THE EVOLUTIONARY HISTORY OF PARTHENOGENETIC CNEMIDOPHORUS LEMNISCATUS (SAURIA,TEIIDAE). I. EVIDENCE FOR A HYBRID ORIGIN

Chromosomes and allozymes were studied from chromosomally distinct unisexual (races B and C) and bisexual (races D and E) populations of the teiid lizard Cnemidophorus lemniscatus, and from selected outgroup taxa (C. murinus, C. nigricolor, Ameiva ameiva, and A. auberi). Karyotyping confirmed the racial identity of individuals and showed that the chromosomal composition of populations at specific localities has remained the same for 20 years. All individuals of both unisexual populations were heterozygous for a pericentric inversion that distinguishes D and E bisexuals. Also, the unisexuals were all heterozygous for 8 of 11 protein loci for which D and E were fixed or nearly fixed for different alleles. Most of these alleles represent derived states relative to the other Cnemidophorus and Ameiva analyzed, and the fixed heterozygote condition at these nine markers provides unequivocal support for the hypothesis that the unisexual C. lemniscatus arose by hybridization between ancestors genetically similar to extant D and E populations. At the remaining three loci for which D and E show fixed differences, the unisexuals were homozygous rather than heterozygous. This suggests that either (1) allozymes have been lost by mutations to null, silent, or convergent mobility states, (2) ancestral genotypes were similar to but not identical with the extant D and E races, and/or (3) limited recombination may occur between unisexual genomes. Allozyme-based genetic distances between D and E were large, suggesting that bisexual races D and E are genetically isolated; each race should be accorded full species status. This conclusion is supported by the absence of any clear biochemical evidence for their monophyly with respect to the other Cnemidophorus examined. Cladistic analyses of 17 phylogenetically informative loci revealed two equally parsimonious shortest trees, one supporting monophyly and the other paraphyly of the C. lemniscatus complex. Further testing of the monophyly of C. lemniscatus requires additional data. With the present study, the evidence that all parthenogenetic Cnemidophorus are of hybrid origin is complete.     

THE HYBRID ORIGIN OF ASTER ASCENDENS (ASTERACEAE)

Aster ascendens Lindl. is a rhizomatous perennial of the Great Basin region of the western United States, with a chromosome base number of x = 13. Populations in the southern and western parts of the range are mainly diploid (n = 13) and those in the north and east are mainly tetraploid (n = 26). This species is postulated to be an amphiploid which has resulted from hybridization between the western Cordilleran A. occidentalis (T. & G.) Hook, and the Great Plains species A. falcatus Lindl. Aster ascendens is intermediate between these two species in most morphological traits, but resembles A. occidentalis in ray color and general growth habit, and A. falcatus in the shape of the outer phyllaries. The karyotype of A. ascendens also exhibits a combination of the features found in A. occidentalis and A. falcatus karyotypes. Of the specimens collected for this study, 13% were putative backcrosses of A. ascendens to one of its parent taxa or to a closely related species; this suggests that there is potentially continuing genetic interchange between the species groups that gave rise to A. ascendens.     

A HYPOTHESIS ON THE EVOLUTIONARY ORIGIN OF THE SQUAMOSAL BONE IN THE THERIAN MAMMALS

Theatheriansandthetherianshavebothinheritedthesquamosa1fromthecyno-dontidancestry,theprimitivepropertyofthesquamosa1intheatherianlineagehavebeenmaintainedfromthebeginning(theLateTriassic,Morganucodontids)totheend(Cretaceous,Triconodontids).Inthetherianlineage,thelowertherianshavethesamesquamosalastheatherians,however,thehighertherianshaveapeculiarsquamosal,so,thechangeofthepropertyofthesquamosaImaytakeplacelntheevolutionaryphasefromthelowertherianmammals(Symmetrodonta,Eupan-totheria,Multitu…     

寒武纪大附肢节肢动物的起源和进化机制分析

在节肢动物早期演化阶段 ,前附肢的演化是提高动物感觉和捕食能力的重要事件。化石和发育生物学实验结果表明节肢动物前附肢 (即口前附肢 )的原型为短棒形触角 ,在此基础上分别演化为长鞭形触角和腿肢形大附肢 ;大附肢与螯肢形态相似 ,为同源构造 ,因此寒武纪大附肢类群可能是现生螯肢类的干支类群。文中结合发育生物学研究进展分析节肢动物演化早期大附肢产生可能的分子进化机制。认为腿肢形大附肢的发生可能与Hox基因或其它未知发育调控因子参与触角或者附肢原型的发育有关 ,从而改变原有前附肢发育中Dll基因、dac基因和hth基因间特异的相互作用 ,导致前附肢获得腿肢特异性发育程序。     

EVOLUTIONARY ORIGIN OF THE CYCLORRHAPHA (DIPTERA): TEST OF ALTERNATIVE MORPHOLOGICAL HYPOTHESES

Abstract— The higher flies, infraorder Cyclorrhapha [=Muscomorpha (McAlpine, 1989)], have undergone enormous radiation since the Cretaceous (∼100 Myr). Rapid morphological evolution in cyclorrhaphans has made their phylogenetic placement with respect to more primitive clades a long-standing problem in dipteran systematics. Of the two most plausible hypotheses, one treats the Cyclorrhapha as sister group to the orthorrhaphous superfamily Empidoidea [=Empidiformia (Hennig, 1948), Orthogenya (Brauer, 1883)], while the other places them within the empidoids. The debate over cyclorrhaphan origin has heretofore focused on homology interpretations for a few character systems, particularly the male genitalia. We provide the first attempt to assemble and quantify all of the available morphological evidence. By cladistic analysis of these data under alternative codings of genitalic features reflecting opposing homology theories, and then excluding these features altogether, we sought to judge which genitalic theory is better supported by the evidence as a whole, and how much the debate matters to resolving cyclorrhaphan origins. Using the analog of a factorial design, we also measured the effect of alternative transformation series in several other controversial characters, of outgroup choice and of successive weighting. Under all manipulations, including both genitalic codings, the Cyclorrhapha originate within the Empidoidea, near the family Atelestidae. However, trees in which the Empidoidea are constrained to be monophyletic are only 1-6 steps longer (out of ∼150), a fit not significantly worse under a permutation test for monophyly. Adult morphological data may not suffice to settle either the placement of Cyclorrhapha or the debate over genitalic homology. Moreover, the issue of genitalic homology does not appear critical to that of cyclorrhaphan origin.     

DIVERGENCE OF MEIOTIC DRIVE-SUPPRESSION SYSTEMS AS AN EXPLANATION FOR SEX-BIASED HYBRID STERILITY AND INVIABILITY

Two empirical generalizations about speciation remain unexplained: the tendency of the heterogametic sex to be sterile or inviable in F₁ hybrids (Haldane's rule), and the tendency of the X chromosome to harbor the genetic elements that cause this sex bias in hybrid fitness. I suggest that divergence of meiotic drive systems on the sex chromosomes can explain these observations. The theory follows from two simple facts. First, sex chromosomes are particularly susceptible to the forces of meiotic drive. Second, divergence of meiotic drive systems can cause hybrid sterility and in viability. The main objection to the theory is that meiotic drive is apparently rare, whereas the observed pattern of hybrid fitness is widespread. I answer this objection by showing that divergence of meiotic drive systems can explain the two generalizations even if large departures from Mendelian segregation are rarely observed.     

RECONSTRUCTING THE COMPLEX EVOLUTIONARY ORIGIN OF WILD ALLOPOLYPLOID TOBACCOS (NICOTIANA SECTION SUAVEOLENTES)

Nicotiana (Solanaceae) provides an ideal system for understanding polyploidization, a pervasive and powerful evolutionary force in plants, as this genus contains several groups of allotetraploids that formed at different times from different diploid progenitors. However, the parental lineages of the largest group of allotetraploids, Nicotiana section Suaveolentes, have been problematic to identify. Using data from four regions of three low‐copy nuclear genes, nuclear ribosomal DNA, and regions of the plastid genome, we have reconstructed the evolutionary origin of sect. Suaveolentes and identified the most likely diploid progenitors by using a combination of gene trees and network approaches to uncover the most strongly supported evidence of species relationships. Our analyses best support a scenario where a member of the sect. Sylvestres lineage acted as the paternal progenitor and a member of either sect. Petunioides or sect. Noctiflorae that also contained introgressed DNA from the other, or a hypothetical hybrid species between these two sections, was the maternal progenitor. Nicotiana exemplifies many of the factors that can complicate the reconstruction of polyploid evolutionary history and highlights how reticulate evolution at the diploid level can add even greater complexity to allopolyploid genomes.     

EVOLUTIONARY SORTING OF CHARACTERS IN A HYBRID SWARM. I: DIRECTION OF SLOPE

Quercus Douglasii and Quercus turbinella subsp. californica intergrade in California in a belt about 250 miles long. In the area of each parental taxon and near Lebec, in the middle of the zone where hybrids are common and where the gene pool is about evenly divided as to origin, the phenotypes were scored on a six-point scale to establish hybrid indices. At Lebec populations on slopes facing at intervals of one-eighth the distance around the compass were scored. The composition of each population was heterogeneous but restricted markedly according to exposure to the sun. Plants on northeast-facing slopes approached but did not equal the high score of Quercus Douglasii in Madera County; those on southwest-facing slopes approached but did not equal the extreme low score of subsp. californica near Victorville. Evolutionary sorting must be very rapid, for it is correlated with slopes of the same hills, and the hybrid swarm must be very sensitive to the environment.     

MOLECULAR TESTS OF THE HYPOTHESIZED HYBRID ORIGIN OF TWO DIPLOID HELIANTHUS SPECIES (ASTERACEAE)

Enzyme electrophoresis and restriction-fragment analysis of chloroplast DNA (cpDNA) and nuclear ribosomal DNA (rDNA) were used to test the hypothesis that both Helianthus neglectus and H. paradoxus are stabilized hybrid derivatives of H. annuus and H. petiolaris. The four species are annuals, diploid outcrossers, and have the same chromosome number. Helianthus annuus and H. petiolaris had the same allele in highest frequency for 16 of the 18 isozyme loci examined and had different majority alleles for only 6-Pgd3 and Pgi2. The two species had divergent rDNAs that could be distinguished by seven diagnostic restriction site mutations and three length mutations, and their cpDNAs could be differentiated by three diagnostic restriction site mutations. The alleles observed in H. neglectus were not a combination of those observed in H. annuus and H. petiolaris. Although H. neglectus had only one unique allele, it possessed none of the three alleles specific to H. annuus. In contrast, it had four of the seven alleles specific to H. petiolaris. Furthermore, H. neglectus had the same rDNA type as H. petiolaris and had the same cpDNA as that found in two populations of H. petiolaris ssp. fallax. These data allowed us to speculate that H. neglectus may be a recent derivative of H. petiolaris ssp. fallax, rather than a stabilized hybrid derivative as originally proposed. In contrast, H. paradoxus combined the alleles of H. annuus and H. petiolaris and had no unique alleles. At Adh2, H. paradoxus was monomorphic for an allele found only in H. petiolaris ssp. fallax, whereas at 6-Pgd3 and Pgi2, it was monomorphic for high frequency H. annuus alleles. Furthermore, H. paradoxus combined the rDNA repeat types of both proposed parents and had the chloroplast genome of H. annuus. These data provide compelling evidence that H. paradoxus, in contrast to H. neglectus, was derived via hybridization.     

EVOLUTIONARY ORIGIN OF A PARTHENOFORM,THE AMAZON MOLLY POECILIA FORMOSA,ON THE BASIS OF A MOLECULAR GENEALOGY

The appearance of vertebrate species that reproduce without genetic recombination has been explained by their origin from a rare hybridization event between members of two distantly related species. For the first recognized vertebrate unisexual, the Amazon molly Poecilia formosa, mostly morphological and biochemical genetic information has been available so far with respect to its evolutionary origin. DNA sequence analyses of transcribed portions of the genome (tyrosine kinase proto-oncogenes) demonstrated its hybrid state unequivocally. Both alleles can be traced in a DNA sequence-based phylogenetic tree to extant species that represent the parental species or that are closely related to the corresponding extinct forms, namely P. mexicana limantouri and a so far taxonomically ill-defined north Mexican subspecies of the P. latipinna/P. velifera complex. A rough estimate from the mutation rates dates the hybridization event further back than would have been predicted on the basis of “Muller's ratchet” for an ecologically successful species.     

UNRAVELING CRYPTIC RETICULATE RELATIONSHIPS AND THE ORIGIN OF ORPHAN HYBRID DISJUNCT POPULATIONS IN NARCISSUS

Evolutionary consequences of natural hybridization between species may vary so drastically depending on spatial, genetic, and ecological factors that multiple approaches are required to uncover them. To unravel the evolutionary history of a controversial hybrid (Narcissus×perezlarae), here we use four approaches: DNA sequences from five regions (four organellar, one nuclear), cytological studies (chromosome counts and genome size), crossing experiments, and niche modeling. We conclude that (1) it actually consists of two different hybrid taxa, N.×perezlarae s.s. (N. cavanillesii×N. miniatus) and N.×alentejanus (N. cavanillesii × N. serotinus); (2) both have been formed several times independently, that is, polytopically; (3) N. cavanillesii was the mother progenitor in most hybridization events. We also address the origin of orphan hybrid populations of N.×perezlarae in eastern Spain, hundreds of kilometers away from N. cavanillesii. Although long‐distance dispersal of already formed hybrids cannot be completely rejected, extirpation of N. cavanillesii via demographic competition is a more likely explanation. Low‐reproductive barriers to fertilization by foreign pollen in N. cavanillesii, molecular footprints of the former presence of this species in the area, active asexual propagation by bulbs in N.×perezlarae, and overlapping ecological niches are consistent with the extirpation scenario.     

THE ORIGIN AND EVOLUTIONARY RELATIONSHIPS OF ‘HUAUZONTLE’ (CHENOPODIUM NUTTALLIAE SAFFORD), DOMESTICATED CHENOPOD OF MEXICO

A New World assemblage of tetraploid Chenopodium species (section Chenopodium, subsection Cellulata) includes two domesticates, C. quinoa of Andean South America and C. nuttalliae of Mexico. Both have been combined into a single species and the Mexican form has been considered as a possible derivative of C. quinoa. The domesticates and related, sympatric weed forms, C. berlandieri of North America and C. hircinum of the Andes, were examined for variation in morphological and biochemical characteristics and also were included in a program of artificial hybridization. Results indicate that the domesticated forms are more closely related to their sympatric weeds than to each other. The Mexican cultigen is placed as a subspecies of C. berlandieri, the taxon from which it most likely evolved under human selection in North America. Possible origins for the Andean weed-crop complex are considered. Southward migration of a North American tetraploid appears to be more likely than independent allotetraploidy in South America. Of the North American tetraploids examined, C. berlandieri var. zschackei of the western U.S. shows closest affinities to the Andean complex.