Classification of the flea families (Siphonaptera): II. Family hystrichopsyllidae (Part 4)

Distribution of the ancestral, homoiologous, and convergent character states over the tagmata and morpho-functional complexes of the flea body in nine subfamilies of the Hystrichopsyllidae is analyzed. The analysis is based upon the states of 111 characters of 73 structures of 3 tagmata in the adult flea body. The homoiologous states (those arising in parallel in close flea infraorders) comprise 35%, the ancestral (the least specialized) ones, 16%, and the convergent ones, 13% of the total number of character states of all the 10 types. The greatest numbers of the ancestral character states in the Hystrichopsyllidae are found in the mesothorax (22.8%) and aedeagus (21.1%); the metathorax reveals 19.3%, and the head, 17.5% of the ancestral character states. The greatest number of homoiologous character states, 34.1% of the total number, is observed in the mesothoracal segment. Smaller fractions of the homoiologous character states are found in the head (24.6%) and metathorax (17.5%). Those character states which arise in the phylogenetically remote infraorders are considered convergent. The greatest number of convergent character states in the Hystrichopsyllidae (55.1%) is observed in the structure of the aedeagus.     

Morphological diversity of the skeletal structures of fleas (Siphonaptera). Part 2: The general characteristic and features of the thorax

Analysis of a considerable amount of data on the anatomy of 27 structures of the thorax in 96 genera of fleas (over 90% of the genera in the world fauna) shows that different flea taxa can be described based on 48 universal and specific characters, whose 203 states reflect the entire known diversity of the flea morphology. Of them, 26 characters with 96 states can be formulated based on universal terms; 22 universal characters with 88 states describe the proportions of the segments and their parts, and also proportions and shapes of their sclerites; 23 specific characters with 107 states describe the structure of the thoracic sclerites and ridges as well as the patterns of their junctions. Judging by the number of characters and their states, the most evolutionarily flexible structures in fleas are the mesosternite and its apodeme together with the upper part of the mesopleural rod. Homoplasies at various levels comprise less than half (40%) of all the character states in the thorax structure. The character states reflecting the phylogenetic closeness of taxa make up about 15%.     

Morphological diversity of the skeletal structures of fleas (Siphonaptera). Part 4: The general characteristic and features of the abdomen

Characters of 25 abdominal structures were analyzed in the fleas of 96 genera representing over 90% of the world fauna. It was shown that different flea taxa could be described based on 16 universal and 12 specific characters, whose 108 states reflect the entire known diversity of the flea abdominal morphology. Of them, 16 characters with 39 states are formulated based on universal terms. Five universal characters with 13 states describe the proportions of various structures, and also the proportions and shapes of their sclerites; 17 specific characters with 69 states describe the structure of the skeletal elements and the patterns of their junctions. Judging by the number of characters (14) and their states (69), the most evolutionarily flexible structures in fleas are the inner sclerites and anchoring structures of the aedeagus, sternite IX in males, and also the spermatheca and tergite I in females. The character states reflecting the possible phylogenetic closeness of taxa comprise 39%, while 61% of the abdominal character states are homoplasies.     

Complex clines: the predictivity of complicated patterns of geographic variation portrayed by multivariate analysis

The stability of complex patterns of geographic variation was investigated by assessing the congruence between multivariate ordinations derived from randomly chosen real characters. Two series of populations were analysed representing two situations with complex patterns of geographic variation. The first, a ‘Eurasian’ series of populations, showed a strongly structured hierarchical pattern, the second, an ‘eastern’ series of populations, showed a more subtle complex pattern of smooth clines and steps. The characters were selected from a total of 81 (Eurasian) or 61 (eastern) within-population independent characters from six different systems. The congruence between ordinations of the geographical populations was measured by the rotational fit statistic, R². Three procedures were used to compare ordinations based on from two to up to 80 characters randomly chosen to give: A, completely independent character sets; B, subsets compared to the total set; and C, potentially overlapping sets. All three procedures showed that congruence between the ordinations was asymptotic in relation to character number. This relationship was described by one of two mathematical models (procedure B did not result in a hyperbolic model as found with simple patterns of geographic variation). Generally speaking, once a sufficient number of characters are used, the complex patterns of geographic variation are stable, reliable and predictive and not substantially influenced by character choice. The strongly structured hierarchical pattern required 15 or so characters to achieve reliability whilst the more subtle patterns required 20 or so characters. However, the addition of further characters does improve reliability in both cases. The greater percentage of variance portrayed by three-dimensional ordinations compared to two-dimensional ordinations is achieved at the cost of lower congruence when a sufficient number of characters are used. If case studies of geographic variation were to adopt these procedures (preferably using completely independent character sets; procedure A) the reliability of their results would be indicated).     

The distribution and systematic relevance of the androecial character oligomery

Localization of the stamens can be approached by a preliminary distinction between two characters, oligomery and polymery, occurring in two different groups of taxa, respectively the oligomerous complex and the polymerous complex. Oligomery is described by four character states standing in a close semophyletic relationship: diplostemony, obdiplostemony, haplostemony and obhaplostemony. Each character state is analysed for its distribution and systematic value. Diplostemony is the synapomorphic character state for the oligomerous line and has arisen once from a polymerous ancestor or in parallel in different lines. Obdiplostemony arises ontogenetically in three different ways. Loss of one whorl leads either to obhaplostemony, or haplostemony; both character states are believed to represent evolutionary steps of no-return. Secondary increases and reductions of the stamens within a whorl are seen as expressions of the intrinsic variability of the character states and should not be homologized with them. Stamen numbers can be increased by the building-up of complex primordia or by secondary receptacular growth. Reductions of stamens affect one or two whorls of stamens and are caused by lack of space, interactions with the gynoecium and zygomorphy. The distribution of the different character states of oligomery is presented on Dahlgrenograms and the androecia of a number of families and their relationships are discussed. The interactions between oligomery and polymery are analysed as guidelines for a global phylogeny of the Magnoliatae.     

How can third codon positions outperform first and second codon positions in phylogenetic inference? An empirical example from the seed plants

Greater phylogenetic signal is often found in parsimony-based analyses of third codon positions of protein-coding genes relative to their corresponding first and second codon positions, even for early-derived ("basal") clades. We used the Soltis et al. (2000; Bot. J. Linn. Soc. 133:381-461) data matrix of atpB and rbcL from 567 seed plants to quantify how each of six factors (observed character-state space, frequencies of observed character states, substitution probabilities among nucleotides, rate heterogeneity among sites, overall rate of evolution, and number of parsimony-informative characters) contributed to this phenomenon. Each of these six factors was estimated from the original data matrix for parsimony-informative third codon positions considered separately from first and second codon positions combined. One of the most parsimonious trees found was used as the constraint topology; branch lengths were estimated using likelihood-based distances, and characters were simulated on this tree. Differential frequencies of observed character states were found to be the most limiting of the factors simulated for all three codon positions. Differential frequencies of observed character states and differential substitution probabilities among states were relatively advantageous for first and second codon positions. In contrast, differential numbers of observed character states, differential rate heterogeneity among sites, the greater number of parsimony-informative characters, and the higher overall rate of evolution were relatively advantageous for third codon positions. The amount of possible synapomorphy was predictive of the overall success of resolution.     

Considerations on the evolution of qualitative multistate traits

Simple models for the evolution of qualitative multistate traits are considered, in which the traits are permitted to evolve in time-dependent versus speciation-dependent fashion. Of particular interest are the means and variances of distances for these traits in evolutionary phylads characterized by different rates of speciation, when alternative characters are neutral with respect to fitness, and when the total number of observable characters is limited to small values. As attainable character states are increasingly restricted, mean distance (D) in a phylad decreases, regardless of whether evolution is a function of time or of rate of speciation. The ratio of mean distances in species-rich and species-poor phylads of comparable evolutionary age (D_R/D_P) remains near one when differentiation is proportional to time, even when attainable character states are severely restricted. D_R/D_P also nears one as a result of restricting character states when differentiation is proportional to rate of speciation, but the effect is not severe unless the number of character states is very small and the probability of change per speciation very large. These and other results are discussed with reference to available data sets on qualitative multistate traits.     

Proteroctopus ribeti in coleoid evolution

Palaeontological data are key elements for inferring ancestral character states and the assembly of character complexes, but cephalopod fossils preserving soft tissues are very rare. The exceptionally well‐preserved, unique specimen of Jurassic Proteroctopus ribeti Fischer & Riou from the Lagerstätte of La‐Voulte‐sur‐Rhône (c. 165 Ma, France) is one of the few fossil octopod related taxa, but is rarely considered in evolutionary studies. In this paper, we used synchrotron microtomography to reappraise its external characters and for the first time, to reveal its internal structures. A unique character association is found with two fins, head fused to the body, eight well‐developed arms with cirri and two rows of oblique suckers, a gladius and absence of an ink sac. The phylogenetic analysis indicates that Proteroctopus is a basal member of the Vampyropoda. However, this result should be interpreted with caution due to the number of unknown character states in the matrix. Contrary to previous assumptions, the phylogenetic position of Proteroctopus, as well as its stratigraphic occurrence, suggest that the arrangement of biserial suckers may be the ancestral condition in Vampyropoda.     

The distribution and the systematic relevance of the androecial characters oligomery and polymery in the Magnoliophytina

In the system of classification sensu Cronquist, the division between the two subclasses Dilleniidae and Rosidae relies on only a few characters. The major distinguishing character is the occurrence of centripetal or centrifugal sequence of stamen development. We question not only the application and the value of this character, but also the maintenance of the Rosidae and Dilleniidae as separately derived natural taxa, because of the lack of solid distinguishing criteria. We have studied the staminal characters in the 192 families of the Dilleniidae and the Rosidae, using Dahlgren's classification diagram.
The study focussed on the following androecial characters: 1) polyandry; i.e. spiral polyandry, complex polyandry (viz. centrifugal, centripetal and lateral polyandry) and cyclic polyandry 2) diplostemony and obdiplostemony 3) haplostemony and obhaplostemony.
In our proposal only two characters are maintained, namely oligomery (diplostemony s.l.) and polymery (true polyandry s.l.), both having a number of character states. Polyandry, as it has been described up to now, is not to be regarded as homologous in all Magnoliophytina. Both characters correspond to two different lines within the Magnoliatae. In addition, the division between the Dilleniidae and Rosidae sensu Cronquist is no longer tenable.
Finally, a number of modifications of Dahlgren's diagram are suggested and a division of the Magnoliophytina in a polymerous group and an oligomerous group is proposed. The Dilleniidae-Rosidae complex is included in the oligomerous group.     

Phylogenetically structured variance in felid bite force: the role of phylogeny in the evolution of biting performance

A key question in evolution is the degree to which morphofunctional complexes are constrained by phylogeny. We investigated the role of phylogeny in the evolution of biting performance, quantified as bite forces, using phylogenetic eigenvector regression. Results indicate that there are strong phylogenetic signals in both absolute and size‐adjusted bite forces, although it is weaker in the latter. This indicates that elimination of size influences reduces the level of phylogenetic inertia and that the majority of the phylogenetic constraint is a result of size. Tracing the evolution of bite force through phylogeny by character optimization also supports this notion, in that relative bite force is randomly distributed across phylogeny whereas absolute bite force diverges according to clade. The nonphylogenetically structured variance in bite force could not be sufficiently explained by species‐unique morphology or by ecology. This study demonstrates the difficulties in identifying causes of nonphylogenetically structured variance in morphofunctional character complexes.     

Ultrastructural observations on the oogenesis of Triops cancriformis (Crustacea,Notostraca)

Summary The first stages of the oogenesis of Triops cancriformis have been studied. At the outset the oocyte is smaller than the nurse cells. Meiosis begins with typical synaptonemal complexes. The significance of these complexes and of some other peculiar structures of germ cells, i.e., pore complexes and annuli within the nucleus, and annulate lamellae within the cytoplasm are discussed. The morphofunctional uniformity of some cytoplasmic structures (annulate lamellae, concentrically arranged ER, and yolk globules) in the oocyte as well as its nurse cells is also discussed.     

Geometric and traditional morphometrics for the assessment of character state identity: multivariate statistical analyses of character variation in the genus Arrenurus (Acari,Hydrachnidia, Arrenuridae)

Geometric and traditional morphometric approaches are tested to describe and reveal taxonomic characters and character states in variation of shape and size of idiosoma morphology in the Arrenuridae. Patterns of variation of idiosoma and glandularia features of males from 11 Mexican species of Arrenurus (Megaluracarus) and two species of subgenus Dadayella were explored with five landmark configurations and three sets of interlandmark distances. Separate principal component analyses (PCA) and canonical variate analyses (CVA) were performed for each data set. The eight multivariate analyses of variance among 13 a priori groups (species) detected significantly different morphometric variants, which were interpreted as different taxonomic character states. Patterns of character state similarity among species were examined with unweighted‐pair grouping method using averages (UPGMA) cluster analyses on Mahalanobis distances. Analyses of five landmark configurations revealed important taxonomical variation in the anterior idiosoma outline (10 character states), the outline of the posterior region or cauda (13 states), the distribution of postocularia, and the second and third pairs of dorsoglandularia (nine states), the fourth pair of dorsoglandularia (three states), and ventroglandularia on the posterior side of idiosoma (nine character states). Multivariate analyses of three sets of distance measurements also resulted in the detection of potential taxonomic characters related to idiosoma size (12 character states), postocularia and dorsoglandularia (13 states), and ventroglandularia (nine character states). Morphometric analyses of distances and shapes provide a formal basis for the interpretation of taxonomic characters, and for the discovery of character states. These characters should be investigated further in a wider sample of species for the phylogenetic systematics of these water mites. In the meantime, idiosoma regions and structures were tested for congruence in a phylogenetic analysis, and were proposed as homologous among the species sampled.     

Can poorly understood new characters support a poorly understood phylogeny? Shell-structure data in Hydrobiid systematics (Mollusca: Gastropoda: Prosobranchia: Hydrobiidae)

Parsimony-based techniques may help to reconstruct phylogeny based on knowledge of character evolution, and vice versa. Problems arise when one deals with poorly known and poorly understood phylogenies and character behaviour, as in such cases it is not easy to avoid circular logic. This study is concerned with such a case: the poorly understood systematics of the family Hydrobiidae; and with a new character set: shell surface and inner structure as studied by SEM. This is based on previous papers by these authors, describing these structures. A total of 25 binary or multistate characters were chosen, all of unknown transformation series and unclear homologies. The shortest tree was found with the HENNIG86 PC program, and further analysis was carried out with MACCLADE. The evolution of all the characters was analysed. Seven characters were excluded as being uninformative and/or having unknown states in some taxa. MPR was rooted based on the outgroup criterion, resulting in a tree that was two steps longer. The latter was compared with four trees found in the literature. The SEM-based tree supports only the tree based on the classification by Ponder and Warén (1988) and does not support the trees constructed following the systems described by Bernasconi (1992) or Radoman (1983). The shell-structure characters seem to show the same mosaic pattern of evolution as shown by the traditionally used characters, extending the character list. It seems possible to support the ‘traditional’ data set with the ‘new’ one using successive approximations, thus avoiding circular logic.     

Fifty years of character compatibility concepts at work

In the mid 19th century,systematic biologists realized that observable similarities and differences among a group of related species could be the basis for hypotheses about the evolutionary relationships among the species and their ancestors.Such hypotheses Can be expressed as characters.A character is comprised of two or more character states of species considered to be similar with respect to a basis for comparison.The states of a character may also be arranged into a character state tree to hypothesize speciation events associated with changes from one character state to another.In the mid 20th century.some systematists realized that sometimes paxrs of characters(or character state trees)could be incompatible as hypotheses,i.e.,they could not both be true.Through the 1950s,'60s and'70s,tests for,and ways to resolve,incompatibilities were used to estimate an ancestor relation based on mutually compatible characters.An estimate was often shown as a diagram connecting ancestors to their immediate descendants(not quite correctly)called a phylogenetic tree.More recently,other applications of compatibility concepts have been developed,including:identify characters that appear to be random in the context of their data set;combine estimates of ancestor relations for subsets of taxa in a larger collection into a single estimate(a so-called supertree)for the whole collection;and interpret geographic patterns in an evolutionary context.     

Identification of novel mitochondrial protein components of Chlamydomonas reinhardtii. A proteomic approach

Pure mitochondria of the photosynthetic alga Chlamydomonas reinhardtii were analyzed using blue native-polyacrylamide gel electrophoresis (BN-PAGE). The major oxidative phosphorylation complexes were resolved: F(1)F(0)-ATP synthase, NADH-ubiquinone oxidoreductase, ubiquinol-cytochrome c reductase, and cytochrome c oxidase. The oligomeric states of these complexes were determined. The F(1)F(0)-ATP synthase runs exclusively as a dimer, in contrast to the C. reinhardtii chloroplast enzyme, which is present as a monomer and subcomplexes. The sequence of a 60-kD protein, associated with the mitochondrial ATP synthase and with no known counterpart in any other organism, is reported. This protein may be related to the strong dimeric character of the algal F(1)F(0)-ATP synthase. The oxidative phosphorylation complexes resolved by BN-PAGE were separated into their subunits by second dimension sodium dodecyl sulfate-PAGE. A number of polypeptides were identified mainly on the basis of their N-terminal sequence. Core I and II subunits of complex III were characterized, and their proteolytic activities were predicted. Also, the heterodimeric nature of COXIIA and COXIIB subunits in cytochrome c oxidase was demonstrated. Other mitochondrial proteins like the chaperone HSP60, the alternative oxidase, the aconitase, and the ADP/ATP carrier were identified. BN-PAGE was also used to approach the analysis of the major chloroplast protein complexes of C. reinhardtii.     

The status of ovary capillaries in fetuses of White rats after irradiation in various periods of its embryogenesis

The peculiarities of reactions of blood capillaries and their cell structures in the ovary of 20-days old foetus to the irradiation in embryogenesis were revealed; it was shown for the first time that the character and direction of changes in principal morphofunctional parameters depended on the dosage, power of gamma-irradiation and the age of the developing organism by the exposure moment.     

THE EFFECT OF ORDERED CHARACTERS ON PHYLOGENETIC RECONSTRUCTION

Abstract Morphological structures are likely to undergo more than a single change during the course of evolution. As a result, multistate characters are common in systematic studies and must be dealt with. Particularly interesting is the question of whether or not multistate characters should be treated as ordered (additive) or unordered (non-additive). In accepting a particular hypothesis of order, numerous others are necessarily rejected. We review some of the criteria often used to order character states and the underlying assumptions inherent in these criteria.
The effects that ordered multistate characters can have on phylogenetic reconstruction are examined using 27 data sets. It has been suggested that hypotheses of character state order are more informative then hypotheses of unorder and may restrict the number of equally parsimonious trees as well as increase tree resolution. Our results indicate that ordered characters can produce more, equal or less equally parsimonious trees and can increase, decrease or have no effect on tree resolution. The effect on tree resolution can be a simple gain in resolution or a dramatic change in sister-taxa relationships. In cases where several outgroups are included in the data matrix, hypotheses of order can change character polarities by altering outgroup topology. Ordered characters result in a different topology from unordered characters only when the hierarchy of the cladogram disagrees with the investigator's a priori hypothesis of order. If the best criterion for assessing character evolution is congruence with other characters, the practice of ordering multistate characters is inappropriate.