Diversification of mandarin citrus by hybrid speciation and apomixis

The origin and dispersal of cultivated and wild mandarin and related citrus are poorly understood. Here, comparative genome analysis of 69 new east Asian genomes and other mainland Asian citrus reveals a previously unrecognized wild sexual species native to the Ryukyu Islands: C. ryukyuensis sp. nov. The taxonomic complexity of east Asian mandarins then collapses to a satisfying simplicity, accounting for tachibana, shiikuwasha, and other traditional Ryukyuan mandarin types as homoploid hybrid species formed by combining C. ryukyuensis with various mainland mandarins. These hybrid species reproduce clonally by apomictic seed, a trait shared with oranges, grapefruits, lemons and many cultivated mandarins. We trace the origin of apomixis alleles in citrus to mangshanyeju wild mandarins, which played a central role in citrus domestication via adaptive wild introgression. Our results provide a coherent biogeographic framework for understanding the diversity and domestication of mandarin-type citrus through speciation, admixture, and rapid diffusion of apomictic reproduction.

The Alternative Distribution of Related Earthworms Aporrectodea caliginosa and A. trapezoides (Oligochaeta, Lumbricidae) in Ukraine as a Case of Geographical Parthenogenesis

Geographical parthenogenesis describes phenomenon when parthenogenetic hybrid forms or species have larger distribution areas or higher abundance than their amphimictic parental species, especially in climatically unfavorable conditions. This phenomenon was studied in Ukraine for the pair species of earthworms Aporrectodea сaliginosa (Savigny, 1826) s. l. We found that the hermaphroditic amphimictic A. caliginosa clearly predominates in the northern and western regions, and the apomictic parthenogenetic A. trapezoides (Duges, 1828) in the southern and eastern regions with a continental arid climate. In the sample sets of A. сaliginosa–A. trapezoides group, usually one of the species sharply predominated, and the equality of their abundance was very rare. The reason for this fact is both the alternative geographical distribution and the ability of A. trapezoides to form populations in habitats unfavorable for A. caliginosa. In general, the situation in this group agrees with the classical model of geographic parthenogenesis and confirms the high adaptive potential of apomictic organisms. This fact once again raises the question of non-adaptive reasons for the exclusion of the apomictic reproduction in highly organized animals.

Population genomics reveal apomixis in a novel system: uniclonal female populations dominate the tropical forest herb family, Hanguanaceae (Commelinales)

The abundance of apomixis in tropical plant genera is poorly understood, and this affects the understanding of speciation and evolution. Hanguanaceae is a tropical monogeneric, dioecious plant family. All but two species are solitary herbs with no capability to spread vegetatively. Viable seeds are often produced when males have not been observed. Our aim was to investigate the presence of apomixis in Hanguana. We used reduced representation genomics to study phylogenetics and genetic variability in all populations of Hanguana in Singapore. We measured genome sizes and estimated ploidy levels in 10 species. Almost all taxa tested were genetically uniform (uniclonal) regardless of the extent of their distribution. The distribution of single clones over distinct localities supports our hypothesis of apomictic reproduction. Only one sexually reproducing native species was detected. Triploid and pentaploid states support our hypothesis that the type of apomixis in Hanguana is gametophytic. Population genomics tools offer a quick and cost-effective way of detecting excess clonality and thereby inferring apomixis. In the case of Hanguana, the presence of male plants is a strong indicator of sexual reproduction, whereas genome triplication is indicative of apomictic reproduction.

Secondary origin, hybridization and sexual reproduction in a diploid- tetraploid contact zone of the facultative apomictic orchid Zygopetalum mackayi

ABSTRACT>Mixed-cytotype populations are ideal to understand polyploid establishment and diversification. We used the orchid Zygopetalum mackayi to understand how facultative apomictic reproduction relates to polyploidy. Sexual diploids and facultative apomictic tetraploids occur under distinct niches, with a contact zone where triploids occur. We hypothesized that facultative apomictic reproduction increases the fitness of tetraploids through reproductive interference between cytotypes. We predict patterns of genetic diversity of allopatric tetraploid populations to be significantly different from contact zone populations as a result of dominant apomictic reproduction in the later. We also describe the contact nature of diploids and tetraploids and the role of the intermediate triploids based on patterns of genetic structure within and among pure and mixed-cytotype populations.>We designed eight microsatellite markers and genotyped 155 individuals from six populations resulting in 237 alleles. We described patterns of genetic diversity and structure within and among populations and cytotypes.>Genotypic diversity is similarly high among all populations and cytotypes. Each cytotype emerged as a genetically cluster, combining individuals from different populations. Triploids clustered in an intermediate position between diploids andtetraploids.>We rejected the hypothesis of reproductive interference between cytotypes of Z. mackayi. Patterns of genetic diversity are incongruent with the occurrence of apomict reproduction in tetraploids. Mixed-cytotype populations originate from secondary contact and triploids are hybrids between diploids and tetraploids and act as a reproductive barrier. We suggest polyploidy rather than facultative apomixis explains higher fitness of tetraploids in this species and, therefore, eco-geographical patterns of distribution.

Desarrollo del gametofito femenino de Tagetes patula (Asteraceae)

The genus Tagetes reproduces sexually by seed, but recent morphological and hybridization studies in Tagetes patula suggest an apomictic type of reproduction (seed development without fertilization). In order to determine the sexual or apomictic origen of the embryo, we have studied megasporogenesis, megagametogenesis and the early development of the embryo. Tagetes patula L. has a typical ovule for the family Asteraceae: anatropous, unitegmic, tenuinucellate and with basal placentation. A single hypodermal archesporial cell develops directly as the megaspore mother cell. Megaspogenesis is normal and embryo sac develops from the chalazal megaspore. The embryo sac is of the Polygonum type. Female ray flowers show irregularities in megagametophyte development such as formation of more than eight nuclei, inverted polarity and incomplete differentiation of the megagametophyte cells in mature flowers. These irregularities do not necessarily prove apomictic reproduction in Tagetes patula.

Chromosome doubling in Paspalum notatum var. saure (cultivar Pensacola)

Paspalum notatum is one of the most important native grassland species of southern Brazil, and the main forage source for livestock. The native ecotypes are tetraploid and have apomictic reproduction, which prevents the registration and protection of plant varieties. However, the cultivated variety Paspalum notatum var. saurae (cultivar Pensacola) is diploid and sexual. The objective of this study was to artificially produce polyploid cv. Pensacola plants, to be used as female parents in crosses with male tetraploid and apomictic parents. Seedlings and seeds were immersed in colchicine solutions for different periods. A tetraploid chromosome number was confirmed in three plants. The reproduction mode of the resulting polyploid plants will be assessed; if confirmed to be sexual, they can be used in crosses with native apomictic plants to breed new intraspecific hybrids.

Have necrohormones a role in embryogenesis?

The recognition of apoptosis (programmed cell death) as an accompaniment of normal development, the products released by the protoplasts undergoing self-destruction being utilized by adjacent living cells, stimulates renewed interest in Haberlandt's concept of "necrohormones" playing a role in apomictic reproduction. Recent work on somatic embryogenesis in carrot shows that regular death of certain cells in embryogenic cultures satifies the criteria of apoptosis. Similar observations have been made with embryogenic cultures of <em>Picea abies</em>. Haberlandt's claim that cell death induced by injury adjacent to an ovule in <em>Oenothera</em> could lead to parthenogenesis, despite conflicting evidence from later experimenters, is worthy of reexamination.