Chromosomal evolution of the Canidae. II. Divergence from the primitive carnivore karyotype

The Giemsa-banding patterns of chromosomes from the arctic fox (Alopex lagopus), the red fox (Vulpes vulpes), the kit fox (Vulpes macrotis), and the raccoon dog (Nyctereutes procyonoides) are compared. Despite their traditional placement in different genera, the arctic fox and the kit fox have an identical chromosome morphology and G-banding pattern. The red fox has extensive chromosome arm homoeology with these two species, but has only two entire chromosomes in common. All three species share some chromosomes with the raccoon dog, as does the high diploid-numbered grey wolf (Canis lupus, 2n = 78). Moreover, some chromosomes of the raccoon dog show partial or complete homoeology with metacentric feline chromosomes which suggests that these are primitive canid chromosomes. We present the history of chromosomal rearrangements within the Canidae family based on the assumption that a metacentric-dominated karyotype is primitive for the group.     

The pattern of phylogenomic evolution of the Canidae

Canidae species fall into two categories with respect to their chromosome composition: those with high numbered largely acrocentric karyotypes and others with a low numbered principally metacentric karyotype. Those species with low numbered metacentric karyotypes are derived from multiple independent fusions of chromosome segments found as acrocentric chromosomes in the high numbered species. Extensive chromosome homology is apparent among acrocentric chromosome arms within Canidae species; however, little chromosome arm homology exists between Canidae species and those from other Carnivore families. Here we use Zoo-FISH (fluorescent in situ hybridization, also called chromosomal painting) probes from flow-sorted chromosomes of the Japanese raccoon dog (Nyctereutes procyonoides) to examine two phylogenetically divergent canids, the arctic fox (Alopex lagopus) and the crab-eating fox (Cerdocyon thous). The results affirm intra-canid chromosome homologies, also implicated by G-banding. In addition, painting probes from domestic cat (Felis catus), representative of the ancestral carnivore karyotype (ACK), and giant panda (Ailuropoda melanoleuca) were used to define primitive homologous segments apparent between canids and other carnivore families. Canid chromosomes seem unique among carnivores in that many canid chromosome arms are mosaics of two to four homology segments of the ACK chromosome arms. The mosaic pattern apparently preceded the divergence of modern canid species since conserved homology segments among different canid species are common, even though those segments are rearranged relative to the ancestral carnivore genome arrangement. The results indicate an ancestral episode of extensive centric fission leading to an ancestral canid genome organization that was subsequently reorganized by multiple chromosome fusion events in some but not all Canidae lineages.     

Chromosomal evolution within the family Estrildidae (Aves) I. The Poephilae

Eleven species of estrildid finches were examined cytogenetically with G- and C-banding. The analysis revealed a preponderance of pericentric inversions at both the inter- and intraspecific levels. In addition, considerable variation in the pattern of heterochromatin distribution, particularly in the sex-chromosomes, was recorded as polymorphisms and interspecific differences. This variation was not found to be associated directly with either speciation or morphological change. Rather, it is argued that only those rearrangements which do not lead to meiotic problems survive in avian lineages.The chromosomal data were also used to clarify systematic relationships within the Poephilae, demonstrating that the monotypic genera Aegintha and Aidemosyne are allied to the Neochmia group.     

Molecular evolution of the leptin exon 3 in some species of the family Canidae

The structure of the leptin gene seems to be well conserved. The polymorphism of this gene in four species belonging to the Canidae family (the dog (Canis familiaris) – 16 different breeds, the Chinese racoon dog (Nyctereutes procyonoides procyonoides), the red fox (Vulpes vulpes) and the arctic fox (Alopex lagopus)) were studied with the use of single strand conformation polymorphism (SSCP), restriction fragment length polymorphism (RFLP) and DNA sequencing techniques. For exon 2, all species presented the same SSCP pattern, while in exon 3 some differences were found. DNA sequencing of exon 3 revealed the presence of six nucleotide substitutions, differentiating the studied species. Three of them cause amino acid substitutions as well. For all dog breeds studied, SSCP patterns were identical.     

Chromosome numbers of fishes. I

The published chromosome numbers for 208 temperate and tropical freshwater fishes have been compiled in alphabetical order within the families. This facilitates the work for ichthyologists interested in the karyotype(s) of a particular species, genus or family as well as giving an overall account of the basic chromosome numbers and karyological trends in fishes.     

Chromosomal evolution in the South American Nymphalidae

We give the chromosome numbers of about 80 species or subspecies of Biblidinae as well as of numbers of neotropical Libytheinae (one species), Cyrestinae (4) Apaturinae (7), Nymphalinae (about 40), Limenitidinae (16) and Heliconiinae (11). Libytheana has about n=32, the Biblidinae, Apaturinae and Nymphalinae have in general n=31, the Limenitidinae have n=30, the few Argynnini n=31 and the few species of Acraeni studied have also mostly n=31. The results agree with earlier data from the Afrotropical species of these taxa. We supplement these data with our earlier observations on Heliconiini, Danainae and the Neotropical Satyroid taxa. The lepidopteran modal n=29-31 represents clearly the ancestral condition among the Nymphalidae, from which taxa with various chromosome numbers have differentiated. The overall results show that Neotropical taxa have a tendency to evolve karyotype instability, which is in stark contrast to the otherwise stable chromosome numbers that characterize both Lepidoptera and Trichoptera.     

Fixation probabilities and effective population numbers in diploid populations with overlapping generations

A finite diploid population, observed at times t = 0, 1, 2,…, is studied. An individual is said to be in age group i at time t if its age is between i and i + 1 units at that time, where i ? 1. It is assumed that the number of individuals in a particular age-sex class is the same for every t and that the probability that a male offspring was produced by a mating of a male in age group i and a female in age group j is p_ij^m (with a corresponding probability p^f_ij for a female offspring), regardless of when the individual is born. The probability of ultimate fixation of an allele A₁ and the inbreeding effective number, for large populations, is calculated under the further assumptions that A₁ is neutral and that mating is random, given the ages of the mates.     

Chromosomal evolution inEpilobium sect.Epilobium (Onagraceae)

WithinEpilobium sect.Epilobium, a cytological analysis of 121 experimental hybrids, involving 40 species, indicates the presence of a widespread BB chromosome arrangement in Eurasia, Africa, and Australasia, as well as in North and South America less commonly. The AA chromosome arrangement, which differs from BB by one reciprocal translocation, occurs in North America, South America, and in at least three European species. The CC arrangement, which differs from AA by two reciprocal translocations, characterizes theAlpinae, a circumboreal group. Distinctive or only partly worked out chromosome arrangements occur in the EuropeanE. duriaei andE. nutans and in the North AmericanE. luteum, E. obcordatum, E. oregonense, andE. rigidum. With earlier results, the chromosome arrangements of some 65 of the estimated 185 species of the section have been established fully or partly.     

Chromosomal features and evolution of Bromeliaceae

New cytological information and chromosome counts are presented for 19 taxa of 15 genera of the Bromeliaceae, among them, data for 15 taxa and five genera are reported for the first time. The basic number x = 25 is confirmed and polyploidy seems to be the main evolutionary mechanism in Bromeliaceae. Most of the analyzed species presented 2n = 50. Polyploids have been detected in Deinacanthon urbanianum with 2n = ca.160 and Bromelia laciniosa with 2n = ca.150. In Deuterocohnia lorentziana we observed individuals with two different ploidy levels (2n = 50 and 2n = 100) growing together in the same pot. Ayensua uaipanensis showed the uncommon number 2n = 46. After triple staining with CMA₃/Actinomycin/DAPI one or two CMA⁺/DAPI⁻ bands could be observed in the studied species (Aechmea bromeliifolia, Greigia sphacelata and Ochagavia litoralis). The role of these features in the evolution of the family is discussed, revealing new aspects of the evolution of the Bromeliaceae.     

I. I. Shmal'gauzen's concept of the evolution of ontogeny

The concept of evolution of ontogenesis by I. I. Shmal'gauzen is presented as a result of reviewing some of his theoretical works. This concept appears to be the most consistent development of classical darwinism on the basis of a basically new (as compared with synthetic theory of evolution) evolutionary-synthetic approach. This latter has been based on the idea of the organism integrity in onto- and phylogenesis and the involvement of the organismic level (ontogenesis), together with the population and biocoenotic ones, in the evolutionary process as a whole.     

Genome size and genome evolution in diploid Triticeae species.

One of the intriguing issues concerning the dynamics of plant genomes is the occurrence of intraspecific variation in nuclear DNA amount. The aim of this work was to assess the ranges of intraspecific, interspecific, and intergeneric variation in nuclear DNA content of diploid species of the tribe Triticeae (Poaceae) and to examine the relation between life form or habitat and genome size. Altogether, 438 plants representing 272 lines that belong to 22 species were analyzed. Nuclear DNA content was estimated by flow cytometry. Very small intraspecific variation in DNA amount was found between lines of Triticeae diploid species collected from different habitats or between different morphs. In contrast to the constancy in nuclear DNA amount at the intraspecific level, there are significant differences in genome size between the various diploid species. Within the genus Aegilops, the 1C DNA amount ranged from 4.84 pg in A. caudata to 7.52 pg in A. sharonensis; among genera, the 1C DNA amount ranged from 4.18 pg in Heteranthelium piliferum to 9.45 pg in Secale montanum. No evidence was found for a smaller genome size in annual, self-pollinating species relative to perennial, cross-pollinating ones. Diploids that grow in the southern part of the group's distribution have larger genomes than those growing in other parts of the distribution. The contrast between the low variation at the intraspecific level and the high variation at the interspecific one suggests that changes in genome size originated in close temporal proximity to the speciation event, i.e., before, during, or immediately after it. The possible effects of sudden changes in genome size on speciation processes are discussed.     

Chromosomal variation and evolution in Lycoris (Amaryllidaceae) I. Intraspecific variation in the karyotype of Lycoris chinensis Traub

In 188 bulbs from five populations of Lycoris chinensis from Anhui province, China, several chromosomal variations have been discovered. Although their frequencies are low, some rearranged chromosomes which are aberrant have been found. The aberrants are: (1) small metacentrics (m′); (2) submetacentrics (sm); (3) subtelocentrics (st); (4) acrocentrics (t); and (5) satellite chromosomes (SAT). All can be easily suspected as being derived from telocentric chromosomes (T type chromosomes). Some individuals having one or more B chromosomes have been found, and intrapopulational variation of B chromosomes in number has also been observed. Because of having B chromosome, L. chinensis has some different chromosome complement numbers: 2n?=?16, 2n?=?16?+?1B, 2n?=?16?+?2B, 2n?=?16?+?3B, and 2n?=?16?+?5B. In addition, a new triploid karyotype composed of 3n?=?24?=?9m?+?11t(2SAT)?+?4T chromosomes has been found. Vegetative propagation is an efficient means of perpetuating the aberrant chromosomes and the triploids.     

Molecular systematics of the Canidae

Despite numerous systematic studies, the relationships among many species within the dog family, Canidae, remain unresolved. Two problems of broad evolutionary significance are the origins of the taxonomically rich canidae fauna of South America and the development in three species of the trenchant heel, a unique meat-cutting blade on the lower first molar. The first problem is of interest because the fossil record provides little evidence for the origins of divergent South American species such as the maned wolf and the bush dog. The second issue is problematic because the trenchant heel, although complex in form, may have evolved independently to assist in the processing of meat. We attempted to resolve these two issues and five other specific taxonomic controversies by phylogenetic analysis of 2,001 base pairs of mitochondrial DNA (mtDNA) sequence data from 23 canidae species. The mtDNA tree topology, coupled with data from the fossil record, and estimates of rates of DNA sequence divergence suggest at least three and possibly four North American invasions of South America. This result implies that an important chapter in the evolution of modern canids remains to be discovered in the fossil record and that the South American canidae endemism is as much the result of extinction outside of South America as it is due to speciation within South America. The origin of the trenchant heel is not well resolved by our data, although the maximum parsimony tree is weakly consistent with a single origin followed by multiple losses of the character in several extant species. A combined analysis of the mtDNA data and published morphological data provides unexpected support for a monophyletic South American canidae clade. However, the homogeneity partition tests indicate significant heterogeneity between the two data sets.     

Chromosomal evolution inEpilobium sect.Epilobium (Onagraceae), III.

A cytological analysis of 22 experimental interspecific hybrids withinEpilobium sect.Epilobium, involving 15 taxa, has indicated that the generalized western North AmericaE. obcordatum has the BB chromosome arrangement. This strengthens greatly our earlier contention that BB is the original chromosome arrangement in the section. The very distinctive northwestern North AmericanE. luteum, only species in the section with cream-colored petals, has the CC arrangement, like the members ofHaussknecht'sAlpinae. This chromosome arrangement apparently originated in the North Pacific region in plants that were of normal stature. The North AmericanE. leptocarpum, which closely resembles the Japanese endemicE. fauriei and, like it, forms gemmae in the axils of the upper leaves, also resembles it in having the CC chromosome arrangement. The AsianE. platystigmatosum has a novel chromosome arrangement derived from BB, which we here designate the EE arrangement.