Combination Of Character States Research Articles

Revisiting the octopicolid copepods (Octopicolidae: Octopicola Humes, 1957): comparative morphology and an updated key to species

A review of the present state of knowledge on the octopicolid copepods (Octopicolidae: Octopicola Humes, 1957) is presented. Characteristic morphological features are illustrated with scanning electron micrographs of Octopicola superba superba Humes, 1957. Comparative morphology analysis led to the conclusion that there is sufficient evidence to justify raising the two subspecies of O. superba to full species rank. A new identification key for the four species of Octopicola Humes, 1957, i.e. O. superba Humes, 1957, O. antillensis Stock, Humes & Gooding, 1963, O. stocki Humes, 1963 and O. regalis Humes, 1974, is proposed after evaluation of the morphological characters which vary more markedly between them. Among other characters, these species differ in the ornamentation of the third antennal segment, maxilla and male maxilliped. They are further distinguished by a combination of several character states concerning the fifth pedigerous somite.

Rediscovery and re-description of Ischnocnema nigriventris (Lutz, 1925) (Anura: Terrarana: Brachycephalidae).

Besides its brief original description in 1925 and information provided by one specimen collected in the 1980s, nothing else is known for Ischnocnema nigriventris (Lutz, 1925). Also, the poor preservation of the type series has hindered the association of this name with any known population of Ischnocnema. Fieldwork in Bertioga municipality, state of Sao Paulo, Brazil, revealed a population of Ischnocnema to which we were able to apply the name Ischnocnema nigriventris. We report this rediscovery, re-describe the species on the basis of the newly found specimens, and describe its mating call. Ischnocnema nigriventris is allocated to the Ischnocnema lactea species series and we propose a diagnosis based on a combination of morphological character states. The species is diagnosed by prominent conical tubercles on the upper eyelid, inguinal region and hidden areas of the hind limbs with yellow mottling in males and orange mottling in females, and by its advertisement call composed of two to four non-pulsed notes, the first one differentiated from the others by its lower intensity and different frequency. Ischnocnema nigriventris is only known from Parque das Neblinas (Bertioga municipality), Paranapiacaba (Santo André municipality), and Boracéia (Salesópolis municipality), localities in the state of São Paulo, Brazil

Micromorfología foliar del género Quercus (Fagaceae) en México

México es el principal centro de diversidad del género Quercus en el Hemisferio Occidental. A pesar de los avances recientes en el conocimiento de los encinos mexicanos, aún existe cierto grado de confusión taxonómica, particularmente dentro de algunos complejos de especies. En este estudio se utilizó microscopía electrónica de barrido para describir las estructuras micromorfológicas foliares (tricomas, ceras epicuticulares y estomas) de ambas superficies foliares (haz y envés) en representantes seleccionados de encinos mexicanos, con el objetivo de evaluar la utilidad taxonómica de estos caracteres. En total se examinaron 27 especies, pertenecientes a las secciones Quercus (encinos blancos) y Lobatae (encinos rojos y negros), incluyendo varios grupos de plantas cercanamente relacionadas con problemas de delimitación taxonómica, así como especies que presentan hibridación. Se encontraron varios tipos de tricomas, tanto los de tipo glandular (simples y bulbosos) como los no glandulares (solitarios, multirradiados, estrellados, estrellados fusionados y fasciculado estipitados). Las ceras epicuticulares se observaron en forma de capas, capas fisuradas, costras, gránulos, placas y placas en rosetas. Los estomas fueron elípticos y se encontraron elevados con respecto a la superficie foliar o al mismo nivel que ésta. Entre los tres tipos de estructuras examinadas, los tricomas fueron los más útiles para propósitos taxonómicos, seguidos por las ceras epicuticulares. Todas las especies tuvieron diferentes combinaciones de estados de carácter para estas estructuras micromorfológicas, lo que permitió la elaboración de claves para identificar a las especies dentro de los grupos problemáticos.

First Discovery of Plasmobatidae (Acari, Oribatida) in Gabon, redefinition and new species of the genusSolenozetesGrandjean, 1932

ABSTRACT The family Plasmobatidae is herewith reported for the first time from Gabon and Solenozetes makokouensis n. sp. is described. The holotype was collected from litter in dense evergreen humid forest, Makokou, Province Ogoove-Ivindo, north-eastern Gabon. The species is described and illustrated based on adult females. The new species is easily distinguishable by the following combination of character states: cerotegumental layer amorphous, with small tuberculations mixed with granulations; ro seta falcate, inserted on forward-extending visorlike protuberance; in seta wrapped in cerotegumental granulate layer; prodorsal anterior zone complex with irregular protuberances; bothridium horseshoe-shaped, internal rings appearing gear-like; sensillus filiform; notogaster anteriorly with semicircular furrow and depressed zone; macropores opening on foveae; notogastral microsculpture irregular foveate pattern; opisthosomal gland apophysis conical with rounded tip; five pairs of notogastral setae. The genus S...

Genetic Identity and Diversity of Perennial Pepperweed (Lepidium latifolium) in Its Native and Invaded Ranges

AbstractPerennial pepperweed is an invasive plant species in North America, native to temperate Eurasia and northern Africa. Effective biological control depends upon correct taxonomic identification. Therefore, we investigated morphological and genetic data (cpDNA sequences and amplified fragment length polymorphisms [AFLP]) in its native range, where the species is at times treated as multiple taxa (L. latifolium, L. affineandL. obtusum). We also analyzed genetic data to determine the number and distribution of haplotypes and genotypes in the invaded range. Using Bayesian analysis, we found three clusters of AFLP genotypes in the native range, but little correlation between these clusters and morphological characters used to distinguish taxa. Also, we found combinations of morphological character states within many native range plants that are incompatible with current species descriptions, offering no support for splittingL. latifoliumsensu lato into three species. In North America 97% of the genetic variation was among populations and there were only eight AFLP genotypes in 288 plants, suggesting few introductions or a severe bottleneck, and little or no creation of new genotypes since introduction. We found plants in the native range that are genetically similar (88 to 99%) to six of the eight invasive AFLP genotypes, suggesting that Kazakhstan and China are origins for much of the North American invasion.

A semantic model for species description applied to the ensign wasps (hymenoptera: evaniidae) of New Caledonia.

Taxonomic descriptions are unparalleled sources of knowledge of life's phenotypic diversity. As natural language prose, these data sets are largely refractory to computation and integration with other sources of phenotypic data. By formalizing taxonomic descriptions using ontology-based semantic representation, we aim to increase the reusability and computability of taxonomists' primary data. Here, we present a revision of the ensign wasp (Hymenoptera: Evaniidae) fauna of New Caledonia using this new model for species description. Descriptive matrices, specimen data, and taxonomic nomenclature are gathered in a unified Web-based application, mx, then exported as both traditional taxonomic treatments and semantic statements using the OWL Web Ontology Language. Character:character-state combinations are then annotated following the entity–quality phenotype model, originally developed to represent mutant model organism phenotype data; concepts of anatomy are drawn from the Hymenoptera Anatomy Ontology and linked to phenotype descriptors from the Phenotypic Quality Ontology. The resulting set of semantic statements is provided in Resource Description Framework format. Applying the model to real data, that is, specimens, taxonomic names, diagnoses, descriptions, and redescriptions, provides us with a foundation to discuss limitations and potential benefits such as automated data integration and reasoner-driven queries. Four species of ensign wasp are now known to occur in New Caledonia: Szepligetella levipetiolata, Szepligetella deercreeki Deans and Mikó sp. nov., Szepligetella irwini Deans and Mikó sp. nov., and the nearly cosmopolitan Evania appendigaster. A fifth species, Szepligetella sericea, including Szepligetella impressa, syn. nov., has not yet been collected in New Caledonia but can be found on islands throughout the Pacific and so is included in the diagnostic key. [Biodiversity informatics; Evaniidae; New Caledonia; new species; ontology; semantic phenotypes; semantic species description; taxonomy.]

A new fossil salamander (Caudata, Proteidae) from the Upper Cretaceous (Maastrichtian) Hell Creek Formation, Montana, U.S.A.

ABSTRACT North American Late Cretaceous salamanders are principally known by isolated atlantes and trunk vertebrae. Here I describe a new genus and species of fossil salamander, Paranecturus garbanii, gen. et sp. nov., based on these elements from the lower portions of the Hell Creek Formation (Maastrichtian), Garfield County, northeastern Montana, U.S.A. It is diagnosed by a unique combination of character states that include the presence of an alar-like process of the atlas and a groove on the posterior face of the neural arch and solid dorsal rib-bearers of the trunk vertebrae. My phylogenetic analysis of 13 caudate taxa and 23 atlantal and trunk vertebral characters recovered P. garbanii as a member of the Proteidae. P. garbanii represents the oldest fossil record of the Proteidae and demonstrates that this lineage was present before the Cretaceous-Paleogene mass extinction event. SUPPLEMENTAL DATA—Supplemental materials are available for this article for free at www.tandfonline.com/UJVP

The oldest dinosaur? A Middle Triassic dinosauriform from Tanzania

The rise of dinosaurs was a major event in vertebrate history, but the timing of the origin and early diversification of the group remain poorly constrained. Here, we describe Nyasasaurus parringtoni gen. et sp. nov., which is identified as either the earliest known member of, or the sister-taxon to, Dinosauria. Nyasasaurus possesses a unique combination of dinosaur character states and an elevated growth rate similar to that of definitive early dinosaurs. It demonstrates that the initial dinosaur radiation occurred over a longer timescale than previously thought (possibly 15 Myr earlier), and that dinosaurs and their immediate relatives are better understood as part of a larger Middle Triassic archosauriform radiation. The African provenance of Nyasasaurus supports a southern Pangaean origin for Dinosauria.

The diversity of cimolestan mammals within the White River Group of South Dakota and Nebraska

A new Late Carboniferous cnemidolestodean insect Xixia huban gen. et sp. nov., is described from the Xiaheyan locality (Zhongwei City, Ningxia, China; Tupo Formation). Its combination of character states and observed variants on the wing venations are indicative of homologies shared with more derived members of the order. In particular it is ascertained that MP runs fused with CuA and CuA + CuPa, but is not usually visible as a distinct vein. The new genus exhibits previously unknown coloration pattern composed of dark patches distributed over the whole forewing (as opposed to regular stripes or rows of spots, previously documented in the group). The comparatively abundant sample makes it one of the best documented cnemidolestodeans to date.

Two new species of the genus Parosphromenus (Teleostei: Osphronemidae) from Sumatra

Parosphromenus gunawani spec. nov. (previously known as P. spec. “Danau Rasau”) and P. phoenicurus spec. nov. (previously known as P. spec. “Langgam”) are described from Sumatra, Indonesia. Parosphromenus gunawani is similar to P. bintan but differs from it by lacking the prominent broad blue band in the caudal fin of males. It is distinguished from the remaining species of the genus by an unique combination of meristic and colour character states. Parosphromenus phoenicurus can be distinguished from all the other species of the genus by the rhombic caudal-fin shape in males (versus round or lanceolate with filaments).

Asexual propagules as an adaptive trait for epiphylly in tropical leafy liverworts (Lejeuneaceae)

Many links between form and function are described in the context of adaptation. Several morphological and life-history traits in the leafy liverwort family Lejeuneaceae (Marchantiophyta) have been hypothesized to be adaptations for living on the surface of leaves of vascular plants (epiphylly). There have been, however, no rigorous tests of these hypotheses. Using a recently published phylogeny of Lejeuneaceae and trait data from published monographs, I tested the correlations of putative adaptive traits with the incidence of epiphylly. Both cross-species and phylogenetic-based analyses of trait data were performed to distinguish the patterns of shared evolutionary history from independent origins of putatively adaptive traits. The rates of transitions between different combinations of character states were also calculated to determine whether traits were more likely to evolve in the presence of epiphylly. Only one trait, production of asexual propagules, was correlated with epiphylly in the phylogenetic-based analysis. The rate of transition to asexual propagules was also significantly higher in the presence of epiphylly. Other traits correlated with epiphylly appeared to be the results of shared evolutionary history among sister taxa and therefore not due to adaptive evolution. The present study distinguished production of asexual propagules from other traits as a key adaptive response to living on the leaf surface. No other putative "adaptive" traits to epiphylly showed evidence of being specific adaptation to epiphylly. The results highlight the importance of phylogenetically controlled methods in determining an adaptive function of traits.

Tyrannosauroid dinosaurs from the Aguja Formation (Upper Cretaceous) of Big Bend National Park, Texas

ABSTRACTRare remains of tyrannosauroid dinosaurs from the Aguja Formation in West Texas indicate the presence here of a relatively gracile species, comparable in form and adult size toAppalachiosaurusor subadult albertosaurines,GorgosaurusandAlbertosaurus. Histologic analysis of one of the specimens indicates that the Aguja tyrannosaur attained an adult size substantially smaller than adult albertosaurines (700 kg, 6·5 m body length). The frontal bone is narrow with a wide orbital slot and a bipartite joint for the postorbital, features thought to be diagnostic of Albertosaurinae; but there is a tall sagittal crest and reduced parietal wedge separating the frontals on the midline, features thought to be diagnostic of Tyrannosaurinae. The tall sagittal crest may be a synapomorphy of Tyrannosaurinae, and the Aguja tyrannosaur is herein referred to that clade. However, the unique combination of character states exhibited by the frontal prevents confident attribution to any known species. The Aguja tyrannosaur provides further evidence that North American Campanian tyrannosauroids were remarkably diverse for such large predators, and that each species was apparently endemic to a relatively small geographic province.

Tramps, narrow endemics and morphologically cryptic species in the epiphyllous liverwort Diplasiolejeunea

Diplasiolejeunea is a pantropical, epiphytic genus of leafy liverworts that occurs from the lowlands to more than 4000m altitude. Phylogenetic analyses of a molecular dataset consisting of three markers (nuclear ribosomal ITS region, plastidic trnL-F region and rbcL gene) and 122 accessions (plus two outgroups, Colura and Cololejeunea) indicate that the evolutionary diversity of Diplasiolejeunea is underestimated by current morphology-based classification. Four morphologically semi-cryptic species have been recovered. The molecular phylogenies support a deep split into a Neotropical and a Paleotropical clade, the latter structured into Australasian, Asian and Afromadacascan lineages. Presented results confirm the ranges of two pantropical species (D. cavifolia, D. rudolphiana), provide evidence for dispersal from the Neotropics into the Paleotropics, indicate speciation along altitudinal gradients and demonstrate extensive morphological homoplasy. We propose a revised supraspecific classification of Diplasiolejeunea into a predominantly Paleotropical subgenus Physolejeunea and predominantly Neotropical subgenera Austrolejeuneopsis and Diplasiolejeunea, the former containing mainly epiphytic species, the latter mainly epiphylls. Several clades are supported by combinations of morphological character states, and could be assigned to sections at some later point. This is the first comprehensive phylogeny of a largely epiphyllous genus of liverworts.

A new species of Laetacara from the northern Brazil coastal floodplains (Teleostei: Cichlidae)

Laetacara flamannellus sp. n. is described from the northern Brazilian coastal floodplains. It differs from all its congeners by a combination of character states: presence of a dark brown or black spot located on dorsal-fin base; presence of a yellow stripe on the middle portion of the dorsal fin, crossing the whole fin; presence of an orange ring around the dorsal-fin base spot; absence of a red pigmentation on belly; absence of a red or purple zone on the anal-fin base; absence of a red zone on the caudal-fin base; cycloid scales on the opercle plate; and trunk scales above upper lateral line with few ctenii. The presence of a deep notch in the dorsal margin of the anterior ceratohyal, often used as diagnostic for Laetacara, is confirmed here for L. flamannellus.

New Genus and Species of Nectar-Feeding Bat from the Atlantic Forest of Southeastern Brazil (Chiroptera: Phyllostomidae: Glossophaginae)

The lowland Atlantic Forest of southeastern Brazil is well known for its biological diversity and numerous endemic taxa. Among bats collected recently at the Reserva Natural Vale and previously in the Floresta Nacional de Goytacazes, in the state of Espírito Santo, are specimens easily recognized as nectar-feeding glossophagines, but that exhibit a suite of morphological traits that preclude assignment to any of the 10 genera currently recognized in the subfamily Glossophaginae. Here we describe a new genus and species based on this material. This new taxon, named Dryadonycteris capixaba, is diagnosed based on both external and craniodental features, including traits not seen in other nectar-feeding phyllostomids, such as similar-sized calcar and foot and inflated maxillary bones. The combination of character states seen in Dryadonycteris suggests that it belongs in the Tribe Choeronycterini, subtribe Choeronycterina, but the mosaic nature of primitive and derived states seen in this taxon precludes easy assessment of its relationships to other choeronycterine genera. Future explicit phylogenetic analyses of morphological data and DNA sequencing studies will be necessary to resolve its phylogenetic position within Choeronycterini.

A new phytosaur (Archosauriformes, Phytosauria) from the Lot’s Wife beds (Sonsela Member) within the Chinle Formation (Upper Triassic) of Petrified Forest National Park, Arizona

ABSTRACT A new phytosaur taxon from Petrified Forest National Park, Arizona, is here described based on cranial material from a single individual. This specimen previously was included in an extensive phylogenetic analysis, and it was found to possess a combination of character states that differs from all known phytosaur taxa in addition to two autapomorphies within the braincase and an autapomorphy of the mandible. The new taxon adds to the taxonomic diversity recognized from the Sonsela Member of the Chinle Formation. The continued increase in phytosaur diversity emphasizes the need to more accurately characterize and identify taxa within a phylogenetic systematic context in order to produce a more refined signal for biostratigraphic correlations, biochronologic inferences, and faunal dynamics during the Late Triassic.

A new Upper Carboniferous stem‐orthopteran (Insecta) from Ningxia (China)

Abstract During the Upper Carboniferous, orthopteran insects (grasshoppers, katydids, and crickets) were represented by numerous species distantly related to crown‐orthopterans, such as lobeattid and cnemidolestodean insects. The panorthopterans, including total‐orthopterans and their closest relatives, are represented by comparatively rarer species in localities of this period. Here we describe Heterologus duyiwuer sp. nov., an infrequent panorthopteran from the Late Carboniferous locality of Xiaheyan Village (Zhongwei City, Ningxia Hui Autonomous Region, China). The only available specimen is composed of an isolated forewing exhibiting a combination of character states previously unknown, in particular the lack of posterior radius (RP) / anterior Media (MA) connection, late branchings of the media (M) and anterior cubitus (CuA), and a branched posterior branch of the posterior cubitus (CuPb). Based on its unusual branching pattern, the composite stem resulting from the fusion of CuA and CuPaα (second anterior branch of CuP) is assumed to be composed of a branched CuA and a simple CuPaα.

†Caruso, a new genus of anglerfishes from the Eocene of Monte Bolca, Italy, with a comparative osteology and phylogeny of the teleost family Lophiidae

†Caruso, a new genus of lophiid anglerfishes, erected to contain †Lophius brachysomus Agassiz from the Eocene of Monte Bolca, Italy, is described and compared osteologically with all known representatives of the family, both extinct and extant. Together with †Sharfia, known from a single specimen also from Monte Bolca, †Caruso is the oldest member of the teleost family Lophiidae known to date, based on articulated skeletal remains. It possesses several autapomorphic features as well as a unique combination of character states that clearly separate it from all other known lophiiform fishes. Evidence is provided to show that †Sharfia is the sister group of all other known lophiid taxa, that †Caruso is most closely related phylogenetically to the extant genus Sladenia, and that these two genera together form the sister group of all remaining members of the family. †Caruso brachysomus is the third valid extinct taxon of the Lophiidae known from articulated skeletal remains. Candidate developmental mechanisms that may have contributed to the origin of the lophiiform body plan are also suggested. The hypertrophic development of the jaws, suspensorium, hyoid apparatus and branchial arches typical of lophiiforms may be correlated, possibly under the influence of discrete sets of genes. The synapomorphic elongation of the pectoral-fin radials might be related to a heterochronic shift of the onset of the expression of the gene sonic hedgehog. Finally, the highly modified spinous dorsal fin may have arisen by submodularization and co-option of the first dorsal-fin module, whereas its forward shift over the neurocranium may represent an anterior shift of the stripes of competency for dorsal-fin formation along the dorsal midline.

Predicting species richness in tropical lichenized fungi with ‘modular’ combinations of character states

A numerical approach is presented to predict the number of potentially undiscovered species in groups of species that exhibit a ‘modular’ combination of character states, that is, nearly ‘free’ combination that is not random but allows for a wide range of cross-combination of character states. The method uses a character correlation index (CCI) based on observed versus expected frequencies of pairwise character state combinations in known species of a group to estimate the degree of potentially positive and negative evolutionary constraints in the co-occurrence of character states. For instance, if a character state combination, such as ‘muriform ascospores’ and ‘presence of norstictic acid’, occurs less frequently than expected by chance, it indicates the presence of a negative evolutionary constraint. If no such constraint is detected, but a particular character combination has not yet been observed in a described species, it can be predicted that yet undescribed species will exhibit that combination. The method can therefore be used to detect ‘gaps’ in the taxonomic record, i.e. character combinations that have not been detected in known species but are likely to occur in undiscovered species. Modular combination of character states appears to be particularly common in Fungi, including lichens, and the method was here used to predict species richness in the lichen genus Graphis, specifically the Graphis scripta group. A total of 42 species is known in this group and, out of a total of 240 possible character state combinations, a further 48 were found likely to represent undiscovered species, suggesting that the real species number in this group is close to 90 or about twice the number now known. This agrees well with previous estimates for undiscovered diversity of tropical lichen species and for the first time gives a taxonomic prediction of how such ‘missing’ species are expected to look like.

Rogerzetes lacouturieri n. gen., n. sp., (Acari: Oribatida: Eremaeozetidae) from Madagascar

ABSTRACT Rogerzetes lacouturieri n. gen., n. sp., collected from Manankaza Forest Station, Ambohitantely, Tananarive, Madagascar, is described and illustrated based on adult specimens. The new genus is distinguishable by the following combination of character states: body flattened; posterior part of lamellae fused, forming plate-like structure with rounded margin partially covering the bothridia; central posterior part plate-like, with tongue-like expansion surpassing dorsosejugal furrow; end of longitudinal fissure round–ovoid; large apical lamellae; naso and prodorsal sensory structures present; tube between le and sensory structure; in absent; prehumeral tecta prominent, partially covering the bothridia; 10 pairs of notogastral setae; anal plate ending in long spine; legs heterotridactylous. The new species is characterized by the following combination of characteristics cuticular microsculpture of foveate pattern; prodorsum with internal well-developed sensory structure; anterior part of notogaster, between dorsosejugal furrow and lenticulus, flat, semi-circular area, and at lower level than surrounding notogaster; prehumeral tecta ear-like; notogastral setae short and rod-like; epimeral setation (3-1-1-2). Eremaeozetes betschi, Fernandez and Cleva, 2009, is transferred to the new genus.