Monophyletic Assemblage Research Articles

IsRhizoplaca(Lecanorales, lichenized Ascomycota) a monophyletic genus?

Rhizoplaca Zopf is a genus characterized by an umbilicate thallus with an upper and a lower cortex, as well as a cupulate hypothecium. It has been considered to be related to Lecanora Ach., the type genus of the Lecanoraceae and, in particular, to the lobate species of this genus. The phylogeny of Rhizoplaca, the monotypic Arctopeltis thuleana Poelt, and a number of representatives of different groups of Lecanora is studied, using sequences from the nuclear ribosomal internal transcribed spacer (ITS) regions. The results suggest an origin for Rhizoplaca species within the large genus Lecanora. A well-supported monophyletic assemblage includes the umbilicate type species Rhizoplaca melanophthalma (DC.) Leuck. & Poelt, the lobate Lecanora novomexicana H. Magn., and five vagrant Rhizoplaca species. Rhizoplaca chrysoleuca (Sm.) Zopf and Rhizoplaca subdicrepans (Nyl.) R. Sant. form a separate well-supported group and Rhizoplaca peltata (Ram.) Leuck. & Poelt is more closely related to Lecanora muralis (Schreb.) Rabenh. Together with data on secondary chemistry, the results show that the umbilicate thallus with a lower and an upper cortex, as well as apothecia with a cupulate hypothecium found in Rhizoplaca and A. thuleana, have developed several times in independant lineages in Lecanora. The thallus morphology in lecanoroid lichens is highly variable and does not necessarily reflect phylogenetic relationships.Key words: Rhizoplaca, Lecanora, Lecanorales, phylogeny, ITS.

Molecular re-assessment of some phylogenetic, taxonomic and biogeographic relationships between the leech genera Dina and Trocheta (Hirudinea: Erpobdellidae)

The three most abundant and species-rich genera of erpobdellid leeches – Erpobdella, Dina and Trocheta – have traditionally been distinguished by their annulation pattern. Erpobdella has five unsubdivided annuli per somite, in Dina the last annulus is widened and subdivided once, and in Trocheta, the last and often also the first annulus is further subdivided. The last two genera are often referred to as the subfamily Trochetinae. The presented molecular phylogenetic analysis, based on mitochondrial 12S rDNA sequences and nuclear ITS2 sequences, has revealed a sister relationship between Trocheta and Erpobdella rather than between Trocheta and Dina. Furthermore, the annulation pattern has proved to be inappropriate for the diagnosis of Dina and Trocheta. The taxon Trocheta bykowskii krasensehas been classified on the basis of its Trocheta-like annulation; however, according to the molecular phylogeny, it belongs to the Dina clade. A pair of diagnostic PCR primers were developed to identify unambiguously individuals with intermediate annulation. In summary, (1) the traditional genus diagnosis of Dina and Trocheta is no longer tenable, and the genera are not always morphologically distinguishable by traditional characters; (2) T. b. krasense Sket, 1968, should be re-classified as Dina krasensis; (3) the subfamily Trochetinae should be annulled because it does not represent a monophyletic assemblage.

Phylogenetic relationships and historical biogeography of melanotaeniid fishes in Australia and New Guinea

Phylogenetic analysis of melanotaeniid mtDNA cytochrome b and tRNA Pro-control region sequence is broadly consistent with the current taxonomy. However, the molecular phylogeny supports the elevation of M. s. australis to full species status and indicates either that it is a composite species or has introgressed with sympatric Melanotaenia species. Phenotypically cryptic mtDNA diversity in north-eastern Australia possibly represents an undescribed species. Six major monophyletic clades present in the phylogeny were strongly supported by morphological data. The clades represent three biogeographic regions. Fish from northern New Guinea form a monophyletic clade, within which Melanotaenia and Glossolepis are polyphyletic. The divergence of this clade from those in southern New Guinea is consistent with the final uplift of the Central Highlands 5 million years BP. North-western New Guinea and associated islands represent another highly divergent, monophyletic clade of a similar age to that in northern New Guinea. The remaining four clades form a monophyletic assemblage restricted to southern New Guinea and Australia: one in northern Australia, one with a disjunct distribution in north-western and eastern Australia, one widespread throughout Australia and southern New Guinea, and one in southern New Guinea with an outlying species in northern Australia. The phylogenetic relationships between Australia and southern New Guinea are consistent with episodic connection via the freshwater Lake Carpentaria during periods of low sea level.

Characterisation of cDNAs for ribulose bisphosphate carboxylase small subunit genes in different species of Avena (Poaceae)

ABSTRACT We report the cloning and sequencing of 14 rbcS cDNAs in six species of Avena (Poaceae) with different genome and ploidy levels. The nucleotide sequences 504 bp long were aligned with the published sequences of cultivated hexaploid oat, wheat, barley, rye, rice and corn and subjected to cladistic and phenetic analyses. The parsimonious analysis generated a tree with a topology very similar to the phenogram generated by the Neighbor-Joining analysis based on the Jukes and Cantor distances. Within the monophyletic assemblage of the tribe Aveneae, consistent clades composed of rbcS clones belonging to different species are recognized. It is suggested that they correspond to orthologous genes belonging to different subfamilies, and that the “within-species˚d homogenisation may have occurred at a slow rate with respect to species evolution. In the monophyletic group of Pooideae, the topologies place barley rbcS sequences closer to wheat and rye than to oat sequences. This grouping agrees with most taxonomic and phylogenetic views.

Molecular phylogeny of the apterygotan insects based on nuclear and mitochondrial genes

Summary Basal hexapodan orders (Apterygota) are traditionally divided in two well defined taxonomic groups, Entognatha and Ectognatha. Entognathy (the enclosed mouthparts condition) is considered the most distinctive character joining the Ellipura (Protura+ Collembola) with the order Diplura, whereas the presence of exposed mouthparts (ectognathy) is the feature shared by Microcoryphia and Zygentoma with the pterygote insects. In spite of the growing interest for the evolutionary history of the Apterygota, there is no complete agreement among the general phylogenetic hypotheses based on the study of morphological characters. In this study we analyzed the DNA sequence of segments of the nuclear Elongation Factor-1α(EF-1α) and of the mitochondrial 12S rRNA genes, and used different methods of phylogenetic reconstruction. The 12S gene seems to be more suited than the EF-1α to resolve some of the most outstanding systematic disputes, whereas the lack of resolution at the deeper nodes does not allow to assess the phylogenetic relationships within Microcoryphia and between ectognathan orders. We have obtained a fairly high support for the monophyly of the orders Diplura and Zygentoma. In the 12S analysis, the Ellipura and the Entognatha form monophyletic assemblages. In addition, the study of the distribution of introns in the EF-1α suggests a relationship between Collembola and Diplura.

Cloning of a Proopiomelanocortin cDNA from the Pituitary of the Australian Lungfish, Neoceratodus forsteri: Analyzing Trends in the Organization of this Prohormone Precursor

The polypeptide hormone precursor, proopiomelanocortin (POMC), was cloned and sequenced from the pituitary of the Australian lungfish, Neoceratodus forsteri, the only surviving species of the oldest extant lineage of lungfish. The Australian lungfish POMC cDNA had an open reading frame that coded for a 255-amino acid precursor. A comparison of POMC sequences from the Australian lungfish and the African lungfish indicated that the deduced amino acid sequences for ACTH, β-MSH, and β-endorphin were over 90% identical. Furthermore, within the open reading frames of the two lungfish POMCs, there was 84% identity at the nucleotide level. Although a γ-MSH-like region was detected in the Australian lungfish POMC cDNA, this sequence contained mutations that have been detected in the γ-MSH sequences of some ray-finned fish and are not found in the γ-MSH sequence of the African lungfish or those of tetrapods. In addition, the sequence of β-endorphin in the two species of lungfish has amino acid motifs that are found in the β-endorphin sequences of cartilaginous fish and ray-finned fish but not in tetrapods. However, maximum parsimony analysis of the entire POMC open reading indicated that the lungfish POMC sequences form a clade with two amphibian POMC sequences rather than with POMC sequences from ray-finned fish. This result is consistent with the accepted view that the sarcopterygians (lungfishes and tetrapods) are a monophyletic assemblage. Analysis of rates of divergence for various POMC sequences indicate that point mutations are accumulating in the lungfish POMC sequences at a slower rate than in either amphibian or mammalian POMC sequences. The phylogenetic implications of these observations are discussed.

Using Character Concordance to Define Taxonomic and Conservation Units

Conservation biologists often apply evolutionarily defensible definitions of conservation units, such as evolutionarily significant units (ESU; Moritz 1994, 1995; Vogler & DeSalle 1994), when assessing the conservation status of populations (Legge et al. 1996; Britten et al. 1997; Pichler et al. 1998; Waits et al. 1998; Zhu et al. 1998; Roman et al. 1999). Conversely, taxonomists often do not explicitly invoke any definition when delineating their fundamental units, species. For example, only 4 of 74 species descriptions (fishes, amphibians, and reptiles) published in Copeia since 1995 identify an underlying species definition. We can only assume, based on the morphological emphasis, that a morphological species concept is applied in most of these instances. Three problems can result from taxonomic practices that delineate species without an operational evolutionary framework. First, the characteristics of species (e.g., reproductively isolated arrays or monophyletic assemblages) are not adequately defined to prevent these evolutionary units from becoming simple arrays of individuals that are distinct along one or a few axes (e.g., families). Second, a fundamental partition (diagnostic character) frequently frames analyses of character variation in subsequent studies, an operational flaw in taxonomic practice (Wilson & Brown 1953) from which population geneticists are not immune. Third, conflicting patterns of distinctiveness among individual characters lead to taxonomic confusion and inevitable controversy over what constitutes a good characteristic for taxon-level descriptions. These problems contribute to taxonomic and conservation controversies such as that outlined by Karl and Bowen (this issue) that consume limited conservation resources. Examining variation among genetically independent characters for common patterns (character concordance) might reduce taxonomic controversy by providing an evolutionary framework within which character variation can be objectively evaluated. The black-green turtle debate illustrates the problem of taxonomy without an evolutionary framework, the inevitable controversy that ensues, and the advantage of a character concordance approach. Black turtles were first recognized as taxonomically distinct because they are not green but nonetheless resemble green turtles in other attributes (Bocourt 1868). Under many species concepts, species are distinct units, but distinctness is a relative assessment that can apply to a variety of hierarchical levels, including individuals (Vogler & DeSalle 1994). Also, arrays of organisms can be distinct for reasons other than a shared history of divergence, for example, when the phenotype is largely determined environmentally. When our expectation is that species constitute distinctive arrays of individuals and our focus is a single trait-color, for example-we risk delineating nonevolutionary units as species.

The South American catfish genus Auchenipterus Valenciennes, 1840 (Ostariophysi: Siluriformes: Auchenipteridae): monophyly and relationships, with a revisionary study

Abstract The Neotropical catfishes of the genus Auchenipterus Valenciennes (1840) are reviewed. The genus is hypothesized to be a monophyletic assemblage on the basis of the shared presence of grooves in the ventral surface of the head that accommodate adducted mental barbels. A possible second synapomorphy, the presence of papillae on the dorsal and medial surface of the ossified maxillary barbel of mature males, is tentatively advanced pending discovery of adult males of three species. Contrary to previous hypotheses which considered Auchenipterus to consist of a maximum of five species, we recognize 11 species, including two previously undescribed forms, A. britskii and A. menezesiAuchenipterus is broadly distributed through the Rio Orinoco, Rio Amazonas, and Rio de La Plata basins, and the coastal drainages of the Guianas, with one species in the Rio Pindare-Mirim and Rio Parnaiba basins of northeastern Brazil. Auchenipterus nuchalis, previously thought to be broadly distributed across the range of the genus, is found to rather have a restricted distribution in the eastern portions of the Amazon basin, the lower portions of the Rio Tocantins, and lower courses of some rivers in Suriname and French Guiana. Citations of A. nuchalis from elsewhere in the range of the genus are of other species. Euanemus Muller & Troschel (1842) and Ceratocheilus Miranda Ribeiro (1918) are considered synonyms of Auchenipterus. Euanemus colymbetes Muller & Troschel (1842) is considered a synonym of Auchenipterus dentatus Valenciennes (1840), and A. paysanduanus Devincenzi (1933) is placed into the synonymy of A. nigripinnis Boulenger (1895). A neotype is designated for Hypopthalmus nuchalis Spix & Agassiz (1829). Lectotypes are designated for Euanemus colymbetes and Auchenipterus nigripinnis.

The South American catfish genus Auchenipterus Valenciennes, 1840 (Ostariophysi: Siluriformes: Auchenipteridae): monophyly and relationships, with a revisionary study

The Neotropical catfishes of the genus Auchenipterus Valenciennes (1840) are reviewed. The genus is hypothesized to be a monophyletic assemblage on the basis of the shared presence of grooves in the ventral surface of the head that accommodate adducted mental barbels. A possible second synapomorphy, the presence of papillae on the dorsal and medial surface of the ossified maxillary barbel of mature males, is tentatively advanced pending discovery of adult males of three species. Contrary to previous hypotheses which considered Auchenipterus to consist of a maximum of five species, we recognize 11 species, including two previously undescribed forms, A. britskii and A. menezesiAuchenipterus is broadly distributed through the Rı́o Orinoco, Rio Amazonas, and Rı́o de La Plata basins, and the coastal drainages of the Guianas, with one species in the Rio Pindaré-Mirim and Rio Parnaı́ba basins of northeastern Brazil. Auchenipterus nuchalis, previously thought to be broadly distributed across the range of the genus, is found to rather have a restricted distribution in the eastern portions of the Amazon basin, the lower portions of the Rio Tocantins, and lower courses of some rivers in Suriname and French Guiana. Citations of A. nuchalis from elsewhere in the range of the genus are of other species. Euanemus Müller & Troschel (1842) and Ceratocheilus Miranda Ribeiro (1918) are considered synonyms of Auchenipterus. Euanemus colymbetes Müller & Troschel (1842) is considered a synonym of Auchenipterus dentatus Valenciennes (1840), and A. paysanduanus Devincenzi (1933) is placed into the synonymy of A. nigripinnis Boulenger (1895). A neotype is designated for Hypopthalmus nuchalis Spix & Agassiz (1829). Lectotypes are designated for Euanemus colymbetes and Auchenipterus nigripinnis.

Mitochondrial DNA phylogeny of the family Cichlidae: monophyly and fast molecular evolution of the neotropical assemblage.

A mitochondrial DNA (mtDNA) phylogeny of cichlid fish is presented for the most taxonomically inclusive data set compiled to date (64 taxa). 16S rDNA data establish with confidence relationships among major lineages of cichlids, with a general pattern congruent with previous morphological studies and less inclusive molecular phylogenies based on nuclear genes. Cichlids from Madagascar and India are the most basal groups of the family Cichlidae and sister to African-Neotropical cichlids. The cichlid phylogeny suggests drift-vicariance events, consistent with the fragmentation of Gondwana, to explain current biogeographic distributions. Important phylogenetic findings include the placement of the controversial genus Heterochromis basal among African cichlids, the South American genus Retroculus as the most basal taxon of the Neotropical cichlid assemblage, and the close relationship of the Neotropical genera Cichla with Astronotus rather than with the crenicichlines. Based on a large number of South American genera, the Neotropical cichlids are defined as a monophyletic assemblage and shown to harbor significantly higher levels of genetic variation than their African counterparts. Relative rate tests suggest that Neotropical cichlids have experienced accelerated rates of molecular evolution. But these high evolutionary rates were significantly higher among geophagine cichlids.

Systématique moléculaire des Mélitées

The holarctic subfamily Melitaeinae of Nymphalid butterflies (old genus Melitaea Hbn.) has been split into various genera by morphological systematics, but this division has been debated. This work presents the results of a preliminary survey of European taxa using two types of molecular criteria: analysis of enzyme variation by electrophoresis (19 loci) and sequencing of a mtDNA fragment ( ND1 gene). The information provided by both kinds of markers are largely congruent. The division of the subfamily into two monophyletic lines corresponding to the genera Euphyryas s. 1. and Melitaea s. l. is validated. The subsequent division of the former genus into two units, Eurodryas and Hypodryas is also confirmed by molecular criteria. Inside rhe old genus Melitaea, a monophyletic assemblage, corresponding to the ‘ Mellicta group’ is disclosed, but the phylogeny of the other species of the group is confused. Uniting them in a ‘Didymaeformia group’ or in a genus Melitaea s. str. risks rhe generation of a paraphyletic assemblage. It remains to be determined whether or not these results are due to insufficient resolution, or whether they reflecr a real evolutionary pattern

A cladistic analysis of Platythelphusa A. Milne-Edwards, 1887, from Lake Tanganyika, East Africa (Decapoda: Potamoidea: Platythelphusidae) with comments on the phylogenetic position of the group

The phylogenetic relationships of the freshwater crab genus Platythelphusa were investigated by means of a cladistic analysis using morphological characters. The platythelphusa were found to be a monophyletic assemblage, this finding supports the recognition of the family Platythelphusidae. Platythelphusa is endemic to Lake Tanganyika, East Africa, and some of its species bear a striking resemblance to certain marine crabs (grapsids). The study does not support the idea that the Platythelphusidae emerged from a relict population of a grapsid or grapsid-like taxon: the evidence supports an hypothesis for the evolution of this family from freshwater crab ancestors. Parsimony considerations position the freshwater crab family Deckeniidae as the sister group of the platythelphusids. In addition, the Palaeotropical freshwater crabs appear to constitute a monophyletic group.

Study of the Larvae of Antiporus strigosulus (Broun) (Coleoptera: Dytiscidae) with Implications for the Phylogeny of the Hydroporini

Description of structures of the third instar of Antiporus strigosulus (Broun) is presented. Based on actual knowledge of larval morphology of the subfamily Hydroporinae, it is confirmed that Antiporus Sharp belongs to tribe Hydroporini. Within the Hydroporini, Antiporus is hypothesized to constitute a monophyletic assemblage along with Nebrioporus Régimbart and Stictotarsus Zimmermann based on two synapomorphies: presence of natatory setae on dorsal margin of femora, tibiae, and tarsi, and urogomphomere 1 which is more than 3.10 times longer than urogomphomere 2. This clade may include also other genera of which information on these character states is not available.

Evolution of the Ribosomal RNA Internal Transcribed Spacer One (ITS-1) in Cichlid Fishes of the Lake Victoria Region

The nucleotide sequences of the first internal transcribed spacer (ITS-1) of the ribosomal RNA gene cluster have been determined for 11 species of closely related endemic cichlid fishes of the Lake Victoria region (LVR) and 6 related East African cichlids. The ITS-1 sequences confirmed independently derived basal phylogenies, but provide limited insight within this species flock. The line leading toPseudocrenilabrus multicolorarose early, close to the divergence event that separated the tilapiine and haplochromine tribes of the “African Group” of the family Cichlidae. In this phylogeny,Astatoreochromis alluaudiand the riverineAstatotilapia burtoniare sister taxa, which together are a sister group to a monophyletic assemblage including both Lake Victoria and Lake Edward taxa. The ITS-1 data support the monophyly of haplochromine genera across lakes. Since Lake Victoria is believed to have been dry between 14,500 and 12,400 BPE, the modern assemblage must have been derived from reinvasion by the products of earlier cladogenesis events. Thus, although the regional superflock is monophyletic, the haplochromines of Lake Victoria itself did not evolvein situfrom a single ancestor.

Developmental Ultrastructure of the Male Gamete ofSelaginella

To evaluate structural homology among streamlined, biflagellated male gametes of Selaginella and bryophytes, we undertook a thorough ultrastructural study of spermatogenesis in Selaginella kraussiana (Kunze) A. Br. As in Lycopodium and bryophytes, centrioles originate as bicentrioles in the spermatid mother cell. In the nascent spermatid, a fibrous connective band is involved in the separation and rotation of the two centriolar constituents of the bicentriole. The locomotory apparatus consists of two staggered monomorphic basal bodies that originate from the centrioles and an underlying multilayered structure (MLS), including a spline of parallel microtubules and subtending elongated lamellar strip. Beneath the MLS lies the anterior mitochondrion, which elongates during development to a length of over 1.5 cellular revolutions by progressive fusion of mitochondria. Spline microtubules (MTs) increase posteriorly from one at the anterior tip to a maximum of 19 at the cell posterior. The angle between lamellar strip plates and spline MTs changes front to back from 85° to 40°. In the late midstage spermatid, the rounded nucleus compacts and elongates in register with spline growth. Individualization of cellular coils begins at the cell anterior and progresses posteriorly leaving a cytoplasmic mass in the region between coils. In the final stages of development, the lamellar strip regresses and the cylindrical nucleus shortens. Concomitantly, the posterior mitochondrion and plastid, containing two starch grains, are positioned alongside the spline at the posterior of the cell. In development, the spermatozoid of Selaginella more closely resembles those of pteridophytes than bryophytes. Common developmental and structural features of the locomotory apparatus among Selaginella and members of the Lycopodiaceae are consistent with a monophyletic lycopsid assemblage.

A phylogenetic analysis of the genus Carica L. (Caricaceae) based on restriction fragment length variation in a cpDNA intergenic spacer region

The phylogenetic relationships among twelve wild and cultivated species of Carica (Caricaceae) were analyzed using restriction fragment length variation in a 3.2-kb PCR amplified intergenic spacer region of the chloroplast DNA. A total of 138 fragments representing 137 restriction sites accounting for 5.8% of the amplified region were examined. Both parsimony and neighbor joining cluster analyses confirmed the close association among South American wild Carica species. However, cpDNA data did not support the traditional monophyly hypothesis for the evolution of Carica. Further, cpDNA analyses showed two basic evolutionary lineages within the genus Carica, one defined by cultivated C. papaya and another consisting of the remaining wild species from South America in a well resolved but poorly supported monophyletic assemblage. This evolutionary split in Carica strongly suggests that C. papaya diverged from the rest of the species early in the evolution of the genus and evolved in isolation, probably in Central America.

The tortoise and the hare: choosing between noncoding plastome and nuclear Adh sequences for phylogeny reconstruction in a recently diverged plant group

Phylogenetic resolution is often low within groups of recently diverged taxa due to a paucity of phylogenetically informative characters. We tested the relative utility of seven noncoding cpDNA regions and a pair of homoeologous nuclear genes for resolving recent divergences, using tetraploid cottons (Gossypium) as a model system. The five tetraploid species of Gossypium are a monophyletic assemblage derived from an allopolyploidization event that probably occurred within the last 0.5-2 million years. Previous analysis of cpDNA restriction site data provided only partial resolution within this clade despite a large number of enzymes employed. We sequenced three cpDNA introns (rpl16, rpoC1, ndhA) and four cpDNA spacers (accD-psaI, trnL-trnF, trnT-trnL, atpB-rbcL) for a total of over 7 kb of sequence per taxon, yet obtained only four informative nucleotide substitutions (0.05%) resulting in incomplete phylogenetic resolution. In addition, we sequenced a 1.65-kb region of a homoeologous pair of nuclear-encoded alcohol dehydrogenase (Adh) genes. In contrast with the cpDNA sequence data, the Adh homoeologues yielded 25 informative characters (0.76%) and provided a robust and completely resolved topology that is concordant with previous cladistic and phenetic analyses. The enhanced resolution obtained using the nuclear genes reflects an approximately three- to sixfold increase in nucleotide substitution rate relative to the plastome spacers and introns.

Molecular Data Confirms the Species Status of the Christmas Island Hawk-Owl Ninox natalis

SummaryDNA sequences from the mitochondrial ND2 gene were used to assess the taxonomic status of the Christmas Island Hawk-Owl Ninox natalis. Sequence variation was examined across 17 specimens representing 9 species and subspecies within the genus Ninox. The comparisons revealed a correlation between taxonomic level and genetic divergence with a clear separation between the levels of sequence divergence observed in comparisons involving individuals, subspecies and species. These observations provided strong endorsement for the use of DNA sequence data to assess the taxonomic status and phylogenetic relationships of N. natalis. Levels of sequence divergence recorded between the three forms of the N. squamipila complex examined (squamipila, hypogramma and natalis) ranged from 4.0% to 4.8%. These were comparable to the levels of divergence observed between obviously distinct species such as N. rufa and N. strenua (5.4%) and were consistently greater than observed among subspecies within the monophyletic N. novaeseelandiae complex (1.5% to 2.3% between novaesee-landiae, leucopsis and undulata). The genetic distinctiveness of all three forms of squamipila was also apparent from phylogenetic analysis of the data. Distance and parsimony methods both failed to identify any of the three forms of squamipila as forming a monophyletic assemblage. Based on the combined evidence (sequence divergence and phylogenetic position) it is concluded that squamipila, hypogramma and natalis each represent separate species in the genus Ninox. These findings have implications for conservation efforts in the region.

Phylogeny of West AfricanCaryedon(Coleoptera: Bruchidae): Congruence between Molecular and Morphological Data

Seed beetles belonging to the Old World genusCaryedonfeed in the seeds of various Caesalpinioideae, Mimosoideae, and Combretaceae. In an attempt to resolve broad phylogenetic relationships within the genus, we obtained 332 base pair sequences of mitochondrial 12S ribosomal DNA and morphological data for the 16 West AfricanCaryedonspecies. Morphological characters were analyzed under maximum parsimony and sequences were compared under maximum parsimony, maximum likelihood, and neighbor joining. Using a partition homogeneity test, we determined that morphological and molecular data sets were combinable. Combined data were analyzed under maximum parsimony. Morphological and molecular trees were congruent at the species group level and total evidence analyses yielded the same topologies as molecular data with each of the three outgroups used. Four main terminal clades are recognized, each corresponding with a group of species generally feeding on the same host plant family, subfamily, genus, or species. The monophyly of legume feedingCaryedonis supported by both data sets, and Combretaceae feeders split in two monophyletic assemblages.

Cladistic analysis, phylogeny and biogeography of the Hawaiian Platynini (Coleoptera: Carabidae)

The 128 known native Hawaiian species of the tribe Platynini are analysed cladistically. Cladistic analysis is based on 206 unit‐coded morphological characters, and also includes forty‐one outgroup taxa from around the Pacific Rim. Strict consensus of the multiple equally parsimonious cladograms supports the monophyly of the entire species swarm. The closest outgroup appears to be the south‐east Asian‐Pacific genus Lorostema Motschulsky, whose species are distributed from India and Sri Lanka to Tahiti, supporting derivation of the Hawaiian platynines from a source in the western or south‐western Pacific. The biogeographic relationships of the Hawaiian taxa are analysed using tree mapping, wherein items of error are minimized. The area cladogram found to be most congruent with the phylogenetic relationships, and most defensible based on underlying character data is {Kauai[Oahu(Hawaii{Lanai[East Maui(West Maui + Molokai)]})]}. This progressive vicariant pattern incorporates progressive colonization from Kauai, and vicariance of the former Maui Nui into the present islands of Molokai, Lanai, West Maui and East Maui. The evolution of flightlessness, tarsal structure, pronotal setation and bursal asymmetry are evaluated in the context of the cladogram. Brachyptery is a derived condition for which reversal is not mandated by the cladogram, although repeated evolution of reduced flight wings is required. Tarsal structure supports Sharp's (1903) recognition of Division 1 as a monophyletic assemblage, but exposes his Division 2 as a paraphyletic group requiring removal of the genus Colpocaccus Sharp. Pronotal setation is exceedingly homoplastic, and is not useful for delimiting natural groups. Left‐right asymmetry of the bursa copulatrix reversed twice independently, resulting in mirror‐image bursal configurations in B. rupicola and Prodisenochus terebratus of East Maui. The amount of character divergence is greater among species comprising Division 1 than among species of its sister group, the redefined Division 2. Based on superior fit of Division 1 relationships to the general biogeographic pattern, a greater speciation rate coupled with more extensive extinction is rejected as the cause for this greater divergence. Intrinsic differentiation in the processes underlying cuticular evolution appears to be more consistent with the observed biogeographic and morphological patterns.