Showing papers in "Zoologica Scripta in 2004"

Phylogenetic analysis of the Balanidae (Cirripedia, Balanomorpha)

Abstract: Pitombo, F. B. (2004). Phylogenetic analysis of the Balanidae (Cirripedia, Balanomorpha). —Zoologica Scripta, 33, 261–276. Our concept of the family Balanidae has undergone many changes since it was originally proposed, and it is now a more circumscribed group. Nevertheless, there are no studies of the phylogenetic relationships among its currently recognized subfamilies: Balaninae, Concavinae and Megabalaninae. Here, a new suite of characters is presented, thus allowing a better evaluation of the systematics of the group. A fourth subfamily, the Amphibalaninae, is proposed to accommodate Amphibalanus gen. nov. and its allies, and Perforatus gen. nov. is proposed and transferred to the Concavinae (all formerly Balaninae). A phylogenetic analysis of the Balanidae was performed. The results showed that Balaninae is basal to the other three subfamilies as well as Concavinae, and Megabalaninae forms a monophyletic group. Hypotheses of character evolution are presented and discussed. As a general pattern, Concavinae and Megabalaninae have the most derived structures, such as the extension of the lateral margin of the sheath, inner face of radii with a longitudinal abutment, basis multilayered (vesicular) and a spur with infolded margins.

18S rDNA phylogeny of Clitellata (Annelida)

Abstract: The phylogeny of Clitellata was analysed using 18S rDNA sequences of a selection of species representing Hirudinida, Acanthobdellida, Branchiobdellida and 10 oligochaetous families. Eleven new 18S sequences of Capilloventridae (one), Haplotaxidae (one), Propappidae (one), Enchytraeidae (two), Lumbricidae (one), Almidae (one), Megascolecidae (two), Lumbriculidae (one), and Phreodrilidae (one) are reported and aligned together with corresponding sequences of 28 previously studied clitellate taxa. Twelve polychaete species were used as an outgroup. The analysis supports an earlier hypothesis based on morphological features that Capilloventridae represents a basal clade of Clitellata; in the 18S tree it shows a sister-group relationship to all other clitellates. The remaining clitellate taxa form a basal dichotomy, one clade containing Tubificidae (including the former ‘Naididae’), Phreodrilidae, Haplotaxidae, and Propappidae, the other clade with two subgroups: (1) Lumbriculidae together with all leech-like taxa (Acanthobdellida, Branchiobdellida and Hirudinida), and (2) Enchytraeidae together with a monophyletic group of all earthworms included in the study (Lumbricidae, Almidae and Megascolecidae). These earthworms are members of the taxon Crassiclitellata, the monophyly of which is thus supported by the data. The tree also shows support for the hypothesis that the first clitellates were aquatic. The position of the single species representing Haplotaxidae is not as basal as could have been expected from earlier morphology-based conclusions about the ancestral status of this family. However, if Haplotaxidae is indeed a paraphyletic assemblage of relict taxa, a higher number of representatives will be needed to resolve its exact relationships with the other clitellates.

Phylogeny of Recent Canidae (Mammalia, Carnivora): relative reliability and utility of morphological and molecular datasets

Abstract: Zrzavý, J. & Řicankova, V. (2004). Phylogeny of Recent Canidae (Mammalia, Carnivora): relative reliability and utility of morphological and molecular datasets. — Zoologica Scripta, 33, 311–333. Phylogenetic relationships within the Canidae are examined, based on three genes (cytb, COI, COII) and 188 morphological, developmental, behavioural and cytogenetic characters. Both separate and combined phylogenetic analyses were performed. To inspect the phylogenetic ‘behaviour’ of individual taxa, basic phylogenetic analysis was followed by experimental cladistic analyses based on different data-partition combinations and taxon-removal analyses. The following phylogeny of the Recent Canidae is preferred: (1) Urocyon is the most basal canid; (2) Vulpes is a monophyletic genus (including Fennecus and Alopex); (3) the doglike canids (DC) form a clade (=Dusicyon + Pseudalopex + Lycalopex + Cerdocyon + Atelocynus + Chrysocyon + Speothos + Lycaon + Cuon + Canis), split into two subclades, South American and Afro-Holarctic, with uncertain position of the Chrysocyon + Speothos subclade; (4) Canis is paraphyletic due to the position of Lycaon and Cuon. Otocyon and Nyctereutes are the most problematic canid genera, causing an unresolved branching pattern of Otocyon, Vulpes, Nyctereutes and DC clades. Reclassification of the two basal species of ‘Canis’ into separate genera is proposed (Schaeffia for ‘C.’ adustus, Lupulella for ‘C.’ mesomelas). Although the morphological dataset ranked poorly in both separate and simultaneous analyses (measured by number of minimum-length topologies, relative number of resolved nodes in the strict consensus of all minimum-length topologies, consistency and retention indices, nodal dataset influence, and number of extra steps required by the data partition to reach the topology of the combined tree), the morphological synapomorphies represent nearly one quarter of all synapomorphies in the combined tree. Among the hidden morphological support of the combined tree the developmental and behavioural characters are conspicuously abundant.

Phylogeny, evolutionary history and taxonomy of the Mustelidae based on sequences of the cytochrome b gene and a complex repetitive flanking region

Abstract: The Mustelidae is a diverse family of carnivores which includes weasels, polecats, mink, tayra, martens, otters, badgers and, according to some authors, skunks. Evolutionary relationships within the family are under debate at a number of different taxonomic levels, and incongruencies between molecular and morphological results are important. We analysed a total of 241 cytochrome b (cyt b) gene sequences and 33 sequences of a complex repetitive flanking region from 33 different species to compile an extensive molecular phylogeny for the Mustelidae. We analysed these sequences and constructed phylogenetic trees using Bayesian and neighbor-joining methods that are evaluated to propose changes to the taxonomy of the family. The peripheral position of skunks in phylogenetic trees based on both loci suggests that they should be considered a separate family, Mephitidae. The subfamily Melinae is the basal group within the Mustelidae and trees based on the cyt b gene suggest that the American badger, Taxidea taxus, should be considered a separate monotypic subfamily, Taxidiinae. Otters classified within the genera Lutra, Amblonyx and Aonyx are grouped within the same clade in cyt b and combined partial cyt b and flanking region trees and show reduced levels of inter specific divergence, suggesting that they could be classified together under a single genus, Lutra. The Bayesian tree based on combined data from both loci supports the idea that subfamily Mustelinae is paraphyletic, as otters (subfamily Lutrinae) are included in this subfamily. Low levels of genetic divergence among European polecat, Mustela putorius, steppe polecat, Mustela eversmannii, and European mink, Mustela lutreola, suggest that these species could be considered subspecies within a single species, Mustela putorius. Our results are consistent with a rapid diversification of mustelid lineages in six different radiation episodes identified since the Early Eocene, the oldest events being the separation of subfamilies and the split of marten (Martes, Gulo) and weasel (Mustela) lineages in the Early Middle Miocene. The separation of New World from Old World lineages and the split of the remaining genera are estimated to have occurred in Late Miocene. The most recent events have been the differentiation of species within genera and this probably occurred in four radiation episodes at the end of Late Miocene, Early Pliocene, Late Pliocene and Pleistocene epochs.

Phylogenetic relationships of basal hexapods among the mandibulate arthropods: a cladistic analysis based on comparative morphological characters

Abstract: In this paper we propose a reappraisal of the relationships between the basal hexapod lineages (the former ‘apterygote’ insects) and the other major groups of mandibulate arthropods. It results from a cladistic analysis including 72 characters based on external morphology, internal anatomy and development. Detailed comments are provided on the various characters used and the scoring of their states. The 35 terminal taxa include 12 hexapods (9 of which are basal ‘apterygote’ representatives), 7 myriapods, 13 crustaceans, and 3 chelicerates taken as outgroups. The results of our analyses are discussed in detail for each of the taxonomic groupings, and compared with those recently obtained by other authors using different approaches based on morphological, palaeontological, developmental or molecular sequence data. Our results support the monophyly of the Mandibulata, Crustacea, Atelocerata (Tracheata) and Hexapoda, but the assemblage of Myriapoda appears poorly supported. A close relationship between Crustacea and Hexapoda, as hypothesized by several authors, is not found in any of our analyses. Within Hexapoda, the Protura and the Collembola appear as independent clades, whereas the two unresolved dipluran taxa are grouped with the monophyletic Ectognatha (Archaeognatha, Zygentoma and Pterygota).

Genetic exchange between anchialine cave populations by means of larval dispersal: the case of a new gastropod species Neritilia cavernicola

Abstract: Despite being limited to caves, many anchialine taxa have disjunct insular distributions, which raises questions about their origins and colonization history. This study deals with the new gastropod Neritilia cavernicola sp. n. (Neritopsina: Neritiliidae) from anchialine caves on two islands in the Philippines that are separated by the deep Bohol Strait and situated 200 km apart along the coastline of Cebu Island. Neritilia cavernicola is an obligate stygobiont and most closely resembles Neritilia littoralis, which lives in interstitial waters of the Nansei-shoto Islands, Japan. Its eggs and larval shells are identical to those of other Neritilia species, despite their different adult habitats. This suggests a marine planktotrophic phase (as occurs in amphidromous riverine species of Neritilia), and consequent migration between islands via ocean currents. Here we present the first genetic structure for anchialine cave organisms; comparisons of 1276 bp sequences from mitochondrial cytochrome oxidase I show no evidence of genetic isolation between the islands. All individuals evidently are part of a panmictic population and the low vagility of adults and their seemingly isolated cave habitats do not limit gene flow in N. cavernicola. This migration model, based on marine larval dispersal, may be widely applicable to anchialine stygobites with insular distributions, as many such organisms (including shrimps, crabs and fishes) are phylogenetically allied to amphidromous species.

The freshwater turtle genus Mauremys (Testudines, Geoemydidae) — a textbook example of an east–west disjunction or a taxonomic misconcept?

Abstract: Barth, D. Bernhard, D. Fritzsch, G. & Fritz, U. (2004): The freshwater turtle genus Mauremys— a textbook example of an east–west disjunction or a taxonomic misconcept? —Zoologica Scripta, 33, 213–221. We compare 1036 bp of the mitochondrial cytochrome b gene (cyt b) from all six Mauremys species with 16 other taxa, representing both currently recognized subfamilies of the Geoemydidae (Geoemydinae and Batagurinae) to contribute a comprehensive dataset towards resolving the conflicting Mauremys taxonomy and phylogeography. Mauremys, a representative of the Geoemydinae, is thought to be an example of a taxon with an east–west disjunction due to Pleistocene glacial extinction, with species occurring in the western Palearctic and species in the eastern Palearctic and Oriental regions. Our results contradict this traditional zoogeographical scheme and the current taxonomy of the Geoemydidae. Mauremys is paraphyletic with respect to two East Asian genera belonging to the Batagurinae: Chinemys and Ocadia. Therefore, Mauremys, as currently understood, clearly represents a taxonomic misconcept. Mauremys+Chinemys+Ocadia contains four well supported clades, two of which —M. japonica+Chinemys+Ocadia and M. annamensis+M. mutica— are confined to eastern Asia. The other two —M. caspica+M. rivulata and M. leprosa— occur in the western Palearctic. Mauremys leprosa may represent an ancient lineage which differentiated before the split between the other western and eastern species occurred. The patchy distribution of the four clades is likely the result of several ancient radiations rather than of a Pleistocene extinction. The sister-group of Mauremys+Chinemys+Ocadia is Cuora, a morphologically highly specialized genus with a complicated shell hinging mechanism.

Molecular and morphological evidence for multiple species within Paranoplocephala omphalodes (Cestoda, Anoplocephalidae) in Microtus voles (Arvicolinae)

Abstract: Haukisalmi, V., Wickstrom, L. M., Henttonen, H., Hantula, J. & Gubanyi, A. (2004). Molecular and morphological evidence for multiple species within Paranoplocephala omphalodes (Cestoda, Anoplocephalidae) in Microtus voles (Arvicolinae). —Zoologica Scripta, 33, 277–290. The present study was designed to test the hypothesis that the anoplocephalid cestode Paranoplocephala omphalodes (Hermann, 1783), a Holarctic parasite of Microtus voles, is a complex of host-specific species, rather than a single host-generalist species, using uni- and multivariate morphometrics and DNA sequence data from the mitochondrial cytochrome oxidase I gene. The phylogenetic methods applied to the mtDNA sequence data showed consistently that the cestodes morphologically recognizable as P. omphalodes include four well-supported monophyletic groups, representing at least three distinct, largely host-specific species. Multivariate morphometrics (discriminant analysis) successfully distinguished the four main mtDNA clades of P. omphalodes-like cestodes. The true P. omphalodes is shown to be a parasite of Microtus arvalis, M. agrestis and Clethrionomys glareolus in Europe. Microtus oeconomus harbours two host-specific, allopatric and possibly conspecific clades, one with a Holarctic and another with an (eastern) Beringian (Alaskan) distribution. The eastern Beringian endemic M. miurus is also parasitized with a host-specific, morphologically divergent species of Paranoplocephala. The cestode clades recognized in M. oeconomus and M. miurus represent 2–3 undescribed species. Molecular phylogenetic analyses supported the monophyly of the ‘northern clade’ of Paranoplocephala spp., an assemblage including P. kalelai from Clethrionomys spp., P. macrocephala from Microtus spp. and all clades of P. omphalodes-like cestodes except those representing the true P. omphalodes from Europe. The intra- and interspecific phylogeny within the northern clade is compared tentatively with the known evolutionary history of the hosts.

Phylogeny of Australasian venomous snakes (Colubroidea, Elapidae, Hydrophiinae) based on phenotypic and molecular evidence

Abstract: Scanlon, John D. & Lee, Michael S. Y. (2004). Phylogeny of Australasian venomous snakes (Colubroidea, Elapidae, Hydrophiinae) based on phenotypic and molecular evidence. — Zoologica Scripta, 33, 335–366. Phylogenetic relationships among Hydrophiinae (Australasian and marine elapid snakes) are inferred using 87 characters from external, skeletal, hemipenial and internal anatomy, ecology, and chromosomes as well as available sequences of two mitochondrial genes (cytochrome b and 16S rRNA). Parsimony analysis of the combined data retrieves many widely accepted clades; while observed bootstrap or branch (Bremer) support for these is often weak, most have never been corroborated previously by a rigorous numerical analysis. Sea kraits (Laticauda) and Solomon Islands elapids are basal to the remaining hydrophiines (Australian terrestrial forms and hydrophiin sea snakes). The latter clade includes three main lineages: a large-bodied oviparous lineage, a small-bodied oviparous lineage, and a viviparous lineage (which also includes the hydrophiin sea snakes, strongly reaffirmed as monophyletic). While the Solomons retain a relictual fauna, New Guinea has less endemism and has been invaded multiple times by Australian lineages, so there is no clear ‘stepping stone’ pattern supporting a northern (Asian, rather than Gondwanan) biogeographical origin.

Phylogenetic relationships of fish leeches (Hirudinea, Piscicolidae) based on mitochondrial DNA sequences and morphological data

Abstract: Utevsky, S. Y. & Trontelj, P. (2004). Phylogenetic relationships of fish leeches (Hirudinea, Piscicolidae) based on mitochondrial DNA sequences and morphological data. — Zoologica Scripta, 33, 375–385. Phylogenetic relationships of fish leeches (Piscicolidae) were deduced from combined mitochondrial DNA sequences of 12S rDNA and COI genes using Bayesian inference and Maximum Likelihood, as well as from a combined molecular-morphological data matrix using Maximum Parsimony. Monophyly of the family was confirmed. The traditional subdivision into three subfamilies, the Platybdellinae, Pontobdellinae, and Piscicolinae, received weak support, but was not challenged by alternative groupings. In contrast to prior classifications, a basal split emerged between the Pontobdellinae on the one hand, and the Platybdellinae and Piscicolinae on the other. The complex coelomic system of pontobdellines is viewed as plesiomorphic, and independent reductions of the coelom in other fish leech groups are hypothesized. According to the inferred phylogeny, seawater was the primary habitat of fish leeches. Eurasian freshwaters were colonized by a species-rich freshwater clade (genera Piscicola, Baicalobdella, Cystobranchus and Caspiobdella) and, probably independently, by the Asian genus Limnotrachelobdella.

Morphological and molecular taxonomic analysis of the Encarsia meritoria species-complex (Hymenoptera, Aphelinidae), parasitoids of whiteflies (Hemiptera, Aleyrodidae) of economic importance.

Abstract: Five closely related species of Encarsia, belonging to the luteola species-group, are analysed taxonomically using morphological and molecular techniques. Four of these belong to the meritoria-complex, and all species have a complicated taxonomic history of repeated misidentification and confusion. Morphological analysis is focused on morphometric characters of the female antennae using principal component and canonical discriminant analyses. DNA sequence data for the D2 region of 28S nuclear ribosomal genes were obtained for 13 populations of the luteola-group, with a further seven population sequences being obtained from GenBank. The combination of morphological and molecular study enables us to resolve the complex to a large extent, and to correct the previous confusion surrounding this group of species. Two species —E. californica and E. dispersa— are described as new. A proposal to place E. brasiliensis (Hempel) in synonymy with E. hispida DeSantis (the more recently described taxon) is presented. A lectotype is designated for E. haitiensis Dozier. Diagnoses or descriptions, and illustrations, of all included species are provided to facilitate the identification of females belonging to this complex. All known data concerning hosts and geographical distribution are presented.

The scientific status of metazoan cladistics: why current research practice must change

Abstract: Jenner, R. A. (2004). The scientific status of metazoan cladistics: why current research practice must change. —Zoologica Scripta, 33, 293–310. Metazoan phylogenetics is bustling with activity. The use of comprehensive morphological data sets in recent phylogenetic analyses of the Metazoa indicates that morphological evidence continues to play a key role in the reconstruction of metazoan deep history. In this paper I review the scientific status of morphological metazoan cladistics from the perspective of cladistic research cycles. Each research cycle consists of three main steps: (1) the compilation of a data matrix (2) the simultaneous evaluation of all possible cladograms in a character congruence test, and (3) the assessment of the relationship between evidence and hypothesis after finding the optimal tree. I identify a striking discrepancy between the sophistication of the analysis of given data sets (Step 2), and their compilation and the interpretation of the results (Steps 1 and 3). The latter two steps deserve far greater attention than is current practice. Uncritical and nonexplicit character selection, character coding, and character scoring seriously compromise Step 1. Careful comparative morphological study prior to data matrix construction is necessary to remedy this problem in future cladistic analyses. Step 2 is the locus of most recent advances in metazoan cladistics through the increasing availability of computing power, and the development of increasingly efficient phylogenetic software that allows analysis of large data sets. Failure to identify problems and errors generated in Step 1 of the research cycle is testament to the general failure of Step 3. Consequently, recent progress in metazoan cladistics is primarily analytical, while the only empirical anchor of the discipline receives surprisingly little attention. Not surprisingly, the first generation of modern metazoan phylogeneticists used computers principally as a relatively quick and easy means to generate abundant phylogenies from morphological data. The next phase should build on this foundation by critically testing these alternative hypotheses by a thorough qualitative reassessment and elaboration of morphological data matrices, and a more critical approach to data selection. A rigorous research program for metazoan cladistics can only be established when the cladistic research cycle is properly completed, and when subsequent research cycles are effectively linked to previous efforts.

Cladistic revision of talitroidean amphipods (Crustacea, Gammaridea), with a proposal of a new classification

Abstract: This paper reports the results of a cladistic analysis of the Talitroidea s.l., which includes about 400 species, in 96 genera distributed in 10 families. The analysis was performed using paup and was based on a character matrix of 34 terminal taxa and 43 morphological characters. Four most parsimonious trees were obtained with 175 steps (CI = 0.617, RI = 0.736). A strict consensus tree was calculated and the following general conclusions were reached. The superfamily Talitroidea is elevated herein as infraorder Talitrida, which is subdivided into three main branches: a small clade formed by Kuria and Micropythia (the Kurioidea), and two larger groups maintained as distinct superfamilies (Phliantoidea, including six families, and Talitroidea s.s., including four). Within the Talitroidea s.s., the following taxonomic changes are proposed: Hyalellidae and Najnidae are synonymized with Dogielinotidae, and treated as subfamilies; a new family rank is proposed for the Chiltoniinae.

Systematics, phylogeny and biogeography of Anterastes (Orthoptera, Tettigoniidae, Tettigoniinae): evolution within a refugium

Abstract: Ciplak, B. (2004). Systematics, phylogeny and biogeography of Anterastes (Orthoptera, Tettigoniidae, Tettigoniinae): evolution within a refugium. —Zoologica Scripta, 33, 19–44. The genus Anterastes, distributed in southeastern Europe and the western part of Anatolia, is revised based on previous materials and numerous specimens collected from new localities. A key to all species is presented. Two new species, A. antitauricus sp. n. and A. ucari sp. n. are described. Anterastes akdaghensis Ramme is placed in synonymy with A. babadaghi Uvarov. Cladistic analysis confirmed the monophyly of Anterastes. The relationships among the species of Anterastes are: A. uludaghensis + ((A. serbicus + A. burri + A. antitauricus sp. n.) + (A. anatolicus + A. tolunayi + (A. niger + (A. babadaghi + A. turcicus + A. ucari sp. n.)))). The biogeography of the genus shows a correlation with its phylogeny. It is assumed that the genus arose from an ancestral stock in northwestern Anatolia in the Pliocene and the later range of this ancestral population expanded and contracted under the effects of the ice ages (glacial and interglacial periods, respectively). It is postulated that speciation within the genus, suggested by phylogenetic analysis, might have occurred when the range of ancestral populations expanded during glacial periods and contracted in subsequent warm periods. The present species may be the product of relict populations remaining in refugia at higher altitudes with alpine or subalpine vegetation in southern Anatolia.

First molecular evidence for reassessing phylogenetic affinities between genets ( Genetta ) and the enigmatic genet-like taxa Osbornictis , Poiana and Prionodon (Carnivora, Viverridae)

Abstract: Gaubert, P., Tranier, M., Delmas, A.-S., Colyn, M. & Veron, G. (2004). First molecular evidence for reassessing phylogenetic affinities between genets (Genetta) and the enigmatic genet-like taxa Osbornictis, Poiana and Prionodon (Carnivora, Viverridae). —Zoologica Scripta, 33, 117–129. The subfamily Viverrinae is a composite group of carnivores comprising the large and plantigrade terrestrial civets (Civettictis, Viverricula and Viverra) and the slender and generally more arboreal genets and genet-like taxa (Genetta, Prionodon, Poiana, Osbornictis), both having Asiatic and African representatives. The problematic phylogenetic relationships between genets and genet-like taxa are addressed for the first time from a molecular perspective through complete cytochrome b gene sequences. We used a large taxonomic sample set including some very rare and crucial species such as Osbornictis piscivora, Poiana richardsonii (museum specimen material) and Genetta johnstoni. The results from parsimony, distance and maximum likelihood analyses do not support the monophyly of the Viverrinae and contradict previous morphological hypotheses. The Asiatic linsangs (Prionodon spp.) are excluded from the Viverrinae and represent either a basal Feliformia or Viverridae. The other genet-like taxa constitute a strongly supported monophyletic African group, in which the African linsang (represented by Poiana richardsonii) is a sister group to the genets. The aquatic genet Osbornictis piscivora is included within the latter clade, and the genus Osbornictis should be considered a junior synonym of Genetta. African and Asiatic terrestrial civets are monophyletic, but their phylogenetic affinities with the genet-like clade are inconclusive using our data set. On the basis of our molecular results, morphological convergences and adaptations to peculiar habitats and ways of life within genets and genet-like taxa are discussed.

A new species of the genus Microbunodon (Anthracotheriidae, Artiodactyla) from the Miocene of Pakistan: genus revision, phylogenetic relationships and palaeobiogeography

Abstract: Lihoreau, F., Blondel, C., Barry, J. & Brunet, M. (2004). A new species of the genus Microbunodon (Anthracotheriidae, Artiodactyla) from the Miocene of Pakistan: genus revision, phylogenetic relationships and palaeobiogeography. — Zoologica Scripta, 33, 97–115. New unpublished remains of small Anthracotheriinae are described. First, materials from the upper Oligocene (MP 30) locality of La Milloque, southwest France, permit a review of the species Microbunodon minimum. Thereafter, fossils from the middle and late Miocene of the Potwar Plateau, Pakistan are attributed to the European genus Microbunodon. Microbunodon milaensis sp. n. from the Nagri Formation (between 10.3 and 9.2 Ma), Pakistan, is described and the species M. silistrensis from the Lower Manchar Formation (between 16 and 15 Ma) and from the Chinji Formation (between 12.7 and 11.5 Ma), Pakistan, is reviewed. The new species represents the last occurrence of the subfamily Anthracotheriinae, around 9.3 Ma. Similar materials from the Bugti and Siwalik Hills were previously considered as a small Anthracotherium. Comparisons with M. minimum from the European late Oligocene lead to a complete revision of the genus and permit definition of a new set of characters, which separate Microbunodon from Anthracotherium. A cladistic analysis reconsiders phylogenetic relationships among Anthracotheriinae, separating an Anthracothema–Anthracotherium clade and an Anthracokeryx–Microbunodon clade. Microbunodon appears to stem from the Asian late Eocene–lower Oligocene genus Anthracokeryx. These results imply a new distribution of the genus Microbunodon showing exchanges between Europe and Asia during the late Oligocene and probably the lower Miocene.

Data incongruence and the problem of avian louse phylogeny

Abstract: Smith, V. S., Page, R. D. M. & Johnson, K. P. (2004). Data incongruence and the problem of avian louse phylogeny. —Zoologica Scripta, 33, 239 –259. Recent studies based on different types of data (i.e. morphological and molecular) have supported conflicting phylogenies for the genera of avian feather lice (Ischnocera: Phthiraptera). We analyse new and published data from morphology and from mitochondrial (12S rRNA and COI) and nuclear (EF1-α) genes to explore the sources of this incongruence and explain these conflicts. Character convergence, multiple substitutions at high divergences, and ancient radiation over a short period of time have contributed to the problem of resolving louse phylogeny with the data currently available. We show that apparent incongruence between the molecular datasets is largely attributable to rate variation and nonstationarity of base composition. In contrast, highly significant character incongruence leads to topological incongruence between the molecular and morphological data. We consider ways in which biases in the sequence data could be misleading, using several maximum likelihood models and LogDet corrections. The hierarchical structure of the data is explored using likelihood mapping and SplitsTree methods. Ultimately, we concede there is strong discordance between the molecular and morphological data and apply the conditional combination approach in this case. We conclude that higher level phylogenetic relationships within avian Ischnocera remain extremely problematic. However, consensus between datasets is beginning to converge on a stable phylogeny for avian lice, at and below the familial rank.

Molecular taxonomy and phylogeny of the 'living fossil' lineages triops and lepidurus (branchiopoda: notostraca)

Abstract: Mantovani B. Cesari M. & Scanabissi F. (2004). Molecular taxonomy and phylogeny of the ‘living fossil’ lineages Triops and Lepidurus (Branchiopoda: Notostraca). —Zoologica Scripta, 33, 367–374. European Triops cancriformis and Lepidurus apus were analysed for 12S and 16S mitochondrial genes and compared to North American and Japanese taxa. There are no cryptic species among European T. cancriformis populations, which are highly homogeneous in comparison to conspecific Japanese samples. T. cancriformis differs from congeneric taxa all over its range, which can be explained by its antiquity. In contrast, the parapatric subspecies L. apus apus and L. apus lubbocki are morphologically conserved and differ substantially at the mtDNA level. The genetic distance values between them are of the same order of magnitude as those observed between American Lepidurus species. Their subspecific status therefore requires further analysis. L. apus apus is more closely related to a L. arcticus sample from Iceland than to L. apus lubbocki. It is also related to a Canadian L. couesii population. Further analyses of populations from the whole range of L. arcticus and the European range of L. couesii are needed to understand the relationships among these notostracan taxa. When considering the two genera, it is clear that Lepidurus is a well supported monophyletic unit, while Triops is polyphyletic, embodying very divergent taxa.

Hippopotamyrus ansorgii species complex in the Upper Zambezi River System with a description of a new species, H. szaboi (Mormyridae)

Abstract: Specimens referable to Hippopotamyrus ansorgii sampled from the Upper Zambezi River System within Caprivi (Namibia) represent a complex of three species, two of which coexist in the Upper Zambezi River, and a third that inhabits a nearby river, the Kwando, with which the Zambezi has been connected during periods of flooding. All three are indistinguishable in terms of their general appearance, but differ consistently in electric organ discharges (EOD), morphology, and molecular genetic characters. All phenotypes display a monopolar, headpositive EOD pulse with specific post- or prepotentials. For H. ansorgii from the Zambezi River (HaZ), pulse duration is less than 0.5 ms (down to 0.205 ms; N = 34); for the syntopic H. szaboi sp. n., it is greater than 0.6 ms (up to 1.8 ms at 10% peak amplitude; N = 19). The parapatric phenotype of H. ansorgii from the Kwando River (HaK) has pulses shorter than 0.215 ms (down to 0.105 ms; N = 36). All three members of the species complex may be distinguished from each other by 7 − 9 anatomical characters, analysed by MANOVA. Based on 22 enzymes and proteins studied, the moderate to high Wright’s fixation index and the significant (P < 0.05) allele differentiation between EOD phenotypes provide additional evidence for incipient speciation. Pairwise analyses of the three different phenotypes showed the two parapatric species of H. ansorgii grouped together, and distinguishable from individuals of H. szaboi . Analyses of the mitochondrial cytochrome b gene revealed that all specimens which were attributed to H. szaboi form a well-supported monophyletic basal clade (bootstrap support 73% or 82%). The genetic distances (uncorrected p distances) between H. szaboi and the two species of H. ansorgii are between 0.6% and 1.7%. Within the derived H. ansorgii clade some phylogeographical differentiation can be seen for fishes from the Zambezi and Kwando Rivers, but the respective groups are not consistent or supported by significant bootstrap values. The question of which of the two parapatric morphological and EOD phenotypes of H. ansorgii recognized in the present paper represents H. ansorgii (Boulenger, 1905) cannot be resolved at present because of the paucity and unclear origin of the historical type material.

Phylogeny of the Aplysiidae (Gastropoda, Opisthobranchia) with new aspects of the evolution of seahares

Abstract: The phylogeny of the Aplysiidae is investigated, based on 37 morphological and histological characters polarized a priori by outgroup comparison. The Aplysiidae represents a monophyletic taxon comprising two distinct clades: Aplysiinae and Dolabellinae + Dolabriferinae + Notarchinae. The traditional classification of Longicommissurata (Aplysiinae + Dolabellinae) is no longer valid since the Longicommissurata are paraphyletic. However, the Brevicommissurata (Dolabriferinae + Notarchinae) form a monophyletic group. Within the Dolabriferinae two sister-groups can be distinguished: Dolabrifera and Phyllaplysia+Petalifera petalifera. The genus Petalifera is paraphyletic. Based on the present phylogeny, several aspects of the evolution of the Aplysiidae are discussed.

Non‐monophyly of the avian genus Seicercus (Aves: Sylviidae) revealed by mitochondrial DNA

Abstract: The phylogeny of all species and nearly all subspecies of Seicercus and representatives of all subgenera in Phylloscopus was estimated based on two mitochondrial genes. According to the gene tree, and supported by non-molecular data, Seicercus belongs in three separate clades. Two of these include only taxa currently classified as Seicercus, while the third comprises S. xanthoschistos and P. occipitalis. These results suggest that both Seicercus and Phylloscopus are paraphyletic. The gene tree suggests two more cases of non-monophyly: (1) the ‘S. burkii complex’ is separated into two different clades, one of which also includes S. affinis and S. poliogenys; (2) two populations of S. affinis intermedius are more closely related to S. affinis ocularis than to a third population of intermedius. A recent proposal to split the ‘S. burkii complex’ into six species is corroborated, as is the recognition of the taxon cognitus as a colour morph of S. affinis intermedius. Our study also revealed unexpectedly large genetic divergences between three different populations of the monotypic S. poliogenys, indicating the presence of cryptic species. Our results underscore the importance of dense sampling at the specific and infraspecific levels in intrageneric phylogenetic studies.

Genetic vs. morphological differentiation of Old World buzzards (genus Buteo, Accipitridae)

Abstract: Kruckenhauser, L., Haring, E., Pinsker, W., Riesing, M. J., Winkler, H., Wink, M. & Gamauf, A. (2004). Genetic vs. morphological differentiation of Old World buzzards (genus Buteo, Accipitridae). — Zoologica Scripta, 33, 197–211. Here, a comprehensive phylogenetic investigation of Old World buzzards of the buteo–vulpinus complex and related taxa using morphological and genetic markers is presented. The morphometric analysis proved useful to discriminate taxa. Nevertheless, phylogenetic relationships cannot be resolved with these characters. Sequence comparisons between the control region and the pseudo-control region revealed that the latter is the most variable section of the mitochondrial genome. Consequently it was used as a marker sequence. In the genetic analysis, almost no sequence variability was found among taxa comprising the buteo–vulpinus complex as well as Buteo rufinus and Buteo oreophilus, suggesting gene flow and/or incomplete lineage sorting. Thus, rapid morphological differentiation in adaptation to different environments was not accompanied by genetic differentiation of the mitochondrial genomes of these taxa. In contrast, the East Palearctic taxa are well differentiated genetically. The ‘superspecies’ concept and taxonomic consequences of our results are discussed.

Evolution of the nervous system in Paraphanostoma (Acoela)

Abstract: Raikova, O. I., Reuter, M., Gustafsson, M. K. S., Maule, A. G., Halton, D. W. & Jondelius, U. (2004). Evolution of the nervous system in Paraphanostoma (Acoela). —Zoologica Scripta, 33, 71–88. According to recent molecular studies, the Acoela are the earliest extant bilaterian group. Their nervous system displays a striking variety of patterns. The aim of the present investigation was to study the variability of the nervous system in a monophyletic group of the Acoela. Six species of Paraphanostoma were chosen for the study. Using immunocytochemical methods and confocal scanning laser microscopy, the immunoreactive patterns of serotonin (5-HT) and the neuropeptide GYIRFamide were described in detail. The study has demonstrated that the brains in Paraphanostoma species, although diverse in detail, still follow the same general pattern. 18S rDNA sequences were used to generate a hypothesis of the phylogeny within the group. Characters of the nervous system revealed in this study were coded and analysed together with 18S rDNA data. Several synapomorphies in the nervous system characters were identified. However, numerous parallelisms in the nervous system evolution have occurred. Data obtained demonstrate that the genus Paraphanostoma is closely related to Childia and should belong to the same family, Childiidae.

Phylogeny of the African murid tribe Otomyini (Rodentia), based on morphological and allozyme evidence

Abstract: Based on a cladistic analysis of 45 morphological (craniodental) and 46 binary allozyme characters, previous systematic treatments of the African murid tribe, Otomyini (laminate-toothed rats), are reviewed. Cladistic analysis of the craniodental data, involving eight outgroup taxa, confirmed the monophyly of the Otomyini, and suggested Pelomys to represent the sister genus of the Otomyini. Craniodental synapomorphies provided strong support for certain basal relationships among Otomyini rodents, reinforcing available palaeontological evidence. However, poor statistical (Bremer decay index) support was obtained for terminal relationships. The data presented revealed a ‘mesic clade’ of southern and eastern African species, with Otomys sloggetti basal to this group. The arid-adapted, southern Africa-endemic species, Parotomys littledalei, P. brantsii and O. unisulcatus, were all placed basal to the ‘mesic clade’, but did not form a separate ‘arid clade’, as suggested by earlier biochemical studies. Two allozyme synapomorphies supported the existence of the ‘mesic clade’, separate from arid-adapted southern African species. A strict cladistic interpretation of the present data did not support the existence of two genera in the tribe, and the two species of Parotomys (whistling rats) should be transferred to Otomys. At the species level, specific identity of O. lacustris and O. barbouri, distinct from O. anchietae, was supported by several autapomorphies, and O. tropicalis burtoni was shown to be included in O. angoniensis rather than O. tropicalis, extending the range of the former species into West Africa.

Comparative morphology of the venom apparatus in the braconid wasp subfamily Rogadinae (Insecta, Hymenoptera, Braconidae) and related taxa

Abstract: The morphology of the venom apparatus intima in representatives of 38 genera of the problematic braconid wasp subfamily Rogadinae and other cyclostome braconids was investigated and a preliminary phylogenetic analysis for the group was performed with the information obtained. Despite the limited number of characters, the data suggest several relationships at various taxonomic levels. The venom apparatus in the Clinocentrini and the Stiropiini is relatively unmodified and similar to that found in other genera previously placed within a broader concept of the Rogadinae (e.g. genera of Lysitermini, Pentatermini, Tetratermini, Hormiini) and also to that of the Betylobraconinae. The presence of a cone of filaments located inside the secondary venom duct near to its insertion on the venom reservoir/primary venom duct is proposed as a synapomorphy for the tribe Rogadini to the exclusion of Stiropiini, Clinocentrini and Yeliconini. Other features of the secondary venom duct and its insertion on the venom reservoir/primary venom duct support a number of relationships between the genera of the Rogadini and also within the large genus Aleiodes. A clade containing 15 Rogadini genera (Bathoteca, Bathotecoides, Bulborogas, Canalirogas, Colastomion, Conspinaria, Cystomastacoides, Macrostomion, Megarhogas, Myocron, Pholichora, Rectivena, Rogas, Spinaria and Triraphis) is supported by the presence of a thickened and short secondary venom duct, whereas the different members of Aleiodes (excluding members of the subgenus Heterogamus) and Cordylorhogas are distinguished by having a recessed secondary venom duct with well-defined and numerous internal filaments. New World Rogas species exhibit a unique venom apparatus and may not be closely related to the Old World ones. Features of the venom apparatus of the enigmatic genus Telengaia and the exothecine genera Shawiana and Colastes suggest that the Telengainae and Exothecinae are both closely related to the Braconinae, Gnamptodontinae, and possibly to the Opiinae and Alysiinae. An unsculptured venom reservoir was found in one specimen of the type species of Avga, A. choaspes, which is consistent with it occupying either a very basal position within the cyclostome braconids or belonging to a recently recognized ‘Gondwanan’ clade that also includes the Aphidiinae

First International Phylogenetic Nomenclature Meeting: a report

Abstract: The First International Phylogenetic Nomenclature Meeting convened in Paris from July 6–9, 2004 in the Museum National d’Histoire Naturelle. This meeting marked a turning point in the history of biological nomenclature — the inauguration of a new society (the International Society for Phylogenetic Nomenclature) that will soon launch a code of phylogenetic nomenclature (the PhyloCode) that will represent a fundamental change in the way in which taxon names are defined.

Phylogenetic nomenclature is compatible with diverse philosophical perspectives

Abstract: Pleijel, F. & Harlin, M. (2004). Phylogenetic nomenclature is compatible with diverse philosophical perspectives. — Zoologica Scripta, 33, 587–591.

Modelling phylogenetic relationships using reticulated networks

Abstract: 89Makarenkov, V., Legendre, P. & Desdevises, Y. (2004). Modelling phylogenetic relationshipsusing reticulated networks. — Zoologica Scripta, 33, 89–96.Most traditional methods of phylogenetic analysis assume that species evolution can be rep-resented by means of a bifurcating tree model. In many phylogenetic situations, however,some of the evolutionary links between species are due to reticulate evolution. For instance,reticulate models can adequately describe such complicated mechanisms as lateral gene trans-fer in bacteria or species hybridization. The theoretical concepts of reticulate evolution devel-oped in the 1980s and 1990s need to be supported by appropriate analytical tools and software.In this paper, we present the main features of a new distance-based method for modelling phy-logenetic relationships among species by means of reticulated networks (RNs). The methoduses the least-squares model to build a RN by gradually improving upon the solution providedby a phylogenetic tree. A computer program facilitating the reconstruction and visualizationof reticulate phylogenies is made available to researchers. In the application section, we illus-trate the usefulness of the method by studying the evolution of honeybees (genus Apis). Themethod for reconstructing RNs has been included in the T-Rex (Tree and Reticulogram Recon-struction) package recently developed by the first-named author.Vladimir Makarenkov. Departement d’informatique, Universite du Quebec a Montreal, C.P. 8888,succursale Centre-Ville, Montreal (Quebec), Canada, H3C 3P8. E-mail:makarenkov.vladimir@uqam.caPierre Legendre, Departement de sciences biologiques, Universite de Montreal, C.P. 6128, succursaleCentre-Ville, Montreal (Quebec), Canada, H3C 3J7. E-mail: pierre.legendre@umontreal.caYves Desdevises, Laboratoire Arago, Universite Pierre et Marie Curie, UMR CNRS 7628, BP 44,66651 Banyuls-sur-Mer Cedex, France. E-mail: desdevises@obs-banyuls.fr

Anatomy and relationships of the Early Cambrian worm Myoscolex

Abstract: Dzik, J. (2004). Anatomy and relationships of the Early Cambrian worm Myoscolex. —Zoologica Scripta, 33, 57–69. Numerous fossil specimens of Myoscolex ateles Glaessner, 1979 from the late Early Cambrian Emu Bay shale of Kangaroo Island, South Australia with phosphatized organic matter-rich tissues show its muscular body wall penetrated by rows of rod-like structures — possible chaetae. The body wall was composed of an external layer with transverse (circular) fibres. This layer was thickest in lateral parts of the body and very thin dorsally. In the ventro-lateral quarter of the body circumference, a belt of longitudinal fibres extended along the body. Longitudinal fibres also occurred in the dorsal region of the body. Along the venter extended a narrow longitudinal belt of probably oblique cords, crossing themselves perpendicularly. In having a virtually smooth, laterally flattened body, Myoscolex closely resembles the slightly geologically younger Pikaia from the Burgess shale of British Columbia, generally believed to be one of the oldest chordates. Being the oldest probable annelid, at least superficially similar to the opheliid polychaetes, Myoscolex may appear not too distant from the ancestor of the phylum. The lateral body flattening of Myoscolex was apparently an adaptation to swimming by undulation of the body in transverse plane, similar to today's errant polychaetes but without using chaetae or appendages in propulsion.

The measurement of test severity, significance tests for resolution, and a unified philosophy of phylogenetic inference

Abstract: Queiroz, K de (2004) The measurement of test severity, significance tests for resolution, and a unified philosophy of phylogenetic inference — Zoologica Scripta , 33 , 463–473 The philosopher Karl Popper described a concept termed degree of corroboration , C , for evaluating and comparing hypotheses according to the results of their tests C is, fundamentally, a comparison of two likelihoods: p ( e | hb ), the likelihood of the hypothesis ( h ) in conjunction with the background knowledge ( b ), and p ( e|b ), the likelihood of b alone C is closely related to the likelihood ratio of nested hypotheses When phylogenetic analysis is interpreted as an attempt to assess C for a phylogenetic tree (the hypothesis, h ), several interpretations have been given for p ( e|b ) Here I describe a new interpretation that equates p ( e|b ) with the probability of the data in the absence of a hypothesis of phylogenetic resolution, that is with the likelihood of an unresolved or polytomous tree Under this interpretation, C for a fully or partially resolved phylogenetic tree is the likelihood of that tree minus the likelihood of the corresponding unresolved tree These same two likelihoods can be used in a likelihood ratio test (LRT) to assess the significance of the degree of corroboration of the hypothesis of phylogenetic resolution This LRT for resolution is closely related to permutation tests for phylogenetic structure in the data, because data that evolved on a true polytomous tree are expected to be phylogenetically randomized It therefore reconciles the interpretation of the evidence ( e ) as the distribution of character states among taxa (rather than the score of the optimal tree) with the interpretation of permutation tests as methods for assessing C Likelihood methods are (contrary to the views of some commentators) central to understanding how Popper’s C applies to phylogenetic hypotheses, and they form the foundation of a unified and inclusive philosophy of phylogenetic inference