Ingroup Clades Research Articles

Phylogeny of North Americas largest cicada radiation redefines Tibicinoides and Okanagana (Hemiptera: Auchenorrhyncha: Cicadidae: Tibicininae).

Tibicinoides, with three small endemic California cicada species, has a confusing, intertwined systematic history with Okanagana that we unravel here. An ingroup including all species of Tibicinoides and the majority (84.7%) of Okanagana species were sampled for six gene regions, polarized with Clidophleps, Okanagodes, Subpsaltria, and Tibicina outgroups, and subjected to Bayesian phylogenetic analysis. Although the ingroup was monophyletic from all outgroups including Tibicina, Tibicinoides rendered Okanagana paraphyletic among two major ingroup clades. To bring classification into agreement with phylogeny, we redescribe and redefine Tibicinoides to include all Okanagana species with a hooked uncus in the male genitalia, all of which grouped with the type T. cupreosparsa (Uhler, 1889) in the first of these clades: T. boweni (Chatfield-Taylor & Cole, 2020) comb. n., T. catalina (Davis, 1936) comb. n., T. hesperia (Uhler, 1876) comb. n., T. mercedita (Davis, 1915), T. minuta (Davis, 1915), T. pallidula (Davis, 1917a) comb. n., T. pernix (Bliven, 1964) comb. n., T. rubrovenosa (Davis, 1915) comb. n., T. simulata (Davis, 1921) comb. n., T. striatipes (Haldeman, 1852) comb. n., T. uncinata (Van Duzee, 1915) comb. n., T. utahensis (Davis, 1919) comb. n., and T. vanduzeei (Distant, 1914) comb. n. Okanagana is redescribed and restricted to the species of the second major clade which contained the type O. rimosa (Say, 1830). We describe two new genera for morphologically distinct orphan lineages: Chlorocanta gen. nov. for C. viridis (Davis, 1918) comb. n. and Hewlettia gen. nov. for H. nigriviridis (Davis, 1921) comb. n. We recognize O. rubrobasalis Davis, 1926 stat. rev. as a species and relegate two former species to junior subjective synonyms: O. noveboracensis (Emmons, 1854) = O. canadensis (Provancher, 1889) and O. occidentalis (Walker in Lord, 1866) = O. lurida Davis, 1919. Tibicinoides and Okanagana together represent a rapid radiation that presents challenges to phylogenetic analysis including suboptimal outgroups and short internodes.

Evolution of diverse host infection mechanisms delineates an adaptive radiation of lampsiline freshwater mussels centered on their larval ecology.

North American watersheds contain a high diversity of freshwater mussels (Unionoida). During the long-lived, benthic phase of their life cycle, up to 40 species can co-occur in a single riffle and there is typically little evidence for major differences in their feeding ecology or microhabitat partitioning. In contrast, their brief parasitic larval phase involves the infection of a wide diversity of fish hosts and female mussels have evolved a spectrum of adaptations for infecting host fish with their offspring. Many species use a passive broadcast strategy: placing high numbers of larvae in the water column and relying on chance encounters with potential hosts. Many other species, including most members of the Lampsilini, have a proactive strategy that entails the use of prey-mimetic lures to change the behavior of the hosts, i.e., eliciting a feeding response through which they become infected. Two main lure types are collectively produced: mantle tissue lures (on the female’s body) and brood lures, containing infective larvae, that are released into the external environment. In this study, we used a phylogenomic approach (ddRAD-seq) to place the diversity of infection strategies used by 54 North American lampsiline mussels into an evolutionary context. Ancestral state reconstruction recovered evidence for the early evolution of mantle lures in this clade, with brood lures and broadcast infection strategies both being independently derived twice. The most common infection strategy, occurring in our largest ingroup clade, is a mixed one in which mimetic mantle lures are apparently the predominant infection mechanism, but gravid females also release simple, non-mimetic brood lures at the end of the season. This mixed infection strategy clade shows some evidence of an increase in diversification rate and most members use centrarchids (Micropterus & Lepomis spp.) as their predominant fish hosts. Broad linkage between infection strategies and predominant fish host genera is also seen in other lampsiline clades: worm-like mantle lures of Toxolasma spp. with sunfish (Lepomis spp.); insect larvae-like brood lures (Ptychobranchus spp.), or mantle lures (Medionidus spp., Obovaria spp.), or mantle lures combined with host capture (Epioblasma spp.) with a spectrum of darter (Etheostoma & Percina spp.) and sculpin (Cottus spp.) hosts, and tethered brood lures (Hamiota spp.) with bass (Micropterus spp.). Our phylogenetic results confirm that discrete lampsiline mussel clades exhibit considerable specialization in the primary fish host clades their larvae parasitize, and in the host infection strategies they employ to do so. They are also consistent with the hypothesis that larval resource partitioning of fish hosts is an important factor in maintaining species diversity in mussel assemblages. We conclude that, taking their larval ecology and host-infection mechanisms into account, lampsiline mussels may be legitimately viewed as an adaptive radiation.

Molecular phylogeny and evolution of Calaphidinae (Hemiptera: Aphididae).

Calaphidinae is the second-largest subfamily in the family Aphididae. Despite their species diversity and some taxonomic controversy, no phylogenetic studies have been conducted on them thus far. Herein, we report the first molecular phylogeny of Calaphidinae and two related lineages, Phyllaphidinae and Saltusaphidinae, based on five genes (3418 bp) for 126 taxa. Maximum parsimony, maximum-likelihood and Bayesian inference phylogenetic analyses were performed on the multilocus dataset. Divergence time estimation, biogeographical reconstruction, ancestral host plant reconstruction and PhyloType analyses were performed to identify evolutionary trends in Calaphidinae. Our phylogenetic results lead to several conclusions: Phyllaphidinae is a sister group to Calaphidinae s.l.; Calaphidinae is paraphyletic with respect to the former "Saltusaphidinae"; the ingroup clade was subdivided into nine newly recognized lineages; and three subtribes of Calaphidinae (Monaphidina, Calaphdina and Panaphidina) and many genera were not recovered as monophyletic. A new classification is proposed with eight tribal divisions that reflect our phylogenetic results, including three new tribes (Pterocallidini trib.n., Pseudochromaphidini trib.n. and Shivaphidini trib.n.) and three new statuses (Saltusaphidini stat.n., Therioaphidini stat.n. and Myzocallidini stat.n.). The ancestral reconstruction results imply that the ingroup taxa's common ancestor originated in the Eastern Palaearctic and might have fed on Fagaceae in the Late Cretaceous. Later, multiple host shifts and an expanding geographical distribution led to the current species diversity of Calaphidinae. Our reconstructions suggest that species diversification cannot solely be explained by speciation via host shifts and that geographical isolation probably also played a key role. Our results provide new insight into the natural classification and history of the host plant associations and biogeography of Calaphidinae s.l.

Corrections to the spellings of two tribe names in the leafhopper subfamily Ledrinae (Hemiptera: Cicadellidae).

In 2009, Jones and Deitz published a tribe-level taxonomic revision and reclassification of the cryptic, arboreal leafhopper subfamily Ledrinae Kirschbaum, 1868 (Hemiptera: Cicadellidae), based on cladistic analyses of 235 morphological features for 75 cicadellid species. Their evolutionary reconstructions found strong node support for a monophyletic ingroup comprising five lineages-each morphologically and geographically cohesive-and also identified numerous traditionally placed taxa (sensu Oman et al 1990) that did not belong. In light of the robustness of their results, the authors recognized the five independent ingroup clades as tribes of Ledrinae, and described three of these as new.

Analisis Filogeni Thiaridae (Mollusca: Gastropoda) Papua Menggunakan Karakter Morfologi

The characteristics  morphology and variations of shell can be used to reconstruct the sequence and phylogenetic of Thiaridae. 34 characters and 2 outgroups of Bellamya sp. from Papua and Sulawesi were used to observe 27 species Thiaridae. Phylogenetic Analysis Using Parsimony (PAUP Software Version 4.0., Swofford) and the neighbour joining cladogram were used to reconstruct the phylogenetic relationship and to group the taxa of Thiaridae based on distribution areas of genera separately. The study indicated that  the taxon of Thiaridae can be groupped into 3 clades. Each clade depicts the taxon population based on their distribution areas in Batanta Island, Sorong, Manokwari, and Bintuni Gulf. All molded into one clade. Biak, Supiori, Serui, Wamena, Merauke, and Jayapura are formed one clade. Thiara, Terebia, and Melanoides are genera which formed a clade each genus. While Stenomelania molded into 3 clades. Morphology character and closed regionsof distribution tended to form ingroup clade and sister species.Key words: characters, phylogeni, Thiaridae, dan Papua.

A New Snake Skull from the Paleocene of Bolivia Sheds Light on the Evolution of Macrostomatans

Macrostomatan snakes, one of the most diverse extant clades of squamates, display an impressive arsenal of cranial features to consume a vast array of preys. In the absence of indisputable fossil representatives of this clade with well-preserved skulls, the mode and timing of these extraordinary morphological novelties remain obscure. Here, we report the discovery of Kataria anisodonta n. gen. n. sp., a macrostomatan snake recovered in the Early Palaeocene locality of Tiupampa, Bolivia. The holotype consists of a partial, minute skull that exhibits a combination of booid and caenophidian characters, being the presence of an anisodont dentition and diastema in the maxilla the most distinctive trait. Phylogenetic analysis places Kataria basal to the Caenophidia+Tropidophiidae, and represents along with bolyeriids a distinctive clade of derived macrostomatans. The discovery of Kataria highlights the morphological diversity in the maxilla among derived macrostomatans, demonstrating the relevance of maxillary transformations in the evolution of this clade. Kataria represents the oldest macrostomatan skull recovered, revealing that the diversification of macrostomatans was well under way in early Tertiary times. This record also reinforces the importance of Gondwanan territories in the history of snakes, not only in the origin of the entire group but also in the evolution of ingroup clades.

Phylogenetic reassessment of the Exophthalmus genus complex (Curculionidae: Entiminae: Eustylini, Geonemini)

The monophyly of the Neotropical entimine weevil genus Exophthalmus Schoenherr, 1823 (Curculionidae: Entiminae: Eustylini Lacordaire) is reassessed. Exophthalmus presently includes more than 80 species, approximately half of which are restricted to either the Caribbean archipelago or the continental Neotropics. The taxonomic composition and position of Exophthalmus have been subject to longstanding disagreements; in particular, authors have questioned the relationship of Exophthalmus to other Caribbean genera such as Diaprepes Schoenherr, 1834 (Eustylini) and Lachnopus Schoenherr, 1840 (Geonemini Gistel), as well as to the speciose Central and South American genera Compsus Schoenherr, 1823, Eustylus Schoenherr, 1842, and Exorides Pascoe, 1881 (all Eustylini), among others. The present study scrutinizes these traditional perspectives, based on a cladistic analysis of 143 adult morphological characters and 90 species, representing 30 genera and seven tribes of Neotropical entimine weevils. The character matrix yielded eight most-parsimonious cladograms (length = 239 steps; consistency index = 66; retention index = 91), with mixed clade support that remains particularly wanting for some of the deeper in-group divergences. The strict consensus supports the existence of a paraphyletic Geonemini ‘grade’ that includes Lachnopus and related Caribbean genera such as Apotomoderes Dejean, 1834, followed by a monophyletic Eustylini in-group clade. Within the latter, a monophyletic South American Eustylini clade - including Compsus, Eustylus, Exorides, and related genera - is sister to a major clade that contains a ‘grade’ of heterogeneous and often misclassified Caribbean members of the Eustylini, Geonemini (Tetrabothynus Labram & Imhoff, 1852 and Tropirhinus Schoenherr, 1823), and Tanymecini Lacordaire (Pachnaeus Schoenherr, 1826), as well as two major clades: one with the majority of Central American Exophthalmus species, and the other with most Caribbean members of Exophthalmus. The Central American Exophthalmus clade is paraphyletic with respect to Chauliopleurus Champion, 1911 (Geonemini) and Rhinospathe Chevrolat, 1878 (Phyllobiini Schoenherr). The Caribbean clade, in turn, contains two subclades: i.e. (1) the Greater Antillean Exophthalmus s.s. clade, including the type species Exophthalmus quadrivittatus (Olivier, 1807); and (2) the primarily Lesser Antillean Diaprepes. The latter genus is therefore nested within Central American and Caribbean species of a highly paraphyletic Exophthalmus, yet may be rendered monophyletic if several Lesser Antillean Exophthalmus species are (re-)assigned to Diaprepes. The results thus provide a suitable basis for a revision of all Exophthalmus species, and furthermore suggest that historical biographic factors, including colonization via temporary continental Neotropics-to-Caribbean land connections, were important in the evolution of major eustyline lineages. Based on these preliminary insights, the following taxonomic and nomenclatural adjustments are made. Compsoricus gen. nov. is erected to accommodate two Puerto Rican species erroneously assigned to Compsus: i.e. the herein designated type species Compsoricus maricao comb. nov. and Compsoricus luquillo comb. nov. Eustylus dentipes comb. nov. is transferred from Compsus. Diaprepes marginicollis Chevrolat, 1880 is reinstated from synonymy under Exophthalmus. Lastly, the following five transfers are proposed: (1) Chauliopleurus Champion, 1911, from Geonemini to Eustylini; (2) Tetrabothynus Labram & Imhoff, 1852, from Geonemini to Eustylini; (3) Tropirhinus Schoenherr, 1823, from Geonemini to Eustylini; (4) Rhinospathe Chevrolat, 1878, from Phyllobiini to Eustylini; and (5) Pachnaeus Schoenherr, 1826, from Tanymecini to Eustylini.

Phylogenetic assessment of the Caribbean weevil genus Lachnopus Schoenherr (Coleoptera:Curculionidae:Entiminae)

The genus Lachnopus Schoenherr (Coleoptera : Curculionidae : Entiminae : Geonemini Gistel), with 66 species described to date, is the largest and most widespread entimine weevil genus in the Caribbean region. The monophyly and internal phylogenetic composition of Lachnopus are tested, using a cladistic analysis of 61 adult morphological characters of 31 ingroup species and five outgroup taxa representing the genera Diaprepes Schoenherr, Exophthalmus Schoenherr, Pachnaeus Schoenherr (all Eustylini), Ischionoplus Chevrolat (Geonemini) and Apodrosus Marshall (Polydrusini). The analysis produced two most parsimonious cladograms with a length of 167 steps, a consistency index of 36 and a retention index of 73. According to the strict consensus and preferred character state optimisations, the examined species of Lachnopus do not form a monophyletic entity, even though most species are placed in a major L. coffeae–L. guerinii ingroup clade, which stands in sister relationship to Ischionoplus. This major clade is constituted by at least three well-recognisable subgroups, each associated with a particular geographic range in the Caribbean region: (1) a widely distributed L. coffeae–L. lineicollis ‘grade’; (2) the Hispaniolan L. proteus–L. mercator clade; (3) and the L. hispidus–L. guerinii clade with species from Cuba, Florida and Jamaica. This study provides a sound phylogenetic basis for future revisions of Lachnopus and related geonemine genera.

Testing the monophyly of the New Zealand and Australian endemic family Conoesucidae Ross based on combined molecular and morphological data (Insecta: Trichoptera: Sericostomatoidea)

Conoesucidae (Trichoptera, Insecta) are restricted to SE Australia, Tasmania and New Zealand. The family includes 42 described species in 12 genera, and each genus is endemic to either New Zealand or Australia. Although monophyly has been previously assumed, no morphological characters have been proposed to represent synapomorphies for the group. We collected molecular data from two mitochondrial genes (16S and cytochrome oxidase I), one nuclear gene (elongation factor 1‐α) (2237–2277 bp in total), and 12 morphological characters to produce the first phylogeny of the family. We combined the molecular and morphological characters and performed both a maximum parsimony analysis and a Bayesian analysis to test the monophyly of the family, and to hypothesize the phylogeny among its genera. The parsimony analysis revealed a single most parsimonious tree with Conoesucidae being a monophyletic taxon and sistergroup to the Calocidae. The Bayesian inference produced a distribution of trees, the consensus of which is supported with posterior probabilities of 100% for 15 out of 22 possible ingroup clades including the most basal branch of the family, indicating strong support for a monophyletic Conoesucidae. The most parsimonious tree and the tree from the Bayesian analysis were identical except that the ingroup genus Pycnocentria changed position by jumping to a neighbouring clade. Based on the assumption that the ancestral conoesucid species was present on both New Zealand and Australia, a biogeographical analysis using the dispersal‐vicariance criteria demonstrated that one or two (depending on which of the two phylogenetic reconstructions were applied) sympatric speciation events took place on New Zealand prior to a single, late dispersal from New Zealand to Australia.

Molecular and morphological systematics of hypogean schizomids (Schizomida:Hubbardiidae) in semiarid Australia

We used molecular and morphological techniques to study troglobitic schizomids inhabiting a variety of subterranean landforms in semiarid Western Australia. The study was designed to explore the taxonomic and phylogenetic status of newly discovered populations of subterranean schizomids. Molecular sequences of the mitochondrial DNA (mtDNA) cytochrome oxidase subunit 1 (COI) and small subunit rRNA (12S) were obtained from a total of 73 schizomid specimens. Populations sampled from boreholes within mesa landforms in the Robe Valley were highly genetically distinct from species of Draculoides Harvey, 1992 found elsewhere in the Pilbara (Cape Range and Barrow Island). Pronounced genetic structuring was also evident at a fine spatial scale within the Robe Valley, with populations from each of the mesas examined exhibiting unique and highly divergent mtDNA lineages. These molecular data were generally supported by small but significant morphological features, usually in the secondary male structures, but some species were represented only by female specimens that possessed more conservative morphologies. The molecular data defined two major in-group clades, which were supported by morphological differences. One clade was widespread and included the type species of Draculoides, D. vinei (Harvey), along with D. bramstokeri Harvey & Humphreys, D. brooksi Harvey, D. julianneae Harvey, D. mesozeirus, sp. nov. and D. neoanthropus, sp. nov. The second clade was restricted to the Robe Valley and deemed to represent a new genus, Paradraculoides, which included four new species P. anachoretus, sp. nov., P. bythius, sp. nov., P. gnophicola, sp. nov. and P. kryptus, sp. nov. (type species).

Phylogeny of Milkweed Tussocks (Arctiidae: Arctiinae: Phaegopterini) and Its Implications for Evolution of Ultrasound Communication

Some have proposed that the evolution of ultrasound as a male courtship cue in milkweed tussocks has arisen stepwise through transitional stages of redundant courtship cues (both pheromone and ultrasound). Furthermore, it has been proposed that ultrasound courtship was evolutionarily linked to a switch to cardenolide-producing larval host plants. To test the hypothesis of a transitional stage and proposed correlation of the behavior with larval hosts, the phylogenetic relationships of the phaegopterines Euchaetes, Cycnia, Ectypia, Pygarctia, and Pareuchaetes were assessed with 88 characters (205 states) derived from adult morphology. The resulting strict consensus placed the Pareuchaetes among the outgroup genera and recovered the remaining four genera as a clade. Euchaetes was not monophyletic; however, removal of Pareuchaetes results in monophyly. Based on this phylogeny, the ancestor of the ingroup clade fed on cardenolide-containing host plants, possessed well developed tymbals and lacked coremata (pheromone distributing structures). Thus, the use of male ultrasound in courtship was correlated with larval feeding on cardenolide hosts. The hypothesis that ultrasound first arose as a redundant courtship cue, with a subsequent loss of the male pheromone, was not supported by our data. Rather, male pheromone courtship was reacquired after ultrasound courtship in two lineages, based on the presence of coremata and behavioral data.

Review of generic limits of the tiger moth genera Virbia Walker and Holomelina Herrich-Schäffer (Lepidoptera: Arctiidae: Arctiinae) and their biogeography

A phylogeny for Holomelina and Virbia was constructed based on adult morphology. This study included 42 of the 70 recognized species in Holomelina and Virbia, two new species, and two elevated synonyms (H. marginata Druce and V. inversia Edwards). The following outgroups were chosen from the Arctiini: Arachnis picta Packard, Hypercompe permaculata Packard, Paracles fusca Walker, Phragmatobia fuliginosa (Linneaus), Pyrrarctia isabella (J. E. Smith), and Spilosoma virginica (Fabricius). Fifty-eight characters (195 states) were described and scored, including several newly discovered characters of the male and female abdomen and genitalia. Three male character systems, the shape and orientation of the basiphallus; the shape, orientation and ornamentation of the endophallus; and the pattern of sclerotization of the eighth male sternite, were important diagnostic characters for smaller clades and species. The maximum parsimony analysis of all taxa with 100 randomized repetitions yielded 779 trees with a length (L) of 1698, a consistency index (CI) of 0.41 and a retention index (RI) of 0.65. Holomelina species were placed in two clades, an H. aurantiaca clade and an H. opella clade that was paraphyletic with respect to Virbia. The ingroup clade comprised of Holomelina and Virbia species was supported by eleven synapomorphies and a Bremer index higher than fifteen. The decay of this node was greater than 10. Taxon jack-knife analyses revealed that results were sensitive to the inclusion of two outgroup species, A. picta and P. fuliginosa. When either was removed the relationships of ingroup taxa became unresolved. When the analysis was restricted to species which represented by both males and females, we recovered an ingroup consisting of Holomelina and Virbia species, but the strict consensus of 123 trees (L = 1356, CI = 0.47, RI = 0.65) resulted in a polytomy of smaller clades. Neither analysis recovered a reciprocally monophyletic Holomelina and Virbia. We place Holomelina as a junior generic synonym of Virbia and provide a species checklist for Virbia. Biogeographic areas were scored from distribution information in a specimen-level database consisting of over 12,000 specimens. The following eight areas were recognized: A. western USA and Canada, B. central USA and Canada, C. eastern USA and Canada, D. southwestern USA and Mexico, E. central America, F. western South America, G. eastern and central South America, and H. highlands of northern South America. A weighted ancestral area analysis was used to determine ancestral areas for five major ingroup clades. We propose replacing the name “probability index” (PI) of Bremer with the term “Weighted Area Index (WAI)” because the PI of Bremer is not based on a statistical distribution and the term is misleadingWe determined that the ancestral area for Virbia was most likely the southwestern United States and Mexico. The species distributed in North America north of Mexico represent a composite fauna with a minimum of two dispersal events from Mexico into the northern latitudes. In contrast, the South American fauna appears to have a single origin.

Origin and diversification of the endemic Hawaiian tree snails (Achatinellidae: Achatinellinae) based on molecular evidence

Tree snails of the endemic subfamily Achatinellinae comprise a diverse and important component of the Hawaiian fauna. In recent decades anthropogenic impacts have resulted in devastating extinction rates in Hawaiian tree snails. To address long-standing biogeographic, systematic, and evolutionary questions we used cytochrome c oxidase subunit I (COI) gene sequences to reconstruct the phylogeny of 23 extant species spanning the range of the subfamily from five Hawaiian Islands. To investigate family-level relationships, data were analyzed from 11 terrestrial pulmonate families. Although nodal support for monophyly of the endemic Pacific family Achatinellidae and endemic Hawaiian subfamily Achatinellinae was strong, bifurcation order among deeper ingroup nodes was not well-supported by bootstrap resampling. We hypothesize that lineage extinction and rapidity of lineage formation may have rendered evolutionary reconstruction difficult using a standard phylogenetic approach. Use of an optimized evolutionary model, however, improved resolution and recovered three main clades. The diversification pattern inferred contradicts the traditional biogeographic hypothesis of a Maui origin of the achatinelline lineage. Taxa comprising the basal ingroup clade ( Achatinella spp.) and seeding lineages for subsequent clades originated on O‘ahu. Therefore it appears that the ancestral colonizing species of achatinellines arrived first on O‘ahu from an unknown source, and that O‘ahu is the Hawaiian origin of the subfamily. Species previously defined by morphological criteria were generally found to be phylogenetically distinct, and the overall colonization pattern follows the island-age progression rule with several instances of generic polyphyly and back-colonization.

High levels of mitochondrial cytochrome b divergence in annual killifishes of the genus Cynolebias (Cyprinodontiformes, Rivulidae)

Phylogenetic relationships based on 324 base pairs of the mitochondrial cytochrome b gene were examined in 14 species of the genus Cynolebias. The monophyly of the genus relative to three outgroup taxa belonging to the family Rivulidae was supported by the sequence data. Bootstrap values corroborated the existence of intrageneric monophyletic units in both parsimony and Neighbour-joining analyses. These include a Cynolebias bellottii–C. wolterstorffi clade, a C. adloffi-2, C. duraznensis, C. viarius and C. adloffi-1 group, and a C. gymnoventris,C. luteoflammulatus pair and the strongly supported assemblage that includes C. prognathus and C. cheradophilus clade. Phylogenetic relationships remain poorly supported for C. nigripinnis, C. affinis and C. alexandri, and unresolved among the previous ingroup clades. Cytochrome b sequences reveal an unexpectedly high level of divergence among species of the genus Cynolebias. Consequently, cytochrome b shows good resolution of recent cladogenetic events but limited phylogenetic information at deeper nodes. High levels of sequence divergence span a broad range withinCynolebias . The highest sequence divergence (c. 28%) occurred among C. antenori and the remaining species of the genus. The minimum divergence value (4.5%) is exhibited by sympatric species C. cheradophilus andC. prognathus .

High levels of mitochondrial cytochrome b divergence in annual killifishes of the genus Cynolebias (Cyprinodontiformes, Rivulidae)

Phylogenetic relationships based on 324 base pairs of the mitochondrial cytochrome b gene were examined in 14 species of the genus Cynolebias. The monophyly of the genus relative to three outgroup taxa belonging to the family Rivulidae was supported by the sequence data. Bootstrap values corroborated the existence of intrageneric monophyletic units in both parsimony and Neighbour-joining analyses. These include a Cynolebias bellottii-C. wolterstorffl clade, a C. adloJfi-2, C. duraznensis, C. viarius and C adloffi-1 group, and a C. gymnoventris, C. luteoflammulatus pair and the strongly supported assemblage that includes C. prognalhus and C. cheradophilus clade. Phylogenetic relationships remain poorly supported for C. nigripinnis, C. affinis and C. alexandri, and unresolved among the previous ingroup clades. Cytochrome b sequences reveal an unexpectedly high level of divergence among species of the genus Cynolebias. Consequently, cytochrome b shows good resolution of recent cladogenetic events but limited phylogenetic information at deeper nodes. High levels of sequence divergence span a broad range within Cynolebias. The highest sequence divergence (c. 28%) occurred among C. antenori and the remaining species of the genus. The minimum divergence value (4.5%) is exhibited by sympatric species C. cheradophilus and C. prognathus.

A trnL-F based phylogeny for species ofPelargonium (Geraniaceae) with small chromosomes

Phylogenetic analysis was performed of 921 positions of trnL (UAA) 5′ exon — trnF (GAA) exon chloroplast DNA regions from 68 representatives ofPelargonium sectt.Campylia, Cortusina, Glaucophyllum, Hoarea, Isopetalum, Ligularia, Otidia, Pelargonium, Peristera, Polyactium, andReniformia, together with five putative outgroup species from sectionsCiconium, Chorisma andJenkinsonia. The total data set therefore comprised 67.2 kb of DNA sequence. Two main ingroup clades were identified: one clade contains sectionsPeristera, Reniformia, andIsopetalum, the other contains sectionsCampylia, Cortusina, Glaucophyllum, Hoarea, Ligularia, Otidia, Pelargonium, Polyactium and two species currently grouped in sect.Peristera. Branching order among five main clades within the latter clade was not resolved. The trnL-F sequence data support monophyly only for sectionsReniformia andHoarea, the remainder of the currently recognized sections ofPelargonium being either paraphyletic or polyphyletic. The data further suggest that sect.Polyactium is diphyletic and that sect.Glaucophyllum is nested within sect.Pelargonium. One relatively derived clade, which represents half of the genus, contains predominantly geophytic and succulent species, occurring in the geographically restricted winter rainfall region of the South African Cape. This pattern is interpreted as reflecting explosive radiation, possibly as an adaptive response to recent aridification in the western Cape.

Molecular systematics and a revised taxonomy of the onocleoid ferns (Dryopteridaceae: Onocleeae)

Nucleotide sequences of the chloroplast-encoded rbcL gene were determined for all five species of the onocleoid ferns (Dryopteridaceae tribe Onocleeae), including both varieties of Onoclea sensibilis, and for outgroup member Blechnum glandulosum. Together with GenBank sequences of three additional onocleoid accessions and four additional taxa representing the outgroup Blechnaceae, these were analyzed cladistically under the optimality criteria of maximum parsimony and maximum likelihood. Maximum parsimony yielded a single most-parsimonious tree with the three accessions of Onoclea sensibilis var. sensibilis left as an unresolved trichotomy. Maximum likelihood yielded a single set of three optimal trees whose only topological variation was in the trivial positioning of the three accessions of Onoclea sensibilis var. sensibilis relative to each other. Thus tree topologies of the onocleoid ingroup under maximum parsimony and maximum likelihood were completely congruent. Matteuccia orientalis and M. intermedia formed the basalmost ingroup clade strongly separated from the remaining taxa and sister to them. Onoclea sensibilis is strongly separated from its sister clade of Matteuccia struthiopteris plus Onocleopsis hintonii, and the two varieties of Onoclea sensibilis are well differentiated from each other. Matteuccia struthiopteris and Onocleopsis hintonii form the least strongly supported clade. Levels of sequence divergence among onocleoid taxa are compared with values from other taxa, and morphological and chromosomal data used in previous, noncladistic studies of the onocleoids are reevaluated in light of this rbcL phylogeny. Matteuccia orientalis and M. intermedia should be recognized in their own genus Pentarhizidium, and the previously recognized varieties of Onoclea sensibilis are supported at least at varietal rank and possibly at the rank of species. Molecular and morphological data bearing on the circumscriptions of Matteuccia and Onocleopsis are equivocal but perhaps most concordant with their continued recognition as monotypic genera.