Evolutionary relationships of the ruficornis and aeneus groups ofspecies of the genus Merodon Meigen, 1803 (Diptera: Syrphidae)

Abstract

<p>U radu, metodom PAGE (poliakrilamid gel elektroforeze), analizirana je gensko-enzimska<br />varijabilnost 11 populacija vrsta ruficornis grupe: M. ruficornis, M. armipes, M. crymensis, M. loewi i M.<br />recurvus; 11 populacija vrsta aeneus grupe: M. aeneus: M. aeneus A, M. aeneus B, M. aeneus C, M.<br />cinereus A, M. cinereus B, M. funestus i M. desuturinus; 7 populacija avidus grupe (M. avidus A i M.<br />avidus B) roda Merodon i 4 populacije vrste Cheilosia vernalis sa teritorije Balkanskog poluostrva.<br />Analizirana je varijabilnost alozima determinisanih alelima 17 lokusa (Aat, Fum, Gpd-1, Gpd-2, Gpi, Had,<br />Hk-2, Hk-3, Idh-1, Idh-2, Mdh-1, Mdh-2, Me, Pgm, Sod-1, Sod-2, Sod-3) vrsta ruficornis grupe, 15 lokusa<br />(Aat, Fum, Gpd-2, Gpi, Had, Hk-2, Hk-3, Idh-2, Mdh-1, Mdh-2, Me, Pgm, Sod-1, Sod-2, Sod-3) aeneus<br />grupe, 16 lokusa (Aat, Ao, Fum, Gpd-2, Gpi, Had, Hk-2, Hk-3, Idh-2, Mdh-1, Mdh-2, Me, Pgm, Sod-1,<br />Sod-2, Sod-3) avidus grupe i 12 lokusa (Fum, Gpd-2, Gpi, Had, Hk-2, Hk-3, Idh-1, Idh-2, Mdh-1, Mdh-2,<br />Pgm, Sod-1) populacija vrste Ch. vernalis.<br />Populaciono-genetičkom analizom vrsta ruficornis grupe utvrđen je dijagnostički značaj Aat,<br />Fum, Had, Hk-2, Hk-3, Mdh-2, Me, Pgm i Sod-1 lokusa. Species-specifičnim alelima identifikovane su<br />vrste i formiran genetičko-biohemijski dihotomi ključ. Analizom alozimske varijabilnosti populacija vrsta<br />aeneus grupe registrovani su kriptični taksoni: M. aeneus A, M. aeneus B, M. aeneus C, M. cinereus A i<br />M. cinereus B. U simpatričkim i alohronim populacijama vrsta M. aeneus A i M. aeneus C registrovani su<br />dijagnostički Had, Sod-1, Me, Aat i Pgm lokusi. Analizom PGM zimograma, u okviru prethodno<br />definisane &quot;prolećne generacije&quot; determinisana je populacija sa Kopaonika taksona M. aeneus B aeneus<br />kompleksa. Determinacija kriptičnih taksona M. cinereus A i M. cinereus B izvr&scaron;ena je na osnovu speciesspecifičnih<br />genotipova Had lokusa. Najveći broj dijagnostičkih lokusa registrovan je između vrsta M.<br />desuturinus i M. funestus, kao i između navedenih vrsta i ostalih vrsta aeneus kompleksa. Utvrđen je i<br />dijagnostički značaj većine analiziranih lokusa: Aat, Fum, Gpd-2, Hk-2, Hk-3, Idh-2, Mdh-2, Me, Pgm i<br />Sod-1. Na osnovu genetičkih markera Aat i Idh-2 lokusa i dijagnostičkih morfolo&scaron;kih karaktera<br />identifikovane su sestrinske vrste M. avidus A i M. avidus B.<br />Stepen genetičke diferencijacije između vrsta u okviru ruficornis i aeneus grupa bio je veći u<br />odnosu na blisko srodne vrste aeneus i cinereus kompleksa i sestrinskih vrsta avidus grupe. Na genetičku<br />identičnost ukazuje 52,94% (ruficornis grupa) i 55,56% (aeneus grupa) vrednosti genetičke bliskosti po<br />lokusu, odnosno, potpuna genetička različitost registrovana je u 25,41% (ruficornis grupa) i 25,49%<br />(aeneus grupa) analiza.<br />Na osnovu prosečne genetičke bliskosti i klaster analize u okviru ruficornis grupe diferencirana je<br />grupa blisko srodnih vrsta M. armipes, M. ruficornis i M. recurvus u odnosu na genetički udaljene vrste,<br />M. crymensis i M. loewi. Dendrogramom genetičkih odnosa između vrsta aeneus grupe formirana je<br />monofiletska grupa vrsta aeneus i cinereus kompleksa, nasuprot genetički udaljenim vrstama M. funestus i<br />M. desuturinus.<br />Utvrđena je veća zastupljenost vrednosti genetičke bliskosti po lokusu, mere genetičke<br />identičnosti (37,44%) i manji procenat pokazatelja genetičke različitosti (47,80%) između vrsta aeneus i<br />avidus grupa u odnosu na vrste ruficornis i avidus grupe (29,84% i 58,09%). Na visok stepen genetičkih<br />razlika između kongeneričkih vrsta roda Merodon ukazuje najveća procentualna zastupljenost vrednosti<br />genetičke bliskosti po lokusu, pokazatelja genetičke različitosti (62,01%) i mali stepen genetičke<br />identičnosti (26,84%).<br />Utvrđen je pleziomorfan karakter Gpi (Gpijl), Hk-2, Hk-3 (Hkc) i Mdh-2 (Mdh-2e) lokusa vrsta<br />ruficornis, aeneus i avidus grupa roda Merodon, kao i predački aleli Gpii, Gpij, i Pgmf vrsta avidus i<br />aeneus grupa. Poređenjem alozima determinisanih alelima 10 lokusa populacija vrste Cheilosia vernalis,<br />suprageneričke out vrste, i populacija vrsta roda Merodon, registrovani su identični aleli samo u Fum i<br />Pgm lokusima.</p>

<p>Gene-enzyme variability of the ruficornis (11 populations: M. ruficornis, M. armipes, M.<br />crymensis, M. loewi and M. recurvus), aeneus (11 populations: M. aeneus: M. aeneus A, M. aeneus B, M.<br />aeneus C, M. cinereus A, M. cinereus B, M. funestus and M. desuturinus) and avidus (7 populations: M.<br />avidus A and M. avidus B) groups of species of the genus Merodon and four Cheilosia vernalis<br />populations from the Balkan peninsula was analyzed using PAGE (polyacrilamide electrophoresis).<br />Allozyme variability of 17 loci in the ruficornis species group (Aat, Fum, Gpd-1, Gpd-2, Gpi, Had, Hk-2,<br />Hk-3, Idh-1, Idh-2, Mdh-1, Mdh-2, Me, Pgm, Sod-1, Sod-2, Sod-3), 17 loci in the aeneus species group<br />(Aat, Fum, Gpd-2, Gpi, Had, Hk-2, Hk-3, Idh-2, Mdh-1, Mdh-2, Me, Pgm, Sod-1, Sod-2, Sod-3), 16 loci in<br />the avidus species group (Aat, Ao, Fum, Gpd-2, Gpi, Had, Hk-2, Hk-3, Idh-2, Mdh-1, Mdh-2, Me, Pgm,<br />Sod-1, Sod-2, Sod-3) and 12 loci in Ch. vernalis populations (Fum, Gpd-2, Gpi, Had, Hk-2, Hk-3, Idh-1,<br />Idh-2, Mdh-1, Mdh-2, Pgm, Sod-1) was evaluated.<br />Diagnostic value of Aat, Fum, Had, Hk-2, Hk-3, Mdh-2, Me, Pgm and Sod-1 loci was determined<br />by the population-genetic analysis of the ruficornis species group. The species were identified using<br />species-specific alleles and genetic-biochemical key was formed. Allozyme variability analysis of the<br />aeneus species group populations enabled discrimination of the cryptic taxa: M. aeneus A, M. aeneus B,<br />M. aeneus C, M. cinereus A and M. cinereus B. In sympatric and alochronic populations of the species M.<br />aeneus A and M. aeneus diagnostic loci were observed (Had, Sod-1, Me, Aat and Pgm). Analysis of the<br />PGM zymogram allowed the identification of the cryptic species M. aeneus B in the previously delineated<br />&quot;spring generation&quot; from Kopaonik. Cryptic taxa M. cinereus A and M. cinereus B were discriminated<br />based on the species-specific genotypes at the Had locus. The largest numbers of the diagnostic loci were<br />registered for differentiating M. desuturinus and M. funestus species, and between these two and other<br />species of the aeneus complex. Diagnostic value was recorded for the analyzed loci: Aat, Fum, Gpd-2, Hk-<br />2, Hk-3, Idh-2, Mdh-2, Me, Pgm and Sod-1. Sibling species M. avidus A and M. avidus B were identified<br />using genetic markers of Aat and Idh-2 loci and diagnostic morphological characters.<br />The degree of genetic differentiation between the species of the ruficornis and the aeneus groups<br />was higher comparing to closely related species of the aeneus and the cinereus complexes and sibling<br />species of the avidus group. Out of the performed analyses, 52.94% (ruficornis group) and 55.56%<br />(aeneus group) of the genetic identity values for loci point to genetic identity, while complete genetic<br />difference was registered in 25.41% (ruficornis group) and 25.49% (aeneus group).<br />Average values of the genetic identity and cluster analysis enabled differentiating the groups of<br />closely related (M. armipes, M. ruficornis and M. recurvus) and genetically distant species (M. crymensis<br />and M. loewi) in the ruficornis group. Based on dendrogram of genetic relationships between the species<br />of the aeneus group, monophyletic group of the aeneus and the cinereus complex species was formed, as<br />opposed to genetically distant species M. funestus and M. desuturinus.<br />Genetic identity among loci between the aeneus and the avidus groups was higher (37.44%) while<br />genetic distance was lower (47.80%) in comparison to the corresponding values for the ruficornis and the<br />avidus groups of species (29.84% and 58.09%). High values of genetic difference (62.01%) and low<br />values of genetic identity (26.84%) indicate great genetic difference between congeneric species of the<br />genus Merodon.<br />Pleziomorphous character of Gpi (Gpijl), Hk-2, Hk-3 (Hkc) and Mdh-2 (Mdh-2e) loci in the<br />ruficornis, aeneus and avidus species groups of the genus Merodon and ancestral alleles (Gpii, Gpij, Pgmf<br />) in the avidus and the aeneus species groups were registered.<br />Comparison of allozymes of the suprageneric out species Ch. vernalis and the populations of the<br />genus Merodon revealed identical alleles only in Fum and Pgm loci.</p>