Dioecious Genus Research Articles

The complete plastid genome sequence of Ilex suaveolens (H. Lév.) Loes, the most abundant medicinal holly in Mount Huangshan

Holly (Ilex L.) is a woody dioecious genus cultivated as pharmaceutical, ornamentals, and industrial materials. Ilex suaveolens (H. Lév.) Loes is an endemic medicinal holly with a predominant distribution in Mount Huangshan, China. In the present work, the complete plastid genome of I. suaveolens was de novo sequenced by high-throughput sequencing technology. The newly-assembled plastid genome holds 37.6% of the overall GC content and a length of 157,857 bp, comprising a large single-copy (LSC, 87,255 bp), a small single-copy (SSC, 18,398 bp), and a pair of inverted repeat (IRs, 26,102 bp) regions. The plastid genome annotation suggested the presence of a total of 89 protein-encoding genes, 37 transfer RNA (tRNA) genes, and eight ribosomal RNA (rRNA) genes. The plastome-mediated phylogenetic topology revealed that I. suaveolens clustered together with I. szechwanenesis and I. viridis in the same clade, and a strong relationship between clades and biogeography was found. These data contribute to the understanding of genetic diversity and conservation study of Ilex in Mount Huangshan.

Comparative Survey of Morphological Variations and Plastid Genome Sequencing Reveals Phylogenetic Divergence between Four Endemic Ilex Species

Holly (Ilex L.), from the monogeneric Aquifoliaceae, is a woody dioecious genus cultivated as pharmaceutical and culinary plants, ornamentals, and industrial materials. With distinctive leaf morphology and growth habitats, but uniform reproductive organs (flowers and fruits), the evolutionary relationships of Ilex remain an enigma. To date, few contrast analyses have been conducted on morphology and molecular patterns in Ilex. Here, the different phenotypic traits of four endemic Ilex species (I. latifolia, I. suaveolens, I. viridis, and I. micrococca) on Mount Huangshan, China, were surveyed through an anatomic assay and DNA image cytometry, showing the unspecified link between the examined morphology and the estimated nuclear genome size. Concurrently, the newly-assembled plastid genomes in four Ilex have lengths ranging from 157,601 bp to 157,857 bp, containing a large single-copy (LSC, 87,020–87,255 bp), a small single-copy (SSC, 18,394–18,434 bp), and a pair of inverted repeats (IRs, 26,065–26,102 bp) regions. The plastid genome annotation suggested the presence of numerable protein-encoding genes (89–95), transfer RNA (tRNA) genes (37–40), and ribosomal RNA (rRNA) genes (8). A comprehensive comparison of plastomes within eight Ilex implicated the conserved features in coding regions, but variability in the junctions of IRs/SSC and the divergent hotspot regions potentially used as the DNA marker. The Ilex topology of phylogenies revealed the incongruence with the traditional taxonomy, whereas it informed a strong association between clades and geographic distribution. Our work herein provided novel insight into the variations in the morphology and phylogeography in Aquifoliaceae. These data contribute to the understanding of genetic diversity and conservation in the medicinal Ilex of Mount Huangshan.

Plastome phylogenomics of the East Asian endemic genus Dobinea

Dobinea is a dioecious genus endemic to East Asia that consists of two extant species: Dobinea delavayi and Dobinea vulgaris. Although the genus is morphologically distinct, its phylogenetic position remains controversial. In this study, we investigated the phylogenetic relationships between Dobinea and related taxa by sequencing the whole plastome DNA sequences for both extant species of Dobinea and comparing them to published plastomes within Sapindales. The complete plastomes of D. vulgaris and D. delavayi were 160,683 and 160, 154 base pairs (bp) in length, including a pair of inverted repeat regions (IRs, 26,889 and 26,759 bp) divided by the large single-copy region (LSC, 87,962 and 87,555 bp) and small single-copy region (SSC, 18,943 and 19,081 bp), and identically encoded 113 unique genes (79 protein-coding genes, 30 tRNAs, and 4 rRNA genes). Plastid phylogenomic analyses showed that Dobinea was a well-supported monophyletic unit and sister to the clade including tribes Anacardieae and Rhoideae, which suggests that Dobinea is a member of Anacardiaceae. In addition, molecular dating inferred D. delavayi and D. vulgaris diverged approximately 10.76 Ma, suggesting the divergence between these two species may have been driven by the intensification of the Asian summer monsoon and the establishment of distinct monsoon regimes in East Asia.

Phylogeny and biogeography of the hollies (Ilex L., Aquifoliaceae)

AbstractThe holly genus, Ilex L., in the monogeneric Aquifoliaceae, is the largest woody dioecious genus (>664 spp.), with a near‐cosmopolitan distribution in mesic environments. We constructed a phylogeny based on two nuclear genes, representing 177 species spread across the geographical range, and dated using macrofossil records. The five main clades had a common ancestor in the early Eocene, much earlier than previously suggested. Ilex originated in subtropical Asia and extant clades colonized South America by 30 Ma, North America by 23 Ma, Australia by 8 Ma, Europe by 6 Ma, and Africa by 4 Ma. South and North America were colonized multiple times. Ilex also reached Hawaii (10 Ma) and other oceanic islands. Macrofossil and pollen records show the genus has tracked mesic climates through time and space, and had a wider distribution before late Miocene global cooling. Our phylogeny provides a framework for studies in comparative ecology and evolution.

Novelty upon novelty visualized by rotational scanning electron micrographs (rSEM): Laboulbeniales on the millipede order Chordeumatida.

Laboulbeniales are highly specific ectoparasitic fungi of arthropods (insects, millipedes, and arachnids). The first Laboulbeniales parasitizing the millipede order Chordeumatida (Diplopoda) were discovered and described as a new dioecious genus of Laboulbeniales, Thaxterimyces, to accommodate the new species T. baliensis. Also the millipede host is a new species and is described as Metopidiothrix sheari. This is the first time Laboulbeniales fungus and its millipede host are described as new together. Males of Metopidiothrix have the most extensive secondary sexual modifications in the entire class Diplopoda. Although nothing is known about the function of these modifications, the unique pattern of Laboulbeniales infection in the new millipede species is obviously related to host sexual behavior. Rotational Scanning Electron Micrographs (rSEM) are used to create a 3D comprehensive model to examine the fungal-host interaction, a more advanced visualization of the ectoparasitic fungus on its host. Laboulbeniales diversity on millipedes is still understudied, and a consistent effort is needed to unveil and understand the extent and diversity of this biological interaction. Due to their minute size and apparently non-detrimental effect on their hosts, Laboulbeniales in general have been largely ignored by mycologists and neglected by generations of entomologists. As a result a significant component of global biodiversity has been strongly underestimated, and a wealth of new discoveries is still to be made both in the field and in existing museum collections.

Synflorescence morphology of grasses with reduced terminal inflorescences: a case study of Jouvea (Cynodonteae, Chloridoideae, Poaceae)

Jouvea is a dioecious genus of grasses with two species. Molecular phylogenetic analyses place it in tribe Cynodonteae, closely related to subtribes Hilariinae and Scleropogoninae, and the genera Allolepis and Sohnsia. The staminate inflorescence in Jouvea is an ordinary grass spike of spikelets, but the pistillate inflorescence is represented by a single spikelet which lacks glumes, has a thick and cylindrical rachilla, lemmas forming a tube that enclose the palea and pistil, stigmas protruding from an apical pore of this tube, and does not have lodicules. Within Cynodonteae, inflorescences reduced to a single spikelet also occur in subtribe Monanthochloinae, in which the small number of flowers of the terminal inflorescence is compensated by an increase in the number of lateral floriferous shoots. We here describe the floriferous shoot system (synflorescence) of Jouvea and compare it to other cynodonteae grasses with reduced inflorescences. Jouvea species display a high number of lateral shoots (trophotagma enrichment axes) growing from the medial and distal zones of the synflorescences. These shoots have prophyllar origins and form clusters of lateral inflorescences. The elevated number of trophotagma enrichment axes of Jouvea may be associated with the extreme reduction in the pistillate terminal inflorescence. In addition, the increase in number of spikelets by the development of prophyllar branches is a unique strategy within tribe Cynodonteae.

Chloroplast genome structure in Ilex (Aquifoliaceae)

Aquifoliaceae is the largest family in the campanulid order Aquifoliales. It consists of a single genus, Ilex, the hollies, which is the largest woody dioecious genus in the angiosperms. Most species are in East Asia or South America. The taxonomy and evolutionary history remain unclear due to the lack of a robust species-level phylogeny. We produced the first complete chloroplast genomes in this family, including seven Ilex species, by Illumina sequencing of long-range PCR products and subsequent reference-guided de novo assembly. These genomes have a typical bicyclic structure with a conserved genome arrangement and moderate divergence. The total length is 157,741 bp and there is one large single-copy region (LSC) with 87,109 bp, one small single-copy with 18,436 bp, and a pair of inverted repeat regions (IR) with 52,196 bp. A total of 144 genes were identified, including 96 protein-coding genes, 40 tRNA and 8 rRNA. Thirty-four repetitive sequences were identified in Ilex pubescens, with lengths >14 bp and identity >90%, and 11 divergence hotspot regions that could be targeted for phylogenetic markers. This study will contribute to improved resolution of deep branches of the Ilex phylogeny and facilitate identification of Ilex species.

Floral trait evolution associated with shifts between insect and wind pollination in the dioecious genus Leucadendron (Proteaceae).

Transitions between animal and wind pollination have occurred in many lineages and have been linked to various floral modifications, but these have seldom been assessed in a phylogenetic framework. In the dioecious genus Leucadendron (Proteaceae), transitions from insect to wind pollination have occurred at least four times. Using analyses that controlled for relatedness among Leucadendron species, we investigated how these transitions shaped the evolution of floral structural and signaling traits, including the degree of sexual dimorphism in these traits. Pollen grains of wind-pollinated species were found to be smaller, more numerous, and dispersed more efficiently in wind than were those of insect-pollinated species. Wind-pollinated species also exhibited a reduction in spectral contrast between showy subtending leaves and background foliage, reduced volatile emissions, and a greater degree of sexual dimorphism in color and scent. Uniovulate flowers and inflorescence condensation are conserved ancestral features in Leucadendron and likely served as exaptations in shifts to wind pollination. These results offer insights into the key modifications of male and female floral traits involved in transitions between insect and wind pollination.

Evidence for a Common Origin of Homomorphic and Heteromorphic Sex Chromosomes in Distinct Spinacia Species.

The dioecious genus Spinacia is thought to include two wild relatives (S. turkestanica Ilj. and S. tetrandra Stev.) of cultivated spinach (S. oleracea L.). In this study, nuclear and chloroplast sequences from 21 accessions of Spinacia germplasm and six spinach cultivars or lines were subjected to phylogenetic analysis to define the relationships among the three species. Maximum-likelihood sequence analysis suggested that the Spinacia plant samples could be classified into two monophyletic groups (Group 1 and Group 2): Group 1 consisted of all accessions, cultivars, and lines of S. oleracea L. and S. turkestanica Ilj. and two of five S. tetrandra Stev. accessions, whereas Group 2 was composed of the three remaining S. tetrandra Stev. accessions. By using flow cytometry, we detected a distinct difference in nuclear genome size between the groups. Group 2 also was characterized by a sexual dimorphism in inflorescence structure, which was not observed in Group 1. Interspecific crosses between the groups produced hybrids with drastically reduced pollen fertility and showed that the male is the heterogametic sex (XY) in Group 2, as is the case in S. oleracea L. (Group 1). Cytogenetic and DNA marker analyses suggested that Group 1 and Group 2 have homomorphic and heteromorphic sex chromosome pairs (XY), respectively, and that the sex chromosome pairs of the two groups evolved from a common ancestral pair. Our data suggest that the Spinacia genus may serve as a good model for investigation of evolutionary mechanisms underlying the emergence of heteromorphic sex chromosome pairs from ancestral homomorphic pairs.

Ndido de las poblaciones ibéricas de Mercurialis annua?

Pannell, J.R ., Santos del Blanco, L. 2014 . Evolution of combined versus separate sexes: What have we learnt from the Iberian populations of Mercurialis annua ?. Ecosistemas 23(3): 13-22. Doi.: 10.7818/ECOS.2014.23-3.03 Most plant species are hermaphrodites, with both male and female functions performed by the same individuals. However, separate sexes (dioecy) have evolved on numerous independent occasions, probably either in response to selection for inbreeding avoidance, or because it pays individuals to specialize in one gender or the other. Although the evolution of dioecy from hermaphroditism tends to be thought of as a one-way path, dioecy has broken down to yield hermaphroditic populations on several occasions. One such case is found in the mainly dioecious genus Mercurialis ( Euphorbiaceae ). In the species complex M. annua , diploids are dioecious, but polyploid populations are variously monoecious or androdioecious (where males co-exist with functional hermaphrodites). This species complex offers rich material for addressing questions concerning the evolution and ecology of combined versus separate sexes, the evolution of secondary sexual dimorphism, which likely contributes to the stability of dioecy in the genus, and the evolution and genetics of sex determination and sex chromosomes. The species also offers itself as a valuable teaching tool for addressing topics ranging from sex-ratio selection to inter-sexual competition.

Floral structure and development in the monoecious palm Gaussia attenuata (Arecaceae; Arecoideae).

Sexual dimorphism, at both the flower and plant level, is widespread in the palm family (Arecaceae), in contrast to the situation in angiosperms as a whole. The tribe Chamaedoreeae is of special interest for studies of the evolution of sexual expression since dioecy appears to have evolved independently twice in this group from a monoecious ancestor. In order to understand the underlying evolutionary pathways, it is important to obtain detailed information on flower structure and development in each of the main clades. Dissection and light and scanning electron microscopy were performed on developing flowers of Gaussia attenuata, a neotropical species belonging to one of the three monoecious genera of the tribe. Like species of the other monoecious genera of the Chamaedoreeae (namely Hyophorbe and Synechanthus), G. attenuata produces a bisexual flower cluster known as an acervulus, consisting of a row of male flowers with a basal female flower. Whereas the sterile androecium of female flowers terminated its development at an early stage of floral ontogeny, the pistillode of male flowers was large in size but with no recognizable ovule, developing for a longer period of time. Conspicuous nectary differentiation in the pistillode suggested a possible role in pollinator attraction. Gaussia attenuata displays a number of floral characters that are likely to be ancestral to the tribe, notably the acervulus flower cluster, which is conserved in the other monoecious genera and also (albeit in a unisexual male form) in the dioecious genera (Wendlandiella and a few species of Chamaedorea). Comparison with earlier data from other genera suggests that large nectariferous pistillodes and early arrest in staminode development might also be regarded as ancestral characters in this tribe.

Experimental Evaluation of Insect Pollination versus Wind Pollination in Leucadendron (Proteaceae)

Premise of research. Evolutionary transitions from insect to wind pollination are thought to have occurred many times during the angiosperm radiation. On the basis of floral features, several such transitions have been postulated for Leucadendron (Proteaceae), a dioecious genus of 96 taxa confined almost entirely to the Cape Floristic Region. To confirm whether these transitions took place, we carried out experimental tests for wind pollination and sampled insect flower visitors in 15 Leucadendron species, representing six clades in the genus. Methodology. In three clades in which an insect-wind shift has been inferred, we sampled species with traits suggestive of either wind or insect pollination. Plants were bagged with fine-mesh exclusion bags that excluded insects but allowed the passage of airborne pollen. Insects visiting female inflorescences were collected for identification and analysis of their pollen loads. Pivotal results. We found that insect exclusion had little effect on seed set of five species conforming to the wind pollination syndrome (L. rubrum, L. salicifolium, L. dubium, L. coniferum, and L. teretifolium), while seed set was strongly reduced by insect exclusion in the remaining 10 species conforming to an insect pollination syndrome. The most common pollinator of the insect-pollinated species was the nitidulid beetle Pria cinerascens. Conclusions. This study provides experimental verification of contrasting insect and wind pollination systems in Leucadendron and will thus enable formal phylogenetic tests of adaptive changes in floral traits, such as color and scent, that may be associated with pollination system transitions in this diverse lineage.

The sex‐linked region in Populus tremuloides Turesson 141 corresponds to a pericentromeric region of about two million base pairs on P. trichocarpa chromosome 19

In the dioecious genus Populus, sex determination has been located to chromosome 19. However, despite a high degree of genome collinearity, various Populus species seem to differ with regard to the location of the sex-determining region on the respective chromosome and the apparent heterogametic sex. In this study, the boundaries of the recombination-suppressed, sex-linked region of the male P. tremuloides clone Turesson 141 were localised by genetic mapping using new SNP and InDel markers. The respective region seems to be located in a pericentromeric position. The corresponding P. trichocarpa genome region spans about two million bp and comprises 65 gene loci, which were bioinformatically evaluated for their potential as candidate genes for sex determination. Three putative transcription factor genes and four genes that are potentially involved in flower development processes, e.g. meristem transition from the vegetative to the reproductive phase, were identified. Populus tremuloides sequence data of the sex-linked region is required for a final search for candidate genes.

The effect of 2n gametes on sex ratios in Actinidia

Sex can sometimes lead to complications. In some crops, 2n gametes have been exploited by plant breeders to transfer genetic variation between taxa of different ploidy levels. However, their role and use in dioecious genera have received relatively little attention. In the dioecious genus Actinidia (kiwifruit), seedling populations usually segregate equally for females and males as sex is determined by an XX female/XY male system. While fertilization involving 2n egg cells is not expected to affect the sex ratios of progenies, fertilization involving 2n pollen is likely to produce progenies with excess males. The extent of sex ratio distortion will depend on the relative contributions of first and second division restitution, and the frequency and location of cross-overs in meiosis. In this study, seedlings recovered from crosses between females of hexaploid Actinidia deliciosa and males of two diploid species, Actinidia chinensis and Actinidia eriantha, included a proportion of pentaploid hybrids presumably derived from fertilization involving 2n pollen. Most of these pentaploids were male, and a proportion of them were likely to be carrying two Y chromosomes. If used as parents in further crosses, males with multiple Y chromosomes are likely to cause distorted sex ratios in their immediate progenies. In dioecious genera such as Actinidia, the effects on sex ratios of different mechanisms of ploidy change need to be taken into account when considering the evolution of polyploidy and the design of breeding strategies involving ploidy manipulation.

Taxonomic Revision of the Neotropical Genus <I>Lacunaria</I> (Quiinaceae / Ochnaceae s. l.)

Abstract A taxonomic revision of the Neotropical dioecious genus Lacunaria (Quiinaceae / Ochnaceae s. l.) is presented including keys to the species and subspecies, descriptions, illustrations, nomenclature, typification, and geographical distribution of each species. Seven species are recognized within the genus, one subspecies, L. jenmanii subsp. subsessilis, is described newly, and the status of Lacunaria decastyla has been changed, transferring it to Lacunaria crenata subsp. decastyla. Leaf pubescence, number of secondary veins, and petal, stamen, and carpel number, although all quite variable, are the most important characters to distinguish the species.

Causes of secondary sexual differences in plants — Evidence from extreme leaf dimorphism in Leucadendron (Proteaceae)

In extreme cases leaves in male plants of the dioecious genus Leucadendron (Proteaceae) are up to an order of magnitude smaller than female leaves. This secondary sexual dimorphism (SSD) in leaf size has previously been suggested to be due to intra-male sexual selection, leading to an increase in male allocation to reproduction in dimorphic species. After critically evaluating previous data provided to support this hypothesis, I suggest on both theoretical grounds and on re-analysis that this argument is unlikely and unsupported. Leaf size dimorphism could theoretically evolve directly due to disruptive ecological selection between genders, leading to niche dimorphism either within or between habitats. I test this ecological causation hypothesis by providing data on specific leaf area (sla) and water use efficiency (δ 13C) of leaves from males and females of several Leucadendron species. Results confirm the expectation of minimal gender differences. I argue that leaf dimorphism is a consequence of selection on flower size and architecture.

Canopy seed storage is associated with sexual dimorphism in the woody dioecious genus Leucadendron

Summary 1. Species of the dioecious South African plant genus Leucadendron vary strikingly in their degree of sexual dimorphism, with males of dimorphic species typically possessing more and thinner branches and smaller leaves than the corresponding females. 2. Many Leucadendron species are serotinous, with females of serotinous species maintaining seeds in transpiring cones for a number of years. Since cones must be kept alive, serotinous females presumably incur higher costs in terms of water loss than non‐serotinous females and, particularly, males. Given that branching architecture is known to affect water conductance, with less resistance in less highly branched plants, we predicted a positive correlation between the degree of serotiny in a species and sexual dimorphism in branching. 3. We tested the effect of serotiny and several other factors on the degree of sexual dimorphism in Leucadendron by conducting a comparative analysis of variation across 49 species using phylogenetic regression. 4. There was a strong positive association between the degree of serotiny and sexual dimorphism. This association is consistent with the idea that the costs of maintaining cones may select females to be less highly branched than males. 5. Synthesis. Serotiny is a strategy of maternal care that contributes to offspring survival after seed maturation. We found that, in Leucadendron, the evolution of increased serotiny corresponds with the evolution of more marked sexual dimorphism. This study exposes a hitherto unidentified association between sexual dimorphism and maternal care in plants and points to a possible additional reproductive burden carried by serotinous plants compared with those that release their seeds at maturity.

The endemic genus Pancheria (Cunoniaceae) in New Caledonia: notes on morphology and the description of three new species

A generic description of the dioecious genus Pancheria (Cunoniaceae) from New Caledonia is accompanied by notes on the morphology of the leaves, stipules, inflorescences, flowers and fruits. Descriptions, illustrations, distribution maps and conservation assessments are provided for each of three new species. Pancheria mcphersonii occurs on Mts Panie, Colnett and Ignambi in north-eastern Grande Terre, in dense low forest and montane scrub on soils derived from mica-schist (non-ultramafic). Pancheria dognyensis grows on the Plateau de Dogny in central New Caledonia and on Roche Ouaieme and Mt Mandjelia in the north-east, in forest and especially at the forest edge, also on non-ultramafic substrates. Pancheria ajiearoana is found in central Grande Terre, in scrub on ultramafic soils. Because all three species are restricted to a small number of areas at high elevation, climate change is one of the threats that justifies giving each of them a preliminary conservation status of Endangered; as global temperatures increase, the area of habitat with a microclimate suitable for these species is likely to decrease.

SKEWED SEX RATIOS, POLLEN LIMITATION, AND REPRODUCTIVE FAILURE IN THE DIOECIOUS SEAGRASS PHYLLOSPADIX.

Dioecious mating systems in angiosperms, in which individuals are unisexual, require that pollen disperses between male and female individuals and may cause dioecious species to be vulnerable to reduced reproductive success via pollen limitation. This potential cost has rarely been documented, in part because of a dearth of studies investigating abiotic pollen dispersal, which tends to be associated with dioecy. I document widespread variation in reproductive success in an ecologically important, habitat-forming group, the seagrasses (marine angiosperms), using two species of the dioecious genus Phyllospadix. At multiple sites in the state of Washington, USA, I demonstrate pervasive male rarity (all sites <24% male by area). Male abundance is shown to predict female reproductive success at two scales. Within sites, the ratio of maturing seeds to total ovules declines rapidly with increasing distance to the nearest male, while among sites, mean seed : ovule ratios scale positively with male abundance. At some sites, less than 1 in 100 ovules mature into seeds. A field experiment conducted at a high pollen availability site shows that manipulating pollen availability could produce the range of seed sets observed across sites, but pollen limitation was not definitively demonstrated. Overall my results are consistent with pollen limitation in Phyllospadix and comprise the first strong evidence of this phenomenon in seagrass populations. In addition, seed production rates predicted local seedling recruitment, demonstrating a measurable demographic consequence of low pollen availability.

Cross-species amplification of microsatellite loci within the dioecious, polyploid genus Actinidia (Actinidiaceae)

Microsatellite marker transfer across species in the dioecious genus Actinidia (kiwifruit) could offer an efficient and time-effective technique for use during trait transfer for vine and fruit improvement in breeding programmes. We evaluated the cross-species amplification of 20 EST-derived microsatellite markers that were fully informative in an Actinidia chinensis mapping family. We tested all 20 markers on 120 genotypes belonging to 21 species, 5 with varieties and/or chromosome races. These 26 taxa included 16 diploids, 7 tetraploids, 2 hexaploids and 1 octaploid, and represented all four taxonomic sections in the genus. All 20 markers showed some level of cross-species amplification. The most successful marker amplified in all genotypes from all species from all sections of the genus, the least successful amplified fragments only in A. chinensis and A. deliciosa. One species, A. glaucophylla, failed to amplify with all but 2 markers. PIC (Polymorphism information content) values were high, with 14 of 17 markers recording values of 0.90 and above. Sequence data demonstrated the presence of the microsatellite in all the amplified products. Sequence homology was less 5' of the microsatellite and increased toward the start codon of the translated region of the EST from which the marker was derived. The data confirm that EST-derived microsatellite markers from Actinidia species show cross-species amplification with high levels of polymorphism which could make them useful markers in breeding programmes.