Relationships among the basal lineages of Noctuidae (Lepidoptera,Noctuoidea) based on eight gene regions

In this study, we clarify the relationships between the basal lineages in the moth family Noctuidae using DNA sequence data from eight independent gene regions. Data matrices (6.4 kbp) are analysed using parsimony and model‐based methods (maximum likelihood and Bayesian inference). Our results support the family Noctuidae as a monophyletic group in which most subfamilies have hindwing vein M2 reduced or absent. Our phylogenetic hypothesis suggests that in the Noctuidae, the plesiomorphic condition is that in which vein M2 arises about one‐third of the way up the discocellular vein between the origins of M1 and M3, mainly parallel to M3, and is of thickness similar to vein M3. Most Noctuidae lineages possess an apomorphic (derived) condition in which hindwing vein M2 is markedly reduced or totally absent, so that the cubital vein appears to be three‐branched and these lineages are hence referred to as ‘trifine’. However, Noctuidae also include a number of lineages in which vein M2 is unreduced, or only slightly reduced, and these are more problematic for morphological association with the family Noctuidae. Our results also show that the subfamily Acronictinae is not closely related to Pantheinae, but instead shows a closer association with Amphipyrinae. Among the major lineages of Noctuidae, we postulate a general trend, with numerous exceptions, in larval host plants from woody plants in the basal groups towards herb feeding in derived groups. Similarly, the major radiations of monocot‐feeding groups within the family Noctuidae are in the higher trifines. The following taxonomic changes are proposed: Thiacidinae, syn. nov., a junior synonym of Pantheinae, and Dyopsinae, stat. nov., are reinstated as a subfamily.     

Further progress on the phylogeny of Noctuoidea (Insecta: Lepidoptera) using an expanded gene sample

Major progress has been made recently toward resolving the phylogeny of Noctuoidea, the largest superfamily of Lepidoptera. However, numerous questions and weakly supported nodes remain. In this paper we independently check and extend the main findings of multiple recent authors by performing maximum‐likelihood analyses of 5–19 genes (6.7–18.6 kb) in 74 noctuoids representing all the families and a majority of the subfamilies. Our results strongly support the six family system of Zahiri et al., with the former Lymantriidae and Arctiidae subsumed within the huge family Erebidae, and Noctuidae restricted largely to the subfamilies with so‐called trifine hindwing venation. Our data also strongly corroborate monophyly of the set of four families with quadrifid forewing venation, to the exclusion of Notodontidae, and removal from the latter of Oenosandridae. Other among‐family relationships, however, remain unsettled. Our evidence is equivocal on the position of Oenosandridae, which are sister group to either Notodontidae alone or to all other noctuoids. Like other recent nuclear gene studies, our results also provide no strong support for relationships among the four quadrifid forewing families. In contrast, within families our analyses significantly expand the list of robustly resolved relationships, while introducing no strong conflicts with previous molecular studies. Within Notodontidae, for which we present the largest molecular taxon sample to date, we find strong evidence for polyphyly for some, or all, recent definitions of the subfamilies Thaumetopoeinae, Pygaerinae, Notodontinae and Heterocampinae. Deeper divergences are incompletely resolved but there is strong support for multiple ‘backbone’ nodes subtending most of the subfamilies studied. Within Erebidae, we find much agreement and no strong conflict with a recent previous study regarding relationships among subfamilies, and somewhat stronger support. Although many questions remain, the two studies together firmly resolve positions for over half the subfamilies. Within Noctuidae, we find no strong conflict with previous molecular studies regarding relationships among subfamilies, but much stronger resolution along the ‘backbone’ of the phylogeny. Combining information from multiple studies yields strongly resolved positions for most of the subfamilies. Finally, our results strongly suggest that the tribes Pseudeustrotiini and Prodeniini, currently assigned to the largest subfamily, Noctuinae, do not belong there. In sum, our results provide additional corroboration for the main outlines of family‐level phylogeny in Noctuoidea, and contribute toward resolving relationships within families.     

Towards resolving and redefining Amphipyrinae (Lepidoptera,Noctuoidea, Noctuidae): a massively polyphyletic taxon

Amphipyrinae have long been a catchall taxon for Noctuidae, with most members lacking discernible morphological synapomorphies that would allow their assignment to one of the many readily diagnosable noctuid subfamilies. Here data from seven gene regions (> 5500 bp) for more than 120 noctuid genera are used to infer a phylogeny for Amphipyrinae and related subfamilies. Sequence data for 57 amphipyrine genera – most represented by the type species of the genus – are examined. We present here the first large‐scale molecular phylogenetic study of Amphipyrinae and the largest molecular phylogeny of Noctuidae to date; several proposed nomenclatural changes for well‐supported results; and the identification of areas of noctuid phylogeny where greater taxon sampling and/or genomic‐scale data are needed. Adult and larval morphology, along with life‐history traits, for taxonomic groupings most relevant to the results are discussed. Amphipyrinae are significantly redefined; many former amphipyrines, excluded as a result of these analyses, are reassigned to other noctuid subfamily‐level taxa. Four genera, Chamaeclea Grote, Heminocloa Barnes & Benjamin, Hemioslaria Barnes & Benjamin and Thurberiphaga Dyar, are transferred to the tribe Chamaecleini Keegan & Wagner tribe n. in Acontiinae. Stiriina is elevated to Stiriinae rev. stat. , Grotellina is elevated to Grotellinae rev. stat. and Annaphilina is elevated to Annaphilini rev. stat. Acopa Harvey is transferred to Bryophilinae, Aleptina Dyar is transferred to Condicinae, Leucocnemis Hampson and Oxycnemis gracillinea (Grote) are transferred to Oncocnemidinae, Nacopa Barnes & Benjamin is transferred to Noctuinae and Narthecophora Smith is transferred to Stiriinae. Azenia Grote (and its subtribe Azeniina), Cropia Walker, Metaponpneumata Möschler, Sexserrata Barnes & Benjamin and Tristyla Smith are transferred to Noctuidae incertae sedis. Hemigrotella Barnes & McDunnough (formerly in subtribe Grotellina) is retained in Amphipyrinae. Argentostiria Poole and Bistica Dyar are retained in Stiriini but removed from incertae sedis position. This published work has been registered on ZooBank: http://zoobank.org/urn:lsid:zoobank.org:pub:4A140782‐31BA‐445A‐B7BA‐6EAB98ED43FA .     

Retransmission of echo-like signals: methods and results of studies on nocturnal moths (Insecta)

Studies of capability to echolocate in nocturnal moths using the method of echo-signal retransmission are described in detail. During retransmission the insect was presented not with the echo from some real object but with electronically generated echo-like signal which appears following the certain delay after insect's own acoustic signal. In that way the artificial echo from non-existent obstacle is produced. In the current study the delay was set to 0.5 ms that corresponds to a distance of about 8 cm from obstacle. At the same time there was no any single movement of any object near the flying insect therefore the activities of sensory systems other than auditory did not cause systematical influence on results obtained. Noctuid moths (Noctuidae, Lepidoptera) are acoustically active insects. They are capable of producing ultrasonic clicks during flight. The most typical response of a moth to the retransmission of echo-like stimuli is an activation of own clicks emission that is often accompanied with rise in click amplitude. Using the activation of emission as a criterion of moth's response to the echo-like stimulation we measured echolocational thresholds in three species: Amphipyra pyramidea (36 dB SPL), Enargia paleacea (31 dB SPL) and Blepharita satura (26 dB SPL). The ability to echolocate was also demonstrated in 20 species of subfamilies Catocalinae, Amphipyrinae, Cuculliinae, Hadeninae, Noctuinae, Heliothinae.     

Phylogenetic relationships of Acronictinae with discussion of the abdominal courtship brush in Noctuidae (Lepidoptera)

We present results of an eight‐gene molecular study of the subfamily Acronictinae and related Noctuidae. Amphipyrinae are recovered as sister to Acronictinae, but with weak support – not surprisingly, the content of the two subfamilies has often been mixed in classifications. Balsinae, previously placed near Acronictinae or within Noctuinae, is recovered within an unresolved polytomy of Cuculliinae, Eustrotiinae, Raphiinae and Dilobinae. Gerbathodes Warren, Moma Hübner and Nacna Fletcher are excluded from Acronictinae. Three genera recently transferred into the subfamily – Cerma Hübner, Chloronycta Schmidt & Anweiler and Comachara Franclemont – are confirmed as acronictines. Lophonycta Sugi (the type genus of Lophonyctinae) is returned to the Acronictinae. Sinocharis Püngeler, formerly considered to be Acontiinae or as the basis of its own subfamily Sinocharinae, is nested within early diverging Acronictinae genera. Both subfamilies are formally synonymized: i.e. Lophonyctinae syn.n. and Sinocharinae syn.n. Nine acronictine genus‐level taxa were found to nest within the nominate genus Acronicta Ochsenheimer: Eogena Guenée, Hyboma Hübner, Hylonycta Sugi, Jocheaera Hübner, Oxicesta Hübner, Simyra Ochsenheimer, Subacronicta Kozhanchikov, Triaena Hübner, and Viminia Chapman. Eogena, Oxicesta, and Simyra, currently treated as valid genera, nest within terminal clades of the genus Acronicta and are here subsumed within the genus: Eogena syn.n. , Oxicesta syn.n. and Simyra syn.n. Four well‐supported species groups within Acronicta are identified: the alni clade, the leporina clade, the nervosa clade and the psi clade. While many previous treatments have stated explicitly that Acronictinae lack abdominal scent brushes, or excluded genera with brushes from the subfamily, we show that well‐developed brushes are present in three early diverging acronictine genera: Cerma, Lophonycta, and Sinocharis. We illustrate and describe the brushes of all three genera, and briefly review the taxonomic distribution of the anterior abdominal courtship brushes in Noctuidae, emphasizing the labile evolutionary distribution of these structures.     

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.     

Theraphosidae phylogeny: relationships of the ‘Ischnocolinae’ genera (Araneae,Mygalomorphae)

The mygalomorph spider subfamily ‘Ischnocolinae’ was originally established as a group based on the presence of divided tarsal scopula. Later, the divided condition of the scopula was considered the plesiomorphic state, which could not support the monophyly of ‘Ischnocolinae’. In Raven 1985, the subfamily was considered paraphyletic, pending a phylogenetic analysis to reinvestigate monophyletic groups. This study comprises such a phylogenetic analysis, based on morphological data, that includes representatives of all genera currently included in ‘Ischnocolinae’ as well as representatives of all other nine Theraphosidae subfamilies (Thrigmopoeinae, Ornithoctoninae, Eumenophorinae, Stromatopelminae, Harpactirinae, Selenogyrinae, Theraphosinae, Aviculariinae and Selenocosmiinae). The family Theraphosidae is considered monophyletic and expanded to include three additional genera previously considered as possible Barychelidae, namely Brachionopus (as Harpactirinae), Trichopelma and Reichlingia (as Ischnocolinae sensu stricto) while ‘Ischnocolinae’ as previously defined does not appear as monophyletic. However, two monophyletic groups were defined as subfamilies to include some former ‘Ischnocolinae’ representatives. The first group includes Acanthopelma rufescens, Trichopelma nitidum, Reichlingia annae, Ischnocolus spp., Holothele rondoni, Holothele culebrae and Holothele aff culebrae and is hereby named as Ischnocolinae (sensu stricto). The other subgroup comprises Sickius longibulbi, Holothele incei, Holothele aff incei, Guyruita spp., Schismatothele lineata, Hemiercus modestus, Holothele colonica and Holothele sp., together established as Schismatothelinae subfam. nov. Several genera included in former ‘Ischnocolinae’ appear as monophyletic (Catumiri, Oligoxystre, Heterothele, Nesiergus, Chaetopelma, Ischnocolus, Guyruita and Plesiophrictus). However, the genera Holothele, Schismatothele and Hemiercus deserve more attention in order to evaluate their intrarelationships and inclusion of species.     

The phylogeny of leaf beetles (Chrysomelidae) inferred from mitochondrial genomes

The high-level classification of Chrysomelidae (leaf beetles) currently recognizes 12 or 13 well-established subfamilies, but the phylogenetic relationships among them remain ambiguous. Full mitochondrial genomes were newly generated for 27 taxa and combined with existing GenBank data to provide a dataset of 108 mitochondrial genomes covering all subfamilies. Phylogenetic analysis under maximum likelihood and Bayesian inference recovered the monophyly of all subfamilies, except that Timarcha was split from Chrysomelinae in some analyses. Three previously recognized major clades of Chrysomelidae were broadly supported: the ‘chrysomeline’ clade consisting of (Chrysomelinae (Galerucinae + Alticinae)); the ‘sagrine’ clade with internal relationships of ((Bruchinae + Sagrinae) + (Criocerinae + Donaciinae)), and the ‘eumolpine’ clade comprising (Spilopyrinae (Cassidinae (Eumolpinae (Cryptocephalinae + Lamprosomatinae)))). Relationships among these clades differed between data treatments and phylogenetic algorithms, and were complicated by two additional deep lineages, Timarcha and Synetinae. Various topological tests favoured the PhyloBayes software as the preferred inference method, resulting in the arrangement of (chrysomelines (eumolpines + sagrines)), with Timarcha placed as sister to the chrysomeline clade and Synetinae as a deep lineage splitting near the base. Whereas mitogenomes provide a solid framework for the phylogeny of Chrysomelidae, the basal relationships do not agree with the topology of existing molecular studies and remain one of the most difficult problems of Chrysomelidae phylogenetics.     

The phylogeny of Sericini and their position within the Scarabaeidae based on morphological characters (Coleoptera: Scarabaeidae)

Abstract. To reconstruct the phylogeny of the Sericini and their systematic position among the scarabaeid beetles, cladistic analyses were performed using 107 morphological characters from the adults and larvae of forty‐nine extant scarabaeid genera. Taxa represent most ‘traditional’ subfamilies of coprophagous and phytophagous Scarabaeidae, with emphasis on the Sericini and other melolonthine lineages. Several poorly studied exoskeletal features have been examined, including the elytral base, posterior wing venation, mouth parts, endosternites, coxal articulation, and genitalia. The results of the analysis strongly support the monophyly of the ‘orphnine group’ + ‘melolonthine group’ including phytophagous scarabs such as Dynastinae, Hopliinae, Melolonthinae, Rutelinae, and Cetoniinae. This clade was identified as the sister group to the ‘dung beetle line’ represented by Aphodius + Copris. The ‘melolonthine group’ is comprised in the strict consensus tree by two major clades and two minor lineages, with the included taxa of Euchirinae, Rutelinae, and Dynastinae nested together in one of the major clades (‘melolonthine group I’). Melolonthini, Cetoniinae, and Rutelinae are strongly supported, whereas Melolonthinae and Pachydemini appear to be paraphyletic. Sericini + Ablaberini were identified to be sister taxa nested within the second major melolonthine clade (‘melolonthine group II’). As this clade is distributed primarily in the southern continents, one could assume that Sericini + Ablaberini are derived from a southern lineage. Plausibly, ancestors of Sericini + Ablaberini and Athlia were separated by a vicariance event, such as the separation of the African plate from the rest of Gondwana, whereas Sericini and Ablaberini probably diversified during the early Tertiary, with dispersal of some basal Sericini to South America.     

Molecular phylogeny of the fishes traditionally referred to Cyprinini sensu stricto (Teleostei: Cypriniformes)

Yang, L., Mayden, R. L., Sado, T., He, S., Saitoh, K. & Miya, M. (2010). Molecular phylogeny of the fishes traditionally referred to Cyprinini sensu stricto (Teleostei: Cypriniformes). —Zoologica Scripta, 39, 527–550. Carps (e.g. Koi) of the genus Cyprinus and Crucian carps (e.g. Goldfish) of the genus Carassius are among the most popular freshwater fishes around the world. However, their phylogenetic positions within the subfamily Cyprininae, relationships with their allies (e.g. Procypris, Carassioides), and the monophyly of the group formed by them and their allies, which is referred as the tribe Cyprinini sensu stricto, are far from clear. Historically, the Cyprinini was defined by different people according to whether a cyprinine fish possessed a spinous anal‐fin ray (or anal spine), the spine was serrated or not, and occasionally, the number of branched dorsal‐fin rays. Some definitions were established without providing any diagnostic characters. In this study, we investigated the monophyly of the tribe Cyprinini sensu stricto, based on four different historical definitions, and explored the phylogenetic relationships of these members in the subfamily Cyprininae. Using five mitochondrial genes as markers, both maximum‐likelihood and Bayesian trees were constructed using the optimal partitioning strategy. Both analyses successfully resolved a monophyletic Cyprininae and recovered seven major clades from this subfamily. The diagnosis limiting the tribe Cyprinini sensu stricto to four genera, Cyprinus, Carassius, Carassioides and Procypris, received most support. We propose that only those cyprinines that possess a serrated anal spine and have no <10 branched dorsal‐fin rays should be considered members of this tribe. Cyprinini is sister to the Sinocyclocheilus clade, a group traditionally considered a barbin, and together they form the ‘Cyprinini‐Sinocyclocheilus’ clade. Procypris forms the basal clade of the Cyprinini, whereas species of Carassius and Carassioides locate at the top.     

Laying the foundations of evolutionary and systematic studies in crickets (Insecta,Orthoptera): a multilocus phylogenetic analysis

Orthoptera have been used for decades for numerous evolutionary questions but several of its constituent groups, notably crickets, still suffer from a lack of a robust phylogenetic hypothesis. We propose the first phylogenetic hypothesis for the evolution of crickets sensu lato, based on analysis of 205 species, representing 88% of the subfamilies and 71% tribes currently listed in the database Orthoptera Species File (OSF). We reconstructed parsimony, maximum likelihood and Bayesian phylogenies using fragments of 18S, 28SA, 28SD, H3, 12S, 16S, and cytb (~3600 bp). Our results support the monophyly of the cricket clade, and its subdivision into two clades: mole crickets and ant‐loving crickets on the one hand, and all the other crickets on the other (i.e. crickets sensu stricto). Crickets sensu stricto form seven monophyletic clades, which support part of the OSF families, “subfamily groups”, or subfamilies: the mole crickets (OSF Gryllotalpidae), the scaly crickets (OSF Mogoplistidae), and the true crickets (OSF Gryllidae) are recovered as monophyletic. Among the 22 sampled subfamilies, only six are monophyletic: Gryllotalpinae, Trigonidiinae, Pteroplistinae, Euscyrtinae, Oecanthinae, and Phaloriinae. Most of the 37 tribes sampled are para‐ or polyphyletic. We propose the best‐supported clades as backbones for future definitions of familial groups, validating some taxonomic hypotheses proposed in the past. These clades fit variously with the morphological characters used today to identify crickets. Our study emphasizes the utility of a classificatory system that accommodates diagnostic characters and monophyletic units of evolution. Moreover, the phylogenetic hypotheses proposed by the present study open new perspectives for further evolutionary research, especially on acoustic communication and biogeography.     

Phylogenetic relationships of the Suidae (Mammalia,Cetartiodactyla): new insights on the relationships within Suoidea

Orliac, M. J., Antoine, P. ‐O., Ducrocq, S. (2010). Phylogenetic relationships of the Suidae (Mammalia, Cetartiodactyla): new insights on the relationships within Suoidea. —Zoologica Scripta, 39, 315–330. In most analyses, both molecular and morphological phylogenies of the Cetartiodactyla support the monophyly of Suoidea. However, the evolutionary history of this superfamily remains poorly known primarily due to long‐lasting debates about the taxonomic content and relationships of the suoid families and subfamilies. Despite their crucial position in the reconstruction of the phylogeny of Cetartiodactyla, Suoidea themselves have received little attention in those phylogenies, and no extensive analysis of the group has been performed so far. We therefore examine the phylogeny of the Suidae through the first phylogenetic analysis of Suoidea, including recent and fossil representatives of all four putative families. The results support the monophyly of the traditional suid subfamilies and indicate the Sanitheriidae as sister taxon to the Suidae clade. The evolutionary history within Suidae reveals its complexity, with major convergences involving important morphological structures such as the auditory region or the upper male canine. Divergent signals gathered from either dental or cranio‐mandibular features are responsible for two long‐lasting unresolved issues within Suoidea: the question of the relationships between ‘Old World’ and ‘New World’ peccaries remaining unsolved, as well as the position and familial status of the mid‐Tertiary tayassuid Perchoerus.     

Multiple nuclear and mitochondrial DNA sequences provide new insights into the phylogeny of South African Lacertids (Lacertidae,Eremiadinae)

Eremiadinae, one of three subfamilies of Lacertidae, are distributed throughout Asia and Africa. Previous phylogenetic studies suggested that one of the main groups of Eremiadinae (the Ethiopian clade) consist of two clades with predominately East‐African and South‐African distribution. Yet, especially the latter one, which includes the genera Pedioplanis, Meroles, Ichnotropis, Tropidosaura and Australolacerta, was not well supported in the molecular phylogenetic analysis. In this study, we analysed the phylogenetic relationships among the genera of the ‘South African clade’ to assess whether this group actually forms a highly supported clade and to address questions concerning the monophyly of the genera. We sequenced sections of the widely used mitochondrial genes coding for 16S rRNA, 12S rRNA and cytochrome b (altogether 2045 bp) as well as the nuclear genes c‐mos, RAG‐1, PRLR, KIF24, EXPH5 and RAG‐2 (altogether 4473 bp). The combined data set increased the support values for several nodes considerably. Yet, the relationships among five major lineages within the ‘South African clade’ are not clearly resolved even with this large data set. We interpret this as a ‘hard polytomy’ due to fast radiation within the South African lacertids. The combined tree based on nine marker genes provides strong support for the ‘South African Clade’ and its sister group relationship with the ‘East African Clade’. Our results confirm the genus Tropidosaura as a monophylum, while Ichnotropis is paraphyletic in our trees: Ichnotropis squamulosa appears more closely related to Meroles than to Ichnotropis capensis. Furthermore, the monophyly of Meroles is questionable as well. Based on our results, I. squamulosa should be transferred from Ichnotropis into the genus Meroles. Also, the two species of Australolacerta (A. australis and A. rupicola) are very distantly related and the genus is perhaps paraphyletic, too. Finally we propose a phylogeographical scenario in the context of palaeoclimatic data and compare it with a previously postulated hypothesis.     

Molecular phylogenetics of heliothine moths (Lepidoptera: Noctuidae: Heliothinae), with comments on the evolution of host range and pest status

Abstract The Heliothinae are a cosmopolitan subfamily of about 365 species that include some of the world’s most injurious crop pests. This study re‐assesses evolutionary relationships within heliothines, providing an improved phylogeny and classification to support ongoing intensive research on heliothine genomics, systematics, and biology. Our phylogeny estimate is based on two nuclear gene regions, namely elongation factor‐1α (EF‐1α; 1240 bp) and dopa decarboxylase (DDC; 687 bp), and on the barcoding region of mitochondrial cytochrome oxidase I (COI; 708 bp), providing a total of 2635 bp. These were sequenced for 71 heliothines, representing all major genera and nearly all recognized subgenera and species groups, and for 16 outgroups representing all major lineages of trifine Noctuidae. Analysis of the combined data by maximum likelihood, unweighted parsimony and Bayesian methods gave nearly identical topologies, and the individual gene trees showed only one case of potentially strong conflict. Relationships among genera and subgenera are resolved with strong bootstrap support. The earliest‐diverging lineages (c. 200 species in total) consist almost entirely of host specialists, reflecting the inferred ancestral heliothine host range under parsimony. The remaining species form a clade – the Heliothis group – that includes most of the polyphages (30% of heliothines) and all of the major pests. Many other species in the Heliothis group, however, are host specialists. Our results extend previous efforts to subdivide this large clade, and show the most notorious pest groups, the corn earworm complex (Helicoverpa) and the tobacco budworm (Heliothis virescens) group, to be closely related, joining with a small oligophagous genus in what we term the major‐pest lineage. Thus, genomic/experimental results from one model pest may extrapolate well to other pest species. The frequency of evolutionary expansion and contraction in host range appears to increase dramatically at the base of the Heliothis group, in contrast to the case for earlier‐diverging lineages. We ascribe this difference provisionally to differential evolutionary constraints arising from contrasting life‐history syndromes. Host‐specific behaviour and crypsis, coupled with low fecundity and vagility, may discourage host‐range expansion in earlier‐diverging lineages. By contrast, in the Heliothis group, the absence of host‐specific traits, coupled with high vagility and fecundity, may more readily permit expansion or contraction of the host range in response to varying ecological pressures such as host species abundance or differential competition and predation.     

利用叶绿体基因组数据解析锦葵科梧桐亚科的系统位置和属间关系

分子系统学研究将传统梧桐科与锦葵科、木棉科和椴树科合并为广义锦葵科,并进一步分为9个亚科.然而,9个亚科之间的关系尚未完全明确,且梧桐亚科内的属间关系也未得到解决.为了明确梧桐亚科在锦葵科中的系统发育位置,厘清梧桐亚科内部属间系统发育关系,该研究对锦葵科8个亚科进行取样,共选取55个样本,基于叶绿体基因组数据,采用最大...     

Morphology, molecules, and character congruence in the phylogeny of South American geophagine cichlids (Perciformes, Labroidei)

Phylogenetic relationships among the Neotropical cichlid subfamily Geophaginae were examined using 136 morphological characters and a molecular dataset consisting of six mitochondrial and nuclear genes. Topologies produced by morphological and combined data under parsimony were contrasted, congruence among different partitions was analysed, and potential effects of character incongruence and patterns of geophagine evolution on phylogenetic resolution are discussed. Interaction of morphological and molecular characters in combined analysis produced better resolved and supported topologies than when either was analysed separately. Combined analyses recovered a strongly supported Geophaginae that was closely related to Cichlasomatinae. Within Geophaginae, two sister clades included all geophagine genera. Acarichthyini (Acarichthys+Guianacara) was sister to the ‘B clade’, which contained the ‘Geophagus clade’ (‘Geophagus’steindachneri+Geophagus sensu stricto, and both sister to Gymnogeophagus) as sister to the ‘Mikrogeophagus clade’ (Mikrogeophagus+‘Geophagus’brasiliensis), and in turn, the Geophagus and Mikrogeophagus clades were sister to the crenicarine clade (Crenicara+Dicrossus) and Biotodoma. The second geophagine clade included the ‘Satanoperca clade’ (Satanoperca+Apistogramma and Taeniacara) as sister to the ‘Crenicichla clade’ (Crenicichla+Biotoecus). Several lineages were supported by unique morphological synapomorphies: the Geophaginae + Cichlasomatinae (5 synapomorphies), Geophaginae (1), Crenicichla clade (3), crenicarine clade (1), the sister relationship of Apistogramma and Taeniacara (4) and of Geophagus sensu stricto and‘Geophagus’steindachneri (1), and the cichlasomine tribe Heroini (1). Incorporation of Crenicichla in Geophaginae reconciles formerly contradictory hypotheses based on morphological and molecular data, and makes the subfamily the most diverse and ecologically versatile clade of cichlids outside the African great lakes. Results of this study support the hypothesis that morphological differentiation of geophagine lineages occurred rapidly as part of an adaptive radiation.     

Multi‐gene phylogenetic analysis of south‐east Asian pest members of the Bactrocera dorsalis species complex (Diptera: Tephritidae) does not support current taxonomy

Bactrocera dorsalis sensu stricto, B. papayae, B. philippinensis and B. carambolae are serious pest fruit fly species of the B. dorsalis complex that predominantly occur in south‐east Asia and the Pacific. Identifying molecular diagnostics has proven problematic for these four taxa, a situation that cofounds biosecurity and quarantine efforts and which may be the result of at least some of these taxa representing the same biological species. We therefore conducted a phylogenetic study of these four species (and closely related outgroup taxa) based on the individuals collected from a wide geographic range; sequencing six loci (cox1, nad4‐3′, CAD, period, ITS1, ITS2) for approximately 20 individuals from each of 16 sample sites. Data were analysed within maximum likelihood and Bayesian phylogenetic frameworks for individual loci and concatenated data sets for which we applied multiple monophyly and species delimitation tests. Species monophyly was measured by clade support, posterior probability or bootstrap resampling for Bayesian and likelihood analyses respectively, Rosenberg's reciprocal monophyly measure, P(AB), Rodrigo's (P(RD)) and the genealogical sorting index, gsi. We specifically tested whether there was phylogenetic support for the four ‘ingroup’ pest species using a data set of multiple individuals sampled from a number of populations. Based on our combined data set, Bactrocera carambolae emerges as a distinct monophyletic clade, whereas B. dorsalis s.s., B. papayae and B. philippinensis are unresolved. These data add to the growing body of evidence that B. dorsalis s.s., B. papayae and B. philippinensis are the same biological species, which poses consequences for quarantine, trade and pest management.     

Origin and diversification of the cryptic ant genus Stenamma Westwood (Hymenoptera: Formicidae), inferred from multilocus molecular data,biogeography and natural history

The genus Stenamma Westwood comprises a group of cryptic, cold tolerant ants that occur throughout the Holarctic and Middle American regions. Traditional approaches to taxonomy and phylogeny are confounded by multiple factors, including the conservative and often convergent morphology of workers and the rarity of reproductive castes in collections. Monophyly of Stenamma and relationships within the genus are uncertain as nearly all previous taxonomic work has been regional in scope. Furthermore, the sister group to Stenamma has not been well established. Here an extensive molecular dataset consisting of ten genes (～8 kb of data), 48 ingroup taxa (20 Nearctic, 6 Palaearctic and 22 Neotropical) and 8 outgroup taxa (6 closely related non‐Stenamma and 2 additional myrmicines) is used to investigate the broad‐scale phylogeny and evolutionary history of Stenamma. Phylogenetic analysis is performed under maximum likelihood and Bayesian frameworks on individual genes and several alternate concatenated datasets, which are used to investigate the effects of inclusion or exclusion of COI and intronic regions. The timing of Stenamma evolution is inferred in beast and ancestral areas are reconstructed using both the s‐diva and DEC methods, as implemented in the programs rasp and lagrange , respectively. Stenamma is revealed as monophyletic with high support and tentatively is sister to a group of New World species placed currently in Aphaenogaster Mayr and Messor Forel. Within Stenamma, two major clades are recovered: a ‘Holarctic clade’ (HOC) and a ‘Middle American clade’ (MAC). The HOC consists of the European S. striatulum Emery sister to two well‐supported groups, the informal ‘debile’ and ‘brevicorne’ clades. The ‘brevicorne’ clade is entirely Nearctic, whereas the ‘debile’ clade includes both Nearctic and Palaearctic representatives. The MAC occurs from the southern United States to northern South America and, with the exception of S. huachucanum Smith, is almost completely isolated geographically from the HOC. It includes a depauperate northern clade and the ‘MAC core’, which is a diverse assemblage of wet forest adapted species found throughout Central America. Divergence dating and biogeographic reconstruction show that Stenamma is most likely to have originated in the Nearctic at the Eocene–Oligocene boundary (～35 Ma) and diversified more rapidly at 16 and 8 Ma for the HOC and MAC, respectively. Potential environmental factors affecting the evolution of Stenamma include the intense global cooling of the late Eocene combined with aridification and mountain building. The phylogenetic results are discussed in relation to the current Stenamma species groups and several new morphological characters are presented to help in identification.     

Phylogeny and systematics of mitriform gastropods (Mollusca: Gastropoda: Neogastropoda)

With about 800 Recent species, ‘miters’ are a widely distributed group of tropical and subtropical gastropods that are most diverse in the Indo‐West Pacific. They include the two families Mitridae and Costellariidae, similar in shell morphology and traditionally treated as close relatives. Some genera of deep‐water Ptychatractidae and Volutomitridae are close to miters in shell morphology, and the term ‘mitriform gastropods’ has been introduced to refer to Mitridae, Costellariidae, and this assortment of convergent forms. The present study aimed at the reconstruction of phylogenetic relationships of mitriform gastropods based on representative taxon sampling. Four genetic markers [cytochrome c oxidase subunit I (COI), 16S and 12S rRNA mitochondrial genes, and H3 (Histone 3) nuclear gene] were sequenced for over 90 species in 20 genera, and the molecular data set was supplemented by studies of radula morphology. Our analysis recovered Mitridae as a monophyletic group, whereas the genus Mitra was found to be polyphyletic. Of 42 mitrid species included in the analysis, 37 formed a well‐supported ‘core Mitridae’ consisting of four major clades, three of them consistent with the subfamilies Cylindromitrinae, Imbricariinae, and Mitrinae, and Strigatella paupercula standing out by itself. Basal to the ‘core Mitridae’ are four minor lineages, with the genus Charitodoron recognized as sister group to all other Mitridae. The deep‐water family Pyramimitridae shows a sister relationship to the Mitridae, with high support for a Pyramimitridae + Mitridae clade. Our results recover the monophyly of the Costellariidae, which form a well‐supported clade that also includes Ptychatractidae, Columbariinae, and Volutomitridae, but not Mitridae. Most derived and diverse amongst Costellariidae are species of Vexillum, characterized by a bow‐shaped, multicuspidate rachidian tooth. Several previously unrecognized deep‐water costellariid lineages are revealed. Their members retain some plesiomorphies – in particular a tricuspidate rachidian tooth – that makes them morphologically intermediate between ptychatractids and Vexillum. The taxa of Ptychatractidae included in the analysis are not monophyletic, but form three well‐supported, unrelated groupings, corresponding respectively to Ceratoxancus + Latiromitra, Exilia, and Exiliodea. None of them shows an affinity to Pseudolividae. © 2015 The Linnean Society of London     

Testudodinium gen. nov. (Dinophyceae), a new genus of sand‐dwelling dinoflagellates formerly classified in the genus Amphidinium

A new genus of sand‐dwelling photosynthetic dinoflagellate, Testudodinium Horiguchi, Tamura, Katsumata et A. Yamaguchi is proposed based on Testudodinium testudo (Herdman) Horiguchi, Tamura, Katsumata, et A. Yamaguchi comb. nov. (Basionym: Amphidinium testudo Herdman) and a new species in this new genus, Testudodinium maedaense Katsumata et Horiguchi sp. nov. is described. Amphidinium corrugatum is also transferred to this genus, making a new combination T. corrugatum (Larsen et Patterson) Horiguchi, Tamura et A. Yamaguchi. These three species are similar to the members of the genus Amphidinium in having an extremely small episome and a dorsoventrally flattened cell body. They are, however, distinguished from the genus Amphidinium seusu stricto by the possession of a distinct longitudinal furrow in the middle of ventral side of the episome. Phylogenetic trees based on small subunit (SSU) rDNA revealed that all three of these Testudodinium species formed a robust clade and, although statistical support is not high, the tree suggests Testudodinium clade is not closely related to Amphidinium seusu stricto clade. The morphological differences together with molecular data support the establishment of a new genus for A. testudo and its related species.