.) ovale, D.(S.) sebastodis, D. (Luxitrema) philippinensis, D. plagiorchis, Pseudochetosoma salmonicola, Overstreetia sodwanaensis, Steganoderma (Steganoderma) formosum, S. (S.) atherinae, S. (Lecithostaphylus) retroflexum, S. (L.) depauperati (Yamaguti, 1970) n. comb., S. (L.) hemirhamphi, S. (L.) nitens, S. (L.) parexocoeti, S. macrophallos, S. oviformis. Most zoogonids were found to exhibit some level of predilection for a particular piscine host group, but little general information on the zoogeography of the group was discovered. Ultrastructural evidence is presented suggesting that the membranous egg-capsule of the zoogonines shows vestiges of the three layers of a normal tanned egg-shell.     

The origin of Lecithodesmus (Digenea: Campulidae) based on ND3 gene comparison

Species of Lecithodesmus (Campulidae) occur almost exclusively in baleen whales throughout a wide geographical distribution. Other campulids occur only in odontocetes and, secondarily, in pinnipeds and the sea otter. Therefore, the ancestor of Lecithodesmus might have either cospeciated with mysticetes during the early divergence of mysticete and odontocete cetaceans or originated later via host switching. We evaluate both possibilities based on a phylogenetic analysis. The ND3 mitochondrial gene sequence of a species of Lecithodesmus was included in a previous partial molecular phylogeny of the Campulidae. Fasciola hepatica and Dicrocoelium dendriticum were used as outgroups. Maximum parsimony, neighbor-joining, and maximum likelihood methods indicated a nonbasal position of Lecithodesmus sp. in the tree, suggesting that the ancestor of Lecithodesmus colonized mysticetes from campulids of odontocetes. This result emphasizes the importance of host-switching processes in the development of the helminth fauna of marine vertebrates, as previously suggested.     

Urotrematidae Poche, 1926 (Platyhelminthes: Digenea) in Chinese freshwater fishes

The genera Sinineobucephalopsis and Sinogastromyzontrema are diagnosed and placed in the family Urotrematidae. Sinineobucephalopsis macrocirrus is redescribed from Glyptothorax fokiensis in Guangdong and Hunan provinces. Urotrema glyptothoraci is considered synonymous with Sinineobucephalopsis macrocirrus. The new combinations Sinineobucephalopsis postlecitha, Sinineobucephalopsis proeilecitha and Sinineobucephalopsis sinipercae are formed for species originally placed in Urotrema and Urotrematulum. Sinogastromyzontrema guangxiensis is redescribed from Sinogastromyzon wui in Guangxi Province. Urotrema parallelorchis is considered a synonym of Sinogastromyzontrema guangxiensis. A key to the species or species-complexes in the family Urotrematidae is given.     

Phylogenetic systematic evaluation of a classification of the Zoogonidae Odhner, 1902 (Cercomeria: Trematoda: Digenea: Plagiorchiiformes)

A recent systematic study of the digenean family Zoogonidae presented a series of cladograms, which are the product of phylogenetic systematic, or cladistic, analysis. However, one of the two subfamilies and nine of the 21 genera recognised in that study lacked putative synapomorphies, a requirement for phylogenetic systematic studies. This study presents a re-analysis of the database for the zoogonids, based on rigorous application of phylogenetic systematic methods. A new phylogenetic tree is presented, which better fits the original data than the published tree (with a consistency index of 52.3% vs. 46.3%). Four subfamilies, three monophyletic and one of uncertain status, and 10 genera could be recognised phylogenetically. This would affect the nomenclatorial status of one-third (26) of the species in the family. However, it is recommended that another analysis, based on more characters, be carried out before nomenclatorial changes are proposed.     

Biometric variability of Hadwenius tursionis (Marchi, 1873) (Digenea,Campulidae) from the intestine of the bottlenose dolphin Tursiops truncatus (Montagu, 1821)

The species discrimination in the family Campulidae is largely based on morphometric characters (absolute measurements and ratios). To assess the variability of this characters we have studied the campulid species Hadwenius tursionis from four Mediterranean bottlenose dolphins Tursiops truncatus from the Western Mediterranean. In H. tursionis most absolute measurements differed significantly across the infrapopulations, showing high coefficients of variation. Only egg dimensions appeared to be fairly conservative. Body ratios were less variable than absolute measurements, but growth patterns did vary significantly between the infrapopulations studied. None of the ratios previously used in relation to other species of the family Campulidae indicated either isometric or allometric growth in this case. Only the ratios concerning the size of the gonads and the shape of the gonads and suckers showed isometry. It is concluded that absolute measurements and ratios should be used with caution in relation to the separation of species of the family Campulidae unless their intrinsic variability can be established beforehand.     

Lepocreadiidae Odhner, 1905 and Aephnidiogenidae Yamaguti, 1934 (Digenea: Lepocreadioidea) of fishes from Moreton Bay,Queensland, Australia,with the erection of a new family and genus

Digeneans of the lepocreadioid families Lepocreadiidae Odhner, 1905 and Aephnidiogenidae Yamaguti, 1934 from Moreton Bay, off southern Queensland, Australia, are recorded, along with the erection of a new family, Gibsonivermidae. Molecular data were generated for all representatives of these families collected during this study and a phylogram for members of the superfamily was generated based on the partial 28S rDNA dataset, placing these species in context with those previously sequenced. This phylogenetic analysis demonstrates that the monotypic Gibsonivermis Bray, Cribb & Barker, 1997 is isolated from all other lepocreadioids and supports the erection of Gibsonivermidae n. fam., which is defined morphologically, based particularly on the uniquely elongated male terminal genitalia, the distribution of the uterus in the forebody and the presence of a uroproct. Mobahincia teirae n. g., n. sp. is reported from Platax teira (Forsskål) in Moreton Bay and off Heron Island and New Caledonia. Recognition of this new genus is based on molecular results and the combination of caeca abutting the posterior body wall and the lack of an anterior body scoop or flanges. The following lepocreadioid species are reported from Moreton Bay for the first time: Bianium arabicum Sey, 1996 in Lagocephalus lunaris (Bloch & Schneider), Diploproctodaeum cf. monstrosum Bray, Cribb & Justine, 2010 in Arothron hispidus (Linnaeus), Multitestis magnacetabulum Mamaev, 1970 and Neomultitestis aspidogastriformis Bray & Cribb, 2003 in Platax teira and Opechona austrobacillaris Bray & Cribb, 1998 in Pomatomus saltatrix (Linnaeus). Bianium plicitum (Linton, 1928) is reported from Torquigener squamicauda (Ogilby) for the first time. Sequences of newly collected specimens of Austroholorchis sprenti (Gibson, 1987) indicate that the species forms a clade with other members of the Aephnidiogenidae, agreeing with its morphology. The phylogenetic status of all newly sequenced species is discussed.     

A multivariate morphometric analysis and systematic review of Pseudonaja (Serpentes,Elapidae, Hydrophiinae)

Pseudonaja is a clade of seven nominal species of elapid snakes distributed throughout Australia and in southern New Guinea. The species‐level systematics of this group is generally considered to be problematic. A recent phylogenetic analysis of mitochondrial DNA sequences for a geographically extensive series of Pseudonaja specimens revealed nine major clades, of which six largely coincide with nominal species (P. affinis, P. guttata, P. inframacula, P. ingrami, P. modesta and P. textilis). The three remaining clades are composed of specimens currently referred to P. nuchalis. This paper presents a multivariate analysis of 30 morphometric variables recorded for 220 specimens, representing the P. affinis, P. inframacula, P. textilis and three P. nuchalis clades (P. guttata, P. ingrami and P. modesta are well‐demarcated species and, accordingly, were not considered). The morphometric data readily separate these putative lineages, affording compelling evidence that they constitute evolutionary species. The names aspidorhyncha and mengdeni are resurrected for two of the three species presently recognized as P. nuchalis. These species, P. affinis, P. inframacula, P. nuchalis and P. textilis are redescribed. © 2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2009, 155 , 171–197.     

The Synthesium Brachycladiidae Odhner, 1905 (Digenea) association with hosts based on nuclear and mitochondrial genes

Synthesium pontoporiae, exclusive parasite of the endangered dolphin, Pontoporia blainvillei, is endemic and restricted to the South Atlantic and belongs to the Brachycladiidae family Odhner, 1905. The study of this family has been limited by the difficulty of accessing the parasites from their marine mammal hosts and as a consequence there is a paucity of genetic information available. Herein we present a genetic analysis using 18S rDNA sequences of S. pontoporiae and S. tursionis and the ND3 mtDNA sequence of S. pontoporiae. The genetic analysis of 18S rDNA sequences of brachycladiids and acanthocolpids determined two major clusters associated with their definitive hosts, marine mammals and fishes, respectively. Considering the tree topology of brachycladiids ND3 mtDNA gene, two clusters were defined, one with the Synthesium species. This work contributes with fundamental genetic information on S. pontoporiae, and suggests a Brachycladiidae genetic evolution related to their hosts.     

The Lepocreadiidae Odhner, 1905 (Digenea) of fishes from the north-east Atlantic: summary paper,with keys and checklists

Results published in seven previous papers on the Lepocreadiidae are summarised and keys are given to the 19 species of lepocreadiid parasite from north-eastern Atlantic fishes. Diagnoses for the family and the two subfamilies represented in the north-eastern Atlantic are presented. Parasite-host and host-parasite lists, arranged in taxonomic order, are given. The NE Atlantic fauna is dominated numerically by members of the subfamily Lepidapedinae, which occur most frequently in gadiform fishes, often in the deep-sea.The three lepocreadiine forms are parasites of cosmopolitan fishes or fish families that predominate in warm waters.     

Contribution to the taxonomy of the Ectocarpales (Phaeophyceae) from the coast of Karachi, Pakistan

BEGUM, M. & KHATOON, N., 1992. Contribution to the taxonomy of the Ectocarpales (Phaeophyceae) from the coast of Karachi (Pakistan). The taxomony and ecology of five species belonging to the order Ectocarpales, representing three genera– Ectocarpus with one species, Hincksia with three species and Nemacystus with one species–are studied and reported as new records for the coast of Karachi, Pakistan.     

Contribution to the taxonomy and phylogeny of Sarcocapnos DC. (Fumariaceae)

 To solve problems concerning the status of the taxa described in the genus Sarcocapnos, we have conducted a study using morphological, pollen morphology (light microscopy), cytogenetic and molecular techniques. Focusing on the last technique, we have sequenced ITS-1 and ITS-2 of nuclear rDNA. The species differ basically according to 5 morphological traits (leaf shape, flower spur, corolla colour, corolla size, and crest of the stigmatic surface). The cytogenetic analyses indicated n=16 to be the standard chromosome number. The ITS analyses showed that the genus is monophyletic, defining two main well-supported clades, one containing S. saetabensis and S. enneaphylla, and one containing the rest of the species. In this second clade, S. speciosa, S. pulcherrima, and S. baetica subsp. ardalii are related, as are S. integrifolia, S. crassifolia subsp. crassifolia, and S. crassifolia subsp. atlantis; S. baetica subsp. baetica forms a trichotomy with the foregoing groups. S. speciosa is shown to be a species separate from S. crassifolia subsp. crassifolia, as in the case of S. baetica with respect to S. integrifolia. Palynologically, the parameters used enabled us to establish clear differences between the taxa, often corroborating the macromorphological and genetic data. The flower spur has been reduced several times in different groups of the genus, for which the classifications established on the basis of this trait are paraphyletic. Received July 16, 2002; accepted December 11, 2002 Published online: March 31, 2003     

A multivariate comparison between three NW. European populations ofArtemisia norvegica (Asteraceae) by means of chemometric and morphometric data

Specimens from Scotland, S. and C. Norway were grown in the botanical garden of Bergen, Norway. Some of the Scottish specimens came from a meristem tissue culture. The specimens were compared by a principal component analysis of lipids and related compounds, and of morphological characters from leaves and flowers. The populations differed from each other, but some overlap was found in leaf characters. The results are discussed in relation to distribution and immigration history, and it is argued that the differences among the populations may have evolved in postglacial time.     

A multivariate approach to assess the taxonomic utility of morphometric characters inDoronicum (Asteraceae, Senecioneae)

A multivariate morphometric study based on 22 characters was carried out on samples of the 26 accepted species of the Eurasian and north-African genusDoronicum, to assess whether quantitative, mainly continuous, characters were of use in suggesting, delimiting and structuring natural groups. The study concentrated on three-medium sized to small groups: Mediterranean (D. plantagineum group), European (D. grandiflorum group), and central Asian. Two questions were addressed for each of these groups: species delimitation within them and the inclusion/exclusion of species of uncertain, but putative close affinities. Principal components analysis and discriminant analysis (canonical variates analysis), gave moderately satisfactory results. The addition of doubtfully-related species to presumably natural groups increased cohesiveness of the groups because the between-species differences were minimized when placed in a broader more diverse context. The relative status of the uncertain species, either as outliers or true members, was appropriately depicted by the scatterplots, but more convincingly solved when the species turned out to be outliers. Discrimination between species, which belong to cohesive groups was effective in some cases at least when using discriminant analysis, and the exceptions involved particularly problematic taxa. It was concluded that morphometric characters are a useful complementary source of information in groups where qualitative (in particular, shared) characters are scarce as is the case inDoronicum.