Phylogenetic relationships of species of Crenicichla (Teleostei: Cichlidae) from southern South America based on the mitochondrial cytochrome b gene

Phylogenetic analysis using Bayesian inference, likelihood and parsimony methods was conducted on 60 complete mitochondrial cytochrome b sequences from 21 species of Crenicichla, including all species known from Uruguay (Crenicichla celidochilus, Crenicichla lepidota Crenicichla minuano, Crenicichla missioneira, Crenicichla punctata, Crenicichla scottii, Crenicichla vittata), Crenicichla compressiceps, Crenicichla empheres, Crenicichla geayi, Crenicichla iguassuensis, Crenicichla macrophthalma, Crenicichla menezesi, Crenicichla notophthalmus, Crenicichla regani, Crenicichla cf. regani, Crenicichla semifasciata, Crenicichla sveni, Crenicichla tendybaguassu, two unidentified species, and also two species of Teleocichla. Bayesian analysis resulted in a trichotomy with three major groups: (1) The C. missioneira species group (C. celidochilus, C. empheres, C. minuano, C. missioneira, C. tendybaguassu, and an undescribed species analyzed); (2) a group of southern species (C. iguassuensis, C. punctata, C. scottii, C. vittata); and (3) a rather heterogeneous group comprising the type species C. macrophthalma, members of the Crenicichla reticulata species group (C. geayi, C. semifasciata), members of the Crenicichla wallacii species group (C. compressiceps, C. notophthalmus, C. regani, C. cf. regani), members of the Crenicichla saxatilis species group (C. lepidota, C. menezesi, C. sveni, C. sp.), and two species of Teleocichla. Parsimony jackknifing resulted in a quadritomy with: (1) C. macrophthalma, (2) Teleocichla, (3) the saxatilis + wallacii group species, and (4) the rest, which include C. geayi and C. semifasciata as sister group to a dichotomy with the C. missioneira group and the remaining southern species. The sequence variation within the C. missioneira group is remarkably minor despite considerable morphological differences, supporting the conclusion that it forms an endemic species flock in the Uruguay River basin. Previously proposed species groups within the speciose genus Crenicichla (more than 90 species known) are partly corroborated. However, C. celidochilus was not previously associated with the C. missioneira species group, and C. vittata has not previously been associated with C. scottii, C. iguassuensis, or C. punctata. Crenicichla lepidota, C. sveni, C. menezesi and C. sp. represent the C. saxatilis group. Species of small size, representing the C. wallacii species group and Teleocichla are characterized by very long branches, and the position of Teleocichla differed considerably between the Bayesian and parsimony trees. This finding does not invalidate Teleocichla but rather suggests that the several monophyletic major clades within Crenicichla may need nominal recognition. A putative hybrid specimen with a morphology combining components from C. vittata and C. scottii, but with a cytochrome b sequence from C. scottii was found in a sample from the Rio Quaraí/Cuareim. Another putative hybrid specimen with a unique morphology but a cytochrome b sequence agreeing with C. scottii was found in a sample from Maldonado, but no other Crenicichla species than C. scottii is known from that locality.     

Seven new calochroid and fulvoid species of Cortinarius

We describe seven new European species of Cortinarius. All species are based on analyses of morphological and DNA sequence data. They all belong to a well-supported clade comprising most species traditionally treated in Cortinarius subgenus Phlegmacium sections Fulvi and Calochroi (i.e. the/Calochroi clade). All taxa are either fulvoid (containing anthraquinoid pigments) or calochroid (without these pigments). Morphological and ecological data are presented for all species and compared with similar species. A dichotomous key is presented for C. calochrous and similar species, including all six newly described calochroid species. The calochroid species C. albertii, C. chailluzii, C. cisticola, C. sancti-felicis, C. selandicus and C. vesterholtii spp. nov., and the fulvoid species C. langeorum sp. nov. are described.     

A new pelagic predatory pike cichlid (Teleostei: Cichlidae: Crenicichla) from the C. mandelburgeri species complex with parallel and reticulate evolution

The Crenicichla mandelburgeri species complex from the Middle Paraná shows parallel evolution of ecomorphs to the unrelated C. missioneira species complex from the Uruguay River. In this article, we describe a new species from the C. mandelburgeri species complex that has evolved a parallel morphology and ecology to an unrelated species from the C. missioneira species complex (C. celidochilus). The new species is a pelagic predator that feeds predominantly on fishes and together with C. celidochilus is the only known pelagic species in the large riverine genus Crenicichla. The new species is endemic solely to a small tributary (the Urugua-í) of the Middle Paraná River where it is sympatric and partly syntopic with two other closely related endemic species that, however, differ strongly in their ecomorphologies (one is a generalistic invertivore and the other a specialized molluscivore). Mitochondrial DNA phylogeny finds the new species nested within the widespread C. mandelburgeri. Reduced genome-representation ddRAD analyses, however, demonstrate that this new species is of a hybrid origin and shares ancestry with C. ypo, one of the two studied sympatric species.

     

Conidial surface ultrastructure of human-pathogenic and saprobicCladosporium species

Freeze-fracturing of outer wall layers ofCladosporium conidia revealed two types of ultrastructure, coinciding with taxonomic characteristics. The outer conidial layers were essentially smooth in the human pathogenic species,C. bantianum, C. carrionii, andC. trichoides. In contrast, mosaic arrays of rodlets on conidia were observed with freeze-fracturing in the saprobic species,C. cladosporioides, C. coralloides, C. herbarum, C. sphaerospermum, andC. variabile. Conidia ofC. elatum were an exception among the saprobic species as they had smooth surfaces. The present study supports the suggestion that the human pathogenicCladosporium species should be transferred to another genus.     

CHLOROPLAST DNA RESTRICTION SITE VARIATION AND THE PHYLOGENY OF COREOPSIS SECTION COREOPSIS (ASTERACEAE)

Restriction site variation in chloroplast DNAs (cpDNAs) of Coreopsis section Coreopsis was employed to assess divergence and phylogenetic relationships among the nine species of the section. A total of fourteen restriction site mutations and one length mutation was detected. Cladistic analysis of the cpDNA data produced a phylogeny that is different in several respects from previous hypotheses. CpDNA mutations divide the section into two groups, with the two perennial species C. auriculata and C. pubescens lacking any derived restriction site changes. The other seven species are united by five synapomorphic restriction site mutations and the one length mutation. These seven species fall into three unresolved clades consisting of 1) the remaining three perennial species, C. grandiflora, C. intermedia, and C. lanceolata; 2) three annual species, C. basalis, C. nuecensoides, and C. nuecensis; and 3) the remaining annual, C. wrightii. The cpDNA data suggest that, although the perennial habit is primitive within the section, the annual species of section Coreopsis have likely not originated from an extant perennial species. The estimated proportion of nucleotide differences per site (given as 100p) for the cpDNAs of species in the section ranges from 0.00 to 0.20, which is comparable to or lower than values reported for other congeneric species. The low level of cpDNA divergence is concordant with other data, including cross compatibility, interfertility and allozymes, in suggesting that species of the section are not highly divergent genetically.     

Phylogenetic relationships among annual and perennial species of the genus <Emphasis Type="Italic">Cicer</Emphasis> as inferred from ITS sequences of nuclear ribosomal DNA

The cladistic analysis of the DNA sequences of the internal transcribed spacers of ribosomal cistrons (ITS1 and ITS2) for 20 species of Cicer L. (among which all the annuals), shows that various sections of the genus are not monophyletic. Annual species do not form a clade: C. arietinum, in fact, is closely related to both C. echinospermum and C. reticulatum, whereas C. bijugum, C. judaicum, and C. pinnatifidum form a separate clade. The annual C. cuneatum is sister group to the perennial C. canariense and both are archaic species within the genus. C. yamashitae is, on the contrary, the only annual species belonging to a group of perennials, within which close relationships are evident between C. graecum and C. montbretii as well as among a group of mainly Asian species.     

<Emphasis Type="Italic">δ</Emphasis><Superscript>13</Superscript>C values and crassulacean acid metabolism in <Emphasis Type="Italic">Clusia</Emphasis> species from Panama

The genus Clusia is notable in that it contains arborescent crassulacean acid metabolism (CAM) plants. As part of a study of CAM in Clusia, titratable acidities were measured in 25 species and ¹³C values were measured for 38 species from Panamá, including seven undescribed species, and 11 species from Colombia, Costa Rica and Honduras. CAM was detected in 12 species. Clusia flava, C. rosea and C. uvitana exhibited ¹³C values or diurnal fluctuations in acidity indicative of strong CAM. In C. croatii, C. cylindrica, C. fructiangusta, C. lineata, C. odorata, C. pratensis, C. quadrangula, C. valerioi and C. sp. D diurnal fluctuations in acidity were consistent with weak CAM but the ¹³C values were C₃-like. All of the species that exhibited strong or weak CAM were in the C. flava or C. minor species groups. CAM was not detected in any member of the C. multiflora species group. Strong CAM species were not collected at altitudes above 680 m a.s.l. On the basis of ¹³C values, the expression of CAM was similar in terrestrial, hemi-epiphytic and epiphytic species and did not differ between individuals of the same species that exhibited different life-forms. This study indicates that phylogenetic affiliation may be a predictor of an ability to exhibit CAM in Clusia species from the Panamanian region, and that weak CAM is probably a common photosynthetic option in many Clusia species. ¹³C value is not a particularly good indicator of a potential of Clusia species growing in the field to exhibit CAM because it appears that the contribution in most species of CAM to carbon gain is generally rather small when integrated over the life-time of leaves.     

Morphological and molecular analyses support the existence of host-specific Peronospora species infecting Chenopodium

About 20 species of Peronospora have been reported to cause downy mildew on Chenopodium, but, particularly in plant pathology literature, only one species, P. farinosa, is considered to be involved. We performed sequence analysis of the ITS rDNA to reveal the phylogenetic relationships of Peronospora specimens from five species of Chenopodium, viz. C. album, C. ambrosioides, C. bonus-henricus, C. hybridum, and C. polyspermum. The five clades corresponded to particular Chenopodium species, and showed a high level of sequence divergence. Differences in the morphology of the conidia and ultimate branchlets also supported the separation of the five groups at the host species level. These results suggest that the names P. variabilis, P. boni-henrici, P. chenopodii, and P. chenopodii-polyspermi should be used for the four downy mildew pathogens specific to C. album, C. bonus-henricus, C. hybridum, and C. polyspermum, respectively. The Peronospora on C. ambrosioides was found to be an independent species.     

Catapyrenium (Verrucariaceae) species from South America

13 species of the lichen genusCatapyrenium are reported from South America. Five species (C. analogicum, C. andicolum, C. exaratum, C. lachneoides, andC. podolepis) are described as new. A key to the species known from South America is presented. Remarks on taxonomy, ecology and distribution of the species are given.Studies on the lichen genusCatapyrenium (Verrucariaceae) III. For second part seeBreuss (1991).     

Phylogeny and character evolution in the Indo-Pacific genus Ctenogobiops (Gobiiformes: Gobiidae)

Ctenogobiops is a genus of Indo-Pacific gobies that form obligate, mutualistic associations with shrimp in the genus Alpheus. This study provides a molecular phylogenetic analysis of eight Ctenogobiops species: C. aurocingulus, C. crocineus, C. feroculus, C. formosa, C. maculosus, C. mitodes, C. tangaroai, and C. tongaensis. We recover two clades within the genus, one consisting of C. feroculus and C. aurocingulus, the second including the remaining species arrayed as follows: (C. tongaensis (C. mitodes (C. formosa (C. maculosus (C. crocineus, C. tangaroai))))). Recovery of C. maculosus and C. crocineus as distinct taxa suggests that these species are not synonymous, although sampling in this study is limited. Species of Ctenogobiops are morphologically very similar to each other, with generally consistent meristic character states present throughout the genus. Recognition of species is based primarily on slight variations in color pattern, shape of the dorsal fin, and size of the gill opening. Comparison of our specimens of C. mitodes with accounts of C. pomastictus confirms that color pattern variations and lateral scale counts are more reliable indicators of species identity than relative dorsal fin spine length, particularly for smaller specimens. We evaluate the distribution of morphological characters in the context of the new phylogenetic hypothesis, and provide a summary of distinguishing characters for Ctenogobiops species. In this case, as in other instances of diverse reef-dwelling fish taxa, molecular data are ideal for inferring phylogenetic relationships, whereas morphological data remain the most expedient way to identify species.     

DNA SEQUENCE VARIATION IN THE RIBOSOMAL INTERNAL TRANSCRIBED SPACER REGION OF FRESHWATER CLADOPHORA SPECIES (CHLOROPHYTA)1

Freshwater species of Cladophora (Chlorophyta) are globally distributed and occupy an unusually wide range of ecological habitats. Delineating species is difficult because most easily observed morphological traits are highly variable and because sexual reproduction has not been clearly documented. Synthesizing ecological data on freshwater Cladophora species is problematic because it is unclear whether freshwater Cladophora species comprise many genetically distinct species or a few ecologically and morphologically variable and/or plastic species. We determined nucleotide sequences of the internal transcribed spacer (ITS) region of the nuclear ribosomal cistron of freshwater Cladophora species from a wide range of habitats and geographic locations. We compared these sequences to those derived from culture collections of C. fracta and C. glomerata, the two most commonly reported freshwater Cladophora species. Cladophora fracta and C. glomerata had very similar ITS sequences (95.3%). All other sequences were identical to those from the C. fracta or C. glomerata culture collections with the exception of one California sample that was similar to both C. fracta (95.6%) and C. glomerata (92.4%). ITS genotypes did not correlate with morphology or geography. This analysis shows that common freshwater Cladophora species comprise very few (possibly one) ecologically and morphologically variable species.     

The Pollen of Swedish Callitriche (Callitrichaceae) - Trends Towards Submergence

The pollen morphology of the six Swedish Callitriche L. species was investigated using LM, SEM and TEM. The surface ornamentation is reticulate in C. stagnalis, C. platycarpa, C. cophocarpa and C. palustris with supratectal elements in the first three species. The ectexine is reduced in C. hamulata, occurring as scattered granules, but completely absent in C. hermaphroditica. None of the species has distinct apertures but C. cophocarpa and C. platycarpa both have leptomata. The ultrastructure of the pollen wall in C. palustris shows regions with lamellated footlayer and an endexine which may function as apertures. Pollen morphology is found to be related to pollination biology; species with leptomata sometimes being pollinated through ephydrophily and species with reduced exine being hyphydrophilous.     

Seed coat anatomy ofCrossostylis (Rhizophoraceae): its evolutionary and systematic implications

The seed coat structure of all 13 species ofCrossostylis was studied to contribute to an understanding of species delimitation and relationships within the genus. The mature seed coat is relatively uniform and consistently constructed mainly by a well-developed exotesta and a well-developed fibrous exotegmen. The species differ in the thickness of the exotesta and exotegmen, the anatomy of exotestal cells, the presence and absence of persistent mesotesta, and so forth. On the basis of comparisons of these characters, close relationships are suggested in the species groups such as:Crossostylis banksiana andC. cominsii; C. biflora, C. raiateensis andC. multiflora; C. gandiflora, C. sebertii andC. imera; and five species in the Fiji Islands. These relationships except for those of Fijian five species are also supported by cladistics as their common characters are evaluated as synapomorphy. Species-level separation ofC. banksiana, C. pedunculata andC. raiateensis each from the closest species is doubted based on the results of seed coat structure.     

CYTOGENETIC STUDIES OF CARTHAMUS DIVARICATUS WITH ELEVEN PAIRS OF CHROMOSOMES AND ITS RELATIONSHIP TO OTHER CARTHAMUS SPECIES (COMPOSITAE)

Carthamus divaricatus (Beg. et Vacc.) Pamp., found only in Libya, has 11 pairs of chromosomes, a new chromosome number in the genus. The species is distinct morphologically. It has yellow, purple, and white corollas, yellow pollen, dark-purple striped anthers, horizontal branches, and strongly divaricate outer involucral bracts. The terminal portion of the middle involucral bracts is dentate and reddish brown. It is self-incompatible. Meiosis is regular in the different corolla-color types of C. divaricatus and their intraspecific hybrids. C. divaricatus was crossed to six species with n = 12, to three species with n = 10, to C. lanatus with n = 22, and to two species with n = 32 chromosomes. The morphological characteristics and cross-ability of the parental species plus the pollen viability, seed-set, and meiotic behavior of the hybrids involving C. divaricatus and other Carthamus species indicated that C. divaricatus is very closely related to species with n = 10, closely related to C. lanatus with n = 22, and less closely related to C. tinctorius with n = 12 chromosomes. C. divaricatus seems to be distantly related to C. nitidus (n = 12). It is proposed that C. divaricatus be included provisionally with 10-chromosome species in Section II. Alternative hypotheses for the development of the three basic chromosome numbers are discussed.     

Allozyme variation and phylogeny in annual species ofCicer (Leguminosae)

Allozymic variation at 30 isozyme loci was examined electrophoretically in nine annual and one perennial species ofCicer. While most of the accessions examined were monomorphic, species can be differentiated on the basis of their enzyme phenotypes. Several groups of species were identified based upon genetic distance values. For example,C. arietinum, C. reticulatum, andC. echinospermum shared the same alleles for most of the loci exmained. PerennialC. anatolicum is also closely related to this group. Similarly,C. judaicum, C. bijugum, andC. pinnatifidum formed another group. Two annual species,C. chorassanicum andC. yamashitae clustered together, whereasC. cuneatum was the most distantly related species. Correlations were found between genetic distances and geographic distribution. Results from enzyme electrophoresis tend to support the previously reported taxonomic treatments based upon crossability and morphological similarity. However,C. yamashitae, which has been classified in the second crossability group, is quite distinct genetically and morphologically from the remaining species of the group. An isozyme gene duplication observed in the genus suggested the monophyletic origin of the species examined in the present study.