In search of historical biogeographic patterns in the western Mediterranean terrestrial fauna

A cladistic biogeographic study of the western Mediterranean terrestrial fauna is made using taxon-area cladograms of ten groups of animals showing high levels of endemicity in the area. The groups analysed are the Nephrotoma flavescens group, the Tipula (Acutipula) maxima group, the T. (Lunatipula) bullata and falcata group, the subgenus T. (Mediotipula) , the T. (Savtshenkia) goriziensis group, the T. (S) signata group (Insecta, Diptera, Tipulidae), the Protonemura corsicana group (Insecta, Plecoptera, Nemouridae), the genus Speonemadus (Insecta, Coleoptera, Cholevidae), and the subgenera Triturus (Palaeotriton) and Triturus (Triturus) (Urodela, Salamandridae). The groups contain a total of 123 species and subspecies. Detailed distribution maps of 94 species and subspecies of Tipulidae included in the study are given. Based on the distributions of 74 endemic species and subspecies, 13 areas of endemism in the Mediterranean are recognized. The geology of the western Mediterranean since the late Oligocene is discussed with reference to a number of maps showing kinematic reconstructions of the area. Five methods for cladistic biogeographic purposes were employed, viz. Brooks Parsimony Analysis, Component Compatibility Analysis, Component Analysis, Three-Area Statements Analysis, and Paralogy-free Subtree Analysis. General area cladograms produced by computer implementations of the five methods show low levels of congruence. Geological area cladograms are fully compatible with some of the results of Brooks Parsimony Analysis only.     

CLADISTIC AND BIOGEOGRAPHIC ANALYSES OF THE WEEVIL GENUS LISTRODERES SCHOENHERR (COLEOPTERA: CURCULIONIDAE)

Abstract— The weevil genus Listroderes Schoenherr is a monophyletic group defined by the body vestiture consisting of subcircular to suboval scales, and comprises 35 species endemic to southern South America. A cladistic analysis of the genus was carried out using 44 characters from external morphology, body vestiture, and male and female genitalia. The curvipes (three species), nodifer (five species) and robustus (four species) species groups and the 23 species of the costirostris species group were considered terminal taxa. Apomorphic states were identified using the genus Hyperoides Marshall as outgroup. The analysis yielded 122 equally parsimonious cladograms, each with 89 steps and a consistency index of 0.42; a successive weighting procedure resulted in nine cladograms (consistency index of 0.69 and retention index of 0.85). In the general consensus cladogram, nodifer-robustus and curvipes-costirostris are two pairs of sister species groups. The costirostris group comprises the subgroups foveatus, angusticeps, costirostris, delaiguei, bimaculatus and elegans , in phylogenetic order. A distributional analysis of the species of Listroderes led to identification of four areas of endemism, namely central Chile, sub-Antarctic, central Argentina and Chaco. A vicariance biogeographic analysis of these areas, based on area cladograms of Listroderes, Hyperoides, Naupactus taeniatulus species group (Coleoptera: Curculionidae), and Lucilia generic group (Asteraceae) was carried out applying the three-area statements technique. According to the general area cladogram the sequence of area-fragmentation is as follows: (central Chile (sub-Antarctic (central Argentina, Chaco))). This sequence is congruent with the history hypothesized for the region, where the uplift of the Andes (Oligocene-Pliocene) led to a progressive aridification, replacing the ancient subtropical forest by open-country environments.     

A Review of Current Theories and Methods in Cladistics and a Cladistic Study of Twelve Lindera Species in Eastern China

Cladistics has become a widely used method for phylogenetic reconstruction．Because of rapid improvement Of cladistic theories and methodologies,and application of new data,especially,molecular data,it is becoming realistic to reconstruct phylogenies of organisms,and to establish natural classifications based on these phylogenies．This paper reviews some current cladistic theories and methods in a practical way,such as choosing characters,defining character states,polarizing characters,analyzing data matrices, calculating consensus cladograms,choosing among multiple equally most parsimonious cladograms,estimating reliability of cladograms,and applying cladograms to classification, character evolution,and biogeography． Based on 36 morphological characters．a parsimony analysis of 12 species representing six sections in subgenus Lindera and an outgroup species from subgenus lteodaphne of the genus Lindera(Lauraceae)was conducted．The results suggest a close relationship between section Lindera and section Sphaerocarpae,which is different from the previous phylogenetic hypothesis within the genus．In the strict consensus cladogram,two species,L．megaphylla and L.chienii,from section Cupuliformes are in the most primitive and the most advanced clades respectively,indicating that the section is polyphyletic．The cladogram also suggests that section Lindera be a polyphyletic group．     

分支系统学当前的理论和方法概述及华东地区山胡椒属十二个种的分支系统学研究

本文概述了当前分支系统学研究中涉及的主要理论和方法,包括性状的选取、性状状态和极性的确定、数据矩阵的分析计算、结果分支图的处理、分支图可靠性的评价及分支图的应用。本文同时以华东地区樟科山胡椒属Linderal2个种的分支系统学研究为例,讨论了用形态性状进行分支系统学研究中可能遇到的问题,也揭示了一些分支系统学与传统的系统学在应用性状推导进化关系上的不同点。对这12个种的分支系统学研究得出了一些不同于传统系统学方法所推测的山胡椒属内的系统发育关系,如分支系统学研究显示山胡椒组和球果组很近缘。在严格一致性分支图上,杯托组的黑壳楠和江浙山胡椒分别位于最原始和最进化的分支,表明这个组是复系类群。分支图也显示山胡椒组可能是复系类群。     

Cladistic analysis of Enkianthus with notes on the early diversification of the Ericaceae

The genus Enkianthus (Ericaceae) has been subject to a cladistic analysis of morphological, anatomical, embryological, and cytological data, using two species of Clethra as outgroup. In order to evaluate the monophyly of Enkianthus , the genera Epigaea and Phyllodoce from the subfamily Rhododendroideae, and the genus Andromeda from the subfamily Vaccinioideae were also included in an initial analysis which yielded 20 equally parsimonious cladograms. The results indicated that Enkianthus is monophyletic, and a subsequent analysis with only the outgroup taxa and 14 species of Enkianthus yielded two equally parsimonious cladograms with a more resolved topology. In order to obtain strictly monophyletic taxa, the current subgeneric classification of Enkianthus is revised, recognizing four sections: Enkianthus (9 spp.), Andromedina (2 spp.), Meisteria (2 spp.), and Enkiantella (4 spp.). The paper provides a key, illustrations of representative taxa, a cladogram, and strict consensus trees.     

Two new species and a phylogenetic analysis of the genus Tectidrilus (0Iigochaeta, Tubifcidae)

Tectidrilus probus sp.n. and Tectidrilus profusus sp.n. are described from offshore localities in southern California. Both species appear closely related to T. diversus Ersëus, 1982, which occurs in the same area, but they differ from the latter by possessing small oesophageal diverticula; T. probus differs also by its very small spermathecal vestibules. Tectidrilus profusus is a highly apomorphic form with elaborate, partly muscular, male ducts and spermathecae. Monophyly of Tectidrilus is supported by the short clitellum and the unisetal 'bundles' in postclitellar segments. Patterns of apomorphic character states within the genus are analyzed under the principle of Camin-Sokal parsimony (irreversibility of character states), giving three equally parsimonious cladograms. In the strict consensus tree of these cladograms, eight of the twelve members of Tectidrilus form an apomorphic group defined by the papillated body wall, but the individual species within this group cannot be unequivocally resolved from each other.     

Phylogenetic revision of the Taenogera Kröber genus-group (Diptera: Therevidae), with descriptions of two new genera

The phylogenetic relationships of the Taenogera genus-group, consisting of nine genera, are examined using cladistic methods. Twenty-one species representing nine genera and one outgroup were compared in a cladistic analysis across 99 states in 44 characters. The genus Taenogera Kröber is revised to contain only T. longa (Schiner), T. nitida (Macquart) and T. notatithorax Mann. Two new genera are described and figured: Actenomeros gen. n., with two species, A. corniculaticaudus sp. n. and A. onyx sp. n.; and Taenogerella gen. n., with four species, Ta. elizabethae sp. n., Ta. platina sp. n., Ta. schlingeri sp. n. and Ta . nigrapicalis (Mann) comb. n. Phylogenetic analysis also supports the removal of Nanexila gracilis (Mann) comb. n. from Taenogera .     

A cladistic analysis of the genus Lopezia (Onagraceae)

Lopezia, a genus of 22 species largely restricted to Mexico, forms a monophyletic group defined by the possession of two stamens in tetramerous flowerS. A cladistic analysis of the genus was performed using 16 characters from morphology, anatomy and embryology. Twenty of the 22 species were considered the terminal taxa. Polarity of the characters was based on the outgroup comparison method, using three alternative outgroups: the genus Fuchsia, Onagraceae excluding Ludwigia, and Epilobieae-Onagreae. Using Fuchsia as outgroup, 29 equally parsimonious cladograms were produced, each with 24 steps and a consistency index of 0.75. A successive weighting procedure was applied, resulting in 15 cladograms with consistency index of 0.85. The strict consensus cladogram defines eight monophyletic groups and supports most of the current sectional classification of Lopezia, with the exception that no synapomorphy defines section Jehlia. The larger sections Lopezia and Pelozia, although well-defined as clades, are not fully resolved internally. All cladograms support section Riesenbachia as monophyletic. The two alternative hypotheses for outgroups produced similar results: 61 most parsimonious trees, reduced after the successive weighting procedure to 15, which are identical to those produced with Fuchsia as outgroup. These results are discussed in the context of data on cytology and pollination biology.     

A CLADISTIC ANALYSIS OF CIRCAEA (ONAGRACEAE)

Abstract— Circaea (Onagraceae), a genus of seven species from the northern hemisphere, forms a monophylctic group defined by the following synapomorphies: two stamens (opposite the sepals), fruits indehiscent capsules, hooked hairs on the fruits, and perianth parts reduced to two. A cladistic analysis of the genus was performed using 22 characters from morphology, anatomy and palynology. The seven species and seven additional subspecies were considered the terminal taxa. Polarity of the characters is based on the outgroup comparison method. Eighteen equally parsimonious cladograms were produced, each with 32 steps and a consistency index of 0.75. A successive weighting procedure was applied, resulting in six cladograms with a consistency index of 0.95. All 24 cladograms share the following seven monophylctic groups: (1) all species except C. cordala and C. glabrescens (stipules deciduous); (2) the group of C. mollis , the three subspecies of C. lutetiana and C. erubescens (exsertcd nectary); (3) C. repens and all six subspecies of C. alpina (single-seeded fruits, stolons terminated by tubers, and viscin threads reduced or absent); (4) the six subspecies of C. alpina (unilocular ovaries with no trace of a second loculc and tardily opening perianth); (5) C. alpina subspecies alpina, micrantha, pacifica , and imaicola (flowers opening on ascending to erect pedicels); (6) C. alpina subspecies alpina, micrantha , and pacifica (translucent leaves); (7) C. alpina subspecies alpina and murantha (glabrous stems). Results of the cladistic analysis support most of the current systematic classification of the genus, except for the relationships among the three subspecies of C. lutetiana .     

Phylogeny of the subfamily Chauliodinae (Megaloptera: Corydalidae), with description of a new genus from the Oriental Realm

Abstract.  A new Oriental fishfly genus, Sinochauliodes gen.n. , is described, including four species: S. fujianensis ( Yang & Yang, 1999 ) comb.n. , S. griseus ( Yang & Yang, 1999 ) comb.n. , S. maculosus sp.n. and S. squalidus sp.n. A cladistic analysis based on adult morphological characters clarified the phylogenetic status of the new genus and allowed the reconstruction of the intergeneric relationships of the subfamily Chauliodinae. Two main clades within Chauliodinae were recognized from the cladistic analysis. The Asian fishflies, together with the two Nearctic genera, Chauliodes and Nigronia , formed a monophyletic lineage, and the new genus was assigned as the sister group to the genus Parachauliodes . The biogeography of the Asian fishflies is discussed.     

Cladistic analysis and biogeography of the cicadas of the Indo-Pacific subtribe Cosmopsaltriina (Hemiptera: Cicadoidea: Cicadidae)

Abstract A consensus cladogram is presented for ninety-seven of the ninety-nine (sub)species of the cicada subtribe Cosmopsaltriina on the basis of a cladistic analysis of forty-nine characters. The consensus cladogram confirms the monophyly of the eight genera constituting the subtribe, and the monophyly of the subtribe as a whole. Cosmopsaltriina occur in Sulawesi, the Moluccas, New Guinea, Cape York, the Solomon Islands, Vanuatu, Fiji, Tonga and Samoa. The subtribe demonstrates a high rate of endemism on genus, species group and species level. The taxon-area and consensus-area cladograms resulting from Brooks parsimony analysis are discussed in relation to the palaeogeography of the area.     

A Cladistic Analysis of the Genus Cyttaria (Fungi-Ascomycotina)*

Abstract—Cyttaria Berkeley (Cyttariaceae, Cyttariales, Class Discomycetes), is a genus of eleven species, seven from South America and four from Australasia (Australia, Tasmania, and New Zealand). Cyttaria is a monophyletic genus defined by the following synapomorphies: fleshy to gelatinous stromata with endostromatic apothecia; and complete lack of fungal chitin in the cell walls, having instead β-1–3-glucan. All Cyttaria are exclusive parasites of Nothofagus species. A cladistic analysis of the genus was performed using 18 characters from macromorphology, micro-morphology, and phenology. Polarity of characters is based on the outgroup comparison method (using the Class Discomycetes as a whole) and on the ontogenetic criterion. A hypothetical outgroup was constructed using all plesiomorphic states. Two equally parsimonious cladograms were produced, each with 40 steps and a consistency index of 0.70. These differed in the position of the South American species C. hookeri and C. johowii. In one cladogram, C. hookeri is the sister group to the rest of the genus, and in the other, both species form a monophyletic group (ascospores ovoid) that is the sister group of the rest of the genus. Our analysis of the two characters causing this difference (position of apothecia on stroma, and shape of ascospores) supports the latter hypothesis. In both cladograms, all of the Australasian species form a monophyletic group (thick ectostroma), and, within that, two subgroups are defined: C. gunnii-C. pallida (papillae present), and C. nigra-C. septentrionalis (ectostroma with black incrustations). The Australasian species form a monophyletic group (spermatangia absent) with C. berteroi , and this last group forms a monophyletic group (conidia absent) with C. espinosae. Cytlaria darwimi and C. exigua form another monophyletic group (very thick ectostroma). Excluding C. hookeri and C. johowii, C. hariotii is the sister group to the remaining eight species.     

Phylogenetic systematics and evolution of Agnotecous in New Caledonia (Orthoptera: Grylloidea, Eneopteridae)

Abstract. Within a framework for historical analysis of Eneopterinae biogeography the New Caledonian endemic cricket genus Agnotecous Saussure, 1878 is revised: the eight already known species are diagnosed and six new species described, A. azurensis Desutter‐Grandcolas sp.n. , A. chopardi Desutter‐Grandcolas sp.n. , A. clarus Desutter‐Grandcolas sp.n. , A. doensis Desutter‐Grandcolas sp.n. , A. meridionalis Desutter‐Grandcolas sp.n. and A. occidentalis Desutter‐Grandcolas sp.n. Four species groups are characterized by male genitalic structures. Identification keys are provided for both males and females. A cladistic analysis was performed using fifty‐eight morphological characters. The two resultant topologies, which differ only in topology of three apical species, support the monophyly of Agnotecous and that of the four species groups. Preliminary hypotheses of Eneopterinae historical biogeography are derived from phylogenetic and distributional data.     

Cladistic analysis of Neuroptera and their systematic position within Neuropterida (Insecta: Holometabola: Neuropterida: Neuroptera)

A phylogenetic analysis of Neuroptera using thirty‐six predominantly morphological characters of adults and larvae is presented. This is the first computerized cladistic analysis at the ordinal level. It included nineteen species representing seventeen families of Neuroptera, three species representing two families (Sialidae and both subfamilies of Corydalidae) of Megaloptera, two species representing two families of Raphidioptera and as prime outgroup one species of a family of Coleoptera. Ten equally most parsimonious cladograms were found, of which one is selected and presented in detail. The results are discussed in light of recent results from mental phylogenetic cladograms. The suborders Nevrorthi‐ formia, Myrmeleontiformia and Hemerobiiformia received strong support, however Nevrorthiformia formed the adelphotaxon of Myrmeleontiformia + Hemerobiiformia (former sister group of Myrmeleontiformia only). In Myrmeleontiformia, the sister‐group relationships between Psychopsidae + Nemopteridae and Nymphidae + (Myrmeleontidae + Ascalaphidae) are corroborated. In Hemerobiiformia, Ithonidae + Polystoechotidae is confirmed as the sister group of the remaining families. Dilaridae + (Mantispidae + (Rhachiberothidae + Berothidae)), which has already been proposed, is confirmed. Chrysopidae + Osmylidae emerged as the sister group of a clade comprising Hemerobiidae + ((Coniopterygidae + Sisyridae) + (dilarid clade)). Despite the sister‐group relationship of Coniopterygidae + Sisyridae being only weakly supported, the position of Coniopterygidae within the higher Hemerobiiformia is corroborated. At the ordinal level, the analysis provided clear support for the hypothesis that Megaloptera + Neuroptera are sister groups, which upsets the conventional Megaloptera + Raphidioptera hypothesis.     

Revision of the genus Pseudabris Fairmaire (Coleoptera,Meloidae), an endemic to the Tibetan Plateau,with biogeographical comments

The mylabrine genus Pseudabris is endemic to the Tibetan Plateau and includes seven species with overlapping or disjunct ranges. The genus is revised: three new species, P. brevipilosa sp.n., P. latimaculata sp.n., P. regularis sp.n., are described and illustrated, and a key to the species is provided. Molecular evidence supports the placement of Pseudabris within the tribe Mylabrini. Results of a morphology‐based cladistic analysis support the existence of two lineages, one centred mainly on the south central plateau, and the second in the eastern area. Faunistics and bionomics of the genus are summarized, focusing on phenology, elevation, habitat preference and host plants. The endemism of the Tibetan Plateau is discussed, with a special focus on the genus Pseudabris.     

Taxonomic revision and phylogenetic relationships of the bee genus Parapsaenythia Friese (Hymenoptera,Apidae, Protandrenini), with biogeographic inferences for the South American Chacoan Subregion

The bee genus Parapsaenythia Friese (Apidae, Protandrenini) is restricted to South America and characterized by the strongly punctate integument and hairy eyes. In the present work, the species of Parapsaenythia are revised, with seven species recognized, of which three are described as new: P. carinulata sp.n. (Argentina, Brazil, Paraguay), P. inornata Moure (Brazil), P. lanata sp.n. (Brazil), P. paspali (Schrottky) (Argentina, Brazil, Paraguay), P. puncticutis (Vachal) (Argentina, Bolivia), P. scutellaris sp.n. (Argentina, Bolivia, Brazil, Paraguay) and P. serripes (Ducke) (Argentina, Brazil). Lectotypes are designated for Psaenythia (Parapsaenythia) argentina Friese, Anthrenoides paspali Schrottky and Parapsaenythia paraguayae Brèthes. Floral records, distribution maps, illustrations and an identification key are provided. The cladistic analysis of 11 terminal species (all species of Parapsaenythia plus four outgroup species), based on 45 characters of external morphology and male terminalia, produced a single most parsimonious tree. Parapsaenythia was recovered as monophyletic, with the relationships among its species as follows: (P. inornata (P. carinulata sp.n. (P. lanata sp.n. (P. scutellaris sp.n. (P. serripes, P. paspali, P. puncticutis))))). Based on these relationships plus the available distributional data, a cladistic biogeographic analysis was performed, and area cladograms for the Chacoan subregion are presented and discussed.     

Revision of the Neotropical genus Chimarrhodella Lestage (Trichoptera: Philopotamidae)

Abstract. A revision and phylogenetic analysis of the Neotropical genus Chimarrhodella Lestage is presented. Four described species are recognized, C.galeata (Martynov), C.peruviana (Ross), C.ulmeri (Ross) and C.nigra Flint. In addition, five new species are described, C.costaricensis sp.n. (Costa Rica), C.tapanti sp.n. (Costa Rica), C.flinti sp.n. (Venezuela), C.pilcopata , sp.n. (Peru) and C.tobagoensis sp.n. (Tobago). These species are organized into two monophyletic species groups, designated here the Galeata Group and the Peruviana Group. The Galeata Group is characterized by having male genitalia with tergum X possessing well developed lateral lobes bearing sclerotized, spine-like processes and includes C.galeata, C.costaricensis, C.flinti, C.pilcopata and C.tapanti. The Peruviana Group is characterized by having male genitalia with tergum X only slightly divided apically, tergum IX bearing hook-like processes posteriorly, and female genitalia with segment VIII much elongated in comparison to the Galeata Group and includes C.peruviana, C.nigra, C.tobagoensis and C.ulmeri. Records of the genus from Costa Rica and Tobago represent significant extensions of the known range for the genus, previously unknown north of Panama. A key to males and known females and a species-level phylogenetic analysis are included.     

Phylogenetic analysis of a group of palaeonemerteans (Nemertea) including two new species from Queensland and the Great Barrier Reef, Australia

Sundberg, P., Gibson, R. & Olsson, U. (2003). Phylogenetic analysis of a group of palaeonemerteans (Nemertea) including two new species from Queensland and the Great Barrier Reef, Australia. — Zoologica Scripta, 32, 279–296.
Based on 18S rDNA nucleotide sequences and morphological characters, we reconstruct the phylogeny for a group of palaeonemerteans estimated to be monophyletic. Two new palaeonemertean species from Queensland and the Great Barrier Reef, Australia are included in the phylogenetic analysis. The results confirm that one of the species, Cephalothrix queenslandica sp. n., is part of the Cephalothrix–Cephalotrichella–Procephalothrix group. These genera are redefined phylogenetically under the name Cephalothrix based on the cladistic analysis. The other species, Balionemertes australiensis gen. et sp. n., is placed in a new genus which forms a sister taxon to Cephalothrix . The morphology of both new species is described in detail.     

Systematics and biogeography of the fishfly genus Parachauliodes (Megaloptera: Corydalidae) endemic to the east Asian islands

Abstract.  The fishfly genus Parachauliodes van der Weele, 1909 comprises six species from Taiwan, Japan, and South Korea, ranging along the east Asian islands. Herein, all the Parachauliodes species are described, including two new species, namely P. asahinai sp.n. and P. niger sp.n . A cladistic analysis based on adult and larval morphological characters was conducted to reconstruct the interspecific phylogeny of the Parachauliodes species. Two main clades within Parachauliodes were obtained from the cladistic analysis, namely ' P. asahinai  + P. continentalis ' and '( P. japonicus  +  P. yanbaru ) + ( P. nebulosus  +  P. niger )'. Based on the present interspecific phylogeny and distribution patterns of Parachauliodes , its ancestor might have invaded the east Asian islands by a land bridge from Taiwan via Ryukyu to the Japanese archipelago about 10−5 million years ago.     

A generic revision of the Phallodrilinae (Oligochaeta, Tubificidae)

The generic classification of the subfamily Phallodrilinae (with a total of 252 species) is revised according to phylogenetic principles. Morphological character patterns are assessed under the principle of maximum parsimony, but due to the poor resolution and instability in the resulting cladograms, the analysis is supplemented with intuitive weighting of synapomorphic similarities. Thirty-one genera are recognized. The former definition of the type genus Phallodrilus Picrantoni, 1902, is revised and the members of this genus are reclassified into 20 taxa, each hypothesized to be monophyletic: Phallodrilus s.str . (revised to contain the type species only), Somalidrilus gcn.n., Milliganius gen.n., Phallodriloides gen.n., Albanidrilus gen.n., Abyssidrilus gen.n., Paraktedrilus gen.n., Uniporodrilus Erséus, 1979, Inermidrilus gen.n., Gianius gen.n., Mexidrilus gen.n., Peosidrilus Baker & Erséus, 1979, Atlantidrilus Erséus, 1983, Thalassodrilus Brinkhurst, 1963, Pirodrilus gen.n., Pirodriloides gen.n., Marionidrilus gen.n., Pectinodrilus gen.n., Pacifidrilus gen.n. and Pseudospiridion gen.n. Nine new species are described from the Northwest Atlantic: Phallodriloides pinnulatus sp.n., Abyssidrilus opulentus sp.n., A. potens sp.n., A. altoides sp.n., Mexidrilus obtusus sp.n., Peosidrilus dalei sp.n., P. aduncus sp.n., P. riseri sp.n., and Atlantidrilus hamulus sp.n. The genus Discordiprostatus Baker, 1982, is synonymized with Nootkadrilus Baker, 1982.