What Do Pneumocystis Organisms Tell Us about the Phylogeography of Their Hosts? The Case of the Woodmouse Apodemus sylvaticus in Continental Europe and Western Mediterranean Islands

Pneumocystis fungi represent a highly diversified biological group with numerous species, which display a strong host-specificity suggesting a long co-speciation process. In the present study, the presence and genetic diversity of Pneumocystis organisms was investigated in 203 lung samples from woodmice (Apodemus sylvaticus) collected on western continental Europe and Mediterranean islands. The presence of Pneumocystis DNA was assessed by nested PCR at both large and small mitochondrial subunit (mtLSU and mtSSU) rRNA loci. Direct sequencing of nested PCR products demonstrated a very high variability among woodmouse-derived Pneumocystis organisms with a total number of 30 distinct combined mtLSU and mtSSU sequence types. However, the genetic divergence among these sequence types was very low (up to 3.87%) and the presence of several Pneumocystis species within Apodemus sylvaticus was considered unlikely. The analysis of the genetic structure of woodmouse-derived Pneumocystis revealed two distinct groups. The first one comprised Pneumocystis from woodmice collected in continental Spain, France and Balearic islands. The second one included Pneumocystis from woodmice collected in continental Italy, Corsica and Sicily. These two genetic groups were in accordance with the two lineages currently described within the host species Apodemus sylvaticus. Pneumocystis organisms are emerging as powerful tools for phylogeographic studies in mammals.     

Nuclear-DNA markers confirm the presence of two anchovy species in the Mediterranean

Morphometric, allozymic, and mitochondrial DNA variability previously indicated that the Mediterranean anchovy Engraulis encrasicolus includes two distinct forms, one of the inshore habitat, and the other one of the open-sea habitat. Here, we showed that the two forms significantly differ by several morphological characters. To test the hypothesis, proposed previously, that the two forms are distinct biological species, we used length-polymorphic, exon-primed intronic PCR markers from the creatine-kinase multigenic family and genetically characterized anchovy samples collected in the northwestern Mediterranean and in the eastern Atlantic. Large genetic differences were found between the two forms (Weir and Cockerham's theta = 0.397 to 0.586). In contrast, geographic variation in the open-sea form at the scale of the eastern Atlantic/northwestern Mediterranean was weak (theta = -0.006 to 0.042). This again demonstrated considerable restriction to nuclear gene flow between inshore and open-sea anchovy populations. In addition to previous results from allozymes, this confirmed their status as distinct biological species, namely Engraulis albidus sp. nov. and Engraulis encrasicolus L.     

Macrogeographic variations in food choice of mainland citril finches Carduelis [citrinella] citrinella versus insular Corsican (citril) finches Carduelis [citrinella] corsicanus

We studied variations in diet composition in sub-populations of mainland citril finches Carduelis [citrinella] citrinella in European mountain chains, compared to closely related insular Corsican (citril) finches Carduelis [citrinella] corsicanus on several Mediterranean Islands with a special emphasis on key food plants used during the breeding period. We found that citril finch sub-populations of the Pre-Pyrenees and the Black Forest rely mostly on mountain pine and dandelion seeds as main food resources during the breeding season. In contrast, insular Corsican finches on the Mediterranean islands of Capraia and Corsica fed mainly on various herb species. Finches in Corsica fed predominantly on shepherd’s purse, while birds from Capraia were choosing rosemary as their main food plant during the breeding period. These observations confirmed the previously expected difference in food choice between the two forms due to the observed increased niche breadth on Corsica. However, Corsican finches on Sardinia showed a food choice more similar to that of the citril finch populations, in that they fed predominantly on black pine seeds during breeding season. Thus we were not able to find clear behavioural differences in diet composition between all studied sub-populations of citril and Corsican finches. To conclude, food choice of both forms is highly adaptive and, in the different breeding areas, distinct key plants may be of special importance during the breeding season. The preference for these plants is likely to be linked to their abundance and their energetic and nutritional content.     

鸟类鸣声行为对其物种分化和新种形成影响

鸟声和鸟类的形态特征一样,具有物种的特异性,在鸟类分类和野外识别方面是有意义的,从生物种的基本概念出发,新种的形成有赖于两个亲缘种群间不能相互婚配和繁殖隔离的产生。鸟类主要的有关特征是在配对形成过程中的听觉和视觉特征,如果出现鸣声特征的差异,而且这种差异已经超出了种间“语言通讯”的变化范围,那么它就有可能与群内其它异笥无法配对,从而被排除到该种群或种的范围之外,然而鸣禽的鸣唱有时在不同种群间变化很大,而在同一种群内比较稳定,从而形成种群的方言,在方言种群之间,如果长期隔离,就会在种群之间形成障碍乃至遗传上的隔离,从而使种群间失去了相互配偶的机会,新的亚种或种可能由此形成,文章最后假设出了基于鸣声行为的鸟类新种形成机理图解。     

New insights into the past and recent evolutionary history of the Corsican mouflon (Ovis gmelini musimon) to inform its conservation

Human-mediated species dispersal across the Mediterranean stretches back at least 10,000 years and has left an indelible stamp on present-day biodiversity. Believed to be a descendant of the Asiatic mouflon (Ovis gmelini gmelinii), the Corsican mouflon (O. g. musimon) was translocated during the Neolithic as ancestral livestock by humans migrating from the Fertile Crescent to the Western Mediterranean. Today, two geographically limited and disconnected populations can be found in Corsica. Whether they originated from distinct founders or one ancestral population that later split remains unknown, although such information is pivotal for the species’ management on the island. We genotyped 109 and 176 individuals at the Cytochrome-b gene and 16 loci of the microsatellite DNA, respectively, to gain insights into the natural history of the Corsican mouflon. We found evidence confirming that the Asiatic was the ancestor of the Corsican mouflon, which should thus be unvaryingly referred to as O. g. musimon, i.e. as a subspecies of the Asiatic mouflon. Haplotype divergence dating and the investigation of genetic structure highlighted a strong and ancient genetic differentiation between the two Corsican populations. Approximate Bayesian Computation pointed to the introduction of a single group of founders as the most reliable scenario for the origin of the entire Corsican population. Later, this ancestral stock would have decreased in number, facing genetic bottlenecks and eventually resulting in two divergent demes. Splitting most likely occurred several hundred years ago. Their shared past notwithstanding, we discuss whether the two relic Corsican mouflon populations should be now considered as distinct management units.

     

A Contrast in Extra-Pair Paternity Levels on Mainland and Island Populations of Mediterranean Blue Tits

Extra‐pair copulation (EPC) is a widespread behaviour amongst birds, yet the rate of occurrence of extra‐pair offspring is highly variable both within and across species. Two populations of Mediterranean blue tits, Parus caeruleus, one on the mainland and the other on the island of Corsica, are characterized by high levels of extra‐pair paternity (EPP), with 14 and 25% of extra‐pair offspring (in 46 and 68% of the nests), respectively. The rate of EPP is significantly higher in the island population. Such high rates on Corsica contradict the predictions of higher mate fidelity and lower levels of EPP on islands. Despite expressing several traits that characterize an insular syndrome, blue tits of Corsica do not exhibit lower genetic variation which could be associated with low levels of EPP on islands. Furthermore, our results do not support any of the hypotheses that have been proposed so far to explain variation in EPP rates at the local level. We tentatively suggest that proximate Mediterranean constraints, especially consistent food shortage, prompt females to seek better breeding opportunities through a two‐step process including high rates of EPC and divorce.     

Temporal pattern of vocalization type usage in singing sessions of male tyrant flycatchers (Tyranninae)

It has been hypothesized that variation (e.g., of repertoire elements) in prolonged vocalization sessions of passerine birds can serve to minimize habituation by conspecifics. The repertoire of vocalization types limited in suboscine passerines, raising the question of how a limited set of elements can create patterns that minimize habituation. This question was studied by computational analysis of recorded singing sessions of 20 suboscine species from the subfamily Tyranninae, family Tyrannidae (tyrant flycatchers). The recordings of 12 of the species included two or more distinct vocalization types (VTs). In these species, the interval between vocalization units when the VT changed was on average shorter than that when the VT remained the same. In addition, when the VT changed, the mean interval length between successive vocalization units differed depending on which VT preceded the interval and which VT followed it. On the other hand, species with just a single VT in the session analyzed showed a surprisingly high degree of absolute difference between adjacent vocalization units with respect to both the length of the vocalization and percentage of time elapsed until peak amplitude. A change in the rhythm of vocalization accompanying a change in VT provided a potential means of drawing a conspecific listener's attention to the change in VT. The results showed that tyrant flycatchers use temporal patterning to achieve a high level of variety in vocalization sessions despite a limited vocal repertoire.     

GENETIC DIFFERENCES BETWEEN TWO GROWTH‐FORMS OF LITHOPHYLLUM MARGARITAE (RHODOPHYTA) IN BAJA CALIFORNIA SUR,MEXICO1

Unattached, nongeniculate, coralline algae or rhodoliths exhibit a range of morphological variability seemingly dependent on environmental factors. Rhodoliths have an extensive fossil record, and environmentally dependent characteristics make them potentially reliable paleoindicators. Species of the rhodolith‐forming genus Lithophyllum Philippi in Baja California Sur, Mexico were recently consolidated into one species. Under the new classification, L. margaritae (Hariot) Heydrich consists of several growth forms presumably reflecting local environmental conditions. We examined the genetic structure of four populations of this species using amplified fragment length polymorphisms (AFLP) to characterize the extent of genetic variation associated with foliose and fruticose growth forms. AFLP band sharing analysis revealed that foliose growth forms exhibited consistently higher intrapopulation similarities (0.75–0.85) than fruticose growth forms (similarity range, 0.55–0.67). This trend was also evident in comparisons of geographically isolated populations. These data indicate that the two morphologies are genetically distinct and that genetic exchange between foliose and fruticose growth forms of L. margaritae may be limited. Consequently, rhodolith growth forms appear to be the result of an interplay between both genetic makeup and environmental conditions.     

Evidence for reproductive isolation between cave bear populations

The European cave bear (Ursus spelaeus), which became extinct around 15,000 years ago, had several morphologically different forms. Most conspicuous of these were small Alpine cave bears found at elevations of 1,600 to 2,800 m. Whereas some paleontologists have considered these bears a distinct form, or even a distinct species, others have disputed this. By a combination of morphological and genetic methods, we have analyzed a population of small cave bears from Ramesch Cave (2,000 m altitude) and one of larger cave bears from Gamssulzen Cave (1,300 m), situated approximately 10 km apart in the Austrian Alps (Figure 1A). We find no evidence of mitochondrial gene flow between these caves during the 15,000 years when they were both occupied by cave bears, although mitochondrial DNA sequences identical to those from Gamssulzen Cave could be recovered from a site located about 200 km to the south in Croatia. We also find no evidence that the morphology of the bears in the two caves changed to become more similar over time. We suggest that the two cave bear forms may have represented two reproductively isolated subspecies or species.     

Genetic diversity and evolutionary history of the Tyrrhenian treefrog Hyla sarda (Anura: Hylidae): adding pieces to the puzzle of Corsica–Sardinia biota

The Corsica–Sardinia archipelago is a hotspot of Mediterranean biodiversity. Although tempo and mode of arrival of species to this archipelago and phylogenetic relationships with continental species have been investigated in many taxa, very little is known about the current genetic structure and evolutionary history subsequent to arrival. In the present study, we investigated genetic variation within and among populations of the Tyrrhenian treefrog Hyla sarda, a species endemic to the Corsica–Sardinia microplate and the surrounding islands, by means of allozyme electrophoresis. Low genetic divergence (mean D_nei = 0.01) and no appreciable differences in the levels and distribution of genetic variability (H_E: 0.06–0.09) were observed among all but one populations (Elba). Historical demographic and isolation‐by‐distance analyses indicated that this diffused genetic homogeneity could be the result of recent demographic expansion. Along with paleoenvironmental data, such expansion could have occurred during the last glacial phase, when wide and persistent land bridges connected the main islands and a widening of lowland areas occurred. This scenario is unprecedented among Corsica–Sardinia species. Together with the lack of concordance even among the few previously studied species, this suggests either that species had largely independent responses to paleoenvironmental changes, or that most of the history of assembly of the Corsica–Sardinia biota is yet to be written. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 103 , 159–167.     

Geographical differentiation, acoustic adaptation and species boundaries in mainland citril finches and insular Corsican finches, superspecies Carduelis [citrinella]

Aim In birds, differentiation of acoustic characters is an important mechanism of reproductive isolation that may lead to an ethological–acoustic barrier, resulting in the formation of new species. We examined acoustic variation in mainland citril and insular Corsican finch populations, with the aim of assessing the degree of acoustic differentiation between both members of the superspecies Carduelis [citrinella] and documenting possible variation between local subpopulations that are geographically isolated. Location We chose study sites throughout the geographical ranges of citril and Corsican finches. For the citril finch, we obtained samples from the Black Forest (Germany), the Cevennes (France) and the Pyrenees (Spain); for the Corsican finch, we obtained samples from the islands Capraia and Sardinia (Italy) and Corsica (France). Methods We analysed frequent contact calls and elements of the perch song. Vocalization patterns of the study populations were compared by means of discriminant and hierarchical cluster analyses. Results There were significant differences in vocalization characteristics of perch songs and contact calls, which permitted unambiguous discrimination of citril and Corsican finch populations. However, we also detected significant differences in contact calls between mainland citril finch subpopulations. There was a pattern of clinal variation in vocalization: short, steeply modulated signals in the northern part of the geographical range (Black Forest) and long, shallowly modulated signals in the southern part (Pyrenees). Main conclusions Acoustically, mainland citril and insular Corsican finches separate well in their contact calls and perch songs. However, variation in the two vocalization patterns between subpopulations of mainland citril finches indicates that acoustic characteristics can evolve very quickly, not only on islands but also on the mainland. Local habitat differences may play a crucial role in the rapid evolution of these signals under full or partial isolation of small subpopulations. To judge the importance of signal variation as a pre‐mating isolating barrier, future studies will have to determine whether members of the distinct subpopulations are able to match their signals to each other if they re‐meet, and whether intraspecific species recognition is still possible.     

Species status and population genetic structure of the flightless chafer beetles Prodontria modesta and P. bicolorata (Coleoptera; Scarabaeidae) from South Island, New Zealand

This study uses isozyme electrophoresis to investigate the systematic status of two rare flightless chafer beetle species, Prodontria modesta and P. bicolorata , currently distinguished solely on the basis of colour. Seven polymorphic loci were analysed for the species in sympatry and allopatry. In sympatry, gene frequencies imply no genetic barrier between the two colour forms. Wright's hierarchical F-statistics were calculated to determine how the genetic variation is partitioned across the geographic range of the two species. Strong geographic structuring occurs at the population level but there is little genetic differentiation attributable to species. Comparisons of morphological measurements are in support of the electrophoretic results. The systematic significance of the two distinct colour forms is thus questionable. Under the biological, recognition or phylogenetic species concepts, the genetic data suggest that there is only one species, polytypic for a colour pattern. Most of the genetic diversity of this group of beetles lies within and among populations, which differ quite markedly over their geographic range, rather than between the two named species.     

Ecology and genetics of interspecific hybridization in the swallowtails, Papilio hospiton Gene and P. machaon L., in Corsica (Lepidoptera: Papiliomdae)

The Corsican swallowtail butterfly, Papilio hospiton , is endemic to Corsica and Sardinia (France and Italy) and included in the list of endangered species by the Washington Convention, It is spread all over Corsica in scattered populations linked to diverse habitats. A study by enzyme electrophoresis showed that the genetic diversity of the species is of the same order of magnitude as that of Papilio machaon from continental France and Corsica. The differentiation between populations is rather low, which is consistent with the high vagility of the adults. Natural hybridization between P. hospiton and P. machaon is frequent, and laboratory crosses show that the hybrids are not sterile. However, developmental perturbations impair the viability of further hybrid progenies. Although limited introgression between the two species is likely to take place, enzyme electrophoresis and PCR-RFLP analysis of mitochondrial DNA show that their gene pools remain distincfTtienetic assimilation by P. machaon therefore does not seem to be a threat for P. hospiton.     

Predicting distribution of <Emphasis Type="Italic">Zygaena</Emphasis> moths on West Mediterranean islands. Implications for biogeography and conservation (Lepidoptera Zygaenidae)

In this paper a conservation biogeography approach has been applied to predict distribution and conservation priorities in West Mediterranean Zygaena moths. The presence/absence matrix data for mainland region and for the three largest islands (Sardinia, Corsica and Sicily) has been analyzed. The analyses have been performed on individual species and revealed that species distribution in the Mediterranean mainland can be largely predicted. However, Sardinia and Corsica islands showed highly impoverished faunas while Sicily did not revealed a lower richness than predicted. Logistic regressions at individual species level showed that several taxa, predicted to be present in Sardinia and Corsica are actually absent. For Sicily the opposite trend was obtained and several unpredicted species actually occur on this island. Conversely to butterflies, ecological traits did not differ between predicted and unpredicted Sicilian species, suggesting that ancient and stochastic colonisations are responsible for the occurrence of several species. Due to the very low probabilities that relict populations could re-colonize islands following possible extinctions, they are suggested to deserve particular conservation efforts. In particular, Z. orana from Sardinia, Z. corsica from Corsica and Z. trifolii, Z. purpuralis and Z. carniolica from Sicily showed distinctive biogeographic patterns and/or particular rarity.     

Genetic variation among Corsican and continental populations of the Eurasian treecreeper (Aves: Certhia familiaris) reveals the existence of a palaeoendemic mitochondrial lineage

In this study we investigated the phylogenetics of the Eurasian treecreeper (Certhia familiaris), a forest passerine with a wide Palaearctic range including Corsica, using three mitochondrial genes and three nuclear introns, and its phylogeographic history using the COI gene. Our phylogenetic results, including eight of the ten sub‐species currently recognized, support the monophyly of C. familiaris with respect to its Indo‐Asian sister species C. hodgsoni. C. familiaris comprises two lineages that diverged during the mid‐Pleistocene (c. 1 Myr): one palaeoendemic lineage has an allopatric range nowadays restricted to the Corsica island and the Caucasus region whereas the second one, more recent and widespread, is distributed over most of Eurasia and in northern China. The most likely scenario that may explain such a pattern is a double colonization of the western Palaearctic from the eastern range of the species. During the middle Pleistocene period, a first lineage expanded its range up into Europe but did not persist through glacial cycles except in Corsica and the Caucasus region. Later, during the upper Pleistocene, a second lineage began to diversify around 0.09 Myr, spreading towards the western Palaearctic from a unique refuge likely located in the eastern Palaearctic [correction added on 6 March 2015 after first online publication: 0.9 Myr amended to 0.09 Myr]. Apart from C. f. corsa, our results do not suggest any distinct evolutionary history for other sub‐species previously described on morphological grounds in Europe. Our study highlights the important conservation value of the Corsican treecreeper and emphasizes the major role of mature pine forests in the evolution of endemic bird taxa in Corsica. © 2015 The Linnean Society of London, Biological Journal of the Linnean Society, 2015, 115 , 134–153.     

Plant genetics: when not to interfere

New evidence suggests that the model plant Arabidopsis has two biochemically distinct pathways that produce genetic crossovers. Studies in several organisms have revealed that one kind of crossover regulation - crossover interference - is applied differently from species to species. Arabidopsis appears to use an interference system similar to that of budding yeast.     

Affiliative behavior, ultrasonic communication and social reward are influenced by genetic variation in adolescent mice

Social approach is crucial for establishing relationships among individuals. In rodents, social approach has been studied primarily within the context of behavioral phenomena related to sexual reproduction, such as mating, territory defense and parental care. However, many forms of social interaction occur before the onset of reproductive maturity, which suggests that some processes underlying social approach among juvenile animals are probably distinct from those in adults. We conducted a longitudinal study of social investigation (SI) in mice from two inbred strains to assess the extent to which genetic factors influence the motivation for young mice to approach one another. Early-adolescent C57BL/6J (B6) mice, tested 4-6 days after weaning, investigated former cage mates to a greater degree than BALB/cJ (BALB) mice, irrespective of the sex composition within an interacting pair. This strain difference was not due to variation in maternal care, the phenotypic characteristics of stimulus mice or sensitivity to the length of isolation prior to testing, nor was it attributable to a general difference in appetitive motivation. Ultrasonic vocalization (USV) production was positively correlated with the SI responses of mice from both strains. Interestingly, several USV characteristics segregated with the genetic background of young mice, including a higher average frequency and shorter duration for the USVs emitted by B6 mice. An assessment of conditioned place preference responses indicated that there was a strain-dependent difference in the rewarding nature of social contact. As adolescent mice aged, SI responses gradually became less sensitive to genetic background and more responsive to the particular sex of individuals within an interacting pair. We have thus identified a specific, genetic influence on the motivation of early-adolescent mice to approach one another. Consistent with classical theories of motivation, which propose a functional relationship between behavioral approach and reward, our findings indicate that reward is a proximal mechanism through which genetic factors affect social motivation during early adolescence.     

Population genetic structure of Tomicus piniperda L. (Curculionidae: Scolytinae) on different pine species and validation of T. destruens (Woll.)

Genetic diversity and population structure of Tomicus piniperda was assessed using mitochondrial sequences on 16 populations sampled on 6 pine species in France. Amplifications of Internal transcribed space 1 (ITS1) were also performed. Our goals were to determine the taxonomic status of the Mediterranean ecotype T. piniperda destruens, and to test for host plant or geographical isolation effect on population genetic structure. We showed that T. piniperda clusters in two mtDNA haplotypic groups. Clade A corresponds to insects sampled in continental France on Pinus sylvestris, P. pinaster and P. uncinata, whereas clade B gathers the individuals sampled in Corsica on P. pinaster and P. radiata and in continental France on P. pinea and P. halepensis. Insects belonging to clade A and clade B also consistently differ in the length of ITS1. Individuals belonging to both clades were found once in sympatry on P. pinaster. Genetic distances between clades are similar to those measured between distinct species of Tomicus. We concluded that clade B actually corresponds to the destruens ecotype and forms a good species, T. destruens. Analyses of molecular variance ( amova ) were conducted separately on T. destruens and T. piniperda to test for an effect of either geographical isolation or host species. Interestingly, the effect of host plant was significant for T. piniperda only, while the effect of geographical isolation was not. Pine species therefore seems to act as a significant barrier to gene flow, even if host race formation is not observed. These results still need to be confirmed by nuclear markers.     

Morphological and genetic evidence supports the existence of two species in the genus Ophelia (Annelida, Polychaeta) from the Western Mediterranean

About 900 individuals of the intertidal polychaete Ophelia cf. bicornis were collected at four sandy beaches in Corsica, ten in Sardinia and one in Tuscany (Western Mediterranean). They were analysed morphologically and genetically to clarify the taxonomic status of the littoral species of Ophelia in the study area and to assess the population genetic structuring. Two morphotypes were distinguished on the basis of the number of nephridiopore pairs (five or six). Genetic distances, F -statistics and analysis of molecular variance based on eight allozyme genetic markers were consistent in showing the two morphotypes to separate at species level. Thus, the two morphotypes which most authors have included in O. bicornis s.l. are not one morphologically variable taxon, but instead constitute two valid species, namely O. bicornis Savigny, 1818, with six pairs of nephridiopores, and O. barquii Fauvel, 1927, with five pairs of nephridiopores. Mechanisms of ecological speciation in a background of gene flow can be taken into account to explain the origin of the two species. Moderate genetic differentiation was found among populations within each species, suggesting that dispersal may not be effective over large distances, or that local selection plays an important role in population differentiation.  © 2004 The Linnean Society of London, Biological Journal of the Linnean Society , 2004, 83 , 101–113.     

Evidence for subdivision within the M molecular form of Anopheles gambiae

The principal vector of malaria in sub-Saharan Africa, Anopheles gambiae is subdivided into two molecular forms M and S. Additionally, several chromosomal forms, characterized by the presence of various inversion polymorphisms, have been described. The molecular forms M and S each contain several chromosomal forms, including the Savanna, Mopti and Forest forms. The M and S molecular forms are now considered to be the reproductive units within A. gambiae and it has recently been argued that a low recombination rate in the centromeric region of the X chromosome has facilitated isolation between these forms. The status of the chromosomal forms remains unclear however. Therefore, we studied genetic differentiation between Savanna S, Forest S, Forest M and Mopti M populations using microsatellites. Genetic differentiation between Savanna S and Forest S populations is very low (F(ST) = 0.0053 +/- 0.0049), even across large distances. In comparison, the Mopti M and Forest M populations show a relatively high degree of genetic differentiation (F(ST) = 0.0406 +/- 0.0054) indicating that the M molecular form may not be a single entity, but could be subdivided into at least two distinct chromosomal forms. Previously it was proposed that inversions have played a role in the origin of species within the A. gambiae complex. We argue that a possible subdivision within the M molecular form could be understood through this process, with the acquisition of inversions leading to the expansion of the M molecular form into new habitat, dividing it into two distinct chromosomal forms.