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Guillermo D'Elía, Ulyses F. We present sytsematics systematic study of Scapteromys populations from Argentina, Paraguay, and Uruguay, based on molecular and morphological evidence. First, based on DNA sequences base pairs from the mitochondrial cytochrome- b gene, we found that Scapteromys populations fall into 2 main clades; 1 formed by Argentinean and Paraguayan populations together with 1 population from western Uruguay, and the other constituted by the remaining Uruguayan populations.
Analysis of molecular variance showed that Morphological analyses corroborated the existence of 2 main morphotypes among Scapteromys specimens. The phylogeographic break identified is mostly congruent with patterns of morphological and chromosomic variation. In light of these results, we propose that S. A la luz de estos resultados, proponemos que S. The genus of swamp rats, Scapteromysbelongs to Sigmo-dontinae, a New World subfamily of cricetid rodents. Define systematics class 11 is distributed through part of the Río de la Plata basin and some adjacent areas in east-central Argentina, south coastal Brazil, southern Paraguay, and Uruguay.
It occupies habitats near watercourses, including large coass, small creeks, ponds, and swamps. In fact, Scapteromys is define systematics class 11 excellent swimmer; it propels itself by horizontal undulations of the tail and by rowing and paddling with the hind and fore feet, respectively Massoia and Fornes Scapteromys also is able to climb trees, a behavior reported as an adaptation to living in flooded areas Barlow ; Sierra de Soriano Scapteromys is mainly nocturnal and feeds primarily on insects and oligochaetes but also hirudines and vegetation Barlow ; Massoia Phylogenetic analyses of nuclear and mitochondrial DNA sequences indicate that this genus is define systematics class 11 of the tribe Akodontini D'Elía ; Smith and Patton Hershkovitz and Massoia established the basis of the current contents of Scapteromys by removing some species previously allocated to the genus and placing them into new genera, Kunsia and Bibimys.
Massoia and Fornes considered Scapteromys aquaticusa taxon described by Thomasto be a subspecies of S. With the advent of cytogenetic studies of sigmodontine rodents, it was shown that Scapteromys has a large amount of chromosomal variation. Further, this variation appeared to be geographically structured. A diploid number 2n of 32 occurred in populations from Argentina Brum et al.
Myers, define systematics class 11. In parallel, our knowledge of Scapteromys also increased directly from fieldwork, which extended the known distribution of the genus. In addition, Reig described the 1st known fossil species of the genus, S. In this study we present a systematic study of Scapteromys populations from Best free website for affiliate marketing, Paraguay, systemattics Uruguay based on molecular and morphological evidence.
First, we analyze the phylogeographic structure Avise et al. Second, we evaluate the agreement among the uncovered phylogeographic pattern and morphological and karyotypic variation. Third, we consider the taxonomic rank of the phylogeographic units found within Scapteromys. Molecular data analyses. Sequences of 4 specimens were obtained from Smith and Patton and D'Elía Negative xefine were included in all experiments.
In all cases, both heavy and light DNA strands were sequenced and compared. Sequences were visualized, reconciled, and translated to proteins to what is the red dot on match mean for stop codons using Sequence Navigator version 1. Map of a portion of the southern cone of South America showing collecting localities of Scapteromys specimens included in this study.
Numbers refer to sites listed in Appendix I. Dotted line encloses the approximate known distribution of the genus compiled from several sources. See Appendix I for population names and specimens included. Sequence alignment define systematics class 11 done with the program Clustal X Thompson et al. Descriptive analyses assessment of base composition, number of variable characters by codon position, observed number of differences between all haplotype pairs, and differences between sjstematics of populations of the cytochrome- b gene sequences were completed with the program MEGA 2.
The search strategy used consisted of heuristic replicates with tree bisection and reconnection and random addition of taxa. Based on the results of D'Elíasamples of Kunsia and Blarinomys Appendix I were used as what are the causes of disability Nixon and Carpenter to polarize character systdmatics define systematics class 11.
Characters were treated as unordered and equally weighted. Relative support of the recovered clades was assessed by performing 1, jackknife replications Farris clasw al. In addition, Bremer support cpass Bremer were computed for those nodes that originate from branches longer than 1 or 2 steps. Only those changes unambiguously optimized regardless of the kind of character transformation used i.
A hierarchical analysis of the distribution of genetic diversity was conducted in the form of an analysis of molecular variance AMOVA— Excoffier et al. Hierarchical levels were defined on the basis of sampling localities and major clades found in the maximum parsimony analysis. Morphologic data analyses. Locality and sex information were recorded as given on specimen tags and collection catalogs. Skulls were segregated into 5 age classes based on molar wear following Barlow's classification.
Examples of the molar occlusal morphology variation related to wear can be consulted in Massoia ; figure 1. A subset of skulls in general, adult individuals were qualitatively scored for those skull features i. In addition, molar morphology and other cranial characters also were evaluated. Holotypes of S. Measurements were taken according to definitions provided by Myers et al. Descriptive statistics mean, standard deviation, and range were derived for locality samples with 5 or more adult individuals 11 samples.
To increase sample sizes for the multivariate analyses of craniodental dimensions, variables that showed significant intersexual differences were discarded and sexes pooled. A subset of intact skulls without missing measurements was used in multivariate analyses. This sample was constituted only of specimens from age class 3 because 1-way ANOVA revealed significant differences in most of the variables systtematics adult age classes results not shown.
Multiple analysis of variance MANOVA was used to test for significant differences between groups systematisc on the basis of sampling localities and major clades found in the maximum parsimony analysis. Principal components were extracted from a variance-covariance matrix and computed using the craniodental variables after transformation to their natural logarithms. Statistics analyses used the program Statistica StatSoft, Inc.
Molecular based analyses. These haplotypes present define systematics class 11 strong base compositional bias, with a marked deficit of guanine, especially in 3rd codon positions. The 21 haplotypes are defined by 55 variables sites of which 10 correspond to 1st codon positions, 7 to 2nd positions, and 38 to 3rd positions. These 55 nucleotide substitutions implied 11 defije acid differences.
Intrapopulation divergence was low, ranging from 0 to 5 define systematics class 11 substitutions 0—0. Intrapopulation divergence accounts for only 2. Values of divergence between haplotype pairs from different populations show a large range of variation, from 0 to 38 observed substitutions 0. Distribution of the 21 cytochrome- b haplotypes what is sanctioned load and connected load in electricity bill in the phylogeographic analysis of population of Scapteromys.
Population numbers are those of Fig. Haplotype numbers correspond to those listed in Fig. A strict consensus analysis of these trees Fig. One clade the western clade includes all Argentinean and Paraguayan populations together with a population from western Uruguay Las Cañas ; the 2nd the eastern clade has a more restricted distribution what is evolutionary relationship in biology includes the remaining Uruguayan populations Fig.
AMOVA systrmatics that most Observed systematiccs among populations of both clades ranges from 3. Relationships among haplotypes within each of the main clades are poorly resolved. AMOVA indicated that only Observed substitutions among populations within each clade ranges from 0. Haplotypes recovered from Kunsia and Blarinomys specimens were used as outgroups. A Numbers above sysematics below branches correspond to define systematics class 11 jackknife and Bremer support values, respectively, of the nodes at their right.
Numbers at the right of haplotype numbers identify populations as detailed in Fig. Nodes A, B, and C are discussed in the text. B The same consensus tree superimposed onto the distribution map of Scapteromys populations studied Fig. Wider lines indicate boundaries of countries. Shaded areas correspond with the define systematics class 11 depicted in the tree. Morphological analysis. The 1st morphotype is characterized by a frontoparietal suture that more or less resembles an open U or V, and it is often associated with a quadrate mesopterygoid fossa that has a bluntly pointed median palatine process Fig.
This morphotype characterized all Argentinean and Paraguayan populations as well as 1 from Uruguay Las Cañas and is present in the holotype of S. A more or less pronounced frontoparietal suture resembling a W, often associated with a rounded mesopterygoid fossa that lacks a median palatine process, characterized systematkcs other morphotype Fig.
This morphotype is common in all but 1 of the Uruguayan populations, and it is represented by the holotype of S. However, there is some degree of variability in these characters. There are, for example, individuals whose frontoparietal sutures are irregular and cannot be scored as either of the 2 morphotypes.
The correspondence between morphotypes and geography also is obscured by the fact that at most of the how to calculate equivalence ratio clade localities and at 2 southern populations of the western clade, both main morphs occur in sympatry Table 2. In these polymorphic populations, however, 1 of the morphotypes is always predominant.
Type of frontoparietal suture, mesopterygoid fossa, and presence or absence of median palatine process scored in Scapteromys specimens from 68 populations. An asterisk indicates that for a population at least 1 of the individuals morphologically studied also was included in the phylogeographic analysis. Suture types are explained in the text and shown in Fig. See Appendix I for locality details.
Dorsal views of Scapteromys skulls anterior at top, posterior at bottom showing different type of frontoparietal suture above and mesopterygoid fossa systekatics.
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