Chromosome plasticity in Ctenomys (Rodentia Octodontidae): chromosome 1 evolution and heterochromatin variation

A chromosome 1 (Cr1) pericentric inversion is described in six of seven species in the genus Ctenomys (tuco-tucos) from Uruguay. The inversion was inferred from G-band analyses of subtelocentric Cr1 hypothesised to be derived from the ancestral metacentric condition. Cr1 varies across species in heterochromatin amount and localisation including a metacentric chromosome without positive C-bands in C. torquatus, a subtelocentric chromosome with heterochromatic short arms in C. rionegrensis, and a subtelocentric chromosome negative after C-banding in five of the species analysed here. Pachytene chromosomes from C. rionegrensis, a species with the highest heterochromatin content, and C. torquatus, one of the species with the lowest heterochromatin content, were analysed in order to assess possible mechanisms of heterochromatin evolution. This analysis revealed the presence of three heterochromatic chromocenters in C. rionegrensis where bivalents converge, while in C. torquatus only one chromocenter was observed. In both species, highly repetitive DNA was observed, localised in chromocenters after “in situ” hybridisation. Heterochromatin associated protein M31 was localised in chromocenters of both species after immuno-detection. The spread of heterochromatin in Ctenomys chromosomes could be produced by chromatin exchanges at the chromocenter level. We propose the exchange of this DNA associated proteins between non-homologous chromosomes in pachytene to be the responsible for the spread of heterochromatin through the karyotypes of species like C. rionegrensis     

Chromosomal polymorphism and small karyotypic differentiation in a group of Ctenomys species from Central Argentina (Rodentia: Octodontidae)

Cytogenetic analysis was performed in six nominal taxa of the genus Ctenomys with a diploid number of 2n=47/48. The studied species were: C. australis (2n=48, FN=76); C. mendocinus (2n=47/48, FN=68/75/76); C. porteousi (2n=47/48, FN=71/72/73); C. azarae (2n=47, FN=71); C. sp. (chasiquensis) (2n=47/48); and C. talarum (2n=48, FN=80). The first three species shared the whole complement, C. talarum shares with them 19 arms from a total of 43 (44%). In all species analyzed constitutive heterochromatin was detected in most short arms, and in several centromeres. Polymorphisms for several pairs involving the heterochromatic short arms together with a complex polymorphism of pair A1 were found in C. azarae, C. sp., C. mendocinus and C. porteousi. Intraindividual variation found in one specimen of C. porteousi involving heterochromatic arms is discussed.These results lead us to propose the inclusion of all species except C. talarum, within a complex called the mendocinus-group. A new case of conservatism for chromosomal number in the genus Ctenomys is found in this group.     

Polymorphism of canonical and noncanonical <Emphasis Type="Italic">gypsy</Emphasis> sequences in different species of <Emphasis Type="Italic">Drosophila </Emphasis><Emphasis Type="Italic">melanogaster</Emphasis> subgroup: possible evolutionary relations

Mobile genetic elements constitute a substantial part of eukaryotic genome and play an important role in its organization and functioning. Co-evolution of retrotransposons and their hosts resulted in the establishment of control systems employing mechanisms of RNA interference that seem to be impossible to evade. However, “active” copies of endogenous retrovirus gypsy escape cellular control in some cases, while its evolutionary elder “inactive” variants do not. To clarify the evolutionary relationship between “active” and “inactive” gypsy we combined two approaches: the analysis of gypsy sequences, isolated from G32 Drosophila melanogaster strain and from different Drosophila species of the melanogaster subgroup, as well as the study of databases, available on the Internet. No signs of “intermediate” (between “active” and “inactive”) gypsy form were found in GenBank, and four full-size G32 gypsy copies demonstrated a convergence that presumably involves gene conversion. No “active” gypsy were revealed among PCR generated gypsy ORF3 sequences from the various Drosophila species indicating that “active” gypsy appeared in some population of D. melanogaster and then started to spread out. Analysis of sequences flanking gypsy variants in G32 revealed their predominantly heterochromatic location. Discrepancy between the structure of actual gypsy sites in G32 and corresponding sequences in database might indicate significant inter-strain heterochromatin diversity. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Dolichopteryx rostrata, a new species of spookfish (Argentinoidea: Opisthoproctidae) from the eastern North Atlantic Ocean

A new species, Dolichopteryx rostrata, is described on the basis of a single specimen (66.2 mm in standard length) collected west of the Hebrides Islands, eastern North Atlantic Ocean. The new species is characterized by an elongate snout and head, small pouchlike eyes, an adipose fin, short dorsal fin base, anal fin base originating under dorsal fin base, a clear longitudinal suborbital brownish band extending forward from behind posterior margin of orbit to snout tip, and 41 (=26 + 15) vertebrae. Total fecundity is low; the ovarian eggs number only 473, despite the ovary having developed ova. Ovarian eggs could be clearly subdivided into an “undeveloped group” (0.1–0.7 mm diameter classes, n = 405) and a “developed group” (0.9–1.3 mm classes, n = 68), based on their frequency distribution. Supplementary material to this paper is available in electronic format at http://dx.doi.org/10.1007/s10228-005-0306-2     

Integrative analysis of chromosome banding,telomere localization and molecular genetics in the highly variable <Emphasis Type="Italic">Ctenomys</Emphasis> of the Corrientes group (Rodentia; Ctenomyidae)

The genus Ctenomys comprises about 70 species with great chromosome diversity. The Corrientes group is one of the most chromosomally variable lineages in the genus, where the diploid number (2n) varies from 41 to 70. In this group, three nominal species and numerous polymorphic and polytypic populations have been described. In order to get insight into the chromosomal evolution of this species complex, we applied different banding and molecular cytogenetic techniques. The results were interpreted in an evolutionary context, based on mitochondrial cytochrome b analyses. Studied samples are representative of the broad chromosomal variability in the group, including specimens with 2n?=?42 to 2n?=?70. Heterochromatin was scarce but concentrated in a few chromosomes. Centromeric DAPI-negative heterochromatin was observed in some autosomal pairs, which differed among populations. Location and amount of DAPI-neutral heterochromatin within the Y chromosome varied among populations. The variable distribution of heterochromatin indicates its dynamic behavior. NORs were detected in one pair of autosomes, which also differed among some populations. Telomeric FISH signals were observed in all complements only at the chromosome ends. The Corrientes group belongs to a clade that also includes C. pearsoni, C. lami, C. minutus, C. ibicuiensis and C. torquatus. Almost all of these species are variable at the chromosomal level, suggesting that this is the ancestral condition of the clade. Within the Corrientes group, the observed low genetic divergence, in contrast with its high chromosomal variability, is indicative of decoupling between the rates of chromosomal and mitochondrial evolution.     

Phylogeographic patterns and conservation units of a vulnerable species,Cabot’s tragopan (<Emphasis Type="Italic">Tragopan caboti</Emphasis>), endemic to southeast China

Cabot’s tragopan (Tragopan caboti) is a pheasant endemic to southeast China and is protected under both national and global legislation due to ongoing decline in population size and increased habitat fragmentation. We investigated the phylogeographic patterns and examined the consistency between evolutionary units and assumed subspecies taxonomy in this species. Six populations across the whole species presenting a wide distribution range were sampled and two mitochondrial DNA segments (control region, CR and cytochrome b, cyt b) were used in this study. The results demonstrated a high level of genetic diversity (h = 0.97 in CR and 0.78 in cyt b) and significant differentiation among populations. Phylogenetic analyses strongly indicated two reciprocally monophyletic clades named the “West group” and “East group” that were not consistent with the present subspecies regional distribution. The divergence time between the two groups was estimated to be around 5.54 × 10⁵–8.7 × 10⁵ years ago, and the expansion times of the two groups were close (about 3 × 10⁵ years ago), indicating the effect of glaciation in intraspecific differentiation. Based on the results of genetic analyses combined with geographic isolation and distinct population history, our findings suggest that two management units (MUs) should be defined in T. caboti for conservation.     

Non-random distribution of the pericentromeric heterochromatin in meiotic prophase nuclei of mammalian spermatocytes

The central or peripheral distribution of condensed chromatin (CC) was studied in pachytene spermatocyte nuclei in Mus domesticus, 2n=40; Pudu puda, 2n=70; Ctenomys opimus, 2n=26 and Octodon degus, 2n=58. Species were chosen according to the morphological characteristics of their chromosomal complements and in particular, the terminal or medial chromosomal localisation of the pericentromeric constitutive heterochromatin. Counts were made by defining the areas corresponding to peripheral and central location in each nuclear section from a series. The null hypothesis (i.e. random distribution of CC) was rejected. In the nuclear sections of Mus domesticus and Pudu puda, 69% and 74% of CC, respectively, was found in the peripheral nuclear space, while in those of Octodon degus and Ctenomys opimus, 69% and 65% of CC, respectively, was found in the central nuclear space. We estimate that if the CC measured in spermatocyte nuclei corresponds mainly to pericentromeric constitutive heterochromatin, the distribution found is consistent with that expected in accordance with the nuclear architecture model for meiocytes (Fernández-Donoso, 1982; Fernández-Donoso & Berrios, 1985). This model proposes a peripheral nuclear localisation for pericentromeric heterochromatin of telocentric bivalents and a relatively central nuclear localisation for pericentromeric heterochromatin of metacentric bivalents. We also discuss some of the biological consequences that could arise from the conservation of such distributions. This revised version was published online in July 2006 with corrections to the Cover Date.     

Lagomorphs from the Miocene of Sandelzhausen (southern Germany)

Three genera of lagomorphs, Prolagus, Lagopsis, and “Amphilagus,” were identified during a revision of the lagomorph material from Sandelzhausen (MN5, Early/Middle Miocene boundary, southern Germany). Evidence of two morphological and dimensional classes were observed at some tooth positions in Prolagus (some p3 show an unmistakable P. oeningensis morphology, others closely resemble P. crusafonti), but not at other tooth positions (e.g., M1–2). Insufficient data from Sandelzhausen precludes identification of two different species of Prolagus from this locality, and to define the characteristics of the possible P. crusafonti-like species. Thus, all Prolagus specimens have been classified as P. aff. oeningensis. The genus Lagopsis is represented by L. cf. penai, whose presence is compatible with a MN5 age. The relative abundance of Lagopsis to Prolagus may indicate relatively cool and wet palaeoclimatic conditions. The largest primitive lagomorph species from continental Europe is present at Sandelzhausen. Morphological and dimensional comparisons with other European primitive lagomorphs exclude any affinity with the genera Eurolagus and Titanomys and with the species included in “Amphilagus ulmensis”. Some common features with “Amphilagus antiquus” were observed, although they are not sufficient for the attribution to this taxon. Until there is a general revision of European primitive lagomorphs, the Sandelzhausen giant lagomorph is classified as “Amphilagus” sp. Its origins, whether from evolution within Europe or migration from Asia, remain unknown.      

Restudy and reassignment ofDentalium antiquum Goldfuss, 1841 (Middle Devonian)

Dentalium antiquum is one of two moderately well-known Devonian “scaphopods” in the German literature. Examination of the type material and a few specimens in other institutions indicates more individual variability than is to be expected in species of the molluscan class Scaphopoda. The species is transferred with question toColeolus Hall, a fossil presumed to be a calcareous “worm” tube.      

Characterization of the cDNAs Encoding Three GnRH Forms in the Pejerrey Fish <Emphasis Type="Italic">Odontesthes bonariensis</Emphasis> (Atheriniformes) and the Evolution of GnRH Precursors

Most vertebrates express two gonadotropin releasing hormone (GnRH) variants in brain tissue but there is an increasing number of fish species for which a third GnRH form has been detected. We characterized the precursors (cDNAs) of all three forms expressed in the brain of the pejerrey (silverside) fish, Odontesthes bonariensis (Atheriniformes): type I (GnRH-I; 440 bp), type II (GnRH-II; 529 bp), and type III (GnRH-III; 515 bp). The expression of these GnRHs precursors was also observed in peripheral tissues related to reproduction (gonads), visual and chemical senses (eye and olfactory epithelium), and osmoregulation (gill), suggesting that in teleost fish and possibly other vertebrates GnRH mediates directly or indirectly many other functions besides reproduction. We also present a comprehensive phylogenetic analysis including representatives of all chordate GnRH precursors characterized to date that supports the idea of two main paralogous GnRH lineages with different function. A “forebrain lineage” separates evolutionarily from the “midbrain lineage” as a result of an ancient duplication (ca. 600 million years ago). A third, fish-only clade of GnRH genes seems to have originated before the divergence of fish and tetrapods but retained only in fish. Phylogenetic analyses of GnRH precursors (DNA and protein sequences) under different optimality criteria converge on this result. Although alternative scenarios could not be statistically rejected in this study due to the relatively short size of the analyzed molecules, this hypothesis also receives support from chromosomal studies of synteny around the GnRH genes in vertebrates. [Reviewing Editor: Dr. Axel Meyer]     

Materials towards the revision of the genus Triplophysa rendahl, 1933 (Cobitoidea: Balitoridae: Nemacheilinae): A revision of nominal taxa of Herzenstein (1888) described within the species “Nemachilus” stoliczkae and “N.” dorsonotatus, with the description of the new species T. scapanognatha sp. nova

This study reviews the nominal taxa within the species “Nemachilus” stoliczkae and “N.” dorsonotatus, described by Herzenstein (1888). N. dorsonotatus, N. d. retropinnis, and N. plagiognathus are reviewed within the species Triplophysa stoliczkae; it is suggested that this species is represented by at least 5 subspecies. N. stoliczkae brevicauda, N. s. leptosoma (synonym, N. s. productus), and N. s. crassicauda are considered as valid species, the last 2 are related to the group including the species T. tenuis, T. choprai, T. tenuicauda, and T. yasinensis rather than with T. stoliczkae. It was shown that the “stoliczkae” species group (sensu Prokofiev, 2001) should be divided into three species groups: “stoliczkae” (s. str.), “tenuis,” and “robusta”; their diagnoses and the species composition are presented. A new species from the “stoliczkae” group is described (T. scapanognatha sp. nova), related to T. tanggulaensis (Zhu, 1982). Original Russian Text ? A.M. Prokofiev, 2007, published in Voprosy Ikhtiologii, 2007, Vol. 47, No. 1, pp. 5–25.     

Chromosome multiformity in the genus Ctenomys (Rodentia,Octodontidae)

The karotypes of eleven species of the South American burrowing rodents, genus Ctenomys are described, and information on the somatic number of two other species of the same genus is given. The studied species are: C. torquatus (2n =68), C. tuconax (2n =61), C. minutus (2n=50), C. talarum (2n= 48), C. porteousi (2n=48), C. cf. minutus (2n=48), C. australis (2n=46), C. azarae (2n=48), C. latro (2n=42), C. magellanicus fueguinus (2n=36), C. tucumanus (2n=28), C. opimus luteolus (2n=26), and C. occultus (2n =22). This extreme intrageneric variation in somatic number is also reflected by a great amount of diversity in chromosome structure. Karyotypes seem to be rather constant at the species level. Autosomal polymorphism has been found in two of the species, namely C. talarum and C. latro. The hypothesis of the superimposition of Robertsoman rearrangements, pericentric inversions and translocations in the evolution of the karyotype of Ctenomys is advanced. The direction of chromosome change, either toward increase or decrease in chromosome number, is discussed. It is emphasized that high chromosome multiformity is correlated in Ctenomys with a rapid and explosive pattern of species diversification; the meaning of the small size of populations in enhancing the role of chromosome rearrangements in the evolution of Ctenomys is discussed.     

Karyotypic and molecular polymorphisms in <Emphasis Type="Italic">Ctenomys</Emphasis><Emphasis Type="Italic">torquatus</Emphasis> (Rodentia: Ctenomyidae): taxonomic considerations

The rodent genus Ctenomys (tuco-tucos) comprises more than 60 described species, and shows extraordinary inter- and intraspecific karyotypic variation. The most widely distributed species of Ctenomys in Brazil is C. torquatus. Although several cytogenetic studies have been done, the karyotypic variability of this species is still poorly known. In this paper we report two new diploid numbers for C. torquatus: 2n = 40 and 2n = 42, both showing AN = 72. The new distribution limits of C. torquatus here reported include localities in the southern, central and western parts of Rio Grande do Sul (RS) State in southern Brazil. The phylogenetic relationship between C. torquatus from Alegrete, RS, and Ctenomys sp. from Corrientes, Argentina, is described by means of mtDNA cytochrome b analysis. Although both entities share similar karyotypes and sperm morphology, these two species are not phylogenetically close.     

Coexistence of congeneric spiny lobsters on coral reefs: differences in conspecific aggregation patterns and their potential antipredator benefits

Den sharing by conspecific spiny lobsters (aggregation) is modulated by chemical attraction but may confer several, not necessarily mutually exclusive, antipredator byproduct benefits: a “guide effect”, which only benefits the individual attracted to a sheltered conspecific; a “dilution effect”, which reduces per-capita risk of predation simply through aggregation; or active “group defense”. Each potential benefit has a different set of predictors (relationships between aggregation and conspecific or predator densities), but conflicting results could suggest the simultaneous operation of more than one benefit. These predictions were tested for coexisting Panulirus guttatus (a reef-obligate) and Panulirus argus (a temporary reef-dweller) using data collected during 11 surveys on fixed sites over a coral reef in Mexico. P. guttatus greatly outnumbered P. argus, but P. argus showed a greater tendency to aggregate. All three benefits of den sharing operated for the more social P. argus, with “group defense” being of the most benefit for larger individuals, and the “guide” and “dilution” effects for smaller individuals recently immigrating into the reef habitat and sharing dens with larger conspecifics. P. guttatus did not display “group defense” and its aggregations appeared to be modulated by the interplay between attraction and aggressive behaviors. This species relied more on solitary crypticity, especially at larger sizes, but appeared to benefit from a “guide effect” at high conspecific densities. In experimental tanks, each species tended to aggregate when tested separately, but when tested jointly, aggregation among P. guttatus was significantly reduced. The experimental results reflect the differential patterns of aggregation between the fore-reef, where P. guttatus dominated, and the back-reef, where coexistence of both species was greater.     

Resolving relationships in some African fungus-growing termites (Termitidae, Macrotermitinae) using molecular phylogeny, morphology, and field parameters

The Macrotermitinae are a large and successful subfamily of fungus-growing termites, characterised by their symbiotic association with white-rot fungi of the genus Termitomyces. The taxonomy of the subfamily, and in particular of the largest genus Odontotermes, is problematic. We used sequences of the mitochondrial 16S gene from termites occurring in East Africa and Malawi to explore the phyletic relationships within the genus Odontotermes and to place the genus in the broader context of other fungus-growing termites. We also interpret this phylogeny in relation to classical morphological taxonomy in the form of absolute and relative dry weights of the sterile castes, and in relation to innate behaviour as shown by nest architecture and fungus comb structure. This work lays the foundations of a complete taxonomic revision of the Macrotermitinae. The phylogeny supports one clear subdivision, here called the “tanganicus” group, within the genus Odontotermes. It also supports the significance of field observations on the structure of fungus combs, as the whole “tanganicus” group builds fungus combs of the sponge type (or modified forms thereof) which are hardly known elsewhere. Other phyletic relationships are less clear, the residual sequences being referred to as the “latericius” group. We recognise three probable miniature species within the “tanganicus” group and another possible one in the residual “latericius group”. Received 16 January 2007; revised 30 October 2007 and 11 March 2008; accepted 4 April 2008.     

A simple chromosomal marker can reliably distinguishes <Emphasis Type="Italic">Poncirus</Emphasis> from <Emphasis Type="Italic">Citrus</Emphasis> species

Several chromosome types have been recognized in Citrus and related genera by chromomycin A₃ (CMA) banding patterns and fluorescent in situ hybridization (FISH). They can be used to characterize cultivars and species or as markers in hybridization and backcrossing experiments. In the present work, characterization of six cultivars of P. trifoliata (“Barnes”, “Fawcett”, “Flying Dragon”, “Pomeroy”, “Rubidoux”, “USDA”) and one P. trifoliata × C. limonia hybrid was performed by sequential analyses of CMA banding and FISH using 5S and 45S rDNA as probes. All six cultivars showed a similar CMA⁺ banding pattern with the karyotype formula 4B + 8D + 6F. The capital letters indicate chromosomal types: B, a chromosome with one telomeric and one proximal band; D, with only one telomeric band; F, without bands. In situ hybridization labeling was also similar among cultivars. Three chromosome pairs displayed a closely linked set of 5S and 45S rDNA sites, two of them co-located with the proximal band of the B type chromosomes (B/5S-45S) and the third one co-located with the terminal band of a D pair (D/5S-45S). The B/5S-45S chromosome has never been found in any citrus accessions investigated so far. Therefore, this B chromosome can be used as a marker to recognize the intergeneric Poncirus × Citrus hybrids. The intergeneric hybrid analyzed here displayed the karyotype formula 4B + 8D + 6F, with two chromosome types B/5S-45S and two D/5S-45S. The karyotype formula and the presence of two B/5S-45S chromosomes clearly indicate that the plant investigated is a symmetric hybrid. It also demonstrates the suitability of karyotype analyses to differentiate zygotic embryos or somatic cell fusions involving trifoliate orange germplasm. During the submission of this paper, we analyzed 25 other citrus cultivars with the same methodology and we found that the chromosome marker reported here can indeed distinguish Poncirus trifoliata from grapefruits, pummelos, and one variegated access of Citrus, besides the previously reported access of limes, limons, citrons, and sweet-oranges. However, among 14 mandarin cultivars, two of them displayed a single B/5S-45S chromosome, whereas in Citrus hystrix D.C., a far related species belonging to the Papeda subgenus, this chromosome type was found in homozygosis. Since these two mandarin cultivars are probably of hybrid origin, we assume that for almost all commercial cultivars and species of the subgenus Citrus this B type chromosome is a useful genetic marker.     

Embryonic β-globin in the non-Antarctic notothenioid fish Cottoperca gobio (Bovichtidae)

Haemoglobins are sensitive to temperature and their properties mirror the thermal conditions encountered by species during their evolutionary histories. This paper provides data on molecular phylogeny of the haemoglobin chains of Cottoperca gobio, a notothenioid fish of sub-Antarctic latitudes, belonging to the basal family Bovichtidae. Unlike most Antarctic notothenioids, C. gobio has two major haemoglobins sharing the β chain. In the molecular phylogenetic analysis, the β chain is included in the clade of the “embryonic” or minor Antarctic globins. Although, in the majority of notothenioids, “embryonic” (minor) α and β globins are expressed in traces or small amounts in the adult stage, in C. gobio the present analysis supports the occurrence of a complete “switch” to exclusive expression of the embryonic β-globin gene in adult fish. The α and β chains sequences have been used to expand our knowledge of the evolution of notothenioid haemoglobins.The protein sequence data reported in this paper will appear in the UniProt Knowledge base under the accession number: P84652 (β chain), P84653 (α ¹ chain).     

Molecular kinship analyses of the agaricoid Russulaceae: Correspondence with mycorrhizal anatomy and sporocarp features in the genus Russula

A data set of LSU DNA sequences of mainly European Russula and Lactarius species was subjected to molecular phylogenetic analysis. Species could be allocated to six clades, with an unresolved phylogeny. One of these clades represents the genus Lactarius. The only analysed species of the section Archaeinae (Russula) was placed basal to both genera. Thus Lactarius appears to be derived from Russula. Russula was divided into four clusters, corresponding to the sections Plorantinae and Nigricantinae, subgenus Heterophyllidia including the section Foetentinae, and a cluster representing the remaining subgenera of the “Genuinae”. Even though the resulting groups can be considered as valid classificatory groups, species associations resulting from molecular analyses neither support the division of Russula into the subgenus Compacta (including the sections Nigricantinae, Plorantinae, and Archaeinae) and the “Genuinae” (including all remaining taxa), nor do they support previously proposed evolutionary lineages within the “Genuinae”. Ribosomal ITS DNA sequences of Russula species were analysed to achieve better infrageneric resolution. The results are discussed in relation to current classification systems and to what is known about the mycorrhizae formed by Russula species. While the systematic value attached to many macroscopic and microscopic sporocarp features was not supported by sequence data, mycorrhizal anatomy is in good correspondence with many of the results from the phylogenetic analysis.