Taxonomic studies on Senecio aquaticus (Asteraceae). A recommendation for the taxonomic status of Aquaticus and Barbareifolius

Aquaticus and Barbareifolius are two subgroups of Senecio aquaticus s. l. (Asteraceae) that have been recognized at various taxonomic levels in European taxonomic literature. In this paper we present the results of morphological studies on S. aquaticus s. l. in order to establish the taxonomic status of these two subgroups. Using multivariate morphometric techniques (discriminant analysis and principal component analysis), we examined the diagnostic potential of 60 macro-morphological characters and studied the delimitation of Aquaticus and Barbareifolius. Seven of the characters studied were included in the discriminant function of the stepwise discriminant analysis and are therefore considered to be the most reliable diagnostic characters of Aquaticus and Barbareifolius. Barbareifolius has, amongst other distinguishing traits, a more divaricate inflorescence than Aquaticus. Because both subgroups are distinct but overlapping clusters in multivariate morphometric space, and because clear differences in their geographical distribution are absent, we recommend the taxonomic status of variety to both groups. Following the rules of nomenclature, Aquaticus and Barbareifolius should therefore be named S. aquaticus Hill var. aquaticus and S. aquaticus Hill var. barbareifolius (Krock.) Wimm. & Grab.  © 2004 The Linnean Society of London, Botanical Journal of the Linnean Society , 2004, 145 , 489–498.     

Delimitation of sympatric Palaemon (Decapoda,Palaemonidae) species of the Laguna Madre,Mexico

Grass shrimps Palaemon mundusnovus, P. pugio and P. vulgaris sympatrically inhabit in the Laguna Madre, Mexico. They exhibit very close morphological similarity and overlap in their diagnostic characteristics, which has hindered certainty in their identification and has raised doubt regarding their taxonomic validity. In this work, we analyse intra‐ and interspecific morphometric and meristic variability through a multivariate analysis, and we determine the molecular variation using mitochondrial sequences of the 12S, 16S and COX1 genes to confirm the validity of the three taxa as having distinct lineages or to recognize a smaller number of species with phenotypic plasticity. Our results corroborate the taxonomic validity of the three species; however, there is intraspecific plasticity, interspecific overlap of characteristics and greater morphological and molecular similarity between the species P. mundusnovus and P. pugio, whereas P. vulgaris was better delimited. These species form a monophyletic group but the phylogenetic relationship obtained is discussed. Telson length was the primary variable in the principal components analysis, whereas the length of the propodus of the second pereiopod was the best discriminant. The range of variation reported in the characteristics linked to the rostrum is extended, and its influence on the separation of these species is recognized. The joint application of multivariate analysis from morphological variables and molecular tools is recommended to clarify the taxonomic status of species featuring close morphological similarity and sympatric distribution.     

Taxonomic notes on Gaudinia (Poaceae) based upon morphological characters and the first record of G. maroccana for Portugal

Gaudinia maroccana is considered a Moroccan endemism. After a qualitative and quantitative study of 13 morphological characters of Gaudinia based upon 43 specimens from western Iberian Peninsula, northwestern Morocco and Azores Islands, with statistical analyses (correspondence analysis, box-plots and cluster), we can conclude that this species also occurs on the coast of Portugal. Some taxonomic remarks on G. fragilis, G. coarctata, G. hispanica and G. valdesii and a key for determination are presented.     

The taxonomic status of some Atlanto‐Mediterranean species in the subgenus Holothuria (Echinodermata: Holothuroidea: Holothuriidae) based on molecular evidence

Molecular and morphological data were used to evaluate the taxonomic status of the species Holothuria tubulosa Gmelin, 1790 , Holothuria stellati Delle Chiaje, 1823 , Holothuria mammata Grube, 1840 , and Holothuria dakarensis Panning, 1939, belonging to the nominate subgenus Holothuria (Holothuria) (family Holothuriidae) from the Mediterranean Sea and Atlantic Ocean. A 16S rRNA marker distinguished three well‐supported clades with clear genetic differentiation amongst them. The morphometric characters, although they reflected the clades, showed great variability, and some specimens from different clades overlapped. The morphological data and the literature suggest that the clades correspond to H. dakarensis (from Cape Verde Islands), H. mammata (from the Atlanto‐Mediterranean area) and H. tubulosa (from the Mediterranean Sea). Holothuria stellati is considered here to be a junior subjective synonym of H. tubulosa. Great morphological intraspecific variation within H. tubulosa and H. mammata explains the confusion in the literature. Holothuria tubulosa includes specimens with distinctive ossicles, but others are similar to H. mammata. In these cases, the presence or absence of Cuvierian tubules proved a reliable indicator to the identity of these species; unfortunately this character is difficult to assess in preserved material. According to the results of discriminant analysis we propose a set of ossicle morphometric variables that permit the optimum assignation of individuals to the clades. Our results present a new perspective on the taxonomic status of species in Holothuria (Holothuria), and show how a molecular approach, combined with a morphological approach, can solve taxonomic problems.     

The taxonomic status of the Black Sea and Marmara Sea populations of the Broadnosed Pipefish Syngnathus cf. argentatus Pallas (Teleostei: Syngnathidae) based on morphological and molecular characters

As the Black Sea and Marmara Sea population of the Broadnosed Pipefish Syngnathus cf. argentatus show some morphological differences from the Mediterranean Sea populations, some authors regard it as an endemic species Syngnathus argentatus Pallas, 1814, while others consider it as a synonym of S. typhle Linnaeus, 1758. The aim of this study is to compare the populations of the Black Sea and Sea of Marmara with the Aegean Sea population, using a combination of morphological and molecular characters, in order to clarify their taxonomic status. Sampling was carried out at three stations in the Black Sea, two in the Sea of Marmara and three in the Aegean Sea, and a total of 24 morphometric and 6 meristic characters were examined. Metric data were analysed by Principal Component Analysis (PCA) and phylogenetic relationships between the populations were analysed using both cytochrome oxidase subunit 1 (COI) gene and cytochrome b (cyt-b) gene sequences. Although constant differences were observed in snout depths between the Black Sea/Marmara Sea and the Aegean Sea populations, other morphological features and genetic analysis did not enable these populations to be differentiated. These findings indicate that S. argentatus is a synonym of S. typhle.     

Taxonomy of the Solidago virgaurea Group (Asteraceae) in Poland,with Special Reference to Variability along an Altitudinal Gradient

The morphological differentiation and taxonomic treatment of lowland and high-mountain morphotypes within the Solidago virgaurea group are controversial. To clarify the taxonomic status of these taxa, we conducted a morphometric analysis of 1,746 individuals from 80 localities along an altitudinal gradient from the lowlands of northern Poland to the Carpathians and Sudetes of southern Poland. Multivariate morphometric analyses, cluster analyses and principal component analyses, were used to examine the morphological differentiation within the S. virgaurea group in Poland. Canonical discriminant analysis was applied to determine the morphological characters that best discriminate among the taxa. The stability of the high-mountain Solidago minuta morphotype was tested in an experimental field established in lowland Poland; individuals transplanted from various mountain sites were cultivated at this site, and the morphotypes remained stable in terms of their floral and vegetative characters. Multivariate analyses revealed two morphologically distinct taxa in the S. virgaurea group, which correspond to lowland S. virgaurea s. str. and high-mountain S. minuta as recognised in some European floras. The most important morphological characters for distinguishing the taxa are the number of tubular florets per capitulum, inner involucral bract width and involucre height. Vegetative and inflorescence characters appear to have less taxonomic value because they changed continuously with altitude. A key for identifying S. virgaurea and S. minuta in Poland is presented.     

Quantitative comparison of taxa and taxon concepts in the diatom genus Fragilariopsis: a case study on using slide scanning,multiexpert image annotation,and image analysis in taxonomy1

Semiautomated methods for microscopic image acquisition, image analysis, and taxonomic identification have repeatedly received attention in diatom analysis. Less well studied is the question whether and how such methods might prove useful for clarifying the delimitation of species that are difficult to separate for human taxonomists. To try to answer this question, three very similar Fragilariopsis species endemic to the Southern Ocean were targeted in this study: F. obliquecostata, F. ritscheri, and F. sublinearis. A set of 501 extended focus depth specimen images were obtained using a standardized, semiautomated microscopic procedure. Twelve diatomists independently identified these specimen images in order to reconcile taxonomic opinions and agree upon a taxonomic gold standard. Using image analyses, we then extracted morphometric features representing taxonomic characters of the target taxa. The discriminating ability of individual morphometric features was tested visually and statistically, and multivariate classification experiments were performed to test the agreement of the quantitatively defined taxa assignments with expert consensus opinion. Beyond an updated differential diagnosis of the studied taxa, our study also shows that automated imaging and image analysis procedures for diatoms are coming close to reaching a broad applicability for routine use.     

Population structure of arctic lamprey <Emphasis Type="Italic">Lethenteron camtschaticum</Emphasis> from the Kol River (Western Kamchatka)

Phenetic diversity is investigated and the taxonomic status of lampreys (Petromyzontidae) from the Kol basin (Western Kamchatka) is verified. Typically anadromous lamprey, anadromous lamprey forma praecox, resident lamprey, smolts, and larvae of lamprey are discovered and described. Their comparative analysis is made by standard morphometric characters. Differences between the forms by plastic characters depend on size and weight. No significant differences are found by major taxonomic characters between the forms of anadromous and resident lampreys which would indicate their taxonomic separation. Phenetic types of spawners are representatives of the population of one complexly structured species represented by life forms realizing different life history strategies (anadromous and resident), and belong to the species Arctic lamprey Lethenteron camtschaticum (Tilesius, 1811).     

How reliable are morphological and anatomical characters to distinguish European wildcats,domestic cats and their hybrids in France?

Phenotypic variation in hybridizing species or subspecies is a prerequisite for allowing conservation ecologists and wildlife managers to identify parental populations and their hybrids in the field. We assessed the reliability of a set of eight morphological (body size and pelage characters) and four anatomical criteria (skull and intestine morphometric measurements) to distinguish between 302 French specimens classified as wildcat, domestic cat or hybrid on the basis of a Bayesian analysis (STRUCTURE) of their multilocus microsatellite genotypes. This aim was achieved by performing a set of multivariate analyses on morphological, anatomical and genetic data sets (Hill and Smith's analysis, co‐inertia analysis and discriminant analysis of principal components). Wildcats and domestic cats were very satisfactorily distinguished, even when using simple non‐invasive morphological criteria easily usable in the field like the morphology of the tail, dorsal line or flank stripes. Using anatomical instead of morphological characters slightly increased the discriminating power. Many more difficulties arose when we tried to distinguish hybrid specimens from both wildcat and domestic ones. Anatomical characters performed better than morphological ones in recognizing hybrids, but the assignment success rate remained very low, about 31.6% and 1.5%, respectively. Overall, the most discriminating characters were two continuous, derived anatomical characters: the cranial index followed by the intestinal index. Classification of specimens in three classes based on their microsatellite genotypes appeared to be inadequate for identifying hybrid specimens, as hybrid specimens seemed to be distributed along an anatomical continuum. With this observation in mind, we assessed the linear relationships between a proxy of the individual level of hybridization (q_ik) and the cranial and intestinal indices, respectively. Both relationships were highly significant. The greatest correlation was found with the cranial index (R² = 60.4%). Altogether, our results suggest that future work should be geared towards enhancing the measure of hybridization using more discriminating molecular markers and improving morphometric skull measurements through the use of modern geometric morphometric methods, using landmarks rather than skull dimension.     

Integrated genetic and morphological data support eco‐evolutionary divergence of Angolan and South African populations of Diplodus hottentotus

The genus Diplodus presents multiple cases of taxonomic conjecture. Among these the D. cervinus complex was previously described as comprising three subspecies that are now regarded as separate species: Diplodus cervinus, Diplodus hottentotus and Diplodus omanensis. Diplodus hottentotus exhibits a clear break in its distribution around the Benguela Current system, prompting speculation that Angolan and South African populations flanking this area may be isolated and warrant formal taxonomic distinction. This study reports the first integrated genetic [mitochondrial (mt)DNA and nuclear microsatellite] and morphological (morphometric, meristic and colouration) study to assess patterns of divergence between populations in the two regions. High levels of cytonuclear divergence between the populations support a prolonged period of genetic isolation, with the sharing of only one mtDNA haplotype (12 haplotypes were fully sorted between regions) attributed to retention of ancestral polymorphism. Fish from the two regions were significantly differentiated at a number of morphometric (69·5%) and meristic (46%) characters. In addition, Angolan and South African fish exhibited reciprocally diagnostic colouration patterns that were more similar to Mediterranean and Indian Ocean congeners, respectively. Based on the congruent genetic and phenotypic diversity we suggest that the use of hottentotus, whether for full species or subspecies status, should be restricted to South African D. cervinus to reflect their status as a distinct species‐like unit, while the relationship between Angolan and Atlantic–Mediterranean D. cervinus will require further demo‐genetic analysis. This study highlights the utility of integrated genetic and morphological approaches to assess taxonomic diversity within the biogeographically dynamic Benguela Current region.     

Morphological analysis ofAlouatta seniculus species group (Primates,Cebidae). A comparison with biochemical and karyological data

Fifteen cranial measurements were taken from wild caught specimens ofAlouatta seniculus seniculus, A. s. stramineus andA. s. macconnelli. A morphological analysis showed sex dimorphism in these three groups. A multivariate analysis discriminated among these taxa; males being more clearly discriminated than females. Our data showed that these taxa can be separated on the basis of quantitative cranial traits, biogeographic distribution, karyological differences, and biochemical characters. We therefore propose a new taxonomic arrangement, changing their taxonomic status to the species level (Alouatta seniculus, A. macconnelli, andA. stramineus).     

Karyotaxonomic study of some species ofPeucedanum (Umbelliferae)

A study of morphometric karyotype characters was carried out in 14 species ofPeucedanum s. lat. (Umbelliferae — Apioideae) and in one species of the related genusLomatium. The differences of the species in their karyotype characters are correlated only to a limited degree with their similarities of dissimilarities in morphological (i. a. carpological) characters, and thus with their taxonomic position. The evolution of these two sets of characters seems to have proceeded not synchronously or even in different directions in the group of platycarpousUmbelliferae united in the genusPeucedanum. Therefore, it is unlikely that morphometric chromosome characters revealed by monochrome staining can be used appropriately in the taxonomic revision of the polymorphous genusPeucedanum.     

Combining geometric morphometrics with pattern recognition for the investigation of species complexes

The parasitoid Hymenoptera contain a large number of species complexes that are as yet unresolved by traditional taxonomic studies. Evolutionary studies as well as biological control programmes often require further investigations which cannot rely on simple qualitative morphological characters; in many cases, particularly with dried specimens housed in museums, molecular approaches cannot be used. Recent developments in geometric morphometrics and statistical exploratory approaches open new perspectives for the objective evaluation of morphological characters in this taxonomic context. In this study, geometric morphometrics and pattern recognition approaches were applied to the wing shape and venation of two closely related braconid species considered to differ by subtle qualitative morphological head characters. Exploratory analyses such as kernel density estimates and Gaussian mixture analyses were used to explore the structure of the data in the multivariate morphometric space. Discrimination techniques (linear discriminant functions and neural networks combined with cross‐validations) were used to estimate the taxonomic value of qualitative characters. Gaussian mixtures highlighted the existence of two non‐overlapping groups. A good congruence was found between one of the two groups and the a priori defined Bassus tumidulus. The misclassification rate was higher for B. tegularis specimens, which also appeared morphometrically heterogeneous. Discrimination between the two a priori defined species was incomplete with misclassification rates higher than, or equal to, 6%. In most cases, the lack of congruence between species and morphometrically defined subgroups could be related to specimens that exhibited ambiguous qualitative character states. In summary, if two entities are present, they still need to be defined morphologically, while B. tegularis heterogeneity calls for further investigation of specimens of known origin and hosts. © 2003 The Linnean Society of London, Biological Journal of the Linnean Society, 2003, 80, 89–98.     

Morphological and molecular analysis of centropagids from the high Andean plateau (Copepoda: Calanoidea)

In small metazoan invertebrates classical taxonomic analyses can be ambiguous due to the limited number of morphological characters available. This difficulty can yield incorrect estimates of species richness or taxa distribution. The Boeckella genus has been described as the dominant taxon of zooplankton assemblages in the Andean biogeographical region. In this genus, taxonomic classification and delimitation of boundaries between species has long been problematic and controversial. Among South American centropagids Boeckella gracilipes has been regarded as one of the most broadly distributed species, its presence having been reported from Ecuador to Tierra del Fuego. However, in the high Andean plateau some centropagid populations identified as Boeckella gracilipes have also been considered as B. gracilipes titicacae or even identified as a different species, namely Boeckella titicacae. In an attempt to resolve the taxonomic status of the Centropagidae family from the high Andean plateau, we combined traditional and multivariate morphological analyses (integral approach) with the molecular phylogenetic approach. The results obtained allow us to conclude that centropagids collected from the high Andean plateau actually represent a different species, B. titicacae, not B. gracilipes. The phylogenetic reconstruction of the South American Centropagidae family indicated that B. gracilipes represents a sister taxon to B. titicacae. The present study stresses the usefulness of integrating alfa-taxonomy with morphometric and molecular approaches in order to resolve species boundaries, to determine geographical distributions and to investigate evolutionary processes.     

Morphological Differentiation Between Geographically Separated Populations of <Emphasis Type="Italic">Neomysis integer</Emphasis> and <Emphasis Type="Italic">Mesopodopsis slabberi</Emphasis> (Crustacea,Mysida)

Morphological variation was examined in Neomysis integer and Mesopodopsis slabberi, two abundant, low dispersal mysid species (Crustacea, Mysida) along the European coasts. Both species dominate the hyperbenthic communities in the northeast Atlantic, and M. slabberi is also widely distributed in the Mediterranean and Black Sea. Three populations of these species were sampled throughout their distribution range; samples of N. integer were collected in the northeast Atlantic Eems-Dollard, Gironde and Guadalquivir estuaries; in the case of M. slabberi, mysids were sampled in two northeast Atlantic estuaries (Eems-Dollard and Guadalquivir) and one Mediterranean site (Ebro Delta). A total of 12 morphometric and 2 meristic characters were measured from 30–64 mysids per sample. Multivariate analysis showed clear morphometric differences between populations of both species. The morphological differentiation within M. slabberi was highly concordant with the available genetic data from mitochondrial loci, pointing to a large divergence between the Atlantic and Mediterranean populations. However, due to some overlap of individuals between the different populations, the present morphometric analysis does not suffice to assign the different populations to a separate (sub)species status. In the case of N. integer, the morphometric patterns showed a divergence of the Gironde population. Differentiation of populations within this mysid, as in M. slabberi, were mainly related to eye and telson morphology. Potential interactions of the mysid morphology and environmental conditions are discussed.     

Morphological diversity and structure of West Mediterranean Abies species

The aim of this study was the biometrical comparison of Abies pinsapo Boiss., Abies maroccana Trab. and Abies tazaotana Côzar ex Villar from Spain and Morocco. The morphological and anatomical analysis of the needle characters of these firs revealed significant taxonomic differences. The results showed considerable distances between regions and very small differences among the Moroccan fir taxa. The examination of the needles and earlier genetic analysis did not provide evidence for distinguishing A. tazaotana at the species level.     

The limited value of traditional morphometric features in stromatoporoid taxonomy

The morphological variation of stromatoporoids, which are solitary organisms, is partitioned into its presumably genetic and environmental components. Potentially heritable, environmentally mediated and residual components of morphological variability were estimated in a test set containing Devonian stromatoporoids of the genus Gerronostromaria from southern Poland using analysis of variance. The taxonomic importance of traditional morphometric features is limited, because they are dominated by the intra‐skeletal component of variance. Conventional metrics were therefore replaced by stereological and textural quantities. Both stereological and textural features are dominated by the inter‐skeletal and inter‐locality components of variation and thus may be valuable in taxonomic and environmental studies of stromatoporoids. Statistical analyses of these characters (principal component analysis and cluster analysis) were performed. Of 13 characters considered most useful in taxonomic studies, only five have been used previously in conventional species definitions.     

Species-level diversification of African dwarf crocodiles (Genus Osteolaemus): A geographic and phylogenetic perspective

The taxonomy of the African dwarf crocodile (genus Osteolaemus) has been disputed since a novel morphotype was discovered in the early 20th Century. Because this poorly-known reptile is widely hunted throughout the forests of Central and West Africa, resolving the existence and extent of taxonomic units has important management and conservation implications. Lack of molecular data from individuals of known origin and historical disagreement on diagnostic morphological characters have hindered attempts to settle one of the most important taxonomic questions in the Crocodylia. In an effort to clarify the evolutionary relationships among dwarf crocodiles, we sequenced three mitochondrial and two nuclear genes using a large sample of dwarf crocodiles from known localities across major drainage basins of forested Africa. Concordant results from Bayesian, maximum likelihood, maximum parsimony and population aggregation analytical methods support a previously recognized division of the dwarf crocodile into a Congo Basin form (O. osborni) and a West African form (Osteolaemus tetraspis), but also reveal a third diagnosable lineage from West Africa warranting recognition as an separate taxonomic unit. Corrected genetic distances between geographic regions ranged from 0.2% to 0.6% in nuclear fragments and 10.0 to 16.2% in mitochondrial COI. Population aggregation, using fixed and alternate character (nucleotide) states to cluster or divide populations, recovered 232 such molecular characters in 4286 bp of sequence data and unambiguously aggregated populations into their respective geographic clade. Several previously recognized morphological differences coincide with our molecular analysis to distinguish Congo Basin crocodiles from the Ogooué Basin and West Africa. Discrete morphological characters have not yet been documented between the latter two regions, suggesting further work is needed or molecular data may be required to recognize taxonomic divisions in cases where putative species are morphologically cryptic. This study highlights the importance of using widespread taxon sampling and a multiple evidence approach to diagnose species boundaries and reveal cryptic diversity.