Geometrid moths of the genus Eupithecia Curtis, 1825 (Lepidoptera,Geometridae): Prerequisites and characteristic features of high species diversity

The prerequisites and characteristics of high species diversity of the genus Eupithecia (Geometridae), the largest in the order Lepidoptera, are discussed. The surge of speciation in this genus was facilitated by the evolutionarily advanced morpho-functional characters of the adults and larvae, as well as by the plasticity of a number of morphological structures, especially those of the genitalia in both sexes, in the ancestral Eupithecia. The wide range of flowering plants has served as a basis for close associations of the larvae predominantly with generative plant organs and for broad adaptive radiation of Eupithecia into a variety of ecological niches in the areas lacking their competitors from other genera of the tribe Eupitheciini. Transition of the larvae of some Eupithecia to facultative or obligate ambush predation was facilitated by adaptation of the antho- and carpophagous species to high-calorie food rich in proteins, lipids, and carbohydrates. Based on the distribution of the genus Eupithecia and its most morphologically primitive species-groups, its East Asian origin is hypothesized. It is noted that many of the largest genera of animals and plants include relatively small-sized species.     

Taxonomy of the Cidariini (Lepidoptera, Geometridae) in Korea (I)

ABSTRACT Among forty species of the Korean Cidariini, a tribe of Larentiinae (Lepidoptera, Geometridae), nineteen species of ten genera are revised: Ecliptopera Warren, Lampropteryx Stephens, Eustroma Hübner, Eveeliptopera Inoue, Lobogonodes Bastelberger, Hysterura Warren, Sibatania Inoue, Eulithis Hiibner, Gandaritis Moore, and Electrophaes Prout. The diagnostic characters and monophyly of each genus are provided. Figures of adults including male and female genitalia, and distribution maps in Korea are also provided.     

Biology and host range of Tecmessa elegans (Lepidoptera: Notodontidae), a leaf-feeding moth evaluated as a potential biological control agent for Schinus terebinthifolius (Sapindales: Anacardiaceae) in the United States

During surveys for natural enemies that could be used as classical biological control agents of Schinus terebinthifolius Raddi (Brazilian pepper), the caterpillar, Tecmessa elegans Schaus (Lepidoptera: Notodontidae), was recorded feeding on the leaves of the shrub in South America. The biology and larval and adult host range of this species were examined to determine the insect's suitability for biological control of this invasive weed in North America and Hawaii. Biological observations indicate that the larvae have five instars. When disturbed, the late instar larvae emit formic acid from a prothoracic gland that may protect larvae from generalist predators. Larval host range tests conducted both in South and North America indicated that this species feeds and completes development primarily on members of the Anacardiaceae within the tribe Rhoeae. Oviposition tests indicated that when given a choice in large cages the adults will select the target weed over Pistacia spp. However, considering the many valued plant species in its host range, especially several North American natives, this species will not be considered further for biological control of S. terebinthifolius in North America.     

Evaluation of pheromone-baited traps for winter moth and Bruce spanworm (Lepidoptera: Geometridae)

We tested different pheromone-baited traps for surveying winter moth, Operophtera brumata (L.) (Lepidoptera: Geometridae), populations in eastern North America. We compared male catch at Pherocon 1C sticky traps with various large capacity traps and showed that Universal Moth traps with white bottoms caught more winter moths than any other trap type. We ran the experiment on Cape Cod, MA, where we caught only winter moth, and in western Massachusetts, where we caught only Bruce spanworm, Operophtera bruceata (Hulst) (Lepidoptera: Geometridae), a congener of winter moth native to North America that uses the same pheromone compound [(Z,Z,Z)-1,3,6,9-nonadecatetraene] and is difficult to distinguish from adult male winter moths. With Bruce spanworm, the Pherocon 1C sticky traps caught by far the most moths. We tested an isomer of the pheromone [(E,Z,Z)-1,3,6,9-nonadecatetraene] that previous work had suggested would inhibit captures of Bruce spanworm but not winter moths. We found that the different doses and placements of the isomer suppressed captures of both species to a similar degree. We are thus doubtful that we can use the isomer to trap winter moths without also catching Bruce spanworm. Pheromone-baited survey traps will catch both species.     

Comprehensive molecular sampling yields a robust phylogeny for geometrid moths (Lepidoptera: Geometridae)

Background

The moth family Geometridae (inchworms or loopers), with approximately 23 000 described species, is the second most diverse family of the Lepidoptera. Apart from a few recent attempts based on morphology and molecular studies, the phylogeny of these moths has remained largely uninvestigated.

Methodology/Principal Findings

We performed a rigorous and extensive molecular analysis of eight genes to examine the geometrid affinities in a global context, including a search for its potential sister-taxa. Our maximum likelihood analyses included 164 taxa distributed worldwide, of which 150 belong to the Geometridae. The selected taxa represent all previously recognized subfamilies and nearly 90% of recognized tribes, and originate from all over world. We found the Geometridae to be monophyletic with the Sematuridae+Epicopeiidae clade potentially being its sister-taxon. We found all previously recognized subfamilies to be monophyletic, with a few taxa misplaced, except the Oenochrominae+Desmobathrinae complex that is a polyphyletic assemblage of taxa and the Orthostixinae, which was positioned within the Ennominae. The Sterrhinae and Larentiinae were found to be sister to the remaining taxa, followed by Archiearinae, the polyphyletic assemblage of Oenochrominae+Desmobathrinae moths, Geometrinae and Ennominae.

Conclusions/Significance

Our study provides the first comprehensive phylogeny of the Geometridae in a global context. Our results generally agree with the other, more restricted studies, suggesting that the general phylogenetic patterns of the Geometridae are now well-established. Generally the subfamilies, many tribes, and assemblages of tribes were well supported but their interrelationships were often weakly supported by our data. The Eumeleini were particularly difficult to place in the current system, and several tribes were found to be para- or polyphyletic.     

Polymorphism in the sperm ultrastructure among four species of lizards in the genus Tupinambis (Squamata: Teiidae)

We describe, for the first time, the ultrastructure of the spermatozoa of four species of the genus Tupinambis (Squamata, Teiidae). We identified seven polymorphic characters within this genus: the presence and shape of the perforatorial base plate, the presence of the epinuclear lucent zone, the presence of a unilateral ridge in the acrosome, the presence of a central density within the proximal centriole, the number of mitochondria and dense-bodies sets, and the shape of mitochondria. We analysed the evolution of the seven polymorphic characters by mapping them onto a current phylogeny of the species of Tupinambis , using the teiids Ameiva ameiva and Cnemidophorus sexlineatus as outgroups. Our results indicate that sperm ultrastucture characters, although of great value for phylogeny at higher taxonomic levels in reptiles and other groups, are poor predictors of phylogeny when considering the species of Tupinambis studied here. We failed to identify evidences that homoplasy in sperm ultrastructure among the species of Tupinambis is due to convergent adaptation, suggesting that the polymorphism may be selectively neutral in this group.     

Integrated Fossil and Molecular Data Reveal the Biogeographic Diversification of the Eastern Asian-Eastern North American Disjunct Hickory Genus (Carya Nutt.)

The hickory genus (Carya) contains ca. 17 species distributed in subtropical and tropical regions of eastern Asia and subtropical to temperate regions of eastern North America. Previously, the phylogenetic relationships between eastern Asian and eastern North American species of Carya were not fully confirmed even with an extensive sampling, biogeographic and diversification patterns had thus never been investigated in a phylogenetic context. We sampled 17 species of Carya and 15 species representing all other genera of the Juglandaceae as outgroups, with eight nuclear and plastid loci to reconstruct the phylogeny of Carya. The phylogenetic positions of seven extinct genera of the Juglandaceae were inferred using morphological characters and the molecular phylogeny as a backbone constraint. Divergence times within Carya were estimated with relaxed Bayesian dating. Biogeographic analyses were performed in DIVA and LAGRANGE. Diversification rates were inferred by LASER and APE packages. Our results support two major clades within Carya, corresponding to the lineages of eastern Asia and eastern North America. The split between the two disjunct clades is estimated to be 21.58 (95% HPD 11.07-35.51) Ma. Genus-level DIVA and LAGRANGE analyses incorporating both extant and extinct genera of the Juglandaceae suggested that Carya originated in North America, and migrated to Eurasia during the early Tertiary via the North Atlantic land bridge. Fragmentation of the distribution caused by global cooling in the late Tertiary resulted in the current disjunction. The diversification rate of hickories in eastern North America appeared to be higher than that in eastern Asia, which is ascribed to greater ecological opportunities, key morphological innovations, and polyploidy.     

Chloroplast DNA restriction site study of Verbesina (Asteraceae: Heliantheae)

Chloroplast DNA variation was examined for 79 species of Verbesina and 24 outgroups. Two independent analyses of the data were performed. An intergeneric study to elucidate the phylogenetic relationships of Verbesina used 22 genera, most of which are regarded in the literature as closely related to Verbesina. Coreopsis and Hymenoxys served as outgroups for this analysis. The 16 6-bp (base pair) restriction endonucleases used in the intergeneric study revealed 263 phylogenetically informative sites. Wagner analyses of these characters resulted in four equally parsimonious trees with a length of 857 steps and a Consistency Index of 0.492. Results from this study indicate that Verbesina is monophyletic, a member of the tribe Heliantheae, and that its sister taxa are the mostly Mexican genera Podachaenium, Squamopappus, and Tetrachyron. The infrageneric study of Verbesina included species belonging to all of its infrageneric taxa, except the monotypic sect. Stenocarpha. The 17 6-bp restriction endonucleases used in the infrageneric study revealed 137 sites 77 of which were phylogenetically informative. Wagner analyses of these characters generated 180 equally parsimonious trees with a length of 158 steps and a Consistency Index of 0.786. The genera Podachaenium, Squamopappus, and Tetrachyron served as outgroups. Two major clades, which correspond to traditional divisions of the genus based on leaf arrangement, are supported by the study. No support was found for the monophyly of seven of the 11 sections examined; further sampling of sections Lipactinia, Ochractinia, Verbesina, and Ximenesia is needed. Results support a major reassessment of the traditional infrageneric classification of the genus and provide the basis for the reevaluation of the sectional taxonomy of Verbesina. These studies support a North American origin for the genus with several independent introductions into South America producing significant diversity, especially in the Andean region.     

Evolution and losses of spines in slug caterpillars (Lepidoptera: Limacodidae)

Larvae of the cosmopolitan family Limacodidae, commonly known as “slug” caterpillars, are well known because of the widespread occurrence of spines with urticating properties, a morpho‐chemical adaptive trait that has been demonstrated to protect the larvae from natural enemies. However, while most species are armed with rows of spines (“nettle” caterpillars), slug caterpillars are morphologically diverse with some species lacking spines and thus are nonstinging. It has been demonstrated that the evolution of spines in slug caterpillars may have a single origin and that this trait is possibly derived from nonstinging slug caterpillars, but these conclusions were based on limited sampling of mainly New World taxa; thus, the evolution of spines and other traits within the family remains unresolved. Here, we analyze morphological variation in slug caterpillars within an evolutionary framework to determine character evolution of spines with samples from Asia, Australia, North America, and South America. The phylogeny of the Limacodidae was reconstructed based on a multigene dataset comprising five molecular markers (5.6 Kbp: COI, 28S, 18S, EF‐1α, and wingless) representing 45 species from 40 genera and eight outgroups. Based on this phylogeny, we infer that limacodids evolved from a common ancestor in which the larval type possessed spines, and then slug caterpillars without spines evolved independently multiple times in different continents. While larvae with spines are well adapted to avoiding generalist predators, our results imply that larvae without spines may be suited to different ecological niches. Systematic relationships of our dataset indicate six major lineages, several of which have not previously been identified.     

A chloroplast phylogeny of Arisaema (Araceae) illustrates Tertiary floristic links between Asia, North America, and East Africa

The evolution of Arisaema is reconstructed, based on combined sequences (2048 aligned bases) from the chloroplast trnL intron, trnL-trnF spacer, and rpl20-rps12 spacer obtained for species from all 11 sections, including sectional type species and geographically disjunct East African and North American/Mexican species. Analyses were rooted with a representative sample of the closest outgroups, Pinellia and Typhonium, to rigorously test the monophyly of Arisaema. Sections in Arisaema are mostly based on leaf, stem, and inflorescence characters and, with one exception, are not rejected by the molecular data; however, statistical support for sectional relationships in the genus remains poor. Section Tortuosa, which includes eastern North American A. dracontium and Mexican A. macrospathum, is demonstrably polyphyletic. The third New World species, A. triphyllum, also occurs in eastern North America and groups with a different Asian clade than do A. dracontium/A. macrospathum. The genus thus appears to have entered North America twice. Fossil infructescences similar to those of A. triphyllum are known from approximately 18 million-year-old deposits in Washington State and can serve to calibrate a molecular clock. Constraining the age of A. triphyllum to 18 million years (my) and applying either a semiparametric or an ultrametric clock model to the combined data yields an age of approximately 31-49 my for the divergence of A. dracontium/A. macrospathum from their Asian relatives and of 19-32 my for the divergence between African A. schimperianum and a Tibetan/Nepalese relative. The genus thus provides an example of the Oligocene/Miocene floristic links between East Africa, Arabia, the Himalayan region, China, and North America. The phylogeny also suggests secondary loss of the environmental sex determination strategy that characterizes all arisaemas except for two subspecies of A. flavum, which have consistently bisexual spathes. These subspecies are tetraploid and capable of selfing, while a third subspecies of A. flavum is diploid and retains the sex-changing strategy. In the molecular trees, the sex-changing subspecies is sister to the two non-sex-changing ones, and the entire species is not basal in the genus.     

Mitochondrial and nuclear phylogenies of Cervidae (Mammalia, Ruminantia): Systematics, morphology, and biogeography

The family Cervidae includes 40 species of deer distributed throughout the northern hemisphere, as well as in South America and Southeast Asia. Here, we examine the phylogeny of this family by analyzing two mitochondrial protein-coding genes and two nuclear introns for 25 species of deer representing most of the taxonomic diversity of the family. Our results provide strong support for intergeneric relationships. To reconcile taxonomy and phylogeny, we propose a new classification where the family Cervidae is divided in two subfamilies and five tribes. The subfamily Cervinae is composed of two tribes: the tribe Cervini groups the genera Cervus, Axis, Dama, and Rucervus, with the Père David's deer (Elaphurus davidianus) included in the genus Cervus, and the swamp deer (Cervus duvauceli) placed in the genus Rucervus; the tribe Muntiacini contains Muntiacus and Elaphodus. The subfamily Capreolinae consists of the tribes Capreolini (Capreolus and Hydropotes), Alceini (Alces), and Odocoileini (Rangifer + American genera). Deer endemic to the New World fall in two biogeographic lineages: the first one groups Odocoileus and Mazama americana and is distributed in North, Central, and South America, whereas the second one is composed of South American species only and includes Mazama gouazoubira. This implies that the genus Mazama is not a valid taxon. Molecular dating suggests that the family originated and radiated in central Asia during the Late Miocene, and that Odocoileini dispersed to North America during the Miocene/Pliocene boundary, and underwent an adaptive radiation in South America after their Pliocene dispersal across the Isthmus of Panama. Our phylogenetic inferences show that the evolution of secondary sexual characters (antlers, tusk-like upper canines, and body size) has been strongly influenced by changes in habitat and behaviour.     

Testing hypotheses of speciation timing in Dicamptodon copei and Dicamptodon aterrimus (Caudata: Dicamptodontidae)

Giant salamanders of the genus Dicamptodon are members of the mesic forest ecosystem that occurs in the Pacific Northwest of North America. We estimate the phylogeny of the genus to test several hypotheses concerning speciation and the origin of current species distributions. Specifically, we test competing a priori hypotheses of dispersal and vicariance to explain the disjunct inland distribution of the Idaho giant salamander (D. aterrimus) and to test the hypothesis of Pleistocene speciation of Cope's giant salamander (D. copei) using Bayesian hypothesis testing. We determined that available outgroups were too divergent to root the phylogeny effectively, and we calculated Bayesian posterior probabilities for each of the 15 possible root placements for this four-taxon group. This analysis placed the root on the branch leading to D. aterrimus, indicating that current distribution and speciation of D. aterrimus fit the ancient vicariance hypothesis and are attributable to the orogeny of the Cascade Mountains rather than recent inland dispersal. Furthermore, test results indicate that D. copei is distantly related to other coastal lineages and likely originated much earlier than the Pleistocene. These results suggest that speciation within the genus is attributable to ancient geologic events, while more recent Pleistocene glaciation has shaped genetic variation and distributions within the extant species.     

Phylogeny and biogeography of the dung beetle genus Phanaeus (Coleoptera: Scarabaeidae)

Abstract The dung beetle genus Phanaeus as currently recognized by Edmonds (1994) consists of 51 species placed in 13 species groups and two subgenera. Here, I examine the phylogeny and biogeography of this genus by analysing the mitochondrial cytochrome oxidase subunit I (530 bp), nuclear large subunit ribosomal RNA (28S, D2 region), and 67 morphological characters for 28 species of Phanaeus. Both maximum parsimony and Bayesian analyses from the combined data yielded well‐resolved trees, although low bootstrap and posterior probability support were found for basal nodes. The phylogenetic hypotheses presented here suggest that the subgenera Phanaeus s.str. and Notiophanaeus should each be elevated to the status of full genus. With the exception of the eucraniine outgroups, the paleano species group of the genus Phanaeus is recovered as sister to all other taxa, including the outgroups Oxysternon, Sulcophanaeus and Coprophanaeus. High bootstrap values and posterior probabilities supported the species groups endymion, tridens and vindex. Biogeographical analyses suggest an ancestral distribution for Phanaeus in the Andes in South America, although numerous dispersal events evidently have produced a complicated biogeographical history.     

A new species of the genus Lightiella: the first record of Cephalocarida (Crustacea) in Europe

A new species of Cephalocarida belonging to the genus Lightiella is described. Like all known species of Lightiella , the new species is characterized by reduction of trunk segment 8, which also lacks both pleura and thoracopods. The diagnostic characters of the species are: (1) one seta on the inner distal corner of the penultimate endopodal segment of second maxilla and thoracopods 1–5; (2) only one claw on the distal segment of the endopod of thoracopod 6. A cladistic analysis of 27 morphological characters was used to estimate the phylogeny of all species of Lightiella , with all other cephalocarid species used as outgroups. The discovery of this species in the Mediterranean fills a gap in the distribution of the genus and of the entire class.  © 2006 The Linnean Society of London, Zoological Journal of the Linnean Society , 2006, 148 , 209–220.     

Phylogenetic relationships of the genus Hoplia Illiger (Scarabaeidae: Hopliinae)

Results of phylogenetic analysis based on 34 morphological characters of 24 species of 11 genera of Hopliinae from Europe, Japan, South Africa, Madagascar, North and Central America, indicates that the genus Hoplia is a monophyletic group with species distributed in Europe, Japan and America. Based in this analysis the Asiatic genus Ectinohoplia is the closest relative of the genus Hoplia, and the South American genus Barybas (Melolonthinae: Macrodactylini) is the sister group of Hopliinae.     

Systematics of the genus Mayazomus (Arachnida: Schizomida): the relevance of using continuous characters and pedipalp setae patterns to schizomid phylogenetics

The schizomid genus Mayazomus Reddell & Cockendolpher, 1995, endemic to south‐eastern Mexico, currently comprises seven species. It was originally proposed to accommodate two species, from Chiapas and Tabasco. Recently, five additional species from Chiapas were described. The monophyly of the genus has never been tested using cladistic analysis. We undertook a phylogenetic analysis using the seven described species of Mayazomus as the ingroup, ten exemplar species representing the most similar New World hubbardiids as the outgroup, and one protoschizomid species to root the tree. The analysis was based on 130 morphological characters (continuous and discrete characters). The resulting topologies recovered Mayazomus as paraphyletic, with Heteronochrus estor Armas & Viquez, 2010, from Guatemala nested within the genus; therefore, we formally propose its synonymy herein. Mayazomus appears to be most closely related to Rowlandius Reddell & Cokendolpher, 1995, a South American genus. This contribution also provides new characters derived from the pedipalp setae with important phylogenetic information; as well as the implementation of morphometric ratios, as continuous characters, to partially codify the shape of the male flagellum. The relationships recovered amongst the outgroups used in this contribution are a reliable baseline for future analyses of the phylogeny of the New World schizomids.     

A Conspectus of Myriophyllum (Haloragaceae) in North America

Thirteen species of the genus Myriophyllum are found in North America; ten are native, three have been introduced. A key to the species is provided and previous conflicting taxonomic treatments are assessed. The possibility of a polyploid series involving four species is considered and dismissed. Eight of the common species are found to exhibit sufficient phenotypic plasticity to account for most if not all the taxonomic problems encountered.     

Latitudinal clines in alternative life histories in a geometrid moth

The relative roles of genetic differentiation and developmental plasticity in generating latitudinal gradients in life histories remain insufficiently understood. In particular, this applies to determination of voltinism (annual number of generations) in short‐lived ectotherms, and the associated trait values. We studied different components of variation in development of Chiasmia clathrata (Lepidoptera: Geometridae) larvae that originated from populations expressing univoltine, partially bivoltine or bivoltine phenology along a latitudinal gradient of season length. Indicative of population‐level genetic differentiation, larval period became longer while growth rate decreased with increasing season length within a particular phenology, but saw‐tooth clines emerged across the phenologies. Indicative of phenotypic plasticity, individuals that developed directly into reproductive adults had shorter development times and higher growth rates than those entering diapause. The most marked differences between the alternative developmental pathways were found in the bivoltine region suggesting that the adaptive correlates of the direct development evolve if exposed to selection. Pupal mass followed a complex cline without clear reference to the shift in voltinism or developmental pathway probably due to varying interplay between the responses in development time and growth rate. The results highlight the multidimensionality of evolutionary trajectories of life‐history traits, which either facilitate or constrain the evolution of integrated traits in alternative phenotypes.     

Morphology and Biology of the Preimaginal Stages of Some Species of the Genus Eupithecia Curt. (Lepidoptera,Geometridae)

Entomological Review - The biology, host plants, and statial distribution of 17 species of the genus Eupithecia Curt. (Lepidoptera, Geometridae) from different regions of Asia (Kazakhstan,...     

Genetic relationships among American species of the genus Prosopis (Mimosoideae, Leguminosae) inferred from ITS sequences: evidence for long-distance dispersal

Aim  The genus Prosopis includes 44 species and has a pseudoamphitropical, disjunct distribution. We aimed to determine whether American Prosopis sections arose in North or South America, and to explain the current distribution of their species on the basis of their genetic relationships.
Location  South-western USA, Mexico, Caribbean Antilles, Peru–Ecuador, central and northern Argentina, south-western Argentina (Patagonia) and Cuyo, south-western Asia and northern Africa.
Methods  Internal transcribed spacer fragments from 21 species of Prosopis were sequenced and the data were used to analyse the phylogenetic relationships using Microlobius and Mimosa as outgroups. Genetic distances were calculated to estimate the degree of divergence. Dispersal–vicariance (DIVA) analysis was conducted to help understand the biogeographical history of the genus.
Main conclusions  The sections Strombocarpa and Algarobia are not monophyletic. Prosopis argentina (section Monilicarpa ) and the species of Algarobia are included in single clade. The phylogeny, DIVA analysis, and the pattern of genetic distances indicate that the ancestral area for the American species was wide, from south-western USA to Central and northern Argentina. Successive vicariance events split this area, and long-distance dispersal episodes (perhaps mediated by birds) led to recolonizations from North to South America, and vice versa .