Statistical approaches to assess intraspecific variations of morphological continuous characters: the case study of the family Phytoseiidae (Acari: Mesostigmata)

Statistical methods for delimiting species with morphological data remain woefully understudied. The present study presents statistical approaches to study the intraspecific variations of continuous characters used in species diagnosis. For this, the data set comprised 14 species of the mite family Phytoseiidae (Acari: Mesostigmata) and 23 continuous characters (seta lengths). The statistical approaches aim to determine (i) how intraspecific variations differ depending on the characters and species considered, and (ii) how to choose the number of specimens to be considered for characterizing intraspecific boundaries. The means of the related confident limits at 95% (rCL₉₅) of the 14 species and the 23 setae are around 20%, suggesting that 95% of the specimens of a same species are included in the interval μ ± 0.2μ. No relationship was found between seta variations and the number of specimens considered for each species, nor for seta variations and their position on the mite body. However, a correlation between seta lengths and rCL₉₅ means was observed. An abacus was proposed to determine the coefficient that must be applied to mean seta length to calculate the lower and upper limits of the interval that will include 95% of the specimens of a same species. The higher the precision around the mean, the higher the number of specimens considered. Furthermore, this number varies considerably depending on the characters and species considered for high precisions. However, for a priori error around the mean of 15%, the number of specimens needed to characterize the intraspecific variation is quite homogeneous and ten specimens would be required. I discuss the utility of such results to resolve some persistent problems in the alpha‐taxonomy of the family Phytoseiidae but also of other taxa. © The Willi Hennig Society 2012.     

New investigation technique for thoracic sclerites of mosquitoes (Diptera, Culicidae) for the correct identification of genera and species

A new technique for studying the external morphology of thoracic sclerites of mosquitoes (Diptera: Culicidae) was developed. According to this method, the shape of sclerites and the position of setae and scales can be examined using SEM or light microscopy even in cases when setae or scales have been lost. The method can be recommended for the damaged material which is often obtained during sampling. The bases of setae usually exceed 100 μm in diameter and thus differ significantly from the bases of scales which are about 30 μm in diameter. Analysis of the structure of sclerites and the arrangement of setae and scales on them in different specimens of one species may reveal intraspecific variation of this character complex. Comparison of species from different genera of the family Culicidae, as well as comparison of closely related species, will probably help to estimate the diagnostic value of thoracic sclerites as morphological characters.     

An integrative morphological and molecular diagnostics for Typhlodromus pyri (Acari: Phytoseiidae)

Tixier, M‐S., Okassa, M. & Kreiter, S. (2011). An integrative morphological and molecular diagnostics for Typhlodromus pyri (Acari: Phytoseiidae). —Zoologica Scripta, 41, 68–78. Phytoseiidae species are predators used in biological control programmes to limit mite pests out‐breaks in the world. Specific diagnostic is thus very important to ensure the success of pest management. This study aims to determine the species status of populations within the main species in vineyards and orchards: Typhlodromus (Typhlodromus) pyri, considering four molecular markers (COI, Cytb mtDNAs, 12SrRNA, ITSS) and 34 morphometrical characters. With the three mitochondrial DNA fragments considered, unexpected high genetic distances were observed between two groups, whereas both nuclear DNA and morphological analyses did not show such distinction. Furthermore, the GYMC and the 4 × rule tests emphasised the existence of a single species. Similarly, cross‐breeding experiments between specimens of the two groups showed no reproductive incompatibility. Therefore, despite the great intraspecific variation emphasised by the three mtDNA markers, we conclude that all the specimens considered belong to the same species. The present study clearly underlines the difficulty to conclude on species status using only mt markers and genetic distances, and stresses the necessity to apply multiple approaches for species definition. It also shows the confidence of the morphology for delineating this species.     

Phylogenetic and taxonomic problems in freshwater Oligochaeta with special emphasis on chitinous structures in Tubificinae

Setae and chitinous penis sheaths are the main characters used to distinguish genera and species of the subfamily Tubificinae. Genital setae and penis sheaths are of functional importance to facilitate copulation. Similar structures in different genera may be homologous or products of parallel evolution (homoiologous). Form and dimensions of the penis sheaths of many tubificine species are very variable. Transspecific overlap of quantitative species characters can make the determination of specimens in the genus Limnodrilus difficult. The configuration of the distal ends of the penis sheaths is an important character to distinguish Limnodrilus species. The definition of the intraspecific variability of this morphological character is problematic.     

Statistical discrimination of sex in Melanoides tuberculata (Gastropoda: Thiaridae)

Systematists may rely on morphometric differences among samples of specimens for the recognition of living and fossil species, even though morphometric differentiation may be caused by non-genetic factors, such as ecophenotypy, differential growth rates and taphonomic mixing. When genetic differences between sexes or among closely related species are expressed as differences in the morphology of the individual or population, potentially valuable information becomes available to the systematist for a variety of genetic and ecological investigations. We have studied the morphology of the freshwater snail Melanoides tuberculata (Muller, 1774) in Israel, where males occur in what would otherwise be normally parthenogenetic (all female) populations. In modern M. tuberculata, sex may be determined by observation of gonadal tissue; in fossil specimens, any classification according to sex must be accomplished using only preservable features of the mineralized shell. Previous research confirmed that in large samples, mean shell shape of male and female snails differed significantly, but the degree of difference was too small to identify the sex of any individual specimen. We apply a three stage process that results with a high degree of accuracy in the discrimination of individual M. tuberculata specimens by sex on the basis of continuous morphological characters: (1) measurement of many aspects of shell morphology of individuals of known sex, and stepwise discrimination to discover which of the variables, if any, contribute to the morphometric differentiation of males from females (one time only, for the species); (2) use of these selected variables in a clustering procedure to make a preliminary assignment of each specimen to sex; (3) use of cluster assignments in a discrimination procedure to optimally predict sex. For species that exhibit morphometric differences between two groups, and for which continuous morphometric variation precludes the a priori recognition of discrete clusters, this sequential procedure may be of broad applicability. These objective methods may be applied to the discrimination within any set of specimens for which the hypothesis of two, and only two, constituent groups may be entertained.     

Morphological and molecular diagnostics of <Emphasis Type="Italic">Phytoseiulus persimilis</Emphasis> and <Emphasis Type="Italic">Phytoseiulus macropilis</Emphasis> (Acari: Phytoseiidae)

This study focuses on the diagnostics of two natural enemy species, belonging to the genus Phytoseiulus in the family Phytoseiidae (sub-family Amblyseiinae): P. macropilis and P. persimilis. These two species are of primary importance in biological control all over the world. However, they are morphologically very similar and specific diagnostics is difficult. This study utilizes mitochondrial molecular markers (12S rRNA and Cytb mtDNA) to differentiate these two species. Morphological analyses showed significant differences between P. persimilis and P. macropilis for 17 morphological characters of the 32 considered. However, despite these significant differences, the ranges of all characters overlap. Only the serration of the macroseta on the basitarsus (StIV) allows the differentiation between P. persimilis and P. macropilis. Despite these small morphological differences, molecular results, for both mitochondrial DNA fragments considered (rRNA and Cytb mtDNA), showed a clear delineation between the specimens of P. macropilis and P. persimilis. This study emphasizes (i) that only one morphological character (serration of the seta StIV) clearly separates these two species, and (ii) the usefulness of an automatical molecular and simple diagnostic tool for accurate differentiation of the two species and ensure the morphological diagnostics. Further studies are proposed, including more DNA sequences especially for P. macropilis.     

Morphological variation versus genetic divergence: a taxonomic implication for Mogannia species (Cicadidae: Cicadinae)

Species showing intraspecific morphological variation tend to be very difficult to identify using morphological characters. One such example is the cicada genus Mogannia where some species show considerable intraspecific variation mainly exhibited by wing pattern and body colouration. Thirty-one variants covering different putative species of Mogannia were recognized and illustrated in the present paper. Molecular data of mitochondrial COI and Cytb sequences were employed to test the level of variation and phylogeny of them. The existence of a ‘barcoding gap’ between intraspecific and interspecific genetic divergences and the reciprocally monophyletic clades indicate that all the closely related variants represent a single species, and that all these variants correspond to six species, respectively. However, the evolutionary relationships of intraspecific variants are not resolved possibly due to insufficient genetic variation among them. Our results indicated that some morphological characters, especially the wing pattern and body colouration, and even the number of apical processes of the aedeagus in a couple of related species, must be used with great caution in delimiting Mogannia species and their relatives. The factors responsible for intraspecific morphological variation and phylogeny of Mogannia spp. are preliminarily discussed.     

Idiosomal setae in the family Phytoseiidae (Acari: Mesostigmata): variation,geographical distribution and taxonomic considerations

Phytoseiidae classification is based on idiosoma chaetotaxy and the assumed evolutionary process is seta suppression. This article aims to determine how depilation could have taken place depending on seta position, subfamily and region. For this, the occurrence of 21 variable setae on dorsal and ventral shields was determined for 1996 species in seven biogeographical regions. The occurrence of eight rare setae assumed to be past relics and 11 rarely absent setae (assumed to be undergoing a loss process) was analysed. The subfamily Phytoseiinae has ‘lost’ the highest number of primitive setae, the subfamily Amblyseiinae has ‘retained’ the highest number and Typhlodrominae has an intermediate position, except for the seta z6. The subfamily Phytoseiinae shows the highest number of setae undergoing a loss process, whereas the subfamily Typhlodrominae has lost these setae in 674 species and subfamily Amblyseiinae in 415 species, making this latter subfamily that with the most retained dorsal setae. According to spatial seta occurrence, it could be hypothesized that Amblyseiinae originated from South Gondwana, Typhlodrominae from the Euro‐America region (Laurasia zone) and Phytoseiinae from the Ethiopian area. The presently admitted classification of Phytoseiidae is discussed with regard to the occurrence of rarely present setae (assumed to have been lost in high frequency). © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 112 , 606–624.     

The comparative morphology of the larynx in snakes

Young, B.A. 2000. The comparative morphology of the larynx in snakes. —Acta Zoologica (Stockholm) 81 : 177–193 The larynx and glottal tube were examined in 10 specimens each of 22 snake species. Qualitative analysis through dissection and clearing and staining revealed distinct morphological variation in the cartilage and intrinsic musculature of the larynx. Quantitative analysis of nine morphometric characters revealed laryngeal sexual dimorphism in three species, significant interspecific differences in the coefficients for every morphometric feature regressed on body size, no significant intraspecific differences in the log transformed means, and poor discrimination of species using principal component analysis. The observed variations in laryngeal morphology did not correlate with the phylogeny of the species, or with habitat preferences.     

Species of the genus Psoroptes (Acari: Psoroptidae): A taxonomic consideration

The biosystematic status of mite species belonging to the genus Psoroptes Gervais, 1841 is difficult to determine by phenotypic methods and has been subject to taxonomic revisions and ongoing debate. At present, the existence of five species, P. cuniculi (Delafond, 1859), P. ovis (Hering, 1838), P. equi (Hering, 1838), P. cervinus Ward, 1915 and P. natalensis Hirst, 1919, is generally accepted. This classification is based mainly on the host species, the localization of the mites on their hosts and morphological characters of male mites. However, a critical review of the literature indicates that the features used to discriminate between the five species are not unequivocal: (a) the localization of mite populations on host animals is not completely strict, (b) the lengths of the outer opisthosomal setae of male mites, which are the main morphological features used for species discrimination, overlap between the five postulated species, and (c) host specificity cannot be deduced from results of transfer experiments. Rather, conspecificity of the members of the genus Psoroptes has to be presumed which is supported by molecular genetic analyses. On these grounds and on rules of priority P. cervinus Ward, 1915, P. cuniculi (Delafond, 1859), P. natalensis Hirst, 1919 and P. ovis (Hering, 1838) are seen as synonyms of P. equi (Hering, 1838).     

Ecological wood anatomy of NepaleseRhododendron (Ericaceae). 2. Intraspecific variation

The intraspecific relationship of selected wood anatomical characters with stem diameter, plant height, and altitude was investigated in four NepaleseRhododendron species:R. anthopogon, R. lepidotum, R. campanulatum, andR. arboreum, i.e., two shrubs, one subtree, and one tree. We studied 23 to 27 specimens for each species.R. anthopogon grew from 3,380 to 4,950 m,R. lepidotum from 2,060 to 4,720 m,R. campanulatum from 2,790 to 4,140 m, andR. arboreum from 1,430 to 3,460 m. Multiple regression analysis and actual distribution of character values show that the wood anatomical characters having a significant correlation with non-anatomical factors differ between species. The number of significant characters are two inR. anthopogon andR. campanulatum, five inR. lepidotum, and nine inR. arboreum and tends to be small in species having smaller altitudinal ranges. Average pore area, most strongly correlated with non-anatomical factors in interspecific variation, is significantly correlated only inR. lepidotum andR. arboreum. The general trends in intraspecific variation among four NepaleseRhododendron species differ from the trends found in interspecific variation within the genus and are characteristic of each species.     

New conditions and intraspecific variation of some muscles of hands and feet of Dendropsophus labialis (Peters, 1863) (Anura,Hylidae)

Systematics of frogs have been based on osteological and molecular characters; however, the morphology of the muscles of hands and feet has proven to be an important complement to these studies, but it has not been sufficiently studied. This study presents undescribed conditions based on the origin, insertion or arrangement of 18 muscles of hands and feet of Dendropsophus labialis, and intraspecific variation. Muscles of four specimens (two males and two females) were examined to observe both sides of each specimen determining new conditions, and comparing them with documented species of Dendropsophus. Four intraspecific variant categories were established: Minor, unique, explosive and mimicking. Mainly mimicking, explosive and unique variations were found. Our results show the need to expand intraspecific studies in more species of Dendropsophus and to assess its value in the phylogeny of the genus. The family Hylidae may be identifiable based upon the combination of character states of some of these muscles: the m. flexor indicis superficialis proprius and the group of tendines superficialis digitorum of hands and foot, the m. extensor brevis superficialis Digiti III of the hand, the m. extensor brevis superficialis Digiti IV in hands and the mm. extensores breves profundi Digiti III in hands.     

维螨属2新种记述(蜱螨亚纲:中气门目:维螨科)

记述维螨属2新种:河南维螨Veigaia henanensis sp.nov.和福建维螨Veigaia fujianensis sp.nov.     

Geographical variation in the species Montivagum dihumerale and speciation in chigger mites (Acari: Trombiculidae)

Intraspecific morphological variation of the chigger mite species Montivagum dihumerale (Traub et Nadchatram, 1967) is studied. Eco-geographic rules of the variation are revealed. General size of mites is found to be increased along with the rise of the high-mountain character of the landscape in the collection localities. The numbers of idiosomal setae are varied independently from the size parameters and geographically close populations are proved to be the most similar by these characters. At the same time, numbers of the setae of different types play the leading role in the discrimination of closely related Montivagum species, while the eco-geographical rules have not been found in this genus at the level of interspecific differences. As a result, the hypothesis is set up, that a significant degree of isolation of local populations separated from each other by high mountain ranges of the Central Asia is the main factor of speciation in the genus Montivagum. Regional character of this speciation mode is confirmed by the comparison with other chigger mites taxa.     

Kallymenia crouaniorum (Kallymeniaceae,Rhodophyta), a new red algal species from the Laminaria hyperborea understorey community

In this paper we describe Kallymenia crouaniorum Vergés & Le Gall, sp. nov. (Kallymeniaceae), a new marine red alga from the north-eastern Atlantic Ocean. rbcL and LSU sequences of this species, previously misidentified in the field as Kallymenia reniformis, diverged from those of other Kallymenia species by at least 7.5% and 5.2%, respectively. Kallymenia crouaniorum also has a set of distinctive vegetative and reproductive characteristics, including a deeply lacerate frond, a short stipe, dentate margins, large cortical cells up to 110 μm in diameter, highly refractive stellate medullary cells with arms up to 1000 μm in length, and a monocarpogonial branch system. Molecular phylogenies inferred from rbcL and LSU data indicated, albeit with weak support, that this new species is a sister taxon of a lineage encompassing the generitype K. reniformis, as well as most species of Kallymenia included in the phylogenetic analysis. The main morphological characters that delineate monocarpogonial Kallymenia species are presented. Moreover, after reviewing the literature and several herbarium specimens, we found in the Weber-van Bosse Herbarium a specimen collected at Roscoff in August 1894 with the anatomical characters of the species described here, confirming that this newly described species has in fact been overlooked and is not a recent introduction.     

Geographical variation in the high-duty cycle echolocation of the cryptic common mustached bat Pteronotus cf. rubiginosus (Mormoopidae)

The use of bioacoustics as a tool for bat research is rapidly increasing worldwide. There is substantial evidence that environmental factors such as weather conditions or habitat structure can affect echolocation call structure in bats and thus compromise proper species identification. However, intraspecific differences in echolocation due to geographical variation are poorly understood, which poses a number of issues in terms of method standardization. We examined acoustic data for Pteronotus cf. rubiginosus from the Central Amazon and the Guiana Shield. We provide the first evidence of intraspecific geographic variation in bat echolocation in the Neotropics, with calls significantly differing in almost all standard acoustic parameters for the two lineages of this clade. We complement our bioacoustic data with molecular and morphological data for both species. Considerable overlap in trait values prevents reliable discrimination between the two sympatric Pteronotus based on morphological characters. On the other hand, significant divergence in the frequency of maximum energy suggests that bioacoustics can be used to readily separate both taxa despite extensive intraspecific variability in their echolocation across the Amazon. Given the relative lack of barriers preventing contact between bat populations from the Central Amazon and French Guiana, the documented acoustic variation needs to be further studied in geographically intermediate locations to understand the potential isolation processes that could be causing the described divergence in echolocation and to determine whether this variation is either discrete or continuous.     

我国华鳊属鱼类形态差异及其物种有效性的研究

为探讨华鳊属鱼类种间和种内居群间的形态差异及其物种有效性,采用多变量形态度量,结合传统分类方法,获得该属鱼类不同地理居群共224尾标本的32个测量性状和7个可数性状数据,并做主成分分析。在可数性状上,海南华鳊各居群间无差异,但明显不同于其他种类;四川华鳊与大眼华鳊差别明显,与伍氏华鳊之间也存在较大程度的差异;大眼华鳊的乌江和珠江居群之间存在显著差异,而伍氏华鳊各地理居群差异不明显。在可量性状上,四川华鳊和大眼华鳊的乌江居群都明显区别于其他种或居群,而大眼华鳊的珠江居群与伍氏华鳊之间以及海南华鳊各居群之间没有明显差异。对上述结果分析认为：四川华鳊和海南华鳊为两个有效的物种;大眼华鳊的乌江标本是一个未被描述的物种,而珠江居群和伍氏华鳊各居群应属同一个种。     

How useful are the recommended counts and indices in the systematics of the Octopodidae (Mollusca: Cephalopoda)

A morphological dataset based on 14 standard counts and indices was constructed for 68 specimens comprising 12 species of octopuses. This was used to construct distance matrices based on morphological characters. These matrices were compared with genetic distance matrices compiled during molecular phylogenetic analyses of the same 12 species using four mitochondrial and two nuclear genes. Mantel tests showed that there was significant congruence between the phenetic and genetic matrices, suggesting that the genetic signal is reflected in the morphological data set. Matrices of geographical distance were constructed for the 12 species based on the latitude, longitude, and depth of capture of 1726 individuals. These matrices never showed significant congruence with genetic data or with morphological data. Multivariate analysis of the morphological dataset suggests that these counts and indices, traditionally used for discriminating between species in cephalopods, do not show great discrimination at species level, but provide excellent discrimination at the generic level, and, as such, might be useful for resolving the generic placement of some problematic taxa. © 2008 The Linnean Society of London, Biological Journal of the Linnean Society, 2008, 95 , 205–218.