Topotypic Material Research Articles

Taxonomic revision of 14 south‐western European species of Isotomurus (Collembola, Isotomidae), with description of four new species and the designation of the neotype for I. palustris

A detailed study of morphological characters in populations of the collembolan genus Isotomurus from southern Europe is presented. Isotomurus is a cosmopolitan genus in which species diagnosis primarily relies on pigmentation patterns and chaetotaxy. Correct diagnosis is often complicated by the high variability of these characters. This has created confusion in the systematics of the genus and has led to descriptions of several varieties, colour forms and subspecies, the taxonomic value of which needs revision. Complete screening of pigmentation patterns and the study of sensorial chaetotaxy in specimens collected in 37 localities across south‐western France, Italy and Switzerland have enabled 14 species to be distinguished; one taxon is elevated to the rank of species (Isotomurus prasinus) and four new species are described: I. hadriaticus, I. aetnensis, I. gallicus and I. pseudopalustris. The neotype for I. palustris is designated in order to clarify the taxonomic status and the type locality for this species whose topotypic material is lost. A redescription of previously known species on the basis of the newly introduced characters is also presented, together with a key to the south‐western European species of Isotomurus. Following this study, the two species I. indipendente and I. italicus came to be junior synonyms of I. palustris and I. fucicolus, respectively. A cladistic analysis of 14 morphological characters provides a phylogenetic reconstruction which is largely concordant with previous taxonomic works.

New species of Heterodrilus (Oligochaeta, Tubificidae) and records of H. maiusculus from the Mediterranean Sea

The exclusively marine genus Heterodrilus is common in sandy sediments, particularly in tropical and subtropical seas. In this paper, H. maiusculus, previously known only from Saudi Arabia, is recorded from Corsica and Italy, and it is shown to have bifid pe‐nial chaetae, a trait confirmed also for paratypic and topotypic material of the species. Two new species are described: H. ursulae sp.n. and H. tripartitus sp.n., both from Italy. Heterodrilus ur‐sulae has trifid anterior chaetae, short, characteristically bent and bipartite atria, and long spermathecal ducts; H. tripartitus has bifid anterior chaetae and spermathecae with distinct vestibules, the latter feature being unique within the genus.

An amended description of the theropod footprint Bueckeburgichnus maximus Kuhn 1958, and its bearing on the megalosaur tracks debate

The ichnospecies Bueckeburgichnus maximus (Kuhn 1958) is amended and illustrated on the basis of type and topotype material, from the Lower Cretaceous of Germany, showing that previous illustrations were morphologically incorrect and misleading. The track is unequivocally that of a large functionally tridactyl theropod, but can not confidently be attributed to a megalosaurid dinosaur, as suggested by ichnological tradition. Bueckeburgichnus has been compared with theropod tracks from the Upper Jurassic of Portugal labeled as Eutynichnium, Megalosaurus, Megalosauripus and Megalosauropus, but the comparisons are tenuous and compromised by invalid ichnotaxonomic treatment of all four ichnogenera. Megalosaurus must be abandoned as an ichnotaxonomic descriptor. The status of Megalosauripus, Megalosauropus and Eutynichnium is under further investigation but preliminary results suggest that Bueckeburgichnus maximus is distinct from any ichnospecies assigned to these two ichnogenera.

Anatomy and systematics of the prosauropod dinosaur Thecodontosaurus antiquus from the upper Triassic of southwest England

ABSTRACT Re-examination of the surviving specimens of Thecodontosaurus antiquus indicates that this plesiomorphic sauropodomorph can be diagnosed on the basis of elongate basipterygoid processes, a relatively short dentary, and a squared posterior process of the ilium. Although much of the original topotype material found in the 1830s in Bristol, England, has now been lost, some 245 specimens remain. These indicate a small, gracile prosauropod, up to 2.5 m in length, distinguished primarily on the absence of derived characters seen in other prosauropods. Although attempts were formerly made to subdivide the Bristol specimens into several dinosaurian, and other, taxa, most appear to pertain to the prosauropod Thecodontosaurus antiquus. The specimens do indicate a clear separation into two morphs, a gracile and a robust form, presumably evidence of sexual dimorphism. A juvenile Thecodontosaurus sp. from South Wales may belong to the same species. A cladistic analysis indicates that Prosauropoda is probably a clade, rather than a series of outgroups to Sauropoda, but support for this conclusion is weak. Echoing other recent cladistic analyses, stronger support is found for the existence of a clade Sauropodomorpha, made up from Prosauropoda + Sauropoda, for the clade Sauropoda itself, and for the clade Eusauropoda within Sauropoda.

Allohyaena (Mammalia: Carnivora): giant hyaenid from the Late Miocene of Hungary

The topotypic material of the giant Late Miocene hyaenid Allohyaena kadici Kretzoi is described. New data on the deciduous dentition shows unambiguously that A. kadici is a hyaenid and not a percrocutid as reported by some previous authors. A. kadici is compared to the large hyaenids Adcrocuta eximia and Crocuta crocuta. These comparisons show that A. kadici has a mixture of primitive characters such as dp4 morphology, retention of m2, long and slender premolars and a large protocone on P4, and derived characters such as a preparastyle on P4, an internal root on P3 and a uniquely derived talonid structure of ml. This combination of features makes A. kadici difficult to classify, but it is considered to probably be most closely related to derived, bone-cracking hyaenids such as Pachycrocuta and Crocuta. A. kadici is rare in the fossil record, being found at only two sites. We suggest that the reason for this rarity is that it had a geographic and stratigraphic range which is poorly sampled in the Miocene fossil record of Europe.

Allohyaena (Marnmalia: Carnivora): giant hyaenid from the Late Miocene of Hungary

The topotypic material of the giant Late Miocene hyaenid Allohyaena kadici Kretzoi is described. New data on the deciduous dentition shows unambiguously that A. kadici is a hyaenid and not a percrocutid as reported by some previous authors. A. kadici is compared to the large hyaenids Adcrocuta eximia and Crocuta crocuta. These comparisons show that A. kadici has a mixture of primitive characters such as dp4 morphology, retention of m2, long and slender premolars and a large protocone on P4, and derived characters such as a preparastyle on P4, an internal root on P3 and a uniquely derived talonid structure of ml. This combination of features makes A. kadici difficult to classify, but it is considered to probably be most closely related to derived, bone-cracking hyaenids such as Pachycrocuta and Crocuta. A. kadici is rare in the fossil record, being found at only two sites. We suggest that the reason for this rarity is that it had a geographic and stratigraphic range which is poorly sampled in the Miocene fossil record of Europe.

Revision of the Charophyte Flora of the Siwalik Group (Neogene-Quaternary) of the Lesser Himalaya, India

On the basis of a re-examination of the topotype material of the Siwalikcharophytes described by Bhatia and Mathur (1978) and on additional materialcollected from six new localities, the taxonomy of the charophyte flora isbrought up to date. In all, 17 taxa are recorded and illustrated. At thespecific level, generic shifts have been made in the case ofChara rantzieni sivalensis,Chara contraria (non Braun et Kuetzing),Tectochara sahnii andSphaerochara pecki. The following taxa were hitherto notknown from the Siwalik Group:Chara globularis globularis Thuillier,C. globularis aspera (Deth. ex Wild.),C. sp. cf. hispida Linné,Lychnothamnus breviovatus Lu & Luo,Lamprothamnium papulosum (Wallr.) Groves,L. succinctum (A.Braun),Nitellopsis (Tectochara)bicarinata (MÄdler), N.(T.) helvetica (MÄdler) andSphaerochara prolifera (Zizermann ex A.Braun).

Neotricula aperta (Gastropoda: Pomatiopsidae), the intermediate host of Schistosoma mekongi: allozyme variation and relationships between Khmer, Lao, and Thai populations

AbstractVariation was studied at 13 electrophoretically detected allozyme loci for samples of six populations of the polytypic snail Neotricula aperta; this was the first time that topotypic material from southern Laos and material from Kampuchea had been so examined. In addition, samples of all three strains of N. aperta from North East Thailand were examined and the findings compared with those of Staub et al. (1990). The samples were taken from the Mekong and Mul rivers of the Lower Mekong Basin. A small sample was also included from a γ‐strain population in the Xé‐Bang‐Fai (XBF) river of central Laos. The γ‐strain population of southern Laos and of Kampuchea has been shown to act as an intermediate host for Schistosoma mekongi.In contrast to the work of Staub et al. (1990) significant multilocus genetic differences (Nei's D, DN= 0.6–1.2) were detected between the three strains from Thailand and no cryptic taxon, that included both α‐ and γ‐strains could be demonstrated. However, there was agreement with the 1990 report in that the β‐strain was divisible into two new taxa, with a reduction in heterozygote deficiency, although the genetic distance was not marked (DN= 0.12). Significant genetic distances were found between γ‐strain snails from the type locality and β‐ and γ‐strain samples from North East Thailand (DN= 2.0 and 1.5, respectively). On the basis of genetic distance measures, the Β‐strain is probably a sibling species of N. aperta and the γ‐strain of North East Thailand may also be a separate species. The population at XBF appears to be N. aperta s.s. The public health implications of the findings are discussed, particularly in the context of Mekong river water resource development.

New and revised Lower Devonian Rugosa from Western North America and Tasmania

Anactolasma yukonanumnew genus and species is described from the Pragian of Yukon Territory and is assigned to the Anactolasmatinae, a new subfamily of the Mucophyllidae Hill. Adult stages ofAnactolasmaresemble gerontic stages ofKobehaMerriam, although the genera have distinct ontogenies.Kobehais revised on the basis of new material of its type speciesK. walcottiMerriam. The Papiliophyllinae Stumm are promoted to family status.Kobeha ketophylloidesMerriam, from the Pragian of Nevada, andEndophyllum banksiJell and Hill, from the Pragian of Tasmania, are restudied using new topotypic and other material. A new endophyllid genus namedMurphyphyllumis erected for them, withE. banksias type species.Asarcophyllum ramosumnew genus and species, from the Emsian of Nevada, is made type species of a new cyathophyllid genus.Spongophyllum nevadenseStumm andS. expansumStumm, both also from the Emsian of Nevada, are placed in the new genus, as isAsarcophyllum coronanew genus and species, from the Emsian of Yukon Territory. Biostratigraphic settings, with emphasis on associated corals, brachiopods and conodonts, are given for the new and revised genera and species.

Restudy of the holotype of Operculodinium centrocarpum (Deflandre & Cookson) wall (Dinophyceae) from the Miocene of Australia, and the taxonomy of related species

Operculodinium centrocarpum (Deflandre & Cookson) Wall 1967 is one of the most commonly reported dinoflagellate cysts (dinocysts) in samples from Tertiary and Quaternary deposits. However, at least two distinct dinocyst morphologies are assigned to this species in the literature at the present time. In addition, two separate species have been erected for one of these distinct morphologies. In order to define criteria with which to differentiate these morphologies, holotype and topotype material of O. centrocarpum (Deflandre & Cookson) Wall 1967 and O. echigoense Matsuoka 1983 were restudied and compared with dinocysts of Protoceratium reticulation (Claparéde & Lachmann) Bütschli 1885. Re‐investigation of the holotype and topotype specimens of O. centrocarpum (Deflandre & Cookson) Wall 1967 (=Hystrichosphaeridium centrocarpum Deflandre & Cookson 1955) and O. echigoense Matsuoka 1983 demonstrated that O. echigoense Matsuoka 1983 is a junior synonym of the former. This restudy also confirmed that O. centrocarpum (Deflandre & Cookson) Wall 1967 (= H. centrocarpum Deflandre & Cookson 1955) is morphologically distinct from the Quaternary dinocysts that are widely attributed to this species in the literature. [These Quaternary forms are referred to in this paper as O. centrocarpum (Deflandre & Cookson) sensu Wall 1967.] Viable cysts that are morphologically identical to the Quaternary forms [i.e., O. centrocarpum (Deflandre & Cookson) sensu Wall 1967] have been incubated to produce the motile dinoflagellate P. reticulatum (Claparéde & Lachmann) Bütschli 1885. We propose that, 1) the latter name should be applied only to large, morphologically distinct dinocysts comparable to those originally described from the Miocene of Australia, and 2) the distinctive Quaternary forms [i.e., O. centrocarpum (Deflandre & Cookson) sensu Wall 1967] should be referred to as the cysts of Protoceratium reticulatum (Claparéde & Lachmann) Bütschli 1885.

The type specimens (Middle Cambrian) of the trace fossil Archaeonossa Fenton and Fenton

The original description of Archaeonossa, from beds considered Early Cambrian but now dated as early Middle Cambrian, included at least two unrelated forms of trace fossils. The ichnogenus and the type ichnospecies are redefined, a lectotype is designated, and the type material is reiilustrated, along with topotype material from Yoho National Park, British Columbia. Although this trace fossil was originally attributed to movement by a gastropod, it is concluded, on the basis of restudy, that the type occurrence should not be used as evidence of activity by Gastropoda.

Streptognathodus (Conodonta) succession at the proposed Carboniferous-Permian boundary stratotype section, Aidaralash Creek, Northern Kazakhstan

Streptognathodus Pa elements dominate conodont faunas from the proposed Carboniferous-Permian boundary stratotype at Aidaralash Creek, northern Kazakhstan. The phyletic development of this genus provides the means for subdividing the late Gzhelian to early Sakmarian prodeltaic section (Beds 9-37) into six zones. The preferred Global Stratotype Section and Point (GSSP) for the base of the Permian System is 27 m above the base of Bed 19 at the first occurrence of Streptognathodus isolatus Chernykh, Ritter, and Wardlaw, 1996. This arbitrarily chosen point in the evolutionary continuum of nodose streptognathodids is recognizable in basinal rocks elsewhere in the Ural trough (lower part of Bed 16 at Usolka) as well as the cyclic shelf succession of the American Midcontinent (Glenrock Limestone Member of the Red Eagle Limestone).Eight new species of Streptognathodus occur in the Aidaralash collections: S. bellus n. sp., S. costaeflabellus n. sp., S. flexuosus n. sp., S. glenisteri n. sp., S. longilatus n. sp., S. rectangularis n. sp., S. sigmoidalis n. sp., and S. tenuialveus n. sp. In addition, three morphotypes with limited occurrence (S. sp. A-C) are left in open nomenclature. New collections of topotype material from the nearby Tabantal River section permit clarification of S. barskovi (Kozur).

Evolution in the Early Cretaceous planktonic foraminiferal genus Blefuscuiana

Improved taxonomic understanding provides important information regarding evolution, biostratigraphic distribution and palaeobiogeography. Three species of Early Cretaceous planktonic foraminifera which are of uncertain identity are Globigerina aptica AGALAROVA, Globigerina infracretacea GLAESSNER and Hedbergella aptiana BARTENSTEIN. Examination of topotype and metatype material enables these species to be better defined and their inter-relationship and stratigraphic range to be established. All three species are assigned to the genus Blefuscuiana (Praehedbergellidae). A new species, B. albiana, the descendant of B. aptiana s.s., is also recognised. © 1996 E. Schweizerbart'sche Verlagsbuchhandlung.

Isozyme analysis ofGalaxiasspecies (Teleostei: Galaxiidae) from the Taieri River, South Island, New Zealand: a species complex revealed

We examined genetic differentiation among 23 samples of non-migratory river galaxias from 17 streams in the Taieri River system, South Island, New Zealand. Four major genetic types were found, two of which occur in narrow sympatry in one location. These were compared with topotypical material representing Galaxias anomalus from the Clutha system (Otago) and G. vulgaris from the Waimakariri system (Canterbury) in order to establish identity. Morphological examination of these four major genetic types revealed consistent concomitant differences. The results suggest that there are at least three species of river galaxias in the Taieri system: G. anomalus, G. vulgaris and at least one previously undescribed species. We propose that the genetic structuring and subsequent speciation of this group has been promoted by the absence of the marine juvenile phase that is found in five other members of the genus native to New Zealand. This structuring may be exacerbated by population fragmentation over the last century owing to the negative influence of introduced trout. The phylogenetic diversity within the river system mirrors the diverse flora and invertebrate fauna of the region, and has conservation implications that parallel those resulting from our improved knowledge of the New Zealand herpetofauna through the application of genetic analysis.

Isozyme analysis of Galaxias species (Teleostei: Galaxiidae) from the Taieri River, South Island, New Zealand: a species complex revealed

Abstract We examined genetic differentiation among 23 samples of non-migratory river galaxias from 17 streams in the Taieri River system, South Island, New Zealand. Four major genetic types were found, two of which occur in narrow sympatry in one location. These were compared with topotypical material representing Galaxias anomalus from the Clutha system (Otago) and G. vulgaris from the Waimakariri system (Canterbury) in order to establish identity. Morphological examination of these four major genetic types revealed consistent concomitant differences. The results suggest that there are at least three species of river galaxias in the Taieri system: G. anomalus, G. vulgaris and at least one previously undescribed species. We propose that the genetic structuring and subsequent speciation of this group has been promoted by the absence of the marine juvenile phase that is found in five other members of the genus native to New Zealand. This structuring may be exacerbated by population fragmentation over the last century owing to the negative influence of introduced trout. The phylogenetic diversity within the river system mirrors the diverse flora and invertebrate fauna of the region, and has conservation implications that parallel those resulting from our improved knowledge of the New Zealand herpetofauna through the application of genetic analysis.

Update on the northern Australian Cambrian spongesRankenella, JawonyaandWagima

New material from the Ordian to early Templetonian (early Middle Cambrian) of the Georgina Basin and Daly Basin in northern Australia allows further observations on the anthaspidellid sponge Rankenella mors and the chambered heteractinide sponge Jawonya gurumal respectively. Explanate specimens of R. mors are found to bear closely spaced, rimmed oscules, and the known range of the species is extended from the Ranken Limestone near Soudan to include the Arthur Creek Formation near Ammaroo. Jawonya gurumal from the Tindall Limestone near Claravale is better preserved than type and topotype material, and demonstrates that the genus is two-walled, and not one-walled as originally described. Furthermore, exopore architecture is much more complex than previously envisaged. The co-occurring related genus Wagima is also considered to be two-walled. Rankenella in the Ranken Limestone flourished in a low-energy, shallow subtidal marine environment subject to episodic higher-energy events that generated ooid shoals and flat-pebble conglomerates. In the Arthur Creek Formation the genus occurs in anaerobic calcimudstone deposited in a low-energy shelf area of limited circulation. Jawonya and Wagima are elements of a spongemollusc association in open shelf onkoid limestones of the Tindall Limestone. Rankenella is among the oldest known intact demosponges. Spiculation in Jawonya and Wagima suggests that the history of the heteractinide astraeospongiid-wewokellid lineage was one of Ordovician reduction or loss of original polyactines in favour of octactines, followed by polyactine reinstatement in the Carboniferous.

The trace fossilFucoides circinatusBrongniart, 1828, from its type area, Västergötland, Sweden

Abstract Topotypical material of the trace fossil Fucoides circinatus Brongniart, 1828, from the Lower Cambrian of south‐central Sweden, consists of vertically oriented spreite burrows. It follows that Fucoides circinatus should not, contrary to common practise, be used for burrows of Zoophycos type and is not a contending type species of Zoophycos Massalongo, 1855. Furthermore, Alectorurus Schimper, 1869, of which Fucoides circinatus is type species by monotypy, should not be included as a synonym of Zoophycos. The type species of Phycodes Richter, 1850, is P. circinatum Richter, 1853, which is distinct from its supposed synonym F. circinatus. Jensen, S. & Bergstrom, J., 1995: The trace fossil Fucoides circinatus Brongniart, 1828, from its type area, Vastergotland, Sweden. GFF, Vol. 117 (Pt. 4, December), pp. 207–210. Stockholm. ISSN 1103–5897.

Prionospio steenstrupi,P. fallaxandP. dubia(Polychaeta, Spionidae): Re-evaluation of identity and status

Abstract Within the genus Prionospio Malmgren, 1867, the ‘steenstrupi-group’ (including the type species P. steenstrupi Malmgren, 1867) is characterised by the possession of four pairs of branchiae, of which the first and fourth pairs are pinnate. Unfortunately the identities of P. steenstrupi and the other northeastern Atlantic species (P. fallax Soderstrom, 1920 and P. dubia Maciolek, 1985) have been much confused. All have been variously misinterpreted or synonymised in the literature. An examination of type, topotypic and other European material indicates that all three species are valid and readily distinguished. Each species is described and figured in detail, and their geographical distributions reassessed. The usefulness of the characters employed to distinguish these and other species is discussed. It is recommended that assessments of intraspecific variability be included in all species descriptions.

An endemic brachiopod fauna from the Middle Ordovician of North Wales

AbstractThe brachiopod fauna from the base of the Bryn Siltstone Formation (middle Caradoc), Glyn Ceiriog, north Berwyn Hills, North Wales, consists of Platystrophia elevata sp. nov., Salopia abbreviata sp. nov., Kiaeromena ungula (M'Coy) and unnamed species of Hesperorthis?, Reuschella and Bicuspina. The dominant brachiopod species, Leptaena (Leptagonia) ungula M'Coy, 1851 is revised and its current assignment to the leptaenid genus Kiaeromena confirmed, based on the examination of type and topotype material from the Bryn Siltstone Formation. Kiaeromena spjeldnaesi nom. nov. is proposed to replace the junior homonym K. ungula Spjeldnæs, 1957 for that congeneric species from the upper Caradoc rocks of Ringerike, southern Norway. The fauna as a whole is unusual within the context of the Anglo‐Welsh province and may represent a rare preserved nearshore community.

The Famennian brachiopod Zilimia polonica (Gürich) and its palaeoenvironmental significance

The rhynchonellid brachiopod Zilimia polonica (Gürich, 1896) is described, based on new topotypic material. The neotype of the species comes from the lower Famennian Upper crepida zone. The crinoidal marls containing Z. polonica and an associated brachiopod assemblage are regarded as cohesive debris flow material, redeposited from shallow-water into an adjacent intrashelf basin. The distribution of Z. polonica suggests that it populated open-marine, moderately shallow environments within the swell and basin topography developed along the margins of the East European and the eastern cratons.