The Genetic and Cytogenetic Relationships Among Subgenera of Chrysolina Motschulsky, 1860 and Oreina Chevrolat, 1837 (Coleoptera: Chrysomelidae: Chrysomelinae)

International Journal of Zoology and Animal Biology ISSN: 2639-216X MEDWIN PUBLISHERS Committed to Create Value for Researchers

The Genetic and Cytogenetic Relationships among Subgenera of Chrysolina Motschulsky, 1860 and Oreina Chevrolat, 1837 (Coleoptera: Chrysomelidae: Chrysomelinae)

Petitpierre E* Review Article Department of Biology, University of Balearic Islands, 07122 Palma de Mallorca, Spain Volume 4 Issue 3

Received Date: May 03, 2021 Eduard Petitpierre, Department of Biology, University of Balearic *Corresponding author: Published Date: May 28, 2021 Islands, 07122 Palma de Mallorca, Spain, Email: [email protected] DOI: 10.23880/izab-16000305

Abstract

Molecular phylogenetic analyses mainly based on mtDNA and nuDNA sequences and/or, secondarily, on chromosomes and Chrysolina and 19 of its closely related genus Oreina, belonging

plantphylogenetic trophic affiliations,trees showed have monophyly been obtained of Chrysolina-Oreina in 84 species of, with four main clades of subgenera with high support values. to 42 of the 70 described subgeneraChrysolinopsis of the former andand toTaeniochrysea all of the seven, and ones the of second the latter. those Bayesian of Chrysomorpha and Maximum, Euchrysolina likelihood, Melasomoptera and Synerga. The third is much large and composed of three subclades of subgenera, Chrysocrosita and ErythrochrysaThe first clade, Colaphosomaincludes subgenera and Maenadochrysa , and Centoptera, Fasta, the subgenera of Oreina and Timarchoptera. These

p plants, at the root of the core of Chrysolina threesubgenera main ofclades Oreina enclose most species with 2n=24(Xy ) male chromosomes and an ancientCh. trophic Timarchoptera affiliation) withhaemochlora Lamiaceae, a ♂ . However, a host shift from Lamiaceae to Asteraceae was detected in all but one and another from Lamiaceae to Apiaceae in Oreina s. str. and in ( tospecies Apiaceae with was 2n( found)=23(XO) in a subclade chromosomes. with Crositops, The fourth Minckia main andclade Threnosoma holds more, and than the a specieshalf of subgeneraCh. soiota, thisand lastis ambiguous should be for ancestral host-plantTimarchoptera reconstruction, and moved with to Paraheliostola nine botanic families. A statisticallystat. ressur. significant thereby these shift twofor asubgenera second association could not be synonymous as it was recently suggested. New trophic shifts to Hypericaceae were clearly substantiated in Hypericia and excludedSphaeromela from subclade , and to Plantaginaceae in another with L. MedvedevColaphodes , Ovosoma, Palaeosticta and probably Taeniosticha. A further great subclade showing a shift to Asteraceae is that of Allochrysolina, Anopachys, Apterosoma, Chalcoidea, Sibiriella and the species Ch. (Pezocrosita) convexicollis. All species in these latter subclades display high chromosome numbers,

p) in Sphaeromela p) in Hypericia p) in Anopachys, Chalcoidea, Colaphodes, Minckia and Ovosoma ♂ Threnosoma namelywere found 2n=32(Xy in the subclades of Arctolina, 2n=38(Xy, Colaphoptera and, 2n=40(XyPleurosticha, whose species have low chromosome numbers, , and 2n( )=47(XO) in . Possible reversal shifts from Plantaginaceae to Lamiaceae and to Asteraceae p Chrysolina s. str. holds species with a relative polyphagy, ♂ Stichoptera 2n=24(Xy ) or 2n=26(Xyp). Near to these previous, the subcladeChrysolina of due to its high variation in chromosome numbers from on four plant families, and). sharing Finally, 2n(the rising) =23(XO) of some chomosomes. subgenera ofFurthermore, Chrysolina the subclade of , mainly affiliated to Scrophulariaceae,p is an exceptionp among the subgenera of 2n=22(Xy ) to 2n=34(Xy to the genus rank was statistically rejected with

highly significant values. Keywords: Chrysolina; Oreina; DNA; Chromosomes; Phylogenies

The Genetic and Cytogenetic Relationships among Subgenera of Chrysolina Motschulsky, 1860 and Oreina Int J Zoo Animal Biol Chevrolat, 1837 (Coleoptera: Chrysomelidae: Chrysomelinae) 2 International Journal of Zoology and Animal Biology

Introduction Chrysolina and Oreina.

Discussion and Conclusions genus Chrysolina and Amale recent genitalia insightful adult and characters detailed taxonomic[1], has prompted review of us the to set up the possible Motschulsky, genetic interrelationships 1860, based on among morphological many of trees obtained from the combined analysis of mtDNA cox1 and Therrnl Bayesiangenes and and nuDNA Maximum H3 gene, Likelihood have shown phylogenetic three gene sequences [2-5], and both mitochondrial and nuclear distinctive most plesiomorphous clades of species feeding DNAthe described [6], together subgenera, with chromosomaltaking into account studies mitochondrial [7-15], and Ch. Chrysolina (Taeniochrysa) americana and Ch. (Chrysolinopsis) gemina, and its near genus Oreina, 2)on another Lamiaceae with plantsthree species (P=0.98 of the [6]: subgenus 1) that Synerga including Weise, orhost-plant less often affiliations two botanic because families most [16-18]. species of 1900, Ch. coerulans, according with Hsiao and Pasteels [5], are highly specific feeders on one Ch. herbacea Ch. viridana Material and Methods Melasomoptera Bechyné, et al. [32], Ch. (Duftschmid, grossa 1825) and (Küster, Chrysolina 1844),Hsiao andtwo Pasteelsspecies of [5], the and subgenus Ch. lucida 19 species of Oreina of the seven described subgenera have of the subgenus Chrysomorpha(Fabricius, 1792), according withCh. beenA geneticallytotal of 84 speciesanalyzed, of either by DNA of 43 sequences subgenera and/ and cerealis cyaneoaurata (Olivier, 1807), one of the subgenus Euchrysolina BechynéMotschulsky, [32], et Ch. al. graminis[33], selected for building a phylogenetic tree: a partial sequence of (Motschulsky, 1860) and another or by karyotypes. Three differentrrnL), molecular a partial sequence markers of were the cox1), the(Linnaeus, subgenus 1758), Maenadochrysa this latter Bechyné species according[32] Ch. affinis with baetica Hsiao the mitochondrial 16S rDNA ( nuH3) [6]. and Pasteels [5], 3)Ch. a affinis third largemesatlantica clade made of three taxa of mitochondrialWe have also used cytochrome the published c oxidase mitochondrial subunit 1 phylogeneticgene ( Ch. femoralis and a fragment from the nuclear histone 3 gene ( subgenus(Suffrian, 1851),Erythrochrysa Bechyné [32], (Kocher, with 1958)Ch. polita and (Olivier, 1790), close toChrysocrosita them the monotypic Bechyné trees based on the small subunit ribosomal RNA (12S) and [32], with Ch. jakowlewi the large subunit ribosomalcox1) [3], RNAand (16S)that on [5], the that large on the subunit large clade(Linnaeus, but in 1758), an intermediate the Asian subgenus position between the subgenus subunit ribosomal RNA (16S) and the cytochrome c oxidase Maenadochrysa (Weise, 1894), and also in the sameCh. insubunit meiotic 1 and gene mitotic ( testis cells, by classical methods, have (Colaphosoma) sturmi ribosomal RNA (16S) only [4]. The chromosomal analyses this last subgenera, and plus those a doubtful of these and two misleading latter taxa,subclade Chrysolina and composed of Ch. (Centoptera) (Westhoff, bicolor 1882), the unique speciesCh. of Oreinagiven chromosome species, reported numbers, in multiple male sex-chromosome publications [6,7,10]. systems, In (Fasta) fastuosa and in some cases detailed karyotypes of Ch. sturmi which has not (Fabricius, been cytogenetically 1775), are also considered as worth data for phylogenetic trees. (Scopoli, 1763). All species of these three p) chromosomes and similar addition, host-plant affiliations to different botanic families clades, except studiedhave been yet, carefully display analyzed, 2n=24(Xy show meta- or submetacentric implementedBayesian, Neighbour-joining in the different published (NJ), Maximum cladograms Likelihood [2-6]. karyotypes [11,19-20,23], and the nine karyotypes which (ML) and/or Maximum Parsimony (PAUP) trees have been Results groupchromosomes of species, and and among bearing them the aribosomal satellited DNA first locus, autosome very [7,23], that could be a marker chromosome for all these were only chromosomally analyzed but not by DNAs, and Chrysolina among a total number of almost likely. Two further species feeding also on Lamiaceae, that p), Ch. (Bittotaenia) salviae The genetic findings, DNA sequences and/or karyotypes, Ch. (Rhyssoloma) fragariae onones 84 [1], species have of been used for our review. The DNA data of sharing also 2n=24(Xy (Germar, Chrysolina500, belonging are tocompiled 43 subgenera, in 63 species 61% of theand whole the chromosomes 70 described previous1824) and clades of Chrysolina. (Wollaston, 1824) in 69 species, both corresponding to 35 subgenera, and in (7,43,44), should probably join to any one of these three the closely related genus Oreina, 19 species of the seven described subgenera, have been analyzed by DNA sequences a combined analysis of the three sequenced genes [6], and closeIn to the the Maximum latter subclade Likehood of phylogenetic Ch. bicolor and tree Ch. obtained fastuosa by, Chrysolina and Oreina [3-6] and/or chromosomes [7-25]. Moreover, the findings on Oreina, of two different subgenera, Chrysochloa believeplant affiliations that this sample in these is enough species representative of for getting andthere Oreina is another s.str. one (P=0.527 [6], made of five species of reliableare also genetictaken from interrelationships several authors among [26-31]. these Therefore, subgenera we of Hope, 1840 p) chromosomes [8,21], but with an additional , all of them but one sharing karyotypes with 2n=24(Xy Petitpierre E. The Genetic and Cytogenetic Relationships among Subgenera of Copyright© Petitpierre E. Chrysolina Motschulsky, 1860 and Oreina Chevrolat, 1837 (Coleoptera: Chrysomelidae: Chrysomelinae). Int J Zoo Animal Biol 2021, 4(3): 000305. 3 International Journal of Zoology and Animal Biology species of Chrysolina, Ch. (Timarchoptera) haemochlora monotypic Sphaeromela sampled species, Ch. brunsvicensis Ch. ♂ geminata Bedel,Ch. 1899,hyperici the first with four DNA diverse(Gebler, in 1823). size, Thiswith lasttwo speciesautosome displays pairs athreefold highly distinct larger Ch. quadrigemina (Gravenhorst, 1807), Ch. karyotype of 2n( )=27(XO) chromosomes, strikingly varians (Paykull, 1799), (Forster, 1771) and subclade. All these (Suffrian, species 1851), are feeding and the on second Hypericaceae with arethan very any heterogeneous,of the remainders Ch. andhaemochlora lacking the and Y-chromosome two species (Schaller, 1783) [3-6,16,17], constitute a well-defined of[11]. Oreina The onfeeding Apiaceae, affiliations and the of other the species three species in this ofsubclade Oreina other species of the remaining subgenera [16,17], out of on Asteraceae, contrary to all the above mentioned species theplants, Moroccan a botanic Ch. affiliationseriatopora which, of the has monotypic been found subgenus in no Atlasiana Bourdonné et Doguet, 1991 [17], which has not this subclade of at least one species of Chrysolina been genetically studied yet. The chromosomes of the three on Lamiaceae plants [5,6,29]. Therefore, the presence in ones of Oreina Hypericia p) elements, Oreina is a different with genus five mostly metacentrics of medium and small size [8,21], while has induced us to apply an Approximately checked species of share 2n=38(Xy from Chrysolina, and the AU test has given a statistically Ch. varians p Unbiased test [34] to see whether displays 2n=32(Xy ) [8,24]. significant value of 0.016, P < 0.05, rejecting this hypothesis species of the subgenus Chalcoidea Ch. [6], so they might be joined, although most recent authors analisThe five updated DNACh. and/or carnifex cytogenetically sampledCh. still Anotherkeep them new separated and very [1,18,35,36]. heterogeneous main clade, the interstincta Ch. janbechynei Motschulsky, Cobos, 1860, 1953 Ch. curvilinea (Linnaeus, 1767), Ch. (Fabricius,marginata 1792), families so far recorded in Chrysolina. Those on Apiaceae 1758) belong (Suffrian, to a clearly 1851), supported subclade [3,5,6], with plantsfourth one,encloses includes the threesubgenera analyzed affiliated species to allin thethe ninesubgenus plant three(= more species Weise, of 1884) two subgenera,and Ch. (Pezocrosita) (Linnaeus, Threnosoma Ch. helopioides convexicollis Ch. carnifex, 1851), Ch. obscurella Ch. timarchoides, and two a bit more distant to the Chalcoidea species, Ch. one of the subgenusMotschulsky, Crositops 1860, Marseul, 1883, Ch. pedestris (Suffrian,, one (Anopachys) aurichalcea(Jacobson, 1901), very close Ch. to (Anopachys) also in the subgenus Timarchoptera(Suffrian, 1851) and Ch. eurina soiota [6], and three in the subgenus Minckia (Gebler, 1825) and Motschulsky, 1860, (Frivaldszky, 1883). All these species feed on Sulcicollis Ch. oricalcia Chalcoidea p Ch. peregrina Strand,Ch. rufoaenea 1835 (= Asteraceae plantsn [6,16,17], and at least theCh. five aurichalcea species of, J. Sahlberg, 1913), (O.F. Müller,Ch. 1776),soiota, ♂ share 2n=40(Xy♂ ) chromosomes [0,12,14,19,21], feeding on this (Herrich-Schäffer,plant family [6,16,17,29]. 1831) Their and chromosome whereas the cytogenetic findings on (Suffrian, 1851) [5,6], all of them, out of maybe none2n( )=31(XO)of the three to 2n( chromosomally)=45(XO) chromosomes, studied species have provedof the all showing high values, thereby the three quoted species subgenusthe existence Apterosoma of several cryptic species [21,40].Caudatochrysa Although ofnumbers Threnosoma and sex-chromosome♂ systems are differentMinckia, butCh. Bechyné, et al. [32] have been surveyed at the DNA level, Motschulsky, 1860 (= oricalcia p), Ch. peregrina p) and Ch. ♂ share 2n( )= 47(XO), those of rufoanenea p), the highest updated number in the Chrysolina 2n=40(Xy and the subfamily Chrysomelinae, 2n=46(Xy plus the unique their similar cytogenetic results of 2n( )=39(XO) and species of the 2n=50(Xy subgenus Cyrtochrysolina a2n=42(XY) probable chromosomesclose genetic resemblance,[13], and their in trophic agreement affiliation with on Asteraceae (Jolivet and Petitpierre, 1976), point out to Ch. marcasitica p) [8,11,21,23], and Ch. pedestris ♂ Kippenberg (2012), the species of Chalcoidea, Anopachys, Apterosoma (sub Threnosoma, though (Germar, having 1824) two 2n=42(Xy large ring-shaped autosome Caudatochrysa)the proposal by and Bourdonné Allochrysolina and Doguet Bechyné, [ 17] 1950, who joinedin his with 2n( )= 47(XO), as those of the subgenus group 9. In relation with this last subgenus Allochrysolina, the molecular phylogeny of the two analyzed species, Ch. bivalents, very remarkable among the remaining medium or fuliginosa Ch. lepida Timarchopterasmall ones, and a tiny X-chromosome. MikhailovParaheliostola [38], and them in a minor subclade within the large one of Anopachys Medvedev,recently also et al. Bienkowski [38], but [1],the molecular synonymyzed phylogeny the subgenera of their and Chalcoidea (Olivier, subgenera 1807) and[6], and also in (Olivier,accordance 1807) with join an Motschulsky (1860) and L. respective typical species, Ch. haemochlora and Ch. soiota, p), although Ch. does not support this view, because when we applied fuliginosa and Ch. lepida the AU Shimodaira test, the hypothesis of monophyly Inidentical relation chromosome to the subgenus number Pezocrosita, of 2n=42(Xy the DNA analysis of only one species, Ch. convexicollis have distinguishable, may not assume karyotypes that [8].the Paraheliostola stat. ressur. as subgenus should be included in this latter clade, because it is fora good both new species subgenus. was statistically rejected [6]. Therefore, L. Medvedev (1992) should be groups [1], and three or four more species of Pezocrosita at Two further subgenera, Hypericia Bedel, 1899 and the aleast, highly should complex be geneticallytaxon made analyzed of 34 species before in sevencoming different to this

Petitpierre E. The Genetic and Cytogenetic Relationships among Subgenera of Copyright© Petitpierre E. Chrysolina Motschulsky, 1860 and Oreina Chevrolat, 1837 (Coleoptera: Chrysomelidae: Chrysomelinae). Int J Zoo Animal Biol 2021, 4(3): 000305. 4 International Journal of Zoology and Animal Biology conclusion. Two species belonging to the subgenus Allohypericia Bechyné, et al. [32], Ch aeruginosa Ch. perforata between the subgenera Chalcoidea and Hypericia, based on subgenus Colaphoptera (Faldermann,Ch. bigorrensis1835) and Pasteels et al. [42] suggested a possible relationship (Gebler,Ch. 1830), globosa plus probably three more of theCh. between the subgenera Allochrysolina and Anopachys based purpurascens Motschulsky, 1860, their defensive toxins, and Hsiao and Pasteels [5] another (Fairmaire,the previous 1865), one of Arctolina and (Panzer, Pleurosticha 1805), subgenera. and The chromosome (Germar, number 1822), of joinCh. tobigorrensis a sister subclade of on mtDNA phylogeny, host-plant affiliation and chemical 1865) and Ch. globosa , p), agrees defense, and both proposals have not been rejected by our with those of Ch. (Arctolina) poretzkyi (Fairmaire,, Ch. Allochrysolina,AU tests (P= 0.066 Chalcoidea and P=0.215, and Pezocrosita, respectively based [6]. Likewise,on adult (Pleurosticha) gebleri (Panzer, 1805) 2n=26(Xy, Ch. (Pleurosticha) Bourdonné [46] proposed a monophyly for the subgenera lagunovi and Ch. (Pleurosticha) sylvatica(Jacobson, whereas that1897) of Ch. L. Medvedev, 1979 purpurascens crassimargo p), is morphology and host-plant affiliations, an hypothesis quite similar. Moreover, the host-plants of Ch. bigorrensis and bethat possible. could not be statistically rejected by our AU test too Ch. globosa (Germar, 1824),Ch. aeruginosa 2n=24(Xy and (P=0.205) [6]. Therefore, these three phyletic assertions can Ch. perforata Chrysolina of Ch. oirota areand Lamiaceae, Ch. Poretzky but. those of s. str. Ch. bankii Ch. are Lamiaceae and Asteraceae, as well as those costalisThe five checked speciesCh. obsoleta of the Brullé, subgenus 1838), Ch. rufa Motschulsky, 1860,Ch.staphylaea (Fabricius, 1775), include sister minor subclades of species mostly feeding Ch. wollastoni (Olivier, 1807, = Ch. rutilans Wollaston, on PlantaginaceaeAnother well-supported plants, such as subclade those of (P=0.91,the subgenera [6]), (Duftschmid,nec 1825), (Linnaeus, 1758) and Taeniosticha Ch. petitpierrei Bechyné, 1957 (= Ch. reitteri 1864 ♂ Gravenhorst, 1807), constitute a highly supported PalaeostictaMotschulsky, Bechyné, 1860, 1952, Ch. diluta Kippenberg, subclade either by DNA molecular systematics [3,4,6] or by 2004 and Ch. kocheri (Weise, 1884), whose feedingsColaphodes are not species,their 2n( Ch.)= (Lithopteroides)23(XO) chromosomes exanthematica (19,21,23;, whichPetitpierre shares et yet known, Ch. haemoptera (Germar, al., 1988;♂ Petitpierre et al., 2004). Nevertheless, in another Ovosoma1824) and (CodinaCh. Padilla,vernalis 1961),pyrenaica it is related to this previous clade or not, because we should Motschulsky, 1860, Ch. susterai Bechyné, (Linnaeus, 1950, though 1758), andthis also 2n( )= 23(XO) chromosomes, we do not know whether last speciesMotschulsky, has only been 1860, studied chromosomally (Dufour,but not the Canarian endemic species Ch. costalis and Ch. wollastoni by1843) DNA. and The probably chromosomes of the species included in this await to DNA analyses. Moreover, Bienkowski [1] placed within the subgenus Rhyssoloma Ch. diluta p) [21] typical species is the Madeira island endemic Ch. fragariae Ch. haemoptera, Ch. vernalis pyrenaica Wollaston, 1854, whose large subclade are known for , 2n=36(Xy a unique species for this subgenus, according with Bechyné and Ch susterai p (Petitpierre,Another species 1981), that would be possibly included in this that we follow too because Ch. fragariae, apart from being subclade, when ,the all DNAof them sequences with 2n=40(Xy were become) [11,20,21,44]. available, is a[32], Madeiran Bienkowski endemic, [33] has and a very Kippenberg unusual [35],longitudinal a viewpoint relief Ch. (Zeugotaenia) limbata

p) chromosomes [7] instead on Plantaginaceae [16,17], it shows a similar chromosome (Fabricius, 1775), because it feeds ♂ p) [8], and shares some morphological in elytra and it shows 2n=24(Xy Ch. features with the subgenus Taeniosticha [37], sub genus fragariaeof 2n( )= that 23(XO) may chromosomesgive an additional of the stronger above support mentioned to numberCraspeda of Motschulsky 2n=42(Xy , 1860), and the subgenus Palaeosticta thisspecies, view. despite of the absence of DNA findings in [37].

The subgenera Arctolina An additional highly evolved subclade is that of the genetically sampled species, Ch. oirota subgenus Stichoptera Ch. poretzky, and Pleurosticha Kontkanen, 1959, with two have been analyzed, either by their DNA sequences and/or by other ones, Ch. gebleri Lopatin,Ch. 1990, lagunovi and their chromosomes [8,9,12,21,23,39], Motschulsky, 1860, with where a highly eight variable species Motschulsky, [33], with three Ch. sylvatica p) of Ch. kuesteri L. Medvedev, et al. [38] p) p) of Ch. sanguinolenta Mikhailov, et al. [31] and (Gebler, 1823), numbers from 2n=22(Xy (Helliessen, 1912) p) chromosomes, that may allow to a possible p) of Ch. colasi Ch. latecincta displaysynonymisation a remarkable of these DNA two resemblance,subgenera. However, with 2n=26(Xy Ch. oirota to 2n=34(Xy Ch. oceanoripensis (Linnaeus, Bourdonné, 1758), a Doguet modal orand 2n=26 Ch. poretzky (XY value of 2n=24(Xy Ch. ripoceanesis (Cobos, 1952), auct. ), and three Ch. lagunovi species(Demaison, with 1896) high and chromosome numbers, Ch. lucidicollis seem to have a double feeding affiliation and Petitpierre, [39] (= Ch. sylvatica grossepunctata p), and Ch. andon Lamiaceae Ranunculaceae, and Scrophulariaceae,respectively. whereas gypsophilae Ch. rossia Mikhailov, 2006 and have only one on Lamiaceae (Lindberg, 1950) with 2n=30(Xy (Küster, 1845) and (Illiger, 1802) Petitpierre E. The Genetic and Cytogenetic Relationships among Subgenera of Copyright© Petitpierre E. Chrysolina Motschulsky, 1860 and Oreina Chevrolat, 1837 (Coleoptera: Chrysomelidae: Chrysomelinae). Int J Zoo Animal Biol 2021, 4(3): 000305. 5 International Journal of Zoology and Animal Biology

p). A distinctive cytogenetic trait of these Allohypericia, Arctolina and Pleurosticha display feedings in three plant families, witha few 2n=32(Xylarge autosome pairs and the remaining small pairs, (P=0.66) [6], while others such as beingeight karyotypesthe number is oftheir these asymmetry large autosome of autosome pairs sizes, inversely with probably also from a Plantaginaceae ancestor, in reversal correlated with the chromosome numbers [9]. The trophic on Asteraceae, Lamiaceae and Ranunculaceae [15,30,31], Stichoptera is quite eclectic, Chrysolina s. str. are paradigmatic in this sense because mainly on Scrophulariaceae, but at least two species only someshifts (P=0.5of them, in as these Ch. bankii,two cases) can [6].feed The on speciesthree plant of subgenus families affiliation of these species of Ch. Plantaginaceae in adults [6,39]. staphylaea on Lamiaceae, and another one with a secondary choice on at[16]. least, In addition,Asteraceae, Ch. Lamiaceae (Euchrysolina) and Plantaginaceae, graminis has a anddouble Finally, the subgenus Sibiriella on two families, Lamiaceae and Ranunculaceae been cytogenetically studied by two species, Ch. montana as the two sampled species of the subgenus Allohypericia, Ch. schewyrewi L. Medvedev, 1999, has Ch.feeding aeruginosa too, on Asteraceaeand Ch. perforata and Lamiaceae, quoted [16,17], before. asIn well all

p p) chromosomes, respectively (Gebler, 1848) and (Jacobson, 1895), with 2n=40(XYon Asteraceae) and plants 2n=40(Xy are also found in species of the subgenus arethese always examples chosen of by a relativeother congeneric polyphagy, species. there areThus, no most new Chalcoidea[8,11], This whichnumber might of 2n=40 place chromosomes Sibiriella phyletically and their close feeding to speciesplant-affiliations, of Chrysolina-Oreina because these have selected possibly botanic maintained families a Sphaerochrysolina common potential genetic basis in their genomes for plant Ch. umbratilis them. Likewise, the monotypic subgenus inKippenberg the previous [35], subgenus, with we could not (Weise, place it 1887),in a molecular having affiliationsReferences in these botanic families. phylogenetic2n=30(XY) chromosomes tree. However, [14], Sphaerochrysolina but without DNA findings,resembles as morphologically, among others, to the subgenera Chrysolina 1. s.str., Colaphoptera and Cyrtochrysolina Bienkowski AO (2019) Chrysolina of the world. [1], but it isChrysolina strikingly 2. Taxonomic review. Livny: G.V. Mukhametov Publ pp: 920. s.str.different by its karyotype♂ of 2n=30Colaphoptera chromosomes and) and the p Dobler S, Mardulyn P, Pasteels JM, Rowell Rahier chiasmatic) XY y sex-chromosomeCyrtochrysolina system, whereas) chromosomes. chemical defense and life history in the leaf beetle genus p p M (1996) Host-plant switches and the evolution of Displays 2n ( )=23(XO), Sphaerochrysolina 2n=26(Xy Oreina. Evolution 50: 2373-2386. on2n=24(Xy Asteraceae plants [29], is clearly 2n=42(Xy different from those Furthermore, the trophic affiliation of 3. Mitochondrial DNA phylogeny and host plant use in palearctic Chrysolina Ranunculaceae and Apiaceae [6]. Garin CF, Juan C, Petitpierre E (1999) of the above subgenera on Lamiaceae, Plantaginaceae, (Coleoptera, Chrysomelidae). Journal of Molecular Chrysolina-Oreina Evolution 48: 435-444. Although Lamiaceae are the first plant-affiliation for Mitochondrial 16S rDNA sequences and their use the genus (P=0.98) [6] the species of 4. Gómez Zurita J, Garin CF, Juan C, Petitpierre E (1999) tohave new extended host-plants their families trophism have tonot eightbeen otherunique botanicevents families [6,15,16,17]. The different shifts from Lamiaceae as phylogenetic markers in leaf-beetles with special subgenera Hypericia and Sphaeromela [3,6] namely, that reference to the subfamily Chrysomelinae. In: Cox ML, et for each of them, except for Hypericaceae selected by the 5. al. (Eds.), Advances in Chrysomelidae Biology pp: 25-38. grouping Oreina s.str. and Ch. (Timarchoptera) haemochlora Chrysolina – Oreina from Lamiaceae to Apiaceae, has taken placeMinckia in a clade and Hsiao TH, Pasteels JM (1999) Evolution of host-plant Threnosoma, and Ch. (Timarchoptera) soiota affiliation and chemical defense in (P=0.37) [6], in a subclade with the subgenera leaf beetles revealed by mtDNA phylogenies. In: Cox ML, Oreina subgenus Chrysochloa (P=0.62) [6]. et al. (Eds.), Advances in Chrysomelidae Biology pp: 321- withAlso, the that subgenera from Lamiaceae Allochrysolina, to Asteraceae Anopachys, in a Apterosoma, subclade of 6. 342. Chalcoidea and Pezocrosita (P=0.54)(Ch. convexicollis) [6] and in another to the molecular systematics and the evolution of host- plantJurado associations Rivera JA, Petitpierre in the genus E (2015) Chrysolina New contributions has put within his “genus” Chalcoidea (P=0.94) (Coleoptera, subgenera[46]. In agreement Allochrysolina, with this Chalcoidea last subclade, s.str. and Bourdonné Pezocrosita [46]. It 7. is interesting to note that some subgenera Motschulsky, subclades, such the Chrysomelidae, Chrysomelinae). ZooKeys 547: 165-192. as that of Stichoptera, shows a double trophic shift from Petitpierre E (1988) A new contribution to the cytogenetics and cytotaxonomy of the Chrysomelinae (Coleoptera, Chrysomelidae). Cytobios 54: 153-159. possibly Plantaginaceae to Scrophulariaceae and Lamiaceae Petitpierre E. The Genetic and Cytogenetic Relationships among Subgenera of Copyright© Petitpierre E. Chrysolina Motschulsky, 1860 and Oreina Chevrolat, 1837 (Coleoptera: Chrysomelidae: Chrysomelinae). Int J Zoo Animal Biol 2021, 4(3): 000305. 6 International Journal of Zoology and Animal Biology

8. 20. of Chrysolina Mots. and Oreina Chrysomelidae,Petitpierre E (1999) Chrysomelinae). The cytogenetics Hereditas and cytotaxonomy 131: 55-62. Petitpierre E (1978) Chromosome numbers and Chevr. (Coleoptera, 31:sex-determining 219-223. systems in fourteen species of 9. Chrysomelinae (Coleoptera, Chrysomelidae). Caryologia of the subgenus Stichoptera of Chrysolina 21. Petitpierre E (2000) The cytogenetics and cytotaxonomySome Chrysolina Oreina Aspects on the Insight on the Insect Biology(Coleoptera,. Narendra Petitpierre E (1981) New data on the cytology of PublishingChrysomelidae). House, In: Delhi, Sobti India, RC, Yadav pp: 181-187. JS, et al. (Eds.), (Mots.) and (Chevr.) (Coleoptera, 22. Chrysomelidae). Genetica 54: 265-272. 10. nine species of the genus Chrysolina chromosomal evolution of Chrysomelinae revisited Petitpierre E (1983) Karyometric diffrences among Petitpierre E (2011) Cytogenetics, cytotaxonomy 157: 67-79. and Cytology 25: 33-39. Mots. (Coleoptera, Chrysomelidae). Canadian Journal of Genetics and 11. (Coleoptera: Chrysomelidae). ZooKeys 23. Chrysolina some representatives of the family Chrysomelidae Petitpierre E, Mikhailov, Yu E(2009) Chromosomal Barabás L, Bezo M (1978) Chromosome count in evolution and trophic affiliation in the genus (Coleoptera: Chrysomelidae). In: Jolivet P, Santiago Blay (Coleoptera). Biologia (Bratislava) 33: 621-625. J, Schmitt M, et al. (Eds.), Research on Chrysomelidae 12. Volume. 2. Brill, Leiden-Boston, pp: 225-234. 24. Barabás L, Bezo M (1979) Prispevok k cytotaxonómii analysis on twelve species of the subfamily Chrysomelinae listaviek (Coleoptera, Chrysomelidae). Biologia Petitpierre E, Mikhailov, Yu E (2013) A chromosomal 25. (Bratislava) 34: 845-850. 61: 193-198. (Coleoptera, Chrysomelidae). Folia Biologica (Kraków) particularlyPetitpierre E,that Segarraof Chrysomelinae C (1985) and Chromosomal Palearctic 13. variability and evolution of Chrysomelidae (Coleoptera) chromosomal analysis of the Chrysolina angusticollis Petitpierre E, Aiko N, Hattori K, Katakura H (2001) A 26. Alticinae. Entomography 3: 403-426. Chromosome Science Chrysolina species complex (Coleoptera, Chrysomelidae). fromBienkowski the Russia AO (2010) and theReview European of the leafcountries beetle ofgenus the 4: 117-120. Motschulsky (Coleoptera, Chrysomelidae) wings. Entomological Review 90: 885-902. 14. ChrysolinaPetitpierre, E, Kippenberg H, Mikhailov Y, Bourdonné former USSR: I. A key to species with developed hind JC (2004) Karyology and cytotaxonomy of the genus 27. Motschulsky (Coleoptera, Chrysomelidae). Chrysolina 15. Zoologischer Anzeiger 242(4): 347-352. fromBienkowski the Russia AO (2011) and theReview European of the leafcountries beetle ofgenus the Motschulsky (Coleoptera, Chrysomelidae) Mikhailov, Yu E (2011) Interrelation of throphic and wings. Entomological Review chromosomal evolution in leaf beetles (Coleoptera, former USSR: II. A key to species with the reduced hind 16. Chrysomelidae). Lesnoy Vestnik 4: 13-20. 28. 91: 645-654. des Chrysolina typesJolivet deP, Petitpierre sélection Etrophique. (1976) Les Annales plantes-hôtes de la connuesSocieté Jolivet P, Petitpierre E, Daccordi M (1986) Les plantes- (Col. Chrysomelidae). Essai sur les hôtes des Chrysomelidae. Quelques nouvelles précisions et additions (Coleoptera). Nouvelle Revue 17. Entomologique de France (N.S.) 12: 123-149. 29. Entomologique 3: 341-357. biosystématique des Chrysolina l.s. Chrysomelidae:Bourdonné JC, Chrysomelinae). Doguet S (1991) Annales Données de la Societé sur la 3.Kippenberg Supplementband. H (1994) 88 Familie Chrysomelidae. In: (Coleoptera: Lohse GA, Lucht W, et al. (Eds.), Die Käfer Mitteleuropas. 30. Goecke & Evers, Krefeld pp: 17-92. 18. entomologique de France (N.S.) 27: 29-64. Supplement Lopatin I K, Mikhailov, Yu E (2010) Contribution to WinkelmanRutilans. Association J, Debreuil des Coléopteristes M (2008) Les Amateurs Chrysomelinae du Sud the knowledge of the leaf-beetle fauna (Coleoptera, de la France France. (Coleoptera, Chrysomelidae). Chrysomelidae) of Eastern Kazakhastan. Entomological 31. Review 90: 877-884. 19. Villelongue- dels-Monts pp: 188. Chrysolina sylvatica Chrysolina species of the genus Chrysolina subcostataMikhailov Yu E (2006) On the types andPleurosticha status of Petitpierre E (1975) Notes on chromosomes of ten (Gebler, 1823) and Mots. (Coleoptera: (Gebler, 1848) from the subgenus PetitpierreChrysomelidae). E. The Genetic Genetica and Cytogenetic45: 349-354. Relationships among Subgenera ofMotschulsky, 1860 (Coleoptera,Copyright© Chrysomelidae). Petitpierre E. Chrysolina Motschulsky, 1860 and Oreina Chevrolat, 1837 (Coleoptera: Chrysomelidae: Chrysomelinae). Int J Zoo Animal Biol 2021, 4(3): 000305. 7 International Journal of Zoology and Animal Biology

et considérations sur le sous-genre Stichoptera

eme 32. Eurasian Entomological ContributionJournal 5: 292-302. à la connaissance du genre Chrysolina Motschulsky, 1860 (Coleoptera, Chrysomelidae). BechynéChrysomelidae). J (1950) Entomologische 7 Arbeiten aus dem Nouvelle Revue d’Entomologie (N.S.) 29: 33-52. Motsch. (Col. Phytophaga in the Chrysolina aurichalcea 40. Fujiyama S, Okamoto K (1996) Two new karyotypes 33. Museum Grey 3: 351-385. Chrysolina (Mannerheim) complex (Coleoptera. Chrysomelidae), 31: 203-209. their relationship and Bienkowskisubgenera, AOspecies, (2001) Asubspecies, study of the genusand synonyms distribution in the Japanese archipelago. Applied Motschulsky, 1860, with a checklist of all the described Entomology and Zoology

(Coleoptera: Chrysomelidae: Chrysomelinae). Genus 41. Gómez-Zurita J, Hunt T, Kopliku F, Vogler AP (2007) (Wroclaw) 12: 105-235. andRecalibrated their insect tree herbivores. of Leaf Plos Beetles One (Chrysomelidae) for the problem of regions using the scale bootstrap indicates independent diversification of angiosperms 34. resamplings.Shimodaira H Technical (2000) Another report No.calculation 2000-35, of Department the p-value 2(4): E360. of Statistics, Stanford University, California. 42. Pasteels JM, Termonia A, Daloze D, Windsor DM (2003) 35. Distribution of toxins in chrysomeline. Proceedings leaf beetles: of the Fifthpossible International taxonomics inferences.Syposium In:on Furththe Chrysomelidae,DG, et al. (Eds.), Kippenberg H (2010) Subfamily Chrysomelinae Latreille, Special Topics in Leaf Beetle Biology 1802. In: Löbl I, Smetana A, et al. (Eds.), Catalogue of Sophia-Moscow, pp: 261-275. Palaearctic Coleoptera Volume 6 Chrysomeloidea. Apollo Iguassu Falls (Brazil), August 2000, Pensoft Publishers, 36. Books, Stenstrup pp: 390-443. Palaearctic Chrysomelidae.

Warchalowski629. A (2010) The 43. Petitpierre E (1982) Chromosomal findings on 22 Volume 1. Natura optima dux Foundation, Warszawa, pp: species of Chrysomelinae (Chrysomelidae: Coleoptera). 37. Chromosome Information Service 32: 22-24. Taeniosticha Craspeda 10 Chromosome numbers and meioformulae of Bourdonné JC (2005) Révision du sous-genreère 44. Petitpierre E, Segarra C, Yadav JS, Virkki N (1988) partie. Nouvelle Motschulsky, Revue d’Entomologie 1860 du genre 21: 297- 353.Motschulsky, 1860 (Coleoptera, Chrysomelidae) Chrysomelidae. In: Jolivet P, Petitpierre E, Hsiao TH, et (N.S.) al. (Eds.), Biology of Chrysomelidae. Kluwer Academic 38. Publishers, Dordrecht / Boston / London, pp: 161-186. Timarchoptera Paraheliostola MikhailovMedvedev, Yu1992 E (2002) of the Consideration genus Chrysolina of the subgenera 45. Yosida T (1944) Chromosome studies in the Coleoptera. Motschulsky, 1860 and L. I. Japanese Journal of Genetics 20: 107-115. Motschulsky (Coleoptera, Chrysomelidae) after description of two 46. Bourdonné JC (2012) Révision du genre Chalcoidea new forms from Khakassia. Eurasian Entomological Motschulsky, 1860 avec description d’une nouvelle 39. Journal 1: 219-228. Chrysolina espèce Chalcoidea (s. str.) hollandei (1ère partie) (Stichoptera) oceanoripensis nova species, endémique (Coleoptera, Chrysomelidae, Chrysomelinae). Nouvelle Bourdonné JC, Doguet S, Petitpierre E (2013) Revue d’Entomologie (N.S.) 28: 99-181.

des dunes francaises des Landes de Gascogne