Cytogenetics, cytotaxonomy and chromosomal evolution of Chrysomelinae revisited (Coleoptera, Chrysomelidae)

Nearly 260 taxa and chromosomal races of subfamily Chrysomelinae have been chromosomally analyzed showing a wide range of diploid numbers from 2n = 12 to 2n = 50, and four types of male sex-chromosome systems. with the parachute-like ones Xy(p) and XY(p) clearly prevailing (79.0%), but with the XO well represented too (19.75%). The modal haploid number for chrysomelines is n = 12 (34.2%) although it is not probably the presumed most plesiomorph for the whole subfamily, because in tribe Timarchini the modal number is n = 10 (53.6%) and in subtribe Chrysomelina n = 17 (65.7%). Some well sampled genera, such as Timarcha, Chrysolina and Cyrtonus, are variable in diploid numbers, whereas others, like Chrysomela, Paropsisterna, Oreina and Leptinotarsa, are conservative and these differences are discussed. The main shifts in the chromosomal evolution of Chrysomelinae seems to be centric fissions and pericentric inversions but other changes as centric fusions are also clearly demonstrated. The biarmed chromosome shape is the prevalent condition, as found in most Coleoptera, although a fair number of species hold a few uniarmed chromosomes at least. A significant negative correlation between the haploid numbers and the asymmetry in size of karyotypes (r = -0.74) has been found from a large sample of 63 checked species of ten different genera. Therefore, the increases in haploid number are generally associated with a higher karyotype symmetry.     

Phylogeographic assessment of mtDNA paraphyly and the evolution of unisexuality in Calligrapha (Coleoptera: Chrysomelidae)

The leaf beetle genus Calligrapha is one of the few examples of animals with several obligate unisexual, female‐only species. Previous work showed that each one arose independently from interspecific hybridization events involving different species. However, all of them clustered in a single mtDNA clade together with some individuals of the parental bisexual species, which appeared as deeply polyphyletic in the mtDNA genealogy of the genus. The dating of these splits using a molecular clock placed them in the Quaternary and it was hypothesized that climatic change during this period may have favored range expansions and secondary contacts required for hybridization. In this work, we test this hypothesis and the origins of unisexuality in Calligrapha examining the diversity of mitochondrial (cox1) and nuclear (wingless, Wg) genes and the Bayesian continuous mtDNA phylogeography of a sample of more than 500 specimens of two bisexual species of Calligrapha at a continental scale and two unisexual species derived from them. Besides a major topological difference, whereby each bisexual species is monophyletic for Wg but paraphyletic for cox1, both gene datasets are consistent with a minimum of seven evolutionary lineages, coherent with geography and consistent with an ordered expansion to occupy their current ranges. The results also imply their survival in well‐established glacial refuges during the Last Glacial Maximum (LGM). Thus, for bisexual C. multipunctata there are two main, southern and northern lineages. The southern lineage expanded its range in two evolutionary branches, to the Rocky Mountains and to the northern Mississippi and Ohio River basins, respectively. The northern lineage has one branch in the Upper Mississippi and one that expanded west to the Pacific Northwest and east to the northeastern North Atlantic, finding refuge in both areas. These major lineages are parapatric in the Northern Great Plains, an area consistent with them having found refuge in the so‐called Driftless region during the LGM. For bisexual C. philadelphica, one northern lineage expanded west from the northern Appalachians and one east and southwest along the axis of the Appalachians, and the timing of events is consistent with their persistence in glacial refugia at both sides of the main Great Lakes lobe of the Laurentide Ice Sheet. There is evidence that the northeastern North Atlantic lineages of both species hybridized at the edge of their ranges after the LGM. The additional, divergent mtDNA lineage of these species shows evidence of range expansions of two lineages, one for each species, in an area south of the Laurentide Ice Sheet, and giving origin to the unisexual species by way of hybridization with other species in the Alleghanian region after the LGM. Interestingly, the individuals of supposedly bisexual species in this clade are all females. This suggests that unisexuality actually predates the origin of unisexual taxa in this system and that some bisexual species in Calligrapha may be species complexes instead, with cryptic species differing in their reproductive mode.     

广西叶甲亚科昆虫种类记述(鞘翅目:叶甲科)

记述中国科学院动物研究所 1 998～ 2 0 0 0年对广西十万大山及大瑶山地区考察所采获叶甲亚科并该所标本馆馆藏叶甲亚科标本及文献记载 ,共 2 0属 35种 ,其中包括1中国新记录属 Pseudolina Jacoby、1新种 Gonioctena( Brachyphytodecta) oculata sp.nov.、2中国新记录种 Pseudolina indica Jacoby,Paropsides bouveri Chen。模式标本保存在中国科学院动物研究所。     

The genera of Chrysomelinae (Coleoptera: Chrysomelidae) in Costa Rica

Keys in Spanish and English are given for the genera of Chrysomelinae known from Costa Rica. For each genus, a list of species compiled from collections in the University of Costa Rica, the National Biodiversity Institute, and the entomological literature is presented. The genus Planagetes Chevrolat 1843 is recorded for the first time from Central America, and the genus Leptinotarsa St?l 1858 is synonymized with Stilodes Chevrolat 1843.     

The morphology of larvae and systematics of the leaf-beetles Chrysolina tundralis and Chrysolina roddi (Coleoptera, Chrysomelidae, Chrysomelinae)

The 1st- and 4th-instar larvae of Chrysolina tundralis and Ch. roddi and the egg of the latter species are described for the first time; the instar-related changes of the larval morphology are discussed. The 4th-instar larva of Ch. tundralis is very similar to that of Ch. septentrionalis, but differs in the smaller sclerite-like plates of the abdominal segments and wider spaces between them (4–7 times as wide as a sclerite-like plate). The 4th-instar larva of Ch. roddi is very similar to that of Ch. pedestris, but differs in the dark brown coloration of the body and a fewer number of setae (9–12) in the dorsolateral areas of the meso- and metathorax. Data on the habitats and host plants of the larvae are given.     

Assessment of the role of Wolbachia in mtDNA paraphyly and the evolution of unisexuality in Calligrapha (Coleoptera: Chrysomelidae)

Calligrapha is a New World leaf beetle genus that includes several unisexual species in northeastern North America. Each unisexual species had an independent hybrid origin involving different combinations of bisexual species. However, surprisingly, they all cluster in a single mtDNA clade and with some individuals of their parental species, which are in turn deeply polyphyletic for mtDNA. This pattern is suggestive of a selective sweep which, together with mtDNA taxonomic incongruence and occurrence of unisexuality in Calligrapha, led to hypothesize that Wolbachia might be responsible. I tested this hypothesis studying the correlation between diversity of Wolbachia and well‐established mtDNA lineages in >500 specimens of two bisexual species of Calligrapha and their derived unisexual species. Wolbachia appears highly prevalent (83.4%), and fifteen new supergroup‐A strains of the bacteria are characterized, belonging to three main classes: wCallA, occupying the whole species ranges, and wCallB and wCallC, narrowly parapatric, infecting beetles with highly divergent mtDNAs where they coexist. Most beetles (71.6%) carried double infections of wCallA with another sequence class. Bayesian inference of ancestral character states and association tests between bacterial diversity and the mtDNA genealogy show that each mtDNA lineage of Calligrapha has specific types of infection. Moreover, shifts can be explained by horizontal or vertical transfer from local populations to an expanding lineage and cytoplasmic incompatibility between wCallB and wCallC types, suggesting that the symbionts hitchhike with the host and are not responsible for selective mtDNA sweeps. Lack of evidence for sweeps and the fact that individuals in the unisexual clade are uninfected or infected by the widespread wCallA type indicate that Wolbachia does not induce unisexuality in Calligrapha, although they may manipulate host reproduction through cytoplasmic incompatibility.     

Subsocial Neotropical Doryphorini (Chrysomelidae,Chrysomelinae): new observations on behavior,host plants and systematics

A summary of literature, documented observations and field studies finds evidence that mothers actively defend offspring in at least eight species and three genera of Neotropical Chrysomelinae associated with two host plant families. Reports on three Doryphora species reveal that all are oviparous and feed on vines in the Apocyanaceae. Mothers in the two subsocial species defend eggs and larvae by straddling, blocking access at the petiole and greeting potential predators with leaf-shaking and jerky advances. A less aggressive form of maternal care is found in two Platyphora and four Proseicela species associated with Solanaceae, shrubs and small trees. For these and other morphologically similar taxa associated with Solanaceae, genetic distances support morphology-based taxonomy at the species level, reveal one new species, but raise questions regarding boundaries separating genera. We urge continued study of these magnificent insects, their enemies and their defenses, both behavioral and chemical, especially in forests along the eastern versant of the Central and South American cordillera.     

A cytogenetic study on three Chilean species of Chrysomelinae (Coleoptera, Chrysomelidae)

Three species of Chilean leaf beetles were chromosomally analyzed. The endemic Araucanomela wellingtonensis displays a male meioformula of 13 + Xyp with 2n = 28 chromosomes and an asymmetric karyotype with two large autosome pairs and 12 medium/small pairs of autosomes and sex-chromosomes, a diploid number which had not been found among the other species of the subtribe Paropsina sensu lato studied to date. Strichosa eburata presents a meioformula of 11 + Xyp, 2n = 24 chromosomes, as occurs in many species of chrysomelines belonging to different subtribes. Furthermore, Phaedon cyanopterum has a 16 + Xyp meioformula, that is 2n = 34 chromosomes, of small size mostly, also in agreement with the karyological findings obtained in all the other congeneric species so far examined. These cytogenetic data are discussed with respect to the previous ones in this subfamily and with other characters of taxonomic and evolutionary value.     

叶甲亚科一新属及两新种记述（鞘翅目：叶甲科）

记述中国叶甲亚科1新属:玉龙叶甲属Yulongedon gen.nov.,包括2新种:靓玉龙叶甲Y.formosus sp.nov.,宝藏神玉龙叶甲Y.jambhalai sp.nov.。     

The mesothoracic intercoxal perforation in Chrysomelinae and its evolutionary significance (Insecta: Coleoptera: Chrysomelidae)

In the thorax of many beetles the posterior parts of the mesoventrite and anterior parts of the metaventrite are flexed inward to form mesocoxal cavities laterally and a transverse double-walled infolding medially. This study focuses on the structure of the median part of the infolding in Chrysomelinae (=Timarchini + Chrysomelini; 42 species studied). Whereas the infolding is intact in Timarchini and very few Chrysomelini, it is almost always perforated in Chrysomelini. The perforation connects the body cavity anterior and posterior to it. It is usually small or absent in wingless Chrysomelinae, but there are exceptions. The phylogenetic interpretation of the intercoxal perforation in the entire Cucujiformia is unclear: among Chrysomelidae, the perforation was found in a representative of Hispinae, but was absent in the cryptocephaline and criocerine species examined; among non-chrysomelid Cucujiformia the perforation was found in an anthribid, but was absent in the examined tenebrionid and erotylid species. From a functional perspective, the perforation likely reduces the movability of the intercoxal area. The origin of this perforation appears as an unusual type of evolutionary structural transformation where ± widely separated (but opposed) parts of the body wall belonging either to two external projections or to a single infolding (depending on the point of view) became attached to each other along their external surfaces, and then dissolved. As a result, the parts of the body cavity on both sides of the perforated walls came to communicate in a new place.     

A new species of the genus Chrysolina Motschulsky, 1860 from Turkey (Coleoptera: Chrysomelidae: Chrysomelinae)

A new leaf beetle species, Chrysolina (Lopatinica) kabalaki sp. n., from Northeast Turkey is described and compared with closely related species. Habitus and male genitalia of the species are photographed. A key is provided for the species.

http://www.zoobank.org/urn:lsid:zoobank.org:pub:89CEE108-138D-4D38-837D-A8C3682BB773     

Revision of the Eastern Asian genera Ambrostoma Motschulsky and Parambrostoma Chen (Coleoptera: Chrysomelidae: Chrysomelinae)

The leaf beetle genera Ambrostoma Motschulsky, 1860 and Parambrostoma Chen, 1934 have been revised and now include 14 species. Two new species from Nepal are described, Parambrostoma kippenbergi sp.n. and P. medvedevi sp.n. Three new synonymies are established: Ambrostoma rugosopunctatum Chen = Ambrostoma (Parambrostoma) laosensis Kimoto & Gressitt, syn.n. , Ambrostoma rugosopunctatum Chen = Ambrostoma daccordii Medvedev, syn.n., Ambrostoma fortunei (Baly) = Ambrostoma quadriimpressum chusanica Gruev, syn.n . One species was transferred from Chrysomela Linnaeus to Ambrostoma Motschulsky: A. superbum (Thunberg), comb.n . All the species now included are described and illustrated. Microcomputer tomography was applied for the first time in a study on chrysomelid beetles. A cladistic analysis based on morphological characters of adults was conducted to reconstruct the intergeneric and interspecific phylogeny of Ambrostoma and Parambrostoma. The results show that the monophyly of both genera is well supported. Ambrostoma is widespread in East Asia, whereas Parambrostoma is restricted to the southern slope of the Himalayas, where a relatively recent and modest speciation took place.     

Molecular systematics of Eumolpinae and the relationships with Spilopyrinae (Coleoptera, Chrysomelidae)

The 3400 species of Eumolpinae constitute one of the largest subfamilies of leaf beetles (Chrysomelidae). Their systematics is still largely based on late 19th century monographs and remains highly unsatisfactory. Only recently, some plesiomorphic lineages have been split out as separate subfamilies, including the southern hemisphere Spilopyrinae and the ambiguously placed Synetinae. Here we provide insight into the internal systematics of the Eumolpinae based on molecular phylogenetic analyses of three ribosomal genes, including partial mitochondrial 16S and nuclear 28S and complete nuclear 18S rRNA gene sequences. Sixteen morphological characters considered important in the higher-level systematics of Eumolpinae were also included in a combined analysis with the molecular characters. All phylogenetic analyses were performed using parsimony by optimizing length variation directly on the tree, as implemented in the POY software. The data support the monophyly of the Spilopyrinae outside the clade including all sampled Eumolpinae, corroborating their treatment as a separate subfamily within the Chrysomelidae. The systematic placement of the Synetinae remains ambiguous but consistent with considering it a different subfamily as well, since the phylogenetic analyses using all the available evidence show the representative sequence of the subfamily also unrelated to the Eumolpinae. The Megascelini, traditionally considered a separate subfamily, falls within the Eumolpinae. Several recognized taxonomic groupings within Eumolpinae, including the tribes Adoxini or Typophorini, are not confirmed by molecular data; others like Eumolpini seem well supported. Among the morphological characters analyzed, the presence of a characteristic groove on the pygidium (a synapomorphy of the Eumolpini) and the shape of tarsal claws (simple, appendiculate or bifid) stand out as potentially useful characters for taxonomic classification in the Eumolpinae.     

Taxonomic review of the genus Gonioctena Chevrolat (Coleoptera: Chrysomelidae: Chrysomelinae) in the Korean Peninsula

A taxonomic review of the genus Gonioctena in the Korean Peninsula is presented. Ten species belonging to three subgenera are recognized. Gonioctena jacobsoni (Ogloblin et Medvedev) and Gonioctnea kamiyai Kimoto are recorded for the first time in Korea. A key, illustrations of diagnostic characters, and the coloration patterns of Korean Gonioctena are provided.     

First report of the invasive Australian tortoise beetle Trachymela sloanei (Coleoptera: Chrysomelidae: Chrysomelinae) in Asia

The Australian tortoise beetle Trachymela sloanei (Coleoptera: Chrysomelidae: Chrysomelinae), an invasive pest of several species of Eucalyptus has been reported for the first time in Asia. It was first detected in Shenzhen, China where it has become established. Currently, T. sloanei is not a destructive forest pest in Shenzhen. Pupae and adults were frequently found infected by an unknown fungus.     

中国扁叶甲属地理分布格局研究(叶甲科,叶甲亚科)

本文运用GIS技术研究扁叶甲属Gastrolina在中国的地理分布格局。以采集地的行政单元转换地理坐标与动物地理区划图叠加后产生叶甲的地理分布图。结果显示此属主要分布在华中区、华北区、华南区、西南区、东北区。根据地理分布图及有关分析,认为华中区中部、华北区西北部是该属的丰富度中心和分布中心,华中区中部为中国扁叶甲多样性中心和分化中心。     

New data on the systematics of leaf-beetle genus Chrysolina (Coleoptera, Chrysomelinae) from the Palaearctic region

Chrysolina przewalskii (Jacobson, 1895) and Ch. lucidula Chen, 1934 are redescribed based on the type specimens. Neotype of Ch. przewalskii is designated. Ch. daccordii Lopatin, 2000 is a new junior synonym of Ch. juldusana (Lopatin, 1962); Ch. daosana Lopatin, 2007 is a new junior synonym of Ch. tani Lopatin, 1998; Ch. burchana Lopatin, 1998 is a new junior synonym of Ch. przewalskii (Jacobson, 1895). Ch. lucidula Chen, 1934 described from a mislabeled specimen is a new junior synonym of the South European Ch. lucida (Olivier, 1807). Ch. hyrcana var. cyanescens (Jacobson, 1894) is synonymized with Ch. circumducta (Ménétriés, 1848) and is merely a color form of the latter species.     

核桃扁叶甲三亚种的分类地位订正 (鞘翅目：叶甲科，叶甲亚科)

通过对核桃扁叶甲3个亚种（指名亚种Gastrolina depressa depressa Baly,淡足亚种G. depressa pallipes Chen和黑胸亚种G. depressa thoracica Baly）的比较形态学研究,结合生物学和生物地理学方面的资料,对其分类地位进行了探讨。从形态上来说,三者的上唇、下唇、后翅和爪的形态区别较小,但是触角、上颚、下颚和受精囊则存在着较大的区别,而且通过超微形态的比较,淡足亚种、黑胸亚种和指名亚种均存在着较为明显的形态区别。动物地理学研究表明,指名亚种和黑胸亚种在我国北纬25°～30°之间有地域重叠,目前仅在部分地区可以通过海拔来区分。但是对于淡足亚种而言,它的分布区与其他两个亚种有着一定的地理隔离,其间并没有重叠区的存在。因此从地理分布上而言,三者也存在着不同。通过上述讨论,本文将核桃扁叶甲三亚种恢复或提升为种,即核桃扁叶甲G. depressa Baly,黑胸扁叶甲G. thoracica Baly和淡足扁叶甲Gastrolina pallipes Chen, stat. Nov.。     

Molecular systematics reveals the origins of subsociality in tortoise beetles (Coleoptera,Chrysomelidae, Cassidinae)

Subsocial behaviour is known to occur in at least 19 insect orders and 17 families of Coleoptera. Within the leaf beetle family, Chrysomelidae, extended maternal care is reported in only 2 of 15 subfamilies: Cassidinae and Chrysomelinae. Although the emergence of subsociality in insects has received much attention, extensive analyses on the evolution of this behaviour based on phylogenetic approaches are missing. Subsociality is recorded in 33 species of tortoise beetles belonging to the tribes Mesomphaliini and Eugenysini. A molecular phylogenetic reconstruction of these tribes and the remaining five Neotropical tribes of cassidine tortoise beetles was used to investigate the evolution of maternal care and to elucidate the phylogenetic relationships among Neotropical cassidine tribes. A phylogeny was constructed using 90 species and three loci from both mitochondrial and nuclear genes (COI, CAD and 28S). Bayesian inference and maximum likelihood analyses based on a concatenated dataset recovered two independent origins, with no evidence of reversal to solitary behaviour. One origin comprises three Mesomphaliini genera tightly associated with Convolvulaceae, and the other consists of the genus Eugenysa Chevrolat (Eugenysini), a small clade embedded within a group feeding exclusively on Asteraceae. A previous hypothesis suggesting dual origins on different host plants was confirmed, whereas other hypotheses based on a phylogenetic reconstruction of Cassidinae could not be sustained. Our analysis also revealed that the tribe Mesomphaliini is a monophyletic taxon if Eugenysini is included, and for this reason, we re-establish synonymy of both tribes. We also provide nine new records of subsociality for tortoise beetles species.