Area Cladograms Research Articles

Molecular Phylogenetics and Biogeography of Tanagers in the GenusRamphocelus(Aves)

Ramphocelustanagers are distributed throughout the Neotropical lowlands. In this paper, mitochrondrial DNA (mtDNA) sequence data from cytochrome b and ND2 genes are used to estimate relationships among seven of nine species of the avian genusRamphocelus.Genetic differentiation is high betweenRamphocelus passerinii passeriniiandRamphocelus passerinii costaricensis,and the two subspecies are diagnosable and distinct from one another both morphologically and genetically. Thus, elevation to species status is recommended. Three clades are supported by both gene sequences; one clade containsR. passerinii, R. costaricensis,andR. icteronotus;the second clade containsRamphocelus carbo, Ramphocelus bresilius,andRamphocelus nigrogularis;the third clade containsRamphocelus sanguinolentus.The degree of saturation was assessed for both genes and saturation of third position of codons occurs by 10–12% uncorrected pairwise sequence divergence. The general area cladogram suggests the following area relationships: Pacific and Caribbean Central America are sister areas, Choco is the sister to the Central American area, and Amazonia/southeastern Brazil is the outgroup area to the Choco/Central American clade.

Ichthyogeography of the Guinea–Congo rain forest, West Africa

SynopsisIchthyogeography is the section of biogeography which seeks to interpret fish biodiversity in terms of present and past distributions and abundance. It can also have practical applications in fisheries management and conservation. There are thought to be more than one thousand species of fish in the 40 or so major rivers of the Guinea–Congo rain forest region, with correspondingly high levels of endemicity. In the Congo or Zaire river basin alone, the proportion of endemic fish species may exceed 80% (from a total of >690), but many more taxonomic and distributional data remain to be gathered.Unlike terrestrial vertebrates, the diverse freshwater fishes of Guinea–Congo evidently have distributions which are closely confined by hydrography. Hence, fish distributions may be of particular help in corroborating or refuting postulated geological events, patterns and processes and in explaining associated aspects of rainforest evolution. From the 19th century until now, the ichthyogeography of Guinea–Congo has been diagnosed largely in terms of presumed post-Miocene geological sequences of fish taxa and their past and recent dispersal in relation to particular hydrological conditions: mainly riverine volume discharge, salinity and temperature. From this, the fish fauna is, by convention, divided into ‘provinces’ established on endemism, palaeogeography and supposed physical or ecological barriers to dispersal. However, in this paper it is argued that the traditional ichthyogeographical accounts which highlight endemism and dispersal are generally flawed. It is argued here that historical patterns of fish distribution can only be fully understood if phyletic (cladistic) data are taken into consideration. While Upper and Lower Guinea and the Zaïre basin may be defined in part on the basis of endemism there is a lack of taxonomic and distributional evidence to show that Guinea–Congo is itself a cohesive ichthyogeographical unit.There is clearly a need for comparisons with fish distributions outside the rain forest zone of Guinea–Congo. African inter-provincial, trans-continental and inter-continental comparisons reveal distribution patterns which may relate more to pre-Miocene rather than post-Miocene geology or present-day rain forest ecology. Continental drift, notably between Africa and South America, probably led to the separation 85 million years ago of previously united fish populations. This may account for recent higher-level phyletic correspondences between the separate rain forest fish faunas of Africa and the neotropics.Last, the so-called ‘marine intrusive’ fishes – which are normally excluded from zoogeographical consideration – merit a careful re-evaluation. While they may be regarded as an inconvenience in developing scientific hypotheses, such intrusives can comprise a remarkable 30% or more of the riverine fish fauna in Guinea–Congo. It seems that the widely accepted ecological divisions between marine and ‘primary freshwater’ fishes are not as clear-cut as has been supposed. In addition, zoogeographically critical marine, trans-Atlantic, phyletic relationships apparently exist. These are probably best interpreted by using area cladograms in the context of ocean basin development, rather than by referring solely to marine fish dispersal and the traditional continental and provincial ichthyogeography of Guinea–Congo.

Trilobite biogeography in the Middle Devonian: geological processes and analytical methods

Phylogenetic patterns of trilobite clades were used to deduce biogeographic patterns during the Middle Devonian, a time of active plate collision between North America (Laurentia) and other plates, coincident with several major episodes of sea-level rise and fall. The mapping of biogeographic states onto phylogenies for asteropyginid and proetid trilobites indicated that during their history these trilobite clades often shifted the areas they occupied, and also underwent vicariant differentiation, followed by range expansion, followed by subsequent vicariance. Biogeographic patterns in these individual phylogenies were evaluated and synthesized using a modified version of Brooks Parsimony Analysis, which is discussed. This method makes it possible using cladistic methods to distinguish between episodes of vicariance and episodes of dispersal. Two types of dispersal are recognized herein: (1) the individualistic responses of certain taxa in a single clade that cannot be generalized, i.e., traditional ad hoc dispersal, and (2) those patterns of congruent range expansion that are replicated across several clades. The latter are not treated as true dispersal, expansion of a taxon's range over a barrier accompanied by diversification, but rather as a result of the temporary removal of barriers to marine taxa, due either to relative sea-level rise or to the collision of formerly disjunct plates. These are interpreted as changes in the structure of areas, and this type of dispersal is referred to as geo-dispersal. Geo-dispersal was found to have occurred in the Middle Devonian trilobite fauna of Eastern North America.Biogeographic analysis indicated that Eastern North America is a strongly supported area, with the Appalachian and Michigan Basins as sister areas. Armorica and the Canadian Arctic are also sister areas. Congruence was found between area cladograms produced by vicariance and dispersal analyses for Middle Devonian trilobites, suggesting that in some cases the geological processes governing vicariance, such as sea-level changes, were the same as those that caused dispersal.

A preliminary phylogenetic and biogeographic analysis of the dyakiidae (Gastropoda: Stylommatophora) and a biogeographic analysis of other sundaland taxa

The phylogeny of the Dyakiidae, an early diverging lineage of the Limacoidea sensu lalo, is reconstructed. In the Dyakiidae a unique transformation series of different genital accessory organs, the so-called “stimulators”, is preserved which is an important argument for the hypothesis that the different stimulator types of the Stylommatophora are homologous. The biogeographic distribution patterns of the genera of the Dyakiidae are summarized and the ancestral areas of the major clades are analysed. The analysis of the area cladograms of the Dyakiidae and of several butterfly and heteroptera genera occurring in Sundaland revealed neither a general pattern of relationship between the areas of endemism in Sundaland, nor a general pattern of relationship between Sundaland and other areas. The area cladogram of the rather immobile Dyakiidae endemic to Sundaland might reflect older events than the area cladograms of the more mobile and widespread butterfly and plant bug genera. The general incompatibility of the area cladograms involving Sundaland taxa suggests that dispersal across barriers has played a major role in the historical biogeography of the analysed groups and challenges the hypothesis of vicariance biogeography that the distribution patterns of organisms are largely due to the fragmentation of an ancestral biota.

A PRELIMINARY PHYLOGENETIC AND BIOGEOGRAPHIC ANALYSIS OF THE DYAKIIDAE (GASTROPODA: STYLOMMATOPHORA) AND A BIOGEOGRAPHIC ANALYSIS OF OTHER SUNDALAND TAXA.

- The phylogeny of the Dyakiidae, an early diverging lineage of the Limacoidea sensu lalo, is reconstructed. In the Dyakiidae a unique transformation series of different genital accessory organs, the so-called "stimulators", is preserved which is an important argument for the hypothesis that the different stimulator types of the Stylommatophora are homologous. The biogeographic distribution patterns of the genera of the Dyakiidae are summarized and the ancestral areas of the major clades are analysed. The analysis of the area cladograms of the Dyakiidae and of several butterfly and heteroptera genera occurring in Sundaland revealed neither a general pattern of relationship between the areas of endemism in Sundaland, nor a general pattern of relationship between Sundaland and other areas. The area cladogram of the rather immobile Dyakiidae endemic to Sundaland might reflect older events than the area cladograms of the more mobile and widespread butterfly and plant bug genera. The general incompatibility of the area cladograms involving Sundaland taxa suggests that dispersal across barriers has played a major role in the historical biogeography of the analysed groups and challenges the hypothesis of vicariance biogeography that the distribution patterns of organisms are largely due to the fragmentation of an ancestral biota.

Cladistic Biogeography of Plants in Australia and New Guinea: Congruent Pattern Reveals Two Endemic Tropical Tracks

?Although congruence among multiple taxa is accepted as the basis of the cladistic approach to biogeography, seldom has congruence been tested for patterns within continents. Moreover, contiguous land areas are subject to confounding factors that might lead to incongru? ence, and areas of endemism are difficult to define. Most previous studies have analyzed single taxa and therefore could not test for congruence. Here, we report the first cladistic biogeographic analysis of multiple plant taxa (ll angiosperm groups) within continental Australasia. We have used Page's (1994, Syst. Biol. 43:58-77) new mapping method implemented in his program COM? PONENT 2.0, minimizing the optimality criterion leaves added, which is the number of terminal branches that need to be added to reconcile all input cladograms with the general area cladogram. A randomization test showed significant congruence, and a jackknife test indicated which parts of the general area pattern are robust. Conflicting patterns shown by widespread species may have been influenced by recant range expansions. At least two independent histories are postu? lated for taxa endemic to the tropics. One appears as a close relationship between the monsoon tropical areas, including southern New Guinea, and the other is a track (a group of areas with a common history) of successive differentiation along the east and south coasts. The Australian wet tropics (the Atherton area), although geographically proximate to the monsoon tropics, are part of the east coast track. Tasmania shows a very early vicariance from the rest of the continent, and the central arid areas appear related to adjacent coastal areas rather than to each other (contra Cracraft, 1991, Aust. Syst. Bot. 4:211-227). The postulated exchange of taxa with southeast Asia following late Miocene contact was not tested because of a lack of suitable taxon cladograms. [Cladistic biogeography; congruence; tree mapping; Australia; New Guinea; angiosperms; trop? ics.1 Congruence is the essence of cladistic biogeography. If general area patterns ex? ist, reflecting a common history among multiple, distantly related taxa, then at? tempts to reconstruct that history are jus? tified. However, congruence cannot be as? sumed a priori; different taxa may respond differently to earth events, depending upon their varying vagility or response to the appearance of barriers to dispersal. A bar? rier to one species may be a suitable hab? itat for another. For example, black soil plains would be a barrier to both the plant genus Persoonia and the bird genus Platycercus, whereas calcareous plains support? ing forest would be a barrier only to Per? soonia. Thus, a vicariance event for one taxon may not be a vicariance event for an4 E-mail: mike.crisp@anu.edu.au. other, depending upon their differing eco? logical tolerances. Neither can congruence be expected for areas having multiple his? tories involving different taxa. Even if con? gruence is demonstrated, no single causal explanation, e.g., vicariance, can be as? sumed (Page, 1988). Nonetheless, congruent patterns have been demonstrated among taxa occurring on different continents, at least in the southern hemisphere (Brundin, 1965; Ro? sen, 1978; Edmunds, 1981; Humphries and Parenti, 1986; Weston and Crisp, 1994; Un? der and Crisp, 1996), and these patterns have usually been explained by the plate tectonic theory of the breakup of the supercontinent Gondwana during the last 160 million years, causing vicariance of widespread ancestral species. However, con? gruence at this level is sometimes viewed as trivial; the present continents are sepa-

HISTORICAL BIOGEOGRAPHY: Introduction to Methods

The five basic historical biogeographic methods are: dispersalism, phyloge­ netic biogeography, panbiogeography, cladistic biogeography, and parsimony analysis of endemicity. Dispersalism derives from the traditional concepts of center of origin and dispersal. Bremer's recent cladistic implementation of dispersalism estimates the relative probability that different areas were part of the ancestral distribution of a group. Phylogenetic biogeography applies the rules of progression and deviation to elucidate the history of the geographical distribution ofa group. Panbiogeography consists of plotting distributions of different taxa on maps, connecting their distribution areas together with lines called individual tracks, and looking for coincidence among individual tracks to determine generalized tracks. Generalized tracks indicate the preexistence of widespread ancestral biotas, subsequently fragmented by geological or climatic changes. Cladistic biogeography assumes a correspondence between taxonomic relationships and area relationships, where comparisons between area cladograms derived from different taxa allow one to obtain general area cladograms. The most important cladistic biogeographic procedures are: com­ ponent analysis, Brooks parsimony analysis, three-area statements, and recon­ ciled trees. Parsimony analysis of endemicity (PAE) classifies areas by their shared taxa, analogous to characters, according to the most parsimonious solution. We think the various methods are not mutually-exclusive alternatives, but some of them can· be integrated in a single biogeographic approach, with the capability of resolving different problems, such as the recognition of spatial

Historical biogeography of the holarctic: Area relationships, ancestral areas, and dispersal of non-marine animals

Based on published phylogenies for 73 groups of Holarctic non-marine animals, interrelationships between the four Holarctic infraregions (western and eastern Nearctic, western and eastern Palearctic) are examined. The study includes analysis of resolved area cladograms, ancestral areas and dispersal indicated by cladistic subordinateness. Area relationships reflecting present continental configurations (Nearctic vs. Palearctic) dominate the material to the extent that one might speak of a general Holarctic area pattern. Paleocontinental (western Nearctic+eastern Palearctic, western Palearctic+eastern Nearctic) and disjunct patterns are relatively more frequent among groups of higher taxonomic rank. The western Nearctic seems to have played a bigger role than the other infraregions as a center of origin. Two computer programs for constructing resolved area cladograms, viz., COMPONENT 1.5 and COMPONENT 2.0, are compared. The three standard assumptions for biogeographical analysis are compared and arguments are presented in favour of Assumption 0.

HISTORICAL BIOGEOGRAPHY OF THE HOLARCTIC: AREA RELATIONSHIPS, ANCESTRAL AREAS, AND DISPERSAL OF NON‐MARINE ANIMALS

Abstract —Based on published phylogenies for 73 groups of Holarctic non‐marine animals, interrelationships between the four Holarctic infraregions (western and eastern Nearctic, western and eastern Palearctic) are examined. The study includes analysis of resolved area cladograms, ancestral areas and dispersal indicated by cladistic subordinateness. Area relationships reflecting present continental configurations (Nearctic vs. Palearctic) dominate the material to the extent that one might speak of a general Holarctic area pattern. Paleocontinental (western Nearctic+eastern Palearctic, western Palearctic+eastern Nearctic) and disjunct patterns are relatively more frequent among groups of higher taxonomic rank. The western Nearctic seems to have played a bigger role than the other infraregions as a center of origin. Two computer programs for constructing resolved area cladograms, viz., COMPONENT 1.5 and COMPONENT 2.0, are compared. The three standard assumptions for biogeographical analysis are compared and arguments are presented in favour of Assumption 0.

Molecules, Morphology, and Area Cladograms: A Cladistic and Biogeographic Analysis of Gambusia (Teleostei: Poeciliidae)

Researchers investigating historical biogeography rely on taxon cladograms to infer the relationships of different areas of endemism. Unfortunately, systematists are often faced with many equally parsimonious taxon cladograms from a single data set or conflicting phylogenetic hypotheses from independent data sets. We present an analysis of congruence between two con? flicting phylogenetic hypotheses of the poeciliid fish genus Gambusia, one based on mitochondrial DNA sequences and the other based on morphological characters. We explore how different phy? logenetic hypotheses alter interpretations of area relationships and propose a set of area relation? ships for the genus. Furthermore, we compared the area relationships depicted for Gambusia with that of two Middle American fish genera (Xiphophorus and Heterandria) with similar distributional limits. Our analysis revealed areas of congruence among taxa from nuclear Central America but areas of incongruence among taxa from the Panuco basin and North American components. We discuss the implications of our findings in light of conventional hypotheses regarding Caribbean biogeography. This study illustrates the importance of considering alternative phylogenetic hy? potheses fully before attempting to interpret the biogeographic history of a taxon or region. (His? torical biogeography; Caribbean biogeography; phylogeny; Gambusia; Poeciliidae; congruence; area cladograms.)

Application of cladistics to terrane history—parsimony analysis of qualitative geological data

Hypotheses of terrane dispersal or accretion can be represented graphically as branching diagrams (cladograms), but an assessment of competing hypotheses of terrane history requires a method of analysis of supporting evidence which resolves the most parsimonious explanation of all available data. Cladistics is a rigorous analytical method first developed for phylogeny reconstruction (i.e. biological history), but applicable to any hierarchical data set. Given appropriate definitions, the various types of geological, geophysical and biological data used to support hypotheses of fragmentation or fusion history for geological regions (terranes) assumed to have had independent geological histories can be organized hierarchically. Terrane fragmentation is equivalent to phylogenetic splitting of biological taxa, and standard algorithms for parsimony analysis may be directly applied. Terrane accretion may be represented as a coalescing area cladogram, and the supporting evidence also forms a hierarchical data set, but with two main differences. The less general attributes historically precede the more general (the reverse applies in phylogeny reconstruction), and the branching points (nodes on the cladogram), unlike hypothetical common ancestors in phylogeny reconstruction, represent defined geographic areas, with a geological structure which can be investigated. In cladistic reconstruction of evolutionary history the common ancestors are hypothetical, and their attributes can only be inferred from the distribution of attributes amongst the terminals (known biological taxa); in contrast, the end product of terrane accretion is a composite structure (geological province) within which juxtaposition of terranes may eliminate some of the possible historical sequences which led to its formation.

Classification, reconstructed phylogeny and geographical history of genera of Pilipalpinae (Coleoptera : Tenebrionoidea : Pyrochroidae)

The 12 genera of Pilipalpinae are classified on the basis of characters of larvae and adults. Three new genera and six new species are here described: Malagaethes, gen. nov. (type species M. lawrencei, sp. nov.); Ranomafana, gen. nov. (type species R. steineri, sp. nov.); Binburrum, gen. nov. (type species Techmessa ruficollis Champion); Binburrum angusticollis, sp. nov.; Binburrum concavifrons, sp. nov.; Cycloderus immaculicollis, sp. nov. and Cycloderus hirsutus, sp. nov. The following new synonymies of specific names are proposed (with valid names given first): Paromarteon constans Lea, 1917 = Eucistela cyanea Carter, 1922; Paromarteon mutabile Blackburn, 1897 = Paromarteon mutabile var. nigripenne Lea, 1920; Temnopalpus bicolor Blackburn, 1888 = Temnopalpus tricolor Lea, 1920; Pilipalpus dasytoides Fairmaire, 1876 = Copobaenus maculicollis Pic, 1942 and Pilipalpus danvini Abdullah, 19646; Exocalopus pectinatus Broun, 1893 = Exocalopus antennalis Broun, 1903. The following subspecies have been elevated to species rank: Paromarteon apicale Lea, Paromarteon fasciatum Lea and Paromarteon parvum Lea. Phylogenetic analysis of 30 structural characters of larvae and adults yielded the following set of incompletely resolved relationships among genera of Pilipalpinae: (((Paromarteon + ((Temnopalpus + Malagaethes) + Pilipalpus + (Ranomafana + (Incollogenius + ((Exocalopus + (Binburrum + (Cycloderus + Morpholycus)) + Techmessodes) + Techmessa))))). The data set contained much homoplasy and several reversals. The historical geographical relationships inferred from the reconstructed phylogeny were compared with geological evidence for the break-up of Pangaea and Gondwanaland. The ancestral stock of Pilipalpinae was widespread on Gondwanaland, and differentiated through its fragmentation. Remnant relict genera persisted on Madagascar, New Zealand, southern South America (Magellanica), and Australia. Brooks Parsimony Analysis was conducted on the data resulting in the following area relationships: (Holarctic + (Madagascar + (New Zealand + (Australia + Chile)))). This agrees generally with accepted geological evidence and is considered support for they hypothesised phylogeny. A single clade (Temnopalpus + Malagaethes) was in disagreement (homoplasous) with the area cladogram, indicating possible incongruence in the data. The area relationships of other Southern Hemisphere groups were compared with Pilipalpinae.

On the Identification of Areas of Endemism

Methodological developments in cladis? tic biogeography have mostly focused on converting taxon-area cladograms into area cladograms and on obtaining general area cladograms (Nelson and Platnick, 1981; Wiley, 1988; Page, 1990; Nelson and Ladiges, 1991). The problem of identifying areas of endemism, although fundamental for any cladistic biogeographic study, has been somewhat neglected. An area of endemism is an area of non?

In Search of a Method for Cladistic Biogeography: An Empirical Comparison of Component Analysis, Brooks Parsimony Analysis, and Three-area Statements

Abstract—Three quantitative cladistic biogeographic methods, namely, component analysis, Brooks parsimony analysis (BPA), and three-area statements (TAS) have been proposed for obtaining general area cladograms from taxon-area cladograms. Available programs implementing these methods include COMPONENT versions 1.5 and 2.0 for component analysis, TAS for three-area statements, and Hennig86 for analysing matrices for both three-area statements and BPA. Ten different data sets were analysed with these programs and items of error were used to evaluate the general area cladograms obtained. None of the computer implementations of the methods compared proved to be more effective than the others."…all methodologies, even the most obvious ones, have their limits…"Feyerabend (1993:231)

BIOGEOGRAPHIC PATTERNS OF MONTANE MEXICAN AND CENTRAL AMERICAN CARABIDAE (COLEOPTERA)

AbstractCarabid taxa inhabiting the montane forests of México and Central America are analyzed using cladistic biogeographic methods. Taxa exhibiting the Nearctic, Meso-American Montane, and Paleo-American distributional patterns are included in the analysis. Widespread taxa are assumed to indicate that the areas of endemism they inhabit are closely related, and the general pattern of area relationships is determined using parsimony analysis. Of the nine areas of endemism recognized in the analysis, the Sierra Madre Oriental is considered the most distinct. The northern Sierra Madre Occidental, Arizona Mountains, and Sonoran Desert comprise one monophyletic set of areas, as do the more southerly areas of the Sierra Transvolcanica, Sierra Madre del Sur, Chiapan–Guatemalan highlands, and Talamancan Cordillera. The southern Sierra Madre Occidental area of endemism is equivocally related to both areas to the north and areas to the south. Biogeographic information derived from widespread species is as congruent with the general area cladogram as information derived from groups with precinctive species, indicating that the same biogeographic barriers often limit distributions of both widespread species and more inclusive clades comprising precinctive species. Taxa exhibiting the Meso-American Montane Pattern are more congruent with the general area cladogram than those representing the Nearctic Pattern, even though they exhibit less fidelity to highland habitats.

IN SEARCH OF A METHOD FOR CLADISTIC BIOGEOGRAPHY: AN EMPIRICAL COMPARISON OF COMPONENT ANALYSIS, BROOKS PARSIMONY ANALYSIS, AND THREE‐AREA STATEMENTS

Abstract— Three quantitative cladistic biogeographic methods, namely, component analysis, Brooks parsimony analysis (BPA), and three‐area statements (TAS) have been proposed for obtaining general area cladograms from taxon‐area cladograms. Available programs implementing these methods include COMPONENT versions 1.5 and 2.0 for component analysis, TAS for three‐area statements, and Hennig86 for analysing matrices for both three‐area statements and BPA. Ten different data sets were analysed with these programs and items of error were used to evaluate the general area cladograms obtained. None of the computer implementations of the methods compared proved to be more effective than the others."…all methodologies, even the most obvious ones, have their limits…"Feyerabend (1993:231)

Ancient and recent patterns of geographic speciation in the oyster mushroom Pleurotus revealed by phylogenetic analysis of ribosomal DNA sequences.

Evidence from molecular systematic studies suggests that many mushroom species may be quite ancient. Gene phylogenies were developed to examine the relationship between reproductive isolation, genetic divergence, and biogeography in oyster mushrooms (Pleurotus). Sequence data were obtained for two regions of DNA from populations belonging to eight intersterility groups (biological species). Phylogenetic analysis of sequences from the 5' portion of the nuclear encoded large subunit rDNA demonstrates an ancient origin for four intersterility groups of broad geographic distribution (world-wide), with a more recent radiation of several intersterility groups that are restricted to the Northern Hemisphere. An expanded analysis using sequence data from the more variable rDNA internal transcribed spacer region also reveals a phylogenetically based pattern of genetic divergence associated with allopatric speciation among populations from different continents in the Northern Hemisphere. The ability of rDNA sequences to resolve phylogenetic relationships among geographically isolated populations within intersterility groups illustrates the importance of biogeography for understanding speciation in Pleurotus. Patterns of geographic distribution among intersterility groups suggest that several species lineages evolved quite early, with recently evolved groups restricted to the Northern Hemisphere and older lineages occurring throughout the world. Based on phylogenetic evidence, analysis of historical biogeography using area cladograms shows that multiple dispersal and vicariance events are responsible for patterns of speciation observed.

The southern Palearctic genus Enneastigma Stein (Diptera: Anthomyiidae): Taxonomy of Mediterranean species, phylogeny and ancestral areas analysis

AbstractEnneastigma is a small genus of myopinine anthomyiids which comprises five species in the Mediterranean subregion (E. compressum Stein, E. rarum sp. n., E. rufitibia sp. n., E. triplex Loew, E. triseriatum Stein) and two species in SE Palearctic part of China (E. pilosiventrosum Fan & Chen, E. shanghaiensis Fan & Chen). All species appear to be univoltine and only active during the cooler parts of the season. The larvae may all be coprophagous. The Mediterranean species are revised with keys to males and females and illustrated descriptions of males, females and eggs. One lectotype is designated. A cladistic analysis was conducted which also included the Chinese species. This resulted in a single, most parsimonious phylogeny. The resulting area cladogram was used to assess the alternative methods for estimating ancestral areas as proposed recently by K. Bremer and F. Ronquist. It is concluded that the best results are obtained by using a refined Ronquist analysis.

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.

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.