Paraphyletic Research Articles

Proper names in twin worlds: Monophyly, paraphyly, and the world around us

This paper addresses the theoretical relevance of monophyletic, paraphyletic and polyphyletic groups under the paradigm of sophisticated scientific realism. The doctrine of metaphysical realism is introduced using the philosophy of Karl Popper as an example, which is then contrasted with scientific realism. A discussion of the nature of causal relations presents an account of counterfactual conditionals. The current state of art casts the theory of phylogenetic systematics in a stark contrast of classes (universals) and individuals (particulars). In practice, however, individuals piggyback on classes, or sets. Natural kinds are introduced in order to overcome this deep dichotomy. The theoretical relevance of natural kinds lies in their explanatory value, and that may change with changing context. It is for this reason that non-monophyletic groups can have explanatory value (their members can function as tokens of causally relevant kinds) within certain domains of evolutionary biology. Explanatory value is maximized by integration of the genealogical hierarchy of species and monophyletic taxa with other areas of evolutionary biology.

Testing the sister-group relationship of the Rhodniini and Triatomini (Insecta: Hemiptera: Reduviidae: Triatominae)

The Reduviidae (Assassin bugs) form one of the largest and morphologically most diverse families of Hemiptera, with well over 6500 recognised species of worldwide distribution (Maldonado Capriles, 1990; Schuh and Slater, 1995). They have been variously grouped into up to 32 subfamilies, of which 22 were recognised by Davis (1969), 21 by Putchkov and Putchkov (1985), 25 by Maldonado Capriles (1990), and 22 by Schuh and Slater (1995). Most are predators on other invertebrates, although the subfamily Triatominae is customarily deWned on the basis of a blood-sucking habit and associated morphological adaptations (Jeannel, 1919; Lent and Wygodzinsky, 1979; Usinger, 1943). Over 130 species of Triatominae are currently recognised, currently grouped into up to 19 genera and 6 tribes (Galvao et al., 2003) of which the Triatomini and Rhodniini are the most numerous and most epidemiologically signiWcant (Table 1). In general, the various subfamilies of the Reduviidae have been assumed to be monophyletic, although in the case of the Triatominae several arguments have been advanced to suggest that these blood-sucking forms may have diVerent origins within the predatory subfamilies, thus representing a polyphyletic or paraphyletic group (SchaVer, 2003). Evidence to support this idea has been drawn from several comparative sources including overt morphology and biogeography (SchoWeld, 1988; SchoWeld and Dolling, 1993), morphometric comparisons (e.g., Dujardin et al., 2000), comparative sensilla arrays (e.g., Catala, 1997), mitochondrial and ribosomal DNA sequence comparisons (e.g., Garcia, 1999; Garcia and Powell, 1998; Lyman et al., 1999; Monteiro et al., 2000; Marcilla et al., 2001; Stothard et al., 1998) and the presence of markedly diVerent antihaemostatic compounds in the saliva of Rhodniini and Triatomini (Ribeiro et al., 1998). In addition, since a degree of at least facultative haematophagy has arisen in a range of other Hemiptera it has been suggested that this may be a relatively simple evolutionary step from an otherwise predaceous habit, so that polyphyletic derivation of haematophagy may be expected within this group (SchoWeld, 2000). Nevertheless, the assumption of monophyly within the Triatominae has persisted—at least implicitly—in several revisions (e.g., Schuh and Slater, 1995), and also in some DNA sequence-based comparisons (e.g., Hypsa et al., 2002). Moreover, all arguments so far advanced to support a polyphyletic concept of the Triatominae have been based on comparisons within the subfamily itself rather than on comparisons with other Reduviidae. We therefore present here an analysis of currently available gene sequence data—16S ribosomal DNA (mitochondrial gene)—for species of Triatominae representing the tribes Rhodniini and Triatomini (as deWned by Lent and Wygodzinsky, 1979) together with representatives of other subfamilies of the Reduviidae.

A molecular analysis of some Eastern Atlantic grouper from the Epinephelus and Mycteroperca genus

Mitochondrial cytochrome b (397 bp) and 16S rDNA (516 bp) sequences analysis was used to investigate the phylogenetic relationships among some Eastern Atlantic Epinephelinae species. Six species of Epinephelus ( E. aeneus, E. caninus, E. costae, E. haifensis, E. marginatus and E. tauvina) and two species of Mycteroperca ( M. rubra and M. fusca) were analysed. Neighbour-joining and maximum-parsimony analysis support the paraphyletic grouping of the Epinephelus and Mycteroperca analysed. The maximum pairwise nucleotide divergence value in cyt b among all taxa was 0.196 between E. aeneus and E. marginatus and the minimum value was 0.006 between E. costae and M. rubra. Meanwhile, in 16S sequence analysis, the maximum value is 0.093 between E. aeneus and E. tauvina and the minimum value is 0.011 between E. marginatus and M. rubra. Molecular clock estimates for the species suggest a divergence time of 20–24 mya, which coincides with the Miocene period. A molecular analysis was also conducted, using other Epinephelinae sequences from GenBank in order to improve our understanding of the phyletic status of the Epinephelus and Mycteroperca species analysed.

Phylogenetic Relationships of Xylella fastidiosa Strains Based on 16S–23S rDNA Sequences

In spite of the lack of resolution of Xylella fastidiosa phylogenetic relationships, parsimony analysis of the 16S-23S rDNA sequence from a wide range of hosts has been evaluated in this research. In order to establish an easier method for sequencing the spacer region completely, a new primer pair was designed. The sequences obtained revealed a higher level of variation than that found in 16S gene sequences, with similarity values ranging from 0.80 to 1.00. The cladogram constructed allowed the clustering of two major clades. From these results it has been possible to recognize the monophyletic grouping of some strains belonging to the same host, possibly representing only one infection process. However, for other hosts there is paraphyletic and polyphyletic grouping. This methodology followed from promising results regarding strain-host clustering. With the parsimony approach, hypothetical genealogical relationship among Xylella strains may be inferred.

Do nine-primaried passerines have nine or ten primary feathers? The evolution of a concept

The number of primary feathers in a bird’s wing has been used as a systematic character since the first half of the nineteenth century. During the years, though, the definition of which feathers to count as a primary has changed and today the species historically denoted as having only nine primaries are instead said to have, for example, nine ‘functional’ primaries. In this study, I investigated the borderline between ‘nine-primaried’ and ‘ten-primaried’ birds to search for a proper definition of the term ‘nine-primaried’. A total of 161 specimens of 104 bird species, mainly passerines, were examined. All species examined had ten primaries although the ‘nine-primaried’ species had primary ten more or less concealed under primary covert nine. The number of primary coverts has decreased over time, with ten primary coverts as the ancestral state within Passeriformes and nine primary coverts among most oscine species. In conclusion, a proper definition of ‘nine-primaried’ might be “with primary ten concealed by primary covert nine”. This definition includes all taxa historically denoted ‘nine-primaried’, i.e. systematically it is a definition of a paraphyletic group. The term ‘nine-primaried’ is thus too inclusive to be of more than very limited systemtic value and, consequently, the New World nine-primaried oscines group might gain from a new denotation.

Evolution of Suessenguthia (Acanthaceae) inferred from morphology, AFLP data, and ITS rDNA sequences

The phylogeny and evolution of Suessenguthia (Acanthaceae), a genus of six species from the Andean foothills and adjacent Amazonia in Bolivia, Peru and western Brazil, are discussed based on morphological and molecular (amplified fragment length polymorphism, ITS rDNA) data. Suessenguthia forms a paraphyletic group at the base of the larger genus Sanchezia. The non-overlapping geographical distribution of closely related species suggests that parapatric or allopatric speciation is the major mode in the genus. A major evolutionary tendency promoting diversification of the group presumably was a change from bee- to hummingbird pollination, resulting in a successive adaptation of flower morphology and inflorescence structure.

On the quality of tree-based protein classification

Phylogenetic analysis of protein sequences is widely used in protein function classification and delineation of subfamilies within larger families. In addition, the recent increase in the number of protein sequence entries with controlled vocabulary terms describing function (e.g. the Gene Ontology) suggests that it may be possible to overlay these terms onto phylogenetic trees to automatically locate functional divergence events in protein family evolution. Phylogenetic analysis of large datasets requires fast algorithms; and even 'fast', approximate distance matrix-based phylogenetic algorithms are slow on large datasets since they involve calculating maximum likelihood estimates of pairwise evolutionary distances. There have been many attempts to classify protein sequences on the family and subfamily level without reconstructing phylogenetic trees, but using hierarchical clustering with simpler distance measures, which also produce trees or dendrograms. How can these trees be compared in their ability to accurately classify protein sequences? Given a 'reference classification' or 'group membership labels' for a set of related protein sequences as well as a tree describing their relationships (e.g. a phylogenetic tree), we propose a method for dividing the tree into monophyletic or paraphyletic groups so as to optimize the correspondence between the reference groups and the tree-derived groups. We call the achieved optimal correspondence the 'accuracy of a tree-based classification (TBC)', which measures the ability of a tree to separate proteins of similar function into monophyletic or paraphyletic groups. We apply this measure to compare classical NJ and UPGMA phylogenetic trees with the trees obtained from hierarchical clustering using different protein similarity measures. Our preliminary analysis on a set of expert-curated protein families and alignments suggests that there is no uniformly superior algorithm, and that simple protein similarity measures combined with hierarchical clustering produce trees with reasonable and often the most accurate TBC. We used our measure to help us to design TIPS, a tree-building algorithm, based on agglomerative clustering with a similarity measure derived from profile scoring. TIPS is comparable with phylogenetic algorithms in terms of classification accuracy and is much faster on large protein families. Due to its time scalability and acceptable accuracy, TIPS is being used in the large-scale PANTHER protein classification project. The trees produced by different algorithms for different protein families can be viewed at http://panther.appliedbiosystems.com/pub/tree_quality/trees.jsp. For every tree and every level of classification granularity we provide the optimal TBC along with the reference classification. The script that evaluates the accuracy of TBC is available at http://panther.appliedbiosystems.com/pub/tree_quality/index.jsp

Paraphyletic group, PhyloCode and phylogenetic species—the current debate and a preliminary commentary

In this essay, three currently hotly debated issues in biological systematics, i.e., the paraphyletic group, the PhyloCode, and the phylogenetic species concept, have been briefly reviewed. (1) It is widely acknowledged that cladistics has made some positive contributions to the study of systematics. In particular, the employment of outgroup analysis for assessing character polarities, the application of synapomorphies to the inference of relationships between taxa, and the use of cladistic methods for reconstructing phylogeny, have all greatly facilitated the improvement of systematic approaches. A fatal flaw in cladistics is its refusal to accept paraphyletic groups. Frankly, we are adherents and practitioners of phyletics, and hence consider paraphyletic groups to be acceptable. For example, an AFLP analysis has shown that Zabelia (Caprifoliaceae) can be included in Abelia, but the members in Zabelia differ from those in Abelia not only in pollen morphology, but also in having persistent petioles dilated and connate at base, thus enclosing axillary buds, characters of adaptive significance obtained possibly when Zabelia members entered a new ecological niche, so we consider that they are better treated as two independent genera, though indeed such a treatment makes Abelia paraphyletic. (2) Some cladists pointed out that as the tool for communication and the system for information storage and retrieval, biological nomenclature is required to be unambiguous, unique and stable. They criticise the Linnaean rank-based system of nomenclature for failing to satisfy such requirements for the naming of clades and species. To address this problem, the PhyloCode is proposed in recent years, in which three definitions for clade naming are given, i.e., the node-based, the stem-based, and the apomorphy-based. We are of the opinion that since the Linnaean binominal system of botanical nomenclature has existed for nearly 250 years, the rejection of this system and the adoption of the PhyloCode would create a state of chaos in botanical nomenclature. This does not mean that there exist no merits in the proposals made by the PhyloCode supporters. We suggest that further studies should be conducted for its practical application. (3) It has been well known that there are many problems with the application of the biological species concept in plants, and thus at the present time the majority of plant systematists actually

Some palaeozoic ‘protorthoptera’ are ‘ancestral’ orthopteroids: Major wing braces as clues to a new split among the ‘protorthoptera’ (Insecta)

Synopsis The Archaeorthoptera is an apomorphy‐based supra‐ordinal clade of insect that mainly comprises the extant order Orthoptera and the fossil orders Caloneurodea and Titanoptera. Our knowledge of this group is, however, incomplete because some taxa assigned to the paraphyletic group ‘Protorthoptera’ have not been accurately described. Here we describe or re‐describe Carboniferous material assigned to Omalia macroptera Van Beneden & Coemans, 1867, Coselia palmiformis Bolton, 1922, Palaeomastax carbonis Handlirsch, 1904, Omaliella ramosa n. gen. & n. sp., Eoblatta robusta (Brongniart, 1885), Ctenoptilus elongatus (Brongniart, 1893) n. comb., Ischnoptera diaphanes n. gen. & n.sp., Protodiamphipnoagaudryi (Brongniart, 1885), Cnemidolestes woodwardi (Brongniart, 1893), Ischnoneura oustaleti (Brongniart, 1885), Narkeminopsis eddi Whalley, 1979 and Bouleites latipennis Lameere, 1917. These taxa are all assigned to the Archaeorthoptera, based on the presence of characteristic features in their wing venation. A new system of homologies of the median system is proposed, which implies the loss of vein MP has occurred in some of these taxa. Our new data demonstrate the weakness of the previous specific, generic, familial and ordinal taxonomies.

NEW DATA ON PALEOZOIC GRYLLOBLATTID INSECTS (NEOPTERA)

The paraphyletic group Grylloblattida, formerly included in the Protorthoptera sensu lato, is of great importance for the phylogeny of some ‘polyneopterous’ orders, because it is currently assumed to contain the sister groups of modern Plecoptera (stoneflies) and Grylloblattodea. We present new discoveries and new observations and interpretations concerning this crucial assemblage. Lodevopterum angustus new genus and species and Depressopterum minutus new species are described from the Upper Permian of the Lodève Basin (Hérault, France). The ‘Grylloblattida’ from Commentry (Allier, France; Upper Carboniferous) are revised: the family Protoblattinidae Meunier, 1909a is restored after the redescription of the type species Protoblattina bouvieri Meunier, 1909a; the family Stenoneuritidae Lameere, 1917 is revised; Palaeocixius fayoli Handlirsch, 1906b and Protoblattina giardi Meunier, 1921 are synonymized with Palaeocixius antiquus Brongniart, 1893, followed by a discussion on the possible familial attributions of this genus. The affinities of the poorly known family Protoperlidae Brongniart, 1885a are discussed after a critical compilation of the available data, i.e., Protoperla boltoni Lameere, 1917 is considered as Grylloblattida incertae sedis and Polyetes elegantissima Meunier, 1921 is transferred into Laurentiaupterum new genus and assigned to Paraphenopteridae new family. Close phylogenetic relationship of Atactophlebia termitoides Martynov, 1928 (Tikhiye Gory, Tatarian Republic, Russia; Upper Permian) and Pinidelia sukatshevae Storozhenko, 1994 (Tyulkino, Perm region, Russia; Upper Permian) is based on a new interpretation of their forewing venation. The recent taxonomy and phylogeny of the Grylloblattida proposed by Storozhenko (1998, 2002) are refuted. The current concepts of Protorthoptera, Plecopteroid Assemblage, and Grylloblattida are compared.

The phylogeny and classification of the tribe Halictini, with special reference to the Halictus genus-group (Hymenoptera: Halictidae)

The monophyly of the cosmopolitan tribe Halictini, including over 2300 currently recognized species, is supported by at least a single manifested synapomorphy shared by all members of the tribe: metasomal tergum VII of the male is modified; this forms a transverse ridge giving a false apex beneath which the tergum is strongly reflexed to the morphological posterior margin. On the basis of the present phylogenetic analysis, the tribe Halictini is subdivided into five subtribes: Halictina (comprised of 7 genera: Echthralictus, Glossodialictus, Halictus, Homalictus, Patellapis, Seladonia, and Thrincohalictus), Sphecodina (4 genera: Eupetersia, Microsphecodes, Ptilocleptis, and Sphecodes), Thrinchostomina (2 genera: Thrinchostoma and Parathrincostoma), Caenohalictina (9 genera: Agapostemon, Caenohalictus, Dinagapostemon, Habralictus, Mexalictus, Paragapostemon, Pseudagapostemon, Rhinetula, and Ruizantheda), and Gastrohalictina (one large and diverse genus: Lasioglossum s. l.). The subtribe Halictina is a paraphyletic group; the remaining four tribes are strictly monophyletic (holophyletic). The monophyly of the Halictus genus-group, comprising the genera Halictus and Seladonia, is supported by two distinct synapomorphies of the male genitalia: (1) dorsal gonostylus simple (not double), flattened, broad, narrowed proximally, and provided with a clump of very coarse bristles on the inner surface; (2) ventral gonostylus sclerotized, relatively thin and long, directed backward. Only generalized members of the group possess both the character states above. In derived members, the clump of bristles and the ventral gonostylus are often lost independently. The sister group of the Halictus genus-group is the genus Thrincohalictus. This is supported by the following synapomorphy found among the tribe Halictini only in Halictus, Seladonia, and Thrincohalictus: the ventral gonobasal rim in the male genitalia is forming a right posterolateral angle with a short projection directed laterally. Another feature characterizing these three genera is the presence (except in the parasitic subgenus Paraseladonia) of posterior bands of tomentum or dense and much plumose appressed hairs on the metasomal terga. However, this character is shared also with Patellapis subgenera Patellapis and Lomatalictus. The following subgeneric classification of the genera Halictus and Seladonia is suggested. The genus Halictus includes 12 subgenera: Acalcaripes (2 species), Argalictus (8), Halictus s. str. (4), Hexataenites (11), Lampralictus (1), Monilapis (29), Nealictus (2), Odontalictus (2), Platyhalictus (14), Protohalictus (13), Ramalictus (2), and Tytthalictus (4). The genus Seladonia comprises 6 subgenera, including 2 new ones: Mucoreohalictus subg. n. (15), Pachyceble (22), Paraseladonia (1), Placidohalictus subg. n. (5), Seladonia s. str. (36), and Vestitohalictus (16). The subgenera of Halictus and Seladonia are keyed. The phylogenetic tree of the subgenera of Halictus and Seladonia is reconstructed with use of 46 morphological characters of adults. All other genera of the tribe Halictini were taken as outgroup. The genus Halictus is ascertained as a strictly monophyletic group based upon a single postulated synapomorphy: dorsal gonostylus with a triangular hair patch as a distal appendage on inner side. All the subgenera of this genus appear as strictly monophyletic groups, with the exception of Monilapis, which is a paraphyletic one in relation to Acalcaripes. The monophyly of the genus Seladonia is supported by three synapomorphies, including a novelty (unique synapomorphy): male dorsal gonostylus with a deep cleft. All the subgenera of this genus are strictly monophyletic with the exception of Placidohalictus, which is a paraphyletic one in relation to Vestitohalictus and Mucoreohalictus. A synonymical catalogue of species and species-group names in the genera Halictus and Seladonia, including 442 names, is provided as an appendix.

Phylogeny and behaviour of the Gollumiellinae, a new subfamily of the ant‐parasitic Eucharitidae (Hymenoptera: Chalcidoidea)

Abstract. Gollumiellinae is proposed as a new subfamily for the Indo‐Pacific genera Gollumiella Hedqvist and Anorasema Bouc̆ek based on analyses of three ribosomal transcript gene regions (28S‐D2 and ‐D3, and 18S‐E23; 1262 aligned base pairs) sequenced for twenty‐eight genera and fifty‐four species of Eucharitidae, and twelve genera and nineteen species of Pteromalidae (Pteromalinae) and Perilampidae (Chrysolampinae and Perilampinae). Gollumiella and Anorasema have been treated as either a monophyletic or paraphyletic group within the Eucharitini (Hymenoptera: Eucharitidae). The monophyly of the Eucharitidae and a sister‐group relationship between Gollumiellinae and Oraseminae + Eucharitinae are supported strongly in parsimony and Bayesian analyses. The molecular phylogeny contradicts previous morphological hypotheses, in which Gollumiella and Anorasema are within Eucharitini. The strength of the molecular hypothesis is explored through evaluations of data alignments that are eye optimized, clustal‐x aligned, eye optimized but with gaps coded as a fifth base, and eye optimized and combined with morphological data. The oviposition behaviour and host associations of G. longipetiolata on tree ferns (Cyathea latebrosa, Cyatheaceae), and the morphology of the planidium and pupa are newly described. Eggs are deposited vertically, with the base anchored into the leaf or petiolar surface. Unlike other Eucharitidae, the eggs are not stalked, but rather tipped with a secretion that may act as an attractant for their ant host, Paratrechina sp. (Formicidae: Formicinae). Various morphological character state optimizations and their implications for convergent morphology, behaviour and host associations are discussed. All results using gene regions treated separately or combined with the morphological data reach the same conclusion: Anorasema + Gollumiella form the sister group of Oraseminae + Eucharitinae, and thus deserve subfamily status. This suggests that very distinct traits such as the fusion of the pronotum and prepectus in adults and the fusion of the first two tergites of the planidia are convergent apomorphies. Molecular data, when strongly supported, can provide new information to unravel convergent from synapomorphic changes, resulting in more robust hypotheses of relationship.

실고기목 어류 (Syngnathiformes)의 분자계통학적 분류

The previous morphology-based taxonomic frameworks within the family Syngnathidae had emphasized the significance of the male brood pouch and reproductive biology in defining the group. However, several different hypotheses had been proposed by different investigators. This study has been carried out to determine the phylogenetic relationships among 19 species belonging to the order Syngnathiformes with three Gasterosteiformes species as outgroup taxa by using the mitochondrial cytochrome b DNA sequences. Phylogenetic analyses based on neighbor-joining distance, maximum parsimony, minimum evolution and maximum likelihood method strongly supported that the family Syngnathidae, the suborder Syngnathoidei and the order Syngnathiformes were all monophyletic group. Although much of previous morphological analyses were supported by our molecular data, there were some significant discrepancies between molecular and morphological work. Such an interesting result was that the weedy seadragon (Phyllopteryx taeniolatus) strongly grouped together with the New Zealand pot-belly seahorse (Hippocampus abdominalis). Considering the markedly different brooding structure between them, this unexpected result might be explained whether by multiple independent origins of brooding structure or by hybridization between the female Hippocampus and other syngnathid species having individual membranous egg compartment. In addition, the suborder Aulostomoidei was paraphyletic group because the shrimpfish (Aeliscus strigatus), belonging to the family Centriscidae, always grouped together with the family Syngnathidae as a sister taxon.

A molecular phylogeny of Oxya (Orthoptera: Acridoidea) in China inferred from partial cytochrome b gene sequences

The grasshoppers of the genus Oxya are well known to damage rice, sugar cane, and other crops, yet their phylogenetic relationships have not been examined with molecular data. In this study, we obtained the 432 bp DNA sequences of the mitochondrial cytochrome b gene from 91 individuals of nine Oxya species and two outgroups ( Gesonula punctifrons and Acrida cinerea). Phylogenetic analyses for the molecular data set were then carried out using the maximum parsimony and neighbor-joining methods. The results showed that the nine Oxya species form four well-supported clades, which include (1) O. intricata and O. flavefemura; (2) O. japonica and O. bicingula; (3) O. agavisa; and (4) O. chinensis, O. brachyptera, O. adentata, and O. hainanensis, respectively. In particular, the monophyly of O. hainanensis and O. agavisa is strongly supported, respectively. However, O. flavefemura and O. intricata, O. bicingula, and O. japonica form paraphyletic groups, respectively, and O. chinensis, O. adentata, and O. brachyptera form a polyphyletic group, suggesting that they should be merged as few as three species.

Larval development of Apiomithrax violaceus (A. Milne Edwards, 1868) (Decapoda: Brachyura: Majoidea: Pisidae) reared in laboratory conditions, and a review of larval characters of Pisidae

Apiomithrax violaceus (A. Milne Edwards, 1868) is a pisid majoid crab occurring in tropical and subtropical coastal waters of the eastern and western South Atlantic. Larval development consists of two zoeal stages and a megalopa. Beginning with the first zoea, the duration of each larval stage at 24°C was 3–8 (5±1), 3–5 (4±0.5) and 9–15 (11±2) days, the megalopa and first crab instar appearing 9–11 (10±1) and 20–27 (23±2) days after hatching, respectively. Larval characters agree with those proposed for the Majoidea, in having nine or more setae on the scaphognathite in the first zoea and well-developed pleopods in the second zoea. However, larvae of A. violaceus do not fit larval pisid features. Zoeal stages differ from most other Pisidae in having lateral spines, a long rostral spine extending beyond the antenna, two spines per telson fork and a dorsolateral process on the third abdominal somite. The megalopa differs in having a spine dorsally on the carapace and on the basial segment of the second pereiopod. Two characters that are potentially unique to Apiomithrax include a zoeal antenna with an exopod that is much longer than the protopod, and a rostral spine that is longer than the dorsal spine. These characters should facilitate the identification of this taxon and could also be useful for phylogenetic studies. A review of larvae of 28 species among 14 genera indicated that there is no apparent single larval character that differentiates the Pisidae, with more limited phylogenetic analyses suggesting that this is a paraphyletic group. Apiomithrax, Eurynolambrus, Pisoides, Rochinia and Scyra have the most divergent morphological characters within the family. The analysis and inclusion of additional taxa is likely to shed more light on the sister-group relationships of the Pisidae. However, based on the extent of morphological interspecific variability of known larvae it is likely that the group, as presently defined by adult morphology, is not monophyletic.

Phylogenetic relationships within the family Talpidae (Mammalia: Insectivora)

AbstractThe phylogenetic relationships within the family Talpidae (Mammalia, Insectivora) were inferred from parsimonious analysis of 55 morphological characters, which were collected from 43 skulls of 22 species, and which included all of the extant genera in the current taxonomy of the Talpidae. Two equally parsimonious cladograms were obtained. The Uropsilinae, Desmaninae and Talpinae subfamilies formed monophyletic clades, and the Desmaninae and Talpinae subfamilies were sister clades. The Talpinae subfamily was subdivided into the following clades: Urotrichus, Dymecodon, Scalopus–Scapanus, Parascalops–Scapanulus, Condylura, Neurotrichus, Scaptonyx and Talpini (Talpa, Euroscaptor, Scaptochirus, Parascaptor and Mogera). The Urotrichini (Urotrichus, Dymecodon and Neurotrichus) and Scalopini (Scalopus, Scapanus, Parascalops and Scapanulus) were considered to be paraphyletic, and were defined primarily by their ancestral characteristics. Since the genus Dymecodon was not clustered with Urotrichus, it was considered to be a valid genus, and not just a junior synonym of the latter. The Asian genus Euroscaptor was found to be a paraphyletic group, and deserves further systematic revision. The evolution from semi‐fossorial habit to fully fossorial habit seems to have occurred three times in the Scalopus–Scapanus, Parascalops–Scapanulus and Talpini clades. The migration of the Talpinae to North America from Eurasia probably occurred on four separate occasions in the Scalopus–Scapanus, Parascalops, Condylura and Neurotrichus lineages.

Molecular data and the evolutionary history of dinoflagellates

We have sequenced small-subunit (SSU) ribosomal RNA (rRNA) genes from 16 dinoflagellates, produced phylogenetic trees of the group containing 105 taxa, and combined small- and partial large-subunit (LSU) rRNA data to produce new phylogenetic trees. We compare phylogenetic trees based on dinoflagellate rRNA and protein genes with established hypotheses of dinoflagellate evolution based on morphological data. Protein-gene trees have too few species for meaningful in-group phylogenetic analyses, but provide important insights on the phylogenetic position of dinoflagellates as a whole, on the identity of their close relatives, and on specific questions of evolutionary history. Phylogenetic trees obtained from dinoflagellate SSU rRNA genes are generally poorly resolved, but include by far the most species and some well-supported clades. Combined analyses of SSU and LSU somewhat improve support for several nodes, but are still weakly resolved. All analyses agree on the placement of dinoflagellates with ciliates and apicomplexans (=Sporozoa) in a well-supported clade, the alveolates. The closest relatives to dinokaryotic dinoflagellates appear to be apicomplexans, Perkinsus, Parvilucifera, syndinians and Oxyrrhis. The position of Noctiluca scintillans is unstable, while Blastodiniales as currently circumscribed seems polyphyletic. The same is true for Gymnodiniales: all phylogenetic trees examined (SSU and LSU-based) suggest that thecal plates have been lost repeatedly during dinoflagellate evolution. It is unclear whether any gymnodinialean clades originated before the theca. Peridiniales appear to be a paraphyletic group from which other dinoflagellate orders like Prorocentrales, Dinophysiales, most Gymnodiniales, and possibly also Gonyaulacales originated. Dinophysiales and Suessiales are strongly supported holophyletic groups, as is Gonyaulacales, although with more modest support. Prorocentrales is a monophyletic group only in some LSU-based trees. Within Gonyaulacales, molecular data broadly agree with classificatory schemes based on morphology. Implications of this taxonomic scheme for the evolution of selected dinoflagellate features (the nucleus, mitosis, flagella and photosynthesis) are discussed.

Chromosomal and molecular characterization of Aethomys kaiseri from Zambia and Aethomys chrysophilus from Tanzania (Rodentia, Muridae).

Aethomys is a common and widespread rodent genus in the African savannas and grasslands. However, its systematics and taxonomy are still unclear as no study has covered the entire range. In fact it might not be a monophyletic genus and perhaps should be split into two subgenera, Micaelamys and Aethomys. In this paper, we present findings based on the cytogenetics and the entire cytochrome b sequence of two species from Zambia (A. kaiseri) and Tanzania (A. chrysophilus), and we compare them with the sequences of a South African species (A. namaquensis) and other allied muroid genera. Comparison of the banded chromosomes revealed complete G-band homology between the autosomes of the two species. However, the X and Y chromosomes clearly differ in size and in C- and G-banding, being much larger in A. kaiseri. Comparison of the cytochrome b sequences places the separation between A. kaiseri and A. chrysophilus at 4.49 Mya, a period of intense speciation in other African muroids. The resulting phylogeny strongly supports the idea of a paraphyletic group, suggesting the need to elevate the previously described subgenera to the genus rank.

The diphyletic nature of the infraorder Thalassinidea (Decapoda, Pleocyemata) as derived from the morphology of the gastric mill

[A comparison of the gastric mill in some representative species of the Thalassinidea, indicates that the Thalassinidea are possibly diphyletic, consisting of the superfamily Callianassoidea Dana, 1852, including the families Callianassidae Dana, 1852, Axiidae Huxley, 1879, Callianideidae Kossmann, 1880, Ctenochelidae Manning & Felder, 1991, and Gourretiidae Sakai, 1999, in which the propyloric ossicle is simple and has a smooth posterior surface; and the superfamily Thalassinoidea Latreille, 1831, including the Thalassinidae Latreille, 1831, Upogebiidae Borradaile, 1903, and Laomediidae De Haan, 1849, in which the propyloric ossicle is triangularly protruded downwards and has a series of fine, transverse septa on its posterior face. Ein Vergleich der Kaumagen ausgewahlter Arten der Thalassinidea deutet an, dass die Thalassinidea moglicherweise diphyletisch sind. Sie bestehen aus zwei Uberfamilien, einerseits den Callianassioidea Dana, 1852, bei denen der propylorische Mittelzahn einfach und dessen Ruckseite glatt ist, andererseits den Thalassinoidea, bei denen dieser Zahn dreieckig vorragt und auf der Ruckseite transversale Septen tragt. Zu den Callianassoidea gehoren die Familien Callianassidae Dana, 1852, Axiidae Huxley, 1879, Callianideidae Kossmann, 1880, Ctenochelidae Manning & Felder, 1991 und Gourretiidae Sakai, 2004; zu den Thalassinoidea die Familien Thalassinidae Latreille, 1831, Upogebiidae Borradaile, 1903 und Laomediidae De Haan, 1849. , A comparison of the gastric mill in some representative species of the Thalassinidea, indicates that the Thalassinidea are possibly diphyletic, consisting of the superfamily Callianassoidea Dana, 1852, including the families Callianassidae Dana, 1852, Axiidae Huxley, 1879, Callianideidae Kossmann, 1880, Ctenochelidae Manning & Felder, 1991, and Gourretiidae Sakai, 1999, in which the propyloric ossicle is simple and has a smooth posterior surface; and the superfamily Thalassinoidea Latreille, 1831, including the Thalassinidae Latreille, 1831, Upogebiidae Borradaile, 1903, and Laomediidae De Haan, 1849, in which the propyloric ossicle is triangularly protruded downwards and has a series of fine, transverse septa on its posterior face. Ein Vergleich der Kaumagen ausgewahlter Arten der Thalassinidea deutet an, dass die Thalassinidea moglicherweise diphyletisch sind. Sie bestehen aus zwei Uberfamilien, einerseits den Callianassioidea Dana, 1852, bei denen der propylorische Mittelzahn einfach und dessen Ruckseite glatt ist, andererseits den Thalassinoidea, bei denen dieser Zahn dreieckig vorragt und auf der Ruckseite transversale Septen tragt. Zu den Callianassoidea gehoren die Familien Callianassidae Dana, 1852, Axiidae Huxley, 1879, Callianideidae Kossmann, 1880, Ctenochelidae Manning & Felder, 1991 und Gourretiidae Sakai, 2004; zu den Thalassinoidea die Familien Thalassinidae Latreille, 1831, Upogebiidae Borradaile, 1903 und Laomediidae De Haan, 1849. ]

The first-instar larva of the genus Arthropterus (Coleoptera : Carabidae : Paussinae): implications for evolution of myrmecophily and phylogenetic relationships within the subfamily

The first-instar larva of the genus Arthropterus W.S. MacLeay, 1838 is described and illustrated. This is the first description of a newly hatched (first-instar) larva in the myrmecophilous tribe Paussini and the first known larva from a basal paussine lineage. A phylogenetic analysis of the subfamily based on larval characters confirms the placement of the genus Arthropterus as a sister-group of the remaining Paussini and supports the hypothesis that Metriini is the sister-group of Ozaenini+Paussini, with 'Ozaenini' as a paraphyletic group. Within this phylogenetic framework, we reassess which larval characters are diagnostic of the tribes Paussini and Ozaenini. Several larval features of Arthropterus, such as the riddled sensilla S-VIII and the fused terminal disk, are interpreted as adaptations to myrmecophily. This interpretation supports the hypothesis that larvae have played an important role in the evolution of myrmecophily within the subfamily Paussinae.