Developments in avian cytotaxonomy: implications for Afro-avian Species

Ejere. VC. 2000. Developments in avian cytotaxonomy: implications for Afro-avian Species. Ostrich 71 (1 & 2): 40.

Karyological studies of the extant species of birds have progressed slower than those of other animal groups. To date, a paltry proportion of less than 20% of about an estimated 9 000 avian species have been karyotyped, such that for most orders, karyological information remains largely scanty. Data accumulated so far, has revealed a lot of interesting features of the typical avian karyotype. Essentially, the karyotype is dichotomous, containing several pairs of fairly big chromosomes (= macrochomosomes) and very small to minute chromosome elements (= microchromosmes). The diploid chromosome number is also variable, ranging from 2n = 40 to 2n = 92 with a mode of 2n = 80 chromosomes observed in a majority of species. Studies have further revealed that chromosomal evolution in birds is highly conservative especially with reference to the first 3 pairs of macrochromosomes. (Group A), while considerable variability in number and morphology occurs in the remaining groups of chromosome elements. These variabilities, as well as the homologies revealed by chromosome banding techniques, have aided the cytotaxonomy of various individual avian groups. In this review, the karyological as well as the phylogenetic relationships of the various avian species so far karyotyped, vis-a-vis the cytotaxonomic implications for the abundant Afro-avian fauna are highlighted. Similarly, the significance of the observed peculiarities in the avian karyotype is briefly discussed. Suggestions are proffered in respect of the application of karyotype analysis technique to bird conservation in Africa. This will focus mainly on the identification of the gender of birds for the singular purpose of breeding rare and endangered species for zoological gardens and related institutions.     

Chromosome painting in<Emphasis Type="Italic"> Callicebus lugens</Emphasis>, the species with the lowest diploid number (2<Emphasis Type="Italic">n</Emphasis>=16) known in primates

Cytogenetic studies have shown that New World primates are karyologically diverse and highly derived. The genus Callicebus is the best example of this karyological diversity, with diploid numbers ranging from 2n=50 to 2n=16. We report on Callicebus lugens, which has the lowest diploid number (2n=16) yet found in the primate order and represents a striking example of extreme karyotypic shuffling. To better understand the genomic rearrangements that have resulted in this extremely low diploid number, we mapped chromosome homologies between C. lugens and humans by in situ hybridization. The total number of hybridization signals was 42, excluding the Y chromosome, with a total of 34 syntenic associations not found in humans. This species has one of the most derived karyotypes among the Platyrrhini. Fusion has been the predominant mode of karyological evolution, although fissions and inversions have also transformed the C. lugens karyotype. Remarkably in such a highly rearranged karyotype, the synteny of 11 human chromosomes (4, 5, 9, 12, 13, 14, 17, 18, 20, 21, and X) was maintained intact, even if most of these human-homologous gene clusters were translocated. Other human syntenies, such as homologues to human chromosomes 10 and 16, were highly fragmented. Comparisons of the C. lugens-human homology map with those of other New World primates have not yet helped establish a phylogenic arrangement between congeneric species or link Callicebus with any other genus.Communicated by S. Henikoff     

Trends in the karyotypic evolution of the Neotropical catfish family Heptapteridae Bockmann 1998 (Teleostei: Siluriformes)

The Heptapteridae family is endemic to the Neotropics and is one of the most representative members of the Siluriformes order in small bodies of water. Due the lack of data for clearly distinguishing their species, only recently was elevated from condition of subfamily to family. However, many doubts about certain species remain yet. There are a relatively large number of cytogenetics studies about heptapterids that permits comparative analyses focused in the karyotype evolution of the group. However, the absence of studies about this theme motivated this work. Were analyzed cytogenetically most representative species of the family: Rhamdia, Imparfinis, Pimelodella, Phenacorhamdia, and Taunaya, and compared with available data from the family. The comparative analysis indicated the predominance of biarmed chromosomes and low variation of the diploid number (around 2n = 58) suggesting that centric inversions were more important than centric fissions in the history of heptapterids. It is corroborated by a remarkable number of NOR phenotypes mainly in Imparfinis and Rhamdia species. A probable condition of Heptapteridae plesiomorphic karyotype would be one composed of 2n = 58 chromosomes of types metacentric and submetacentric mainly, with simple NOR at the sub terminal position. The reduction of diploid number, it would possibly through successive events of chromosome fusion with deletions and inversion rearrangements responsible for NOR variation within the certain genus. The occurrence of B chromosomes is from a derived event, possibly of recent origin, without phylogenetic implications.     

七种木蓝属植物的染色体研究

本文采用根尖压片法对豆科（Leguminosae）木蓝属（IndigoferaL.）植物的7个种：多花木蓝（Indigofera amblyantha Craib）、河北木蓝（I.bungeana Walp.）、滇木蓝（I.delavayi Franch.）、腺毛木蓝（I.scabrida Dunn）、四川木蓝（I.szechuensis Craib）、刺序木蓝（I.silvestrii Pamp.）、尖叶木蓝（I.zollingeriana Miq.）的染色体数目和核型进行了研究。结果表明：除了尖叶木蓝染色体数目为2n=4x=32（四倍体）外,其余6种木蓝的染色体数目均为2n=2x=16（二倍体）。尖叶木蓝和刺序木蓝核型分类为2A型,其余5种木蓝的核型均为1A型。种间核型差异很小。供试种主要包含中部着丝点区染色体。滇木蓝、腺毛木蓝、刺序木蓝、四川木蓝和尖叶木蓝的核型分析为首次报道。     

Extreme degree of chromosome number variability in species of the spider genus Scytodes (Araneae, Haplogynae, Scytodidae)

Genus Scytodes includes most species of the spider family Scytodidae. Until now, 187 species of the genus have been described. In spite of this great diversity, only three Scytodes species were karyotyped so far. The present paper provides for the first time karyotype analysis of two synanthropic species, Scytodes fusca and Scytodes itapevi. Furthermore, new data on karyotype of Scytodes globula are also provided using conventional and differential cytogenetical procedures. The diploid number in the genus Scytodes varied considerably, namely from 2n = 13 to 2n = 31. The diploid number found in S. globula (2n♂ = 13) is the lowest in haplogyne spiders with monocentric chromosomes. Except S. globula, this number has been found only in one haplogyne spider with monocentric chromosomes, namely Ochyrocera sp. (Ochyroceratidae). On the contrary, the diploid number of S. fusca (2n♂ = 31) is one of the highest diploid numbers recorded in haplogyne spiders. The degree of intrageneric variation found in the genus Scytodes is the highest recorded in araneomorph spiders with monocentric chromosomes so far. Some karyotype characteristics (diploid number, chromosome morphology, total chromosome length, and distribution of constitutive heterochromatin) allowed us to postulate a close relationship between S. globula and S. itapevi. According to the karyotype data, S. fusca is not closely related to these two species. This conclusion corroborates a recent taxonomic work that grouped S. globula, S. itapevi, and other four Scytodes species in the ‘globula group’.     

Karyotype morphology in Hyperiella dilatata Stebbing 1888 (Amphipoda: Hyperiidae) from the Ross Sea (Antarctica)

Summary The chromosome complement and some karyological features were investigated in the pelagic amphipod Hyperiella dilatata Stebbing 1888 from the Ross Sea (Antarctica). The diploid karyotype consists of 48 metacentric and 10 submetacentric elements (2n = 58). The presence of secondary constrictions and supernumerary chromosomes is described. Available chromosome numbers of Hyperiidea exhibit a wide range of distribution, among which Hyperiella dilatata is the closest to the modal number of other amphipods.     

Molecular genetic and karyological analysis of antlered sculpins of <Emphasis Type="Italic">Enophrys diceraus</Emphasis> group (Cottidae)

Molecular genetic and karyological analyses of antlered sculpin, Enophrys diceraus, from the Sea of Japan and the Sea of Okhotsk were carried out. The karyotype of this species was studied for the first time. On the basis of karyological analysis, it was established that E. diceraus from the Sea of Japan and the Sea of Okhotsk was polymorphic in terms of the number of chromosomes and their morphology (2n = 36, 35, and 37, NF = 40). It was suggested that the karyotype with 35 chromosomes could have been produced as a result of Robertsonian translocation; the karyotype with 37 chromosomes could have been produced by crossing of individuals with different number of chromosomes. On the basis of the molecular genetic analysis of the mitochondrial COI and 16S rRNA genes, considerable differences between E. diceraus from the Sea of Japan and the Sea of Okhotsk corresponding to the level of interspecies genetic variability were established. It is concluded that E. diceraus from the Sea of Japan belongs to another species, most likely, E. namiyei.     

A new karyotype for <Emphasis Type="Italic">Cavia magna</Emphasis> (Rodentia: Caviidae) from an estuarine island and <Emphasis Type="Italic">C. aperea</Emphasis> from adjacent mainland

We describe a new karyotype for Cavia magna Ximenez, 1980 from an estuarine island and the karyotype of Cavia aperea Erxleben, 1777 from an adjacent mainland. The species have differences in diploid number (2n), autosomal fundamental number, quantity, and distribution of heterochromatin as dissimilar distributions of the nucleolus-organizing regions (Ag-NORs). The C. aperea karyotype has a diploid number of 64 as previously reported for C. aperea and most other Cavia species. In contrast, this new C. magna karyotype exhibits a variant diploid number of 2n = 62, considering that previous work reported a karyotype of 2n = 64 for C. magna. The discovery of a distinct diploid number within C. magna represents the first record of intra-specific chromosomal variation in a species of Caviidae. The diploid number of 2n = 62, heterochromatin quantity, Ag-NOR distributions, and inversed X chromosome from this population of C. magna are as seen in the geographically proximate (Cavia intermedia Cherem Olimpio and Ximenez; intermediate Cavy). These data provide further evidence supporting C. magna as the sister species of C. intermedia.     

Repetitive DNA Sequences and Evolution of ZZ/ZW Sex Chromosomes in Characidium (Teleostei: Characiformes)

Characidium constitutes an interesting model for cytogenetic studies, since a large degree of karyotype variation has been detected in this group, like the presence/absence of sex and supernumerary chromosomes and variable distribution of repetitive sequences in different species/populations. In this study, we performed a comparative cytogenetic analysis in 13 Characidium species collected at different South American river basins in order to investigate the karyotype diversification in this group. Chromosome analyses involved the karyotype characterization, cytogenetic mapping of repetitive DNA sequences and cross-species chromosome painting using a W-specific probe obtained in a previous study from Characidium gomesi. Our results evidenced a conserved diploid chromosome number of 2n = 50, and almost all the species exhibited homeologous ZZ/ZW sex chromosomes in different stages of differentiation, except C. cf. zebra, C. tenue, C. xavante and C. stigmosum. Notably, some ZZ/ZW sex chromosomes showed 5S and/or 18S rDNA clusters, while no U2 snDNA sites could be detected in the sex chromosomes, being restricted to a single chromosome pair in almost all the analyzed species. In addition, the species Characidium sp. aff. C. vidali showed B chromosomes with an inter-individual variation of 1 to 4 supernumerary chromosomes per cell. Notably, these B chromosomes share sequences with the W-specific probe, providing insights about their origin. Results presented here further confirm the extensive karyotype diversity within Characidium in contrast with a conserved diploid chromosome number. Such chromosome differences seem to constitute a significant reproductive barrier, since several sympatric Characidium species had been described during the last few years and no interespecific hybrids were found.     

Gene mapping of 28S and 5S rDNA sites in the spined loach Cobitis taenia (Pisces, Cobitidae) from a diploid population and a diploid–tetraploid population

We compare the chromosomal 28S and 5S rDNA patterns of the spined loach C. taenia (2n = 48) from an exclusively diploid population and from a diploid–polyploid population using 28S and 5S rDNA probe preparation and labelling, and fluorescence in situ hybridization (FISH). The 5S rDNA was located in two to three chromosome pairs, and separated from the 28S loci for the males and one female (F1) from the diploid population. Loaches from a diploid–polyploid population, and one female (F2) from the diploid population were characterized by at least one chromosome pair with 5S and 28S overlapping signals. The fishes differed mainly in their number of 28S rDNA loci, located on 3–6 chromosomes. All individuals from both populations were characterized by one acrocentric chromosome bearing a 28S rDNA signal on the telomeres of its long arm. The number of major ribosomal DNA in the karyotype of C. taenia by FISH was always higher than the number of Ag-NORs. Our data confirm the extensive polymorphism of NORs in both populations, as already has been observed in closely related Cobitis species, and less polymorphic 5S rDNA pattern. However, this preliminary result highlights the need for a wider scale study.     

Reinvestigation of the chromosomes of a male owl monkey (Aotus trivirgatus) and its hybrid son

A male owl monkey, probably belonging to Aotus trivirgatus nigriceps, was found to have 51 chromosomes. Since the Y chromosome is lacking, the odd diploid number probably is the result of a Y-autosome translocation. However, there are two autosomes from different pairs that theoretically could contain the Y. This specimen produced a male young with a female A. trivirgatus griseimembra, having a karyotype with 54 chromosomes. The diploid number of the hybrid is 52, including 28 paired and 24 unpaired elements. Comparison of the paternal, maternal and hybrid karyotypes using Q- and C-banding techniques permit conclusions on the rearrangements (fusions, reciprocal translocations, inversions and insertions) that could possibly have led to the karyological differences between both subspecies.     

A cytogenetic study on three Chilean species of Chrysomelinae (Coleoptera, Chrysomelidae)

Three species of Chilean leaf beetles were chromosomally analyzed. The endemic Araucanomela wellingtonensis displays a male meioformula of 13 + Xyp with 2n = 28 chromosomes and an asymmetric karyotype with two large autosome pairs and 12 medium/small pairs of autosomes and sex-chromosomes, a diploid number which had not been found among the other species of the subtribe Paropsina sensu lato studied to date. Strichosa eburata presents a meioformula of 11 + Xyp, 2n = 24 chromosomes, as occurs in many species of chrysomelines belonging to different subtribes. Furthermore, Phaedon cyanopterum has a 16 + Xyp meioformula, that is 2n = 34 chromosomes, of small size mostly, also in agreement with the karyological findings obtained in all the other congeneric species so far examined. These cytogenetic data are discussed with respect to the previous ones in this subfamily and with other characters of taxonomic and evolutionary value.     

Cytology of Encephalartos hildebrandtii A. Br. & Bouche

The chromosomes of Encephalartos hildebrandtii A. Br. &Bouche were studied from root tip cells (2n–27), and theplant was found to be a triploid on the basic chromosome number9. Karyotype analysis has shown that the somatic complementconsists of two sets of chromosomes, a diploid set of 18 chromosomesin nine paris, and a haploid set of 9, and therefore it is suggestedthat this is an allotriploid. This is the first report of a polyploid species in the genusEncephalartos, and also in the whole of the Cycadales.     

Banded karyotype of spined loach Cobitis taenia and triploid Cobitis from Poland

The present work provides new data on the banding pattern of diploid Cobitis taenia and its triploid hybrid females, which belong to the diploid–polyploid complex in the Vistula River tributary. C-banding, silver-staining (Ag), and fluorescent staining with chromomycin A₃ techniques were used to describe the diploid and triploid karyotype. The karyotype of Cobitis taenia of 2n=48 was characterised by one pair of NOR-bearing subtelocentric chromosomes and at least four chromosomes with CMA₃-positive sites. The C-positive heterochromatin was present in the centromeres of almost all chromosomes and the pericentromeric regions of several metacentric and submetacentric chromosomes. The triploid females of 3n=74 had two pairs of chromosomes with active NORs. The NORs-sites were located terminally on two biarmed and two uniarmed chromosomes. The CMA₃-staining revealed at least six A₃-positive sites. The C-banded and A₃-stained triploid karyotype was composed of haploid set of Cobitis taenia and diploid set of unidentified species, so heterochromatin pattern confirmed the possibility of their hybrid origin. The characteristics of banded diploid and triploid karyotype, and the hypothetical karyotype of an unknown species of 2n=50 is discussed. This revised version was published online in July 2006 with corrections to the Cover Date.     

On the highest chromosome number in mammals

The mitotic and meiotic chromosomes of the semiaquatic rodent Ichthyomys pittieri (Rodentia, Cricetinae) from Venezuela were analyzed by means of conventional staining and several banding techniques. The diploid chromosome number of this rare species is 2n = 92, which is the highest value known for mammals. It is assumed that this exceptionally high chromosome number is the result of repeated centric fissions. The karyotype of I. pittieri was compared with that of Anotomys leander, for which a diploid number of 2n = 92 has also been reported. The karyological relationships existing within the Neotropical Cricetidae are summarized.     

The Role of Fusion in Ant Chromosome Evolution: Insights from Cytogenetic Analysis Using a Molecular Phylogenetic Approach in the Genus Mycetophylax

Among insect taxa, ants exhibit one of the most variable chromosome numbers ranging from n = 1 to n = 60. This high karyotype diversity is suggested to be correlated to ants diversification. The karyotype evolution of ants is usually understood in terms of Robertsonian rearrangements towards an increase in chromosome numbers. The ant genus Mycetophylax is a small monogynous basal Attini ant (Formicidae: Myrmicinae), endemic to sand dunes along the Brazilian coastlines. A recent taxonomic revision validates three species, Mycetophylax morschi, M. conformis and M. simplex. In this paper, we cytogenetically characterized all species that belongs to the genus and analyzed the karyotypic evolution of Mycetophylax in the context of a molecular phylogeny and ancestral character state reconstruction. M. morschi showed a polymorphic number of chromosomes, with colonies showing 2n = 26 and 2n = 30 chromosomes. M. conformis presented a diploid chromosome number of 30 chromosomes, while M. simplex showed 36 chromosomes. The probabilistic models suggest that the ancestral haploid chromosome number of Mycetophylax was 17 (Likelihood framework) or 18 (Bayesian framework). The analysis also suggested that fusions were responsible for the evolutionary reduction in chromosome numbers of M. conformis and M. morschi karyotypes whereas fission may determines the M. simplex karyotype. These results obtained show the importance of fusions in chromosome changes towards a chromosome number reduction in Formicidae and how a phylogenetic background can be used to reconstruct hypotheses about chromosomes evolution.     

Karyotype analysis and new chromosome number reports in <Emphasis Type="Italic">Silene</Emphasis> species (sect. <Emphasis Type="Italic">Auriculatae</Emphasis>, Caryophyllaceae)

Karyotype study was performed in 13 populations of 11 Silene species (sect. Auriculatae L., Caryophyllaceae) growing in Iran. All the species studied showed 2n = 2x = 24 chromosome number supporting the earlier report on S. meyeri, while the chromosome number of S. palinotricha, S. sojakii, S. gertraudiae, S. elymaitica, S. pseudonurensis, S. dschuparensis, S. eriocalycina, S. araratica, S. prilipkoana and S. commelinifolia are new to science. The chromosomes were mainly metacentric or sub-metacentric and their size varied from 1.10 μm in S. pseudonurensis to 7.11 μm in S. dschuparensis. The species studied differed significantly in the total size of the chromosomes, the size of the short arms and the long arms, indicating the role of quantitative genomic changes in the Silene species diversification. They also differ in their karyotype formulae indicating the occurrence of structural changes in their chromosomes. The Silene species were placed in 1A, 2A, 1B and 2B classes of Stebbins karyotype symmetry showing symmetrical karyotypes. Clustering of the species based on karyotype features grouped the species of S. palinotricha, S. prilipkoana, S. commelinifolia, S. eriocalycina, S. meyeri, S. araratica and S. Sojakii together while the species of S. gertraudiae and S. elymaitica showed more similarity and were placed close to each other.     

Clarification of the systematic position of Cercariaeum crassum Wesenberg-Lund, 1934 (Digenea), based on karyological analysis and DNA sequences

Chromosome set and rDNA sequences of the larval digenean Cercariaeum crassum were analysed in order to clarify its systematic position and possible adult form. Parasites were obtained from the sphaeriid bivalve Pisidium amnicum, collected in Lithuanian and Finnish rivers. The karyotype is shown to consist of five pairs (2n = 10) of large, up to 14 μm, chromosomes. Complement, composed of a low diploid number of exclusively bi-armed elements, presumably arose through Robertsonian fusions of acrocentric chromosomes. Consistent with a Robertsonian-derived karyotype, one or two small, metacentric, mitotically stable B chromosomes were detected in the cells of parthenitae isolated from some host individuals. A phylogenetic analysis using rDNA internal transcribed spacer 2 (ITS2) and 28S sequences corroborates the allocation of C. crassum to the family Allocreadiidae. In neighbour-joining and maximum parsimony phylogenetic trees C. crassum clusters into one clade with Allocreadium spp., and is the closest sister group in relation to A. isoporum; the level of rDNA sequence divergence between them (2.67% for ITS2 and 1.16% for 28S) is consistent with the level expected for intrageneric variation. The present study adds significant information to a database for establishing species-specific characters for confident characterization of different developmental stages of allocreadiid species, clarification of their life cycles and evaluation of intra- and interspecific variability.