The sperm structure of <Emphasis Type="Italic">Galloisiana yuasai</Emphasis> (Insecta,Grylloblattodea) and implications for the phylogenetic position of Grylloblattodea

The sperm ultrastructure of the Grylloblattodea Galloisiana yuasai was described and the sperm characters were comparatively examined in several orthopteroid insect orders for inferring the phylogenetic placement of the Grylloblattodea. The spermatozoa of G. yuasai are joined in bundles (spermatodesms) containing 200 units. Major features of these spermatozoa include a monolayered acrosome, a 9+9+2 axoneme with 16-pfs accessory microtubules and expanded intertubular material, and an evident “centriole adjunct”. The diffused material observed between the axoneme and the mitochondrial derivatives is considered to be an extension of the three connecting bands observed in other orthopteroid taxa, similar to what happens in some orthopteran lineages. The presence of the connecting bands, even though modified in G. yuasai, suggests that the Grylloblattodea are to be placed in a clade with Mantophasmatodea, Mantodea and Orthoptera.     

A SIBLING NEW SPECIES OF GLYCASPIS TAYLOR (HOMOPTERA: PSYLLIDAE) AND REINSTATEMENT OF THREE SYNONYMS

The first two sibling species recognised in the genus Glycaspis Taylor are recorded from different eucalypt hosts; Glycaspis (Glycaspis) propensa sp.n. is described from the "box" group and is a sibling of G. (G.) dulcieana Moore from the "ironbark" group. The previously synonymised species G. mirabilis Moore, G. vellerosa Moore and G. convallaria Moore are reinstated as valid species.     

Phylogeny and biogeography of ice crawlers (Insecta: Grylloblattodea) based on six molecular loci: designating conservation status for Grylloblattodea species

Ice crawlers (Insecta: Grylloblattodea) are rarely encountered insects that consist of five genera representing 26 species from North America and Asia. Asian grylloblattids are the most diverse, but North American ice crawlers (genus Grylloblatta) are known for their adaptation to cold conditions. Phylogenetic relationships among grylloblattid species and genera are not known. Six genes were sampled in 35 individuals for 18S rRNA, 28S rRNA, histone 3, 12S rRNA, 16S rRNA, and cytochrome oxidase II from 21 populations of Grylloblatta, three populations from Japan (genus Galloisiana), and three populations from Russia (genus Grylloblattina). Phylogenetic analysis of these data with two mantophasmid outgroups in POY supported monophyletic genera, with Grylloblatta as sister to Grylloblattina. Grylloblatta was shown to contain two major lineages: a clade in Northern California and Oregon and a clade in Washington and Oregon. One new species and six candidate species are proposed. IUCN Red List Conservation Criteria were implemented to designate conservation status for each lineage.     

Taxonomic revision of three species of the genus Genarchopsis (Digenea: Hemiuroidea: Derogenidae) in Japan by molecular phylogenetic analyses

The taxonomic status of three nominal species of Genarchopsis (G. goppo Ozaki, 1925; G. gigi Yamaguti 1939; and G. fellicola Shimazu, 1995) (Digenea: Hemiuroidea: Derogenidae) was investigated by molecular phylogenetic analyses using partial sequences of the genomic ITS-1 region and the mitochondrial COI. The analyzed samples were divided into four groups: Lake Biwa, West Japan, Central Japan and G. fellicola. The Lake Biwa group, a sister taxon to the other three groups, was interpreted as G. gigi, so we concluded that G. gigi is valid; thus, this species is resurrected taxonomically. The specimens from the type host caught near the type locality of G. goppo were included in the West Japan group, so this group was regarded as G. goppo sensu stricto. Because the phylogenetic position of the Central Japan group could not be confirmed, it was identified tentatively as G. goppo, even though this species thus becomes paraphyletic. The taxonomic validity of G. fellicola was reconfirmed. The divergence time of G. gigi is discussed in relation to the geological history of Lake Biwa and the origin of host species.     

A Further Numerical Taxonomic Study of the Rhodochrous Group

A computer taxonomy was carried out as an aid to clarify the relationships between the genera Mycobacterium, and Nocardia as well as the ‘Rhodochrous’ group. The last group contained slightly acid-fast organisms isolated from sputa and soil and also included strains received as “Mycobacterium rhodochrous.” The group had been proposed as the genus Gordona by the present author. The rhodochrous group formed a cluster at the 87% similarity level and was clearly separated from all strains of the genera Mycobacterium and Nocardia. The cluster of the rhodochrous group was divided into seven subclusters at the 90% similarity level, and six of the seven corresponded to six species of the genus Gordona: G. bronchialis; G. rubropertincta (synonym G. rubra); G. terrae; G. aurantiaca; G. rhodochroa; G. rosea. If one accepts the concept that a phenetically well-defined cluster is a species, each of the subclusters of the rhodochrous group should be regarded as a species and the entire group as a genus. Discrepancies between the results of numerical classification and the results of lipid analyses remain as a problem to be solved in the future.     

Embryonic development of Galloisiana yuasai Asahina, with special reference to external morphology (insecta: Grylloblattodea)

The embryogenesis of Grylloblattodea, one of the most primitive of the polyneopteran orders, is described using Galloisiana yuasai with special reference to external morphology. The egg membranes are characterized by an endochorion crossed by numerous vertical aeropyles and a fairly thin vitelline membrane, features shared by Mantophasmatodea. The inner layer formation is of the fault type. Serosal elements in the amnioserosal fold differentiate into hydropylar cells, to function in water absorption together with specialized amniotic structures, i.e., an amniotic strand and a thickened amnion. The germ band is of the short germ type. The germ band immerses deep into the yolk after its full elongation along the egg surface, and in this respect blastokinesis closely resembles that of Mantophasmatodea. The embryological features, i.e., those on egg membranes and blastokinesis, may suggest a closer affinity of Grylloblattodea and Mantophasmatodea. Appendages, ectodermal invaginations, and sternal and pleural sclerites are discussed in the light of serial homology, to provide a new basis for elucidating the insect body plan. Appendages are divided into the proximal coxopodite and distal telopodite, the former being divided further into the subcoxa and coxa. Subcoxal and coxal elements are identified in the mandible as well as in the abdominal appendages. The subcoxa is divided into the epimeron and episternum by the pleural suture in thoracic segments. Likewise, in the abdominal segments the subcoxa is divided into two, although the homologs of the epimeron and episternum are not sclerotized, and in the labial segment the subcoxal derivative or the postmentum is divided into the submentum and mentum. Two coxal endites bulge out from the medial side of the gnathal appendages. The mandibular molar and incisor, maxillary lacinia and galea, and labial glossa and paraglossa are serially homologous with each other. In the thoracic segments the original embryonic sternum or "protosternum" is largely replaced by subcoxal elements, and merely remains as a small anterior presternum and a posterior spinasternum. A major part of the venter is represented by the derivatives of the episternum such as an extensive basisternum, katepisternum, and trochantin and the medial element of the epimeron. The pleuron is derived from the episternal elements or the anepisternum and preepisternum, which bears a spiracle in the mesothorax and metathorax, and the lateral element of the epimeron. The homolog of the preepisternum in the prothorax is the cervical sclerite, but with no spiracle developed. A median ventral invagination arises in the thoracic segments as a spina, and the homolog of the spina develops into the eversible sac in the first abdominal segment.     

Postembryonic development and homology of external genitalia in Galloisiana nipponensis (Caudell et king) (Notoptera : Grylloblattidae)

The postembryonic development of genital and postgenital segments of Galloisiana nipponensis (Notoptera : Grylloblattidae) have been described in detail. The coxite, stylus, epiproct and paraproct are already observed in the first nymphal instar, but the sexual characters are almost indistinguishable in the 1st to 4th nymphal instars, because the genital segments in these stages are not yet differentiated. In the 5th nymphal instar in the male, the paired primary phallic lobes start to appear at the hind margin of the 9th sternum. The rudiments of ventral valvulae in the female originate at the hind margin of the 8th sternum. In the 6th nymphal instar, the rudiments of inner valvulae begin to appear on the 9th sternum, in addition to the ventral valvula rudiments. The long dorsal valvulae are formed from the extension of nymphal coxite, while the nymphal styli are more reduced than those of younger instars.The ventral valvulae of the 8th abdominal segment are thought to be homologous with the inner valvulae of the 9th segment, judging from their site and mode of development. Further, the inner valvulae may also be homologous with the primary phallic lobes (phallomeres) of the male. The external genitalia in G. nipponensis originate from 2 structures: the dorsal valvulae (considered of appendicular origin), and the phallomeres; ventral and inner valvulae are regarded as mere integumentary outgrowths.     

Functional Morphology of the Phallosome in Glossina (Diptera, Glossinidae) and Its Evolutionary Implications

Current views on the phylo-genetjc relations of the three species groups within the genus Glossina are critically reviewed. In respect of the male genitalia, the fusca group is here regarded as more specialized than the morsitans group. The detailed phallosome morphology of G. austeni is compared with that of G. fuscipes , on the basis of their mode of action in living flies. The comparison is extended to fusca group species, using dead material only. The presumed phylogeny of Glossina is presented in outline; the evolution of the genus has involved a progressive invasion of riverine and later of forest habitats, from an original semi-arid grassy woodland habitat.     

DNA base composition and DNA relatedness among species of Trichosporon Behrend

The nuclear DNa of 28 species (30 strains investigated) of yeasts classified currently or previously in the genus Trichosporon. was analysed for its molar percentage of guanine + cytosine (mol% G+C). This criterion, together with biochemical characteristics, suggested the separation of the organisms studied into two groups. The first group, which appears related to the Ascomycetes, includes thirteen species with a G+C content lower than 50 mol% (34.7–48.8), and lacks urease (except T. margaritiferum). The second group appears related to the Basidiomycetes and includes fifteen species with a G+C content higher than 50 mol% (57–64) and has the ability to hydrolyse urea.A DNA homology experiment with T. beigelii and twelve other species of the second group showed very low values of complementarity with T. beigelii-labeled DNA. All these species must be considered as taxa other than T. beigelii.     

GRACILARIA HUMMII SP. NOV. (GRACILARIALES,RHODOPHYTA), A NEW NAME FOR THE AGAROPHYTE “GRACILARIA CONFERVOIDES” HARVESTED IN NORTH CAROLINA DURING WORLD WAR II1

Gracilaria hummii Hommers. et Freshwater is proposed as a new name for the inshore cylindrical species found in North Carolina that was treated as Gracilaria confervoides (L.) Grev. during World War II, and more recently as G. verrucosa (Huds.) Papenf. Molecular evidence places G. hummii in the Gracilis‐group in Gracilaria together with G. gracilis (Stackh.) Steentoft, L. M. Irvine et Farnham, the name currently applied to specimens formerly identified as G. confervoides and G. verrucosa. G. hummii differs from G. gracilis in possessing shallower male conceptacles in which the spermatangial filament originates from a surface cortical cell rather than from a subcortical cell. The cystocarps are similar, except that the gonimoblasts of G. hummii are attached to the base of the pericarp by numerous, prominent thickened terminal tubular cells and because terminal tubular cells are absent above the midregion of the cystocarp or in the vicinity of the ostiole. The gonimoblasts are subtended by a bundle of longitudinally oriented, thick‐walled secondary filaments of a type that has not been described before in Gracilaria but that may be characteristic of some other species. G. hummii occupies a basal position in the Gracilis‐group and is distinct from all other cylindrical North Carolina Gracilaria species, according to the molecular and morphological evidence.     

New species of mole crickets of the Gryllotalpa gryllotalpa group (Orthoptera: Gryllotalpidae) from Israel, based on morphology, song recordings, chromosomes and cuticular hydrocarbons, with comments on the distribution of the group in Europe and the Mediterranean region

Two sibling species of mole crickets of the Gryllotalpa gryllotalpa group inhabit Israel. Both are described as new species based on differences in their morphology, acoustic behaviour, chromosome number, cuticular hydrocarbon pattern and habitats. The species with males having 2n = 23 chromosomes, described in the past as the 'Dead Sea race', is designated as G. marismortui sp.n. It is an endangered species. In morphology and chromosome number it is similar to G. cossyrensis Bacetti & Capra from Italy, but differs in the composition of cuticular hydrocarbons. Gryllotalpa marismortui occurs in a limited geographical area along the shore of the Dead Sea. Another species, with 2n = 19 chromosomes, is described as G. tali sp.n. and is found throughout Israel. The calling songs of the males of the new species differ markedly. Females of the respective chromosomal numbers discriminate in favour of the homospecific call. Their habitats are strikingly different: G. marismortui lives in hypersaline soil along the Dead Sea shore (total chloride 187.25 meq %), G. tali inhabits freshwater soil (9.31 meq %). The distribution of the twelve known sibling species and one chromosomal race (not yet described) of the G. gryllotalpa group from Europe and the eastern Mediterranean is summarized. The zoogeography and phylogeny of the group are discussed.     

Multilocus phylogeny and species delimitation within the genus Glauconycteris (Chiroptera,Vespertilionidae), with the description of a new bat species from the Tshopo Province of the Democratic Republic of the Congo

The genus Glauconycteris Dobson, 1875 currently contains 12 species of butterfly bats, all endemic to sub‐Saharan Africa. Most species are rarely recorded, with half of the species known from less than six geographic localities. The taxonomic status of several species remains problematic. Here, we studied the systematics of butterfly bats using both morphological and molecular approaches. We examined 45 adult specimens for external anatomy and skull morphology, and investigated the phylogeny of Glauconycteris using DNA sequences from three mitochondrial genes and 116 individuals, which in addition to outgroup taxa, included nine of the twelve butterfly bat species currently recognized. Four additional nuclear genes were sequenced on a reduced sample of 69 individuals, covering the outgroup and Glauconycteris species. Our molecular results show that the genus Glauconycteris is monophyletic, and that it is the sister‐group of the Asian genus Hesperoptenus. Molecular dating estimates based on either Cytb or RAG2 data sets suggest that the ancestor of Glauconycteris migrated into Africa from Asia during the Tortonian age of the Late Miocene (11.6–7.2 Mya), while the basal diversification of the crown group occurred in Africa at around 6 ± 2 Mya. The species G. superba is found to be the sister‐group of G. variegata, questioning its placement in the recently described genus Niumbaha. The small species living in tropical rainforests constitute a robust clade, which contains three divergent lineages: (i) the “poensis” group, which is composed of G. poensis, G. alboguttata, G. argentata, and G. egeria; (ii) the “beatrix” group, which contains G. beatrix and G. curryae; and (iii) the “humeralis” group, which includes G. humeralis and a new species described herein. In the “poensis” group, G. egeria is found to be monophyletic in the nuclear tree, but polyphyletic in the mitochondrial tree. The reasons for this mito‐nuclear discordance are discussed.     

Four new species of Gyrodactylus von Nordmann, 1832 (Monogenea,Gyrodactylidae) on gobiid fishes: combined DNA and morphological analyses

Four Gyrodactylus species parasitising four closely related gobiid species in European coastal waters were studied and compared with G. arcuatus Bychowsky sensu Bychowsky & Poljansky (1953) from Gasterosteus aculatus . These were G. gondae n. sp. from Pomatoschistus minutus and P. lozanoi , G. flavescensis n. sp. from Gobiusculus flavescens , G. arcuatoides n. sp. from P. minutus and G. branchialis n. sp. from P. microps. Combined molecular and morphological analyses, as well as morphometric and statistical methods, were used. The ssrRNA V4 region and the complete ITS rDNA region were sequenced. Genetically the four new species are clearly distinct from G. arcuatus . From a morphological point of view, the haptoral hard parts of G. gondae n. sp., G. flavescensis n. sp. and G. arcuatoides n. sp. are related to those of G. arcuatus, while these parts of G. branchialis n. sp. are different, but related to those of G. quadratidigitus Longshaw, Pursglove & Shinn, 2003. For the latter two species, a new species group is formed. The V4 and ITS sequence analyses, however, indicate a close relationship between G. branchialis and the three G. arcuatus-like species.     

A brief evolutionary excursion comes to an end: the genetic relationship of British species of Gentianella sect. Gentianella (Gentianaceae)

 There is much taxonomic confusion in the genus Gentianella section Gentianella, five species of which are found in Britain. Gentianella anglica is a British endemic restricted in its distribution to the chalk downland of southern England. It is considered to be threatened because of habitat loss, and due to hybridisation with the closely related, widely distributed G. amarella. The Red Data Book species, G. uliginosa, shares morphological similarities with the other two species but has a very restricted distribution. Individual plants from populations across the range of these three species were analysed using AFLPs and universal cpSSRs to determine the degree of genetic variability within and between them, and to look for evidence of hybridisation. Samples of the two other Gentianella section Gentianella species found in Britain, G. campestris and G. germanica, were also included in the study. The analysis of chloroplast SSRs was not informative for G. anglica, G. amarella and G. uliginosa, while G. campestris and G. germanica differed from each other and the other three species at a single locus. Principal co-ordinate analysis of the AFLP data revealed only three distinct groups: one group contained G. campestris samples, another contained all samples of G. germanica, and a third contained all individuals from the other three species. In mixed populations of G. anglica and G. amarella, individuals of the two species were genetically more similar to each other than they were to individuals of the same species from other populations. It is proposed that G. anglica may be an early flowering form of G. amarella that has been created and maintained as a consequence of former grassland management practices. Received May 20, 2002; accepted October 28, 2002 Published online: March 31, 2003     

Phylogenetic systematics and historical biogeography of the Neotropical electric fish Gymnotus (Teleostei: Gymnotidae)

Phylogenetic interrelationships of the Neotropical electric fish genus Gymnotus are documented from comparative study of phenotypic data. A data matrix was compiled of 113 phenotypic characters for 40 taxa, including 31 recognized Gymnotus species, six allopatric populations of G. carapo, two allopatric populations of G. coropinae, and three gymno‐tiform outgroups. MP analysis yielded 15 trees of equal length, the strict consensus of which is presented as a working hypothesis of Gymnotus interrelationships. Diagnoses are presented for 26 clades, including three species groups; the G. cylindricus group with two species restricted to Middle America, the G. pantherinus group with 12 species in South America, and the G. carapo group with 16 species in South America. The basal division of Gymnotus is between clades endemic to Middle and South America. Both the G. pantherinus and G. carapo groups include trans‐Andean sister‐taxon pairs, suggesting a minimum date for the origins of these groups in the late Middle Miocene (c. 12 Ma.). The geographically widespread species G. carapo is paraphyletic. Analysis of character state evolution shows characters of external morphology are more phylogenetically plastic and provide more phylogenetic information in recent branches than do characters of internal morphology, which themselves provide the more information in deeper branches. Nine regional species assemblages of Gymnotus are recognized, none of which is monophyletic. There are at least two independent origins of Gymnotus species in sediment rich, high conductivity, perennially hypoxic whitewater floodplains (varzea´) derived from an ancestral condition of being restricted to low conductivity non‐floodplain (terra firme) black and clearwater rivers and streams. These phylogenetic, biogeographic and ecological patterns suggest a lengthy and complex history involving numerous instances of speciation, extinction, migration and coexistence in sympatry. Evolution in Gymnotus has been a continent‐wide phenomenon; i.e. Amazonian species richness is not a consequence of strictly Amazonian processes. These patterns are similar to those of other highly diverse groups of Neotropical fishes and do not resemble those of monophyletic, rapidly generated species flocks.     

Gymnopilus maritimus (Basidiomycota, Agaricales), a new species from coastal psammophilous plant communities of northern Sardinia, Italy, and notes on G. arenophilus

The new species Gymnopilus maritimus is described from coastal plant communities of Juncus maritimus, growing on sandy soil or on decaying plants, from northwestern Sardinia (Italy). The distinguishing features of G. maritimus are: (1) an unusual habitat, (2) robust basidiomata, (3) mild taste, and (4) big and strongly warted spores. The new species is compared with the micromorphologically similar species G. fulgens sensu auct. Brit. p.p. and the biogeographically and ecologically similar species G. arenophilus, as well as with other European species. A photograph of fresh material, drawings of the main micromorphological features, and FESEM and optical microscope microphotographs of basidiospores are added. Furthermore, some notes on micromorphological characters of G. arenophilus are presented and its distribution area enlarged with a record from France. A key for the European species of Gymnopilus morphologically, ecologically, and/or biogeographically related to G. maritimus is presented. The phylogeny inferred from ITS rDNA sequences revealed that G. maritimus represents an independent species and that it is not related to G. arenophilus or G. fulgens. It is the sister group of the clade containing G. imperialis and G. spectabilis, but with a bootstrap support below 50%. The characters shared by the species in this clade are: (1) robust basidiomata, (2) pileus fibrillose or scaly-fibrillose, and (3) spores longer than 8 μm, dextrinoid and strongly warted. Gymnopilus imperialis and G. spectabilis differ by the basidiomata with membranous ring in the stem, living on conifers or decaying wood, and having narrower or wider spores, respectively. Taxonomical novelties: Gymnopilus maritimus Contu, Guzm.-Dáv., A. Ortega & Vizzini     

Status and relationships of the extinct giant Canary Island lizard Gallotia goliath (Reptilia: Lacertidae), assessed using ancient mtDNA from its mummified remains

Ancient mitochondrial DNA sequences (378 base pairs of cytochrome b and 368 of 12S rRNA) extracted from a mummified extinct giant lizard, Gallotia goliath , from eastern Tenerife, Canary Islands, were used to assess the species status and relationship of this form within the genus. G. goliath is clearly a member of the G. simonyi group of the western Canary islands (Tenerife, La Gomera, El Hierro and La Palma) and is not closely related to the giant G. stehlini of Gran Canaria. Contrary to recent opinion, it is phylogenetically distinct, within the G. simonyi group, from the extant G. simonyi of El Hierro and also from the recently discovered live G. gomerana on La Gomera and from G. intermedia in north-western Tenerife. It may be the sister taxon of either all the other members of the G. simonyi group or of G. intermedia . The phylogenetic distinctness of G. goliath makes Tenerife unique among oceanic islands in having had one giant and two medium-sized lizard species that were probably substantially herbivorous, the others being G. intermedia and G. galloti . Gallotia shows great community differences on other islands in the Canaries, two having a single small species, one a single giant, and three a giant and a medium-sized form. © 2003 The Linnean Society of London, Biological Journal of the Linnean Society , 2003, 80 , 659–670.     

GENETIC DIVERSITY,INTROGRESSION, AND INDEPENDENT DOMESTICATION OF OLD WORLD CULTIVATED COTTONS

Gossypium arboreum L. and G. herbaceum L. are the diploid species of cultivated cotton. Little is known regarding the time and place of domestication of either species. Because G. arboreum is known only as a cultigen, others have proposed that it arose from domesticated G. herbaceum during the more than 5,000-year history of Old World cotton cultivation, with wild G. herbaceum subsp. africanum (Watt) Mauer as the putative ancestor of both species. An alternative hypothesis is that the two species have independent origins from progenitors that diverged prior to domestication. The relative merits of these opposing hypotheses were evaluated using data derived from starch gel electrophoresis of enzymes. One hundred and three accessions of G. arboreum and 31 accessions of G. herbaceum were examined for allelic variation at 40 allozyme loci. All measures of genetic variability demonstrated that G. arboreum contains greater diversity than G. herbaceum, although both species have relatively low levels of allozyme variation. In contrast to expectations based on morphology and other chemical data sets, the two species are highly differentiated with respect to allozyme composition. Gossypium arboreum and G. herbaceum each contain a large number of unique alleles and are fixed or nearly fixed for alternate alleles at 8 loci. Five allozyme loci have alleles that are rare in one of the two species but common in the other. Based on restricted occurrence of these alleles to areas with a long history of sympatric cultivation and the geographic distribution of a null allele, we suggest that a significant portion of the allelic diversity in both species results from historical, bidirectional interspecific introgression. The interspecific genetic identity estimate (0.74) is markedly lower than for documented progenitor-derivative and crop-ancestor species pairs. Based on these data, as well as previous cytogenetic data and the observation of F₂ breakdown in interspecific crosses, we suggest that cultivated G. arboreum and G. herbaceum were independently domesticated from divergent ancestors.     

Isozyme criteria in the testing of phyletic relationships between species of Gobius and related eastern Atlantic-Mediterranean genera (Teleostei: Gobüdae)

Horizontal starch gel electrophoresis was employed to investigate allozyme variation at 19 isozyme loci, in nine closely-related Atlantic-Mediterranean gobiid fish species, to assess genetic relationships, and to compare this with a phyletic hypothesis based on morphological apo-morphies. The species examined were Gobius niger, G. auratus, G. cruentatus, G. paganellus, Mauligobius maderensis, Zosterisessor ophiocephalus, Thorogobius ephippiatus, Padogobius martensii and P. nigricans . Various phenetic and cladistic analyses were performed on isozyme and morphological data. The phenetic and cladistic results from morphological data, and the cladistic results from isozyme data, were largely comparable, with P. martensü and P. nigricans forming a sister group to all the other taxa, species of Cobius forming a crown group within the latter, and stem lines formed by Mauligobius, Zosterisessor and Thorogobius , respectively. The isozymic estimates of genetic distance differed from these chiefly in the relative positions of Zosterisessor and G. auratus , which appear less similar to other species of Gohius. G. auratus shows many alleles which are unique within the genus Gobius ; possible explanations for this are put forward. The position of nigricans as congeneric with Padogobius martensü is confirmed by both phenetic and cladistic analyses of isozyme data.