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Karyological studies of Mediterranean stick-insects belonging to the genera Clonopsis and Leptynia (Insecta Phasmatodea)

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Abstract

Karyotypes of three species of Clonopsis and of various populations of Leptynia hispanica and L. attenuata complexes were analyzed. The taxonomic status of the latter complexes is still uncertain. These genera include bisexual and parthenogenetic polyploid species and populations. The chromosomal numbers are: 2n = 32 in bisexual C. algerica, 2n = 22 in bisexual C. maroccana, 3n = 54 in telytokous C. gallica, 2n = 38 in bisexual Leptynia hispanica, 3n = 57 in telytokous triploid populations of L. hispanica, 4n = 76 in telytokous tetraploid populations of the same complex, 2n = 36 in a bisexual taxon of L. attenuata complex, 2n = 38 in an other bisexual taxon and 4n = 76 in a parthenogenetic tetraploid taxon of the same complex. All the karyotypes are described. It is postulated that pericentric inversions and centric fusions have played a relevant role in chromosomal evolution of these genera. Because hybridization is the more frequent mode of speciation in these stick-insects, the polyploidy, connected with parthenogenetic reproduction is probably an allopolyploidy.

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... The phasmid genus Pijnackeria Scali 2009b is a Spanish monophyletic taxon embodying four diploid bisexual and two polyploid parthenogenetic species (3n, 4n), all sharing the same haploid chromosome set with n = 19 (Brock 1991(Brock , 1993Bianchi 1992¸Bianchi andMeliadò 1998;Scali 2009a;Scali et al. 2013). ...
... P. hispanica (Pantel 1890) (Fig. 1), the nominal species of the genus, appears to be a very successful tetraploid hybrid parthenogen, ranging from the Sierra Nevada (southern Spain) up to the Sistema Central Mountains, northwestwards, and to the Serrania de Cuenca, northeastwards. Here, its distribution area overlaps that of the similarly successful triploid hybrid parthenogen P. masettii Scali et al. 2013 which is also distributed up to the southern French districts of Var, Herault and Basses Alps (Bianchi 1992;Ghiselli et al. 2007;Scali 2009a, b;Scali et al. 2013) (Fig. 2). The remaining four taxa, namely P. lucianae (Scali et al. 2013), P. barbarae (Scali et al. 2013), P. lelongi (Scali et al. 2013) and P. originis (Scali et al. 2013), are diploid and show much more limited distribution areas in the south-eastern Iberian Peninsula, clearly suggestive of relict distribution (Fig. 2). ...
Article
The stick-insect genus Pijnackeria includes four diploid bisexual and two polyploid (3n, 4n) parthenogenetic species. Earlier analyses of the tetraploid parthenogen P. hispanica using mitochondrial markers allowed tracing its maternal ancestry to Pijnackeria originis, while no maternal nuclear contribution was found, thus suggesting an androgenetic and hybrid origin. The recently described Pijnackeria recondita—showing, among other features, a specific antennal structure linking it to the tetraploid parthenogen—prompted us to check whether the new species could be the unknown paternal ancestor of P. hispanica. In this work, we use karyology and molecular analysis of the mitochondrial gene cytochrome c oxidase subunit 2 (cox2) and the nuclear gene elongation factor 1 subunit α (ef1-α) to investigate the origin of such a complex tetraploid hybrid parthenogen. The molecular analysis supported P. recondita as being a paternal ancestor of P. hispanica, but also suggested that two more fathering species have to be taken into account: P. barbarae and the unknown paternal ancestor of the triploid hybrid P. masettii. Therefore, P. hispanica is apparently a polyphyletic chimeric androgen, which we propose to indicate as an androgenetic complex. Our data also revealed that P. hispanica is between 1.96 Myr and 3.31 Myr old, making it the oldest parthenogenetic taxon discovered among insects.
... Polyploid (3n and 4n) telytokous species of Leptynia (Heteronemiidae) are studied via analysis of karyotypes and DNA markers (Bianchi 1992;Passamonti et al. 2004;Ghiselli et al. 2007;Milani et al. 2009). Comparison with bisexual populations suggested hybridization as a possible mechanism that maintains polyploidy in parthenogenetic Leptynia hispanica (Bol ıvar, 1878) and Leptynia attenuate Pantel, 1890. ...
... Comparison with bisexual populations suggested hybridization as a possible mechanism that maintains polyploidy in parthenogenetic Leptynia hispanica (Bol ıvar, 1878) and Leptynia attenuate Pantel, 1890. In 4n populations of L. hispanica, allotetraploidy (duplication of both maternal and paternal chromosome sets each derived from different parental species) was proposed (Bianchi 1992). Recently, Leptynia genus has been splitted up with description of a new genus, Pijnackeria, on the basis of morphological, molecular and cytogenetic criteria. ...
Article
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In hexapods, unlike the majority of animals, development without fertilization is a common phenomenon. They evolved a striking diversity of unisexual reproductive types that include a variety of modes starting from spontaneous parthenogenesis in females to the production of impaternate males with different variants in between. Many reports about parthenogenetic species have accumulated over time. Here, we present a review of various parthenogenetic hexapod groups with a particular focus on their chromosome systems and ploidy level. We show that conclusions about the reproductive mode often lack solid evidence and sometimes inefficiently demonstrate how parthenogenesis is maintained in corresponding groups. In this review, basal hexapods (Protura, Collembola, Diplura), primarily wingless insect groups (‘Apterygota’) and non-holometabolous insects are listed with references to a variety of their unisexual reproductive modes.
... n. (Figs 5-6) The new species has been localized in the Ronda district (Sierra de Grazalema), where it colonizes small patches of shrubs and grasses (Genista, Cystus, Ru- bus). This is a genetically and chromosomally well differentiated taxon (Figs 2-4), long waiting its formal description (Bianchi, 1992;Bianchi and Meliado, 1998;Passamonti et al., 2004). ...
... Derivatio nominis. The species is dedicated to the distinguished Italian student of circum-Mediterranean stick-insects Anna Paola Bianchi, of Rome University La Sapienza, who first pointed out the occurrence of this southern Spanish Leptynia taxon with the same chromosome number as L. caprai, but with a clearly differing karyotype structure for some large pairs of the set (Bianchi, 1992). ...
Article
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Leptynia specimens were analyzed by karyotype analysis, mitochondrial gene sequencing and SEM of bodies and eggs. Here we describe a new species, Leptynia annaepaulae, and three subspecies of L. attenuata Pantel (L. attenuata attenuata, L. attenuata iberica, L. attenuata algarvica). The phylogeny of the genus Leptynia is congruent with a karyotype trend toward a reduction of chromosome number and the shift from the shared XX/X0 sex chromosome formula to the unusual XX/XY one. Chromosome repatterning appears to occur ahead of genetic differentiation, following a chromosome model of cladogenesis. Chromosome and genetic differentiation, in turn, appears to precede morphological distinction, thus realizing a condition of incipient species for most of the Leptynia taxa. Actually, morphological analyses revealed that, only rarely clear cut differences exist among and between taxa, while, more often, just trends in the differentiating traits occur, since the investigated characters generally suffer from some overlapping: In this study, only the 10th:9th ratio value and the subanal vomer appear to be diagnostic for L. annaepaulae against all other Leptynia taxa. As a consequence, the subanal vomer as well as cercus tooth features with egg chorion traits are not sharply diagnostic for the remaining co-generic taxa; however, comparisons are quite helpful in reducing uncertainties. A likely phylogeographic scenario for the genus supports that Leptynia ancestors spread from Northern Africa into Southern Spain where an ancestral taxon originated L. annaepaulae (2n = 40/39, XX/X0, with 2 large dibrachial pairs). Later on, a northbound colonization, should have originated L. caprai (2n = 40/39, XX/X0, all acrocentrics), from which L. montana (2n = 38, XX/X0) and L. attenuata (2n = 36, XY/XX) originated; supporting instances of chromosome repatterning have been actually observed. In this connection we like to stress that, particularly in stick insects, androgenesis has been a preferential pathway to quickly make homozygous those odd chromosome rearrangements likely responsible for low fitness in the heterozygotes.
... The genus Leptynia was formerly known to comprise two well-differentiated species: Leptynia hispanica Bolivar, 1878, and Leptynia attenuata Pantel, 1890. Both taxa have now to be considered as species complexes (Bianchi 1992;Scali 1996;Bianchi & Meliado 1998;Passamonti et al. 1999). However, within each complex, morphology does not allow clean discrimination, since no morphological character shows a neat divergence (Scali 1996). ...
... However, within each complex, morphology does not allow clean discrimination, since no morphological character shows a neat divergence (Scali 1996). Within the L. hispanica complex, bisexuals, triploid and tetraploid parthenogenetic populations have been found (Nascetti et al. 1983;Bianchi 1992), suggesting that polyploidy played a central role in the evolution of the species complex. This complex is therefore a good experimental system for studying these evolutionary and reproductive mechanisms. ...
Article
The Leptynia hispanica stick insect species complex includes bisexuals, triploid and tetraploid parthenogenetic populations, suggesting that polyploidy has played a central role in the evolution of this complex. An analysis of karyotype, mitochondrial DNA (cox2) and nuclear DNA (ef1-alpha) markers was carried out to clarify phylogenetic relationships and microevolutionary/phylogeographical patterns of the L. hispanica complex. Our analyses suggested a subdivision of bisexual populations into four groups, tentatively proposed as incipient species. Moreover, triploids and tetraploids showed two independent origins, the latter being more ancient than the former. From ef1-alpha analysis, triploids showed hybrid constitution, while the hybrid constitution of tetraploids is likely, but more data are needed. We suggest that L. hispanica is a case of 'geographical parthenogenesis' with parthenogenetic strains colonizing large peripheral ranges, and bisexuals confined to glacial refuge areas. Moreover, the age, wide distribution and competitive advantage of polyploids over diploids, demonstrate their significance in the evolution of the L. hispanica species complex.
... In the 80s, as collecting progressed, it became increasingly clear that both L. attenuata and L. hispanica had to be considered species-complexes: the former included only Mendelian taxa, the latter also polyploid parthenogens of hybrid origin (Nascetti et al. 1983;Brock 1991Brock , 1993Bianchi 1992;Lelong 1992). ...
Article
Recently, the Iberian stick insect genus Pijnackeria has been erected by splitting Leptynia Pantel on the basis of several distinguishing features. In addition to Pijnackeria hispanica, the tetraploid all-female type species, molecular, karyological and SEM investigations led to the recognition of four bisexual and one triploid unisexual new species. Bisexuals' karyotypes (2n = 37/38) differ for minute traits and the haploid set is repeated, with few differences, three or four times in the polyploids that appeared to be of hybrid origin. Diagnostic morphological traits were found among body size parameters, antennal articles, male cerci, ovipositor valve and egg chorionic features. All species commonly feed on the broom Sarothamnus scoparius, but habitat disturbance appeared to induce food plant shifts. Moreover, trends from bisexuality to unisexuality through spanandry, probably related to habitat disruption, have been witnessed. The diploid species (Pijnackeria lucianae, Pijnackeria barbarae, Pijnackeria lelongi and Pijnackeria originis) have small ranges, while the polyploid hybrids (Pijnackeria masettii and P. hispanica) spread through Spain and Southern France, featuring a clear geographic parthenogenesis scenario, by colonizing wide areas and likely displacing their ancestors, or even leading them to extinction. Cyclic climatic changes and natural or anthropic habitat fragmentation may have been also of relevance in shaping present-day distribution.
... Furthermore,Nascetti et al. (1983)proposed that both L. attenuata and L. hispanica could actually represent species-complexes and that the all-female populations were triploid and tetraploid forms related to bisexual L. hispanica; they also suggested that the parthenogenetic tetraploids originated by heterospecific hybridization. These hypotheses, based on preliminary electrophoretic gene–enzyme investigations, were supported through karyological findings byBianchi (1992); she also provided evidence that the triploid parthenogenetic populations should also have originated by hybridization. Thus, the new collections led to a much better definition of range, population structure and ecology of the two species-complexes. ...
Article
The Iberian phasmid species Leptynia attenuata and L. hispanica (Pantel, 189024. Pantel , P. J. 1890 . Notes Orthoptérologiques. II Les Phasmides d'Europe et des pays limitrophes. . Anales Sociedad Española de Historia Natural , 19 : 371 – 404 . View all references) are actually species‐complexes, each embodying a few specific taxa. Morphological, karyological and molecular investigations concordantly revealed that considerable differences exist between the two complexes, each representing a monophyletic group. Significant differentiating body traits are: the presence in the “attenuata” males of a well‐developed subanal vomer, which is lacking in those of the hispanica‐complex; the sharply pointed abdomen tip in the “hispanica” females, quite different from the rounded abdomen ending of the attenuata‐complex females. Also the egg structure and SEM chorionic pattern differ neatly. Karyotypes of the “attenuata” taxa mainly differ by chromosome repatternings entraining number reductions, while those of the hispanica‐complex maintain the same chromosome number in sexual taxa and add similar haploid sets in polyploid parthenogens. Also molecular investigations neatly separate the complexes and support specific differentiations of taxa within each complex; therefore, a splitting of the genus Leptynia is felt necessary. Caudell wrongly reported the designation by Pantel of L. hispanica as the type species of the genus, while Kirby validly indicated Leptynia attenuata as the type species; therefore, the subsequent Leptyniella proposition for the Leptynia attenuata complex by Bolìvar, is not valid either. Thus in the genus splitting, the Leptynia designation must be maintained to the “attenuata” complex, while the new name Pijnackeria is here given to the former “hispanica” taxa. This splitting is in full accordance with the seminal description by Pantel for the occurrence of the subanal vomer in the L. attenuata males and the presence of the pointed female abdomen in “hispanica” taxa, plainly settling a rather confusing nomenclatural situation. It also significantly updates the Iberian phasmid systematics. The formal description of taxa belonging to the reshaped Leptynia and to the new genus Pijnackeria will be dealt with in papers to follow.
... The model is well in accordance with the present knowledge of the formation of triploid hybrids in vertebrates (SUOMALAINEN et al. 1987) and plants (LEWIS 1980). Evidence for the association of allopolyploidy with parthenogenetic reproduction has also been presented in stick insects, in Phasmatodea by BIANCHI (1992). ...
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A novel method was applied to study chromosomes and their behavior in female meiosis in three parthenogenetic psocid species (Psocoptera). Two of these species Valenzuela flavidus (Steph.), Caeciliusidae and Peripsocus subfasciatus (Rambur), Peripsocidae displayed a triploid chromosome number 3x=27 while the third species Trichadenotecnum majus (Kolbe) was diploid (2n=18). In all species meiosis consisted of only a single, mitosis-like maturation division, i.e. meiosis was of apomictic type. Models for the formation of polyploid parthenogenetic forms are discussed.
Article
To Michael White. SUMMARY -Karyotypes of several bisexual populations of the Leptynia attenuata complex were analysed. This complex includes four karyotypically differentiated taxa: Leptymtl attenuata s.str., Leptymtl montana, Leptynia caprai and Leptynia sp. The chromosome numbers of the four species are: 2n=36 (XY; XX) for Leptynia attenuata, 2n=38 (X0; XX) for Leptynia montana, 2n=40 (X0; XX) for Leptymtl caprai and 2n=40 (X0; XX) for Leptynia sp. The evolutionary relationships between the four species are discussed .
Article
Karyological and allozymatic characterizations of recently collected samples of the Iberian Leptynia attenuata complex support the occurrence of three genetically differentiated groups, for which parallel morphological observations evidenced only partially diagnostic characters. The groups are: the Portuguese population of Fóia (Serra de Monchique), referred to as the nominal taxon, L. attenuata; the Spanish populations of the Sistema Central, referred to as L. montana; the populations of the Toledo district, referred to as L. caprai. These taxa seem to represent a case of incipient speciation, with chromosomal and genetic differentiation ahead of the morphological one. Chromosome repatternings, affecting autosomes as well as sex chromosomes, appear to go together with the evolutionary events.
Chapter
This chapter illustrates the variety of parthenogenetic mechanisms within three genera of phasmids (stick insects) occurring around the Mediterranean and the close ties between sexuals and asexuals. The link between parthenogenesis and polyploidy appears to provide an escape from cytogenetic disruption. When compared to diploids, both auto-polyploids and allo-polyploids can take advantage of higher mutation rates and the resulting increase of heterozygosity; they can also escape the accumulation of mutations. Detailed investigations of the karyotype structure, reproduction modes and phylogenetic relationships reveal a wide variety of reproductive modes including repeated backcrosses to parental species. Diploid hybrids can also reproduce hemiclonally, with one parental genome being eliminated and the hybrid being produced anew in each generation. However, also the reversal route from hybrids to the fathering species occurs through androgenesis, which resembles sexual reproduction with most of its genetic traits. All of these reproductive modes can be considered as patterns of “reticulate” evolution. Automicts, apomicts and hybrids with different amounts of recombination and levels of polyploidy (from diploids to tetraploids) have been found. The genetic diversity in these asexuals is kept high in mitochondrial DNA, rDNA cistrons, satellite DNA and coding genes. Other addressed issues, clearly linked to phasmid parthenogenesis are the peculiarities of spindle structure and the microtubule and γ-tubulin patterns for the egg development: these might be one of the reasons why phasmids are so exceptional in their variety of reproductive modes.
Article
An in-depth analysis of the Leptynia attenuata species complex has been performed by cytochrome oxidase subunit 2 (cox2) gene sequencing as well as karyotype and allozyme analysis. The whole set of data allows to largely resolve the taxonomy of the group and suggests an overall trend of chromosomal repatternings through a progressive reduction of the chromosome number. A previously suggested new species has been also confirmed on a genetic basis. Data are discussed in order to depict a phylogenetic and phylogeographic scenario fitting the observed genetic relationships between the different species of the group. Chromosome rearrangements are proposed as the major speciation driving force within the group and androgenetic reproduction is suggested as a shortcut to overcome the problem of fixing chromosomal rearrangements that are strongly underdominant in heterozygotes.
Chapter
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The problem of speciation remains one of the most intriguing areas of evolutionary biology. It is now recognised that there is not just one, but perhaps many possible speciation mechanisms (Dobzhansky 1972). To a large degree, the mode of speciation that occurs in any given instance is a function of the biological and population characteristics of the group of organisms concerned. However, independently of their mode of origin, all fully differentiated species have one basic characteristic in common—i.e., their reproductive or genetic isolation from all other species. To appreciate the fundamentals of the speciation process, we should therefore try to trace the evolution of this single distinguishing feature in each of the possible speciation patterns. This will require a consideration of all the possible modes of speciation, a knowledge of the succession of stages and key events for each, the preconditions which might apply, and any particularly unique aspects. The origin of the species gap is the feature of central interest. Thus, it is most important to assess those events which lead to reproductive isolation.
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Seven major races, with diploid numbers ranging from 26 to 40, and three types of sex-chromosome mechanism were found in the Australian phasmatid Didymuria violescens (Leach). The differences between chromosome complements are mainly due to translocations between autosomes, and to translocations between autosomes and sex chromosomes. The geographic pattern of chromosome variation and the characteristics of hybrids implicate chromosomal rearrangements in mechanisms of speciation.
Article
An increasing number of animal species of hybrid origin has been detected in recent years in different groups. In the present paper many of such cases are reviewed in detail and the mechanisms involved in their evolution are examined. The origin of an animal hybridogen requires: a) an interspecific hybridization, and b) changes in the maturation divisions allowing the transmission of the hybrid genome to the next generation. The latter are not caused by hybridization itself, and are certainly rare events; on the contrary, the role of high rates of interspecific hybridization should be taken into account, and the conditions for their occurrence are discussed.Animal hybrid species reproduce in different ways: thelytokous parthenogenesis, gynogenesis, hybridogenesis, self-fertilization, or fission. The consequences of these reproductive strategies are examined. Two phenomena are considered particularly relevant for the evolutionary success of animal hybridogens: 1. hybrid species being generally unisexual, their reproductive potential is doubled with respect to that of their bisexual ancestors (“demographic advantage”); 2. interspecific hybridization dramatically enhances heterozygosity (“heterotic advantage”). The short- and long-term effects of these phenomena on animal hybrid species are discussed.
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Following the recent discovery of seven new species, ten specific taxa are now recognized within the genus Bacillus: 1. B. rossius; 2. B. grandii; 3. B. whitei; 4. B. lynceorum; 5. B. atticus; 6. B. diplocarius; 7. B. carius; 8. B. rhodius; 9. B. cyprius; 10. B. creticus. Furthermore, eight subspecies have been defined within B. rossius. The distribution and the ecology of the species are summarized. On the basis of ootaxonomy, karyology, electrophoretic allozyme analysis and vitellin comparisons, the phyletic relationships among taxa within the genus are suggested, with particular emphasis on the natural thely‐tokous hybrids, B. whitei diploid and the triploid B. lynceorum. The main evolutionary mechanisms at work here appear to be: interspecific hybridization, polyploidy, chromosome repatterning and the switch to parthenogenetic reproduction. Gradual genetic differentiation has produced subspecific taxa in B. rossius.
Article
A wide diversity in chromosome complement is found in two species of phasmids of the primitive group Prisopini—Prisopus ariadne Hebard and Prisopus berosus Westwood. P. ariadne has a diploid male complement of 28, comprising 13 pairs of relatively large mediokinetic autosomes and Neo XY sex chromosomes. P. berosus, 2n =49, has relatively small autosomes most of which are mediokinetic, and retains the XO—XX sex mechanism. Chromosomal polymorphism in this species is suggested by the presence of an unequal pair of autosomes and a structural differentiation in the X in one of two males studied.The relative amount of DNA per nucleus in male germ cells (Peulgen cytophotometry) shows a significant difference in total chromosomal content between the complements of the two species.These data are discussed with reference to the cytotaxonomy of phasmids.
Article
The process which restores the zygoid number of chromosomes in the thelytokously parthenogenetic stick insectBacillus rossius Rossi, is described. During the first 15 days after oviposition approximately 200 haploid (A+X=18 chromosomes) pre-blastoderm cells develop in the egg-cortex. During the second period of 15 days up to about 2000 blastoderm cells develop in which were found: haploid mitoses, C-mitoses (A+X=18 chromosomes), endomitoses, diploid mitoses (2A+2X=36 chromosomes) and pycnosis. The next 10 days the number of pycnotic nuclei decreases gradually and the blastoderm cells multiply by diploid mitosis only. Thereafter, a diploid germ band is formed either directly or after a diapause. It is concluded that C-mitosis in haploid cells restores the somatic number of chromosomes (automictic parthenogenesis), that a portion of the cells in C-mitosis degenerates and that endomitosis leads to the formation of some polyploid cells only. This development is discussed in relation to the occurrence of geographic parthenogenesis and of impaternate males.
Article
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