Evolutionary pattern of karyotypes and meiosis in pholcid spiders (Araneae: Pholcidae): implications for reconstructing chromosome evolution of araneomorph spiders

Background Despite progress in genomic analysis of spiders, their chromosome evolution is not satisfactorily understood. Most information on spider chromosomes concerns the most diversified clade, entelegyne araneomorphs. Other clades are far less studied. Our study focused on haplogyne araneomorphs, which are remarkable for their unusual sex chromosome systems and for the co-evolution of sex chromosomes and nucleolus organizer regions (NORs); some haplogynes exhibit holokinetic chromosomes. To trace the karyotype evolution of haplogynes on the family level, we analysed the number and morphology of chromosomes, sex chromosomes, NORs, and meiosis in pholcids, which are among the most diverse haplogyne families. The evolution of spider NORs is largely unknown. Results Our study is based on an extensive set of species representing all major pholcid clades. Pholcids exhibit a low 2n and predominance of biarmed chromosomes, which are typical haplogyne features. Sex chromosomes and NOR patterns of pholcids are diversified. We revealed six sex chromosome systems in pholcids (X0, XY, X1X20, X1X2X30, X1X2Y, and X1X2X3X4Y). The number of NOR loci ranges from one to nine. In some clades, NORs are also found on sex chromosomes. Conclusions The evolution of cytogenetic characters was largely derived from character mapping on a recently published molecular phylogeny of the family. Based on an extensive set of species and mapping of their characters, numerous conclusions regarding the karyotype evolution of pholcids and spiders can be drawn. Our results suggest frequent autosome–autosome and autosome–sex chromosome rearrangements during pholcid evolution. Such events have previously been attributed to the reproductive isolation of species. The peculiar X1X2Y system is probably ancestral for haplogynes. Chromosomes of the X1X2Y system differ considerably in their pattern of evolution. In some pholcid clades, the X1X2Y system has transformed into the X1X20 or XY systems, and subsequently into the X0 system. The X1X2X30 system of Smeringopus pallidus probably arose from the X1X20 system by an X chromosome fission. The X1X2X3X4Y system of Kambiwa probably evolved from the X1X2Y system by integration of a chromosome pair. Nucleolus organizer regions have frequently expanded on sex chromosomes, most probably by ectopic recombination. Our data suggest the involvement of sex chromosome-linked NORs in achiasmatic pairing.

Evolutionary dynamics of rDNA clusters on chromosomes of buthid scorpions (Chelicerata: Arachnida)

We examined the distribution of genes for major ribosomal RNAs (rDNA) on holokinetic chromosomes of 74 species belonging to 19 genera of scorpions from the family Buthidae using fluorescence in situ hybridization (FISH). Our analysis revealed differences between the two main evolutionary lineages within the family. The genera belonging to the ‘Buthus group’, with a proposed Laurasian origin, possess one pair of rDNA mainly in an interstitial position, with the only exceptions being the terminal location found in some Hottentotta and Buthacus species, possibly as a result of chromosome fissions. All the remaining buthid ‘groups’ possess rDNA found strictly in a terminal position. However, the number of signals may increase from an ancestral state of one pair of rDNA loci to up to seven signals in Reddyanus ceylonensis Kovařík et al., 2016. Despite the differences in evolutionary dynamics of the rDNA clusters between the ‘Buthus group’ and other lineages investigated, we found a high incidence of reciprocal translocations and presence of multivalent associations during meiosis in the majority of the genera studied. These phenomena seem to be typical for the whole family Buthidae.

Comparative FISH mapping of ribosomal DNA clusters and TTAGG telomeric sequences to holokinetic chromosomes of eight species of the insect order Psocoptera

Repetitive DNAs are the main components of eukaryotic genome. We mapped the 18S rDNA and TTAGG telomeric probe sequences by FISH to meiotic chromosomes of eight species of the order Psocoptera considered a basal taxon of Paraneoptera: Valenzuela burmeisteri (Brauer, 1876), Stenopsocus lachlani Kolbe, 1960, Graphopsocus cruciatus (Linnaeus, 1768), Peripsocus phaeopterus (Stephens, 1836), Philotarsus picicornis (Fabricius, 1793), Amphigerontia bifasciata (Latreille, 1799), Psococerastis gibbosa (Sulzer, 1766), and Metylophorus nebulosus (Stephens, 1836). These species belong to five distantly related families of the largest psocid suborder Psocomorpha: Caeciliusidae, Stenopsocidae, Peripsocidae, Philotarsidae, and Psocidae. We show that all the examined species share a similar location of 18S rDNA on a medium-sized pair of autosomes. This is the first study of rDNA clusters in the order Psocoptera using FISH. We also demonstrate that these species have the classical insect (TTAGG)n telomere organization. Our results provide a foundation for further cytogenetic characterization and chromosome evolution studies in Psocoptera.

Cyperaceae in Mexico: Diversity and distribution

<p><strong>Background</strong>: Cyperaceae, with about 5,500 species and 90 genera worldwide, are the third largest family among Monocots. A unique combination of morphological and karyotypical features, among which stand holokinetic chromosomes, favors a rapid evolution and diversification and a high level of endemism in some groups. Preliminary checklists of Mexican sedges have been published but an updating of the taxonomy and nomenclature of the group for the country is required.</p><p><strong>Questions</strong>: How many and which species and genera of Cyperaceae are in Mexico?, what patterns of geographic distribution display those species?, which are the main gaps in the systematic knowledge in the family?</p><p><strong>Study site and years of study</strong>: Mexico, 1990 to 2016.</p><p><strong>Methods</strong>: A database of Mexican Cyperaceae was generated with basis in literature review, study of herbarium specimens (11 herbaria in Mexico and the United States) and field work, the last mainly focused on <em>Carex</em>. Diversity and endemism level were calculated. Besides, we analyzed in different space scales their distributional range.</p><p><strong>Results</strong>: Our dataset includes 460 species and 20 infraspecific taxa in 21 genera that belong to 10 of the 17 tribes of the family. Subfamily Cyperoideae includes almost 100 % of the Mexican sedges, as only one representative of subfamily Mapanioideae is known for the country. At the generic level, a drastic reduction in number (21) in comparison to previous inventories (27) results of recent phylogenetic and taxonomic rearrangements. The most diverse genera are <em>Carex</em> (138 taxa) and <em>Cyperus</em> (125), followed by <em>Rhynchospora</em> (65) and <em>Eleocharis</em> (57). Sedges in Mexico are found from sea level to above 4,300 m, in all types of vegetation. The highest diversity was found for Chiapas (237 taxa, 52 % of the total) and Veracruz (206 taxa, 45 %), followed by Oaxaca and Jalisco. Two genera (<em>Cypringlea</em> and <em>Karinia</em>) and 111 species or infraspecific taxa are endemic to Mexico (24 %), 43 of them micro-endemic (only known from one state in the country). Endemism increases to 57 % when the biogeographic extension known as Megamexico is included. Forty six names are excluded from the Mexican flora.</p><strong>Conclusions</strong>: Regardless of the addition of taxa and refining of the databases, the checklist presented here is still preliminary. Collection deficiencies and insufficient taxonomic revision for Mexican sedges are reflected in gaps in their knowledge. There are at least 45 undescribed species; including them the richness of Mexican sedges would exceed 500 species. Many complexes of species are in need of taxonomic revision, mainly in <em>Carex</em> but also in<em> Bulbostylis</em>, <em>Cyperus</em>, <em>Eleocharis</em>, <em>Rhynchospora</em> and <em>Scleria</em>. To advance in the inventory and better understanding of the diversity of Mexican Cyperaceae, we propose some research topics to be addressed in the short term.<p> </p>

Cytological evidence for automictic thelytoky in parthenogenetic oribatid mites (Acari, Oribatida): Synaptonemal complexes confirm meiosis in Archegozetes longisetosus

Diplo-diploid parthenogenesis (thelytoky) is a widespread phenomenon in the mite taxon Sarcoptiformes, and is unusually frequent in the suborder Oribatida, where it characterizes almost 10% of extant species, including whole genera and families. Based on molecular and cytological data, terminal fusion automixis with an inverted meiotic sequence based on holokinetic chromosomes has been suggested as the reproductive mode in these mites. However, unequivocal structural evidence for meiosis is missing. The model organism Archegozetes longisetosus, a thelytokous member of the parthenogenetic family Trhypochthoniidae, was studied to gain ultrastructural insight in oocyte progression and meiotic processes. In this study, ovarian nuclear organization of its tritonymphal instar was examined by transmission electron microscopy (TEM). Numerous synaptonemal complexes were observed in the ovary, unequivocally confirming automixis (meiotic thelytoky) in oribatid mites for the first time. No recombination nodules were found. The nuclei are transcriptionally active in late prophase. Inverted meiosis is discussed as a result of the spatial arrangement of chromatid segregation.

Comparison of different cytogenetic methods and tissue suitability for the study of chromosomes in Cimex lectularius (Heteroptera, Cimicidae)

In the article we summarize the most common recent cytogenetic methods used in analysis of karyotypes in Heteroptera. We seek to show the pros and cons of the spreading method compared with the traditional squashing method. We discuss the suitability of gonad, midgut and embryo tissue inCimexlectulariusLinnaeus, 1758 chromosome research and production of figures of whole mitosis and meiosis, using the spreading method.The hotplate spreading technique has many advantages in comparison with the squashing technique. Chromosomal slides prepared from the testes tissue gave the best results, tissues of eggs and midgut epithelium are not suitable. Metaphase II is the only division phase in which sex chromosomes can be clearly distinguished. Chromosome number determination is easy during metaphase I and metaphase II. Spreading of gonad tissue is a suitable method for the cytogenetic analysis of holokinetic chromosomes ofCimexlectularius.

Non-Random Distribution of 5S rDNA Sites and Its Association with 45S rDNA in Plant Chromosomes

5S and 45S rDNA sites are the best mapped chromosome regions in eukaryotic chromosomes. In this work, a database was built gathering information about the position and number of 5S rDNA sites in 784 plant species, aiming to identify patterns of distribution along the chromosomes and its correlation with the position of 45S rDNA sites. Data revealed that in most karyotypes (54.5%, including polyploids) two 5S rDNA sites (a single pair) are present, with 58.7% of all sites occurring in the short arm, mainly in the proximal region. In karyotypes of angiosperms with only 1 pair of sites (single sites) they are mostly found in the proximal region (52.0%), whereas in karyotypes with multiple sites the location varies according to the average chromosome size. Karyotypes with multiple sites and small chromosomes (<3 µm) often display proximal sites, while medium-sized (between 3 and 6 µm) and large chromosomes (>6 µm) more commonly show terminal or interstitial sites. In species with holokinetic chromosomes, the modal value of sites per karyotype was also 2, but they were found mainly in a terminal position. Adjacent 5S and 45S rDNA sites were often found in the short arm, reflecting the preferential distribution of both sites in this arm. The high frequency of genera with at least 1 species with adjacent 5S and 45S sites reveals that this association appeared several times during angiosperm evolution, but it has been maintained only rarely as the dominant array in plant genera.