Molecular phylogeny of Neotropical bioluminescent beetles (Coleoptera: Elateroidea) in southern and central Brazil

Bioluminescence in beetles is found mainly in the Elateroidea superfamily (Elateridae, Lampyridae and Phengodidae). The Neotropical region accounts for the richest diversity of bioluminescent species in the world with about 500 described species, most occurring in the Amazon, Atlantic rainforest and Cerrado (savanna) ecosystems in Brazil. The origin and evolution of bioluminescence, as well as the taxonomic status of several Neotropical taxa in these families remains unclear. In order to contribute to a better understanding of the phylogeny and evolution of bioluminescent Elateroidea we sequenced and analyzed sequences of mitochondrial NADH2 and the nuclear 28S genes and of the cloned luciferase sequences of Brazilian species belonging to the following genera: (Lampyridae) Macrolampis, Photuris, Amydetes, Bicellonycha, Aspisoma, Lucidota, Cratomorphus; (Elateridae) Conoderus, Pyrophorus, Hapsodrilus, Pyrearinus, Fulgeochlizus; and (Phengodidae) Pseudophengodes, Phrixothrix, Euryopa and Brasilocerus. Our study supports a closer phylogenetic relationship between Elateridae and Phengodidae as other molecular studies, in contrast with previous morphologic and molecular studies that clustered Lampyridae/Phengodidae. Molecular data also supported division of the Phengodinae subfamily into the tribes Phengodini and Mastinocerini. The position of the genus Amydetes supports the status of the Amydetinae as a subfamily. The genus Euryopa is included in the Mastinocerini tribe within the Phengodinae/Phengodidae. Copyright © 2013 John Wiley & Sons, Ltd.     

Molecular phylogeny and phylogeography of flightless beetles Parechthistatus gibber and Hayashiechthistatus inexpectus (Coleoptera: Cerambycidae) inferred from mitochondrial COI gene sequences

To elucidate the speciation patterns of two endemic flightless cerambycid beetles, Parechthistatus gibber and Hayashiechthistatus inexpectus, molecular phylogenetic analysis was carried out. A 1144 bp region of the cytochrome oxidase subunit I gene was sequenced for individuals from 51 local populations of these species. There were nine haplotype lineages of P. gibber, and H. inexpectus was included within a P. gibber lineage. These lineages were highly divergent and occurred in different regions. Based on previously published molecular change rates for the COI gene (1.5–2.3% per million years), the time of divergence of P. gibber COI haplotypes was inferred to be 3.0–4.6 million years ago, in the Pliocene.     

Contributions to molecular systematics of water scavenger beetles (Hydrophilidae,Coleoptera)

Phylogenetic relationships within Hydrophilidae were examined by analyses of separate and combined nuclear and mitochondrial markers (28S rRNA, 18S rRNA, 16S rRNA, 12S rRNA, COI and COII genes). The preferred (Bayesian) tree topology suggests a sister group relationship between Spercheidae and Hydrophilidae, supporting the ‘hydrophilid lineage’; Epimetopidae are placed on the base of the ‘helophorid branch’, the monophyly of Sphaeridiinae is highly supported, nested deeply within Hydrophilidae closest to Enochrus, making Hydrophilinae and Acidocerini paraphyletic; Hydrobius appears as sister taxon to (Hydrochara + Hydrophilus) without a closer relationship to Acidocerini; the hydrophiloid–histeroid sister group relationship is confirmed. The topology of several taxa remains contradictory, and requires further investigations with a larger taxon sampling and additional molecular markers.     

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.     

Ground beetles (Coleoptera: Carabidae) as bioindicators

One of the primary goals of research on bioindicators is to identifyspecies or other taxonomic units that would reliably indicate disturbances inthe environment, and reflect the responses of other species or the overallbiodiversity. However, there is no perfect bioindicator and selecting the mostsuitable one depends to a great extent on the goal of the survey. In this paperwe examine the suitability of carabids as bioindicators. Carabids are frequentlyused to indicate habitat alteration. They have been used in grasslands andboreal forests where species number and/or abundances have been noted to changealong a habitat disturbance gradient. A common trend is that large, poorlydispersing specialist species decrease with increased disturbance while smallgeneralist species with good dispersal ability increase. Some species are notaffected by moderate disturbance. There is, however, not enough research todetermine how suitable carabids are for biodiversity studies, or how well theyrepresent the response of other species. We conclude that carabids are usefulbioindicators, but as crucial understanding of their relationship with otherspecies is incomplete, they should be used with caution.     

大步甲的分子系统与进化模式

大步甲主要分布于北半球,全世界大约有1000余种。因大步甲后翅退化,不能飞行,其移动能力和扩散范围都受到很大程度的限制,因此,容易产生地理隔离和遗传分化,从而成为研究物种分化和生物多样化的好材料。近年来,通过对大步甲分子系统的详细研究,不仅很大程度上解明了大步甲各分类群间的系统发育关系,而且对大步甲的系统演化过程和形态进化模式也有了较深的理解,获得了很多重要的见解。在此做一个简单的综述。     

Global diversity of water beetles (Coleoptera) in freshwater

The global diversity of True Water Beetles, False Water Beetles and Phytophilous Water Beetles (sensu Jäch, 1998a. In Jäch &; Ji (eds), Water Beetles of China, Vol. II. Zoologisch-Botanische Gesellschaft in Österreich and Wiener Coleopterologenverein, Wien: 25–42.) is assessed. Facultative Water Beetles, Parasitic Water Beetles and Shore Beetles (sensu Jäch, 1998a. In Jäch &; Ji (eds), Water Beetles of China, Vol. II. Zoologisch-Botanische Gesellschaft in Österreich and Wiener Coleopterologenverein, Wien: 25–42.) are here classified as “paraquatic” and are thus not included in the assessment. It is estimated that about 18,000 species of aquatic Coleoptera are roaming the earth at present. About 12,600 (70%) of these are already described (deadline: October 2005). About 30 beetle families have aquatic representatives, and in 25 of these families at least 50% of the species are to be considered as aquatic. Six families are supposed to include 1,000 or more aquatic species: Dytiscidae (3,908 described species/5,000 estimated), Hydraenidae (1,380/2,500), Hydrophilidae (1,800/2,320), Elmidae (1,330/1,850), Scirtidae (900/1,700) and Gyrinidae (750/1,000). Scirtidae and Hydraenidae are regarded as the least explored families, the number of described species in each of these two families probably will be almost doubled in the future. The Palearctic (ca. 3,350 described species/ca. 3,900 estimated), the Neotropical (2,510/3,900) and the Afrotropical Region (2,700/3,750) harbour almost the same number of water beetle species, followed by the Oriental (2,200/3,580) and the Australian/Pacific Realm (1,340/2,100). The Nearctic (1,420/1,550) is by far the poorest region in terms of water beetle diversity.     

Molecular phylogeny of Macrolycus (Coleoptera: Lycidae) with description of new species from China

Macrolycus is a genus of net‐winged beetles with 69 species distributed in the eastern Palearctic and northernmost part of the Oriental region. The first molecular phylogeny of Macrolycus was produced using an rrnL + tRNA‐Leu + nad1 mtDNA fragment. The major lineages and species limits were identified with morphology and molecular data. We propose that Cerceros is a subgenus of Macrolycus to enable identification of all adult specimens in the genus without DNA sequencing. Two species groups are proposed in Macrolycus s. str. and six in Cerceros. Additionally, twelve Macrolycus species are newly described from China: M. aquilinus, M. baihualingensis, M. bicolor, M. guangxiensis, M. jianfenglingensis, M. kuatunensis, M. lizipingensis, M. parvus, M. phoeniceus, M. rhodoneurus, M. rosaceus and M. sichuanensis. Macrolycus holzschuhi is proposed to be a junior subjective synonym of M. jeanvoinei. The highest diversity of Macrolycus is found in southern China. The species from the main islands of Japan are placed in two species groups: M. excellens is a sister to remaining species of the M. murzini group and the M. flabellatus group is a monophylum of closely related species in a sister position to the M. bicolor group.     

中国北方常见芫菁分子物种界定(鞘翅目: 芫菁科)

【目的】应用线粒体COI和核CAD基因片段探讨自动条形码间隔探索(automatic barcode gapdiscovery, ABGD)、广义混合Yule溯祖模型(generalized mixed Yule coalescent, GMYC)、贝叶斯泊松树进程(Bayesian Poisson tree processes, bPTP)和贝叶斯系统发育和系统地理分析(Bayesianphylogenetics and phylogeography, BPP) 4种分析方法在芫菁科(Meloidae)昆虫分子物种界定中的适用性。【方法】分别基于COI, CAD和COI+CAD串联序列数据集,应用ABGD, GMYC, bPTP和BPP 4种方法对中国北方芫菁科常见的6属(沟芫菁属Hycleus、斑芫菁属Mylabris、豆芫菁属Epicauta、绿芫菁属Lytta、星芫菁属Megatrachelus和短翅芫菁属Meloe)18个形态种进行分子物种界定,并与形态学鉴定结果进行比较。【结果】利用COI+CAD串联序列数据集所得物种界定结果与形态鉴定结果一致;COI数据集使用ABGD和GMYC方法的界定结果与形态鉴定结果一致,而bPTP划分的物种数较形态鉴定结果多;基于CAD序列在3种单基因物种界定方法的结果中,除GMYC与形态划分一致外,其余均显示部分结果与形态划分不同。【结论】在芫菁科分子物种界定中,多基因联合序列、多种界定方法分析所得结果优于单一基因片段和界定方法的分析结果。本研究的结果为芫菁科昆虫的分子物种界定和整合分类提供了数据支持和参考。     

Phylogeny of minute carabid beetles and their relatives based upon DNA sequence data (Coleoptera, Carabidae, Trechitae)

The phylogeny of ground beetles of supertribe Trechitae is inferred using DNA sequences of genes that code for 28S ribosomal RNA, 18S ribosomal RNA, and wingless. Within the outgroups, austral psydrines are inferred to be monophyletic, and separate from the three genera of true Psydrina (Psydrus, Nomius, Laccocenus); the austral psydrines are formally removed from Psydrini and are treated herein as their own tribe, Moriomorphini Sloane. All three genes place Gehringia with Psydrina. Trechitae is inferred to be monophyletic, and sister to Patrobini.Within trechites, evidence is presented that Tasmanitachoides is not a tachyine, but is instead a member of Trechini. Perileptus is a member of subtribe Trechodina. Against Erwin's hypothesis of anillines as a polyphyletic lineage derived from the tachyine genus Paratachys, the anillines sampled are monophyletic, and not related to Paratachys. Zolini, Pogonini, Tachyina, and Xystosomina are all monophyletic, with the latter two being sister groups. The relationships of the subtribe Bembidiina were studied in greater detail. Phrypeus is only distantly related to Bembidion, and there is no evidence from sequence data that it belongs within Bembidiina. Three groups that have been recently considered to be outside of the large genus Bembidion are shown to be derived members of Bembidion, related to subgroups: Cillenus is related to the Ocydromus complex of Bembidion, Zecillenus is related to the New Zealand subgenus Zeplataphus, and Hydrium is close to subgenus Metallina. The relationships among major lineages of Trechitae are not, however, resolved with these data.     

Proventricular trituration in adult carabid beetles (Coleoptera: Carabidae)

Adult carabids that ingest particles of food have a muscular proventriculus lined internally with an intima that bears sclerotized projections. In experiments where six species of carabid were allowed to feed upon Drosophila, the average size of cuticular remains found in post-proventricular regions of the gut was generally significantly smaller than those in the crop. This reduction in size of cuticular particles is likely to be due to proventricular trituration rather than prolonged exposure to digestive secretions. There is no evidence that material is triturated by the proventriculus and returned to the crop, the main seat of digestion in carabids, so proventricular trituration is unlikely to enhance digestion. By crushing large, potentially abrasive, cuticular particles, the proventriculus may serve to protect midgut epithelial tissue.     

The phylogenetics and biochemistry of host-plant specialization in Melitaeine butterflies (Lepidoptera: Nymphalidae)

Butterflies in the tribe Melitaeini (Lepidoptera: Nymphalidae) are known to utilize host plants belonging to 16 families, although most host-plant records are from four families. Of the 16 host-plant families, 12 produce secondary plant metabolites called iridoids. Earlier studies have shown that larvae of several melitaeine species use iridoids as feeding stimulants and sequester these compounds for larval defense. I investigate the evolutionary history of host-plant use in the tribe Melitaeini by testing a recent phylogenetic hypothesis of 65 species representing the four major species groups of the tribe. By simple character optimization of host-plant families and presence/absence of iridoids in the host plants, I find that plant chemistry is a more conservative trait than plant taxonomy. The ancestral host plant(s) of the entire tribe most likely contained iridoids and were likely to be in the plant family Plantaginaceae. A major host shift from plants containing iridoids to plants not containing iridoids has happened three times independently. The results show that the evolution of host-plant use in melitaeines has been (and still is) a dynamic process when considering plant taxonomy, but is relatively stable when considering host-plant chemistry.     

Lamellicorn beetles (Coleoptera: Scarabaeoidea) in Korea and Mongolia

This work was undertaken as an initial part of the collaborative effort between Korean and Mongolian researchers to study the terrestrial and aquatic insects in these countries. An analysis of data on faunal composition, taxonomic diversity and biogeographical structure of lamellicorn beetles in Korea and Mongolia was conducted in order to characterize the faunal relations between two countries. The Korean fauna of lamellicorn beetles is almost two fold greater in both generic diversity and species richness than that of Mongolia. The species compositions of lamellicorn beetles in two countries are substantially different from each other and the relative levels of faunal overlap are insignificant. Not only the faunal diversity and composition, but also the biogeographical structure of lamellicorn beetles in Korea and Mongolia display dissimilar patterns, except for their similar degree of endemism. Additional data on communities of dung beetles in the grasslands of central Mongolia, as well as a brief discussion on conservation issues of the endangered species in Korea, especially Gymnopleurus mopsus, are given.     

Molecular phylogeny of the tribe Erotini with description of a new genus from China (Coleoptera: Lycidae)

Erotini is a small tribe of net‐winged beetles known from the northern temperate zone. We investigated relationships and limits of tribes and genera using a molecular phylogeny inferred from rrnL, cox1 and nad5 mtDNA fragments. Lopheros Leconte, 1881 and Pseudaplatopterus Kleine, 1940 were inferred in current molecular analyses as terminal lineages within Erotini. Therefore, we consider Lopherotini Kazantsev, 2012 and Pseudaplatopterina Kazantsev, 2012 as junior synonyms of Erotini Leconte, 1881. The Platycis genus group comprises 19 species representing several genus‐group taxa previously placed in Platycis s. l. Konoplatycis Nakane, 1969 was recovered as a deeply rooted lineage of Erotini and we inferred another independent lineage for which we propose a new genus Sinoplatycis gen. nov. Additionally, we studied the diversity of Platycis s. l. in Asia, found their highest diversity in the temperate forest habitats of Eastern Asia and inferred the paraphyletic character of the subgenus Erotides s. str. Five new species are described from China: Sinoplatycis cardinalis sp. nov. , S. zhani sp. nov. (type species of Sinoplatycis gen. nov. ), Erotides brunnescens sp. nov. , E. pusillus sp. nov. and E. slipinskii sp. nov. The molecular phylogeny shows deep splits of Japanese species with respect to the continental Eurasian fauna and the role of shallow seas and mountain systems in the diversification of the Platycis genus group.     

基于线粒体基因组序列的鞘翅目肉食亚目水生类群系统发育分析

【目的】明确肉食亚目(Adephaga)水生类群线粒体基因组的基本特征,并基于线粒体基因组序列分析肉食亚目水生类群的系统发育关系。【方法】基于Illumina HiSeq X Ten测序技术测定了圆鞘隐盾豉甲Dineutus mellyi和齿缘龙虱Eretes sticticus的线粒体全基因组序列,对其进行了基因注释,并对其tRNA基因二级结构进行了预测分析。加上已公布的鞘翅目(Coleoptera)肉食亚目水生类群17个种的线粒体基因组序列,对该类群共19个种线粒体的蛋白质编码基因(protein-coding genes, PCGs)开展了比较基因组学分析,包括AT含量、密码子偏好性、选择压力等。基于13个PCGs的氨基酸序列和核苷酸序列,利用最大似然法(ML)和贝叶斯法(BI)分别构建鞘翅目肉食亚目水生类群的系统发育关系,并通过FcLM分析进一步评估伪龙虱科(Noteridae)和瀑甲科(Meruidae)的系统发育位置。【结果】圆鞘隐盾豉甲和齿缘龙虱的线粒体基因组全长分别为16 123 bp(GenBank登录号: MN781126)和16 196 bp(GenBank登录号: MN781132),都包含13个PCGs、22个tRNA基因、2个rRNA基因和1个D-loop区(控制区)。19个肉食亚目水生类群线粒体基因组PCGs的碱基组成都呈现A+T偏好性,在密码子使用上也都偏向于使用富含A+T的密码子;在进化过程中13个PCGs的进化模式相同,都受到纯化选择。基于线粒体基因组13个PCGs的氨基酸序列的肉食亚目水生类群的系统发育关系为(豉甲科Gyrinidae+(沼梭甲科Haliplidae+((壁甲科Aspidytidae+(两栖甲科Amphizoidae+龙虱科Dytiscidae))+(水甲科Hygrobiidae+(瀑甲科Meruidae+伪龙虱科Noteridae)))))。【结论】研究结果表明,豉甲科是肉食亚目水生类群的基部类群,接下来是沼梭甲科和龙虱总科;伪龙虱科和瀑甲科互为姐妹群,并一起作为龙虱总科内部的一个分支;两栖甲科与龙虱科具有更近的亲缘关系。     

The relationship between the ventral nerve cord,body size and phylogeny in ground beetles (Coleoptera: Carabidae)

The ventral nerve cord in the family Carabidae (considered in the widest sense! exhibits variations in the degree of fusion of thoracic and abdominal ganglia. There are usually three discrete thoracic ganglia and between one and seven discrete abdominal ganglia, the number differing between tribes. Of the 44 tribes and 177 species examined, 38 tribes contained species showing no differences in the degree of ventral nerve cord consolidation. However, the remaining six tribes showed variations in the degree of ventral nerve cord consolidation between genera (Lebiini, Cychrini, Nebriini, Scaritini, Licinini and Brachinini), whilst one genus showed variations between species (Leistus, Nebriini). No variation in ventral nerve cord consolidation was observed in conspecifics. The degree of ventral nerve cord consolidation is inversely proportional to overall body length. With respect to phylogeny, the degree of consolidation of the nerve cord docs not consistently support the traditional Carabinae-Harpalinae subfamily division. However, the Harpalinae always have four or less discrete abdominal ganglia (with the sole exception of the Broscinij, whilst the Carabinae exhibit almost the whole range of variations. Thus the Harpalinae (or the major pari of it) may be a monophyletic group, but this is not true of the Carabinae. Trends in the degree of ventral nerve cord consolidation for the various tribes were noted, and phylogenetic implications were evaluated wherever possible.     

An unexpected clade of South American ground beetles (Coleoptera,Carabidae, Bembidion)

Phylogenetic relationships of the Antiperyphanes Complex of the genus Bembidion are inferred using DNA sequences from seven genes (two nuclear ribosomal, four nuclear protein coding, and one mitochondrial protein coding). Redefined subgenera within the complex are each well-supported as monophyletic. Most striking was the discovery that a small set of morphologically and ecologically heterogeneous species formed a clade, here called subgenus Nothonepha. This unexpected result was corroborated by the discovery of deep pits in the lateral body wall (in the mesepisternum) of all Nothonepha, a trait unique within Bembidion. These pits are filled with a waxy substance in ethanol-preserved specimens. In one newly discovered species (Bembidion tetrapholeon sp. n., described here), these pits are so deep that their projections into the body cavity from the two sides touch each other internally. These structures in Bembidion (Nothonepha) are compared to very similar mesepisternal pits which have convergently evolved in two other groups of carabid beetles. The function of these thoracic pits is unknown. Most members of subgenus Nothonepha have in addition similar but smaller pits in the abdomen. A revised classification is proposed for the Antiperyphanes Complex.     

Cost of shield defence for tortoise beetles (Coleoptera: Chrysomelidae)

Abstract.

• 1 Tortoise beetle larvae possess a shield composed of exuviae and faeces which functions as an effective defence against some invertebrate predators.
• 2 In the laboratory, Charidotella bicolor, Deloyala guttata and Chelymorpha cassidea larvae with their shields experimentally removed did not exhibit enhanced performance (i.e. decreased development time, increased body mass, or higher survival) compared to control larvae with shields intact.
• 3 Disturbance caused during shield removal did not adversely affect larvae because performance did not differ among controls (undisturbed larvae with intact shields), disturbance controls (shield removal simulated, but shield left in place), and larvae with shields removed.
• 4 Larvae without shields did not exhibit compensatory feeding to reconstruct the shield following its removal.
• 5 In a field experiment in which predators were excluded, larvae with shields removed did not develop faster than controls; in fact, survival was slightly reduced (10%) for larvae without shields and may have resulted from desiccation.
• 6 For slow-moving tortoise beetle larvae, the cost of bearing the shield is minimal. Thus, larval shields, composed of recycled waste products, provide an inexpensive mode of protection from some natural enemies.