Life forms and strategies of lichen-feeding darkling beetles (Coleoptera,Tenebrionidae: Helopini)

Trophic relations of lichen-feeding darkling beetles of the tribe Helopini in Turkey, the Caucasus, and Anti-Atlas are considered. This group is the most diverse in relict Anatolian forests of Cedrus libani, Abies cilicica, Pinus nigra, Juniperus spp., and Quercus cerris, where multitaxon lichenophagous assemblages exist. Three life strategies for attenuation of interspecific competition are distinguished: (1) differentiation of the consumed lichens, (2) separation of the life forms based on the types of the lichens consumed, and (3) segregation of circadian activity in adults. We assume that adaptation of the Helopini larvae to the soil life promoted their wide distribution in different landscapes by elimination of the limiting factor for completing their life cycles, namely the need for ligneous substrate of old rotting trees. The possibility of using lichen-feeding beetles of the tribe Helopini in ecosystem and lichen indication is considered.     

台湾和日本拟步甲在纬度分布上的比较研究

台湾和日本处在不同的地理纬度上,同属岛屿海洋性气候,前者地域面积远小于后者,拟步甲的物种多样性却大于后者。为弄清楚这些科学问题,作者采用G-F指数对从台湾到日本不同纬度梯度上的拟步甲多样性分布格局进行了比较分析,得到如下初步结论：(1) G-F指数从大到小依次是：台湾(21°N-25°N)(0．826)>日本(24°N-45°N)(0．824)>日本Ⅱ纬度区(30°N-35°N)(0．792)>日本Ⅰ纬度区(24°N-30°N)(0．765)>日本Ⅲ纬度区(35°N-40°N)(0．761)>日本Ⅳ纬度区(40°N-45°N)(0．603);(2)台湾拟步甲属的多样性(DG)、族的多样性(DF)和G-F指数(DG-F)均最高,分别是4．263、24．464和0．826;(3)各纬度上拟步甲的物种分布情况：台湾(21°N-25°N)(541种)>日本(24°N-45°N)(489种)>日本Ⅰ纬度区(24°N-30°N)(257种)>日本Ⅱ纬度区(30° N-35° N)(231种)>日本Ⅲ纬度区(35° N-40° N)(172种)>日本Ⅳ纬度区(40° N-45° N)(60种)。研究数据显示, G-F指数能较好地反映台湾和日本各地拟步甲族、属的多样性。其物种多样性在纬度上的分布表现为从南向北递减的趋势,并对其基本原因进行分析。作者首次基于台湾和日本两个岛屿拟步甲物种多样性的比较分析,对现有岛屿生物多样性的有关理论提出个人看法,认为岛屿生物地理学的“物种-面积关系理论”中的“岛屿面积越大,物种数量就越多”可能存在一定的局限性,不一定能客观地反映种类众多的现生岛屿昆虫物种多样性的实际情况。     

The taxocene of lichen-feeding darkling beetles (Coleoptera,Tenebrionidae: Helopini) in a forest-steppe ecotone

The taxocene of lichen-feeding darkling beetles from the tribe Helopini is studied in the steppes of the south of Rostov Province. The species of lichens grazed on by the representatives of this tribe are identified for the first time and dependence of circadian activity and spatial distribution of adults on temperature and air humidity is shown. Global warming and human activities may have caused degradation of steppe lichen communities that formed the dietary base for the lichen-feeding darkling beetles in natural ecosystems. As a result, 3 out of 4 beetle species completely left steppe biotopes for tree shelterbelts. This caused a change in the trophic relationships (transition from epigeic to epiphytic lichens) and in the spatial distribution of darkling beetles, promoted the establishment of competitive interactions, and increased the duration of circadian and seasonal activity due to the lengthening of the periods with optimum temperature and air humidity for each species.     

Spatial structure of community of darkling beetles (Coleoptera: Tenebrionidae) in the Negev Highlands, Israel

The distribution of niches in resource space of darkling beetles of Negev desert, Israel were studied using discriminant analysis Seventeen environmental variables (substrate structure and vegetation) were measured at 375 plots within 15 grids The first three discriminant axes account for 77% of the variance and reflect a soil structure gradient from sand to clay (the first axis), a gradient of increasing productivity (the second axis), and a type of vegetation architecture (the third axis) Abundance and biomass of tenebrionids increased linearly along the first, the second and the third axes Species richness and diversity had a slanting parabola shape along the substrate axis and increased along the productivity axis There were two distinct spatial guilds of tenebrionid species One guild was composed by species that preferred sandy-gravel soils, and another guild consisted of species that preferred clay soils Niche overlapping of the "sandy-gravel" species was lower than that of the "clay" species Niches of the "sandy-gravel" species were randomly distributed in the resource space Those of the "clay" species formed a tight cluster with most niches placed in the center of the space, representing the centrifugal type of organization The conformity of the guild structure with the different types of organization was supported by compansons between empirical data and stochastic models The border between these two guilds matches with proposed earlier biogeographical boundary (Ramon-Zohar line)     

Cytogenetics of the darkling beetles Zophobas aff. confusus and Nyctobates gigas (Coleoptera, Tenebrionidae)

Males of Zophobas aff. confusus and Nyctobates gigas (Tenebrionidae) collected in the State of Pernambuco, Brazil, were studied through conventional staining, C-banding, silver nitrate impregnation (AgNO(3)), and the base specific fluorochromes CMA(3) and DAPI. Z. aff. confusus was found to have 2n = 20 (9+Xyp) while N. gigas exhibited 2n = 18 (8+neoXY). Large pericentromeric blocks of constitutive heterochromatin (CH) were detected throughout the autosomal complement of the two species, except in one autosomal pair of N. gigas in which no heterochromatic block was observed. The sex chromosomes of both species were almost totally heterochromatic. Double staining with CMA(3)/DA (distamycin) and DAPI/DA marked CH in Z. aff. confusus. However, DAPI staining was more intense. N. gigas was found to possess blocks of CH-positive CMA(3) and homogeneous DAPI. AgNO(3) staining also revealed differences between the two species. In Z. confusus an NOR was observed in the sexual bivalent Xyp and N. gigas was found to have an autosomal NOR.     

Multiple-chromosome sex systems in the darkling beetles Blaps gigas and Blaps gibba (Coleoptera, Tenebrionidae)

We have studied mitotic and meiotic chromosomes in the males of two species of Blaps: B. gigas and B. gibba. Karyological characteristics such as the occurrence of a multivalent configuration at diakinesis and two types of metaphase-II spreads support the notion that multiple-chromosome sex systems involving five chromosomes in B. gigas and eight chromosomes in B. gibba have developed in these species. Results obtained by means of silver staining and C-banding techniques suggest that the complex sex systems occurring in B. gigas and B. gibba may have originated from exchanges of terminal ribosomal genes among the Y chromosome and some autosomes.     

A new genus and species of darkling beetles of the tribe Helopini (Coleoptera,Tenebrionidae) from Turkey

A new genus and species Microdocnemis xerophilicus gen. et sp. n. of the tenebrionid-beetle tribe Helopini is described from southwestern Turkey. The new genus belongs to the cylindrinotoid group of genera and is closely related to the genus Odocnemis Allard, 1876 in the structure of the male genitalia and female genital tubes and in the denticulation of the inner margin of the fore tibia. Microdocnemis differs from Odocnemis in the following characters: body very weakly flattened dorsally and ventrally; apical part of elytra, body ventrally, and epipleura with hairs; anal sternite with double bordering; apical margins of tibiae with short thick spines. The only species of the new genus inhabits xerophytic stony biotopes and has been found under stones; the species of the genus Odocnemis live on tree trunks covered with lichens.     

Reevaluation of Blapimorpha and Opatrinae: addressing a major phylogeny‐classification gap in darkling beetles (Coleoptera: Tenebrionidae: Blaptinae)

The taxonomic concepts of Blapimorpha and Opatrinae (informal and traditional, morphology‐based groupings among darkling beetles) are tested using molecular phylogenetics and a reassessment of larval and adult morphology to address a major phylogeny‐classification gap in Tenebrionidae. Instead of a holistic approach (family‐level phylogeny), this study uses a bottom‐up strategy (tribal grouping) in order to define larger, monophyletic lineages within Tenebrioninae. Sampling included representatives of 27 tenebrionid tribes: Alleculini, Amarygmini, Amphidorini, Blaptini, Bolitophagini, Branchini, Cerenopini, Coniontini, Caenocrypticini, Dendarini, Eulabini, Helopini, Lagriini, Melanimini, Opatrini, Pedinini, Phaleriini, Physogasterini, Platynotini, Platyscelidini, Praociini, Scaurini, Scotobiini, Tenebrionini, Trachyscelini, Triboliini and Ulomini. Molecular analyses were based on DNA sequence data from four non‐overlapping gene regions: carbamoyl‐phosphate synthetase domain of rudimentary (CAD) (723 bp), wingless (wg) (438 bp) and nuclear ribosomal 28S (1101 bp) and mitochondrial ribosomal 12S (363 bp). Additionally, 15 larval and imaginal characters were scored and subjected to an ancestral state reconstruction analysis. Results revealed that Amphidorini, Blaptini, Dendarini, Pedinini, Platynotini, Platyscelidini and Opatrini form a clade which can be defined by the following morphological features: adults—antennae lacking compound/stellate sensoria; procoxal cavities externally and internally closed, intersternal membrane of abdominal ventrites 3–5 visible; paired abdominal defensive glands present, elongate, not annulated; larvae—prolegs enlarged (adapted for digging); ninth tergite lacking urogomphi. To accommodate this monophyletic grouping (281 genera and ～4000 species), the subfamily Blaptinae sens. nov. is resurrected. Prior to these results, all of the tribes within Blaptinae were classified within the polyphyletic subfamily Tenebrioninae. The non‐monophyletic nature of Terebrioninae has already been postulated by previous authors, yet no taxonomic decisions were made to fix its status. The reinstatement of Blaptinae, which groups ～50% of the former Tenebrioninae, helps to clarify phylogenetic relations among the whole family and is the first step towards a complete higher‐level revision of Tenebrionidae. The Central Asian tribe Dissonomini (two genera, ～30 species) was not included in Blaptinae due to a lack of representatives in the performed phylogenetic analyses; however, based on morphological features, the tribe is listed as a potential addition to the subfamily.     

A comprehensive molecular phylogeny of tiger beetles (Coleoptera,Carabidae, Cicindelinae)

Tiger beetles are a remarkable group that captivates amateur entomologists, taxonomists and evolutionary biologists alike. This diverse clade of beetles comprises about 2300 currently described species found across the globe. Despite the charisma and scientific interest of this lineage, remarkably few studies have examined its phylogenetic relationships with large taxon sampling. Prior phylogenetic studies have focused on relationships within cicindeline tribes or genera, and none of the studies have included sufficient taxon sampling to conclusively examine broad species patterns across the entire subfamily. Studies that have attempted to reconstruct higher‐level relationships of Cicindelinae have yielded conflicting results. Here, we present the first taxonomically comprehensive molecular phylogeny of Cicindelinae to date, with the goal of creating a framework for future studies focusing on this important insect lineage. We utilized all available published molecular data, generating a final concatenated dataset including 328 cicindeline species, with molecular data sampled from six protein‐coding gene fragments and three ribosomal gene fragments. Our maximum‐likelihood phylogenetic inferences recover Cicindelinae as sister to the wrinkled bark beetles of the subfamily Rhysodinae. This new phylogenetic hypothesis for Cicindelinae contradicts our current understanding of tiger beetle phylogenetic relationships, with several tribes, subtribes and genera being inferred as paraphyletic. Most notably, the tribe Manticorini is recovered nested within Platychilini including the genera Amblycheila Say, Omus Eschscholtz, Picnochile Motschulsky and Platychile Macleay. The tribe Megacephalini is recovered as paraphyletic due to the placement of the monophyletic subtribe Oxycheilina as sister to Cicindelini, whereas the monophyletic Megacephalina is inferred as sister to Oxycheilina, Cicindelini and Collyridini. The tribe Collyridini is paraphyletic with the subtribes Collyridina and Tricondylina in one clade, and Ctenostomina in a second one. The tribe Cicindelini is recovered as monophyletic although several genera are inferred as para‐ or polyphyletic. Our results provide a novel phylogenetic framework to revise the classification of tiger beetles and to encourage the generation of focused molecular datasets that will permit investigation of the evolutionary history of this lineage through space and time.     

Sperm morphology and phylogeny of lagriids (Coleoptera,Tenebrionidae)

The systematics of tenebrionids remain unclear, principally at the subfamily level, as is the case of the Lagriinae. Considering that sperm morphology has contributed to the various insect group phylogenies, in this work we describe the structure and ultrastructure of these cells in Lagria villosa. Sperm in this species exhibit a strong morphological similarity to those of Tenebrio molitor and Tribolium castaneum, the only two species of Tenebrionidae with previously described sperm. In tenebrionids, the flagellar components offer good diagnostic characters, e.g. the symmetry of mitochondrial derivatives in L. villosa differentiates sperm of this species from those of Te. molitor and Tr. castaneum. However, the lateral association of the nucleus with flagellar components, the form of accessory bodies, and the presence of material connecting the axoneme to the accessory bodies and mitochondrial derivatives indicate that the three species form a sister group. Therefore, the sperm morphology of L. villosa support lagriid beetles as a subfamily (Lagriinae) of Tenebrionidae.     

Higher‐level phylogeny of diving beetles (Coleoptera: Dytiscidae) based on larval characters

A comprehensive higher‐level phylogeny of diving beetles (Dytiscidae) based on larval characters is presented. Larval morphology and chaetotaxy of a broad range of genera and species was studied, covering all currently recognized subfamilies and tribes except for the small and geographically restricted Hydrodytinae, where the larva is unknown. The results suggest several significant conclusions with respect to the systematics of Dytiscidae including the following: monophyly of all currently recognized subfamilies, although Dytiscinae when considered in a broad context is rendered paraphyletic by Cybistrinae; currently recognized tribes are monophyletic except for Agabini, Hydroporini and Laccornellini; inter‐subfamily and inter‐tribe relationships generally show weak support, except for a few well supported clades; three distinct clades are recognized within Dytiscinae [Dytiscini sensu lato (i.e. including the genera Dytiscus Linnaeus and Hyderodes Hope), Hydaticini sensu lato, and Cybistrini]; and recognition of Pachydrini as a distinct tribe. Other less robust results include: Methlini sister to the rest of Hydroporinae; relative basal position of Laccornini, Hydrovatini and Laccornellini within Hydroporinae; close relationship of Agabinae and Copelatinae; Matinae nested deep within Dytiscidae, as sister to a large clade including Colymbetinae, Coptotominae, Lancetinae and Dytiscinae sensu lato; the sister‐group relationship of Agabetini and Laccophilini is confirmed. The results presented here are discussed and compared with previous phylogenetic hypotheses based on different datasets, and the evolution of some significant morphological features is discussed in light of the proposed phylogeny. All suprageneric taxa are diagnosed, including illustrations of all relevant synapomorphies, and a key to separate subfamilies and tribes is presented, both in traditional (paper) format and as an online Lucid interactive identification key.     

Molecular insights into the phylogeny of Blapstinina (Coleoptera: Tenebrionidae: Opatrini)

The monophyly of the North and South American endemic subtribe Blapstinina (Tenebrionidae: Opatrini) is tested through phylogenetic analyses using five molecular markers [nuclear ribosomal 28S (28S), cytochrome oxidase subunit II (COII), arginine kinase (ArgK), carbamoyl-phosphate synthetase domain of rudimentary (CAD), wingless (wg)]. Representatives of several opatrinoid subtribes were taken into consideration, including other geographically overlapping endemic genera, namely Ammodonus, Ephalus and Pseudephalus (all previously considered representatives of Ammobiina). A comparative study of morphology was performed to assess resulting phylogenetic hypotheses. Analyses support the monophyly of Blapstinina; however, they also strongly indicate that Ammodonus should be included within the subtribe. Mecysmus is nested within Blapstinus and therefore, a new synonymy, Blapstinus (= Mecysmus syn.n. ), and the following combinations are introduced: Blapstinus advena comb.n. , B. angustus comb.r. , B. laticollis comb.n. , B. parvulus comb.n. , B. tenuis comb.n. Morphological analysis showed a close affiliation between Ephalus and Pseudephalus. Based on these results, Pseudephalus is synonymized with Ephalus [Ephalus (= Pseudephalus syn.n. )], and the following combination is introduced: Ephalus brevicornis comb.n. Recovered topologies also strongly support transferring Ephalus stat.n. into Opatrina, making the distribution of Opatrina amphi-Atlantic.     

Higher‐level phylogeny of longhorn beetles (Coleoptera: Chrysomeloidea) inferred from mitochondrial genomes

Cerambycidae (longhorn beetles) and related families in the superfamily Chrysomeloidea are important components of forest ecosystems and play a key role in nutrient cycling and pollination. Using full mitochondrial genomes and dense taxon sampling, the phylogeny of Chrysomeloidea with a focus on Cerambycidae and allied families was explored. We used 151 mitochondrial genomes (75 newly sequenced) covering all families and 29 subfamilies of Chrysomeloidea. Our results reveal that (i) Chrysomelidae (leaf beetles) are sister to all other chrysomeloid families; (ii) Cerambycidae sensu stricto (s. s.) is polyphyletic due to the inclusion of other families that split Cerambycidae into a ‘lamiine’ clade comprising Lepturinae sensu lato (s. l.) + (Lamiinae + Spondylidinae) and a ‘cerambycine’ clade comprising Dorcasominae + (Cerambycinae + Prioninae s. l.); (iii) the subfamilies within the two clades of Cerambycidae s. s. were monophyletic, except for the placement of Necydalinae nested in Lepturinae, and the placement of Parandrinae within Prioninae (now considered as tribes Necydalini and Parandrini, respectively); (iv) smaller families were grouped into two major clades: one composed of Disteniidae+Vesperidae and the other composed of Orsodacnidae + (Megalopodidae + Oxypeltidae); (v) relationships among the four major clades were poorly supported but were resolved as ((cerambycines + (Disteniidae + Vesperidae) + Orsodacnidae + (Megalopodidae + Oxypeltidae)) + lamiines. Divergence time analyses estimated that Chrysomeloidea originated ca. 154.1 Mya during the late Jurassic, and most subfamilies of Cerambycidae originated much earlier than subfamilies of Chrysomelidae. The diversification of families within Chrysomeloidea was largely coincident with the radiation of angiosperms during the Early Cretaceous.     

Pythiopina,an enigmatic subtribe of darkling beetles (Coleoptera: Tenebrionidae: Pedinini): taxonomic revision,microtomography, ecological niche models and phylogenetic position

Morphological, anatomical, and distributional data concerning the S outh A frican endemic beetle subtribe P ythiopina (T enebrionidae: P edinini) are revised. Five species, representing two genera, are recognized. Included in this total is one new species (Meglyphus mariae K amiński sp.n. ). The following species are placed in synonymy: Meglyphus ciliatipes [=Meglyphus calitzensis syn.n. ]; Meglyphus laenoides [=Meglyphus andreaei syn.n. ; =Meglyphus namaqua syn.n. ]. Microtomographic models for all valid P ythiopina species, including the holotype of the newly described species, are presented and analysed. Endoskeleton morphology (specifically characters of the tentorium and metendosternite) proved to be informative at the specific and generic levels. An identification key is provided to all known species of the subtribe. Environmental niche models are presented for the majority of species. A molecular phylogeny of P edinini based on six genetic loci (28S : D 1–D 3 region; 28S : D 4–D 5 region, COII , A rgK , CAD 2, wg) was also produced to explore the phylogenetic position of P ythiopina. This analysis is the first to include representatives of all seven subtribes of P edinini, and supports a sister relationship between P ythiopina and the P alaearctic subtribe D endarina. Results also suggest the existence of a second pair of sister taxa within P edinini (in addition to M elambiina) with an amphitropical A frican distribution. This published work has been registered in ZooBank, http://zoobank.org/urn:lsid:zoobank.org:pub:285AD87A‐46B1‐4FE9‐BC57‐949EA1F70D49 .     

鞘翅目高级阶元分子系统学： 研究现状及存在的问题

鞘翅目是世界上物种最丰富的类群, 分为原鞘亚目（Archostemata Kolbe, 1908）、 藻食亚目（Myxophaga Crowson, 1955）、 肉食亚目（Adephaga Schellenberg, 1806）和多食亚目（Polyphaga Emery, 1886）。随着分子生物学的发展,分子系统学的技术被广泛应用于鞘翅目系统学研究中。本文综述了鞘翅目高级阶元的分子系统学的研究进展及存在问题。基于分子生物学手段, 分子分类学家提出了关于鞘翅目高级阶元分子系统学很多假说, 分子分析结果支持鞘翅目的4个亚目各为单系, 而亚目间的系统关系还不统一。基于分子手段对于亚目内的系统发育关系的研究也有了一定的进展, 比如： 分子系统学结果支持肉食亚目的水生类群和陆生类群分别为单系, 水生类群为一次起源。目前, 鞘翅目高级阶元分子系统学的研究还不够成熟和完善, 主要表现为： 材料选择有限且不均衡、 基因数目和适合度不理想, 以及一些关键节点研究的欠缺。     

Climatic correlates of body size in European tenebrionid beetles (Coleoptera: Tenebrionidae)

Tenebrionidae are one of the largest families of beetles and are known for their adaptations to hot and dry climates. An increase in body size also increases the volume/surface area ratio, which reduces transpiration, and hence water loss. If an increase in body size is an important adaptation in tenebrionids to cope with increasing aridity, we expect a correlation between body size and climatic gradients in the major tenebrionid clades. Alternatively, we can postulate that arid climates do not drive body size evolution, but rather select, from a wider fauna containing species of any size, those that have larger bodies. In this case we expect that drier regions will host faunas that contain, on average, larger species. To test the first hypothesis, we correlated inter-specific body size variation in the main tenebrionid clades with climatic gradients in Europe. We found only weak trends. To test the second hypothesis, we regressed mean body size of European country faunas against climatic characteristics. We found a strong increase in body size in southern faunas experiencing hot and dry climates. Therefore, increase in body size is not a major adaptation in tenebrionid evolution, but climate is an important filtering factor that determines a prevalence of larger species in southern Europe.     

Resolving phylogeny at the family level by mitochondrial cytochrome oxidase sequences: phylogeny of carrion beetles (Coleoptera, Silphidae)

We investigated the phylogenetic relationships of carrion beetles (Coleoptera, Silphidae) using 2094 bp of their mitochondrial cytochrome oxidase subunit I and II and tRNA leucine gene sequences. Shorter fragments of this gene region previously have been used to establish generic relationships in insects. In this study, they provided more than sufficient resolution, although the third positions of the protein-coding sequences reached saturation for the deeper divergences. This first published phylogeny for the Silphidae comprises 23 species from 13 genera sampled across the geographic range of the family. In addition, we included species from three related families as outgroups. One of these families, the Agyrtidae, was, until recently, included in the Silphidae, but its resolution here justifies its current position as a separate family. The silphid subfamilies Nicrophorinae and Silphinae are monophyletic in all analyses. All genera for which several species were sampled are supported as monophyletic groups, with the exception of the genus Silpha. European and North American representatives of two Nicrophorus species described from both continents are supported as each others' closest relatives. The lineage that colonized Gondwanaland and that most likely originated in the Palearctic is the most basal within the Silphinae.     

The oldest opatrine terrestrial darkling beetle (Coleoptera: Tenebrionidae: Tenebrioninae) from the Paleocene of Menat (France)

The oldest terrestrial darkling beetle of the opatrine lineage, Palaeosclerum pohli gen. et. sp. nov. (subfamily Tenebrioninae) is described from the Middle Paleocene of Menat (France). The complex of characters, combining the peculiar shape of body (robust, adapted to a fossorial lifestyle), structure of fossorial protibia, 5-segmented antennal club demonstrates a considerable similarity of the new genus to the extant representatives of the ‘sclerina’ generic group of the subtribe Opatrina. Palaeosclerum gen. nov. is compared with recent genera Sclerum Dejean, 1834, Platynosum Mulsant and Rey, 1859 and Eurycaulus Fairmaire, 1868.     

Home range size in two darkling beetle species (Coleoptera,Tenebrionidae) from Southern Kazakhstan

The repeated collecting of marked darkling beetle species, including the diurnal species Tentyria gigas and the nocturnal species Cyphogenia gibba from Kazakhstan demonstrated that their home ranges constituted 457 and 360 m², respectively. More than 500 beetles from an area of 2.5 ha were marked by engraving numbers on their elytra. Beetles were observed during two seasons at a distance covering 2 km from the site of release. Sex was determined in a quarter of all the beetles. In females and males of T. gigas, the home range constituted 700 and 552 m², respectively; in C. gibba, 599 and 343 m², respectively. Small home ranges of these species can be explained by the presence of numerous predators, occasional death of beetles under cattle’s hoofs, and a great number and high density of shelters.     

Molecular phylogeny of rootworms and related galerucine beetles (Coleoptera: Chrysomelidae)

The Galerucinae (Coleoptera: Chrysomelidae) sensu stricto (true galerucines) comprise a large assemblage of diverse phytophagous beetles containing over 5000 described species. Together with their sister taxon, the flea beetles, which differ from true galerucines by having the hind femora usually modified for jumping, the Galerucinae sensu lato comprises over 13 000 described species and is the largest natural group within the Chrysomelidae. Unlike the flea beetles, for which robust hierarchical classification schemes have not been erected, an existing taxonomic structure exists for the true galerucines, based mostly on the works of the late John Wilcox. In the most recent taxonomic list of the Galerucinae sensu stricto, five tribes were established comprising 29 sections housing 488 genera. The majority of the diversity within these tribes is found within the tribe Luperini, in which two genera, Monolepta and Diabrotica, are known to contain over 500 described species. Here, we extend the work from previous phylogenetic studies of the Galerucinae by analysing four amplicons from three gene regions (18S and 28S rRNA; COI) representing 249 taxa, providing the largest phylogenetic analysis of this taxon to date. Using two seven‐state RNA models, we combine five maximum likelihood models (RNA + DNA for the rRNAs; three separate DNA models for the COI codon positions) for these partitions and analyse the data under likelihood using Bayesian inference. The results of these two analyses are compared with those from equally weighted parsimony. Instead of choosing the results from one optimality criterion over another, either based on statistical support, tree topology or philosophical predisposition, we elect to draw attention to the similar results produced by all three analyses, illustrating the robustness of the data to these different analytical methods. In general, the results from all three analyses are consistent with each other and previous molecular phylogenetic reconstructions for Galerucinae, except that increased taxon sampling for several groups, namely the tribes Hylaspini and Oidini, has improved the phylogenetic position of these taxa. As with previous analyses, under‐sampled taxa, such as the Old World Metacyclini and all sections of the subtribe Luperina, continue to be unstable, with the few taxa representing these groups fluctuating in their positions based on the implemented optimality criterion. Nonetheless, we report here the most comprehensive phylogenetic estimation for the Galerucinae to date.